MEMOIRS
OF THE
QUEENSLAND MUSEUM
VOLUME 39
BRISBANE
PART 2
20 JULY, 1996
PHYLOGENETIC STATUS OF THE IRRAWADDY DOLPHIN ORCAELLA
BREVIROSTRIS (OWEN IN GRAY): A CLADISTIC ANALYSIS
PETER W. ARNOLD AND GEORGE E. HEINSOHN
Arnold, P.W. & Heinsohn, G.E. 1996 07 20: Phylogenetic status of the Irrawaddy Dolphin
Orcaella brevirostris (Owen in Gray): a cladistic analysis. Memoirs of the Queensland
Museum 39(2): 141-204. Brisbane. ISSN 0079-8835.
Examination of Orcaella brevirostris from Queensland has provided new information on
colour pattern, external morphometrics, skull morphology, variation in the tympanoperiotic
bones and postcranial skeleton. Facial anatomy is described for the first time.
Cladistic analyses, incorporating the new information, investigated the phylogenetic position
of Orcaella. Our results provide no support for the separation of beluga and narwhal into
two different families (Kasuya, 1973) nor for the proposal that Orcaella and Delphinapterus
are closely related (Kasuya, 1973; Pilleri et al.,1989). The position of delphinoid families in
our cladograms is consistent with previous synoptic classifications (Slijper, 1962, fig.36;
Heyning,1989; Barnes,1990). Our results offer no support for classifications which widely
separate delphinids and phocoenids (Shimura & Numachi,1987; Lint et al., 1990; Pilleri et
al.,1989).
We suggest that Orcaella is a delphinid sensu lato. Comparison of characters in the two
nearest outgroups (phocoenids and monodontids) suggest Orcaella (and other *blunt-headed'
genera) represent the most primitive Delphinidae. However, we cannot rule out the alterna-
tive that extensive convergence occurred. Extensive neoteny of the skull in Orcaella
suggests one means by which the many apparently primitive features could occur.
O Cetacea, Orcaella, periotic bone, tympanic bone, facial anatomy, neoteny, cladistics.
Perer W. Arnold, Museum of Tropical Queensland, 70-84 Flinders Street, Townsville,
Queensland 4810; George E. Heinsohn , Department of Zoology, James Cook University of
North Queensland, Townsville, Queensland 4811; received 8 November 1995.
The phylogenetic status of the Irrawaddy dol-
phin Orcaella brevirostris (Owen in Gray,1866)
remains in doubt (Marsh et al.,1989). At least
eight taxonomic hypotheses are implied in
various proposed classifications (Fig.1). Orcael-
la was initially classified as a porpoise in
Phocoena. It has also been placed in a more
strictly defined Delphinidae (dolphins), Del-
phinapteridae (with beluga Delphinapterus
leucas (Pallas,1776)), Monodontidae (with
beluga and narwhal Monodon monoceros Lin-
naeus,1758) or in the monotypic Orcellidae.
This paper addresses classification of Orcaella.
We re-describe the skull, emphasizing neotenic
characters which have confounded previous com-
parisons between Orcaella and other odon-
tocetes. We re-examine the tympanoperiotic
bones which were poorly known and describe the
facial anatomy which was unknown. We assess
colour patterns for possible taxonomic charac-
ters. New data on external morphometrics and the
post-cranial skeleton are presented.
Incorporating this data we evaluate characters
that have been used in odontocete classification.
Most classifications have been based on one
anatomical system (Fraser & Purves,1962; Mead,
1975; Kasuya, 1973; Pilleri et al.,1989) and have
produced partially conflicting classifications.
Only Heyning (1989) used computerized
phylogenetic analysis on a range of characters.
Although he argued for a phocoenid-delphinid
clade, his cladograms left relationships of the
Monodontidae, Phocoenidae and Delphinidae
unresolved. Gretarsdottir & Arnason (1992) also
left the relationship of these 3 families un-
resolved. de Muizon (1988) examined extant and
fossil taxa, using a comprehensive range of char-
acters to produce 4 arrangements of the
Monodontidae, Phocoenidae & Albireonidae,
Kentriodontidae and Delphinidae, which were
still only a subset of the possibilities. We present
a cladistic analysis of the Monodontidae,
Phocoenidae and Delphinidae s. l. Within this
context we discuss affinities of Orcaella.
MATERIALS
Specimens of Orcaella were either found
stranded and dead, or recovered after drowning
in shark nets near Townsville. Skeletal material
142 MEMOIRS OF THE QUEENSLAND MUSEUM
Phocoena Other Delphinidae
* Orcaella (including Delphinapterus Phocoenidae Monodontidae Ziphiidae Physeteridae Delphinidae
Monodon) Monodon * Orcaella
Delphinapterus
A Owen, 1866
Delphinapterinae Orcaellinae E Linteral, 1990
Delphinapterus * Orcaella
|
on Spr EADEM EM PM Delphinnptorntae Ziphiidae Monodontidae Stenidae Phocoenidae Delphinidae
(Monodon Monodon Sousa * Orcaella
Delphinapterus | Sotalia
Steno
B Kasuya, 1973
F Fraser and Purves. 1962
Monodontinae Delphinapterinae — Orcaellinae Phocoenidae Delphinidae *
Monodon Delphinapterus & — * Orcaella Monodontidae
+ Denebola Monodon
Delphinapterus
L I | Phocoenidae Delphinidae
Monodontidae
Globicephalidae
Orcaellidae Grampidac
* Orcaella Grampus
+extinct
* includes Stenidae of Fraser & Purves, 1962
C Barnes, 1984; Gaskin, 1982; Evans, 1989
G Nishiwaki, 1963, 1966
Delphinidae Phocoenidae
Monodontidae Phocoenidae Delphinidae
Monodontidae Delphinapteridae Iniidae Pontoporidae Platanistidae Monodon * Orcaella
* Orcaella Delphinapterus
Delphinapterus
D Pilleri, Gihr and Kraus, 1989 H Heyning, 1989; de Muizon, 1988; Gretarsdottir and Arnoson, 1992
FIG. 1. Taxonomic hypotheses of phylogenetic relationship of Orcaella to other toothed whales. A indicates that
Orcaella was placed within Phocoena, which was more broadly defined than at present. The Delphinidae was
also more inclusive, containing both the beluga Delphinapterus and narwhal Monodon. The dashed lines in D
separate different lineages as envisioned by Pilleri et al. (1989). The taxonomic relationships in F are derived
from the table in Fraser & Purves (1962); the most generalized taxon is Ziphiidae on the left, and increasing
specialization is inferred as one moves to the right. The dashed lines indicate a separation of taxa: the
Physeteroidea (Physeter, Kogia) and Platanistoidea (Platanista, Inia, Pontoporia, Lipotes) were interspersed
between the Monodontidae and Stenidae. The references indicated in H are only a few of the more recent ones
supporting placement of Orcaella in the Delphinidae.
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
is deposited in the Queensland Museum (Pater-
son, 1986, 1994) (QMJ or QMJM). Most are
currently held at the Museum of Tropical
Queensland, Townsville; those held in Brisbane
are indicated by an asterisk. Specimens collected
by James Cook University staff are registered
MM or CET, which numbers have been quoted
in the literature; these numbers are used par-
ticularly when discussing anatomical material. If
skeletal material from these specimens has been
registered in the Queensland Museum, both the
MM and QMJM numbers are listed below. Ab-
breviations for other collections are: CMN:
Canadian Museum of Nature, Ottawa, Ontario,
Canada; NSMNH: Nova Scotia Museum of
Natural History, Halifax, N.S., Canada; UBC:
Dept. Zoology, University of British Columbia,
Vancouver, B.C., Canada; VA: Vancouver
Aquarium, Vancouver, B.C., Canada. Measure-
ments of beluga and narwhal are of condylobasal
length and are approximate.
Orcaella brevirostris (Owen in Gray,1866)
Queensland: QMJM4740; QMJM4735
(MM1004); QMJM4700 (MM006); QMJM4704
(MMO012); QMJM4708 (MM021); QMJM4709
(MM025); QMJM4712 (MM030); QMJM4714
(MM032); QMJM4721 (MM 061); QMJM4725
(MM081); QMJM4726 (MM082); QMJM4727
(MM088); MM092; MM1003; QMJM11342,
QMJM11343; MM1015; MMO16.
Phocoenidae
Neophocaena phocaenoides (Cuvier,1829)
Saudi Arabia: A. Preen,private collection.
Phocoena phocoena (Linnaeus, 1758) Nova
Scotia, Canada: NSMNH973.Z.309.1; NSMNH
unregistered, Oct. 26, 1982; NSMNH971.-
Z.300.1; NSMNH unregistered, Crescent Beach
1977; NSMNH973.Z.310.1.
Monodontidae
Delphinapterus leucas (Pallas,1776) Quebec,
Canada: Trois Pistoiles, May 11, 1983, 573 mm;
DL4.85, 400 mm; DL2.86, 468 mm; Northwest
Territories, Canada: CMN 19556, Collinson Inlet;
CMN29997, 548 mm; CMN29998-30000, Bel-
cher I, Hudson Bay, 505, 549 and 552 mm respec-
tively; one unregistered.
Monodon monoceros (Linnaeus, 1758) Canada:
CMN32278-32280, Baffin I, latter 508 mm;
Arctic Biological Station MM65(at CMN);
Arctic Biological Station MM66, 580 mm (at
CMN); Koluktoo Bay, Baffin I: UBC 9467, 285
mm; Holman I, Northwest Territories: VA, un-
registered; VA, unregistered (no data).
143
Delphinidae
Delphinus delphis Linnaeus, 1758
*QMJM2033, Gold Coast; *QMJM2776,
Moreton Bay.
Feresa attenuata Gray, 1874 *QMJM825,
Kingscliff, NSW.
Globicephala macrorhynchus Gray, 1846
*0MJMS354; CET1001, Mackay.
Globicephala melas (Trail, 1809)
*QMJM4480, Point Lookout; *QMJ15.2104.
Grampus griseus (Cuvier, 1812) *QMJM9542,
N. Stradbroke I; *QMJM3858, Moreton I.
Lagenodelphis hosei Fraser, 1956 *QMJM
2749, Fraser I.
Lagenorhynchus acutus (Gray, 1828) NSMNH
unregistered.
Lagenorhynchus albirostris (Gray, 1846)
NSMNH 72.2.343.8.
Peponocephala electra | (Gray,1846)
*QMJM2144, Moreton I; *QMJM6577, N.
Stradbroke I; *QMJM7854; QMJM4702, Mis-
sion Beach; QMJM4730, Crystal Creek, N of
Townsville.
Pseudorca crassidens (Owen,1846)
*QMJM14210; *QMJM937, Townsville;
MM1028.
Sousa chinensis (Osbeck, 1765) QMJM4701,
Magnetic I; QMJM4703, Magnetic I;
QMJM4711, Townsville; QMJM4717, Magnetic
I; QMJM4728; QMJM4731, Pallarenda,
Townsville; QMJM4737, Rowes Bay,
Townsville.
Stenella attenuata (Gray, 1846) *QMJM6433,
Moreton I.
Stenella coeruleoalba
*QMJM3859.
Stenella longirostris (Gray, 1828) QMJM4716,
QMJM4718, QMJM4719, all from off Michael-
mas Cay, near Cairns.
Tursiops truncatus (Montagu, 1821)
*QMJM8859; QMJM4713, juvenile, Magnetic I;
QMJM4715, Magnetic I; QMJM4724, Magnetic
I; MM91A, Palm I; MM1018.
(Meyen, 1833)
METHODS
Colour pattern is described from photographs
of acaptive from Cairns ( Dawbin,1972; Leather-
wood & Reeves,1983; Mitchell,1975:911 ); a
1.86m ? (MM334) and 2.19m 9 (MM335) from
Cairns; a 2.15m 9 (MM30), 1.87m 9 (MM21)
and 2.2 m 9? (MM25) from near Townsville
(Talbot & Steene,1984; unpubl. photographs).
Based on dentinal layers in teeth, MM25 was
estimated at 9 years and MM30 an estimated 8
144 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 2. Colour pattern of MM30, a 2.15 m long ? from Townsville.
Fig 4. Ventral colour pattern of MM25, a 2.2 m long 9 from Townsville.
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
years in age (Heinsohn,1979). Lengths of both
animals exceeded the minimum length of con-
firmed sexually mature 9 9 in the Queensland
population (Marsh et al.,1989); MM335 was
pregnant.
Qualitative features of the skull were compared
with Tursiops (Rommel,1990). Standard skull
measurements are as in Perrin (1975).
The angle of the posterior process of the tym-
panic was determined by temporarily fixing the
bulla ventral side upwards on a desk. One arm of
a compass was aligned along the meridional axis;
the second arm was swung to lie over the lateral
edge of the process. The angle so formed was
traced onto paper and measured with a protractor.
The periotics were temporarily fixed dorsal
side up on the stage of a compound microscope
and the positions of the three cochlear apertures
drawn using a camera lucida. Care was taken to
orient the bulla in the same way to avoid parallex
problems. The periotic triangle of Pilleri et al.
(1989) was created by drawing lines between
outlines of the apertures on the drawing. Angles
and lengths of sides of the triangles were deter-
mined from the drawings.
Three specimens of Orcaella (MM333, 1.34m
9 from Mackay, MM334, MM335, 9? 9 from
Ellis Beach, Cairns) were dissected to examine
the upper respiratory tract and facial region. The
frozen head of MM334 was sectioned lon-
gitudinally on a band saw. The blowhole was set
well to the left so the first cut was just to the left
of the median line, and the second c.2cm further
left. The latter section passed through the tym-
panoperiotic bones and pterygoid region. Facial
musculature was not examined, but the relative
proportions of muscle, connective tissue and
*melon' were assessed. The 'melon' was dif-
ferentiated from connective tissue by its lesser
vascularisation (Mead,1975) and the more fatty
appearance relative to muscle and connective
tissue.
Phylogenetic analyses used Hennig86, version
1.5 (Farris,1988). The implicit enumeration op-
tion was chosen to find all of the most par-
simonious trees. The implicit enumeration
method is time consuming for large data sets so a
subset of delphinid genera were analysed. Repre-
sentatives of most delphinid genera were ex-
amined; character states within the genera used
to generate the cladograms cover the range of
variation within the Delphinidae. Wherever pos-
sible, characters were reduced to binary values to
avoid some of the problems with multistate char-
acters. All characters were set to non-additive.
145
In this analysis, only extant families of odon-
tocetes were considered as outgroups, although
character states in fossil taxa have been con-
sidered in certain cases. We have not examined
fossil material, and many features used have not
been described in the literature available to us. To
include them would result in many missing
values, which can cause problems. The
Kentriodontidae, which has been considered a
potential sister group to delphinids, phocoenids
and monodontids, cannot be clearly defined (de
Muizon,1988) and may be polyphyletic. de
Muizon (1993) considered the peculiar
Odobenocetopsidae the sister group to the
Monodontidae, but it is so highly modified that it
is not relevant to our discussion.
Among extant odontocetes, the river dolphins
Platanista, Lipotes, Inia and Pontoporia are con-
sidered the closest living relatives of the
Monodontidae-Phocoenidae-Delphinidae (de
Muizon,1988; Heyning,1989; Barnes,1990). The
first two authors also demonstrated that Platanis-
ta is separate from the other genera; de Muizon
(1990) placed it in a separate superfamily. In the
present comparisons, the taxonomic status of the
river dolphins can be left unresolved, but
Platanista was considered separately from /nia
and Pontoporia, and was used as the outgroup.
Character states of features used in this analysis
were also determined for Berardius, a primitive
ziphiid (Moore, 1968) and Physeter, to determine
polarity of characters. In cases where the charac-
ler was not present in either physeterids or
ziphiids, the character state in the fossil taxon
Eurhinodelphoidea was considered to help deter-
mine the polarity. de Muizon (1990) considered
the Eurhinodelphoidea as the sister taxon to Del-
phinida (his taxon, which encompasses all genera
considered here, except Platanista). Reasoning in
these cases is therefore not that the character is
primitive because it occurs in a fossil taxon but
because it occurs in a putative sister group, the
members of which happen to be all extinct.
Polarity decisions for all characters are justified
in Appendix 2.
COLOUR PATTERN
RESULTS. A broad grey or blue-grey cape
(‘spinal field’ of Mitchell, 1970) extends from the
top of the head and back downwards about half
the distance to the level of the eye, continuing
posteriorly to just behind the base of the dorsal
fin, from which it tapers as a wedge along the
upper half of the tail stock (captive animal: Daw-
146
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 5. Dorsal view of QMJM4721, with 312.8 mm
condylobasal length and 11.5 dentinal layers, Note
broad transverse width of neurocranium; super-
numerary bone ( triangle) infilling the postnarial
fossa; conspicuous triangular interparietal bone
(arrow); dorsal extension of parietal bones,
posterolateral to the interparietal bone; wide separa-
tion of exoccipital and frontal bones; weakly ex-
pressed telescoping (wide expanse of frontal bone
exposed); prominent spina mesethmoidalis (open
arrow); poorly developed nuchal crest.
bin, 1972; MM30: Fig. 2; Talbot & Steene,
1984:300; MM25: Fig.3). The dark grey is more
extensive on the melon of MM25 than MM30
(Figs 2,3). Grey extends onto the lower jaw and
throat region in the captive animal (Leatherwood
& Reeves, 1983:154) and MM25 (Fig.4) but in the
latter, a lighter grey throat patch extends as far
back as the axilla of the flippers (Fig.4). The
abdominal field from the flipper region to the
genital region in MM25 and MM30 (Figs 2,4) is
white. The flanks between the dark cape and
white abdominal field are light grey to brownish-
FIG. 6. Dorsal view of QMJM4740. Note triangular
area of spongy bone at base of rostrum, extending
forward to just in front of spina mesethmoidalis;
nodular nasal bones (open arrow); supernumerary
bone; poor development of mesethmoid plate
(posterior margin indicated by arrow) and consequent
exposure of frontal bones anterior to nasal bones;
wide exposure of frontal bone (triangle) between the
interparietal and maxillary bones.
grey (Talbot & Steene,1984; Figs 2,3); the light
grey extends onto the tail stock.
MM334 has the subtle 3-tone pattern, but the
dorsal fin on both sides is lighter than the back.
This does not appear to be a post-mortem effect.
DISCUSSION. Underwater photographs of O.
brevirostris from the Mahakan River, Indonesia
(Tas'an & Leatherwood, 1984; unpubl. photos by
Dr. A. Preen) do not show the three tone colour.
This may be a real difference, or simply reflect
the difficulty in differentiating subtle shading
with the rapid loss of contrast underwater.
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
Anderson (1879, pls 25, 25a) described O.
brevirostris as ‘dark slaty-blue, nearly black, and
very little paler on the ventral surface’. It is un-
clear how long his specimens had been dead;
postmortem darkening of cetaceans càn be sub-
stantial (Pilleri,1976). Anderson (1879) noted
that living O. fluminalis (=brevirostris) were
more lightly coloured . The dark grey to black
colouration of Orcaella needs to be confirmed on
living animals.
The subtle 3-tone colouration of Queensland
Orcaella may be found in juveniles of several
dolphin genera, becoming fainter or lost in adults.
However, given the sizes and ages of the animals
examined, the colour patterns we describe appear
to be characteristic of adult animals.
Mitchell (1970) considered the three-tone pat-
tern as the ‘common baseline from which to
interpret most of the patterns found within the
Delphinidae'. Its occurrence in Orcaella thus can
not be used as evidence for relationship to other
delphinid genera. The extension of grey onto the
lower jaw and throat in Orcaella is similar to the
pattern in Globicephala, Pseudorca, Pepono-
cephala and Grampus. None of the Orcaella had
a throat chevron, midventral stripe or genital
patch as occurs in those genera (Minasian et al.,
1987; Mitchell 1970), although MM25 had a
vague light grey throat patch similar to a throat
chevron. The throat chevron is not restricted to
the ‘blunt- headed’ whales; Tursiops truncatus cf
aduncus may have a distinct throat chevron
(G.J.B. Ross pers. comm.). The photographs of
Orcaella are not of sufficient quality to show
unequivocally more specialised features such as
spinal blaze, bridles or flipper stripe. In this case,
colour pattern is of little taxonomic use.
SKULL
RESULTS
REDESCRIPTION OF SKULL. Values
presented below are based on 14 animals from
central Queensland, Australia with condylobasal
lengths from 297.7—334.8mm and an estimated
age (based on dentinal layers) of 3-28 years. An
additional skull of a newborn, 232.1mm con-
dylobasal length, was examined for non-men-
sural characters, but measurements of this
specimen were excluded from the descriptive
statistics.
Unless otherwise indicated, values are percent-
ages of condylobasal length, based on measure-
ments on the left side of the skull. Orcaella has
147
cranial asymmetry as is general in delphinids; this
will be considered in detail in a separate publica-
tion.
DORSAL ASPECT. The neurocranium is broad
(Fig.5), 65.6% (62.6—68.095) at the postorbital
process of the frontal which is usually the widest
point of the skull. Zygomatic width is 65.4%
(63.2-68.3%).The lacrymal bones extend for-
ward from the maxillary bones to form the lateral
border of prominent antorbital notches, 4.096
(2.8-4.5%) deep.
The rostrum is 44.4% (43.2-45.5%)long and
37.6% (34.4-39.8%) wide at the base. Premaxil-
laries as a percentage of rostrum width are 50.5%
(47.0-57.6%) at the base, 62.8% (60.0-65.9%) at
0.25 length, 61.2% (58.1—65.196) at 0.5 length
and 65.0% (56.0-73.1%) at 0.75 length. The
spongy, triangular area of the rostrum, bounded
laterally by 2 oblique ridges, extends just beyond
the proximal quarter of the rostrum (Fig.6).
The premaxillaries continue onto the cranial
vault, showing distinct asymmetry at the point of
maximum width: the left premaxillary is 4.3%
(2.4-5.7%) wide and the right 8.5% (7.6-9.1)%
wide at this point. There is a prominent exposure
of the frontals between and behind the ascending
processes of the maxillaries (Fig. 5).
The vertex is composed mainly of the frontals,
with a median suture deflected to the left (Fig.5).
It forms an ill-defined ridge running along the
body axis between the ascending processes of the
maxillaries. The nasal bones are peculiar nodules,
often two on each side of the vertex (Fig.6), from
7.3-15.6mm long by 6.4-14.0mm wide. The
nasal bones sometimes coalesce, but always
retain their nodular appearance (Fig. 7). The
postero-medial pair of nasal nodules are at the
apex of the vertex while the anterolateral nasal
nodules are on the anterior face of the vertex. The
anterior face of vertex has a shallow postnarial
pit, usually filled in by a supernumerary bone
6.0-22.0mm long by 5.4—17.8mm wide (Figs 5—
T).
The mesethmoid plate is generally poorly
developed (Figs 5-8), leaving much of the
anterior face of the vertex exposed (Figs 6,8). An
elongate, shallow fossa occurs laterally in this
space, between the apical nasals and the meseth-
moid plate (Fig.8). When the mesethmoid plate is
more extensive, the fossa excavates its postero-
lateral margin, leaving a median extension of the
plate which reaches back to the supernumery
bone on the vertex (Fig.7).
The frontals are always distinctly separated
148
FIG. 7. Oblique anterodorsal view of vertex, QM
JM4714. Note nodular, partially coalesced nasal
bones (open arrow); development of mesethmoid
plate (posterior margin indicated by triangle); and
maxillary intrusions (arrow) along anterior margin of
superior nares.
from the supraoccipital by prominent dorsal ex-
tensions of the parietal bone and a triangular
interparietal bone (Figs 5,8).
Anteromedial borders of the superior nares are
edged by maxillary intrusions 8.2% (5.8-10.0%)
long and 2.4% (1.3-3.5%) wide (Figs 5-8). These
intrusions are weakly size dependent (r=0.649 for
left maxillary intrusion length vs condylobasal
length). A very prominent spina mesethmoidalis
extends anterior to the bases of the antorbital
notches and widely separates the premaxillaries
(Figs 5,6,8). Braincase width is 49.8% (47.0—
54.0%) across the parietals and 51.3% (48.2—
55.2%) across the squamosals.
LATERAL ASPECT. The skull is deep, 57.296
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 8. Dorsal view of QMJM4735. Note poor
development of mesethmoid plate (posterior margin
indicated by arrow); shallow lateral fossa between
mesethmoid plate and depressions for the nasal bones
(open arrow); prominent interparietal bone.
(55.3-60.6%) from vertex to the left paroccipital
crest.
The orbit is prominent (Fig. 9), 15.8% (15.0—
17.1%) between the preorbital and postorbital
processes of the frontal bone. The jugal is stout,
fitting within a prominent notch on the anterior
face of the zygomatic arch. The maxillary bone
over the lacrymal and preorbital process of fron-
tal is raised to a variable extent; when it is exten-
sively developed it imparts a concave profile to
the supraorbital plate of the maxillary bone.
The temporal fossa is bounded dorsally and
posteriorly by a weakly developed temporal crest.
The majority of the fossa is bounded medially by
parietal and base of the squamosal (Fig.9). The
zygomatic arch of the squamosal is prominent,
with an extensive mastoid section laterally and a
wide post-glenoid space (Fig.9). The paroccipital
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
149
* tie
FIG. 9. Lateral view of QMJM4721. Note prominent orbit; robust jugal bone (broken); oblique orientation of
occipital condyles; poorly developed temporal crests; ventral orientation of paroccipital process of exoccipital;
fissure (triangle) between exoccipital and prominent mastoid portion of zygomatic arch (open arrow).
process of the exoccipital is directed outwards
and ventrally, with minimal intrusion anteriorly
onto the zygomatic arch. A deep, usually Y-
shaped fissure separates the mastoid portion of
the squamosal from the paroccipital process of
the exoccipital (Fig. 9). The occipital condyles
are prominent and directed ventrally at c.45° to
the skull axis (Fig.9). Braincase length is 43.1%
(42.0-44,9%).
VENTRAL ASPECT. The posterior of the palate
contains triangular lateral lobes of the palatine
bones, each with a prominent foramen (Fig.10)
which forms the anterior end of a channel exten-
deding dorsally and backwards to the pterygo-
palatine fossa.
The lateral lobe of the palatines extends
posterolaterally as a wing-like process (Figs
10,11) 11.0.26 (9.3-12.5%) long along its anterior
margin, and underlying an extensive preorbital
cavity formed between the maxillaries and pres-
phenoid/frontal bones (Fig.11). This cavity ex-
tends dorsally as a lobe, more extensively
developed on the right than the left side.
The pterygoid hamuli are distinctly separated
by triangular medial lobes of the palatine bones
(Figs 10,11). The palatines flank the vomer and
spina mesethmoidalis, which is often incomplete-
ly covered by the vomer. Sometimes the vomer is
visible between the sutures of the palatines and
maxillary bones (Fig.12) but in most cases it is
indicated only by a pit.
A medial flange extends from the pterygoid
hamuli (Figs 10-12), bringing them to within
1.0% (0.6—1.5%) of one another and almost com-
pletely covering the inferior nares.
The lacrymal bones are massive (Figs 10,12),
12.4% (10.2-15.0%) long by 14.4% (13.4—
15.1%) wide. The frontal bones form a prominent
obliquely transverse ridge (Fig. 11) without an
optic groove or channel. The optic foramen is
incompletely separated from the anterior lacerate
foramen by a short vertical bony bridge formed
by the fused presphenoid-orbitosphenoid bones.
The alisphenoid bone is a prominent plate (Fig.
12), forming the anterior margin of the foramen
ovale (posterior margin of foramen provided by
an extension of the basioccipital).
The zygomatic arch has an extensive tym-
panosquamosal recess (Fig. 12). The falciform
process is reduced, 3.9% (3.0-5.1%, n=10) long
by 3.7% (2.8-4.9%, n=7) wide. It runs parallel to
the alisphenoid, rather than ventrally and is not
closely associated with the periotic bone. The
postglenoid space contains a wide groove for the
auditory meatus and a prominent triangular mas-
150
P
FIG. 10. Ventral view of QMJM4708. Note triangular
anterior portion of lateral lobe of palatine, containing
palatine foramen (arrow); posterior wing-like exten-
sion of lateral lobe of palatine (triangle); complete
separation of medial and lateral lobes of palatines by
pterygoid (open arrow); separation of pterygoid
hamuli by medial lobes of the palatines; medial flan-
ges on pterygoid hamuli.
toid pad (Figs 11,12), the latter providing ar-
ticulation for the tympanoperiotic bones. Be-
tween the base of the squamosal, just medial to
the mastoid, and a ventral extension of the parie-
talis a deep pit (Figs 11,12). In QM JM4709, this
is one end of a channel which opens on the back
of the skull in the suture between the squamosal
and parietal bones. The cranial hiatus is wide in
young animals, but could be completely infilled
in older animals.
MANDIBLE (Fig. 13). Mandibles are 76.7%
(75.4-78.6%) long, with coronoid depth 23.6%
(22.3-25.196). The mandibular fossa is 32.8%
(29.9-35.4%)long, while the mandibular sym-
physis is 7.7% (5.6-11.4%) long by 8.3% (6.5—
9.0%) deep.
Alveolar/tooth counts (mean, rounded to whole
number, followed by range in parentheses) are
18 (17-20y/ 18 (16-20) nz14
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 11. Ventral view of QMJM4721. Note wing-like
posterior extension of lateral lobe of palatine; medial
flange of pterygoid hamuli (triangle);transverse fron-
tal ridge anterior to optic channel; large pre-orbital
space for dorsal extension of pre-orbital lobe (curved
arrow); mastoid pad of zygomatic arch (broad arrow);
prominent groove for external auditory meatus (thin
arrow); deep pit medial to mastoid pad (open arrow).
17 (15-19)/ 17 (16-18) n=13.
Total alveolar/tooth counts are 66—78. Ap-
parently the central teeth erupt first; in MM334
the erupted tooth count is only 15/13 11/14.
COMPARISON WITH TYPE SPECIMEN.
Owen's (1866) description of the vertex and the
accuracy of his pl.9, fig.2 are confirmed by the
holotype (R. Sabin pers comm.; Fig.14). The
most significant differences in the vertex, as com-
pared with Australian material, are the 2 elongate
antero-posteriorly compressed nasal bones (cf
multiple nodular nasal bones) and well-
developed mesethmoid plate, abutting on the
nasal bones (cf reduced mesethmoid plate).
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
FIG, 12. Ventral view of QMJM11342. Note vomer
insunk between medial lobes of palatine (arrow);
incomplete separation of medial and lateral lobes of
the palatines; wing-like posterior extension of
palatine; medial flange on pterygoid hamuli;
prominent alisphenoid; extensive tympano-
squamosal recess of zygomatic arch (triangle); mas-
toid pad and groove for external auditory meatus.
FIG. 13. Lateral and medial views of mandible of
QMJM4708.
151
FIG. 14. Dorsal view of BM(NH)1865.4.20.1,
holotype of Orcaella brevirostris.
DISCUSSION
NEOTENIC FEATURES OF THE SKULL.
Winge (1921) noted that Orcaella ‘gives the im-
pression of being a dwarf form with noticeably
large braincase in proportion to the face’. This
appearance is due to massive development of the
skull roof. Comparison of newborn and mature
Orcaella (Fig. 15) with newborn and adult
Peponocephala (Fig.15) and Feresa (Fig.15)
show this to be a retained juvenile feature. In all
these cases, the interparietals and parietals form
major components of the skull roof and much of
the frontal bone is exposed in the newborn
animal. In adult Peponocephala and Feresa, the
parietals have been excluded from the roof of the
skull and the interparietal is reduced or complete-
ly obscured by other bones; telescoping of the
maxillaries has advanced to cover much of the
152
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 15. Neoteny in Orcaella. Note similarity in development of interparietal (arrow), dorsal extension of parietal
(open arrow), extensive exposure of frontals (weak telescoping of maxillary and premaxillary bones) in the
newborn Peponocephala, Feresa and Orcaella (A,B,C). In adult Peponocephala and Feresa (D,E) the dorsal
exposure of the interparietal is reduced or obliterated; the dorsal extensions of the parietals are similarly
overridden; the frontals become covered by the maxillaries and premaxillaries. In contrast, adult Orcaella (F)
retains juvenile features in the form of extensive dorsal exposure of the interparietal & dorsal extensions of
parietals, weak telescoping of skull. Peponocephala redrawn from Dawbin et al. (1970), Feresa from Perrin &
Hubbs (1969). Juvenile Orcaella is QMJM11343, a disarticulated skull which was re-assembled; positions of
the bones could be determined by their impressions on the braincase.
frontal bone. Similar change with growth is evi-
dent in Pseudorca (Cowley,1944, pl.1), Globi-
cephala melas (Fraser,1950, pl.2), Monodon
(Eales,1950, fig. 15; van Beneden & Gervais,
1868-1879, pl.45, fig. 1) and Phocoena phocoena
(van Beneden & Gervais,1868-1879, pl.43,
fig.5). In adult Orcaella, however, dorsal ex-
posure of the interparietal and parietals is main-
tained, and there is only weak telescoping (Figs
5,14,15; Marsh et al.,1989, fig.3) of the skull,
much as in the newborn or juvenile.
This discrepancy between Orcaella and other
genera can further be demonstrated by comparing
proportions of interparietal length and width in
Orcaella and Pseudorca of known age. Orcaella
maintains the proportional length and width of
the interparietal in the oldest animals examined
(Fig.16). Pseudorca, while having a greater dor-
sal exposure of the interparietal than most odon-
tocetes, shows a significant decrease in
interparietal length after two growth laminae and
width after 8-14 dentinal laminae (Purves & Pil-
leri, 1978, fig.16).
Nuchal and temporal crests are poorly defined
in Orcaella, which is general in juvenile skulls
(as in newborn and adult Pseudorca
(Cowley,1944, pl.1), Peponocephala (Dawbin et
al.,1970, fig.7) or Feresa (Perrin & Hubbs,1969,
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
fig.4)). Barnes (1985) noted these features as
paedomorphic in phocoenids.
Also evident from the growth series of odon-
tocetes is the progressive development of the
mesethmoid plate on the front of the braincase. In
adult Orcaella, the mesethmoid can completely
cover the anterior face of the braincase (Fig.14).
In Queensland material much of this area is left
exposed. The mesethmoid plate is poorly
developed in a specimen of Orcaella from Mel-
ville Bay, Northern Territory (Johnson, 1964,
pl.14). Two specimens in the Western Australian
Museum also have a poorly developed meseth-
moid plate (J.L. Bannister, pers. comm.). This
suggests that poor development of the meseth-
moid is another retained juvenile feature which
may be best expressed in Australian animals. If it
were a truly primitive character, one would not
expect to see any examples in which the meseth-
moid completely covered the front of the brain-
case.
Telescoping of the skull (Miller, 1923) is weak
in newborn odontocetes, with the ascending
processes of the maxillaries still far forward,
leaving a large amount of the frontals exposed. In
most genera, telescoping is increasingly ex-
pressed in older animals, but in Orcaella teles-
coping resembles that of juvenile odontocetes,
with the frontals widely exposed.
The short rostrum of Orcaella may be a
retained juvenile feature. According to Tomilin
(1967), the mean rostrum length as a proportion
of condylobasal length for adults and 'young' of
Grampus were 0.501, 0.457; Globicephala
0.507, 0.474; Pseudorca 0.481, 0.475; and Or-
cinus 0.500 (adult d), 0.471. The mean value for
proportion of rostrum length in Orcaella (0.444)
is thus closer to the values for juveniles of other
blunt-headed dolphins (although similarto values
for adults of some phocoenids).
Most features listed as neotenic relate to the
roof of the braincase. DeBeers (1937 in Pilleri et
al., 1982) contrasted the dermal bones of the skull
roof and the substitution bones, preformed in
cartilage, at the base of the skull. He further noted
that it was the dermal bones in which differentia-
tion and growth reflected the growth of the brain,
wwhile the bones at the skull base were inde-
pendent of brain growth. Thus neotenic features
should be most obvious in the dorsal aspect of the
skull, as documented here for Orcaella.
Lloze (1982) documented the os wurmiens
(‘small supernumerary bones...situated between
various bones of the cranium") in Orcaella; they
also occur on some Queensland specimens. Lloze
153
suggested that they may be space ‘fillers’, as-
sociated with the growth of the braincase. The
braincase is proportionately larger in Orcaella
than in most odontocetes (Lloze, 1982). An en-
larged braincase is generally recognised as a
neotenic feature (Gould, 1977) and we suggest it
is a retained juvenile feature in Orcaella.
Oeschlager (1986) noted a proportional
decrease in the mastoid portion of the squamosal
with age in Tursiops and Lagenorhynchus, which
suggests that the large postglenoid space in Or-
caella is a neotenic feature. We think this is
unlikely for two reasons. First, a proportional
decrease in the mastoid was not obvious in a small
series of Tursiops skulls available to us.
Moreover, the posteroventral orientation of the
paroccipital process of the exoccipital in Orcael-
la and the large basicranial space, as well as the
extent of the postglenoid space, exceeds that in
newborn odontocetes of other delphinoid genera
which we have examined, with the exception of
Neophocaena.
Deflexion of the occipital condyles (Fig.9),
otherwise known only in Neophocaena, may
reflect the enlargement of the braincase, but it
probably is not a retained juvenile feature as there
is no indication from growth series of Orcaella or
other genera that the condyles move from à
ventral to posterior position.
IMPLICATIONS OF NEOTENY. Orcaella's
skull is characterised by large braincase and short
rostrum as in Gray's (1866) diagnosis. If these are
retained juvenile features as we suggest, then it
may not be valid to make comparisons of Orcael-
la with adults of other genera for taxonomic
purposes. It is not surprising that published at-
tempts have been equivocal since juvenile skulls
of odontocete genera are more similar than are
skulls of adults.
de Muizon (1988) placed Orcaella in the
Globicephaliinae, on dilation of the premaxillae
at the tip of the rostrum. However, the premaxil-
lary width of Orcaella at 0.75 rostrum length is
comparable to that in delphinids (pers. obs.) and
is closer to that of juvenile Globicephala.
de Muizon (1988) and Barnes (1990) con-
sidered a well-developed mesethmoid plate to be
diagnostic features of delphinids. Although the
Orcaella mesethmoid plate can completely fill
the posterior border of the nares (Fig.14), it is
usually poorly developed in Australian
specimens, leaving much of the anterior slope of
the vertex exposed. de Muizon (1988) and Barnes
(1990) suggested that the greater development of
154
percentage condylobasal length
0 5 10 15 20
number of dentinal layers
FIG. 16. Variation in width (closed circles) and length (open circles) of interparie-
tal in Orcaella as percentage of condylobasal length, compared with age as
expressed by dentinal layers in teeth. The newborn animal QMJM11343 was
not aged, and is arbitrarily placed at 0.5 years. Note the minimal reduction in
both width and length of the interparietal.
the mesethmoid plate pushed the nasals to an
apical position on the vertex and resulted in
transverse compression of the nasal bones.
Despite the weaker development of the meseth-
moid plate in our material, the posterior pair of
nasal bones of Orcaella are apical, so position of
the nasals may not be linked to development of
the mesethmoid. The nasals are, however,
reduced in size, nodular, and not compressed.
They differ from the type of O. brevirostris (Fig.
14) and from O. fluminalis (Anderson, 1879, pl.
42, fig.2). van Beneden & Gervais (1868-1879,
pl. 64, fig.2a) illustrated 2 reduced nodular nasals
in Orcaella from the Mekong River; Lloze (un-
publ. MS) also noted that ‘the rudimentary nasal
bones are formed by two little bones which fuse
with advancing age’. Thus reduction of the nasals
may be a generic feature, although it seems par-
ticularly well shown by Queensland specimens.
Reduction of the nasals seems to occur in some
Cephalorhynchus hectori (van Beneden,1881)
(pers. obs.) and Perrin (pers. comm.) noted a
Stenella frontalis (Cuvier, 1829) with two nasals
on the right. Perhaps the nodules represent mul-
tiple sites of ossification (G.J.B. Ross pers.
comm.) which may remain discrete. If so, this
could be another neotenic feature.
MEMOIRS OF THE QUEENSLAND MUSEUM
The postnarial fossa, in-
filled by a supernumary
bone, and shallow lateral
fossae on the anterior slope
of the vertex have not been
described in Orcaella, al-
though the former was il-
lustrated by Johnson
(1964, pl. 14). This may be
because of the greater
development of the
mesethmoid in animals ex-
amined to date from
southeast Asia. Postnarial
fossae have been described
in Kentriodontidae, but
these appear to be different
from the present case,
based on de Muizon (1988,
fig.21). The lateral fossae
are similar to fossae of
phocoenids, Delphinap-
terus and Monodon (de
Muizon, 1988, fig.25).
These fossae in Orcaella
do not, however, provide
evidence for its close
relationship to phocoenids
or monodontids. Rather, they probably reflect the
weak mesethmoid plate in our material, leaving
much of the vertex exposed. In a juvenile Tur-
siops (QMJM4713), with the mesethmoid still
weakly developed, there are similar shallow im-
prints or fossae between the mesethmoid plate
and nasals.
25 30
BASICRANIAL CHARACTERS. Since the
basal skull bones are less subject to modification
by brain growth, perhaps they offer better char-
acters for assessing relationships of Orcaella.
Unfortunately, positions of the sphenoidal bones
are rarely evident from photographs and we have
been able to examine only a limited series of
specimens, usually without juveniles.
In ventral view (Figs 10-12), enclosure of the
palatine foramen within the triangular lateral
lobes of the palatine is characteristic. This ap-
pears to be the minor palatine foramen of Evans
& Christensen (1979). In sagittal section (Fig.33)
the palatine contains a highly vascularised ex-
cavation. In Tursiops (Rommel, 1990) and in
specimens of Sousa, Globicephala, Pseudorca,
Stenella, Delphinus, Peponocephala, Grampus,
and Feresa the minor palatine foramen is not
visible and there is only the major palatine
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
TABLE 1. Summary statistics of measurements, based
on 21 tympanoperiotic bones from 11 animals. Num-
bering of characters corresponds to sequence in Ap-
pendix 1, which contains full data and details of
measurements.
1)Standard length tympanic bone |34.82 33.05-36.5
(2)Tip to posterior end of inner [95.42 93.15-
posterior prominence 99.71
(3)Distance from posteroventral 58.07-
66.26
tip of outer posterior prominence
to tip of sigmoid process
53.94-60.98
(4)Distance from posteroventral
tip of outer posterior prominence
to conical process
G)Widin bulla at level of
(7)Width across inner and outer 53.94-62.61
posterior prominences
22.52-36.19
5.28-8.89
(8)Depth interprominential notch
(10) Width posterior branch of
lower tympanic aperture
31.00-38.20
31.97-41.38
(13)Standard length periotic bone
(14) Thickness superior process
at level of upper tympanic
aperture (excluding spongy bone
shelf over aequeductus vestibuli)
(15)Width across cochlear
rtion and superior process at
evel of tym pert
(19)Length of cochlear portion of
periotic
(22)Standard length as
percentage of standard length of
tympanic bone
38.95-46.62
(23)separation of foramen 9.08-19.01
singulare and aequeductus
Fallopii (n=12)
(25) Anteropisterior length of
undus of the internal auditory
meatus
(26)Width of foramen ovale (n=9) | 5.55 3.74-7.00
(27) Width of foramen rotundum 8.85 7.54-10.29
(28) Width of head malleus (n=9) |10.86 8.53-12.02
foramen on the suture of the palatine lobe with
the maxillaries. However, both foramina may be
present in delphinids (Perrin pers. comm.) and
their development may be a variable feature.
The lateral and medial lobes of the palatines are
widely separated and the maxillary can directly
contact the pterygoid hamuli, totally separating
the medial and lateral lobes of the palatines
(Fig.10). de Muizon (in press) notes that separa-
tion of medial and lateral lobes of the palatines is
unusual among odontocetes. The separation of
the pterygoids in Orcaella is by the medial lobes
of the palatines, which surround and may over-
ride the vomer so that it is no longer visible from
Ac
FIG. 17. Diagram of periotic of Orcaella in dorsal view
showing features of the periotic triangle. Abbrevia-
tions: cF=canal for facial nerve (=aquaeductus Fal-
lopi); Av=aquaeductus vestibuli (=ductus
endolymphaticus); Ac=aquaeductus cochleae; alpha,
beta and gamma indicate angles as defined by Pilleri
et al., 1989.
below (but see Fig. 12). This resembles the con-
struction in delphinids such as Sousa, Tursiops
and Stenella, and is distinct from other odon-
tocetes with widely separated pterygoids (Pilleri
et al., 1982). In phocoenids and Delphinapterus,
the pterygoid hamuli are relatively small and do
not deeply excavate the palatine, so the connec-
tion between median and lateral lobes is broad. In
the long-snouted dolphins, although the
pterygoids deeply excavate the palatines the latter
are displaced forward onto the rostrum so that
medial and lateral lobes of the palatine remain
connected. Partial to complete separation of the
lobes can occur in Grampus (Tomilin, 1967) and
Feresa (QMJMS825). Perrin (pers. comm.) noted
complete separation of the medial and lateral
lobes in a juvenile Globicephala macrorhynchus;
the lobes are also separate in a Southern Hemi-
sphere G. melas (QMJ15.2104). This separation
may result from deep excavation of the
pterygoids abutting onto a shortened rostrum of
the skull, but it is not a simple relationship.
Monodon (Tomilin, 1967; pers. obs.) and the
delphinid Pseudorca (adult and newborn:
Cowley, 1944) have well developed pterygoid
hamuli and a short rostrum but the median and
lateral lobes of the palatines remain broadly con-
nected.
156
TABLE 2. Angles of the periotic triangle, measured
from camera lucida drawings. Abbreviations: PA,
AW: different observers; L=left; R=right; #1, #2:
measurements by same observer.
SPECIMEN NUMBER a |B
QMJM4706 (AW) |88 |62
QMJM4706 R(AW) |82 |63 [35 f
40.5
QMJM4705 | L(AW) |86
R(AW)
L(AW)
R(AW)
R(PA)
R(AW)
L(AW)
QMJM4705
QMJM11343
QMJM 11343
QMJM4708
QMJM4709
QMJM4740
Orcaella is unusual in its separation from the rest
of the braincase. A posterior extension of the
palatine is found in Globicephala, Feresa, and
Pseudorca (Purves & Pilleri, 1978, fig. 18) where
it forms a bridge under the optic canal. However,
in those genera, the palatine wing is oriented
vertically (in adults examined), whereas in Or-
caella it is parallel with the bones of the
basicranium, as a free wing-like structure, which
appears to be flattened against the bones of the
skull. The basicranium also appears to be flat-
tened. The relatively long paroccipital crest, al-
most at right angles to the main axis of the skull,
creates a very large basicranial space, which may
cause a more lateral (flattened) orientation of the
bones of the skull base. For instance, the fal-
ciform process of Orcaella is flattened against the
braincase. This differs from the delphinids ex-
amined (Tursiops, Sousa, Globicephala, Pseudo-
rea, Stenella, Delphinus, Peponocephala,
Grampus, and Feresa) which all have a strong
ventral deflexion of the falciform process, curv-
ing around the anterior of the periotic bone.
An extensive cavity between the maxillaries
and frontal/presphenoid can accommodate a dor-
sal extension of the preorbital lobe. This feature
resembles the extension in phocoenids (Fraser &
Purves, 1962). However, there is a similar expan-
sion of the lobe in delphinids such as Pseudorca
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 3. Lengths of sides of the periotic triangle,
measured from camera lucida drawings.
WAV-
SPECIMEN NUMBER Av-cF | Av-Ac | Ac/Av-
cF
[ QuuMaz9 | (aw) | so | 40 | 678 |
| Quo | | so | 36 | 720
| 39 |
59
50
39
48
54
QMJM4708 | (AW) 39 | 1000
QMIM4704 | (PA) | 48 | 44 | 917
QMJM11343 | (aw) | 60 | 4 66.7
0
QMJM11343 37
QMJM4705
QMJM4740
QMJM4721
(pers. obs.) and juvenile Globicephala macro-
rhynchus (Perrin pers. comm.).
The pterygoid hamuli have medial flanges
(Figs 10-12) which can almost contact distally
although there usually is a distinct gap. The flan-
ges are also visible in a specimen from the North-
ern Territory (Johnson, 1964, pl.15). This
contrasts with the very widely separated
pterygoid hamuli, without flanges, in SE Asian
specimens (Owen, 1866; Anderson, 1879; Pilleri
& Gihr, 1973-1974).
The mastoid portion of the squamosal is well-
developed both ventrally and laterally. Ventrally,
there is a mastoid pad for attachment of the tym-
panoperiotic bones (Fig. 11; Owen, 1866, pl. 9,
fig. 3; Anderson, 1879, pl. 42, fig. 3) and a con-
spicuous groove for the external auditory meatus.
Laterally, the large lateral rugose mastoid portion
of the zygomatic arch may reflect a broad attach-
ment base for the sternomastoid muscles, possib-
ly associated with flexibility of the head. The
zygomatic arch of the squamosal is thus well-
developed, and intermediate between Monodon
and Delphinapterus on the one hand and
phocoenids & delphinids on the other. Thus, con-
trary to Heyning (1989), Orcaella does not show
extreme reduction of the zygomatic arch. This
reduction of the arch was the only synapomorphy
listed by Heyning (1989) for the clade
Phocoenidae-Delphinidae. The taxonomic sig-
nificance of the well-developed arch in Orcaella
is considered as part of a cladistic analysis of the
delphinoid taxa, but note that a relatively well-
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
157
FIG. 18. Lateral view of left tympanoperiotic bones, QMJM4709. Note anterior spine of bulla; posterolateral
orientation of posterior processes of bulla and periotic (directed towards viewer); triangular depression in front
of sigmoid process (open arrow); posterior branch of lower tympanic aperture (triangle); concave ventral margin
of bulla, and tubercle on anterior process of periotic (arrow).
developed arch also occurs in Neophocaena,
within the Phocoenidae (pers. obs.) and in the
delphinid Orcinus (Heyning, 1989).
The deep, elongate pit between the squamosal
and parietal (Fig.11) has been noted by de
Muizon (in press) in ‘some delphinids’; it occurs
in Sousa chinensis and in several delphinids (Per-
rin pers. comm.). It appears to be the channel for
a blood vessel which exits on the back of the
braincase, between the squamosal and parietal.
CONCLUSIONS. Neotenic features of the skull
roof in Orcaella include the retained dorsal ex-
posure of the interparietal and dorsal extensions
of the parietals, poorly developed telescoping of
the skull with large exposure of the frontal bones,
and weak development of the mesethmoid plate
on the anterior of the cranium (the last character
possibly restricted to Australian animals). The
relatively large braincase and short rostrum,
which substantially affect the appearance of the
skull, may also be neotenic features. Retention of
juvenile features in Orcaella may compromise
attempts to establish relationships based on the
comparisons of its skull with adults of other
genera; it would be more apppropriate to compare
it with juveniles of other odontocetes. However,
juvenile skulls of various genera are very similar
with diagnostic features often only well ex-
pressed in adults.
The basicranium may be less subject to
neoteny. The skull base of Orcaella resembles
that of Pseudorca, Globicephala, Feresa and
Grampus (we have not been able to examine in
detail skulls of Orcinus). In particular, there is a
similar excavation of the palatines by the
pterygoid hamuli, which can lead to separation of
the medial and lateral lobes of the palatine (shared
with Feresa, Globicephala and Grampus);
posterior extension of the lateral lobe of the
palatine (shared with Globicephala,Feresa,
Pseudorca) and a transverse widening of the skull
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 19. Ventral view of left bulla, QMJM4709. Note
swollen outer posterior prominence (open arrow);
deep interprominential notch (arrow); spongy bone
along midline of bulla; posterolateral orientation of
and spongy bone distally on posterior process.
(as in Pseudorca, Globicephala). However ex-
cavation of the palatines and orientation of the
posterior wings of the lateral palatines may be
more similar to juveniles of genera such as
Globicephala than to adults (i.e. may be
neotenous). The cranial sinuses also align Or-
caella with the bulbous-snouted dolphins (Fraser
& Purves, 1962). However, in all these genera the
cranial sinuses show few specializations; they
may be linked by shared primitive features. A
prominent feature is the well-developed
zygomatic arch of the squamosal and ventral
attachment of the tympanoperiotic bones which
sets Orcaella apart from delphinids although the
arch is similarly developed in Neophocaena.
Taxonomic utility of basicranial features needs
to be examined further. It has generally proved
impossible to demonstrate whether a feature is
neotenic or in a generalised, unspecialised state.
FIG. 20. Ventral view of left bulla, QMJM4700. Note
angulate inner posterior prominence (triangle); deep
interprominential notch; posterolateral orientation of
posterior process (arrow).
One option is to compare a wider range of
juvenile delphinid genera than was available to
us in this study. Another option is to assess
polarity of basicranial features, based on out-
group analyses involving criteria other than
cranial morphology. We present such analyses
later in this paper.
TYMPANOPERIOTIC BONES
(Table 1; Fig.17; Appendix 1)
TYMPANIC BULLA. Standard length 34.8mm
(33.0-36.5). The anterior tip can be drawn out into
a spine-like process (Figs 18, 30). The outer
posterior prominence is swollen and hemispheri-
cal, with no lateral compression (Fig. 19). The
inner posterior prominence is sharply angulate
(Figs 20, 21); a weak ventral keel extends more
than half way to the tip (Fig 20). The inter-
prominential notch is deep (Figs 19,20). There is
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
FIG, 21. Dorsal view of left bulla, QMJM4700. Note
angulate inner posterior prominence (triangle);
prominent accessory ossicle (arrow); ridges and
grooves of articular facet (open arrow) which is
directed posteriorly (associated with posterior orien-
tation of periotic process).
no median furrow. A spongy, weakly developed
ridge extends along the anteromedial border of
the interprominential notch, continuing forward
to the tip along the midline (Fig. 19). There is no
compression: the width of bulla at the level of the
sigmoid process is 57.8 (53.9-61.0)% of standard
length. The posterior process is well developed,
variable in direction from almost posterior to
distinctly posterolateral. The mean angle of the
lateral margin is 34.9° (17°-47.5° to long axis of
bulla) (Figs 18-20). Although the articular facet
with the periotic can be more posteriorly oriented
(Fig. 21), the posterior process is usually oblique-
ly oriented laterally (Figs 19,26). Distally the
posterior process consists of spongy bone (Fig.
19). The facet with the posterior process of the
periotic is ridged and grooved (Fig. 21).
159
In lateral view, the ventral margin is slightly
concave (Fig. 18). A deep triangular area anterior
to the sigmoid process is bounded on its ventral
side by a low, wide ridge which gives rise on its
anterior margin to a shallow vertical groove (Fig.
18). The accessory ossicle is prominent (Fig. 21).
In medial view, the border of the involucrum is
low, not rising anteriorly to form a distinctly
curved margin (Fig. 22). The elliptical foramen is
closed, but minute perforations are often present
in the area usually occupied by the foramen (Fig.
23).
PERIOTIC BONE
DORSAL ASPECT. Standard length 35.7mm
(31.0-38.2). Anterior, superior and posterior
processes are in a straight line (Fig. 24). The
superior process can be directed medially as a
shelf of spongy bone, partially hiding the tractus
spiralis foraminosus and completely obscuring
the internal aperture of the aquaeductus Fallopi
(canal for facial nerve) (Fig. 24). A similar
medially directed shelf of spongy bone can cover
the aperture of the aquaeductus cochleae and
aquaeductus vestibuli (ductus endolymphaticus)
(Fig. 25). The aquaeductus vestibuli is set at the
base of a large funnel-shaped depression bounded
by spongy bone (Figs 24,25). The tractus spiralis
foraminosus is prominent (Fig. 25). The
aquaeductus Fallopi is directed anteriorly at the
level of the anteriomost margin of the tractus
spiralis foraminosus (Fig. 25). The foramen sin-
gulare is at the posterior margin of the tractus
spiralis foraminosus and separated from the
aquaeductus Fallopi by a long, obliquely oriented
crista transversa which has a secondary ridge
continuing along the anterior margin of the fun-
dus of the internal auditory meatus.
The posterior process forms an acute back-
wardly pointing triangle in dorsal view (Figs
24,26; Tables 2,3). At its maximum extent it
covers only about 0.75 of the posterior process of
the tympanic bulla, and tapers to a tip on the
posteromedial side of the process of the bulla
(Fig. 26).
LATERAL ASPECT. In external view, the
anterior process is square and truncate, directed
ventrally where it fuses with the bulla between
the accessory ossicle and the sigmoid process
(Figs 18,27). A distinct tubercle occurs on the
upper half of its anterior face, near the level of the
anterior margin of the cochlear portion of the
periotic (Figs 18,25).
160
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 22. Medial view of left bulla, QMJM4700. Note low medial margin of involucrum, not raised anteriorly to
form sinusoidal border.
The posterior process bends sharply ventrad, to
fuse with the posterior process of the tympanic
bulla (Fig. 27). The distal half of the external
surface of the posterior process, where it articu-
lates with the squamosal, is spongy bone (Figs
24,26).
In medial view the periotic is oriented obliquely
ventrad and is closely apposed to the involucrum
of the bulla, although never touching (Fig. 25).
The apertures of the aquaeductus vestibuli and
aquaeductus cochleae are directed medially, that
of the aquaeductus Fallopi anteromedially. The
groove for the stapedius muscle is prominent
(Fig. 28).
VENTRAL ASPECT. The greatest diameter of
the cochlear portion is at right angles to the main
axis of the periotic (Fig. 28). On the postero-
medial quadrat of the cochlea is an oblique
groove which runs obliquely anterior (Fig. 28). It
occurs where the periotic is closely apposed to the
bulla and may be a channel for a blood vessel
(possibly associated with corpus cavernosum;
note vascularisation of the region in Fig. 33). The
anterior-posterior length of the articular facet (1n-
cluding the basal smooth portion (Fig. 28) and
distal spongy portion) is 47.7-54.1% of the
periotic length(n=5).
ARTICULATION OF PERIOTIC AND TYM-
PANIC BONES. In posterior view, the articula-
tion of the posterior process of the periotic with
that of the tympanic is irregular (Fig. 23). The
facet for attachment with the mastoid portion of
the squamosal is entirely spongy bone (Figs
29,30). Anteriorly, the articulation is between a
triangular wedge of the periotic and the
squamosal. Posteriorly, the connection is be-
tween the the squamosal and the posterior half of
the posterior process of the tympanic bone (Figs
23,26).
DISCUSSION
COMPARISONS OF VALUES WITH
KASUYA (1973). The morphometrics of this
series (Table 1, Appendix 1) generally confirm
values presented by Kasuya, based on specimens
from unspecified localities held at the United
States National Museum and Zoological Survey
of India. The bulla is wider across the posterior
processes in Queensland specimens. There is a
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA 161
FIG. 23. Posterior view of left tympanoperiotic bones,
QMJM4709. Note irregular margin of posterior
processes of bulla and periotic; closed elliptical
foramen, with a few pores visible (triangle). The open
arrow and closed arrow indicate spongy bone of bulla
and periotic respectively, which articulates with the
mastoid portion of the zygomatic arch.
major discrepancy in the depth of the inter-
prominential notch (mean=30.3 in present series
vs 11.7 reported by Kasuya). This must involve a
difference in measuring technique but the inter-
prominential notch is deep in our specimens (Figs
19, 20). Our value for height of the bulla, from
sigmoid process to ventral keel, is higher than
reported by Kasuya (1973). His measurement
was taken obliquely and incorporated not only
height but also width of the bulla. Since the latter
is wider in our material than reported by Kasuya,
our values for ‘height’ should also be greater.
TAXONOMIC CHARACTERS OF THE TYM-
PANOPERIOTIC BONES. Characters used by
Kasuya to differentiate the families of delphinoid
odontocetes (Table 4) are considered for Orcael-
la: (1) direction of posterior process of bulla. This
is not usually posterior as in Delphinapterus and
the phocoenids, but variable from almost
posterior to posterolateral (Figs 19,20), with a
mean angle along the lateral margin of ap-
proximately 35°. Although the apex of the
process is directed posteriorly, the lateral margin
is always deflected and the articular facet is usual-
ly distinctly posterolateral (Fig. 19). The mean
angle of deflection of the posterior process is
FIG. 24. Dorsal view of left periotic, QMJM4700. Note
that anterior (arrow), superior and posterior processes
are in line and that the latter is posteriorly directed.
Note spongy bone shelf overarching the tractus
spiralis foraminosus (open arrow), and wide funnel-
like depression, surrounded by spongy bone, which
contains the aquaeductus vestibuli (triangle).
comparable to the deflection in delphinids such
as Sousa chinensis (e.g. 33^, 36.5? in MM1020).
The greater variability in the orientation of the
posterior process in our series of Orcaella brings
into question its usefulness as a character to
separate families. As such, it weakens the case for
linking Orcaella and Delphinapterus in the Del-
phinapteridae.
(2) direction of posterior process of periotic.
This is directed posteriorly, but the posterior
orientation is found not only in Delphinapterus
but also in phocoenids.
(3) width of posterior branch of lower aperture
of tympanic membrane (LTA). Even with our
larger series of specimens, there is no overlap in
162
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 4. Tympanoperiotic characters used by Kasuya (1973) to classify odontocetes. Families are as given
by Kasuya (1973). Unless otherwise stated, data for Orcaella are from this study; values for other species are
from Kasuya (1973).
direction of
posterior process
tympanic bulla
direction of
posterior process
of periotic
Monodontidae
Monodon
lateral
width of posterior
branch of lower
tympanic aperture
facet of posterior
processes of bulla
d periotic
compression of
bulla
ventral keel
interbromine ntial
notch depth
ridged (this paper)
strongly
compressed
39.6 (37.5-41.4)
well developed
$$$}
Delphinidae
sterolateral to
ateral
sterolateral to
ateral
Delphinapteridae
Delphinapterus Orcaella
Phocoenidae
variable, mean =
35? (17°-48°)(this
posterior
paper)
posterior
+
posterior
posterior
means from 2.5
(Stenella
attennuata) to 7.2
(Tursiops cf gilli)
ridged (Yamada in
Kasuya 1973)
strongly —
compressed in
Globicephala &
Grampus
well developed in
Globicephalinae
(sensu Kasuya)
mean =10.0
(Globicephala) to
16.6 (Lissodelphis)
anterior spine of
tympanic bulla
medial margin of
involucrum
strongly
sinusoidal, high
anteriorl
variable, present
in Globicephalinae
sinusoidal, esp in
Globicephalinae
+ —R——— ——
nin
hininae,closed
lobicephala
Grampus, Orcinus,
Sousa, Sotalia
Cephalorhynchus
o
Del
in
elliptical foramen
ranges of values between Orcaella (LTA: 5.3-
8.9%) and Delphinapterus (LTA: 10.1-11.8%:
Kasuya, 1973). This character thus offers no sup-
port for linking Orcaella and Delphinapterus in
the Delphinapteridae. Rather, the values for Or-
caella overlap with both Monodon (mean = 7.596:
Kasuya, 1973: Appendix 2) and at least some of
the delphinids, such as Tursiops (2.2-9.2%:
Kasuya, 1973) and Sousa (7.3% : Kasuya, 1973).
The LTA is wide in Delphinapterus and
phocoenids where the posterior process of the
tympanic is oriented posteriorly. If there is a
lateral shift of the process, this must impinge on
the LTA which is just anterior to the process.
Thus one could expect a narrow LTA in taxa
where the posterior process has shifted laterally;
Neophocaena
10.3; Phocoena |mean = 6.5
7.5, 10.6, 10.3; Gi - 8.9)
Phocoenoides this paper)
8.9,10.5
smooth ridged (this paper)
not compressed
mean = 57.8
(53.9-61.0)
not compressed
mean = 58.9 not compressed
(58.0-59.0)
low (in Phocoena)
to well developed
(in Neophocaena,
Phocoenoides)
Neophocaena
mean=14.1;
Phocoena
mean=14.6, 16.2
Phocoenoides
mean=17.0, 17.6
mean=11.7
mean=15.1 (Kasuya)
may be present
absent absent | (thís aper)
:
low, most anterior
height in
Neophocaena
sinusoidal low throughout
closed closed
this is as observed. Characters (1) and (3) may
thus be linked.
(4) articular facets of posterior processes. These
are ridged in Orcaella but this is also true in
delphinids and Monodon, as well as Delphinap-
terus.
(5) compression of bulla. When the
width/length ratio was 0.5 or less , Kasuya (1973,
fig.75) considered the bulla to show strong lateral
compression. Within the Delphinidae of Kasuya,
there are genera (e.g. Globicephala, Grampus)
which showed strong compression, while others
showed no compression of the bulla. Thus the
strong compression of the bulla in Monodon but
not in Delphinapterus can not be used to separate
them at family level.
(6) development of ventral keel. Kasuya
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
FIG. 25. Medial view of left tympanoperiotic bones,
QMJM4709. Note spongy bone over cF (arrow) and
Av (open arrow); position of Ac (triangles); large
tractus spiralis foraminosus; tubercle on anterior
process of the periotic; close association of cochlear
portion of the periotic with medial porion of the bulla.
(1973:54) noted that the development of the
ventral keel and the median furrow could be
linked with the extent of compression of the bulla.
This may also apply to the width of the inter-
prominential notch. To the extent that these char-
acters are linked to compression of the bulla, they
can not be considered valid family characters.
(7) medial margin of involucrum. The margin
is low throughout in Orcaella, which contrasts
with the sinusoidal form of Monodon, Delphinap-
terus and most delphinids.
The other characters in Table 4 either vary
widely within a single family (e.g. closure of
elliptical foramen) or are found among several
families (anterior spine of bulla) and thus are of
limited use in defining families.
PERIOTIC TRIANGLE. Pilleri et al. (1989)
defined the periotic triangle, which is formed by
connecting the apertures of the canal for the facial
nerve (aquaeductus Fallopi), aquaeductus ves-
tibuli and aquaeductus cochleae. They recog-
nized 6 types, including a monodontid triangle in
FIG. 26. Dorsal view of left tympanoperiotic bones,
QMJM4709. Note posterior orientation of posterior
process of periotic; posterolateral orientation of
posterior process of tympanic and expanse of spongy
bone (arrow) which articulates with the zygomatic
arch .
163
Monodon, Delphinapterus and
Orcaella. The monodontid tri-
angle was defined by (1) apertures
of aquaeductus cochleae and
aquaeductus vestibuli ‘roughly at’
the same level, hence angle alpha
almost a right angle triangle; (2)
distance between the two aper-
tures (AvAc) roughly equal to that
between the aquaeductus ves-
tibuli and canal for facial nerve
(AvcF); and (3) angles beta and
gamma approximately equal (i.e.
45°).
Considering each character of
the monodontid triangle:
(1) alpha almost a right angle
This was only approximately so
for Delphinapterus (108°: Pilleri
et al., 1989), but more so for Monodon
(99°,90°,90° Pilleri et al., 1989 ) and for Orcaella
(mean=85.4° (72°-102°); Pilleri et al., 1989:
164 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 27. Lateral view of left periotic, QMJM4700. Note squared off form of anterior process (to the left); ventral
deflexion of posterior process to form triangular wedge resting on top of posterior process of tympanic.
86°,84°). However the ‘delphinid triangle’ is ap-
proximately a right angle; the mean value of
alpha from Pilleri et al. (1989) for Monodon and
Delphinapterus was 96.8° vs 97.1° for delphinids.
(2) AvAc approximately equal to AvcF, thus
AvAc/AvcF approximately equal to 1.0. We had
to calculate these figures for Delphinapterus and
Monodon from Pilleri et al. (1989, fig.34). The
line from AvAc was equal to that from AvcF in
Delphinapterus and was about 0.8 of the length
of AvcF in Monodon. Based on 15 measurements
in our series, the line AvAc was about 0.70
(mean=0.73) the length of AvcF, which cannot be
considered ‘approximately equal’. In our study,
there is considerable variation between ob-
servers, with the means for two sets of observa-
tions as 0.68 (n=10) and 0.82 (n=5); moreover,
the range is large (from 0.54-1.00 in the first
series; 0.70-0.92 in the second). In defining the
‘delphinid’ triangle, Pilleri et al. (1989) stated
that the line Ac-Av is ‘less’ than Av-cF. Given
the unclear boundaries contrasting this statistic
for the ‘monodontid’ and ‘delphinid’ triangle,
FIG. 28. Ventral view of left periotic, QMJM4700.
Note strong ventral deflexion of anterior process
(upper right); great transverse width of cochlear por-
tion; oblique groove on postero-medial border of
cochlear portion (open arrow); fenestra ovalis (tri-
angle); prominent groove for stapedial muscle, run-
ning obliquely below fenestra ovalis; ridged, compact
basal portion of posterior process (arrow).
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
FIG. 29. Oblique ventral view of skull, QMJM4720,
incompletely cleaned. Connective tissue (broad
arrow) holds tympanoperiotic bones in original,
ventral position against mastoid pad (open arrow) of
zygomatic arch. Also note falciform process of the
zygomatic arch (triangle).
and the extensive variability within Orcaella, the
ratio AvAc/AvcF must be interpreted with cau-
tion.
(3) beta and gamma roughly equal; i.e. gamma
/beta about 1.0. In measuring periotics of 9
animals, variability and observer bias were exten-
sive. Our mean values (beta 58.3 (47—70); gamma
36.3 (30-54) correspond closely to those of Pilleri
et al. (1989): beta 61,62; gamma 33,34. The mean
value for beta was much larger than that given by
Pilleri et al. (1989) for monodontids (mean=49.5)
and delphinids (mean=54.7). The value for
gamma was intermediate between that for
monodontids (41.9) and delphinids (28.2) (Pilleri
et al., 1989, table 16). Given the wide overlap in
values (e.g. gamma: Orcaella 30-54, Delphinap-
terus and Monodon 32-48, delphinids 18-38) the
165
FIG. 30. Oblique ventral view of skull, QMJM4714,
with tympanoperiotic bones re-attached against mas-
toid pad (broad arrow) of zygomatic arch. Note
anterior spine of bulla (small arrow); ventral
deflexion of paroccipital process of exoccipital, with
shallow depression on anterior face (open arrow), and
falciform process flattened against the skull base.
data provide equivocal support for a relationship
between Orcaella and monodontids.
The mean of gamma/beta for Delphinapterus
and Monodon is 1.01 (Pilleri et al. (1989)), but
our mean for Orcaella is 0.64 (n=16). The ratios
from Pilleri et al. (1989) are even lower (0.54,
0.55). Again, there is considerable variation
within and between observers in our specimens
(Ist series mean=0.59 (0.43-1.00, nz11); 2nd
series mean=0.78 (0.60—0.92, n=5)). However
neither our series nor that of Pilleri et al. (1989)
can be considered ‘approximately the same’.
Variability in monodontid triangle values seem
extensive even for Delphinapterus and Monodon,
but our figures and those of Pilleri et al. (1989)
for Orcaella correspond only poorly to the defini-
tion of the monodontid triangle. The closest cor-
respondence is in the angle alpha (mean=85.4° for
Orcaella, which approximates 90°). However the
delphinid triangle is also approximately 90°.
166
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 31. Near sagittal section of head, MM334, a 1.86m 9 . Note moderate development of melon (open arrow),
prominent rostral muscles (triangle), and extent of connective tissue (arrow) at the front of the upper jaw.
Evidence from the so-called monodontid triangle
that Orcaella is closely related to either Del-
phinapterus or Monodon is equivocal at best.
ADDITIONAL CHARACTERS OF PERIOTIC
BONES. Details of the tractus spiralis
foraminosus, such as a prominent crista transver-
sa, resemble Neophocaena and Pontoporia (Per-
rin pers. comm.). The groove in the periotic,
which may be the impression of a blood vessel
(Fig. 33), has not been described in other odon-
tocetes, and thus can not be used as a taxonomic-
character.
ATTACHMENT OF TYMPANOPERIOTIC
TO SQUAMOSAL. Kasuya (1973) and Kleinen-
berg et al. (1969) noted that the periotic of the
beluga was firmly sutured to the squamosal; this
was considered characteristic of Delphinapterus
and Monodon by Tomilin (1967). Heyning
(1989) did not find the periotic sutured to the
squamosal in beluga he examined nor is it sutured
in beluga and narwhal examined by one of us
(PA). Kasuya (1973) described how the dorsal
surface of the periotic is grooved to articulate
with the squamosal. We could not confirm the
type of articulation in beluga and it is not evident
in published photographs where the tym-
panoperiotics seem to have been re-attached in a
variety of orientations. In a juvenile narwhal
(UBC9467) the periotic is attached to the
squamosal by a thin wedge of the posterior
process bounded ventrally by the posterior
process of the tympanic. In narwhal and beluga,
the main attachment seems to be by the periotic,
while in Orcaella the attachment is shared be-
tween the tympanic and the periotic. The attach-
ment in all three genera appears to be by
connective tissue on a mastoid pad on the ventral
side of the zygomatic arch within a large post-
glenoid space (Figs 29,30); this contrasts with
phocoenids and delphinids where the attachment
is in a cavity formed by the squamosal, exoccipi-
tal and basioccipital (Kasuya 1973; pers. obs.).
Ventral attachment of the tympanoperiotics of-
fers the most convincing evidence that Orcaella
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
167
FIG. 32. Opposing section to that in Fig. 31, showing development of melon (open arrow), connective tissue and
rostral muscles. Also note partial section of the inferior nares (arrow).
may be related to Delphinapterus but the feature
is shared with Monodon (placed in a separate
family by Kasuya) and may be a primitive fea-
ture.
CONCLUSIONS. 1. Orientation of the posterior
process of the tympanic bulla is variable in Or-
caella, but the lateral margin is usually deflected
approximately 35? from the main axis of the
bulla. The deflection of the process is more com-
parable to delphinids such as Sousa than to Del-
phinapterus. A larger series of measurements for
the posterior branch of the lower aperture of the
tympanic membrane shows that, contrary to
Kasuya (1973), itis within the range of delphinids
rather than that of beluga. These findings weaken
arguments for beluga and Orcaella being
cofamilial. The other characters (Table 4) provide
equivocal evidence for the Delphinapteridae.
2. Linking Monodon, Delphinapterus and Or-
caella in one family is inconsistent. Including
Monodon and Delphinapterus in one family im-
plicitly rejects the orientation of the posterior
processes of the bulla and periotic, the compres-
sion of the bulla and the width of the posterior
branch of the lower tympanic aperture (LTA) as
family characters, since they differ between the
two genera. However, orientation of the posterior
processes and width of the LTA were the main
characters used by Kasuya (1973) to unite Or-
caella and Delphinapterus in the Delphinap-
teridae.
3. Our data and that of Pilleri et al. (1989) on
the periotic triangle provide either no support or
only equivocal support for linking Orcaella with
either Delphinapterus or Monodon.
4. Tympanoperiotics of Orcaella are attached
by connective tissue to a prominent mastoid pad
on the ventral surface of the zygomatic process.
A similar ventral connection occurs in Del-
phinapterus and Monodon, although there ap-
pears to be a greater involvement of the posterior
process of the tympanic bulla in the articulation
of Orcaella. In phocoenids and delphinids the
attachment of the posterior processes is within a
cavity formed by the squamosal, exoccipital and
basioccipital bones.
168
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 33. Section approximately 2 cm left of Fig 32. Note well developed rostral muscles; blowhole ligament with
cartilaginous inclusion (small arrow); diagonal membrane (open arrow); vascularised excavation of palatine
bones (triangle); tympanoperiotic bones in cross-section (broad arrow) bordered by acoustic fat anteriorly and
peribullary sinuses dorsally and posteriorly.
FACIAL ANATOMY
EXTENT OF MELON. MM333 from Mackay
has a gape of 130 mm. The anterior boundary of
the melon starts about 30 mm behind the tip of
the upper jaw; the anterior 30 mm (or c.23% of
gape length) is blubber and dense connective
tissue. The melon is bordered ventrally by well-
developed rostral muscles. The near sagittal sec-
tions of MM334 from Ellis Beach (Figs 31-33)
show the well-developed rostral muscles and the
limited extent of the melon.
Mead (1975) demonstrated that the superficial-
ly similar ‘bulbous-headed’ dolphins are
anatomically quite distinct. For instance, Gram-
pus has an extensive melon while Pseudorca has
a preponderance of dense connective tissue in the
forehead. Orcaella is closest to the generalized
delphinid, with neither melon nor connective tis-
sue developed to a high degree. It resembles
genera such as Lagenorhynchus or even Tursiops
as much as it does any of the bulbous headed
species.
The bulbous shape of the forehead is one of the
reasons for the supposed relationship of Del-
phinapterus and Orcaella. Anatomy of the
forehead in these two genera is distinct, however,
except perhaps for the well-developed rostral
musculature (Figs 31-33 for Orcaella; Heyning
(1989:33) for Delphinapterus). There is a con-
spicuous melon in the beluga, which runs to the
front of the forehead (Pilleri et al., 1980, fig.13).
The profile of the beluga's forehead is quite mal-
leable and soft. Changes in the forehead are well
documented for this species. In contrast, the
anterior forehead of Orcaella is exclusively
dense connective tissue, and it was not possible
to deform the profile of the forehead in specimens
we examined. Although the 'facial expression' is
*changeable' (Martin,1990), the profile of the
head did not vary on captive Orcaella observed
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
FIG. 34. Oblique dorsal view of section in Fig 32,
partially dissected to reveal right vestibular sac (open
arrow), with folded walls. Also note lip of the nasal
plug (arrow).
at the Jaya Ancol Oceanarium (Dr. A. Preen pers.
comm.).
Thus the resemblance between Orcaella and
Delphinapterus appears superficial. There is at
least as great a similarity between Orcaella and
the phocoenid Neophocaena, which suggests
head shape is a shared primitive character.
RESPIRATORY TRACT. The blowhole forms a
crescent, with the horns directed anteriorly; its
width is 1.5-2.2% of the standard body length.
The blowhole is displaced towards the left in
MM333, 334.
The spiracular cavity continues ventrally from
the blowhole as a transverse slit, which is sur-
rounded by dense connective tissue anterior to the
vertex of the skull.
The vestibular sacs extend laterally to
posterolaterally from the spiracular cavity. In
MM333 they appear to be collapsed and cover
less than half the area of the vestibular sacs in the
larger MM334. In both specimens, however, the
right and left vestibular sacs are approximately
169
FIG. 35. Enlarged view of section in Fig 31. Note
muscles around nasal sacs; the darkly pigmented dor-
sal vestibular sac (small arrow); elongate tubular
nasofrontal sac (open arrow); blowhole ligament with
cartilage (broad arrow); nasal plug with prominent lip
(below blowhole ligament) entering the inferior ves-
tibule; extensive premaxillary sac (triangle).
equal in size. They are lined with a darkly pig-
mented epithelium and the walls have slight con-
centric folds in MM334 (Fig.34) and MM335.
The vestibular sacs are connected to the
spiracular cavity by a medial ventral slit.
The nasofrontal sacs are immediately below the
vestibular sacs. The anterior portion of the
nasofrontal sac is tubular; in MM335 the right
naso-frontal is 14mm in diameter and the left 10
mm. The anterior portion of the nasofrontal sacs
bend posterolaterally, then medially to form a
U-shaped tube. The lateral wall of the right
nasofrontal sac has a series of perforations and
trabeculae, but no extensions from the nasofron-
tals. The nasofrontal sacs of MM333 are col-
lapsed, about 2.5 mm diameter for the left horn;
details are not obvious. In both specimens, the
nasofrontal sacs enclose an area comparable to
that covered by the vestibular sacs. In MM334,
the right nasofrontal sac also has trabeculae.
170
FIG. 36. Oblique dorsal view of section in Fig. 32. The
vestibular sac has been partly removed. The dull
probe marks passageway of spiracular cavity from
level of vestibular sac to exit between the blowhole
ligament (open arrow) and nasal plug (arrow). The pin
indicates anterior portion of naso-frontal sac. The lip
of the nasal plug has been drawn forward to reveal
inferior vestibule. The premaxillary sac (triangle) is
also more apparent due to displacement of the nasal
plug. There is no indication of posterior septum of
blowhole ligament nor posterior nasal sac.
Much of the anterior portion is collapsed and
visible in section as a slit (Fig. 35).
In MM335, a probe could be passed from the
posterior portion of the nasofrontal sac ventrally
to connect with the spiracular cavity; this portion
is termed the inferior vestibule (see discussion).
The inferior vestibule is bounded anteriorly by
MEMOIRS OF THE QUEENSLAND MUSEUM
the blowhole ligament, which runs laterally on
both sides from the nasal septum. In the sagittal
section of MM334, the inferior vestibule accom-
modates the lip of the nasal plug and runs behind
the blowhole ligament, apparently in connection
with a poorly defined posterior portion of the
nasofrontal sac. A small cartilage inclusion is
present in the blowhole ligament of MM334 (Figs
35,36). There is no indication of a posterior sep-
tum of the blowhole ligament (as defined by
Curry, 1992) in any of the specimens examined.
The cavity behind the blowhole ligament and
connecting with the posterior portion of the
nasofrontal sacs is not subdivided by a fold of
tissue; i.e. there is no indication of a posterior
nasal sac.
The prominent nasal plugs have a conspicuous
lateral lip, which fits into the inferior vestibule
(Figs 35,36). In MM333, these are 5 mm wide or
19% and 15% of the total width of the left and
right nasal plugs respectively. The melon enters
the right but not the left nasal plug.
The premaxillary sac is a thin-walled sac with
darkly pigmented tissue (Figs 33,35,36). In
MM335, the area of the premaxillary sacs is about
double (1.7—2.1 times) that of the vestibular sacs.
In MM333, in which the vestibular sacs appear
collapsed, the premaxillary sacs cover an area
about 5—7 times that of the vestibular sacs.
No accessory sacs were found.
Between the inferior vestibule and skull in the
sagittal sections of MM334 is a distinct connec-
tive tissue sheet which appears to be the diagonal
membrane (Fig. 33).
Anderson (1879) noted maxillary (=vestibular)
sacs, 'naso-facial' sacs entering the common
spiracular cavity and ‘large’ premaxillary sacs in
Orcaella brevirostris from India. His description
was not detailed enough to compare with more
recent studies on the respiratory tract.
Schenkkan (1973), Mead (1975), Heyning
(1989) and Heyning & Mead (1990) reviewed the
variations in the upper respiratory tract and as-
sociated nasal sacs. The cladistic analysis of
Heyning (1989) was especially affected by char-
acters of the facial region (24/40 characters), with
11/40 characters referring to the respiratory tract
alone. He identified the loss of the ‘posterior
nasal sac' as a synapomorphy for the Del-
phinidae. This feature needs to be considered in
more detail. The inferior vestibule ‘forms a com-
munication between the spiracular cavity and the
nasofrontal sac. Hence, if you have both of those
features, you, by definition, have the "inferior
vestibule” (Mead pers. comm., 8.3.94). Heyning
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA 171
TABLE 5. Data for cladograms.
Platanista O—000000 10001 1010—0 1002000000 1000000
Pontoporia 101010000100000 101000000020 100010100010
illustrated in Tursiops by Lawrence &
Schevill (1956, fig. 20b) than to Mead’s
(1975) fig. 4 of the same genus. Sagittal
sections of Orcaella (Figs 31-33) cor-
respond closely to those in Lawrence &
1 101000000 10010010100001002 1100000100010
1000102101 10000100—111020101110100111
Schevill (1956, figs 3, 4).
Whereas the anatomy of the
Globicephala
1000102101 10000100—101010111101100111
110110212001100101010000101101110111111
110110212001100101010010111101110111111
100101211000100101101000001211100101111
100111101000100101101000001211100101111
100101212000100101000000001211101101111
respiratory tract is well-documented for
delphinids (Schenkkan,1973; Mead,
1975), phocoenids (Curry,1992) and
ziphiids (Heyning,1989), the situation
is less clear in other odontocetes, in-
cluding the narwhal and beluga. For the
former, the only information is a
Orcaella
(1989) used the term ‘posterior nasal sac’ in its
simplest form to refer to a dorsal extension of the
inferior vestibule, as in the ziphiid Mesoplodon
(Heyning, 1989:10). In Berardius and Hyper-
oodon, there is a separation of the posterior cavity
into an anterior chamber which receives the
nasofrontal sac and a caudal chamber which was
referred to as the posterior nasal sac. The separa-
tion is by a transverse fleshy fold, the ‘hintere
klappe' of Kukenthal 1893 according to Heyning
& Mead (1990). Curry (1992) re-described the
facial anatomy of species of Phocoena and
Phocoenoides, and figured a posterior nasal sac
separated from the chamber receiving the
nasofrontals by an extensive sheet of connective
tissue which she referred to as the 'posterior
septum of the blowhole ligament'. As a result of
the well-developed septum, the upper respiratory
tract of phocoenids is well forward of the vertex
of the skull. This is shown by diagrams and
photographs in Heyning (1989), Curry (1992),
Reidenberg & Laitman (1987, fig. 3b) and
Schenkkan (1973). This contrasts with the more
posterior placement of the upper respiratory tract
in delphinids which lack or have a reduced
posterior septum (e.g. Pseudorca : Mead 1975)
and lack a posterior nasal sac (Mead, 1975; Heyn-
ing, 1989). The contrast is shown well by com-
paring the sagittal section of Phocoena
(Reidenberg & Laitman, 1987, fig. 3b) with Del-
phinus, Grampus and Globicephala (Reidenberg
& Laitman, 1987, figs 2a,2b,3a), and Orcaella
(Fig. 31).
In our Orcaella, the nasofrontal sacs are
clearest in sagittal sections. In all specimens the
tubular form of the posterior portion of the naso-
frontal sac is ill-defined and there is an extensive
connection between it and the inferior vestibule.
In this respect, the connection is closer to that
100111211000100101000010011111100100111
diagram (Huber,1934), which has been
discussed by Mead (1975) and Heyning
(1989). The occurrence of a posterior
nasal sac in narwhal is based on this diagram,
even though it is difficult to relate certain other
details in the diagram, e.g. the ‘lateral sacs’
(=vestibular sacs) and the nasofrontals, to the
pattern in other odontocetes. The diagrams and
description of the respiratory tract of beluga by
Kleinenberg et al. (1969) more clearly suggest a
posterior nasal sac but again are open to inter-
pretation. Fortunately, Heyning (1989) was able
to dissect a beluga and confirmed that the *in-
ferior vestibule divides dorsally into a rostral
nasofrontal sac and caudally into a posterior nasal
sac'. The upper respiratory tract appears to be
more anteriorly situated in beluga, as in
Phocoena, if we are interpreting correctly the
features in the photograph of a sagittal section
(Pilleri et al., 1980, fig. 13). This is consistent
with the suggestion above that the anterior posi-
tion of the respiratory tract is correlated with a
posterior nasal sac and well-developed posterior
septum. The diagram of the head of a narwhal
(Raven & Gregory,1933) also suggests that the
respiratory tract is far forward, and Schenkkan
(1973), based on Huber's ‘very clear diagrams’,
noted that the blowhole and nasal tract were
‘relatively more anteriorly’ situated in narwhal
than in other odontocetes.
The position of the nasal sacs and upper
respiratory tract in our Orcaella is consistent with
that in delphinids (Lawrence & Schevill, 1956;
Mead, 1975) and contrasts strongly with the
phocoenid pattern (with a well-developed
posterior septum of the blowhole ligament)
(Curry, 1992). The orientation of the respiratory
tract is not as well documented in monodontids,
but appears to be closer to phocoenids and thus
differs from what we observed in Orcaella.
Heyning (1989) noted that in the beluga, the
172
FIG. 37. Analysis One; all characters used. Cladogram
length 61; consistency index 70; retention index 75.
Abbreviations (for Figs 37-47) are: Pg(o): Platanista
gangetica (outgroup); Ig: Inia geoffrensis, Pb: Pon-
toporia blainvillei; Mm: Monodon monocerus; DI:
Delphinapterus leucas; Np: Neophocaena
phocaenoides; Pp: Phocoena phocoena; Gm:
Globicephala melas, Ob: Orcaella brevirostris; Tt:
Tursiops truncatus; Sc: Sousa chinensis.
vestibular sacs have apertures on the anterior
aspect of the vestibule, as in phocoenids. In our
Orcaella, the vestibular sac connects with the
spiracular cavity by a transverse slit as in Tur-
siops (Lawrence & Schevill, 1956, fig. 20a).
Thus, structure of the upper respiratory tract in
Orcaella is unlike that in either beluga or
narwhal.
The balance of evidence suggests that the
posterior nasal sac is a feature of ziphiids, beluga
& narwhal, and phocoenids but that it is lost in
delphinids as indicated by Heyning (1989). This
is one of the most soundly based morphological
synapomorphies for the Delphinidae. Thus the
apparent lack of a posterior nasal sac in Orcaella
supports its inclusion in the Delphinidae and ar-
gues against its close relationship with Del-
phinapterus. The position of the upper respiratory
tract in Orcaella, close to the vertex of the skull,
MEMOIRS OF THE QUEENSLAND MUSEUM
is also consistent with the pattern in delphinids.
The larger premaxillary sacs relative to vestibular
sacs in Orcaella is similar to the pattern in del-
phinids (Schenkkan, 1973). ‘Lateral lips on the
nasal plugs' was also inferred by Schenkkan
(1973) to be a specialised feature, best developed
in delphinids; Orcaella has well-developed
lateral lips on the nasal plugs. However, this
feature is inadequately known in other odon-
tocetes, including beluga and narwhal.
The form of the vestibular sacs and, more espe-
cially, trabeculae in the right nasofrontal sac of
Orcaella resemble Globicephala (Mead, 1975).
The trabeculae in Globicephala were considered
*extremely unusual' by Mead, and their occur-
rence in Orcaella and Globicephala, along with
the bulbous head, suggests a relationship between
the two genera. However, the melon structure is
different and other criteria need consideration in
assessing this relationship.
PHYLOGENETIC ANALYSIS
(Table 5)
RESULTS. In the first analysis all characters
were used and treated as non-additive. The two
cladograms produced (Fig. 37a,b) link beluga and
narwhal as a sister group to Phocoena, Neo-
phocoena, the delphinid genera and Orcaella.
The latter was placed with the delphinid genera,
although the two cladograms differed in detail. A
comparison of ancestral states in the two
cladograms was extracted by option ‘hcl’ of Hen-
nig86. This shows Orcaella linked with Sousa
and Tursiops because only the atlas and axis were
fused. Globicephala was closer to Phocoena and
Neophocaena as it has at least 3 cervical ver-
tebrae fused. This is a weak character on which
to base such a separation, but there is no a priori
basis to choose between the cladograms. How-
ever, except for the position of Orcaella and
Globicephala, the two cladograms are identical.
In the second analysis, mandibular symphysis
length, rostrum length, fusion of cervical ver-
tebrae, presence/absence of olecranon process,
roofing of temporal region were masked through
option 'cc' because they may be homoplastic.
The one cladogram produced (Fig. 38) was iden-
tical to Fig. 37b.
As is general practice (Forey et al.,1992), inap-
plicable values were coded '-, the same as miss-
ing values. Platnick et al. (1991) cautioned that
while missing and inapplicable values are treated
the same computationally, they are logically dif-
ferent (missing values can eventually be coded as
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
173
FIG. 38. Analysis 2; characters showing homoplasy
masked. Cladogram length 52; consistency index 69;
retention index 72.
0, 1, etc but there is no way to logically code
inapplicable values). They further suggested that
cladograms resolved on the basis of inapplicable
values may be questionable. In our analysis, this
could apply to the subdivision of the pterygoid
sinuses, which is a valid character for dolphins
and porpoises but inapplicable to monodontids.
In analysis 3 homoplastic (as in Analysis 2) and
Pg(o) Pg(o)
FIG. 40. Analysis 4; characters showing homoplasy or questionable polarity
masked. Cladogram length 33; consistency index 69; retention index 68.
FIG. 39. Analysis 3; characters showing homoplasy
and inapplicable characters masked. Cladogram
length 48; consistency index 68; retention index 73.
inapplicable characters (relating to subdivision of
the pterygoid sinuses) were masked.
In the single cladogram (Fig. 39), Tursiops and
Globicephala were linked, with Sousa and Or-
caella more distantly linked within the terminal
cluster of branches. The most important feature
is that Monodon and Delphinapterus were still a
group distinct from Phocoena, Neophocaena,
Orcaella and the delphinid genera. Thus, struc-
ture of the other cladograms
was not being driven by inap-
plicable characters. The char-
acters of the pterygoid
sinuses were retained in sub-
sequent analyses, because
they do allow clearer separa-
tion of the delphinid genera
without unduly affecting the
major branches of the
Ttt cladogram.
Polarity of a number of the
characters is equivocal. In
Analysis 4, the homoplastic
characters (Analysis 2) and
equivocal characters (orbit in
front of nares, convex profile
of skull, length of zygomatic
arch of squamosal, disap-
pearance of superior lamina
of pterygoid, orientation of
posterior process of periotic,
orientation of posterior
process of tympanic, form of
sigmoid process, presence/-
absence of lateral furrow,
breadth of lower tympanic
aperture, retraction of
premaxillaries from nasals,
174
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 41. As in Analysis 4, but with attachment to
mastoid pad masked. Cladogram length 31; consis-
tency index 70; retention index 67.
shape of anterior process of periotic) were
masked.
In all 4 cladograms produced (Fig. 40A-D),
Monodon and Delphinapterus were separated
from Phocoena, Neophocaena, Orcaella and the
delphinid genera. However, in two cladograms
(Fig. 40A,B) Orcaella was separated from
Pg(o)
FIG. 42. As in Analysis 4, but with all characters of
tympanoperiotic bones retained, despite equivocal
polarity. Cladogram length 42; consistency index 69,
retention index 71.
FIG. 43. Analysis 6; characters showing homoplasy
and multistate attributes masked. Cladogram length
40; consistency index 75; retention index 78.
Phocoena, Neophocaena and all the delphinids.
Analysis of the ancestral states suggested that this
is based on the ventral attachment of the tym-
panoperiotic bones to the mastoid pad of the
zygomatic arch. This was confirmed by re-run-
ning the analysis with that character masked (Fig.
41A,B) producing two cladograms identical to
Fig. 40C,D.
Because of the significance given to the tym-
panoperiotic bones, Analysis 5 retained tym-
panoperiotic characters, despite the equivocal
polarity. The cladogram (Fig. 42) was identical to
Figs 37B,38,40C,41A. Even with all tympano-
periotic characters retained, Monodon and Del-
phinapterus are linked, while Orcaella is linked
with Phocoena, Neophocoena and the delphinids.
Coding continuously varying multistate char-
acters (e.g. rostrum length, breadth of lower tym-
panic aperture) involves a more or less arbitrary
decision on where to set the limits of the various
categories. This introduces a bias so in Analysis
6 multistate characters were masked. Analysis
with characters exhibiting homoplasy and multi-
state characters masked (i.e. comparable to
Analysis 2) produced 2 trees (Fig. 43A,B) with
Monodon and Delphinapterus as a sister group to
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
175
FIG. 44. Analysis 7; characters showing homoplasy or questionable polarity or multistate attributes masked.
Cladogram length 30; consistency index 73; retention index 73.
Phocoena, Neophocaena, the delphinids and Or-
caella. In both cases Orcaella was linked with the
delphinids as the most derived taxa. This analysis,
with a shorter length of 40 (due to fewer at-
tributes), has the highest consistency index (0.75)
and retention index (0.78) of all analyses.
In Analysis 7 any multistate characters not al-
ready deleted because of homoplasy or ques-
tionable polarity were masked. This is
comparable to Analysis 4, and produced the same
FIG. 45. Nelson consensus tree based on Analysis 7.
set of cladograms (Fig. 44A-D). The length was
shorter (30 vs 33) because of the fewer characters
but the consistency and retention indices were
lowerthan in Analysis 6. A Nelson consensus tree
was determined using option 'nelsen' of Hen-
nig86 (Fig. 45).
Analysis 7 and Analysis 4 were apparently
driven by ventral attachment of the tym-
panoperiotic in Orcaella. Analysis 8 was the
same as Analysis 7 except that attachment of the
tympanoperiotic bones was also masked. The 2
cladograms (Fig. 46A,B), which were the same
as Fig. 41A,B had consistency index (0.75) com-
parable to Analysis 6, but lower retention index
(0.73). The Nelson consensus tree is Fig. 47.
DISCUSSION
TAXONOMIC RELATIONSHIPS OF
MONODONTIDAE-PHOCOENIDAE-DELP
HINIDAE. The cladograms consistently
separated phocoenids and delphinids from the
monodontids Delphinapterus and Monodon, and
could even intersperse the phocoenids with the
delphinids; this suggests that the phocoenids and
176
FIG. 46. Analysis 8; as in Analysis 7 but with ventral
attachment of the tympanoperiotic bones masked.
Cladogram length 28; consistency index 75; retention
index 73.
delphinids together form the sister group to the
monodontids. This extends previous studies
(Heyning, 1989; Gretarsdottir & Arnason, 1992;
Milinkovitch et al., 1994) in which the Monodon-
tidae-Phocoenidae-Delphinidae remained an un-
resolved trichotomy.
We could not demonstrate any synapomorphies
for the delphinid-phocoenid clade. Heyning
FIG. 47. Nelson consensus tree based on Analysis 8.
MEMOIRS OF THE QUEENSLAND MUSEUM
(1989) indicated one, 'extreme reduction of
zygomatic process of squamosal'. He considered
the zygomatic process reduced in Orcaella, but
we have shown that the zygomatic arch is sub-
stantial with a prominent mastoid section and that
the mastoid pad provides the attachment point for
the tympanoperiotic bones. On this character,
Orcaella would be excluded from the phocoenid-
delphinid clade. This occurred in our cladogram
when ‘attachment of the tympanoperiotic bones’
was not masked. We suggest that form of the
zygomatic arch is a reversal and not primitive in
Orcaella; this is based on a series of synapomor-
phies it shares with the Delphinidae (Implications
for previous classifications, below). Heyning
(1989) characterized the zygomatic arch of the
delphinid Orcinus as 'substantial' an implied
reversal which he associated with the handling of
large prey by killer whales. The poorly developed
mesethmoid plate in Orcaella is possibly primi-
tive, linking it more with phocoenids than del-
phinids. We suggest, however, that the poorly
developed mesethmoid plate is a neotenic fea-
ture, most apparent in Australian material.
Monodontids and phocoenids have a suite of
characters including short rostrum, no beak, short
mandibular symphysis, poorly developed
mesethmoid plate, widely separated pterygoid
hamuli, and relatively unspecialized pre- and
post-orbital lobes (especially so in monodontids).
As our cladograms suggest that these families are
the closest living relatives of delphinids, by out-
group comparison we would expect the features
just listed to also occur in the most primitive of
the delphinids. This suggests that blunt-headed
Pseudorca, Orcinus, Globicephala and Orcaella,
as well as Cephalorhynchus (which shows many
parallels to phocoenids) are the most primitive
delphinids. The alternative suggestion is that
long-beaked delphinids such as Sotalia and Sousa
are the most primitive extant delphinids (e.g.
Kellogg, 1928; Fraser & Purves, 1962, using the
Stenidae) and that the blunt-headed genera are
convergent. Three lines of evidence can test the
alternatives: (1) molecular data; (2) more exten-
sive fossil material and (3) anatomical studies to
determine whether the morphology of blunt-
headed odontocetes is homologous. The
molecular phylogeny of Milinkovitch et al.
(1994, fig.1) is consistent with our cladograms,
in that monodontids and phocoenids are
separated from the delphinids as potential sister
groups. To that extent their results support the
possibility that blunt headed dolphins are primi-
tive. Anatomical studies (Mead, 1975; herein)
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
suggest, however, that the facial anatomy may be
quite diverse and non-homologous in ‘blunt-
headed’ dolphins, The anatomy of the facial
region is generally considered to reflect adapta-
tions to sound production (Heyning, 1989; Heyn-
ing & Mead, 1990) and features of the skull may
reflect such modifications. Thus the possibilities
of convergence are extensive. The demonstration
of neotenic features of the skull in Orcaella
provides a mechanism whereby more generalized
features may be expressed, without implying that
the taxon is primitive. The polarity of each char-
acter used in classification will therefore have to
be examined before the alternative interpretations
can be properly evaluated.
IMPLICATIONS FOR PREVIOUS CLAS-
SIFICATIONS. Despite differences between our
various analyses, the following were consistent:
1, Monodon and Delphinapterus were linked; 2,
Orcaella was not linked with Delphinapterus (3)
Phocoena and Neophocaena were linked; 4,
Phocoena, Neophocaena, Orcaella and the del-
phinids were the sister group to Monodon and
Delphinapterus; and 5, Tursiops and Sousa were
linked, as the most derived branch.
Fourteen characters were not used (Appendix
2), usually because of insufficient comparative
data. Available information for these characters
is consistent with the results just outlined, except
for the immunological and electrophoretic data of
Lint et al. (1990) which is discussed later. The
linking of Tursiops and Sousa may reflect the
limited subset of delphinids used and is the most
weakly supported of our conclusions. The other
results have more general implications for the
classifications in Fig. 1:
1. OWEN (1866) (Fig. 1A). Orcaella is never
linked with Phocoena and Neophocaena in our
cladograms. When Owen described O. brevi-
rostris, the concept of Phocoena was broader
than it is today. Orcaella has a dorsal extension
of the pre-orbital lobe, as in phocoenids. How-
ever, this extension was intermediate in develop-
ment between that in phocoenids and that shown
by delphinids such as Pseudorca (pers. obs). Or-
caella has none of the other synapomorphies of
phocoenids (e.g., premaxillary boss, spatulate
teeth, folded vestibular sacs).
2. KASUYA (1973) (Fig. 1B). Monodon and
Delphinapterus were consistently linked, even
when all tympanoperiotic bone characters were
included (Fig.41). Our results do not support their
family level separation (Kasuya, 1973); they
validate Gray’s (1821) Monodontidae. This fami-
177
ly is as diagnosed by Tomilin (1967, as Del-
phinapterinae) and Rice (1984), except that the
periotic bones do not appear to be fused to the
squamosal as stated by the former author. The
fossil Denebola Barnes, 1984 would also be in-
cluded in the family, based on the anterior posi-
tion of its orbits and the extensive border of the
nares by the maxillary bones. Other fossils
presumed to be monodontids (e.g. in Pilleri et al.
1989) need to be re-evaluated.
Orcaella and the delphinids were linked in our
cladograms based on the absence of a posterior
nasal sac and the lateral orientation of the
posterior process of the tympanic bulla. We have
shown elsewhere that characters of the tympano-
periotic bones proposed as the basis for the Del-
phinapteridae by Kasuya (1973) and Pilleri et. al.
(1989) are variable or provide only equivocal
support. Moreover, Orcaella shared none of the
synapomorphies identified for Delphinapterus
(anterolateral margins of superior nares bordered
by maxillaries, curvature of flippers & convex
profile of skull), nor the undivided cranial
sinuses, retention of lateral and superior lamina
of the pterygoid bones & anterior position of the
orbits with associated prominent palatine bridge
under the orbit. Hence we found no support for
the Delphinapteridae Kasuya, 1973. Other fea-
tures used to unite Orcaella with Delphinapterus
appear to be either primitive characters or have
proved to be variable when a larger series of
specimens were examined. In the first category is
the flexibility of the head and cervical sinus,
shared not only with beluga but also with river
dolphins, such as /nia, and Neophocaena. Despite
statements to the contrary, this flexibility is not
associated with separate cervical vertebrae: while
these are separate in monodontids and river dol-
phins, Orcaella has the atlas and axis fused as in
most delphinids.
The light colour may be a feature of riverine
populations, but the Queensland animals at least
have a three-tone colour pattern such as in Tur-
siops, rather than the more diffuse pattern of
monodontids. The lack of a beak and abbreviate
melon are primitive features shared not only with
beluga but also phocoenids such as Neo-
phocaena. Moreover, the melon in Orcaella ap-
pears distinct from that in beluga. Similarities in
general appearance led Mitchell (1975) to sug-
gest ‘a phyletic relationship’ between Orcaella
and *the Arctic white whale', but we believe that
the features are either shared primitive characters
or convergent. We similarly suggest that ventral
attachment of the tympanoperiotics in Orcaella
178
is a primitive feature shared with monodontids
and river dolphins such as Platanista, and thus
has no taxonomic significance .
Characters which are more variable than
originally described include most of the features
of the tympanoperiotic bones, such as the
posterior orientation of the posterior process of
the bulla, width of posterior branch of lower
tympanic aperture and the supposedly diagnostic
statistics of the periotic triangle. The orientation
of the processes and width of the LTA also char-
-acterize phocoenids, as much as Delphinapterus.
Our larger series of specimens indicates that an
olecranon process is usually present in Orcaella
(Fig. 48A,B; cf. de Muizon, 1988), as in many
delphinids but not Delphinapterus or Monodon
which lack an olecranon process (Tomilin, 1967).
The deltoid tuberosity of the humerus is more
extended in Orcaella than in most delphinids
(associated with the greater length of the
humerus) but is not as subdistal as in monodon-
tids (de Muizon, 1988, fig. 22). It, therefore,
provides only equivocal evidence that Orcaella
is a monodontid (cf. de Muizon, 1988).
Phocoena and Neophocaena were consistently
linked most closely with Orcaella and delphinids
Tursiops, Sousa and Globicephala; this does not
support wide separation of the Phocoenidae and
Delphinidae as in Fig. 1B.
3. BARNES (1984), GASKIN (1982)(Fig. 1C).
Both authors linked the phocoenids and del-
phinids as in our cladograms. However, their
linking Monodon, Delphinapterus and Orcaella
in the same family contrasts with our findings.
Including Monodon and Delphinapterus in the
same family implicitly rejects the orientation of
the posterior processes of the bulla and periotic,
the compression of the bulla and the width of the
lower tympanic aperture as valid family charac-
ters, since these all differ in the two genera.
However, orientation of the posterior processes
and width of the tympanic aperture were the two
main characters used to unite Orcaella and Del-
phinapterus in the Delphinapteridae.
As discussed above, other characters which
link these genera are shared primitive features,
most of which could be equally used to support a
relationship between Orcaella and the
phocoenids.
Miller (1923) placed Delphinapterus and
Monodon in different subfamilies based on dif-
ferences in dentition, the pterygoid hamuli and
extent to which the alisphenoid was overspread
by the superior lamina of the pterygoids. Except
for dentition, these features are not known for
MEMOIRS OF THE QUEENSLAND MUSEUM
Denebola, the only other monodontid accepted in
this paper. Although Monodon and Delphinap-
terus are distinct, we question the need for sub-
family separation.
4. PILLERI et al. (1989) (Fig. 1D). This clas-
sification suffers from the same inconsistencies
noted in the last section. In addition, the
phocoenids and delphinids were widely
separated, the former being placed with the river
dolphins Platanista, Inia and Pontoporia. None
of our cladograms support this classification:
phocoenids differ from the river dolphins in
profile of the tympanic bulla, shape of the sig-
moid process, loss of lateral furrow, symmetry of
vestibular sacs, roofing of the temporal region,
position of the orbits relative to nares and com-
plexity of the cranial sinuses. We consider the
delphinids and phocoenids closely related based
on morphology (de Muizon, 1988, 1990; Heyn-
ing, 1989), and ribosomal DNA (Milinkovitch et
al, 1993). This relationship is also consistent
with the molecular phylogeny in Milinkovitch et
al. (1994), which included the river dolphin /nia.
5. LINT et al. (1990) (Fig. 1E). Our results
agree with placing Orcaella in the Delphinidae.
The major discrepancy in their classification is
the extreme separation of phocoenids and del-
phinids. As indicated in the previous section, this
is not supported by any of our cladograms and is
inconsistent with most previous classifications. It
was based on a combination of immunological
and electrophoretic results, the latter incorporat-
ing the data of Shimura & Numachi (1987).
Shimura & Numachi (1987) used a limited range
of species; the ziphiid Berardius was the only
non-delphinid taxon compared with phocoenids.
It is thus possible that ziphiids and phocoenids
were linked in the phenogram because they both
differed from delphinids, rather than because they
were closely related. Even with the extra species
considered by Lint et al. (1990), the analysis was
biased heavily towards delphinids, which may
affect their conclusions on relationships among
higher odontocete taxa.
6. FRASER & PURVES (1962) (Fig. 1F). We
place Orcaella in the Delphinidae, as proposed
by Fraser & Purves (1962). There are two aspects
of their classification which we would question.
The first is the linking of Sousa, Sotalia and Steno
in the Stenidae, which is considered a more primi-
tive taxon than the Phocoenidae. Grouping Steno,
Sotalia and Sousa as the Stenidae (Fraser & Pur-
ves, 1962) is based on inconsistent or unclear
criteria. The rugose teeth of Steno, proposed as a
primitive character, is not shared by the other two
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
genera, The elongate rostrum and elongate man-
dibular symphysis, appear to be variable features
in other odontocete families (Heyning, 1989) and
may not be primitive. The length of rostrum ap-
pears to be particularly subject to reversals and
convergences: a long rostrum occurs not only in
less derived taxa such as Platanista but in more
specialized taxa, such as Stenella, which we con-
sider among the most derived of delphinids. Pre-
vious authors (Kellogg, 1928) emphasized the
resemblance of kentriodontids and long-snouted
dolphins such as Sotalia (closely related to
Sousa). In light of the much more primitive form
of the kentriodontid skull (e.g. lacking asym-
metry), the resemblance may be superficial and
dependent on variable features such as rostrum
length and length of mandibular symphysis.
Fraser & Purves (1962) pointed out the mixture
of supposedly primitive characters and an ad-
vanced cranial sinus system in Sotalia and Sousa.
We have not examined Sotalia but our Sousa
chinensis skulls suggest close coalescence of pre-
and post-orbital lobes to surround the optic nerve,
as inferred by Fraser & Purves (1962). In addi-
tion, the excavation of the bones in the sphenoidal
region of Sousa skulls we examined was complex
and similar to that in more advanced delphinids
such as Tursiops and Stenella. The consistent
grouping of Tursiops and Sousa as the most
derived group in our cladograms further suggests
a more derived condition for Sousa than recog-
nized in the systems of Kellogg(1928) and Fraser
& Purves (1962). We have to recognise the
limited range of delphinids considered in our
analysis, however, so that our results may be
biased.
In most of our cladograms, the phocoenids ap-
peared as less derived than the delphinids. In Figs
40B & 44B the phocoenids were aligned to Tur-
siops and Sousa, but the cladogram was un-
resolved. The more generalized state of
phocoenids is further supported by studies of base
pair length of repetitive DNA: the phocoenids
shared the 1750 base pair length with other taxa
such as ziphiids, monodontids, etc whereas the
delphinids had a unique base pair length of
around 1580. The phocoenids were separated,
with monodontids, from the delphinids in the
molecular phylogeny of Milinkovitch et al.
(1994). The phocoenids, nonetheless, show
specializations (e.g. dorsal extensions of the
preorbital pterygoid lobe, morphology of the
nasal region which was considered by Klima &
van Bree (1985) to be more derived in Phocoena
than in other odontocetes examined). These
179
specializations may reflect a long period of
separation from the delphinids. The large number
of synapomorphies for the Phocoenidae suggests
it is a conservative body plan, although many of
the characters are variably expressed throughout
the family (Perrin pers. comm.; Appendix 2). Our
results do not support the classification (Fraser &
Purves, 1962) which placed the Phocoenidae be-
tween the Stenidae (including Sousa) and the
Delphinidae.
An even greater disparity occurs between our
results and the position of the Monodontidae in
Fraser & Purves' (1962) classification. They con-
sidered the undivided form of the pterygoid
sinuses a primary feature and the basis for their
Superfamily Monodontoidea. As argued else-
where, this feature needs to be confirmed by
dissection. Beluga and narwhal appear to be more
primitive than Phocoena, Neophocaena and del-
phinids (Figs 37-47). However, Fraser & Purves
(1962) suggested that they are more primitive
than all odontocetes other than ziphiids.
Monodon and Delphinapterus are consistently
linked with Phocoena, Neophocaena and del-
phinids (Figs 37—47) arguing for their retention
within a single taxon, such as the Delphinoidea
(Heyning, 1989). Moreover, characters linking
monodontids with Phocoena, Neophocaena and
delphinids are derived features whereas those
linking them to river dolphins are primitive or of
questionable polarity (e.g. anterior position of
orbits). Although Heyning (1989) and de Muizon
(1988) did not resolve relationships between
monodontids, delphinids and phocoenids, their
analyses showed they form the most derived
group of living odontocete taxa. Grouping
monodontids, phocoenids and delphinids is also
supported by some chemical data, such as the
distribution of isovaleric acid in acoustic fat
(Litchfield et al., 1975; Appendix 2). This group-
ing was also evident in molecular phylogenies
(Milinkovitch et al., 1994), de Muizon (1988) and
Heyning (1989) indicated a series of synapomor-
phies for physeterids and river dolphins
(Platanista, Inia, Pontoporia, Lipotes), all of
which would have to be considered reversals if
Fraser & Purves' (1962) position of monodontids
was accepted. Our results support de Muizon
(1988, 1990) and Heyning (1989), indicating that
the undivided pterygoid sinuses must be con-
sidered a reversal rather than a primary feature in
monodontids. This removes the basis for separat-
ing beluga and narwhal in their own superfamily.
7. NISHIWAKI (1963, 1964, 1972) (Fig 1G).
Nishiwaki (1963, 1964) proposed the Orcellidae
180
(‘Orcaelidae’ of Nishiwaki 1972), with: 1, only
atlas and axis fused; 2, size less than 12 feet (4
m); 3, no beak and less than 20 teeth in each row
of the upper jaw. The first two characters are also
consistent with the Delphinidae as defined by
Nishiwaki. Although he characterized delphinids
as having a distinct beak, he included genera such
as Cephalorhynchus where the beak is reduced or
absent. The only character separating Orcaella
from delphinids is tooth number, but this seems a
questionable basis for family separation given the
variability in tooth number within other families
(e.g. phocoenids). This character would no longer
hold if Grampus, put in its own family by
Nishiwaki, is considered a delphinid (Mead,
1975). Moreover, fossil Tursiops have as few as
14 teeth per side of each jaw (Barnes, 1990).
While we have reservations about the validity
of the characters used by Nishiwaki, Orcaella
does have characters (e.g. attachment of the tym-
panoperiotics to the mastoid pad of the zygomatic
arch, the development of the zygomatic arch)
which separate it from the delphinids and
phocoenids. These characters could indicate a
distinct family for Orcaella. We reject this option
because Orcaella shares several derived features
with the Delphinidae, including loss of posterior
nasal sac, 1580 bp length of repetitive DNA,
maintenance of the right premaxillary close to the
nasals, apical position of nasals, development of
the mesethmoid plate (de Muizon, 1988; Heyn-
ing, 1989; Barnes, 1990; Gretarsdottir & Ar-
nason, 1992), although the last feature may vary
between populations. These features imply a
commonality of the most recent ancestor. Al-
though some of the cladograms (e.g. Fig. 37B)
indicate a basal position among delphinids for
Orcaella, none suggested that it is a distinct fami-
ly. The most parsimonious conclusion is that the
zygomatic arch and attachment of the tym-
panoperiotic bones in Orcaella are reversals. The
other characters which separate Orcaella from
delphinids are either unique to Orcaella, or
probably neotenous; in either of these cases they
are of no value in assessing relationships between
taxa. Toemphasize them, rather than evidence for
most recent common ancestor, promotes a system
based on discordancy, characterised by an ever
increasing number of monotypic taxa.
8. SLUPER (1936), FORDYCE (1984), DE
MUIZON (1988), HEYNING (1989), etc. (Fig.
1H). We agree with the ‘classic’ placement of
Orcaella in the Delphinidae s.l. Orcaella was
commonly (Figs 40c, 41a, 43a, 46b) placed as the
least derived in the series Orcaella, Globicephala
MEMOIRS OF THE QUEENSLAND MUSEUM
and (Tursiops + Sousa). However, equally com-
monly (Figs 40d, 41b, 43b, 46a) there was an
unresolved trichotomy of Orcaella,
Globicephala and (Tursiops + Sousa). Gray
(1866, 1871), Anderson (1879), Fraser & Purves
(1962), Mead (1975), de Muizon (1988) and
Heyning (1989) placed Orcaella with the 'blunt-
headed' dolphins such as Orcinus, Pseudorca and
Globicephala. We have indicated skull features
(separation of the medial and lateral lobes of the
palatines, posterior extension of the lateral lobe
of the palatine bone) which link Orcaella with the
*blunt-headed' dolphins, especially Globi-
cephala and Pseudorca. What is unresolved is
whether these features are evidence of relation-
ship or whether they are convergent, reflecting
shared constraints on morphology (e.g., short
rostrum). Fraser & Purves (1962) suggested a
series of increasing specialization from Pseudo-
rca to Orcinus, Orcaella, Globicephala and
Feresa. This was based on what we consider
minor variations in the relative development of
the pre- and post-orbital lobes, which have been
largely inferred from skulls and not confirmed by
dissection. In addition there can be considerable
variation in the sinuses within taxa, e.g. Pseudo-
rca (Fraser & Purves, 1962; Purves & Pilleri,
1978, fig. 18). Pseudorca was apparently placed
as most primitive because of the more extensive
development of the bony lateral lamina of the
sinuses. However, this is formed from the
palatine bone, rather than the lateral lobe of the
pterygoid bone, as in monodontids and some river
dolphins which were considered less derived by
Fraser & Purves (1962). The lamina may not be
homologous among these genera.
de Muizon (1988) placed Orcaella in the
Globicephaliinae based on 'dilation of the
premaxillae at apex of rostrum'. He illustrated the
tip of the snout of Orcaella, composed entirely of
premaxillae. However, the maxillae do not reach
the tip in delphinids such as Tursiops or Sousa
(pers. obs). Moreover, neither the premaxillae nor
the rostrum show the broad transverse expansion
as in Pseudorca, and especially Globicephala
melas. Premaxillae/rostrum width at 0.75 length
in Orcaella was 65.0% (56.096—73.196), which is
comparable to Lagenorhynchus (Sergeant &
Fisher, 1957) or Tursiops (Ross, 1977) in the
Delphiniinae sensu de Muizon.
Cladograms, which place phocoenids and
monodontids as the nearest living relatives of
delphinids, suggest by outgroup analysis that fea-
tures such as short rostrum of skull, widely
separated pterygoid hamuli and unspecialized
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
cranial sinuses are primitive within the del-
phinids. Similarities of Orcaella to phocoenids,
especially Neophocaena, as well as monodontids,
suggests it is one of the most primitive of the
delphinids. Dudok van Heel (in Kamminga et al.,
1983) suggested that Orcaella was one of the
‘oldest “modern” species.."forced" inshore by
more recent newcomers’. Although we have
reservations about invoking competition to ex-
plain delphinid distributions, our data are consis-
tent with Orcaella being a less derived taxon (at
least within the Delphinidae, s./.). Such an inter-
pretation reinforces the generalized nature of
genera such as Cephalorhynchus (which shows
many parallels to phocoenids), as well as Pseu-
dorca, Orcinus and Orcaella. The generalized
features in these genera make it difficult to estab-
lish natural groupings of such genera based on
shared derived features. Moreover, Mead (1975)
indicated the wide morphological diversity in the
facial anatomy of ‘blunt-headed’ dolphins which
suggests that they may not be a natural group.
Grampus and Peponocephala, in particular, fit
poorly with the other genera at least based on the
cranial sinus anatomy. Given the generalized and
apparently primitive characters which link them,
it is difficult to assess whether Pseudorca, Or-
cinus and Globicephala form a natural grouping,
and whether Orcaella is related to any of them.
Orcaella, for instance, has relatively few teeth,
but they are not enlarged as in the other genera
(e.g. Pseudorca, which has enlarged teeth even in
the foetus: Pilleri & Purves, 1978, fig. 17).
Cladistic analyses assume that convergence is
not so widespread that it masks underlying
phylogenetic patterns; by outgroup analysis fea-
tures of Orcaella are assumed to be primitive.
The evidence for convergence in toothed whales
is overwhelming, however, for characters such as
rostrum length and probably mandibular sym-
physis length. A lack of knowledge of the func-
tional significance of many of the other features
used in cetacean classification (e.g. facial
anatomy) prevents a balanced consideration of
the extent of convergence in delphinids and other
odontocetes. Neoteny in Orcaella further compli-
cates interpretation; it provides a mechanism
whereby apparently primitive features could
occur in a more derived species. We conclude that
Orcaella is not a monodontid or delphinapterid
(sensu Kasuya, 1973). The balance of evidence
suggests it is a delphinid s. /. However, given the
ambiguity of characters used to define groupings
within the Delphinidae and limitations in the
comparative material of delphinids available to
181
us, we did not consider it worthwhile to pursue
our cladistic analyses further (e.g. by using dif-
ferent combinations of delphinid genera) in order
to place Orcaella within the family. Recent DNA
studies may provide important new evidence on
the relationships of toothed whales. However,
rigorous phylogenetic techniques on a wider
range of characters than were available for this
study, should be rewarding. Such studies will also
provide the necessary tests for taxonomic
hypotheses generated by DNA and other
molecular techniques.
ACKNOWLEDGEMENTS
We thank H. Marsh, B. Dowd, R. Kenchington,
A. Spain, A. Birtles, M. Devereux, G. Hopper, E.
Drom, L. Price, M. Downes, A. Witting, T. Hein-
sohn, R. Heinsohn, A. Haffenden, J. Lanyon, A.
Preen, K. Saalfeld (all at James Cook University),
T. McNamara (Queensland Dept. Primary In-
dustries) & K.Adler (Queensland Dept Environ-
ment & Heritage) for help with carcass salvage.
We thank J. Hoey (Great Barrier Reef Aquarium)
for access to freezer space. We thank Bill Dowd
for careful preparation of the skeleton of
MM1015. Robert Patterson and Steve Van Dyck
provided access to the Queensland Museum col-
lection. For similar help, we would like to thank
F. Scott (Nova Scotia Museum of Natural His-
tory, Halifax, Nova Scotia), D. Balkwill
(Canadian Museum of Nature, Ottawa, Ontario),
J. Ford and education staff at the Vancouver
Aquarium (Vancouver, British Columbia), and
D. Canning (Dept. of Zoology, University of
British Columbia). A. Preen (James Cook
University) provided a specimen of
Neophocaena and made observations for us on
live Orcaella at the Jaya Ancol Aquarium, Jakar-
ta, Indonesia. We thank J.L. Bannister for ex-
amining Orcaella in the Western Australian
Museum and R. Sabin for examining the
holotype. Preparation of teeth and counting of
dentinal growth rings were carried out by N.
Goudberg, under the direction of Helene Marsh
(James Cook University). A. Williamson helped
measure tympanoperiotic bones. We also thank
Helene Marsh for access to literature. We thank
Bruce Cowell (Queensland Museum), R.
Yeldham (James Cook University) and M.
Pulsford (Natural History Museum) for photog-
raphy and J. True (Museum of Tropical
Queensland) for Fig. 1. We thank C. de Muizon
(Institut Francais d'Etudes Andines, Lima, Peru)
and B. Kamminga (Delft University of Technol-
182
ogy, Delft, The Netherlands) for papers and com-
ments on Orcaella. For helpful comments, we
are grateful to D. Bellwood (James Cook Univer-
sity), D. Gaskin (University of Guelph), J. Mead
(Smithsonian Institution), C. Wallace (Museum
of Tropical Queensland), W.F. Perrin (Southwest
Fisheries Science Centre, La Jolla, California)
and G.J.B. Ross (Australian Biological Resour-
ces Study, Canberra).
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APPENDIX 1: Measurements of tympano-
periotic bones (after Kasuya, 1973). Measure-
ments 1 and 13 in mm. Measurements 2-10 as %
of standard length of tympanic bulla. Measure-
ments 14-19, 23-28 as % of standard length of
periotic.
TYMPANIC BULLA
1. Standard length of tympanic bulla, distance from
anterior tip to posterior end of outer posterior
prominence
2. Distance from anterior tip to posterior end of inner
posterior prominence
3. Distance from posteroventral tip of outer posterior
prominence to tip of sigmoid process
4. Distance from postero-ventral tip of outer posterior
prominence to tip of conical process
5. Width of tympanic bulla at level of the sigmoid
process
6. Height of tympanic bulla, from tip of sigmoid
process to ventral keel
7. Width across inner and outer posterior prominences
8. Greatest depth of interprominential notch
9. Width of upper border of sigmoid process
10. Width of posterior branch of lower tympanic aper-
ture
PERIOTIC
13. Standard length of periotic, from tip of anterior
process to posterior end of posterior process, measured
on a straight line parallel with cerebral border
14. Thickness of superior process at the level of upper
tympanic aperture
15. Width of periotic across cochlear portion and supe-
rior process, at the level of upper tympanic aperture
16. Least distance between the margins of the fundus
of internal auditory meatus and of aperture of ductus
endolymphaticus (aquaeductus vestibuli)
17. Least distance between the margins of the fundus
of internal auditory meatus and aperture of aquaeduc-
tus cochleae
18. Length of articular facet of the posterior process of
periotic for the posterior process of tympanic bulla
19. Anteroposterior diameter of cochlear portion
22. Length of periotic shown by the percentage of
length of tympanic bulla
23. Separation of foramen singulare and aquaeductus
Fallopi
24. Angle of lateral margin of posterior process of
tympanic bulla to sagittal axis
MEMOIRS OF THE QUEENSLAND MUSEUM
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PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
25. Maximum width of fundus of the internal auditory
meatus
26. Width of foramen ovale
27. Width of foramen rotundum
28. Width of head of malleus
APPENDIX 2: CHARACTER STATES
1. Vestibular sacs. Heyning (1989) demonstrated
that vestibular sacs are a feature of all odontocetes
except Physeteridae (including Kogia), Ziphiidae
and Platanista. Schenkkan (1973) described ves-
tibular sacs in ziphiids, but we follow the inter-
pretation by Heyning, who had access to a wider
range of material and used supplementary criteria
such as position of the sacs relative to muscle
layers. Lack of vestibular sacs is primitive by out-
group analysis, as suggested by Heyning (1989).
2. Folded vestibular sacs. This is a specialized
feature of the Phocoenidae, as demonstrated by
numerous authors. It was coded as - in Platanista
which lacks vestibular sacs; 0 for those genera
with unfolded vestibular sac (primitive condi-
tion), and 1 for Phocoena and Neophocaena, with
folded sacs.
3. Hypertrophied right vestibular sac. Schenkkan
(1973) and Heyning (1989) noted this specializa-
tion of the river dolphins (except Platanista). It
was coded - for Platanista without vestibular
sacs, 0 for those with symmetrical sacs (primi-
tive) and 1 for /nia and Pontoporia.
4. Orbit in front of nares. Miller (1923) noted this
feature in Monodon and Delphinapterus; it oc-
curs in all the river dolphins. It appears to be
present in Physeter (van Beneden & Gervais
1868-79; Kellogg, 1928), but not in Kogia simus
(Perrin pers. comm.) It is not so in extant ziphiids,
butitis in Squaloziphius, which was considered
to be a ziphiid by de Muizon (1990) (but see
Fordyce & Barnes, 1994).
Miller (1923) noted the posterior opening of the
antorbital canal near the anterior of the orbit in
delphinids (based on Delphinus), as in terrestrial
mammals. Since the antorbital canal was more
posterior in those species where the orbit was in
front of the nares, he argued that this was the
specialized condition. However, the posterior
opening of the antorbital canal may also be
situated more posteriorly relative to the orbits in
phocoenids and Pseudorca, which do not have
the orbits in front of the level of the nares (pers.
obs.). This compromises Miller's argument and
187
the polarity remains equivocal. In the juvenile
narwhal UBC 9467, the orbit was already anterior
to the nares, so there is no indication of the
polarity state from this ontogeny.
Polarity is equivocal based on outgroup
analysis, comparative anatomy and ontogeny.
The more anterior position of the orbit is con-
sidered primitive based on river dolphins and
physeterids, and possibly ziphiids (the last based
on Squaloziphius, which Fordyce & Barnes
(1994) considered ‘more reminiscent of’ the Eur-
hinodelphidae than Ziphiidae). If they are right,
orbit in front of the nares may be a primitive
feature by outgroup analysis since the Eur-
hinodelphoidea is a possible sister group to the
Delphinida sensu de Muizon. Coding the anterior
position of the orbits as primitive makes the
fewest assumptions about the relationships of the
river dolphins and monodontids .
5. Separation of pterygoids
Physeter, Berardius and Platanista all have
medially appressed pterygoids, and by outgroup
analysis this is considered the primitive state.
Separation of pterygoids is coded as 1.
The palatines and vomer in Orcaella resemble
those in delphinids where there has been a com-
pression of the palatines and median vomerine
spine by the pterygoids. It therefore may not be
homologous with the situation in phocoenids,
monodontids and some primitive delphinids. All
species with widely separated pterygoids are
nonetheless scored 1.
6. Posterior nasal sac
We accept Heyning’s (1989) argument that a
posterior nasal sac is primitive. Its occurrence in
the narwhal is from Huber (1934), but Heyning
confirmed it in Delphinapterus. We have not
been able to confirm the loss of this sac in Sousa,
but it is scored as lacking the sac, as this is a
consistent feature of delphinids examined.
7. Mandibular symphysis length
Usually expressed in binary form (short/long)
but animals examined fell into 3 ranges: >50%
mandible length (in Physeter and the river dol-
phins), 30-50% (Sousa and Steno (Perrin pers.
comm.)) and «3046. The polarity is equivocal by
outgroup analysis, as Physeter has an elongate
mandibular symphysis, but it is shorter (<50%) in
Kogia and <30% in ziphiids. Coding is 0 =>50%,
1230-5046 and 2=<30%, but since all characters
were treated as nonadditive no direction of
specialization is implied.
188
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 48A-C. Variability in development of olecranon process (arrow, in A) in 3 specimens of Orcaella from
central Queensland, Note the well-developed process in A,B. Also note the elongate deltoid tuberosity of the
humerus (open arrow in A).
8. Rostrum length
An elongate rostrum (>60% skull length) was
considered primitive, by outgroup analysis. A
wide range of fossil odontocetes also have an
elongate rostrum. Neophocaena has a particular-
ly short rostrum (<40%), while all other genera
considered in this study are in the 40-50% range.
All are coded 1, however, to reduce the com-
parisons to binary values. As discussed else-
where, this feature seems particularly subject to
convergences and possible reversals.
9. Atlas and axis separate
The polarity of this character is equivocal by
outgroup analysis. Physeter has a unique arran-
gement, where the atlas and axis are separate, but
cervicals 2-7 are fused. Extant ziphiids have at
least 3 cervicals fused, including the atlas and
axis. However, de Muizon (1988) noted that
most Mio-Pliocene cetaceans had separate cervi-
cals. This implies that there may have been rever-
sals in some lineages (separate cervicals to par-
tially fused to separate) and thus the character is
subject to homoplasy. In the initial analysis, the
unfused condition was considered primitive and
coded 0 in the analysis. Where only the atlas and
axis are fused, the character is coded as 1, while
it is 2 for cases when 3 or more cervicals are
fused. This extent of fusion of cervical vertebrae
has been used (e.g. Nishiwaki, 1963) to define
families so it is included as a multistate variable.
However, we recognize that the number of fused
cervical vertebrae is variable even within a
species. Fischer (quoted in Tomilin, 1967) noted
that in Tursiops there could be fusion of the first
two, or the third-fourth and fifth, or even the sixth
and seventh cervical vertebrae. Rommel (1990)
also noted variability in fusion of the cervicals but
attributed at least some of the fusions to
pathological conditions. There will also be
variability when animals of different ages are
compared.
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
10. Olecranon process
This process on the ulna is considered primitive
by outgroup analysis. It is present in ziphiids,
although it is less developed in Berardius than in
Mesoplodon, Ziphius and Hyperoodon (True,
1910; Tomilin, 1967). It is well-developed in
Physeter, baleen whales and many fossil taxa.
The loss of the olecranon (e.g. in Delphinapterus,
Monodon: Tomilin, 1967) must then be con-
sidered a derived feature. However, the olecranon
process appears variable even among the del-
phinids and phocoenids; this may be another fea-
ture subject to reversals, i.e homoplasy.
de Muizon (1988) considered that Orcaella
lacks an olecranon process, as in monodontids.
Our more extensive series of specimens shows
that although the process may be obscure (Fig.
48C), it is generally well-developed (Fig.
48A,B). This is not simply an age dependent
feature, as specimens with an olecranon process
had 9 and 17 dentinal layers in the teeth.
11. Convex facial profile
This is a specialized feature of extant monodon-
tids and some phocoenids (Perrin, pers. comm.).
It is considered a derived feature, as by de
Muizon(1988), Heyning (1989) and Barnes
(1990). de Muizon (1988) and Marsh et al. (1989)
noted that Orcaella has a concave supracranial
profile, as in delphinids, but unlike monodontids.
This argument is weakened by lack of a convex
facial profile in the fossil monodontid Denebola
brachycephala Barnes,1984 and juvenile Mono-
don (Eales, 1950; pers. obs. UBC9467). The latter
point is relevant, given the retention of numerous
juvenile characters in the skull of Orcaella.
12. Premaxillary boss
This is generally recognized (e.g. Rice, 1984;
Heyning, 1989) as a specialized feature of
phocoenids and is so coded.
13. Bony lateral lamina of pterygoid
This is equivocal by outgroup analysis as the
Physeterida (Ziphiidae and Physteridae) shows a
reduction in the lateral lamina overall. The bony
lamina of the pterygoid is present in Platanista.
Consideration of fossil cetaceans demonstrates a
reduction from a bony lamina as the primitive
condition to an increasing development of a
membranous lamina (Oelschlager, 1990; de
Muizon, 1988). More particularly, the bony
lamina is found in fossil platanistoids and in the
Eurhinodelphoidea (de Muizon, 1988), a possible
sister group to the Delphinida. It is coded, with
189
0 where the bony lamina is well developed and 1
where it is reduced (in Phocoena, fide Fraser &
Purves (1962) but not confirmed in the small
series of Phocoena we examined) or absent. This
feature may be particularly variable within the
phocoenids (Perrin pers. comm.).
14. Pneumatised maxillary crest
This is a specialized feature of Platanista and
fossil relatives (Heyning, 1989; de Muizon, 1990).
15. Palatines covered by pterygoids
This is a specialized feature of Platanista
(Heyning, 1989). It is a special case of the more
general trend for separation of the lateral and
medial lobes of the palatine by the pterygoid, also
seen to a lesser extent in some individuals of
Berardius and Kogia (de Muizon, in press), as
well as Orcaella, Globicephala and juveniles of
other genera, e.g. Feresa.
16. Acquisition of lateral lamina of palatine
This character, used to define the Delphinida by
de Muizon(1988), is a specialized feature as-
sociated with excavation of the palatine by the
pterygoids.
17. Nasopharyngeal sac
This is a specialized feature of Platanista. No
analagous structure has been documented in other
odontocetes, although Anderson (1879) noted
what he thought was a rudiment of this structure
in a foetal Orcaella.
18. Pterygoid sinus undivided
Fraser & Purves (1962) illustrated the cranial
sinuses in a variety of odontocetes and used the
variations in its development as a basis for clas-
sification of cetaceans. The undivided sinus was
considered a feature of physeterids and ziphiids
(including Berardius, although that genus was
not included in their summary fig. 22) and
Platanista. This is the primitive condition by
outgroup analysis.
Fraser & Purves (1962), de Muizon (1988) and
Heyning (1989) noted the undivided pterygoid
sinus in Delphinapterus and Monodon. This con-
dition is even more primitive than that in river
dolphins /nia and Pontoporia, in which there are
distinct pre- and post-orbital lobes (Heyning,
1989). It was the basis for placing the monodon-
tids in their own superfamily Monodontoidea
(Fraser & Purves, 1962) and presents difficulties
in interpretation (de Muizon, 1988; Heyning,
1989). The latter author, while retaining the
monodontids in the superfamily Delphinoidea,
commented ‘I am perplexed as to how such a
primitive air sinus system could have evolved
within the Monodontidae’. de Muizon (1990),
who considered both fossil and extant taxa, noted
that if the condition in monodontids was con-
sidered primitive, then reversals would have to be
invoked in the Eurhinodelphoids, Lipotes, Pon-
toporia, Inia and the delphinoids. As with all
previous authors, we have had access only to
skeletal material, and have inferred the air sinus
distribution from this. Although the lateral lamina
of the pterygoid makes a particularly convincing
boundary in Monodon (pers. obs.; see also Pilleri
et al., 1982, figs 2b, 3), the extent of the lateral
lamina in Delphinapterus is more variable. Pseu-
dorca shows comparable variation in the
development of the bony lateral lamina of the
palatine. Purves & Pilleri (1978, fig. 18) il-
lustrated considerable variation in the cranial
sinuses of Pseudorca, apparently associated with
the variation in the bony lateral lamina so there
may be comparable variation in Delphinapterus.
In addition, dissections of specimens have not
always confirmed the pattern of sinuses inferred
by Fraser & Purves (1962) from the skull, e.g.
re-examination of [nia and Pontoporia by Heyn-
ing (1989) resulted in different interpretations of
the anterior sinus. It is thus highly desirable that
the condition of the cranial sinuses in both Del-
phinapterus and Monodon be re-examined by
dissection of fresh material. In the absence of
such information, we accept the undivided con-
dition of the pterygoids in monodontids as sug-
gested by previous workers and supported by our
examination of skulls.
19. Coalescence of pre- and post-orbital lobes of
pterygoid sinus to surround optic nerve
Fraser & Purves (1962) documented this as a
feature of the delphinids such as Tursiops, Gram-
pus, Stenella, and Delphinus. Based on their
work, deMuizon (1988) used the coalescence of
the sinuses as the basis for the subfamily Del-
phininae, within Delphinidae. This is a special-
ized feature. Coding is - for those genera which
have undivided sinus , 0 for well-separated pre-
and post-orbital lobes and 1 for coalescence.
20. Dorsal extension of pre-orbital lobe of
pterygoid sinus.
This is a specialized feature of the phocoenids
(Fraser & Purves, 1962; Heyning, 1989), but is
not shown uniformly throughout the family (Per-
rin, pers. comm.). There is an incipient extension
MEMOIRS OF THE QUEENSLAND MUSEUM
of this lobe in Orcaella (pers. obs.), as already
noted by Fraser & Purves (1962) but this is coded
as absent in the present analysis, as delphinids
such as Pseudorca show a similar but less ad-
vanced condition. Codes are - for undivided
sinus, O for limited extension and 1 for well-
developed extension of the lobe.
21. Anterior sinus
This is equivocal by outgroup analysis, as the
cranial sinus is undivided in ziphiids and
physeterids. Monodon and Delphinapterus
would also be coded - as their sinus is undivided.
Fraser & Purves (1962) indicated an anterior
sinus in /nia, based on skull morphology, but
Heyning (1989) could not confirm this by dissec-
tion. /nia is thus coded as 0 (anterior sinus ab-
sent) in this analysis. In the absence of contrary
information, the configuration indicated by
Fraser & Purves (1962) has been accepted. Elon-
gation of the anterior sinus is considered the
derived state (Fraser & Purves, 1962).
22. Curvature of flippers
The flippers of mature males of beluga
(Vladykov, 1943) and narwhal ( Reeves &
Tracey, 1980; Martin, 1990) curve upwards at the
anterolateral margin. Although this seems a
minor character, it is consistent. The deformation
of the flipper involves a curvature of the second
and third digits (Vladykov, 1943), and may be
associated with the relatively larger amount of
connective tissue to bone in the flippers of beluga
and narwhal than in delphinids. It is a specialized
feature of these two genera.
Mature males of both species have a strongly
convex posterior edge to the flukes (Vladykov,
1944; Leatherwood & Reeves, 1983); in narwhals
the anterior margin may curl forward so that the
flukes appear to be ‘back to front’. Although there
appears to be similar fluke form in the two
species, this is not treated as a separate character.
23. Cervical sinus
The indentation of the nape to form a neck-like
constriction is a specialized feature, found in
Platanista (Anderson, 1879), Inia (Hoyt, 1984:
68), Neophocaena (Hoyt, 1984:75), Orcaella and
some ziphiids (Perrin pers. comm.).
24. Anterolateral margin of nares formed by max-
illaries
This is a specialized feature of extant and fossil
monodontids (de Muizon, 1988; Barnes, 1990).
de Muizon (1988) pointed out that it is not shared
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA 191
by Orcaella, which has the small maxillary in-
trusions characteristic of many delphinids.
25. Spatulate teeth
This is a specialized feature of phocoenids
(Rice, 1984; Heyning, 1989), but variably de-
veloped within the family (Perrin pers. comm.).
26. Length of squamosal
Heyning (1989) considered 2 characters as-
sociated with the temporal region: roofing over
by the frontal bones and length of the squamosal.
The length of the squamosal has generally been
considered ‘short’ or ‘long’. Examination of a
range of genera (from illustrations in the literature
as well as examination of skulls) suggested that
they form 3 categories : 0-30%, 30-50% and
>50%, which are coded as 0, 1 and 2 respective-
ly. Polarity is equivocal by outgroup analysis:
Physeter is 1 (Miller, 1923), Berardius 0
(Tomilin, 1967) and the river dolphins 2. The
character is treated as non-additive so no direc-
tion of specialization is implied. Heyning (1989)
characterized the zygomatic arch of Orcaella as
‘extremely reduced’, a feature he considered a
synapomorphy of the delphinids. However, it is
about 35% of the cranial vault length in Orcaella
(pers. obs.) and 39-42% in Neophocaena. This
approaches the values for Monodon (c. 44%) and
is much higher than that for delphinids such as
Tursiops (21-24%) or Sousa (23%). Roofing of
the temporal region is discussed later (character
39).
27. Disappearance of superior lamina of ptery-
goid in orbital region
This is equivocal by outgroup analysis; it was
considered an advanced character by Fraser &
Purves (1962). This is consistent with the reduc-
tion of bony margins to the cranial sinuses as
indicated by Oelschlager (1990). de Muizon
(1988) noted the strongly developed superior
lamina of the pterygoid in Platanistoidea and the
Eurhinodelphoidea. The superior lamina is thus
considered primitive while its disappearance is
treated as the derived condition in this analysis.
28. Orientation of posterior process of periotic
The process is oriented strongly ventrad
towards the posterior process of the tympanic or
posteriorly, except in delphinids where it is
posterolateral to strongly lateral (Kasuya, 1973).
The latter is considered specialized (coded 2);
ventral orientation (Platanista) is coded as 0;
posterior orientation as 1.
29. Orientation of posterior process of tympanic
Polarity is equivocal by outgroup analysis; the
posterior orientation was considered primitive by
Kasuya (1973) and is accepted as such here. We
observed the orientation of the process in Orcael-
la to be posterolateral, not posterior as indicated
by previous authors (e.g. Kasuya, 1973).
30. Sigmoid process of tympanic bulla
Four forms of the sigmoid process are recog-
nized, based on Kasuya (1973) and Pilleri et al.
1989: square (Physeter), flail-shaped (Berar-
dius), short (river dolphins) and long & L-shaped
(all others). In our analysis, the short sigmoid
process of the river dolphins was coded 0 and the
L-shaped process 1. No direction of specializa-
tion is implied and, in any case, the character
reduces to a binary feature in the present analysis.
31. Lateral furrow of tympanic bulla
The polarity of this feature is equivocal by
outgroup analysis: it is absent in Physeter (and
Kogia), but present in ziphiids and the river dol-
phins. It is generally present in fossil odontocetes,
including most physeteroids (de Muizon, 1988).
It is considered primitive, as by Kasuya (1973).
32. Width of posterior branch, lower tympanic
aperture of bulla.
The lower tympanic aperture is not present in
physeterids, but is less than 1046 standard length
of the tympanic bone in Berardius and all river
dolphins except Pontoporia. The branch is over
10% in Delphinapterus (Kasuya, 1973, based on
very small sample size) and phocoenids. The
wide branch is considered a specialization, by
outgroup analysis, but this is equivocal (see
below). Although boundaries set on continuous
variables will always be more or less arbitrary,
the value of 1046 did appear to be relatively well
defined. It also is consistent with use by Kasuya
(1973), whose study on tympanoperiotic bones
has been very influential. Coding is 0 for « 10%
and 1 for >10% standard length of tympanic.
A lateral shift of the posterior process will
compress to some extent the posterior branch of
the lower tympanic aperture. Thus there may be
a purely structural reason for the fact that taxa like
phocoenids and beluga, with a posteriorly
directed posterior process, have a wide aperture
while the delphinids, with postero-lateral to
lateral process of the tympanic, have a narrower
aperture. Thus this character may not be inde-
pendent of character 29, and it could be argued
that the primitive condition was a wide aperture,
192
associated with a posteriorly directed process
(considered the primitive condition, see character
29).
In Orcaella the LTA width is <10% (see sec-
tion on Tympanoperiotic bones).
33. Compression of bulla
Compression of the bulla (width 50% or less of
standard length of tympanic bulla) occurs in
Platanista, Monodon and Globicephala. Lack of
compression is considered primitive by outgroup
analysis.
34. Sutural connection of tympanoperiotic to
skull
Heyning’s (1989) interpretation is accepted
here, There is a sutural connection in ziphiids and
physeterids. This is considered primitive.
Platanista shows an intermediate condition, with
partial fusion to the skull, while all others con-
sidered show no sutural connection to the skull.
This feature may be subject to some variation
within species. Kleinenberg et al. (1964) and
Kasuya (1973) reported that the periotic was su-
tured to the skull in Delphinapterus, but neither
Heyning (1989) nor we found this to be the case
on belugas we examined (see discussion of tym-
panoperiotic bones for further details).
35. Retraction of premaxillaries from nasals
The anterior retraction of the premaxillaries is
a specialized feature of phocoenids ( de Muizon,
1988; Heyning, 1989; Barnes, 1990). The river
dolphins supposedly show a gradation of lateral
retraction of the premaxillaries, but this is not
coded here because it is difficult to quantify and
appears to overlap with retraction in some del-
phinids. The general condition in which the left
premaxilla is retracted away, while the right
premaxilla maintains close contact with the
nasals, is considered a feature of the delphinids,
as by previous authors (e.g. de Muizon, 1988;
Heyning, 1989; Barnes, 1990), but it is variable.
36. position of attachment of tympanoperiotic to
skull
In Delphinapterus, Monodon and Orcaella, the
dorsal surface of the posterior processes of
periotic and tympanic bones are involved in the
connection to the mastoid portion of the
squamosal. In the phocoenids, which also have a
large mastoid portion of the zygomatic arch, and
the delphinids, where the mastoid portion is com-
pressed by the exoccipitals, the processes of the
tympanoperiotic join in a cavity formed by the
MEMOIRS OF THE QUEENSLAND MUSEUM
basioccipital, exoccipital and squamosal, more
dorsally placed than in the three genera just men-
tioned. The attachment seems to be a distinctly
different form in ziphiids and physeterids, how-
ever the more ventral attachment is similar to the
situation in Platanista. The type of attachment in
Delphinapterus, Monodon and Orcaella is thus
considered primitive.
37. Profile of tympanic bulla
de Muizon (1988) and Pilleri et al., (1989)
indicate that the bulla in outer lateral view may
have a convex or concave profile. The former
characterizes Physeteridae, Ziphiidae and all the
river dolphins; it is thus primitive by outgroup
analysis. Monodontids, phocoenids and del-
phinids have a concave profile.
38. Shape of anterior process of periotic
de Muizon (1988) listed a square, almost rec-
tangular, profile of the anterior process as a char-
acter of the Delphinoidea (Monodontidae,
Phocoenidae, Delphinidae) and said that it was a
consistently recognizable character. Kasuya
(1973), however, was less clear in this separation.
He noted that the process was curved and rod-like
in Physeter and Platanista, pyramidal in ziphiids
(although elongate in Berardius), and flat & rec-
tangular in Inia, Pontoporia, Delphinapterus and
Monodon, phocoenids and delphinids. Kasuya’s
scheme is used here. There may be some varia-
tion, however, in the form of the anterior process
in Monodon: UBC9467 has an elongate, curved
process.
39. Roofing of temporal fossa
The outgroup analysis is equivocal in defining
polarity. In Physeter and Berardius it is roofed
over, but in some fossil physeterids (e.g.
Diaphorocetus: Kellogg, 1928) and various fossil
taxa (Agorophiidae, Squalodontidae) as well as
Kogia (Perrin pers. comm.) and the river dolphins
it is open, exposing the wide zygomatic arches.
Following Heyning (1989), this is considered the
primitive state, with roofing of the temporal
region derived. It should be noted that the reason
for considering the feature primitive is not be-
cause it occurs in a fossil species, but because it
is present in at least some representatives of the
outgroup, which just happen to be fossil species.
CHARACTERS NOT CONSIDERED
Heyning (1989) included a number of charac-
ters specific to ziphiids (e.g. throat grooves,
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
elevated vertex) or physeterids (e.g. spermaceti
organ, distal sac, lack of one nasal bone). Since
the inter-relationships of the Physeteridae and
Ziphiidae are not under examination, and these
characters are irrelevant to the Delphinida (sensu
de Muizon, 1988), they have not been included.
Additional characters not considered are listed
below.
1. Bicipital ribs
Slijper (1936, 1962) considered an elevated
number of bicipital ribs as a primitive condition.
He envisioned that reduction of bicipital ribs was
linked to a need for flexibility of the thorax in a
diving mammal (Slijper, 1962). This argument is
weakened by the finding of Rommel(1990) that
the elastic ligaments allowed significant flexure
of the bicipital ribs in Tursiops. The number of
bicipital ribs (7-8/12-13 ribs) in Orcaella (see
Appendix 4) is higher than in most delphinids,
comparable to Delphinapterus (8/11-12: Slijper,
1936) but lower than in phocoenids (11/13-14 in
Neophocaena; 10/12-14 in Phocoena: Nishi-
waki, 1963). Insufficient information was avail-
able to use this character.
2. Lip on nasal plug
Schenkkan (1973) illustrated the increasing
development of a lateral lip on the plug from
Kogia, Pontoporia, ziphiids through to Phocoena
and delphinids. Insufficient information was
available for other genera to use this character.
Similarly, there was insufficient information to
assess any patterns in the extension of the melon
into the right or left nasal plug.
3. Proportion of premaxillary sac to vestibular
sac
Schenkkan (1973) documented a gradation
from large vestibular sacs in Pontoporia, Inia and
Phocoena to small sacs in Delphinus and Stenel-
la. Conversely, the premaxillary sacs assumed
greater relative importance from the river dol-
phins to Delphinus and Stenella. We could not
calculate values for monodontids or some of the
delphinids not considered by Schenkkan, so have
not included the character.
4. Coat of cement on tooth
Lonnberg (1910) contrasted the teeth of Del-
phinapterus with those of Phocoena, Steno,
Globicephala and Delphinus. The former dif-
fered from the porpoise and dolphins examined
in the strong development of a cement coat, con-
stituting the bulk of the tooth. As pointed out by
Lonnberg, a similar tooth structure occurs in
Physeter, suggesting that this may be the primi-
tive condition. It is not associated with the size of
the teeth; Orcinus has a typical delphinid tooth.
193
The cement coat on teeth of Kogia may be exten-
sive, building up to form a convex profile to the
tooth in old animals of both species in the genus
(Ross pers. comm.). Among ziphiids, the situa-
tion is more variable: in Hyperoodon and Ziphius
most of the tooth is cement, but dentine predom-
inates in teeth of Berardius (Ross pers. comm.).
Platanista has a substantial cement component in
the teeth, but Kasuya (1972) noted that it 1s dis-
tinct from the pattern in Physeter. A cement coat
extends the entire length of the erupted tusk in
Monodon (Reeves & Tracey, 1980) but it is un-
clear if it is as well developed as in Delphinap-
terus or Physeter. There were insufficient data to
establish polarity of the character, either by out-
group analysis or other arguments. A similar
situation exists for accessory cusps, which occur
in juvenile Delphinapterus (Stewart & Stewart,
1989) and are sporadic in other odontocetes.
5. Phalangeal formula and flipper shape
de Muizon (1988) noted that there is a greater
number of phalanges in digits 2 and 3, relative to
the other digits in delphinids, whereas there is
greater uniformity in monodontids. However, a
comparison of phalangeal formula for Del-
phinapterus, Monodon and various dolphins sug-
gest there is considerable overlap. In particular,
the formula is similar in Orcinus (1:2; II:6-7;
III:4-5; IV: 3-4; V: 2-3: Nishiwaki, 1963)to the
monodontids (Monodon: 1:1-2; II: 5-8; III: 4-6;
IV:2-4; V: 2-3; Delphinapterus: 1:1-2; IT: 6-9; III:
4-5; IV:2-4; V: 2-4: Nishiwaki, 1963). Eales
(1953) noted that phalangeal number decreases
with age in the narwhal and beluga; the curvature
of the flipper in adult males may be related to this
fact. However, it can not be used as a character
for the monodontids, as this reduction was also
found in Phocoena (Eales, 1953). The flipper
does tend to be more squared-off in Physeter,
ziphiids, platanistids and monodontids, than it is
in phocoenids and delphinids, but there is varia-
tion in form among the latter (e.g. Orcinus, a
delphinid, has broad paddle-shaped flippers espe-
cially developed in the male).
6. Position of delto-pectoral tuberosity
de Muizon(1988) considered the distal position
of this tuberosity a feature of delphinids and
phocoenids. He further indicated that the
tuberosity was subdistal in Orcaella, linking it
more with the monodontids. On our specimens
(Fig. 48 A,B,C), the tuberosity was elongate and,
while not distal, it was not as clearly subdistal as
figured for Delphinapterus and Monodon by de
Muizon (1988, fig. 22).
194
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 6. External morphometrics for 22 specimens of Orcaella brevirostris from central Queenland. Numbers
in the left hand column refer to the following measurements. Values for 2-17 are percentages of total length.
1. Total length (m); 2. Tip of upper jaw to eye; 3. Tip of upper jaw to gape; 4. Tip of upper jaw to ear (auditory
meatus); 5. Length eye to ear; 6. Tip of upper jaw to anterior base flipper; 7. Tip of upper jaw to blowhole; 8.
Tip of upper jaw to tip of dorsal fin; 9. Tip of upper jaw to umbilicus; 10. Tip of upper jaw to genital aperture
(centre); 11. Tip of upper jaw to anus; 12. Length of anterior margin flipper; 13. Length flipper from tip to axilla;
14. Maximum width flipper; 15. Height dorsal fin; 16. Length of dorsal fin base; 17. Width of tail flukes.
1.34 | 1.86
ee a
[ 1 [o9 | 1.05 | 134 |
5|6/|,7]|8]|9,]|10|1
1.87
[670 | 963 | 110 | - | 6.10 | 6.10 | 6.80 |
| 6.60 |
6.10
4.30
12.30
6.80
5.60
5
| 6 |z. : 60 | 23.00 | 17.50] -
7 | 9.78 | 11.90] 11.50] 6.10 | 1016| 760 |. - |
| 8 | 57.14 | 60.00 | 58.20 | 57.50
| 9 [4780|4860|4450| - [4490] - |
10 | 63.19 | 55.20 | 60.70 | 60.50 | 60.40]. - p
64.20
12.60 13.70
6.60
4.00
13.90
6.40
4.30
3.77
6260| - | - | 55.20] 59.40 | 54.80 | 57.00 | 54.90 | 58.10 | 57.60 | 63.00
56.60
| 11 [6560|6670|64.40|6420|6420| - | - [66.50] 60.80 | 66.50 | 66.50 | 66.50 | 64.20 | 67.70 | 64.80 |
| 12. |2033 | 20.00] 17.40 | 16.40 | 17.10 | 17.10 | 15.10 | 17.90 | 15.60 | 15.90
14.40
18.60
6.50
18.30
11.40
19.50
6.50
5.70 11.20
44.20 | 42.30 | 44.70 | 42.40] - |
61.60 | 54.90 | 54.40 | 63.60 | 54.34
44.50
5240) -
| 16.70 | 18.80 | 19.80 | 16.60 | 16.00 |
19.80 | 16.60
14.90 | 16.70 | 16.30
13.70 | 15.40
16 | 7.69 - 7.50 | 9.60
21.40 | 29.00 | 25.10 | 23.70
7. Impression of ‘duplicated apex of nasofron-
tal sac' on skull
de Muizon (1988) documented fosetta or shal-
low depressions on the anterior face of the brain
case above the nares in phocoenids and
monodontids. He considered these the impres-
sions of a duplicated apex of the nasofrontal sacs
(apparently the same as the 'posterior nasal sac'
of Heyning, 1989). We noticed similar fosetta in
Orcaella but there was no indication of a
posterior nasal sac in our dissections of Orcaella
(see Facial Anatomy). We therefore reject the
occurrence of such fosetta as evidence for a
posterior nasal sac.
8. Development of mesethmoid plate and
elevation of nasals
de Muizon (1988) and Barnes (1990) linked
these two characters, which they considered diag-
nostic of delphinids. Orcaella examined herein
have a poorly developed mesethmoid plate,
which does not impinge on the reduced nasals.
However, the nasals are still elevated on the ver-
tex, suggesting that the characters are not linked.
Elevation of the nasals may be a more consistent
character for the delphinids, although there are
similarities in the position of the nasals in
Monodon, Delphinapterus and the delphinid
26.40 | 33.20 | 30.20 | 32.30 | 30.70 :
Pseudorca. These characters are supporting fea-
tures of the delphinids but are not incorporated in
the analysis, because of their variability (even
between populations of Orcaella in the case of
the development of mesethmoid plate).
9. Compression of vertebral centra
de Muizon (1988) suggested that delphinids
showed a tendency for compression of the ver-
tebral centra. This is linked to higher numbers of
vertebrae, and possibly a greater flexibility of the
spinal column. Low numbers of vertebrae (under
60) does appear to be a primitive character (see
Table in Watson, 1985), but quite unrelated
species show elevated numbers of vertebrae (e.g.
Phocoenoides among phocoenids, Lissodelphis
and some Lagenorhynchus in delphinids). Due to
this variability, the feature was not included.
Similarly, de Muizon (1988) suggested that the
Phocoenidae and Delphinidae have more elon-
gated transverse processes of the lumbar ver-
tebrae, contrasted with more triangular processes
in other Delphinida, such as river dolphins (ex-
cept Platanista) and Delphinapterus. However,
there again seems to be much variation in del-
phinids. The transverse processes of a North At-
lantic Globicephala melas are intermediate
between those of Delphinapterus and delphinids
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
TABLE 6 (continued)
fe atte [20 fa fe |
2.25
6.40 9.10
|
1 12. 4
| - | 400] - | 3.50 |
| 18.50| 19.60 | 17.30 | 19.90
51.30 | 53.20
11 [64.40 | 65.20/66.20| - [45.80 | 54.00 64.70)
| - |19.10| 16.70] 16.90] 17.80 | 17.30 | 15.70 |
12.80
| - | 750 | 580 | 7.10 | 6.70 | 6.60 | 6,20
— pim 4.40
820. 6.30
8.90
i
such as Lagenorhynchus acutus with elongate
processes (pers. obs.).
10. Palatine bridge under optic channel
de Muizon (1988) considered this a feature of
the monodontids; we observed it in beluga and,
even more strikingly so, in narwhal but have not
used it as a character because posterior develop-
ment of a palatine wing is not restricted to
monodontids. It is equally developed in the del-
phinid Pseudorca, where the lateral boundary is
formed exclusively of palatine bone which ex-
tends almost to the squamosal. The posterior ex-
tension of the palatine, in combination with the
more anterior position of the orbit in monodon-
tids, is the basis for the obvious bridge underlying
the optic channel in Delphinapterus and
Monodon. It is particularly striking in the latter
because of the strong depression of the skull in
that genus;this ensures that the palatine wing is
closely appressed to the optic channel. Orcaella
represents the opposite extreme where the
posterior extension of the palatine extends out-
ward, well away from the basicranial bones, as a
free wing-like structure.
Since we consider the palatine bridge to be
associated with the more anterior position of the
17
195
orbits (character 4), we do not include the palatine
bridge as a separate character.
11. Immunological and electrophoretic dis-
tance
Lint et al. (1990) demonstrated a close relation-
ship of Monodon with Delphinapterus, based on
immunological distance; Orcaella was well
separated from these genera and grouped with the
delphinids, Orcinus and Lagenorhynchus. Inia
was closer to the monodontids than delphinids,
but was not closely linked to either. These
relationships were supported by electrophoretic
data, and combined with the latter to give a
phylogeny of odontocetes (Lint et al., 1990: fig.
2d) in which delphinids were widely separated
from phocoenids. These data have been discussed
elsewhere, but were not used in our analysis
because there were insufficient data for all
species.
12. Base pair length of repetitive DNA
Gretarsdottir & Arnason (1992) compared the
base pair lengths of highly repetitive DNA in a
variety of odontocetes. Representatives of most
families examined, including beluga and
narwhal, had a primitive base pair length of about
1750 bp. The delphinids, however, had a bp
length of about 1580, while Orcaella had a base
pair length of 1583.
These data support Orcaella being a delphinid,
but could not be incorporated in the analysis
because comparable data were not available for
other species; most importantly there were no
data for any of the river dolphins.
13. Isovaleric acid and acoustic fat
Litchfield et al. (1975) compared odontocetes
for distribution of lipids in both acoustic (melon
and mandible fatty bodies) and non-acoustic (e.g.
blubber) tissues. They separated two groups:
ziphiids, physeterids and ‘Platanistidae’
(Platanista,Inia) without isovaleric acid in the
acoustic tissue, and the Monodontidae, Pho-
coenidae and Delphinidae with isovaleric acid.
They further subdivided the latter group, with
delphinids having over 3% of the lipids as waxy
esters, while the monodontids and phocoenids
had exclusively triglycerides.
Morris (1985) argued that isovaleric acid can
not be considered a by- or waste product of stand-
ard physiological processes but represents a
specialization, produced at considerable
physiological cost. Presence of isovaleric acid
has been linked to development of the melon, and
the function of fat in the melon as an acoustic lens.
As pointed out by Litchfield et al (1975), some
other mechanism must be used by the river dol-
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 7, External morphometrics for 11 specimens of Orcaella from SE Asia, Numbers in the extreme left
hand column are measurements, as in Table 6. Specimen numbers 1, 3, 5, 7 & 10 are from Anderson (1879);
2, 6, & 9 from Tas'an et al., 1980; 4, 8 & 11 from Pilleri & Gihr, 1973-1974.
1 2 3
l 086 | 153 | 180
| 2 | - | 980 | - | 88 | - |
8.00 73
12.80 9.80 12.10
59.20 56.00 | 55.80
22.20 | 14.50 20.60
| - | 73 | - [sse | - | - | 64 | - |
2220 | 2120 | 2000 | 1990 | - |
Ree ee EC ELEM
[| - | - | - | - |
REET i - | - | - [8w]
| 330 | 350 | - | - | 420 | - |
23.9° , 20.80
| oso | - | - | 1090 | 11.10
5740. | 60.40 | 61.90
lao
Fes | - | --
| - 7320| 67.60.
66.50"
- |-
- - 74.50
9.69
jotes:- 1. to anterior of base; 2. curved along back; 3. un
phins, but distribution of isovaleric acid suggests
that itis a derived feature of the more specialized
delphinoids. It was not used in the analysis be-
cause of missing values for a number of species.
14. Pseudaliid nematode parasites
At least 28 nominal species of pseudaliid
nematodes have been reported from the cranial
sinuses and respiratory tract of odontocetes and
they are perhaps the most promising group of
parasites to use in assessing inter-relationships of
the hosts.
Except for Delamurella Gubanov, 1952 in
Berardius from the northwest Pacific, pseudaliids
are restricted to the Delphinida sensu deMuizon
(1988). This record is apparently based on a
single report; more information is necessary to
evaluate the relationship of Delamurella to other
pseudaliids.
The phocoenids have the widest range of
pseudaliid parasites: P. phocoena has 5 species,
Phocoenoides dalli 6 and Neophocaena
phocaenoides 5 nominal species, with limited
overlap of parasites between host species.
Pseudaliusis apparently restricted to phocoenids
(but see anomalous records below). Arnold &
Gaskin (1975) maintained Torynurus Baylis &
Daubney, 1925 distinct from Pharurus; at that
26.10
by subtraction of published values; 6. in centre of flipper; 7. *depth through center'; 8. indirectly calculated; 9
to anterior of eye; 10. 'anterior of base to posterior of tip'.
BI RED ETE
alt x
9.50 7.03
2.40
2.40
10.40
10.40
530 | 6.60
- | 2390 |
27.30
specified flipper length; 4. to middle of fin; 5. calculated
time the genera were known from phocoenids
(Phocoena, Phocoenoides) and monodontids
respectively. Descriptions of additional species
from Neophocaena and Phocoenoides (Petter &
Pilleri,1982; Kuramochi et al.,1990) have blurred
distinctions between these genera. The diagnostic
characters of Pharurus, Torynurus, Stenurus and
Pseudostenurus need to be re-evaluated; study of
new material of Pseudostenurus from
Neophocaena is crucial to such a review. The
species of Halocercus in Phocoenoides and
Neophocaena also need review.
If the restricted definition of Pharurus as
proposed by Arnold & Gaskin (1975) is con-
firmed, then this genus is restricted to monodon-
tids ( P. pallasi in Delphinapterus, P. alatus in
Monodon). The other genera reported from
monodontids (Stenurus, Halocercus) have also
been reported in phocoenids and delphinids.
Identifications of pseudaliids in delphinids are
often incomplete and poorly documented. Del-
phinus delphis has the largest number (4) of
pseudaliid species reported from any delphinid
host, all in Halocercus or Skrjabinalius; some of
these species require confirmation. Halocercus
and Skrjabinalius have the widest reported dis-
tributions within the Delphinidae, having been
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
reported from Delphinus, Stenella, Tursiops,
Lagenorhynchus, Sotalia, Sousa, Peponocephala
and Cephalorhynchus. Skrjabinalius is known
only in the Delphinidae. Distinctions between the
species of Halocercus in particular are often
poorly documented and many records need con-
firmation. Stenurus ovatus has been recorded in
Tursiops and Lagenodelphis, while S.
globicephalae has been reported from
Globicephala melas, G. macrorhynchus,
Lagenorhynchus acutus, Grampus griseus,
Peponocephala electra and Feresa attenuata.
Unidentified species of Stenurus have been
reported from Cephalorhynchus hectori and Lis-
sodelphis peronii. No pseudaliids have been
reported from Orcinus or Orcaella, nor have we
found any pseudaliids in the Orcaella we necrop-
sied.
Analysis at species level is complicated by the
probability of misidentifications in the literature,
as noted above. For instance, Stenurus minor has
been reported from Delphinapterus leucas,
Phocoena phocoena, Phocoenoides dalli and
Grampus griseus. The last record is based on a
single report by von Linstow (1910). This pre-
dates the description of Stenurus globicephalae,
which is well-known from various blunt-headed
genera, including G. griseus (Arnold & Gaskin,
1975; paragraph above). Additional ques-
tionable or anomalous records include T. con-
volutus and Pseudalius inflexus in Lageno-
rhynchus acutus and Pseudostenurus auditivus in
Pseudorca.
Until necessary revision is done on pseudaliids
and questionable records evaluated, the distribu-
tion of pseudaliid parasites in odontocetes offers
little help in assessing inter-relationships of the
hosts.
APPENDIX 3:
EXTERNAL MORPHOMETRICS
RESULTS
General features correspond well with descrip-
tions of Orcaella brevirostris from Anderson
(1879), Lloze (1982) and Marsh et al. (1989).
Among 37 animals from central Queensland,
the longest d is 2.70m , the next longest being
2.35m. The longest 9 is 2.30m; two animals of
unknown sex are 2.35 and 2.49m long.
Body proportions are available for a smaller
sample (19 animals 1.86-2.35m and 3 animals
0.91-1.34m long). These are compared with
measurements in the literature (Tables 6 & 7).
197
There are anomalous values for animal 21 (Table
6); the measurements are not used in the calcula-
tion of the descriptive statistics.
Dimensions of the head (e.g. rostrum to eye,
rostrum to flipper, rostrum to gape, rostrum to
auditory meatus) are larger in the small animals
(0.91-1.34m total length). The only clear sexual
dimorphism is in the more posterior position of
the genital aperture in 9 9 (61.6-63.6% total
length, plus one anomalous value of 54.9%) than
in males (52.4-56.6%). Even with the anomalous
value included, the difference is statistically sig-
nificant (596, Mann-Whitney U test). There was
almost total overlap between sexes for all other
characters.
There is considerable variability in the meas-
urement of the dorsal fin base, which is difficult
to clearly define. The wide variation in fluke
widths may reflect post-mortem shrinkage in
some of the stranded animals, as well as errors
introduced by calculating total width by ex-
trapolating from one undamaged fluke.
DISCUSSION
Our data and other records (e.g. Pilleri & Gihr,
1973-1974) suggest that dd grow larger than
9 9, but the numbers of animals examined is
small. More material is necessary to confirm the
possible sexual dimorphism in body length.
Morphometrics for 11 SE Asian specimens
were collated from the literature (Table 7). For
our comparisons, we do not separate O.
‘fluminalis’ and O. brevirostris. Smaller animals
(0.91-1.34m long) are considered separately.
Animals from Indonesia measured by Tas'an et
al. (1980) had much higher values for the meas-
urement 'tip of upper jaw to the umbilicus' than
all animals from Queensland. This feature was
not routinely measured by other authors, however
Lloze (MS) noted that in two specimens 1.9 and
2.0m long from the Mekong, the umbilicus oc-
curs ‘5-6cm in front of the midlength ofthe body’.
This gives a figure of approximately 47% total
length, which is consistent with the Queensland
sample. Thus the values reported for Indonesian
animals do not necessarily indicate variation be-
tween Queensland and SE Asian animals.
Some measurements appear to be taken in dif-
ferent ways. The distance between upper jaw and
blowhole will vary according to whether it is
taken in a direct line (using calipers) or curved
over the body. The measurement of the dorsal fin
base also seems to be subject to wide variation.
Published values for flipper length (Table 7) were
measured along the anterior margin, through the
198
TABLE 8. Length-weight data for Orcaella
brevirostris, combining original data from
Queensland with literature records. The record from
Anderson, 1879 was a near term foetus from India.
The records from Tas’ an et al., 1980 were consecutive
weighings of acaptive Indonesian animal. Records of
Lloze were from the Mekong River. *= the animal
was weighed in pieces, so weight is underestimated.
SOURCE
Anderson, 1879
Tas'an et al., 1980
Tas'an et al., 1980
original
Tas'an et al., 1980
Lloze in Marsh et al., 1989
Lloze in Marsh et al., 1989
*original
*original
Marsh et al., 1989
centre of the flipper or at an unspecified location.
In spite of these inconsistencies, there is general
agreement in proportions of animals over 1.86 m
long, although the 2.75m male from Thailand
(Bonhote in Pilleri & Gihr, 1973-1974) had
anomalously high values for rostrum to flipper
and rostrum to dorsal fin, as well as a low value
for fluke width. Height of the dorsal fin is consis-
tently higher (4.4% (3.4-5.3%), n=18) in
Queensland animals, compared with the SE
Asian animals (2.9% (2.3-4.5%), n=9); this is
highly significant (p =0.002%, Mann-Whitney U
test). The ‘height’ of the dorsal fin in a 2.29 m ó
(Anderson,1879) was 7.03% (Table 7), however
the measurement was taken from the anterior of
the base of the fin to the posterior tip of the fin.
This implies that it was an oblique measurement,
not comparable to the standard height, and the
value is not included for comparison. The dorsal
fin of Qld animals appears higher, with a more
convex anterior margin and shorter base than
animals from Kalimantan held at Jaya Ancol
Oceanarium (based on photos taken by Dr A.
Preen). The posterior margin of the fin of Qld
animals is also much straighter (Fig. 3) than the
distinctly emarginate border illustrated by Ander-
son (1879, pl. 25, fig. 4), and is closer to the 2.2
m d O. ‘fluminalis’ illustrated by that author (pl.
25a, fig. 1). In Qld specimens the tip of the fin is
usually acute, however it could be rounded as in
some SE Asian animals (e.g. Lloze, 1982, Fig. 1).
MEMOIRS OF THE QUEENSLAND MUSEUM
General accounts (e.g. Morzer-Bruyns, 1971;
Sylvestre, 1993) of the Irrawaddy Dolphin give
the average weight as 100 kg; Martin (1990)
noted that ‘typically’ weight was 90-150kg.
There appear to be very few published values of
weights (Table 8). The weights of three animals
from Qld between 2.14-2.25m long varied be-
tween 114-133kg.
These are underestimates as the animals were
weighed in pieces, with loss of body fluids and
blood. Another animal 2.17 m long, also weighed
in pieces, was reported to weigh 190 kg (Marsh
et al., 1989). This value was confirmed from the
original data sheets, but in light of other measure-
ments, appears to be too high. More data are
needed to provide an accurate length-weight
relationship.
APPENDIX 4: POST-CRANIAL SKELETON
AXIAL SKELETON
The vertebral formula for 6 central Qld
specimens is C7, T12-13, L9-12, Ca28-30 = 58-
60, with 17-20 chevrons and up to two nodules
below the caudals. In JM511, there is a small
tubercle on the last caudal vertebra; if this repre-
sents a fused vertebra the total count becomes 61.
CERVICAL VERTEBRAE (Fig.49)
The first 2 cervical vertebrae are fused. The
combined neural processes of C1-C2 are usually
bifid at the top (Fig. 49). The transverse proces-
ses of C1-C2 are low tubercles, barely extending
beyond the condylar facets in anterior view (Fig.
49A). The condylar facets are well separated
ventrally.
Diapophyses and parapophyses are developed
to a variable degree on cervicals 3-7; in QMJM
4729 (MM92) they are obvious only on cervical
4 (Fig. 49A,B). The last cervical vertebra is
distinct in its high neural process, well developed
transverse processes and costal facet on the
centrum.
THORACIC VERTEBRAE (Fig. 50)
The anterior eight thoracic vertebrae are of
similar width; the transverse processes increase
in width from T8, with the last thoracic vertebra
distinctly wider than the others. On several
specimens the anterior border of the transverse
processes of the last thoracic vertebra has a dis-
tinct antrorse spine (Fig. 50B). In QMJMS11, the
transverse processes of the last (12th) thoracic
vertebra have distinct facets for articulation with
the ribs. MM333 has 13 thoracic vertebrae, each
with a fossa or swelling of the tip of the transverse
processes.
The first metapophysis appears on T7-8 ; the
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
199
FIG. 49. Cervical vertebrae of QMJM4729, a 2.12 m long d. A, Anterior view. B, Posterior view. Note bifid
spinous process and poorly developed transverse processes of fused cervicals 1 & 2.
last postzygopophysis on T8-9. The last costal
facet is on T7.
LUMBAR VERTEBRAE (Fig. 51)
The transverse and neural processes reach
their greatest lengths on lumbars 1-3. In MM16,
maximum length of the transverse process is
95mm, that of the neural process 90mm. In
QMJM4752 the last 2 lumbars have reduced
metapophyses, but they have no mammillary
processes or ridges.
CAUDAL VERTEBRAE (Figs 52,53)
The transverse processes are directed anterior-
ly from C22, and by Ca3 the tips of the transverse
processes reach in front of the anterior face of the
centrum. The greatest widths of the transverse
processes are on Cal-5. From about Cal0, the
transverse processes are sharply bent forward
(Fig. 52B). The last transverse process is on
Cal6-17. The last neural process is on Cal9-21;
up to Cal7-18 have metapophyses. In
QMJM4729 there is one transitional, globular
vertebra and the last 9 caudals are dorsoventrally
compressed (Fig. 53A,B); in QMJMS511 there
are two globular, transitional vertebrae.
CHEVRONS (Fig.54)
QM JMS511 has 20 definite chevrons, with pos-
sibly a 21st. The next most complete set of chev-
rons isin MM1015, with 19 chevrons plus 2 small
nodules. QMJM4729 and 4752 have 17 chevrons
but no nodules.
The first chevron has unfused, asymmetrical
arms, with a strongly developed anterior spine
and 2 dorsal attachment points on the larger left
arm (Fig. 54). In MM334 the chevron is enclosed
in thick ligament. The anterior projection and the
two dorsal projections are each linked to the
posteroventral face of a vertebral centrum; i.e the
chevron span three vertebrae. Subsequent chev-
rons have fused, equilateral arms, presenting
more typical Y or V shapes in anterior view.
RIBS
QMJMS511, QMJM4752 and MM333 have 13
ribs (one pair reduced in QMJM4752 and
200 MEMOIRS OF THE QUEENSLAND MUSEUM
it,
Wideeer à
ena
FIG. 50. Thoracic vertebrae of QMJM4729. A, Lateral view. B, Dorsal view. Note
antrorse spine (arrow) on transverse process of last thoracic vertebra.
FIG. 51. Lumbar vertebrae of QMJM4729. A, Lateral view. B, Dorsal view. Note
the maximum development of transverse processes on anterior lumbar vertebrae.
QMJMS11) of which 8 are
bicipital. MM1015 and
MM?2 have 12 ribs, 7 of
which are bicipital.
STERNUM (Fig. 55)
The sternum is variable
in shape with no apparent
sexual dimorphism. A
conspicuous foramen is
consistently present in the
manubrium between the
facets for the first two pairs
of sternebral ribs. The
anterior facets are latero-
dorsal in position. In
MM1015 there are 5 pairs
of articular facets, but 7
pairs of sternebral ribs; the
fifth and sixth sternebral
ribs articulate with the last
facet while the last ster-
nebral rib pair is free.
HYOID APPARATUS
(Fig.56)
The fused basi-thyro-
hyals form a slender ar-
cuate transverse bar with
no obvious sexual dimor-
phism. The stylohyals are
stout, slightly curved and
flattened rods.
PELVIC BONES
The small pelvic bones
are variable but generally
are elongate with a lateral
tubercle anteriorly. Not
enough material from
sexed individuals is avail-
able to determine sexual
dimorphism. The pelvic
bone of MM333 is short
and broad, but larger in-
dividuals of both sexes
have more elongate pelvic
bones.
SCAPULA (Fig. 57)
The scapula is distinctly
longer than deep, with a
distinct anterolateral ridge
marking the supraspinatus
fossa (terminology from
Rommel, 1990), which is
22.3-27.8% of the maxi-
mum scapula length. The
acromion process is con-
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA 201
sistently larger than the
coracoid process, ex-
panded distally to form a
broad triangle, which can
have a slightly emarginate
or concave distal margin.
FLIPPER BONES
The humerus is elongate
(Fig.48). There is a broad,
laterally placed humeral
condyle and a low medial
‘common tubercle’. The
delto-pectoral tuberosity
on its anterior margin can
extend to the distal end but
is usually subdistal to the
epiphyseal suture (Fig.
48).
The olecranon process is
usually prominent (Fig.
48A,B). However, the ulna
can become partially fused
with the humerus, with no
signs of an olecranon
process (Fig. 48C). The 5
carpal bones are usually
distinct; what appear to be
the radiale and inter-
medium can be fused (Fig.
48).
DISCUSSION
COMPARISONS WITH
SOUTHEAST ASIAN
ANIMALS
VERTEBRAL
COLUMN. The number
of vertebrae from Qld
animals is lower than in SE
Asian animals (58-60
(61?) vs 62-63).
Anderson (1879:409)
gave a formula for Orcael-
la of C7, T12-13, L13-14,
Sa2, Ca27-28 = 62-63. For
O. 'fluminalis' the formula
was C7, T13, L16, Ca26
=63 (actually 262, Ander-
son, 1879:366). Given the
variability in published
figures (see below) and the
limited number of animals
FIG. 52. Anterior caudal vertebrae of QMJM4729. A, Lateral view. B, Dorsal
view. Note strong anterior projection of transverse processes.
FIG. 53. Posterior caudal vertebrae of QMJM4729. A, Lateral view. B, Dorsal
view.
“$4513 383323228350...
FIG. 54. Chevrons of MM1015, lateral view.
not, however, related simply to differences in the
examined, it is impossible to assess the sig- sizes of animals examined. Our specimens are
nificance of these differences. Differences are
202
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 55. Dorsal views of sternum from: A, QMJM4752; B, QMJM4729 & C, QMJM4735. Note broadly concave
anterior margin, anterior foramen, dorsolateral orientation of facets foranterior sternebrae, and compound nature
of sternum (A,B).
1.3-2.4 m long, which overlaps the lengths (1.8-
2.3 m) of Anderson's (1879) specimens.
There is difficulty in assessing whether ver-
tebrae are thoracic or lumbar when there are
reduced ribs free from the vertebral column. In
QMJMS11, there are 13 pairs of ribs (last
reduced) but only 12 vertebrae with articular
facets. Functionally, there are 13 thoracic ver-
tebrae, but only the first 12 would be recognized
as thoracic, based on vertebral morphology. We
ignored these riblets in the vertebral formula, as
did Lloze (in deSmet, 1977).
Anderson (1879) noted that sacral vertebrae of
O. brevirostris have transverse processes, with
contracted bases and distal expansions. We do not
see this distinction in our material. Sacral ver-
FIG. 56. Fused basi-thyrohyal bones, QMJM4709.
tebrae are not generally recog- nized in cetaceans
(deSmet,1977;Rommel, 1990).
We follow Rommel (1990) in defining the 1st
caudal as that vertebra with a chevron posterior
to its centrum (cf deSmet, 1977). This is compli-
cated in Orcaella by the elongate, possibly com-
pound, form of the 1st chevron. In MM333 and
MM334 which we dissected, the two dorsal
projections of the left chevron are each associated
with a separate vertebra. In MM334, the anterior
projection is also closely associated with a third
vertebra. In MM333, the underside of the first
centrum is intermediate, lacking the sharp keel
typical of lumbar vertebrae and the broad trian-
gular facet for the attachment of chevrons seen on
most caudals. In MM334, the vertebra linked
with the anterior projection of the first chevron
has a sharp keel and would have been classified
as lumbar on this basis; the next two vertebrae
have broad tubercular facets typical of caudal
vertebrae. Only the vertebrae associated with
dorsal projections of the chevron are considered
as caudal vertebrae.
Anderson (1879:412) noted that in O. brevi-
rostris the chevrons begin between the 34th and
35th vertebrae. Using Rommel's (1990) defini-
tion, the first caudal vertebra would be the 34th.
For O. ‘fluminalis’, Anderson (1879: 408) noted
that chevrons started between the 34th and 35th
vertebrae (although elsewhere he gave the posi-
tion as between L12-13).
PHYLOGENY OF THE IRRAWADDY DOLPHIN, ORCAELLA
203
FIG. 57. Lateral view of right scapula. A, QMJM4720. B, QMJM4726. Note well developed acromion process,
with triangular distal expansion.
The revised formulae for O. brevirostris and O.
‘fluminalis’ would thus be C7, T13, L13, Ca28-
29=62-63 and C7,T13, L13, Ca29=62. The for-
mula for an Orcaella from the Mekong River was
C7, T12, L15, Ca28=62 (de Smet, 1977, adjusted
for the difference in defining caudal vertebrae).
Lloze (MS, and in Marsh et al., 1989) gave a
formula of C7, T13, L16, Ca26-27=62-63.
CHEVRONS
In our material only the first chevron is un-
fused; Anderson (1879) noted that the first two
chevrons are unfused, as are those from chevron
20 posteriorly. Anderson did not describe the
shape of the first chevron but noted it articulated
*to the anterior and posterior surfaces of the 34th
vertebra’. This implies that there were 2 dorsal
projections, as in our material.
RIBS
Anderson (1879), in describing O. ‘fluminalis’,
noted that there were 13 ‘true’ ribs with ‘a free
floating’ rib “considerably removed from the ver-
tebral column'. In his diagnosis of the genus he
noted 12-13 vertebral ribs , with 1-2 free ribs.
Both statements imply a total of up to 14 ribs.
204
Lloze (MS; in deSmet, 1977) also noted 13 pairs
of ribs, with a 14th pair free from the vertebral
column. MM333 has 13 pairs of ribs, but other
Australian specimens have 12 vertebral ribs,
sometimes with a 13th free pair of reduced riblets.
It is unlikely that the free riblets have been over-
looked in those animals where only 12 pairs of
ribs were collected.
STERNUM
This is variable (Fig. 55). Our specimens more
closely resemble the description and illustration
for O. ‘fluminalis’ in Anderson (1879, pl. 43, fig.
10), than O. brevirostris (pl. 43, fig. 5). Although
Anderson did not illustrate a foramen in the
manubrium, he noted it in O. brevirostris. None
of the sterna illustrated by Anderson or Pilleri &
Gihr (1973-1974) are compound, whereas most
of Qld specimens have a smaller posterior piece
as well as the manubrium (Fig. 55A,B). MM334
has a median slit running from the posterior end
to the level of facets for the second pair of ster-
nebral ribs. This is an even more extensive
development of the *deep wide notch' than noted
in O. ‘fluminalis’ by Anderson (1879).
PELVIC BONES
The elongate pelvic bones of Qld animals cor-
respond well to the illustration in Anderson
(1879, pl. 42, fig. 11). The pelvic bones illustrated
by Pilleri & Gihr (1973-1974, pl. 27, fig. 4)
appear to be stylohyals.
SCAPULA
The scapula from Qld specimens correspond
more to Anderson's (1879, pl. 43, fig. 11) illustra-
tion of O. ‘fluminalis’, but given the variability
in our small series there is no basis to separate O.
‘fluminalis’ and O. brevirostris. The major dis-
MEMOIRS OF THE QUEENSLAND MUSEUM
tinction in our material from previous descrip-
tions is that the acromion process is generally
larger than the coracoid process (Fig. 57) which
is opposite to the situation in SE Asian animals
(Anderson, 1879; Pilleri & Gihr, 1973-1974, pl.
26).
APPENDIX 5: NOMENCLATURE
The author citation for O. brevirostris is
generally given as (Gray, 1866). It is generally
recognized, however, that the first description of
the species was by Sir Richard Owen. This incon-
sistency is apparent in the synonymy given by
Hershkovitz (1966), where in the space of 5 lines
the authority was given as 'Orcaella brevirostris
Gray' but the type species was given as 'Orcaella
brevirostris Owen, by monotypy'.
Owen gavean extensive description of the skull
of Phocaena (Orca) brevirostris, which was read
June 20, 1865. The account was not published,
however, until the following year. The volume is
dated 1869, but independent evidence (Zoologi-
cal Record 1867) indicates that Owen's article
was available in 1866; Johnson (1964) gave the
publication date as August 15, 1866. In the mean-
time (March 1866, according to Johnson (1964))
Gray included an account of ‘Orca brevirostris’
in his Catalog of the seals and whales in the
British Museum. He attributed Phocaena (Orca)
brevirostris to the unpublished account by Owen
(‘Zool. Trans v, ined’) and included (also at-
tributed to Owen) an extensive extract of the
description and a figure by that worker.
The description and species name were at-
tributed to Owen by Gray. Thus following Ride
et al. (1985:91, article 50) and ICZN Recommen-
dation 51B, we consider the appropriate author
citation to be O. brevirostris (Owen in Gray,
1866).
A NEW SPECIES OF POMPONATIUS DISTANT FROM AUSTRALIA
(HEMIPTERA:HETEROPTERA:COREIDAE:ACANTHOCORINI)
H. BRAILOVSKY AND G.B. MONTEITH
Brailovsky, H. & Monteith, G.B. 1996 07 20: A new species of Pomponatius Distant from
Australia (Hemiptera:Heteroptera:Coreidae: Acanthocorini). Memoirs of the Queensland
Museum 39(2):205-210. Brisbane. ISSN 0079-8835.
The Australian Pomponatius Distant is redescribed and the distribution of its type species
P.typicus Distant is extended to the Northern Territory and to New Guinea. P. Iuridus sp.
nov. is described from eastern and northern Australia. Distribution maps and a key are
included. Food plants are Melaleuca and Callistemon (Myrtaceae). _] Hemiptera, Coreidae,
Pomponatius, taxonomy, biological control.
H. Brailovsky, Instituto de Biologia UNAM, Departamento de Zoologia, Apdo. Postal #
70153, México, 04510 D.F.; G.B. Monteith, Queensland Museum, P.O. Box 3300, South
Brisbane, Queensland 4101, Australia; received 5 February 1996.
Pomponatius Distant, 1904 of the Acan-
thocorini was described from coastal Queensland
and is known only from the type species, P.
typicus Distant, 1904. It has been mentioned as a
garden pest, under the common name of the cal-
listemon tip bug, damaging cultivated species of
the large Australian native plant genera
Melaleuca and Callistemon (Jones & Elliot,
1986).
Because Melaleuca quinquenervia (Cav.) S.T.
Blake, has become a severe weed pest of wetlands
in Florida (Balciunas, 1990) the United States
Department of Agriculture undertook an inten-
sive search among native Australian insects for
possible biocontrol agents. This search identified
Pomponatius typicus, under the common name of
the tip wilting bug, as one possible biocontrol
agent (Balciunas & Burrows, 1994).
Museum collections were found to contain a
second undescribed species of Pomponatius
which widely overlaps in distribution and food
preference with the type species. The new species
is described below and the generic range is ex-
tended to the Northern Territory and to New
Guinea.
The following abbreviations indicate institu-
tions where specimens are deposited or which
generously lent material: Australian Biological
Control Laboratory, Townsville (ABCL);
Australian National Insect Collection, Canberra
(ANIC); Museum and Art Gallery of Northern
Territory, Darwin (NTM); Department of
Primary Industries, Mareeba (MDPI); Depart-
ment of Primary Industries, Brisbane (QDPI);
The Natural History Museum, London (BMNH);
California Academy of Sciences, San Francisco
(CAS); Canadian National Collections, Ottawa
(CNC); Colección Entomológica del Instituto de
Biología, Universidad Nacional Autónoma de
México (IBUNAM); Australian Museum, Syd-
ney (AMS); Queensland Museum, Brisbane
(QM); University of Queensland Insect Collec-
tion, Brisbane (UQIC); University of California,
Riverside (UCR).
All measurements are in millimetres.
Pomponatius Distant, 1904
Pomponatius Distant, 1904:265.
DESCRIPTION.Body elongate, narrowed
posteriorly.
Head. Wider than long across eyes, nearly pen-
tagonal, not produced beyond the antenniferous
tubercles and dorsally flat; tylus apically up-
turned to form a small horn, with apex subacute
or truncate; jugum unarmed, thick, apically
globose, equal or slightly shorter than tylus; inner
margins of antenniferous tubercles with con-
spicuous and large lobe apically rounded and
widely separated; antennal segment I robust,
thickest and always longer than head, segments
II and III cylindrical, regularly incrassate and
segment IV always fusiform or antennal seg-
ments I and II regularly incrassate and segment
III the thickest and gradually widening; segment
IV the shortest, segment III shorter than I and II
and segment II usually shorter than I; ocelli ses-
sile; preocellar pit deep; eyes moderately large,
semiglobose and sessile or moderately large and
compressed at lateral margins; postocular
tubercle relatively small; bucculae rounded,
short, not extending beyond anterior margin of
eyes; rostrum reaching anterior margin or middle
206
FIG. 1.Dorsal view of Pomponatius luridus sp. nov.,
male, length 11.5mm.
third of mesosternum; rostral segment I the
shortest, segment III shorter than I and IV and
segment IV the longest or subequal to I; frons
with a deep central longitudinal incision.
Thorax. Pronotum trapeziform, wider than
long, moderately declivent; collar not clearly
marked; frontal angles produced forward as long
and broad or produced as medium sized and
slender conical projections; anterior margin con-
cavely sinuate; anterolateral margins clearly
emarginate or slightly upwardly reflected and
nodulose; humeral angles rounded, not expanded
and dorsally tuberculate; posterolateral margins
slightly concave and smooth; posterior margin
expanded on a short lip with the posterior border
straight and smooth; calli not elevated, separated
along midline by short longitudinal furrow.
Anterior lobe of metathoracic peritreme elevated
and reniform, posterior lobe sharp, small;
mesosternum with a medial groove, extending to
anterior third.
Legs. Femora apically incrassate, densely
tuberculate and ventrally armed with a distinct
tooth near apex; tibiae terete, conspicuously sul-
cate.
MEMOIRS OF THE QUEENSLAND MUSEUM
Scutellum. Triangular, longer than wide, flat,
transversely striate; apex subacute or truncate.
Hemelytra. Macropterous, almost reaching the
apex of the last abdominal segment; costal mar-
gin emarginate; apical margin almost straight or
clearly sinuate, with inner third concave and outer
third straight; apical angle obtuse; hemelytral
membrane elongate with several longitudinal
veins or small with reticulate venation.
Abdomen. Connexival segments complete, not
extended as short spines; abdominal sternite
without medial furrow.
Male genitalia. Genital capsule: Posteroventral
margin projected in a broad or slender large-sized
triangular lobe (Figs. 2F, 2G). Parameres: Body
simple and broad or with middle third narrow,
neck-like; apical projection with anterior lobe
convex and continuous with body and posterior
lobe conspicuously curved or dilated and ending
in a sharp or blunt short projection (Figs. 2A-E).
Female genitalia. Abdominal sternite VII with
plica and fissura; plica triangular, short, reaching
anterior third of sternite VII. Genital plate:
Gonocoxae I square, shorter than paratergite IX,
in caudal view closed, in lateral view slightly
convex; paratergite VIII triangular, with spiracle
visible; paratergite IX elongate, triangular, larger
than paratergite VIII.
DIAGNOSIS. Body elongate and narrowed
posteriorly, antennal segments I to III regularly
incrassate, antenniferous tubercles with a distinct
lobe on the inner margins, posterior margin of
pronotum expanded on a short lip and femora
apically incrassate with a distinct tooth near apex.
REMARKS. Kumar (1965) studied an undeter-
mined species of Pomponatius and described the
male and female genitalia as well as the glands
associated with the internal male organs.
Pomponatius luridus sp. nov.
(Figs 1,2C-G, 4)
MATERIAL EXAMINED. HOLOTYPE d:
QMT13987: Central Qld: Mt. Moffatt NP, East
Branch, Maranoa R, (Top Moffatt Camp), 26.1x.1986,
Monteith, Yeates & Thompson, in QM. PARATYPES:
Qld: QMT13988, 9, same data as holotype (QM);
QMT13989, 9, Levers Plateau, via Rathdowney, 6-ii-
1966, F.R. Wylie (QM); QMT13990,d, Mt Tam-
borine, 27.x.1957, S. Breeden (QM); 9, St George,
13.v.1973, J Hodgson (IBUNAM); 183 9? , In-
dooroopilly, xi. 1979, G.Gordh (UCR + IBUNAM); 6,
Einasleigh Riv., 35 m. E. of Georgetown, 31.v.1972,
G.B. & S.R. Monteith (UQIC); 9, 62 km W. Mt
Garnet, 31.xii.1978, R.I. Storey, on Melaleuca sp.
NEW SPECIES OF POMPONATIUS FROM AUSTRALIA
f) ji
F G
FIG. 2.Male genitalia. A-B, parameres of Pomponatius
typicus Distant. C-E, parameres of Pomponatius
luridus sp. nov. F-G, dorsal and lateral views of
genital capsule P. luridus sp. nov. (Scale lines in mm.)
(MDPD; 1¢ 1 9, Biggenden, 12.xii.1971, H. Frauca
(ANIC); 1d. 599 , Degilbo Ck., Biggenden,
4.xii.1971, H. Frauca (ANIC); 1 d, Mullet Ck., 31 mi.
N. Bundaberg, 27.iv.1959, T.G. Campbell (ANIC); d,
Calliope R. Xing, N. of Gladstone, 11.1965, E.J. Reye,
on bottle brush (ANIC); c, Brisbane, 24.viii. 1976,
B.K. Cantrell (QDPI); 9, Kingaroy, 7. xi.1983, J.
Wessels, on Melaleuca armillaris (QDPI); 28 , Stan-
thorpe, 6.xii.1926 (QDPI); 9, Indooroopilly,
15.1.1962, J.H. Barrett (QDPI); 9, James Cook Univ.,
Townsville, 4.ix.1989, S.J. Newman, on Callistemon
viminalis (ABCL); 8 , Chelmer, Brisbane, 15.ix.1992,
M. Purcell, on Callistemon viminalis (ABCL). NSW:
9 , Oatley, 15.vi.1950, Deuquet (AMS); , Enfield, Syd-
ney, E.P. Ramsay (IBUNAM); 29 9, Rivertree,
10.viii.1921 (QDPI). NT: d, Edith Falls, Katherine
Gorge NP, 6.xii.1980, M.B. Malipatil & I. Archibald
207
(NTM); 1 d, No locality, 25.ix.1896, A.L. Schrader
(AMS).
DESCRIPTION. MALE. Dorsal coloration. Pale
yellow, with punctures dark orange and following
areas creamy yellow: anterolateral margins of
pronotum and apex of scutellum; antennal seg-
ment I orange yellow with tubercles reddish, seg-
ment II orange yellow with apical third and
tubercles reddish, segment III with apical half and
apical join pale yellow and basal half including
the tubercles dark orange reddish and segment IV
pale yellow; following areas black: longitudinal
stripe close to the anterior third of anterolateral
margins of pronotum, and anterior third of costal
margin of corium; connexival segments and ab-
dominal sternite VII pale yellow with punctures
dark orange; abdominal segments I to VI bright
orange. Ventral coloration: Pale yellow with pink
diffuse areas and with following areas black:
apex of rostral segment IV, one small discoidal
spot on propleura and mesopleura, two lon-
gitudinal series of discoidal spots running lateral-
ly to the midline on mesosternum and irregular
spots on mesocoxa, metacoxa, mesotrochanters
and metatrochanters; tubercles of femora black or
reddish brown or pale pink.
Structure. Antennal segments I to III regularly
incrassate; eyes moderately large, semiglobose,
sessile but not compressed at lateral margins.
Pronotum: Frontal angles produced forward as
long and broad conical projection; anterolateral
margins clearly emarginated. Hemelytra: Apical
margin clearly sinuate, with inner third concave
and outer third straight; hemelytral membrane
elongate with several longitudinal veins.
Genital capsule. Posteroventral margin
projecting as a large broad triangular lobe (Fig.
2F,G). Parameres (Fig. 2C-E).
FEMALE. Coloration. Similar to male. Ab-
dominal segments IV to VII bright orange with
two pale brown discoidal or irregular spots, close
to the connexival segments; segment VIII dark
orange and segment IX dark reddish brown with
longitudinal stripe orange, close to midline; con-
nexival segments I to VII pale reddish brown with
upper margin mostly pale yellow; connexival
segments VIII and IX mostly yellow; abdominal
sterna and genital plates pale yellow with follow-
ing areas black: irregular spots on sterna III to VII
and close to midline as well as the internal margin
of fissure and the area close to upper margin of
plica.
VARIATION. 1, Antennal segment IV with dis-
208
FIG, 3.Dorsal view of Pomponatius typicus Distant,
female, length 15.1mm.
tal half pale yellow and basal half pale orange
yellow. 2, Humeral angles black or dark brown.
3, Endocorium with black discoidal spot some-
what diffuse. 4, Scutellar disc with or without
median black stripe. 5, Abdominal sterna and
genital plates of female pale yellow, with pale
pink marks or with only the internal margins of
fissure and the areas close to the plica black. 6,
Abdominal segment VIII of female bright orange.
MEASUREMENTS. ófirst, then 2: Head
length: 1.08, 1.32; width across eyes: 1.36, 1.56;
interocular space: 0.80, 0.91; interocellar space:
0.25, 0.27; length antennal segments: I, 2.40,
2.72; II, 2.48, 2.64; III, 2.08, 2.24; IV, 2.00, 1.88.
Pronotum: Total length: 2.12, 2.76; width across
frontal angles: 1.20, 1.44; width across humeral
angles: 2.40, 3.08. Hemelytra: Total length of
hemelytral membrane: 3.76, 4.40. Scutellar
length: 1.00, 1.40; width: 0.92, 1.32. Total body
length: 11.52, 14.65.
MEMOIRS OF THE QUEENSLAND MUSEUM
DISTRIBUTION (Fig. 4). From Sydney along
coastal and inland Queensland to the Northern
Territory.
FOOD PLANTS. See discussion below.
ETYMOLOGY. Latin luridus, pale yellow; for its light
coloration.
Pomponatius typicus Distant, 1904
(Figs 2A-B, 3, 4)
Pomponatius typicus Distant, 1904: 266.
MATERIAL EXAMINED. NSW: Bulahdelah,
11.xi.1932 (IBUNAM); Hat Head, nr. Kempsey,
3.1.1970 (ANIC). QLD: HOLOTYPE 2, Townsville
(BMNH); Acacia Ridge, 16.1.1973; Lockerbie Area,
Cape York, 13- 27.iv.1973; Jardine River road cross-
ing, 16-27.ix.1974; Brisbane, 24.x.1916; Lakefield
NP, 75 km N. of Laura, 15-28.vi.1980; Kuranda,
3.x.1920; Nth Keppel Island, track to Mazie Bay,
3.ix.1987 (QM); Archerfield, 19.ix.1964; St Lucia,
6.ix.1965; Bowen, 2.vi.1965; Brisbane, 20.v.1959
(UQIC); Black Mt Rd (Kuranda), 24.1.1970; Cairns,
22.vii-24.xii.1969 (CNC); Townsville 5.ii.1945
(CAS); Watalgan Range, via Rosedale, 6.iv.1974;
Baldwin Swamp Fauna Reserve, Bundaberg,
27.xi.1971; Goodwood, Isis Shire, 5.iv. 1974; Gum-
dale, Brisbane, 11.xi.1968; Ladysmith Yard at Turnoff
Lagoon, 18.ix.1930 (ANIC); Bridge Ck, Cook High-
way, 29.v.1974, on Melaleuca; Kurrimine Beach Rd,
26.vi.1974; Mareeba, 1.11.1979, at light; Home Hill,
28.vii.1947 (MDPI); Chandler, Brisbane, 29.i1.1976,
damaging new growth of Callistemon sp.; Mareeba,
29.ix.1981; Townsville, viii. 1984, on Callistemon
viminalis, nr. Gamboola H.S., 23.vii.1982, (QDPI);
Coombabah, 14.11.1990 & 4.ix.1990, on tips of
Melaleuca quinquenervia; Burpengary, 12.xi.1992,
feeds on M. quinquenervia, Chelmer, 15.ix.1992, on
M. quinquenervia; Centenary Park, Cairns, 8.v.1991,
tips of M. quinquenervia; 16.9 km ESE of Tully,
31.viii.1994 and 2.viii.1994, ex M. quinquenervia;
same locality, 27.vi.1994, ex M. dealbata; 14.3 km S.
Tully, 31.viii.1992; 21.8 km S. Tully, 4.xi.1993, ex M.
dealbata; 15 km ESE Tully, 27.xi.1994, ex M. quin-
quenervia; James Cook Univ., Townsville, 17.1v.1990
& 18.1.1993, feeding on Callistemon viminalis, same
locality, 4.vii.1991 and 2.11.1992, ex. Melaleuca quin-
quenervia (ABCL). NT: 48 ml. SW of Daly River,
30.viii.1968; 9 km NE of Mudginbarry HS, 26.v.1973
(ANIC); Wildman Rivercashew plantation, 17.x.1989,
ex adjacent Melaleuca regrowth; Mary R., 37 ml. E of
Pine Creek, 9.vii.1971; Darwin, 9.viii.1983, ex Lep-
tospermum (NTM); Crystal Falls - Biddlecombe Cas-
cades, Nitmiluk NP, 17-18.ii1.1995, ex Melaleuca
viridiflora (QM); Howard River, Darwin, 24.x.1986,
ex Melaleuca nervosa (ABCL). PNG: Rouku,
Morehead River, Western District, PNG, 19.iii-
28.v.1962 (ANIC).
NEW SPECIES OF POMPONATIUS FROM AUSTRALIA
NEW GUINEA
Port Moresby
Brisbane
QUEENSLAND
ansaa F E
I
|
NORTHERN TERRITORY |
|
i}
l
l
\
1
1
\
I
1
\
1
i
\
\
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a NEUE rend
|
\
|
Newcastle
@ Pomponatius typicus |
| New SouTH WALES
A Pomponatius luridus
LLL—————
FIG. 4.Distribution map of Pomponatius species.
DIAGNOSIS. Antennal segment III gradually
widening and uniformly pale orange yellow,
hemelytral membrane short, with reticulate vena-
tion, anterolateral margins of pronotum slightly
reflected, and propleura and mesopleura without
black discoidal spot. Genital capsule:
Posteroventral margin projected in a large slender
triangular lobe. Parameres (Fig. 2A-B).
DISTRIBUTION (Fig. 4). This species was first
described from Townsville (Distant, 1904) and
later recorded from Rockhampton (Blóte, 1935)
and Heathlands in Cape York Peninsula (Cassis,
1993). It is now known to be widespread in
Australia from northern NSW along the coast to
north Queensland. It is also now recorded from
the Northern Territory and New Guinea.
Genitalia of specimens from the Northern Ter-
ritory and southern New Guinea agree with those
from eastern Australia.
FOOD PLANT RECORDS
Pomponatius feeds by sucking sap from young
stems at the ends of branches, especially when the
apical shoot has a flush of soft new growth. Their
feeding causes wilting and sometimes death of
209
the terminal shoot and its young leaves. This
effect is often useful evidence for locating
specimens on plants and has lead to their common
name of ‘tip wilting bugs’.
All records for Pomponatius species are from
plants of the Family Myrtaceae, as follows:
Pomponatius luridus — Melaleuca sp. (Mt Gar-
net); Melaleuca armillaris (Kingaroy); Callis-
temon viminalis (Townsville, Chelmer); on bottle
brush (Calliope R.).
Pomponatius typicus — Melaleuca quinquener-
via (Coombabah, Burpengary, Chelmer, Cairns,
Tully District, Townsville); Melaleuca dealbata
(Tully district); Melaleuca viridiflora (Nitmiluk
NP); Melaleuca nervosa (Darwin); Melaleuca sp.
(Cook Highway, Wildman R); Callistemon
viminalis (Townsville) Callistemon sp. (Chel-
mer); Leptospermum sp. (Darwin).
Almost all records are from the closely related
Melaleuca and Callistemon. The only confirmed
Callistemon is C.viminalis which is sometimes
assigned to Melaleuca. The only other genera for
which records exist are Leptospermum (one
record from Darwin which may have been a
garden plant and for which actual feeding has not
been confirmed) and a prolonged infestation of a
garden plant of Calothamnus in Brisbane by a
species of Pomponatius which was not identified
(GBM obs.). Calothamnus is related to
Melaleuca but is naturally restricted to south-
western Australia beyond the known range of
Pomponatius.
Circumstantial evidence for Pomponatius
feeding on other plant species in the form of
characteristically damaged growing tips has been
noted by the USDA survey staff on Melaleuca
arcana, M. argentea, M. bracteata and Callis-
temon polandii (Burrows pers. comm.).
Although both species of Pomponatius have
been recorded from both Melaleuca and Callis-
temon it would appear from inferences which can
be drawn from the data that, in the wild, Pom-
ponatius luridus is virtually restricted to Callis-
temon viminalis, and Pomponatius typicus is
similarly restricted to ‘paper bark tea trees’ of the
M. leucadendra complex as defined by Blake,
1968. For P. luridus the record from M. armillaris
is from a garden plant outside its natural range
while many of the collection localities where
plant associations were not recorded are habitats
where C. viminalis is abundant, e.g. Calliope R.,
Maranoa R., Degilbo Ck, Mullet Ck. Similarly
with P. typicus all Callistemon records are from
unnatural suburban situations, while most natural
occurrences of the bug are from habitats where
FIG. 5. A living 2 of Pomponatius luridus sp.nov. on
its foodplant Callistemon viminalis in Brisbane.
paperbarks predominate. This includes the PNG
record.
This apparent food plant dichotomy between
the two Pomponatius species is suggested as the
mechanism which maintains their specific in-
tegrity throughout almost identical distributions.
KEY TO THE KNOWN POMPONATIUS
SPECIES
1.Antennal segment III not noticeably more swollen
than II, and bicoloured with apical half paler
than basal half; hemelytral membrane elongate,
longer than 3.20mm, with several longitudinal
veins, and with its basal margin strongly sinuate;
dark longitudinal stripes on prosternum about
same width as rostrum; apical lobe of paramere
not dilated and without a small projection (Figs
2C-E) . . Pomponatius luridus sp. nov. (Fig. 1)
Antennal segment III noticeably more swollen
than II and not bicoloured; hemelytral
membrane short, less than 3.20mm, with reticu-
MEMOIRS OF THE QUEENSLAND MUSEUM
late venation and with its basal margin uniform-
ly curved; dark longitudinal stripes on proster-
num at least twice width of rostrum; apical lobe
of each paramere dilated and ending in a sharp
or blunt projection (Fig. 2A,B)
Pomponatius typicus Distant (Fig. 3)
ACKNOWLEDGEMENTS
Thanks to the following individuals and institu-
tions for loans and other assistance: J. Balciunas
(ABCL), T.A. Weir (ANIC), G. Brown (NTM),
R. Storey (MDPI), J. Donaldson (QDPI), G. Cas-
sis (AMS), Janet Margerison-Knight (BMNH),
Norman D. Penny (CAS), Michael D. Schwartz
(CNC), Margaret Schneider (UQIC) and Saul
Frommer (UCR). We are especially grateful to J.
Balciunas for access to the melaleuca insect sur-
vey data accumulated by his group. Biol. Ernesto
Barrera, Cristina Urbina, and Felipe Villegas
(IBUNAM) prepared the dorsal view illustrations
and genitalia drawings. The Consejo Nacional de
Ciencia y Tecnologia, Mexico (CONACT)
provided financial assistance to HB.
LITERATURE CITED
BALCIUNAS, J. 1990. Australian insects to control
melaleuca. Aquatics 12:15-19.
BALCIUNAS, J. & BURROWS, D. 1994. USDA
Australian Biological Control Laboratory 1994
Annual Report. 32pp.
BLAKE, S.T. 1968. A revision of Melaleuca
leucadendron and its allies (Myrtaceae). Con-
__ tributions of the Queensland Herbarium 1:1-114.
BLOTE, H.C. 1935. Catalogue of the Coreidae in the
Rijksmuseum van Natuurlijke Histoire. Part 1.
Coreinae, First Part. Zoologische Mededelingen
18: 181- 227.
CASSIS, G. 1993. A report on the Heteroptera and
Dermaptera from the survey of Cape York Penin-
sula. pp. 109-123. In Cape York Peninsula Scien-
tific Expedition, Wet Season 1992 Report, Vol. 2.
Royal Geographical Society of Queensland.
DISTANT, W.L. 1904. Rhynchotal Notes. 22. Heterop-
tera from north Queensland. Annals and Magazine
of Natural History, ser. 7 13: 263-276.
JONES, D. & ELLIOT, R. 1986. ‘Pests, diseases and
ailments of Australian plants’. (Lothian Press:
Melbourne) 333p.
KUMAR, R. 1965. Aspects of the Morphology of
Coreoidea and their value in its higher classifica-
tion. Proceedings of the Royal Society of
Queensland 76(3): 27-91.
REVISION OF THE AUSTRALIAN GENUS MICROTROPESA MACQUART
(DIPTERA: TACHINIDAE: TACHININI)
CHRISTOPHER J. BURWELL
Burwell, C.J. 1996 07 20: Revision of the Australian genus Microtropesa Macquart (Diptera:
Tachinidae:Tachinini) Memoirs of the Queensland Museum 39(2): 211-226. ISSN 0079-
8835.
The endemic Australian genus Microtropesa Macquart is revised and its 14 species keyed.
M. danielsi sp. nov. and M. longimentum sp. nov. are described. Microtropesa nigricornis
Macquart and M. intermedia Malloch are confirmed as valid. Microtropesa flavitarsis
Malloch is considered a junior synonym of M. obtusa (Walker). The female of M. viridescens
Paramonov and the male of M. canberrae Paramonov are described for the first time. The
first instar larva of M. intermedia and the genitalia of males of most species are figured,
O Diptera, Tachinidae, Microtropesa, Australia.
Christopher J. Burwell, Department of Entomology, University of Queensland, Brisbane,
Queensland 4072. Current address: Queensland Museum, PO Box 3300, South Brisbane,
Queensland 4101; received 7 December 1995.
Australian Tachinini are dominated by the en-
demic Microtropesa Macquart (Crosskey, 1973),
specimens of which are up to 16 mm long and
often with striking colour patterns on the thorax
and abdomen. Malloch (1928, 1929, 1930)
described five species and provided a key. Hardy
(1939) dealt with a number of species described
by earlier authors but omitted by Malloch and
synonymised several names (many incorrectly).
Hardy incorrectly synonymised Tasmaniomyia
Townsend with Microtropesa (Paramonov,
1951; Crosskey, 1973). Paramonov (1951)
described four new species and provided a useful
but incomplete key (three species discussed in the
text were omitted from the key). Paramonov was
unsure of the status of M. intermedia but thought
it probably conspecific with M. nigricornis.
These species have been regarded as distinct
(Crosskey, 1973; Cantrell & Crosskey, 1989).
Paramonov (1951) was also unsure of the identity
oftwo species described by Walker (1853, 1858).
METHODS
Male genitalia were examined by detaching the
terminal abdominal segments, immersing them in
cold 1096 KOH for 24—48 hours, washing in
water and dissecting in 70% ethanol. The
preparations were stored in glycerol in separate
microvials. These were pinned next to the
specimen and given duplicate labels.
Drawings of male genitalia and larvae were
made with the aid of acompound microscope and
drawing tube. Other drawings were prepared
using a dissecting microscope and camera lucida.
Measurements of body length were taken from
pinned specimens using a micrometer eyepiece.
TERMINOLOGY
Terminology follows Crosskey (1973) and the
following abbreviations are used: Al, A2, A3 —
Ist, 2nd and 3rd antennal segments; acr, acros-
tichal seta; dc, dorsocentral seta; fr, frontal seta;
ia, intra-alar seta; mm, median marginal seta; ov,
outer vertical seta; pfr, parafrontal; pf, parafacial;
pro-orb, proclinate orbital seta; prst-acr,
presutural acrostichal seta; prst-dc, presutural
dorsocentral seta; prst-ia, presutural intra-alar
seta; stp, sternopleural seta; St1, St2 - 1st and 2nd
abdominal sternites; T1+2, Ist apparent ab-
dominal tergite (= fused 1st and 2nd tergites); T3,
T4, T5 — 2nd, 3rd and 4th apparent abdominal
tergites. Head height is defined as the vertical
dimension of the head, in lateral view, from the
ocellar triangle to the peristome (Fig. 1C).
Abbreviations for depositories are as follows:
Australian Museum, Sydney (AM); Australian
National Insect Collection, Canberra (ANIC);
Biological and Chemical Research Institute, Syd-
ney (BCRD); British Museum (Natural History),
London (BMNH); Institut für Pflanzenschutz-
forschung (formerly Deutsches Entomologisches
Institut), Eberswalde, Germany (DED; G.
Daniels collection, Brisbane (GD); Muséum Na-
tional d'Histoire Naturelle, Paris (MNHN);
Queensland Department of Primary Industries,
Brisbane (QDPI); Queensland Museum, Bris-
bane (QM); South Australian Museum, Adelaide
(SAM); University of Queensland Insect Collec-
212
tion, Brisbane (UQIC); Western Australian
Department of Agriculture, Perth (WADA).
SYSTEMATICS
Order DIPTERA
Family TACHINIDAE
Tribe TACHININI
Microtropesa Macquart, 1846
Microtropesa Macquart, 1846:313; Engel, 1925:344;
Malloch, 1928:614; Malloch, 1929:286; Malloch,
1930:99; Hardy, 1939:33; Paramonov 1951:761;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:761.
Gerotachina Townsend, 1916:152. Type species
Tachina obtusa Walker, 1853, by original designa-
tion. Synonymy by Hardy, 1939:33.
Tasmaniomyia Townsend, 1916; Hardy, 1939:33
(misidentification).
TYPE SPECIES. Musca sinuata Donovan, 1805, by
monotypy.
DIAGNOSIS. Eyes bare, pf haired; A2 elongate,
almost as long as A3 or longer (Figs 1C,D, 3C,
3G, 4D, 5A); 2 with 2 pro-orb, absent in d; &
without ov; palpi normal. Thorax with at least
4+4 dc, 1+2 ia; propleuron haired; 2 or 3 (rarely
4) stp; pleurotergite with long, dense hairs; cell
R5 open (Fig. 5B), rarely closed (Fig. 3H). Ab-
domen wider than thorax, normally subglobose
with whitish or grey pollinose markings; T3 with
or without mm, if present up to 5 pairs; sternites
exposed, often with spiniform setae; aedeagus
without epiphallus.
Females differ from males in having wider fore
and mid tarsi, shorter claws and a conspicuously
wider vertex.
REMARKS. The long, dense hairs on the
pleurotergite distinguish Microtropesa from all
other Tachinini except the New Guinean
Paratropeza Paramonov, which is distinguished
by an inflated abdomen with fused T3 and T4 and
deeply excised T5 and by very strong spiniform
setae on the abdomen and scutellum.
BIOLOGY. Microtropesa flaviventris Malloch
parasitises the noctuid moths Mythimna
(Pseudaletia) convecta (Walker) and Persectania
ewingii (Westwood) (Crosskey, 1973). Hosts of
other species of Microtropesa are unknown.
About 70% of specimens of Microtropesa in
collections are males. This is due in part to the
habit of males of some species of frequenting
prominent mountain peaks and hilltops. Males of
MEMOIRS OF THE QUEENSLAND MUSEUM
M. sinuata and M. violacescens have been ob-
served resting on the ground in exposed, sunny
situations at the summit of hilltops and sometimes
on prominent rocky outcrops.
The genus occurs mainly in the southern half of
the continent (Fig. 7) with only M. danielsi sp.
nov. and M. violacescens known north of 23°S.
KEY TO SPECIES OF MICROTROPESA
1.Mentum of proboscis longer than head height (Fig.
3G); wing with cell R5 closed (Fig. 3H)
Pid ork SE ght 2 As, d longimentum
Mentum of proboscis shorter than head height
(Figs 3C, 5A); wing with cell RS open (Fig. 5B)
2
2. Base of wings with dark brown or black mark-
ings; legs black
Base of wings yellowish or orange, without dark
brown or black markings; legs with at least
femora yellowish or orange ......... 4
3.Hairs on palpi mostly yellow ..... viridescens
Hairs on palpi black ............ skusei
4.Dorsal surface of scutellum with yellow or orange-
yellow DASE. ir 250.0. ap aa mea 5
Dorsal surface of scutellum with black or dark
Brown Hains octe x, «eho phy elu a ais 7
5.Pf with black hairs extending below level of vibris-
sae, at most with a few orange-yellow hairs
ventrally near eyes; head, in lateral view, with
epistome projecting weakly (Fig. 1C); fr, de and
acr very weak and long, hair-like canberrae
Pf with black hairs (if present) not extending
below level of vibrissae, usually at least ventral
third (often much more) of pf with orange-yel-
low hairs; head, in lateral view, with epistome
projecting more strongly (Figs 1D, 5A); fr, dc
and acr strong, distinctly bristle-like
6.T3 of d , from above, with pair of submedian,
grey, pollinose, anterior spots, rarely absent in
9; d with grey, pollinose triangle on T4 about
as wide as distance between basal scutellar
setae, narrower in 9 ; prescutum usually with dis-
tinct pattern of whitish pollinose stripes and
SPOUS~ ese T P pope herd EL sinuata
T3 of d, from above, without grey, pollinose
spots; d with grey, pollinose triangle on T4
about as wide as distance between subapical
scutellar setae; prescutum with thin, shifting
whitish pollinosity a little thicker surrounding
anterior prst-dc; ? unknown ...... danielsi
7.Dorsal surface of T5 with pair of dark-brown or
black, submedian, spots separated from anterior
margin and sometimes from similar ventral pair
of dark spots by grey pollinosity (T5 often with
bases of some setae black) (Fig. 3D)
latigena
` Dorsal surface of T5 without dark-brown or
REVISION OF MICROTROPESA
black spots or if present, spots extend to anterior
margin (Fig. 5C) or ventral surface of T5 (Fig.
AC. T ge jet hee Reunion riha n
8.T5 in caudal view with ground colour completely
obscured by thick, grey pollinosity . campbelli
T5 in caudal view with ground colour not com-
pletely obscured by thick, grey pollinosity (eg.
Figs 4C, 5C) 9
9.T3 without mm; mesoscutum with uniform, thin,
whitish pollinosity .............. 10
T3 with mm; mesoscutum with distinct pattern of
whitish, pollinose stripes and/or spots, at least
onprescutum . a pd eppi ee ne 11
10.Ground colour of abdomen yellowish with dark
dorsomedial vitta (may cover most of dorsum of
abdomen in 9); pf with black hairs ventrally
a Vea BoP a Ai dedi es, IG a flaviventris
Ground colour of abdomen red-brown to black;
pf with orange-yellow hairs ventrally
violacescens
11.Abdomen with whitish, pollinose markings
restricted to median triangle or ' V' on T3; T4
and T5 with uniform, orange pollinosity; vertex
of d very narrow, distance between eyes about
twice width of ocellartriangle . . ochriventris
Abdomen with whitish or grey, pollinose mark-
ings on T3, T4 and T5; T4 and T5 without
orange pollinosity; vertex of d wider, distance
between eyes approaching four times width of
ocellartriangle ..............-.. 12
12.Thoracic pleura with black hairs restricted to dor-
sal mesopleuron, remainder with pale, yellow
hairs; prescutum with three whitish, pollinose
vittae, one median, and two submedian enclos-
ing prst-dc obtusa
Thoracic pleura with black hairs on pleurotergite,
sternopleuron and most of mesopleuron; pres-
cutum with median whitish, pollinose vitta, but
each submedian vitta replaced by pair of
whitish, pollinose spots enclosing anterior and
posterior prst-de .......... sess 13
13.Pf and genae golden pollinose; d with apex of
cerci pointed, surstyli hook-like; 9 often with
dark areas on dorsal T4 and T5 connected
Chg AD ual ey ek I pat RA intermedia
Pf silver pollinose, genae golden pollinose; d
with apex of cerci blunt, surstyli blade-like; 9 al-
ways with dark areas on dorsal T4 and T5
separated by grey pollinosity (Fig. 4C)
nigricornis
Microtropesa campbelli Paramonov, 1951
(Fig. 7A)
Microtropesa campbelli Paramanov, 1951:768;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:761.
MATERIAL EXAMINED. HOLOTYPE 2,
213
Blundell’s, near Canberra, Australian Capital Ter-
ritory, 30.11.1931, T.G. Campbell (ANIC).
DIAGNOSIS. Body length: 10.3 mm. A3 ovoid,
orange; pf with black hairs; genae with yellow
hairs, except dorsally hairs black; prescutum with
pattern of whitish pollinosity (precise pattern
obscured by discolouration); wings light grey,
bases orange-yellow; ground colour of abdomen
yellowish brown except most mesal areas on T3
black; T3 and T4 with thick, grey, pollinose tri-
angles, apex of triangle on T4 very broad enclos-
ing most mesal setae of marginal row; T3 with
single pair of weak mm; T5 entirely grey, pol-
linose with very weak, hair-like setae.
REMARKS. M. campbelli is the only species in
which T5 is entirely grey pollinose. Only the
holotype is known.
Microtropesa canberrae Paramonov, 1951
(Figs 1A-C, 7A)
Microtropesa canberrae Paramanov, 1951:771;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:761.
MATERIAL EXAMINED. HOLOTYPE 2, Black
Mountain, Canberra, ACT, 18.v.1942, T.W. Pickard
(ANIC). OTHER MATERIAL EXAMINED. NSW:
Paddy's R., nr Marulan (18, ANIC). S AUST: Talia
(18, SAM).
DIAGNOSIS. Body length: 11.7-13.3 mm. ¢
cerci and surstyli as in Fig. 3A,B. Fr, dc and acr
long, hair-like; very long hairs on pf, genae and
thorax; pf with extensive black hairing, extending
below level of vibrissae; epistome weakly
projecting (Fig. 1C); A3 ovoid, orange; hairs on
thoracic pleura almost entirely orange-yellow;
hairs on dorsal scutellum orange-yellow; wings
grey, bases orange-yellow; ground colour of ab-
domen reddish-brown to black; T3 with (d) or
without (9) pair of anterior, submedian grey,
pollinose spots; T4 with median, grey, pollinose
triangle; T5 grey pollinose except for pair of large
dark spots extending on to T4; sternites and inner
edges of tergites without pollinosity; T3 without
mm; fore tarsi of female wide (cf Fig. 5D).
REMARKS. M. canberrae, M. sinuata and M.
danielsi form a group of very similar species. M.
canberrae can be distinguished from the latter
two species by a weakly projecting epistome (Fig.
1C), parafacials with extensive black hairing,
extending below the level of the vibrissae, long
214
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. A-C, Microtropesa canberrae (Talia, S.A.); D-G, M. danielsi (D-F, 3km NNW Palmer R. crossing, Qld;
G, 32 km S Theodore, Qld). A, B, E, F, caudal and lateral views of d cerci and surstyli. C, D, lateral view of
ő heads, vestiture omitted. G, caudal view of apex of d cerci and surstyli. HH, head height. A, B, E-G to same
scale.
and hair-like fr, dc and acr and very long hairs on
the parafacials, genae and thorax.
Males, which have not been previously
described, are similar to females but differ as
follows: vertex narrower; fore and mid tarsi not
widened; ground colour of abdominal tergites
reddish-brown (black in 9); T3 with a pair of
anterior, submedian, grey, pollinose spots (absent
in 9); median, grey, pollinose triangle on T4
wider. Male terminalia of M. canberrae closely
resemble those of M. danielsi.
Microtropesa danielsi sp. nov.
(Figs 1D-G, 7A)
MATERIAL EXAMINED. HOLOTYPE QM26036,
3, 3 km NNW Palmer River crossing, 16°04’S.,
144°47°E., Queensland, 19.v.1989, G. & A. Daniels
REVISION OF MICROTROPESA
(QM). Terminalia removed and stored in glycerol in
microvial pinned next to holotype, with duplicate label.
PARATYPES 8 d, same data as holotype (one with
terminalia removed and stored in glycerol in microvial
pinned next to paratype, with duplicate label); 1 d,
same data as holotype except 6.x.1989, L. Ring; 2 d,
same data as holotype except 11.ix.1991, R.C.
Manskie; 1 ¢ , Mt Moffatt Nat. Pk, Mt Moffatt summit,
25°04’S 148°03’E, 23.xi.95; C.J. Burwell (QM26037);
2 d, sandstone hilltop 32 km S Theodore, 25° 10'S
150° 00° 4.x.1991, E.G. Daniels (all UQIC unless
indicated).
DIAGNOSIS. d cerci and surstyli as in Fig.
1E-G. Pf with extensive yellow hairing; epistome
strongly projecting (Fig. 1D); A3 ovoid, orange;
hairs on thoracic pleura almost entirely orange-
yellow; hairs on dorsal scutellum orange- yellow;
mesonotum with thin covering of shifting whitish
pollinosity, thicker surrounding anterior prst-dc;
wings grey, bases orange-yellow; ground colour
of abdomen dark, reddish-brown to black; T3
without grey, pollinose spots; T4 with median,
grey, pollinose triangle about as wide as distance
between subapical scutellar setae; T5 grey pol-
linose except for pair of large dark spots extend-
ing on to T4; sternites and inner edges of tergites
without pollinosity; T3 without mm.
DESCRIPTION. MALE. Head (Fig. 1D). ground
colour orange-yellow; genae and pf golden pol-
linose; pfr and sometimes dorsal pf with black
hairs, those on rest of pf and genae orange-yel-
low; A3 orange, ovoid; Ist aristal segment shorter
than 2nd, 3rd thickened basally.
Thorax. ground colour of mesoscutum black
except posterior yellow intrusion between rows
of dc; mesoscutum with thin covering of shifting,
whitish pollinosity, thicker surrounding anterior
prst-dc in dorsal view; hairs on mesoscutum
mainly black, orange-yellow posteriorly; scutel-
lum orange with orange-yellow hairs; hairs on
pleura orange-yellow.
Wings. dark grey, bases orange-yellow.
Legs. orange-yellow, tarsi darkened.
Abdomen. ground colour of tergites dark red-
brown, black medially; T3 without mm; dorsal
surface of T3 immaculate; T4 with median, grey,
pollinose triangle about as wide as distance be-
tween subapical scutellar setae; T5 grey pollinose
except for pair of large dark spots extending onto
T4; sternites and inner edges of tergites without
pollinosity; hairs on abdomen black except some
yellow hairs on ventral T1+2, Stl and St2.
Body length. 10.6-13.0 mm.
FEMALE. Unknown.
REMARKS. M. canberrae, M. danielsi and M.
sinuata form a group of very similar species. M.
danielsi is most easily distinguished from M.
canberrae by extensive yellow hairing on the
parafacials.
M danielsi is distinguished from most
specimens of M. sinuata by the thinly pollinose,
unpatterned mesonotum. Males of M. danielsi
differ from males, and most females of M. sinuata
by the absence of grey, pollinose spots on T3.
Males of M. danielsi also differ from those of M.
sinuata by a narrower grey, pollinose triangle on
T4 and consistently longer apices of the surstyli
(Fig. 1E-G).
ETYMOLOGY. For Greg Daniels, co-collector of the
holotype.
Microtropesa flaviventris Malloch, 1930
(Figs 2A,B, 7C)
Microtropesa flaviventris Malloch, 1930:101;
Paramonov, 1951:772; Crosskey, 1973:135;
Cantrell & Crosskey, 1989:761.
MATERIAL EXAMINED. HOLOTYPE. 9, Nar-
romine, NSW (ANIC). OTHER MATERIAL EX-
AMINED. QLD: Taroom (1 ? , QDPI); Jondaryan (19,
UQIC); Wellcamp (1d, 19, QDPI); Nobby (1d,
QDPI); 16 km N Boonah (1 9 , QM); 5 km NE Leyburn
(1d, QDPI). NSW: nr Bourke (1G , ANIC); 4 mile Ck,
W of Wollomombi (19, ANIC); Bogan R. (19,
ANIC); Mt Boppy, nr Cobar (28 , ANIC); Taree (19,
BCRI); Barrington Tops (19, ANIC); Round Hill
Fauna Reserve (1d, AM); Orange (12, ANIC); Gos-
ford Dist. (19, 1d, BCRI); Newbridge (1d, 19,
ANIC, 98, 69, BCRI); Penrith (38, 19, BCRI);
Cowra (18, BCRI); Sydney (18, BCRI); Beverly
Hills, nr Sydney (1d, AM); Mt Boyce, Blue Mtns (19,
AM); Sutton Forest (16, BCRD; Berry (18, BCRI);
nr Yass (1G, ANIC); Urana (19, ANIC); Gevogery
[Gerogery?] (19, ANIC); Albury (1d, BCRI); Mt
Gladstone, nr Cooma (16, ANIC); Nimmitabel (19,
ANIC). ACT: Black Mtn (58, 1 9, ANIC); Canberra
(298,39, ANIC); Blundell's, Canberra (12, ANIC);
Mt Gingera (1d, 12, ANIC). VIC: Strathmerton (1d,
ANIC); Hume Weir via Wodonga (18, UQIC);
Alexandra (3 2, ANIC); Genoa (3¢, ANIC); Ballarat
(1d, ANIC); Eagle Point, S of Bairnsdale (1 2, ANIC).
DIAGNOSIS. Body length: 8.4-11.4 mm. ¢
cerci and surstyli as in Fig. 2A, B. Hairs on pf
black, those on genae mostly pale yellow, some-
times with a few black hairs dorsally; mesonotum
in caudal view blue-black with thin covering of
whitish pollinosity; wings light grey, almost
hyaline, base yellowish; ground colour of ab-
domen yellowish, excavation of T1+2 blue-
216
MEMOIRS OF THE QUEENSLAND MUSEUM
0.025 mm
FIG. 2. A, B, Microtropesa flaviventris; C-E, M. intermedia. A-D, caudal and lateral views of d cerci and surstyli.
E, lateral view of cephalopharyngeal skeleton of 1st instar larva. AS, accessory sclerite; DC, dorsal cornu; IR,
intermediate region; MH, mouth hook; S, sclerite of salivary gland; VC, ventral cornu. A-D to same scale.
black, continued as dorsomedial vitta extending
to T5; T3 and T4 in caudal view each with indis-
tinct, median triangle of thin, whitish pollinosity
which hardly obscures ground colour, sometimes
only anterior base of triangle present on T3; T5
with extensive thin, whitish pollinosity, some-
times with pair of yellow spots without pol-
linosity in caudal view; T3 without mm.
REMARKS. M. flaviventris most closely
resembles M. obtusa and M. ochriventris from
which it can be distinguished by the uniform, thin
pollinosity on the mesonotum and the absence of
mm on T3. The blue-black, abdominal vitta of
M. flaviventris is broader in females than males.
In females it is about as broad as the distance
between the basal scutellar setae, while in males
it is, at most, a little wider than the distance
between the subapical scutellar setae.
Hardy (1939) incorrectly considered M.
flaviventris a junior synonym of M. obtusa.
Microtropesa intermedia Malloch, 1930
(Figs 2C-E, 7E)
Microtropesa intermedia Malloch, 1930:100; Hardy,
1939:35; Paramonov, 1951:775; Crosskey,
1973:135; Cantrell & Crosskey, 1989:761.
MATERIAL EXAMINED. HOLOTYPE. d,
Eidsvold, Queensland, 20.iv.1924, Bancroft (ANIC).
OTHER MATERIAL EXAMINED. QLD: Yeppoon
(16, ANIC); Theodore (19, UQIC); Electra State
Forest, 25 km S Bundaberg (18, ANIC); Bluff Ra.
foothills, Biggenden (1 9, ANIC); SW Bluff Ra., Big-
genden (1d , ANIC); Rockpool Gorge, Mt Walsh Nat.
Pk, Biggenden (2d, 19, ANIC); Coalstoun Lakes, nr
Biggenden (3d, ANIC); S Boolboonda Ra., via Mt
Perry (1 9, ANIC); nrSeary's Ck, Cooloola (19 , QM);
Noosa Nat. Pk (19, UQIC); Tibrogargan Ck, Glas-
shouse (19, UQIC); Beeberum [Beerburrum?] (18,
UQIC); Point Lookout, North Stradbroke Is. (1d,
UQIC); Brisbane (18, 29, QM); St. Lucia, Brisbane
(18,19,UQIC,29, ANIC); Stradbroke Is. (1 9 , QM);
Stockyard Ck, SE Capalaba (1d , UQIC); Sunnybank,
Brisbane (26, ANIC); 4 km WNW Mt Cotton (19,
UQIC); The Blunder, Brisbane (1 2, ANIC); 16 km N
Boonah (12, UQIC); Amiens State Forest, nr Stan-
thorpe (1 2, UQIC). NSW: Clyde Mtns, nr Braidwood
(19, ANIC); Alpine Ck, Kiandra (29 , ANIC). ACT:
Canberra (18, ANIC); Blundell’s (1 8, ANIC); Lees
Ck, Brindabella Ra. (18, 19, AM); Bendora (19,
ANIC); Mt Gingera (18, 39, ANIC); Cotter R. (1d,
ANIC). VIC: 2 miles [3.5 km] NW Porepunkah (1d,
ANIC). Tasmania: Rupert Point, Pieman R. (1d,
ANIC); 9 km SE Miena (13 ? , UQIC); Derwent Bridge
(16, ANIC). Other: no data (19, QDPI, 19, ANIC).
REVISION OF MICROTROPESA
217
FIG. 3. A-D, Microtropesa latigena; E, F, M. near latigena ; G, H, M. longimentum holotype. A, B, E, F, caudal
and lateral views of ó cerci and surstyli. C, G, lateral view of 9 heads, vestiture omitted. D, caudal view of 4th
apparent abdominal tergite (TS). H, apex of wing.
DIAGNOSIS. Body length 11.7-13.5 mm.
ó cerci and surstyli as in Fig. 2C,D. Pf and genae
golden pollinose; A3 black, base yellow or
orange-yellow; mesonotum with bronze reflec-
tion and distinct pattern of whitish pollinosity
which shifts markedly when viewed dorsally and
then caudally; mesonotum with median, pol-
linose vitta, broken at transverse suture, extend-
ing to base of scutellum in most specimens;
prescutum, in dorsal view, with a pair of pollinose
spots enclosing anterior and posterior setae of
each row of prst-dc; wings grey, bases orange-
yellow; hairs on ventral surface of basal node of
R4+5 black, sometimes with a few yellow hairs;
ground colour of T3 and T4 red-brown to black,
both with median, grey, pollinose triangles; T5
grey pollinose with pair of large, dark spots; T3
with one or more pairs of mm.
REMARKS. M. intermedia is very similar to M.
nigricornis from which it is best separated by the
golden pollinose pf and genae, the bronze reflec-
tion and shifting pollinose markings on the
mesonotum and the colour of the hairs on the
ventral node of R4+5. Some specimens of M.
intermedia are similar to those of M. nigricornis
in that the grey, pollinose markings on the ab-
domen are distinct. However, in other specimens
of M. intermedia the pollinose triangles on T3 and
T4 are thin with poorly defined margins. In addi-
tion, some females of M. intermedia can be
separated from those of M. nigricornis by the
218
dark spots on T5 extending to the anterior margin
of the tergite. The male genitalia of M. intermedia
(Fig. 2C,D) have the apex of the cerci pointed and
the surstyli blade-like and differ markedly from
those of M. nigricornis (Fig. 4A,B).
Ist-instar larvae were recovered from the uterus
of a female M. intermedia. Their banded appear-
ance and strongly arched dorsal cornua of the
cephalopharyngeal skeleton resemble the larvae
of Cuphocera (Tachinini) and the Linnaemyini
described by Cantrell (1988). They differ in the
distinctly serrate edges of the mouth-hooks (Fig.
2E). Body length: 0.76 mm (n = 10).
Microtropesa latigena Paramonov, 1951
(Figs 3A-D, 7A)
Microtropesa latigena Paramonov, 1951:769;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:761.
MATERIAL EXAMINED. HOLOTYPE 9, Kal-
goorlie, WA, August, L.J. Newman (ANIC). OTHER
MATERIAL EXAMINED. NSW: Broken Hill (18,
AM). SAUST: Coonamoree [? Coonamoranie] (1d ,
ANIC); Stuart Ra. (19, ANIC); 37 miles [60 km] ENE
White Well [White Wells] (2d , ANIC); 19 miles [31
km] ENE Eucla (12, ANIC). WA: Mt Squires (19,
SAM); Nanambinia HS, SW Balladonia (1 2, ANIC).
DIAGNOSIS. Body length: 10.9-12.6 mm. ĝ
cerci and surstyli as in Fig. 3A,B. A3 ovoid,
brown (darker in d), base yellow; pf very wide
(Fig. 3C), with black hairs; hairs on genae mostly
black; anterior mesonotum with distinct pattern
of whitish pollinosity, prescutum with acr
enclosed in median, pollinose vitta bordered
laterally by pair of black, submedian vittae ex-
tending on to scutum, narrowly interrupted at
transverse suture; each row of prst dc enclosed in
pollinose vitta bordered laterally by black vitta
extending on to scutum, broadly interrupted at
transverse suture; wings light grey, bases
brownish-yellow; ground colour of T3 and T4 in
9 black, in d reddish-yellow, black medially; T3
and T4 with median, grey, pollinose triangles
usually with sharply defined margins; T5 grey,
pollinose with pair of small, submedian, black or
dark-brown spots posterodorsally (Fig. 3D),
often second pair of dark spots posteroventrally;
bases of some setae on T5 black; T3 with one or
two pairs of mm (weaker in d).
REMARKS. A male and female in ANIC from
near Alice Springs are similar in most respects to
M. latigena, to which they key, but the abdominal
ground colour of the male is darker, the mm on
MEMOIRS OF THE QUEENSLAND MUSEUM
T3 are extremely weak in the female and absent
in the male, the pollinose triangles on T3 and T4
have poorly defined margins and the cerci and
surstyli of the male differ slightly (Fig. 3E,F).
They probably represent an undescribed species.
Microtropesa longimentum sp. nov.
(Figs 3G,H, 7A)
MATERIAL EXAMINED. HOLOTYPE @&, Lake
Surprise area, Simpson Desert, 135° 5’E., 26? 4’S., S
AUST, 13.ix.1971, T.F. Houston (SAM).
The holotype has some setae, hairing and pollinosity
abraded, however it differs radically from the other
known Microtropesa and description is warranted.
DIAGNOSIS. M. longimentum is recognisable
by its greatly elongated mentum (Fig. 3G) and
closed cell R5 in the wing (Fig. 3H).
DESCRIPTION. FEMALE. Head (Fig. 3G).
Ground colour yellow; viewed laterally, pfr, pf
and genae thickly whitish pollinose; pfr with fine
black hairs; pf and genae with sparse, strong,
black hairs, many seta-like; A3 quadrangular,
wider at apex, brownish; Ist aristal segment elon-
gate, almost as long as 2nd; 3rd aristal segment
about twice as long as combined length of 1st and
2nd, basal third thickened; proboscis with men-
tum longer than head height; labellum reduced.
Thorax. Ground colour of mesonotum black,
except posterior margin, posterior intrusion be-
tween rows of dc and lateral margin enclosing ia
yellow; scutellum yellowish; pattern of whitish
pollinosity on mesonotum as follows; narrow,
median, pollinose vitta between acr; pollinose
vitta enclosing each row of prst-dc; prst-ia and
presutural seta enclosed in pollinosity; humeral
callus pollinose; pollinosity on scutum partially
abraded but with large, median patch enclosing
both rows of acr; hairs on mesonotum and scutel-
lum black.
Wings (Fig. 3H). Cell R5 closed; wings light
grey, bases yellowish-brown.
Legs. Brownish-yellow, apical tarsal segments
darkened.
Abdomen. Ground colour of tergites, in dorsal
view, shining-black; T3 and T4 each with thick,
grey, pollinose, anterior band with median, trian-
gular extension reaching posterior margin; T5
grey pollinose except for pair of large,
posterodorsal black spots; venter of abdomen
grey pollinose, thin on sternites, yellow ground
colour showing through; T3 with pair of very
strong mm.
REVISION OF MICROTROPESA
219
FIG. 4. A-C, Microtropesa nigricornis; D-F, M. obtusa; G, H, M. ochriventris. A, B, E-H, caudal and lateral
views of d cerci and surstyli. C, caudal view of 4th apparent abdominal tergite (T5). D, d antenna. A, B, E-H
to same scale.
Body length. 10.6 mm.
MALE. Unknown.
ETYMOLOGY. Latin noun in apposition, for the long
mentum.
Microtropesa nigricornis Macquart, 1851
(Figs 4A-C, 7F)
Microtropesa nigricornis Macquart, 1851:199, pl. 21,
fig. 5; Townsend, 1939:14; Hardy, 1939:35; Engel,
1925:344; Paramonov, 1951:774; Crosskey,
1971:278; Crosskey, 1973:135; Cantrell &
Crosskey, 1989:761.
MATERIAL. LECTOTYPE éd (designated Crosskey,
1971:278), TAS (MNHN) [not examined but notes on
important characters provided by Dr. L. Matile of
MNHN]. MATERIAL EXAMINED NSW: Gibraltar
Ra. Nat. Pk (26,29 , UQIC); NE [New England?] Nat.
Pk (1d, ANIC); 6 km NE Bilpin (1d, AM); 5 km E
Bilpin (16, 29, AM); Jinki Ck, Blue Mtns (19, AM);
3 km S Mount Wilson (12, AM); Mt Tomah (28,
AM); 3 miles [5 km] SW Mt Tomah (28 , AM); Mount
Victoria (1 8, ANIC); Hat Hill (10d, 1 9,AM); Black-
heath (1d, ANIC); Lawson, Blue Mtns (3d, AM);
Goondera Ridge, Royal Nat. Pk (22 , AM); Clyde Mts,
nr Braidwood (1d, ANIC); Olson's [Olsen's?]
Lookout, Snowy Mtns (18, ANIC); Sawpit Ck, Mt
Kosciusko (1 9, ANIC). ACT: Canberra (1 8, ANIC);
Mt Coree (58, 19, ANIC); Blundell's (16, ANIC);
Lees Ck, Brindabella Ra. (18, 1 9, AM); Mt Franklin
(3d, ANIC). TAS: Granite Point Reserve, Bridport
(52, UQIC); Mt George (1 8, UQIC); Turners Beach,
nr Ulverstone (1d, UQIC); Don R., nr Don (29,
UQIC); Hobart (1d, ANIC); Tasmania (26, 19,
ANIC).
220
DIAGNOSIS. Body Length 11.8-12.9 mm. d
cerci and surstyli as in Fig 4A,B. Pf silver pol-
linose; genae golden pollinose; A3 black, base
yellow or orange-yellow; mesonotum black with
distinct pattern of whitish pollinosity which shifts
little when viewed dorsally and then caudally;
mesonotum with median, pollinose vitta, broken
at transverse suture, extending to base of scutel-
lum in most specimens; prescutum with pair of
pollinose spots enclosing anterior and posterior
setae of each row of prst-dc; wings grey, bases
orange-yellow; hairs on ventral surface of basal
node of R4+5 yellow, sometimes with a few black
hairs; ground colour of T3 and T4 red-brown to
black, both with median, grey, pollinose tri-
angles; T5 grey pollinose with pair of large, dark
spots; T3 with one or more pairs of mm.
REMARKS. M. nigricornis is very similar to M.
intermedia from which it can be distinguished by
the silver pollinose pf, the black mesonotum with
stable pollinose markings and the colour of the
setae on the ventral surface of the basal node of
R4+5. The grey pollinose markings on the ab-
domen of M. nigricornis are always thick with
sharply defined margins and the dark spots on T5
are always bordered anteriorly by pollinosity in
both sexes (Fig. 4C). Male genitalia of M.
nigricornis (Fig. 4A,B) have the apex of the cerci
blunt and the surstyli hook-like and differ
markedly from those of M. intermedia (Fig.
2C,D).
Microtropesa obtusa (Walker, 1853)
(Figs 4D-F, 7B)
Tachina obtusa Walker, 1853:274.
Echinomyia stolida Walker, 1858:196, Synonymy by
Austen, 1907:330.
Gerotachina obtusa Townsend, 1916:152; Townsend,
1932:40.
Microtropesa flavitarsis Malloch, 1929:288; Malloch,
1930:100; Paramonov, 1951:773; Crosskey,
1973:135; Cantrell & Crosskey, 1989:761.
Synonymy by Hardy, 1939:36.
Microtropesa obtusa (Walker); Hardy, 1939:36;
Paramonov, 1951:771; Crosskey, 1973:135;
Cantrell & Crosskey, 1989:761.
MATERIAL EXAMINED. LECTOTYPE 9 desig-
nated Townsend (1932:40), NSW (BMNH). Holotype
of E. stolida d, NSW (BMNH). Holotype of M.
flavitarsis 6, TAS (AM). OTHER MATERIAL EX-
AMINED. ACT: Canberra (3d, 39, ANIC). S AUST:
Roseworthy (19, SAM). WA: Kojarena (19, ANIC);
Geraldton (18 , 1 9, ANIC); Bunbury (18, ANIC, 28,
WADA).
MEMOIRS OF THE QUEENSLAND MUSEUM
DIAGNOSIS. Body length: 11.2-12.5 mm. &
cerci and surstyli as in Fig. 4E,F. Pf black haired;
genae with numerous black hairs dorsally, rest
with pale yellow hairs; A3 of 9 ovoid, of ó
triangular, produced towards face distally (Fig.
4D); anterior mesonotum with distinct pattern of
whitish pollinosity; acr enclosed in median pol-
linose vitta bordered laterally by pair of black
submedian vittae which are narrowly interrupted
at transverse suture; each row of prst-dc enclosed
in pollinose vitta bordered laterally by black vitta
which is broadly interrupted at transverse suture;
ground colour of abdomen orange-yellow with
black, dorsomedial vitta; T3 and T4 in caudal
view each with median triangle of thin, whitish
pollinosity which usually obscures ground
colour; T5 with pair of large spots without pol-
linosity in caudal view; T3 with 2-4 pairs of
strong mm.
REMARKS. Microtropesa obtusa most closely
resembles M. flaviventris and M. ochriventris.
The numerous black hairs on the genae, the pat-
tern of pollinose vittae on the mesonotum and the
strong mm on T3 distinguish M. obtusa from M.
flaviventris. The pattern of whitish pollinosity on
the abdomen, especially T4 and T5 distinguishes
M. obtusa from M. ochriventris.
The triangular A3 of male M. obtusa is charac-
teristic.
Microtropesa ochriventris Malloch, 1929
(Figs 4G,H, 7D)
Microtropesa ochriventris Malloch, 1929:287, fig.2;
Malloch, 1930:100; Paramonov, 1951:776;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:762.
MATERIAL EXAMINED. HOLOTYPE 2, Allyn
Range, Barrington Tops, NSW, February 1925, Syd-
ney University zoological expedition (ANIC).
OTHER MATERIAL EXAMINED. QLD: Pomona
(1d, UQIC); Wild Horse Mtn, nr Beerwah (8d,
QDPI); Mt Tibrogargan (18, UQIC); Mt Mitchell, nr
Aratula (1d, UQIC). NSW: hilltop, 24 km W Grafton
(18, UQIC); 25 km W Grafton (1 2, QDPI), Barwick
R., NE [New England?] Tableland (1c , ANIC); Point
Lookout, nr Ebor (38, ANIC); 12 km SW Ebor (29,
UQIC); Allyn Ra., Barrington Tops (22, ANIC); Hat
Hill (1 9, AM); Kuringai [Ku-ring-gai] Chase Nat. Pk
(19, BCRI); Kanangra Boyd Nat. Pk (1d, BCRD;
Nowra (19, ANIC); 9 miles [14.5 km] NW Braidwood
(19, ANIC); Rutherford Ck, Brown Mtn, nr Nim-
mitabel (19, AM). ACT: Black Mtn Reserve (19,
ANIC); Canberra (1 8, ANIC); Mt Coree (18, ANIC);
Lees Ck, Brindabella Ra. (1 8, AM); Mt Gingera (19,
ANIC). Victoria: Bogong, via Mt Beauty (19, UQIC);
REVISION OF MICROTROPESA
2mm
0.8 mm
D
221
0.4 mm
FIG. 5. Microtropesa sinuata. (E, F, Mt Tinbeerwah, Qld; G, 25 km W Bridport, Tas.; H, Isla Gorge, Qld; I,
Swan R, W.A.). A, lateral view of ó head, vestiture omitted. B, apex of wing. C, caudal view of 4th apparent
abdominal tergite (T5). D, dorsal view of 9 fore tarsus. E, F, caudal and lateral view of d cerci and surstyli.
G-I, caudal views of apices of d cerci and surstyli. E-I to same scale.
Victoria (12, WADA). Tasmania: Strzelecki Peaks,
Flinders Is. (1d, ANIC); Strahan (1c , ANIC); Hobart
(12, ANIC). South Australia: 2 km SE of Montacute
(19, ANIC). Other: Gordon [N.S.W. or Tas.?] (19,
ANIC).
DIAGNOSIS. Body length 10.7-12.3 mm. d
cerci and surstyli as in Fig. 4G,H. Vertex of d
very narrow, less than twice width of ocellar
triangle; Pf black haired; genae with numerous
black hairs dorsally, rest with pale yellow hairs;
A3 of d and 9 ovoid; mesonotum with bronze
reflection; anterior mesonotum with distinct pat-
tern of whitish pollinosity; acr enclosed in
median pollinose vitta bordered laterally by pair
of black submedian vittae which are narrowly
interrupted at transverse suture; black submedian
vittae coalesce anteriorly; each row of prst-dc
enclosed in pollinose vitta bordered laterally by
black vitta which is broadly interrupted at
transverse suture; ground colour of abdomen
orange with black, dorsomedial vitta which is
largely obscured by pollinosity; T3 with median
triangle or ' V" of whitish pollinosity; rest of ab-
domen with orange pollinosity (thicker on T5)
except for pair of orange spots with black mesal
edges on T3; T3 with one or more pairs of mm.
REMARKS. M. ochriventris can be separated
from the similar M. flaviventris and M. obtusa by
the extensive orange pollinosity on the abdomen.
The very narrow vertex of males is characteristic.
Hardy (1939) incorrectly considered M.
ochriventris a junior synonym of M. obtusa.
222
Microtropesa sinuata (Donovan, 1805)
(Figs 5, 7G)
Musca sinuata Donovan 1805:[166], pl. [41].
Microtropesa sinuata (Donovan); Macquart,
1846:313; Macquart, 1851:199; Engel, 1925:344,
fig.B; Townsend, 1932:40; Townsend, 1939:13;
Hardy, 1939:34; Paramonov, 1951:776; Crosskey,
1973:135; Cantrell & Crosskey, 1989:762.
Tachina bura Walker, 1849:760. Synonymy by Hardy
1939:34.
Microtropesa ignipennis Brauer, 1899:510 (unavail-
able name published in synonymy with sinuata
Donovan, not subsequently made avail-
able);Crosskey, 1971:278.
Microtropesa latimana Malloch, 1929:287, fig. 3a;
Malloch 1930:100. Synonymy by Hardy 1939:34.
Some early references are given by Hardy (1939:34).
MATERIAL. The holotype of M. sinuata is not in
London as stated by Townsend (Crosskey, 1971) and
is presumed lost. Lectotype of T. bura d , designated
by Crosskey (1973:163), Tasmania, Rev. J. Ewing
(BMNH) [examined]. Holotype of M. latimana 9,
Queensland (AM) [examined]. OTHER MATERIAL
EXAMINED. QLD: Kroombit Tops (2d, 19, UQIC,
18, QM); Mt Scoria, 6 km S Thangool (48, UQIC);
Expedition Ra. (18, AM); Marlong Arch, Mt Moffatt
Nat. Pk (2d, QM); Mt Moffatt summit, Mt Moffatt
Nat. Pk (46, QM); Mt Moffat [Moffatt] Nat. Pk (1d,
UQIC); Isla Gorge Nat. Pk (58, UQIC); Mt Woowoon-
ga, 54 miles [87 km] SW Bundaberg (1d, ANIC);
Woowoonga Ra., SW Bundaberg (18, ANIC); Bluff
Ra., Biggenden (18, ANIC); Mt Tinbeerwah, nr
Cooroy (2d, UQIC, 66, QDPI); Cooroy (1d, AM);
Beacon Hill, 16 km W Imbil (18, UQIC); 16 km SW
Kumbia (18, UQIC); Sunday Ck Fire Tower, Jimna
State Forest (18, QM); Yarraman (18, UQIC); Wild
Horse Mtn, nr Beerburrum (26 , UQIC); Mt Glorious
(18, ANIC); Brisbane (68, QDPI, 1d, ANIC);
Toowong (1d, QM); 13 km N Dunwich, North
Stradbroke Is. (18, UQIC); Flinders Peak (138, 49,
UQIC, 18, QM); Mt Edwards, nr Aratula (3d , UQIC);
Bollon (1d, ANIC); Mt Mitchell, nr Aratula (2d,
UQIC); Miami, G. [Gold] Coast (19, UQIC); Mt
Greville, nr Aratula (23, UQIC); Lamington Nat. Pk
(5d, UQIC); [Lamington?] Nat. Pk (19, QM);
Amiens, nr Stanthorpe (28 , AM, 1 9, ANIC); Amiens
State Forest (18, UQIC); Glen Aplin (38, UQIC); Mt
Marlay, nr Stanthorpe (18, UQIC); Queensland (19,
AM). NSW: Bald Rock Nat. Pk, 25 km SE Stanthorpe
(18, UQIC); Gibraltar Ra. Nat. Pk (18, 19, UQIC);
Grafton (26 , ANIC, 1d, AM); 24 km W Grafton (1d ,
UQIC); 25 km W Grafton (1d, UQIC, 28, QDPI); 15
km W South Grafton (18, UQIC); 20 km W South
Grafton (1d , UQIC); 24 km W South Grafton (1d,
UQIC); Mt Kaputar Nat. Pk (28 , AM); 14 km SW Ebor
(19, UQIC); New England Nat. Pk (18, ANIC); Ural-
la (1d, ANIC); Dangar Falls (38, ANIC); Warrum-
bungle Nat. Pk, nr Coonabarabran (1 2, ANIC); Cobar
(1d, UQIC); Mt Boppy, nr Cobar (78, ANIC, 1d,
MEMOIRS OF THE QUEENSLAND MUSEUM
AM); Mt Gibraltar (1d, AM); Barrington R. (19,
ANIC); W end Dilgy Ck Circle Road, N section Bar-
rington Tops State Forest (19, AM); Mt Arthur, 3
miles [5 km] W Wellington (1d, ANIC); Round Hill
Fauna Reserve (59, AM); Toronto (18, UQIC, 2d,
1 9, ANIC); Orange (1d, ANIC); Clarence, Blue Mtns
(39, AM); 3 km NE Bilpin, nr Kurrajong (12, AM);
nr Mt Bell (18, AM); Mt York, Blue Mtns (1 9$, AM);
Mt Banks, Blue Mins (16, AM); Blackheath (29,
ANIC); Wentworth Falls (29 , ANIC); Lindfield (19,
ANIC); Ashfield (19, AM); Baldy Bill Fire Rd,
Kanangra Plateau (19, AM); Macquarie Fields (19,
BCRI); Goondera Ridge, Royal Nat. Pk (1d, AM);
Wee Jasper (1 9, ANIC); Clyde Mtns, nr Braidwood
(128, ANIC); Clyde Mtns, E slope (18, ANIC); Tal-
bingo (1d, ANIC); Durras North (1d, ANIC);
Roseville (1 8, BCRI). ACT: Black Mtn (1 8, ANIC);
Canberra (1d, 12, QDPI); Mt Coree (19, ANIC);
Blundell's (12, ANIC); Queanbeyan (18, UQIC);
Lees Spring, Brindabella Ra. (19, ANIC). VIC:
Wyperfield Nat. Pk (1d, AM); Mt William, Grampians
(16, ANIC); Gisborne (26, ANIC, 18, QM); Killara
(1d, ANIC); Melbourne (1d, SAM); Mt Dandenong
(18, ANIC). TAS: Mt William Nat. Pk (1d , UQIC);
25 km W Bridport (18, UQIC); Lefroy (1d, SAM);
Freycinet Nat. Pk (1d, ANIC); Tasmania (2d, 19,
SAM). South Australia: Sleaford Bay (22, ANIC); Mt
Lofty (1d, 19, SAM). WA: 6 miles [9.5 km] SW
Mullewa (19, ANIC); Three Springs (19, WADA);
Moora (1d, WADA); Toodyay (1d, SAM); [?]
Cunderdin (19, QM); Perth (19, ANIC); Crawley,
Perth (1 9, ANIC); Swan R. (18,29, ANIC); Bedfor-
dale (19, WADA); Gleneagle (19, WADA); Mt
Ragged (1d, ANIC); 24 miles [39 km] E Pingrup (1 9,
ANIC); Margaret R. mouth (19, SAM); Stirling Ra.
(16, WADA); Albany (18, ANIC). Other: 1590 (18,
QDPI); no data (19, QM, 19, SAM, 18, 19, QDPI).
DIAGNOSIS. Body length: 10.2-14.6 mm. d
cerci and surstyli as in Fig. SE-I. Pf usually with
extensive yellow hairing, rarely with extensive
black hairing but never extending below level of
vibrissae and at least ventral third of pf with
yellow hairs; epistome strongly projecting (Fig.
5A); A3 ovoid, orange; hairs on thoracic pleura
almost entirely orange-yellow; hairs on dorsal
scutellum orange-yellow; prescutum with
whitish pollinosity developed into pattern of
stripes and spots; wings grey, bases orange-yel-
low; ground colour of abdomen dark, reddish-
brown to black; T3 usually with pair of anterior,
submedian, grey, pollinose spots (rarely absent in
9); T4 with median, grey, pollinose triangle
wider than distance between subapical scutellar
setae in d; T5 grey pollinose except for pair of
large dark spots extending on to T4; sternites and
inner edges of tergites without pollinosity; T3
without mm.
REVISION OF MICROTROPESA
A
223
FIG. 6. A, B, Microtropesa violacescens; C, D, M. viridescens. A-D, caudal and lateral views of d cerci and
surstyli.
REMARKS. M. canberrae, M. danielsi and M.
sinuata form a group of very similar species.
Most specimens of M. sinuata differ from M.
canberrae by extensive yellow hairing on the
parafacials. Rarely the parafacials are black
haired as in M. canberrae, however, in M. sinuata
black hairs never extend below the level of the
vibrissae.
Most specimens of M. sinuata can be separated
from M. danielsi by the pattern of whitish pol-
linose spots on the prescutum of the mesonotum.
Males and most females of M. sinuata differ from
males of M. danielsi by the pair of anterior, sub-
median, grey, pollinose spots on T3. Males of M.
sinuata also differ from those of M. danielsi by a
wider grey, pollinose triangle on T4 and consis-
tently shorter apices of the surstyli (Fig. 5E-I).
Microtropesa skusei Bergroth, 1894
(Fig. 7B)
Microtropesa skusei Bergroth, 1894:73; Engel,
1925:345; Paramonov, 1951:767; Crosskey,
1973:135; Cantrell & Crosskey, 1989:762.
MATERIAL. HOLOTYPE 2, Coomooboolaroo,
near Duaringa, Queensland (most likely depository
Zoological Museum, Helsinki, not located by Crosskey
(1973), probably lost).
DIAGNOSIS. As for M. viridescens except hairs
on palpi black.
REMARKS. Bergroth (1894) described M.
skusei from one or more females which have not
been located. Females of M. viridescens match
the original description of M. skusei except for the
hairs on the palps. According to Bergroth (1894)
the hairs on the palps of M. skusei are black while
those on the palps of M. viridescens are yellow,
occasionally with a few black hairs.
Hardy (1939) incorrectly considered M. skusei
a junior synonym of M. sinuata.
Microtropesa violacescens Enderlein, 1937
(Figs 6A,B, 7B)
Microtropesa violacescens Enderlein, 1937:441;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:762.
Microtropesa sinuata; Malloch, 1928:614; Malloch,
1929:287, figs 1, 3b; Malloch, 1930:100.
Microtropesa fallax Hardy, 1939:35; Paramonov,
1951:777. Synonymy by Crosskey, 1973:136.
MATERIAL. Lectotype of M. violacescens ó (desig-
nated Crosskey, 1973:161), Herberton, QLD (DEI)
[not examined]. Holotype of M. fallax 9, Brisbane,
Queensland [not located, probably lost]. MATERIAL
EXAMINED. NT: 7 km NNW Cahills x-ing, E Al-
ligator R. (19, ANIC); Manton Dam, 52 km SSE
Darwin(1d , ANIC). QLD: Mt White, Coen (1 8, AM);
Cooktown (1d , ANIC); 12 km N Palmer R. x-ing (19,
UQIC); 8 km SSW Atherton (7d, 19, UQIC);
Bluewater, Paluma Ra., NW Townsville (19 , UQIC);
Biloela (18, QDPI); Marlong Arch, Mt Moffatt Nat.
Pk(13d,QM); Mt Moffat [Moffatt] Nat. Pk (1 8, UQIC,
3d, QM); sandstone hilltop 32 km S Theodore (108,
224 MEMOIRS OF THE QUEENSLAND MUSEUM
e M.ochriventris
. campbelli
. canberrae
*remp
ISSIE
. danielsi og
. latigena Ta
A . longimentum
M. intermedia
^ M. obtusa
+ M. skusei Qa
e M. violacescens ^ 08
u M. viridescens
e M. nigricornis
e M. flaviventris
è V IIG
FIG. 7. A-G, Distribution of Microtropesa spp. in Australia.
REVISION OF MICROTROPESA
UQIC); Isla Gorge Nat. Pk (1 d, UQIC); Mt Walsh Nat.
Pk, north peak (1d, ANIC); Bluff Ra. (1d, ANIC);
Gayndah (1d, AM); Coast Ra., Biggenden (1d,
ANIC); Buderim Mtn, nr Mooloolah (1 8, BCRI); Mt
Beerwah, via Glasshouse (26, UQIC); Glasshouse
Mtns Lookout (18, UQIC); Moreton Is. (1, QM);
Brisbane (22, QM, 2d, QDPI); Palmerston (19,
QDPI); Mt Gravatt (19, QDPI); Millmerran (29,
ANIC, 1¢, UQIC); Cunnamulla (1d , AM); Amiens,
nr Stanthorpe (18, AM); Stanthorpe (18, QDPI, 19,
UQIC); Palingyard (12, ANIC). New South Wales:
Graman (1d, BCRI); Warrumbungle Nat. Pk (2d,
AM); Bogan R. (22,62, ANIC); Stephens Peak (3d,
QDPI); Round Hill Fauna Reserve (26, 19, AM, 19,
GD); 6 km NE Bilpin, nr Kurrajong (19, AM);
Hazelbrook (19, ANIC); East Minto (1d, AM).
Western Australia: Geraldton (1 d, SAM, 18, WADA,
13,19, ANIC); Rottnest Is. (3d, 1 9$, ANIC, 12,19,
WADA); Garden Is. (19, ANIC); White Lake, Rock-
ingham (19, ANIC); Cape Peron, 40 km SW Perth
(38, UQIC); 10 km N Mandurah (4d , UQIC); Augus-
ta (28 , UQIC); King George Sound (2 2, AM), Other:
no data (16, 49 , QDPI).
DIAGNOSIS. Body length: 12.0-16.0 mm. d
cerci and surstyli as in Fig. 6A, B. A3 ovoid,
orange, apex sometimes brown; pf and genae
mostly with orange-yellow hairs, those on dorsal
pf sometimes black; mesonotum, in dorsal view,
with thin covering of whitish pollinosity; dorsal
surface of scutellum with black hairs; wings grey,
bases orange-yellow; ground colour of ab-
dominal tergites dark, reddish-brown to black
(often with bluish reflection); abdomen, in caudal
view, usually with grey, pollinose markings as
follows; T3 with anterior, pollinose band inter-
rupted medially; T4 with median, pollinose tri-
angle extended laterally as anterior band, T5 with
pair of large, dark spots separated from anterior
margin by pollinosity; T3 without mm.
REMARKS. In many specimens the grey, pol-
linose markings on the abdomen are thin and
weak and only fully apparent when the abdomen
is seen in caudal view. The anterior, pollinose
band on T3 can be represented only by a pair of
submedian spots, or is rarely absent.
Microtropesa viridescens Paramonov, 1951
(Figs 6C,D, 7B)
Microtropesa viridescens Paramonov, 1951: 765;
Crosskey, 1973:135; Cantrell & Crosskey,
1989:762.
MATERIAL EXAMINED. HOLOTYPE d, Canber-
ra, ACT, 7.11.1948, Paramonov (ANIC). OTHER
MATERIAL EXAMINED. Queensland: Mt Moffatt
225
summit, Mt Moffatt Nat. Pk (3d, QM). NSW: Bogan
R. (18, ANIC). S AUST: 5 km SW Mt Sarah HS, N
Oodnadatta (19 , ANIC); 20 [miles? 2 32 km] SW Port
Augusta (12, ANIC). WA: 15 km NE Mt Singleton
(19, ANIC); Pinjarra (1d , ANIC).
DIAGNOSIS. Body length: 10.8-12.0 mm. 3
cerci and surstyli as in Fig. 6C, D. Pf and genae
with orange-yellow hairs; palpi mostly yellow
haired, occasionally with a few black hairs;
mesonotum with uniform, thin covering of
whitish pollinosity; wings hyaline with black
markings at base; ground colour of thoracic
pleura, scutellum and abdomen dark-brown or
black; legs black; T3, T4 and T5 each with silver,
pollinose, anterior band interrupted medially.
REMARKS. Females of M. viridescens, which
have not been previously described, are similar to
males but differ as follows: vertex, fore and mid
tarsi wider; mesonotum with coppery-green
reflection (greenish in d); abdominal ground
colour shining black, with bluish (not greenish)
reflection.
The only apparent difference between M.
viridescens and M. skusei is the colour of the hairs
on the palps. It is likely that M. viridescens is a
junior synonym of M. skusei especially given that
M. viridescens occurs at Mt Moffatt, about 200
km from the type locality of M. skusei.
ACKNOWLEDGEMENTS
I thank the following curators for the loan of
specimens: Dr D.K. McAlpine (AM), Dr P.S.
Cranston (ANIC), Dr M.J. Fletcher and Dr G.R.
Brown (BCRI), Mr G. Daniels (GD), Dr B.K.
Cantrell (QDPI), Mr E.C. Dahms (QM), Dr E.G.
Matthews (SAM); Miss M.A. Schneider (UQIC)
and Mr K.T. Richards (WADA). Thanks also to
Dr L. Matile (MNHN) for examining the types of
M. nigricornis.
I am grateful to Dr E.M. Exley and Mr G.
Daniels for their critical comments and to Mr P.
Avern and Mr G. Thompson for their technical
assistance. I am especially indebted to Dr B.K.
Cantrell for his friendly and helpful supervision.
LITERATURE CITED
AUSTEN, E.E. 1907. The synonymy and generic posi-
tion of certain species of Muscidae (ses. lat.) in the
collection of the British Museum, described by the
late Francis Walker. Annals and Magazine of
Natural History ser. 7, 19: 326-347.
BERGROTH, E. 1894. Ueber einige australische Dip-
226
teren. Stettiner entomologische Zeitung 55: 71-
yey
BRAUER, F. 1899. Beitráge zur Kenntniss der Mus-
caria Schizometopa. I. Bemerkungen zu den
Originalexemplaren der von Bigot, Macquart und
Robineau-Desvoidy beschriebenen Muscaria
Schizometopa aus der Sammlung des Herm G.H.
Verrall. Dritte Folge. Sitzungsberichte der
Akademie der Wissenschaften in Wien 108: 495-
529,
CANTRELL, B.K. 1988. The comparative morphology
of the male and female postabdomen of the
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tions of some first-instar larvae and pupae. Inver-
tebrate Taxonomy 2: 81-221.
CANTRELL, B.K. & CROSSKEY, R.W. 1989.
Tachinidae. Pp. 733- 784. In Evenhuis, N.L. (ed.),
Catalog of the Diptera of the Australasian and
Oceanian Regions. (Bishop Museum Press:
Honolulu and E.J. Brill: Leiden).
CROSSKEY, R.W. 1971. The type-material of
Australasian, Oriental and Ethiopian Tachinidae
(Diptera) described by Macquart and Bigot. Bul-
letin of the British Museum (Natural History).
Entomology 25: 251-305.
1973. A conspectus of the Tachinidae (Diptera) of
Australia, including keys to the supraspecific
taxa and taxonomic and host catalogues. Bulletin
British Museum (Natural History). Entomology
Supplement 21: 1-221.
DONOVAN, E. 1805. An epitome of the natural history
of the insects of New Holland, New Zealand, New
Guinea, Otaheite, and other islands in the Indian,
Southern, and Pacific Oceans’. (Rivington: Lon-
don). 167p.
ENDERLEIN, G. 1937. Dipterologica. IV. Sit-
zungsberichte der Gesellschaft naturforschender
Freunde zu Berlin 1936: 431-443.
ENGEL, E.O. 1925. Uber Rutiliidae sensu lat. (Dipt.).
Zoologische Jahrbucher (Syst.) 50: 339-376.
HARDY, G.H. 1939. Notes on Australian Muscoidea
IV. The genus Microtropeza and some
Phaoniinae. Proceedings of the Royal Society of
Queensland 50: 33-39.
MEMOIRS OF THE QUEENSLAND MUSEUM
MACQUART, J. 1846. Diptéres exotiques nouveaux
ou peu connus. Supplément. Mémoires de la
Société royale des sciences, de l'agriculture et des
arts de Lille 1844: 133-364.
1851. Diptéres exotiques nouveaux ou per connus.
Suite du 4.e supplément publié dans les mémoires
de 1849. Mémoires de la Société royale des scien-
ces, de l'agriculture et des arts de Lille 1850:
134-294,
MALLOCH, J.R. 1928. Notes on Australian Diptera.
17. Proceedings of the Linnean Society of New
South Wales 53: 598-617.
1929. Notes on Australian Diptera. 20. Proceedings
of the Linnean Society of New South Wales 54:
283-343.
1930. Notes on Australian Diptera. 23. Proceedings
of the Linnean Society of New South Wales 55:
92-135.
PARAMONOV, S.J. 1951. Notes on Australian Diptera
(6-8). 8. A review of the genus Microtropeza
Macq. (Tachinidae). Annals and Magazine of
Natural History (12)4: 761-779.
TOWNSEND, C.H.T. 1916. New genera and species of
Australian Muscoidea. Canadian Entomologist
48: 151-160.
1932. Notes on Old-World oestromuscoid types.
Part 2. Annals and Magazine of Natural History
(10)9: 33-57.
1939. Manual of myiology in twelve parts. Part 8.
Oestroid generic diagnoses and data.
Microtropesini to Voriini. 408 Pp. (Itaqua-
quecetuba: Sao Paulo).
WALKER, F. 1849. List of the specimens of dipterous
insects in the collection of the British Museum.
Part 4. Pp. 688-1172. (British Museum: London).
1853. Diptera. Part 4. In 'Insecta saundersiana: or
characters of undescribed insects in the collection
of William Wilson Saunders, Esq., F.R.S., F.L.S.,
&c. Vol. 1’. Pp. 253-414. (Van Voorst: London).
1858. Characters of undescribed Diptera in the col-
lection of W.W. Saunders, Esq., F.R.S., &c
[part]. Transactions ofthe Entomological Society
of London (N.S.) 4: 190-235.
GEOMETRICAL STUDY OF A CAST OF LEPTOPHLOEUM AUSTRALE (McCOY)
WALTON FROM QUEENSLAND
H. TREVOR CLIFFORD
Clifford, H.T. 1996 07 20: A geometrical study of a cast of Leptophloeum australe (McCoy)
Walton from Queensland. Memoirs of the Queensland Museum 39(2): 227-230. Brisbane.
ISSN 0079-8835.
Geometrical study of a cast of Leptophloeum australe (McCoy) Walton confirms that during
its compression there has been no increase in diameter. [ ] Leptophloeum, axis, cast,
compression.
H. Trevor Clifford, Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101,
Australia; 1 February 1996.
Axial casts of the lepidophyte Leptophloeum
australe (McCoy) Walton have been recorded in
Australia from Queensland (Carruthers, 1872)
and Victoria (McCoy, 1874). Herein an addition-
al cast is described from Queensland. The
specimen is of particular interest because over
most of its surface the outlines of leaf-cushion
bases are clearly defined, (Fig. 1) thereby ena-
bling the length of its diameter, prior to compres-
sion, to be determined with reasonable accuracy.
This length therefore provides a bench mark
against which diameters as estimated by other
methods may be evaluated. .
All methods employed assume the cast to have
been cylindrical prior to its compression, a view-
point justified by the uniformity in size and shape
of the leaf-cushions on its surface. Furthermore,
the cast was assumed to be elliptical in transverse
section which assumption has been confirmed by
direct observation. Although casts are often
reported to be elliptical in section (Pant & Srivas-
tava, 1995) there appears to be no previous pub-
lication in which the contention has been
confirmed by a direct comparison of the observed
section shape with that of the ellipse calculated
from the maximum and minimum widths of the
cast.
MATERIAL
The cast (QMF3275) was collected by Leich-
hardt during his expedition from Moreton Bay to
Port Essington and is the only specimen in the
Queensland Museum known to have been col-
lected by the explorer whose misfortune it was to
lose, by misadventure or necessary abandonment,
most of his collections when near to his journey’ s
end (Leichhardt, 1847). The specimen is labelled
‘Clarke River’ and so most probably has its
Provenance in the Clarke River Basin (Draper et
al., 1993). Because the cast was collected as a
surficial boulder its precise age is not deter-
minable. However, the taxon is well represented
by impressions in the Ruxton Formation whose
age, based on conodonts, is Late Devonian to
Early Carboniferous.
Two available casts (Table 1) are entirely
mineral in composition and a thin section of
QMF3204 revealed it to be a fine sediment
dominated by rock fragments and quartz. Other
minerals present are muscovite, orthoclase,
plagioclase, sericite and undifferentiated iron
oxides. The unabraided condition of the grains
and their composition indicates the cast formed
in young sediments with a proximal igneous and
metamorphic provenance. The cement is
dominated by silica and clay with subordinate
iron oxides (Alex Cook per. comm. ).
RESULTS
From measurements of the specimen (Table 1)
it is possible to make several independent esti-
mates of the diameter of the cast before its com-
pression. That the outline of the cast in transverse
section approximates closely to an ellipse may be
confirmed by reference to Fig. 2 where the ellipse
calculated on the basis of the maximum and min-
TABLE 1. Five attributes of two casts of Leptophloeum
australe.
ATTRIBUTE DIMENSIONs (mm)
QMF3275 | QMF3204
Perimeter 283
Maximum width 117
Minimum width
Leaf-cushion width
No. leaf-cushions encircling
228
A
MEMOIRS OF THE QUEENSLAND MUSEUM
B
I S e ag AH A a
f T 1000 i} [T 1200 13500 I Da
FIG.1. Cast of Leptophloeum australe (QMF3275). A, Lateral view. B, End view.
imum widths of the cast is superimposed upon its
original outline.
Four approaches to the estimation of the
diameter of the cast prior to its compression will
now be considered:
1. Perimeter of Cast. If it is assumed that during
compression the perimeter of the cast is un-
changed the diameter of a circle with the same
perimeter is readily calculated (Table 2).
2. Cross Sectional Area of Cast. If the cross
sectional area of the cast is unaffected by com-
pression the areas of the elliptical section of the
cast and that of its precompression circular sec-
tion will be the same. Accepting the maximum
and minimum width of the cast as axis-lengths the
area of the elliptical section may be calculated
and from this the diameter of a circle of similar
area determined (Table 2).
3. Maximum Width of Cast. Assuming that the
vertical compression of a horizontal cylindrical
cast may occur without any lateral expansion the
maximum width of the cast is the same as the
diameter of the uncompressed cast (Table 2).
4. Leaf-cushion number x Leaf-cushion width.
TABLE 2. Four estimates of the diameter of a
presumed cylindrical cast of Leptophloeum australe
as determined from certain attributes of its com-
pressed cast (QMF3275).
ESTIMATED
BASIS OF ESTIMATE DAT
(mm)
]. Perimeter 90
2. Cross sectional area TI
3. Maximum width 117
4. Cushion number x cushion width 107
The number of leaf-cushions encircling the cast
is half the number of the vertical rows in which
they are arranged (Fig. 3). The recognition of
these rows is more reliable than that of adjacent
leaf-cushion boundaries wherever the surface of
the cast is irregular or the leaf-cushions are
strongly compressed as in regions of maximum
curvature. Therefore, the number of leaf-
cushions encircling the cast was determined as
half the number of vertical rows of leaf-cushions
on the surface of the cast. Multiplication of leaf-
cushion number by leaf-cushion width provided
a perimeter for the uncompressed cast from
which its diameter was calculated (Table 2).
DISCUSSION OF RESULTS
Each of the 4 methods employed provided a
different estimate of the diameter of the original
cast (Table 2). These differences are readily ac-
counted for if it is assumed that during compres-
sion a cylindrical cast was deformed into one
elliptical in transverse section, with the major
axis of the ellipse being of the same length as the
diameter of the cylinder.
Such a situation is illustrated in Fig. 4 where
sections of a theoretical cast, prior to and sub-
sequent to its compression, are superimposed.
The ellipse is similar to that of Fig. 2 because the
lengths of the major and minor axes of the two
are the same. Along the perimeter of both the
circle and ellipse solid circles mark the positions
of leaf-cushion margins. The number of leaf-
cushions is the same as that on the cast being
studied and their positions on the ellipse are ver-
tical projections of those on the circle. Therefore
GEOMETRY OF CRUSHING LEPTOPHLOEUM AUSTRALE
229
FIG. 2. Outline of cast of Leptophloeum australe (QMF3275) with the perimeter of theoretical ellipse
superimposed. A-D are places on cast surface at which leaf-cushion widths (Table 3) were measured.
the ellipse may be regarded as a compression of
the circle without a change of its diameter.
Both the perimeter and the area of the ellipse
will provide measurements which will lead to
underestimates of the diameter of the cir-
cumscribing circle, and the shorter the minor axis
of the ellipse the poorer will be the estimate.
Furthermore, of the two parameters under dis-
cussion the perimeter of the ellipse will always
FIG. 3. Semidiagrammatic drawing of the disposition
of leaf-cushion outlines on the surface of cast of
Leptophloeum australe (QMF3275). A= direction of
stem apex; CB=leaf-cushion base; CM=leaf-cushion
margin; CW=leaf-cushion width; VC=vascular
cicatrix..
provide a closer estimate of the perimeter of the
inscribing circle than will the area of the ellipse.
This statement is especially true of strongly com-
pressed casts. Towards the limit of compression,
as the length of its minor axis approaches zero,
the perimeter of an ellipse approaches a value of
twice the length of its major axis but the area of
the ellipse approaches zero. Hence it is clear that
ellipses whose minor axes are short in com-
-———«- leaf-cushion
FIG. 4. Transverse sections of 2 theoretical casts on
whose perimeters the margins of the leaf-cushion
outlines have been marked by solid circles — circle,
before compression; ellipse, after compression. The
major and minor axes of the ellipse are the same
length as in Fig. 2.
230
TABLE 3. The areas and perimeters of an ellipse with
different minor and major axis ratios expressed as
proportions of its circumscribing circle.
Ellipse as proportion if its
circumscribing circle
Length of minor axis
Length of major axis
parison to their major axes provide poor informa-
tion for predicting the diameter of the cir-
cumscribing circle.
However, the situation is different if the length
of the minor axis of the ellipse equals or exceeds
half the length of the major axis. Over this range
of values both the area (exactly) and the perimeter
(closely) of the ellipse are linearly related to the
area of the circumscribing circle. Whereas for
high values of the ratio of the lengths of the minor
and major axes, both the perimeter and the area
of the ellipse are useful predictors of the diameter
of the circumscribing circle, for low values of the
ratios neither is useful but the perimeter is the
better estimator (Table 3). This observation is
confirmed by the estimates of diameter (Table 2).
It is also clear (Fig. 4) that whereas the leaf-
cushion bases around the perimeter of the circle
are all of the same width their projections on to
the ellipse vary in width. Whereas leaf- cushion
B (Fig. 4) is almost the same width as that of A,
the width of leaf-cushion D is much less than that
of C. If the leaf-cushion is bisected by the minor
axis of the ellipse its width will be almost exactly
that of its width before projection from the circle.
Therefore, provided the leaf-cushions measured
are situated close to where the minor axis of the
ellipse meets its surface their widths combined
with their number provide an accurate basis for
estimating the diameter of the uncompressed cast.
Because only one cast is available it is not
possible to determine whether the diameters of
the uncompressed cast as estimated from the
maximum width of the compressed cast (117mm)
and by the leaf-cushion number x width method
(107mm), are statistically different. However, the
data indicate that if there has been any lateral
spread of the cast during compression the exten-
sion has been slight for the ratio of the former to
the latter is 1.09 which value is close to unity.
Further support for the view that the width of
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 4. The widths of leaf-cushion at 4 positions
on the surface of a cast of Leptophloeum australe
(QMF3275) as measured directly and as determined
by projection from a circular cast onto one elliptical
intransverse section with a major axis the same length
as the diameter of the circular cast.
Leaf-cushion width (mm)
rr
4.0503
2-04 | 8o |
11.9+0.2
13.408 —
the cast has not increased as a result of compres-
sion comes from a comparison of the measured
and predicted widths of leaf-cushions. Measure-
ments were made on four areas of the cast surface
and the predicted widths were taken from similar
positions on the ellipse (Fig. 2; Table 4). Standard
errors can be attached to the means of the meas-
urements because there are several leaf-cushions
available on similar areas of the cast but there is
only one predicted value for corresponding parts
of the ellipse. Only on the surface of greatest
curvature does the predicted value of leaf-
cushion width differ significantly from that
measured. Such close agreement between the two
sets of values is further support for the hypothesis
that compression of the cast has occurred without
any lateral extension.
Although casts of Leptophloeum australe are
rare, impressions of their axes are abundant. Most
are flat and irregular in outline, but a few are
parallel-sided thereby resembling impressions of
complete axes. However, it is not reasonable to
assume that the impression revealed on a flat
surface derives from a whole axis. It may repre-
sent any portion of the surface, the remainder of
which may be buried in the rock on either side of
the plane (along which the rock split) to reveal the
impression. Cleavage across a cast near to its
surface and parallel to its length would expose
impressions with parallel sides but whose widths
are much less than the diameter of the cast. That
this situation is common is suggested by the many
impressions whose leaf-cushion bases have
widths typical of casts whose diameters are
greater than the width of the impression.
CONCLUSION
Although it has been widely accepted that com-
pression of plant tissues and casts usually occurs,
without a concomitant increase in their width at
GEOMETRY OF CRUSHING LEPTOPHLOEUM AUSTRALE
right angles to the force applied, the process has
been subject to little theoretical or experimental
study.
Using a series of projections similar to that
employed above, Walton (1936) compared the
shapes of some solids pre- and post- compression.
His approach was qualitative and non-ex-
perimental. However, according to Harris (1974:
144) Walton’s observations were underpinned by
a series of experiments in which he had com-
pressed ‘various solid plant organs - plant stems,
apples and the like in wet sand in a power press
so constructed as to allow surplus water to drain
away’. Apples so compressed were converted
into hemispheres filled with sand but with their
diameters unchanged.
The pioneering study of Walton (1936) has
been extended by Harris (1974), Niklas (1978)
and Rex & Chaloner (1983). Harris (1974) em-
bedded hollow balls of wax or plastic, in a variety
of matrices which were then compressed. After
compression the balls were approximately hemis-
pherical and circular in outline. None of the com-
pressions showed any obvious evidence of
horizontal extension. Simulated compression
studies of Niklas (1978) were quite extensive but,
unfortunately, the procedures adopted were not
adequately described. However, he demonstrated
that whereas hollow and solid, but dehydrated
stems compress without lateral extension
hydrated solid stems ‘show a maximum increase
of 1096 diameter'. Experimental procedures
adopted by Rex & Chaloner (1983) are fully
explained and so their results can be critically
assessed. Foam rubber was used to represent
plant material and saw dust the embedding
matrix. Pressure was applied either with a single
piston or a parallel system of independent pistons
each of which was spring loaded, but with the
whole set controlled by a single screw mechan-
ism. After compression, transverse sections of the
original cylinders (stems), differed in shape ac-
cording to the piston system employed. Nonethe-
less in both systems and for several cylinder
diameters the maximum width of the stem fol-
lowing compression was equal to or slightly less
than the original diameter. That is, the results are
in accord with the predictions of Walton (1936).
It is not possible to compare with certainty the
results obtained from the several experimental
studies because of the diversity of materials
employed and the differences in procedures
adopted. Furthermore, these results are not direct-
231
ly comparable with the observational data
reported above for none of the experimenters
included casts in their studies. Nonetheless, the
application of pressure to embedded material did
not result in its lateral extension except for one
report by Niklas (1978) . The similarity ofthe pre-
and post-compressional diameters of the Leich-
hardt cast (QMF3275) suggest that even when
subjected to pressures much greater than those
employed in the laboratory plant axes do not
expand lateral to the force applied.
ACKNOWLEDGEMENTS
Thanks are extended to Mary Wade, Alex Cook
and Vincent Hart for helpful discussions during
the preparation ofthis paper and to Natalie Camil-
leri and Maryanne Venables for assistance in
preparing the diagrams. The photographs were
taken by Gary Cranitch.
LITERATURE CITED
CARRUTHERS, W. 1872. Notes on fossil plants from
Queensland. Quarterly Journal of the Geological
Society of London 28: 2-9
DRAPER, J.J., SCOTT, M. & WITHNALL I.W. 1993.
Clarke River Basin and associated rocks.
Queensland Geology 4: 203-219.
HARRIS, T.M. 1974. Williamsoniella lignieri: its pol-
len and the compression of spherical pollen grains.
Palaeontology 17:125-149
LEICHHARDT, L. 1847. ‘Journal of an overland ex-
pedition in Australia from Moreton Bay to Port
Essington during the years 1844-1845’, (T. & W.
Boone: London). 544p.
McCOY, F. 1874. Lepidodendron (Bergenia) australe
(McCoy). Pp. 37-39. In 'Prodromus of the
palaeontology of Victoria. Decade 1’. (Geological
Survey of Victoria: Melbourne).
NIKLAS, K.J. 1978. Morphometric relationships and
rates of evolution among Palaeozoic vascular
plants. Evolutionary Biology 11: 509-543.
PANT, D.D. & SRIVASTAVA, P.C. 1995. Lower Car-
boniferous plants from Wallaranna Series of Pun-
jab-Kashmir Himalaya. Palaeontographica B,
235: 23-49.
REX, G.M. & CHALONER, W.G. 1983. The ex-
perimental formation of plant compression fossils.
Palaeontology 26: 231-252.
WALTON, J. 1936. On the factors which influence the
external form of fossil plants: with descriptions of
the foliage of some species of Palaeozoic equi-
setalean genus Annularia Sternberg. Philosophi-
cal Transactions of the Royal Society of London
226B: 219-237.
232
POSSIBLE AFFINITIES BETWEEN VARANUS
GIGANTEUS AND MEGALANIA PRISCA. Memoirs of the
Queensland Museum 39(2):232. 1996:- Molnar (1990)
described two frontals, and a parietal, of a giant Pleistocene
varanid at King Creek, eastern Darling Downs. The material
was assigned to Megalania prisca, the only varanid of com-
parable size. This identification is probably correct since
undoubted remains of Megalania occur in the same deposits.
Molnar (1990) noted that the frontals and parietal of the
King Creek varanid exhibited many unusual features, which
could not be found in any varanid skulls examined, and were
thus presumably derived within varanids. Among these fea-
tures were the prominent sagittal crest along the median suture
between the frontals, and the parallel transverse ridges extend-
ing at right angles to this crest. Both these features are also
found in Varanus giganteus (Fig. 1) and are absent in other
species of Varanus (Molnar, 1990) and in the nearest outgroup
taxa, Lanthanotus and Heloderma (Rieppel, 1980; Pregill et
al., 1986; Estes et al., 1988). They are thus derived within
Varanus suggesting affinities between the King Creek varanid
and V. giganteus. Molnar (1990) noted that, in the King Creek
varanid as in V. giganteus (Fig.1), the sagittal crest and
parallel transverse ridges were confined to the frontals, and
did not extend onto the parietals. This phylogeny is based on
very incomplete material and only two characters.
Megalania prisca, Varanus giganteus, V. salvadori and V.
komodoensis are the 4 largest known varanids (Pianka, 1995).
Despite the latter two not being Australian natives, all 4 belong
to a discrete radiation of Australian monitors, the ‘gouldii
species group' (Baverstock et al.,1993). If. Megalania prisca
has affinities with V. giganteus and thus belongs within the
gouldii species group Megalania will have to be synonymised
with Varanus. Relationships within the gouldii species group
are not yet well established (Baverstock et al. 1993).: there isa
distinct possibility that, when relationships within this radiation
are resolved, V. giganteus, V. salvadori, V. komodoensis, and
Megalania prisca will form a clade. If so, this would mean that
the four largest varanid species represent a single discrete
radiation of giant predatory lizards.
Ithank the Australian Research Council for funding, Jenny
Clack and Ray Symonds for loan of the figured specimen, and
Malcom Ricketts for photography.
Literature Cited
BAVERSTOCK, P.R., KING, D., KING, M., BIRRELL, J.
& KRIEG, M. 1993. The evolution of the species of the
Varanidae: Microcomplement fixation analysis of
serum albumins. Australian Journal of Zoology 41:
621-638.
MEMOIRS OF THE QUEENSLAND MUSEUM
ESTES, R., DE QUEIROZ, K. & GAUTHIER, J. 1988.
Phylogenetic relationships within Squamata. Pp. 119-
281 In Estes, R. & Pregill, G. K. (eds). ‘Phylogenetic
relationships of the lizard families'. (Stanford Univer-
sity Press: Stanford).
MOLNAR, R.E. 1990. New cranial elements of a giant
varanid from Queensland. Memoirs of the Queensland
Museum 29: 437-444.
PIANKA, E.R. 1995. Evolution of body size: varanid lizards
as a model system. American Naturalist 146: 398-414.
PREGILL, G., GAUTHER, J. & GREENE, H.W. 1986. The
evolutlon of helodermatid squamates, with description
of a new taxon and an overview of Varanoidea. Trans-
actions of the San Diego Society of Natural History 21:
167-202.
RIEPPEL, O. 1980. The phylogeny of anguinomorph lizards.
Denkschrifte Schweizerbartsche Naturf. Gesellshaft
94: 1-86.
M.S.Y. Lee, School of Biological Sciences, University of Syd-
ney, NSW 2006, Australia; 10 December 1995.
FIG. 1. Skull of Varanus giganteus (University Museum of Zoology, Cambridge R9586) in (A) dorsal, and (B) right laterodorsal
view, showing the sagittal crest and dermal sculpture on the frontals between the orbits. Scale bar = 3cm.
COGGERIA NAUFRAGUS GEN. ET SP. NOV., A SAND-SWIMMING SKINK FROM
FRASER ISLAND, QUEENSLAND
P.J. COUPER, J.A. COVACEVICH, S.P MARSTERSON AND G.M. SHEA
Couper, P.J., Covacevich, J.A., Marsterson, S.P. & Shea, G.M. 1996 07 20: Coggeria
naufragus gen. et sp. nov., a sand- swimming skink from Fraser Island, Queensland. Memoirs
of the Queensland Museum 39(2):233-241. Brisbane. ISSN 0079-8835.
Coggeria naufragus gen. et sp. nov. is a distinct lygosomine in the Sphenomorphus group.
It has a digital formula of 3/3. A sharp snout and more than 40 lingually-directed maxillary
teeth are key distinguishing characters. The new skink is a sand-swimmer, which feeds on
worms. It is known only from Fraser Island, SEQ, a World Heritage Site. [ ]$5phenomorphus
group, Scincidae, rainforest species, World Heritage Site, Fraser Is.
P.J. Couper & J.A. Covacevich, Queensland Museum , .P.O Box 3300, South Brisbane,
Queensland 4101, Australia; S.P. Marsterson, Queensland Department of Environment &
Heritage, P.O. Box 21, Yungaburra, Queensland 4872, Australia; G.M. Shea, Department
of Veterinary Anatomy, University of Sydney, New South Wales 2006, Australia; 1 December
1995.
On Fraser Island in June, 1991, Mike West
found a small sand-swimming skink while dig-
ging. It was sent to the Queensland Museum for
identification. At first glance, the damaged
specimen, resembled Saiphos equalis (Gray,
1825). Several unsuccessful attempts were made
to find more specimens. During summer 1994/95,
a party from the Queensland Museum (PJC and
JAC) and the Queensland Department of En-
vironment and Heritage (SM, Rod Hobson, Keith
Twyford, other staff and volunteers) collected
several more specimens.
Like other World Heritage Sites in Queensland,
Fraser Is. has been the focus of considerable
research and management effort. The island's
diverse reptile fauna (Barry & Campbell, 1977;
Covacevich & Couper, 1991) in many habitats,
including rainforest, was thought to be well
known. That a secretive, very distinct skink from
Fraser Is. rainforest should be discovered in 1991
indicates that there are still elements of the
Queensland reptile fauna about which we know
little.
The skink is assigned to the Sphenomorphus
group within the Lygosominae. The Sphenomor-
phus group has a single frontal bone; palatine
bones in contact on the ventral midline;
ventrolateral ridges of the frontal each with a
short process, and frontal separated from the pala-
tine by an extensive section of prefrontal; nine
premaxillary teeth; an open Meckel’s groove; iris
virtually as dark as pupil; parietal scales in con-
tact behind the interparietal; posteriorlateral edge
of each parietal bordered by two temporals and a
nuchal, and a greatly enlarged medial pair of
preanal scales (Greer, 1970, 1979, 1986a).
MATERIALS AND METHODS
All measurements were taken using Mitutoyo
electronic callipers. Supraciliaries, supralabials,
infralabials, and subdigital lamellae on the hind
toes were counted on both sides. The following
meristic characters have been used:- snout-vent
length (SVL); axilla to groin (AG); tail length,
vent to tip (TL); forelimb, axilla to tip of longest
digit (L1); hindlimb, groin to tip of longest digit
(L2); forelimb to snout, from axilla to tip of snout
(L1-S); head length, tip of snout to posterior
margin of parietals (HL); head width, measured
level with the posterior margin of the parietals
(HW); head depth, measured level with the
posterior margin of the parietals (HD); snout, tip
to anterior margin of orbit (S); eye to ear-crease,
posterior margin of orbit to mid lateral margin of
ear-crease (EE). Osteological characters are
based largely on a single, cleared and stained
specimen, QMJ59670, and supplemented by data
(vertebral counts) from X-rays of QMJ57431,
59237, 59312, 59361, 59468-69 and 59671.
SYSTEMATICS
Coggeria gen. nov.
TYPE SPECIES. Coggeria naufragus sp. nov.
ETYMOLOGY. For Harold Cogger, former Curator of
Reptiles and Deputy Director of the Australian
Museum, for his many contributions to knowledge and
conservation.
DIAGNOSIS. Elongate body (Fig. 1), reduced
limbs (front and rear limb 4.396 and 7.796 of SVL,
234
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. Coggeria naufragus gen. et sp. nov.
respectively), snout wedge-shaped in profile
(Figs 2,3); nasals slightly enlarged; prefrontals
separated; supraoculars 3, first only in contact
with frontal; last 2 supraoculars partially
separated by a supraciliary; supraciliaries 5, first
contacting frontal; supralabials usually 6, fourth
below eye; postsupralabial single; ear opening
absent; upper secondary temporal overlapped by
lower. Osteology: maxilla-frontal contact; pre-
and postfrontals in contact above orbit; postorbi-
tal absent; distinct narrowing of skull at premaxil-
lary-maxillary junction; maxillary teeth greater
than 40, with long axis of tooth running trans-
versely and crowns directed lingually; dentary
teeth more than 45; pterygoid teeth absent;
presacral vertebrae 47-50; manus lacking inter-
medium, distal carpals 1 and 5 and metacarpals 1
and 5, and has phalanges reduced to 0.2.3.3.0; pes
with astragalus and calcaneum fused, lacking dis-
tal tarsals 1 and 5, metatarsals 1 and 5, and with
phalanges reduced to 0.2.3.3.0 (Fig. 4); sternal
ribs 2; mesosternal ribs 1; ischia forming acute
angle at symphysis with shafts paralleling those
of pubes. Parietal peritoneum lacking pigment.
Other elongate genera of the Sphenomorphus
group (Anomalopus, Calyptotis, Coeranoscin-
cus, Ophioscincus, Saiphos, Lerista) share some
of the apomorphies of Coggeria in varying com-
binations (Table 1). However, many of these
apomorphies are associated with burrowing, and
may point to parallel evolution rather than close
relationships (Greer & Cogger, 1985). Anatomi-
cal variation and phylogenetic relationships in the
Sphenomorphus group, particularly in the non-
Australian members, remain poorly known, and
a well-corroborated cladistic phylogeny is not
available.
Coggeria shares many apomorphies with
Coeranoscincus (15; 18) and Ophioscincus (17),
which are closely associated geographically.
However, Coeranoscincus differs from Coggeria
in having: teeth fang-like, posteriorly curved and
sharply pointed; snout conical; ischial shaft
weakly developed or absent. Ophioscincus dif-
fers from Coggeria in having: supraciliaries 3-4;
supralabials 5; limbs 2% of SVL or shorter;
phalanges absent on both manus and pes. Tooth
shape and a high number of maxillary teeth of
Coggeria set it apart from Anomalopus, Calyp-
totis, Coeranoscincus, Ophioscincus, Saiphos
and Lerista, all of which have fewer than 26
maxillary teeth, with a generally upright or
posteriorly- curved orientation (Cogger, 1992;
Greer,1983, 1986b, 1989; Greer & Cogger, 1985;
Storr, 1971).
Coggeria naufragus sp. nov.
(Figs. 1-5)
MATERIAL EXAMINED. HOLOTYPE QMJ59361,
E of Central Stn workshop (25728'42"' S, 153°03’21"E)
SEQ. PARATY PES QMJ5743 1, between Leading Hill
& Lake Garrawongera, behind Poyungan Valley
(25?23'S, 153°05’E); QMJ59237, N of Central Stn
workshop (25?28'37"S, 153?03'15"E); QMJ59312,
COGGERIA, A NEW SKINK FROM FRASER ISLAND
235
FIG. 2. Coggeria naufragus gen. et sp. nov. showing wedge-shaped snout.
QMJ59468, W of Central Stn workshop (25?28'38"S,
153?03'15"E); QMJ59469, Ikm S of Pile Valley,
1.5km E of Central Stn (25?28'56"S, 153°04’20"E);
QMJ59670, QMJ59671 Central Stn, E of QDEH
workshop (25?28'23"S, 153?03'23"E). OTHER
MATERIAL: QMJ60232 (tail only), N of Central Stn
workshop (25?28'37"S, 153?03' 15"E). All from Fraser
Island.
ETYMOLOGY. Latin naufragus, castaway,
shipwrecked.
DIAGNOSIS. As for genus.
DESCRIPTION. SVL(mm)=62-127 (n=7, mean
98.2). Proportions as % SVL: AG=73.4-76.8
(n=7, mean 75.2); TL=76.5—-83.1 (n=3, mean
79.2); L123.1-4.3 (n=7, mean 3.8); L226.2-7.7
(n=7, mean 6.7); L1-S=20.3-22.7 (n=6, mean
21.4); HL27.5—9.8 (n=7, mean 8.2); HWz5.3-
6.0 (n=6, mean 5.6); HD=4.1-4.5 (n=5, mean
SUPRACILIARIES
PRETEMPORALS
POST-
INFRALABIALS
POSTOCULARS
4.4); S=3.3-4.0 (n=7, mean 3.5); EE-4.4—5.6
(n=7, mean 4.9).
Head indistinct from neck; snout wedge-
shaped in profile. Nasals large, moderately
spaced. Nostril positioned anteroventrally in
nasal. Prefrontals large, moderately spaced. Fron-
tal 1.2 times as long as wide; contacting prefron-
tals, frontonasal, frontoparietals, first
supraoculars and first supraciliaries. Frontoparie-
tals paired, in broad contact. Interparietal free,
parietal eye present. Parietal scales in broad con-
tact behind the interparietal. Enlarged nuchal
scales 4-5 pairs. Two nuchals in direct contact
with posterior edge of parietal scales. Loreals 2,
first larger. Supralabial scales 6-7 (n=16, mean
6.1); where 6, fourth below eye; where 7, fifth
below eye. Postsupralabial single. Infralabials 6.
Postmental contacts 2 infralabials on each side.
Three pairs of enlarged chin scales; first pair in
contact, second pair separated by 1 longitudinal
ventral scale row, third pair separated by 3 lon-
PRIMARY TEMPORAL
2ND TEMPORALS
|
SUPRAOCULARS PRETEMPORALS
FIG. 3. Head scalation of the holotype (QM J59361) of Coggeria naufragus gen. et sp. nov.
236
M
D
$ i
FIG. 4. Number and configuration of the bones in the
manus (A) and pes (B) of C. naufragus.
gitudinal ventral scale rows (terminology follows
Greer, 1989:152). Lower eyelid movable and
scaly. Preoculars 2. Presuboculars 1. Suboculars
2. Supraoculars 3. Supraciliaries 5. Postoculars 2.
Pretemporals 2. Primary temporals 1. Secondary
temporals 2, upper the largest and overlapped by
lower. External ear opening completely covered
Jy.
FIG. 5. Ectopterygoid, pterygoid and palatine area in
C. naufragus.
MEMOIRS OF THE QUEENSLAND MUSEUM
by scaly epidermis, its former position indicated
by an elongate, vertically-oriented, shallow
depression.
Body elongate, with smooth scalation. Mid-
body scale rows, 22-24 (n=8, mean 22.8).
Paravertebral scales, from anteriormost nuchal to
posterior margin of hindlimb 88-100 (n=7, mean
92.7); slightly enlarged. Number of scales in a
direct line between mental and preanal scales
100-111 (n=7, mean 105.9). Medial pair of
preanal scales enlarged, overlapping outer
preanals. Limbs short, tridacty]. Subdigital lamel-
lae on hindlimb - first toe 2-3 (n= 16, mean 2.3),
second toe 3-5 (n=16, mean 3.3), third toe 3-4
(n=16, mean 3.6). Original tail tapered distally,
terminating sharply.
Skeletal features. Premaxillary teeth 9 (n=1).
Maxillary teeth 43/43 (n=1). Dentary teeth 48/49
(n=1). Frontal single. Vomers fused. Palatines in
contact along ventral midline. Palatal rami of
pterygoids with slight recurved processes. Ectop-
terygoid process absent (Fig. 5). Parietal foramen
present. Postorbital bone absent. Supraorbital
fenestra small, almost obliterated by close ap-
position of supratemporal arch to parietal. The
hyoid apparatus is illustrated in Fig. 6.
Presacral vertebrae 47-50 (n=8). Postsacral
vertebrae 45 (n=1). Complete inscriptional chev-
rons 11 (n=1). Sternal/mesosternal ribs 2/1 (n=1).
Manus comprising radiale, ulnare and pisiform;
centrale; distal carpals 2-4; metacarpals 2-4, and
phalanges 0.2.3.3.0. Pes comprising fused
astragalus and calcaneum; distal tarsals 2—4;
metatarsals 2-4; phalanges 0.2.3.3.0. (Fig. 4).
cb1
FIG. 6. Hyoid apparatus of C. naufragus. bh = basihyal,
cbl = first ceratobranchial, ch = ceratohyal, gh =
glossohyal process, hh = hypohyal, tr = tracheal car-
tilages.
COGGERIA, A NEW SKINK FROM FRASER ISLAND 237
TABLE 1. Character states (derived +, plesiomorphic -) in reduced-limbed genera of the Sphenomorphus group.
Data for genera other than Coggeria are from text and illustrations of Cogger (1992), Greer (1983, 1986b,
GENERA
CHARACTER: DERIVED
STATE
1989), Greer & Cogger (1985) and Storr (1971). For other material examined see Appendix 1.
snout profile sharp * -
nasal slightly enlarged + +
loreal single - +/-
prefrontals absent - +/-
supraoculars 3 or less + +/-
first supraciliary contacting frontal
| last 2 supraoculars partially
separated by a supracili
supraciliaries <7
supralabials <7
supralabial below eye, not 5th
postsupralabial single
secondary temporals: lower
overlaps upper
nasal-prefrontal bone contact - +/-
pre & postfrontals .
approach/contact above orbit t +
postorbital absent + +
- +
premaxillary teeth <9 +(<7)
:
Coggeria |Anomalopus e dni E Seis Saiphos Calyptotis | Lerista
- id , +/-
+ + - +
+/- oe
- H- 4
+ + -
+ + +
+- - -
+(4-6) +(3-4) +/-(5-7)
H6 +(5) +(6)
+(4 +(3) +(4
> + -
+/- - -
+ + + +/- -
+(<2) +(S11) | +/-(<25) | +/-(<21) |
(a)
+(S2) +(<14) +/-(<33) | +/-(<33)
+(18-24) | +(18-22) | +(18-24) | +(16-24)
n L t 3: . (5
+ + + š +- ©
E
E +/-
+(8) +/-(<9)
-(<26 -(<21)
-(<26) -(<22)
+(252) +(243)
sternal ribs <3
mesosternal ribs <2
phalanges (manus), not 23453
phalanges (pes), not 23454
| parietal peritoneum unpigmented ii
ischial & pubic shafts parallel "
+
+(22) +(S2)
+1433) ©
4-2) ©
"m
+/- (c)
+ 00000
+ 00000
+ 02330 4 3 |-/- 23453
+ 02330|+ 23453 |+/- 23454
23443 02454
23444 02350
238
Measurements and scale counts for holotype
(QMJ59361). SVL=106.5mm; AG=78.4mm; TL
=88.5mm; L1=4.6mm; L2=6.8mm; L1-S=
21.6mm; HL=8.4mm; HW=5.7mm; HD=
4.8mm; S=3.5mm; EE=5.6mm. Maximum
length of frontal 2.3mm; maximum width of fron-
tal 2.0mm; four pairs of enlarged nuchal scales;
supralabial scales 6; Midbody scale rows 22;
paravertebral scales 88; number of scales in direct
line between the mental and anal shields 100;
subdigital lamellae on hindlimb (both sides of
body), first toe 2, second toe 3, third toe 3.
Pattern. Dorsum light tan. Some scales with
dark brown spots, forming broken, longitudinal
lines on body and tail; head with dark brown
mottlings. Ventral and lateral surfaces greyish,
heavily marked with black flecks; sharply demar-
cated from dorsum by a dorsolateral row of black
spots (1/scale), beginning behind the eye and
running the full length of tail. Dorsal scales im-
mediately adjacent to the black dorsolateral zone
paler than those of the rest of the dorsum, in
juveniles forming a stripe bordering the black
dorsolateral zone; stripe less clearly defined in
larger animals. Limbs grey, heavily mottled with
black.
COMPARISON. C. naufragus can be confused
with only Australian lygosomines with a digital
formula of 3/3 (Coeranoscincus reticulatus,
Hemiergis decresiensis, southwestern popula-
tions of H. peronii, some species of Lerista and
Saiphos equalis). C. naufragus is separated from
C. reticulatus by snout shape in profile (sharp vs
bluntly conical) and number of maxillary teeth
(240 vs «26); from H. decresiensis, south-
western populations of H. peronii and Lerista
spp. by lower eyelid (scaly vs clear window).
From Lerista, it is further distinguished in lacking
an external ear opening; from Saiphos, in having
prefrontals and 2 loreals (vs 1).
HABITAT. Fossorial species associated with tall
forest communities on sandy substrates. The type
series was collected from three main vegetation
communities: closed Syncarpia hillii, Satinay
forest (Fig. 7A); mixed S. hillii and Eucalyptus
pilularis, Blackbutt forest (Fig. 7B), and open E.
pilularis forest (Fig. 7C). The S. hillii and mixed
S. hillii / E. pilularis forests contain a Backhousia
myrtifolia (Carrol) understorey and Macrozamia
miquelii (Wild Pineapple) ground cover. The
open E. pilularis forest has a Monotoca scoparia
(Prickly Broom Heath) understorey. These
forests have a long history of harvest, and mod-
MEMOIRS OF THE QUEENSLAND MUSEUM
ification by fire. They occur as a broad, broken,
central strip on Fraser Is., between latitudes
25°08’—25°44’S, and are illustrated as vegetation
types one to four on the Fraser Is. vegetation maps
(Department of Forestry 1979, 1985). C.
naufragus has been collected only between
25?20'-25?30'S. The association between C.
naufragus and tall forest communities may
reflect collecting effort, or population variations
between different habitats. For many years,
Anomalopus pluto, another fossorial skink, was
believed to be associated with monsoon forests
because this was present at its type locality. Sub-
sequent collections have shown A. pluto to occur
in a wide variety of vegetation types. Substrate,
rather than vegetation type, appears to determine
the distribution of A. pluto (Couper, 1992).
HABITS. C. naufragus is difficult to find during
the cooler months and in dry periods. Most in-
dividuals (preserved and released) were caught
during an extensive pit-trapping programme be-
tween | December 1994 and 15 March 1995.
Surface activity occurs during the summer
months in periods of high humidity, or immedi-
ately following rain. Most pit-trapped specimens
were captured at night. Two specimens have been
recorded while digging. The first of these,
QMJ57431, was found during winter (13-15 June
1991). The second, a tail only (QMJ60232), was
found 20cm below the surface, during an exces-
sively dry period (17 Nov. 1994).
C. naufragus is an adept ‘sand-swimmer’.
Specimens held briefly in captivity for photog-
raphy, in sand-filled ice-cream containers,
‘disappeared’ rapidly, evading capture repeated-
ly and easily. We surmise that this species is
largely subterranean, and that it burrows deeply
during cold and dry times.
No data are available on its breeding habits.
However, a specimen (TL 80mm, SVL ap-
proximately 45mm) is considerably smaller than
any specimens in the type series and was con-
sidered a juvenile. It was collected 15 August
1995, at Pile Valley, 2km E of the type locality
and released.
DIET. Faecal and gut samples from QMJ59670,
and a faecal sample from QMJ59671 show that
C. naufragus is a worm feeder. Both faecal
samples contained humus-rich soil (presumed to
be worm gut contents), with a few sand grains and
numerous oligochaete setae (L. Cannon pers.
comm.). The gut sample contained similar
239
COGGERIA, A NEW SKINK FROM FRASER ISLAND
Syncarpia hillii forest.
3
2.5
$5
s
PU
Ta
uc
23
22
M
an
ri
x25
E E
Ù
5
st.
S. hillii and Eucalyptus pilularis fore:
mixed
FIG. 7. Vegetation communities with which C. nau,
B,
240
material, with what appeared to be worm dermal
tissue.
ACKNOWLEDGEMENTS
We thank Mike West (Fraser Island Safaris) for
collecting the first specimen of C. naufragus, and
bringing it to our attention; Keith Twyford &
David Redman (Queensland Department of En-
vironment and Heritage) for supporting our
fieldwork and implementing a trapping
programme to obtain additional material; the
Kingfisher Bay Resort for providing support
during the initial search for specimens; Luke Bar-
rowcliffe, Ben Garrett, Rod Hobson, Dan
Haipola, Josh Hastings, Celia Jobson, Moyra
McRae, Anika Tauchmann, Noel Wedding, Lyn
Willsher (Queensland Department of Environ-
ment and Heritage staff and volunteers) for
patrolling pit traps; Jeff Wright for collecting a
road-killed specimen, and for preparing our
photographic plates; Mrs K. Reynolds (Univer-
sity of Queensland) for X-raying specimens; Ms
H. Janetzki for assisting with the final proof-read-
ing of the manuscript; Ms D. Case for preparing
Table 1; Steve Wilson for his valuable advice and
Fig. 2 and Lauren Keim for her assistance in the
laboratory.
LITERATURE CITED
BARRY, D.H. & CAMPBELL, P.R. 1977. A survey of
the mammals and herptiles of Fraser Island, with
comments on the Cooloola Peninsula, North
Stradbroke, Moreton and Bribie Islands. Oc-
casional Papers in Anthropology 8: 147-178.
(Anthropology Museum: University of
Queensland).
GOGGER, H.G. 1992. Reptiles and amphibians of
Australia. 5th ed. (Reed:Sydney). 775pp.
COUPER, P.J. 1992. Anomalopus pluto Ingram, a poor-
ly known skink from Cape York Peninsula.
Memoirs of the Queensland Museum 32(1): 54.
COVACEVICH, J.A. & COUPER, P.J. 1991. The rep-
tile records, pp 45-140. In Ingram, G.J. and Raven,
R.J. (eds). An atlas of Queensland's frogs, reptiles,
birds and mammals. (Board of Trustees,
Queensland Museum: Brisbane). 391 pp.
COVACEVICH, J.A. & MCDONALD, K.R., 1993.
Distribution and conservation of frogs and reptiles
of Queensland rainforests. Memoirs of the
Queensland Museum 34(1): 189-199.
DEPARTMENT OF FORESTRY AND
QUEENSLAND NATIONAL PARKS AND
WILDLIFE SERVICE, 1979. Fraser Island,
central sheet 1:50,000, edition 1 (Government
Printing Office: Brisbane).
MEMOIRS OF THE QUEENSLAND MUSEUM
1985. Fraser Island, south sheet 1:50,000, edition 2
(Government Printing Office: Brisbane).
GREER, A.E. 1970. A subfamilial classification of
scincid lizards. Bulletin of the Museum of Com-
parative Zoology. 139(3): 151-184.
1974. The generic relationships of the scincid genus
Leiolopisma and its relatives. Australian Journal
of Zoology Supplementary Series 31: 1-67.
1979. A phylogenetic subdivision of Australian
skinks. Records of the Australian Museum 32(8):
339-371.
1983. The Australian scincid lizard genus Calyptotis
De Vis: resurrection of the name, description of
four new species, and discussion of relationships.
Records of the Australian Museum 35: 29-59.
1986a. Lygosomine (Scincidae) monophyly: a third,
corroborating character and a reply to critics.
Journal of Herpetology 20(1): 123-126.
1986b. Diagnosis of the Lerista bipes species group
(Lacertilia: Scincidae), with a description of a
new species and an updated diagnosis of the
genus. Records of the Western Australian
Museum 13(1): 121-127.
1989. The biology and evolution of Australian
lizards. (Surrey Beatty and Sons Pty Ltd: Chip-
ping Norton, Sydney). 264pp.
GREER, A.E. & COGGER, H.G. 1985. Systematics of
the reduce-limbed and limbless skinks currently
assigned to the genus Anomalopus (Lacertilia:
Scincidae). Records of the Australian Museum
37(1): 11-54.
HUTCHINSON, M.N., 1993. Family Scincidae. Pp.
261-279 In Glasby, C.J., Ross, G.J.B. & Beesley,
P.L. (eds). Fauna of Australia. Vol 2A. Amphibia
& Reptilia. (Australian Government Publishing
Service: Canberra). 439pp.
SADLIER, R.A., 1990. The scincid lizard genus Nan-
noscincus Günther: a revaluation, Memoirs of the
Queensland Museum 29(2): 487-494.
SHEA, G.M., 1990. The genera Tiliqua and
Cyclodomorphus (Lacertilia: Scincidae): generic
diagnosis and systematic relationships. Memoirs
of the Queensland Museum 29(2): 495-519.
STORR, G.M., 1971. The genus Lerista (Lacertilia,
Scincidae) in Western Australia. Journal of the
Royal Society of Western Australia 54(3): 59-75.
COGGERIA, A NEW SKINK FROM FRASER ISLAND
APPENDIX 1. Specimens examined in
preparation of Table 1 and indicated on the table
by supercript.
(a) Ophioscincus truncatus (QMJ28853,
QMJ28620, QMJ40988). (b) Lerista bougainvil-
lii (AMR91671, AMR88302), L. microtis
(AMR47895), L. lineopunctulata (AMR64384),
L. labialis (AMR104024). (c) Lerista bougainvil-
lii (AMR91671, AMR88302), L. microtis
(AMR47895), L. punctatovittata (AMR104137),
L. lineopunctulata (AMR64384), L. labialis
(AMR104024). (d) Calyptotis ruficauda and C.
scutirostrum after Greer, 1983; C. temporalis
based on (AMR60765), C. thorntonensis based
on (AMRS56575). (e) Lerista bougainvillii
(AMR91671, AMR88302), L. microtis
(AMR47895), L. punctatovittata (AMR 104137),
L. lineopunctulata (AMR64384). (f) Calyptotis
ruficauda and C. scutirostrum after Greer, 1983;
C. temporalis based on AMR60765 and C.
thorntonensis based on AMR56575. (g)
Anomalopus mackayi count based on
AMR13138. (h) Calyptotis lepidorostrum
(AMR59246); counts for C. temporalis and C.
thorntonensis inferred from X rays of AM type
series. (1) Anomalopus gowi (AMR63130), A.
leuckartii | (A&MR43949), A. mackayi
(AMR13138), A. pluto (AMR94362), A. swan-
soni (AMR5186), A. verreauxii (AMR6437);
Coeranoscincus frontalis (AMR3823), C.
reticulatus (AMR4795); Ophioscincus
cooloolensis (QMJ27384), O. ophioscincus
(AMR47642), O. truncatus (AMR866); Saiphos
equalis (AMR7242); Calyptotis lepidorostrum
(AMRS59246), C. ruficauda (AMR52339), C.
scutirostrum (AMR43061); C. temporalis and C.
thorntonensis (X rays of type series). (j)
AMR104138. (k) AM X rays - Calyptotis
lepidorostrum (A&MR90305), C. scutirostrum
(AMR76120), C. temporalis (unregistered), C.
thorntonensis (AMR56603), and C. ruficauda
(AMR69547). (1) Not applicable; ischium not
projecting in either Coeranoscincus frontalis
(AMR89278) or C. reticulatus (AMR6375). (m)
AMR104138. (n) AM specimens: Calyptotis
lepidorostrum (AMR90305, X ray), C. ruficauda
(AMR69547), C. scutirostrum (AMR76120), C.
temporalis (X ray), C. thorntonensis
(AMR56603, X ray). (o) Anomalopus leuckartii
(QMJ33156), A. verreauxii (QMJ57097), A.
brevicollis (QMJ33122), A. gowi (QMJ45361),
A. pluto (QMJ54040, QMJ54083, QMJ54213),
A. swansoni (QMJ42773); Calyptotis
lepidorostrum (QMJ57061), C. ruficauda
(QMJ26024), C. scutirostrum (QMJ30616), C.
temporalis (QMJ31794), C. thorntonensis
(QMJ58111); Saiphos equalis (QMJ56908).
242
A BANDY BANDY WITH A DIFFERENCE. Memoirs of
the Queensland Museum 39(2):242: - The Bandy Bandy,
Vermicella annulata, is well known for its distinct colour and
pattern (Shine, 1980). It is widespread in Queensland (Ingram
& Raven, 1991). It is represented in the Queensland Museum
by 210 specimens of which 209 are distinctly marked with
uniform 'rings' of black and white (Fig.1).
On 26 April, 1995, we found an unusually coloured and
patterned bandy bandy on Retro Station, via Capella MEQ
(22°51'47"S 147°54’45"E). This specimen, now QMJ 59875
(Fig.2), was under a hollow log in Poplar Box (Eucalyptus
populea) woodland, with a sparse shrub understorey. The
introduced species Buffle Grass (Cenchrus ciliaris) and
Parthenium Weed (Parthenium hysterophorus) were present.
The collection site is an ‘island’ surrounded by paddocks of
sunflowers and mung beans.
The specimen (TL 39cm) is unusual in being fawn-grey on
black (rather than white on black); having a distinct vertebral
stripe on the anterior third of its body (rather than 'rings'
only); having a ‘broken’ vertebral stripe posteriorly connect-
ing sets of 2-4 ‘rings’ (rather than ‘rings’ only); and in having
its venter virtually unmarked by the 'rings' (rather than con-
sistently ringed in white). In other respects (e.g. head shape,
scale counts) the specimen is unremarkable (Cogger, 1992).
Acknowledgements
The specimen was collected during a survey of reptiles of
the Brigalow Biogeographic Region in Queensland for the
Endangered Species Program of the Australian Nature Con-
servation Agency, Canberra. Lindsay Vagg provided access
to Retro Station. Jeff Wright provided the photographs.
Literature cited
Cogger, H.G. 1992. Reptiles & Amphibians of Australia
(Reed Books: Chatswood, Sydney). 775pp.
Ingram, G.J. & Raven, R.J. (eds) 1991. An atlas of
Queensland's frogs, reptiles, birds and mammals.
(Board of Trustees, Queensland Museum: Brisbane).
391pp.
Shine, R. 1980. Reproduction, feeding and growth in the
Australian burrowing snake Vermicella annulata. Jour-
nal of Herpetology 14(1): 71-77.
P.J. Couper, J.A. Covacevich, Queensland Museum, Box
3300, South Brisbane, Queensland 4101, Australia; K.R. Mc-
Donald, Queensland Department of Environment and
Heritage, P.O. Box 834, Atherton, Queensland 4873,
Australia; 2 December 1995.
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig.1A, Vermicella annulata from Acacia Ridge, Brisbane,
SEQ, showing typical, highly distinct, regular black and
white colour and pattern. B, V. annulata, from Retro Sta-
tion, via Capella, MEQ, showing unusual colour (fawn-grey
on darker grey) and pattern (some longitudinal lines with
irregular bands).
ASPIDITES RAMSAYI (BOIDAE) IN THE BRIGALOW BIOGEOGRAPHIC REGION OF
QUEENSLAND: OCCURRENCE, CONSERVATION STATUS
AND POSSIBLE BILBY ASSOCIATIONS
J.A. COVACEVICH AND P.J. COUPER
Covacevich, J.A. & Couper, P.J. 1996 07 20: Aspidites ramsayi (Boidae) in the Brigalow
Biogeographic Region of Queensland: occurrence, conservation status, and possible Bilby
associations. Memoirs of the Queensland Museum 39(2): 243-246. Brisbane. ISSN 0079-
8835.
Aspidites ramsayi is widespread in central, northwestern, southwestern and parts of eastern
Australia. In southwestern Western Australia, an isolated population in what is now
‘Wheatbelt’ is ‘endangered’ and A. ramsayi is ‘specially protected’ under state legislation.
In western New South Wales, A. ramsayi is ‘endangered’ at the state level. In the southern
Brigalow Biogeographic Region (BBR) of Queensland, it now occurs as an apparently
isolated population. Because brigalow habitats are poorly protected in reserves, because
much of the habitat of A. ramsayi in the BBR is grazed and farmed and because the area is
likely to be subjected to further modification, prospects for A. ramsayi in the area are poor.
Despite the fact thatA. ramsayi is not accorded special conservation status in Queensland,
we contend that this species in the BBR is ‘vulnerable’.
The common name ‘Bilby Snake’ is used for A. ramsayi in the Yuleba-Surat area of the BBR.
This may point to a former close association there between A. ramsayi and the Bilby,
Macrotis lagotis, an endangered species. [ ] Aspidites, Bilby, vulnerable species, endangered
species, Brigalow.
J.A. Covacevich & P.J. Couper, Queensland Museum, Box 3300, South Brisbane,
Queensland 4101, Australia; received 2 February 1996.
Aspidites ramsayi (Macleay, 1882) is
widespread in arid and semi-arid Australia. It is
known from an isolated population in south-
western WA, and from the Broome area, WA, in
a broad band across WA, the southern NT, north-
ern SA to southwestern and southern Qld, and
western NSW (museum records; Wilson &
Knowles, 1988; Cogger, 1992; Cogger et al.,
1993; Pearson, 1993; Sadlier, 1994). The isolated
population of A. ramsayi in southwestern WA is
endangered, and is 'close to extinction'... (Cog-
ger et al., 1993). Its range *... now largely coin-
cides with the northeastern wheat belt of Western
Australia' (Cogger et al., 1993). There have been
two recent (1992, 1996) records from the western
edge of the Nullabor Plain (D. Pearson, pers.
comm.). Probable reasons for the decline of this
population are identified as ‘... a combination of
factors, including clearance of habitat for agricul-
ture and grazing, and crop production.’ ... (Cog-
ger et al., 1993). Pearson (1993) suggested that
predation by foxes and cats may promote the
decline also. In western NSW, A. ramsayi is
endangered at the state level and is apparently
‘lost’ from grazing areas; ... ‘the most recent
sightings ... are from within reserves, so land
reservation could have a mitigating effect on the
processes causing the decline. ... (This is) ... a
wide- ranging species so habitat fragmentation
from clearing and heavy grazing could adversely
affect (it).' ... (Sadlier & Pressey, 1993; Sadlier,
1994).
In Queensland, A. ramsayi occurs in the dry
subtropics, between the Qld-NT and Qld-S A bor-
ders, and the Yuleba-St.George areas in south
central Qld (Fig.1). In the western part of its range
(near Bedourie-Birdsville-Thargomindah), A.
ramsayi is associated with desert and semi-desert
grasslands and chenopod woodlands on red sandy
soils and stony downs of the Channel Country
Complex (Sattler, 1986). As these areas are spar-
sely grazed and as there are no firm plans to
intensify this land use, no threats to the continued
survival of A. ramsayi in this part of its range are
evident. In the eastern part of its range (Roma-
Yuleba-Surat-St. George), A. ramsayi occurs on
black soils and in stony ridge country, in
Brigalow (Acacia harpophylla) woodland and
grasslands, of the Brigalow Biogeographic
Region (Sattler, 1986), and just west of this
region. These populations now appear to be iso-
lated from that in the far west of the state,
presumably as a result of European land use in the
Mitchell Grasslands and Mulga Lands of the in-
244
FIG. 1. Queensland showing Brigalow Biogeographic
Region and distribution of the Woma/'Bilby Snake’,
Aspidites ramsayi. Squares 2 museum specimens.
Triangles = reliable sight records,
tervening area. Six specimens of A. ramsayi have
been recorded from the interface of the ‘western’
and ‘eastern’ populations: 1915, QMJ944 Avon-
dale Station 23°36’S 143°16’E; 1922,
SAMRO01101 Minnie Downs 25°02’ 145°52’;
1949, QMJ 7454 Jundah 24°05’ 143°04’; 1987,
sight only, Mariala N.P. 26°05’ 145?04' (P.Mc-
Crae, pers. comm.); 1995, sight only Morven -
Augathella road 26°06’S, 147°14’E, (C.Dollery,
pers. comm.); 1 April, 1996, sight only, 20km N
Charleville on Augathella road 26*17'146?17*
(C. Dollery, pers. comm.). Examination of all
Queensland records of A. ramsayi suggests that
present ‘western’ and ‘eastern’ populations were,
but are no longer, continuous.
Specimens from western and eastern localities
are biochemically indistinct (S. Donellan pers.
comm.). Three of these records are between 40
and 80 years old. The remainder are recent, but
from very close to the BBR. It seems reasonable
to suggest A. ramsayi no longer occurs in the
500km stretch between the Betoota (25°42’
140°50’) area in the west, and Charleville-Mor-
ven-Augathella (26°25’S,147°07’ E-25°48’S,
146°35’E) to the east. For conservation, the
‘eastern’ population of A. ramsayi should be con-
MEMOIRS OF THE QUEENSLAND MUSEUM
sidered virtually confined to the southern BBR
and extreme eastern portion of the Mulga Lands.
There are parallels between the populations of
A. ramsayi in and on the margins of Queensland's
BBR and southwestern WA. Both appear to be
isolates. Both occur in areas prized for agricul-
ture/grazing. Although much of the natural
habitat of A. ramsayi in southwestern WA has
been cleared, large areas suitable for the species
persist in the northern Wheatbelt, and from the
eastern edge of the Wheatbelt to the Nullabor
Region and the goldfields (D. Pearson pers.
comm.) The natural habitat of the south central
Qld (i.e. ‘eastern’) populations of A. ramsayi still
is subject to grazing and some agriculture, but
what appear to be a healthy populations of A.
ramsayi occurs in the area. Notwithstanding this
apparent, current security, there are no grounds
for complacency about the status of these popula-
tions. Their habitat is not protected in any nation-
al parks, although State Forests 328 (Yuleba) and
75, 41, 60 and 48 (in the St George area) are
important potential refuges for the species,
should plans to expand agriculture in the area
proceed.
Several authors have commented in the last
decade on the plight of BBR plant communities.
All agree that the area has been overcleared and
that representation of habitat diversity in reserves
is very poor (Sattler & Webster, 1984; Gasteen,
1985; Sattler, 1986, 1993; Davie etal., 1994). The
remaining woodland of the Morven-Roma-
Yuleba-Surat-St. George area of the BBR where
the ‘eastern’ populations of A. ramsayi occur is
good grazing land. Much of it has potential for
agriculture. If patterns of development elsewhere
in the BBR ensue, barring implementation of
clearing restrictions currently being discussed
(e.g. Central Queensland News 28.04.95), it
seems reasonable to suggest A. ramsayi should be
considered ‘at risk’ in the region. The potential
threat is emphasised by the apparent fate of
populations in south-western WA and western
NSW.
Our data suggest that A. ramsayi in and on the
western margins of the BBR of Queensland
should be recognized as a ‘vulnerable’ species
(Ingram & Raven, 1991). That is, A. ramsayi
there is ... ‘not presently endangered but at risk
over a longer period of time through continued
depletion ... largely occur (s) on sites likely to
experience changes in land use which would
threaten the survival of the species in the wild’.
... (Ingram & Raven, 1991).
STATUS OF ASPIDITES RAMSAYI IN QLD BRIGALOW BELT
BILBY ASSOCIATIONS?
A. ramsayi is known widely by the common
name *Woma' (Wilson & Knowles, 1988; Cog-
ger, 1992); in the Roma-Y uleba-Surat area, it has
been known for many years as the 'Bilby Snake’.
The name ‘Woma’ is not in general use in this
area (J.Harland, R. Allwood pers. comm., March,
1995).
The Bilby, Macrotis lagotis Reid, 1837, is an
endangered species, once widely distributed in
arid and semi-arid Australia, but now with a
restricted range. In Qld, it is confined to a few
isolated, small areas of the southwest (Gordon et
al, 1990; Southgate, 1990; Department of En-
vironment & Heritage, 1995). The use of the
common name ‘Bilby Snake’ for A. ramsayi
would be unremarkable if there were not a cor-
relation between the former ranges of both
species, including the BBR of Queensland. Dis-
tribution maps (Southgate, 1990; Gordon et al.,
1990; Ingram & Raven, 1991) for M. lagotis, and
A. ramsayi (Wilson & Knowles, 1988; Ingram &
Raven 1991; Cogger, 1992) show that these two
species at one time occurred widely, often in the
same areas. However, Womas occur in several
areas from which Bilbies are not known (e.g. the
Pilbara, southern Kimberley, the Gibson Desert
west of the Clutterbuck Hills to near Shark Bay
in WA, in the area south of Perth, and in much of
SA). A Queensland Museum specimen of M.
lagotis, QMJ90, attests to its presence in the
southern BBR. This is a current, but potentially
threatened, stronghold of A. ramsayi. The M.
lagotis is from Surat, and was collected in 1912,
by H. Donaldson. Local, longtime naturalists do
not know now anything of the Bilby in the
Yuleba-Surat area beyond the local name, 'Bilby
Snake’ (R. Allwood, T. Broughton, J. Harland
pers. comm., March 1995).
Whether or not there was an association be-
tween the Bilby and so called Bilby Snake in this
area is uncertain. Womas are known from 'Bilby
areas’ of Queensland's Channel Country (P.
McRae, pers. comm.), but we have no observa-
tions of direct association of the two species in
the BBR. A. ramsayi is known to prey equally on
reptiles and mammals (Shine, 1991) and, in the
Y uleba-Surat area, to occur on sandy soils and in
'stony country ... and go down sink holes'
(R.Allwood pers. comm., March 1995). It does
not seem unreasonable to surmise that 'Bilby
Snakes' may, at least in this area, have utilized
Bilby burrows for shelter, and Bilbies as food, at
least enough to have given rise to their local
245
name. Several recently collected (and released),
or road-killed specimens of ‘Bilby Snakes’ in the
BBR have been mammal feeders, containing hare
and rabbit remains. (R.Allwood, J.Nixon-Smith
pers. comm.). With the demise of the Bilby and
the rise of rabbits, the snakes may have been
forced to prey on common introduced mammals.
Many landholders have reported finding ‘Bilby
Snakes’ in and near rabbit warrens recently (C.
Eddie pers. comm.).
ACKNOWLEDGEMENTS
Mr C. Eddie, Mr P. McCrae and Mr C. Dollery
(Queensland Department of Environment), Mr
Trevor Beetson of Primary Industries (Forest Ser-
vices), Mr S. & Mr R. Irwin, Mr W. Irwin, Mr J.
Harland, Mr R. & Mrs B. Allwood, Mr J. Nixon-
Smith, Mr T. & Mrs J. Broughton and the
Curators of Reptiles of the state museums in
Australia have assisted us to map the occurrence
of A. ramsayi. Field work in the southern
Brigalow Biogeographic Region of Queensland
was funded the Australian Nature Conservation
Agency through its Endangered Species Pro-
gram. Our grant was administered by the
Queensland Department of Environment &
Heritage (now Department of Environment).
LITERATURE CITED
COGGER, H.G. 1992. Reptiles and amphibians of
Australia. (Reed Books: Chatswood, Sydney).
775p.
COGGER, H.G., CAMERON, E.E., SADLIER, R.A.
& EGGLER, P. 1993. The action plan for
Australian reptiles. (Australian Nature Conserva-
tion Agency: Canberra). 254p.
DAVIE, J.S., BARRY, S. & MORGAN, G. 1994. Plan-
ning for nature conservation in rurual environ-
ments: the Brigalow Belt in central Queensland.
Pp. 69-89. In Hynes, R.A. & Johnson, R.W. (eds).
Rural Queensland: a sustainable future. The ap-
plication of geographic information systems to
land planning and management. Proceedings of a
symposium held 23-24 November, 1989. (The
Royal Society of Queensland: Brisbane). 151p.
DEPARTMENT OF ENVIRONMENT &
HERITAGE, 1995. A rescue plan for the Greater
Bilby (Macrotis lagotis) in Queensland. Un-
published report of the Conservation Strategy
Branch, 25p. (Department of Environment &
Heritage: Brisbane).
GASTEEN, W.J. 1985. The Brigalow lands of eastern
Australia - agricultural impact and land use poten-
tial versus biological representation and stability.
Pp. 45-49. In Gasteen, J., Henry, D. & Page, S.
(eds). Agriculture and conservation in inland
246
Queensland. (Wildlife Preservation Society of
Queensland: Brisbane).
GORDON, G., HALL, L.S. & ATHERTON, R.G.
1990. Status of bandicoots in Queensland. Pp.37-
42. In Seebeck, J.H., Brown, P.R. Wallis, R.L. &
Kemper, C.M. (eds). Bandicoots and Bilbies:
(Surrey Beatty & Sons: Chipping Norton).
INGRAM, G.J. & RAVEN, R.J. 1991 eds. An atlas of
Queensland frogs, reptiles, birds and mammals.
(Queensland Museum Board of Trustees: Bris-
bane). 391p.
PEARSON, D.J. 1993. Distribution, status and conser-
vation of pythons in Western Australia. Pp. 383-
395. In Lunney, D. & Ayers, D. (eds).
Herpetology in Australia. A diverse discipline.
(Royal Zoological Society of New South Wales:
Sydney). 474p.
SADLIER, R.A. & PRESSEY, R.L. 1994. Reptiles and
amphibians of particular conservation concern in
the western division of New South Wales: a
preliminary review. Biological Conservation 69:
41-54.
SADLIER, R.A. 1994. Conservation status of the rep-
tiles and amphibians in the Western Division of
New South Wales - an overview. Pp. 161-167. In
Lunney, D., Hand, S., Reed, P. & Butcher, D.
(eds). Future of the fauna of Western New South
Wales. (The Royal Zoological Society of New
South Wales: Mosman, Sydney). 245p.
MEMOIRS OF THE QUEENSLAND MUSEUM
SATTLER, P.S. 1986. Nature conservation in
Queensland: planning the matrix. Proceedings of
the Royal Society of Queensland 97: 1-21.
SATTLER, P.S. 1993. Towards a nationwide biodiver-
sity strategy: the Queensland contribution.
Pp.313-325. In Moritz, C. & Kikkawa, J. (eds).
Conservation Biology in Australia and Oceania
(Survey Beatty & Sons: Chipping Norton, Syd-
ney).
SATTLER, P.S. & WEBSTER, R.J. 1984. The conser-
vation status of Brigalow (Acacia harpophylla)
communities in Queensland. Pp.149-160. In
Bailey, A. (ed.). The Brigalow Belt of Australia.
(The Royal Society of Queensland: Brisbane).
SHINE, R. 1991. Australian snakes. A natural history.
(Reed Books Pty Ltd: Balgowlah). 223p.
SOUTHGATE, R.I. 1990. Distribution and abundance
of the Greater Bilby Macrotis lagotis Reid (Mar-
supilia: Peramelidae). Pp.293-302. In Seebeck,
J.H., Brown, P.R., Wallis, R.L. & Kemper, C.M.
(eds). Bandicoots and Bilbies. (Surrey, Beatty &
Sons: Chipping Norton). 392p.
THOMAS, M.B. & MCDONALD, W J.F. 1989. ‘Rare
and threatened plants of Queensland'. 2nd ed.
(Department of Primary Industries: Brisbane).
WILSON, S.K. & KNOWLES, D.G. 1988. Australia's
reptiles. A photographic reference to the terrestrial
reptiles of Australia. (William Collins Pty Ltd:
Sydney). 447p.
LERISTA ALLANAE (SCINCIDAE: LYGOSOMINAE):
60 YEARS FROM EXHIBITION TO EXTINCTION?
J.A. COVACEVICH, P.J. COUPER AND K.R. MCDONALD
Covacevich, J.A., Couper, P.J. & McDonald, K.R. 1996 07 20: Lerista allanae (Scin-
cidae:Lygosominae): 60 years from exhibition to extinction? Memoirs of the Queensland
Museum 39(2): 247-256. Brisbane. ISSN 0079-8835.
Lerista allanae (Longman, 1937) is known from 12 specimens collected between 1929 and
1960, from 3 localities (Retro, Logan Downs and Clermont), in a small section of
Queensland’s central Brigalow Biogeographic Region. The species is/was confined to ‘black
soil’ downs. The area from which L.allanae is known has undergone dramatic change since
first settlement; itis now farmed or grazed, so little natural vegetation remains.Many searches
by specialists have failed to find L.allanae. We conclude that L.allanae may be extinct, while
acknowledging that this is impossible to prove. If it is not extinct, L.allanae is critically
endangered. Despite its ‘obscurity’, the possible extinction/endangered status of L.allanae
is a matter of concern, notwithstanding the extremely high economic value of agriculture in
the Brigalow Biogeographic Region. The apparent demise of L.allanae may herald loss of
other reptile species.
We recommend seeking the co-operation of land holders to protect remnants of natural
vegetation in its range; recognising the vital importance of remaining natural grasslands like
Gemini National Park; implementing a pit-trapping programme to increase the chance of
re-locating the species; and examining the need to protect, at least, the type localities of
Brigalow Biogeographic Region species where they still occur. [ ]Leristaallanae, extinction,
endangered species, Brigalow Biogeographic Region.
J.A. Covacevich & P.J. Couper, Queensland Museum, P.O. Box 3300, South Brisbane,
Queensland 4101, Australia; K.R. McDonald, Queensland Department of Environment, Box
834, Atherton, Queensland 4883, Australia; received 4 April 1996.
Lerista allanae (Longman, 1937) is a burrow-
ing skink from a small area of Queensland's black
soil downs in the Central Brigalow Biogeo-
graphic Region. It is known only to taxonomists,
and has been collected by, at most, a handful of
people, between 1929 and 1960. No photographs
are known of L.allanae in life.
Thirteen museum specimens comprise the total
knowledge base of this species, one of 11 species
of Australian reptiles recognised as ‘endangered’
(Cogger et al., 1993).
SPECIMEN HISTORY
The 13 specimens of L.allanae were collected
in 3 localities, all in close proximity to one
another, in the undulating downs (= gently un-
dulating plains formed on Tertiary basalt, Per-
mian shales, sandstone and unconsolidated
sediments of the Oxford Land System) of the
central Brigalow Biogeographic Region (J. Mc-
Cosker pers. comm.): ‘20 miles W of Capella’ =
Retro Station 22°51°53"S 147°53’43"E*, Aug.
1929+, J.R. Slevin, (AMR13819); 6 Aug. 1929,
J.R. Slevin, (CAS77101, ‘missing’ 14 Feb,
1988); 12 Dec, 1936, J.R. Slevin, (CAS77099);
13 Aug, 1929, J.R. Slevin (CAS77100); Retro
Station, Sept, 1936-Nov 1937, Maida (Mrs Percy
C.) Allan, (QMJ6180 holotype, QMJ6040,
QMJ6179 paratypes, QMJ6238, QMJ6308,
QMJ6429, QMJ6430); Logan Downs approx.
22?25'S 147°55’E, 16 Nov, 1948+, H.Wom-
ersley, (SAMR2823); Clermont 22°50’S
147°38’E, Jan, 1960+, C.Vallis (QMJ12232),
(Couper & Ingram, 1992; Cogger et al., 1993;
Shea, 1993; Covacevich & Couper, 1994; J.V.
Vindum, pers. comm).
Heber Longman described Rhodona allanae,
naming it for the collector of the first specimens
sent to him. However, nearly a decade earlier,
J.R. Slevin from the California Academy of
Sciences, had collected specimens of this species.
This material, from ‘20 miles W of Capella’
(=Retro Station) was not included by Longman
*(Latitude and longitude calculated at Retro homestead, in the absence of precise data on collection locality/ies.
+Date of registration, unless identified as a date of collection. Registration occurs soon after collection, generally.
248
in his type series. Longman’s types (Covacevich
& Couper, 1994) were sent to him by Mrs Percy
(Maida) Allan, wife of the manager of Retro, The
correspondence between Longman and Mrs
Allan (Queensland Museum Archives) is a lively
record of his excitement at ‘the find’ on Retro,
and her keen interest in natural history. It also
sheds some light on the colour of L.allanae and
on its habitat preferences. (e.g. *...an elongated,
grey, dark - spotted skink... in litt., H. Longman
to Mrs P.C. Allan, 20 May, 1936; 'This is a very
rare and interesting lizard, and I hope that other
specimens can be obtained and forwarded. It is
the first of this particular kind to be obtained in
Queensland, and it is probably new to science...
in litt., Longman-Allan 10 Sept., 1936; '...In this
lot of lizards there is a very good specimen of the
elongated one, the ‘Retro’ lizard, they are not
easy to find, but have asked the man who brings
in new soil for garden to keep a lookout as they
seem to be down a few inches in black soil under
tussocks of grass and so far we have had four from
that one patch. .. in litt, M. Allan to Heber
Longman, 18 June, 1937; *... I was particularly
pleased to have two additional specimens of the
elongated grey, dark-spotted skink lizards. These
will probably be described as new in the next
number of our Memoirs. They are allied in some
respects to a North Queensland species named
after Sir Hubert Wilkins, and to a West Australian
species. Congratulations on this discovery. ...' in
litt., Longman to Allan 20 May, 1937).
Longman was keen to exhibit the new reptile.
..'As the type and two paratypes should be
preserved downstairs to avoid fading, as far as
possible, I hope that you will be able to find other
specimens for exhibition and for exchange with
one or two other museums.’ ... (in litt., Longman
to Allan, 18 June, 1937). Specimen QMJ6238,
registered 28 June, 1937, was placed on display
in the galleries of the Queensland Museum,
presumably soon after registration. It was
removed from exhibition, faded, but otherwise in
good condition, on 18 Aug., 1954.
Data from the registers, correspondence and
literature on the habitat of L.allanae are scant.
Slevin's (1955) description of Retro, from his
visit there in 1929 and 1936, *... open grasslands
and scattered gum trees with moderately heavy
groves of tea tree and an occasional bottle tree.
Both black and red soil are in evidence. Though
the surrounding country did not look attractive as
a collecting ground it proved to be excellent, with
a large number of species.’ ...; and Mrs Allan's
description of the skink's association with ‘tus-
MEMOIRS OF THE QUEENSLAND MUSEUM
socks of grass’ and being found ‘down a few
inches in black soil’ are all that is known. That
four specimens of the 10 known from Retro came
from *one patch', that only 12 have been lodged
in museums, and that both Longman and Mrs
Allan allude to its ‘scarcity’ suggest that L.al-
lanae may have been always both ‘rare’ and
highly-localized in the black soil downs of the
Capella-Clermont area. These data also suggest
that the species was difficult to find. The exact
collection site for these specimens was not
referred to in Longman’ s description of L.allanae
(1937), and exact collection sites for the
specimens from Logan Downs and Clermont also
are imprecise and devoid of habitat notes.
Slevin’s collections in 1929 and 1936 suggest
that L. allanae may have been scarce always.
Slevin was a skilfull collector. California
Academy of Science records show he collected
1200 reptile specimens of 33 species on his two
field trips to Retro. Only three specimens of the
then undescribed L. allanae were collected.
Reduction in size and intensification of land use
on Retro have gone ‘hand in hand’, since the early
1920’s. This pattern, now being implemented
more quickly than ever on the central downs of
the Brigalow Biogeographic Region, is not new
for rich black soil downs in Qld. In southeastern
Qld, similar pressures to increase farm produc-
tion have resulted in dramatic changes in ver-
tebrate and plant species diversity on the Darling
Downs (Covacevich unpubl. data; Fensham pers.
comm.).
L. ALLANAE VS L. COLLIVERI
The recent separation of L. colliveri Couper &
Ingram, 1992 from L. allanae was based on what
could be regarded as minor external differences.
Lerista colliveri was described from 37 spec-
imens which had been identified as L. allanae.
These specimens account for distribution maps
(Cogger, 1975, 1979, 1983, 1986, 1992; Ehmann,
1992; Wilson & Knowles, 1988) which suggest
that L. allanae occurs beyond the Capella-Cler-
mont area to the vicinity of Hughenden and the
Townsville area.
With Couper & Ingram's (1992) separation of
L. colliveri, L. allanae is again a narrowly-
restricted species, which has not been reported or
collected since 1960, despite many searches.
Comparison of the recent redescription of L. al-
lanae from the type specimens and of L. colliveri
(Couper & Ingram, 1992) and the observations of
Shea (1993) shows they differ in colour pattern
LERISTA ALLANAE
and in the number of long toe lamellae. In L.
colliveri the forelimb varies from a nubbin to a
style. L. allanae has no forelimb.
With these data in mind, we recognise that the
differences between L. allanae and L. colliveri
could be seen as too minor to warrant species
recognition. If this were so, L. colliveri would be
treated as a junior synonym of L. allanae and, as
colliveri were collected relatively recently (1977-
1985), some of the concern about L. allanae
would abate.
However, separation of species within Lerista
from only slight or even without external dif-
ferences, has well established precedence. Char-
acters used by Greer et al. (1983) to separate L.
cinerea, L. storriand L. vittata from one other are
also subtle.
In separating L. emmotti from L. punctatovit-
tata Ingram et al., (1993) identified only one
character as diagnostic, a didactyle forelimb vs a
monodactyle forelimb or monostylus. Estimates
of genetic distance derived from allozyme
electrophoresis showed that the two 'forms' war-
ranted recognition as distinct species, despite the
fact that development of the toes in degenerate-
limbed skinks is highly variable. In other genera
also, subtle morphological differences have been
used to support species delineations based essen-
tially on allozyme variation (e.g. Daugherty et al.,
1990a; Patterson & Daugherty, 1990). Regarding
the Oligosoma nigriplantare species complex of
New Zealand, Daugherty et al. (19902) observed:
‘A clear pattern of genetic and specific diver-
gence is overlain by a highly variable pattern of
color and morphological variation, often inde-
pendent of species boundaries. ...’ and that this
frustrated traditional taxonomic methods.
Another recent New Zealand example highlights
the importance of examining the distinctness of
L. allanae and L. colliveri. Daugherty et al.
(1990b) analysed the conservation implications
of recognising three taxa of tuatara, where only
one had been known.
As L. allanae has not been collected since 1960,
the possibility that genetic distance from L. col-
liveri can be determined from allozyme
electrophoresis seems remote. However, given
patterns elsewhere; that there are morphological
differences between L. allanae and L. colliveri;
and that the two are geographically isolated, we
consider L. allanae distinct from L. colliveri. We
also recognise the need for what might be termed
‘taxonomic caution’ in dealing with potentially
endangered taxa; and that ...“Taxonomies are not
irrelevant abstractions, but essential foundations
249
of conservation practice’ (Daugherty et al.,
1990b).
RETRO: SIZE AND LAND USE
Retro was established in or about 1861, from
the amalgamation of a series of ‘runs’ in the
Leichhardt Pastoral District. Initially, Retro in-
cluded close to 400 km? (Queensland Department
of Lands, 1866). By the 1880s it extended NE
from the Capella-Clermont road, and straddled
Retro Ck and its tributaries, (Queensland Depart-
ment of Lands, 1884) (Fig.1). Through partition
and sales, both as a result of government policy
and market vagaries, Retro's size has diminished
dramatically. By about 1920, a major section of
Retro had been excised near the Clermont-Capel-
la road (Queensland Department of Lands, ca.
1920); by the 1940s further excisions on the SE
boundary near Abor Ck and on the NE boundary
had been made (Queensland Department of
Lands, circa 1940s a,b); by 1951, Retro had been
reduced to c.72,000 acres (Queensland Depart-
ment of Lands, 1951). Retro is now a holding of
2208.774 ha. Former ‘Retro Station’ now forms
part of Dakota, Carramah, Meelia, Hazeldene,
Weimby Downs, Mt Oscar, Salt Springs,
Amatunga, Penaddi, and Grenada holdings
(Alick, 1995, map 3).
Intensification and simplification have charac-
terized land use on Retro. It was a sheep station
originally. In 1891, 71000 sheep were shorn on
the station (L.Vagg pers. comm.). On Retro, and
on most other sheep holdings in the Capella area
of the black soil central downs, sheep-grazing
changed natural pastures to such an extent that
‘new’ grasses and weeds bearing seeds contam-
inated ‘the clip’ beyond use. This forced a change
from sheep to cattle grazing in the area (J. Mc-
Cosker, pers. comm.). Cattle grazing progres-
sively gave way to mixed cattle and grain
production, with the availability of machines to
hasten clearing and improved grain markets in the
1940s-1950s. In 1951, Retro was held by The
Queensland British Food Corporation, and was
used primarily for grain production. In 1996,
Retro is ‘broadscale’ farmed, with some irriga-
tion, to produce sunflower seeds, mungbeans and
sorghum.
There are no data on early vegetation on Retro,
but from the early 1940s (soon after the Allan
collections of L.allanae), detailed description of
Retro's ‘natural’ vegetation is available. At that
time, Retro was primarily a cattle station. The
'Retro Freehold Lands' map, Queensland
250 MEMOIRS OF THE QUEENSLAND MUSEUM
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Gemini N.P.
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CAPELLA
Fig. 1.Retro Station, the type locality of Lerista allanae, showing progressive size reduction since the 1860s.
(///= approximate extent in the 1860s;
= boundary during the 1940s; —
= boundary, 1995). Also shown
are sites searched in attempts to locate L. allanae between 1970 and 1996. (hand collecting, 1970-1994 E,
1995-1996 A; pit-trapping, 1995-1996 x). Two pit-traps and one hand-collecting site for 1996 are not shown.
All are on Kevricia, 35km ESE of Capella. 1cm-4.8km.
Department of Lands, c. 1940s describes the
country and vegetation from north to south in
detail: ‘Scattered timber and Stone, Stony
Coolibah Ridges, Heavy Scrub with Stony Hills,
Some Slope Gullies and Stone, Plain, Ti-tree,
Scrubby and Gravelly Ridges, Plain with lightly
scattered Bauhinia, Black Soil Fairly Heavy
Scrub, Open Tableland Coolibah and Blood-
wood, / Open Plain, inter alia’ ... Only one tiny
area near the homestead, now abandoned, but still
standing (Fig.2) was cleared. This is described as
‘Old Cultivation’. Retro contains no ‘natural’
vegetation save for islands of Poplar Box
(Eucalyptus populnea) woodland with sparse
shrub understorey and open Coolibah (E. or-
gadophila) woodlands. Buffle Grass (Cenchrus
ciliaris), an introduced pasture grass, and the
Parthenium weed (Parthenium hysterophorus)
are common in areas not under crops.
SEARCHES FOR L.ALLANAE, 1968-1995
Between 1968 and 1995 there have been many
attempts to find L.allanae on Retro, the area
formerly part of ‘old’ Retro, and in ‘suitable’
habitats nearby. Other reptile specimens have
been collected during searches aimed at estab-
lishing if L. allanae still occurs (Fig.1) . These
LERISTA ALLANAE
251
Fig. 2.The abandoned homestead Retro Station, a relic of the 19th century, where Maida Allan, who collected
the type specimens of L. allanae, lived in the 1930s.
searches, which have intensified since 1992, have
not been methodical in terms of either seasonal or
habitat sampling. However, experienced collec-
tors have searched the area in ‘good’ reptile times
(summer and spring) many times. They have
found other burrowing, secretive or elusive, small
skinks (e.g. Lerista fragilis, Anomalopus
brevicollis, Glaphyromorphus punctulatus,
Menetia greyii and M. timlowi) in the area, but
L.allanae has not been located. Lerista spp. and
other burrowing skinks (e.g. Anomalopus spp.)
are not usually collected by non-specialists.
(Since 1985, 125 Lerista spp. have been added to
Queensland Museum reference holdings. Only
seven of these, i.e. 6%, were collected by people
other than herpetologists. Numbers for
Anomalopus spp. are similar, i.e. 8/62, 1396). It
is, therefore, not surprising that, despite recent
radio and newspaper publicity about L.allanae,
no specimens have been collected 'accidentally'
by ‘amateurs’
The Longman-Mrs Allan correspondence had
not been used as a basis for target-searching for
L.allanae prior to 1995, Using Mrs Allan's
descriptions; following consultations with Ms J.
McCosker (of Department of Environment,
Emerald) regarding soil and vegetation types,
especially grasslands; using data from maps of
‘old’ Retro's boundaries; and bearing the Logan
Downs and Clermont collection localities in
mind, 4 sites were chosen for intensive L.allanae
searches, both by hand and using pit traps. Pit-
trapping is acknowledged widely to be a reliable
way of sampling reptiles, including species often
not found by hand- searching (Morton et al.,
1988). Many types of fences, traps and designs of
pit-traps have been used, with varying success
rates. In an arid habitat, it has been shown that , .
.'A simple straight line of pit-traps and a drift
fence with buckets approximately 7m apart is the
most effective . . .' (Hobbs et al., 1994). This
method was adopted.
In Gemini National Park, 2 pit-traps lines of 12
buckets each were set at different times, early
summer and late summer-early autumn. The first
(22°30°25"S 147?5]'06"E) was set from 15-21
Feb., 1995; the second (22?29' 15" 14752 14"),
from 15-20 April, 1995. Gemini National Park is
one of only three small national parks within
100km of Retro. Here, open grasslands, once
extensive in the central Brigalow Biogeographic
Region, have been protected from invasion by, or
252
deliberate replacement with, introduced grasses.
This is the only such locality in the area. Gemini
National Park supports Blue Grass (Dichanthium
sericeum) Downs, with open-groved Bauhinia
hookeri, and scattered Eucalyptus spp., on dry,
cracking, self-mulching, red-black soils. The in-
troduced weed (Parthenium hysterophorus) is
common on its margins, but has not invaded areas
where the native grass is dense. Gemini National
Park lies about 30km NW of the former boundary
of ‘old’ Retro. It adjoins Logan Downs, the
locality from which the 1948 specimen of L.
allanae was collected. (Surveys for L. allanae
were not conducted on Logan Downs because
permission to work there could not be obtained).
On Meelia, between 21-26 April, 1995, a pit-trap
line was set about 8km SE of the old homestead
on Retro, at 22^57' 12"S, 147°54’30"E. Meelia
lies within the boundary of ‘old’ Retro Station.
This site is one of the few, minute remnants of
uncleared land remaining in the area. The pit-trap
line was set in red-black, cracking clay soil with
open-groved B. hookeri and some Blue Grass (D.
sericeum). Much of Meelia has been cleared for,
or modified by, cattle grazing but the pit-trap site
is grazed only intermittently, and supports native
vegetation. Two pit traps were set on Kevricia at
23*14'77"S, 148716 71"E and 23*14'79"S, 148°
16' 19"E between 12-16 Feb., 1996. The former
(site 1) was set in a 30m wide strip of Brigalow
forest on deep, black, cracking soils. It had not
been cleared, but was disturbed by cattle and
adjoined a ploughed paddock. The latter (site 2),
an area of dense Brigalow regrowth, also grazed
and trampled by cattle, had been cleared once, in
1981. Kevricia is some 60km from Retro. It was
chosen for sampling because of its similarity to
some of Retro's former vegetation and because
of the scarcity of such habitats in the area. A
pit-trap was also set at Peak Downs (22^56'29"S,
148?04'60"E), 12-16 Feb., 1996. This was set in
cleared, grassed, black soil adjoining both the old
and new homesteads. (Peak Downs was estab-
lished in the same era as Retro and, like Retro
which it once adjoined, has been divided into
many small holdings. Like Retro also, Peak
Downs was a significant early collecting locality
for reptiles (Cogger et al., 1983).
Results of pit-trapping in these sites (Table 2)
indicate that burrowing and cryptic reptile
species still occur in the area from which L.
allanae is known.
Further, Queensland Museum register provides
a good record of incidental reptile collecting un-
dertaken on and near Retro since 1968. Hand
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 2. Summary of results from six pit-trap lines
set in 1995- 1996, at four locations close to Retro, in
search of L. allanae.
| : Gemini evricia| Peak
| Species NP Downs
|-— dh ab ase
| Gemmato hora
| nobbi E +
vittatus
willi x
" *
D. williamsi
Geyhra catenata
Heteronotia binoei + +
Glaphyromorphus
punctulatus
Lerista fragilis
Menetia greyii
M. timlowi i ESSO RC
Carlia pectoralis | | |+ +] |
collections by Queensland Museum staff/col-
leagues have been made at the following
localities (Fig.1): Gibson Downs (22'49'S,
148°12’E); Highland Plains (22°40’S, 148°08’E);
8km from Clermont on Clermont-Mackay road
(22°45’S, 147°38’E); Gaylong, via Capella
(22°46’S, 148?09'E); 500m ENE of old Corry
turnoff (22°45’S, 148°02’); Glencoe Stn
(23°08’S, 148°14’E); 8-10km E of Capella at
Gregory Mine turnoff (22°58’S, 148°03 E); Retro
(22°52’S, 147°54’E); 20km N of Capella on Cler-
mont road (22°59’S, 14751 E); 17km NW of
Capella on Clermont road (23°00’S, 147°54’E);
Gemini NP (22°30’25"S, 147?01' 51"E); Mazep-
pa NP (22°11°51"S, 147718'48"E); Moonda
Siding (22°59°52"S, 147^53' 16"E); Peak Ra. NP
(22°30°25"S, 147?01'51"E); Huntly Downs
(22252'35"S, 147T07'E); Prairie Stn
(23?06'46"S, 147°47°36"E); Meelia (22°57’S,
147°54’); and Peak Downs (22°56’S, 148°04’E).
That L. allanae has not been found despite
recent pit-trapping and many hand-collecting
searches by specialists in the last 28 years does
not mean it is extinct. However, as other burrow-
ing species known only to specialists have been
found, and as L. allanae has not, is reason for
concern. It seems reasonable to suggest that, if L.
allanae survives, the sites on Gemini National
Park (‘secure’ under legislation); Meelia and
Kevricia (both freehold) are potential refuges for
the species.
+
+
+
+
LERISTA ALLANAE
L. ALLANAE, POSSIBLY EXTINCT?
Extinction is difficult, if not impossible, to
prove. The case of Tiliqua adelaidensis parallels
that of L. allanae in many ways. Both are known
from narrowly-restricted localities which have
been changed by grazing or farming; both
were/are known from only very small series of
specimens (20 T. adelaidensis vs 13 L. allanae);
for both, many unsuccessful searches by ex-
perienced herpetologists have been undertaken;
and both species are secretive. T.adelaidensis
was classed as ‘possibly extinct’ or ‘extinct’ by
many authorities (Ehmann, 1982, 1992; Burtonet
al., 1986; Cogger, 1992; Hutchinson, 1992a,b).
On 14 Oct., 1992, 33 years after it had been seen
last, a specimen of T.adelaidensis was found in
the gut of a Brown Snake, Pseudonaja textilis
(Armstrong & Reid, 1992).
The prospect that Lallanae will, like
T.adelaidensis, be rediscovered is not good. L.al-
lanae was collected/seen last in 1960. The three
known localities for L.allanae have undergone
dramatic change, including clearing of forests;
replacement of native grasses with introduced
species, some of which are aggressive weeds; the
replacement of grazing with intensive farming;
and heavy use of pesticides to combat insect
pests, particularly locust. All will have reduced
the species’ chances of survival. Thus it does not
seem unreasonable to regard L.allanae as possib-
ly extinct. L.allanae is a secretive species, like all
Lerista spp., and is unlikely to be collected by
farmers or graziers, and sent to a museum for
identification, if records for other Lerista spp. are
any guide; it may have some unusual life habit
that has assisted its survival in the face of what
appear to be major assaults on its only known
habitat; and it is possible that the only known
habitat for L.allanae may not be its true stron-
ghold. This review confirms that L. allanae is, at
least, endangered. In any terms it needs urgent
conservation attention. L. allanae ‘fits’ almost
perfectly the definition of ‘endangered’ (Cogger
et al., 1993): '*... Taxa in danger of extinction and
whose survival is unlikely if the causal factors
continue to operate ... whose numbers have been
reduced to a critical level or whose habitats have
been so drastically reduced that they are deemed
to be in immediate danger of extinction. Also
included are taxa that may now be extinct but
have definitely been seen in the wild during the
past 50 years and have not been subject to
thorough searching’... .
So little data are available on L.allanae that its
253
critical population level and habitat are unknown,
but the area from which it was collected has been
drastically altered. It may be extinct now, but has
been seen in the last 50 years (last in 1960). It has
been subject to many searches, including some
that would be termed methodical, if not
‘thorough’.
THE IMPORTANCE OF L.ALLANAE
If L.allanae is extinct, it is the first Australian
reptile species to be so since European settlement.
This species is neither visible nor ‘appealing’. It
has no charismatic quality. It has been seen by
only a handful of people, and it is poorly known
by even specialist naturalists and herpetologists.
It has no special status in the area from which it
was described.
The reasons usually proposed for striving to
maintain biodiversity are that: extinction can dis-
rupt an ecosystem and cause the demise of other
species; reduced diversity makes the world less
interesting; the present generation has a respon-
sibility to leave a world of maximum diversity =
quality, for future generations; we may want
properties (e.g. drugs) from species; and species
have intrinsic value and a ‘right’ to exist (Elliot,
1980). The possible extinction of L. allanae could
not disrupt the modified central downs of the
Brigalow Biogeographic Region or cause the ex-
tinction of other species. Its possible loss could
not be said to diminish seriously the lives of
future generations; and its extinction is unlikely
to result in the loss of some magical compound
for the betterment of mankind. The loss of L.al-
lanae may, like the canary in a coal mine, be a
warning. Being narrowly restricted, rare and from
an area now intensely farmed and grazed, it may
be an indicator that other Brigalow species are
about to decline. Many other species either con-
fined to or occurring mainly in the Brigalow
Biogeographic Region, are already recognised as
rare and vulnerable (McDonald et al., 1991), and
the conservation status of other Brigalow reptile
species is of concern (e.g. Aspidites ramsayi after
Sadlier, 1994; Sadlier & Pressey, 1994; Cov-
acevich & Couper, this memoir).
Much of Queensland's Brigalow Biogeo-
graphic Region is rich grazing and agricultural
land. Much of it has already been modified exten-
sively and few reserves have been set aside for
the protection of Brigalow communities (e.g. Sat-
tler & Webster, 1984; Gasteen,1985; Sattler,
1986; Davie et al., 1994). It is now well known
that nothing, not even stringent habitat protec-
254
tion, guarantees species protection (e.g. frogs in
high altitude rainforests of Queensland; Ingram
& McDonald, 1993). However, the corollary
seems to be true for the Retro-Clermont-Logan
Downs Brigalow habitats. Clearing of native
forests comes close to ensuring species loss.
Emerald 65km S of Retro, is the main commer-
cial centre for the central downs. It ‘... is anticipat-
ing an economic boom which will triple the value
of production in the area’... which ‘...has beef
cattle, cotton farms, citrus orchards, wheat and
sorghum ...(and) coalmines. The key is a $235
million dam ... on the Comet River to provide
1,300,000 megalitres of water for new coalmines
and agricultural expansion. ’... (italics ours), Hay
(1995). In the face of such ambitious plans for
economic benefits from increased development
and clearing in the central downs, whether or not
L.allanae survives can be seen as trivial, That one
cost of such economic benefits could also be the
extinction of additional reptile (and other)
species, can be seen also as inevitable and trivial.
If, however, the pursuit of an ideal - to maintain
biodiversity - in the Brigalow Biogeographic
Region is worthwhile, the current status of L.al-
lanae is not trivial. Rather, it becomes a matter of
concern, necessitating, at least, restriction of
clearing in the area.
RECOMMENDATIONS
].Recognise that, inherently, L.allanae is worth
preserving, if it still exists.
2.Consult with landholders in the Retro-Logan
Downs area to encourage their co-operation to
protect/preserve remnant stands of native vegeta-
tion from cattle, clearing and pest plants.
3.Recognise that Gemini N.P. is the best refuge
in which L.allanae may still survive. The par-
ticular importance of this small park should be
recognised, and special effort be directed towards
its management (e.g. protection from Parthenium
hysterophorus, illicit grazing of cattle, fire).
4.Continue efforts, particularly through pit-
trapping to locate L. allanae. Whether such ef-
forts should be urgent, major, and involve
substantial research money and time, is a matter
for consideration by protection authorities (i.e.
Queensland Department of Environment and the
Australian Nature Conservation Agency).
5.Recognise that type localities and habitats of
other narrowly restricted species in the Brigalow
Biogeographic Region may be altered or
threatened by continued development. Where
species still occur at their type localities or in
MEMOIRS OF THE QUEENSLAND MUSEUM
small pockets of what may always have been a
narrow range and where these areas still support
native vegetation, every effort should be made to
protect these areas, at least, from clearing.
ACKNOWLEDGEMENTS
R. Sadlier, Australian Museum, Sydney, J.V.
Vindum, California Academy of Sciences, San
Francisco and A. Edwards, South Australian
Museum, Adelaide provided data from their hold-
ings; Bill Kitson, Department of Lands, Brisbane
located and interpreted old maps of Retro;
Lindsay Vagg, the owner of Retro, gave us access
to and information about Retro; Juliana Mc-
Cosker and Claire Smith, Queensland Depart-
ment of Environment, Emerald and Charleville,
helped us in the field, provided us with data on
possible study sites, and explained the Capella-
Clermont area land systems to us; Rod Fensham,
Queensland Herbarium, provided data on clear-
ing of Brigalow vegetation in southern
Queensland; H. Milne, Peak Downs, Rob
Donaldson, Meelia, P. and C. Murphy, Kevricia
and B. Garside, Hazeldene generously gave us
access to their properties and assisted us in the
field. Fieldwork was funded by the Australian
Nature Conservation Agency through its En-
dangered Species Program. Our grant was ad-
ministered by the Queensland Department of
Environment and Heritage (now Department of
Environment). Glen Ingram, Queensland
Museum and Glen Shea commented on the
manuscript.
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TWO NEW SPECIES OF FALSE SPIDER CRABS (CRUSTACEA: BRACHYURA:
HYMENOSOMATIDAE) FROM NEW CALEDONIA
P.J.F. DAVIE AND B. RICHER DE FORGES
Davie, P.J.F. & Richer de Forges, B. 1996 07 20: Two new species of false spider crabs
(Crustacea: Brachyura: Hymenosomatidae) from New Caledonia. Memoirs of the
Queensland Museum 39(2): 257-262. Brisbane. ISSN 0079-8835.
Two new species of Hymenosomatidae are described from estuarine habitats on the west
coast of New Caledonia. Odiomaris estuarius sp. nov. is most closely allied to the endemic
New Caledonian O. pilosus (A. Milne Edwards, 1873). Neorhynchoplax euryrostris sp. nov.
is unique in the genus with its prominent, broad, unilobate rostrum. [ ]Brachyura,
Hymenosomatidae, Odiomaris, Neorhynchoplax, New Caledonia.
Davie, P.J.F., Crustacea Section, Queensland Museum, P.O. Box 3300, South Brisbane,
Queensland 4101, Australia; Richerde Forges, B., Department of Oceanography, ORSTOM,
BP A5 Noumea, New Caledonia; 1 August 1995.
The Hymenosomatidae has been relatively lit-
tle studied, probably due to their small size and
cryptic habit. Lucas (1980) and Lucas & Davie
(1982) described 13 species from Australia, in-
dicating that family diversity had probably been
seriously underestimated. Ng & Chuang (1996)
provided the first overview of the Southeast
Asian fauna, recognising 24 species in 10 genera
(including 8 new species), making 15 new species
described from Southeast Asia since 1988 (Ng,
1988, 1991; Chuang & Ng, 1991). Thus the New
Caledonian fauna could be expected to harbour
some undescribed species. Ng & Richer de For-
ges (1996) reviewed the hymenosomatid fauna of
New Caledonia, establishing two new genera and
describing two new species. During mangrove
and intertidal collecting in December 1993 we
found 2 new species living under logs and rocks
in two estuaries north of Noumea, on the west
coast of New Caledonia.
SYSTEMATICS
Abbreviations used in the text are: MNHN,
Muséum national d'Histoire naturelle, Paris;
OM, Queensland Museum, Brisbane; c.b.,
carapace breadth; G1, male first gonopod.
Class CRUSTACEA
Order DECAPODA
Suborder BRACHYURA
Family HYMENOSOMATIDAE
Odiomaris estuarius sp. nov.
(Figs 1, 2A, C)
MATERIAL EXAMINED. HOLOTYPE MNHNB
25278, d (4.6 x 4.3 mm), Dumbea estuary, New
Caledonia, 8.12.1993, P. Davie & B. Richer de Forges.
PARATYPES MNHN-B25275, 4 8 (4.7 x 44; 4.0 x
3.6; 3.6 x 3.4; 2.7 x 2.7 mm), 4 9 (5.4 x 5.0; 4.1 x 3.9;
4.0 x 3.5; 3.8 x 3.4 mm). QMW20576, 5 d (4.8 x 4.4;
4.1 x 3.9; 4.0 x 3.7; 3.0 x 3.0; 2.5 x 24 mm), 5 9? (6.2
x 5,8 4.9 x 4.4; 4.3 x 4.0; 3.6 x 3.3; 3.4 x 3.1), same
data as holotype.
DESCRIPTION. Carapace subcircular, width
1.0-1.14 (mean=1.08) times length (including
rostrum); dorsal carapace surface flat to slightly
convex, with gastro-cardiac, cervical and
thoracic grooves; thoracic grooves short, angled
laterally; anterolateral and posterolateral angles
not indicated; carapace rim broad, minutely
granular; branchiostegites nearly vertical, slop-
ing out ventrally near last walking legs; rostrum
spatulate; eyes prominent, corneas swollen; pos-
tocular lobes relatively small in size, not
prominent in dorsal view, affording little protec-
tion for corneas; lower margin of orbital cup with
inner and outer blunt knobs; antennules con-
cealed in dorsal view when flexed; interanten-
nular septum narrow; rostral keel indistinct,
rostrum broadly rounded in frontal view; epis-
tome short; upper ridge of pterygostomial region
prominent, becoming broken posteriorly,
elevated on either side of pterygostomial and
branchiostegite junction anterior to cheliped
bases; surfaces of pterygostome and branchios-
tegite granular; Milne-Edwards' apertures nor-
mal.
Mouth field wider than long, almost completely
filled by 3rd maxillipeds; ischium of 3rd maxi-
llipeds shorter than merus along lateral edge, palp
not reaching ischio-merus junction, exopod only
visible proximally. Length of chelipeds in avail-
able males, and females, subequal to carapace
258
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. Odiomaris estuarius sp. nov., holotype male. A, dorsal view; B, frontal view. Scale line = 1 mm.
breadth; only slightly stouter than largest walking
legs; outer surfaces sparsely setose, and minutely
granular; propodus of largest male (holotype) not
inflated, similar to females, without ventral keel;
fingers curved, without marked proximal gape,
cutting margins bearing minute teeth only.
Length of walking legs c. 1.4 times carapace
width in both males and females, legs moderately
thick, dactyli distinctly longer and more slender
than propodi, densely setose, slightly curved,
with distinct subterminal tooth.
Female abdomen oval, convex, with 2 deep,
sinuous, submedial grooves running entire
length, defining a convex central region; seg-
ments 1-5 progressively longer and broader, with
base of telson forming greatest width; telson ar-
cuate, c. 1.75 times longer than segment 5; eggs
c. 0.4 mm diameter. Male abdomen with seg-
ments 1 and 2 shorter and broader than segments
3-5, segment 3 next shortest, segments 4 and 5 of
similar length, tapering to telson, telson bluntly
triangular, c. twice length of segment 5; interca-
lated plates laterally at base. Male G1 stout, with
2 distinct distal processes, a longer corneous
process and a shorter lobular elongation of stem;
conspicuously setose particularly on the disto-ab-
dominal face.
Body coloured light brown (alcohol preserved
specimens), often with a thin caked-on layer of
very fine sediment; feathered setae, often thick
with sediment, around mouth frame, Milne-
Edwards’ aperture, on walking legs, around edge
of female abdomen.
HABITAT. Under logs in estuary.
DISTRIBUTION. Only known from the type locality
on the west coast of New Caledonia.
ETYMOLOGY. For its estuarine habitat. Its only con-
gener lives in freshwater.
REMARKS. Odiomaris Ng & Richer de Forges,
1996, was erected for a freshwater species from
New Caledonia, O. pilosus (A. Milne Edwards,
TWO NEW FALSE SPIDER CRABS, NEW CALEDONIA
259
FIG. 2. A, C: Odiomaris estuarius sp. nov., holotype, A, first gonopod, C, abdomen. B, D: Neorhynchoplax
euryrostris sp. nov., holotype, B, first gonopod, D, abdomen. Scale line A, B = 0.2 mm; C, D = 0.5 mm.
1873), which had been included by Lucas (1980)
in Amarinus. Odiomaris estuarius sp. nov. fits
well with the generic diagnosis of Ng & Richer
de Forges (1996). The two genera appear close
but the most important characters separating
Odiomaris are: 1, G1 more slender, with two
distinct distal processes, a longer corneous
process and a shorter lobular elongation of the
stem; and 2, the elongated triangular telson of the
male abdomen which is significantly longer than
wide at base (breadth c. 0.9 or less times length);
whereas in Amarinus the telson is more-or-less
rounded and short, being much wider than long
(breadth 1.2 or more times length).
Odiomaris estuarius differs from O. pilosus by:
1. The numerous thin spinules (or stiff setae) that
cover the carapace margins, rostrum, pterygos-
tomial regions, walking legs and chelipeds,
which are so characteristic of O. pilosus, are
absent; instead the carapace has only soft short
fringing setae, and the legs and chelipeds have
longer, feathered setae. 2. The rostrum is marked-
ly wider and more spatulate than in O. pilosus;
also the carapace rim which continues across
260
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 3. Neorhynchoplax euryrostris sp. nov., holotype male. A, dorsal view; B, frontal view. Scale line = 1 mm.
behind the rostrum, is relatively flat, and does not
project forward onto the rostrum as a rounded
tongue as it does in O. pilosus. Seen in ventral
view the rostrum is broad and rounded, and does
not form a narrow V-shape as in O. pilosus. 3. The
eyes are large with the corneas swollen; the pos-
tocular lobes are relatively small in size, not
prominent in dorsal view, and afford little protec-
tion for the corneas. In O. pilosus the eyes are
relatively smaller and the postocular lobes are
prominent and project forward almost as far as
the anterior edge of the cornea. 4. The posterior
carapace grooves are almost indiscernible on O.
estuarius, whereas on O. pilosus the transverse
cardiac-intestinal groove is very distinct, and the
cardiac region itself is strongly defined. 5. It
seems that O. estuarius is a much smaller species
(present specimens less than 6.4 mm c.b.) than O.
pilosus (which from the literature attains at least
18 mm c.b.). The females of O. estuarius have a
mature abdomen from 5.4 mm, and the males
have mature gonopods from as little as 2.5 mm
c.b., although the largest male (4.6 mm) does not
yet have inflated chelae.
Neorhynchoplax euryrostris sp. nov.
(Figs 2B, D, 3)
MATERIAL EXAMINED. HOLOTYPE MNHN
B25276, 3 (3.5 x 3.4 mm), Dumbea, New Caledonia,
8.12.1993, P. Davie & B. Richer de Forges.
PARATYPES QMW19911, 3 9 (3.1 x 2.9; 2.8 x 2.6;
2.8 x 2.6), 1 9 (3.0 x 2.8 mm). QMW20573, 1 9 (2.0
x 1.8 mm), 2 ovig. 9 (3.3 x 3.1; 2.7 x 2.6 mm), Le Cap.
MNHN-B25277, d (2.7 x 2.7 mm) 3 ovig. 9 (3.1 x
2.85; 3.3x 3.1; 2.8 x 2.6 mm), all same data as holotype
except as noted.
DESCRIPTION. Carapace width in males sube-
qual to length (including rostrum); slightly wider
in females (1.04-1.11 times length). Dorsal sur-
face flattened, with gastric, cardiac and
postbranchial regions convex; gastrocardiac, cer-
vical and thoracic grooves well defined; bran-
chiostegites sloping out slightly towards leg
TWO NEW FALSE SPIDER CRABS, NEW CALEDONIA
bases; anterolateral margin with two low, blunt,
triangular prominences; posterolateral angle with
strong, curved, forwardly directed spine; rostrum
unilobate, margins moderately converging over
posterior two-thirds, more sharply converging
anteriorly to prominent acute medial lobe;
anterior rostral margin with short, curved fringing
setae, and two or more long setae on apex of
medial rostral lobe; eyes short, comea swollen,
fully visible in dorsal view; postocular lobes well-
formed; antennules longer than rostrum when
unflexed, basal segment with blunt lateral lobe
bearing setae; interantennular septum a pro-
minent narrow ridge, extending to base of medial
rostral lobe; rostral keel clearly defined, rounded;
antenna with long, very fine, flagellum; no anten-
nal spine; pterygostomial region with strong dor-
sal ridge.
Ischium and merus of third maxilliped with
strong setae along inner edges, ischium with dis-
tinct, acute, anteromedial lobe, palp stout, long,
reaching ischio-merus junction, exopodite and
epipodite conspicuous. Male chelipeds greater
than 1.5 times longer than carapace width, much
stouter than walking legs; propodus especially
inflated and expanded ventrally; merus with con-
spicuous spine on outer ventral margin at about
distal third, fingers slightly curved, meeting over
distal half, small gape proximally; dactyl with
large basal molar, and smaller medial tooth; fixed
finger with about six small teeth over proximal
half, largest medially; outer surface of palm and
fingers of holotype male conspicuously setose;
largest male with inflated chelae at 3.5 mm
carapace width, smaller (2.7 mm) male with
claws still immature. Female chelipeds stouter
than walking legs but much smaller and less
setose than male chelipeds, fingers meeting along
most of length, each with row of even low teeth;
walking legs long and slender, length of 2nd
walking leg c. 2.5 times carapace width, dactyli
slender and curved distally, with strong,
recurved, subterminal tooth, but otherwise un-
armed along ventral edge; walking legs setose,
especially on ventral surface of dactylus.
Female abdominosternal region typical of
Neorhynchoplax species, without pleopods, eggs
apparently brooded within abdomen and
cephalothorax cavity; segment 1 broad, segment
2 very short, segments 3-5 fused, forming major
part of abdomen, telson broad, laterally with
prominent, blunt, locking lobes, lateral margins
more-or-less straight, a rounded apical lobe dif-
ferentiated; female genital aperture subovate,
with raised broad anterior and outer shoulder,
261
sited on sternum just posterior to base of cheliped;
male abdomen with segments 1 and 2 short, fused
article of segments 3-5 expanding moderately to
maximum abdomen width then tapering uniform-
ly to telson, telson triangular, apically rounded;
G1 little curved, moderately stout, twisted, with
a distinct thin terminal portion tapering to a point,
and projecting towards sternum, setation as
figured.
Body colour light brown to brown (alcohol
preserved specimens); short setae on lateral
carapace walls, sternum, chelipeds, walking legs
and abdomen, with tendency to accumulate silt
particles and take on clubbed appearance.
HABITAT. Estuarine to tidal freshwater; in crevices in
rotting logs at edge of water at low tide; and under
stones in freshwater.
DISTRIBUTION. Only known from the type material
from the western coast of New Caledonia.
ETYMOLOGY, For its broad unilobate rostrum.
REMARKS. Ng & Chuang (1996) and Ng &
Richer de Forges (1996) restrict Elamenopsis A.
Milne Edwards, 1873, to only a few species most
closely resembling the type, E. lineatus A. Milne
Edwards, 1873, which is transversely broad with
short walking legs, and very different in ap-
pearance from Neorhynchoplax euryrostris sp.
nov. They resurrected Neorhynchoplax for the
bulk of species included in Elamenopsis by Lucas
(1980) and subsequent authors. Neorhynchoplax
euryrostris is easily recognised by the shape of
the rostrum. Only 4 species of Elamenopsis
(sensu Lucas, 1980) have a unilobate rostrum viz.
E. lineata A. Milne Edwards, 1873, E. inermis
(Takeda & Miyake, 1971), E. nasalis (Kemp,
1917) and E. minima Lucas & Davie, 1982. Of
the three species that properly belong in Neo-
rhynchoplax, E. nasalis and E. minima are very
different in having thin spine-like rostrums; and
E. inermis has only a very short, very narrow,
triangular rostrum.
One feature which Neorhynchoplax euryrostris
and N. inermis have in common, which does
suggest a close alliance, is the strongly twisted
G1, although it differs markedly in detail between
the two species.
ACKNOWLEDGEMENTS
Peter Davie is very grateful to the French Em-
bassy for providing funds to help undertake the
field collecting in New Caledonia. Ms Alison Hill
262
is especially thanked for drawing the dorsal and
frontal views of the two new species. Peter Ng
provided valuable criticism of the manuscript.
LITERATURE CITED
CHUANG, C.T.N. & NG, P.K.L. 1991. Preliminary
descriptions of one new genus and three new
species of Hymenosomatid crabs from
southeastern Asia (Crustacea: Decapoda:
Brachyura). Raffles Bulletin of Zoology 39(2):
363-368.
KEMP, S. 1917. Notes on the Crustacea Decapoda in
the Indian Museum. 10. Hymenosomatidae.
Records of the Indian Museum, Calcutta 13 (Part
5, No. 15): 243-379.
LUCAS, J.S. 1980. Spider crabs of the family
Hymenosomatidae (Crustacea: Brachyura) with
particular reference to Australian species: sys-
tematics and biology. Records of the Australian
Museum 33(4): 148-247.
LUCAS, J.S. & DAVIE, P.J.F. 1982. Hymenosomatid
crabs of Queensland estuaries and tidal mud flats,
including descriptions of four new species of
MEMOIRS OF THE QUEENSLAND MUSEUM
Elamenopsis A. Milne-Edwards and anew species
of Amarinus Lucas. Memoirs of the Queensland
Museum 20(3): 401-419.
NG, P.K.L. 1988. Elamenopsis mangalis sp. nov., a new
species of mangrove-dwelling hymenosomatid
crab from Singapore (Crustacea, Decapoda,
Brachyura). Crustaceana 55(3): 274-278.
1991. Cancrocaeca xenomorpha, new genus and
species, a blind troglobitic freshwater
hymenosomatid (Crustacea: Decapoda:
Brachyura) from Sulawesi, Indonesia. Raffles
Bulletin of Zoology 39(1): 59-63.
NG, P.K.L & CHUANG, C.T.N. 1996. The
Hymenosomatidae (Crustacea: Decapoda:
Brachyura) of Southeast Asia, with notes on other
species. Raffles Bulletin of Zoology, Suppl. 3.
NG, P.K.L. & RICHER DE FORGES, B. 1996. The
Hymenosomatidae (Crustacea: Decapoda:
Brachyura) of New Caledonia, with descriptions
of two new genera and two new species. Memoirs
of the Queensland Museum 39(2): 263-276.
TAKEDA, M. & MIYAKE, S. 1971. A new freshwater
crab of the family Hymenosomatidae from the
Palau Islands. Researches on Crustacea 4,5: 1-7.
THE HYMENOSOMATIDAE (CRUSTACEA: DECAPODA: BRACHYURA) OF NEW
CALEDONIA, WITH DESCRIPTIONS OF TWO NEW GENERA AND TWO NEW
SPECIES
P.K.L. NG AND B. RICHER DE FORGES
Ng, P.K.L. & Richer de Forges, B. 1996 07 20: The Hymenosomatidae (Crustacea:
Decapoda: Brachyura) of New Caledonia, with descriptions of two new genera and two new
species. Memoirs of the Queensland Museum 39(2): 263-276. Brisbane. ISSN 0079-8835.
All 5 previously described species of hymenosomatids were examined. Micas gen. nov. is
established for Elamena minuta A. Milne Edwards, 1873, and Micas falcipes sp. nov., whilst
Odiomaris gen. nov. is established for Elamena pilosa A. Milne Edwards, 1873. Specimens
previously referred to E. truncata (Stimpson, 1858) are here described as Elamena vesca sp.
nov. O Brachyura, Hymenosomatidae, New Caledonia, Indo-West Pacific.
Ng, P.K.L., Department of Zoology, National University of Singapore, Kent Ridge, Sin-
gapore 119260, Republic of Singapore; Bertrand Richer de Forges, Department of Oceanog-
raphy, ORSTOM, BP A5 Noumea, Nouvelle Calédonie; received 1 August 1995.
Five hymenosomatid crabs have been reported
from New Caledonia: Elamenopsis lineata A.
Milne Edwards, 1873, Elamena truncata
(Stimpson, 1858), Halicarcinus minutus (Milne
Edwards, 1873), Halicarcinus keijibabai
(Takeda & Miyake, 1971) and Amarinus pilosus
(Milne Edwards, 1873) (Lucas, 1980).
While the first author was revising the hymeno-
somatids of Southeast Asia (Ng & Chuang,
1996), the second author provided a collection of
6 species of New Caledonian hymenosomatids.
The study of this material, as well as the types of
Elamenopsis lineata and Elamena minuta,
resulted in a reappraisal of the taxonomy of the
New Caledonian Hymenosomatidae. Halicar-
cinus minutus and Amarinus pilosus are referred
to new genera, Micas and Odiomaris, respective-
ly. They both differ markedly from Halicarcinus
White, 1846, and Amarinus Lucas, 1980, respec-
tively. A new species of Micas is also described.
Comparisons with Elamena truncata (Stimpson,
1858) from Taiwan also show that E. truncata A.
Milne Edwards, 1873, is separate. We review the
5 previous species, including necessary new taxa;
2 further New Caledonian species are added by
Davie & Richer de Forges (1996).
Specimens examined are deposited in the
Queensland Museum (QM), Brisbane, Australia;
Muséum national d'histoire naturelle (MNHN)
Paris, France; Amsterdam Museum (ZMA), The
Netherlands; and Zoological Reference Collec-
tion (ZRC), Department of Zoology, National
University of Singapore. Measurements provided
are of the carapace width and length respectively.
G1 and G2=d first and second gonopods, respec-
tively. Terminology follows Melrose (1975).
SYSTEMATICS
Class CRUSTACEA
Order BRACHYURA
Family HYMENOSOMATIDAE
Elamenopsis A. Milne Edwards, 1873
Elamenopsis A. Milne Edwards, 1873: 324; Lucas,
1980: 190 (partim).
TYPE SPECIES. Elamenopsis lineatus A. Milne Ed-
wards, 1873, by monotypy.
REMARKS. Elamenopsis contains E. lineata and
E. ariakensis (Sakai, 1969). Lucas (1980)
synonymised Neorhynchoplax Sakai, 1938, with
Elamenopsis. Ng & Chuang (1996) redefined
Elamenopsis, restricting it to species with a
unilobed rostrum, laterally oval carapace and
short ambulatory legs; and resurrecting Neor-
hynchoplax Sakai, 1938, for the other Elamenop-
sis species recognised by Lucas (1980).
Elamenopsis lineata A. Milne Edwards, 1873
(Fig. 1)
Elamenopsis lineatus A. Milne Edwards, 1873: 324, pl.
18 fig. 4; Kemp, 1917; 250; Tesch, 1918: 26, pl. 1
fig. 5, 5a-c; Seréne & Umali, 1970: 58, pl. 5 fig. 11.
Elamenopsis lineata: Lucas, 1980: 192, figs. 3j, 5j, 8e,
10j; Chuang & Ng, 1994: 87; Ng & Chuang, 1996.
MATERIAL EXAMINED. HOLOTYPE MNHN
B651 9 (2.6 x 2.0 mm), Dotio, New Caledonia, M.
Batema. ZMA uncat., 1 d (3.8 x 2.8 mm), Sulawesi,
Indonesia, Siboga Expedition. QM W2341, 8,2 9,5
of Dunwich, SEQ. 15.7.1962, F, Vohra. ZRC
1994.4242, ex. QM W2337, à, 2 9, S of Dunwich,
SEQ, 12.7.1962, F. Vohra.
264
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. Elamenopsis lineata. A, holotype 9 (after Milne Edwards, 1873: pl. 18 fig. 4). B, holotype 2 (MNHN
651), New Caledonia. C-F, d (2.7 x 2.0 mm) (QM W2341) (after Ng & Chuang, 1996). G, 9 (3.4 x 2.5 mm)
(ZRC 1994.4242) (after Ng & Chuang, 1996), Australia. A, overall view. B, dorsal view of carapace. C, left
third maxilliped. D, right third ambulatory leg. E, ¢ abdomen. F, left Gl. G, 2 abdomen. Scales: A-F = 0.05
mm. G = 0.5 mm.
REMARKS. This species was treated in detail by
Ng & Chuang (1996). Milne Edwards (1873)
described it from the Dotio river, New Caledonia,
but it has not been reported from there since.
DISTRIBUTION AND HABITAT. New
Caledonia, Australia, Sulawesi (Tesch, 1918) and
Philippines. Milne Edwards (1873) reported the
species from a brackish water habitat while Tesch
(1918) reported his specimen from coral reefs,
which record was questioned by Lucas (1980)
and Ng & Chuang (1996). Lucas (180: 193) noted
that the species can be common on muddy sub-
strates in Australia.
Halicarcinus White, 1846
Halicarcinus White, 1846: 178; Lucas, 1980: 176
(complete synonymy).
TYPE SPECIES. Cancer planatus Fabricius, 1775, by
original designation.
HYMENOSOMATIDAE OF NEW CALEDONIA
REMARKS. Halicarcinus minutus (A. Milne Ed-
wards, 1873)[=Micas gen. nov. herein] and H.
keijibabai (Takeda & Miyake, 1971) have been
reported from New Caledonia. Lucas (1980)
redefinition of Halicarcinus is followed here.
Halicarcinus keijibabai (Takeda & Miyake,
1971)(Fig. 2)
Rhynchoplax keijibabai Takeda & Miyake, 1971: 165,
figs 1, 2.
Halicarcinus keijibabai: Lucas, 1980: 164.
MATERIAL EXAMINED. ZRC 1994.4283, d , inter-
lidal region, low tide at OUEMO, Nouméa, New
Caledonia, 2.7.1992, B. Richer de Forges.
REMARKS. This species was described from
New Caledonia (1 d and 1 9 ) (Fig. 2A, B) and
has not been reported elsewhere. Our ó agrees
with Takeda & Miyake's (1971) detailed descrip-
tion and figures. Characters (e.g. G1 and d ab-
domen) not figured by Takeda & Miyake (1971)
are illustrated here. Halicarcinus keijibabai is
very close to H. coralicola (Rathbun) but the
postocular tooth is less pronounced, the base of
the rostrum is fused with the postocular lobes
forming a broad band above the eyestalks, the
anterior lateral angle lacks an acute tooth, the
subhepatic/pterygostomial regions have only 1
(vs. 3) teeth, and most significantly, the Gl is
straighter and not medially curved (Fig. 2G).
Other differences in the carapace structure and
form of the ambulatory dactylus noted by Takeda
& Miyake (1971: 168) however, are subject to
variation and cannot be used. Lucas (1980: 164)
noted that the medial rostral lobe in H. coralicola
lacks long terminal setae, but this is incorrect as
this character is present in both species.
DISTRIBUTION AND HABITAT. New
Caledonia. Under rocks in the intertidal area, in
waters no more than 1 m deep. Same habitat for
H. coralicola in SE Asia and Japan (Chuang &
Ng, 1994; Ng & Chuang, 1996).
Micas gen. nov.
TYPE SPECIES. Elamena minuta A. Milne Edwards,
1873.
DIAGNOSIS. Carapace rounded; dorsal surface
smooth, grooves all well delineated, reaching to
lateral margins of carapace; cardiac region with
distinct longitudinal groove. Eyes visible dorsal-
ly. Rostrum trilobed, lobes short; median lobe
265
may or may not be separated from carapace by
crest. Ambulatory legs long, slender; dactylus
with 2 subterminal teeth. Male abdominal seg-
ments 3 and 4 fused; 9 abdominal segments 2-5
fused. G1 slender distally, stout basally, twisted
medially.
REMARKS. Kemp (1917) commented that
Elamena minuta should be placed in Halicar-
cinus. It has been collected only twice previously
(A. Milne Edwards, 1873; Takeda & Nonumura,
1976) and d & have not been described.
H. minutus has few of the important features of
Halicarcinus. The complete longitudinal cardiac
groove on the carapace is evident in no other
hymenosomatid, although H. hondai (Takeda &
Miyake) has a partial one (Lucas, 1980: 184, fig.
3H). The G1 of H. minutus is twisted twice, once
medially and again, more strongly 3/4 from the
base, with the tip slightly bifurcated. The am-
bulatory dactyli of M. minutus are very elongate,
much longer than for other Halicarcinus species
and have only two subterminal teeth, lacking the
series of sharp teeth present on many Halicar-
cinus species (with the exception of the enigmatic
H. filholi (De Man)). Like most Elamena, the ó
abdomen also has segments 3 and 4 fused, quite
unlike most Halicarcinus in which all the seg-
ments are free. The same is true for the 9 ab-
domen, which in M. minutus has segments 2-5
completely fused, with no sutures visible (most
9 9 of Halicarcinus have all segments free).
Unlike both Halicarcinus and Elamena, the 3
chelipeds are also short and not elongate. The
carapace of H. minutus also bears a close
resemblance to many species of Neorhynchoplax
s.s., but the third maxillipeds are quadrate, cover-
ing at least 3/4 of the mouthfield.
Micas falcipes sp. nov., is also described from
New Caledonia. A NE Australian species, H.
afecundus Lucas, 1980, is also assigned to Micas.
It has the same short trilobed rostrum, lon-
gitudinal groove on the cardiac region, elongate
ambulatory dactyli and 2 abdominal condition
(segments 2-5 fused) as M. minutus.
ETYMOLOGY. Latin mica, grain; for the small size
of the type species. Gender masculine.
Micas minutus (Milne Edwards, 1873)
(Fig. 3)
Elamene minuta Milne Edwards, 1873: 324 (partim),
pl. 18 fig. 5.
Elamena minuta: Tesch, 1918: 21 (partim).
Halicarcinus minutus; Takeda & Nonumura, 1976: 65,
266 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 2. Halicarcinus keijibabai. A, holotype d (1.7 x 2.0 mm [excluding rostrum]) (after Takeda & Miyake,
1971, fig. 1). B, allotype 9 (1.7 x 1.8 mm [excluding rostrum]) (after Takeda & Miyake, 1971, fig. 2). C-G, d
(2.3 x 2.7 mm) (ZRC 1994.4283), New Caledonia, A, B, overall view. C, d abdomen. D, left third maxilliped.
E, left third ambulatory leg. F, left third ambulatory dactylus. G, left G1. Scales = 0.5 mm.
HYMENOSOMATIDAE OF NEW CALEDONIA 267
»
FIG. 3. Micas minutus. A, B, D, E, lectotype 3 (3.4 x 3.5 mm) (MNHN B656Sa) (A, after Milne Edwards, 1873,
pl. 18 fig. 5). C, 9 (after Takeda & Nonumura, 1976, fig. 2). F-H, 9 (2.9 x 2.6 mm) (ZRC 1994.4250b). A,
overall view. B, C, carapace. D, & abdomen. E, left G1. F, right third ambulatory leg. G, left third ambulatory
dactylus. H, 2 abdomen. Scales: B, C, F, H = 1.0 mm. D, G 2 0.5 mm. E = 0.25 mm.
268 MEMOIRS OF THE QUEENSLAND MUSEUM
W
FIG. 4. Micas falcipes sp. nov. A-E, paratype 9 (2.9 x 2.6 mm) (ZRC 1196.477). F, paratype 9 (3.1 x 2.6 mm)
(MNHN B24915b). I-M, paratype d (2.8 x 2.7 mm) (MNHN B24915a). A, carapace. B, rostrum. C, G, left
third maxilliped. D, J, right third ambulatory leg. E, K, right fourth ambulatory leg. F, 9 abdomen. H, left chela.
I, carpus and merus of left cheliped. L, d abdomen. M, left G1. Scales: A, F, G, J-M = 1.0 mm. B-E, H, I = 0.5
mm.
HYMENOSOMATIDAE OF NEW CALEDONIA
fig. 2; Lucas, 1980: 177 (partim); Chuang & Ng,
1994: 88 (partim).
MATERIAL EXAMINED. LECTOTYPE. MNHN-
B656Sa, d (3.4x 3.5 mm), New Caledonia, M. Marie.
OTHER MATERIAL: ZRC 1993.6512, 9 (3.5 x 3.3
mm), ZRC 1994.4250a, 9 (ovigerous) (4.1 x 3.9 mm),
intertidal region, low tide at OUEMO, Nouméa, New
Caledonia, 2.7.1992, B. Richer de Forges.
REMARKS. Elamena minuta was described
briefly by Milne Edwards (1873) and although he
did not indicate the number of specimens he had
examined, he provided measurement of one
specimen 3.0 mm in carapace width (sex not
stated). His figure is very schematic and provides
almost no information on the carapace lateral
margins, third maxillipeds, ambulatory dactylus,
abdomen or Gl. The condition of the cardiac
region and posterior lateral spine was neither
figured or mentioned. In the MNHN are three
dried specimens which had been labelled as
Elamena minuta. All were collected by M. Marie
and labelled as types. The largest specimen (our
lectotype), a d 3.4 by 3.5 mm, carries a label
indicating that it was the specimen figured by
Milne Edwards. The other two specimens
(paralectotypes) belong to M. falcipes sp. nov.
The lectotype is still in good condition and shows
a clearly defined longitudinal groove on the car-
diac region and well developed posterior lateral
spines.
Takeda & Nonumura (1976: fig. 2) figured a 9
(Fig. 3B) with a prominent posterior lateral spine;
their specimen agrees very well with the lec-
totype d.
DISTRIBUTION AND HABITAT. New Cale-
donia. Under rocks in the intertidal zone, along
the shore or on the fringing reef of small coral
islands.
Micas falcipes sp. nov.
(Fig. 4)
Elamene minuta Milne Edwards, 1873: 324 (partim).
Elamena minuta: Tesch, 1918: 21 (partim).
Halicarcinus minutus: Lucas, 1980: 177 (partim);
Chuang & Ng, 1994: 88 (partim).
MATERIAL EXAMINED. HOLOTYPE
MNHMB25300 (ex ZRC 1993.6511), 8 (3.2 x 3.1
mm), intertidal region, low tide at OUEMO, Nouméa,
New Caledonia, 2.7.1992, B. Richer de Forges.
PARATYPES ZRC 1994.4250b, d (3.5 x 3.3 mm), 9
(ovigerous) (2.9 x 2.6 mm), MNHN B24915b, d (2.8
x 2.7 mm), 9 (3.1 x 2.6 mm), intertidal region, low tide
at OUEMO, Nouméa, New Caledonia, 2.7.1992, B.
269
Richer de Forges. PARATYPE, ZRC 1996.477, 9 (2.9
x 2.6 mm), station 10, shallow water in bay, OUEMO,
Nouméa, New Caledonia, 19.4.1995, B. Richer de
Forges. MNHN B656Sb, 2 à (2.5 x 2.3 mm, 2.5 x 2.3
mm), New Caledonia, M.E. Marie.
DESCRIPTION.Carapace width (at posterior
pair of angular lobes) greater than length; dorsal
surface almost flat, smooth, gastrocardiac
grooves deep, distinct; cervical and thoracic
grooves shallow; cardiac region with longitudinal
groove which is deeper on anterior half; pterygos-
tomian region with low knob. Anterolateral mar-
gin with low angle, never spiniform;
posterolateral margin gently convex; with dis-
tinct, unarmed swelling on posterior lateral angle.
Rostrum trilobate, lobes subequal in size; tip of
lateral below tip of median lobe; median lobe
appears continuous with the dorsal carapace sur-
face, crest separating it from carapace very low.
Eyes visible dorsally.
Third maxillipeds almost completely cover
mouth field when closed; ischium distinctly
shorter than merus along outer lateral edge; inner
lateral edge of merus lined with fine setae, inner
lateral margins meeting when closed; palp longer
than merus; exopod longer than merus, reaching
distal edge of merus.
Chelipeds equal, stouter than ambulatory legs;
surfaces smooth; fingers c.1/3 length of inflated
palm; fingers laterally flattened, slightly curved
inwards; cutting edges serrated, with quadran-
gular tooth at proximal portion of dactylus; tips
of fingers sharp, pointed.
Ambulatory legs slender, cross-section of
merus subcircular; distal dorsal edge of merus
and carpus with well developed but rounded
tooth; dorsal margin of propodus convex, median
height (measured from dorsal to ventral margins)
distinctly higher than distal and proximal heights;
margin of dactylus lined with dense, fine setae;
strongly curved, tip hooked, with 2 subequal sub-
terminal teeth; subdistal margin smaller and less
distinct tooth in between tip and subterminal
tooth.
ő abdomen triangular; segments 3 and 4 com-
pletely fused without trace of sutures; all other
segments free; telson more than 2 times length of
segment 6, lateral margins gently convex. Gl
slender, elongate, twisted medially. Female ab-
domen rounded, segments 2-5 fused, without
trace of sutures, telson c. half width of fused
segments 2-5, lateral margins sinuous; lateral
margins of fused segments 2-5 tapering gradually
towards segment 1.
270 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 5. Odiomaris pilosus. A, 3 (11.8 x 10.1 mm) (ZRC 1994.425 1a). B, 9 (8.8 x 7.8 mm) (ZRC 1994.4251b),
New Caledonia.
HYMENOSOMATIDAE OF NEW CALEDONIA
REMARKS.Micas falcipes sp. nov. resembles M.
minutus but can be separated by: the rounder
carapace (mainly because the anterolateral mar-
gins are more convex), a posterior lateral swelling
(sometimes dentiform, never spiniform), the
median rostral lobe continuous with the carapace
surface (vs. separated by a crest), the distal angle
of the dorsal margin of the ambulatory merus
more strongly produced, the dorsal margin of the
ambulatory propodus convex, the ambulatory
dactyli proportionately stouter and shorter, the ĝ
telson proportionately longer, with the lateral
margins gently convex and the G1 proportionate-
ly shorter. The posterior lateral swelling in M.
falcipes seems to be reliable in separating M.
minutus. M. minutus has well-developed, sharp
spines, whereas M. falcipes is never spiniform.
DISTRIBUTION AND HABITAT. New
Caledonia. One specimen was collected under a
sponge in 0.2m of water. The others are from
shallow waters in the intertidal region. It is typi-
cally brown to black in life, with the posterior part
of the postbranchial region sometimes white.
ETYMOLOGY. Latin falx and pes, for the sickle-
shaped ambulatory dactylus. A noun in apposition.
Odiomaris gen. nov.
TYPE SPECIES. Elamena pilosa Milne Edwards,
1873.
DIAGNOSIS. Carapace laterally oval, distinctly
broader than long; grooves on dorsal surface
well-defined. Third maxilliped quadrate;
anteroexternal angle of merus rounded, not
auriculiform; ischium broad, expanded. Am-
bulatory legs relatively short, dactylus with one
weak subterminal tooth. Male abdomen evenly
triangular; telson triangular, lateral margins al-
most straight, base with distinct, movable inter-
calated plates. G1 relatively slender, distal part
with well-developed pectinated process and a
smaller, weakly chitinised process.
REMARKS. Lucas (1980) referred E. pilosa
Milne Edwards, 1873, to Amarinus (type E. ?
lacustris Chilton), with all congeners from brack-
ish or freshwater. Lucas (1980: 198) noted that
the G1 of Amarinus is *... stout, curved at base,
otherwise with little curvature, terminal portion
with fine setae in tufts or scattered, terminating in
one or several lobes'. Holthuis (1968), however,
had figured the G1 of E. pilosa (as a species of
Halicarcinus) which is unlike that of any known
271
Amarinus, being distinctly more slender, the dis-
tal part possessing 2 distinct processes. Ten
species of Amarinus are known (Lucas, 1980;
Lucas & Davie, 1982; Ng & Chuang, 1996) and
all have very similar G1s, being short and stout.
Comparing E. pilosa with Amarinus reveals
several other differences. In E. pilosa the
anteroexternal angle of the merus of the third
maxilliped is not auriculiform (vs. distinctly
auriculiform), the d' abdomen is shaped dif-
ferently, the structure (including the telson with
the almost straight lateral margins) being evenly
triangular (vs. broadly triangular abdomen, the
telson being semicircular), G1 is more slender,
the distal part having a well-developed pectinated
process and a smaller, weakly chitinised process
(vs short, stout and without the two pectinated
processes). E. pilosa also differs in having the
carapace laterally oval (vs. circular to subcir-
cular) and the inner lateral margin of the ischium
of the third maxilliped is broad and expanded (vs.
narrow). While E. pilosa has the intercalated
plates at the base of the telson, which is distinctive
of Amarinus (cf. Lucas, 1980), the differences
noted here require generic separation. This
decision is further validated by a second species
from New Caledonian estuaries (Davie & Richer
de Forges, 1996). Intercalated plates on the d
telson is a character shared by the closely related
Amarinus, Odiomaris, and the monospecific
Australian Hymenosoma Lucas, 1980.
In E. pilosa, the infraorbital tooth is well-
developed and visible in dorsal view, whereas
Amarinus typically has the infraorbital tooth
weak or almost indiscernible, and not visible in
dorsal view. Similarly the dorsal carapace
grooves of E. pilosa are well-defined, with the
posterior ones reaching the edge of the carapace,
and the intestinal region marked by a short but
distinct longitudinal groove. These characters are
not considered to be of generic significance as the
second species of Odiomaris (Davie & Richer de
Forges, 1996), does not have a well developed
infraorbital tooth, and the dorsal carapace
grooves are poorly defined posteriorly, such that
there is no obvious longitudinal groove on the
intestinal region.
ETYMOLOGY. Latin odium, dislike and maris, sea;
for the type species' freshwater habitats. Gender mas-
culine.
272 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 6. Odiomaris pilosus. A, D, F, d (after Milne Edwards, 1873, pl 18, fig. 6). C (after Holthuis, 1968, fig.
3a). B, E, G-J, d (11.8 x 10.1 mm) (ZRC 1994.4251a). A, overall view. B, dorsal view of carapace. C, frontal
view showing epistome. D, front view showing mouthparts. E, right cheliped. F, right chela. G, left third
maxilliped. H, right third ambulatory leg. I, right third ambulatory dactylus. J, right fourth ambulatory leg.
Scales = 1.0 mm.
HYMENOSOMATIDAE OF NEW CALEDONIA
273
FIG. 7. Odiomaris pilosus. A, 3 (after Milne Edwards, 1873, pl. 18, fig. 6). E, 9 (after Milne Edwards, 1873,
pl. 18, fig. 6). B-D, d (11.8 x 10.1 mm) (ZRC 1994.4251a). A, d sternum and abdomen. B, abdomen. C,
D, left Gl. E, 2 abdomen. C, ventral view. D, dorsal view (setae not drawn). Scales = 1.0 mm.
Odiomaris pilosus (Milne Edwards, 1873)
(Figs 5-7)
Elamene pilosa Milne Edwards, 1873: 322, pl. 18 fig.
6; Kemp, 1917: 247.
Elamena pilosa: Tesch, 1918: 21; Roux, 1926: 229,
figs 55, 56.
Halicarcinus pilosus: Holthuis, 1968: 117, fig. 3.
Amarinus pilosus: Lucas, 1980: 198; Chuang & Ng,
1994: 87.
MATERIAL EXAMINED. ZRC 1994.4251, 8 (11.8
x 10.1 mm), 9 (8.8 x 7.8 mm), Boghen River, New
Caledonia, 17.1.1993, B. Richer de Forges.
REMARKS. This material agrees with the
description and figures of Milne Edwards (1873),
Roux (1926) and Holthuis (1968). Roux (1926)
and Holthuis (1968) described the short and very
stiff setae on the carapace as 'spinules/spines',
but in our fresh specimens, they were flexible and
not stiff. Thus, we refer to them as setae.
Our d was dissected and its gonads preserved
in glutaraldehyde for ultramicroscopic study of
the sperm.
DISTRIBUTION AND HABITAT. New Cale-
donia. With Trigonoplax unguiformis this is
largest of known hymenosomatids. It is common
in New Caledonian rivers from the estuary inland
for several km, in shallow freshwaters, under or
between rocks in areas with rapid currents.
Elamena Milne Edwards, 1837
Elamena vesca sp. nov.
(Fig. 8)
Elamene truncata Milne Edwards, 1873: 323 (junior
homonym of Trigonoplax truncata Stimpson,
1858).
? Elamena truncata: Gordon, 1940: 67, fig. 5; McNeill,
1968: 47; Lucas, 1980: 171, figs 2D, 6D, 8B, 10H.
MATERIAL EXAMINED. HOLOTYPE
MNHNB22843, d (6.0 x 5.4 mm), intertidal region,
274 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 8. Elamena vesca sp. nov. A, C-E, G-I, holotype ¢ (6.0 x 5.4 mm) (MNHN). B, F, paratype 9 (7.2 x 6.2
mm) (ZRC 1993.6498). paratype 9 J, K (7.5 x 6.8 mm) (MNHN), New Caledonia. A, B, dorsal view of carapace.
C, left third maxilliped. D, d right cheliped. E, d abdomen. F, 2 abdomen. G, H, left G1. I, G1 apex. J, right
third ambulatory leg. K, right ambulatory leg dactylus. Scales: A-I = 0.5 mm. J, K = 1.0 mm.
HYMENOSOMATIDAE OF NEW CALEDONIA
low tide at OUEMO, Nouméa, New Caledonia,
2.7.1992, B. Richer de Forges. PARATYPES ZRC
1993.6498-6499, ZRC 1994.4285, 4 2, MNHN
B22844, 9 , same data as holotype; MNHN, 2 8,2 9,
QM W,2 6, 2, ZRC 1996.478, 3 3, 9, Nouméa, Anse
Vata, New Caledonia, 18.4.1995, B. Richer de Forges.
DESCRIPTION. d : Carapace width (at posterior
pair of angular lobes) subequal to length; dorsal
surface gently convex, smooth with no distinct or
very poorly defined cervical, thoracic and
gastrocardiac grooves, with dark brown pigmen-
tation sometimes forming 2 eye-like spots;
lobules at anterior lateral angle and at pterygos-
tomian region equally rounded; anterolateral and
lateral angles poorly defined, rounded. Rostrum
truncated with ventral rostral keel partially
visible dorsally. Eyes visible dorsally.
Third maxillipeds cover 3/4 of mouth field
when closed; ischium shorter than merus along
outer lateral edge; dense short setae occupying
entire length of inner lateral edge of ischium;
inner lateral edge of merus lined with dense setae,
longer than that on ischium; inner lateral margins
meeting when closed; palp subequal in length
with merus; exopod longer than merus, with long
setae more sparse than that found on merus lining
the inner lateral edge.
Chelipeds equal, stouter than ambulatory legs;
surfaces smooth; fingers c.1/3 length of inflated
palm; fingers laterally flattened, slightly curved
inwards; cutting edges serrated, with quadran-
gular tooth at proximal portion of dactylus; tips
of fingers sharp, pointed.
Ambulatory legs slender, cross-section subcir-
cular; distinct tooth at distal dorsal edge of merus
and carpus; dactylus laterally flattened, straight
proximally with distal portion more curved with
subterminal tooth; tip sharply hooked; smaller
and less distinct tooth in between tip and subter-
minal tooth; ventral edge of dactylus lined with
row of short setae; carpus shorter than propodus
and merus which is longer than former.
Abdomen triangular, 5-segmented, segments 3
and 4 fused with no distinct suture; all the other
intersegmental sutures distinct; width greatest at
proximal end of fused segment; telson subequal
in length to segment 5; proximal 1/3 length of
lateral edge straight, distal 2/3 progressively con-
cave, sides of telson gently concave and tapering
rapidly to slightly rounded tip.
G1 slender, strongly curved, tapering gradually
to pointed tip; 8 subterminal setae with spinules
spanning 1/2 setal length; 4 subterminal, equally
spaced protrusions on left side; distal portion
slightly curved; middle portion with 90° turn.
275
9 : Carapace at posterior lateral angle, broader
than long; pigmentation much darker and denser
with no distinct spots on dorsal surface of
carapace, carapace at anterior and posterior
angles raised forming slight ridges (in mature 9 ).
Abdomen with no fused segments, all interseg-
mental sutures distinct, covers entire sternum,
reaching base of legs, broader than long, tip con-
cave (in mature 9 ) or slightly pointed (in imma-
ture 9). Cheliped slender, not stouter than
ambulatory legs; fingers spatulate, outer cutting
edges serrated, tip of fingers sharp.
REMARKS. Elamena vesca sp. nov. cannot be
separated easily from E. truncata unless adult ó s
are available. Despite many reports of Elamena
truncata (Stimpson, 1858) in the literature, a clear
description is only available from Ng & Chuang
(1996). For E. truncata, the lateral angles are well
marked and distinct, the posterior lateral angle
being almost tooth-like, whereas in E. vesca, all
the angles are more rounded. The fingers of the
3 chela in E. vesca are also proportionately
longer than those of E. truncata, which has
shorter fingers and a more stocky palm. The
ambulatory legs of E. vesca, especially the dac-
tyli, are also proportionately longer compared to
E. truncata. The d abdomen of E. truncata has a
proportionately longer and more rounded telson,
and the lateral margins of segment 5 are more
strongly concave than in E. vesca. The Gls of E.
vesca and E. truncata differ markedly, with that
of E. vesca being shorter, more strongly bent and
the tip curved upwards (almost straight in EF.
truncata) (Ng & Chuang, 1996).
Milne Edwards (1873) described this New
Caledonian species as Elamene truncata, without
figures. He was apparently not aware of
Stimpson's (1858) description of Trigonoplax
truncata from Japan. Kemp (1917: 273) com-
mented *... That both authors have used the same
specific name is presumably due to a remarkable
coincidence.' The species are in fact different,
and Milne Edwards' (1873) name is a junior
homonym of Stimpson's. The species is here
described as new, using fresh specimens as types.
Australian specimens of E. truncata are tenta-
tively referred to E. vesca (Ng & Chuang, 1996).
The Australian specimens however, differ from
E. vesca in the d abdomen as well as G1. The c
abdomen of ‘E. truncata’ of Gordon (1940: 69,
fig. 5d) and Lucas (1980: 245, fig. 8B) differ from
that of E. vesca (Fig. 7E) in having fused seg-
ments 3 and 4 longer and the telson longer than
segment 5 (subequal in length for E. vesca).
276
Gordon’s (1940) & abdomen has a proportionate-
ly longer segment 5 and fused segments 3 and 4
compared to that of Lucas (1980). Also the lateral
margins of fused segments 3 and 4 are strongly
convex in Gordon’s figure but only gently convex
in Lucas’. The Australian specimens need
taxonomic review but are definitely closer to E.
vesca than E. truncata as defined by Ng &
Chuang (1996).
DISTRIBUTION AND HABITAT. New
Caledonia and possibly Australia. Cryptic, inter-
tidal on rocky shores, in shallow water (about 0.5
m). Our specimens were collected under rocks.
ETYMOLOGY. Latin vesca, weak; alluding to the
appearance of the species.
ACKNOWLEDGEMENTS
We are grateful to Daniéle Guinot (MNHN)
and S. Pinkster (ZMA) for loan of specimens.
John Lucas and Peter Davie kindly read the
manuscript and offered useful suggestions. Chris-
tina Chuang prepared figures 1B-G and 8A-F, H,
I. The study was partially supported by
RP900360 to the first author from the National
University of Singapore.
LITERATURE CITED
CHUANG, C.T.N. & NG, P.K.L. 1994. The ecology
and biology of Southeast Asian false spider crabs
(Crustacea: Decapoda: Brachyura:
Hymenosomatidae). Hydrobiologia 285: 85-92.
DAVIE, P.J.F. & RICHER DE FORGES, B, 1996. Two
new species of spider crabs (Crustacea:
Brachyura: Hymenosomatidae) from New
Caledonia. Memoirs of the Queensland Museum
39(2):257-262.
GORDON, I. 1940. On some species of the genus
Elamena (s.s.) (Crustacea, Decapoda). Proceed-
ings of the Linnean Society of London 152(1):
60-78.
HOLTHUIS, L.B. 1968. On Hymenosomatidae (Crus-
tacea Decapoda Brachyura) from fresh water, with
the description of a new species. Beaufortia
15(195): 109-121.
KEMP, S. 1917. Notes on Crustacea Decapoda in the
Indian Museum. 10. Hymenosomidae. Records of
the Indian Museum 13: 243-279.
LUCAS, J.S., 1980. Spider crabs of the family
Hymenosomatidae (Crustacea; Brachyura) with
particular reference to Australian species: sys-
tematics and Biology. Records of the Australian
Museum 33(4): 148-247.
LUCAS, J.S. & DAVIE, P.J.F. 1982. Hymenosomatid
crabs of Queensland estuaries and tidal mud flats,
MEMOIRS OF THE QUEENSLAND MUSEUM
including descriptions of four new species of
Elamenopsis A. Milne Edwards and a new species
of Amarinus Lucas. Memoirs of the Queensland
Museum 20(3): 401-419.
MCNEILL, F.A. 1968. Crustacea, Decapoda and
Stomatopoda. Scientific Reports of the Great Bar-
rier Reef Expedition 7(1): 1- 98.
MELROSE, M.J. 1975. The marine fauna of New
Zealand: Family Hymenosomatidae (Crustacea,
Decapoda, Brachyura). Memoirs of the New
Zealand Oceanographic Institute 34: 1-123.
MILNE EDWARDS, A. 1873. Recherches surla Faune
Carcinologique de la Nouvelle Caledonie.
Nouvelles Archives du Muséum d'Histoire
naturelle 9: 155-332.
MILNE EDWARDS, H. 1837. Historie Naturelle des
Crustacés, Comprenant l'Anatomie, la
Physiologie et al Classification de ces Animaux.
Volume 2. Libraire Encyclopaedie de Roret, Paris,
458 pp.
NG, P.K.L. & CHUANG, C.T.N. 1996. The
Hymenosomatidae (Crustacea: Decapoda:
Brachyura) of Southeast Asia, with notes on other
species. Raffles Bulletin of Zoology, Supplement
3, in press.
ROUX, J. 1926. Crustacés Décapodes d'eau douce de
la Nouvelle-Calédonie. In F. Sarasin & J. Roux,
Nova Caledonia 4: 181-240.
SAKAI, T., 1938. Studies on the Crabs of Japan. 3.
Brachygnatha, Oxyrhyncha. (Yokendo Co.:
Tokyo). pp. 193-364.
SERENE, R. & UMALI, A.F. 1970. The family
Raninidae and other new species of brachyuran
decapods from the Philippines and adjacent
regions. Philippine Journal of Science 99(1/2):
21-105.
SHEN, C.J. 1932. The brachyuran Crustacea of North
China. Zoologica Sinica (A) 9(1): 1-320.
STIMPSON, W. 1858. Prodomus descriptionis
animalium evertebratorum, quae in Expeditione
ad Oceanum Pacificum Septentrionalem, a
Republica Federate missa, Cadwaladaro Ringgold
et Johanne Rodgers Ducibus, observatit et
descripsit. V. Crustacea Ocypodoidea. Proceed-
ings of the Academy of Natural Sciences of
Philadelphia 10: 93-110.
TAKEDA, M. & MIYAKE, S. 1971. Two new
hymenosomatid crabs of the genus Rhynchoplax
from the West and South Pacific. Researches in
Crustacea 4/5: 1-7.
TAKEDA, M. & NUNOMURA, N. 1976. Crabs col-
lected by the Melanesia Expedition of the Osaka
Museum of Natural History, 1958. Bulletin of the
Osaka Museum of Natural History 30: 61-92.
TESCH, T.T. 1918. The Decapoda Brachyura of the
Siboga Expedition. I. Hymenosomidae,
Retroplumidae, Ocypodidae, Grapsidae, and Ger-
carcinidae. Siboga Expeditie 39c(82): 1-148.
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series | 18: 176-178.
TWO NEW FRESHWATER CRABS IN AUSTRALOCARCINUS DAVIE, WITH
REMARKS ON TROGLOPLACINAE GUINOT AND GONEPLACIDAE MACLEAY
(CRUSTACEA:DECAPODA:BRACHYURA)
P.J.F. DAVIE & D. GUINOT
Davie, P.J.F. & Guinot, D. 1996 07 20: Two new freshwater crabs in Australocarcinus Davie,
with remarks on Trogloplacinae Guinot and Goneplacidae Macleay (Crus-
tacea:Decapoda:Brachyura). Memoirs of the Queensland Museum 39(2); 277-287. Brisbane.
ISSN 0079- 8835.
Two new species, A. kanaka and A. palauensis, of the previously monotypic Australocar-
cinus Davie are described from New Caledonia and Palau Islands, respectively, The three
species are separated on dentition of the anterolateral margins. Direct development of the
young without free-living stages is confirmed for A. kanaka. Australocarcinus is placed in
the previously monotypic almost completely freshwater Trogloplacinae Guinot which is
restricted to the tropical West Pacific. Their apparent closest relatives are marine, mostly
deepwater Chasmocarcininae Seréne (Gonoplacidae). Trogloplax has strong troglobitic
adaptations and is presumed to have evolved from an Australocarcinus-like ancestor. The
Trogloplacinae is separated from the Chasmocarcininae by structure of the antennular region,
and differences in length and shape of the male gonopods. C] Brachyura, Goneplacidae,
Australoplax, Trogloplax, freshwater.
P.J.F. Davie, Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101,
Australia; D. Guinot, Laboratoire de Zoologie (Arthropodes), Muséum national d'Histoire
naturelle, 61 rue de Buffon, Paris cedex 05, France; 15 April 1996.
Trogloplax Guinot, 1986, was erected for a
freshwater troglobitic crab from New Britain. Its
type, T. joliveti, has a remarkable appearance,
blind, and with many adaptive features for a cave
environment. Because of its many peculiar fea-
tures Guinot (1986) erected the Trogloplacinae,
in the Goneplacidae for it.
Australocarcinus Davie, 1988, was established
for a north Queensland estuarine species, A.
riparius Davie, 1988, also peculiar in the com-
bination of morphological characters, in par-
ticular the unusual sternal plate, gonopods, and
putative direct development of the young.
While Trogloplax shared the unusual structure
of male sternite 8, its strong troglobitic adapta-
tions made it difficult to be sure if the two genera
were closely related. Close comparison shows the
2 genera to be related in the Trogloplacinae. We
provide a redefinition of the subfamily and better
understanding of its relationships within the
Goneplacidae.
Two additional species of Australocarcinus are
described herein. One comes from a freshwater
stream in northern New Caledonia and the
second, from Palau, was discovered by P.J.F.D.
amongst an unidentified collection of grapsids on
loan from the USNM to Michael Türkay at the
Senckenberg Museum, Frankfurt. Also a large
population of A. riparius from a freshwater rain-
forest stream in north Queensland is reported.
This discovery, the records from freshwater of the
New Caledonian species and the confirmation of
direct development, show that trogloplacines are
a very old freshwater group, probably derived
from a marine ancestor, with at least A. riparius,
showing estuarine tolerance. Jamieson & Guinot
(1996) examine the ultrastructure of the sper-
matozoan of A. riparius and discuss possible
generic relationships.
Abbreviations used in the text: c.b.=carapace
width; G1,G2=male first and second gonopods;
MNHN, Muséum national d'Histoire naturelle,
Paris; QM, Queensland Museum, Brisbane;
SMF, Senckenberg Museum, Frankfurt; USNM,
United States National Museum, Washington.
Measurements given in the text are of the
carapace breadth (measured at the widest point)
followed by length. Leg segments were measured
in a straight line to give maximum dorsal length,
and so are not always the maximum possible
length.
SYSTEMATICS
Family GONEPLACIDAE MacLeay, 1838
subfamily TROGLOPLACINAE Guinot, 1986
Trogloplacinae Guinot, 1986:307; 1987:25; 1988:22;
1994:167; Guinot & Geoffroy, 1987:18.
278
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. Australocarcinus kanaka sp. nov., 9 paratype, 11.1 x 9.5 mm (QMW21389), dorsal view. Scale line in
mm.
DIAGNOSIS (emended from Guinot, 1986).
Carapace rounded; sometimes poorly calcified;
anterolateral margin cristate, entire or toothed;
front with or without shallow median indentation,
without latero-external notch. Eyes relatively
small. Antennules folded completely into fosset.
Antenna lying in orbital hiatus. Buccal frame
quadrangular; third maxilliped wide, together al-
most completely closing buccal cavity; exopod
broad, with flagellum. Sternal plate very broad,
with all sutures interrupted; large part of sternite
8 exposed; a supplementary transverse suture in
middle of sternite 8, parallel to suture 7/8, form-
ing a supplementary plate. Sterno-abdominal
cavity deep. Male abdomen with segments 3-5
fused. Abdominal locking mechanism in normal
position. Vulvae of female very large, occupying
position near extremity of sternal suture 5/6.
Penis very long, lying in covered channel on
sternite 8, only uncovered next to the coxa of P5;
finally protruding as long soft papilla. Chelipeds
with minor heterochely and heterodonty. Gl
stout, moderately tapering, with an apical aper-
ture; G2 about as long as Gl, with flagellum
about same length as peduncle.
REMARKS. The Trogloplacinae belong to the
Heterotremata showing a coxosternal disposition
of the male sexual opening (Guinot, 1978,
19792).
Australocarcinus Davie, 1988
Australocarcinus Davie,1988:259.
TYPE SPECIES. Australocarcinus riparius Davie,
1988, by original designation.
DIAGNOSIS. Carapace smooth, glabrous,
regions poorly defined; anterolateral margins
convex, with or without rounded teeth,
posterolateral margins subparallel, posterolateral
facet delimited. Frontal margin shallowly
sinuous, formed of 2 rounded lobes, moderately
deflexed, without preorbital lobes or teeth; fron-
to-orbital border c. 0.5-0.6 times carapace width.
Orbits small, unarmed, with slightly raised rim.
Eyestalks short, moveable, with well developed
corneas; completely retractable within orbit.
Chelae robust, similar but one slightly larger.
Legs long, slender, hirsute, second pair longest.
Male abdomen with segments 3-5 fused, segment
3 expanded laterally, subequal in width to seg-
ment one, neither covering sternum between last
pair of legs. Sternal segment 8 in male with a
closed, invaginated channel carrying penis, such
TWO NEW FRESHWATER CRABS IN AUSTRALOCARCINUS
279
FIG. 2. Australocarcinus kanaka sp. nov., 9 paratype, 11.1 x 9.5 mm (QMW21387), showing hatched megalopae
under the abdomen. Scale line in mm.
that appears to be formed of 2 discrete plates;
female of normal form. G1 stout, straight, taper-
ing to simple apex; G2 as long as first, slender,
narrowed in width over distal half, ending in
simple apex.
Australocarcinus riparius Davie, 1988
(Fig. 7)
Australocarcinus riparius Davie, 1988:260, figs 1-3.
MATERIAL EXAMINED. QMW18234, 1268 8
(8.3x6.7-12.9x10.2 mm), 29 9(10.2x8.2, 11.3x
9.5mm), Mclvor River at Isabella-MclIvor road cross-
ing, 15°07.2’S, 145?04.4" E, freshwater, under rocks in
pools in drying river bed, fringing rainforest, DO2 0.9
ppm, altitude c. 30m, 18.11.1992, P. Davie & J. Short.
DESCRIPTION. See Davie (1988).
HABITAT. Freshwater and estuarine. The col-
lection site at the MclIvor River, is within rain-
forest at about 30m altitude, and they were found
under rocks in pools in the drying river bed.
Previous records were from estuaries where it is
apparently common in salinities up to 20 p.p.t.
(Davie, 1988).
DISTRIBUTION. This record extends the dis-
tribution 400km further north along the eastern
Australian coast, from the Murray River, near
Cardwell, and Hinchinbrook Island.
Australocarcinus kanaka sp. nov.
(Figs 1-4, 6, 8)
MATERIAL EXAMINED. HOLOTYPE
MNHNB25279, d (11.1x 9.5 mm), Cold’ Amoss, near
Ouégoa, New Caledonia, in freshwater stream, 100 m
altitude, 13.11.1993, R. Raven. PARATYPES
QMW21387, 9 with megalopae (11.1 x 9.5 mm), data
as for holotype. QMW21388, ¢ (10.5 x 9.0 mm), 29 ?
(12.0 x 10.0; 13.1 x 10.8 mm), data as for holotype.
QMW21389, 9 (11.1 x 9.5 mm), d (6.4 x 5.5 mm),
Col d’Amoss, near Ouégoa, New Caledonia, in fresh-
water stream, 100 m altitude, 13.05.1984, G. Monteith
and D. Cook. QMW20577, d (3.6 x 3.1 mm),
Ouekoula, near Ouémou, upper drainage of the
Ouémou River (draining to the west coast), small trick-
les originating from seeps, 230 m altitude, 13.07.1993,
P. Bouchet. MNHN-B25280, 29 @ (11.2x 9.2; 12.4 x
10.4 mm), Pangou, upper drainage of the Ouaiéme
River (draining to the east coast), in wet mosses in
splash zone of very small stream forming rapids, 300
m altitude, 13.07.1993, P. Bouchet. MNHNB25281, ó
(8.7 x 7.3 mm), Ouéné, near Pangou, upper drainage of
the Ouaiéme River, seepage and small trickle, under
dense secondary vegetation, 400 m altitude,
14.07.1993, P. Bouchet. MNHNB25282, 3d d (3.5 x
3.0; 4.0 x 3.6; 7.5x 6.2 mm), 22 9 (7.2 x 6.3; 8.4 x 7.2
mm), Koumac, 3 Creeks, New Caledonia.
DESCRIPTION. Carapace subrectangular;
greatest width behind exorbital angles; 1.16-1.17
times broader than long. Carapace convex
anteriorly, flat from side to side posteriorly,
slightly convex anteriorly. Regions poorly
defined, cardiac and metagastric regions defined
280 MEMOIRS OF THE QUEENSLAND MUSEUM
1mm
0.2mm
2mm
2mm
utu
ES nn CMBRENINER 9D
FIG. 3. Australocarcinus kanaka sp. nov. A-C, F, 9 paratype, 11.1 x 9.5 mm. D, E, G, d paratype, 6.4 x 5.5 mm
(QMW21389). A, right side of carapace in dorsal view. B, dactylus of third maxilliped showing stout comb-like
setae. C, third maxilliped. D, sternum of male. E, male abdomen. F, right chela. G, sternites 7 and 8 of male
showing fissure to form a supplementary plate.
TWO NEW FRESHWATER CRABS IN AUSTRALOCARCINUS
FIG. 4. Australocarcinus kanaka sp. nov., & holotype,
11.1 x 9.5 mm, QMW20577. A, gonopod 1. B,
gonopod 2. Scale line = 0.5 mm.
by shallow grooves; posterolateral facet
delimited from behind last anterolateral tooth.
Lateral margins subparallel, or slightly divergent
posteriorly; straight. Anterolateral margins
regularly convex; cristate; with two teeth behind
the exorbital angle. Exorbital angle effaced. First
anterolateral tooth blunt, placed about halfway
along anterolateral margin. Second anterolateral
tooth blunt; similar in size to first in small male
but less obvious on holotype. Front moderately
deflexed; with or without shallow median emar-
gination; lateral angles rounded; no pre-orbital
teeth; lateral margins diverging posteriorly; fron-
to-orbital border c. 0.5 times width. Carapace
281
surface evenly punctate, smooth, naked, long
feathered setae restricted to side walls. Upper
orbital border smooth; moderately concave.
Lower orbital border straight, slightly concave at
inner end; smooth. Inner angle of lower orbital
border effaced, reaching about halfway up basal
antennal segment; lower orbital border con-
tinuous with lower edge of outer orbital tooth as
a slightly raised rim. Antennal flagellum entering
orbit c.1.5 times length of orbit. Orbital hiatus
open. Basal antennal segment short, not touching
front; unarmed. Inter-antennular septum narrow.
Third maxilliped with merus and ischium sub-
equal. Merus slightly wider than long; outer mar-
gin straight; antero-external angle not produced,
broadly rounded; about equal in length to is-
chium. Suture between merus and ischium
horizontal. Ischium quadrate; inner margin
smooth. Palp articulating at inner distal margin of
merus; relatively long, dactylus armed with long,
stout, comb-like bristles apically. Exopod
moderately broad.
Chelipeds subequal, right slightly the larger on
holotype; large and robust; borders granulate;
row of longer feathered setae on anterior and
posterior borders; carpus with a broad spine at
inner angle. Outer surface of palm smooth, with
coarse punctations. Outer surface of palm naked.
Inner surface of palm smooth. Immovable finger
moderately long; flattened on outer surface; with
a broad, shallow, longitudinal groove, bearing
close cropped setae; length cutting edge c. 0.4
times length propodus. Ventral border of chela
straight. Dorsal surface of dactyl smooth,
rounded; distal third of dactyl with broad groove
like fixed finger. Fingers pointed, slightly curved
inwards; a narrow gape between cutting margins
proximally.
Walking legs relatively long, compressed,
slender; second pair the longest. Longest leg c.
1.8-1.9 times maximum carapace width. Merus
of third leg c. 3.7-4.25 times as long as wide
(paratype and holotype respectively); carpus c.
2.7-2.8 times as long as wide; propodus c. 2.5-2.7
times as long as wide; dactylus c. 1.5-1.6 times
length of propodus. Dactyli cylindrical, densely
covered in felt of setae; terminating in acute
chitinous tips. Anterior margin of merus with a
subdistal shoulder; unarmed terminally. Leg seg-
ments smooth; fringed with short, feathered,
setae, longer on infero-distal borders of propodi.
Male abdomen moderately broad; 5 free seg-
ments; 3-5 fused; 1 and 3 widest, subequal. First
segment not covering entire width of sternum
between 4th pereiopods; narrow. Segments 3-5
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 5. Australocarcinus palauensis sp. nov., 2 holotype, 7.1 x 5.8 mm (USNM). A. carapace in dorsal view.
B, third maxilliped. C, left chela. D, abdomen. Scale line = 1 mm.
tapering, basally bulbous. Segment 6 c. 2 times
wider than long. Telson longer than penultimate
segment; c. 0.8 times longer than wide; apically
rounded. Male gonopods Fig. 4. Sternum broad.
HABITAT. Freshwater habitats - in a freshwater
stream; in small trickles originating from seeps;
in wet mosses in splash zone of very small stream
forming rapids; and in seepage and small trickles
under dense secondary vegetation. Recorded up
to 400m above sea level.
ETYMOLOGY. A noun in apposition for the native
peoples of the islands of the South West Pacific.
Australocarcinus palauensis sp, nov.
(Figs 5, 8)
MATERIAL EXAMINED. HOLOTYPE USNM, 9
(7.1 x 5.8 mm), Addeido River, in small fast tributary,
Babelthuap Id., Palau Islands, 05.03.1946, D.S. Frey.
DESCRIPTION. Carapace subrectangular;
greatest width behind exorbital angles; 1.22 times
broader than long. Carapace convex anteriorly,
flat from side to side but slightly convex towards
the margins. Regions poorly defined, cardiac and
metagastric regions indistinctly defined;
posterolateral facet not defined. Lateral margins
subparallel; straight. Anterolateral margins
regularly convex; cristate; without teeth. Exorbi-
tal angle effaced. Front moderately deflexed;
bilobed; lateral angles rounded; no pre-orbital
teeth; lateral margins diverging posteriorly; fron-
to-orbital border c, 0.55 times carapace width.
Carapace surface evenly punctate, smooth. Dor-
sal surface naked, long feathered setae on side-
walls. Upper orbital border smooth; moderately
TWO NEW FRESHWATER CRABS IN AUSTRALOCARCINUS
FIG. 6. Australocarcinus kanaka sp. nov., megalopa
(QMW21387). Pleopodal membrane still attached
around base of telson. Scale line 2 0.5 mm.
concave. Lower orbital border shallowly sinuous;
smooth; inner angle effaced, reaching about
halfway up basal antennal segment; continuous
with lower edge of outer orbital tooth as a slightly
raised rim. Antennal flagellum entering orbit;
both flagellums missing. Orbital hiatus open.
Basal antennal segment short, not touching front,
unarmed. Inter-antennular septum narrow.
Third maxilliped with merus and ischium sub-
equal. Merus wider than long; outer margin
slightly convex; antero-external angle not
produced, broadly rounded; c. 0.85 times length
of ischium. Suture between merus and ischium
horizontal. Ischium quadrate; inner margin
smooth. Palp articulating at inner distal margin of
merus; relatively long, dactylus armed with long,
stout, comb-like bristles apically. Exopod
moderately broad.
Chelipeds slightly unequal, right slightly the
larger; large and robust; borders granulate; row
283
of long feathered setae on anterior and posterior
borders; carpus with a broad, blunt spine at inner
angle. Outer surface of palm smooth, with coarse
punctations; naked. Inner surface of palm
smooth. Immovable finger moderately long, flat-
tened on outer surface; without obvious lon-
gitudinal groove. Length cutting edge c.0.4 times
length propodus. Ventral border of chela slightly
concave at base of fixed finger. Dorsal surface of
dactyl smooth, rounded. Fingers pointed; slightly
curved inwards; a narrow gape between cutting
margins proximally, fingers with teeth poorly
differentiated, but large, backwardly directed
molar near base of dactyl.
Walking legs relatively long; compressed;
slender; second pair the longest, c. 1.8 times
maximum carapace width. Merus of third leg c.
3.5 times as long as wide; carpus c. 2.6 times as
long as wide; propodus c. 2.4 times as long as
wide; dactylus c. 1.4 times length of propodus.
Dactyli cylindrical, densely covered in a felt of
setae; terminating in acute chitinous tips.
Anterior margin of merus with subdistal
shoulder; unarmed terminally. Leg segments
smooth; fringed with short, feathered setae,
longer on infero-distal borders of propodi.
HABITAT. From a fast flowing tributary of the
Addeido River suggesting a freshwater habitat.
ETYMOLOGY. For the Palau Islands.
REMARKS ON AUSTRALOCARCINUS
DAVIE
The three known species are very similar but
differ conspicuously by the dentition of the
anterolateral margins. Australocarcinus riparius
has 4 large, prominent blunt teeth; A. kanaka has
only 2 low, hardly projecting, blunt lobes; and A.
palauensis has no anterolateral teeth, but simply
a continuous crest. The telson of A. kanaka is
much shorter than on A. riparius. A. riparius also
differs from the other two species by having the
outer margin of the merus of maxilliped 3 much
more strongly convex.
Davie (1988) speculated that direct develop-
ment occurred in A. riparius because 18 juvenile
crabs were collected with the adult female. This
reproductive strategy has now been confirmed for
A. kanaka as one female (QMW21387) has 17
megalopae still attached to its pleopods (Figs
2,6). The megalopae have no obvious yolk
reserve. Ovigerous females of A. riparius carry
about 70 eggs, so there seems to be significant
attrition during the developmental stages. The
284
FIG. 7. Australocarcinus riparius Davie, 1988 (d, 11.6 mm c.b., QMW 18234),
scanning electron micrograph of the inside of sternite 8, showing invaginated
channel in which the penis lies.
holotype female of A. palauensis also shows large
vulval openings and therefore direct development
can similarly be inferred for this species.
RELATIONSHIP OF AUSTRALOCARCINUS
AND TROGLOPLAX
Many of the most unusual characteristics of
monotypic Trogloplax (Guinot, 1986, 1987)
(type T. joliveti), can be attributed to its caver-
nicolous adaptations. These are common, to a
greater or lesser extent, in all cave dwelling
arthropods. They are: 1) the soft, poorly calcified
carapace; 2) dorsoventral flattening; 3) the
reduced narrow eyestalks with the corneas lack-
ing faceting and pigment (Guinot, 1988, fig. 16A-
C); 4) the loss of pigmentation of the carapace and
pereiopods; 5) exceptionally elongated and thin
pereiopods, especially the ambulatory legs; 6) the
two long spines of the chelipeds. (Guinot,
1986:165).
Despite strong superficial differences there are
fundamental characters that mark a close
relationship between Trogloplax and
Australocarcinus: 1) the carapace lacks setae, is
rounded, and the anterolateral borders are
anteriorly converging and cristate. 2) the buccal
cavity is large and quadrangular. 3) the maxi-
llipeds are short and broad, with the exopods
relatively broad and with a flagellum. 4) the ster-
nal plate is broad with the sutures 4/5 to 7/8
MEMOIRS OF THE QUEENSLAND MUSEUM
interrupted, and with ster-
nite 8 broadly uncovered.
5) the Gl is stout and
straight, and with an apical
opening. 6) the G2 is sub-
equal in length to G1, and
with the two parts (flagel-
lum and peduncle) sube-
qual in length. 7) the male
abdomen has segments 3-
5 fused. 8) the sterno-ab-
dominal cavity is deep in
the male. 9) there is feeble
heterochely and feeble
heterodonty. 10) the an-
tenullar flagellae fold
transversely into fossae
under the frontal margin.
11) a sternal crest sur-
rounds the telson of the
male. 12) the outer face of
sternite 8 is composed of
two plates separated by a
suture which marks an
medial] invaginated canal,
in which the penis lies.
[ Openin g
for penis
REMARKS ON THE GONEPLACIDAE
Guinot (1969a,b,c, 1971) showed that the
Goneplacidae MacLeay, 1838, sensu Balss
(1957) constitutes an heterogeneous assemblage,
and must be re-appraised. Evolutionary lineages
in the family are the goneplacine (-car-
cinoplacine) line, a pilumnine line, the panopeine
line, and the euryplacine line. The Geryonidae
were considered very distant from the
Goneplacidae. The Rhizopinae Stimpson, 1858,
still needs to have some questions resolved. This
subfamily may need to be restricted to the type
species, R. gracilipes Stimpson, 1858, which ap-
pears to constitute a separate lineage, a little
different from the pilumnine line (sensu stricto)
because the male Gl, while similar, is not of
typical pilumnid form with the strongly recurved
tip (Guinot, 1969a, fig. 110), but the G2 is effec-
tively the same as in the Pilumnidae. Ng (1987)
reviewed the constitution of the Rhizopinae, and
it appears that all the genera he included have the
G1 with a pilumnine recurved tip, and the short
sinuous G2 also typical of pilumnids. We think it
possible that the Rhizopinae (sensu Ng, 1987)
may still be heterogeneous with Rhizopa itself
forming a separate lineage. If this is so, then the
name Typhlocarcinopsinae Rathbun, 1909, is
TWO NEW FRESHWATER CRABS IN AUSTRALOCARCINUS
available for Typhlocarcinops and perhaps some
other genera. Typhlocarcinops has gonopds that
are not of the classical pilumnid type (sensu Ng,
1987).
We consider that primitive goneplacid genera
of the pilumnine line cannot be separated
phylogenetically from the xanthoid Pilumnidae
(Pilumnus). In the same way the panopeids Cyr-
toplax Rathbun, 1914, Tetraplax Rathbun, 1901,
Glyptoplax Smith, 1870, and Cycloplax Guinot,
1969, etc., cannot be separated phylogenetically
from the panopeids like Panopeus H. Milne Ed-
wards, 1834, but they form a natural grouping for
which the name Eucratopsinae Stimpson, 1871 (=
Prionoplacinae Alcock, 1900) may be used.
The Goneplacidae contains the Goneplacinae
Miers, 1886, Carcinoplacinae, Miers, 1886,
Chasmocarcininae Seréne, 1964, Trogloplacinae
Guinot, 1986, and the Euryplacinae Stimpson,
1871. Many genera are still not satisfactorily
placed in these subfamilies and a revision is ur-
gently needed.
AFFINITIES OF THE TROGLOPLACINAE.
The most revealing character in helping to under-
stand the affinities of the Trogloplacinae is the
disposition of the penis and gonopore. Typically
in coxosternal crabs (sensu Guinot, 1978, 1979a),
the ejaculatory canal opens on the coxa of P5 and
there is a long penis sometimes visible near the
aperture on the coxa but with most of its length
covered by the complete junction of sternites 7
and 8 and with the distal part long and emerging
to enter the base of the G1 (Guinot, 1979a, fig.
51F; 1979b, figs 2,3). Sternite 8 of the Troglo-
placinae has an invaginated channel in which the
penis lies (Fig. 7).
The Trogloplacinae as Guinot (1986) has al-
ready indicated, belongs to the Goneplacidae and
is most closely related to the Chasmocarcininae
Seréne, 1964 (Chasmocarcinus Rathbun, 1898) a
marine, mostly deepwater group (Seréne,
1964a,b). All Chasmocarcininae (Chasmocar-
cinus, Camatopsis Alcock & Anderson, 1899,
Chasmocarcinops Alcock, 1900, Hephthopelta
Alcock, 1899, Scalopidia Stimpson, 1858) have
the penis lying in either an enclosed or open
groove in sternite 8 (Felder & Rabelais, 1986),
such that sternite 8 has an intercalated plate
anteriorly, a fact first observed for Chasmocar-
cinus by Rathbun (1914).
The type species of Hephthopelta (H. lugubris
Alcock) is a female, and not well known, but a
new species of Hephthopelta we have examined
(being described by Davie & Richer de Forges)
285
has an open suture in sternite 8, such that the penis
is visible. Glaessner & Secretan (1987) and
Tavares (1992) described exactly such a condi-
tion for the fossil American Eocene crab Fal-
conoplax Van Straelen, 1933. This genus had
been attributed to the “Tymolinae’ by Glaessner
(1969)(= Cyclodorippidae), but Tavares (1992)
re-attributed this fossil to the Goneplacidae sensu
lato, agreeing with Van Straelen’s (1933) original
placement. We believe that Falconoplax belongs
to the Chasmocarcininae. The first record of a
fossil Chasmocarcinus was reported from the
Eocene of Antarctica, in shallow water sediments
(Feldmann & Zinsmeister, 1984).
Seréne (1964a), with some reserve, placed
Megaesthesius Rathbun, 1909, in the Chasmocar-
cininae. We exclude it from the subfamily but
acknowledge its relationships must be inves-
tigated further.
In spite of the similarity in the coxosternal plate
between the Trogloplacinae and the Chasmocar-
cininae, there are some notable differences. The
major difference is that in the Chasmocarcininae
the basal antennular segment is very swollen and
completely fills the antennular fosset, such that
the flagellum is excluded and cannot be folded.
The gonopods of the two subfamilies are similar
in that the G1 is stout and tapering and the G2 is
long and not sigmoid, however there are also
differences. In the Chasmocarcininae the G2 is
noticeably shorter than the G1 and the flagellum
is short (length varies a little depending on the
species). In the Trogloplacinae the G2 is as long
or slightly longer than the G1, and the flagellum
occupies about half or slightly more of the length.
Considering the close relationships with the
exclusively marine Chasmocarcininae, and the
presence of at least one estuarine tolerant species
in Australocarcinus, derivation of the completely
freshwater Trogloplax from a marine stock is a
logical assumption.
BIOGEOGRAPHY. The trogloplacines are so far
only known from the western Pacific (Fig. 8).
Trogloplax occurs in two freshwater subter-
ranean caves on New Britain, PNG (Guinot &
Geoffroy, 1987). The two caves are 250 km apart,
and the crabs were found 300m into the caves.
Australocarcinus is now known from three
species in north Queensland, New Caledonia, and
Palau; it seems probable that other species will be
found on the West Pacific Islands. Although little
is known of the reproductive patterns of the
trogloplacines, Davie's (1988) supposition of
direct development for Australocarcinus riparius
286
f fe MERE re @ New Britain
S.) E
1 r x
= =~
Queensland e. à
N New Caledonia
FIG. 8. Distribution of Australocarcinus and Troglo-
plax. Star = A. palauensis sp. nov.; hollow box = A.
kanaka sp. nov.; solid circle — A. riparius Davie,
1988; diamond = Trogloplax joliveti Guinot, 1986.
has been confirmed for A. kanaka. If the other
species of Australocarcinus and Trogloplax also
have direct development, this should reinforce
their allopatric distributions and contribute to
small area speciation. One factor which could
allow genetic interchange over a limited range is
the high salinity tolerance exhibited by A.
riparius, which allows it to survive under es-
tuarine conditions. Estuarine corridors along the
coast between river systems, during flood condi-
tions, could allow dispersal of adults and the
maintenance of the species over long distances,
as occurs on the NE coast of QLD. Such a disper-
sal method however is only likely to be of local
importance. There seems to be no obvious disper-
sal or vicariant explanation that conveniently ex-
plains the observed distribution patterns of the
genera and species of the Trogloplacinae.
ACKNOWLEDGEMENTS
We would like to acknowledge the help of a
number of people. Michael Tiirkay,
Forschungsinstitut Senckenberg, Frankfurt, kind-
ly gave PD access to loan material from the
USNM in his care, amongst which was found one
of the new species. Clare Bremner and Alison
Hill executed Figures 3 and 5 respectively, and
Kylie Stumkat took the SEM photograph.
Bertrand Richer de Forges of ORSTOM,
Noumea, New Caledonia, gave PD hospitality
and stimulating discussion during his visitto New
MEMOIRS OF THE QUEENSLAND MUSEUM
Caledonia, and made valuable comments on the
manuscript. Philippe Bouchet of the Muséum
national d'Histoire naturelle, Paris, collected
many specimens from inaccessible places. Peter
Ng, National University of Singapore, is thanked
for his thorough review of the paper.
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MILNE EDWARDS, H. 1834. Histoire naturelle des
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1914. A new genus and some new species of crabs
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288
FEEDING HABITS OF THE RING-TAILED GECKO,
CYRTODACTYLUS LOUISIADENSIS. Memoirs of the
Queensland Museum 39(2); 288. 1996:- ‘Geckos can be vora-
cious predators and will actively pursue and kill their prey.
Generally they attack .... arthropods ... head size governs the
size of prey. '... (King & Horner, 1993). Notwithstanding the
fact that most Australian geckos are opportunistic arthropod
feeders, some supplement their diets with plant exudates
(Greer, 1989; Bauer, 1990; Ehmann, 1992; Couper et al.,
1995) and a few large species prey also on vertebrates (Bauer
& Sadlier, 1994). The following Australian species are known
to prey on vertebrates: Nephrurus asper (including N. amyae
and N. sheai), N. deleani, N. laevissimus, N. levis, N. milli, N.
stellatus, N. vertebralis, N. wheeleri, Qedura marmorata, O.
ocellata, Phyllurus platurus and Pseudothecadactylus
lindneri (Bauer, 1990). All are relatively large (maximum
SVLs 80-135mm).
Recent observations made near Cooktown, NEQ, show
that another large Australian gecko, Cyrtodactylus
louisiadensis (maximum SVL 130mm, after Couper & Greg-
son, 1994), feeds on vertebrates, In July, 1994 in a house near
Jensen's Crossing, Endeavour R. (15°25’S 145"04'E), a large
(exact measurement not recorded), male C. louisiadensis was
observed stalking, then ‘ambushing’ a frog (K.J. & M. Jago).
The gecko, alerted by the frog's movement, ran quickly down
the wall of the house till it was about .3m from the frog (Litoria
pallida) on the floor. Tt then ‘jumped’ to the floor and ran
quickly a few steps to pounce on the frog which was ingested
rapidly. Predation by C. louisiadensis on L. pallida there is
apparently a fairly regular event, because it was observed (K.J.
& M. Jago) again soon afterwards (September, 1994). On
three subsequent occasions, specimens of L pallida were
placed (K.J. & M. Jago) on the floor about 1 m from resting,
but alert C. louisiadensis, low on the house walls. On all
occasions their movement triggered the same ‘ambush’ and
voracious ingestion response by the geckos, In early Novem-
ber, 1995 on the granite boulders of the Black Trevethan Ra.
(15°40’S, 145?14'E), a specimen of C. louisiadensis (SVL
93.6mm, now QMJ60620) was found (L.R., H.J. & D. Cook)
with a struggling specimen of the gecko, Nactus galgajuga
(SVL 30.8mm, now QMJ61096) in its mouth. The latter had
been seized midbody.
As other species of Cyrrodactylus (C. caspius and C.
cavernicolus, from Turkestan, and Borneo respectively) are
known to prey on vertebrates (Bauer, 1990), itis not surprising
that C. louisiadensis also does so, However, predation on
frogs is unusual amongst Australian geckos, having been
reported only once, by Pseudothecadacrylus linderi (Husband
& Irwan, 1995). The stomach contents of all specimens of C.
louisiadensis in the collection of the Queensland Museum
have been examined. They contain only arthropod remains:
Class Chilopoda, unidentified (from QMJ38198); Class
Arachnida, unidentified (QMJ27083, QMJ53634),
Heteropoda jugulans (QMJ38330), Yiinthi chillagoe
(QMJ45365), Lychas sp. (QMJ38331), Liocheles sp.
(QMJ45365); Class Insecta Order Orthoptera, unidentified
(QMJ24493), Family ?Gryllacrididae (QMJ27256), Family
Stenopelmatidae (QMJ2431, QMJ55367), Family Gryllidae
(QMJ2431, QMJ38197), Order Blattodea Family Blattidae
Methana sp. (QMJ19327), Family Blaberidae Laxta sp.
(QMJ38197, QMJ38198, QMJA5365), Calolampra sp. 1
MEMOIRS OF THE QUEENSLAND MUSEUM
(QMJ60863, QMJ60865), Calolampra sp. 2 (QMJ60328),
Order Lepidoptera, unidentified moth (QMJ38198), Order
Hymenoptera Family Formicidae Oecophylla smaragdina
(queens) (QMJ60869), Family Vespidae (QMJ38197), Order
Coleoptera, unidentified larva (QMJ30062), Family
Tenebrionidae (QMJ 30063), Family Elateridae (QMJ60869),
Family Curculionidae (QMJ30063), nematode presumed to
be a parasite (QMJ48084), unidentified possibly moulted skin
(QMJ32323, QMJ52850). This is an unremarkable sample of
large, mostly nocturnal, arthropods. Two specimens (from
QMJ60863 and QMJ60865 from Peach Ck, MclIlwraith Ra.
13?45'30"S 145819'30"E) are interesting. They contain wings
and other parts of a very large species of the cockroach genus
Calolampra. The species is probably undescribed and is rare
in collections. It would be expected to be an “under)bark’
species which would forage on trunks of trees.Bauer & Sadlier
(1994) discuss the relationship between feeding on vertebrates
by Rhacodactylus auriculatus and the enlarged, caniniform
teeth unique to this species. They conclude that this dentition
may reflect the significance of skinks in the diet of this
species, and special requirements for subjugation and han-
dling of such prey. The teeth of C. louisiadensis are small.
However, the anterior maxillary teeth are ‘caniniform’ and
thus well-suited to capturing and holding large arthropods,
and to grasping struggling, small vertebrates as the oppor-
tunities present.
Literature Cited
Bauer, A.M. 1990. Gekkonid lizards as prey of invertebrates
and predators of vertebrates. Herp Review 21 (4): 83-87.
Bauer, A.M. & Sadlier, R.A. 1994. Diet of the New
Caledonian gecko Rhacodactylus auriculatus
(Squamata, Gekkonidae). Russian Journal of Herpetol-
ogy 1(2): 108-113.
Couper, P.J., Covacevich, J.C. & Wilson, S.K. 1995. Sap
feeding by the Australian gecko Gehyra dubia.
Memoirs of the Queensland Museum 38(2): 396.
Couper, P.J. & Gregson, R.A.M. 1994. Redescription of
Nephrurus asper Gunther, and description of N. amyae
sp. nov. and N. sheai sp. nov. Memoirs of the
Queensland Museum 37(1): 53-67.
Ehmann, H. 1992, Encyclopedia of Australian animals The
National Photographic Index of Australian Wildlife,
The Australian Museum. (Collins, Angus & Robertson:
Sydney). 495pp.
Greer, A.E. 1989, The biology and evolution of Australian
lizards. (Surrey Beatty; Chipping Norton). 264pp.
Husband, G. & Irwin, D. 1995. A previously unrecorded prey
item for the Giant Cave Gecko (Pseudothecadactylus
lindneri). Herpetofauna 25(2): 58-59.
King, M. & Horner, P. 1993. Family Gekkonidae. pp. 221-
233. In Glasby, C.J., Ross, G.J.B. & Beesley, P. (eds).
Fauna of Australia. Volume 2A. Amphibia & Reptilia.
(Australian Government Publishing Service: Canber-
ra). 439pp.
J.A. Covacevich, P.J. Couper, G. Monteith, K. Jago, H.
Janetzki & L. Roberts, Queensland Museum, P.O. Box 3300,
South Brisbane, Queensland 4101, Australia; April 1996.
ULTRASTRUCTURE OF THE SPERMATOZOON OF AUSTRALOCARCINUS RIPARIUS
(CRUSTACEA: BRACHYURA: GONOPLACIDAE: TROGLOPLACINAE)
B.G.M. JAMIESON AND D. GUINOT
Jamieson, B.G.M. & Guinot, D. 1996 07 20: Ultrastructure of the spermatozoon of
Australocarcinus riparius (Crustacea: Brachyura: Goneplacidae: Trogloplacinae). Memoirs
of the Queensland Museum 39(2): 289-296. Brisbane. ISSN 0079-8835.
The spermatozoon ofthe freshwater crab Australocarcinus riparius is heterotreme. However,
absence of a recognizable acrosome ray zone is not a general heterotreme feature, though
constant for thoracotremes. This zone is also unrecognizable in potamoids and in corystoids.
The convex ring at the posterior end of the inner acrosome zone is similar to the xanthid ring,
characteristic of the Xanthidae and Panopeidae, but homology is doubtful. Other xanthid
features such as the accessory opercular ring and opercular overhang are absent from the
sperm of A. riparius and relationship with xanthoids is not supported spermatologically. In
its pointed shape, the perforatorium resembles that of corystoids but there the operculum is
perforate. No special similarities to sperm of potamoids, which similarly have an obligatory
freshwater existence at all stages of the life cycle, are apparent. The sperm of A. riparius
shows no clear affinity with those of any other Heterotremata although confirming its
inclusion in that group. In producing more than one spermatozoon per spermatophore, A.
riparius, having marine relatives, may be less evolved along the path of lecithotrophy than
are the potamoids, lacking marine cofamilials. LlAustralocarcinus riparius, Goneplacidae,
Trogloplacinae, spermatozoon, lecithotrophy.
B.G.M. Jamieson, Zoology Department, The University of Queensland, Queensland 4072,
Australia; D. Guinot, Laboratoire de Zoologie (Arthropodes), Muséum national d'Histoire
naturelle, 6] Rue de Buffon, 75231 Paris, Cedex 05, France; received 21 November 1995.
Davie (1988) described Australocarcinus
riparius for a curious crab collected from a soft
estuarine mud bank of the Murray River, NE
Queensland. It has been found living in fresh-
water in rainforest (Davie & Guinot, 1996). This
cryptic crab, excavating its own burrows, is
remarkable in having direct development (the
ovigerous female has only some 70 large eggs)
and maternal care. Davie (1988) assigned it to the
marine Goneplacidae. Guinot (1986) described
Trogloplax joliveti from caves on New Britain.
This specialised cavernicolous crab, with loss of
pigmentation, thin cuticle, very long legs, and
blind, nevertheless is a goneplacid for which
Guinot (1986) erected the Trogloplacinae. Davie
& Guinot (1996) describe 2 freshwater species of
Australocarcinus (from New Caledonia and
Palau) and show that the Australian epigean
Australocarcinus riparius) and the insular-sub-
terranean hypogean Trogloplax are closely re-
lated in the Trogloplacinae. This, incidentally,
furnishes additional evidence for the ancient
faunal relationship of North Australia and
southern Papua-New Guinea/ New Britain.
The Trogloplacinae have pleopod 2 longer than
pleopod 1, a very large sternal plate, with all the
sutures 4/5 to 7/8 incomplete, and a peculiar plate
on sternite 8 which covers the penis at its origin
from the coxa of pereiopod 5 (P5) until its final
emergence in the middle of the lateral margin of
sternite 8. The disposition of the male pores can
be considered as coxo-sternal (Guinot, 1978,
1979a, b). The Trogloplacinae fall in that part of
Heterotremata possessing a wide thoracic ster-
num. The Trogloplacinae, which have extant
cofamilial marine relatives (Chasmocarciniinae)
are derived from a former marine stock. They
exemplify how true freshwater crabs (e.g. the
potamoids), which lack contemporary marine
cofamilials, might have originated from marine
precursors during the Tertiary.
Classically the diverse Goneplacidae (Guinot,
1969), are intermediate between xanthids (or re-
lated forms) and other crabs such as grapsids.
The sperm of Australocarcinus is the first to be
investigated in a goneplacid and in a heterotreme
with a coxo-sternal disposition of the male genital
pore and is here described as a first step in aiding
resolution of the relationships within the
Goneplacidae.
MATERIALS AND METHODS
COLLECTIONS. The mature male specimen
was collected by Peter Davie and John Short from
freshwater rainforest habitat in the McIvor River
perforatorium
perforatorial tubules
nuclear arm
thickened ring
cytoplasm
MEMOIRS OF THE QUEENSLAND MUSEUM
operculum
subopercular zone
inner acrosome zone
outer acrosome zone
nuclear plasma membrane
FIG. 1. Australocarcinus riparius, Semidiagrammatic sagittal section of a spermatozoon traced from a transmis-
sion electron micrograph.
at Isabella-McIvor road crossing, north
Queensland in November 1992.
HISTOLOGY. The male reproductive material
(both testes including the ducts of the vasa
deferentia) was removed from the fresh crab and
immediately fixed in cold glutaraldehyde for
more than 2 hours at 4°. The gonad tissue was
processed in the Zoology Department, The
University of Queensland, by the fixation proce-
dure (outlined below) for transmission electron
microscopy. This was carried out in a Lynx -el.
Microscopy Tissue Processor (Australian
Biomedical Corporation, Ltd., Mount Waverley,
Victoria, Australia).
Portions of the fixed testis (c.l mm3) were
rinsed in 0.2 M phosphate buffer (pH 7.2) (3
rinses in 15 min), postfixed in phosphate buffered
1% osmium tetroxide for 80 min; similarly
washed in buffer and dehydrated through ascend-
ing concentrations of ethanol (40-100%). After
being infiltrated and embedded in Spurr’s epoxy
resin, thin sections (500-800 A thick) were cut on
a LKB 2128 UM IV microtome with a diamond
knife. Sections were placed on carbon-stabilized
colloidin-coated 200 jum mesh copper grids and
stained (according to Daddow, 1986) in
Reynold's lead citrate for 30 s, rinsed in distilled
water, then 6% aqueous uranyl acetate for 1 min,
lead citrate again for 30 s and a final rinse in
distilled water. Micrographs were taken on an
Hitachi H-300 transmission electron microscope
at 80 kV and a JEOL 100-S transmission electron
microscope at 60 kV.
RESULTS
For a comparative account and explanation of
the various components of the brachyuran sper-
matozoon see Jamieson (1991a, 1991b, 1994)
and the Discussion. The last paper contains a
diagram of these components.
GENERAL MORPHOLOGY (Figs 1-3; Table
1). Each of the many spermatophores in the testes
contains several to many spermatozoa. As such
the spermatophores constitute coenospermia. An
acrosome vesicle forms most of the volume. The
acrosome is concentrically zoned but lacks the
concentric lamellation of thoracotremes; it is
capped apically by a dense operculum and is
ensheathed in a thin cytoplasm which in turn is
embedded in the nucleus. The acrosome vesicle
is centrally penetrated by a cylindrical per-
foratorial column from the posterior end to im-
mediately below the opercular complex. The
nuclear material forms several marginal projec-
SPERM ULTRASTRUCTURE, AUSTRALOCARCINUS RIPARIUS 291
tions. The subspherical acrosome is typical of the
Eubrachyura (Heterotremata + Thoracotremata).
A chromatin-containing posterior median
process of the nucleus, seen in homolids, Ranina
and some majids is absent. The nucleus consists
of uncondensed, fibrous chromatin, and forms a
cup surrounding the acrosome as in all other
brachyurans. A thin layer of cytoplasm which
intervenes between nucleus and acrosome as in
other brachyurans, forms a small mass containing
the centrioles at the posterior end of the per-
foratorial chamber. Cytoplasmic islets lateral to
the acrosome and embedded in the chromatin
(Figs 1, 2A, B, 3F) contain lamellae and bodies
identifiable by homology with other crabs as
degenerating mitochondria, although no cristae
have been observed.
ACROSOME. The subspheroidal core of the
spermatozoon consists of the concentrically
zoned acrosome which is capped by, and in-
cludes, the opercular complex (Figs 1, 2A, 3A).
The acrosome is invested by an acrosomal
membrane underlain by a moderately electron
dense sheath, the capsule. The mean length of the
acrosome, from the apex of the operculum to the
base of the capsule is 3.83 jum (SD=0.10, n=4);
the mean width is 4.29 pm (SD=0.09, n=4). The
acrosomal membrane and capsule are in-
vaginated to cover an elongate subacrosomal or
perforatorial chamber, the contents of which are
the perforatorium (Figs 1, 2A, B, 3A-D). The
anterior tip of this chamber abuts on the posterior
face of the subopercular zone of the opercular
complex. The perforatorium has a pronounced
anterior taper and has the outline, in longitudinal
sagittal section of an elongate triangle with a
rounded base (Figs 1, 2A, 3A). It consists of a
moderately electron dense matrix and is chiefly
remarkable for numerous microtubule-like struc-
tures, perforatorial tubules, which appear to arise
near, if not from, the posterolateral walls of the
perforatorial chamber and extend chiefly
anteriorly (Figs 2A, 3A, D). These tubules are
closely adpressed to each other and form large
bundles in which the tubules, in transverse sec-
tion (Fig. 3C,E), form a quasi-crystalline array,
leaving a narrow central core of the perforatorium
of matrix material.
The central, subacrosomal axis of the acrosome
formed by the perforatorial chamber is sur-
rounded by a moderately electron dense layer, the
inner acrosome zone (Figs 1, 2A, B, 3A-E) which
extends from the subopercular zone at the
anterior end of the acrosome almost to the
posterior end of the acrosome, reaching the thick-
ened ring. The inner acrosome zone tapers
anteriorly, following the outline of the per-
foratorium. Approximately its posterior 1/3, im-
mediately anterior to the thickened ring, is
widened to form a convex ring around the per-
foratorium (Figs 1, 2A, 3D); the inner acrosome
zone, anterior to this ring, narrows so that there is
a deep constriction between it and the ring. In
some micrographs the convex ring appears to be
separate from, although overlapping with, the
anterior inner acrosome zone. It is not possible to
determine whether the convex ring is the
homologue of a xanthid ring. Acrosome ray zone,
typical of heterotreme sperm, absent.
An outer acrosome zone (Figs 1, 2A, B, 3A-E)
surrounds the inner acrosome zone and the base
of the perforatorial chamber, being several times
wider than the inner zone. This outer zone ex-
tends to the convex margin of the acrosome,
being bounded by the capsule. It is uniform in
structure and moderately electron dense, though
paler than the inner acrosome zone, and, like
other heterotreme sperm, does not display the
concentric lamellae which are characteristic of
thoracotreme sperm.
At the anterior pole of the acrosome, as in all
other brachyurans and in paguroids there is a
dense circular, imperforate, cap-like operculum
(Figs 1, 2A, 3B). In longitudinal section it has an
almost flat base and a low triangular anterior
surface. Only a very thin layer of the outer,
anterior surface is strongly electron dense. The
much thicker lower zone is the subopercular
zone. The operculum has a mean width of 2.35
pam (SD=0.06, n=4).
Accessory ring, present lateral to the oper-
culum in xanthoids and, though differently orien-
tated, in thoracotremes, is not present nor is there
an opercular overhang. Periopercular rim absent.
At the posterior pole of the acrosome (Figs 1,
2A, 3A, D, G, H) the capsule is interrupted, as in
all brachyurans, by invagination of the acrosome
membrane and capsule as an orifice which opens
into the columnar subacrosomal chamber. A
thickened ring which is visible on each side of the
subacrosomal invagination in most heterotremes
and many thoracotremes is strongly developed
(Figs 1, 2A, 3A, D). It is more extensive on the
posterior, peripheral aspect of the acrosome than
where it skirts the inside of the base of the sub-
acrosomal chamber.
CYTOPLASM. The cytoplasm of the sperm
forms a thin, scarcely discernible layer of ir-
292 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 2. Australocarcinus riparius, transmission electron micrographs of spermatozoa. A, sagittal section. B,
transverse section. Abbreviations: ce=centriole; cy=cytoplasm; ia=inner acrosome zone; m=putative degenerat-
ing mitochondrion; ms=membranous structure; n=nucleus; na=nuclear arm; npm=nuclear plasma membrane;
oa=outer acrosome zone; o=operculum; p=perforatorium; pt=perforatorial tubules; so=subopercular zone;
tr=thickened ring.
SPERM ULTRASTRUCTURE, AUSTRALOCARCINUS RIPARIUS 293
FIG. 3. Australocarcinus riparius, transmission electron micrographs of spermatozoa. A, sagittal section (LS).
B, transverse section (TS) through the anterior region of the perforatorium. C, TS through the base of the inner
acrosome zone proper, showing the quasi-crystalline array of perforatorial tubules. D, LS through the per-
foratorial invagination, showing the basal origins of the perforatorial tubules. E, TS through the convex ring,
showing the quasi-crystalline array of perforatorial tubules. F, TS, showing a large membranous (lamellar)
structure. G, Oblique section at the base of the perforatorium, showing two mutually perpendicular centrioles.
H, longitudinal section of a centriole. Abbreviations as in Fig. 2.
294
regular thickness, ensheathing the acrosome ex-
cepting its opercular region (Figs 1, 2A, B, 3A,
4A, B). Membrane complexes and putative
degenerating mitochondria which extend as is-
lands far into the nucleus material are presumably
derived from the cytoplasm but continuity of the
large membranous structures of this type (Figs
2B, 3F) with the periacrosomal cytoplasmic layer
has not been demonstrated. Periacrosomal
cytoplasm is continuous with a mass lying at the
posterior pole of the perforatorial chamber and
the material within the posterior perforatorial
chamber may also be regarded a cytoplasm. No
cytoplasm extends into the nuclear arms.
CENTRIOLES. The basal cytoplasm of the per-
foratorial chamber at the level of the posterior end
of the acrosomal capsule contains centrioles (Fig.
3D, G, H). Two centrioles, at right angles (Fig.
3G) are each of normal length (Fig. 3H) and not
elongate as in potamoids.
NUCLEUS. As in other brachyurans, the nuclear
material is located in the lateral arms and in the
cup-shaped structure around both the acrosome
and its cytoplasmic sheath. C.1/3 of the length of
the spermatozoon consists of the nuclear material
which lies posterior to the acrosome (Figs 1, 2A).
A nuclear envelope is not present between the
chromatin and the periacrosomal cytoplasmic
sheath, but some scattered vesicles (Fig. 3G) and
components of the membranous structures pos-
sibly represent residues of the nuclear envelope.
The external, surface of the cell is bounded by
a moderately dense membrane which may repre-
sent fused nuclear and plasma membranes, here
termed the nuclear plasma membrane (Figs 1, 2A,
B, 3A). The general chromatin consists of a dif-
fuse network of electron dense filaments in a pale
matrix as in other brachyurans. In longitudinal
(Fig. 2A) and transverse sections (Fig. 2B), the
nucleus is deeply and irregularly incised by what
are presumed to be the bases of the nuclear arms
but discrete nuclear arms have not been observed.
DISCUSSION
Varuna litterata and Cardisoma carnifex return
to brackish or marine water to breed, and during
larval growth, and therefore might be expected to
differ in fertilization biology from A. riparius,
which has no marine stages though, as a
goneplacid, having extant cofamilial marine rela-
tives. One or other of these species might also
differ from freshwater potamoids which have
MEMOIRS OF THE QUEENSLAND MUSEUM
neither marine stages nor contemporary
cofamilial marine relatives. Sperm morphology
might reflect different fertilization biology while
any unusual similarities between the sperm of
Australocarcinus and potamoids which differed
from those of crabs with marine or estuarine
stages might be related to the internal physiologi-
cal environment of nonmarine crabs. However,
such correlates, if they exist, are not apparent.
It is presumably coincidental that all 5 genera
in Table 1 lack a recognizable acrosome ray zone
as, although absence is an unusual feature for
heterotremes (in the first 3 genera), absence is an
ancestral feature (thoracotreme synapomorphy)
in Varuna and Cardisoma. The acrosome ray
zone is absent in all thoracotremes (a pre-
dominantly marine group), in which (in contrast
with true freshwater crabs) freshwater or ter-
restrial species always have contemporary rela-
tives in the sea. Absence of the acrosome ray zone
in corystoids (pers. obs.), a marine group, as in
thoracotremes, suggests that absence cannot
simply be attributed to a freshwater existence.
Further lack of correlation of absence of an
acrosome ray zone with the mode of life is seen
in Varuna litterata, lacking this zone and
reproducing in estuarine and marine conditions
whereas A. riparius, and the potamoids, also lack-
ing it, reproduce in freshwater. It is difficult to be
certain whether acrosome rays are present or
absent, as in Potamonautes perlatus (Jamieson,
1993).
The convex ring at the posterior end of the inner
acrosome zone in A, riparius is not seen in other
freshwater crabs. It is similar to the xanthid ring,
of the Xanthidae and Panopeidae, but homology
is doubtful. Other xanthid features such as the
accessory opercular ring and the opercular over-
hang are absent from the sperm of A. riparius and
relationship with xanthoids cannot be supported
spermatologically. In its pointed shape, the per-
foratorium resembles that of corystoids but there
the operculum is perforate.
An opercular perforation in Potamon,
Potamonautes (homoplasic with majids) and in
Cardisoma (where it is an ancestral, synapomor-
phic, thoracotreme condition), but not in Varuna,
or the wholly nonmarine Australocarcinus, again
cannot be attributed to a nonmarine habitat.
The thickened ring is exceptionally well
developed in Australocarcinus. Reduction of the
thickened ring occurs in Potamon and
Potamonautes and is extreme in grapsids (though
moderately developed in Varuna) and in Car-
disoma, Although reduction probably relates to
SPERM ULTRASTRUCTURE, AUSTRALOCARCINUS RIPARIUS
295
TABLE 1. Spermatozoal characters of Brachyura as in freshwater (*) and terrestrial crabs (#).
1.Acrosome len
2.Acrosome zonation
3. Operculum
4. Opercular projections into
suboperculum
5. Operculum-capsule continuity
th/width
*Australo-
carcinus
riparius
Goneplacidae
(This stud
*Potamonautes
perlatus
Potamidae
(Jamieson,
1993)
*Potamon
fluviatile
Potamidae
Guinot et al. in
press
concentric
imperforate
absent
discontinuous
0.9
concentric
imperforate
absent
discontinuous
0.8
concentric
perforate
absent
discontinuous
6. Operculum thickness
7. Opercular width
8. Periopercular rim
9. Accessory ope
10. Subopercular protuberance
11. True acrosome ray zone
12. Outer acrosome zone
13. Anterolateral pale zone
14. Flangelike lower zone
15. Xanthid ring
16. Perforatorium
17. Head of perforations
18. Corrugations of perforatorial
chamber
rcular ring
———— V
thin
moderately
wide
absent
absent
absent
not ragged
absent
absent
absent?
preequatorial
noncapitate
absent
moderate
moderately
wide
well developed
absent
absent
presence
uncertain
not ragged
absent
absent
absent
preequatorial
noncapitate
absent
moderate but
thicker
moderately
wide
absent
absent
absent
not ragged
absent
absent
absent
noncapitate
absent
#Cardisoma
carnifex
Gecarcinidae
*Varuna
litterata
Grapsidae
concentric concentric
imperforate but
indented
perf orate with
apical button
absent absent
discontinuous | discontinuous
moderate moderate
not wide extremely wide
absent
absent | — 1 |
absent absent
absent
modified
elongate
absent
absent
preequatorial preequatorial
noncapitate noncapitate
absent absent
19. Lateral arms
several
several
several
several several
20. Lateral arms
21. Centrioles
22. Posterior median process of
nucleus
23. Thickened ring
nuclear only
not elongate
24. Concentric lamellae
25. Capsular chambers
26. Capsular projections
27. Capsular flange
28. Spiral acrosome zone
29. Opercular overhang
peculiarities of the acrosome reaction (Medina,
1992; Medina & Rodriguez, 1992) it is not a
absent
absent
absent
absent
absent
absent
common feature of freshwater crabs.
A. riparius produces large eggs and is suspected
to have maternal care but its spermatophores
contain several to many spermatozoa, the
coenospermial condition. This contrasts with the
well developed
elongate
nuclear only
nuclear only nuclear |
nuclear only
elongate
present not seen
absent
vestigial
absent
vestigial
resent
P
(reduced)
absent
absent
absent
absent
absent
absent
absent
cleistospermia
absent
absent
absent
absent
absent
present?
cleistospermia
lateral vesicle absent
absent absent
absent absent
absent absent
absent absent
absent? resent
coenospermia
production of spermatophores with single sper-
matozoa, cleistospermia, in Potamon and
Potamonautes. This contrast need not invalidate
the hypothesis (Guinot et al., in press) that the
cleistospermial condition is an adaptation to
peculiar features of fertilization biology in
potamoids and, particularly, to the large size and
296
small numbers of eggs, which correlates with
their known lecithotrophic, direct development.
It was considered possible that sperm from
potamoid spermathecae are delivered singly to
the eggs. Production of single-sperm sper-
matophores (cleistospermia) was conjectured to
be a device preventing polyspermy in individuals
in which wastage of the small numbers of large
eggs (and incidentally of spermatozoa) has to be
minimized. Possibly A. riparius, having marine
relatives, is less evolved along the path of
lecithotrophy than the potamoids, lacking marine
cofamilials.
Sperm of A. riparius shows no clear affinity
with those of other Heterotremata although con-
firming its inclusion in that group.
ACKNOWLEDGEMENTS
We thank Peter Davie, Queensland Museum,
for collecting and identifying A. riparius. L. Dad-
dow and D. Scheltinga, Zoology Department,
University of Queensland, are thanked for tech-
nical assistance with electron microscopy. C. C.
Tudge prepared the line drawing. This research
was supported by an Australian Research Council
Grant to BGMJ and by the Muséum National
d'Histoire Naturelle, Paris.
LITERATURE CITED
DADDOW, L. 1986. An abbreviated method of the
double lead stain technique. Journal of Submicro-
scopic Cytology 18: 221-224.
DAVIE, P.J.F. 1988. A new genus and species of
Goneplacidae (Crustacea:Brachyura) from
Queensland, Australia. Memoirs of the
Queensland Museum 25: 259-264.
DAVIE, P.J.F. & GUINOT, D. 1996. Two new fresh-
water crabs in Australocarcinus Davie, with
remarks on the Trogloplacinae Guinot and the
Goneplacidae MacLeay (Crustacea: Decapoda:
Brachyura). Memoirs of the Queensland Museum
39: 277-287.
GUINOT, D. 1969. Recherches préliminaires sur les
groupements naturels chez les Crustacés
Décapodes Brachyoures. 7. Les Goneplacidae.
Bulletin du Muséum National d'Histoire
MEMOIRS OF THE QUEENSLAND MUSEUM
Naturelle, Paris (2) 41: 241-265 ; (2): 507-528 ;
(3): 688-724.
1978. Principes d'une classification évolutive des
crustacés décapodes brachyoures. Bulletin
Biologique de la France et de la Belgique 112:
211-292.
1979a. Données nouvelles sur la morphologie, la
phylogenése et la taxonomie des Crustacés
Décapodes Brachyoures. Mémoires du Muséum
National d'Histoire Naturelle, Paris (A) Zool.
112: 1-354.
1979b. Probléme pratique d'une classification
cladistique des Crustacés Décapodes
Brachyoures. Bulletin de I’ Office National des
Péches. Tunisie, 3: 33-46.
1986. Description d'un Crabe cavernicole aveugle
de Nouvelle-Bretagne (Papouasie-Nouvelle
Guinée), Trogloplax joliveti gen. nov. sp. nov. et
établissement d'une nouvelle sous-famille
Trogloplacinae subfam. nov. Comptes Rendus
Hebdomadaires des Seances de l' Academie des
Sciences, Paris (3), 303: 307-312.
GUINOT, D., JAMIESON, B.G.M. & TUDGE, C.C. in
press. Spermatozoal ultrastructure and relation-
ships of the freshwater crabs Potamon fluviatile
and Potamon ibericum (Crustacea, Brachyura,
Potamidae). Journal of Zoology, London.
JAMIESON, B.G.M. 1991a. Ultrastructure and
phylogeny of crustacean spermatozoa. Memoirs
of the Queensland Museum 31: 109-142.
199]b. Sperm and Phylogeny in the Brachyura
(Crustacea). Pp. 967-972. In B. Baccetti (eds).
‘Comparative Spermatology 20 Years After’
(Raven Press: New York).
1993. Ultrastructure of the spermatozoon of
Potamonautes perlatus sidneyi (Heterotremata,
Brachyura, Crustacea). South African Journal of
Zoology 28: 40-45.
1994, Phylogeny of the Brachyura with particular
reference to the Podotremata: evidence from a
review of spermatozoal ultrastructure.
Philosophical Transactions of the Royal Society
B 345: 373-393.
MEDINA, A. 1992. Structural modifications of sperm
from the fiddler crab Uca tangeri (Decapoda)
during early stages of fertilization. Journal of
Crustacean Biology 12 (4): 610-614.
MEDINA, A. & RODRIGUEZ, A. 1992. Structural
changes in sperm from the fiddler crab, Uca
tangeri (Crustacea, Brachyura), during the
acrosome reaction. Molecular Reproduction and
Development 33: 195-201.
NEW SPECIES AND RECORDS OF ASTIEAE (ARANEAE: SALTICIDAE)
FROM AUSTRALIA AND PAPUA NEW GUINEA
JOANNA GARDZINSKA
Gardzinska, J. 1996 07 20: New species and records of Astieae (Araneae: Salticidae) from
Australia and Papua New Guinea. Memoirs of the Queensland Museum 39(2): 297-305.
Brisbane. ISSN 0079-8835.
Arasia eucalypti sp. nov. and Helpis minitabunda (Koch) are first records of each genus from
Papua New Guinea. Helpis gracilis sp. nov. and Tauala athertonensis sp. nov. are described
from NEQueensland. [ ] Astieae, Arasia, Helpis, Tauala, Australia, Papua New Guinea.
Joanna Gardzinska, Zaklad Zoologii WSR-P, Prusa 12, 08-110 Siedlce, Poland; received 7
December 1995.
Wanless (1988) listed 6 genera of Astieae com-
prising 32 species, to which Zabka (1995) added
monotypic Megaloastia mainae; all are of Aust-
ralian origin. Some genera (Astia, Helpis, Arasia)
are associated with Eucalyptus forests, others
(Sondra, Jacksonoides, Tauala) are rainforest
dwellers. The genera are endemic to Australia
except Helpis which extends to New Zealand. It
is not surprising to find some in Papua New
Guinea, an island with a common geological his-
tory with Australia, especially in its southern part
with similar floristic characteristics (Eucalyptus
savannah) to NE Australia (Zabka, 1990, 1991,
1993).
MATERIAL AND METHODS
Material from PNG is deposited in the
Queensland Museum, Brisbane (QM). Compara-
tive material was borrowed from the Australian
Museum, Sydney (AMS). Measurements are in
millimetres. The drawings were made using a
grid system. Dissected epigynes were digested in
lactic acid. Abbreviations used are:
AEW=anterior eyes width, ag=accessory gland,
AL=abdominal length, CH=cephalothorax
height, CL=cephalothorax length, co=copulatory
opening, CW=cephalothorax width, e=embolus,
EFL=eye field length, fd=fertilisation duct,
id=insemination duct, mk=membranous keel,
PEW=posterior eyes width, s=spermatheca,
sr-seminal reservoir, ta=retrolateral tibial
apophysis, tg=tegulum, tl=tegular lobe.
Order ARANEAE
Family SALTICIDAE
Tribe ASTIEAE
Arasia Simon, 1901
Astia Koch, 1880: 1158. (part)
Arasia Simon, 1901: 432, 436-438; Roewer, 1954:
968; Bonnet, 1955: 632; Brignoli, 1983: 626; Wan-
less, 1988: 81-84, 115-119; Davies & Zabka, 1989:
206, 210.
TYPE SPECIES. Astia mollicoma Koch, 1880, by
original designation.
Arasia eucalypti sp. nov.
(Figs 1A-E, 2A-D)
MATERIAL EXAMINED. PNG: HOLOTYPE,
QMS28896, ő, National Capital District, Waigani,
university campus, under Eucalyptus bark, 15.07.1988,
D. J. Court, M. Zabka. PARATYPES, QMS28897, 9,
same data except 28.06.1988; QMS28808, 5 9,2 d;
QMS30725 2; QMS30726 9, 2 juveniles, same data
except 25.06.88-14.09.88; QMS28899 9, Central
Province, Sinumu Dam., 1.05.88, D.J. Court.
QMS30723, 9, d, 2 juveniles, Port Moresby,
25.06.88, D.J. Court, M. Zabka; QMS30724, 9, d,
Boroko, 14.09.86, D.J. Court.
DESCRIPTION. Compared to A. mollicoma
(Fig. 1F), larger, body lighter coloured (Figs 1A,
2A). Tegulum of the palpal organ more robust,
seminal reservoir curved, membranous keel
accompanying embolus not distinctive (Fig.
1D,E,G,H). Distal part of the insemination ducts
in epigynes shorter and wider apart (Fig. 2D,E).
MALE. Cephalothorax (Fig. 1A,B) wide and
flat, orange beige with narrow darker margin,
covered with scattered stiff brown hairs and white
fine marginal hairs. Eyes surroundings black;
light guanine spots in the centre of the eye field;
eyes fringed by white hairs. Abdomen (Fig. 1A)
pale greyish, with pattern of darker markings and
white guanine spots, with scattered white and
brown hairs. Spinnerets pale grey. Clypeus
orange beige with numerous white hairs.
Chelicerae of similar colour, plurident, with 3
promarginal and 4 retromarginal teeth (Fig. 1C).
298
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. A-E, Arasia eucalypti sp. nov., 3. A, dorsal view. B, cephalothorax, lateral view. C, cheliceral dentition.
D,E, palpal organ, ventral and retrolateral views. F-H, Arasia mollicoma (Koch, 1880), d . F, dorsal view. G,H,
pedipalp in ventral and retrolateral views. (G,H from Davies & Zabka, 1989).
Maxillae and labium orange with lighter tips.
Sternum pale yellow with darker margins. Venter
yellow-grey. Legs yellowish-orange or pale yel-
low, anteriors darker, clothed in white and am-
bery hairs, spines moderately strong; tibia I with
6 prolateral and 6 retrolateral spines, metatarsus
I with 3 prolateral and 3 retrolateral spines.
Pedipalps yellow with long white and ambery
hairs, their structure shown in Fig. 1D,E.
Dimensions. CL 2.70, CW 2.50, CH 0.80, EFL
1.10, AEW 1.60, PEW 1.40, AL 3.90.
FEMALE. Cephalothorax (Fig. 2A) similar in
colour and shape to that in the 6, however, ab-
domen little darker. Chelicerae robust with 2 or 3
promarginal and 5 retromarginal teeth (Fig. 2B).
Legs yellow-orange, darker on tarsi and metatar-
si; tibia I with 5 prolateral and 5 or 6 retrolateral
spines, metatarsus I with 3 or 4 prolateral and 4
retrolateral spines. Palps yellow with darker tips,
clothed in long whitish hairs. Epigyne (Fig.
2C,D) is similar to that in A. mollicoma (see
arrows on Fig. 2E), except for different course of
NEW ASTIEAE, NEQ AND PNG 299
FIG. 2. A-D, Arasia eucalypti sp. nov., 9 . A, dorsal view. B, cheliceral teeth. C, epigyne. D, internal structures.
E, Arasia mollicoma, internal genitalia (E from Davies & Zabka, 1989).
insemination ducts (Proszynski, 1984, 1987; Helpis Simon, 1901: 432, 436-438; Waterhouse, 1912:
Davies & Zabka, 1989). 125; Neave, 1939:600; Roewer, 1954: 969; Bonnet,
Dimensions. CL 3.60, CW 3.30, CH 1.30, EFL 1957: 2150; Brignoli, 1983: 627; Wanless, 1988:
1.40, AEW 2.00, PEW 1.90, AL 6.00. 81-84, 94-102; Davies & Zabka, 1989: 206, 212.
TYPE SPECIES. Astia minitabunda Koch, by original
DISTRIBUTION. Southern PNG, on Eucalyptus designation.
tree trunks.
Helpis minitabunda (Koch, 1880)
Helpis Simon, 1901 (Figs 3,4,5F)
Astia Koch, 1880: 1160.(part) Astia minitabunda Koch, 1880: 1160.
300
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 3. Helpis minitabunda (Koch, 1880), d. A, dorsal view. B, cephalothorax, lateral view. C, cheliceral teeth.
D,E, pedipalp, ventral and retrolateral views.
Helpis minitabunda Simon, 1901: 431, 432, 436, 438;
Rainbow, 1911: 280; Roewer, 1954: 969; Bonnet,
1957: 2151; Proszynski, 1971: 417; Wanless, 1988;
94-98; Davies & Zabka, 1989: 206, 212.
MATERIAL EXAMINED. PNG: QMS28900, 30728,
3, 9, Mt. Hagen, Hagen Park Hotel, under Eucalyptus
bark, 4.07.1986, D.J. Court; QMS28901, d , Calanthe,
Goroke, Norman Cruttwell's residence, 11.04.87, D.J.
Court.
DESCRIPTION. Male. Cephalothorax (Fig.
3A,B) moderately low, pale brown-orange,
darker marginally with orange guanine spots in
the centre of eye field and scattered whitish and
pale brown hairs. Surroundings of eyes black
with fine white hairs. Abdomen (Fig. 3A) grey
with pattern of white guanine spots and grey-
brown markings. Spinnerets grey. Clypeus,
chelicerae (Fig. 3C) and maxillae and labium
orange-brown. Sternum pale orange-brown with
darker margins. Venter light grey with light spots.
First legs orange with brown markings on femora,
tibiae and patellae; other legs paler; tibia I with 3
prolateral and 3 retrolateral spines, metatarsus
with 2 prolateral and 2 retrolateral ones. Palpal
organ (Figs 3D,E).
Dimensions. CL 2.70, CW 2.10, CH 1.40, EFL
1.40, AEW 1.80, PEW 1.60, AL 3.70.
FEMALE. Cephalothorax (Fig. 4A) yellow-
brown, paler in the centre; eye region with yellow
guanine spots. Surroundings of lateral eyes I, II,
and III black with white hairs. Abdomen pale
grey with brown and white spots. Clypeus orange
brown with rows of white hairs. Chelicerae of
similar colour with 5 promarginal and 6 retromar-
NEW ASTIEAE, NEQ AND PNG
301
FIG. 4. Helpis minitabunda (Koch, 1880), 9 . A, dorsal view. B, cheliceral teeth. C, epigyne. D, internal structures.
ginal teeth (Fig. 4B). Maxillae and labium as in
the d, Sternum orange-brown. Venter light grey.
Legs similar in colour and spination to those in
the c . Pedipalps generally yellow, darker distal-
ly, clothed in white hairs. Epigyne (Fig. 4C,D).
Dimensions. CL 3.80, CW 3.30, CH 1.50, EFL
1.50, AEW 2.20, PEW 2.00, AL 7.20.
COMPARISON. H. minitabunda differs from
other species by the shape of embolus and tegular
lobe in the ó palpal organ (Figs 3D,E, 5D,E, G).
Accessory glands of the epigyne more distant
from the spermathecae (Figs 4D, 6D,E).
DISTRIBUTION. NSW, QLD, TAS, NZ, PNG.
Helpis gracilis sp. nov.
(Figs 5A-E, 6A-D)
MATERIAL EXAMINED. Holotype: d, New South
Wales, Morton National Park, SW Nowra, Sally Creek
Valley, under bark of snow gums, 23.04.1988, M.
Zabka. Allotype: 9 , same data. Paratype 9 , same data.
All AMS.
DIAGNOSIS. Shape of embolus and tegular lobe
distinctive within the genus (Fig. 5D-G). In com-
parison to H. minitabunda the accessory glands
more distant from the spermathecae (Figs
4D,6D).
MALE. Cephalothorax (Fig. 5A,B) chestnut-
brown with paler median stripe, eye field dark
orange. Surroundings of eyes black, covered with
brown hairs. Abdomen (Fig. 5A) pale grey,
darker laterally, covered with brown hairs. Spin-
nerets grey. Clypeus orange-brown, fringed by
white hairs. Chelicerae relatively long, pale
brown, with 5 promarginal and 6 retromarginal
teeth (Fig. 5C). Maxillae and labium brown with
paler tips, clothed in brown hairs. Sternum orange
with narrow darker margins. Venter dark grey.
Legs: anteriors orange-brown with brown mark-
302
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 5. A-E, Helpis gracilis sp. nov., 3. A, dorsal view. B, cephalothorax, lateral view. C, cheliceral teeth. D,E,
pedipalp, ventral and retrolateral views. F, Helpis minitabunda (Koch, 1880), pedipalp, ventral view. G, Helpis
occidentalis (Simon, 1901), pedipalp, ventral view, (F from Davies & Zabka, 1989; G from Wanless, 1988).
ings, especially around the spines, others yellow
with brown markings; tibia I with 3 prolateral and
3 retrolateral spines, metatarsus I with 2
prolateral and 2 retrolateral ones. Pedipalps
brown-beige with dense, rather long hairs on
basal segments (Fig. 5D,E).
Dimensions, CL 3.10, CW 2.75, CH 2.30, EFL
1.40, AEW 2.15, PEW 1.85, AL 4.50.
Female. Cephalothorax (Fig. 6A) orange-
beige-brown clothed in short white and brown
hairs, with orange spots on the eye field. Sur-
roundings of eyes black. Abdomen pale yellow-
grey with pattern of darker spots, covered with
brown hairs. Spinnerets pale yellow-grey.
Clypeus pale brown with white long hairs.
Chelicerae of similar colour, with 4 teeth on
NEW ASTIEAE, NEQ AND PNG
303
FIG. 6.A-D, Helpis gracilis sp. nov., 9. A, dorsal view. B, cheliceral teeth. C, epigyne. D, internal structures.
E, Helpis occidentalis (Simon, 1901), internal genitalia.
promargin and 6 on retromargin (Fig. 6B). Max-
illae and labium pale brown with pale yellow
tips. Sternum light yellow with brown margins.
Venter pale yellow-grey, centrally darker. Legs:
first pair pale brown-orange with some darker
spots, others yellow with brown markings;
spination of leg I as in the c. Pedipalps yellow,
clothed in white hairs. Epigyne (Fig. 6C,D) very
similar to H. occidentalis (Fig. 6E).
Dimensions. CL 3.40, CW 2.70, CH 1.60, EFL
1.50, AEW 2.20, PEW 1.95, AL 4.65.
DISTRIBUTION. Morton National Park, New
South Wales.
Tauala Wanless, 1988
Tauala Wanless, 1988:81-84, 120-133; Davies &
Zabka, 1989: 206, 209.
TYPE SPECIES. Tauala lepidus Wanless, 1988, by
original designation.
Tauala athertonensis sp. nov.
(Fig. 7A-E)
MATERIAL EXAMINED. NE Queensland: Holotype
QMS28903, 9, near Atherton, dry Eucalyptus forest,
on grass, 10.11.1987, M. Zabka.
DIAGNOSIS. Course of insemination ducts dis-
tinctive (Fig. 7E,F).
DESCRIPTION. Female. Cephalothorax (Fig.
7A,B) beige-brown with scattered short brown
hairs and black surroundings of eyes. Abdomen
pale grey with pattern of darker spots, clothed in
ambery hairs. Spinnerets pale grey. Clypeus
orange, edged in short hairs. Chelicerae dark
orange with 3 teeth on promargin and 7 on
retromargin (Fig. 7C). Maxillae orange-brown
304
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 7. A-E, Tauala athertonensis sp. nov., 9. A, dorsal view. B, cephalothorax, lateral view. C, cheliceral teeth.
D, epigyne. E, internal structures. F, Tauala alveolatus (Wanless, 1988), internal genitalia.
with paler inner margins, labium similar, with
paler tips. Sternum yellow with darker margins.
Legs yellow with brown markings on femora,
tibiae and patellae; tarsi and metatarsi darker;
tibia I with 4 prolateral and 3 retrolateral spines,
metatarsus I with 2 prolateral and 2 retrolateral
ones. Pedipalps pale yellow. Epigyne (Fig. 7D,E)
with long, narrow insemination ducts, oval
spermathecae accompanied by the accessory
glands. Diagnostic characters marked with ar-
rows,
Dimensions. CL 1.90, CW 1.50, CH 0.90, EFL
0.90, AEW 1.50, PEW 1.20, AL 2.50.
DISTRIBUTION. Near Atherton, NEQ.
ACKNOWLEDGEMENTS
I am grateful to D.J. Court (Boroko, Singapore)
for allowing me to study his PNG collection.
Mark Harvey, J. Proszynski and M. Zabka
provided critical comments on the typescript.
LITERATURE CITED
DAVIES, V. TODD & ZABKA, M.1989. Illustrated
keys to genera of jumping spiders (Araneae: Sal-
ticidae) in Australia. Memoirs of the Queensland
Museum 27(2): 189-266.
PROSZYNSKI, J.1984. Diagnostic drawings of less
known Salticidae (Araneae) - an atlas. Zeszyty
Naukowe Wyzszej Szkoty Rolniczo-
Pedagogiczney w Siedlcach I: 1-177.
1987. Diagnostic drawings of less known Salticidae
NEW ASTIEAE, NEQ AND PNG 305
(Araneae) 2 - anatlas. Zeszyty Naukowe Wyzszej
Szkoty Rolniczo-Pedagogiczney w Siedlcach 2:
1-172.
WANLESS, F. R. 1988. A revision of the spider group
Astieae (Araneae: Salticidae) in the Australian
region. New Zealand Journal of Zoology 15: 81-
172.
ZABKA, M. 1990. Remarks on Salticidae (Araneae) of
Australia. Acta Zoologica Fennica 190: 415-418.
1991. Studium taksonomiczno-zoogeograficzne
nad Salticidae (Arachnida:Araneae) Australii.
Zeszyty Naukowe Wyzszej Szkoty Rolniczo-
Pedagogiczney w Siedlcach. Rozprawa naukowa
32.
1993. Salticidae (Arachnida: Araneae) of New
Guinea - a zoogeographic account. Bollettino
dell' Accademia Gioenia di Scienze Naturali 26:
389-394.
1995. Salticidae (Arachnida:Araneae) of oriental,
Australian and Pacific regions. 11. A new genus
of Astieae from Western Australia. Records of
the Western Australian Museum, Supplement 52:
159-164.
306
REPRODUCTIVE CHARACTERISTICS OF FEMALE
FROGS FROM MESIC HABITATS IN QUEENSLAND.
Memoirs of the Queensland Museum 39(2): 306. 1996:-
Fecundity has important implications for the life history
strategies and population dynamics of all animals, yet few data
have been published on the fecundity of Australian frogs
(Tyler 1989). Fecundity is herein defined in the strict sense of
the word: ‘the number of eggs produced by an individual’
(Lawrence, 1989). The primary objective of this paper is to
present new information on the reproductive characteristics
of mesic frogs from eastern Queensland, some of which are
now missing or their populations have declined (Ingram &
McDonald, 1993; Richards et al., 1993; Hero, 1996). This
paper is a supplement to the existing papers relating to fecun-
dity of Australian frogs (reviewed by Tyler, 1989). We present
egg counts and mean egg sizes (10 eggs measured / female)
from clutches either dissected from museum specimens or
layed in captivity. In museum specimens, half of the gonad
was removed and the number counted doubled to give the
total. Female body size is also provided because it may be an
important determinant of fecundity. Specimens were dis-
sected at the Queensland Museum in March 1996 and the
museum registration numbers are provided where available.
The results are presented in Table 1. The amphibians
exhibit a wide variety of egg numbers and egg diameters. A
cursory examination of the data suggests a positive associa-
tion between body size and egg numbers, however, there is no
obvious phylogenetic influence. A thorough examination of
these relationships, using a complete data set, is currently in
progress.
Thanks to J. Covacevich and P. Couper, Qld Museum for
providing space, access to the specimens, and good company
during our time at the museum. Richard Retallick and Chris
Reid assisted in the preparation of this manuscript.
Literature Cited
Hero, J.M. 1996. Where are Queensland's missing frogs ?
Wildlife Australia Magazine. in press, June II 1996.
Ingram, G.J. & McDonald, K.R. 1993. An update on the
declining of Queensland's frogs. Herpetology in
Australia. Eds. D. Lunney & D. Ayers. Transactions of
the royal Society of New south Wales 297-303.
Lawrence, E. 1989. Henderson's dictionary of biological
terms. (Longman Scientific & Technical: Hong Kong).
645p.
Richards, S.J., McDonald, K.R. & Alford, R.A.. 1993.
Declines in populations of Australia's endemic tropical
rainforest frogs. Pacific Conservation Biology 1:66-76.
Tyler, M.J. 1989. Australian Frogs. ( Viking O'Neil: Mel-
bourne). 220p.
Jean-Marc Hero & Sheree Fickling, Wet Tropics Manage-
ment Authority; Cooperative Research Centre for Tropical
Rainforest Ecology and Management; Zoology Department
James Cook University, Townsville, Queensland 4811; 6 May
1996
MEMOIRS OF THE QUEENSLAND MUSEUM
Table 1: Reproductive characteristics of female frogs
from mesic habitats in Qld. Mean egg size is from a
sample of 10 eggs / female (* indicates n=5 eggs).
Measurements in mm. MED=Mean egg diameter
9 SVL |SOURCE
L. infrafrenata
L. lesueuri 1612 61731
x 2240 1.5 64.15 |61730
L. lorica «150 36092
L. nannotis 216 1.98 55.5 41298
M 136 2.93 51.75 |41301
st 160 - 51.05 30901
L. nyakalensis | |86 2.5 32.9 55596
pa — moe
s 90 1.9 37.35 55592
—
L. revelata 787 - 41.6 rs. obs.
aE LO (pers. ops. |
L. rheocola 63 2.6 37.1 32105
5 46 2.4 35.0 25153
L. xanthomera |1598 1.5 54.1 43160
“ 1454 1.4 54.2 36006
Myobatrachidae
Adelotus brevis |216 33.75 |28280
I $ eim
"i 214 1,5* 35.35 |27774
Mixophyes 904 25 82.6 29256
fasciolata
22951
43855
4184
422
108.4
84.6
M. iteratus
M. schevilli
Ranidae
Rana daemeli —|.|2372 84.0 26211
A NEW GENUS AND SPECIES OF ANT-ASSOCIATED COCCID (HEMIPTERA:
COCCIDAE: MYZOLECANIINAE) FROM CANTHIUM LAM. (RUBIACEAE)
PENNY J. GULLAN AND AIMORN C. STEWART
Gullan, P.J. & Stewart, A.C. 1996 07 20: A new genus and species of ant-associated coccid
(Hemiptera: Coccidae: Myzolecaniinae) from Canthium Lam. (Rubiaceae). Memoirs of the
Queensland Museum 39(2): 307-314. Brisbane. ISSN 0079-8835.
The adult female and first-instar nymph of Torarchus endocanthium gen. et sp. nov.
(Hemiptera: Coccidae: Myzolecaniinae) from Queensland, is described. This coccid is
known only from inside hollow, swollen stems (ant domatia) of plants of the genus Canthium
Lam. (Rubiaceae), where it lives as a trophobiont in the nests of ants of a Podomyrma species
(Formicidae: Myrmicinae). The first-instar nymph has typical coccid features, but the adult
female is unusual, being distinguished within the Coccidae by its multilocular disc-pores and
microducts resembling bilocular pores on the dorsum, and rounded, projecting anal plates
on the anterior edge of a setose bulge. [ ] Coccidae, Formicidae, Rubiaceae, ant-plant
association.
Penny J. Gullan & Aimorn C. Stewart, Division of Botany & Zoology, The Australian
National University, Canberra, ACT 0200, Australia; received 15 December 1995.
A new ant-plant association (Monteith, 1989,
1990) from marginal rainforest areas of coastal to
central Queensland involves trees of the Can-
thium odoratum-C. buxifolium complex
(Rubiaceae) and ants of an undescribed
Podomyrma species (Formicidae: Myrmicinae).
The ant colonies occur within specialised swell-
ings in the living stems (ant domatia) (Fig. 1),
which the ants hollow out by perforating the wall
and removing the pith. The ants never occur away
from their host trees; likewise, the trees are rarely
found without these ants. Monteith and co-
worker Paul Flower established that the ants ob-
tained their nutrition largely from scale insects,
also called coccoids (Hemiptera: Coccoidea),
living within the ant domatia (Fig. 1). The feeding
stylets of these coccoids presumably tap the
phloem of the host plant and the coccoids’ ex-
creta, called honeydew, is consumed by the ants.
It is not known whether the ants ever eat the
coccoids.
These coccoids of Canthium belong to two
families - the Pseudococcidae (mealybugs) and
the Coccidae (soft scale insects or coccids). The
mealybugs are Pseudococcus longispinus (Tar-
gioni Tozzetti) and two undescribed, closely re-
lated species, probably belonging to Crisicoccus
Ferris or Paracoccus Ezzat & McConnell. The
undescribed mealybug species might be specific
to the Podomyrma-Canthium association al-
though they belong to a group with no known
association with ants, whereas P. longispinus is
cosmopolitan, polyphagous, and one of the most
pestiferous mealybugs in Australia (Williams,
1985; Williams & Watson, 1988). The soft scale
insects belong to an undescribed species of Coc-
cidae with unusual morphology, suggestive of an
obligate relationship with the ants. Individual
adult coccids have their convex venter closely
fitted into pits gnawed by the ants into the inner
surface of the domatium. Their dorsum is ridged
and covered in setae, and their anal area appears
modifed to facilitate honeydew removal by ants.
The eversible anal tube is strongly developed and
surrounded by a pair of rounded, dorsally project-
ing anal plates so that the whole complex forms
a prominent mound. These coccids have been
collected only from ant chambers inside the stems
of Canthium odoratum (Forster f.) Seemann and
may be dependent on this ant-plant association,
FIG. 1. Sectioned hollow stem of Canthium odoratum
showing ant workers and larvae (Podomyrma sp.) on
the right and an adult female of Torarchus endocan-
thium on the left.
308
although they are not found in all plants that
house colonies of Podomyrma. To date, the coc-
cids have been collected only in SEQ coastal
areas and one more inland site, although the ant-
plant association is more widespread, Where coc-
cids are absent, the domatia house mealybugs,
often in large numbers and sometimes of more
than one of the above species. In older stems, the
mealybugs mostly reside in the ant-gnawed pits
but may feed anywhere within younger stems.
Sometimes domatia house both coccids and
mealybugs.
Scale insects have been reported previously
from inside the swollen stems of Rubiaceae in
Africa (Bequaert, 1922). Those collected with
Crematogaster laurenti Forel (Formicidae: Myr-
micinae) inside hollow stems of Psydrax subcor-
data (DC.) Bridson (formerly Plectronia
laurentii De Wild.) from Zaire were described as
Hemilecanium recurvatum by Newstead (1910).
This species is morphologically very different
from the coccid from Canthium in Queensland
(types of H. recurvatum (BMNH) have been ex-
amined) and belongs to a different subfamily. In
Australia, no coccids or mealybugs have been
reported previously from Canthium, although
species of Myzolecanium Beccari (formerly
placed in Cryptostigma Ferris; see Qin and Gul-
lan, 1989; Gullan, Buckley & Ward, 1993) and
Alecanopsis Cockerell (Green, 1924) have been
described from ant nests in living hollow stems
of other plants. Alecanopsis and Myzolecanium
belong to the same subfamily as the coccids from
Canthium but are very different morphologically.
This paper describes the coccid from Can-
thium. Features of the adult female place it in the
Myzolecaniinae Hodgson, 1994. It is atypical for
a coccid in having both multilocular disc-pores
and microducts resembling bilocular pores on the
dorsum, and in having anal plates which are very
rounded and dorsally projecting, together form-
ing a mound when the anal tube is retracted (i.e.
when the plates are closed). In most other coccids,
each anal plate is triangular, posteriorly directed
and lies level with the dorsal surface when the
anus is retracted. Furthermore, in this new species
the anal area is located on the anterior edge of a
prominent bulge which bears very long setae in a
central depression. The first-instar nymph is more
typically coccid-like (Miller, 1991), except that
each stigmatic cleft has only a single stout stig-
matic seta. It differs in this regard from known
first-instar nymphs of other Myzolecaniinae (Ray
& Williams, 1980; Qin & Gullan, 1989; Sheffer
& Williams, 1990).
MEMOIRS OF THE QUEENSLAND MUSEUM
METHODS AND ABBREVIATIONS
Terminology follows Hodgson (1994, 1995).
To prepare adult females and nymphs as micro-
scope slide-mounts, body contents were cleared
in cold 10% w/v potassium hydroxide (KOH)
solution overnight, the cuticle was stained in acid
fuchsin in acid alcohol, dehydrated in 3 changes
of absolute ethanol and 1 of absolute propan-2-ol
and then placed in 3 changes of xylene prior to
mounting in Canada balsam. Scale insects were
prepared for scanning electron microscopy
(SEM) after preservation and storage in 80%
ethanol. Each specimen was dehydrated in a
graded ethanol series, dewaxed in xylene,
rehydrated through a graded ethanol series into
distilled water, post-fixed in 1% aqueous osmium
tetroxide, washed in distilled water and sonicated
briefly to remove any black precipitate, critical
point dried, glued onto a metal stub with nail
varnish and coated with gold palladium under
vacuum. Specimens were then examined and
photographed using a Cambridge S360 SEM.
Abbreviations used for the depositories are:
ANIC, Australian National Insect Collection,
CSIRO, Canberra; BMNH, The Natural History
Museum, London; QM, Queensland Museum,
Brisbane. Each listed scale insect is mounted on
a separate microscope slide, unless otherwise
specified.
SYSTEMATICS
Torarchus gen. nov.
TYPE SPECIES. Torarchus endocanthium sp. nov.
DIAGNOSIS. Adult female with rounded-elon-
gate elevations on dorsum in a definite arrange-
ment; dorsal cuticle with both multilocular
disc-pores and microducts resembling bilocular
pores; anal plates rounded and projecting,
situated on anterior edge of raised area of cuticle.
First-instar nymph of typical coccid form, but
with single stout seta per stigmatic cleft. All
stages living in hollow stems of host plant at-
tended by ants.
DESCRIPTION. Features believed to be
taxonomically significant at generic level are
highlighted. The species description provides the
best summary of this monotypic taxon.
Adult female broader than long, with a row of
rounded-elongate elevations or ridges dorsally on
each side of midline, these become less apparent
after slide-mounting; setae flagellate, clustered
NEW ANT-ASSOCIATED COCCID IN CANTHIUM
309
FIG. 2. SEMs of cuticular features of Torarchus endocanthium: A, anal area of adult 9 , showing rounded anal
plates and long setae on raised area behind anus (scale 50 jum); B, enlargement of anal area of another adult 9 ,
showing anal plates and partially everted anus (scale 50 jum); C, multilocular disc-pores and openings of
microducts on dorsolateral surface of adult 9 (scale 10 jum); D, dorsal view of anal area of first-instar nymph,
showing anal plates and rugose cuticle (scale 25 jum).
dorsally on ridges, in irregular double row mar-
ginally and sparsely scattered ventrally, but with
a group of very long (1.5-2.5 times length of anal
plates), robust setae posterior to anal plates on
raised area; dorsal pores of 3 kinds: 1, disc-pores
with 4-8 loculi, 2, microducts which resemble
bilocular pores when viewed end-on in light
microscope (Fig. 3h) and 3, simple pores; dorsal
tubular ducts and dorsal tubercles absent; anal
plates projecting and lobe-like with cluster of
apical setae; spiracles much larger than legs;
spiracular disc-pores present in bands between
body margin and each spiracle, with 5 loculi;
pregenital disc-pores with 5 or 6 loculi; ventral
tubular ducts absent but small microducts scat-
tered ventrally; eyespots absent; legs and anten-
nae reduced.
First-instar nymph oval; derm with rugose
sculpturing; dorsum lacking setae, pores, ducts
and tubercles; anal plates elongate triangular with
rounded angles; margin with row of slender setae,
each stigmatic cleft with single stout stigmatic
seta; venter with few setae on head and thorax,
short setae in 2 submarginal rows on abdomen,
longer setae in 2 submedial longitudinal rows on
abdomen; antennae 6-segmented; legs well
developed, fore legs each with single digitule.
COMMENTS. This genus shares several fea-
tures with others of the Myzolecaniinae; for ex-
310
ample, Cyclolecanium Morrison, Megasaissetia
Cockerell, Neolecanium Parrott and Pseudo-
philippia Cockerell also possess some form of
dorsal pore or microduct with a bilocular ap-
pearance and that of N. imbricatum (Cockerell)
even has an inner duct (Hodgson, 1994) similar
to that of T. endocanthium. The dorsal ridges of
T. endocanthium are of similar shape but different
number and arrangement from those of Coccus
tumuliferus Morrison (Morrison 1921), which is
not a true species of Coccus Linnaeus (Coccinae)
but undoubtedly a member of the
Myzolecaniinae. Furthermore, the anal plates of
C. tumuliferus are of somewhat similar shape to
those of T. endocanthium but with different ar-
rangement and number of setae; however, the
dorsal pores and setae and ventral ducts of C.
tumuliferus are very different from those of T.
endocanthium. The presence of multilocular
disc-pores and microducts resembling bilocular
pores on the dorsum of Torarchus easily distin-
guishes it from Akermes Cockerell, Alecanopsis
and Myzolecanium, which are the other genera of
the Myzolecaniinae found in Australia; further-
more, it differs from Myzolecanium in having
very shallow stigmatic clefts and from Akermes
and Alecanopsis in having no stigmatic setae.
Torarchus keys out in Hodgson (1994, pp. 91-92)
to the couplet containing Akermes and Alecanop-
sis, but does not fit the description of either.
ETYMOLOGY. Latin torus, round elevation or bulge,
Greek archos, anus; refers to the shape of the plates
surrounding the anus and location of the anal area on a
cuticular bulge.
Torarchus endocanthium sp. nov.
(Figs 1-4)
MATERIAL EXAMINED, HOLOTYPE,
QMT13986, adult ? ,Qld, Mt Crosby, off Crosby Rd
& Bunya St., 27? 32'S, 152? 48 E, 9.iv. 1989, P. Flower.
PARATYPES adult ? 9 only, QLD: 7 adult 9 9, same
data as holotype; 1 adult 9, 2 slides of first-instar
nymphs, Mt Crosby, 18.viii.1988, P. Flower; 1 adult
9, 1 slide of first-instar nymphs, Mt Moffatt Nat. Park,
Kenniff' s Lookout, 24° 55'S, 147° 59'E, 13.xii.1987,
G. Monteith, G. Thompson & D. Yeates; 7 adult 9 9,
3 immature 9 9, 6 slides of first-instar nymphs,
Auburn Gorge, SW Mundubbera, 25? 43’S, 151° 03’E,
late March 1989, G. Monteith; 3 adult 9 9, Keysland,
20 km NW of Wondai, late March 1989, G. Monteith.
Location of paratypes: 4 adult ? 2 and 2 slides
of first-instar nymphs in ANIC, 1 adult 9 in
BMNH, remainder in QM (QMT26022-26035).
DIAGNOSIS. As for genus.
MEMOIRS OF THE QUEENSLAND MUSEUM
DESCRIPTION. Adult 9 (10 specimens
measured). Live material. Body of young 9 yel-
low, with sparse coating of white, powdery wax.
Mounted material. Body transversely oval,
rounded on each side, 1.3-1.8 times wider than
long, rather flattened but with elevated areas on
dorsum; stigmatic clefts shallow; anal cleft fused.
Length 1.5-2.6 mm, width 2.3-4.4 mm.
Dorsum. Derm membranous except for narrow
crescent of sclerotisation around anterior margin
of anal plates and light sclerotisation at margin in
stigmatic cleft; with 7 radial ridges dorsally on
each side of body, forming an elevated submedial
area following curve of margin, with anterior-
most and posteriormost ridges smallest; a
rounded raised area lies posterior to anal plates
(Fig. 2A) with central depression marking posi-
tion of anal cleft. Dorsal setae flagellate, 15-100
pm long in submarginal areas and around anal
plates, longer (70-150 pm) medially and
clustered on ridges; a group of 13-26 very long
(230-500 um), robust setae posterior to anal
plates on raised area. Dorsal pores of 3 kinds: 1,
multilocular disc-pores 5.5-7.5 jum in diameter
with 4-8 (mostly 6, Figs 2C, 3i) loculi, scattered
in marginal and submarginal area; 2, microducts
resembling bilocular pores (Figs 2C, 3h), 3-4 jum
in greatest dimension, densely distributed over
entire dorsum; 3, simple pores (Fig. 3f) 3-4 pm
in diameter, scattered over dorsum. Preopercular
pores absent. Dorsal tubular ducts absent. Dorsal
tubercles absent. Anal plates lobe-like, rounded
apically (Figs 2A, 2B, 3d), each 150-210 jum
long, 110-150 jum wide when measured in natural
position (distortion during mounting common);
with 5-6 small setae apically on each plate. Ano-
genital fold with 2 pairs of setae in hypopygial
position, a pair of larger setae at each corner of
anterior margin and 1 pair laterally. Anal ring
58-68 um in diameter, probably with 6 pairs of
setae, 65-115 zm long, rather flattened (Fig. 3e),
difficult to see when retracted inside anal tube
(Fig. 2B).
Margin. Marginal setae flagellate (Fig. 3g), 25-
150 jum long, in irregular double row around
entire margin except absent in stigmatic clefts
and posteriorly where fused anal cleft joins mar-
gin. Stigmatic clefts very shallow with small area
of light sclerotisation; lacking stigmatic setae.
Eyespots apparently absent.
Venter. Derm membranous; only abdominal seg-
NEW ANT-ASSOCIATED COCCID IN CANTHIUM
311
FIG. 3. Adult 9 of Torarchus endocanthium. Enlargements: A, quinquelocular disc-pore from stigmatic furrow.
B, ventral microduct. C, pregenital disc-pore with 6 loculi. D, anal lobes viewed dorsally (on left) with anal ring
and anal ring setae indicated by dashed lines, and ventrally (on right) showing setae and supporting bar of
ano-genital fold, E, anal ring seta. F, simple pore. G, marginal setae. H, lateral and end-on views of dorsal
microduct. I, multilocular disc-pore.
mentation discernible, Ventral setae flagellate,
15-75 um long, sparsely scattered. Pregenital
disc-pores (Fig. 3c) 8-10 pm in diameter with 5-7
(mostly 5-6) loculi, distributed around vulva on
posterior segments and in an irregular line reach-
ing to metathoracic spiracle of each side. Stig-
matic furrows each with quinquelocular
disc-pores (Fig. 3a), 5-6 jum in diameter, in a band
from margin to spiracle. No preantennal pores
present. Ventral microducts (Fig. 3b) each with
outer ductule 4-5 um long and inner ductule with
conspicuous innermost end, each microduct ap-
pearing as a small slightly oval pore when viewed
end-on, scattered throughout venter. Ventral
tubular ducts absent. Spiracles well developed,
with conspicuous muscle plate to each peritreme;
anterior spiracle plus peritreme 150-200 um
long, 110-135 i.m wide; posterior spiracle plus
peritreme 150-200 jum long, 120-140 um wide.
Legs reduced, each 90-130 ium long, trochanter
fused with femur, fusion of tibia and tarsus partial
to complete; each claw small, without denticle;
312
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 4. First-instar nymph of Torarchus endocanthium, with anus everted. Enlargements: A, quinquelocular
disc-pore from stigmatic furrows. B, stigmatic area with stigmatic seta in cleft, 2 marginal setae and quin-
quelocular disc-pores. C, apex of hind leg showing digitules. D, dorsal (on left) and ventral (on right) views of
anal plates. E, anal ring with setal positions indicated by blackened spots; F, dorsal view of anal lobes and
everted anal ring; G, marginal seta.
claw digitules longer than claw but shorter and
broader than tarsal digitules; tarsal digitules 22-
25 jum long. Antennae reduced, with 5-6 seg-
ments at most, segmentation often indistinct; total
length 95-150 jum; with fleshy setae 13-25 jum
long on apical segments and flagellate setae 15-
65 jum long on all segments. Clypeolabral shield
350-390um long, 270-320um wide. Labium 1-
segmented, 140-150 um long, c.150 jum wide.
First-instar nymph (10 specimens measured).
Mounted material. Body oval, 463-560 j.m long,
270-315 wm wide. Derm membranous
throughout but with rugose sculpturing dorsally
and ventrally, most clearly visible in SEM (Fig.
2D). Segmentation not readily apparent (but ob-
vious in whole nymphs under SEM).
Dorsum. Setae, pores, ducts and tubercles absent.
NEW ANT-ASSOCIATED COCCID IN CANTHIUM
Anal plates (Fig. 4d) elongate triangular with
rounded angles, 40-48 jum long, 15-24 jum wide;
anterolateral margin 22-26 jm long,
posterolateral margin 25-30 jum long; dorsal sur-
face with scattered microspines. Each plate with
4 dorsal setae, 3 on apex of plate, 1 on mesal
margin; median seta on apex robust, 190-225 um
long, about half length of body. Ano-genital fold
with 1 pair of anterior margin setae and 1 pair of
lateral margin setae. Anal ring (Fig. 4e) ap-
proximately circular, 18-22 jum in diameter, with
about 14 irregularly shaped pores and 6 setae,
45-68 jum long, with 2 setae distinctly shorter and
more slender than other 4 (Fig. 4f).
Margin. Marginal setae (Fig. 4g) 10-15 jum long,
slender, tapering to a point, usually curved, point-
ing in posterior direction, distributed as follows:
8 around head between anterior stigmatic clefts,
4 between each pair of stigmatic clefts, and 18 on
abdomen. Stigmatic clefts moderately developed
(Fig. 4b), each with single stout stigmatic seta
10-16 wm long, bordered by 2 slender marginal
setae, | anterior and | posterior to each stigmatic
seta. Eyespots present just above level of antennal
scape.
Venter. Ventral body setae slender, of 2 lengths:
submarginal setae short, 2-5 jum long, in 2 lon-
gitudinal rows each of 7 on each side of abdomen
(2 pairs of setae per segment), | seta between
anterior and posterior stigmatic clefts and | pair
at apex of head; submedial setae 20-45 jum long,
in 2 longitudinal rows of 6 each on abdomen (1
pair per segment), setae longer posteriorly, and 1
pair between antennae. Spiracles plus peritremes
15-21 jum long, 6-10 jum wide. Stigmatic furrows
each with quinquelocular disc-pores (Fig. 4a),
2-3 pm in diameter, in single row; anterior
spiracular disc-pore bands each with 4-6 pores;
posterior spiracular disc-pore bands each with
4-5 pores. Other pores and ducts absent. Legs
well developed, 210-245 pm long, without
tibiotarsal sclerotisation or free articulation; | or
afew flagellate setae on each segment; 2 knobbed
claw digitules (Fig. 4c) per leg; 2 knobbed tarsal
digitules (Fig. 4c) per leg, except fore legs each
with only 1 digitule; tarsal claw with small den-
ticle. Antennae well developed, 6-segmented,
130-160j.m long; segment III longest, 33-45 jum;
5 fleshy setae (1 on IV, 1 on V, 3 on VI), 13-25
pam long, and about 15 hair-like setae, 10-53 jum
long. Mouthparts with clypeolabral shield 83-90
pum long, 63-77 jum wide; labium 1-segmented,
313
30-36 jum long, 45-50 jum wide, with 4 pairs of
setae; stylets looped, total length 500-660 um.
COMMENTS. The first-instar nymphs of most
Coccidae possess 3 stout stigmatic setae per stig-
matic cleft. The nymph of T. endocanthium has
only one differentiated stigmatic seta per stig-
matic cleft (presumably homologous with the
median stigmatic seta of other coccids). The 2
slender setae that border each stigmatic seta in
nymphs of T. endocanthium appear identical to
the remainder of the marginal setae but may be
homologous with the 2 stout lateral stigmatic
setae that normally accompany the longer median
stigmatic seta in other coccids.
ETYMOLOGY. Greek endon, within or inside, with
the name of the host plant Canthium.
ACKNOWLEDGEMENTS
We thank Geoff Monteith, Queensland
Museum for drawing our attention to this scale
insect, for loan of the specimens and for Fig. 1.
We acknowledge the facilities and technical sup-
port of the Electron Microscopy Unit, ANU.
Chris Hodgson, Geoff Monteith, Katie Strong
and Doug Williams made helpful comments on
the manuscript. Robert Hoare advised on the use
of Latin and Greek to form the new names. Diane
Bridson provided the name of the African
Rubiaceae.
LITERATURE CITED
BEQUAERT, J. 1922. Ants in their diverse relations to
the plant world. Bulletin of the American Museum
of Natural History 45: 333-584.
GREEN, E.E. 1924. On some new species of Coccidae
from various sources. Bulletin of Entomological
Research 15: 41-48.
GULLAN, P.J., BUCKLEY, R.C. & WARD, P.S. 1993.
Ant-tended scale insects (Hemiptera: Coccidae:
Myzolecanium) within lowland rain forest trees in
Papua New Guinea. Journal of Tropical Ecology
9: 81-91.
HODGSON, C J. 1994. "The Scale Insect Family Coc-
cidae: An Identification Manual to Genera.' (CAB
International: Wallingford) 639p.
1995. Observations on the structure of the spiracles
of adult female Coccidae. Israel Journal of En-
tomology 29: 47-55.
MILLER, D.R. 1991. Superfamily Coccoidea. pp. 90-
107. In Stehr, F.W. (ed.) ‘Immature insects.
Volume 2.” (Kendall/Hunt: Iowa).
MONTEITH, G.B. 1989. Ant symbiosis in the plant
genus Canthium in Queensland. News Bulletin of
314
the Entomological Society of Queensland 17(3):
31-32.
1990. The plant-ant connection. Wildlife Australia
27: 6.
MORRISON, H. 1921. Some nondiaspine Coccidae
from the Malay Peninsula, with descriptions of
apparently new species. The Philippine Journal of
Science 18: 637-677.
NEWSTEAD, R. 1910. On two new species of African
Coccidae. Journal of Economic Biology 5: 18-22.
QIN, T.K. & GULLAN, P.J. 1989. Cryptostigma Ferris:
acoccoid genus witha strikingly disjunct distribu-
tion (Homoptera: Coccidae). Systematic En-
tomology 14: 221-232.
RAY, C.H., Jr. & WILLIAMS, M.L. 1980. Description
of the immature stages and adult male of Pseudo-
MEMOIRS OF THE QUEENSLAND MUSEUM
philippia quaintancii (Homoptera: Coccoidea:
Coccidae). Annals of the Entomological Society
of America 73: 437-447.
SHEFFER, B.J. & WILLIAMS, M.L. 1990. Descrip-
tions, distribution, and host-plant records of eight
first instars in the genus Toumeyella (Homoptera:
Coccidae). Proceedings of the Entomological
Society of Washington 92(1): 44-57.
WILLIAMS, D.J. 1985. 'Australian Mealybugs'.
(British Museum (Natural History): London)
431p.
WILLIAMS, D.J. & WATSON, G.W. 1988. ‘The scale
insects of the tropical South Pacific region. Part 2.
The Mealybugs (Pseudococcidae)’. (C.A.B. Inter-
national: Wallingford) 260p.
THE ATLANTIC-MEDITERRANEAN BIVALVE, CORBULA GIBBA (OLIVI)
(CORBULIDAE: MYOIDEA) IN PORT PHILLIP BAY, VICTORIA
JOHN M. HEALY AND KEVIN L. LAMPRELL
Healy, J.M. & Lamprell, K.L. 1996 07 20: The Atlantic-Mediterranean bivalve, Corbula
gibba (Olivi) (Corbulidae: Myoidea) in Port Phillip Bay, Victoria. Memoirs of the
Queensland Museum 39(2): 315-318. Brisbane. ISSN 0079-8835.
Established populations of the common European corbulid bivalve, Corbula (Varicorbula)
gibba (Olivi, 1792) are recorded for the first time from Australia within Port Phillip Bay,
Victoria. Aside from a tendency to exhibit pink radiating bands, the Victorian specimens are
indistinguishable from typical French and British populations. The species appears to have
been introduced to Australia sometime after the mid- 1980's, as it was not recorded in benthic
surveys of Port Phillip Bay up to 1972 or encountered by local collectors prior to 1983.
Although we cannot identify the precise geographical origin of the Victorian population on
morphological features, the recent plague-level outbreak of the polychaete Sabella spallan-
zanii in Port Phillip Bay suggests a Mediterranean origin. Presumably this species has been
spread via release of ballast water. C] Corbula gibba, Mollusca, Bivalvia, introduced species.
John M. Healy, Department of Zoology, University of Queensland, 4072, Queensland,
Australia; Kevin L. Lamprell, Malacology Section, Queensland Museum, P.O. Box 3300,
Queensland 4101, Australia; received 16 January 1996.
Accidental or in some cases deliberate intro-
duction of foreign marine species, has, and con-
tinues to be a serious problem in ports throughout
the world. While certain species may be
transported as adults attached to ships' hulls, a
number of species are apparently being conveyed
as larval stages in ballast water (Carlton, 1985,
1987, 1989; Williams et al., 1988; Carlton et al.,
1990). To survive such long voyages, these larvae
are necessarily hardy and represent species com-
mon within their native habitat. The recent
plague-proportion spread of the Mediterranean
tubiculous polychaete Sabellaria spallanzanii
throughout Port Phillip Bay — an event now
threatening the viability of a major scallop fishery
— has highlighted the importance of under-
standing changes occurring in local shallow
water faunas around the Australian coastline.
Within the Mollusca, accidentally introduced
species of bivalves have in recent years posed a
serious challenge to industry and/or the ecologi-
cal health of marine and freshwater habitats. The
fouling damage incurred by the spread of the
European "zebra mussel’ Dreissena polymorpha
throughout the lakes of the United States (Hebert
et al., 1989; Topping, 1991) is one notable ex-
ample of the serious ecological and potential
economic impact of an introduced bivalve
species.
In recent years authors have documented the
sudden appearance of foreign bivalve species in
Australian waters including the Japonic mytilids
Musculus imus (Bartsch) and Musculista sen-
housia (Benson) to Western Australia (Kendrick
& Slack-Smith, 1982; Slack-Smith & Brearley,
1987), the Japonic ostreid Crassostrea gigas (evi-
dently a deliberate introduction) and the New
Zealand venerid Paphia largilliertii . During our
studies of Australian Corbulidae (basket clams),
our attention was drawn to an apparently un-
known species from Port Phillip Bay by Mr. R.
Burn of the Museum of Victoria. This corbulid
had not been taken in two extensive benthic sur-
veys of the bay between 1958 and 1972, nor had
it been gathered by local collectors prior to 1983
(R. Burn pers comm.). Thus we suspected an
introduced species. We could find no closely
comparable Asian or American species. How-
ever, there was very close resemblance between
the mystery Australian corbulid and the common
Atlantic-Mediterranean Corbula (Varicorbula)
gibba (Olivi, 1792). We illustrate and describe
Australian specimens of C. gibba and compare
them to Atlantic material adding a discussion on
the implications of this introduction.
MATERIALS AND METHODS
Australian material (all Port Phillip Bay, Vic-
toria): Hampton Beach - several specimens (coll.
G. Macaulay). Mornington (20 miles from
Hampton), from mud on the back of scallop
trawlers docked on 3 Nov. 1991 (preserved in
10% formalin in sea water; coll. G. Macaulay)
(K.Lamprell collection). Southern Port Phillip
Survey 1986-1990 Marine Research Group of
316
Victoria (Museum of Victoria) - Lots NMVF
60444, 60541-60551 all dredged in 6-16m of
water.
European material: France: 2 specimens from
Bretagne (P. van Pel); England: alcohol-
preserved specimens dredged off Shellness, Beer
and Milford Haven (all England) (Natural His-
tory Museum, London). Ireland: alcohol-
preserved specimens dredged off Dublin (Natural
History Museum, London).
SYSTEMATICS
Superfamily MYOIDEA
Family CORBULIDAE
Corbula (Varicorbula) gibba Olivi,1792
Corbula nucleus Lamarck,1818.
DESCRIPTION. Shell to length of 13.5 mm,
solid, posterior side longer than anterior; right
valve much larger and encompassing the left
valve marginally, with umbone extending
beyond the margin of the left valve; both valves
with a well defined postero-umbonal ridge.
Sculpture: right valve with well developed, flat,
moderately wide, concentric ridges; interstices
narrow; left valve with fine, closely set, raised,
concentric ridges crossed by several raised, radial
ridges which extend from the umbones to the
margins. Colour: white, with variable brown or
reddish radial rays; internally white or deep
purple; periostracum on right valve, thin, light
brown, left valve light brown towards the um-
bones, thick, dark brown, concentric concentric
layers marginally.
HABITAT. In sandy mud, obtained from scallop
trawlers and by diver in littoral mud and sand.
DISTRIBUTION. Widely distributed throughout
the Atlantic Ocean, Mediterranean Sea and ex-
tending into the Black Sea (Yonge, 1946; Tebble,
1966; Hrs-Brenko, 1981); Port Phillip Bay, Vic-
toria.
COMPARISONS. We find no characters to
separate the Port Phillip Bay specimens from
Atlantic and Mediterranean Corbula gibba, other
than a tendency to reach a smaller size (Fig. 1).
Conceivably this could be an environmentally
induced effect due to suboptimal nutrient levels
or growth conditions. Commonly the Port Philip
specimens showed pink-purple colouration on
the inside of the valves and/or one or more
coloured rays externally, whereas most Atlantic
MEMOIRS OF THE QUEENSLAND MUSEUM
and Mediterranean material examined showed no
or slight traces of colour (some colouration and
external coloured rays present in examined
material from Bretagne, France). Again this may
be due to suboptimal conditions for Port Phillip
Bay animals or reflect retention or exaggeration
of juvenile colouration. Specimens with no trace
of purple colour were also common in the Port
Phillip Bay material, and for this reason we do not
regard colour differences between Australian and
Atlantic material as highly significant (unfor-
tunately very few of the Atlantic and Mediter-
ranean specimens available were of the juvenile
size-class).
DISCUSSION. Corbula (Varicorbula) gibba has
an extensive geographical range in the Atlantic
(Norway south to Angola) and the Mediterranean
(including the Black Sea [Hrs-Brenko, 1981]).
Suter (1913) reported the species (as C. nucleus)
from the Chatham Islands, east of the South Is-
land of New Zealand, but this needs to be con-
firmed. In many areas C. gibba may be the most
abundant subtidal bivalve (for example, off the
English coast (Yonge, 1946), some workers in
fact using the species for population analyses
(Hrs-Brenko, 1981). Corbulid species can sur-
vive long periods in the ballast water of ocean
going vessels, and then go on to generate heavy
or at least significant populations in foreign har-
bours (e.g. the Chinese corbulid Potamocorbula
amurensis in San Francisco Harbour - Carlton et
al., 1990). It is not possible to identify the precise
geographical origin of the Victorian population
of C. gibba on morphological features. The recent
heavy infestation of the Mediterranean tubeworm
Sabella spallanzanii throughout large areas of
Port Phillip Bay does however suggest a possible
source (via ballast water) for these bivalves.
Although Poramocorbula amurensis is now ap-
proaching plague proportions in San Francisco
Bay (reaching densities of over 10000/ m2) and
out-competing endemic species for settlement
space (Carlton et al., 1990; Nichols et al., 1990),
there is no direct evidence to suggest that Corbula
gibba will cause similar problems in Port Phillip
Bay. Nevertheless C. gibba is known to occur in
enormous numbers at certain localities in the
Atlantic (2450/m?) and can inhabit a variety of
sediment types (coarse sand to fine mud, but
preferring sand) and are capable of thriving even
in substantially polluted areas (Hrs-Brenko,
1981). For these reasons we feel that the popula-
tion of C. gibba in Port Phillip Bay should con-
tinue to be monitored, particularly in relation to
EUROPEAN CORBULA GIBBA IN PORT PHILLIP BAY 317
FIG. 1. Corbula (Varicorbula) gibba (Olivi, 1792). A-C from silty mud 2-4m, Sandringham Harbour, Port Phillip
Bay. D-F, Bretagne, France. G-H, dredged off Shellness, England. A, external view of right valve (valve length
10.8 mm). B, internal view of right valve (valve length 10.8 mm). C, external view of left valve (valve length
9.9 mm). D, external view of right valve (valve length 12.6 mm). E, internal view of right valve (valve length
13.5 mm). F, internal view of left valve (valve length 12.0 mm). G, external view of right valve (valve length
12.5 mm). H, paired valves viewed from left valve (length of right valve 11.5 mm).
318
the impact of this species on the settlement suc-
cess of native bivalves within the Bay. Bearing in
mind the problems caused by the Asian corbulid
Potamocorbula amurensis in the San Francisco
area, vessels operating out of Port Phillip Bay
should not take on ballast water within the Bay.
Similarly, the practice of emptying ballast water
from foreign vessels in Port Phillip Bay should be
discontinued.
ACKNOWLEDGEMENTS
We thank Sue Boyd, Museum of Victoria, Bob
Burn, Geelong, G. Macaulay, Hampton, Kathie
Way, The natural History Museum, London and
Peter van Pel for loan of material. Bob Burn also
very kindly shared his knowledge of corbulids
from Victoria. Photographs of type Corbula
nucleus Lamarck (MHNG 1082/83) (taken by Mr
C Paton) were kindly supplied by the Museum
d'Histoire Naturelle Geneva. This study was sup-
ported by a grant-in-aid from the CSTRO Division
of Fisheries and a Keith Sutherland Award (to
K.L.). An Australian Research Fellowship
enabled the participation of J.H.
LITERATURE CITED
CARLTON, J.T. 1985. Transoceanic and interoceanic
dispersal of coastal marine organisms: the biology
of ballast water. Oceanography and Marine Biol-
ogy Annual Review 23: 313-371.
1987. Patterns of transoceanic marine biological
invasions in the Pacific Ocean. Bulletin of Marine
Science 41: 452-465.
1989. Man's role in changing the face of the ocean:
biological invasions and implications for conser-
vation of near- shore environments. Conservation
Biology 3: 265-273.
CARLTON, J.T., THOMPSON, J.K., SCHEMEL, L.E.
& NICHOLS, F.H. 1990. Remarkable invasion of
San Francisco Bay (California, USA) by the Asian
clam Potamocorbula amurensis. 1. Introduction
and dispersal. Marine Ecology Progress Series 66:
FORBES, E. & HANLEY, S. 1853. A history of British
Mollusca and their shells. Vol. 1,4. (John Van
Voorst: London). pp 180-185.
HEBERT, P.D.N., MUNCASTER, B.W. & MACKIE,
G.L. 1989. Ecological and genetic studies on
MEMOIRS OF THE QUEENSLAND MUSEUM
Dreissena polymorpha (Pallas) : a new mollusc in
the Great Lakes. Canadian Journal of Fisheries
and Aquatic Science 46: 1587-1591.
HRS-BRENKO, M. 1981. Population studies of Cor-
bula gibba (Olivi), Bivalvia, Corbulidae, in the
northern Adriatic Sea. Journal of Molluscan
Studies 47 (1): 17-24.
KENDRICK, G.W. & SLACK-SMITH, S. 1982. Mus-
culus imus (Bartsch) in Western Australia.
Australian Shell News 37-38: 4.
NICHOLS, F.H., THOMPSON, J.K. & SCHEMEL,
L.E. 1990. Remarkable invasion of San Francisco
Bay (California, USA) by the Asian clam
Potamocorbula amurensis. II. Displacement of a
former community. Marine Ecology Progress
Series 66: 95-101.
REEVE, L.A. 1843-1844. Monograph on the genus
Corbula. In L.A. Reeve (ed.) Conchologica
Iconica; or, illustrations of the shells of mollus-
cous animals 2:5 pls. (pl. 1, Aug. 1843; pl. 2 Sept;
pl. 3 Jan. 1844; pl. 4 April; pl. 5 May).
SLACK-SMITH, S.M. & BREARLEY, A. 1987. Mus-
culista senhousia (Benson, 1842); a mussel
recently introduced into the Swan River Estuary,
Western Australia. (Mollusca: Mytilidae).
Records of the Western Australian Museum 13
(2): 225-230.
SUTER, H. 1913. Manual of the New Zealand Mollus-
ca. (John Mackay: Wellington) 1120p.
TEBBLE, N. 1966. British bivalve seashells. A hand-
book for identification. (British Museum (Natural
History):London).
TOPPING, J. 1991. Zebra mussels: North American
environmental and economic nightmare of the
nineties. Biome 11(1): 2pp.
VOKES, H.E. 1945. Supraspecific groups of the
pelecypod family Corbulidae. Bulletin of the
American Museum of Natural History 86 (1):
1-32.
WILLIAMS, R.J., GRIFFITHS, F.B., VAN DER
WAL, EJ. & KELLY, J. 1988. Cargo vessel
ballast water as a vector for the transport of non-
indigenous marine species. Estuarine and Coastal
Shelf Science 26: 409-420.
YONGE, C.M. 1946. On the habits and adaptations of
Aloidis (Corbula) gibba. Journal of the Marine
Biological Association of the United Kingdom
26(3): 258-276.
ZHUANG, Q. & CAI, Y. 1983. Studies on the Cor-
bulidae (Bivalvia) off Chinese coasts. Transac-
tions of the Chinese Malacological Society 1:
57-68.
EVOLUTION AND ZOOGEOGRAPHY OF AUSTRALIAN FRESHWATER TURTLES
BEN MANNING AND CHRISTOPHER P. KOFRON
Manning, B. & Kofron, C.P. 1996 07 20: Evolution and zoogeography of Australian
freshwater turtles. Memoirs of the Queensland Museum 39 (2): 319-331. Brisbane. ISSN
0079-8835.
The contemporary turtle fauna of Australia comprises freshwater and marine species but no
terrestrial taxa. The literature on evolution and zoogeography of the Australian freshwater
turtles (Chelidae) is reviewed. Because opposing conclusions were reached in several of
these studies, we critically examine each of them. We accept the phylogeny of Georges &
Adams (1992); Elseya and Emydura are not synonymous and Elseya consists of the E.
dentata and E. latisternum species groups (generically distinct). However, Pseudemydura
umbrina shares common ancestry with other Australian short-necked turtles and is their
extant sister taxon. The E. latisternum group is the sister taxon of the non-Pseudemydura
Australian short-necked turtle genera.
Understanding pre-Cretaceous dispersal of the ancestral lineage on the South American-An-
tarctic-Australian supercontinent is important for ascertaining relationships among
Australian freshwater turtles. Increasing aridification during the Pleistocene, resulting in
isolation within river drainages, appears to have augmented allopatric speciation.
We recognize 23 extant species of Australian chelid turtles in 7 genera: Chelodina expansa,
C. longicollis, C. novaeguineae, C. oblonga, C. rugosa, C. steindachneri and one un-
described species; Elseya dentata and 3 undescribed species; Elseya latisternum and three
undescribed species; Elusor macrurus; Emydura macquarrii (inclusive of E. australis, E.
krefftiiand E. signata), E. subglobosa, E. victoriae and 2 undescribed species; Pseudemydura
umbrina; and Rheodytes leukops. Relationships among genera remain enigmatic.
O Australia, evolution, turtles, zoogeography.
Ben Manning & Christopher P. Kofron, Department of Zoology, University of Queensland,
Brisbane, Queensland 4072, Australia. (Present address CPK: Department of Environment,
Queensland National Parks and Wildlife Service, Far Northern Region, PO Box 2066,
Cairns, Queensland 4870, Australia); received 2 November 1995.
The reptilian order Testudines contains about
257 living species of turtles (Ernst & Barbour,
1989) in two suborders, Cryptodira and
Pleurodira. Cryptodira contains 10 families and
about 203 species, and Pleurodira two families
and about 54 species. Cryptodiran turtles with-
draw the head by vertical flexure of the neck into
an S, and the pelvic girdle does not attach to the
plastron. Pleurodirans withdraw the head by hor-
izontal flexure of the neck, and the pelvic girdle
attaches to the plastron. Each suborder is charac-
terised by its own specialised cervical vertebrae.
Although cryptodirans are more derived than
pleurodirans, the former did not stem from the
latter (Ernst & Barbour, 1989). The earliest fos-
sils of cryptodiran turtles are Middle Jurassic, 50
million years older than the earliest pleurodiran
fossils (Legler & Georges, 1993a). However, the
fossil record is poor, and pleurodiran turtles
probably originated earlier (Gaffney, 1979; Bull
& Legler, 1980; Ernst & Barbour, 1989). The two
suborders are probably derived from separate an-
cestral lineages, each of which presumably could
not withdraw the head because of non-specialised
cervical vertebrae (Gaffney, 1977).
Pleurodira contains the Pelomedusidae and
Chelidae. Pelomedusids inhabit Africa, Mada-
gascar and South America, but previously also
Europe, Asia and North America (Frair, 1980).
They retain mesoplastral bones in the shell and a
squamosal bone in the skull; thus, Pelomedusidae
is plesiomorphic to Chelidae (McDowell, 1983),
which lacks mesoplastral and squamosal bones.
The contemporary fauna of Australia com-
prises freshwater and marine turtles, with a com-
plete absence of terrestrial species. The
Australian freshwater turtles are pleurodirans of
the Chelidae, with exception of one cryptodiran,
Carettochelys insculpta of the Carettochely-
didae. The latter species inhabits northern
Australia and New Guinea, and it is the sole
extant member of its family. As recently as the
Eocene, Carettochelydidae had a broad distribu-
tion in the Northern Hemisphere (Legler & Geor-
ges, 19932). The living marine turtles of Australia
are cryptodirans of the Cheloniidae and Der-
mochelyidae.
320
TABLE 1. Recent primary research papers that recog-
nize the two nominal turtle genera Elseya and
Emydura as separate, or as congeneric
Separate recognition
Cann & Legler (1994)
White & Archer (1994)
Georges (1993)
Georges & Adams (1992)
Legler (1985)
Congeneric recognition
Gaffney (1991)
Gaffney, Archer & White
1989)
McDowell (1983)
Gaffney (1981)
Frair (1980)
Legler & Cann (1980
Burbidge, Kirsch & Main
(1974) -
Legler (1981
Australia was inhabited also by freshwater
cryptodirans of the living Trionychidae (Pliocene
- Gaffney & Bartholomai, 1979; Willis, 1993),
and possibly Emydidae (middle Miocene; Mur-
ray & Megirian, 1992). A primitive non-marine
cryptodiran was reported from the Early Cret-
aceous of Victoria (Gaffney et al., 1992), and also
primitive terrestrial turtles that are neither cryp-
todiran or pleurodiran (Rich & Rich, 1989).
Australia was inhabited as recently as the Pleis-
tocene by giant horned terrestrial turtles (shells
1.5m length) of the extinct eucryptodiran
Meiolaniidae, which also occurred in South
America, Madagascar, Lord Howe Island and
New Caledonia (Mittermeier, 1984; Archer et al.,
1991). The fossil turtles of Australia were
reviewed by Gaffney (1981, 1991) and Molnar
(1991).
Outside Australia and New Guinea, the Chel-
idae inhabit only freshwaters east of the Andes in
South America (Iverson, 1992), but a fossil
specimen was found in Antarctica (Pritchard &
Trebbau, 1984). Although chelids are not the
dominant turtle family in South America, they are
comparable in diversity to those in Australia.
Consequently, constructing a phylogeny of fresh-
water turtles requires understanding of evolution-
ary relationships between the two geographically
distant groups, together with the Pelomedusidae.
Australian chelids separate into two broad mor-
phological and ecological groups (Goode, 1967;
Cann, 1978), long-necked species and short-
necked species. Chelodina contains long-necked
species that spear or ambush their food
(Pritchard, 1984; Legler & Georges, 1993b).
They have 4 claws on the forelimbs, and a neck
at least 2/3 the length of the shell (Cogger, 1992),
with which they strike out rapidly and capture
prey (Legler, 1993). Elseya, Elusor, Emydura,
MEMOIRS OF THE QUEENSLAND MUSEUM
Pseudemydura and Rheodytes are short-necked
active foragers (Legler & Georges, 1993b; Cann
& Legler, 1994). These have 5 claws on the
forelimbs, and a neck less than 1/3 the length of
the shell (Cogger, 1992).
South American chelids also separate into long-
necked sit-and-wait predators and short-necked
active foragers (Ernst & Barbour, 1989). Like
Australian long-necked Chelodina, South Am-
erican long-necked Chelus and Hydromedusa
have 4 claws on the forelimbs. Also, like Aust-
ralian short-necked genera, South American
short-necked Phrynops and Platemys have 5
claws on the forelimbs. The questions are posed:
How are the geographically-distant Australian
and South American chelids related? Are the
similar groups a result of ecological convergence,
or do they share common ancestry?
Australian chelids are morphologically conser-
vative (Cogger, 1992), which has hindered the
recognition of valid taxa and obscured their true
relationships. Discoveries of 3 new genera and 11
new species (Legler & Cann, 1980; Georges &
Adams, 1992; Cann & Legler, 1994), of which 1
genus and 9 species await description, have made
their phylogeny even more enigmatic. Ecologies
of most species are poorly known. There is con-
troversy regarding relationships and validity of
taxa, especially the nominal genera and species
of Elseya and Emydura (Table 1). The taxonomy
is in a state of flux. Cogger (1992) recognized 15
species in 5 genera; Georges & Adams (1992)
recognized 23 species in 7 genera.
This paper has 2 purposes; a review of the
literature on evolution and zoogeography of
Australian freshwater turtles and a review of op-
posing published conclusions. We recommend
acceptance of one of the proposed phylogenies.
EVOLUTION OF THE CHELIDAE
The most important evolutionary and
zoogeographic studies of Australian freshwater
turtles are by Burbidge et al. (1974), Gaffney
(1977), Legler & Cann (1980), Legler (1981),
McDowell (1983), Georges & Adams (1992),
and Cann & Legler (1994). The zoogeographic
background was established by Burbidge et al.
(1974), McDowell (1983), and Pritchard & Treb-
bau (1984). The systematic studies by Legler &
Cann (1980), Legler (1981), McDowell (1983),
and Cann & Legler (1994) approach an ideal
sampling strategy. Georges & Adams (1992)
sampled extensively and used sensitive bio-
AUSTRALIAN FRESHWATER TURTLES
FIG. 1. Relationships within the Chelidae, based upon
26 cranial characters (Gaffney, 1977). Black dots
indicate Australian genera, open circles South
American genera.
chemical techniques capable of detecting cryptic
relationships.
Among the 3 types of systematic methods
(evolutionary, phenetic, cladistic), the last has the
strongest Popperian scientific basis. Cladistics
attempts to test hypotheses by designating char-
acters as either plesiomorphic (ancestral) or
derived (descendent). Derived characters shared
between taxa are considered evidence of common
ancestry unless homoplasy is demonstrated (Hen-
nig,1966; Brooks & McLennan, 1991). The in-
group is the taxon whose evolutionary relation-
ship is being tested. It is compared to a reference
taxon, the outgroup, which is usually considered
the sister taxon of the ingroup (Brooks & Mc-
Lennan, 1991). Determination of the outgroup is
a crucial and often most subjective decision in
cladistics (Hennig, 1966; Maddison et al., 1984).
Gaffney (1977) undertook a cladistic analysis
of living chelids using 26 cranial characters (Fig.
1). On the basis of skull simplification, he desig-
nated Emydura (inclusive of Elseya) as the out-
group of all other chelid genera. By assigning
equal weight to all character states, Gaffney
(1977) assured that all changes had equal prob-
ability. The Australian long-necked Chelodina
were shown to be more closely related to South
American long-necked Chelus and Hydromedusa
than to Australian short- necked Emydura and
Pseudemydura, The South American short-neck-
ed Phrynops and Platemys were positioned be-
tween Chelodina and the Australian short-necked
genera.
Although Gaffney (1977) recognized Elseya
and Emydura as congeneric (as did Frair [1980]
and McDowell [1983]), this view is not common-
ly followed (White & Archer, 1994). However,
Gaffney et al. (1989) claimed that the two genera
are not adequately diagnosed.
Pseudemydura shares no derived skull charac-
ters with either the Australian and South
American long-necked turtles or Australian
short-necked turtles, but instead posseses 7 uni-
que skull characters (Gaffney, 1977). Gaffney
(1977) erected the monogeneric Pseudemy-
durinae and considered Pseudemydura as sister
taxon to other chelids. Pseudemydura is autapo-
morphic among chelids in lacking temporal
emarginations of the skull (Gaffney et al., 1989),
and P. umbrina is unique among turtles by its
behavior of excavating the egg chamber with
forelimbs rather than hindlimbs (Kuchling,
1993). However, several derived characters are
shared between Pseudemydura and other
Australian short-necked turtles, such as reduced
neural bones and cornified head scutes (Mc-
Dowell, 1983). The intergular scute extending
posteriorly to separate the humeral scutes was
considered unique and derived in Pseudemydura
(Burbidge et al., 1974), but McDowell (1983)
observed it also in Elseya dentata and Emydura
australis. Thus there may be no need for the
Pseudemydurinae,
Because changes in chromosome morphology
are quantum changes (King, 1985), cladistic
analyses using karyotypes are free from problems
associated with measuring continuous variables
(King, 1985). Mutations detected by G- and C-
banding can be treated as analogues of mor-
phological character states (King, 1985). Evol-
utionary relationships of the cryptodirans were
studied in this manner by Bickham & Baker
(1976) and Bickham & Carr (1983). Also using
G- and C-banding, Bull & Legler (1980) con-
ducted chromosomal studies of 13 of the 14
pleurodiran genera. However, chelids do not
respond well to this staining technique and a
cladistic analysis was not done.
Even so, Bull & Legler (1980) gave valuable
insight into evolution of chelids. For example, the
ancestral karyotype is thought to be the one that
most species share (King, 1985). The majority of
chelids are uniform in number (2n = 50) and
bands of their chromosomes, which supports the
outgroup designation of Emydura (and Elseya)
322
ANTARCTICA
FIG, 2. The position of Australia and Antarctica during
the mid Cretaceous (after Rich & Rich [1989]). Black
areas in Australia indicate the seaway extant during
the Cretaceous.
by Gaffney (1977). The Cryptodira existed
during the Early Cretaceous. Because the level of
chromosomal variation among cryptodiran fam-
ilies is less than that between the two pleurodiran
families, pleurodirans probably originated prior
to the Cretaceous (Bull & Legler, 1980).
McDowell (1983) used osteological and soft
organ characters to study relationships of the
Chelidae and Pelomedusidae. He concluded that
Elseya and Emydura were the most pelomedusid-
like and, thus, plesiomorphic of the Chelidae,
which further supported Gaffney’s (1977) out-
group designation. Specifically, Elseya dentata
has a longitudinal ridge on the maxilla, a charac-
ter it shares only with the pelomedusid Podoc-
nemis of South America.
Frair (1980) conducted a biochemical study of
pleurodirans using total protein electrophoresis
and immunoelectrophoresis with phenetic anal-
yses, sampling all Chelidae and Pelomedusidae.
He confirmed that each family is monophyletic,
as first proposed by Gaffney (1975) and sup-
ported the arrangement of Gaffney (1979). Frair
(1980) also showed the Chelidae to be more
closely related to the Madagascan pelomedusid
Erymnochelys than to the South American pelo-
medusid Podocnemis and Peltocephalus. Thus it
appears that the lineage giving rise to the
MEMOIRS OF THE QUEENSLAND MUSEUM
Chelidae occurred in both South America and
Africa when joined 115 million years ago. Erym-
nochelys madagascariensis is a relict species
(Pritchard & Trebbau, 1984) and is probably most
like the ancestral condition.
Pritchard & Trebbau (1984) reviewed earlier
zoogeographic hypotheses of the Chelidae, as
well as Pelomedusidae and other living and ex-
tinct South American turtles. In light of recent
geological and evolutionary studies, along with a
fossil chelid from Antarctica of Miocene age,
they suggested the Chelidae could have
originated in Antarctica. Pritchard & Trebbau
(1984) argued that passive dispersal across vast
stretches of ocean is possible only for terrestrial
turtles. Thus an Antarctic landbridge or narrow
sea should be considered in zoogeographic
hypotheses of Australian and South American
chelids (Fig. 2).
The hypothesis that the Australian long-necked
turtles (Chelodina) did not originate from
Australian short-necked turtles (Gaffney, 1977)
is supported by recent biochemical data. Georges
& Adams (1992) studied the evolutionary
relationships of Australian chelids using al-
lozyme electrophoresis (Hillis, 1987). Their ex-
tensive data set included 54 independent loci of
277 specimens of all species (except
Pseudemydura umbrina) from 76 populations
through Australia and New Guinea.
Georges & Adams (1992) employed 3 sys-
tematic methods. Their phenetic analysis incor-
porated principal co-ordinates, which gave
representations of genetic distance. The cladistic
analysis using PAUP (Swofford, 1985) produced
unresolved cladograms because only 14 of 54 loci
could be used as characters. A parsimony method
(Farris, 1972; Baverstock & Schwaner, 1985)
using distance-Wagner and Fitch-Margoliash
procedures, which are not widely accepted, also
produced unresolved cladograms. Although the
several generated phylogenies varied, relation-
ships of some taxa remained constant.
Georges & Adams’ (1992) results suggest that
divergence between the Australian long- and
short-necked turtles is about twice that within
either of the two groups; and the level of diver-
gence among the short-necked species is substan-
tially greater than that among the long-necked.
These results support Gaffney’s (1977)
phylogeny in which South American genera are
intermediates, and short-necked turtles are
probably more primitive.
McDowell (1983) presented a simple dispersal
theory to explain Gaffney’s (1977) results. The
AUSTRALIAN FRESHWATER TURTLES
ancestor of Emydura entered South America
from Australia. The lineage diverged, giving rise
to South American short-necked turtles, from
which evolved a long-necked form that dispersed
back into Australia.
Neural bones are considered an important tax-
onomic character for chelids. They were reported
absent in all living Australian chelids, except
Chelodina oblonga (Burbidge et al., 1974; Mc-
Dowell, 1983). Neural bones were reported in
South American chelids, which McDowell
(1983) considered evidence for convergence of
the long-necked condition. However, Rhodin &
Mittermeier (1977) reported low frequencies of
neural bones in certain populations of both short-
and long-necked Australian chelids.
The Australian long-necked Chelodina oblon-
ga and South American chelids possess well-
developed neural bones, which in C. oblonga
vary from 5-8 (Burbidge et al., 1974; Rhodin &
Mittermeier, 1977). Neural bones (although
reduced in size) persist at low frequencies in
certain populations of Elseya latisternum, an un-
described Elseya sp. from the Manning River
(NSW), C. longicollis, C. novaeguineae and C.
siebenrocki. Of two fossil specimens of Emydura
from mid- Tertiary deposits in Tasmania, one has
one reduced neural bone and the other none (War-
ren, 19693). Apparently the character should be
assessed as absent/reduced in size, or well
developed (Pritchard, 1984).
The fossil record of chelids is poor, beginning
in the Early Cretaceous of Victoria (Warren,
1969b) with a specimen identified first as extant
Emydura macquarrii but later as Chelycara-
pookus arcuatus. It has well-developed neural
bones (Burbidge et al., 1974), and is considered
ancestral to short-necked turtles in which neural
bones are absent or reduced. Early Cretaceous
chelid fossils have been found in NSW and in
western QLD (a Chelycarapookus-like form:
R.E. Molnar, pers. comm.).
Chelycarapookus arcuatus was a river-dweller
(Molnar, 1991) and probably had a broad dis-
tribution in eastern Australia. With vulcanism
forming the Great Dividing Range later during
the Cretaceous (Galloway & Kemp, 1984), and
with increasing aridification during the Pleis-
tocene, the contemporary turtle fauna may have
evolved by allopatric speciation.
McDowell's (1983) zoogeographic hypothesis
explains certain phylogenetic relationships. The
following model is adapted from his, considering
that at times Antarctica had a warmer environ-
323
ment. During the Early Cretaceous, the ancestral
lineage including Chelycarapookus inhabited the
southern supercontinent. During the late Eocene,
Australia was isolated by sea, with members of
the Chelycarapookus lineage left in Australia and
in the Antarctic-South American landmass. The
trend in neural bones in the Australian lineage
was for reduction and loss, giving rise to Elseya,
Elusor, Emydura, Pseudemydura and Rheodytes.
The South American lineage retained well-
developed neural bones, evolving into the South
American short-necked and long-necked turtle
genera. However, the ancestral lineage of South
American long-necked turtles also dispersed
back into Australia, giving rise to Australian
long-necked turtles. The general trend for neural
bones in this group was for reduction and loss, but
well- developed neural bones were retained in the
immediate lineage to Chelodina oblonga. Mor-
phologically and biochemically, C. oblonga is
plesiomorphic in its genus (Burbidge et al., 1974;
Georges & Adams, 1992).
Why did only the ancestral South American
long-necked turtle lineage disperse back to
Australia from the Antarctic-South American
landmass, and not also the ancestral South Am-
erican short-necked turtle lineage? The answer
might be elucidated by examining ecologies of
the two groups. Unlike any South American
short-necked turtle genus, in addition to inhabit-
ing freshwaters, the South American long-necked
Hydromedusa also inhabits estuaries (Ernst &
Barbour, 1989). Australian long-necked Chelo-
dina (C. longicollis, C. expansa) has greater resis-
tance to evaporative water loss than Australian
short-necked Emydura (E. macquarrii; Chess-
man, 1984). Chelodina inhabits permanent and
temporary freshwaters (Chessman, 1984), and
can also tolerate saline waters. During drought,
Chelodina walks overland and finds other water,
or buries into substrate and aestivates (Goode,
1967; Cann, 1978; Grigg et al., 1986). Chelodina
rugosa can inhabit highly saline ephemeral
swamps (Ehmann, 1992), and C. longicollis can
tolerate brackish water for prolonged periods
(Smith, 1993). On the other hand, all Australian
and South American short-necked chelids inhabit
only permanent freshwaters, except Pseud-
emydura which inhabits ephemeral swamps and
aestivates. Thus it appears that the lineage giving
rise to Australian long-necked turtles had greater
potential for dispersal across land and/or a narrow
sea between Australia and the Antarctic-South
American landmass.
324
EVOLUTIONARY RELATIONSHIPS
AMONG AUSTRALIAN FRESHWATER
TURTLES
Ina study of evolutonary relationships of Aust-
ralian chelids, the ideal sample would include all
species; with specimens from throughout a
species’ range, of various sizes and both sexes to
include the variations of geography, ontogeny
and sexual dimorphism. Legler & Cann (1980),
Legler (1981), McDowell (1983), Georges &
Adams (1992), and Cann & Legler (1994) ap-
proach this ideal sampling strategy.
Using cladistic analysis of cranial characters,
Gaffney (1977) diagnosed distant chelid genera,
but he could not separate Elseya from Emydura.
Because the skull and jaw of chelids are probably
correlated conservative characters, the data set
appears inadequate for diagnosing closely related
genera. In addition, the data were from small
samples, often single specimens.
Burbidge et al. (1974) diagnosed Elseya from
Emydura on nuchal and intergular scutes: nuchal
scute usually absent in Elseya, present in Emy-
dura; intergular scute smaller and more rectan-
gular in Elseya, larger and less rectangular in
Emydura. Elseya was reported to have an elon-
gate snout. However, data for Elseya and Emy-
dura were from single specimens of Elseya
latisternum and Emydura macquarrii, their sole
representatives of the two genera.
Burbidge et al. (1974) also used phenetics to
analyze serological data of Australian chelids.
The first 3 dimensions of the principal compon-
ents described 85% of the variation. Among the
short-necked turtles, Elseya dentata clustered
with Elseya novaeguineae; Emydura australis,
Emydura krefftii and Emydura subglobosa
clustered; and the two groups together formed a
closely associated group. Elseya latisternum,
Emydura macquarrii and — Pseudemydura
umbrina were outliers; however, after omitting
the anti-Elseya latisternum data, Elseya latister-
num clustered with Elseya dentata and Elseya
novaeguineae. The long-necked turtle species
(Chelodina) clustered together, but with C.
oblonga an outlier within the genus.
These results showing P. umbrina and C.
oblonga as outliers to the short-necked and long-
necked turtles, respectively, are accepted. Their
sera react strongly and they have different mor-
phologies. However, the designation of Elseya
latisternum as an outlier is treated with caution
because its serum reacted weakly.
Bull & Legler (1980) reported the short-necked
Elseya, Emydura and Rheodytes to have identical
MEMOIRS OF THE QUEENSLAND MUSEUM
karyotypes (2n=50 and gross chromosome mor-
phology), and Pseudemydura differed only
slightly by having chromosome pairs 6 and 10
acrocentric, Pelomedusid turtles differed greatly,
with 2n=26-36 and 5 large macrosomes.
Using total protein electrophoresis, Frair
(1980) observed only one type of electrophoretic
pattern among two species of Elseya and two
Emydura. The electrophoregrams of Elseya lati-
sternum and Elseya novaeguineae were similar,
but the former longer and with its cathodal line
closer to the negative pole. The electrophore-
grams of Emydura signata and Emydura sub-
globosa were similar to Elseya novaeguineae, but
the latter slightly more negative. Among
Chelodina, there were two electrophoretic pat-
terns: 3 anodal lines for C. longicollis and C.
novaeguineae; and 1 for C. expansa, C. rugosa
and C. siebenrocki. Observing little variation be-
tween Elseya and Emydura relative to that within
Chelodina, Frair (1980) recognized Elseya and
Emydura as synonymous.
Frair (1980) concluded the results of Bull &
Legler (1980) (similar chromosome numbers and
bands of Elseya and Emydura) supported their
synonymy. However the chromosomes of
Pseudemydura umbrina and Rheodytes leukops
are also similar in number and bands to Elseya
and Emydura, but Pseudemydura and Rheodytes
are separate genera because of their unique mor-
phologies.
Frair (1980) also used the observations of
Webb (1978) on basking of captive Australian
turtles to support synonymy of Elseya and
Emydura: Elseya and Emydura basked regularly,
whereas Chelodina never basked. However,
basking is prevalent in the South American short-
necked chelids Phrynops and Platemys (Lacher
et al., 1986; Monteiro & Diefenbach, 1987); in
Pelomedusidae (short-necked turtles) (Miller,
1979); and catholically in the suborder Cryp-
todira (short-necked turtles), especially the
Emydidae (Ernst & Barbour, 1989). Because
basking behavior is widespread in short-necked
turtles, it has little value in supporting synonymy
of Elseya and Emydura. Rather, it probably indi-
cates ecological differences between short-neck-
ed and long-necked turtles.
Legler (1981) examined more than 3000
specimens of Australian chelids and considered
geographic distributions to elucidate relation-
ships. He stated that Elseya dentata (and at least
3 undescribed species) and Elseya latisternum
(and at least 1 undescribed species) belonged
to separate genera, but no diagnoses were
presented.
AUSTRALIAN FRESHWATER TURTLES
Pseudemydura
steindachneri
novaeguineae
longicollis
oblonga
(Mann)
rugosa
expansa
paopao
El. latisternum
El. (Gwydir)
El. (Bellingen)
El. (Manning)
Em. subglobosa
Em. (Sleisbeck)
Em. victoriae
Em. (Daly Mission)
Em. macquarii
El. dentata
El. (Sth Alligator)
El. novaeguineae
El. (Johnstone)
El. (Burnett)
Short-necked alpha
Rheodytes
FIG. 3. The phylogeny of Australian freshwater turtles
(and Elseya novaeguineae), based on allozyme
electrophoresis (Georges & Adams, 1992). Short-
necked alpha is Elusor macrurus.
McDowell (1983) examined at least 5 spec-
imens from various localities of all Australian
short-necked turtle species, except Emydura sig-
nata. Because there were no significant differen-
ces among the characters he tested, McDowell
(1983) placed Elseya novaeguineae in synonymy
of Elseya dentata; and Emydura albertisii,
Emydura krefftii and Emydura subglobosa in
synonymy of Emydura australis. Also, his results
showed Elseya dentata (and Elseya novae-
guineae) to be more closely related to Emydura
australis than to Elseya latisternum, so he recog-
nized Elseya and Emydura as synonyms. How-
ever, the biochemical results of Georges &
Adams (1992) argue against these synonymies.
In addition, there are pronounced color differen-
ces among the nominal species of Emydura. In
life Emydura subglobosa, commonly called the
painted turtle, has a crimson plastron and yellow
on the face (Legler, 1981). Living Emydura sp.
(Daly, Nicholson and Roper Rivers) has red on
the face and legs, and Emydura krefftii yellow
across the head (Cann, 1978; Cogger, 1992). But
the significance of color pattern in diagnosing
short-necked turtles remains unknown.
On allozyme characters, Georges & Adams
(1992) could not distinguish between Emydura
325
macquarrii, E. krefftii (including Cooper Creek
and Fraser Island populations) and E. signata, all
of which they considered to be E. macquarrii.
Georges (1993) regarded E. macquarrii as
polytypic or a species complex, with unresolved
systematic positions of distinctive populations in
Cooper Creek and Fraser Island and the taxa
referred to as E. krefftii and E. signata.
Although Elseya latisternum was an outlier to
other Elseya species (and Emydura), neither Bur-
bidge et al. (1974) nor Frair (1980) suggested that
Elseya was paraphyletic. In serological studies
like Frair’s (1980), the variation of electrophore-
grams is misconstrued by multiple protein
analysis (Baverstock & Schwaner, 1985). How-
ever, monovalent techniques with antisera for
single proteins are more amenable to genetic
interpretation because the strength of cross reac-
tion is directly proportional to differences in the
gene locus encoding the protein (Baverstock &
Schwaner, 1985). Georges & Adams (1992) con-
sidered single locus protein variation and conse-
quently has the same objectivity as monovalent
immunological methods.
Georges & Adams' (1992) principal co-or-
dinates analysis of the first 3 dimensions des-
cribed 6346 of the variation. Australian chelids
clustered into 5 groups, which were recognised
by Legler (1981). Elseya novaeguineae, which
McDowell (1983) placed in synonymy of Elseya
dentata, had a divergence level similar to that of
Elusor and Rheodytes to the Elseya dentata
group. This species also lost its affinities with the
Elseya dentata group in higher dimensions.
The highest levels of divergence were within
Elseya and Chelodina; that within Emydura was
relatively small, suggesting its 5 species are rela-
tively young. Divergence between Elseya and
Emydura was less than that within Elseya. The
Elseya dentata group was closer to Emydura than
to the Elseya latisternum group, which was estab-
lished also by McDowell (1983). However, in-
stead of recognizing Elseya and Emydura as
congeneric, Georges & Adams (1992) considered
the Elseya latisternum group to comprise a
separate genus, their rationale that synonymiza-
tion would have to include also Elusor and
Rheodytes. The genetic distances between the
provisional genera (Elseya dentata group, Elseya
latisternum group, Emydura) are similar to, if not
greater than, that between many of the cryp-
todiran emydid genera, especially the batagurines
(Sites et al., 1984).
In regard to Australian long-necked Chelodina,
the phylogenetic results of Georges & Adams
326
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 2. Distribution of Australian freshwater turtles (Georges & Adams 1992) and Elseya novaeguineae.
From Cogger (1992), Ehmann (1992), Georges & Adams (1992), Iverson (1992), Legler & Georges (1993b),
Georges (1993). ?=systematic position of the population is unresolved. Zoogeographic regions from Fig. 4
sp. (Gwydir
R)
Emydura T I
(Cooper C. + l
Diamantina R.) 1
l
|
i}
D
(1992) aligned with the species groups estab-
lished by their prinicpal components analysis. In
general, their interpretation of relationships
agreed with Burbidge et al. (1974) and Legler
(1981); however, contrary to Legler (1981), the
level of divergence between the C. expansa and
C. longicollis groups did not warrant separate
generic recognition.
Georges & Adams (1992) tentatively placed
Elusor and Rheodytes outside the Elseya dentata,
Elseya latisternum and Emydura radiation (Fig.
3). Although the cladograms of Georges &
Adams (1992) were unresolved, their phenetic
analysis suggests both Elusor and Rheodytes
have their closest affinities with the Elseya den-
tata group. Similarly, the PAUP-generated
cladogram of Cann & Legler (1994), based upon
35 morphological and 16 allozyme characters
treated equally and compared to Chelodina as the
outgroup, suggests Elusor and Rheodytes form
the sister group to the Elseya dentata group.
Cann & Legler (1994) also performed multi-
discriminate analyses with 31 mensural charac-
SOUTH- WESTERN CENTRAL SOUTHEASTERN EASTERN | nommen — | NEW
WESTERN GUINEA
WESTERN EASTERN WESTERN
sp. (Liverpool
+ Mann R.)
E oe WIWeS — P
sp. (Bellingen
R.)
sp. (Manning
R.)
macquarrii |
novaeguineae
sp. (South
Alligator R.)
dentata
novae-
guineae
sp. (Johnstone
R.)
latisternum
1
! (Fraser L)
1
ters from large samples (n=41-1000) of the El-
seya dentata group, Elseya latisternum group,
Elusor, Emydura and Rheodytes. The data were
treated as percentages of carapace length. In the
resulting 3D scatter plots, the groups separated
equidistant from each other but with Emydura
separate from all. Consequently, Cann & Legler
(1994) recognized the Elseya dentata group, El-
seya latisternum group, Elusor and Rheodytes as
the Elseya group of genera.
The primary weakness of Cann & Legler's
(1994) multidiscriminate analysis is that a
species' morphology is adaptive to its environ-
ment. For example, extrapolation from Cann &
Legler (1994) would suggest that Australian
short-necked chelids are more closely related to
short-necked emydid turtles than to Australian
long-necked chelids. Also, sample groups must
be of similar sex and age classes to negate sexual
and ontogenetic differences in morphology. For
example, 9096 of Cann & Legler's (1994) sample
of Elusor macrurus were small juveniles. If this
species experiences ontogenetic changes in mor-
AUSTRALIAN FRESHWATER TURTLES
FIG. 4, The zoogeographic regions of Australian fresh-
water turtles (modified from Burbidge et al., 1974).
Species within each region are listed in Table 2.
phology, then comparison against a sample of
another species except of similar composition is
meaningless.
The relationship of Pseudemydura umbrina to
other Australian chelids remains obscure. Its en-
dangered status and small population size (Kuc-
hling & DeJose 1989) limit certain avenues of
research (e.g. Georges & Adams 1992). Nonethe-
less, P. umbrina shares several derived characters
with other Australian short-necked turtles, but
none with South American genera or Australian
long-necked Chelodina (McDowell, 1983). Thus
Pseudemydura may be the sister taxon to other
Australian short- necked turtles.
REPRODUCTIVE PATTERNS OF
AUSTRALIAN FRESHWATER TURTLES
Examining the ecology of a species can help
elucidate its evolutionary history: knowing func-
tions of structures makes the grading of character
states more accurate, and it gives insight into
selective pressures for higher or lower rates of
speciation (Shine, 1985). Legler (1985) recog-
nized 2 patterns of reproduction among Aust-
ralian chelids, one having evolved in the tropics
and the other in temperate environments.
The two reproductive patterns were defined by
egg size and hardness, time of nesting and length
327
of incubation (Legler, 1985). The tropical
strategy invoked large hard eggs being deposited
during the dry season (winter) with long incuba-
tion; and the temperate strategy, smaller flexible
eggs deposited in spring or summer with relative-
ly short incubation.
A temperate origin was indicated (Legler,
1985) for the Elseya latisternum group (E. lati-
sternum and at least 3 undescribed species),
Chelodina longicollis group (C. longicollis, C.
novaeguineae, C. steindachneri) and Emydura. A
tropical origin was indicated for the Elseya den-
tata group (E. dentata, E. novaeguineae and two
undescribed species) and C. expansa group (C.
expansa, C. oblonga, C. rugosa).
However, the two patterns are not realistic and
the criterion for egg hardness was only subjec-
tively defined. For example, the C. expansa and
E. dentata groups lay the largest eggs (Legler,
1985), but these species are the largest Australian
freshwater turtles. Also, larger eggs may neces-
sarily be harder because of structural constraints.
In cryptodirans there is positive correlation be-
tween adult size and egg size; egg size and hatc-
hling size; and for hard shelled eggs, egg size and
shell thickness (Ewert, 1979). Also, some tropical
species lay soft shelled eggs, and some both hard
and soft (Moll & Legler, 1971; Ewert, 1979).
Legler (1985) was selective of data from pre-
vious authors, ignoring that which lessened dis-
tinction between the two patterns. For example,
C. oblonga (proposed tropical origin) nests
during summer (Burbidge, 1967, 1984) and C.
expansa (proposed tropical origin) sometimes
during autumn (Goode & Russell, 1968; Georges,
1984); and eggs of C. longicollis (proposed
temperate origin) undergo long incubation, up to
200 days (Cann, 1978). Thus it appears that addi-
tional reproductive patterns exist. Also, if the
contrasting tropical and temperate environments
are strong selective pressures on reproduction,
why do some species of each group inhabit the
other environment? Within Emydura (proposed
temperate origin), E. subglobosa, E. victoriae
and 2 undescribed species inhabit only the
tropics; and E. macquarrii (inclusive of E.
australis, E. krefftii and E. signata) inhabits both
temperate and tropical environments (Cogger,
1992; Iverson, 1992). Georges et al. (1993) pre-
viously questioned the fit of Legler's (1985)
reproductive patterns to species groups.
However, according to the centre of diversity
rule (Ross, 1974), Emydura probably evolved in
the tropics, and also the Elseya dentata, C. expan-
sa (C. expansa, C. rugosa and an undescribed
328
species in the NT, sensu Georges & Adams
[1992]) and C. longicollis groups. With 3 species
in the temperate zone and Elseya latisternum
inhabiting both temperate and tropical zones, the
E. latisternum group probably evolved in temp-
erate environments. In addition, the groups for
which the centre of diversity rule predicts tropical
origins all have representatives and/or closely
related species in New Guinea, dispersal north-
ward having been possible by Pleistocene land-
bridges (Burbidge et al., 1974). On the other
hand, the Elseya latisternum group (predicted
temperate origin), with E. latisternum ranging
from northern NSW (temperate) to the tip of Cape
York Peninsula, has no close relative in New
Guinea.
ZOOGEOGRAPHY OF AUSTRALIAN
FRESHWATER TURTLES
Zoogeographic regions for Australian fresh-
water turtles were defined by Burbidge et al.
(1974), which in general corresponded to those
for freshwater fish (Whitley, 1959). However, we
also recognize Central and Eastern zoogeo-
graphic regions for freshwater turtles (Fig. 4,
Table 2). The Central Region is in the arid zone
of interior Australia and contains the Cooper
Creek and Diamantina River population of large
macrocephalous Emydura, which Ehmann
(1992) and Legler & Georges (1993b) reported to
be an undescribed species. Although Georges &
Adams (1992) could not distinguish the popula-
tion by allozyme electrophoresis from E. mac-
quarrii, Georges (1993) regarded it as a dis-
tinctive population with unresolved systematic
position. The Eastern Region encompasses the
Fitzroy, Burnett, Mary and Brisbane river drain-
ages, to which are endemic only an undescribed
species of the Elseya dentata group (Georges &
Adams, 1992), Elusor macrurus and Rheodytes
leukops. Our Central and Eastern regions cor-
respond to Whitley's (1959) Sturtian and Kref-
ftian regions, respectively. Zoogeographic
regions for freshwater turtles are almost identical
to those for freshwater fish. Also, like some fresh-
water fish, some freshwater turtles inhabit both
northern Australia and southern New Guinea.
The vicariance hypothesis of Burbidge et al.
(1974) suggests that ancestors of each species
group were distributed widely throughout north-
ern and eastern Australia during the Pleistocene
epoch when the climate was cooler and wetter.
The existing chelid fauna resulted by allopatric
MEMOIRS OF THE QUEENSLAND MUSEUM
speciation, directed by isolation of river drain-
ages with increasing aridification.
Southwestern Australia is inhabited by en-
demic Chelodina oblonga and Pseudemydura
umbrina. This region became isolated by forma-
tion of a broad inland sea from the Gulf of Car-
pentaria to the coast of S AUST during the Early
Cretaceous (Rich & Rich, 1989; Fig. 2). Burbidge
et al. (1974) proposed that the eastern Chelodina
ancestor evolved into the form lacking neural
bones, and the eastern Pseudemydura became
extinct. Prior to the Cretaceous sea, Elseya and
Emydura had tropical northern distributions.
Later during the Cretaceous the sea receded, leav-
ing a vast waterless desert equally impenetrable.
Alternatively, Pritchard & Trebbau (1984) sug-
gested two separate invasions from Antarctica,
one each into eastern and western Australia.
The distributions of freshwater turtle and fish
species groups support the vicariance hypothesis
of Burbidge et al. (1974). The species within each
group are largely allopatric, with sympatry be-
tween species of different groups (Legler, 1981;
Iverson, 1992). In addition, there are fossils and
a relict population of Emydura in the arid zone of
SW QLD and adjacent S AUST (Gaffney, 1979;
Ehmann, 1992); and fossils of Pseudemydura
(early to middle Miocene) from Riversleigh, on
the Gulf of Carpentaria (Gaffney et al., 1989).
Burbidge et al. (1974) showed C. oblonga to be
morphologically and serologically an outlier to
other Chelodina. Similarly, the phenetic analysis
(3D) of Georges & Adams (1992) showed C.
oblonga to be an outlier within the C. expansa
group (although this was not maintained at higher
dimensions), and their phylogenetic analyses
consistently placed C. oblonga as pleisiomor-
phic. These results together with retention of
well- developed neural bones (Burbidge et al.,
1974) indicate that C. oblonga approaches the
ancestral condition, further supporting Bur-
bidge's et al. (1974) vicariance hypothesis.
The hypothesis of Burbidge et al. (1974) re-
quires the short-necked turtle genera to have a
northern distribution prior to formation of the
inland Cretaceous sea. However, the Elseya
latisternum group probably had a southeastern
origin, and thus potential for dispersal into south-
western Australia. The following hypotheses
may explain the absence of the Elseya latister-
num group from southwestern Australia. (1) The
E. latisternum lineage previously inhabited also
southwestern Australia but has since become ex-
tinct there. (2) The southern Australian environ-
ment during the Cretaceous was not favorable for
AUSTRALIAN FRESHWATER TURTLES
dispersal of the E. latisternum lineage westward
into southwestern Australia. (3) The E. latister-
num lineage evolved after the Cretaceous, and
since then the southern Australian environment
has not been favorable for dispersal westward.
Hypotheses (2) and (3) appear most likely be-
cause members of the E. latisternum group live
in the headwaters of rivers, even above major
waterfalls in the Great Dividing Range of eastern
Australia (Legler, 1981). Low flat land, instead
of elevated ranges, between SE and SW Australia
may have been a barrier to dispersal of this group.
On the other hand, if Emydura had a temperate
distribution prior to formation of the inland
Cretaceous sea, the genus could have dispersed
westward into southwestern Australia.
Rheodytes leukops Legler & Cann (1980) and
Elusormacrurus Cann & Legler (1994) belong to
monotypic genera and with highly restricted ran-
ges. Rheodytes leukops is endemic to headwaters
of the Fitzroy River, E of the Great Dividing
Range in central eastern Queensland. The species
is highly specialised for inhabiting rapids: it feeds
on invertebrates in rapids, and it can obtain all of
its oxygen through its enlarged cloacal bursae
(Legler & Georges, 1993b). The other short-
necked turtles feed in slow-moving or standing
water (Legler & Georges, 1993b), and their cloa-
cal bursae are less developed. Elusor macrurus is
endemic to the Mary River drainage of SE QLD,
also E of the Great Dividing Range, and its biol-
ogy is poorly known (Cann & Legler, 1994)
CONCLUDING STATEMENT
Although we now have a better understanding
of Australian chelids, 1 genus and 9 extant
species await description (Georges & Adams,
1992), and the systematic positions of several
populations of Emydura remain unresolved:
Cooper Creek, Fraser Island, and the taxa referred
to as E. krefftii and E. signata. Intergeneric
relationships remain enigmatic. However, it ap-
pears that Pseudemydura umbrina shares com-
mon ancestry with other Australian short-necked
turtles and is their sister taxon. Also, it appears
the Elseya latisternum group is the sister taxon of
the non-Pseudemydura short-necked turtles, as
suggested by data from Georges & Adams (1992)
and Cann & Legler (1994).
ACKNOWLEDGEMENTS
We are especially grateful for the assistance of
Gordon C. Grigg, Barne G.M. Jamieson and
329
Colin J. Limpus. The following persons also
engaged in discussion and/or read and com-
mented on the manuscript: David T. Booth, An-
gela Chapman, Christine James, Ralph E.
Molnar, Craig Moritz and R. Anthony Thulborn.
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PERINATAL SKELETAL INJURIES IN TWO BALAENOPTERID WHALES
ROBERT A. PATERSON AND STEPHEN M. VAN DYCK
Paterson, R.A. & Van Dyck, S.M. 1996 07 20: Perinatal skeletal injuries in two balaenopterid
whales. Memoirs of the Queensland Museum 39(2): 333-337. Brisbane. ISSN 0079-8835.
Two recently born balaenopterid whales (Balaenoptera acutorostrata and Megaptera
novaeangliae) stranded on the coast of southern Queensland exhibited similar pericranial
andrib lesions considered to result from compression injury. Birth trauma, perhaps associated
with caudal presentation, is considered the most likely cause of the lesions. L] Balaenopterid
whales, skeletal injuries, birth trauma.
Robert A. Paterson & Stephen M. Van Dyck, Queensland Museum, P.O. Box 3300, South
Brisbane, Queensland 4101, Australia; received 22 February 1996.
The Queensland Museum (QM) cetacean col-
lection contains skeletal material of 11 juvenile
balaenopterids of which 7 are minke whales
Balaenoptera acutorostrata, 3 are humpback
whales Megaptera novaeangliae and | a blue
whale B.musculus musculus. Among these,
pericranial and rib lesions are evident in 2 very
immature specimens, 1 minke whale and 1
humpback whale. This paper describes the
pathology and speculates on its cause.
x
DESCRIPTIONS
QMJM7301. A 2.9 m long 2 minke whale
stranded at the Big Sand Hill, Moreton Island
(27°13’S, 153°22’E) on 11.vi.87. Its flipper and
body colouration was typical of the diminutive or
Type 3 form (Best, 1985) and was illustrated in
Paterson (1994). The umbilicus was healed. Su-
perficial ‘cookie-cutter’ lesions similar to those
described in other examples of this species (Wil-
liamson, 1975; Arnold et al., 1987) were noted as
FIG. 1. Minke whale QMJM7301. Radiograph demonstrating numerous bilateral ventral rib fractures. The
sternum has not been included.
a
well as predatory rakes near the dorsal fin.
Bilateral rib fractures were palpable but there
were no wounds or scars superficial to them.
Abundant callus (some was lost during prepara-
tion) was evident both by direct inspection and
radiological examination (Fig. 1). After prepara-
tion of the skull a raised area of shell-like ossifica-
tion 6.0 cm long and 2.5 cm in greatest diameter
was noted on the lateral aspect of the supra-orbital
process of the left frontal (Fig. 2).
QM JM7303. A 4.2 m long d humpback whale
stranded at Moon Point, Fraser Island (25°14’S,
153°00’E) on 17.10.89. It was frozen soon after
death and transported to the QM. The umbilicus
was healed. Numerous bites and rakes, con-
sidered to be due to shark attack, were noted.
They included a large fresh right axillary wound.
Rib fractures (Fig. 3) were not as severe as those
in QMJM7301. During dissection a cystic
pericranial lesion measuring 16.0 cm long and 7.0
cm in greatest diameter was noted on the lateral
aspect of the supra-orbital process of the right
frontal (Fig. 4). There were no soft tissue or
cutaneous abnormalities superficial to the lesion.
Its raised periosteal edge was biopsied and his-
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 2. Minke whale QMJM7301 (Left). Dorsal view
of skull (98cm long) demonstrating periosteal new
bone formation on the lateral aspect of the supra-or-
bital process of the left frontal bone. (Right) Close-up
profile view of the same region.
tological examination (Fig. 5) demonstrated
periosteal new bone formation superficial to a
sub-periosteal cyst consistent with trauma several
weeks prior to death (J. Musgrave pers.comm.).
DISCUSSION
There are very limited data concerning the time
and place of birth of Type 3 minke whales. The
two smallest examined in South Africa by Best
(1985) were 1.92 m and 2.54 m long and they
stranded at latitude 34°S in May and July respec-
tively. The former had a raw and completely
unhealed umbilicus and was considered to be a
very recent live birth (Best, 1985). QMJM7301
was 2.9 m long, and stranding occurred at 27°S in
June. Its skeleton was extremely immature com-
pared with the 6 other juvenile minke whale
skeletons in the QM collection and this suggests
a very young age, probably less than 8 weeks.
There are extensive data concerning the time
and place of birth of Southern Hemisphere
humpback whales. The modal length at birth is
4.3 m (Chittleborough, 1958) and most births
occur in the vicinity of latitude 20°S between late
July and mid September (Townsend, 1935; Chit-
tleborough, 1965; Paterson & Paterson, 1989).
PERINATAL SKELETAL INJURIES IN BALAENOPTERID WHALES
335
FIG. 3. Humpback whale QMJM7303. Radiograph demonstrating healing undisplaced bilateral ventral rib
fractures. The sternum has not been included. (The vertical ‘tracks’ in some ribs on the left of the photograph
result from drilling to insert numbering wires).
QMJM7303 was 4.2 m long and stranded at
25°S in October on the western side of Fraser
Island, a region frequented by humpback whales
during the southern migration (Corkeron et al.,
1994). QMJM7303 was probably less than 6
weeks old when it died.
Radiological opinion was sought in an attempt
to date the rib fractures. They were considered to
have occurred approximately 6-8 weeks before
death and were likely to have resulted from com-
pression and not from blunt trauma (J.P. Masel,
pers. comm.) The position and extent of the
bilateral rib fractures are consistent with com-
pression injury described in human neonates by
Caffey (1973). The extensive fractures in
QMJM7301 are considered to represent a ‘central
flail’ a term used in human trauma when multiple
rib fractures occur on both sides of the sternum
(Hunt & Schwab, 1992). Such an injury in
humans is often life-threatening and may require
assisted ventilation. However, some healing was
evident in the fractures of QMJM7301 and its
death was not due to acute chest trauma.
The lateral aspect of the supra-orbital process
of the frontal would be susceptible to injury,
particularly in an immature animal, if dorsolateral
compression occurred. Pericranial injury, similar
to that in QMJM7301 and QMJM7303, occurs in
c.2% of human neonates and is termed cephal-
haematoma. It usually results from cranial
moulding during parturition and, as an isolated
finding, is not associated with mortality or per-
sistent morbidity (Caffey, 1973). The degree of
new bone formation in the pericranial lesion of
QMJM7301 (Fig.2) suggests a longer period of
survival than QMJM7303 in which new bone
FIG. 4. Humpback whale QMJM7303. Dorsal view of
skull (which measures 1 13cm in length) demonstrat-
ing periosteal new bone formation on the lateral
aspect of the supra-orbital process of the right frontal
bone.
formation was limited to the edge of the lesion
(Fig. 4).
Most data concerning foetal presentation and
the mechanism of parturition in cetaceans derive
from studies of captive odontocetes. Caudal
presentation is usual (McBride & Kritzler, 1951;
Slijper, 1962), a presentation considered disad-
vantageous in large domestic animals. Arthur
(1964) noted that in such animals the foetus is
wedge-shaped when the presentation is cephalic
and this serves to progressively dilate the birth
canal during its passage, whereas in caudal
presentation compression of the foetal abdomen
causes expansion of the ribs and the costal arch
engages abruptly. Also, the foetal occiput, often
MEMOIRS OF THE QUEENSLAND MUSEUM
the broadest foetal part, makes an abrupt [pelvic]
engagement. Caudal presentation has been noted
in a humpback whale (Dunstan, 1957) and such
presentation in mysticetes could be associated on
occasions with the difficulties described by Ar-
thur (1964), although the pelvic structures of
cetaceans and large terrestrial animals differ con-
siderably. Hartley (1983) listed contusions to the
cranial periosteum and rib fractures among the
postmortem findings in a large series of foal
perinatal mortalities.
We conclude on the basis of available evidence
that the pericranial and rib injuries in QMJM7301
and QMJM7303 were sustained during parturi-
tion. However, it remains conjectural if they con-
tributed to the premature deaths of these whales.
ACKNOWLEDGEMENTS
We received valuable assistance from many
persons. Patricia Paterson assisted with the dis-
section and retrieval of QMJM7301. Vic Hislop
retrieved QMJM7303, arranged its freezing, and
transported it to the QM. John Masel, Director of
Metropolitan Paediatric Radiology in Brisbane,
dated the rib fractures and John Musgrave of
Sullivan, Nicolaides and Partners prepared and
diagnosed the histological specimens from
QMJM7303. Deidre Pyecroft gave us the benefit
of her veterinary experience and searched the
relevant literature. Bruce Cowell and Jeff Wright
of the QM prepared the photographs. Sophie
Kupis and Stephen Marmo, of the Jindalee Medi-
cal Centre and Royal Brisbane Hospital respec-
tively, took the radiographs.
LITERATURE CITED
ARNOLD, P., MARSH, H. & HEINSOHN, G. 1987.
The occurrence of two forms of minke whales in
east Australian waters with a description of exter-
nal characters and skeleton of the diminutive or
dwarf form. Scientific Reports of the Whales Re-
search Institute, Tokyo 38: 1-46.
ARTHUR, G.H. 1964. ‘Wright’s veterinary obstetrics
(including diseases of reproduction)'. (Bailliere,
Tindall & Cox : London).
BEST, P.B. 1985. External characters of southern minke
whales and the existence of a diminutive form.
Scientific Reports of the Whales Research In-
stitute, Tokyo 36: 1-33.
CAFFEY, J. 1973. ‘Paediatric X-ray diagnosis’. (Year
Book Publishers : New York).
CHITTLEBOROUGH, R.G. 1958. Breeding cycle of
the female humpback whale Megaptera nodosa
(Bonnaterre). Australian Journal of Marine and
Freshwater Research 9: 1-18.
PERINATAL SKELETAL INJURIES IN BALAENOPTERID WHALES
337
FIG. 5. Humpback whale QMJM7303. Histological section (x 100) from edge of supra-orbital lesion demonstrated
in Fig. 4. Thickened periosteum (A) is seen superiorly; spicules of vertically arranged osteoid seams (B) with
intervening connective tissue are seen centrally and a layer of fibrous tissue (C) lines a sub-periosteal cyst (D)
at the lower left of the photograph. The appearances are those of periosteal separation with some new bone
formation superficial to a post-traumatic sub-periosteal cyst.
1965. Dynamics of two populations of the
humpback whale Megaptera novaeangliae
(Borowski). Australian Journal of Marine and
Freshwater Research 16; 33-128.
CORKERON, P.J., BROWN, M., SLADE, R.W &
BRYDEN, M.M. 1994, Humpback whales,
Megaptera novaeangliae (Cetacea : Balaenop-
teridae), in Hervey Bay, Queensland. Wildlife
Research. 21; 293-305,
DUNSTAN, D.J. 1957. Caudal presentation at birth of
a humpback whale, Megaptera nodosa (Bon-
naterre). Norsk Hvalfangst-Tidende. 46: 553-555.
HARTLEY, W.J. 1983. Some aspects of foal mortality
in New South Wales. Proceedings postgraduate
committee in veterinary science, University of
Sydney. 65: 207-212.
HUNT, D.M. & SCHWAB, F.J. 1992, Chest trauma. Pp
77-100. In Rosen, P., Doris, P.E., Barkin, R.M.,
Barkin, S.Z. & Markovchick, V.J. (eds), ‘Diag-
nostic radiology in emergency medicine'. (Mosby
: St. Louis).
MCBRIDE, A.E. & KRITZLER, H. 1951. Observa-
tions of pregnancy, parturition and post-natal be-
haviour in the bottlenose dolphin. Journal of
Mammalogy. 32(3): 251-266.
PATERSON, R.A. 1994, An annotated list of recent
additions to the cetacean collection in the
Queensland Museum. Memoirs of the Queensland
Museum. 35(1) : 217-233.
PATERSON, R. & PATERSON, P. 1989. The status of
the recovering stock of humpback whales Megap-
tera novaeangliae in east Australian waters.
Biological Conservation. 47: 33-48.
SLIJPER, E.J. 1962. ‘Whales’. (Hutchinson : London).
TOWNSEND, C.H. 1935. The distribution of certain
whales as shown by log book records of American
whaleships. Zoologica 19: 1-50.
WILLIAMSON, G.R. 1975. Minke whales off Brazil.
Scientific Reports of the Whales Research In-
stitute, Tokyo 27: 37-59,
338
SURVIVAL OF A LARGE CROCODYLUS POROSUS
DESPITE SIGNIFICANT LOWER JAW LOSS. Memoirs
of the Queensland Museum 39(2): 338. 1996:- On 25 May,
1995 a large (SVL 195.5cm; TL 424.5cm), male crocodile
(Crocodylusporosus) was caught in the Nicholson River, NW
Qld (17°44’S, 139?31'E). It was captured in a mesh trap, set
under the supervision of the Department of Environment and
Heritage. The crocodile had attacked and killed a dog. A
significant portion of its dentary, including the symphysis to
the region of the 10th and 11th tooth alveoli, had been lost.
Theright partial dentary, which closed first, was 7.5cm shorter
than the left, resulting in an uneven closing of the jaws. The
tongue had also been partially amputated, but had healed with
a large egg-shaped growth at the anterior edge. This was
apparently an area of scar tissue resulting from the trauma or
from the inclusion of a bone fragment in surrounding tissue.
The tongue appeared to be hypertrophied, probably as a result
of its continued involvement in the closing and manipulation
of the two dentary fragments. Rather than a normal, flat
appearance, buccal edges were greatly enlarged, giving the
tongue a pillowed appearance. This crocodile had sustained
several other injuries, including scarring on the trunk in front
of the hind limbs and amputation of the first and second
metatarsals of the right foot. Many teeth were also missing
from both lower jaws. (It was not possible to ascertain whether
these had fallen out with age or wear, or had been broken
during aggressive encounters with other crocodiles).
A wide variety of abnormalities and injuries have been
recorded amongst crocodiles (lordansky, 1973). Some studies
of injuries have focussed on particular species (e.g.
Crocodylus niloticus by Cott, 1961; C. johnstoni by Webb &
Manolis, 1983 and C. porosus by Webb & Messel, 1977).
Many injuries sustained by crocodiles result from conflicts
related to social behaviour and territoriality. There have been
few studies of large crocodiles in the wild. Of 1345 specimens
ofC. porosus examined in one study, only 10 had snout- vent
lengths greater than 150cm because, in wild populations,
juveniles predominate (Webb & Messel, 1977). The least
commonly recorded injuries in large crocodiles are those to
the head. They account for only 25% of scarring in large
crocodiles (TL >150cm, Webb & Messel, 1977). This is not
surprising, because many crocodiles sustaining significant
head injuries would die. One example (Webb & Manolis,
1989) was found dead, having lost the dorsal portion of its
snout during combat.
H 61
LDF 43.5
MEMOIRS OF THE QUEENSLAND MUSEUM
The injuries sustained by the Nicholson R. crocodile were
significant (Fig. 1). No estimate of when the jaw loss occurred
can be made. The specimen was in good condition, with large
fat reserves at the base of the tail and neck. It had been known
in the area by locals, because of his distinctive head, for at
least 18 years. Despite this, there had been no reports of a
damaged lower jaw. The crocodile apparently survived large-
ly by exploiting a cattle station rubbish dump where cattle
offal, road-killed wallabies and feral pig carcasses were
dumped. To do so, it walked close to 50m from the river. It
would locate a carcass by smell, and go straight to it, regard-
less of the difficulty of the terrain. The crocodile would grasp
the carcass in its jaws and return to the water by the easiest
path, to feed. Abundant tracks indicated that this crocodile
actively moved about on land several nights each month.
I thank Terri, Bob and Lyn Irwin; Dave and Jenny Hansen;
Noel Oliver of Escott Station; Brad Jones; Lee Pang and
Roger Bilney, for their assistance in compiling this informa-
tion. Jeanette Covacevich, Queensland Museum helped
prepare it for publication.
Literature Cited
Cott, H.B. 1961. Scientific results of an inquiry into the
ecology and economic status of the Nile crocodile
Crocodylus niloticus in Uganda and northern Rhodesia.
Transactions of the Zoological Society of London 29:
211-337.
Glasby, C.J., Ross, G.J.B. & Beesley, P.L. (eds) 1993. Fauna
of Australia. Vol. 2A. Amphibia & Reptilia. (Australian
Government Publishing Service : Canberra).
lordansky, N.N. 1973. The skull of the Crocodilia. Pp. 201-
262. In C. Gans & T.S. Parsons, (eds). Biology of the
Reptilia. Vol. 4. (Academic Press: New York).
Webb, G.J.W. & Manolis, S.C. 1983. Crocodylus johnstoni in
McKinlay River Area, N.T. V. Abnormalities and In-
juries. Australian Wildlife Research 10, 407-420.
1989. Crocodiles of Australia. (Reed: Sydney).
Webb, G.J.W. & Messel, H. 1977. Abnormalities and injuries
in the esturarine crocodile, Crocodylus porosus.
Australian Wildlife Research 4: 311-319
Stephen Irwin, Queensland Reptile Park, Glasshouse Moun-
tains Tourist Drive, Beerwah, Queensland 4519, Australia;
23 february 1996.
FIG. 1. Lateral view of injury with details of measurements taken at time of capture. Head (H). Left dentary
fragment (LDF). Right dentary fragment (RDF). All measurements in centimetres
FOUR NEW RECORDS OF SURGEONFISHES (PERCIFORMES: ACANTHURIDAE)
FROM THE GREAT BARRIER REEF
KENDALL D. CLEMENTS AND JOHN E. RANDALL
Clements, K.D. & Randall, J.E. 1996 07 20: Four new records of surgeonfishes (Perciformes:
Acanthuridae) from the Great Barrier Reef. Memoirs of the Queensland Museum 39(2):
339-342. Brisbane. ISSN 0079-8835.
Four species of surgeonfishes are recorded from the outer Great Barrier Reef for the first
time. They are: Acanthurus maculiceps (Ahl), Naso minor (Smith), Naso thynnoides (Valen-
ciennes) and Ctenochaetus tominiensis Randall. C] Great Barrier Reef, surgeonfishes.
K.D. Clements, School of Biological Sciences, University of Auckland, Private Bag 92019,
Auckland, New Zealand; J.E, Randall, Bishop Museum, PO Box 19000A, Honolulu, Hawaii
96817-0916; received 3 May 1996.
Four species of surgeonfishes new to Australia
were found along the outer shelf reefs of the
northern Great Barrier Reef(GBR): Acanthurus
maculiceps (Ahl), Naso minor (Smith), N. thyn-
noides (Valenciennes), and Ctenochaetus
tominiensis Randall. All except N. minor occur
around New Guinea (Allen & Swainston 1993;
Randall 1994). Prior to this report, 36 species of
surgeonfishes were recorded from the GBR and
Coral Sea (Randall et al. 1990). The Acan-
thuridae (surgeonfishes) consists of 73 species,
mostly from the Indo-Pacific region. The most
distinctive feature of the family is the sharp spine
or spines on the caudal peduncle.
SYSTEMATICS
Material is housed in the Australian Museum
(AMS), and Bernice P. Bishop Museum (BPBM).
Family ACANTHURIDAE
Acanthurus maculiceps (Ahl, 1923)
(Fig. 1)
Hepatus maculiceps Ahl,
Talassia, New Britain).
MATERIAL. AMSI37145-001, 213mm SL, No Name
Reef (Reef 14-139), 14?38' S, 145°38’E, dense coral
growth near reef crest, adjacent to the reef pass, 4m,
spear, K.D. Clements, 16 November 1995.
DIAGNOSIS. Dorsal rays IX,25; anal rays III,
23; pectoral rays 17 (including rudimentary up-
permost ray); anterior gill rakers 20; posterior gill
rakers 22; 14 upper and 18 lower teeth; body
moderately deep for genus, depth 2.1 in SL, com-
pressed, width 2.95 in depth; head length 3.3 in
SL; dorsal profile of head strongly convex; snout
length 4.3 in SL; orbit diameter 4.8 in head
1923:36, (type locality,
length; caudal peduncle depth 8.45 in SL; ninth
dorsal spine longest, 6.35 in SL; caudal fin lunate,
the caudal concavity 5.3 in SL; caudal spine 3.0
in head length. Stomach thickwalled. Colour in
alcohol: dark brown with a horizontal black band
(narrowing at each end) at upper end of gill
opening from behind eye to 1.5 orbit diameters
posterior to gill opening; a narrow black margin
at edge of caudal spine socket; numerous faint
small pale spots on head; a transverse whitish
band on chin; dorsal fin with 9 narrow pale stripes
alternating with dark brown, and a narrow black
margin; anal, caudal, and pelvic fins dark brown;
pectoral fins dark brown, with a pale spot distally
on upper half.
REMARKS. This species is known from Christ-
mas Island (Indian Ocean), Ryukyus, Philippines,
Indonesia, Palau, Marianas, Marshall Islands,
Kiribati, Tuvalu, Samoa, Line Islands, and Caro-
line Islands (Randall, 1956; Myers, 1989; Jones
et al., 1993. It is distinguished from other sur-
geonfish in colour by a combination of the white
caudal spine with a black margin, the patch of
yellow on the distal third of the pectoral fin, the
black lozenge-shaped mark extending backwards
from the upper edge of the operculum, numerous
small pale yellow spots on the head, and an ir-
regular white bar at the base of the caudal fin.
Ctenochaetus tominiensis Randall, 1955
(Fig. 2)
Ctenochaetus tominiensis Randall, 1955: 163. (type
locality, Gulf of Tomini, Sulawesi).
MATERIAL. BPBM14726, 64.5mm SL, NW side of
horseshoe-shaped reef at S boundary of Pandora
entrance, 11?30'S, 144°00’E, rich coral bottom, 14m,
multiprong spear, R.C. Steene, 20 November 1972.
DIAGNOSIS. Dorsal rays VIII,26; anal rays
340 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 1. Acanthurus maculiceps (Ahl,1923), AMSI37145-001, No Name Reef, Great Barrier Reef.
FIG. 2. Ctenochaetus tominiensis Randall, 1955, BPBM14726, Pandora Reef, Great Barrier Reef.
IIL24; pectoral rays 16 (including short splint- expanded incurved tips, 25 in upper jaw and 28
like upper ray); anterior gill rakers 20; posterior in lower; distal half of expanded part of upper
gill rakers 21; teeth slender and flexible with teeth smooth-edged, lower half with 3 denticula-
NEW SURGEONFISH RECORDS, GREAT BARRIER REEF
tions; expanded outer part of lower teeth with 3
denticulations (including the tip, much the
broadest); edges of lips strongly papillate; body
very deep, depth 1.75 in SL, strongly compressed,
width 3.1 in depth; head length 3.0 in SL; snout
length 4.2 in SL; orbit diameter 5.4 in head
length; caudal peduncle depth 9.1 in SL; eighth
dorsal spine longest, 5.5 in head length; caudal
fin lunate, the caudal concavity 4.2 in SL. Colour
in alcohol: head, body and paired fins brown;
caudal fin light brown (may be whitish in life);
spinous portion of dorsal fin brown, with faint
curving oblique pale bands alternating with
brown bands, becoming horizontal in outer
posterior part of soft portion of fin; soft portion
of dorsal and anal fins brown with a broad distal
pale zone (bright orange in life) except for a
narrow black margin; a black spot at rear base of
dorsal fin and a less distinct blackish one at rear
base of anal fin; anterior nostril white.
REMARKS. This species is known from In-
donesia, Philippines, PNG, and the Solomon Is-
lands. The dorsal and anal softray counts of the
GBR specimen are higherthan the recorded range
(Randall, 1955, table 1), but fin-ray counts of
BPBM material are equally high. C. tominiensis
is distinguished within the genus by the broad
orange borders of the soft dorsal and anal fins.
Naso (Axinurus) minor (Smith, 1966)
(Fig.3)
Axinurus minor Smith, 1966:638 (type locality, Pinda
Reef, Mozambique).
DIAGNOSIS. See Randall, 1994.
REMARKS. Underwater photographs of this
species were taken on the outer reef slope at the
north end of No Name Reef (Reef 14-139:
14?38'S, 145?38' E) in 20-30m on 18/1/95, and at
the south end of the same reef in 25-30m on
23/1/1995 (Fig. 3). Schools of N. minor were
Observed at the N end of Hick's Reef (Reef 14-
086: 14°26’S, 145728" E), in 15-25m on 24/1/95.
These observations involved schools of 15-20
individuals moving rapidly over the shelf at the
base of the escarpment. The schools appeared to
move up onto this shelf from deeper water. N.
minor is common on the seaward side of outer
shelf reefs in the northern section of the Great
Barrier Reef. The lack of any previous Australian
record is probably due to the limited diving on
these deepslope habitats, and because Naso minor
is difficult to approach underwater. We could not
collect specimens by spear and our photographs
could only be taken from a distance. This species
is known from the Philippines, Indonesia,
Reunion, and Mozambique (Randall, 1994).
Naso (Axinurus) has a single fixed keel-like
spine on each side of the caudal peduncle (Ran-
dall, 1994). Other species of Naso have two fixed
caudal spines. Naso minor may be distinguished
within the genus by the combination of a single
black caudal spine and a bright yellow caudal fin
and by the male frequently adopting a distinct
countershading pattern (Randall, 1994, pl. 2B).
Naso thynnoides (Valenciennes, 1835)
(Fig.4)
Axinurus thynnoides Cuvier, 1829: 225 (nomen
nudum). Valenciennes in Cuvier and Valenciennes,
1835:299, pl. 293 (type locality, Dorey Harbour,
New Guinea).
DIAGNOSIS. See Randall, 1994.
REMARKS. Underwater photographs of this
species were taken at the north end of Hick's Reef
FIG. 3. Underwater photograph of Naso minor (Smith, 1966), No Name Reef, Great Barrier Reef.
(Reef 14-086: 14°26’S, 145?28' E) in 15-25m on
24/1/95 and 25/1/1995. On both occasions a
group of approximately 6 individuals were
moving rapidly over the slope. On the first oc-
casion these fish were schooling with a much
larger group of N. minor. Attempts to collect
specimens by spear were unsuccessful. The N end
of Hick's Reef is unusual for outer reefs in the
vicinity as the shelf between the base of the
escarpment and the outer dropoff is relatively
shallow (cf. 20-35 m in other places). We have
not seen N. thynnoides elsewhere on the GBR.
N. thynnoides may be distinguished within the
genus by a single caudal spine and a series of
narrow, dark bars on the body (Randall 1994).
The caudal fin in the specimens of N. thynnoides
in our photos is grey, not distinctively yellow as
in N. minor or blue as in N. caeruleacauda (Ran-
dall 1994). Colour of N. thynnoides photographed
at Hick's Reef is very similar to Randall, 1994,
pl.1A. Itis known from the western Indian Ocean,
Philippines, Amami-Oshima Islands, Ryukyu Is-
lands, Indonesia, Solomon Islands, Tuvalu,
Caroline Islands, and Maldive Islands (Jones et
al., 1991; Randall, 1994).
ACKNOWLEDGEMENTS
We thank the Lizard Island Research Station
for facilities, Australian Museum for loan of
specimens, Lynda Axe for help in the field, Roger
C. Steene for the specimen and photograph of C.
tominiensis, Ian MacDonald for help with
photography, and Howard Choat for logistical
support and helpful comments. KDC was sup-
ported by an ARC Postdoctoral Fellowship and
a Special Investigator Award to Howard Choat.
MEMOIRS OF THE QUEENSLAND MUSEUM
LITERATURE CITED
AHL, E. 1923. Ichthyologische Mitteilungen 3. Ueber
eine neue Art der Gattung Hepatus. Zoologisches
Museum Berlin, Mitteilung, 11: 36-37.
ALLEN, G.R. & SWAINSTON, R. 1993. Reef fishes
of New Guinea. (Christensen Research Institute:
Madang).
CUVIER, G. 1829. Le règne animal distribué d’après
son organisation, pour servir de base à l'histoire
naturelle des animaux et d'introduction à
l'anatomie comparée, ed. 2, 2. ( Déterville: Paris).
CUVIER, G. & VALENCIENNES, M.A. 1835. His-
toire naturelle des poissons. 10. (Chez F.G. Lev-
rault: Paris).
JONES, G.P., KALY, U.L. & CLEMENTS, K.D. 1991.
Preliminary records of the coral reef fishes of
Tuvalu. South Pacific Journal of Natural Science
11: 40-57.
KUITER, R.H. 1992. Tropical reeffishes of the Western
Pacific: Indonesia and adjacent waters. (Penerbit
PT Gramedia Pustaka Utama: Jakarta).
MYERS, R.F. 1989, Micronesian reef fishes. (Coral
Graphics: Guam).
RANDALL, J.E. 1955. A revision of the surgeon fish
genus Ctenochaetus, Family Acanthuridae, with
descriptions of five new species. Zoologica 40:
149-168.
1956. A revision of the surgeon fish genus Acan-
thurus. Pacific Science 10: 159-235.
1994, Unicornfishes of the subgenus Axinurus (Per-
ciformes: Acanthuridae: Naso), with description
of a new species. Copeia 1994: 116-124.
RANDALL, J.E, ALLEN, G.R. & STEENE, R.C.
1990. Fishes of the Great Barrier Reef and Coral
Sea. (Crawford House Press: Bathurst).
SMITH, J.L.B. 1966. Fishes of the subfamily Nasinae
with a synopsis of the Prionurinae. Ichthyological
Bulletin of the Department of Ichthyology,
Rhodes University, Grahamstown 32: 634-682.
FIG. 4. Underwater photograph of Naso thynnoides (Valenciennes, 1835), Hick's Reef, Great Barrier Reef.
NEW LAND SNAILS FROM BOGGOMOSS ENVIRONMENTS IN THE DAWSON
VALLEY, SOUTHEASTERN QUEENSLAND (EUPULMONATA: CHAROPIDAE AND
CAMAENIDAE)
JOHN STANISIC
Stanisic, J. 1996 07 20: New land snails from boggomoss environments in the Dawson
Valley, southeastern Queensland (Eupulmonata: Charopidae and Camaenidae). Memoirs of
the Queensland Museum 39(2):343-354. Brisbane. ISSN 0079-8835.
Land snails, Elsothera hewittorum sp. nov. (Charopidae) and Adclarkia dawsonensis gen. et
sp. nov. (Camenidae) are described from on and near mound spring environments in the
Dawson Valley, southeastern Queensland. These mound springs, locally known as bog-
gomosses, are significant perennially moist habitats, in a predominantly dry environment.
Relationships and biogeographic significance of the species are assessed. The roles of the
Dawson Valley and the scattered mesic habitats occurring there are examined in regard to
past episodes of land snail migration. It is concluded that the boggomosses are central to
dispersal of wet-adapted biota in the region. O Mollusca, Eupulmonata, Charopidae,
Camaenidae, biogeography, boggomosses.
John Stanisic, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,
Australia; 25 August 1995.
The Dawson Valley, southeastern Queensland
(SEQ) is located in relatively snail-poor
countryside.The microphyll vine forests and
semi-evergreen vine thickets of the coastal and
sub-coastal ranges to the east support some of the
richest land snail communities of eastern
Australia with up to 40 species recorded at several
sites (Stanisic,1994). However, in southern
Queensland land snail diversity decreases rapidly
with increased distance from the coast and sites
within the Dawson Valley support «10 species.
Climate of the Dawson Valley is sub-humid,
being transitional between the eastern humid and
western semi-arid regions (Nix,1977). Rainfall is
600-700mm annually and mesic refugia which
are liable to support land snails are scarce. Much
of the Dawson Valley has been cleared for cul-
tivation and grazing. In the remaining natural
areas, dry open forest and woodland com-
munities, dominated by Acacia harpophylla
(brigalow), are conspicuous (Johnson, 1984) (Fig.
1C,D). Highly developed rainforest does not
occur, but semi-evergreen vine thicket charac-
terised by Brachychiton rupestre, is sparsely scat-
tered in the region (Speck, 1968). This vegetation
community is usually found on basalt-derived
brown and grey-brown loams and clays. Besides
well-drained soils, vine thickets also favour the
limited rocky outcrops in the area (Fig. 1A,B).
These outcrops act as moisture reservoirs and
provide protection from fires. Other moist
microhabitats are found along drainage lines (Fig.
1F) where weathered alluvium and outcropping
Tertiary rocks furnish suitable conditions for the
maintenance of small discontinuous pockets of
‘rainforest-derived’ flora (pers. obs.). In this
context new land snails on and around the bog-
gomoss on Mount Rose Station (Fig. 1E), NE of
Taroom, Dawson Valley raise interesting ques-
tions on significance of scattered mesic refugia to
land snail survival in marginally dry areas.
BOGGOMOSSES
The boggomosses are a series of small, elevated
peat bogs or swamps scattered among the wood-
land communities near Taroom, on the Dawson
River. They are fed by mound springs which are
leakages from aquifers of the Great Artesian
Basin (Wilson, 1995). Water forces its way to the
surface through faults in the underlying
sandstone. Where this water percolates above the
ground a unique mesic habitat is formed. Sedges,
tall grasses and ferns form a green ground-carpet
of vegetation; sometimes an understorey of
smaller trees is developed. Large gums are a
feature. In many cases the dominant large tree
species is Eucalyptus coolabah (coolibah) whose
size and shallow root system combined with the
soft, moist substrate in which it lives, makes it
vulnerable to wind damage. Thus a feature of the
boggomosses is a large amount of ground-strewn,
branch and whole tree debris which provides a
diverse array of litter-zone microhabitats. AI-
though a number of the boggomosses have been
degraded to varying degrees by stock, and in
some instances fire, they are still significant
habitats for moisture loving biota.
MEMOIRS OF THE QUEENSLAND MUSEUM
LAND SNAILS OF DAWSON VALLEY BOGGOMOSS
SYSTEMATICS
Class GASTROPODA
Order EUPULMONATA
Suborder STYLOMMATOPHORA
Family CHAROPIDAE
Elsothera Iredale, 1933
Elsothera Iredale, 1933:53; Iredale, 1937a:324 Iredale,
1937b: 24; Stanisic,1990:160; Smith, 1992:187.
TYPE SPECIES. Helix sericatula Pfeiffer,1850; by
original designation.
REMARKS. Elsothera is characterised by
species which have strongly, radially ribbed ,
brown to greyish shells (often with darker
periostracal streaks) , with or without an um-
bilicus and with a protoconch that has pre-
dominantly radial ribs and low, moderately
spaced spiral cords. The complex vas deferens/
epiphallus junction, expanded vas deferens and
penis with very enlarged apical bulb and reduced
sheath is a combination of characters peculiar to
Elsothera sensu Stanisic,1990. Elsothera pre-
viously extended from SE S AUST into N NSW.
The new species significantly extends the range
of the genus. As presently understood, it is
probably the most ecologically diverse genus of
the Charopidae, inhabiting rainforest, wet and dry
sclerophyll forest, and woodland.
Elsothera hewittorum sp. nov.
(Figs 2A-C; 3A-F; 4A-D)
ETYMOLOGY. For the Hewitt family, owners of
Mount Rose Station, Taroom.
MATERIAL EXAMINED. HOLOTYPE
QMMO56283, Taroom, c.45km NE on Mt Rose Stn,
SEQ (25°27°15"S,150°01'15"E), under log beside
boggomoss, woodland. Collected J.Stanisic, 23 May
1995. Height of shell 2.52mm, diameter 5.21mm, H\D
ratio 0.51, D\U ratio 3.49, whorls 4 3M. PARATYPES
QMMO56281, 6 specimens, same data as holotype.
DESCRIPTION. Shell (Fig. 2) small, diameter
4.87-5.29mm (mean 5.06mm) with 4 1/2-5 (mean
4 3/4) evenly coiled whorls. Apex and spire
slightly elevated, height of shell 2.27-2.94mm
(mean 2.57mm), SP\BWW ratio 0.10-0.20 (mean
0.13),H/D ratio 0.47-0.61 (mean 0.51).
345
Fig. 2. A-C, holotype of Elsothera hewittorum sp. nov.,
QMMO56283, Mt Rose Stn. Scale line=3mm.
FIG. 1. Vegetation communities in the Taroom area, Dawson Valley, SEQ. A,B, vine thicket on rocky outcrop,
Mt Rose Stn; C, open woodland, Boggomoss Stn; D, brigalow\eucalypt association, Boggomoss Stn; E,
boggomoss, Mt Rose Stn, the type locality of Adclarkia dawsonensis sp. nov.; F, remnant vine thicket along
edges of Cabbage Tree Creek, near Taroom.
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 3. Shell and radular details of Elsothera hewittorum sp. nov. , QMMOS6283, holotype, Mt Rose Stn. A,
apical sculpture. B, body whorl sculpture. C, close-up of a major rib to show extent of microsculpture. D,E,
central and inner lateral teeth. F, marginal teeth. Scale lines as marked.
Protoconch somewhat dull, of 1 1/4 whorls, ap-
proximate width 620(m. Apical sculpture (Fig.
3A) of evenly and moderately spaced, fine wavy,
spiral cords and weak, slightly curved low radial
ridges, mainly on the second half of the
protoconch, becoming more pronounced toward
the protoconch-teleoconch boundary. Post-
nuclear sculpture (Fig. 3B) of crowded, pro-
minent, weakly protractively sinuated radial ribs,
each with a well developed periostracal blade.
Ribs on body whorl 144-176 (mean 157). Micros-
culpture (Fig. 3B) of fine radial riblets, 4-8 be-
tween each pair of major ribs, and fine, crowded
spiral cords; microsculpture continuous on major
ribs (Fig. 3C). Umbilicus wide, deep cup-shaped,
diameter 1.29-1.64 (mean 1.46mm), D/U ratio
LAND SNAILS OF DAWSON VALLEY BOGGOMOSS 347
a
cert
or uu
Re RRR se as
d pee UA DUE
Tir
FIG. 4. Reproductive anatomy of Elsothera hewittorum sp. nov. , QMMOS6283, holotype, Mt Rose Stn. A,
reproductive system.B, vas deferens-epiphallus junction. C, penis interior. D, ovotestis. Abbreviations outlined
in text. Scale lines as shown.
3.19-3.78 (mean 3.49). Sutures impressed,be-
coming more so as the last whorl slowly des-
cends. Aperture subcircular, lip simple. Colour of
shell greyish-brown with occasional darker
periostracal streaks. Based on 5 measured
specimens (QMMOS56281, QMMO56283).
Penis (P) (Fig. 4A) relatively long, with a
prominent apical bulb. Penial sheath (PS) only in
the bulb region. Penis internally (Fig.4C) with a
series of low spongy thickenings distally (PPS)
and a prominent fleshy, irregularly shaped
pilaster proximally (PPM); basal region of penial
chamber separated from apical region by a low
circular collar-like thickening. Epiphallus (E)
long,muscular and twisted about the penis, inter-
nally with longitudinal thickenings, entering
penis apically through a simple pore (EP). Penial
retractor muscle (PRM) attached to penial bulb
adjacent to the penis-epiphallus junction.Vas
deferens (VD) initially an expanded tube with
prominent glandular walls, descending to the
peni-oviducal angle, then ascending and twisting
about the penis prior to becoming a thin tube and
entering the epiphallus through a fleshy, circular
collar (Fig.3B). Vagina (V) c.1/2 the length of the
penis. Free oviduct (UV) large, swollen. Prostate-
uterus (DG\UT) without unusual features, Atrium
(Y) short. Spermatheca (S) with an enlarged base,
long and thin stalk (SS) and large, ovate head
appressed to the surface of the albumen gland
(GG). Hermaphroditic duct (GD) typical.
Ovotestis (G) (Fig. 3D) two clumps of finger-like
alveoli oriented parallel to the plane of coiling
and embedded in the apical whorls of the diges-
tive gland (Z). Animal colour grey. Radula (Fig.
3D-F) with tricuspid central and lateral teeth
(central slightly smaller) in which the mesocone
is long and lanceolate, ecto- and endocones short
and acutely pointed; marginal teeth multicuspid
due to ectoconal splitting. Based on one dissected
specimen (OMMO56281).
COMPARATIVE REMARKS. Elsothera hewit-
torum sp.nov. resembles E. funerea (Cox,1868)
from NSW (type locality: Mudgee) and E. nesana
Iredale,1937 (type locality:Port Lincoln, South
Australia). [The latter species was considered a
synonym of E. murrayana (Pfeiffer, 1864) by
Smith (1992) but has a tighter coiling pattern and
much finer ribbing]. Compared with £. funerea,
E. hewittorum is smaller with less elevated spire
and apex, has smaller whorls, finer adult ribbing,
larger umbilicus and almost obsolete radial ribs
on the the protoconch (Fig. 3A). E. nesana has
similarly disposed radial ribs but has larger
whorls,more prominent radial ribs on the
protoconch and a basally flattened body whorl, £.
nautilodea (Cox,1866) and E. genithecata
Stanisic,1990 from NE NSW are readily distin-
guished from the new species by their closed
umbilici. E. sericatula (Pfeiffer,1850) from
central coastal NSW has a closed umbilicus and
radial ribs on the entire protoconch. E. murrayana
has more loosely coiled whorls and widely
MEMOIRS OF THE QUEENSLAND MUSEUM
spaced bold radial ribs. The combination of very
small, brown shell with dense radial ribbing,
slightly raised spire and relatively wide umbilicus
distinguishes E. hewittorum from other land
snails of the Dawson Valley.
RANGE AND HABITAT. Known only from the
type locality. More westerly species of this com-
plex (E.funerea, E.nesana, E.murrayana) inhabit
drier woodland environments with a preference
for mesic refuges such as rocky outcrops, par-
ticularly limestone (Stanisic unpubl. data).
E.hewittorum has a similar ecologic preference.
REMARKS. Generic assignment of E. hewit-
torum is based on structural similarity of its
reproductive system to that of Elsothera by
Stanisic (1990). However, the anatomical consis-
tency of species of Elsothera (Iredale, 1937a,b;
Stanisic,1990; Smith, 1992) underpins a wide
variation in shell form. Shell shape has a strong
ecological correlation (Solem & Climo, 1985;
Stanisic, 1990) and the variability in this character
in Elsothera reflects the diverse ecological
preferences of its members. E. hewittorum is
closest to E. funerea and E. nesana with which it
shares a greyish-brown shell with very open um-
bilicus and apical sculpture of radial ribs and
prominent spiral cords. The three species also live
in similar habitats.
Family CAMAENIDAE
Subfamily CAMAENINAE
Adclarkia gen. nov.
ETYMOLOGY. For Adam Clark of Taroom,
TYPE SPECIES. Adclarkia dawsonensis sp. nov.
DIAGNOSIS. Shell large, relatively thin, with
weakly elevated spire and apex, comparatively
small umbilicus and weakly reflected lip. Apical
sculpture of scattered subcircular pustules. Adult
sculpture of very weak radial growth ridges and
moderately spaced elongate pustules ; micros-
culpture of fine periostracal ridgelets. Genitalia
elongate,with a long tubular penis, short epiphal-
lic flagellum and proto-sheath enveloping penis,
epiphallus and epiphallic flagellum. Penial
retractor muscle inserted on epiphallus. Penis
internally with prominent pustules and a wrinkled
tubular verge with a terminal pore. Head wart
present,
COMPARATIVE REMARKS. The shell of
LAND SNAILS OF DAWSON VALLEY BOGGOMOSS
Adclarkia bears little resemblance to other large
Queensland camaenids. The relatively thin, al-
most monochrome shell is in direct contrast to the
larger, striped and solid-shelled camaenids of
coastal regions. Taxonomic significance of these
external differences is supported by the reproduc-
tive anatomy, which in Adclarkia has numerous
irregularities not seen in moist-forest camaenids
further east (Solem, 1992a). In particular the
proto-sheath and pattern of penial pustules con-
trasts with the muscular penial sheath and very
complicated penial sculpture in Sphaerospira
Morch, 1867, Bentosites Iredale, 1933 and Hadra
Albers, 1860. Meridolum Iredale, 1942 and Ther-
sites Pfeiffer, 1842 (see Solem, 1992a) from coas-
tal regions of southern New South Wales and
southern Queensland have a protosheath rather
than a prominent penial sheath and pustulose
penial wall sculpture. However, the shells of
these two genera have different form and micros-
culpture. Only M.gilberti (Pfeiffer, 1846) has
shell surface pustules but these are round and
crowded and lack the periostracal scales of A.
dawsonensis. Galadistes Iredale, 1938 from in-
land northern New South Wales and Pallidelix
Iredale, 1933 from the Expedition Range in
southern Queensland have yet to be examined
anatomically. These genera have pustules on the
adult shell but differ in form (circular) from those
of Adclarkia. They also have quite different shell
shape (globose) and apical sculpture (very
crowded pustules).
Cupedora lredale,1933 from eastern South
Australia most resembles Adclarkia in anatomy
and shell. Both genera have a proto-sheath and
similar pustulation in the penis. However, in
Cupedora the pustulations in the penial chamber
are largely confined to the upper half with
prominent longitudinal ridges in the lower half.
In particular the shell sculpture of Cupedora (see
Solem, 1992a, pl. 26, figs c,d), apart from the
shape of the pustules, is very similar to that of
Adclarkia. Main differences are the the greater
calcification in shells of Cupedora and their ten-
dency to have reddish chestnut bands in most
cases.
Adclarkia dawsonensis sp. nov.
(Figs 5, 6A-C, 7A-H, 8A-D)
ETYMOLOGY. For the Dawson Valley.
MATERIAL EXAMINED. HOLOTYPE QM
MO56284, Taroom, c.45km NE, on Mt Rose Stn, SEQ
(25?27' 15"S, 150701" 15"E), in litter beneath sandpaper
fig on boggomoss. Collected by J. Stanisic, J. Johnson,
J. Short, 23 May 1995. Height of shell 14.58mm,
349
FIG. 5. Adclarkia dawsonensis sp. nov., QM
MO56284, holotype, Mt Rose Stn.
diameter 21.68mm, HD ratio 0.67, D\U ratio 9.86,
whorls 5 1/8. PARATYPES QMMO56280, 4 sub-
adults, 12 juveniles, same data as holotype;
QMM04239, | adult, Theodore, Dawson Valley, SEQ,
under rubbish in damp situation; QMMO6779, 2
adults, Theodore, Dawson Valley, SEQ, in garden
crawling after rain, ex S. McKay; QMMO56289,1
adult, 7 juveniles, same locality as holotype, under
logs, C. Eddie, 11 July 1995,
DESCRIPTION. Shell (Fig. 6A-C) comparative-
ly thin, diameter 21.68—25.74mm (mean
23.82mm) with 5 1/8—5 5/8 (mean 5 1/2 whorls.
Apex and spire very slightly elevated, height of
shell 14.58-16.62mm (mean 15.80mm), H\D
ratio 0.65—0.67 (mean 0.67). Apical sculpture
(Fig. 7A) of radially disposed, moderately
spaced, irregularly elongate pustules. Spire and
body whorl with densely scattered, elongate pus-
tules (Fig.7B) and weak, radial growth ridges; in
unworn specimens a small periostracal scale sits
atop each pustule (Fig. 7C) . Microsculpture of
fine, crowded ridgelets (Fig. 7B,C). Shell
periphery rounded with last whorl slowly des-
cending. Aperture subcircular. Lip weakly
reflected, columellar margin dilated and partially
covering the umbilicus. Umbilicus small, width
2.34—3.24mm (mean 2.63mm), D/U 7.15—10.55
(mean 9.26). Shell colour light-brown to
greenish-yellow horn, occasionally with a nar-
row, red subsutural band and a small, red circum-
umbilical patch; lip white. Based on four
measured specimens (QMMOS6284,
QMM04239, QMMO6779).
Genitalia (Fig. 8A) with long vagina (V) and
short free oviduct (UV). Vagina internally with
high, thin, longitudinal pilasters. Spermatheca
(S), with stalk swollen basally and head ap-
350
FIG. 6. A-C, Shell of Adclarkia dawsonensis sp. nov.
QMMO56284, holotype, Mt Rose Stn. Scale in mil-
limetres.
MEMOIRS OF THE QUEENSLAND MUSEUM
pressed to base of albumen gland (GG). Prostate-
uterus (DG\UT) without unusual features. Ter-
minal male genitalia enveloped in a sheath of
connective tissue (PS). Penis (P) slightly longer
than vagina, muscular; internally (Fig. 8B) with
longitudinal rows of elongate pustules (PP)
which become more crowded in the lower half of
the penis before giving rise to fleshy longitudinal
pilasters (PPL) in the lower quarter; apically with
a wrinkled, tubular papilla (PV). Epiphallic pore
(EP) terminal on the papilla. Epiphallus (E) a
reflexed, muscular tube, about half the length of
the penis, becoming enlarged about halfway
along its length, internally with longitudinal
thickenings; a short finger-like flagellum (EF) is
located at the epiphallus-vas deferens junction.
Vas deferens (VD) a thin tube descending from
the prostate-uterus to the peni-oviducal angle,
reflexing apicad of the angle. Penial retractor
muscle (PRM) inserted on the epiphallus at the
point where it reflexes. Talon (GT) short, finger-
like, embedded in the surface of the albumen
gland. Hermaphroditic duct (GD) weakly con-
voluted, not swollen. Albumen gland (GG) elon-
gate, rusty-brown. Ovotestis (G) (Fig. 8C) several
clumps of long, creamy-coloured, finger-like al-
veoli, in the apical whorls of the digestive gland
(Z). Atrium (Y) without unusual features. Animal
colour light brown to white with varying amounts
of grey in the neck region, on the sides of the foot
and above the tail. Surface of lung-roof and vis-
ceral coil with prominent, irregular black pig-
mentation (Fig. 5). Mantle mustard-orange. Head
wart (Fig. 8D) located between the bases of the
superior tentacles. Radula (Fig. 7F-H) with basi-
cally unicuspid central and lateral teeth, central
slightly smaller, with very tiny ectocones situated
half-way up the main cusp shaft. Lateral teeth
with prominent anterior flare, reduced in central
tooth. Lateromarginal teeth tricuspid with en-
docone located high up on the mesocone. Mar-
ginal teeth multicuspid with ectocone splitting
into several minor cusps. Jaw (Fig. 7D,E) with
several prominent central ribs, considerably
reduced on the side and microscopically,
transversely striated. Based on one dissected
adult (QMMO56284).
FIG. 7. Shell, jaw and radular details of Adclarkia
dawsonensis sp. nov., QMMOS56284, holotype, Mt
Rose Stn. A, apical sculpture. B, pustules and fine
ridgelets on body whorl. C, body whorl pustules with
periostracal scales. D, jaw. E, microsculpture on ribs
of jaw. F,G, central and inner lateral teeth. H, mar-
ginal teeth. Scale lines as marked.
LAND SNAILS OF DAWSON VALLEY BOGGOMOSS 351
352 MEMOIRS OF THE QUEENSLAND MUSEUM
«^
xt
LO e.. Ç, à
do NOME A
6S 9 1
EC
2 EE- m.
ie
AHAC
gj I aca. S
Giese
Ses
N2)
YS
AD
WN
SY
AS
FIG. 8. Reproductive anatomy of Adclarkia dawsonensis sp. nov., QMMOS6284, holotype, Mt Rose Stn. A,
reproductive system. B, penis interior. C, ovotestis. D, head wart. Abbreviations outlined in text. Scale lines as
marked.
LAND SNAILS OF DAWSON VALLEY BOGGOMOSS
COMPARATIVE REMARKS. A number of
other large camaenids occur in the Dawson Val-
ley. Sphaerospira mattea (Iredale, 1933) has a
similarly shaped shell but is yellowish-white with
several reddish-brown chestnut bands. No sur-
face pustulation is present. The animal is reddish-
grey with a red mantle. Xanthomelon
pachystylum (Pfeiffer, 1845) is much larger than
A.dawsonensis and has a brownish-yellow, heavy
and globose shell without an umbilicus. The
animal is light grey in colour and lacks the
prominent dark pigmentation of A. dawsonensis.
However, most confusion is liable to arise
through the occurrence of a local, as yet unnamed
rhytidid which has a shell of similar size and
shape to that of A.dawsonensis. This snail is
readily recognised by its yellowish, more flat-
tened shell with rather large umbilicus and sculp-
ture of prominent radial ribs.
RANGE AND HABITAT. Taroom to Theodore,
in the Dawson Valley. It is a free sealer living in
litter and under logs.
REMARKS. Two lots of shells from Theodore
(QMMO4239, QMMO6779) have been in the
Queensland Museum since the mid-1970's but
attempts to find live specimens in the Theodore
area have proved fruitless. The thin shell of
A.dawsonensis indicates that it is not an inhab-
itant of rock talus but prefers living amongst
vegetation and soil (Stanisic pers. obs.). This
makes it especially prone to extermination
through habitat destruction by fire and land clear-
ing. Considering the degree of habitat modifica-
tion which has taken place in the Dawson Valley
it is possible that the distribution of A. dawsonen-
sis is now extremely restricted.
DISCUSSION
The Dawson Valley is depauperate in land
snails compared with areas of closed forest fur-
ther east, yet it would appear that it has played an
important part in the past dispersal of terrestrial
molluscs now more diverse elsewhere. On the
basis of similarities in shell form and reproduc-
tive anatomy E. hewittorum and A. dawsonensis
appear related to taxa further south. If this view
is supported by much-needed revisionary studies
of related taxa in the drier areas of western New
South Wales the occurrence of these two species
in the Dawson Valley region has interesting
biogeographic implications.
Solem (1992b) contended that colonisation of
eastern South Australia by the Camaeninae, in-
cluding Cupedora, was from the north through
what is now coastal and subcoastal Queensland,
and then through New South Wales. Their ances-
tors came from Papua New Guinea as post-
Miocene immigrants (Bishop,1981). Finding a
possible relative of this South Australian group
of camaenids in the Dawson Valley region lends
support to Solem's contention. The thin shell of
A.dawsonensis would not be unexpected in an
ancestor which lived in scattered moist refugia on
alluvial flats and along drainage lines. The more
complicated penial surface sculpture of
Cupedora is probably due to sympatric species
interactions but nevertheless can be readily
derived from the simple pattern of pustules and
longitudinal ridges seen in A. dawsonensis.
Cupedora has adapted to living in a restricted
moisture regime and is most frequently as-
sociated with rocky slopes, crevices and talus.
The heavier more robust shell of its members is
typical of species with this habitat preference.
E. hewittorum provides additional support for
the claim that the Dawson Valley region was an
important corridor for snail movement in the past.
Apart from the rainforest inhabiting E.
genithecata there is no other record of the genus
in Queensland. Intensive collecting in the
Chinchilla area has failed to locate it in that
region. This identifies E. hewittorum as a north-
ern outlier of a group which has a nearly con-
tinuous distribution from northern New South
Wales through western Victoria and into eastern
South Australia. The Charopidae is a Gondwanan
family mainly confined to rainforests (Stanisic,
1990). Elsothera is the only group which has
managed to diversify into the drier eucalypt
forests. Presumably the dispersal was from wetter
forests in the east or from ancestors isolated in
mesic refugia as rainforests retreated to the coas-
tal mountain ranges. In either case this radiation
most probably occurred sometime in the Pliocene
when conditions were wetter (Kemp,1981) and
were more likely to facilitate the dispersal of
these tiny animals. The extensive distribution of
the genus suggests an early radiation.The aggres-
sive drying of the Pleistocene (Galloway &
Kemp,1981) would have fragmented widespread
populations leading to extinctions where mesic
refugia did not provide ameliorating micro-
climates. The close resemblance of E. hewittorum
to E. funerea from northern NSW suggests
derivation from a common ancestor and its
similarity to E. nesana from South Australia at-
tests to the extent of the initial radiation. It is
354
probably not coincidental that the evolutionary
track spanned by these charopids mirrors that of
A. dawsonensis and Cupedora.
Mesic refugia such as those associated with
drainage lines in the Dawson Valley have probab-
ly played an important part in the endurance of
land snails in the region. Their future importance
to the survival of wet-adapted biota such as land
snails should not be underestimated. The unique,
perennially moist boggomosses of Taroom are an
integral part of this intricate survival story. It is
unlikely that A. dawsonensis and E. hewittorum
are restricted to the boggomoss environment on
Mount Rose Stn. A.dawsonensis appears to
favour drainage lines (Theodore is also on the
Dawson River) and is probably spread over the
alluvial flats of the Dawson Valley by floods.
Following extensive land clearing in the Dawson
Valley, opportunistic colonisation of the few
remaining mesic refugia such as the boggomos-
ses is now probably an essential part of the long
term viability of this species. The single occur-
rence of E. hewittorum makes any speculation
about survival in preferred habitat less informa-
tive, however, the same general principles of
dispersal and colonisation would apply.
The boggomosses, particularly those with
structured vegetation communities, form an im-
portant part of the scattered archipelago of mesic
refugia in the Dawson Valley. Considering the
damage which has been caused to these habitats
in the past, by both stock and fire, it will be
necessary to implement more sympathetic
management practices in the future to ensure their
continued existence and the survival of the biota
which they support.
ACKNOWLEDGEMENTS
Ithank Adam Clark and Ian Hewitt of Taroom
for advice and hospitality. 1 thank Kylie Stumkat
for the SEMs and the Queensland Museum
Photography Section.
LITERATURE CITED
BISHOP, M. J. 1981. The biogeography and evolution
of Australian land snails. Pp. 924-954. In Keast,
A. (ed.), Ecological biogeography of Australia.
(Dr W. Junk: The Hague).
GALLOWAY, R. W. & KEMP, E. M. 1981. Late
Cainozoic environments in Australia. Pp. 52-80.
MEMOIRS OF THE QUEENSLAND MUSEUM
In Keast, A. (ed.), Ecological biogeography of
Australia. (Dr W. Junk: The Hague).
IREDALE, T. 1937a. A basic list of the land Mollusca
of Australia. Australian Zoologist 8: 287-333.
1937b. An annotated checklist of the land shells of
south and central Australia. The South Australian
Naturalist 18:6-56.
JOHNSON, R. W. 1984. Flora and vegetation of the
Brigalow Belt. Pp. 41-59. In Bailey, A. (ed.), The
Brigalow Belt of Australia. (Queensland Depart-
ment of Primary Industries: Brisbane).
KEMP, E. M. 1981. Tertiary palaeogeography and the
evolution of the Australian climate. Pp. 32-49. In
Keast, A. (ed.), ‘Ecological biogeography of
Australia’. (Dr W. Junk: The Hague).
NIX, H. A. 1977. Land classification criteria: climate.
Pp. 27- 40. In Land units of the Fitzroy region,
Queensland. Land Research Series No. 39. (Com-
monwealth Scientific and Industrial Research Or-
ganisation, Australia: Melbourne).
SMITH, B.J. 1992. Non-marine Mollusca. Pp.1-405. In
Houston, W.W.K. (ed.), 'Zoological catalogue of
Australia'. (Australian Government Printing Ser-
vice: Canberra).
SOLEM, A. 19922. Camaenid land snails from southern
and eastern South Australia, excluding Kangaroo
Island. Part 1. Systernatics, distribution and varia-
tion. Records of the South Australian Museum,
Monograph Series 2:1-338.
1992b. Camaenid land snails from southern and
eastern South Australia, excluding Kangaroo Is-
land. Part 2. Biogeography and covariation.
Records of the South Australian Museum,
Monograph Series 2: 339-425.
SOLEM, A. & CLIMO, F.M. 1985. Structure and
habitat correlations of sympatric New Zealand
land snail species. Malacologia 26: 1- 30.
SPECK, N.H. 1968. Vegetation of the Dawson-Fitzroy
area. Pp. 157-173. In Perry, R.A. (ed.), Lands of
the Dawson-Fitzroy area, Queensland.Land Re-
search Series No. 21. (Commonwealth Scientific
and Industrial Organisation, Australia: Mel-
bourne).
STANISIC, J. 1990. Systematics and biogeography of
eastern Australian Charopidae (Mollusca, Pul-
monata) from subtropical rainforests. Memoirs of
the Queensland Museum 30:1-241.
1994. The distribution and patterns of species diver-
sity of land snails in eastern Australia. Memoirs
of the Queensland Museum 36: 207-214.
WILSON, B. 1995. Artesian springs of the Great Ar-
tesian Basin, Pp. 1-22. Unpublished report for the
Queensland Department of Environment and
Heritage and the Australian Nature Conservation
Agency.
A NEW CAMAENID LAND SNAIL FROM THE WET TROPICS BIOGEOGRAPHIC
REGION, NORTHEASTERN QUEENSLAND (EUPULMONATA:CAMAENIDAE)
JOHN STANISIC
Stanisic, J. 1996 07 20: Anew camaenid land snail from the Wet Tropics Biogeographic
Region, northeastern Queensland (Eupulmonata:Camaenidae). Memoirs of the Queensland
Museum 39(2): 355-363. Brisbane. ISSN 0079-8835.
A new camaenid land snail, Monteithosites helicostracum gen. et sp. nov., is described from
the rainforested summits of Bakers Blue Mountain and Hanns Tableland, NE Queensland.
These localities are western outliers of the Wet Tropics rainforest massif. The species
displays periostracal sculpture and genital anatomy which suggest that its relationships are
with camaenids much further to the south. Biogeographic implications of this discovery are
discussed. [ ] Mollusca, Camaenidae, Wet Tropics, biogeography.
John Stanisic, Queensland Museum, P.O, Box 3300, South Brisbane, Queensland 4101,
Australia; received 20 May 1996.
A number of large-sized camaenid land snails
are known from the Wet Tropics region of north-
eastern Queensland (NEQ). Smith (1992) placed
these in Hadra Albers, 1860, Sphaerospira
Morch, 1867, Spurlingia Iredale, 1933 and Jack-
sonena Iredale, 1937. Monteithosites helico-
stracum gen. et sp. nov. from the summits of
Bakers Blue Mountain (Mtn) and the Hanns
Tableland, southwest of Mount Molloy, NEQ is
a biogeographically significant addition to this
fauna.
Bakers Blue Mtn and the Hanns Tableland
(Fig.1) are elevated western outliers of the main
Wet Tropics rainforest massif. They are separated
from the more expansive Carbine and Windsor
Tablelands in the north by the valley of the
Mitchell River but are interconnected by a series
of low ridges and hills which continue through to
the Herberton Range in the south. The lower
slopes generally support eucalypt woodland.
However the peaks in these outliers exceed
1000m and their summits, where rainfall is ade-
quate, support rainforest. The dominant rain-
forest type 1s araucarian notophyll vine forest
with austral conifers (Agathis robusta, Araucaria
cunninghamii) (Tracey, 1982). Floristic relation-
ships of this rainforest appear to be with Mt Lewis
in the north (Godwin, 1986). Rainforest on Hanns
Tableland is confined to the northern end and to
scattered valleys and fireproof niches. The peaks
of Bakers Blue Mtn are cooler and wetter than
those of the Hanns Tableland and consequently
support larger areas of rainforest. Nix (1991)
identified these outliers as distinct biogeographic
units within the Wet Tropics region.
M.helicostracum gen. et sp. nov. provides im-
portant new information on these isolated refugia
and their historical relationship to other parts of
the Wet Tropics.
SYSTEMATICS
Class GASTROPODA
Order EUPULMONATA
Suborder STYLOMMATOPHORA
Superfamily CAMAENOIDEA
Family CAMAENIDAE
Previous taxonomic studies of the large
Camaenidae of the Wet Tropics, NEQ have been
largely shell-based. Pilsbry (1894) grouped all
these species under Thersites Pfeiffer, 1855 in a
revision that was partially based on genital
anatomy. Iredale (1933, 1937) abandoned
Pilsbry’s conservative system and introduced a
large number of new species and genera, solely
on the basis of shell features. Some of these taxa
have somewhat doubtful status (Burch, 1976) and
have yet to be critically revised. Pace (1901) and
Solem (1979) presented anatomical data on the
very large Hadra bipartita (Ferussac, 1822).
Smith (1992) produced an updated but critically
untested listing of taxa. Stanisic et al. (1994)
produced a preliminary report on the land snails
of the Wet Tropics which presented distribution
data for a large number of described and un-
described camaenids; the new species described
herein was identified as Camaenidae WT 11.
The description of this new land snail has en-
tailed a precursory anatomical examination of
several other camaenid species in order to place
itin systematic context. These revisionary studies
will be more fully pursued elsewhere but do
356
Lis
^ - -
lmt Windsor ! N
} Tableland \
[4
13 P f
1. Z Mt Carbine
^j Tableland
Cape Tribulation
[
4
ILS
Mt Lewis Nee
Bakers Blue $c is 15 KA
Mtn ` V^ ^5
A “TNR
Hanns $e. \ * ipe
Tableland }
reeba
17*00' Me
<
)$ Atherton A
€ Tableland x
Innisfail
=
18°00'
~
Sef
19°00" Kilometres
145°00'
FIG. 1. Distribution of Monteithosites helicostracum
sp.nov. - € and the biogeographic units of the Wet
Tropics (after Nix, 1991). [1, Finnigan; 2, Thornton;
3, Windsor; 4, Carbine; 5. Macalister; 6, Bakers Blue;
7, Hanns; 8, Malbon Thompson; 9, Atherton; 10,
Hinchinbrook; 11, Seaview; 12, Paluma].
identify M. helicostracum as an unusual member
of the Camaenidae of the Wet Tropics.
Monteithosites gen. nov.
ETYMOLOGY. For Dr Geoff Monteith and in refer-
ence to the similarity to Bentosites from mideastern
Queensland (MEQ).
MEMOIRS OF THE QUEENSLAND MUSEUM
TYPE SPECIES. Monteithosites helicostracum sp.
nov.
DIAGNOSIS. Shell large (to 33.88mm in
diameter), yellow with few to many, dark red-
dish-brown spiral bands; lip dark brown and
reflected. Shell sculpture of very fine, crowded,
radial thread-like periostracal wrinkles and
prominent, widely spaced spiral periostracal
cords. Penis short, stout, with a thick sheath and
highly muscularised walls; verge absent.
Epiphallus with a weakly expanded ascending
arm; entering penis apically through a simple
pore. Vas deferens and epiphallus bound to penial
sheath. Animal with reddish mantle and tail and
dark brown tentacles.
COMPARATIVE REMARKS. The unusual
periostracal sculpture readily separates Mon-
teithosites from other large camaenids of the Wet
Tropics. Hadra (sensu Smith, 1992) has
bicoloured shells with reduced banding and either
almost smooth periostracal sculpture or, in the
case of H. bellendenkerensis (Brazier, 1875) and
H. beddomae (Brazier, 1878), fine radial threads
with prominent radially disposed, zig-zag
periostracal thickenings; Spurlingia and Jack-
sonena have comparatively drab shells with
strongly rugose sculptures and prominent
periostracal scales; Sphaerospira Morch, 1867
(sensu Smith, 1992) contains a large number of
species with banded shells occurring from SEQ
to NEQ and previously included in 5 genera
(Iredale, 1937). Unpublished studies by the
author suggest that this genus, as defined by
Smith (1992), is polyphyletic. A primary division
is expressed in animal colour. The greater propor-
tion of species have animals with orange to red-
dish-orange mantle tissue and tails, and orange to
brown tentacles e.g. S. yulei (Forbes, 1851) from
MEQ. A smaller number have all-over grey to
black animals e.g. S. fraseri (Griffith & Pidgeon,
1833) from SEQ. The animal of Monteitosites
bears a strong, external resemblance to the former
group particularly those species from MEQ
which were previously included in Bentosites
Iredale, 1933 and Varohadra Iredale, 1933 (see
Iredale, 1937).
Periostracal sculpture in Sphaerospira s.1.
varies from a simple microscopic pattern of very
fine, crowded, radial periostracal, thread-like
wrinkles (S. yulei), to one in which these fine
wrinkles are supplemented by coarse, wavy or
zigzag, radial periostracal ribs (S. fraseri) not too
dissimilar from those in some Hadra spp. The
NEW CAMAENID LAND SNAIL GENUS FROM THE WET TROPICS
FIG. 2. Monieithosites helicostracum sp. nov. Shell of
holotype, QMMO57242. Scale line = 10mm.
latter pattern is seen in both black-mantled
species (SEQ to NEQ) and red-mantled species
from the southern parts of the Wet Tropics (NEQ)
only. Monteithosites displays yet a third level of
variation. However the underlying microsculp-
ture of radial thread-like wrinkles in Mon-
teithosites is identical to that in the red-mantled
forms from MEQ and NEQ suggesting that the
periostracal cords may be merely a functional
equivalent of the zigzag to wavy types seen in
other hadroid camaenids and of limited value in
establishing relationships.
Genital anatomy relates Monteithosites to red-
mantled Sphaerospira s.l. from MEQ and NEQ
rather than Sphaerospira s.l. from SEQ or Hadra
from the Wet Tropics. The comparatively weakly
muscular epiphallus of Monteithosites contrasts
with the strongly muscular, elongate condition of
Sphaerospira s.l.(Solem, 1992) from SEQ and
Hadra (Solem, 1979). Hadra is also charac-
terised by well-developed epiphallic caeca. S.
U3
Un
-
yulei from MEQ has the epiphallus reduced to
such an extent that the ascending arm is narrow
and barely differentiated from the vas deferens
(Solem, 1992); a penial verge and tiny epiphallic
flagellum are present. This species also has the
simple epiphallic entry and sheath- bound vas
deferens of Monteithosites. Similar genital
anatomy is found in other red-mantled
Sphaerospira from MEQ and the southern parts
of the Wet Tropics. In contrast S. fraseri (and
other black-mantled species) have a strongly
developed epiphallus, no accessory epiphallic
structures and usually lack penial verges (Bishop,
1978; Solem, 1992; Stanisic, unpubl.). The penis
of Monteithosites appears to be highly specialised
and finds no correlatives among hitherto il-
lustrated species.
Monteithosites helicostracum sp. nov.
(Figs 2-5)
ETYMOLOGY, Latin helico, spiral, referring to the
spiral periostracal cords.
MATERIAL EXAMINED. HOLOTYPE
QMM057242, Bakers Blue Mtn, c.17km W Mt Mol-
loy, NEQ (16°42’S, 145°10’E), rainforest, under logs,
Collected G.Monteith, D.Cook, 11-12 September,
1981. Height of shell 25.54mm, diameter 33.88mm,
H/D ratio 0.754, whorls 6 1/8. PARATYPES
QMMO048172, 1 adult, 1 juvenile, same data as
holotype; QMMO57240, 1 adult, 1 juvenile, Bakers
Blue Mtn, upper slopes, NEQ (16°43’21"S,
145°10°15"E), microphyll vine forest/basalt talus,
under logs, J.Stanisic, G.Ingram, 1 Jul 1995;
QMMO56834, 7 adults, Hanns Tableland, N end, NEQ
(16°49’S, 145?1 I E), rainforest, 950- 1000m, G.Mon-
teith, G. Thompson, 11-14 Dec 1995. OTHER
MATERIAL QMMO48181, 3 juveniles, Bakers Blue
Min, c.17km W Mt Molloy, NEQ (16°42’S, 145?10' B),
rainforest, 800- 1000m, ANZSES Expedition, 30 Dec
1989-9 Jan 1990.
DESCRIPTION. Shell (Fig. 2A,B) large,
diameter 29.26-33.88mm (mean 31.66mm), with
5 3/4-6 1/8 (mean 5 7/8-) whorls. Apex and spire
moderately elevated, height 20.56-25.54mm
(mean 22.71mm). H/D ratio 0.699-0.762 (mean
0.727). Protoconch c.1 1/2 whorls. Apical sculp-
ture of crowded, slightly curved, weak radial
growth ridges. Spire and body whorl (Fig. 3) with
very fine, crowded, radial thread-like periostracal
wrinkles and conspicuous, more widely spaced,
spiral, coarse, periostracal cords. Umbilicus bare-
ly open, covered by the reflection of the collumel-
lar lip. Sutures weakly impressed. Whorls evenly
rounded. Last whorl descending rapidly in front.
358
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 3. Monteithosites helicostracum sp. nov., QMMO48172. A,B, details of adult shell sculpture; C,D, details
of radula. Scale lines as marked.
Aperture lunately ovate; lip strongly reflected.
Shell colour yellow-horn with dark reddish-
brown spiral bands; subsutural and peripheral
bands moderately wide sometimes split into a
series of narrower bands; basally a series of nar-
row bands sometimes coalesced into a single
broader band; several narrow intermediate brown
bands may also be present. Umbilical chink and
lip dark brown. Parietal callus shiny, dark choco-
late-brown. Area behind lip with a dark brown
suffusion which may extend rearwood for 1/8
whorl. Based on 9 measured adults
(QMMO48172, QMMO56834, QMMO57240,
QMMO57242).
Genitalia (Fig. 4A) with ovotestis (G, Fig. 4B)
comprised of several clumps of finger-like lobes
of alveoli lying in the apical whorls of the diges-
tive gland (Z). Hermaphroditic duct (GD) strong-
ly convoluted; talon (GT) short, finger-like.
Spermatheca (S) with head lying at the base of the
albumen gland (GG); stalk (SS) long, bound to
the prostate-uterine surface. Uterus (UT) white,
without unusual features; prostate (DG) a series
of light brown acini appressed to the uterine
surface. Free oviduct (UV) short. Vagina (V)
short, about 1/2 length of penis, internally with
numerous longitudinal thickenings; spermathe-
cal entrance a simple pore. Epiphallus (E) with
short thick descending arm and narrow, weakly
expanded ascending arm, about 2/5 length of
penis, internally with a large longitudinal pilaster;
entering penis apically through a simple pore
(EP). Ascending arm of epiphallus and vas
deferens bound to penial sheath by connective
tissue. Vas deferens (VD) a thin tube, barely
differentiated from the epiphallus. Penis (P, Fig.
4C) relatively short, stout, strongly muscularised
with a thick sheath (PS). Penial wall (PW) ex-
tremely thick and penial lumen reduced to a very
narrow tube. Internally, penis with a conspicuous,
NEW CAMAENID LAND SNAIL GENUS FROM THE WET TROPICS
359
SN tp
ae
FIG. 4. Monteithosites helicostracum sp. nov., QMMO56834. A, genitalia; B, ovotestis; C, details of penis
interior. Scale lines as marked.
central longitudinal pilaster (PPM) and several,
narrower minor longitudinal pilasters (PP); lower
penial chamber with short, rectangular pustules;
verge absent. Atrium (Y) very short and without
unusual features. Animal (Fig. 5) grey to dark
brown with reddish tail and mantle and dark
brown tentacles. Head wart present. Radula (Fig.
3B, C) with broadly unicuspid central and lateral
teeth, central tooth slightly smaller; anterior flare
weak, basal plate prominent. Marginal teeth
tricuspid with ectocone low down and endocone
high up on mesoconal cutting edge; basal plate
and anterior flare reduced. Jaw typically
camaenid. Based on two dissected specimens
(QMMOS57242, QMMOS56834).
ae
ee EIL Teen dn
nm
FIG. 5. Monteithosites helicostracum sp. nov., Hanns Tableland, QMMO56683.
COMPARISONS. The shell of M. helicostracum
comes closest in appearance to that of some
Sphaerospira (sensu Smith, 1992) from NEQ and
MEQ. It resembles S. saxicola (Iredale, 1937)
from the Bowen area, MEQ which possesses fine
thread-like periostracal wrinkles, but lacks any
major periostracal elements. Some species in the
S. sardalabiata (Cox, 1871) complex (sensu
Smith, 1992) from the southern parts (Paluma,
Seaview and Cardwell Ranges) of the Wet
Tropics bear an overall similarity to the new
species in size and shape. However,
M.helicostracum is distinguished from these
species by its spiral periostracal cords. S. sar-
dalabiata s.l. has coarse, wavy to zigzag, radial
periostracal thickenings (Stanisic, unpubl.).
Sphaerospira spp. from NEQ and MEQ also
show some anatomical similarity to M.
helicostracum in reduction of the epiphallus,
penial structures, form of the radula (Pilsbry,
1894; Solem, 1992; Stanisic, unpubl.) and animal
colour. However the rather unusual penis of M.
helicostracum distinguishes it from all of these
species (Stanisic, unpubl.).
HABITAT AND DISTRIBUTION. Under logs
in araucarian notophyll vine forest with austral
conifer emergents on Bakers Blue Mtn and Hanns
Tableland, NEQ.
REMARKS. The adult periostracal sculpture of
coarse spiral cords and the penial specialisation
of M. helicostracum is a combination of charac-
ters not seen in other hadroid camaenids. The
MEMOIRS OF THE QUEENSLAND MUSEUM
periostracal spiral cords
may be functionally
analagous to the more wide-
ly occurring zigzag to wavy
variety but the unusual
penis is more difficult to
correlate. Sphaerospira
rockhamptonensis (Cox,
1873) from The Caves area,
Rockhampton, MEQ has a
penis with thick muscular
walls and reduced lumen
| (Stanisic, unpubl.). How-
ever this species has a dark-
grey to black animal with a
greatly enlarged epiphallus
and is not closely related. It
has a simple periostracal
sculpture of very fine
wrinkles. An analagous
reduction in penial lumen
width (in this case caused
by a dramatic narrowing of the entire penis) was
noted by Solem in Turgenitubulus pagodula
Solem, 1985, an unrelated camaenid from NW
Australia. In this instance more fundamental
characters indicated the species' relationship to a
widespread genus with otherwise strongly con-
sistent genital anatomy. Similarly the general pat-
terns of shell, animal, and anatomical characters
displayed by M. helicostracum suggest a relation-
ship with those species with banded shells and
orange to red coloured animals from NEQ and
MEQ. The simple internal structure of the penial
chamber and absence of a verge may be the result
of spatial constraints associated with the overall
reduction in size of the penial lumen. However,
this proposition needs to be more thoroughly
tested through a comprehensive revision of all the
hadroid camaenids.
DISCUSSION
The relationships of M. helicostracum to other
hadroid camaenids from eastern Queensland still
need to be fully confirmed but appear to rest with
a widespread radiation consisting of rainforest-
bound species now separated by large tracts of
drier countryside. These species are part of
Sphaerospira s.l. and are characterised by having
banded shells and animals with orange to red
mantles. In the southern part of the Wet Tropics
these species inhabit the drier rainforest between
Innisfail and Townsville; in MEQ they occur in
drier araucarian vine forests and vine thickets
NEW CAMAENID LAND SNAIL GENUS FROM THE WET TROPICS
between Mackay and Bowen; and in SEQ possib-
ly more distantly related species (also with red-
dish mantles) live in a variety of dry rainforests.
These rainforests are peripheral to core wet,
upland refugia and presumably land snails in
these environments would have been more
strongly affected by climatic shifts. The Wet
Tropics and MEQ radiations of Sphaerospira s.l.
are separated by an extensive relatively dry cor-
ridor between Bowen and Townsville, NEQ
where coastal ranges are absent and orographic
rainfall is low (Dick, 1974). In S Queensland a
dry corridor in the St Lawrence-Gladstone area
separates SEQ and MEQ radiations. Similarly,
Bakers Blue Mtn and Hanns Tableland are
separated from the main massif of the Wet
Tropics by dry valleys and ridges (Nix, 1991).
Thus biogeographic interpretation of the history
of M. helicostracum and its allies would appear
to involve climate-induced fragmentation of rain-
forest communities. This view is supported by
Bishop (1981) who suggested that the history of
Sphaerospira s.l. was intimately linked to the
deterioration of the climate and involved sub-
division of once more extensive ranges.
Itis generally accepted that the Camaenidae are
a northerly derived element of the Australian land
snail fauna (McMichael & Iredale, 1959; Bishop,
1981; Solem, 1992). In the east, dispersal would
have been from the New Guinea region across the
Torres Strait following collision of the Australian
continental plate with the Asiatic and Pacific
Ocean plates (Doutch, 1972). Sphaerospira s.l.
today extends from the Wet Tropics to northern
NSW indicating a long history of dispersal. The
moisture dependence and relative low vagility of
land snails are important limitations to their dis-
persal and distribution in eastern Australia
(Bishop, 1981; Stanisic, 1994). The present day
confinement of these hadroid camaenids to rain-
forest indicate that tracts of dry sclerophyll forest
are effective dispersal barriers. The contraction
of rainforest which led to the creation of barriers
to their dispersal began in the Miocene with the
onset of major aridity episodes (Kemp, 1981).
These barriers would have reached their acme in
the Plio-Pleistocene when more rapid climatic
change isolated mesic communities in montane
refugia (Galloway & Kemp, 1981). The effects
on Sphaerospira s.l. should have been dramatic
and it is not surprising that current distribution
patterns are complex and that differentiation of
the group appears to have occurred on opposite
sides of major arid corridors in the St Lawrence-
361
Gladstone and Bowen-Townsville areas (Bishop,
1981).
M. helicostracum shows a level of differentia-
tion which suggests that its isolation from its
allies has been long-term, possibly in the same
time frame as the separation of the red-mantled
NEQ and MEQ species groups. Recent fluctua-
tions in rainforest distribution (c. 120,000 yBP)
within the Wet Tropics (Kershaw, 1981) do not
appear to have had an effect on the distribution of
M. helicostracum. Nix (1991) proposed a pos-
sible connection between the Bakers Blue
biogeographic unit and other units to the north
and east during cool-wet and warm-wet phases of
the last 10,000 years, yet M. helicostracum
remains isolated in the two outliers. Either the
proposed connection did not exist or was too brief
to allow dispersal of this species.
It is probably significant that Monteithosites
and its allies are absent from the core of the Wet
Tropics. The present day preference of
Sphaerospira s.l. with reddish to orange coloured
animals for drier rainforest types often dominated
by hoop-pine (Araucaria cunninghamii) (unpubl.
data) suggests a disposition for a very particular
environmental regime. This is most notable in
MEQ and SEQ where araucarian microphyll vine
forest is more widespread. A. cunninghamii (and
its associated dry rainforest type) is relict in NEQ
(Tracey, 1982) although it has shown consider-
able fluctuation in occurrence since the Plio-
Pleistocene (Kershaw, 1981).
The absence of Sphaerospira s.l. with orange
to red-mantled animals from the Atherton
biogeographic unit (now dominated by Hadra
and its allies) and from other moist upland refugia
(higher peaks of the Wet Tropics, NEQ and Eun-
gella, MEQ) might indicate that this group only
ever occupied marginally wet rainforest types
peripheral to the larger massifs. Equally it may
indicate displacement from wetter types by the
dark animal hadroids which now dominate the
truly moist refugia from SEQ to MEQ. Lack of
knowledge about the inter-relationships of these
species currently makes it difficult to be sure of
the accuracy of either alternative. However, some
speculation is possible.
The bioclimatic analyses of Kershaw & Nix
(1988) support the view that there was no recent
connection between Bakers Blue Mtn and Hanns
Tableland. In contrast, presence of M.
helicostracum on both these outliers indicates a
past connection which most likely predates the
time scale of their analyses. Connections between
the outliers and with the Atherton unit in the south
362
were less likely during recent times but could
have been present in the Plio-Pleistocene when
araucarian vine forests may have been in com-
parative ascendency (Kershaw et al., 1991).
However, absence of related camaenids from the
Herberton Range (a subunit of the Atherton unit)
which has summits in excess of 1000m and ex-
tensive areas of rainforest, might suggest that the
group never colonised this region. These summits
should have provided moist refugia during the
driest periods of the Plio-Pleistocene. Yet
presence of the probable nearest relatives of M.
helicostracum in the Cardwell, Seaview and
Paluma Ranges points to a NW-SE dispersal
track on the drier western edge of the Wet
Tropics. In these circumstances it is a strong
possibility that the suitable dispersal corridors
which may have existed on the western edge of
the Atherton have been lost since the Miocene.
The preference of M. helicostracum and its pur-
ported allies for a very specific environmental
regime which is now restricted in the north would
seem to provide support for this proposition. It is
possible that the differentiation and speciation
within the group needs to be viewed as occurring
along very subtle environmental gradients
defined by rainforest structural types. A revision
of the hadroid camaenids of the Wet Tropics is
needed to more fully understand past dispersal
routes but indications are that these land snails
have the potential to more accurately define cur-
rent hypotheses.
LITERATURE CITED
BISHOP , M.J. 1978. Anatomical notes on the
Australian camaenid land snail Sphaerospira
fraseri (Griffith & Pidgeon). Journal of the
Malacological Society of Australia 4: 9-21,
1981. The biogeography and evolution of Australian
land snails. Pp. 924-954. In Keast, A. (ed.),
Ecological biogeography of Australia. (W. Junk
: The Hague).
BURCH, J.B. 1976. Outline of classification of
Australian terrestrial molluscs (native and intro-
duced). Journal of the Malacological Society of
Australia 3:127-156.
DICK, R.S. 1974. Frequency patterns of arid, semi-arid
and humid climates in Queensland. Capricornia 1:
21-30.
DOUTCH, H.F. 1972. The palaeogeography of north-
ern Australia and New Guinea and its reelevance
to the Torres Strait area. Pp. 1-10. In Walker, D.
(ed.), Bridge and barrier: the natural and cultural
history of Torres Strait. (Australian National
University:Canberra).
GALLOWAY, R.W. & KEMP, E.M. 1981. Late
Cainozoic environments, Pp. 52-80. In Keast, A.
MEMOIRS OF THE QUEENSLAND MUSEUM
(ed.), Ecological biogeography of Australia. (W.
Junk: The Hague).
GODWIN, M. 1986, Vegetation of northern part of
Portion 1 Layland. Unpublished report for the
Queensland Department of Environment and
Heritage. (Queensland Department of Environ-
ment and Heritage).
IREDALE, T. 1933. Systematic notes on Australian
land shells. Records of the Australian Museum
19:37-59.
1937. A basic list of the land Mollusca of Australia.
Part II. Australian Zoologist 9:1-39.
KEMP, E.M. 1981. Tertiary paleogeography and the
evolution of the Australian environment. Pp.33-
49. In Keast, A. (ed.), Ecological biogeography of
Australia. (W, Junk: The Hague).
KERSHAW, A.P. 1981, Quaternary vegetation and en-
vironments. Pp. 81-101. In Keast, A. (ed.),
Ecological biogeography of Australia .(W. Junk:
The Hague).
KERSHAW, A.P. & NIX, H.A. 1988. Quantitative
palaeoclimatic estimates from pollen data using
bioclimatic profiles of extant taxa. Journal of
Biogeography 15: 589-602.
KERSHAW, A.P., SLUITER, I.R., McEWAN
MASON, J., WAGSTAFF, B.E. & WHITELAW,
M. 1991. The history of rainforest in Australia-
evidence from pollen. Pp. 1-16. Werren, G &
Kershaw, P. (eds), The rainforest legacy.
Australian national rainforest study. (Australian
Government Publishing Service: Canberra).
McMICHAEL, D.F. & IREDALE, T. 1959, The land
and freshwater Mollusca of Australia. Pp.224-
245. In Keast, A., Crocker, R.L. & Christian, C.S.
(eds), Biogeography and ecology in Australia. (W.
Junk: The Hague).
NIX, H.A. 1991. Biogeography: pattern and process.
Pp. 11-39. In Nix, H.A. & Switzer, M.A. (eds),
Rainforest animals. Kowari 1. (Australian Nation-
al Parks and Wildlife Service: Canberra).
PACE, S. 1901. Note on the anatomy of Thersites
(Hadra) bipartita (Fer). Proceedings of the
Malacological Society of London 4:205- 207.
PILSBRY, H.A. 1894, Manual of Conchology, 2, 9:
49-160.
SMITH. B.J. 1992. Non-marine Mollusca. Pp.1-405. In
Houston, W.W.K. (ed.), Zoological catalogue of
Australia. Vol.8. (Australian Government Print-
ing Service:Canberra).
SOLEM, A. 1979, Camaenid land snails from Western
and central Australia (Mollusca: Pulmonata:
Camaenidae). I. Taxa with trans- Australian dis-
tributions. Records of the Western Australian
Museum, Supplement 10:1-142.
1985. Camaenid land snails from Westren and
central Australia (Mollusca: Pulmonata:
Camaenidae). V. Remaining Kimberley genera
and addenda to the Kimberley. Records of the
Western Australian Museum, Supplement
20:707-981.
1992. Camaenid land snails from southern and east-
NEW CAMAENID LAND SNAIL GENUS FROM THE WET TROPICS 363
ern South Australia excluding Kangaroo Island. 1994. A preliminary report on the land snails
Records of the South Australian Museum, occuring in the Wet Tropics, northeastern
Monograph Series 2:1-338. Queensland. Unpublished report for the Wet
STANISIC, J. 1994. The distribution and patterns of Tropics Management Autority. (Queensland
species diversity of land snails of eastern Museum : Brisbane).
Australia. Memoirs of the Queensland Museum TRACEY, J.G. 1982. The vegetation of the humid
36: 207-214. tropical region of north Queensland. (CSIRO:
STANISIC, J., EDDIE, C., HILL, A., & POTTER, D. Melbourne).
364
FRUIT EATING BY THE GECKO GEHYRA DUBIA IN
TOWNSVILLE. Memoirs of the Queensland Museum
39(2): 364. 1996:-This note describes opportunistic observa-
tions of fruit-eating by G. dubia occupying a residence in
suburban South Townsville. At approximately 1745hr on 14
March 1996 a G. dubia was observed feeding on ripe sugar
bananas suspended within the residence. Individual bananas
had ripened on the bunch and had partially detached, being
suspended by one strip of peel only. The ripe flesh was
exposed where the skin had given way, and it was on this flesh
that the gecko was feeding; tearing large chunks of the fruit
off with it's mouth and consuming them. On three subsequent
occasions, as the fruit became completely detached from the
bunch and was stored on a kitchen table, geckoes were startled
from amongst them although fruit consumption was never
observed before the gecko(s)? took off in alarm. We do not
know if they were the same or different individuals. No
invertebrates were seen on the bananas at these times and we
suspect that the geckcoes were once again feeding on the fruit.
Greer (1989) summarised published observations of
Gecko diet. Non-animal foods variously eaten by Christinus
guentheri, C. marmoratus, Heteronotia binoei and
Lepidodactylus lugubris, include sap, nectar, jam, honey and
sugar. Couper et al. (1995) added to the list of sapfeeding
MEMOIRS OF THE QUEENSLAND MUSEUM
Geckos when they described sapfeeding by Gehyra dubia in
the wild. The only published reference to fruit-eating by an
Australian gecko that I am aware of is King & Horner's (1993)
observation of G. australis feeding on pulpy fruit. As such,
this note provides further evidence of fruit-eating by another
species of Australian gecko in the wild.
Literature Cited
Couper, P.J., Covacevich, J.A. & Wilson, S.K. 1995. Sap
feeding by the Australian gecko Gehyra dubia.
Memoirs of the Queensland Museum 38(2): 396.
Greer, A. 1989. Gekkonidae, Geckos. In Greer, A. (ed). The
Biology and evolution of Australian lizards. (Surrey
Beatty & Sons: Chipping North, NSW).
King, M. & Horner, P. 1993. Family Gekkonidae. Pp. 221-
233. InGlasby, C.J., Ross, G.J.B. & Beesley, P.L. (eds),
Fauna of Australia vol 2a, Amphibia and Reptilia.
(Australian Government Publishing Service: Canber-
ra).
Scott Burnett & Jacqueline Nolen, 9 Nelson St, South
Townsville 4810, Queensland; 23 April 1996.
REVISION OF AUSTRALOXENELLA HOWDEN & STOREY IN AUSTRALIA
(COLEOPTERA: SCARABAEIDAE: APHODIINAE)
R.I. STOREY & H.F. HOWDEN
Storey, R.I. & Howden, H.F. 1996 07 20: Revision of Australoxenella Howden & Storey in
Australia (Coleoptera: Scarabaeidae: Aphodiinae). Memoirs of the Queensland Museum
39(2): 365-380. Brisbane. ISSN 0079-8835.
Australoxenella Howden & Storey, the only Australian member of the aphodiine tribe
Stereomerini, is revised. Eleven species are described of which the following are new:
concinna, kalpara, midgee, mirreen, moogoon, peckoriim, teeta, wurrook, zborowskii.
Relationships between species are discussed. Most new specimens were taken in flight
interception traps. No information is available on the biology of these suspected ter-
mitophiles. [.] Coleoptera, Scarabaeidae, Aphodiinae, Australoxenella, taxonomy.
R.I. Storey, Department of Primary Industries, PO Box 1054, Mareeba, Queensland 4880,
Australia; H.F. Howden, Department of Entomology, Canadian Museum of Nature, P.O.
Box 3443, Station D, Ottawa, Ontario KIP 6P4 CANADA; received 20 September 1995.
Australoxenella Howden & Storey, 1992 was SYSTEMATICS
erected for two unusual specimens representing
two species of aphodiine Scarabaeidae taken in Order COLEOPTERA
the far north of the Northern Territory. The Family SCARABAEIDAE
Stereomerini was erected for Australoxenella, Subfamily APHODIINAE
Bruneixenus Howden & Storey, and Stereomera
Arrow and Termitaxis Krikken, previously
placed in the tribe Rhyparini. Collecting in NT
and in Queensland since 1992 has yielded 96
specimens of Australoxenella, including nine
new species. Most were caught in flight intercep-
tion traps, although several were from malaise
traps and one from a berlesate. Habitat ranged
from mixed eucalyptus woodland to dense rain-
forest. With revision of Australian Aphodiinae in
progress (Stebnicka & Howden, 1994, 1995)
revision of Australoxenella was necessary in light
of the new material. Although Howden & Storey
(1992) speculated that all Stereomerini were ter-
mitophiles, no direct evidence of this life habit in
Australoxenella has surfaced.
METHODS AND MATERIALS
Terminology follows Howden & Storey (1992).
Scale bars on SEM photographs are in milli-
metres. Collection abbreviations used are:
ANIC-Awustralian National Insect Collection,
Canberra, A.C.T.; HAHC-H. & A. Howden Col-
lection, Ottawa, Ontario; ISEA-Institute of Sys-
tematics and Evolution of Animals, Cracow,
Poland; NTMA-Northern Territory Museum of
Arts and Sciences, Darwin; QMBA-Queensland
Museum, Brisbane; QPIM-Department of
Primary Industries, Mareeba, Queensland; SBPC
—S. B. Peck Collection, Ottawa, Ontario.
Australoxenella Howden & Storey, 1992
Australoxenella Howden & Storey, 1992:1811.
TYPE SPECIES. Australoxenella humptydooensis
Howden & Storey, 1992.
DESCRIPTION. Head. Dorsally broad, flat-
tened, feebly convex, about twice as wide as long.
Clypeus anteriorly with broadly inflexed edge,
obtusely angulate medially, nitid. Head dorsally,
with distinct median and lateral grooves, length
of median groove variable, lateral grooves slight-
ly deeper, c. 1/3 of distance to outer edge of gena,
length of lateral grooves variable, lateral grooves
slightly convergent anteriorly for apical 1/4—2/3
length; surface of head dorsally, except for
grooves, with close, appressed, circular scales
giving granulate appearance. Gena not obviously
delimited, inner edge indicated by small, non-
granulate marking near summit of convex area.
Pronotum. Strongly, irregularly convex with
transverse sulcus near middle, extending each
side to at least lateral thirds, variable in length,
depth and width; area anterior to sulcus with 5 or
usually 7 longitudinal grooves, inner 5 grooves
variable in depth and width, rounded ridges be-
tween grooves variable, inner 2 ridges often
elevated and forming rounded prominence just
anterior to transverse sulcus; equivalent median
surface posterior to sulcus with only trace of
median groove, always strongly convex and more
strongly elevated than any other part of dorsum;
3 variable lateral posterior grooves on each side
parallel, and extending into sulcus; pronotum
laterally with sides almost parallel; anterior
angles slightly acute, posterior angles almost 90°,
abruptly rounded; pronotal surfaces with close,
appressed, circular scales, except grooves and
sulcus. Scutellum minute, surface shining.
Elytron. Moderately to strongly ridged on disc,
each elytron with sutural and 2-3 lateral intervals
elevated with shallow to deep U-shaped depres-
sions between; distinct striae absent or with one
stria below feeble humeral umbone, usually
merging with lateral bead near middle of lateral
edge; epipleuron broadly inflexed, parallel-sided
or tapering towards apex, often longitudinally
concave near metacoxa; elytral apices not greatly
modified, explanate in some species, conjointly
broadly rounded; covered by close, appressed,
circular scales, density variable, always
numerous along crests of ridges and outside
fourth ridge, scattered to dense on U-shaped
depressions; depressions dull, alutaceous. Meta-
thoracic wings long, functional.
Pygidium, Surface ventral, flattened, wider
than long, with broadly rounded apex anterior in
position.
Ventral surface. Antenna 9-segmented, club
3-segmented, equal in length to basal 6 segments
of antenna combined; area surrounding antenna
deeply concave. Mentum medially concave,
anterior margin almost straight with small
median projection. Maxillary palpus apparently
3-segmented, apical segment lanceolate. Man-
dibles thin, blade-like. Eye small, somewhat tri-
angular, approximately 10 facets long x 18 wide,
not visible dorsally, covered by pronotum when
head retracted. Prosternum laterally with
alutaceous to subnitid surface, anterior edge near
antennae with a raised marginal bead; median
prosternal process anteriorly strongly elevated,
apex expanded, process posteriorly lanceolate,
medially carinate, with obtusely angled sides.
Mesosternum moderate in size, triangular, nar-
rowed laterally. Metasternum between meso-
coxae narrow, blade-like, posterior to mesocoxae
abruptly widened, flat, shallowly concave in
posterior median 1/2, anteriorly behind meso-
coxae with transverse marginal bead, surface
alutaceous to nitid; metacoxae contiguous. Ab-
domen with 4 (d) or 5 (9) segments visible
medially, 6 visible laterally, apical segment
longer in midline than other segments combined.
Legs. Femora wide, flattened. Protibia with 2
MEMOIRS OF THE QUEENSLAND MUSEUM
teeth on outer edge, one apical, one subapical,
subapical occasionally obsolete. Meso- and
meta-tibiae flattened, approximately twice as
long as wide; outer narrow edge of each tibia with
irregular longitudinal rows of punctures, each
puncture with a minute yellowish seta; each tibia
with 2 minute apical spurs, may be large setae;
small apical setae also present. Protarsus 5-seg-
mented, with several conspicuous setae at ventral
apex of each segment; claws normal. Meso - and
meta-tarsi 4-segmented, appearing to be 3-seg-
mented, basal segment shorter than second seg-
ment; claws reduced in thickness.
KEY TO SPECIES OF AUSTRALOXENELLA
1 Pronotum anteriorly with median, longitudinal
groove similar in width to groove on each side
and nearly as wide as adjacent ridges; anteriorly
with ridge on each side of median groove at
most only slightly higher than adjacent ridge
(Fig. 1A); elytral epipleura nearly constant in
width to apices (except teeta sp.nov.)
AWE LAE MH LIUC (Group 1) 2
Pronotum anteriorly with median, longitudinal
groove narrow, more so than lateral grooves;
anteriorly with ridge on each side of median
groove more strongly elevated posteriorly than
adjacent outer ridge (Fig. 5C); elytral epipleura
noticeably narrowing towards apices.
p peat nigale ge iaaa ION COND in (Group 2) 7
2 (1)Each elytron with sutural and three lateral
ridges and a single stria in basal outer quarter
Each elytron with sutural and two lateral ridges,
stria in basal outer quarter absent (Fig. 1B);
NOld ga gos nog BRS concinna sp. nov.
3 (2)Pronotal ridges anterior to transverse sulcus
lacking clusters of erect spur-like setae... . 4
Pronotal ridges anterior to transverse sulcus each
with four clusters of closely appressed, erect
spur-like setae (Fig. 1C); N Qld
Me iuacta WA eat ote zborowskii sp. nov.
4 (3)Pronotum in lateral view with median conical
prominence posterior to transverse sulcus much
more strongly elevated than area anterior to sul-
cus; lateral pronotal and elytral margins lacking
distinct fringe of yellowish setae
Pronotum in lateral view with median conical
prominence posterior to transverse sulcus only
slightly more elevated than area anterior to sul-
cus; lateral pronotal and elytral margins with dis-
tinct fringe of yellowish setae (Figs 1E, 1F);
N Qld teeta sp. nov.
5 (4)Elytron with crests of longitudinal ridges on
disc narrow, subacute (Figs 3B, 3D)
Elytron with crests of longitudinal ridges on disc
REVISION OF AUSTRALOXENELLA
rounded, broad; Bathurst Island, NT
DS 0.44 A. bathurstensis Howden & Storey
6 (5)Elytral surface anteriorly between suture and ad-
jacent raised (sutural) ridge with 2 rows of
scales; next depression between sutural ridge
and first lateral ridge with close, irregular,
transverse rows of five scales; narrower species,
length to width ratio 2.2:1 (Fig. 3B); NT
295, 81 T S MES D reis © moogoon sp. nov.
Elytral surface near base between suture and ad-
jacent raised (sutural) ridge with 3 irregular rows
of scales; next depression between sutural ridge
and first lateral ridge with close irregular,
transverse rows of 6-8 scales; broader species,
length to width ratio 2:1 (Fig. 3D); NT
wurrook sp. nov.
7 (1)Pronotum in lateral third posterior to transverse
sulcus transversely convex anterior to posterior
margin
Pronotum in lateral third posterior to narrow
transverse sulcus broadly, shallowly concave be-
tween sulcus and posterior margin (Fig. 4A);
N Qld midgee sp. nov.
8 (7)Posterior prosternal lanceolate process with
central carina blade-like, very little expanded at
middle (Fig. 2F); elytral U- shaped depressions
with regular transverse rows of scales in fine
grooves, remaining surface without scales or
setae (Fig. 2E; N Qld ..... kalpara sp. nov.
Posterior prosternal lanceolate process with
central carina expanded at middle (Fig. 5E);
elytral U-shaped depressions with scales not
grouped in transverse rows in fine grooves; NT
9 (8)Metasternum medially just posterior to
mesocoxae with narrow transverse ridge with
declivous posterior edge forming anteriorly-
directed abrupt, obtuse angle (Fig. 5E); third
lateral ridge on each elytron with inner parallel
linesofscales crie seme nn 10
Metasternum medially just posterior to
mesocoxae with broadened transverse ridge,
posterior edge of ridge almost straight or feebly,
anteriorly, arcuate medially (Fig. 4E); third
lateral ridge on each elytron lacking inner paral-
lel lines of scales (Fig. 4D); NT
peckorum sp.nov.
10 (9)Meso- and meta-femora ventrally with
numerous oval, appressed scale-like setae; elytra
in lateral view with sutural ridges distinctly ar-
cuately elevated at posterior third (Fig. 5B); NT
Pa-c.cield An. ae mirreen sp. nov.
Meso- and meta-femora ventrally with scale-like
selae present only near posterior edges, with
scatlered, slender setae elsewhere; elytra in
lateral view with sutural ridges only feebly ar-
cuately elevated at posterior third (Fig. 5D); NT
A. humptydooensis Howden & Storey
Australoxenella concinna sp. nov.
(Figs 1A, 1B, 6A)
ETYMOLOGY. For the beautiful appearance.
MATERIAL EXAMINED. HOLOTYPE QMBT
13424 (sex not determined) from 17°24’S, 145°41’E,
Westcott Rd, Topaz, Qld, 6.xii.1993-25.ii.1994, Mon-
teith, Cook, Janetzki, RF intercept 680m.
DESCRIPTION. Length 2.9mm, greatest width
2.0mm. Dark reddish brown dorsally and ventral-
ly.
Head. Median groove reaching anterior edge,
lateral grooves reaching anterior edge and slight-
ly converging for anterior 1/2; surface of head
with small oval-elongate marking above each
eye; anterior edge of head between genae with a
fringe of short, adjacent, yellowish, flattened
setae.
Pronotum. Four central anterior rounded ridges
equal in size, none elevated; median, first and
second lateral pairs of grooves equal in width and
depth, third lateral pair of grooves fine, joining
anterior pronotal edge c. 1/3 distance from
anterior angles to centre line; posterior median
groove not visible in anterior 1/2, adjacent
rounded ridges slightly elevated, crest acute, ad-
ditional fourth pair of lateral grooves present,
strong from centre of sculptured area to posterior
angles; transverse sulcus strong narrow, running
to lateral edge of pronotum, slightly widened
posteriorly for inner 1/2; widened area of sulcus
bare, anterior edge of sulcus and most grooves
with short adjacent yellowish setae, groups of
longer similar setae in centre of sculptured area;
small clusters of spur-like setae on anterior ridges
and posterior prominence, not well developed;
lateral pronotal edges from anterior angles to
posterior angles with a distinctive fringe of ad-
jacent, yellowish, flattened setae.
Elytron. Broad, flattened, explanate; with
sutural and 2 lateral ridges, ridges weakly
elevated, shortened, sutural longest c.2/3 length
of elytron, second lateral shortest c.1/2 length of
elytron; U-shaped depressions very shallow,
barely detectable; entire dorsal surface of elytron
with close, appressed, circular scales, crest of
each ridge with a distinct row of adjacent, ap-
pressed, circular scales; lateral margin of elytron
with a distinct fringe of adjacent, flattened, yel-
lowish setae, this in the form of a double row on
basal 1/6 which could indicate position of a ves-
tigial stria; epipleuron wide, slightly narrowed in
apical 1/6, nitid, punctures and setae fine,
stronger in apical 1/2.
Pygidium. Nitid with small, setate punctures.
368 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG.1. A,B, Australoxenella concinna sp. nov. A, head and pronotum, lateral view. B, pronotum and elytra, dorsal
view. C,D, Australoxenella zborowskii sp. nov. C, head and pronotum, lateral view. D, elytra, dorsal view. E,F,
Australoxenella teeta sp. nov. E, head and pronotum, lateral view. F, elytra, dorsal view.
Ventral surface and legs. Abdominal segments
nitid, posterior 1/2 of large segment with small
setate punctures; metasternum nitid, central area
with small setate punctures; meso- and meta-
femora nitid, ventral surfaces with fine setate
punctures, more numerous on posterior 1/2;
meso- and meta-tibiae nitid, a few fine setate
punctures on outer 1/2, lower lateral margin of
each metatibia with a row of short, adjacent,
flattened setae.
REVISION OF AUSTRALOXENELLA
COMMENTS. Australoxenella concinna sp.
nov. is closest to A. zborowskii sp. nov., sharing
with it the setal fringes on the pronotum and
elytra, and the clusters of spur-like setae on the
pronotal sculpturing. It differs from that species
in the additional fringe of setae on the anterior
margin of the head, the pronotal clusters being not
as strongly developed, the elytra having circular
scales over the entire surface and the lack of a
third lateral pair of elytral ridges. Australoxenella
concinna is the most southerly species of the
genus; it was trapped in dense rainforest (G.
Monteith, pers. comm). Several hundred trap-
months using flight interception traps by the
senior author in many similar Atherton
Tablelands sites over many years have failed to
capture further specimens.
Australoxenella zborowskii sp. nov.
(Figs 1C, ID, 6A)
ETYMOLOGY. For Paul Zborowski, collector of
several species described in this paper.
MATERIAL EXAMINED. HOLOTYPE ANIC113,
sex not determined, from 11° 41'S, 142° 42'E, 14km
ENE of Heathlands, Qld, 21.i.-19.ii.1994, P.
Zborowski, flight intercept trap.
DESCRIPTION. Length 2.6mm, greatest width
1.4mm. Reddish brown dorsally and ventrally.
Head. Three grooves all reaching anterioredge,
lateral grooves converging slightly for apical 1/3
length; surface of head with narrow, elongate,
nitid marking above each eye.
Pronotum. Sculptured area anterior to
transverse sulcus wider, about 1/2 width of
pronotum, grooves strong, intergroove rounded
ridges of equal height, not raised around median
groove, third lateral pair of grooves weak, almost
reaching anterior angles; prominence around
median groove posterior to transverse sulcus
strong, apex acute, additional fourth pair of
lateral grooves present, running from middle por-
tion of sulcus almost to posterior angles;
transverse sulcus constant in width, narrowing
towards apices, almost reaching lateral edges of
pronotum; grooves nitid, middle region of sulcus
with prominent yellow setae; pronotal disc with
spur-like clusters of erect, yellow setae - four
groups on each of the four narrow anterior ridges,
five each on adjacent wider ridges, less obvious
clusters on posterior central prominence and
posterior lateral raised areas; lateral pronotal
edges from anterior angles to just past posterior
angles each with a distinct fringe of short, ad-
jacent, flattened, yellowish setae.
369
Elytron. Disc flattened, explanate, ridges not
very high and U- shaped depressions shallow;
with sutural and three lateral ridges, sutural ridge
very slightly stronger, third lateral ridge strong,
running from base to c. 3/8 distance along lateral
margin of elytron; area of elytron outside stria
c.1/2 as wide as in most other species; dorsal
surface outside third lateral ridge with close, ap-
pressed, circular scales, scales also numerous
along remaining margin of elytron and up to
apices of elytral ridges, crest of each elytral ridge
with a single row of overlapping scales along
entire length resulting in a carinate appearance;
U-shaped depressions between lateral ridges with
regular transverse, single, occasionally double,
rows of scales joining ridges; lateral margin of
elytron from base to apex with a distinct fringe of
short, adjacent, flattened yellowish setae; lateral
stria also with a reduced fringe row; epipleuron
broad, slightly narrowed at apex, bare, nitid.
Pygidium. Smaller than in other species, with
scattered setate punctures.
Ventral surface and legs. Meso- and meta-
tibiae with a few minute setate punctures,
posterior outer margin with a short setal fringe;
ventral surfaces of meso- and meta-femora with
scattered, medium, setate punctures.
COMMENTS. This species is known from a
single specimen taken in notophyll vine forest in
a flight interception trap (P. Zborowski, pers.
comm.). Characters separating A. zborowskii sp.
nov. are discussed under A. teeta sp. nov. and A.
concinna sp. nov. The distinctive pronotal and
elytral fringes of flattened setae are found only in
three Queensland species, A. zborowskii, A. teeta
and A. concinna.
Australoxenella teeta sp. nov.
(Figs 1E, IF, 6A)
ETYMOLOGY. An Aboriginal word for insect.
MATERIAL EXAMINED. HOLOTYPE ANIC114,
sex not determined, from 13? 39'S, 142? 40'E (GPS),
2km N Rokeby, Qld, 15.ii.-18.1i1.1994, P. Zborowski
& M. Shaw, flight intercept trap.
DESCRIPTION. Length 2.6mm, greatest width
1.3mm. Reddish brown dorsally and ventrally.
Head. Median groove almost reaching anterior
edge, lateral grooves slightly shorter and con-
verging for anterior 1/2; surface of head with
small elongate, nitid marking above each eye.
Pronotum. Anterior rounded ridges adjacent to
median groove weakly elevated posteriorly, third
lateral pair of grooves weak, almost straight to
370 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG.2. A-C, Australoxenella bathurstensis Howden & Storey. A, head and pronotum, dorsolateral view. B, elytral
apex, dorsolateral view. C, posterior half, ventral view. D-F, Australoxenella kalpara sp. nov. D, head and
pronotum, lateral view. E, elytron, dorsal view. F, mesosternum, ventral view. (Figs 2A,B from Howden &
Storey, 1992)
anterior angles; ridges adjacent to median groove — ly; transverse sulcus not widened at apices, sulcus
posterior to transverse sulcus forming anelevated reaching 3/4 distance to lateral edge of pronotum
prominence, median groove not visible anterior- from centre line; sulcus medially and centre of
REVISION OF AUSTRALOXENELLA
sculptured area with dense, yellow setae; lateral
pronotal edges from anterior angles to just past
posterior angles with a distinct fringe of short,
adjacent, flattened, yellowish setae.
Elytron. Disc with sutural and three lateral
ridges, ridges not strong, sutural ridge most
developed, third lateral ridge reduced and meet-
ing margin of elytron about 1/2 distance to apex;
lateral stria reaching margin of elytron about 3/8
distance to apex; dorsal surface outside of third
lateral ridge with close, appressed, circular
scales, crests of other ridges also with scales
including a row of adjacent scales on crest of
sutural and first lateral, U-shaped depressions
between lateral ridges with spaced, transverse,
single rows of scales joining ridges, surface dull,
bare between; margin of elytron from base to
apex with a distinct fringe of short, adjacent,
flattened, yellowish setae; epipleuron narrowing
only slightly towards apex, surface bare, subnitid.
Pygidium. Nitid with only scattered setate
punctures.
Ventral surface and legs. Tibiae wide, flat-
tened, nitid, impunctate on broad surfaces;
femora broad, nitid, ventral surfaces of meso- and
meta-femora with numerous setate punctures.
COMMENTS. Australoxenella teeta sp. nov.
was taken 1n open forest in a flight interception
trap set in red earth woodland dominated by
Eucalyptus tetradonta and E. mesophila (P.
Zborowski, pers. comm.) The border fringe of
flattened setae on the lateral edges of the
pronotum and elytra are well developed only in
A. teeta, A. concinna sp. nov. and A. zborowskii
sp. nov. All three species are nitid ventrally and
have the prominence on the pronotal surface
anterior to the transverse sulcus very reduced or
absent. Australoxenella zborowskii and A. con-
cinna have the elytra more explanate, the setal
fringes slightly longer, the sculptured area in
front of the pronotal transverse sulcus wider and
less raised and prominent clusters of spur-like
setae on the anterior pronotal ridges, allowing
easy separation from A. teeta.
Australoxenella bathurstensis Howden &
Storey, 1992
(Figs 2A-C, 6B)
Australoxenella bathurstensis Howden & Storey,
1992:1813.
MATERIAL EXAMINED. HOLOTYPE
NTMAI287, from Cape Fourcroy, Bathurst Island, NT,
26.x.-3.xi.1979, P. Horner & I. Archibald.
371
DESCRIPTION. Length 3.2mm, greatest width
1.6mm. Brown dorsally, reddish brown ventrally.
Head. Median groove reaching 3/4 distance to
anterior edge, lateral grooves slightly shorter and
slightly converging for anterior 1/2; surface of
head with narrow, elongate marking above each
eye.
Pronotum. Median, first and second lateral
pairs of grooves anterior to transverse sulcus
strong, equal in depth and width, third lateral pair
of grooves fine, joining anterior edge of
pronotum c.1/3 distance from anterior angles to
centre line, anterior four central ridges strong,
rounded, equal in height and width; median
groove posterior to transverse sulcus barely
visible on posterior half only, first lateral pair
stronger, second and third lateral pairs strongest,
ridges either side of median groove forming
strong prominence, crest rounded; transverse sul-
cus narrow, barely widened toward centre; area
between anterior and posterior sculpturing with
groups of appressed, yellow setae.
Elytron. Broad, flattened, explanate; sutural
and three lateral ridges not strong, crests low,
rounded, third lateral ridge forming lateral mar-
gin ofelytron just past 1/2 distance to apex; lateral
stria joining lateral margin of elytron just before
1/2 distance to apex; close, appressed, circular
scales cover surface outside third lateral ridge and
along crests of all four ridges, U-shaped depres-
sions with few scattered scales mostly in basal
1/2, surface of depressions dull; epipleuron
widened to apex, dull to subnitid on apical 1/3,
circular scales along outer margin.
Pygidium. Subnitid with scattered small setate
punctures,setae fine, flattened.
Ventral surface and legs. Abdominal segments,
metasternum, and meso- and meta- femora sub-
nitid with scattered small punctures each with a
fine seta, meso- and meta-tibiae subnitid, with
setate punctures only near outer margins.
COMMENTS. Australoxenella bathurstensis
Howden & Storey can be distinguished from the
other species of Group 1 by the form of the elytra,
with all ridges low and rounded, with associated
scales unordered, and largely bare U-shaped
depressions. It is still known only from the
holotype.
Australoxenella moogoon sp. nov.
(Figs 3A, 3B, 6B)
ETYMOLOGY. An Aboriginal word for beetle.
MATERIAL EXAMINED. HOLOTYPE ANIC115,
sex not determined, from 33km E of Jabiru, Arnhem-
372
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG.3. A,B, Australoxenella moogoon sp. nov. A, head and pronotum, lateral view. B, elytra, dorsal view. C,D,
Australoxenella wurrook sp. nov. C, head and pronotum, lateral view. D, elytra, dorsal view,
land, NT, 15-23.xii.1993, S. & J. Peck, 93-130, 5 FITs,
Podocarp Canyon, rainforest. PARATYPES (5) in
HAHC, QPIM, same data as Holotype.
DESCRIPTION. Length 2.4—2.5mm, greatest
width 1.3-1.4mm. Reddish brown dorsally,
slightly lighter ventrally.
Head. Median groove almost reaching anterior
edge, lateral grooves slightly shorter and slightly
converging for anterior 1/2; surface of head with
narrow elongate marking above each eye.
Pronotum. Anterior rounded ridges adjacent to
median groove only very slightly elevated
posteriorly, four ridges and median, first and
second lateral pairs of grooves equal in ap-
pearance, third lateral pair of grooves fine reach-
ing anterior pronotal edge c.1/3 distance from
anterior angles to centre line; ridges either side of
median posterior groove forming a subacute
prominence, anterior portion of median groove
scarcely visible, other grooves strong; transverse
sulcus strong, slightly widened to c.2/3 distance
to apices; apical 1/3 of anterior grooves, second
and third lateral posterior grooves, widened sec-
tion of sulcus bare, nitid; apices of both anterior
and posterior ridges near centre of pronotum with
close, appressed, yellowish setae.
Elytron. Broad, flattened; disc with sutural and
three lateral ridges, sutural, first and second
lateral ridges similar in development, not strongly
raised, third lateral ridge stronger joining lateral
margin of elytron c.2/3 distance to apex; lateral
stria joining margin of elytron c.1/2 distance to
apex; crests along length of all ridges each with
single row of adjoining, circular scales, less dis-
tinct on third lateral; rest of surface with circular,
appressed scales densest outside third lateral
ridge, fairly dense inside third lateral towards
second lateral ridge, rest of surface with scales
most numerous towards crests of ridges;
epipleuron wide to apex, subnitid with scales only
towards outer margin.
REVISION OF AUSTRALOXENELLA
Pygidium. Nitid with scattered, small setate
punctures.
Ventral surface and legs. Abdominal segments,
metasternum, mid and hind legs nitid, abdominal
segments, centre of metasternum and meso- and
meta-femora with scattered small, setate punc-
tures.
COMMENTS. Australoxenella moogoon sp.
nov. is a member of Group | species group and
does not have a pronotal prominence anterior to
the transverse sulcus and has the elytral ridges
low and capped with a single row of adjacent
circular scales. It is closest to A. wurrook sp. nov.
and the two species can be separated by charac-
ters listed in the key to species. Australoxenella
wurrook and A. moogoon were taken at the same
locality and series of flight interception traps but
not necessarily in the same trap.
Australoxenella wurrook sp. nov.
(Figs 3C, 3D, 6B)
ETYMOLOGY. An Aboriginal word for flat.
MATERIAL EXAMINED. HOLOTYPE ANIC116,
sex not determined, from 33km E of Jabiru, Arnhem-
land, NT, 15-23.xii.1993, S. & J. Peck, 93-130, 5 FITs,
Podocarp Canyon, rainforest. Paratype in HAHC, same
data as Holotype.
DESCRIPTION. Length 2.8—3.1mm, greatest
width 1.6—1.7mm. Dark reddish brown dorsally,
slightly lighter ventrally.
Head. Median groove reaching anterior edge,
lateral grooves falling short of anterior edge and
slightly converging on anterior 1/3; surface of
head with moderately long, narrow marking
above each eye; anterior edge of dorsal surface of
head with an indistinct fringe of very short flat-
tened setae.
Pronotum. Four central anterior rounded ridges
of equal height, median, first and second lateral
pairs of grooves of equal width and depth, third
lateral pair of grooves fine reaching anterior
pronotal edge c.1/3 of distance from anterior
angles to centre line; pronotal surface adjacent to
posterior median groove elevated, crest of
prominence acute, median groove not visible in
anterior 1/2, strong in posterior 1/2; transverse
sulcus long, narrow, visible for 4/5 distance to
lateral pronotal edges, surface of sulcus bare,
nitid; pronotal surface except for sulcus and small
area between anterior and posterior sculpturing,
covered in close, appressed, circular scales,
denser in five central anterior grooves, groups of
dense, yellowish setae at anterior ends of
373
posterior grooves; lateral pronotal edges between
anterior and posterior angles with indistinct
fringe of very short, flattened setae.
Elytron. Wide, flattened, length to width ratio
about 2:1; sutural and three lateral ridges visible,
not high, sutural ridge not raised near posterior
end, equivalent in height to first and second
lateral ridges, third lateral ridge weaker, shorter,
almost reaching lateral margin of elytron c.3/8
distance to apex then running parallel to margin
towards apex; lateral stria strong, short, reaching
lateral margin of elytron just before third lateral
ridge; entire surface of elytron densily covered
with close, appressed, circular scales, these in the
form of a single row of adjacent scales along crest
of each elytral ridge; epipleuron broad all the way
to apex, nitid, with scales only towards outer
margin.
Pygidium. Smaller, nitid with scattered setate
punctures.
Ventral surface and legs. Ventral surface nitid,
last abdominal segment and meso- and meta-
femora moderately punctate, punctures with
short, fine setae.
COMMENTS. Australoxenella wurrook sp. nov.
is closest to three new Queensland species, A.
zborowskii s, A. concinna, and A. teeta. It lacks
the elytral fringes of flattened setae found in the
three Queensland species though it does have
very short fringes on the anterior of the head and
lateral pronotal edges. The circular scales in the
U-shaped elytral depressions are not arranged in
evenly spaced transverse rows. The pronotal
clusters of spur-like setae on A. zborowskii and A.
concinna are absent in A. wurrook, and the elytra
of A. teeta are much less flattened than A. wur-
rook. The two specimens of A. wurrook were
taken in flight interception traps at Podocarp
Canyon near Jabiru, N.T., in rainforest. A.
moogoon sp. nov. was also taken at this site but
not necessarily in the same traps as A. wurrook.
Australoxenella midgee sp. nov.
(Figs 4A, 4B, 6A)
ETYMOLOGY. An Aboriginal word for small.
MATERIAL EXAMINED. HOLOTYPE ANIC117,
sex not determined, from 11? 51'S, 142? 38'E, 12km
SSE of Heathlands, Qld, 16.1.1992, T. A. Weir & I.D.
Naumann, Berlesate ANIC 1214 closed forest litter.
DESCRIPTION. Length 2.4mm, greatest width
1.3mm. Dark reddish brown dorsally and ventral-
ly.
Head. Median groove reaching anterior edge,
374
lateral grooves slightly shorter and slightly con-
verging for anterior 1/2; surface of head with
narrow, elongate, nitid marking above each eye.
Pronotum. Anterior rounded ridges adjacent to
median groove strongly elevated, the resulting
prominence as high as prominence posterior to
transverse sulcus, first lateral pair of anterior
grooves barely visible, third lateral pair of
anterior grooves very weak, meeting anterior
pronotal edge c.1/3 distance from anterior angles
to centre line; posterior grooves not strong, ridges
adjacent to median groove forming an elevated
prominence; transverse sulcus not widened at
apices; pronotal areas posterior to sulcus and
adjacent to median sculpturing depressed, con-
cave, the lateral margins of depressed areas form-
ing costae parallel to lateral pronotal edges.
Elytron. Disc with sutural and three lateral
ridges, sutural and third lateral ridges strongest,
sutural ridge slightly raised in posterior 1/3;
lateral stria not quite reaching lateral margin of
elytron c.1/2 distance to apex; dorsal surface out-
side of third lateral ridge with close, appressed,
circular scales, crests of other ridges with similar
scales; U-shaped depressions between ridges
with regular, transverse, rows of scales joining
the ridges, apparently in slight depressions, sur-
face bare, dull between; epipleuron narrowing
towards apex, surface dull, bare except with
scales along outer margin.
Pygidium. Dull with small dense setate punc-
tures covering entire surface.
Ventral surface and legs. Abdominal segments,
legs and centre part of metasternum dull with
numerous setate punctures, setae less scale-like
than those on dorsal surface; meso- and meta-
tibiae and femora less flattened than in other
species of the genus.
COMMENTS. Australoxenella midgee sp. nov.
and A. kalpara sp. nov. are the Group 2
Queensland species closest to A. humptydooensis
Howden and Storey. Australoxenella midgee can
easily be separated from the latter species by the
shape of the pronotum, the transverse rows of
scales on the elytral depressions, and narrower
tibiae. The only known specimen was taken in a
berlesate sample of litter from closed forest.
Australoxenella kalpara sp. nov.
(Figs 2D-F, 6A)
ETYMOLOGY. An Aboriginal word for bed of a
river.
MATERIAL EXAMINED, HOLOTYPE ANIC118,
sex not determined, from 15° 11'S, 143? 52' E (GPS),
MEMOIRS OF THE QUEENSLAND MUSEUM
Hann River, Qld, 18.xii.1993- 14.1.1994, P, Zborowski
& E.D. Edwards, flight intercept trap.
DESCRIPTION. Length 2.3mm, greatest width
].2mm. Dark reddish brown dorsally, slightly
lighter ventrally.
Head. Median groove almost reaching anterior
edge, lateral grooves slightly shorter and slightly
converging for anterior 1/2; surface of head with
small, narrow, elongate, dull marking above each
eye.
Pronotum. Anterior rounded ridges adjacent to
median groove strongly elevated, the resulting
prominence almost as high as prominence
posterior to transverse sulcus, median and first
lateral pair of grooves only slighty reduced, third
lateral pair of grooves very finely impressed,
reaching anterior pronotal edge c. 1/3 distance
from anterior angles to centre line; transverse
sulcus broad, joined by similar depression run-
ning from posterior pronotal edge just outside
pronotal sculpturing posterior to sulcus;
transverse sulcus and wide grooves from
posterior pronotal edge bare, dull, area between
two pronotal prominences with close, appressed,
yellowish setae.
Elytron. Disc with sutural and three lateral
ridges all strong, sutural ridge elevated slightly
on posterior 1/3, crest of third lateral ridge with
fine groove along inner edge resulting in a
doubled appearance; lateral stria strong, meeting
lateral margin of elytron c.1/2 distance to apex;
crests of sutural, first and second lateral ridges
each with single row of adjacent circular scales,
additional scattered scales along outer edges of
these ridges, close, appressed, circular scales
from the third lateral ridge to lateral margin of
elytron except along centre of third lateral ridge,
just inside the lateral stria and just inside basal 1/4
of margin; U-shaped depressions between ridges
with regular transverse rows of adjacent, circular
scales, rest of surface bare, dull; epipleuron
strongly narrowing towards apex, surface dull,
bare except scales along outer margin.
Pygidium. Subnitid with numerous punctures
separated by about one diameter, each with a
small elongate scale.
Ventral surface and legs. First four abdominal
segments and metasternum dull, last visible ab-
dominal segment, meso- and meta-femora and
tibiae subnitid; abdominal segments, femora and
centre portion of metasternum with scattered,
elongate, appressed setae or scales; carinate crest
of lanceolate posterior prosternal process not
widened near centre.
REVISION OF AUSTRALOXENELLA 375
0.25
E 4 P * 0.25
FIG.4. A,B, Australoxenella midgee sp. nov. A, head and pronotum, lateral view. B, elytra, dorsal view. C-F,
Australoxenella peckorum sp. nov. C, head and pronotum, lateral view. D, elytra, dorsal view. E, mesosternum,
ventral view. F, male genitalia, dorsal view.
COMMENTS. Australoxenella kalpara sp. nov. of the central crest of the prosternal lanceolate
is closest to A. midgee sp. nov., also from Cape process in A. kalpara. The habitat was riverine
York Peninsula. The two species canbe separated open woodland dominated by Melaleuca and
by pronotal sculpture, reduced punctation on the Leptospermum (P. Zborowski, pers. comm.).
pygidium in A. Kalpara and the blade-like shape
376
Australoxenella peckorum sp. nov.
(Figs 4C-F, 6B)
ETYMOLOGY. For the collectors of most of the new
specimens in this study, Stewart and Jarmila Peck of
Ottawa, Canada.
MATERIAL EXAMINED. HOLOTYPE ANIC119,
sex not determined, from Kakadu N.P., Kapalga Res.
Stn, NT, 11-25.xii.1993, S. & J. Peck, 93-118 FIT
eucalypt woodland. PARATYPES same data as
Holotype, (10); N.T.: Kakadu N.P., Kapalga Station
Gabarlgu, 25. xii.- 7.1.1994, S. & J. Peck, 93-137 FIT
rainforest, (2); Kakadu N.P., S. Alligator River, Gun-
garee rainforest trail, 12-25.xii.1993, S. & J. Peck,
93-120 FIT I, (3), same data except 93-121 FIT II, (5),
same data except 25.xii.-6.1.1994, S, & J. Peck, 93-141
FIT I, (1). Paratypes in HAHC, QPIM, NTMA, SBPC.
DESCRIPTION. Length 2.3-3.0mm, greatest
width 1.2-1.6mm. Reddish brown dorsally and
ventrally.
Head. Median groove reaching or almost
reaching anterior edge, lateral grooves reaching
2/3 distance to anterior edge, converging slightly
for apical 2/3; surface of head with small elongate
marking above each eye.
Pronotum. Anterior rounded ridges adjacent to
median groove raised posteriorly to form a
prominence almost as high as prominence
posterior to transverse sulcus, median and first
lateral pairs of grooves reduced, third lateral pair
of grooves fine, middle 1/3 often not visible
reaching anterior edge of pronotum c.1/3 distance
from anterior angles to centre line; posterior
prominence strong, apex rounded, median groove
weakly visible on posterior half only, first pair of
lateral grooves not very deep; transverse sulcus
strong, widened almost entire length.
Elytron. Disc with sutural and three lateral
ridges, sutural ridge only slightly raised
posteriorly, second lateral ridge not strong, third
lateral strong reaching almost to apex of elytron,
crests of sutural, first and second lateral ridges
rounded; lateral stria strong, reaching lateral mar-
gin of elytron almost 1/2 distance to apex; close,
appressed, circular scales on elytron largely con-
fined to area outside third lateral ridge and near
crests of other ridges, U-shaped depressions and
area near apex almost bare, dull; epipleuron nar-
rowed towards apex, dull, bare except near outer
margin which has close, appressed, circular
scales.
Pygidium. Dull with scattered, medium punc-
tures, setae indistinct.
Ventral surface and legs. Abdominal segments
dull with scattered, medium-sized punctures,
setae indistinct; metasternum dull, punctures and
MEMOIRS OF THE QUEENSLAND MUSEUM
setae of central portion indistinct; meso- and
meta-femora with scattered medium-sized setate
punctures; metasternum medially just posterior to
mesocoxae with broadened transverse ridge, the
posterior edge of which is almost straight to feeb-
ly, anteriorly, arcuate medially.
Male genitalia as in Fig. 4F.
COMMENTS. The shape of the posterior edge of
the transverse ridge of the metasternum which is
an almost straight line (instead of an anteriorly
directed, abrupt, obtuse angle) allows separation
of A. peckorum sp. nov. from the other members
of Group 2 Australoxenella species. The long
series was taken at several sites within Kakadu
National Park, N.T., using flight interception
traps set in eucalyptus woodland and rainforest.
Australoxenella mirreen sp. nov.
(Figs SA, 5B, 6B)
ETYMOLOGY. An Aboriginal word for south.
MATERIAL EXAMINED. HOLOTYPE ANIC120,
sex not determined, from Litchfield N.P., Pethricks
Rainforest, NT, 8-30.xii.1993, S. & J. Peck, 93-110
FIT deep rainforest. Paratypes same data as Holotype,
(1); N.T.: Litchfield N.P., Wangi Falls, 8- 30.xii. 1993,
S. & J. Peck, 93-113 FIT deep rainforest, (1). Paratypes
in HAHC.
DESCRIPTION. Length 2.4-2.6mm, greatest
width 1.3-1.4mm. Dark reddish brown dorsally
and ventrally.
Head. Median groove reaching 3/4 distance to
anterior edge, lateral grooves slightly shorter and
slightly converging for anterior 2/3; surface of
head with small oval marking above each eye.
Pronotum. Anterior rounded ridges adjacent to
median groove raised posteriorly to form a
prominence almost as high as prominence
posterior to transverse sulcus, median groove
slightly reduced towards apex of prominence,
first and second lateral pairs of grooves strong,
third lateral pair of grooves finer, reaching
anterior pronotal edge c.1/3 distance from
anterior angles to centre line; posterior
prominence strong, apex rounded, median groove
not visible for anterior 1/3, first lateral pair of
grooves slightly stronger, second lateral pair
stronger still; transverse sulcus strong, widened
for most of length.
Elytron. Disc with sutural and three lateral
ridges strong, sutural ridge strongly raised
posteriorly, three lateral ridges about same
height, third lateral ridge slightly longer joining
lateral margin of elytron c.4/5 distance to apex;
REVISION OF AUSTRALOXENELLA 377
FIG.5. A,B, Australoxenella mirreen sp. nov, A, head and pronotum, lateral view. B, elytron, lateral view. C-F,
Australoxenella humptydooensis Howden & Storey. C, head and pronotum, lateral view. D, dorsolateral view.
E, mesosternum, ventral view. F, male genitalia, ventral view. (Fig.5D from Howden & Storey,1992).
lateral stria strong, joining lateral margin of number in U-shaped depressions and especially
elytron c.2/3 distance to apex; surface outside near apex; epipleuron narrowed near apex, sur-
third lateral ridge and near crests of allridges with face dull bare, except near outer margin which
close, appressed, circular scales, these reduced in has close, appressed, circular scales.
378
Pygidium. Dull with indistinct scattered setate
punctures.
Ventral surface and legs. Abdominal segments
dull with indistinct scattered setate punctures;
metasternum dull, centre portion with indistinct
setate punctures; meso- and meta- femora dull
with scattered setate punctures.
COMMENTS. Australoxenella mirreen sp. nov.,
from Litchfield National Park, west of Adelaide
River, is similar to A. humptydooensis from the
Darwin area. The two species are separable with
difficulty by the characters mentioned in the key
to species.
Australoxenella humptydooensis Howden &
Storey, 1992
(Figs 5C-F, 6B)
Australoxenella humptydooensis Howden & Storey
1992:1813.
MATERIAL EXAMINED. HOLOTYPE
QMBAT12146, 6km E Humpty Doo, 6-19.x.1990, R.I.
Storey, at U.V. light. Other material (51): N.T.: Dar-
win, CSIRO McMillans Rd, 1-25.xii.1993, S. & J.
Peck, mixed eucalypt woodland 93-87 FIT, (40), same
data except 25.xii- 10.1.1994, mix. euc. woodlot 93-148
FIT, (3); 50km S Darwin, Berry Spg.Pk, 4-27.xii.1993,
S. & J. Peck, 93-99 rainforest malaise, (7), same data
except 27.xii.-3.1.1994, 93-154 rainforest FIT, (1).
Specimens in ANIC, HAHC, ISEA, QPIM, QMBA,
SBPC,
DESCRIPTION. Length 2.2-2.6mm, greatest
width 1.1—1.4mm. Dark reddish brown dorsally
and ventrally.
Head. Median groove reaching close to anterior
edge, lateral grooves slightly shorter and slightly
converging for anterior 1/2; surface of head with
small oval-elongate marking above each eye.
Pronotum. Median anterior groove slightly
reduced, especially posteriorly, apical 1/2 of third
lateral pair of grooves fine, joining anterior
pronotal edge c.1/3 distance from apical angles to
centre line, two median anterior, rounded ridges
raised posteriorly to form a prominence, not as
high as prominence posterior to transverse sul-
cus; median groove posterior to sulcus fine,
sometimes not visible in anterior 1/2, rounded
ridges either side of median groove expanded to
form a prominence, with crest rounded;
transverse sulcus deep, widened posteriorly for
c.3/4 length.
Elytron. Disc with sutural and three lateral
ridges, sutural ridge slightly raised in middle 1/3,
first lateral ridge strong, basal 1/2 highest, second
lateral ridge weaker, third lateral ridge weak,
MEMOIRS OF THE QUEENSLAND MUSEUM
joining lateral margin of elytron c.4/5 distance to
apex; lateral stria reaching margin of elytron c.1/2
distance to apex; close, appressed, circular scales
on surface of elytron outside of third lateral ridge,
just inside third lateral ridge except for a fine, bare
groove on basal 1/2, second lateral ridge rounded
with scales on outside, inside bare with fine
groove, crest of first lateral ridge rounded with
scales over most of length, a slight inner groove
on apical 1/3, sutural ridge rounded with scales
over entire length. U-shaped depressions and
apex of elytron dull with only scattered scales;
epipleuron wide, narrowing towards apex, dull
with scales along outer margin.
Pygidium. Subnitid with scattered small punc-
tures each with a short appressed, elongate scale.
Ventral surface and legs. Abdominal segments,
metasternum, and mid and hind legs subnitid,
femora, centre of metasternum and abdominal
segments with scattered fine punctures, each with
a short, appressed, flattened seta, more scale-like
on abdominal segments and apical margins of
metafemora.
Male genitalia as in Fig. 5F.
COMMENTS. Australoxenella humptydooensis
is closest to A. mirreen which was taken slightly
south of the former, in Litchfield National Park.
Australoxenella humptydooensis was taken in
numbers using both flight interception and
malaise traps, in open and closed forest habitats.
DISCUSSION The 11 species of Australoxenella
fall into two groups separated in the first couplet
of the key. Group 1 consists of A. bathurstensis,
A. zborowskii, A. concinna, A. teeta, À. wurrook,
and A. moogoon . All have the sculptured area of
the pronotum in front of the transverse sulcus of
a constant height with the centre five grooves of
equal depth and the four ridges of equal height;
all but A. teeta have the elytral epipleura wide to
the apex; the epipleura are subnitid to nitid in all
species; in A. zborowskii, A. concinna, A. wur-
rook and A. moogoon the elytra are relatively
broad, the ridges and U-shaped depressions not
strongly developed, and the crest of each ridge is
marked by a distinctive single row of adjacent
circular scales. Group 2, containing A. midgee, A.
kalpara, A. peckorum, A. mirreen and A.
humptydooensis, always has the middle two
anterior pronotal ridges developed into a
prominence similar to the posterior prominence
but lower; the elytral epipleura are always dis-
tinctly narrowed towards the apex and not strong-
ly nitid; the elytra are more convex in cross
REVISION OF AUSTRALOXENELLA
379
FIG.6. Distribution of Australoxenella spp. A, North Queensland: (@ ) A. concinna sp. nov.; (O ) A. zborowskii
sp. nov.; (solid semicircle) A. midgee sp. nov.; (*) A. kalpara sp. nov.; (Å) A. teeta sp. nov. B, Northern
Territory: (8 ) A. bathurstensis Howden and Storey; ((_)) A. wurrook sp. nov.; (solid semicircle ) A. moogoon
sp. nov.; C ).A. peckorum sp. nov.; (O ) A. mirreen sp. nov.; (Å) A. humptydooensis Howden and Storey.
section and the ridges and U-shaped depressions
more strongly developed; the crests of the elytral
ridges usually do not have a distinctive row of
adjacent circular scales. Both species groups
occur in N.T. and Qld.
The following are comments on characters
employed by Howden & Storey (1992): 17. Ab-
sence of striae. Australoxenella was rated
apomorphic (striae absent) in this character
despite all but one of the new species and the two
original species having an elytral feature which
was thought to be a 'stria'. The exact derivation
of this stria-like feature is unknown.
18. Less than five abdominal segments at mid-
line. The abdominal segments in Australoxenella
are greatly compacted and reduced at the mid-line
(except the large last visible segment), and dis-
cerning the exact number is difficult. The number
varies with sex, four in males and five in females.
Except for these features, all new species other-
wise agree with the character-states table for
Australoxenella of Howden & Storey (1992).
Bordat & Howden (1995) described three new
genera in the tribe from Borneo and discussed the
phylogeny of the Stereomerini, removing the
South American Termitaxis Krikken.
Specimens of Australoxenella have been taken
in closed (rainforest) and open forest situations
(c.1/3 in the former). Six species were taken only
in closed forest, two only in open forest, two in
both, with no information available for one
species. Both habitat types produced specimens
in N.T. and Qld. As most specimens were ob-
tained using flight interception traps, it would
seem that collection site data reflect habitat
preferences of the beetles. There was some
evidence in the long N.T. series that catches were
higher in traps where some odour of decay was
evident, the traps having been run for several
weeks.
Australoxenella is still rare in collections. Six
species are only represented by the holotype.
However, the long series of A. humptydooensis
and A. peckorum indicate that at least these spe-
cies can be common under some circumstances.
There is no information available on the biol-
ogy of Australoxenella. Howden & Storey (1992)
speculated that all Stereomerini were termito-
philes and the unusual morphology, specialised
pronotal setae in some species and rarity all point
to the genus being inquiline. The only direct
evidence of termite species being the hosts, was
that the only known South American specimen of
the tribe, the holotype of Termitaxis holmgreni
Krikken was taken in the nest of a termite (Krik-
ken, 1970). Bordat & Howden (1995) suggested
380
that Termitaxis should be removed from
Stereomerini until the holotype could be closely
examined. Though some work has been under-
taken investigating termite nests and their as-
sociated fauna in tropical areas of Australia, no
Australoxenella specimens have been found.
ACKNOWLEDGEMENTS
We are thankful for the loan of specimens to:
Dr Stewart Peck, Ottawa; Dr Geoff Monteith,
QMB; Mr Tom Weir, ANIC; and Dr Graham
Brown, NTMA. Anne Howden and John
Donaldson made useful comments on the
manuscript.
LITERATURE CITED
BORDAT, P. & HOWDEN, H.F. 1995. Trois nouveaux
genres, trois nouvelles espéces de Stereomerinae
MEMOIRS OF THE QUEENSLAND MUSEUM
de Bornéo (Coleoptera, Aphodiidae). Bulletin de
la Société Entomologique de France 100:11-20.
HOWDEN, H.F. & STOREY, R.I. 1992. Phylogeny of
the Rhyparini and the new tribe Stereomerini, with
descriptions of new genera and species (Coleop-
tera; Scarabaeidae; Aphodiinae). Canadian Jour-
nal of Zoology 70: 1810-1823.
KRIKKEN, J. 1970. Termitaxis holmgreni gen. nov.,
sp. nov., a blind flightless termitophilous scarab
from Peru (Coleoptera: Aphodiidae). Proceedings
of the Kongliga Nederlandse Akademie
Wetenschappen Series C 73: 469-476.
STEBNICKA, Z.T. & HOWDEN, H.F. 1994. A
revision of the Australian genus Podotenus A.
Schmidt (Coleoptera: Scarabaeoidea: Aphodiini).
Invertebrate Taxonomy 8:17-62.
1995. Revision of Australian genera in the tribes
Aphodiini, Aegialiini and Proctophanini
(Coleoptera: Scarabaeidae: Aphodiina). Inver-
tebrate Taxonomy 9: 709-766.
A NEW SPECIES OF SCHIZOTREMA (CUMACEA: NANNASTACIDAE) FROM
MORETON BAY, QUEENSLAND
D.J. TAFE AND J.G. GREENWOOD
Tafe, D.J. & Greenwood, J.G. 1996:07:20: A new species of Schizotrema (Cumacea:
Nannastacidae) from Moreton Bay, Queensland. Memoirs of the Queensland Museum 39(2):
381-389. Brisbane. ISSN 0079-8835.
Both sexes of Schizotrema nudum sp. nov. are described and figured. It differs from other
species of the genus in lacking spines on the carapace and on the dorsal surface of the
abdomen. The species also differs from the closely related S. leopardinum Hale, in the relative
lengths of exopod and endopod of the uropod. Eleven species are now known in this genus.
Five (S. aculeatum Hale, S. depressum Calman, S. leopardinum Hale, S. nudum sp. nov. and
S. resimum Hale) occur in Australian waters. [ ] Crustacea, Cumacea, Nannastacidae,
Australia, Schizotrema.
D.J. Tafe, Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101,
Australia; J.G. Greenwood, Department of Zoology, University of Queensland, St Lucia,
Queensland 4072, Australia; received 20 February 1996.
During a study of cumaceans in Moreton Bay,
on the central east coast of Australia, a new
species of Schizotrema Calman, 1911 was cap-
tured. The genus is one of 18 in the Nannas-
tacidae. Schizotrema is differentiated from the
closely related Schizocuma by the pseudorostrum
which forms widely spaced inhalent siphons.
All specimens were taken using a sledge-
mounted net of 500 jum mesh towed within 0.4m
ofthe substratum at night. Although 700 samples
were collected from 40 sites widely distributed
within the Bay region only three specimens of the
new species were taken, and all came from the
same location. One specimen of Schizotrema
aculeatum Hale, 1945, was also taken from the
same location. A male and female of the new
species were dissected, drawn and described and
the other male used for S.E.M. study. Descrip-
tions were carried out with reference to Felgen-
hauer (1992), Hale (1936, 1945, 1949), Jones
(1963) and Watling (1989). In descriptions of
appendages lateral refers to that margin of the
appendage facing the lateral body surface and
medial refers to that margin directed toward the
midline of the body. Long plumose natatory setae
on thoracic appendages have been truncated in
some figures
SYSTEMATICS
Class CRUSTACEA
Subclass MALACOSTRACA
Superorder PERACARIDA
Order CUMACEA
Family NANNASTACIDAE
Schizotrema Calman, 1911
Schizotrema-‘Calman, 1911:360; Stebbing, 1913:165.
DIAGNOSIS. Anterolateral angle of carapace
well developed; eyes divided into two separate
groups; pseudorostrum forming two widely
separated inhalent siphons; second to fifth
pereionites well developed, usually with plate-
like lateral extensions; no pleopods and no free
telson.
Eleven species are now known in this genus.
Five (S. aculeatum Hale, S. depressum Calman,
S. leopardinum Hale, S. nudum sp. nov. and S.
resimum Hale) occur in Australian waters and
only S. aculeatum Hale and S. nudum sp. nov. are
known from Moreton Bay.
KEY TO SPECIES OF SCHIZOTREMA
(Adapted from Hale 1949)
1. Carapace broad and depressed. Peduncle of
uropod longer than telsonic somite
a i er, Tere eee: depressum Calman
Carapace not or little depressed, Peduncle of
uropod much shorter than telsonic somite .. 2
2. Exopod of uropod at least 0.3 times as long as en-
dopod (not including terminal spines) .... 3
Exopod of uropod at most 0.25 times as long as
edopod. « ov tae. sews cues Ei eS
3. Last pedigerous and first pleon somite each with
one pair of outstanding dorsal spines . . . . . E
Such somites with clumps of small spines or no
Spies 7 ^1 2140 asl A hye ee PORE edt 5
4. Spines on carapace aculeate . . aculeatum Hale
382
Spines on carapace spatulate sakaii Gamo
5. Anterolateral angle of carapace produced as a
stout cylindrical process... ..... 6
Anterolateral angle of carapace not produced as a
stout cylindrical process
6. Telsonic somite wider than long in dorsal view
Parai i tats fate rigs bifrons Calman
Telsonic somite longer than wide in dorsal view
atlanticum Bacescu & Muradian
7. Uropod with spine of exopod not quite reaching
distal end of endopod (excluding terminal spine)
ee rere gu vts ua macrodactylus Fage
Uropod with spine of exopod clearly reaching
beyond distal end of endopod (excluding ter-
minal spine)
8. Surface of carapace with reticulate pattern of in-
dentations; anterolateral corner of carapace an-
gular and without spine . nudum sp. nov,
Surface of carapace for the most part smooth;
anterolateral angle of carapace produced as a
slender spine ......... sordidum Calman
9. Carapace with dorsal and lateral spines
resimum Hale
HOM rat d abs 10
10. Carapace with mottled colour pattern, and with
anterolateral angle broad, not at all down-bent
Bab kPa tie Hoe Roe d leopardinum Hale
Carapace without mottled colour pattern, and
with anterolateral angle produced and strongly
down-bent bidens Fage
Schizotrema nudum sp. nov.
(Figs 1-4)
MATERIAL EXAMINED. HOLOTYPE
QMW20468, adult (S.L., 1.6 mm), perm. slide mount,
Horseshoe Bay, Peel Island, 27°30'S, 153°22’E, site
31, D. Tafe, 17.6.1990, 2-3 m, sand, 34 p.p.t. salinity,
19°C water temperature. PARATYPES QMW20469,
ovig. (S.L., 1.4mm), same data as holotype, perm. slide
mount; QMW20470, adult (S.L., 1,7 mm), S.E.M.
mount, site 34, otherwise same data as holotype.
DESCRIPTION OF MALE. Standard length 1.6
mm, (measured from anterior end of pseudo-
rostrum to posterior end of telsonic somite, ex-
cluding spines and setae). Cuticle well calcified.
Carapace with reticulate pattern of indentations;
last three pedigerous and first three abdominal
somites with paired dorsal nodules but without
obvious spines (Fig. 1A, B). Carapace twice as
long as combined length of exposed pedigerous
somites, almost 1/3 total length of animal;
posterior margin of carapace slightly raised dor-
sally, produced posteriorly to cover whole of first
pedigerous somite and dorsum of second;
anterolateral corner of carapace angular and
MEMOIRS OF THE QUEENSLAND MUSEUM
devoid of spines. Two prominent ocular regions
located anteriorly on dorsolateral margins, each
composed of 3 separate, hemispherical lenses
(Fig. 1C, D).
Pleural parts of second to fifth pedigerous
somites expanded (Fig. 1D); pleural region of
second pedigerous somite almost as wide as
carapace, 3-5 gradually decreasing in width; ab-
dominal somites not expanded laterally. Dorsal
region of telsonic somite flattened, approximate-
ly as wide as long, and longer than peduncles of
uropods (Fig. 1E).
First antenna (Fig. 2A) with first segment of
peduncle longer than second and third segments
together, three large setae distolaterally, row of
short, simple setae laterally; second segment with
two large setae distolaterally, tubercle with one
plumose and two simple setae distomedially;
third segment subequal in length to second, bear-
ing ] simple and 2 plumose setae distomedially,
and accessory flagellum distolaterally; accessory
flagellum bearing 4-5 simple setae; main flagel-
lum 3-segmented, first and second segments
slender and subequal in length, third smaller, with
2 long, annulated aesthetascs terminally and 2-3
fine setae subterminally.
Second antenna (Fig. 2B) with 3 short proximal
segments; fourth segment longer than combined
1-3. Third and fourth segments with lateral mar-
gins setose. Flagellum elongate, multiarticulate.
Mandible (Fig. 2C) molar process cylindrical
with flattened masticating surface. Incisor with
3-4 teeth on inner edge; lacinia mobilis present
on left mandible only; row of 5-6 closely spaced
stiff setae on inner margin of mandible between
distal teeth and molar process.
First maxilla (Fig. 2D) with 2 well developed
terminal endites carrying at their distal ends a
number of hamate setae; larger lobe with 7-9
setae, smaller 3-4 setae, with longest proximal,
slender and pappose. Appendage with backward-
ly directed endopodite posteriorly; palp unilobed,
with 2 long, slender setae, each with fine setules
almost invisible at x400.
First maxilliped (Fig. 2E) basis with a row of
slender simple setae medially and a hamate seta
distomedially. Endopod 4-segmented; first seg-
ment short, with 3 setae distomedially; second
segment rectangular with scattered setae; third
segment shorter and narrower, with 3 strong dis-
tal setae one of which is long and serrulate, lateral
margin of segment convex, with simple setae;
fourth segment circular, with 2 fine subterminal
setae.
Second maxilliped (Fig. 2F) basis subequal in
NEW CUMACEAN FROM MORETON BAY
Ww
oo
UJ
FIG. 1. Schizotrema nudum sp. nov., adult male. A, lateral view. B, dorsolateral view. C, carapace, lateral view.
D, carapace, anterolateral view. E, telsonic somite and uropods, dorsal view. SEMs with JEOL 6400.Scale bars
= 100um
384 MEMOIRS OF THE QUEENSLAND MUSEUM
f O-1mm ] 0-5 mm r O-1mm 1
ACDEF B GHI
FIG. 2. Schizotrema nudum sp. nov., adult male. A, first antenna. B, second antenna. C, right mandible and portion
of left mandible. D, first maxilla. E, first maxilliped. F, second maxilliped. G, third maxilliped. H, first pereiopod.
I, second pereiopod.
NEW CUMACEAN FROM MORETON BAY
length to merus, long, plumose seta distomedial-
ly, setae on medial margin and ridge on lateral
margin; ischium extremely short, without setae;
merus geniculate, expanded centrolaterally,
prominence bearing several long setae dis-
tolaterally, single large plumose seta arising from
the central anteromedial surface; carpus slightly
shorter than merus, longer than propodus, with
row of plumose setae along medial margin, scat-
tered simple setae on lateral surface; propodus
almost as wide as long, with 3 stout serrulate setae
distally, with 3-4 slender setae subdistally; dac-
tylus short with strong terminal seta.
Third maxilliped (Fig. 2G) basis inflated, sub-
equal in length to rest of limb; laterodistal lobe
with 2 stout and 1 fine plumose setae, two
plumose setae on distomedial region of basis;
ischium of endopod reduced, with simple seta
distolaterally; merus broad with a stout, plumose
seta distolaterally, row of short setae on convex
lateral margin; carpus subequal in length to merus
but narrower, long plumose seta laterally, scat-
tered short setae distomedially; propodus longer
than merus, carpus, and dactylus, simple seta
distolaterally, 3 plumose setae distally; dactylus
slender with 3 stout, curved terminal setae and 1
slender curved subterminal seta, 1 medial and 2
lateral simple setae. Exopod with basal segment
bearing 12 ridges with microspines, following
segment naked and elongate, 4 short distal seg-
ments each bearing 1 pair of long plumose setae.
First pereiopod (Fig. 2H) with inflated basis
and 5 endopod segments. Endopod longer than
basis, terminal segments progressively longer
from ischium to propodus, dactylus shorter than
propodus and subequal in length to terminal seta;
setae on all segments (except terminal seta) short
and simple; propodus and dactylus with rows of
fine setae on medial margins; terminal seta of
dactylus slightly curved with setules on distal
medial margin, 2 smaller terminal setae and 2
subterminal setae. Exopod with 2 basal segments
of similar length, first broad and fringed laterally
bearing 12 ridges with microspines; second
naked and elongate; segments 3-7 reduced in
length, each bearing a pair of long, plumose setae.
Second pereiopod (Fig. 21) basis inflated, as
long as rest of limb. Endopodal ischium reduced;
carpus subequal in length to dactylus, twice as
long as merus and propodus; terminal dactylar
seta straight and longer than segment, 2 shorter
terminal setae, | lateral and 2 medial setae; sur-
face of dactylus and propodus covered with
microspines. Exopod with 2 subequal proximal
segments, first bearing 12 ridges with
385
microspines laterally; segments 3-7 very
reduced, each bearing a pair of long, plumose
setae.
Third pereiopod (Fig. 3A) endopod segments
slender, together at least 1.5 times as long as
inflated basis. Endopodal carpus longer than
other segments, almost twice combined length of
ischium and merus; dactylus arcuate, bearing 2
setae. Exopod with 2 subequal proximal seg-
ments, first bearing 10 ridges with microspines
laterally; segments 3-7 very reduced, each bear-
ing a pair of long, plumose setae.
Fourth pereiopod (Fig. 3B) similar to third ex-
cept terminal 5 segments of endopod together
twice as long as inflated basis; exopod with 2
subequal proximal segments, first naked and
second bearing 7 pairs of minute ridges with
microspines; 4 reduced terminal segments each
bearing a pair of long, plumose setae.
Fifth pereiopod (Fig. 3C), with slender basis
and terminal segments, without exopod. Carpus
longer than other segments, 1.5 times length of
propodus and twice combined length of ischium
and merus; dactylus arcuate as with third and
fourth pereiopods.
Telsonic somite (Figs 1E, 3D) as wide as long,
with 6 scattered setae on each side of the dorsum
but without spines. Dorsal profile as figured.
Peduncle of uropod (Figs 1E, 3D) with lateral
margin 0.6 times length of telsonic somite, 0.4
times length of endopod exclusive of its terminal
spine. Exopod at least as long as peduncle and
0.45 length of endopod, with its long terminal seta
reaching nearly to midlength of terminal endopod
seta; | short distolateral terminal exopod seta.
Terminal seta of endopod about half as long as
ramus, with subterminal lateral setule; 2-3 short
distomedial endopod setae. Medial margin of en-
dopod serrate, with microspines.
DESCRIPTION OF FEMALE. Standard Length:
1.4 mm. Cuticle well calcified. Carapace and
abdominal somites with reticulate pattern of in-
dentations and paired dorsal nodules as in male.
First antenna (Fig. 3E) with first segment of
peduncle longer than combined length of seg-
ments 243, 3 long setae distomedially, row of fine
setae medially; second segment with 2 long setae
distomedially, tubercle with 3 simple setae dis-
tolaterally; third segment subequal in length to
second, with accessory flagellum bearing 3-4
setae; main flagellum 3-segmented with 2 annu-
lated aesthetascs and 1 long seta mounted ter-
minally; combined length longer than 3 segments
of peduncle.
386
Mandible (Fig. 3F) molar process cylindrical
with flattened masticating surface. Distal end
with 4 teeth on inner edge; lacinia mobilis on left
mandible only; row of 5 closely packed spines on
inner margin of mandible between distal teeth
and molar process.
First maxilla (Fig. 4A) with 2 well-developed
terminal endites bearing several hamate setae;
larger lobe with 8 distal setae and a subterminal
seta laterally; smaller lobe with 4 distal setae,
longest one proximal, slender and plumose.
Posteriorly appendage with a backwardly
directed endopodite; unilobed palp with 2 slender
distal setae.
Second maxilla (Fig. 4B) with 3 lobes, all with
setae along distal margins; largest, most distal
lobe with 13-14 setae, longest one plumose;
smaller inner lobe with 11-12 simple setae, outer
lobe with 6-7 delicate, inwardly directed setae.
First maxilliped (Fig. 4C) basis with row of
simple setae laterally and stout setae medially;
endopod 4-segmented, first segment short, with
one long plumose seta and 2 stout setae medially;
second segment rectangular, with hamate setae
medially and slender setae laterally; third seg-
ment narrow, with 3 strong distal setae, one long
and plumose; lateral margin of third segment
convex, with simple setae; fourth segment cir-
cular, with 2 fine subterminal setae.
Second maxilliped (Fig. 4D) basis longer than
merus and ischium combined, long plumose seta
distomedially; row of short setae medially and
short simple seta distolaterally; ischium rudimen-
tary and naked; merus geniculate, lateral margin
convex, with several fine setae, single large
plumose seta arising from the central
anteromedial surface; carpus longer than merus
and propodus, with 4 large plumose setae and
numerous simple setae medially, row of simple
setae midlaterally; propodus almost as wide as
long, 3 strong, serrulate and 5-6 slender setae
distally, directed medially, medial margin with
rounded setuled lobe, 3 midlateral setae; dactylus
slender, shorter than propodus, directed medially
with strong terminal spine. Exopod with 5 long,
plumose setae.
Third maxilliped (Fig. 4E) basis as long as next
3 segments combined, tubercle with 2 long
plumose setae distolaterally, 2 plumose setae dis-
tomedially; ischium short with plumose seta dis-
tomedially; merus longer than ischium but
shorter than carpus, plumose seta distolaterally
and small simple seta distomedially; carpus
shorter than propodus, plumose seta distolateral-
ly, 3-4 small setae distomedially; propodus nar-
MEMOIRS OF THE QUEENSLAND MUSEUM
row proximally, lateral margin convex, three
slender plumose setae distomedially, row of short
setae on medial margin, 3-4 short setae on distal
margin; dactylus slender, twice as long as wide,
with 2 large curved terminal setae and 2-3 smaller
subterminal setae. Exopod with 2 proximal seg-
ments subequal in length; 2 reduced terminal
segments each bearing a pair of long plumose
setae.
First pereiopod (Fig. 4F) basis inflated; 5
slender endopod segments; ischium subequal in
length to merus with 1 medial and 2 lateral setae;
merus with 3 setae on convex lateral margin and
2 on medial margin; carpus twice length of merus
with 2 lateral and 3 medial setae; propodus sube-
qual in length to carpus and at least 1.5 times as
long as dactylus, 4 setae laterally and 2 dis-
tomedially, row of fine setae along medial mar-
gin; dactylus slender, with curved terminal seta
as long as dactylus, 2 shorter slender terminal
setae and 2 subterminal setae. Exopod with 2
proximal segments subequal in length; 3 reduced
terminal segments each bearing a pair of long
plumose setae.
Second pereiopod (Fig. 4G) basis inflated,
longer than segments 1-4 of endopod, scattered
simple setae only; ischium reduced and naked;
merus longer than ischium and propodus but
shorter than carpus and dactylus, 3 lateral and 2
medial setae; carpus subequal in length to dac-
tylus, twice as long as propodus, 1 long and 2
short distolateral setae, row of fine lateral setae;
propodus short with row of fine lateral setae;
dactylus twice as long as propodus with major
terminal seta straight and longer than segment, 2
shorter terminal setae, surface of dactylus
covered with microspines. Exopod with 2
proximal segments subequal in length, 2 reduced
terminal segments each bearing a pair of long,
plumose setae.
Pereiopods 3-5 (Fig. 4H-J) with slender basis
and 5 slender endopod segments; dactylus ar-
cuate with seta on inner margin; carpus longer
than propodus and nearly twice as long as merus
and ischium together; no exopod.
Telsonic somite (Fig. 4K) as wide as long, with
9-10 short setae on dorsal surface, with 2 slender
setae distomedially.
Peduncle of uropod (Fig. 4K) with outer margin
more than half as long as telsonic somite (0.65 X)
and half as long as endopod exclusive of its
terminal seta, | slender distomedial seta; exopod
shorter than peduncle and about 1/3 as long as
endopod, with its stout terminal seta reaching
beyond the distal end of endopod segment, 2
NEW CUMACEAN FROM MORETON BAY 387
SSE
Ze
2L
SESE
la
D
ZS
SS
See
=
27
LZ
À
N
À
|
i
oe
FIG. 3. Schizotrema nudum sp. nov. A-D, adult male. A, third pereiopod. B, fourth pereiopod. C, fifth pereiopod.
D, telsonic somite and uropods. E,F, ovigerous female. E, first antenna. F, right and left mandibles.
388 MEMOIRS OF THE QUEENSLAND MUSEUM
J
pr.5
I 0:1mm l 9:Tmm !
AJK BCDEFGHI
FIG. 4. Schizotrema nudum sp. nov., ovigerous female. A, first maxilla. B, second maxilla. C, first maxilliped.
D, second maxilliped. E, third maxilliped. F, first pereiopod. G, second pereiopod. H, third pereiopod. I, fourth
pereiopod. J, fifth pereiopod. K, telsonic somite and uropods.
NEW CUMACEAN FROM MORETON BAY
small subterminal setae; endopod with 2 stout
terminal setae, the outer one 2/3 as long as the
ramus and 3 times as long as the inner one,
bearing a long subterminal setule laterally and
row of microspines medially, 1 fine terminal seta,
2 dorsolateral and 2 dorsomedial setae, and 3
distomedial ridges bearing microspines.
DISCUSSION. Schizotrema nudum sp. nov. is
distinguished within the genus by its lack of
spines on the general surface of carapace and
abdomen, and by the paired nodules on the dorsal
surfaces of most free somites of the pereion and
pleon. The uropods also distinguish S. nudum
from other Australian species. Schizotrema
nudum id distinguished from S. aculeatum, S.
bifrons and S. sakai which have on spination
alone. The latter three species have characteristic
spines on the carapace and free somites.
Specimens of S. aculeatum were also taken at
night in Horseshoe Bay.
Like S. nudum, S. leopardinum is completely
devoid of stout spines on the cuticle. However,
the relative proportions of the rami of the uropod
enable the two species to be easily distinguished.
The female of S. leopardinum has not been
described but the male uropod has an exopod only
0.25x the length of the endopod, compared to
0.45x in S. nudum. Also the differences in colour
and surface texture make the two species readily
distinguishable. The bold pigment patches of S.
leopardinum persist even in spirit preserved
material (Hale, 1949).
Schizotrema resimum is distinguished from S.
nudum by its smaller size, conspicuous body
spination and very small exopod on the uropod.
The dorsum of its carapace exhibits rather large
spines, many of which bear a brush of minute
setae distally.
Schizotrema depressum exhibits a broad,
depressed carapace which distinguishes it from S.
nudum and all other members of the genus (Hale,
1949).
The new species was compared with type
material of S. aculeatum and S. leopardinum, and
non-type material of S. depressum, all in the
South Australian Museum.
389
ETYMOLOGY. Latin nudum naked; referring to
the lack of conspicuous body spines and striking
colour patterns.
ACKNOWLEDGEMENTS
We gratefully acknowledge B. Koh and R.
Crudgington for assistance in the field, J. Short
and P. Mather for constructive comments and P.
Davie, L. Watling and G. Poore for reviewing the
manuscript. Sampling and analytical equipment
was supplied by the Zoology Department,
University of Queensland, and funding of
fieldwork and electron microscopy was supplied
by Reckitt & Colman Pty Ltd.
LITERATURE CITED
CALMAN, W. T. 1911. On new or rare Crustacea of
the Order Cumacea from the collection of the
Copenhagen Museum. Part 2. The Families Nan-
nastacidae and Diastylidae. Transactions of the
Zoological Society of London 18: 341-398.
FELGENHAUER, B. E. 1992. External anatomy and
integumentary structures. Pp. 7-43. In Harrison,
F.W. & Humes, A.G. (eds). Microscopic
Anatomy of Invertebrates. Vol. 10: Decapoda
Crustacea. (Wiley-Liss: New York).
HALE, H.M. 1936. Cumacea from a South Australian
reef. Records of the South Australian Museum 5:
404-438.
1945. Australian Cumacea. No.9. The family Nan-
nastacidae. Records of the South Australian
Museum 8: 145-218.
1949, Australian Cumacea. No. 16. The family Nan-
nastacidae. Records of the South Australian
Museum 9; 225-246.
JONES, N. S. 1963. The marine fauna of New Zealand:
crustaceans of the order Cumacea. New Zealand
Oceanographic Institute Memoir 23: 1-82.
STEBBING, T.R.R. 1913. Cumacea (Sympoda). Das
Tierreich, 39 (R. Friedlander & Son: Berlin).
210p.
WATLING, L. 1989. A classification system for crus-
tacean setae based on the homology concept. Pp.
15-26. In Felgenhauer, B.E., Watling, L. &
Thistle, A.B. (eds). Crustacean Issues 6: Function-
al morphology of feeding and grooming in Crus-
tacea. (A.A.Balkema: Rotterdam).
REPTILE DIVERSITY IN A CALLITRIS FOREST IN
CENTRAL QUEENSLAND'S BRIGALOW BELT.
Memoirs of the Queensland Museum 39(2) 390, 1996:- The
clearing of much of Queensland's Brigalow Biogeographic
Region for agriculture and grazing has been well documented
(Sattler & Webster, 1984; Gasteen, 1985), but the effects of
this.on animal diversity and populations is unknown. To date
no comprehensive review of distribution and status of the
Brigalow Biogeographic Region's reptile species has been
completed, although such a study is current (Covacevich,
Couper & McDonald pers.comm.). To further that work T
monitored, a long term pil trap lo assess reptile species diver-
sity in a small, relatively untouched forest near Emerald,
MEQ.
The study site (2340 S, 148°06'E) is 100m trom the full
supply level of Fairbairn Dam. It was maintained by staff of
the Camp Fairbairn Outdoor Education Centre of the
Queensland Education. Department, with assistance from
visiting school children, The trap design followed Hobbs et.al,
(1994). Twelve buckets were set in à N-S line, over 50m. The
study site was located in sand amongst Callitris columellaris,
with scattered pockets of Cassia ovata.
The study was conducted over 199 days, from Aprl all
December 1995; specimens were collected on 72 days over 4
seasons, 170 specumens of 24 species were collected. 23
voucher specimens have been lodged in the Queensland
Museum reference collection (QMJ61410-61412, 61423,
61426, 61436-61441, 61444-61454, 61459). Most reptile ac-
livity occurred during October, November and May. In these
months 77.5% of the specimens were trapped (Fig 1).
The following species were collected during the study:
Diplodactylus conspicillatus x7, D. stelndachaeri x17, D.
taenicauda x7, D, viliatus x11, Heterunotia binoet x12,
Neplirueus asper 83, Lialis burronis x3, Carlia munda &Y, C,
pectoralis x7, Cryploblepharay carnabyi x27, Ctenotus
robuxtus x6, C. strauchii x13, C.taeniolatus x13, Lerista
fragilis x3, L punctatovittata x9, Menetia greyli x2, Morethia
boulengeri x13, M. taentopleura x3, Varanus teistis x1, Ram
phoryphlops sp. x4, Demansia pscimmoplris x1, Rhiroplo-
cephalus Tostai xl, Simoselaps australis x2, Vermicella
annulata x1. in addition to reptiles the following were col-
lected (and released) from the pit irap: spiders; scorpions,
centipedes; millipedes; bush cockroaches, frogs (Lim.
nodynastes ormilus), ianmnals (Tachyelossus aculeatus,
Pseudomys delicatulus).
Construction of the Fairbulm
Dam commenced in 1968 and was
completed in Decemher 1972. 11
flooded c.16000ha of native
forests, most of which had heen
grazed or modified. Native forests
boundyng iis waters are now vii-
tually the only non-agricultural
land in the Emerald district, That
24 species of small reptiles were
found in this small patch of Cal-
liris Forest in a 9 month period
suggests thal the diversity of small
reptiles has been maintained,
despite considerable modification
of habitat, It also highlights the
value of pit-trapping in surveys. I
have been based at Fairburn Dom
fur 5 years, and spend some time
everyday in the field. Prior to this
study, only 7 reptile species had D
heen recorded incidentally in the
Outdoor Education Centre's lease,
an aren of 19 ha, No methodical
Dscem ber
Novemaber
October
Septem ber
August
July
June
May
April -
Fig.l.
MEMOIRS OF THE QUEENSLAND MUSEUM
hand collecting or searching for reptiles had been undertaken.
No significant range extensions for reptiles were made,
However, the study confirms D. raenicauda, C. pectoralis, C.
taeniolatus and M. taeniopleura near the limit of their range
in the Emerald area (Covacevich & Couper, 1991). Two
species on this site are of special conservation concern cur-
rently. D. (aenicauda and V. annulata are classified *rare or
insufficiently known’ by Cogger et. al. (1993).
The Australian Nature Conservation Agency funded field
work by Jeanette Covacevich, Patrick Couper and Keith Me-
Donald in the Brigalow Biogeographic Region. They en-
couraged me to monitor the trap and report on this study. My
colleagues Bruce Davis and Jillian Ryan from Camp Fairbaim
Outdoor Education Centre and the many staff and students
that visited the centre during the study made the daily task of
checking the traps a pleasurable, educational experience,
Jeanette Covacevich assisted me in preparing this note.
Literature Cited
Cogger, H.G, Cameron, E. Sadlier, R. & Eggler, P. (eds) 1993,
The action. plan tor Australian reptiles. (Australian
Nature Conservation Agency; Canberra) 254 pp,
Covacevich, J.A & Couper, P.J., 1991. Reptiles. In Ingram,
0J. & Raven, RJ, (eds). An atlas of Queensland's
frogs, reptiles, birds and mammals. (Queensland
Museum: Brisbane).
Gasteen, W.J. 1985. The Bngalow lands of eastern Australia
- agricultural impact and land use potential versus
biological representation and stability, Pp. 45-49. In
Gasteen, J., Henry. D. & Page, S. (eds). Agriculture and
conservation in inland Queensland. (Wildlife Preserva-
tion Society of Queensland: Brisbane). 114p,
Hobbs, T.J, Monon, S. R., Masters, P. & Jones, K.R, 1994,
Influence of pit-trap design on sampling of reptiles in
arid spinifex grasslands. Wildlife Research 2): 438-90,
Saler, P.S. 1986. Nature conservation in Queensland: plan-
ning the matrix, Proceedings of the Royal Society of
Queensland 97: 1-21.
Sattler, P.S. & Webster, R.J. 1984 The conservation status of
Brigalow (Acacia harpophylla) communities in
Queensland. Pp. 149-160. In Bailey, A. (ed.) The
Brigalow Belt of Australia. (The Royal Society of
Queensland; Bnsbane),
Russell Deer, Camp Fairbairn Outdoor Education Centre,
Box 419, Emerald, Queensland 4720, Australia; 20 April
1996.
No. of Reptiles trapped versus Month
Number of specimens trapped per month.
THE BODOTRIIDAE (CRUSTACEA: CUMACEA) OF MORETON BAY,
QUEENSLAND
DJ. TAFE AND J.G. GREENWOOD
Tafe, D.J. & Greenwood, J.G. 1996:07:20. The Bodotriidae (Crustacea: Cumacea) of
Moreton Bay, Queensland. Memoirs of the Queensland Museum 39(2): 391-482. Brisbane.
ISSN 8835-0079.
Of 29 species of the Bodotriidae (Cumacea) collected from 40 sites in Moreton Bay
(1989-1993) 22 are new records for the region and 18 are new species. Of the new species
5 are left in open nomenclature because insufficient material ts available on which to base à
description, The 9 of Glyphocuma halei, previously unknown, is also described, Keys are
given to the subfamilies, genera and species of Bodotriidae from Moreton Bay. The keys to
species of Cyclaspis, Glyphocuma, and Leptocuma, sre broadened 10 include all Australian
species, Where a high degree of sexual dimorphism exists within species, such as those in
the genus Glyphocumua, separate keys to the sexes are constructed. L ] Crustacea. Moreton
Bay, Australia, Peracarida, Zooplankton.
D.J. Tafe, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia;
JG. Greenwood, Department of Zoology, University of Queensland, St Lucia, Queensland
4072, Australia; received 20 January 1996,
This is the first major taxonomic investigation
of Cumacea in Queensland waters, Isolated or
small collections have been studied in works b
Hale (1944a,b, 1948, 19493), Stephenson et al.
(1978) and Stephenson (1980a,b). The most sig-
nificant taxonomic works on Australian Cumacea
were published between 1928 and 1953 by H.M.
Hale. In 1953 he listed 160 species from the W,
S and E coasts of Australia.
The Bodotriidae is more diverse than all other
families (94 species) in Australian waters (Baces-
cu, 1988, 1990, 1992b). Cyclaspis is by far the
most species rich genus and its genetic centre is
in the Australasian region (Bacescu, 1992b). The
Nannastacidae (45 species) and Gynodiastylidae
(36 species) are also well represented around the
Australian coastline, while the Diastylidae (15
species), Leuconidae (2 species) and Lam-
propidae (2 species) are poorly represented,
Highest diversity in all families occurs on the
Pacific Coast, particularly near rivers, while 1he
S coast has the fewest species (Hale, 19532).
Five families of Cumacea are represented in the
species which have previously been collected
from Moreton Bay. They are the Bodotriidae,
Nannastacidae, Gynodiastylidac, Diastylidae and
Leuconidae. The Bodotriidae 1s most diverse, as
in other areas around Australia (Hale, 19532). Its
dominant genus is Cyclaspis. Sixteen bodotriid
species were recorded for Queensland waters by
Hale between 1928 and 1951]. Greenwood &
Johnston (1967) added one and Stephenson et al.
(1978), Stephenson (1980a,b), Bacescu (1990,
1992b) added 6 more. This paper describes fur-
ther new species, and summarises the distribution
of bodotriid species in Moreton Bay (Figs 1, 2).
Sexual differences within a species include
sculpturing and. armature of the body, and the
number of spine-like setae and fine setae on ap-
pendages. They also include more basic differen-
ces which enable greater swimming capacity in
the male and a marsapium in the 9 (Jones, 1963).
Specimens described were collected between
1989 and 1993. 787 plankton samples were taken
from 40 sites (Fig. 1). Most were taken by sledge-
net sampler (approx. 8556 at all sites), some by
plankton net (approx. 10% at sites 28, 29, 31), and
the remainder by re-entry tray (sites 28, 32) and
light-trap (site 24).
Two methods of specimen preparation for the
SEM were trialled, using 9 ? of Bodotria armata
sp. nov, (Fig, 8). The freeze-substitution method
(right hand sidc) eliminated the problem of salt
crystallisation and therefore proved to be the
most satisfactory method of SEM preparation
(Taufe, 1995),
Measurements of somites and segments are
taken along midlines of the entire sections, nut
just tht exposed portions. Length measurements
of whole specimens are recorded as standard
lengths, measured from the antenor end of the
pseudorostrum to the posterior end of the telsonic
somite, excluding spine-like setae and fine setae.
TERMINOLOGY. Terminology (Fig. 3) follows
the more recent works on the group (e.g., Day,
1975, 1278a,h, 1980; Watling, 1989, 1991a.b:
Felgenhauer. 1992; Roccatagliata, 1989, 1993),
392 MEMOIRS OF THE QUEENSLAND MUSEUM
et ie y
Moreton X
S i
a Pumicestone Passaze Boy (V
^ A DE
X slan
Lradhbroke
ts Land
153°00'E 20°
riG, 1. Satellite photograph of Moreton Bay showing sampling locations (supplied courtesy Ross Quinn through
Sunmap, from Landsat imagery, Australian Centre for Remote Sensing, Canberra).
BODOTRIIDAE OF MORETON BAY 393
BRIBIE \
Nono NN a —5
5E O7 | Y
ISLAND |
FIG. 2. Map of part of Moreton Bay showing sampling locations in upper Pumicestone Passage (from Queensland
Boating Safety Chart, Harbours and Marine, Brisbane).
394
X 7 P*eucorestium
E ynibe.
Wier AW. Ahlat lobe
K uy L Darapnte
Branrhini ordo
Werinpnd t
4 Fj td
Man wrewniiè T
T
———
7
pP Freeones
j c N
y = Fieonie v
zm.
Flagellum af Antenne e
Panne A
Jropod peduncle eme
FIG. 3. Diastylis rathkei. A, ? in dorsal view. B, adult
d in lateral view (after Sars, 1900).
As compared to older papers pereiopod replaces
peraeopod, segment replaces joint, pereionite
replaces pedigerous somile or thoracic somite,
pleonite replaces pleon somite or abdominal
somite and telsonic somite replaces anal or sixth
pleon somite. In descriptions of carapace fea-
tures, such as ridges and tubercles, the terminol-
ogy employed by Hale (1944a:66) has been
retained. In descriptions of appendages and their
setation, Watling (1989) and Felgenhauer (1992)
are followed. In relation to appendages lateral
refers to that margin/border of the appendage
facing the lateral body surface and medial refers
to that margin directed toward the midline of the
body.
The classification of substratum given in the
distribution section of each species follows the
U.S. Department of Agriculture (USDA) clas-
sification of grain sizes [1.e. coarse sand (2,000-
1,000 jum), medium sand (1,000-500jim), fine
sand (500-125yum), coarse silt (125-634um)
silmud (63-21.m)], as given in Gee & Bauder
(1986) and Giere et al. (1988). Geographic arcas
of the Australian marine environment are shown
on Fig. 4. The classification of abundance is:
uncommon- individuals taken in total of all
MEMOIRS OF THE QUEENSLAND MUSEUM
samples; common-10-100 individuals; very
common= 100-1,000; abundant=1,000 in-
dividuals.
ABBREVIATIONS. Abbreviations used in-
clude: L.V. (lateral view), D.V. (dorsal view),
V.V. (ventral view), D.L.V. (dorsolateral view),
V.L.V. (ventrolateral view), A.L.V. (antero-
lateral view), P.L.V. (posterolateral view), SEM
(scanning electron microscope), PSM (per-
manent slide mount), S.L. (standard length), QM
(Queensland Museum), AM (Australian
Museum), SAM (South Australian Museum).
RESULTS
Eight genera and 29 species (18 new) of
Bodotriidae were taken from Moreton Bay. Thir-
teen of those species are described, the remaining
five species lacking sufficient specimens to
enable full descriptions. The previously un-
described ? of Glyphocuma halei Greenwood &
Johnston is described. Twenty-two of the 29
species are new records for Moreton Bay.
Taxonomic keys are either updated to include
the new species, or newly constructed where
necessary. The keys to species of Cyclaspis,
Glyphocuma and Leptocuma are extended to in-
clude all known Australian species (Indo-
Australasian species in the case of Cyclaspis).
Where a high degree of sexual dimorphism exists
(e.g., in Glyphocuma), keys to the sexes are kept
separate.
Adults of Cyclaspis ornosculpta sp. nov. ex-
hibit à high degree of sexual dimorphism, typical
for members of the exsculpta group. Changes to
morphology during the development of C. ornos-
culpta sp. nov. are documented for both sexes.
KEY TO THE FAMILIES OF CUMACEA IN
MORETON BAY
(Adapted from Hale, 1946a; Jones, 1963, 1976)
1.No independent telson. Endopod of uropod with |
or 2 segments 2
Telson present. aed of raetiped with 2 or 3
segments
2,In both sexes the {att 4 pairs of péreiopodt iiher
without exopods or with very small rudimentary
exopods, de with 5 pairs of pleopots
eag aai ehe a debe w. , . Bodotriidae
In the ? at most the last 3 pairs of perciopods
without exopods, in male only the last, d with D
or 2 pairs of pleopods 4
3.d with one or two pairs of pleopods, ? third maxi-
BODOTRIIDAE OF MORETON BAY
Indian Ocean
Pd gt
^ONW iz
n; oceanic ; *
d)
5 7NW coast
7
>
"et.
ANZ
FeO) NW cisci
Central
W erasi
. N D^
oceanic y*
M Xs
nee
i Un
t S Gulfs coast
/ SW coust t » "
S oceanic
s>—0 *
@Bodotriidae
* Nannastacidae
*Gynodiastylidae
ADiastylidae
BLeuconidae
* Cyclaspis
Luke Eyre basin
- --— à (o:
395
Coral Sea
a Gulf of
Carpentaria
NE oceanic
x
s“
NE
creak
I Great X
l s Harrier
f Rer w
l ^ j
l NE evastal
e$
^ "E
re vi j
E
Tullius
River
hasta
sanadg Jo "ON
>
Mirciy-Darting hai
.-— 95
» » Central
N.S, W. Si E coast
KETIN, w
+
[j
l
=
|
|
|
‘
S Gulfs
& (3 M. SE oceanic
S S
= >
Ll
Tasman Sea £^
Q-«-
(o
OP»
FIG. 4. States, standard drainage divisions, coastal zones within the 200m bathymetric contour, and the 200
nautical mile Australian fishing zones (courtesy Crustacean Section, Australian Museum).
lliped with exopod (except Paradiastylis)
EP Diastylidae
d with no trace of pleopods. ? third maxilliped
lacksexopod .......... Gynodiastylidae
4.Endopod of uropod |-segmented, 9? has last 3
pairs of pereiopods without exopads, in d only
last pair. 3 without pleopods
PUM aerate, OF teeth TO Nannastacidae
Endopod of uropod 2-segmented. 9 has last 2
pairs of pereiopods without exopods, in & only
last pair. d. with 2 pairs of pleopods
Leuconidae
Family BODOTRIIDAE T, Scott, 190] emend.
Kurian, 1951
DIAGNOSIS. No free telson. Pleopods in males
only, with an outer process on endopod, usually
5 pairs, occasionally 2 or 3 pairs. Mandibles
narrow at base. Endopod of uropod 1- or 2-seg-
mented.
REMARKS, The family is distinguished from the
Diastylidae and Gynodiastylidae by the lack of an
independent telson, and from the Nannastacidae
and Leuconidae by the 5 pairs of pleopods in the
male (except for the Mancocuminae which are
not known from Australian waters), There are at
least 317 species of Bodotriidae described
worldwide, 94 of which occur in Australian
waters (including the new species described
below). Australian species are distributed
amongst the genera as follows: Bodotria, 4;
Cyclaspis, 58; Eocuma, 1; Iphinoe, 1; Gaus-
sicuma, 1; Gephyrocuma, 4; Glyphocuma, 6;
Leptocuma, 10; Picrocuma, 3; Pomacuma, 3;
Sympodomma, 2; Zenocuma, |.
The family is divided into 3 subfamilies; the
Bodotriinae Scott, 1901, have exopodites on only
the first pair of pereiopods, the Vaunthomp-
soniinae Sars, 1878 have exopodites on at least
the first 2 pairs of pereiopods; the Mancocuminae
396
Watling, 1977 have exopodites on at least the first
3 pairs of pereiopods.
KEY TO SUBFAMILIES AND GENERA OF
AUSTRALIAN BODOTRIIDAE
(Adapted from Jones, 1976 and Hale, 1944b)
|.Exopods on first pereiopods only . Bodotriinac.2
Well developed or rudimentary exopods on at
least first to third pereiopods
1232254282042 23429 Vaunihompsoniinac.5
2.Five free pereionites, Endopod of uropod 2-seg-
mented Iphinoe Bare
Four free pereioniles. Endopod of uropod 1 or 2-
segmented 3
3.Carapace with strong lateral ridges separating dor-
sal fram lateral regions throughout length
IAE TM n Bodotria Goodsir
Carapace without strong lateral ridges separating
dorsal from lateral regions throughout length 4
4.Uropods with peduncle more than half the length
of the rami. Carapace without lateral homs, En-
dopod of uropod l-segmented . Cyelaspis Sars
Peduncle less than half the length of the rami.
Carapace with lateral horns. Endopod of uropod
2-segmented Eocuma Marcusen.
5.First pereiopods with joints curiously expanded 4
First pereiopods not so modified
6.First antenna strongly geniculate, with joints of
peduncle sub-globose Gephyrocuma Hale
First antenna not strongly geniculate, joints not at
all globose
7 Telsonic somite well produced posteriorly. Basis
of first pereiopod with distal lobe
eb als fae de Me pe d Pomacuma Hale
Telsonie somite subtruncate, scarcely produced
posteriorly. Basis of first perciopod with no dis-
tal lobe Zenocuma Hale
8.Second pereiopod wilh a distal brush of setae on
propodus and dactylus, but no spine-like setae
V cent ot find BEE n Spice ap Leptocuma Sars
Second pereiopod without brushes of setae on ter-
minal segments but with spine-like setae on at
leastdactylus. -e llle. 9
9, Dorsal plate of telsanic samile subiruncate
posteriorly and not at all produced between
bases of uropods
Dorsal plate of telsonic somite rounded or an-
gular posteriorly and produced between bases of
WODE 2-0. 2 inii. ef. oon, 11
10.Dorsal plate of telsonic somite truncale posterior-
ly, Endopod of pleopods with narrow external
process Cumupsis Sars
Dorsal plate of telsonic somite excavated
posteriorly. Endopod of pleopods without exter-
nal process Heterocuria Miers
11. Third maxilliped with merus much longer than i$-
e 8 omo ee ee € Low UU * on ok
mys due ddr eder o t
MEMOIRS OF THE QUEENSLAND MUSEUM
chium but shorter than cure
b uenis uit. Vaunthompsonia Bate
Third maxilliped with merus subequal in length
to ischium and carpus 12
I2Ocularlobepresent .......... sess 13
Ocularlobeabsent .... sss se 14
13 Fourth pereiopod of male with exopod
Lied.ua4att6ejmxnd Glyphocuma Hale
Fourth peeeippod of male without exopod
, Sympodomma Stebbing
[4;Pseudorostrol lobes meeting in front of ocular
lobe. Anal portion of telsonic somite much
shorter than rest of somite— . Bathycuma Hansen
Pseudorostral lobes not meeting in front of ocular
lobe. Anal portion of telsonic somite as long as
rest of somite Gaussicuma Zimmer
Subfamily BODOTRIINAE Scott, 1901
DIAGNOSIS. Exopods present on first pair of
pereiopods; first pereionite usually concealed;
endopod of uropod often undivided.
Bodotria Goodsir, 1843
DIAGNOSIS. Cuticle strongly calcified.
Carapace with strong lateral ridges separating
dorsal from lateral regions throughout length.
First pereionite not visible in dorsal view, second
is long. In both sexes only first pereiopod with
exopod. Second pereiopod basis and ischium not
distinctly separated. Endopod of uropod one or
2-segmented, with distal segment always shorter,
Bodotria maculosa Hale, 1944
(Figs 5A-F, 6A,C,E)
Bodotria maculosa Hale, 1944b:226, figs 1,2, Hale,
19493:107, fig. 1.
MATERIAL EXAMINED. HOLOTYPE
SAMC2365, adult d, 4.2mm long, Dangerous Reef,
Spencer Gulf; PARATYPES SAMC2448, d', 3.5mm
long, off Port Hacking, NSW, 50 m, on sand; AM
unreg., Lizard Island, Qld; QMW20471, ovig. 9, S.L.
2.9mm, SEM mount, Horseshoe Bay, 27305,
153°21' E, sile 31, D. Tafe, 11 Oct. 1990, 2-3m, sand,
35.5ppt salinity, 24.5°C water temperature.
DIAGNOSIS. Carapace with median dorsal
ridge; lateral carina prominent, below which is
secondary carina which curves up postenorly to
meet primary lateral carina; dorsal surface with
coarse squamose-reticulate patterning formed by
large, shallow pits; line of shallow pits immedi-
ately above secondary latera] carina. Antennal
notch deep and narrow. Pseudorostral lobes wide,
truncate anteriorly, and reach apex of ocular lobe.
BODOTRIIDAE OF MORETON BAY 397
FIG. 5. A-F, Bodotria maculosa. A-C, à d . A, form from SA, LY. B, form from NSW, LY . C, telsonic somite
and uropod of S.A, form, DV. (A-C after Hale, 1944b). D-F, gravid 2 9. D, form from Western Australia, LV.
E, form without lateral carina on pleon, from WA, LV. F, telsonic somite and uropod of latter WA form. DV.
(D-F after Hale, 19492), G, Bodotria sp. nov. 2, gravid 9, LV.
398 MEMOIRS OF THE QUEENSLAND MUSEUM
SPS4F — SkU
FIG. 6. A, Bodotria maculosa Hale 9. carapace LV, shows squamose-reticulate pallerning on carapace and
absence of lateral carinae on pleonites. B, Bodotria armata sp. nov. ° carapace LV, shows lateral carinae on
pleonites and absence of squamose-reticulate patterning on carapace. C, Bodorria maculosa Hale 9 carapace
DLV, shows prominent lateral carina. D, Bodotria armata sp. nov. 9 carapace DLV, shows overlapping scales
and absence of lateral carina. E, Bodotria maculosa Hale 9 uropods VV, shows relatively long, cylindrical
peduncles. F, Bodotria armata sp. nov. 9 uropods VV, shows relatively short, angular edged peduncles.
BODOTRIIDAE OF MORETON BAY
Colour. Orange or brown with numerous black
chromatophores dotted over entire body,
S.L. Adult d 3.5-4.5mm, Adult Ŷ 2.9-3.1mm.
HABITAT AND DISTRIBUTION. On sand; 2-
50m depth. Central and Lower East Coast, S,
Gulfs Coast, Central and Lower West Coast (Fig.
3).
REMARKS. Bodotria maculosa resembles B.
arenosa Goodsir, 1843 and B. pumilio Zimmer.
1921 (=B. similis Calman,1907), but is easily
distinguished by the elevated dorsal carinae of the
last 3 pedigerous somites (Fig. 5A,D). It is further
distinguished from B. arenosa by the wider
carapace in dorsal view, and the relatively shorter
peduncle of the uropod (Figs 5C,F, GE); from H.
pumilio by the larger adult size (pumilio 2-
2.25mm). The Moreton Bay specimen agrees
closely with the type series and with the Lizard
Island specimens,
Hale noted considerable variation in the sculp-
ture of the carapace and size range of adults
(1944b:226), He described two males, one from
NSW (Fig. 5A) and 1 from SA (Fig.5B). The
NSW form is smaller, has more pronounced
squamose pitting of the carapace, and has more
slender thoracic appendages with longer spine-
like setae and fine setae, Two 9 9 forms were
described from WA (Hale, 19492). One form ex-
hibits strong lateral carinae on the pereionites and
pleonites (Fig. 5D) while in the other form such
carinae are totally lacking (Fig. 5E). When both
forms are adult as these are, such differences
indicate that more than one species may be in-
volved, Hale designated a type for both the SA
und NSW forms so it will be necessary to erect a
lectotype for the SA form to be the true B.
maculosa should further studies conclusively
separate the SA, WA and NSW forms as separate
species.
Hale's figure (1949, fig. 1B). of pereiopod 4
is probably incorrectly labelled and should read
'prp.5', since his own drawings of the whole
specimen show long basal segments on
pereiopods 1-4.
Bodotria armata sp. nov,
(Figs 6B,D,F, 7-11)
MATERIALEXAMINED_HOLOTY PEQMW20472
adult d (S.L. 3.2mm), PSM 353. Horseshoe Bay,
27°30'S, 153?21'E, site 31, D. Tafe, 11 Oct, 1990,
3.3m, sand, 35.5 ppt salinity, 745°C water tempera-
ture, PARATYPES OMW20473 ovig, 9, alkuype,
S.L, 2.9mm, PSM #54, same data as holotype;
QMW 20474 adult d, S.L. 3.0mm, PSM #51, same data
as holotype; QMW20475 ovig. 2, S.L. 2.7mm, PSM
#52, same data as holotype; QMW20476 ovig. 9 S.L,
2.6mm, SEM mount, same location as holotype, 7
April 1991; QMW20477 adult d , S.L. 3.1mm, SEM
mount, sume location as holotype, 7 April 1991;
QMW20478 adult d, S.L. 3.0mm, PSM #27, same
location as holotype, 4 Feb. 1993; QMW20479 avig.
9, S.L. 2.4mm, PSM #28, same location as holotype,
4 Feb. 1993,
DESCRIPTION. MALE. Integument calcified,
covered with small, rounded overlapping scales,
Carapace 0,25 S.L.; with strong median dorsal
ridge. almost straight in lateral view; lateral
ridges prominent, extending length of carapace;
carapace 0.74 as wide as long, lateral margins
rounded in dorsal view. Antennal notch deep and
narrow, Pseudorostral lobes wide, joining
anteriorly to ocular lobe which is as wide as long
(Fig.7A-G), Pereionite | almost fully concealed,
Percionite 2 as long as fourth or fifth, longer than
third; second to fifth pereionites with strong
median dorsal and lateral ridges (Fig. 7C,D,G).
Pleon robust and tapering in dorsal view: all five
pleonites with median dorsal ridges but without
defined lateral ridges: first 4 pleonites and tel-
sonic somite subequal in length. fifth pleonite 1.5
times as long as fourth (Fig. 7A-C,E,F.H). Tel-
sonic somite projecting postenorly over bases of
uropods, Posterior margin rounded with median
notch in dorsal view (Figs 7B.E,H, 8). First an-
tenna 3-segmented with terminal segmented
Magellum: first segment geniculate, longer than
next two segments combined; second segment
shorter and stouter than third, with two fine setae
distomedially, one distolaterally; third segment
with three fine distolateral setae; first segment of
flagellum twice as long as second, with three
proximolateral setae; second segment with two
acstelascs and (wo fine setae distally (Fig. 8A),
Third maxilliped as in B. maculosa. First
pereiopod with carpus reaching level of antennal
tooth of carapace; basis c.0,5 as long again as rest
of appendage, distal margin not produced, with 2
slender setae; ischium very short, c. 0.33 as long
as merus, both segments devoid of long setae;
carpus longer than merus and more than twice as
long as propodus, which is subequal in length to
dactylus; dactylus with 3-4 slender setae distally
and strong terminal spine-Jike seta, subequal in
length to segment: all segments of endopod with
short spine-like setae on posterior surface; Ex-
opod well developed; 2 slender proximal seg-
ments imd 5 short distal segments; all segments
except basal segment with 2 long setae distally
400 MEMOIRS OF THE QUEENSLAND MUSEUM
SPsS4nJ SKU u
rE
BODOTRIIDAE OF MORETON BAY
(Fig. 10A), Pereiopods 2-5 5-segmented (exclud-
ing coxa), ischium not separated from basis; car-
pus and dactylus subequal in length; terminal
spine-like seta longer than dactylus; small scat-
tered setae on all segments (Fig. IOB-E).
Pereiopod 2 with basal segment slightly longer
than combined length of remaining segments,
with 4-5 plumose setae and brush of small setae
on lateral margin; merus subequal in length to
carpus and twice as long as propodis; carpus and
dactylus each with three strong spine-like setae
distally (Fig. 10B). Pereiopod 3 with basis longer
than combined length of remaining segments:
merus shorter than each of remaining segments.
with 2 long setae distally; carpus shorter than
propodus, with one long seta distally; propodus
with spine-like seta distally, extending to distal
margin of dactylus, which itself bears two ter-
minal spine-like setae subequal in length (Fig.
10C). Pereiopod 4 with basis almost as long as
remaining segments combined, with
proximolateral seta, distolateral seta and lateral
brush of fine setae; merus 0.5 as long as carpus,
with 2 distolateral setae; carpus shorter than
propodus, with distolateral seta; propodus with
proximolateral seta and distomedial spine-like
seta, extending to distal margin of dactylus; dac-
tylus with 2 terminal spine-like setae subequal in
length (Fig. 10D). Pereiopod 5 with basis much
shorter than remaining segments combined, with
distolatera] seta; merus shorter than carpus, with
distolateral setà; carpus shorter than propodus,
with distolateral seta; propodus with distomedial
spine-like seta, extending to distal margin of dac-
tylus, which itself bears two terminal spine-like
setae subequal in length (Fig. 10E). Peduncle of
uropod 1.6 times as long as telsonic somite, lined
with plumose setae on whole length of inner
margin; endopod single-jointed, slightly longer
than telsonic somite or exopod, with 9-10 trun-
cated spine-like setae on inner margin, a short
subterminal and long terminal spine-like seta;
inner and outer margins more or less serrate;
exopod with 11-12 plumose setae on inner mar-
gin, long terminal spine-like seta and short ter-
minal simple seta (Fig. SB,C).
FEMALE. Integument calcified and covered with
any
FIG. B. Bodotria armata sp, nov., holotype adult 3 . A,
first antenna, B, uropod, DV, C. rami of uropod, DV.
small, rounded overlapping scales, as in the male
(Fig.6B,D). Carapace more than 0.25 S.L.; with
strong median dorsal and lateral ndges, as in
male; carapace 0.85 as wide as long, lateral mar-
gins rounded in dorsal view. Antennal notch and
pseudorostral lobes as in d (Figs 6B,D, 9B,D.H).
First perefonite almost fully concealed by second,
which is longer than percionites 3-5; all visible
pereionites have strong median dorsal and lateral
ridges, the dorsal ridges being distinctly keel-like
(Figs 6B,D, 9B). Pleon robust, all 5 pleonites with
median dorsal ridge and first 3 with defined
lateral ridges; first 4 pleonites and telsonic somite
subequal in length. fifth pleonite 1.5 times as long
FIG. 7. Bodotria armara sp. nov, 3, A,B, whole mount adult LV, shows relative lengths of carapace and somites.
C,E, whole mount subadnit LV, shows pronounced dorsal lobes of pereionites, lateral carinae on pleonites and
mild lateral carinae on pleonites 3-5, D, carapace adult LV, shows almost straight dorsal edge, overlapping
scales and absence of lateral carina, F, carapace adult DV, shows maximum width in mid-region of carapace
and median dorsal ridge extending over full length. G, carapace adult DLV, shows dorsal ridge extending over
full length and absence of squamose- reticulate patterning. H, whole mount adult DV, shows median dorsal
ridge extending continuously along carapace, pereion and pleon.
402 MEMOIRS OF THE QUEENSLAND MUSEUM
SP34F IAKU
BODOTRIIDAE OF MORETON BAY
| ~
a | d TN
\ y / ^ | Vl y
Nd LLLI t My
\ | ~ | /
e, Wy [a
ue Dn iJ J
FIG, 10. Badotria armata sp. nov., holotype adult c.
A, first pereiopod, B, second pereiopod. C, third
pereiopod. D, fourth pereiopod. E, fifth pereiopod.
as fourth (Figs 6B. 9H). Telsonic somite project-
ing posteriorly over bases of uropods (Fig. 1 1B).
First antenna 3-segmented with terminal seg-
mented flagellum, as in male. Third maxilliped,
first, third and fifth pereiopods as in male. Basis
of second pereiopod has only 2 plumose setae,
carpus only with 2 distal spine-like setae, Basis
of fourth pereiopod much longer than remaining
segments combined (Fig. LLA). Peduncle of
uropod 1.5 times as long as telsonic somite,
without plumose setae on inner margin but with
small scattered setae on posterior surface; en-
dopod single-jointed, slightly longer tham tel-
sonic somite and exopod, with | spine- like seta
and 6-8 short simple setae on inner margin, à short
subterminal and long terminal spine-like seta;
403
inner and outer margins more or less serrate;
exopod with 11 plumose setae on inner margin,
long terminal spine-like seta and short terminal
seta (Figs 6F, 1 1B,C).
Colour. Orange or brown with numerous black
chromatophores in both sexes.
S.L. Adult ¢ 3.0-3.2mm. Adult 9 2.4-2.9mm.
HABITAT AND DISTRIBUTION. On sand; 2-
4m; sites 15, 30, 31, 34 and 35. Both sexes are
abundant at site 31, and at times outnumber al]
other cumacean species.
REMARKS. Bodorria armata resembles B.
muculosa but is distinguished by the lack of
squamose-reticulate patterning on the carapace.
Also the scales of the carapace are more
pronounced (Figs 6D, 7D, 9D), and the overall
size is smaller (0,7x in compared specimens) than
that of B. maculosa. Minor differences in the 9
include relatively short, angular uropodal pedun-
cles and lateral carinae on the pleonites (Fig.6).
Minor differences in the d include second seg-
ment of first antenna shorter than third; dactylus
of first pereiopod subequal in length to propodus;
terminal spine-like setà of second pereiopod
longer than dactylus; uropod with 1-2 fewer inner
marginal spine-like setae.
The fourth pereiopod of the 9 has a much
longer basis than B, maculosa (Hale, 1949a, fig.
1B). While B. armata differs from both forms of
B. maculosa described by Hale it more closely
resembles the NSW form.
ETYMOLOGY. Latin.armara, armour, for the armour-
like overlapping scales on the carapace.
Bodotria sp. nov. |
(Fig. 5G)
MATERIAL. QMW20480 ovig. (S.L. 2.2mm), in
70% ethanol, site 31, D. Tafe, 11 Oct. 1990, 2-3m,
sand, 35.5 ppt salinity, 24.5°C water temperature;
specimen poorly preserved.
Colour. Whitish with scattered black pigment
spots.
S.L. Adult 9 2.2mm.
HABITAT AND DISTRIBUTION, On sand;
3m; 2 9 9from site 31, Moreton Bay.
FIG. 9. Bodotria armata sp. nov. 9. A, carapace DV, shows encrusting salt crystals. B, carapace DV, shows
surface free of salt crystals, C, ocular lobe DV, shows obscured ocular region. D, ocular lobe DV, overlapping
scales of ocular region visible. E, lower carapace LV, shows clinging salt crystals, F, lower carapace LV, shows
overlapping scales of lower carapace. G, whole mount LV, shows salt crystals over whole body surface. H,
whole mount LV, shows clean body surface with dimpled texture of carapace fully visible.
404
FIG. 11. Bodotria armala sp. nov., allotype ovig, ?.
A, fourth pereiopod. B, uropod, DV, C, rami of
uropod, DV.
REMARKS. Bodotria sp. nov. 1 cannot be ade-
quately described until more specimens are
secured. The species resembles B. armata sp.
nov. but the carapace is shorter, broader in dorsal
view and not strongly calcified (Fig. 5G); the
scales on the carapace are not pronounced, the
colouration of the carapace is whitish with scat-
tered black pigment spots, even after being
preserved in 70% ethanol for two years (cf
brownish without pigment spots in B. armala),
and the uropods are more delicate.
Cyclaspis Sars, 1865
DIAGNOSIS. Cuticle strongly calcified. First
pereionite seldom visible in dorsal view, Ar-
MEMOIRS OF THE QUEENSLAND MUSEUM
licular pegs nearly always present on sides of
pleonites. Only first pereiopods bear exopod. d
with 5 pairs of pleopods. Peduncle of uropods
never much shorter than rami; endopod 1-seg-
mented.
REMARKS. 93 species world wide (Bacescu,
1988); found in all oceans, though most species
(6095) inhabit shallow water (10-50m) in
temperate latitudes of Southern Hemisphere.
KEY TO INDO-AUSTRALASIAN SPECIES
OF CYCLASPIS
(Adapted from Hale, 1944a)
SECTION |
Sides of carapace without ridges or tubercles in
either sex.
Viewed from above the lateral contour of the
carapace is always evenly curved. Carapace
usually smooth except for the reticulate pattern-
ing, but sometimes slightly roughened owing to
granules or pits.
|. Front margin of carapace with an acute, forwardly
directed spine on each side, below antennal
angle caprella Hale
No spines at front of carapace
e. breee om m on t n t9 n8
2.Pseudorostral lobes meeting for an appreciable dis-
lance in front of ocular lobe , . (picta group) 3
Pseudorostral lobes barely or not meeting in front
of ocular lobe (levis group) 1 !
3,Carapace smooth and delicate, with fine median
dorsal ridge for full length of carapace . ... 4
Carapace not as above
4.Carapace with a low median dorsal projection at
posterior end. Rami of uropod subequal in
length gibba Hale
Carapace with smooth dorsal profile throughout
length. Exopod of uropod distinctly longer than
endopod sp. nov. 2
5.Curapace with many longitudinal rows of minute
granules. Peduncle of uropod not longer than tel-
sonic somite .........- costata Calman
Carapace smooth. Peduncle of uropod much
longer than telsonic somile
5.Both rami of uropod with at least one articulated
terminal spine-like seta
Both rarni of uropod without terminal spine-like
setae
FIG. 12. Cyclaspis eretata Hale d . A, whole mounts LV and DV, shows relative lengths of carapace and somites.
B, whole mounts (part) DLV, shows carapace and percianites with dorsal carinae. C, carapace LV. shows
antennal notch, depth and length of carapace. D, carapace DLV, shows median dorsal ridge on anterior hall.
E, carapace DV, shows cretations, apertures and oscular region. F, carapace (dorsal region) DV, detail of
apertures in mid-dorsal region. G, Pereionites DV, shows brush of setae on margin of third pereionite. H,
uropods DV, shows relative lengths of peduncle and rami.
BODOTRIIDAE OF MORETON BAY 405
1 mr
“Sah
7H SU
406
7.First pereiopod short, with carpus not reaching
level of antennal tooth. Rami of uropod barely
half as long as peduncle, exopod with 2 outer
marginal spine-like setae . - . . . picta Calman
First pereiopod long, with carpus reaching level
of antennal tooth. Rami of uropod about 2/3 as
long as peduncle, exopod with 2 outer marginal
spine-like selae . 0... ee ee sp. nov. |
8.Exopod bears 4 mucrone. Carpus of first
pereiopod 1/3 as long again as propodus
Me qa aar zh lucida Hale
Exopod without mucrone. Carpus of first
pereiopod not longer than propodus . . . . .
9.Peduncle of uropod shorter than exopod. Terminal
spine-like seta of second pereiopod at least 1/3
as long again as dactylus daviei sp. nov.
Peduncle of exopod subequal or longer than ex-
opod. Terminal spine-like seta of second
pereiopod subequal in length to dactylus . . 10
10.Setae of third to fifth pereiopods long; 5 on car-
pus, the longest reaching for nearly hall of their
length beyond tip of dactylus - mollis Hale
Setae of third to fifth pereiopods short; 3 on car-
pus, none reaching beyond tip of dactylus
MEI. CUESTA. fulgida Hale
] L.Endopod of uropod with at least 1 articulated ter-
minal spine-hke seta or mucrone, Carapace
heavily calcified and pitted
alveosculpta sp. nov.d
Endopod of uropod with apex acute and without
articulated terminal spine-like setae or
muérones. Carapace not beayty pacified or
puéd oi ea ear i oe ge oe
12.Exopod of uropod with apex acute and tacking
terminal mucrones . 2. s. aaan ee 3
Exopod of uropod with one or more articulated
terminal mucrones = -. i -- e- -i 30
13.Carapace with numerous fine longitudinal striae
D Lad doe aet ae pert a ee tas strigilis Hale
Carapace without longitudinal striae . . . . . 14
14,Carapace with a low median dorsal projection at
posterior end 15
Carapace without median dorsal projection at
pesteigrend 4.2.2.2... a 76
15.Carapace with median dorsal ridge distinct for
whole length and with a conspicuous pit on each
side alongside posterior median projection.
Pedunele of uropad longer than rami
epatis uen edidit bag a» io alle sheardi Hale
Carapace with median dorsal ridge obsolete for
posterior half of length: no conspicuous pits al
posterior end, Peduncle of uropod shorter than
tali fb: e eae eu. nijobergi Zimmer
16.Carapace not globose, somewhat laterally com-
pressed in both sexes. Uropods slender, the
peduncle longer than the telsonic somite. . 17
Carapace globose in both sexes. Uropads slont,
MEMOIRS OF THE QUEENSLAND MUSEUM
the peduncle shorter than, or barely as long as,
telsonic somite... +e) ene onn n 26
17.Propodus of first peretopods almost as long as
merus and carpus together 18
Propodus of first pereiopods subequal in length
to carpus 19
18.Inner margin of endopod of uropod with a row of
setae, followed by 7- 8 slender spine-like setae
(adulld ) 2... llle levis Thomson
Inner margin of endopod of uropod with 3 to 6
proximal spine-like setae, followed by a row of
15-23 shorter spine-like setae (both sexes)
cretaia Hale
VIIJ
19.Carapace roughened with fine granules
Ce ha ER ey ge is granulosa Hale
Carapace notasabove |... ee ,20
20.Carapace with two mid-dorsal depressions, one
either side of dorsal ridge
ade dedii qe de Pe andersoni sp. nov.
Carapace not as above... . . , CE Pa AJ
21.Carapace somewhat compressed laterally with
strong dorsomedial ridge Poyagiiont length and
nootherridges . 2... ee ee es 22
Carapace not ás above .. oono 00.04 23
22.Peduncle of uropod shorter than rami. First
pereiopod with group of spinc-like setae (d ) or
fine setae (9 ) on medial bulge of basis, S.L.
AGM a manaa ie a wh A sallai sp. now,
Peduncle of uropod longer than rami. First
pereiopod without group of spine-like setae (d. ]
or fine setae (d ) on medial bulge of basis, SL.
SAT wad DU uu. cooki sp. nov.
23. Basis of first pereiopods with a large apical tooth-
like projection, reaching to distal a of
WS M. ttn q tust al a lupi 24
tooth m osm smash th deade boanie. ‘of apie Sa 25
24.Rami of uropod longer than peduncle (subadult
GORY oc solar dese formosae Zimmer
Rami of uropod equal in length to peduncle
(ovigerous do)... herdmani Calman
25.Pcduncle of nropod longer than rami. S.L. <6mm
au. Ra 9 acie -Ivu-T- a pe tt concinna Hale
Peduncle of uropod shorter than rami. S.L.»8mm
VE aed addet d doe candidoides Bacescu
26,Size small, ovigerous 9 3,5mm. Ocular lobe di-
lated anteriorly, with prominent circular dark len-
secs i. 7 . pusilla Sars
Size large, ovigerous 9 Tram or more. Ocular
lobe not dilated anteriorly but somewhat trian-
gular, with lenses pale and elongate .... 27
27, Caripace overhanging second pereionite
posteriorly. Third to fifth perciopods with long
Sela "ova. ehh ee i. globosa Hale
Carapace not overhanging second pereionite
posteriorly. Peretopods 3-5 with short setae
BODOTRIDAE OF MORETON BAY
28.Carapace coarsely pitted, slightly rugose. Pleon
robust. Dactylus of pereiopod 2 with longest ter-
minal spine-like seta shorter than propodus and
dactylus together... -. .. ... Clarki Hale
Carapace smooth or with extremely fine reticula-
ton, Pleon slender. Dactylus of pereiopod 2
with longest terminal spine-like seta as long as
propodus and dactylus together
29.Opuc lobe extends in front of pseudorostral
lobes. Peduncle of uropod has setae on inner
margin stocki Bacescu
Optic lobe does not extend in front of pseudo-
rostra] lobes. Peduncle of uropod is devoid of
setae on inner margin ....... pinguis Hale
30,Peduncle of uropods less than 1.5 times as long
as telsonic segment and subequal to rami. Ex-
opod with at least 2 elongate unequal mucrones
ardimal end. st moet fl. Laas 4 3l
Peduncle of uropods about twice as long as tel-
sonic segment and longer than rami. Exopod
wilh | or 2 short mucrones or spine-like setae al
distal end (less than 4x as long as broad). . . 32
31.Exopod of uropod with two unequal mucrones at
distal end; d with spine-like setae on basis of
first pereiopod; ? uropod without selae on inner
margin of exopod sublevis Hale
Exopod of uropod with 3 unequal mucrones al
distal end; d without spine-like setae on basis of
first pereiopod; ? uropod with selae on inner
marginofexopod....... tranteri sp. nov.
32. Exopod of uropod with one or more mucrones 33
Exopod of uropod with one or more spine-like
setae
33, Peduncle of uropod al most hall as long again as
rami. Endopad of uropod has proximal half of
inner margin naked or with plumose setae (adult
Oye Pie reu Nah. . pura Hale
Peduncle of uropod 2/3 as long again as rami, En-
dopod of uropod has 5-6 short spine-like setae
on proximal half of inner margin (adult d )
nitida Hale
34.Basis of first pereiopods only 3/4as long as rest
of limb, with a long apical tooth, reaching distal
margin of isehium hornelli Calman
Basis of first perciopods subequal! in length to
resi of limb, with apical tooth short, reaching
only mid length of ischium . . . . . colloni Hale
e jer e s t n oigo n
SECTION 2
Sides af carapace never smooth, but with ridges
or tubercles, or both. Viewed from above the
lateral contour of the carapae, owing to the sculp-
ture, is rarely evenly curved, particularly in the
g
| Sides of carapace almost smooth, with no
anterolateral tubercle, ridge or other projection
below pseudorostral suture
Sides of carapace never almost smooth. with al
407
least one anterolateral tubercle, ridge or toath-
like projection below pseudorostral suture . . 6
2,Carapace with a prominent mid-dorsal tooth aver
base ofocular lobe ..... uniplicata Calman
3.A slight but obvious incision in dorsal margin of
carapace at middle of length, Exopod of uropod
with no apical spine- like seta, but with mucrone
sabulosa Hale
No incision in dorsal margin of carapace at mid-
dle of length. Exopod of uropod with slender d"
calspine-like setā : s: 2.2:
4.Two ridges on each side of carapace, Propodus oy
first pereiopods subequal in length to dactylus
argus Zimmer
One ridge on each side of carapace, Propodus of
first pereiopods much longer than dactylus. . 5
5.Carapuce with postenor transverse ndge faint,
short and confined to posterior half of carapace.
ast $603 56 BA tee thamsoni Calman
Carapace without posterior transverse ridge but
with fine ridge curving obliquely forwards from
median dorsal ridge 10 antero-inferior margin
ba cae tae a elise als spilotes Hale
6. Chiapase fully encircled by a collar-like ridge
cingulata Calman
Curapace not encircled by a collar-like ridge , 7
7.A quadrilateral area on each side of carapace,
defined by ridges or tubercles (distinct and
depressed in 9 , often indistinct in £)
oti pe batt Eua (exsculpta group) 8
No quadrilateral area on side of carapace . . 26
8.Carapace with anterior transverse ridge crossing
dorsal midline (distinct in 9, oflen indistinct in
d) eie eoa dre. tomis
Carapace with anterior transverse ridge not cross-
ing dorsal midline 5.5... 23
9,Posterior transverse ridge of carapace with 2-3 dis-
tinet branching ridges. Propodus of pereiopod 1
as long as combined length of ischium, merus
and carpus
ea ee SE oe fu indoaustralica Bàcescu
Posterior transverse ridge of carapace without dis-
tinet branching ridges. Propodus of pereiopod 1
nol as lang as combined Jength of ischium,
merus and carpus ... 2.6... ee eee 10
10.Carapace heavily calcified and composed of a
reticulate network of deep cralers
d ars ade. d RA agrenosculpta sp, nov,
Carapace may be heavily calcified and even
pitled but not composed of reticulate network of
detroralBis iov. ero. anni peek. l1
11.Post-ocular tubercle on midline of carapace, im-
mediately in front of anterior transverse ridge.
Surface of carapace studded with blunt spines
anteriorly tribults Hale
No post-ocular tubercle or blum spines on
anterior of carapace
408 MEMOIRS OF THE QUEENSLAND MUSEUM
—'18arMm
—— 168rm
236 13mm
m
BODOTRIIDAE OF MORETON BAY
12.Texture of carapace scale-like. Peduncle of
uropod much longer than rami, more than twice
as longinsubadult?........ bovis Hale
Texture of carapace pitted. Peduncle of uropod
subequal in length to rami .......... 13
13.Carapace with reticulate pattern of shallow
roundéd pits" sca uu. LES eA ees 14
Carapace with honeycomb structure of deep an-
BUlaT Pits? oaral gra uke + cae els is 19
14.Carapace with two lateral carinae running for-
wards from anterior transverse ridge to front
margin ofcarapace........ exsculpta Sars
Carapace without such carinae
15.Carapace with median dorsal ridge produced
posteriorly as keel-like lobe with notch
eve de eae cA e EIS gea a alveosculpta sp. nov.
Carapace with median dorsal ridge produced
posteriorly as rounded keel-like lobe without
HOW TA c, 255 2 20M A, el poy. S 16
16.Dorsolateral carinae present between two
transverse ridges of carapace ........ 17
No dorsolateral ridges between transverse ridges
17.Transverse ridges of carapace with denticula-
tions. SL. 9-]1mm . . . .. prolifica Bácescu
Transverse ridges of carapace smooth. S.L.
4-6MM on 54 7 98s. ornosculpta sp. nov.
18.Dactylus of first pereiopod at least - as long as
propodus candida Zimmer
Dactylus of first pereiopod less than 2/3 as long
as propodus............. usitata Hale
19.Carapace with pits separated by thick walls
Qa delat E «He chaunosculpta sp. nov.
Carapace with pits separated by network of thin
walls 2f 39443222: E b V .20
20.Longitudinal ridge runs obliquely from below an-
tennal tooth almost to end of carapace
FAQS RESTER. PEE mawsonae Hale
a re Moho ee Sogn he ey see Lh 21
21.Carapace with median dorsal keel forming 3
lobes posteriorly; posterior transverse ridge
rising on either side of median notch into
laminar tooth with sharp apex bending forward
anddownward ....... persculpta Calman
Carapace with median dorsal keel forming | lobe
posteriorly; posterior transverse ridge rising on
either side of median line to form blunt forward-
ly directed tooth. ............0.. 22
409
22.Carapace with small posterolateral lobes either
side of median dorsal lobe; exopod of uropod
without setae supersculpta Zimmer
Carapace without posterolateral lobes either side
of median dorsal lobe; exopod of uropod with
ROTA Ir. o rus Gerang tue agrenosculpta sp. nov.
23.Cephalothorax and pleon covered with small
spine-like setae; no ridges on back or sides of
carapace aspera Hale
Cephalothorax and pleon not covered with small
spine-like setae; well defined ridges on sides of
[apr M PORTE ARA 24
24.Posterior transverse ridge crosses dorsal midline
3 tke SPA MOM LEE australis Sars
Posterior transverse ridge does not cross dorsal
midline p TE 04e. A acy chy. "n 25
25.Quadrangular area on side of carapace with 4
prominent tubercles . . . . .. elegans Calman
Quadrangular area on side of carapace with 1 or
2 prominent tubercles... . .. similis Calman
26.Carapace with tubercles or ridges posterior to the
anterolateraltubercles............ 27
Carapace without tubercles or ridges posterior to
the anterolateral tubercles. ......... 32
27.Side of carapace with 3 obliquely transverse
cdrinae rg eS oe PAS TS Eussdis 28
Side of carapace with 1 transverse curved carina
OR NONE P ta an yaza coe es net ed chee
28.Carapace with reticulate pattern of deep angular
piis. cc Se et ae mawsonae Hale
Carapace without deep angular pits ...... 29
29.Carapace with mid-dorsal projection at hinder
MATIN. solis uel triplicata Calman
Carapace without mid-dorsal projection at hinder
MASIN o v, r SER ESSA sibogae Calman
30.Carapace covered in minute sparse spinules with
a well marked spinose median dorsal ridge for
whole length; on each side are four rounded
tubercles; short ridge leading back from antennal
notch (d) cana Hale
Carapace not as above; longitudinal ridge run-
ning from antennal tooth to about middle of
carapace length
31.Dorsum of carapace, as seen from side, rising
abruptly to an angular peak at middle of length
b p a oe EZ. Ma Bee AB.) shy LM simula Hale
Dorsum of carapace smoothly rounded
ose pes Pa oT ev oe coelebs Calman
FIG. 13. Cyclaspis cretata Hale 2. A, whole mounts DV, shows relative lengths of carapace and somites. B,
whole mount DV, shows carapace, pereionites and pleonites with median dorsal carinae. C, carapace DLV,
shows articular peg and median dorsal carina of first pleonite. D, carapace DV, shows ocular region, apertures
and absence of setae on margin of third pereionite. E, carapace DLV, shows median dorsal ridge on anterior
half and median dorsal depression on posterior half. F, carapace DLV, shows apertures in the bases of both
anterior cretations and collar-like first pereionite. G, carapace DLV, shows anterior cretations with apertures
and posterior cretations without apertures. H, uropods DV, shows relative lengths of peduncle and rami.
410 MEMOIRS OF THE QUEENSLAND MUSEUM
LED EZ
aN
H
is
i
È
i
FIG. 14. Cyclaspis species. A-E, Cyclaspis cretata. A, adult d, cephalothorax, LV. B, 8, DV. C, gravid 9,
cephalothorax, LV. D, DV. (A-D after Hale, 1948). E, paratype 3 uropod. F-I, Cyclaspis fulgida, type 9 . F,
LV. G, cephalothorax, DV. H, chromatophores of integument. I, uropod, DV. J-P, Cyclaspis strigilis. J-L,
type d. J, cephalothorax, DV. K, LV. L, uropod, DV. M-P, paratype 2. M, uropod, DV. N, first pereiopod.
O, cephalothorax, DV. P, cephalothorax, LV (E-P after Hale, 19442).
BODOTRIIDAE OF MORETON BAY
FIG. 15. Cyclaspis usitata gravid 2. A, first antenna.
B, mandible. C, pereiopod 4, D, first maxilla. E,
second maxilla. F, uropods, DV. G, rami of uropod,
32.Ocular lobe narrow, much longer than wide. Pere-
jopod | with dactylus about as long as carpus
ET ems sau Ros cert munda Hale
Ocular lobe as wide as long, much longer than
wide. Pereiopod | with dactylus less than half as
long as carpus pruinosa Hale
Cyclaspis cretata Hale, 1944
(Figs 12, 13, I4A-E)
Cyclaspis cretata Hale, 1944a:91, figs 19,20. Hale.
1948:4, fip. 2.
Cyclaspis ? cretata Stephenson et al., 1978:210.
MATERIAL EXAMINED. HOLOTYPE SAMC2418
d (6mm long), Cronulla, NSW, 2.5m, K. Sheard.
submarine light, Sept. 1942, PARATYPES
SAMC2366, C2368, C2370, C2371 d, 9,4.2-5.3mm
long, Spencer Gulf, SA, 6m, K. Sheard, Feb. 1941.
OTHER MATERIAL QMW?20481 adult d, S.L.
4.6mm, SEM mount, Rainbow Channel, site 26,
27°27'S, 153723" E, D. Tafe, 16 Feb. 1989, 6m, on sand,
411
34 ppt salinity, 25°C water temperature; QMW20482
adult 9 , S.L. 4.2mm, SEM mount, same data as above:
QMW20483 adult d, S.L. 4.5mm, PSM #41, Horse-
shoe Bay, site 31, 27^30' S, 153721 E, D. Tafe, 14 Aug.
1991, 2-3m, on sand, 27.2 ppt salinity, 16.5°C water
temperature.
Colour. Whitish with faint brown mottling;
sparse black chromatophores sometimes present
on carapace and abdomen.
S.L. Adult d 4.2-6,0mm. Adult 9 4.0- 5.3mm.
HABITAT AND DISTRIBUTION. On sand
often around coral reefs; 2-25 m; from S. Gults
Coast and Lower West Coast (Fig. 4). NSW:
Cronulla. SA: Kangaroo Is. Memory Cove, Spen-
cer Gulf (Hale, 19443). OLD: Sandy Cape, Noosa
River, off Moreton Island (Hale, 1948), Moreton
Bay (common in sandy areas). WA; Rottnest
Island, Shark Bay, South Passage, Thomson Bay,
Abrolhos Islands, Turtle Bay, off Walabi Island
and Andaman Island (Hale, 1948),
REMARKS. Moreton Bay specimens matched
Hale's type specimens but are slightly smaller
than his NSW and SA specimens. Both sexes are
distinguished from other species by the pattern of
pitting on the posterior dorsal surface of the
carapace (Figs 12A-F, 13A-G, 14A-D). 8 d and
2 9shavc4 depressions in the mid- dorsal region
of the carapace, located in the bases of the
anterior pits (Figs 12D-F, 13D-G). d d have a
tuft of short setae on the dorsum of the third
pereionite (Figs 12G, 14A), which is not present
on the 9 (Figs 13C-F, 14C). The uropods of the
Moreton Bay d 6 (Fig. 12H) are identical to
those figured by Hale (Fig. 14E). The uropods of
the ? have not been previously figured; those
shown here (Fig. 13H) match the original descrip-
tion.
Cyclaspis fulgida Hale, 1944
(Fig. I4F-I)
Cyclaspis fulgida Hale, 1944a:80, figs 9,10. Hale,
1948:24, fig. 13. Hale, 1953a:72.
MATERIAL EXAMINED. HOLOTYPE
SAMC?424, ovig. V, length 5.75mm, K. Sheard, sub-
marine light, Sept. 1942, 2.5m, Cronulla, NSW.
OTHER MATERIAL QMW20484, adult d, S.L.
3.9mm, PSM #55, Horseshoe Bay, site 31, 27730 S,
153?2I E, D. Tale, 13 Aug. 1990, 2-3m. on sand, 27
ppt salinity, 16.5*C water temperature.
Colour. Whitish, sooty black chromatophores
sometimes present on carapace, pereiopods and
abdomen.
S.L. Adult d 3.9-5.5mm. Adult 9? 5.75mm.
412
FIG, 16, Cyclaspis usitata subadult d. A, first
pereiopod. B, second pereiopod. C, uropods, DV.
HABITAT AND DISTRIBUTION. On sand; 2-
5m; Central East Coast, Lower East Coast, and
Lower Wesi Coast (Fig. 4). NSW: Cronulla
(Hale, 19442). WA: Garden Island, Careening
Bay (Hale, 1948). Qld: Moreton Bay (Hale,
1948).
REMARKS. Moreton Bay specimens match
Hale'stype specimens from Cronulla. The cuticle
is smooth and scarcely calcified and the carapace
ovoid in dorsal view. It has a shallow lateral
depression on each side of a low median dorsal
ridge, which continues towards the posterior as a
faint double ridge. The ocular lobe is prominent
and slightly elevated. The pseudorostral lobes
meet in front of the ocular lobe. The colour of the
MEMOIRS OF THE QUEENSLAND MUSEUM
cuticle is white with scattered, sooty black
chromatophores. This species resembles C. mol-
lisbutthe ocularlobe is more prominent, the basis
of the first pereiopod has a well developed dis-
tomedial tooth and the endopod of the uropod has
13 (cf, 8) medial spine-like setae (Fig. 141). Com-
mon in Moreton Bay.
Cyclaspis strigilis Hale, 1944
(Fig. 14J-P)
Cyclaspis strigilis Hale, 1944a:83, figs 11-14.
MATERIAL EXAMINED. SAMC2412-2413 d, 9,
off Fraser Island, Qld, 24720' S, 153°02°E. (Warreen'
Mar. 1938); AMP22642 d, 9, North Head, Sydney;
QMW20485 ovig. 9 ,S.L. 4.7mm, PSM #20, Pumices-
tone Passage, site #12, 26°49°S, 153^8 E, J. Green-
wood, 26 Jan 1990, 2m, on sand, 34.9 ppt salinity,
27.3*C water temperature.
Colour. Colourless to whitish, with a few
brown chromatophores on carapace, pereiopods
and abdomen.
S.L. Adult à 4.4mm. Adult 93.6-4.7mm.
HABITAT AND DISTRIBUTION. On sand; I-
7 m; NE and Central E Coast. Qld: Fraser Island
area (Hale, 1944a), Moreton Bay.
REMARKS. 9 first pereiopod of Moreton Bay
specimen with inner apical angle of basis rounded
rather than pointed as shown by Hale (Fig. 9N).
S.L. of Moreton Bay specimen is 30% larger than
the type specimen. Otherwise the specimen
agrees exactly with the original description.
Cyclaspis usitata Hale, 1932
(Figs 15. 16)
Cyelaspis usitata Hale, 1932:549, fig.1; Hale,
1944a:122, figs 43,44.
MATERIAL EXAMINED. HOLOTYPE
SAMC1841, 2, S.L. 10mm, St. Vincent Gulf, SA.
OTHER MATERIAL AMG917, 2; QMW20486 ovig.
9, S.L. 6mm, PSM #12, Pumicestone Passage, site
#12, 26°49'S, 153°8"E, J. Greenwood, Mar. 1992, 2 m,
on sand, 35.4 ppt salinity, 24.5°C water temperature;
FIG. 17. Cyclaspis alveosculpta sp. nov. d. A, whole mount LV, shows relative lengths of carapace and somites.
Also shows affects of partial digestion in fish, B, carapace LV, shows shape of carapace in lateral view, C,
Anterior carapace LV, shows reduction of anterior transverse ridge and pattern of pits in integument. D, carapace
ALY, shows reduction of anterior and posterior transverse ridges and position of ocular lobe, E, carapace LV,
detail of integument showing reticulate pattern of scales, F-H, Cyclaspis cooki sp. nov. 9 F, carapace LV, shows
delicate, finely reticulate integument and prominent sub-acule antennal tooth. G, Pleonites 1-3 LV, shows
articular notches and median dorsal ridge. H, Pleonites 1-3 DLV, shows fine median dorsal ridge of pleon.
BODOTRIIDAE OF MORETON BAY
413
MEMOIRS OF THE QUEENSLAND MUSEUM
i8BmPm
BODOTRIIDAE OF MORETON BAY
QMW20487 sub-adult , S.L. 5.4mm, PSM #42,
Pumicestone Passage, same data as above;
QMW20488, sub-adult d, S.L. 6mm, PSM #14, off
Macleay Island, 27°36’S, 153?22'E, D. Tafe, 9 Nov.
1989, 1.5 m, on fine sand, 34 ppt salinity, 25°C water
temperature.
Colour. Whitish to yellow.
S.L. Adult &: 6-8mm, Adult 9: 5-9mm.
HABITAT AND DISTRIBUTION. On sand; 1-
"Im; S Gulfs Coast, Lower and Central E Coast.
SA: St. Vincent Gulf. NSW: Jervis Bay (Hale,
1944a). Qld: Moreton Bay.
REMARKS. The Moreton Bay form is smaller
than the SA and NSW forms but similar in size to
the WA form. The first antenna, fourth pereiopod
and uropod of the 9 (Fig.15) match those of Hale
(1944a:123). The d has not as yet been described.
The subadult ¢ from Moreton Bay cannot be
described as the d of the species, considering the
extent of morphological change which occurs in
all members of the exsculpta group during
maturation. Hale (1948) suggested that C.
mjobergi Zimmer may well be the d of C. usitata.
Certainly the description of C. mjobergi (d)
given by Zimmer (1921) closely resembles the
above subadult d of C. usitata (3), however, the
length of the former species (type) is recorded as
14mm. Smaller specimens (8-9mm) of C.
mjobergi (à ) from St. Vincent Gulf (Hale, 1944a:
88), differ in segmentation and setation of the
pereiopods to the above subadult c of C. usitata
(Fig. 16).
Hale (1944a:122) recorded a large number of
99 of C. usitata, c.7mm in length, from
Brighton, SA. He also recorded an ovigerous 9,
6mm long, from Shark Bay, WA (1948:41). The
types of C. candida and C. mjobergi (Zim-
mer, 1921), both appear to be too large (12.5 and
14mm respectively) to be considered the d of C.
usitata.
Cyclaspis cooki sp. nov.
(Figs 17F,G,H, 18-20)
MATERIAL EXAMINED. HOLOTYPE
QMW20520, adult d, S.L. 5mm, PSM #48, Pumices-
tone Passage, site 12, 26°49’S, 153^8' E, J. Greenwood,
415
FIG. 19. Cyclaspis cooki sp. nov. A-B, holotype adult
d. A, first antenna. B, first pereiopod. C, second
pereiopod. D, third pereiopod. E, fourth pereiopod. F,
fifth pereiopod.
4 May 1990, 2m, coarse sand, 33.3 ppt salinity, 23.5°C
water temperature. PARATYPES QMW20521, ovig.
9, allotype, S.L. 2.5mm, PSM #47, same data as
holotype; QMW20522, adult d', S.L. 3.06mm, SEM
mount, off Coochiemudlo Island, site 35, 5m, sand,
27°34'S, 153°21"E, D.Tafe, 17 June 1990, 34 ppt
salinity, 18°C water temperature; QMW20523, adult
9, S.L. 2.4mm, SEM mount, same data as above.
DESCRIPTION. MALE. Integument lightly cal-
cified, delicate and easily broken, even when
freshly caught; finely reticulate and sparsely
pitted (Fig. 18A,C,E). Carapace length 0.32 S.L.
and 0.48 as wide as long in dorsal view, with
FIG. 18. Cyclaspis cooki sp. nov. A, d DV, shows relative lengths of carapace and somites. B, 9 LV, shows
relative lengths of carapace and somites. C, d carapace DV, shows shape and texture of carapace. D, 9 carapace
DV, shows shape and texture of carapace. E, & carapace DLV, shows strong median dorsal ridge extending
throughout. F, 9? carapace DLV, shows strong median dorsal ridge extending throughout. G, d uropods DV,
shows relative lengths of peduncle and rami. H, 9 uropods VV, shows relative lengths of peduncle and rami.
416
FIG. 20, Cyclaspis cooki sp, nov. A-B, holotype adult
d. A, uropods and telsonic somite, DV. B, rami of
uropod, DV. C-D, allotype ovig. V. C, uropods and
telsonic somite, DV. D, rami of uropod, DV.
strong median dorsal ridge throughout length,
lateral margins evenly rounded, dorsal edge
slightly arched, pseudorostral lobes barely reach-
ing apex of ocular lobe; sides of carapace devoid
of ridges or sculpture; antennal notch moderate
and visible in dorsal view; antennal tooth sub-
acute, no antennal ridge, pseudorostral lobes
wide, joining just anterior to ocular lobe which is
almost as wide as long, rounded, with 9 lenses, 3
central surrounded by 6 outer lenses (Fig. 18C,E).
Pereion 0.48 as long as carapace; first pereionite
fully concealed; second pereionite a narrow col-
lar posterior to carapace; lateral section shorter
than each of remaining 3 overlapping percicnites
(Fig. I&A,E).
MEMOIRS OF THE QUEENSLAND MUSEUM
Pleon robust with median dorsal ridge and well-
developed lateral articular processes; first 4
pleonites subequal in length. fifth pleonite 1.3
limes as long as fourth (Fig. 18A). First antenna
3-segmented with terminal segmented flagellum;
first segment somewhat geniculate, longer than
second and third segments. combined; second
segment 0.9 times third segment, with 2 fine setae
distolaterally; third segment with 2 slender setae
distomedially and one distolaterally; first seg-
ment of flagellum twice as long as second, which
has 2 aestetascs and 2 fine setae distally (Fig. 19).
All pereiopods 7-segmented, with terminal spine-
like setae longer than dactylus, except in first
pereiopod; some spine-like setae have rows of
fine spinules distally (Fig. 19). First pereiopod
with basis 1.2 times length of remaining segments
combined, with plumose seta distolaterally; is-
chium 0.6 times length of merus which is 0.7
times length of carpus; carpus subequal in length
to propodus, which is 1.4 times length of dac-
tylus, with 1 medial and 2 distal setae; dactylus
with 2 slender terminal spine-like setae, | almost
as long as itself, 1 terminal seta and 2 stout
sublerminal setae; exopod well developed, wide
proximal segment with 1 short plumose seta dis-
tolaterally and 8 shorter distal segments, each
with 2 long setae (Fig. 19). Second pereiopod
with basis 0.8 times length of remaining segments
combined, with minute distomedial and dis-
tolateral setae; ischium 0.4 times length of merus,
with plumose seta distomedially; merus 1.7 times
length of carpus, with plumose seta distomedial-
ly; carpus 1.4 times length of propodus, with 2
spine-like setae distomedially and | spine-like
seta distolaterally; propodus 0,5 times length of
dactylus, which has 2 terminal spine-like setas,
onc slighily longer than itself, ] small terminal
seta and | subterminal spine-like seta (Fig. 19).
Third to fifth pereiopods with merus longer than
ischium, carpus as long as combined length of
propodus and dactylus, the longer of which is
propodus: basis with 1-3 plumose setae medially;
ischium with 3 setae distomedially; merus with
sets distomedially; carpus with 2 spine-like setae
and | fine seta distolaterally; propodus with
spine-like seta and minute seta distally; dactylus
FIG. 21. Cyelaypis tranteri sp. nov. d. A,B, whole mount LV, shows relative lengths of carapace and somites.
C. carapace LV, shows subacute antennal tooth and first pereiopod. D, carapace DV, shows maximum width
in mid-region and median dorsal ridge visible along anterior 2/3 of carapace. E, carapace LV, shows fine
reticulate pattern of pits, and relative Jength and depth of carapace. F, Pereiopods LV, shows very long basal
segment of second pereiopod. G, uropods DV, shows relalive lengths of peduncle and rami H, Distal end of
exopod DV, shows two setose terminal mucrones and one small curved terminal seta.
BODOTRIIDAE OF MORETON BAY 417
igm
weg. 14mm
igearm
R
—— 185Pm _ =e a= 1, T m 5138 15mm
z
il00Hp _»
12mm
H
SPIM
418 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 22. Cyclaspis tranteri sp. nov. 3. A, distal end of
exopod of uropod DV, shows two setose terminal
mucrones at apex of exopod. B-D, distal end of ex-
opod of uropod (part) DV. E, distal end of exopod of
uropod (part) DV, detail of mucrones showing con-
voluted appearance of each shaft distally and fibrous
appearance of setules.
BODOTRIIDAE OF MORETON BAY
with terminal spine-like seta, terminal seta and
subterminal seta (Fig. 19D-F). Third pereiopod
with basis slightly shorter than remaining seg-
ments combined; ischium 0.4 times length of
merus; merus 0.7 times length of carpus;
propodus 1.7 times length of dactylus (Fig. 19D).
Fourth pereiopod with basis 0.7 times length of
remaining segments combined; ischium 0.4 times
length of merus; merus 0.65 times length of car-
pus; propodus 1.7 times length of dactylus (Fig.
19E). Fifth pereiopod with basis 0.5 times length
of remaining segments combined; ischium 0.4
limes length of merus; merus 0.7 times length of
carpus; propodus 1.7 times length of dactylus
(Fig. I9F). Peduncle of uropod 1.8 times as long
as telsonic somite, lined. with 14-15 plumose
setae On inner margin; endopod 0.9 times as long
às peduncle, subequal in length to exopod, with 6
slender spine-like setae und 11-12 stout spine-
like setae on proximal 3/4 of inner margin, apex
pointed, without spine-ljke setae or mucrones;
exopod with 6-8 plumose setae on proximal 2/3
of inner margin, apex pointed, without spine-like
setae or mucrones (Fig. 19A,B).
OVIGEROUS FEMALE. Integument lightly cal-
cified, finely reticulate and sparsely pitted, as in
d (Fig. 18B,D,F). Carapace length 0.35 S.L. with
strong median dorsa! ndge and sides devoid of
ridges or sculpture, as in g; width 0.48 times
length in dorsal view; antennal looth subacute
and extending to anterior extremity of carapace;
ocular lobe and pseudorostral lobes as in d. (Figs
I7F, I8D,F), Pereion 0.4 times as long as
carapace. First pereionile fully concealed by
second, both produced ventrally to form the mar-
supium; pereionites with dorsal ridge, dor-
solateral margin of fifth with articulation notch
(Fig. 18B). Pleon robust, all five pleonites with
dorsal ridge and lateral articulation notches; first
4 pleonites and telsonic somite subequal in
length, fifth pleonite 1.5 times as long as fourth
(Figs 17G.H. 18B). Telsonic somite projecting
posteriorly over bases of uropods (Figs 18B,
20C). First antenna 3-segmented. with terminal
segmented flagellum, us in d . Pereiopods as in 3
except: merus of second pereiopod has stout
spine-like seta distolaterally; carpus of third to
fifth pereiopods cach have 3 distal spinc-like
setae rather than 2 spine-like setac and | minute
seta. Peduncle of uropod 1.6 times as long us
telsonic somite, without plumose setae on inner
margin; endopod 0.75 times as long as peduncle,
subequal in length to exopod, with 5 spine-like
setae on proximal 2/3 of inner margin, apex
419
pointed, without spine-like setae or mucrones;
exopod with 2 plumose setae on proximal 1/3 of
inner margin, apex pointed, without spine-like
setae or mucrones (Fig. 20C,D).
Colour. Cream with small black
chromatophores speckled on carapace and ab-
domen.
S.L. Adult g 3.0-3, 5mm. Adult 9 2.4-2.5mm.
HABITAT AND DISTRIBUTION, Medium and
coarse sand; 1-5m water depth; sites 12, 34, 35 18
Moreton Bay.
REMARKS. Cyclaspis cooki resembles C pura
Hale from S Aust and C. juxta Hale from WA, but
the rami of the uropods are longer relative to the
peduncle and 1he setation of the pereiopods and
uropods is different (Figs 19, 20). Both sexes are
common in Moreton Bay.
ETYMOLOGY, For Steye Cook, Queensland
Museunt, who collected the types.
Cyclaspis tranteri sp. nov.
(Figs 21-23, 241,3}, 25)
MATERIAL EXAMINED. HOLOTYPE
OMW20489, adult d, S.L. 2.7mm. PSM #3-
PARATYPES QMW20490, ovig. 9. allotype, S.L.
2.45mm, PSM #2, sume dala asholetype; QMW 20491,
adult g, S.L. 2 5mm. PSM #4; QMW20497, adult F,
S.L. 2.5mm. PSM 31; QMW?20493, adult g, S.L.
2.6mm. SEM mount; QMW20494, ovig. 9, S.L.
2.4mm, SEM mount: all types from Raby Bay, site 32,
27°30°S, 153°18'E, D. Tale, 22 July 1989, Sm, sandy
mud, 34 ppt salinity, 24.8"C water temperature same
data as holotype
DESCRIPTION. MALE. integument thin and al-
most membranous, with fine reticulate pattern of
pits (Fig.21 A-E). Carapace (0.27 S.L. twice us
long as deep; in profile the dorsal margin evenly
curved except for the protruding ocular lobe, with
mild post-ocular depression; median dorsal ridge
is Visible along anterior 2/3 of carapace: in dorsal
view carapace js widest in mid-region and tapers
slightly towards the front and rear; antennal notch
widely open, antennal tooth subacute and visible
in dorsal view, no antennal ridge; pseudorostyil
lobes tapering anteriorly and joining just below
ocular lobe (Fig. 21D), join not visible in dorsal
view; ocular lobe as wide as long. rounded with
distinct corneal lenses, and located at anterior
extremity of carapace (Figs 21C-E, 24A, ED).
Percion 0,55 times as long as carapace; firs
pereionite. fully concealed; second pereionite
forming a narrow collier posterior to carapace; as
420 MEMOIRS OF THE QUEENSLAND MUSEUM
E =
___ a M Ul 8 ach. a 5 13mm
SKU
BODOTRIIDAE OF MORETON BAY
long as third but shorter than
fourth and fifth pereionites
(Figs 21C,D, 24A,E,1). Pleon
slender and long, no dorsal or
lateral ridges; first 4 pleonites
subequal in length, each with
small lateral articular notches;
fifth pleonite 1.5 times as long
as fourth (Fig. 21A,B). Tel-
sonic somite subequal in length
to fourth pleonite, with
posterodorsal projection (Figs
21B. 24F, 25B). First antenna
asin C. sublevis. Second anten-
na with flagellum reaching to
posterior end of peduncle of
uropod (Fig. 21B). AI
pereiopods 7-segmented, with
terminal spine-like setae
longer than dactylus. First
pereiopod with carpus reach-
ing beyond level of antennal
tooth; length of basis 1.2 times
rest of appendage, with dis-
tomedial margin produced into
blunt tooth and with plumose
distolateral seta; ischium 0.5
times length of merus which is
0.6 times length of carpus; car-
pus 0.9 times Jength of
propodus and subequal in
length to dactylus; propodus
1.1 times length of dactylus,
with 2 medial and 2 dis-
tomedial setae; dactylus with 2
slender spine-like setae and J
seta terminally, 1 spine-like
seta and | seta subterminally
and | seta on medial margin;
exopod well-developed, very
broad proximal segment bear-
ing plumose seta distally, and 8
short distal segments, each with 2 long, plumose
setae (Figs 21C, 24A,H, 25A). Second pereiopod
as in C, sublevis except that terminal spine-like
seta is distinctly longer (1.3 times) than dactylus
and basis is c. twice length of remaining segments
combined (Fig. 2] A,F). Pereiopods 3-5 as in C.
sublevis, Peduncle of uropod 1.3 times length of
telsonic somite, with 20-21 plumose setae on
LV.
421
FIG. 24, Cyclaspis species, A-H, Cyclaspis sublevis, A, type G, LV. B, type
ovig. 9, LV. C, telsonic somite of F. LV. D, ? cephalothorax, DV. E, d
cephalothorax, DV. F, 3 uropod and telsonic somite, DV. G, 9 uropod and
telsonic somite. and m, mucrones of exopod. H, d first pereiopod. (A-H,
alter Hale, 1948). 1-J, Cyclaspis iranteri sp. nov. I, adult G, LV. J, ovig. 9,
inner margin in 2 rows; endopod subequal in
length to peduncle and 0.95 times length of ex-
opod, with 4 plumose spine-like setae and 11
naked spine-like setae on inner margin, and with
2 minute spine-like setae close to the acute distal
end; exopod with 5 plumose setae on inner mar-
gin, 2elongate unequal terminal mucrones (setac)
and | minute curved terminal seta (Figs 21G,H,
22, 24F, 25B,C).
FIG. 23, Cyelaspis tranteri sp. nov. V. A, whole mount LV, shows relative lengths of carapace and somites. C,
whole mount DV, shows bulbous carapace and slender pleon. E, carapace DV, shows maximum width in
postenor region and fine reticulate pattern of pits. F, carapace LY, shows relative length and depth of carapace.
G, uropods DV, shows relative lengths of peduncle and rami. H, Distal end exopod of uropod DY, shows two
terminal setose mucranes and one small nan-setose (convoluted) seta,
422
OVIGEROUS FEMALE. Integu-
ment thin and almost
membranous, with fine reticu-
late pattern of pits, as in ó
(Fig. 23A-E). Carapace 0.29
S.L. almost as deep as long; in
profile the dorsal margin has a
slight even curve except for
the protruding ocular lobe;
dorsal ridge is barely visible
along anterior 1/2 of carapace;
in dorsal view carapace is
widest in posterior 1/2 and
tapers anteriorly; antennal
notch less widely open than in
d ;antennal tooth subacute, no
antennal ridge; pseudorostral
lobes tapering anteriorly and
joining just below ocular lobe
(Figs 23A-F, 24B,D,J).
Pereion 0.6 times as long as
carapace; first pereionite nar-
row but visible; second
pereionite meets carapace
dorsally and continues smooth
curve of dorsal profile:
pereionites | 3-5 taper
posteriorly in dorsal view
(Fig. 23A-F). Pleon very
slender, no dorsal or lateral
ridges; first 4 pleonites sube-
qual in length, each with
minute latera! notches; fifth
pleonite 1.4 times as long as
fourth (Figs 23A-D, 24B,]).
Telsonic somite subequal in
length to fourth pleonite and
0.7 times length of fifth
pleonite, with posterodorsal
projection (Figs 23B,D, 24C).
First antenna as in d. Seg-
mentation of pereiopods as in
ó ; endopod of first pereiopod without distal tooth
and exopod with slender basis (Fig. 15D); seta-
tion of pereiopods 2-5 as in d. Peduncle of
uropod 1.3 times length of telsonic somite,
without setae on inner margin; endopod 1.1 times
length of peduncle, 0.9 times length of exopod,
with 4-7 naked spine- like setae on inner margin
and 2 minute spine-like setae close to the acute
distal end; exopod with 3-4 plumose setae on
inner margin, 2 elongate unequal terminal
mucrones (setae) and | minute curved terminal
seta (Figs 23G,H, 24G, 25E,F).
Colour. Cream and translucent.
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 25. Cyclaspis tranteri sp. nov, A-C, holotype adult d . A, first pereiopod.
B, uropods, DV, C, rami of uropod, DV. D-F, allotype ovig. 9. D, first
pereiopod, E, uropods, DV. F, rami of uropod.
S.L. Adult d. 2.4-2.7mm. Adult 9 2.3-2.5mm.
HABITAT AND DISTRIBUTION. In 4-7m over
silt and fine sand; Raby Bay, off Cleveland Point.
REMARKS. C. tranteri closely resembles C.
sublevis Hale from Broome, WA, and the Haw-
kesbury River, NSW (AMP28613). However, the
new species has 3 mucrones on the exopod of the
uropod, rather than 2 mucrones, and a long ter-
minal spine-like seta (longer than dactylus) on the
second pereiopod. In addition the d has no spine-
like setae on the medial bulge of the first
BODOTRIIDAE OF MORETON BAY
TABLE 1. Uropod spination of C. sublevis and C.
tranteri.
C. sublevis C. tranteri
cor [colon | 8 | èe |
C2997 | C2998
| 19 | 6 | 202 | 5 |
6 | 14-15
| s | o | 56 | 34 |
Exopod mucrones
pereiopod. The standard lengths of both d s and
9s are, on average, 8 to 1096 smaller than C.
sublevis. Spination of uropods compares with that
of C. sublevis (Hale, 1948:10, figs 5, 6) (Table 1)
The 2 larger mucrones are setose filaments
(Fig. 22). The third naked and smaller filament is
present in both sexes (Figs 21H, 23H).
C. sublevis belongs to the levis group and most
closely resembles the much larger New Zealand
C. calmani Hale (=levis Calman not Thomson)
(Hale,1948), but in C. calmani the basis of the
first pereiopod has no distal tooth and the rami of
the uropod are relatively shorter. There remains
confusion as to whether C. levis Thomson and C.
calmani Hale are two separate species or varia-
tions within the one species.
Thomson's original description and figures of
C. levis had a number of inaccuracracies (Cal-
man,1907; Hale,1944a; Jones,1963) wit the
major points of confusion being relative propor-
tions of segments of the first pereiopods and
presence or absence of terminal spine-like setae
on the rami of the uropods. Hale (1944a) con-
cluded that the differences between Thomson's
and Calman's specimens were consistent, in spite
of the confusion, and erected a new species, C.
calmani, which he thought was related to both C.
levis and C. sublevis. In the key we follow Baces-
cu (1988) in synonymising calmani and levis.
Peduncle setae
Exopod setae
(captions for two following pages)
423
Specimens of C. tranteri can be distinguished
from C. sublevis by the setae on the inner margin
of the exopod of the uropod in the 9 and by
structural differences of the first pereiopod in the
3. They include the absence of spine-like setae
on the basal segment of the endopod and a broad
basal segment on the exopod. These 2 species are
distinguished within the genus by the proportion
and spination of the uropod. Also the slender
dactylus of pereiopods 3-5 separates C. tranteri
and C. sublevis within the levis group. C. tranteri
was abundant over sandy mud.
ETYMOLOGY. For David Tranter, for his contribu-
tion to zooplankton research in Australia,
Cyclaspis ornosculpta sp. nov.
(Figs 26-30)
MATERIAL EXAMINED. HOLOTYPE
QMW20495, adult d, S.L. 5.04mm, PSM #57,
Pumicestone Passage, site 5, 26°52’S, 153°7’E, J.
Greenwood, 28 Jan 1991, 3m, sand, 36 ppt salinity,
31°C water temperature. PARATY PES. QMW20496,
ovig. 9, allotype, S.L. 4.76mm, PSM #58, Pumices-
tone Passage, site 3, 26°54’S, 153°5’E, otherwise same
data as holotype; QMW20497, adult d, S.L. 4.3mm,
SEM mount, same data as holotype; QMW20498,
ovig. 9, S.L. 4.25mm, SEM mount, same data as
holotype.
DESCRIPTION. MALE. Integument calcified,
with reticulate pattern of shallow pitting (Fig.
26A-D). Carapace 0.3 S.L. twice as long as deep;
in profile the dorsal margin is slightly convex
with mild post-ocular depression; median dorsal
ridge is visible along length of carapace and
pronounced on anterior half; in dorsal view
carapace is widest in region of anterior transverse
FIG. 26. Cyclaspis ornosculpta sp. nov. 3. A, whole mount (part) DV. B, whole mount (part) DV, shows relative
lengths of carapace and somites. C, whole mount (part) LV. D, whole mount (part) LV shows lateral articulation
notches of pleonites. E, Anterior carapace A.DV shows ocular lobe and reticulate pattern of pits. F, carapace
LV shows reduced anterior and posterior transverse ridges. quadrilateral area barely discernible. G, carapace
DV shows median dorsal ridge along length. H, uropods DV shows relative lengths of peduncle and rami.
FIG. 27. Cyclaspis ornosculpta sp. nov. A, carapace adult d LV, shows reduced sculpture. B, carapace ovigerous
9 LV, shows accentuated sculpture. C, carapace adult d DLV, shows reduced sculpture. D, carapace ovigerous
9 DLV shows accentuated sculpture. E, carapace late subadult d DLV, shows slightly reduced sculpture. F,
carapace late subadult 9 DLV shows retention of sculpture. G, carapace early subadult ¢ DLV, shows moderate
sculpture. H, carapace early subadult $ DLV, shows moderate sculpture.
MEMOIRS OF THE QUEENSLAND MUSEUM
424
BODOTRIIDAE OF MORETON BAY
18080rnm
A62
12mm
MEMOIRS OF THE QUEENSLAND MUSEUM
SP22MWM
Spear SKU § 13mm BP22F ^ SkU
7— sa)
A !
BODOTRIIDAE OF MORETON BAY
ridge; in lateral view posterior transverse ridge
visible only as fine line behind slightly depressed
quadrilateral area; antennal notch a short
groove;antennal tooth subacute, no antennal
ridge; pseudorostral lobes tapering anteriorly and
joining just below ocular lobe, join not visible in
dorsal view; ocular lobe as wide as long, rounded,
and located at anterior extremity of carapace (Fig.
26E-G). Pereion 0.5 times as long as carapace;
first pereionite fully concealed; second pereionite
forms a narrow collar posterior to carapace, as
long as third but shorter than pereionites 4 and 5;
tufts of short setae on posterodorsal margins of
last 3 pereionites (Fig. 26A-D). Pleon robust, no
dorsal ridges; first four pleonites subequal in
length, each with lateral notch in posterior margin
and oblique carina extending anteriorly and
ventrally from notch; fifth pleonite 1.6 times as
long as fourth, with 5 short spine-like setae on
posterodorsal margin (Fig. 26A-D). Telsonic
somite shorter than fifth pleonite and longer than
fourth, with posterodorsal projection (Fig.
26B,D). First antenna 3-segmented with terminal
segmented flagellum; first segment somewhat
geniculate, longer than second and third segments
combined; second segment subequal in length to
third, with fine seta distomedially; third segment
with 2 terminal and 2 subterminal setae; first
segment of flagellum 1.6 times as long as second,
which bears 2 aesthetascs and 2 fine setae distally
(Fig. 29A). All pereiopods with terminal spine-
like setae longer than dactylus; first and third to
fifth pereiopods 7-segmented, second 6-seg-
mented since ischium is fused with basis.
Pereiopod 1 with carpus reaching beyond level of
antennal tooth; length of basis 1.1 times rest of
appendage, distal margin not produced, with
plumose distomedial seta and numerous small
lateral setae; ischium 0.9 times length of merus,
with plumose distomedial seta; merus 0.6 times
as long as carpus, with small distolateral projec-
tion; carpus subequal in length to propodus which
is 1.2 times-length of dactylus, with group of 5
distomedial setae; dactylus with 2 slender spine-
like setae and | seta terminally, and 5 setae along
medial margin; exopod well-developed, slender
proximal segment with 3-4 short setae dis-
tolaterally and 8 short distal segments, each with
2 long setae (Fig. 29B). Pereiopod 2 with fused
427
FIG. 29. Cyclaspis ornosculpta sp. nov, holotype adult
G. A, first antenna. B, pereiopod 1. C, pereiopod 2.
D, pereiopod 3. E, pereiopod 4. F, pereiopod 5.
basis and ischium (join barely discernible) slight-
ly longer than combined length of remaining
segments, with 4 plumose setae laterally and 2
plumose setae distomedially; merus 1.8 times
length of carpus, with plumose seta distomedial-
ly; carpus 1.2 times length of propodus, with 2
strong spine-like setae and rounded process dis-
tally; dactylus 2.3 times length of propodus, with
1 long and 2 short spine-like setae distally, each
with a subterminal seta (Fig. 29C). Pereiopods
3-5 with merus longer than ischium, propodus
longer than dactylus and carpus longer than com-
bined length of propodus and dactylus; basis with
4-6 plumose setae medially; ischium with 3 setae
distomedially; merus with seta distomedially;
carpus with 3 setae laterally and 2 spine-like setae
distolaterally; propodus with spine-like seta dis-
FIG. 28. Cyclaspisornosculpta sp. nov. A, juvenile d LV,. B, juvenile 9 LV,. Sexes indistinguishable on external
morphology. C, early subadult d LV. D, early subadult ? LV. Pleopods developing in d, transverse ridges
developing in 9. E, Late subadult d LV. F, Late subadult 9 LV, Posterior transverse ridge higher in 9. pleon
slender in 2, robust in d. G, d LV. H, Ovigerous 9 LV. Pleopods and second antennae fully developed in d,
marsupium developed in 9. sculpture of carapace reduced in d , accentuated in 9.
FIG. 30. Cyclaspis ornasculpta sp. nov. A-B, holotype
d. A, uropods and lelsonic somite, DV. B, rami of
uropod, DV. C-D, allotype ovig. 9 . C, uropods and
telsonic somite, DV. B, rami of uropod, DV.
tolaterally; dactylus with terminal spine-like seta
and 1-2 terminal setae; some spine-like setae
have rows of fine spinules distally (Fig. 29D-F).
Pereiopod 3 with basis subequal in length to
remaining segments combined; ischium 0.6 times
length of merus; merus 0.9 times length of carpus,
with distomedial seta; propodus 1.3 times length
of dactylus (Fig. 29D). Pereiopod 4 with basis 0.8
times length of remaining segments combined;
ischium 0.4 times length of merus which is 0.9
times length of carpus; propodus 1.3 times length
of dactylus (Fig. 29E). Pereiopod 5 with basis 0.7
times length of remaining segments combined;
ischium 0.5 times length of merus which is 0.9
times length of carpus; propodus 1.5 times length
of dactylus (Fig. 29F). Peduncle of uropod sube-
qual in length to telsonic somite, with 15-16
plumose setae on inner margin; endopod 0.8
MEMOIRS OF THE QUEENSLAND MUSEUM
times length of peduncle and 0.95 times length of
exopod, with 12 plumose setae and 6-7 short
spine-like setae on inner margin, apex pointed,
with serrate inner edge, outer margin with 3
plumose setae: exopod with 12 plumose setae on
inner margin, 1 long terminal spine-like seta and
2 minute terminal setae, outer margin with row of
fine setac (Figs 26H. 30A.B).
OVIGEROUS FEMALE. Integument calcified. with
reticulate pattern of shallow pitting on raised
arcas of carapace and abdomen (Fig. 27B,H).
Carapace 0.35 S.L. almost as deep as long; in
profile the dorsal margin is raised in regions of
transverse ridges and depressed in postocular
region; median dorsal ridge is strong and visible
along length of carapace, forming rounded
process at posterior extremity: in dorsal view
carapace is widest in region of posterior
transverse ridge; anterior and posterior transverse
ridges and dorso- and inferolateral ridges well
developed: depressed quadrilateral region well
defined by ridges; antennal notch, antennal tooth
and pseudorostral lobes as in d (Figs 27B,D,
28H). Pereion 0.5 times às long às carapace; first
pereionite almost fully concealed; second
perionite with rounded dorsal projection,
pereionites 3-5 with low dorsal profile and taper-
ing posteriorly (Figs 27B, 28H). Pleon robust but
much more slender than in d', no dorsal or lateral
ridges; first 4 pleonites subequal in length, each
with lateral articulation notches; fifth pleonite 1.7
times as long as fourth, with 4 short spine-like
setae on posterodorsal margin (Figs 27B, 28H).
Telsonic somite shorter than fifth pleonite and
longer than fourth, with posterodorsal projection
(Figs 28H, 30C). First antenna as in d. All
pereiopods with terminal spine-like setae longer
than dactylus; first and third to fifth pereiopods
7-segmented, second 6-segmented as in d . Seg-
mentation of pereiopods same as in d ; carpus of
pereiopods 3-5 with 2 rather than 3 setae Jaterally.
Peduncle of uropod subequal in length to telsonic
somite, with 9-10 plumose setac on inner margin;
endopod 0.8 times length of peduncle, 0.95 times
length of exopod, with 2 plumose setae on
proximal 1/2 of inner margin, apex bluntly
pointed, with serrate inner edge, inner and outer
margins with fine scale-like teeth: exopod with
8-10 plumose setae on inner margin, long ter-
minal spinc-like seta and minute terminal seta,
outer margin with scale-like teeth and scattered
fine setae (Fig. 30C,D).
brown, black
Colour. Cream to
BODOTRIIDAE OF MORETON BAY
chromatophores dotted on carapace and ab-
domen.
S.L. Adult 3 4.3-5.0mm. Adult 9 4.3-4, 8mm.
HABITAT AND DISTRIBUTION. Most com-
mon over silt and fine sand but were also present
over medium and coarse sand in summer in 1-9m
of water; 12 sites in Pumicestone Passage, sites
2] and 36. Their occurrence in sledge net samples
was highly seasonal, with peak abundances in
summer.
REMARKS. Cyclaspis ornosculpta exhibits
sexual dimorphism in the adult similar to that in
C. elegans Calman, 1907 which species it
resembles but can be readily distinguished from
by the 2 transverse ridges on the carapace in
dorsal view. While early juvenile stages of d d
and 99 of the new species are alike, late
juveniles and adults are readily separated by the
pleopods in the d and ornate carapace sculpture
in the 2.
Adult ds have well-developed second anten-
nae and 5 pairs of pleopods, typical of d d of the
Bodotriidae (Fig. 26C,D). The heavily sculptured
appearance of the carapace of the adult Ẹ is
absent in the adult ¢ (Fig. 27A-D). The quadri-
lateral area of the adult d is not distinctly
recessed like that of the 9 and the posterior
transverse ridge is barely visible.
Late subadult d d have partially developed
pleopods and reduced sculptunng of the carapace
compared to late subadult ? ? (Fig. 27E,F). The
stocular depression of the dorsa! median ridge
1s more acute in the 9, as is the height of the
posterior transverse ridge. Both features are most
highly developed in the adult 9. Early subadult
d d and 9 9 cannot be distinguished on external
morphology until pleopods begin to develop in
the d (Fig. 27G,H).
Comparison of the developmental stages of
both sexes (Fig. 28) shows the gradual develop-
ment of sexual dimorphism. The sex of juveniles
js indistinguishable on external morphology
alone (Fig. 28A,B) but early subadult d d can be
distinguished from 9 9 by having a more robust
abdomen and the appearance of pleopod buds
(Fig. 28C,D). The quadrilateral areas of both
sexes become more developed at this stage. Late
subadult d d exhibit a slight reduction in the
definition of the quadrilateral area and transverse
processes while the reverse occurs in late sub-
adult 9 9 (Fig.28E,F). The most marked changes
occur in the final stage of development when the
carapace of the d elongates and simultaneously
429
FIG. 31. Crane andersoni sp. nov. holotype d. A,
first antenna. B, pereiopod 1. C, pereiopod 2. D,
pereiopod 3, E, pereiopod 4. F, pereiopod 5.
loses. definition of the quadrilateral area and
transverse processes. The 9, on the other hand,
develops calcified posterolateral tubercles and a
posterior tubercle along the median dorsal ridge.
The dorsal margin of the second pereionite also
becomes elevated as a tubercle. The postocular
depression becomes more acute in the 9 and
develops into an angular kink in some post-
ovigerous 9 9. In contrast the post-ocular
depression in the ó remains mildly concave and
may even flatten out slightly. The carapace of the
d, like that of the 9, calcifies on maturity Fig.
28G,H).
Calman (1907) noted that the subadult d of C.
elegans more closely resembles the adult 9 than
it does when full grown. It appears from his
description and figures that development follows
a similar pattern to that of C. ornosculpta.
ETYMOLOGY. Orno-, a contraction of ornate, and the
exsculpia species group ta which it belongs.
430
FIG. 32. Cyclaspis andersoni sp. nov. A-B, holotype
G. A, uropods and telsonic somite, DV. B, rami of
uropod, DV. C-D, allotype ovig. 2. C, uropods and
telsonic somite, DV. B, rami of uropod, DV.
Cyclaspis andersoni sp. nov.
(Figs3 1-35)
MATERIAL EXAMINED. HOLOTYPE
QMW20499, adult d, S.L. 2.9mm, PSM #23,
Pumicestone Passage, site 10, 26°50°S, 153?7'E, J.
Greenwood, 26 Feb 1991, 2m, sand, 34 ppt salinity,
27.2C water temperature. PARATYPES
QMW20500, ovig. 9 , allotype, S.L. 3.5mm, PSM #56,
Horseshoe Bay, 27°30’S, 153?21' E, D. Tafe, 2-3m,
sand, 13 Aug. 1990, 27.2 ppt salinity, 16.5°C water
temperature; QMW20501, adult &, S.L. 2.8mm, SEM
mount, off Dunwich, 27?29'S, 153°22’E, D. Tafe, 4m,
sand, 17 April 1990, 33 ppt salinity, 26.5°C water
MEMOIRS OF THE QUEENSLAND MUSEUM
temperature; QMW20502, ovig. 9, S.L. 2.3mm, SEM
mount, same data as above; QMW20503, adult d , S.L.
2.85mm, in 70% ethanol, Pumicestone Passage, site 10,
26^50'S, 153?7' E, J. Greenwood, 25 Jan 1990, 2m,
sand, 34.9 ppt salinity, 27.3°C water temperature;
QMW20504, ovig. 9, S.L. 2.4mm, in 70% ethanol,
same data as above.
DESCRIPTION. MALE.Integument thin, lightly
calcified, with small, even reticulate patterning
(Fig. 33A-E). Carapace 0.33 S.L. with mild
median dorsal ridge on anterior 1/2 and median
dorsal recess on posterior 1/2; curvature of
carapace is smooth, without lateral ridges, but
with middorsal depressions either side of the
median dorsal ridge; carapace 0.65 as wide as
long, lateral margins evenly curved in dorsal
view; antennal notch a short, shallow groove;
antennal tooth subacute, no antennal ridge; pseu-
dorostral lobes wide, joining just anterior to
ocular lobe which is as wide as long, rounded,
with five lenses (Fig. 33A- E,H). Pereion 0.45 as
long as carapace; first pereionite partially con-
cealed by second, which forms a narrow collar
posterior to carapace, shorter than each of
remaining 3 pereionites which are overlapping,
with low dorsal profile; tuft of very short setae on
posterodorsal margin of third pereionite (Fig.
33B,D). Pleon robust, no dorsal or lateral ridges;
first 4 pleonites subequal in length with small
lateral articulation notches; fifth pleonite 1.5
times as long as fourth (Fig. 33A,B). Telsonic
somite shorter than fifth pleonite and subequal in
length to fourth, with posterodorsal projection
and shallow dorsal notch (Fig. 33A,F). First an-
tenna 3-segmented with terminal segmented
flagellum; segment 1 geniculate, longer than seg-
ments 2+3; segment 2 longer than 3, with fine seta
distomedially; segment 2 with 2 thick and 2 thin
setae distolaterally, and 1 fine seta distomedially;
segment 1 of flagellum slender and 4 times as
long as second which has 2 aesthetascs and 2 fine
setae distally (Fig. 31A). All pereiopods 7-seg-
mented, with terminal spine-like setae longer
than dactylus. Pereiopod 1 with carpus reaching
beyond level of antennal tooth; length of basis 1.5
times rest of appendage, rounded tooth and
plumose seta on distal margin; ischium 0.3 times
FIG. 33. A-H Cyclaspis andersoni sp. nov. 8. A, B, whole mount LV, shows relative lengths of carapace and
somites. C, carapace LV, shows mid-dorsal indentations either side of median dorsal ridge. D, carapace DV,
shows ocular lobe, collar-like second pereionite and maximum width of carapace in mid-region. E, carapace
DLV, shows median dorsal ridge on anterior half and median dorsal recess on posterior 1/2 of carapace. F,
uropods DV, shows posterodorsal projection and shallow dorsal notch on telsonic somite. G, uropods DV,
shows relative lengths of peduncle and rami. H, anterior carapace DLV, detail of antennal notch and ocular
lobe. I, Cyclaspis alveosculpta sp. nov. d DV, shows aperture in dorsum of carapace.
BODOTRIIDAE OF MORETON BAY
431
432
length of merus which is subequal in length to
carpus; propodus 1.4 times length of dactylus,
with 1 medial and 2 distal setae; dactylus with 2
slender terminal spine-like setae, 1 longer than
itself, and 2 stout subterminal setae (Fig. 31B).
Pereiopod 2 with division between basal segment
and ischium barely visible; basis 0.9 times length
of remaining segments combined, with small dis-
tomedial seta; ischium 0.3 times length of merus,
with plumose seta distomedially; merus 1.7 times
length of carpus, with plumose seta distomedially
and stout spine-like seta distolaterally; carpus 1.2
times length of propodus, with spine-like seta
distomedially and spine-like seta distolaterally;
propodus 0.65 times length of dactylus, with
small distomedial seta; dactylus slender with 2
terminal spine-like setae, 1 longer than itself, 1
small terminal seta and 1 subterminal spine-like
seta (Fig. 31C). Pereiopods 3-5 with merus longer
than ischium and propodus longer than dactylus;
basis with 2-4 plumose setae medially; ischium
with 3 setae distomedially; merus with seta dis-
tomedially; carpus with 3 spine-like setae dis-
tolaterally; propodus with spine-like seta and
minute seta distally; dactylus with terminal spine-
like seta, terminal seta and subterminal seta; some
spine-like setae have rows of fine spinules distal-
ly (Fig. 31D-F). Pereiopod 3 with basis subequal
in length to remaining segments combined; is-
chium 0.65 times length of merus which is 0.65
times length of carpus; propodus 1.3 times length
of dactylus (Fig. 31D). Pereiopod 5 with basis 0.7
times length of remaining segments combined;
ischium 0.5 times length of merus which is 0.6
times length of carpus; propodus 1.2 times length
of dactylus (Fig. 31E). Pereiopod 5 with basis 0.5
times length of remaining segments combined;
ischium 0.7 times length of merus which is 0.55
times length of carpus; propodus 1.6 times length
of dactylus (Fig. 31F). Peduncle of uropod 1.8
times as long as telsonic somite, lined with 16
plumose setae on inner margin; endopod 0.75
times as long as peduncle and 0.95 times as long
as exopod, with 12-13 spine-like setae on
proximal 2/3 of inner margin and 2-4 fine setae
on outer margin, apex pointed, with 2 minute
subterminal spine-like setae; exopod with 2
MEMOIRS OF THE QUEENSLAND MUSEUM
plumose setae on proximal 1/3 of inner margin
and 2-3 fine setae on outer margin, apex pointed,
with 3 minute subterminal spine-like setae (Fig.
33F,G, 32A,B).
OVIGEROUS FEMALE. Integument lightly cal-
cified and covered with fine reticulate pattern, as
in the d (Fig. 34A-E). Carapace 0.29 S.L. with
strong median dorsal ridge on anterior 1/2 and
dorsal groove on posterior 1/2; middorsal depres-
sions either side of ridge; carapace 0.68 as wide
as long, lateral margins evenly curved in dorsal
view; depth 0.8 times length of carapace in lateral
view; antennal notch and pseudorostral lobes as
in ó (Fig. 34A-E). Pereion 0.44 times as long as
carapace; pereionite 1 almost concealed by
second, both produced ventrally to form the mar-
supium; pereionites 3-5 short, without ridges, and
tapering in dorsal view; fifth with lateral articula-
tion notch (Fig. 34A,B). Pleon robust and sub-
cylindrical, devoid of dorsal or lateral ridges; first
4 pleonites subequal in length with lateral ar-
ticulation notches; fifth pleonite 1.5 times fourth
with rounded lateral process overlapping telsonic
somite (Fig.34A,B). Telsonic somite 0.55 times
fifth pleonite, posterodorsal projection and shal-
low middorsal notch (Fig. 34F-H). First antenna
3-segmented with terminal segmented flagellum,
as in d. Pereiopods 1-5 as in d except ischium
of 3-5 have 4 rather than 3 setae distally. Peduncle
of uropod 1.6 times as long as telsonic somite,
without plumose setae on inner margin; endopod
0.75 times as long as peduncle and 0.95 times as
long as exopod, with 6-7 spine-like setae on
proximal 2/3 of inner margin, apex with fine point
and 2 minute subterminal spine-like setae; exo-
pod with 2 plumose setae on proximal 1/3 of inner
margin, apex with fine point and 2 minute subter-
minal spine-like setae (Fig. 32C,D, 34F-H).
Colour. Cream with black chromatophores
dotted on carapace and abdomen.
S.L. Adult d 2.8-2.9mm. Adult 9 2.3- 3.5mm.
HABITAT AND DISTRIBUTION. Most com-
monly over silt and fine sand but were also over
medium and coarse sand in summer in 1-5mm of
FIG. 34. Cyclaspis andersoni sp. nov. A,B, whole mount, ovigerous 9, LV, shows relative lengths of carapace
and somites. C, whole mounts d (left) and 9 (right), LV, shows relative sizes of both sexes. D, carapace
ovigerous 9 LV, shows relative length and depth. E, carapace ovigerous ? DLV, shows median dorsal ridge
on anterior half and median dorsal recess on posterior 1/2. F, uropods second ovigerous 9 DV, shows
posterodorsal projection and shallow dorsal notch bearing 2 minute apertures on telsonic somite. G, uropods
ovigerous 9 VV, shows relative lengths of peduncle and rami. H, uropods third ovigerous 9 DV, shows
posterodorsal projection and shallow dorsal notch bearing 2 minute apertures on telsonic somite.
BODOTRIIDAE OF MORETON BAY
433
434 MEMOIRS OF THE QUEENSLAND MUSEUM
y 4d
Seu
n.
D
BODOTRIIDAE OF MORETON BAY
cA
436
TABLE 2. Setation of 3 uropods of C andersoni and
C. nitida.
Fig. 33G) (Hale, 19443)
|Pedunclesetation | — 16long | l3long.Sshor |
Exopod setation | 2lmg | 7 long
L- | ameme | ameme |
water; sites 1-3, 5, 6, 8-12, 15, 26, 28, 31, 33, 34,
35, and 37.
REMARKS. Cyclaspis andersoni is related ta C.
nitida Hale, recorded from NSW and WA. The
main differences are in carapace shape and seta-
tion of the uropods. Hale (19442) described the
3 only of C. nitida so only the d of C. andersoni
can be compared. Both sexes of C. andersoni
have middorsal depressions either side of the
midline, behind which the midline is recessed
(Figs 33E, 34E). C. nitida has a thin longitudinal
median dorsal ridge the length of the carapace
and shows no indication of middorsal depressions
(Hale, 1944a:110). The ocular lobe in C. ander-
soni has 5 lenses (Fig. 33D) compared to 11 in C.
nitida. Both species have a fine reticulate pattern-
ing over the carapace, with black pigment spots,
but the new species has a concentration of black
spots on the middorsal region of the carapace and
their setation differs (Table 2), The differences
in the uropods alone distinguish the species. The
uropods of the 9 of C. anderson lack spines
compared to the á (Fig. 34F-H).
Cyclaspis andersoni can be distinguished from
C. cretata Hale and C. strigilis Hale, two related
species found in SE QLD, by the patterning of the
carapace and overall body size. C. andersoni is
much smaller than C. cretata (Fig. 35A-D) and
C. strigilis, The carapace has à more granular
surface texture than C. crelata (Fig. 35E-H)
which species has a squamose reticulate pattern-
MEMOIRS OF THE QUEENSLAND MUSEUM
ing on the posterodorsal region (Fig. 35E,F).
while C. srrigilis has numerous oblique striae on
the dorsal and lateral surfaces. C. andersoni dif-
fers from both of the latter species in having
relatively short uropodal rami (Fig 35C,D).
Cyelaspis andersoni was the second most abun-
dant cumacean species taken by sledge-pet at
sites 10-12.
SEM photographs of cumaceans vary in quality
depending on the fragility of the specimen (de-
gree of caleification), the state of preservation
and the method used for SEM preparation.
Specimens shown in figs 33 and 34 were freshly
caught and transferred live to liquid nitrogen,
using the freeze-suhstitution method (Tafe,
1995). Specimens in Fig. 35 all suffered some
degree of shrinkage during critical point drying.
ETYMOLOGY. For Prof Don Anderson, Sydney
University,
Cyclaspis alveosculpta sp, nov.
(Figs 17A-E. 331, 36-39, 70E-G)
MATERIAL EXAMINED, HOLOTYPE
OMW 20505, adult d, S.L. 7.5mm, PSM #61, Middle
Banks, 27"12'S, 153^18' E, S. Cook, Sept. 1972, 8m,
sand, 35 p.p.t. salinity, 23^C water temperature,
PARATYPES QMW8536, ovig. 9, allotype, S.L.
7.2mm, same data as holotype; QMW20506, ovig, 2,
S.L. 6.8mm, SEM mount, Horseshoe Bay, site 31,
27°30°S, 15372 1E, D. Tafe, 17 April 1990, 3 m, sand,
33 TAE salinity, 25"C wnter temperature;
OMW30507, subadult d, S.L. 6mm, SEM mount,
same data as above; QMW20508, ovig. 9, S.L; 7mm,
PSM #11, Raby Bay, 27°30'S, 153" | 8'E, D. Tafe, 9
Nov. 1989, 5 m, sandy mud, 35 p.p.t. salinity, 25°C
waler temperature, QM W20509, adult d, S.L. 7.8mm,
PSM #66, same dala as holotype; QMW20510, 2 adults
3d, SL. 7.4, 7,5mm, PSM #60, off Goat Island,
27°31'S, 153°22'E, D. Tafe, 17 April 1990, from gnis
of Apogonidae; QMW20511, subadult d , S.L. 7.2mm,
PSM #59, Raby Bay, 27°30°S, 15318 E, D. Tafe, 22
July 1989.
(captions for ji gures on previous (wo pages)
FIG. 35. A, VLV, left to right, andersoni d , eretatad , undersoni 9 , ererata F. B, DLV, left to right, andersoni
d, cretatad , andersoni 9, eretata V . C, C, eretata d (lop); anderseni d (bottom) LV. D, C. eretata 9 (top);
andersoni 9 (bottom) LV. comparison of ovigerous ? ? in lateral view. E,F, C. crerara sp. nov. LY. E, 3
carapace, shows smooth texture of inlegument, F, 9 carapace shows smooth lexture with cretations, G,H, C.
andersoni sp. nov., carapace, LV, shows small, even reticulate patterning of integument, G, d. H, 9
FIG. 36, Cyclaspis alvensculpta sp. nov, A.B, subadult d, LV, shows reduced transverse ridges (cf. 9) and
partially developed pleopods. C, uropods of subadult d VV, shows relative lengths of peduncle and rami, D,
carapace of subadult ¢ LV, shows pitting of integument and pasterodorsal lobes. E,F, ovigerous ? , LV, shows
relative lengths of carapace and somites. G, carapace of ovigerous 9 LV, shows relative length and depth of
carapace, and shape of pareat lobes. H, Anterior carapace ALV, shows reticulate pattern of pitting and
anterior position of ocular lobe.
BODOTRIIDAE OF MORETON BAY
DESCRIPTION. MALE.
Integument strongly cal-
cified with pattern of shal-
low pitting (Figs 17A-E,
37). Carapace 0.3 S.L.,
1.75 times as long as deep,
without defined anterior
and posterior transverse
ridges though both regions
are slightly raised on lateral
surfaces; median dorsal
ridge pronounced
throughout carapace length
and smoothly convex in
profile, with slight pos-
tocular depression; anten-
nal notch a short groove;
antennal tooth subacute, no
antennal ridge; pseudo-
rostral lobes tapering
anteriorly and joining just
below ocular lobe, join not
visible in dorsal view;
ocular lobe at anterior ex-
tremity of carapace;
posterior extremity with
small dorsal lobe, not
raised above line of dorsum
(Fig. 37C). Subadult 6 d
with anterior and posterior
transverse ridges well
defined and median dorsal
ridge produced posteriorly
to form 2 raised, plate-like
lobes (Fig. 17A-D).
Pereion 0.5 as long as
carapace; first pereionite
concealed by second,
which is partially fused
with carapace, dorsal
profile that of carapace;
dorsal profile of remaining
pereionites continuous with pleon, fourth and
fifth with lateral articulation notches (Fig. 37C).
Pleon robust, median dorsal line visible on last
two pleonites; each pleonite with lateral articula-
tion notches, first four pleonites subequal in
length, fifth 1.6 timesas long as fourth (Fig. 37C).
Telsonic somite shorter than pleonite 5 and
longer than 4, with posterodorsal projection (Fig.
37 A,D). First antenna 3-segmented with terminal
segmented flagellum. All pereiopods 7-seg-
mented. Pereiopod 1 with carpus reaching
beyond level of antennal tooth; length of basis
subequal to rest of appendage, with 20-30 stout
437
FIG. 37. Cyclaspis alveosculpta sp. nov. 6. A,B, paratype d (removed from gut
of Leiognathus moretoniensis Ogilby). A, uropods and telsonic somite, DV
(some inner marginal spines and setae missing). B, rami of uropod, DV. C,D,
holotype d. C, LV. D, uropod and telsonic somite, DV.
spine-like setae on proximomedial bulge, 2
plumose distal setae and numerous fine lateral
setae; ischium 0.8 times length of merus; merus
0.5 times as long as carpus; carpus 0.9 times
length of propodus, with 2 fine medial setae;
propodus with 2 distomedial and 4 short medial
setae; dactylus 0.85 times length of propodus,
with 2 slender spine-like setae and 1 fine seta
terminally, 1 spine-like seta and 1 fine seta sub-
terminally and 5 setae along medial margin; ex-
opod well-developed, enlarged proximal
segment with 3-4 short setae distolaterally and 8
short distal segments, each with 2 long setae (Fig.
38A). Second pereiopod with basis 0.75 times
438
FIG. 38. Cyclaspis alveosculpta sp. nov. holotype d,
A, pereiopod 1. B, pereiopod 2. C, pereiopod 3. D,
pereiopod 4. E, pereiopod 5.
combined length of remaining segments, with 4
plumose setae laterally and 1 distolaterally; is-
chium 0.3 times length of merus, with 3 setae
distomedially; merus as long as combined length
of carpus and propodus, with 1 plumose seta
distomedially and 1 distolaterally; carpus twice
as long as propodus, with 2 strong spine-like setae
and rounded process distally; dactylus 3 times
length of propodus, with 2 terminal spine-like
setae (one at least 1.7 times longer than itself),
one subterminal spine-like seta and one subter-
minal seta (Fig. 38B). Pereiopods 3-5 with merus
longer than ischium, carpus longer than merus,
propodus longer than dactylus and terminal
spine-like seta longer than dactylus; basis with
4-6 plumose setae medially; ischium with 4 setae
distomedially; merus with seta distomedially;
carpus with 2-3 setae laterally and 3 spine- like
setae distolaterally; propodus with spine-like seta
and minute seta distally; dactylus with terminal
spine-like seta longer than itself, terminal seta
MEMOIRS OF THE QUEENSLAND MUSEUM
and subterminal seta; some spine-like setae have
rows of fine spinules distally (Fig. 38C- E).
Pereiopod 3 with basis 0.8 times length of
remaining segments combined; ischium 0.4 times
length of merus which is 0.8 times length of
carpus; propodus 1.2 times length of dactylus
(Fig. 38C). Pereiopod 4 with basis 0.7 times
length of remaining segments combined; ischium
0.35 times length of merus which is 0.8 times
length of carpus; propodus 1.2 times length of
dactylus (Fig. 38D). Pereiopod 5 with basis 0.65
times length of remaining segments combined;
ischium 0.35 times length of merus which is 0.9
times length of carpus; propodus 1.2 times length
of dactylus (Fig. 38E). Peduncle of uropod 0.8
times length of telsonic somite, with plumose
setae lining whole inner margin (15-22 setae in
fully mature adult); endopod 1.2 times length of
peduncle, subequal to length of exopod, with
17-30 plumose setae (2 rows) and 10-15 short
spine-like setae on inner margin, apex channelled
and slightly curved inwards, with subterminal
constriction; exopod with 12-16 plumose setae on
inner margin, apex channelled and slightly
curved inwards, with subterminal constriction.
Fine scattered setae on telsonic somite, peduncle
and rami (Fig. 37). Subadult d uropod with
shorter peduncle with fewer setae and endopod
with shorter setae and spine-like setae (Fig. 36C,
39A).
OVIGEROUS FEMALE. Integument strongly cal-
cified with pattern of shallow pitting, as in d (Fig.
36E-H). Carapace length 0.32 S.L. as long as
deep including marsupium, with strong anterior
and posterior transverse ridges; in profile median
dorsal ridge is slightly convex with postocular
depression and raised areas in regions of
transverse ridges; median dorsal ridge is
produced posteriorly to form notched, plate-like
lobe; antennal notch a short groove; antennal
tooth subacute; pseudorostral lobes tapering
anteriorly and joining just below ocular lobe, join
not visible in dorsal view; ocular lobe at anterior
extremity of carapace (Fig. 36E-H). Pereion 0.5
times as long as carapace. First pereionite a nar-
row band, visible in lateral view; pereionites 1
and 2 produced ventrally to form the marsupium,
second also produced dorsally to form plate-like
lobe; pereionites 3 and 4 with lateral overlapping
lobes; fifth with lateral articulation notches and
well-developed dorsolateral carinae (Fig. 36E-
H). Pleon robust, all 5 pleonites with dorsolateral
carinae and lateral articulation notches; first 4
pleonites and telsonic somite subequal in length,
BODOTRIIDAE OF MORETON BAY
fifth pleonite 1.6 times as long as fourth (Fig.
36F). Telsonic somite projecting posteriorly over
bases of uropods (Figs 36F, 39D). First antenna
3-segmented with terminal segmented flagellum,
as in d. Pereiopods as in d except: basis of first
pereiopod with only | stout spine-like seta on
reduced medial bulge, basis of exopod also has
reduced bulge compared to d ; carpus of second
pereiopod has only 1 strong spine-like seta distal-
ly; ischium of pereiopods 3-5 with only 3 distal
setae, carpus with only 1-2 lateral setae (Fig.
39B,C). Peduncle of uropod 0.7 times as long as
telsonic somite, with 6-7 plumose setae on inner
margin; endopod 1.4 times as long as peduncle,
subequal in length to exopod, with row of minute
scales and spine-like setae on innér margin, apex
channelled with subterminal constriction; ex-
opod with 8-9 plumose setae on inner margin,
apex channelled with subterminal constriction
(Fig. 39D).
Colour. Cream to fawn.
S.L. Adult d 7.2-7.8mm. Adult 2 7.0-7.5mm.
HABITAT AND DISTRIBUTION. Most com-
monly over medium and coarse sand in 1-5m of
water; sites 11, 12, 15, 26, 30, 31, 32 and 36.
REMARKS. Cyclaspis alveosculpta closely
resembles C. usitata Hale, from NSW and S
AUST. The most obvious differences relate to the
ridges and tubercles of the carapace. Also the
dactylus of pereiopods 1 and 2 are relatively
longer in C. alveosculpta and the d has many
more spine-like setae on the basis of the first
pereiopod, Hale (1932:550; 1944a:123) only
described and figured the 9 of C. usitata though
he later suggested (Hale, 1948:40) it to be the 2
of C. mjobergi, described by Zimmer (1921) from
ód only. It is obvious that the posterior
transverse ridge and posterior dorsomedial
tubercle of the adult 9 of C. usitata are poorly
developed compared those of C. alveosculpta.
The posterior median dorsal ridge of the carapace
is raised in the adult ĝ of C. mjobergi but not in
C. alveosculpta. The dactylus of pereiopod 1 is
less than 2/3 as long as the propodns in C. usitata
and C. mjobergi but at least 2/3 as long in C.
alveosculpia.
Cyclaspis alveosculpta superficially resembles
C. munda Hale from southern NSW, but differs
in having spine-like setae on the basis of
pereiopod | and a very long terminal spine-like
sela on pereiopod 2. Roccatagliata (1989)
described C. sculptilis from Brazil, which
resembles C. alveosculpta , but the location and
439
D
RT 3
c f
\ f | < x
FIG. 39, Cyclaspis alveasculpta sp. nov. A, uropods
and telsonic somite of subadult g , DV. B-D, paratype
ovig. 9 . B, pereiopod 1. C, pereiopod 2. D, uropods
and telsonic somite, DV.
definition of the posterior transverse ridges on the
carapace are different.
d and 9 of C. alveosculpta were recorded in
the same area, both strongly calcified with reticu-
late pitting of the carapace. The carapace of the
subadult d' has similar sculpture to the adult %
(Fig. 70E-G), but loses it during the last stage of
development (cf. C. ornosculpta, Fig. 27). All
adults and subadults have 2 dorsal depressions,
behind the anterior transverse ridge of thc
carapace on either side of the median ridge (Figs
331, 70E,G).
8 adult d d of C. alveosculpia were taken from
gut contents of fish (Leiognathus maretoniensis,
Apogonidae) trawled (17 April 1990) between
sites 30 and 31, depth 5-15 m, on sand (S.L. fish
2-7cm), Some setae and spine- like setae have
been broken off the uropodal rami (Fig. 37A,B)
during the digestive process.
440 MEMOIRS OF THE QUEENSLAND MUSEUM
imm
Mu 2mm
BODOTRIIDAE OF MORETON BAY
ETYMOLOGY. Latin alveus, pil, and exseulpta
species group.
Cyclaspis chaunosculpta sp. nov.
(Figs 40-43)
MATERIAL EXAMINED. HOLOTYPE
QMW320512, adult d, S.L. 10.32mm, PSM #65, Mid-
dle Banks, 27°12°S, 153*18'E, S. Cook, Sept. 1972,
Bm, sand, 35 p.p.t. salinity, 23°C water temperature.
PARATYPES QMW20513, ovig. 2, allotype, S.L.
9.5mm, PSM #63, sume data as holotype; OMW 20514,
subadult d , S.L. 5.2mm, SEM mount, Horseshoe Bay,
Site 31, 27°30" S. 153721 E, D. Tale, 17 April 1990, 2-3
m, sand, 33 p.p.i. salinity, 25"C water temperature.
QMW8536, 3 adult 9. 9, S.L, 9,0-9.7mm, same data
iis holotype.
DESCRIPTION. MALE. Integument strongly cal-
cified with pattern of pitting which resembles the
porous structure of a sponge (Fig. 41 A). Carapace
0.3 S.L. depth 0.55 times length, with poorly
defined anterior and posterior transverse ridges:
profile of median dorsal ridge may be slightly
raised in vicinity of both transverse ridges, with
shallow post-ocular depression and small
posterior lobe; antennal notch a short groove;
untennal tooth subacute, no antennal ridge; pseu-
dorostral lobes tapering anteriorly and joining
just below ocular lobe, join not visible in dorsal
view; ocular lobe at anterior extremity of
carapace (Fig. 41A). Pereion 0.65 as long as
carapace; pereionite | concealed; pereionite 2
without dorsal lobe; pereionites 3-5 with dor-
solateral carinae, tufts of setae on posterodorsal
margins and dorsal profile continuous with that
of pleon; third and fourth with posterolateral
overlapping lobes and fifth with lateral articula-
tion notch (Fig. 41A). Pleon robust; each pleonite
with lateral articulation notches, first 4 pleonites
with dorsolateral carinae and subequal in length,
fifth 1.45 times as long as fourth (Fig. 41A).
Telsonic somite 0.66 times length of fifth pleonite
and subequal to fourth, with swollen posterodor-
sal projection (Fig. 41A). First antenna 3-seg-
mented with terminal segmented flagellum. All
pereiopods 7-segmented. Perciopod 1 with car-
pus reaching beyond level of antennal tooth; basis
44l
FIG. 41. Cyclaspis chaunosculpta sp. nov, type
specimens. A, holotype d, LV. B, allotype ovig. 9,
LV.
L1 times rest of appendage, with 20-25 siout
spine-like setae on pronounced proximomedial
bulge, 2 plumose distal setae and rounded distal
lobe which extends at least 1/3 Jength of ischium:
ischium 0.6 times length of merus, which ts 0.6
times length of carpus; carpus 0.8 times length of
propodus; propodus with 3 distomedial and 5
short medial setae; dactyJus 0.55 times length of
propodus, with 2 slender spine-like setae and |
fine seta terminally, ] spine-like seta and 3 fine
seta subterminally and 3 setae along medial mar-
gin; exopod well developed, proximal segment
with distal bulge bearing 3-4 short plumose setae,
FIG. 40. Cyclaspis chaunosculpta sp. nov. subadult d . A,B, whole mount DV, shows relative lengths of carapace
and somites. C, carapace AL V, shows anterior position of ocular lobe and shape of antennal notch. D, carapace
DV, shows maximum width in region of anterior transverse ndge. integument strongly calcified with pattern
of pitting (resembles porous structure of sponge). E, carapace PLV, shows structure of posterior transverse
ridge. F, carapace and Pereion PLV, shows dorso lateral projections of pereion. G, carapace DLV, shows median
dorsal ridge and anterior and posterior transverse ridges. H, Uropod DV , shows relative lengths of peduncke
and rami.
442
FIG. 42. Cyclaspis chaunosculptà sp. nov., A,B,
holotype adult 3. A, pereiopod 1. B, pereiopod 2.
C,D, allotype, ovig. 9. C, pereiopod 1. D, pereiopod
2.
8 short distal segments, each with two long setae
(Figs 41A, 42A). Pereiopod 2 same as C. alveos-
culpta except ischium has 2 setae distomedially,
dactylus is 2.5 times length of propodus and main
dactylar spine-like seta 1s 1.4 times dactylus (Fig.
42B). Pereiopods 3-5 same as C. alveosculpta
except basis has 3-5 plumose setae medially and
dactylar spine-like seta is 0.9 times length of
dactylus. Uropod same as C. alveosculpta except
peduncle of uropod 0.7 times length of telsonic
somite, with 16- 18 setae lining inner margin,
endopod 1.3 times length of peduncle, with 30-40
plumose setae (2 rows) and 9-11 short spine-like
setae on inner margin, apex finely channelled and
incurved, with subterminal constriction; exopod
with 12-14 plumose setae on inner margin, apex
finely channelled and incurved (Fig. 43A-C),
OVIGEROUS FEMALE. Integument strongly cal-
cified with pattern of deep pitting, as in & (Fig.
41 B). Carapace length 0.3 S.L. as deep as long
including marsupium, with strong anterior and
posterior transverse ridges; in profile median dor-
sal ridge is convex with postocular depression
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG, 43. Cyclaspis chaunosculpta sp. nov. holotype d.
A, uropod and telsonic somite, DLV. B, distal end of
exopod. C, distal end of endopod.
and raised areas in regions of transverse ridges;
median dorsal ridge produced posteriorly to form
rounded plate-like lobe almost as high as median
dorsal hump; antennal notch, antennal tooth,
pseudorostral lobes and ocular lobe as d (Fig.
41B). Pereion 0.5 times as long as carapace;
pereionite J a narrow band, visible in lateral view;
percionites | and 2 produced ventrally to form
marsupium, second also produced dorsally to
form plate-like lobe; pereionites 3-5 with dor-
solateral carinae, tufts of setae on posterodorsal
margins and dorsal profile continuous with that
of pleon; pereionites 3 and 4 with posterolateral
overlapping lobes and fifth with lateral articula-
uon notch (Fig. 41B). Pleon robust, first 4
pleonites subequal in length, with dorsolateral
carinae and lateral articulation notches; fifth
pleonite 1.5 times as long as fourth; telsonic
BODOTRIIDAE OF MORETON BAY
somite projecting posteriorly over bases of
uropods (Fig. 41B). First antenna 3-segmented
with terminal segmented flagellum, as in d.
Pereiopods às in d except: basis of pereiopod 1
without enlarged distal lobe or spine-like setae on
reduced medial bulge; carpus of perciopnd 2 has
l strong and 1 weak spine-like seta distally; is-
chium of pereiopods 3-5 with 3-4 distal setae,
carpus has only 1-2 lateral setae in addition to 3
distal spine-like setae (Fig. 42C,D), Uropod same
as 9 of C alveosculpta, exceptexopod has 11-12
plumose setae on inner margin.
Colour. Cream.
S.L. Adult & 10.3mm. Adult 7 9.0-9.7mm.
HABITAT AND DISTRIBUTION. Most com-
nion over medium and coarse sand in 1-5m of
water, Middle Banks and Horseshoe Bay in
Moreton Bay.
REMARKS. Cyclaspis chaunosculpta closely
resembles C. supersculpra Zimmer, 1921, from
NW Australia. Both species have a reticulate
pattern of deep pits on the carapace and ab-
dominal segments: however, C. supersculpta also
has lateral bulges either side of the median dorsal
ridge at the posterior extremity of the carapace.
There isno sign of such bulges in juvenile or adult
specimens of the new species. Close examination
af the holotype of C. xupersculpta shows that the
posterolateral projection on the fifth pleonite is
only about half as long as shown by Zimmer
(1921, fig. 8). The size of this projection is similar
lo that of the new species. C. chaunosculpta also
has more strongly developed transverse ridges
than C. supersculpra. A comparison of 9?
Juveniles of both species (Fig. 43D,E) shows
these differences in carapace structure. The sub-
adult d of C. chaunosculpta has strong transverse
ridges on the carapace, unlike the adult d. (Fig.
40).
C. chaunosculpta also resembles C. aspera
Hale, which has been recorded off Cofts Harbour,
NSW, The most obvious differences relate to the
absence, in C, ¢haunosculpta, of spinules on
carapace und somites. Also the anterior
transverse ridge is wider than the posterior one in
dorsal view, whereas in C, aspera the posterior
ridge is wider (Hale, 19442a:125, figs 45-46),
C. chaunosculpta can he distinguished from C.
candida and C. mjobergi by the shorter peduncle
of the uropod relative to its rami. It also differs
from the latter species in lacking small tubercles
on the middorsal region of the carapace (Zimmer,
1921),
443
C. chaunosculpta can be distinguished from C
alveoxculpta by the deeper, sponge-like pitting of
the carapace, shorter dactylar spine-like seta of
the second pereropod and larger overall size.
ETYMOLOGY. Greek chaunos, porous, sponge-like
and the exyculpta species group.
Cyclaspis agrenosculpta sp. nov,
(Figs 44-47)
MATERIAL EXAMINED, HOLOTYPE
QMW 20516, adult d , S.L. 10.8mm, PSM #62, Middle
Banks, 27^12'8, 153°18’E, S, Cook, Sept, 1972, 8m,
sand, 35 p,p.t. salinity, 23°C water temperature.
PARATYPES QMW20517. avig. 9 , ullotype, S.L.
9.8mm, PSM #64, same data as holotype; OMW20518,
subaduli 2, S.L, 7.4mm. SEM mount, Horseshoe Bay,
sile 31, 27° 3078, 153° 21°E, D. Tafe, 10 Jan. 1993,
2-3m, sand, 34 ppt salinity, 26°C water temperature.
DESCRIPTION. MALE. Integument strongly cal-
eified with a network of angular pits interspersed
hy thin calcified ndges (Fig. 45A). Carapace 0,29
S.L. 1.95 times as long as deep, with poorly
defined anterior and posterior transverse ridges;
profile of median dorsal ridge almost straight.
with slight postocular depression and slight
posterior hump; antennal notch a short groove:
antennal tooth rounded, no antennal ridge; pseu-
dorostral lobes tapering anteriorly and joining
just below ocular lobe, join not visible in dorsal
view; ocular lobe at anterior extremity of
carapace (Fig. 45A). Pereion 0.6 as long as
carapace, pereionite | fully concealed; pereionite
2 forming a collar behind carapace with dorsal
lobe us high as dorsum of carapace; pereionites
3-5 with dorsolateral carinac, tufts of setae on
posterodorsal margins and dorsal profile con-
tinuous with that of pleon; third and fourth with
posterolateral overlapping Jobes and fifth with
lateral articulation notch and raised dorsum (Ttg.
45A). Pleon very robust and calcified, cach
pleonite with lateral articulation notches and dor-
solateral carinae, first 4 subequal in length, afik
1.6 times as long as fourth (Fig. 45A). Telsomc
somite 0.66 times length of fifth pleonite and
subequal to fourth, wiih small mid-dorsal hump
and posterodorsa) projection (Fig. 45A). First
antenna 3-segmented with terminal segmented
flagellum; first segment somewhat geniculate.
All perciopods 7-segmented. First pereiopod
with carpus reaching beyond level of antennal
tooth; length of basis 1.4 times rest of appendage,
with 17-19 spine-like setae and small protrusion
on pronounced proximomedial bulge, 2 plumose
distal setae and rounded distal lobe extending 1/4
444 MEMOIRS OF THE QUEENSLAND MUSEUM
BODOTRIIDAE OF MORETON BAY
length of ischium; ischium 0.8 times length of
merus, which is 0.55 times length of carpus;
carpus 0.9 times length of propodus; propcdus
with 2 slender spine-like setae and 2 setae dis-
tomedially; dactylus 0.65 times length of
propodus, with 2 slender spine-like setae and |
fine seta terminally, 1 spine-like seta and 3 fine
seta subterminally and | seta along medial mar-
gin; exopod well-developed, proximal segment
with distal bulge bearing 5-6 short plumose setae,
8 short distal segments, each with 2 long setae
(Fig. 46A). Pereiopod 2 same as C, alveosculpta
except ischium has 2 distal setae, smaller of the
two carpal spine-like setae reaches distal end of
dactylus, dactylus is 2.3 times length of propndue
and main dactylar spine-like seta is |.] times
dactylus (Fig. 46B), Pereiopods 3-5 same as C.
alveosculpra except basis has 3-7 plumose setae
medially, carpus has 1 lateral seta in addition to
3 distal spine-like setae and dactylar spine-like
seta is 0.75- 0.85 times length of dactylus.
Peduncle of uropod 1.2 times length of telsonic
somite (Fig. 45A), with plumose setae lining
whole inner margin (26-30 setae in fully mature
adult); endopod 0,9 times length of peduncle or
exopod, with 25-30 plumose setae (2 rows) and
18 short spine-like setae on inner margin, apex
bluntly pointed with subterminal constriction; ex-
opod with 18-20 plumose setae on inner margin,
apex channelled, slightly curved inwards (Fig.
47A).
OVIGEROUS FEMALE. Integument strongly cal-
cified with a network of angular pits interspersed
by thin calcified ridges, as in d (Fig. 45B).
Carapace length 0.3 S.L. almost as deep as long
including marsupium, with strong anterior and
posterior transverse ridges; in profile median dor-
sal ridge is slightly concave between transverse
ridges and slightly convex behind posterior
transverse ridge, with postocular depression and
small raised lobe at posterior extremity; antennal
notch a short groove; antenna! tooth subacute, no
antennal ridge; pscudorostral lobes and ocular
lobe as in ġ (Fig. 45B). Pereion 0.6 times as long
as carapace; pereionite | a narrow band, visible
in lateral view; pereionites 1 and 2 produced
445
is
^
ESTNE
LL finn
FIG, 45. Cyclaspis agrenosculpra sp, nov, types. A,
holotype d, LV. B, allotype ovig. ¥, LV.
ventrally to form the marsupium, second also
produced dorsally to form plate-like lobe almost
as high as preceding lobe; fifth with lateral as-
ticulation notches and welldeveloped dor-
solateral carinae (Fig. 45B). Pleon heavily
calcified but slender. all 5 pleonites with dor-
solateral carinae; first 4 pleonites subequal in
Jength, with lateral articulation notches; fifth
pleonite 1.6 times as long as fourth (Fig. 45B).
Telsonic somite 0.66 times length of pleonite 5
and subequal to fourth, with small middorsal
hump and posterodorsal projection, as in d. (Fig,
45B). First antenna 3-segmented with terminal
FIG. 44. Cyclaspis agrenosculpta sp. nov. subadult 8 . A,B, whole mount LV, shows relative lengths of carapace
nnd somites, C, carapace DLV, shows structure of anterior and posterior transverse ridges. aperture located
behind anterior ridge. D, carapace PLV, shows recessed median dorsal ridge between posterior transverse ridge
and posterior of carapace. E, Structure of carapace DV, shows dorsal aperture and network of calcified pits
interspersed by thin chitinised ridges. F, carapace ADV, shows ocular lobe at anterior extremity and profile of
posterior transverse ridge, G; Detail of dorsal aperture DV, shows aperture lined with numerous plate-like lobes.
H, urepods VV, shows relative lengths of peduncle and rimi.
446
FIG. 46. Cyclaspis agrenosculpta sp. nov., A,B,
holotype d. A, pereiopod 1. B, pereiopod 2. C-D,
allotype ovig. 2. C, pereiopod 1. D, pereiopod 2.
segmented flagellum, as in d . Pereiopods as in d
except: basis of first pereiopod without enlarged
distal lobe or spine-like setae on reduced medial
bulge, though protrusion is present on bulge;
carpus of pereiopod 2 has 1 strong and 1 weak
spine-like seta distally; pereiopods 3-5 with basis
with 4-8 medial setae, carpus with 2-3 lateral
setae and dactylar spine-like seta 0.83-0.93 times
length of dactylus (Fig. 46C,D). Peduncle of
uropod 1.1 times as long as telsonic somite, with
7-9 plumose setae on inner margin; endopod sub-
equal in length to peduncle and 0.9 times length
of exopod, with a single spine-like seta on middle
of inner margin, apex bluntly pointed; exopod
with 16-17 plumose setae on inner margin, apex
channelled and slightly curved inwards (Fig.
47B).
Colour. Cream.
S.L. Adult d 10.8mm. Adult 9 9.8mm.
HABITAT AND DISTRIBUTION. Most com-
mon over medium and coarse sand in 1-10m of
water; from Middle Banks and Horseshoe Bay,
Moreton Bay; uncommon on the western but
common on the eastern side of Moreton Bay.
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 47. Cyclaspis agrenosculpta sp. nov., A, uropod
and telsonic somite of holotype d , DV, and distal end
of endopod. B, uropod and telsonic somite of allotype
ovig. 9, DV, and distal ends of both rami.
REMARKS. C. agrenosculpta resembles C.
australis Sars (1887:12, pl. 1, figs 1-20) from VIC
and C. tribulis Hale (1928:34, figs 3,4) and C.
mawsonae Hale (1944a:119) from S AUST.
However, the new species is distinguished from
all3 by the pattern of deep, rectangular pits on the
carapace. Stephenson et al. (1978:208, 1980:259;
Bacescu, 1988:69 recorded C. agrenosculpta
9 9 as C. tribulis. amd d d as C. mawsonae
(Stephenson et al., 1978:210; Bacescu, 1988:61).
C. tribulis and C. mawsonae are not known in
QLD contrary to Bacescu (1988). Two depres-
sions are located in the dorsal surface of the
carapace of C. agrenosculpta, roughly in the
same positions as for C. alveosculpta (Fig. 44C-
G). The subadult 9 of C. agrenosculpta closely
resembles the adult except for the transverse
ridges of the carapace, which are not as well
BODOTRIIDAE OF MORETON BAY
developed (Fig. 44A,B). The
uropod of the subadult 9 is
basically the same as in the
adult except for the slightly
shorter peduncle and smaller
inner marginal spine- like seta
of the endopod.
Cyclaspis agrenosculpta
most closely resembles C.
chaunosculpta sp. nov., taken
from the same area. They can
be readily distinguished by
the sculpture pattern of the
carapace and the relative
peduncle length of the
uropod.
Cyclaspis agrenosculpra
can be distinguished from C.
candida and C. mjobergi by
the height and shape of the
second pereionite. It also dif-
fers from the latter species in
lacking small tubercles on the
mid-dorsal region of the
carapace (Zimmer, 1921).
ETYMOLOGY. Greek agrenon,
net, and the exscu/pla species
group.
Cyclaspis daviei sp. nov.
(Figs 48-50)
MATERIAL EXAMINED.
HOLOTYPE QMW20521, ovig.
9. S.L. 3.0mm, PSM #19, Horse-
shoe Bay, site 31, 27°30°S,
153°21°E, D. Tafe, 4 Feb. 1993,
2-3m, sand, 34 p.p.r. salinity,
26°C water temperature.
QMW20522, ovig. 9, S.L. 2.6mm, in 70% ethanol,
same data as above; QMW 20523, ovig. ?, S.L. 2.5mm,
in 7096 ethanol, same data as above,
DESCRIPTION. OviGEROUS FEMALE. Integu-
ment thin, lightly calcified, with small, even
reticulate patterning (Fig. 48A). Carapace 0,33
S.L. with mild median dorsal ridge on anterior 1/2
and median dorsal recess on posterior 1/2; curva-
ture of carapace is smooth, without lateral ridges;
carapace 0.65 as wide as long, lateral margins
slightly rounded in dorsal view. Antennal notch
a short, shallow groove; antennal tooth subacute,
no antennal ridge, Pseudorostral lobes wide, join-
ing just anterior 1o ocular lobe which is as wide
as long, rounded, with 11] lenses (Fig. 48A).
Pereion 0.5 times length of carapace; pereionite
447
FIG, 48. A,B, Cyclaspis daviei sp. nov. paratype ovig. 9. A, LV. B, first
antenna. C, Cyclaspis andersoni sp. nov. paratype ovig. 9, LV.
1 fully concealed by second, which forms a nar-
row collar posterior ta carapace; shorterthan each
of remaining 3 pereionites which are overlapping
(Fig. 48A). Pleon robust, no dorsal or lateral
ridges; first 4 pleonites subequal in length, fifth
pleonite 1.5 times as long as fourth (Fig. 48A).
Telsonic somite shorter than fourth pleonite, with
posterodorsal projection. First antenna 3-seg-
mented with terminal segmented flagellum, first
segment somewhat geniculate, longer than
second and third segments combined; second
segment longer than third, with 3 small setae
distomedially; third segment with 2 small setae
distolaterally and 1 fine seta distomedially; first
segment of flagellum 2.5 times as long as second,
which bears 2 aesthetascs and 2 fine setae distally
(Fig. 48B). All pereiopods 7-segmented, with
448
FIG. 49, Cyclaspis daviei sp. nov. holotype 9. A,
pereiopod 1. B, pereiopod 2. C, pereiopod 3. D,
pereiopod 4. E, pereiopod 5.
terminal spine-like seta longer than dactylus, ex-
cept for first pereiopod (Fig. 49). Pereiopod 1
with carpus reaching beyond level of antennal
tooth; length of basis 0.8 times rest of appendage:
ischium 0.6 times length of merus which is 0.7
times length of carpus and with small distal
process; carpus 0,7 times length of propodus
which has 2 small distal setae; dactylus 0.7 times
length of propodus with 2 slender terminal spine-
like setae, 1 almost as long as itself, and 2 ter-
minal setae (Fig. 49A). Pereiopod 2 with basis
subequal in length to that of remaining segments
combined, with rows of small setae along medial
and distal margins; ischium 0,4 times length of
merus, with plumose seta distomedially; merus
1.4 times length of carpus, with plumose seta
distomedially and spine-like seta distolaterally;
carpus 1.3 times length of propodus, with spine-
like seta distomedially and spine-like seta dis-
tolaterally; propodus 0.5 times length of dactylus
which is slender with 2 terminal spine-like setae,
the longer one 1.4 times longer than itself, 2 small
MEMOIRS OF THE QUEENSLAND MUSEUM
terminal seta and 1 subterminal spine-like seta
(Fig. 49B). Pereiopods 3-5 with merus twice as
long as ischium and propodus at least 1.6 times
as long as dactylus; basis with seta distomedially;
ischium with 2 setae distomedially; merus with
seta distomedially; carpus with 1-2 spine-like
setae distolaterally, small seta distomedially and
small seta proximomedially; propodus with
spine-like seta and minute seta distally; dactylus
with terminal spine-like seta, terminal seta and
subterminal seta (Fig. 49C-E). Pereiopod 3 with
basis 0.95 times length of remaining segments
combined; ischium 0.5 times length of merus
which is 0.75 times length of carpus; propodus
1.6 times length of dactylus (Fig. 49C), Pereiopod
4 with basis 0.7 times length of remaining seg-
ments combined; ischium 0.5 times length of
merus which is 0.9 times length of carpus;
propodus 1.7 times length of dactylus (Fig. 49D).
Pereiopod 5 with basis 0.6 times length of
remaining segments combined; ischium 0.5 umes
length of merus which is 0.7 times length of
carpus; propodus 1.6 times length of dactylus
(Fig. 49E), Peduncle of uropod 1.2 times length
of telsonic somite, without plumose setae on
inner margin; endopod 1.1 times as long as
FIG. 50. Cyclaspis daviei sp. nov. holotype 9. A,
uropods and telsonie somite, DV. B, rami of uropod,
DV.
BODOTRIIDAE OF MORETON BAY
peduncle and 0.95 times as
long as exopod, with 10-11
short spine-like setae on
proximal 2/3 of inner margin
and 4-5 short setae on dorsal
surface, apex pointed, with 2
minute subterminal spine-like
setae; exopod with 5 plumose
setae on proximal 1/2 of inner
margin and 3-4 short setae on
dorsal surface, apex pointed,
with 2 minute subterminal
spine-like setae (Fig. 50).
Colour, White, translucent
with many black
chromatophores on carapace
and abdomen.
S.L. Adult 9 3.0mm.
HABITAT AND DISTRIBU-
TION. Most common over
fine sand in 1-5m of water;
from sites 12 and 31 in
Moreton Bay.
REMARKS. C. daviei most
closely resembles C. ander-
soni but can easily be distin-
guished by the peduncle of the
uropod, which is shorter than
the rami. Also the dactylus on
pereiopod 2 is longer, the seta-
tion of the ischium of
pereiopods 3-5 is different, as
is the setation of the uropodal
rami, and there are no indenta-
tions on the middorsal region
of the carapace,
ETYMOLOGY. For Peter Davie,
Queensland Museum.
449
FIG. 51. Cyclaspis sallai sp. nov, holotype d. A, first antenna. B, pereiopod
Cyclaspis sallai sp. nov.
(Figs 51-53)
MATERIAL EXAMINED. HOLOTYPE
QMW20524, adult d, S.L. 5.2mm, PSM #49,
Pumicestone Passage, site 12, 26°49°S, 153°8’E, J.
Greenwood, 24 April 1990, 2m, coarse sand, 30.8 ppt
salinity, 18.7°C water temperature. PARATYPE
QMW20525, ovig. ? , allotype, S.L. 4.5mm, PSM #50,
same data as holotype,
DESCRIPTION. MALE. Integument smooth and
calcified, with minute scattered spine-like setae
|, C, pereiopod 2. D, pereiopod 3. E, pereiopod 4. F, pereiopod 5.
on surface. Carapace resembles that of Cyclaspis
cooki except ocular lobe has 10 lenses, 4 central
and 6 outer. Pereion 0.5 same as in C. cooki. Pleon
robust and same as in C. cooki except fifth
pleonite 1.4 times as long as fourth. First antenna
3-segmented with terminal segmented flagellum;
first segment somewhat geniculate, longer than
second and third segments combined, with rows
of fine setae on anterior (medial) surface; second
segment 1.2 times third segment, with 4 fine setae
distomedially and 2 setae distolaterally; third seg-
450
FIG. 52. Cyclaspis sallai sp. nov. holotype d. A,
uropods and lelsonic somile, DV. B, rami of uropod,
DV.
ment with 3 slender setae distomedially and 2
distolaterally; first segment of flagellum twice as
long as second, which has two aesthetascs and 2
fine setae distally (Fig. 51A). All pereiopods 7-
segmented, with terminal spine-like setae at least
as long as dactylus; some spine-like setae have
rows of fine spinules distally (Fig. 51B-F).
Pereiopod | with basis I.I times length of
remaining segments combined, with simple seta
distomedially, plumose seta disto-laterally, and
clump of 8 stout spine-like setae on bulge of
medial region; ischium 0.65 times length of
merus which is 0.5 times length of carpus; carpus
0.8 times length of propodus, which is 1.7 times
length of dactylus, with ] medial and 2 distal
setae; dactylus with 2 slender terminal spine-like
setae, one as long as itself, 2 terminal setae and
stout subterminal seta; exopod well developed,
MEMOIRS OF THE QUEENSLAND MUSEUM
wide proximal segment, 8 shorter distal seg-
ments, each with two long setae (Fig. 51B).
Pereiópod 2 with basis 0.8 times length of
remaining segments combined, with small dis-
tomedial and distolateral setae; ischium 0.4 times
length of merus, with plumose seta distomedially;
merus 1.4 times length of carpus, with plumose
seta distomedially and stout spine-like seta dis-
toJaterally; carpus 1.4 times length of prapodus,
with 2 spine-like setae distomedially and spine-
like seta distolaterally; propodus 0.6 times length
of dactylus, wilh fine seta distomedially; dactylus
has 2 terminal spine-like setae, the longer one 1.4
times longer than itself, 2 small terminal setae and
1 subterminal spine-like seta; all segments have
fine scattered setae (Fig. 51C). Pereiopods 3-5
with merus longer than ischium, carpus as long
as combined length of propodus and dactylus, the
longer of which is propodus; basis with 1 seta
distomedially and fine scattered setae on surface;
ischium with 2 long and 1 minute setae dis-
tomedially; merus with seta distomedially; car-
pus with 2 spine-like setae and 1 fine seta
distolaterally; propodus with spine-like seta and
minute seta distally; dactylus with terminal spine-
like seta, terminal seta and subterminal seta (Fig.
51D-F). Pereiopod 3 with basis 1.1 times length
of remaining segments combined; ischium 0.5
times length of merus which is 0,8 times length
of carpus: propodus 1.7 times length of dactylus
(Fig. 51D). Pereiopod 4 with basis 0.95 times
length of remaining segments combined; ischium
0.5 times length of merus which is 0.8 times
length of carpus; propodus 1.8 times length of
dactylus (Fig. 31E). Pereiopod 5 with basis 0.6
tines length of remaining segments combined;
ischium 0.4 times length of merus which is 0.8
times length of carpus: propodus 1.8 times length
of dactylus (Fig. 51F). Peduncle of uropod 1.1
times às long às telsonic somite, lined with 21-22
plumose setae on inner margin (15-16 long, 6-7
short); endopod 1.1 times as long as peduncle,
subequal in length to exopod, with 6-7 slender
spine-like setae and 10-11 stout spine-like setae
on proximal 2/3 of inner margin, the most distal
stout spine-like seta being well spaced from the
others; apex pointed, without spine-like setae or
mucrones; exopod with 6 plumose setae on
proximal 2/3 of inner margin, apex pointed,
without spine-like setae or mucrones; fine scat-
lered setae on surfaces of telsonic somite,
peduncle and rami (Fig. 52A,B).
OVIGEROUS FEMALE. Integument smooth, lightly
calcified, with minute scattered setae on carapace
BODOTRIIDAE OF MORETON BAY
and pleon, as in d. Carapace length 0.35 S.L.
without distinct dorsal or lateral carinae; width
0.48 times length in dorsal view; antennal tooth
subacute and extending to anterior extremity of
carapace; ocular lobe and pseudorostral lobes as
in d. Pereion 0.4 times as long as carapace.
Pereionite 1 fully concealed by second, both
produced ventrally to form the marsupium;
pereionites with dorsal ridge, dorsolateral margin
of fifth with articulation notch. Pleon robust, all
five pleonites with dorsal ridge and lateral ar-
ticulation notches; first 4 pleonites and telsonic
somite subequal in length, fifth pleonite 1.5 times
as long as fourth. Telsonic somite projecting
posteriorly over bases of uropods. First antenna
3-segmented with terminal segmented flagellum;
first segment geniculate, as in d. Pereiopods as
in d except: pereiopod 1 has smaller spine-like
setae on medial region of basis and 6 reduced
terminal segments (rather than 7) on the exopod;
carpus of pereiopod 2 has 1 (rather than 2) spine-
like setae distomedially (Fig. 53A). Peduncle of
uropod 1.3 times as long as telsonic somite,
without plumose setae on inner margin; endopod
atleastas long as peduncle and subequal in length
to exopod, with 2 minute setae and 6 stout spine-
like setae on proximal two-thirds of inner margin,
the most distal stout spine-like seta being well
spaced from the others; apex pointed, without
spine-like setae or mucrones; exopod with 5
plumose setae on proximal two-thirds of inner
margin, apex pointed, without spine-like setae or
mucrones; fine scattered setae on surfaces of tel-
sonic somite, peduncle and rami, as in d (Fig.
53B,C).
Colour. White to fawn with small black
chromatophores speckled on carapace and ab-
domen.
S.L. Adult d 5.2mm. Adult 9 4.5mm.
HABITAT AND DISTRIBUTION. Most com-
mon over medium and coarse sand in 1-4m of
water; from sites 11 and 12 in Moreton Bay. Both
sexes are common in Pumicestone Passage,
Moreton Bay.
REMARKS. C. sallai most closely resembles
type specimens of C. cooki; however, C. sallai is
50% larger, more robust, with a covering of fine
setae on the pereiopods and uropods. Pereiopod
l also has spine-like setae on the medial bulge of
the basis and a relatively short dactylus,
pereiopod 2 has a strong distal spine-like seta on
the merus and a relatively short dactylus, and the
uropod has rami at least as long as the peduncle.
451
FIG. 53. Cyclaspis sallai sp. nov. allotype ovig. 9. A,
pereiopods 1. B, uropods and telsonic somite, DV. C,
rami of uropod, DV.
Cyclaspis sallai resembles C. juxta Hale from
S AUST, but the rami of the uropod are much
longer relative to the peduncle, and the setation
of the uropod is different. The basis of pereiopod
l has an angular projection on the distomedial
margin which may appear rounded or pointed,
depending on the orientation of the appendage.
ETYMOLOGY. For Michael Salla, who assisted in the
field.
Cyclaspis sp. nov. 1
(Fig. 54A,B)
MATERIAL EXAMINED. QMW20515, ovig. 9,
S.L. 1.9mm, in 70% ethanol, off Dunwich, site 28,
27°29’S, 153°22’E, D. Tafe, 20 May 1989, 4 m, sand,
34 p.p.t. salinity, 24°C water temperature; uncommon.
REMARKS. Further specimens are required in
order to describe the species. Cyclaspis sp. nov.
452
/ 4
N
j Y
/ À
Fog
i
1
FIG. 54. Cyclaspis species. A-B, Cyclaspis sp. nov. 1,
gravid 2. A, LV. B, uropods, ielsonic somite and
pleonite 5, DV. C-E, Cyclaspis sp. nov. 2. C, ovig. 9,
LY. D, subadult ?, LV. E, uropod of ovig. ?, DV.
| superficially resembles C, strigilis Hale and. C.
fulgida Hale, respectively from Fraser Island,
Qld, and Cronulla, NSW. However, it is easily
distinguished by terminal spine-like setae on the
endopod and exopod of the uropod. Also the
carapace does not exhibit the reticulate pattern of
sooty black chromatophores, typical of C. ful-
gida, or the numerous oblique striae, typical of C.
strigilis.
The dorsal line of the carapace of the new
species is almost straight in lateral view. The
ocular lobe is slightly raised, similar to that of C.
stocki (Bacescu, 1990), and the pseudorostrum is
MEMOIRS OF THE QUEENSLAND MUSEUM
54A). Pleonite 5 is 1.5 times the length of pleonite
6 (telsonic somite) and subequal in length to the
peduncle of the uropod; the peduncle has 6 short
medial spine-like setae and the endopod has 6
medial and 3 terminal spine-like setae. The en-
dopod is subequal in length to the exopod and
distinctly shorter than the peduncle (Fig. 54B).
Cyclaspis sp. nov, 2
(Fig. 54C-E)
MATERIAL EXAMINED. QMW20519, ovig. 9,
S.L. 2.7mm, in 70% ethanol, off Coochiemudio Island,
site 34, 27732' S, 153720' E, D. Tafe, 17 June 1990, 4m,
sand, 35 p.p.t. salinity, 18°C water temperature,
QMW 20520, subadult 9, S.L. 2.6mm, same data as
above; uncommon.
REMARKS. Further specimens are required to
describe this species. Cyclaspis sp. nov. 2
resembles C. gibba Hale (1944a:75, figs 4-5)
from Jibbon, NSW. Both species are of similar
length (2.6-3,0mm), both have a smooth finely
reticulate, ovoid shaped carapace with delicate
median dorsal ridge, large antennal notch and
subacute antennal tooth. However, adult and sub-
adult 9? 9 of C. gibba have more prominent
ocular lobes in lateral view than C. sp. nov. 2.
Setation of the uropods is similar in both species
but the exopods are longer, relative to the en-
dopods, in C. sp. nov. 2 (Fig. 35E).
Eocuma Marcusen, 1894
Cyclaspis Kossman, 1880:85.
Eocuma Marcusen, 1894:170. Day, 1978a:168,
DIAGNOSIS. Cuticle strongly calcified and brit-
tle, carapace with lateral horns, at least in 9 . First
pereionite and sometimes second firmly united
with carapace. Basis of first pereiopod prolonged
distomedially. Second pereiopod with basis and
ischium fused. Uropods with peduncle much
shorter than rami; inner ramus l-segmented.
Genus includes 23 species worldwide, most of
which inhabit warm, shallow (5-50m) waters.
Only E. agrion has been recorded from
Australian waters.
curiously upturned as 2 pointed filaments (Fig.
FIG. 55. Eocuma agrion Zimmer d . A,B, whole mount LV, shows relative lengths of carapace and somites. C,
carapace LV, shows curved dorsal profile of carapace, D, anterior carapace LV, shows unusual pseudorostral
region. E, carapace DV, shows reviculate pattern of shallow pits and well developed lateral horns, F, lateral hom
on carapace LV, shows scale-like surface texture of horn. G, anterior carapace DV, shows wide ocular lobe
and pseudorostral projection. H, uropod LV, shows very short peduncle relative to rami,
m
BODOTRIIDAE OF MORETON BAY
|
|
i
4
4
454
Eocuma agrion Zimmer, 1914
(Figs 55, 56A-C)
Eocuma agrion Zimmer, 1914:176, figs 1-2. Hale,
1944b;229, figs 3-4. Hale, 1949a;109, Stephenson
et al., 1978:208.
MATERIAL EXAMINED. QM'W20526, adult d , S.L.
6.1mm, SEM mount, Horseshoe Bay, site 31, 27? 30'S,
153° 2V'E, D. Tafe, 11 Oct 1990, 4m, sand, 35.5 ppt
salinity, 24.5°C water temperature, QMW20527, adult
d. S.L. 6.3mm, in 70% ethanol, same dala as above.
QMW20528, subadult 9, S.L. 5.9mm, SEM mount,
same data as above, QMW20529, subadult 9, S.L.
5.7mm, in 70% ethanol, same data as above,
DISTRIBUTION.Known from Central East
Coast, Lower East Coast and Lower West Coast
(Fig. 4). QLD: Moreton Bay (Hale, 19492); sites
15, 26, 28, 31, and 33, Moreton Bay (herein).
NSW: Cronulla (Hale, 1944b), WA; Fremantle,
Herald Bight and Broadhurst Bightin Shark Bay,
Onslow, Dampier Archipelago, Garden Island
(Hale, 1944b, 19492). Common in Moreton Bay.
d d are taken far more commonly in light-trap
samples than ? 9.
REMARKS. Moreton Bay specimens match £,
agrion in the very long and flexible pleon (Fig.
55A,B). Living specimens are yellowish with a
reliculate pattern of pitting on the carapace (Fig.
55C,D.E). The ocular Jobe is much wider than
long, with | large anterior lens and 4 smaller
postenor lenses (Figs 55G, 56A). The carüpace
has large lateral horns in both sexes (Figs 55F,
56A.C). The uropods are held wide apart and the
rami of each are also spread (Figs 55, 56B).
Subfamily VAUNTHOMPSONIUNAE Sars,
1878
DIAGNOSIS. Exopods on at least first 3 pairs of
perejopods. Always 5 pereiomtes exposed and
endopod of uropod 2-segmented. Second antenna
of 9 often 3-segmented and in most genera third
segment is distinct.
Gephyrocuma Hale, 1936
Gephyrorima Hale, 1936b-412. Hale: 1954b:247.
MEMOIRS OF THE QUEENSLAND MUSEUM
DIAGNOSIS. Ocular lobe wide and not distinctly
separated from frontal lobe, lenses very large.
Antennal notch so widely open that no distinct
incision nr antennal angle is evident. Pleon
reduced, at most 2/3 as long as carapace in Z,
shorter in €. First antenna strongly geniculate,
with segments of peduncle globose. Basis of third
maxillipeds without external apical lobe but with
very large inner lobe. Basis of pereiopod I dis-
tinctly rwisted, with no distal inner lobe. Exopods
of pereiopods | and 2 well-developed, rudimen-
tary on 3 and 4. Uropods with short peduncle and
with endopod 2- segmented, the first segment
much longer than the second.
REMARKS. Four species are known from
Australia: G. pala from Gulf St. Vincent, SA; G.
repandum from Cronulla, NSW and Careening
Bay, WA;G. simile from Shark Bay, WÁ; and G.
sp. nov. 1 from Moreton Bay.
KEY TO THE AUSTRALIAN SPECIES OF
GEPHYROCUMA
(Adapted from Hale, 1944b)
LExapod of third pereiopod ]-segmented 2
Exopod of third pereiopod 2-segmented. ... 3
2, Second endopod segment of uropod with six inner
spine-like setae in d, none in
behi piegi eii om repandum Hale
Second endopod segment of uropod with one
inner spine-like seta in d and F .. . sp- pov. |
3.Pleon at most barely longer than pereionites
together. First segment of endopod of uropod
less than twice as long as second . . . pala Hale
Picon at least half as long again as pereionites
together. First segment of endopod of uropod at
least 4 times as long as second . . . simile Hale
Gephyrocuma repandum Hale, 1944
(Figs 56D-H, 57H)
Gephiveacuma repanda Male, 1944b:248, figs 15, 16,
17B. Hale, 19492:110, figs 2, 3.
MATERIAL EXAMINED, QMW20530, adult d, S.L.
2.3mm, SEM mount, Pumicestone Passage, site 12,
26°49°S, 153"08' E, J. Greenwood, 14 Mar 1991, 2m,
sand, 34.2 ppt salinity, 27.0°C water temperature,
QMW20531, adult d, S.L. 2.2mm, in 70% ethanol,
same data as above, QM W20532, adult 9, S.L, 1.9mm,
FIG, 56. Eociuma and Gephyrocuma species. A-C, Eocuma agrion, d . A, anterior portion of carapace, DV. B,
LV. C, cephalothorax, DV. D-H, Gephyrocuma repandum. D, type d, LV. E, paratype d uropod, DV and Ei,
terminal spines of endopod. F, ovig. ? , LV. G, 9 uropod, DV. H, ovig. 9 cephalothorax, DV. I-J, Gephyrocuma
similis, \ type d uropod; DV and li, distal segment of endopod. J, type ovig. 9 uropod, DV. (A-E, Hale, 1944b.
F-J, Hale, 19493).
BODOTRIIDAE OF MORETON BAY 455
456
SEM mount, same data as above, QMW20533, adult
2 S.L. 2.0mm. in 70% ethanol, same data as above.
DISTRIBUTION. Lower and Central East Coast
and Lower West Coast (Fig. 4). NSW: Cronulla
(Hale, 1944b). Qld: sites 6-9, 10-12 and 31 in
Moreton Bay. WA: Garden Island (Hale, 1949).
REMARKS. Moreton Bay specimens match G.
repandum from Cronulla, NSW; cuticle thin and
smooth with blackish chromatophores (Fig.
56D.F); carapace with dorsal margin evenly and
slightly convex; ocular lobe much broader than
long (Fig. 56D.F,H); pedigerous somites all ex-
posed, together 2/3 as long as carapace (Fig.
S6D.F); pleon more than 2/3 as long as
cephalothorax (Fig. 57H); exopod of third
pereiopod 1-segmented (Fig. 56D,F); d uropods
stout, peduncle only c.1/2 as long as exopod, with
a row of long plumose setae on inner margin;
endopod a little longer than exopod, segment 1
with spinules on inner margin, segment 2 with
inner row of 6 stout spine-like setae and ] ter-
minal spine-like seta; exopod with 8-10 long
plumose setae on inner margin and 3 unequal
terminal spine-like setae (Fig. 36E); 9 uropods
stout, peduncle shorter than in d, with no long
inner setae; endopod with denticles but no spine-
like setae on inner margin; exopod with 1 inner
seta and 3 terminal spine-like setae (Fig. 36G). G.
repandum was taken in relatively large numbers
throughout the year ar sites 10, 11 and 12, in
combination with G, barbarae . lt is the most
abundant cumacean recorded at site [2 in
Pumicestone Passage.
Gephyrocuma sp, nov. |
(Figs S7A-G, 70H)
MATERIAL EXAMINED. QM W20534, adult d, S.L.
1.8mm, SEM mount, Pumicestone Passage, site 12,
26°49°S, 153"08'E, J. Greenwood, 4 May 1990, 2m,
coarse sand, 33.3 ppt salinity, 23.5°C water tempera-
ture. QM W20535, adult 9, S.L. l. 5mm. data same as
above.
REMARKS. Further specimens arc required to
describe the species. Gephyrocuma sp. nov. |
resembles G. repandwumn Hale and G. similis bul
MEMOIRS OF THE QUEENSLAND MUSEUM
differs in shape of pereiopod 1, maxillipeds 3 and
selation of the uropods (Fig. 57A-D,G). The
menus and carpus of maxilliped 3 much more
robust than in G. repandum and the uropods are
devoid of plumose setae (Fig. 57E,G). The en-
dopod and exopod are more robust than in G.
repandum (Fig. SGE,G) and G. similis (Fig.
561,J), the endopod of both d and 9 have 5-6
short inner spine-like setae on the first segment,
1 on the second and 1 terminal; exopod with no
inner spine-like setae and 3 unequal terminal
spine-like setae (Fig. 57D,G). The carapace is
covered with a reticulate pattern of crescent
shaped scales, between which are scattered pits
with short sensory filaments (Figs 57F, 70H).
Glyphocuma Hale, 1944
Glyphocuma Hale, 1944b:268.
GENOTYPE. Synipedomma bakeri Hale,
19362:397.
DIAGNOSIS. Pseudorostral lobes not extending
in front of ocular lobe, which is narrow. Basis of
maxilliped 3 with large external laterodistal lobe,
dentate on medial edge and reaching distal end of
merus. $ with exopods on pereiopods 1-4, those
of fourth pair sometimes small, Female with ex-
opods on pereiopods 1-3 only. First antenna with
accessory flagellum 2-segmented, Second anten-
naof 9 3-segmented, with conical distal segment
distinct. Mandibles elongate, with long row of
spine-like setac (up to 20). Telsonic somite
produced posteriorly, its apex rounded and slight-
ly excavate.
REMARKS. This genus ts close to Sympodomma
but differs in having an exopod on the fourth
pereiopod of the d, and in having the merus of
maxilliped 3 less expanded externally. It
resembles Heterocuma but in that genus the crest
of the carapace is not incised in the 2. Maxilliped
3 of Glyphocuma has the carpus widened as in
Cyclaspis, the terminal segment of the second
antenna of the 9 is tiny, the telsonic somite is
very different, and the segments of the flagellum
of the ¢ second antenna are extremely short.
Glvphocuma contains 5 species, all of which
FIG. 57. A-G, Gephyrocuma sp. nov. 1. A, d. LV shows relative lengths of carapace and somites, B, d LV
shows relative lengths of carapace and somites. C, £ DV shows maximum width in anterior region of carapace.
tapers posteriorly, D, 2, DV, shows maximum width in posterior region of carapace. E, d, VLV shows robust
third maxillipeds. F, 3, DV, detail af integument of carapace showing crescent shaped scales and minute
filaments. G, d uropod, DV, shows robust, sparsely spinuled rami. H, Gephyrocuma repandum Hale d VLV
shows numerous setac on pereiopods and rami of uropods,
BODOTRIIDAE OF MORETON BAY 457
@kY 390E1 8816766 SP.23
458
MEMOIRS OF THE QUEENSLAND MUSEUM
BODOTRIIDAE OF MORETON BAY
are Australian and two of which (G. halei, G.
serventyi) have been previously recorded from
Queensland.
SEXUAL DIMORPHISM. The ovigerous 9 and
immature d have crest of the carapace finely or
coarsely serrate, or incised with resultant angular
projections, or strongly ridged. Adult d's have the
armature of the dorsum obliterated and the anten-
nal notch widely open. They also have the
anterolateral portion of pereionite 4 developed as
a lobe which may overlap pereionite 3,
KEY TO AUSTRALIAN FEMALES OF
GLYPHOCUMA
(Adapted from Hale, 1944b)
| Anterior half of crest of carapace cut into 6 or
more smallteeth . 2... eee 2
Anterior half of crest of carapace smooth or with
1 or 2 incisions, but no teeth .. . 2. eS
2.Carapace twice as long as deep, with dorsal teeth
inconspicuous; antennal notch narrow: ocular
lobe projecting well beyond pseudorostral lobes
and with corneal lenses not confined to anterior
portion bakeri (Hale)
Carapace less than twice as long as deep, with
dorsal teeth large; antennal notch wide; ocular
lobe not projecting beyond pseudorostral lobes
and with small corneal lenses restricted to
anterior portion >... . <... dentatum Hale
3, Anterior half of crest of carapace smooth, Exopod
of uropod with at least 10 plumose setae on
inner margin halei Greenwood & Johnston
Anterior half of cresi of carapace with | or 2 in-
cisions. Exopod of uropod with at most 6
plumose on inner margin... .. sss 4
4.Carapace slender, with 2 dorsomedial incisions,
the second with 2 or 3 denticles; ocular lobe nar-
row, more than twice as long ss wide, apically
rounded in dorsal view ingequale Hale
Carapace robust, with | dorsomedial incision and
2 or 3 denticles; ocular lobe as wide as long. àpi-
cally angular in dorsal view
Poh. A Ah servenrvi Hale
KEY TO AUSTRALIAN MALES OF
GLYPHOCUMA
(Adapted from Hale, 1944b)
1, Body slender, the carapace more than twice as
lonppasdcep 2 ..:3 5.014243 po: 2
Body robust, the carapace less than twice as long
as deep
PEEL LLL LLLLLLLLLLLLLLLLLLLLI
2.Exopod of uropod without spine-like setae on
lateral margin. Exopod of fourth perciopod with
flagellum 2-segmented bakeri Hale
Exopod of uropod with spine-like setae on latera!
margin, Exopod of fourth pereiopod with flagel-
lum 4- or 5-segmented 3
3.Dorsal edge of carapace smooth; exopod of fourth
pereiopod with flagellum 4-segmented
halei Greenwood & Johnston
Dorsal edge of carapace sinuate; exopod of
fourth pereiopod with flagellum 5-segmented
inaequale Hale
4.Ocular lobe narrow, more than twice as long as
wide, with corneal lenses confined to anterior
end which is rounded dentatum Hale
Ocular lobe as wide as long, with corneal lenses
rcaching to posterior end which is angular
serventyi Hale
Glyphocuma dentatum Hale, 1944
(Figs 58A-D, 59A-G)
Glyphocuma dentatum Hale, 1944b:273, Figs 33, 34.
MATERIAL EXAMINED. QMW20536, subadull 9,
S.L. 4mm. SEM mount, Horseshoe Bay, site 31, 27°
30'S, 153° 21'E, D. Tafe, 7 April 1991, sand, 3m, 33.2
ppt salinity, 24.8°C water temperature; QMW20537,
subadult d, S.L. 5.5mm, in 70% ethanol, data same as
above.
DISTRIBUTION. Lower and Central East Coast
(Fig. 4). NSW: Port Hacking, Ulladulla, Eden,
46-100m: on mud (Hale, 1944b). Qld: sites 28, 31
and 36 in Morcton Bay.
REMARKS. Moreton Bay specimens match G.
dentatum from Port Hacking, NSW. The degree
of dentation of the dorsal edge of the carapace
was found to be variable (Figs 58A-D, 59A-G).
Common in parts of Moreton Bay.
Glyphocuma halei Greenwood & Johnston,
1967
(Figs S8E-F, 60)
Glyphocuma halei Greenwood & Johnston, 1967:93,
figs 1-2.
MATERIAL EXAMINED. PARATYPES
FIG. 58. A-D, Glyphocuna dentatim 2. A,B. LV, shows relative lengths of carapace and somites, C, carapace
LV , shows 7 slender teeth on dorsal margin. D, whole mount LV, shows anterior end of carapace. E-H,
Glyphocuma halei 9, B, uropods DLV, shows relative lengths of peduncle and rami. F, uropod rami DLV,
shows spination of rami. G,H, subadult LY, shows smooth dorsum Of carapace and relative lengths of segments.
460 MEMOIRS OF THE QUEENSLAND MUSEUM
Tb te! ts
FIG, 59, A-G, Glyphocuma dentorum, A, lois pe ovig. 9, LV. B, 9 cephalothorax, DV. C, paratype ? uropod.
D, juvenile d cephalothorax, DV. E, allatype ó ocular lobe, DV. F, allotype 8 cephalothorax, LV. G, subadult
& uropod. H-O, Glyphocuma bakeri. H-K, type 2. H, LV. ], ocular lobe, DV. J, cephalothorax, DV. K, uropod.
L-O, d. L, LV. M, cephalothorax, DV, N, ocular lobe, DV. O, uropod. (A-G and L-O, Hale, 1944b. H-K, Hale,
19363).
BODOTRIIDAE OF MORETON BAY
QMW 20538, adult 2, allorype, S.L.
10mm, in 70% ethanol, Horseshoe
Bay, site 31, 27^30'S, 153°21'E, D.
Tafe, 7 April 199], sand, 3m, 33.2
ppt salinity, 24.8°C waler tempera-
ture; QM W 20539, subadult 9 , S.L.
3.6mm, SEM mount, off Macleay
Island, site 36, 27°36'S, 153722 E,
D. Tafe, 8 Nov. 1989, sand/silt. 2m,
33.2 ppt salinity, 26,8°C water
temperature; QMW20540, adult 3,
S.L. Himm, in 70% ethanol, off
Coochiemudlo Island, site 34,
27732'8, 153720' E, D. Tafe, 17 June
1990, sand, 5m, 34,8 ppt salinity,
18.8°C water temperature;
QMW20541, subadult d, S.L.
8.5mm, in 70% ethanol, data same
as above.
DESCRIPTION. FEMALE. ln-
tegument calcified with fine
granular texture (Fig. 58G,H).
Carapace length 0.25 S.L. with
strong median dorsal ridge, the
anterior 1/2 of which bears fine
crenulations though the dorsal
profile is smooth and slightly ar-
ched, sides devoid of ridges or
sculpture, as in d; depth 0.57
times length in lateral view; an-
tennal notch widely open; ocular
lobe and pseudorostral lobes as
in d. (Figs 58G,H, 60F). Pereion
0.86 times as long as carapace.
Pereionite 1 visible only above
lateral midline; pereionites
without lateral ridges,
posterolateral margin of
pereionite 5 produced as lobe
which overlaps pleonite 1, as in
d (Fig. 58G,H). Pleon robust, all
5 pleonites without lateral ridges
or lateral articulation notches;
first 4 pleonites and telsonic
somite subequal in length, fifth
pleonite 1.35 times as long as
fourth (Fig. 58G,H). Telsonic somite projecting
posteriorly over bases of uropods (Figs 58H,
60G). First antenna 3-segmented with terminal
segmented flagellum; first segment geniculate, as
in d . Pereiopods as in d. Peduncle of uropod 1.2
times as long as telsonic somite, with 14-16 naked
spine-like setae along length of inner margin;
endopod 0.75 times as long as peduncle and 0.9
times as long as exopod, with 9-11 spine-like
setae on inner margin and | spine-like seta on
$61
FIG. 60. Glyphocuma halei. A-E, 3. A, DV, B, cephalothorax, LV. C,
carapace, DV. D, anterior of carapace, LV. E, uropod and telsonic somite,
DV. (All after Greenwood & Johnston, 1967). F-G, 9. F, cephalothorax,
LV, G, uropod, DV.
outer margin of proximal segment, 10-11 spine-
like setae on inner margin of distal segment and
3 terminal spine-like setae, the longest of which
is as long as the segment; distal segment slightly
longer than proximal segment as in d; exopod
with 11-13 plumose setae on inner margin, 12-14
spine-like setae on outer margin and 3 terminal
spine-like setae, the longest of which is at least
1/2 as long as exopod; terminal spine-like setae
462
of endopod and cxoped bear tiny mucrones at
apex (Figs S8E,F, 60G).
Colour. Cream with scattered brown
chromatophores on carapace and abdomen.
S.L. Adult d. 9-11 mm. Adult ? 10mm.
HABITAT AND DISTRIBUTION. Most com-
mon over medium and coarse sand in 1-4m of
water; from Central East Coast (Fig. 4). Old:
Waterloo Bay in Moreton Bay (Greenwood &
Johnston, 1967); sites 12, 31, 34 and 36 in
Moreton Bay,
REMARKS. d specimens matched G. halei from
the same region. 9 9 have not previously been
found. The adult 9 from Moreton Bay resembles
9 s of G. dentatum from Port Hacking, NSW, but
differs in the structure and setation of the uropods
(Figs 58E,F, 59C, 60G). In d d and the adult 9
of G. halei the distal endopod segment of the
uropod is longer than the proximal segment (Fig.
60E,G), whereas in G. dentatum it is shorter (Fig.
59C.G).
G. halei also resembles G. bakeri, however,
both sexes of G. halei bear at least 12 outer
spine-like setae on the exopod of the uropod. The
same segment in G, bakeri has no outer spine-like
setae (Fig. 59K,O). Also, as noted by Greenwood
& Johnston (1967:98), the rami of the uropod arc
relatively longer in G. halei, and the exopod of
the fourth pereiopod has 4 segments in G. halei,
2in G. bakeri. The general body shape of G. halet
is very similar to G. bakeri in lateral view (Fig.
59H,L), but the carapace is not as slender in
dorsal view (Fig. 591,J,M,N),
The uropods of the G. halei *' carry terminal
spine-like setae on the endopod and exopod, each
of which bear distinct lerminal spinules (Fig.
60G). Spinules are not present on the terminal
spine-like setae of F uropods nf G. denrarum or
G, bakeri (Fig. 3TC,K).
Leptocuma Sars, 1873
Leptocuma Sars, 1873-24. Hale, 1936b; 409, Hale,
1944b:251.
DIAGNOSIS. Pseudorpstral lobes extend in front
of ocular lobe, not meeting; mandible robust,
MEMOIRS OF THE QUEENSLAND MUSEUM
with at least 9 spine-like setac: busis of the mix-
illiped 3 not produced distally; pereiopod 2 with
brush of distal setae on the propodus and dac-
tylus, but no spine-like setae; antenna | with
accessory flagellum 1-segmented; telsonic
somite produced posteriorly with apex angular;
maxilliped 3 with ischium short and merus not as
long as carpus; ocular lobe wide, moderate or
large in size; second antenna of & with segments
of flagellum elongate; pereionite 3 of ? produced
forward on each side to form lobe overlapping
pereionite 2; anterolateral parts of pereionite 4 of
d similarly expanded to override pereionite 3.
REMARKS. The genotype, L kinbergii, was
described fromthe Ẹ , taken in the South Atlantic
off South America. Only two other species have
since been recorded from the Americas, L.
forsmani Zimmer,1943 and L. patagonicum Roc-
catagliata,1993. Hale (1944b) described 2
Australian species, L. pulleini (QLD, NSW, SA)
and L. sheardi (SA); d & of these 2 species have
5 pairs of pleopods and the exopod of perciopod
4 is rudimentary, as in the 9. Hale described L.
intermedium (NSW), L. nichollsi (WA), L,
abstipum (NSW), L. serriferum (NSW, WA) and
L. vicarium (NSW) from Australia. The
American species were keyed out by Roc-
catagliata (1993). The Australian species fall into
two well defined groups.
KEY TO AUSTRALIAN SPECIES OF
LEPTOCUMA
(Adapted from Hale, 1944b)
1.First pereiopod wilh a prominent simple spine-like
sela on distomedial margin of basis, preceded by
several shorter spine-like setae, and with a well-
developed brush of sctae at distal end of
propodus. Sciae of pereiopods 3-5 very
numerous, Uropod with first segment ofen-
dopod shorter, or barely longer, than second.
Over I3mm in length ....-...2.-.- 2
Pereiopod } with a serrate spine-like sela on dis-
tomedial margin of basis, preceded by 1 longer
spine-like seta, also serrate; wilh sparse setae at
distal end of propodus. Sctae of perciopods 3-5
not very numerous. Uropod with first segment
of endopod much longer than second. Less than
Bwm in length.. 2.2. nn 3
FIG. 61. Leptocuma barbarae sp. nov. d. A,B, whole mount LY shows relative lengths of carapace and somites.
C, whole mount LV, shows relative lengths of telsonic somite and pleonite 5. D, whole mount DV, shows width
tapering from carapace to telsonie somite. E, carapace and pereionites 1-3 DV, shows maximum width in
mid-region of carapace, F, carapace and pereionites 1-3 LV, shows relative length and depth of carapace. G,
uropods DV. shows relative lengths of peduncle and rami. H, Pereionites 2-3 LV, shows slender spines on
anterolateral margins of pereionites 2-3.
BODOTRIIDAE OF MORET
12mm
12mm
464 MEMOIRS OF THE QUEENSLAND MUSEUM
SP42F sky — PORE
— 188B8Hmn
65 13mm
SP42F
BODOTRIIDAE OF MORETON BAY
2, Second pereiopixl with carpus 2/3 as long again
as merus pulleini Hale
Second pereiopod with carpus subequal in length
tp mëma of Se ee vicarium Hale
3.Carapace with strong median dorsal ridge extend-
ing from ocular lobe almost to first pedigerous
Sd wp. SS 820 ee oo eS s ;
Carapace with at most scarcely distinguishable
median dorsal ridge ........... .. 5
& à hb c» 9 we me eg ^4
4.Anterolateral margins of pereionites 2 and 3 with a
row of spine-like setae. Pereiopod 1 has 2 con-
spicuous serrate spine- like setae on distornedial
SPAS A oy ARs ae c s kennedyi sp. nov.
Anterolateral margins of percionites 2 arid 3
without spine-like selae. First pereiopod has 4
conspicuous serrate spine-like setas on dis-
tomedial margin nichollsi Hale
5.Dorsal margins of pereionites undulating in lateral
view, One of the terminal spine-hke setae of en-
dopod of uropod geniculate ( 9) or hooked (d ).
Pleon with obvious lateral and dorsal carinae
p deu Er Siu d obstipum Hale
Dorsal margins of pereionites smooth. Terminal
spine-like setae of endopod of uropod straight or
barely curved. Pleon smooth or with scarcely dis-
tinguishable traces of carinae =. 6
6.Size under 5mm. Uropod with second segment of
endopod much more than half length of first — 7
Size about 7mm. Uropod with second segment of
endopod about hall length of first oo... LI
T. Anterolateral margins of second and third
pedigerous segments with row of short spine-
hkésétae we mm barbarae sp. nov.
Anterolateral margins of second and third
pedigerous segments without spine-like setae
die eas D M, qug u ; . serriferum Hale
8.First pereiopod with propodus much longer than
dactylus. Second pereiopod with propodus and
dactylus subequal in length sheardi Hale
First pereiopod with propodus scarcely longer
than daclylus. Second pereiopod with dactylus
fully 1/3 as long again as propodus
intermedium Hale
= Arat w Roth m A eA m Are
Leptocuma barbarae sp. nov.
(Figs 61-64)
MATERIAL EXAMINED. HOLOTYPE
QMW20542, adult d, S.L. 3.3mm, PSM #16, Hopse-
shoe Bay, sile 31, 27°30'S, 153?21' E, D. Tafe, 4 Feb.
1993, 3m, sand, 34.2 ppt salinity, 26°C waler tempera-
tur. PARATYPES QMW20543, ovig. 2, allotype,
S.L. 3.6mm, PSM #17, same data as holotype;
QMW20544, adult d, S.L. 3.6mm, PSM #43, Taa-
galooma, site 15, 27^11'S, 153*19' E, D. Greenwoexd,
14 April 1990, &m, sand, 35.5 ppt salinity, 24°C water
temperature; OMW20545, ovig. 2, S.L. 3.7mm, PSM
#44, same data as above; QMW20546, ovig. F, S.L.
3.5mm, PSM #18, same data as holotype; QMW 20547,
adult d , S.L. 3.2mm, SEM mount, Horseshoe Bay, site
31, D. Tafe, 7 April 1991, 2m, sand, 33.2 ppt salinity,
25.5°C water temperature; QM W20548, ovig. €, S.L.
3.1mm, SEM mount, same dala as above.
DESCRIPTION. MALE. Integument lightly cal-
cified, with fine granular texture (Fig. 61A,B).
Carapace length 0.26 S.L. and depth 0.6 times
length in lateral view with lateral margins evenly
rounded: dorsal edge slightly arched: mild
median dorsal ndge present on anterior 1/2 of
carapace; lateral surfaces devoid of ndges or
sculpture; greatest width at middle of Jength in
dorsal view: antennal notch a shallow concavity,
no antennal tooth or antennal ndge; pseuderostral
lobes wide, extending anteriorly beyond ocular
lohe which is wider than long (Fig. 61A-F).
Percion robust, 0.9 times length of carapace: sides
of pereionite | concealed by pereionite 2, which
bears 7-8 slender spine-like setae on anterolateral
margin; pereianite 3 overlaps second laterally
and also bears 7-8 spine-like setae on
anterolateral margin; pereionite 4 overlaps third
and fifth laterally. and bears 4-5 spine-like setae
on each of overlapping lobes; pereionite 5 over-
laps pleonite | laterally and bears 4-5 spinc-like
setae on overlapping lobe; dorsolateral carinae
pereionites 3-5 (Fig. 61A-F,H). Pleon and tel-
sonic somite 0.5 S.L. without dorsal or dor
solateral cannae; first + pleonites subequal in
length, each with posterolateral overlapping
lobes beating 4-5 small spine-like setae: fifth
pleonite 1.3 times as long as fourth. Telsonic
somite subequal in length to fourth pleonite, with
posterodorsal projection which is U-shaped in
dorsal view (Fig. 61A-D,G). First antenna us in
L serriferum except peduncle has tuft of about 12
fine setae proximally, third segment has group of
7 setae distally. all of which extend beyond tirst
segment of flagellum. Mandible as in L ser-
FIG. 62. Lepiocuma barbarae sp. nov, ovig. 9, A,B, whole mount LV, shows relative lengths of carapace and
somites. C, whole mount LV, shows relative lengths of telsonic somite and peduncle of uropod. D, carapace
and pereioniies 1-3 DV, shows maximum width of carapace in posterior region. median dorsal ridge barely
visible, granular texture. E, carapace and pereionites 1-3 DLY, shows median dorsal ridge visible on anterior
half of carapace, F, carapace and perejonites 1-4 LV shows lateral lobes of pereionite 3 overlapping pereionites
2 and 4, slender spines on anterolateral margins of pereionites 2-3. G, uropods DV, shows spination relative
lengths of peduncle and rami.
466
FIG. 63, Leptocuma barbarae sp. nov. holotype adult 6. A, pereiopod 1, B,
pereiopod 2, C, pereiopod 4.
riferum except there are 10-11 (rather than 12)
spine-like setae. Third maxilliped as in L. ser-
riferum except setae along medial margin of basis
are all plumose and of similar length. All
pereiopods 7-segmented (Fig. 63). Pereiopod | as
in L. serriferum except dactylus is 0,82 times
length of propodus and dactylar spine-like seta is
0.72 times length of dactylus (Fig. 63A).
Pereiopod 2 as in L. serriferum except setae on
MEMOIRS OF THE QUEENSLAND MUSEUM
medial margin of basis are
plumose, propodus is 0.8 times
length of dactylus and has 8
medial setae (Fig. 63B).
Pereiopods 3-5 with merus
longer than ischium, carpus
slightly longer than merus and
longer than combined length of
propodus and dactylus;
propodus twice length of dac-
tylus; dactylar spine-like seta at
least 1.5 times length of dac-
tylus; basis with 2 setae distal-
ly, 2-3 medially and 2-3
laterally; ischium with 2-3
spine-like setae and 2-3 setae
distally; merus with 1-2 spine-
like setae distally and 1-2 setae
medially; carpus with 2-3
spine-like setae distally and 1-2
setae laterally; propodus with
spine-like seta and minute seta
distally; dactylus with terminal
spine-like seta, terminal seta
and subterminal seta. Exopod
of pereiopod 3 well-developed
with bulbous basis, like those
of pereiopods 1 and 2; exopod
of pereiopod 4 rudimentary and
1-segmented, Pereiopod 4 as in
L. serriferum except basis of
endopod has 3 medial and 2
lateral setae (Fig. 63C).
Peduncle of uropod 1.35 times
length of telsonic somite, lined
with 18-19 spinose spine-like
setae (2 rows) on inner margin;
endopod subequal in length to
peduncle and to exopod, with
15-16 spinose spine-like setae
on inner margin of proximal
segment, 5-6 spinose spine-
like setae on inner margin of
distal segment (increasing in
length distally) and 3 terminal
spine-like setae, the longest of
which is subequal in length to
distal segment, distal segment 0.6 times length of
proximal segment; dorsal surface of both en-
dopodal segments bear numerous fine setae; ex-
opod with 9-IO plumose setae and 2 slender
spine-like setae along inner margin, outer margin
with short subterminal spine-like seta, 3 terminal
spine-like setae, the longest of which is 0.5 times
length of exopod; terminal and subterminal
BODOTRIIDAE OF MORETON BAY
spine-like setae of both rami bear
minute apical setae; (Figs 61G,
64A).
OVIGEROUS FEMALE. Integument
lightly calcified, with fine
granular texture, as in d (Fig.
62A-E). Carapace length 0.24
S.L. and depth 0.8 times length in
lateral view with lateral margins
evenly rounded; dorsal edge
slightly arched; median dorsal
ridge is obsolete on posterior 1/2
and barely visible on anterior 1/2;
lateral surfaces devoid of ridges
or sculpture; greatest width at
posterior of length in dorsal view;
antennal notch a shallow con-
cavity, no antennal tooth or anten-
nal ridge; pseudorostral lobes
wide, extending anteriorly
beyond ocular lobe which is wider
than long (Fig. 62A-E). Pereion
robust, 1.25 times length of
carapace; sides of pereionite 1 '
concealed by pereionite 2, which
bears 7-8 slender spine-like setae
on anterolateral margin;
pereionite 3 overlaps second and
fourth laterally and bears 9-11
spine-like setae on anterolateral
margin; pereionite 4 overlaps fifth
laterally and fifth overlaps
pleonite 1 laterally; dorsolateral
carinae pereionites 3-5 (Fig. 62A-
F). Pleon and telsonic somite 0.48
S.L. without dorsal or dorsolateral
carinae; first 4 pleonites subequal
in length, each subcylindrical
with V-shaped posterolateral
margins; fifth pleonite 1.5 times
as long as fourth. Telsonic somite
subequal in length to fourth
pleonite, with posterodorsal
projection which is U-shaped in
dorsal view (Fig. 62B,C,G). First
antenna and mandible as in L. serriferum except
mandible has 10-11 (rather than 12) spinc-like
setae. Third maxilliped as in L. serriferum except
setàe along medial margin of basis all plumose
and of similar length. Pereiopod 1 as in L. ser-
riferum except dactylus 0.82 times length of
propodus; dactylar spine-like seta 0.58 times
length of dactylus.Pereiopod 2 as in L. serriferum
except propodus with group of 10 (rather than 5)
467
FIG. 64. Leptocuma barbarae sp. nov. A, uropods and lelsonic somite of
holotype d', DV. B, uropods and telsonic somite of allotype ovig. 9,
DV,
distal setae. Pereiopods 3-5 as in L. serriferum
except basis of fourth with 4 medial and 4 lateral
setae. Peduncle of uropod 1.3 times length of
telsonic somite, lined with 10-12 spine-like setae
on inner margin; endopod 1.2 times length of
peduncle and subequal in length to exopod, with
11-15 spine-like setae on inner margin of
proximal segment, 5-7 spine-like setae on inner
margin of distal segment, 3 terminal spine-like
468 MEMOIRS OF THE QUEENSLAND MUSEUM
BODOTRIIDAE OF MORETON BAY
setae, the longest of which is at least 0,75 times
length of distal segment, distal segment at least
0.5 times length of proximal segment; dorsal
surface of both endopodal segments bear
numerous fine setae; exopod with 5-7 short setae
and 2 slender spme-like setae on distal half of
inner margin, short subterminal spine-like seta on
outer margin, 3 terminal spine-like setae, the
longest of which is 0.5 times length of exopod;
terminal and subterminal spine-like setae of both
rami bear minute apical setae (Fig. 62G. 64B).
Colour. Pale yellow with conspicuous blackish
chromatophores which may appear as dark bands
on carapace,
S.L. Adult d 3.2-3 5mm. Adult 2 3.1-3.6mm.
HABITAT AND DISTRIBUTION. Most com-
monly over medium and coarse sand in 1-8m of
water: from sites. 15, 28, 30, 31, 34, and 35 in
Moreton Bay. In late summer L. barbarae pre-
dominated in the water column at night at site 31.
REMARKS. The Moreton Bay specimens
resemble L serriferum from Cronulla, NSW hut
differ in the spination of pereionites, pleonites
and uropods. Fresh specimens of both L. bar-
barae and L. serriferum (Hale, 1944b:261) are
coloured pale yellow with conspicuous blackish
chromatophores, which in L. barbarae may ap-
pear às bands on the carapace. The most striking
difference between the two species is the
presence, in L. barbarae, of short spine-like setae
on the anterolateral margins of pereionites 2 and
3 and on the posterolateral margins of pleonties
1-4. The spination of the ? uropod is also reduced
compared to L. serriferum (Table 3)
The standard length of L. barbarae ( 9 : 3.6mm,
3: 3.4mm) is a little smaller than L. serriferum
( 9. 4.4mm. d : 42mm).
ETYMOLOGY. For Barbara Koh, who assisted in the
field.
Leptocuma kennedyi sp. nov
(Figs 65-67)
EXAMINED.
MATERIAL HOLOTVYFE
469
TABLE 3, / uropod setation of L. serriferum and L
barbarae.
| Pedunclespinaion | — 15 — | — 1012. |
18 11-15
[Endopod2spinsion| — 11. | $53 |
QMW 20549, adult d, S.L. 4.3mm, SEM mount, Tan-
galooma, site 15, 27^11' S, 153*19'E, D. Greenwood,
14 April 1990, 8m, sand, 35.5 ppt salinity, 24°C water
temperature, PARAT YPES: QMW20550, ovig. ¢, al-
lotype, 5.L. 4.2mm, SEM mount, same data as above:
QMW20S551, adult 3, S.L. 3.5mm, PSM #15, Horse-
shoe Bay, site 31, 27*30' S, 153°21'E, D. Tafe, 4 Feb.
1993, 3 m, sand, 34.2 ppt salinity, 26°C water tempera-
ture.
DESCRIPTION. MALE. Integument lightly cal-
cified, finely granular and sparsely pitted with
minute indentations (Fig. 65A.C}. Carapace
length 0.26 S.L. and depth 0.6 times length in
lateral view with lateral margins evenly rounded;
dorsal edge slightly arched and sinuate; darsum
with strong median ridge throughout length;
lateral surfaces devoid of ridges or sculpture;
greatest width at middle of length in dorsal view;
antennal notch a moderate concavity, no antennal
tooth or antennal ridge; pseudorostral lobes wide,
extending anteriorly beyond ocular lobe which is
wider than long (Fig. 65A,C.E). Pereion robust,
11.9 times length of carapace: sides of pereionite
1 concealed by pereionite 2. which bears 9
slender spine-like setae on anterolateral margin;
percionite 3 overlaps second laterally and bears
7-8 spine-like setae on anterolateral margin;
pereionite 4 overlaps third and fifth laterally, and
bears 3-4 spine-like setae on anterolateral mar-
gin; pereionite 5 overlaps first pleonite laterally
(Fig. 65A,C). Pleon and telsonic somite 0.5 S.L.
with faint dorsolateral carinae; first 4 pleonites
subequal in length, each with poorly developed
posterolateral overlapping lobes; fifth pleonite
1.6 times as long as fourth. Telsonic somite sub-
equal in length 1o fourth pleonite, with
posterodorsal projection which is V-shaped im
dorsal view (Fig. 65A,G). First antenna as in L.
FIG. 65, Leptocuma kennedyt sp. nov, A, whole mount d. LV, shows relative lengths of carapace and somites,
B, whole mount ? LV, shows relative lengths of carapace and somites. C, carapace and pereionites 1-5 & LV.
shows relative length and depth of carapace. slender spines on anterolateral margins of pereionites 1-3. D,
carapace and pereionites 1-3 2 LV, shows relative length and depth of carapace. slender spines visible on
anterolateral margins of pereionites 1-2. E, carapace and pereionites 1-3 d. DV, shows strong median dorsal
ridge and sparse pitting of carapace, F, carapace and pereionites 1-3 9 DLV, shows strong median dorsal Ry
and sparse pitting of carapace, G, Uropod d DV, shows relative lengths of peduncle and rami. H, Uropod
VV, shows relative lengths of peduncle and rami.
470
FIG. 66. Leptocuma kennedyi sp. nov. paratype d. A, pereiopod |. B,
pereiopod 2. C, pereiopod 4.
serriferum except peduncle has tuft of about 20
fine setae proximally, third segment has group of
5 setae distally all of which extend beyond first
segment of flagellum. Mandible as in L. ser-
riferum except there are 10-11 (rather than 12)
spine-like setae. Third maxilliped as in L. ser-
riferum except setae along medial margin of basis
are all plumose and of similar length. Pereiopod
l as in L. serriferum except dactylus is 0.78 times
length of propodus and dactylar spine-like seta
MEMOIRS OF THE QUEENSLAND MUSEUM
0.66 times length of dactylus
(Fig. 66A). Pereiopod 2 as in L..
serriferum except setae on
medial margin of basis
plumose, propodus 0.6 times
length of dactylus and with 8
distal setae (Fig. 66B).
Pereiopods 3-5 with merus
longer than ischium, carpus
slightly longer than merus and
longer than combined length of
propodus and dactylus;
propodus twice length of dac-
tylus; dactylar spine-like seta at
least 1.5 times length of dac-
tylus; basis with 2 setae distal-
ly, 2-3 medially and 2-3
laterally; ischium with 2-3
spine-like setae and 2-3 setae
distally; merus with 1-2 spine-
like setae distally and 1-2 setae
medially; carpus with 2-3
spine-like setae distally and 1-2
setae laterally; propodus with
spine-like seta and minute seta
distally; dactylus with terminal
spine-like seta, terminal seta
and subterminal seta. Peduncle
of uropod 1.5 times length of
telsonic somite, lined with 20-
22 spinose spine-like setae (2
rows) on inner margin; en-
dopod subequal in length to
peduncle and to exopod, with
14-16 spinose spine-like setae
on inner margin of proximal
segment, 5-6 spinose spine-
like setae on inner margin (in-
creasing in length distally) of
distal segment, 3 terminal
spine-like setae, the longest of
which is subequal in length to
distal segment; distal segment
0.6 times length of proximal
segment; dorsal surface of both
endopodal segments bear
numerous fine setae; exopod with 6 plumose
setae and 2 slender spine-like setae along inner
margin, outer margin with 7 short spine-like
setae, 3 terminal spine-like setae, the longest of
which is 1/3 the length of exopod; outer edge
spine-like setae of exopod and terminal spine-like
setae of both rami bear minute apical setae (Fig.
65G, 67 A,B).
BODOTRIIDAE OF MORETON BAY
OVIGEROUS FEMALE. Integument lightly cal-
cified, finely granular and sparsely pitted with
minute indentations, as in d (Fig. 65B,D).
Carapace length 0.24 S.L. and depth 0.63 times
length in lateral view with lateral margins evenly
rounded; dorsal edge slightly arched and sinuate;
strong median dorsal ridge on anterior 3/4 of
dorsum; lateral surfaces devoid of ridges or sculp-
ture; greatest width at posterior of length in dorsal
view (Fig.Pl 25F); antennal notch very small and
subacute; pseudorostral lobes wide, extending
anteriorly beyond ocular lobe which as wide as
long (Fig. 65B,D,F). Pereion robust, 1.1 times
length of carapace; sides of pereionite 1 con-
cealed by second pereionite, which bears 14-15
slender spine- like setae on anterolateral margin;
pereionite 3 overlaps second and fourth laterally
and bears 9-11 spine-like setae on anterolateral
margin and 1-2 on posterolateral margin;
pereionite 4 overlaps fifth laterally and fifth over-
laps first pleonite laterally; no dorsolateral
carinae on pereionites (Fig. 65B,D,F). Pleon and
telsonic somite 0.49 S.L. with faint dorsolateral
carinae; first 4 pleonites subequal in length, each
subcylindrical with small nipple-like projections
on posterolateral margins; fifth pleonite 1.6 times
as long as fourth. Telsonic somite subequal in
length to fourth pleonite, with posterodorsal
projection which is V-shaped in dorsal view (Fig.
65B,H). Peduncle of uropod 1.3 umes length of
lelsonic somite, lined with 5-6 spine-like setae on
inner margin; endopod 1.1 times length of
peduncle and subequal in length to exopod, with
10-11 spine-like setae on inner margin of
proximal segment, 4-5 spine-like setae on inner
margin of distal segment, 3 terminal spine-like
setae, the longest of which is as long as distal
segment, distal segment 0.5 times length of
proximal segment; dorsal surface of both en-
dopodal segments with numerous fine setae: ex-
opod with up ta 5 short setae and 2 slender spine-
like setae on distal half of inner margin, short
subterminal spine-like seta on outer margin, 3
terminal spine-like setac, the longest of which is
0.5 times length of exopod; terminal and subter-
minal spine-like setae of both rami bear minute
apical setae (Fig. 65B,H),
Colour White with small blackish
chromatophores dotted on carapace and pereion.
S.L. Adult d 4.3mm. Adult 2 4.2mm.
HABITAT AND DISTRIBUTION. Most com-
mon over medium and coarse sand in ]-5m of
water; sites 15, 28, 31 and 34 in Moreton Bay.
471
FIG. 67. Leprocuma kennedyi sp. nov. paratype adult
d. Á, rami of uropod, DV. B, uropods and telsonic
somite, DV.
REMARKS. Leprocuma kennedyi most closely
resembles L. nichollsi but has rows of spine-like
setae on the anterolateral margins of pereionites
2 and 3. Also the first endopodal segment of the
uropod is fully twice as long as the second and
has up to 18 inner marginal spine-like setae (cf.
1.5X with 9 spine-like setae in L. nichollsi; Hale,
19493:117).
L. kennedyi is larger than L barbarae and
smaller than L. imermedia. lt also differs from the
above two species in having a strong median
dorsal ridge, extending from the ocular lobe al-
most to the first pereionite. Like L. barbarae it
has a row of short spine-like setae in both sexes
on the anterolateral margins of pereionites 2 and
3. Unlike L. barbarae (Fig. 61G, 62G) the ex-
opod of the d uropod has a row of outer spine-
like setae (Fig. 65G). Like L. intermedia the body
is large and tapering from carapace lo lelsonic
somite, However, the second endopod segment
of the uropod (ds only) of L kennedyi is rela-
tively longer than that of L. intermedia (Hale,
1944h:265).
ETYMOLOGY. For John Kennedy, Queenslan
Museum, who assisted in the Field.
Picrocuma Hale, 1936
Picrocuma Hale, 1936h:415. Hale, 1945c: 177,
DIAGNOSIS, Pseudorostral lobes meeting in
front of wide ocular lobe. First antenna with
3-segmented peduncle and rudimentary acces-
sory flagellum. Mandible with spine-like seta
row, only 4-5 spine-like setae; incisor portion
greatly elongated, with cutting edge tridentate.
Third maxilliped with exopod and similar to first
pereiopod; basis not produced apically.
Pereiopods 1-3 with well-developed exopods in
2. Uropod of both sexes with endopod 1-seg-
mented. Pereionite 2 longer than others.
GENOTYPE. Picrocuma poecilotum.
REMARKS. Salient features of genus are struc-
ture of mandible and unspecialised third maxi-
lliped.
Picrocuma poecilotum llule, 1936
(Fig. 68)
Picrocuma poecilorum Hale, 1936b:415, figs Ta-c, Ba-
i. Hale, 19454:178, figs 22, 23.
MEMOIRS OF THE QUEENSLAND MUSEUM
MATERIAL EXAMINED. SAMC2006, ovig. F,
holotype, S.L. 1.9mm, Wynyard, Fossil Reef, Tas-
mania, on stones, Hale, 1936b. OTHER MATERIAL
QMW20552, adult d, S,L. 1.3mm, in 70% ethanol,
Tangalooma, site 15, 27°11°S, 153°19°E, D. Green-
wood, 14 April 1990, &m, sand, 35.5 ppt salinity, 24°C
water temperature; QMW20553, adult ? , S.L. 1,2mm,
in 70% ethanol, sarne data as above; QM W20554., adult
d, S.L. 1.4mm, SEM mount, same dala as above;
QMW20555, adult 2, S.L. 1.3mm, SEM mount, same
data as above.
DISTRIBUTION. North East Coast, Central East
Coast, Tasmanian Coast and S. Gulfs Coast (Fig.
4). QLD: Lizard Is., Myora Bight in Moreton
Bay, on sand (Hale, 1945a:180); sites 15, 28. and
31 in Moreton Bay. Tas: Wynyard, Fossil Reef,
on stones (Hale, 1936b:417). SA: Sellicks Beach,
Gulf St. Vincent, on stones (Hale, 1936b:417].
REMARKS. P. poecilotum from Moreton Bay
matches the types (SAMC2006) from Tasmania.
Standard lengths of adult and subadult ? 9 are
1.3 and 0.95mm respectively, smaller than those
from Tasmania (Hale, 1936b).
Both sexes are described (9? Hale;1936b; d
Hale,1945a). Hale expressed some uncertainty
about the fact that adult 9 ? could be up to 40%
larger than adult d 8 , and that 9 9 from Moreton
Bay have the carapace less swollen than the type
9. He also pointed out (1936b:417) that, al-
though the species was by far the commonest
cumacean at Sellicks Beach, SA, all were imma-
ture. The ? s he examined from Moreton Bay had
‘the carapace less swollen than in the type’
(Hale,1945a:180), and the d lacked sensory fila-
ments on the first antenna that occurred on the
same appendage of immature ¢ specimens from
SA. It appears that he has combined two species
into his description of P. poecilotum, the d being
of a different species. Both species commonly
occur in parts of Moreton Bay. P. poecilotum
(Fig. 68) has a shorter, more anteriorly tapering
carapace, longer uropodal peduncles and deeper
textured integument than P. crudgingtoni (Fig.
69).
FIG. 68. Picrocuma poecilorum Hale. A,B, whole mount 9 , showing relative lengths of carapace and somites.
A, LV. B, DV. C, carapace and pereion ? LV, shows arched dorsum and relative lengths of pereionites. D,
carapace and pereion 9 ALV, shows arched dorsum and anteriorly tapering pseudorostral lobes. E. whole mount
d DV, shows relative lengths of carapace and somites. F, uropods d DV, shows relative lengths of telsonic
somite, peduncle and rami. G, uropods $ DV, shows relative lengths of telsonic somite, peduncle and rami. H,
uropod rami $ DV, shows spination and surface texture of rami.
473
BODOTRIIDAE OF MORETON BAY
474
Picrocuma crudgingtoni sp. nov.
(Fig, 69)
MATERIAL EXAMINED, HOLOTYPE
QMW 20556, adult G, S.L. 1.1mm, in 70% ethanol,
Hays Inlet, site 17, 27*17'S, 153°09"E, D. Tafe, 12 Oct.
1985, 2m, silt, 33 ppt salinity, 24°C water temperature.
PARATYPES QMW20557, ovig. 9, allotype, S.L.
1.3mm, in 70% ethanol, same data as holotype,
QMW20558, adult d, S.L. 1.2mm, SEM mount,
Pumicestone Passage, site 1, 26°55°S, 153705 E, J.
Greenwood, 14 Dec. 1990, 1.2m, silt, 33.5 ppt salinity,
29°C water temperature; QMW2(559, adult 9, S.L.
1.2mm, in 70% ethanol, same dala as above,
DESCRIPTION. MALE. Integument lightly cal-
cified, composed of triangular, overlapping
scales giving à smooth appearance (Fig. 69).
Carapace 0.3 S.L. and 0.6 times as wide as long
m dorsal view. with lateral margins evenly
rounded, widest al midlength; depth 0.5 times
length in lateral view with dorsum smoothly ar-
ched, no median dorsal or lateral carinae; anten-
nal notch shallow, no antennal tooth or ridge;
pseudorostral lobes wide, roundly truncate, jom-
ing for a distance in front of ocular lobe equal to
0.15 times length of carapace; ocular lobe twice
as wide as long (Fig. 69A-E). Pereion 0,9 times
length of carapace; 5 percionites exposed, first
narrows laterally, second is longest, twice as long
as third or fourth, fifth longer than fourth, third to
fifth tapering in dorsal view (Fig. 69A-E). Pleon
and telsonic somite 0,45 S.L. and 1.5 times length
of carapace; robust structure, no median dorsal or
lateral carinae: no Jateral articular processes; first
4 pleonites increasing slightly in length, fifth
pleonite 1.5 times as long as fourth; telsonic
somite 1.3 times wider than long, posteriorly
rounded and slightly produced (Fig. 69A-F). First
antenna as in d of P. poecilolum, 3-segmented
with terminal flagellum; first segment somewhat
geniculate, as long as second and third segments
combined; second segment subequal in length to
third, which bears a group of short setae distally;
flagellum segment with a group of 7 sensory
filaments distally, All pereiopods 7-segmented,
except pereippod 2 which has no distinct ischium,
Structure of pereiopods as in d of P. poecilalum
MEMOIRS OF THE QUEENSLAND MUSEUM
(Hale, 19452, p.179). Peduncle of uropxxd. 1.4
times length of telsonic somite, very broad, its
width 0.45 times its length, and without spination;
rami short and robust; endopod 0.85 times length
of peduncle and slightly longer than exopod, with
5-6 inner marginal and 2 unequal terminal spine-
like setae. the longer 0.35 times ramus length;
exopod with 2 unequal terminal spine-like setae,
the longer 0.4 times ramus length (Fig. 69F,H).
OVIGEROUS FEMALE, Integument lightly cal-
cified, composed of smooth triangular, overlap-
ping scales, as in d.
Carapace length 0.25 S.L. and 0.7 times as wide
as Jong in dorsal view with lateral margins evenly
rounded and widest at posterior of length; depth
0.6 limes length in lateral view with dorsum
smoothly arched, no median dorsal or lateral
carinae; antennal notch shallow, no antennal
tooth or ridge; pseudorostral lobes wide, roundly
Inincate, joining for a distance in front of ocular
lobe equal to 0.2 times length of carapace; ocular
lobe and pseudorostral lobes as in &. Percion as
long as carapace; 5 pereionites exposed, first
narrows laterally, second is longest, twice as long
as third or fourth, fifth longer than fourth, third to
fifth tapering in dorsal view. Pleon and telsonic
somite 0.5 S.L. and 1.7 times length of carapace;
robust structure, no median dorsal or lateral
earinae; no lateral articular processes; first 4
pleonites increasing slightly in length, fifth
pleonite 1.5 times as long as fourth; telson
somite 1,3 times wider than long, posteriorly
rounded and slightly produced. First antenna 3-
segmented, with terminal flagellum; first seg-
ment somewhat geniculate, às long as second and
third segments combined; second segment sube-
qual in length to third, which bears a group of
short setae distally; flagellum with à group of 7
sensory filaments distally. Pereiopods 7- seg-
mented, except pereropod 2 which has no distinct
ischium, Structure of pereiopods same as in d.
Uropod as in d except peduncle is 1.3 times
length of telsonic. somite and inner margin of
endopod has 3 spine-like setae.
Colour. White with mottled brown
chromatophores,
FIG. 69, Pi¢rackma eridgingroni sp, nov, d. A, whole mounts DV & LV. B, whole mount LV, shows relative
lengths of carapace and somites. C, whole mount DV, shows body tapering from mid-region of carapace to
pleonite 4, D, carapace und pereionites 1-5 LV, shows dorsum smoothly arched dorsum, shallow antennal notch
and absence of lateral carinae. E, carapace and pereionites 1-4 DV, shows relative lengths of percionites in
dorsal view and absence of median dorsal ridge . F, uropod DV, shows relative lengths and widths of telsonic
somite, peduncle and rami. G, anterior carapace DV, shows pseudorostral lobes wide, roundly truncate, joining
for a distance in front of ocular lobe. H, uropod rami DV, shows spination and 1exture of rami.
BODOTRIIDAE OF MORETON BAY
—— 166rm
| ms
475
MEMOIRS OF THE QUEENSLAND MUSEUM
476
BODOTRIIDAE OF MORETON BAY
S.L. Adult d 1.0-1.2mm. Adult 9 1.1-1.3mm.
HABITAT AND DISTRIBUTION. Most com-
mon over fine sand in 1-4m of water; from all
sites in Pumicestone Passage, and sites 16, 17,
and 31 in Moreton Bay; abundant at sites 1, 2, 3,
9, 10, and 11 in Pumicestone Passage.
REMARKS. P. crudgingtoni resembles P.
poecilotum, but lacks the acutely pointed
carapace and pronounced dorsal hump. The
hump is located in the region of pereionite 2 of
the latter species, which is three times longer than
pereionite 3 and half as long as the carapace (Fig.
68C,D). In P. crudgingtoni it is only twice as long
as pereionite 3 and 1/4 as long as the carapace;
the peduncle of the uropod is about twice as long
as wide (Fig. 69F), compared to at least three
times as long as wide for P. poecilotum (Fig.
68F,G). The armature of the uropodal endopod is
also different in the new species.
The carapace of P. crudgingtoni has a smooth
appearance, being composed of flattened trian-
gular, overlapping scales (Fig. 69G). In contrast
the carapace of P. poecilotum is composed of
rounded, overlapping scales giving a roughened,
pebbled appearance (Fig. 68C,D).
P. crudgingtoni has a small, streamlined but
robust exoskeleton which, like that of P.
poecilotum, is well adapted to living amongst
sand grains in areas subject to currents or wave
action. Both species are very common in silt/fine
sand areas of Moreton Bay subject to tidal cur-
rents.
ETYMOLOGY. For Robert Crudgington, who as-
sisted in the field.
Pomacuma Hale, 1944
Pomacuma Hale, 1944b:241.
GENOTYPE. Pomacuma cognata Hale, 1944b.
DIAGNOSIS. FEMALE. Carapace with pseudo-
rostral lobes extending in front of moderately
477
large ocular lobe and meeting in midline; anten-
nal notch closed but not fused. Five pereionites
exposed; first pereionite short. Pleon longer than
carapace and pereion combined; telsonic somite
well produced posteriorly with distal margin
rounded. Third maxilliped with well-developed
exopod. First 3 pereiopods with well-developed
exopods; pereiopod 4 with rudimentary 1-seg-
mented exopod bearing few setae. Basis of
pereiopod 1 widened distally with large lobe on
distal end; lobe produced to articulation of is-
chium and merus. Carpus of pereiopod 2 much
shorter than merus. Endopod of uropod 2- seg-
mented, distal segment very short; medial margin
of exopod with plumose setae.
MALE. Second antenna reaching to end of pleon.
Thoracic exopods as in 9 . Five pairs of pleopods.
REMARKS. This genus is related to Zenocuma
but differs in structure of basis of third maxilliped
and first pereiopod, and in pseudorostrum and
telsonic somite. Two species of Pomacuma have
been previously recorded off eastern Australia.
KEY TO AUSTRALIAN SPECIES OF
POMACUMA
1, Pleon ridged. i. 1. aes eden oo ge 8d 2
Pleon not ridged
2.Sides of carapace tapering anteriorly in dorsal
view. Pereionite 4 with only posterior overlap-
ping lobe in lateral view cognatum Hale
Sides of carapace almost parallel in dorsal view.
Pereionite 4 with anterior and posterior overlap-
ping lobes in lateral view ....... sp. nov. 1
Pomacuma australiae (Zimmer, 1921)
(Fig. 71A-D)
Vaunthompsonia australiae Zimmer, 1921:4, figs 1-7.
Leptocuma australiae; Hale, 1936b:408.
Pomacuma australiae Hale, 1944b:244, figs 12-14.
Hale, 1949a:110.
MATERIAL EXAMINED. SAMC2480,3 3, 9, S.L.
8.7-9.0mm, Fraser Island, Qld (Hale,1944b);
FIG. 70. A-D, Pomacuma sp. nov. |. A, d DLV, whole mount shows dorsal and lateral carinae extending along
pereion and pleon. B, d LV, whole mount showing relative lengths of carapace and somites. C, 6 DLV, carapace
truncate anteriorly and almost parallel sided; median dorsal ridge pronounced. D, d LV, carapace and
pereionites 1-4 showing basis of pereiopod 1 widened distally, pereiopod 4 with lateral lobes which overlap
pereiopods 3 and 5. E-G, Cyclaspis alveosculpta sp. nov. E, subadult d DLV, showing strong transverse ridges
and reticulate pattern of pitting, F, subadult d VV, pleonite 5 showing immature pleopod 5. G, subadult d
ALV, showing high arches of posterior transverse ridge,and strong median dorsal and dorso lateral ridges. H,
Gephyrocuma sp. nov. 1, d, DV, detail of integument anterolaterally showing a sensory filament, protruding
from between cuticular plates.
478 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 71. A-D, Pomacuma australiae. A-C, 9. A, LV. B, cephalothorax, DV. C, uropod, DV. Ci, distal ends of
rami. D, d uropod, DV. Di, distal ends of rami. E-G, Pomacuma cognatum, E-G, type ovig. 9. E, LV. F,
anterior portion of carapace, DV. G, uropod, DV. Gi, distal ends of rami, (All after Hale, 1944b).
BODOTRIIDAE OF MORETON BAY
QOMW 20560, adult d , S.L. 8.5mm, in 70% ethanol, off
Coochiemudlo Island, site 34, 27°32'S, 153°20'E, D
Tafe, 17 June 1990, 7.5m, silt, 33 ppt salinity, 20°C
water temperature. QMW20561, adult 9, S.L. 8.1 mm,
in 70% eihanal, data as above,
DISTRIBUTION. Central and Lower East Coast,
Central and Lower West Coast (Fig.4). Old:
Fraser Island (Hale, 1944b:246); sites 31 and 34
in Moreton Bay. NSW: Jibbon, Wata Mooli,
Eden, Ulladulla (Hale, 1944h:246). WA: Cape
Jaubert, Shark Bay (Zimmer, 1921;4),Uncom-
mon in Moreton Bay.
REMARKS. Moreton Bay specimens match the
types. Two adults and à juvenile were taken by
sledge-net during the day and are slightly smaller
butotherwise the same as those deseribed by Hale
(Fig. 43A-G). The standard length of the & is
8.5mm (cf Hale, 1944b: 9.0mm) and the 9 is
8.1mm (cf Hale, 1944b: 8.7mm).
Pomacuma sp. nov. |
(Fig. 70A-D)
MATERIAL EXAMINED, QMW20562, subadult d,
S.L. 4.7mm, SEM mount, Raby Bay, site 32, 27° 30'S,
153° 18'E, D. Tafe, 17 June 1990, 1300 hours, sledge-
net, Sm, sandy mud, 33.8 ppt salinity, 18.7°C water
temperature. Uncommon,
REMARKS. Further specimens are required to
describe the species. The Moreton Bay specimen
is subadult but totally unlike any other species of
Pomacuma described to date. It bears. little
resemblance to the other two Australian species,
P. australiae and P, cognatum, P. cognatum was
recorded from the same region hy Stephenson et
al. (1978) and Stephenson (19805); the
anterolateral angle of its carapace is quite dif-
ferent to that of Pomacuma sp. nov. 1. The
carapace of the new species has a strong median
dorsal ridge extending from the ocular lobe al-
most to the first pereionite (Fig. 71C). The ocular
lobe is wider than long with pseudorostral lobes
meeting in front of it. Perciónites 2-4 are virtually
as Wide as the carapace in dorsal view (Fig. 70C).
The pereionites and pleonites have obvious
lateral and dorsal carinae. (Fig. 70A-D). The
uropod has a peduncle at least twice as long as the
rami. The anterolateral margins of pereionites 2
and 3 are wilhout spine-like setae. Pereiopod 1
robust with the basis widened distally and extend-
ing to The anterolateral curvature of the carapace:
it bears a forwardly directed Jobe on the distal
end, typical of the genus.
476
DISCUSSION
This study shows the Bodotriidae to be the
dominant family in Moreton Bay, both in terms
of the number of species and the abundance of
individuals. This is consistent with findings on
the E, W and S coasts of Australia. Hale (1937,
1943, 1945b, 19532) stated that, with regard to
the number of species, the bodotriids outnumber
the other families on the Indian Ocean and
southern Australian coasts, but on the Pacific side
the diastylids are equally well represented. How-
ever, his Diastylidae included the Gynouras-
tylidac. There is little doubt the Bodotriidae is ihe
most diverse family on the Pacific coast of
Australia.
The Bodotriidae is represented by 59 species
off the Pacific coast at latitudes of less than 35°S,
compared to 19 species in the Southern Ocean
(Bacescu, 1988). It appears from cumaccan
records m other parts of the world (Sars, 1865;
Calman. 1904, 1907, 1911, 1912; Zimmer, 1921;
Stebbing, 1912, 1913; Bacescu, 1988; Day, 1975,
1978a; Ledoyer, 1993) that the family is nega-
lively amphipolar. No species of Bodotriidae are
yet known from latitudes greater than 70° and
only 6% of records are from latitudes greater than
50° (Day, 1978a,b). Brandt (1993) and Ledoyer
(1993) showed that cumaceans are abundant on
the sea floor in both Arctic and Antarctic seas,
Ledoyer (1993) recorded population densities of
74 individuals per cubic metre in the Weddell
Sea between depths of 200-1200m; however, he
did not record any bodotriids,
Endemism is very high within the Bodotnidae,
Less than half (42%) of the 81 species recorded
around the Australian coastline occur in waters of
more than one coastal zone (Fig. 4), and only 17%
ure recorded in both Indian and Pacific Oceans
(Baceseu, 1988), The two subfamilies of
Bodotriidae recorded in Australian waters are
disproportionately represented. The Bodotrunie
exhibit a much higher diversity of species (54
species) and a much higher rate of occurrence
than do the Vaunthompsoeniinae (27 species), lar-
gely due lo the high diversity of Cyclaspis.
The greatest worldwide diversification of
species within the Bodotriidae occurs in southern
tropical und temperate latitudes (Diy, 1975,
19782). JF it were not for the large number of
species of Cyclaspis in Australasia, the
Bodotriinae would be a mainly tropical group.
The Vaunthompsoniinae is mainly a temperate
ARN
group, with 485% of species occurring between
20° and 50°S (Day, 19783).
The vast majority of known species (87%) of
Bodotriidae are from depths of less than. 200
metres, indicating that ihe family belongs
predominantly to the shelf fauna. A number of
new species of Bodotriidae and other families
have been recorded from depths greater than
200m (Jones & Sanders, 1972: Day, 1978a;
Gama. 1989; Roccatagliata 1989: Jones. 1990;
Watling, 1991a,b; Roccatagliata & Heard, 1992;
Reccutagliata, 1993), indicating that the apparent
lack of deep water species is at least partly due to
scarcity of collecting.
In South African waters Ihe numbers of in-
dividuals are highest within the Bodotriinae but
diversity within this subfamily is much lower
than in Australia (Day, 1978a). The ecological
reasons for this difference is not clear, but the
immediate cause, according to Day (19784), is
the effective colonisation of South African cdas-
tal waters by just a few species. Iphinoe africana
and J, stebbingi account for over half the in-
dividuals of the Bodotriinue taken from around
South Africa, with a further 10
species nf this genus being recorded from 1he
region. In Australia /phinoe has not gained as
great a foothold, the only recorded species being
i. pellucida, off NSW and Tasmania.
Cyclaspis has been highly successful in
colonising Australian coastal waters. It accounts
for 60% of bodotriid species, and is widespread
in its. distribution, Hale (1953a) found that
Cyclaspis individuals were the most abundant in
light-trap catches off Garden Island, W.A. He
encountered them everywhere where there was a
sandy substratum. Bacescescu (1990, 1992b)
described 4 new species of Cyclaspis from the
northern waters of Australia. In the present study
Cyclaspis predominated in catches over sand and
silUmud substrata, but Nannastacus was more
common in seagrass beds (Hale, 1949b, 1953).
Males of both of these genera by far outnumbered
9 9 in Hale's light- trap catches, as in the present
sledge-net catches. It appears that d d. of these
two genera are much more active in the water
column at night than are 9 9 ,
Some species of Bodotriidae were highly
restricted in their geographical distributions
within Moreton Bay. Gephvrocuma repandum,
the most abundant species by far at the northern
entrance to Pumicestone Passage, was not
recorded elsewhere in Moreton Bay or in the
upper estuary. Leprocuma barbarae, the most
abundant species in Horseshoe Bay, was not
MEMOIRS OF THE QUEENSLAND MUSEUM
recorded 8km to the W in Raby Bay. Cyclaspis
tranteri the most abundant species in Raby Bay,
was not recorded in Horseshoe Bay even though
over 200 samples were taken there spanning all
seasons. Such findings lend support to the view
that cumaceans generally have restricted disper-
sal pattems compared to other peracarids,
The richest areas of those sampled in Moreton
Bay were generally the shallow water, marine
arcas with sandy substrata. High numbers of in-
dividuals were consistently taken after sunset in
such areas, using à sledge-net sampler at times of
slack nde. Despite this the single largest sample
was taken at slack high water over a mud sub-
stratum. This was an unusually large haul (87
cumaceans per m-4) for the middle of the day. It
was mainly due to a swarm of Cyclaspis franteri
which accounted for 92% of cumacean numbers
taken. Even more unusual was the fact that jhe
haul was taken over a mud substratum. Repeated
samplings al the same location (site 32) on other
occasions failed to yield numbers of such mag-
nitude, Similar sledge-net hauls over sandy sub-
strata often yielded catches of at least 50
cumaceans per m?, when taken at night on a slack
ride.
ACKNOWLEDGEMENTS
We gratefully acknowledge Barbara Koh and
Robert Crudgington for assistance in the field,
Joan Greenwood for assistance in the laboratory
and Lilli Staheli for assistance with the manu-
script. Critical reviews by Les Watling, Darling
Marine Center, University of Maine, and Peter
Jell and P. Davie. Queensland Museum, have
greatly improved the manuscuscript. In particular
we would like to thank the Queensland Museum,
for financial support to the senior author during
final preparation of the manuscript. The Zoology
Department, University of Queensland, is
thanked for supplying sampling equipment and
funding. Finally, the fieldwork and electron
microscopy could not have been undertaken
without the significant financial suppart of Reck-
itt & Colman Pty Ltd-
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ANNUAL PATTERN OF ACTIVITY OF THE BROWN TREE SNAKE (BOIGA
IRREGULARIS) IN SOUTHEASTERN QUEENSLAND
KARINA H. BULL AND JOAN M. WHITTIER
Bull, K.H. & Whittier, J.M. 1996 07 20: Annual pattern of activity of the Brown Tree snake
(Boiga irregularis) in southeastern Queensland. Memoirs of the Queensland Museum 39(2):
483-486. Brisbane. ISSN 0079-8835.
Dates of collection of Boiga irregularis from SE Queensland in the Queensland Museum
indicate adult snakes were most commonly collected in January, February and March, the
warmest and wettest months. Juvenile snakes were less numerous than adults, but were
represented in all months of the year, with peaks in January and April. Few females were
collected during final stages of egg development, and few gravid females are represented.
These observations support field observations that suggest adults are most active in the warm,
wet summer months and that females are relatively inactive during late ovarian development
and when gravid. O Seasonality, Reptilia, Colubridae, Boiga, introduced species,
Queensland.
Karina H. Bull and Joan M. Whittier, Centre for Conservation Biology and Department of
Anatomical Sciences, University of Queensland, Queensland 4072, Australia; 20 June 1996.
The activity of reptilian species may be con-
fined to certain months of the year. In tropical
regions, where temperatures are relatively warm
and stable throughout the year, many species are
active all year round. However, some reptiles in
these areas have depressed activity during the
tropical *winter'. This is believed to be a conse-
quence of the low levels of humidity related to the
dry season, rather than a consequence of tempera-
ture (Shine, 1991a). Even in temperate regions,
high humidity seems to stimulate greater snake
movement. The apparent relations of snake ac-
tivity to seasonal changes in temperature and
rainfall have prompted this investigation into the
annual activity cycle of the arboreal brown tree
snake, Boiga irregularis, in SE Queensland.
B. irregularis is a colubrid from coastal
Australia, Papua New Guinea and islands in NW
Melanesia and Indonesia (Cogger, 1992). On
Guam (13?N), where it has been introduced, the
snake causes a major impact on the power supply
of the island by climbing on electrical wires and
producing electrical shorts (Fritts et al., 1984).
The Naval Public Works Command maintain a
log of these occurrences, which has allowed
evaluation of activity patterns of the snakes over
several years. The seasonal incidence of B. ir-
regularis is linked with monthly rainfall similar
to other arboreal tropical snakes (Henderson et
al., 1978). Peak activity occurs in May, June and
July, the months that generally show an increase
in rainfall after 7 months of steadily declining
precipitation (Fritts et al., 1984).
Although activity of B. irregularis in
Australasia has not been documented it has been
inferred from study of its reproductive habits
(Shine, 1991b; Whittier & Limpus, 1996). During
late spring and summer B. irregularis is en-
countered in SE Qld, including urban areas. How-
ever, in the winter it is more secretive. Being
nocturnal and arboreal means that the behaviour
of B. irregularis is likely to be strongly influenced
by seasonal changes in the weather. It is a ther-
moconformer during its activity period.
To observe general fluctuations in B.
irregularis" annual activity pattern, snakes were
examined from the Queensland Museum (Appen-
dix 1). This study was based around the general
premise that the seasonal distribution of the
species in the museum's collection is repre-
sentative of the abundance of B. irregularis in
natural populations. Therefore, fluctuations in the
number of museum specimens over the year
would indicate changing activity levels of the
snake. Collections of B. irregularis from PNG
and the wet tropics of Australia differ in being
collected as biased samples on specific field trips.
SE Qld specimens have been obtained generally
from casual encounters, usually by the general
public (J. Covacevich, pers. comm.). In the
Queensland Museum's collection, reliable col-
lection dates (as opposed to registration dates)
began in the 1970's.
Observations of B. irregularis and by inference
its activity level changes through the year (Figs
1-2). Adult snakes (>75cm) are most numerous
in January, February and March. These months
are both the warmest and wettest (Fig. 3). Adult
snakes are least frequent during May-September,
the cooler, drier months of the year. Numbers of
INDIVIDUALS CAPTURED
484
MEMOIRS OF THE QUEENSLAND MUSEUM
Adults
Months
[] Juveniles
FIG. 1. Monthly incidence of adult and juvenile specimens of Boiga irregularis from SE Qld.
snakes only start to significantly increase in Oc-
tober, the first month of increased precipitation
after 7 months of steadily declining rainfall.
Temperatures begin their upward turn in Septem-
ber, a month earlier than the end of the dry season.
Juvenile snakes («75cm) are less numerous
than adults, however juveniles experience less of
a decline in activity over winter, relative to their
numbers (Fig 1). They persist at a more or less
stable level throughout the year, although are
most abundant in January and April.
Males are always more common than females
in all months of the year (Fig.2). Both sexes
experience declines in activity during April-Sep-
tember, however male snakes appear to emerge
prior to females; greater numbers of males are
found in October but greater numbers of females
are found in December. Females also are under-
represented in February, and are found in higher
numbers in late summer/autumn in March and
A pril.
Seasonal fluctuations in the activity of B. ir-
regularis in SE Qld appear linked with tempera-
ture and rainfall. When minimum temperatures
fall below 15?C, snakes reduce their level of
activity. These minimum temperatures represent
the night time temperatures that occur when these
nocturnal snakes are foraging and moving about.
Although temperatures begin to increase in Sep-
tember, no significant increase in snake activity
is observed until October. This may be due to the
snake's minimum temperature threshold; that is,
the need for temperatures above 15°C to stimulate
activity. Otherwise, the onset of rains in October
may explain appearance of snakes in this month.
Sexual differences in activity levels of this
snake tend to skew the sex ratio of captured
snakes. This is probably due to the more secretive
nature of females, especially those in the final
stages of egg development (September-Decem-
ber) and egg laying (particularly February).
Males also tend to be more active in searching for
mates and as a consequence may travel further
distances than females and appear more active.
The large increase in male activity in October
may reflect this, as October is the time when
spring matings are known to occur (Whittier &
Limpus, 1996).
The temporary disappearance of females in the
population during February is likely to be due to
the occurrence of egg laying during this month.
Females must be clandestine at this time, as very
few are captured with enlarged ovulated follicles
( from 227). Following egg laying, females may
INDIVIDUALS CAPTURED
SEASONAL ACTIVITY PATTERN OF THE BROWN TREE SNAKE
485
Months
O Females
FIG. 2. Monthly incidence of male and female specimens of Boiga irregularis from SE Qld.
become active again, feeding to gain fat stores for
winter dormancy. Similar observations of inac-
tivity in gravid females have been made in Guam
(R.T. Mason pers. comm.)
As incubation takes a minimum of 65 days in
B. irregularis (Shine, 1991b), the small increase
in juveniles in May could represent the emer-
gence of hatchlings. These juveniles must grow
and maintain themselves in the months following
hatching and, as this does not occur until autumn,
it explains the persistence of juveniles over the
winter months.
The increase in acquisitions during July is mis-
leading as it is caused by a group of 3 adult males
and 2 juvenile males found in a winter aggrega-
tion. They were among 30-40 specimens of B.
irregularis, Dendrephalis punctulatus and
Morelia spilota variegata (Covacevich & Lim-
pus, 1973). This winter aggregation of mixed
species appeared not to be associated with breed-
ing but with thermoregulatory behaviour
(Covacevich & Limpus, 1973).
ACKNOWLEDGEMENTS
We thank Patrick Couper, Jeanette Covacevich
and Glen Ingram, Queensland Museum for their
enthusiastic support of our investigations. We
also thank R.T. Mason for information about the
Guam snakes.
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petofauna 6(2): 16-20.
FRITTS, T.H., N.J. SCOTT & SAVIDGE, J.A. 1987.
Activity of the arboreal Brown Tree Snake (Boiga
irregularis) on Guam as determined by electrical
outages. Snake 19: 51-58.
HENDERSON, R.W., J.A. DIXON & SOINI, P. 1978.
On the seasonal incidence of tropical snakes. Mil-
waukee Public Museum Contributions in Biology
and Geology 7: 1-15.
SHINE, R. 1991a. Australian Snakes: A Natural His-
tory. (Reed Books: Sydney).
1991b. Strangers in a strange land: Ecology of the
Australian colubrid snakes. Copeia 1991: 120-
131,
WHITTIER, J.M. & LIMPUS, D. 1996. Reproductive
patterns of a biologically invasive species: the
brown tree snake (Boiga irregularis) in eastern
Australia. Journal of Zoology (London) 238: 591-
597.
486 MEMOIRS OF THE QUEENSLAND MUSEUM
200
100
AVERAGE MINIMUM TEMPERATURE ( C)
AVERAGE RAINFALL (mm)
J F M A M J J A S Oo N D
----- t--- Min. Temp. Months ——t"—- Av. Rainfall
FIG. 3. Monthly mean minimum temperature (°C) and mean rainfall (mm) for Brisbane.
APPENDIX 1. Queensland Museum QMJ30965; QMJ31853; QMJ32238;
specimens of Boiga irregularis examined for this QMJ32247; QMJ32266; QMJ35332;
study. QMJ36056; QMJ36100; QMJ36754;
QMJ11181; QMJ11269; QMJI1337; QMJ36961; QMJ37146; QMJ37174;
QMJ11495; QMJ11577; QMJ13512; QMJ38166; QMJ40003; QMJ40278:
QMJI4315; QMJI4503; QMJ14744; QMJ40292; QMJ40293; QMJ40936;
QMJ15554-QMJ15557; QMJI6104; QMJ40949; QMJ41383; QMJ41393;
QMJI8IO!; QMJ18532; QOMI20295; Qwr44240: (QMJA46579; QMJ46860.
ume Motori TAES QMJ47015: | QMJ47312-QMJ47314.
QMJ22716; QMJ23163; QMJ23774; QMJ47320; QMJ47384; QMJ47396;
QMJ25382: QMJ25523: QMJ26906. QMJ47920; QMJ48104; QMJ48121;
QMJ28388; QMJ28392: QMJ28393; QMJ48128; QMJ48582; QMJ48599;
QMJ28477; QMJ28865; QMJ29007; QMJ48606; QMJ49851; QMJ49894;
QMJ30284; QMJ30960; QMJ30963; QMJ49906.
MEMOIRS OF THE QUEENSLAND MUSEUM
NEPHRURUS ASPER (SQUAMATA: GEKKONIDAE):
SPERM STORAGE AND OTHER REPRODUCTIVE
DATA. Memoirs of the Queensland Museum 39(2); 487.
1996:- Data on size at maturity, sexual dimorphism and
seasonal reproductive patterns for Nephrurus species were
provided by How et al. (1990). They examined over 1000
specimens, of which 70 were assigned to N. asper. This taxon
has since been shown to be a composite of three species, N.
asper Günther, N. amyae Couper and N. sheai Couper (Couper
& Gregson, 1994),
The N. asper specimens examined by How et al, were from
the collections of the Western Australian, South Australian
and Northern Territory Museums, and are largely N. amyae
and N. sheai (Couper & Gregson, 1994), Similarly, reproduc-
tive records for N. asper (Gow, 1979; Bedford & Christian,
1993) also relate to N. sheai and N. amyae, respectively, based
on locality. Wagner & Lazik (1996) provided reproductive
data for N. asper and N. levis. From their account it remains
unclear whether their N, asper colony contains N, asper s.s.,
the recently described taxa (Couper & Gregson, 1994),
hybrids, or a composite of two/three prickly knob-tail species,
N. asper s.s. is confined to Qld (Couper & Gregson, 1994).
Two specimens collected on 25 April 1995 from Dipperu
National Park (21?53'53"S, 148?43'03"E), 130km from the
type locality, provide knowledge of reproduction in N. asper.
The following observations confirm repetitive clutches for
this species (found in other Nephrurus species by How et al.,
1990, based on the simultaneous presence of oviducal eggs
and yolking ovarian follicles, and in *N. asper' and N. levis by
Wagner & Lazik (1996) and record sperm storage for the first
time (previously inferred for only N. milii by How et al.
(1990), on the basis of asynchrony between peak testis size
and vitellogenesis).
The Dipperu specimens were transferred to Brisbane and
housed in a glass tank (76 x 30cm). The two geckos avoided
contact, and always used separate sheltering sites, The female
appeared to be dominant and would frequently displace the
male from his sheltering site, forcing him to seek alternate
cover. The close confinement of these geckos was evidently
stressful to the male, which was found dead on 6 Sept. 1995
(QMJ60375). On the morning of 4 Dec. 1995, the female
remained in the open during daylight hours, and excavated a
cavity in the moist soil surrounding the water bowl, Two eggs
were laid. These were immediately removed and placed in
moist vermiculite. The female retreated to cover when the
eggs were removed, but emerged at night to fill in the nest
hole and excavate an extensive cavity beneath a paving stone
in the corner of her tank. The eggs measured 31.03x17.17mm
(QMJ61614) and 30.04x17,18mm. Only the second of these
survived incubation to hatch on the 28 March, 1996, Incuba-
tion took 115 days at a max/min temperature range of 31°-
21°C. The hatchling measured 46.29mm (SVL) and weighed
3.2g. On the afternoon of 26 Jan. 1995, the female laid a
second clutch of two eggs. These also were transferred to
487
moist vermiculite for incubation. The second clutch of eggs
was smaller than the first, measuring 26.68x15.64mm and
26.21x15.10mm and weighing 3.7g and 3.3g, respectively.
The female weighed 28.5g after the eggs were layed. The
relative clutch mass (RCM (1) after Greer, 1989) equalled
24.696. The second clutch hatched on 2 June 1996. Incubation
took 129/130 days at the same max/min temperature range as
that for the first clutch. The hatchlings measured 44.76mm
and 46.40mm (SVL) and weighed 2.5g and 3.0g, respectively.
These observations provide data on egg and hatchling
sizes, and incubation period for N. asper. The recorded RCM
(1) for N. asper (24.696) corresponds closely with that of N.
amyae (24.8%, Bedford & Christian, 1993). Given that the
male N. asper died 140 days before the female produced her
second clutch of eggs, it is apparent that females of this species
have the ability to store sperm. As the male was obviously
stressed, and actively avoided the female, it seems likely that
the female was already carrying sperm from a copulation prior
to her capture (277+ days prior to laying the second clutch of
epgs).
The specimens on which these observations are based were
collected during à survey of reptiles of the Brigalow
Biogeographic Region in Queensland for the Endangered
Species Program of the Australian Nature Conservation
Agency, Canberra. I thank Jeanette Covacevich and Glenn
Shea for suggesting improvements to the manuscript.
LITERATURE CITED
Bedford, G. & Christian, K. 1993. Egg size of the Prickly
Knob- tailed Gecko (Nephrurus asper, Günther, 1876)
with a preliminary comparison of investment per
progeny among geckos. Dactylus 1(4):38-41.
Couper, P.J, & Gregson, R. A.M. 1994, Redescription of
Nephrurus asper Günther, and description of N, amyae
sp. nov. and N. sheai sp. nov. Memoirs of the
Queensland Museum 37(1): 53-67.
Gow, G. 1979. Notes on the biology of Nephrurus asper.
Northern Territory Naturalist 1(2): 19-20.
Greer, A.E. 1989. The biology and evolution of Australian
Lizards, (Surrey Beatty & Sons: Chipping Norton).
264p.
How, R.A., Dell, J. & Wellington, B.D. 1990. Reproductive
and dietary biology of Nephrurus and Under-
woodisaurus (Gekkonidae) in Western Australia.
Records of the Western Australian Museum 14(4):
449-459.
Wagner, E. & Lazik, C. 1996. Husbandry and reproduction of
Australian geckos of the genus Nephrurus. Reptiles:
guide to keeping reptiles and amphibians 4(5): 56-67.
Patrick J, Couper, Queensland Museum, PO Box 3300, South
Brisbane, Queensland 4101, Australia; 20 May 1996.
CONTENTS (continued)
HEALY, J.M. & LAMPRELL, K.L.
The Atlantic-Mediterranean bivalve, Corbula gibba (Olivi) (Corbulidae: Myoidea)
in Port Phillip Bay, Victoria... 2.2... 6... e ccc ec eee te ham gas 315
MANNING, B. & KOFRON, C.P.
Evolution and zoogeography of Australian freshwater turtles |... ciis es 319
PATERSON, R.A..& VAN DYCK, S.M.
Perinatal skeletal injuries in two balaenopterid whales, ,.......-..,-.+-+ POT ete EA 333
CLEMENTS, K.D. & RANDALL, J.E.
Four new records of surgeonfishes (Perciformes: Acanthuridae) from the Great Barrier Reef ,....,... 339
STANISIC, J, a
New land snails from Boggomoss environments in the Dawson Valley, southeastern
Queensland (Eupulmonata: Charopidae and Camaenidae) ........... 00.000 aurreri eee eee 343
STANISIC, J,
A new camaenid land snail from the Wet Tropics Biogeographic Region, northeastern Queensland
(Eupulmonata: Camaenidae), .... +... cee ee aae ak ehh beans 355
STOREY, R.I. & HOWDEN, H.F.
Revision of Australoxenella Howden & Storey in Australia (Coleoptera: Scarabaeidae: Aphodiinae). . . 365
TAFE, D.J. & GREENWOOD, J.G.
A new species of Schizotrema (Cumacea: Nannastacidae) from Moreton Bay, Queensland.........,. 381
TAFE, D.J. & GREENWOOD, J.G.
The Bodotriidae (Crustacea: Cumacea) of Moreton Bay, Queensland ............0. 20000 cece eee 391
BULL, K.H. & WHITTIER, J.M.
Annual pattern of activity of the Brown Tree Snake (Boiga irregularis) in southeastern Queensland ., . 483
NOTES
LEE, M.S.Y.
Possible affinities between Varanus giganteus and Megalania prisca... 232
COUPER, P.J., COVACEVICH, J,A. & McDONALD, K.R
A Bandy Bandy with a difference ...... 0. cence Rm 242
COUPER, P.J., COVACEVICH, J-A., MONTEITH, G.B., JAGO, K., JANETZKI, H. & ROBERTS, L.
Feeding habits of the ring-tailed gecko, Cyrtodactylus louisiadensis. ........ 0s 0b reece ee Crime 288
HERO, J.M. & FICKLING, S.
Reproductive characteristics of female frogs from mesic habitats in Queensland ...........2,5....23- 306
IRWIN, S.
Survival of a large Crocodylus porosus despite significant lower jaw loss .......... 22... oe p 338
BURNETT, S. & NOLEN, J.
Fruit eating by the gecko Gehyra dubia in Townsville. |..... i ciere na han 364
DEER, R.
Reptile diversity in a Callitris forest in central Queensland's brigalow belt ..... 2.0.2.6. .00 005-4000 390
COUPER, P.J.
Nephrurus asper (Squamata: Gekkonidae): sperm storage and other reproductive data ............-. 487
CONTENTS
ARNOLD, P.W. & HEINSOHN, G.E.
Phylogenetic status of the Irrawaddy Dolphin Orcaeila brevirostris (Owen in Gray):
KNQCWITUYUEUTS TCR ORT TTE TTT TET TII CIE oY EE es hae We ods bb ok COT OS 141
BRAILOVSKY, H. & MONTEITH, G.B.
A new species of Pomponatius Distant from Australia (Hemiptera: Heteroptera:
ORI Acanthocor e erue ueceicterk moe Xe e rkeexee nre re mer) ew» e 205
BURWELL, C.J.
Revision of the Australian genus Microtropesa Macquart (Diptera: Tachinidae: Tachinini).......... 211
CLIFFORD, H.T.
Geometrical study of a cast of Leptophloeum australe (McCoy) Walton from Queensland .......... 227
COUPER, PJ., COVACEVICH, J.A., MARSTERSON, S.P. & SHEA, G.M.
Coggeria naufragus gen. et sp. nov., a sand-swimming skink from Fraser Island, Queensland ....... 233
COVACEVICH, J.A. & COUPER, P.J.
Aspidites ramsayi (Boidae) in the Brigalow Biogeographic Region of Queensland: occurrence;
conservation status and possible bilby associations. ...........00. cece cece ene 243
COVACEVICH, J.A., COUPER, P.J. & MCDONALD, K.R
Lerista allenae (Scincidae: Lygosominae): 60 years from exhibition to extinction? ................ 247
DAVIE, P.J.F. & DE FORGES, R.
Two new species of false spider crabs (Crustacea: Brachyura: Hymenosomatidae)
from: New caledoni& «i6. cesses rem rhe Y RR RA ES EE Ea Ea HR IER ERG 257
NG, P.K.L. & DE FORGES, R.
The Hymenosomatidae (Crustacea: Decapoda: Brachyura) of New Caledonia,
with descriptions of two new genera and two new species ........0.0. 0. cece essen 263
DAVIE, P.J.F. & GUINOT, D.
Two new freshwater crabs in Australocarcinus Davie, with remarks on the Trogloplacinae Guinot
and Goneplacidae MacLeay (Crustacea: Decapoda: Brachyura). .....,..seesespessesossss 277
JAMIESON, B.G.M. & GUINOT, D.
Ultrastructure of the spermatozoon of Australocarcinus riparius
(Crustacea: Brachyura; Gonoplacidae: Trogloplacinae) 2.0.6... eussssros eeror cee eee nee 289
GARDZINSKA, J.
New species and records of Astieae (Araneae: Salticidae) from Australia and Papua New Guinea .... 297
GULLAN, P.J. & STEWART, A.C.
A new genus and species of ant-associated coccid (Hemiptera: Coccidae: Myzolecaniinae)
from: Canthium Lam, (Rübiscéeae). oo. ee sek ke be pe e hh bribra serais gi 307
(continued inside cover)
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