Memoirs
OF THE
Queensland Museum
Brisbane
June, 1978
Volume 18
Part 2
Volume 18
Part 2
Memoirs
OF THE
Queensland Museum
Published by Order of the Board
Printed in Australia by The Courier-Mail Printing Service, Campbell Street. Bowen Hills Q, 4006.
THE
QUEENSLAND MUSEUM
Gregory Terrace
Fortitude Valley
Brisbane Qld 4006
Telephone: 52 2716-8
TRUSTEES
J. C. H. Gill, B.A., LL.B. (Chairman)
I. G. Morris, C.M.G. (Vice Chairman)
Emeritus Professor S. A. Prentice, M.E.E., B.Sc.
Professor J. M. Thomson. D.Sc.
D. M. Traves, O.B.E., B.Sc.
R. I. Harrison. M.B.E., E.D., B.Comm.
A. F. Hinchy, A.A.U.Q.
A. Bartholomai, M.Sc., Ph.D. {ex officio, Secretary)
SCIENTIFIC STAFF
Director: A. BARTHOLOMAI, M.Sc., Ph.D.
Deputy Director: B. M. Campbell, M.Sc. (Editor)
Senior Curators: E. C. Dahms, B.Agr.Sc., B.Sc., Curator of Insects
Patricia Mather. Ph.D., D.Sc., Curator of Higher Invertebrates
M. C. QUINNELL, B.A.(Hons), Curator of Anthropology/ Archaeology
D. Robinson, B.Sc.(Hons), Ph.D., Curator of History and Technology
Mary Wade, B.Sc.(Hons), Ph.D., Curator of Geology
Curators: M. ARCHER, B.A., Ph.D., Curator of Mammals
M. J. Bishop, M.A., Ph.D., Curator of Molluscs
L. R. G. Cannon, B.Sc., Ph.D,, Curator of Lower Invertebrates
Jeanette CovaCEVICH, B.A., Curator of Reptiles
Valerie Davies, M.Sc., D.Phil., Curator of Arachnids
R. G. Hardley, B.A., (Hons), Curator of Australian Ethnography
G. J. Ingram, B.Sc., Curator of Amphibia
R. J. McKay, B.A., Curator of Fishes
R. W. Monroe, B.Sc., Curator of Crustacea
I. G. Sanker, B.Sc., Dip. Ed., Curator of Industrial Technology
P. P. Vernon, Ornithologist
Mem. QdMus. 18(2): 127-43, pis. 23-7. [1978]
THE MACROPODIDAE (MARSUPIALIA) FROM THE
ALLINGHAM FORMATION, NORTHERN QUEENSLAND.
RESULTS OF THE RAY E. LEMLEY EXPEDITIONS, PART 2.
Alan Bartholomai
Queensland Museum
ABSTRACT
The fossil Macropodidae from the Bluff Downs fauna of the Allingham Formation, northern
Queensland, of lower Pliocene age, are described. Eight taxa have been recognized, although
inconclusive evidence exists to suggest that additional taxa may be defined when broader
collections become available. Potoroines are currently unknown and slhenurines are very poorly
represented- Macropodines predominate. Four species are new. Troposodon bluffensis,
Protemnodon snewini. Macropus (Prionotemnus) narada and M. iOsphranter) pavana.
Two species, M. i?P.) dryas and T. minor are known also from the Chinchilla Sand, although
the latter species has a wide temporal range being well represented in Pleistocene deposits.
The occurrence none the less supports the previously ascribed late Pliocene age for the
Chinchilla Sand.
A preliminary statement on the fauna from the
Allingham Formation and the definition of the
Formation are provided in Archer and Wade
(1976). The unit is regarded as being of lower
Pliocene age because it is overlain in part by the
Allensleigh ‘How' of the Nulla Basalt, dated
radiometrically at between 4 and 4 5 M.y. This
then represents a minimal age for the Formation,
although from what is already known of the fauna,
actual time of deposition was probably not much
earlier. Continuing investigation of the deposits
has been made possible through the support of Dr
Ray E. Lemley as part of a joint Queensland
Museum and South Dakota School of Mines
project. Larger collections of macropodids have
resulted, enabling the present study to be
undertaken. Other groups will form the basis for
additional studies.
The deposits are considered to be of great
importance and they represent only the second
Pliocene, mammal-bearing site in Queensland.
The reasonably precise minimal age and the
known diversity of the fauna suggest that results
will assume some significance in correlation of
Australian lacustrine and terrestrial deposits.
All measurements throughout are in milli-
metres.
STHENLRINAE
Sthenurus sp.
(Plate 23, fig. 1)
Mati-.rim. Examined: F9104, partial, isolated
right (breadth 12‘6).
DISCUSSION; Only one fragmentary tooth of
Stenurus Owen has been recovered from the
Allingham Formation. Its morphology suggests
that it is an ML Insufficient features are preserved
for more than generic determination to be
attempted. However, of species currently known,
the tooth is believed to be from a form close to
Sthenurus antiquus, described by Bartholomai
(1963) from the late Pliocene Chinchilla Sand.
Ornamentation and height of the lophs is similar,
although in the Allingham specimen, the strength
of the ridge which ascends into the median valley
from the metaconc is considerably weaker than in
the maxilla of S. antiquus figured by Bartholomai
(1963, fig. 10). As in S. antiquus, the base of the
crown is swollen posteriorly and the low midlink
is positioned above the centre of the median valley.
However, the strong ridge curving posterolingual-
ly from the hypocone overlaps, rather than meets,
the weaker ridge from the metacone.
128
MEMOIRS OF THE QUEENSLAND MUSEUM
Although upper teeth are unknown in S.
notabilis Bartholomai, also from the late Pliocene
Chinchilla Sand (Bartholomai 1963). size and
crown height would preclude this species from
close relationships with the Allingham speci-
men.
Among Pleistocene species of Sthenurus
reviewed by Bartholomai (1963) and Tedford
(1966), or described by Merrilecs (1965, 1967),
the Allingham specimen appears to be within the
size range of S. areas De Vis, but has less
pronounced accessory ridging. S. orientalis
Tedford of comparable size, is known only from
its lower dentition, while S. occidentalis Glauert,
S. brownei Merrilees and S. gilli Merrilees are
all smaller species.
MACROPODINAE
Troposodon minor (Owen, 1877)
(Plate 23, figs. 2-4)
Sthenurus minor Owen, 1877, pp. 352-61, pi. 37 , figs.'
1-3, pi. 38, figs. 1-4.
Troposodon minor (Owen); Bartholomai, 1967, pp.
23-32, figs. 1-3 (synonymy).
Matpriai. Examim-D: F9046, isolated left P^;
F9047, isolated partial left M.; F9048. isolated partial
left M 3 ; F9049, isolated left DP\ F9050. isolated right
M,; F9051, isolated left DPh F9052, isolated partial
right M’; F9053, isolated partial right M|.
DISCUSSION: The genus Troposodon
Bartholomai is currently represented in Australian
Upper Cainozoic deposits by two species, T. minor
(Owen) and T. kenti Campbell. Of these, T,
minor is by far the most commonly recorded
species, being present in sufficient numbers in
collections from the Darling Downs deposits to
enable Bartholomai (1967) to assess the
populations statistically.
Within the Darling Downs deposits, Barth-
olomai (1967) has shown T. minor to be present
in both the Pleistocene fluviatile deposits and the
Chinchilla Sand ot late Pliocene age. Samples
were compared using Student’s I Test and no
significant size or morphological features were
found to justify separation. Turnbull and
Lundelius (1970) have also suggested that an
isolated, partial lower molar. MNV-P26422, from
the post-early Pliocene fossil soil deposits at
Grange Burn, Hamilton, Victoria, may represent
T. minor, a possibility which cannot be discounted
on the basis of the description provided.
T. kenti from the ?early Pleistocene Katipiri
Sands, has been distinguished from T. minor by
Campbell (1973) on the basis of its larger size,
TABLE I: Measurements eor Troposodon minor
(Owen, 1877) '
Specimen
u
Ml
M.
M 3
DP3
F9046
12-8x51
F9047
—
—
— x9-3
F9048
—
—
—
— xlO-2
F9050
—
11-0x6-9
F9053
—
— x7-3
—
F9049
—
—
—
—
10 - 2 x 8-1
F9051
—
—
—
—
10-7x7-0
supported by the less ornamented nature of the
apical shelves of the molar teeth and minor
morphological differences in the premolars.
Representation of material larger than T. minor
in the Chinchilla Sand material, but insufficient
for description, was noted by Bartholomai (1967).
No additional material has as yet been recovered
to support or refute the suggestions by Campbell
(1973) that these specimens may represent T.
kenti in the Chinchilla Sand, or that they may
reflect extreme variation in T. minor.
Although the sample available from the
Allingham Formation is small, comprising only
isolated teeth, a reasonable range of dental
morphology is represented and some variation
within the population is observable. Measur-
ements for this material appear in Table 1.
Slight size differences which are evident
between teeth from the Allingham Formation and
those from the Chinchilla Sand are attributable
to the small sample size in the former and the
fact that the Chinchilla Sand sample itself is a
relatively small one for statistical purposes.
Nevertheless, the Allingham sample falls within
limits which one would expect for the Chinchilla
Sand sample, based on large sample assessments
for macropodines provided in Bartholomai (1971,
1973, 1975). Structurally, the Allingham sample
generally presents coarser accessory ridging in
anterior molars and in DP^ than is usual in
Darling Downs specimens, but this feature is
variably present in some individuals from the
Darling Downs, including the Juvenile specimen
figured by Batholomai (1967, fig. 2).
No differences are therefore present which
enable separation of the Allingham material from
T. minor, although T. kenti is clearly excluded
by both size and morphological considerations.
This is of particular interest because of the age
of the Allingham Formation, which indicates a
minimal temporal spread for T. minor from at
least the mid Pliocene to the late Pleistocene, the
longest recognisable range for an Australian
marsupial species.
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
129
Troposodon bluffensis* sp. nov.
(Plate 23, figs. 5, 7-10)
Mattrim. Examined F9054. holotype, isolated
right M**. adult, Allingham Formation of mid Pliocene
age, Bluff Downs Station, Allingham Creek.
In addition, F9055, an isolated left M 4 , F9056, a
partial isolated left M‘*; F9057, an isolated right M 4 ;
F9058, an isolated right P 2 and F9059, an isolated left
P^ have been used in the description of the species.
Specific Diagnosis: a small species of
Troposodon. much smaller than T. minor: P^
small, with low longitudinal crest; upper molars
lacking a forelink or with this present in very
incipient form, and lacking a ridge from the
paracone to the labial extremity of the anterior
cingulum; midlink high with major contribution
from near midpoint of melaloph crest. P^ small,
low crowned, markedly L-shaped; lower molars
with reduced accessory ridging on apical shelf and
with base of crown swollen posteriorly without
production of posterior cingulum.
DESCRIPTION; Upper dentition known only
from P^ and M**.
P^ elongate, subcrcscentic in basal outline, with
labial margin concave and lingual convex, broader
posteriorly than anteriorly; longitudinal crest low,
more elevated posteriorly than anteriorly,
moderately concave iabially, transected by three,
weak sets of vertical labial and lingual ridges,
these becoming weaker posteriorly. Apex of
paracone about one-fifth distance along crown
from anterior limit. Hypocone moderately low,
positioned posterolingual to metacone and
connected to it by strong descending ridge; weak
ridge curves posterolabially to meet extremely
weak posterior ridge from metaconc near base of
crown, without production of posterior fossette;
strong anterior ridge from hypocone ascends to
form well-defined lingual cingulum to point
opposite paracone; cingulum sinuous in lingual
view, converging only slightly anteriorly in
occlusal view; lingual basin broad, smooth.
Anterior ridge from paracone extending well
anteriorly; base of crown slightly swollen
anterolingually.
small, subrectangular in basal outline,
slightly constricted across median valley; lophs
low, moderately bowed anteriorly, with melaloph
narrower than prololoph. Anterior cingulum low.
*Named for Bluff Downs Station, the property on which
the Allingham Formation occurs.
moderately broad, anteriorly broad convex, nearly
flat in labial moiety, but ascending slightly
iingually; forelink absent or present in very
incipient form; labial margin of cingulum
unconnected to paracone by accessory ridge;
midlink high, strong, curving posterolabially from
prolocone, then curving posteriorly abruptly to
unite with strong ridge from near midpoint of
crest of mctaloph; Junction marked by shallow
cleft. Weak ridge ascends poslerolingually from
paracone towards midlink, defining unornamented
apical shelf; median valley shallowly U-shaped,
transversely flaitcncd, with lingual extremity
marked by low ridge; strong ridge curves
posterolabially from hypocone towards base of
crown; weaker ridge from metacone ascends to
disappear below ridge from hypocone with
production of posterolabial indentation; accessory
ridges from hypocone and metacone ascend across
posterior face of melaloph to unite near midline,
defining unornamented apical shelf. Base of crown
narrowly swollen posteriorly, this separated below
by shallow groove.
Lower dentition known only from Pt and
M4.
P 2 relatively small, subcrescentic in occlusal
view, slightly broader posteriorly than anteriorly;
longitudinal crest relatively low, markedly
L-shaped, being flexed Iingually in its posterior
extension. Anterior cuspid set posteriorly at about
one-half distance along crown from anterior
margin, with anterior ridge from cuspid angled
anteriorly and marked by low cuspule about
one-half distance to crown base; low, anterolabial
cingulum developed from this cuspule to unite
with labial ridge from anterior cuspid. Longitudin-
al crest transected by two weak sets of vertical
labial and lingual ridges; lingual extension of crest
descends slightly to posterolingual cuspid, this
having strong posterior ridge and stronger anterior
ridge descending from it. Base of crown swollen
Iingually and Iabially.
M 4 small, subrectangular in basal outline,
moderately strongly constricted across talonid
basin; lophids low, moderately strongly convex
posteriorly, with prololophid broader than
hypolophid. Trigonid basin moderately broad, its
length approximately equalling distance between
lophids. Forclink high, strong, curving antero-
lingually from protoconid across trigonid to point
on anterior cingulum labiad to midline; anterior
cingulum moderately high, broad, rounded
anterolabially, more angular anterolingually,
descending markedly Iabially and Iingually from
forelink; strong accessory ridge curves anter-
olabially from metaconid towards forelink.
130
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 2: Mhasuri MhNTS for Troposodon bluffensis
SP. NOV.
Specimen
M'*
P 2
M 4
F9054.
holotype
lI-7x8-5
F9056
—
— x9-l
—
—
F9055
—
—
—
12 - 0 x 8-2
F9057
—
—
—
12-7x8-2
F9058
1 0-6x5-]
—
—
—
F9059
—
—
70x3-5
—
defining apical shelf: weak accessory ridges
present on apical shelf, descending anteriorly from
near protolophid crest. Talonid basin narrowly
U-shaped, descending slightly labially and
lingually from midlink. Midlink strong, moderate-
ly high, curving slightly anterolinguaily from
hypoconid to near midpoint of posterior of
protolophid; slight ridge curves towards midlink
from enloconid without production of apical shelf.
Base of crown swollen posteriorly but without
production of posterior cingulum.
DISCUSSION: Although at present known
from a very small sample comprising only isolated
cheek teeth, the material is so distinct both in its
size and morphology as to conclusively justify its
separation as a distinct species of Troposodon
Bartholomai within the Allingham Formation.
Compared with the Chinchilla Sand sample of
T. minor (Owen) of late Pliocene age, the
Allingham material falls well below the lower
observed limits for size in equivalent teeth and,
indeed, is well below the statistical limits for that
population. The morphological distinction of
molars in T. bluffensis lies in the greater
simplicity of links and accessory ornamentation
and in the more generalized basic structure
overall.
The Pt is more complex anteriorly and
anterolabially than in T. minor, but it has more
subdued ridging associated with the longitudinal
crest. Ridges from the postcrolingual cuspid,
especially the anterior ridge, are stronger and
better-defined. The upper permanent premolar is
less triangular in basal outline than in T.
minor.
That two species of Troposodon should be
represented within the Bluff Downs deposits is not
surprising, taking into account the macropodine
representation within other fossil and recent
faunas and the fact that the Allingham Formation
rellects both Huviatile and lacustrine sedimenta-
tion (Archer and Wade 1976) with attendant wide
sampling of endemic and derived specimens.
It is considered likely that in view of its more
generalized nature. T. bluffensis stands closer to
the ancestral form from which Troposodon was
derived than do either 7. minor or 7. kenti.
In addition to the material here referred to 7.
bluffensis. there exists an isolated right P3, F9060,
figured in Plate 23, fig. 8. which may belong with
this species. The tooth is clearly of the
Troposodon type, having the posterior extension
of its longitudinal crest acutely angled lingually,
giving the tooth a marked L-shaped appearance
in occlusal view. It differs from P3 in 7. minor
in being shorter (10- 1x5-2) and in having its
anterior margin near vertical. Although the
longitudinal crest is lower anteriorly than
posteriorly, the crown is relatively high in
comparison with 7. minor and with the P-, referred
to 7. bluffensis. While it is considered highly
unlikely that a further species of Troposodon is
represented in the Allingham Formation, the
specimen is maintained as doubtfully referred to
7. bluffensis until more complete mandibular
material is collected to confirm its taxonomic
position.
Petrogale sp.
(Plate 23, fig. 6)
Matf.rial Examinrd: F7794, isolated left Mt;
F7795, isolated right M 3 ; F9043, isolated left M 2 :
F9044, isolated left M 4 ; F9045, isolated left M-,.
DISCUSSION: The isolated molars here
referred to Petrogale sp. were regarded by Archer
in Archer and Wade (1976) as c.f. Thylogale sp.,
but were not figured. Petrogale and Thylogale are
difficult to separate on the basis of isolated cheek
teeth, particularly molars. However, it appears
that Petrogale molars are more like those in
Macropus than are those in Thylogale. In
Thylogale, the lophids are very narrow longitudin-
ally, even towards the crown base, while
transversely, the lophid crests are more rectilinear.
The midlinks are angled more anterolinguaily
from the hypoconid in Thylogale.
In size, the referred sample is similar to P.
penicillata, but insufficient is known of the
morphology of the Allingham sample to justify
more discrete treatment at this tine.
The isolated right DP\ F7785, mentioned by
Archer in Archer and Wade (1976) as being
difficult to distinguish from corresponding teeth
in Thylogale (e.g. 7. stigmatica) does show
greater morphological similarity to Thylogale
than to Petrogale. Rather than promote additional
speculation, its position is left doubtful until more
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
131
TABLE 3: Measurements for Peirogale sp.
Specimen
M 2
M 3
M 4
F7794
6-2x41
F7795
—
7-lx4-l
F9043
6-3x4- 1
F9044
—
—
8-2x5-2
F9045
6-9x4 1
—
—
adequate samples become available. Thylogale is
known from post-early Pliocene deposits at
Grange Burn, Hamilton, Victoria (Turnbull and
Lundelius 1970).
Protemnodon snewini* sp. nov.
(Plate 24. figs. 1-4; Plate 25, fig. 1)
Protemnodon sp. Archer 1976 (partim), pp. 390-1,
pi. 57a.
Materiai Examined: F9061, holotype, right
mandibular ramus with I|, P 3 -M 4 , adult, Allingham
Formation of mid Pliocene age, Bluff Downs Station,
Allingham Creek.
This specimen is possibly associated with the partial
skull, F9074, and a partial, disarticulated skeleton,
F9075, all located close to one another at the same
stratigraphic level within the deposits.
In addition, F7810, a partial right maxilla with
M'-M^ shattered M^-M"*, adult; F7809. an isolated left
M^; F9067. an isolated, partial right M 4 ; F7788, an
isolated left P^; F9068. an isolated right M 3 , F781 1, an
isolated right M**; F9069, an isolated right M’; F9070,
a partial, isolated left Pj; F9071, an isolated left M^
F9072, an isolated left P-; F9073, an i.solated right P-;
F7786, a right P^, aged; F9062, an isolated right P^;
F9063, an isolated partial right M 4 ; F9064, an isolated
right M|; F9065, an isolated left M 3 : F9066, isolated
partial left M^; F7824, an isolated right P^; have been
used in the description of the species.
Specific Diagnosis: A small species,
generally smaller than P. anak. Mandible
comparatively very shallow with an elongate
symphysis ascending at about 10 ^ to base of
mandible; lateral groove extends to below
posterior root M 3 ; menial foramen close to
diastemal crest about one-quarter distance along
diastema from anterior root P 3 . Condyle reduced
transversely.
Ii shallow. Cheek tooth row slightly bowed
labially in occlusal view. P 3 narrow, elongate,
slightly longer than M 4 ; crown low, with straight
longitudinal crest transected by four sets of
slightly anteriorly directed labial and lingual.
* Named for Mr W. Snewin who, with Mr J. Barrett,
discovered the Allingham site.
ridges. Lower molars slightly constricted across
talonid basin, w'ith lophids low, links labiad to
mid-line; lingual moiety of trigonid unswollen;
anterior cingulum broad, extended anterolingually
into distinct, subangular corner to crown; talonid
broadly U-shaped; posterior cingulum poorly
developed.
P- elongate, with longitudinal crest low,
transected by three or four sets of vertical labial
and lingual ridges; lingual cingulum well
developed, with shallow lingual basin narrowing
slightly anteriorly. P^ narrow, elongate, slightly
constricted basally at posterior one-third;
longitudinal crest nearly straight, low, transected
by three or four sets of vertical labial and lingual
ridges; lingual cingulum very low, continuing to
anterior of crown base, sinuous; lingual basin very
narrow, shallow. Upper molars with low lophs,
unconstricted, or slightly constricted across
median valley in posterior molars; strong ridge
ascends from paracone to limit of broad, flat,
anterior cingulum; midlink very low; posterior
ridge from paracone very weak; metaloph
relatively broad in M*^-
Description-. Mandible very shallow,
relatively thick; base of symphysis ascending
anteriorly at an angle of approximately 10 “ to
base of mandible; symphysis elongate, shallow, not
ankylosed, rugose; geniophyal pit shallow, near
posterior symphysial limit; diastema elongate with
diastema! crest angular posteriorly, more rounded
anteriorly; ventral margin of ramus rounded
between symphysis and extremely weak diagastric
ridge and process. Mental foramen small, ovate,
near diastemal crest, about one-quarter distance
along crest from anterior root P 3 . Ramus with
shallow lateral groove extending posteroventrally
to below posterior root M 3 , somewhat disrupted
by roots of P 3 : posterior of groove approximately
mid-way between alveolar margin and base of
ramus. Diagastric process separated from base of
angle by very shallow post-diagastric sulcus,
bounded above by shallow diagastric fossa. This
fossa separated above from shallow, broad
depression opening posteriorly into pterygoid
fossa. Post-alveolar shelf short, with shelf angle
reasonably well defined, leading to post-alveolar
ridge ascending to disappear rapidly on medial
wall of coronoid process, above large mandibular
foramen. Masseteric crest low, approximately at
level of alveolar margin; masseteric foramen
moderately large with deep masseteric fossa.
Angle of mandible broadly inflected, produced
posteriorly and dorsally into crest. Anterior of
coronoid process ascends posterodorsally at about
132
MEMOIRS OF THE QUEENSLAND MUSEUM
13 ° from vertical. Condyle very reduced, longer
than broad, planar transversely and gently convex
longitudinally, supported posteriorly by narrow
shelf.
I) elongate, deeply rooted; slightly curved in
lateral view, markedly curved in occlusal view,
developing subhorizontal transverse facet of wear,
but gently convex wear facet longitudinally with
upper incisors; mesial wear facet present along
ventral margin by approximation with other lower
incisor; root compressed, oval in section; crown
subquadrantal in section, tapering and blade-like
anteriorly, enamelled laterally, this produced
dorsolabially and ventrolingually into flanges;
crown slightly enamelled ventromcsially.
P3 relatively elongate, robust, moderately broad;
crown subovate in basal outline with longitudinal
crest secant, nearly straight but strongly flexed
lingually in its posterior extension; crest transected
by two sets of near vertical labial and lingual
ridges with production of cuspules at crest;
strength of ridges and cuspules decreases
posteriorly; anterior cuspid of crest well defined;
anterior of crown straight, with ridge from
anterior cuspid curving lingually towards base of
crown; lingual base of crown swollen; broad, short,
vertical, basal ridge present posterolabially.
DP3 is not preserved.
P3 elongate, narrow, deeply rooted; crown
subovate in basal outline with longitudinal crest
secant, slightly sinuous, and slightly flexed
lingually in its posterior extension; crest transected
by four sets of slightly anteriorly directed labial
and lingual ridges, with production of cuspules at
crest; strength of ridges and cuspules decreases
posteriorly; anterior cuspid of crest well defined,
but posterior cuspid less well developed; anterior
of crown straight, near vertical extending well
anterior to anterior root; labial base of crown
slightly swollen, but not as strongly as posterior
moiety of lingual base.
Mi<M 2<M3<M4: molars subrectangular in
basal outline, slightly to markedly constricted
across talonid basin; lophids low, slightly convex
posteriorly with hypolophid broader than protolo-
phid in M] and M^, approximately equal in M3
and narrower in M4. Trigonid basin broad, its
length approximately equalling distance between
lophids. Forelink low, moderately strong, curving
slightly anterolingually from protoconid across
TABLE 4: Measurements for Mandibles of Protemnodon snewini sp. nov.
Specimen
li depth
P 2
P 3
M,
M 2
M 3
M 4
F9061,
holotype
— xIO -6
—
15-0x4-9
9-8x6-6
12-3x8-3
13-3x9-3
13-8x9-0
F7824
—
8-7x4-7
F9065
—
—
—
_
12 - 8 x 8-6
F9067
—
—
—
— x8-5
F9068
—
—
13-2x8-7
F9070
—
—
— )^5-I
F9063
—
—
—
_
— x9-7
F9064
—
—
—
9-9x7-l
—
—
TABLE 5: Measurements for Maxillae of Protemnodon snewini sp. nov.
Specimen
P 2
P 2
M'
M 2
M3
M*!
F9074,*(rt.)
—
15-2x6-6
9-9x8-7
— xlO-4
13-1x10-9
13-5x10-5
F9074,*(lt.)
—
—
10 - 0 x 8-8
12-3x10-2
13-1x10-9
13-9x]0-7
F7810
—
■ —
—
13-8x11-6
14-0x11-7
F7811
—
—
—
14-5x11-8
F9069
—
—
11-3x9-9
F7788
10-9x5-8
F9072
10-7x6-2
F7786
10-7x5-4
F7809
—
—
13-0xU-4
F9071
—
—
—
12-1x11-4
F9062
—
16-6x6-6
F9073
—
15-7x6-6
—
—
—
—
'Associated with holotype
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
133
trigonid to point on anterior cingulum well labiad
to midline. Anterior cingulum low, broadly curved
labially but produced anterolingually into well
defined corner; near horizontal lingualiy, but
descending reasonably strongly labially. Midlink
low, strong, decreasing in strength in posterior
molars, descending almost directly from hypocon-
id, anterolingually to point at base of protolophid,
well labiad to midline; slight ridge descends
posterolabially from metaconid. Talonid basin
broadly U-shaped, descending at low angle
labially and lingualiy from midlink; posterior
cingulum poorly defined, frequently represented
only by series of low tubercles.
Upper incisors and DP^ not preserved.
P- elongate, moderately narrow, robust,
subovate in basal outline, somewhat broader
posteriorly than anteriorly; longitudinal crest low,
secant, very slightly concave labially; crest
transected by three or four sets of vertical labial
and lingual ridges with production of cuspules at
crest; apex of paracone positioned about
one-quarter length of crown from anterior limit.
Hypocone moderately strong, united to metacone
by strong, high, descending anlerolabial ridge;
posterior ridge from hypocone curves labially to
unite with posterolingua! ridge ascending from
metacone below crown base; shallow posterior
fossette formed between this and ridge connecting
posterior cusps; anterior ridge from hypocone
ascends rapidly to above crown base to form low
lingual cingulum, converging slightly anteriorly to
position anterior to level of paracone; slight
anterolingual ridge from paracone connects with
anterior of lingual cingulum; cingulum in lingual
view slightly sinuous. Lingual basin shallow,
occasionally transected by weak extensions of
ridges from longitudinal crest. Base of crown
slightly tumid labially.
P3 moderately elongate, narrow, robust,
subovate in basal outline, slightly constricted at
posterior one-third, but otherwise with labial and
lingual margins subparallel; crown with longi-
tudinal crest low, secant, slightly concave labially;
apex of paracone about one-fifth distance along
crown from anterior limit; crest transected by
three or four sets of near vertical labial and lingual
ridges with production of cuspules; strength of
ridges decreases posteriorly. Hypocone moderately
high, united to above metacone by high, strong
labial ridge; strong ridge from hypocone ascends
sharply labially to unite with posterolingual ridge
from metacone; well defined posterior fossette
developed between this and ridge connecting
posterior cusps. Anterior ridge from hypocone
ascends sharply to form very low lingual
cingulum, subparallel to longitudinal crest to
above paracone, then curving sharply to anterior
of crown; more prominent ridges from cuspules
along crest cross very shallow lingual basin to
cingulum; strong ridge from paracone unites with
cingulum; cingulum markedly sinuous in lingual
view, moderately tuberculate. Slight anterolabial
cingulum present. Base of crown very slightly
tumid labially.
M'<M-<M^<M^; molars subrectangular to
subovate in occlusal view, usually slightly
constricted across median valley especially in
posterior molars; lophs low, slightly bowed
anteriorly, with mctaloph broader than protoloph
in ML approximately equal in M^ and slightly
narrower in M**, anterior cingulum low, broad,
short, slightly descending labially; forclink absent,
but variable slight ridges occasionally pass from
base of protoloph towards cingulum; cingulum
generally flattened anteriorly; low ridge usually
descends from paracone to labial limit of
cingulum; midlink very low, its strength
decreasing in posterior molars, curving poster-
olabially from protocone to unite with weak ridge
from near crest of metaloph lingual to midline
above median valley. Weak ridge ascends
posteriorly from paracone, then curves abruptly
lingualiy near median valley to unite with weak,
anterolingual ridge from metacone about onc-half
distance from midlink to labial margin. Median
valley relatively narrowly V-shaped labially and
lingualiy; base of median valley slightly ascending
labially and lingualiy from midlink with labial
moiety sometimes pocketed. Strong ridge curves
posterolabially from hypocone to posterolabial
base of crown, uniting with weaker ridge from
metacone; this delimits very shallow posterior
fossette somewhat labiad to midiine.
DISCUSSION; A partial right mandibular
ramus, F78I2, referred to Protemnodon sp. by
Archer in Archer and Wade (1976), has been
recovered which may not be referrable to P.
sne^\nm. being generally larger than the P. snewini
sample. Measurements appear in Table 6 . This
specimen retains DP 3 and M| and was figured by
Archer in Archer and Wade (1976, pi. 57, fig.
b). The DP^, especially, is morphologically distinct
from P. chinchiiiaensis and P, devisi from the
Chinchilla Sand, in having its protolophid very
restricted and with the forelink and midlink
descending anteriorly and posteriorly from the
proloconid. The labial base of the protolophid
comprises two weak ridges uniting to ascend to
the protoconid. The forclink is very high and the
labial and lingual moieties of the trigonid basin
134
MEMOIRS OF THE QUEENSLAND MUSEUM
descend at high angles from this link. The first
molar possesses a strong posterior cingulum, as
does DPjt, and a weak accessory link is present
close to the near central midlink in the lingual
moiety of the talonid basin. The anterior cingulum
is linked by a moderately strong connecting ridge
to the metaconid. A further specimen, F9077, an
isolated right M 2 , also shows larger size than P.
snewini, has the strong posterior cingulum and
stronger more central links. The possibility that
these specimens indicate the existence of a second
species of Protemnodon from the Allingham
Formation would not be surprising, bearing in
mind the evidence of two species within the
Chinchilla Sand and three within the Pleistocene
deposits of the eastern Darling Downs.
It is obvious from a cursory examination of the
material that P. snewini presents morphological
features which bear comparison with P.
chinchillaensis from the late Pliocene Chinchilla
Sand (Bartholomai 1973) and P. otibandus from
the mid Pliocene Otibanda Formation of the
Bulolo Valley, Papua New Guinea (Plane 1967).
P. devisi, also from the Chinchilla Sand, is
excluded by considerations of molar size alone,
but docs possess permanent premolars which
accord better with P. snewini than do those in P.
chinchillaensis which are actually and relatively
much longer in comparison with molar lengths
(Figure 1). Permanent upper premolars in P.
chinchillaensis are considerably wider teeth and
are generally more robust. The lower incisors in
12’0 _
10-0 .
9-0
z
t-
Q
<
lU
a:
CO
8-0
LJ
LU
is:
UJ
z
o
60 .
z
LU
§
0 ,
5-0 .
4-0
9-0
10-0
15-0
16-0
17-0
18-0
LOWER CHEEK TEETH LENGTH
Fig. 1; Scatter diagram illustrating the proportional relationships of permanent lower cheek teeth in Protemnodon
snewini sp. nov. ( + ) compared with those in Protemnodion chinchillaensis Bartholomai (x). Solid line links
mean values from P 3 -M 4 in P. snewini; heavy dashed line links similar mean values for P. chinchillaensis.
BARTHOLOMAI; ALLINGHAM FORMATION MACROPODIDAE
135
TABLE 6: Measurements for Protemnodon sp.
Specimen
DPj
M|
M 2
F7812
F9077
9-7x4-9
ll-5x7-l
14-0x9-3
P. snewini
are typically
macropodine
, whereas
those in P. chinchillanesis (as in F707I) are
considerably more spatulate. Upper molars in P.
snewini have lower midlinks and broader
metalophs on fVI'^. Compared with P. otibandus,
P. snewini has a shallower, longer mandibular
ramus with its coronoid process less vertical and
with the condyle considerably reduced transverse-
ly. The cheek tooth row is straighten, links are
more labially positioned and are lower, the
anterior cingulum is flatter and broader,
possessing a distinct anierolingual corner, and the
lower premolar is less robust. in P. otibandus
is much more robust, while in upper molars,
midlinks are higher and the anterior cingulum is
not as flattened as in P. snewini.
The left maxilla, attributed by Plane (1972) to
P. otibandus, from the marine lower Pliocene
(Kalimnan) Jemmy's Point Formation, Victoria,
has been suggested previously to be P. devisi by
Bartholomai (1973). Accessory ridging in this
specimen, particularly from the paracone but also
from metacone into the median valley, is stronger
in posterior molars than general in P. snewini. The
Jemmy's Point specimen is larger, being
comparable with dimensions in P. devisi.
Turnbull and Lundelius (1970) have figured
and described an isolated left molariform tooth
which appears referable to Protemnodon. This
specimen, PM 4429, suggested to be a DP-^ by
Turnbull and Lundelius (1970) is from a fossil soil
at Grange Burn, Hamilton, Victoria, dated as
post-early Pliocene. Although direct comparison is
impossible because DP^ is unknown in the
Allingham sample, accessory ridging from the
paracone and metacone into the median valley
appears stronger than in P. snewini, while the
specimen must be larger.
Associated parts of a left hind limb have been
located in the Allingham excavations, indicating
the possibility of recovering more complete
skeletons in the deposits. Although these remains
were not found with cranial specimens, it is
apparent from the proportions of the metatarsal
IV and the preserved elements of digit IV that
they are from a species of Protemnodon. At this
time, it can only be assumed that they are from
P. snewini. Numerous isolated elements from the
hind limbs of macropodines have been located
and, among these, several are morphologically
acceptable to represent elements missing in the
associated limb. These have been included in the
illustration (Fig. 2) and comprise the calcaneum,
cuboid, astragalus, metatarsal V and the proximal
phalange of digit IV.
Fig. 2: Partial right pes, possibly of Protemnodon
snewini sp. nov., all F9075, except calcaneum, cuboid,
astragalus, metatarsal V and proximal phalange of
digit IV, x!/2.
136
MEMOIRS OF THE QUEENSLAND MUSEUM
The ratio of total length of metatarsal IV The most remarkable feature in the preserved,
(102-5) to its median width (20-5) is 500. associated part of the hind limb, is the structure
Compared with P. otihanda, the ratio for mt IV of the tibia which, as shown in Fig. 3, is highly
for which is 518, provided in Plane (1967), the curved mesially. While slight displacement,
bone is relatively broader than in the Awe particularly along fractures in the distal moiety
material. The median phalanx in digit IV is 26-0 of the bone, lends to emphasize this feature, the
long by 23'9 broad, somewhat larger than in P. tibia is much more curved than in other
otihanda and relatively broader. macropodines. Whether the specimen is aberrant
cannot be ascertained at this time but a
pathological cause is suspected.
Fig. 3; Tibia and fibula, possibly of Proiemnodon
snewini sp. nov., F9075, associated with pes illustrated
in Fig. 2, x'/4.
Macropus (Prionotemnus) narada sp. nov.
(Plate 25, figs. 2-3)
Matfriai Examinho. F9105, holotype, partial
left maxilla with DP-^-M’, P-^ missing, juvenile,
Allingham Formation of mid Pliocene age. Bluff Downs
Station, Allingham Creek.
In addition. F9106, isolated left DP-L and F9107,
isolated right have been used in the description of
the species.
Specific Diagnosis: A relatively small
species. Upper molars and DP^ with well-defined
forelink and strong anterior ridge from paracone
to anlerolabial margin of anterior cingulum; fine,
relatively weak ridge passes posterolingually from
paracone towards mid-point of labial moiety of
median valley; midlink high, strong, with major
contribution from near mid-point of hypoloph
crest.
Description; Known only from upper
molariform and molar cheek teeth.
DP^ relatively small, molariform, subovale in
bas^l outline, slightly constricted across median
valley; lophs moderately high, anteriorly bowed,
with metaloph broader than protoloph. Anterior
cingulum relatively high and broad, short,
ascending at moderate angle lingually; strong
forclink present between cingulum and base of
protoloph below mid-line; strong, high ridge
ascends anteriorly from paracone to anterolabial
margin of anterior cingulum; slight ridge curves
posterolingually from paracone into median
valley. Strong, high ridge curves postcrolabially
from protoconc across median valley as midlink
to unite with strong ridge from near middle of
metaloph crest. Median valley ascends labially
and lingually from midlink; valley sharply
V-shaped. Strong ridge ascends postcrolabially
Irom hypocone to meet much weaker, posterior
ridge from melacone near base of crown. Posterior
lossette developed below midline.
M' and preserved; molars subrectangular in
basal outline, slightly constricted across median
valley, with metaloph broader than protoloph in
M* but slightly narrower in M-"*; lophs moderately
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
137
high, anteriorly bowed. Anterior cingulum
relatively high, often broad, short, ascending at
moderate angle lingually; strong forelink present
at midline, very strong, high ridge ascends
anteriorly from paracone to anterolabial margin
of anterior cingulum. Relatively weak ridge
ascends posterolingually from paracone towards
median valley; variable, weak ridge from
metaconc ascends into median valley; strong, high
ridge curves gently anterolabially from protocone
to unite with short, high ridge from near centre
of metaloph crest as midlink. Median valley
sharply V-shaped, ascending slightly labially and
lingually from midlink. Strong ridge curves
posterolabially from hypocone to unite with weak,
posterior ridge from metaconc near base of crown;
slight fossette formed at midline.
Discussion. Although only represented in
the Allingham Formation by a very small sample
of maxillary remains, Macropus {Prionoteninus)
narada sp. nov. is quite distinct from M. dryas,
the other possible Prkmotemnus present in the
deposits. M- dryas lacks the well-defined and
strong ridge which ascends from the paracone to
the anterolabial limit of the anterior cingulum,
present in M, narada It also lacks the accessory
ridges ascending posteriorly from the paracone
and sometimes anteriorly from the metaconc.
Within the Chinchilla Sand, the only recorded
Prionotemnus is P. palankarinnicus Stirton, first
described from the Pliocene Mampuwordu Sands
of the Tirari Desert, South Australia. Upper
molars of this species, studied in the collections
of the University of California, Berkeley, lack
forelinks, but possess the strong anterior ridge
from the paracone to the anterior cingulum.
Compared with the recorded Pleistocene species
from the fluviatile deposits of the eastern Darling
Downs, A/, narada is generally slightly larger than
individuals within M. agilis siva (De Vis) and is
about the same size as M. thor (Dc Vis), both
of which were revised in Bartholomai (1975).
Morphologically, the teeth are closer to those in
M agilis siva but possess a much better-developed
forelink. The forelink in M. thor is subdued and
accessory ridges into the median valley from the
paracone and metaconc are lacking. The anterior
cingulum is near-planar in the labial moiety.
Upper teeth in other Pleistocene species, M.
gouldi (Owen) and M. piltonensis Bartholomai
are unknown at this time.
The lower first molar, F7784, recorded and
figured by Archer in Archer and Wade (1976,
pL57, fig. f) may relate to M. narada. It is unlike
any other macropodid recovered from the
Allingham Formation and, purely by elimination,
and its close occlusal relationships with the upper
molars described here, could well indicate the
morphology to be expected in the lower molars
of A/, narada. Until it is certain that only two
species of A/. (Prionotemnus) are present in the
Allingham, its taxonomic position must remain in
doubt.
TABLE 7; Measurements for Macropus (Prionotem-
nus) narada
SP. NOV.
Specimen
DP3
M>
M 2
M 2
F9105,
holotype
F9106
F9107
9-lx7-2
90x7-1
10-lx7'9
—
12-5x9-0
Macropus (?Prionotemnus) dryas
(De Vis, 1895)
(Plate 26, fig. 6; Plate 27, figs 1-3)
Halmaturus dryas De Vis, 1895, pp. 109-11, pi. 17,
figs. 11-5.
Macropus (Prionotemnus) dryas (De Vis): Barth-
olomai, 1975, pp. 229-32, pi. 25, figs. 1-4
(synonymy).
Macropus sp. c.f. M. dryas (De Vis): Archer in
Archer and Wade, 1976, p.391, pi. 57, figs. c-d.
Maiiriai Examined: F7823, partial right
mandibular ramus with P^-Mj. P 3 excavated,
juvenile (figd. Archer. 1976, pi. 57, figs. c-d). F9078,
isolated right M^; F9079, isolated right M^; F9080,
isolated partial left M 2 : F908I, isolated left Mt’ F9082,
isolated right M^; F9083, partial isolated left M^
F9084. isolated left Ph F7828, isolated left Ph F7780,
partial left maxilla with M--M^, juvenile; F7781,
isolated left Ph F9085. partial left mandibular ramus
with M 1 -M 3 , aged; F9086, partial right mandibular
ramus with P 3 -M 4 ; adult; F9087. partial right
mandibular ramus with M 2 , juvenile; F9088. partial left
mandibular ramus with base of I), P 3 -M 1 , juvenile;
F9089, partial left mandibular ramus with M 3 -M 4 ,
adult; F9090, isolated left M^; F909I, isolated right P 2 ;
F9092, isolated partial right M^; F9093, isolated partial
left M^; F9094, isolated left M 2 ; F9096, isolated right
DP 3 ; F9097. isolated right Py. F9098, isolated partial
right M 2 : F9103, isolated left M'; F9095, isolated
partial left DP 3 ; F9099, isolated right M^: F9100,
isolated partial left M'; F9I01, isolated partial right
DP 3 ; F9102, isolated left P 3 ; F7790, isolated right P^;
F7782, partial right mandibular ramus with M] M 3 ,
juvenile; F7783, isolated right P 3 ; F7791 isolated left
P^.
DISCUSSION: The sample of Macropus
(? Prionotemnus) dryas (De Vis) from the
138
MEMOIRS OF THE QUEENSLAND MUSEUM
Allingham Formation is numerically the largest
among macropodids from the deposits, suggesting
that this species was the most abundant grazing
herbivore in the fauna. Bartholomai (1975)
revised M. dryas, previously recorded only from
the late Pliocene Chinchilla Sand in the western
Darling Downs, where the species was well
represented but not dominant within the
assemblage present.
Observed measurements for cheek teeth in the
Allingham sample often fall towards the lower
observed limits or even outside the lower limits
of comparable teeth in the Chinchilla Sand
material. No other morphological differences are
evident which might be considered significant in
the separation of the Allingham Formation sample
from that in the Chinchilla Sand. With the larger
sample now available, the possible structural
distinctions noted by Archer in Archer and Wade
(1976) arc considered to be within the range of
variation for M. dryas. or are likely to be
encountered in the species, bearing in mind the
variation exhibited by modern species of
Macropus (Bartholomai 1971).
Two specimens, F7790 and F7791, have been
referred to M. dryas. both of which arc P-. This
tooth was not present in the available Chinchilla
Sand material at the time of Bartholomai’s ( 1 975)
revision. These possess a single vertical set of
labial and lingual ridges transecting the
TABLE 8: MFAStREMnNTS for Mandibles or Macropus {?Prionotemnus} drvas (De Vis.
1895)
Specimen
P2
DP 3
P 3
M,
M 2
M 3
M 4
F7823
8-5x3-8
8-8x5-0
11-3x4-0
9-9x6-5
F9078
—
—
—
— ■
12-0x7-2
—
—
F9079
—
—
—
—
11-6x7-0
—
—
F9080
—
—
—
—
— x8-3
—
—
F9081
—
—
—
—
12-5x80
—
—
F9082
—
—
—
—
11-9x7-1
—
—
F9085
—
—
—
8-8x6-3
10-0x7-6
12-2x8-2
—
F9086
—
—
11-3x3-9
9-lx5-7
1 1 -Ox—
12-7x8-0
13-8x8-1
F9087
—
—
—
—
11 -5x—
—
—
F9088
—
—
10-5x3-6
8-7x5-9
—
—
—
F9089
—
—
—
—
—
12-4x8-2
13-4x8-3
F9090
—
—
—
—
—
13-0x8-4
—
F9091
7-6x3-9
—
—
—
—
—
—
F9092
—
—
—
—
—
— x8-3
—
F9093
—
—
—
—
— x7-9
—
—
F9094
—
—
—
—
12-2x7-5
—
—
F9096
—
8-6x4-3
—
—
—
—
—
F9097
—
—
11-7x4-2
—
—
—
—
F9095
—
8-0x4-4
—
—
—
—
—
F9099
—
—
—
—
11-9x7-7
—
—
F9101
—
— x4-5
—
—
—
F9102
—
—
11-4x3-9
—
—
—
—
F7782
—
—
—
—
11 -7x—
13-6x7-7
—
F7783
—
11-2x4-0
—
—
—
—
TABLE 9:
Measurements for
Maxillae of Macropus
1895)
(?Prionolemnus} dryas
(Dl: Vis.
Specimen
P2
DP3
P3
M'
M2
M3
M4
F9084
_
13-7x6-0
F7828
—
—
13-8x6-4
—
F7780
—
—
—
—
11-1x9-3
11-8x9-4
F7781
■ —
—
13-3x6-3
—
F9103
—
—
—
10-3x7-9
F9i00
—
—
— x8-2
F7790
8-2x5-4
—
F77gi
8-3x5-l
—
__
—
—
—
—
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
139
longitudinal crest, have a low hypocone and a
variably present lingual cingulum. This structure
can be reasonably W'cll developed compared with
the situation in where it is represented only by
a series of low' tubercles closely adpressed to the
base of the longitudinal crest.
Macropus (Osphranter) pavana sp. nov.
(Plate 26, figs. 1-5)
Mathriai. Examined: F9108, holotype, partial
left maxilla with DP-^-M', P-^ missing, juvenile,
Allingham Formation of mid Pliocene age. Bluff Downs
Station, Allingham Creek.
In addition, F9109, a partial left mandibular ramus
with M 2 -M 4 . adult; F9110, isolated right M-; F9111,
isolated right M^; F9112, isolated right M'; F9113,
isolated right Pb F9il4, isolated right P^; F9115
isolated right M-; F9116, isolated left P^; F9117,
isolated left M^; F7773, isolated right M^; have been
used in the description of the species.
Specific Diagnosis-. A relatively small
species with deep ramus, relatively narrow below
anterior cheek teeth; P 3 relatively elongate with
trenchant longitudinal crest transected medially
by single set of vertical labial and lingual ridges
and with crest slightly curving lingually
posteriorly. Lower molars with midlink well labiad
to midline; hypolophid with variable oblique
groove. DP^ with high secant ridge from paracone
to anterolabial limit of anterior cingulum; forelink
strong; accessory link present mid-way between
midlink and labial margin of median valley. P-^
with longitudinal crest slightly cleft at anterior
one-third and w'ith single medial set of vertical
labial and lingual ridges transecting crest;
hypocone very low; posterolingual fossette and
lingual cingulum very reduced, cingulum degener-
ating to series of tubercles anteriorly. Molars with
relatively strong forelinks; anterior molars with
slight accessory link mid-way between midlink
and labial margin of median valley; this is variably
absent in posterior molars.
Description: Upper incisors, P-, and
not preserved.
DP^ molariform, subovate in basal outline,
slightly constricted across median valley, with
metaloph much broader than protoioph. Lophs
high, slightly convex anteriorly. Anterior cin-
gulum moderately high, narrow, short, with
well-defined forelink well linguad to mid-line from
base of protoioph to cingulum; cingulum near
planar, labiad to forelink, ascending at very high
angle lingually. Strong high ridge ascends
anteriorly from paracone to anterolabial margin
of cingulum as secant crest. Well-defined, high,
strong ridge ascends posterolabially from
proloconc across median valley as midlink, uniting
with ridge from near mid-point of metaloph, below
median valley; relatively strong ridge ascends
posteriorly from paracone towards median valley,
producing slight accessory link across floor of
valley mid-way between midlink and labial
margin. Valley narrowly V-shaped labially and
lingually, near planar transversely. Strong ridge
ascends posteriorly from hypocone, curving
labially to near poslerolabial margin of crown,
uniting with slight posterior ridge from metacone;
fossette present at midline. Base of crown swollen
slightly labially and lingually.
P^ relatively large, subovate in basal outline,
broader posteriorly than anteriorly; high,
relatively short, longitudinal crest transected by
median set of vertical labial and lingual ridges
with production of cuspule at crest; slight cleft
present along crest about one-third distance from
anterior cusp. Hypocone very low, connected to
metacone by weak anterolabial ridge; stronger
ridge curves posterolabially to meet posterolingual
ridge from metaconc, defining shallow posterolin-
gual fossette; anterior ridge from hypocone
ascends abruptly forming low cingulum poster-
olingually, this degenerating anteriorly into basal
tubercles, lingual basin very restricted.
M'<M molars subovate in basal outline, very
slightly constricted across median valley; lophs
high, somewhat rotated with metaloph crest more
convex anteriorly than protoioph in unworn teeth;
metaloph broader than protoioph in anterior
molars. Anterior cingulum relatively high, narrow,
short; well-defined forelink present linguad to
mid-line, between base of protoioph and equally
well-developed, short, transverse ridge across
cingular shelf. Labial moiety of cingular shelf near
planar, lingual moiety ascending at high angle
from forclink; slight ridge ascends anteriorly from
paracone to anterolabial margin of cingulum.
Strong, high ridge curves posterolabially from
protocone to unite with strong ridge from near
centre of hypoloph as midlink; posterior ridge
from paracone very reduced; low accessory link
developed across median valley and mid-way
between midlink and labial margin, this
occasionally lacking by M^; median valley near
planar transversely, sharply V-shaped longitudin-
ally. Anterior ridge from melacone to accessory
link very weak in M', sometimes lacking by M^.
Strong, slightly flared ridge curves posterolabially
from hypocone to posterolabial base of crown.
140
MEMOIRS OF THE QUEENSLAND MUSEUM
there uniting with weak posterior ridge from
mctacone; fosselte developed at mid-line; base of
crown usually slightly swollen.
Mandible imperfectly known. Ramus deep,
relatively narrow below anterior cheek teeth;
post-alveolar shelf elongate with angle poorly
developed; ventral margin of ramus rounded
posterior to symphysis; diagastric ridge and
process very weak; diagastric process separated
from base of angle by very shallow post-diagastric
sulcus, bounded above by very shallow diagastric
fossa: this fossa separated above from broad
shallow depression opening posteriorly into
pterygoid fossa; mandibular foramen large;
masseteric crest raised to about level of occlusion
of check teeth.
Lower incisor, and DP 3 are unknown.
P 3 relatively elongate, subovate in basal outline,
somewhat broader posteriorly than anteriorly.
Longitudinal crest trenchant with anterior cuspid
well-defined but with posterior cuspid less
well-defined; crest ascends posteriorly, being
transected mesially by set of weak, vertical, labial
and lingual ridges with production of low cuspule
at crest. Posterior extension of crest curves very
slightly lingually before descending to crown base.
Anterior surface from anterior cuspid near
vertical. Base of crown narrowly swollen labially,
lingually and anteriorly.
M|<M 3 < M 3 <M 4 ; molars subovate to subrec-
tangular in basal outline, slightly constricted
across talonid basin; lophids high, convex
posteriorly, somewhat anteriorly rotated in labial
view, with hypolophid broader than protolophid in
M| and Mt and slightly narrower in M 3 and M4.
Trigonid basin relatively broad, its length
approximately equalling distance between lophids.
Forelink high, strong, curving anterolingually
from proloconid across trigonid basin to point
labiad to mid-line on high anterior cingulum;
trigonid basin slopes labially and lingually from
forelink and posteriorly from cingulum; anterior
cingulum more expanded anterolingually; very
slight ridge descends anteriorly from metaconid;
slight anterolabial fossette formed in trigonid.
Midlink high, strong, curving anterolingually from
hypoconid to unite with well-developed ridge from
proloconid above talonid basin; junction occasion-
ally markedly flexed; variable, high, accessory
ridge present running anterolingually from
hypoconid part of midlink. Talonid basin narrowly
V-shaped labially, broadly U-shaped lingually,
slightly descending labially and lingually from
midlink; floor of talonid sometimes with low
transverse fold labially and lingually from
midlink. Posterior of hypolophid with moderate.
near-vertical groove, rarely pocketed towards its
base. Hanked by low, vertical ridges. Crown
somewhat Hexed about labial extremity of talonid
basin.
DISCUSSION: Taxonomic separation of
species of Macropus Shaw is frequently extremely
difficult on the basis of dental morphology alone
and normally, single characters are of little use
for this purpose. This certainly is true of the
material from the Allingham Formation, here
referred to Macropus {Osphranter) pavana sp.
nov.
Archer in Archer and Wade (1976) correctly
assigned part of the material to M. (Osphranier)
suggesting it should be compared with M. woodsi
Bartholomai (1975) a species from the Chinchilla
Sand with which M. pavana has obviously close
relationships. With the larger sample now
available as a result of subsequent collecting,
separation of the material from M. woodsi and
from M. (Osphranier} pan De Vis, 1895, also from
the late Pliocene Chinchilla Sand has been
possible.
M, pavana is smaller than M. pan and has its
P^ relatively narrower, with a lower hypocone and
much reduced lingual cingulum and basin.
Development of an accessory link across the
median valley in upper molars appears restricted
to M' and sometimes M^ whereas this structure
frequently occurs in incipient for^m in posterior
molars in M. pan. P 3 is similar in these species,
but that in M. pavana lacks the prominent
posterolingual cuspule present in M. pan.
Compared with M. woodsi. in which the upper
permanent prcmolar has yet to be described, M.
pavana. is of similar size but upper molars lack
well-defined accessory links across the median
valleys of posterior molars P 3 is larger than in M.
woodsi. and rather than possessing a trifid,
crescentic longitudinal crest, has a much
straightcr, trenchant structure. Lower molars
possess a stronger midlink which includes a much
greater contribution from near the protocone and
the posterior surface of the protolophid.
Three isolated premolars from the Allingham
Formation mentioned by Archer in Archer and
Wade (1976) as resembling M (Osphranier) have
been assigned elsewhere within Macropus in this
paper, with the exception of F7789. which is
clearly of the M. (Osphranier) type. This specimen
has a bifid, more curved longitudinal crest than
in the referred P 3 of M. pavana. It also possesses
an anterolingual ridge from the posterior cuspid.
It is possible that additional M. (Osphranier)
species may be present in the Allingham
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
141
TABLE 10: Measurhments for Maxii.lae of
Macropus {Osphranter} pavana SP. NOV.
Specimen
DP3
P^
M'
M 2
F9I08,
holotype
10-2x7-5
11-9x9-8
F9110
—
—
—
13-2x10-7
F91 13
—
9-8x5-6
—
—
F91 14
—
9-3x5-7
—
—
F9112
—
—
10-4x8-5
—
F9115
—
—
—
12-6x9-6
TABLE 11: Measurements for Mandibles of
Macropus {Osphranter} pavana SP. nov.
Specimen
P, M,
M.
M 3 M 4
F9109
13-9x9-2
16-2x10-717-4x10-0
F9116
8-6x5-0 —
—
— —
F9111
— —
13-7x8-6
— —
F9117
— —
—
16-4x10-1 —
F7773
— —
—
— xlO-3 ^
Formation, and whether this specimen reflects
extreme variation in M. pavana or whether it
represents an additional taxon must remain
unresolved until further collections are forthcom-
ing from the deposits.
CONCLUSIONS
Comparison of the macropodids from the
Allingham Formation with those from other
Pliocene deposits in Australia is largely limited by
the nature of other occurrences, the extent to
which faunal elements have been researched and
the lack of diversity in the faunas yet reported.
The notable exception is the Chinchilla Sand in
the western Darling Downs, south-eastern Queens-
land, a formation believed to be of late Pliocene
age. This provides a substantia! basis for
comparison, although other deposits in Victoria
and in the Tirari Desert of central Australia give
tantalizing glimpses of elements present in those
parts of the continent in Pliocene times. The Awe
fauna in Papua New Guinea (Plane 1967)
contains taxa which also warrant attention in this
regard. An undescribed, presumably Pliocene
fauna from Merriwah in New South Wales
contains reasonable diversity and will prove
interesting for comparison when study on it has
been completed.
The Macropodidae from the Chinchilla Sand
were studied originally by De Vis (1895).
Progressive revisions of the different taxa
represented appear in numerous papers by
Bartholomai (1963, 1966, 1967, 1973a, 1973b,
1976). A total of 1 1 macropodid species has been
recorded, although as with the Bluff Downs fauna,
fragmentary remains suggest the presence of
additional forms. Among sthenurines, the most
common species is Slhenurus antiquus. Tropo-
sodon minor, Protemnodon chinchillaensis, P.
devisi. Macropus {Osphranter} pan. M. (O.)
woodsi and M. (?Prionotemnus) dryas are very
common among the Macropodinae. Other
macropodids are present in relatively low
numbers. Similar to the Allingham Formation,
potoroincs have yet to be located.
The only macropodids common to both faunas
are T minor and M. f? PJ dryas. Of these, T.
minor is known from wide temporal and
geographical ranges and is not considered
particularly useful in correlation. M. (? P.) dryas,
however, has been recorded previously only from
the Chinchilla Sand. This fact, taken in
conjunction with the known minimal age for the
Allingham Formation, the presence of a
presumably more primitive Troposodon in those
sediments and the general similarities of the taxa
where comparisons are possible, is supportive of
the laic Pliocene age currently attributed to the
Chinchilla Sand. Close relationships are apparent
between Chinchilla and Allingham diprotodontids
(Archer 1976).
Detailed comparisons have been providco
within the text above with taxa described from
other Australian and Papua New Guinea deposits.
For the most part, conclusions other than
taxonomic cannot be drawn at this time.
Although it is apparent that grazing ma-
cropodids predominated in the assemblage present
in the Allingham Formation, suggesting that open
sclerophyll and open grasslands habitats were
present in the area during deposition, detailed
palaeoecological assessment should await results
of studies on other groups represented.
LITERATURE CITED
Arc HER. M., 1976. Bluff Downs local fauna. In Archer,
M. and Wade, M., Results of the Ray E. Lcmley
Expeditions, Part 1. The Allingham Formation and
a new Pliocene vertebrate fauna from northern
Australia. Mem. Qd Mus. 17: 379-97.
Archer. M. and Waix-, M., 1976. Results of the Ray
E. Lcmley Expeditions, Part 1. The Allingham
Formation and a new Pliocene vertebrate fauna
from northern Australia. Mem. Qd Mus. 17 :
379-97.
Bartholomai. A., 1963. Revision of the extinct
macropodid genus Sthenurus Owen in Queensland.
Mem. Qd Mus. 14 : 51-76.
1966. The type specimens of some of De Vis’ species
of fossil Macropodidae. Mem. Qd Mus. 14 :
115-25.
142
MEMOIRS OF THE QUEENSLAND MUSEUM
1967. Troposodon. a new genus of fossil Ma-
cTopodidae (Marsupialia). Mem. Qd Mus. 15 :
21-33.
1971. Morphology and variation in the cheek teeth
in Macropus gigantcus Shaw and Macropus agilis
(Gould). Mem. Qd Mus. 16: 1-18.
1973a. The genus Protemnodon Owen (Marsupialia:
Macropodidae) in the upper Cainozoic deposits of
Queensland. Mem. Qd Mus. 16: 309-63.
1973b. Fissuridon pearsoni. a new fossil macropodid
(Marsupialia) from Queensland. Mem. Qd Mus.
16 ; 365-8.
1975. The genus Macropus Shaw (Marsupialia:
Macropodidae) in the upper Cainozoic deposits of
Queensland. Mem. Qd Mus. 17: 195-235.
1976. The genus Wallahia Trouessart (Marsupialia:
Macropodidae) in the upper Cainozoic deposits of
Queensland. Mem. Qd Mus. 17 : 373-7.
Campwi ii. C. R., 1973. A new species of Troposodon
Bartholomai; from the early Pleistocene Kanunka
fauna. South Australia (Macropodidae: Mar-
supialia). Rec. S.A. Mus. 16 : 1-18.
Dr Vl.s. C. W.. 1895. A review of the fossil jaws of the
Macropodidae in the Queensland Museum. Proc.
Linn. Soc. N.S.W. 10 : 75 -133.
Mr RRii HI S. D., 1965. Two species of the extinct genus
Sthenurus Owen (Marsupialia: Macropodidae)
from south-eastern Australia, including Sthenurus
gilli sp. nov. J. R. Soc. W. Au.st. 48: 22-32.
1967. South-western Australia occurrences of
Sthenurus (Marsupialia: Macropodidae) including
Sthenurus hrownei sp. nov. J. R. Soc. W. Aust. 50:
65-79.
OwKN. R. 1877. Researches on the Fossil Remains of
the Extinct Mammals of Australia with a Notice
of the Extinct Marsupials of England. 2 vols. XV
+ 522 pp. (J. Erxlebcn: London).
Pi AN[\ M. D., 1967. Stratigraphy and vertebrate fauna
of the Otibanda Formation, New Guinea. Bull. Bur.
Min. Resour. 86: 1-64.
1972. A New Guinea fossil macropodid (Marsupialia)
from the marine Pliocene of Victoria, Australia.
Mem. Nat. Mus. Vic. 33: 33-6.
Tedford, R. H., 1966. A review of the macropodid
genus Sthenurus. Bull. Dept. Geol. Uni. Calif. 57:
1-72.
Ternbuh, W. D. and Lundeuus. E. L., 1970. The
Hamilton fauna. A late Pliocene mammalian fauna
from the Grange Burn, Victoria, Australia.
Fieldiana: Geology 19 : 1-163.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 23
Fig 1 A-B: Sthenurus sp., F9104, stereopair of occlusal view of partial
right M", x2.
Fig 2A-B: Troposodon minor (Owen), F905J, stereopair of occlusal
view of isolated right DP\ x2.
Fig. 3A-B: Troposodon minor (Owen), F9046, stereopair of occlusal
view of isolated left P^, x2.
Fig 4A-B: Troposodon minor (Owen) F9050, stereopair of occlusal
view of isolated right Mj, x2.
Fig. 5A-B; Troposodon hluffensis sp., nov., F9058, lateral and
stereopaired occlusal views of isolated right P2, x2.
Fig. 6A-B: Perrogale sp., F7794. stereopair of occlusal view of isolated
left M., x2.
Fig, 7, 7A'B: Troposodon bluffensis sp. nov., F9057, lateral and
stereopaired occlusal views of isolated right M 4 , x 2 .
Fk:. 8, 8A-B: ? Troposodon hluffensis sp. nov., F9060, lateral and
stereopaired occlusal views of i.solalcd right P3, x2.
Fig 9, 9A-B: Troposodon bluffensis sp. nov., F9054. holotype. lateral
and stereopaired occlusal views of isolated right M‘*, x2.
Fig 10, lOA-lOB: Troposodon bluffensis sp. nov., F9059, lateral and
stereopaired occlusal views of isolated left P2, x2.
BARTHOLOMAI; ALLINGHAM FORMATION MACROPODIDAE
Plate 23
MEMOIRS OF THE QUEENSLAND MUSEUM
Platf 24
Fig. 1. lA-B: Protemnodon snewini sp. nov., F9074, lateral view y}h
and stereopaired occlusal views xl of partial adult skull.
Fig 2, 2A-B; Protemnodon snewini sp. nov., F7788, lateral and
stereopaired occlusal views of isolated left P^, x2.
Fig 3, 3A-B: Protemnodon snewini sp. nov., F7824, lateral and
stereopaired occlusal views of isolated right P 2 , x2.
Fig. 4: Protemnodon snewini sp. nov., F906I, holotype, lateral view
of right mandibular ramus, x'/ 2 .
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
Plate 24
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi ATI- 25
Fig. lA-B: Proiemmuion snewini sp. nov., F9061, hololype, stereopair
of occlusal view of right mandibular ramus with 1], P3-M4, xl.
Fig. 2, 2A-B: Macropus (Prionoiemnus) narada sp. nov., F9105,
holotype, lateral and stereopaired occlusal views of partial left
maxilla with DP* M', xl.
Fig. 3, 3A-B: Macropus (Prionoiemnus) narada sp. nov., F9107,
lateral and stereopaired occlusal views of isolated right M^, x2.
BARTHOLOMAl: ALLINGHAM FORMATION MACROPODIDAE
Plate 25
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate-: 26
Fig. 1, lA-B: Macropus (Osphranter) pavana sp. nov., F9108,
holotype, lateral and stereopaired occlusal views of partial left
maxilla with DP-^-M', xl.
Fig. 2, 4A-B: Macropus (Osphranter) sp., F7789, lateral and
stereopaired occlusal views of isolated left P3, x2.
Fig 3, 2A-B: Macropus (Osphranter) pavana sp. nov., F91 14, lateral
and stereopaired occlusal views of isolated right P-\ x2.
Fig 4, 3A-B: Macropus (Osphranter) pavana sp. nov., F91 16, lateral
and stereopaired occlusal views of isolated left P3, x2.
Fig. 5, 5A-B: Macropus (Osphranter) pavana sp. nov., F7773, lateral
and stereopaired occlusal views of isolated right M3, x2.
Fig. 6, 6A-B: Macropus {? Prionotemnus) dryas (De Vis), F9084,
lateral and stereopaired occlusal views of isolated left P-^, x2.
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
Plate 26
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 27
Fig, 1. lA-B: Macropus {? Prionotemnusj dryas (De Vis). F9086,
lateral and occlusal views of partial right mandibular ramus with
P3-M4, xl.
Fig. 2, 2A-B: Macropus f? Prionotemnusj dryas (De Vis). F7780,
lateral and stereopaired occlusal views of partial left maxilla with
M— M3, xl.
Fig. 3, 3A-B: Macropus (? Prionotemnusj dryas (De Vis), F7790,
lateral and stereopaired occlusal views of isolated right P2, x2.
BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE
Plate 27
Mem. Qd Mus. 18(2): 145-9 [1978]
THE ROSTRUM IN PALORCHESTES OWEN
(MARSUPIALIA: DIPROTODONTIDAE)
RESULTS OF THE RAY E. LEMLEY EXPEDITIONS, PART 3
Alan Bartholomai
Queensland Museum
ABSTRACT
The rostral area of the skull is examined in all described species of the genus Palorchestes
Owen and is shown to be characterized by massive dorsal excavation, associated with recorded
and inferred reduction of the nasals and extreme elongation of the anterior of the palate.
These features, taken in conjunction with the development of very large infraorbital foramena
and the anteriorly directed facial area are interpreted as supporting a tapir-like proboscis in
P. painei and probably in all known species of Palorchestes. The symphysis of the lower jaws
is elongate in all forms and is narrow and deeply channelled dorsally, suggesting the presence
of a long, flexible tongue which could have acted in conjunction with a proboscis in facilitating
ingestion of herbage.
The genus Palorchestes is reasonably well-
known in literature but has never been strongly
represented in collections by specimens which
show details of the cranial morphology.
Palorchestes was defined by Owen (1874) as a
macropodid and it was not until much later
(Woods 1958) that its current systematic position
within the Diprotodontidae was recognized. Three
species have now been defined within the genus,
these comprising the type species, P. azael Owen
1874, from Pleistocene sediments, P. parvus De
Vis 1895, from the Chinchilla Sand of late
Pliocene age and P. painei Woodburne 1967, from
the Waite Formation of late Miocene or early
Pliocene age.
Stirton et al. (1967) have attempted a
phylogeny of the Diprotodontidae and have used
Tate’s (1948) subfamily, the Palorchestinae, for
Palorchestes and for Ngapakaldia Stirton and
Pitikantia Stirton from the Etadunna Formation
of late Oligocene or early Miocene age. Pitikantia
does not have crania preserved which present the
rostral area. It is certain that Ngapakaldia did
not have the type of cranial specialization
observed in Palorchestes, and from what is
currently known of Pitikantia, it is unlikely that
dorsal excavation of the rostrum was a feature of
that genus also.
The presence of an extensive modification to the
rostral area of the skull in Palorchestes has been
widely noted (Woods 1958; Woodburne 1967) but
it was not until preparation of a near-complete
cranium of P. painei. collected from the Waite
Formation during the Ray E. Lcmiey expedition
by the Queensland Museum in 1974, that the full
extent of this structure and its possible
significance became apparent. Less complete, but
equally convincing material for both P. azael and
P. parvus in the Queensland Museum collections
also indicates the presence of comparable rostral
structure in those species.
The Rostrum in Palorchestes Painei
Woodburne 1967
A reasonably complete description of the
cranium of P. painei is provided in Woodburne
(1967). The rostral area in QMF 9179 and QMF
9178 comprises more complete material on which
the following supporting and supplementary
comments are provided.
The rostral area is very elongated and relatively
narrow and generally tapers anteriorly in lateral
view. The incisor tooth row is projected more
ventrally than the horizontal plane of the cheek
teeth. Although slightly incomplete, the premaxil-
lae are broader transversely than the anterior of
146
MEMOIRS OF THE QUEENSLAND MUSEUM
the maxillae giving the premaxillary alveolar
margin a somewhat spatulate appearance. The
palatal sutures between the premaxillae and
maxillae are clearly defined, extending anter-
olalerally from the posterior border of the incisive
foramina along their lateral borders then turning
abruptly posterolaterally to the diastemal margins
well posterior to the foramena, then continuing
laterally, posterodorsally to near the narial
notches. The premaxillae thus form broad wedges
bordering the anlerodorsal margins of the narial
opening. Low, dorsomedial premaxillary spines
are present, approximately above the level of
r~.
Ventrolateral diastemal margins are angular
and the lateral surfaces posteriorly arc gently
concave above the margins. In occlusal view, the
incisors present a broadly U-shaped outline.
Dorsal margins of the thin posterior processes of
the prcmaxillac are acute anteriorly, becoming
sharply rounded and elevated to a higher angle
to the horizontal. These margins are slightly
rotated mcsially. The anterior of the palate is
moderately concave transversely with a medial
sulcus extending anteriorly from the incisive
foramena.
The maxillae form the bulk of the remainder
of the rostrum. The palate is gently concave
longitudinally posterior to the premaxillary-max-
illary suture and is also concave transversely with
an accentuated medial sulcus extending from the
incisive foramena posteriorly to about the level of
M“. Laterally, the maxillae comprise long, deep
bones wedging anteriorly to the angular diastemal
margins. Towards the orbital margins, the lateral
surfaces of the maxillae are sharply curved
laterally, forming broad, near-vertical, transverse
surfaces comprising an anteriorly directed facial
region. These surround extremely large, eliptical,
infraorbital foramena on each side, but that on
the right side is much larger than on the left in
QM F9179. However, in QM F91 78 each foramen
is relatively smaller. Lengths of the infraorbital
canals arc very short, opening posteriorly into
variable, but large maxillary foramena.
The lacrymals extend moderately onto the
facial region. Low frontal crests extend laterally
as more pronounced ‘brow' ridges, associated with
the frontals, nasals and lacrymals, with the
development of well-defined ventrolateral grooves
anteriorly below the ridges. The nasals are very
short, with nasal spines extending only slightly in
advance of the narial notches.
The nasal cavity is deep anteriorly and very
deep posteriorly, surrounded anteriorly by the
premaxillae and posteriorly by the maxillae.
premaxillae and nasals. A bony nasal septum is
present posteriorly. The floor of the cavity is
occupied by fragments of bone, largely represent-
ing the remains of the vomer and parts of the
septum. Anteriorly, the vomer is strongly and
moderately deeply, inverted T-shaped in
section.
The Rostrum in Palorchestes Parvus De Vis
1895
Woods (1958) has provided a basic description
of the known cranial remains of P. parvus and
the following comments, based on QM F789,
amplify that study.
The rostrum is very elongate. It is relatively
broader and tapers to a greater extent anteriorly
in lateral view than in P. painei. The premaxillae
are much broader transversely than in P. painei
but are apparently not much broader than the
anterior of the maxillae. The anterior alveolar
border is flat and the incisors form a nearly
rectilinear occlusal surface. Anterior to moderate-
ly high, medial, premaxillary spines, the dorsal
surface is considerably extended and shelf-like,
curving gently laterally to the alveolar margins.
Above the level of the incisive foramena and
posterlateral to the premaxillary spines, the dorsal
surfaces are deeply and variably excavated,
leaving sharp, thin flanges of bone lining the nasal
cavity anteriorly; lateral to these excavations, the
bones are more robust, being broadly rounded
dorsally at the anterior of the excavations and
more acute posterolaterally. The palate is broadly
concave transversely at the alveolar margin, but
broad, ventral protruberences occur lateral to the
•incisive foramena in the area of the premaxill-
ary-maxillary suture. Diastemal margins are
flanged posterior to P. becoming less so towards
the maxillae. The premaxillary-maxillary sutures
extend anterolaterally from near the posterior of
the incisive foramena, then turning abruptly
posterolaterally to the diastemal border and
proceeding posterodorsally leaving thin wedges of
bone extending as the dorsal rims of the narial
opening. These wedges are considerably shallower
anteriorly than in P. painei: the dorsal rims are
somewhat directed mcsially. Little of the anterior
of the maxillae remains.
Although the infraorbital foramena are not
preserved, part of one maxillary foramen is
present, showing it to have been wide and
presumably large. The infraorbital canals must
have been short. The dorsal surface of the cranium
is unknown, as is the anterior surface of the orbital
area. No nasals are preserved, but the structure
BARTHOLOMAI: THE ROSTRUM IN PALORCHESTES
147
Fig. 1; Cranial remains in Palorchestes. A, P. painei (F9179), B, P. parvus (F789); C, P. azael (F3837), showing
lateral extent of rostral modification. Scale in cms.
of the premaxillae suggests these must have been
extensively retracted.
The Rostrum in Palorchestes Azael Owen
(1874
Owen (1874) describes the nature of the
rostrum in P. azael. Existing descriptions are
supplemented by the juvenile specimen, QM
F3837.
The rostrum is elongate and relatively narrow,
more like that in P. painei than in P. parvus.
The premaxillae are broad transversely, being
much broader than the anterior breadth of the
maxillae. The incisive alveolar border is broadly
rounded and the upper incisors present a broad
U-shaped occlusal outline. The medial premaxil-
lary spines are relatively low and the dorsal
surfaces anterior to these are broadly convex
transversely to the alveolar margins. Posterior to
P, the disastemal margins are flanged for a short
distance, then rounded from about the level of the
premaxillary-maxillary sutures. The palate is
anteriorly shallowly concave transversely, with a
deep, medial sulcus extending to the alveolar
margin from the incisive foramena. The foramena
are positioned well back and may be confluent.
The premaxillary-maxillary sutures extend
anterolaterally from near the posterior of the
incisive foramena, then turn abruptly posterolater-
ally continuing posterodorsally from the diastemal
margin. The posterior extensions of the dorsal
parts of the premaxillac exist as broad wedges of
bone capping the maxillae and bordering the
narial opening, similar to those in P. painei. These
are somewhat directed mesially. The palate has
a relatively deep sulcus medially, posterior to the
incisive foramena. Below the premaxillae
processes, the maxillae anteriorly are laterally
concave. Hollowing of the portion remaining of
the facial area suggests that transverse, planar
areas existed anterior to the orbits. No trace
remains of the infraorbital foramena, canals or
maxillary foramena. Although dorsal aspects of
the skull are unknown, the nasals are interpreted
as having been extensively retracted, based on the
observed morphology of the premaxillae and the
narial opening.
148
MEMOIRS OF THE QUEENSLAND MUSEUM
The Mandibular Symphysis in Palorchestes
Owen 1874
Woodbnrne (1967, figs. 23-4) has illustrated a
specimen with a near complete symphyseal area
in P. painei. The area is elongate, narrow, but
deeply channelled, flaring anteriorly to accom-
modate large I,. The symphysis is also
characterized by general down flexing of the
anterior extremity of the dentaries which imparts
a broadly longitudinally convex curviture to the
dorsal symphyseal surface. Although this area in
P. parvus is known only from a single specimen
with no preserved teeth (QM F9I80), no doubt
exists regarding its identity. The structure is very
similar to that in P. painei. In P. azael, a similar
situation exists. The specimen QM F774, figured
by Woods (1958, fig. 3), has a nearly complete
symphyseal area. Although the ventral border is
generally downflexed with regard to the bases of
the rami, the anterior flexing present in both P.
painei and P. parvus is lacking.
DISCUSSION
Although the dorsal surface of the cranium has
not, as yet, been recovered in either P. parvus or
P. azael, it is reasonable to assume from the
known structure in P. painei and the morphology
that is represented in the later material, that the
rostral area of the skull in all three species was
essentially similar. Detailed differences have been
noted above, but none of these is considered to
have had any marked effect on the general rostral
structure or its overall functional significance.
The massive dorsal excavation of the rostrum
is computable with the presence in life of either
an extensive rhinarium with anterodorsally
directed nostrils or a proboscis, possibly similar to
that in the eutherian tapiroids. Of these
interpretations, the latter is considered most likely
for the following reasons. The vomer apparently
carried a cartillagenous septum well anteriorly,
consistent with terminal nostrils directed anterior-
ly; the facial area of the skull is very flattened
close to the rostrum and directed almost
perpendicularly and at right angles to the axis of
the skull; a prominent groove is present below the
‘brow' ridge and this, together with the preceeding
point are both features presumably associated
with the implantation of strong muscles; the
variable, but large size of the infraorbital
foramena, infraorbital canals and maxillary
foramena are associated with the passage of an
expanded infraorbital artery, vein and nerve,
sufficient to supply a structure much larger and
more functional than a rhinarium.
The degree of convergence w'ilh eutherian
animals like tapiroids and litopterns is only
superficial and Palorchestes has achieved its
specialized cranial structure largely through
modification of the premaxillae, rather than
through maxillary modification. The deep, curved
sulcus around the narial notch in Tapirus has been
only partly achieved in Palorchestes, while the
rounded smooth anterodorsal aspect of the
premaxillae has not been duplicated at all. Nasals
in Palorchestes are retracted to a greater extent
than in Tapirus suggesting that any proboscis in
Palorchestes was not as well supported.
Fig, 2: Reconstruction of head of Palorchestes, based mainly on P. painei.
BARTHOLOMAI: THE ROSTRUM IN PALORCHESTES
149
Retraction of the nasals in marsupials has also
been reported by Bartholomai (1973) in the
macropodids Protemnodon roechus Owen and
possibly also P. brehus (Owen). These animals,
however, are presumed to have possessed large
rhinaria rather than functional proboscises.
The morphology of the mandibular symphysis
in Palorchestes suggests that the tongue must
have been long, narrow and Hexible. Associated
with the long diastema and the possible presence
of a proboscis, these features w'ould have been of
benefit to Palorchestes in the gathering and
ingestion of herbage before cropping by the broad
series of upper and lower incisors.
The apparent consistency of these features and
the peculiarity of the rostral morphology in
particular, in Palorchestes suggest that the
ancestry of Palorchestes was probably not
associated with either Ngapakaldia or Pitikantia
from the late Oligocenc or early Miocene
Etadunna Formation (Stirton 1967). Presumably
these forms are closer to the generalized structural
ancestors from which all three genera had evolved
independently.
LITERATURE CITED
Bartholomai, A. 1973. The genus Protemnodon
Owen (Marsupialia: Macropodidae) in the Upper
Caino/.oic deposits of Queensland. Mem. Qd Mus.
16 : 309-63.
Df Vis, C. W.. 1895. A review of the fossil jaws of the
Macropodidae in the Queensland Museum. Proc.
Linn. Soc. N.SAV. 10 : 75-133.
OwFN. R., 1874. On the fossil mammals of Australia.
Part IX. Phil. Trans. 164 : 783-803.
SriRroN. R. A., 1967. The Diprotodontidae from the
Ngapakaldi fauna. South Australia. Hull. Bur. Min.
Resour. 85: 1 44.
SriRTON. R. A., WOODBIRNB. M. O., and Pi.anr. M.D.,
1967. A phylogeny of the Tertiary Diprotodontidae
and its significance in correlation. Bull. Bur. Min.
Resour. 85: 149-60.
Tati. G. H. H., 1948. Studies on the anatomy and
phylogeny of the Macropodidae (Marsupialia).
Results of the Archbold Expeditions. No. 59. Bull.
.■Uner. Mus. Nat. Hist. 91 : 337-351.
WooDHL R\i , M. O.. 1967. The Alcoota fauna, central
Australia. An integrated palaeontological and
geological study. Bull. Bur Min. Resour. 87:
I 187.
WooD-S. J. T., 1958. The extinct marsupial genus
Palorchestes Owen. Mem. Qd Mus. 13 : 177-93.
Mem. Qd Mus. 18(2): 151-4. [1978]
AN UNDESCRIBED SPECIES OF ROCK DWELLING
CRYPTOBLEPHARUS (LACERTILIA: SCINCIDAE)
J. COVACEVICH and G. J. INGRAM
Queensland Museum
ABSTRACT
A new species of Cryptoblepharus (C. fuhnij is described from granite boulders of the
Melville Range, near Cape Melville, northeastern Queensland, and is compared with its nearest
relatives C. virgatus and C. litoralis.
The genus Cryptoblepharus Wicgmann was
resurrected and redefined by Fuhn (1969a, b) to
contain a single species, C. houtonii (Desjardin),
and its twenty-one subspecies. These subspecies
included C. houtonii virgatus which had been
described as Ahlepharus virgatus from Cooktown,
north-eastern Queensland (Garman 1901).
Ablepharus houtonii litoralis had been described
from the Innisfail area by Merlcns (1958). Cogger
(1973, 1975) elevated it to specific status {C.
litoralis) following Arnold's (1966) suggestion.
Cogger has treated virgatus as a subspecies of C.
houtonii noting that — . . some (subspecies) may
represent distinct species while others may be
minor variants . . (p. 258). Storr (1976) has
treated Ablepharus houtonii clarus (Storr 1961)
from south-western Western Australia as
Cryptoblepharus virgatus clarus. considering C.
virgatus from eastern Australia a distinct species
because this form and the Mauritius C. houtonii
(Desjardin) were unlikely to be conspecific,
confirming Carman's original description of the
species. In north-eastern Queensland, two species
of Cryptoblepharus, C. virgatus and C litoralis,
are currently recognised.
In November 1970, one of us (JC) working with
C. Tanner and T. Tcbble observed a strikingly
marked, dark Cryptoblepharus common on the
black rocks of the exposed boulders of the Melville
Range, Cape Melville, Cape York Peninsula,
north-eastern Queensland. The lizards were very
agile and alert and could be collected only with
the aid of a pistol and dust shot.
A typical specimen of C. virgatus (QM J20565)
was collected at the same time on a tree growing
amongst the boulders ^on which the dark
Cryptoblepharus was common. The latter differs
from both C. virgatus (with which it is
synchronosympatric) and C. litoralis (which
occurs only on the foreshore in north-
eastern Queensland and New Guinea) meristical-
ly, and in colour, pattern, and external
morphology. No other members of this genus have
the striking achromatic pattern of C. fuhni.
Differences observed are sufficiently distinct to
warrant recognition of this skink as a new species,
C. fuhni. C. fuhni is named to acknolwedge the
contribution to herpetology of Dr Ion Fuhn.
Cryptoblepharus fuhni
Hoior't'P!- QM J20566 Melville Range, Cape
Melville, Cape York. NE.Q. {14°]6'S, 144'G0'E).
Collected J. Covaccvich. C. Tanner and T. Tcbble. 30
Nov 1970.
pARATYPFiS: QM J20515-6, J20567-71, same data s
holotype.
Diagnosis
A long-legged, rock-dwelling Cryptoblepharus
distinguished from all other species of
Cryptoblepharus by its striking achromatic
pattern of white spots and dashes on a black
background (Fig. la). C. fuhni may be
distinguished further from C. virgatus by
midbody scale count (23-26 vs 20-23) and
number of lamellae under the fourth toe (22-26
vs 19-22); and from C. litoralis usually by the
number of lamellae under the fourth toe (22-26
vs 20-22). See fig. la, b, c and Table 1.
152
MEMOIRS OF THE QUEENSLAND MUSEUM
Distribution
Known only from the granite boulder 'black'
mountains of the Melville Range, Cape Melville,
north-eastern Queensland.
Description of Holotype
Snout-vent length (mm) 46-0. Head width
(%SVL) 7-0. Hind limb length (%SVL) 25-0. Tail
lost. No supranasals, but nasal scales divided.
Fig. !: A. Cryptoblepharus fuhni (J20569, on granite boulders, Melville Range, Cape Melville, NE.Q.)
B. Cryptoblepharus virgatus (J20565, on tree, Melville Range, Cape Melville, NE.Q.)
C. Cryptoblepharus litoralis (J20434, on granite boulders, Lizard Island, NE.Q.)
COVACEVICH AND INGRAM; SPECIES OF ROCK DWELLING CRYPTOBLEPHARUS
153
Rostral and frontonasal in broad contact.
Prefrontals large, meeting with a medium suture,
contacting the frontonasal, anterior and posterior
loreals, first supraciliary, first supraocular and
frontal. Anterior and posterior loreals large,
subequal. Frontoparietals and interparietal fused,
forming a large kite-shaped scale with a narrow
anterior half. Parietals large, forming a medium
suture along midline. Four large supraoculars, the
second ones the largest which just touch in the
midline. Supraciliaries five on each side, the first
the largest. Three enlarged upper ciliarics forming
a hood over lop of large transparent palpebral
disc. The latter covers nearly all the eye. No
moveable eyelids. Palpebral disc completely
surrounded by three rows of small scales except
for upper margin where there is only one row
between disc and upper ciliary hood. Seven upper
labials, fifth subocular. Six lower labials. Ear
aperture obvious (0*6 mm wide), tympanum
sunken, small rounded lobules around edge. Eight
preanal scales, central pair enlarged. Limbs
TABLE 1; Comparison of Colour, Pattern, Body Proportions, and Scale Counts of C. fuhni, C. virgatus
AND C. litoralis
Feature
Species
C. fuhni (fig. la)
C. virgaius (fig. lb)
C. litoralis (fig. Ic)
colour and pattern
black basically with a
striking pattern of while
spots and dashes which
vary in size and which
form two paravertebral
lines from neck to tail
base; lamellae and palmar
surfaces black.
brown basically with well
defined white latero-dorsal
lines from nostril to tail;
two black paravertebral
lines and a brownish verte-
bral line; head copper
brown; laterally black with
white speckling; lamellae
and palmar surfaces white.
black basically w'ith grey-
green speckling and
blotches which may form
indistinct latcrodorsal
bands; white speckles pre-
sent dorsally and laterally
and on legs and tail;
lamellae and palmar sur-
faces black.
hind leg length % SVL
minimum 5 1 .4
maximum 43.8
maximum 48.4, 39.02
(Mertens, 1958)
mean head % SVL
14.2
13.8
13.2
mid body scale rows
23-26
usually 20-23, 20 (Gar-
man, 1901)
23-28, 24-28 (Mertens,
1958)
lamellae under 4th toe
22-26
19-22
20-22, 18 (Range not
described by Mertens
1958)
154
MEMOIRS OF THE QUEENSLAND MUSEUM
pentadactyl, well developed and greatly overlap-
ping when adpressed. Toes long. Subdigital
lamellae black, undivided and smooth, 24 under
fourth toe. Palmar tubercles black and rounded.
Midbody scales in 24 rows. Dorsal, lateral and
ventral scales smooth; dorasls larger than vcntrals
which are larger than laterals. Colour in
preservative, a contrasting achromatic pattern of
white spotting and dashes on a black ground
colour. Head black with brownish white speckling,
labials flecked with larger speckles. Dorsal surface
of neck, trunk and tail black with a series of spots
and large dashes forming dorsolateral lines
beginning at back of eye and continuing down tail;
also two series of smaller dashes forming two
paravertebral lines from neck to base of tail.
Lateral surface of neck and trunk, black with a
series of small spots and dashes forming parallel
lines; these become a series of large dots down tail.
Dorsal and lateral surfaces of legs and toes black
with crossing barrings of white dots. Ventral
surface cream, except for hands and feet which
are black.
Description of Paratypes
As for the holotype except as follows:
Snout-vent length (mm): 35-47 (N = 7, x 40-6,
SD4-58). HW (%SVL): l3 s-15-6 (N = 5, k 14-2,
SDO-88). Lenth of hind limb (%SVL); 514-57-5
(N = 7, X 54-2, SD 2-07) tail length (%SVL):
158
(N = 1). Midbody scale rows 23-26 (N = 7, x
24- 1, SD 1 46). In three of the paratypes the frontal
narrowly contacts the fused interparietal-
frontoparietals but in J20569 the second
supraoculars form a short suture in the midline.
In the smaller paratypes there is a tendency for
the dorsolateral dashes to join to form lines, and
for the head and lateral surfaces to be suffused
with brown.
Comparison with other Closely Related
Species*
C.fuhni may be distinguished from C. virgatus
and C. litoralis by colour, pattern, body
proportions and external features. These features
are summarised in Table 1.
Remarks
Description of C. fuhni brings the number of
known lygosomid skinks restricted to bare
boulder habitats to six in Queensland. These are
Carlia coensis (Mitchell), C. mundivensis
(Broom), Carlia spp. nov. (two species, Ingram
*based on ten specimens of C. virgatus and C litoralis
(including topotypes) from the Queensland Museum
reference collection and on the type descriptions of these
two species.
and Covacevich, pers. observ.), Lampropholis sp.
nov. (Rawlinson, pers. comm.) and Crypto-
blepharus litoralis. These lygosomid skinks
share most of the following characters
when considered in relation to their congeners:
high number of midbody scales; large size; black
or near black colour, often with whitish dashes or
flecks; habit dorsoventrally flattened; prominent
eyes; large supraoculars; long limbs and digits;
high lamellae count for fourth toe; black palms
and lamellae; agility and fast movement. All
species are posturing heliotherms {sensu
Rawlinson, 1974). They do not emerge from
crevices and caverns until ambient temperatures
are high (usually between 9-10 am) and they
thermoregulate by changing body posture while
resting on exposed basking sites. They forage in
the sunlight and when temperatures are too high
(towards noon), basking ceases and foraging may
be continued in shaded areas. During the hottest
part of the day skinks are usually inactive,
sheltering until late afternoon when a brief search
for food is usually resumed.
ACKNOWLEDGEMENTS
Mr Bruce Campbell has constructively cri-
ticised this work. Miss Eleurctte Briggs prepared
the figure from photographs taken by Mr Allan
Easton.
LITERATURE CITED
Arnold. J. 1966. ‘A taxonomic study of the lygosomid
skinks of Queensland'. Unpublished M.Sc. thesis
(University of Queensland: Brisbane).
Cogger. H. G. 1973. Classification of Australian skinks.
Herpetofauna 6(2): 7-14.
1975. 'Reptiles and Amphibians of Australia’, (Reed:
Sydney).
Garman. S. 1901. Some reptiles and batrachians from
Australasia. Bull. Mus. Comp. ZooL Harv. 39;
1-14, pis. 1-2.
FtJUN. L 1969a. Revision and redefinition of the genus
Ablepharus Lichenstein, 1823. (Reptilia, Scin-
cidae) Revue roum. Biol. (Zoologies 14 : 23-41.
1969b. The 'polyphyletic' origin of the genus
Ablepharus (Reptilia, Scincidae): a case of parallel
evolution. Z. ZooL Syst. EvolForsch. 7: 67-76.
Mertens, R. 1931. Ablepharus boutonii (Desjardin)
und seine geographische Variation. ZooL Jb.
(Syst.). 61 : 63-210.
1958. Neue Eidechsen aus Australien. Senckenbere.
bioL 39: 51-5.
Rawlinson, P. A. 1974. Biogeography and ecology of
the reptiles of Tasmania and the Bass Strait area.
In W. D. Williams (ed) ‘Biogeography and ecology
in Tasmania.’ (Junk; The Hague).
Storr. G. M. 1976. The genus Cryptoblepharus
(Lacertilia, Scincidae) in Western Australia. Rec.
West. Aus. Mus. 4 : 53-63.
Mem. Qd Mus. 18(2) 157-64 [1978]
THE NATURE OF THE MOLAR-PREMOLAR BOUNDARY IN
MARSUPIALS AND A REINTERPRETATION OF THE
HOMOLOGY OF MARSUPIAL CHEEKTEETH
Michael Archer
Queensland Museum
ABSTRACT
Recent studies of dental ontogeny and abnormalities in marsupials indicate that all systems
of homology in current use are incorrect, in part because all are based on the evidently
erroneous assumption that true post-canine tooth replacement occurs in marsupials.
A new terminology is presented which accounts for all morphological and ontogenetic data,
including the apparent phenomenon of Zahnreihen in dasyurid dentitions. This concept is three
premolars, Pl-3, a deciduous first molar. Ml, and four permanent molars M2-5. Marsupials
are therefore regarded to have three premolars, five molars, and no true post-canine milk-teeth.
Some marsupials and in particular some kangaroos have six or more molars but these have
been additions to the posterior end of the molar series.
Owen (1840-5) introduced stability into dental
terminology by defining premolars as those
post-canine teeth anterior to, and including, the
posterior-most tooth having a milk-tooth predeces-
sor. Teeth posterior to this tooth were regarded
as molars. Application of this concept to
marsupials has been complicated by interpretation
of ontogenetic evidence. Spurious swellings along
the free edge of the dental lamina have sometimes
been interpreted as incipient or vestigial tooth
buds representing replacement teeth. These
interpretations have often given rise to conflicting
terminologies. More recently, ontogenetic studies
(Berkovitz 1967a, Archer 1974) of structurally
primitive marsupials have confirmed earlier
research (e.g. Woodward 1893) suggesting that
the replaced tooth. Ml (in the terminology used
here, but dP4 in the terminology of Thomas 1888,
Table 1) does not give rise to the tooth germ of
the replacing tooth P3 as it should if it were a
member of the P3 premolar tooth family. For this
reason, it has been suggested (Archer 1974) that
cheekteeth in dasyurid marsupials cannot be
classified as premolars or molars using Owen’s
(1840-5) system, and that recognition of
Zahnreihen (in the sense of Woerdman 1921) or
tooth developmental sequences may provide a
means for classification of marsupial teeth
independent of the phenomenon of tooth
replacement.
Examination of marsupial groups reveals many
differences in dental morphology and patterns of
tooth development. An attempt has been made
here to clarify these patterns using tooth
morphology and ontogeny and to interpret
homology of cheekteeth.
Terminology of crown morphology is set out
elsewhere (Archer 1975a, 1976b, 1976c) or, if
different from this, follows Bensley (1903) and
Stirton (1967). Terminology of post-canine
cheektooth number follows in part Owen (1845)
and Stirton (1955), and not Thomas (1888) whose
system I have previously used. It differs from
Stirton’s nomenclature in that the deciduous
post-canine cheektooth is called Ml. The
nomenclature used here is therefore Pl-3, Ml,
and M2-5. Owen (1845) and Stirton (1955)
regard that there are three adult premolars, PI,
P2, and P3, and the deciduous tooth in the
cheektooth row of polyprotodonts is called dP3
(see Table 1). This deciduous tooth is called Ml
in this paper. Marsupial names are used in the
sense of Ride (1970), Laurie and Hill (1954),
Clemens (1966), and Kirsch (1968). Specimen
number prefixes used are as follows: J, JM, or
F, Queensland Museum.
158
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 1: Postcanine Cheektooth Nomenclature used in This Work, and that of Thomas (1888) used
IN Previous Works, with Diagrams of Upper and Lower Cheekteeth of Seven Representative Marsupial
Groups.
Present
notat ion — ►
Upper
PI P2 P3 Ml M2 M3 M4 M5
Lower
PI P2 P3 Ml M2 M3 M4 M5
^ Example
Didelphids
'SB S3
Didelphis
Petaurids
o esj • ©ppe?
Pseudocheirus
Burramyids
^ ^ s . OOo -
- ^ - < 53 ) 0 © -
Distoechurus
Phalangerids
^
- -
Trichosurus
Diprotodontids
- -
- <3 CH)
Palorchestes
Potororoines
- m
Hypsiprymnodon
Macropodines
Dendrolagus
Thomas's 11888}
notation
PI P3 P4 dP4 Ml M2 M3 M4
PI P3 P4 dP4 Ml M2 M3 M4
TABLE 2: Notational Systems for Marsupial Post-canine Cheeckteeth inferred from
Ontogenetic Studies and Compared with Thomas (1888).
Author
Group examined
Thomas (1888)
PI
P3
P4
dP4
Ml
M2
M3
M4
Marsupials
Berkovitz (1967)
PI
P2
P3
Ml
M2
M3
M4
M5
Didelphids
Archer (1974)
PI
P2
P3
Ml
M2
M3
M4
M5
Dasyurids
Wilson and Hill (1897)
PI
P2
P3
dP3
MI
M2
M3
M4
Peramelids
Berkovitz (1968)
PI*
P2
P3
MI
M2
M3
M4
M5
Phalangerids
Bolk (1929)
PI
P2
P3
Ml
M2
M3
M4
M5
Phalangerids
Berkovitz (1966)
PI
P2*
P3
Ml
M2
M3
M4
M5
Macropodids
*Iniliate but do not form part of functional dentition.
Ontogeny
Recent ontogenetic work (Table 2) has revealed
that in the dasyurid Antechinus jlavipes there
appear to be eight post-canine tooth families each
of which has only one generation (Archer 1974).
They develop in a time sequence as two distinct
series; PI. P2, P3; and Ml, M2, M3, M4. M5
(PI, P3 P4, dP4, Ml-4 of Archer 1974). These
series or Zahreihen are also morphologically
uniform such that the teeth of the first are
premolariform and those of the second molar-
iform. Berkovitz (1967a), similarly demonstrates
that in at least one didelphid, the tooth regarded
here as P3 develops from the dental lamina
between P2 and Ml and is therefore not a
replacement tooth for Ml.
Berkovitz (1966) also shows that a similar
situation exists in at least one macropodid. It may
also occur in one phalangerid (Bolk 1929,
Berkovitz 1968) but the evidence is not clear.
Kirkpatrick (1969) demonstrates a developmental
relationship between the teeth regarded here as
, M2, M3, M4 and M5 in several
macropodids, and an apparently close relationship
between P2 and P3. He suggests M2-4 are
successional teeth in the Ml family.
These studies demonstrate Ml to be part of
a molariform cheektooth series and evidently
unrelated to P3 in at least three marsupial
tamihes, including didelphids and dasyurids which
are structurally ancestral to other marsupial
groups. ^
ARCHER: MARSUPIAL CHEEKTEETH HOMOLOGY
159
Morphology
Bensley’s (1903) important examination of
marsupial cheektooth morphology serves as a basis
for further comparisons, and in particular, a closer
examination here of the P2, Ml, P3, M2 region.
Ride (1961) realizes the importance of this region
in determining cusp homology in macropodids and
it also seems to be the important region in
interpreting homology in macropodid and
phalangerid cheekteeth.
To avoid lengthy descriptions, the juvenile and
adult P1-M5 region of the dentition of
representative marsupials are shown in outline
form in Table 1 and attention is given below only
to particular aspects of these dentitions.
Individual Cheekteeth
(P2, P3, Ml, M2)
P2: Because there has been no satisfactory
demonstration of more than three premolars in
marsupials, it is possible that P2, in all marsupials
with three premolars in the adult dentition, are
homologous teeth. In dasyurids with less than
three, it is always P3 which has been lost (Archer
1976a). In phalangeroids with less than three
upper premolars (e.g. phalangerids, macropodids,
and diprotodontids) homology of the anterior
premolar is uncertain. Berkovitz (1966) demon-
strates that in at least one macropodid the anterior
adult premolar is the second of three teeth to
develop on the dental lamina posterior to the
canine. The first tooth develops but later
disappears. However, Berkovitz (1968) demon-
strates that in at least one phalangerid, the
anterior adult premolar is the first of three to
develop on the dental lamina posterior to the
canine. The second tooth develops but does not
persist. Homology of lower antemolar teeth in
phalangeroids is extremely uncertain, most early
ontogenetic studies having misinterpreted true
milk incisor teeth for vestigial tooth families (e.g.
Woodward 1893).
Kirkpatrick (1969) has examined tooth
development in some macropodids and concludes
that P2 is a milk-tooth which is later replaced by
P3, a second generation tooth in the P2 tooth
family. This view is not supported by other
ontogenetic studies on macropodids such as those
of Berkovitz (1966) where P3 develops from the
dental lamina between P2 and MI. The
interpreted differences may result from post-
initiation degenerative changes in the dental
lamina or by shifts in relative position due to
migration of tooth buds or development of the free
edge of the dental lamina. Before the actual
homologies of P2 and P3 in macropodids can be
determined, ontogenetic studies should be carried
out on potoroine macropodids where P2 and P3
are large and the possible masking effects of tooth
migration arc reduced.
Morphology of P2 in all marsupials in which
it has not been lost is either premolariform or
caniniform. In some potoroine macropodids, and
some caenolestoids, it is a plagiaulacoid sectorial
tooth, in Phalanger it is caniniform. In no group
is it molariform.
P3: Ontogenetic evidence in didelphids,
dasyurids, phalangerids, and macropodids sug-
gests P3 is the posterior member of a
premolariform Zahnreihe. It develops from the
dental lamina anterior to Ml and posterior to
P2. Abnormal teeth interpretable (Archer 1975)
as P** (P5 of Archer 1975) in some macropodids,
and inferred to develop posterior to P^ on the
dental lamina, are also premolariform.
Morphologically, P3 is never molariform
although the posterior end may become secondari-
ly molarized in quadritubercular or lophondont
forms such as phascolarctids, macropodids, and
diprotodontids. In potoroine macropodids, caen-
olestoids, some burramyids and incipiently in some
phalangerids, P3 tends towards or is a well-
developed plagiaulacoid sectorial tooth. Broom
(1896) suggests the sectorial premolars of
Burramys (a burramyid) and Hypsiprymnodon (a
macropodid) are completely unlike those of
phalangerids because the serrations are on
opposite ends of the tooth. This observation seems
of little import in view of the fact that in other
macropodids (e.g. some Bettongia and Potorous)
the serrations occur in the middle of the tooth
without actually reaching the anterior end. In
some of these forms (as noted by Ride 1956) the
smooth anterior portion of the sectorial premolar
is longer than the smooth posterior portion, the
opposite of the condition found in
Hypsiprymnodon. In caenolestoids a comparable
range of morphology suggests position of
serrations is not significant in diagnosing groups
above the generic level.
Ml: Ontogenetic studies (Archer 1974) of
dasyurids suggest that Ml (called dP4 by Archer
1974) is the most anterior member of a posterior
molariform Zahnreihe which includes MI-M5.
Kirkpatrick (1969) has similarly suggested that
Ml and M2-5 (his dP4 and Ml -4) in
macropodids develop as related series of teeth,
although he does not interpret the relationship as
a Zahnreihe. Sequence of tooth development in
the macropodid Setonix noted by Berkovitz
(1966) is similar to that in dasyurids.
160
MEMOIRS OF THE QUEENSLAND MUSEUM
From a survey of M 1 morphology (to be
published), it is also apparent that these teeth, in
ail marsupials in which they are not reduced to
vestiges, although rarely premolariform,
are frequently molariform. Therefore morphology
and ontogeny is used here to conclude that the
marsupial deciduous cheektooth is actually the
first molar, i.e. Ml, there being no true
post-canine tooth replacement. In contrast is the
more traditional view of Owen (1840-5) and most
later workers who believe that this tooth is a true
milk-premolar that secondarily has become
molariform. Indirect evidence for Owen's view is
the well-known fact (e.g. Butler 1952) that in
many cutherian groups dP3 has undergone
molarization to either increase the number of
functional molariform teeth in juveniles, or to shift
anteriorly the molariform-premolariform boun-
dary. 1 do not think this is the case in marsupials
for four reasons. First, the oldest known
(Cretaceous) marsupials have an extremely
well-developed molariform Ml (e.g. Clemens
1966) which might not be the case if molarization
of a deciduous premolar was a secondary
development unless secondary molarization
occurred extremely early in primitive marsupials.
Secondly, except for didelphids, macropodids, and
some phalangerids, the marsupial Ml is almost
invariably too small to be functional and it seems
improbable that it would secondarily evolve
molariform characters when it never really has a
chance to function as a molar. In at least some
modern didelphids (Archer 1976) Ml is
comparable in complexity to the same tooth in
Cretaceous didelphids, thereby providing no
evidence for secondary molarization. In ma-
cropodids it is probable that M 1 has become
secondarily molarized, but there is no evidence
that this process of molarization in macropodids
started with a premolariform Ml. Third, there
appears to be a repetitive basic crown pattern in
Ml in distantly related groups including some
dasyurids, phalangerids, and diprotodontids which
suggests the possibility that an ‘archetypal’
molariform pattern may persist rather than
develop polyphyetically in teeth which are free
from heavy selective pressure. Fourth, recent
ontogenetic evidence indicates M 1 is part of the
molariform tooth series and not a predecessor to
P3.
The improbability of a tooth family relationship
between Ml and P3 is further indicated by the
various ways in which tooth reduction occurs in
this position. In many dasyurids, thylacinids,
peramelids, and diprotodonts such as phascolarc-
tids and Petaurids (Archer 1975), Ml is tiny or
absent while P3 is large. Yet in other diprotodonts
such as some phalangerids. Ml is only slightly
smaller than P3, and in macropodine ma-
cropodids, Ml is markedly larger than the small
P3. Ziegler (1971) has suggested that in mammals
in general reduction of premolar number occurs
first by loss of the permanent tooth and only later
by loss of the deciduous tooth. For this reason,
the inconsistent pattern of tooth reduction in
marsupials would suggest there is no true
milk-tooth in the postcanine cheektooth row.
Morphology of M| in potoroine macropodids
is similar to that of M 2 in many non-macropodid
diprotodonts such as Phalanger and Trichosums.
Broad aspects of this similarity have been noted
by Bensley (1903) and Ride (1961) both of whom
regard it as occurring in non-homologous teeth in
the two groups. Ride (1961) also regards the cusps
involved in the compressed trigonids of these
similar teeth to differ. The principal cusp on the
Irigonid of Mi in phalangerids (see below) is
regarded here (and by Ride 1961) to be the
protoconid. However, Ride (1961) regards the
principal cusp of M, in Hysiprynuiodon to be
the metaconid, a conclusion based on his
interpretation of a small cupsule on the posterior
slope of the main cusp as the protoconid, and of
the apparent topographic serial homology of this
cuspule with a cuspule on M^ in the position of
a protoconid. If Ride is right, the apparent
similarity between M^ of phalangerids and Mj
of macropodids is the result of convergence. It is
suggested below that the anterobuccal cusp on M 2
of petaurids and phascolarctids is, as Bensley
(1903) concludes, a ncomorph or protostylid, the
protoconid having shifted lingually. Phascolarctids
are also regarded by some authors (Winge 1941,
Archer 1976) as structurally ancestral to other
diprotodonts. Therefore it is possible that the
anterobuccal cusp in M 2 of Hypsiprymnodon (and
other macropodids) is the homologue of the
phascolarctid protostylid, and not the protoconid.
The liny cusp observed by Ride (1961) on M|
of Hypsiprymnodon may be the serial homologue
of this protostylid, the high cusp on that tooth
again being the protoconid. The compressed
condition of the trigonid of M^, and relatively
slight development of the anterobuccal cusp in
Hysiprymnodon might then be regarded as
structurally ancestral characters. Pressure to
molarize Mi and M; has resulted in enlargement
of this cusp in M 2 of all and M, of most other
macropodids.
Further support for the possibility that the
protoconid is anlerolingual on trigonids of M, in
macropodids is provided by Berkovitz(1967b). In
ARCHER: MARSUPIAL CHEEKTEETH HOMOLOGY
161
an ontogenetic study of crown development in
Seloni.x he shows that although the anterobuccal
cusp of Ml, interpreted by him to be the
protoconid, develops first, the anterolingual cusp
of M| develops first. Accepting Ride's (1961)
interpretation of this cusp as the metaconid,
Berkovitz concludes that ontogeny of M, in
Seionix is not therefore recapitulating phylogeny
because the protoconid is generally regarded as
the original trigonid cusp. Although the principal
of ontogeny recapitulating phylogeny in marsupial
tooth cusps has been questioned elsewhere
(Archer 1975), in the present case the early
development of the lingual cusp is more consistant
with its interpretation as a protoconid than a
metaconid.
Although these points arc made in order to
indicate that the cusps of molariform teeth with
compressed trigonids in different groups of
diprotodonts may be homologous, it does not
necessarily indicate that the teeth themselves are
homologous.
M2: Previous confusion in interpretation of M^
cusp homology has resulted from examination of
worn molars. Ride (1961) points out that Benslcy
(1903) must have had only worn specimens of
Hypsiprymnodon resulting in his failure to
correctly interpret the number of cusps on Mi.
Ride (1961) and Bensley (1903), however, seem
to have had only worn specimens of
Phascolarctos. Because I regard the morphology
of Phascolarctos to be basic to at least an
understanding of other diprotodont groups
(Archer 1976), it is of interest here to brieHy
describe the morphology of its Mi (e.g. J 13278).
The tooth have five principal cusps forming apices
of crests. The talonid has a buccal hypoconid and
a lingual entoconid. The cristid obliqua crosses
from the hypoconid to the tip of the tallest
trigonid cusp, the protoconid. This cusp is just
lingual to a medial position on the trigonid. The
protoconid is connected by a posterolingual crest
to the metaconid. A paracristid extends anteriorly
from the protoconid to the anterior end of the
tooth, the topographic position of a missing
paraconid. Buccal to the cristid obliqua,
protoconid, and paracristid, a well-developed
acce.ssory crest, with a neomorphic cusp at its
apex, extends from the anterior base of the
hypoconid to the anterior tip of the tooth. Bensley
(1903) suggests that this anterobuccal cusp is a
new development and not homologous with the
protocone of succeeding molars. I entirely agree,
and do not regard this interpretation as a violation
of the concept of serial homology, which leads
Ride (1961) to suggest that the only cusp he
observes on the anterolingual end of Mi is the
metaconid and the anterobuccal cusp is the
protoconid. Ride also regards the anterobuccal
cusp in Psendocheiriis, which is smaller, to be the
protoconc. Here again, details of unworn molars
as well as a comparison of the Mi of most species
of Pseudocheirus, make it reasonably clear that
the anterobuccal cusp in Pseudocherus is the
homologue of the protostylid in Phascolarctos and
not the protoconid. In Pseudocheirus. as in
Phascolarctos. the protoconid is the high
anterolingual cusp, with a posterolingual, variably
cuspid crest representing the metaconid.
The Ml of Hypsiprymnodon differs from that
tooth in phalangerids, phascolarctids and
petaurids, but is adequately illustrated and
described by Ride (1961). In other macropodids,
morphology of M 2 is similar to Hypsiprymnodon
but lacks the slight lateral compression of the
trigonid.
It is clear that in all macropodids the
anterobuccal cusp on M 3 .S is the protoconid, as
concluded by Ride (1961). The alternative view,
proposed by Bensley (1903), that this cusp is the
homologue of the phascolarclid protostylid of Mi,
is not acceptable. However, Ride's conclusion that
there is no evidence for suggesting Mi in
macropodids was ever other than quadrituber-
cular, is doubtful because the anterobuccal cusp
on Ml may not be the protoconid.
Ml of all diprotodonts except most macropodids
and all diprotodontids has a laterally compressed
trigonid. As a result of this compression, the
paracristid assumes a longitudinal orientation by
lingual displacement of the protoconid. In almost
all diprotodonts, the degree of compression
correlates w ith the degree of sectorial development
of P 3 . A culmination of this trend may be seen
in ihylacoleonids where the trigonid of M 2 is a
massive, longitudinal shearing crest. The opposite
extreme is found in some diprotodontids and
macropodids where Pj is frequently almost round
and tubercular and the trigonid of Mi lacks any
compression. The fact that in Phascolarctos
attempts to molarizc the Mi trigonid result in
development of a new cusp, rather than a buccal
shift of the protoconid, indicate the stability of the
laterally compressed trigonid in diprotodonts. In
polydolopid (and possible abderitine) caenoles-
toids, the compressed trigonid of M 2 is not clearly
formed in the same way as it is in diprotodonts.
The analogue of the short longitudinal paracristid
may be a new development unrelated to the actual
positions of the protoconid and paraconid.
162
MEMOIRS OF THE QUEENSLAND MUSEUM
Concepts Of Cheektooth Homology
In Polypkotodont Marsupials
As indicated above from ontogenetic and
morphologic data, dasyurids and didelphids have
eight postcanine cheekteeth which are most
appropriately interpreted as Pl-3, Ml and M2-5
(where Ml is the nomeclatural equivalent of the
dP3 of Stirton 1955). There is nothing about
perameloids to suspect they differ from this basic
polyprotodont pattern. Wilson and Hill (1897)
have shown that P3 appears to develop from
dental lamina lingual to the developing Ml but
their data do not show that it could not be
comparable with the dasyurid situation described
by Archer (1974) where P3 appears to develop
from the dental lamina between P2 and Ml, and
only secondarily comes to lie lingual to Ml as
the tooth buds grow and crowd the developing
tooth row as a whole.
Stirton (1955) employs the tooth nomenclature
PI-3, dP3. Ml 4. This nomenclature is also used
by almost all modern Americans (e.g. Woodburne,
Tedford, Clemens, Marcus, Lillegraven, Camp*
bell) as well as by some Australian zoologists (e.g.
Bartholomai. Marshall, Plane) who do not follow
Thomas (1888). Although based on the concept
of tooth replacement, it purports to avoid implying
that a particular premolar tooth family has been
lost in marsupials. As Mahoney and Ride (1975)
point out, no system of numbering can avoid
implying homology, and it could be concluded
from Stirlon's terminology that marsupials have
lost the original P4 of their common ancestor with
placental mammals, even though Stirton did not
intend to imply this concept.
The common alternative is the system of
Thomas (1888) which is PI, P3-4, dP4, Ml-4. It
is used by many zoologists in Australia and
England (e.g. Ride, Archer in previous works,
Mahoney, Berkovitz, Merrilecs, etc.) and some
American zoologists (e.g. Tale). Because of its
wide use among Australian zoologists, I adopted
it in earlier works. However, it is based on two
apparent misconceptions: that the homologue of
the placental P2 is missing from the marsupial
tooth row (the lack of acceptable evidence for this
is reviewed by Archer 1974, 1975); and that Ml
(dP3 of Stirton) is a true milk-tooth.
Ziegler (1971) also accepts the apparently
erroneous concept of cheektooth replacement in
marsupials, but differs from Thomas (1888) in
regarding marsupials to have lost PI, accordingly
designating the functional adult premolars P2, P3
and P4. l.undelius and Turnbull (e.g. 1973) also
use this system but regard homology of P2 to be
doubtful.
Reasons for not accepting any current concept
of loss of a particular premolar family in
structurally primitive marsupials arc given
elsewhere (Archer 1975). Although it docs seem
probable that ancestral marsupials lost a premolar
family which they must have shared in their
common ancestor with placentals, there is as yet
no conclusive morphological, ontogenetic, or
palaeontological evidence for this loss.
Concepts Of Cheektooth Homology
IN Diprotodont Marsupials
Reports of the significance of residual lingual
and spurious buccal traces of dental lamina in
diprotodonts arc not considered here. They are
adequately reviewed by Berkovitz (1966) who
concludes that most lingual downgrowths are
merely residual free ends of dental lamina and do
not represent vestigial replacement teeth.
In all diprotodont marsupials there are fewer
teeth in the total dentition than in any
polyprotodont. Most of the reductions in number
involve incisors, canines and prcmolariform teeth.
Petaurids have the highest diprotodont tooth
number and, at least in their upper postcanine
cheekteeth, the number is identical with
polyprotodont marsupials. Further, they show the
same manner of apparent tooth replacement as
polyprotodonts. They differ from most polypro-
todonts in that the M‘ is very tiny (Archer 1975).
In some other diprotodont families, such as the
diprotodontids and phalangerids, M' is much
larger and functional. Therefore in at least these
diprotodont groups, morphological as well as the
limited ontogenetic data support the cheektooth
homology of PI-3, Ml, M2-5.
Macropodid diprotodonts are unique among
marsupials in that P3 during eruption replaces the
teeth regarded here as P2 as well as Ml. This
raises the possibility that the macropodid Ml is
not homologous with M 1 of other marsupials, a
possibility however which is negated by the
ontogenetic evidence that in macropodids, as in all
marsupials. Ml is the first molar. Similarly,
comparisons of the phalangerid M-, with the
macropodid M, reveal at least a basically similar
trigonid construction and could be regarded as
evidence that the teeth are homologous. However,
it these two teeth were regarded as homologues,
it would be necessary to ignore the ontogenetic
data which indicates that the macropodid M| is
a first molar and the phalangerid M^ is a second
molar. The posterior deciduous cheekteeth in the
two groups, i.e. Ml, also show similarities in
trigonid construction and if these teeth are
regarded as homologues, there is no conflict with
ontogenetic data.
ARCHER; MARSUPIAL CHEEKTEETH HOMOLOGY
163 .
In summary, there is no evidence for suggesting
that the homology of the cheekteeth in
diprolodont marsupials differs from that of
polyprotodont marsupials. Even kangaroos, with
their unique number of deciduous cheekteeth, lack
any true postcanine milkteeth and are thus
essentially similar to other marsupials.
Caenolestoids
There is no evidence for apparent tooth
replacement in living caenolestids in over 150
specimens of Caenoloestes, Orolestes and
Rhyncholestes examined by the author in museum
collections. Similarly, there does not appear to be
any evidence for tooth replacement in the great
variety of known fossil caenolestoids, including
those forms with large plagiaulacoid premolars. If
tooth replacement of the sort which occurs in
other marsupial orders does occur in caenoles-
toids, it must occur very early in ontogenetic
development. Assuming this is the case, the
maximum caenolestoid postcanine cheektooth
dentition would be PI -3, Ml (not yet observed),
M2-5. If tooth replacement does not occur, then
it is possible that caenolestoids represent a unique
order of marsupials all members of which have
no more than seven postcanine cheekteeth.
Discussion
All morphologic and ontogenetic evidence in
polyprotodont and diprotodont marsupials leads to
the conclusion that there are three prcmolar tooth
families and five molar tooth families the first of
which is deciduous.
This conclusion has brought me to an impasse.
If these data are acknowledged but not used, there
is a possibility that in using an alternative and less
probable system, comparisons of particular teeth
within the various marsupial groups and between
marsupials and placenlals may be meaningless.
For this reason, although I have previously used
the system of Thomas (1888), I intend to use the
nomenclature PI -3, Ml, and M2-5 until it is
shown to be wrong or less probable than an
alternative system.
ACKNOWLEDGEMENTS
Dr A. Bartholomai and Mr B. Campbell
(Queensland Museum) read and constructively
criticised a draft of this work. Ms R. Owen and
Mrs C. Farlow (Queensland Museum) typed
drafts of the manuscript.
Early concepts from which this work has
developed were discussed with Dr W. D. L. Ride
(then Director of the Western Australian
Museum) and the present work owes much to his
stimulation.
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1961. The chcck-tcclh of Hypsiprymnodon moschatus
Ramsay 1876 (Macropodidae: Marsupialia). J.
Roy. Soc. West. Aust. 44: 53-60.
1970. ‘A guide to the native mammals of Australia’,
xiv and 249 pp. (Oxford Univ. Pr.: Melbourne).
Stirton. R. a.. 1955. Late Tertiary marsupials from
South Australia. Rec. S. Aust. Mus. II:
247-68.
Tati-.. G. H. H., 1 947. On the anatomy and classification
of the Dasyuridae (Marsupialia). Bull. Amer. Mus.
Nat. Hist. 88: 97-156.
Tmoma.s, O.. 1888. ‘Catalogue of the Marsupialia and
Monotremata in the collection of the British
Museum (Natural History)', xiii and 401 pp.
(British Museum (Natural History) : London).
TiRMitii. W. D., LtNDiiK.s. E. L., 1973. The
mammalian fauna of Madura Cave, Western
Australia Part 1. Fieldiaana {Geology} 31: 1-35.
Wi\GF. H., 1941. ‘The inlcrrrelationships of the
mammalian genera', Vol. 1, xii and 418 pp. (C. A.
Reit/cls Forlag : Kobenhavn).
Woodward. M. F., 1893. Contributions to the study of
mammalian dentition. Part 1. On the development
of the teeth of the Macropodidae. Proc. Zooi Soc.
Land. 1893: 450 73.
Zit'Xii.i-R. A. C., 1971. A theory of the evolution of
thcrian dental formulas and replacement patterns.
Q. Rev. Biol. 46: 226-49.
Mem. Qd Mus. 18 ( 2 ): 165-77, pis. 28-9. [1978]
RECENT LOCAL FAUNAS FROM EXCAVATIONS AT
HERVEYS RANGE, KENNEDY, JOURAMA, AND
MOUNT ROUNDBACK, NORTH-EASTERN QUEENSLAND
Michael Archer
Queensland Museum
and
HELEN BRAYSHAW
James Cook University
ABSTRACT
The non-human remains from four excavations carried out by H. Brayshaw in north
Queensland are identified. Bone damage noted is interepreted to be caused by smashing,
charring, cooking, chewing, rodent gnawing, insect and soil damage, manufacture into tools,
and possibly incision.
Human remains suggest cannabilism. Humans included in the deposits are generally juvenile,
but do not appear to represent a normal mortality curve. As a result, it is suggested that
there was killing with a selective bias. There also appears to be a selective bias in the particular
human skeletal elements present.
One animal from the Herveys Range deposit represents a diprotodont marsupial evidently
unknown to European zoologists. Other vertebrate species represent those found today in
sclerophyl forest, savannah woodlands with rocky areas, and streams. E.xcept for large bats,
volant and gliding animals, such as marsupial gliders and birds, are unexpectedly absent.
Kangaroos are the most abundant animals represented in the deposits.
As part of a regional archaeological survey four
small excavations were undertaken along the coast
of the Herbert/Burdekin district, north Queens-
land, at Herveys Range, Kennedy, Jourama and
Mount Roundback (see Fig. 1). A report of these
excavations is given in Brayshaw (1977). Lithic
material, shells, and bones were found at all four
sites; human bone occurred in the deposits at
Herveys Range, Kennedy and Jourama, but not
at Mount Roundback. Bones recovered from the
excavations were placed in bags and taken to
Townsville for initial sorting where ribs, isolated
mammal vertebra, central portions of limb shafts,
and phalanges were removed. The remaining
potentially identifiable specimens were sent to
Brisbane where they were allowed to soak for a
few minutes in water, and then gently cleaned
with a damp wad of cotton. This precaution was
taken to avoid causing accidental marks on soft
bones. The best preserved, most complete and
taxonomically diverse specimens were from the
Herveys Range deposit and for this reason, were
identified first. Bones from the other sites were
then compared with the Herveys Range material
and, unless they clearly differed, were referred to
the same species. The abundant human remains
were isolated and analysed separately. A copy of
W. Wood's list of identified human materia! is
lodged in the Queensland Museum library. A
summary of this report is presented here as an
Appendix.
Vernacular and scientific names of mammals
follow the usage of Ride (1970). Dental
nomenclature is that used by Archer (1978).
Fossil kangaroo names are those used by
Bartholomai (1975).
Representative specimens are registered in the
Queensland Museum fossil vertebrate collection.
The remainder of the material will be lodged in
the collections of James Cook University.
Stratigraphic level designations cited in the text
are given as follows: H indicates Herveys Range;
C indicates Jourama; E indicates Mount
Roundback; F indicates Kennedy; excavation
squares in each site are given as roman numerals,
e.g. I to XII; depth is indicated by numbers 1 to
166
MEMOIRS OF THE QUEENSLAND MUSEUM
OUNK ISLAND
▼ EXCAVATED SITE
LAND .
ABOVE.
.610 METRES
KENNEDY
HINCHINBROOK
ISLAND
PALM ISLANDS
JOURAMA
MAGNETIC ISLAND
TOWNSVILLE'
HERVEYS
RANGEcC)
mount’ .jBOwen
ROUNDBACK k
Vi
CHARTERS
TOWERS
ISLAND
.COLLINSVILLE
MACKAY
LOCATION
QUEENSLAND
Fig 1. Locations of the Kennedy, Herveys Range, Mount Roundback and Jourama excavations in north-eastern
Queensland.
12 where each number represents a 5 centimetre
spit or interval. Thus H V 7 indicates the 30-^5
cm spit of square V, at Herveys Range.
NON-HUMAN REMAINS
Herveys Range
Species identified from the Herveys Range
excavation arc shown in Table 1. In this table, no
attempt has been made to itemize the fauna by
squares. The numbers represent minimum number
of individuals based on the largest number of any
common osteological or dental unit. Question
marks indicate laxa doubtfully identified.
Comments about Taxa
Fish: The fish has not been identified. It is
invariably represented by broken dentary
fragments. Although they arc not burned, one
fragment may have been cooked. The entire fish
would be relatively small, probably smaller than
30 cm in length. Other diagnostic and more
abundant fish remains such as vertebrae are easily
recognised and their absence suggests fish were
not a common food item.
Lizards: There are at least two kinds of
lizards represented: The Frilled Lizard
(Chlamydosaurus kingii} and the Blue-Tongue
Skink (Tiligua scincoides). Both are represented
by broken dentary or maxillary fragments, and in
some cases postcranial elements such as pelvis and
femur fragments. None show clear evidence of
burning, cooking, or tooth marks.
The Eastern Blue-tongue Skink is similar to the
Shmgle-back IJzard (Trachydosaurus rugosus)
and has been differentiated here by the fact that
its teeth rapidly enlarge in overall size from front
to back, the rate of enlargement being much
greater than that in the Shingle-back. One
Blue-tongue specimen (in H IV 10) is much
smaller than the others (in H VI 2) but probably
represents a juvenile rather than another taxon.
These occurrences of Blue-tongue and Frilled
lizards are within the known ranges for the
species. Both are recorded from open forest
situations.
Snakes: Vertebral and cranial fragments
represent at least one kind of large Python.
Regrelably, comparison has not been possible with
either the Black-headed Python {Aspidites
melanocephalus) or the Water Python (Liasis
fuscus). In comparison with other large pythons
{Morelia spilotes, M. areas, and Liasis
amethysfinusl the Herveys Range material
including a large isolated right quadrate (in H VII
2) most resembles the Carpet Snake (Morelia
spilotes). One of the five identifiable fragments
has been burned (a vertebra), another cooked (the
right quadrate), and another (vertebra in H VI
I) has been broken and probably chewed. Carpet
snakes are known from open forest situations and
the Herveys Range occurrence is within the known
range for the species.
TABLE 1: Fauna from the Excavation at Herveys Range, 1974
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND 167
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Fauna
Fish
Chiamydosaurus kingii
Tiliqua scincoides
Morelia sp.
Isoodon macrourus
Trichosurus cf. T. vulpecula
Petrogale cf. P. penicillata
Wallabia bicolor
Macropus agilis
M. parryi
M. giganteus
Small rodent
Meserubriomys gouldii
Macroderma gigas
Pteropus scapulatus
Canis familiar is
Homo sapiens
Unknown animal
No. of laxa
Squares excavated
168
MEMOIRS OF THE QUEENSLAND MUSEUM
Bandicoots; The species represented is the
Brindled Bandicoot {Isaodott macrourus}. It is
markedly larger than Cape York specimens of the
otherwise similar Brown Bandicoot {Isoodon
obesulus} and comparable in size to south-eastern
Queensland Brindled Bandicoots. The only clearly
referable specimen (in LI VII I) is an undamaged
dentary. There is nothing to indicate its presence
in the deposit is necessarily the result of human
activity. The Brindled Bandicoot is known from
open forest situations and the present occurrence
is within the known range of the species.
Brush-Tailed Possums: We have not
found a consistani morphological character which
separates molar teeth of Brush-tailed Possums
{Trichosurus vulpecula) from the Bobuck {T.
caninus). Because Brush-tailed Possums frequent
all habitats except rainforest which is the
preferred habitat of the Bobuck, identification of
the species could be an important aspect in the
interpretation of palaeoecology. However, the
specimens from Herveys Range are not inconsis-
lant with interpretation as Brush-tailed Possums
and in view of the presence of open forest in the
area today, and the absence of any other rainforest
animal remains in the deposit, the simplest
conclusion is that the specimens probably do
represent Brush-tailed Possums rather than
Bobucks.
Brush-tailed Possum remains were present on
most stratagraphic levels. Every specimen is
broken, most show evidence of having been
cooked, but none are burned. Some have fractures
or marks which suggest chewing (c.g. in H VII
surface). Most specimens represent young adults.
There were no juveniles or pouch-young. Herveys
Range is within the known distribution of
Brush-tailed Possums.
Rock-WALLABIES; There are differences of
opinion about the number of species of
Rock-wallabies (Pelrogale) in north Queensland.
On the basis of Ride’s (1970) remarks about
distribution, the species in the area of Herveys
Range today is likely to be the Brush-tailed
Rock-wallaby (P. penicillataj. Other authorities
suggest additional forms (some representing taxa
included by Ride in P. pencillata) occur in or near
this area including godmani, puella, and inornafa.
We have attempted to differentiate the molar
teeth of these forms using modern Queensland
Museum specimens and could find no consistant
way of separating small samples referable to
Godman's Rock-wallaby {P. godmani} from large
samples referable to the Brush-tailed Rock-
wallaby (P. penicillata). Because all forms of
Petrogale occupy similar rocky habitats, it
probably docs not make any significant difference
which species is represented in terms of
palcoecological interpretation. Nevertheless, the
only Rock-wallaby specimens in the Queensland
Museum from the Townsville area represent the
Brush-tailed Rock-wallaby, and they are very
similar to the Herveys Range material. For these
reasons the Herveys Range specimens arc
regarded as Petrogale cf. P. penicillata.
Rock-wallabics arc the most common animal in
the deposit. Every one of the 45 specimens is
smashed, cooked or burned. One dentary (in H
VII Surface) has tooth marks on it that suggest
a non-human carnivore. Most dentaries have I|
broken off at the base such that the enamel face
projects as a hard transverse edge beyond the
dentine. This suggests the dentaries were intended
to be. or were actually used as, tooth engravers
(Mr K. Akerman, pers. comm.). One left dentary
(in H III 5) is possibly abnormal. Although
basically similar to other Rock-wallaby specimens,
the last molar has a peculiar post-link and a
posterior cingulum, characters otherwise unknown
in Rock-wallabies. It may represent a super-
numerary fifth molar (Archer 1975). Another
specimen (in H VII 1), a right dentary. has what
appears to be an impacted projecting down into
the alveolus for the missing Rl|.
Swamp Wallabies; Swamp Wallabies
(Wallahia hicolor) are uncommon in the deposit
being represented by only five specimens. Of
these, three have premolars and are positively
identified. Two others arc referred with some
confidence, but not certainty, on the basis of molar
morphology. The fifth specimen is an isolated 1|
and, although only doubtfully referred to this
species, cannot be matched with any other
wallabies identified from this deposit. All
specimens are smashed but none are clearly
cooked or burned. One of the dentaries has
I] broken off in the manner described above for
Rock-wallabies and may have been intended for
use as a tooth engraver. The Herveys Range
occurrence for Swamp Wallabies is within their
known range and habitat requirements.
Agile Wallabies: Although called a wallaby
(Ride 1970), Agile Wallabies {Macropus agilis]
are very large kangaroos and the most common
large animal in the Herveys Range deposit, being
represented by at least 17 individuals and 33 skull
or dental fragments. All specimens are smashed,
cooked or burned. All dentaries have I, missing
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND 169
or broken off in the fashion described above for
Rock-wallabies. Almost half of the specimens
whose age has been determined represent juveniles
(P3 hadn't erupted). This occurrence of Agile
Wallabies is within their known range and habitat
requirements.
WhiptailS: Whiptails (Macropus parryi)
sometimes also called Pretty-face Wallabies, are
larger than Rock-wallabics but smaller than Agile
Wallabies. They are as rare in the Herveys Range
deposit as the comparable sized Swamp Wal-
labies. All specimens arc broken, but none arc
obviously cooked, burned or chewed. Whiptails
have a molar morphology similar to Agile
Wallabies but arc immediately distinguished by
their much smaller and bicuspid P,. One Herveys
Range specimen (in H IV Surface), the proximal
end of an ulna, has been identified as a Whiplail
because it exactly matches ulnae of Whiptails and
is clearly dissimilar to ulnae of all other
marcopodid species known from the Herveys
Range deposit. One of three dentary specimens (in
H VII 1) is only tentatively identified as a
Whiptail. It differs from other Herveys Range
specimens and Queensland Museum material in
possessing a very narrow anterior cingulum and,
judging by the roots, relatively large P^ Unless
it represents a taxon otherwise unrecognized from
this deposit, it probably represents an extreme
variation in Whiptails. The Herveys Range
occurrence of Whiptails is within their known
distribution and accords with their habitat
requirements.
Great Grey Forester: This is the largest
species { Macropus giganteus) represented. Un-
fortunately, all of the very large kangaroo remains
are too incomplete to identify with any certainty
and it is possible that Antilopincs (M. antilopinus)
or Euros (M. rohustusj are represented among the
unidentified material. The only upper molar has
a small but well-developed forelink and well-
developed buccal accessory cuspules in the
mid-valley. The first character favours identifica-
tion as the Great Grey, because the vast majority
(there are exceptions) of Antilopines and Euros
lack a well-developed forelink. The second
character is ambiguous, being present in some
fossil species (e.g. Macropus (Osphranter) pan
and M. (OJ woodsij of the group of kangaroos
including Antilopines and Euros, but evidently not
in these two modern species of the group. For the
present, the remains are referred to the Great
Grey Forester, but with reservations. All of the
specimens are broken but only one (in H VII
Surface) is obviously also burned. Great Grey
Foresters are likely to occur today in the Herveys
Range area, as it is within their known range and
habitat requirements. Because Rock-wallabies
were (or are) so abundant, it is probably
reasonable to assume the habitat would also suit
Euros.
Unknown Animai.: The single most intriguing
fragment from the Herveys Range deposit is a left
dentary fragment in H III (5). It appears to be
mammal because the tooth alveoli are deep and
completely unlike those of fish or reptiles
examined. The absence of small incisor alveoli
anterior to the large caniniform alveolus suggests
it is not a polyprotodont marsupial. It also
resembles no known placental mammal. In all
probability it represents a diprolodont marsupial
somewhat similar to the Green Ringtail Possum
fPseu{/oc/ic/rus archeri) which has a very short
diastema and a relatively steeply inclined I,. The
Herveys Range specimen differs from these in
completely lacking a diastema and in being larger.
There is little point in speculating further about
its affinities because it is such a small fragment.
However, it certainly suggests the presence of an
as yet undiscovered (by European zoologists)
north Queensland mammal. Whatever it repre-
sents, it was evidently used by the Aborigines as
a food because, like all the other bones in this
deposit, it has been thoroughly smashed.
Small Rodents: One broken specimen (in H
VI 2) retaining only M 3 , appears to be a small
pseudomyine rodent. It clearly does not represent
a species of Raitus. There is no evidence that it
was cooked.
Black-footed Tree-rat: This very large
rodent ( Mesembrioniys gouldii) may be represent-
ed by an isolated LI]. The distribution of the tooth
enamel appears to differentiate it from the north
Queensland Giant White-tailed Rat (Uromys
caudintaculatus) and the common Water Rat
(Hydromys chrysogasterj. Although it is similar
to the Gaint White-tailed Rat, the tooth enamel
in Tree-rat specimens in the Queensland Museum
extends farther around the buccal side of the
tooth. This represents a significant range
extension southwards for the Black-footed
Tree-rat. although it is just within the range of
the Gaint White-tailed Rat. Because the
identification is based on such limited material,
we are hesitant to regard it as anything other than
a tentative identification, and one which requires
substantiation by much better material. The
170
MEMOIRS OFTHEQUEENSLAND MUSEUM
Black-fooled Trcc-rat normally inhabits woodland
savannah while the Giant White-tailed Rat
inhabits rainforest. This is added support for the
possibility that the tentative identification is
correct.
Ghost Bat: Only one specimen (in H V 3)
represents this large carnivorous bat
(Macroderma gigas). There is nothing to indicate
that it was cooked. Its presence in the deposit
suggests but does not necessarily mean that it was
eaten by Aborigines. Ghosts Bats are nocturnal,
extremely alert, and do not hesitate to fly during
the day if disturbed. This would probably make
them very difficult animals to catch. Ride (1970)
considers that their diurnal roosts are invariably
in caves or tunnels. The Hervey's Range
occurrence is within their known modern
range.
Red Flying Fox: This relatively small Flying
Fox (Pteropus scapulatus) is represented by one
broken specimen (in H VII 4) which suggests the
species was eaten, albeit infrequently. Like the
Ghost Bat it is nocturnal but roosts in trees and
probably would be an easier species to catch. One
specimen (in H VI surface), a left denlary, lacks
the Ml normally found in this species. Only one
modern specimen in the Queensland Museum
shows the same condition. This distribution record
is within the known range and choice of habitat
for the species.
Dingo: Dingoes (Canis familiaris) are repre-
sented by only three teeth. Two of these appear
to represent broken adult canines, one of which
(in H VI Surface) may have been burned. The
third tooth, undamaged, appears to have been an
incompletely erupted LdP 4 , thus presumably
broken out of a dentary rather than naturally
dropped out or shed. The two adult canines are
split dorso-vcntrally, a type of fracture that
usually occurs after a tooth is heated, or allowed
to age over a long period in a skull. If the Dingoes
had broken the teeth during life, we would expect
the break to have been transverse along the
horizontal plane. Dingoes are known from almost
all areas of Australia.
Aborigines: Aboriginal remains are spread
throughout the deposit and the great majority are
thoroughly broken. Some are also burned. Of
particular interest is the symphysis of a very small
child which is thoroughly burned as well as
broken. Details of the human remains are given
below.
Bone Damage
Damaged bones noted may be grouped into
seven categories: broken bones; bones clearly used
as implements; burned bones; cooked bones;
decalcified bones; soil or insect damaged bones;
tooth-marked bones.
Broken Bones: Of over 150 identifiable bone
fragments, less than a few dozen of the smallest
such as larsals are intact. In addition to
identifiable fragments, there are hundreds of
smaller fragments too smashed to be
identified.
Very few of these show gouges suggesting points
of violent impact. From this we conclude either
that the bones were crushed individually with a
blunt instrument, such as a flat rock, or were
crushed while still part of the whole carcase, the
skin and flesh thereby protecting the bones
surfaces from impact damage.
Some fragments appear to have had tooth
crowns deliberately smashed off the roots (e.g. the
cheekteeth of a kangaroo maxillary fragment in
H VII 10). Almost all kangaroo dentaries have
It broken off in a manner suggesting the dentaries
were intended for use as tooth engravers (see
above). Alternatively or concordantly the incisor
tips may have been used elsewhere, perhaps as
ornaments or points for spears. The very low'
number of isolated lower incisor crowns compared
with the high number of dentaries lacking this
tooth, is evidence for this possibility.
Obvious Bone Implements: Two bone frag-
ments (one in H VI 1, and one in H VII 4) were
obviously used as awls, needles, or spatulas. Both
may have been manufactured from the dense part
of a kangaroo fibula.
Burned Bones; Burning is evidenced by the
black, charred texture of a bone surface.
Fragments that do show burning are normally
thoroughly burned. It is possible that some bone
fragments encountered while eating may have
been tossed back into the fire. Although it seems
highly unlikely that carcases would be cooked
until charred as a preparation for eating, some
bones forming extremities such as teeth or
phalanges could have been burned well prior to
the flesh being cooked.
Cooked Bones: Interpretation of alteration in
a bone due to cooking is less easy than recognition
of alteration due to burning. Bones have been
interpreted here as cooked if they show a
particular brown colouration such as might result
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND 171
if they were contacted by hot oils or other
by-products of heated flesh. One humerus
fragment (in H VII 5) has two distinctly charred
ends. The central portion however, is brown and
merges gradually into the black of the charred
ends. This is the brown colour regarded in other
bones to indicate alteration due to cooking.
Other processes could produce a brown colour
in the bone, such as acids in the soil in which the
bones are burned. No attempt was made here to
distinguish these possible causes.
Decalcified Bones: A few bone fragments
(such as one in H VI Surface) have a very soft
whitish texture and can easily be scratched with
a fingernail. Possibly these bones were charred
and subsequently bleached. Alternatively, they
may have passed through the alimentary tract,
either of an Aboriginal or a Dingo. Bone
fragments of this type are very uncommon in the
deposit.
Possibly Insect-damaged Bones: One
dentary fragment (in H VII 1) of an Agile
Wallaby has pits on its surface which suggest
either corrosion or the activities of invertebrates,
such as beetle larvae. Burned, softened bone is
often subject to insect destruction. Damage of this
sort has been observed on bones from other similar
deposits.
Tooth-marked Bones: Tooth marks are
relatively uncommon on bones from the Herveys
Range deposit compared with those from the
Jourama deposit. Three sorts have been recog-
nized: fine, sharp, conical depressions or scratches
possibly produced by Dingoes or children (e.g. in
H VII surface); blunter, deeper conical depres-
sions possibly produced by adult Aboriginals with
blunter canines (e.g. in H VIII 5); marks clearly
attributable to rodents (e.g. in H VII 4). In the
case of rodent gnawing, the implication is that at
least some bones were left unburned on the living
surface. The particular bone gnawed by rodents
had also been smashed and apparently chewed by
another carnivore such as a Dingo.
The absence of more extensive chewing suggests
the possibility that food was relatively abundant
in the Herveys Range area than in the Jourama
area.
Horizontal Distribution of Bones
IN THE Deposit
It was evident that for any given level (i.e.
surface, level 1, and so on) bones were more
abundant towards the rear of the shelter. On the
surface level, H HI (the farthest from the shelter
back, and directly under the drip line) had no
identifiable fragments; H IV had 2; H V had 6;
H VI had 4; and H VII (rear of shelter) had 17.
On level 1, H III had 4; H IV had 1; H V had
5; H V! had 8; and H VII had 26. This suggests
cither that Aboriginals centred their activities at
the rear of the shelter to avoid high temperatures
or rain or they may have centred their activities
forward of the rear and tossed unwanted bone
fragments to the rear of the shelter, possibly to
keep them out from under foot.
Kennedy
Unlike the material from the Herveys Range
deposit, very few of the bones from the Kennedy
deposit are identifiable. Accordingly the non-
human fauna is discussed as a whole.
The only two recognizable taxa arc as follows:
fish (unidentified), levels I, 2 and 4; Frilled
Lizard {Chlamydosaurus kingii), level 5, 7 and
possibly 1. All bones are smashed or merely
broken and several arc clearly burned. Other types
of bone damage include corrosion pits (in F II 2),
and decalcification (in F II 7). Comments relative
to this kind of damage are noted above in
reference to the Herveys Range deposit. The
presence of only the Frilled Lizard, rather
than the Eastern Blue-tongue lizard which is the
more abundant of the two at Herveys Range, is
puzzling but the samples are too small for this
difference to have statistical significance. The
abundance of lizard rather than mammal species
is perhaps more meaningful, and is an even more
radical departure from the Herveys Range
deposit, but we do not know what it means. The
relatively greater abundance of fish indicates
either a proportionately greater amount of time
spent fishing, or else a greater abundance of fish.
But again, it is impossible to interpret the true
significance of these site differences because of the
small size of the Kennedy sample.
Mt Roundback
As in the case of the Kennedy site, the
identifiable non-human bone fraction of the Mt
Roundback deposit is small.
There 'are six identifiable non-human taxa:
molluscs (pelecypod, type unidentified), 5-10 cm,
represented by a small shell fragment; amphibians
(at least one large species, possibly a large hylid),
surface, represented by isolated limb bones; snake
(possibly an elapid?), 5-10 cm, represented by a
small vertebra; Brush-tailed Possum {Trichosurus
vulpecula, but see applicable comments about
taxonomy given above for the Herveys Range
172
MEMOIRS OFTHE QUEENSLAND MUSEUM
specimens), surface and 5-10 cm. represented by
an isolated lower incisor and a dentary fragment;
Rock-wallabies {Petrogale sp.), 10-15 cm and
20 25 cm, represented at least by maxillary and
dentary fragments; and a large Kangaroo,
probably a Euro (Macropus cf M. robustus), 5-10
cm, represented by at least an isolated upper
molar.
The only departure from species also present in
the Herveys Range deposit is the possible Euro.
Identification of the single large tooth is based on
the fact that although there is a forclink. it is very
tiny, a condition common to Euros but not Great
Grey Foresters which normally have a well-
developed forelink. Many other bone fragments in
the Mt Roundback deposit represent very large
kangaroos, but because they are mostly postcran-
ial fragments, they are unidentifiable.
Bone damage
All bones are broken, with rare exception.
Some, but not all, of the frog bones on the surface
arc undamaged. None of the bone fragments on
the surface, but some within the deposit are
burned. One pelvic fragment (in E6, 15-20 cm)
appears to have been decalcified and/or burned.
Another pelvic fragment (in E7, surface) shows
corrosion pitting of the sort noted in the Herveys
Range deposit. One bone fragment (in E4,
surface) shows damage caused by teeth, but there
arc no clear indications of what carnivore was
responsible. Kangaroos will even chew bones
though they don't leave conical tooth depressions
near the broken ends.
JOURAMA
The non-human remains from the Jourama
deposit represent few taxa, and none otherwise
unrepresented in the Herveys Range deposit.
However some aspects of bone damage and the
abundance of Aboriginal remains in this site make
it unique.
Taxa
There are four identifiable non-human taxa:
mollusc (pelecypod, cyrenid, Batissa or Cyrena,
which occurs in the lower reaches of tropical
rivers), level 3; Rock-wallaby {Petrogale sp.),
possibly represented by specimen on level 2; Agile
Wallaby ( Macropus agi/is), level 4 and possibly
level 2; and the Dingo (Canis faniiliaris), level 1
and a specimen not allocated to a level.
In terms of composition, this assemblage more
cosely resembles that from Herveys Range than
from Kennedy.
Bone damage
The material from this deposit is unique in that
almost half of the level samples contain bone
fragments showing clear evidence of tooth marks.
In many of these chewed specimens (e.g. in C III
4, C 111 5, and C III 6). the entire surface of the
bone fragment is disfigured by gouges, fractures
and tooth impressions, even to the extent of
disfiguring the edges of the fragmented bone.
Nothing, unfortunately, indicates whether the
carnivores were Aborigines. Dingoes or both. We
suspect Aborigines because nothing like this kind
of damage is present in a large sample of bones
deliberately fed to captive Dingoes (an experiment
carried out for us by Mr A. Boorsboom.
Queensland University). Of course, much depends
on how chewing behaviour changes as a function
of hunger. In the case of the experimentally fed
Dingoes, the animals, although hungry, were
certainly not starving. One dense limb bone
fragment (in C III 4) shows a unique kind of
damage. It has what appears to be a deep incision
that is clearly not attributable to rodents. Dingoes
or Aboriginal teeth. It superficially resembles a
wedge-shaped incision present in a fossil bone
from Mammoth Cave, W'estern Australia
(Archer. Merrilees and Crawford, in preparation)
which has been attributed to Aboriginal activity.
In the present case, no attempt has yet been made
to find evidence of sawing or chipping. An
alternative cause of the damage could conceivably
be spalling, although a percussion point is not
obvious. Other types of bone damage in this
deposit arc of the same kind as noted for bones
in the Kennedy and Mt Roundback deposits.
HUMAN REMAINS
Herveys Range
On the basis of the sample identifications
provided by Dr W. Woods, there are 13 juveniles
and 17 adults represented. There appears to be
a heavy mortality between the ages of 10 to 12
years. There is no clear evidence for very old
individuals. Adults that have been aged are
generally middle-aged. Only one infant was noted.
Many human bones, including juveniles, show
charring and scratch marks. Virtually no human
bones except some hand and foot elements are
unbroken. These facts suggest that the human
remains represent a food eaten by the inhabitants
of the Herveys Range shelter. The nature of the
bone breaks and charring is similar to that shown
by the non-human bones in the same deposit.
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND 173
Disproportionate Representation: In most
levels, humans are represented mainly by fool
bones, hand bones, skull fragments and teeth.
Pelves are very rare and limb bones uncommon,
although the latter may be at least partially
represented by numerous unidentifiable bone
splinters. The calcaneum and aslragulus are also
generally absent. Reasons for this disproportionate
representation are not obvious. Possibly some
human carcases were dismembered elsewhere, and
the only portions brought to the site were wrists
and hands, lower parts of feel and entire heads.
Although unlikely, it is difficult to otherwise
explain the representation.
DISEASE: Only arthiritis was noted.
JOURAMA
All samples considered, there are 15 juveniles
and 9 adults represented. Of the juveniles there
are two mortality peaks, one between the ages of
3 to 5, and another, the larger of the two, between
8 to 13. The adults for which age estimates have
been made are young adults, except one
middle-aged or old individual. One possibly new
born infant is present. All except some of the
smallest bones are broken, some including
juveniles and adult bones arc charred, and others
marked. Smashed teeth, periotic bones and
calcanea indicate considerable force involved in
breakage. One bone shows what appears to be
traces of red pigment, suggesting possible
ceremonial use of this human bone.
Curious concentrations of bone fragments were
encountered during excavation. These include at
least broken limb, vertebral, foot, mandibular and
cranial eiements but no given cluster represented
a whole individual. The almost vertical orientation
of some long limb bone fragments in these clusters
suggests the remains were either placed in
excavated pits, perhaps as burials, or fell into
sleep-sided crevices in the shelter’s surface. It is
obvious from photographs taken at the lime that
some of these bones were deposited as small
fragments while others show fractures developed
or widened in situ which resulted in their removal
as yet smaller fragments. Some skulls had
complete, undistorled cranial vaults. Because
these were heavily invested with plant roots, they
were removed in pieces during excavation.
Representation: There is some unequal
representation generally of recognizable limb
bones, and commonly of pelves. Foot and hand
bones, and teeth are common.
Mt Roundback
Only three levels produced identifiable human
remains. In the entire deposit, five individuals arc
represented: two unaged juveniles; an adult; a
middle-aged male; and an unaged individual.
Some bones arc charred or marked, and all except
some small bones are broken. Representation of
bone elements is reasonable even considering the
small number of samples.
Kennedy
Human material was only identified in one
level, and represents one unaged individual.
Comparisons of Human Remains
between Sites
The two well-represented sites, Herveys Range
and Jourama, provide the only really meaningful
data for intersite comparisons.
Humans as Food; In general, the Herveys
Range deposit has a much lower percentage of
human bone, in terms of all bone present, than
Jourama. This suggests, assuming humans were
a food source, the Jourama people ate more
people. However, the assumption that all the
human remains represented food for the Jourama
people is questionable because of the evidently
intact nature of some crania. If these particular
individuals were eaten, why weren’t their heads
smashed in order to obtain the brains? The
apparent absence of entire skulls, but obvious
abundance of skull fragments, many of which
were burned, at Herveys Range docs not conflict
with the notion that the humans in that deposit
were a source of food.
Possible interments: At Jourama the
presence of bone clusters of dominalely human
bone, and at least some intact skulls, suggests
burial or at least emplacement in pits, a
phenomenon not obviously present in the Herveys
Range deposit. The non-cranial bones were,
however, almost all broken prior to burial, and at
least one of these broken bones has what appears
to be traces of red pigment. Possibly the bones
or carcases were broken, cooked or burned and
some bones decorated for ritualistic purposes prior
to burial in pits. Whatever the reason, this aspect
of the Jourama deposit appears to differ from that
of Herveys Range.
Age of Humans Represented: In the
Herveys Range deposit there arc more adults than
juveniles represented, while at Jourama there are
far more juveniles than adults represented. The
174
MEMOIRS OFTHE QUEENSLAND MUSEUM
reasons for this difference are obscure and much
would depend on ihe reason the humans were part
of each deposit. In both deposits it seems unlikely
that the ages of the individuals could represent
the structure or mortality curves of the population
unless some bias were involved, such as selective
killing. If they did represent a mortality curve,
surely more infants would be represented. The
Jourama deposit revealed only one possibly
neonate; there were none from Herveys Range.
GENERAL REMARKS CONCERNING
ALL FOUR SITES
Habitats Indicated
The faunas from all four sites suggest they were
obtained from sclerophyl forest or savannah
woodland, with rocky areas and streams. Even the
Kennedy fauna, although extremely impoverished,
contains the Frilled Lizard which (Cogger 1975)
only inhabits dry sclerophyl forests and
woodlands.
The Herveys Range deposit contains the highest
proportion of mammal remains and the lowest
proportion of fish or shells. This suggests it was
farther from water than the other three sites.
Bias in Species Representation
With a few exceptions, the majority of arboreal
and volant species likely to be in the vicinity of
all four sites are unrepresented in the deposits.
These include three genera of gliders, pigmy
possums, most bats and all birds. In fact the only
arboreal marsupial represented, the Brush-tailed
Possum, is also known to spend a considerable
amount of time on the ground. This suggests the
Aborigines were either hunting more in the open
savannah areas, or else had few skills enabling
them to recover arboreal animals. The total lack
of birds is even more puzzling because many spend
a considerable amount of time on the ground.
Also totally missing are the dasyurids, the
native mainly terrestrial carnivores which range
from a mouse to a cat in size. Over 10 kinds could
have been available to them and it is not at all
clear why they aren't represented in the
deposit.
Some of these absences may be explained by
chance sampling of excavation sites. More
extensive examination would be expected to yield
additional laxa.
ACKNOWLEDGEMENTS
Dr Wally Woods (University of Queensland)
supplied the age analysis and identification of the
human remains which formed the basis of the
Appendix. Mr A. Boorsboom contributed
Dingo-chewed bones for comparison. Mr K.
Ackerman kindly collected and provided bones
from Aboriginal campsites from north-western
Australia. Mr A. Easton took the photographs.
LITERATURE CITED
Most of the specific references regarding mammal
distributions are not noted above because most are based
on Queensland Mu.scum records.
ARCUttR, M., 1975. Abnormal dental development and
its significance in dasyurids and other marsupials.
Mem. Qci Mus. 17: 251 65.
1978. The nature of the molar-premolar boundary in
marsupials and a reinterpretation of the homology
of marsupial cheekteeth. Mem. Qd. Mus. 18(2):
157-64.
Bartuoi oma}. a., 1975. The Genus Macropus Shaw
(Marsupialia: Macropodidae) in the upper Cain-
ozoic deposits of Queensland. Mem. Qd Mus 17(2):
195-235.
Brayshaw. H. C., 1977. 'Aboriginal material culture in
the Herbert/Burdekin district, north Queensland.'
PhD Thesis, .lames Cook University.
CoGGFR. H. G., 1975. ‘Reptiles and Amphibians of
Australia’. (A H. and A. W. Reed: Sydney).
Ridi\ W. D. L., 1970. ‘A guide to the native mammals
of Australia.' (Oxford Univ. Press: Melbourne).
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND 175
APPENDIX
Summary or Human Rrmains Ir^HNTir-ii t) by
W. Woods, witd Commi;nts
Herveys Range
Surface At least two individuals: one adult, one
(21 fragments) juvenile. Charring of skull and post-
cranial fragments. Most parts of
skeleton are represented except pelvis
and limbs. Dominance of hand, foot and
skull fragments. Most bones broken.
Only one tooth present.
Spit 1 At least four individuals: baby 6-9
(36 fragments) months; juvenile lO-Il years; middle
aged adult. Charring and smashing of
teeth and bones, including those of
juveniles. Missing arc pelvis, femur,
tibia, ulna. Dominance of hands, feet,
skull and teeth. Some teeth of juveniles
(VII) have roots which are charred,
suggesting they may have been smashed
out of skulls before being burned.
Spit 2 At least three individuals: child approx-
(30 fragments) imately 3 years; child 10- 1 2 years; adult
(age ?). Charring and smashing of bones
— particularly noteworthy is smashed
tympanic region which is very dense part
of skull requiring considerable force to
break. Missing are almost all limb bones
except fibula, pelvis, sacrum, and
scapula. Dominance of hand and foot
elements, skull fragments, teeth and
some vertebrae.
Spit 3 At least two individuals: an adult,
possibly male, of 30+ years: a juvenile
(age ?). Charring only noted on adult
skull fragment (HVII), and a juvenile
scapula fragment (HVII). Adult had
arthritis in one foot. Missing are most
portions of pelvis, and major limb bones.
Dominance of fool and hand bones,
tooth and skull fragments.
Spit 4 At least two individuals: a middle aged
(22 fragments) adult; juvenile. Charring noted only on
two juvenile skull fragments (HVII).
Small samples but still no remnants of
limb bones, pelves, scapulae, sacra.
Dominance of skull fragments, teeth,
foot and hand bones.
Spit 5 At least one individual: a ?juvenile male.
(1 fragment) No charring noted. One zygomatic
(skull) bone only of human remains
identified.
Spit 6 At least two individuals: possibly a
(3 fragments) juvenile of approximately 12 years; one
adult. No charring noted. Rib, tooth and
toe bone, only three fragments.
Spit 7 At least one individual: one adult. Small
(10 fragments)sample, but no burning noted. Fragmen-
tation common. Some marking on tibial
fragments. Missing are pelvis, femur,
fibula, arm (except hand elements) and
most of axial skeleton. Present are skull
fragments, scapula fragment, hand
elements, tibial fragments, and atlas
fragment.
Spit 8 At least two individuals: a juvenile
(3 fragments) approximately 12 years; an adult (age
?). Small sample, but no charring noted.
Present arc finger, vertebra fragment
and spine of scapula.
Spit 9 At least two individuals: a juvenile
(8 fragments) approximately 6 years; an adult (age?).
Charring of skull, vertebra. Missing are
pelvis, limbs, feet, hands: only represent-
ed by skull fragments, isolated teeth,
vetebral fragment.
Spit 10 At least two individuals: a juvenile
(4 fragments) (age?); an adult (age?). No charring
noted. Only represented by fragments of
skull, foot and femur.
Spit 1 1 One adult (age?). No charring noted.
(5 fragments) Represented only by fragments of rib,
femur, skull and one trapezoid.
Spit 12 At least two adults (age?). No charring
(8 fragments) noted. Represented are fragments of rib,
foot bones, vertebrae, and some hand
bones.
Spit 13 At least two adults (age?). Charring of
(16 fragments) skull fragment. Missing are all limbs,
pelvis, axial skeleton (except skull).
Dominance of foot and hand
elements.
Spit 14 At least one individual: an adolescent or
(16 fragments) young adult. No charring noted. Missing
are limbs (except hands and feet),
pelves, sacrum. Dominance of foot and
hand elements, and less so, skull
elements.
Spit 15 Possibly a single adult. No charring
(15 fragments) noted. Missing arc all major limb bones
except femur, pelvis, sacrum, scapula
and skull. Dominance of hand and foot
elements. For the first time a calcaneum
(fragments) is represented.
Jourama
Surface One individual (age?). No damage
(1 fragment) noted. One right cuboid only present.
Spit 1 At least two individuals: one juvenile
(11 fragments) 8-10 years; one adult (age?). Charring
of a long bone fragment. A broken
periotic bone indicates considerable
force involved in smashing skull. Some
long bones were evidently cut, or at least
have cut marks. Missing are most bones
but representation includes skull, teeth,
hand and at least one long bone.
176
MEMOIRS OF THE QUEENSLAND MUSEUM
Spit 2 At lo^st four individuals; juvenile
(32 fragments) approximately 4 years; juvenile approx-
imately H years; juvenile approximately
13 years; adult in early twenties.
Charring of juvenile skull fragments and
a mandible. A juvenile ulna and radius
show heat marks (cooking without
charring?) and scratches. Missing are
pelves and possibly limb bones of adult.
Some limb bone fragments arc present
but, except for juvenile radius and ulna,
arc evidently unidentifiable. Dominant
are isolated teeth.
Spit 3 At least four individuals: a juvenile
(115 fragments)approximalcly 10 years; a juvenile
approximately 16 years; a subadult
female; and an adult (age?). Charring
noted on some skull fragments only, i.e.
uncommon. Breakages other than nor-
mal smashing includes scratch marks on
ileum of young female; on rib fragments;
juvenile long bones; metatarsal has piece
removed (?); metacarpal with scratch
marks; scratches on adult femur; scratch
marks on rib fragments and on young
adult mandible fragment. One mandible
fragment suggests teeth were removed
(or naturally lost?) from bone after
death. Calcaneum is smashed, a feat
taking considerable force. All elements
represented by at least one fragment, but
scarce arc pelvic fragments, vertebrae
and limb bones. Common are isolated
teeth, foot and hand bones.
Spits 3 & 4 At least two individuals: a child of
(32 fragments) approximately 8 years; a Juvenile of
approximately 10 years. No charring
observed. Most long bones show
scratches. One long bone shows traces of
?red pigment. There are no obvious
missing elements, except there are no
hand or foot bones — the reverse of the
usual situation.
At least two individuals: a child of
approximately 10-11 years; an adult
(age?). Charring noted on mandible
fragments; long bone fragments; a
patella; skull fragment. Splitting or
scratching noted on rib fragments, long
bone fragments, etc. A cuboid and
calcaneum are broken, considerable
force being required. Some teeth are also
broken — also requiring considerable
Spit 4
(77 bones - -
nearly 100
fragments)
Spit 5
(87 +
fragments)
Spit 5
north end
(20 + small
fragments)
Spit 6(a)
(46 + many
fragments)
Spit 6(b)
(60 + many
fragments)
Spit 7
(6 fragments)
Kennedy
Spit 9 (FII)
force. Most elements are represented,
including pelves. Dominance of hand
and foot bones, and isolated teeth.
At least five individuals; an infant
(possibly just new born); a juvenile of
approximately 4-5 years; a juvenile of
approximately 11-12 years; two young
adults. Charring noted in skull frag-
ments and long bone fragments. Most
bones are represented except pelvis and
identifiable long bones, except femur —
some fragments of long bones may
represent others. Dominance of isolated
teeth only.
Probably a single adult male. A talus is
charred. Scratch marks noted on
fragments of femur, clavicle and ribs.
Representation reasonable. Missing are
pelvis, arms and hands, although scapula
and clavicle present. Legs and feet
dominate.
At least three individuals: a juvenile
approximately 3-4 years; an adult male
(age'Q; an adult female (age?). Charred
bones include scapula fragments and
long bone fragments. Splitting and/or
scratching noted on femur fragment,
long bone fragments, also on fragments
of pelvis and scapula. Most bones are
represented except arms (hands repre-
sented). Several pelvic fragments pre-
sent, including juvenile and adult.
At least three individuals: two juveniles;
a middle aged male. Charring is not
spccificially noted but many of the bones
are said to show evidence of heat
exposure. Scratching and/or splitting
are noted in tibial, fibular, femoral,
radial, ulnar and clavical fragments.
One vertebra shows advanced arthritis.
Representation is reasonably even — no
obvious imbalance.
At least two individuals: a juvenile of
approximately 4- 5 years; and an adult
(age?). No charring noted. Scratch
marks on maxillary fragments. Sample
small, so unequal representation not
surprising. Fragments all cranial and
mandibular.
Not clearly more than one individual
(age?). Fragments of right pubis and
ribs, also foot bones present.
MEMOIRS OF THE QUEENSLAND MUSEUM
Platb 28
Representative specimens from the Kennedy and Jourama
excavations: A, F9199, five pieces of bone from Jourama, C III 5,
which have been thoroughly chewed and/or cut, XI; B, two views of
F9200, Jourama, C III 4, showing possible incision (i) in profile and
plan view, XI; C, F9201, Herveys Range, H VII 5, humerus fragment
showing charred ends and what is referred to in this paper as a cooked
medial region, that is it is discoloured but not charred, X2; D, F9184,
Herveys Range. H VI! I, Macropus cf. M. parryi (see text), left
dentary with crown of 1] broken off, possibly intended to be a scraper,
XI; E, F9188, Hervey's Range. H VII surface, Petrogale sp. cf. P.
penicillata, juvenile right dentary showing conical depression
interpretated as a tooth mark (t) XI; F, F9I89, Herveys Range,
H VII 2, Trichosurus cf. T. vulpecula. right maxillary fragment, XI;
G, F9198, Herveys Range, H III 4,? Mesembriomys gouldii, LI|, XI;
H, F9205, Herveys Range, H VI 2, small rodent, XI; I, F9I97,
Herveys Range. H VII 1, Canis famiUaris. LdP4, XI; J, F9195,
Herveys Range, H V 3, Macroderma gigas, left dentary fragment;
XI; K, F9I96, Herveys Range, H VII 4, Pteropus scapulatus, left
maxillary fragment, XI ; L, F9I92, Herveys Range, H VI surface, P.
scapulatus. left dentary fragment showing no alveoli for otherwise
variably present M3. XI; M-N, F9204, Herveys Range, H III 5,
anterior end of a left dentary of an unknown but apparently new
diprotodont marsupial, stereopair occlusal view (M) and lateral view
(N), X2; O, F9206, Herveys Range, H VI 3, Homo sapiens, charred
fragment of a juvenile right dentary, X2; P, F9207, Herveys Range,
H VII surface, Chlamydosaurus kingii. fragment of a right maxilla,
XI; Q, F9203, Herveys Range, H VI 2, Tiligua scincoides, fragment
of a left dentary, XI; R. F9202. Herveys Range, H VII 2, isolated
right quadrate of a large python skull, XI; S, F9194, Herveys Range,
H VI 1. palatine of a large python cf. Morelia sp., XI; T, F9208,
Herveys Range, H III 6, dentary fragment of a fish, XI.
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND Plate 28
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 29
A. General view of the Herveys Range Site. The rock shelter excavated
is under the breakaway of the large boulder in the central
background.
B. Bone cluster in excavation at Jourama, G III 5. Most bones visible
are human and, with the possible exception of the dentary (lower
left) and astragulus (upper middle), all show marked predepositional
smashing. Limb bones at upper right are not in the round, each
being represented by only one side of the shaft. Ruler is one foot
long. Photo taken looking WNW.
ARCHER AND BRAYSHAW: FAUNAS FROM EXCAVATIONS IN NORTH QUEENSLAND Plate 29
Mem. QdMus. 18(2): 179-83, pi. 30. [1978]
PTERASTERICOLA VIVIPARA N. SP., A PARASITIC
TURBELLARIAN (RHABDOCOELA: PTERASTERICOLIDAE) FROM THE
CROWN-OF-THORNS STARFISH, ACANTHASTER PLANCI
L. R. G. Cannon
Q ueensland Museum
ABSTRACT
Pterastericola vivipara n. sp. feed on epithelial cells of the pyloric caeca of Acanthaster
planci. Ciliated larvae may hatch from eggs within these small red worms and burrow through
the parent’s body. It is the second member of this genus described and the first record of
this family (Pterastericolidae) from the Southern Hemisphere.
Turbellaria have frequently been reported from
the gut of echinoderms, principally from
holothurians and echinoids (Jennings 1971, 1974).
Four species have been found in asteroids. These
are the rather aberrant Acholades asterias
Hickman and Olsen, 1955 (F. Acholadidae) which
lives encysted in the walls of the tube feet of
Coscinasterias calamaria in Tasmania and three
species in the family Pterastericolidae, viz.,
Pierastericola fedotovi Beklemishev, 1916,
Triloborhynchus astropectinis Bashiruddin and
Karling, 1970 and T. psilastericola Jespersen and
Lutzen, 1972. All three pterastericolids occur in
the pyloric caeca of asteroids from northern
European waters.
No endo-parasites have as yet been described
from the Crown-of-Thorns Starfish, Acanthaster
planci, which has received so much attention over
the past decade in relation to destruction of living,
hard corals in the Indo-Pacific (Chesher 1969;
Vine 1970; Endean 1973). Investigation of the
Crown-of-Thorns along the Great Barrier Reef for
associated parasites and commensals (Cannon
1972, 1975) led to the discovery of the worm
herein described.
MATERIALS AND METHODS
Starfish were transported alive as described by
Cannon (1973) to Brisbane for examination in the
laboratory. Worms were only found in the pyloric
caeca although all internal organs were examined.
Routinely worms were detected by gently
squashing sections of pyloric caecum between
glass while being examined with a stereoscopic
microscope. Worms removed from caeca were
examined alive in filtered seawater. Whole mounts
were prepared from worms fixed in cold 10%
Formal/Saline, cold A.F.A. and cold Gilson’s
fluid, and stained with either Mayer’s or
Kornhauser’s haemalum. Sections at Ip were
obtained from cold, Bouin’s fixed worms,
embedded in 60°C paraffin. These sections were
stained with Haemaxtoylin and eosin. Measur-
ements were made with a micrometer eyepiece,
drawing was with the aid of a camera lucida.
Pterastericola vivipara n. sp.
(Fig. 1, Plate 30)
Material examined
Hoi.otype: from pyloric caecum of Acanthaster
planci Linnaeus, Centipede Reef, 9/vi/l974, L.
Cannon, QM G10313 (wholemount).
ParatypeS; from pyloric caecum of A. planci,
Centipede Reef, 9/vi/i974, L. Cannon, QM G10314-5
(wholemounts) G103I6 (sections).
Other material examined included live specimens
gently squashed under coverslip pressure.
Diagnosis
Small pyriform turbellarian, epidermis comple-
tely ciliated, lacking rhabdoids; mouth sub-
terminal, pharynx bulbous, doliiform, but gut
dorso-ventrally orientated when fixed, intestine
sac like; lacking protonephridia; gonopore single,
ventral; ovary and testis single, compact, lateral;
vitellaria in two lateral bands, male system with
180
MEMOIRS OF THE QUEENSLAND MUSEUM
copulatory armature of hook and stylet; ciliated
larvae may hatch within and burrow out of
parent.
Host: Acanthaster planci Linnaeus
(Asteroidea).
Habitat: Pyloric caeca.
Locality: Great Barrier Reef, Australia.
Description
Worm (Fig. 1) pyriform tapering to blunt point
posteriorly; slightly dorso-ventrally flattened when
swimming or crawling, usually a ball in situ;
anterior parenchyma with red pigment particular-
ly in large specimens; measuring x ± s = 722
± 87 X 590 ± 90// (range: 630-891// x.
387-702//^ N = 14); protonephridia lacking.
Body Wall and Musculature: Entire worm
covered with ciliated epidermis 7 to 8// thick, cilia
4 to 6// long, basement membrane thin; (hot
fixation causes epidermis to lift clear of
membrane); rhabdoids lacking; clusters of large,
deeply staining globules at base of cilia cover
surface of young worms, but become increasingly
scattered as worms mature. Body wall muscula-
ture of outer circular fibres, inner longitudinal
fibres and between scattered oblique fibres;
dorso-ventral muscles more common anteriorly
than posteriorly; prominent musculature sur-
rounds pharynx and cirrus pouch which are highly
mobile in life.
Parenchyma: Large poorly staining cells,
often vacuolated, fill posterior to give a foamy
appearance, but only in life; these vacuolated cells
readily take up aqueous neutral red and rapidly
shrink in hypertonic salt solution; other large cells
throughout about 18 x 12// staining with
haematoxylin grey to blue and containing
vacuoles. Sub-cpidermal glands common through-
out surface parenchyma, their ducts anastamose
discharging basic secretion through goblet-like
ducts between epithelial cells; several, deeply
eosinophilic glands in young worms run from
mid-fore body to anterior tip.
Nervous System: Bilobed neural mass mid
way between mouth and anterior end; several
anteriorly directed nerve tracts; two, principal,
lateral nerve cords run posteriorly; sense organs
not detected.
CANNON: PARASiTIC TURBELLARIAN FROM ACANTHASTER
181
Digestive System (Plate 30,a): Mouth
without lips, sub-terminal, ventral about 120//
from anterior end; short, buccal cavity lined with
large cuboidal cells with finely granular cytoplasm
leads to large, muscular pharynx x ± s = 111
± 14 X 93 ± 15// (range: 89 - 133 x 71 - 116//,
N = 14); pharynx dorso-ventrally orientated m
fixed material, opening into small oesophogial
cavity lined with large, weakly eosinophilic,
columnar cells which can extend through pharynx;
large, saccate gut lying in dorsal body extends
slightly forward from pharynx as well as extending
posteriorly; gut separated from parenchyma by
thin layer of cells below tall, columnar
gastrodermis; three types of gastrodermal ceils (a)
phagocytic cells with intracellular food vacuoles,
(b) secretion cells with evenly stained cytoplasm
and (c) secretion cells with foaming cytoplasm;
gut outpockets in large specimens.
Male Reproductive System; Single testis
lateral, just posterior to pharynx; sperm
production synchronous; simple vas deferens leads
directly to cirrus pouch where, passing through a
narrow opening, it swells to form a seminal vesicle
which has a narrow opening at base of male
cuticular armature; armature (Plate 30, b), a plate
with two chilinous horns — one terminating in
a closed stylet x ± s = 34 ± 3// long (range 31 -
38//; N = 5), the other is hooked x ± s = 40 ±
3// long (range 38 — 45//, N = 5) with a narrow
pore at end, opening away from stylet; sperm and
seminal fluids are discharged through pore of
hook; armature extends into a common genital
atrium which opens mid-ventrally Just posterior to
mouth; two sets of glands have long ducts which
lead to and enter cirrus pouch beside vas deferens
and coil about seminal vesicle; the less common
glands have granular cytoplasm, the more
common ones are strongly eosinophilic, these have
brown, foamy cytoplasm in life and secrete
droplets (cementing?) through hook.
Female Reproductive System: Single, bipar-
itite ovary with inner layer of small, dividing cells
and outer region of larger, maturing oocytes;
ovary lateral, posterior to testis; oocytes seen
passing down oviduct to ootype/mehlis gland
complex; oviduct joined first by duct from seminal
receptacle and then by joint vitelline ducts where
they enter ootype; deeply staining basophilic cells
surround ootype; uterus not evident in large
specimens, eggs apparently pass into parenchyma;
female pore with a short muscular region behind,
opens in common genital atrium: some eggs seen
extruded from pore, others fill parenchyma, in ail
stages of development, many hatching within
worm; vitcllaria paired, extending vcntrally and
laterally at level of common genital pore to
margins of body where they expand dorsally and
a little posteriorly; ovary and testis on right, thus
left vitellarium is a little more extensive: seminal
receptacle a simple, large, club-shaped sac filled
with sperm in large specimens, but appears to
open to the ventral surface through a small pore
in small, less mature worms; muscle bundles (a
pscudovagina) can sometimes be seen (Plate 30,
c). Eggs sub-spherical to spherical measuring x ±
s = 84 ± 6 X 81 ±7// (range 76 - 98 x 71 -9H^r,
N = 1 I ) with golden tanned walls; some eggs laid,
others crumble (Plate 30, d) and ciliated larvae
which hatch have been observed live (moving
about within the parent) and in fixed and
sectioned specimens (Plate 30, e).
Remarks
This worm is similar to Pterastericola fedotovi
Beklemishev, 1916 which was partially redes-
cribed by Karling (1970). The worms share the
same general shape, pigmentation and ciliation.
Reproductive structures are more anterior and
vitcllaria less extensive than in P. fedotovi.
however. Both testis and ovary are lateral in P.
vivipara whereas the ovary in posterior and
median in P. fedotovi. The male cuticular
armature of P. vivipara has two sub-equal spines
about 30 to 40// long and about 10 to 15// apart.
The armatures of P. fedotovi illustrated by
Beklemishev (1916) and by Karling (1970) do not
curve similarly but they are larger, about 80//;
unlike that of P. vivpara the hook appears shorter
than the spine. Host and locality differences
between P. fedotovi and P. vivipara are
considerable.
Habitat and Distribution
Worms were found restricted to A. planci and
were found only in starfish from the central Great
Barrier Reef (Cannon, 1975). They were found
more commonly in winter, but showed no
correlation of incidence with size or sex of host.
Cannon (1975) showed too, that though the
worms occurred randomly throughout the caeca
and fed upon the lipid rich caccal epithelial cells
of A. planci, they were not pathogenic.
Biology
Worms may be seen as small, red balls tightly
pressed in outpockets of the caeca similar to the
habit of Trilohorhynchus astropeciinis
(Pterastericolidae) described by Bashiruddin and
Karling (1970). Though infection levels are rarely
182
MEMOIRS OF THE QUEENSLAND MUSEUM
high, when they are, large numbers of worms of
all sizes occur in the caeca from unpigmented
juveniles to adults containing many eggs. Eggs
may be laid, or may hatch within the parenchyma
of the parent worm from which juveniles
presumably break free (hence the specific name).
Thus large populations no doubt build up within
one starfish; transfer between hosts via eggs may
be possible.
Larval worms are small and ciliated when first
they hatch and measure x±s = 81 ± 10x65
± Afi (66 — 99 X 57 — 73^, N = 10). Deeply
staining eosinophilic glands lie laterally and open
anteriorly. The function of the glands may be to
aid hatching, but the movement of worms through
the parent's body suggests they may be used to
aid escape. Juvenile worms have not been observed
in the coelomic fluid of the starfish, nor have adult
worms, so penetration of the host caecal
epithelium seems unlikely.
Discussion
The present species brings the total to five
turbellarians recorded from Asteroidea. All are
rhabdocoels contained within three genera, —
Triloborhynchus, Pterastericola and Acholades.
It is perhaps noteworthy that each genus has been
recorded from hosts in different orders of
Asteroidea; Triloborhynchus has been found in
starfish of the O. Phanerozonia, Pterastericola
from hosts of the O. Spinulosa, and Acholades
from O. Forcipulata. The latter monotypic genus
is unlike the others in either morphology or
habitat being encysted within the walls of the tube
feel of a Tasmanian starfish (Hickman and Olsen,
1955). Bashiruddin and Karling (1970) and
Kariing (1970) maintained Triloborhynchus and
Pterastericola should be contained in the F.
Pterastcricolidae despite the reduced ciliation and
posterior adhesive organ in the former genus,
characteristics said by Karling (1970) to link
Triloborhynchus with the Monogenea. Stunkard
and Corliss (1951) considered Pterastericola
related to the F. Umagillidae. The true affinities
of these worms must await more detailed
study.
It is not clear if the pseudovagina of
Pterastericola functions in copulation (Karling,
1970). Unlike Karling (1970) who had only
mature specimens of P. fedotovi, I had immature
as well as mature specimens of P. vivipara and
in one not fully gravid worm the pseudovagina
appeared to open to the surface, whereas in
sections of mature specimens no opening was seen.
Karling (1970, Fig. 10) observed a break in the
epidermis of P. fedotovi. It seems possible that the
vagina is functional in young animals, but
atrophies after copulation or as the worm matures.
Alternately the musculature of the pseudovagina
may serve as anchoring tissue for the stylet during
copulation, enabling the sperms to be innoculated
through into the parenchyma. Certainly Beklemi-
shev (1916) reported wandering sperms in the
parenchyma of P. fedotovi and hypodermic
impregnation is known among the rhabdocoels
(Hyman 1951, p.l25).
The peculiar habit of eggs hatching within the
parent body so that young are released into the
parenchyma has been described for another
Australian turbellarian Avagina vivipara
Hickman (1956) from a heart urchin
Echinocardium cordatum. This habit clearly
shortens the life cycle and decreases dispersal,
enabling large populations to build up in the one
host. Although Hickman (1956) gave few details
it seems likely the ciliated young of both these
worms wander through the parent tissues until
they eventually break free into the lumen of the
gut of the host. According to Hyman (1951,
p-135) members of the rhabdocoel group
Typhloplanida often produce two kinds of eggs —
thick shelled resistant ones and thin shelled
subitaneous eggs from which young may hatch
within the parent. Such eggs are presumed an
adaptation to quickly increase the populations of
worms at favourable times, but in temporary or
unpredictable environments. Echinoderm popula-
tions are known to form dense aggregations
(Reese, 1966). Worms able to multiply within
their hosts when these hosts are rare would be
advantaged. Perhaps this mode of reproduction
alternates with normal egg laying when host
populations are dense, similar to the Typhoplanids
(Hyman 1951). On one occasion eggs were found
in mucous trails left by infected starfish in
aquaria. The method of feeding of Acanthaster
would favour reinfection from mucous covering
the substrate over which it everts its stomach.
ACKNOWLEDGMENTS
This work was made possible by a grant from
the Advisory Committee into Research on the
Crown-of-Thorns Starfish. Technical assistance
was ably rendered by Con Boel and Biomarine
Pty. Ltd., Townsville. Thanks are extended to J.
C. Pearson for help and discussion.
LITERATURE CITED
Bashiruddin, M. and Karling, T. G., 1970. A new
entocommensal turbellarian from the sea star
Astropecten irregularis. Z. Morph. Okol. Tiere 67:
16-28.
CANNON: PARASITIC TURBELLARIAN FROM ACANTHASTER
183
Beklemishev. V., 1916. Sur les turbellaries parasites de
la cote Mourmanne. I! Rhabdocoela. Trudy
leningr. Obshch. Estest. 45: 60-79.
Cannon. L. R. G., 1972. Biological associates of
Acanthaster plana . Pp 9-19 in Crown-of-Thorns
Seminar, University of Queensland. 25 August
1972 (Department of Education and Science:
Canberra).
1973. An artificial gill for the Crown-of-Thorns?
Turtox News 50: 26-7.
1975. Observations on a parasitic turbellarian
from Acanthaster planci . P.p 39-54 in ‘Crown-
of-Thorns Seminar Proceedings.’ Brisbane, 6
September 1974. (Aust. Gov. Printing Service:
Canberra).
Chesher. R. H., 1969. Acanthaster planci : Impact on
Pacific coral reefs. Westinghouse Electric Corpora-
tion Report to U.S. Dept, of Ini. Pub. No.
187631.
Endean. R., 1973. Population explosions of Acanthaster
planci and associated destruction of hermatypic
corals in the Indo-West Pacific Region. In Jones,
D. A. and Endean. R, (Eds.) ‘Biology and Geology
of Coral Reefs. II.’ (Academic Press: New
York).
Hickman. V. V., 1956. Parasitic turbellaria from
Tasmanian Echinoidea. Pap. Proc. R. Soc. Tasm.
90: 169-81.
Hickman, V. V. and Olsen. A. M., 1955. A new
turbellarian parasite in the sea star Coscinastarias
calamaria Gray. Pap. Proc. R. Soc. Tasm. 89:
55-63.
Hyman. L. H., 1951. ‘The Invertebrates: Platyhel-
minthes and Rhynchocoela. The acoelomate
bilateria.’ Volume II. (McGraw-Hill; New
York).
Jennings. J. B., 1971. Parasitism and Commensalism
in the Turbellaria. Adv. Parasitol. 9; 1-32.
1974, Symbioses in the Turbellaria and their
implications in the study of the evolution of
parasitism. In Vernberg. W. B. (Ed.). ‘Symbiosis
in the Sea'. (University of South Carolina Press:
Columbia).
Karling. T. G., 1970. On Pterastericola fedotovi
(Turbellaria), commensal in Sea Stars. Z. Morph.
Okol. Tiere 67: 29-39.
Reese, E. S. 1966. The complex behaviour of
echinoderms. In Boolootian. R. A. (Ed.)
‘Physiology of Echinodermata’. (John Wiley
(Inlerscicnce): New York).
Stunkard. H- W. and Corliss. J. O., 1951. New
species of Syndesmis and a revision of the family
Umagillidae Wahl. 1910 (Turbellaria: Rhab-
docoela). Biol. Bull. mar. biol. Lab., Woods Hole.
101: 319-34.
Vine, P. J., 1970. Densities of Acanthaster planci in the
Pacific Ocean. Nature, Lond. 228: 341—2.
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi ATF 30
(a) Section through gut of P. vivipara in situ (Scale is 100//).
(b) Male copulatory armature (Scale is 50//).
(c) Pscudovagina in sub-adult specimen (Scale is 100//).
(d) Egg capsule after hatching within parent (Scale is 75//).
(e) Section of adult with larva in tissues (Scale is 100//).
CANNON: PARASITIC TURBELLARIAN FROM ACANTHASTER
Plate 30
Mem. QdMus. 18(2): 185-212. [1978]
THE MACROBENTHOS OF THE
MIDDLE BANKS AREA
OF MORETON BAY*
W. Stephenson,
S. D. Cook,
and
S. J. Newlands
Zoology Department, University of Queensland
ABSTRACT
This study was undertaken to quantify certain aspects of the benthic biota of the Middle
Banks area, with a view to predicting the effects of sand removal for use as Till' in enlarging
Brisbane Airport. Fifty-seven sites lying roughly in a rectangle 9 km x 1 -5 km were sampled
at 3 month intervals from September 1972 to June 1974. Each sample comprised duplicate
catches with an OTm^ Smith-Mcintyre grab. The northern sites lie in the area of the proposed
excavation and the southern ones (in deeper water) in a prawn fishing area. There is a sharp
division between coarser northern and finer southern sediments, and suggestions of a progressive
coarsening of sediments during the study.
A total of 468 ‘species' were originally sorted from the collections, a noticeably richer and
more diverse biota than Bramble Bay (on the western side of Moreton Bay). To reduce the
size of the data matrix to analyse, the first classifications were undertaken with successive
lots of 50 taxa x 57 sites x 8 times. Sites were clearly separable into northern and southern
subareas and the same broad area pattern was generated by abundant and less abundant
species. Subsequent analyses involved separate consideration of the two sub-areas, and
eventually 12 site-groups were recognised with their characterising species. Effects on the biota
of current dredging of a navigational channel are detectable.
There were problems in separating microtopographical changes, due to inaccurate relocation
of sites, from chronological changes. It appears that of the species investigated about 10%
have recurrent seasonal patterns, about 40% have sequential patterns, and about 45% have
obscure patterns, possibly influenced by microtopography. The .sequential patterns are akin
to those obtained in preflood data from Bramble Bay, with occupancy of appropriate areas
for tolerably brief periods (e.g. 6 months) and then giving way to other .species. Few- of the
sequential breaks appear related to abiotic conditions.
It is suggested that almost the entire biota is in a continual stale of flux, with most species
either annuals or ‘subannuals', with few attaining sexual maturity in the area, with the
population dependent upon recurrent recruitment, and with intermittent but severe predation
by fish as an important possibility. The consequences of temporary human disturbance to the
area are discussed.
A proposal to enlarge Brisbane Airport would
involve Tilling* a considerable area with fine sand
from the Middle Banks portion of Moreton Bay.
The present work was requested by the (then)
Commonwealth Department of Housing and
Construction and by the Queensland Department
of the Co-ordinator General hopefully to predict
the effects of removal of the sand on the
benthos.
There is already human interference in the
Middle Banks area, with the south-western portion
kept dredged to 12 m for shipping purposes. This
breaks an otherwise W shaped area of banks
shallower than 10 m and allows the considerable
Manuscript accepted October 1976.
186
MEMOIRS OF THE QUEENSLAND MUSEUM
COMBOYURO POINT
SHARK SPIT
CENTRAL BANKS
27°10'S
EAST CHANNEL
TANGALOOMA POINT
MIDDLE BANKS
153 10 E
STRADBROKE
ISLAND
27^30'S
BRAMBLE BAY
OLD P LE L GHT
SERPENTINE CREEK
MORETON
ISLAND
BRISBANE RIVER
NORTH
Fig. 1: Moreton Bay showing localities mentioned in the text, sampling area enclosed in rectangle. Depths from
Admiralty chart 236 of 1973: > 20 m dark stipple, 10-20 m pale stipple, 5 m thin contin. line, high water
mark thick contin. line.
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
187
tidal currents in the area more easy access to the
deeper waters to the south.
The area initially selected for dredging lay on
the eastern edge of the Middle Banks, abutting
on the East Channel (see Fig. I). The effects of
dredging this area are likely to be numerous. First,
benthic organisms and their environment would be
removed from a relatively small area, and there
would be an interval before this was recolonised.
Secondly, the local topography would alter as
sediments filled the excavated area. Thirdly, if the
excavation led to generally increased current
velocities in the area as a whole, it could lead to
general erosion. While such erosion could
facilitate navigation it could affect the benthos
more widely, and possibly could affect the
adjacent shores of Moreton Island. Fourthly,
material suspended in the water during dredging
would be distributed by the tides. The most
important effects could well be due to the slower
flood tide which would carry material into the
deeper waters to the south. Here there is known
to be a rich benthos (Stephenson, Williams and
Fig. 2: Location of sites within sampling area. Depths from 1972 survey by Queensland Dept, of Harbours and
Marine: 10 fm broken line, 5 fm thin contin. line, high water mark thick contin. line. Sites as solid circles,
beacons or buoys as crosses.
MEMOIRS OF THE QUEENSLAND MUSEUM
Lance 1970) and there is also a seasonal fishery
for peneid prawns.
After considering the above, we advised against
dredging near the existing dredged break in the
banks because it was fell this would give maximal
erosion and most widespread biotic effects and it
is gratifying to record that this advice was
accepted. The position tentatively selected was in
the vicinity of our sampling sites 22, 25, and 26,
(see Fig. 2).
A past survey of the area (Stephenson, Williams
and Lance 1970) was made with a naturalists
dredge and subsequent work at Peel Island in
Moreton Bay (Stephenson, Williams and Cook
1974) has shown that this failed to reveal a
substantial fraction of the benthos. Present
experience showed that the van Veen grab used
in near-concurrent studies in Bramble Bay
(Stephenson. Raphael and Cook 1976, Stephen-
son, Cook, and Raphael 1977) was ineffective
at Middle Banks except in some areas during the
brief periods of slack tide. Hence an 0- 1 m^
Smith-Mcintyre grab was employed and revealed
a surprisingly rich and varied biota.
Past experience at both Peel Island and
Bramble Bay has shown appreciable time changes
in the benthos and to make even approximately
effective predictions of the likely effect of sand
dredging at Middle Banks, it was felt that
sampling should continue at three month intervals
for at least two years. Sampling began in
September 1972 and before the work was due to
be completed two natural catastrophes had
occurred. These, a major flooding of the Brisbane
River in January 1974 followed by a cyclone in
March 1974, produced very marked effects on the
biota of Bramble Bay, but preliminary examina-
tion of the present catches suggested only slight
effects on the Middle Banks benthos, and so the
project was concluded after the two years.
Biotic Sampling
Because of the rapid tidal currents, the effects
of dredging will be greater to the north and south
of the dredged area than to the east and west.
Hence the sampled area was roughly in the form
of a rectangle ca 9 km x U5 km. At both northern
and southern extremities there were sufficient
sighting marks for sites to be arranged on a grid
ca 0-4 km apart but in the centre of the area due
to a paucity of marks they lay on traverses ca 1
km apart.
Fifty-eight sites were originally sampled but
because one (site 39) was overlooked on one
occasion, for present purposes we consider 57 sites,
with original sites 40-58 renumbered 39-57.
(Labelled specimens to be deposited in the
Queensland Museum carry the original number-
ing.) The positions of the final sites are shown in
Fig. 2.
Locations of sites were by horizontal sextant
angles on conspicuous objects in the vicinity
(landmarks or beacons) and inaccurary in
relocating sites is believed to be not greater than
50 m.
At each site on each occasion, duplicate catches
were made with the O-I Smith-McIntyre grab.
Catches were sieved through a final aperture of
1 mm and the two catches pooled, preserved and
the biota separated from the residual sediment at
base. The resultant biota is referred to as a
sample.
Samples were obtained on eight occasions from
September 1972 at three month intervals until
June 1974, and these are referred to below as
times 1-8.
Considerable difficulties were encountered in
identifying species, this reflecting the paucity of
work in similar environments in eastern Australia.
Reference collections were established for what
were adjudged to be species, and these were later
referred either to named specimens collected in
previous benthic surveys or to group experts
(acknowledge later). In several cases the reference
specimens belonged to more than one species, and
these polyspecific taxa are indicated later. There
were a few cases of ‘oversplitting’ and these wore
not apparent until all the numerical analyses were
completed.
The Area of Sampling
1. General Description: The northern exit
from Moreton Bay consists of alternate banks and
channels which, south of the narrowest portion,
run generally from a north-east to south-west
direction. This portion has been called the
‘north-eastern corrugations’ by Stephenson,
Williams and Lance (1970). The main features
from west to east are Western Banks, Pearl
Channel, Central Banks, Main Channel, Middle
Banks, and East Channel. Larger vessels leave
Moreton Bay through a dredged cutting in Middle
Banks, then via the East Channel until it fuses
with the Main Channel. North of the narrowest
portion there are extensive banks to the north-west
of Comboyuro Point, and the main navigational
channel runs to the NNW.
The southern part of the sampled area lies in
the deepest portion of Moreton Bay, earlier called
the ‘basin’ by Stephenson, Williams and Lance
(1970). Depths are almost 30 m and here the tidal
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
189
current on the sea floor is likely to be negligible.
The substratum contains obvious silt.
The general topography of the northern exit
from Moreton Bay is suggestive of a basin c. 30
m deep which has been partially filled by sand.
Maxwell (1970) states that wind and stream
erosion as well as undercutting of the shore line
have been responsible for large quantities of sand
being carried westwards into Moreton Bay.
Presumably Moreton Island has supplied the sand
for the north-eastern corrugations.
While over a long period the Middle Banks area
has become partially filled with sand, there is
evidence that this has not been a continuous and
progressive process. Three sets of soundings of the
Middle Banks area have been made. The first
made in 1865 show a considerable area of the
Middle Banks shallower than 10 fm. The second
soundings were made in 1946 and show an
enlarged shallow area towards the south. A third
set of soundings made by the Queensland
Department of Harbours and Marine in relation
to the Airport Scheme in 1972 show a reduction
of the southern shallow area. This must be due
to dredging the navigational channel. However the
later soundings show that the portion of the East
Channel deeper than 10 fm has become more
extensive (sec Fig. 3), and it seems likely that sand
has been removed to the north. Throughout tidal
currents are strong and underwater observations
showed sand ripples c. 0-5 m high on the bottom
in the vicinity of site 12 and these conditions
probably apply to most of the northern sites. Here
the sediment surface will be partially mobile and
is very ‘clean’.
2. SEDIMENTS: These were collected using the
01 m2 Smith-Mcintyre grab, with hand selection
of an aliquot. (It was not appreciated at the times
of collection that hand selection causes error, see
Stephenson. Cook and Raphael 1977.) Samples
were obtained in September 1972, September
1973 and March 1974. Analyses were by wet
sieving as described in previous papers, using
sieves of the following apertures in mm: 2-00, 1-00,
0-50, 0-25, 0-125, 0 063. The retained grades are
described repectively as gravel = shell grit, very
coarse sand, coarse sand, medium sand, fine sand,
and very fine sand, with the material passing the
finest sieve described as mud.
The data comprise a three-dimensional matrix
of percentages of 7 sediments grades x 57 sites
X 3 times. This was classified by a hierarchical
agglomerative method (Bray-Curtis, group
average) differing from the methodology for biotic
data (see later) only in that sedimentary data were
not transformed (they are already standardised by
Fig 3: Depths of southern % of sampling area on
successive surveys; top 1865 survey, middle 1946,
bottom 1972. Depths: > 10 fm, stipple; 6 fm, thin
contin. line; 3 fm, dotted line; high water mark, thin
contin. line. Arrow, true north; scale line 1 km.
sample total). The method is identical with that
used by Stephenson, Cook and Raphael (1977).
Classification of sites by sediments is shown by
dendrogram on Fig. 4a, and of times by sediments
on Fig. 4b.
The sites-classificalion gives an initial dicho-
tomy at ca 70% dissimilarity and the main groups
were accepted at 10-25% dissimilarity. In contrast
the times-classification gives an initial dichotomy
at ca 6% dissimilarity and the groups finally
accepted were at ca 4% dissimilarity.
In the sites classification of sediments three
main groups were first accepted, and the
subgroups of two of these explored for topographic
coherence. Only one showed this and hence four
sediment site-groups were finally accepted, and on
Fig. 5 these groups are map-plotted. They form
a general sequence from site-group 1 in the
south-west, through site-group 2 as a transverse
southern band to site-group 4 in the north.
Site-group 3 consists of flanking sites mostly
190
MEMOIRS OF THE QUEENSLAND MUSEUM
forming a western wedge between site-groups 2
and 4.
Mean compositions of the sediments in the four
site-groups are given in Table I. In site-group
1 the modal grade of sediment is fine sand,
followed by mud; in site-group 2 it is fine sand
followed distantly by medium sand while in
site-group 3 it is again fine sand but more closely
followed by medium sand. The northern sediment
site-group (4) differs markedly from the
remainder in having medium sand as the modal
group.
The mean composition of sediments in the three
times arc given in Table I. This shows a
progressive change towards coarseness over the
period of investigation. It had been anticipated,
following analyses on sediments in Bramble Bay,
that the floods of January 1974 would have caused
increased mudiness. By the time post-flood Middle
Banks sediments could be collected there had been
a cyclone. Wave action due to the latter could
have been responsible for removal of mud and
increased coarseness. Against this there is a
distinct suggestion of progressive change.
TABLE 1: Mean Percentages of Grades of Sediment in Sediment Site-groups and
Sediment Times-groups.
Sediment grade
Sediment
1 2
Site-groups
3 4
1 (Sept 72)
Times
5 (Sept 73) 7
(March 74)
Mud
15-7
8-4
6-9
1-4
5-2
5-0
4-4
Very fine sand
4-4
1-9
1-7
0-1
I-O
1-2
1-0
Fine sand
76-0
78*7
59-8
13-4
40-6
36-9
33-8
Medium sand
3-6
12-9
31-7
83-6
52-0
56-1
59-6
Coarse sand
O-I
0-2
0-3
0-7
0-5
0-6
0-5
Very coarse sand
01
0-1
01
0-3
0-2
0-2
0-3
Gravel/shell grit
01
0-2
0-2
0-6
0-5
0-3
0-4
38,41
39,40
20,28
1, 2, 3, 4, 5, 6
42,43
46,48
33,34
7, 8, 9.10,11,12
44,45
49,50
56
13,14,15,16,17,18
47
51,52
15,20,21,22,23,24
53,54
25,26,27,29,30,31
1
55
1 1
1 1
32,35,36,37,57
1
2
3
4
Fig 4: Dendrograms showing classifications using sediment compositions as attributes: A of 57 sites, B of 3
times.
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
191
Fig 5: Topographic distribution of the four sediment
site-groups.
Further analyses were conducted, to determine
in which sediment site-group the chronological
changes were greatest. First, grades of sediments
were reduced to five by fusion of coarse sand, very
coarse sand and gravel. Next, in each sediment
site-group the variance due to the three times in
each sediment grade was obtained, and variances
were then summated over grades. Results for
sediment site-groups 1-4 were sequentially as
follows: 10-08, 8-58, 220-52 and 42-06. There is
greater variation in the two northern sediment
site-groups (3 and 4) than in the two southern
ones (1 and 2) with extremely high variance in
sediment site-group 3.
3. Hydrography: (a) Currents. No precise
data are available on the extent of penetration of
Pacific Ocean water into Moreton Bay and its
mixation with the waters of the Bay. General
observation suggests that surface water from
outside the Bay penetrates to well beyond the
sampled area on each tidal cycle.
Also no precise data are available on tidal
currents. It is known that they flood from and ebb
to the north and that surface currents are
generally in excess of 1 km/hr apart from periods
of slack water.
(b) Salinities. Data are available for Shark Spit
(27°]7'S., 1 53^22 "E.) from a survey made by the
CSIRO Division of Fisheries and Oceanography.
The data from the deepest samples (c. 29 m) taken
at Shark Spit have been reviewed by Stephenson,
Cook and Raphael (1977). From August 1972
to July 1973 the salinity averaged 34-5°/oo and
ranged from 33-l°/oo to 35-3°/oo. The variations
showed a marked inverse relationship to rainfall
in the catchment of the Brisbane River in the
previous month.
From October 1973 to September 1974 the
average salinity was 33 ^“/oo and the range of
32-0°/oo to 35-l'^/oo. There was a period of
relatively low salinity (c. 32-I®/oo) from February
1974 to August 1974 evidently due to thcieffects
of the excessive runoff from the flood of January
1974 being prolonged by the rainfall associated
with the cyclone of March 1974.
It is clear that the effects of the extreme
climatic disturbances on the salinities were
prolonged but relatively minor.
(c) Temperatures. Data equivalent to the above
on water temperatures are: August 1972 to July
1973, mean 22-25°C, range 17-2-26-6'^C; October
1973 to September 1974, mean 21-79'^C, range
I5-9~25*8°C. Comparison with data average over
the period 1931-50 at the Old Pile Light near the
mouth of the Brisbane River suggests that from
December 1972 until about December 1973
temperatures at Shark Spit were consistently
about half a degree warmer than average.
Biotic Data — Analyses
Involving all Species
Many species were present in so few samples
and in such small numbers that they are excluded
from later analyses. These rarer species are
important in aspects such as faunistics, species
richness and diversity, and these and similar topics
are dealt with in the present section.
Faunistic Composition of the Samples: A
total of 468 taxa were obtained originally, and this
reduced to 463 because of ‘oversplitting’. Fifteen
of the listed taxa are known to be polyspecific.
All but the rare species are listed in the Appendix,
together with the code numbers used throughout.
192
MEMOIRS OF THE QUEENSLAND MUSEUM
The number 463 compares with 420 taxa
(excluding amphipods) from grab sampling an
area of c. 3 km- near Peel Island (Stephenson,
Williams and Cook 1974), 182 from an area of
30 km- in Bramble Bay (Raphael 1974) and 98
from Serpentine Creek (Stephenson and Campbell
1977).
Of the 463 taxa 31-8% were polychaetes, 27-9%
crustaceans (10-6% decapods and 7-2% am-
phipods), 23-5% molluscs (10-8% gastropods and
12-5% bivalves) and 6-3% echinoderms. The
recording of 18 species of cumaceans and 14
species of fish was noteworthy. Amongst the latter
were small specimens of dubiously benthic
species, and several crustaceans (e.g. Lucifer)
were also doubtfully benthic. Unless clearly
non-benthic (e.g. some crustacean larvae, and
some planktonic molluscs) all taxa which were
caught are listed.
Distribution Patterns of Rarer Species:
Rarer species are here taken as those in which
only one or two individuals were recorded in the
survey. By taking recordings of each individual
and summating over times we can determine
whether the rare species were concentrated in any
patterns of sites, and conversely by summating
over sites we can search for patterns in times.
The number of rarer recordings in sites ranged
from zero to 10, and map plotting suggested a
pattern in the data somewhat comparable to that
obtained in the sediment site-groupings. Mean
numbers in sediment site-groups were: site-group
1 (South-west) 3-57, group 2 (southern transverse
band) 4 00, group 3 (flanking middle) 8*20 and
group 4 (northern) 2-35. Overall the northern area
has few rare recordings, the southern area an
intermediate number, and the middle flanking
portion (with the most unstable sediments) has the
largest number.
The number of rare recordings in times ranged
from 18 to 29 with September values (mean 28-5)
noticeably greater than the remainder.
Sites x Times Analyses: The total data
which were available for analysis formed a three
dimensional matrix of 468 taxa (5 for species) x
57 sites {q for quadrats) x 8 limes (t). By
summaling over species we derived q x Lmatricies
with dimensions of 57 x 8 and two such
summations were considered. The first involved
the number of species {sj per sample; this is a
simple measure of diversity and in Hulbert's
(1971) terms it is the species density appropriate
to the particular sampling parameters. The second
summation was the total number of individuals (n)
of ail species, and is the numerical equivalent of
biomass. Numerous sample measures other than
direct summations are available provided species
become anonymous, and we used the standardized
Shannon diversity (to log base 10) which we
designate d. These three measures s, n, and d were
previously used in work on Bramble Bay by
Stephenson, Cook and Raphael (1976) (albiet
with different symbolism) and Stephenson, Cook
and Raphael (1977) and because the sampling
procedures differed only in the type of grab
employed comparisons can be effected. It is
appreciated that with polyspecific and fractionat-
ed taxa, the value of s and d are somewhat
inaccurate.
We are interested firstly in means over all
samples. These are for s 3214, for n 170-23, and
for d 1-143. They compare with 12-2, 158-4, and
0-74 respectively for preflood conditions in
Bramble Bay (Stephenson, Raphael and Cook
1976). Clearly the Middle Banks grounds as a
whole carried a richer and more diverse biota than
did Bramble Bay in the preflood sampling.
Secondly we are interested in the variation of
s, n and d values between sites and between times.
This requires first averaging by rows and by
columns in each of the q x t tables, and secondly
assessing the variability in row means and in
column means. Variation was expressed as ranges
and variances and results are given in Table
2 .
These results show there is greater heterogenei-
ty in sites than in times, with the ratios of
variances being 4-72 for s, 2-94 for n and 8-67
for d. While a fraction of the differences may be
due to greater ‘random’ variation in the sites data
which are meaned over eight times compared with
TABLE 2; Ranges and Variances of Mean Values of 5. n and d in Sites and in
Times.
Sites
Range
Means
Variance
Times
Range
Means
Variance
s
20-10-57-13
82-30
24-67-38-30
17-42
n
67-75-392-88
8393-4
85-44-217-65
2854-6
d
0-978-1-394
0-00702
1104-1-186
0-00081
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
193
times data over 57 sites, the greater importance
of sites in the overall heterogeneity is believed to
be a real phenomenon. The Bramble Bay work
showed a most marked time effect due to the flood
of January 1974 (Stephenson, Cook and Raphael
1977) but this is not apparent in the present
data.
Table 2 shows reasonable constancy in
diversities (d) from place to place and from time
to time. This suggests that changes in numbers
of species and number of individuals ‘balance out’
in the diversity formulation as noted previously in
the postflood data from Bramble Bay and also in
work on the Serpentine Creek benthos by
Stephenson and Campbell (1977).
Significant positive correlations were obtained
between s. n and d for the different sites
Spearman rank correlation coefficients were:
between s and n +0-83, between s and d +0-68
and between n and d 4-0-33.
Data on time trends are given in Table 3.
Spearman rank correlation coefficients between
the three sets of values were: s and n +0-98 (very
highly significant), between s and d +0*24 and
between n and d +0-13 (neither significant). This
suggests that chronological changes in s and n
are again sufficiently parallel almost to cancel out
effects in diversity.
Table 3 shows high values of n in each of the
two Septembers, and this follows the pattern
obtained in an earlier study at Peel Island
(Stephenson, Williams and Cook 1974). The
results are also similar in that lowest values occur
in March of a non-flood year. Values of s follow
the same tendency, but no general tendency is
apparent in the diversity data.
Classificatory Techniques
General Techniques: The general approach is
classificatory, beginning with a three dimensional
matrix of s (species) x q (sites or quadrats) x
t (times). By summation over times we drive an
s X q matrix and by classification obtain
site-groups and their related species-groups;
TABLE 3: Ml A\ Valui s oi
similarly by summation over sites we derive an 5
X t matrix and hence times-groups and their
related species-groups. The classification of the
two derived matrices (viz. s xq and sx t) followed
the methods used by Stephenson, Raphael and
Cook (1976) and this involved: transformation of
data using iog,o (n+1), Bray-Curtis dissimilarity
measure, group average sorting. For entity (i.e.
site or time) classification data were not
standardised; for species classification the
transformed data were standardised by totals prior
to deriving inter-species dissimilarities.
Past experience has shown that the main
objection to this method lies in the species
classification where rarer species form isolated
and ‘dominating’ species-groups. It was hoped that
by operating in the first instance within narrow
ranges of species abundances that this objection
would be less serious (see later). In fact the
species groupings which were obtained throughout
still left much to be desired.
Following earlier work (Stephenson, Raphael
and Cook 1976; Stephenson and Campbell 1977;
Stephenson, Cook and Raphael 1977) we take the
view that species should be placed in the same
species-group if they characterise groups of
entities (sites or times) in a similar way. A
satisfactory entity classification then becomes a
necessary pre-requisite to a satisfactory species
classification. Because of occasional misclassifica-
tions, some re-allocation of entities may be
necessary, involving either the occasional and
obvious cases (e.g. Stephenson and Williams
1971) or by procedures such as REMUL (Lance
and Williams 1975). In the present cases there
were no obvious misclassifications and re-
allocations have not been made.
If the entities are sites then the next step is to
replace the species x sites matrix by the much
smaller species x site-group matrix — this contains
the mean recordings of the species. By inspection
we can immediately see which species may
positively characterise a given site-group by
occurring there in proportionally the largest
numbers.
n A\l) d A1 Dll I I RI N’T Tivii s
(Sept 72)
2
(Dec 72)
3
(March 73)
Times
4
(June 73)
5
(Sept 73)
6
(Dec 73)
7
(March 74)
8
(June 74)
s
32-88
28-23
24-67
32-65
38-30
35-32
33-54
32-53
n
185-21
128-54
85-44
171-05
271-65
191-61
162-25
165-09
d
1-115
1-144
M04
1-127
1-139
1-186
1-172
1-163
194
MEMOIRS OF THE QUEENSLAND MUSEUM
However the variability of recordings within the
site-groups may be such that differences between
means are misleading. To locate such cases, as in
work of the recent past, we use a ‘pseudo-F test’
and employ such terms as ‘conformity’ and
‘noticeably different’ instead of ‘significant’, using
the 0 05 probability appropriate to the F test as
a cut-off point.
For a given array of species recordings arranged
in site-groups there are many possibilities for
testing, for example each site-group set against all
the remainder, each pair of site-group sets against
the remainder etc. It has been found more
economical to perform pseudo-F testing by a
programmable hand calculator on those sets
suggested by perusal of the data, than to test all
by a computer-based programme.
Pscudo-F testing reveals species which do not
conform to the entity-groupings. When species
with a range of abundances are considered we may
expect non-conformers to include common species
which are evenly distributed amongst the
entity-groups and which must not be lost sight of.
We may also expect them to include rarer species
insufficiently concentrated in any entity group for
the results to be meaningful.
Data Reduction: In benthic work there are
typically many rare species which add little to the
ecological understanding of an area beyond the
contributions they make to analyses like those of
the previous section. There is a more pragmatic
reason for their exclusion: they greatly increase
computing costs and their inclusion may result in
the computer being unable to accept the total
data. In the present case, to achieve the limiting
matrix of 199 x 57 x 8 would mean excluding
species occurring 1 2 times or less, and it was felt
this might be too severe. Because there are
objections to most of the methods of data
reduction previously employed we here attempted
alternative approaches. The first consisted of
dealing with the species in successive lots of 50
and compared the classifications obtained. This
showed, inter al, that the sites were divisible into
two markedly dissimilar groups. We then
considered the two groups of sites separately —
by reducing the q dimension in the matrix the
critical s dimension becomes automaticallv
reduced.
Analyses on Lots of Fifty Species
The potential advantages of this method are
first that the ‘stopping rule’ as regards species is
indicated by the data themselves. It occurs at that
point in the successive analyses when either no
patterns or ‘nonsensical’ patterns emerge. This
involves only a minimum of subjectivity. Secondly
it shows whether the patterns revealed by the
abundant and less abundant species are similar,
and thirdly the results should be less dependent
on the data transformation which is employed.
This is because there is a narrower range of
abundances in a matrix; in practice this relative
independence increased as we moved to the less
abundant group of species. It was anticipated that
the main disadvantage of the method would be in
comparing the results of the different analyses but
in the event this was only a minor problem.
We deal first with ail j x ^ analyses, then all
s "x. t analyses.
Species x Sites Analyses: We first consider
the site classifications, then the conforming
species.
(a) Sites — first 50 species: The dendrogram of
the classification (Fig. 6A) shows remarkably
clear separation into two site-groups which consist
of .southern and northern sites. Apart from a single
site (29) the boundary follows the topography of
the area, and coincides with the 10 fm depth
contour on the ‘lip’ of the Middle Banks. Three
subordinate groupings are evident within the
southern subarea, and although these have
topographic coherence we do not discuss them
further at this stage.
(b) Sites — second 50 species: The dendrogram
(Fig. 6B) again shows clear separation into
northern and southern site-groups, although less
clearly than in the previous dendrogram. The two
main site-groupings are identical with the previous
ones. Three subordinate groupings are again
evident in the southern subarea but their
constituent sites are not identical with the previous
ones.
(c) Sites — third 50 species: The northern and
southern site-groups were identical with those
above.
(d) Sites — fourth to sixth lots of 50 species:
In the fourth lot the results were only broadly
similar to the above with one subgroup mostly of
topographically northern sites (7, 20, 23, 24, 29,
35) in the main southern group which also
included site 36. In the fifth lot excluding two
vacant sites (9, 27), the original groups were
revealed excepting that site 31 was in the southern
group. In the sixth lot of species the south and
north pattern was not evident. There were eight
isolated sites and three groups at a 98%
STEPFJENSON ET AL; MACROBENTHOS OF MIDDLEBANKS
195
1 ^
1
75
B
1
55 _
r r--^
35
1
29,35
38,41
28,33
1, 2, 3, 4, 5, 6
38,41
39,45
28,29
1, 2, 3, 4, 5, 6
50
42,43
34,39
7, 8, 9,10,11,12
42,43
48.51
33,34
7, 8, 9,10,11,12
44,45
40,46
13,14,15,16,17,18
44,47
52,53
35,40
13,14,15,16,17,18
47
48,49
19,20,21,22,23,24
54
46,49
19,20,21,22,23,24
51,52
25,26,27.30.31,32
50,55
25,26,27,30,31,32
53,54
36,37,57
56
36,37,57
55,56
1 1
1 1
1 1
1 1
1 1
1 1
1 1
1 1
1 a
1b
Ic
2
1a
lb
1c
2
Fi(i 6: Dendrograms of site
classifications using species
recordings as
attributes:
A first 50
speies, B second 50
species.
dissimilarity level with only one group showing
approximate topographic coherence. The stopping
point is clearly at or about the fifth lot of species
(species occurrence >6).
The distinctiveness of the northern and southern
subgroups decreased progressively from the first
to the fifth lots of species. This is indicated in
Table 4,
TABLE 4: Levbls of Dhndrogram Fusions for
Spi.c iis Lots with Dfci.ininc; Abundancfs
Species
lot
Approx level of
lowest fusions
Approx level of
highest dichotomy
First 50
10
60
Second 50
20
70
Third 50
30
85
Fourth 50
40
90
Fifth 50
50
100
(e) Conformity of species to main site-
groupings: For species in the first to third lots,
the main site-groups are identical. For species in
the fourth to sixth lots of 50 it seems more
appropriate to test conformity to the general
pattern than to the fragmenting remnants of
it.
In these cases the rationale of testing for
conformity is altered. We obtain groups from
elsewhere and test on the lines of a ‘construction
set’ of data and a ‘validation set’. Instead of
‘noticeable differences’ of species distributions we
might now approximate more closely to ‘sig-
nificant differences' excepting that the problems
of non-normal distribution with series of zero
recordings became more acute.
The sixth lot of 50 species takes in only some
of the species in which three specimens were
recorded and omits others so the present testing
was extended to all occurring three times. Below
three recordings there can be no conformity in the
sense we have used the term.
Results of conformity tests to the southern and
northern groups of sites covering all species except
the less abundant ones are given in the Appendix.
It will be noted that more species positively
characterise the southern site-group than the
northern one. With decreasing abundance of
species there are increasing proportions which fail
to conform. Thus for 6 and 7 recordings the
percentage not conforming is 29, for 5 recordings
36, for 4 it is 56 and for 3 it is 65.
(f) The main ‘communities’: If we view a
benthic community as an area and the species
which characterise it positively, then we have two
main communities. It is of interest that identical
areas show up in analyses on the first, second and
third sets of species and an almost identical one
in the fifth set. Clearly the abundant and less
abundant species follow the same broad area
pattern.
196
MEMOIRS OF THE QUEENSLAND MUSEUM
B
Fig. 7: Dendrograms of time classifications: A first 50 species, B third 50 species.
There are very many conforming species, 158
in the southern community and 80 in the northern
one, and to assist comprehension these data must
be compressed. If we list only the five commoner
species which conform to each of the areas we
arrive at a southern community of Prionospio,
tanaid 1, Solemya, Callianassa and
Poecilochaetus and a northern community of
Urohaustorius, Amphiura octacantha, Con-
cholestes, mysid 4 and amphipod 3. Four of the
northern species are small crustaceans which are
possibly mobile; mobility could be an advantage
in this generally turbulent area.
Conforming species are possibly different from
numerically dominant species — a conforming
and dominant species in the southern area may
well be a dominant species (but clearly not a
positively conforming one) in the northern area.
Purely on a dominance basis the southern area is
a community of Prionospio, tanaid 1, Solemya,
Poecilochaetus and Discobotellina while the
northern area is a Urohaustorius, Prionospio.
Amphiura octacantha. Concholestes and mysid 4
community.
Species x Times Analyses; Time classifica-
tions of the first and third lots of 50 species are
shown as dendrograms on Fig. 7, and these are
representative of the remainder. No two
classifications were identical and the only common
grouping was of times 7 and 8. The times were
grouped in a sequential or near-sequential order
throughout, and there were never close groupings
of the equivalent seasons of the two years.
By comparing Figs. 6 and 7 it is evident that
dissimilarities between times are much less than
between sites.
With the lack of constancy in time patterns,
tests of conformity of species to these patterns are
on a somewhat shifting basis. Taking three
times-groups in each case gave only 75 conforming
species compared with 238 conforming to two
site-groups. Further consideration of times-
groupings is deferred until the next section.
Biotic Data — Separate Analyses
OF Northern and Southern Areas
GENERAL: For reasons given earlier it was
decided to subdivide the study area into southern
and northern subareas and to analyse each
separately. Additional reasons are (a) by working
within these respective communities heterogeneity
is greatly reduced, (b) special attention can be
given to the northern area which would be most
immediately involved in human interference, (c)
it is possible that time changes differ in the two
areas.
The problem about subdivision is that three
different criteria can be used and each gives
slightly different results. Using sediments site 20
is included in the southern group, but its biotic
and topographic affinities are with the northern
group. Using topography site 29 is in the northern
group, but using biotic affinities it is in the
southern. Site 36 alters affinities between one
biotic classification and another. Eventually an
overlapping division was decided upon with sites
29 and 36 in both groups. The northern group then
consists of sites 1-27, 29-32, 36, 37 and 57 (34
in all) while the southern group consists of sites
28, 29, 33-36, 38-56 (25 in all).
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
197
A
35 _
30 .
B
38,41
29,35
39.45
28,33
29,36
15,16
42.43
50
48 51
34,40
17,20
44.47
1
1 !
52,53
54
I I
46,49
55,56
I 1
1 1
1
1
4
2
3
6
10
1, 2
5,13
21,22
26,31
3. 4
14,19
23,24
32
7, 8
25,27
9,10
30
11,12
37,57
12
J L
11
J L
Fig. 8: Dendrograms of site classifications in A southern and B northern sub-areas. (Site-group 12 is disregarded
in the main analysis.)
Fig. 9: Topographic distribution of the 11 main
site-groups. Site-groups 4 and 6 overlap and their
individual sites are shown by the symbols
indicated.
Species Considered: An arbitrary cut-off level
was adopted, and species occurring less than 10
times in the reduced data were excluded. Because
we now look for finer patterns and hence more
site-groups, a higher cut-off level than previously
was thought desirable. It was confirmed later as
being approximately correct. The southern
analyses involved 159 species and the northern
ones 115.
Species x Sites Analyses: (a) Sites classifica-
tions. The dendrograms of site-classifications are
given m Figure 8. We first consider the marginal
sites 29 and 36 common to the two analyses. Site
36 joins the southern area dendrogram at a much
higher dissimilarity level than for the northern
area and is henceforth considered as a northern
site. Site 29 is ambiguous in its affinities and is
treated as both a northern and southern site.
In Figure 8A four site-groups are readily
recognisable at a dissimilarity level of c. 30% and
these show topographical coherence. They are
map plotted in Figure 9 and numbered 1-4 in a
S-N direction. In Figure 8B at just over 30%
dissimilarity there are five groups of very
dissimilar sizes. The largest group divides at a
slightly lower level to give groups of 11 and 14
sites and of these the former gives a chained series
and is retained as a unit. The latter spans the area
of greatest interest where excavation may occur,
and divides into three topographically coherent
sub groups at about 25% level. These subgroups
were retained. In all eight northern groups were
tentatively accepted and are map plotted as groups
5-12 (see Fig. 9). Site-group 12 consists of the
single site 6 which is marginal to the area of
198
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 5: Species Conforming to the Four
Southern Site-groups.
Category
Species
Most in
site-group 1
25, 16, 19, 90. Ill, 134,
(-6)
Approx
coequal most
site-groups 1 and 2
14, 29, 98, 96, 118, 124, 170
(-7)
Most in
site-group 2
156, 176 ( = 2)
Approx
coequal most
site-groups 2 and 3
1, 9, 15, 37, 66, 75, 139, 146,
181, 212 (=10)
Most in
site-group 3
3, 4, 45, 65, 94. 108, 93, 172,
137, 173, 175, 219 (=12)
Approx
coequal most
site-groups 3 and 4
52, 48, 68, 63. 64, 32, 61, 53,
34, 20, 84, 83, 88, 103, 22, 85,
126, 135, 128, 151, 113, 131,
49, 143, 147, 161, 168 ( = 27)
Most in
site-group 4
51, 8, 2, 7, 23, 21, 81, 46, 18,
43, 106, 58, 56, 150, 129, 6, 44,
57, 70 (=19)
Most in site-groups
I, 2, 3 (least in 4)
24, 27, 33, 36, 38, 60, 95, 127,
159 ( = 9)
Most in site-groups
2, 3. 4 (least in 1)
26, 30, 39, 35, 97, 74, 107, 117,
77, 144, 141 (=11)
Species arc ranked from those with highest to those
with lowest recordings in each group.
TABLE 6: Species in the Seven Northern
Site-groups.
Category
Species
Approx
coequal most
site-groups 5 and 6
31, 3 ( = 2)
Most in
site-group 6
1, 10, 4, 52, 9, 30, 15, 63, 26,
13, 11, 40, 83, 37, 41, 91, 17
( = 17)
Most in site-groups
5, 6, 7
138 (=1)
Most in site-groups
6 and 7
5 (=1)
Most in
site-group 7
49, 199 ( = 2)
TABLE 6: cont
Category
Species
Most in site-groups
7 and 8
160 (=1)
Most in
site-group 8
54, 86, 104 ( = 3)
Most in site-groups
7, 8, 9
7, 57, 70, 91, 163, 93, 109
( = 7)
Most in site-groups
7, 8, 9, 11
20, 59 ( = 2)
Most in site-groups
6, 7. 8, 9
23, 77 ( = 2)
Most in
site-group 9
6, 8, 102, 89, 116, 203 (=6)
Most in
site-group 10
21, 47, 55, 155, 182 ( = 5)
Most in
site-group 1 1
43, 99, 136, 140 ( = 4)
Most in site-groups
10 and 1 1
2 (=1)
Most in site-groups
9, 10, 1 1
32 (=1)
Most in site-groups
9 and 1 1
105 (=1)
Most in site-groups
5, 6, 7, 8 (least
in 9, 10, 11)
39 (=1)
Most in site-groups
5, 6, 7, 8, 9 (least
in 10, 11)
34 (=1)
Most in site-groups
5, 6, 7, 8, 9, 10
(least in 11)
113 ( = 1)
Most in site-groups
6, 7, 8, 9, 10, II
(least in 5)
18 (=1)
Most in site-groups
5, 7, 8, 9, 10, 11
(least in 6)
22 (=1)
Species are ranked from those with highest to
those with lowest recordings in each group.
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
199
interest and is disregarded in subsequent
analyses.
(b) Species-groupings, southern sub-area. Of
the 159 species considered, 42 did not conform to
any rational arrangement of site-groups. Of the
six species occurring 11 and 10 times, only one
conformed indicating a correct choice of cut-off
point.
The conforming species are listed in Table 5
which shows that in general more species
characterise two site-groups than one. If we
include species characterising one, two or three,
site-groups, then 22 species characterise site-group
I, 39 site-group 2, 69 site-group 3 and 57
site-group 4. To describe communities in terms of
these numbers of characterising species is not
helpful. As stated earlier an alternative method
is not to consider conforming species but instead
a few of the numerically dominant species in each
site-group and this we follow.
(c) Species-groupings, northern sub-area. Of
the 115 species considered, 54 did not conform to
any rational arrangement of site-groups. Of the
four species occurring less than 12 times, only one
conformed, again confirming the choice of cut-off
level.
The conforming species are listed in Table 6,
which shows that site-group 6 has the largest
number of uniquely characterising species (17);
this agrees with the isolated position of site-group
6 on the dendrogram (Fig. 8B).
Many of the species conform to combinations
of site-groups, with the largest number (17)
involving site-groups 7 and 8 followed by
site-groups 7, 8 and 9 (16spp.). This confirms the
similarity of these site-groups, again as shown in
the dendrogram.
Counting species which characterise more than
one site-group, then 7 characterise site-group 5,
27 site-group 6, 21 site-group 7, 20 site-group 8,
23 site-group 9, 10 site-group 10 and 1 1 site-group
II.
(d) Overall view of site-groups. Table 7 shows
the site composition of each of the eleven main
site-groups considered, and their more abundant
characterising species. Data from the previous q
X / analyses are added to this table. They comprise
the mean number of individuals of all species per
site in a site-group per time (n), the mean number
of species (s) and mean standardised Shannon
diversity (to log base 10) {d}.
Site-group 1 occupies the extreme SW. of the
sampled area where sediments are predominantly
fine sand (76%) with the highest percentage of
mud (ca 16%), and in relatively deep water (ca
31 m). It may be described in terms of dominants
as a Schizaster — Nucula — Prionospio
community and within the southern area has the
lowest average population (197), lowest species
density (36), and lowest diversity (M2).
Site-group 2 forms an arc to the north of
site-group 1 in slightly shallower water (mean
depth c. 30 m). It was not clearly distinguished
by its sediments but these are predominantly of
fine sand (79%) followed by medium sand (10%).
It may be described as a Prionospio -
Aglaophamus — Callianassa community, has a
high average population (278) and for the
southern sub-area a relatively low species density
(38) and diversity (1T7).
Site-group 3 (depth c. 22 m) is parallel to and
to the north of site-group 2. It does not precisely
follow the topography of the area, lying at a slight
angle to the 20 m contour and is mostly deeper
than this. It is not clearly distinguished by its
sediments. These are predominantly fine sand
(69%) followed by medium sand (24%). It may
be described as a Prionospio — tanaid --
Solemya community. It has the highest recorded
population (320) and number of species (48), and
the average of the diversity in the total area
(1-20).
Site-group 4 (depth ca 16 m) consists of three
sites straddling and somewhat askew to the 20 m
contour just south of the southern lip’ of the
sampled area. Its sediments are highly variable,
site 50 having predominantly fine sand (78%), and
both sites 29 and 35 having predominantly
medium sand (76% and 66% respectively). It
could be described as a Rhizammina —
Discobotellina — Prionospio — Solemya
community but for the fact that the first named
species is markedly seasonal. The population
(215) is below the average for the southern
site-groups but much in excess of any northern
groups. The species density (46) is almost that of
site-group 3, and the diversity (1-29) is the highest
of the southern site-groups. Almost identically
high diversity also occurs in site-group 6 (depth
c. 9 m) which again contains site-group 29. It can
be described as a Prionospio — Urohaustorius
community, has the lowest population (79) of any
site-group, the lowest species density (24) but the
highest diversity (1-56). Site-group 5 is at the
western edge of the Middle Banks lip, with a
modal sediment of medium sand (82%). It is a
Prionospio — Urohauslorius — tanaid communi-
ty and if reckoned as within the northern sub-area
has its highest population (143), and species
density (38), and has a high diversity (1-30),
second only to site-group 6 (depth c. 1 1 m).
200
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 7: Drtah s OF Site-groups.
Site-group
I
2
3
4
5
Constituent
sites
38, 41, 42
43, 44, 47
39, 45, 48, 51,
52, 53, 54
28, 33, 34, 40,
46, 49, 55, 56
29, 35, 50
37, 57
Most
abundant
species
(occurrences*)
25(22), 12(22),
1(21), 10(18),
16(17), 19(14),
14(14), 11(11)
1(72), 11(17),
5(14), 9(14),
3(13), 4(12),
3(11)
1(77), 3(32),
4(30), 9(16),
13(13), 15(12)
28(27), 10(20),
1(16), 4(14),
3(10)
1(11),
2(9)
h*
197
278
320
215
79
}*
36
38
48
46
24
d*
1-12
M7
1-20
1-29
1-56
Main
sed. grades
76% fine sand
16% mud
79% fine sand
10% med. sand
69% fine sand
24% med. sand
Variable
87% med. sand
Mean depth
(m)
30-5
29-9
22-3
16-2
1 1-0
* Rounded
None of the remaining site-groups appear to be
distinguished by their sediments, ail having
predominantly medium sand present as in the two
previous cases.
Site-group 7 (depth c. 15 m) includes two sites
(31, 32) in or close to the northern end of the
dredged navigational channel. It can be described
as a Concholestes — Callianassa — Prionospio
community, and has almost the lowest population
of any site-group (81), and also low species
richness (25) and diversity (Ml).
Site-group 8 (depth c. 14 m) lies on the eastern
slopes of the Middle Banks in the area from which
sand might be dredged and may be described as
a Concholestes — Urohaustorius — mysid 4
community. It has a relatively low population
density (91) but somewhat higher species richness
(30) and diversity (M8) than adjacent
site-groups.
Site-group 9 lies in the NE. of the sampled area,
in generally deeper water than adjacent sites
(depth c. 19 m) and may be described as a
Concholestes — Amphiura octacantha — mysid
4 community. It has a higher population than
adjacent site-groups (125); species richness is low
(25) as is diversity (109).
Site-group 10 lies in the W. of the sampled area
in a generally shallow and turbulent area (depth
c. 8 m). It forms a Urohaustorius —
Platyischnopus — Prionospio community with
low population (113), species density (25) and
diversity (103).
Site-group 1 1 is the northernmost area sampled.
It lies in a turbulent area adjacent to but deeper
than that of site-group 10 (depth c. 14 m). It
forms a Urohaustorius — Amphiura octacantha
— Prionospio community, again with low
population (96), species richness (25) and
diversity (Ml).
SPECifs X Times Analyses: The dendrograms
of time classification for the southern sub-area
(Fig. 10A)and northern sub-area (Fig. lOB) are
alike in showing groupings which are predomin-
antly of sequential times and in not following
seasonal patterns. The sequential pattern is
slightly better developed in the northern sub-area
and also the inter-time dissimilarities are
generally lower.
By dealing with sub-areas the dissimilarity
levels in the times dendrograms have become
roughly equated to those of the sites dendrograms
and hence times now become relatively more
important. Perusal of the two-way coincidence
table of time-groups with species showed that less
than a third of the species conformed noticeably
to the time-groups of the dendrograms.
As an alternative to the dendrogram groupings
of times, these were dissected into chronologically
meaningful groupings and the conformity of
species to these groupings was investigated.
Approximately half the species analysed con-
formed to these groupings. Some of the remaining
species occurred at random (see later) and it was
concluded here there were no chronological
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
201
TABLE 7: Cont.
6
7
8
9
10 11
29, 36
26. 31, 32
21, 22, 23
24, 25, 27, 30
5, 13, 14,
19
15, 16, 17, 20 1, 2, 3, 4, 7,
8, 10, 11, 12, 18
1(24). 7(10), 5(10), 7(19), 2(8), 7(27), 6(21), 2(34), 21(12), 2(16), 6(14),
2(6), 1(8), 6(6), 8(8), 6(6), 8(14), 1(7), 1(10), 18(7), 1(10), 8(9)
3(5) 20(6), 1(6) 23(6) 8(5)
143
81
91
125
113
96
38
25
30
25
25
25
1-30
11 1
1-18
1-09
1-03
Ml
82% med. sand
—
> 80% med. sand
—
—
9-1
14-6
14-0
19-2
7-6
14-0
changes. For the residue while recordings were too
heterogeneous to be statistically at random, the
groupings of times to which the species conformed
did not make obvious chronological sense.
Finally the species were divided into four broad
categories, and each of these further subdivided.
Throughout the categories were non-overlapping
and sometimes this involved difficult decisions
regarding allocation. The categories were: A —
those occurring at random, and hence showing no
chronological pattern. For the commoner species
randomness was tested by using summed
recordings in each of the eight times. For species
occurring with fewer than 16 individuals, the limit
of formal use of is exceeded. We have still
applied the mechanics of the X “ test, which might
now be designated a ‘pseudo- X 2 test’ and describe
the species as ‘pseudo-random’; B — those with
Fig. 10: Dendrograms of time classifications within A
sequential patterns of various types. One sequence
of times was dissected from the remainder and the
pseudo F test applied to the Iog,o (« + 1)
transformations of the two sets of recordings. The
first dissection point separated times 1-6 from 7
and 8, and we thus determine the species with
noticeably decreased numbers after the flood and
those with noticeably increased numbers. The
second point was between times 1-4 and 5-8,
separating the two years of study, and we thus
derive species occuring in high numbers in the
second year and in the first year respectively. The
remaining sequential species followed a variety of
time combinations and we first separated those
with three sequential noticeably high recordings
('9 month species’) from those with two in
sequence ('6 month species’). Finally we listed
species with a sequence of two noticeably low
B
southern and B northern sub-areas.
202
MEMOIRS OF THE QUEENSLAND MUSEUM
recordings; C - those with seasonal patterns with
noticeably high recoi dings in the same season of
both years (mostly in September) and those with
noticeably low recordings, D — non-random
species giving apparently 'nonsense' chronological
patterns; these are presented in order of
decreasing likelihood that they give conceptual
sense. The first group contains species with a
single outstandingly low value, the second species
Tabu- 8; Conkormity or Specics to Prescribed
Time-groups.
Random
Random
South: 29, 53, 125, 137, 156 ( = 5 = 3%)
North: 15, 72, 73, 83, 87, 116, 130, 134, 136, 142
(=10=9%)
Pseudo-random
South: no, 171, 180, 190, 191, 192, 219, 224
(=8=5%)
North: 17, 38, 109, 117, 149, 169, 189 ( = 7 = 6%)
Sequential
Fewer after flood
South; 25. 30, 41 ( = 3 = 2%)
North: 22, 42, 110 ( = 3 = 3%)
More after Jlood
South: 4. 56, 58. 66, 105, 127, 146, 149, 161, 184
(=10=6%)
North: 31, 59, 86, 146, 203 ( = 5 = 4%)
More in 1st year
South: 96, 131 ( = 2=1%)
North: 89. 199 ( = 2=2%)
More in 2nd year
South: 9, li 19, 20, 23, 26, 38, 49, 55, 65, 75, 109,
ill. 133, 147, 212 ( = 6=12%)
North: 49, 66. 125, 162, 182 ( = 5=4%)
9-month species
Soulh; 6, 59, 83, 95, 128 ( = 5 = 3%)
North: 18, 23, 54, 55, 74 ( = 5=4%)
6-month species
South: 1, 2, 7, 8, 10, 13, 14, 16, 21, 24, 27, 33, 39,
44, 50, 64, 71, 74, 106, 107, 120, 154, 166,
197 (=24=15%)
North: 2, 6, 7, 1 1, 13, 24, 37, 67, 69, 70. 71, 82, 92
(=13 = 11%)
Sequential low recordings
South: 3, 31, 32, 48 (=4=3%)
North; 9, 20, 26. 32, 57 ( = 5=4%)
Seasonai.
Seasonal high recordings (mostly Septembers)
South; 12, 28. 45, 54, 61, 81, 97, 101, 108, 118, 126,
129, 135 (=13 = 8%)
North: 21, 63. 77, 121 (=4=3%)
Seasonal low recordings (various)
South, nil (=0=0%)
North: 5, 8, 56, 58 ( = 4 = 3%)
O BSC l IRE
Single outstanding low
South: 36, 37, 40, 69 (=4)
North; 20, 35, 39, 43, 44, 46, 50 ( = 7)
Oscillating
South; 11, 18, 35.. 42, 43. 46, 47, 52, 57, 60, 62, 67,
70, 76, 80, 84, 85, 88. 90, 91. 92, 93, 94, 98,
104, 113, 115, 117, 122, 132, 139, 141, 143,
152, 153, 168, 170, 172, 176, 204, 209, 216
( = 42)
North- 1, 4, 10, 12, 19, 40, 41, 45, 47, 48, 53, 62,
65. 75, 78, 79, 85, 91, 93, 99, 102, 104, 105,
112, 113, 114, 132, 140, 141, 147, 155. 158,
160, 163, 185, 187 ( = 36)
Single outstanding high
South: 5, 17, 22, 34, 51, 68, 77, 8?, 103„ 124, 138,
144, 145, 150, 151, 159, 173, 175, 178, 181,
186, 196, 211 ( = 23)
North: 3, 27, 43, 52, 100, 119, 138, 148, 164
(-9)
with oscillating values and typically with alternate
high and low recordings, while the last group
contains species with a single outstandingly high
value.
Details of the species in each of these categories
in southern and northern sub-areas are given in
Table 8. Summated percentages of random and
pseudorandom species are 8% S., 15% N.; of
sequential species 40% S., 33% N.; of seasonal
species 8% S., 7% N., and of obscure time patterns
43% S., and 45% N None of the differences
between the two sub-areas were significant.
The flood of 1974 produced noteworthy changes
in less than 10% of the species, in marked
contradistinction to Bramble Bay (Stephenson,
Cook and Raphael 1977) More species
occurred in noticeably high numbers after the
flood (15) than in noticeably low numbers. Apart
from flood-affected species, 21 species occurred in
higher numbers in the second year of study
compared with four in the first year.
DISCUSSION
Site PATTERNS Four groups of sites were
clearly recognisable in the southern sub-area and
seven groups were less clearly recognisible in the
northern sub-area. There is no accepted level at
which these site-groups can be designated as
communities and subcommunities.
Characteristics of each of these site-groups have
been given in the text (especially in Table 6) and
we now discuss all except the most northerly ones
(site-groups 10 and 11). The southern groupings
(site-group 1-4) are in deeper areas of fine sand
furthest from oceanic water and have relatively
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
203
high populations, species densities and diversities.
The individual groups within the southern area do
not precisely follow either the topography or
sedimentology of the area and one can only
assume that a combination of hydrographic
parameters is responsible for their delineation. A
likely combination would be distance from oceanic
water and current velocity. The highest popula-
tions and species densities occur on the relatively
steep slopes just south of the Middle Banks, and
further south populations and species densities are
reduced and other ‘communities’ can be
recognised. Within the confines of our study area
the areas of highest populations and species
densities do not form an ecotone between south
and north; there is no doubt the populous area
belongs to a southern site-group. One suspects on
the basis of a study of the dredged biota of
Moreton Bay by Stephenson, Williams and Lance
(1970), that our most south-western site-group 1
may extend considerably beyond our study area,
in which case the populous area of site-group 3
(and to a lesser extent site-group 2) become
something of especial local interest.
Taking site-groups 2 and 3 together, in terms
of population density we have an average of c.
1500 animals per square metre contrasting with
r. 1000 further south and c. 600 further north.
These are not high values within either a Moreton
Bay or a more global context. For example inshore
at Bramble Bay, Stephenson, Raphael and Cook
(1976) give preflood mean values of up to and
3000 organisms per square metre, and quote both
higher and lower figures from the literature. On
the other hand both species density (c. 43) and
diversity (r. 1-2) are much higher than were
obtained with a van Veen grab at Bramble Bay
(c. 12 and 0-8 respectively). The dense
macrobenthos in site-groups 2 and 3 corresponds
with an area in which prawn trawling is
concentrated. Presumably the factors responsible
for the dense benthos, plus the benthos itself,
produce a concentration of prawns. If so the
prawns and the trawling could both be expected
to take their loll of the benthos.
It is undesirable that future engineering works
should have any permanent deleterious effects on
the areas of site-groups 2 and 3. We can obtain
some insight on the likelihood of these occurring
by attempting an evaluation of the effects of
engineering works now in operation — viz.
dredging of the navigational channel (between the
beacons in Fig. 2). Sites 40, 50 and 56 lie
immediately south of this channel, and they do
not appear to show any biotic differences
compared with neighbouring sites. In fact two of
these sites (40 and 56) are in the area of greatest
population density (site-group 3).
Site-group 5 is of especial interest because the
two sites lie in (site 37) and in immediate
proximity with (site 57) the southern end of the
dredged channel. They are characterised by the
lowest population density (79) and species density
(24) obtained in the entire survey, and it is
difficult to believe other than that dredging has
been responsible. It is of interest to note that this
site-group has the highest standardi.sed Shannon
diversity (mean value 1*56), and this provides a
good example of how misleading diversity
measures can be.
Site-groups 4 and 6 lie respectively just south
of the topographic boundary between southern
and northern areas, and on or just north of the
sedimentary boundary. They are intermediate as
regards population density (ranging from c. 1050
to c. 750 individuals per square metre) but have
higher species densities and diversities than
northern sites. Site-group 4 is of especial interest
because two of its characterising species
{Rhizammina and Discobotellina) are arenaceous
Foramenifera, although one of these
iRhizammina) is markedly seasonal.
Discobotellina is widespread in Moreton Bay
(Stephenson, Williams and Lance 1970) and in
different areas is in association with a great
variety of other species. There is a reasonably
close parallel with dominance of other species of
areaceous Foramenifera off the W. African coast
(Buchanan 1960).
In site-group 7 one site (32) lies in the northern
end of the dredged navigational channel, one lies
very near this end (31) and the third (26) c. 1 ‘/2
km away. Populations densities and species
densities are approximately as low as those of
site-group 5, and again it seems that dredging is
responsible.
Site-groups 7, 8 and 9 show considerable biotic
similarity and form a series running northwards
from the dredged area on the western side of the
Eastern Channel. There is a progressive increase
in population density from 7 to 9, and it is possible
this is due to distance from the dredging.
Timh-PatternS: Analyses which involved
summations over all species indicated seasonally
high populations and species densities in each of
the Septembers and further that values were
higher in the second year than the first one.
Analyses on separate species gave obscure
sequential groupings of times rather than seasonal
groupings, and suggested a progressive shift in the
biota.
204
MEMOIRS OF THE QUEENSLAND MUSEUM
The latter analyses, while generally more
informative, run into a major problem in
interpretation which has been raised in an earlier
study (Stephenson, Raphael and Cook 1976).
Because of inaccurate relocation of sites on
successive samplings different local concentrations
of species are likely to be sampled on different
occasions. It could be argued that because the
values considered involve summated recordings at
many sites (25 and 34 in the main analyses)
‘random’ microtopographical variations should
cancel out. Against this species may be
concentrated in certain areas, and a single
exceptional patch within these areas could bias the
results. The importance of microtopographical
patterns has been demonstrated by Poiner (1977)
working north of Peel Island. In the present study
we can only estimate the relative importance of
chronology and microtopography. We assume that
when species conform to apparently meaningful
chronological changes these changes are real, and
that when the chronological picture is decreasingly
meaningful there is an increasing possibility of a
microtopographical effect.
Almost 10% of the species considered
conformed to seasonal patterns and in most cases
showed noticeably high populations in the two
Septembers. High September values have been
noted in two previous local studies — near Peel
Island (Stephenson, Williams and Cook 1974) and
at Bramble Bay before the 1974 flood (Stephen-
son, Raphael and Cook 1976) and can now be
regarded as an established feature in the benthic
calendar. Sequential patterns of one sort or
another occurred in about 40% of the species
analysed. Because there is a grow'ing interest in
non-seasonal changes in benthos (e.g. Stephenson,
Williams and Cook 1974; Buchanan, Kingston
and Sheader 1974; Eagle 1975; Stephenson,
Raphael and Cook 1976; and Stephenson, Cook
and Raphael 1977) and because this was the
largest category of species with recognisable
patterns, special attention was devoted to it.
The general approach we followed was to look
for changes in biota w'hich are paralleled by
abiotic changes and to assume in the first instance
that the former are due to the latter. Where no
such parallels occur we must fall back upon biotic
interactions either between benthic species
(Rhoads and Young 1970; Eagle 1975) or between
benthic organisms and nektonic predators
(Stephenson and Searles 1960; Levings 1972,
1974; Mills 1975). In the present case the most
severe abiotic changes for nearly a century were
probably the floods of January 1974 and resultant
prolonged dilution of the Middle Banks area
(Stephenson, Cook and Raphael 1977). Almost
10% of the species showed noticeable changes
between the prefiood times (times 1-6) and the
postflood ones (times 7-8). While this is more
than for any other sequential period, it is a
miniscule effect compared with that on Bramble
Bay (Stephenson, Cook and Raphael 1977).
Moreover at Middle Banks the Hoods had a
‘beneficial’ effect with 15 species occurring in
noticeably higher numbers after the Hood and only
six before. The effect was most noticeable in the
southern area, where ten species increased in
number after the Hood. The simplest explanation,
that this is due to a supply of sedimentary food
material, as MacGinitie (1939) suggested after a
Californian Hood, appears unlikely. There was less
small particulate sediment in the area after the
Hood than before.
The remaining sequentially occurring species
mostly involved noticeably high numbers for only
two consecutive sampling periods i.e. for 6 months
or slightly longer. It appears that a sequence of
species occupied (in quantity) the different areas
of sampling each for tolerably brief periods and
were then replaced by others. This is suggestive
of rapid recruitment and rapid replacement, as
Stephenson, Raphael and Cook (1976) suggested
at Bramble Bay. It implies that biotic interactions,
whose nature is as yet unknown, are an underlying
phenomenon. Superimposed on the transitory
aspects of the data there is an underlying trend
from lime 1 to time 8 (as shown by the
dendrograms) suggesting an overall biotic shift.
The somewhat scanty sedimentary data suggest a
parallel shift towards coarser sediments and it is
tempting to interrelate the two.
The largest single category of species revealed
by the times analyses (40-45% of the species
considered) fail to give immediate chronological
sense. There are two possible explanations,
microtopographical patchiness or real chron-
ological changes on a 3 monthly lime bases. For
example species with a single outstandingly low
value could reHect a single occasion of severe
predation followed by replacement during the next
three months; species with oscillating values could
reflect irregular and repeated recruitment; while
species with a single outstandingly high value
would be showing settlement in 3 months and
disappearance during the next three months.
Present data do not permit an easy choice between
the alternative microtopographic or rapid
chronological changes; current work suggests both
are about equally applicable on an 8 week time
basis.
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
205
Continual Biotic Flux: Wc have noted the
likelihood of tolerably rapid changes in at least
40% of the species — the sequential ones. We have
suggested that rapid changes may be a feature of
a greater proportion of the species.
If the bulk of the biota consisted of perennial
species we would not expect the biota to show
rapid changes, and what might appear as such
would be due to topographical patchiness. If the
bulk of the species were annuals, rapid changes
would be inevitable. Thus in theory we could
argue from life-spans to the reality of the
chronological changes. Unfortunately we have no
direct information upon the life-spans of species
from the area apart from some data on
Discobotellina (Stephenson and Rees 1965a, b)
which suggests that larger specimens may be three
years old. In general we can only assume that
relatively large size equates to an age exceeding
one year, and amongst the 100 most abundant
species the only cases of relatively large sized
species are: spp. 5, 10, 12, 14, 24, 25, 29, 39, 41,
42, 49, 69, 87, 95 and 98 (i.e. 15 in all). Amongst
the less abundant species the proportion of large
species is much lower, and that of annuals or
subannuais must be very high. As Eagle (1975)
has noted when populations consist of single year
classes we can expect instability. He relates this
to low diversity, but as indicated by Stephenson,
Williams and Cook (1974) it could go a long way
to explaining high diversity in a time context.
The overall impression was of a biota dominated
by small individuals, of which very few species
would attain sexual maturity within the study
area. This was particularly so in the northern
sub-area; of the 15 larger species listed only one
characterised the northern sub-area. If species
disappear from within the area before attaining
sexual maturity, predation by fish seems a likely
cause. Levings (1972, 1974) has shown how
seasonal variations in predation by a single species
of fish can produce complex changes in a
relatively simple benthic biota off the Atlantic
coast of Canada. In the present study area there
are probably some 20 species of benthic feeding
fish (personal communication from Professor J.
M. Thomson) and their predation could account
for the various and sometimes rapid quasi-
seasonal declines in population which were
observed.
Non-seasonal increases in population are more
difficult to explain. One possibility is that many
species have prolonged breeding seasons (which is
not unlikely in a subtropical situation) and can
occupy vacant space at any period within a
prolonged settlement period. A balance between
prolonged settlement and variable predation could
well explain the cases of oscillation noted above.
It is clear that much more data are required
before this suggestion can be confirmed.
Meanwhile the evidence that the benthic
population is in an overall state of flux seems
tolerably strong. It carries some important
consequences as regards a temporary human
disturbance of the area. Excavation in the
proposed area is unlikely to cause major
destruction of breeding populations because few
species remain there long enough under natural
circumstances to breed. Also, if the normal
method of maintenance of the biota is by
settlement of one of a great variety of species, it
seems probable that this process will only be
briefly interrupted, and that a biota roughly
comparable with the original should soon be
re-established. We might postulate that the
settling biota will be more simplified than
originally because a larger area and a more vacant
area than that provided after fish predation will
be available. After some predator-prey oscillations
possibly of a few months duration a situation very
comparable with the original could be
expected.
ACKNOWLEDGMENTS
We are deeply grateful to the (then)
Commonwealth Department of Housing and
Construction and to the Queensland Department
of the Co-ordinator General for sponsoring the
study and providing financial support. We are also
most grateful to (a) the CSIRO Division of
Fisheries and Oceanography for access to their
currently unpublished hydrographic data, (b) the
survey branch of the Queensland Department of
Harbours and Marine for access to their 1972
soundings of the area and for helpful advice on
chartings, and (c) to the Hydrographer of the
Royal Australian Navy for survey datings and
copies of older charts of the area.
Especial thanks are due to the numerous
individuals who have assisted in species identifica-
tions. In addition to those who named past
reference collections, acknowledged in previous
papers, these comprise: Dr A. N. Baker, National
Museum of New Zealand (ophiuroids and
echinoids); Prof. J. H. Day, lately of Zoology
Dept., University of Cape Town (cirratulid and
orbiniid polychaetes); Dr S. Edmonds, Zoology
Dept., University of Adelaide (sipunculids); Dr J.
Haig, University of S. California (pagurids); Dr
P. Hutchings, Australian Museum (nephtyid and
terebellid polychaetes); Mr M. G. Johnston,
Zoology Dept., University of Queensland
206
MEMOIRS OF THE QUEENSLAND MUSEUM
(cumaceans); Dr J. D. Kudenov, Dept, of
Fisheries and Wildlife, Victoria (aphroditid and
spionid polychaetes); Dr J. G. B. Poore, same
address, (Callianassa and anthurid isopods); Dr
C. R. Smalley, Zoology Dept., University of W.
Australia (alpheids); Prof. J. M. Thomson,
Zoology Dept., University of Queensland (fish);
and Miss V. A. Wadley, CSIRO Division of
Fisheries and Oceanography (peneids). Not all
material in these groups was identified by these
experts and the authors accept responsibility for
any errors which may have occurred.
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Schizammina two genera of arenaceous foramin-
ifera from the tropical Atlantic, with a description
of a new species. J. Linn. Soc. (Zool.) 44:
270-7-
Buchanan. j. B., Kingston. P. F. and Sheadkr. M.,
1974. Long-term population trends of the benthic
macrofauna in the offshore mud of the Northum-
berland Coast. J. mar. Biol. U.K. 54:
785-95.
Eagle, R. A., 1975. Natural fluctuations in a soft
bottom benthic community. J. mar. Biol. V.K.
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Hurlbert. S. H., 1971. The nonconcept of species
diversity: a critique and alternative parameters.
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Lance, G. N. and Williams W. T., \915.REMIJL: a
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Levings. C. D., 1972. ‘A Study of Temporal Change
in a Marine Benthic Community, with Particular
Reference to Predation by Pseudopleuronectes
americanus (Walbaum). (Pisces: Pleuronectidae).
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Scotia. pp.201.
1974. Seasonal changes in feeding and particle
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icanus). Trans Am. Fish Soc. 103: 828-32.
MacGinitie, G. E., 1939. Some effects of fresh water
on the fauna of a marine harbor. Amer. Midi. Nat.
21: 681-6.
Maxwell, W. G. H., 1970. The sedimentary framework
of Moreton Bay, Queensland. Aust. J. mar. Freshw.
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Mills. E. L. 1975. Benthic organisms and the structure
of marine ecosystems. J. Fish. Res. Bd. Canada.
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PoiNER. 1. R., 1977. Microvariation in the fauna of a
sublittoral sand bank, Moreton Bay, Queensland.
Aust. J. Ecol. 2: 297-308.
Raphael, Y. L, 1974. ‘The Macrobenthic Fauna of
Bramble Bay, Moreton Bay, Queensland." M.Sc.
thesis (Unpublished), Zoology Department, Univer-
sity of Queensland.
Rhoades. D. C. and Young. D. K., 1970. The influence
of deposit feeding organisms on sediment stability
and community trophic structure. J. mar. Res. 28:
150-78.
Stephenson. W., and Campbell, B. M., 1977. The
macrobenthos of Serpentine Creek. Mem. Qd Mus
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Stephenson. W., Cook. S. D. and Raphael, Y. L, 1977.
The effect of a major flood on the macrobenthos
of Bramble Bay, Queensland. Mem. Qd Mus. 18(1):
95-118.
Stephenson. W., Raphael. Y. 1. and Cook.S. D., 1976.
The macrobenthos of Bramble Bay, Queensland.
Mem. Qd Mus 17(3): 425-47.
Stephenson. W. and Rees. M., 1965a. Ecological and
life history studies on a large foraminiferan
(Discobotellina biperforata Collins, 1958), from
Moreton Bay, Queensland. I. Life cycle and nature
of the test. Pap. Dep. Zool. Univ. Qd 2:
207-23.
1965b. Ecological and life history studies on a large
foraminiferan (Discobotellina biperforata Collins,
1958), from Moreton Bay. Queensland. 11.
Aquarium observations. Pap. Dep. Zool. Univ. Qd
2: 239-58.
Stephenson. W. and Searles, R. B., 1960. Experimen-
tal studies of the cology of intertidal environments
at Heron Island. L Exclusion of fish from beach
rock. Aust. J. mar. Freshw. Res. 11: 241-67.
Stephenson. W. and Williams, W. T., 1971. A study
of the benthos of soft bottoms, Sek Harbour, New
Guinea, using numerical analysis. Aust. J. mar.
Freshw. Res. 22: 11-34.
Stephenson. W., Williams, W. T. and Cook. S. D.,
1974. The benthic fauna of soft bottoms. Southern
Moreton Bay. Mem. Qd Mus. 17: 73-123.
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1970, The macrobenthos of Moreton Bay. Ecol.
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STEPHENSON ET AL; MACROBENTHOS OF MIDDLEBANKS
207
APPENDIX
Species from Middle Banks benthos with indications of their systematic positions. Code numbers are in hierarchical
order of abundance.
Code
No.
Species
Systematic
position
Number Main
Collected site group
1
Prionospio sp. 1
Polychaeta: Spionidae
12802
S
2
Vrohaustorius sp. — polyspecific
Amphipoda: Haustoriidae
3394
N
3
tanaid 1
Tanaidacea
3094
S
4
Solemya terraereginae (1 redale)
Bivalvia: Solemyidae
3075
S
5
Callianassa arenosa Poore
Crustacea: Callianassidae
2776
S
6
Amphiura octacantha H. L. Clark
Ophiuroidea: Amphiuridae
2741
N
7
Concholesies sp. polyspecific
Amphipoda: Corophiidae
2674
N
8
mysid 4
Mysidacca
2390
N
9
Poecilochaetus sp.
Polychaeta: Trochochaetidae
2258
S
10
Discobotellina biperforata Collins
Foraminifera: Aslrorhizidae
2131
s
11
Aglaophamiis verrilli McIntosh
Polychaeta: Nephtyidae
2040
s
12
Nucula spp. — polyspecific
Bivalvia: Nuculidae
1922
s
13
Prionospio sp.4
Polychaeta: Spionidae
1837
s
14
Brissopsis luzonica Gray
Echinoidea: Spalangidae
1343
s
15
Magelona sp.2 (‘cincta’)
Polychaeta: Magelonidae
1341
s
16
aloidid
Bivalvia: Aloididae
1274
s
17
Magelona sp.3 (‘capensis’)
Polychaeta; Magelonidae
1217
s
18
amphipod 3
Amphipoda
993
N
19
Golfingia trichocephala (Sluiter)
Sipuncula: Golfingiidae
927
s
20
Eunice cf. indica Kinberg
Polychaeta: Eunicidae
886
N
21
Platyischnopus sp. — polyspccific
Amphipoda: Haustoriidae
852
N
22
gastropod (red and white)
Gastropoda
819
N
23
amphipod 4
Amphipoda
792
N
24
bivalve 1
Bivalvia
775
S
25
Schizaster lacunosus (Linnaeus)
Echinoidea: Hemiasteridae
703
S
26
Goniada emerita Audouin and Milne Edwards
Polychaeta: Glyceridae
688
s
27
Leptomya pura Angas
Bivalvia: Semelidae
673
s
28
Rhiiammina cf. algaeformis H. B. Brady
Foraminifera: Aslrorhizidae
668
N
29
Xenophrhalmoides dolichophallus Tesch
Decapoda; Goneplacidae
654
s
30
carid 2
Decapoda: Caridae
648
s
31
Dispio sp.2
Polychaeta: Spionidae
643
N
32
juv. fibularid
Echinoidea: Fibularidae
632
N
33
‘Dentaliuni’ sp.
Scaphopoda
564
S
34
Tharvx sp.A (undescr.)
Polychaeta: Cirratulidae
466
N
35
amphipod 6
Amphipoda
433
S
36
Paralacydonia paradoxa Fauvel
Polychaeta: Lacydoniidae
432
s
37
Terebellides sfroemi Sars
Polychaeta: Terebellidae
428
s
38
Haploscoloplos hifurcatus Hartman
Polychaeta: Orbiniidae
423
s
39
Branchiostoma nioretonensis Kelly
Cephalochordata
412
s
40
Sthenelais sp.
Polychaeta: Aphroditidae
409
s
41
Echinocardium cordatum (Pennant)
Echinoidea: Spatangidae
405
s
42
Neosolen vaginoides (Lamarck)
Bivalvia: Culiellidae
403
N
43
Mesanthura sp.l
Isopoda: Anlhuridae
396
N
44
amphipod 5
Amphipoda
380
N
45
orbiniid spp. — polyspccific
Polychaeta: Orbiniidae
360
s
46
Caulleriella sp.A + Tharyx sp.B — polyspccific
(both undcscr.)
Polychaeta: Cirratulidae
349
N
47
Lucifer hanseni Nobili
Decapoda: Sergestidae
344
N
48
lumbrinerid 1
Polychaeta: Lumbrineridae
322
N
49
Epigonichthys cultellus Peters
Cephalochordata
298
N
50
Glycera prashadi Fauvel
Polychaeta: Glyceridae
293
N
5!
Ophiura kinbergi Ljungman
Ophiuroidea: Ophiolepidae
290
S
52
Amphipholis loripes Koehler
Ophiuroidea: Amphiuridae
269
s
208
MEMOIRS OF THE QUEENSLAND MUSEUM
Code
No. Species
53 nemertean (pink) — polyspecific
54 Micronephlhys sphaerocirrata Wesenberg-Lund
55 nemertean (pale) — polyspecific
56 Marphysa sp.
57 Mesanthura sp.2
58 Magelona cf. papillicornis Muller
59 polycirrinae — polyspecific
60 bivalve 2
61 Amphioplus depressus (Ljungman)
62 anthurid (undescr. genus 2, sp.2)
63 amphipod 15 — polyspecific
64 bivalve 8
65 ?Notomastus cf. aberans Day
66 = species 34 (Tharyx sp.A)
67 ?Noiomastus latericeus Sars
68 polyzoan 1
69 Sipunculiis aequabilis Sluiter
70 terebellid (undescr. genus)
71 amphipod 14
72 anthurid (undescr. genus 2, sp. 1)
73 Thalenessa sp.
74 Poly dor a sp. 1
75 Owenia fusiformis Delle Chiaje
76 Sthenelepis cf. japonica (McIntosh)
77 Aricidea sp.
78 bivalve 4
79 bivalve 1 2
80 bivalve 14
8 1 bivalve 6
82 Chaetozone sp.A (undescr.)
83 Scalibregma injlatum Rathke
84 amphipod 8
85 nemertean (orange banded) — ?polyspecific
86 Eunice sp.2
87 Polycarpa tinefor (Quoy and Gaimard)
88 Chaetozone sp.B (undescr.)
89 Paraonides sp.
90 Paphia sp.
91 phoronid — ?polyspecific
92 amphipod 9
93 Cyclaspis iribulis (Hale) complex
94 Euclymene sp.
95 Pectinaria antipoda Schmarda
96 Alpheus distinguendus de Man
97 Naticu cf. colliei Recluz
98 Raphidopus ciliaius Stimpson
99 Lumbrineris sp.l
100 gastropod 10
101 Eocuma agrion (Zimmer)
102 bivalve 7
103 gastropod 5
104 Loimia medusa (Savigny)
105 Apseudes sp.
106 Aspidosiphon inquilinus Sluiter
107 Fulvia sp.
108 amphipod 20
109 nereid 1
110 isopod 5 — polyspecific
Systematic Number Main
position Collected site group
Nemertea
264
S
Polychaeta: Nephtyidae
262
Nemertea
256
Polychaeta: Eunicidae
227
N
Isopoda: Anthuridae
219
N
Polychaeta: Magelonidae
212
N
Polychaeta: Terebellidae
208
S
Bivalvia
188
s
Ophiuroidea: Amphiuridae
184
s
Isopoda: Anthuridae
180
N
Amphipoda
179
s
Bivalvia
171
s
Polychaeta: Capitellidae
168
s
Polychaeta: Cirratulidae
165
s
Polychaeta: Capitellidae
162
s
Polyzoa
161
s
Sipuncula: Sipunculidae
157
s
Polychaeta: Terebellidae
151
N
Amphipoda
137
s
Isolda: Anthuridae
134
N
Polychaeta: Aphroditidae
133
N
Polychaeta: Spionidae
129
Polychaeta: Oweniidae
117
S
Polychaeta: Aphroditidae
109
s
Polychaeta: Paraonidae
109
s
Bivalvia
106
N
Bivalvia
102
N
Bivalvia
95
S
Bivalvia
94
s
Polychaeta: Cirratulidae
93
N
Polychaeta: Scalibregmidae
90
s
Amphipoda
90
s
Nemertea
85
s
Polychaeta: Eunicidae
84
N
Tunicata: Styelidae
84
s
Polychaeta: Cirratulidae
82
s
Polychaeta: Paraonidae
80
N
Bivalvia: Veneridae
80
s
Phoronidae
80
s
Amphipoda
78
Cumacea: Bodotriidae
78
s
Polychaeta: Maldanidae
75
s
Polychaeta: Pectinariidae
75
s
Decapoda: Alpheidae
75
s
Gastropoda: Naticidae
74
s
Decapoda: Porcellanidae
73
s
Polychaeta: Eunicidae
71
N
Mollusca: Gastropoda
70
Cumacea; Bodotriidae
69
s
Bivalvia
69
N
Gastropoda
68
s
Polychaeta: Terebellidae
66
s
Tanaidacea
66
Sipuncula: Aspidosiphonidae
66
s
Bivalvia: Cardiidae
65
s
Amphipoda
64
s
Polychaeta: Nereidae
63
s
Crustacea: Isopoda
63
N
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
209
Code
No. Species
1 1 1 bivalve 29
1 1 2 Scoloplos johnstonei Day
113 Leptanthura cf. australis Haswell
114 Matuta inermis Miers
115 amphipod 28
1 16 Thermiste sp.
117 Conuber conica (Lamarck)
1 18 Tellina sp.l
119 Diogenes cf. rectimanus Miers
120 Natica sp.l
121 bivalve 16
122 amphipod 13
1 23 gastropod 8
1 24 Protankyra sp.
1 25 Sigalion sp.
126 Dimorphostylis australis (Foxon)
127 aphrodilid 3
1 28 Harmothoe sp.
129 mysid I
130 Drilonereis sp.
131 capitellid spp. — polyspecific
132 carid 4
133 bivalve 22
134 Lumbrineris sp.2
135 amphipod 18
1 36 Syllidia sp.
137 Nereis jacksoni Kinberg
138 amphipod 17
139 Chloeia flava (Pallas)
140 onuphid 1
141 carid 3
142 Veletuceta hedleyi (Lamy)
143 amphipod 21
144 gastropod 3
145 bivalve 3
146 phyllodocid 3
147 sabellid 2
1 48 isopod 3
149 amphipod 1 1
150 amphipod 16
151 gastropod 2
152 Leonnates stephensoni Ruillier
153 Muraenicthys godeffroyi Regan
1 54 isopod 2
155 bivalve 5
156 Tellina texturata Sowerby-
157 Theora lata Hinds
158 bivalve 17
1 59 Spiochaetopterus sp.
160 Pomacuma cognata (Hale)
161 nereid 2
162 cumacean 6
163 Austrolepidoda schmitti Efford and Haig
164 platyhelminth 1
165 amphipod 10
166 isopod 1
167 anemone 1
168 bivalve 9
Systematic Number Main
position Collected site group
Bivalvia
63
S
Polychaeta: Orbiniidae
57
N
Isopoda: Anthuridae
56
Crustacea; Calappidae
56
N
Amphipoda
51
S
Sipuncula: Golfingiidae
51
N
Gastropoda: Naticidae
50
S
Bivalvia: Tellinidae
50
S
Decapoda: Paguridae
49
N
Gastropoda: Naticidae
49
S
Bivalvia
49
N
Amphipoda
48
S
Gastropoda
47
Holothuriidae: Synaptidae
47
s
Polychaeta: Aphroditidae
46
Cumacea
46
s
Polychaeta: Aphroditidae
44
s
Polychaeta: Aphroditidae
44
s
Mysidacea
44
s
Polychaeta: Eunicidae
43
N
Polychaeta: CapitelHdae
40
s
Decapoda: Caridae
40
Bivalvia
40
s
Polychaeta: Eunicidae
39
N
Amphipoda
39
s
Polychaeta: Hesionidae
36
N
Polychaeta: Nereidae
36
S
Amphipoda
36
Polychaeta: Amphinomidae
35
s
Polychaeta: Eunicidae
35
N
Decapoda: Caridae
35
Bivalvia: Glycymeridae
35
N
Amphipoda
34
S
Gastropoda
34
s
Bivalvia
34
s
Polychaeta: Phyllodocidae
33
Polychaeta: Sabellidae
33
s
Isopoda
33
N
Amphipoda
33
Amphipoda
33
s
Gastropoda
33
s
Polychaeta; Nereidae
32
s
Pisces: Myridae
32
s
Isopoda
31
s
Bivalvia
31
N
Bivalvia: Tellinidae
29
s
Bivalvia: Semelidae
29
Bivalvia
29
N
Polychaeta: Chaetopteridae
28
S
Cumacea: Bodotriidae
26
N
Polychaeta: Nereidae
25
S
Cumacea
25
N
Decapoda: Albuneidae
25
N
Platyhelminthes: Polycladida
24
N
Amphipoda
24
Isopoda
23
S
Actiniaria
22
Bivalvia
22
210
MEMOIRS OF THE QUEENSLAND MUSEUM
Code
No. Species
169 amphipod 24
170 bivalve 11
171 Spiochaetopterus cf. vitrarius (Ehlers)
172 Glyphocurna sp.
173 Penaeus plehejus (Hesse)
174 Polyonyx iransversus (Haswcll)
175 phyllodocid 7
176 Sternaspis scutaia (Renier)
177 Metapenaeus hennettae Racek and Dali
178 gastropod 16
179 phyllodocid 1
180 Glyphocurna halei Greenwood and Johnston
181 Chaetopterus variopedatus Renier
182 Moira lelhe Lutken
183 capitellid 1
184 tanaid 6
185 Trachypenaeus fulvus Dali
186 Philine angasi Crosse and Fisher
187 gastropod 27
188 Cyclaspis mawsonae (Hale)
1 89 isopod 4
190 Dosinia cf. sculpta Hanley
191 bivalve 10
192 Ophiocentrus sp.
193 phyllodocid 2
194 amphipod 26
195 juv. carid 1
196 bivalve 13
197 bivalve 18
198 gastropod 20
199 gastropod 21
200 Ophiodromus sp.
201 Dorvillea sp.
202 Travisia cf. forbesii Johnston
203 maldanid 1
204 maldanid 2
205 tanaid 5
206 Apanthura sp.
207 amphipod 29
208 Hexapus granulijerus Campbell and Stephenson
209 Modiolus sp.
210 bivalve 21
21 1 bivalve 30
212 Natica sp. 4
213 gastropod 25
214 Pista spp. — polyspecific
215 Laonice sp.
216 Merinides sp.
217 Cyclaspis ? crelala (Hale)
? granulos (Hale)
218 anlhurid (undescr. genus 1 sp.l)
219 Doxander viiiaius Linnaeus
220 nemertean (green)
221 aphroditid 1
222 Diplocirrus sp.
223 ^Pontophilus sp.l (doubtfully benthic)
224 Phos sculptilis Watson
225 phyllodocid 6
Systematic
Number
Main
position
Collected site group
Amphipoda
21
Bivalvia
21
S
Polychaeta: Chaetopteridae
20
S
Cumacea
20
s
Decapoda: Peneidae
20
s
Decapoda: Porcelianidae
20
Polychaeta: Phyllodocidac
19
Polychaeta: Slernaspidae
19
s
Decapoda: Peneidae
19
Gastropoda
19
s
Polychaeta: Phyllodocidae
17
Cumacca: Bodotriidae
17
s
Polychaeta: Chaetopteridae
16
s
Echinoidea: Hemiasteridae
16
N
Polychaeta: Capitcllidae
15
Tanaidacca
15
s
Decapoda: Peneidae
15
Gastropoda: Philinidae
15
s
Gastropoda
15
N
Cumacea: Bodotriidae
14
Isopoda
14
N
Bivalvia: Veneridae
14
S
Bivalvia
14
s
Ophiuroidea: Amphiuridae
14
Phyllodocidac
13
Amphipoda
13
Decapoda: Caridae
13
Bivalvia
13
s
Bivalvia
13
s
Gastropoda
13
N
Gastropoda
13
N
Polychaeta: Hesionidae
12
Polychaeta: Runicidae
12
Polychaeta: Opheliidae
12
Polychaeta: Maldaiiidae
12
N
Polychaeta: Maldanidae
12
S
Tanaidacea
12
s
Isopoda: Anthuridae
12
N
Amphipoda
12
N
Decapoda: Goneplacidae
12
S
Bivalvia: Mylilidae
12
s
Bivalvia
12
Bivalvia
12
s
Gastropoda; Naticidae
12
s
Gastropoda
12
Polychaeta: Tercbellidae
11
s
Polychaeta: Spionidae
11
s
Polychaeta: Spionidae
11
s
Cumacea: Bodotriidae
1 1
N
Isopoda; Anthuridae
1 1
Gastropoda: Strombidae
1 1
s
Nemertca
10
Polychaeta: Aphroditidae
10
s
Polychaeta: Flabelligeridae
10
s
Crustacea: Crangonidae
9
N
Gastropoda; Photidae
10
s
Polychaeta: Phyllodocidae
9
N
STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS
211
Code
No. Species
226 = species 216 (Nerinides sp.)
227 Matuta spp. juv. not M. inermis} — polyspecific
228 bivalve 1 5
229 bivalve 24
230 Sthenolepis sp.
231 Pseudeurythoe microcephala (Fauvel)
232 nereid 4
233 eunicid 1
234 Travisia sp.2
235 bivalve 19
236 Luidia sp.
237 Rhynobrissus pyramidalis Agassiz
238 Glossobalanus sp.
239 polyzoa 2
240 nemertean (brown)
241 Eupanthalis sp.
242 Etenone sp.
243 Nephtys vikingensis Paxton
244 spionid 7 — polyspecific
245 Ophelina cf. acuminata Oersted
246 tanaid 3
247 amphipod 22
248 amphipod 27
249 Betaeus spp. — polyspecific
250 oyster 1
251 Amphiura catephes H. L. Clark
252 Nematonereis sp.
253 Polydora sp.2
254 bivalve 20
255 blenny I (doubtfully benthic)
256 phyllodocid 15
257 Arabella sp.
258 maldanid 3
259 sabellid 3
260 amphipod 19
261 Eucrassatella cumingii (Adams)
262 Mactra sp.
263 Tellina sp.2
264 Natica sp.3
265 gastropod 33
266 Mogula mollis Herdmann
267 Euleanira sp.
268 phyllodocid 5
269 phyllodocid 8
270 phyllodocid 10
271 nereid 3
272 Prionospio sp.3
273 amphipod 25
274 Alpheus spp. (larva) — polyspecific
275 Albunea cf. microps Miers
276 Limaria sp.
277 bivalve 25
278 bivalve 26
279 bivalve 31
280 gastropod 13
281 gastropod 15
282 Pentaceraster australis (Luther)
283 enteropneust
Systematic
position
Number Main
Collected site group
Polychaeta: Spionidae
9
S
Decapoda: Calappidae
9
N
Bivalvia
9
N
Bivalvia
9
S
Polychaeta: Aphroditidae
8
S
Polychaeta: Amphinomidae
8
S
Polychaeta: Nereidae
8
S
Polychaeta: Eunicidae
8
N
Polychaeta: Opheliidae
8
N
Bivalvia
8
N
Asleroidea: Luididae
8
S
Echinoidea: Spatangidae
8
N
Balanogiossidae
8
S
Bryozoa
7
Nemertea
7
s
Polychaeta: Aphroditidae
7
s
Polychaeta: Phyllodocidae
7
Polychaeta: Nephtyidae
7
N
Polychaeta: Spionidae
7
s
Polychaeta: Opheliidae
7
Tanaidacea
7
s
Amphipoda
7
Amphipoda
7
s
Decapoda: Alpheidae
7
s
Bivalvia: Ostreidae
7
Ophiuroidea: Amphiuridae
7
S
Polychaeta: Eunicidae
6
s
Polychaeta: Spionidae
6
S
Bivalvia
6
s
Pisces: Blenniidae
7
N
Polychaeta: Phyllodocidae
5
N
Polychaeta: Eunicidae
5
Polychaeta: Maldanidae
5
s
Polychaeta: Sabellidae
5
Amphipoda
5
S
Bivalvia: Eucrassatellidae
5
N
Bivalvia: Mactridae
5
N
Bivalvia: Tellinidae
5
N
Gastropoda: Naticidae
5
S
Gastropoda
5
Tunicata: Molgulidae
5
Polychaeta: Aphroditidae
4
s
Polychaeta: Phyllodocidae
4
Polychaeta: Phyllodocidae
4
s
Polychaeta: Phyllodocidae
4
s
Polychaeta: Nereidae
4
Polychaeta: Spionidae
4
Amphipoda
4
Decapoda: Alpheidae
4
s
Decapoda: Albuneidae
4
Bivalvia: Limidae
4
s
Bivalvia
4
s
Bivalvia
4
s
Bivalvia
4
Gastropoda
4
Gastropoda
4
Asteroidea: Oreasteridae
4
Enteropneusta
4
212
MEMOIRS OF THE QUEENSLAND MUSEUM
Code
No. Species
Systematic
position
Number Main
Collected site group
284 Paraplagusia unicolor (Macleay)
Pisces: Cynoglossidae
4
S
285 Leanira yhleni Malmgren
Polychaeta: Aphroditidae
3
s
286 aphroditid 7
Polychaeta: Aphroditidae
3
287 phyllodocid 4
Polychaeta: Phyllodocidae
3
288 phyllodocid 12
Polychaeta: Phyllodocidae
3
289 phyllodocid 14
Polychaeta: Phyllodocidae
3
290 phyllodocid 19
Polychaeta: Phyllodocidae
3
s
291 syllid 1
Polychaeta: Syllidac
3
292 spionid 6 — polyspecific
Polychaeta: Spionidae
3
N
293 = species 216 (Nerinides sp.)
Polychaeta: Spionidae
3
294 Ophelina gigaritea Rullier
Polychaeta: Opheliidae
3
295 capiteilid 2
Polychaeta: Capitellidae
3
296 sabcllid 6
Polychaeta: Sabellidae
3
s
297 unknown polychaete
Polychaeta
3
298 Gynodiastylis sp.
Cumacea: Diastylidae
3
299 tanaid 7
Tanaidacea
3
s
300 mysid 2
Mysidacea
3
301 mysid 3
Mysidacea
3
302 Squilla laevis Hess
Crustacea: Stomatopoda
3
s
303 Lysiosquilla perpasta (Hale)
Crustacea: Stomatopoda
3
304 ?Pontophilus sp.3
Decapoda: Crangonidae
3
N
305 Conchocetes artificiosus (Fabricius)
Decapoda: Dromiidae
3
306 Dorippe australiensis Miers
Decapoda: Dorippidae
3
307 Actumnus squamosus de Haan
Decapoda: Xanthidae
3
s
308 Achaeus lacertosus Stimpson
Decapoda: Majidae
3
s
309 Sanguinolaria sp.
Bivalvia: Sanguinolariidae
3
310 gastropod 1
Gastropoda
3
3 1 1 gastropod 7
Gastropoda
3
312 gastropod 19
Gastropoda
3
N
3 1 3 gastropod 22
Gastropoda
3
N
314 gastropod 26
Gastropoda
3
315 gastropod 29
Gastropoda
3
316 Artiphipholis squamata (Delle Chiaje)
Ophuiroidae: Amphiuridae
3
S
317 Lovenia camorata H. L. Clark
Echinoidea: Spatangidae
3
318 holothurian 5
Holothuroidea
3
Mem. QdMus. 18(2): 213-7. [1978]
A NEW SPECIES OF PONTELLOPSIS (COPEPODA,
CALANOIDA) FROM MORETON BAY, QUEENSLAND
J. G. Greenwood
University of Queensland
ABSTRACT
A new species, Pontellopsis tasmanensis sp. nov., found in coastal waters of eastern Australia
is described and figured, and the relationship of this to allied species is discussed.
A study of the zooplankton of Moreton Bay,
Queensland (Greenwood 1976) revealed the
presence of males and females of a species of
Pontellopsis which could not be allocated to any
known species and which is described here.
All specimens were collected with either
Clarke-Bumpus or 40 cm diameter conical nets of
nylon mesh with an aperture size of 195 microns.
Dissections were made in polyvinyl alcohol lightly
stained with chlorazol black, and drawings made
with the aid of a Wild M20 drawing tube.
Pontellopsis tasmanensis sp. nov.
(Figs 1 a-j, 2 a-f)
Materiai. Examined
HolotypE: Queensland Museum, W520I, J, Rainbow
Channel, Moreton Bay (27° 20’S., 153° 15’E.), SE.Q.,
coll. 8. vii. 1964, J. G. Greenwood (Station 1 of
Greenwood 1976).
ALLOTYPE: W5202, same data as Holotype.
Paratypes: W5203, 5c3c3, Sjj, same data as
Holotype.
P. tasmanensis was taken on 45 occasions during
1963-66 from Rainbow Channel, central Moreton Bay,
and Cleveland Channel (Stations. I, II, and 111 of
Greenwood 1976). Males were taken on 25 occasions,
females on 29 occasions, males with females on 9
occasions. Material other than the type series was
examined in less detail and discarded.
Description of Female (Fig. 1)
Body very robust. Total body length varying
between 142-1 *69 mm (av. 10 individuals 1-56
mm). Lenth ratio of metasome to urosome 2-98:1;
metasome length to width ratio 1-62:1.
Metasome with fourth and fifth thoracic
segments fused; posterior margins of fifth segment
produced symmetrically into small rounded
posterolateral processes. (Fig. 1 a, b, c).
Urosome two-segmented, genital segment
approx. 1-47 limes length of anal segment. Genital
segment asymmetrical, right postero-dorsal region
produced into rounded or broadly triangular lobe
(Fig. 1 d, e) which extends laterally to maximum
distance at level of posterior third of segment. This
lobe truncated abruptly posteriorly, at right angles
to body axis. Degree of development of this lobe
shows some variability. Most individuals as in Fig.
1 d, but others with larger lobes, to the maximum
found as in Fig. 1 e. Forms with larger lobes
sometimes with a small seta at tip of lobe and
similar small seta on mid-ventro-lateral surface.
No other spines or projections present on genital
segment. Anal segment produced postero-dorsally
into large, slightly asymmetrical triangular
supra-anal plate reaching to posterior level of
furcal rami (Fig. 1 d, e). Furcal rami symmetrical,
length c. twice width.
Fifth legs asymmetrical (Fig 1 f, g). Right
exopod with stout spine developed from inner
border slightly distal to midpoint, not reaching
end of ramus; corresponding spine absent from left
exopod. Both exopods terminate in single point.
Right exopod (Fig 1 h) with two subterminal
spines on outer border, the more distal the larger;
two minute spinules spaced more proximally along
outer border. Left exopod (Fig I i) with cluster
of three subterminal spines, two forming lateral
borders of small plate on which third is mounted;
214
MEMOIRS OF THE QUEENSLAND MUSEUM
GREENWOOD: A NEW SPECIES OF PONTELLOPSIS
215
two minute spinules more proximally along outer
border, first being just proximal to the ‘plate'.
Endopods similar, bifid terminally, but left
endopod narrower and longer, about 1-3 times
length of right endopod.
Description of Male (Fig. 2)
Body very robust. Total body length varying
between 1-28-1 -54 mm (av. 10 individuals 1-36
mm). Length ratio of metasome to urosome
2-58:1, metasome length to width ratio 1-96:1.
Metasome with fourth and fifth thoracic
segments fused; posterior margins of fifth segment
rounded on left side, right side produced as
elongate spine-like process reaching to mid or
posterior border of fourth urosomal segment (Fig.
2 a, b).
Urosome five-segmented, segments 1-5 in
proportions 40:15:17:12:16 (total 100).
Genital segment symmetrical, with single small
seta on right postero-lalcral margin (Fig. 2 c).
Second segment with patch of minute tubercles
on right postero-lateral surface. Third segment
slightly asymmetrical, slightly swollen on right
side, apex of swelling with minute tubercles.
Fourth and fifth segment without spines or
tubercles.
Fifth legs as figured (Fig 2 d, e, f). Right leg
with thumb-like process from base of proximal
exopod segment (Rel) similar in length to
finger-like distal exopod segment (Re2) thumb
slightly expanded or spathulate distally; large seta
on Rel distal to base of thumb. Re2 with
excavated lateral surface bearing one larger, one
smaller seta; small terminal, longer subterminal
seta; minimum distance between origins of
thumb and finger similar to length of segment.
Left leg with distal protopod segment (B2) and
Rel of similar length. Disto-lateral margin of Rel
with stout spine extending c. two-thirds length
Re2. Re2 short, c. half length Rel; length twice
width; lobe on medial face clothed with fine setae;
one long one short seta terminally.
Remarks
During much of the survey in which these
specimens were taken, the male and female forms
described here were regarded as belonging to
different species, the male being provisionally
related to P. tenuicauda and the female regarded
as a new form related, by structure of the fifth
legs particularly, to P. macronyx, P. scotti and
P. tenuicauda. Subsequent realisation of the
coincidence in frequency and occurrence pattern
of these two strongly suggested conspecificity.
There were, for example, no female P. tenuicauda
yet the apparent males occurred on 25 occasions.
More detailed examination of the two forms
showed some differences between these males and
those of P. tenuicauda as described by Mori
(1937), and considerable differences between the
present females and those of P. tenuicauda sensu
Mori (1937). and particularly sensu Giesbrechl
1892 (from which Mori’s also show differences).
Designation of the new species was therefore
necessary.
Both male and female share some characteris-
tics with several related species, but are
distinguished from them on combinations of
features. These distinguishing features are
discussed below, firstly for females, subsequently
for males.
FEMALES: The asymmetrical fifth legs of female
P. tasmanensis show some resemblance to those
of the following species: P. pexa Scott, P.
yamadae Mori, P. macronyx Scott, P. scotti
Sewell, P. tenuicauda (Giesbrecht):
P. pexa differs in that the right exopod there
terminates in three small spines, not a single point,
and although a spine is developed from the inner
border it arises distally and extends well beyond
end of the ramus (Scott 1909; p 173, pi. 54 fig.
General body form is similar to that of P.
yamadae. but although the female fifth legs of
that species are asymmetrical they differ from the
present form in having one (or more) large inner
spines from both exopods (Mori 1937, p. 98, pi.
47 fig. 6; pi. 48 fig. 13. Brodsky 1962, p. 147, pl.
48). P. yamadae is also a much larger calanoid
(2-8 mm, Mori 1937; 2-45 mm, Brodsky
1962).
Form of the fifth leg is very close to that of
P. macronyx and P. scotti. However in those
species the endopods are of similar length and the
exopods are not (reverse of P. tasmanensis). the
right exopod being shorter than the left (Scott
1909, p. 173, pi. 54 fig. 6; Sewell 1932, p. 388,
fig. e). P. macronyx further differs in armature
of the genital segment, which has paired
dorso-lateral spines both anteriorly and posterior-
ly, in having a smaller anal plate, and it is a
slightly larger copepod (1-73-1-97 mm), (Scott
Fig 1 Pontellopsis tasmanensis sp. nov., female, a, lateral view; b, dorsal view; c, Th5 and urosome with P5
in situ lateral view d, urosome with typical development of genital segment; e, urosome with maximal,
development of genital segment (as in ‘e’ above); h, detail of terminal region right P5 exopod; i, detail terminal
region left P5 exopod, j, proximal portion of right antennule.
216
MEMOIRS OF THE QUEENSLAND MUSEUM
1909; Sewell 1932; Kasturirangan 1963). P. scotti
differs additionally in having: an elongate anal
segment without extended anal plate, and on
which the furcae are set asymmetrically; paired
lateral spines anteriorly on the genital segment,
the posterior end variously developed but typically
with a swelling on the left side (elongate in some),
some with rod-like spine on right side (Sewell
1932, fig 129a)-
Closcst similarity in fifth leg structure is with
P. tenuicauda. However in that species the genital
segment does not develop a bulbous postcro-lateral
projection, having instead two lateral spines on the
right side (some variability is apparent in
development of these spines, cf. Giesbrecht 1892,
pi. 41 fig. 43, and Mori 1937, pi. 46 fig. 8). P.
tenuicauda also has a small anal plate (not
reaching mid-region of furcae), and very small
Fi(.. 2. Pontellopsis tasmanensis sp. nov., male, a, dorsal view; b, lateral view; c, urosomal segments 1-4; d, P5;
e, terminal portion left P5; f, anterior view, right P5.
GREENWOOD: A NEW SPECIES OF PONTELLOPSIS
217
fifth thoracic segment lobes (see espec. Giesbrecht
1892).
Sherman (1964) took females of a Pontellopsis
sp. from the central South-Pacific which could not
be identified with published descriptions but
which were similar to those of P. macronyx. No
further details of that species were given by
Sherman, but his females must be similar also to
those of P. tasmanensis.
MALES: P. tasmanensis males bear close
resemblance to those of P. scotti, P. yamadae and
P. tenuicauda.
They differ from P. scotti in relative lengths of
the 'thumb and finger' (Re2) of the right fifth leg;
absence of the central swelling found in the right
fifth thoracic segment process of P. scotti (Sewell
1932, p.390, fig. 129 b, f).
P. yamadae males are almost twice the size of
the present forms (2-5 mm, Mori 1937, p. 99) and
differ in some further details: Right fifth leg
‘thumb’ shorter than ‘finger' in P. yamadae
(slightly longer in P. tasmanensis) and without
spathulate tip; distal spine on Rel of left fifth leg
very short in P. yamadae.
P. tenuicauda is only slightly larger than P.
tasmanensis (c. ! -65 mm, Mori 1937, p. 98) and,
according to Mori’s description and figure, closely
similar. It differs in that: The right fifth thoracic
segment process is longer, reaching to posterior
border of anal segment; the second urosomal
segment is asymmetrical with pronounced swelling
on right side similar to that on the third segment;
a large seta is borne on the base of the thumb;
the thumb is not spathulate distally. Mori (1937,
pi. 46 fig. 10) did not show any spine distally on
the left Rel, but this may have been an
omission.
LITERATURE CITED
Brodsky, K. A., 1962. On the fauna and distribution
of Calanoida of surface waters in the north-western
part of the Pacific Ocean. Issled. dalnevost. Morei
SSSR 8: 91-166.
GiLSBRtCHT. W., 1892. Systemalik und Faunistik dcr
Pelagischen Copepoden dcr Golfes von Neapei.
Fauna Flora Golf. Neapei, Mongr. 19: 1-831.
Grei-nwood. J. G., 1976. Calanoid copepods of Moreton
Bay (Queensland) I. Families Calanidae, Eucalan-
idae and Paracalanidae. Proc. R. Soc. Qd 87:
1-28.
Kasturirangan, L. R., 1963. A key for the
identification of the more common planktonic
copepoda of Indian coastal waters. Ind. Nat. comm,
on Oceanic Res. Publ. 2: 1-87.
Mori. T., 1937. The pelagic Copepoda from the
neighbouring waters of Japan, pp. 1-150.
(Reprinted 1964 by Soyo Co. Inc; Tokyo).
Scott. A., 1909. The copepoda of the Siboga
Expedition, Part 1. Free-swimming, littoral and
semi-parasitic Copepoda. Sibioga Exped. Mongr.
29a: 1-323.
Sewell. R. B. Seymour. 1932. The Copepoda of the
Indian Seas. Mem. Indian Mus. 10: 223-407.
Sherman. K., 1964. Pontellid copepod occurrence in the
central South Pacific. Limnol. Oceanogr. 9(4);
4 ' 76 - 84 .
Mem. Qd Mus. 18(2); 219-31, pis. 31-8. [1978]
ASTHENOPSIS WHITEHOUSE, 1939 (TRIEOBITA, MIDDLE CAMBRIAN)
IN NORTHERN AUSTRALIA
Peter A. Jell
University of Queensland
ABSTRACT
Comparisons of the Trilobite Asthenopsis with other genera, previously confused with it,
show it to be a distinct taxon containing six Australian species. Apart from the type species,
Asthenopsis levior Whilehouse, 1939, five are described as new {queenslandica. rhinostrongyla.
unquinsepta. butorosa, and opalensis). Another new species is recognized but not named and
another population is compared with A. rhinostrongyla. Some remarks are also made on muscle
insertions.
Since Whilehouse erected the genus in 1939
and introduced Asthenopsis levior as type, no new
species have been assigned and those European
species placed there by Whilehouse may now be
more correctly placed in other genera. Of the
species from China referred by Walcott (1913) to
Solenopleura, by Kobayashi (1935) to
Solenoparia. and by Whilehouse (1939) to
Asthenopsis. none belong to Asthenopsis. No
references to Asthenopsis have appeared since
1939 other than the short note (Opik 1967, p.l84)
indicating its ptychoparioid affinities and several
cursory identifications at the generic level.
Terms used in the systematics are those defined
by Harrington, Moore, and Stubblefield {in
Moore 1959), glabella being used to include the
occipital ring. Notation for muscle areas of
glabella and points on facial suture follow Richter
and Richter (1940). 1 have referred to any sagittal
or exsagiltal dimension in terms of length and any
transverse dimension in terms of width.
Figured material is housed in the Common-
wealth Palaeontological Collection (CPC),
Department of Geology, Australian National
University (ANU), or Department of Geology,
University of Queensland (UQF). Locality
numbers referred to are in the registers of the
Bureau of Mineral Resources, Canberra (M, H,
or MNF), Department of Geology, Australian
National University (ANU), or Department of
Geology, University of Queensland (UQL).
Genus Asthenopsis Whilehouse, 1939
Type Species; Asthenopsis levior Whilehouse,
1939 by original designation.
Generic Diagnosis
Ptychoparioid with glabella of slightly variable
length, three or four pairs of lateral glabellar
furrows, an anteromedial glabellar muscle scar,
slightly tapering, straight to very slightly waisted
lateral glabellar margins. Furrow Ip long and
forked adaxially. Fixed cheeks w'idc (0-7 - 0-9 of
basal glabellar width). Eye ridges paired or
tripled. Anterior axial furrow weakly or not
impressed, always much weaker than lateral axial
furrow, and shallowing through ontogeny.
Fossulae deep. Brim 0-2 - 0-5 of glabellar length.
Border furrow with wide pits anterior to the
fossulae. Palpebral lobes short and near midlength
of cephalon or slightly posterior to it. Free cheek
with high eye socle, with strong anterior doublural
projection, and genal spine that is commonly
deflected slightly laterally to break the cephalic
marginal curve at the level of the posterior of the
palpebral lobe.
Thorax of 14 segments. Pleural lips either
rounded, broadly squared or spinose. Pygidium
relatively small (c. 0-3 of cephalic length),
transverse, and moderately vaulted to flat in
pleural areas. Border furrow poorly to not
impressed. Doublure no wider than border, usually
220
MEMOIRS OF THE QUEENSLAND MUSEUM
very much less. Ornament highly variable from
species to species including smooth, finely
papillose, and tuberculate (both fine and
coarse).
Comparison
Solenopleura Angelin, 1854 with holometopa
as type is a distinct genus characterized by the
lack of preglabellar field, glabella occupying a
larger percentage of the cranidial surface, fixed
cheek 0-6 or less of basal glabellar width
(canaliculata is probably not congeneric), almost
aspinose cheek, and more quadrangular pygidium.
Jincella Snajdr, 1957 with S. prantli Ruzicka,
1944 as type is distinguished from Asthenopsis by
the short convex border, short preglabellar field,
well impressed anterior axial furrow, rather
conical glabellar shape, narrow fixed cheeks, well
impressed glabellar furrows, and long narrow
posterior limb. Solenoparia Kobayashi, 1935 with
Ptychoparia toxeus Walcott, 1905 (Walcott 1913,
pi. 19, fig. 10) as type is distinguished by its short
convex border, narrow fixed cheeks, longer
palpebral lobes, glabella occupying larger part of
cranidial area, and less transverse pygidium.
Kounamkites Lermontova in Chernysheva, 1956
based on virgatus Chernysheva, 1956 is distin-
guished by the short border relative to brim,
strongly diverging anterior branches of suture,
well impressed anterior axial furrow, undeflected
genal spines, and less transverse pygidium.
The content and validity of the several genera
mentioned above are not commented on herein as
1 have had no first hand experience with them.
However, they are sufficiently known to be sure
of their distinction from Asthenopsis.
Eosoptychoparia kochihei (Walcott, 1911)
from Manchuria represents a very similar genus
even to the deflected genal spine but distinctions
are evident. I shall describe E. kochibei more fully
in a forthcoming paper.
Musculature
As illustrated for Triarthrus (Cisne, 1974, fig.
2) the ventral longitudinal muscles of Asthenopsis
are interpreted as running forward to the two wide
slightly deeper areas of the anterior border furrow
situated directly in front of the fossulac. These
muscles that run back to the major ventral
endoskeletal bars to which attach many of the
muscles of the mouth parts and other appendages,
are probably strongly digitate anteriorly (by
homology with the cephalocarid Hutchinsonieila,
see Hessler 1964) with many small attachment
sites on the cuticle.
In passing it should be noted that a large
percentage of trilobites have some structures in or
near the anterior border furrow (i.e. low anteriorly
on the sagittal or exsagittal glabellar profile) that
can be readily interpreted as ventral longitudinal
muscle attachment sites. Those lacking such
structures are not precluded from having muscle
insertions in this position but are simply
interpreted as not showing them in any distinctive
way. In the Crepicephalidae the two or three wide
distinctive pits in the border furrow are formed
by several smaller circular pits providing
apodemes for the digitations of the ventral
longitudinal muscles. This structure also suggests
that broad depressions and discrete pits may be
analogous. Many post-Cambrian trilobites exhibit
strong apodemes or muscle scars well forward on
the cephalon that would be ideally placed for
attachment of the ventral longitudinal muscles.
The dorsal longitudinal muscles of Asthenopsis
are interpreted as anchoring anteriorly to the
glabellar furrows and posteriorly to the apodemes
of the transaxial furrows. While these muscles
form single sheaths the full length of the animal,
they are also attached to each segmental
apodemal area to allow intersegmental mechan-
isms to function.
A very small muscle scar is situated
anteromedially on the glabella. I am unable to
find the description of any other ptychoparioid
referring to a similarly situated muscle scar. Jell
(1975, p, 23) interpreted a muscle scar in this
position on a species of Pagetia and discussed its
possible function as a hypostomal diductor muscle
by direct homology with the living notostracan
Triops. In many dalmanitaceans a pit is present
anteromedially on the glabella (Campbell in
press) which may be homologous with that of
Pagetia and indeed with that of Asthenopsis.
Affinities
The taxonomic position of Asthenopsis is not
clear. Originally placed in the Solenopleuridae it
has been regarded as correctly assigned except for
the statement by Opik (1967, p. 184) that it may
be regarded as a ptychopariid. I agree with Opik
and believe that in the light of new species
described herein the genus can be confidently
divorced from any close relationship with the
group of genera that have come to be known as
the Solenopleuridae. I shall not make any
assignment at this stage as I am at present
engaged in a numerical taxonomic study of
ptychoparioids and do not wish to pre-empt those
results.
JELL; MIDDLE CAMBRAIN TRILOBITA
221
Asthenopsis levior Whitehouse, 1939
(Fig. la; Plate 31, figs. 1-9; Plate 32, figs.
1-5)
Asthenopsis levior Whitehouse, 1939, p. 214,
pi. 22, figs. 17-20.
MaTI RIAI Ewminfd
Hol.oTVPi' UQF3337, an external mould of a
complete exoskeleton the counterpart of which is missing
from the University of Queensland collections, from V
Creek at the Undilla to Camooweal Road crossing.
Otiiir Mati RIAI Four cranidia (C), I free cheek
(FC), 2 pygidia (P), and 1 thorax (T) from M41 at the
base of an isolated hill of Split Rock Sandstone 4-8 km
east of Douglas Creek on the Old Burketown Road (lat.
19° 27-5 'S., long. 138° 37 'E); 3C, 2P. and one complete
from M247 at Barkly No. 7 Bore west of Yelvertoft;
2C from the left bank of Douglas Creek halfway from
the Old Burketown Road to the O’Shannassy River; 3C
from 12 km south of Douglas Spring on the ‘Morslone’
to ‘Undilla’ Road; IC and 2T from left bank of Douglas
Creek at ‘Morstone’; 2C from M52 1-3 km west of M41;
IFC from Ml 39 just west of Top Hands Waterhole on
the upper reaches of Harris Creek; 5C. I P, and 3T from
UQL469 on top of hill immediately east of the crossing
of Harris Creek by the Camooweal to Thorntonia Road;
3C and IFC from UQL3510 15 km south-west of the
Thorntonia airstrip on the Kangaroo Flat road. This
includes the figured specimens CPC 17064-8 and
UQF68711-4 (Pis. 31, 32). Whitehouse's originally
figured specimens were also examined and are refigured
herein (PI. 31 figs. 1, 4, 6, 7).
All localities are in the V Creek Limestone except
M52 which is in the Mail Change Limestone. The age
of the material is the Zone of Ptychagnostus
nathorsti.
Diagnosis
Outline regularly oval. Surface ornament
lacking or of minute papillae with rare slightly
larger tubercles. Glabella extending 0-7 of
cephalic length, with tapering straight or convex
lateral margins, with truncated anterior and
angular anterior corners. Three pairs of lateral
glabellar furrows, smooth to very poorly
impressed, with furrow Ip long and wide, furrows
2p and 3p successively shorter and narrower.
Occipital node at midlength of ring. Border flat
to weakly convex long and tapering laterally.
Facial suture with slightly convex anterior branch
in exsagittal line or slightly convergent. Eye ridges
variable in prominence, paired. Palpebral lobe
short, just behind midlength of glabella. Caecal
ornament on extra-glabellar parts of cephalon.
Thorax of 14 segments. Pleural furrows long, well
impressed, running almost to pleural tips. Pleurae
downturned in fulcral line, with spinose tips.
Pygidium about 0T5 of total length. Axis with
three rings and terminus. Pleural areas highly
vaulted with four pairs of ribs. Posterior margin
concave.
Description
The holotype designated by Whitehouse (1939,
p. 278) has a cephalon 9 mm long but other
specimens have been collected with cephala up
to 16 mm long so the type is not a fully mature
individual. This description is based on the more
mature material and its variations from the
morphology of the holotype are discussed below
under DISCUSSION OF Holotype.
Exoskeleton almost twice as long as wide,
moderate overall convexity with steeper slopes
towards the margin. Cranidium two or three times
length of pygidium. Overall shape oval but
somewhat more rounded anteriorly than
posteriorly.
Cranidium moderately vaulted, twice as wide as
long, with moderately steep slope to border furrow
in lateral profile. Glabella tapering forward to half
its basal width, with straight or slightly convex
sides, truncated almost transverse anterior.
Anterior axial furrow very poorly impressed
(represented almost entirely by a change of slope
in lateral profile), never as well impressed as
lateral axial furrow. Axial furrow very broadly V
shaped in section, with deep fossulae anteriorly,
floor raised adjacent to lobe Ip, running directly
past occipital ring to margin. Occipital ring five
times as wide as long, with prominent sagittal
node at the midleigth, posterior margin convex,
tapering laterally into a low ridge that crosses the
axial furrow into the posteroproximal corners of
the fixed cheek. Occipital furrow poorly
impressed, slightly deeper laterally, with smooth
surface of apodemal pit extending up onto lobe
Ip and occipital ring. Glabellar furrows three
pairs in number, defined mainly as smooth areas
on an otherwise weakly ornamented glabella.
Furrow Ip long, directed at 30° to transverse line
out of axial furrow then posteriorly at 60° to
transverse line in a shorter adaxial terminally
expanded section, with medial swelling just out of
axial furrow. Furrow 2p shorter and narrower
than Ip, of same general shape but with much
narrower unexpanded adaxial part and directed
less posteriorly. Anteromedian glabellar muscle
scar present.
Brim downsloping much more steeply and twice
as long at facial suture as axially, convex
becoming more so laterally, very slightly
depressed from fossulae forward to border furrow,
with prominent anastamosing caeca originating
out of the eye ridge and running forward into the
222
MEMOIRS OF THE QUEENSLAND MUSEUM
Fkj. 1 A: Reconstruction of Asthenopsis levior Whitehouse, 1939 based
on holotypc and UQF44314.
border. Border furrow poorly to moderately
impressed, with a pair of very wide indistinct pits
directly anterior to the fossulae (ventral
longitudinal muscle attachment sites), transverse
except for slight convexity between wide pits.
Anterior border moderately convex often accen-
tuated by sharp posterior slope but always with
gently sloping anterior, highest point at midlength.
Eye ridges double, prominent, continuing through
the fossulae and around front of glabella as low
parafrontal band, directed posteriorly at approx-
imately 20° to transverse line. Palpebral lobe 0-25
of cephalic length, abaxially upturned, and
posteriorly downturned, moderately arcuate,
opposite furrow Ip and lobe 2p. Palpebral furrow
moderately impressed, shallowing at midlength.
Fixed cheeks 0*82 basal glabellar width,
moderately convex, rising gently to palpebral lobe.
Posterior limb wide and long, almost rectangular
in shape except for slightly convex and posteriorly
directed anterior margin, sloping steeply abaxial-
ly, with faint caecal system posterolaterally from
palpebral lobe. Posterior border furrow lengthen-
ing and deepening laterally, turned slightly
forward at extremity. Posterior border lengthening
and flattening laterally, with marked ridge
forming high point for most of its width but
meeting the posterior margin two thirds of
distance from fulcrum to u'. Posterior margin
straight to fulcral line (used in sense of Opik,
1967, p.57) then convex. Facial suture with o- n
only 0-6 of /?- v variable from convex and
converging to straight and exsagittal, (S at
midlcngth of palpebral lobe, e at posterior of
furrow Ip. Free cheek with long anterior extension
where facial suture cuts obliquely across border,
y^at posterior of border furrow. Margin evenly
curved to base of rounded gcnal spine, then
deflected slightly abaxially down spine. Border
wide, flat to weakly convex. Eye socle low, of
JELL: MIDDLE CAMBRAIN TRILOBITA
223
B
Fig. IB: Reconstruction of Asthenopsis queenslandica sp. nov. based
on hoiotype and CPC 17072.
constant height. Genai field exhibiting very low
anastamosing caeca.
Rostral plate and hypostoma unknown at
present. Thorax of 14 segments each 12 times as
wide as long. Articulating half ring slightly lower
than axial lobe, shorter than furrow or lobe, with
prominent transverse ridge near posterior forming
high point and dividing smooth relatively flat
anterior part from steeper granulose posterior
part. Articulating furrow with posterior wall
steeper and higher, with fine papillose ornament
anteriorly. Axial lobe expanding slightly laterally
to occupy full segmental length, flat (sag.) with
transverse posterior margin. Accessory apodemal
pits low abaxially, joining articulating furrow and
posterior margin. Pleural furrow occupying
40-50% of fixed pleura, swinging forward and
shallowing adaxially, meeting axial furrow a short
distance from margin, with very shallow posterior
arm adaxially giving a forked appearance.
deepening and shortening on free pleura before
finishing at the midlength near the base of the
pleural spine. Pleural strips of equal length except
on free pleura where anterior strip shortens as it
runs obliquely behind wide moderately long
smooth facet. Pleural spines stout, curving slightly
posteriorly. Segments not overlapping at all on
fixed pleura (i.e. margins abutting) but overlap-
ping almost all facet on free pleura. Segments
transverse throughout.
Pygidium almost three times as wide as long,
well vaulted, with rounded margin except for
slightly concave section across axis posteriorly.
Articulating half ring extending in front of
otherwise transverse margin, with fine papillose
ornament posteriorly. First axial ring highest part
of pygidium, with posterior excavation on axial
three quarters cutting out most of segmental
length (sag.) with weak accessory apodemal pits
in exsagittal line close to axial furrow. Second
224
MEMOIRS OF THE QUEENSLAND MUSEUM
axial ring with almost imperceptible accessory
apodeme and smaller posterior excavation. First
and second iransaxial furrows moderately
impressed with wide undepressed apodemal areas.
Posterior transaxial furrows possibly three in
number, progressively weaker, and continuous.
Axis parallel sided with rounded posterior, with
moderately steep posterior slope to border and
margin. Pleural areas with three well impressed
pleural furrows and one poorly impressed
interpleural furrow anteriorly, furrows deepest in
fulcral line. Caecal ornament running out from
axial furrow along pleural strips, through border
furrow and into the border. Facets wide, relatively
short, triangular, smooth and steeply sloping.
Anterior pleural strip of first segment short and
convex to fulcrum then longer, less convex and
oblique beyond. Border furrow poorly impressed
more so behind axis. Border flat and of uniform
width. Ornament of fine papillae on axial rings,
terminus, pleural ribs and border of some
specimens, furrows smooth.
Discussion of Holotype
As already mentioned the holotype specimen
represents an immature individual so it varies in
several minor respects from the description given
above. With the recognition of these morphogene-
tic changes further outlined under Morphogeny
below comes the realization that the species has
considerable intraspecific variation between
individuals of the same and only slightly different
size.
The holotype is crushed with only slight
displacement of the free cheeks that do however,
obscure the course of the facial suture. It has fine,
close-spaced papillae with an occasional very
widely spaced tubercle. Details of occipital ring
and glabellar posterior are lost as the exoskeleton
of that part has remained in the counterpart.
Fossulae are deeper so the frontal area is more
convex (exsag.) but less steeply sloping. The
caecal network on the brim and eye ridges is less
prominent. The palpebral furrow is abaxially
convex and the lobe more strongly upturned.
Fixed cheek width and basal glabellar width
almost the same. On the thorax the ornament is
coarser axially but less apparent on the pleura. On
the pygidium the posterior margin is straight, the
interpleural furrow is very weak, the axis does not
reach so close to the margin, and the pleural areas
are not so vaulted.
Morphogeny
The smallest individual with a complete thorax
(Plate 31, fig. 2) has 14 segments and a cephalon
4 mm long so that it must be a very early
holaspide. A slightly smaller cranidium (Plate 31,
fig. 3) 3-5 mm long is similar to that of the
complete individual. While the cepahlon is shorter
than 5 mm the fixed cheeks are more convex, the
axial furrow is deeper, the ornament is coarser,
the furrows forward from the fossulae are more
evident, the palpebral lobes are relatively long
extending from lobe Ip to furrow 3p or half
glabellar length, pleural tips are not spinose but
simply pointed.
Individuals with cranidia 5-10 mm long are
characterized by the holotype described above but
it should be noted that most individuals of this
size range have coarser papillae than the
holotype.
Affinities
Asthenopsis levior is quite variable from
locality to locality but only in minor details of
glabellar shape, ornament, relief, and anterior
course of facial suture. As no uniform variation
of these characters can be observed and as the
variation is not as great as between this and other
species of the genus described below no other
species have been separated from the relatively
distinct (within the genus) levior.
Asthenopsis queenslandica* sp. nov.
(Fig. lb; Plate 33, figs. 1-8; Plate 34, fig. 1)
Material Examined
Holotypf: CPC 1 7074, a complete specimen from
M243 on the flood plain of Whistler Creek, 0-8 km from
its junction with the Buckley River, 42 km west of
Yelvertofl Homestead (Lat. 20° 04' S.. long. 138° 30'
E): V Creek Limestone. Zone of Ptychagnostus
nathorsti.
Other Material; Three complete specimens, 14C,
2FC and 3P including CPC17070-3 and 17075-8 (Pis.
33 and 34) from M243; 2C from UQL467 4-8 km west
of Redbank Creek, just west of laterite ridge between
Redbank and Harris Creeks; 20C, IFC, 8P, and several
thoracic segments from UQL463 top of hill 6 km south
of Thorntonia Homestead on left bank of West
Thornton River.
Both UQL463 and 467 are in the V Creek Limestone
with 463 belonging to the Euagnostus opimus Zone and
467 to the Ptychagnostus punctuosus Zone.
Diagnosis
Asthenopsid with relatively square anterolateral
glabellar corners, low cephalic convexity generally
and in the anterior margin, four pairs of smooth
♦Named for the state of Queensland.
JELL: MIDDLE CAMBRAIN TRILOBITA
225
to very poorly impressed lateral glabellar furrows,
anteriorly placed occipital node, relatively long
brim, doubled eye ridges, relatively narrow fixed
cheeks, straight slightly converging anterior
branches of facial suture, fine close spaced
papillose ornament with sparse small tubercles
superimposed. Pleural tips squared. Transverse
pygidium having flat pleural areas, no border
furrow, and an evenly curved margin.
Description
Cranidium four times as long as pygidium, very
weakly vaulted. Glabella standing above flat to
slightly abaxially rising fixed cheeks in anterior
profile, subquadrale tapering only slightly forward
from furrow Ip, with angular anterolateral
corners and only slightly convex anterior margin.
Occipital ring with small sagittal node anteriorly.
Occipial furrow well impressed, short, and smooth
(apodemal part) with steeper anterior than
posterior wall laterally and shallow, slightly
longer, with papillose ornament and steeper
posterior than anterior wall axially. Glabellar
furrow 4p may be present low in the axial furrow.
Glabellar ornamented with fine papillae except on
the apodemal areas. Brim downsloping slightly
more steeply at the facial suture than axially. Eye
ridges only slightly elevated, divided into two
parallel ridges. Fixed cheeks 0-73 of basal
glabellar width, rising gently to palpebral lobe
with fine papillose ornament and a very few
slightly larger tubercles interspersed, with very
faint caecal network radiating from lobe Ip and
from the rear of the eye. Posterior limb elongate
triangular in shape, sloping steeply down
abaxially. Facial suture with ji - 7 straight and
very slightly converging, tS behind midlength of
palpebral lobe.
Free cheek relatively flat, not vaulted, ji at
posterior of border. Border convex anteriorly,
flatter posteriorly, with one or two continuous
parallel terrace lines near the margin, with
papillose ornament. Genal spine with flat dorsal
and slightly rounded ventral surfaces. Thorax of
14 segments each 15 times as wide as long. Each
pleuron with fine papillae and single row of
tubercles (up to 7). Free pleura with rounded
anterolateral and semisquared posterolateral
corners. Anterior fixed pleural margin just under
posterior of next anterior segment but with
margins directly opposed in same plane for short
distance at fulcrum then overlapped on the free
pleural facets. Segments becoming slightly
anteriorly convex near the pygidium.
Pygidium with an unbroken marginal curve,
and a low profile with axis standing only slightly
above flat pleural areas. Furrows all shallower
than in A. levior, decreasing in depth posteriorly
and laterally. Border furrow not impressed.
Border may be defined as area beyond pleural
furrows. Fine papillae on areas out of furrows and
off muscle scars.
Morphogeny
Although no small specimens with a full
complement of segments are available some
comments on the morphological changes with
increased cranidial size are pertinent. The adults
described above all have cranidial length of 10-12
mm whereas two other groups exist with cranidial
lengths of 3 mm and 5-6 mm respectively.
The 3 mm cranidia (Plate 33, figs. 2, 4) have
a second order of larger tubercles .scattered over
the entire exoskeleton giving the same ornament
as in mature individuals of A. rhinostrongyla.
They have an anterolaterally rounded glabella,
almost no glabellar furrows, better impressed
anterior axial furrow, convex fixed cheeks Just
slightly more than half basal glabellar width and
with no apparent division of the eye ridge.
The 5-6 mm cranidia (Plate 33, fig. 7) have
the same ornament as the 3 mm cranidia, the
anterolaterally rounded glabella, very poorly
impressed glabellar furrows, virtually no anterior
axial furrow, convex fixed cheeks 0-6 of basal
glabellar width, and paired eye ridge.
Astbenopsis rhinostrongyla* sp. nov.
(Fig. 2A; Plate 34, figs. 2-7;
Plate 35, figs. 1, 2)
Matf-rial Examined
Holo TYPE CPC 1 7080. a cranidium from H 1 38 on the
Huckitla 1:250,000 Geological map at 22°35’ south
latitude, 136° 02’ east longitude on Arthur Creek,
Northern Territory; Arthur Creek Beds, Euagnostus
opimus Zone.
Other MATF.RtAL: Seven C, 3 FC, 1 T, and 3 P
including CPC 17079, 17081-6, (Pis. 34 and 35) from
H138.
Diagnosis
Asthenopsid with rounded anterolateral glabel-
lar corners, flat unarched anterior border,
moderate convexity, anterior branches of facial
suture diverging forward and abaxially convex,
relatively long pelpebral lobes, ornament of very
close spaced papillae over entire surface except
apodemal areas, palpebral lobes and furrows, and
coarse perforated tubercles interspersed sparsely
^Rhinos (Gr.) nose, strongylus (Gr.) rounded, refers
to the rounded glabellar anterior.
226
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig. 2: Reconstruction of cephalon and pygidium of Asthenopsis species.
A: Asthenopsis rhinostrongyla sp. nov. based on the
holotype, CPC 17085, and CPC 17086.
C: Asthenopsis butorasa sp. nov. based on the
holotype, ANU30646, and ANU30647.
B; Asthenopsis opalensis sp. nov. based on the
holotype, CPC17100, and CPC17102.
D: Asthenopsis unquinsepta sp. nov. based on the
holotype, CPC17087, and CPC17088.
JELL; MIDDLE CAMBRAIN TRILOBITA
227
over the same areas. Pleural tips rounded.
Transverse pygidium with flat pleural areas.
Description
Cranidium with moderately steep anterior slope
in lateral profile. Glabella subquadrate tapering
only very slightly forward to 0-8 of its basal width.
Occipital furrow with markedly deeper apodemal
pits, almost dumb-bell shaped apodemal areas
extending onto lobe Ip and occipital ring. Four
pairs of lateral glabellar furrows smooth, not
depressed. Furrow Ip with adaxial part not
expanded. Furrow 3p separated from axial furrow.
Furrow 4p very indistinct, narrow and smooth.
Occipital node situated anteriorly. Axial furrow
distinctly shallower adjacent to lobe Ip and
occipital ring. Eye ridge quite low and formed of
two trunks. Fixed cheeks 0-8 of basal glabellar
width, slightly convex and horizontal. Posterior
limb steeply downsloping. Facial suture with
(> - ji convex and diverging. Posterior border
furrow lengthened slightly laterally. Posterior
border only moderately convex and slightly longer
laterally, with its ridge not markedly distinct and
not obvious beyond fulcrum. Ornament of papillae
and sparse perforated tubercles over cranidium
except on furrows, palpebral lobe and distal parts
of posterior limb.
Free cheek doublure without ornament,
extending only slightly adaxial to long anterior
dorsal projection. Thorax of at least 12 segments
(exact number unknown). Anterior half of half
ring smooth but posterior half sloping into furrow,
moderately papillose. Up to 14 large perforated
tubercles on each pleural strip. Posterolateral
pleural corner well rounded. Facet more than half
segmental length.
Pygidium transverse with second transaxial
furrow poorly impressed. Axis with relatively
gentle posterior slope to flat border area. Pleural
areas almost flat with one well and one poorly
impressed pleural furrow separated by long flat
rib bearing a poorly impressed interpleural furrow.
Anterior strip of first segment lengthening
markedly at and beyond fulcrum. Ornament of
papillae everywhere except in furrows and close
packed on high parts with large sparsely scattered
tubercles over the same areas.
Asthenopsis sp. cf. A. rhinostrongyla sp. nov.
(Plates 38, figs. 3-7)
Material Examined
Two C with FC, IFC, and 3T including CPC17103
to 17105 and UQF68715 to 68716 from Ml 56 (located
on Camooweal 1:250,000 Geological map) on the middle
reaches of Opal Creek in the Age Creek Formation,
Ptychagnostus punctuosus Zone, of the Currant Bush
Limestone.
Description
Those features not consistent with A. rhinos-
trongyla are referred to comparatively in this
description. Glabella with better impressed
anterior axial furrow producing a distinct break
in lateral profile. Axial furrow poorly impressed
throughout. Palpebral lobes narrower, markedly
upturned laterally. Fixed cheeks almost as wide
as the base of the glabella. Facial suture with o
much closer to /i . Ornament, especially the
papillae, more subdued, larger tubercles fewer in
number. Free cheek with terrace lines prominent
near margin. Border slightly narrower. Genal
spine slightly more abaxially deflected. Pygidium
unknown at present.
Remarks
The nature of the glabella, anterior border and
palpebral lobes along with the slightly different
ornament make reference to A. rhinostrongyla
difficult. However, when a larger population of
that Northern Territory species is studied the
limits of variation may be such that this material
will be assignable. It should be pointed out that
the cephalic convexity, anterior border arch,
subtle ornament, and in one instance (Plate 38,
fig. 6) short frontal area also ally this species to
A. opalensis. However, shape of the glabella and
its furrows, eye ridges, shorter palpebral lobes,
and straight anterior parts of facial suture
distinguish that species.
Asthenopsis unquinsepta* sp. nov.
(Fig. 2D; Plate 35, figs. 3-8)
Material Examined
HolotypE; CPC17089, a cranidium from M157 on a
sharp bend near the second creek junction on Opal
Creek, latitude 19° 25' S, longitude 138° 35' E.; Age
Creek Formation, Zone of Ptychagnostus
punctuosus.
Other Material: Five C, 2 FC, and 2P including
CPC 1 7087, 17088, and 17090-2 (PL 35) from
M157.
Diagnosis
Asthenopsid with glabella tapering forward to
half its basal width, glabellar anterior truncated,
anterolateral glabellar corners quite angular,
furrow Ip with prominent domed area at its
*From the latin numerals 1, 5, and 7, the material
being from M157.
228
MEMOIRS OF THE QUEENSLAND MUSEUM
midiength abaxially, frontal area relatively long
and fixed cheeks of moderate width, anterior
branch of facial suture straight and converging
slightly forward, eye ridges doubled, ornament on
border and glabellar of fine tubercles but on
frontal area and fixed cheeks of coarser close
spaced tubercles. Pygidium moderately vaulted.
Description
Glabella tapering anteriorly to half its basal
width, sharply truncated anteriorly by change in
ornament, anterior axial furrow not impressed
even by a change of slope. Occipital ring less than
four times as wide as long with medially placed
node. Occipital furrow poorly impressed on axial
half, with very steep anterior wall into pronounced
apodemal pits laterally. Furrow Ip with domed
abaxial elevation prominent. Glabellar ornament
of fine tubercles. Axial furrow very poorly
impressed throughout, with diverticulum from
lobe Ip conspicuous. Ventral longitudinal muscle
pits poorly impressed. Frontal area with
prominent caecal network standing well above
surface. Eye ridges doubled. Fixed cheeks almost
flat. Palpebral lobe flat, only very slightly
upturned and opposite furrow and lobe 2p. t - lc
directed posteriorly at moderate angle to
transverse line, crossing posterior border oblique-
ly. Posterior limb relatively long and subtriangular
in shape. Posterior border furrow increasing in
length beyond fulcral point. Ornament on anterior
border, on glabella except furrows of fine
tubercles closely spaced, absent on palpebral
lobes, posterior border furrow, and border furrow;
on fixed cheeks, posterior limb, and frontal area
ornament of close packed coarser perforated
tubercles, with occasional ones larger than the
rest.
Free cheek with ornament on genal field similar
to that of frontal area. Doublure extending
adaxially, with terrace lines only on marginal
curve, absent on ventral dorsally sloping part.
Thorax with each articulating half ring having
distinct transverse ridge near its midlength,
anterior wall of articulating furrow rising very
steeply to this ridge. Ornament of coarse tubercles
on pleural strips not arranged in a single row.
Pleural extremity squared, with rounded anterior
corner and right angled to slightly pointed
posterior corner. Pygidium with pleural areas
moderately vaulted, anteriorly almost as high as
axis that is sunken between pleural areas, axis well
above pleural areas posteriorly. Axis lightly
tapering to a well rounded posterior, almost flat
transversely between apodemal pits, with
apodemal pits becoming more adaxially placed
posteriorly. Facets narrow and short. Ornament of
coarse close packed tubercles on prominences and
border.
Morphogeny
An individual with cranidium 6-5 mm long
(Plate 35, fig. 6, lower) has 14 segments and can
be interpreted as an holaspide. Another with
cranidium 3-5 mm long (Plate 35, fig. 6, upper)
has 1 1 segments of an incomplete thorax and may
also be an holaspide. However, some mor-
phological differences between these individuals
and the mature ones (1 1 mm cranidia) are worth
noting.
At 3-5 mm ornament is fine with a few larger
tubercles, the axial furrow is better impressed
(possibly from compaction), anteriorly rounded
glabella, fi— converges more strongly forward,
and fixed cheeks arc narrower.
At 6-5 mm the ornament is of moderately
coarse close spaced tubercles, axial furrow is well
impressed, fixed cheeks are narrower, and more
convex, caecal ornament is subdued, /; - (S
converges forward, and thoracic pleura have a line
of up to seven large tubercles.
Asthenopsis butorosa* sp. nov.
(Fig. 2C; Plate 36, figs. 1-9)
MaT[-RIA1. EXAMINFI)
Hoi.gtvpE: ANU30649, a cranidium from
ANU 10325, 5-6 km west of Chummy Bore on the
Thorntonia to Camooweal Road; Currant Bush
Limestone, Ptychagnostus punctuosus Zone.
Other Material: One complete, 1 T, and 3 P
including ANU30645-8 (PI. 36) from ANU10325; 6C,
3 FC, and 2 P including CPC 17093-6 (PI. 36) from
Mi 61 at the base of the V Creek Limestone 12-8 km
east of Morstone Homestead (marked on Camooweal
1:250,000 Geological Map), Ptychagnostus punctuosus
Zone.
Diagnosis
Asthenopsid with highly vaulted cephalon,
rounded anterolateral glabellar corners, poorly
impressed anterior axial furrow represented by
considerable change in slope, markedly laterally
tapering anterior border, anterior branch of facial
suture convex diverging forward, with very steeply
sloping frontal area more than twice as long at
facial suture as sagittally, with relatively close
packed ornament of coarse tubercles on border.
* Bu (Lat.) large, torus (Lat.) rounded protuberance,
refers to the coarse tubercular ornament.
JELL: MIDDLE CAMBRAIN TRILOBITA
229
frontal area, fixed cheeks, glabella, occipital ring,
thoracic and pygidial axes and pleura.
Description
Cranidium (up to 16 mm long) with highly
vaulted cheeks and glabella, with steeply sloping
frontal area (especially anterolaterally) and
posterior limb, and with glabella standing well
above convex cheeks in anterior profile. Glabella
tapering slightly forward to 0-75 basal width, with
straight to very slightly waisted (at furrow Ip)
lateral margins, with anterior axial furrow
represented by a considerable change of slope (up
to 40°) and hence more distinct than in any other
species of the genus. Occipital furrow poorly
impressed over axial half, apodemal area
dumb-bell shaped extending up onto the rear of
lobe Ip and occipital ring.
Glabellar furrows smooth, Ip not expanded at
adaxial end and without abaxial dome shaped
elevation. Anteromedial muscle scar almost in
anterior axial furrow. Anterior border most
convex of any in genus, tapering strongly laterally.
Border furrow well impressed with prominent
axial and lateral anterior convexities. Frontal area
moderately convex, almost vertically downsloping
at facial suture, with depressions extending
forward from axial to border furrows. Fixed cheek
width almost equal to basal glabellar width,
moderately convex. Facial suture with ji - (S
abaxially convex and diverging forward. Or-
nament on glabella (except furrows), frontal area,
fixed cheeks (except palpebral lobe and furrow),
posterior limb, and posterior border of coarse (up
to 0-3 mm diameter) high tubercles, on anterior
border and palpebral furrow of slightly finer
tubercles, with tubercles on fixed cheeks becoming
more widely separated towards the axial
furrow.
Free cheek moderately vaulted, with quite high
eye socle. Genal spine with convex dorsal surface,
and slight adaxial curve in distal portion.
Ornament on genal field of close spaced tubercles
superimposed on caecal ornament.
Thorax of at least 14 segments (no complete
mature thorax known). Axial lobe with coarse
tuberculate ornament. Pleural furrow unforked
adaxially. Pleural strips with single row of
tubercles becoming less distinct on free pleurae.
Free pleura with squared to slightly pointed
posterolateral corner, with finer tuberculate
ornament than elsewhere. Pygidium just over
twice as wide as long. Pleural area with three or
four pleural furrows and one interpleural furrow.
Facets short, narrow, and directed strongly
posteriorly. Fulcra prominently raised. Marked
posterior excavation on first axial ring.
Morphogeny
Cranidium 3-5 mm long (Plate 36, fig. 1) much
less vaulted, with coarse but sparse ornament,
anterior part of facial suture converging
forward.
Cranidium 5*5 mm long (Plate 36, fig. 3)
vaulted, with coarse, closer spaced ornament,
anterior part of facial suture convex and diverging
laterally, fixed cheeks higher than glabella.
Asthenopsis opalensis* sp. nov.
(Fig. 2B; Plate 37, figs. 1-4;
Plate 38, figs. 1, 2)
Material Examined
Holotype: CPC17098, a cranidium from MNFI5,
1.6 km south of the junction of Opal Creek and the
O’Shanassy River; Mailchange Limestone,
Ptychagnostus punctuosus Zone.
Other Material: Six C, 1 FC, and 1 P including
CPC17097, and 17099-102 (Pis. 37 and 38) from
MNF15.
Diagnosis
Asthenopsid with anterolaterally rounded
glabella standing well above fixed cheeks,
moderate cephalic convexity, four pairs of lateral
glabellar furrows with smooth or punctate
surfaces, distinctive occipital apodemal pits,
anteriorly placed occipital node, eye ridges of
three parallel trunks, straight strongly convergent
anterior branches of facial suture, very fine close
packed papillose ornament overlain by sparse
larger tubercles. Pygidium transverse, weakly
vaulted, with papillose ornament.
Description
Cranidium moderately convex, with glabella
standing above convex fixed cheeks, a moderately
steep frontal area, and a moderately arched
anterior border. Glabella with rounded anter-
olateral corners, convex anterior, with straight
only slightly tapering lateral margins, with four
pairs of lateral glabellar furrows and an
anteromedial muscle scar. Occipital furrow
shallow over axial half. Occipital apodemal pit
distinctive, with wavy slightly raised smooth areas
both anteriorly and posteriorly giving an hour
glass shape with long axis transverse. Furrow 3p
slightly arcuate, directed very slightly to the
*Named for Opal Creek on which it is found.
230
MEMOIRS OF THE QUEENSLAND MUSEUM
posterior adaxially and not connected to axial
furrow. Furrow 4p short narrow, directed slightly
anteriorly out of fossula very close to glabellar
anterior. Glabellar furrows either smooth or finely
punctate. Axial furrow crossed by low distinct
diverticula at occipital ring and lobe Ip, 2p and
3p. Fossulae not at all distinct. Frontal area flat
short and downsloping axially but convex
laterally, with both slope and length (to twice
sagittal length) increasing laterally. Eye ridges
consisting of three parallel trunks (not visible in
all specimens) with the median one highest and
joining the anterior one just abaxial to the fossula.
Parafrontal band low, arising out of fossula, not
visible on all specimens. Palpebral lobe flat only
slightly upturned laterally, anterior opposite
anterior of lobe 2p, posterior level with anterior
of lobe Ip. Palpebral furrow straight to slightly
abaxially convex, deepending posteriorly. Fixed
cheeks 0-9 of basal glabellar width, slightly convex
and horizontal, with sparse tubercles over an
extremely faint anastomosing caecal network
radiating from the axial furrow posteriorly. Facial
suture with I'i - ji straight, converging strongly
anteriorly, t - concave adaxially then straight.
Posterior border much longer and flatter beyond
fulcrum. Free cheek moderately vaulted, with ji
in border furrow. Border flat throughout. Genal
field with sparse, coarse tubercles superimposed
on faint caecal network.
Pygidium transverse, with well rounded margin,
and flat pleural areas. Two pleural furrows and
an interpleural furrow present. Anterior pleural
strip of first segment lengthening well inside
fulcrum, weakly convex throughout. Marked
posterior excavation in first axial ring.
Asthenopsis sp. nov.
(Plate 32, fig. 6)
Material Examined
One complete, but damaged, exoskeleton (CPC 17069)
from 12 km south of Douglas Spring on the Morstone
to Undilla Road. It occurs in the V Creek Limestone
in the Ptychagnostus nathorsti Zone in association with
Asthenopsis levior and Papyriaspis lanceola.
Description
This specimen, being somewhat damaged
anteriorly and distorted by several cracks, cannot
be satisfactorily compared with other species on
cranidial features. Only those features that are at
variance with the description of A. levior above,
are referred to here.
It has a flatter border on the free cheek. The
genal spine is not deflected. The pygidium has
only a weakly convex pleural area and a poorly
defined border. The thorax has 16 segments, and
a tuberculate pleural ornament. The segments are
individually shorter than their corresponding
number in A. levior whether matching them (i.e.
14 against 14) from the anterior or posterior.
Remarks
Although this specimen could be included in A.
levior as representing a population well away from
the topotype population 1 believe that the
undcficcted genal spine alone, is sufficient to
separate it at the species level. In the absence of
other material I am reluctant to erect a new name
but am confident further collecting will necessitate
the assignment of one.
Distinguishing features of the five new northern
Australian species described herein are outlined in
Table 1.
LITERATURE CITED
Campbell. K. S. W., (in press). Trilobiles of the
Haragan, Bois d'Arc and Frisco Formations. Bull.
Okla geol. Surv.
CiSNE. J. L., 1974 Trilobites and the origin of the
arthropods. Science 186: 13-18.
Hfssler, R. R., 1964. The Cephalocarida. Comparative
skeletomusculature. Mem. Conn. Acad. Arts Sci.
16: 1-97.
Jell, P. A., 1975, Australian Middle Cambrian
codiscoids with a review of the superfamily.
Palaontographica A 150: 1-97.
Kobayashi. T., 1935. The Cambro-Ordovician forma-
tions and faunas of south Chosen. Palaeontology.
Part 3. Cambrian faunas of south Chosen with a
special study on Cambrian trilobite genera and
families. J. Fac. Sci. Tokyo Univ. (sect. 2) 4:
49-344.
Moore, R. C., Editor. 1959. ‘Treatise on Invertebrate
Paleontology’. Part O, Arthropoda 1 (Geological
Society of American and University of Kansas:
Boulder and Lawrence).
Opik, a. A., 1967. The Mindyallan fauna of
north-western Queensland. Bull. Bur. Miner.
Resour. Geol. Geophys. Aust. 74: 1-404.
Richter, R. and Richter, E., 1940. Die Sankianda —
Stufe von Andalusien, eine fremde Fauna im
europaischen Ober-Kambrium. Abh. Senkenberg
Naturf. Ges. 450: 1-88.
Wai ( on . C. D., 191 1. The Cambrian faunas of China.
Smithson misc. Colins 57: 69-109.
Whitehouse, F. W., 1939. The Cambrian faunas of
north-eastern Australia. Mem. Qd Mus 11*
179-282.
TABLE 1: Distinguishing Features of Species of Asthenupsis
JELL: MIDDLE CAMBRAIN TRILOBITA
73
is
(X
"O
M
u
73
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J3
—
OiD
o
CQ
s
ffi
231
MEMOIRS OF THE QUEENSLAND MUSEUM
Platf 31
Asthenopsis levior Whitehouse, 1939
(1, 4, 6, 7, 8 from the type locality on V Creek)
Fig. 1: Thorax and pygidium, UQF3340, (figured by Whitehouse,
1939, pi. 22, fig. 20), x 1-2.
Fig. 2: Latex cast of damaged early holaspid exoskeleton, CPC 17064,
X 6, from M4L
Fig. 3: Early holaspid cranidium, UQF68711, x 10, from
UQL3510.
Fig 4: Pygidium, UQF3339, (figured by Whitehouse, 1939, pi. 22,
fig. 19), X 2.
Fig. 5; Damaged cranidium, UQF68712, x 4-5; b, anterior oblique
view, from UQL3510.
Fig 6: Damaged cranidium, UQF3338. (figured by Whitehouse, 1939,
pi. 22, fig. 18), X 2.
Fig. 7: Latex cast of slightly damaged Holotype exoskeleton,
UQF3337, (figured by Whitehouse, 1939, pi. 22, fig. 17), x 2-5;
a, lateral oblique view, b, dorsal view.
Fig. 8: Large slightly damaged complete exoskeleton, UQF44314, x
1-5.
Fig. 9: Ventral view of left free check, CPC17065, x 3 from
M139.
Illustrations are dorsal views unless otherwise stated.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 31
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi ATI- 32
Asthenopsis levior Whitehouse, 1939
Fig. 1: Damaged cranidium, CPC 17066, x 2-5, from M247.
Fig. 2: Cranidium, UQF68713, x 1-5, from UQL469.
Fig. 3: Damaged pygidium exhibiting well developed caeca passing into
the border, UQF68714, a, x 2-5, b, x 7, from UQL469.
Fig. 4; Cranidium, CPC 1 7067, x 3, from Douglas Creek halfway from
the Old Burketown Road to the O'Shanassy River.
Fig. 5: Cranidium, CPC 17068, x 3, same locality as Fig. 4.
Asthenopsis sp. nov.
Fig. 6; Damaged exoskeleton, CPC 1 7069, x 2-2, from 12 km south of
Douglas Creek on the Morstone to Undilla Road.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 32
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 33
Asthenopsis queenslandica sp. nov.
(All material is from M243)
Fig. 1: Cranidiiim, CPC17070, a, x 4. b, x 6-5.
Fig 2; Small cranidium, CPCI7071, x 9.
Fig. 3: Right free cheek, CPC17072, x 3.
Fig. 4: Small cranidium, CPC17073. x 7.
Fig. 5: Holotype exoskeleton, CPC 17074, x 3.
Fig. 6: Large damaged exoskeleton without free cheeks, CPC 17075,
X 2-5.
Fig. 7: Large damaged exoskeleton without free cheeks, CPC 17076,
X 5.
Fig. 8: Pygidium, CPC17077, x 6-5.
JELL; MIDDLE CAMBRAIN TRILOBITA
Plate 33
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 34
Asihenopsis queenslandica sp. nov.
Fig. 1: Cranidium, CPC17078, x 4-5, a, anterior view, from
M243.
Asthenopsis rhinostrongyla sp. nov.
(All material is from HI 38)
Fici. 2: Small cranidium, CPC17079, x 5-5.
Fig. 3: Hololype cranidium, CPC 1 7080, x 3-5, a, anterior view.
Fig. 4: Ventral view of right free check, CPC19081, x 4-5.
Fig. 5: Thoracic fragment, CPC17082, x 2-5, b, lateral view.
Fig. 6: Latex cast of cranidium, CPC17083, a, x 4-2, b, lateral oblique
view, X 4.
Fig 7: Left free cheek, CPC17084, x 4.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 34
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATH 35
Asthenopsis rhinostrongyla sp. nov.
Fi(i, 1; Left free cheek, CPC 17085, x 5, b, lateral view, from
HI 38.
FKi, 2; Pygidium, CPC17086, x 6, from H138.
Asihenopsis unquinsepta sp. nov.
(All material is from Ml 57)
Fig. 3: Pygidium, CPC17087, x 3.
Fki. 4: Left free check, CPC 17088, x 4.
Fig. 5: Holotype cranidium, CPC 17089. x 3, a, lateral view.
Fig. 6: Two small individuals slightly damaged, CPC 17090, x 3.
Fig. 7: Latex cast of left free cheek, CPC 17091, x 4.
Fig. 8: Cranidium and thorax, CPC17092, x 2-5.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 35
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 36
Asthenopsis butorosa sp. nov.
Fig. 1: Latex cast of complete (less free cheeks) early holaspide,
ANU30645, x 5. from ANUI0325.
Fig 2: Pygidium, CPC17093, x 3-5, from M161.
Fig, 3: Small cranidium, CPC17094, x 7, b, lateral oblique view, from
from Ml 61.
Fig. 4: Latex cast of pygidium and damaged thorax, ANU30646, x
3, from ANU 10325.
Fig. 5; Cranidium, CPC 17095, a, lateral oblique view, x 4, b, dorsal
view, X 4, c, anterior oblique view, x 5, from M161.
Fig. 6; Right free cheek, ANU30647, x 5, from ANU10325.
Fig. 7: Latex cast of pygidium, ANU30648, x 5, from
ANU10325.
Fig. 8; Holotype cranidium, ANU30649, x 2-8, from ANU10325.
Fig. 9: Cranidium, CPC17096, x 3, from M161.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 36
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 37
Asthenopsis opalensis sp. nov.
(All material is from MNF15)
Fig. 1: Cranidium. CPC17097, x 3.
Fig, 2: Holotype cranidium, CPC17098, a, anterior view, x 3, b,
4.
Fig. 3: Cranidium, CPC17099, x 4, a, anterior oblique view.
Fig. 4: Cranidium, CPC17100, a, x 5-5, b, x 6, c, x 2*5.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 37
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 38
Asthenopsis opalensis sp. nov.
Fig. 1: Cranidium. CPCI7101, x 3, from MNF15.
Fig. 2; Pygidium, CPC17102, x 4-5, from MNF15.
Asthenopsis sp. cf. A. rhinostrongyla sp. nov.
Fig, 3: Cranidium (mostly exfoliated), CPC17103, x 3, a, lateral view,
b, from MI 56.
Fig. 4: Right free cheek, CPC17104, x 4-5, from M156.
Fig. 5: Pygidium, UQF68715, x 5, from UQL447.
Fig. 6: Cranidium, CPC17105, x 3, a, anterior view.
Fig. 7; Cranidium, UQF68716, x 4-5, b, anterior oblique view.
JELL: MIDDLE CAMBRAIN TRILOBITA
Plate 38
Mem. QdMus. 18(2): 233-43, pi. 39. [1978]
PARANCHISTUS PYCNODONTAE SP. NOV., A NEW PONTONIINE SHRIMP
ASSOCIATED WITH AN OSTREID BIVALVE HOST
A. J. Bruce
Heron Island Research Station, Gladstone
ABSTRACT
Paranchistus pycnodontae (Crustacea, Decapoda, Pontoniinae), a new species of shrimp from
Heron Island, Queensland, is described and illustrated. The species is most closely related
to P. spondylis Suzuki, from which it is distinguished. It is the second species of this genus
to be found in Australian waters and the first to be found in association with an ostreid host,
Pycnodonta hyotis L.
The genus Paranchistus was first designated by
Holthuis (1952), and at present includes four
species, all of which are now known to be
associates of large marine bivalve molluscs. Of
these four species, only one, Paranchistus
armatus (H. Milne-Edwards), has so far been
recorded from Australian waters. This species is
an associate of the giant clam, Tridacna gigas
(L.), and has been reported from Undine Reef,
Cape Tribulation, and from Chapman Island,
Queensland (McNeill 1968, Bruce 1975). Recently
a single example of another species of this genus
was found in association with an ostreid bivalve,
and the specimen is now described here as new
species.
Paranchistus pycnodontae sp. nov.
(Figs. 1-5, Plate 39)
Material Examined
HolotypE; Queensland Museum W7337, 5 , non-
ovigerous; Heron Island, Capricorn Group, Queensland;
No. 2442, 3 m, central lagoon, 17 August 1976, coll.
D. Fisk.
Description
A medium sized pontoniine of moderately
slender build. Carapace smooth. Rostrum well
developed, strongly compressed, with feebly
developed carina and extending anteriorly to
middle of intermediate segment of antennular
peduncle; rostrum horizontal, dorsal and ventral
margins straight and subparallel, tapering to an
acute tip distally; distal third of upper border with
five small subequal, acute teeth, with numerous
setae in interspaces; ventral border sparsely setose
with a single very small acute tooth below third
dorsal tooth. Orbit feebly developed, supraorbital
spines absent; inferior orbital angle slightly
produced, broad in dorsal view. Antennal spine
small, slender, marginal, not exceeding inferior
orbital angle. Hepatic spine distinctly projecting,
more slender than antennal spine and lower and
more posterior, in a small fossa and mobile.
Anterolateral angle of carapace bluntly
rectangular.
Abdominal segments smooth. Third segment
not produced posteriorly in dorsal midline. Length
of sixth segment 013 x depth, 015 x length of
fifth segment. Pleura 1-3 only slightly expanded,
4-5 smaller, bluntly rounded posteriorly, 6 with
posteroventral angle blunt, posterolateral angle
slightly more acute.
Telson c. 1-5 x length of sixth abdominal
segment, 2-2 x longer than broad; lateral margins
almost straight, converging posteriorly; anterior
width c. 2-7 X width at level of lateral pair of
posterior spines. Two pairs of small, subequal,
submarginal dorsal spines at 0-6 and 0-7 of telson
length. Posterior telson margin rounded, without
median point, with three pairs of spines; lateral
pair small, similar to dorsal spines, situated on
dorsal surface of telson, slightly in advance of
posterior margin; intermediate pair very stout, c.
0-14 X telson length, with attenuated distal ends,
submedian pair uniformly tapering, setulose, c.
0-75 X intermediate spine length.
234
MEMOIRS OF THE QUEENSLAND MUSEUM
Eyes normal, cornea globular, hemispherical,
slightly oblique, with distinct accessory spot;
podophthalmite slightly flattened, c. 1-2 x longer
than broad, distinctly wider than diameter of
cornea.
Antennules normal; peduncle exceeds rostrum
by most of intermediate segment; proximal
segment c. 1-3 x longer than proximal width,
medial border straight, without ventral spine;
styiocerite slender, acute, exceeding half segment
length; lateral border feebly convex, medially
convergent, with very short subrectangular
distolateral spine; anterolateral margin strongly
produced, distinctly angulated; intermediate and
distal segments subequal, obliquely articulated, c.
0-7 X length of proximal segment. Upper
flagellum short, biramous, proximal three
segments fused; shorter free ramus with two stout
segments, longer with nine slender segments; four
groups of aesthetascs. Lower flagellum slender,
sixteen segments.
Antenna with robust basicerite, with feeble
distolateral tooth; ischiocerite, merocerite
normal; carpocerite slender, exceeding middle of
scaphocerite, c. 3-2 x longer than wide. Sca-
phocerite well developed, extending well beyond
antennular peduncle: lateral border very feebly
convex, with acute distolateral spine, not
exceeding anterior margin of lamina. Lamina
broad, c. 2-2 x longer than width, anterior
margin broad, slightly truncate. Flagellum short,
slender, c 2 x postorbital carapace length.
Mouthparts generally similar to other species of
Paranchistus. Mandibles moderately robust,
without palp; molar process stout, with several
large blunt distal teeth, incisor process slender,
three acute teeth distally, central tooth
smallest.
Maxillula normal; palp with feebly developed
lateral lobe, medial lobe normal with small hooked
terminal seta; upper and lower lacinia broad,
densely setose with strongly setulose seta; short,
stout, feebly dentate spines distally on upper
lacinia, spiniform setae on lower.
Maxilla with stout, proximally swollen palp,
medial and lateral borders setose, distally slender,
non-setose. Basal endite well developed, broad,
deeply cleft, each lobe with numerous long,
slender, setulose setae. Scaphognathite broad, c.
2 6 X longer than width; anterior lobe broad.
Fig. 1: Paranchistus pycnodontae sp. nov., holotype female, Heron Island, Queensland. Scale in mm.
BRUCE: A NEW PONTONIINE SHRIMP
235
Fig 2’ PcirQfichistus pycnodontQ^ sp. nov., holotype: A, anterior carapace, rostrum, and antennae, dorsal aspect,
B, anterior carapace and rostrum, lateral; C, orbital region, dorsal; D, distal rostrum; E, inferior orbital angle
and antennal spine; F, eye, dorsal; G, antennular peduncle; I, antenna; J, distolateral tooth of scaphocerite; K,
telson; L, posterior telson spines; M, uropod; N, distolateral spine of exopod of uropod,
236
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig. 3: Paranchistus pycnodontae sp. nov., holotype: A, mandible; B, maxillula; C, maxilla; D, first maxilliped-
h, second maxilliped; F, third maxilliped; ^
BRUCE: A NEW PONTONIINE SHRIMP
237
20mm
C
0-5m m
BF
0-25 mm
HI
Fig. 4: Paranchistus pycnodontae sp. nov., holotype: A, first pereiopod; B, chela of first pereiopod; C, second
pereiopod; D, chela of second pereiopod; E, fingers of chela; F, fixed finger of chela; G, third pereiopod; H,
distal propod and dactyl of third pereiopod, lateral; I, idem, dorsal aspect of dactylus; J, accessory dactylar
tooth of fourth pereiopod;
238
MEMOIRS OF THE QUEENSLAND MUSEUM
First maxilliped with subcylindrical palp,
distomcdially with setulose setae, exceeding
anterior margin of endite but not caridean lobe;
basal endite broad, rounded, medial border
straight, confluent with coxal endite, junction
indicated by minute protuberance, with numerous
slender finely setulose setae; dorsal surface with
oblique row of long, coarsely setulose setae; coxal
endite sparsely setose. Exopod well developed,
flagellum with four plumose distal setae only,
caridean lobe small, broad. Epipod triangular,
deeply bilobed.
Second maxilliped normal; dactylar segment c.
3 X longer than broad, numerous slender, finely
setulose setae medially. Exopod with four
plumose setae distally; epipod small, irregularly
subrectangular; podobranch absent.
Third maxilliped moderately slender, reaching
proximal end of carpocerite; ischiomerus feebly
separated from basis, junction indicated medially
by small notch: ischiomerus slightly tapering
distally, moderately bowed, r. 4 x longer than
proximal width, medial border feebly setose, short
simple setae; penultimate segment c. 0-6 x
ischiomerus, 3-2 x longer than width, with longer
stouter setae medially and laterally; distal segment
similarly setose, c. 0-5 x ischiomerus, tapering, c.
3-5 X width. Basis broad, sparsely setose medially,
as long as wide: coxa stout, medially rounded, well
developed epipod laterally. Exopod slightly
exceeds ischiomerus, four plumose distal setae.
Arthrobranch rudimentary.
Coxae of third maxillipeds widely separated;
fourth thoracic slernite broad, unarmed.
First pereiopods slender proximally, stouter
distally, exceeding carpocerite by carpus and
chela; chela robust, palm subcylindrical, slightly
compressed; fingers subequal to palm, broadly
subspatulate, distally rounded with fine denticula-
tions along distal third of cutting edge medially
and two thirds laterally, slightly gaping
proximally on medial side, densely setose. Palm
r. 1-7 X longer than width, transverse rows of
cleaning setae proximally. Carpus proximally
slender, width increasing x 3 distally, r. 5 x
longer than distal width, cleaning setae dis-
tomedialiy; mcrus subequal to carpus, slightly
bowed, c. 6-4 x longer than width; ischium
compressed, c. 2-6 x width, c. 6-5 x merus,
medial border with row of short setae; basis 0-33 x
merus, sparsely setose along medial margin, coxa
robust, small setose ventromedian process
present.
Second pereiopods well developed, slender,
subequal, similar. Palm of chela subcylindrical,
smooth, slightly compressed distally, c. 3-4 x
width; fingers well developed, c. 0-6 of palm
length, dactylus strongly curved, overreaching
fixed finger, tip acutely hooked, cutting edge
sharp, entire except for small tooth at 0-3 of
length, outer surface with numerous short, erect
setae: fixed finger similar, proximal half of cutting
edge with eight small teeth, distal four subacute,
proximal four rounded, distal part of cutting edge
feebly denticulate. Carpus c. 0-47 x palm length,
2 X longer than distal width; distal margins feebly
excavate, unarmed. Merus c. 0*75 x palm length,
4 X longer than width, uniform, with small
distovenlral tooth. Basis and coxa normal.
Ambulatory pereiopods slender, third extending
beyond scaphocerite by 0-3 of carpus and chela;
dactyl about 0-27 x propod length; unguis distinct
from corpus, dorsoventrally compressed, hastate,
surface covered with dense felt of short processes
giving furry appearance; corpus w'ith single well
developed, slender acute accessory spine; short
distal sensory setae present. Propod c. 6-6 x longer
than width, uniform, with pair of distoventral
spines and single preterminal ventral spine.
Carpus 0-63 propod length, c. 3-2 x longer than
distal width, unarmed; merus c. 1-27 x propod
length; ischium c. 1-0 x propod length, c. 5 x
longer than distal width; basis and coxa robust,
all unarmed. Pereiopods 4-5 similar: 3, 5 subequal,
4 slightly shorter. Accessory spines on dactyls of
pereiopods 4. 5, with small additional denticle on
posterior margin.
Uropods slightly exceed telson tip, protopodite
with large acute posterolateral tooth; exopod
broad, c\ 2-3 x longer than wide, broadly rounded
distally, with small mobile spine only at posterior
end of convex lateral border; endopod c. 3 x
longer than width.
MEASUREMENTS: (In millemetres)
Total length (approx.) 18.7
Rostrum and carapace 6.2
Postorbital carapace 4.5
Second pereiopod chelae 3.8 (left and right)
COLOURATION: Semi-lransparenl with numer-
ous small round evenly distributed red chromato-
phores, all over body, ambulatory pereiopods,
antennae and caudal fan. Similar but slightly
larger chromatophores over second pereiopods.
Chromatophores absent from antennal flagella
and propods of ambulatory pereiopods.
HOST; Pycnodonta hyotis L. (Ostreidea)
ASSOCIATES: One speeimen of Onuxoden
parvibrachium (Fowler) (Pisces: Carapidae), a
species not previously known from Australia.
BRUCE: A NEW PONTONIINE SHRm?
239
TABLE 1: Hosts oi Si‘i(iis oi P\r\\(Mishs wn omiir Pomomim. Assoc iahs
Paranchistus spp.
Hosts
Other associates
P. armatus (H. Milne-
Edwards)
Tridacna gigas (L.)
Anchistus niiersi (De Man)
Conchodytes tridacnae Peters
P. ornatus Holthuis
Atrina vexillum Born
Anchistus custos (Forskal)
Conchodytes hiunguiculatus (Paulson)
P. nohilii Holthuis
Spondylus gaederopus L.
—
P. spondylis Suzuki
Spondylus harbatus Reeve
—
P. pycnodontae sp.nov.
Pycnodonta hyotis L.
Platypontonia hyotis Hipeau-Jacquotte
Kig. 5: Paranchistus pycnodontae sp. nov., holotype,
colour pattern.
Systematic Position of P. pycnodontae
Paranchistus pycnodontae appears to be most
closely related to P. spondylis Suzuki, and may
be distinguished from that species by the
sub-spatulate fingers of the first pereiopods. In all
other features the morphology of the two species
appears to be very similar. Other minor
differences include: In P. spondylis (a) the palm
of the chela of the second pereiopods appears to
be stouter; (b) the accessory spines of the
ambulatory pereiopods are shorter and stouter; (c)
the accessory spines of the dactyls of the fourth
and fifth pereiopods are without denticles; (d) the
minute processes on the anterior aspect of the
dactyls of the walking legs are perpendicular to
the surface; (e) the lateral posterior telson spines
appear to be marginal; (f) and the intermc^diate
posterior telson spines arc of uniform taper.
DISCUSSION
All species of the genus Paranchistus Holthuis
are known to be associates of bivalve molluscs.
The genus is not represented outside tropical
Indo-West Pacific waters. The host molluscs
belong to the families Tridacnidae, Pinnidae,
Spondylidae and now also include the Oslrcidae.
P. pycnodontae is the only shrimp so far known
to associate with Pycnodonta hyotis in Australian
waters, but in the western Indian Ocean
Hipcau-Jacquotte (1971) has recorded the
occurrence of a different pontoniine shrimp,
Platypontonia hyotis Hipcau-Jacquotte, which is
also known to associate with Pterostrea imhricata
(Lam.) in Japanese waters (Suzuki 1971). The
only other ostreid bivalve known to have a
pontoniine associate is Lopha cristagalli (L.), the
host of Platypontonia hrevirostris (Miers),
(Bruce 1968).
The hosts of the species of Paranchistus are
summarized in Table 1, which also indicates other
pontoniine associates of the same host.
240
MEMOIRS OF THE QUEENSLAND MUSEUM
In addition to the morphological characters that
distinguish P. pycnodontae from P. spondylis, it
may be noted that the colour patterns also appear
to be distinctive. In the figures of the latter given
by Suzuki (1971), the specimen shows a
comparatively coarse pattern of larger and more
sparsely distributed red chromatophores than is
found in P. pycnodontae. Most of the pontoniine
associates of the bivalve molluscs present a colour
pattern of uniformly distributed chromatophores
over most of the body surface and appendages.
These may be very small and numerous, as in
Anchistus custos, Paranchistiis ornatus or
Conchodytes tridacnae. In P. spondylis and P.
pycnodontae they are larger and more numerous,
and in Anchistus demani and A. miersi, they are
conspicuously larger and much less numerous. In
complete contrast, the colour pattern of
Platypontonia hyotis consists mainly of large
spots and bands of red.
The dactyls of the ambulatory pereiopods are
of particular interest as, in the genus
Paranchistus. they exhibit a wide range of
morphological variations of great value in
distinguishing the species. P. arniatus presents a
simple condition, with a simple, well developed
unguis distally and a broad accessory tooth. There
is no trace of ornamentation on the unguis (Bruce
1975). In P. ornatus the unguis is still distinct
from the corpus and the outer surface is covered
with transverse rows of short spines which become
obsolescent towards the tip. The ventral aspect of
the unguis is also transversely ridged. The corpus
has distal sensory setae present, but the accessory
spine is absent, the region of the spine being
provided with a low eminence covered with small
tubercles. The pit described by Holthuis (1952)
appears to be a vesicle below this eminence. In
P. pycnodontae, the arrangement is essentially
similar except that a well developed accessory
spine is present and the unguis is very much
larger. A comparable vesicular structure and
sensory setae are also present. P. nohilii and P.
spondylis have dactyls very similar to P.
pycnodontae.
Fujino (1975) has provided scanning electron
photomicrographs of the dactyl of the walking legs
in Anchistus miersi (De Man). Anchistus
Borradaile is a genus very closely related to
Paranchistus Holthuis, from which it is separated
principally by the complete absence of an hepatic
spine. In Anchistus a comparable range of
variation in dactylar morphology is also present
and in species such as A. demani Kemp and A.
miersi, with A. gravieri Kemp representing a less
well developed stage, and the dactyls very closely
resemble those found in some Paranchistus spp.
SEM photomicrographs clearly show that the
outer surface of the unguis of the dactyls is
densely covered with large numbers of short erect
spinules. In Anchistus the anterior aspect of the
unguis has been reported as scoop-shaped (Kemp
1922; Fujino 1975) but in Paranchistus spp. this
surface appears to be convex rather than concave.
The precise function of the spinulalions of the
unguis is still obscure, but is presumably related
to the grip of the shrimp upon its host. The
function of the vcntrally situated tooth on the
corpus would appear to enable a good grip upon
the host's tissues to be obtained during traction
by the limb. The distally directed spinules of the
unguis may function by preventing the penetration
of the dactyl too far into the host's tissues and
thereby avoiding excessive damage to the host in
a commensal relationship.
ACKNOWLEDGMENT
I am grateful to Dr Paxton for the identification
of the associated fish.
LITERATURE CITED
Bruci;. a. J., 1968. Notes on some Indo-Pacific
Pontoniinae. XII. The re-cxamination of the types
of Pontonia brevirostris Miers, 1844, with the
designation of a new genus Platypontonia
(Decapoda, Natantia). Cru.staceana 15(3); 289-97,
figs. 1-3.
1975. Pontonia armata H. Milne-Edwards
(Decapoda Natantia, Pontoniinae), — a correction.
Crustaceana 29(1); 49-54, figs. 1-3.
Fi jino, T., 1975. Fine features of the dactylus of the
ambulatory pereiopods in a bivalve associated
shrimp, Anchistus miersi (De Man), under the
scanning electron microscope (Decapoda, Natantia.
Pontoniinae). Crustaceana, 29(3); 252-54, figs. 1,
pi. 1.
Hiprau-Jacqliottk. R., 1971- Notes de faunistique et
de biologic marines de Madagascar. V.
Platypontonia hyotis nov. sp., (Decapoda Natantia,
Pontoniinae). Crustaceana. 20(2): 125-40, figs.
1-7.
Holthlis, L. B., 1952. The Decapoda of the Siboga
Expedition. Part XL The Palaemonidae collected by
the Siboga and Snellius Expeditions with remarks
on other species. IF Subfamily Pontoniinae. Siboga
Exped. Mon., 39a***: 1-252, figs. I-IIO, tab. 1.
Kfmp. S., 1922. Notes on Crustacea Decapoda in the
Indian Museum. XV. Pontoniinae. Rec. Indian
Mus. 24: 113-288, figs. I-I05, pis. 3-9.
Mac Neill. F. A., 1968. Crustacea, Decapoda and
Stomatopoda. Great Barrier Reef Expedition.
1928-29. Sci. Rep. 1(1): 1-98, figs. 1-2, pis.
I -2.
Suzuki, H., 1971. On some commensal shrimps in the
western region of Sagami Bay. Researches Crust.
4: 1-25, figs. 1-12, pis. 1-3.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 39
Paranchistus pycnodontae sp. nov., holotype, in Pycnodonta hyotis L.
BRUCE: A NEW PONTONIINE SHRIMP
Plate 39
Mem. Qd Mus. 18(2): 243-63. [1978]
GASTROINTESTINAL NEMATODES FROM AQUATIC AUSTRALIAN SNAKES
H. 1 . JONES
Department of Parasitology, University of Queensland
ABSTRACT
Three new nematodes are described, and another three species are recorded from six species
of aquatic snake (Acrochordidae and Colubridae). Four nematodes are from genera which
predominantly parasitise vertebrates of other Phyla or Orders, but which inhabit a similar
aquatic environment to these snakes.
This paper is the first in a proposed series on
the gastrointestinal parasites of Australian snakes.
Little work has been done on the nematode
parasites of the Australian tropica! and sub-
tropical aquatic snakes, most of it being confined
to identifications and host records (Johnston and
Mawson 1941. 1948), with no indication of
intensity or prevalence of infection. This paper is
concerned with the gastrointestinal nematodes
from 26 Australian aquatic Colubrid and
Acrochordid snakes preserved in the Queensland
Museum, Brisbane. Representatives from all six
Australian species in this group were examined,
namely Acrochordids javanicus, Acrochordus
granulatus, Cerberus rhynchops, Myron ri-
chardsoni. Fordonia leucobalia and Enhydris
polylepis.
The nematodes recovered are as follows:
Subclass ADENOPHOREA
Order ENOPLIDA
Superfamily DIOCTOPHYM ATOIDEA
Eustrongylides acrochordi sp. nov.
(Fig. I, Table 1)
Material Examined
HolotypE: Queensland Museum G 1 0275, ^ , collected
by H. Jones from stomach of Acrochordus javanicus.
QM J2319I, collected by J. Covacevich and C. Tanner
near Coen, North Queensland, June 1973.
Paratype: QM G 1 0276, ^ , (incomplete), same data
as holotype.
Diagnosis
Double row of six apical papillae, marked
terminal striations and row of lateral punctations,
nerve ring anterior, oesophagus long and
voluminous, anus terminal. Vagina opens into the
rectum, forming a cloaca. No tail.
Description
Worm long, cylindrical and partly coiled. Very
marked striations near both ends, diminishing
until scarcely visible near centre of worm. Two
lateral rows of very small punctate markings,
situated in the striations, proceed posteriorly from
anterior end, disappearing within a few mm. Two
lateral rows of four evenly spaced small papillae
at caudal end, replaced anteriorly by punctate
markings as at anterior end, gradually disappear-
ing. Body of fairly uniform thickness, tapering
slightly at posterior end but more markedly at
anterior end. Mouth elongated dorsovcnlrally,
surrounded by six spined papillae, two laterally
and four submedially. Another row of six rounded
papillae below these. Another 12 much smaller
papillae, two just bclow' each lateral spined
papilla, one just anterior to each lateral rounded
papilla, and the remaining six at intervals between
the two main rows of papillae.
A short straight pharynx leads into a wide
undulating oesophagus; this increases slightly in
width throughout its length, is without a bulb, and
measures 0-2 x the total length of the worm. Nerve
ring surrounds oesophagus just behind its origin.
No excretory pore seen. A wide rectum, flattened
244
MEMOIRS OF THE QUEENSLAND MUSEUM
Fi(i 1. hustrongylides acrochordi sp. nov. Holotype ^ G10275, A, anterior end; B, anterior extremity, en face;
C, Anterior end and oesophagus; D, posterior end, dorsal; E, posterior end, lateral, showing insertion of vagina
into rectum.
JONES: NEMATODES FROM AQUATIC SNAKES
245
TABLE 1: Body Measurements (In mm) of Holotype
Immature Female Eustrongylides acrochordi sp.
nov.
Total length 84-50
Width at mid-point 103
Width at 2-0 mm from front 0*56
Head width at 1st row of papillae 015
Head width at 2nd row of papillae 0-22
Width at beginning of oesophagus 0-28
Oesophagus length 15-90
Oesophagus proportion of length 1/5
Oesophagus width near beginning 01 8
Oesophagus width c. 1 /4 along 0-28
Oesophagus width c. 3/4 along 0*42
Pharynx length 0-19
Length of anterior papilla spines 0*01
Nerve ring fr. ant. extremity 0*24
Nerve ring fr. oesoph. beginning 0 03
Caudal papillae distance 0 08
Punclation spacing at anterior end 0-04
Punctation spacing proceeding posteriad 0*13
Anus width 010
Rectum length 0-38
Rectum width 0*13
Post, intestine-rectum 1 00
Width of uterus 4 mm fr. tail 0-15
Width of uterus near rectum 0 06
Lumen of uterus 0*04
Uterine wall thickness 0*01
Distance of post, edge of vagina fr. anus 0*08
dorsoventrally, leads into the terminal anus. This
is widened laterally and is surrounded dorsally
and venlrally by a rugose ridge. There is no tail.
Ovary not seen. The vagina passes posteriorly and
just before the rounded posterior end of the worm
turns sharply medially and enters the rectum. The
uterus was not fully developed and contained no
eggs.
Discussion
Both the holotype and the paratype (which
lacked the anterior end) were lying freely in the
stomach among a large number of Tanqua
ophidis. The worms are characteristic of the
genus, but differ from all described species in that
the vagina opens into the rectum, forming a
cloaca. This feature is sufficient to ascribe them
to a new species, even though neither worm is
mature. In all other Eustrongylides species the
vulva opens terminally, very close to the anus. A
cloaca has been described from only one other
group of adenophorean nematodes, the free-living
Lauratonema species (Gerlach 1953).
All previous records of Eustrongylides species
have been from large aquatic birds, from the
proventriculus or a.ssocialed glands, and it is
possible that this snake was acting as a paratenic
host.
Subclass SECERNENTEA
Order OXYURIDA
Superfamily OXYUROIDEA
Spironoura fordoniae sp. nov.
(Fig. 2, Table 2)
Material Hxaminfi^
Holotype Queensland Museum G 10277, collected
by H. Jones from rectum of Fordonia leucobalia, QM
J23200, collected by J. BredI from Edward River, North
Queensland, 1973.
ALLOTYPES; QM G 10278, 2 ^ (poor condition),
same data as holotype.
ParatypeS: QM G10279, 3 ^ ^,1 55 , same data
as holotype; QM G 1 0280, 8 2 55 , (poor
condition), collected by H. Jones from stomach of F.
leucobalia. QM J23924, collected by J. Bredl from
Edward River, North Queensland, 1973.
Diagnosis
Head with three lips surrounded by six papillae.
Vestibule present. Pharynx short, oesophagus with
hour-glass formation at posterior end, stout equal
alatc spicules, conspicuous oblique precaudal
ventral muscle bands, ten pairs of caudal papillae,
one unpaired precloacal papilla, lateral alae from
cervical region to near posterior end, vulva just
past mid-length.
Description
Worm tapering towards either end. No
constriction behind head. Body finely striated.
Lateral alac commence in cervical region and run
most of the length of body, diminishing in size
and finally disappearing in last quarter of body.
Tail slender and finely pointed in both sexes.
Mouth with three lips, with two papillae at base
of each. Short vestibule leading into muscular
pharynx, length about 2-5 x width. Oesophagus
muscular, long, gradually increasing in diameter
to a prcbulbar swelling (preceded by a
constriction) and a large spherical muscular bulb.
Intestine straight and wide lumcned. Nerve ring
0-25-0-20 from front of oesophagus. Small but
distinct cervical papillae about 0-66 along
oesophagus, excretory pore about 0-75 along
oesophagus.
MALE: Length rather less than females. Caudal
end coiled venlrally. Oblique precaudal ventral
muscle bands conspicuous; no ventral sucking disc.
No caudal alae. Ten pairs of sessile caudal
papillae, arranged as three regularly spaced
ventrolateral precloacal papillae, three closely set
pairs of paracloacal papillae, and four pairs of
caudal papillae of which the most anterior two are
246
MEMOIRS OF THE QUEENSLAND MUSEUM
Eld 2. Spironoura fordoniae sp. nov. Holotype 3 G10277. A, anterior end; B, anterior extremity, en face; C,
anterior end and oesophagus. D, posterior end, ventral; E, posterior end, lateral; F-I, sections at level of upper
oesophagus, lower oesophagus, mid-body and near posterior end, respectively.
TABLE 2; BOD't Mlasi revients {IN mm) or Spironoura fordoniae. sp nov and Camallanides cerberi sp
JONES: NEMATODES FROM AQUATIC SNAKES
247
248
MEMOIRS OF THE QUEENSLAND MUSEUM
lateral and the most posterior two ventral. One
unpaired precloacal papilla. Spicules equal in
length, curved and stout, with wide alae reaching
almost to the bluntly pointed tips. Gubcrnaculum
chitinized and elongated, with a pointed posterior
end.
FEMALE: Vulva on a slight protrusion, just past
mid-length. Vagina directed anteriorly. No caudal
papillae. Eggs large, elongated, with thick shells,
unernbryonated.
Dicussion
These worms differ from all previously
described members of the genus in the possession
of lateral alae. As absence of alae is given as part
of the generic diagnosis (Yorke and Mapicstone
1926; Yamaguli 1961 ), this must now be amended
to read; alae present or absent.
The genus Spironoura is confined to the
digestive tract of fish, amphibia and reptiles.
However of the approximately 27 species which
have been described from reptiles (Yamaguti
1961) all but three arc from chelonians; only S.
rnascula (Rudolphi 1819) and S. nitida
(Travassos 1920) have been described from
snakes. The only previous Australian record of the
genus is S. elseyae Johnston and Mawson 1941
from the turtle Elseya dentata.
Order ASCARIDIDA
Superfamily ASCARIDOIDEA
Family ANISAKIDAE
Goezia sp.
Mati-riai Examinld
Queensland Museum G 10292, 2 collected by H.
Jones from stomach of Acrochordus granulatus, QM
J28740. collected by J. Covacevich and P. Filewood, Iron
Range, North Queensland, June 1976.
It was not possible to assign these to a species,
but they appeared to be the same as specimens
recently recovered from Australian sea-snakes and
crocodiles, to be described by Sprent (in press).
Apart Irom these, the only previous record of this
genus from a reptile has been G. gavialidis,
(Maplestone 1930), from an Indian CJavial. All
other species have been from fish.
Order SPIRURIDA
Suborder CAMALLANINA
Superfamily CAMMALLANOIDEA
Caniallaiiides cerberi sp. nov.
(Fig. 3, Table 2)
Material Examined
Hoi OTVnr Queensland Museum G 1 028 1, (3, collected
by H. Jones from upper oesophagus of Cerberus
rhynchops, QM J23630, collected by J. BredI, Edward
River, North Queensland, June 1973.
At l.OTYt’i-;: QM GI0282, j, same data as
holotypc.
ParatypeS: QM GI0283, 3 (5c?, 11 $5, and 9
incomplete specimens, same data as holotype; QM
G 1 0284, I , oesophagus of Enhydris polylepis, QM
J20282, collected by S. Sterling near Cairns, North
Queensland, 1970.
Diagnosis
Chitinised buccal valves with 12 14 internal
longitudinal ridges. Dorsal and ventral chitinous
rods projecting posteriorly from edge of buccal
capsule. Four small perioral papillae. Seven or
eight pairs of pedunculate precloacal papillae and
eight or nine pairs of postcloacal pedunculate
papillae in males. Unequal spicules. No guber-
naculum. Vulva on a pedunculate prominence;
female tail slender and ending in a slight knob.
Viviparous.
Description
Fixed worms pale grey in colour, with a black
streak running the entire length due to blood or
blood products in intestine. Fairly uniform width,
anterior end rounded, tail tapering. All specimens
were collapsed so diameter could not be measured
accurately. Head capsule chitinized, brown.
Cuticle finely striated. Mouth elongated dorso-
ventrally, with four inconspicuous sessile papillae,
one beside each corner. Buccal capsule consists of
two buccal valves, each of which consists of two
chitinous masses separated by a broad longitudin-
al groove. A smaller chitinous body anterior to
each mass, from the median aspect of which is
a small projection at the worm's anterior
extremity. Twelve to lourteen longitudinal ridges
inside each valve, only the central ones of which
continue to base of capsule. Chitinous ring at base
of buccal capsule. Thin curved chitinous bodies at
dorsal and ventral edges of capsule run through
ring at base of capsule to anterior end of
oesophagus; from exterior aspect of each an
irregularly-shaped chitinous bar extends
posteriorly.
Oesophagus divided into two portions; anterior
muscular portion stout, with pronounced swelling
towards posterior end. Posterior glandular portion
almost cylindrical in shape, widening slightly
towards posterior end, lumen narrow and tortuous.
Intestine broad lumened, filled with blood or
blood products, running directly to anus. Nerve
ring surrounds anterior portion of oesophagus,
JONES: NEMATODES FROM AQUATIC SNAKES
249
Fig. 3. Callamanides cerberi sp. nov. Hololype S G10281, Allotype $ G10282. A, anterior end, lateral; B, anterior
end, dorsal; C, en face; D, male tail, ventral; E, female tail, lateral; F, vulva, lateral; G, embryo in
uterus.
250
MEMOIRS OF THE QUEENSLAND MUSEUM
excretory pore near posterior end of muscular
oesophagus. Very small bristle-like cervical papilla
seen in one female only.
MALE: Less than half length of female. Well
developed caudal alae, terminating just before tip
of pointed tail. Alae connected anteriorly (not in
all specimens), supported by an inconstant number
of papillae; six to nine regularly-spaced pedun-
culate papillae precloacally, and eight or nine
smaller irregularly spaced papillae between cloaca
and tip of tail. Spicules unequal in size and
dissimilar in shape. Larger right spicule (projects
from cloaca in all specimens) alate, tapering to
fine slightly curved point. Smaller left spicule not
alate, but also with a finely curved tip. No
gubernaculum.
FEMALE: Mean length 27 mm. Tail long and
tapering, terminating in an almost club-shaped
knob, with or without a conical tip. Vulva situated
just anterior to midpoint of body on rounded
pedunculate appendage directly posteriorly.
Vulval aperture on dorsal side of appendage,
against body of worm, surrounded by two small
lips. Muscular vagina runs anteriorly some
distance before opening into opposed uterine
tubes. All specimens contained many larvae, many
of which contained numerous black granules,
indicating that they were obtaining nutriment
from the females’ blood intake.
Discussion
Five species of Camallanides are recognised at
present, all from the Indian subcontinent; C
prashadi Baylis and Daubney 1922, C. piscatori
Khera 1954, C. ptyasi Khera 1954, C. dhamini
Deshmukh 1968, and C. hemidenta Majumdar
1965. C. prashadi was recovered from Naja
bungarus and Ptyas mucosus, as well as from
Bungarus fasciatus by Baylis (1929), and from
a frog, Rana tigrina by Karve (1930), C. piscatori
from Natrix piscator, C. ptyasi and C. dhamini
from Ptyas mucosus and C. hemidentata from a
freshwater fish, Channa striatus. In addition,
Gupta (1959) recorded a single unidentified
female Camallanides from a sea-snake,
Hydrophis cyanocinctus. All were found in the
intestine of their hosts. The five snake species
from which these are recorded are not found in
Australia, but their distribution overlaps with both
that of Fordonia leucobalia and Cerberus
rhynchops in the Indo-Malaysian archipelago. C.
prashadi has been identified from C. rhynchops
from Thailand (British Museum, unpublished)
Camallinides cerberi differs from those species
described in being larger, in having four instead
of six apical papillae, in the absence of a
gubernaculum, and in the rounded anterior end.
In addition, it differs from C. prashadi in the
shape of the dorsal and ventral chitinous bodies
which do not project forwards as in that species,
in the thicker post-directed chitinous rods, and in
the wider groove separating the chitinous masses
of each valve. W'hcther or not C. piscatori and
C. ptyasi have dorsal and ventral chitinous bodies
seems uncertain (Deshmukh 1968). C. cerberi
differs from C. prashadi and C. dhamini in having
a relatively shorter and more bulbous vulval
appendage; the vulval aperture is on the doral side
of this appendage, and not on the ventral side as
in C. prashadi. C. ptyasi and C. piscatori. In C.
dhamini there are 14-16 longitudinal internal
buccal ridges, and less inequality in spicule length.
These specimens are assigned to a new species on
the basis of these various features. Differences in
the number and position of the caudal papillae,
however, have not been taken into account; they
have been used to some extent in differentiating
species in this genus but, at least in C. cerberi,
they are variable. C. piscatori and C. dhamini
were described from one male, C. ptyasi. C.
hemidenta, and the redescription of C. prashadi
(by Agrawal 1967) were from two males. In view
of this, and the fact that three of these (C.
prashadi, C. ptyasi and C. dhamini) have been
described from the same host. Ptyas mucosus. it
seems desirable that a greater number of
specimens be examined before differentiation
based on these papillae can be relied upon.
Further study may question the validity of these
species.
Suborder SPIRURINA
Superfamily GNATHOSTOMATOIDEA
Tanqua ophidis Johnston and Mawson 1948
Material Examined
Queensland Museum G 10285, G 1 0286, G 10287,
G 1 0288 and G 10293 from Acrochordus javanicus QM
J23192, J23718, J23189, J23I91 and J23718 respective-
ly (see Table 4); 3 6 S and 1 incomplete specimen in
author’s collection, stomach of Enhydris polylepis.
J23215.
Apart from two in the oesophagus and two in
the intestine, all specimens were in the stomach,
in numbers ranging from seven to 179. Sexes were
approximately equal, with a large number of
young worms in the two most heavily infected
JONES: NEMATODES FROM AQUATIC SNAKES
251
Fig. 4. Heliconema longissima (Ortlepp 1922), G 10291. A, anterior end, lateral; B, male tail, ventral; C, right
spicule; D, left spicule.
252
MEMOIRS OF THE QUEENSLAND MUSEUM
snakes; clearly it is a prevalent parasite in this
species. It was originally described from a
freshwater snake, Amphiesma mairii, and from A.
javanicus (see Johnston and Mawson 1948), and
has not been reported since then. E. polylepis is
therefore a new host record.
Superfamily PH YSALOPTEROIDEA
Heliconema longissima (Ortlepp 1922)
(Fig. 4, Table 3)
Material Examined
Queensland Museum G10291. 37 stomach, 3
oesophagus, 1 rectum Fordonia leucohalia; J 10262;
G10290, 12 stomach and 11 proximal small intestine,
F. teucobalia. J877; G 10289, one from stomach, F.
leucohalia. .123200.
The characters and measurements of the worms
agree in almost every particular with the original
description by Ortlepp (1922) and the redescrip-
tion by Ogden (1969), the only differences being
that in these specimens there were never more
than nine longitudinal tesselated ridges on the
ventral surface of the male caudal region, instead
of 12 in Ogden’s description, and in the females
the vulva was always just posterior to the
midlength of the body.
Discussion
In the measurements of this species by Ortlepp
(1922), Li (1934) from Anguilla pekinensis from
China and Ogden (1969), the vulva was always
slightly anterior to the midlength of the body.
These differences, however, together with the
lesser number of tesselated ridges in the male, are
not thought sufficient to warrant a new species,
and may be host-induced variations.
There has been uncertainty about the hosts of
the original material examined by Ortlepp (1922),
which had been labelled ‘snakes, Australia’.
Chabaud and Campana-Rouget (1956) in
suggesting that the genus Orlleppina erected for
the type species by Schultz (1927) was
synonymous with Heliconema, suggested that the
original specimens had been wrongly labelled, and
that they had probably come, as had all other
known members of the genus, from eels, and this
view was supported by Ogden (1969). The finding
of these specimens in aquatic snakes now suggests
that maybe the original labelling was correct. The
present specimens are therefore the first certain
records of the genus which were not recovered
from eels.
DISCUSSION AND CONCLUSIONS
The nematode species recovered were:
Eustrongylides acrochordi sp. nov., Spironoura
fordoniae sp. nov., Goezia sp., Camallanides
cerberi sp. nov., Tanqua ophidis Johnston and
Mawson 1948 and Heliconema longissima
(Ortlepp 1922). In 24 of the 26 snakes the
stomachs were empty; two Fordonia leucohalia
TABLE 3; Body Measurements on mm) of Heliconema longissima
(ORTLEPP).
Males (4)
Females (11)
range x ± &
Length
17.85 -27.37
28.27 -38.55
32.19 ±3.19
Maximum width
0.192- 0.350
0.402- 0.542
0.499±0.069
Nerve ring*
0.231- 0.315
0.259- 0.350
0.298±0.026
Cervical papilla*
0.231- 0.315
0.245- 0.329
0.290±0.025
Excretory pore*
0.366- 0.413
0.406- 0.546
0.4I4±0.049
Muscular eosophagus
0.371- 0.497
0.455- 0.532
0.425±0.028
length
Glandular oesophagus
2.47 - 3.62
2.84 - 4.14
3.33 ±0.29
length
Muscular oesophagus
0.063- 0.105
0.070- 0.112
0.091±0.006
width
Glandular oesophagus
0.119- 0.189
0.140- 0.210
O.I78±O.OI4
width
Tail length
0.231- 0.350
0.091- 0.203
0.145±0.028
Spicule length, left
0.482- 0.616
—
—
Spicule length, right
0.231- 0.259
—
—
Vulva position!
50-66
56±2
* from anterior end
t % of body length from anterior end
Host Qld. Museum Locality and Date Tanqua Eustrongylides Camallanides Heliconema Spironoura Goezia
(Habitat and diet) Cat. No.
JONES: NEMATODES FROM AQUATIC SNAKES
253
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254
MEMOIRS OF THE QUEENSLAND MUSEUM
(J877 and J 10262) contained remains of large
Crustacea.
The differing habitats of the snakes probably
reflect the different composition of their food, and
hence their largely different parasites. Data in
Table 4 suggest that T. ophidis and E. acrochordi
probably have fresh-water life cycles, S.
fordoniae, Goezia sp. and H. longissima marine
life cycles, and C. cerheri an estuarine life
cycle.
These are the first records of snakes as hosts
for parasites in the genera Eustrongylides and
Heliconema. Goezia have recently been reported
from sea-snakes and crocodiles (Sprent, in press),
and the great majority of species in the genus
Spironoura are found in fish or chelonians. The
usual hosts for Spironoura spp. (fish and turtles),
Eustrongylides spp. (aquatic birds) and
Heliconema spp. (eels) are ones which live in
similar environments to these snakes, and with
which they may have common food and hence
sources of infection. The presence of these
parasites in snakes illustrates their physiological
opportunism in adapting to phylogenetically
different hosts which inhabit the same environ-
ment and may be involved in similar food-chains
to their more usual hosts. Both £. acrochordi and
S. fordoniae show morphological features not
found in other members of these genera (cloaca
in one; lateral alae in the other), suggesting that
they may be evolving away from the main pattern
of forms found in these genera.
ACKNOWLEDGMENTS
I would like to thank Miss. J. Covacevich for
her great help in allowing me access to and
facilities for the examination of the snakes in the
Queensland Museum, and to Professor J. F. A.
Sprent for helpful advice and discussion.
LITERATURE CITED
Agrawal. V., 1967. Some new Camallanoidea
(Spirurida) nematodes from fishes, amphibians and
reptiles. Annls. Parasit. hum. comp. 42: 327-42.
Baylis. H. a., 1929. On the Hosts of Camallanides
prashadi Baylis and Daubney 1922 (Nematoda).
Ann. Mag. nat. Hist. (10)4: 50.
Baylis. H. and Daubney R., 1922. Report on the
parasitic nematodes in the collection of the
Zoological Survey of India. Mem. Ind. Mus. 7:
263-347.
Chabuad. a. G. and Campana-Rouget. Y., 1956. Le
genre Ortleppina Schulz 1927, parasite d‘Apodes, et
non de Serpents, est Synonyme du genre
Heliconema Travassos 1919. Ann. Parasit. hum.
comp. 31: 308-9.
Deshmukh. P. G., 1968. Camallanides dhamini n.sp.
from Rat Snake Ptyas mucosus Riv. Parassit. 29:
119-22.
Gerlach, S. a. Von, 1953. Lauratonema nov. gen.
Vertreter einer neuen Familie mariner Nematoden
aus Dem Kustengrundwasser. Zool. Am. 151:
43-52.
Gupta, S. T., 1959. Nematode Parasites of Vertebrates
of East Pakistan. III. Camallanidae from fish,
amphibians and reptiles. Canad. J. Zool., 37:
771-9.
Johnston. T. H. and Mawson. P. M., 1941, The
Gallard collection of Parasitic Nematodes in the
Australian Museum. Rec. Aust. Mus. 21: 1 10-15.
Johntson. T. H. and Mawson. P.M., 1941-47. Some
parasitic nematodes in the collection of the
Australian Museum. Rec. Aust. Mus. 21: 9-16.
Johnston. T. H. and Mawson. P. M., 1948. Some new
records of Nematodes from Australian snakes. Rec. i
S. Aust. Mus. 9: 101-6.
Karve. j. N., 1930. Some parasitic nematodes of frogs !
and toads. Ann. Trop. Med. Parasit. 24: 481-91.
Khera. S., 1954. Nematode Parasites of some Indian
Vertebrates. Indian J. Helminth. 6: 27-133.
Li. H. C., 1934. Report on a collection of parasitic
nematodes mainly from North China pt. 2.
Spiruroidea. Trans. Am. Microsc. Soc. 53: 174-95.
Majumdar. N., 1965. Camallanides hemidenta sp.n.
(Nematoda: Camallanidae) occurring in Channa
striatus (Bloch). Zpol. Am. 174-175: 222-5.
Maplestone. P. a., 1930. Parasitic nematodes obtained '
from animals dying in the Calcutta Zoological
Gardens — Parts 1-3. Rec. Ind. Mus. 32: 385-412.
Ogden. C. G., 1969. A revision of the genus Heliconema
Travassos 1919, Physalopteridae (Nematoda). J.
nat. Hist. 3: 423-31.
Ortlepp, R. j., 1922. The Nematode Genus
Phvsaloplera Rud. Proc. Zool. Soc. London: j
999-1 107.
Rudolphi. C. a., 1819. ‘Entozoorum synopsis cui
accedunt mantissa duplex et indices locupletissimi”
81 1 pp. (Berolini).
Schulz. R., 1927. Die familie Physalopteridae Leiper
1908. Nematodes und die principen ihre Klassifica- j
tion. pp 287-312 In ‘Sammlung Helminth Arbeiten t
Prof. K. I. Skrjabin gewidment’. (Moscow).
Sprent. J. F. A. (in press) Synopsis of the Ascaridoidea:
Terranova, Goezia, and Paraheterotyphlum. J.
Helminth.
Travassos. L., 1920. Genero Florencioia Trav. 1919.
Arch. Esc. Sup. Agric. Med. Vet. 4: 21-4.
Yamaguti, S., 1961. ‘Systema Helminthum. HI The
Nematodes of Vertebrates.’ (Interscience Publishers
Inc.: New York).
Yorke, W. and Maplestone, P. A., 1926. The
Nematode Parasites of Vertebrates.’ (Hafner
Publishing Company: New York).
Mem. Qd Mus. 18(2): 255-63, pi. 40. [1978]
SOME PARASITIC BARNACLES (RHIZOCEPHALA: SACCULINIDAE) FROM
PORTUNID CRABS IN MORETON BAY, QUEENSLAND
W. J. Phillips
Department of Parasitology, University of Queensland
ABSTRACT
One new species of Sacculina and two new species of Heterosaccus are described and
additional morphological features of Sacculina granifera Boschma 1973 revealed by scanning
electron microscopy are recorded.
Only three sacculinids have been reported from
Australian waters. Boschma (1933) described
Sacculina duracina from Parthenope longimanus
(Leach) at Port Molle, Queensland, and more
recently S. granifera from Portunus pelagicus
(Linnaeus) from Moreton Bay, Queensland
(Boschma 1973). A further unnamed Sacculina
sp. from Thalamita sima H. Milne-Edwards from
Sydney was mentioned by Haswell (1888). S.
granifera was described only recently although it
is a serious parasite of the commercial sand crab
(P. pelagicus) and has been known for many years
(Thomson 1951). In the course of an investigation
into the biology of this association, S. granifera
and three other sacc-uiinids found on different host
species were examined primarily to determine
whether the other crab species served as reservoir
hosts of S. granifera. This paper contains
descriptions of three previously unnamed sacculin-
ids and further observations on the morphology
of S. granifera.
MATERIALS AND METHODS
Externae of the parasites were fixed in 5%
formalin, Bouin's Huid or 70% alcohol. For
sectioning, a small part of the mantle and parts
of the visceral mass containing colleteric glands,
cypris cell receptacles and ducts were removed,
dehydrated and embedded in wax. Sections were
cut at 7 - 10/^ and were stained in borax carmine
or chlorazol black E. Retinacula were observed by
mounting small pieces of mantle in balsam and
examining the inner surface. Pieces of mantle
taken from young, clean externae were prepared
for scanning electron microscopy by dehydration
in graded alcohols and xylol and coating with gold
dust. All measurements are in millimetres unless
otherwise stated, and follow the convention
illustrated in Fig. la where A = anterior to
posterior dimension, B = dorsal to ventral
dimension (The parasite lies on one side), and C
= the left to right dimension i.e. the thickness
of the parasite between the crabs abdomen and
cephalothorax. Abbreviations used in figures are
as follows:
col. gl.
colleteric gland
ex.
excrescences
ext. cut.
external cuticle of mantle
g. coat.
granular coating of mantle
j-
junction of male duct and
cypris cell receptacle
1. can.
lymph canals
l.c.r.
left cypris cell receptacle
l.m.d.
left male duct
mant.
mantle
me.
mesentery
m.o.
mantle opening
pap.
papillae
ped.
peduncle
r.e.r.
right cypris cell receptacle
r.m.d.
right male duct
rods
supporting rods of retinaculum
sept.
septae
vise. m.
visceral mass.
256
MEMOIRS OF THE QUEENSLAND MUSEUM
Sacculina amplituba nov. sp.
(Fig. la - j)
Material Examined
Hoitn't Ri Queensland Museum W7145, ex Matuta
granulosa, Main channel, Moreton Bay, collector W.
Phillips, 6. iv. 1973.
Par \t VEi s. W7146, a whole specimen, W7147, serial
sections, same data as holotype.
Diagnosis
Panduriform parasites occurring singly. Cypris
cell receptacles and ducts outside visceral mass,
completely separated, receptacles large, roughly
globular without septae passing abruptly into
ducts; left receptacle terminates on the right side
of the mesentery; diameters of ducts widen until
equal to that of receptacles, septate. Tubes of
colleteric glands arranged in approximately seven
rows, greatest division shows 150-160 tubes.
External cuticle covered with spiny excrescences
of 35// length formed of hyaline chiton differing
from that of the main layers. Retinacula not
present. Parasitic on Matuta granulosa Miers.
Description
Body form: Mature externae are ovoid to
rectangular in shape (Fig. la) and brownish in
colour except for a pale patch surrounding the
posterior stalk. Fine lines can be seen on the
mantle, the opening of which is anterior on top
of a short muscular tube. Although some
specimens were examined other than those
designated types, none w'cre found on crabs
bearing more than one externa. Dimensions of
holotype: A=15, B = 21,C = 8 and a whole
paratype A = 10, B = 15, C = 6.
Cypris cell receptacles; The cypris cell
receptables are in the anterior region of the stalk
completely separated from the viscera! mass (Fig.
lb,f-h). They are roughly globular and without
septae and open into the ducts by a narrow
aperture (Fig. le-g). The ducts widen till they
become as wide as the receptacles (Fig. Id) then
narrow near the opening into the mantle cavity.
The terminal portion is free in the mantle cavity
(Fig. Ic). The internal surface of the duct bears
septae (Fig. Id). The left cypris cell receptacle
terminates on the right side of the mesentery (Fig.
Ih).
Colleteric glands: The colleteric glands are
cushion shaped bodies in the middle portion of the
left and right sides of the visceral mass (Fig. lb).
In transverse section in the area of greatest
division of tubes, there are 1 50- 1 60 lubes in up
to seven rows. Fig. Ij shows 155 lubes.
Mantle: The external cuticle of the mantle is
75-100// in thickness and bears clusters of spines
35-40// in length. The spines are united at their
bases in clumps of two to seven; they are of a
hyaline chiton different from the cuticle of the
mantle (Fig. li). As with all saccuiinids, granular
matter collecting round the excrescences may
partially obscure them (Fig. li). No retinacula
were observed.
Remarks
The name Sacculina amplituba nov. sp. is
proposed because the male ducts widen to a
diameter equal to that of the receptacles. In
related species the ducts are narrower than the
receptacles. 5'. amplituba is similar to 5. beauforii
Boschma 1949 from Scylla serrata (Forsk.), S.
leptodiae Guirin-Ganivit from Xantho exeratus
(H. Milne-Edwards) and S. vankampeni Boschma
from Ozius rugulosus Stimpson, but differs from
these in having (i) ducts as wide as the receptacles
whose ends lie free in the mantle cavity as opposed
to ducts not as wide as the receptacles and without
free ends and (ii) the left receptacle terminating
on the right rather than the left. Furthermore, the
colleteric glands of S. beauforti divide into
approximately 50 lubes and those of S. leptodiae
into less than 20 (Boschma 1949) and so are quite
different from the glands in S. amplituba which
divide into more than 100 tubes. S. beauforti has
the cuticle of the mantle of 120-130// in
thickness (Boschma 1955) considerably thicker
than that of iS. amplituba (75-100// thick).
Apart from the morphological distinctions, S.
amplituba should be considered a new species
since each of the most similar species all occur
in a different host genus and the work of Fratello
(1968) on chromosomes indicates a high degree
of host specificity among saccuiinids.
Sacculina granifera Boschma, 1973
The most important characteristic of S.
granifera is the structure of the excrescences of
Fig, 1; Sacculina amplituba: a, whole parasite — left side. A, B and C are the anterior to posterior, dorsal to
ventral and left to right dimensions respectively; b, left side with left side of mantle removed, scale = 4 mm;
c-h, transverse sections of posterior part of parasite showing male organs, scale = 1 mm; i, transverse section
of external cuticle of mantle showing excrescences, scale = 40//; transverse section of colleteric gland in
region of greatest division of tubes, scale = 300// .
PHILLIPS: PARASITIC BARNACLES FROM CRABS
257
258
MEMOIRS OF THE QUEENSLAND MUSEUM
the external cuticle described as irregularly
globular (Boschma 1973). Scanning electron
micrographs (Plate 40. Pig. l.b) reveal that each
of the globular excrescences bears many small
spines. F'urther more, Boschma described the
collctcric glands of N. granifera as dividing into
40 lubes in one row. Specimens examined by me
show the collctcric glands to be divided
consistently into up to 100 lubes in the region of
greatest division.
Hetcrosaccus lunatus nov. sp.
(Fig. 2 a-i)
MATI RlAl EXAMIM D
Hoioiapi Queensland Museum W7148, ex
Charyhdis callianassa, Moreton Bay, collector R.
Bishop. Feb. 1975.
Pnrm'iPIs Queensland Museum W7149 ex Charyb-
dis callianassa. Moreton Bay, collector R. Bishop, Nov.
1974, three whole specimens on one host. Queensland
Museum W7150. ex C. callianassa. Moreton Bay,
collector W. Phillips, 24/v/1973, sections.
Diagnosis
Kidney shaped parasite, more than one of which
may occur on the one host. Cypris cell receptacles
within visceral mass and deeply crescentic,
terminating near colleleric glands and containing
sponge-like meshwork. Receptacles open widely
into straight ducts also with sponge-like
mcshw'ork; ducts open near stalk; receptacles and
ducts surrounded by lymph spaces. Collctcric
glands posterior, with 60-70 lubes in five to six
rows in region of greatest division. External cuticle
with papillae 15/; long and 10// apart, of same
material as mantle. Retinacula rounded flaps of
tissue occurring singly each with four to six rods.
Parasite of Charyhdis callianassa (Herbst).
Description
Body form: The externa (Pig. 2a) is kidney-
shaped having a wide mantle opening with poorly
developed musculature. The numbers of cxternac
per crab varies from one to three, the exlernae
being smaller where there arc more than one on
one host. Dimensions of the holotype: A = 10,
B = 19, C = 7; and of whole paratypes (i) A
= 10, B - 20, C = 9, (ii) A = 8, B = 13, C
= 6, (iii) A = 8, B = 11, C 6.
Cypris cell receptacles: The male organs are
confined to the posterior dorsal portion of the
visceral mass w'hcre they are surrounded by lymph
canals. The rest of the visceral niass is without
pronounced lymph canals. The ducts which open
close to the stalk (Fig. 2,h) are of a consistent
diameter of about 400// and have a sponge-like
structure for most of their length. The lumen of
each duct is 40-50// wide. The ducts are not
significantly convoluted. They pass gradually into
the receptacles which arc only slightly wider than
the ducts (Fig. 2 d- g). The receptacles have a
deep curvature and terminate near the colleteric
glands (Fig. 2g).
Colleteric glands: The colleleric glands are
cushion-shaped bodies lying in the posterior half
of the visceral mass near the middle. In the region
of greatest division there are 60-70 tubes in five
to six rows (Fig. 2i). There is no chitinous lining
but villiform projections arc present on the
inside.
Mantle: The external cuticle of the mantle is
15-20// thick and is covered with papillae
approximately 15// long and 10// apart (Fig. 2c).
The papillae are projections of the cuticle of the
mantle and may be partially or wholly obscured
by a granular coating (Fig. 2b). Scanning electron
micrographs (Plate 40, Fig. b) clearly show the
shape and distribution of the papillae and the
granular coaling.
Retinacula (Fig. 2c) are single rounded flaps
of tissue about 20// wide each containing four to
six rod-like bodies. The retinacula are about
200// apart.
Remarks
The name Helerosaccus lunatus nov. sp. is
proposed because of the deeply crescentic nature
of the cypris cell receptacles.
Heterosaccus multilacinensis nov. sp.
(Fig. 3 a-i)
Mati'riai Examimd
Hoi.otypi. Queensland Museum W7151, sections, ex
Carybdis iruncata, East Moreton Bay, collector W.
Phillips 6/iv/l973
pARATYPiS: W7152. cx C truncQta. four specimens
on host, same data as holotype.
Diagnosis
Kidney-shaped parasite more than one of which
may occur on the one host. Cypris cell receptacles
Fi(. 2: Helerosaccus lunaius: a, whole parasite — left side, scale = 3 mm; b, transverse section of external cuticle
of mantle, scale = 10//; c, internal retinacula, scale = 25//; d-h, transverse sections of posterior part of
parasite showing male organs, scale =• 1 mm; i, transverse section of colleteric glands in region of greatest
division of tubules, scale = 200// .
PHILLIPS: PARASITIC BARNACLES FROM CRABS
259
260
MEMOIRS OF THE QUEENSLAND MUSEUM
shallow crcsccnlic, not surrounded by lymph
spaces; ducts slightly convoluted opening away
from stalk; meshwork present in receptacles but
not in ducts. Collcteric glands in posterior part of
visceral mass, region of greatest division of tubes
shows 40-60 lubes in four to five rows. External
cuticle with papillae 15//' long, close together at
base and separated by 5// at tip. Papillae of same
material as cuticle. Retinacula of one to six
rounded Haps on a common base with one rod per
flap. Parasite of Charybdis truncata
(Fabricius).
Description
Body form: The externae (Fig. 3a, b) are
kidney-shaped with a wide mantle opening
showing poorly developed musculature. The
number of externae per crab may be more than
one. Dimensions of paratypes (i) A = 10, B =
17, C = 4, (ii) A = 9, B = 10, C = 7, (iii) A
= 10. B = 19, C = 6, (iv) A = 18, B = 14,
C = 4.
Cypris cell receptacles: The male organs are
confined to the posterior dorsal portion of the
visceral mass and are not surrounded by lymph
spaces although lymph spaces are seen commonly
in the visceral mass (Fig. 3 a,d-g). The ducts
which open close to the stalk are somewhat
convoluted and do not possess a sponge-like
structure (Fig. 3 e-g). They have a diameter of
220 240// and the lumen is 50-70// in diameter.
The ducts pass gradually into the receptacles
which have a shallow curvature and do possess a
sponge-like structure. The receptacles arc two to
three times as wide as the ducts (Fig. 3 d,e).
Collcteric glands; The cushion shaped collcteric
glands arc situated in the posterior part of the
visceral mass towards the middle. The region of
greatest division of tubes shows 40-60 tubes in
four or five rows (Fig. 3h). There is no chitinous
lining, but the lumen is lined with villiform
structures.
Mantle: The external cuticle of the mantle is
approximately 50// thick with papillae 15// long
which are extensions of the cuticle (Fig. 3b). The
papillae touch at the bases and taper to a blunt
point. They arc about 5// apart at the tips.
Retinacula (Fig. 3c) consist of one to six
rounded flaps of chiton on a common base, each
flap containing two or three rods which may
project out of the flaps and appear to extend into
the base. The flaps are from 18-20// long and
spaced about 200// apart.
Remarks
The name Heterosaccus multilacinensis nov. sp.
is proposed because of the many flaps on each
retinaculum.
Boschma (1963) described Heterosaccus
papillosus from several species of the genus
Charybdis including C. callianassa. It is similar
to, yet distinct from, the two species described
here. //. lunatus differs from //. papillosus in
having (i) wider receptacle ducts opening close to
the stalk, (ii) deeper curvature of the receptacles,
(iii) greater division of the lubes in more
posteriorly placed collcteric glands, (iv) longer
more closely placed papillae on the mantle, and
(v) retinacula consisting of flaps with rods rather
than spindles. H. multilacinensis differs from H.
papillosus in having (i) ducts opening close to the
stalk rather than at a distance from the stalk, (ii)
shallower curvature of the receptacles, (iii) greater
division of tubules in more posteriorly placed
collcteric glands, (iv) papillae closer together and
of a different shape, and (v) retinacula consisting
of many Haps w ith rods. H. multilacinensis differs
from //. lunatus in having (i) narrower more
convoluted male ducts without a sponge-like
structure, (ii) less division of the tubes of the
collcteric glands, (iii) longer, thicker, more closely
packed papillae, and (iv) retinacula consisting of
several Haps on one stalk.
DISCUSSION
Undoubtedly the rhizoccphalan fauna of
Australia will be found to be more numerous than
is presently recorded. The influence of these
parasites on the biology of their hosts is
considerable and where the hosts are of
commercial importance they produce considerable
losses to the industry. As yet no sacculinid has
been found to parasitize the commercial mud crab
Scylla serrata in Australia although S. beauforti
and Loxothylacus ihlei Boschma 1949 have been
described from this species in Indonesia.
Discrimination of species of rhizoccphala is
difficult since insufficient is known of the details
of their life cycles or the variability of
characteristics with the age and maturation of the
externa. Scanning electron micrographs of the
exterior surfaces of the mantle of young externae
should provide accurate and detailed information
on one rather important taxonomic feature, viz.
the form of the cuticular excrescences. Clear
differences can be seen between Sacculina
amplituba and Heterosaccus lunatus (Plate
40).
PHILLIPS: PARASITIC BARNACLES FROM CRABS
261
Fig. 3: Heterosaccus muliilacinensis: a, whole parasite, left side, with left side of mantle removed, scale -
3 mm; b, papillae of external surface of mantle, scale =10//; c, retinacula, scale = 25//; d-g, transverse
section of posterior part of parasite showing male organs, scale = 1 mm; h, transverse section of colleteric glands
in region of greatest division of tubes, scale = 300//.
262
MEMOIRS OF THE QUEENSLAND MUSEUM
Probably the most promising means of
discriminating species would be chromosome
techniques as described by Fratello (1968) who
was able to discriminate several species previously
described as one.
ACKNOWLEDGMENTS
The sacculinids described in this paper were
related to a post-graduate honours project in the
Department of Parasitology, University of
Queensland. 1 would like to express my thanks to
Professor J.F.A, Sprent for permission to use
departmental facilities.
I would also like to thank Dr L.R.G. Cannon
for assistance in preparing the manuscript; the
C-S.LR.O.. for use of its boat for collecting
specimens. Mr J. Hardy of the Electron
Microscope Unit and Mr J. Mines for advice on
histological matters. The late Dr H. Boschma of
Leiden Museum, Netherlands also provided
valuable information concerning this work.
LITERATURE CITED
Boschma, H., 1933. The Rhi/occphala in the collection
of the British Museum. J. Linn. Soc. 38:
473 552.
1 949. Sacculina beauforti and Losolhylacus ihlei, two
Rhi/ocephala of the crab Scylla serraia (Forsk.).
Bijdr. Dierk. 28: 41-6.
1955. The described species of the family Sacculin-
idac. Zool. Verb.. Leiden. 27: 1-76.
I 967. A Rhi/ocephalan parasite of the crab Charyhdis
callianassa (Hcrbsl.). Prov. K. ned. Akad. Wet.
66C; 132 3.
1973. Sacculina granifera nov. spec., a rhizocephalan
parasite of the crab Portunus iPortunusj pelagicus
(Linnaeus) from the coast of Queensland. Proc. K.
ned. Akad. Wei. 76C: 313-8.
Frathi.io. B-, 1968. Cariologia c lassonomia dci
Sacculinidi (C'irripedi, Ri/ocefali). Caryologia 21
(4): 359-67.
H \s\\ 1 t I , W’.A., 1 888. On Sacculina infesting
Australian crabs. Proc. Linn. Soc. N.S.W. 3:
171 1-2.
Thomson, J.M., 1951. Catch composition of the sand
crab fishery in Morelon Bay. Aust. J. mar.
Fre.shwat. Re.^. 2: 237-44.
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATH 40
A: External surface of mantle of Sacculina granifera 1. x500; 2
x2500
B: External surface of mantle of Heterosaccus lunatus 1. x450; 2
xl350.
PHILLIPS; PARASITIC BARNACLES FROM CRABS
Plate 40
Mem. Qd Mus. 18(2): 265-71, pis. 41-42 [1978]
THE OLD BURIAL VAULT
AT NORTH QUAY, BRISBANE
J. C. H. Gill
Queensland Museum
ABSTRACT
A brick structure accidentally uncovered in 1972 during construction of the Riverside
Expressway, Brisbane, opposite the eastern side of the junction of Herschel Street and North
Quay, has been determined to have been a burial vault dating from the penal settlement days
of Morcton Bay and Brisbane Town. This probably represents the last convict structure to
be located in this area. It has been assessed as fully as possible from the limited record able
to be made at the time of discovery, as within an hour of this it had been destroyed
totally.
In the Queensland Museum are eleven sand
stock bricks, registered H9750 to H9760 inclusive,
collected by the Museum on 23 August 1972; their
in situ provenance is recorded as ‘from suspected
grave site on Brisbane River bank 100 yards
[90 m] west of Tank Street’. They provide a
slender base on which to assert they were part of
a structure dating from the convict days of
Brisbane Town, yet there is evidence which
affords positive indications of the time when the
bricks were used, of the structure of which they
were part, and of the use to which that structure
was put.
The narrative which follows will incorporate the
historical evidence. Then the structure as such will
be dealt with as far as it is possible to make a
reconstruction from the relics available.
Historical Evidence
On 23 August 1972 a bulldozer working on the
Brisbane Riverside Expressway construction
opposite the junction of North Quay and Herschel
Street dislodged some bricks buried in the soil,
and disclosed an aperture in the sloping bank
behind which lay a buried chamber. Examination
revealed a barrel-vaulted, brick structure, over
which a coating of lime mortar had been applied
internally.
Main Roads Department officers supervising
the construction halted work and informed the
Royal Historical Society of Queensland of the
find. The author was contacted and requested
Queensland Museum assistance to assess the
historical value of the discovery. Very limited ^ime
was allowed for investigation before construction
work resumed. The urgent situation precluded
detailed archaeological investigation. Fortunately
a local television station, QTQ 9, filmed external
and internal views, and later donated the film to
the Queensland Museum. Rough sketches were
made of the structure and some bricks were taken
for later examination. As nothing further could be
done, the Main Roads Department officers were
informed and within minutes the bulldozer
operator, who had kept his motor running all this
time, had destroyed the structure. A press
photographer arrived too late to do other than
photograph a few bricks lying where the structure
had been, resulting in coverage in the ‘Courier-
Mair, on 24 August 1972, p. 3.
With the wisdom of hindsight it seems to have
been a rather inglorious episode. Yet the element
of surprise and the pragmatic attitude of the
people concerned with the site gave no opportunity
for complete survey. Furthermore the nature and
purpose of the structure was at first not clear. The
situation could not compare, for example, with the
discovery, again per medium of a bulldozer, on
30 March 1974, of an early brick vault in what
had been the earliest official burial ground in
266
MEMOIRS OF THE QUEENSLAND MUSEUM
Fk. 1. Plan based on Assistant Surveyor R. Dixon's 1840 survey, with outlines of modern streets superimposed.
Sydney from 1793 to its closure on 27 January
1820. The purpose of the structure was clear —
it contained a collapsed wooden coffin within
which were bone fragments and Sydney
University archaeologists had three days to
assemble a team to survey it. Resulting public
interest gave the vault a further reprieve and in
the end it was dismantled for reconstruction at the
Old Sydney Town Project. Finally a thoroughly
researched monograph was published (Birming-
ham and Liston 1976).
The site of Brisbane's earliest burial ground was
investigated previously by Gill (1959-60, 1974),
who recalled that the structure was only yards
away from isolated memorials on North Quay
known as ‘the children’s graves’.
Relics from Brisbane’s penal settlement era are
few. The absence of a burial ground from this era
is particularly notable when one considers the
plethora of early memorials in the other states of
Australia. Controversy has raged as to the site of
Brisbane’s earliest burial ground. Yet it has been
proved beyond doubt that the penal settlement’s
earliest and only burial ground was that shown
in the town maps of Assistant Surveyor Robert
Dixon (1840)* and Surveyor Henry Wade
(1842)t situated on the north bank of the
Brisbane River at the northern end of the William
Jolly Bridge in the area bounded by the present
day Skew Street and Eagle Terrace.
One of the causes of controversy was the
presence of three (at least) memorials on the river
bank at North Quay directly opposite the eastern
corner of North Quay and Herschel Street,
immediately above a prominent rock face which
rises abruptly out of the river. The Royal
Historical Society of Queensland has a photo-
graph (said to have been taken about 1876) which
shows memorials in situ at this point (Plate 41,
fig. I). This photograph, acknowledged to Mr
H.A. Jones was donated in 1921 and was
reproduced in the ‘Brisbane Courier' on 25
October 1913 at p. 12. It w^as captioned: ‘The
Soldiers Graves on North Quay. A controversy
has for some time been waged in the “Courier”
as to the exact location of the soldiers’ graves in
the old burial ground on North Quay. Our photo
shows their position (indicated by a X).' The
memorials have long been gone from North Quay
but they are still to be seen in Toowong General
Cemetery. On 5 October 1881 the memorials at
North Quay and any remains (under licence
* Queensland Survey Office Map M.T.3
t Queensland Survey Office Maps 6 and B.il82
GILL: OLD BURIAL VAULT
267
issued by the Colonial Secretary on 23 August
1881) were removed to Toowong.*
Only three memorials and remains were
removed. Whether there were any more burials on
this site is a matter of doubt.
The three memorials all commemorate young
children — William Henry Roberts, son of
Charles Roberts of the Commissariat Department
who died of illness on 15 November 1831 aged
3 years and 2 months), Peter, son of Private Peter
McCauley of H.M. 17th Regiment of Foot,
accidentally drowned on 5 January 1832, aged 5
years and 8 months; and Jane, daughter of John
Pittard, former Colour Sergeant of the 57th
Regiment, who died on 29 January 1833, aged 12
months and 13 days (Plate 41, fig. 2).
The McCauley and Pittard memorials are the
usual headstone and footstone types; the Roberts
memorial is a catafalque or table monument type
with the inscription on the flat top. The memorials
show evidences of the ravages of time. In 1962
the Brisbane City Council took steps to refurbish
them. The decay of the first two memorials has
been arrested, but the inscription on the Roberts
memorial is now lost and is, in fact, covered by
a coat of stucco. Fortunately it was recorded in
its entirety in 1913 with some errors which, except
for the age (5 instead of 3 years) were corrected
by Gill (1959-60).
Assistant Surveyor Dixon in his 1840 map, in
addition to the official burial ground, showed two
small squares, habitually used by surveyors to
indicate isolated graves, at a point corresponding
to the location of The children’s graves’ (Gill
1959-60).
Steele (1975) indicates that the Reverend John
Vincent, in his initial report to Archdeacon Scott,
complained that the settlement’s ‘burial ground is
at a very remote distance, is too small and so very
difficult to be dug . . . that two men with the
utmost exertion are scarcely able to prepare a
grave in six or seven hours’.
If the children were buried where the memorials
were located, it would have required the cutting
of kists in solid rock, and the purpose of the
structure unearthed on 23 August 1972 now
becomes clear. It was a burial vault sited on or
slotted into the rock in which bodies were placed
whilst sorrowing relatives were allowed to place
memorials around it, a much easier process than
cutting a full-size grave into the rock each
time.
Evidence of the existence of the vault was
available, but the disappearance of the structure
beneath the earthen embankment of the widened
North Quay in the latter part of the nineteenth
century and the presence up to 1881 of the table
monument to the Roberts’ child led to the belief
that references to the vault were references to the
table monument.
The removal of the memorials from North
Quay on 5 October 1 88 1 excited no comment from
the press of the day. In fact it was not until
Friday, 21 October 1881 that the ‘Brisbane
Courier’ on page 2 remarked:
‘The old landmarks of Brisbane are rapidly
disappearing and although by many the removal of
these mementoes of early days may be regarded with
some lingering feelings of regret the improvement
which will result in the general appearance of the city
must be a matter of satisfaction*.
The disappearance of the old convict barracks and
old buildings on the corner of Queen and Albert
Streets is favourably commented upon, whilst the
fact that the dreary looking goal at Petrie Terrace
will soon become a thing of the past also is
editorially approved. Then:
‘Another “landmark" that has disappeared from
within the boundaries of the city are the old graves
on the North Quay. In these graves were the remains
of three bodies that have been under the ground for
some fifty years. By the order of the Colonial
Secretary, the remains with the tombstones that were
over them, have been removed to the cemetery at
Toowong, that being considered a more fitting place
for them, where it will be less trouble to keep the
graves in order, and where they will be subjected to
le.ss desecration*.
It was not on-the-spot reporting by any means and
it is not surprising that the vault escaped notice
by the press. There the matter rested for 32 years
whilst improvement works along the North Quay
greatly altered the old landscape.
Professor F. W. S. Cumbrae Stewart through
the columns of the ‘Brisbane Courier’ of 30
September 1913 (p. 4) drew attention to the
childrens memorials in Toowong Cemetery,
expressed the belief that they had been removed
thence from North Quay in 1876, and sought
information as to where the gravestones originally
stood on North Quay.
* John Oxley Library MS.O.M. 65/18, Queensland State Archives [QSA.] COL/A319, 3694; QSA. COL/G15,
p. 585, 81/1233. , „ . u u
t John Oxley Library ibid. Mr Melville the sexton at Toowong Cemetery in 1940 says it appears to be 3 but
could be 5 years, on account of weathering of the stone. In 1881 (59 years earlier) the age is given as 3 years
and 2 months. The stone was then 50 years old and the inscription much more legibde.
268
MEMOIRS OFTHE QUEENSLAND MUSEUM
Between 30 September and 5 November 1913
the ‘Brisbane Courier' published fourteen letters
on the subject of the ‘Old Burial Ground' (Gill
1959-60).
On 7 October 1913 the ‘Brisbane Courier’ (p.
4) published a letter from a Mr Thos. Illidge who
stated he was on the spot when the remains were
dug up by Mr D. Hannah of Paddington.
Published with the letter was an interview by a
Courier reporter with Mr David Hannah of
Rosalie who said he had assisted in the removal
of headstones on the river bank from North Quay
to Toowong. It is significant that Mr Hannah did
not refer to the removal of remains.
On 16 October 1913 (at p. 7) ‘Sixty Years a
Queenslander' wrote about ‘the monumental
Slone' which in earlier years marked the spot
known to old residents as the ‘soldiers' graves'. He
referred to the difference of opinion which seemed
to exist as to the exact site. This he regarded as
quite natural for during the last few years (i.c.
prior to 1913) ‘surface appearances have changed
or become obliterated’. He also says:
‘In the 'fifties and 'sixties of last century much of
the river frontage along the North Quay was covered
with a tangled growth of lantana. which could only
be penetrated by a bird-nesting boy or a billygoat. As
the former, 1 have often “worked my passage"
through this scrub and my memory fixes the site of
the graves as about midway between Tank and
Herschel Streets and just above a prominent rock
which here rises abruptly over the river.’ (Gill
1959-60).
He was almost on the correct site and indicates
that with the profusion of under-growth making
the site inaccessible the general public had come
to refer to them as the ‘Soldiers' Graves’.
On 20 October 1913 (p. 6) the ‘Brisbane
Courier’ published a letter from a Mrs Lucy
Sonnonschein (nee Wynn) of Warwick. She
said:
‘The old burial grounds were situated opposite the
present Roma Streets goods shed gates. After the
burial ground was removed, the land was built upon
and occupied by Davies (Davis) the Jew. Joseph Jayes,
Bill and Joe Jewell, carpenters, Joshua Peter Bell, and
Bulger, a wine merchant. New shops have replaced
these dwellings. This burial ground was known as the
“Soldiers’ Graves". On the bank of the river in North
Quay there was a large vault, opposite Paddy Pacey's
milking yard, which marked the spot where some
officer was buried.' (Gill 1959-60).
The existence of the burial vault, rediscovered
in 1972, being unknown, the author as already
slated believed this to be a reference to the table
monument to the Roberts' child.
Yet on 5 November 1913 the ‘Brisbane Courier’
published (in the Courier Home Circle Sup-
plement at p. 6) the final two letters in the burial
ground debate. Mr A. E. Campbell of Rockhamp-
ton refers to the sketch in the ‘Courier' of 25
October of the “soldiers' grave'*.
‘Yes. that is about the spot, only high up, about
ten yards from the fence, [the X was placed too low
on the photograph], and there is a high stone face
projecting towards the river. It was the grave of an
officer who went out riding and his horse carried him
against a tree and broke his neck. I think his name
on the lop slab was “Lawson", The tomb must have
been shifted in the early ’sixties. The bottom end near
the river was broken, and it acted as a cave. Many
a one slept there for the night, for it was a cosy camp
for a cold night.’ (Gill 1959-60).
Here again the author believed the reference w-as
to the table monument; but it would have made
cramped quarters and access would have been
grossly difficult. The mean dimensions of the
monument are only 1-905 m by 0-990 m by
0-852 m high. The brick vault was in a different
category — in the 'fifties and 'sixties it was buried
beneath lantana; any remains in it would have
been moved in 1881; (in any event the remains
of young children would not have the durability
of those of adults); the access, if the end near the
river had broken away, would have been easier,
and air circulation and animal scavengers would
have removed any associations with its grisly past
by the time ‘down and outs' began to camp in
it.
It is considered that these reports provide
sufficient evidence of the existence of a vault.
In addition to its existence we also have
evidence ol its use. .A macabre story recounted by
J. J. Knight (1895) furnishes this. Talking of the
offences for which a convict could be sent to the
triangle Knight says:
‘A very common offence was the purloining of a
few cobs of corn and potatoes, the chief ingredients
of the convicts' much prized “fiddle-cake", the love
for which caused many sore backs at the triangle, and
weary legs on the treads. These two things having been
obtained, the corn was ground on an improvised grater
made usually of a piece of tin or zinc, in which holes
had been punched.'
Alter describing the grating and recipe for the
delicacy, Knight goes on
‘But it was often more difficult to obtain the
material for the grater than the corn and potatoes,
and in one case at least a convict finding this so
resorted to a very questionable means of gratifying
his desires. The overseer at Eagle Farm was
unfortunate enough to lose one of his children, and
the body having been placed in a tin box, was laid
in one of the vaults on the river bank near Herschel
Street. This fact was. of course, known to the
prisoners, and a day or so after the funeral one who
had so urgently desired the possession of a “grater"
GILL: OLD BURIAL VAULT
269
effected an entrance to the vault and taking out the
dead body of the child found in the box the material
for the manufacture of this necessary implement of
the “fiddle-cake" maker. Fortunately for the
sacrilegist, he was not found out, and was enabled to
make many specimens of that delicacy, which was
described by one who had often partaken of it as being
“better than any pie going".’
Apart from anything else this indicates that a
measure of improvisation was required still in the
undertaking line. If there were no coffins readily
available for children of officials one wonders
what sort of hurriedly thrown together boxes were
used for convict burials.
Knight talks of one of the vaults on the river
bank near Herschel Street. The available evidence
indicates there was only one such vault. It appears
to have carried no marks of identification (the
adjacent memorials made this unnecessary). Mr
Campbell does mention the name Lawson, but he
appears to be confusing the inscribed Roberts'
monument with the vault. In Plate 41, fig. 1 the
Roberts' monument is plainly discernible but the
vault is not, although Mr Campbell has no doubt
about the location of the latter. It was because
the vault carried no identifying marks or
inscription that all the stories about soldiers’
graves and officers' burial places came into being.
The correct function of the vault probably
occurred to no one; the memorials were there and
it would not be known generally that they were
sited on solid rock. Once North Quay was widened
in the late nineteenth century the footpath on the
river side of the roadway was cut off from the
steep river bank by a wooden picket fence of
sturdy construction. Apart from wooden stairway
accesses to the Commercial, Brisbane and
Brisbane Grammar School Rowing Club
Boatsheds there was no access and for upwards
of half a century until World War II the North
Quay river bank from the O'Connor Boathouse
(Commercial Rowing Club) upstream to the Grey
Street (William Jolly) Bridge became a terra
incognita.
Even the erection of air raid shelters in 1942
resulting in the partial disappearance of the fence
followed by its nearly complete demolition in the
post-war period brought about no interference
with the river bank itself.
It was not until the construction commenced of
the Riverside Expressway section of the South
East Freeway that the riverbank began to be
altered substantially. In August, 1972 the work
began on the expressway exit to Herschel Street.
This meant carving away a section of the
riverbank; before the bulldozer got down to the
rock, it blundered into the old vault with the
results already indicated. The vault could well
have been coeval with the Windmill and the
Commissariat store*, and may well be the last of
the Brisbane Town convict structures which
remained to be rediscovered.
Description of the Vault
The vault itself was constructed of sand-stock
bricks (as they are commonly known) with mean
approximate dimensions of 205-9 mm x 63-5 mm
X 107-9 mm. The dimensions of the vault as
estimated by Mr A. Sweetser, Queensland
Museum, in a drawing he made (Figure 2) are
given as 1-53 m x 1-83 m with a barrel vaulted
roof rising above 1-53 m high side and end walls.
The entrance at the northern end had been bricked
up after any remains had been cleared from the
vault. Two wooden slabs lay on the floor covered
by rubble and a partly burned piece of hardwood
plank lay near the southern end of the vault. There
was some evidence of dampness in the earth floor
but no signs of the vault ever having been
inundated with water; the run-off from North
Quay to the Brisbane River was steep enough to
prevent water from gathering and lying in the
vault. The only wall of the structure to show up
in full width in the movie film was the northern
wall, and estimating from the known lengths of
the bricks collected from the site the mean width
of the wall could be 1-72 m, 019 m more than
Mr Sweetser's estimate (Figure 2).
The side wails were grooved for their entire
lengths at two levels, which indicated an intention
to have the coffins resting on planks (fitted into
the grooves) above ground level and provisions for
an upper layer of coffins if needed.
An analysis of the lime mortar used to bind the
bricks and plaster the interior of the vault has
been made by the Queensland Government
Chemical Laboratory. The analytical report
(Appendix I) established the source of the burnt
lime used in the mortar as Ipswich; a clear link
with the penal settlement days.
As regards the bricks, what were known as the
old and new brick kilns respectively both were
located within 230 m of the vault as shown on
Figure i (Steele 1975). So the raw materials for
the brickwork were close at hand.
Plate 42. fig. 1 shows the intrusion of some tree
roots, but the lack of moisture had inhibited the
growth of these.
Owing to the gross interference with ground
levels, even before the discovery of the vault as
♦Both completed in 1829.
270
MEMOIRS OF THE QUEENSLAND MUSEUM
well as subsequently, no soil profile could be
taken. However it was obvious to the eyewitness
that the vault had been covered by fill from
elsewhere than the river bank itself and that the
river bank proper began at the base of the
structure.
Apart from the planks within the vault no
timber beams or lintels were noted in the walls
of the structure; it appears to have been small
enough not to require limber to span any openings
or archwork which the bricks themselves could not
support. No other features remain to be remarked
upon.
Thus a few feet of film and eleven bricks at
the Queensland Museum, with this account, will
be the only record of a structure dating from the
early penal settlement days of Brisbane Town.
LITERATURE CITED
Birminomam. J., and Liston, C., 1976. Old Sydney
burial ground 1974 — emergency excavation.
Studies in Historical Archaeology No. 5, vii + 40
pp.
Gll.L, J. C. H., 1959 60. Some investigations into the
site of Brisbane’s earliest burial ground. J. R. Hist.
Soc. Qd. 6 (2): 388-417.
1974. Dc Mortius. /^7Wiiso\. P. D. (Edit.) Moreton
Bay Sesqui - Centenary 1824-1974 (Library Board
of Queensland: Brisbane).
Kmout. j. j., 1895. In the Early Days, xi + 390 pp.
(Sapsford and Co.: Brisbane).
STHta.i:. J. G., 1975. ‘Brisbane Town in Convict Days
1824-1842.' xxii + 403 pp. (University of
Queensland Press: Brisbane).
FiG. 2. Diagram of the vault, after a sketch by Mr A. Sweetser held in the Queensland Museum.
GILL: OLD BURIAL VAULT
271
APPENDIX 1
Government Chemical Laboratory
William Street,
Brisbane, Q. 4000
16th November 1976
Dr. D. J. Robinson
Qld. Museum
Gregory Terrace &
Bowen Bridge Road,
BRISBANE.
Ref: DR;DF HlO/5
Dear Sir,
On Tuesday 8lh November, soft portions of the white
material in mortar samples H9255 and 9578 provided
by the Museum were scraped out and this material on
analysis gave carbon dioxide, 9% acid soluble silica and
6% magnesium as magnesium oxide. Aslo present is
calcium oxide, iron oxide and alumina.
The Geological Survey Office was contacted and
confirmed that the results of an early analyses of the
Limestone Hill Limestone was
Loss of Ignition
43.8%
Total Silica
5.7%
Iron oxide
0.9%
Alumina
1.1%
Calcium oxide
27.5%
Magnesium oxide
20.6%
This indicated that this mineral is an impure mixture
of calcium and magnesium carbonates. Other areas of
limestone in the Ipswich area have similar compositions.
The presence of acid soluble silica and the magnesium
oxide indicates that the burnt lime used in the mortar
sample probably came from the Ipswich district which
seems to be the only source of limestone in the local
area other than sea shells. The Ipswich deposits have
variable compositions but all seem to have more than
5% magnesium carbonate.
Microscopic and hand lens examination showed no
shell particles and it is not expected that shells burnt
to calcium oxide would contain acid soluble silica and
magnesia.
Warwick and Gympie limestones are much lower in
magnesia.
D. Mathers
Director
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 41
Fig. 1 : The memorials in situ ca. 1 876 - photograph in Royal Historial
Society of Queensland collections.
Fig, 2; Children’s memorials at Toowong Cemetery.
GILL; OLD BURIAL VAULT
Plate 41
1
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 42
Fig. 1: Interior of vault-photograph from QTQ 9 16 mm film.
Fig. 2: Riverside Expressway showing remains of bold, rocky face
above which the vault was located.
GILL: OLD BURIAL VAULT
Plate 42
Mem. QdMus. 18(2): 273-319, pis. 43-103 [1978]
THE CORAL GENUS ACROPORA (SCLERACTINIA:
ASTROCOENIINA; ACROPORIDAE) IN THE CENTRAL
AND SOUTHERN GREAT BARRIER REEF PROVINCE
Carden C. Wallace
Queensland Museum
ABSTRACT
The genus Acropora (excluding subgenus Isopora Brook) is revised from zonation studies
on two reefs, and incidental collecting from many other localities in the Great Barrier Reef
Province. As a result of field studies, many species are placed in synonymy, and a resulting
forty species are redescribed. The revision includes description of field characteristics of the
species, many of which were previously known only from small skeletal series. Problems of
identification and delimitation of the various species are discussed.
In the Great Barrier Reef Province, as in most
Indo-Pacific reef areas, the regions of densest
coral growth are regions dominated by the
branching coral genus Acropora. Over 300 species
have been described for this genus. Some 80 have
been recorded from the Great Barrier Reef, and
many more would have been added to this list had
the reef slopes and deeper waters been more
accessible to early collectors. The original 300
rtiay be reduced to around 70 by synonymy, and
the 40 species here treated are considered to
represent at least 90% of the branching Acropora
in the Great Barrier Reef Province.
The following is a working paper on the
taxonomy of this neglected genus. It is based on
regional studies in the Great Barrier Reef region,
and in particular on line transect studies carried
out at Big Broadhurst Reef (Wallace 1975,
Wallace and Dale 1977) and Bushy-Redbill Reef
(Wallace and Lovell 1977, Wallace unpublished).
It is an attempt to open the study of the genus
to workers in all aspects of coral study and to this
end it concentrates on field description of the
species.
it is clear to the author after some experience
with the genus that few species can be fully
understood by simple morphological studies of
colonies, even with the addition of ecological
information. Even the population cannot be
considered a sufficient unit for study as there are
often other key species in the surrounding
assemblage which may look like, physically
interact with, or affect the members of the
population in some way. This paper seeks to
indicate such problems.
Taxonomic History
The name Acropora Oken, 1815 was officially
validated in 1963 (Boschma 1961, China 1963).
It came into general use with Verriil (1902),
although not without some argument and
nostalgia for the previously widely used
Madrepora Lamarck: ‘it is with a feeling of regret
that we are forced to abandon the use of a generic
name which remained unquestioned by four
generations of authors during the golden days of
systematic zoology’ (Mayor 1924, p.vii).
Many authors have described species in this
genus. Few have attempted to synonymise or
group species, fewer still to define and standardise
terms. The most notable single contribution to the
taxonomy of the genus was that of Dana (1846)
who gave very careful attention to his specimens
collected during the United States Exploring
Expedition. His arrangement of species according
to colony shape and radial corallite shape without
defined subgeneric status remains the simplest and
most acceptable treatment of the genus.
Brook (1893) produced the only true mono-
graph of the genus, summarizing all previous work
and re-describing all described species. He ordered
the species into subgeneric groups, attempting to
274
MEMOIRS OF THE QUEENSLAND MUSEUM
take into account all skeletal characters, but his
subgenera have not found general acceptance:
indeed, current synonyms often span the
groupings. Brook unfortunately did not examine
Dana’s specimens, although he did see most of the
European located types, and most Dana species
can be shown to have a Brook synonym. Verrill
(1902) also split many Dana species needlessly.
Nemenzo (1967) revived the use of group names,
although not affording subgeneric status to his
groups.
Of the twentieth century authors, Vaughan
(1918), Hoffmeister (1925), Crossland (1928.
1952) and Wells (1954) have treated the genus
in detail, with ecological data, although not
concentrating exclusively on Acropora.
Detailed underwater studies of Acropora were
first made by J. Verwey in the Bay of Batavia
in the 1930’s, and the taxonomic results of his
study are still unpublished (see Umbgrove 1939,
p.56, 1940, p.303; Wells 1954, p.4I4). In
discussions with Dr Verwey I have found that we
are usually in agreement in our general assessment
of species limits (though obviously not always on
names). His opinions pre-date those of this paper
by some forty years, and it is hoped that his work,
which is to be a complete monographic treatment
of the genus, will soon be available.
Characters and Terminology
(For coral terminology not specifically referrable
to Acropora see Moore, Hill, and Wells
1956).
Acropora lacks most of the skeletal characteris-
tics used in the species identification and study
of variability of other corals, e.g. columella,
dissepiments, septal structures. The main
character defining the genus is the method of
branching: an axial polyp builds a corallite of
increasing length, and buds off radial polyps from '
its growing tip. Any radial polyp has the potential :
to take up an axial role. A variety of branching
patterns occur, and these (with commonly applied
terminology) are summarized in Figure 1.
The polyps have twelve tentacles, one of which
(overlying a directive septum) is longer than the
others. Two cycles of simple trabecular septa can
be expressed in the corallites, although further
cycles may be apparent in the walls. The walls,
regarded as synapticulothecac (Wells 1956) have
recently been shown to have septal contribution
(Ricart y Menendez and Freidman 1977). See
plate 43 and Figure 1 for a summary of these
features).
The axial corallites are usually described by
measurements, (in this paper expressed as inner
and outer diameter) and by septal development
(here expressed as maximum observed develop-
ment, to an approximate fraction of the radius,
e.g. to 1/3R). The radial corallites are described
according to their shapes, summarized in Figure
2 .
‘Coenosteum’ as described for Acropora
commonly refers to both external corallite and
intercoraliite features. The walls of corallites are
often clearly costate, in which case the
intercoraliite material is usually a spongy
reticulum, with or without spines. Sometimes
there is little or no difference between the
appearance of the two areas. This is usually when
the coenosteum is a dense arrangement of
elaborated spines. With electron microscopy i
different categories of spines can be seen and the '
coenosteum can be more accurately described j
(Wallace and Grimmer, in preparation). j
i
Variability
All Scleractina are capable of broad mor-
phological variety, but in Acropora this can be
caespitose caespito-corymbose table plate
Fig. 1: Common colony shapes in Great Barrier Reef Acropora and their nomenclature.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
275
particularly subtle. A number of features
contribute: the axial method of growth allows a
continuum of shape possibilities, which may be
exploited differently by the species, the popula-
tion, the colony, or even parts of the colony. The
simplified skeletal structure allows the application
of only a limited descriptive terminology, and
description of small variations is always difficult.
Detailed measurements for analysis of variation
often show similar variability within the colony to
that in the entire population (Wallace, unpub-
lished, for A. millepora and A. aspera; similarly
high variability in Porites is shown by Brakel
1977). In fact the absolute variation in size of
most skeletal features within the entire genus is
relatively small for a coral group.
In this much speciated genus there is a tendency
for numbers of species to co-occur. Often
differences between species can be seen in
population strategies and slight differences in
habitat preference.
Despite the above, some of the taxonomic
confusion so repeatedly quoted in the literature
disappears when the taxonomist enters the water.
This is particularly so with reef slope species
where growth form and other features can often
be seen to alter gradually with depth. It can be
staled unequivocally that the most complex
species laxonomically are those occurring on the
shallow reef flat, and none of these can be
considered to be fully described in the present
paper.
Some generalizations can be made, and should
be born in mind when interpreting the taxonomic
section.
(1) Some reef regions support predominantly
characteristic colony shapes, which can be
attributed to physical parameters (in
particular low-profile corymbose shapes on
the outer reef flat where exposure is great
(low water depth); small flat plates on the
deep reef slope where light availability is
limited). In these areas the species
composition can be expected to include (a)
species capable of only that shape; (b)
species having the shape within a range of
phenotypic variability; (c) morphs of
polymorphic species.
(2) All species which extend down the reef
slope exhibit a gradual flattening out of
shape with depth, and the ratio of radial
to axial corallite numbers decreases. These
flat colonies can be recognised for what
they are if the observer investigates
shallower sections of the populations. (In
skeletal collections they are more difficult
to sort). A few species are characteristically
flat but with high radial to axial corallite
ratio, and do not extend into deep water.
At least one species {A. granulosa) is
apparently adapted to low-light situations,
and has a characteristic distribution which
includes shallow but shaded situations.
tubular, round opening tubular,
oblique opening
tubular tubular, tubular, nariform
appressed dimidiate opening opening
(tubo-nariformJ
tubo-nariform dimidiate
with rostrate
development
(front)
lipped
(labellate)
flaring lip
cochleariform sub-immersed
immersed
Fig. 2; Radial corallite shapes in Great Barrier Reef Acropora and their nomenclature.
276
MEMOIRS OF THE QUEENSLAND MUSEUM
(3) Some basically arborescent species which
occur on the reef flat may exhibit diverse
growth forms adapted to different
habitats.
(4) The greatest variety of colony shape seems
to occur in areas with good water cover and
reasonable water circulation, such as deep
‘middle reef flat' areas and leeward patch
reefs. Many species can be thought of as
achieving their most ‘characteristic’ shape
in these areas. Here also many colonies
achieve colossal size.
Many species have characteristic colouration, at
least locally, which can be used as a guide to
identification but it should never be supposed that
this is the only possible colour for the species.
Methods
Field Studies: The observations in this paper
are the result of a variety of studies on reefs in
the Centra! and Southern Great Barrier Reef
Province (sec Maxwell 1968). They concentrate
on reef slope and other below-L.W.M. areas, as
the corals here have previously been least
observed. On Big Broadhurst Reef (18°55’S;
147o44^E) observations and collections were made
serially across the outer reef flat and down the
reef slope (see Wallace 1975; Wallace and Dale
1977). Similar techniques were used on the reef
Oat at Redbill Island {20^57’S; 150"05’E)
(Wallace, unpublished). Collections with habitat
data were made at Bushy-Redbill Reef (see
Wallace and Lovell 1977), Darlcy Reef (19'^24’S;
148°05’E), Bowden Reef (19^02'S; 147^06’E),
Prawn Reef (19“02’S; 148°05’E), Viper Reef
(18°53’S; 148°10’E), Tryon Reef (23°15’S;
151^47’E), Lady Musgrave Reef (23°55’S;'
152°24’E), and Heron Island reef (23°27’S;
151°55’E). Detailed samples of populations of A,
miilepora and A. aspera for a morphometric study
were made at Heron Island. A reference collection
was made in the Fiji Islands, type locality of many
of the species. Occasional reference is also made
to observations by the author at Enewetak,
Marshall Islands and Tongatabu, Tongan Islands.
A recent opportunity to study specimens at James
Cook University (JCU) collected at Lizard Island
(14^40’S; 145''28’E) and the Palm Island group
(18‘"40'S; 146°33’E) allowed further notes to be
added for some species.
Other workers contributed specimens from
various parts of the Great Barrier Reef Province
and the Murray Islands. Information on this
material is given to the text, and locations of reefs
can be seen in Maxwell (1968, figs. 17A-D).
Other material examined in the Queensland
Museum included the collections of the Great
Barrier Reef Committee (G.B.R.C.) and
specimens mentioned by Stephenson and Wells
(1956).
Type Material: Types were studied in the
Smithsonian Institution (USNM), Yale Peabody
Museum (YPM), British Museum of Natural
History (BM), Mu.seurn National d’Histoire
Naturellc, Paris (PM), University of the
Philippines (UP). Fragments of type material
were received on loan from the first three
institutions, and photographs of type specimens
were received from the Museum fiir Naturkunde,
Berlin (MNB), the USNM and the BM.
Collections were examined at the Rijksmuseum
van Naturlijke Historie, Leiden (Verwey collec-
tions), Zoological Museum Amsterdam, and the
University of the South Pacific, Fiji.
Laboratory Studies: Most specimens were
stored as cleaned skeletons (preferably macerated
and cleaned in water). Small fragments of some
specimens have been preserved entire in buffered
formalin. Observations on skeletons were made
with a Wild M5 binocular microscope with linear
measuring eyepiece. W'here possible, particularly
with type material, five observations per character
were made for each specimen, but expediency j
prevented this procedure being used for all (
specimens. Additionally, representative specimens
of each species were examined with scanning
electron microscopy, and a standardized
procedure for photographing axial and radial
corailites and inter-corallite spines was followed.
This information is sometimes used to clarify
descriptions, but it is mainly to be used for a study
further defining species groups and pursuing a
phylogenetic grouping of them (Wallace and
Grimmer, in preparation).
Use of the Text
Any taxonomic text has two main categories of
readers: those interested in the taxonomic
interpretation as such, and those requiring an
identification tool. For this genus, I suspect the
second category will be in the majority. I
recommend that no attempt be made to identify
single specimens without field observations, and
conversely that field work (even when done
without taxonomic emphasis) include casual
assessment of affinities of colonies. Any field notes
which allow specimens to be later grouped as
series are useful.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
277
For observing skeletal specimens with an
incident light microscope a finger held between
the light source and the specimen (to diffuse the
light) enables corallite shape and septal features
to be viewed clearly.
The photographic plates are considered to form
a major part of the taxonomic description, because
they (1) display field appearance of colony and
habitat, (2) indicate size and shape of corallites,
and where possible coenosteai texture, (3) indicate
variety in all features. A fourth and most
important role intended for the plates is that of
a visual key. The identification process will be
simplified if the user scans the plates before
attempting to identify material. Reference to plate
numbers is made once only for each
description.
Where synonymy lists of other authors have
been accepted without investigation these are
quoted as (synonymy) after the author reference.
Check-lists and catalogues (e.g. Ralhbun 1887)
are not used for synonymy lists. In this context
Faustino (1927), being mainly a reiteration of
previous descriptions, is regarded as a check list
unless it makes an addition to information on a
species.
The arrangement of species is partly in
accordance with previous arrangements, and
reflects my own views of affinities. Strongly
supported affinities are grouped as species-groups.
As both detailed morphological and phylogenetic
affinities of species will be pursued in a later paper
this arrangement must be considered a working
guide only.
One species group, that including A. palifera
(Lamarck) (i.e. the subgenus Isopora of Brook,
1893), has been omitted as this is under scrutiny
in population studies by Potts (1976, 1977) and
will be given separate taxonomic treatment (Potts
and Wallace, in preparation).
Specimens quoted in the text are registered
specimens, and in most cases further unregistered
material and spirit specimens are located in the
Queensland Museum (QM). Smalt reference
collections will be lodged in the USNM, BM, and
MNB.
Depth ranges recorded for species apply to the
areas studied, and these may differ in other
areas.
Finally, some of the taxonomic format used by
other authors has been omitted. For example,
many authors describe undersurface details of
colonies. As these depend on general colony shape
and are reductions of upper surface features their
usefulness is questionable. Radial corallite
measurements are not given, as a statistically
acceptable assessment of these which is compar-
able across different shapes has not been arrived
at (the illustrations and previous authors’
descriptions give an indication of size range).
SYSTEMATICS
Family ACROPORIDAE Verrill, 1901
Acroporidae Verrill, 1901, p.I63; Wells, 1936, p.99;
Wells, 1956. P.F372.
Massive or ramose colonies by extralentacular
budding; hermatypic. Corallites small, synap-
ticulolhecale or septothecate, pseudocostate or
costate, slightly differentiated from coenostcum.
Septa non-exsert, in 2 cycles, formed by simple
spiniform trabeculae projecting inw'ard and
upward from vertical mural trabeculae, commonly
fusing to form laminae. Columella absent or
trabecular and weak. Dissepiments thin and
tabular when developed. Coenostcum extensive,
light reticulate, flaky, generally spinose or striate
on surface. (Modified from Wells 1956,
P.F372).
Genus Acropora Oken, 1815
Acropora Oken, 1815, p.66; Verrill, 1901, p.l64;
Verrill 1902, p.208 (synonymy); Vaughan, 1918,
p.l59; Wells, 1956, p.F373 (synonymy); Nemenzo,
1967, p.47.
Typi Spi-CII S: Millepora muricata Linnaeus, 1758.
Acroporidae which are ramose, rarely massive
or encrusting; branching with an axial or leading
corallite larger than the more numerous radial
corallites budded from it; united by light,
reticulate, spinose or pscudocostate or costale
coenostcum. Columella and dissepiments absent.
(Modified from Wells 1956, p.F374).
The ‘Acropora robusta' GROUP
The four species Acropora palmerae. A.
robusta. A. rotumana, and A. abrotanoides are
very similar in corallite morphology but not colony
shape. A. palmerae, described as a completely
encrusting species (Wells 1954) can bear short
vertical branches (Wallace, in prep.). A.
rotumana is stalked, with thick alciform
branching units or even forming a sturdy
arborescent table. A. robusta can vary phenoty-
pically from encrusting to open branching. A,
abrotanoides, still poorly categorized, occurs as
clumps of mainly vertical branches.
278
MEMOIRS OF THE QUEENSLAND MUSEUM
Ignoring this last species, the other species have
strong habitat preferences: A. palmerae for rocky
(typically algal ridge) locations; A. rotumana for
deeper-water reef flat areas, A. robusta for the
shallow outer reef. On Enewetak atoll in the
Marshall islands A. palmerae and A, rotumana
arc very abundant and A. robusta rare {Wallace,
in prep.). On the Central Great Barrier Reef
where there is no distinct algal ridge, A. palmerae
is absent, any encrusting colonies being recognis-
able as members of an A. robusta population: A.
rotumana appears as a rare species.
Within each species there is also a phenotypic
component to radial corallitc variation: the radials
are best developed on the most freely branching
parts of the colony.
These features, and the added fact that the
clumsy colonics never lend themselves to the
collection of a truly representative sample, have
led to the naming of many species from single or
few fragments. As well as the species placed in
synonymy here, a number of more obscure
species-names possibly belong with these species,
e.g. A. cyclopea (Dana), A. cuspidata (Dana), A.
conigera (Dana), A. smithi (Brook). With the
current lack of geographic studies on Acropora it
is difficult and unwise to lump these.
The separation of the three species treated here
is maintained on the basis of their apparent
distinctness in Great Barrier Reef populations,
and pending further regional studies.
Acropora robusta (Dana, 1846)
(Plates 44, 45)
Madrepora robusta Dana. 1846, p.475, pi. 39, fig.3, 3a,
pi. 31, figs. 3a-c: Brook, 1893, p.42 (synonymy).
Madrepora pacifica Brook, 1891, p.465; 1893, p.39,
pi. 30, fig.B.
Acropora pacifica: Crossland, 1952. p.202, pl.31, fig. 2,
pi. 32, fig. 2; Nemenzo, 1967, p.54, pM8, fig.4.
Madrepora decipiens Brook, 1892, p.456; 1893, p.51,
pi. 14, figs. B to D.
Acropora decipiens: Vaughan, 1918, p.l65, pi. 67, figs.
2, 2a, 2b; Nemenzo, 1967, p.60.
Material Examined
USNM: Fiji Islands, U.S. Expl. Exped., A. robusta
holotype 297.
BM: Samoa, Rev. Whitmore, A. pacifica holotype
1875.10.2.13.’ Rocky Is., G.B.R. Saville Kent A.
decipiens syntypes 1892.6.8.82; 1892.6.8.85.
QM: Big Broadhurst Reef: SW. side, reef slope: 4m,
27.111.1973, C.W., G10190; 2-2m, Il.x.1973, C.W.,
G10250; l-5m, 18.x. 1973, C.W., G10191; Im,
26.111.1973, C.W,, G10194; SW. side. Outer reef flat:
Il.x.1973, C.W.. G10246. 18.x. 1973, C.W., G10248;
22.X.1973. C.W., G10247.
Bushy-RedbilLReef; SE. side, reef crest, 27.xii.1972,
C.W., G10193; S side, reef crest; 27.xii.1972, E. Lovell.
G10I92; 25.xii.l972, E. Lovell, GI0257; W. side, reef
crest, 22.xii.1972, C.W'., G 10258; W. side, reef slope,
I9.xii.l972, E. Lovell, G10259; NW. side, reef crest,
30.xii.l972, C.W., G10I93; NNE side, reef slope,
30.xii.l972, C.W., G10193; NNE side, reef slope,
30.xii.l972, C.W., G10197.
Bowden Reef:' SW. side, reef crest, 15.vii.l972, C.W.,
G 101 98, G 10199; slope of opening in SW. side,
26.vii.1972, C.W., GI0256.
Bramble Cay: Dec. 1924, C. Hedley, G 10201; Oct.
1924, C. Hedley, G 1 0262.
Coates Reef. 13.viii.l924. ‘Geranium’, G10261.
Feather Reef, ILviii.1924, ‘Geranium’, G10260.
Flinders Reef, Moreton Bay. W. side, 6.ix.l973, C.W.,
G 10200.
Masthead Reef, NNE. side, reef crest, August 1974,
J. Buhman, Gi0254, G10255.
Field Diagnosis
Colonies have stout branches which can be
vertical cones, distorted humps, or freely
branching horizontal units on different parts of
the same colony. Sections of the colony can be
completely encrusting. Those parts of the colony
on solid substrate have the solid vertical cones, but
these extend horizontally and branch freely where
small holes in the reef, or the reef edge itself,
allow. The low, distorted rounded humps are
formed in very shallow (e.g. reef crest) areas.
Prominent tubular radial corallites mixed with
shorter ones are obvious at the tips of the cones
and branches. Common colours are bright green
with deep pink branch-tips and pink-brown,
yellow-brown or cream.
Laboratory Diagnosis
Laboratory difficulties occur because usually
only a piece of the heavy colony is collected. Two
very different fragments are illustrated in Plate
45 figs, a and c. Crossland’s plate 32, fig.2, shows
another shape the branches can take.
Branching pattern: This is described above. The
free branches are usually 20 to 35 mm diameter,
but branchlets as narrow as 10 mm can occur;
maximum length measured is 250 mm. The cones
may be as thick as 40 mm at the base. The humps
are either solid or encrusting, taking the shape of
the substrate.
Axial corallites: Outer diameter 2-5 to 3 0 mm;
inner diameter L2 to 1-5 mm. Septation:
primaries to 3/4R, secondaries to I /4R, usually
incomplete. The axial corallites are not strongly
different from large radial corallites, and on the
rounded humps they may be absent altogether.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
279
Radial corallites: Tall and short radial corallites
mixed. The differences between the two are
emphasised at the growing tips where the tall
radials may be 5 mm long. On the low humps the
differences may disappear entirely. The tall
radials are tubular, with rounded, oblique or
dimidiate openings. Their angle of extension from
the branch decreases from 90*^ at the base to about
45° at the tips. The short radials usually have only
a small section of outer wall present. Septation
is usually well-developed in the tall radials
(primaries to 1 /2R, secondaries to 1 /3R), and less
developed in the short radials (sometimes only the
directives visible).
Coenosteum: Costate on corallites, reticulate
with simple spines in between.
Within Reef Distribution
The species occurs only in the shallow outer
flat-reef crest area, and flourishes at the
windward (SE.) end of the reefs.
Identification Difficulties and History
In the field this species is conspicuous and
distinctive. In the laboratory the colony is poorly
represented by fragments and there is the
temptation to find a ‘best fit’ with one of the many
described robust species, few of which have been
described from the field. Only the most certain
synonyms have been included; some of the robust
species mentioned in the introduction may also be
this species. The more slender free-growing
branches resemble A. intermedia, and specimens
of the two species can be confused in the
laboratory (but never in the field). A. palmerae
Wells may be synonymous. An encrusting colony
of A. robusta with few vertical branchlets is
illustrated in colour in Roughly 1936 (plate 6
upper).
Geographic Distribution
Indo-Pacific: China Sea, Philippines, Great
Barrier Reef, Fiji Islands, Samoa, Marshall
Islands (C.W.).
Acropora rotumana (Gardiner, 1898)
(Plate 46)
Madrepora rotumana Gardiner, 1898, p.258, pi. 23,
fig.2.
Acropora rotumana: Hoffmeister, 1925, p.69; Wells,
1954, p.419, p.ll2, rig.1-3, pl.ll3, figs.4, 5.
Acropora tutuilensis (part) Hoffmeister, 1925,
p.7L
Material Examined
USNM: Pago Pago Harbor, Tutuila, Samoa A.
tutuilensis no. 4 (mentioned Hoffmeister, 1925),
Rongerik Atoll 44477; Rongerlap Atoll 44478; Bikini
Atoll. 44469, 44470, 44472, 44473, 44474, Enewetak
Atoll 44475: A. rotumana (mentioned Wells, 1954).
QM: Big Broadhurst Reef: SW. side, outer reef flat:
14.X.1973, C.W., GI0233; 22.X.1973, C.W., G10234,
G10235; E. side, reef slope, Im, 21.X.1973, C.W.,
G10236.
Bowden Reef: SW. side, reef crest, 15.vii.l972, C.W.,
G10263.
Field Diagnosis
Mature colonies are about 1-5 m in height and
width. They have a sturdy tree-like appearance
due to the small number of very stout branches
which have some proximal vertical growth
component, then become horizontal for most of
their extent, and proliferate into small branchlets
at the tips. This is not the full range of shapes
possible in the species. At Enewetak Atoll, for
example, it can form a stalked table (Wallace, in
preparation). Colours: deep pinkish-brown or
apple green. (See Gardiner 1898 for detailed field
diagnosis).
Laboratory Diagnosis
The fragments collected from this species are
usually branch-tips with a greater or lesser
amount of branch remaining. They often have a
‘distorted' appearance, due to the irregular
secondary branching pattern.
Branching pattern: described above.
Axial corallites: Outer diameter 2-0 to 2-5 mm
(in the G.B.R. specimens); inner diameter 0-8 to
1-2 mm. Septation: primaries present to 1/2R, few
secondaries, up to 1 /4R.
Radial corallites: Tall and short radial corallites
mixed. Tall radials extend from branch at 45° to
90° and are tubular with round, oblique, dimidiate
or nariform openings. Short radials usually have
less than half the wall developed, and range down
to immersed. Primary septa not well developed,
up to 1/3R; secondaries absent or a few just
visible.
Coenosteum; Costate or pseudocostate on
radials, reticulate with simple spines in
between.
Within Reef Distribution
Occasional colonies found just below the reef
top on the upper reef terrace or in surge channel
openings on the outer reef. Elsewhere (e.g. on
Enewetak Atoll) it occurs in deep-water reef flat
situations.
Identification Difficulties and History
This is one of the few Acropora species to be
originally described from the field as well as from
280
MEMOIRS OF THE QUEENSLAND MUSEUM
skeletal material. Both Hoffmeister and Wells
regard it as a good species because of its
distinctive colony shape. Gardiner states it to be
‘by far the most abundant coral on the reefs of
Rotuma’. It cannot be distinguished from A.
robusta or A. abrotoftoides on radial corallite
shape.
Geographic Distribution
Pacific Ocean: Great Barrier Reef, Rotuma,
Samoa, Marshall Islands.
Acropora abrotanoides (Lamarck, 1816)
(Plate 47)
Madrepora abrotanoides Lamarck, 1816, p.280;
Brook, 1893, p.56 (synonymy).
Acropora abrotanoides: Crossland, 1952, p.204;
Wells, 1954, p.418, pl.l23, figs. 1, 2.
non Acropora abrotanoides: Vaughan, 1918, p.I66,
pi. 68, fig. 2.
Madrepora deformis [non Michelinl; Dana, 1846,
p.484, pL43, fig. 1.
Madrepora danai Milne Edwards and Haime, 1860,
p.560.
Madrepora danae: Brook, 1893, p.57 (synonymy).
Acropora danai: Wells, 1954, p.418, pi. Ill, figs.
4-6.
Materiai. Examined
USNM: Tahiti, U.S. Exp). Exped., A. danai holotype
303 (M. deformis of Dana); Marshall Islands, J. W.
Wells. A. danai 45175 (mentioned Wells 1954).
YPM; Tahiti, fragment of type (labelled A. deformis)
4162 (mentioned Verrill, 1864)
QM: Big Broadhurst Reef, patch reef in lagoon,
28.iii.1973, C.W., G10238.
Bushy-Rcdbill Reef, NW. side, reef crest, 22.xii.1972,
C.W., G 10239.
Bowden Reef, slope of opening in SW. side,
26.vii.1972, C. W., G10240-2.
Viper Reef, patch reefs in lagoon, L5m., 16.vii.l972,
C. W., G10243.
Great Astrolabe Reefs, Fiji Islands, 1974, C. W.,
G 10244.
Field Diagnosis
Colonies occur as tufts of sturdy vertical to
oblique branches with appearance roughened by
the presence of numerous incipient branchlets.
Colour: yellow-brown.
LABORATORY DIAGNOSIS
Branching pattern (taken from five specimens):
From a basal area (usually dead) a small number
of sturdy branching units is given off. Each
branching unit has a small number of main
branches which are vertical or almost so, and the
branches may branch again. In addition, each
branch is roughened by the presence of incipient
branches. The longest branch is 150 mm, and the
widest 40 mm.
Axial corallites (on main branches): Outer
diameter 2-5 to 3-8 mm; inner diameter 0-8 to 1-2
mm. Scptation: primaries present to 1/3R,
secondaries sometimes complete to I/4R.
Radial corallites: Tall and short radials mixed.
Tall radials tubular with round, oblique or
dimidiate openings. Those with round openings
usually can be recognised as incipient axials,
themselves budding radials from their
circumference.
Coenosteum: Costate on radials, reticulate with
simple spines in between.
Within Reef Distribution
This species seems to be restricted to upper reef
areas, mainly the outer reef flat and crest, and
the tops of patch reefs in deep lagoons.
Identification Difficulties and History
Brook describes his specimens in detail, and
apparently examined Lamarck’s type, which I
have been unable to locate. AH other authors had
meagre or immature specimens (e.g. Vaughan’s
Three small immature specimens'). There are no
morphological characters supporting the separa-
tion of A. danai. A. listen (Brook) may also be
a synonym, but its validity could be tested by a
study on the Tongan reefs.
This species still remains categorized only as a
group of specimens, with no field identity. The
fact that it eludes categorization in the field
suggests that it is not a ‘good’ species. The
abundance of incipient branches is a poor species
character, but this is at present the main feature
defining this species.
Geographic Distribution
Indo-Pacific: Singapore, Great Barrier Reef,
Marshall Islands, Tahiti.
Acropora intermedia (Brook, 1891)
(Plates 48, 49)
Madrepora intermedia Brook, 1891, p.463; 1893,
p.3L pl.l, fig. C.
Acropora intermedia: Crossland, 1952, p.200, pi. 32,
fig. 1; Stephenson and Wells, 1956, p.l6; Pillai and
Schcer, 1976, p.24, pi. 2, fig. 1.
Material Examined
BM: Maidive Islands, A. intermedia syntype
1886.11.22.6.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
281
QM: Big Broadhurst Reef, SW. side, reef slope: 2m,
11.X.1973. C. W., GIMOO; 2m, 26.iii.1973, C. W.,
G1 1295; 3m, 28.iii.1973, C.W., GI1310; 5m, 25.ii.1973,
C.W., GI1302; 5.3m, 13.x. 1973, C.W., Gn576; 6m,
25.iii.1973, C. W., G1I301; 6 1m, 13.X.1973, C. W.,
Gn582; 6-3m, 13.X.1973. C. W., G1I579; 7-6m,
13.x. 1973, C. W., G1I575; 7-7m, I4.X.1973, C. W.,
Gl 1578, Gil 580; 8m, 26.iii.1973, C. W., Gl 1300; 8-3m,
I5.X.1973, C. W., 011581; 8-6m, 15.X.1973, C. W.,
011577; SW. side, outer reef flat: 1 Lx. 1973, C. W.,
G11308, G11583; 22.X.1973, C. W., G1 1327, G11584;
SW. side, floor of surge channel opening: 20.x. 1973, C.
W., G11307, GU309, GI 1585.
Bowden Reef, SW. side, upper reef slope, 26.vii.1972,
C. W., G11316.
Bushy-Redbill Reef, S. side, reef slope, 3m,
27.xii.1972, C. W., G11311; adjacent Redbill Is., reef
crest, 18.xii.l972, C. W., G11305; NW. side, first reef
crest, 22.xii.1972, C. W., G11312, G11313; NNE. side,
reef slope, Jan. 1973, C.W.. G11391.
Heron Is., W. side, reef flat, 6.vii.l973, Y. Loya,
G11297.
Field Diagnosis
Colonies vary from small clumps to small
thickets of around 3m diameter and are
sturdy-branched arborescent. The radial corallites
are a mixture of tall and short tubular and are
obvious under water. Colours are cream to pale
brown, pale green or greenish brown or bright
blue.
Laboratory Diagnosis
Branching pattern: Branches are given off at
wide angles (45° to 90°) and branch width varies
from 15 to 25 mm. Branches usually taper
strongly.
Axial corallites: From barely exert to 2 mm
exert. Outer diameter 3 0 to 4-0 mm (in QM
specimens); inner diameter 11 to 1-5 mm.
Septation: primaries well developed, often
reaching 3/4R, secondaries usually present, up to
1/4R.
Radial corallites: Tall and short radial corallites
are mixed. The tall radial corallites are tubular,
up to 5 mm long, with round, oval or dimidiate
openings and usually extend from the branches at
90° to 45°. Lower wall may be slightly thickened.
Primary septa are developed up to 2/3R, and
secondaries present, up to 1/4R. The smaller
radials are tubular appressed or emergent, or
sub-immersed, and oriented with their openings
facing in random directions to the branch.
Coenosteum: Strongly costate on radial
corallites, open reticulate with simple spines
between radials.
Within Reef Distribution
Middle and outer reef flat, upper reef slope to
about 12 m, surge channel floors and sandy
bottoms around patch reefs.
Identification Difficulties and History
This species appears to be morphologically
similar in radial corallite and coenosleal features
to the A. robusta group. Many specimens are
difficult to place exactly in either this species, A.
formosa or A. grandis, and clearly all three
species require further field study.
Geographic Distribution
Indo-Pacific: Maldives, Great Barrier Reef.
Acropora grandis (Brook, 1892)
(Plate 50 A,B)
Madrepora grandis Brook, 1892, p.457; 1893, p.42,
pi. I, figs. A, B.
Acropora grandis: Crossland, 1952, p.202, pl.31, figs.
1, 3, 4,
Material Examined
BM: Palm Island, Saville-Kent, A. grandis syntype
1892.6.8.60; Herring Island, Bowen, Saville-Kent. A.
grandis 1892.6. 8. 314; Rocky Island, Saville-Kent, A.
grandis var. 1892.6.8.261.
QM; Big Broadhurst Reef, SW. side, reef slope: 6m,
25.iii.1973, C.W., 011320; 6-Im, I3.X.I973, C.W.,
GII366; 7m, 27.iii.1973, C.W., G11299; 7-lm,
13.x. 1973, C.W., GI1367; 7-7m, 14.x. 1973, 011368.
Bushy-Redbill Reef, NW, side, first reef crest,
22.xii.1972, C.W., 011306, G1I315, G11317.
Darley Reef, patch reef in lagoon, 4 m, 24.iii.1973,
C.W., 011298.
Field Diagnosis
Openly arborescent colonies have scattered
radial corallites of mixed size with large obvious
openings, and may be brown, bright blue, bright
purple or even greenish-brown with blue tips.
Laboratory Diagnosis
Branching pattern: Branches commonly spread
at from 60° to 40° . Branches are from 10 to
30 mm thick, and may be as long as 30 cm
without branching. They usually taper
gradually.
Axial corallites: Up to 3 mm exert; outer
diameter 2-5 to 3-5 mm; inner diameter 1-0 to
1-7 mm. Septation: both cycles may be present,
the primary septa up to 1/3R, the secondary up
to 1/4R.
Radial corallites: Two sizes of radial corallite
occur. Towards the branch tips the large radials
reach 3 mm length, but elsewhere they are much
282
MEMOIRS OFTHE QUEENSLAND MUSEUM
shorter than this. The openings are round to oval,
and are directed straight out from the branch, or
nearly so. The septa are poorly developed: both
cycles may be completely absent, or primary sepia
may be present up to 1/4R, plus a few
secondaries.
Coenostcum: The surface has a light, crumbly
appearance, the coenostcum being costate or
reticulate on the radial corallites and open
reticulate between.
Identification Difficulties and History
This species has only been recorded from the
Great Barrier Reef, and in my experience it is not
common on the outer reefs, but is more
characteristic of the fringing reefs of the
continental islands. Crossland treats the species in
detail. Although this species is morphologically
different from A. formosa and A. intermedia in
many features, there remains a suspicion that it
may be related to one or other of these, and all
three species require further study in the field.
Geographic Distribution
Recorded from the Great Barrier Reef only.
Acropora formosa (Dana, 1846)
(Plates 51, 52)
Madrepora formosa Dana, 1846, p.473, pl.3I, fig. 2a,
2b, pi. 38, rig.4.
Acropora formosa: Hoffmeister. 1925, p.55. pi. 8,
figs. I -3 (Synonymy); Wells, 1950, p.35; 1954.
p.4I5, pi. 102, figs.1-9. pi. 103. figs.I-5, pi. 104, fig.4
(synonymy); Stephenson and Wells, 1956, p.14
(synonymy); Ncmenzo, 1967. p.6l, pl.21, fig.3;
Pillai and Schecr, 1974, p.453; Pillai and Scheer,
1976, p.23.
Madrepora hrachiata Dana. 1846, p.474, pi. 38, fig.3,
3a, 3b.
Madrepora gracilis Dana, 1846, p.482, pi. 41. fig 3
3a, 3b.
Madrepora nohilis Dana, 1846. p.48l, pi. 40, fig.3,
3a.
Acropora nohilis: Hoffmeister, 1925, p.59, pl.l I, fig.l
(synonymy); Wells, 1954, p.416. pi. 104, figs.l, 2;
Nemenzo, 1967, p.62, pl.21, fig. 5.
non Acropora nohilis: Pillai and Scheer. 1974, p.453.
figJc.
Acropora laevis (part) Crossland, 1952. p.230, pi. 45,
figs.l, 2.
M \ 1 1 Ki \i E\ \\ii\i I)
USNM: Fiji Islands. U.S. Expl. Exped., A. formosa
syntypes 888, 282; Fiji Islands, U.S. Expl. Exped., A.
gracilis holotype 333; Sooloo Sea, U.S. Expl. Exped.,
A. hrachiata holotype 295; Singapore, U.S. Expl.
Exped., A. nohilis holotype 427.
QM: Big Broadhurst Reef: SW. side, reef slope:
1-6 m, Oct. 1973. C.W.. GI1388: 2 m, 20.x. 1973,
C.W.. G1I339; 2 m, II.x.1973, C.W., GI1343; 2 m!
26.iii.l973, C.W., GI1398, G11405; 2-3 m, 20 x 1973
C.W., GI1349, GII353, G11360; 2-6 m. 22.x.l973i
C.W., G1I346; 2-8 m, I2.X.1973, C.W., G11356;
3-8 m. 12.x. 1973, CAV.. GII335, GII347: GII362;’
5- 3 m, 13.X.1973, C.W.. GI1365; 5-7 m.
23.x. 1973, C.W.. GII348, GII364; 6 m, 25.iii.I973,
C.W„ GII320; 6 m, 13.X.I973, C.W., G11336;
6- 1 m, 13. X. 1973, C.W.. GI1363, G1I373, 6-3 m,’
20.x. 1973. C.W., GI1334: 6-3 m. 23.x. 1973, C.W.,
GI1361; 6-3 m, 13.x. 1973, C.W., G11369, G11371;
7- 6 m, I3.X.1973, C.W., GII367; 7-7 m, 14x1973
C.W.. GI1333, G11341, GI]368, G1I372: 7-8 m’
I4.X.1973, C.W., GII330. GI1337, Gl]358, Gn37o!
8- 1 m, 22.x. 1973, C.W., GII340; 8-7 m. I5.X.1973,
C.W., G11352; 12-4 m, 23.X.1973. C.W., GII363;
12-5 rn, 17. x. 1973, C.W., Gl 1350, G1 1355; 28.iii.l973i
C.W,, G 1 1396; SW side, outer reef flat: 20.x. 1973,
C.W., GI1331, G1I354; ll.x.1973, C.W., G1I359;
1 8.x. 1973, C.W., Gl 1344; SW. side, reef crest: 1-3 m,
22. x. 1973, C.W., GII345; 1-3 m, 23.x. 1973, C.W.]
G11351; SW. .side, surge channel, 7-5 m 20x 1973
C.W., GI1342.
Bowden Reef: SW. side, upper reef slope: 26.vii.1972,
C.W., GII318. GI13I9. Gl]322, GII323, GI1325;
floor of opening in SW. side, 5 m, 26.vii 1972 C w'
GI1380, G1138I, 011375.
Bushy-Redbill Reef: NW. side, first reef crest:
23. xii.t972, E. Lovell, Gl 1374, G11376, 01 1377;
22.xii.l972. C.W., GII306, G1I378, G1I379, G1I384
G11385.
Darley Reef, patch reef in lagoon: 1 m, 24.iii.1973,
C.W., GII401; 4 m, 22.iii.1973, C.W., GI1404.
Viper Reef, .luly 1972, C.W., GI1326, G11394.
Field Diagnosis
This is the common, thicket-forming arbores-
cent species with numerous small tubular radial
corallites. commonly coloured cream to brown,
less often bright blue or blue-tipped.
Laboratory Diagnosis
Branching pattern: Branching is always openly
arborescent. Branch diameters vary from slender
(10 mm) to thick (35 mm).
Axial corallites: From barely exert to 1-5 mm
exert. Outer diameter 2-0 to 2-8 mm; inner
diameter 0-8 to 1-2 mm. Septation: Primary septa
well developed, up to 1/2R; secondary septa
usually present, but poorly developed, up to
1/4R.
Radial corallites: Tubular, varying from
sub-immersed to 5 mm, prominent, with openings
from round to sharply oblique, the corallites
oriented at anything from 90° to the branch to
fully appressed. Septal development varies greatly,
but most commonly secondary septa are absent or
only poorly represented.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
283
Coenostcum: Coslate or neatly arranged simple
spines on radial corallites, reticulate with simple
spines between.
Within Reef Distribution
Reef crest and slope to position of surge channel
floor (10-12 m), lagoonal floors and lagoonal
patch reefs (may even form entire basis of
lagoonal patch reefs), middle reef flat, occasional
small colonies on outer reef flat, fringing
reefs.
Identification Difficulties And History
The interpretation that I have placed on this
species follows that of Wells (1954) and
Stephenson and Wells (1956) preceded by
Hoffmeislcr ( 1 925). The reader is referred to their
discussions. The species, as interpreted, may well
be a mixture and there is certainly no satisfactory
interpretation of variability (for example slender
and sturdy branched thickets may occur adjacent
to each other). There is some suggestion (J.
Collins, pers. comm.) that the sturdy branched
colonies may be old colonies. The colonies offer
no readily perceplable clues in the field. If there
are several species involved, the mosaic of their
variabilities may be adding to the confusion.
This species requires individual attention, and
the only assistance that can be offered at present
is in distinguishing from the field the other ‘easily
recognizable’ arborescent species; A. haimei, A.
microphthalma, A. intermedia, A. aspera, A.
pulchra and A, splendida. A. acuminata Verrill
does not seem to occur in the areas studied.
Geographic Distribution
Indo-Pacific: Ceylon, Cocos-Keeling, Amboina,
New Ireland, Great Barrier Reef and Torres
Straits, Fiji Islands, Samoa, Marshall Islands,
Tahiti. (Further Indian Ocean localities in Pillai
and Scheer, 1974).
Acropora splendida Nemenzo, 1967
(Plates 53, 54)
Acropora splendida Nemenzo, 1967, p.51, pi. 17,
fig-2.
Matkriai Examiniuj
UP: Hundred Islands, Pangasiman, 1959, A. de la
Cruz, holotype C931.
USP; Fiji Reefs 3916.
QM: Big Broadhursl Reef, SW. slope: 23-5 m,
17.x. 1973, C.W., G8715; 15 m, 16.x. 1973, C.W., G8716;
9-7 m, 15.x. 1973, C.W., G8714; 8 m, 28.iii.1973, C.W.,
G8704-6; 7-H m, 15.x. 1973, C.W., G87I3; 7-5 m,
15.x. 1973, C.W., G87I2; 7 m, 26.iii.l973. C.W., G870],
G8702; 6-9 m, 14.x. 1973, C.W., G8710, 08711; 6-6 m,
14.x. 1973, C.W., G8708, G8709; 6 m, 25.iii.1973, C.W.
G8699; 5 m. 26.iii.l973. C.W., G8700; 4-9 m,
13.X.I973, C.W., G8707; 3 m, 27.iii.1973. C.W.,
G8703.
Bowden Reef: Lagoon slope, Nov. 1972, R. Pearson,
G8696; floor of opening in SW. end of reef, 26.vii.1972,
C.W., G8697.
Darley Reef, patch reefs in lagoon, 4 m, C.W.,
G8698.
Field Diagnosis
Colonics arc large (commonly around Im
diameter) bowl to bracket shaped, the shape given
by long openly arborescent branches which curve
and divide to varying extent, depending on their
position within the colony. The bowl shape occurs
in specimens on flat surfaces and attachment
becomes lateral with steepness of the attaching
surface. With increasing depth the colonies
become flatter and smaller. A specimen collected
at 23-5 m (G8715) is completely horizontal.
Common colours are dark olive green with paler
green tips to the branches, dark blue with paler
blue tips, brown with pale blue lips, brown or
cream.
Laboratory Diagnosis
Branching pattern; As the colonies arc large,
fragments are usually collected. These are often
easily confused with other large arborescent
species. Specimens from the centre of the bracket
usually have long curving branches with little
secondary branching; those from the edge zone arc
more proliferous, with shorter branches and some
fusions, and may be flat underneath.
Axial corallites: Outer diameter 2*0 to 3*5 mm;
inner diameter 0-8 to 1-5 mm. Septation:
primaries to 2/3R, all or some secondaries present
to 1 /2R.
Radial corallites: Tubular, projecting from
branches at 90'" (less towards tips) with openings
round, oval, nariform, or dimidiate. Radials are
usually evenly sized and distributed but smaller
radials may be present amongst these, and their
openings may be variously directed. Septal
development varies, but primaries arc usually
present to 1/3R, and some secondaries visible.
Coenostcum: Usually neatly costate on radial
corallites, open reticulate with simple or laterally
flattened spines in between.
Within Reef Distribution
Reef slope from a few metres below crest to
limits of Acropora distribution (species no. 3 in
Wallace 1975); deep water lagoonal patch reefs;
fringing reefs.
284
MEMOIRS OF THE QUEENSLAND MUSEUM
Identification Difficulties and History
Nemcnzo’s species was based on a single
fragment, and he was not aware of the shape of
the colony. The species seems to have been
neglected in all other literature, although it is
possible that fragments may have been identified
with other arborescent species. This is the only
Acropora with an ‘arborescent bracket’ shape and
it is very distinctive in the field. Laboratory
specimens without field notes become unnecessari-
ly confused with other arborescent ‘problems’.
Geographic Distribution
Indo-Pacific; Philippines, Great Barrier Reef,
Fiji Islands, (common in Fijian fringing reefs,
from my personal observations), Palau (M.
Pichon, pers. comm.).
Acropora horrida (Dana, 1846)
(Plates 55, 56)
Madrepora horrida Dana, 1846, p.472, pi. 39, fig.2,
2a; Verrili, 1864, p.4I; Brook, 1893, p.l88
(synonymy).
Acropora horrida: Wells, 1954, p.417, pi. 107,
fig.l.
Madrepora angulaia Quelch, 1886, p.l60; Brook
1893,p.l95.
Madrepora inermis Brook, 1891, p.462; 1893, p.194,
pi. 29, figs. A, B.
Acropora inermis: Wells, 1954, p.43T
MaTI'RIAI EXAMlNtD
USNM; Fiji Islands, U.S. Expl. Exped., A. horrida
holotype 291.
YPM; Fiji Islands, U.S. Expl. Exped., A. horrida
fragment of type 2013.
BM; Zamboangana. ‘Challenger' Exped., A. angulata
holotype 1886.12.9.233; South Seas, purchased, A.
rVif'/'w/.y syntypes 1841.12.11.6, 1841.12.11.7.
QM: Big Broadhurst Reef: SW. side, reef slope: 7-3
m, I5.X.I973, C.W., G8755; 7-6 m, 23.X.I973, C.W.,
G8757, G8758; 5 m, 27.iii.1973, C.W., G8759: 8 m,
28.iii.1973, C.W., G8761; 4 m, 28.iii.l973, C.W.,
G8763; SW. .side, surge channel, 8-5 m, 20.x. 1973,
C.W.. G8756.
Bowden Reef: Slope of opening in SW_ side:
26.vii.1972, C.W., G8764-7; 1-5 m, 26.vii.1972, C.W.,
G8768; 1-2 m, 26.vii.I972, C.W., G8769; 25.vii.1972,
C.W., G8771; floor of opening in SW. side, 26.vii.I972,
C.W., G8770.
Bushy-Redbill Reef: NW. side, floor outside slope: 12
m, 30.xii.I972, C.W., G9076; 8 m, 3I.xii.l972, C.W.,
G9077; NW. side, middle reef flat, 22.xii.I972, C.W.,
G9078; 20.xii.l972, C.W., G9079; NW. side, reef
patches, 12 m, 14. vi. 1975, C.W., G9080; W. side, reef
slope: 8 m, 3.vi.l975, C.W., G9081; Dec. 1972, C.W.,
G8774; adjacent Redbill Is., reef slope, 5 m, 2.vi.l975
C.W., G9084.
Darley Reef, patch reef in lagoon: 22.iii.l973, C.W.,
G8773; 6 m, 22.iii.1973, C.W., G8762, 23.iii.1973,
C.W., G8760.
Heron Island, W. side, reef slope, July 1973, CW
G8773.
Fiji Islands, Great Astrolabe Reefs (Kadavu), W.side
of Yaukuvi Levu fringing reef, l.ii.l974, C.W.,
G9085.
Field Diagnosis
Occurs as sprawling arborescent to shrubby
patches. Branches are slender and have a ragged
appearance due to the scattered distribution of the
radials and their poorly-formed walls. The polyps
of this coral are usually extended, an unusual
feature in Acropora. The colour is usually light
powder-blue to light grey.
Laboratory Diagnosis
Branching pattern: Branching is primarily
open-arborescent. Specimens from loose substra-
tum areas tend towards horizontal primary
growth; those From firmer substrates are more
upright. Main branches are 7 to I 5 mm in width,
and taper gradually towards the tips. Secondary
branchlets are irregularly sized and placed. They
can be short, scattered twigs which alter the shape
of the colony little, or bundles of branchlets which
give a shrub-like appearance, or bottlebrush
formations of subequal length arranged all over
the branches.
Axial corallites; Outer diameter 1-6 to 2-3 mm;
inner diameter 0-6 to 1-2 mm. Septation: Primary
septa present, up to 2/3R, secondary cycle poorly
developed, usually a few septa just visible.
Radial corallites: Scattered, sub-immersed to
emergent tubular, sometimes appressed, with
round openings. Walls are thin and very fragile
around the opening. Primary septa developed, up
to 1/3R, secondary septa usually absent, or one
or two just visible.
Cocnosteum: Open reticulate with simple to
forked spines, both on corallites and between.
Within Reef Distribution
This species occurs where there is some constant
water cover, down to at least 15 m, and apparently
favours high sediment areas, i.e. deeper reef flat
areas, reef slopes, rubble and sandy bottom areas
around patch reefs and on surge channel fioors.
It occurs with A. vaughani in the deeper
areas.
Identification Difficulties and History
The species is distinctive in the field because
of the extended polyps, ragged appearance and
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
285
unusual colouring. The extended polyps also give
it a silky sheen. Laboratory differences from A.
vaughani are explained by Wells 1954 (p.417).
Brook’s A. inermis specimens arc apparently
slender branches of this species. Quelch’s A.
angulata is named from a single branch lip.
Geographic Distribution
Indo-Pacific: Philippines, Great Barrier Reef,
Fiji Islands, Marshall Islands.
Acropora vaughani (Wells 1954)
(Plate 57)
Acropora vaughani Wells, 1954, p.416, pi. 105, fig. 1,
pi. 106, figs. 1-8, pi. 107, figs. 2-6.
Material Examined
USNM; Bikini lagoon, 18 fms, holotype 44452;
Seaward slope, Bikini Atoll, 21-25 fms, paratype
44457.
QM: Big Broadhurst Reef, SW, side, reef slope: 9-8m,
16.X.1973, C.W., G10264; 8Tm, 14.X.1973, C.W.,
G10265; 7m, 13.x. 1973, C.W., G10266; 6-lm,
13.X.1973, C.W., G10267, G10268, 3m, 27.iii.1973,
C.W., G10269; 7m, 28.iii.l973, C.W., G10270; 8m,
28.iii.I973, C.W., G10271.
Bushy-Redbill Reef, NNE. side, off-reef floor, 12m,
30.xii.l972, C.W., G10272.
Field Diagnosis
Occurs as scattered ‘shrubby arborescent’
patches, particularly on areas of loose bottom
material (sand or rubble). Scattered short final
branchlets are given off from the main branches.
Colour is usually cream or pale brown.
Laboratory Diagnosis
Branching pattern: Branching is basically
arborescent, but the length and positioning of
secondary branches varies greatly. Branch widths
are from 10 to 20 mm. Secondary branches (or
‘branchlets’) can be regularly arranged along the
main branches, or scattered. In general, the
slender main branches tend to be more proliferous
than the sturdy ones.
Axial corallites: Outer diameter: 1-6 to 2-5 mm;
inner diameter: 0-6 to 0-9 rnm Septation: both
cycles usually complete, the primaries up to 2/3R,
secondaries up to 1 /4R.
Radial corallites: Tubular with round or
occasionally oval to nariform openings. They are
unequal in length and orientation (usually
oriented within a range 45° to 90°, except towards
basal areas, where they may be appressed).
Coenosteurn: Spines are distributed evenly over
corallites and inter-corallite areas. In heavily
calcified specimens these have elaborated tips; in
less calcified specimens the tips are simple to
laterally flattened.
(For more detailed laboratory diagnosis see
Wells, 1954).
Within Reef Distribution
From about 3m to 15m on reef slope,
deep-water lagoons, and patch reef areas, on
rubble or sandy floors, fringing reefs. Usually it
occurs with A. horrida (Dana), from which it can
easily be distinguished, on the Great Barrier Reef,
because A. vaughani is more sturdy, does not
usually extend the polyps in daylight, and is
commonly cream or pale brown, while A. horrida
is pale blue or grey. The upper range is more
limited than for A. horrida, which can occur in
some reef flat situations.
Identification Difficulties and History
The appearance of the branchlets can be very
varied (see Wells), but with good field notes the
species can be identified without confusion with
other arborescent species.
Geographic Distribution
Pacific; Marshall Islands, Great Barrier
Reef.
Acropora pulchra (Brook, 1891)
(Plates 58, 59, 60C)
Madrepora pulchra Brook, 1891, p.468; 1893, p.44,
pl.28, fig. A.
Madrepora pulchra var. stricta Brook, 1893, p.44,
pl.28, fig. B.
Madrepora pulchra var. alveolata Brook, 1893, p.45,
pl.28, fig. C.
Acropora pulchra: Vaughan, 1918, p.l62, pi. 66, figs.
3, 3a; Crossland, 1952, p.203; Stephenson and
Wells, 1956, p.l7.
Acropora pulchra var. stricta: Crossland, 1952, p.204,
pi. 34, fig. 2.
Acropora pulchra var. alveolata: Vaughan, 1918,
p.I62, pi. 66, figs. 1, 2; Crossiand, 1952, p.203.
Material Examined
BM: Keeling Is., A. pulchra holotype 1884.2.16.1.
QM: Bushy-Rcdbill Reef, adjacent Redbill Is., reef
flat: 7.vi.l975, C.W., Glil02, 011105; ll.vi.l975,
C.W., GI1097. 011103, 011104; I2.vi.l975, C.W.,
G1I093, 011095, 011096, GlllOO, GlllOl.
Heron Island, W. side, reef flat: 2.vii.l973, Y Loya,
01 II 17; 3.vii.l973, C.W\. 011120; July 1973,
C.W., 011115; 011119, lLxii.1973, C.W., GllllI,
011116, 011121, 011122; 15.xii.l973, C.W.,
011118.
286
MEMOIRS OF THE QUEENSLAND MUSEUM
Fiji Islands. Great Astrolabe Reefs. Feb. 1974, C.W.,
Gil 123-5.
Low Isles, August 1954, W. Stephenson (Stephenson
and Wells 1956): G2617-20; G2699-G2712, G2715,
G2716, G2721-5.
North West Island, C. Hcdley, 3-4 June 1924,
GB.R.C. 170.
Field Diagnosis
Colonies of this species are arborescently
branching, and can vary in overall shape from
open thickets to compact corymbose form. Small
radial corallites with oblique apertures give the
branch surface a smoother appearance than that
of other arborescent or corymbose species. Colour
is pale to dark brown, often with pale blue
tips.
Laboratory Diagnosis
Branching pattern: The angle of emergence,
length, and width of branches varies widely, as the
plates show. Branch widths are from 7 to 15 mm,
and the overall general colony form can be
anything from an arborescent thicket to a neatly
caespito-corymbose clump.
Axial corallites; 1 to 2 mm exert; external
diameter 2-0 to 3-5 mm, internal diameter 0*6 to
1-2 mm. Septation: primary septa well developed,
up to 2/3R, secondary septa absent or poorly
represented, occasionally all present, up to
1/4R.
Radial corallites: Mixed sizes, all very small,
project at 90^^ from branch, upper wall poorly
developed, lower wall extended and lip-like. The
opening is oval to dimidiate. Primary septa may
be present to 2/3R, or may be just visible;
secondary septa not usually visible, or a few may
be present to 1 /4R.
Coenosteum: Costate on radial corallites, openly
reticulate to costate between. General appearance
is light and spongy.
Within Reef Distribution
This is a reef flat species, occurring particularly
on the inner and middle reef flat. It occurs with
A. aspera, often in close association, as plate 60C
shows.
Identification Difficulties and History
Brook (1893) described the radial corallities
well and his descriptions should be consulted. The
main problem with this species is seen when it
occurs with A. aspera. The two species show
similar colony shape variations, apparently in
response to similar environmental conditions.
Although the radial corallites are different in both
shape and size, colonies sometimes appear to have
a mixture of characteristics of both species. This
problem was encountered on Redbill reef flat, at
places where populations of the two species
intermingled: some colonies suggest a hybrid
appearance. It is possible that some of Vaughan’s
A. pulchra specimens were A. aspera.
Geographic Distribution
Indo-Pacific: Cocos-Keeling Islands, Great
Barrier Reef.
Acropora aspera (Dana, 1846)
(Plates 60, 61, 62)
Madrepora aspera Dana, 1846, p.468, pi. 38, fig. 1,
la, lb; Brook, 1893, p.62 (synonymy).
Acropora aspera: Crossland, 1952, p.205, pi. 33, figs.
2, 3; Ncmenzo, 1967, p.65.
Madrepora hebes Dana, 1846, p.468, pi. 35, fig. 5;
Brook, 1893, p.l28 (synonymy).
Acropora hebes: Vaughan, 1918, p.I74, pi. 73, figs. 2,
2a, pi. 74, figs. 1. 2, 2a, 2b, pi. 13, fig. 6;
Hoffmeister, 1925, p.57. pi. 9, figs. 3a. 3b; Wells,
1950, p.36; Crossland, 1952, p.217; Wells, 1954,
p.423, pi. 104, fig. 3; Stephenson and Wells, 1956,
p.l4; Nemenzo, 1967, p.64, pi. 22. fig. 1.
Madrepora cribripora Dana, 1846, p.470, pl.31, fig.
1, la-lc; Brook, 1893, p.l23.
Madrepora manni Quelch, 1886, p.l50, pi. 9, fig. 1,
la; Brook, 1893, p.63 (synonymy).
Acropora ntanni: Fauslino, 1927, p.263, pi. 85, figs. 6,
7; Nemenzo. 1967, p.63.
Acropora tuzonica Verrill, 1902, p.231, pi. 36c, fig. 4,
pl.36F, fig. 9.
?Madrepora arabica Milne Edwards and Haime,
1860, p.l45; Brook, 1893, p.66 (synonymy).
MaTIRIM E.XAMIM'D
USNM; Fiji Islands. U.S. Expl. Exped., A. aspera
holotypc 285; Fiji Islands, U.S. Expl. Exped., A. hebes
syntypes 287, 286; Rewa River Mouth, Fiji Islands, U.S.
Expl. Exped., A. cribripora holotypc 289.
YPM; Manilla Bay, Luzon. Philippines A. luzonica
holotypc 1809.
PM: Mer Rouge, A. arabica holotypc 331.
QM: Big Broadhurst Reef: S. side, surge channel on
outer flat, 2 m, 21.x. 1973, C.W., GI0928; patch reef
in lagoon, 28.iii.l973, C.W., G10929, GI0930; SW. side,
reef slope, 18.X.1973, C.W., G10962.
Bushy-Redbill reef: W. side, middle reef flat,
14.vi.l975, C.W., G 10947; N. side, inner flat-algal bank
area, 6.vi.l975. C.W., G10937-41, G10943, G10960.
G1096I; adjacent Redbill Is., reef fiat, 7.vi.l975, C.W.,
G 10944-6.
Bowden Reef. SW. side, reef slope, 1 m, 15.vii.l972,
C.W., GI0925.
Heron Island; July 1973, C.W., GI0849-G10854;
G 1091 3— G 1 091 7; G10921; W. side, inner reef flat.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
287
lLxii.1973, C.W., GI0860-2; W. side, reef flat:
15.xii.l973, C.W., G10886-92; 17.xii.l973, C.W.,
G10893, GI0896; N. side, reef Hat, ILxii.1973, C.W.,
G10855; N. side, reef flat (solid area), 14.xii.l973,
C.W., G10883-5; N. side, inner reef flat, lLxii.1973,
C.W., G10856. G10857; N. side, middle reef flat,
lLxii.1973, C.W., G10858; S. side, outer reef flat,
12. xii.l973, C.W., G 10865-72; SW.side, outer reef Hat.
13. xii.l973, C.W., G10873-81; WSW. side, outer reef
flat, 12,xii-1973, C.W., G 10864; SE. side, outer reef flat,
17.xii.l973, C.W.. GI0897-902.
Low Isles. 1975, C. Limpus, G1093L
Masthead Reef, reef flat, August 1974, J. Buhmann,
G 10926.
Michaelmas Cay, 1975, C. Limpus, G10932,
G10933.
Maer Island, Murray Island Group, N. side, outer reef
flat, 17.vii.l974, G. Ingram, G10927.
Bramble Cay, Oct. 1924, C. Hedley, G.B.R.C. 7,
93-7, 149-52, 168, 169, 21 1.
Feather Reef, I Lviii.1924, ‘Geranium', G.B.R.C.
181.
Ellison Reef, 25.vii.1924, Paradice, G.B.R.C. 2-6,
54-67, 176.
Surprise and Flora Reefs, Sept. 1924, ‘Geranium’,
G.B.R.C. 8, 9, 12, 13, 16, 19-21, 24, 28.
Fiji Islands: Makaluvau Reef, 10. i. 1974, C.W.,
G10955, G 10957; Great Astrolabe Reefs, Feb. 1974,
C.W.. G 10949-54.
Field Diagnosis
This is a basically arborescent species in which
the colonies show considerable phenotypic
flexibility. Different colonies or parts of a colony
can have long, slender spreading branches with
scattered radial corallites or shorter thicker
branches, even to the extent of appearaing
corymbose, with crowded corallites. Radial
corallites are in two sizes and have poorly
developed walls. Colours are commonly pale
blue-grey, green-grey, or cream, less commonly
bright blue. It is a characteristic of this species
on the Great Barrier Reef that specimens, when
collected, exude large quantities of mucous.
Laboratory Diagnosis
Branching pattern: Ranges from open branch-
ing (at 90°) to closed branching, the branches
being given off vertically to obliquely to give a
corymbose appearance. Branch widths vary from
10 to 15 mm.
Axial corallites: From non-exert to 3 mm exert.
Outer diameter 3-5 to 4-5 mm; inner diameter 1-3
to 1-8 mm. Septation: Both septal cycles usually
present, the primaries up to 2/3R, the secondaries
up to 1/3R.
Radial corallites: Two sizes of radial corallite
are mixed. These are best regarded as large
diameter’ and ‘small diameter’, although the
‘large diameter’ radials are also always more
prominent. The ‘large diameter’ radials have
round openings, upper wall undeveloped, and
lower wall short and thickened, or extended
horizontally as a lip, which may be rounded or
pointed. The ‘small diameter’ radials are
sub-immersed to immersed. In general colonies
with short, thick branches tend to have the radial
corallites crowded with short, thick lips, and
colonies with long slender branches have radial
corallites scattered, with thin pointed lips. On the
larger radials primary septa are usually visible up
to 1/3R; and secondary development varies, but
often some septa are present, up to 1/4R.
Coenosteum: Open reticulate with simple to
laterally flattened spines between radials, costate
on radials.
Within Reef Distribution
This is a reef flat species, occurring from the
inner reef flat to outer flat, but not on elevated
outer flat platforms. It can occur in very silty and
coral-depauperate areas and shallow lagoonal
situations.
Identification Difficulties and History
The two species ‘A. hebes' and A. aspera are
both ‘well known’ and typical specimens of the two
species appear very different from each other.
However, the relationship of the two can be easily
established in the field. The placing of less known
synonyms is more difficult, and possibly there are
other species names still to be included. Two
major problems of identification occur with this
species. The first is in separating some corymbose
specimens from sturdy specimens of A. millepora,
the second in separating some arborescent colonies
from arborescent A. pulchra. (see discussions for
these species).
Geographic Distribution
Indo-Pacific: ?Red Sea, Cocos-Keeling, Philip-
pines, Great Barrier Reef, Fiji Islands, Samoa,
Marshall Islands, Tongan Islands (C.W.).
The ‘Acropora corymbosa’ Group
A number of species have radial corallites with
scale-like lips, and a disproportionately large
number of species descriptions have been written
to cover these species.
In the following list there is barely a species
which has not been placed in synonymy or
compared with, or mistaken for some other species
in the list, and of the ‘real’ species that exist, each
has been placed under a number of the commonest
288
MEMOIRS OF THE QUEENSLAND MUSEUM
names. The main species concerned are: A.
corymhosa (Lamarck), A, efjlorescens (Dana), A.
cytherea (Dana), A. spicifera (Dana), A.
hyacinthus (Dana), A. surcidosa (Dana), A.
millepora (Ehrcnberg), A. prostrafa (Dana), A.
subulata (Dana), A. convexa (Dana), A. armata
(Brook), A. reticulata (Brook). A, arcuata
(Brook), A. rythereilo (Verrill). A conferta
(Quelch), A. pectinata (Brook), A. recumhens
(Brook), A. squamosa (Brook).
The most widely applied name on this list is the
oldest. A. corymhosa (Lamarck). Three con-
tenders for this name are the species I am
identifying as A. hyacinthus, A. cytherea and A.
millepora. Older interpretations of this species
tended to favour an A. cytherea-Wke appearance.
A specimen regarded as the type (of A.
corymhosa) was seen and described by Brook
(1893) who points out the similarity between A.
cytherea and this. Professor J. Wells has kindly
shown me photographs of (apparently) this type,
sent to him some time ago by the Paris Museum.
Neither my own search of the Lamarck collection,
nor a later search by Dr J. P. Chevalier could
locate a type.
My opinion is that the currently accepted view
of this species (c.g. Wells 1954) is morphologically
between A. millepora and A. hyacinthus. While
it is possible that this is a true species, at least
some of the specimens so identified are other
species: for example Stephenson and Wells 1956
mention two specimens, of which one (G2626) is
a small reef flat specimen of A. hyacinthus, and
the other (G2623) a stunted A. millepora.
The species has been mentioned by many other
authors since 1893, e.g. von Marenzeller (1907),
Vaughan (1918), Hoffmeister (1925, 1929), Thiel
(1932), Crossland (1952), Ncmenzo (1967), Pillai
and Scheer (1976). and is regarded as having a
wide Indo-Pacific distribution. In the interests of
taxonomic stability, I am not attempting to trace
its synonyms on the basis of a regional study, but
I feel a caution must be taken that the
interpretation of this species by the various
authors may not coincide. Hopefully, with
extensive regional studies, this group will be given
a more complete taxonomic treatment.
The species which I interpret as A. hyacinthus
and A. cytherea are dominant members of the
reef-front assemblage. They arc the early
colonizers and often influence the effective shape
of some reef areas. Special problems are
associated with the separation of these species at
their overlapping limits. A. spicifera has not been
present in my areas of study. A. millepora is a
common reef-flat species which has usually (on
the Great Barrier Reef) been identified as its
synonym, A. squamosa.
Acropora hyacinthus (Dana, 1846)
(Plates 63, 64A-C, 65, 66A,B)
Madrepora hyacinthus Dana. 1846, p.444, pi. 32,
fig. 2; Brook, 1893, p.I07 (synonymy).
Acropora hyacinthus: Th\e\. 1932. p.i23, pi. 1 6. fig. 2;
1933. p.20; Stephenson and Wells. 1956. p.16;
Nemenzo. 1967. p.|l5, pi. 33, fig.I; Pillai and
Scheer, 1976, p-29.
Acropora hyacinthus (?part): Hoffmeister, 1925,
p.64. pL]3, fig.3: Wells, 1954. p.42L
? Madrepora conferta Quelch, 1886. p.l64, pi. 10,
fig.3; Brook, 1893, p.l08.
Madrepora pectinata Brook, 1892, p.460* 1893 p 95
pl.27, fig.D, E.
Madrepora recumhens Brook, 1892, p461- 1893
p.I06, pl.27. fig.F,
Acropora pectinata: Vaughan. 1918, p.l72, pl.71,
fig.I, la-lc, 2; Thiel, 1932, p.ll9, pi. 14, fig.4
(synonymy).
Acropora corvmbosa (part): Stephenson and Wells
1956, p.l2-
Madrepora surculosa var. turbinata Dana, 1846,
p.446: Brook, 1893, p.200 (synonymy).
Madrepora turbinata: Verrill, 1864, p.'42.
Acropora turbinata: Verrill. 1902, p.242.
Note: This .synonymy does not include all references
to synonyms, as their interpretations are difficult to
trace.
Matfriai Examinkd
USNM: Fiji Islands, U.S. Expl. Exped., A. hyacinthus
holotype 246; Tahiti, U.S. Expl. Exped., A. surculosa
(var. turbinata) synlype 251.
YPM; Tahiti, A. surculosa var. turbinata type
fragment 2017.
BM: Fiji Reefs, Challenger, A. conferta holotype
1885.2.1.12; Thursday Is., Saville Kent, A. pectinata
syntype 1892.6.8.155; low Woody Is., G.B.R . A.
recumhens (var.) 1 892. 6. 8.161.
QM: Big Broadhursl Reef; SW side, reef slope: Im,
! Lx. 1973, C-W . G9860, G986I; Im. 13.X.1973, C.W.,
G9862; l-3m. 22.X.197.3, C.W.. G9863; L5m, 11.x. 1973,
C.W., G9859; 2-7m, I2.X.I973, C.W., G9864; 3m.
29.iii.1973, CW., G9872; 5-3m, 13.X.I973. C.W..
G9865:6 3m. 13.x. 1973, C.W.. G9866: 8Tm. 15.x. 1973
C.W.. G9867; Oct. 1973. C.W., G9868: patch reef in
lagoon. 2 Lx. 1 973, C.W., G9869; E. side, reef slope, 4m.
2 Lx. 1973, C W.. G9870.
Bowden Reef, opening in outer reef. 26.vii.1972,
C.W., G9877. G9878: SW side, reef crest, 0-6m.
I5.vii.l972, C.W.. GI0716.
Bushy-Rcdbill Reef; S. side, outer flat, 3Lv.l975,
C.W., G9874: NW.side, top of patch reef off reef edge,
19.xii.I972, E. Lovell, GI0727; W. of Redbill Is., edge
of surge channel, 0-5m, I9.xii.I972, C.W., GI0232.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
289
Darley Reef, patch reef in lagoon, 24.jii.1973 C W
G9876.
Great KeppcI Island, E. side, fringing reef, Dec. 1975
R. Woolley, G 1 0723, G 10726.
Heron Island: reef flat, W. side: 2.vii.l973, Y. Loya.
G9875; 6.vii.l973, Y. Loya. G10720. July 1973, C.W.[
G107I8, GI0722; N. side, 9.viii.l950, W. Stephenson,
G9684; reef flat. April 1954, E. Grant, G2680; N. side,
14.V.I954, W. Stephenson, G2681-3.
Lady Musgrave Reef, W. side, outer slope,
16.vii.197L 1. Neuss, G6268.
Tryon Reef, E. side, reef flat, 12.vii.l971. C.W.,
G6267; NW. side, reef slope, I8.xii.l971, I. Neuss
G6478, G6480, G6494.
Viper Reef, patch reef in lagoon, I m, 16.vii.l973
C.W., G10717.
Bramble Cay, Oct. 1924, C. Hedley, G.B.R.C. 34-9
86-92.
Coates Reef, 13.vii.l924, ‘Geranium’, G.B.R.C. 141,
195.
Flora Reef. 21. v. 1924, Dr Paradice, G.B.R.C. 138.
Flinders Reef. Moreton Bay, NW. margin, 9.viii-I972,
E. Lovell, G6984; NW. slope, 2Lvi.l973, E. Lovell,
G7297, G7298.
Field Diagnosis
Tabular to plate like colonies to over 3m
diameter, formed by the developmental process
described by Floffmeister (1925). Colonies are
sturdy in texture, and the vertical branchlets have
a rosette like appearance due to the regular
arrangement of labellate corallites around them.
Colours are blue with pink growing edge (common
reef flat colouration), pale to dark brown, pink-,
yellow-, green- or blue-brown.
Laboratory Diagnosis
Branching pattern: From a central to lateral
stalk, branching is horizontal, with branches
anastomosing to a greater or lesser extent.
Secondary branchlets are single or in groups of
up to seven, from 1 mm to 22 mm long, and from
3 mm to 8 mm wide (commonly around 10 mm
long and 4 mm wide). In very shallow reef flat
situations, the branchlets may be reduced to mere
buds, but very wide (up to 8 mm). The longest
branchlets are in lagoonal specimens. In this
species and in A. cytherea, layers may be added
to the original plate.
Axial corallites: Up to 1-5 mm exert; outer
diameter 1-4 to 2 0 mm; inner diameter 0-6 to LI
mm. Septalion: primaries present to 2/3R,
secondaries absent, or a few present to 1/4R.
Radial corallites: Tubular appressed, with only
lower (or outer) wall developed (2/3 circumfer-
ence or less) to form a rounded lip. Radials are
arranged evenly and closely around the branchlet,
giving a rosette-like appearance, typical of this
species. Septation may be strongly developed, ail
primaries being present to I /4R, plus a few
secondaries, or so weakly developed that only the
directive septa can be distinguished. On horizontal
branches immersed corallites occur.
Coenosteum: Coslatc on radial corallites,
reticulate with occasional laterally-flattened
spines between.
Within Reef Distribution
This species occurs in lagoons, deep water reef
flat situations, shallow outer reef flat, and on the
reef slope to about 30 m (species no. 4 in Wallace
1975).
Identification Difficulties and History
Dana’s type of A. hyacinthus (illustrated by
Hoffmeister 1925) is a young specimen just in the
Vasiform’ stage of development. This compares
closely with Juveniles of the species I have
described, which is a sturdier species than A.
cytherea. Much reef flat material of the species
(well calcified) has probably been identified as A.
corymbosa, while the interpretations of A.
hyacinthus (e.g. by Hoffmeister) tend to be of a
lighter-textured form. Whether his material was
of A. cytherea or of a geographic variant of A.
hyacinthus I am not able to judge. The deepest
occurring colonies of A. hyacinthus are light
textured with reduced radial corallite lips. These
could be confused with A. cytherea from shallower
depths but from a comparable depth A. cytherea
has the distinctive ‘A. reticulata’ form. A. conferta
(Quelch) has slightly smaller axial corallite
dimensions than those given here, but on radial
corallite characteristics it appears to be a
synonym. This is one of the most successful
Indo-Pacific species of Acropora: it is an early
colonizer, and occurs with great abundance in
some areas, Hoffmeister's detailed treatment of a
species under this name, without parallel detailed
treatment of A. cytherea (or 'A. reticulata'),
leaves unsolved a taxonomic puzzle. A study of
these two species on the Samoan reefs would be
extremely useful.
Geographic Distribution
Widespread Indo-Pacific distribution from the
Mascarene Archipeligo (G. Faure pers. comm.) to
Tahiti. Exact localities cannot be quoted because
of taxonomic confusion.
Acropora cytherea (Dana, 1846)
(Plates 63, 64A, D, 66C, D, 67)
Madrepora cytherea Dana, 1846, p.441, pi. 32, fig. 3a,
3b; Brook, 1893, p.99 (synonymy).
290
MEMOIRS OF THE QUEENSLAND MUSEUM
Acropora cytherea: Crossland. 1952, p.215.
Madrepora efflorescens Dana. 1846, p.441, pi. 33,
fig. 6; Brooic, 1893, p.35 (synonymy).
Acropora efflorescens: Pillai and Scheer, 1976, p.26,
pl.3, fig.3.
Madrepora armala Brook, 1892, p.452; 1893, p.lOO,
pi. 10, figs. A, B (synonymy).
Madrepora reticulata Brook, 1892, p.461: 1893, p.68,
pi. 4, figs. A, B.
Madrepora reticulata var. cuspidata Brook, 1893,
p.69.
Acropora reticulata: Wells, 1954, p.422, pi.] 10,
figs. 4-6, pi. 114, figs.1-6 (synonymy); Pillai and
Scheer, 1976, p.28, pl.7, fig.l,
Madrepora arcuata Brook. 1893, p.l02, pi. 12; Studer,
1901, p.395.
Acropora cytherella Verrill 1902, p.253, pi. 36, fig. 7,
pi. 36a, fig.7. pl.36F, fig.l. (synonymy).
Acropora corymbosa 'cytherea Form’: von Maren-
zeller, 1907, p.32. pl.l, figs.I. 2; pl.2, fig.3.
Acropora hvacinthus (part): Hoffmeister, 1925, p.64;
Wells, 1954, p.421.
MaTI KIAI E\AMINt:D
USNM: Tahiti, U.S. Expl. Exped., A. cytherea
syntype 226; Samoa, A. hyacinthus (id. Hoffmeister),
no. 14 Mayor Collection.
YPM: Fiji Islands, U.S. Expl. Exped., A. cytherella
holotype 2007; East Indies, A. efflorescens ?type
1799.
BM: Singapore, A. arrttata syntypes 1850.1.16.1,
1857.4.6.1; Amirante Islands, A. reticulata syntype
1882.10.17.131; Navigator Island, A. arcuata syntypes
1862.1.27.5, 1875.10.2.9.
QM: Big Broadhurst Reef; S W. side, reef slope: I m,
13. x. 1973, C.W.. G9841, G9842; 1-3 m, ll.x.1973,
C.W., G9837, G9838; 2 m. 26.iii.1973, C.W., G9853;
2-3 m, I2.X.I973, C.W., G9839, G9840; 3 m,
28.iii.1973. C.W., G9856; 5 m, 27.iii.1973, C.W.,
G9855; 7 m, 13.X.I973, C.W., G9843; 8-1 m,
14. X.1973, C.W., G9845; 8-2 m, I4.X.1973, C.W.,
G9844; 8-3 m, I5.X.1973, C.W., G9847; 8-7 m,
15. X.I973, C.W., G9846; 9-7 m, 15.X.1973, C.W.,
G9849; 10-6 m, 6.x, 1973, C.W., G9848; SW. side, surge
channel floor, 5 m, 20.X.I973, C.W., G9850.
Bushy-Redbill Reef. NW. side, reef slope, 21.xii.l972,
C.W.. G9857.
Tryon Reef: NNE. side, outer slope. 21.xii.I97], I.
Neuss, G6482; NW. side, outer slope, 6-10 m,
19.xii.l97I. I. Neuss, Ci6479.
Viper Reef, patch reef in lagoon. 1 m from top,
16. vii.l972. C.W., G107I7.
Flinders Reef, Moreton Bay: 1973, E. Lovell, G7290,
G7291; NW. margin, 3-12 m, 9.viii.l972, E. Lovell,
G6999, G7032; W. margin, 10 m, 10.iv.l972, E. Lovell,
G7303, G7309.
Solitary Islands, 1972, J. Veron et al., G7057.
G7058.
Field Diagnosis
Tabular to plate-like colonies to over 3 m
diameter, formed by the developmental process
described by Hoffmeister (1925) for A. hyacinth-
us. Colonies have a light crumbly texture. Axial
corallites are usually obviously exert, and contrast
with slight scale-like radial corallites. Horizontal
branches are reticulated to fully anastomosed.
Colours are cream, pale brown, pink-, yellow-,
green- or blue-brown or -grey.
Laboratory Diagnosis
Branching pattern: From a central to lateral
stalk, branching is horizontal, with branches
anastomosing to a greater or lesser extent.
Secondary branchiets are vertical and single or in
groups of up to six, from long and slender (up
to 20 mm length, width around 3 mm at this
length), to short bundles of tubular axials
approximately 5 mm long with occasional
proliferation of radial corallites at their base. In
the series of specimens from lesser to greater
depths on the Big Broadhurst Reef, a transition
occurs from long branchiets with non-proliferous
tips (identifiable with ‘A, arcuata’ and A.
armata), through shorter branchiets with
proliferous tips, the proliferations often partially
naked of radial corallites, to very short branchiets
with few radial corallites (identifiable as ‘A.
reticulata’). All specimens are openly reticulated
and lightly calcified.
Lagoonal colonies are more anastomosed and
heavily calcified and the branchiets are similar to
those of deep reef slope colonies. Specimens from
the geographically extreme southern locations
(Flinders Reef, Moreton Bay and the Solitary
Islands) have the main branches strongly
anastomosed, and bundles of short thick
proliferous branchiets: they identify with A.
cytherea sens, strict.
Axial corallites: From 1-0 to 5-0 mm exert.
Outer diameter 1-3 to 1-7 mm in Great Barrier
Reef specimens. 1-6 to 2-5 mm in the Flinders
Reef — Solitary Island specimens; inner diameter
0*7 to 10 mm. Septation: primary septa present,
up to 2/3R; secondary septa often absent, never
fully present, up to 1 /4R.
Radial corallites: Tubular appressed with only
lower (or outer) wall (1/2 circumference or less)
developed, to form an elongate lip. In A. cytherea
s.s. forms these lips are thickened: on the reef
slope they are light structured and may be reduced
to single or double points. Septal development of
radials very poor: usually only directives can be
seen, sometimes a few other primaries are present
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
291
as fine points. On horizontal branches immersed
coralliles occur.
Coenosteum: Costate on radial corallites,
reticulate with laterally flattened spines in
between.
Within Reff Distribution
On the Great Barrier Reef this is a reef slope
species (species no. 5 in Wallace 1974), but
occasional colonies occur in lagoonal and deeper
water reef flat areas. On other reefs (e.g.
Enewetak Atoll) it has a similar distribution
(C.W.).
Identification Difficulties and History
Two special taxonomic problems are associated
with this species: (1) In combining A. cytherea
with A. reticulata, and (2) In distinguishing
between lightly structured colonies of A.
hyacinthus and colonies of A. cytherea with good
branchlel development.
A. cytherea s.s. occurs in geographically
extreme locations (Tahiti, Solitary Islands,
Moreton Bay). The temptation is to retain A.
reticulata, which best describes the species in its
main tropical range.
The second problem can also be staled as a
problem of interpretation of A. arcuata and A,
armata. These and A. cytherea have been
combined with A. hyacinthus by recent authors.
On A. arcuata and A. armata my own
observations are of a co-occurrence of these
‘species’ with definite A. hyacinthus at the same
depth on the Big Broadhurst reef front, and their
graduation into typical 'A. reticulata' with depth.
Crossland’s works are important to the ecological
documentation of this species: in 1928 he
identified A. cytherea as A. hyacinthus, in 1931
reversing the identification. In 1952 he notes that
both species are common in Tahiti.
This species, as I have defined it, is easily
separated from A. hyacinthus in the field by its
light crumbly skeleton. The two species co-occur
on the reef front, A. cytherea extending a little
deeper, and A. hyacinthus extending more into
reef flat areas.
Geographic Distribution
The species has a wide Indo-Pacific distribution
from the Mascarene Archipeligo (G. Faure, pers.
comm.) to Tahiti.
Acropora millepora (Ehrenberg, 1834)
(Plates 68, 81 D)
Heteropora millepora Ehrenberg, 1834, p. 109.
Madrepora millepora: Brook, 1893, p.Il6
(synonymy).
Acropora millepora: Verrill, 1902, p.257; Thiel, 1932,
p.l24, pi. 18, fig. I, pi. 19, fig.l; Nemenzo, 1967,
p.94, pi. 28, figs.l, 2.
Madrepora spathulaia Brook, 1891, p.469; 1893,
p.l2I, pi. 32, fig.B.
Madrepora squamosa Brook, 1892, p.463; 1893,
p.l20, pl.20, fig. B;
Acropora squamosa: Vaughan, 1918, p.l73, pi. 72,
figs.l, 2, 2a, 3; Crossland, 1952, p.216; Stephenson
and Wells, 1956, p.l8.
Acropora sarmentosa: Vaughan, 1918, p.l7A, pi. 72,
fig.4.
MAThRiAL Examined
BM: Claremont Is., Great Barrier Reef, Saville Kent:
A. millepora (mentioned Brook, 1893) 1892.6.8.165; A.
squamosa syntype 1892.6.8.163; Treasury Is., Solomon
Islands, D.A. Guppy, A. spathulata holotype
1884.12.11.25.
MNB: A. millepora holotype, 854 (photograph).
QM; Big Broadhurst Reef: patch reef in lagoon,
28.111.1973, C.W., G1 1052-6; E. side (weather side),
channel in outer flat, 21.X.1973, C.W., G1I055.
Bushy-Redbill Reef: NW. side, outer reef fiat,
23.xii.1972, C.W., G11047, G11049; NW. side, reef
slope, 23.xii.1972, C.W., G11046.
Bowden Reef, SW. side, reef crest, 26.vi.1972, C.W.,
G11064.
Darley Reef, patch reef in lagoon, 1 m, 18.vi.l972,
C.W., G1I057; 3 m, 22.iii.1973, C.W., G11058; 4 m,
22.111.1973, C.W., G11059.
Heron Island: N. side, reef flat, 14.xii.l973, C.W.,
GI0996-G1 1001; S. side, outer reef flat, 12.xii.l973,
C.W., G 10973, G 10976-83; SW. side, outer reef flat,
I3.xii.l973, C.W., GI0985-93; W. side, reef flat:
15.xii.l973, C.W., Gl 1002-5; 17.xii.l973, C.W.,
G1 1006-8; SE side, reef flat, 17.xii.l973, C.W.,
Gl 1009-16.
Great Keppel Is., Dec. 1975, R. Woolley, G11061.
Maer Island, Murray Island group, N. side, inner reef
flat: 16.vii,1974, G. Ingram, G1I060; 18.vii.l974, G.
Ingram, Gl 1062.
Masthead Reef, reef flat, Aug. 1974, J. Buhmann,
GI1063.
Michaelmas Cay, 1974, C. Limpus, G11067.
Low Isles, 1974, C. Limpus, G 11066, G 11068.
Fiji Islands, Great Astrolabe Reefs, Feb. 1974, C.W.,
Gl 1033-43.
Field Diagnosis
Low corymbose or stalked corymbose colonies,
commonly with neat round outline; branches
terete or slightly tapering, covered by evenly sized
and closely arranged radial corallites with flaring,
scale like lips. Commonest colouration (on both
the Great Barrier Reef and Fijian reefs) is a dull
to brilliant green with dull to bright orange branch
tips. The colonies can also have multiple
colouration, being predominately blue or pink,
292
MEMOIRS OF THE QUEENSLAND MUSEUM
with touches of other pastel colours and a grey
sheen, or bright orange to pale cream.
Laboratory Diagnosis
Branching pattern; Branches are given off
vertically to obliquely from a central to lateral
region which may be consolidated into a stalk. In
some colonies from sandy situations a growing
point is barely recognizable, and the growth is
bush like and relatively indeterminate. A small
amount of secondary branching occurs, and the
top of the colony is in a single plane. Branch width
varies from 7 to 13 mm.
Axial corallites: Barely exert. Outer diamter 24
to 3-9 mm; inner diameter 0-9 to 1*6 mm.
Septation: primary cycle fully developed, up to
i/2R; secondary cycle usually represented, but
not all septa developed, up to 1 /4R.
Radial corallites: No upper wall is developed,
the lower half of the wall is expanded as a rounded
lip, and the outer edges of this lip may Bare away
from the opening of the corallite. The primary
septa are often well developed, up to 2/3R,
secondaries absent or a few present to 1 /4R.
Coenosteum; Coslate on radial corallite lips,
reticulate with simple spines between radials.
Within Reei Distribution
Reef flat, from sandy middle reef flat to
consolidated outer flat, occasionally to a few
metres below the reef top; tops of lagoonal patch
reefs and fringing reefs.
Identification Difficulties and History
The usual name applied locally to this species
is 4. squamosa, but the material from the Great
Barrier Reef identified as A. millepora by Brook
(apparently the last person to observe and describe
Ehrenberg’s type) belongs to this species, and in
fact forms a series from the same locality as his
A. squamosa types. Verrill (1902) synonymised
the two, but he has not been followed by later
authors.
Two species regarded as being similar to A.
millepora (or A. squamosa*) may now be
dismissed: A. sarmentosa was misinterpreted by
Vaughan (1918), and is a clearly definable
species; A. aspera (= A. hebes) with many
synonyms is a highly versatile reef flat species
which in its low corymbose form appears very
similar to this species. A. millepora has a more
restricted reef flat distribution than A. aspera,
and extends closer to the reef crest. In aggressive
interactions observed at Heron Island and on
Bushy-Redbill Reef A. millepora tissues appear
to be inclined to overgrow those of A. aspera.
Small colonies of the species have- probably been
identified as ‘4. corymbosa* by some authors. 4.
imbricata (Ehrenberg) is another possible
synonym.
Geographic Distribution
The most definite records of this species are
only from the Great Barrier Reef, Solomon
Islands, Fiji Islands (C.W.) and Marshall Islands
(C.W.). Other less certain records extend the
range to Ceylon, the Philippines, New Ireland,
New Hebrides.
Acropora delicatula (Brook, 1891)
(Plate 69)
Madrepora delicatula Brook, 1891, p.461; 1893,
p.l09, pi. 28, figs. D, E.
Acropora delicatula: Wells, 1954, p.420, plT15,
figs.l, 2.
non Acropora delicatula; Stephenson and Wells,
1956, p.l2.
Material Examine^d
BM; Solomon Islands, Guppy, 4. delicatula holotype
1884.12.11.23.
QM; Big Broadhursl Reef, SW.side, reef slope: 2 m,
26.111.1973, C.W., G11442; 6 m, 25.iii.1973, C.W.,
Gll44i; 6 m, 27.iii.1973, C.W., G1I439; 8 m,
26.111.1973. C.W., G11436; 8-1 m, 14.x. 1973, C.W.,
GI1434; 8 6 m, 15.x, 1973, C.W., 011438; 13-4 m,
17.X.1973, C.W., G11437; 17 m, 23.X.1973, C.W.,
G1 1435.
Bushy-Redbill Reef, NW. side, reef slope, 3-7 m,
21.X11.1972, C.W., G11440.
Field Diagnosis
Shallow caespitose or corymbose-plate colonies
have slender vertical branchlets with small,
scalc-like radial corallites. Colours are cre.*m, pale
brown or greenish brown.
Laboratory Diagnosis
Branching pattern; From an attachment that is
lateral or nearly so, branching is at first
horizontal, then branchlets or bundles of
branchlets are given off vertically upwards. The
branchlet bundles can be as tall as 40 mm, and
branchlet width is around 5 mm. A small amount
of branching from the undersurface may
occur.
Axial corallites: 0-5 to I *5 mm exert. Outer
diameter 1-5 to 2 mm; inner diamter 0-6 to 0*9
mm. Septation: primary septa well developed, up
to 3/4R, secondary cycle usually absent, or a few
septa may be present up to 1/4R.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
293
Radial corallites: Appressed tubular, nariform
or dimidiate, the outer wall only developed, and
extended as a Up which is correspondingly
rounded, elongate or spade-shaped. The septal
development varies, but commonly all primary
septa are visible, up to I /4R, and two or three
secondaries just visible.
Coenosteum: Costate or rows of simple spines
on radial corallites, neat rows of simple spines, or
reticulate with scattered spines, between.
Within Reef Distribution
I have observed only a very few colonies, all in
reef slope situations. The species is much more
commonly represented from fringing reefs, and a
series in J.C.U. is from the Lizard Island and
Palm Islands reefs.
Identification Difficulties and History
This species is poorly documented, and working
with a poor series I am unable to resolve some
problems. A. subulata (Dana), may be a senior
synonym. Stephenson and W'ells’ specimen from
Low Isles (QM G2648) is A. tenuis.
Geographic Distribution
Great Barrier Reef, Solomon Islands, Marshall
Islands.
Acropora haimei (Milne Edwards and Haime,
1860)
(Plates 70, 71)
Madrepora haimei Milne Edwards and Haime, 1860,
p.l51; Brook, 1893, p.77 (synonymy); von
Marenzeller, 1907, p.51 pi. 16 figs. 45-48.
Acropora haimei: Vaughan, 1918, p.l63, pi. 70, figs. 3,
3a. 3b, pi. 66. figs, 4. 5; Crossland, 1952. p.207.
pl.33, fig. I, pi 35. fig. !: Ros.si. 1954, p.48;
Stephenson and Wells, 1956, p.l4; Nemenzo. 1967.
p.82. pi. 25. fig. 1.
Material Examined
QM: Big Broadhurst Reef: SW. side, reef slope: 9-8
m, 16.X.1973, C.W., G10216, G102I7; 9-6 m, 15.x. 1973,
C.W., G10215; 8-7 m, 15.X.1973, C.W., G102I3,
G102I4; 8-6 m, 15.X.1973, CW., GI0212; 8-3 m,
15.X.1973, C.W., G10211; 8 m, 14.X.1973, C.W.,
G102I0; 7-8 m, 23.X.1973, C.W., G10206-9; 7 m,
13.X.1973, C.W., G10205; 5-6 m, 23.X.1973, C.W.,
G10204; 6 m, 13.x. 1973, C.W., G10224; 5-7 m,
23.X.1973, C.W., G10225; 5 ni, 27.iii.1973, C.W.,
G10182, G10221; 5 m, 25.iii.1973, C.W., G10181; 4-2
m, 22.X.1973, C.W., G10222; 3-8 m, 12.X.1973, C.W.,
G10180; 1-6 m, 22.X.1973, C.W., G10223; 15 m,
12.X.1973, C.W., G10179; 1-3 m, ll.x l973, C.W.,
G10202; 1-3 m, 14.X.1973, C.W., G10203; 1 m,
26.iii.1973, C.W., G10194; SW. side, surge channel
opening, 10 m, C.W., G10183; patch reef in lagoon,
21.x,1973, C.W., G10184, G10185.
Bushy-Redbill Reef; NNE. side, reef slope, 3 m,
30.xii.l972, C.W., G10186, G10187, Gl 1303; NW. side,
reef crest, 30.xii.l972, C.W„ G10195, G10227, G10228;
W side, reef crest, 18.xii.l972, C.W., G10226.
Bowden Reef, opening in SW. side of reef, 26.vii.1972,
CW., G10188, G10I89, G10229-31.
Field Diagnosis
This is a medium sized compact arborescent
species, which occurs as scattered turf like
patches, bushes or thickets. It is distinguished
from other arborescent species by very open radial
corallites. Colour is usually cream or pale
brown.
L aboratory Diagnosis
Branching pattern: Branching is open and
irregular. Main branches are up to 15 mm wide
and tapering.
Axial corallites: Outer diameter 2-2 to 3-5 mm;
inner diameter 0-8 to 1-2 mm. Septation: primary
septa present up to 2/3R; secondary septa all
present, or mostly present, up to 1/3R.
Radial corallites: Tubular, extend at from 45°
to 90° from branch; openings cochleariform; the
upper part of the corallite wall (about 1/3 of the
diameter) is thinner and shorter than the rest of
the wall, and the thicker lower portion often flares
slightly. Septal development is usually very
marked: primaries can extend to R, and a full set
of secondaries can be present. A good key to the
identification of this species is the presence, on the
proximal part of branches, of well formed
cochleariform radials with stiong septal
development.
Coenosteum: Costate on ladials, reticulate with
simple or laterally flattened spines in between.
Identification Difficulties and History
Although following Crossland’s interpretation,
I am still not fully satisfied that A. haimei is the
correct identification of this species. Milne
Edwards and Hairne’s type cannot be located.
Other authors, particularly von Marenzeller.
describe a greater variety of colony shapes than
I have seen, including shallow water reef flat
forms which approach corymbose. To a worker
with some experience in the field, this species
becomes easy to distinguish from other arbores-
cent species because of its large open radial
corallites.
Whhin Reef Distribution
The species occurs as small patches on middle
and outer reef flats, but achieves its greatest
294
MEMOIRS OF THE QUEENSLAND MUSEUM
abundance on the reef slope (to about 10 m), in Heron Island, W. side, reef flat, 6.vii.l973, Y. Loya,
surge channel openings and on sandy bottoms in G11457.
deep lagoons and around broken reef- patch Masthead Reef, SW. side, upper reef slope, Aug.
ai-goc 1974, J. Buhmann, G11419.
Geographic Distribution
Indo-Pacific: Red Sea, ?Diego Garcia, Maidive
Islands, Ceylon, Mauritius, Singapore, Great
Barrier Reef, Fiji Islands.
Acropora tenuis (Dana, 1846)
(Plates 72, 73)
Madrepora tenuis Dana, 1846, p.451; Ortmann, 1888,
p.l52; Brook, 1893. p.83 (synonymy).
? Madrepora eurystuma Klunzinger, 1879, p. 16, pi.
L fig. 8, pi. 4, fig. 7, pi. 9, fig. 12.
Madrepora eurystoma: Brook, 1893, p. 137
(synonymy).
Madrepora macrostoma Brook, 1891, p.464; 1893, p.
105, pi. 19, fig. B.
Madrepora bifaria Brook, 1892, p.453; 1893, p.llO,
pi. 30, fig. A.
Madrepora kenti Brook, 1892, p.458; 1893, p. 110,
Pl- II, fig. B.
Madrepora dilatata Brook, 1893, p.8I.
Madrepora anihocercis Brook, 1893, p. 106, pl.l3, fig.
C (synonymy).
?Acropora anihocercis: Nemenzo, 1967, p. 109.
Materiai. Examined
USNM; Fiji Islands, U.S. Expl. Exped., Madrepora
tenuis holotype 259.
BM; Koseir, Red Sea, A. eurystoma 1886. 10.5.5;
Diego Garcia, G. C. Bourne, A. eurystoma 1891.4.9.1.
(mentioned Brook, 1893); Mauritius, A. macrostoma
holotype 1878.2.4.7; Java, A. bifaria holotype
1859.12.12.2; Thursday Island, Saville-Kent, A. kenti
holotype 1892.6.8.202; Palm Island, Saville-Kent, A.
anthocercis syntype 1892.6.8.235; Rocky Island,
Saville-Kent, A. anthocercis syntypes 1892.6.8.236,
1892.6.8.237.
QM: Big Broadhurst Reef, SW. side, reef slope; 1-9
m, 22.X.1973, C W., GII413; 3-3 m, 22.x.l973» C.W.,
G11429; 4 m, 24.iii.1973, C.W., 011431; 6 m,
27.111.1973, C.W., G11428; 7 m, 28.iii.1973, C.W.,
GI1430; 7-8 m, 14.X.1973, C.W., G11456; 8 m,
26.111.1973, C.W., G11433; 8 m, 14.X.1978, C.W.,
011420; 8 1 m, 14.X.I973, C.W., G11424; 8-2 m,
14.X.I973, C.W., G11422, GII423; 8-6 m, 15.x. 1973,
C.W., G1 1425; 9-2 m, 15.X.1973, C.W., G1 1421; 9-8 m,
16.X.1973, C.W., G11426, GU432.
Bushy-Redbill Reef: W. side, outer reef flat, Dec.
1972, C.W., G114I7; NW. side, reef slope: 19.xii.l972,
E. Lovell, G11412, G11415, G11416; NW. side, floor
outside slope, 22.xii.1972, C.W., G1I427.
Darley Reef, patch reef in lagoon, 22.iii.1973, C.W.,
G11414, G11418.
Field Diagnosis
Colonies are Thick plates’ or caespito-
corymbose. The radial corallites are evenly
arranged and have distinctive, lip like, flaring
outer wall and strong septal development, which
can often be seen with the naked eye. Once learnt
in the field, this species is always easily
recognised. The commonest colour is cream, less
comrnonly colonies are greenish-blue or bright
blue. Polyps are often partly extended during the
day, and in cream colonies these are often bright
orange (axial polyps) and bright purple (radial
polyps).
Laboratory Diagnosis
Branching pattern: From a growing area which
may be central to lateral, branching is horizontal,
then secondary branchlets or groups of branchlets
are given off vertically to obliquely, their tips in
one plane. Branching may also occur from the
lower surface but these lower branches are never
as long as the upper. Branchlet widths are from
6 to 8 mm. Shallow water specimens are Thick
plates’ of up to 110 mm vertical depth; deeper
water specimens may be thin plates as little as 35
mm thick.
Axial corallites: Up to 2 mm exert. Outer
diameter: 1-9 to 3-0 mm; inner diameter 0-8 to
1-2 mm. Septation; both cycles usually developed,
the primaries up to 2/3R, the secondaries up to
1/3R.
Radial corallites: Tubular, ascending, the inner
wall less developed (both in thickness and length)
than the outer; the outer wall flaring, so that the
opening appears large and round or slightly oval.
Towards the middle of the branches these radials
have a classic 'cochleariform' appearance. Both
septal cycles are usually developed, and in some
reef flat and lagoonal specimens these may fill the
corallite. The size of the radials is very even over
the corallum. With increasing water depth the
corallite wall becomes less flaring, the ‘cochlear-
iform’ appearance is lost, and the corallites are
more scattered.
Coenosteum: Either strongly costate or in rows
of simple spines on the radial corallites, reticulate
with simple spines between.
Within Reef Distribution
Reef flat (deeper water areas), lagoonal patch
reefs, reef slope to about 10 m.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
295
Identification Difficulties and History
The synonymy list for A. tenuis is large, and
possibly still incomplete. The synonyms differ in
features such as length of branches and degree of
calcification. In the field, the colonies look quite
similar to those of A. aculeus (Dana). Some field
experience is necessary to distinguish them. The
main differences are:
(a) wide flaring lip and larger radial coralliles
in A. tenuis:
(b) colour pattern: A. tenuis is usually cream
or pinkish brown while A. aculeus has a
distinctive double colouration;
(c) A. tenuis very often has the polyps
expanded, as noted.
The shape of the radial corallites shows some
similarity to that in A. haimei, and strong
similarity to that in A. striata Verrill, which does
not occur in the Great Barrier Reef regions
studied, but is a common species in the Marshall
Islands. One specimen from Lizard Island in the
J.C.U, with bottlebrush growth form is possibly
A. striata.
Geographic Distribution
Indo-Pacific: Mauritius, Red Sea, Diego
Garcia, Indonesia, Philippines, China Sea, Great
Barrier Reef, Fiji Islands.
Acropora tubicinaria (Dana, 1846)
(Plate 74)
Madrepora tubicinaria Dana, 1846, p.451, pi. 32, fig.7;
Brook, 1893, p.l39 (synonymy).
Acropora tubicinaria: Verrill, 1902, p.2I9; Wells, 1954,
p.423, pi. 122, figs.3-5.
Material Examined
USNM: Fiji Islands, U.S. Expl. Exped., A.
tubicinaria holotype 258.
QM: Bushy-Redbill Reef: W. side, inner reef flat:
28.V.1975, C.W., G1 1071-6, G11080; Jan. 1973, C.W.,
G1 1077; 18.xii.l972, C.W., Gl 1081; NW. side, first reef
crest, 22.xii.1972, C.W.. G11082; S. side, microatoll
zone, 2.i.l973, C.W., G11078, G11079, G11083.
Masthead Reef, August 1974, J. Buhmann, G11088.
Great Keppel Island, Dec. 1975, R. Woolley,
Gn089.
Field Diagnosis
Colonies are small rounded caespitose clumps
(largest seen 25 cm diameter) in which the
openings of the radial corallites are obvious.
Colour is brown, occasionally with blue tips.
Laboratory Diagnosis
Branching pattern: From a central growing area
branches are given off vertically to obliquely, and
these branch again, sometimes infrequently,
sometimes frequently. As colonies mature the base
and basal parts of branches may die, and portions
of the colony become separate from each other.
Branches are from 8 to 10 mm thick, and may be
truncate or strongly tapered.
Axial corallites: Non exert. Outer diameter 1-8
to 3-2 mm; inner diameter 1-0 to 1-2 mm.
Septation: both septal cycles developed, primaries
up to 3/4R, secondaries up to 1 /4R.
Radial corallites: Short tubular, usually upper
part of wall less developed than lower, opening
circular. Usually both septal cycles are developed,
but just visible, although on parts of the corallum
primaries can reach 1/2R and secondaries
1/4R.
Coenosteum: On some colonies lines of simple
spines are arranged both on and between radials.
More commonly the coenosteum is costate or in
lines of spines on the radials, and reticulate with
simple spines between.
Within Reef Distribution
On Bushy-Redbill Reef, this species is common
in the coral depauperate inner reef flat (recorded
as A. digitifera by Wallace and Lovell 1977).
With A. palifera and species of Porites it
dominates this zone and it is rare on other parts
of the reef.
Identification Difficulties and History
This species is rarely mentioned in the
literature. On my evidence and that of Wells, it
tends to specialize in certain reef zones (which
may be characterized by poor coral cover), but
is common in these areas. It is rare in collections
brought to the QM for identification, and 1 have
not yet seen it from fringing reefs. A. striata
Verrill mentioned by Wells (1954) as a ‘related
form’ does not occur on the Great Barrier
Reef.
Geographic Distribution
Great Barrier Reef, Fiji Islands, Marshall
Islands, Tahiti.
Acropora aculeus (Dana, 1846)
(Plates 75, 76)
Madrepora aculeus Dana, 1846, p.450, pi. 32, fig. 6;
Brook, 1893, p.I04 (synonymy).
Acropora aculeus: Faustino, 1927, p.269; Nemenzo,
1967, p.lI4.
Madrepora tubigera Horn, 1860, p,435; Verrill, 1864,
p.4!; Quelch, 1886, p.l61; Brook, 1893, p.79 ( +
further synonymies); Verrill, 1902, p.239, pi. 36,
figs. I, 2-2b. pi. 36a, figs.l, 2-2b; pl.36f, fig. 8.
296
MEMOIRS OF THE QUEENSLAND MUSEUM
Acropora tuhigera: Crossland, 1952, p.208.
''f Madrepora ruina Studer. 1878, p.533, pi. 2, figs. 6a,
6b; Brook. 1893, p.82. ‘
Acropora nano: Wells, 1950, p.39, pi. 10, figs. 3, 4.
Acropora nana Nemenzo, 1967, p.85.
Madrepora patula Brook, 1892, p.460; 1893.
p.l 1 1, pi. 9, fig.E.
Acropora paiula: Crossland, 1952, p.215; Stephenson
and Wells, 1956, p.l6; Nemenzo, 1967, p.l02.
‘^Madrepora elegantvla Ortmann, 1889, p.507. pi. 12,
ng.5.
Madrepora elegantula: Brook, 1893, p.l 15.
Mathrial Examinhd
USNM; Fiji Islands. U.S. Expl. Expd., A. aculeus
holotype 257; Cocos-Keeling, A. nana (mentioned Wells,
1950) 44322,
VTM; A. tuhigera fragment of type 1483.
BM: Port Denison, Great Barrier Reef, A. patula
holotype? 1892.6.8.274.
QM; Big Broadhurst Reef: SW, side, reef slope:
I3.X.I973. C.W., G9093; 3 m, 28.iii 1973. C W., G9097;
5 m, 27.iii.1973, C.W., G9I07; 51 m, 23.X.1973,
C.W., G9086. G9087; 6 m. I3.X.I973, C.W., G9092; 6 1
m, 13-X.1973. C.W., G9090; 7 m, 28.iii 1973. C.W..
G9106;8 m, 26.iii 1973, C.W., G9108; 8-2 m, 14 x. 1973.
C.W., G9094: 8-5 m, 23.X.I973, C.W., G9100; 8-7 m,
15.x. 1973. C.W., G9095; 9-2 m, 23.X.1973, C.W.,
G9088; 10 rn, 20.X.I973, C.W.. G9099; 10 m, 23.X.1973,
C.W.,G9101; I()-6 m. 16.x. 1 973, C.W., G9102; 13-3 m.
23.x.I973,C.W.,G9089;25 m, 1 7.x. 1973, C.W., 09091;
25 1 m. 18.X.1973, C.W.. G9I04; SW. side, surge
channel Boor. 10 m, 20.x. 1 973. C.W., G9098,
G9108.
Bushy- Redbill Reef; W. side, middle reef flat:
8.vi-t975, CW, G9122-4: 14.vi.l975, C.W., G9125.
G9126; W side, outer reef flat, 31. vi. 1975, C.W.,
G9I16; W. side, reef slope, 1-8 m, 3.vi.l975, C.W ,
G91 18, C;9120; NW. side, middle reef Hat, 30.xii.I972,
C.W., G9114; NW. side, patch reefs; I9.xii.l972, E.
Lovell. G9542; 15 m, 16.vi.l975. C.W., G9117; NW.
side, reef slope, 3 7 m, 21.xii.l972, C.W., G9115;
adjacent Redbill Is., reef crest: 19.xii 1972, CW,
G9113; I.vi.l975. C.W., G9119; adjacent Redbill Is.,
reef slope, 20.xii.l972, C.W., G11463.
Darley reef, patch reefs in lagoon: 3 m, 18.vii.l972,
C.W., G9I10; 3 m, 22.iii.1973, C.W., G9I09; 4 m.
22.iii.1973, C W., G9111.
Prawn reef, patch reef in lagoon, 15.vii.l972, C.W.,
09112.
Field Diagnosis
This species occurs as Thick plates’ or
caespito-corymbose units. The colony can be one
plate with a loose central or side attachment, or
a number of tiers of plates can develop. The single
plates occur on the outer reef flat and deeper
water parts of the reef slope: the layering develops
in deep water middle reef flat areas and on the
reef crest. Small round colonies can occur on the
outer reef flat. Colours are blue, grey, green, or
brown on lower parts of branches, with tips of
branches yellow, lime green, pale blue or
brown.
Laboratory Diagnosis
Branching pattern: From a usually lateral
attachment, main branches are horizontal and
secondary branchlets or bundles of branchlets are
given off horizontally to obliquely on both sides
of them. Branchlets are slender (3 to 7 mm). The
‘stout branches’ of Dana was either a slip of the
pen or a reference to the main stem. Small shallow
reef flat specimens grow as vertical branchlets
directly from an encrusting base.
Axial corallites: Outer diameter 1-8 to 2-4 mm;
inner diameter 0-8 to 10 mm. Septation: primary
septa well developed, up to 2/3R. secondaries
usually partly but poorly developed, up to
1/4R.'
Radial corallites- Tubular appressed to partly
appressed, with round to slightly oval openings at
90*^ or more to ihe branch. Primary septa
developed up to 1 /2R, and some secondaries
visible. Radials are usually evenly sized and
distributed on branches and show little variation.
Wall can be thickened.
Coenosteum: I.ines of simple spines on and
between radials, or sometimes a spongy appear-
ance between radials.
Within Reef Distribution
Middle and outer reef flats, reef crest, reef slope
to 20 m, lagoonal patch reefs.
Identification Difficulties and History
Much of the variability in this species is related
to colony size. Some idea of the scale of this
variability can be seen by comparing Well’s A.
nana specimen (1950, pi. 10, figs.3, 4) and my
Plate 76, Fig.D. Other features such as thickness
of wall, angle of radial lip, and completeness of
radial septation, vary little, but are responsible for
the various species in synonymy. A. elegantula
(Ortmann) as interpreted by Brook (1893) (BM
1892.12.5.18) is a heavily calcified specimen of
this species.
Geographic Distribution
Indo-Pacific: Cocos-Keeling Islands, Philip-
pines, Great Barrier Reef, Fiji Islands, Samoa,
Marshall Islands.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
297
Acropora cerealis (Dana. 1846)
(Plate 77)
Madrepora cerealis Dana, 1846, p.460, pi. 35, fig.2;
Brook, 1893, (part) p.91 (synonymy).
Acropora cerealis: Faustino, 1927, p.266, pi. 86, figs.l .
3; Nemenzo, 1967, p.83, pi. 25, fig.2,
Madrepora hysirix Dana, 1846, p.476, pi. 40, fig. 1,
pi. 31, fig. 5; Brook, 1893, p.l76 (synonymy).
Acropora hystrix: Wells, 1954, p.425; pi. 125,
figs. 1-4.
Madrepora tizardi Brook. 1893, p.89, pl.Il, figs. C,
D. (synonymy).
Acropora tizardi: Wells, 1954, p.425, pi. 1 25, figs. 5,
6; Nemenzo. 1967. p.l03
Material Examined
USNM: Sooloo Sea, U.S. Expi. Exped., A. cerealis
synlype 269; East Indies, U.S. Expl. Exped., A. cerealis
syntype 270, Fiji Islands, U.S. Expl. Exped., A. hvstrix
holotype 298.
YPM: Fiji Islands, U.S. Expl. Exped., A. hystrix
fragment of type 2039.
BM: Tongatabu, J. J. Lister, 1891.3.6.9; Amboina,
Challenger, 1885,2.1.1; A. cerealis (Brook 1893
mentioned specimens); Tizard Bank, A. tizardi syntype
1889.9.24.1 15.
QM: Big Broadhurst Reef, SW. side, reef slope: 4-5
m, 12.X.1973, C.W., G9532; 5-1 m, 14.X.I973, C.W.,
G9539; 6 m, 13.x. 1973, C.W., G9535; 6-3 m.
23.X.1973, C.W., G9538, G9531; 8-6 m. 15x1973,
C.W., G9533; 8*7 m. 23.X.1973, C.W., G9529; 10 m.
23.X.I973, C.W., G954I; 14*2 m, 23.x4973. C.W.,
G9530; 13.X.I973. C W., G9537; Oct., 1973, C.W.,
G9540.
Bowden Reef, slope of opening in SW. side, 3 m,
26.vii.1972, C.W., G9519. G9520.
Bushy-Redbill Reef, NW. side, reef crest, 22.xii.1972,
C.W., G9543.
Darley Reef, patch reefs in lagoon: 5 m, 24.iii,1973,
C.W., G9524; 1 m, 22.iii.1973, G9525, G9528; 3 m.
!8.vii.l972, C.W., G9526.
Viper Reef, patch reefs in lagoon, 3 m, 6.vii.l972,
C.W.. G9521. G9523.
Field Diagnosis
Colonies can be untidy caespitose clumps in
lagoonal situations, through caespito-corymbose
on the upper reef slope, to corymbose plates on
the deeper reef slope. Radial corallites are narrow
tubo-nariform, with tendency for outer edge to be
hooked upwards ('A. tizardi’}. with elongate
oblique opening (‘A. hystrix) tubo-nariform
tending to nariform (A. cerealis s.s.). The
resultant appearance is a colony similar to A.
nasuta, but with ‘spinier’ appearance and slender,
usually more complexly branching, branchlets.
Colour is usually cream to pale brown.
Laboratory Diagnosis
Branching pattern; From a central to lateral
attachment, main branches are given off vertically
(mainly in lagoonal colonies) to horizontally.
Secondary branching is on all sides of vertical
branches or vertically to obliquely upwards from
horizontal branches. There is a tendency for
further secondary branching. Branchlet widths are
around 10 mm, but much of this width is given
by the spreading radial corallites.
Axial corallites: Outer diameter 1-8 to 21 mm,
inner diameter 0-7 to 0-9 mm. Septation: primary
septa well developed, up to 3/4R, secondaries
absent or a few present to 1 /4R.
Radial corallites: The shape of the radial
corallites is nariform to tubo-nariform, but the
opening varies from rounded to slightly oval
(lagoonal specimens) to accentuated elongate. The
lower wall may be slightly thickened or may be
extended as a ‘hook’. In lagoonal specimens,
radials tend to be scattered. In reef-slope
specimens with elongate radials these tend to be
arranged in neat rows along the branches. Septal
development varies, but primaries are always
strongly visible, and secondaries usually at least
partially visible.
Coenosteum: Costate or dense lines of spines on
corallites. reticulate with simple spines in
between.
Within Reef Distribution
Deep water reef flat areas, reef slope (to 15 m
in present study), lagoonal patch reefs.
Identification Difficulties and History
Wells (1954) commented on the similarity
between A, tizardi and A. hystrix. Unfortunately
A. cerealis sens, strict, is the most difficult of the
synonyms to place correctly. Without the
accentuated oblique radial corallile opening it
approaches A. nasuta in general appearance.
Deeper reef slope specimens are very lightly
structured, and are easily confused with specimens
of A. tenuis from similar localities
Geographic Distribution
China Sea, Philippines, Great Barrier Reef, Fiji
Islands, Marshall Islands, Tongan Islands.
Acropora nasuta (Dana, 1846)
(Plate 78)
Madrepora nasuta Dana, 1846. p.453, pi. 34, rig.2;
Brook, 1893, p.73 (synonymy).
Acropora nasuta: Verrill, 1902. p.257; Hoffmeister,
1929, p.364; Wells, 1954, p.424. pi. 1 13, figs. 5, 6;
pi. 124, figs. 1-3; Nemenzo, 1967, p.88, pi. 26,
ng.3.
298
MEMOIRS OF THE QUEENSLAND MUSEUM
Madrepora nasuta var. crassilahia Brook, 1 893,
p.74.
Acropora nasuta crassilahia: Wells, 1954, p.425,
pi. 1 24, fig. 4.
Acropora nasuta var. crassilahia: Nemenzo, 1967,
p.89, pi. 26, fig.2.
Madrepora effusa Dana, 1846, p.455; Brook, 1893,
p.76 (synonymy).
Acropora effusa: Verriil, 1902, p.229, pi. 36, fig.l6,
16a, pl.36B, fig.7, 7a.
Madrepora cymhicyathus Brook, 1893, p.86
(synonymy).
Acropora cymhicyathus: Hoffmeister, 1925, p.63,
pi. 13, figs.2a, 2b; Wells, 1954, p.425; pi. 124,
figs. 5-7; Stephenson and Wells, 1956, p. 12.
Matf.rial Examined
USNM: Tahiti, U.S. Expl. Exped., A. nasuta holotype
260.
YPM: Point Pedro, Ceylon, A. effusa holotype
8147.
QM: Big Broadhurst Reef: patch reef in lagoon:
21.X.1973, C.W., 011212; 28.iii.1973, C.W.. G11227;
SW. side, outer reef fiat, 22.iii.!973, C.W„ G11209;
SW. side, reef crest: 1-5 m, Il.x.1973, C.W., GI1216;
1-5 m, 22.X.I973, C.W., G11213; 2 m, ll.x.1973, C.W..
G1 1215, 011274; SW. side, reef slope: 7 m, 28.iii.1973,
C.W., G11224; 7-6 m, 13.X.1973, C.W., G11217.
Bowden Reef, SW. side, reef crest, 1 m, 15.vii.l972,
C.W., G1I21I.
Bushv-Redbill Reef; adjacent Redbill Is., reef flat:
18.xii.r972, C.W., GI1204; 7.vi.l975, C.W., 011222;
12.vi.l975, C.W., 01 1223; W. side, middle reef fiat;
5.vi.l975, C.W., GI1226; 27.vi.l975, C.W., 01 1221,
01 1228; NW. side, first reef crest, 22.x. 1973, C.W.,
011214.
Parley Reef, patch reef in lagoon, 3 m, 22.iii.!973,
C.W., GI1218.
Heron Island: W. side, outer reef flat, 7.vii.l973,
C.W., 011209; W. side, reef flat, .luly 1973, C.W.,
011205; W. side, reef slope, 7 m, 7.vii.l973, Y. Loya,
011203.
Palm Islands, 1939, T. C. Marshall, 011206-8.
Field Diagnosis
Corymbose to tabular colonies, the branchlets
covered evenly and neatly with nariform corallites
with elongate openings. Colour is most commonly
cream to pale brown, usually with pale blue tips,
but can also be yellowish or greenish.
Laboratory Diagnosis
Branching pattern: Branches arise from an
encrusting plate, or a sturdy stalk. Secondary
branches are vertical in the centre of the colony,
and more curved towards the edges; they are not
usually very proliferous.
Axial corallites: Outer diameter 2 0 to 3*0 mm;
inner diameter 0-5 to 0-9 mm. Septation: primary
septa present, up to 3/4R, secondary septa
anything from all absent to all present up to '
1/4R.
Radial corallites: Nariform, dimidate, or
tubo-nariform, opening at 90" to branch or less.
The walls of the corallites may be thickened.
Primary septa present up to 1/3R, secondary
septa poorly developed, usually less than a full
cycle present, up to 1/4R.
Coenosteum: Laterally flattened or forked
spines are arranged densely on radial corallites,
sometimes joining as costae. Between radials
coenosteum is reticulate.
Within Reef Distribution
Occurs on most parts of the reef flat, reef crest,
upper reef slope (to about 8 m in studied areas).
This is sometimes the only Acropora (other than
A. palifera) occurring in shallow lagoonal patch
reefs in inner lagoons adjacent to coral cays.
Identification Difficulties and History
The close similarity of A. nasuta and A.
cymhicyathus has been commented on by other
authors. Although not well represented in the
literature, this is a common Acropora on the
Great Barrier Reef, occurring in a variety of
habitats. Colonies are often very regularly formed,
and easily recognised, although it can sometimes
be confused with more regular colonies of A.
cerealis. It is a good photographic subject, usually
being identified in popular texts as ‘A.
surculosa'.
Geographic Distribution
Indo-Pacific: Ceylon, Great Barrier Reef, Fiji
Islands, Samoa, Marshall Islands, Tahiti.
Acropora diversa (Brook, 1891)
(Plates 79, 80A, B)
Madrepora diversa Brook. 1891. p.46I; 1893, p.l41,
pi. 16, fig.B.
Acropora diversa: Wells, 1954, p.424, pl.l 17, figs. 3-6;
Stephenson and Wells, 1956. p.l3.
Madrepora concinna Brook. 1891, p.460; 1893, p.I65,
pi. 17.
Acropora otteri Crossland, 1952, p.229, pl.43, figs.l,
2, pl.44, figsJ, 2.
Material Examined
BM: Diego Garcia, G. C. Bourne, A. diversa holotype
1891.4.9.4; Mauritius, A. concinna syntypes 1878.2.4.3,
1878.2.4.8; Great Barrier Reef Expedition, A, otteri
syntypes 1934.5.14.17; 1934.5.14.76; June Reef, outer
moat A. otteri (? also a syntype) 1934.5.14.315.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
299
QM: Big Broadhurst Reef: SW. side, outer reef flat*
22.X.1976, C.W.. GM232. GI1233; 18.x 1973, C.W.,
G1 1261, Gl 1263, G1 1269, Gn279, 011280; 22,x.l973,
C.W., G1I272, G11285; SW. side, reef slope: 1-3 m,
22.X.1973, C.W., G11245; 1-5 m. 18.x. 1973, C.W.,
G11266; 3 m, 27.iii.l973, C.W., 011254; 3-3 m.
22.X.1973, C.W., 011237; 4-5 m, 23.X.1973. C.W.,
G1 1273; 51 m, 23.x. 1973, C.W., G11236, G11270; 5-7
m, 23.x. 1973, C.W., G1I241; 6 m. 27.iii.1973, C.W.,
011252; 7 m, 28.iii.1973, C.W., 0112.59; 7-1 m,
13.X.1973, C.W.. 011276; 7-8 m, C.W., 011267; 8 m,
28.iii.1973, C.W., 011246, 011258; 8-1 m, i4.x.l973,
C.W., 011231; 8-2 m, 14.x. 1973, C.W., 011230,
01 1243, G1 1285; 8-6 m. 15.x. 1973, C.W-, 01 1240; 8-7
m, 15.X.1973. C.W., 01 1264; 9-6 m, C.W., 01 1265; 9-7
m. 15.X.1973, C.W.. 011271; 9-8 m, 16.x. 1973, C.W.,
011239; 10-6 m, 23.x. 1973, C.W.. 011275; 12-5 m,
I7.X.1973, C.W., 011242; 14-2 m, 23.X.1973, C.W.,
01 1235; SW. side, reef crest; 1-5 m, 11.x. 1973. C.W.,
011278; 1-6 m. 22.X.1973, C.W., 011284; 22.x 1973.
C.W., 011234.
Bowden Reef, SW. side, reef crest; 0-75 m,
1 5.vii.l972, R. Pearson, Gl 1 268; 1 m, 15.vii.l972, C.W.,
011248, G11251, 011260; SW. side, reef slope:
26.vii.1973. C.W., 011238.
Bushy-Redbill Reef, NW. side, reef slope: Jan. 1973,
C-W , 011249; 19.xii.l972, E. Lovell. 01 1257; GI 1262;
011282.
Darley Reef, patch reef in lagoon: 1 m, 24.iii.1973,
C.W., G11250; 5 m, 19.vii.l972, C.W., 011244,
011256; 24.iii.1973, C.W., 011253, 011255.
Field Diagnosis
Corymbose, caespito-corymbose, tabulate to
plate-like colonies have a mixture of tall and short
tubular radial corallites which is usually
distinctive. A variety of colours occur, vis. cream,
yellow-brown, blue-grey, blue-brown.
Laboratory Diagnosis
Branching pattern: From an attachment region
which is central to lateral, branches are given off
upwards vertically to obliquely. These may be as
short as 20 mm to as long as 70 mm, and may
branch proliferously or rarely. Branch widths vary
from 7 to 20 mm.
Axial corallites: 1 to 3 mm exert. Outer
diameter 2-4 to 3-2 mm (slightly larger in A.
diversa type); inner diameter 0-8 to 1-1 mm.
Septation: both cycles usually present, the first
cycle up to 3/4R, the second up to 1/3R.
Radial corallites: Tall and short corallites are
mixed, in a pattern which may be regular (tall
radials arranged in vertical rows) or very
irregular. The form of the tall and short radials
is similar — tubular, with round, or slightly oval
openings, sometimes tubo-nariform. The outer
wall may be thickened. The septal development
is usually poor, primary septa being present, up
to 1/3R, but usually less than this, the secondary
cycle usually only partly present, up to 1/4R.
Coenosteum: Simple pointed to laterally
flattened spines are densely arranged on the radial
corallites, sometimes forming costae. Between the
radials their arrangment is less dense.
Within Reef Distribution
Outer reef flat, reef crest and reef slope to about
15 m, lagoonal patch reefs.
Identification Difficulties and History
Amongst my material there is great variability
in radial corallite features such as length, shape
of opening, ratio of short to long, as well as the
expected colony-shape variability. Both in the field
and in the laboratory I find areas of overlap and
difficulty in distinguishing some specimens from
A. nasuta and others from A. variabilis. A study
concentrating on these three species would be
profitable.
Geographic Distribution
indo-Pacific: Mauritius, Diego Garcia, Great
Barrier Reef, Marshall Islands.
Acropora variabilis (Klunzinger, 1879)
(Plate 80C, D)
Madrepora variabilis Klunzinger, 1879, p.17, pl.l,
fig.lO, pl.2. figs.I, 5, pi. 5, figs.l, 3, pl.9, fig.l4;
Brook, 1893, p.l61.
Acropora variabilis: von Marcnzeller, 1907, p.49,
pi. 15, figs. 4B44; Vaughan, 1918, p.l81, pi. 80.
figs.2. 3, 3a, 3b; Faustino, 1927, p.276; Wells, 1950,
p.38; 1954, p.428, pi. 128, figs.l, 2, pi. 130, figs. 1,
2; Rossi, 1954, p.52; Stephenson and Wells, 1956,
p.l9; Scheer and Pillai, 1974, p.23, pi. 8, fig-2; Pillai
and Scheer, 1976, p.31.
Acropora variabilis var. pachyclados: Crossland,
1952, p.222, pi 38, figs.l, 6.
Matf.riai Examined
MNB: Koseir, Klunzinger, A. variabilis var.
pachyclados 2118; Koseir, Klunzinger, A. variabilis var.
cespiiofoliata 2120 (Klunzinger mentioned specimens,
examined as photographs only).
QM- Big Broadhurst Reef, SW. side, outer reef flat:
11.X.I973, C W.. GII291; 18.x 1973, C.W..
011287-90.
Field Diagnosis
Colonies may be small round clumps of vertical
to oblique branches or stalked corymbose colonies
to about 30 cm across. Branches are covered by
appressed tubular corallites with round openings.
The sizes of the radial corallites may be similar
or extremely variable. Colour may be brown.
300
MEMOIRS OF THE QUEENSLAND MUSEUM
lavender-brown, greenish-brown, or yellow or
cream with purple corallites.
Laboratory Diagnosis
As this species has been well described by other
authors, and my series is poorly representative of
the species, further description is not given. Refer
to Vaughan 1918 for tables of measurements of
skeletal features.
Within Reef Distribution
Shallow outer reef flat (particularly where an
elevated platform is present), reef crest. A large
series in the JCU comes from the fringing reefs
of the Palm Islands and Lizard Island.
Identification Difficulties and History
Although this species is poorly represented in
my areas of study, evidence suggests it may be
common (a) on windward outer reef flat platforms
(as small round clumps) and (b) on reef edges of
the fringing reefs of continental islands (as
caespito-corymbose colonies). The series in JCU
from Lizard Island and the Palm Island group is
in the second category. An allied species is A,
valida (Dana) (see Hoffmeister, 1925, p.6()). This
latter species may also occur on the Great Barrier
Reef, but is not represented in the QM collections,
except by two specimens collected in the Fiji
Islands, and closely comparable with Dana's
type.
Geographic Distribution
Indo-Pacific' Red Sea, Nicobar Islands,
Cocos- Keeling, Philippines, Great Barrier Reef,
Marshall Islands.
Acropora hiimilis (Dana, 1846)
(Plates 81, 82, 83)
Modrepora humilis Dana, 1846, p.483, pl.31, fig. 4,
pi. 41, fig. 4.
Acropora humilis: Wells, 1954, p.425, pi. 1 00, fig. 1.
pLl26, figs.l^, pi. 1 27, figs.3, 4, pL128. figs. 3-5
(synonymy), Rossi, 1954. p 50; Slephen.son and
Wells, 1956, p.l5; Pillai and Scheer, 1976, p.32.
This species has been given extensive taxonomic
treatment by Wells 1954. He combined 17 species
(some with additional synonyms designated by
earlier authors), and concluded that three broad
forms could be recognised. These he considered
to be characteristic of different reef localities and
related, in particular, to water level. Although a
large suite of specimens is on hand at the QM,
their description is withheld until a further study
concentrating on this complex species can be
carried out. For general purposes, this species is
probably the best known and most easily
recognised Acropora. Some additional informa-
tion is added below on Well’s first facies, forma
alpha {A. samoensis, A pelewensisj.
Acropora humilis (Dana, 1846) forma a Wells,
1954
MatfriaI- Examined
QM‘ Big Broadhurst Reef, SW. side, reef slope: 8 6
m, 14.X.1973, C.W , G1 132, Gl 135; 8-6 m, 15.X.1973,
C.W., 011131; 8-7 m. 15.X.1973, 011130; 9-8 m,
16.x. 1973, G11134; 10-4 m, 16.x, 1973, C.W.. G11133;
12-5 m, I7.X.I973. C.W., G11I37.
Bowden Reef, SW side, upper reef slope, 26.vii.1972,
C.W., G11I89.
Darlev Reef, patch reef in lagoon: 24.iii.l973, C.W..
Gl 1 128; 3 m, 22.iii.l973, C.W., GI 1129; 22.iii.1973, 5
m, C.W., GI16IL
Field Diagnosis
Colonies may be low arborescent (with
shrub-like growth) caespito-corymbose. or plate-
like. Branches are terete (non tapering) or slightly
tapering, with wide axial corallites, and large
thick-walled radial corallites. Colours are
cream-brown, sometimes with pale blue tips, or
pale blue or lavender.
LABORArORY DIAGNOSIS
Branching pattern: Superimposed on the colony
shape described above is usually a high degree of
budding. The branches may be as broad as 20 mm,
but in the plate like colonies (which occur on reef
slopes) and particularly in deeper-water
specimens, branches can be as narrow as 8
mm.
Axial corallites: About 2 mm exert. Outer
diameter 3-0 to 5 0 mm; inner diameter 11 to 1-6
mm. Septation: both cycles present, primaries up
to 3/4R, secondaries up to 1 /2R.
Radial corallites: Tubular or appressed tubular,
with round or oval openings, or nariform, the outer
wall thickened. Primary septa are usually present,
up to 1 /3R; secondary septa not usually fully
developed, but some present up to 1/4R.
Coenosteum: Costale to reticulate both on and
between radial corallites, with spines that are
laterally flattened and sometimes slightly
elaborated at the tip.
Within Reef Distribution
This form extends down the reef slope and
sloping surfaces of lagoonal patch reefs.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
301
Identification Difficulties and History
Deep water specimens, because of the narrow
and proliferous branches, are barely recognisable
as A. humilis. and begin to have a similar general
appearance to A. sarmenlosa.
Geographic Distribution
A. humilis has a wide Indo-Pacific
distribution.
Acropora digitifera (Dana, 1846)
(Plate 84)
Madrepora digitifera Dana, 1846, p.454; Brook,
1893, p.75.
Acropora digitifera: Verrill, 1902, p.228, pi. 36, fig. 12,
pl.36B, fig.3; Vaughan, 1918, p 175, pi. 13, fig, 7;
pi. 76, figs. I, la, 2; Wells, 1954, p-427, pi. 127,
figs.l, 2; 1955. p.9; Stephenson and Wells, 19^6,
p.l3 (synonymy).
Acropora fraierna Verrill, 1902. p.247, pi 36, fig. 18,
pi 36B, fig.9.
Materia! Examined
YPM: A. digitifera type (fragment, ex. Boston Soc.
Nat. History) 4192; Tahiti, U.S. Expl. Exped., A
fraierna holotype 2032.
QM: Big Broadhurst Reef, SW. side, outer reef flat:
lLx.1973, C.W., G11170; 18.X.I973, C.W., GI1169,
G11171, Gni72; 22.X.I973. C.W., G11168, 011187.
Bushy-Redbill Reef, adjacent Redbill Is., reef flat,
I8.xii.I972, C.W., GII176; 2.vi.l975, C.W G11173,
GI1I74; 5.vi.l975, C.W., 011175; 10.vi.]975, C.W.,
G11188; Il.vi.l975, C.W., G11177.
Heron Island, W. side, reef flat. July 1973, C.W.,
G11180.
Oyster Cay, C. Limpus, 1975, G11179
Fiji Islands: Great Astrolabe Reefs, Yaucuve Leva
fringing reef, Jan. 1974, C.W., G11183, G11184,
G1II86; Vaga Bay, Beqa, patch reefs, 15. i. 1974, C. J.
Wallace, G1 1 182.
Field Diagnosis
Colonies are corymbose, with central to lateral
attachment, and tapering branches. Appearance is
of a diminutive, ‘neat’ A. humilis. Colour is
commonly cream to pale brown, with or without
blue branch tips.
Laboratory Diagnosis
Branching pattern; From a growing area which
may be central to lateral, and more or less
stalk-like, main branches grow horizontally, and
give off secondary branches or bundles of
branches vertically to obliquely. These may taper
to a point, or be terete, and are from 8 to 20 mm
greatest width
Axial corallites: Non exert. Outer diameter 2-8
to 3-8 mm; inner diameter 0-8 to 11 mm.
Septation: both cycles developed, primaries up to
2/3R, secondaries up to 1/4R.
Radial corallites: A size gradation occurs from
branch tip to proximal and small corallites are
interspersed with large. Shape is dimidiate, or
tubular with oval to dimidiate opening, radials
spreading at 90° from branch. Outer wall is
thickened.
Coenosteum: Costate or with regular lines of
spines on corallites, spongy with spines in
between.
Within Reef Distribution
Limited to the shallow middle reef and outer
reef flat pavement (‘A. digitifera' from inner reef
flat as interpreted by Wallace and Lovell 1977 is
A. tubicinaria.)
Identification Difficulties and History
On some reefs (e.g. Enewetak, Marshall
Islands) this species is well marked and easily
recognised. On the Great Barrier Reef, where A.
digitifera occurs with A. humilis. some colonies
cannot be definitely assigned to one or other
species on morphological grounds. As Stephenson
and Wells (1956) point out, the main difference
is in dimensions. A further analysis of one must
include the other
Geographic Distribution
Great Barrier Reef, Moreton Bay (Queens-
land), Fiji Islands, Marshall Islands, Tahiti.
Acropora multiacuta Nemenzo, 1967
(Plate 85)
Acropora multiacuta Nemenzo, 1967, p.l33, pl.39,
figs.l, 2. 3; Scheer and Pillai 1974, p 24, pi. 6,
fig-4.
Material Examined
USNM: (donated): Darley Reef, shallow patch reefs
in lagoon, 1 m, 24.iii.l973, C.W.
BM: (donated): Darley Reef, shallow patch reefs in
lagoon, I m, 24.iii.l973, C.W.
JCU: Philippine Islands, 1975: 3 m, M. Pichon.
2509/75; 5 m, M. Pichon, 2525/75.
QM: Darley Reef, shallow patch reefs in lagoon: 1
m, 19.vii.l972, C.W., G6721, G6722; upper surface,
23.iii.l973, C.W., G10465, G10470, G10471: upper
surface, 24.iii.I973, C.W., G10466, GI0467. GI0469;
1 m, 24.iii.I973, C.W., G10464; 3 m, 22.iii.1973, C.W.,
G 10468.
302
MEMOIRS OF THE QUEENSLAND MUSEUM
Field Diagnosis
Irregular caespitose to caespito-corymbose
colonics from an encrusting central to lateral base,
with axial corallites sturdy and prominent,
sometimes to the extent of the entire branch or
one side of the branch being naked of radial
corallites. Colour is whitish blue with pale blue
polyps. The largest colony seen to date is 19*5 cm
diameter.
Laboratory Diagnosis
Only twelve Great Barrier Reef specimens have
been examined and these show much variation,
particularly in the extent of radial corallite
presence on the branches, the amount of
secondary branching, and the length of the main
branches. Some dimensions such as the width of
the branches are not included in the diagnosis for
this reason.
Branching pattern; From the encrusting base,
the main branches arise vertically to obliquely,
and may be variously curved. The longest branch
in the present collection is 10 cm. At the base of
main branches smaller branches may occur. These
do not always alter the colony shape, but in the
side attached specimens it appears they would
contribute to the development of a bracket shape
as the colony matured. Some specimens have
prolific incipient branching along the main and
secondary branches.
Axial corallites: (On the main branches) outer
diameter 3-5 to 6-5 mm; inner diameter 10 to 2-0
mm. Seplalion; both cycles present, occasionally
a third cycle partially developed, primaries up to
1/3R, secondaries up to 1 /4R. On the basal and
incipient branches axials arc narrower and
sometimes compressed so that the opening is oval.
ThCvSe are described as radial corallites by
Nemenzo.
Radial corallites: Scattered, nariform, tubo-
nariform, or partly appressed tubular; often
oriented with opening down or across the branch.
Septa not developed at all, or primaries just
visible.
Coenosteum: Densely echinulate on both radial
corallites and inter-corallite areas; the spines
laterally flattened, occasionally some pseudocos-
tatc development on the axial corallite.
Within Reef Distribution
The only populations seen have been on the
upper surface and edges of patch reefs, just below
low water. The colonies occur on the surface or
in depressions in the irregular reef surface, and
the main branches may be curved to maintain an
overall vertical orientation. The naked areas of
branches are always upwards.
Identification Difficulties and Notes
Nemenzo used the species grouping Alticyathus
for this species and for his A. fastigata (1967,
p.l34). The latter may be a synonym, or may be
A. digitifera. I have not been able to locate
Nemenzo’s types in the UP collections. On the
Great Barrier Reef, the species has only been seen
from the Darley Reef lagoon, and it cannot yet
be fully categorised.
Geographic Distribution
Indo-Pacific: Nicobar Islands, Philippines,
Great Barrier Reef.
Acropora clathrata (Brook, 1891)
(Plates 86, 64C).
Madrepora clathrata Brook, 1891, p.459; 1893, p.49,
pi. 5, pl.6, Hg.A, B.
Madrepora orbicularis Brook, 1892, p.460; 1893,
p.37, pi. 2 (synonymy);
Madrepora vasiformis Brook, 1893, p.37, pi. 26, fig.
A (Synonymy).
Acropora vasiformis: Pillai and Scheer, 1976, p.27,
pi. 3, fig. 4.
Acropora tutuilensis (part) Hoffmeister, 1925, p.71,
plate 19, figs, la-le.
Material Examined
BM: Mauritiu.s, A. clathrata holotype 1893.4.7.78;
Ceylon, A. orbicularis holotype 1883.3.24.7; Rodriguez,
A. vasiformis holotype 1876.5.5.92.
USNM: Pago Pago Harbour, Tutuila, Samoa, A.
tutuilensis no. 1; ?nos. 2, 3. (Mayor collection)
(mentioned Hoffmeister, 1925).
QM: Big Broadhurst Reef, SW. side, reef slope: 5 m,
25.111.1973, C.W., G9753; 8 m, 26.iii.1973, C.W.,
G9752; I m, 27.iii.1973, C.W„ G9748; 3 m, 27.iii.I973.
C.W., G9749; 4 m, 27.iii.I973, C.W., G975I; 5 m,
27.111.1973, C.W., G9750: 1 m, 13.x. 1973, C.W., G9742,
G9746; 6-3 m, 13.x. 1973, C.W., G9735; 7 m, 13.X.1976.
C.W., G9743; 71 m, 13.X.1973, C.W.. G9740, G9741;
7-3 m, 14.X.1973, C.W., G9739; 8 m, I4.X.1973, C.W.,
G9744, 8-1 m, 14.X.1973, C.W., G9738; 19-7 m,
15.X.1973, C.W., G9737. 4 m. 20.X.I973, C.W., G9745;
10 m, 20.x. 1973, C.W., G9759; 1*9 m, 22.X.1973, C.W.,
G9747; 3*3 m, 22.X.1973, C.W., G9736.
Bowden Reef: slope of opening in SW. side: 0-5 m,
26.vii.1972. C.W., G9760; I m, 26.vii.1972, C.W.,
G9761; SW. side, reef slope, 1 m, 15.vii.l972, C.W.,
G9762.
Bushy-Rcdbill Reef: NW. side, reef slope, Dec. 1972,
C.W., G9754; NNE. end, 30.xii.l972, C.J.W., G9755;
S. end. reef slope, 3 m, 3.vi.l975, C.W., G9756; reef
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
303
slope adjacent Redbill Is., 0-5 m, l.vi.l975, C.W.,
G9758; 1 m, 2.V.1975, C.W., G9757.
Parley Reef: patch reef in lagoon, 3 m, 18.vii.l972,
C.W., G9763.
Field Diagnosis
Horizontal plate (reaching approx. 1-2 m
diameter), developing by unilateral expansion
from a vasiform shape with a single attachment,
which becomes lateral as the colony develops. The
plate has a flat appearance, due to the orientation
of the secondary branchlels in the plane of the
plate or obliquely to it. Common colours are
pinkish brown, apple green and yellowish
green.
Laboratory Diagnosis
Branching pattern; From the attaching stalk,
branching is oblique to horizontal. Secondary
branches arise in the same plane as primary
branches, or obliquely to them, forming a single
layer or a series of interlacing layers. All branches,
except those towards the edge zone, are of similar
diameter. Anastomosis of branches varies, so that
anything from an open network to a solid plate
(with branches either fused to the plate or
projecting obliquely from it) is possible.
Axial corallites: Outer diameter 1-5 to 2*2 mm,
inner diameter 0-6 to 0-9 mm. Septation: primary
septa present, up to 1 /3R, secondaries usually not
visible, or a few present to less than 1/4R.
Radial corallites: A number of shapes are
possible, and specimens may possess all, some, or
only one of the possible types, viz. tubular, with
round, oval, or dimidiate openings, tubo-nariform,
nariform, rostrato-nariform, dimidiate, sub-
immersed or immersed. Immersed corallites
usually only occur along lines of fusion of
branches. The other types project from the
branch, at from 90° to 45°, occasionally less.
There is a tendency for differential thickening or
extending of the longer wall sometimes with
bizarre extensions such as long points on nariform
and horns on dimidiate corallites. Plate 86
shows:
(D) all uniform size, nariform or dimidiate
with differential outer wall development.
(B) mixed size, tubular with round, oval, and
dimidiate openings, immersed and
sub-immersed.
(F) similar to B, but corallites more
appressed and scattered.
Septation: septa are never well developed;
usually only the directive septa are visible, or a
few other primaries present as small spines. As
the corallites are rarely appressed, they appear
well spaced and there is always some coenosteum
between.
Coenosteum; On unthickened corallites this
may be visibly porous and is usually costate, on
thickened radials it is composed of compactly
arranged flattened spines, sometimes pseudocos-
tate. Between corallites it is reticulate.
Identification Difficulties and History
This coral occurs with the other large
horizontal plate Acropora {A. hyacinthus and A.
cytherea), from which it is easily separated by its
flattened appearance and by the lack of scale-like
corallite lips. Identification difficulties occur at
the laboratory stage, where so many different
combinations of colony features and corallite size
and shape are seen. This variety is demonstrated
in the synonymy: A. clathrata s.s. has narrow
branches, openly reticulated, and radial corallites
are tubular, with a variety of sizes and
opening-shapes; 'A. orbicularis’ and ‘A. vasifor-
mis’ are almost solid plates, ‘A. vasiformis' with
rostrate thickening of many radial corallites, A.
orbicularis’ with unthickened radials in a variety
of shapes. Hoffmeister’s no. 1. specimen of A.
tutuilensis is a partly fused plate of this species.
His no. 4 is probably A. rotumana. All the type
specimens can be compared with specimens in the
QM series. A. stigmataria (Milne Edwards and
Haime) (see Brook 1893, p.5U), may be a senior
synonym but its type cannot be located for
examination.
Within Reef Distribution
The species appears to be restricted to sloping
surfaces or good water cover. Very large
specimens can occur around low water mark on
the edge of surge channels, on the upper reef
slope, and on the floor of surge channel openings
(see Plate 64C).
Geographic Distribution
Indo-Pacific: Mauritius, Rodriguez, la Reunion
(G. Faure pers. comm.), Ceylon, Seychelles (?),
Great Barrier Reef, Samoa.
Acropora divaricata (Dana. 1846)
(Plates 87, 88)
Madrepora divaricata Dana, 1846, p.477, pl.41, fig. 2;
Milne Edwards and Haime, 1860, p.l40; Brook,
1893, p.64.
Madrepora tenuispicata Studer, 1880, p.20, figs. la,
lb; Brook, 1893, p.96.
304
MEMOIRS OF THE QUEENSLAND MUSEUM
Acropora lenuispicata: Piliai and Scheer, 1974, p.455
fig. 4b.
'1 S-iadrepora complanata Brook, 1891 p 459- 1893
p.70, pl.4, fig.C.
Acropora complanata. Pillai and Scheer, 1976. p.28.
pi. 7. fig,2,
?Madrepora complanata var. informis Brook 1893
p.7L
Material Examined
USNM: Fiji Islands, U.S. Expl. Exped., A. divaricaia
holotype 299.
YPM: Fiji Islands, U.S. Expl. Exped., A. divaricata
fragment of type 2008.
BM: Seychelles, H M.S. Alert, A. complanata
syntypes 1882.10.17.140, .147, .148; Macclesfield Bank,
13 fathoms, A complanata var. informis syntypes
1892.10.17.71, .72, .73.
Hessisches Landmuseum: Acropora tenuispicata
(colour transparencies only) (mentioned Pillai and
Scheer 1974).
QM; Big Broadhurst Reef: SW. side, reef slope; 12-4
ni, 23.x. 1973, C.W.. G9I62; II m, 16,x.l973, C.W.,
G9166; 9-8 rn, 16.X.1973, C.W.. G9167; 9-6 m,
15.x. 1973, C.W., G9164; S-7 m, 15.x. 1973, CW,
G9170. G9I73; 8-6 m, I5.X.1973, C.W., G9I74, G9175;
83 m, 15.x. 1973, C.W., G9I63, G9172; 81 m]
I4.X 1973, C.W., G9161. G9168; 8 m, 14 x 1973, C.W.,
G9160. G9I65. 09178; 7-8 m, 14.X.1973, C W., G917U
G9I76, G9177; 7-6 m, 28.iii.1973, C.W., G9180; 7 1 m,
13 X.1973, C.W., G9169; 7 m, 28.iii.1973, C.W., G9181,
G9188, 6 m, 26.iii,1963, C.W., G9I79; 6 m, 25 iii 1973
C.W . G9184.
Bushy-Redbii! Reef: W. side,, reef slope; 8 in.
3.viT975, C.W., G9186; 2 m, 3.vi.l975, C.W., G9185;
NW. side, reef patches, 12 m, I4.vi.l975. CW
G9187.
Darley Reef: patch reefs in lagoon; 7 m, 22.iii.1973,
C.W, G9182; 3 m, 18.vii.l972, C.W., G9I84-
22.iii.l973, C.W., G10219.
Fiji Islands, Great Astrolabe Reefs (Kadavu), W. side
of Yaukuve Levu, fringing reef, l.ii,1974, C.W., G9781
G9782.
Fiflu Diagnosis
Occurs as bracket like colonies with central to
lateral attachment, ‘corymbose’ in having all
branches reaching up to a horizontal plane, and
‘caespitose’ in having divaricate branching within
the boundaries of the colony shape. Reaches
approximately 50 cm diameter. Branches appear
rough because of projecting radial corallites.
Colour is usually a drab dark brown or dark
brown with blue tips.
Laboratory Diagnosis
Although the species is easily recognised in the
field, skeletal fragments can be confusing.
Branchlet dimensions and radial corallite shape
are variable amongst colonies in a single
population, and pieces taken from different parts
ol the same corallum may appear different
because of their orientation.
Branching pattern; From a single area of
attachment, branching is central to lateral.
Periferal branches contribute to an oblique
undersurface; inside these is a network of short
branchlets at wide angles, the final branchlets
being erect or nearly so, and ending in a horizontal
plane. Branchlet widths vary from 7 rnm to 15
mm.
Axial corallites; Outer diameter 2-3 to 3 0 mm;
inner diameter 0-8 to 1-1 mm. Septation: primary
septa present, up to 1/2R, secondary septa poorly
developed, but usually some present, up to
1/4R.
Radial corallites: Shape and size of the radial
corallites changes along the branches. On upper
branchlets they are prominent (up to 3 mm long),
usually extending at from 45^^ to 90°. They are
usually tubular on branch tips, passing through
tubo-nariform to nariforin, then rounded to
sub-immersed proxirnally. The prominent radials
are sometimes extended by rostrate development
(see Plate 88 C, D). Within the sequence from
distal to proximal, radials are usually evenly
graded and neatly arranged, but they can be
unevenly graded, so that branches appear ragged
(Plate 88 A), and downward directed radials can
occur anywhere along the branch.
Within Reef Distribution
This coral occurs in the higher diversity (for
Acropora) parts of the reef, where no particular
colony shape predominates: middle reef slope,
deeper outer reef Oats, patch reefs in deep lagoons
and leeward broken reef areas. The species,
although common, has a drab appearance when
alive, and is easily missed when subjective
observation techniques are used.
Identification Difficulties and History
1 he radial corallites of this species are similar
to those of the flat plate species, Acropora
clathrata- I have not found specimens Acropora
divaricata deep enough to form flat plates but am
confident that A. complanata (Brook), a flat plate
species dredged from deep water, represents deep
water flattening of A divaricata. For A.
tenuispicata (Studer) I have drawn on Pillai and
Scheer’s (1974) interpretation of this species. The
type is not in the MNB, where the remainder of
the Studer types are located. The combination of
a determinate bracket with divaricate branching
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
305
is unique in the Acropora. The shape has not been
emphasised in previous descriptions. A series in the
JCLI from Lizard Island fringing reefs shows more
rounded colony shape, with slender branches,
often naked of radial corallites on their upper
surface.
Geographic Distribution
I ndo- Pacific: Seychelles, Singapore, Fiji Is-
lands, Great Barrier Reef.
Acropora sarnientosa (Brook, 1892)
(Plate 89)
Madrepora sarmentosa Brook, 1892, p.462; 1893,
p.I27, pi. 22
Acropora sarmentosa: Nemenzo, 1967, p.90, pi. 26,
fig-4.
non Acropora sarmentosa: Vaughan, 1918, p. 174,
pi. 72, rig.4, pi. 73, fig.l.
Acropora rosaria form I: Crossland, 1952, p.224,
pl.40, fig.3.
Material Examined
BM: Port Denison, Saville Kent, A. sarmentosa
syntype 1892.6.8.228.
UP: Little Balatero Cove, Puerto Gaelera, Oriental
Mindoro, Nemenzo 259; Muelle, Puerto Gaelera,
Oriental Mindoro, Nemenzo 324 (mentioned Nemenzo,
1967).
QM: Big Broadhurst Reef. SW. side, reef slope: 17m,
24.X.1973, C.W., G9064, G9065; I5-7m. 23.X.I973,
C.W., G9062; 15-4m, I7.X.1973, C.W.. G9056; 9-7m,
I5.X.1973, C.W., G9059; 8 Im, 14.x. 1973. C.W., G9055;
8m, 28.iii.l973, C.W., G9054; 7-6m, 13 x. 1973, C.W.,
G9057; 6-8m, 20.x. 1973, C.W., G9058; 41m, 22.x. 1973,
C.W., G9061.
Bowden Reef, slope of opening in SW. side, l-2m,
23.viii.1972, C.W., G9066.
Bushy-Redbill Reef, NW. side, reef slope:
30.xii.l972, E. Lovell, G9071; 3-7m, 2l.xii.l972, E.
Lovell, G9073; NW. side, reef crest, 30.xii.l972. C.W.,
G9070; NW side, patch reefs; 12m, 15, vi. 1975, C.W.,
G9075; 12m, 14.vi.l975, C.W., G9060: SW. side, reef
crest, 24.V.I975, C.W , G9063; outer reef flat adjacent
Redbill Island- Lvi.1975, C.W., G9072; 22.xii.l972, E.
Lovell, G9074.
Darley Reef, patch reefs in lagoon. 6m, 22.iii.1973,
C.W., G9067; 4m, 22.iii.1973, C.W., G9068; 2.5m,
24.iii.1973, C.W., G9069.
Heron Island, W. side, reef flat, 6.vii.l973, Y. Loya,
G9704.
Ellison Reef: dredged 9 fathoms, 25.vii 1924, Dr
Paradice, G9709; 24.vii.1924, C. Hedley, G9707.
Fiji Islands, Great Astrolabe Reefs: Qasilabe fringing
reef, 4^6m. 4.ii.l974, C.W., G97I0; Yaukuve Levu, W.
side, fringing reefs, 8m, l.ii.l974, C.W., G9705.
Field Diagnosis
Colonies of this species usually have few
(commonly 2 or 3) thick rounded branching units,
consisting of a central horizontal to oblique
branch or branches, with vertical branchlet
bundles evenly distributed, but longer on the
upper side. Colonies of more than 50 cm across
are unusual, and attachment is usually from the
side. The overall appearance is smooth — axial
corallites are not exert, radial corallites are large,
evenly distributed, uniformly sized and not
projecting; secondary branching patterns are
regular and branchleis terete. The usual
colouration on the Barrier Reef is two-toned, and
cryptic — most often a dull greenish grey or
-brown, with pale brown or pink tips to the
branchleis. This colouration occurs also in the
Fijian reefs. On patch reefs in deep lagoonal
situations, the colony can assume a sturdy rounded
shape with central attachment.
Laboratory Diagnosis
Branching pattern; From a side attachment,
branching is horizontal or oblique, with two to
several mam branches. Vertical to acute
branchleis occur at regular intervals along the
main branches; these usually branch again, one to
several limes, and are shorter and narrower on the
under surface. Upper surface branchleis are 5 to
9 mm wide.
Axial corallites: Outer diameter 3 0 to 4 0 mm,
up to 7 mm in lagoonal specimens; inner diameter
10 to 2-0 mm. Septation: 12 septa or slightly less,
usually well developed (primaries to 3/4R,
secondaries to 1/2R)-
Radial corallites: All of similar size, neatly and
evenly arranged around the branchlet, appressed
tubular. (Brook described them as ‘swallow-nest
shaped’). In shallow water specimens the wall is
thick, and corallites are densely packed around the
branchlet. With increasing depth, radial corallites
become more scattered, thinner walled, and the
wall may flare a little, and appear lip like.
Coenosleuni: Spines which may be laterally
flattened or slightly elaborated are arranged
evenly both on radials and between: sometimes
radial walls are costate.
Within Reef Distribution
Reef slope from crest to limits of depth of
Acropora distribution (species no. 18 in Wallace
1975); deeper water diverse reef flats and lagoonal
patch reefs; sporadically in shallow outer reef
situations where there is space for outward
growth; fringing reefs. The species is never very
306
MEMOIRS OF THE QUEENSLAND MUSEUM
abundant, but is usually present in most reef
habitats.
Identification Difficulties and History
In the field, lower reef slope specimens of this
species and of A.Jlohda may appear similar. The
two-toned colouration, thicker branchiets and
smoother general appearance of A. sarmentosa
can distinguish it. Vaughan (1918) suggested that
A. sarmentosa may be a growth form of A.
squamosa (= A. milleporaj. His specimens were
not A. sarmentosa but were indeed ‘A. squamosa’.
His misidentificalion of this species may account
for it being overlooked in much of the literature.
Both Brook and Nemenzo describe their
specimens well.
Geographic Distribution
Indo-Pacific: Philippines, Great Barrier Reef,
Fiji Islands.
Acropora florida (Dana, 1846)
(Plates 90, 91, 92)
Madrepora florida Dana, 1846, p.466, pi. 37, fig.],
non Madrepora florida: Brook, 1893, p.53.
Madrepora gra\>ida Dana. 1846, p.470. Brook, 1893,
p.59 (synonymy).
Acropora gravida: Neinen/o, 1967, p.l07, pi. 31, fig. 3;
Scheer and Pillai, 1974, p.l8, pl.5, fig.l; Pillai and
Scheer, 1974, p.453.
Madrepora mirabilis Quelch, 1 886, p. 1 59, pi. 1 0, fig. 5;
Brook, 1893, p.i25.
Madrepora cornpressa Bassett-Srnith, 1890, p.452;
Brook, 1893, p.60, pi. 33, fig.F.
Madrepora affinis Brook, 1893, p.60, pi. 28, fig.F
(synonymy).
Acropora affinis: Crossland, 1952, p.205, pi. 34, fig.l;
Nemenzo, 1967, p.77.
Acropora vermiculata Nemenzo, 1967, p 108 pi 31
fig.4. ■ ’
Material Examined
USNM: Fiji Islands, U.S. Expl. Exped., A. florida
holotype 282.
YPM: Fiji Islands. U.S. Expl. Exped., A. florida
fragment of type 2002.
BM: Tizard Bank, China Sea, 5 fthms, A. cornpressa
holotype 1889.9.24.117; Darnley Island, J.B. Jukes, A.
affinis syntype 1846.7.30.29; Banda, Challenger, A.
mirabilis holotype 1885.2.1.14; Claremont Is., G.B.R.,
Savilie Kent, A. ornata var. (id. Brook)
1892.6.8.112.
QM: Big Broadhurst Reef: S W side, reef slope:
15-3ni, i7.x.l973, C.W., G8649; lL3m, 17.x, 1973,
C.W., G8653; 9 3m, I6.X.I973, C.W., G8647; 9-3m,
17.x. 1973, C.W., G8646; 8-6m, 17.X.1973, C.W., G8b48,
G8654; 8-6m. i6.x.l973. C W,, 08661; 8ni, 28.iii.l973,
C.W., G8664; 7-9m, I5.X.I973, C.W„ 08650; 7-4m,’
15.x. 1973, C W . G8656, G8662; 6 6m, 14 x. 1973, C W.,
(18655, G8657, G8667; 6m, 27.iii 1973, C W.. G8665; '
6m, 25.iii.I973, C W., G8666; 5-8m. )3.x.I973, C.W..' i
G865I; 5-2m, I3.X.I973. C.W., G8645; 3m. 25,iii.I973’ 1
C W., G8669; 2*3m. 1 Lx. 1975, (' W., G8644; 2m !
25.iil.1973, C.W., G866I, 08663, G8668, G8670; S W. ■
side, surge channel, 8-5m, 20.x 1973, G8658, G8659; S !
W. side, outer reef fiat, 18.X.I973, G8652. ’ ^
Low Isles, June 1974, C. Limpus, G8672.
Murray Islands (Maer Is.), 18.vii.l974, G. Ingram
G867L
Bowden Reef, S. end, reef slope, 3-3m 15vii 1972
C.W., G6726.
Darley Reef, lagoonal patch reef, 3-5m, 19.vii 1972
C.W.. G6725.
Viper Reef, lagoonal patch reef, l-5m, 16viil972
C.W G6724.
Fiji Islands, Makaluvau Reef, outer reef flat
10.1,1974, C.W., G10273.
Field Diagnosis
This is cl sturdy open arborescent species, in
which the surface of the branches is covered by
short secondary branchiets. It is highly variable
in two aspects, ihe shape of the colony and the
density and prominence of the secondary ;
branchiets. The colony shape changes with depth I
and slope of the attaching surface. In shallow '
water and flat substrate it forms a rounded, open j
colony with central attachment; on sloping j
surfaces the attachment is more lateral, and the I
branches lend to extend horizontally. With !
increasing depth the branches become flatter. The
secondary branchiets vary from evenly distributed
with even lengths to scattered and variously sized
and may even be undeveloped on some branches.
The most irregular colonies occur in fringing reef
situations.
Laboratory Diagnosis
Branching pattern: Sturdy main branches divide
sparsely to form an upright bush. The shape of
the colony and the cross-sectional shape of the
branch become Hatter with increasing water
depth. The branches usually proliferate towards
the branch lips into shorter branches. The surface
of the branches is covered with short branchiets.
Except on completely vertical branches the
branchiets arc shorter on the under-surface (to
absent on horizontal branches).
Axial corallites: Outer diameter 2-0 to 3 0 mm;
inner diameter 0 8 to 1-4 mm Septation; primary
septa present, up lo 2/3R; secondary septa usually
all developed, or at least 3 present, up to
1/2R.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
307
Radial corallites: Evenly sized and distributed,
I appresscd tubular with round opening, with wall
which may flare slightly. Septation: primary septa
present, up to 1 /2R, some to all secondaries
present, up to 1 /4R.
Coenosteum; Costatc or reticulate on radial
I corallites, reticulate in between, with scattered
j simple spines.
Within Reef Distribution
' Entire reef slope from reef crest to limits of
Acropora growths; deep water lagoonal patch
reefs, fringing reefs. Occasionally on middle or
. outer reef flat. It is well known as an early
I recolonizer on the fringing reefs near Townsville,
N.Q. (D. Tarca, pers. comm.).
' Identification Difficulties and History
; The variability of the species is expressed in the
I synonymy. Brook misunderstood A. florida: his
specimens in the BM are possibly A. rotumana.
I A. florida s.s. is atypical of the species as it
appears on the Great Barrier Reef in that the
corallites are larger and the branchlets poorly
developed: G6725 is closest to this. The species
is best described by A. affinis, which has evenly
I sized and distributed branchlets; its similarity to
A. gravida was apparent to Brook. A. cotnpressa
is named from a flat plate specimen of this species.
A. mirabilis is named for a single aberrant
specimen which is probably this species under
some environmental stress.
Geographic Distribution
Indo-Pacific: Singapore, Celebes, Philippines,
Strait of Malacca, Nicobar Islands, Great Barrier
Reef, Fiji Islands, Enewetak Atoll (C.W.).
The ‘Acropora echinata’ Group
I
t The species Acropora echinata, A. suhglabra.
! A. carduus and A. longicyathus, with their
^ synonyms, have in common their shape, radial
corallite structure and coenosteal structure.
The growth forms are comparable within the
group, and nothing exactly similar is found outside
the group. The branches are commonly labelled
"bottlebrushh secondary branchlets, or bundles of
branchlets, are given off evenly around the main
branches, giving a round brush like unit. The
growth of a colony is indeterminate, the units
being capable of openly branching or closely
proliferating, giving a variety of form in collected
specimens. The colonies tend to occur on rubble
or sandy floors, and the bases of branch units are
usually dead. If the colonies occur in deep sloping
conditions, they are small and approach a
flattened form.
The radial corallites are round tubular
appressed or partly appressed, tending to develop
into axial corallites. Their numbers relative to the
number of axials are lower than in most other
groups. In all species, the number of radial
corallites per axial decreases distally on the
branch unit, so that long naked axials may occur
around the base.
The spines of the coenosteum have multiple tips,
and the coenosteal appearance on and between
radials is similar.
The main differences among the species are in
the corallite dimensions and the amount of
elaboration of the coenosteal spines.
These species occur only in the sheltered, deeper
parts of reefs, where almost any Acropora species
can survive. They appear to have poor differentia-
tion of radial corallites from axials, and if this is
accompanied by a similar lack of functional
differentiation it may account for their lack of
colonizing success.
■Acropora echinata (Dana, 1846)
Madrepora echinata Dana, 1846, p.464, pi. 36, fig.l,
la; Brook, 1893, p.I85 (synonymy).
Acropora echinata: Vaughan, 1907, p.l58, pis. 49, 50;
Wells, 1954, p.423, pi. 135, figs. 1-4, pi. 136, figs
1-6 (synonymy); Nemenzo, 1967, p.l26; Pillai and
Scheer, 1976, p.33, pl.ll, figs.l, 2.
Material Examined
USNM: Fiji Islands, U.S. Expi. Exp., A. echinata
syntype 275.
UP: Batangas Channel, Puerto Galera, Oriental
Mindoro C1064 (mentioned Nemenzo, 1967).
I have not seen this species in the central and
southern Great Barrier Reef. A single specimen
in James Cook University was collected at Lizard
Island on a sandy bottom. A large specimen in
the Queensland Museum bearing no locality data
has been presumed to be from the Great Barrier
Reef. The species is well described and illustrated
by other authors. Further notes are given with A.
subglabra.
Geographic Distribution
Philippines, Suiu Sea, Great Barrier Reef, Fiji
Islands, Samoa, Marshall Islands.
Acropora subglabra (Brook, 1891)
(Plate 94A, B)
Madrepora subglabra Brook, 1891, p.470; 1893,
p.l86, pi. 29, fig.c (synonymy).
308
MEMOIRS OF THE QUEENSLAND MUSEUM
Acropora suhglahra: Thiel. 1933. p.24.
M. subglahra var. rugixsa Brook, 1893, p.l87.
Acropora suhglahra var. rugosa: Nemcnzo, 1967,
p.l25, pi. 35, fig.2.
Madrepora procumbens Brook, 1893, p. 188, pi. 29,
fig.d.
Acropora procumbens; Th\c\, 1932, p.l30, pi. 10, fig.2;
Ncmenzo, 1967. p.I27-128, pi. 35, fig.l.
Material Examined
BM: A. procumbens syntype 1843.3.6.131; South
Seas, A. subglabra syntype 1841.12.11.1.
UP: A. subglabra var. rugosa C731 (mentioned
Nemenzo, 1967).
QM: Lizard Island fringing reef, June 1973, R.
Pearson. G 1 07 13, G 107 1 4.
Field Diagnosis
Sprawling, shrubby bottlebrush colonies, of very
slender proportions, colour pale brown or pinkish
brown.
Laboratory Diagnosis
Branching pattern: Main branches may have
any orientation from vertical to horizontal.
Secondary branchlets distributed evenly around
main branches, up to 30 mm long, undersurface
branchlets being shorter than upper surface
branchlets in horizontal branches.
Axial corallites: Outer diameter 0-9 to 1-5 mm;
inner diameter 0-5 to 08 mm. Septation:
primaries complete, to 3/4 R, secondaries absent
or some present to less than I /4R.
Radial corallites; Scattered appressed tubular,
tubo-nariform or nariform, up to 3 mm long.
Non-appressed tubular radials are incipient
axials.
Coenosteum: Lines of elaborate spines both on
and between radial corallites.
Within Reef Distribution
The species apparently does not occur in the
Central and Southern Great Barrier Reef
province, but is present further north, in situations
below the reef flat.
Identification Difficulties and History
This species requires field study. Brook’s
description of A. procumbens was on the basis of
‘contracted (axial corallite) apertures’, but in fact
the internal axial diameter of the type is only an
average 0-1 mm less than that of the A. subglabra
type. A. echinata (Dana) is a similar species of
slightly larger dimensions, and it is probable that
this is a (senior) synonym.
Geographic Distribution
Indo-Pacific: Singapore, Philippines, Banda,
Great Barrier Reef.
Acropora carduus (Dana 1846)
(Plates 93A, 94C, D)
Madrepora carduus Dana, 1846, p.464, pi. 36, fig. 2;
Brook, 1893, p.I78 (synonymy).
Acropora carduus: Faustino, 1927, p.277, pi. 93,
figs.l, 2; Nemenzo 1967, p. 123-4, pi. 34, fig. 3.
Material Examined
USNM; Sooloo Sea, U.S. Expl. Exped., Acropora
carduus paratype? Ill (N.B. Rathbun (1887) records
278 from Fiji as type).
QM: Big Broadhurst Reef, S W. slope: 6 m,
25.111.1973, C.W., G10729. G10730; 6-3 m. 23.X.1973,
C.W., GI0738; 7-8 m, I4.X.1973, C.W., G10733; 8 m,
26.111.1973, C W, G10731; 8 m, 14.X.I973, C.W.,
GI0734; 9-6 m, 15.x. 1973, C.W., G10736; 9-8 m,
16.x. 1973, C.W.. GI0737; 12-4 m, 23.x. 1973, C.W.,
GI0732; 16-2 m, 23.X.I973, C.W., G10735.
Bowden Reef. SW. side, reef slope. 2 m, 26.vii.i972,
R. Pearson, G10741. G10742.
Bushy-Redbill Reef: NW. side, sandy floor around
patch reefs. 15 m, 15.vi.l975, C.W., G 10739; W. side,
reef crest, 2 m, 3.vi-l975, C.W . G 10740.
Fiji Islands, Great Astrolabe Reefs, Jan. 1974, C.W
G 10744.
Field Diagnosis
Arborescent to shrubby, bottlebrush branched
colonies of dimensions intermediate between those
of A. subglabra and A. longicyathus. Colour;
cream, pale brown, or pink-brown.
Laboratory Diagnosis
Branching pattern: Growth indeterminate, main
branches covered by short branchlets, which are
evenly distributed, more or less equal in size and
extending at 45° to 90° from the branch. Total
diameter of this ‘bottlebrush' is 15 to 30 mm,
with branchlets up to 12 mm long and 5 mm wide.
Main branches may be proliferous, giving a
shrubby appearance, or sparsely branching, giving
an arborescent appearance: usually both types of
branching occur within a colony. In deep water
(e.g. specimen G 10735) the branchlets develop on
the upper surface of the colony only, and the
colony is effectively reduced to a small plate.
Axial corallites: From non exert (budding at
tip) to 7 mm exert. Outer diameter 1-0 to 2 0 mm;
inner diameter 0-5 to 0-8 mm. Septation: primary
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
309
septa well developed, up to 3/4R, secondary septa
absent, or some present, to 1/4R.
Radial coralliles; On branchlets, scattered
appressed tubular, sometimes approaching nar-
iform, with round openings. On some colonies
radials crowded, partly appressed tubular and
extending out from branchlets. On the main
branches radials are sub-immersed to immersed,
or in some cases tubular appressed. Primary septa
poorly to well developed to I/2R, secondaries
usually absent except in immersed corallites,
where a few may be present. Radial corallites are
best represented on distal parts of branches. At
the bases of branches most have developed into
long tubular axials.
Coenosteum: finely echinulate on radial
corallites and between. Occasionally spines are
arranged in rows or even joined as costae.
Within Reef Distribution
Reef slope, from below the area of rough water
surge to about 20 m; deep water reef flat areas,
sandy floors of lagoons and reef-edge patch
reefs.
Identification Difficulties and History
Being a deeper water species, A. carduus is
poorly represented in older collections, and hence
presents few historical problems. > It occurs with
A. longicyathus. and they can be easily separated
by obvious differences in corallite dimensions.
Geographic Distribution
Indo-Pacific: Philippines, Sulu Sea, Great
Barrier Reef, Fiji Islands.
Acropora longicyathus (Milne Edwards and
Haime, 1860)
(Plates 93B, 95)
Madrepora longicyathus Milne Edwards and Haime,
1860, p.l48: Brook. 1893, p.l87 (Synonymy).
Acropora longicyathus: Nemenzo, 1967, p.l28, pi. 35,
figs. 3, 4.
Madrepora prolixa Verrill, 1866, p.22.
Acropora prolixa: Verrill, 1902, p.237, pi. 36, figs. 3,
3a, pl.36A, Figs. 3, 3a; pl.36F, fig.l4; Hoffmeister,
1925, p.65, pi. 16; Crossland, 1952, p.226.
Material Examined:
PM: A. longicyathus type 303A.
USNM: Ousima, A. prolixa syntype 414.
QM; Big Broadhursl Reef, S W. side, reef slope: 6m,
25.iii.1973, C.W., G10763; 7m, 27.iii.l973, C.W.,
G10767; 8m, 26.iii.1973, C.W., G10762, G10764; 9m,
15.X.1973, C.W., G10751, G10752; 9-6m, 15.x. 1973,
C.W., GI0753; 9 7m, 15.X.1973, C.W., G10754; 9-8m,
16.x. 1973, C.W., G10755; 10-3m, 16.X.1973, C.W.,
G10756; I0-6m, 16.X.1973, C.W.. G10757, G10758;
12m, 16.x. 1973, C.W.. G10759; 12-5m, 17.X.1973,
C W.. G10760; surge channel, 10m, 20.X.1973, C.W.,
GI076L
Bushy-Redbil Reef: W. side, outer flat, 31.V.1975,
C.W.,G]0745; reef slope near Redbill Is., 3m, l.vi.l975,
C.W., G10746; W. side, patch reefs (sandy floor), 8m,
3.vi.l975, C.W., G10747, G10748; N W. side, patch
reefs (sandy floor) I2m, 14. vi. 1975, C.W., G10749,
G10750.
Field Diagnosis
Arborescent to shrubby, bottlebrush branched
colonies of sturdy dimensions. Colour: cream, pale
to dark brown, or blue-brown.
Laboratory Diagnosis
Branching pattern: Growth indeterminate, main
branches covered by evenly distributed proliferous
branchlets or bundles of branchlets. These are
directed at 45° to 90° to the branch, and may be
as short as 5 mm all over branch, or all several
centimetres long and much branched. Greatest
total branch width is 80 mm, smallest (except for
branch tips) 40 mm. Deep-water colonies can
have little undersurface development, and
approach a plate like shape.
Axial corallites: Sometimes exert (to 10 mm)
near base of branch, more often one side naked
of radial corallites, other side with 3 or 4 from
their outer rim. Outer diameter 2-1 to 2*8 mm;
inner diameter 0-8 to 13 mm. Septation; primary
cycle present to 3/4R, secondary cycle present, or
at least partly developed up to 1 /4R.
Radial corallites: On branchlets, appressed or
partly appressed tubular with round openings.
When fully appressed, radials are scattered, only
a few to each axial corallite: however, branchlets
can have radials touching. In the first case, radials
on main branches are immersed or (more usually)
sub-immersed; in the second, main branch radials
are usually similar to those of branchlets.
Coenosteum: Neatly echinulate on and between
corallites, the spines laterally flattened with
simple to forked or more elaborate lips.
Within Reef Distribution
Reef slope, from below the area of rough water
surge to about 20m; deep water reef flat areas,
sandy floors of lagoons and leeward patch
reefs.
Identification Difficulties and History
The two species A. longicyathus and A. prolixa
have been combined by other authors: Verrill’s
310
MEMOIRS OF THE QUEENSLAND MUSEUM
specimens were small branch tips. This species is
not well represented in collections.
Geographic Distribution
Philippines, Ousima, Samoa, New Guinea,
Great Barrier Reef.
The ‘Acropora .squarrosa’ Group
The group of species A. elseyi. A. rosaria, A.
squarrosa and A. granulosa has in common (1)
dense cchinulate cocnosleum of elaborate spines,
(2) large tubular to nariform radial corallites, and
(3) a tendency for some branchlcts to have an
upper surface naked of radial corallites. The group
is linked to the \4. echinata' group which has,
however, less differentiated radial corallites.
The synonyms adopted here may be controver-
sial. Most of the species involved seem to be well
documented in the literature, but they still present
extraordinary identification problems. The
synonymies are presented as hypothe.ses requiring
further testing.
The oldest name for the group, A. squarrosa
(Ehrenberg) has been treated by a number of
authors (c.g. von Marenzeller 1907, Vaughan
1918, Wells 1954), but the interpretations are
various and sometimes confusing. A. granulosa.
with its synonyms here listed, links this group to
the ‘A. echinata' group.
A. nncrophthalma, as here interpreted, is
included in the group because of its cocnosteal
structure.
Acropora microphthalma (Verrill, 1869)
(Plate 96)
Madrepora microphthalma Verrill, 1869, pp.83, and
102 .
Acropora microphthalma: Verrill, 1902, p.232,
pl.36C, fig.l, 36E, fig. 1 5.
Acropora microphthalma: Wells, 1954. p.429,
pi. 126, figs. 7 9 (synonymy),
non Acropora microphthalma: Stephenson and Wells,
1956, p.lO.
Acropora laevis (part) Crossland, 1952, p.230.
Mate-rial Examini-d
YPM: Ryuku Islands, A. microphthalma holotype
774 (fragment).
QM: Big Broadhurst Reef, SW. side, reef slope; 3m,
28.111.1973, C.W.. G8684; 4m, 13.x. 1973, C.W., G8688,
G8689; 6m. 25.iii.1973, C.W., G8685; 6-4m, 14.x. 1973,
C.W„ G8693; 7-8m, 15.X.1973, C.W., G8691; 8m.
26.111.1973, C.W., G8683, G8686; 8m, 28.iii.1973, C.W.,
G8682; 91m, 16.X.1973, C.W., G8690; 9-4m, 16.X.1973,
C.W., G8692.
Bowden Reef, SW. side, reef crest, 0-3m, 24.vii.I972,
C.W., G8680.
Field Diagnosis
An aborcsccnt species forming small clumps (up
to 100 cm across) the branches having slender
dimensions, and radial corallites being small and
crowded. Colour is most commonly a whitish-
cream. Although because of its size it is not a
conspicuous species, it is the most delicate
arborescent coral in the Great Barrier Reef area,
and is easily recognized.
Laboratory Diagnosis
Branching pattern: Branching is open arbores-
cent, with many short branchlcts being given off
towards branch tips at 45*^ to 90° to the main
branches. The greatest branch width measured in
the collections is 14 mm, the smallest (excluding
branchlcts) 5 mm.
Axial corallites: Outer diameter 1-8 to 2-3 mm;
inner diameter 0-8 to 1 00 mm. Septation:
primary septa strongly developed, up to 3/4R,
secondaries absent, to prc.scnl up to 1/4R.
Radial corallites: Extend at about 45°, short
tubular with round to oval opening to tubo-
nariform, outer wall slightly thickened. First
septal cycle well developed, up to 2/3R, second
cycle usually partially represented. Radials arc
uniform in size and evenly distributed, giving a
fine-grained appearance to the branches.
Cocnosleum: When spines are well developed,
they have elaborate tips and are distributed both
on corallites and between. However, lightly
calcified coralla may have simple pointed spines
and a generally spongy appearance.
Within Reef Distribution
Middle reef flat, deeper reef flat areas, sandy
floors around patch reefs, reef slope to surge
channel floor.
Identification Difficulties and History
Interpretation of A. microphthalma from
VerrilTs poor series is difficult and previous
identifications have linked it with A. exilis. The
A. microphthalma of Stephenson and Wells
(1956) (no. G2697, QM) is in my opinion A.
nasuta.
The species as here interpreted is distinctive,
particularly in the field, and the fragment I have
seen of VerrilTs type compares well with the tips
of the larger dimensioned specimens. At least one
specimen of Crossland's A. laevis series (BM
1934.5.14.67) is this species.
Geographic Distribution
Ryuku Islands, Great Barrier Reef, Fiji Islands
(C.W.), Enewetak Atoll (C.W.).
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
311
Acropora elseyi (Brook, 1892)
(Plate 97 A, B.)
Madrepora elseyi Brook, 1892, p.456; 1893, p. 172,
pl.ll, figs. E, F.
Acropora elseyi: Crossland, 1952, p.223; Pillai and
Scheer, 1976, p.31, pL9, fig.l.
Madrepora exilis Brook, 1892, p.457; 1893, p.l72,
pi. 10, figs. C, D.
Acropora exilis: Crossland, 1952 (part), p.223, pi. 39,
fig. 4; Stephenson and Wells, 1956, p.l3, pi. 1(b)
(synonymy).
Material Examined
BM: N. Australia, J. Elsey, A. elseyi syntypes
1857.11.18.214, 215, 216, 217, 218; Thursday Is.,
Saville-Kcnt, A. elseyi 1892.6.8.241-4, 6; Rocky Is.
Saville-Kcnt, A, elseyi 1892.6.8.247 (mentioned Brook,
1893). Port Denison, Savillc-Kent, A. exilis syntypes,
1892.6.8.103, 104, 106.
QM: Big Broadhurst Reef, E. side, side of reef flat
channel, 2Lx.i973, C.W., G10802.
Bushy-Redbill Reef: W. side, middle reef flat, Im,
27.V.1975, C.W., GI0793; G10794; W. side, outer reef
flat, 8.vi.l975, C.W., G10797; N W. side, first reef crest,
23.xii.1972, C.W., GI0798; adjacent Redbill Is., reef
crest, 19.xii.l972, C.W., G 10795; adjacent Redbill Is.,
reef slope, 19.xii.l972, C.W., G10796.
Field Diagnosis
Low bushy to caespitose colonies with
bottlebrush-lype branches. Colour usually bright
yellow, yellow-brown or lime green-brown.
Laboratory Diagnosis
Branching pattern: Main branches surrounded
by regularly placed short branchlets which may
be of equal or unequal length. Main branches up
to 15 mm wide, branching units up to 100 mm
wide, branchlets up to 8 mm wide.
Axial coralliles: From non-exerl to 2 mm exert.
Outer diameter 1-6 to 3-2 mm; inner dimaeter 0*6
to 10 mm. Septation: primary septa well
developed, up to 3/4R, secondary septa absent or
few present to less than 1/4R.
Radial corallites: On both main branches and
branchlets, radials are tubular with round
openings, becoming round tubular distally.
Radials are usually evenly distributed on
branches, and almost touching, sometimes upper
surface of small branchlets is naked of corallites.
Septation: primaries well developed, up to 1/2R,
secondaries absent or a few just visible.
Coenosteum: Neatly and densely costate both
on and between corallites.
Within Reef Distribution
This is one of the few Acropora occurring in
the shallow longitudinal patch reefs perpendicular
to the reef edge, forming the windward edge of
lagoons, and is characteristic of these areas. It also
occurs in deeper reef flat areas, and reef crest and
upper reef slope on some leeward reef areas. It
is a common species of fringing reefs of
continental islands.
Identification Difficulties and History
Both species in synonymy were described from
the Barrier Reef, A. elseyi being slightly sturdier
than ‘A. exilis’. The species will be better
understood when studied on fringing reefs.
Geographic Distribution
Indo-Pacific: Maidive Archipelago, Great
Barrier Reef.
Acropora rosaria (Dana 1846)
(Plate 97 C, D)
Madrepora rosaria Dana. 1846, p.465, pi. 36, fig. 3.
Madrepora rosaria var. diffusa Brook, 1893,
p. 1 80.
Acropora rosaria: Vaughan, 1918, p.l84, pi. 82, figs. 2,
2a, 2b; Weils, 1954, p.428, pi. 130, figs. 3, 4;
Stephenson and Wells, 1956, p.l8.
Acropora rosaria (part): Crossland, 1952, p.224,
pi. 40, figs.l, 4.
Madrepora syringodes (part) Brook, 1892, p.463;
1893, p.l77, pi. 33, fig.E.
Material Examimt:)
USNM: Fiji islands, U.S. Expl. Exped., A. rosaria
holotype 281; paratype 933.
QM: Palm Islands, 1939, T. C. Marshall, G9192,
G9193, G9I94, G9I95, G9196, G9197, G9200,
G9202.
Field Diagnosis
Not determined.
Laboratory Diagnosis
Branching pattern: Usually ‘bottlebrush’
branching units arc formed, a central branch of
up to 20 mm diameter bearing branchlets at even
intervals all around.
Axial corallites: Exert 10 to 2 0 mm. Outer
diameter 2-5 to 3-8 mm; inner diameter: 0-8 to
2-4 mm. Septation: first cycle well developed, up
to 3/4R, second cycle usually present, or most
septa present, up to 1/3R.
Radial corallites: Tubular, partly or fully
appressed, or nariform. Walls thick, primary septa
developed up to 1/3R and secondary septa partly
developed, up to I/2R.
Coenosteum: Densely arranged spines, some-
times laterally flattened, both on and between
corallites.
312
MEMOIRS OF THE QUEENSLAND MUSEUM
Within Reef Distribution
The specimens described were collected on the
fringing reefs of the Palm Island group.
Identification Difficulties and History
This species requires further study. The only
material easily identifiable with A. rosaria
available to me was collected in habitats not
comparable with those I have studied. I am thus
unable to pursue the possibility of a close
relationship between this species and A.
squarrosa. I consider it probable that the figured
syntype of Brook's A. syringodes
(BM 1892.6.8.209 from Palm Islands) is A.
rosaria. Interpretations of A. syringodes are
difficult to follow and seem to span a number of
species, and it is likely that the latter species, as
described, is a mixture.
Geographic Distribution
Great Barrier Reef, Fiji Islands, Samoa,
Marshall Islands.
Acropora squarrosa (Ehrenberg, 1834)
(Plates 98, 99, 100)
Heteropora squarrosa Ehrenberg, 1834, p.l!2.
Madrepora squarrosa: Brook, 1893, p.65
(synonymy).
Acropora squarrosa: von Marenzeller, 1907, p.46,
pi. 1 4, figs. 36-39; Vaughan. 1918, p.l84, pi. 83, figs.
2, 2a, 2b; Wells. 1954, p.427, pi. 129, figs. 1, 2;
Rossi, 1954, 1954, p-52; Nemenzo, 1967, p.69,
pl.21, fig.4; Pillai and Scheer 1976, p.31.
Acropora murrayensis Vaughan, 1918. p. 183, pi. 82,
figs.l, la. lb; Nemenzo, 1967, p.71, pi 23, fig.2.
Madrepora syringodes (part) Brook, 1892, p.463;
Brook, 1893, p. 177 (not illus.).
Madrepora cancellata Brook, 1893, p. 166, pi. 32,
fig.C.
Acropora cancellata: Crossland, 1952, p.225, pl.41,
figs.3, 4.
Material Examinrd
USNM: Murray Islands, A. murrayensis holotypc.
BM: A. syringodes 1893.4.7.163 (mentioned Brook,
1893); Louisadc Archipelago, 15 fathoms, A. cancellata
holotype 185L9.29.39.
QM: Big Broadhurst Reef, SW. side, reef slope: 1-3
m, 23.X.1973, C.W., GI0839; 1 -9 m, 23.x.i973, C.W.,
G10840; 6-3 m, 13.X.1973, C.W., GI0826, G10827; 6-3
m. 23.X-1973. C.W.. G1084I; 7 m, 13.x. 1973, C.W.,
G10825; 7-8 ni. I4.X.I973. C.W.. G10828, G10830; 8
m. 14.X.I973, C.W'., G10829; 81 m, 14.X.1973, C.W..
GI0824; 8-2 m, 14.X.1973, C.W., G10831; 8-7 m,
15.X.1973, C.W., G10832; 9 m, I5.X.1973, C.W.,
G10833, G10834; 9-2 m, 25.X.1973, C.W., G10842; 10-6
m, 16.X.I973, C.W., G10836; 11-4 m, 25.X.1973, C.W.,
G10843: 12*9 m, 23.X.1973, C.W., G10845; 13-4 m,
17. X.1976, C.W., G10837; 30 m, I8.x.i973, C.W.,
G10838; E. side, reef slope, 4 m, 2Lx.l973, C.W..
G 1 0844.
Bushy-Rcdbill Reef: Reef crest adjacent Redbill Is.:
18. xii.l972, C.W., G10768, G10770; 19.xii.l972, C.W.,
G10769; Lvi.I975, C.W., G10771; reef slope adjacent
Redbill Is.: 20.xii.l972, E. Lovell, G10772, G10773:
20.xii.I972, C.W., G9I89; W. side, middle reef flat; 1
m, 8.vi.l975. C.W., GI0774, G10775; I4.vi.l975, C.W.,
G10776; NW side, first reef crest: 22.xii.I972, E.
Lovell. G10777; 22.xii.1972, C.W., GI0778, G10779;
NE. side, D. Hadley. I. i. 1973, G10780.
Bowden Reef, SW\ side, reef slope: 24.vii.1972, C.W,,
GI0789; 26.vii.1972. C.W., GI0788.
Darley Reef, patch reef in lagoon: Im, I8.vii.l972,
R Pearson, GI0782, GI0785; 1 m, 22.iii.I973, C.W.,
G10783, GI0784, G10786, GI0787. G10790; 3 m,
I8vii.l972, C.W.. GI0846; 3 m, 24.iii.1973, C.W.,
G10791; 4 m, 22.iii.1973, C.W., G10792; 6 m,
22.iii.1973, C W.. G10847; 7 m, 22.iii. 1973, C.W.,
G 10846.
Viper Reef, patch reef in lagoon, 2 m, 16.vii.l972,
C.W., G1078L
Field Diagnosis
Colonies can be bottlebrush branching,
caespitose to plate like, some of the variation being
due to depth gradients. The surface of the
branches has a general ‘globular’ appearance due
to rounded, thickened corallites, and a whitish
glow behind the general colouration due to dense
coenosteum showing through semi-transparent
tissue. Common colours are pale blue, lavendar,
cream-brown and yellow-brown.
Laboratory Diagnosis
Branching pattern: Bottlebrush branching
colonies have tapering branching units, and the
branchlets extend widely from the main branch
(90° or just less). In caespitose colonies there is
a tendency tow'ards development of a bottlebrush
unit (incipient branchlets developed evenly around
the branches) and even in plate-like colonies the
short upright branches bear indications of
incipient branchlets along their length. In most
specimens of this species branchlets which are
naked of radial corallites on their upper surface
can be seen.
Axial corallites; From barely exert to 2*5 mm
exert. Outer diameter 2-6 to 3-8 mm; inner
diameter 0-7 to 1-4 mm. Septation: primaries
usually well developed (up to 3/4R), secondaries
completely absent to fully present, up to
1/3R.
Radial corallites: Tubular appressed to
nariform, with round to oval openings at 90° or
less to branch. Wall is thickened so that corallite
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
313
may appear round tubular, or distorted rostrate
hooks or horns may be developed. Septa are
usually poorly developed; primaries can usually be
just detected, and sometimes a few second-
aries.
Coenosteum: Laterally flattened spines with
elaborated tips are arranged densely and evenly
over both the corallites and the inter-corallite
region.
Identification Difficulties and History
The material examined shows extraordinary
variation in colony shape, branch size, and radial
corallile distribution. However, I am unable to
separate the collection into units on any of these
characters. The evidence suggests a species where
the role of the radial corallite is very flexible. The
‘easiest’ identification in this group is A.
murrayensis s.s,, which was well described by
Vaughan. This occurs particularly on sandy floor
situations in lagoons and deep water reef flat
areas. On the reef crest and upper reef slope it
can also occur, but with shortened ‘botllebrush’
branches. The caespitose to plate-like forms occur
mainly on sloping surfaces, and on deep parts of
the reef slope small flat plates approaching A.
granulosa are formed, these being identifiable
with A. cancellata s.s. At least one of Brook’s
syntypes of A. syringodes (BM 1893.4.7.163)
from unknown locality, compares closely with
caespitose specimens in the present collection. The
biggest problem is in interpreting A. squarrosa s.s,
which I have taken (following Vaughan 1918) to
be caespitose, approaching corymbose, with very
regularly arranged radial corallites and relatively
thick branches. I suspect A. austera specimens are
sometimes identified to this species.
Geographic Distribution
Indo-Pacific; Red Sea, Seychelles, Maldives,
Minicoy, Philippines, Louisade Archipelago,
Great Barrier Reef, Marshall Islands.
Acropora granulosa (Milne Edwards and Haime,
1860)
(Plates 101, 102)
Madrepora granulosa Milne Edwards and Haime,
1860, p.l56; Brook, 1893, p.l89 (synonymy).
Madrepora speciosa Quelch, 1886, p.l63, pi. 10, fig.l;
Brook, 1893. p.l91.
Madrepora clavigera Brook, 1892, p.455; 1893, p. 183,
pi. 9, figs. A, A'.
Acropora clavigera: Crossland, 1952, p.226, pi. 40,,
fig. 2, pl.42, fig.3.
Madrepora rayneri Brook, 1892, p.461: 1893, p.l91,
pi. 8, fig.A.
Acropora rayneri: Wells, 1954, p.431, pi. 134, fig. 6,
pi. 137, figs. I, 2, pi. 139, figs.l, 2.
Material Examined
PM: Acropora granulosa holotype 328a.
BM: Acropora clavigera holotype 1851.1 1.14.28. Fiji,
Acropora rayneri syntypes, 1862.2.4.30, 1862.2.4.44.
QM; Big Broadhurst Reef, SW. slope: Oct., 1973,
C.W., G1I483; 5 m, 27.iii.1973, C.W., G11476; 7 m,
26.iii.l973, C.W.. 011474; 7 m, 27.iii.1973, C.W.,
GI 1477, G1 1493; 8 m, 28.iii.1973, C.W., G1 1492; 8 m,
24.X.1973, C.W., G1I481; 10 m, 23.x. 1973, C.W.,
GI1494; 10-4 m, 16.X.1973. C.W., G11487; 17-2 m,
17.X.1973, C.W., GH480; 25 m, 17.x. 1973, C.W.,
011485.
Bushy-Rcdbill Reef: adjacent Redbill Is., reef crest,
I m, Lvi.1975, C.W., G11490; 20.xii.l972, E. Lovell,
Gl 1491; NW. side, patch reefs, I5.vi.l975, 12 m, C.W.,
G11488.
Darley Reef, patch reef in lagoon, 2-5 m, 19.vii.l972,
C.W., G6723.
Feather Reef, reef slope, 10 m, 24.x. 1972, R. Pearson,
G11479, G11482.
Field Diagnosis
Colonies are side-attached thin plates, with
anastomosing horizontal branches and short
vertical branchlets or groups of branchlets. Axial
corallites are long, tapering or rounded, and may
be sinuous. Radials are relatively few and
.scattered.
Laboratory Diagnosis
Branching pattern; described above.
Axial corallites: From barely exert to as long
as 10 mm bare of radials. Outer diameter 1-2 to
2-5 mm; inner diameter: 0-6 to 2-2 mm. Septation;
primaries well developed, up to 3/4R, secondaries
absent or poorly represented, to less than
1/4R.
Radial corallites: Appressed tubular to nariform
with round to slightly oval opening, directed at
90^ or less to branch. Radials, except at growing
edge of colony, are few, and inconspicuous, the
appearance of the colony being dominated by the
axial corallites.
Coenosteum; Spines with pointed or laterally
flattened tips are densely and evenly arranged on
and between corallites.
Within Reef Distribution
Reef slope and sides of lagoonal patch reefs,
usually at depths greater than 20 m, but can be
much shallower on very steeply sloping surfaces,
for example G 11490 is a specimen from 1 m on
the side of a sleep and narrow surge channel
opening.
314
MEMOIRS OF THE QUEENSLAND MUSEUM
Identification Difficulties and History
The conibinalion of the various species in
synonymy seems barely possible from Ihe type
material, and also from much of my own material
which identifies strongly with one or another
species. A more acceptable combination would be
two species, A. speciosa = A. rayneri and A.
granulosa = A. clavigera. However some large
specimens show a mixture of characteristics of all
four species. The differences amongst the
described species are in dimensions and the degree
of tapering of the axial corallilc; ‘A. rayneri' is
very slight, and A. speciosa' sturdier, both with
tapering axials, A. granulosa s.s. and ‘A.
clavigera' both have thick, rounded, non-tapering
axials. In all the material there is little difference
in inner axial diameter, axial septation, radial
corallite shape and coenosteal spines. The
locations of the specimens give no clue to the
morphological differences. All categories except
A. speciosa' (one lagoonal specimen) co-occur on
the reef slope. Their different appearances
(whether genotypically or phenotypically deter-
mined) cannot be explained by reef slope
gradients, and are probably related to rnicrohabi-
tat features.
Despite the variety in the species, extreme care
must be taken in assigning a specimen to this
species, as deeper water specimens of many other
species appear (as cleaned specimens) similar to
this species.
Geographic Distribution
Indo-Pacific: Mascarene Archipeligo (G. Faurc,
pers. comm.), Louisade Archipelago, Great
Barrier Reef, Fiji Islands, Marshall Islands.
Acropora austera (Dana, 1846)
(Plates 103, 65C)
Madrepora austera Dana, 1846, p.478; Brook, 1893,
p.56 (synonymy); Verrill, 1902, p.266, pi 36, fig.
10, pi. 36B, fig. 1.
Materiai. Exmaim d
YPM: Acropora austera (fragment of type) 4190.
QM: Big Broadhursl Reef, S W. side, reef slope: 7-8
m, I4.X.1973. C.W., G 108 1 3, G 10828; 8-0 m, 14.X.I973,
C.W., G10814; 8-2 m, 14.x. 1973. C.W., G10815; 9-0 m,
15.x. 1973. C.W., G10816.
Bowden Reef. SW. end. reef slope, 26.vii.I972, C.W..
G10817.
Bushy-Redbill Reef: adjacent Redbil! Is., reef crest:
2.vi.l975. C.W.. G10806, GI0807; 4.vi.l975, C.W.,
G 10808; S. side, reef slope, 3 m, 27.xii.1972, C.W.,
G10809, G10810; W, side, patch reef, 8 m, 15. vi. 1975,
C.W., G10811; N W. side, patch reef, 19.xii.l972, E.
Lovell, G 108 12.
Darley Reef, patch reef in lagoon, 24.iii.1973, C.W.,
G10818, G10819.
Viper Reef, patch reef in lagoon, 16.vii.l972, C.W.,
G 1 0820.
Fiji Islands, Great Astrolabe Reefs, .Ian. 1974, C.W.,
GI0821, G10822.
Field Dlagnosis
Patchy, irregularly branching colonies of bushy,
bottlcbrush or caespitose appearance; large,
irregular-length rounded radial corallitcs with
very large openings. Colour cream to pale brown
or dirty yellow. Extended polyps may be bright
orange (axials) and purple (radials).
Laboratory Diagnosis
Branching pattern: Main branches are up to 40
mm width. These give off secondary branches of
such irregular spacing, angle, and length, that the
colony may appear caespitose, bottlcbrush, or low
arborescent. Although growth is apparently
indeterminate, colonies do not often reach more
than about 1 m across, occurring usually in areas
of high density cover.
Axial corallites: From barely exert to 3 mm
exert. Outer diameter 2-4 to 3-8 mm; inner
diameter 1-0 to 1*5 mm. Septation: all septa
usually present, primaries up to 2/3R, secondaries
up to 1 /2R.
Radial corallites: On secondary branches
radials are tubular, nariform or tubo-nariform, of
mixed length. Similar corallites on main branches
tend to become appressed, then rounded, distally.
The shape of the opening approaches a square,
and the lower (or outer) wall is sometimes
thickened accentuating the angularity of the
opening. Primary and most or all secondary septa
are developed and, as Dana comments, these
usually slope towards the centre deep in the
corallite.
Coenostcum: Reticulate with elaborated spines
on and between radial corallites.
Within Reee Distribution
Reef crest and upper slope, top and sides of
lagoonal patch reefs. This species seems to occur
particularly where there is a bend or edge on the
reef surface (see plate 65C).
Identification Difficulties and History
This species is neglected in the literature,
although my experience has shown it to be widely
distributed.
Geographic Distribution
Indo-Pacific: Singapore, Philippines, Great
Barrier Reef, Fiji (C.W.), Enewetak Atoll
(C.W.).
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
315
Acropora brueggemanni (Brook, 1893)
(Plate 50C, D)
Madrepora brueggemanni Brook, 1893, p. 145, pi. 24.
(synonymy).
Acropora brueggemanni: Crossland, 1952, p.221.
Madrepora brueggemanni var. uncinata Brook, 1893,,
p.l46, pi. 35, fig. E.
Acropora brueggemanni var. uncinata: Nemenzo,
1967, p.55.
Material Examined
BM: Singapore, A. brueggemanni syntype
1878.4.1.1.
JCU: Palm Island Group, T. Done, 4 specimens;
Lizard Island Group, M. Pichon and T. Done, 6
specimens.
QM: Big Broadhurst Reef, SW. side, reef slope, 5 m,
27.iii.l973, C.W., G11495; 6 m, 27.iii.1973, C.W.,
G1 1496-8.
Palfrey Islet, Lizard Island group, 30.vii.l977, P.
Hutchings, G1 1499-506.
Field Diagnosis
Sturdy arborescently branching colonies occur
as either small clumps or extensive thickets. Axial
corallites are obviously large and bulbous and
there may be several axials at or near the branch
tip. General appearance approaches that of A.
palifera (see Wells 1954, p.430), but a definite
branching pattern is achieved. Colour is pale
brown to pale apple green.
Laboratory Diagnosis
Branching pattern: The branches may be round
or irregular in cross section, from 15 to 30 mm
diameter, and either tapering or truncate. The
angle of branching is usually wide.
Axial corallites: Outer diameter 2-0 to 8 0 mm;
inner diameter 0-8 to 1-4 mm. In general, when
there is a single axial corallite, this is wider than
the members of a bundle of axials. Septation: Both
septal cycles usually developed, the primaries up
to 3/4R, secondaries up to 1/3R.
Radial corallites: From short (barely emergent)
to appressed tubular, with round openings.
Primary septal cycle usually well developed, up to
1 /2R; secondary cycle poorly developed, up to
1/4R.
Coenosteum: Dense arrangement of elaborated
spines both on and between radials.
Within Reef Distribution
In my work, I have encountered this species
only as a rare member of the upper reef slope
assemblage. It is very common on the fringing
reefs of the Lizard Island and Palm Island
Groups, usually on sandy or poorly consolidated
substrates.
IDENTIFICATION DIFFICULTIES AND HISTORY
Both Brook and Crossland note an approach to
the characteristics of subgenus Isopora in this
species. Strong affinities with A. palifera can be
seen in the radial corallite structure, coenosteal
texture, and tendency to multiple axial corallites.
It is considered in the present paper in order to
avoid field-identification problems.
Geographic Distribution
Singapore, Philippines, Great Barrier Reef.
ACKNOWLEDGMENTS
I wish to express my gratitude to the following:
Prof. Jan Verwey (whose taxonomic decisions on
Acropora were made many years ago but whose
search for perfection still keeps him from
publication) for his generosity in discussing his
work, and Mrs Verwey for her hospitality. Prof.
John Wells for encouragement, constant assis-
tance and hospitality. Other workers in coral
taxonomy, in particular Maya Wijsman-Best,
Richard Randall, Brian Rosen, Michel Pichon,
Gerard Faure, John Veron, and members of the
N.S.F. Enewetak Taxonomic Workshop (1976)
for advice and discussion. Micky Watkins and
Hugh Hope who took us to the reef. Robert
Pearson whose unpublished observations con-
tributed to the ground work of the study, and
Colin Limpus who made large contributions of
specimens and observations. Museum colleagues
who helped in the field (Roily McKay, Terry
Tebblc, Dianne Gleeson, David Joffe); in separate
field collecting (Glen Ingram, Jan Buhmann) and
back-up work (Alan Easton and staff, photo-
graphy; Peter Berryman and staff, artwork); many
members of technical and clerical staff, and Bruce
Campbell and Lester Cannon for advice on the
manuscript. Peter Beveridge and family and Bruce
Carlson for hospitality in Fiji. The following
Museum Curators and their staff for assistance
with specimens and hospitality: Dr K. Reutzler
and Dr F. Bayer (USNM), Dr P. F. S. Cornelius
(BM), Dr J. P. Chevalier (PM), Dr W. Hartman
(YPM), Dr D. Kuhlmann (MNB), Dr M.
Wijsman-Best (RNHL), Dr Abad-Santos (LP),
Dr Van Soest (Zoological Museum Amsterdam).
Neville Coleman for considerable assistance with
coloured underwater records. D. and M. Tarca of
Townsville. Many other friends in the field, but
in particular Ed Lovell, Robin Elks, Harvey
Walsh and Carl Wallace. John Hardy and Bob
Grimmer of the University of Queensland
MEMOIRS OF THE QUEENSLAND MUSEUM
Electron Microscope Unit. Michel Pichon and
Terry Done for hospitality at James Cook
University.
The study was supported by grants from the
Advisory Committee on Crown-of-Thorns starfish
and Queensland Museum Funds.
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1-23.
1956. Scieractinia. pp.F328-F478 in Moore, R. C.
(ed.) ‘Treatise on Invertebrate Paleontology’ part F.
Coelenterata. (Geol. Soc. Amer).
318
MEMOIRS OF THE QUEENSLAND MUSEUM
INDEX TO SPECIES
Page
Plates
A. abrotanoides
280
47
A. aculeus
295
75, 76
A. aspera
286
48, 60a, 81
A. austera
314
65. 103
A. breuggemanni
315
50
A. carduus
308
93. 94
A. cerealis
297
77
A. clathrata
302
53. 64, 75, 86
A. cyiherea
289
44, 53, 64, 66, 67, 70
A. delicatula
292
69
A. digitifera
301
48, 65, 84
A. divaricata
303
87, 88, 93
A. diversa
298
79, 80
A. echinata
307
A. elseyi
311
97
A. florida
306
64, 90, 91, 92, 93
A. formosa
282
51, 52, 53, 64, 65, 75
A. grand is
281
50
A. granulosa
313
101, 102
A. haimei
293
53, 70, 71
A. horrida
284
55, 56
A. humilis
300
44. 65, 81, 82, 83
A. hyacinthus
288
44. 53, 64, 65, 66, 70,
A. intermedia
280
48, 49
A. longicyathus
309
93. 95
A. microphthalma
310
96
A. millepora
291
43, 68, 81, 84
A. multiacuta
301
85
A. nasuta
297
78
A. pulchra
285
58, 59, 60
A. robusta
278
44, 45, 81
A. rosaria
311
97
A. rotumana
279
46
A. sarmentosa
305
89
A. splendida
283
53, 54, 90, 93
A. squarrosa
312
98, 99, 100
A. subglabra
307
94
A. tenuis
294
43, 72, 73
A. tubicinaria
295
74
A. variabilis
299
44, 80
A. vaughani
285
57
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
319
Fig. 3: Diagrammatic representation of Plate 43C indicating features used in the description of species.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 43
Features of Acropora displayed by scanning electron microscopy.
A. Side view of a branch tip (of A. mitlepora) from which tissues
have been removed to display skeleton.
B. Side view of a branch tip (of A. tenuis) from which skeleton
has been removed, to display soft tissues. The specimen is in
a retracted state, so that tentacles are withdrawn.
C. Face view of skeletal branch tip of A. millepora.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 43
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi.ATi-; 44
Acropora robusta (Dana, 1846) on Bushy-Rcdbill Reef.
A. Large colony on SW. reef crest showing distorted humps, some
vertical cones, and some free branching. Other corals in the
picture are Pocillopora.
B. (Left hand side) colony on SW. reef crest with vertical cones
and free branching. Other Acropora in the picture are A.
hyacinthus (upper centre), A. variabilis (centre) and several A.
humilis.
C. Small colony on SE. (weather) side of reef (arrow), with A.
cuneata (centre), A. humilis (upper centre) and Pocillopora.
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATF. 45
Acropora robusta (Dana. 1846)
A, B. Free branching portion of colony, G 10191 . Big Broadhurst
reef. SW. slope, 1-5 m, colour green, pink tips.
C, D. Sturdy vertical humps from colony, G 10259, Bushy-Redbill
Reef, W. slope. 10 m.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 45
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 46
Acropora rotumana (Gardiner, 1898)
A, B. Specimen from branch lip, G10235, Big Broadhurst Reef,
SW. side, outer reef flat, colour pale green.
C, D. Two branches of colony, G10263, Bowden reef, SW. side,’
reef crest, colour pink-brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 46
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl,ATl^ 47
Acropora ahroianoides (Lamarck, 1816)
A, B. G 10239 Bushy-Redbill reef, NW. side, reef crest, colour
yellow-brown.
C, D. G 10238 Big Broadhurst reef, upper surface of a patch reef
in the lagoon, colour yellow-brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 47
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 48
Acropora intermedia (Brook, 1891) on Bushy-Redbill Reef.
A. Large thicket in deep water middle reef flat on leeward (W.)
side.
B. Small patches with A. digitifera (centre) on reef crest, SW.
side.
C. Small patch (left) with A. aspera on inner middle reef flat, SW.
side.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 48
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 49
Acropora intermedia (Brook, 1891)
A. B. G1 1400 Big Broadhurst Reef, SW. slope, 2 m, colour bright
blue.
C, D. G11295 Big Broadhurst Reef, SW. slope, 2 m, colour pale
green.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 49
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 50
A. B. Acropora grandis (Brook, 1892) G11298 Darley reef, patch
reef in lagoon, 4 m, colour creamy brown.
C, D. Acropora brueggemanni (Brook, 1893) G 11498 Big
Broadhurst Reef, SW. slope, 6 m, colour brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 50
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 51
Acropora formosa (Dana, 1846) on the reef.
A. Large thickets on sandy bottom in deep middle reef flat,
Bushy-Redbili Reef, leeward (W.) side.
B. Thicket (foreground) with diverse Acropora assemblage in
background, base of upper reef terrace. Big Broadhurst Reef
SW. side.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 51
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 52
Acropora formosa (Dana, 1846)
A, B. G11339, Big Broadhurst reef, 2 m, colour pink-grey.
C, D. G11381 Bowden Reef, floor of opening in SW. side, 5m.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 52
MEMOIRS OF THE QUEENSLAND MUSEUM
Plath 53
Acropora splendida Nemenzo, 1967 on Big Broadhurst Reef, SW.
side.
A. Colony on gently-sloping upper reef terrace (about 7 m)
surrounded by flat-plate species. The distinctive ‘arborescent-
bracket’ shape can be seen.
B. Colony in a mixed assemblage at about 10 m (centre of plate).
This colony is flatter than the previous, and its shape is somewhat
disguised by another arborescent colony (A. formosa, lower left
and right). The small flat colony upper left of A. splendida is
A. clathrata: the large plate above this is A. hyacinthus, and
above it to the right is A. cytherea. The small dense bushy colony
(upper left) is A. haimei.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 53
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATF 54
Acropora splendida Nemenzo, 1967.
A, B. Specimen from inner part of large bracket, G8699, Big
Broadhurst Reef, SW. slope, 6 m, colour brown, pale blue
tips to branches.
C, D. Specimen from outer part of bracket, G8701, Big Broadhurst
Reef, SW. slope, 7 m, colour olive green, pale green tips to
branches.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 54
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 55
Acropora horrida (Dana, 1846)
A. Shrub like formations (centre) with soft corals in deep water
leeward (W) middle reef flat on Bushy-Redbill Reef.
B. ‘Bottlebrush* branch from C, below.
C. Sprawling arborescent formation on sandy bottom, 15 m depth,
ofUreef floor. Bushy-Redbill Reef, NW. side.
* 4 %
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 55
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 56
Acropora horrida (Dana, 1846)
A, B. Piece of slirub-like colony, G9078, Bushy-Redbill Reef, NW.
side, middle reef flat, colour lavender-grey.
C, D. Piece of colony, open arborescent, proliferous with short
branchlets, 09077, Bushy-Redbill Reef, NW. side, floor
outside slope, 8 m, colour pale blue.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 56
MEMOIRS OF THE QUEENSLAND MUSEUM
pLATi-; 57
Acropora vaughani Wells, 1954.
A, B. G 10270, Big Broadhurst Reef, SW. slope, 7 m, colour pale
brown.
C, D. G 1027 1 Big Broadhurst Reef, SW. slope, 8 m, colour pale
brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 57
MEMOIRS OFTHEQUEENSLAND MUSEUM
Pl,ATE 58
Acropora pulchra (Brook, 1891)
A, B. G1 11 18, Heron Island reef flat. Low corymbose colony with
slender branches.
C, D. Gill 16 Heron Island, W. side, reef flat, colour
cream-brown, blue tips. Corymbose colony with thick
branches.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 58
MEMOIRS OF THE QUEENSLAND MUSEUM
P\ .\Ti[ 59
Acropora pulchra (Brook, 1891)
A, B. Glini Heron Island Reef flat. Compact arborescent
colony.
C, D. Specimen from reef flat adjacent to Redbill Island. Open
arborescent colony.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 59
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl,ATE 60
Acropora aspera (Dana, 1846) on the reef.
A, B. Microatoll type development of corymbose colonies on middle
reef flat. Heron Island reef, W. side. (Scale divisions are 2
cm).
C. Large thicket (left half of picture) in contact with thicket
of A. pulchra (right hand side) on inner middle reef flat,
Bushy-Rcdbill Reef, SW. side.
rfM-
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 60
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 61
Acropora aspera (Dana, 1846)
A, B. G1096I, Bushy-Redbill Reef, N. side, inner flat-algal bank
area, colour brown, blue tips.
C, D. G 10930 Big Broadhurst Reef, patch reef in lagoon, colour
pale green. (Specimen typical of ‘A. hebes').
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 61
\
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 62
Acropora aspera (Dana, 1846)
A, B. G 10929 Big Broadhurst Reef, patch reef in lagoon, colour
pale green.
C, D. G 10921 Heron Island reef flat, colour pale brown. (Specimen
typical of A. aspera 5 . 5 .)
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 62
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 63
Large flat plates (Acropora hyacinthus. A. cytherea and A. clathrata)
and arborescent species, on Big Broadhursl Reef. (N.B. species
cannot be distinguished).
A. On upper reef terrace.
B. On surge channel floor. Note the change in arborescent to flat
plate species ratio between this and the reef terrace in the
background.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 63
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 64
Large flat plate species on the reef.
A. Acropora hyacinthus (upper left), A. cytherea (lower left), A.
clathrata (centre). With A. florida (left of centre) and
arborescent species on the surge channel floor (approx. 10 m)
at Big Broadhurst Reef.
B. A. hyacinthus with other Acropora in the Darley Reef lagoon.
Note oblique additions to the original flat plate.
C. A. clathrata (centre) with A. hyacinthus (right) and A. formosa
(left) on the reef crest at Bushy-Redbill Reef.
D. Three juvenile A. cytherea colonies on the reef crest at
Bushy-Redbill Reef.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 64
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 65
Acropora hyacinthus (Dana, 1846) on Bushy-Redbill Reef.
A. Deep middle reef flat area on leeward (W.) reef (1-5 m at low
tide^ with A. formosa.
B. Outer reef flat, SW. side. Several colonies with A. digitifera
(centre and lower centre) and A. formosa.
C. Reef crest, SW. side. Two colonies (arrows) with A. austera
(centre left), A. digitifera (centre) and A. humilis (centre
right).
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 65
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 66
Acropora hyacinthus (Dana, 1846) and A. cytherea (Dana, 1846)
A, B. A. hyacinthus portion of colony, G9865 Big Broadhurst Reef,
SW. side, reef slope, 5-3 m, colour pink-brown.
C, D. A. cytherea portion of colony, G9856 Big Broadhurst Reef,
SW. side, reef slope, 3 m, colour pink-brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 66
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 67
Acropora cytherea (Dana, 1846)
A. G7291 Flinders Reef, Moreton Bay, SE. Queensland.
B. G9855 Big Broadhurst Reef, SW. slope, 5 m, colour pale pinkish
brown.
C. G9849 Big Broadhurst Reef, SW. slope, 9*7 m, colour
yellow.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 67
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 68
Acropora millepora (Ehrenberg, 1834)
A, B. G11064 Bowden Reef, slope of opening in SW. side, 2 m,
colour orange.
C, D. G11049 Bushy-Redbill Reef, NW. side, outer reef flat,
colour pale brown, yellow tips.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 68
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 69
Acropora delicatula (Brook, 1891)
A, B. GI1441 Big Broadhurst Reef, SW. slope, 6
brown.
C. G11435, Big Broadhurst Reef, SW. slope, I
i, colour pale
m.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 69
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 70
Acropora haimei (Milne Edwards and Haime, 1860)
A. Compact arborescent clump on gentle reef slope on SW. side
of Big Broadhurst Reef, about 5 m depth, (centre of picture)
with flat plate species A. hyacinthus and A, cyfherea, and other
arborescent species.
B. Turf like patch on middle reef flat at Bushy-Redbill Reef, SW.
side, with soft corals, Seriatopora (lower left) and Acropora
palifera (lower right).
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 70
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 71
Acropora haimei (Milne Edwards and Haime, 1860)
A, B. G 10222 Big Broadhurst Reef, SW. slope, 4-2 m,
cream.
C, D. G 102 1 7 Big Broadhurst Reef, SW. slope, 9-8 m,
colour
colour
cream.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 71
20mm
10mm
30mm
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATE 72
Acropora tenuis (Dana, 1846) on the leeward (W.) side of
Bushy-Redbill Reef.
A. Corymbose plate (arrow) on outer middle reef flat. With sponges
and soft coral.
B. Layered colony in middle reef flat area, sandy bottom. With A.
palifera, Pocillopora and Seriatopora.
C. Layered colonies on reef crest.
m
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Platc 72
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 73
Acropora tenuis (Dana, 1846)
E. G11418 Darley Reef, patch reef in lagoon, colour pale
brown.
GI1428 Big Broadhurst Reef, SW. slope, 6 m, colour
pink-grey.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 73
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 74
Acropora tubicinaria (Dana^ 1846)
A, B. G11072, Bushy-Redbill Reef, W. side, inner reef flat, colour
pale brown.
C, D. G 11077, Bushy-Redbill Reef, W. side, inner reef flat, colour
pale brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 74
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 75
Acropora aculeus (Dana, 1846) on the leeward side of Bushy-Redbill
Reef
A. Plate like colony on middle reef flat (arrow) with soft corals
and sponges.
B. Layered colony on reef crest (bottom of picture), with A.
formosa (left), lA. haimei (right), A. palifera (centre), A.
clathrata (plate) and other corals.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 75
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 76
Acropora aculeus (Dana, 1846)
A, B. From Big Broadhurst Reef, SW. slope, about 10 m. Small,
thin plate specimen.
D. Large tiered plate colony in deep water middle reef flat area,
Bushy-Redbill Reef, W. side.
C. Portion of a large tiered plate.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Platf 76
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 77
Acropora cerealis (Dana, 1846)
A, B. G9519 Bowden Reef, slope of opening in SW. side, 3m,
colour blue.
C. D. G9529 Big Broadhurst Reef, SW. slope, 8-7m, colour
cream.
E, F. G9526 Darlcy Reef, patch reef in lagoon, 3m, colour pale
brown-lavender.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 77
^ o
MEMOIRS OF THE QUEENSLAND MUSEUM
Plath 78
Acropora nasuta (Dana, 1846)
B. Colonies in the deep middle reef Hat of Bushy-Redbill Reef,
. side with sandy bottom and soft corals.
11224 Big Broadhurst Reef, SW. slope, 7m, colour
grey.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 78
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi.ATi-: 79
Acropora diversa (Brook, 1891)
A, B. Gl 1246 Big Broadhurst Reef, SW. slope, 8m, colour purple,
cream tips.
C, D. G11245 Big Broadhurst Reef, SW. slope l-3m, colour
cream.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 79
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 80
A, B. A. diversa (Brook, 1891) G 11236 Big Broadhurst Reef, SW.
slope 5-1 m, colour blue-brown.
C, D. A. variabilis (Klunzinger, 1879) G11290 Big Broadhurst
Reef, SW. side, outer reef flat, colour cream.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 80
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate-. 81
Acropora humilis (Dana, 1846) and A. millepora (Ehrenberg, 1834)
on the reef.
A. Reef edge at Big Broadhurst Reef, SW. side, looking back across
the reef flat. A. rohusta and A. hyacinthus in the foreground:
most of the other colonies are A. humilis.
B. A. humilis colony on sandy bottom, deep middle reef flat,
Bushy-Rcdbill reef, W. side. This colony could be regarded as
forma a — . forma />*.
C. Outer reef flat on windward side (SE.) of Bushy-Redbill Reef.
Colony in centre may be regarded as A. humilis forma ; . and
that to its left as A. humilis forma P . To the right is another
low A. humilis. and other corals are A. palifera and
Pocillopora.
D. A. millepora (centre) on reef flat adjacent to Redbill Island;
to its left is A. humilis forma V , and another small A. millepora
colony. To the right is A. aspera.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 81
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi.Aii- 82
Acropora humilis (Dana, 1846)
A, B. G1 161 1 Darley Reef, patch reef in lagoon, 5m, colour pale
brown, lavender tips. Forma a Wells.
C, D. G11189 Bowden Reef, slope of opening in SW. side. Forma
(\ Wells.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 82
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATH 83
Acropora humilis (Dana, 1846) (Specimens not mentioned in
text).
A. B. 011 150 Big Broadhurst Reef, weather (E.) side, 2m, colour
blue. A. humilis forma v Wells.
C. D. Gil 142 Big Broadhurst Reef, SW, slope, 2m. Forma ^ »
forma ft Wells.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 83
MEMOIRS OF THE QUEENSLAND MUSEUM
Plath 84
Acropora digitifera (Dana, 1846)
A. Colonies on outer reef flat adjacent to Redbill Island (small
colony centre left is A. milleporaj
B, C. G1 1 169 Big Broadhurst Reef, outer reef flat, colour cream,
blue tips.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 84
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 85
Acropora multiacuta Nemenzo, 1967. All specimens from Darley
Reef, patch reef in lagoon, all pale blue.
A. G1047!.
B. G10468.
C. D. G6721.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 85
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 86
Acropora clathrata (Brook, 1891)
A, B. G9762, whole colony, Bowden Reef, SW. side, reef slope Im,
colour lavender-brown.
C, D. G9753, portion of colony, Big Broadhurst Reef, SW. side,
reef slope, 5ni, colour yellow-brown.
E, F. G9750, portion of colony. Big Broadhurst Reef, SW. side,
reef slope, 5m, colour brown, with mauve tips to
branchlets.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 86
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 87
Acropora divaricata (Dana, 1846) (C is a whole colony, others are
portions of colonies).
A. G9162 Big Broadhurst Reef, SW. slope, 124m, colour brown,
blue tips.
B. G9181 Big Broadhurst Reef, SW. slope, 7m, colour brown, blue
tips.
C. G9182, Darley Reef, patch reef in lagoon, 7m, colour brown,
lavender tips.
D. G9173 Big Broadhurst Reef, SW. slope, 8-7m, colour
brown.
E. G9174 Big Broadhurst Reef, SW. slope, 8-6m, colour brown,
blue tips.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 87
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 88
Acropora divaricata (magnified views of specimens in plate 87)
A. G9162, see Plate 87A.
B. G9181, see Plate 87B.
C. G9173, see Plate 87D.
D. G9174, see Plate 87E.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 88
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 89
Acropora sarmentosa (Brook, 1892)
A. B. G9057, Big Broadhurst Reef, SW. slope, 7-6 m, colour
cream-brown.
C, D. G9059, Big Broadhurst Reef, SW. slope, 9-7 m, colour green,
pink-grey tips.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 89
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 90
Acropora florida (Dana, 1846) on Big Broadhurst Reef.
A. In mixed assemblage on floor of surge channel at 10 m (centre).
Rounded, open-arborescent colony. Branches are round in
cross-section, secondary branchlets are evenly distributed and
sized. Coral in bottom left corner is A. splendida.
B. Colony on reef slope, at about lOm (arrow). Branches here are
more or less horizontal, and are flattened in cross-section.
Secondary branchlets arc differentially developed on the upper
surface.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 90
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 91
Acropora florida (Dana, 1846)
A, B. G8651. portion of colony. Big Broadhurst Reef, SW. slope,
5- 8m, colour brown.
C. G8665, whole colony. Big Broadhurst Reef, SW. slope, 6m,
colour pink-brown.
D. G8657, portion of colony. Big Broadhurst Reef. SW. slope,
6- 6m, colour green-brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 91
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 92
Acropora JJorida (Dana, 1846)
A, B. G8655 Big Broadhurst Reef, SW. slope, , 6-6m, colour pale.
pinkish brown. (This specimen resembles A. florida
C, D. G8647 Big Broadhurst Reef, SW. slope, 9-3m (flat plate
specimen).
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 92
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 93
Acropora echinata group on Big Broadhurst Reef, SW. side.
A. On surge channel floor at 10m. A. carduus (Dana, 1846).
(arrow) with a small table (7^4. cytherea) and A. Jlorida
(surrounding).
B, On gentle reef slope, about 12m. A. longicyathus (Milne
Edwards and Haime, I860) (arrow). Above this is A. splendida.
and A. divaricaia, upper right.
WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 93
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 94
A, B. Acropora subglahra (Brook, 1891) G10714 Lizard Island
fringing reef, colour pinkish brown.
C, D. Acropora carduus (Dana, 1846) G10731 Big Broadhurst
Reef, SW. slope, 8m, colour brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 94
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi. ATI-: 95
Acropora longicyathus (Milne Edwards and Haime. I860)
A. C. G 10767 Big Broadhurst Reef, SW. slope, 7 m, colour
brown.
Portion of colony from similar location.
B.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 95
MEMOIRS OF THE QUEENSLAND MUSEUM
Pi. ATE-; 96
Acropora microphfhalnia (Verrill. 1869)
A, B. G8685. Big Broadhurst Reef, SW. slope, 6m, colour
brown.
C, D. From Big Broadhurst Reef, SW. slope.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Pi.ATt: 96
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 97
A, B. Acropora elseyi (Brook) G 10793, Bushy-Redbill Reef, W.
side, middle reef flat, 1 m, colour yellow.
C, D. Acropora rosaria (Dana) G9202, Palm Islands, 1929, T. C.
Marshall.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 97
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 98
Acropora squarrosa (Ehrenberg, 1834) on the reef.
A. Bottlebrush branching colony on reef crest, Bushy-Redbill Reef,
W. side, colour blue (arrow).
B. Bottlebrush branching colony, (with shortened brushes) on edge
of hard/soft coral patch, middle reef flat, Bushy-Redbill Reef,
W. side, colour yellow-brown (arrow).
C. Caespitosc ‘incipient bottlebrush' colony on lagoonal patch reef,
Darley Reef (arrow).
D. Bottlebrush branching colony on sand, deep middle reef flat,
Bushy-Redbill Reef, W. side, colour lavender-blue.
E. Bottlebrush thicket in deep water reef flat at Wistari reef, near
Heron Island, (photo courtesy D. Ross Robertson).
F. Caespitose colony on lagoonal patch reef, Darley reef, colour pale
blue.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 98
MEMOIRS OF THE QUEENSLAND MUSEUM
Pl.ATE- 99
Acropora squarrosa (Ehrenberg, 1834)
A, B. G10782 specimen comparable to 'A. murrayensis’ Darley
Reef, patch reef in lagoon, 1 m, colour lavender.
C, D. G 10842 specimen approaching A. syringodes’ (part). Big
Broadhurst Reef, SW. side, reef slope, 9-2 m, colour
cream.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 99
MEMOIRS OF THE QUEENSLAND MUSEUM
PiATi: 100
Acropora squarrosa (Ehrenberg, 1834)
A, B. G 10846 low cacspitosc specimen. Dariey Reef, patch reef in
lagoon, 3 m, colour patches ol pale yellow, lavender and
blue.
C, D, E. G10845 Hal plate specimen (D is undersurface). Big
Broadhurst Reef, SW. side, reef slope, 12-9 m, colour pale
blue-brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 100
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 101
Acropora granulosa (Milne Edwards and Haime, I860). Specimens
with blunt axial corallites.
A, B. Gl 1487, Big Broadhurst Reef, SW. side, reef slope, 10-4 m,
colour cream-grey.
C, D. GI1492, Big Broadhurst Reef, SW. side, reef slope, 8 m,
colour brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 101
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 102
Acropora granulosa (Milne Edwards and Haime, 1860). Specimens
with tapering axial corallites.
A, B. G6723 (comparable with 'A. speciosa'}. Darley Reef, patch
reef in lagoon, 2*5 m, colour pale brown.
C, D. GI 1476 (comparable with ‘A. raynerV). Big Broadhurst Reef,
SW. side, reef slope, 5 m, colour pale brown.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 102
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 103
Acropora austera (Dana, 1846)
A, B. G 10828 Big Broadhursl Reef, SW. slope, 7-8 m, colour
cream.
C, D. G10810 Bushy-Redbili Reef, S. side, reef slope, 3m, colour
lavender.
WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF
Plate 103
I
CONTENTS
Pag(
Bartholomai. Alan
The Macropodidae (Marsupialia) from the Allingham formation, northern Queensland. Results of
the Ray E. Lemley Expeditions, Part 2 127
Bartholomai. alan
The rostrum in Palorchestes Owen (Marsupialia: Diprotodontidae). Results of the Ray E.
Lemley Expeditions, Part 3 ]45
CovACEVicH. J. and Ingram. G. J.
An undescribed species of rock dwelling Cryptoblepharus (Lacertilia: Scincidae) 151
Archfr, Michall
The nature of the molar-premolar boundary in marsupials and a reinterpretation of the
homology of marsupial cheekteeth 157
Archlr. MICHAEL and Brayshaw. Helen
Recent local faunas from excavations at Herveys Range, Kennedy, Jourama,
and Mount Roundback, north-eastern Queensland 165
Cannon. L. R. G.
Pterastericola vivipara n. sp., a parasitic turbellarian (Rhabdocoela: Pterastericolidae)
from the Crown-of-Thorns starfish, Acanthaster planci 179
Stephenson. W., Cook, S. D., and Newlanos. S. J.
The macrobenthos of the Middle Banks area of Moreton Bay 185
Greenwood, J. G.
A new species of Pontellopsis (Copepoda, Calanoida) from Moreton Bay, Queensland 213
Jeli . Peter A.
Asihenopsis Whitehouse, 1939 (Trilobita, Middle Cambrian) in northern Australia 219
Bruc E. A. J.
Paranchistus pycnodontae sp. nov., a new pontoniine shrimp associated with an
ostreid bivalve host 233
Jones. H. I.
Gastrointestinal nematodes from aquatic Australian snakes 243
Philups, W. j.
Some parasitic barnacles (Rhizocephala: Sacculinidae) from portunid crabs in Moreton
Bay, Queensland 255
Gh l. j. C. H.
The old burial vault at North Quay, Brisbane 265
Wallace. Carden C.
The coral genus Acropora (Scleractinia: Astrocoeniina: Acroporidae) in the central!
and southern Great Barrier Reef Province 273
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