MEMOIRS

OF THE

EUM

MUS

QUEENSLAND

VOLUME 42

BRISBANE

PART 1

30 JUNE 1997

MEMOIRS

OF THE

(QUEENSLAND MUSEUM

BRISBANE

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SOME RECENT SPECIES OF THE GENUS ANASKOPORA WASS, 1975 (BRYOZOA:

CRIBRIOMORPHA) FROM QUEENSLAND
P.W. ARNOLD AND P.L. COOK

Arnold, P.W. & Cook, P.L. 1997 06 30: Some Recent species of the genus Anaskopora Wass,
1975 (Bryozoa: Cribriomorpha) from Queensland. Memoirs of the Queensland Museum
42(1): 1-11. Brisbane. ISSN 0079-8835.

Several species of the genus Anaskopora have small, globular colonies, composed principally
of frontally budded autozooids, kenozooids and avicularia. Hitherto, all known species have
been Tertiary fossils from Victoria. A. parkeri sp. nov. is a Recent species here described
from the Queensland continental slope, Colonies possess long rhizoids, which are inferred
to have also occurred in fossil species with the same mode of growth and colony form.
Cellepora doliaris Maplestone, 1909, a rare conical species with more regular frontal
budding, originally found from New South Wales, hàs been referred in the past to the genus
Reginella. It is. here redescribed from specimens collected on the continental slope of
Queensland, and is reassigned to Anaskapora. The characters of the genus Reginella, which
have been somewhat misunderstood, are discussed and the type species, R, furcata (Hincks,
1882), is redescribed. CT Bryozoa, Anaskopora, Reginella, taxonomy, morphology.

Peter W. Arnold, Museum of Tropical Queensland, 70-84 Flinders Street, Townsville,
Queensland, 4810, Australia; Patricia L.Cook, Associate, Museum of Victoria, Melbourne,

Victoria, 3000, Australia: 6 March, 1997.

Colonies of cribriomorph Bryozoa tend to be
encrusting and unilaminar, although a few fossil spe-
cies were inferred to have had a ‘free-living’ habit.

In this paper, we report the discovery of a new
species of deep-sea free-living cribriomorph an-
chored by rootlets. We place the new species in
the genus Anaskopora. Wass (1975) revised the
genus Corbulipora MacGillivray (1895) and de-
scribed several fossil species from the Victorian
Tertiary, some of which he referred to a new
subgenus Anaskopora. Recent revision of
Corbulipora has shown that its species occur in
multiple growth phases (see Bock & Cook,
1994), and that the species of Anaskopora are
generically distinct (see Bock & Cook, in press).

The discovery of a Recent species of An-
askopara makes a significant contribution to the
interpretation of the colony structure of fossil
species, several of which can now confidently be
inferred to have been anchored by rhizoid sys-
tems, Understanding of this mode of growth also
allows the inference that the fossil species lived
in and upon the upper layers of particulate sea-
bottoms, and were part of a 'sand-fauna' (see
Cook, 1981) of similarly adapted bryozoan species.

METHODS

Specimens were bleached in sodium hypochlo-
rite solution and coated with AuPd or carbon for
SEM. Abbreviations used: AM, Australian Mu-
seum, Sydney: BMNH, Natural History Mu-

seum, London; MOV, Museum of Victoria, Mel-
bourne; MTQ, Museum of Tropical Queensland,
Townsville; QMG, Queensland Museum, Bris-
bane,

Morphological terms used in the descriptions
are defined in Bassler (1953) and Boardman &
Cheetham (1983).

SYSTEMATICS

Class GYMNOLAEMATA Allman, 1856
Order CHEILOSTOMATIDA Busk, 1852
Infraorder CRIBRIOMORPHA Harmer, 1926
Family CRIBRILINIDAE Hincks, 1879

Anaskopora Wass, 1975

Anaskopora Wass, 1975:170. Bock & Cook 1996: in
press.

TYPE SPECIES. Cribrilina elevata MacGillivray,
1895,

DESCRIPTION. Colony encrusting, originating,
on very small substrata, often globular-to-coni-
cal. Autozooids with septulae in the vertical
walls, some or all of which become surrounded
by à partially calcified chamber with an upper,
cuticle-covered window. Zooids budded laterally
and distally, or from chambered pores (see below)
as interzooidal frontal buds. Interzooidal
kenozooids and avicularia, together with rhizoids
in some species, produced in the same way. Au-

2 MEMOIRS OF THE QUEENSLAND MUSEUM

lozooids with a costate pericyst, with lacunae:
pelmatidia rare or ubsent. Secondary, calcified
orifice with paired spines and a raised, distal,
fimbriated plate. Avicularia distal or distolateral;
occasionally proximal, rostrum rounded or sub-
triangular, with paired condyles. Ovicells un-
known.

The globular species of Anaskopora, in pårtic-
ular, possess a kind of pore chamber, here called
à ‘chambered pore’ to distinguish it from other
forms of dictellae. This originates as a calcified
chamber surrounding one or more septulae in the
vertical walls ol zooids. The chamber is uncalci-
fied on the frontal side, and the cuticle covering
this window is able to expand intussusceptively
so that the chamber may enlarge to form inter-
zooial Kenozooids, autozooids, aviculana and
rhizoids. Chambered pores occur in Cerbulipora,
but are not the regular source of frontally budded
zooids, as they are in the globular species of
Anaxkopora, Some specimens of species de-
scribed by Bock & Cook (in press), and a few of
the colonies of A. parkeri (see below), originate
upon very small shell substrata. These are rapidly
covered by encrusting zooids with chambered
pores al their comers. These pores produce an
interzooidal, frontally budded series of zooids 1n
subsequent astogeneti¢ generations, which com-
pletely supersede any further encrusting growth.
In Anaskopora doliaris. frontal budding of this
kind seems to form the entire colony, and is very
regular, so that the conical form is produced.

Anaskopora parkeri sp. nov.
(Figs 1 A.B, 2A,B, 3)

MATERIAL. EXAMINED. HOLOTYPE: QMG-
21282, in alcohol, 'Cidaris UU Sin 42.2, 17721778.
146"48,52'E, [5.v, 1986, 296-202m, epibenthic sledge
LL/2 inch inner liner), at MTQ. PARATYPES:
QMG21283, one colony mounted on SEM stub, same
data as holotype; QMG21284, one colony mounted on
SEM stub, same data as holotype; QMG21285, 55
colonies in alcohol, same data as holotype:
MOV F80820, 9 colonies in alcohol, same data as ho-
lotype. ADDITIONAL MATERIAL: QMG212B8, 71
colonies in alcohol, same data as holotype;
QNIG21289, 2 colonies in alcohol, "Cidaris 1" Stn 43.2,
VI*34,58'S, 146°53,21°E, 15,v.1986, 458-500m,
epibenthic sledge (1/2 inch liner), at MIO.

ETYMOLOGY. For the late Shane Parker, of the South
Australian Museum,

DESCRIPTION, Colonies 0.50-5. 0mm in diam-
gler, encrusting small foraminiferans or com-
pletely free-living (Fig. 1A), anchored by large
rhizoids (Fig. 1B). Autozootds with frontal shield

of 13-24 costae alternating with rows of 7-8 lacu-
nac (Fig. 2A). Calcified orifice with one pair of
flattened, sometimes terminally hifid, lateral-oral
spines (Fig. 2A), paired lateral condyles and a
raised distal plate with 5-8 fimhriations (Fig. 2B)
und a pit at the base of the outer surface (Fig. 3).
Operculum golden-brown, with a marginal scle-
rite, filling the secondary orifice, Septulae 2-3
distal, 3-4 lateral, becoming surrounded by cham-
hered pores (Fig. 3) which develop frontally as a
series of 4-6 small kenozooids with cuticular
frontals, surrounding each zooid. Avicularia disto-
lateral (Figs 2A, 3), developed from chambered
pores; occasionally paired or even proximal, Sub-
rostral chambers raised, rounded (Fig. 3); mandi-
bles with a marginal sclerite, oriented laterally.
Zooids budded in alternating interzooidal series
from chambered pores, oriented irregularly, col-
ony forming an ectoproctolith (Figs LA, 2A, 3).
Interzooidal kenozooids becoming extrazooidal.

Colony and zooid dimensions are given in
Table 1.

REMARKS. Å. parker! closely resembles A,
comuta (MacGillivray, 1895), which was re-
ferred to Corbulipora by Wass (1975) but reas-
signed to Anaskopora by Bock & Cook (in press),
The two species have similar colony structures,
and the autozooids of A. cornuta have numerous
straight rows of costae, alternating with small
lacunae, like those of A. parkeri. A. cornuta dil-
fers in the distal position of the avicularium,
which is not nearly as prominent as that of A,
parkeri, The two species are so alike that it seems
certain that the Victorian Tertiary species was a
direct ancestor of the Recent form from Queens-
land. 1t may also be inferred with a high degree
of confidence that the colonies of A. cornuta
possessed rhizoids and lived in a similar micro-
environment to (hat inhabited by A. parkert.

Of 75 colonies of A. parkeri from Stn 42.2, 20
had well developed rhizoids which were 2-3mm
long and, when turgid, 0.33-0.38mm wide. The
colonies exhibited a developmental series; the

smallest had 6 zooids, the largest approximately

50 autozooids visible at the surface. Where the
youngest stages encrusted small foraminifera
there was a large, cuticle-covered cavity next to
the primary zooid; in two cases, with a large
thizoid beside it. Whether the cuticle-covered
cavity is the ancestrula is not known, The rhizoids
themselves terminated in numerous rootlets
which had shell fragments and foraminiferans
adhering to the cuticle. One colony was the sub-
stratum for a small colony of a tubuliporid cyclo-

RECENT SPECIES OF ANASKOPORA 3

FIG. 1. A-B, Anaskopora parkeri sp. nov. A, QMG21283, frontally budded colony, x 32; B, Sketch of colony
with rhizoids, x 15. C-D, A. doliaris (Maplestone). C, QMG21286, colony from frontal side, x 21; D,
QMG21287, colony from basal side, x 31.

4 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-B, Anaskopora parkeri sp. nov. A, QMG21284, general view of zooids, x 58; B, QMG21283, enlarged
view of one zooid, x 100. C-D, A. doliaris (Maplestone). C, QMG21286, general view of zooids, x 42; D,
QMG21287, enlarged view of one zooid, x 66.

RECENT SPECIES OF ANASKOPORA 3

FIG. 3, Anaskopora parkeri sp. nov, QMG2]283. Au
tazooids showing details of orifices and frontal buds,
x 37.

stome. Two others had the horny skeleton of
Stephanoscyplius (SCyphozoa: Coronatae) grow-
ing on them; Lagaiij (1963) described a similar
case in which å hydroid grew on the globular
colonies of Fedora nodosa.

The specimens of A. parkeri were accompanied
by other rooted species associated with sand» fau-
nas. These included Parmidaria sp., Parasticho-
pora vanna (see Cook & Chimomdes, 1981) and
Conescharellina sp.

Anaskopora doliaris (Maplestone)
(Figs 1C,D, 20,D, 4A,B)

Cellepara daliaris Maplestone, 1909: 272, Pl. 77, Figs
10 A,B.

Reginella doliaris Brown 1958: 53, Hastings 1964;
254, Pl. 1, Figs 1-3. Pls 2 and 3

LECTOTYPE (indicated here); AMU201, 22 miles
(34.4km) East of Port Jackson, New South Wales, 80
fath. (146m). HMCS ‘Miner’, labelled ‘TYPE’.
PARALECTOTYPE: BMNH1909.11.12.14, as
above.

MATERIAL. EXAMINED. QMG21286, G21287,
‘Cidaris 1' Stn 432, 17734.58' S, 14675321 E,
15.v.1986, 458-500m, epibenthic sledge (1/2 inch
inner liner). 2 colonies on SEM stubs; at MTQ,

DESCRIPTION. Colony conical, one autozooid
thick (Fig. 1D), inferred to be anchored by basal
rhizoids, Autozooids erect, deeply immersed and
oriented with the distal part of the orifice towards
the ancestrula and the centre ot rhe colony (Fig.
IC). Each autozooid with a series of simple
septulae in the lateral, distal and proximal walls
(Fig. 4B). all septulae becoming included in a
senes of chambered pores which develop fron-
tally inlo new uutozooids or aviculana (Fig. 4B).

The budding of autozooids is regular and radial,
each astogenetic generation of autozooids alter-
nating with one of kenozooids (Fig. 2C). Au-
tozooid frontal shield with about 25 closely
spaced costae, the distal costae thickened as an
apertural bar which may be weakly bifid or thick-
ened to form a median ridge (Figs 2C,D). Costae
alternate with regular rows of 7-8 fine lacunae on
each side of the mid-line. Secondary calcified
orifice rounded, with a pair of flattened, curved
bifid to trifid oral spines (Fig. 2D), a raised.
fimbriated distal plate with 3 denticles (Fig. 4A)
and distal pit (Fig. 4A), and a pair of condyles.
Avicularia usually paired, oral (Fig. 2C), or sin-
gle, proximal (Figs 1D, 4B); rostra raised, with
paired condyles (Fig. 2C). Basal surface of cone
hollow, with laree paired foramina surrounded by
kenozooids and avicularia budded round the
proximal ends of the successive generations of
erect autozooids (Fig. 1 D).

Colony and zooid dimensions are given in
Table 1.

REMARKS. Maplestone (1909) described the
colony as encrusting, but there was no sign of any
substratum in specimens we examined. Of the 39
species listed by Maplestone (1909) associated
with A, doliaris from New South Wales, 1] were
new Io science and 15 had a sand-fauna (see
Cook, 1981) adapted colony form. Of these, 10
had rhizoids (7 species of Conescharellina, | of
Zeuglopora, 1 of Sphaerapora, and | of An-
askopora), and 5 were free-living, lunulitiforra
species (1 of Lunularia, 3 of Selenaria and 1 of
Orionella spp.).

Hastings (1964) gave a very full description
and interpretation of colony growth and relation
ships in C. daliaris. She noted the development
of kenozooids from septulae, and figured and
described the distal fimbriated plate. However,
her account was written before the explanation of
frontal budding by Banta (1972) and of reversed
frontal budding by Cook & Lagaaij (1976). Has-
tings used the structure of the ascophoran genus
Conescharellina in her comparisons. The bud-
ding pattern is analogous in the two exainples,
although the type of frontal bud produced is quite
different in Conescharellina, Hastings (1964:
258; 1966: 68) noted the similarities in develop-
ment of intercalated avicularia and kenozooids in
C. doliaris and Membraniporella agassizii Smitt
(1873: 11, Pl. 5, Figs 103-106), from Florida. M.
agassizii is ereet and quadriserial; the autozooids
have costae without either pelmatidia or lateral
costal fusions (see also Ristedt, 1979, Pl, 2, Figs

h MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Anaskopora doliaris (Maplestone). A,
QMG21286, details of orifices, x 90; B, QMG21287,
details of kenozooids, x 55.

1,2). The gymnocystis distinct earl y in ontogeny,
but later changes produce avicularia and
kenozooids surrounding the autozooids which
eventually obscure all but the orifices on the
frontal surface of branches. The origin of the
kenozooids and avicularia has not been de-
scribed, but may be inferred ta be from cham-
bered pores similar to those in Anaskopora. They
appear to strengthen the delicate, erect branches.
Hastings (1964) also emphasized the similarities
in interzooidal budding from the pore-chambers
of Hipporhoa with the budding of kenozooids in
C. doliaris, The similarities in pore-chamber and
chambered-pore structure are discussed by Bock
and Cook (in press).

Anaskopora doliaris is known from only 4 col-
onies. The lectotype and paralectotype were both
from the *Miner' collections, one specimen hav-
ing been sent by Maplestone to the then British
Museum (Natural History) as a ‘cotype’. We infer
that the two colonies of Anaskopora doliaris from
the ‘Cidaris 1' collection were anchored hy rhi-
zoids like those of A. parkeri, which probably
emanate from the large, paired foramina on the

basal surface (Fig. 2C). It differs from A. parkeri
and the other globular, fossil species, A. elevara,
A. cornuta and A. rotundata (see Bock & Cook,
in press), in the regularity of its reversed frontal
budding and hollow basal surface, It resembles
other species of Anaskopora in the mode of de-
velopment and distribution of interzooidal
kenozooids and avicularia, and the presence of
distal fimbriated plate in the autozooids. The
reasons for its removal from the genus Reginella
are related to the history of the concepts and the
characters of that genus which are discussed
below.

Reginella Jullien, 1886

The attribution of Cellepora doliaris
Maplestone, 1909 to the genus Reginella by Has-
tings (1964) seems to have been the result of a
series of misconceptions which have had several
repercussions in the later interpretation of Aus-
iralasian and other cribriomorph species.

Reginella was introduced by Jullien (1886:
605) ås a genus of Costulidae (=Cribrilinidae), for
C. furcata Hincks (1882: 470, PI. 20, Fig. 5), a
Recent species from the coast of British Colum-
bia. No other species was included. Jullien's def-
inition was not based upon the examination of
specimens, but was an interpretation of Hincks's
description and figure. Canu & Bassler (1929:
243) provided a literal English translation of
Jullien's generic description, which mentioned
that the frontal shield was composed of ‘velumi-
nous ribs much in relief, and that the ‘intercostal
furrows traverse entirely the zooecium and sepa-
rate completely each pair of transverse ribs'.
Pelmatidia were described as decreasing in size
from the margin along each costa, and the lacunae
were noted as each occupying ‘the middle of a
calcareous polygonal cell. Osburn (1950: 179,
Pl. 28, Fig. 3) redescribed R. furcata from Cali-
fornian specimens, and noted that septulae were
present, that the brooding zooid orifices were
dimorphic, that the large ovicells were punctate
and had a median keel, and that the oral spines
were often absent. Hastings (1964: 253) also re-
described K. furcata from Hincks's British Co-
lumbian material and noted that Jullien’s
‘polygonal cells’ were *no more than an effect of
light and shade’, There are, howeyer, other dis-
crepancies between Hincks’s (1882) description
and Jullien's (1886) definition. Hincks empha-
sized that the costae were only ‘slightly raised",
and that the intercostal *erooves! were ‘shallow’
and were arranged ‘radiating ta the median line’.

RECENT SPECIES OF ANASKOPORA

TABLE 1, Comparative measurements of colony, autozooid and avicularium in Anaskopora parkeri and A.
doliaris, Original measurements calculated from scanning electron micrographs. Width of lacuna in microns;
all other measurements in millimetres. The frontal shield length marked by an asterisk is a probable misprint in

Hastings (1964). Autozooid length is considered the same as frontal shield width.

Colony dimensions

Auiozooid length

1.06, 1.06, 1.08 0.69, 0.72, 0:76, 0.68

Fromal shield length, width 0.1* 4 0.5

Aperture antozooid

0.95. 1.01 L, 0.66, 0.70 W
0.21, 0.22, 0.22 L, 0.22, 0.24,
0.24 W

0.49, 0.60 L. 0.41 W
0.14, 0.15 L, 0.17, 0.18,0.21 W

Apertural bar width

Number ordenticleson | of denticles on
fi (Number ordenticleson | late

0.15
03,04.04,04L.02,02 W

0.05, 0.07
0.1, 0.1 L, 0.06, 0.06 W

0.07, 0.07, 0,10

Width of costae

ih of pit at base of fimbriated
Number of costae 9-13 (one side only)

Number ot lacunae per side
Width of lacunae

Height of oral àvicularium

Width of basal kenozooidal

chambers

10-25

0,03. 0.03, 0.03, 0.04, 0.06, 0,06,
0,06,0.09

CO — |

0.07. 0.07, 0.09,0 09

5-8

ET |= 0.18,0.19 =E aiw

Aperture ot oral avicularium 0.16, 0.15 L, 0.12. 0.10 W

0.17, 0.17, 0.17 L, 0.13, 0.13,
0.13 W

D, 16, 0.19, 0,29

nor applicyble

This is in contrast to Jullien's prominent costae,
and furrows which traversed the zooid horizon-
tally. Cook (1985: 123) also examined Hincks's
specimens of R. furcata and noted that the costae
were broad and flattened, with pelmatidia and
small lacunae, and remarked ‘the relationship of
R. furcata with the various species subsequently
assigned to Reginella requires investigation’.
The type and other specimens of R. furcata have
been re-examined, and a short description is
given below,

Reginella furcata (Hincks)

Cribrilina furcata Hincks, 1882: 470, PI. 20, Fig. 5.

Reginella furcara O'Donoghue & O'Donoghue 1926:
52; Osburn 1950: 179, PI. 28, Fig. 3: Hastings 1964:
253; Soule et al, 1995: 123, Fig. 42 A-C,

MATERIAL EXAMINED. BMNH1886.3.16,18,
Queen Charlotte Islands. Hincks Coll. on 4 shell frag-
ments; 1921. [1.17.12 and 1968,1.18,100, Departure
Bay, Vancouver Island, O'Donoghue Coll;
1986.9.10.3 Cannon Beach. Western Washington, L:
Pitt Coll.

RETRO. Autozooids encrusting, colony

riginating from a membraniporiform ancestrula
with long marginal spines. Zooidal gymnocyst
very narrow, vertical walls shallow. Frontal

shield of 11-17 flattened costae, each with 3-4
pelmatidia, Suboral bar variously thickened,
sometimes with å minute central mucro. Costae
irregularly fused with a shallow central suture;
the distal pairs of costae extending horizontally,
the proximal 5-7 pairs converging distally and
medially, Intercostal furrows narrow, with 4-6
rounded lacunae. Six evanescent oral spines, one
lateral pair often remaining, and becoming thick-
ened, or extended frontally and bifurcated termi-
nally, Zooids communicating through 2 distal and
3 jateral groups of septulae, each enclosed in a
very shallow, slit-like chamber at the base of the
gymnocyst. Ovicells large, hyperstomial, with a
median suture line and 16-20 small, rounded or
irregular pores scattered over the surface; fertile
orifices very slightly wider than autozooid ori-
fices. Avicularia absent.

REMARKS. Osburn (1950) illustrated a single
zooid of his Californian material of R. furcata.
The drawing is misleading, as it resembles
Jullien's concept of Reginella rather more than
Hincks's description of R. fureata. The zooid
appears to be at a late ontogenetic stage, with
thickened, but not prominent, costae and the lines
of lacunae traverse the frontal shield horizontally,
with no obvious median suture line. Pelmatidia
were reported to be present but were not

8 MEMOIRS OF THE QUEENSLAND MUSEUM

illustrated. Osburn (1950: 180, Pl. 28, Fig. 4 and
PI. 29, Fig. 3) also illustrated another Californian
species as Reginella mucronata, which had orig-
inally been introduced for Pleistocene material
from the same region by Canu & Bassler (1923:
92, PI. 35, Fig. 4). Canu & Bassler (1923) referred
it to the genus Metracolposa Canu & Bassler
(1917). Canu & Bassler's (1923) retouched pho-
tographs show zooids very similar to that
illustrated by Osburn for R. furcata, rather than
the rounded zooids with large lacunae he
illustrated for R. mucronata. Hastings (1964:
253) considered that these two species were iden-
tical, although Soule (1959: 46), who had exam-
ined both fossil and Recent material, had
regarded R. mucronata as distinct. The genus
Metracolposa was originally introduced for the
Eocene species M. robusta Canu & Bassler
(1917: 35, Pl. 3, Fig. 6; and 1920: 308, Pl. 43, Figs
1-7), from North Carolina. M. robusta and the
other Eocene species described by Canu &
Bassler were all illustrated by heavily retouched
photographs, many of elongated zooids with un-
interrupted lines of lacunae extending across the
frontal shield, with little or no indication of a
median suture line. M. robusta has large, erect,
bilaminar colonies and zooids with paired oral
avicularia. The ovicells are very large and ridged
centrally, but not punctate like those of R. furcata.
Generally, M. robusta, the type species, and the
other Eocene forms assigned to Metracolposa,
have little in common with, and are distinct from,
Reginella. We accept that the Pleistocene-to-Re-
cent M. mucronata is congeneric with R. furcata,
but this provides no justification for regarding the
genus Metracolposa as a synonym of Reginella.
That idea had been very tentatively suggested by
Osburn (1950) but was later greatly extended by
Hastings (1964) in order to include other species
with avicularia within the generic description
(see below).

Jullien's (1886) definition and the studies just
cited so modified the generic concept of
Reginella that it now included characters quite
unlike those of its type species. This would ex-
plain the implicit referral of Cribrilina vas
Brown, 1954, a Pliocene species from New Zea-
land, to Reginella by Brown (1958), when dis-
cussing a somewhat similar species from the
Tertiary of Victoria, Reginella maplestonei
Brown, 1958. Neither species has avicularia, but
small, peristomial ovicells were illustrated in R.
maplestonei. Powell (1967: 221, Pl. 2, Fig. 6, Fig.
4) redescribed R. vas from Recent New Zealand
specimens, noting its elongated zooids, absence

of pelmatidia, and small, imperforate, peristomial
ovicells; all characters completely unlike those of
R. furcata. Brown (1958) also included the Re-
cent, Antarctic Cribrilina projecta Waters, 1904,
which has small oral avicularia and distinct
pelmatidia, in his concept of Reginella. Hastings
(1964) considered that C. projecta was not con-
generic with R. furcata, but gave no evidence for
her conclusion. Moyano (1985) introduced the
genus Dendroperistoma for C. projecta.

The characters of Cellepora doliaris
Maplestone, 1909 also became involved in the
Reginella problem, when Brown (1958:53) men-
tioned that Hastings (presumably in litt.), "has
also pointed out a very close resemblance in the
nature of the frontal shield between C. vas and
Cellepora doliaris Maplestone'. Hastings, how-
ever, did not redescribe C. doliaris for a further 6
years. When she did, she remarked (1964: 258,
footnote 1), with a reference to Brown (1958),
that Brown had told her in 1955 that he had
‘referred Cribrilina alcicornis and Cellepora
doliaris to Reginella’ when, in fact, Brown (1958)
had never mentioned C. alcicornis at all (see
below).

C. alcicornis Jullien (1883: 508, Pl. 14, Figs
23-25) is a deep-water, encrusting species from
the Northeastern Atlantic. Autozooids have cos-
tae with no pelmatidia, and four, large, branched
oral spines. Interzooidal avicularia and large
interzooidal kenozooids are budded among the
autozooids, and the elongated ovicells are not
punctate. C. alcicornis is therefore unlike R.
furcata, and it is significant that Jullien (1886) did
not include C. alcicornis when introducing
Reginella, and that later, Calvet (1907:399) and
Prenant & Bobin (1966: 578, Fig. 200), continued
to refer the species to Cribrilina. More recently,
both d'Hondt (1974: 47, Fig. 7) and Harmelin
(1978: 178, Pl. 1, Fig. 3, Figs 3-4), who examined
additional Atlantic material, also maintained C.
alcicornis in Cribrilina. Harmelin noted that one
of Hastings's ‘points of resemblance’ between C.
doliaris and C. alcicornis was based on the mis-
taken assumption that Jullien's (1886) phrase
‘grande ponctuations referred to the interzooidal
kenozooids, whereas in fact, it referred to the
intercostal lacunae. He remarked "Le changement
de genera ne semble pas justifié pour cette
espèce’.

Quite apart from this confusion, it is unfortun-
ate that Hastings (1964: 252) extended Osburn’s
(1950) tentative synonymy of Metracolposa with
Reginella, including some of the characters of the
Eocene M. robusta, such as avicularia, in her

RECENT SPECIES OF ANASKOPORA 9

concept of. Reginella. Thus the presence of nu-
merous avicularia in C. doliaris, which are totally
absent in Å. furcara, did not preclude its referral
to Reginella, because avicularia had become å
"generic character". In the same way, the presence
of kenozooids with the avicularia in C. doliaris,
was mitigated by the tacit assignment of
Cribrilina alcicornis, which has both, ta
Reginella, although neither were present in R.
furcata. The presence of avicularia and
kenozooids in both C. alcicornis and C. doliarts
is interesting. but the structures differ completely
in their origins and general morphology between
the 1Ww0 species. In any case, neither has anything
10 do with the characters of the genus Reginella,
Two North American species which have been
referred tu Keginella also require further investi-
gaton. Reginella floridana (Srnitl, 1873) has
been described by Winston (1982) from Florida,
and by Cook (1985) from West Africa. No
ovicells have ever been found in any specimen of
(bis species. R. repangulata Winston &
Håkansson, 1986, from an interstitial Floridan
fauna, has ovicells somewhat similar to those of
R. furcara.

It is obvious that by this time the original char-
acteristics of the genus have become completely
submerged by the accumulation of additional or
alternative features derived from the diversity of
species included. This is illustrated by Gordon
(1984: 63, Pl. 20, Figs D-E), who redescribed
Reginella vas, and introduced a new, unisenal
encrusting species from New Zealand, R.
stolonifera Gordon (1984: 63, Pl. 20, Figs A-C),
which had large, branched oral spines like those
of C. alcicornis. The concept of the genus was
now extended to include all the disparate forms
mentioned above, from the Arctic, Antarctic, At-
lantic and Pacific Oceans, from deep and shallow,
even interstitial waters, and from the Tertiaries of
North America, Australia and New Zealand. The
generic characters now encompassed: colonies
encrusting, erect and bilaminar, and conical; zo-
vids uniserial to contiguous and frontally budded,
avicularia present or absent; ovicells hyperston-
jal and peristomial with or without pores or fenes-
trae; pore chambers present or absent, To these
may be added, pelmatidia and kenozooids present
or absent. The only consistent characters are 'slit-
like intercostal lacunae’, and “costae arranged in
straight rows across the zooid, often traversing
the mid-line without interruption’. Although
these features occur late in ontogeny in Å, vas and
R. stolonifera, neither occurs in R. furcata.

In conclusion, it is obvious that Reginella
should be restricted to Recent and Pleistocene
species from the North American ànd Japanese
Pacific coasts, and that Merracolposa should be
retained only for the American Eocene species.
Soule, Soule & Chaney (1995) give. notes on
Reginella furcata and introduce å new genus.
Reginelloides, for R. stolonifera Gordon, 1984.
Cribrilina alcicornis, C. vas, Reginella
maplestonei, R. floridana and R. repangulam
should also he re-investigated, and new generic
groups should be introduced for their reception
where necessary.

The colonies of R. furcata from Vancouver
Island examined by one of us (PLC) included
about 20 with ancestrulae present. The an-
cestrulac had small pore-chambers distally and
laterally, and 12 lung spines surrounded the ope-
sia. Mawatan (1988: 149, Figs 9-14) hus de-
scribed specimens from Hokkaido, Japan, as
Reginella furcata, and discussed previous Japan-
ese records in detail. These include the species
described as Lyrula multipora by Sakakura
(1935: 109, PI, 8, Fig. 7), and as Figularia multi-
parabySilén (1941: 117, Figs 178-180). All these
Japanese records differ trom AA, furcata very
slightly, in the complete lack of oral spines, even
in early ontogeny, the raised mucros beside the
orifice which give it å subtriangular appearance,
and the lack of any dimorphism in the fertile
orifices. Osburn (1950) described several nomi-
nal taxa from the Arctic to the Mexican coasts of
North America and referred them to Reginella.
They all resemble R. furcata in possessing
straight, converging rows of lacunae, paired oral
spines and large ovicells with a central suture and
punctate surface. Until the actual limits of varia-
non of these species, especially those from Cali-
fornia, like R. mucronata, have been analysed, it
is probably best to maintain them, together with
R. multipora, as distinct species of Regineffa.

ACKNOWLEDGEMENTS

We thank Dr PE. Bock (Royal Melbourne
Institute of Technology) for his help and interest.
Ms M. Spencer Jones (Natural History Museum,
London), for the loan of specimens of Reginella

furcata and Dr P. Berents (Australian Museum)

for confirming details of Maplestone's type of C.
doliaris. The 'Cidaris' expeditions on RN.
‘Franklin’ were partially funded by an Australian
Manne Science and Technology Grant to Prof. M.
Pichon,

10 MEMOIRS OF THE QUEENSLAND MUSEUM

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CALVET, L. 1907. Bryozoaires. Expeditions
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COOK. P.L. & LAGAAN, R. 1976, Some Tertiary and
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12 MEMOIRS OF THE QUEENSLAND MUSEUM

SYNONYMY OF CTENOTUS MONTICOLA STORR,
1981 AND CTENOTUS HYPATIA INGRAM AND
CZECHURA, 1990, Memoirs of the Queensland Museum
42(1): 12. 1997:- Cienotus monticola and C. hypatia are
mediurn-sized. Both are classed "poorly known" (McDonald
er al, 1991), The former has å ‘a very restricted distribution

.. with a maximum geographic distribution of less than

100kn". The latter is"... known only from the type collection",
C. monticala was described from à series of specimens col-
lected *77km W af Mareeba, Queensland (17°02'S,
145920 EI The type locality of C. hypatia is ' grantte gorge,
15km W of Mareeba, (17°00'S, 145" 1 7' EJ , These localities
are both on Granite Ck, approximately 6km apart (J. Cov-
acevich pers. comm. ). That, and the fact that the type descrip-
tions of C. monticola and C. hypatia sre very similar.
prompted a re-examination of the type material of each to
determine whether or not they are distinct.

Re-examination of the (ype specimens and type descrip-
lions indicates that C, monticola and C. hypatia are conspe-
cific (Table 1). Almost without exception, the measurements
and scale characters of C. hypatia fall well within the range
nf values. provided by Storr (1981) for C. monticola. The
differences in elongation, indicated by the chin-vent and

aravertebral counts are inconsequential. Greater variation is
requently recorded in a single species (e.g, C. astiemis Horner,
1995 & C. stuartt Horner 1995). Additionally, both type
specimens share the following characters: nasals separated;
nasal groove absent, rostral and fronronasal in narrow to
moderate contact; second loreal 1.2 times as wide as high;
prefrontals large and separated, frontal long and narrow con-
tacting the prefrontals, the frontonasul, the first 3 supra-
oculars, and the frontopanetals, 4 supraoculars, 2nd the
largest; Ist supraciliary the largest, 4th to penultimate consid-
erably smaller than others; Sth supralabial subacular, ear
aperture large; toes compressed,

Colour and pattern are known to be useful in distinguishing
some species of Crenotus (e.g, C. areanus Czechura. E
Wombey, 1982 and C. robustus Storr, 1970). However, this
is not the case with C, monticola and C. hypatta, Colour and
pattern. of the type specimens of C. monticola (5) and C.
kypatia (1) have been detailed by Storr (1981) und Ingram &
Czechura (1990). Variution in the former fully covers the
fatter.

Thus, in every respect (meristics, sealation, colour/pattern)
there are no significant differences between C, monricola and
C. hyparia. Ctenotus mouticola Storr, 1981 is, therefore, a
senior synonym of C. hyparia Ingram & Czechura, 1990,

Acknowledgements
We thank Jeanette Covacevich for reading and suggesting
Improvements to this nole,

Literature Cited

CZECHURA, ON E WOMBEY, J. 1982, Three new striped
skinks, (Crenoms, Lacenilia, Scincidae) from Queens-
land. Memoirs of the Queensland Museum 20(3): 639-43,

HORNER, P. 1995. Two new species of Crenotus (Reptilia.
Scincidae) from the Northern Territory. The Beagle,
Records of the Museums and Art Galleries of the North-
em Territory 12:75-88.

TABLE |, Comparison of SE of C. monticola
(QMJ39468) and C, hypatia (QM 42092). Values in paren
theses are from the type descriptions.

Species

Character C. monticola

(QMJ39468) (QM142092)
Snout-vent (mm) 53 (51-61)
i (SNL) 44 (44) |
215 (215)

13 (NA) 12012)

20 (NA) 19 (NA)

KU 0r 8) 8 (9)

Upper ciliaries

[Presuboculars — | 2(2.rarely 3) |

ae canc ^

| Upper labials ZO)
| Ear lobules 2 or 3 (3-5) 3 or 4 (3-4)
E arg I or 2 (NA)

Largest ear lobule JL usually langest)|
nuchal scales
26 (24-28) 28 (28)

Midbody scale rows
76 (NA) N2(753

No. of scales from
chin-vent

Paravertebral scale
rows

Subdigital lamellae 18/19 (17-19) 18/19 (20)

Structure of moderately wide | narrow callus
subdigital lamellae | callus (narrow to (keeled)
moderately wide
callus)

&7(NA) T2(NA)

INGRAM, G. J. & CZECHURA, G.V. 1990. Four new spe-
cies of striped skinks from Queensland. Memoirs of the
Queensland Museum 29(2): 407-410,

McDONALD. K.R., COVACEVICH, J.A., INGRAM, G.J.
& COUPER, P.J. 1991. The status of frogs and reptiles.
Pp 338-345. In Ingram, GJ, & Raven, R.J. (eds), “An
Atlas of Queensland's Frogs, Reptiles, Birds and
Mammals", (Queensland Museum: Brisbane). 391 pp.

STORR, G.M. 1970. The genus Crenoius (Lacertilia :
Scincidae) in the Northem Territory. Journal of the
Royal Society of Western Australia 32: 97-108.

19831. Ten new Crenotus (Lacertilia: Scincidae) from Aus-
tralia. Records of the Western Australian Museum 2):
125-146,

P.J, Couper, Queensland Museum, PO Box 3300, South Bris-

bane, Queensland 4107, Australla. Pianka, E.R., Department

af Zoology, The University of Texas at Austin, Austin, Texas,

78712-1064, USA: 23 May 1997

A NEW GENUS OF HIPPOLYTID SHRIMP (CRUSTACEA: DECAPODA;
HIPPOLYTIDAE) FOR THOR MALDIVENSIS BORRADAILE

AJ. BRUCE

Bruce, A.J. 1997 06 30: A new genus of hippolytid shrimp for Thor maldivensis Borradaile.
Memoirs of the Queensland Museum, 42(1) 13-23. Brisbane, ISSN 0079-8835,

Å new genus Thorina, is designated to accommodate the hippol ytid shrimp Thor maldivensix
Borvadaile 1915, The new genus is distinguished from Thor particularly by the short,
unidentate rostrum and the absence of an appendix masculina from the male second
hate with marked sexual dimorphism of the first pereiopods. First recorded from thé

aldive Islands, the species is sparsely recorded throughout the Indo-West Pacific region
and newly recorded from Tanganyika, Mauritius, Seychelle Islands, Western Australia, the
Great Barrier Reef, Papua New Guinea, Tonga, Cook and Society Islands. [ ] Natantia,
Hippolytidae, Thorina , Indian Ocean, Pacific Ocean.

AL Bruce, Crustacea Section, Queensland Museum; P.O. Box 3300, Sauth Brisbane,

Queensland, 4101, Australia: 17 May [996.

In his study of the hippolytid shrimps of the
Albaiross Expedition, Chace (in press) indicated
ihe anomalous systematic position of Thor
maldivensis Borradaile 1915. He clearly defined
the characters of the genus Thor Kingsley s. sit.
and lists five major characters that distinguish T.
maldivensis trom all other species of the genus
Thor, omitting it from his key to the species of
that genus. A new monospecific genus for the
reception of this species is here formally estab-
lished. The original description of Thor
maldivensis provided by Borradaile (1915) con-
sisted of a brief diagnosis only but a short more
detailed account with an illustration of å d exam-
ple was published in Borradaile (1917). This ac-
count remains the most detailed available of this
species, which has been only infrequently re-
corded in the zoological literature.
Abbreviations used: CL, postorbital carapace
length; MNHN, Museum national d'Histoire
Naturelle, Paris; NTM, Museums and Art Galler-
ies of the Northern Territory, Darwin; QM,
Queensland Museum, Brisbane.

SYSTEMATICS

Class CRUSTACEA
Order DECAPODA
Suborder NATANTIA
Infraorder CARIDEA
Family HIPPOLYTIDAE

Thorina gen. nov,

DIAGNOSIS. Small hippolytid shrimps of sub-
cylindrical body form. Rostrum very short, slen-
der. acute, not exceeding proximal segment of

antennular peduncle, with single small dorsal
tooth only. Carapace without carinae; supraor-
bital, non-articulate antennal spines present, he-
palic and pterygostomial spines absent.
Antennular peduncle with proximal segment with
moveable plate distally, upper flagellum brush-
like. Scaphocerite mid-laterally unarmed, Eyes
well-developed, cornea hemispherical, With
elongate acute median process anterior to lirst
thoracic sternite. Mandible without palp, with
incisor process, maxillipeds with epipods and
well developed flagella on exopods. Pereiopods
without arthrobranchs, epipods or exopods. First
pereiopod with fingers greater than 1/3 of palm
length: fingers without interlocking terminal
spines, merus with ventral teeth, chelae usually
greatly hypertrophied in some d d. Second pe-
relopod with carpus with 6 segments. Propods of
last 3 pereiopods not multiarticulate. Abdomen
non-cannate, without dorsal teeth, with anterior
3 pleura rounded, posterior 2 posteroventrally
acute, 6th abdominal segment without articulated
posteroventral plate. d second pleopod without
appendix masculina and appendix interna, Telson
with 3 pairs of dorsal spines, 3 pairs of posterior
spines.

TYPE SPECIES. Thor malilivensis Borradaile, 1915.

ETYMOLOGY. From Thor, a hippol ytid generic name
first used by Kingsley, 1878, and -ina (Latin), diminu-
tive. Gender, feminine.

SYSTEMATIC POSITION. Closely related 10
Thor Kingsley, with which most generic charac-
ters are shared and emphasised by the presence of
a triangular mobile plate on the distal segment of
the antennular peduncle, a feature otherwise

Li MEMOIRS OF THE QUEENSLAND MUSEUM

found To eceur only in Thor, Distinguished fram
Thor by the greatly reduced rostrum with only å
single dorsal tooth, marked sexual dimorphisn of
first pereiopods, reduced sexual dimorphism ot
Id pereiopods, presence of 3 pairs of posterior
telson spines instead of 1-2 pairs, and particularly,
the complete absence of an appendix masculina
and appendix interna on the à 2nd pleopod. The
strongly distolaterally spinose meri of the hyper-
trophied å Let pereiopods are also characteristic,
but it may be noted that a single small articulated
spine may be present in the West Atlantic species
Thor dobkini Chace (1972). In Thorina, the spines are
non-articulate denticular processes.

Thorina maldivensis comb, nov,
(Figs 1-6]

ror maldiversis. Borradaile, 1915: 208-209; 1917:
301-407, på. 56 fig, 6; Kemp, 1916: 391; Edmond-
sen, 1925: 6; 1946: 252, 253, fig. 153d; Holthuls,
1953: 53-54; Bruce, 1976: 51; Kumezaki, et al.,
1988: 8], col, pl.; Chace, in press.

MATERIAL EXAMINED. QMW 19914, 13. ] ovig.
9, stn DF,37, Heron Island, Capricorn Islands,
Queensland, 3.0m, 16 October 1976, coll, D.F. Fisk.
QMW21438, 13, 1 ovig. 9, stn AJB/162, Jadini,
Kenya, 22 December 1972. c. 0.4m, lagoon, in
Acropora, coll. A.J. Bruce. QMW21439, Id, Aldabra,
Seychelle Islands, 3 Navember 1964, from coral in
channel reef, intertidal, coll. AJ. Bruce. QM W21 440,
2 spms, macerated, Ras Iwaline, Mombasa, Kenya, 8
February 1972, lagoon, in Millepora, coll. AJ. Bruce.
QMW21441, 1, 1 ovig, 9, sin 140, Kirwetu, Kenya,
346,7 S 39?50.9'E, low water spring tide level, 6
November 1971, coll, A.J, Bruce, reef flat, under dead
coral, QMW21442, 4 ovig. ?, Astove Island,
Seychelle Islands, R.V, Manihine, Cr. 312, 20 August
1970, reef flat, under dead coral, coll. A.J. Bruce.
QOMW21443, 34, 30vig. 2 Farquhar Island, Seychelle
Islands, R.V. Manihine, Cr.336, sin AJB/60, 26 Febru-
ary 1972, coll. AJ, Bruce. QM W21444, 2 spms (1
ovig. 9), stå AJB/107, Ras Iwatine, Mombasa, Kenya,
4"01,3'5 39*44.0' E, 1m, 27 February 1971, lagoon, in
Srylophora, coll. A.J. Bruce. QM W21445, 1 spm (3 7),
sin AJB/99, Ras Twatine, Mombasa. Kenya, 4*04,0'5
19944, YE, 0,5m, | January 1971, lagoon, In
Pacillopara, coll. A.J. Bruce. QM W21446, I ovig. 9,
sin AJB/157, Jadini, Kenya, 4"19,0'5 39935,5'E,
2.001, 19 March 1972, in corals, coll, AJ. Bruce. OM
W21447, 1d, 1 ovig. F, stn AIB/I66, Bamburi,
Mombasa, Kenya, 4900,5'5 39995.0'E, outer reef
creat, 18 August 1973, coll. A.J, Bruce, QM W21448,
HV, 2 juvs, stn AJB/119, Ras Iwatine, Kenya, 47
D1.15'839*43.8' E, 2m, 27 July 1971, edge of deepreet
channel, in Pavona, coll, AJ, Bruce: QMW21449, | 9,
stn AJB/I38, Jadini. Kenya, 4°21.5'S 39*34,5 E, 0-
2m, nter lagoon, in coral. 3 November 1971, coll, al
Bruce. (author's collection), I spm, sm ATB/181a,

Tamannd, Mauritius, intertidal pools, 24 May 1974,
coll. AJ. Bruce, QMW21540, 1d, 3 op, ?, sth
AJB/140, Vipengo, Kenya, 6 November 197 1 reef Nat,
coll. AJ. Bruce. (author's collection), 2 spms, stn
AJB/209, South Patch, Motupore Island, Papua New
Guinea, 9°34.27'S 147*12,65' E, 10-20m, seaward reef
slope, scuba, 6 November 1980, coll. J.M. Lowry.
QMW?21451. (og 2, stn AJB/141, Tutia Reef,
Mafia Island, Tanganyika, 2m, 14 November 1971,
voll, AJ. Bruce, NTMCr 009156. 14, stn RH 92 12,
Cartier Reef, Western Australia, 127226'$
123?32,2"E, 10-15m, 9 May 1992, coll. J.R, Hanley,
BC. Russell, NTMCr004374, 13, 1 ovig, 2,2 juv.,
Masausi Bay, Tutuila, American Samoa, reef l.
N.4m. 22 July 1986, ‘Operation Raleigh", coll. M. Ricli-
mond. NTMCr.004375, 13, Omuta Landing, Mitiaro,
Cook Islands, reet lagoon, 2-3m, 11 July 1986, "Omer:
ation Raleigh’, coll. M. Richmond. NTMCr 004376, I
ovig. "d, Oholei, Tongatapu, Tonga, reef lagoon, 1m,
14 August 1986, ‘Operation Raleigh’, coll. M. Rich-
mond, MNHNNa-6627, 29 (1 ovig.), sin S-20, Tahiti,
Society Islands, coll. O, Odinetz, 1982.

DESCRIPTION. Small hippolytid shrimp ol
moderately slender, subeylindrical body form.
Carapace smooth, with short acute rostrum,
reaching to midlength of proximal segment of
antennular peduncle in dorsal view, without dis-
unci dorsal carina, with single small acute toath
at about level of tip of inferior orbital angle in
lateral view, ventral carina shallow, concave, lat-
eral carinae broadly divergent, continuous with
supraorbital margin, giving rostrum triangular
appearance in dorsal view, with small acute sti-
praorbital teeth, posterior 10 level of dorsal tooth;
inferior orbital angle slightly produced, rounded,
with small acute antennal spine ventrolaterally;
hepatic and pterygostomial spines absent,
branchiostegite with anterolateral angle broadly
rounded.

Abdomen smooth, with 3rd segment slightly
produced posterodorsally, non-carinale; 6th seg-
ment c. 1.33 times longer than maximal depth, c.
0.7 of carapace length in d, 0.55 in 2, with
posterolateral and posteroventral angles acute,
posteroventral angle non-articulate; first 3 pleura
broadly rounded, not markedly enlarged in
ovigerous ? 2, 4th and Sub posteriorly produced,
posteroventral angles broadly acute.

Telson c. 1.5 times length of 6th abdominal
segment in d, L3 times in 2, subequal (0 cara:
pace length in g, c. 0.73 in H. 2.6 times longer
(han anterior width, lateral margins sublineur,
posteriorly convergent, posterint margin angular
with acute median process, willy å pairs of small
dorsal spines at 0,4, 0.6 and 0.8 of telson length,
posterior margin. wilh 3 pirs of spines, lateral
spines small, subequal to dorsal spines, interme-

A NEW GENUS OF HIPPOLYTID SHRIMP 15

wr

JNQA
E

m

Si
Kaes

)

FIG. |. Thorina maldivensis (Borradaile). d. Aldabra Island (upper) and ovigerous ?, Heron Island (lower).

Scale in mm.

diate spines c. 3.0 times as long as lateral spines,
submedian spines slender, plumose, twice length
of lateral spines.

Antennule with proximal segment broad, about
as wide as long, with stout acute tooth at 0,6 of
ventromedial margin, stylocerite elongate, acute,
exceeding length of peduncle, with small an-
teroverted process proximodorsally; statocyst ob-
solete; intermediate segment short, broad, with
acute non-articulated lateral process; distal seg-

ment short, broad, with articulated triangular
plate dorsolaterally: upper flagellum with proxi-
mal 12-13 segments stout with dense tufts of
aesthetascs dorsolaterally, distal flagellum with c.
5slender segments; lower flagellum slender, with
c. 20 segments.

Antenna with basicerite stout, with broad acute,
ventrolateral tooth; carpocerite stout, sub-
cylindrical, extending to about end of antennular
peduncle, flagellum short, filiform c. 3.5 times

16 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Thorina maldivensis (Borradaile). Ovigerous 9, Heron Island. A, anterior carapace and appendages,
lateral; B, same, dorsal; C, rostrum and anterior carapace, lateral; D, antennule, dorsal; E, same, peduncle,
ventral; F, antenna; G, eye, dorsal; H, 2nd pereiopod; I, 3rd pereiopod; J, uropod; K, telson; L, same, posterior
spines, dorsal spine (inset).

A NEW GENUS OF HIPPOLYTID SHRIMP 17

M erok —
Ye

be f

C

In m

FIG. 3. Thorina maldivensis (Borradaile). Ovigerous Ẹ , Heron Island. A, mandible; B, same, molar process; C,
same, incisor process; D, maxillula; E, same, palp; F, maxilla; G, same, palp; H, Ist maxilliped, median process
of anterior sternite stippled; I, 2nd maxilliped; J, 3rd maxilliped; K, same, terminal spines of distal segment of

endopod.

18 MEMOIRS OF THE QUEENSLAND MUSEUM

carapace length, Seaphocerite exleniing well he-
vend stout part of upper antennular flagellum, c.
2,5 times longer than maximal width, situated at
c. 0,3 of length, tapering distally to rounded distal
lamella, distinctly exceeding tip of stout dis-
tolateral tooth, situated at c. 0.8 of straight lateral
margin length.

Eye well developed, with large well pigmented
globular cornea, diameter c. 0.33 of carapace
length in måle, 0.4 in 9, with conspicuous dorsal
ocellus; stalk stout, compressed.

Mouthparts generally similar to those of Thor
species. Large acute compressed transverse wian-
gular median plate, anterior to Ist thoracic sler-
mie, occluding space between coxal endites of
first maxilliped, Mandible without palp; molar
process stout, obliquely truncate distally, with
marginal setae and small denticles; incisor pro-
cess slender, distally oblique, with f small acute
teeth. Maxillula with short, feebly bilobed palp,
upper lube with slender simple seta, lower lobe
with stouter spiniform seta; upper lacinia larger,
oval, with numerous short spines distoventrally.
lower lacinia small, short, with several long
spines distally. Maxilla with slender, tapering
palp, with single spiniform terminal seta, short
preterminal dorsal seta and medial sela; distal
endie well developed, deeply bilohed, densely
sciase medially; proximal endite feebly devel-
oped, slightly bilobed, with few long setae,
Scaphognathite well developed. c. 3.0 times
longer Than central width. posterior lobe small,
anterior lobe large, with median margin slightly
voneave. First maxilliped with 2-segmented palp,
distal segment c, 2.5 times longer than wide,
sparsely setose medially, proximal segment
broader than distal, medial border convex,
sparsely selose; basal endite broad, angular, me-
thal margin with dense short setae; coxal endite
convex, medial Margin with sparse. coarse, long
plumose setae; exopod with well developed fa-
gellum. with numerous plumose setae distally,
caridean lobe small, with numerous plumose
marginal setae; epipod large, triangular. feebly
tnlobed. Second maxilliped with dactylar seg-
ment short, narrow, with numerous short stout
spines, propodal segment large, broad, distomed-
ial horder with numerous long spines; carus and
ischiomerus normal; basis with medial margin
excavate, dorsal and ventral medial borders with
numerous long slender setae, exopod normal,
coxa medially produced, laterally with small
elongate epipod, bearing small podobranch ante-
riorly. Third maxilliped robust, exceeding car-
pocerite by about terminal and half penultimate

segment in d and terminal segment only in 2. In
?, terminal segment c. 73 times longer than
central width, subcylindrical, slightly flattened
ventrally with numerous transverse rows of short
spines, distally obliquely truncate, with 6 stoul
spines, penultimale segment c, Då of Terminal
segment length, twice as long as wide, with
groups ol short spiniform setae medially, long
setae laterally; ischiomeral segment fused with
basis, combined segment subequal to terminal
segment length, moderately bowed, compressed
proximally, expanded distally, lateral margin
with c, 9 spiniform setae, distolateral angle with
single long Straight spine with short acute tooth
medially; hasis short. e. 0.2 of antepenultimate
segment length, medial margin slightly bilobed,
lateral border with small robust rounded epipod;
without arthrobranch. 3 third maxilliped similar
ta 9, exceeding carpocerite by penultimate and
terminal segments,

First perciopods showing marked sexual di-
morphism, small in 9 2, greatly hypertrophied in
some å 3, In 9 9, exceeding basicerite by about
carpus and chela; chela c. 0.5 of carapace length,
with palm subcylindrical, slightly swollen proxi-
mally, with few short pectinate cleaning setae
proxrmoventrally, feebly compressed distally, c.
2.4 times longer than proximal depth, smooth,
fingers about 1/2 palm length, stout, tapering
distally, with sharp medial culting edges, dactyl
with 4 stour non-cornified terminal leeth, fixed
finger with 3, central tooth enlarged with laminar
lateral expansion; carpus c. 0.4 of chela length,
2.5 limes longer than distal width, tapered proxi-
mally, smooth, unarmed, with several cleaning
setae distoventrally; merus c, 0,95 of chela
length, widest at midlength, 2.8 times longer than
central width, with acute ventrolateral tooth dis-
lally, proximal ventral margin with single small
spiniferous tubercle; ischium c. 0.55 of chela
length, compressed, 2.2 times longer than wide,
slightly broader distally than proximally, with
single small preterminal distoventral spinule;
busis and coxa without special features; without
exopod. Hypertrophic male chelae slightly un-
equal, similar, c. 0.25 of carapace length, with
palm subevlindrical, without cleaning setae pro-
ximoventrally, subuniform, €. 5.5 tines lunger
than central depth, ventromedially tuberculate,
fingers c. 0.3 of palm length, stout, deflexed, with
numerous long stilt setae, tapering distally, with
sharp strongly concave culling edges. with stout
hooked Lips crossing distally, dactylus with single
large acute tooth proximally, fixed finger with
smaller acute tooth more distally; carpus c. 0.5 of

A NEW GENUS OF HIPPOLYTID SHRIMP 19

X] N
i
` 4

LA mm

|

An

FIG. 4. Thorina maldivensis (Borradaile). First pereiopods. A, ovigerous 9, Heron Island, CL 2.2 mm; B, d,
Heron Island, CL 2.3mm; C, å, Aldabra Island, CL 2.2mm; D, d, Astove Island, CL 2.4mm.

chela length, 3.3 times longer than distal width,
tapered proximally, feebly tuberculate ventrome-
dially, unarmed, without cleaning setae dis-
loventrally; merus c. 0.9 of chela length,
subcylindrical, 6.0 times longer than central
width, ventrolateral surface tuberculate, with nu-
merous small acute distally directed tubercles,
with several (3-6) large stout acule preterminal
teeth distolaterally, distoventral angle rounded;
ischium obliquely articulated with merus, c. 0.27
of chela length, compressed, distally expanded,
2.0 times longer than greatest width, dorsally
carinate proximally with c. 8-9 denticles or tuber-
cles and long stiff erect spiniform setae; basis and
coxa robust, without special features; without
exopod. Intermediate males with similar but
smaller and less robust chelae, less spinose and
tuberculate, more closely resembling 9 chelae.
Second pereiopods slender, exceeding car-
pocerite by carpus and chela in 9, by chela and
3 distal segments of carpus in d. chela with palm

subcylindrical, c. 2.3 times longer that central
width, with few setal tufts distally, fingers c. 0.6
of palm length, slender tapering, 3.0 times longer
than proximal depth, cutting edges sharp, medial,
tips with 3 acute spines on dactyl, 2 spines and
short tooth on fixed finger; carpus 2.5 times chela
length, 12 times longer than wide, 6-segmented,
segments in ratio of 2: 1: 3,4: 1.9: 1.1: 1.6, first 2
segments poorly separated, distal segment with
transverse row of long serrulate setae distoventr-
ally. merus c. 1.75 times chela length, 6.3 times
longer than central width, 6.0 times longer than
wide, simple; ischium 0.9 of merus length, 5.3
times longer than wide, with 2 long simple spini-
form setae proximoventrally; basis and coxa
without special features.

Ambulatory pereiopods moderately robust,
third pereiopods with slight sexual dimorphism.
9 third pereiopod exceeding carpocerite by pro-
pod and dactyl; dactyl c. 0.27 of propod length,
2.6 times longer than proximal depth, stout, com-

20 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Thorina maldivensis (Borradaile). Ovigerous 9 , Heron Island. A, first pereiopod chela and distal carpus;
B, fingers of chela, lateral; C, same, medial; D, same, tip of fixed finger; E, same, medial ischiomeral spines;
F, 2nd periopod, chela; G, same, fingers; H, same, distal fingers, dactyl above; I, same, medial ischial spine; J,
2nd pereiopod, chela. K, third pereiopod, propod and dactyl. d. Heron Island. M, 3rd pereiopod, propod and
dactyl. N, major 2nd pereiopod, fingers. O, minor 2nd pereiopod, same.

pressed, distal end c. 0.6 of proximal depth, with ` propod c. 0.8 of carapace length, 7.5 times longer
smaller dorsal ungual spine and larger, stouter than wide, uniform, slightly bowed, ventral bor-
distoventral spine, each c. 0.3 of length of dorsal der with 2 larger distoventral spines, 5 ventral
border of corpus, ventral margin with two spines; spines, decreasing in size proximally, dorsal mar-

A NEW GENUS OF HIPPOLYTID SHRIMP 21

FIG. 6. Thorina maldivensis (Borradaile). Ovigerous 9 , Heron Island. A, 3rd pereiopod, dactyl. d. CL 2,3mm,
Heron Island. B, 3rd pereiopod, dactyl. C, Ist pleopod. D, same, endopod. E, 2nd pleopod. F, 3rd pleopod,
endopod. d, CL 2.4mm, Aldabra Island. G, 3rd pereiopod, dactyl. H, Ist pleopod. 1, 2nd pleopod.

gin sparsely setose; carpus c. 0.36 of propod
length, unarmed; merus robust, c. 0.72 of propod
length, 3.2 times longer than central width, with
articulated distolateral spine; ischium c. 0.68 of
meral length, 2.5 times longer than distal width,
narrower proximally, unarmed; basis and coxa
robust, without special features; without exopod.
4th and 5th pereiopods similar to 3rd. propods
slightly longer than 3rd pereiopod propod length,
meri shorter and more slender, 4th c. 0.95 and 5th

0.85 of 3rd merus length, Male 3rd pereiopod
exceeding carpocerite by propod and dactyl, with
slight sexual dimorphism; propod slightly ex-
panded distally in smaller specimen, with 3 pairs
of distoventral spines, dactylus as in 2, with 2
ventral spines; in larger d, propod scarcely ex-
panded distally, with 2 groups of 4 and 5 dis-
toventral spines, dactylus with 3 additional spines
on proximal ventral margin.

22 MEMOIRS OF THE QUEENSLAND MUSEUM

Ovigerous 9 pleopods without special fea-
tures, basipodite enlarged and expanded, rami
short, broadly expanded, 2nd to 5th endopods
with appendix interna. Male Ist pleopod with
basipodite robust, broad, ventromedially con-
cave, coxopodite with uncinate distoventral pro-
cess; endopod sublanceolate, c. 0.8 of exopod
length, 1.6 times longer than proximal width,
medial margin straight, setose, lateral margin
convex, with short plumose setae; exopod c. 3.3
times longer than wide, with plumose marginal
setae; 2nd pleopod larger, endopod c. 0.8 of exo-
pod length, with plumose marginal setae, lacking
appendix masculina and appendix interna, exo-
pod normal; third pleopod normal, with appendix
interna at c. 0.4 of medial margin length.

Uropod with protopodite with large acute lat-
eral tooth; exopod c. 2.5 times longer than wide,
lateral margin straight, sparsely setose, with small
acute distal tooth, with large mobile spine medi-
ally; endopod 0.95 of exopod length, 3.0 times
longer than wide.

TYPES. The type material is deposited in the
collection of the Zoology Museum, Cambridge,
United Kingdom. 3 lots of specimens are held
(pers. comm., R.C. Preece, 16 August 1995),
consisting of the following: (i) 1 specimen in
reasonable condition, from Minikoi, Laccadive
Islands; (ii) 1 specimen in reasonable condition,
from Salomon Island. (iii) 2 specimens, with
loose appendages, from Holulé Island, Malé,
Maldive Islands. 2 lots of specimens, (i) and (iii),
are noted as co-types. The specimens from
Minikoi and Holulé, are noted on their record
cards with "Gardiner Colln. Ann. Mag. Nat. Hist.
(8) XX, p. 208. Percy Sladen Trust. Exp. XXII pt
3. p 400'. The Salomon Island specimen is anno-
tated only “Gardiner Colln.* All have the acces-
sion number AR 3.1920.

MEASUREMENTS (mm). å, Astove Island:
postorbital carapace length, 2.3; carapace and
rostrum, 3.3; total body length, 12.5; major first
pereiopod chela, 5.5; minor first pereiopod chela,
4.8; 3rd pereiopod propod, 1.8; same, merus, 2.3.
Ovigerous 9 , Heron Island: postorbital carapace
length, 2.2; carapace and rostrum 3.1; total body
length, 11.8; Let pereiopod chela, 1.2, 3rd perei-
opod propod, 1.7; same, merus, 1.9.

COLOURATION. The Jadini, Kenya, specimen
was noted to be an opaque white, except for the
transparent posterior 4th, 5th and 6th abdominal
segments and caudal fan; pereiopod bases also
white, meri banded with white; antennule and

antenna white, filiform flagella transparent. The
Mitiaro specimens were noted by the collector as
"translucent, with pink and white mottled head
and legs, green tinge to lower abdomen and tail
and ‘translucent, with pink predominating, mot-
tled white head and legs', the Tutuila specimens
as 'transparent, with green tinge on lower abdo-
men, mottled green- pink head and legs', and the
Tongatapu specimens as 'transparent, with pink
tinge and pink mouthparts, red/white eggs, dark
body organs'. A good colour photograph is pro-
vided by Kamizake et al. (1988). This shows a
semi-translucent pale yellowish shrimp, heavily
mottled with fine red speckling and scattered
whitish patches, particularly at bases of pleopods
and base and tips of caudal fan.

REMARKS. The exact habitat of Thorina
maldivensis has not been established but most
personally collected specimens have been ob-
tained from shallow intertidal coral reef pools
with a wide variety of corals and other coelenter-
ates and algae. As many were collected by the use
of poison, the precise niches occupied were not
observed. Several specimens were collected from
coral heads but these associations may have been
accidental. Hayashi (1986) records this species
from under coral blocks. The species appears to
be generally uncommon, but Edmondson (1946)
reported that in Hawaiian waters the species ‘is
common among brown seaweeds near the shore'.
The colouration of the Jadini specimen does not
appear cryptic and suggests a commensal life-
style. Where several specimens were collected
together the association is more likely to be genu-
ine. The Papua New Guinea specimens from 10-
20m represent the maximum bathymetric range
for this species.

The d specimen from Cartier Reef had only
un-hypertrophied first pereiopods but the distal
propod and dactylus of the 3rd pereiopods were
distinctly sub-prehensile, the ventral dactylus
with accessory spines. The d from Tutuila pos-
sessed only part of a single first pereiopod, lack-
ing the chela. The merus was rather feebly
spinulate, with blunt spines, and the spiniform
setae along the dorsal margin of the carpus were
feebly developed, mostly short and slender, with
only a single distal long spiniform seta.

The transverse triangular median sternal pro-
cess that appears to lie anteriorly to the sternite of
the first maxilliped appears unusual but it is rather
difficult to discern its exact relationships. No
similar feature seems to have been reported in
other hippolytids but a similar, though much

A NEW GENUS OF HIPPOLYTID SHRIMP 23

smaller and narrower process, more antero-pos-
teriorly orientated, is also present in Thor
amboinensis (De Man).

DISTRIBUTION. Throughout most of the Indo-
West Pacific region. Type localities: Malé Atoll,
Maldive Islands; Minikoi, Laccadive Islands, and
Salomon Island. Also known from Kenya, Tan-
ganyika*, Mauritius*, Seychelle Islands*, Mald-
ive Islands, Andaman Islands, Cartier Reef*,
Great Barrier Reef*, Papua New Guinea*,
Ryukyu Islands, Marianas Islands, Marshall Is-
lands, Tonga*, Cook Islands*, Kiribati, Society
Islands*, and Hawaiian Islands. (* — new localities).
Tanganyika is used in its zoogeographical mean-
ing and not in a political sense.

LITERATURE CITED

BORRADAILE, L. A. 1915. Notes on Carides, Annals
and Magazine of Natural History, (8)15: 205-213.
1917, On Carides from the western Indian Ocean.
Transactions of the Linnean Society of London,
Zoology, (2)17: 397-412,

BRUCE, A.J. 1976. A Report on a small Collection of
Shrimps from the Kenya National Marine Parks at
Malindi, with Remarks on selected Species.
Zoologische Verhandlingen, Leiden, 154; 1-72.

CHACE, E.A. 1972. The Shrimps of the Smithsonian-
Bredin Caribbean Expeditions with a Summary of
ihe West Indian Shallow-water Species
(Crustacea: Decapoda: Natantia). Smithsonian
Contributions to Zoology, 98: i-x, 1-179.

In press. The Caridean Shrimps (Crustacea:
Decapoda) of the Albatross Philippine Expedi-
tion 1907-1910, Part 7: Atyidae, Eugonat-
onotidae, Rhynchocinetidae, Bathypalaemonidae,
Processidae and Hippolytidae. Smithsonian Con-
tributions to Zoology.

EDMONDSON, C.H. 1925. Crustacea of Central Trop-
ical Pacific. Marine Zoology of Tropical Central
Pacific. (Tanager Expedition Publication No. 1,
1915), Bulletin of the Bernice P. Bishop Museum,
Honolulu, 27: 3-62.

1946. Reef and Shore Fauna of Hawaii. Special
Publication, Bernice P. Bishop Museum, Hono-
lulu, 22: i-iii, 1-381.

HAYASHI, K. 1986. An Annotated List of Shrimps
(Alpheidae and Palaemonidae Excluded) Col-
lected from the Gilbert and Solomon Islands. Pro-
ceedings of the Japanese Society of Systematic
Zoology, 32: 17-29,

HOLTHUIS, LB 1953. Enumeration of the Decapod
and Stomatopod Crustacea from Pacific Coral
Islands. Atoll Research Bulletin, 24: 1-66.

KEMP, S. 1916. Notes on Crustacea Decapoda in the
Indian Museum, VII. Further Notes on Hippolytidae.
Records of the Indian Museum, 12: 385-405.

KINGSLEY, J.S. 1878. Notes on North American Car-
idea in the Museum of the Peabody Academy of
Sciences at Salem, Massachusetts. Proceedings of
the Academy of Sciences of Philadelphia, 1878:
89-98.

KAMEZAKI, N., NOMURA, K., HAMANO, T. &
MISAKI, H 1988, A Guide Book of Marine Ani-
mals and Plants of Okinawa, Crustacea (Macrura
and Anomura), 8: 1-232, col. pls. Shinsei Guide
Book Series. (Southern Press: Okinawa, Japan).

74 MEMOIRS OF THE QUEENSLAND MUSEUM

NEW DATA ON LERISTA INGRAMI, A RARE SKINK
FROM SOUTHERN CAPE YORK PENINSULA, AUS-
TRALIA. Memoirs of the Queensland Museum 42(1): 24
(997:- Lerista ingrami Storr, 1991 is poorly known. It &
treated as ‘rare’ (McDonald etal., 1991) arid was kriown, prior
to Sept., 1996, from 10 specimens (the holotype and nine

ralypes) from what amounts to a single locality — the
oredune of the beach hetween the fondi of the Mclvor R,
und Cape Flattery. on southern Cape York Peninsula. L
ingrami is one of å few species of Australian skinks (4/320
species) not photographed in life. The type locality of this
species has been visited recently (13 Sept, 1996), As s result,
we can clarify some minor discrepancies with collection daia
af the type specimens; present phorographs of a live specimen
of L ingrami and of iis type locality: describe the colour of L
ingrami m lifes and comment on prey of L. ingrii.

Published data on the type locality and the collection
tocality of the pararypes of L. ingrami is at Variance shghtly
with 1nformation recorded in the Queensland Museum regis-
ter. Storr (1991) records:

"Holotype 132396 in Queensland Museum collected on 27
July 1976 by G.J. Ingram near beach north of Melver River,
Queensland, in 15°07'S, 1457] 5E",

'Paratypes Queensland: 7km N of mouth of Melver River
(QM20644-5 1) and 5km N (QM20653)".

The QM register entries for these specimens are:
9QMI32396 (holotype) op mission road near beach, north of
Melvor R., 13 July, 1976, G.J. Ingram’, (Date of registration
and entry 27 July, 1976).
^QMJ20644-51 (paratypes) approx. din N of Melvar R. mouth
Cooktown 1:250000 328089, 20 Nov., 1970, JAC, T Pi Teb-
ble, C. Tanner; ‘buried in sand under log on first dune’.
QMJ20653 (paratype) approx. Am. N of Melvor R. mouth.
Cooktown 1:250000 318090, 20 Nov., 1970, JAC, T.P. Teb-
lle, C. Tanner: "buried in sand of first dune - underlogs'. (Date
of registration and entry 6 Jan., 1971).

For the last-mentioned, sometime in or about 1986, lati-
tude/longifude were calculated from an atlas and added to the
register in pencil, 15905'5 145?]4" E.

Following advice from Gå, Ingram (collector of the holo-
type of L. ingrami) and using å ‘Magellan Global Positioning
System (GPS), we have calculated the type locality for L.
imgrami( Fig. D) at L57070]"S 145*14'42"E, very close to the
locality calculated for/by Storr (but nor entered in the QM register)
at the Ome of his description of the species - 15°07 145915".

We collected topotypic specimens QMJ62430-1,
QMJ62443, One has heen photographed (Fig. 2). All were
found in the middle of the day, ‘at rest’, on slightly moist sand
under dead coconuts, Once disturbed, they invariably sought
escape into the sand, Both mid body (20 x3) and supraciliary

FIG. 1 The type locality of Lerissa grani Starr, 1991.
(Photograph, JAC)

=
NE d

FIG. 2 Leriste ingrami. (Photograph, Jell Wright, OM),

counts (3x2, 4x1) for these specimens fall within the range
given for the species by Starr ( 1991).

L. ingrami was described from old (15-21 years) spirit
Material, so its colour in life was not known. Storr's descrip-
non (1991) can be augmented: body upper surface a shiny pink
or grey-beige, profusely marked by tiny dark brown dors; head
scales dark-edged, Å sharp-edged dark brown lateral stripe
runs from the rostral to the base of the tail, tapering posteri-
orly. Upper surfaces of limbs the same colour as dorsum, also
with dark brown (minute) dots, Venter (SV) pale grey-white,
Tail pale to very bright orange ventrally and dorsally,

There is little information on the prey of Lerista spp.
Wilson & Knowles (1988) report `-. Small arthropods, their
eggs und larvae probably constitute the bulk of their diets ..."
L. bipes and L. muelleri are the only species from sandy
localities for which prey have been reported, and there 15 only
one other reference ta prey of Lerista (a Tasmanian record of
Hewer & Mollison, 1974). Smith (1976) records orthopterans
from L bipes, and proturans and collembolans from 4.
muelleri. Faecal pellets from the recently-collected L ingrami
have been examined by Dr G, Monteith, Senior Curator (Ento-
mology) atthe Queensland Museum. They contain remains of
two smäll scorpions (Arachnida: Scorpionida) and two small
seed bugs (Insecta! Lygaeidae). Both, he advises, are common
and active amongst leaf litter,

Tissue samples were taken from QMJ62430 und
QM16243 1 and sent to the tissue library of the South Austra-
lian Museum, Adelaide. Only minor external morphological
characters (e.g., mid-body scales, lamellae ot the third (oe,
supracilary scales and colour) separate L. ingrami from L.
orientalis (De Vis, 1889) and L. senulata Storr, 1991,

Literature Cited

Hewer, A. & Mollinson, B.C. 1974. Reptiles and amphibians
of Tasmania, Tasmanian Yearbook No.8 (1974): 50-60),

McDonald, K.R Covacevich, J,A., Ingram, G.J. & Couper,
PJ. 199]. The status of frogs and reptiles. Pp. 338-345,
In Ingram. G.J, & Raven, R.J. (eds) An atlas of
Queensland's frogs, reptiles, birds and mammals.
(Board of Trustees, Queensland Museum, Brisbane),

Smith, L.A, 1976, The reptiles of Barrow Island. Western
Australian Naturalist 13(6): 125-36,

Storr, G.M, 1991. Revision of Lerista orientalis (Lacertilia:
Srincidae )ofnorthem Australia Records of the Western
Australian Museum 15(2); 413-417

Wilson, S.K. & Knowles, D.G. 1988, Ausiralia's reptiles, A
photographic reference to rhe terrestrial repriles of Aus-
tralia. (William Collins Pty Lid: Sydney).

LA, Covacevielt, PJ Couper & L Roberts, Queensland Mu-

seum, PO Box 3300, South Brisbane, Queensland 4101, Aus-

traller 3 April 1997

TWO NEW SPECIES OF FRESHWATER ATYID SHRIMPS (CRUSTACEA: DECAPODA:
ATYIDAE) FROM NORTHERN QUEENSLAND AND THE DISTRIBUTIONAL ECOLOGY

OF THE CARIDINA TYPUS SPECIES-GROUP IN AUSTRALIA
SATISH CHOY AND JONATHAN MARSHALL

Choy, 8. & Marshall, J. 1997 06 30: Two new species of freshwater atyid shrimps (Crustacea:

Decapoda. Atyidae) from northern Queensland and the distributional ecology of the Caridina

topus [o ed in australia, Memolrs of rhe Queensland Muxeun AY I): 25-36. Brisbane. ISSN
79-8835.

Two new species of freshwater atyid shrimps are described froni tropical Queensland and
the four members of the Caridina typus-group now known from Australia are reviewed.
Caridina confusa sp, nov. is a slender animal with a relatively long, dorsoventrally flattened,
naked rostrum, found predominantly in open grassland streams of the Atherton Tableland.
Caridina spinula sp. nov., distinguished by its spiniform pterygostomian angle. is found in
small secondary rainforest streams on northeastern Cape York Peninsula. Although both
these new species look superficially like C. zebra Short. 1993, they can be distinguished by
à Combination of characteristics such as rostrum length, shape of the prerygostotniau angle,
Jength-depth ratio of the sixth abdominal segment and the shape of the protopod of the
uropod. C. zebra is found predominantly in primary rainforest streams of thé Atherton
Tableland and C. rypus is found in coastal tropical streams. All species are allopatric, except
for slight overlap in the distribution of C. zebra and C. confusa in some tnthroncipenically
disturbed streams of the upper Barron and the upper North Johnstone catchments. [|
Crustacea, Atyidae, Caridina, Queensland, distriburion,

S, Choy, Resource Sciences Centre, Department of Natural Resources, 1345 Ipswich Road,
Rocklea, Queensland 4106, Australia; J. Marshall, Faculty of Environmental Sciences,

Griffith University, Nathan, Queensland 4111, Australia: 11 December 1996.

Caridina zebra Short, 1993 is a tropical mon-
tane species belonging to the Caridina rypus spe-
cies-group (Short, 1993), This group is
characterised by a short, dorsally unarmed ros-
trum, the presence of epipods on the 1st four pairs
of pereiopods and the presence ol an appendix
interna on the endopod of the Ist pleopod of d d.
Short (1993) reported C. zebra from the monrane
streams of [he wet tropical rainforest areas of the
Tully, Herbert and Johnstone River catchments,
at altitudes of 400-900m. A smaller population of
C. zebra, is also known 1o occur in the lower
Koolmoon Creek (alt. 150m). à tributary of the
Tully River (Hughes et al, 1996). Caridina typus
Milne Edwards, 1837 has been reported from
coastal lowland streams at Cooktown and on
Dunk Island (Roux, 1926; Riek, 1953; Short,
1993),

Re-examination of Short’s material from the
Wet Tropics-Atherton Tableland area (Short,
1993: 62) indicated the presence of two distinct
morphological groups, one group possessing a
longer rostrum, from anthropogenically dis-
turbed grassland areas and the other possessing a
shorter rostrum, generally from relatively undis-
turbed rainforest areas, This was confirmed by
examination of recently collected material trom

the same area (17-I8^S, 145-146°E). Although
the distribution of these two groups tend to over-
lap slightly, they are morphologically and ecolog-
ically distinct. They each warrant species status.
Since the short rostrum form is the holotype of C.
zebra Short, 1993, its status is maintained. The
long rostrum form is described as a new species,
Caridina confusa.

A third morphological and geographically iso-
lated group was recently collected from the
streams in the Mcflwraith Range, Cape York Pen-
insula (13°35-44°S, 143?20' E). This is also de-
scribed as a new species, Caridina spinula,

MATERIAL AND METHODS

Specimens from the Queensland Museum are
provided with catalogue numbers with the prefix
OM. Unless otherwise stated all material was
collected by the first author and various col-
leagues from Griffith University and the Queens-
land Department of Natural Resources (formerly
part Department of Primary Industries). All ex-
amined specimens will be deposited in the
Queensland Museum. Collection was made using
a standard pond net and all samples immediately
preserved. The abbreviations used are: 68. sixth
abdominal segment length; A1P, antennular pe-

26 MEMOIRS OF THE QUEENSLAND MUSEUM

dunele length; AZP, antennal peduncle length;
CL, post-urbital carapace length; RL, rostrum
length; SC, scaphocerite length; SL, body length
from the post-orbital margin of the carapace to
the tip of the telson; ST, stylocente length; T,
telson length.

The format of the description and morphologi-
cal terms follow Choy (1991). Although prob-
lems in the terminology of cuticle spination and
setanon still exist, we have followed the termi-
nology of Felgenhauer (1992),

SYSTEMATICS

Caridina typus Milne Edwards, | 837

Cundinu typus Milne Edwards, 1837: p, 263, pl. 25615,
figs. 4, 5; Holthuis, 1965, p, 10, fig. 3.

Candina typus typica Bouvier, 1925, p.250, figs. 272-
295

Caridina typa Roux, 1926, p. 201; Rick, 1953, p. 117.

MATERIAL EXAMINED, Proserpine River below
Proserpine (20?24,2'S, 148 31.1 E), 19.10.94, LR.
Paterson, 15 d d 12.54. 111m CL), 7 9 9 (4.8-5-1mm
CL): QM W4795, Brandy Creek near Proserpine
(207208, 148"38 E), 214.75. R Monroe; P. File-
wood, ovigerous v (7.81mm CL), non-ovigerous 9
(5.82mm CL); QM W 14241, Lindeman 13., west side,
small creek near goll course (20278, 149?02' E),
27,3,83, 1, Short, 33 (4.03-4.09mm CL), 9 9 15.5-
7.2mm CL).

DIAGNOSIS. Rostrum short, laterally cam-
pressed, with 1-5 teeth of ventral margin; diaer-
esis with more than 20 hamate setae; intermediate
setae on posterior telsoni¢ margin longer than
lateral pair, plumose, with sclerotinous plug; egg
small (length «0.52mm).

REMARKS. Caridina typus has å wide ranging
distribution, extending from eastern and southern
Africa through the Indian Ocean islands, South-
cast Asia jo Japan, Australia and through to
French Polynesia, In Australia it has been col-

tected only from the northeastern coastal areas of

the mainland and from the nearby islands, be-
tween latitudes 15-2175. Although it seems to be
confined to the lower reaches of coastal and is-
land Australian streams, C. typus has been col-
fected trom altitudes of 300m elsewhere (Choy,
1991).

Caridina zebra Short. 1993
Curidina zebra Short, 1993. p. 62 (in part],

MATERIAL EXAMINED. All SNE listed hy
Short. 1995, p. 62. ADDITIONAL MATERIAL.

Upper Tully Catchment: QMW1I7118, Tully River
above Koombooloomba Dam (17°49"S, 145?35' E.
720m), 23.10,91, numerous specimens:
Koombooloomba Creek (17?51.5'S, 145°35.9'E,
790m), 30.10.93, 25 specimens; Echo Creek
(17°59,5'S, 145?38,3 E, 830m), 30.10.93, 50 speci-
mens; Carpentar Creek (17°53.3°S, 145?35 Y'E,
750m), 30.10.93, 23 specimens; Costigan Creek
(17°56°S, 145"37'E, 770m), 3,12.94, 11 specimens
(1.8-4,2mm CL); QMW17119, Koolmoon Creek
(179445, 145734" E), 25.7.90, 4 specimens; Koolmoon
Creek nt Walter's Waterhole (17°44,11°S, 145*34' E.
760m), 30,10.93, 7 specimens; Koolmoon Creek near
Tully River confluence (17^44.9'S, 145?37.1'E,
150m), 31.10.93, 6 specimens, Upper Herbert Catch-
ment: QMW17116, Blunder Creek (17736'S,
145?33'E), 28.11.90, 3 specimens; Blunder Creek
(17947.5'8, 145*32.2'E, 750m), 3.12.94, 1 4, 2
ovigerous NN (4,08-5.6mm CL); Rocky Creek
(1744,7'S, 145931,3'E, 760m), 30.10,93, 34 spéci-
mens; Cameroon Creek, 3.12.94, 14 specimens (1.4-
4.2mm CL);millstream River al Diversion Weir
(V7?40'S, 145*26' E, 720m), 12.10.94; 2 specimens;
Upper North Johnstone Catchment: QMW19261,
Malanda Falls, (17°20.2°S, 145?43,8'E, 750m),
211,93, 30 specimens (including ovigerous ? 2 1;
QMW 19285, Raspberry Creek, 3-4 km above Malanda
Falls (17°22.75'S, 145E33.6 E), 14.11.91, J. Short, P.
Davie, A. Humpherys, silt & para. grass (Urochloa
mutica) infested stream; Raspberry Creek (17?25,5' S,
145728, 8&' E), 31.10.93, 100 specimens; North
Johnstone River (17?40' S, 145739" E, 650m), 5.8.93, 2
9 9. I g; Whaca Creek (177248, 145°38'E, 650m),
7.11.94, 2 9 9; Ithaca Creek at Bauld Rd (17°23'S,
145*38'E, 685m), 13.10.94, 50 specimens; Thiaki
Creek at edge of rainforest (17725'S, 145?32' E, 795m),
5.12.94, 9 specimens (3.4-4, 8mm CL); Thiaki Creek in
rainforest, 13.10.94, 15 specimens; North Beatrice
River at Palmerston Highway (17°32'S, 145*36'F,
720m), 13,10.94, 25 specimens; Henrieta Creek at
Palmerston Highway (17?36'S, 145°45°E, 360m),
14.10.94, 18 specimens; QMW18722, Goolugan
Creek, 23,10,91, numerous specimens; Goolagan
Creek (17936.3'8, 145°45.5'E, 370m), 32 specimens;
Upper Barron Catchment: QMW3078, Atherton lap
water (179168, 145?29'Ej, 14.5.62, QDPI, 2 d d;
Barron River at The Crater (199175, 145?29 E),
7.12.94, 10d G, 8 ? 9 (Zovigerous) (2.0-5.0mm CL);
Barron River at Hemmings Road, 28.11.94, 3 juve-
niles; Wright's Creek (179048, 145?45'E), 4.12.94, 4
?9 LI ovigerous); Peterson Creek (17"17'S,
145"36 E), 1994-95, numerous specimens; Kauri
Creek (169545, 145738 B), 1994-95, numerous spec-
imens; Prior's Creek, I 9; Upper Gwynne Creek in
small pocket of rainforest (17^23.3' 8, 145%30.3'E),
5.12.94,2 F 9, 19 d 8 (3.6-4.0mm CL).

DIAGNOSIS. Body (Fig. 1b) rotund, may have
black and white transverse banding; rostrum (Fig.
2a-f} short, extending to base of 3rd antennular
peduncle (RL«0,5CL), dorsoventrally com-

NEW FRESHWATER SHRIMPS 27

FIG, |. a, Caridina confusa sp. nov,, 3; b, C. zebra
Short 1993, d c, C. spinula sp. nova d. Scale I mm.

pressed, may have an apical tooth (Fig. 2e);
pterygostomian angle acute but not spiniform;
dorsal telsonic spines (Fig. 3a) on posterior 0.66
of telson; posterior telsonic margin (Fig. 3f, g)
usually with median spine, sub-lateral pair of
setae sigmoid, longer than intermediate setae,
selalion numerically variable; protopod of uro-
pod (Fig. 3a, d) elongate, spinate; eggs large
(length «1.32mm); found mainly in primary
rainforest streams on the Atherton Tableland
along the Lamb-Francis-Card well Ranges.

REMARKS, Specimens of C. zebra from some
locations (streams in the upper Barron Catch-
ment) may seem to have a fairly long rostrum
(Fig. 2a, d, f), thus resembling C. confusa sp. nov.
However, the rostrum length relative to the cara-
pace length as well as other features (Table 2),
such as the relative lengths of the antennular
peduncle, scaphocerite, sixth abdominal segment
and the telson are characteristic of this species.
Caridina zebra is one of only two species of atyid
shrimps in the primary rainforest streams of the
upper Tully (the other being Australatya
striolata), where it is very abundant. In other
catchments (Barron, N Johnstone and Herbert),

it is found mainly in the rainforest reaches of
streams. However, it may be common in some
disturbed streams such as Raspberry, Ithaca und
Prior's Creeks. Unlike the upper Tully, which has
an extensive relatively undisturbed forested area,
the upper reaches of the Barron and N. Johnstone
Rivers have been generally cleared and converted
to pasture. Streams running through these open
prassland are inhabited predominantly by C. con-
fusa sp. nov. Only a small area of the upper
Herbert catchment is in the wet (simple notophyll
vine to complex mesophyll vine) forest; the rest
is in dry sclerophyll forest. The abundance ol C.
zebra in these dry-zone streams is low; these
streams are inhabited by another atyid, Paratya
australiensis. Ti is interesting Io note that only
specimens from some of the rainforest streams of
the upper Tully have the black and white trans-
verse banding on their bodies (see Short, 1993),
In all other areas these animals are translucent
brown, with scattered tiny reddish and bluish-
green chromatophores, similar to individuals of
the IWO new species.

Caridina confusa sp. nov.

Caridina zebra Short, 1993, p. $2 (in part):
QMW18841, QMW18720.

MATERIAL EXAMINED. HOLOTYPE. QMW-
21906 ovigerous H. 4.8mm CL, 2.56mm RL, 19.2mm
SL, Thiaki Creek al Seamark Road crossing (17°23.5'S,
145°32.5°E, 750m), stream flowing through open
grazing land, fringing para grass, water depth 0.3-
0.5m, velocity Q.3ms `, silty substrate, hand-netted
amongst edge para grass, 26.8.94, S. Choy, M. Hopper.
ALLOTYPE, QMW21907 adult c, 3.4mm CL.
2.0mm RL, 14.2mm SL, same locality data as holotype,
PARATYPES, Upper North Johnstone Catchment:
QMW21908 upper North Johnstone River near Brom-
field Swamp (17722.5'S, 145°31.3'B, 700m). open
grazing land, fringing para grass, water depth 0,3-0,5m,
hand-netled, 4.12.94, 21 3 d, 50 9 ' (13 ovigerous),
(2.6-5.0mm CL); QMWI18841], Thiaki Creek
(17°24.9'S, 145?35.3'E, 750m), water depth 0.2m,
electro-fished, 1992, Queensland DPI Fisheries
Johnstone Rivers Survey, I d (4.5mm CL);
QMW18720, small creek about 6 km SW of Malanda
(17722. S, 145°33.6°E, 750m), fringing para grass,
waler depth 0.2-1.5m, electro-fished, 1992, Queens-
land DPI Fisheries Johnstone Rivers Survey, å
(3.9mm CL), 2 9 9 (4.3, 4.5mm CL); QMW 18725.
Thiaki Creek, macrophyte area, sume data as
OMW 18841, [3 3 d. (3.2-4.8mm CL), 16 9 9 (3.8-
6.1mm CL); QMW21909 Thiaki Creek at Seamark
Road crossing (17°23.5'S, 145932.5'E, 750m), open
grazing land, [ringing para grass, water depth 0.3-(1. 5m,
hand-netted, 26.8.94, c, 100 specimens; QMW21910
Thiaki Creek at downstream edge of rainforest, Ining-

28 MEMOIRS OF THE QUEENSLAND MUSEUM

ing para grass, depth 0.3-0.5m,
hand-netted, 5.12.94, c. 50 spec.;
QMW21917 Ithaca Creek in
rainforest pocket (17?24.7'S,
145?30.3' E), leaf litter, water
depth 0.3-0.5m, hand-netted,
5.12.94, 6 33,7 9 ?(2 oviger-
ous). Upper Barron Catchment:
QMW21911 Gwynne Creek at
Gillies Road crossing (17°20.3’S,
145°31.1°E, 750m), open grazing
land with small pocket of riparian
rainforest upstream, fringing para
grass, water depth 0.3-0.6m, hand-
netted, 5.12.94, c. 200 specimens
(1.28-5.84mm CL); QMW21912
upper Gwynne Creek in rainforest
pocket (17?23.3'S, 145?30.3'E,
720m), hand-netted, leaf litter,
water depth 0.2-0.4m, 5.12.94, 14
specimens (4.0-5.4mm CL).

DIAGNOSIS. Body slender;
rostrum long, reaching tip of
antennular peduncle (RL
>0.5CL), dorso-ventrally com-
pressed, may be armed with
one dorsal tooth; antennular
peduncle long (A1P>0.5CL),
antennal peduncle long
(A2P>0.6CL), stylocerite long
(ST>0.4CL), scaphocerite long
(SC>0.8CL), sixth abdominal
segment long (6S>0.5CL) with
acute postero-ventral margin;
telson long and slender
(T>0.6CL); dorsal telsonic
spines confined to posterior
half of telson, median spine on
posterior telsonic margin ab-
sent; protopod of uropods acute
but not spinose.

DESCRIPTION. Body (Fig. la) small, sub-
cylindrical; d d in collection up to 4.9mm CL,
9 9 up to 6.2mm CL.

Cephalothorax (Figs. 2i, j; 4a, b) rotund, gla-
brous, breadth c. 0.7 CL, depth c. 0.7 CL; rostrum
long, 0.43-0.76 CL, length 12-16 X height, curv-
ing downward or sigmoid, reaching base to tip of
distal segment of antennular peduncle, asetose,
dorsoventrally compressed, a dorsal tooth may be
present. Antennal spine short, strong, placed on
lower orbital angle; pterygostomian angle obtuse,
pterygostomian spine absent. Eyes large, c. 0.25
CL, corneal diameter c. equals eyestalk length,
retinal pigmentation present. Antennular pedun-
cle shorter than scaphocerite, 0.6-0.7 CL;

FIG. 2. a, C. zebra, from Wright's Ck, ovig. 9; b, C. zebra, from Kauri Ck,
ovig. 9;c, C. zebra, from upper Gwynne Ck, 9 ; d, C. zebra, from Peterson
Ck, d; e, C. zebra, from upper Tully River, 9; f, C. zebra, from upper
Gwynne Ck, 9; g, C. spinula, 9; h, C. spinula, 3; i, C. confusa, from
Gywnne Ck, ovig. 9; j, C. confusa, from Gwynne Ck, d. Scale = 1mm.

stylocerite length 0.7 X proximal antennular seg-
ment length; anterolateral angle of proximal seg-
ment acute, reaching to about 0.15 X intermediate
segment length; intermediate segment 0.7 X
proximal segment length, about 1.7 X distal seg-
ment length; all segments with submarginal plu-
mose setae; distal segment fringed laterally and
apically with plumose setae. Antennal peduncle
0.5-0.6 X scaphocerite length; scaphocerite
slightly longer than antennular peduncle, 0.8-1.0
CL, outer margin straight to slightly concave,
asetose, ending in strong subapical spine, length
3.5 X width, distal lamella and inner margin with
plumose setae. Branchial formula typical for
genus.

Mandibles dimorphic, without palp; right man-
dible with 6-8 strong, sharp incisor teeth laterally;

NEW FRESHWATER SHRIMPS 29

FIG. 3. a, C. zebra, telson and protopod of uropod, 3,
4.8mm CL; b, C. confusa, telson and protopod, d ,
432mm CL; c, C. confusa, posterior body; d, C.
zebra, posterior body; e, C, zebra, dactylus of 3rd
pereiopod of 9 ; f, C. zebra, dactylus of 3rd pereiopod
of dg, h, C. zebra, posterior margin of telson. Scales
= Imm (a-d), 0.1 mm (e-h).

medially two groups of setae. one group with bent
hamate setae, other group with finer straight plu-
mose setae; molar process ridged; left mandible
with 6-8 strong teeth; medially three groups of
setae, molar process ridged.

Maxillula with simple palp. slightly expanded
distally, with long plumose setae distally, few
simple setae proximally; lower lacinia with
broadly rounded margin, bearing several rows of
plumose and simple setae; upper lacinia broadly
elongate, inner edge straight, with several rows
af strong spiniform, hamate, denticulate and plu-
mose setae, outer and lower inner margins with
plumose setae.

Maxilla with slender tapering palp, shorter than
upper endite cleft, setose; margin and submargin
of upper and middle endite with simple, hamate,
plumose and denticulate setae; lower endite with
hamate setae; scaphognathite with regular row of
long plumose setae on distal margin, with shorter
hamate ones continuing down proximal iriangu-

lar process which has e. 11 long simple setae,
some with prominent dilation at base.

First maxilliped with broadly triangular lamel-
lar palp, ending in pointed tip, margins with plu-
mose setae; ultimate and penultimate segments of
endites indistinctly divided; inner margin of ulti-
mate segment with long denticulate setae, long
rows of plumose. simple and hamate setae sub-
marginally, transverse rows of plumose setae
proximally; exopod flagellum distinct, well-de-
veloped, with submarginal and marginal plumose
setae; caridean lobe narrow, with marginal and
submarginal plumose setae.

Second maailliped with dactylar and propodal
segments of endopod fused; inner margins of ull
three proximal segments with long simple, ha-
mate and plumose setae; exopod long, narrow
with marginal long plumose setae distally and
shorter ones proximally.

Third maxilliped reaching beyond tip of an-
tennular peduncle; endopod three-segmented,
basal segment length c. 7 X width; penultimate
segment length c. 7 X width, c. 0.9 X basal
segment length, with transverse rows of spini-
form hamate setae; distal segment c. 0,9 X as long
as penullimate segment, ending in large claw-like
apical hamate seta surrounded by simple and
plumose ones, behind which there are 7-9 hamate
setae on distal 3rd of posterior margin, clump of
serrate and pappose setae proximally; exopod
reaching about 0.5 of 2nd endopod segment, dis-
tal margin with long plumose setae.

First pereiopod (Fig. 4c) reaching tip of basal
antennular segment; chela length 1.7-2.2 X
width, movable finger 1,! X as long as palm,
length 2.9-3.1 X width; finger tips rounded, with-
oul hooks, setal brushes well developed. Carpus
attached to chela venirally, excavated distodors-
ally, length 1.3-1.6 X width, 0.6-0.8 X chela
length, 0.94 X merus length. Merus compressed,
0.6 X as wide as carpus. Ischium length 0.41 X
merus length. Epipod present.

Second pereiopod (Fig. 4d) reaching up of 2nd
segment of antennular peduncle, more slender
and longer than Ist pereiopod. Chela length 2..4-
2.6 X width; movable finger length 4.9-5.1 X
width, 1.5 X as long as palm; finger tips without
hooks, setal brushes well developed. Carpus sub-
conical, length 3.5-4.8 X width, 1.0-1.3 X chela
length, 1.2 X merus length. Ischium length 0.67
X merus length. Epipod present.

Third pereiopod over-reaching antennular pe-
duncle tip by about 0.33 distal propodus. Dacty-
lus sexually dimorphic in adults (cf. Figs, 3e and
3f), length c. 3.8 X width, c. 0.2 X propodus

30 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. a-e, Caridina confusa sp. nov.; a, cephalothorax; b, anterior cephalothorax, dorsal view; c, Ist pereiopod;
d, 2nd pereiopod; e, margin of abdomen and telson, lateral view; f-q, C. spinula sp. nov.; f, margin of abdomen
and telson, lateral view; g, cephalothorax; h, rostrum and orbital margin, dorsal view; i, Ist pereiopod; j, 2nd
pereiopod; k, 3rd pereiopod; I, dactylus of 3rd pereiopod; m, 5th pereiopod; n, dactylus of Sth pereiopod; o,
posterior margin of telson; p, Ist pleopod of adult d ` q, endopod of 2nd pleopod of adult 3. Scales = Imm (a,
g), 0.5mm (b-f, h-k, m), 0.1 mm (l, n, o-q).

NEW FRESHWATER SHRIMPS H

length, ending in prominent claw-hke hamate
sela surrounded by simple setae, behind which
posterior margin bears 4-6 shorter spiniform ha»
mate setae, these being more robust and upright
1n adult d d. Propodus length 9-11 X width,
posierior margin and lateral surface bearing two
rows of small spiniform hamate setae. Carpus
length 0.8 X propodus length, distal projectian
feebly developed, posterior and lateral surfaces
with up to 10 small hamate setae, more spiniform
setation in adult d 3. Merus 1.1-1.4 X length of
carpus, with 2-3 strong, movable spiniform hå-
maie setae along posterior margin. Ischium 0.2 X
length of merus. Epipod present.

Fourth pereiopod reaching tip of 2nd segment
10 Up of 3rd segment of antennular peduncle,
Morphologically similar to 3rd pereropod.

Fifth pereiopod reaching tip of 2nd segment to
tip of 3rd segment of antennular peduncle, Dac-
tylus unguiculate, compressed, length c. 3.8 X
width, ending in claw-like apical hamate seta,
bearing comb-like row of 55-65 hamale setae
gradually increasing in length distally on poste-
nor margin, Propodus length 8-10 X width, 3.6 X
dactylus length, bearing two rows of 15-20 short
hamate setae on posterior margin. Carpus length
0.5-0.6 X propodus length, bearing 2-7 short ha-
mate setae, distal projection. well-developed.
Merus distinctly shorter (0.6-0.8 X) but broader
(1.8 X) than propodus, bearing 2-4 large spini-
form hamate setae. Ischium c. 0.4X length of
merus, with simple setae. Epipod absent,

Abdomen (la) well developed, rotund, gla-
brous, c. 2.8 X CL; sixth abdominal segment
clongate, c. 0.6 X CL, length-depth ratio c.1.7;
protopod of uropod (Figs. 3c, 2e) acute, aspinose;
telson (Fig. 3b) narrow, length c. 0.7 X CL, dorsal
spination (3-5 pairs) confined to posterior half af
telson; posterior telsonic margin rounded, 3-4
pairs of spine-like setae, decreasing in size inte-
riorly. median spine absent, diaeresis on telsonie
exopod 10-16,

First 3 pleopodal endopod with well developed
appendix interna arising sub-distally. Appendix
interna of 2nd. Å pleopodul endopod reaching
beyond middle of appendix masculina, with
many retinaeulae distally. Appendix masculina
subcylindrical, long hamate setae distally and on
inner lateral margin,

Live colour, translucent brown,

ETYMOLOGY. The specific epithet ischosen to high-
light the fact that this new species can be confused wilh
Caridina zebra

REMARKS. Caridina confusa sp, nov. possesses
& longer rostrum and is a much more slender and
elongate animal than C. zebra. Wis found primar-
ily in open grassland streams, flowing through
pastural land of the upper Barron (Gwynne
Creek) and the upper North Johnstone (Ithaca and
Thiaki Creeks) catchments, Smaller numbers are
found, together with C. zebra, in the small, rem-
nant rainforest areas of these streams. Despite
extensive sampling, C. confusa sp. nov, has not
been found anvwhere else.

Caridina spinula sp. nov,

MATERIAL EXAMINED. HOLOTYPE. QMNW-
21913 adult $, 4, 6mm CL, | 2mm RL, 17,6mm SI...
eust-flowing first order tributary of Leo Creek, near
crossing Of Leo Creek Mine Road (13"44.6'5.
143921 S E]. Nesbit River catchment, Mcllwraith
Range, Cape Yurk, alt. c. 450m, 12.7.95, J, Marshall,
N, Phillips. ALLOTYPE. QMW21914 adult 4,
3.2mmCL, 0.8mm RL, 12.4mm SL, same locality data
as holotype, PARATY PES. QMW219155d 2,7 2?
(2 OVigerOus), same data as holotype; QMW21422,
west-[lowing headwaters of Peach Creek (143*20'E,
13244" 8), Archer River catchment, Mellwraith Range.
Cape York Peninsula, alt, c. 500m, 15.11.94, K, Mc-
Donald, 7 33 (3,8-4.6mm CL), 34 9 (1 oun"
(2.6-5.4mm CL. ovig. ? 4.9mm).

DIAGNOSIS. Body rotund, rostrum short (RL
«0.4CL), dorsoventrally compressed but slightly
elevated towards dorsal carina, may have å tooth
on the ventral margin, reaching ta tip ol basal
antennular segment; pterygostomian angle
broadly angular to acute and spiniform; dorsal
telsonic spines. confined to posterior half of tel-
son, posterior median spine may be present; eges
large (length >1.32mm).

DESCRIPTION. Body (Fig, le) small, sub-
cylindrical; 3 å in collection up tà 2.7mm CL,
? 9 upto $.4mm CL,

Cephalothorax (Figs. Ic; 2g, hy 4g, h) round,
glabrous, breadth c, 0,8 CL, depth 0.6-0.8 CL,
rostrum short 0.20-0.33 CL, length 5-10 X height,
curving downward, may be elevated medially,
reaching tip of basal segment of antennular pe-
duncle, may be setose dorsodistally, dorsoven-
trally compressed, a rostral tooth may be present
on the ventral margin. Antennal spine short,
placed on orbital angle; pterygostomian angle
broadly angular (in Peuch Creek specimens) or
acure, spiniform (in Leo Creek specimens), Eyes
large, c. 0.2 CL. corneal diameter c. equal eve-
stalk length, retinal pigmentation present, At-
tennular pedunele shorter than scaphocerite,
0.6-0,7 CL; styloverite length 0,7 X proximal

32 MEMOIRS OF THE QUEENSLAND MUSEUM

antennular segment length; anterolateral angle of
proximal antennular segment acute, reaching lo
about 0.15 X intermediate segment length; inter-
mediate segment 0.7 X proximal segment length,
about 1.7 X distal segment length; all segments
with submarginal plumose setae; distal segment
fringed laterally and apically with plumose setae.
Antennal peduncle 0.5-0.6 X scaphucerite length;
scaphocerite slightly longer than antennular pe-
duncle, 0.6-0.7 CL. outer margin straight to
slightly concave, asetose, ending in strong sub-
apical spine, length 2.9-3.0 X width, distal la-
mella and inner margin with plumose setae.
Branchial formula typical for genus.

Mandibles dimorphic, without palp; right man-
dible with 5-6 strong, sharp incisor teeth laterally;
medially two groups of setae, one group with bent
hamate setae, other group with finer straight plu-
mose setae; molar process ridged; teft mandible
with 5-6 strong teeth; medially three groups of
setae, molar process ndged

Maxillula with simple palp, slightly expanded
distally, with long plumose setae distally, few
simple setae proximally; lower lacinia with
broadly rounded margin, bearing several rows of
plumose and simple setae; upper lacinia broadly
elongate, inner edge straight, with several rows
of strong spiniform, hamate, denticulate and plu-
mose setae, outer and lower inner margins with
plumose setae,

Maxilla with slender tapering palp, shorter than
upper endite cleft, setose; margin and submargin
of upper and middle endite with simple, hamate,
plumose and denticulate setae; lower endite with
hamate setae; scaphognathite with regular row of
long plumose setae on distal margin, with shorter
hamate ones continuing down proximal triangu-
lar process which has c. I1 long simple setae,
some with prominent dilation at base.

First maxilliped with broadly triangular lamel-
lar palp, ending in pointed tip, margins with plu-
mose setae; ultimate and penultimate segments of
endites indistinctly divided; inner margin of ulti-
mate segment with long denticulate setae, long
rows of plumose, simple and hamate setae sub-
marginally, transverse rows of plumose setae
proximally; exopod flagellum distinct, well-
developed, with submarginal and marginal plu-
mose setae; caridcan lobe narrow, with marginal
and submarginal plumose setae,

Second maxilliped with dactylar and propodal
segments of endopod fused; inner margins of all
three proximal segments with long simple harnate
and plumose setae; exopod long, narrow with

TABLE |. Morphometric ratios (range) of pereiopods
ot Caridina zebra Short, 1993, C, confusu sp. nov.
and C. spinula sp. nov. D, dactylus; P, propodus; C,
carpus; M, merus; L, length; W, width. 1, 2, 3 and 5
refer to the corresponding pereiopods.

[øm ivret!
reen "rg
CLEN

KEE

M3L/P3 11-13 LIU

Sars

PSL/PSW 115-145 RO-11.0

CSL/CSW 42357 4.146 2,9-5,8

MSL/M5W 5.7-7.5 5.6-6.0

CSL/PSL äng — [0505 0405 |

DES 1805

o EE — [pm A

marginal long plumose setae distally and shorter
ones proximally.

Third maxilliped reaching beyond tip of an-
iennular peduncle; endopod three-segmented,
basal segment length c. 6.5 X width; penultimate
segment length c. 8 X width, c. 0.9 X basal segment
length, with transverse rows of spiniform hamate
setae; distal segment c, 0.9 X as long as penulti-
male segment, ending in large claw-like apical
hamate seta surrounded by simple and plumose
ones, behind which there are 7-9 hamate setae on
distal third of posterior margin, clump of serrate
and pappose setae proximally; exopod reaching
about 0.5 of 2nd endopod segment, distal margin
with long plumose setae. l

First pereiopod (Fig. 4i) reaching tip of basal
antennular segment; chela length 1,9-2.2 X
width, movable finger 1,1 X as long as palm,
length 2.0-3.4 X width; finger tips rounded, with-
out hooks, setal brushes well-developed. Carpus
attached to chela ventrally, excavated distodors-
ally. length 1.6-2.5 X width, 0.7-1.0 X chela
length, 1,0 X merus length, Merus compressed,

NEW FRESHWATER SHRIMPS 33

TABLE 2. Distinguishing characteristics of Caridina zebra Short, 1993, rounded by simple setae, behind
C. confusa sp. nov. and C, spinul sp. nov. CL, post-orbital carapace which posterior margin bears 4-5

length.

mer ZW

Thstrihitian mainly in rainforese
covered streams of
the upper Tully,
Herbert, Johnstone
and Barron
catchments

Rostnun tà tip of Znd segment
of antennat
peduncte, length.
0.21-0.50 CL

0.58 + 0,01 CL.

lengilr
Antennal peduncle ,
length

Stylocerive
aptennular peduncle
‘segment, length 0,30
+0,04CL

Lenyth of ft
abdominal segruent

Length-depth ratio of | 1.26 + 0.07
Gih abdom. segment

DAR t LOU

short, broad, 0.53 +
0.02 CL. anterior
telsonic width 0.44 +
11.01 1alsonic length,
posterior (elsonic
width 0.19 4 0.01
Telsonic length

Dorsal telsonit to pasterior wo

Dev. epp size (mm). | 0,70-0.80.x 1.22-1.32

Scaphocerite length | 0.61 + 0,02 CL. 0,94 + 0,03 CT. 0,62+ 002 CL

Plerygostamian angle | bluntly angular bluntly angular bhunthy angular to
acutá, spiniform

Protopod of uropod

Undey, egg (mm) 0.63-0.81 x 1.02-1.21 | U,63-0,85 x 0.98-1.36 | (,8O-.NS x 1,30-1,44
(1.73-0.88 x 1.22-1.44 | not avatlable
Number oggs/ female | 38-57, mean 45 28-74, inean 51,5

shorter spiniform hamate setae,

these being more robust and up-
e edad cu right in adult d 3. Propodus length
peche? af streams ol" | streams of Nisbet und 10,5- 13.0 X width, posterior mar-
the upper Johnstone | Archer egtetiments in gin and lateral surface bearing
and Barron the Melllwraith rows of small spiniform hamate
catchment: Range, Cape Vork setae. Carpus length 0.6-0.8 X pro-
fo beyond tip of 2nd | la tip of Ist segrnent podus length, distal projection fee-
segment of of antenfular bly developed, posterior and
meire) epe pm lateral surfaces with 1 large and up
1.692 0.011 CL 055+ 001 CT. lo 5 small hamate setae, more

spiniform setation in adult d d.
Merus 1.6-1.9 X length of carpus,
with 2-4 strong, movable spini-
Form hamare setae along posterior
margin. Ischium 0.3 X length of
metus. Epipod present.

Fourth pereiopod reaching tip of
2nd segment to tip of 3rd segment
of antennular peduncle, morpho-
logically similar to 3rd pereiopod-

Fifth pereiopod (Fig. 4m, n)
reaching tip of 2nd segment tå tip
of 3rd segment of antennular pe-

mot tn tip of Ist not to tip of lat
antennular peduncle | antznnular peduncle
segment, length O45 | segment, Inget 0,32
200505 s001CL

0.50 0.11 CL

LOR + (1:03

1.65 = 0.08

elongate, spinose

elongate, narrow. short, broad, 0,54 £

0,75 -- 0.04 CL, 0.05 CL, anyenor 4 ull
anterior telsanic telsonic width DA): duncle. Dacty lus un guicu late,
widih(39 001 ` | 001 telsonic lengib. compressed, length c. 4.0 X widti,

delsomte length.
posterior telsonic
width L5 500!
telsonic length
confined to posterior

posterior telsenic
width 1.20 + 0.01
islsonre lengir

ending in claw-like apical hamate
seta, bearing comb-like row of 45-
55 hamate setae gradually increas-
ing in length distally on posterior

Jo postener puo

spination ihiras of telson half of telson Hints of telson å 3. F
Telsonic margin methan sping may be | median spine absent | median spinte may be ina Tii. Propodus length 12-18 X

present »-— width, 3.3 X dactylus length, hear-
1622 ing rows of 10-15 short hamate

setae on posterior margin. Carpus
length 0.4-0.6 X propodus length,
bearing 2-7 short hamate setae,
distal projection well-developed.

17-18, mean 17,5

0.6 X as wide as carpus. Ischium length 0.41 X — Merus shorter (0.8-0,9 X) but broader (1.5 X)

merus length. Epipod present.

Second pereiopod (Fig. 4j) reaching tip of 2nd
segment of antennular peduncle, more slender

than propodus, bearing 2-4 large spinifurm ha-
male setae. Ischium c. 0.3 X length of merus, with
simple setae. Epipod absent.

and longer than Ist pereiopod, Chela length 2.4- Abdomen (1c) well-developed, rotund, gla-

2.9 X width; movable finger

length 3.2-4,5 X brous, c. 3 X CL; 6th abdominal segment elon-

width, 1.6 X as long as palm; finger tips without pate, c. 0.5 X CL, length-depth ratio c.1,8;
hooks, setal brushes well-developed. Carpussub- ` protopod of uropod acute, aspinose; telson (Fig.
conical, length 5.8-6.1 X width, 1.4-2.0 X chela — 40) broad, length c. 0.5 X CL, dorsal spinåtion
length, 1.1 X merus length. Ischium length 0.48 — (3-5 pairs) confined to posterior 0.66 of telson;
X merus length. Epipod present. posterior telsonic margin rounded, 4-5 pairs of

Third pereiopod (Fig. 4k, 1) over-reaching an-
tennular peduncle tip by about 0.33 distal pro-
podus. Dactylus sexually dimorphic in adults,

spine-like setae, decreasing in size anteriorly, me-
dian spine may be present; diaeresis on telsonie
exopod 16-22.

length c. 2.6 X width, c. 0.25 X propodus length, First å pleopodal endopod (Fig. 4p) with well-

ending in prominent claw-like

hamate seta sur» developed appendix interna arising sub-distally.

34 MEMOIRS OF THE QUEENSLAND MUSEUM

GE interna of 2nd 3 ver X2 endopod
4q) reaching beyond middle of appendix
e with many retinaculae distally. Ap-
pendix masculina subcylindncal, long hamate
setae distally and on inner lateral margin.
Live colour, translucent brown.

ETYMOLOGY, The specific name refers to the dis-
linctive, spiniform pterygostomian angle which may be
present in some specimens. No other Australian Car-
idina species exhibits this spiniform plerygostomian
angle.

REMARKS. Although C. spinula sp. nov. looks
very much like C. zebra, there are distinct mor-
phological differences (Tables | and 2). The
spiniform pterygostornian angle m specimens
from Leo Creek is ulso very distinctive. This
species is currenily known only from the
Mellwraith Range in the Cape York Peninsula
und, despite extensive sampling, has not been
found on the Atherton Tablelands around the
Lamb-Francis-Cardwell Ranges (where C. zebra
and C. confusa sp. nov. are found), The only other
likely area of its occurrence between the Atherton
Tableland and the McIlwraith Range that we have
not sampled is the Cape Tribulation/Daintree
area. However, none were found here during
other sampling trips (J. Short and B. Herbert.
pers. comm.), The Leo Creek tributary from
where Caridina spinula sp. nov. was collected
consists of a series of riffles and pools no more
than 3m long, Im wide and 0.3m deep, flowing
over a substrate of bedrock with some sand and
gravel, Discharge at time of sampling was ap-
proximately 1 Ls. Vegetation in the area is trop-
ical mesophyll rainforest with a closed canopy at
only 6-8m and some emergent vegelation. The
low canopy suggests that this area may have
recently been disturbed (David Hanger, pers.
comm.). Density of the shrimp was low (1-2 D
Other ånimals recorded from the collection site
include the prawn Macrobrachium tolmerum
(Decapoda: Palaemonidae) and the frogs Litoria
genimaculata, L. longirestrus (Anura: Hylidac),
Sphenophryne gracilipes (Anura: Microhylidae)
and Rana daemeli (Anura: Ranidae).

DISCUSSION

The Caridina typus species-group.
characterised by its short, dorsally unarmed ros
trum, can be identified using the key of Choy
and Horwitz (1995) (to couplet 6, p. 52), The
four species can then be identified using the
following key.

]. Rostrum long, extending beyond tip of second
segment of antennular peduncle, 0.4-0.8 times
carapace length; stylocerite long, 0.4-0.5 times
carapice length; sixth abdominal segment long,
0.55-0.59 times carapace length

die CP A Caridina confusa sp.nov
Rosin rum short, not reaching tip of second seg-
ment of antennular peduncle, 0.2-0.5 times cara-
pace length; stylocerite and sixth abdominal
segment short, «0,4 and «0.5 times carapace
length respectively

2. Rostrum somewhat laterally flattened, with 1-5
teeth on ventral margin; posterior telson margit
angular, median plumose setae on posterior lel-
son margin with selerotinous plug: eggs small,
<0.6mm long C. rop

Rostrum somewhat dorsoventrally flattened, ven-
tral teeth usually absent although one may be
present; posterior telson margin rounded, me-
dian plumose setae without sclerotinous plug;
eggslarge,20.8mmlong .....

3. Rostrum short, not extending beyond up of first
segment of antennular peduncle; length to depth
ratio of sixth abdominal segment >1.4, number of
eggs curried by female «25 C. spinula sp. nov.

Rostrum relatively long, usually extending be-
yond tip of first segment of antennular pedun-
cle, length 10 depth ratio of sixth abdominal
scgment «1.4, number of eggs carried by fe-
male 230 . ea omms rms Ca EDU

Caridina adr Short, 1993, C. confusa sp. nov.
and C. spinula sp, nov, may also be separated on
the basis of distinguishing characters given in
Tables 1 und 2. It is emphasised that individual
characters may be highly variable ånd so a com-
bination of characters should be used to confirm
the identity of keyed out specimens. The relation-
ship between carapace length, rostrum, and the
sixth abdominal segment lengths are linear for all
three species (Fig. 5). The slopes and intercepts
of the regression lines are significantly different
for C. confusa when compared to the other two
species (P>0.05). All three species tend to be
allopatric. It i$ only in and near the short rainfor-
est reaches of Gwynne and Thiaki Crecks that C.
zebra and C. confusa are sympatric. Although C.
zebra has been collected from some open, para
grass infested and anthropogenically disturbed
reaches of streams (e.g., Raspberry, Ithaca und
Prior's Creeks) it does not appear 10 be very
tolerant of these conditions. It seems to prefer the
rainforest reaches of stream. C. zebra is a rotund
animal, more commonly found on sandy, silty
and/or leaf litter beds of riparian-covered rainfor-
est streams and is particularly abundant in the
more elevated, cooler areas (c. 800-950m), where
fish predators and crustacean competitors are ah-

NEW FRESHWATER SHRIMPS 35

4 T——T T T T 4 T T T T T
Caridina zebra C. zebra
3r 4 €
"e y=0.084+0.317x, r-0.718, P«0.05 E
£ £
e 2
2 e
n e E
5 oO
3 i
1 £
E
D
[0] L 1 1 1 a
1 2 3 4 5 6 7 e 1 1 1 L

andi
1 2 3 4 5 6 7

Carapace Length (mm)
Carapace Length (mm)

4 [——T T T

Caridina confusa C. confusa Ü

Rostrum Length (mm)
bv]
Sixth Abd. Seg. Length (mm)
N
T
1

^ y=-0.124+0.607x, r-0.842, P<0.05

[9] L L L L l
1 2 3 4 5 6 7 [0] 1 Ge 4 i -
2 3 4 5 6 7

Carapace Length (mm)
Carapace Length (mm)

4 T T T T T 4 T T 1 T
Caridina spinula C. spinula

L J €
€ d E
E y=0.177+0.196x, r=0.648, P<0.05 ro
£ 2
a, vod
ger J ?
E o
2 3
E Ed
a PA
1 x
[n]

[e] L L L 1 L [6] l L L L 1

1 2 3 4 5 6 7 1 2 3 4 5 6 7
Carapace Length (mm) Carapace Length (mm)

FIG. 5. Correlation between carapace length, rostrum length and sixth abdominal segment length of Caridina
zebra, C. confusa sp. nov. and C. spinula sp. nov. All regressions are significant (P<0.05). The slopes and
intercepts of the regression lines of the appropriate variables between C. confusa and C. zebra and between C.
confusa and C. spinula are significantly different (ANOCOVA, P>0.05).

36 MEMOIRS OF THE QUEENSLAND MUSEUM

sent, C. confusa sp. nov. is a more elongate,
slender animal more commonly found amongst
the bank vegetation (particularly para grass) of
streams flowing through open grassland areas of
the Atherton Tableland.

It is likely that both the species were present in
these previously rainforested areas. The modified
environment may be favouring C. confusa, hence
its predominance in these areas.

ACKNOWLEDGEMENTS

Stuart Bunn, Chris Marshall and David Hur-
wood of Griffith University, Brett Herbert of the
Department of Primary Industries, Walkamin and
Mark Hopper of the Department of Natural Re-
sources, Mareeba assisted in collecting the sam-
ples. Peter Davie allowed access to the
Queensland Museum collection and John Short
assisted in many ways, including helpful discus-
sion. Funding from the Co-operative Research
Centre for Tropical Rainforest Ecology and Man-
agement and the Wet Tropics Management Au-
thority (through Griffith Universily), the
Monitoring River Health Initiative Program and
the Department of Natural Resources made pos-
sible the expeditions to collect samples. Stuart
Bunn, Jane Hughes and John Short provided con-
structive criticism of the draft manuscript.

LITERATURE CITED

BOUVIER, EL 1925. Recherches sur la morphologie,
les variations, la distribution géographique des
Crevettes de la famille des Atyidés. Encyclopédie
Entomologique, Sér. A 4: 1-370.

CHOY, S, 1991. The atyid shrimps of Fiji with descrip-
tion of a new species. Zoologische Medelingen
65(27): 343-362.

CHOY, S. & HORWITZ, P. (1995). Preliminary key to
the species of Australian shrimps (Atyidae) found
in inland waters. Pp. 51-54. In Horwitz, P, "A
preliminary key to the species of Decapoda
(Crustacea: Malacostraca) found in Australian in-
land waters. Identification Guide No. 5.' Co-oper-
ative Research Centre for Freshwater Ecology,
Albury.

FELGENHAEUR, B.E. (1992). External anatomy and
integumentary structure. Pp. 7-43. In Harrison,
EW E Humes, A.G. (eds), ‘Microscopic anatomy
of invertebrates’ Vol. 10. (Wiley-Liss: New York).

HOLTHUIS, L.B. 1965. The Atyidae of Madagascar.
Memoires du Museum national d'Histoire
Naturelle (N.S.) (A), Zoologie 33(1): 1-48.

HUGHES, J.M., BUNN, S.E, HURWOOD, D.A.,
CHOY, S.C. & PEARSON, R.G. 1996. Genetic
differentiation among populations of Caridina
zebra (Decapoda; Atyidae) in the tropical rainfor-
est streams, northern Australia. Freshwater Biol-
ogy 36: 287-296.

MILNE EDWARDS, H. 1937. Histoire naturelle des
Crustaces, comprenant l'anatomie, la physiologie
et la classification de ces animaux 2: 1-532, Atlas,
pp. 1-532, pls. 1-42.

RIEK, E.F. 1953. The Australian freshwater prawns of
the family Atyidae, Records of the Australian Mu-
seum 23: 111-121.

ROUX, J. 1926. An account of the Australian Atyidae.
Records of the Australian Museum 15(3): 237-
254.

SHORT, J.W. 1993. Caridina zebra, a new species of
freshwater atyid shrimp (Crustacea: Decapoda)
from northeastern Queensland rainforest. Mem-
oirs of the Queensland Museum 34(1): 61-67.

GASTROPODS FROM THE BURDEKIN FORMATION, MIDDLE DEVONIAN, NORTH
QUEENSLAND

ALEX G. COOK

Cook, Alex G. 1997 06 30: Gastropods from the Burdekin Formation, Middle Devonian,
north Queensland. Memoirs of rhe Queensland Museum 42(1): 37-49. Brisbane, ISSN

0079-8835.

Twenty one taxa of gastropods are described from the Middle Devonian Burdekin Formation,
north Queensland. New amongst these are Burdekinostoma burdekininensis gen. et. sp. nov.,
Euryzone burdekinensis sp.nov., Anomphalus pajelli sp. nov., Didymalgia bartholomai gen.
el sp. nov., Murchisonia (Murchisonia) jackjelli sp. nov., Palaeozygopleura machenryi sp.
nov., and Trinema heideckeri gen. et sp. nov. The fauna contains typical Old World Realm
genera but is specifically strongly endemic. [ ] Gastropods, Devonian, Queensland, Burde-

kin Formation,

Alex G, Cook. Queensland Museum, PO Box 3300, South Brisbane, Queensland 410],

Australia; 10 March 1997.

Burdekin Formation is å shallow marine car-
bonate sequence exposed within the Burdekin
Subprovince (Wyatt & Jell, 1967: Henderson,
1980; ). The formation consists of, inter alia, a
reefal and biostromal complex, with associated
lagoonal. inter-reef and offshore facies (Cook,
1995). It contains diverse fossil assemblages
which include rugose corals (Zhen 1991, 1994),
tabulate corals, stromatoporoids (Cook, 1994),
sponges, molluses (Cook, 1993a) and conodonts
(Talent & Mawson, 1994). Large, thick-shelled
gastropods from the underlying and interdigitat-
ing. siliciclastic-dominated, Big Bend Arkose
and basal units of the Burdekin Formation are
Burdikinia burdekinensis (Etheridge, 1917),
Amphelissa carinatum (Heidecker, 1959),
Labrocuspis nodosa Heidecker, 1959, and
Fletcherviewta septata Cook, 1993b.

This paper is concerned with a diverse gastro-
pod faunule from a stratigraphically higher
unit in the Burdekin Formation in coralline
packstones interpreted to represent shallow-
water coralline thickets seaward of a biohermal
structure (Cook, 1995), The fauna has been
silicified and was recovered by bulk dissolution
in acelic acid. Both large and small gastropods
were retrieved, in addition to corals, brachiopods
and bivalves. All material comes from
QML1094. near Little Rocks, 27km NW of Char-
ters Towers.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA
Class GASTROPODA
Order ARCHAEOGASTROPODA
Superfamily BELLEROPHONTOIDEA
Family BELLEROPHONTIDAE McCoy, 1851
Subfamily TROPIDODISCINAE Knight, 1956

Tropidodiscus Meck & Worthen, 1866
Tropidodiscus sp.
(Fig. 1A-C)

MATERIAL EXAMINED. QMF35513, QMF35514.

DESCRIPTION. Shell minute, approximately
2.5mm in diameter, 0.8mm wide; involute with
moderately wide umbilicus. Whorl profile
sharply angular, without obvious crest. Relics of
fine, numerous growth lines present.

Bellerophon de Montfort, 1810
Bellerophon (Bellerophon) de Montfort, 1810

Bellerophon (Bellerophon) sp. Å
(Fig. 1D,E)
MATERIAL EXAMINED. QMF34916, QMF34902,

DESCRIPTION. Small to medium-sized, spher-
ical, involute, 5.5mm wide; narrow deep umbili-
cus. Whorl profile evenly rounded; margin
bearing weak crest. Aperture unknown. Orna-
ment consists of many fine growth lines,

REMARKS. The few specimens possess similar
characteristics to the type, B.(B.) vasulites de
Montfort, 1810 from the Middle Devonian of

38 MEMOIRS OF THE QUEENSLAND MUSEUM

Germany as figured by Knight (1941). The nu-
merous growth lines and overall form are similar,
but the Burdekin form is much smaller (Knight,
1941: pl. 11) and the selenizone-bearing crest is
weaker, possibly due to imperfect silicification.

Bellerophon (Bellerophon) sp. B
(Fig. IEG)

MATERIAL EXAMINED. QMF34908.

DESCRIPTION. Spherical, medium-sized, ap-
proximately 10mm in diameter, 9mm high; invo-
lute with deep, wide umbilicus. Whorl profile
wide and rounded with weak crest; slight flatten-
ing of inner whorl surface adjacent to penultimate
whorl, extending approximately one third across
the inner whorl profile.

REMARKS. The specimen is larger and has a
wider whorl profile than Bellerophon
(Bellerophon) sp. A, which is a result of the
flattened area on the whorl profile abutting the
penultimate whorl.

Superfamily EUOMPHALOIDEA
de Koninck, 1881
Family EUOMPHALIDAE de Koninck, 1881

Straparollus de Montfort, 1810
Straparollus (Euomphalus) Sowerby 1814

Straparollus (Euomphalus) sp.
(Fig. 1Q)

MATERIAL EXAMINED. QMF34965.

DESCRIPTION. Medium-sized, planispiral,
21.4mm wide, 8.2mm high. Suture in a channel.
Whorl profile square with prominent angulation
above and below gently convex midwhorl sur-
face. Weak spiral line adumbilically to the upper
angulation which may mark the position of the
UD Aperture rounded, but quadrate. Base flat-
tened.

Straparollus (Serpulospira) Cossman, 1916
Straparollus (Serpulospira) sp.
(Fig. IR)

MATERIAL EXAMINED. QMF34966.

DESCRIPTION. Shell planispiral, disjunct,
13.6mm wide, 4.6mm high. Whorl profile
rounded, aperture rounded. No growth lines pre-
served.

Suborder PLEUROTOMARIINA
Cox & Knight, 1960
Superfamily PLEUROTOMARIOIDEA
Swainson, 1840
Family RAPHIOSTOMATIDAE Koken, 1986
Subfamily OPHILETINAE Knight, 1956

Burdekinostoma gen. nov.

TYPE SPECIES. Burdekinostoma burdekinensis sp.
nov.

ETYMOLOGY. For the Burdekin River, and alluding
to the Raphiostomatidae.

DIAGNOSIS. Minute, low-spired to lenticular,
gradate shell, widely phaneromphalous, having a
commonly channeled suture and prominent,
gently inclined, flattened area on the upper whorl
surface bordered by 2 strong cords, inferred to
contain the selenizone and also commonly con-
taining weak threads; midwhorl profile generally
subrounded but stepped with several cords.

REMARKS. Placement of this striking if minute
genus is difficult, as it superficially resembles
members of the Ophiletinae, Raphiosomatinae
and Euomphalidae. Placement depends on the
interpretation of the inclined surface on the upper
whorl face, which is inferred to contain the
selenizone. In particular the raphiostomine
Wisconsonella Blodgett (1988) from the Eifelian
of North America is a similar taxon to the Burde-
kin material, sharing the gradate shell form, but
lacking the additional cords and the channeled
suture, Placement within the Raphiostominae is
restricted by the widely phaneromphalus nature
of this snail. Alternatively the channeled suture
may be interpreted as containing a sinus, which
would ally the genus to the Euomphalidae, of
which Poleumita Clarke & Ruedemann and Cen-
tifugus Bronn are similar cord-bearing forms.
Neither of these placements is satisfactory. Place-
ment in the Ophiletinae best satisfies the interpre-
tation of the selenizone, accomodates the
channelled suture and the phaneromphalous con-
dition.

Burdekinostoma burdekinensis sp. nov.
(Fig. 2J-P)

MATERIAL EXAMINED. HOLOTYPE: QMF34961.
PARATYPES: QMF35522-QMF35530.

ETYMOLOGY. From the Burdekin River.
DIAGNOSIS. As for genus.

GASTROPODS FROM THE BURDEKIN FORMATION 39

FIG. 1. A-C. Tropidodiscus sp. SEM images. A,B, QMF35512, side and oblique views respectively, x 16; C,
QMF35513, side view, x 16. D-E. Bellerophon (Bellerophon) sp. A. QMF34916, apertural and side views
respectively. F, G. Bellerophon (Bellerophon) sp. B. QMF34908, side and broken apertural views respectively.
H-P, Euryzone burdekinensis sp. nov. H-J, Holotype QMF34917, apical apertural and basal views respectively,
x 4,4; K,L, Paratype QMF 34969, basal and apical views, x 3.2; M,N, Paratype QMF34964 apical and basal
views x 6.3: O, Paratype, QMF35536 oblique side view, x 22.5, SEM image; P, Paratype QMF35536 apical
view x 22,5, SEM image. Q, Straparollus (Euomphalus) sp. QMF34965 apical view, x 1.8. R, Straparollus
(Serpulospira) sp. QMF34966, apical view x 2.3.

40 MEMOIRS OF THE QUEENSLAND MUSEUM

DESCRIPTION. Shell small to minute, less than
4.5mm wide and 1.5mm high, very low-spired,
somewhat gradate, widely phaneromphalous,
with deep umbilicus. Whorl profile series ot an-
gular surfaces but giving an overall rounded pro-
file. Suture with adjacent, wide channel. Upper
whorl face bears numerous spiral threads, of at
least 2 orders of intensity. Three prominent
threads on smaller specimens; bordering ihe su-
tural channel, another high on the whorl profile
and another at the edge of the upper whorl face
giving way to the steeper mid whorl surfaces, the
last 2 inferred to contain the selenizone. At least
2 threads on the midwhorl, creating å vertical
surface at the midwhorl periphery: at least 3
revolving cords on the lower whorl face. The
upper whorl face bears up to 7 minor spiral
threads, in addition to the major cords.

REMARKS. This striking taxon is easily differ-
entiated by Euryzone burdekinensis sp. nov. by
the numerous spiral threads and more angular
whorl profile.

Family GOSSELETINIDAE Wenz, 1938
Sublumily COELOZONINAE Knight, 1956

Euryzoue Koken, 1896
Euryzone burdekininesis sp. nov.
(Fig. (HP

MATERIAL EXAMINED. HOLOTYPE: QMF24917.
PARATYPES: QMF34918-QMF34927.QMF34964,
QMF34969.

DIAGNOSIS. Small to minute member of genus
with sinus on upper whorl profile bouded by fine
ihreads,

DESCRIPTION, Shell discoidal, minute to small,
up to 11.9mm wide, 3,6mm high, generally
smaller (Table 1). Whorls abut, with impressed
sulure. Base with moderately deep umbilicus.
Whorl profile rounded, bearing two fine cords
which border the sinus. Growth lines fine, numer-
ous. Shell repair in evidence on holotype.

REMARKS, The species is characterised by its
discoidal shape and the striking presence of the
sinus on the upper whorl face and its bordering
cords. The most similar member of the genus to
the Burdekin material is Euryzone perilitornara
Linsley 1968 from the Middle Devonian An-
derdon Limestone, Michigan, but its selenizone
is slightly lower on the whorl profile, and it is à
less planispiral than the Burdekin species. The
type species E. delphinuloides (Schotheim) from

TABLE |. Measurements lor Euryzene burdekinensis
Sp. DON,

QMF34969

QMF34964 20
urn il

the Middle Devonian Stringocephalus Lime-
stone, Germany, has a turbiniform shell and a
wider selenizone.

Suborder TROCHINA Cox & Knight, 1960
Superfamily ANOMPHALOIDEA Wenz, 1938
Family ANOMPHALIDAE Wenz, 1938

Anomphalus Meek & Worthen, 1567

TYPESPECIES. Anomphalus rotulus trom the Middle
Pennsly vanian, St Davids Limestone, Illinois, USA, by
original designation,

Anomphalus pajelli sp. nov.
(Fig. 2A-T)

MATERIAL EXAMINED. HOLOTYPE: QMF34850,
PARATY PES: OMF34851-OMF343858.

ETYMOLOGY, For Peter A. Jell.

DIAGNOSIS. Smooth-shelled, medium-sized
member of genus with very weak sutures and
prominent thickening of inner lip which creates a
hemiomphalus base.

DESCRIPTION. Medium-sized, low-spired,
rotelliform hemiomphalus gastropod, up tà 11.5
mm high, 13.5 mm wide (Table 2). Sutures flush,
commonly difficult to detect, whorls embrace
high on upper whorl surface. Umbilicus narrow
obscured or occluded by thickening at the inner
labrum, otherwise base rounded, Whorl profile
rounded with periphery at midwhorl. Aperture
rounded. Shell thick, inner labrum particularly
thickened forming partial umbilical plug. Shell
smooth, very faint growth lines on the base of
QMF34851. There is some variation in the taxon
with respect to where the whorls embrace. On
QMF34851 the whorls embrace further down the
whorl face, producing a slightly more naticiform
shell shape. Despite silicà replacement the

smooth shell has an almost polished appearunce

on some specimens.

GASTROPODS FROM THE BURDEKIN FORMATION 4l

FIG. 2. A-I. Anomphalus pajelli sp. nov. A-C, Holotype QMF34850, apertural, apical and basal views, x 9. D-F,
Paratype QMF34852, apertural, basal and apical views, x 2.2. G-I, Paratype QMF34851 apical, apertural and
basal views, x 2.5. J-P, Burdekinostoma burdekinensis gen. et sp. nov. J,K, Paratype QMF35523, basal and
apical views, SEM images, x13.5; L.M, Paratype QMF 35522, oblique apertural and apical views, SEM images,
x 15; N, QMF 35525 side view, x 13.5; O,P, Holotype QMF34961, x 5.4, apical and side views respectively.

42 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Measurments for Anomphalus pajelli sp.
nov,

| — QMF34850 13.1
OMFMRSI | us |

REMARKS. Both Knight (1941) and Knight et
al. (1960) noted the variable umbilicus in this
genus. The Burdekin material is assigned to the
genus on the basis of gross shape, lack of orna-
ment and the hemiomphalus condition, The type
species Å. rotulus Meek & Worthen as figured by
Knight (1941) has a slightly lower whorl profile
and more prominently incised sutures. An-
omphalus umbilicoliratus Batten (1966) from the
Lower Carboniferous Hotwells Limestone, En-
gland is clearly phaneromphalous and has spiral
ornament on the base. Anomphalus
helicinaeformes (Schlotheim) from the Middle
Devonian Stringocephalus Limestones near
Gladbach, Germany has a fully developed umbil-
ical plug (see Knight 1941: pl 63), but in overall
aspect is most similar to the Burdekin species,
and is closely related. Specimens illustrated by
Knight (1941) show minor variation in spire
height for A. helicanaeformis (Schlotheim).
Naticopsis margheriti Mansuy (1912), may also
be an Anomphalus, albeit with å slightly more
naticiform shell.

Suborder NERITOPSINAE Cox & Knight, 1960
Superfamily NERITOIDEA Rafinesque, 1815
Family NERITOPSIDAE Gray, 1847

Naticopsis M'Coy, 1844

Naticopsis (Naticopsis) sp.
(Fig. 4A)

MATERIAL EXAMINED. QMF34913,

DESCRIPTION. Very small, 4.9mm wide, Smm
high, naticiform, anomphalous; suture adpressed
to slightly impressed; whorl profile rounded, with
periphery well above midwhorl. Upper whorl
surface flatter that lower whorl face. Final whorl
dominated by strong collabral cords, increasing
in intensity towards the aperture. Early whorls
lack this ornament. Parietal and inner lip heavily
thickened; aperture ovate. Base rounded.

REMARKS. The specimen strongly resembles
Natica nexicosta (Phillips) of Whidbome (1892),

from the Middle Devonian at Lummaton, En-
gland, but the latter possesses strong collahral
cords on early whorls. Naticopsis ( ?Naticopsis)
sp. (Rollins, Eldridge & Spiller, 1971), from the
Middle Devanian Marcellus Formation, New-
York, shows a similar ornament, but the threads
are fine and inclined on the whorl face,
Straparollus corrugatus (Stauffer, 1909), which
Linsley (1968) placed within ”Ysonema, and Rol-
lins, Eldridge & Spiller (1971) assigned to
Naticopsis (Naticopsis), from the Middle Devon:
jan of Ohio also shows strong collabral ornament,
but not restricted to the mature whorls. ?/sarte me
corrugatus (Stautfer) from the Middle Devonian
Anderdon Limestone (Linsley, 1968) is a mi-
nutely illustrated taxon. Linsley (1968) describes
the early whorls as smooth suggesting affinity lo
the Burdekin specimen. Positive generic identifi-
cation of Linsley's taxon is impossible without
viewing his material. 1 place Natica nexicosta
(Philips) of Whidborne and Srraparollus
corrugatus Stauffer both within Naticopsys
(Naticopsis) alongside this distinct, if poorly pre-
served and unresolved, taxon from the Burdekin
Forinaton.

Superfamily ORIOSTOMATOIDEA Wenz, 1938
Family ORIOSTOMATIDAE Wenz, 1938

Didymalgia gen nov.
TYPE SPECIES. Didvmalgia bartholamai sp. nav.

ETYMOLOGY, For ‘Didymalgia’ hill, informal name
of laterite profile near the Fletcherview locality,

Didymalgia bartholomai gen. et sp. nov.
(Fig. 3)

MATERIAL EXAMINED. HOLOTYPE; QMF34896.
PARATYPES: QMF34876-OMF34895, QMF34915.

ETYMOLOGY. For Alan Bartholomai.

DIAGNOSIS. Turbiniform, widely phan-
eromphalous shell with numerous revolving
cords and strong comarginal cords forming and
cancellate structure, whorl profile angular,
formed by series of inclined surfaces.

DESCRIPTION. Medium-sized, turbinilorm,
widely phaneromphalus shell, up to 9.5mm high
and 11.1 mm wide (Table 3), Whorl face adorned
with strong revolving cords and slightly less
strong comarginal ribs, producing a cancellate
shell decoration, Whorl profile series of angu-
lar surfaces with a subsutural shelf, steeply
inclined upper whorl face, angular periphery,

GASTROPODS FROM THE BURDEKIN FORMATION 43

FIG, 3, Didymalgia bartholomai gen. et sp. nov. A-D, Holotype QMF34896, x 3 apertural, basal (x 2.8), apical
and side views (x 2.8). E-G, Paratype QMF34875. E, apertural view x 5; F, apical views x 5; G, side view x 4.

LJ, Paratypes QMF34876, x 3 apical and side views.

inclined lower surface leading to the phan-
eromphalous base, sutures embrace just below
periphery which is situated slightly below mid-
whorl. Slight subsutural shelf, 4 major revolving
cords on the upper whorl face, a major cord at the
periphery, and 7 to 8 cords on the lower whorl
face and base. Two orders of cords on the lower
whorl face. Aperture rounded to subrounded,
slightly wider than high. Thick colabral cords are
numerous almost orthocline, and are not de-
flected by a sinus or selenizone.

REMARKS. Oriostoma sp. aff O. gerbaulti
Oehlert of Blodgett & Johnson (1992) from the
Eifelian of Nevada is grossly similarto the Burde-
kin material but has a channeled subsutural zone.
Gyronema multinodosa Blodgeu & Johnson from
the Bifelian of Nevada, which could be
accomodated in Kitikamispira, has fewer cords
and more prominent nodes on the shell surface.
Placement of the genus within the Ori-

ostomatidae is problematic, in that it extends the
range of the family beyond their supposed
Eifelian demise (Blodgett et al. 1990), but the
strong ornament, lack of selenizone on the
whorl face and trochiform, phaneromphalus shell
form make the placement appropriate, if contro-
versial.

TABLE3. Data for Didymalgia bartholomai gen. et sp.

nov.
Height (mm)
9 i ia

Specimen
QMF34896

A
QMF34876
72

H MEMOIRS OF THE QUEENSLAND MUSEUM

Suborder MURCHISONIINA Cox & Kmght, 1960
Superfamily MURCHISONIOIDEA Koken, 1896
Family MURCHISONIIDAE KOKEN, 1896

Murchisonia d' Archaic & de Verneuil 1841

Murchisonia (Murchisonia) jackjelli sp. nov.
(Fig. 4B-H)

MATERIAL EXAMINED. HOLOTYPE: QMF34905.
PARATYPES; QMF34897-QMF34900, QMF34902,
QMF34903, QMF34929, QMF34932, QMF34933,
QMF34935-QMF34940, QMF34942-QMF34943,
QMF34946-QMF34948, QMF34952-QMF34953.

ETYMOLOGY. For J.S. Jell.

DIAGNOSIS. Medium-sized member of subge-
ius witharelatively wide selenizone, subrounded
whorl profile and up to 5 threads on the adult
lower whorl surface,

DESCRIPTION. Medium-sized to large, high-
spired shell, up to 41.5mm long and |1.6mm wide
at base. Suture impressed, whorls embrace at
approximately lower one third ot the whorl
height. Whorl profile subrounded but with 2 weak
peripheral cords bordering a wide selenizone, Up
to 5 additional weaker spiral threads on the lower
whorl face with 2 extremely faint threads on the
upper surface. In juvenile specimens cords bor-
dering the selenizone are stronger on early
whorls, and the lower whorl face lacks the multi-
ple threads of the adult whorls, instead 1 or 2
threads are more prominent. Aperture ovate with
minor thickening of the inner lip.

REMARKS. Poor preservation of the material
significantly impedes the study of this species.
Clearly the presence of the wide midwhorl
selenizone denotes the subgenus, and the numer-
ous weak threads on adult whorls distinguish a
new taxon. The width of the selenizone and the
numerous threads may suggest Carboniferous
Stegecoelia, butthe selenizone is at the periphery
negating that assignment. In the Middle Devon-
ian Murchisonia (Murchisonia) underwent sig-
nificant radiation into a multitude of unusual shell
forms (Knight et al. 1960; Blodgett et al., 1990)
and the included species described to date are
numerous (e.g., Whidborne, 1892). Taxa with
similar multiple threads on the whorl face are. M.
(M.) vicariana Whidborne, M. (M.)
loxanemoides Whidborne, both from the Middle
Devonian of Southern England, but the former
hàs nodose threads and the latter a thinner, lower
set selenizone.

Murchisonia (Murchisonia) sp, Å,
(Fig. AI-J)

MATERIAL EXAMINED. QMF34930.

DESCRIPTION. Small, 10.5mm high, 5.3mm
wide, high-spired, with eight whorls present. Su-
ture strongly impressed, whorls embrace well
below midwhorl; angular whorl profile with 2
prominent spiral cords bordering a selenizone.
Other decoration not preserved.

REMARKS. A selenizone on the midwhorl, bor-
dered by 2 cords is the classic murchisoniid con-
dition and the specimen is here allocated to the
subgenus on this basis. The species resembles M.
(M.) fermioni Tassell, 1982, from the Emsian ot
New South Wales, and also known from the
Givetian in the Broken River Province, but the
material is too poorly preserved to be confident
of any specific assignment.

Murchisonia (Murchisonia) sp. B.
(Fig. 4K)

MATERIAL. EXAMINED. QMF35520.

DESCRIPTION, Shell small, 6mm high, 2mm
basal width, high-spired, with 7 whorls present.
Sutures impressed, whorls embrace just below
periphery. Whorl profile angular with 2 strong
cords bordering a selenizone at the periphery. the
upper cord slightly stronger. Upper whorl face
gently sloped with a weak spiral cord close to the
main omament on the margin. Lower whorl pro-
file obscured, aperture and base unknown.

REMARKS. The bordered selemzone identifies
the subgenus. It is higher-spited than Murchisonia
(Murchisonia.) sp. A. and has.a significantly dif-
ferent whorl profile than Murchisonia
(Murchisonia) jackjelli sp. nov.

Superfamily CRASPEDOSTOMIDOIDEA
Wenz, 1938 |
Family CODONOCHEILIDAE Miller. 1889

Mitchellia de Koninck, 1876
?Mitchellia sp.
(Fig. 4L,M)
MATERIAL EXAMINED. QMF35515, QMF35516,

DESCRIPTION. Two fragmental specimens,
neither of which have a gerontic stage or aperture
preserved. Small high-spired gastropod with im-
pressed suture; whorls embrace just below mid-

GASTROPODS FROM THE BURDEKIN FORMATION 45

FIG. 4. A, Naticopsis (Naticopsis) sp. QMF34913, oblique apical view, x 5. BH. Murchisonia (Murchisonia)
Jackjelli sp. nov. B, C, Holotype QMF34905, side views x 2.8; D, Paratype QMF34900, side view x 1.5; E,
Paratype QMF34897, side view x 2; F, Paratype QMF34903, side view x 2.6; G, Paratype QMF34899 apertural
view x 1.6; H, Paratype QMF35520, side view x12. I,J, Murchisonia (Murchisonia) sp. A. QMF34930, x 3.5.
I, side view; J, apertural view. K Murchisonia (Murchisonia) sp. B. side view x 10. L, M, ?Mitchellia sp. L,
QMF35515, x 20; M, QMF35516 x 20.

whorl. Whorl profile rounded, face bearing nu- lian Emsian genus on the basis of gross shell
merous, (up to 10) spiral threads of equal inten- shape. It lacks the characteristic aperture needed
sity. Aperture not preserved, but inner whorl for more definite assignment to the taxon.
profile ovate with long axis vertical.

REMARKS. Generic assignment is impossible
without more material, but the material is ques-
tionably assigned to the common eastern Austra-

46 MEMOIRS OF THE QUEENSLAND MUSEUM

Order CAENOGASTROPODA Cox, 1959
Superfamily LOXONOMATOIDEA Koken, 1889
Family PALAEOZYGOPLEURIDAE Horny 1955

Palaeozygopleura Horny, 1955

Palaeozygopleura machenryi sp. nov
(Fig. 5A-F)

MATERIAL EXAMINED. HOLOTYPE: QMF35531.
PARATYPES: QMF35532-QMF35525, QMF35539,

ETYMOLOGY. For Colin McHenry.

DIAGNOSIS. Small to minute member of genus
with moderately impressed suture and only
slightly sigmoidal ribs.

DESCRIPTION. Shell high-spired, many
whorled, up to 8 present, up to 4mm long and
1.6mm wide. Sutures impressed, whorls embrace
well below midwhorl; whorl profile rounded. Pe-
riphery at midwhorl. Whorl face bears many
slightly sigmoidal ribs, obvious on all adult
whorls. Protoconch poorly preserved, but smooth
and dextral.

REMARKS. The material is most similar to P.
joanni Linsley from the Middle Devonian An-
derdon Limestone of Michigan, but in the Burde-
kin species the ribs are less numerous and the
suture less impressed. It differs from the single
specimen Palaeozygopleura sp. described below
by the more impressed sutures and coarser orna-
ment.

Palaeozygopleura sp.
(Fig 5G)

MATERIAL EXAMINED. QMF35519,

DESCRIPTION. Minute, 3.6mm high, 1.5mm
wide, high-spired, nearly pupiform with shal-
lowly impressed suture, gently rounded whorl
profile. Ornament consists of fine, prosocline
threads on the whorl surface.

REMARKS. The single specimen has weaker
sutures than P. machenryi sp. nov, and finer pro-
socline ornament.

Family TURRITELLIDAE Woodward, 1851

Trinema gen. nov.

TYPE SPECIES. Trinema heideckeri sp. nov. from the
Middle Devonian Burdekin Formation, north Queens-
land.

ETYMOLOGY. Tri-, three, nema (Greek) thread.

DIAGNOSIS. Small to medium-sized, anompha-
lous, high-spired, many whorled shell; whorl pro-
file rounded, bearing three prominent midwhorl
cords. Suture impressed; a subsutural surface is
bounded below by a weak thread which forms the
upper limit of the deep sinus, in turn bounded
below by the highest of the three midwhorl cords.
Finer threads are present on the lower whorl face.
Growth lines are prosocline above the sinus and
opisthocline below. Whorls embrace just below
the higher of the weak threads on the lower whorl
face.

REMARKS. Placement in the family is indicated
by the overall shell form and the position of the
sinus above the midwhorl cords (see Knight et al.
1960: I317). The taxon is differentiated by the
only other Devonian turritelid (Linsley 1978),
Acanthonema Sherzer & Grabau, 1908 by lack-
ing the distinctive nodose cords. In the Late Car-
boniferous Orthonema Meek & Worthen, 1861
(see Knight 1934, 1941, Knight et al., 1960) there
are 2 prominent midwhorl cords, widely sepa-
rated below the similarly positioned sinus. In
Callispira Nelson,1947, the sinus is also posi-
tioned high on the whorl face, but the whorl face
is less rounded and there are many more stronger
cords (5-6) on the whorl face. There is a consid-
erable gap in the taxonomic record of the
Turritellidae between the appearance of
Acanthonema in the Early Devonian (Sherzer &
Grabau, 1908), their Middle Devonian record
(herein) and the numerous Carboniferous records
of Orthonema (Knight, 1934). Such a discontinu-
ity in the record may be resolved by analysis of
Late Devonian age faunas.

Trinema heideckeri sp. nov.
(Fig. SH-K)

MATERIAL EXAMINED. HOLOTYPE: QMF34954.
PARATYPES: QMF34941, QMF34945, QMF34955.

DIAGNOSIS. As for genus.

DESCRIPTION. Material somewhat fragmental,
Shell small- to medium-sized, anomphalous,
high-spired, up to 12 whorls present. Whorl pro-
file rounded overall, slightly more convex on
lower whorl face; periphery at midwhorl. Suture
impressed with subsutural face bordered below
by weak spiral thread. Midwhorl face dominated
by 3 strong spiral cords the middle of which is
slightly weaker. Between the uppermost of the
strong peripheral cords and the weak cord below

GASTROPODS FROM THE BURDEKIN FORMATION 47

FIG. 5. A-F. Palaeozygopleura machenryi sp. nov., all SEM images. A, Paratype QMF35532, x 19; B,
QMF35533 x 17; C, QMF35534, x 21; D, QMF35535, x 19; E, QMF35538, x 15; F, Holotype QMF35531, x
17. G. Palaeozygopleura sp. QMF35519, x 15. H-K, Trinema heideckeri gen. et sp. nov. H, Paratype
QMF34941, x 4.3; I, Holotype QMF34954, side view x 5; J, Paratype QMF34955, x 2.5; K, Paratype. L,
Paratype QMF34945, small specimen, x 19 SEM image. L, Trinema sp. QMF34931, x 4,3. M, ?Subulites
(Subulites) sp. QMF34934, x 4.2. N, ?Subulties (Fusispira) sp. A. QMF34912, x 6.1. O, P. ?Subulties
( Fusispira) sp. B, QMF35517, x 9, x 17 respectively.

48 MEMOIRS OF THE QUEENSLAND MUSEUM

the suture lies the trace of the deep sinus, indi-
cated by the growth lines, Three weak spiral cords
are present on the lower whorl face. Growth lines
fine, numerous, prosocline above the sinus and
opisthocline below. Whorls embrace just below
the uppermost fine thread on the lower whorl
face.

REMARKS. Trinema heideckeri is superficially
resemblant of Murchisonia species described
below, bul is clearly distinguished by the 3 mid-
whorl cords and the placement of the sinus.
Trinema sp. below is differentiated by the posi-
tion of the suture.

Trinema sp.
(Fig, 5L)

MATERIAL EXAMINED. QMF34931.

DESCRIPTION. Single fragmentary specimen,
high-spired; sulure impressed, whorl face domi-
nated by three prominent spiral cords; whorls
embrace at lowermost of these; weak spiral
thread on uppermost whorl face. Sinus positioned
above the highest thick cord. Growth lines fine
and numerous.

REMARKS. The specimen is distinct from
Trinema heideckeri sp. nov. in that the whorls
embrace higher on the whorl face, at the level of
the lower of the three major midwhorl cords.
Whether this 1s an abberrant form or a new taxon
is unclear.

Superfamily SUBULITOIDEA Lindstrom 1884
Family SUBULTIDAE Lindstrom 1884

Subulites Emmons, 1842

?Subulites (Subulites) sp.
(Fig. 5M)

MATERIAL EXAMINED. QMF34934.

DESCRIPTION. Small, high-spired, fusiform
shell, | 1.3mm high, 4.2mm maximum width, su-
tures slightly impressed; sutural angle moderate.
Whorl profile gently curved. Weak, fine, prosocl-
ine, growth lines. Aperture unknown, but shell
form suggests presence of anterior notch; inner
lip unknown.

REMARKS. Questionable assignment is made
on the basis of the general shell form, but it cannot
be conclusive without data on the columella and
the aperture.

?Subulites (Fusispira) sp. A.
(Fig. 5N)

MATERIAL EXAMINED. QMF34912.

DESCRIPTION, Fusiform, small, rapidly ex-
panding shell, approximately 4mm high and 2mm
wide. Whorls embrace just below periphery
which is situated just below midwhorl. Whorl
profile smoothly rounded. Growth lines, aperture
and inner lip unknown.

REMARKS. Lack of material prevents accurate
assignment of this taxon. It differs from the ma-
terial described below in having no subsutural
angulation,

?Subulites (Fusispira) sp. B.
(Fig. 50,P)

MATERIAL EXAMINED. QMF35517, QMF35518.

DESCRIPTION. Small, fusiform, maximum
height 4mm, width 2.2mm. Whorl profile
rounded except for prominent subsuturål surface
and angualtion high on upper whorl face. Sutures
impressed, whorls embrace just below periphery
which is slightly below the midwhorl.

REMARKS. Inadequate material and poor pres-
ervation of lip, ornament and aperture prevents
assignment of this material. It differs trom ?Sub-
ulites ( Fusispira) sp. A. in having a prominent
subsutural surface and angulation.

FAUNAL AFFINITIES

Presence of species of Murchisonia,
Bellerophon and Palaeozygopleura, Euryzone
and Anomphalus demonstrates Old World Realm
affinities, but these are at generic level only. It is
unclear whether there are links to Chinese and
Russian faunas of this age, as they have been so
sparingly described, and microgastropods of this
age are little-recorded in the literature. There is à
general taxonomic continuance with respect to
Early Devonian (Emsian) faunas described by
Tassell (1982 ), but this is only on a generic level
with the genera represented in both being long-
ranging and widespread. Thus there is à strong
endemism to the fauna, even when compared to
Giveuan gastropods from the adjacent Broken
River Province (Cook & Camilleri, 1997), with
no common species apart from Burdikinia
burdekinensis and Labrocuspis nodosa, This may
be a function of the strongly embayed palaco-
geographic setting of the Burdekin Subprovince,

GASTROPODS FROM THE BURDEKIN FORMATION 49

in comparison to the open marine conditions of

the Broken River Province during the Givetian.

LITERATURE CITED

BATTEN, R.L. 1966, The Lower Carboniferous gåstro-
pod fauna from the Hotwells Limestone of Comp-
ton Martin, Somerset. (Palaeontographical
Society, London).

BLODGETT, R.B. 1988. Wiscensonella, à new
raphiostomatinid gastropod genus from the Mid-
dle Devonian of Wisconson. Journal of Paleontol-
ogy 6203); 442-444,

BLODGETT, R.B. & JOHNSON, J.G. 1992, Early
Middle Devonian (Bifelian) gastropods of Central
Nevada. Paliiontographiea Abt A. 222 (4-6): 85-
134,

BLODGETT, RR. ROHR, D.M. & BOUCOT, AJ,
1990. Early and Middle Devonian gastropod hio-
geography. Pp. 277-284. In MeKerrow. WS &
Scotese, C.R. (eds) Palaeozoic palacogeography
and biogeography. Geological Society Memoir
12. (The Geological Society: London).

COOK, A.G. 1993a. Two bivalves From the Middle
Devoman Burdekin Formåtion. Memoirs of the
Queensland Museum 33(1): 49-53,

1993b. Flercherviewia seprata: A new high-spired
seplate gastropod from (he Devonian of North
doe ge Journal of Paleontology 67(5): 816-

1994, Stomaloporoid palaeoecology and and faunas
of the Burdekin Subprovince. Ph.D. Thesis (un-
publ.) James Cook University of North Queens-
land.

1995. Sedimentology and depositional environ-
ments of the Middle Devonian Big Bend Arkose
and Burdekin Formation, Fanning River Group,
Burdekin Subprovince, north Queensland, Aus-
tralia, Memoirs of the Queensland Museum
38(1); 53-91.

COOK, A.G. & CAMILLERI, N. 1997. Devonian gastro-
pods from the Broken River Province, north
Queensland. Memoirs of the Queensland Muscum
42(1):56-79, this issue.

ETHERIDGE, R. 1917. Descriptions of some Queens-
land Palaeozoic and Mesozoic fossils. 3. A re-
markable univalve from the Devonian limestone
of the Burdekin River, Queensland. Geological
Survey of Queensland Publication 260: 13-16.

HEIDECKER, E. 1959. Middle Devonian molluscs
from the Burdekin Formation of North Queens-
land, University of Queensland Papers, Depart-
ment of Geology 5(2): 1-11,

HENDERSON, R.A. 1980. Structural outline and sum-
mary geological history for northeastem Aus-
tralia, Pp. 1-26, In Henderson, R. A. &
Stephenson, P. J, (eds) The geology and geophys-
ics of northeastern Australia. (Geological Society
of Australia, Queensland Division, Brisbane),

KNIGHT, J.B, 1934, The gastropods of the St Louis,
Missouri. Pennsylvanian outlier, VIN: The

Turretellidae. Journal of Paleontology 8(4): 333-
447,

1941. Paleozoic gastropod genotypes. Geological
Society of America Special Papers 32.

KNIGHT, J.B., COX, LR. KEEN, A.M., BATTEN.
R.L., YOCHELSON, E.L. & ROBERTSON, R.
1960, Kvien Descriptions. Pp. 1169-1331. In
Moore, R.C. ted.) Treatise on invertebrate palcon-
tology, Part I. Mollusca I. (Geological Society of
America & University of Kansas Press: Lawrence,
Kansas),

LINSLEY, R.M. 1968. Gastropods from the Middle
Devonian Anderdon Limestone. Bulletins of
American Paleontology 54 (244): 331-465.

MANSLIY, H, 1912. Etude geologique de Yun-nan Ori-
ental, Part Hl, Paléontologie. Mémoires du Service
Géologique de l'Indo-Chine. 1(2).

MEEK, F.B, & WORTHEN, A.H. 1861. Descriptions
of new Carboniferous fossils from Illinois and
other western states. Proceedings of the Academy
of Sciences of Philadelphia, 1860: 447-472,

NELSON, L.A, 1947, Two new genera of Palaeozoic
Gastropoda. Journal of Paleontology 21: 460-463.

ROLLINS, H.B., ELDREDGE. N. & SPILLER, J.
1971. Gastropoda and Monoplacophora of the
Solsville Member (Middle Devonian, Marcellus
Formation) in the Chenango Valley, New York
State Bulletin of the American Museum af Natu
ral History 144: 129-170,

SHERZER, WU & GRABAL, W. 1908, New Upper
Siluric fauna from saurhern Michigan. Bulletin of
the Geological Sociaty of America 19: 540- 553,

STAUFFER, C.R. 1909. The Middle Devonian of Ohio,
Geological Survey of Ohio Bulletin 10.

TALENT, J.T. & MAWSON. R. 1994. Conodonts in
relation to age and environmental framework of
the Burdekin basin (Mid-Devonian), north-
eastern. Australia, Courier Forsehungsistitut
Senckenberg. 168. 61-81.

TASSELL, C.B. 1982. Gastropods from (he Early
Devonian "Receptaculites" Limestone, Taemas,
New South Wales. Records of the Queen Victoria
Museum Launceston 77.

WHIDBORNE, GF. 1889-92. A monograph of the
Devonian fauna of the South of England, Volume
1, the fauna of the limestone of Lummuton,
Wolborough, Chircombe Bridge and Chudleigh.
(Palaeontographical Society, London).

WYATT. DH & JELL, 1.5. 1967. Devonian ol the
Townsville Hinterland, Queensland, Australia,
Pp. 99.105. In Oswald, D.H. (ed.) International
symposium on lhe Devonian System, Calgary,
1967, (Alberta Society of Petroleum Geologists:
Alberta).

ZHEN. Y.Y. 199]. Devonian rugose coral faunas and
biostratigraphy of the Fanning River Group, north
Queensland. Ph.D. Thesis (Unpubl.). University
of Queensland.

1994, Givetian rugose corals from the northern mar-
gin of the Burdekim Basin, north Queensland.
Alcheringa 18: 301-343.

50 MEMOIRS OF THE QUEENSLAND MUSEUM

PROBABLE PREDATION SCAR ON A DEVONIAN
BRACHIOPOD. Memoirs of the Queensland Museum 42(1):
50. 1997:- A silicified pedicle valve of the brachiopod
Warrenella sp. was recovered from limestone samples of the
Burdekin Formation by acetic acid dissolution. The specimen
derived from Queensland Museum Locality QML 1094, near
Little Rocks, Fletcherview Station, and is registered in the
Queensland Museum Palaeontology collection QMF35128.
Stratigraphy and data concerning the Givetian age of the
Burdekin Formation are outlined by Cook (1995),

The valve (Fig. 1) bears a small depression fracture near
the right anterior margin, 9mm wide and approximately 2mm
deep. This fracture bears a strong resemblance to scars docu-
mented in Carboniferous brachiopods by Alexander (1981)
and Gutteridge (1989). Alexander (1981) attributes such
crushed valves to seized but uningested individuals being
grazed upon by sharks. Gutteridge (1989) similarly attributed
such scars to predation by sharks,

Given the minimal amounts of vertebrate fossil material
recovered from the entire Burdekin Formation (only four
incomplete and unidentifiable bone fragments), it is unclear
what likely predators could have been responsible for this
scar.

Literature Cited

ALEXANDER, R.R. 1981. Predation scars preserved in
Chesterian brachiopods: probable culprits and evolu-
tionary consequences for the articulates. Journal of
Paleontology 55(1): 192-203.

COOK, A.G. 1995. Sedimentology and depositional environ-
ments of the Middle Devonian Big Bend Arkose and
Burdekin Formation, Fanning River Group, Burdekin

FIG. 1. Warrenella sp., QMF35128, arrow indicates probable
predation scar, x 1,3. Specimen blackened with colloidal
graphite and whitened with ammonium chloride for photog-
raphy.

Subprovince, north Queensland, Australia. Memoirs of
the Queensland Museum 38(1): 53-91.
GUTTERIDGE, P. 1989. Shark predaüon on Dinantian
brachiopods from the Derbyshire Dome. Mercian Ge-
ologist 11 (4): 237-244.
A.G. Cook, Queensland Museum, PO Box 3300, South Bris-
bane, Queensland 4101; 23 December 1996.

REVIEW OF FOSSIL GASTROPODS BURDIKINIA KNIGHT 1937 AND AMPHELISSA
ETHERIDGE 1921

ALEX G. COOK

Cook, A.G, 1997 06 30: A review of the gastropod Burdikinia Knight 1937 and Amphelissa
Etheridge, 1921. Memoirs of the Queensland Museum 4211): 51-54, Brisbane. ISSN 0079-

8835.

Burdikinia is here placed within the Helicolomidae. Burdikinia aviønoides (Etheridge), from
the Middle Devonian of New South Wales is formally included in the genus. Austerum
Heidecker 15 å junior synonym of Amphelissa Etheridge. [ ] Gastropods, Devonian, Helicotomidae.

Alex G. Cook. Queensland Museum. P.O. Box 3300 South Brisbane Queensland 4101,

Australia; | May 1997.

The Middle Devonian gastropod Burdikinia
Knight (1937), with type species B. burdekinensis
(Etheridge, 1917) is a distinctive, robust, nodose
taxon regarded as endemic to northeastern Aus-
tralia, Erected by Knight (1937) to correct the
original assignment of the type species to Poly-
amma, the generic name was a corruption or
misspelling of *Burdekin' either River or Lime-
sione. Despite several studies and reports of
Burdikinia burdekinensis including taxonomy
(Etheridge, 1917; Knight, 1937; Knight, 1941;
Heidecker, 1959; Knight et al., 1960) and ecology
(Cook, 1993) there has been no conclusion
reached on its higher taxonomic placement.
partly due to the almost ubiquitous poor preser-
vation of Burdikinia specimens, Thus the genus
has been relegated to the genera inquirenda
(Knight et al., 1960). Discovery of a moderately
well-preserved specimen of B. burdekinensis fi-
nally allows this enigmatic genus to be assigned
to a family.

The posthumous publication of Etheridge
(1921) of 2 gastropods from the Timor Lime-
stone, Upper Hunter Valley, NSW has been ig-
nored by most studies of Devonian gastropods.
Etheridge (1921) erected Polyamma axionoides
on the basis of 2 worn specimens forwarded to
him by the Geological Survey of New South
Wales. The only subsequent reference to this
taxon is that of Pedder, Jackson & Ellenor (1970)
who listed the taxon, assigning the species to
'Burdekinia' (sic).

Etheridge (1921) also erected Amphelissa
isisensis, Which Pedder, Jackson & Ellenor
(1970) listed as Euomphalus isisensis. Austerum
Heidecker bears striking similarity lo Amphelissa
and is synonymised.

SYSTEMATIC PALAEONTOLOGY

Superfamily EUOMPHALIDAE de Koninck, 1881
Family HELICOTOMIDAE Wenz, 1938

Burdikinia Knight 1937

TYPE SPECIES. Polyamma burdekinensis Etheridge
1917 hy original designation, from the Middle Devon-
ian (Givetian) Fanning River Group, probably lower
Burdekin Formation or Big Bend Arkose.

SPECIES INCLUDED. Burdikinia burdekinesis
(Etheridge, 1917), Middle Devonian (Givetian),
north Queensland; Burdikinia axionoides
(Etheridge. 1921) from the Middle Devonian
(Givetian) Timor Limestone, New South Wales.

DISTRIBUTION. Burdekin Subprovince. N
Queensland; Broken River Province, N Queens-
land; Timor Area, Upper Hunter, New South Wales.

REMARKS, Poor preservation has been the sin-
gle contributing factor to the lack of understand-
ing of the systematic position of Burdikinia.
Burdikinia burdekinensis occupied high-energy,
nearshore environmments (Cook, 1993). Conse-
quent abrasion of shell material obliterated all but
gross shell detail. Subsequent recrystallisation of
much of the available material also obscured
taxonomic features, such as the position of the
sinus (if any) or selenizone (if any). In addition
specimens are usually not conducive to prepara-
tion due to the lack of mechanical (and chemical)
difference between surrounding matrix and the
recrystallised shell.

Burdikinia axionoides (Etheridge, 1921) differs
from the type species, being more squat, and
lacking the prominent cords on the lower whorl
face. The lectotype of B. axionoides, (Fig. 1D,E)
here designated as MMF16014, shows the sinus
10 be on the flattened upper whorl face, between

52 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A-C, Burdikinia burdekinesis (Etheridge), QMF35452. A, apical view, arrow shows sinus position, X1;
B, side view, X1; C, basal view. D-E. Burdikinia axionoides (Etheridge), Lectotype MMF16014 X1. D, apical

view, arrow shows sinus; E, side view.

the suture and the nodose cord. The only other
specimen of the original suite examined by
Etheridge (1921), MMF16015 is poorly preserved
and lacks any particular diagnostic features.

Burdikinia is here placed in the Helicotomidae
because of the dominance of spiral ornament, and
particularly the prominent nodose carina border-
ing the flattened upper whorl surface, and the
presence of the shallow sinus on the upper whorl
surface between the suture and the bordering
carina. The anomphalous base is unlike other
Helicotomidae, but reflects great shell thickening
as a probable ecological adaptation.

Burdikinia burdekinensis (Etheridge, 1917)
(Fig. 1A-C)

MATERIAL EXAMINED. HOLOTYPE: GSQF926.
NEW MATERIAL: QMF35452 from Big Bend, near
Charters Towers, NQ.

ADDITIONAL DESCRIPTION. See also
Etheridge (1917) and Heidecker (1959). Growth
lines are fine and numerous; prosocline from the
uppermost and peripheral spiral cord, strongly
recurved about the base towards the axis; these
lines are uninterrupted across the lower whorl
surface. The weak sinus is located on the upper
whorl face regarded here as a subsutural ramp.
The nodes on the spiral cords are solid (as noted
by Heidecker) in deference to Knight et al.
(1960).

REMARKS. The new material places the sinus

accurately and refutes any presence of a
selenizone.

Amphelissa Etheridge 1921

Amphelissa Etheridge 1921: 2; Knight 1941: 34.
Austerum Heidecker 1959: 6.

REVIEW OF FOSSIL GASTROPODS 53

FIG. 2. Amphelissa isisensis Etheridge. A-C, Holotype MMF16011, X1, apical, basal and apertural views
repsectively. D-E, MMF16024, x1. D, apical view; E, side view, note fine growth lines.

TYPE SPECIES. By monotypy. Amphelissa isisensis
Etheridge 1921 from the Middle Devonian Timor
Limestone, NSW.

OTHER SPECIES INCLUDED. Amphelissa
carinatum (Heidecker) 1959, from the Middle
Devonian Burdekin Formation, north Queens-
land.

REMARKS. Amphelissa presented problems to
Knight (1941) who was unable to access type
materials, due to their misplacement. Conse-
quently, and due to the poor quality of the original
illustrations, Knight (1941) regarded the genus as
probably widely phaneromphalous. On the holo-
type of A. isisensis (Fig 2A-C) the base is flat-
tened, and the umbilicus is very shallow, wide and

partially plugged by shell, thus hemiomphalus.
This is an identical condition to the base of the
holotype of Austerum carinatum Heidecker. In A.
isisensis there are well-preserved, fine, numerous
growth lines (Fig. 2D,E), which are not preserved
on specimens of the north Queensland species.
Heidecker (1959) may have been unaware of
the taxon described by Etheridge (1921) from the
Timor Limestone which has gone largely ignored.
Amphelissa was suppressed by Knight et al.
(1960), who synonymised it with Srraparollus
(Euomphalus). Amphelissa Etheridge cannot be
accommodated in Straparollus (Euomphalus)
given the hemiomphalus condition. Amphelissa
is thus resurrected as a valid genus. Amphelissa
carinatum (Heidecker) differs from the type in

54 MEMOIRS OF THE QUEENSLAND MUSEUM

being higher-spired. The holotype of Amphelissa
isisensis, designated by Knight (1941), is regis-
tered in the Geological Survey of New South
Wales Collection as MMF16011. The paratypes
designated by Knight (1941) are registered under
MMF16012.

ACKNOWLEDGEMENTS

I am grateful to Dr Ian Percival for loan of
material from the Geological Survey of New
South Wales collections, to Robert Blodgett for
suggestions to the manuscript and to Rae Sheri-
dan for allowing the specimen to leave the edu-
cational collections under her control.

LITERATURE CITED

COOK, A.G. 1993 Fletcherviewia septata, a new, high-
spired gastropod from the Devonian of north
SE Journal of Paleontology 67 (5): 816-

1.

ETHERIDGE, R. 1917, Descriptions of some Queens-
land Palaeozoic and Mesozoic fossils. 3. A re-
markable univalve from the Devonian limestone

of the Burdekin River, Queensland. Geological
Survey of Queensland Publication 260: 13-16.
1921. Palaeontologia Novae Cambriae
Meridionalis- Occasional descriptions of New
South Wales fossils, No. 8. Records of the Geo-
logical Survey of New South Wales 10 (1): 1-11.

HEIDECKER, E. 1959. Middle Devonian molluscs
from the Burdekin Formation of North Queens-
land. University of Queensland Papers, Depart-
ment of Geology 5(2): 1-11.

KNIGHT, J.B. 1937. Genotype designations and new
names for invalid homonyms among Paleozoic
gastropod genera. Journal of Paleontology
11:709-714.

1941. Paleozoic gastropod genotypes. Geological
Society of America Special Papers 32.

KNIGHT, J.B., COX, L.R., KEEN, A.M., BATTEN,
R.L., YOCHELSON, E.L. & ROBERTSON, R.
1960. Systematic Descriptions. Pp. 1169-1331. In
Moore, R.C. (ed.) Treatise on invertebrate paleon-
tology, Part I. Mollusca 1. (Geological Society of
America and University of Kansas Press: Law-
rence).

PEDDER, A.E.H., JACKSON, J.H., & ELLENOR,
D.W. 1970. An interim account of the Middle
Devonian Timor Limestone of northeastem New
South Wales. Proceedings of the Linnean Society
of New South Wales 94 (3): 242-272.

MIDDLE DEVONIAN GASTROPODS FROM THE BROKEN RIVER PROVINCE,

NORTH QUEENSLAND
ALEX G, CODK AND NATALIE CAMILLERI

Cook, A.G. & Camilleri, N, 1997 06 30: Middle Devonian gastropods from the Broken River
Province, north Queensland. Memoirs of the Queensland Museum. 4201): 55-79. Brisbane.
ISSN 0078-8835.

Twenty seven taxa of gastropods are described from the Eifelian and Givetian sequences of
the Broken River Province, north Queensland with four new genera and ten new species.
New taxa are Denayella lomandraensis sp. nov., Gyronema simpsoni sp. nov.. Frillbeastia
queenslandicus gen, el sp. nov. Brokenriveria pharlapensis gen. et sp.nov., Gemininodosa
langi gen. et sp. nov., Murchisonia (Murchisonia) wandovalensis sp. nov., M. (M.) lawlessi
sp. nov, Palaeozygopleura dodgeyi sp. nov., Australoxa tasselli gen. et sp. nov. and
Leptogyma queenslandicus sp. nov. Three gastropod communities are recognised in the
Givetian rocks of the province, the Murchisonia community, inhabiting a biostromal envi-
ronment, the Labrocuspis community inhabiting high-energy coarse siliciclastie environ-
mens and the Brokenriveria community inhabiting an open, muddy, carbonate shelf, [ ]
Gasrropods, Givetian, Eifelian, Broken River Province, Queensland.

Alex G Cook & Natalie Camilleri. Queensland Museum, PO Box 3300 South Brisbune,

410], Australia; I April 1997.

The Broken River Province. located approxi-
mately 200km W of Townsville, north Queens-
land consists. of two subprovinces; Graveyard
Creek Subprovince and Camel Creek Sub-
province (Withnall & Lang, 1993). The Grave-
yard Creck Subprovince contains inter alia, a
thick, widely-outcropping sequence of Silurian to
Middle Devonian, dominantly shallow marine
units.

Middle Devonian sequences within the Grave-
yard Creek Subprovince contain diverse fossil
assemblages which have been the subject of in-
tense taxonomic and biostratigraphic study (Jell
etal, 1993), Detailed studies of conodonts (Maw-
son, 1987; Mawson & Talent,1989; Mawson ct
al., 1988), fish remains (Turner, 1982, 1995; de
Pomeroy, 1994) and corals (Wyatt & Jell, 1967)
have provided a substantial biostratigraphic
database for tàxonomic studies within the prov-
ince. We follow the summary biostratigraphic
scheme presented by Jell et al. (1993).

Until now gastropods have remained unstudied
within the Broken River Province. Previous work
on Devonian gastropods from north Queensland
åre Etheridge (1917), Heidecker (1959), and
Cook (1993, 1995). Gastropod faunas in south-
castern Australia have received substantially
more attention, more recently through the works
of Tassell (1976, 1977, 1978, 1980, 1982).

This paper is concemed with gastropods col-
lected from Eifelian and Givetian units of the
Broken River Group, namely the Burges Forma-
pon, Dosey Limestone and. Papilio Mudstone,

Material collected over a number of years by staff
at The University of Queensland, Macquarie Uni-
versity and the Queensland Museum contained å
small number of gastropods scattered over å large
number of localities in the Broken River Prov-
ince. Subsequent larger collections made by the
authors in 1994 and 1995 have revealed å more
diverse assemblage. All material is deposited in
ihe Queensland Museum (prefix QMF), and lo

calities are presented in the appendix (QML).

List of Localities. QML541; ‘Calceola’ stop 6.
2nd creek upstream from Broken River Gorge.
Broken River Province. Burges Formation. Eitel-
ian. Collected P.A. Jell. QML1016: Hill above
crossing, S side of river near 'PharLap' prospect,
Broken River, near old GSQ camp site. 19*28' S,
144743" E. Papilio Mudstone, Givetian. Collected
A. Cook, N, Camilleri. QML 1018: Low rise, Ikm
S of Storm Dam, 200m W of road at [9732 92'5,
144°40.51'E. Papilio Mudstone, Givetian. Col-
lected A, Cook, N. Camilleri. QML1019: Ridge
of silicified (silerete)- replaced Dosey Limestone,
100m north of road 500m E of Storm Dam 190.
QML1083. Nuggery Gully, E of Gorge-PharLap
Rd, basal Papilio Mudstone, just above and
adjacent to top of N most Dosey Limestone,
19°27.79'S, 144°44.86'B. Papilio Mudsiume,
Giverian. Collected A, Cook, N. Camilleri, P
Lawless, D. Case, S. Dodgey-Hocknull, P.
Dodgey-Tierney, R, Lootsma, 1995: QML 1090.
Above 'PharLàap' crossing on Broken River, N

side of River on washed oui track approx. 40m

NE of QML1016, 19998,718, 144744.05' E,

56 MEMOIRS OF THE QUEENSLAND MUSEUM

Papilio Mudstone, Givetian. Collected QM Party
1995. QML1092: 500m NW of type section creek
for Dosey & Lomandra Limestones. 150m above
Dosey Limestone. 19°29.98’S, 144?43.87'E.
Papilio Mudstone, Givetian. Collected QM Party
July 1995. A. Cook, P. Lawless, C. McHenry,
September 1995. SD21: E side of Dosey Syncline
in tributary of Dosey Creek. (see Mawson &
Talent, 1989; 221) Papilio Mudstone, Givetian.
SD43e: Storm Dam area, Papilio Mudstone,
Givetian. SD108: Approx 1.5km NE of Spanner
Hill, Papilio Mudstone or Lower Mytton Forma-
tion. (see Mawson & Talent 1989: 211) Givetian.

GASTROPOD ASSEMBLAGES

Three Givetian gastropod communities are
recognised; | Labrocuspis community,
Brokenriveria community and Murchisonia com-
munity (Table 1), corresponding to differing sed-
imentologic regimes. The Labrocuspis
community is found in conglomerates and coarse
sandstones interpreted as high energy facies at the
basal Papilio Mudstone near ‘Nuggetty Gully’
area and from similar sandstones in the upper-
most Papilio Mudstone or lowermost Mytton
Formation in the ‘Spanner Hill’ area. This occur-
rence is strikingly similar to the Burdikinia-
Labrocuspis-Austerum faunule noted by Cook
(1993), which occurs in high energy nearshore
facies ofthe Big Bend Arkose (Givetian, Burdein
Subprovince).

Brokenriveria community occurs within car-
bonate mudstone facies of the Papilio Formation,
interpreted as shallow, low energy, open marine,
impure carbonate shelf. Murchisonia community
is derived from within the uppermost Dosey
Limestone and is associated with corals,
stromatoporoids, sponges from a biostromal en-
vironment.

Eifelian gastropods are insufficiently repre-
sented in the collected fauna to comment on their
commmunity arrangement.

FAUNAL AFFINITIES

Comment has already been made on the simi-
larity of Labrocuspis community in the Burdekin
Subprovince and the Broken River Province. In
both are also found the bivalve Tanaodon
louderbacki, which is also known from the Givet-
ian of Guangxi (Pojeta et al., 1985). The presence
of Murchisonia species, Soleniscus, palaeo-
zygopleurids and Plaryceras suggests affinity
with Old World Realm faunas, but sufficiently
removed to develop distinct endemism at the

TABLE 1. Constituent taxa of Givetian gastropod com-
munities. r = rare, c = common, a = abundant, s =
single occurrence.

———
===>

Species

Brokenriveria

Murchisonia
community
Labrocuspis
community
community

wn

Bellerophon (Bellerophon) sp. A.
Bellerophon (Bellerophon) sp. B. 5

Straparollus (Straparollus) sp. 8

Straparollus (Euomphalus) sp. A r
Straparollus (Euomphalus) sp. B E
Labrocuspis nodosa Heidecker | c
Omphaoltrochid indet. E

Denayella lomandraensis sp. nov. r

Frillbeastia queenslandicus
| gen. et. sp. nov.

Brokenriveria pharlapensis sp. nov. a

Gemininodosa langi gen. et sp. nov. a

Platyceras(Platyceras) sp. 5

| platyceratoid indet. 8
Burdikinia burdekinensis (Etheridge) r

Murchisonia ( Murchisonia)
wandovalensis sp. nov.

Murchisonia (Murchisonia) lawlessi
sp. nov.

Muchisonia (M.) sp. cf. M. (M.)
fermioni Tassell

Murchisonia (M.) sp. s

murchisoniid indet. r

Stylonema? sp. s

Australoxa tasselli gen, et sp. nov. r

Palaeozygopleura dodgeyi sp. nov. r

Soleniscus sp. cf. S. subcostata
Schlotheim

`
Leptogyma queenslandicus sp. nov. c
| Mitchellia striatula de Koninck s

species level. There are, however closely related
species of Brokenriveria gen. nov. and possibly
Frillbeastia gen. nov. known from Germany as
argued below. There are other generic level affin-
ities with Eifelian faunas in Nevada (Blodgett,
1992) and Alaska (Blodgett & Johnston, 1992)
and Givetian faunas in Yunnan (Mansuy, 1912),
Guangxi (Wei & Pan, 1988). Taxonomic hang-
overs from southeastern Australian faunas of
Emsian age are evidenced by the co-occurence of
Mitchellia, Leptogyma, and Murchisonia (M. ) sp.
cf. M. (M.) fermioni Tassell. A dearth of taxo-
nomic work on southeast Asian and Russian gas-
tropods of Middle Devonian age makes
conclusive assessment of the taxonomic affinities
impossible.

GASTROPODS FROM THE BROKEN RIVER PROVINCE 57

SYSTEMATIC PALAEONTOLOGY

Class GASTROPODA Cuvier, 1797
Order ARCHAEOGASTROPODA Thiele, 1925
Superfamily BELLEROPHONTOIDEA
M'Coy, 1851
Family BELLEROPHONTIDAE M'Coy, 1851
Bellerophon Montfort, 1808
Belleropohon (Bellerophon) Montfort, 1808

Bellerophon (Bellerophon) sp. A.
(Fig. LA, B)

MATERIAL EXAMINED. QMF32642 from SD21,
QMF32644 trom QML1018.

DESCRIPTION. Medium-sized, isostrophic
shell, up to 20mm wide and 20mm diameter;
doubly phaneromphalous. Whorl profile gently
rounded, with a weak ridge mid-whorl, Smooth,
fine, growth lines, nearly straight; shell rapidly
expanding: lip and aperture unknown.

REMARKS. Poor preservation of this material
does not allow confident specific assignment to
ane of the many species of Bellerophon
(Bellerophon). It is assigned to this subgenus on
the basis of the rounded whorl profile, absence of
spiral ornament and simplicity of growth lines.

Bellerophon (Bellerophon) sp. B
(Fig. IC, D)

MATERIAL EXAMINED. QMF33621 from
QML1092.

DESCRIPTION. Small, isostrophic shell, 10mm
wide, 10mm diameter; rapidly expanding. aper-
ture flared. Selenizone, prominent upon weak
ridge. No growth lines preserved.

REMARKS. The single, poorly preserved speci-
men cannot be assigned to a species, It is more
rapidly expanding ànd smaller than Bellerophon
(Belleraphon) sp. A.

Superfamily EUOMPHALOIDEA
de Koninck, 1881
Family EUOMPHALIDAE de Koninck, 1881
Straparollus de Montfort, 1810
Straparollus (Straparollus) de Montfort, 1810

Straparollus (Straparollus) sp.
(Fig. 1E, F)

MATERIAL EXAMINED. QMF33103 from
QML1019,

DESCRIPTION, Medium-sized, trochifrom
shell, approximately 5.5mm high, 10mm wide;
apical angle approximately 110°, sutures strongly
impressed. Whorl profile strongly rounded with
slight shoulder. Ornament consists of numerous
fine growth lines, slightly prosocline. There is à
very slight flexure in the growth lines on the weak
shoulder suggesting the sinus. Just below mid-
whorl near the aperture, there is an apparent shell
repair.

REMARKS. The specimen is confidently as-
signed to the subgenus on the basis of the whorl
profile, overall shape and ornament. The number
of species of this subgenus is large and confident
assigment 10 species is unwise due to the lack of
material, and the degree of variation present in
known taxa (Linsley & Yochelson, 1973). The
specimen differs from S.($.) ater (Spitz) of
Jhaveri (1969) from the lower Devonian ol the
Carnic Alps, which is much more squat. 5. (S\)
kokeni (Spitz) of Jhaveri (1969) has a much
more prominent shoulder. Of the taxa described
by Linsley & Yochelson (1973), S. (5.) laevis
(Archiac E Vemeuil), from the Middle Devonian
of Germany, is more flattened, but their S. (S.)
?laevis is more resemblant of the Broken
River specimen. S.(S.) cyclostomus (Hall) of
Linsley & Yochelson (1973) from the Middle
Devonian of North America has a more promi-
nent sinus.

Straparollus (Euomphalus) Sowerby, 1814

Straparollus (Euomphalus) sp. A
(Fig. 1G)

MATERIAL EXAMINED. QMF33104, QMF33352
from L1019.

DESCRIPTION. Small, planispiral shell up 10
12mm in diameter, 4mm maximum height. Both
specimens are preserved as basal moulds. Whorl
profile subrounded to subquadrate, with a weak
angulation between the midwhorl and basal sur-
faces obvious in the last whorl. No obvious
growth lines preserved, but there is a faint hint of
fine collabral lines on QMF33104. Protoconch
unknown.

REMARKS. This material cannot be identifed
further due 10 inadequate preservation.

58 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A,B, Bellerophon (Bellerophon) sp. A., QMF32642 x 2.2. A, side view; B, apertural view. C,D,
Bellerophon (Bellerophon) sp. B, QMF33621 x 2.7. C, side view; D, view of selenizone. E,F, Straparollus
(Straparollus) sp., latex moulds from QMF34267 x 3.1. E, apertural view; F, apical view. G, Straparollus
(Euomphalus) sp. A, QMF33352 x 3.1, basal view. H, Straparollus (Euomphalus) sp. B, QMF34752 x 2.2. All
specimens (and those on subsequent figures) whitened with ammonium chloride for photography.

GASTROPODS FROM THE BROKEN RIVER PROVINCE 59

Straparollus (Euomphalus) sp. B.
(Fig. 1H)

MATERIAL EXAMINED. QMF34752 from
QMLI1018.

DESCRIPTION. Large, planispiral, 42mm in di-
ameter; 4 whorls preserved in cross section and
external mould, the first 2 preserved whorls are
septate; septa gently concave; collabral growth
lines preserved on the final whorl are fine and
numerous.

REMARKS. The planispiral form, simplicity of
growth lines and the septation identifies the sub-
genus, but a species cannot be assigned.

Family OMPHALOTROCHIDAE
Knight, 1945

Labrocuspis Heidecker,1959
Labrocuspis Heidecker, 1959: 6; Kase,1989:149.

TYPE SPECIES. Labrocuspis nodosa Heidecker,
1959, by original designation, from the Middle Devon-
ian (?late Eifelian- Givetian), Big Bend Arkose, north
Queensland.

DIAGNOSIS. Large, anomphalous trochiform
gastropod; suture deep and impressed, whorl pro-
file rises from the suture to a keel and descends
to a variably developed peripheral buttress.

REMARKS. Labrocuspis is presently restricted
to two taxa; the type and L. kobayashii (Kase &
Nishida, 1988), from the Eifelian Nakazato For-
mation, north east Japan. Kase (1989) assigned
the genus to Omphalotrochidae. Heidecker
(1959) remarked on the differential development
of the peipheral buttress, it only being fully ex-
pressed in larger forms.

Occurrences of the genus recorded by Heidec-
ker (1959), Kase (1989) and Cook (1993) high-
light association of this genus with neashore to
shoreline, often high energy siliciclastic deposi-
tion. Cook (1993) suggested strong ecological
control for the genus within the Big Bend Arkose
and Burdekin Formation. Material here described
is from coarse-grained siliciclastic units, includ-
ing conglomerates and very coarse to granular
sandstones which display cross and planar lami-
nation. Thus a high-energy shallow water envi-
ronment is suggested.

Labrocuspis nodosa Heidecker, 1959.
(Fig. 2A-C)

Labrocuspis nodosa Heidecker, 1959: 6, Pl. 1 fig. 2a-d,
Pl. 3, fig. 2a-b.

MATERIAL EXAMINED. QMF16547, QMF33316
locality imprecise, Broken River Province, collected
M. Wade. QMF33315 from SD108, Upper Papilio
Formation or Mytton Formation, collected J. Jell.
QMF33582 from QML1089; QMF33580,
QMF33581, QMF33583 from QML1083. There is
some doubt as to the exact origin of specimens
QMF16547, QMF33315 and QMF33316. All were
collected at the same time, with J. Jell (pers. comm.)
recording the origin as Locality SD108, within the
Papilio Formation. However the coarser grained lithol-
ogy would suggest the nearby outcrops of Mytton
Formation.

DISTRIBUTION. Middle Devonian (?late Eifel-
ian- Givetian), north Queensland. The species is
apparently endemic to the region. Very poorly
preserved material from the Laroona Formation,
Burdekin Subprovince may belong to this taxon,
which would extend the age range to the Ems-
ian.

DESCRIPTION. Moderately large, thick-
shelled, dextrally-coiled, trochiform gastropod
up to 63mm high and 36mm wide, with an apical
angle up to 120°; suture deep and impressed;
sutural slope approximately 10°. Whorl profile
rises sharply from the suture to a gently rounded
carina, and slopes to a very gently convex profile.
Upper whorl profile is gently convex, but breaks
midwhorl to produce a gently concave profile
below the midwhorl producing a buttress on the
lowermost surface. The carina is conspicuously
nodose particularly in final whorls. Base flat, with
a wide callus pad. Aperture quadrate. Growth
lines preserved on base, none preserved on whorl
surface.

REMARKS. Labrocuspis nodosa was only pre-
viously recorded in the nearby Burdekin Sub-
province, Givetian, north Queensland
(Heidecker, 1959). The Broken River material is
relatively poorly preserved but is inseparable
from L. nodosa Heidecker from the Big Bend
Arkose and Burdekin Formation, having the no-
dose carina, variable development of the buttress,
and strongly developed callus pad. It differs from
L. kobayashii (Kase & Nishida) in the degree of
sutural impression, the nodose carina, and hence
the upper whorl profile.

60 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG.2. A-C, Labrocuspis nodosa Heidecker, 1959, QMF33583 x 1, apertural, apical and basal views respectively.
D, Omphalotrochid indet., latex mould from QMF33687 x 2, apical view. EF, Burdkininia burdekininnsis
(Etheridge, 1917), QMF33579 x 2. E, apertural view, F, apical view,

GASTROPODS FROM THE BROKEN RIVER PROVINCE GI

Omphalotrochid indet,
(Fig. 2D)

MATERIAL EXAMINED. QMF330687 from
QML1019.

DESCRIPTION. Mould of an upper surface of å
large, shallowly trochiform shell, 25mm in diam-
eter. Apical angle apporximately 140°, Suture
deep, impressed forming a channel in lust whorl.
Growth lines fine, collabral.

REMARKS, The specimen superficially resem-
bles Orecopia murrayi Tassell, 1978 from the
Lower Devonian Bell Point Limestone, Victoria,
but cannot be confidently assigned to a genus as
no basal structures are known. The specimen
lacks the nodose carina of Labrocuspis nodosa
Heidecker. The specimen is left in open nomen-
clature pending more material.

Superfamily PLEUROTOMARIOIDEA
Swainson, 1840
Family RAPHISTOMATIDAE Koken, 1896
Subfamily RAPHISTOMATINAE Koken, 1896

Denayella Blodgett & Johnson, 1992

TYPE SPECIES. Denayella housei Blodgett & John-
son, 1992. from the Eifelian Denay Limestone of Ne-
vada, USA,

Denayella lomandraensis sp. nov.
(Fig. 3 A,B)

MATERIAL EXAMINED. HOLOTYPE: QMF33650
from QML 1092, Papilio Mudstone, Givetian, Broken
River Province. PARATYPES: QMF33651-33672,
QMF34215-34222 from QML 1092, QMF33619 from
QML1090.

DIAGNOSIS. Small member of genus with
flattenedbase and weak inductural deposit.

DESCRIPTION. Small (Table 2) lenticular, an-
omphalus. Whorl profile sharply rounded, upper
surface gently sloping, lower surface steeper. Pe-
riphery on upper 1/3 of whorl profile. sharply
rounded bearing 2 indistinct cords delimiting a
probable selenizone, within which the lunulae are
nol preserved. Lower cord is more obscure than
the upper and on many specimens they are not
preserved due to abrasion, Aperture rhomboid to
subrounded, with a v-shaped sinus on the labrum.
Shell thick especially at columella. There is a
distinct thickening of shell on the columella. Base
convex, but not sharply so. Very weak, fine, col-

TABLE 2. Measurments for Denayella lomandraensis
sp. nov.

Specimen

labral growth lines, otherwise surface smooth and
unornamented.

REMARKS. The material is placed in Denayella
due to its raphistomid-like appearance, indistinct,
but present, seleneizone and weak inductural de-
posit on the columellar lip. It differs from the
type species in having a more flattened base and
a weaker deposit on the columellar lip. It is more
similar to Denayella sp. of Blodgett & Johnson,
1992 with respect to the flatter base. The species
differs from members of Arizonella Stoyanow,
1948 by its lack of à prominent selenizone, and
well developed collabral ornament. It differs
from Buchelia and Raphistoma due to the lack of
cords on the upper whorl face. Umbotropis
mesoni Tassell 1982, from the Receptacutites
Limestone of New South Wales is clearly phan-
eromphalus, and has more prominent spiral
cords on the upper whorl face rather than the
periphery.

ETYMOLOGY. For Lomandra Creek.
Subfamily LIOSPIRINAE Knight. 1956

Frillbeastia gen. nov.

DIAGNOSIS. Small, trochiform with flattened
base; angular periphery with selenizone bounded
below by frilled carina; upper whorl face with 2
strong threads, the lowermost of which is nodose;
strong prosocline ornament; planispiral pro-
toconch.

TYPE SPECIES. Frillbeasria queenslandicus sp. nov.

DISTRIBUTION, Middle Devonian (Givetian),
uppermost Dosey Limestone, Broken River
Province, north Queensland, ?Middle Devonian
(Givetian), Germany (Sandberger & Sandberger,
1850-6).

REMARKS. The genus is similar to Arcsrra
Stoyanow. 1948 from the early Late Devonian in
Arizona, having a distinct frill, but is less lentic-
ular in shape, possesses stronger collabral orna-
ment, strong spiral cords with nodes, and the

62 MEMOIRS OF THE QUEENSLAND MUSEUM

proioconch is planispiral. Superficially similar is
Astralites Whiteaves, 1892 from the Middle
Devonian of Canada, which lacks the prominent
selenizone and spiral ornament, Members of the
Luciellidae Knight, 1956 such as Luciella de
Koninck, 1883 and Epiptychia Perner 1907 have
the frill above the selenizone, rather than below
in Liospirinae, Zalozone Linsley, 1968 possesses
à double frilled selenizone, and Tylozone Linsley,
1968 has a sclenizone bordered by an upper frill.
Frillbeastia is placed in Liospirinae due to closer
similarity to Arastra, but is by far the most trochi-
form member of the subfamily. Linsley (1968)
placed Arasira within the Raphiostomatinae,
with Zalozone. The base of Frillbeastia is wide
and hence js retained in Liospirinae with the
wide-based Arastra.

Within the genus could be included Lirrorina
alata Sandberger & Sandberger, 1850-1856 from
the Middle Devonian (Givetian) of Germany.
The taxon illustrated (Sandberger & Sandberger,
1850-56: pl. 24: fig. 14) shows the characteristic
frilled keel, a similar base, and numerous cords
on the upper surface. It differs from E queenslandi-
cus by having numerous equal strength cords, it
lacks the strong growth lines and the nodose cord
above the selenizone whichis not identifiable on
the illustrations of the German taxon,

ETYMOLOGY. For the informal name given in the
field, reflecting the character of the selenizone.

Frillbeastia queenslandicus sp. nov.
(Fig. 3C-G)

MATERIAL EXAMINED. HOLOTYPE:
QMF33687, from QML1019 uppermost Dosey Lime-
stone Givetian, Broken River Province. PARATYPES:
QMF33695, QMF34265 from QML 1019,

DIAGNOSIS. As for genus.

DESCRIPTION, Small, trochiform, nearly coni-
cal shell, complete specimen is 16mm wide,
| Imm high with a spire angle of approximately
60°. Base flattened, bearing spiral cord, narrowly
phaneromphalus or anomphalous, Whorl face or-
nate and angular with suture just below frill on
midwhorl. Upper whorl face possesses 2 major
threads, | close to the periphery, and another
weaker thread on the mid-upper whorl face,
These increase in strength throughout growth.
The lowermost thread possesses stout rounded
nodes, the upper thread hus vestigal nodes pre-
served on the holotype. The angular shell periph-
ery is decorated by a simple, fine spiral thread
which is the upper boundary for the selenizonc.

Below the selenizone is hordered by a rhythmi-
cally folded carina, thus producing a frill. The
lower whorl profile possesses a single thread,
slightly nodose. Collabral ornament is prosocl-
ine. Protoconch nearly planispiral, 2-3 whorls
with simple comarginal ornament; a vestige of a
selenizone is recognisable on the 3rd whorl,

REMARKS. Despite the small number of speci-
mens the material is signficantly distinct to war-
rant erection of à new genus. One of the
specimens has less distinct nodes and cords on the
upper whorl face, and shows signs of abrasion,

ETYMOLOGY. For the state of Queensland,

Suborder TROCHINA Cox & Knight, 1960
Supertamily PLATYCERATOIDEA Hall, 1859
Family HOLOPEIDAE Wenz, 1938
Subfamily GYRONEMATINAE Knight, 1956

Gyronema Ulrich in Ulrich & Scofield, 1897

TYPE SPECIES. Trochonemu (Gyronema)
pulchellum Ulrich and Scofield, 1897 from the Middle
Ordovician, Minnesota, United States of America.

DIAGNOSIS. See Knight et al, (1960),

Gyronema simpsoni sp. nov.
(Fig. 4C,D)

MATERIAL EXAMINED, HOLOTYPE; QMF32082
from QML541. Burges Formation, Eifelian, Broken
River Province. PARATYPES; QMF32055,
QMF32058 from QML541.

DIAGNOSIS, Very large Gyronema with charac-
teristic thickened spiral cords, flattened upper-
most whorl surface forming a low shoulder, 2
prominent cords on upper whorl surface, the
highest being an angular carina, 6 prominent
cords on final whorl.

DESCRIPTION, Large, turbiniform, narrowly
phaneromphalus gastropod, with prominent thick
spiral ornament and impressed sutures, Shell
height up to 47mm, and width up to 32mm. with
an apical angle of approximately 20° External
whorl profile is dominated by spiral cords, the
uppermost forming à prominent angular carina
high on the whorl profile which is divided into
two surfaces. Uppermost third of the whorl pro-
file is a gently concave shoulder, which slopes
shallowly from thesuture to the carina; mid whorl
is nearly vertical, and the lower whorl profile
gently convex with both marked by thick spiral
cords. Sutures formed at midwhorl, slightly

GASTROPODS FROM THE BROKEN RIVER PROVINCE 63

above the 3rd spiral cord, thus
obscuring all but 2 uppermost
cords on early whorls. Aperture
generally rounded, slightly
vertically extended with abax-
ial angulation above midwhorl.
Growth lines unknown. The 2
paratypes are somewhat
crushed and distorted, but retain
the characteristic spiral cords.

REMARKS. Material has the
typical form of the genus, but
differs from the type species by
the more flattened shoulder, in
addition this taxon is several
times larger than the genotype
figured hy Knight (1941). In
size and form this species is

comparable to G. bellense of

Tassell from the Early Devon-
ian of Victoria but G. bellense
has far more numerous spiral
cords. G. lirata (Hall) of Rol-
lins, Eldredge & Spiller (1971),
from the Middle Devonian
Marcellus Formation, New
York is smaller (17mm high),
with more spiral cords on the
upper whorl surface, however
it does possess a similar shoul-
der to G. simpsoni. Gyronema
scaliforme Zytlenok,1976
from the Devonian of Belorus'
possesses more cords than the
Broken River species. The
species is close to G. or-
mistoni Blodgett, 1992 from
the Eifelian of Alaska, but
lacks the prominent basal cords
of that taxon.

ETYMOLOGY. For Andrew
Simpson,

Brokenriveria gen. nov.
TYPE SPECIES. Brokenriveria
pharlapensis sp. nov

DIAGNOSIS. Small turbini-
form gyronematid with 2 or-
ders of nodose spiral cords.

FIG. 3. A.B, Denayella lamandraensis sp. nov., Holotype QMF33650 x 6.
A, apertural view; B, apical view. C-G, Frillbeastia queenslandicus gen. et
sp. nov. C-E, latex mould of Holotype QMF33687 x 3.6. C, oblique side
view; D, basal view; E, side view. F, latex mould of Paratype QMF34265,
x 3,1, side view, G, latex mould of Paratype QMF33695, x 4.5, oblique
view showing protoconch.

Queensland, Middle Devonian, Rhineland
(Goldfuss, 1844).

DISTRIBUTION. Middle Devonian (Givetian) REMARKS. Lack of a selenezone precludes as-
Papilio Mudstone, Broken River Province, north signment of this material to grossly similar genera

64 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. A,B. Murchisoniid indet. A, latex mould of QMF33100, side view x
1.8. B, latex mould of QMF34259, oblique side view x 2.7. C, D. Gyronema
simpsoni sp. nov. Holotype QMF32082, x 1.3. C, apertural view; D, side
view. E-G, Platyceras (Platyceras) sp. QMF32641, x 2.7. E, apical view;
F, side view; G, basal view.

Bembexia Oehlert, Nodonema
Linsley, Or other
Pleurotomarioidea. The pro-
socline growth lines across the
shell, the strong cords and the
thicker shell suggest that this
material can be accommodated
within the Gyronematidae. The
genus differs from other
gyronematids in the possession
Of two orders of spiral cord,
rather than one on Yunnania
Mansuy, and Gyronema Ul-
rich. Robert Blodgett (pers.
comm) has kindly drawn my
attention to a similarly or-
namented taxon Turbo
caelatus Goldfuss, 1844 from
the Middle Devonian of Ger-
many, which should be in-
cluded in the genus, but differs
in the number of cords on the
upper whorl face.

ETYMOLOGY. For the Broken
River.

Brokenriveria pharlapensis
sp. nov. (Fig. 5)

MATERIAL EXAMINED. HO-
LOTYPE: QMF32234 from
QML1016, Papilio Mudstone,
Givetian, Broken River Province.
PARATYPES: QMF32235-
32301 from QML1016.

DIAGNOSIS. As for genus.

DESCRIPTION. Small to me-
dium-sized, turbiniform gas-
tropod, up to 17mm high and
wide (Table 3) with an apical
angle of approximately 105°.
Whorl profile rounded, but
dominated by two orders of nu-
merous nodose spiral cords.
Suture impressed, whorls em-
brace at the midwhorl. Periph-
ery situated at midwhorl. Six
major threads on the whorl sur-
face, 1 on the upper whorl face,
another at midwhorl, the re-
mainder on the lower whorl
face. Major cords are inter-
spaced with 1 less prominent
nodose spiral thread, with the
exception of the uppermost

GASTROPODS FROM THE BROKEN RIVER PROVINCE 65

FIG. 5. Brokenriveria pharlapensis sp. nov. A, B, Holotype QMF32234, x 3. A, apertural view; B, apical view.
C-E, paratype, QMF32268, x 2.8. C, side view. D, apical view. E, apertural view. F, QMF32273, x 2.8, apertural
view. G, H. Paratype QMF32272, x 2.7. G, side view; H, apertural view. I, paratype QMF32274, apertural view
x 2,8.

major thread which has 2 minor threads between
it and the suture. Growth lines of 2 orders; numer-
ous, fine, prosocline, slightly coarser growth lines
intesect with spiral cords to produce nodes. Ap-

erture rounded, outer lip not preserved, inner lip
thickened, and in some specimens reflected
slightly. Shell relatively thickened for size. Pro-
toconch unknown.

66 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Measurements for Brokenriveria phar-
lapensis sp. nov.

Specimen height (mm) Kueder A weng
angle (°)
QMF32234 14.6 70
QMF32237 112 o |
QMF32239 14.0 70
QMF32272 143 70
QMF33625 12.5 75 |
QMF33629 13.5 65
QMF33628 13.2 75
| QMP366 | 131

REMARKS. This species differs from
Gemininodosa langi sp. nov. by its lack of very
large nodes on the shell periphery. The specimen
of indeterminite platyceratoid described below
has a more rounded whorl profile, greater expan-
sion rate and less prominent spiral ornament.

ETYMOLOGY. For ‘Pharlap’ Crossing of the Broken
River.

Gemininodosa gen. nov.

TYPE SPECIES. Gemininodosa langi sp. nov. from
the Middle Devonian Papilio Mudstone, Broken River
Province, north Queensland.

DIAGNOSIS. Small to medium-sized, turbini-
form, minutely phaneromphalus, with numerous
spiral cords; upper whorl surface bears 2 spiral
rows of large rounded nodes.

REMARKS. The lack of a sinus or selenizone,
dominance of spiral ornament and the turbini-
form shape indicates placement within the
Holopeidae Wenz. Superficially the genus is like
Oriostoma Munier-Chalmas but does not possess
the wide umbilicus. The distinct nodose ornament
renders the genus unlike any other members of
the family but is reminiscent of the node- (and
selenizone-) bearing Pleurotomariitoidea, such as
Nodonema Linsley, 1968 and Glyptomaria
Knight, 1945.

ETYMOLOGY. For the twin nodes adorning the shell.

Gemininodosa langi gen. et. sp. nov.
(Fig. 6A-I)

MATERIAL EXAMINED. HOLOTYPE: QMF3361 1
from QML1092, Papilio Mudstone, Givetian, Broken
River Province. PARATYPES: QMF33608-33610,
QMF33612-33618, from QML1092, QML33638-
33644 from QML1090.

DIAGNOSIS. As for genus.

DESCRIPTION. Medium-sized, turbiniform,
minutely phaneromphalus, up to 16mm wide and
16mm high (Table 4) with an average spire angle
of 75-115°. Sutures impressed, whorls embrace
at midwhorl. Whorl profile overall rounded, flat-
tened adjacent to the suture, rounded at midwhorl
and lower whorl surface. Midwhorl dominated by
2 rows of large nodes, one at the edge of the
flattened upper whorl surface the other near the
periphery. A third less distinct set of nodes occurs
below the midwhorl. Whorl surfaces are or-
namented with many spiral cords, with at least 5
major cords on the lower whorl surface, 4 on the
mid whorl, and 3 on the upper whorl surface.
Growth lines fine, numerous and opisthocline;
continuing across the midwhorl undeflected by
any selenizone. Protoconch unknown.

REMARKS. This species differs from
Brokenriveria pharlapensis and the indeterminite
platyceratoid described below by possessing the
twin row of nodes on the upper surface. The
grossly similar Nodonema granulatum Linsley
from the Middle Devonian Anderdon Lime-
stone, is much smaller (holotype 6mm high),
and possesses a distinct selenizone.
Kitikamispira ukalundensis Cook, 1995, from
the Emsian Ukalunda Beds of Queensland, and
Kitikamispira kanekoi Kase & Nishida, 1988
from the Eifelian, Nakazato Formation, Japan
both have nodes on all the spiral cords, and these
spiral cords are strong and of equal intensity.

ETYMOLOGY. For Simon Lang.

Family PLATYCERATIDAE Hall, 1859
Platyceras Conrad, 1840

Platyceras (Platyceras) Conrad, 1840

TYPE SPECIES. Pileopsis vetusta, from the Lower
Carboniferous of Queens County, Ireland by subse-
quent designation of Tate (1869).

Platyceras (Platyceras) sp.
(Fig. 4E-G)

MATERIAL EXAMINED. QMF32641, from SD43e.

DESCRIPTION. Small, horn-shaped rapidly ex-
panding shell, 5.1mm high, 11.5mm maximum
width; first whorl in contact, second disjunct;
whorl profile ovate, aperture broken, but basal lip
deflected strongly downwards near apertural

GASTROPODS FROM THE BROKEN RIVER PROVINCE 67

margin. Ornamentconsistsoffine,faintcollabral
growth linesanddiffusecoarser growth rugae.

REMARKS. Substantial variation in members of
the subgenus render it unwise to nominate å spe-
cies on the basis of a single specimen. Platyceras
(P. sp. A.of Tassell (1982) from the Earl y Devon-
ian of Taemas has more disjunct whorls whereas
the Broken River specimen has the first whorl in
contact. Platyceras (P.) mansfieldense Tassell,
1977 from the Early Devonian Loyola Limestone
has similar ornament and has the first whorl in
contact, bur is a significantly larger form

PLATYCERATOIDEA gen. e! sp. indet,
(Fig. 6), K)

MATERIAL EXAMINED. QMF34224 from
QML1092,

DESCRIPTION. Turhiniform, medium-sized,
13mm high and 1 3mm wide. average spire angle
approximately 110°. Sutures impressed, with
whorls embracing slightly above midwhorl,
Whorl profile rounded. Ornament consists of nu-
merous spiral cords of equal strength, and fine
weaker, slightly prosocline collabral growth
lines.

REMARKS. The single distinct specimen lacks
the two orders of spiral threads characteristic of
Brokenriveria pharlapensis and does not bear the
twin cords of Gemininodosa langi. The turbini-
form, shape, lack of sinus and simple ornament
places the specimen in the superfamily, and it 18
possibly a platyceratid, but further assignment 18
impossible without å wider selection of material.

Superfamily and Family indet.
Burdikinia Knight, 1937

Burdikinia burdekinensis (Etheridge, 1917)
(Fig. 2E,F)

Polyamma burdekeninesis 1917 Etheridge: 16; PI. 3
figs 1,2.

Burdikinia burdekinesis (Etheridge) Knight 1937: 711
(Etheridge) Knight 1941: 63; PL 57, figs 3a-h ;
Heidecker, 1959: 5, PI 2, fig. 2, PI, 3, fig.
3a,b: Knight et al. 1960: 1309; fig. 205, 4a,b.

MATERIAL EXAMINED. QMF33578, QMF33579
from OML1083.

DESCRIPTION. Large, low-trochiform shell up
to 35mm high, 55mm wide; apical angle of 130°.
Suture channelled with keel on abaxial margin,
whorls embrace on upper whorl face. Upper

TABLE4. Measurements tor Gemininodosa langi gen.
et sp. nov,

Specimen | Height imm) | Width(mm) app M
est I i42 11.0
| QOME33608 | 14,6 164 ]

QMF33638

OMP33641

Ce

109

whorl surface sloping and slightly concave, lower
whorl surface rounded, Peripheral angulation al
slightly above midwhorl. Aperture subrectangu-
lar. Base rounded, heavily abraded, with poorly
preserved prosocline growth lines. Relicts of
nodes preserved on periphery and on sutural keel.

REMARKS. The material is heavily abraded,
lacking the hasal ornament characteristic of the
taxon, but it is otherwise indistinguishable from
the holotype, and other material collected from
the Burdekin Subprovince. A review ofthe higher
taxonomy of this distinct gastropod is needed.

Superfamily MURCHISONIOIDEA
Koken, 1896
Family MURCHISONIIDAE Koken, 1896
Murchisonia D^ Archaic & De Verneuil, 1841
Murchisonia (Murchisonia)

Murchisonia (Murchisonia) wandovalensis
sp. nov. (Fig. TA-G)

MATERIAL EXAMINED. HOLOTYPE: QMF33680
from QML.1092, Papilio Mudstone, Givetian, Broken
River Province. PARATYPES: QMF33675-
QMF33679, QMF33681-QMF33685, QMF34129
QMF34184, QMF34195-QMF34202 from L1092.

DESCRIPTION. Small, high-spired, turbiniform
gastropod, up to 11mm high and IImm wide
(Table 5), with an apical angle of c. 55°. Suture
impressed. Whorl profile angular with wide, mid-
whorl, peripherally placed selenizone bordered
by prominent 2 spiral cords. Upper whorl face

steep, slightly concave. Lower whorl face
rounded, bearing a spiral cord 1/3 below ihe
selenizone. Whorls embrace at this lowermost
cord, just below the mid-whorl, Growth lines
fine, numerous, collabral; concave 1n the
selenizone; somewhat sinusoidal between the
lower bordering cord of the selenizone and the
spiral cord on the lower whorl face, Base rounded
aperture subrounded with slit at midwhorl.

68 MEMOIRS OF THE QUEENSLAND MUSEUM

GASTROPODS FROM THE BROKEN RIVER PROVINCE 69

TABLE 5. Measurments for Murchisonia (Murchisonia)

Height (mm)

Width(mm)

QMF33676 å 7.2

QMF33680

QMF33685
QMF34202

DIAGNOSIS. Small, somewhat turbiniform
Murchisonia (Murchisonia), with additional spiral
cord on lower whorl face, just below selenizone.

REMARKS. The material is similar to M.(M.)
fermioni Tassell 1982 from the Early Devonian
Receptaculites Limestone, New South Wales,
however M. (M.) fermioni has a more impressed
suture and lacks the spiral cord on the lower whorl
face. From other species of the subgenus it differs
in the less high-spired form, and generally, the
presence of the lower whorl face cord.

ETYMOLOGY. For Wandovale Station.

Murchisonia (Murchisonia) lawlessi sp. nov.
(Fig. 8 A-I)

MATERIAL EXAMINED. HOLOTYPE: QMF33704
from QML1019, uppermost Dosey Limestone, Givet-
ian, Broken River Province. PARATYPES:
QMF33089, QMF34258, QMF33096, QMF33345,
QMF33347, QMF33393, QMF33700, QMF34263
from QML 1019.

DIAGNOSIS. Medium-sized, high-spired mem-
ber of subgenus, with ridge on upper and lower
whorl faces, more prominent on upper, but not
defining a distinct cord.

DESCRIPTION. Medium-sized, high-spired an-
omphalus gastropod, up to 18.4mm high,
12.0mm wide at base (Table 6) with an apical
angle of approximately 22-30°. Whorl profile
angular with peripheral selenizone bordered by
two cords. Upper whorl face shallowly sloping
with a low, rounded, spiral ridge, just below the
suture. Suture impressed, whorls embrace well
below midwhorl at lower spiral ridge. Selenizone
narrow, located at midwhorl. Lower whorl face
rounded and convex, with weaker ridge, slightly
lower and wider. Base rounded, aperture rounded
with slit at peripheral margin.

TABLE 6. Measurements for Murchisonia (Murchisonia)

approx. apical
angle (°)

lawlessi sp. nov.

Specimen

QMF33700
QMF33345

REMARKS. The species is differentiated from
other Murchisonia (Murchisonia) in the area by
the ridge on the upper and lower whorl faces in
addition to those bordering the selenizone.

Murchisonia (Murchisonia) turris de Koninck
from the Early Devonian Receptaculites Lime-
stone, New South Wales lacks the two additional
ridges and is more high-spired. Murchisonia
(Murchisonia) anderdoniae Linsley 1968, from
the Middle Devonian Anderdon Limestone,
North America, has a similar whorl profile and spire
architecture, but is much smaller and lacks the upper
and lower spiral ridges of M.(M.) lawlessi.

The material is superficially resemblant of Ul-
richospira kanekoi Kase & Nishida from the
Eifelian Nakazato Formation, Kitikami Moun-
tains, Japan, but members of Ulrichospira have
the selenizone high on the whorl, rather than
midwhorl in this taxon.

ETYMOLOGY. For Phil Lawless.

Murchisonia (Murchisonia) sp. cf.
M.(M.) fermioni Tassell, 1982
(Fig. 7H,I)

MATERIAL EXAMINED. QMF33701, QMF34266,
QMF33264 from QML1019.

DESCRIPTION. Small to medium-sized, turbini-
form shell, up to 16.1mm high, 10.6mm wide,
with a pleural angle of 35°. Whorl profile acutely
angular, upper whorl face steep, lower flattened.
Periphery at midwhorl bearing narrow selenizone
bordered by 2 spiral threads. Suture impressed,
whorls embrace slightly below selenizone. Base
rounded. Aperture unknown. Growth lines fine,
numerous, collabral, slightly opisthocline.

REMARKS. This species is separated from
M.(M.) wandovalenesis by the lack of spiral or-

FIG. 6. A-I, Gemininodosa langi gen et. sp. nov. A,B. Holotype QMF33611, x 3.3. A, apical view; B, side view.
C-E, Paratype QMF33642, x 3.7. C, oblique apical view; D, side view; E, apertural view. F,G, Paratype
QMF33638, x 3.4. F, apical view; G, Side view. H,I, Paratype QMF32182 (crushed specimen), x 4. H, apertural
view; I, apical view. J,K, Platyceratoidea gen. et sp. indet. QMF34224, x 3.7. J, Side view; K, apertural view.

70 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7. A-G, Murchisonia (Murchisonia) wandovalensis sp. nov. A, Paratype QMF33676, apertural view x 5.
B.C, Holotype QMF33680, x 4,7. B, side view; C, apertural view, D,E, Paratype QMF33685, x 4.5. D, apertural
view; E, oblique side view. F,G, Paratype QMF34202 x 4.5. F, oblique side view; G, apertural view. HI,
Murchisonia (Murchisonia) sp. cf. M. (M.) fermioni Tassell, 1982. H, latex mould of QMF33701, x 3, side
view. I, latex mould of QMF34266, x 3.5, side view.

nament additional to that bordering the
selenizone and the more angular whorl profile.
Murchisonia (A sp. is a higher-spired shell. The
species is resemblant of M.(M.) fermioni Tassell,
1982 from the Early Devonian Receptaculites
Limestone, New South Wales, but Tassell's taxon
is slightly smaller, for this reason we tentatively
refer the limited material to that taxon.

Murchisonia (Murchisonia) sp.
(Fig. 8J)

MATERIAL EXAMINED. QMF33686 from
QMLI019.

DESCRIPTION. Medium-sized, many-whorled,
high-spired slender shell, 22mm height, 8mm basal
width, with an apical angle 20°. Suture impressed
with fine, weak, spiral ridges on the adjacent whorl
surfaces. Whorl profile angular with peripheral,

GASTROPODS FROM THE BROKEN RIVER PROVINCE 71

midwhorl, concave selenizone bordered by
prominent spiral cords of equal strength. Upper
whorl surface begins flattened, thence is gently
concave to periphery. Lower whorl face weakly
concave. Base unknown. Growth lines unknown.
Inner whorl surface rounded in cross-section,
but aperture unknown.

REMARKS. The specimen closely resembles
Murchisonia (Murchisonia) akidota Linsley
1968, from the Middle Devonian Anderdon
Limestone, North America, but Linsley's taxon is
much smaller, and has a more prominent spiral
ridge on the lower suture, rather than weak lines
above and below. Similarly Murchisonia
(Coelocaulis) procera Oehlert 1888 from the
Early Devonian of Saint-Germain-le-Fouilloux,
France and Murchisonia (Murchisonia) scuplta
(Perner, 1907) from the Late Silurian of Bohemia
are larger and lack the ridges adjacent to the
suture.

murchisoniid indet.
(Fig. 4A,B)

MATERIAL EXAMINED. QMF33097, QMF33100,
QMF34259 from QML1019,

DESCRIPTION. Medium-sized, moderately
high-spired, turbiniform gradate shell up to
28mm high and 20mm wide, with an apical angle
of c. 30°. Upper whorl surface with prominent
sutural ramp sloping very gently to peripheral
rounded keel. Midwhorl surface wide and verti-
cal, with selenizone bordered by 2 weak threads.
Lower whorl face rounded, but poorly known in
the material. Suture slightly impressed, situated
at lower part of midwhorl surface. Base un-
known. Collabral growth lines, fine, numerous,
closely spaced, prosocline on sutural ramp and
above selenizone, opisthocline below selenizone.

TABLE 7. Measurements for Australoxa tasselli gen.
et sp. nov.

approx. apical
angle (°)
18.0* 8.6 20
18.0* 92 22

Specimen Height (mm) | Width(mm)

QMF33586
QMF33587

QMF33588 6.5 19
QMF33589 16.9* 10.0 23
QMF33590 | 210* 11.5 25
QMF33708 | 168* 72 20
QMF33712 | Gm 57 20

REMARKS. Blodgett (pers. comm.) has exam-
ined photographs of this material and suggests
that the Broken River specimens are congeneric
with a new species of murchisoniid from north
America. There is not enough material to confi-
dently erect a genus or species.

Order CAENOGASTROPODA Cox, 1959
Superfamily LOXONEMATOIDEA
Koken, 1889
Family LOXONEMATIDAE Koken, 1889
Stylonema Perner, 1907

Stylonema? sp.
(Fig. 10A)

MATERIAL EXAMINED. QMF33102 from QML1019.

DESCRIPTION. High-spired; many-whorled;
small gastropod, 6.9 mm high; 1.4mm wide; api-
cal angle sharp; approximately 12°; sutures
grooved; sutural slope high; whorl profile
rounded; aperture unknown; growth lines and
sinus unknown.

REMARKS. Poor preservation and lack of mate-
rial prevents assignment to species. The whorl
profile and grooved sutures suggest reference to
Stylonema but definite assignment awaits further
material.

Australoxa gen. nov.

TYPE SPECIES. Australoxa tasselli sp. nov.

DIAGNOSIS. Medium-sized, high-spired; angu-
lar whorl profile; growth ornament of prominent
opsithocline ribs. Angular periphery bears shal-
low but sharp sinus. Lower whorl profile with
spiral cord.

REMARKS. The coarse ribbing suggests affini-
ties to Palaeozygopleuridae, but the shallow sinus
suggests placement within the Loxonematidae.
The angular whorl profile is grossly suggestive of
Donaldiella, but the growth parameters are sig-
nificantly different for that genus as indicated by
the differing growth lines. We place the genus in
the Loxonematidae, but the genus appears inter-
mediate between the two families and is derived
from a time of great change in both.

ETYMOLOGY. Latin, Austral, southern, loxa, imply-
ing loxonematid affinifites.

73 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. A-1, Murchisonia (Murchisonia) lawlessi sp. nov. A, latex mould of holotype QMF33704, x 3.1, side
view. B, latex mould of paratype QMF33092, apertural view x 4. C, latex mould of paratype ES x 3.5.

D, latex mould of paratype QMF34260, side view x 3.6. E, latex mould of paratype QMF34258, x 3,5, side
view. F, latex mould of paratype QMF33700, x 3.5, side view. G, latex mould of paratype OME33096, x 3.6,
apertural view. H, latex mould of paratype QMF33089, apertural view, x 4 last whorl abraded. 1, latex mould
of paratype QMF33707, x 4, side view. J, Murehisonia (Murchisonia) sp. QMF33686, x 3.5.

GASTROPODS FROM THE BROKEN RIVER PROVINCE 73

TABLE 8. Measurments for Palaeozygopleura
dodgeyi sp. nov,

Ree n e

oo
| ommo Lo Le La |

Australoxa tasselli sp. nov.
(Fig. 9E-L)

MATERIAL EXAMINED. HOLOTYPE: QMF-
33586, from QML. 1092, Papilin Mudstone, Givetian,
Broken River Province, PARATYPE: QMF33587-
QMF33603, QMF33708-QMF33711. fram
QMLI092.

DIAGNOSIS. Ås for genus.

DESCRIPTION, Medium-sized, high-spired, an-
omphalous, up to slightly more than 22mm high,
i2mm basal width, with a pleural angle of 19-25?
(Table 7). Up to 7 whorls present in the material,
but in general the apical whorls are missing.
Whorl profile angular, with periphery slightly
below midwhorl. Lower whorl face convex.
Upper whorl face slightly convex to nearly flat
with numerous strong opisthocline ribs which
continue to the lower whorl face, only deflected
at the periphery. Sinus expressed as a shallow, but
sharp U-shaped deflection of the ribs on the pe-
ripheral angulation, A weaker spiral cord is pres-
ent on the lower whorl face. Suture impressed;
whorls embrace just below lowermost spiral cord.
Base of holotype has subdued ribbing, otherwise,
base rounded. Finer collabral growth lines pre-
served. Aperture unknown, but growth lines sug-
gest it is rounded with a sharp, shallow
invagination in the outer lip. Protoconch un-
known.

REMARKS, This abundant taxon has the charac-
teristic ribbing and angular whorl profile separat-
ing it from other loxonematids. There is
superficial similarity to Loxonema magnificum
Spitz, 1907 and L. ingens Frech from the Lower
Devonian of the Carnic Alps. Both species are
much larger, L. ingens lacks the angulation, hav-
ing nodes instead and has finer ribbing, L. mag-
nificum, whilst poorly preserved, shows relicts of
nodes rather than à sinus-bearing angulation. An-
other superficially resemblant taxon is Trochus?
lamellasus Lindstrom, 1884 from the Silurian of
Gotland, but that taxon has a different whorl
profile, a flattened base and is clearly not a
loxonematid,

ETYMOLOGY. For Chris Tassell.

Loxonematidae indet.
MATERIAL EXAMINED. QMF32062 from QML54 1.

DESCRIPTION, Poorly preserved, large, many-
whorled, high-spired shell, 35mm long, approxi-
mately 15mm wide at with an apical angle of c.
5-1017. Sutures impressed, sutural slope moder-
alely high, approximately 15°. Coarse ribs sug-
gested in the specimen, other external shell
features not preserved.

REMARKS. Overall shape, sutural slope and
impression suggests affinity to the
Loxonomatidae Koken, The specimen resembles
Loxonema angelicum d'Orbigny of de Koninck
(1877; pl. 4, fig. 9), from the Early Devonian of
New South Wales but is too poorly preserved [or
reliable comparison.

Family PALAEOZYGOPLEURIDAE
Horny, 1955
Palaeozygopleura Horny, 1955

Palaeozvgopleura dodgeyi sp. nov.
(Fig, 9B-D)

MATERIAL EXAMINED. HOLOTYPE: QMF33604
from QML1092. PARATYPES: QMF33605-33607
from QML 1092, QMF33620 from QML 1090.

DIAGNOSIS. Small to medium-sized member of
the genus, with thick opisthocyrt ribs.

DESCRIPTION. Small to medium-sized, high-
spired gastropod which grew to slightly more
than 19mm high and 7-9mm wide, with an apical
angle of approximately 20? (Table 8). Base
rounded, anomphalus; sutures impressed, whorl
profile rounded with surface bearing numerous,
thick, opisthocyrt ribs which extend fron suture
to suture, with a slight subsutural shelf on the
uppermost whorl face. Finer collabral lines are
present between rugae on QMF33620, not pre-
served on other specimens. Aperture not pre-
served, whorl profile and growth lines suggest
that this was rounded, with broad sinus.

REMARKS. The species is smaller than. and
lacks the change in rib spacing of L. alticestatum
Tassell 1982 from the Early Devonian of New
South Wales. L. australis (Cresswell) of Tassell
(1978) is also similar, but has fmer, more numer-
ous rihs. The type species P. alinae (Perner) from
the Lower Devonian of the Prague Basin, has
finer ribbing, and is significantly smaller, This

74 MEMOIRS OF THE QUEENSLAND MUSEUM

GASTROPODS FROM THE BROKEN RIVER PROVINCE 75

TABLE 9. Measurements for Leprogvma queerislandi-
Cus sp. nay.

QMF31673 :
QME33674

OMF342(4-
QMF34209

species lacks the protoconch but is at the upper
range of size diagnosed for the genus by Horny
(1955). Both P. sibleyense Linsley and P. joanni
Linsley from the Middle Devonian Anderdon
Limestone of North America, are smaller, P.
Joanni has finer ribbing. P. muoni Tassell, 1982
from the Recepraculires Limestone of New South
Wales lacks the prominent ribbing, and is smaller.

ETYMOLOGY. For Paul ‘Dodgey’ Tierney and Scott
"Dodgey' Hocknull.

Family PSEUDOZYGOPLEURIDAE
Knight, 1930

PSEUDOZYGOPLEURIDAE gen. et. sp. indet.
(Fig. 9A)

MATERIAL EXAMINED. QMF33584 from
QMLIOSS,

DESCRIPTION. The specimen is 2 whorls of a
large, high-spired anomphalus shell. Basal width
is 21.7mm, with the height of the 2 whorls
26.4mm. Whorl profile rounded; periphery
slightly below midwhorl with the whorl surface
dominated by thick, widely-spaced, slightly sig-
moidal ribs. Suture impressed, embracing whorl
slightly below periphery. Base somewhat conical.
Preparation of specimen revealed no preserved
fine growth lines. Shell moderately thick.

REMARKS. Given the large size and the promi-
nent ribbing, the specimen belongs to the family,
but cannot be further indentified. Lexonema al-
tacostatum Tassell 1982 from the Recepraculites
Limestone of New South Wales is similar, but its
ribs are less sigmoidal.

Superfamily SUBULITOIDEA Lindstrom, | 884
Family SUBULITIDAE Lindstrom, 1884
Subfamily SOLENISCINAE Wenz, 1938

Soleniscus Bayle
TYPE SPECIES. Soleniscus typicus Meek.& Wortlien,
1861.

Solenisncus sp. cf. Solensiscus subcostata
Schlotheim (Fig, 10 E,F)

MATERIAL. EXAMINED. QMF33692-QMF33694
from QML 1019,

DESCRIPTION. Medium sized, fusiform, up to
25mm high, 16mm wide, with an apical angle of
c. 35°. Suture slightly impressed; whorls embrace
slightly below midwhorl. Whorl profile rounded,
periphery slightly above midwhorl. Surface or-
namented by numerous sinusoidal growth lines.
Aperture and base unknown.

REMARKS. Macrochilina Bayle was placed in
synonymy with Soleniscus Meek & Worthern by
Knight et al. (1960) but forms ascribed tọ that
genus have a generally squatter form than those
given to Soleniscus. A strikingly similar, but
much larger form is Macrochilina arculata
Sclotheim of Mansuy (1912) from the Givetian
of Yunnan, Macrochilina subcostata Schlotheim
of Whidborne (1892) is of similar size and orna-
mentation and the Broken River material is re-
ferred to that taxon. Following Knight et al.
(1960) both these taxa should be accommodated
within Soleniscus. Perhaps separation of elongate
and squat froms of the genus can be accommo-
dated in 2 sungenera Soleniscus (Solensiscus)
Meek & Worthen, and Solensicus (Macrochilina)
Bayle.

Leptogyma queenslandicus sp. nov.
(Fig. 10G-J)

MATERIAL EXAMINED. HOLOTYPE:
QMF33673, from QML1092, Papilio Mudstone,
Givetian, Broken River Province. PARATY PES:
QMF33674, QMF34203-QMF34213 all from
QML1092,

DIAGNOSIS. Small, turhiniform, Leptogyma
with very fine growth lines and weak wide sinus.

FIG. 9. A, palaeozygopleurid indet. QMF33584, apertural view x 2.2. B-D, Palaeozygopleura dodgeyi sp. nov.
B,C. Holotype QMF33608 x 3,5. B, apertural view; C, side view. D, Paratype QMF33607 side view x 3.5, E-L,
Australoxa tasselli gen. et sp. nov. EE Holotype QMF33586, x 3.3. E, side view; F, apertural view. G, Paratype
QMF233589, x 3,5 side view. H-J, Paratype QMF33708. x 3.6. H, apertural view; I, side view: J, side view. K,
Paratype QMF33588, x 3.5, side view. L, Paratype QMF33590, x 3.1, side view.

76 MEMOIRS OF THE QUEENSLAND MUSEUM

GASTROPODS FROM THE BROKEN RIVER PROVINCE 77

DESCRIPTION. Small, turbiniform gastropod,
up to 7.7mm high and 6.6mm wide (Table 9),
apical angle c. 25°. Sutures slightly impressed,
whorls join at periphery. Final whorl very large
in proportion to earlier part of conch. Protoconch
unknown. Whorl profile rounded, periphery
below midwhorl, upper whorl face occupies 2/3
of profile. Growth lines very fine, numerous,
closely spaced with a weak wide sinus. Base
rounded. Columellar lip thickended, probahle
minor fold.

REMARKS, Leptogyma Knight, 1936 is sepa-
rated from Auriptogyma Perner, 1903 by the thin-
ner, unfolded columella 1n the latter. Thickening
on the columellar lip confirms the generic identi-
fication. Leprogyma australis Tassel| 1982, from
the Early Devonian of New South Wales is higher
spired than the Broken River material.

ETYMOLOGY. For the state of Queensland,

Family CODONOCHEILIDAE Miller, 1889
Mitchellia de Komnck, 1876

TYPE SPECIES. M. siriatula de Koninck, 1876, from
the limestone of the Yass District, New South Wales,
by original designation,

DIAGNOSIS. See de Koninck 1877, or the 1898
translation.

DISTRIBUTION. Early Devonian (Emsian),
Receptaculites Limestone, Taemas, New South
Wales. The destroyed holotype specimen was
from black argillaceous limestone from the Yass
District, New South Wales and is most probably
Early Devonian; Middle Devonian (Givetian)
Papilio Formanon, Broken River Province, north
Queensland.

REMARKS. Knight et al. (1960) regarded
Mitchellia as a junior synonym of Scoliostoma
Braun, but as pointed out by Tassell (1982) the
gerontic growth stage of the aperture is twisted
downwards and, not upwards and is more con-
stricted where it joins the main spire.

Mitchellia striatula de Koninck, 1877
(Fig. 10, B-D)

MATERIAL EXAMINED. QMF32643 from SD21.

DESCRIPTION. Medium-sized. high-spired,
dextrally-coiled shell; early whorls not present in
this specimen which is 38mm high and has 15mm
maximum width. Apical angle is approximately
20°. Sutures moderately impressed; sutural slope
approximately 15°, Whorl profile rounded, and
rounded base. Aperture broken but. laterally con-
stricted at final growth stage to an elongate oval
shape. Ornament consists of more than 12 spiral
threads. At final growth stage the threads are
deflected downwards, indicating deflection of
apertural growth.

REMARKS. The aperture is incomplete, broken
at the point of deflection, but it is clearly con-
stricted and the spiral threads show a marked
downward deflection, identical to the more com-
plete but smaller silicified specimens described
hy Tassell (1982). The specimen is at least double
the size of those recorded by de Koninck (1876)
and Tassell (1982), but is retained in the species
due to Ihe morphological equivalence.

ACKNOWLEDGEMENTS

We thank Peter Jell for his encouragement and
assistance throughout this project. We thank John
Jell and John Talent for providing material from
Iheir collections. Phil Lawless, Scott Hocknull,
Paul Tierney, Colin McHenry, Donna Case and
Reints Lootsma are thanked for field assistance.
Robert Blodgett and Chris Tassell are thanked fur
their helpful comments on the manuseript.

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FIG. 10. A, Srylonema? sp. latex mould of QMF33102 x 7. B-D, Mirchellia striatula (de Koninck, 1876)
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'Novyie vidy iskolamyich shivotnyich I rastenti
byelorussii”. [New species of fossil animals and
plants from Belorissia]. (Akademi Nauk BSSR;
Minsk).

80 MEMOIRS OF THE QUEENSLAND MUSEUM

PTYCTODONT JAW FROM THE BROKEN RIVER
PROVINCE, NEQ. Memoirs of the Queensland Museum.
42(1): 80. 1997:- A single ptyctodont gnathal recovered from
the Papilio Mudstone (Middle Devonian, Givetian), Broken
River Province, north Queensland, represents the first macro-
fossil evidence for the family in the Middle Devonian of
Australia whilst both Early and Late Devonian ptyctodontds
are known (Long, 1991). Possible ptyctodontid microfossils
have been described from Emsian and Givetian units in the
Broken River Province by De Pomeroy (1994), The new
specimen is a gnathal with a well preserved tritoral surface.
The specimen was recovered from QML1018, SW of Storm
Dam, Wando Vale, where outcrops of the Papilio Mudstone
form part of an extensive muddy carbonate platform succes-
sion whose sedimentology has been detailed by Lang et al.
(1993). Jell at al. (1993) place the Papilio Mudstone at Storm
Dam within the varcus to hermanni-cristatus Conodont Zone.

Systematic Palaeontology
Order PTYCTODONTIDA Gross 1932
Family PTYCTODONTIDAE Woodward 1891
Ptyctodus Pander 1858
?Ptyctodus sp.
(Fig. 1)

1996 ?ptyctodontid indet., De Pomeroy: 431, fig. 6b, ?0,P

Material. QMF35438, from QML1018, SW Storm Dam,
Broken River Province, NQ. Papilio Mudstone, Middle
Devonian, Givetian. Collected A.Cook & N. Camilleri.
Description. Right? superognathal 44mm long; functional
edges steep and narrow with a short anterior cutting edge.
External surface relatively smooth with faint curving linea-
tion. Large depression on the internal surface. Gnathal widens
posteriorly to double width at about two-thirds along its length
(from 4mm to 8mm across); dental plate with smoothly con-
cave, sharp biting edge. Tritor 3mm wide, 17mm long with
approximately 5 tritoral dentine plates per millimetre.
Remarks. Ptyctodonts were predominantly marine bottom
feeders with a small gape, bordered with a series of few strong
beak-like dental plates (Denison, 1978). Ptyctodontid
gnathalia do not consist of bone but are pure dentine. Well
worn dental surfaces of this specimen appear to be preserved
as vivianite. Ptyctodont toothplates are commonly found as
isolated elements in the Middle and early Upper Devonian of
North America and Europe. In the last century many were
given specific names but cladistic analysis of the late twenti-
eth century has revised and suppressed many of these taxa.
Detailed study of such toothplates may prove them to be useful
biostratigraphic tools.

Acknowledgments
We thank E.D. McKenzie for assistance with preparation.

Literature Cited

DENISON, R. 1978. Placodermi. In Schultze, H.-P. (ed.)
“Handbook of Paleoichthyology', Volume 2. (Gustav
Fischer Verlag: Stuttgart).

DE POMEROY, A.M. 1996. Biostratigraphy of Devonian
microvertebrates from Broken River, north Queens-
land. Records of the Western Australian Museum 17:
417-437.

JELL, J.S., SIMPSON, A., MAWSON, R. & TALENT, J.A.
1993. Biostratigraphic summary. Pp. 239-245. In
Withnall, I.W. & Lang, S.C. (eds) ‘Geology of the
Broken River Province, north Queensland”. Queens-
land Geology 4.

FIG. 1. ?Ptyctodus sp., x 1.2. A, external (labial) view. B,
internal (lingual) view. C, occlusal view. D, anterior view.

LANG, S.C., JORGENSON, P., BLAKE, P., HUMPHRIES,
M, FIELDING, C.R., JELL, J.S., WITHNALL, I.W. &
DRAPER, J.J. 1993. Stratigraphy and sedimentology of
the Broken River Group. Pp 79-128. In Withnall, I.W.
& Lang, S.C. (eds) ‘Geology of the Broken River
Province, north Queensland'. Queensland Geology 4.

LONG, J.A. 1991. The long history of Australian fossil fishes.
Pp. 336-428. In Vickers-Rich, P., Monaghan, J.N.,
Baird, R.F. & Rich, T.H. (eds) ‘Vertebrate Palaeonto-
logy of Australasia'. (Pioneer Design Studios with
Monash University: Melbourne).

Susan Turner & Alex G. Cook, Queensland Museum PO

Box 3300 South Brisbane QLD 4101 Australia; 20 Decem-

ber 1996.

SYMBIOTIC STROMATOPOROID-NAUTILOID ASSOCIATION, MIDDLE
DEVONIAN, NORTH QUEENSLAND

ALEX G. COOK AND MARY WADE

Cook, A.G. & Wade, M. 1997 06 30: Symbiotic stromatoporoid-nautiloid association,
Middle Devonian, north Queensland. Memoirs of the Queensland Museum 42(1): 81-89.

Brisbane. ISSN 0079-8835.

The stromatoporoid Clarhrocoilona spissa encrusts specimens of Diademoceras obtained
from the Middle Devonian (Givetian) Papilio Mudstone, Broken River Province and Burde-
kin Formation, Burdekin Subprovince, north Queensland. Stromatoporoid growth com-
menced and flourished while nautiloids were in an upright living position. Diademoceras,
here described for the first time in Australia, is considered upright benthonic to barely
nektobenthonic. [ ] Nautilida, Devonian, stromatoporoid, symbiosis, Queensland.

Alex G.Cook & Mary Wade, Queensland Museum, PO Box 3300, South Brisbane, Queens-

land 4101, Australia; 10 March 1997.

Devonian nautiloids from northeastern Aus-
tralia are poorly known, despite the seminal
works of Teichert (1940) and Teichert &
Glenister (1952). Large and diverse faunas occur
in the Burdekin Basin (Jell, pers. comm.) but
are less diverse in the adjacent Broken River
Province.

Five fragmentary specimens of Diademoceras
were recovered from WSW of Storm Dam,
(QML1017 and BRJ62), from the Papilio Mud-
stone, Broken River Province, north Queensland.
Three specimens were recovered from the Burde-
kin Formation, Golden Valley area, near Fanning
River, Burdekin Subprovince, NQ. This is the
first description of Diademoceras from eastern
Australia. Clathrocoilona spissa is a widespread
encrusting stromatoporoid (Cook, 1994). Speci-
mens from Papilio Mudstone were weathered
from lime-mudstone units interpreted as having
been deposited on a shallow-water, open marine,
muddy shelf (Lang et al, 1993). The Papilio
Mudstone contains àn abundant fauna including
corals, brachiopods, stromatoporoids and con-
odonts, indicative of a Givetian age (Jell et al.,
1993). Material from the Burdekin Formation
was retreived from carbonate mudstone and
packstone units, interpreted by Cook (1995) as
representing deposition on a shallow to moder-
ately deep (5-60m) carbonate shelf within the
geographically restricted Burdekin Basin. The
Burdekin Formation also contains a diverse as-
semblage of corals, stromatoporoids, brachio-
pods and molluscs. Sparse conodont faunas
(Talent & Mawson, 1994) also indicate a Givetian
age.

STROMATOPOROID OVERGROWTHS

Five ofthe eight Diademoceras specimens pos-
sess a sheath of encrusting stromatoporoid, two
others have partial encrustation and the remain-
ing specimen has been abraded. Cut specimens
show attached corals (several auloporid and ru-
gose corals), but growth of these was dominated
by that of the stromatoporoid associates. Polished
blocks and thin sections were prepared to reveal
growth detail of the encrusters. The
stromatoporoid was identified as Clathrocoilona
spissa (Lecompte, 1951). Each sheath consists of
many growth phases of C. spissa, punctuated by
growth inhibition and termination surfaces (sensu
Kazmierczak, 1971). Growth was thickest on the
ventral margin of the nautiloid, thinner inside its
open coil. Many growth phases completely en-
veloped the shell indicating that development of
some phases was uninhibited by the nautiloid's
resting position on the substrate. These must have
grown while this part of the shell was raised
above the substrate. Other growth was more spas-
modic. Cresentic nodes on the shell flanks of
Diademoceras developed at the aperture. They
are likely to have protected siphons for the usual
paired inhalent water currents during pauses in
growth but became overgrown after the shell
grew further (Fig. 4c.) Thus we conclude that at
times the nautiloid positioned itself upright and
stromatoporoid growth commenced during the
life of the nautiloid. The nautiloid, encumbered
with such stromatoporoid encrustation, would
have hardly been capable of significant motion in
the water column, as observed by Wade (1988)
who mistook poorly preserved encrusting
stromatoporoid growth for part of a thick shell
wall. The availability to epizoans supports the

82 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Diademoceras submammilatum (Whiteaves) x 0.5. A-C, QMF32218, prior to sectioning. A, dorsal view;
B, ventral view; C, side view. D, E, QMF33877. D, dorsal view; E. ventral view.

suggestion that the form was benthonic or barely
nektobenthonic. Observed growth interruptions
would have occurred through limited partial
burial, shell rotation with growth or some other

benthic adjustments. Some growth interruptions
may be due to changes in sea-floor sedimentation
effecting the vitality of the stromatoporoid. Fur-
nish & Glenister (1964) have suggested that all

SYMBIOTIC STROMATOPOROID-N AUTILOID ASSOCIATION 83

Nautilida were nektobenthonic and posthumous
floaters, Considering the mass of encrusting
stromatoporoid, it is unlikely that significant post
mortem transport occurred here, except some by
wave action.

Encrustation upon nauitloids has been reported
hy Teichert (1964) who stated that 'irresepective
of normal bouyancy requirements such shells
could not have floated appreciably after the ani-
mal died' (Teichert,1964: K125). Frey (1988)
recorded bryozoan encrustation upon
Treptoceras duseri, a michelinoceratid from the
Ordovician of Ohio. These bryozoan sheaths
were uniformly thin, and Frey (1988) concluded
that whilst Treproceras duseri was nektonic, he
could not determine whether encrustation took
place during the life of the nautiloid.

SYSTEMATIC PALEONTOLOLOG Y
Phylum PORIFERA Grant
Subphylum STROMATOPOROIDEA
Nicholson & Mune
Order STROMATOPORELLIDA Stearn
Family STROMATOPORELLIDAE Lecompte
Clathrocoilona Lecompte 1951

Clathrocoilona spissa (Lecompte) 1951

REMARKS. Cook (1994) has described C. spissa
from the neighbouring Burdekin Subprovince
and noted its presence within Givetian strata of
Ihe Broken River Province. The material here is
adequate for identification, displaying the char-
acteristic irregular architecture, ovcluded irregu-
lar galleries in both tangential and vertical
section, and tripartite laminae.

Phylum MOLLUSCA
Class CEPHALOPODA
Subclass NAUTILOIDEA Agassiz
Order RUTOCERATIDA Flower & Kummel

REMARKS. Nautilida are often characterised by
their thin siphuncular walls, in which layering is
difficult to detect (Flower, 1964), although all
Nautiloidea may be observed or inferred to have
had two layers applied to a basal membrane
which is rarely seen in fossils (Wade, 1988),
Siphuncles in the Diademoceras material de-
scribed here are clearly layered, consisting of two
thick layers and possibly one thin layer (see
below).

Family RUTOCERATIDAE Hyatt, 1884

REMARKS. Most workers post-dating Flawer &
Kummel (1950) have agreed that Rutoceratidae,
or an inclusive larger taxon, is intermediate be-
tween Oncocerida and Nautilida (Kummel,
1964). Although Teichert (1967, 1988) changed
his view of the taxonomic status of Rutoceratidae
as the basal family to the basal suborder uf
Nautilida, Flower (1964: 1988) persisted in
recognising Rutoceratida between Oncocerida
and Nautilida, with Rutoceratidae as the basal and
nominate family. Diademoceras, which Flower
(1949) assigned to the Rutoceratidae, has open
coiling and a nodose shell. respectively rare and
almost unheard of in the Oncocerida. These arc
rather commonplace in Nautilida. They occur
here wilh a thick outer (supportive) layer in the
connecting rings, and thin inner (osmotic pump)
layer. Turek & Marek (1986) have found
oncocerid muscle scars in Ptenoceras, assigned
to Rutoceratidae, order not stated. Rutoceratidae
thus appear to be ‘not yet’ Nautilida. The material
we have is inadequate for major taxonomic revi-
sion, Faced with the choice of withdrawing à
moslly unseen and inadequately described
Rutoceratina to the Oncocerida or accepting
Flower's original evaluation of their status, we
accept his placement,

Diademoceras Flower, 1945

Diademoceras Flower, 1945; 677; Flower 1949: 74;
Kummel, 1964: 418; Zhuraleva, 1974: 124.

TYPE SPECIES. Diademoceras palmeri Flower,
1949. by original designalion from the Middle Devon-
ian (Givetian), Cherry Valley Limestone of New York.

REMARKS. Flower (1945) erected the genus,
but did not describe or figure the type material,
Flower (1949) fully diagnosed and described it.
He further remarked that additional taxa of
Diademoceras were found within the Manitoban
Limestone, of which D. submammilatum
(Whiteaves) was the only described species.
”Diademoceras ajense Zhuraleva, 1974, from the
Givetian of the southern Urals, is based on frag-
mentary material and is poorly known.
Diademoceras ventrolobatun Lai & Zhang, 1988
from the Middle Devonian (Givetian) Qiziqao
Formation is the most recently described member
of the genus.

84 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Diademoceras submammilatum (Whiteaves) x 0.5. A,B,F, QMF32219, prior to sectioning. A, dorsal
view; B, ventral view; F, side view. C-E, QMF32217. C, dorsal view; D, ventral view; E, side view.

SYMBIOTIC STROMATOPOROID-NAUTILOID ASSOCIATION 85

Diademoceras submamilatum
(Whitcaves, 1891) (Figs [-4)

Gyroceras submamillatum Whiteaves 1891:
10, figs, 1à,b.

Diademoceras submamillanus (Whiteaves) Flower
1949: 75.

MATERIAL EXAMINED, QMF11896, collected J,
Jell, from the Papilio Formation, SW of Storm Dam;
QMF32212, QMF32217-9; from QML1018, collected
N. Camilleri & A. Cook, Ikm S of Storm Dam, Papilio
Formation, Broken River Province, north Queensland.
QMF33877, QMF33879, QMF33880 collected D,
Johnson & R. Henderson, from the Burdekin Forma-
tion, Golden Valley area, ‘Fanning River’ Station, NQ.,

PRESERVATION. The nautiloid material is
heavily recrystallised. Stylolites are common,
and there is widespread loss of shell by solution,
Septa in particular, may have been lost by arago-
nite solution because the calcite chamber fill ts
well preserved leaving adjacent crystallised
chamber fills separated by thin spaces. Fre-
quently septa are represented only by a thin,
black, linear iron mineralisation, Since Allison
(1988) established the sequence of early diage-
netic minerals as francolite, iron sulfide, calcite,
itis reasonable to trust shell-like shapes and po-
sitions of thin layers of iron mineralisation. In thin
section these are represented by black lines which
not only follow either surface of the septa, but
may mimic septal or shell wall laminae when the
original structure is lost by solution or
recrystallisation. Here, as is common in open
marine environments, phosphatisation was
elided. Fibrous drusy calcite lined the chambers
in both sectioned specimens, but this was sporad-
ically replaced by coarsely crystalline calcite
which filled the remainder of the cumeral space.

Every thin section is deficient in shell material
in some areas. All specimens were Ireed by sur-
face weathering and have undergone some mod-
erm erosion and breakage. Some breakage
presumably predates burial, since no whole
whorls could be fitted together and body cham-
bers are poorly preserved, but the original empti-
ness of some siphuncles and chambers is
evidence against significant post-inortem Move-
ment on the sea floor although it was well above
the fair weather wave base.

DESCRIPTION. Shell large, vyrtoconic, up 10
[8cm high and 9em wide (Table |) representing
up to à complete volution. Coiling was exogastic
and open for there is no trace of dorsal contact on
fragments of any diameter. Whorl broadly de-
pressed, ovate in transverse section, with a height

107, pl.

TABLE |, Morphometric data for Diademoceras suh-
mamillatum (Whiteaves) from the Papilio Mudstone
and Burdekin Formation.

Node

i
height Won, | Volutión .
(mm
150 5 5 3 2
164 j 22

Specimen spacing
(mm)
OMF! 1896

QMF32212

QMF12207| 182

38

GE
Co:

125
omr as [|

to width ratio of 3:4, The smooth, broadly
rounded arch of the venter and dorsum continues
across the ventro- and dorsolateral areas to the
rather narrowly rounded lateral areas (Figs 1-3).
These are intermittently wholly tuken up us the
sites of large, anteriorly-facing flanged siphons,
like those of Prenoceras. As in Ptenoceras these
are closed hy subsequent growth which excluded
detritus and thus formed so-called spines. Their
detailed structure will be discussed below afler
shell walls. Posteriorly on the whorl the siphons
are represented by small peaks instead of large
spines, as previous descriptions have discussed.
Dorsolaterally the walls are more depressed thàn
ventrolaterally, ihey are almost flattened slopes.
rather than curves, so that the dorsum can appear
bluntly triangular overall. The siphuncle is near
ventral, with constricted, dorsally sub-
cyrtochoanitic necks which grade laterally ina
suhorthochonanitic necks ventrally (Fig. 4B].
The connecting rings are thick and obviously
two-layered. Moderate expansion of the connect-
ing ring within the chambers is consequent upon
neck shape. A thick outer layer dorsally is devel-
oped from most of the tip of the swollen septal
neck, but is more obscure in origin ventrally (Fig,
4B). This outer layer ends against the previous
septum, A thin inner layer arises from the whole
inner edge of the septal neck and adheres to the
thick outer layer to its termination around the
previous septal neck, thereafter the thin inner
layer adheres to the constricted previous neck,
and curved ad-posteriorly outward unti] i1 con-
tacts the inner layer of the previous connecung
ring. In this fashion the inner layers ure connected
lo one another throughout the siphuncle's length,
but the thick outer layers are localised to cach
chamber. Because of the curvature of the inner

86 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. Diademoceras submammilatum (Whiteaves) x 0.5. A-C, QMF 32212. A, side view; B,ventral view; C,
dorsal view. D-F, replica of QMF11896.

layer, it did not always match the previous neck ` sediment-filled space between the bent dorsum of
(or stay matched after death), there is often a the previous neck and the inner layer of the con-

SYMBIOTIC STROMATOPOROID-NAUTILOID ASSOCIATION 87

FIG. 4. Diademoceras submammilatum (Whiteaves), A, QMF33880, show-
ing longitudinal ribs, faint growth lines and ventral sinus; B, Photo negative
of QMF32218 slightly oblique through siphuncle, with encrusting
Clathrocoilona spissa (Lecompte), x 1.9.

necting ring; this space appears in many longitu-
dinal views of the siphuncle, and a similar space
may appear more rarely ventrally.

The septa are swollen toward the dorsum of the
neck and within it, but were otherwise thin. They
are recrystallised wherever preserved. The septal
flaps appear to have been long and relatively
thick but are nowhere clearly seen. The shell wall

appears to be two-tayered,
sometimes separated by a dark
line ànd otherwise by a gap.
Poor preservation and abun-
dance of stylolitisation within
these structures does not allow
us to determine with confi-
dence whether this layering is
primary or of diagenetic origin.
It has the appearance of typical
marine phreaüc cement, The
outer layer is coarsly prismatic,
almost fibrous and not nearly
as prone to recystallisation as
the inner Jayer. The inner layer
is thinner in smaller cross-sec-
tions, but locally varied in
thickness,

External ornament, as de-
scribed by Flower (1949), is
poorly known on most of the
specimens as five of the eight
specimens are completely
sheathed in encrusting
stromatoporoid. QMF33880
shows strong, narrow, longitu-
-> dipal ribs on the shell surface,

` and they are slightly reflected
on the internal mould (Fig.
4A). The ribs can be detected
by changes of shell thickness,
in cross-sections of other spec-
imens and also on the internal
moulds of some specimens. Su-
ture with broad, slight ventral
saddle, and broad ventrolateral
lobes, with narrow saddle on
the sharp umbilical angle;
nearly straight across the dor-
sum. Growth lines of QMF-
33880 indicate a ventral sinus.

The shell possesses a row of
short, rounded, thickly
cresentic nodes oriented con-
cave forward, relatively regu-
larly spaced in any individual,
up to 38mm apart. Sections in-
dicate blocky calcite growth
within the spines indicating they were hollow.

REMARKS. Whiteaves (1891) described à
poorly preserved taxon which is of similar size,
possesses similarly spaced relicts of nodes and
nearly straight sutures and hence is undifferenti-
able from the Broken River material. The type

88 MEMOIRS OF THE QUEENSLAND MUSEUM

species, D. palmeri, is a little smaller than
Whiteaves' taxon, and has weak ventral lobes but
may prove to he conspecific. D. ventrolobatum
Lai & Zhang from the Middle Devonian of
Guangxi, is significantly smaller, Wade (1988)
misinterpreted the worn stromatoporoid encrust-
ing QMF11896, the first specimen collected, in-
tepreting this sheath as very thick layered original
shell, because it faithfully reproduced the paired
spines and shell outlines, and the stromatoporoid
is very poorly preserved on that specimen,

Il the outer wall lamina was not always present.
forming symmetric lateral siphons and minor
structures, and passing below every kind of
epizoan, could be suspected of being an epizoan
100,

Separation of the hollow spines from the cham-
ber by shell wall growth preceeding septa forma-
uon is observed, and validated by the shapes of
exposed chamber fills which show the smooth,
low rises under the nodes, Only the clean node-
fills witness to front walls to the nodes, and the
valcite growth, interpreted as phreatic growth,
testifies to their shape as do the epizoan over-
growths. The smooth shell wall bases were per-
haps added to the living chamber soon after the
shell's siphons were closed, by reactivation of the
mantle. All cephalopods repair by mantle reacti-
vation, so this method is not unusual, and the
gradual forward movement of the growing body
would bring a smooth curve of body adjacent to
the space within the closed siphon. This. would
allow a slight bulge, and account for the observed
shape of the walls,

The undoubtably aragonitie nautiloid shell
recrystalhsed so completely, an explanation of
the relatively good preservation of the micro-
structure of the stromatoporord is required. Such
differential preservation in these specimens
would suggest that the original stromatoporoid
mineralogy differed from that of the nautiloid,
and by inference was probably calcitic.
Stromatoporoids have been variously inferred as
having skeletons which were calcitic (Galloway,
1957; Kershaw, 1990; Rush & Chafetz, 1991) or
aragonitic (Stearn, 1975: Stearn & Mah, 1987).
This material shows circumstantial evidence that
the stromatoporoid was.calertic, but demonstrates
no more than association of preserved culcite
phases than Kershaw (1990),

ACKNOWLEDGEMENTS

We thank John and Peter Jell for encourage-
ment and some material. We thank Bob Hender-

son for providing the Burdekin Formation mate-
rial. The photography and library sections of the
Queensland Museum are thanked for their gener-
ous assistance.

LITERATURE CITED

ALLISSON, P.A. 1988. The role ol'anoxia in the decay
and mineralisation of proteinaceous microfossils.
Paleobiology 14: 139-154.

COOK, A.G. 1994. Middle Devonian stromatoporoid
faunas and palaeoecology of the Burdekin Sub-
province, Ph.D Thesis (Unpubl.). James Cook
University. 264p.

1995, Sedimentology and depositional environ-
ments of the Middle Devonian Big Bend Arkose
amd Burdekin Formation, Fanmng River Group,
Burdekin Subprovince, north Queensland, Aus-
tralia. Memoirs of the Queensland Museum
38CI): 53-91.

FLOWER, R.H, 1945. Classification of Devonian nau-
tiloids. American Midland Naturalist 33(3): 675»
724.

1949. New genera of Devonian nautiloids. Journal
of Paleontology 2301): 74-80.

1964. Nautiloid shell morphology. New Mexico
Bureau of Mines and Mineral Resources Memoit
13: 1-78,

1988. Progress and changing concepts in cephalo-
pod and particularly nautiloid phylogeny and dis»
tribution. Pp. 17-24. In Wiedmann, I. &
Kulimann, J. 2nd Intemational Cephalopod Sym-
posium. Cephalopods Present and Past. OH.
Schindewolf Symposium, Tubingen 1985.
(Schweizerbart'sche Verlagbuchhandlung, Stutt-
garn).

FLOWER, RH & KUMMEL, B. 1950. A classificalton
of the Nautiloidea, Journal of Paleontology 24(5):
604-616.

FREY, R.C. 1988. Paleaecology of Treptoceras duseri
(Michelinoceratida, Proteoceratidae) from Late
Ordovician of southwestern Ohio. New Mexico
Bureau of Mines and Mineral Resources. Memoir
44: 79-101.

GALLOWAY, J.J. 1957, Structure and classification of
the Srromatoporoidea. Bulletins of American Pa-
leontology 37 (164): 345-471.

JELL, J.S., SIMPSON, A., MAWSON, R. & TALENT,
LA. 1993. Biostraligraphic summary. Pp, 234
245, In Wilhnall, LW, & Lang, S.C. Geology of
the Broken River Province, north Queensland,
Queensland Geology 4. (Department of Minerals
and Energy: Brisbane).

KAZMIERCZAK, J. 197]. Morphogenesis and sys-
tematics of the Devonian Stromatoporoidea from
the Holy Cross Mountains, Poland. Palaeo-
ntologia Polomca 26: 1-144.

KERSHAW, S. 1990. Stromatoporoid palacobiology
and taphonomy in à Silunan biostroma on Got-
land, Sweden. Palaeontology 33(3): 681-705,

SYMBIOTIC STROMATOPOROID-NAUTILOID ASSOCIATION 89

KUMMEL, B. 1964. Nautiloidea-Nautilida. Pp. K383-
K457. In Moore, R.C. (ed.) Treatise on Inverte-
brate Paleontology. Part K: Mollusca 3.
(Geological Society of America and University of
Kansas Press: Lawrence, Kansas).

LAI CAIGEN & ZHANG ZHENXIAN, 1988. Nauti-
loids from Early and Middle Devonian of Hunan
and Guangxi. Professional Papers of Stratigraphy
and Palaeontology 21: 29-51.

LANG, SC JORGENSEN, P., BLAKE, P.,
HUMPHRIES, M., FIELDING, C., JELL, J.S.,
WITHNALL, LW, & DRAPER, J.J. 1993. Stra-
tigraphy and sedimentology of the Broken River
Group. Pp. 79-128. In Withnall, LW. & Lang, S.C.
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oO MEMOIRS OF THE QUEENSLAND MUSEUM

NEW INFORMATION ON THE NARROWLY-
RESTRICTED SKINK, NANGURA SPINOSA. Memoirs
af thè Queensland Museum $3(1) YO. 1997:- Surveys of
vertebrates Of dry rantorests of soul and mideastern Queens-
land during 1992 (Horsup et. al, 1993) led to the discovery of
ihe skink Nenguro spinosa Covacevieh, Couper & James,
1993, ‘This pn distinetive, burrowing skink which was found
in å 300m section of dry, gertly sloping creek bank in semi
evergeeen vine ket iñ Nangur State Forest (26075
151*58'E), 20 km north of Murgon, SEQ. This was, until
recently, the only known locality for N. SDUHOSR;

On the 27 February, 1997 one of us (GA) observed å tail
protruding from å burrow on å road embankment in Oak view
Site Forest, SHO (26°07 24S 1527 TONE), An adult skink
wits removed from the burrow, It was photographed (QM
iransparencies NR 360 - NR 373) and released, While being
handled. the specimen shed its tail. This has been registered
m the Queensland Museum (QMJ62787), where our identifi-
cation of the skink as V, spinasa was confirmed Oak view S.P
is i new locality for N, spinosa. The collection sile is approx»
imately 40km east of Nangur S.F.

Additional searches along the road embankment for N.
splnasa i Oak view S E on 28 February and G March, 1997
revealed 22 ‘active’ burrows. Two further burrows were ob-
served 7m and 27m up-slope from the road. Thirty-six speci-
mens were seen 23 adults, E sub-adulrand 12 juveniles Bach
burrow was occupied primarily by a single adult (n= L8). One
was pceupied hy 2 Se one by 2 adults and 3 juveniles;
three hy I adult and 2 juveniles; and three by T adult and I
juvenile, Ouc adult, I sub-adult and 2 juvenile N spinosa were
weighed and (heir snouuyent (SVL) recorded, The adult mea:
sured 92,0mm S VL. and weighed 26gm, the sub-adult mea-
sured 78.2 mm SVL and weighed 14.1 gm; juveniles
suspected To be neonates (with umbilical scars) weighed
I SED (d= 0, n = 2) and had an average 378mm SVL (s.d,
«lå,n=2)

The vegetation at all sites was Araucarian Notophyll Vine
Forest (Horsup et al., 1993) am Quarternary alluvials, at an
altitude of approx. 600m. Burrows in the road embankmert
(mean road embankment height = 106, 90m, sd. = 266, n =
23) were in three clusters along 1,4km of road and were sel
inco the bank ar an average height of 55 jem (sd = 25,5, n=
23) above the road. Burrow entrances were usually remote
trom ground cover (n = 15), associated wilh rocks (n — 5) or
associated with tree bases or surface roots. (n = 4). The only
exception was a burrow ar the base of an old termite mound,
Average size of burrow entrances was & lem wide (s.d. — 3.3.
n=24) and 4.9cm high (s.d. 2 2.1, n 2 23). Each burrow hada
shoot ‘resting platlorm' near the entrance, with an averaye
width of HL, lem (s.d, 24,7, n 223) and a length of 8. Germ (s, d.
= 36, n 2 23). All had north-eusterly aspects.

During daylight at Oak view S.F., we observed individuals
either with tails visible at burrow entrances, heads slightly
promidimg from burrow enuances or on the resting platforms.
One juvenile was pbserved approximately 15em beyond a
resting platform" When approached, i1 rerreated into the
burrow. Nocturnal observations of road side burrows between
21:22 and 22:09hrs revealed 16 N. spinosa with tails visible
ut burrow entrances. The ambien! temperature ar this time was
2)°C Specimens of N, spinosa apparently ‘rest’ ut burrow
entrances from where they ‘ambush’ prey (Coyacevich er al...
1993; Wilson, 1994), The presence of individuals at burrow
entrances in Oak view S.F. may relate to either thermoregula-
ror or foraging behaviour.

Faecal samples ot N. spinosa from Nangur S.F. contained
u diverse range of anhropod remains (Covacevich et al.,
1993). Locating visible evidence of faecal or feeding remains

au the Oakview site was hindered by heavy rains, A beetle
carapace was the orly possible prey remnant found close ro à
burrow,

Searches of road embankments in both ap ar and differ
ent soils and with similar aspects and vegetation in Oakview
SF, faded to reveal further evidence of N. spinosa. Much of
the forest has been planted with Hoop Pine, Araacarta
ennminghanii, leaving remnant strips of Araucarian
Notophyll Vine Forest between plantation. compartments;
Larger remnants of NVF occur only on the steeper slopes
where plantation establishment was not practical. The pres-
ence of Pwo active burrows on slopes away from the road
embankment shows that this species 19 nor restricted to road
enibankments or the creek banks reported by Covacevich et
al, (1993),

N. spinosa was w "species known only from (he pe
collection’, and ‘rare In Australia, bur nor currently consid-
ered endangered or vulnerable ...' (Covacevicher al, 1993),
Under the Name Conservation (Wildlife) Regulation 1994,
this species was "rate" in Queensland, With the discovery of
this second population, the status of N. spinosa should be
changed to that of å ‘species witha very restelered disrribunon
in Australla and with a maximum geographical distribution
of less than 100km `. ` It remains “a species... rare in
Australia .,. not currenily considered endangered or valier-
able 4 (Thomas & McDonald, 1989), Clearing of rainforest
has ceased on Queensland state forests. However, dry
rainforests have no legislative protection (Covacevich & Me-
Donald, 1993), On freehold land they are still cleared and
survive only as remnants, In state forests, vine thickets con-
linue to suffer deleterious changes due to inappropriate fire
regimes, disturbance by domestic stock and Hoop Pine tog:
ging. Further work is required to define the distribution of N.
spinosa and to investigate relationships between supporting
vegutalion, sopls and aspect, This information would be usetul
in refining the conservation status of N, spinosa and rm devis-
ing management plans to conserve this species,

Acknowledgments

Thanks are duc 10 Geoffrey Smith and Teresa Eyre for
assistance with the preparation of this note. Funding lor this
work was provided through the Forest Wildlife Section’s
involvernent in the Comprehensive Regional Assessment
Vertebrate Fauna Survey program In Queensland, Jeanette
Coyacevich and Patrick Couper assisted us to prepare these
data for publication

Literature cited

Covacevich, J.A., Couper, PJ. & James, C, 1993, A new
skink, Nangura spinosa gen. et sp. nov, from å dry
vainforesr of southeastem Queensland. Memoirs of the
Queenstand Museum, 34(1): 159-167.

Covacevich, J, A, & Mcdonald, KR 1993. Distribution and
conservation of [rugs and reptiles of Queensland
rainforests, Memairs of the Queensland Museum
341}: 189-199.

Horsup, A., James, C. & Porter, G. 1993. Vertebrates of dry
ramtorest of south and mideastem Queensland, Mem-
oirs of the Queensland Museum, 34(1); 215-228

Thomas, M.B. & McDonald, W.J.F. 1989. ‘Rare and threat»
enged Plants of Queensland’. 2nd ed. (Depanmenr of
Primary Industries: Brisbane),

Wilson, S 1994. Unknown lizard. Geo. 16(3): 68-76,

D. Hannah, G. Agnew, B. Hamley-& E. Hogan, Forest Wildlife

Necnon, Resource Sciences Centre, Deparment of Natural

Resources, PO Box 631, Indooroopilly, Queensland, 4068,

Australia; 31 March 1997.

A NEW SPECIES OF SALTUARIUS (LACERTILIA: GEKKONIDAE) FROM
GRANITE-BASED, OPEN FORESTS OF EASTERN AUSTRALIA

PJ, COUPER, CJ. SCHNEIDER AND L.A, COVACEVICH

Couper, P.J., Schneider, C.J, & Covacevich, J.A. 1997 06 30: A new species of Saltuarius
(Lacertilia: Gekkonidae) from granite-based, open forests of eastern Australia. Memoirs of
the Queensland Musewun 4241y. 91-96, Brisbane. ISSN 0079-8835,

Saltuarius wyberba sp. nov., from the *granite-belt of southern Queensland and northern
New South Wales, is separated from three of the four previously-known members of the
genus in lacking male preanal pores. From the fourth, S. swaini (an obligate rainforest
species), it is distinguished by smaller size, greater dorsoventral compression, shape and
spinosity ofthe attenuated tail tip, colour and pattern. Analysis of mitochondrial cytochrome-
b sequence data shows that S. wyberba sp, nov. represents an evolutionary lineage indepen-
dent of the S$. swaini populations of southeastern Queensland, Salmarius Sqamata,
Gekkonidae, granite-based forests, Queensland.

JJ Couper & J.A, Covacevich, Queensland Museum, PO Box 3300, South Brisbane,
Queensland 4101; C. J. Schneider, Zoology Department, University of Queensland, Queens-

land 4072, Australia 6 January, 1997.

In their recent revision of Saltuarius, Couper et
al, (1993) recognised four species: S. cornurus, S.
salebrosus, 5. occultus and S. maint, That
Saltuarius specimens from southeastern
Queensland's ‘granite belt" differ morphologi-
cally (size, proportion) ånd in colour and pattern
from those from rainlorests of southeastern
Queensland is well documented (Covacevich,
1975; Couper et al., 1993), Couper et al. (1993)
assigned them to one of three ‘forms’ of S, swaini.
The 'heavily-blotehed' form was recognised
from 'dry eucalypt/granite habitats centred on the
Stanthorpe area, SEQ, and the New England Ta-
bleland, NSW'. The rainforest populations af
Saltuarius in southeastern Queensland belong to
S. swaini, Those from the open forests of the
Stanthorpe aréa, SEQ are not simply a ‘form’ of
8. swaini. Previously-recognised morphological
differences, supported by biochemical data, indi-
cate that they belong to a distinct new species.
The status of Saltuarius specimens from the New
England Tableland, NSW remains enigmatic be-
cause of scant biochemical data from these pop-
ulations, Selruarius specimens from the granite
habitats of SEO are easily-separated from S.
swaini. Recognition of this new species necessi-
tates modification to (he description of S. swaini,
and to the previously published list of specimens
referred to that species by Couper et al. (1993).

Morphological characters follow Covacevich
(1975) and Couper et al, (1993). Skeletal defini-
tions follow Bauer, (1990), The following abbre-
vialions apply: snout to vent length (SVL); tail
length (T), from posterior margin of cloaca to tip
of tail; anenuated tip of original tai] (TT); head

length (HL); head width (HW); head depth (HD)
lower jaw to top of head, between eyes; som
length (S). Additional measurements include -
length of front leg (L1) axilla to tip of longest
digit; length. of hind leg (L2) groin to tip of
longest digit; neck length (NL) axilla to posterior
margin of ear, Specimen designators: Australian
Museum (AMR), Queensland Museum (QMI)
and South Australian Museum (SAM). Compar-
ative material is listed in Appendices I and 2.

SYSTEMATICS

Saltuarius wyberba sp. nov.
(Figs 1-3)

MATERIAL EXAMINED. HOLOTYPE: QMJOL54 I
(Fig, 1), Girraween NP, Granite Areh Trail, 1-5km
from Bald Rock campground, granite boulders in open
forest (28950, 15175604) SEQ; C. Schneider, P.
Couper, M. Lara & J. Girling; 11 Nov 1995, Tissues
from this specimen have been lodged with the SAM.
PARATYPES: AMR92121, R92123, 6km W of
Amiens (28°34°S, 151%46'E) SEQ; AMR98332,
Approx 1.5km NW of Amiens (28734' S, 151°46'E)
SEQ; QMJ35401, Boonoo Fulls, via Tenterfield
(28"48'S, 152*10'E) NSW; QMI25374, Gimiween
NP. via Stanthorpe (28*50'S, 151"55'E) SEQ:
QMJI28648-49, Girraween area, nr Wyberba
(28°50°S, 151°S5°E) SEQ. QMJ29115-J29117,
Stanthorpe area, ?Girraween (24°M)'S, 151?55'E)
SEQ: QMJ30677, Stanthorpe. Aztec Temples, nr
Underground R (28"50'8,152*05"E) SEQ: QMJ-
27349, Girraween NP, nr (28°50'S,151°55'E) SEQ,
QMJ51633-151636, Girraween NF, Natural Arch
track (28°50'S,151°55'B) SEQ: QMJ51093,
Girraween NP, edge, outside park (28°50'S, 151°S6'E}

92 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. The holotype of Saltuarius wyberba sp. nov. (QMJ61541).

SEQ; QMJ61539-40, J61542-45, Girraween NP,
Granite Arch Trail, 1-5km from Bald Rock camp-
ground (28°50’, 151?56.04') SEQ; QMJ50345,
Girraween NP (28?51'S, 151°5F E) SEQ; QMJ54847,
Bookookooara, Boonoo SF (28°51’S, 152?1l'S)
NSW; QMJ30420, Wyberba, nr (28"52'S, 151?52' E)
SEQ.

ETYMOLOGY. W yberba is a rail-siding on the west-
ern boundary of Girraween National Park, in the
Stanthorpe area, SEQ. The name is reported to be of
Aboriginal origin, and said to mean ‘at the end of the
mountain' (Harslett & Royle, 1980). The epithet is to
be treated as a noun in apposition.

DIAGNOSIS. Saltuarius wyberba sp. nov. is dis-
tinguished from Saltuarius spp. except S. swaini,
in lacking male preanal pores. S. wyberba is
separated from S. swaini by size (max SVL
109.2mm vs 131mm); in being more dorsoven-
trally compressed (mean HD 38% HW n=26 vs
45% n=35), and by the shape and spinosity of the
attenuated tail tip (finely tipped with only minute
tubercules, Fig. 2a vs bluntly tipped with large
tubercules, Fig. 2b). Colour pattern also dis-
tinguishes these two species. S. wyberba is grey
to tan, with a pale vertebral stripe and heavy
brown to black blotching. S. swaini is grey or
mid-dark brown with lichen-like, dark-edged,

dorsal blotches or grey/brown with a pale verte-
bral streak. S. wyberba has a wide, open V-shaped
marking between the eyes (Fig. 3a). In S. swaini
this is a narrow deep V (Fig. 3b).

DESCRIPTION. SVL(mm): 75.4-109.2 (n 2 27,
mean = 95.2). Proportions as % SVL: T = 68.4-
80.9 (n = 11, mean = 72.4); TT = 26.8-32.4 (n=
11, mean = 29.4); L1 = 40.1-51.6 (n = 25, mean
= 44.9); L2 = 51.0-59.7 (n = 24, mean = 55.3);
HL = 26.4-29.5 (n = 27, mean = 27.8); HW =
20.5-23.7 (n = 27, mean = 22.3); HD = 7.2- 9.7
(n = 26, mean = 8.5); S = 11.0-12.9 (n = 27,
mean = 12.1); NL = 15.6-21.9 (n = 25, mean =
19.4).

Head. Large, depressed, triangular, distinct
from neck; covered in small granules which are
intermixed with large rounded to conical tuber-
cules; skin of head coossified with skull; rostral
completely divided by a single deep groove (n
= 26), or almost completely divided (n = 1);
rostral contacting nostril; ear opening ellipti-
cal, vertical, much less than half as large as eye;
supralabials 13-18 (n = 54, mean = 14.6); in-
fralabials 10-14 (n = 54, mean = 12.2); Tongue
colour in life, grey/purple.

Neck. Moderate to broad, 31-48% HW.

pell PE OT AE GT KJENT diele Ak BE DAT IAE FEED ET ART A piii tibt EE EE LE EET

A NEW SALTUARIUS FROM EASTERN AUSTRALIA

Body. Moderate, depressed, covered in small
granules; dorsal granules intermixed with larger
conical tubercules; tubercules moderate to large
on back, flanks and neck; lower flank tubercules
small to large, sometimes associated with a lateral
flange running from axilla to groin; basal scales
surrounding flank tubercules slightly enlarged;
granules on throat noticeably smaller than those
on chest and belly.

Preanal pores. Absent.

Limbs, Long, covered in pointed tubercules
dorsally; digits strong, compressed distally; sub-
digital lamellae (fourth toe) 18-24 (n= 54, mean
— 20.4).

Original tail. (n 2 11) Depressed, broad and
contracted at base and attenuated at tip; anterior
flared portion surrounded by an undulating flange
which bears slender, sharply pointed tubercules
around its margin; dorsal surface of tail (except
along midline of flared portion)covered in large
conical tubercules which are particularly pro-
nounced on the attenuated tip: tail tip slender
and free of tubercules, or with only minute tuber-
cules; number of rows of enlarged spinose tuber-
cules anteriorly across the attenuated tip 4-6 (n =
11, mean = 5.5%; attenuated tip accounts for 38-
45% of total tail length; ventral surface smooth
with a shallow groove along the midline of atten-
uated tip.

Regenerated tail. (n= 11) Depressed, broad and
leat-like contracted at base and only just attenu-
ated at tip; tail margin is a broad, thin flange which
bears minute spinose tubercules around the edges;
tail free from spinose tubercules on both dorsal
and ventral surfaces; ventral surface without any
indication of a shallow groove along the midline.

Skeletal Features. Supraocular portion of frontal
grooved; anterior process of interclavicle not present;
epipubic cartilage expanded: presacral vertebrae
25; sacral vertebrae 2; lumbar vertebrae 2; Ist
autotomy septum 6; abdominal vertebrae bearing
reduced ribs 4; rib free cervicals 3; cervical ver-
tebrae not elongate; sternal ribs 3; mesosternal
ribs 2: based on QMJ291 15 (alizarin stained).

Pattern (in spirit), Dorsum tan or grey; heavily
marked with dark brown/black or grey blotches
on head, body and limbs; a narrow vertebral

FIG. 2, Attenuated, original tail tips of the Saltuarius
species. A, S. wyberba (QMJ61541), with a finely
tipped tail, and only minute tubercules; B, S. swaini
(QMJ5649), with a bluntly tipped tail, and large lu-
bercules.

94 MEMOIRS OF THE QUEENSLAND MUSEUM

A

ILLE EE EL E E E E rr b br E TES CL TET HERE RAN TOM! CM

FLEET!

FIG. 3. Markings between the eyes in the Salruarius species, A, S. wyberba (QMJ61541), with a wide, open "V;

B, S. swaini (QMJ51638), with a narrow, deep 'V^.

stripe, broken by four irregular tan or grey
blotches, extends from neck to base of tail; a
wide, open V-shaped marking between the eyes
(Fig. 3a); labials light grey, mottled with dark
brown; toes prominently marked by alternating
pale and dark crossbands. Venter cream with clus-
ters of dark brown granules which often form
irregular bars below the infralabials, and on the
anterior margin of the thigh. Original tail grey to
lan above, marked with four to six irregular pale
crossbands which extend to the ventral surface on
the attenuated tail tip; cream to pale grey below,
mottled with brown, and with a series of pale
blotches along the midline of the anterior flared
portion, Regenerated tail pale grey, or tan with
darker marbling.

Measurements and Scale Counts, Holotype
(QMJ61541); SVL(mm): 97.56, T = 69,55, TT=
26.19, LI = 43.0, L2 = 53.6, HL = 26.0, HW =
21.0, HD =8.2, $= 11.6, NL = 17.2, supralabials
13/13, infralabials 13/12, subdigital lamellae (4th

lI b IC.

GENETICS

Analysis of mitochondrjal cytochrome-b se-
quence data supports the recognition of
Salluarius populations from granite habitats in
the Stanthorpe area (28?40'S, 151°56 E) SEQ, as
disunct from A swaini sensu stricto. 372 base
pairs of the 5' end of the light strand (and corre-
sponding heavy strand) of the cytochrome-b gene
were sequenced from 7 individuals from
Girraween National Park (28°50’S, 151?55' E),
near Stanthorpe, and compared with homologous
sequences from all species of leaf-tailed geckos
including 2 individuals representing S. swaini
from Lamington National Park (287]4'8,
153°08'E) and Mt Tamborine (27°58’S,
153°1 UE). Amplification of target DNA was
achieved from CsCl gradient purified mtDNA
and/or total genomic DNA extracts with primers
Ph-] and MVZ04 (primer sequences available
from CJS). PCR was performed in 25 | reactions
with 1.5mM MgCl, 0.5U Taq polymerase (Pro-
mega), 1X Promega Thermobuffer, 60M each
dNTP, 0.2M each primer, and 30 cycles with 45

A NEW SALTUARIUS FROM EASTERN AUSTRALIA 95

seconds at 94°C, 45 seconds at 45°C, and 45
seconds at 72°C. Automated sequencing of dou-
hle-stranded. products followed manufacturer's
(ABI) suggested protocols. Analyses are those of
one of us (CIS), unpub, data, Phylogenetic anal-
ysis reveals that Girraween sequences form a
strongly supported sister group to a monophyletic
group — composed of the Lamington-Mt Tam-
bourine sequences (100% of bootstrap replicates
in parsimony analysis with all characters unor-
dered and equally weighted). Sequences From the
Girraween group differ from the Lamington-Mt
Tamborine group at 13.8% of sites, a level of
difference similar to that among species of the
closely related genus Phyllurus (P. isis, P.
neprhys, and P. ossa differ at 10,0-13,9% of ho-
mologous sites). These data, in combination with
morphological differences indicate that the
Girraween NP Salruarius populations represent
an evolutionary lineage independent of the
Lamington-Mt Tamborine populations,

Tissue samples from Sualtuarius specimens
from localities in New South Wales are scant,
However, tissues (AMR 141964-5, tissue sample
numbers NR878-9) were obtained from speci-
mens in {wo populations of Saltuarius at
Chelundi State Forest (30*01'07"S, 152°30°02"E
& 30*03'04" S, 152*21'36"E), NSW. Chelundi
SF is near Guy Fawkes NP, approximately
I45kms SW of Girraween NP. Cytochrome-b
sequences from these individuals differ slightly
from each other (0.5%) and form a sister group to
the Saltuarius populations of Girraween NP,
SEQ. Importantly, these sequences differ from
the Girraween samples at approximately 11% of
sites and from Lamington-Mt Tamborine, SEQ
sequences ål 12.6-13.1% ol sites. Given the level
of sequence differences and the discontinuity of
suitable habitat, it seems unlikely that the
Chelundi and Girraween populations are conspe-
cific, However, in the absence of comprehensive
data, their status remains uncertain.

REMARKS

Populations of S. wvberba from the granite-
based forests ol the Stanthorpe area, SEQ are
morphologically and genetically distinct from
populations of .S, swa/nr occurring in the
rainforests of southeastern. Queensland. Speci-
mens QMJ53984, AMR141964-65, AMR43870,
AMR 123490 and AMR 149768, listed in Appen-
thx 2, are of uncertain stàtus, Morphology and
colourpatiern of these specimens readily sepu-
rate them from 5. yweind, Although they are sim-

ilar in Some respects to S. wyberba (colour/pat-
lem) they have been excluded from the type series
because morphological variations set them apart
from S, wyberba from SE Queensland, (Speci-
men QMJ53984, Mann R. Nature Reserve, NSW,
has large, scattered eranules intermixed among
the small granules of the chin. There is nn evi-
dence of this character in any of the specimens
from the Stanthorpe area). Saltuarius specimens
similar in colour and pattern to S. wyberba have
been recorded from dry, open forests associated
with granite as far south as Armidale, 30°31°S,
151°40'E (H. Hines pers, comm.). The possihility
that isolates of granite-based forest in N NSW
may support several species of Saltuarius is not
without precedent, In å recent review, Couper et
al. (1993) described three new species of Pi:
llurus from rainforests in coastal, mid-eastern
Queensland. All occur in extremely close prox-
imily on rainforest-covered mountains, their pup-
ulations being separated by narrow corridors
(8-30k m) of open forest. Two of these species (P
nepthys and P. ossa) may be sympatric in the
Clarke Ra. A parallel situation may occur in the
granite-based, open forest habitats of SE Queens-
land and N NSW {28°40 -30°31 S). Here, ex-
posed granile occurs as isolated outerops and
distinct gorge systems ina ‘sea’ of dry, open forest
(New England hardwoods), It seems possible that
each separate ‘island’ may also support a distinct
Saltuarius sp. Assessment of the stalus of these
Saltuarius populations hinges on extensive col-
lecting and genetic sampling throughout the
‘granite belt” of SE Queensland and N NSW.

Although S. swaini sensu stricto appears ro he
a rainforest species, il may occur in sympatry
with S. wvberha in granite-based open forests in
the Amiens region (28734' S, 151746" E), SEQ. A
single specimen (LAMR92122) morphologically
indistinguishable [rom S. svant, has been col-
lected from *6km NW Amiens’ (Couper et al.,
1994). Available data suggest S. swaini iS All
obligatory rainforest species, with the exception
of this enigmatic specimen. There is no reason Là
question data associated with AMR92122, Rass
Sadher of the Australian Museum (in litt, 23
April, 19496) regards the collector of this speci-
men as supplying data that was "usually better
than average’, Specimen AMRY2122 has been
formalin-fixed. Attempts lo extract useable DNA
for å genetic assessment of this specimen were
unsuccesstul.

96 MEMOIRS OF THE QUEENSLAND MUSEUM

S, SWAINI SENSU STRICTO

With the recognition of S. wyberba, the follow-
ing changes apply to the description of S. swaini
Wells & Wellington (Couper et al, 1993). Mate-
rial examined: specimens QMJ35401, QMJ-
24250, QMJ27349, QMJ25374, QMJ28648-9,
QMIJ29115-7. QMJ30677, QMJ51093, QMI-
51633-6, QMJ54847, QMISO0345, QMJ30420,
AMR92121, AMR92123, AMR98332, AMR-
110510 are S. wyberba, not S. swaini. Specimens
QM153984, AMR141964-65, AMR43870.
AMR 123490 and AMR 149768 (Appendix 1) are
Saltuarius sp. incerta cedis. Morphology: S.
swaini has a deep head (mean head depth 45%
HW) and the attenuated tail tip has large spmes,
and terminates. bluntly (Fig, 2b), Meristics: the
range for each measurement provided by Couper
et al. (1993) remains unchanged (all measure-
ments lor S. wyberba, except HD, fall within the
range previously given for S. swaini). Colour/pat-
tern: two principle colour forms exist: 1) grey-
medium brown with both paler and darker
blotches in the base colour; these blotches are
edged with brown or black lines to give a "lichen-
like" effect; often with a pale vertebral streak. 2)
grey or mid-dårk brown with a pale vertehral
streak,

ACKNOWLEDGEMENTS

The authors thank the Co-operative Research
Centre for Tropical Rainforest Ecology and Man-

APPENDIX 1.

Specimens of Salruarius swaini examined in the
current study. All localities åre for Queensland unless
otherwise indicated.

QMI398, J2409, 12933-34, 13254, J4439, 18183,
18359, J8861. 110440, 312257, J51095 Mi Tamborine,
(27°55'S, 153*10'E); 14819, Mt Tamborine, Eagle
Heights (279555, 153^12" E); 1148 Canungra Ck
(27°58'S, 153°09 E); 13215 Canungra (28?0l'S,
15391 lE); 14198, J5690 Mudgeeraba (28"05'S,
153*22'E); J5649 Flying Fox Valley, Beechmont
(28"08'S, 153?12'E); J3313 Tallebudgera (28"08'S,
153?26E'); 15382 Lamington NP (28^12'S,
153?05'E); 18646 Lamington NP Binna Burra
(28?12'S, 153*11"E); 151094 Mt Superbus SF, via
Warwick, (28°13 S, 152?28' E); J51637-40 O'Reilly's,
Lamington NP (28*]4' S, |53*08' E); J23937 near Mt
Ballow, (28716'S, 152*37' E); 18074, 18075 (allzann
stained), J8099 Mt Clunic, via Boonah (28"18'5,

agement and the Centre for Conservation Biol-
ogy, University of Queensland; the Queensland
Museum: Ross Sadlier (Australian Museum),
Harry Hines, Guy Hodgson (NSW National Parks
and Wildlife Service), Noel Cortinas, Kate
Couper, Amy Couper, Jenny Faulkner, Janie Girl-
ing. Marcia Lara, Mary Mulcahy, Simon and
Jenny Ormsby Jeff Wright, Steve Wilson and
Lauren Keim, who assisted us in the preparation
of this work,

LITERATURE CITED

BAUER. A.M. 1990. Phylogenetic systematics and bio-
geography of the Carphodactylini (Reptilia:
Gekkonidae). Bonner Zoologische
Monographien 30: 1-217,

COUPER, P.J., COVACEVICH, J.A. & MORITZ, C.
1995. A review of the leaf-tailed geckos endemic
to eastem Australia: a new genus, four new spe-
cies, and other new data. Memoirs of the Queens-
land Museum 34(1); 95-124.

COUPER PJ., COVACEVICH, J.A. & MORITZ, C.
1994, Designation of the type species of
Saltuarius, and other data on the genus. Memoirs

of the Queensland Museum 35(1): 26.

COVACEVICH, J. 1975. A review of the genus Phy-
llurws (Lacertilia: Gekkonidae). Memoirs of the
Queensland Museum. 17(2): 293-303.

HARSLETT, I. & ROYLE, M. 1980. They came to the
plateau (International Colour Productions:
Stanthorpe, Qld) 213pp.

152?32"E); J1143 Tweed R. (28*]8'S, 153727 E)
NSW; J5757 Chillingham, Murwillumbah (28°19'S,
153"17'E) NSW; 110565 Mt Lindesay (28?23'S,
152*43'E) SEQ: 54846 Bray's Ck, Border Ranges NP
(28°24'S, 153?03 E) NSW; J9054 Bulahdelah, 96km
NE Newcastle (32°25'S, 152*12" E) NSW,

APPENDIX 2.

Specimens of uncertain status from granite habitats
in NSW, QMJ53984, Teapot Ck, Narrow Pass Fire
Trail, Mann River Nature Reserve (29°45°S,
152?02'E) NSW; AMR141964, Chaelundi SF, (30°01'
07"S, 152?30'02"E) NSW; AMR141965, Chaelundi
SF, Sundew Lookout (30°03°04"S, 152?21'36"E)
NSW; AMR43870, 35km E. of Guyra (30"15'5,
152*00'E) NSW; AMRI23490, Tullawudjah CK,
NSW; AMR149768, Black Ck, 6. Ikm SE along Black
Hole Trail, Curramore SF (29*30'30"S, 152°11'24"B)
NSW.

NEW SPECIES OF RHIZOPINE CRABS (CRUSTACEA: BRACHYURA)
FROM NORTHERN AUSTRALIA

PETER J E DAVIE AND ANDREW HUMPHERYS

Davie, P.1.F. & Humpherys A, 1997 06 30: New species of rhizopine crabs (Crustacea:
Decapoda: Pilumnidae) from northern Australia. Memoirs of the Queensland Museum 42(1):

97-103. Brisbane. ISSN 0079-8835.

Two new species of rhizopine crabs are described from Australia. Cryptolutea arafurensis
sp. nov. is described from Darwin and the Gulf of Carpentaria. It is distinguished by the
degree of carapace granulation and the prominent anterolateral teeth. Heteropilumnus
longisetum sp. nov. is only known from the North-West Shelf. It is related to other
Heteropilumnus species with a coat of long fine setae on their carapace, legs and claws. C]
Brachyura, Pilumnidae, Rhizopinae, Cryptolutea, Heteropilumnus, Australia.

P.J.F Davie & A. Humpherys, Queensland Museum, P.O Box 300, South Brisbane, Queens-

land 4 101, Australia; 3 April 1997.

Scientific cruises conducted by the Common-
wealth Scientific and Industrial Research Or-
ganisation (CSIRO) in northern Australian waters
in recent years have produced a large number of
brachyuran specimens. Among the material de-
posited in the Queensland Museum from these
cruises are several species of pilumnid crabs of
the subfamily Rhizopinae, Two of these species
are considered new to science and are treated in
this paper.

In the most recent revision of rhizopine taxon-
omy, Ng (1987) has redefined the concept of the
Rhizopinae and removed it from the
Goneplacidae and placed it in the Pilumnidae. A
number of genera formerly considered rhizopine
have been excluded, and Heteropilumnus,
Pseudolitochira, Luteocarcinus, Rhizopoides and
Zehntneria have been added to the group (see also
Ng (1990) and Ng & Davie (1991)). It is in the
context of this revised concept of the Rhizopinae
that the new specimens from northern Australia
are treated.

The Rhizopinae hàve been poorly documented
in Australian waters and many specimens have
remained unidentified or unrecorded. While this
preliminary paper documents two new species, à
later paper will attempt to fully document the
Australian rhizopine fauna.

Specimens have been deposited in the Queens-
land Museum (QM) or the Northem Territory
Museum (NTM). All measurements are in
millimetres (mm) and are of maximum carapace
width followed by length, unless otherwise
stated.

SYSTEMATICS

Family PILUMNIDAE Ortmann, 1893
Subfamily RHIZOPINAE Miers, 1886

Cryptolutea arafurensis sp. nov.
(Figs 1. 2)

MATERIAL EXAMINED. HOLOTYPE: NTM
Cr001279, å (16.7 x 13.4mm), Ludmilla Ck., Darwin,
in mangroves, at low water, J. Hanley, 26.2.1982.
PARATYPES: NTM Cr001279, å (13.7 x 10.8mm).
CSIRO Torres Strait, 7.3.1989. QMW21401, 3 d d
(18.0 x 14.3.19.5 x 15.6, 17.2 x 14.1mm), 2 9 2 (16,0
x 12.6, 16.9 x 13.3mm), 13°02'S, 13922.2 E, Gulf of
Carpentaria, 58m, dredged, CSIRO F.R.V. Sher
Surveyor, 24.11.1991. QMW21402, å (13,0 x
10.7mm), 2 ? 9 (16.6 x 13.2, 13.1 x 10.7mm), data as
forQMW21401. QMW21403,7 d å (165x 13,2, 16.6
x 13,5, 15.1 x 12.0, 17.2 x 13.7, 13.7 x 11,2, 15.5 x
12.3, 14.7 x 1 i. 5mm), 12°10.5'S, 139"56.7'E, Gulf of
Carpentaria, 59m, dredged, CSIRO F.R.V. Southern
Surveyor, 24.11.1991, QMW21400, d (14.2 x
11.7mm), 2 9 (15.5x 12.4, 16.8x 13.6mm), 13°25.6'S.
138'36' E, dredged 54m, CSIRO F.R.V. Southern Sur-
veyor, 24.11.1991.

DESCRIPTION. Carapace subquadrate, c. 1.22
times broader than long, somewhat vaulted ante-
riorly. Dense covering of tomentum present on
entire dorsal surface. Carapace surface minutely
punctate in central regions; minutely granular
towards. lateral and posterior margins or nearly
smooth in some specimens. Dense scattered tufts
of tomentum, of varying density, particularly in
anterior third and towards the lateral margins.
Carapace regions relatively poorly defined; over-
lying fine pubescence obscuring surface detail in
some specimens. Metagastric grooves well indi-
cated together with forked frontal groove. Sub-

98 MEMOIRS OF THE QUEENSLAND MUSEUM

GMBREMNER 20

0.5mm

FIG. 1, Cryptolutea arafurensis sp. nov., holotype, NTM Cr001279, å (16.7 x 13.4mm). A, frontal view; B,
frontal and anterolateral margins; C, 3rd maxilliped; D, sternum; E, F, 1st gonopod; G, abdomen; H, coxal plate

of pereiopod 5; I, 2nd gonopod.

hepatic lobes well marked. Front deflexed, fron-
tal margin bilobed, a median cleft with groove
present; margins fringed with fine setae. Orbits
small and shallow but. visible from above. In-
fraorbital margin entire, with internal angle
somewhat acute. Ocular peduncles short, bul-
bous, immoveable; cornea darkly pigmented and
terminal. Antennules relatively short, folding into
oblique fossae. Antennae with basal joint short,
flagellum in contact with orbits. Epistome of

moderate length, shallow. Anterolateral margins
convex, lobes partially obscured by fringing to-
mentum; exorbital angle rounded, first anterolat-
eral lobe obtuse, scarcely indicated by shallow
notch; following two lobes much more acute and
separated by fairly wide U-shaped notches.
Posterolateral margins sub-parallel. Subhepatic
and pterygostomial regions smooth, but with a
few clumps of setae.

NEW SPECIES OF RHIZOPINE CRABS 99

Chelipeds subequal, somewhat robust; right
cheliped usually larger than left; merus compact
and high, trigonal; upper and lower margins fringed
with fine setae; outer surface minurel y granular;
acute tooth present on upper margin. Corpus sub-
ovate, outer and upper surface granulate with
scattered, long setae; inner surface smooth; prom-
inent obtuse tooth present on interior margin;
outer surface covered in fine setae. Palm broad,
high and flat, upper and lower margins somewhat
keeled; outer surface with conspicuous rows of
tuberculate granules, more closely aggregated to-
wards upper margin and sometimes sparse or
nearly absent in some specimens; woolly tomen-
tum overlying portions of upper and lower mar-
gins and adjoining areas. Index straight or slightly
deflexed, with median carina, distal 1/3 usually
smooth; lower margin fringed with fine setac.
Gape with strong dentition, Dactylus
downcurved, with a comb of setae on upper prox-
imal margin, distal end without setae, Hooked
tips of chelipeds overlap when closed, Distal 2/3
of index finger and dactylus dark sepia in colour
(afler preservation).

Ambulatory legs of moderate length, relatively
stout and fringed with fine setae, Dactyli styli-
form, heavily setose. Large projecting serrate
plate present on upper surface of coxae of all 4
pairs.

Buccal cavern sub-quadrate, Third maxillipeds
incompletely close buccal cavern, Ischium quad-
rangular, outer surface smooth. Merus sub-quad-
rate, smaller than ischium; anteroexternal angle
strongly produced.

Thoracic sternum punctate. d abdomen evenly
tapered and heavily tomentose; with 7 distinct
segments; segment 1 covering entire interspace
between last pair of ambulatories; segment 2
much shorter; segment 3 longer again, with some-
what acute lateral angles; segments 4-6 with con-
cave margins, telson triangular with rounded
mex.

Male genital pore coxal, exposed penis lying in
channel between 4th and Sth thoracic sternites.
Male gonopod sigmoid-shaped, with hooked la-
pering terminal portion,

DISTRIBUTION. Darwin, N.T. and the Gulf of
Carpentaria; from low-water to 59m depth.

ETYMOLOGY. Name refers to the Arafura Sea.

REMARKS, This relatively large species is the
5th in the genus Cryprolutea to be described. This
genus is characterised by the presence of à serrate
plate on the coxae of the ambulatory legs.

Cryptolutea erafurensis sp. nov, is relatively
common in the samples from the Gulf of Car-
penlaria, being the dominant! species of
Rhizopinae found. It is, however, highly variable
in the degree ol granulation on the carapace and
cheliped palm, ånd in the extent of setation, Sev-
eral specimens exhibited a markedly pubescent
carapace whilst others were almost devoid of
setae. Similarly, granulation in varying degrec
was present on the carapace of most specimens,
but completely absent on some.

Cryptolutea arafurensis sp. nov. is distinguish
able from its congeners by the prominent antero-
lateral teeth, and by the degree of carapace
granulation typically present. In C. lindemanen-
sis Ward, 1936, the carapace is relatively smooth
and the anterolateral margins have less prominent
teeth; the anjerolateral lobes of C. reschi (Serene,
1971) are more tuberculate and less clearly sepa-
rated as teeth than those of C. arafurensis. In
addition the carapace of C. teschi appears more
areolate or rugose than the present species. C.
granulosa (MacGilchrist, 1905) ts more evenly
granulose, particularly on the outer surface of the
palm, than the present species, and the dense
tomentum on the dorsum 15 absent,

C. sagamiensis (Sakai, 1935) appears closest to
C. arafurensis but it can be distinguished by the
following characters: 1) the anterolateral teeth are
more prominent, with the last 2 lobes narrower
and showing à tendency to unite with each other;
compared with the last 2 lobes being prominent,
somewhat acute, and clearly separated from each
other in C. arafurensis, 2) the larger number of
granules on the outer surface of the cheliped
palm, 3) in C. sagamiensis the merus and carpus
of the chelipeds are covered in rudimentary gran-
ules, whereas in C. argfurensis the carpus, in
particular, is covered in prominent vesiculous
granules, 4) according to the text-fig. 16a) of
Sakai (1935), the merus and ischium of the 3rd
maxillipeds are more rounded in C, sagamiensis
than in C, arafurensis.

Heteropilumnus longisetum sp. nov.
(Fig. 3)

MATERIAL EXAMINED. HOLOTYPE;
OMW 21423, d (7.5 X 5.3mm), North West Shelf, W,
Australia, 19'29,6'S, 11853,2'E, trawled 37-3801,
25.10.1933, T. Ward (CSIRO), PARATYPE,
QMW14449, då (5,6 x 4.0mm). North West Shelf,
02BéBT, 81-82m,

DESCRIPTION OF HOLOTYPE. Carapace 1.4
times broader than long: more or less Bal across

100

ØKOEG E

PETTTTTFTTTETTTTLTTTTTTTTTTET TÅ

MEMOIRS OF THE QUEENSLAND MUSEUM

ERTA LEFI T TETT

FIG. 2. Cryptolutea arafurensis sp. nov., holotype, NTM Cr001279, å (16.7 x 13.4mm). A, frontal view of

chelipeds; B, dorsal view. Scale lines in mm.

the mesogastric and branchial regions although
becoming convex laterally; convex fore and aft
over frontal third. Entire surface covered in fine
silky setae; moderately long posteriorly, becom-
ing much longer anteriorly; longest setae form
transverse fringe across frontal region, and are
about 1/3 or more length of carapace. Regions

relatively poorly defined, with only
meta/mesogastric region being moderately dis-
tinct. Anterior to frontal fringe surface with only
very tiny setae; front deflexed, bilobed; most
prominent medially; sinuous with obtuse, sub-
acute preorbital angle. Supraorbital margin sinu-
ous, microscopically granular, without fissures.

NEW SPECIES OF RHIZOPINE CRABS

Infraorbilal margin also minutely granular;
slighily raised below proximal margin of cornea;
inner margin without enlarged tooth or lobe, inner
angle rounded, meeting distal margin of busal
antennal joint. Antenna with basal segment just
touching front; flagellum about as long as frontal
setae; each segment armed with 2 ar 3 short setae
distally; terminal segment with few very long
setae distally which make antennae appear much
longer. Anterolateral margin consisting of 4 teeth,
including exorbital angle. Exorbital angle small,
sub-acute, with few small accessory granules on
outer margin; 2nd tooth of similar size, leading
Margin very short, trailing margin long and with
few microscopic granules; 3rd tooth most prom-
inent, acute; 4th tooth smallest. clearly defined
but litle bigger than large acute granule. Greatest
carapace width between 3rd anterolateral teeth.
Posterolateral margin longer than anterolateral,
straight, converging posteriorly, Posterior margin
straight, with slightly raised rim.

Third maxilliped with merus c. 1.25 times
wider than long and 2/3 as long as ischium; distal
margin concave; ischium about as long as wide,
inner margin produced, broadly convex and
slightly crenulated,

Stemum and abdomen entirely covered with
moderately long setae; d abdomen reaching
about half distance across fused 3rd and 4th tho-
racic sternites. d abdomen as figured: telson with
convex margins reaching à blunt point, longer
than other segments and c. 1.7 times length of
penultimåte segment; segments I and 3 widest.
similar in width, occupying entire space between
coxae of last pair of legs; 2nd segmente, 0.7 times
width of 3rd. å Ist pleopod sigmoid; apical beak
relatively long and acutely pointed, projecting
laterally rather than recurved; armed with series
of fine bristles on inside edge of distal curve.

Chelipeds subequal, right slightly more
swollen; merus short, trihedral, unarmed except
for row of sharp granules on upper inner border,
and few smaller granules on ventral anterior mar-
gin. Carpus unarmed, inner angle bluntly pointed,
covered in fine setae which are short proximally
but very long distally; surface smooth beneath
setae, without obvious sculpturing. Palm exclud-
ing fixed finger c. 1.25 times longer than high:
fingers relatively long, fixed finger c. 1.1 limes
height of palm, markedly downturned, particu-
larly on left cheliped; both fingers with tips
pointed, margins wilh about 5 large, triangular,
sharply crested teeth; dark colouring on teeth
entire distally then extending obliquely bavk-
wards. towards base of gape. Entire outer face ul

m

Im covered in very long, fine, soll setae; sur-
ace scattered with granules difficult to see except
along outer margins of fingers where they be-
come more conspicuous, especially on Jeri chela.
Walking legs missing.

Variation. The smaller paratype male shows some
significant variations from the much larger holo-
type, andis in possession of several walking legs.
The carapace anterolateral angles are cut into
much more prominent, triangular teeth, and the
outer margin of the 2nd tooth appears compara-
üvely shorter. The dark colouring on the fingers
of the chelae is restricted to the distal ends, und
does not extend backwards along the gape. The
tip of the 1st pleopod is not as long and acute.
Walking legs moderately long and slender; length
of longest c. 1.6 times width of carapace, width
of merus c. 0.3 umes length; dactylus longer than
propodus, tip curved and acute. Propodi, carpi
and dactyli bear very long silken setae, particu-
larly on outer borders. All segments unarmed
except for a few sharp, irregular granules on the
upper margin of the meri,

DISTRIBUTION. Only known from the North-
West Shelf, Western Australia; in 37-82 1 depth.

ETYMOLOGY, Named in reference to the long setae
covering the carapace, legs, and claws, [tis used us i
noun in appesipion.

REMARKS. Hereropilumnus isa relatively large
(18 recognised valid species) and taxonomically
confused genus that is in need of å full revision,
Despite this there are only a restricted number of
species within Hererapilumnus that resemble H,
longiserum sp. nov. in bearing a coat of long fine
setae on their carapace, legs and claws. The clos-
est in appearance to H. Jongiserum sp. nov. are H.
ciliatus (Stimpson, 1858), H. lanuginosus
(Klunzinger, 1913), H, splendidus (de Man,
1929) and H. rrichøphoroides (de Man, 1895].
Comparing our new species with Ihe figures of
Heteropilumnus ciliatus given by Shen (1936:
text-lig. 2 [as H. cristadentatys Shen, 1936]) and
Sakai (1976: pl. 176, fig. 3) Heteropilumnus
longisetum differs from H. ciliatus by the much
lower, less prominent, anterolateral teeth; the
more extensive covering of longer setae particu-
larly over the posterior half of the carapace, and
the outer distal margin of the meris of the 3rd
maxilliped being not produced.

It differs from the original figures of H.
lanuginosus (Klunzinger, 1913: pl. 2, lig. 18; pl.
7, fig 4) by lacking the broad looth on the initer
margin of the infraorbital margin, H splendidus

102 MEMOIRS OF THE QUEENSLAND MUSEUM

EN N Y

3 e

A N W \ d | I) d
(Wl (ities e D d
RA

10
kb Ai il j
Va |

de

Wi NNK

Imm

AR d
y
Za
A li, |
m
D
På

Å IDE

Ca A WA ^;

mm P tutt,
MN dA

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BARE

Imm /

Imm

GMBREMNER 8

FIG. 3. Heteropilumnus longisetum sp. nov., holotype, QMW21423, d (7.5 x 5.3mm). A, dorsal view of carapace;
B, C, Ist gonopod; D, 3rd maxilliped; E, left chela; F, frontal view of orbit and antennae; G, abdomen; H
sternum.

NEW SPECIES OF RHIZOPINE CRABS

can be easily separated because of its lack of
clearly defined anterolateral teeth (see de Man
(1929: pl. 1, fig 3-3c; pl. 2, fig. 3d-f). Finally it
differs from H. trichophoroides (cf. figures of de
Man (1895: 13, fig. 8a-e) and Rathbun (1910: fig.
40, pl. 1, fig. 8 [as Pilumnus borradailei Rathbun,
1909]) because the posterolateral margins are less
convergent; the frontal margin is less produced;
the regions on the carapace are much less defined;
and the setae of the carapace are much longer.

ACKNOWLEDGEMENTS

We are grateful to Dr A.J. Bruce and Karen
Coombes of the Northern Territory Museum, and
to Trevor Ward of C.S.LR.O., for the sending
study specimens; Clare Bremner drew the excel-
lent figures. The work could not have been done
without the financial support of the Australian
Biological Resources Study to P. Davie for the
study of Australian xanthoid crabs.

LITERATURE CITED

KLUNZINGER, C.B, 1913. Die Rundkrabben
(Cyclometopa) des Roten. Meeres. Nova Acta
Academiae Caesarea Leopoldino-Carolinae
Germanicum Naturae Curiosorum 99(2): 103-
402, figs 1-14, pås 5-11 [1-7].

MACGILCHRIST, A.C. 1905. Natural History Notes
from the R.I.M.S.S. “Investigator”, Capt, T.H.
Heming, R.N, (retired), commanding. Series IL,
No. 6. An account of the new and some of the rarer
Decapod Crustacea obtained during the surveying
seasons 1901-1904. Annals and Magazine of Nat-
ural History 15(87): 233-68.

MAN, J.G. de 1895. Bericht über die von herrn
Schiffscapitàn Storm zu Atjeh, an den westlichen
Küsten von Malakka, Borneo und Celebes sowie
in der Java-See gesammelten Decapoden und
Stomatopoden (Part 1), Zoologische Jahrbücher,
Jena. Abteilung für Systematik 8 (Brachyura -
Daclea to Cleistostoma): 485-609, figs 1-15.

1929, On a Collection of Decapod and Stomatopod
Crustacea from Pulau Berhala, an Islet situated in

the Straits of Malacca. Bijdragen tot de
Dierkunde. Amsterdam & Leiden 26: 1-26, pls
1-3.

NG, PK.L, 1987. The Indo-Pacific Pilumnidae II. A
revision of the genus Rhizopa Stimpson, 1858,
and the status of the Rhizopinae Stimpson, 1858
(Crustacea, Decapoda, Brachyura). Indo-Malayan
Zoology 4: 69-111, figs 1, 2, pL J, table 1.

1990. Luteocarcinus sordidus, new genus and spe-
cies, from mangrove swamps in peninsular Ma-
laysia (Crustacea: Decapoda: Brachyura:
Pilumnidae: Rhizopinae). Proceedings of the Bi-
ological Society of Washington 103(1); 95-9,

NG, P.K.L. & DAVIE, EJ.F. 1991. The Indo-Pacitic
Pilumnidae VIL Notes on Heteropilumnus
sasekumari (Seréne, 1971) and Cryptolutea Ward,
1936 (Crustacea: Decapoda: Brachyura). Mem-
oirs of the Queensland Museum 30(3): 517-24.

RATHBUN, M.J. 1909, New crabs from the Gulf of
Siam. Proceedings of the Biological Society of
Washington 22: 107-14,

1910, The Danish Expedition to Siam 1899-1900.
V. Brachyura. Kongelige Danske
Videnskabernes. Selskabs Skrifter. Kyobenhavn
7(4); 301-68 (1-68), text-figs 1-44, pls 1-2, 1

map.

SAKAI, T. 1935, New or Rare Species of Brachyura,
collected by the "Misago" during the zoological
survey åround the [zu-Peninsula. Science Reports
of the Tokyo Bunrika Daigaku. sect. B, 2(32):
63-88, figs 1-17, pls 6-8.

SERENE, R. 1971, Observations préliminaires sur des
Brachyoures nouveaux ou mal connus du Sud-Est
asiatique (Crustacea Decapoda). Bulletin. du
Muséum national d'Histoire Naturelle, Paris Ser.
2, 42(5 [1970 (19711: 903-18, pls 1-6.

STIMPSON, W. 1858. Prodromus descriptionis animal-
ium evertebratorum, quae in Ex peditiane ad Oce-
anum Pacificum Septentrionalem, å Republica
Federata missa, Cadwaladaro Ringgold et
Johanne Rodgers Ducibus, observavit et descripsit
W. Stimpson. Pars. V. Crustacea Ocypodoidea.
Proceedings of the Academy of Natural Sciences
of Philadelphia 10; 93-110 (39-56).

WARD, M. 1936. Crustacea Brachyura from the coasts
of Queensland. Memoirs of the Queensland Mu-
seum XI(1): 1-13, 3 (ss.

104

COMBAT AND COPULATION IN OXYURANUS
MICROLEPIDOTUS (ELAPIDAE). Memoirs of the
Queensland Museum 42(1); 104. 1997;- Shine & Covacevich
(1983) reviewed the ecology of Oxyurarus spp., largely from
examination of museum specimens. However, while they
were able to report body size and aspects of reproduenon and
feeding, combat berween males and copulation were not
described, Male vs male combat in Australian elapids is well
known (Shine, 1991). It has been reported most recently (in
Demansia vestigiata) by Covacevich et al, 1994. Such
behaviour occurs in taxa characterised by male size superior-
ity and js believed to be a strategy for forcing rival males away
from a receptive female, defending against homosexual court-
ship and obtaining food ahead of a rival (Shine et al, 1981),
For Oxyuranus spp. accounts of combat/copulation are
scant, Combat between å å O. scutellatusin the wild has been
photographed (Shine, 1991) ‘during the mating season’, and
rape. has been observed in August and early December
(Shine & Covacevich. 1983). Neither has been described
previously in Q. microlepidotus. In July-September, 1996
oth combat and copulation were observed in wild specimens
of O, microlepidotus In Astrebla National Park, 26km SE of
No. 2 bore (at approximately 24"07' S 140*31'E), SWQ.
Combat å 3. Two specimens were encountered at approxi-
mately 17:22 hours. on 2 September. 1996 (Fig. 1; OM
transparencies NR386, NR387, NR388). The snakes were
fully entwined, ‘plaited’, as reported in black snakes by Shine
(1978). Both spogimans had their heads and approximately
40cm of their bodies raised from the ground and were fre-
quently and alwenately lashing outat each other. Their mouths
were not and did not open. The snakes often came together
with one head ir: on the other. Their bodies were tightly
coiled. What could be termed 'head-raising behaviour" fre-
quently resulted in their falling suddenly to the ground, twist-
ing and repeatedly rolling over one another. This behaviour
continued for 30 minutes, when the snakes separated and
moved Slowly away, One went into a Bilby (Macraris lagotis)
hurrow nearby. The other followed the first to the burrow
entrance, apparently searching. This second specimen then
enicred the same burrow, After 20 minutes one specimen
surfaced from a different entrance, approximately 2m from
the first, The other re-emerged from the initial burrow en-
tance and å similar ritual of entanglement and head-raising
ensued for five minutes, Both specimens then entered the first
burrow. About one minute later, the head of one specimen
appeared from the second entrance, The specimen emerged
front the burrow and moved across the plain rapidly, This
occurred at 18: 10 hours as the last 20-30em of the other snake
was "disappeanng' down the first entrance, A (at time there
as little Light and observations ceased.
Copulation & 9. On 6 August, 1996, two specimens were
observed 2m from a Bilby burrow at 13:30 hours, One (pre-
sumed d was ‘draped’ in a series of coils over the sides of
the other (presumed 9). which was motionless, They were
“joined” genitally (Fig. 2; QM transparency NR389). The
(presumed) å frequently was contracting its body, giving the

FIG. L Male Gay ranis mocrolepiduiws combi. (P, fMeRae)

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 2.0, microlepidotus in copulation. (P. McRae)

impression that it was attempting to maintain its "superior"
position. Infrequently, every 5-10 minutes, the body of the
(presumed)3 contracted in a series of ‘quivers’, also along
the length of the body. After 10-15 seconds these culminated
in a short, sharp series of (presumed) orgasmic contractions.
This behaviour continued for about I hr. Then the specimens
moved slowly to and down the Bilby burrow, Two to three
seconds later, one specimen emerged from the burrow and
moved quickly to the mating site. It then returned to the
burrow. This occurred twice during the next 10 minutes. On
the last occasion, the specimen remained in the burrow, Both
specimens were below ground when observations ceased nr
1500 hours.

These observations both concur with and vary slightly
from previous reports of other elapid species, They conform
with an observation (Shine, 1978) that elapids which are
heavy and museolar hold their bodies upright dunng å dá
combat. That d å combat occurs in O. microlepidotus may
be at variance with the observation that such behaviour is
known only where 3 d are larger than ? ? (Shine, 1991). The
difference between the body sizes of dd vs 99 O
microlepidotus has not been shown to be statistically signili-
vant (Shine & Covacevich, 1983). However, the sample
(d 813, 7 96) was small. Copulation by O. microlepidotus
in the wild in August is consistent with the timing of this
behaviour in captive specimens.

Literature Cited

Covacevich, J.A., Roberts, L. E McKinna, L, 1994, Male
combat in the black whip snake, Demansia vestigiata
Mernoirs of the Queensland Museum 3701). 52,

Shine, R. 1978. Sexual size dimorphism and male combat in
snakes. Oecologia 33: 269-278.

1991. ‘Australian snakes. A natural history". (Reed Books

Pry Ltd: Sydney). 223pp

Shine, RH. E Covacevich, J. 1983, Ecology of highly venomous
snakes; the Australian genus Oxyuranus (Elapidae),
Journal of Herpetology 17: 60-90

Shine, R., Grigg, GC. Shine, TG & Harlow, P. 1981. Mating
and male combat in the Australian blacksnake,
Pseudechis porphyriacus (Serpentes, Elapidae). Jour:
nal of Herpetology 1501): 101-107,

P.D. McRae, Queensland Department of Environment, PO

Box 149, Charleville, Queensland 4470, Australia; JA. Ca

acevich, Queensland Museum, PO Box 3300, Soulli Brisbene,

Queensland 4 101. Australia; 7 April 1997,

TEMPORAL SEGREGATION OF HUMPBACK WHALES DURING MIGRATION IN

SOUTHERN HEMISPHERE WATERS *
W.H. DAWBIN

Dawbin, W.H. 1997 06 30: Temporal segregation of humpback whales during migration in
southern hemisphere waters. Memoirs of the Queensland Museum 42(1): 105-138. Brisbane.
ISSN 0079-8835.

All available data on post-war catches and some pre-war catches of humpback whales in the
southern hemisphere between 6698 and 198 have been examined for evidence that some age
or reproductive categories migrate north, and south early in the season, while others follow
in sequence later. The categories that showed significantly different time sequences between
groups in non-polar waters were as follows:- all immature humpbacks (no significant
difference between males and females), mature males (regardless of size), and mature females
subdivided according to reproductive stage as pregnant, lactating or resting. Based on the
average intervals between the mean dates for each category at whaling localities between
41°S (Cook Strait, New Zealand) and 1°S (Congo), lactating females accompanied by
weaning 'yearlings' migrate north earliest and are followed by immature humpbacks, mature
males, together with resting females and finally pregnant females at 12, 20, 23 & 31 days
later respectively. During the return migration southwards, mixed females (including those
in early pregnancy) occur first together with immature whales, and are followed by mature
males and females in early lactation 10 and 16 days later. Reasons for believing these time
intervals to be minimal are discussed. In general, humpbacks appear to return south in the
same order in which they travelled north, but some females change status so that those that
travel north early when near the end of lactation, may return south early as pregnant animals.
Others that travel north late as pregnant animals, return south late as cows accompanied by
young calves. Mature females, when pregnant, appear to spend a prolonged period in
Antarctic waters but when suckling a calf they spend a substantially reduced interval in cold
waters. Antarctic catches appear to have been taken over too short a season to demonstrate
sequences during the entry and exit of humpbacks to Antarctic waters. [ ] Humpback,
Megaptera , southern hemisphere, migration, segregation.

W.H. Dawbin, School of Biological Sciences, University of Sydney, Sydney, New South Wales
2006 . Present address: Australian Museum, 6-8 College Street, Sydney, New South Wales,
2000, Australia; 23 December 1996.

The published information on the main features
of humpback whale migrations between warm
water breeding regions and cold water feeding
regions have been described and reviewed by a
number of workers. These are referred to by
Dawbin (1966), so the present account will be
confined to a discussion of segregation during the
migratory cycle of humpbacks.

Ever since the International Whaling Statistics
began publishing tables which classified the sex
and sizes of whales caught in all main whaling
grounds, it has been clear that there is a marked
disparity in the composition of humpback catches
from Antarctic regions as compared with those
from temperate and tropical regions. While fe-
males usually predominate in Antarctic catches,
males tend to predominate in catches from
warmer waters. This change of composition sug-
gests that the humpback population does not re-
main homogeneous throughout the season, but
undergoes some degree of segregation into sex

and possibly age groups at certain phases of its
seasonal movements. Mackintosh (1942) has dis-
cussed evidence of changes in the composition of
humpback catches (inter alia) during the Antarc-
tic season, but accounts of sequential changes in
composition in other southern hemisphere re-
gions have tended to describe movements in
rather localised areas.

Segregation between whale categories may
Occur in space (geographical segregation) or in
time. Geographical segregation is here regarded
as theentry of one sex, age or other specified class
within a species, to an area which is not entered
by other classes of that species. The classes are
then clearly separated in space. In time segrega-
tion, one or more classes enter an area at an earlier
date than the other classes, but the latter enter the
same area at a later date. At any one moment the
classes may be separated in space but they all
pass through the same area although at different
times.

* This work was completed in 1963, the year in which the International Whaling Commission banned the taking

of southern hemisphere humpback whales.

106

Sperm whales provide an excellent example of
geographical segregation. Mature males only are
found at high latitudes while females and imma-
ture animals do not occur south of about 45°S.
There are differences in sex ratios and in the
proportion of mature and immature humpbacks
represented in catches at different latitudes, but
there are no known cases of such marked geo-
graphical segregation between categories of
humpbacks as occurs with sperm whales. How-
ever, at many localities there are regular differ-
ences in the catch composition with change in
season.

It is suggested that many of these cases may be
the result of some humpback categories migrat-
ing to warm waters early in the season while
others tavel at varying intervals later. Any such
tendency would change the relative proportions
of those remaining after the start of migration as
well as cause sequential changes in the composi-
tion of the stock migrating past any fixed point.

The object of the present study is to identify the
reproductive stage of each humpback and relate
this to the arrival time of each animal at specified
latitudes during the migratory cycle. It is an at-
tempt to specify the reproductive or age catego-
ries that tend to migrate earliest in the season and
to determine the time interval between that of
their peak density and that of categories which
migrate at a later date. As part of time segregation,
the order in which specified categories travel and
the mean interval between categories is referred
to in this account as *time sequence'. It is this as
distinct from geographical segregation that is the
main subject of the present study.

MATERIAL AND METHODS

With the exception of sighting records on cows
accompanied by calves, humpbacks cannot be
classified accurately into size, sex, or reproduc-
tive stage categories by observations at sea.
Catch data on animals processed at whaling sta-
tions or factory ships have been published regu-
larly in International Whaling Statistics, but the
data are grouped into monthly categories or into
size categories unrelated to dates. The object of
the present study has therefore necessitated an
analysis ofthe available data on individual hump-
backs for those cases that specify the length, sex,
stage of pregnancy, geographical position and
date of catch.

MATERIAL EXAMINED. Data have been obtained
for each of 65,600 humpbacks caught in the southern
hemisphere. Of these 38,100 were caught in post-war

MEMOIRS OF THE QUEENSLAND MUSEUM

years and 27,500 were caught in pre-war years. Both
samples include catches from Antarctic through to
equatorial waters.

From temperate and tropical waters the post-war data
relate to 28,243 humpbacks as follows:- Cook Strait,
New Zealand 1922; Great Barrier Island, New Zealand
264; Tonga 84; (data and gonads collected by the writer
and field collaborators); Albany, West Australia 699;
Carnarvon, West Australia 5889; Point Cloates, West
Australia 3321; Byron Bay, New South Wales 1040;
Moreton Island, Queensland 6206; Norfolk Island 880;
Durban, Natal 854; Madagascar, pelagic 2014; Congo
4071; (data provided on microfilm by Mr E. Vangstein,
Sandefjord with the permission of the managers of the
respective whaling companies. Valuable supplemen-
tary data on Australian catches was provided by Dr
R.G. Chittleborough). Pre-war data includes 14,515
humpbacks as follows:- Coast of Natal 2621; Mada-
gascar 1223; Western Australia land station 3426;
Western Australia, pelagic 7244. Professor J.T. Ruud
provided the pre-war records together with summaris-
ing tables and graphs prepared by himself and collab-
orators.

Antarctic catches in post-war years include 9857
humpbacks taken in late December and early February.
In the 1932/33 to 1938/39 seasons, catching in Antarc-
tic waters occurred from early Novemberto late March,
and data on 12,985 humpbacks caught between 65°S
and 66°S have been obtained from Mr E. Vangstein
who provided duplicate I.B.M. cards for each animal.
Shore whaling in subantarctic waters has, in recent
years, been confined to South Georgia where hump-
backs have formed a negligible part of the catch. There
were substantial catches earlier from South Georgia
and other coastal waters of the Falkland Islands De-
pendencies and records exist for those caught between
1910 and 1917. These data are lodged in the British
Museum (Nat. Hist.) where Dr F.C. Fraser kindly al-
lowed me to examine them. South American hump-
back catches have been small and secondary to those
of other species, so they have not been included in the
present study.

METHODS. Gonads and other material and data
obtained in New Zealand and Tonga were suffi-
cient to classify the local catches in a number of
ways that could be tested for statistically signifi-
cant differences. Subdivisions into sex, and size
categories showed little difference in the date of
peak density of small males compared with small
females, but there were substantial and inconsis-
tent differences between the dates of peak density
of large males as compared with large females.
Classification based on reproductive data showed
little or no difference between the time sequence
of immature males and immature females, but
suggested that the timing of mature females could
be related more closely to their stage of breeding
than to their size. Females at the end of lactation,
resting females and pregnant animals followed

TEMPORAL SEGREGATION OF HUMPBACK WHALES

different time sequences of which that followed
by resting females was most similar to the time
sequence of mature males (Dawbin, 1960).

Except for the specified pregnant females, data
on the reproductive condition of humpbacks are
listed rarely in catch returns. In New Zealand
catches the ovaries. from 611 females and testes
weights of 630 males indicated that females be-
came mature at 39'6" and males al 38'. Those
from which gonads had not been obtained were
therefore classified as mature or immature ac-
cording lo whether they were above or below
these lengths. Mature females whose mammary
glands gave evidence of recent activity and those
recorded as accompanied by a ‘yearling’ were
classified as being at the end of lactation, Preg-
nant females are easy 10 recognize during pro-
cessing as the foetuses of northbound humpbacks
are large at New Zealand latitudes, The remain-
ing mature females include resting animals and
almost certainly include some that were at the end
of lactation but lor which there is inadequate data
to allocate them to this category.

From the large Australian catches,
Chittleborough (1958) has examined numerous
gonads. and found a close correlation between
length and state of maturity using 36'9" as mean
length for maturation of males and 38'6" as mean
length for females, These lengths have therefore
been used in this study to classify Australian
catches and in view of the large samples on which
the criteria are based, they have also been applied
to data from all regions beyond Australasia. Be-
cause of the slight discrepancy between the esti-
mated maturation lengths in Australia and New
Zealand, data from other regions have also been
analysed on the New Zealand criteria and by
arbitrarily varying the lengths from 37 to 39 feet
for males and 38 to 40 feet for females. The
results did not produce significant changes in the
calculated time sequences so the original criteria
on maturity have been retained in the present
analysis.

Some separation of mature non-pregnant fe-
males into stages of lactation or resting has been
possible only for certain Australian catches, Dr
Chittleborough has kindly provided data from
four seasons' catches in Western Australia relat-
ing the proportion of mature females whose
mammary glands showed signs of recent activity
to that of other non-pregnant mature females. The
pre-war Western Australia catches made before
the minimum size length of 35 feet was estab-
lished. include many whales whose length indi-
cales thal they were almost certainly yeartings

107

travelling with mothers near or at the end of
lactation, In other regions non-pregnant mature
females have been classified as a mixed group
including resting and unspecified animals (shown
as R & U in tables) of which an unknown propor-
tion would be in late lactation.

With the above reservations, catch dala on cach
humpback was used to classify the animal as
immature (male or female); mature male: lactat-
ne. resting, pregnant or unspecified mature fe-
male.

Catches from relatively fixed positions (land
stations or coastal factory ships) were classified
as above ard then grouped for each category into
weeks for each season, As differences between
lime sequences per season at any one locality
proved non-significant, the data for successive
seasons were pooled for each week as shown in
the appended tables,

The weeks were specified starting from Ist
May which is the earliest commencing dale for
the most southern shore station under consider-
ation (Cook Strait, 4175). To facilitate compara-
ble analyses for each station, the weeks were
arbitrarily numbered commencing from May 1
and the same week code system has been used for
all non-polar whaling localities, ie, May 1-7,
May 8-12; May 13-21, 3; etc, The numbering for
Antarctic catches commences with November
6-12, Ly eic.

From the weekly frequencies of each hump-
back category per locality, the mean date, stan-
dard deviation and 95% confidence limits af the
mean were calculaled. The limits used were two
standard errors on either side of the mean for
samples exceeding 30 and Yates correction in the
few cases where samples have been smaller than
30. The results have been presented in the graph-
ical form (with slight modifications) described by
Hubbs and Hubbs (1953), In the present graphs
(Figs 23 & 25) the mean date is indicated by a
white line within the black area that represents the
95% confidence limits on either side. Range has
been omitted since this is determined frequently
by a tonc early or late whale and is therefore noi
relevant tò the main theme of this study,

As pointed out by Hubbs & Hubbs, consider-
able reliance can be placed on the significance of
the difference between samples, if the corre-
sponding rectangles (referring to the total range
of four standard errors of the mean) are only
slightly separated, or if the overlap is not more
than about 33% of the length of the shorter of the
two fectangles, An approximate idea of the sig-
nificance of an observed difference can be ob-

lus

lnined by inspection of Fig. 4 in Hubbs ani
Hubbs. In all doubtful cases, t tests were applied
ro determine whether or not a difference was
significant at the 5% level or less, Coded means
hive been reconverted to calendar dates in Tables
12, 14 & 19.

Antarctic pelagic catches have been subdivided
firstly into groups according to latitude and sec-
ondly into sectors of longitude subdivided into
latitudes before classification as above.

While the tables list the humpback groupings
used in the analyses, the graphs far clarity show
fortnightly groupings Io avoid some of the
weekly random fluctuations or the alternative ol
smoothing the data. Graphs of percentage con-
position per fortnight are included as a guide to
the probable composition of catches concentrated
into selected portions of the humpback season
and as å comparison with earlier dala which have
heen published in the form of percentage changes
during the whaling season. It must be emphasised
that such dara are of very doubtful value sf apphed
to the season as a whole. For example, Fig. 2
suggests a fairly continuous replacement of
immature numpbacks by mature animals
throughout the season, but Fig. I shows that this
is the result of å relatively small time difference
in the periods of peak density of immature com-
pared with mature whales. Other figures illustrate
many comparable cases,

The separation of northbound from southbound
humpbacks has, in the absence of individual data
on direction of travel, depended on sighting re-
cords of the proportion of all humpbacks travel-
ling north or south during the migratory season.
Sources of error in such data are discussed helow.,
Sampling Error. Catches are not completely ran-
dom samples of the populations from which they
are drawn, and the main sources of sampling error
are considered below,

a! Size. The minimum length of 35 feet speci-
fied by the International Whaling Commission in
1937 normally gives complete protection to the
‘yearling’ group that range from about 28 to 31
feet in length. Data on yearlings must therefore
be obtained from catches made prior to. 1937 or
from sightings at sea, Members of this group are
seen very frequently accompanying their mothers
in higher temperate latitudes, but they usually
become independent in the tropics.

A proportion of immature whales of other age
groups included in the 30 to 35 feet length range.
are also protected by the size regulation, bur those
between 35 feet and the size of sexual maturity
can be caught. However, they are likely lo be

MEMOIRS OF THE QUEENSLAND MUSEUM

under-represented due tu gunner selection for
whales of greater size. The degree to which this
pccurs has varied between stations and seasons
due to changes in relative availability of larger
whales, and the effects in some cases of operating
under quotas. Within the group of immature
whales which are caught, there is no known
means of distinguishing the sexes at sea, SO, as
females reach å greater size than males before
becoming sexually mature, they would be ex-
pected to form a higher proportion in the catch of
immature whales than that in which they occur in
the whole migraung population of immature
whales,

Mature females grow to a greater maximum
size than mature males and gunner selection for
the largest whales would he expected to increase
the relative representation of this group in the
catch compared with the population as å whole.
However, this effect 18 counterbalanced to vary-
ing degress by the next factor,

b) Presence of Calves. Cows accompanied by
suckling calves are given protection by the Inter-
national Whaling regulations. and this protection
is of greatest importance among southbound
humpbacks returning from the breeding grounds,
and among those in the Antarctic feeding grounds
where the calves are still suckling, It is less im-
portant in the temperate zones where northbound
calves are in the process of being weaned and are
almost certainly capable of continuing their exis-
tence independently of the mother, so the mother
is often hunted. However, some whalers have
actively avoided the mothers of these weaning
yearlings, and the difference in policy relative 10
this group has provided some marked differences
in catch composition between stations.

c) Pregnant Whales, It is generally assumed thal
pregnant females cannot be identified as such at
sea and no regulations relating to this class of
whales have heen made. However, the proportion
of pregnant whales changes during the northward
migration because of the birth of calves at various
latitudes en route. Towards the end of northward
movement a high proportion of females which
were in late pregnancy when they left the Antare-
tic seas, have undergone partutition becoming
recognizable as cows accompanied by young
calves and they then receive full protection. By
the time the southward migration commences all
Whales Which had left the Antarctic in late preg-
nancy have delivered calves and, except for those
which have lost their calves, this whole group 1s
protected during their return migration and sub-
sequent slay m Antarctic waters.

TEMPORAL SEGREGATION OF HUMPBACK WHALES

There is some evidence of an additional com-
plicating factor in certain regions where whales
close to partutition are encountered. At least one
gunner (the Master gunner at Norfolk Island) has
stated that he can recognize females in very late
pregnancy by their slower movements and gen-
eral behaviour, and he is therefore able to select
against such animals. The lack of females in late
pregnancy in Norfolk Island catches at periods
when this group would be expected to pass the
island, lends some support to his contention.

Another group of pregnant whales are those
which are southbound shortly after conception in
the tropics and consequently the foetuses are very
small during the southward migration. There are
relatively few tropical and temperate zone re-
cords of this group partly because the greater part
of southern hemisphere shore whaling is concen-
trated on northbound animals and partly because
the very early foetuses are difficult to find in the
brief examination which is all that is possible at
many shore stations.

d) Curtailment of Season. Catches in some re-
gions have been restricted to limited parts of the
humpback season by specifying dates between
which catching is permitted or by imposing quo-
tas which, when filled, close the season. Antarc-
tic catches since 1949 have been made in periods
ranging from four to fourteen days in any one
season, so have been too short to demonstrate
time sequences within a season. There is even
some evidence that the three of four month catch-
ing periods between 1931 and 1939 were proba-
bly insufficient to sample adequately the
sequence of humpbacks during arrival and depar-
ture from the feeding grounds.

Quotas setting the upper limit of catch were
applied to all Australian whaling stations during
post-war years. When humpbacks were abun-
dant, the existence of quotas encouraged compa-
nies to delay whaling until there had been
numerous sightings that would assure the gunners
of regular catches with good opportunities of
selecting large animals. The earliest humpbacks
were therefore under-represented in catches.
When quotas were filled quickly the late hump-
backs were not sampled. In some cases catches
sampled part only of the northbound stock and in
other seasons part of the southbound as well as all
but the earliest northbound whales were sampled.

e) Specification of northbound and southbound
humpbacks. At whaling localities north of 30°S,
there is some overlap between the last north-
bound humpbacks and the earliest southbound
humpbacks, but the catch data do not specify the

109

direction followed by individuals caught. Fortu-
nately there is some information giving the ap-
proximate dates on which equal proportions of
humpbacks are sighted travelling north compared
with others travelling south past each locality
except Congo, The latter, however, is probably
close to the tropical end point of migration of the
local humpbacks. All humpbacks caught prior to
the date on which equal proportions travel north
and south are classified as northbound and those
after it are classified as southbound. Some of the
humpbacks that travel north late and some of
those that travel south early are therefore allo-
cated incorrectly.

f) Effects of Sampling Errors. The sampling
errors due to size selection, protection of cows
accompanied by calves and the changing propor-
tions of pregnant animals during migration,
change substantially the degree of representation
of several humpback categories at various lati-
tudes, but there is no evidence that it modifies
significantly the mean dates calculated for the
categories specified in this study. So long as the
type of selection at any one station remains fairly
constant throughout the season, the degree of
under- or over-representation of each category
should also remain relatively steady throughout
the season. The effect of under-representation is
a tendency to increase slightly the standard error
of the mean rather than the position of the mean.
Providing each category is homogeneous and
represented by numbers adequate for analysis,
catch samples appear to be a satisfactory guide to
the time of migration past each locality.

The apparent migration times of heterogeneous
groups are, however, changed whenever there is
a difference in selection between groups con-
tained within the complex. Catches of mature
females for example can rarely, if ever, be subdi-
vided completely into lactating, resting and preg-
nant categories. There is frequently a
heterogeneous group which includes resting and
an unknown but probably variable proportion of
unspecified animals (R & U group in tables), that
are selected differently within a single locality.
Among northbound whales, R & U includes fe-
males near or at the end of lactation, resting
females and negligible number of unrecognized
pregnant animals.

Even at stations where the data and material
allow classification of most mature females into
lactating, resting or pregnant, it is reasonably
certain that some recently lactating animals are
unrecognized as such and are classified as resting.
As lactating or recently lactating animals tend to

110

travel northwards earlier than other categories,
the merging of some of these with resting animals
displaces the apparent mean date forwards hy an
unknown amount. However, the displacement 18
presumably greatest when no separation of lactat-
ing and resting animals is possible, and least
when a high proportion of lactating animals can
be specified separately from resting females,
This is further complicated by changes in status
of some mature females with latitude and with the
dilferences in gunner selection referred to in sec-
tion (b) above, To indicate this diagrammatically
in Fig. 23, confidence limits have been omitted in
the case of mixed females, bat calculated means
and standard deviations are included to account
for all specimens, The actual values obtained are
retained for reference in Table 12. Values for
those ‘resting’ females from which a high propor-
tion of lactating animals could be separated are
shown in Fig. 23, but the above reservations on
the relative homogeneity of this group should be
borne in mind.

The mixture of all categories of mature Females
in contrast to mature males has been used ire-
quently in past comparisons and has been shown
in Figs 1-22 as a dotted line. These figures suggest
great variability in seasonal trends between local-
ities, hut examination of dara on the subdivisions
within all mature females shows that this appar-
ent variability is largely the result of changes in
the relative proportions of the subdivisions.

Among southbound mature females there is å
mixed group that includes resting females and
same in early pregnancy, but there is no evidence
that gunner selection operates differently he-
tween the two categories at any one locality or
between different localities. As the available data
are insufficient ro separate these categories, esti-
mated values lor the mixed group are included in
Fig. 25 as well as Table 14, but the group ts clearly
not as homogeneous as any of the other south-
bound categories specified.

Any curtailment of season that delays catehing
until after some of the early humpbacks have
passed the catching zone results 1n à calculated
mean date that is late compared with mean date
which would have been obtained if the early
animals had been represented in the catch, Simi-
larly, if the season ends before the last category
of humpbacks has passed, the calculated mean
date for the latter will be displaced forwards.
Both early and late curtailment of the season
therefore Causes displacement towards the gen-
eral mean of the total catch, Incorrect allocation
of some individuals into northbound or south-

MEMOIRS OF THE QUEENSLAND MUSEUM

bound categories in cases of overlap between the
two migrating streams also tends to minimize
calculated differences between the timing of cat-
egories, Differences thal nevertheless prove
highly significant are likely to be real and to be
greater than indicated in this study.

RESULTS

The time sequence followed by specified cate-
gories of humpbacks as they travel to and from
tropical breeding regions has been studied frorn
samples obtained al a range of latitudes. Those in
temperate and tropical waters have been consid-
cred first followed by an examination of those in
Antarctic latitudes. The samples from temperate
and tropical regions show no difference in trends
between separate breeding stocks off west Africa,
cast Africa, west Australia and east of Australia
so the coastal whaling localities have been con:
sidered in order of latitude from south to north
(except for Tonga) regardless of longitude. This
arrangement approximates the sequences in
which lines of latitude are traversed by north-
bound humpbacks. These form the largest pro-
portion of the catch sample al all temperate and
tropical latitudes except at Tonga where south-
bound humpbacks only are caught. Catch sam-
ples from the latter have threfore been discussed
after a consideration of those from other non-
polar localities, followed hy the results obtained
rom Antarctic waters,

TEMPERATE AND TROPICAL WATERS

Cook Strait, New Zealand, 4175. IL has been
shown previously (Dawbin, 1956) that catches in
Cook Strait are taken almost exclusively from
whales during one stage of migration, Le, norih-
bound towards the breeding areas. Southbound
humpbacks pass through Cook Strait exceedinglv
rarely. Lookouts are posted at high vantage points
some weeks before the humphacks are expected
to traverse Cook Strait, and catching cominences
immediately after the first few humpbacks have
been seen. Operations continue untl a week or
more has passed without sighting a humpback-
The catch, therefore, includes samples from start
to end of the nonhward migration past Cook
Strait, and the majority of those sighted have been
caught.

The small hoat catching methods and the days
lost through bad weather ensures thal gunners
catch whales primarily through availability with
little possibility of selection, except for rejection
of the smallest whales.

TEMPORAL SEGREGATION OF HUMPBACK WHALES II)

TABLE 1. Weekly catches of specified humpbacks,
1947-1960. Cook Strait, NZ, 4198.

omme | | mane ë |

Stan of week
All immatures
All humpbyeks

ino d

I

2
2
il
3
2
2

b

— wn à D

raat) ae [ee [am [aoe ENE

Samples have been obtained through the
catches of each of the seasons 1947 to 1960
totalling 1922 humpbacks. This is supplemented
by sighting data on others, including the highly
significant group of weaning yearlings observed
as accompanying their mothers.

Females in late lactation form the earliest cate-
gory at Cook Strait (Figs 1-2, Table 1). Most have
heen caught before mid-June, and none later than
mid-July in any of 14 seasons. During this period
the size of the local humpback stock appears to
have been fairly stable, so the number of females
in late lactation (as defined above) should have
been approximately the same as those in late
pregnancy, except for a slight reduction caused
by the natural mortality of some calves before
weaning. The 120 recorded compares with 164 in
late pregnancy, and suggests thal a few in late
lactation may have been unrecognised and
grouped among the 216 recorded as resting and
unspecitied. However, no ‘yearlings’ accompa-
nying their mothers have been observed after
mid-July, so it seems probable that the time se-
quence indicated. by the recorded females in late
lactation is substantially correct.

Immature males and lemales (Table 1) show no
significant difference in mean dates. They have

therefore been combined in Figs 1 and 2 which
shows the number and percentage frequency per
fortnight respectively, It is evident that largest
numbers occur about one week later than late
lactating fernales, and about two weeks earlier
than those of mature males, Immature animals
have been represented from the beginning of each
season, but none have been caught during the
final two weeks in any season. Percentage Ire-
quences (Fig. 2) show that immature animals
form å steadily decreasing proportion of the catch
per fortnight as the season progresses, but Fig. I
shows that this is the result of a relatively small
lime difference between the attainment of peak
density by immature animals compared with the
categories of mature humpbacks.

Mature males form the largest single category
of humpbacks at Cook Strait. None have been
caught before mid-May, but the group has been
represented during the remainder of the season,
Peak density occurs in late June with a mean date
of June 29.

Resting females, including a few unspecified
animals that may have been in late lactation, have
followed a very similar ime sequence ro that of
mature males. There is no significant difference
in mean dates or duration of season between these
categories, but there have been nearly four times
as many mature males as resting females 1n the
catch.

Females in late pregnancy have formed the last
category in all seasons during which they have
been represented, None have occurred in May
and the mean date (July 8) is about one week later
Ihan those of mature males or resting females.
Pregnant females are therefore approximately
one month later than those in late lactation.

Great Barrier Is., New Zealand 36°S, As at Cook
Strait, the humpback catches from waters near
Great Barrier Island include a high proportion of
all those sighted, and represent all categories
except those of smallest sizes. The last north-
bound humpbacks leave the area some five or six
weeks hefore the return of the carliest southbound
animals, so there is no diffculty in separatng
animals on direction of travel. Data with gonads
and orher material have been obtained during the
three seasons 1959-1961 relating to 231 north-
bound and 33 southbound animals, but the mate-
rial does not include mammary gland samples or
observations on yearlings accompanying their
mothers. It is therelore impossihle ti specify It-

males in Tate lactation.

112 MEMOIRS OF THE QUEENSLAND MUSEUM

NUMBER OF WHALES

250

COOK STRAIT N.Z.
AIS

IMMATURE

MATURE MALES

TOTAL MATURE FEMALES
END OF

RESTING FEMALES
LATE PREGNANT

us 1575 29/5 12/6 26/6 10/7 247 718 ave EA 18/9 2/10 160 aono
FORTNIGHTS COMMENCING

FIG. 1. Fortnightly catches of specified humpback categories, 1947-1960. Cook Strait, NZ, 41°S.

ioo COOK STRAIT NZ.
COOK STRAIT NZ.

PERCENTAGE OF TOTAL CATCH

EI

IMMATURE

MATURE MALES

vs 15/5 295 12/6 26/6 10/7 24/7 HI 218 an 18/9 2/0 16/10 30/10
FORTNIGHTS COMMENCING......

FIG. 2. Fortnightly percentages of specified humpback categories, 1947-1960. Cook Strait, NZ, 41°S.

TEMPORAL SEGREGATION OF HUMPBACK WHALES

TABLE 2. Weekly catches of specified humpbacks,
1959-1961. Great Barrier Island, NZ, 36°S.

113

TABLE 3. Weekly catches of specified humpbacks,
1952-1958. Albany, WA, 35*S.

Immature
E P
1 E $
3 E: Si
5 E E
5 5 =
Å z z
22/5 2
29/5 Za ai] 1 3 9
5/6 2 | 10 | i 7 28
126 | 21 | 3 | 24 | 14 7 45
196 | 11 | 5 | 16 | 17 5 | 1 | 39
26660 4 | 2 | 6 | 13 5 | 2 | 26
37: | 2 | 2 |] a4 | 9 3 | 3 | 19
mol 5 | 1 | 6 | 24 4 | 3 | 37
17/7 1 1 | 9 6 | 1 | 17
24/7 3 1 | 4
317 4 1 5
7/8
14/8
21/8
28/8
4/9
11/9 a | 2 2 4
18/9
29 | 2 | 3 | 5 5 10
2110
9/10 2 2
1600 1 | 2 | 3 | 1 2 6
23/10 2 3 | 4 4 10
30/10 1 1 1
Total | 58 | 28 | 86 |m] 56 | 11 [264]

The earliest identifiable category comprises the
immature animals (Figs 3-4, Table 2) which are
represented in catches from the start of the season
in late May until about mid-July, but none have
been caught in the last two weeks of the north-
bound season. The mean date is 11 days earlier
than that for mature males.

Mixed mature females include both resting and
unspecified lactating females, and are repre-
sented throughout the season. The estimated
mean date is mid-way between those for imma-
ture animals and mature males, so is relatively
earlier than resting females at Cook Strait. This is
almost certainly the result of including all the late
lactating females, since this group elsewhere
tends to travel earlier than other mature females.

Mature males form the largest group and are
represented throughout the season. The mean
date is 11 days after that of immature animals, and

Immature Mature

på œ OF OF 2
1 «|f. z|z2|s|à
ha E S | «|| È
8 : E
EI < <
29/5 1 1 1 2 0 4
5/6 2 6 8 1 6 1 0 16
12/6 8 10 18 7 16 2 0 43
19/6 | 10 11 21 19 10 6 0 56
26/6 | 13 8 21 36 8 5 0 76
3/7 14 17 31 39 8 11 6 95
10/7 | 17 9 26 46 9 19 4 104
17/7 9 4 13 48 5 20 7 93
24/1 5 9 14 33 3 17 14 81
31/7 3 1 4 29 2 17 13 65
7/8 1 2 3 11 5 12 31
14/8 2 1 3 10 5 7 25
21/8 0 0 0 6 3 l 10
28/8 0 1 1 3 1 1 6
are 79 | 164 |

8 days earlier than for pregnant females. Pregnant
females form a small sample, but all were ob-
tained after the first month of whaling, and the
group was clearly the last northbound category.

Southbound humpbacks caught between mid-
September and late October form a small sample
in which there is no demonstrable difference in
timing between mixed females and immature
humpbacks, but the five mature males obtained
were all caught after mid-October. (Fig. 25, Ta-
bles 2, 14).

Albany, WA, 35?S. The catch is taken from an
exclusively northbound population, as hump-
backs returning south from Western Australian
coastal breeding areas follow a route some dis-
tance west of Albany. Although quotas have been
applied in each season, catching has been carried
on throughout most of the local humpback season
before the quotas were filled. Except for protec-
tion of undersized whales, the Albany catch ap-
pears to be a representative sample of the local
stock throughout its season of migration past the
locality. Catching has commenced about three
weeks later than at the higher latitude of Cook
Strait. Catches in the seven seasons 1952-1958
include 699 humpbacks.

Females classified as at or recently in late lac-
tation form the earliest category (Figs 5, 6, 23,
Tables 3 and 14) with peak density in mid-June

114 MEMOIRS OF THE QUEENSLAND MUSEUM

40-

GREAT BARRIER IS. N.Z.
36's — IMMATURE
— MATURE MALES
(—— TOTAL MATURE FEMALES
O----O RESTING
LATE PREGNAN’

20:

NUMBER OF WHALES

us 15/5 29/5 12/6 26/6 0/7 24 7/8 2ve 4n 18/9 210 1640 300
FORTNIGHTS COMMENCING......

FIG. 3. Fortnightly catches of specified humpback categories, 1959-1961. Great Barrier Is., NZ, 36°S.

GREAT BARRIER IS. NZ.
36'S
IMMATURE

mms MATURE MALES

tM TOTAL. MATURE FEMALES

O- ---O RESTING
år ---Àà LATE

80:

e

PERCENTAGE OF TOTAL CATCH
D

8

5 15/5 205 12 2646 Lo 24/7 KO 240 Ku E 20 16/10 30/0
FORTNIGHTS COMMENCING

FIG. 4. Fortnightly percentages of specified humpback categories, 1959-1961. Great Barrier Is., NZ, 36°S.

TEMPORAL SEGREGATION OF HUMPBACK WHALES 115

999 ALBANY W.A.

IMMATURE

MATURE MALES

NUMBER OF WHALES

26/6 10/7 24/7 KO
FORTNIGHTS COMMENCING......
FIG. 5. Fortnightly catches of specified humpback categories, 1952-1958. Albany, WA, 35°S.

993 ALBANY W.A.

35's — IMMATURE
-—— MATURE MALES

PERCENTAGE OF TOTAL CATCH

US 15/5 29/5 12/6

26/6 10/7 24/7 7^ 21/8 an 18/9 2/0 16/10 30/0
FORTNIGHTS COMMENCING......

FIG. 6. Fortnightly percentages of specified humpback categories, 1952-1958. Albany, WA, 35°S.

116

TABLE 4. Weekly catches of specified humpbacks,
1948-1954. Durban, Natal, 30°S.

| Immature | Mature "w
a O+ er o ZS
$lelo E to I 2 E 3
e E bad ebe:
: E å
a < <
15/5 0 1 1 0 0 l
22/5 2 0 2 0 0 2
29/5 5 2 7 1 1 9
5/6 3 5 8 6 6 20
12/6 | 10 13 23 10 4 37
19/6 | 21 17 38 20 12 70
26/6 | 28 37 65 31 15 111
3/7 22 24 46 42 15 103
10/7 | 18 20 38 37 15 90
17/7 | 14 23 37 24 21 82
24/7 | 14 16 30 29 16 75
31/7 7 12 19 6 10 35
7/8 7 6 13 4 5 22
14/8 2 3 5 2 10
21/8 6 8 14 6 4 24
28/8 2 10 12 2 2 16
4/9 0 0 0 2 i 3
11/9 3 2 5 5 5 15
18/9 II 12 23 13 10 46
25/9 | 12 12 24 19 7 50
2/10 l 5 6 7 6 19
9/10 3 l 4 4 3 11
16/10| 0 0 0 l 0 H
23/10| 0 0 0 l 0 1
30/10| 0 0 0 l 0 l
Total | 191 | 229 | 420 | 273 161 854

and none during the last four weeks of the season.
The mean date (June 28) is about one week earlier
than that of immature whales. The latter are rep-
resented throughout the season, but of 164
caught, seven only were caught during the last
four weeks of the season. Mature males have also
been caught through the season, but peak density
occurred during July and the mean date was 10
days later than for immature whales. Resting and
unspecified females had a time sequence similar
to that of mature males. The small difference in
mean dates is not significant.

Females in late pregnancy have not been re-
corded during the first five weeks of the season,
and their mean date is about two weeks later than
that of mature males. They have occurred one
month later than late lactating females, and they

MEMOIRS OF THE QUEENSLAND MUSEUM

comprise the last category to migrate northwards
past Albany.

Durban, Natal 30°S. Humpback catches from
Durban during post-war years have differed from
those made near any of the other temperate and
tropical localities under consideration in being
secondary to catches of other species. Sperm and
fin whales have formed the major part of the
catch, and both of these species tend to occur at a
greater distance from shore than most hump-
backs. While the writer has no personal knowl-
edge of the local conditions, the total catch
composition suggests that humpbacks are caught
as opportunity arises between catches of the more
valuable and intensively sought sperm and fin
whales, so the humpback catch is possibly not a
consistent sample of the locally migrating stock.

The composition of humpback catches is un-
usual in the very high proportion of immature
animals, which on the present criteria represent
nearly half the total catch, and on the criteria used
in International Whaling Statistics they form
about two thirds of the total catch. The absence
of foetus records suggests that pregnant females
traverse waters closer to shore or further out to
sea than the main catching zone, or that the data
on pregnant females is incomplete. The overall
low percentage of mature females shows that
incomplete recording cannot be the full explana-
tion, and may not even be part explanation.

The writer has no substantial data on which to
separate northbound from southbound animals,
but the bimodal peaks in the present and Mat-
thews (1937) data indicate that the change occurs
during August. Trial analyses were carried out on
data from the start of season to August 13, August
20, August 27 and September 3, with little differ-
ence in sequence resulting. Mid-August (actually
up to August 13) has been used in the calculations
shown in Table 12 & Fig. 23. The post-war data
relates to 854 humpbacks caught during the 1948-
1954 seasons, and includes 667 which have been
classified as northbound.

Immature whales form the earliest identifiable
category, followed by mature males, then mixed
females, but the time intervals between the mean
dates of these categories (Table 12) are too small
to be significant at the 596 level.

Pre-war catches generally included a higher
proportion of humpbacks than other species, so
they are likely to be a more representative sample
of the local humpback stock. Catches by one
company at Durban in 1934 and 1935 included
547 that have been classified as northbound since

TEMPORAL SEGREGATION OF HUMPBACK WHALES 117

"© DURBAN S.AFRICA
30

IMMATURE

NUMBER OF WHALES

us. 15/5 295 SR 26/6 "o7 24/7 UL ave 4m 18/9 2/10 16/10 30/0
FORTNIGHTS COMMENCING......

FIG. 7. Fortnightly catches of specified humpback categories, 1948-1954. Durban, Natal, 30°S.

99] DURBAN _S. AFRICA

30°S —— IMMATURE

PERCENTAGE OF TOTAL CATCH

vs 155 DO Rb 266 Lo 24/7 U ave ar eo 20 wo 30/10
FORTNIGHTS COMMENCING......

FIG. 8. Fortnightly percentages of specified humpback categories, 1948-1954. Durban, Natal, 30°S.

118

TABLE 5. Weekly catches of specified humpbacks,
1954-1961. Byron Bay, E Australia, 298.

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6. Weekly catches of specified humpbacks,
1956-196]. Norfolk Island, 2995.

Ímmature Mature

ia » OF 4

«| å aj 3 E E n E
E E $ å E

| = x a x x

3 3

6 1 xi 5

3 6 21 34 15
3 4 25 12 4l 4l
3 6 46 15 l 68 3/7 52
| 4 44 18 1 7 10/7 8 59
4 5 76 17 3 | 101 WT| 4 2 36
1 3 52 2 7 9I 24/7 | 4 6 7 42
1 2 63 35 3 | 103 37| 2 2 36 12 52
2 2 37 27 8 74 7/8 5 l 9 42
1 I 17 16 6 40 14/8 | l 4 2 31
A 5 15 13 3 36 21/8 | 1 9 20
1 1 9 14 24 28/8.| 2 3 G 43
7 10 10 ål l 52 4/9 5 2 20 32
i 4 19 18 41 WA) 3 15 70
4 6 26 26 58 18/9 5 4 9 58
6 16 | 25 3 [ 55 259 | 6 9 5 56
3 7 30 g 1 46 Wo} 5 4 9 84
3 7 19 I 27 SAO] 2 I 3 63
I I 21 22 16/10 2 2 29
3 4 17 2 23 23/10 2 37

2 18 1
304 | 35 | 1040 2

they were caught prior to mid-August. Among
these, the order of appearance was the same as in
post-war years, but the differences in mean dates
were larger and were very highly significant.
During the three weeks from mid-May into early
June, immature animals only were caught. The
mean date for immature whales (July 6) was 8
days earlier than for mature males, and two weeks
earlier than for mixed females, but there are no
datà on which to subdivide rhe latter into lactat-
ing, resting and pregnant,

There are bimodal fluctuations within the post-
war catches of southbound humpbacks (i.e., after
mid-August) (Table 4, Figs 7-8) that cannot be
explained by the writer. However, the pre-war
catch sample from Durban in 1934 and 1935 was
unimodal, and showed that immarure whales pre-
cede mature males and mixed females on the
southward migration.

Byron Bay, EA 29°S8, Norfolk Island, 29°S,
Moreton Island, EA 27?8. Catch samples from
stations of eastern Australia and Norfolk Islànd
have shared substantial similarities due to quotas
and selection factors in common. The samples
used. for analysis include the following seasons
and catches:- Byron Bay 1954-1961, 1040; Nor-
folk Island 1956-1961, 880; Moreton Island
1952-1961, 6205, Although variations in the size
of catch at each locality tend to mask some of the
similarities (Figs 9, 11, 13), the changes in per-
centage composition per fortnight at each are
strikingly similar (Figs 10, 12, 14).

At each locality, the quotas during most of the
seasons considered in this study were markedly
smaller than the number of humpbacks sighted.
This has made it possible for gunners to select
more stringenily than occurs at localities where
there is only asmall excess of sightings compared
to catch.

TEMPORAL SEGREGATION OF HUMPBACK WHALES 119

150

BYRON BAY EA.
29's
IMMATURE
135.

— MATURE MALES

tM TOTAL MATURE. FEMALES

O----O RESTING
&----À LATE PI

120

105

NUMBER OF WHALES
e

b
vm

30:

Us 15/5 29/5 12/6 7/8 21/8 4/5 18/9 2ho 16/10 3010

2 107 24/7
Portia Ng contenen.. AP

FIG. 9. Fortnightly catches of specified humpback categories, 1954-1961. Byron Bay, E Australia, 2998.
999 BYRON BAY E.A.

———— IMMATURE
G
295
-e MATURE MALES
90
um TOTAL MATURE FEMALES
O----O RESTING
BO
&----AÀ LATE PI
70
I
S
e
Ki
o 60-
å
4
E
o
e
50.
u
o
ul
o
Zap
z
MI
o
a
E
a

us 15/5 29/5 12/6

26/6 10/7 24/7 7/8 ?2y8 KO 18/9 2/10 16/0 30/10
FORTNIGHTS COMMENCING......

FIG. 10. Fortnightly percentages of specified humpback categories, 1954-1961. Byron Bay, E Australia, 2998.

120 MEMOIRS OF THE QUEENSLAND MUSEUM

!'09-] NORFOLK IS.
C298. e — IMMATURE

SO

80.

70:

E)
Q

8

NUMBER OF WHALES

HO 15/5 29/5 12/6

295 17 24/7 7/8 EI) ay 18/9 -o 16/10 30/0
FORTNIGHTS COMMENCING

FIG. 11. Fortnightly catches of specified humpback categories, 1956-1961. Norfolk Is., 29°S.

| NORFOLK IS.
29% — IMMATURE

PERCENTAGE OF TOTAL CATCH

VS 15/5 29/5 12/6 26/6 1007 24/7 718 2us 4/9 18/9 A0 ho 3010
FORTNIGHTS COMMENCING

FIG. 12. Fortnightly percentages of specified humpback categories, 1956-1961. Norfolk Is., 29°S.

TEMPORAL SEGREGATION OF HUMPBACK WHALES

TABLE 7. Weekly catches of specified humpbacks,
1952-1961. Moreton Island, E Australia, 27^5,

Immature

to Dn

Start of week
All humpbacks

å

6
[Tei] 384 | 436 | s20 [as02| | 1347 [ 146 [e205

From each station, it has been the practice 10
avoid adults accompanied by yearlings, whether
or not the adults may have been females which
were al the point of weaning offspring capable of
independent existence. Late lactating females
have therefore been largely absent from catch
samples,

Selection for size has been intense, especially
during the early part of each season when gunners
are usually hopeful of fulfilling the quota from
lurge whales only. From each station, the whales
caught during the earliest one or two weeks of the
season have all been mature. Among the total
catch of northbound humpbacks, the percentage
uf immature animals caught has been about one
quarter of that al non-quota localitiesin the south-
em hemisphere (6, 12 & 11 for the former, 25-50
for the latter).

Since there is some indication that the under-
representation of immature animals. changes
within each season according to the whalers’ as-
sessment of progress in filling the quotas, the
available catch data may be a misleading sample
of the local time sequence for immature whales,
The calculated mean dates at each locality (Fig.
23, Table 12) show no significant differences
from those of mature males. The latter are consis-
tently earlier than mixed mature females which,
due to selection against late lactating females, are
mainly ‘resting’ animals.

Pregnan females are unrepresented in catches
from Norfolk Island, but at Byron Bay and
Moreton Island they form the last northbound
category, The mean dates are approximately two
weeks later than those tor mature males at both
localities.

Humpbacks classified as southbound have in-
cluded those caught after August 21 at Byron Bay
and Moreton Island, and after September 4 al
Norfolk Island, since equal proportions of north-
bound and southbound whales were sighted at the
respective localities on these dates.

Mixed females (early pregnant and resting)
form the earliest southbound category at each
locality, but the differences in time in relation to
immature whales is non-significant, except at
Byron Bay (Fig, 25, Table 14), Mature males are
later than immature whales by amounts which ure
highly significant in each locality (Moreton Is., 5
days; Byron Bay, 8 days; Norfolk Is., 14 days),
Females in early lactation are unrepresented in
catches, but Chittleborough (1962) has shown
that this category 1s the last to travel south along
ihe coast of eastern Australia.

Carnarvon, WA 25°S, Point Cloates, WA 22^8. At
both these stations the quotas per season were
larger than those for Byron Bay, Moreton Island
and Norfolk Island. The catch samples to be
discussed here include Carnarvon 1950-1958,
5889, and Point Cloates 1949-1955, 3321. Catch:
ing at the latter locality ceased after the 1955
season. The length frequencies of catches indi-
cates that gunner selection was usually less in-
tense than at the latter stations and the catch
samples used (1950-1958 at Carnarvon and 1949-
1955 ut Point Cloates) appear to be representative
of the northbound population, except for hump-
backs under 35 feet in length (Figs 15-18, Tables
8-9), The sample classified as northhound in-
cludes all those caught before August 28 at Car-
narvon and August 2! al Point Cloates
(Chittleborough, 1953).

122 MEMOIRS OF THE QUEENSLAND MUSEUM

10001 MORETON IS. E.A.
27's IMMATURE

emmemmm MATURE MALES

* TOTAL MATURE FEMALES

RESTING
LATE

NUMBER OF WHALES

107 24/7 18 2v8 4/9 18/9 2/10 oO 30NO
FORTNIGHTS COMMENCING......

FIG. 13. Fortnightly catches of specified humpback categories, 1952-1961. Moreton Is., E Australia, 27°S.

Ee
99 MORETON (IS. E.A. IMMATURE
275

— MATURE MALES

90. - TOTAL MATURE FEMALES

O----O RESTING
år ---À LATE Pi

OF TOTAL CATCH

PERCENTAGE
a

8

us 15/5 29/5 12/6 266 197 24/7 78 208 AR 18/9 aho 1640 3010
FORTNIGHTS COMMENCING......

FIG. 14. Fortnightly percentages of specified humpback categories, 1952-1961. Moreton Is., E Australia, 2798.

TEMPORAL SEGREGATION OF HUMPBACK WHALES

TABLE 8. Weekly catches of specified humpbacks,
1950-1958. Carnarvon, WA, 25°S.

TABLE 9. Weekly catches of specified humpbacks,
1949-1955. Point Cloates, WA, 22°S.

Immature Mature
1 g S In. | Seok oe
E fo : “o 3 ` É Å
: E E
a x <
3 2 8 0 0 13
5/6 5 8 13 9 42 4 0 68
12/6 | 14 24 38 60 95 11 3 207
19/6 | 30 17 47 61 97 47 2 254
26/6 | 26 31 57 139 | 94 55 1 346
3/7 40 43 83 175 | 67 92 6 423
10/7 | 35 47 82 186 | 49 133 5 455
17/7 | 27 25 52 | 208 | 33 133 2 428
24/7 | 20 27 47 | 237 | 23 138 2 447
31/7 | 28 30 58 | 272 15 134 5 484
7/8 31 21 52 | 289 492
14/8 | 23 31 54 | 293 466
21/8 | 36 14 50 | 278 440
28/8
4/9
9
9
3
2
0
0
0
0

The number of females at the end of lactation
have been calculated from mammary gland data
kindly provided by Dr R.G. Chittleborough. Lac-
tating females form the earliest category with
mean dates (Fig. 23, Table 12) at Carnarvon 20
days and at Point Cloates 11 days prior to those
for immature animals. The latter are 10 and 8 days
earlier than the mean dates for mature males. The
above time intervals are all very highly signifi-
cant, but there is no significant difference be-
tween the mean dates for mature males and
resting females.

Pregnant females form the last northbound
group with mean dates 6 and 4 days later than
mature males and about one month later than
females at the end of lactation.

Catches off Western Australia during 1925-
1928 were taken without quotas or length restric-
tions and therefore include a considerable
number of the small size clases that are absent

Mature

Immature

Start of week
All humpbacks

= O| Allimmatures

19/6 2) 18 33
26/6 | 17 24 41 26
3/7 22 38 60 68
10/7 | 13 21 34 74
177 | 24 60 84 | 123
24/7 | 20 27 47 130
31/7 | 17 32 49 157
7/8 18 27 45 170
14/8 7
21/8 | 26
28/8 | 25
4/9 22
11/9 | 18
18/9 5
25/9 | 4
2/10 H

0

9/10
Total | 244

from post-war catches. Those of less than 30ft in
length are of special interest, since they form the
size classes represented among weaning
*yearlings'. In a total catch of 3,426 there were
174 examples occurring as follows:- 10 in June,
115 in July, and 49 in August. This represented
21%, 13% and 496 of the catch in these months
and the ‘yearlings’ were clearly the earliest ho-
mogeneous group to pass the locality. Subsequent
data have shown that most yearlings accompany
the mother during the major part of the northward
migration. Although the individual females can-
not be specified from the data obtained during the
above seasons, the time sequence of the yearlings
is completely consistent with the occurrence of
late lactating females as the earliest category in
post-war catches. Immature animals (excluding
yearlings) were the next group followed by ma-
ture males and mixed females. There are no data
on pregnant animals. Thus all the categories
which can be recognised in the 1925-1928
catches are consistent in time sequence with those
in the 1949-1958 cathces.

124 MEMOIRS OF THE QUEENSLAND MUSEUM

CANARVON W.A.
25's

IMMATURE
MATURE MALES

* TOTAL MATURE FEMALES
@----@ END OF
O----O RESTING
LATE PREGNANT

42

w

WHALES
P
o
o

OF
H

NUMBER

VS 15/5 29/5 12/6 26/6 om 24m 7/8 ave als 18/9 2ho 1640 30/!0
FORTNIGHTS COMMENCING......

FIG. 15. Fortnightly catches of specified humpback categories, 1950-1958. Carnarvon, WA, 258.

1099 CANARVON W.A.

25's IMMATURE

— MATURE MALES
— TOTAL MATURE FEMALES
@----@ END OF

80 O----O RESTING

LATE PREGNAN

BI
o

PERCENTAGE OF TOTAL CATCH
b
o

w
[*]

UO IS/5 2915 12/6 26/6 10/7 24/7 HO) 2/8 alo 18/9 20 1640 30N0

FORTNIGHTS | COMMENCING...

FIG. 16. Fortnightly percentages of specified humpback categories, 1950-1958. Carnarvon, WA, 25°S.

TEMPORAL SEGREGATION OF HUMPBACK WHALES 125

POINT CLOATES W.A.
22's

— IMMATURE
— MATURE MALES

Lo SE LL CQ QS MT TOTAL MATURE FEMALES

@----@ END OF

O----O RESTING

LATE PREGNANT

24

210.

180:

NUMBER OF WHALES
5
o

a
o

60

15 15/5 29/5 12/6 26/6 977 24/7 We 21/8 Als 18/9 210 Kiel 30/10
FORTNIGHTS COMMENCING......

FIG. 17. Fortnightly catches of specified humpback categories, 1949-1955. Point Cloates, WA, 2208.

POINT CLOATES W.A.
22°S

IMMATURE

MATURE MALES
90:

* TOTAL MATURE FEMALES

e----e BDO
O----O RESTING
A ss sch LATE PREGNANT

e

yw

D

PERCENTAGE OF TOTAL CATCH

w

us s5 2915 126. Zeie I7 247 7/8 21/8 KO 18/9 2/0 eno 30/0
FORTNIGHTS COMMENCING......

FIG. 18. Fortnightly percentages of specified humpback categories, 1949-1955, Point Cloates, WA, 22°S,

During the southward migration, immature an-
imals have been the earliest recognisable cate-
gory to travel past Western Australian land
stations, while aerial observations by
Chittleborough (1953) have shown that cows ac-
companied by young calves comprise the last

group.

Madagascar, Pelagic, 16°-26°S. The catches
from the coastal waters of Madagascar have been
made by catchers operating with a factory ship
that moved between 26°S & 16°S with 22°S as
modal position, Irregularities of movement
through this distance may have had some effects
on catch composition, but there are no data indi-
cating selection apart from rejection of under-
sized whales. There are no data on which to
specify lactating females. The catch in 1948 and
1949 totalled 2014 humpbacks (Table 10, Figs
19-20) of which 1819 caught before September
11 have been classified as northbound. This date
has been derived from descriptions of the above
seasons by Angot (1951).

Immature animals form the earliest recognis-
able category to the extent that they occur in
approximately equal numbers to those of mature
males between mid-June and mid-July, but occur
in smaller numbers than mature males during the
remainder of the season. This results in a calcu-
lated mean date of July 28 (Table 12, Fig. 23) for
immature animals compared with July 31 for
mature males. While this interval of 3 days is
small, it is nevertheless significant at the 1%
level. Mixed females, including unspecified late
lactating and resting animals show no significant
difference in timing compared with mature
males.

Pregnant females have a mean date 16 days
later than mature males and are the last north-
bound category.

The catches of southbound humpbacks (i.e.,
after September 11) were made during 3 to 4
weeks only. This period is insufficient to establish
the time sequence of southbound animals passing
Madagascar.

Congo, 1°S. There is little difference in latitude
between Gabon and Sáo Tomé Is. and the writer
is unaware of significant differences in selection
at the two localities, Catches have therefore been
pooled under the heading ‘Congo’, and amount
to 4,071 humpbacks taken during 1949 to 1952
(Figs 21-22, Table 11). As in the other post-war
catches, animals under 35 feet in length are rarely
represented. There are no data which can be used
to specify animals that have been lactating re-

MEMOIRS OF THE QUEENSLAND MUSEUM

cently, so mature females include a mixture of the
above and resting animals. Most females that
were in late pregnancy when at higher latitudes
have delivered their calves and became protected
as cows accompanied by young calves before
reaching 198. The catch of animals in late preg-
nancy has therefore been very small.

The Congo region is at or near the northern limit
of the west African southern hemisphere hump-
backs, so there is no clearly recognisable passage
of northbound and southbound animals past the
locality. Many specimens presumably remain for
some weeks in the general breeding area before
departing southwards, and there are no data on
which to separate northbound from southbound
animals. The total catch per season has therefore
been used to calculate mean dates which, in these
cases, represent the approximate mid-point be-
tween the beginning of northbound arrivals and
the end of southbound departures for each cate-
gory. This gives an earlier date for animals in late
pregnancy relative to other categories since the
former includes only northbound animals.

Immature animals form the earliest category
(Figs 21-23) with the mean date on July 29 as
compared with August 11 for mature males and
mixed females (Table 12). Females in late preg-
nancy occur 10 days later on mean dates, and
form the last category despite the non-represen-
tation of their southbound equivalents, (i.e., cows
accompanied by young calves).

Tonga, 22°S. Unlike catches from all the preced-
ing localities, those from Tonga have been taken
predominantly from southbound animals, (Fig.
24, Table 13). The local whalers still use open
boats, hand harpoons and hand lances, and with
this equipment they concentrate, like many nine-
teenth century whalers with similar gear, on cows
accompanied by young calves. This category is
protected elsewhere by International Whaling
regulations, and is unrepresented in other catches,
While the catch from Tonga is small (82 in 5
years) it is nevertheless of special interest.

Hunting commences when cows with calves
are first sighted, and these, together with imma-
ture whales up to about 38 feet, comprise the
subsequent catch. However, observations at
Foveaux Strait, NZ, and along both west and east
coasts of Australia indicate that cows and calves
form the last southbound group.

The delay in hunting until cows and calves
appear makes it highly probable that early south-
bound immature whales will be unrepresented in
the catch. Nevertheless, the immature whales

TEMPORAL SEGREGATION OF HUMPBACK WHALES 127

2001 MADAGASCAR IMMATURE

— MATURE MALES

O----O RESTING
år ---À LATE

160:

NUMBER OF WHALES
©

E

40:

VS 15/5 2/5 126 78 2U8 4/9 18/9 2/0 Wu 300

26/6 10/7 24/7
FORTNIGHTS COMMENCING

FIG. 19. Fortnightly catches of specified humpback categories, 1948-1949. Madagascar, 22°S.

' MADAGASCAR
225

IMMATURE

-— MATURE MALES

* TOTAL MATURE FEMALES

` O----O RESTING
dr === LATE PI

CATCH

PERCENTAGE OF TOTAL

us 15/5 295 12/6 206 10/7 24/7 We 2/8 4ls 18/9 Si 16/10 3010
FORTNIGHTS COMMENCING......

FIG. 20. Fortnightly percentages of specified humpback categories, 1948-1949. Madagascar, 22°S.

TABLE 10. Weekly catches of specified humpbacks,
1948-1949. Madagascar, pelagic, 16°S-26°S.

| Immature | Mature
P or s
loloj] 5|% ` 5 å $
3 E 3 zz E| Ê
5 E S E
E S E
H E =
12/76 | 5 1 6 13 9 0 28
19/6 | 0 2 2 n 2 0 15
26/6 | 17 | 14 | 31 | 44 28 0 | 103
3/7 | 36 | 40 | 76 | 69 64 1 210
10/7 | 28 | 35 | 63 | 47 53 1 164
17/7 | 34 | 30 | 64 | 73 37 3 177
24/7 | 27 | 33 | 65 | 91 68 4 | 228
31/7 | 24 | 27 | 51 83 68 4 | 206
718 | 21 32 | 53 | 66 54 9 | 182

caught during operations for cows and calves
have a calculated mean date of August 29, which
is 16 days earlier than for cows and calves. If
allowance is to be made for immature whales that
precede cows and calves, then the above differ-
ence in time would have to be increased. Table 13
shows that (except for one animal) cows and
calves have been caught for approximately one
month longer than immature humpbacks.

ANTARCTIC WATERS. It has been shown else-
where (Dawbin, 1966) that the composition of the
post-war catch samples show no significant dif-
ference between those caught in late December
and those in early February. Pre-war samples in
the 1932-1933 to 1938-1939 seasons included
13,357 catches between early November and late
March. These were distributed between the south
Atlantic Ocean and the eastern portion of the
South Indian Ocean. This area includes regions
with substantial differences in sea temperatures
at equivalent latitudes and dates.

On the hypothesis that humpbacks would enter
Antarctic sectors with warmest water earlier than
sectors with colder water, a separation of the data

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 11. Weekly catches of specified humpbacks,
1949-1952. Congo, 198.

Immature | Mature
gl, IK de AA
: AERE
dä =
E S S
1 0 1 0 2
22 21 19 1 63
37 39 34 1 111
87 73 46 0 206
79 117 81 0 277
99 139 77 0 315
110 | 153 85 2 350
105 | 157 101 3 366
106 | 167 93 3 369
84 143 103 3 333
74 123 77 4 273
66 136 93 10 | 310
35 121 79 7 242
34 121 81 4 240
20 | 105 65 1 191
13 103 56 4 176
10 53 46 1 110
5 51 27 0 83
3 23 14 1 41
1 2 5 0 8
991 | 1847 1188 | 45 | 4071

into sectors of 20° longitude was made. Only two
of these (20°E to 39°E and 80°E to 99°E) con-
tained samples adequate for analysis. However,
in the former sector cold water extends further
north than in the latter at the same dates. Table 19
shows that by using mean dates as an index,
humpbacks occur about three weeks later than in
the latter sector.

Within each sector one would expect changes
in local catch composition during the period that
each category of humpbacks arrives or departs
from Antarctic waters. In view of the consistently
slow rate of humpback migration through a wide
range of latitudes (Dawbin, 1966) and the rela-
tively consistent difference in time sequence be-
tween categories in temperate and tropical waters
as described previously, it seemed probable that
time sequence changes would occur first at more
northerly latitudes, e.g., 55°S and would be more
or less duplicated at a later date in higher lati-
tudes, e.g., 60°S or 66°S. To test this hypothesis
the catch samples within sectors were subdivided
into three degree lines of latitude. In the absence

TEMPORAL SEGREGATION OF HUMPBACK WHALES

—TQ—Q—— IMMATURE
-— MATURE MALES

270-

210:

8 E

NUMBER OF WHALES

a
o

60:

ils 15/5 29/5 2% 26/6 kal 247 KU
FORTNIGHTS COMMENCING.....

FIG. 21. Fortnightly catches of specified humpback categories, 1949-1952. Congo, 198.

IMMATURE

— MATURE MALES

90 + TOTAL MATURE FEMALES

O----O RESTING
so &----A LATE P

70:

E
o

PERCENTAGE OF TOTAL CATCH

us 15/5 295 12/6 26 10/7 24n KU 2V8 4/9 18/9 2o
FORTNIGHTS COMMENCING...

FIG. 22. Fortnightly percentages of specified humpback categories, 1949-1952. Congo, 198.

tM TOTAL MATURE FEMALES

O----O RESTING
år ---À LATE PI

ave a 18/9 2/0 o

ho

129

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 12, Estimated mean date with standard deviation and the 95% confidence limits of the mean for specified
categories af northbound humpbacks. * Since mixed females form a heterogeneous group, these data are nol
siriclly comparable with those relating to (he more [iiia ai categories. 7

0.19

july | 0.37
| myg | 1034 Laus 035

Immature
Mixéd 7 R+U EE fae unf oe |
Maures | June 30
| Peg? | mys | 1036 [amu | ose |
24 (231)

893
immature 9.98

omge [a f E
ESA

0.20

Durban, Natal, 3095.
037

July 9 EEE
July 12

P ur [nga | iir [onn] 0.19

Byron Bay, E Australia, 29°S

Iess | my4 | 990 Tool 100 |

geed Loes 10.35 035

[Mies o Kol July is | 1135* [264200] 0374
duger | ms Leg 06 |
yo | 1075 | 2.631689) | 020

Norfolk Ts., 29°S

Immature July 25
Mixed $ El Aug 2 | 13.98* |3.30(99e| oso |

of recognisilbe trends within this grouping, the
data were pooled into two main groups of six
degree lines including 55°S-60°S and 61°S-66°S
(both inclusive) within each sector,

A few humpbacks only were caught within
these sectors and latitudes duirng the 1932-1933
and 1933-1934 seasons, so these were eliminated
in an attempt Lo reduce error due to seasonal
variation. After these steps, the remaining sam-
ples between 5595 and 66°S included 5030
humpbacks in the sectors between 20°S to 3975

= Is, E pus m

Lee Lal Eed

mean limits

mem nøly 11 | 1072 0.25
[Mixed 9 R+U| July 15

Carnarvon, WA, 25°S
| Lat? | unez | 897 | 2091523 | 008 |
[ immaure | luyis | 1083 [3100636)| 035 |
EES
| Mawes | July28 | 1320 |2.81(2209[ 0.12 |
ENE
| Point Ces, WA, 22.5%
La? | myo | wa [210076 | 031 |

Immature 0,22
[ Resting © | auly28 | 1323 198075)
| Peg ? | Aug! | 1380 | 28340159)
ar roret ie
Madagascar, 1675- 2995

mme | døds | 1320 12950529] 025

0.27

Mature d
meet

iama T Are] Aug I1 em fimus] om |
Aug 2 Ces Dames
BEE EES

and 3938 in the sector between 80°E and 99*E

The weekly catch results are shown in Tubles

15-18, and the fortnightly numbers and percent-
age groupings in Figs 26-27.

The results appear to be so inconsistenl,
whether considered by the above sectors and
latitude groupings, or à variety of subdivisions
within these, that the writer is unable to suggest
any general trend in the order of arrival and
departure of different humpback categories. It is
clear that in Antarctic waters humpbacks, in gen-
eral, are caught earlier in the sector of warmer
waters (80°E to 99°E) then in the colder water sector
and earlier in low latitudes (55°S to 60°S) than in
higher latitudes (see also Table 19), However,

TEMPORAL SEGREGATION OF HUMPBACK WHALES

Table 13. Weekly catches of specified humpbacks,
1957-1961. Tonga, 22°S.

===

Immature | =
ESSET :
EG S a
z E E
E z z
24/7 1 l
31/7 1 2 3
7/8 2 1 3
14/8 5 2 7
21/8 3 7 10
28/8 3 8 11
4/9 4 2 6
11/9 2 9 1
18/9 I 12 13
25/9 1 2 3
2/10 2 3 5
9/10 4 4
16/10 i 1
23/10 1 2 3
30/10
6/11 1 1
Total | 26 56 T | &

even this finding may be more a description of the
movement of factory ships in relation to the edge
of the pack ice than a true indication of humpback
movements.

If differences in actual mean dates for all hump-
backs at each of the four regions are disregarded
for the purposes of discussion, it is still clear from
Figs 26-27 and Table 19, that there are very small
and inconsistent differences in the relative timing
of humpback categories within each region.

Based on mean dates, there is a maximum
difference of nine days between the earliest and
last of the four identifiable humpback categories.
Even within this limited period, the apparent se-

quence on mean dates is different within each of

the four groupings (Table 19). The catch compo-
sition also differs substantially in fortnightly fre-
quency (Figs 26-27). If the catch is an adequate
sample of the local humpback composition, then
the above results indicate that movements of dif-
ferent humpback categories in Antarctic waters
between late November and mid-March are
mainly random.

There are several factors that may contribute to
this negative result. In Antarctic waters, the
movements of factory ships and chasers have
been directed primarily towards catching fin and

131

TABLE 14. Estimated mean date with standard devia-
tion and the 9546 confidence limits of the mean for
specified categories of southbound humpbacks.

Confidence
limits

Week
code mean

Mean date

Tonga, 2295

Immature

3.36 (28)
3.01 (56)
3.29 (84)

0.72

Moreton Is., E
Mixed 9 R+U

1.83 (183)

Immature Sept 8 1.95 (249)
Mature d Sept 13 2.22 (545) 0.18
All 2.13 (977) 0.13

Byron Bay, E Australia, 29°S

Mixed ? R+U 1.70 (100) 0.58
Immature 2.20 (57) 0.45
Mature d Sept 22 21.22 2.47 (191) 0.35
2.48 (348) | 027

Norfolk Is., 295

Mixed 9 ec ‘Sept24 | 21.44 [186060)] 030
Immature | Sept24 | 2152 | 1.86(66) | 046
Matured | Oct8 | 23.52 |1.86(213)| 025

Al | Oet1 | 2246 | 2.12(439)| 020

Great Barrier Is., NZ, 36°S
Immature | Oe? 2339 | 23603 | 141 |

Mixed $ Bail | Oct 8 23.47

blue whales with humpbacks as a rather second-
ary objective. In coastal areas such as the offshore
waters from Durban, it has already been argued
that such selection might significantly affect
catch composition. However, there is no substan-
tial evidence to show that such differential selec-
tion will invalidate humpback sampling on the
open seas distant from shore lines.

As a more simple hypothesis, I suggest that the
catch sample has been taken after a large propor-
tion of the humpbacks had already reached Ant-
arctic waters, and that sampling ended before any
significant emigration from the region had com-
menced.

The season during which southbound hump-
backs pass many tropical and temperate localities
is known, and at an overall migration rate of 15?
per month (Dawbin, 1966), it can be shown that
in most sectors more than half of all humpbacks
could be expected south of 55°S by late Novem-
ber. If the sequence followed in warmer waters is
still maintained during progress into high lati-
tudes, then all main categories of humpbacks

should already be represented south of 55*S be-
fore December, with the possible exception of
cows accompanied by calves. Cows accompa-
nied by calves receive full protection, and there-
fore do not contribute to the catch. The absence
of samples during much of the period of entry into
high latitudes would contribute greatly to a mask-
ing of trends.

The period of emigration from Antarctic waters
as calculated from the time of appearance of
northbound humpbacks at lower latitudes, indi-
cates that very few animals are likely to pass north
of 55°S until late April, i.e., after catch sampling
in high latitudes has ceased. Similar calculations
for 60°S and other high latitudes are consistent
with the view that the catch sampling in Antarctic
waters during 1934-1939 occurred during months
in which entry and exit of humpbacks was too
small to become evident.

LAT. s?

co
so

IMMATURE
MATURE MALES
ENO OF LACTATON
RESTING

moten pesing | FEMALES
& LACTATING

LATE PREGNANT

CONGO

SSS

MADAGASCAR lô-

St CLOATES 22
WA

SARA

CARNARVON 25
WA,

I
EEN LEE
[

MEMOIRS OF THE QUEENSLAND MUSEUM

Catches were made during considerably longer
seasons in the Falkland Island Dependencies re-
gion 1910 to 1917. Data from these catches have
been examined, but they include such wide dif-
ferences between individual companies operat-
ing in proximity to each other that the writer has
failed to establish consistent trends.

It should be noted that these early records were
made at å time when many whales were dismem-
bered alongside anchored factory ships, thus add-
ing difficulties in obtaining accurate length
measurements, and complicating any examina-
tion for foetuses. Inspectors concerned primarily
with catch examination were not required by
International regulations until 1937.

CONCLUSIONS

Northbound humpbacks have been sampled by
catches between 41°S and 175 and include mate-

LOTT 4071

1819

TT D TET

1996

COO
EE c

SNNNNNNNNNNNNNNNN

Treneren myr

4523
NETT

ESS SSS

MORETON IS. 27|
E.A.

NORFOLK IS. 2

BYRON BAY 29
EA

QURBAN 30)

ALBANY as
W.A

SSS I

GT. BARRIER IS. 36
N.I.

CON
V IM,
AGER, n ELLA

D

COOK STRAIT 4
NI

SS

EIES

VIVI VEI)

a a
D mees

sener

5217

COT ` TEE
877A
in NE ER"

ESS

rpm

1922

FIG. 23, The period of passage of specified northbound humpbacks as shown by catch composition in temperate
and tropical waters. Mean date for each category shown as vertical white line. 95% confidence limits for the
mean shown in black. One standard deviation either side of the mean shaded.

TEMPORAL SEGREGATION OF HUMPBACK WHALES

TEEN

NUMBER OF WHALES

UD 15/5 29/5 (2/6 26/6 10 24/7 21/8 4h

FORTNIGHTS COMMENCING......

18/9

IMMATURE

EARLY LACTATING
FEMALES

2/0 16/10 30/10

FIG. 24. Fortnightly catches of southbound immature humpbacks and lactating females, 1957-1961. Tonga, 22°S.

MIXED RESTING &
IMMATURE P de PREGNANT FEMALES
ECH MATURE MALES DID EARLY LACTATING
LAT. S?
TONGA 22| EL i pU

Cm [ Enn

MORETON IS. 27
EA.

BYRON BAY 29
EA
NORFOLK IS. 29

GT. BARRIER IS. 36
N.Z.

198 178 24/8 3/8 79 we 2/9 289 5/0

12/10

IYO 29/0 2l

84

977

348

FIG. 25. The period of passage of specified southbound humpbacks as shown by catch composition in temperate
and tropical waters. Mean date for each category shown as vertical white line. 95% confidence limits for the

mean shown in black. One standard deviation either side of the mean shaded.

134

TABLE 15. Weekly catches of specified humpbacks,
1934/35 to 1937/38. Longitudes 20°E-39°E, latitudes
55°S-60°S.

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 16. Weekly catches of specified humpbacks,
1934/35 to 1937/38. Longitudes 20°E-39°E, latitudes
61°S-66°S.

rial relating to at least four segregated breeding
stocks. These stocks include one or more groups
that pass between east Australia and Tonga, one
along west Australia, another past Madagascar
and another along West Africa to the Congo.
Despite the sampling differences between locali-
ties as described in the text, catches from all these
stocks share similarities in the time sequence
followed by specified categories of humpbacks
(Fig. 23, Table 12).

Females at the end of lactation and still accom-
panied by weaning ‘yearlings’ occur earliest at all
localities from which the data allow identification
of this group. At Cook Strait, Albany, Carnarvon
and Point Cloates, they occur at an average of 12
days earlier than immature whales. There is no
regular difference in timing between immature
males and immature females. Mature males fol-
low immature whales at Cook Strait, Great Bar-
rier Island, Albany, Durban, Carnarvon, Point
Cloates, Madagascar and Congo by an average of
8 days later. As described previously, there is no
significant difference in timing between these
categories at the three stations where quotas and

Immature Mature | Immature Mature
e o o P ^ or o E
E D . G E 2 ` E
$ *o o 5 to 2 ob 8 $ *o o 5 fo 2 E] 8
3 E + È B. E: E M È a
5 E m E 5 E P E
E E 2 E E -
= Zz a » x «x
à | St | å
6/11 1 1 412 1 (17? 4
wit} 3 | 5 ad -x- He 11/12 I I 1
201 1 1 1 18/2] 1 1 1
ami} 6 | 7 | 13 | 2 | 24| 2 | 2512 | 1 1 | nl LEN
412 | 15 | 17 | 32 | 88 | 112 | 11 | 243 m 1 | 9 | 0 | 6 | 11 1 3 | 3
1/12| 63 | 104 | 167 | 186 | 186 | 33 | 572 si | 2 | 3 | 5 | 7 | 9 | 4 | 2s
18/12| 86 | 77 | 163 | 164 | 121 | 20 | 468 in} a | 2 | 3 | 9 | 12 | 7 | 31
25/12| 49 | 51 | 100 | 182 | 118 | 29 | 429 21| 6 | 5 | 11 | 25 | 37 | 2 | 85
Vi | 53 | 56 | 109 | 156 | 96 | 30 | 391 29) | 9 | 10 | 19 | 26 | 19 | I5 | 79
an | 47 | 66 | 113 | 156 | 102 | 19 | 390 52 | 29 | 37 | 66 | 119 | 93 | 30 | 308
isl | 34 | 30 | 64 | 106 | 73 | 21 | 264 122 | 20 | 32 | 52 | 60 | 65 | 26 | 203
228 | 15 | 16 | 31 | 46 | 49 | 20 | 146 19/2 | 40 | 41 | 81 | 124 | 66 | 28 | 299
295 | 30 | 29 | 59 | 207 | 101 | 57 | 424 2602| 5 | 6 | 11 | 28 | 8 | 6 | 53
sa | 5 | 13 | 18 | 42 | 20 | 16 | 96 58 | 10 | 8 | 18 | 28 | 15 66
172 | 1 1 SL a9) de L 23| 5 | 2 | 7 | 13 | 12 | 2 | 34
192 | 9 | 11 | 20 | 11 | 9 | 8 | 48 93| 5 | 4 | 9 | 4 | 1 14
262 | 11 | 17 | 28 | 20 | 17 | 4 | e 26/3 62 | 19 | 12 | 102
58 | 13 | 14 | 27 | 10 | 15 | 1 | 53 Total | 135 | 162 | 297 | sis | 376 | 151 | 1348 |
12/5 1 1 |2
Total | 440 | 514 | 954 | 1403 | 1049 | 276 | 3682 | intense selection for large animals has resulted in

marked under-representation of immature ani-
mals (Byron Bay, Norfolk Island, Moreton Is-

TABLE 17. Weekly catches of specified humpbacks,
1934/35 to 1937/38. Longitudes 80°E-99°E, latitudes
55°S-60°S.

Immature Mature

a or oF Mi

Blof o] El] o| zial?
EI EI È a
5 E e E
E S E
16/11 2 3 5 3 10 16 34
13/11 1 6 18 25
20/11 8 15 16 49 28 108
27/11 5 23 28 21 84 13 146
4/12 26 31 57 104 138 50 344
11/12 19 32 51 100 85 49 285
18/12 | 21 32 53 132 146 56 387
25/2 4 12 16 55 64 51 186
1/1 4 9 13 30 45 33 121
8/1 4 18 22 43 45 9 119
Total 93 167 260 505 667 323 | 1755

TEMPORAL SEGREGATION OF HUMPBACK WHALES 13

TABLE 18. Weekly catches of specified humpbacks,
1934/35 to 1937/38. Longitudes 80*E-99*E, latitudes
61°S-66°S.

tn

TABLE 19. Estimated mean date with standard devia-
tion and the 95% confidence limits of the mean for
specified categories of Antarctic humpbacks.

TM Mature Mean date Week SD (n) Confidence:
" œ m E code mean limits
2Zleisl Biel äl al? 80°-99°E, 55°-60°S
& E e | & | $ : 233 (323) | 026
: E d Mem Colonel
ei == s * Immature | ei [isg] 025 |
27111 1 1 Matre å | Dec 18
4/12 3 3 2 1 l 7 621 |2020755| 0.10
HA2| 3 6 9 I 2 9 | 2 80°-99°E, 61°-66°S
giz} 12 | 20 | 32 | 36 | 17 | 23 | 108 Preg. 9 2.92 (314)
25112 | 16 | 17 | 33 | 59 | 53 | 23 | 168 Mature å
mi | 12 | 11 | 23 | 38 | 18 | 35 | 114 Immature | Jan 27 12.34 0.25
an | 17 | 25 | 42 | 55 | 65 | 42 | 204 Non-preg. 9 | Jan28 | 12.51 [2.81 (589)
ist | 28 | 33 | 61 | 64 | 53 | 39 | 217 Al ` | Jan26 1222 |290Q185| 0.12
98 | 100 | 72 | 42 | 312 20*-39%E, 55°-60°S
55 | 78 | 65 | 33 | 231 8.77 |3.17 (1049)
57 | 100 | 61 | 23 | 241 3.33 (954)
53 | 84 | 95 | 14 | 246 3.01 (1403)
74 | 62 | 54 | 12 | 202 3.24 (276)
24 | 21 | 21 | 9 | 75 All 3.18 (3682)
s | 13 | 9 8 | 35 20°-39°E, 61°-66°S
569 | 713 | 587 | 314 | 2183 | | Non-preg. 9 nen I 14.55 2
Preg. 9 Feb 13 1477 | 2.73051)
land). However, there is no case in which mature Immature | Feb 14 1488 | 2.49 (297)
males have preceded immature whales by an Mature 3 Feb 19 15.62 2.85 (515)
amount that is significant at the 5% level. All a Feb 15 1506 |2.76(1339)

Mature females that are predominantly in the
resting condition have occurred at an average of
only 3 days after mature males so can be regarded
as following a similar time sequence to the latter.
The apparent timing of mixed females is variable
due to the unknown but certainly different pro-
portions of late lactating and resting animals in-
cluded in this group at different localities. For this
reason it is not strictly comparable with the more
accurately specified categories. In general, mixed
females have a mean date which falls after those
for immature whales and before those for preg-
nant females. Pregnant females have been repre-
sented at all localities except Norfolk Island and
Durban. In all cases where they are represented,
they occur as the last northbound category. The
average intervals are 11 days after mature males,
19 days after immature animals and 31 days after
lactating females.

Due to sampling errors discussed previously,
the above time intervals are not regarded as hav-
ing the precision implied by expressing the results
in days, but the results should be a reasonable
indication of the order of time difference between
categories. For the reasons given previously, it is

probable that the differences described here are
minimal and likely to be greater in the migrating
population than shown by catch samples.
Southbound humpbacks form a smaller and
more complex sample which has been discussed
at greater length elsewhere (Dawbin, 1966).
Based on mean dates, mixed females together
with immature animals, form the earliest south-
bound categories (Fig. 25, Table 14). In these
cases, mixed females include those in very early
pregnancy unseparated from resting females.
Many of these females would correspond to those
that travelled north early when at the end of
lactation. If these females also returned south
early, then many would be incorrectly included
amongst the northbound whales which overlap
with southbound up to the time that 50% are
sighted travelling north and 5046 travelling south.
This, combined with a lack of data on which to
separate early pregnant from resting females im-
pedes attempts to estimate a mean time interval
for these females (see also Dawbin, 1966), but
they clearly precede mature males and cows with

136

— oiri

LONGITUDE S
LATITUDES. E

gt e min

MEMOIRS OF THE QUEENSLAND MUSEUM

LI rl
$---.

— t—

LONGITUDES SO
LATITUDES ,.... ët 00"

FIG. 26. Fortnightly catches of specified humpback categories, Antarctic, 1934/35 to 1938/39 seasons. Longi-
tudes 20^E to 39°E and 80°E to 99°E, latitudes 55°S to 60*S and 61°S to 6678 for each longitude group.

calves and the data are not inconsistent with the
view that many may precede immature hump-
backs.

Immature humpbacks travel south about 10
days earlier than mature males and the latter in
turn precede cows accompanied by young calves,
Subtraction of the 10 days’ difference between
immature animals and mature males as above,
from the 16 days' difference between immature
animals and cows with calves at Tonga appears to
indicate a difference of 6 days only (figure used
by Dawbin, 1966) between mature males and
cows with calves. However, reasons for believing
that the calculated interval at Tonga is an under-
estimate have been discussed previously. In addi-
tion, the observations of Chittleborough near
Point Cloates 2295 (1953) and Bryon Bay 28°S
(1962) together with those of the writer in
Foveaux Strait 4875, have shown that cows ac-
companied by calves comprise considerably
more than 5066 of all humpbacks at these latitudes
during the final three or four weeks of the south-
ward migration past each locality. Thus any non-

lactating females and late immature animals to-
gether with mature males, are considerably out-
numbered hy cows and calves at this stage. The
latter, which comprise the females that travelled
north last as pregnant animals, are clearly the last
category to return south. The available data on the
time sequence of southbound humpbacks is not
inconsistent with the view that humpbacks return
south in approximately the same sequence as they
follow during the migration north.

Antarctic catch samples, when subdivided ac-
cording to latitudes and sectors of longitude, in-
dicate differing time sequences between samples.
There is, therefore, no consistent trend recognis-
able in the present analysis of Antarctic catches,
Since most of the samples were obtained after late
November, it seems probable that they were ob-
tained from a population that had already
stabilised considerably as a result of the arrival of
most catchable categories into the Antarctic sam-
pling zone. This hypothesis is consistent with the
expected arrival dates based on known mean
dates in lower latitudes and a migration rate of

TEMPORAL SEGREGATION OF HUMPBACK WHALES

rena cr mm. ue

zim

LONGTUDES
LATITUDES er-se

PeUrwmwor De wm cams

137

FIG. 27, Fortnightly percentages of specified humpback categories, antarctic, 1934/35 to 1938/39 seasons.
Longitudes 20°E to 39°E and 80°E to 99°E, latitudes 55°S to 60°S and 61°S to 6678 for each longitude group.

about 15? per month. On the same evidence, no
consistent changes in composition would be ex-
pected before the northward departure of some
humpbacks in mid to late April, i.e., considerably
later than the period during which catch sampling
occurred. In the absence of consistent trends in
Antarctic samples, the order in which the speci-
fied humpback categories travel past each local-
ity and the time intervals berween the earliest
and each of the later categories have been esti-
mated exclusively from temperate and tropical
data.

The estimated humpback sequences differ
markedly from those of fin whales described by
Laws (1961) who discusses extensive Antarctic
data and the findings of previous workers, espe-
cially Mackintosh (1942). Laws concludes that
both the north and south migrations of older fin
whlaes and of pregnant females are in advance of
those of other groups, and that sexually immature
animals are later. Pregnant humpbacks appear lo
travel south in advance of other humpbacks, but
return north after other categories. Immature

humpbacks are the second group in both south
and north migrations, followed by mature males
together with resting females, while lactating fe-
males travel south last and return north in advance
of other humpbacks.

There is no consistent evidence that different
humpback categories either migrate at different
speeds or stay for significantly different lengths
of time in the breeding areas. The data indicate
that most categories depart also from the feeding
areas in the same order as they arrive. Most
categories therefore spend more or less equal
periods in Antarctic waters. The main exception
are breeding females which, when pregnant,
spend a prolonged period in the feeding regions
and, when lactating, spend an abridged period,
while resting females stay for approximately the
same length of time as mature males and imma-
ture humpbacks.

The implications of these findings in relation to
the breeding cycle and the environmental condi-
Dons encountered during migration are discussed
more full y elsewhere (Dawbin, 1966).

138

ACKNOWLEDGEMENTS

Mr E. Vangstein with the permission of the
managers of southern hemisphere whaling com-
panies has provided a major part of the post-war
daily catch data. Dr R.G. Chittleborough has sup-
plied much supplementary data on Australian
catches. Professor J.T. Rudd supplied catch data
on pre-war temperate and tropical catches, and Dr
F.C. Fraser on catches in the Falkland Island
Dependencies area. Dr N.A. Mackintosh has en-
couraged and discussed the study throughout. Mr
G.T. & J.H. Perano and staff at the Cook Strait
Whaling Station have assisted greatly in the col-
lection of much material as well as catch data
from New Zealand catches. I have greatly appre-
ciated this assistance, and I wish to thank all those
who have helped in this study.

LITERATURE CITED

ANGOT, M. 1951. Report scientifique sur les expedi-
tions baleinieres autour de Madagascar. (Saisons
1949 et 1950). Mem. Inst. Sci. Madagascar 6:
439-486.

CHITTLEBOROUGH, R.G. 1953. Aerial observations
on the humpback whale Megaptera nodosa

MEMOIRS OF THE QUEENSLAND MUSEUM

(Bonnateere), with notes on other species. Aust. J.
Mar. & Freshwater Res. 4: 219-226.

1958. Australian catches of humpback whales 1957.
C.S.LR.O. Aust. Div. Fish. Oceanogr. Rep. 17:
1-21.

1962, Australian catches of humpback whales 1961.
C. S.I R.O. Aust. Div. Fish. Oceanogr. Rep. 34:
1-13.

DAWBIN, W.H. 1956, The migrations of humpback
whales which pass the New Zealand coast. Trans.
Roy. Soc. N.Z. 84: 147-196.

1960. An analysis of the New Zealand catches of
humpback whales from 1947-1958. Norsk.
Hvalf. Tid. 2: 61-75.

1966. The seasonal migratory cycle of humpback
whales. Pp 145-170. In Norris, K.S. (ed.)
"Whales, dolphins and porpoises'. (Univ. Calif.
Press: Berkley and Los Angeles).

HUBBS, C.L. & HUBBS, C. 1953. An improved graph-
ical analysis and comparison of series of samples.
Syst. Zool. 2: 49-57.

LAWS, R.M. 1961. Reproduction, growth and age of
plen fin whales. Discovery Reports 31: 327-
486.

MACKINTOSH, N.A. 1942. The southern stocks of
whalebone whales. Discovery Reports 22: 197-
300.

MATTHEWS, L.H. 1937. The humpback whale,
Megaptera nodosa. Discovery Reports 17: 7-92.

TWO NEW GENERA OF ACANTHODRILINAE (MEGASCOLECIDAE,
OLIGOCHAETA) FROM CAPE YORK PENINSULA AND THE TORRES STRAIT

G.R. DYNE

Dyne. G.R. 1997 06 30: Two new genera of Acunthodrilinac (Megascolecidac, Oligochacta)
from Cape York Peninsula and the Torres Strait, Memoirs of the Queensland Museum 42(1):
139-157. Brisbane. ISSN 0079-8835.

Two new meronephric genera of Australian acanthodrile earthworms from Cape York
Peninsula and the Torres Strait are described: Neodiplorrema gen nov., with seven new
species, and Torresiella gen nov. with å single new species. All are considered to be local
derivatives of the holonephric endemic Australian genus Diplotrema. Six species of
Neodiplatrema show the acanthodrilin condition of the male genital terminalia, with male
pores on segment XVIII and a pair of prostate pores on each of segments X VIL and XIX, but
one species exhibits the mieroscoleeim reduction in which male pores have been translocated
forwards to unite with a single pair of prostate pores, in XVIL, Two species of Neodiplorrema
have modifications of the nephridial system which pose difficulties for the existing classifi-
cation of the Megascolecidae. Torresiella shows the rare balantine reduction in which the
male pores have been translocated posteriorly to unite wilh a single pair of prostate pores,

in XIX. [ ] Acanthodrilinae. Megascolecidae, Neodiplotrema, Torresiella., aligochaete.

G.R. Dyne, Environment Australia, GPO Box 787 Canberra ACT 2600, Australia; 24 April

1997.

Prior to the present work, the Australian genera
of earthworms of the subfamily Acanthodrilinae
have been exclusively holonephric, with a pair of
stomate nephridia in each typical body segment.
In contrast, Lee (1959, 1970) clearly demon-
strated close affinity between holonephric New
Zealand acanthodrile genera and those with more
than one pair of nephridia per segment, the
meronephric condition, and concluded that local
transition from holonephry to meronephry had
occurred. Two meronephric genera of Australian
acanthodriles are here described which are con-
sidered to be similarly localised derivatives of the
endemic holonephric genus Diplorrema.

Diplotrema has been demonstrated to have an
extensive range, from the vicinity of Narrabri,
New South Wales, to northern Queensland and
across northern Australia to the Kimberley region
of Western Australia. (Mackenzie & Dyne, 1991:
Jamieson & Dyne, 1976; Dyne, 1979). The pre-
cise northerly limit of the genus in Cape York
Peninsula is unknown, the most northerly defini-
tive record coming from the vicinity of Weipa
(Dyne, 1984). It is thus uncertain whether
Neodiplatrema effectively replaces Diplotrema
from a Certain point northwards. However, the
two genera are sympatric in in the Cape Melville
National Park and Mcllwraith Range (Jamieson,
1997, this volume).

Despite recent efforts to improve knowledge of
the biota of the region generally (e.g.. Glasco et
al., 1995), the earthworm fauna of Cape York

Peninsula and the Torres Strait remains poorlv
understood. Apart from species of the
pheretimoid genus Begemius described by Easton
(1982), this paper constitutes the only account of
endemic earthworm taxa from the region.

Abbreviations used, ANIC — Australian National
Insect Collection, coll. = collected by, gm = gen-
ital marking, GD = Author's collection, H = ho-
lotype, P = paratype, pr. p = prostalic porophore,
p.5.p. = penial seta pore, QM = Queensland Mu-
seum, Sp. p = spermathecal pore. Scale-bars =
Imm in all figures.

Neodiplotrema gen. nov.

DIAGNOSIS, Setae 8 per segment, Prostates 2
pairs, tubular, their pores on XVII and XIX, or,
exceptionally, a single pair, on XVII. Male pores
present in XVII, or, rarely (microscolecin condi-
tion), closely associated with a single pair of
prostatic pores on XVII. Spermathecal pores 2
pairs, in 7/8 and 8/9, or a single pair, in 8/9 only.
Gizzard very well-developed, in V. Calciferous
glands present or absent, Meronephric, avesicul-
ate; anterior tufted nephridia present, Holandric,
or, exceptionally, metandric. Testis-sacs absent.
Penial setae invariably present; genital setae usu-
ally present, but occasionally lacking.

DESCRIPTION. Small to large terrestrial worms
(48-465mm in length) with 130-500 segments.
Body circular in cross-section. Prostomium pro-

E

Tore

d
Bamaga

Neodiiouenia

FIG. 1. Map showing recorded occurrences of
Neodiplotrema and Torresiella.

epilobous to epilobous. Dorsal and median ven-
iral groove absent, Dorsal pores commencing
fram 8/9 to 11/12, Setae closely paired, com-
mencing on If; ventral intersetal distance (aa) not
greatly different from that (bc) separating the
lateral setae: ventral and dorsal setal couples of
similar width; dorsal median intersetal distance
(dd) > 45% of the body circumference in the
forebody. Setae of XVIII, or the ventral setal
couples lacking; ventral setal couples of XVII
and XIX modified as enlarged penial setae, or,
rarely, only those of XVII thus modified (in N.
deminutionis); ventral setal couples of IX modi-
fied as enlarged genital setae, or undifferentiated.
Nephropores numerous and usually inconspicu-
ous. Clitellum saddle-shaped, or annular, occupy-
ing XIII to XVI-XVII. Two pairs of prostatic
pores, equatorial on XVII and XIX, coincident
with the penial seta orifices, or (D. deminutionis)
à single pair of combined male and prostatic
pores on XVIL Male pores a pair of openings
in seminal grooves (the latter well-developed
or very faint), presetally, or far antenad, in XVIII
(rarely in the microscolecin arrangement), Acces-
sory genital markings present in some of the
segments VII-XXXII, usually intersegmental,
rarely absent. Female pores paired, presetal, me-
dian to lateral of ab, in XIV, Spermathecal pores

MEMOIRS OF THE QUEENSLAND MUSEUM

2 pairs, in 7/8 and 8/9, in or near ab, exceptionally
in 8/9 only.

Some pre-clitellar septa thickened. Dorsal
blood vessel single, continuous onto the pharynx;
last hearts in XIII, some of the commissurals
posteriad of and including X latero-oesophageal,
dorso-ventral commissural vessels commencing
in VI-VII; supra-oesophageal vessel limited to
the oésophagus; subneural vessel absent. Gizzard
large, often strongly muscular, in V. Calciferous
glands absent, or developed as sessile pouches in
the posterior oesophageal region. Intestine com-
mencing in XVI-XX, a dorsal typhlosole well
developed. Body astomate, exonephric; caudally
nephridial bodies enlarged, each with a pre-septal
funnel; extensive tufting present in all or some of
segments III-V, Enteronephry not developed.
Holandric, sperm funnels free in X and XL or
metandric, the funnels in X absent. Seminal ves-
icles present in EX and XII, or in XI and XIL
Prostatic glands tortuous tubes usually restricted
to their segment of origin; posterior pair usually
conspicuously smaller than the anterior set, or (D.
deminutionis) totally lacking; prostatic ducts
moderately long to long, muscularised, Penial
seta follicles usually with copulatory muscula-
ture. Ovaries (conjoined oocytic strings) and me-
dium-sized oviductal funnels present in XIII;
ovisacs. Spermathecae subequal, or the posterior
pair slightly the larger; the single diverticulum is
invariably sessile, and provided with nurnerous
intramural sperm chambers.

DISTRIBUTION, Cape York Peninsula: appar-
ently restricted to monsoonal semi-deciduous
vine-forests in the Lockerbie, Iron Range and
Weipa areas; islands of the Torres Strait (Fig. 1).

TYPE SPECIES. Neodiploirema tumida (designated).

ETYMOLOGY. Differing from Diplotrema in a novel
feature (meronephry).

REMARKS. The morphological similarities of
Neodiplotrema to the Australian acanthodriline
genus Diplotrema are so close, including even the
possession of anterior genital setae in some spe-
cies, that origin of the Neodiplotrema from the
latter seems indisputable.

List of species of NEODIPLOTREMA

1) N. deminutionis sp. NOV., 2) N, exigua sp.
nov., 3) N. lacisbrontoi sp. nov. 4) N. occidentalis
sp. nov., 5) N, raveni sp. nov., 6) N. tumida sp.
nov., 7) N. varionephrica sp. nov.

TWO NEW GENERA OF ACANTHODRILINAE 141

KEY TO SPECIES OF THE GENUS
NEODIPLOTREMA

|. Male apparatus acanthodrilin: 2 pairs of prostatic
pores in XVII and XIX, male pores separate, on
NANI, aphan s rer aco puede e Lote ANA 2
Male apparatus microscolecin: a single pair of
prostatic pores associated with the male pores on
RNI A p lI eden å e H N. deminutionis
2. Nephridia in the mid-body consisting of a promi-
nent megameronephridium, and 3 small
micromeronephridialbodies ........,.3
Nephridia in the mid-body uniform in size or
medianmost nephridium not greatly enlarged ` . 4
3. Genital setae present, intestine commencing in
ME «—. tege ine uS uua N. varionephrica
Genital setae absent, intestine commencing in XX
.sg di t. A EAA OF ene: N. exigua
4. Male organs metandric . . . - .. N. occidentalis
Male organs holandne (though there may be
some reductionin X) . - 21 arena ve rå 5
5. Genital seiae well-developed, in IX; prominent
genital markings present in the region XVIII-
XXI N. tumida
Genital setae absent or rudimentary. Genital
markings absent from the region XVILL-XXI . 6
6. Moderately large worms (150-200mm); ventral
setal couples of XVII lacking, oesophagus with
conspicuous outpouchings in XIV-XV
N. lacisbrontoi
Very large worms (>350mm); veniral setal couples
of XVIII present; oesophageal pouching absent
N. raveni

Neodiplotrema deminutionis sp. nov.
(Figs 2, 11B)

TYPE LOCALITY. 10°46'S, 142*34'E,, on the
eastern side of Lake Bronto, Cape York Peninsula, in
open eucalypt forest in dark sandy soil. Coll. R, Raven,
4 Feb 1975,

MATERIAL EXAMINED. HOLOTYPE AND 8
PARATYPES: ANIC GD.95.31.1

DESCRIPTION. Length 71+, 140mm. Width
2.9, 27mm. Segments 215+, 311 (Holotype, P2).
Uniformly circular in cross-section throughout;
pigmentless buff in alcohol, Prostomium prolob-
ous, first dorsal pre-9/10. Setae in regular longi-
tudinal rows, closely paired throughout; ventral
setal couples of XVIII normal, those of XVII
modified as enlarged penial setae; genital setae
lacking. Nephropores inconspicuous. Clitellum
not developed in any of the specimens available
for study. Combined male and prostatic pores in
distinct longitudinal slits on raised papillae, in

FIG. 2. Neodiplotrema deminutionis genital field,

XVII, coincident with the openings of the penal
setae, Serjal sectioning reveals that the vasa def-
erentia and prostatic ducts do not actually fuse,
but open contiguously al the common pore. Fe-
male pores visible as tiny, transverse slits preset-
ally. in ab, usually with swollen rims. Accessory
markings a series of unpaired, median, in-
trasegmental pads; a small ellipsoidal pad in IX
(H, P4-5); a larger one in X, extending across aa
(H, P1-2, P4-5, P8); a similarly sized swelling in
XI (H, P1, P6) (all the above markings have
shallow, transversely concave centres); a large

142

pad is located in XV, transverse width = aa (H,
Pl, P3, P5-6), with a larger marking in XVI,
across bb (all specimens excepting P7); another
series of similar markings is present in Lhe post-
genital region: a diminutive tumescence in XTX
(H, Pl, P6), a much larger swelling in XX (across
aa) (H, Pl, P6), similarly in XXI (H, P1-6, P8),
and a further small marking in XXII (H, PI, P2,
P3, P4, P5, P6, P8), In some of the above speci-
mens, the accessory markings are faintly devel-
oped.

Septa: 10/11, 11/12 slightly thickened, 6/7-9/10
moderately strongly muscularised, 5/6 slightly
thickened, encapsulating the gizzard. Dorsal
blood vessel single, continuous onto the pharynx.
Supra-oesophageal vessel present VII-XIX. ad-
herent to the roof of the oesophagus, and receiv-
ing vessels from oesophageal blood plexi,
particularly in XII-XIV, Last hearts in XT, those
m X-XID receiving thin connectives from the
dorsal vessel, and more strongly developed ones
from the supra-oesophageal vessels; commissur-
als Xll and XH much larger than the remainder,
which decrease in size anteriorly, those anterior
to X dorsoventral only, Gizzard large, shiny, but
compressible in V, with a slight dorsoventral
compression, Oesophagus rather narrow, con-
sincted intersegmentally, with conspicuous rugae
on its inner walls, and extending from VI to XVI.
Intestinal origin, commencing with abrupt expan-
sion. in XVIL(H). XVI? (P1); somewhat paler and
apparently less muscular åntertor of XXIV. An
extremely well-developed rugose dorsal
typhlosole commences in XXIL Meronephric
throughout; considerable tufting in IV, with many
highly coiled loops, and composite ducts sent 10
the pharynx; the remainder of the nephridia com-
prised of very numerous astomate exonephric
loops scattered on the body wall and septa, larger
and more numerous in the oesophageal region,
Caudally, the bodies again enlarged, each appar-
enily with a distinct preseptal nephrostome; no
enteronephry demonstrable.

Holandric; 2 pairs of small, iridescent funnels
iu X and XI, attached ventrally to septa 10/11 and
11/12; large, subequal, racemose seminal vesicles
in XI und XII, antertorly attached to septa 11/12
and 12/13, composed of large, conspicuous
loculi, Vasa deferentia obvious as closely paired,
slightly iridescent tubes tortuously adherent ta
the body wall on either side of the oesophagus;
the paired ducts become fused in the parietes, in
XVIL Prostatic glands a pair of fattened, loosely
coiled tubular organs lying in segment XVII with
long, muscular (shiny) ducts opening to the exte-

MEMOIRS OF THE QUEENSLAND MUSEUM

rior slightly ventral of the vasa delerentia, Penal
sela follicles large, gently curving, copulatory
musculature well developed, inserted near the
mid-dorsal line; the setae with a gradual bend, the
shaft divided into an ectal region devoid of orna-
mentation, with an intervening secuon bearing
scattered solitary or clustered thorn-like spines
with rather broad bases, and narrowing to a fine
point (these stand out at an angle to the shaft); the
entire ectal portion is approximately equal to the
ornamented section, and these together constitute
about half of the total length of the seta. Length
mature seta 2.35mm; midshaft diameter 25.5jm
(mean of 3). Ovaries, consisting of thin sheaves
of small oocytes, together with medium-sized
diaphanous funnels, are present XIII, ovisacs ab-
sent. Spermathecae a single pair, divided into a
stalked, saceiform ampulla, and a broadly U-
shaped diverticulum which is completely sessile,
and which occupies the entire dorsal aspect of the
short duet; the walls of the diverticulum are stud-
ded with numerous iridescent intramural sperm
chambers, the ampullal ‘stem’ is swollen at its
junction with the diverticulum, Length tight sper-
matheca of IX 2.6mm (apex of ampulla to pare),
Genital seta follicles absent.

ETYMOLOGY. Referring to the diminished
imicrosenlecin] condihion

REMARKS. As this species is the only known
microscolecin Neudiplorrema, identification
should not prove difficult, particularly when the
distinctive array of genital markings is alsa con-
sidered. The lalter are atypical of the genus in
being intrasegmental. N, deminutionis lives in
sympatry with N. /acisbrontoi and N. vari-
oneplirica, but neither of these species appears to
be convincing as an acanthodrilin precursor of the
former.

Neodiplotrema exigua sp. nov.
(Figs 3, LIC)

TYPE LOCALITY 10485, 14228'E, Lockerhie
East. soil over rocky substrate, in dense semi-decidu-
ous vine-forest. Coll. R, Raven, 1 Feb 1975,

MATERIAL EXAMINED. HOLOTYPE. QMGH2889.
PARATYPE, QMGH2890. OTHER MATERIAL.
(0°48'S, [42*28'E, Lockerbie Enst, tinder logs near
‘Mango Tree camp’, Coll. R. Raven, 30 Jan 1975 (3
immature-semi-mature specimens, not designated as
types, ANICGD95,4,3)

DESCRIPTION. Length 48, 64mm. Width (mid-
clitellar) 1. 7mm, Segments 149, 136 (Holotype,
Pl), Uniformly cireular in cross-section,

TWO NEW GENERA OF ACANTHODRILINAE

FIG. 3, Neodiploirema exigua genital field.

pigmentless buff in alcohol. Prostomium prolob-
ous; first dorsal pore 11/12 (imperforate), Setae 8
per segment, commencing in II; ventral setal cou-
ples of XVIII present; those in XVII and XIX
modified as enlarged penial setae; genital setae
lacking. nephropores not visible externally,

143

Clitellum very faint, barely distinguishable. Male
pores located presetally, in definite seminal
grooves, on. XVIII, lateral of b-lines. Prostatic
porophores 2 pairs, in XVII and XIX, each on a
distinct papilla, and intimately associated with
(usually) protuberant penial setae, The slightly
convex male field is delimited laterally hy the
seminal grooves (joining the prostatic pores of a
side), with the prostatic porophores defining its
corners. Female pores small slits presetal in XIV,
in ab. Spermathecal pores 2 pairs of simple de-
pressions, m 7/8 and 8/9, in ab. Accessory mark-
ings 2 pairs of rounded tumescences with oculate
centres present intersegmentally, in 10/11 and
11/12. A further series of three more closely
paired markings (of similar shape) present in
14/15 and 16/17, shghtly lateral of b-lines; the
markings in 14/15 are separated by a distinct
depression, the remainder to.a much lesser extent;
a diffuse and indistinct pad extending from the
posterior edge of the male field to the anterior
portion of segment XXTis seen to be of width bb;
2 further unpaired median markings are located
in 21/22 and 22/23, the former the smaller, with
à multi-oculate central area, the marking in 22/23
longer, almost paired, with central depressions.
Septa: 5/6 delicate, 6/7-7/8 slightly thickened,
8/9-12/13 moderately muscularised, 13/14
slightly affected, remainder diaphanous. Dorsal
blood vessel single, continuous onto the pharynx;
last hearts in XIII, those in X-XUI considerably
larget than the remainder, with connectives from
both the dorsal and supra-oesophageal vessels;
the latter vessel moderately developed, its limits
not determinable. Gizzard large, though com-
pressible, in V; oesophagus narrow, VI-XIX, but
with three pairs of obvious pouches in XVI-
XVIII, that closely resemble stalked calciferous
glands; these are complexly vascularised, and
contain conspicuous lamellae; the three pouches
on each side of the oesophagus are clearly
interconnected, but communication of cach of the
pouches independently to the oesophagus could
not be demonstrated. Intestinal origin XX, a ro-
bust, though flattened, dorsal typhlosole is pres-
ent after XXV. Meronephric; the mid-body
segments each containing a pair of large
megameronephridia with preseptal nephro-
stomes, their terminal ducts discharging to a pair
of ventral, longitudinal ureters that run the entire
length of the body, opening at the anus; the
megameronephridia decrease gradually in size
posteriad, and disappear abruptly approximately
25 segments from the posterior end; in addition
to the above, each intestinal segment contains 3

pairs of small, astomate, exonephric loops, dis-
tributed evenly on each side of the nerve-cord, the
dorsalmost body just below the mid-dorsal line;
these persist (perhaps with some slight trend to-
wards a size increase) to the caudal extremity. A
large pair of tufted nephridial bodies is present in
V, with a much smaller set in VI.

Holandric; large, iridescent spermatic funnels
and sperm masses seen in X and XI; 2 pairs of
palmate seminal vesicles composed of large,
slightly iridescent loculi present in IX and XII,
the former somewhat smaller and compressed.
Prostatic glands 2 pairs of simple tubular struc-
tures in XVII and XIX, the anterior pair the larger;
the ducts are weakly muscular, and short. Penial
seta follicles proportionately large for the species,
each follicle with a distinct retractor ligament
inserted on the dorsolateral aspect of the body
wall. The setae fairly flat, gently curving, the ectal
region with a distinct sinusoidal bend; the ectal
1/4-1/3 is ornamented with clusters of 1-4 irreg-
ular, jagged teeth. Length mature seta 2.4mm;
midshaft diameter 53.4um (mean of 3). Ovaries,
comprising several strings of medium-sized oo-
cytes, together with small, pleated oviducal fun-
nels, present in XIII; minute rounded structures
on the posterior face of septum 13/14 are ques-
tionably ovisacs. Spermathecae rather small,
each organ consisting of an ovoid ampulla and
sacciform, blunt diverticulum (with some irides-
cence); duct very short. Length right spermatheca
of IX 0.6mm (base of ampulla to pore). Genital
seta follicles absent.

ETYMOLOGY. Referring to the distinctive nephridial
system.

REMARKS. This small species has undoubted
affinities with N. varionephrica, both species
possessing a remarkable nephridial system which
incorporates a combination of micromeronephri-
dia and megameronephridia, the latter with lon-
gitudinal ureters. This arrangement (as described
in detail above) is reminiscent of that described
by Bahl (1942) for the dichogastrin
Hoplochaetella khandalensis, though in that spe-
cies, the longitudinal ureters are located dorsally,
and the megameronephridia continue to the ex-
treme caudal segments. Bahl, in summarising
nephridial modifications in earthworms, de-
scribes the condition whereby the common excre-
tory canals open at the junction of the body wall
and the gut (i.e., the anus) as ‘the first steps, so to
speak, in the enteronephric direction’ (Bahl
(1947). An anatomical arrangement of the excre-
tory system whereby longitudinal ureters termi-

MEMOIRS OF THE QUEENSLAND MUSEUM

nate in the proctodaeal region is also seen in the
Western Australian genus Austrohoplochaetella
and the African species, Millsonia anomala.
Jamieson (1974) detected significant intraspe-
cific variation in the occurrence of meronephry
in the Tasmanian species Cryptodrilus poly-
nephricus, a further indication of the lability of
this character-state.

The occurrence of this relatively advanced ne-
phridial system in Neodiplotrema raises problems
for the existing classification (Jamieson, 1971),
in which, by definition, N. varionephrica and N.
exigua would be placed in the Dichogastrini,
within a different subfamily to that in which the
bulk of Neodiplotrema species would be as-
signed. Apart from the nephridial modifications,
which together with the absence of genital setae,
serve to distinguish it from its congeners, N.
exigua is otherwise unremarkable, there being
little to justify the erection of a higher taxonomic
category (i.e., a subgenus) to accommodate it and
N. varionephrica.

Neodiplotrema lacisbrontoi sp. nov.
(Figs 4, 11D)

TYPE LOCALITY. 10?46'S, 142°34’E, dense
rainforest at northern end of Lake Bronto, in
openings in forest canopy, black sandy soil, dense
leaf and twig litter, some ground cover, 1 m deep.
Coll. R. Raven, 4 Feb 1975.

MATERIAL EX AMINED. HOLOTYPE. QMG21 1957.
PARATYPE. QMG21 1958 (4 specimens).

DESCRIPTION. Length 165+, 140mm (Holo-
type, P1). Width 4.9mm (H). Form circular in
cross-section throughout, pigmentless buff in al-
cohol. Prostomium strongly furrowed. First dor-
sal pore at 9/10. Setae 8 per segment, closely
paired, the ventral setal couples of XVIII absent;
those of XVII and XIX modified as enlarged
penial setae; the ventral setal couple on the left
side of X very slightly enlarged, but cannot be
regarded as functional genital setae. Nephropores
not visible externally. Clitellum annular, fairly
strongly developed over XIII-XVII (P1), setae
visible. Male pores are situated just posterior to
intersegment 17/18, lateral to b-lines; each sur-
mounts a tiny papilla in a distinctly demarcated
seminal groove, which connect the porophores on
each side. Prostatic openings, 2 pairs, in XVII and
XIX, not on distinct papillae, but rather at the
anterior rim of small, circular concavities (more
pronounced in the anterior pair), which form the
four corners of a roughly square male field. In

TWO NEW GENERA OF ACANTHODRILINAE

FIG. 4. Neediplotrema lacisbronroi genital field.

undissected specimens, this region appears
slightly depressed with respect to the remainder
of the body surface. Female pores are inconspic-
uous slits slightly pre-setal in a of XIV. Sperma-

thecal pores 2 pairs in 7/8, 8/9, each with å
conspicuously puckered rim. Accessory mark-
ings a very narrow tumid strip in XXI in aa with
central depression; a similar marking in 20/21
without depression; a large tumescent patch in the
upper portion of segment XVI, in bb; a papilli-
form swelling in X, centred on the left side ventral
setal couple, but extending laterally in both direc-
tions, as well as anteriad and posteriad (P1 right
side only, P2 present over both setal couples
(reduced in size), P3 both sides, P4 right side
only; å similar marking in IX, centred on ab, but
confined within 2 adjacent intrasegmental fur-
rows (H only); a pair of papilliform swellings in
VII, similarly disposed, but (H) conjoined by å
median tumid band, P3 small papilla left side
only, P4 left side only,

Septum 5/6 moderately thickened, septa 10/11,
11/12 strongly so, 6/7-9/10 very much thickened
and muscularised. Dorsal hlood vessel single,
continuous onto the pharynx, where it divides
repeatedly. Last hearts in XIII; those in XI] and
XIII the only large heart-like commissurals, the
remainder being much reduced, and decreasing
in size anteriorly; commissurals in X-XIH have
thin connectives to both the dorsal and supra-oc-
sophagesl vessels; the latter is visible in segments
X-XVI only, though ill-defined in segments X
and XI. Gizzard very large, cylindrical and highly
muscular, though somewhat compressible, in V,
Oesophagus narrow, with conspicuous lateral
pouching in XIV-XVI (in PI only present in XIV
and XV); these outpouchings are internally la-
mellate, but their function is conjectural (valcif-
erous, digestive, etc.); a further small dilatation
of the oesophagus is present in XIIL Intestine
commences. with abrupt expansion in XVIII
(XVII-P1), a definite double-ridged dorsal
typhlosole present from XIX posteriad, being
maximally developed from segment XXX. Ne-
phridial system meronephric, in the forebody
consisting of numerous, simple astomaie, ex-
onephrie loops in each segment (more than 30 jn
the intestinal region); Caudally De, the last 35-
40 segments, the nephridial bodies of the last 15
or sa segments being considerably enlarged. No
enteronephry demonstrable; tufting present in 1V,
presumably enteronephric, but the composite
ducts not traceable; the nephridial loops noticea-
bly longer in V and the following oesophageal
segments than in the intestinal region.

Holandric; 2 pairs small, highly convolute non-
iridescent sperm funnels in X and XI, the former
the larger; 2 pairs rounded, racemose seminal
vesicles in X1 and XII, the latter being obviously

146

the larger; (vice versa in PI). Vasa deferentia
readily traceable on the body wall as a pair of
closely associated tortuous tubes entering the pa-
rietes in XVIL Prostates, 2 pairsin XVI and XIX,
each a narrow, highly coiled tube, restricted to its
segment of origin, with a long, thick, looped
muscular duct; the anterior prostates are invari-
ably the larger pair. Medium-sized penisetal fol-
licles, containing few reserve setae, present im
XVII and XIX, associated with the prostatic
ducts; extensive copulatory musculature (i.e.
connective ligaturing of the follicles to the body
wall, for the eversion and/or retracuon of the
penial setae durimg copulation) present, The setae
robust, stout, somewhat flattened, the ectal end
often twisted, the tip invariably recurved; the
ectal 1/3 is ornamented with incomplete, stag-
gered circlets of short, irregular toothlets. Length
of mature seta 0.88mm; midshaft diameter
3L7um (mean of 3). Ovaries fan-shaped, consist-
ing of many strings of small oocytes, and these,
logether with medium-large oviducal funnels, are
located in XIII. Spermathecae 2 pairs, in VII and
IX, each comprising a sacciform ampulla that is
produced into a long digitiform projection (di-
rected anteriad im situ), and a sessile, U-shaped
diverticulum studded with large, iridescent, intra-
mural sperm chambers; the diverticulum is em-
bedded in the wall of the sac-like portion of the
ampulla; length of right spermatheca of IX
2.9mm. Brain crescentic, with broad fusion of the
supra-oesophageal ganglion; 2 quite distinct pro-
stomial nerves arise close together at the point
where the commissure widens to form the dorsal
ganglion; these innervate different regions of the
prostomium. Peristomial nerve single. but
branching a short distance from its point of origin
at the commissure (approximately midway be-
tween the sub- and supra-oesophageal ganglionic
masses), Spermathecal genital setae absent;
slightly enlarged setae are associated with the
genital marking on left IX, but these are not
specialised, and appear to be non-functional,

ETYMOLOGY. For the type-locality, Lake Bronto.

REMARKS. This species is distinguished from
other forms in lacking genital setae, in having the
male pores in an anterior position (near 17/18),
and in exhibiting oesophageal pouching in the
region XIV-XVI. Its closest affinities appear to
lie with the much larger N. raveni, though this
relationship is by no means close.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Neodiplotrema occidentalis genital field.

Neodiplotrema occidentalis sp. nov.
(Figs 5, IIF)

TYPE LOCALITY. 12?35'S, 141955'E, Weipa, north
bank of the Mission River, Cape York Peninsula, in
semi-deciduous vine-forest, litter layer 4-5em deep,
over black ‘bauxite’ soil, Coll. R. Raven, 7 Feb 1975.

MATERIAL EXAMINED, HOLOTYPE. GD 75/44,
PARATYPES, 2 aclitellate specimens. GD 75/45.

DESCRIPTION, Length 34+ (posterior ampu-
tee), 59mm, Width (midclitellar) 4.4, 4.2mm,
Segments ?, 135 (Holotype, Pl). Circular in
cross-section throughout, pigmentless buff in al-
cohol, preservation poor (some maceration). Pro-
stomium pro-epilobous, first dorsal pore 18/19
(H, P1). Setae 8 per segment, closely paired.
Ventral setal couples of XVIII present: those of
XVII and XIX enlarged as penial setae; genital

TWO NEW GENERA OF ACANTHODRILINAE 147

setae lacking. Nephropores not externally recog-
nizable. Clitellum tumid, saddle-shaped, better
developed dorsally, extending over segments
XIII-XIX, ihe ventro-lateral limits indefinite
Male potes not definitely demonstrable (due 1o
partial maceration), hut presumed to lie within the
shallow seminal grooves linking the prostatic
pores of a side; the latter are atop turnid mounds
that are connected across the ventrum by raised
glandular strips. Each prostatic papilla bears 3
conspicuous pores: the two more closely associ-
ated openings marking the site of exit af the
penial setae (as evidenced by the occasional pro-
truding seta), whilst the remaining orifice is the
prostatic opening proper. Female pores a pair of
obvious slits close to seta b in XIV, Spermathecal
pores inconspicuous in 7/8 and 8/9, aligned with
ab, Accessory markings a series of single, median
pads with paired pore-like centres; a marking in
XVI, with the oculate dimpling closely paired, in
the setal arc; a set of markings decreasing in
lateral width posteriad, commenoes in XIX, im-
mediately posterior to the prostatic porophores
(confluent with the tumid strip linking the latter),
and extends to XXII, the eye-like margins
widely-spaced, not always present on the right
side; a further slight swelling is present medianly
in XXIL The small protruberances in VIII, which
overhand the spermathecal pores are apparently
not associated with genital seta follicles.

Septa 5/6 diaphanous, 6/7 slightly thickened,
7/8-9/10 moderately thus, remainder delicate.
Dorsal blood vessel single, continuous onto the
pharynx; last hearts in XIII, these commissurals
beingthe only ones of any significant size, though
those in XII are fairly robust; commissurals in
XI-XIHHE demonstrably latero-oesophageal, with
the supra-oesophageal vessel visible in segments
X-XIII only, Gizzard large and firm, dolioform,
m 5/6, oesophagus narrow, its length partially
restricted by the posteriorly transgressive piz-
zard; intestine commences in XVI, no definite
Lyphlosole noted. Meronephric throughout; a pair
of very large tufts present in I, their composite
ducts running anteriad to discharge into the buc-
cal cavity; in the mid-body, each segment pos-
sesses approximately 10/12 scattered astomale.
cxonephric loops; caudally, the medianmost ne-
phridium is enlarged as a megameronephridium
(on euch side), having à large. preseptal nephro-
stome; its ducts were not traceable owing to poor
fixation; the remaining nephridia are smaller, as-
tomate, and reduced in number (apparently only
3 on each side, regularly spaced).

Metandric; a single pair of faintly iridescent
spermatic funnels that are nonetheless quite large,
together with presumed testicular tissue (attached
lo postenor face al septum 10/11), seen in XE the
funnels are situated on septum LI/12, above the
ventral body wall, with the vasa deferentia run-
ning down the septum and onto the peritoneum.
where they aré traceable as single, lazily winding
ducts. A single large pair of seminal vesicles that
are finely racemose are located in XU, Prostinc
glands 2 pairs of very thin, fattened tubes, re-
stricted to XVI and XIX, the anterior pair obvi-
ously the larger; the ducts are straight and poorly
muscularised, Large, conjoined penisetal falli-
cles containing numerous reserve Selig, are assu-
ciated with the ectal partion of the prostatic
glands and their ducts; copulatory musculature is
reduced to thin ligaments; the setae gently curv-
ing, tapering gradually to a rather blunt tip; the
ectal extremity (approx, 1/5) ornamented with
mcomplete rows of very short, jagged toothlets.
which become sparser proximally. Length of ma-
ture seta l.63mm; midshall diameter 23. Zum
(mean of 2), Ovaries small, with minute funnels,
in XII, ovisacs not seen. Spermathecae 2 sub-
equal pairs, in VIH and IX, each consisting of a
bent, tubular ampulla, and sessile, reniform diver-
ticulum, with ?radially arranged intramural
sperm chambers containing moumerable brightly
iridescent Necks, Length right spermatheca of TX
].2mm (base of ampulla to pore),

ETYMOLOGY, For the type-locality at the western
extremity af the known generic distribution,

REMARKS. The metandric condition of the måle
gonads, and the lack of genital setae, together
with certain nephridial peculiarities, serve 10
characterise N, occidentalis, Wt is not known
whether this species is linked through intermedi-
ate populations to the Lockerbie and Iron Range
endemics, or is an interpluvial isolate, Senii-de-
«iduous notophyll vine-thickets are very re-
stricted in the Weipa region, usually located on
well-drained sites where freshwater aquilers are
close to the surface; if M. occidentalis is an obli-
gate closed forest inhabitant, its range must pec-
essarily be strictly limited,

Neodiplotrema raveni sp. nov
(Figs 6, 11G)

TYPE LOCALITY, 10*48'S, 142*28'E, Lockertue
East. Cape York Peninsula, collected through formalin
expulsion of very rocky, red lateritie soil; Coll, R.
Raven, 2 Feb 1975.

145

FIG. 6. Neodiploirema raveni genital field.

MATERIAL EXAMINED. HOLOTYPE. QMGH295 |
PARATYPES, Paratypes I and 3 OMG2! 1959,
paratypes 2und4, ANIC. GD 95,9 4,

DESCRIPTION. Length 465, 398mm. Width
(midelitellar) 9.0mm, 6.8mm. Segments 457, 499
(H, PI). Circular in cross-section, pigmentless
biscuit in alcohol, Prostomium pro-epilobous,
peristomium much fissured, First dorsal pore
10/11 (H), 9/10 (P1, 4). Setae 8 per segment. in
regular longitudinal rows throughout; ventral seta
couples of XVII present; those of X VILand XIX
modified as enlarged penial setae; those of TX
replaced by genital setae. Nephropores not seen.
Clitellum saddle-shaped, weakly developed, over
L/2XII-XVIH. Male pores inconspicuous, in the
seminal grooves at 17/18, well lateral of b-lines;
prostatic pores 2 pairs, in XVII and XIX, coinci-
dent with the penial seta orifices, on distinct
mounds; the seminal grooves joining the pores of
aside aré very narrow, but deep, and are slightly
convex laterally. The porophores lie in four dis-

MEMOIRS OF THE QUEENSLAND MUSEUM

ünetly sunken areas, which are transversely
linked by two fairly broad channels, in XVII and
XIX, the former of which is interrupted by à
prominent cuneate tumescence in 16/17, The
male genital area, bounded by the seminal
grooves and the above-mentioned depressions, is
concave with respect to the remainder of the body
surface, Female pores inconspicuous openings
well ventral of a, near 13/14. Spermathecal pores
considerable, bul located in deep intersegmental
grooves, in 7/8 and 8/9, in ab. Accessory mark-
ings extensive forose tissue associated with the
genital setae, in 1X, forming a characteristic
dumbbell-shaped tumid area (all specimens); a
broad, pad present in X (H, P1-2), occasionally a
similar marking in XI (P2); a series of elliptical
pad-like markings (extending across bb) with
transversely depressed centres seen in XXI (H -
the smallest of the three markings), in XXII (H.
P1/3, P2 - indistinct), and in XXIII (H, PI, P2 -
indistinct): a broad pad, occupying much of the
longitudinal dimension of XV, extends across bb
(H; PI-2 J.

Septa: 5/6 moderately thickened, 6/7-10/11
much augmented with thick musculature (7/8-
10/11 the strongest), 12/13 slightly thickened,
remainder delicate. Dorsal blood vessel! single,
continuous onto the pharynx; last hearts in XIII,
only the commissurals in X1-XIII large, heart-
like, and receiving connectives from both the
dorsal and supra-oesophageal vessels. The latter
connectives are the more substantial; the supri-
oesophageal vessel was noted in segments X-XIII
only. Gizzard enormous, cylindrical, and virtu-
ally incompressible, in V; oesophagus completely
suppressed by the posteriad encroachment of the
gizzard in VI-X; thereafter highly vaseular, but
lacking pouching or calciferous glands, Intestine
commences in XVIII, a strong dorsal typhlosole
beginning abruptly at XIX. Meronephric
throughout, nephridia commencing in Il; very
conspicuous, profusely divided tufts present in ITI
and IV, their composite ducts running anteriorly,
but not traced. The oesophageal region with large
numbers of astomate, exonephric, sessile loops
restricted to the parietes or the septal bases; these
are fewer 1n number in the intestinal region, with
4 distinct concentration of bodies near the ventral
nerve-cord: caudally, there is å multiplication and
enlargement of the nephridial bodies, each appar-
ently with its own preseptal nephrostome.

Holandric; wispy testis material and small-me-
dium spermatic funnels (diaphanous, translucent-
non-functional) present in X and XT, testis-sacs
absent; large, acinous, very linely divided semi-

TWO NEW GENERA OF ACANTHODRILINAE

nal vesicles present in XI and XII, the latter mass
the larger. Prostatic glands 2 pairs of simple tubu-
lar organs restricted to, and extending laterally in,
their segments of origin; the anterior pair is the
larger. Bach gland possesses a fairly long, coiled,
muscular duct. The a and b follicles of the penal
setae are indistinguishable. forming 2 pairs of
stout, strongly curved bundles joined to the body
wall in a number of places by copulatory liga-
ments. The setae broad, flattened, attenuating
abruptly near the ectal end, the ectal extremity
invariably recurved; the shaft with a faint scatter-
ing of very find clusters of short toothlets distally.
Length of mature seta 3.43mm; midshaft diame-
ter 123.94m (mean of 2). Medium-sized, pleated
oviducal funnels seen in XII, small ?ovisacs
attached to the posterior face of septum 12/13.
Spermathecae 2 pairs, in VIU and IX, each organ
consisting of a rounded diverticulum sessile on
the body wall (duct virtually non-existent), and à
regular, ovoid ampulla connected to the latter by
a distinct, narrow peduncle; length right sperma-
theca of IX 3.8mm. Genital seta follicles a single
pair, in IX, each containing several yellowish,
slightly curved setae; these are omamented over
their ectal 1/3 not with scalloping, but with irreg-
ular transverse dentate grooves that are at the
most 1/3 shaft circumference long. Length of
mature seta 2.77mm: midshaft diameter 79. Aum
(mean of 3).

ETYMOLOGY, For the collector, Robert Raven.

REMARKS. The lack of spermatozoal irides-
cence in the spermathecae or testis-segments in
any of the specimens examined points to a
parthenogeneuc mode of reproduction for the
species. N. raveni is a very large, deep-burrow-
ing, distinctive species which is very readily iden-
tified, even in a comparatively juvenile star
because of the peculiar bipartite spermathecae. It
possibly also exists in the Torres Strait Islands 1n
suitable habitats, but because of its burrowing.
habits, is would not be readily collected.

Neodiplotrema tumida sp. nov.
(Figs 7-8, 11E,H)

TYPE LOCALITY. ca. 10*48'S, 142728" E, Lockerbie
East, Cape York Peninsula, Coll. R. Raven. Holotype
and paratypes 1, 3, 4: 2 Feb 1975, paratypes 5 and 6:
30 Jan 1975; paratypes 7-10, 11-17: 1 Feb 1975:
paratypes 10, 18, 19: 3 Feb 1975,

109358, 142913 E, Green Hill, Thursday Island, in
damp black clayey soil between rocks in vine-forest,
paratypes 20-25, Coll. G. Berry, 7 Dec 1975.

FIG. 7. Neodiplorrenia tumida genital field.

149

150

MATERIAL EXAMINED. HOLOTYPE. ANIC
GD.95.9.8. PARATYPES. PI, P3, P4, ANIC
GD95.9.8; P5-6, ANIC GD.95.9.6; P11-13; P14,
ANIC GD.95.9.7; P10, P18 & P19 (desiccated) ANIC
GD.95.9.9,

DESCRIPTION. Length 149, 142.5mm. Width
(midclitellar) 4.5, 3.4mm. Segments 282, 242
(Holotype, P1). Form circular in cross-section
throughout, pigmentless buff in alcohol, clitellum
in some specimens pink (P2, P4-6). Prostomium
prolobous; first dorsal pore 8/9 (H, P1-2, P4) or
9/10 (P3). Setae 8 per segment, in regular rows
throughout; ventral setal couples of XVIII absent;
those in XVII and XIX modified as penial setae;
those of IX as genital setae. Nephropores not
conspicuous. Clitellum strongly-developed, sad-
dle-shaped, over XIII-XVI (H. P1, P2-4); dorsal
pores and intersegmental furrows distinct, setae
obscured. Male pores slightly presetal, in narrow
but deep seminal grooves that link the prostatic
pores of a side; the latter are located in lateral
depressions, separated by a central cuneiform
tumescence; the posterior margins of the depres-
sions in XVII and XIX are filled with 2 pairs of
large, rounded swellings (anterior pair the larger),
extending laterally to the seminal grooves,
slightly beyond b-lines; segment XVIII slightly
raised and traversed by deep intrasegmental fur-
rows, and longitudinally seminal grooves, which
turn ventrally to b-lines in XVIII. Female pores
inconspicuous in a transverse furrow, just median
of a-lines near intersegment 13/14, in XIV. Sper-
mathecal pores very obvious on protuberant lips
in b-lines, in 7/8 and 8/9. Accessory markings, a
very large, swollen mound filling segment XI
between the intersegmental furrows, and extend-
ing laterally beyond b-lines by a distance approx-
imately equal to 2bc (all specimens); a similar
tumescence extends over 3 segments (XXVIII-
XXX), with some encroachment on segments
XXVII and XXL, it is dissected by the inter-
segmental furrows of the segments it occupies
(see thus in H, P7); XXVIII-XXIX only in P4, P8;
XXIX-XXX only in P1, P2, P5-6, P9, P16, P17,
P18); XXX-XXXI only in P3; a swollen mound
associated with genital setae in IX, with similar
proportions to the accessory markings in XI (H,
P1-19). Accessory marking in XXXII only (P12,
P12, P15, P19); XXVIII-XXIX only (P13);
XXIX-XXX only (P11); XXXI-1/2XXXII only
(P10); XXIX-1/2XXX1 only (P14).

Septa 11/12 moderately, 6/7-10/11 strongly
mmuscularised, 5/6 only slightly thickened. Dor-
sal blood vessel single, continuous onto pharynx,
where it divides repeatedly in III and IV. Supra-

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Neodiplotrema tumida genital field.

oesophageal vessel present, 1/2XI-XIV. Last
hearts in XIII, those in XI-XIII the largest, receiv-
ing connectives from dorsal and supra-oesopha-
geal vessels, the latter sending the larger
connectives in XII and XXIII; the remaining
commissurals decreasing in size anteriorly, and
dorsoventral only. Gizzard large, highly muscu-
lar, and barrel-shaped, in V, with conspicuous

TWO NEW GENERA OF ACANTHODRILINAE

midlength furrow. Oesophagus rather wide, thin-
walled, in VI-XVI, vascularised lo any extent
only in XI-X V1, calciferous glands or pouching
absent. Intestinecommencing in XVII, with a low
dorsal typhlosole commencing early, well-devel-
oped by XXV. Meronephric throughout; the pre-
intestinal region with scattered astomate
exonephric loops adherent to the body wall and
septa, becoming more numerous in the intestinal
region posteriad of XV (here the nephridial bod-
ies are attached exclusively to the body wall);
caudally with slight enlargement of the nephridial
bodies each with a conspicuous pre-septal neph-
rostome (at least 8 counted on each side); robust
pharyngeal tufting present in IN and IV, the com-
posite ducts not traced.

Holandric; small, iridescent, compactly plicate
funnels and compacted sperm masses present in
X and XI, loosely packed seminal vesicles with
small, scattered loculi present in XI and XI,
Prostane glands small, fattened, tightly coiled
tubular organs with origins in XVII and XIX, the
latter pair much reduced (”becoming vestigial); å
very long, coiled, musculur duct enters a Jobulate
glandular mass on each side in XVII, but these
enter the parietes directly in XIX. Penisetal folli-
cles conspicuous, and attached by ligaments to
the body wall in X VTL the hundles reduced in size
in XIX; the setae are fairly long, and bent into an
arc; the distal portion is often twisted with respect
to the main axis of the seta, the extreme ectal tip
is invariably sharply recurved, uniform, the im-
mediately posteriad portion of the shaft obviously
thickened; the ectal of the shaft is ornamented
with short, discontinuous rows of fine Leeth, the
rows arranged in a crude alternation. length of
mature seta 2,29mm; midshaft diameter 77,.0p.m
(mean of 2), Small ovaries, and medium-sized
plicate funnels seen in XII, ovisacs (7) attached
to the posterior face of septum 12/13. Sper-
mathecae 2 pairs in VII and TX, the posterior pair
readily perceived as the larger; each organ con-
sists of a large, ovoid, bipartite ampulla, and å
flattened, sessile, multi-loculate diverticulum oc-
cupying the dorsal aspect of the smaller ampullal
portion; duct extremely short; length of right
spermatheca of IX 2.7mm. A pair of small genital
seta follicles present in IX (directed anteriorly,
lying beneath the spermathecae); specialised
glands lacking; the setae fairly straight, the ectal
1/5 distinctly omamented with deep notching, the
proximal rims of which are armed with irregular,
sharp, triangular teeth, Length of mature seta
229mm; midshall diameter 52.440m (mean of 3).

151

ETYMOLOGY. For the large ventral genital tumes-
Comes,

REMARKS, The Thursday Island specimens dil-
fer in a number of respects from the Lockerbie
forms, almost lo the extent that subspecific rank
might be warranted, There are, however, unusual
morphological features common to both that sig-
nify close relationship, The major divergences
from the type description exhibited by the Thurs-
day Island forms are as follows: Length i40-
175mm; width (midelitellar) 4.4-4,7mm:
segments 313-324; first dorsal pore 9/10-11/12.
Accessory markings a large, tumid pad extending
across cc, and filling segment LX longitudinally
(P20-21); a pair of mounds or composite bipartite
lumescences posr-setally in. XI, extending later-
ally io mid-be (P20-21) and XV (P20. bai å
large, median tumid pad occupying most nf seg-
ment XXV HI and the extreme anterior portion of
XXIX (P20-21). Supra-oesophageal vessel seen
in X-XVI; intestine commences in XIX:
typhlosale commences immediately, bul enlarges
abruptly in XXXI as a complex, 4-folded struc-
ture; posterior prostatic glands not much reduced
(as for mainland specimens); spermathecal am-
pulla bent, slightly bulbous ectally, the diverticu-
lum a sessile tuherele on the bulbous portion of
the ampulla. Length right spermatheca of IX
(base of ampulla te pore 2.6mm).

The presence of genital setae in IX, the location
of the male pores in. 17/18, the occurrence of à
large genital pad in the vicinity of XX VIDI-XXXI,
and the terminally bulbous, hooked penal setae
are all characteristic of N, neida, and may be
used in combination to confirm identity. Inuna-
wre material from Horn Island may also be refer-
able Lo this species, a further indication that it is
comparatively widespread, Little divergence has
apparently occurred since the presumed separi-
tion of the mainland from Thursday Island popu-
lations after the drowning, of Torres Strait,
approximately 6,500-8,000 years bp, The dura-
tion of insular isolation is approximately equiva-
lent to that experienced by mainland and Melville
Island populations of Diplotrema ridet in the
Northern Territory, though overall morphological
divergence is slightly more pronounced in the
latter case

Neodiplotrema varionephrica sp. nov,
(Fig, 9)

TYPE LOCALITY. 10*46'5, 142?34'E, Eastern side
of Lake Bronto, approximately 10 km from the tip of
Cape Yark Peninsula, in dark sandy soil in open Dear:

Ivyprus dominated forest, with dense ground cover. Coll.
R. Raven, 2 Feb 1975.

MATERIAL EXAMINED. HOLOTY PE. QMGH2952,
PARATYPES. QMGH2953. OTHER MATERIAL.
Lake Bronto, on easrern side of lake, open eucalypt
forest, much ground cover, dark sandy soil, One spec-
imen registered as QMG211960; 11 further specimens
registered as QMG211961.

DESCRIPTION. Length 43, 42-- mm (posterior
amputee), Width (midelitellar) 3.1, 3.0mm. Seg-
ments 155, 110+ (Holotype, P1). Form circular m
cross-section throughout, pigmentless grey-buff
in alcohol, elitellum a slight pinkish-brown. First
dorsal pore 11/12 (H, P1), prostomium pra-
epilobous. Setae 8 per segment, commencing in
W, in regular longitudinal rows throughout; ven-
tral couples of XVIII absent or much reduced;
those of XVII and XIX modified as enlarged
penial setae; those of X replaced by genital setae.
Nephropores not visible externally. Clitellum an-
nular, more pronounced dorsally, over XIH-
XVI dorsal pores obscured, intersegmental
furrows only visible ventrally. Male pores small
shts located in the relatively broad. seminal
grooves linking the porophores of a side; the male
openings are well lateral of b-lines, located ap-
proximately midway between the setal arc and
17/18; prostatic pores 2 pairs, in XVII and XIX,
each on a slight protruberance overhanging a
deep transverse fissure, Female pores small slits
barely presetal, median of a-lines by the distance
ab I in XIV. Spermathecal pores in 7/8 and 8/9,
in a-lines, the slit-like pores inclined at å slight
angle 10 the plane of the intersegmental furrows,
Accessory markings il conspicuous tumescence
associated with the genital setae present in X (H,
P1-left side only, P? left side only). A series of 6
pairs of ellipsoidal intersegmental markings with
oculate. centres commencing in. 10/11 (H, Pl-
right side only, P2, P3-lett side only, P4-11), in
14/12 (H, PT-11), in 12/13 (H, PS-right, P6-right,
P9, PLI), in 13/14 (H, P1-7, P9-11), in 14/15 (H,
P1-5, P6-right, P7-11). in 13/16 (H, PL-10) and in
16/17 (H, PI), A broader tumid pad extends
laterally to b-lines to fill segment XIV between
the intersegmental markings at 13/14 and 14/15
(H, P1-2, P3-weak, P4- 11); å similar transverse
marking máy be present in XX (to h-lines) seen
in P3-4, P7, P9 (weak), PI0-11; a transversely
elliptical pad in 21/22, extending laterally to
slightly beyond b-lines rs often present (PL P5-6,
P10-11), with a similar marking less commonly
occuring in 22/23 (P6-7 only); exceptionally,
there is an additional marking similar to those in
the clitellar region in 20/21 (P10, right side only).

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. Neodiplotrema varionephrice genial Field.

Genital seta location variations: located in VII
(P3, lelt side only, PS, left side only). in VHI
(P4-5), and in IX (P3, left side only), No acces-
sory markings- corresponding to genital seta lota-
lions were detected in P5-7, P9-11

No septa strongly muscularised, but 6/7- 10/11
slightly thicker than the remainder. Dorsal blood
vessel single; last hearts in XII, those in X-XIIH
with thin connectives received from the dorsal
and supra-oesophageal vessels; the latter vessel
is detectable in IX-XHI. Gizzard large, well-de-
veloped, though compressible, in V. with a salter
proventricular portion; oesophagus fairly short,
somewhat compressed due to the posteriad trans-
gression of the gizzard (septa to. 12/13 directed

TWO NEW GENERA OF ACANTHODRILINAE

posteriad as a result of the latter); oesophagus
well vascularised, with conspicuous rugae on its
inner walls; 3 pairs of large, lateral outpouchings
present in XVI-XVII, containing definite lamel-
lae, but all interconnected, with a single narrow
opening leading to the oesophagus proper in
XVII. Intestinal origin at posterior end of XVIII,
a strongly developed dorsal typhlosole com-
mencing in XXIV. Meronephric; large tufts de-
veloped in V, their composite ducts passing
anteriorly to the buccal cavity; smaller clusters of
loops seen in IV and ?III; the oesophageal region
with approximately 6 small astomate, exonephric
loops on each side; in segment XX, a conspicuous
megameronephridium developed on each side,
with a large preseptal nephrostome, and long
excretory duct discharging into a thin-walled ure-
ter running the length of the body on either side
of the ventral nerve-cord, in ab; in addition, 3
small micromeronephridial loops are retained on
each side of a segment: one in be, with minute
ducts entering the parietes in b-lines, an interme-
diate lateral loop above d, and a dorsal body close
to the middorsal line (the latter are all astomate
and exonephric); this arrangement persists to the
extreme caudal segments, the ureters apparently
discharging at the anus; the megameronephridia
are lost some 30 segments from the posterior end.

Holandric; compacted sperm masses and me-
dium-sized iridescent funnels seen in X and XI;
2 pairs of seminal vesicles, consisting of large,
loosely associated loculi, present in IX and XII,
the posterior set much larger. Prostatic glands 2
pairs of relatively small, simple tubular structures
with short, muscular ducts, restricted to XVII and
XIX; penisetal follicles rather small, but densely
packed with reserve setae; a single band-like
ligature passes across the prostates to link the
follicles to the body wall near the mid-dorsal line.
The setae gently curving, the tip invariably with
a very distinctive trilobate appearance, the ectal
1/4 of the shaft bearing an irregular scattering of
thorn-like (slightly recurved) spines. Length of
mature seta 1.04mm; midshaft diameter 44.3um
(mean of 3). Ovaries consist of flabelliform clus-
ters of oocytes; these and medium-large oviducal
funnels are present in XIII; no ovisacs seen. Sper-
mathecae consisting of an ovoid ampulla and à
short, blunt diverticulum containing a number of
iridescent intramural pockets. Length right sper-
matheca of IX 1.2mm. Genital seta follicles usu-
ally located in IX, no glandular structures
associated; the follicles have some copulatory
musculature; the setae are fairly straight, or-

153

namented over the ectal 1/2-1/3 with regular
notching.

ETYMOLOGY. Referring to the highly diverse ne-
phridial system.

REMARKS. This species is closely allied to N.
exigua, the major synapomorphic character being
the peculiar nephridial arrangement (mixed
mega- and micromeronephridia in the mid-body,
with ureters). N. varionephrica may be distin-
guished on the basis of its very distinctive penial
setae, which, unlike any known Australian acan-
thodrile, possess trifid tips. Other somatic char-
acters, such as the position of the first intestinal
segment, and the presence or absence of genital
setae may also be used in conjunction with penial
seta morphology to identify the species.

Genus Torresiella nov.

DIAGNOSIS. Setae eight per segment, prostates
a single pair in XIX; male pores a single pair
combined with the latter, on XIX. Spermathecal
pores a single pair, ventrolateral, in 7/8. Wholly
meronephric, with astomate exonephric nephri-
dia throughout; tufting present in the pharyngeal
region. Gizzard well-developed, in V. Calciferous
glands absent. Holandric, testis-sacs absent. Pe-
nial setae and genital setae present.

DESCRIPTION. As for the type-species.
TYPE-SPECIES. T. singularis (monotypic).

ETYMOLOGY. Referring to the type locality in Torres
Strait.

Torresiella singularis sp. nov. Dyne
(Figs 10, 11A)

TYPE LOCALITY. 10?37'S, 42°17°E, Horn Island,
Torres Strait, 0.4km east of the airstrip, in moist clay
near eucalypts, beside a narrow creek. Coll. R. Raven,
27 Jan 1975.

MATERIAL EXAMINED. HOLOTYPE. QMGH2936.
PARATYPE. QMGH2937,

DESCRIPTION. Length 74, 92mm. Width (mid-
clitellar) 3.4, 3.3mm, Segments 164, 202 (ap-
proximate due to maceration). Uniformly circular
in cross-section, pigmentless grey in alcohol.
Prostomium prolobous, peristomium with a dor-
sal cleft. First dorsal pore 8/9. Setae 8 per seg-
ment, commencing in II; caudal setae
conspicuously enlarged with respect to the other
somatic setae; ventral setal couples of XLX mod-
ified as enlarged penial setae; those of XVII and

194

FIG. 10. Torresiella singularis genital field.

XVIII lacking. Nephropores not externally rec-
ognizable. Clitellum not developed. Combined
male and prostatic pores in ab on XIX. coincident
with the penial seta orifices. The combined pores
are located on low mounds, the anterior and pos-
terior approaches of which have a darker, glandu-
lar appearance; the male field generally
depresses, with a conspicuous furrow at 18/19,
overhung anteriorly by a lightly tumid region

MEMOIRS OF THE QUEENSLAND MUSEUM

across bb. Accessory markings slight tumid
swelling associated with the development of gen-
ital setae usually present in VIII, below the sper-
mathecal pores, Female pores minute points
presetally, in an intrasegmental furrow, median of
a-lines, in XIV. Spermathecal pores a single pair,
in ab, in 7/8, conspicuous as expanded, rimless
orifices.

Septa: 5/6 delicate, 6/7, 7/8, 8/9 with a slight-
moderate thickening. 9/10-10/11 moderately
muscularised, 11/12 slightly so. Dorsal blood
vessel single, continuous onto the pharynx; last
hearts in XIII, those in X-XIII larger than the
more anterior commissurals, and with connec-
tives to both the dorsal and supra-oesophageal
vessels (the remainder dorsoventral only); supra-
oesophageal vessel weakly developed, widest in
XIV. not traceable anteriad of VII. Gizzard mod-
eralely large, muscular, dolioform, compressible,
in V; oesophagus in VI-XVI, fairly wide, well
supplied with blood vessels, dilating slightly in-
trasegmentally; expanded into broad oul-
pouchings in XIII-XTV, not demarcated from the
lumen, or calciferous gland-like. Intestine com-
mences with gradual expansion in XVII, à
strongly developed dorsal typhlosole present
after XXV, Meronephric throughout; numerous,
scattered, astomate, exonephric loops present on
the parietes throughout, more numerous in the
intestinal region. Caudally, with some slight in-
crease in size, but with no evidence of nephro-
stomes or enteronephric development. A small
tuft occurs in IV.

Holandric; 2 pairs of small, slightly iridescent
spermatic funnels in X and XI, with 2 pairs of
small, loosely compacted seminal vesicles in IX
and XII; vasa deferentia non-iridescent, clearly
visible only in the 3 segments immediately pre-
ceding the prostatic segment; still paired on each
side in XVIII, fusing in XIX, and entering the
parietes simultaneously with the prostatic duct,
"7fusing with the latter at this point. Prostatic
glands small organs, situated far laterally in XIX,
extending into XX, with a few loose coils in the
horizontal plane; the muscular duct of uniform
diameter, long and straight, perhaps as long or
longer than the uncoiled glandular portion, enter-
ing the parietes in XIX, through a glandular mass.
Penial seta follicles conspicuous, a and b compo-
nents distinguishable, each with only 2-3 yellow-
ish setae; the follicles are attached to the body
wall by a large band of retractor musculature
passing across the prostatic glands, and attached
by several strands in 20/21. The setae with a very
straight shaft, ectally with a characteristic unci-

TWO NEW GENERA OF ACANTHODRILINAE l

un
un

FIG. 11. A, Spermatheca of Torresiella singularis. B, Spermatheca of Neodiplotrema deminutionis. C, Sperma-
theca of Neodiplotrema exigua. D, Spermatheca of Neodiplotrema lacisbrontoi. E, Spermatheca of
Neodiploirema varionephrica. F, Spermatheca of Neodiplotrema occidentalis. G, Spermatheca of
Neodiplorrema raveni. H, Spermatheca of Neodiplotrema tumida.

nate appearance: ornamentation restricted to a
small region behind the point where the shaft
begins to curve sharply, consisting of densely
packed, short clusters of jagged teeth. Ovaries
small, the ovarian funnels diaphanous, medium-
sized, located in XII; a pair of botryoidal ovisacs
present, attached to the posterior face of septum
13/14. Spermathecae a single pair, in VIII, each
consisting of a pyriform ampulla, divided into 2
sections by a transverse constriction, and two
discrete, clavate diverticula joining the ectal
ampullal region; duct indistinguishable from the
latter. Length of right spermatheca 2.1mm (the
diverticula may be terminally bifid). Genital seta
follicles present in VIII, with attached mus-
culature, but no glandular structures, The setae
angular in cross-section, fairly straight, the ectal
1/2-1/3 of the shaft conspicuously ornamented
with dentate notching.

ETYMOLOGY, Referring to the monotypic nature and
the rare balantine condition of the genus,

REMARKS. T. singularis is the only known
balantine Australian acanthodrile.

DISCUSSION

Neodiplotrema contains species with nephrid-
ial systems that by previous definition — having
a pair of stomate meronephridia median to asto-
mate micromeronephridia caudally — would be
considered to place it in the Dichogastrini, in the
subfamily Megascolecinae. The genus is here
excluded from the Dichogastrini because of the
obvious independent origin of its excretory
apomorphy, as evidenced by the undoubted close
relationship of its species to those of the
holonephric genus Diplotrema, in the subfamily
Acanthodrilinae (Tribe Acanthodrilini). The clas-
sificatory problem raised by the convergent de-
velopment of complex meronephric systems in
the Acanthodrilini (sensu Jamieson, 1971) dem-
onstrated in this paper will need to be addressed
more broadly, with particular attention given to
the validity of the Dichogastrini as a grouping.
Jamieson (1978) has already shown, in a cladistic
analysis, that Dichogaster (Dichogastrini)
grouped with a meronephric Diplotrema sp. (now
Neodiplotrema; Acanthodrilinae) and stated that
this grouping tended to confirm his suspicion that

"hase: Dichagastrint with acanthodrilin male
pores (India und Africa) are descended from
Acanthodrillinae and are disrinct from
dichogastrins with megascolecin male pores
(Oriental and Australia)".

Torrestella is also meronephric and appears to
be a related to the Diplarrema-Neodiplotrema
assemblage. Balantine reduction (male pores mi-
grating. posteriorly to unite with a single pair of
prostate pores, in XTX), which distinguishes Tor-
resiella from all other Australian avanthodriles, is
å much less common phenomenon than is the
micmscolecine transformation (male pores mi-
grating forwards to unite with a single pair of
prostate pores, in XVII), The term denves from à
meronephric West African species described by
Michaelsen in 1898 for which he erected a new
genus, Balanta, This was on the basis of the
combined male aid prostatic pores being located
on XIX, an arrangement that had not previously
been recorded. Only two years later, in. "Das
Tierreich’, Michaelsen (1900) suppressed
Balama in Dichogasrer, as its only species, B.
ehrhardti, bad close apparent alfinities to other
members of that genus, despite rhe acanthodriline
apparatus of the latter.

Other balantine genera include BHalanreodrilus
(monotypic) recorded from Yucatan Caves by
Pickford (1938), and Sylvodrilus, å New Zealand
taxon, Partial balantine reduction is known front
Udeina (U. montanus) and Pickfordia (P.
hemibalantina Omodeo, 1958); in these species,
the posterior prostates are retained, but the male
pore has not migrated, remaining in XVII. In
Sylvodrilus, the male pores have shifted 10 the
posterior part of XVIL, and in Balanteodrilus,
they are located in 18/19. The most advanced
degree of transformation is thus to be found in
Torresiella, in which the male and prostatic ducts
are intimately associated (?fused) behind the
combined pore. This condition is approached in
Dichogaster ehrhardti, where, according to
Michaelsen (1898): '... der Samenleiter ... in die
vordere Flache einer winzigen, atrium-arligen
Hypodermis-Einsenkung ausmundend, in deren
Grunde der Prostataporus liegt -

Why the balantine condition should be so rare
is not known. In the acanthodrile genera exam-
ined by the author, there is a distinct tendency for
the anterior prostatic glands to be conspicuousl y
larger than the posterior organs, Correspondingl y,
the anterior spermathecae are often smaller than
those posterior. Again, there is no sausfactory
explanation tor these observations, but they do
indicate a certain predisposition to the

MEMOIRS OF THE QUEENSLAND MUSEUM

mucroscolecin reduction, Intermediate stages in
reduction suggest that in either reduction, elimi-
nation of one of the prostatic sets is a gradual
process. The migration of the terminal end of the
vas deferens must be largely influenced by the
differential effects of the two prostatic pairs. This
is preses to be an embryonic phenomenon, the
relative size ot the prostatic primordia having å
deterministic effect on the length of the vas def-
erens and the positioning of the male pore (the
‘balanced’ effect of subequal prostatic sets result-
ing in ån equatorial or slightly presetal position
ot the male pore, as is commonly the case). A
gradualist interpretation, where the male pore
mügrates progressively over many generations
owing to some external selective force, though
difficult Lo accept. appears to be appropriate in
this instance, In cases where the male pore fails
to migrate despite elimination of cither prostatic
pair, stabilising selection or some other influence
may be involved.

ACKNOWLEDGMENTS

The author is indebted to Prof. B.G.M, Jamie-
son, University of Queensland, for considerable
practical assistance and stimulating discussion in
the preparation of this paper.

REFERENCES

BAHL, KN. 1942, Studies on the structure, develop-
ment and physiology of the nephridia of
Oligochaeta. Pan 1. General introduction and the
nephridia of the Octochaetinae. Quarterly Journal
of Microscopical Science 83; 423-457.

1947. Excretion in the Oligochaeta. Biological Re-
views 22: 109-147.

DYNE, G.R. 1979, A new species of Microscolex
(Diplatrema) from New South Wales. Proceed-
ings of the Linnean Sociery of New South Wales
10301): 37-41.

1984. Systematics and Zoogeography of Australian
meguscolecoid earthworms. Unpublished Ph. D,
thesis. University of Queensland.

EASTON, E. 1982. Australian Pheretimoid earthworms
(Megascolecidae: Oligochaeta): a Synopsis and
the Descripuon of a New Genus and Five New
Species. Australian Journal of Zoology 30: 711-735

GLASCO, D.G., BOLTON, M.P. & BRYETT, AJ.
1995. Fauna distribution modelling for Cape York
Peninsula. Cape York Peninsula Land Use Strat-
egy, Office of the Co-ordinator General of
Queensland, Brisbane and Department of Envi-
ronment, Spon and Territories, Canberra,

JAMIESON, B.G.M. 1971. A review af the
Megascolecoid earthworm genera of Australia.
Part I - Reclassificalion and checklist of the

TWO NEW GENERA OF ACANTHODRILINAE

Megascolecoid genera of the world. Proceedings
of the Royal Society of Queensland 82(6): 75-86.

1974. The indigenous earthworms (Mega-
scolecidae: Oligochaeta) of Tasmania. Bulletin of
the British Museum (Natural History) Zoology
26(4): 201-328.

1978. Phylogenetic and phenetic systematics of the
opisthoporous Oligochaeta (Annelida:
Clitellata). Evolutionary Theory 3: 195-233.

1997, this volume. Some new and previously known
species of earthworms from Cape York Peninsula
(Annelida: Oligochaeta: Megascolecidae). Mem-
oirs of the Queensland Museum 42(1): 233-270.

LEE, K.E. 1959. The earthworm fauna of New Zealand.
New Zealand Department of Scientific and Indus-
trial Research Bulletin 130 : 1-486.

1970. Application of numerical taxonomy to the
classification of Megascolecidae (Annelida:
Oligochaeta). Pedobiologia 10: 257-264.

157

MACKENZIE, N.L. & DYNE, G.R. 1991. Earthworms
of rainforest soils in the Kimberley, Western Aus-
tralia. In Mackenzie, N.L., Johnston, R.B. &
Kendrick, P.G. (eds) Kimberley Rainforests of
Australia. (Surrey-Beatty & Sons: Sydney).

MICHAELSEN, W. 1898. Die Oligochåten der
Sammlung Plate. Zoologische Jáhrbucher Suppl.
4, Heft 2: 471-480.

1900. Das Tierreich, Vermes, Lief. 10,
‘Oligochaeta’. (Friedlander und Sohn: Berlin).

OMODEO, P. 1958. La Réserve Naturelle Intégrale du
Mont Nimba. I. Oligochétes. Mémoires de
l'Institut francais d' Afrique Noire. 53: 9-109.

PICKFORD, G.E. 1938. Earthworms from Yucatan
caves. Publications of the Carnegie Institution
491: 71-100.

ECHOLOCATION AND ROOST SELECTION IN
SEMON'S LEAF-NOSED BAT HIPPOSIDEROS
SEMONI. Memoirs of the Queensland Museum 42(1); 158.
1997:- Semon's Leaf-nosed Bat Hipposideros semeni occurs
from Cape York Peninsula to Townsville, with un isolated
record W of Calliope (Schulz & de Oliveira, 1995), Hall &
Richards (1979) noted this bat roosting in tree hollows and
deserted buildings in rainforest areas. Hall (1995) noted its
raosang im caves, mines and rock overhangs and an array of
artificial roost sites, including the door handle of a car, a
clothes closet, an oven and a picture rail. Ail recorded roosts
have been of individuals. (Hall, 1995) except fora 3 and %
ipgether in a cave (Schulz & de Oliveira, 1995). Little is
known abour roost selection and ne maternity sites have been
located. Known raosts near Coen have been destroyed through
inining (Richards & Hall, 1994). The species is known from
10 Subterranean roosis and is classified nationally as rare
(Hall, 1995; Richards & Hall, 1994),

We reports echolocation and roost selechøn in å tower
karst on Kings Plains Station (15938". 144958"), SW of Cook-
town, 26-28 June, 1996, The tower consisted of 2 outcrops of
Sha, surrounded by semi-evergreen notophyll vine thicket up
io 200m wide, Caves and fissures (32) were investigated, by
å single person using a Petzl headlamp. Ruusting bats were
identified without being disturbed; flying hats were captured
with a hand net, idennfied and released

Three individuals were roosting singly in rock fissures with
å southerly aspect within å radius of 30m of each other, The
first was a nulliparous 7, forearm AN 6mm and weight 9.2gm,
roosting in a narrow fissure 2.1m high and 0 6m wide, 5m
from the entrance. Relative humidity was 6% higher than at
the entrance; temperature was the same, No other bats were
roosting in this fissure. The second was a & in non-breeding
condition roosting 7m from the entrance in a narrow fissure
1.2m high and 0.5m wide. This fissure was deep, with two
chambers at least 25m long; at the extremity of one chamber
were 3 roosting Dusky Leaf-nosed bats, Hipposideros eter,
The H. semoni roost site in this fissure had a relative humidity
2% lower than outside the enance while rhe temperature was
3.27€ wanner. The third individual could not be caught as it
was roosting in å tight fissure D. Am wide, 5m from a near
vertical entrance 35m by U. Am wide, The relative humidity
in (his roost was 756 lower than outside the entrance.

Beneath 2 of the roosts were remains of hontsman spiders
and beetle elytra suggesting they may have been prey items
The probable taking of hunisman spiders suggests thut (his
species may he a partial 'eleaner', taking prey from surfaces
such as rock faces oc free trunks. The captured d and Y were
in non-breeding condition, Other bats roosting in nearby
fissures and short caves where no H semoni were located were
the Eastern Horseshoe bar, Rhinolophus megaphyllus, H. uter
and the Common Sheathtail bat, Taphozous georgianus.

Echolocation culls of the captured produced 284 ulmo-
sonic sequences on 2 tapes, using the Anabat I System
(Corben, 1989y and à Realistic VSC-2001 cassette recorder;
135 sequences outside the entrance of the cave (98 perching
in a burtertly net, and 37 while being held), and 149 sequences
aller 115 release into the cave, Upon release, the bat hung on å
rock wall and emitted ultrasonic pulses. Both recording ses-
sans lasted I hour, sequences outside the cave were fram 4
hours before dusk, whereas those inside were done on the hou
prior to dusk. The bat was then left alone for I minutes, to
record Hs emergence calls, The bel emerged al 19:07 when it
was dark hur mooalit, without emitting echolocation calls.
Summons & Stein (1980), characterised the echolocation calls
uf hipposiderid bats us having a Constant Frequency (CF)
contponent, Followed and often preceded by Frequency Mod-
ulaled components (FM). Each of these sequences was exant-
ined withio Anahat H and recalibraued from a recorded A0K Hz

MEMOIRS OF THE QUEENSLAND MUSEUM

calibration tone. The CF components were also measured,

Regardless of where tbe recordings were made, all sequences

showed a CF component at 78KH2, 9K Hz higher than hand-

held Rhinolophus megaphyllus in the same Sëch (Fig. 1).

which fits its frequency range recorded in SE Qld (M.C. de

Oliveira, unpubl, data), The sequences of H. semoni from

Kings Plains were 2-4KHz lower than using å (OMC bat

detecior and different software call analysis package (Coles,

1993). The single roost of H. semoni was reminiscent of roost

sites of the Northem Leafnosed bat, H. stenotis (Schulz &

Mekhorst, 1985, 1986) H senors specimens were also

encountered as individuals roosting close to disused mineshalts

in serni-dark conditions,

Acknowledgements

We thank Lana Little for directions and details on the first
record at Kings Plalps; David Hannah for assisting in Del,

work;the manager of Kings Plains for allowing access and G,

Smith for logistical support.

Literature vited

COLES, R., 1993, Echolocation and foraging of Australian
Horseshoe Bats (Rhinolophoidea). Fifth International
Theriological Conference, Sydney, Abstracts,

CORBEN, C., 1989, Computer-based call analysis for
microbat identification. Macroderma, 7,

HALL, L.S. 1995. Semon's Leaf-nosed Bat Hippasideros
semoni. Pp, 461-462, In Strahan, R. (ed.) "The mam:
mals of Australia’. (Reed Books: Sydney).

HALL, L.S. & RICHARDS, GC. 1979, Bats of Easter Aus-
talia. Queensland Museum Bookler No, 12. (Queens-
lund Museum: Brisbane).

RICHARDS, G.C. & Hall, L.S. 1994, An Action Plan for Bat
Conservauorn in Australia, Draft. Report to Aust. Na-
ture Conservation Agency, Canberra.

SCHULZ, M. & DE OLIVEIRA, M.C, 1995. Micro-
chiropteran fauna of Kroombit Tops, central Queens-
land, including à discussion on survey techniques,
Ausiralian Zoologist 30: 71-77.

SCHULZ, M. & MENKHORST, K. 1985, Notes on the Lesser
Warty-nosed Horseshoe bat (Hipposideros steretis).
Aust. Bat Res, News. 20: 14-16.

1986, Roost preferences of vave-dwelling bats at Pine
Creek, Northern Territory. Macrodernia 2: 2-7.

SIMMONS, J.A. & STEIN, RH A. 1980. Acoustic Imaging in
Bat Sonar: Echolocation signals and the Evolution of
Echolocation. J, Comp, Physiol, 135; 61-84,

MC. de Oliveira d M. Schulz, Fauna Conservation and

Ecology Section, Dept. of Natural Resources, PO Box 631,

fndoeravpillv, Queensland, 4068, Australia: 3 April 1996

fo. (Fs
oe ees «Rhee
60

Frequency (KHr]
ki
=

50

Time (ms)

EIG, 1 Schematic representation of ullmasonie signal of Å.
semoni (Hise) and R. megaphyllus (Rhine), Kings Plains Sm.

BIOMETRICS OF THE BIRDS OF PARADISE (AVES: PARADISAEIDAE): WITH
OBSERVATIONS ON VARIATION AND SEXUAL DIMORPHISM

CLIFFORD B. FRITH AND DAWN W. FRITH

Frith, C.B. & Frith, D.W. 1997 06 30: Biometrics of the birds of paradise (Aves:
Paradisaeidae): with observations on variation and sexual dimorphism. Memoirs of the
Queensland Museum 42(1): 159-211. Brisbane. ISSN 0079-8835.

The Australasian bird of paradise family Paradisaeidae is presently considered to consist of
42 species, The family includes species groups and species exhibiting morphological,
ecological, zoogeographical and behavioural characters of fundamental interest to many
disciplines of science. Over recent years there has been an exponentially rapid increase in
interest in bird of paradise biology and systematics at all taxonomic levels. Biometrical data
presented here are by far the most comprehensive and directly comparative gathered for the
Paradisaeidae at the family, generic, specific and subspecific levels. Mean values and ranges
of measurements of each subspecies are given, and interspecific and intraspecific size
variation and sexual dimorphism discussed. These data and others are used to assess the
validity of bird of paradise subspecies discussed by recent authors. [ ] Paradisaeidae, Birds
of Paradise, systematics, sexual dimorphism, morphometrics.

Clifford B. Frith and Dawn W. Frith, Honorary Research Fellows of the Queensland
Museum, ‘Prionodura’, P.O. Box 581, Malanda, Queensland 4885, Australia; 11 February 1997.

The Australasian bird of paradise family
Paradisaeidae is presently considered to consist
of 42 species constituting 16 genera and involv-
ing a total of approximately 100 subspecies
(Gilliard, 1969), to which a few have been added
more recently. Birds of paradise include species
groups and species exhibiting morphological,
ecological, zoogeographical and behavioural
characters of fundamental interest to many dis-
ciplines of science (Diamond, 1981, 1986;
Beehler, 1989; Johnsgard, 1994). Thus there is a
large literature about birds of paradise including
at least nine major monographs, in addition to
more general works and hundreds of scientific
publications (Frith, 1979). Moreover, over recent
years there has been an exponentially rapid in-
crease in interest in their biology and systematics
at all taxonomic levels (Bock, 1963; Gilliard,
1969: Diamond, 1972, 1986; Pruett-Jones &
Pruett-Jones, 1988, 1990; Beehler, 1989; Sibley
& Ahlquist, 1990; Clench, 1985, 1992; Cracraft,
1992; Christidis & Schodde, 1991, 1992; Frith,
1992; Frith & Cooper, 1996; Frith & Frith, 1990,
1992, 1993a,b, 1994, 1995, 1996a,b, in press).

Birds of paradise attract considerable attention
because of the elaborate to bizarre plumage of
adult males that are related to the polygynous
mating system of most sexually dimorphic (and
some sexually monomorphic) species and prom-
iscuity in males. For details of their general ap-
pearance and biology see Gilliard (1969), Cooper
& Forshaw (1977), Beehler et al. (1986) and
Coates (1990). Promiscuous adult male birds of

paradise show remarkable interspecific diversity
of highly colourful and ornate plumages, which
they use in highly ritualized and complex solitary
or communal (lek or exploded lek) courtship
displays at traditional sites, perches or courts
(Diamond, 1986; Beehler, 1989). A remarkable
feature of these birds, and stressing the close
genetic relationships between them, is the marked
diversity of intra- and inter-generic hybrids
(Stresemann, 1923, 1930, 1934; Mayr, 1941;
Gilliard, 1969; Fuller, 1979, 1995). The vast ma-
jority of known hybrid specimens are in full or
subadult male plumage (Fuller, 1979), but re-
cently two hybrids have been described in female
plumage (Frith & Frith, 1996a, b). One of the
latter, a unique hybrid between Lophorina
superba and Parotia carolae, was until recently
erroneously known as Lophorina superba
pseudoparotia.

Birds of paradise are so conspicuously signifi-
cant to sexual selection theory that they were
described and discussed at length by Wallace
(1869) and Darwin (1871), and innumerable au-
thors since (see references in Frith, 1979; Dia-
mond, 1981; Beehler, 1989; Johnsgard, 1994). In
attempting to understand the evolutionary origins
and functions of polygynous mating systems and
the role and influence of sexual selection within
them, it is important to know the physical attri-
butes of the species concerned. In particular one
must be aware of fundamental sexual dimor-
phism (in characters other than secondary sexual
ones such as adult male plumage). Sexual dimor-

160

phism in size is particularly significant as it is
usually conspicuous in polygynous vertebrates in
which males are typically larger than females, but
is reversed in polyandrous species in which fe-
males may be larger than their multiple mates
(Darwin 1871). Notwithstanding the enormous
literature on the birds of paradise, no study has
been devoted to comprehensively demonstrating
and reviewing variation in size between the spe-
cies, sexes and their subspecies. No publication
presents more than the vaguest of measurements
for only the smallest of samples. Most of them in
fact fail to indicate any sample sizes.

The monograph by Gilliard (1969) is the stan-
dard modern text for comparative bird of paradise
measurements at the species level, but only size
ranges for some basic characters are provided
therein. No sample sizes are given and few, if any,
are presented for the vast majority of subspecies.
Cooper & Forshaw (1977) presented measure-
ments for ‘five or more specimens’ of nominate
subspecies of each species only. The only sub-
groups for which reasonable data have been pre-
sented in recent years are those of the
Glossy-mantled Manucode Manucodia ater
(Gilliard, 1956) and of all Paradisaea species
(LeCroy, 1981). As the latter two studies were
based predominantly upon specimens in the
American Museum of Natural History, they un-
derstandably present relatively small sample
sizes compared to those presented here.

Several authors have discussed the validity of
various bird of paradise subspecies with respect
to plumage morphology and relative sizes
(Gilliard, 1969; Schodde & McKean, 1972, 1973;
Diamond, 1972; Coates, 1990; Cracraft, 1992).
Cracraft dramatically revised the systematics of
the family by applying the *phylogenetic species
concept’ (Cracraft, 1992 and references therein).
This in large part involved Cracraft reviewing the
subspecies acknowledged by Gilliard (1969) and
elevating the vast majority of them to 'species'
level. Cracraft (1992) considered 25 of the bird
of paradise subspecies recognised by Gilliard
(1969) to be not *diagnosably distinct’, and there-
fore reduced them to synonomy in accord with his
phylogenetic species concept. He recognised one
‘species’ named since Gilliard’s monograph
(Epimachus fastuosus ultimus, Diamond, 1969)
and named two new ‘species’ himself (Manuco-
dia [keraudrenii] aruensis and M. [keraudrenii]
diamondi).

All of the above authors expressed the view that
a number of subspecies (valid or invalid *species'
of Cracraft, 1992) were possibly or probably in-

MEMOIRS OF THE QUEENSLAND MUSEUM

valid but that more comparative material, or other
types of evidence, were required for firm conclu-
sions. While larger series of skins of some taxa
were available to us in a single collection, or were
gathered together at a single institution, for com-
parative review of plumage morphology this was
not possible for the majority of the subspecies.
Thus, we seek here to primarily use the consider-
able biometrical data we accumulated to assess
the validity of bird of paradise subspecies pre-
sented by Gilliard (1969), Diamond (1972) and
subsequent authors (Cracraft, 1992; and refer-
ences therein). The value of observed differences
in average size between populations has limita-
tions given that size is a continuously varying
character. It is more useful to plot individual
specimens over the geographical range of a taxon
in order to perceive any clinal variation in size,
but this approach is most rarely applied and is
well beyond the scope of the present study. It is
clear, however, that biometric data available prior
to this study are grossly inadequate and have led
to numerous erroneous statements conceming the
sizes of some characters of some taxa and others
concerning supposed sexual dimorphism in size
(see below).

The present re-evaluation of bird of paradise
taxa in the light of significant biometrical data
sets is timely both in terms of complimenting and
assessing Cracraft's (1992) revision of plumage
morphology. Moreover, it provides basic evi-
dence supporting the subspecific taxonomy to be
followed in a forthcoming monograph of the bi-
ology of the group (Frith & Beehler, in prep.).
This is desirable as it will not be possible to
present and discuss the significant supporting
biometrical data in that forthcoming work. Ac-
cepted genera and their sequence are those of
Beehler & Finch (1985), with the addition of
those taxa extralimital to New Guinea and there-
fore not dealt with in that publication. As taxa
accepted by Gilliard (1969) form the basis for
contemporary studies, we follow here his system-
atics and scientific names at the species and sub-
species level (more recently described taxa
accepted). The latter are presented in chronolog-
ical order of description.

Data sets herein provide a significant resource
to students of the systematics and biology of the
group and also to those interested in sexual selec-
tion and size dimorphism in an avian group (in-
cluding both polygynous and monogamous
mating systems) or avian biometrics in general.
Beyond that, a data set of this magnitude for an
entire avian family will prove of value to biology

BIOMETRICS OF THE BIRDS OF PARADISE

students with an interest in more general studies
of systematics, speciation, sexual and other di-
morphism.

METHODS

During a receni world tour of larger collections
of bird of paradise skins the authors examined all
sexed specimens from a recorded locality. Mu-
seum specimens were examined at or from (on
loan to us) the following institutions: Queensland
Museum, Brisbane; Australian National Wildlife
Collection, CSIRO, Canberra; Australian Mu-
seum, Sydney; Museum of Victoria, Melbourne;
South Australian Museum, Adclaide; Western
Australian Museum, Perth; American Museum of
Natural History, New York; The Natural History
Museum, London; Museum of Vertebrate Zool-
ogy, University of Califomia, Berkeley; The
Field Museurn, Chicago; Museum of Compara-
tive Zoology, Cambridge; Bernice Pauahi Bishop
Museum, Honolulu; Royal Ontario Museum, To-
ronto, Academy of Natural Sciences, Philadel-
phia; The Carnegie Museum of Natural History,
Pittsburgh; Peabody Museum, Yale University,
New Haven; Delaware Museum of Natural His-
tory, Wilmington; National Museums & Galler-
ies, Liverpool; Nationaal Natuurhistorisch
Museum, Leiden; Swedish Museum of Natural
History, Stockholm, Zoologisk Museum
Københavns Universitet, Københaven; Museum
national D'Histoire Naturelle, Paris; Staatliches
Museum für Tierkunde, Dresden; Zoologische
Staatssammlung, München; Zoolagische Mu-
seum Berlin; Museum Alexander Koenig,
Staatliches Museum für Naturkunde, Stuttgart;
Forschungsinstitut und Naturmuseum
Senckenberg, Frankfurt; Zoologisches Institut
und Zoologishes Museum, Hamburg; Museum
Zoologicum Bogoriense, Bogor; National Mu-
seum and Art Gallery of Papua New Guinea, Port
Moresby.

A sample of at least 25 individuals for each sex
and age class for each subspecies accepted hy
Gilliard (1969) was measured where possible.
Measurements were all taken in the same stan-
dard way with the same instruments by CBF.
"Wing length’ is the flattened and straightened
chord and was measured with a stopped steel
decimal rule, "Tail centrals’ is the maximum
length of the longest of the central pair of rectrices
from the point of feather entry into the skin to its
tip, and ‘tail length’ the same but to the tip of the
longest tail feather other than the central pair. Tail
measurements were made with an unstopped

Ip

small steel decimal rule. When the central and
outer tail rectrices are the same length. or nearly
so («3mm difference), only the longer measure-
ment (usually tail length) is presented as ‘tail
length’, unless there is intrageneric variation, and
then both ‘tail centrals' and ‘tail length’ arc given.

Other measurements were taken with steel elec-
ironic digital vernier calipers to the nearest whole
decimal point and checked and zeroed daily. “Bill
length! is from the union of the bill with the
foreskull to the up of the upper mandible, and "hill
width’ from the anterior nostril edge. “Total head
length’ is the maximum distance from the back of
the skull to the tip of the upper mandible. It was
measured only from specimens retaining suffi-
cient of the rear skull to permit it. It is possible
some specimens that appeared complete in this
respect may not in fact have been so and therefore
the total head length figures presented must be
considered minimum ones. "Tarsal length" is from
the intertårsål joint tò the lower edge of the last
undivided scute (scale) before the toes diverge-
Where possihle, all of these measurements were
laken from a total of 5677 museum specimens.
Weights, recorded upon death of the bird, were
noted from the labels of measured specimens and
also from those of an additional 408 skins thal we
did not measure (total specimens being 6035). Wc
also include additional weights (n — 298) ob-
tained from living birds by ourselves or others
trapping and releasing them.

In certain bird of paradise groups species have
peculiar or unique feathers that provided addi-
tional measurements useful for a examination of
possible intraspecific vanalion. In Manucodia
keraudrenii we measured the length of "ear tufts’
of feathers from their posterior base to the lip ol
the longest with å small unstopped steel rule,
Likewise we measured the structurally similar
tuft of feathers at the base of the “flag’-tipped
occipital plumes in adult male Parotia species
and the analagous single elongate ‘ear’ feather
found in female-plumaged Preridophora alberti.
The maximum length of the longest occipital
plumes of both Parotia species and Pr. alberti and
the crest length of Cnemophilus macgregorii were
likewise measured. The maximum length of the
modified upper wing covers or "standards" of
Semioptera wallacei are also presented. The
length of pectoral flank plumes (of Selencidis,
Paradisaea, etc.) are rather more subjeclive and
give only an indication of relative flank plume size.
This involved laying the maie specimen on its
back with the end of a steel rule at the the up of the
tail and subjectively assessing the average length of

the bulk of plumes (excluding odd finer and
longer feather tips) projecting beyond the tail tip.

In Tables 1-42 we present mean values, stan-
dard deviations, ranges and sample sizes for each
measured character of each sex and age class of
all subspecies, and for each species as a whole.
Data for markedly sexual dimorphic species in
which males may have an adult, subadult (trace
to almost complete adult male plumage intruding
into female plumage) and immature male (fe-
male-type) plumage are presented separately for
each male age group. Data for females of those
species whose plumage tS generally similar at all
ages, are presented collectively, The latter are
referred to as adult females despite the fact that
samples wil! inevitably include some individuals
less than adult given the great similarity in plum-
age, but data for individuals smaller than adult
size (1.2, juveniles Lo immatures) are excluded.
Data for basically monomarphic species, in
which adult males and adult females are generally
similar but have a discernibly different immature
plumage, are given separately lor cach immature
sex. A single exception is Loboparudisea sericea
for which we combine data for immatures and
subadults of both sexes as the very few specimens
of birds in the latter plumage did not warrant their
separalion,

In the species accounts we cite the original
description and type locality of each taxon alter
the species name, if monotypic, or after each
subspecies name. Early synonyms are not pre-
sented. We discuss the biometric data for the
species as a whole, and for subspecies where it is
necessary 10 point out differences in size and
proportions berween them. We do nol here fully
describe plumages of nominates or describe and
discuss distributions of the various laxa unless
pertinent as these are widely available (Gilliard,
1969; Cooper & Forshaw, 1977; Cracraft, 1992)
and will be presented anew in a forthcoming
monograph (Frith & Beehler, in prep.).

Because bird of paradise measurements pre-
sented in Gilliard (1969) and Cooper & Forshaw
(1977) are those refered to in the contemporary
ornithological literature we make specific abser-
vations upon them as and when required, Our
relatively large sample sizes enable us to criti-
cally evaluate previously published assessments
of several size-related characters and have in
some cases found them misleading or errancous.
Gilliard (1969) and Cooper & Forshaw (1977)
present 'culmen length’ as opposed to bill length.
Culmen length is that of the exposed culmen and
is taken from the point where forehead feathers

MEMOIRS OF THE QUEENSLAND MUSEUM

no longer cover the culmen to the bill tip. Thesis
not a measurement of a discrete structure and 15
far more subjective (and variable) than ‘bill
length’ (from anterior margin of the skull to the
bill tip). Thus the culmen measurements of Gilliard
(1969) and Cooper & Forshaw (1977) are consis-
tently shorter than ours and we will not allude to
them again unless to note their inherently mis-
leading nature (see Lophorina superba below).

The following abbreviations are used below: A
= adult; SA = subadult; I = immature; MWL =
mean wing length; MTL = mean tail length;
MTCL= mean tail central length; MLL = mean
tarsal length; THL = total head length; MBL =
mean bill length; MBW = mean bill width: MW
= mean weight; METL = mean ear tuft length;
MOPL = mean occipital plume length; MFPL =
mean flank plume length,

RESULTS AND DISCUSSION

PARADISAEIDAE
CNEMOPHILINAE

Cnemophilus macgregorii Crested Bird ol
Paradise (Table 1).

A9 similarly sized to or slightly smaller than
Ad, and of similar proportions, MWL and MW
hy 4 & 7% respectively. MTL similar in AF, Ad
and Id. Thus, å do not acquire a progressively
shorter tail with age (as in C. loriae). MTL as å
proportion of MWL similar in both sexes, 80 &
82% respectively. MLL and MBL of A 9 3 & 9%
shorter than in Ad respectively, but similar in
proportion to MWL in both sexes, Crest lengths
of Ad and SAG average 34 + 4mm (n = 51) &
32 t3mmí(n -9)respectively, but shorter in (Dd ,
and A9, averaging 19 + 6mm (n = 22) & 17 €
4mm (n = 25) respectively.

1) C. m. macgregorii De Vis, 1890. Annual
Report of British New Guinea 1888-89: 62, Mt
Knutsford (erroneous!y Mt Musgrave in Gilliard
1969), Owen Stanley Mis. Wing lengths of 116-
121mm for Ad and 115-I20mm for AG pre-
sented by Cooper & Forshaw (1977) are
inexplicably long given that our measurements
are of maximised wing lengths.

2) C. m. sanguineus Iredale, 1948, Australian
Zoologist 11: 162. Kumdi, Mt Hagen District.
Like nominate hut generally more red, being or-
ange-red about head, less red on back to orange
rump, underparts with less copper-red suffusion,
MWL shghtly larger. but MTL slightly shorter,
than nominate.

BIOMETRICS OF THE BIRDS OF PARADISE

3) C. m. kuboriensis Mayr & Gilliard, 1954.
Bulletin of the American Museum of Natural
History 103: 361. Mt Orata, Kubor Mis. Up-
perparts of A d. very like sanguineus but slightly
paler, less saturated with red and underparts morc
black, less suffused with red-brown. Characters
supposedly disunguising this subspecies from
sanguineus have been doubted (Diamond, 1972;
Bechler in Coates, 1990; Cracraft, 1992); we
found a Mt Giluwe specimen of C. m. sanguineus
showing these characters. MWL and MTL of
kuborensis slightly shorter than sanguineus, but
samples too small for meaningful comparisons.
Morphology does suggest, however, that
kuborensis is invalid and should be synonomised
with sanguineus,

4) C. m. subspecies, Gilliard (1969) was
brought an A å specimen in the field said to come
from the Kraetke Mis, where the species is unre-
corded, which was paler than Wahgi Highlands
birds. He felt that if the specimen was not faded
it might represent an unknown subspecies, An
Ad specimen in Zoologishe Museum, Hamburg
from Malingdam, near Mt Goliath, Irian Jaya, we
examined also fits this description but could pos-
sibly be the result of immersion in alcohol. lis
wing, tail, tarsal and bill lengths measure 115, 86,
43.6 & 29.4mm respectively; bill width 6.3mm
and weight 90g. These measurements are within
the ranges of other subspecies,

Cnemophilus loriae Loria's Bird of Paradise
(Table 2)

AN similar to or only fractionally smaller than
Ad , and of similar proportions, MWL similar but
MW 5% lighter. MTL of AN and Id slightly
longer (3 & 5% respectively) than Ad. Thus, å
acquire a progressively shorter tail with age.
MTL 71 and 74% of MWL in Ad and A9
respectively, so proportionately longer in A 9.
MLL and MBL of AN negligibly (2%) shorter
than in AG , but similar in proportion to MWL in
both sexes.

1) Cnemophilus I. loriae Salvadori, 1894. An-
nals Museo Civico Genova, ser 2. 14: 151.
Moroka, Owen Stanley Mts. Iridescence on ter-
tials of Ad blue,

2) Cnemophilus |. amethystina Stresemann,
1934. Omithologische Monatsberichte 42: 144.
Schraderberg, Sepik Mis. Iridescent upper sur-
faces of tertials of Ad deep violet-purple, but A?
as other subspecies, Similar in size lo nominate,
but MWL and MTL slightly longer than in other
subspecies.

163

3) Cnemophilus l. inexpectata Junge, 1939,
Nova Guinea, new series 3: 77. Bijenkorf, Oranje
Mis. Iridescence of tertials of Ad more green
than the blue of loriae or the violet-purple of
amethystina but AN as other subspecies, Similar
in size lo nominale, with MTL only slightly
shorter than nominate and (more so) amethvsrina.

Loboparadisea sericea Yellow-breasted Bird
of Paradise (Table 3)

A (uniquely in the Paradisaeidae) slightly
larger than, but similarly proportioned to, Ad,
MWL and MW (sample small) by 2 & 14%
respectively. MTL of A? and Id slightly langer
(4 & 7% respectively) than Ad. Thus, å acquire
a progressively slightly shorter tail with age.
MTL 59 & 61% of MWL in Ad and AH respec-
tively. MLL similar in length and in proportion to
MWL in both sexes, MBL of AN 5% longer than
in AF, but similar in proportion to MWL in both
sexes,

DL. s. sericea Rothschild, 1896. Bulletin of the
British Ornithologists’ Club 6; 16, "Dutch New
Guinea’. Restricted to the Weyland Mts (Mayr,
1941). 9 said to be larger than 3 (age unspeci-
fied) in the Weyland Mis bul not so on Mt
Karimui (Diamond, 1972). Certainly 9 from the
Weyland Mts have longer wings and tails than
those on Mt Karimui but Diamond's samples are
too small (Ad x 1; Id x 1, 9€ x 2) for meaningful
comparisons.

2) L. s. aurora Mayr, 1930. Omithologische
Monatsberichte 38: 147. Dawong. Herzog Mis,
Plumage of upperparts significantly brighter
(paler), more brown-yellow, the crown far paler
and more greenish and less brown, but underparts
similar to sericea. Larger than nominate, MWL
and MTL of Ad by 3 & 7% respectively. While
not discussing it, Cracraft (1992) appeared to
reject Diamond's (1972) attribution of birds from
the Mt Karimui area, Papua New Guinea to the
nominate (where we have also included them)
and considered them to be L s. aurora, I, how-
ever, they are aurora, their measurements would
be at the lower end of the size range for this
subspecies.

PARADISAEINAE

Macgregoria pulchra Macgregor's Bird of
Paradise (Table 4)

A® markedly smaller than, but similarly pro-
portioned to, Ad, MWL, MTL and MW (sample
smalliby 11, H & 26% respectively. Lå and SAG

have not been collected or are difficult to differ-
entiate and so it is unknown if MTL increases or
decreases with age. MTL 76% of MWL in both
sexes. MLL and MBL of A 9 8 & 6% shorter than
in AG respectively, but similar in proportion to
MWL in both sexes.

This sexual size dimorphism is distinctly differ-
ent from all three members of the subfamily
Cnemophilinae and is greater than the majority of
polygynous Paradisaeinae. Its extent is unusual in
a monogamous bird of paradise (Rand, 1940;
Beehler in Coates 1990).

1) M. p. pulchra De Vis, 1897. Ibis 1897: 251,
pl. 7. Mt Scratchley, south-eastern New Guinea.
Wing lengths of 188- 193mm for A9 (Cooper &
Forshaw, 1977) are inexplicably large given our
measurements are maximised wing lengths.

2) M. p. carolinae Junge, 1939. Nova Guinea
(New Series) 3: 82. Oranje Mts. Proportionally
different from nominate, MWL and MTL shorter
by 5 & 6% and 17 & 16% and MLL longer by 5
& 4% in Ad and AN respectively. Weights of
carolinae heavier than pulchra, but samples too
small for meaningful comparisons.

Lycocorax pyrrhopterus Paradise Crow
(Table 5)

AN similar to or slightly smaller than Ad in
size, proportions and notably weight, MWL,
MTL and MW by 3, 4 & 12% respectively. I å
have not been collected or are difficult to differ-
entiate and so it is unknown if MTL increases or
decreases with age. MTL 73% of MWL in both
sexes. MLL and MBL of A 2 & 5% shorter than
in Ad respectively, but similar in proportion to
MWL in both sexes.

I) L. p. pyrrhopterus (Bonaparte, 1851). Con-
spectus Generum Avium 1 (1850): 384. Gilolo I.
Concealed bases of primaries with no white. MTL
75% of MWL in both sexes.

2) L. p. morotensis Schlegel, 1863. Ibis 1863:
119, *Mortag'. Much like nominate but paler,
slightly more brownish above but slightly darker
below. Concealed bases of primaries extensively
white. Significantly larger than the other two
subspecies in all measured characters. MTL 71 &
69% of MWL in Ad and A? respectively, and
thus similar to obiensis but proportionally shorter
than nominate. MWL and MTL of Ad 15 & 9%
and 7 & 8% longer than in Ad pyrrhopterus and
obiensis respectively; these figures slightly less
(12 & 4%, 6 & 3%) for AL.

3) L. p. obiensis Bernstein, 1864. Journal für
Ornithologie 12: 410. Obi Is. Generally like nom-

MEMOIRS OF THE QUEENSLAND MUSEUM

inate but distinct in being much darker overall,
much more glossy blue-green, darker and more
blue-black on crown and uppertail. Far more dif-
ferent from nominate than morotensis and possi-
bly approaching species status (Cracraft, 1992).
Concealed bases of primaries with trace of white
only. Oddly Lambert (1994) incorrectly wrote of
obiensis that birds have a whitish streak above
and behind the eye. Larger than nominate in all
measured characters. MTL 7196 of MWL in both
sexes and thus proportionately shorter than nom-
inate.

Manucodia atra Glossy-mantled Manucode
(Table 6)

AN similar to or slightly smaller than Ad in
size, proportions and notably weight, MWL,
MTL and MW by 4, 6 & 13% respectively. Id
and I? smaller than respective adults in MWL
(6%) & MTL (7 & 5%), and lighter (sample
small). Thus, tail length increases with age in both
sexes. MTL 84 & 83% of MWL in Ad and A9
respectively. The tail/wing index for both sexes
combined ranges from 84-89%, these figures
similar to those given (82-8896) by Coates
(1990). MTL longer than other Manucodia spe-
cies, excluding M. comrii. MLL and MBLof A9
3 & 5% shorter than in Ad respectively, but
similar in proportion to MWL in both sexes.

Since a thorough study by Gilliard (1956), re-
sulting in the acceptance of the three subspecies,
only Cracraft (1992) has reviewed the species. In
assessing plumage variation, from the phyloge-
netic species concept point of view, Cracraft con-
sidered there to be no diagnostic characters
within M. atra populations. We agree with
Gilliard, however, that it would be misleading to
include all populations in a single taxon in the
face of what in fact are quite gross plumage
differences apparent at the extremes of the range
of this species and the clear differences in size
demonstrated here. We therefore follow
Gilliard's treatment until genetic studies provide
definitive answers to this complex problem. Our
measurements of wing and tail for the species as
a whole agree closely with those of Gilliard
(1956); but his exposed culmen is not comparable
with our bill length and his tarsal length range is
larger than ours, indicating differing measuring
techniques.

1) M. a. atra (Lesson, 1830). Voyage of the
Coquille, Zoology 1: 638. Dorey, north-western
New Guinea.

BIOMETRICS OF THE BIRDS OF PARADISE

2) M. a, alrer Rothschild & Hartert, 1903.
Novitates Zoologicac 10: 84. Sudest I. Flanks and
belly markedly more violet than atra. Notably
larger than nominate, Ad by 12 E 15% and AS
9 & 12% in MWL and MTL respectively. MLL,
THL and MBL of A 3 exclusively larger than
nominate and nearly so in A?.

3) M. a. subalter Rothschild & Hartert, 1929.
Bulletin of the British Ornithologists' Club 49;
110. Dobbo, Aru Is. Said to average much more
purple and violet with oil-green rare on adults
(Gilliard, 1969). On average larger than nomi-
nate, but smaller than alter, An analysis more
thorough than is possible here is required in order
to lest the possibility that birds are clinal in size
from larger ones in the extreme south-east of
Papua New Guinea to smaller ones toward the
extreme southeastern-most part of the range of
the generally smaller nominate on the Gulf of
Papua.

Manucudia chalybata Crinkle-collared
Manucode (Table 7)

M. chalybata (Pennant, 1781), Specium
Faunula Indica, in Forster's Indian Zoology
1781; 40 (based on Daubenton, Planches
Enluminées, pl. 634). New Guinea, restricted to
the Arfak Mts, Monotypic. AN similar to or
slightly smaller than Ad in size, proportions and
weight, MWL, MTL and MW by 3, 4 & 3%
respectively. Id and I? smaller than respective
adults in MWL (2 & 495) and MTL (6 & 4%), and
lighter (sample small). Thus, tail length increases
with age in both sexes. MTL 82 & 81% of MWL
in Ad and A 9 respectively. The adult lail/wing
index for both sexes combined ranges from 8 I-
84%, these figures within the range of those (78-
86%) presented by Coates (1990). MLL and MBL
of AN 2 & 5% shorter than in Ad respectively,
but similar in proportion to MWL in both sexes.

The tarsal lengths of 41-45mm for Ad and of
39-46mm for AY of Cooper & Forshaw (1977)
do notagree with our measurements. This species
is difficult to differentiate from M, jobiensis in
almost all external morphology as they are gen-
erally similar and overlap in most measurements,
but the latter species does have à (5-696) shorter
tail proportionate to wing length.

Manucodia comrii Curl-crested Manucode
(Table 8)

AG slightly smaller than AG and of similar
proportions, MWL and MTL by 5 & 4% respet-
tively: Id and I? smaller than respective adults,

165

but samples 100 small for meaningful com-
parsions, MTL 71% of MWL in both Ad and A 8.
The adull tal/wing index for both sexes com-
bined ranges from 70-76%, MÅL and MBL of A ?
3 & 6% shoner than in Ad respectively, but
similar in proportion 10 MWL in both sexes.

Gilliard's (1969) Ad tarsa) length of 57 mm
and Cooper & Forshaws' (1977) of 54-59 mm for
both sexes are very long.

1) M. c. camrii Sclater, P.L. 1876, Proceedings
of the Zoological Society of London 1876: 459.
Huon Gulf in error for Fergusson I.

2) M. c. trobriandi Mayr, 1936, American Mu-
seum Novitates 869: 3, Kaileuna, Trobriand 1
Plumage as nominate with MWL and MTL
shorter by 7 & 9% and 6 & 7 % in A d and AG
respectively. Thus, this subspecies is on average
smaller than the nominate with limited overlap in
size ranges of wing and tail in each adult sex,
Taking this and the two island populations’ allop-
atry into account it seems on balance more useful
to recognise their differences than lo conceal
them, notwithstanding their possibly question-
able status (Cracraft, 1992).

Manucodia jobiensis Jobi Manucode
(Table 91

AN similar lo or slightly smaller than Ad in
size, proportions and notably weight, MWL,
MTL and MW by 4, 5 & 21% respectively. Id
and F9 smaller than respective adults in MWL (3
& 2%) and MTL (4 & 6%), but samples small.
Thus, tail length increases with age in both sexes.
MTL 76 & 75% of MWL in Ad and A 9 respec-
tively, The adult tail/wing index for both sexes
combined ranges from 72-76%, these figures
within the range of those (69-78%) presented by
Coates (1990). MLL and MBL of A? 3 & 5%
shorter than in Ad respectively, but similar in
proportion to MWL in both sexes.

We can generally reconcile our measurements
with those for the species by Gilliard (1969), hut
not at all with Cooper & Forshaws' (1977) sizes
of 168- 173mm for wing and 42-44mm far tarsus
for Ad and 165-172mm for wing and 40-45mm
lor tarsus for A 9 of nominate, which are exclu-
sively toa small (wing) or too large (tarsus),

1) M. j jobiensis Salvadori, 1876. Annals
Museo Civico Genova 7 (1875): 969, Wonapi,
Jobi (= Yapen) I, Only known from Yapen I in
Geelvink Bay, west Irian Jaya.

2) M. i. rubiensis Meyer, 1885. Zeitschrili für
die gesamte Ornithologie 2: 374. Rubi, Geel-
vink Bay. Now known from the mainland coast

166

of northern New Guinea adjacent to Yapen I
eastward to the Astrolabe Bay area of Papua New
Guinea and south in the west to the upper Setekwa
R and in the east the upper Ramu R. MWL and
MTL shorter than nominate, by 3 & 6% and 3 E
2% in Ad and A 9 respectively. Thus, the main-
land birds are on average smaller than those on
Yapen I (Gilliard, 1969). As samples of the latter
are small (n= 6), and wing and tail lengths ranges
of mainland birds in fact entirely overlap those of
the nominate island form, the validity of this
subspecies remains in doubt. Gilliard (1969) de-
scribed rubiensis as similar to jobiensis (Ó wing
length of latter 177-179mm, tail 132- 142mm),
but smaller (3 rubiensis wing length 168-
174mm, tail 127-133mm). Present data are inid-
equate to justify the retention of rubiensis and on
the basis of this we agree with Cracralt (1992)
that it should be synonomised with jobiensis, but
more data are clearly required.

Manucodia keraudrenii Trumpet Manucode
(Table 10)

AF similar to or slightly smaller than Ad in
size, proportions and notably weight, MWL,
MTL and MW by 4, å & 12% respectively. Lä
and I? smaller in MWL (5 & 39) and lighter
(sample small) than respective adults, but only
Id MTL negligibly shorter (2t ), Thus, tail length
increases only shghtly with age in 8, those of 12
and A? being similar. MTL 78% of MW Lin both
sexes. The adult tail/^wing index for both sexes
combined ranges from 73-78%, these figures
being within the range of thuse (70-8155) pre-
sented by Coates (1990). MLL and MBL of AP
3 & 5% shorter than in Ad respectively, hut
similar in proportion to MWL im both sexes.
Propertionale bill length is remarkably consis-
tent, MBL 21% of MWL in all subspecies except
hunsteint and gouldi for which it is 20%.

A difficult species complex within which
Gilliard (1969) acknowledged eight subspecies,
which Cracraft (1992) reduced to seven
(‘species’) by synonomising M. k. mayeri with M,
k. purpureoviolaceus, To these, Cracraft (1992)
added two newly described forms. Cracraft's
(1992) efforts notwithsianding, more collecting
and study of this species are required to meaning-
fully understand distributions and intraspecilic
variation given the patchiness of collecting on
mainland New Guinea and the subjective nature
of plumage characters used (colour and quality of
refracted light in the form of iridesence). The
length of ihe laceolate "ear tulls” are also used in

MEMOIRS OF THE QUEENSLAND MUSEUM

subspecific diagnosis (Gilliard, 1969; Cracrafi,
1992). Data for 10 subspecies are presented, eight
as acknowledged by Gilliard (1969) and the twa
deseribed by Cracraft (1992),

1) M. $. keraudrenii (Lesson & Garnot), 1826.
Bulletin Scientifiques Naturelles (Ferussac) 8:
110. Dorey, Vogelkop, north-western New
Guinea. METL 19 + 4mm (n = 24) & 18+ 2mm
(n 213) in Ad and AF respectively, MTL 78%
of MWL in adults.

21M, k. gouldi (Grey), 1859, Proceedings of the
Zoological Society of London note, p. 158. Cape
York. Like nominate but plumage iridescence
more green, less purple (particularly so op the
upperwing and tail) Ear tuft feathering more
narrowly pointed, METL 34 + 6mm (n = 33) &
29 * 4mm (n = 16) in Ad and AG respectively.
This is much longer than in keraudrenii, but not
longer than in purpureoviolaceus (pace Cracrafi,
1992). MTL proportionately long, 81% of MWL
in adults,

3) M. k jamesii (Sharpe), 1877. Catalogue of
Birds in the British Museum 3: 181, Aleya, Hall
Sound, British New Guinea. Throat and breast
dark metallic blue, washed with green, lacking
purple of nominate, Ear tufis longer than in nom-
inate, METL 27 + 5mm (n2 25) mm & 24 X 7mm
(n 2 25) in Ad and AF respectively. MTL 78%
of MWL in adults.

4) M, E huristeini (Sharpe), 1882, Journal of the
Linnean Society of London 16: 442. East Cape,
New Guinea in error for Normanby L Back, rump
and uppertail are dark bluish-purple (less green)
and the ear tuft feathers are less blue and mare
green than nominate. Generally like nominate,
but much larger overall, MWL and MTL by 18 &
12% and 20 & 15% in Ad and A 9 respectively.
Ear tufts also longer, METL 26 + 3mm (n = 11)
& 22 + 3mm (n=8)in Ad and AN respectively.
MTL 74% of MWL in adults, so proportionately
the shortest of all subspecies,

5) M. k. purpureeviolaceus (Meyer, 18851.
Zeitschrift für die gesammte Ornithologie 2: 375,
pl. I5. Astrolabe Mts. Generally like nominate
but larger with back, breast and belly intensely
iridescent violet-purplish. Ear tufts notably long,
METL 38 + 6mm (n = 35) & 34+ 4mm (n = 29)
in AG and AH respectively. MTL 76% of MWL
in adults.

6) M. k. neumanni (Reichenow, 1918). Journal
fiir Ornithologie 66: 438, Lordberg. Like similar-
sized nominate but lower back. rump. uppertail
and wings dark black-bluish washed iridescent
deep violet-purple, not more greenish. Breast and
belly dark metallic blue, METL 13 + imm (n =

BIOMETRICS OF THE BIRDS OF PARADISE

9)& 11 + Imm(n-8)in Ad and A ® respectively
and similar to those of an otherwise overall larger
adelberti. MTL 80% of MWL in adults.

T) M. Kk. mayri (Greenway, 1942), Proceedings
of the New England Zoological Club 19: 51. Wau,
Morobe District, north-eastern New Guinea.
Near-identical in size/plumage to
perpureoviolaceus and ear tufts also notably
long. METL 38 + I Imm (n = 7) & 37 + 6mm (a
=4)in Ad and AN respectively. MTL 78% of
MWL in adults, The supposed evidence of a
higher wing-tail index in mayri than in
purpureoviolaceus (Gilliard, 1969) is meagre
given limited numbers of specimens of the l'or-
mer, As no other characters including ear tult
length appear to differentiate them, we concur
with Cracraft (1992) that mayri should be syn-
onomised with purpureoviolaceus.

8) M. K. adelberti (Gilliard & LeCroy, 1967).
Bulletin of the American Museum Natural His-
tory 138: 72. Nawawu, Adelbert Mis, Generally
similar to nominate but like M. E neumanni, with
upper wings and tail more green (not, or far less,
blue to purple). Ear tufts shorter than other forms
except neumanni, METL 13 * 2mm (n = 10) &
14mm (n= 1) long in Ad and A9 respectively.
MTL 80% of MWL in adults.

9) M, E aruensis (Cracraft, 1992), Cludistics 8:
10, Wanoem Bay, Kobror I, Aru L West Irian.
Much less green than nominate and differ from
aujacent mainland New Guinea jamesii by being
generally much darker and bluer, less green, Up-
perparts, particularly back, suffused purple, and
lanceolate head feathering deeper, more purple,
cobalt blue than in jamesii: Ad slightly larger
than nominate in MWL (5%) and imore so in MTL
(105€). Ear tuft length most similar to nominate,
METL 22 + Amm (n =4) in Ad. and thus much
shorter than jamesii. MTL proportionately the
longest, 82% of MWL in adults.

10) M. k. diamondi (Cracrafi, 1992), Cladistics
8: 12. Awande, near Okapa, Eastern Highlands
District. Papua New Guinea. Similar to
purpureoviolaceus but defined as distinct from it
in havmg back, upperwings and tail with strong
violet-purple sheens, the breast and helly dark
metallic blue with little or na violet-purple, and
lanceolate head feathering bluish-green as op-
posed to greenish-blue washed violet-purple. Ear
tufts slightly longer than in purpureoviolaceus,
METL 40 x 9mm (n= 11) & 34 t Imm (n 2 4) in
Ad and AG respectively, and longer than in other
subspecies. MTI. 76% of MWL in adults,

167

Paradigalla carunculata Long-tailed
Paradigalla (Table 11)

AS? markedly smaller than Ad in most mea-
surements, MWL and MTL by 11 & 5% respec-
tively. Tails of Iĝ similar in length to A 9. MTL
shorter than MTCL in A 9 (596), Id (7%) and Ad
(18%). Thus, d acquire à progressively longer
tail with age, the central pair increasing consider-
ably in length (20%) and at a far greater rate than
the remainder. MTL 71 & 76% of MWL in A å
and AY respectively (contra P brevicauda}, so
proportionately longer in the $. MLL and MBL
of AY 7 & 2% shorter than in Ad respectively.
but às à proporüon ot MWL negligibly (2 & 3%)
longer.

Wing lengths of 152-157mm for Ad and 152-
159mm for AN of Cooper & Forshaw (1977) are
dramatically/exclusively shorter than ours for the
former and near exclusively shorter for the latter
sex,

1) F c carunculata Lesson, 1835, Histoire
naturelle des oiseaux de paradis et des épirnaques
1835: 242. Arfak Mts,

2) P. c. intermedia Ogilvie-Gram, 1913. Bulle-
tin of the British Ornithologists’ Club 31: 105.
Utakwa R, Nassau Ra, at 5.500 ft. The status of
the long problematical form P c. intermedia has
been recenily discussed in detail, and we note
here only that it is an invalid taxon as the only
three known specimens have been shown to be
relatively young individuals of P brevicauda
(Frith & Frith, in press}. Its biometrical data are
therefore included within P brevicauda.

This species is distinctively different from 2
brevicauda in size, relative proportions and in
sexual dimorphism of these characters. Younger
individuals of carunculara have a shorter tail than
adults, whereas in P brevicauda younger birds
have a considerably longer one than adults.

Paradigalla brevicauda Short-tailed
Paradigalla (Table 12)

P brevicauda Rothschild & Hartert, 1911.
Novitates Zoologicae 18: 159. Mr Goliath, cen-
tral Dutch New Guinea. Monotypic. Synonym: P
b. intermedia (see P. c. intermedia), A similar
toor slightly smaller than Ac, MWL and MW hy
5 & 6% respectively. MTCL only 2 & Imm
shoner thar MTL in Ad and A9 respectively.
A? MTL. however, is considerably longer (28%)
than AG. MTLot Id also much longer (47%) than
Ad, and 19 also have a longer (31%) tail (han
AN (sample small). Thus, d. acquire a progres-
sively shorter (32%1 tail. the central pair also

168

decreasing in relative length (35%) with age.
MTL 34% of MWL in Ad, but 45% in AY, so
proportionately much longer in 9. MLL of A
5% longer than in Ad, but as a proportion of
MWL the same. MBL similar in both sexes, hut
as a proportion of MWL negligibly (255) longer
in A? than Ad.

A stated average tail length of 90mm for this
eet 18 loo long (pace Cooper & Forshaw,

977).

Ptiloris paradiseus Paradise Riflebird
( Table 13)

P. paradiseus Swainson, 1825. Zoological
Journal 1: 479, No type locality (= northern New
South Wales), Monotypic. AN average smaller
than Ad, MWL and MW (sample small) by 10 &
26% respectively, A? also slightly smaller (3%)
in both MWL and MTL than I and lighter (9%,
but small sample), MTL of A ? shorter (7%) than
in Ad, but as å proportion of MWL negligibly
(256) longer. MTL of Id (sample small) also
shorter (4%) than Ad. MTCL 4, 3 & 9% shorter
than MTL in A ?, Id and Ad respectively. Thus,
3 acquire à slightly overall longer tail with age,
unlike other Prileris species, but subsequent tail
centrals decrease slightly in actual length with
age. MILL of A 9 3% shorter than in Ad, but as å
proportion to MWL similar in both sexes. MBL
of AG 8% longer than Ad and as a proportion of
MWL 7% longer,

For discussion of ‘reversed’ sexual dimorphism
in bill length, see Frith (in press),

Ptiloris victoriae Victoria's Riflebird
(Table 14)

P. victoríae Gould, 1850. Proceedings of the
Zoological Society of London 1849: 1 1H. Barnard
Is, North Queensland, Monotypic, AY average
smaller than Ad, MWL and MW by 10 & 18%
respectively. A9 also slightly smaller in MWL
and MTL (3 & 47%) than Id and lighter (7%),
MTL of A9 shorter (4%) than in Ad, but as å
proportion of MWL slightly (4%) longer. MTCL
only 3% shorter in A? and Id, bul 5% shorter in
Ad. Thus, although very little variation in tail
length between Ad and LÅ, tail centrals do de
crease slightly in length with age. MLL ol A ? 64%
smaller in Ad, hut as à proportion of MWL
similar in both sexes. MBLof A F 4% longer than
Ad and as a proportion of MWL 5% longer.

Cooper & Forshaws' (1977) A9 tarsal length
range pf 34-39 mm is exclusively longer than we

MEMOIRS OF THE QUEENSLAND MUSEUM

found. For a discussion of ‘reversed’ sexual di-
morphism in bill length, see Frith (in press).

Ptiloris magnificus Magnificent Riflebird
(Table 15)

AN markedly smallerthan A d , MWLand MW
by 19 & 34% respectively. MWL of AN exclu-
sively shorter than Ad. AY considerably smaller
in MWL and MTL (12 & 8%) than Id , and much
lighter (2796) — more so than the other two
Ptiloris species. MTL of A Ẹ shorter (3%) than in
Ad. but as a proportion of MWL much (10%)
longer. MTL of Id longer (7%) than AG. MTCL
and MTL similar in A ? and Ig, but 5% shorter
in Ad. Thus, d. acquire a progressively shorter
(6%) tail, tail centrals decreasing (11%) in length
ata greater rate with age. MLL and MBL of A ?
10 & 11% shorter than in Ad respectively, but as
aproportion of MWL negligibly (2 & 3%) longer.

Tarsal lengths of Cooper & Forshaw (1977) are
almost longer and are exclusively longer than
ours for Ad and 9 respectively.

1) P. m. magnificus (Vieillot, 1819), Nouveau
Dictionnaire d'Histoire Naturelle, nouveau edi-
tion, 28: 167, ‘Nouvelle Guinée’, restricted to
Dorey, Vogelkop.

2) P. m. alberti Elliot, 1871. Proceedings of the
Zoological Society ol London, p. 583. Cape York,
Australia. MWL of Ad c.Ilmm shorter and
MTL, MLL fractionally less, and bill narrower
and conspicuously more decurved than in other
lwo Subspecies. Extent of culmen base feathering
is intermediate between that of nominate magnifi-
cus and intercedens. MTEL of Ad relatively
shorter (8mm) than rest of tail than in other two
subspecies (both 5mm).

3) P. m. intercedens Sharpe, 1882. Journal of
the Linnean Society of London, Zoology 16: 444.
Milne Bay and East Cape, south-castern New
Guinea. In almost all mean measurements of both
sexes and all age classes this subspecies ts all but
identical to nominate except for MBL being c
4mm less. Bill straighter than that of alberti.

There arc striking differences in d advertise-
ment call between the subspecies (first noted by
Hunstein, in Sharpe 1891) but also within the
populations of £ m. alberti (pers. obs.). In P. m.
magnificus and P. m. alberti the culmen 18 un-
feathered along the ridge while in A m. inter-
cedens only asmall proportion of the culmen base
is unfeathered, The flank plumes in P. m. magnifi-
cus and P m. alberti are longer than the tail but
in È m. intercedens they are equal to or shorter
than the tail, In view of these differences it has

BIOMETRICS OF THE BIRDS OF PARADISE

heen suggested that P. m, intercedens might rep-
resent a distinct sibling species (Beehler &
Swaby, 1991), Because the feathered condition of
the culmen base in Australian P. em. alberti is
intermediate between intercedens and nominate
magnificus and that the Australian population
exhibits considerable geographical variation in
vocalization within its small range
(MacGillivray, 1918; Frith & Beehler, in prep;
pers obs.), however, we here treat jritercedens as
a subspecies. A hybrid specimen between P. m.
intercedens and F. m. magnificus, was collected
at Putei (CSIRO 4112), just to the east of the
Purari R. The subspecies P. m. intercedens is
confined to the east of this river on the New
Guinea south coast and P. m, magnificus other-
wise known from well to the west of it (Beehler
& Swahy, 1991),

Semioptera wallacel Standardwing Bird of
Paradise (Table 16)

A9 average smaller than Ad, MWL and MW
by 8 & 17% respectively. MTL of AG longer
(595) than in Ad and as a proportion of MWL 8%
longer. MTL of Id also longer (7%) than Ad.
MTL and MTCL similar in A ? and Id, but 15%
shorter in Ad. Thus, d acquire å progressively
shorter tail, tail centrals decreasing considerably
(21%) in length and more so with age, MLL of
A9 6% shorter than in Ad. but as a proportion
to MWL similar in both sexes. MBL of AN 3%
shorter than in Ad’, but as a proportion of MWL
negligibly (2%) longer. Standards average 154+

13mm (n = 58) long in Ad.

1) S. w wallacei Gray, 1859. Literary Gazette
(new senes) 39: 406. Near Labuha Village, Bac-
hian (= Bacan) I.

2) S. w, halmaherae Salvadori, 1881. Or-
nitologia della Papuasia e della Molucche 2: Tor-
ino, p. 73. Halmahera. Similar to nominate but
erown and nape (and in some individuals also
mantle) with rich pinkish coppery-purple irides-
cence. MWL as in the nominate, MTL longer and
other measurements slightly smaller.

Seleucidis melanoleuca Twelve-wircd Bird of
Paradise (Table 17)

AN average slightly smaller than Ad, MWL
and MW by 6 & 11% respectively. MTL in AS
much (and exclusively) longer (49%) and as a
proportion of MWL far greater (64%) than in Ad
(40%), MTL of IG similar to AN. MTCL negli-
gibly shorter than MTL in both sexes and all ages.
Thus, d acquire a progressively and grossly

shorter (3596) tail with age. MLL and MBL of A 7
7 & 9% shorter than in Ad respectively, bit
similar in proportion to MWL in both sexes.
MFPL 266 t 35mm (n 2 48) in Ad,

1) S m. melanoleuca (Daudin, 1800). Traité
d'Ornithologie (Lesson) 2: 278. Waigiou, in errur
for Salawaii or the Vogelkop,

2) S. m. auripennis Schlüter, 1911. Falco 7; 2.
Dallmannshafen (2 Wewak), German New
Guinea. Similar to but on average generally
smaller (c, 6%), more so in MBL, than nominate
and underparts of 9. darker, more brownish, and
more heavily barred.

Epimachus albertisi Buff-tailed Sicklebill
(Table 18)

A® similar to Ad in size and weight, MWL
only 34 smaller. MTL of A 9 also shorter (4%)
and has the same (84%) proportionate length to
MWL as Ad. MTL of Id, however, negligibly
(25) longer than Ad. Thus, surprisingly, d de-
crease tail length very slightly with age, but data
are equivocal. MLL similar in both sexes and in
the proportion to MWL. MBL of AH 4% longer
than in Ad and as a proportion of MWL 3%
longer.

Tail length and tail centrals of this species and
E. bruijnii are of similar lengths in both sexes and
all ages, and tail length is shorter than Wing
length. These two species. because af the consis-
tent proportional and tail growth differences be-
tween them and the rwo larger Epimachus species
(fastuosus and meyeri), were long treated as the
separate penus Drepanornis (Gilliard, 1969). We
here follow current usage (Diamond, 1972;
Beehler & Finch, 1986; Beehler et al, 1986) but
note the above differences suggest Cracraft
(1992) was correct in treating Drepanornis as
valid,

DE o albertisi (Sclater, June 1873), Nature 8:
151 and (1873) Proceedings of the Zoological
Society of London p. 558, pl. 47, Hatam, Arfak
Mis, MTL 87% of MWL in adults and proportion-
ately longest m this subspecies.

2) E. a. cervinicauda Sclater 1883. Proceedings
of the Zoological Society of London, p. 578.
Vicinity of Port Moresby. Upperparts in hmh
sexes brown with slightest of chestnut suffusion,
uppertail coverts and tail quer chestnut than an
nominate. Underparts of 9-plumaged birds pale
buff barred strongly with paler brown than al-
bertisi. Smaller than nominate, MTL, 82% of
MWL 82% in adults.

170

3) E. a. geisleri (Meyer, 1893). Abhandlungen
und Berichte des Koniglichen Zoologishcen und
Anthropologisch-Ethnographischen Museums
zu Dresden 4: 15. Sattelberg, Huon Peninsula.
MWL and MTL fractionally smaller than nomi-
nate; those of Ad fall within the ranges given for
the nominate but those of A 9? 9 are exclusive,
but samples too small (n = 9) for meaningful
comparisons. Diamond (1972), Cooper &
Forshaw (1977), Coates (1990) and Cracraft
(1992) concluded that geisleri should be combined
with the nominate, notwithstanding the slightly
smaller size of the former. We concur with this.

4) E. a. inversus Rothschild, 1936.
Mitteilungen aus dem Zoologischen Museum,
Berlin 21: 188. Mt Kunapi, Weyland Mts. Sam-
ples too small (n = 5) for significant biometrical
comparisons, but measurements similar to nomi-
nate. Diamond (1972), Cooper & Forshaw
(1977), Coates (1990) and Cracraft (1992) con-
cluded that inversus should be combined with the
nominate, and we concur.

By synonomising geisleri and inversus with
albertisi, MTL and MWL of both Ad and A 9 of
the resulting albertisi are still larger than those of
cervinicauda and MTL 87% of MWL and propor-
tionately still longer.

Epimachus bruijnii Pale-billed Sicklebill
(Table 19)

E. bruijnii (Oustalet, 1880). Annales des
Scientifiques Naturelles, Paris ser. 6, 9, 1. and
1880 Bulletin de I’ Association Scientifiques,
France, p. 172. Coast of Geelvink Bay between
136930" and 137° of longitude. Monotypic. A9
similarto Ad in size and weight, MWL and MW
by 3 & 9% smaller respectively. MTL in adults of
both sexes almost identical with a similar (69 &
70%) proportionate length to MWL. MTL of Iĝ,
however, longer (5%) than Ad. Thus, surpris-
ingly, d tail length decreases very slightly with
age, but data are equivocal. MLL and MBLof A 9
3 & 5% shorter than in Ad respectively, but as a
proportion to MWL similar in both sexes.

The adult tail length of 16cm for the species by
Cooper & Forshaw (1977) is far too long.

Epimachus fastuosus Black Sicklebill
(Table 20)

A9 markedly smaller than AG, MWL, MTL
and MW by 15, 44 & 30% respectively. MTL of
AG and A9 100 & 32% proportionately longer
than MWL, as in E. meyeri. MTL of A9 and Iĝ
24 & 23% shorter than MTCL respectively, but

MEMOIRS OF THE QUEENSLAND MUSEUM

in Ad this difference far greater (45%). Thus, d
acquire progressively longer tails with increasing
age, the tail centrals more than doubling in length.
MILL and MBLof A ? 7 & 3% shorter thanin AG,
but as a proportion of MWL slightly (3 & 5%) longer.

It is not true that Id can be told apart from 9
in the field by their longer and thicker bills (con-
tra Majnep & Bulmer 1977).

1) E. f. fastuosus (Hermann, 1783). Tabula
affinitatum animalium (Argentorati), p. 195
(based on Daubenton, Planches Enluminées, pls
638-639). New Guinea, restricted to Arfak Mts
(Hartert 1930).

2) E. f. atratus (Rothschild & Hartert), 1911.
Novitates Zoologicae 18: 160. Mt Goliath, Or-
anje Mts, Dutch New Guinea. Ad darker on
underparts and A? more olive, less rufous, on
uppertail than fastuosus. Ad measurements sim-
ilar to nominate, but some variation between A 9 ,
notably in tail length (6% shorter).

3) E. f. stresemanni Hartert, 1930. Novitates
Zoologicae 36: 34. Schraderberg, Sepik Mts. Ad
darker on underparts than nominate but a Leiden
Museum specimen of stresemanni from Anggi
Gita, Arfak Mts, is intermediate in this character.
This subspecies is considerably larger than the
nominate in every respect. It is also larger than
atratus, with which it has been synonomised,
MWL, MTL and MW (sample small) of Ad
being 946, 17% & 18% longer respectively. This
notwithstanding, the subspecies is considered in-
valid given observations of Gilliard & LeCroy
(1961), Diamond (1969) and Cracraft (1992) con-
cerning the clinal nature of otherwise on average
larger stresemanni. When stresemanni is syn-
onomised with atratus, however, the latter be-
comes a taxon of much larger individuals, still
larger than the nominate.

4) E. f. ultimus Diamond, 1969. American Mu-
seum Novitates 2362: 31. Summit of Mt
Menawa, Bewani Mts, Sepik District, Mandated
Territory of New Guinea. Differs from nominate,
but resembles atratus (and stresemanni), in being
more black, less brown, on underparts of Ad and
more olive, less rufous, on uppertail of A 9 , Bill
nearly exclusively shorter (sample small) than
individuals of other subspecies

Epimachus meyeri Brown Sicklebill
(Table 21)

A9 markedly smaller than AG, MWL, MTL
and MW by 14, 20 & 29% respectively. MTL of
Ad and A 9 39 & 28% longer than MWL respec-
tively. MTL of A9 and Iĝ is 32 & 35% shorter

BIOMETRICS OF THE BIRDS OF PARADISE

than MTCL respectively, but in Ad 64%. Thus,
å acquire progressively longer tails with increas-
ing age, the tail centrals more than doubling in
length. MLL of A 9 9% shorter than in Ad, but
similar in proportion to MWL in both sexes. MBL
of A? 3% longer than in Ad, but as a proportion
of MWL 6% longer.

Cooper & Forshaws' (1977) tarsal lengths of
58-62mm and 51-59mm for nominate Ad and 9?
respectively are much larger than we found, with
those of the former sex exclusively so. It is not
mue that Id can be told apart from 9 in the field
by their longer and thicker bills (contra Majnep
& Bulmer, 1977).

1) E. m. meyeri Finsch, 1885. Zeitschrift für die
gesammie Ornithologie 2; 380. Hufeisengebirge,
south-eastern New Guinea. Flank plumes are
fawn brown. Ad and A9 larger than other sub-
species with respect to MWL, MTL. MTCL,
MTL and MW.

2) E. m. albicars (Van Oort, 1915),
Zoologische Mededelingen, Leiden 1: 228
Treubbivak (2366m), Treub Mtis, Central New
Guinea. Like nominate but slightly smaller, and
flank plumes of Ad paler than both meyeri and
bloodi in being white.

3) E. m. megarhynchus Mayr & Gilliard, 1951.
American Museum Novitates 1524; 10.
Gebroeders Mis, Weyland Ra, Dutch New
Guinea, 6000-7000ft. The subspecies
megarhynelius was erected on the basis of å larger
bill than the others, but bill lengths of the only
three specimens known (o 82, 9 79, 83mm) in
fact fall within the range of all subspecies (sec
measurements); see also Cracraft (1992). This
subspecies is now considered synonomous with
albicans (Cracraft, 1992), with which we concur,

4) E. m. bloodi Mayr & Gilliard, 1951, Ameri-
cun Museum Novitates 1524: 10. Mt Hagen, Cen-
tral Highlands, Mandated Territory of New
Guinea, 8300ft. Like nominate but a good deal
smaller and lighter, Flank plumes of Ad påler,
more dirty whitish, than dirty pale brownish of
meyeri.

Astrapia nigra Arfak Astrapia
(Table 22)

A, nigra (Gmelin, 1788). Systema Avium 1: 401 .
‘Oceanic Islands’, restricted to the Arfak Mts,
New Guinea. Monotypic. AN slightly smaller
than Ad, MWL by 895. MTL 116 & 52% longer
than MWLin Ad und A9 respectively. MTL in
A and Id shorter than in Ad by 35 & 33%
respectively, MTL of AY, Id and Ad 17, 19 &

171

30% shorter than MTCL respectively. Thus, d
tail length progressively increases considerably
with age, MTL and MTCL increasing by 50 &
74% respectively, In this respect this species is
similar to A. rothschildi and unlike their conge-
ners. MLL of A9 4% shorier than in Ad, but
similar in proportion to MWL in both sexes. MBL
similar in bath sexes, hut as a Aw gie to MWL
negligibly (2%) longer in AN.

Astrapia splendidissima Splendid Astrapia
(Table 23)

AN closer to Ad in general size than most bird
of paradise species, MWL similar but MW 11%
less — unlike all congeners. MTL 27 & 39%
longer than MWL in Ad and AN respectively.
MTL of À ? and Id longer than Ad by 8 & 12%
respectively. MTL of A9, 13 and Ad 11, 13.&
20% shorter than MTCL respectively. Thus, å
tail length decreases slightly with age, mare so
MTL (11% shorter) than MTCL (4% bg
MLL of A ? negligibly (2%) shorter than in Ad,
but similar in proportion to MWL in both sexes.
MBL similar in both sexes, and also as a propor-
tion of MWL. It is presumably as à result of the
reduced overall body size of this species that
MBL is large proportionate to MWL (3095) com-
pared to other Astrapia species (20-25%).

I) A. s, splendidissima Rothschild, 1895,
Novilates Zoologicac 2: 59, pl. 5. ‘Probably
Charles-Louis Mis’ but instead almost certainly
from Weyland Mts (Mayr, 1941).

2) A. s. helios Mayr, 1936. American Museum
Novitates 869; 3. Mt Goliath, Oranje Ra, Dutch
New Guinea. Like the nominate but crown, neck
and dorsal collar of Ad more bluish and less
golden-green, and spatulate tips of cental mil
feather pair broader. Female-plumaged birds
slightly darker above, Birds of both sexes and all
age classes differ from nominate by having exten:
sive, unconcealed, white bases to underside of
outer primaries except outermost two, this char-
acter being previously overlooked (Mayr, 1936;
Gilliard, 1969; Cracraft, 1992). These characters
and the fact that individuals of helios are on
average larger than the nominate, particularly in
overall wil length (central pair= 15mm longer in
Ad and 8mm in A9), contradict Cracraft's
(1992) view that helios is invalid.

3) A. s. elliottsmithi Gilliard, 1961. American
Muscum Novitates 2031: 3. Mt Hal, 7,200f1 asl,
Victor Emanuel Mts, Mandated Territory of New
Guinea. This subspecies resembles relics, but sts
tail is even longer (central pair = 17mm longer in

172

Ad and 12mm in AS). We agree with Cracraft
(1992) in synonomising elliottsmithi with helios
and concur with his reservations, as the slightly
larger average size of this subspecies appears no
more than the extreme of a west 10 east clinal
increase im the species and because its white
primary bases are of the helios kind and are unlike
those of the nominate, Synonomising these two
subspecies increases the average size of resulting
helios, especially tail length, the central pair
being 20mm longer in Ad and 12mm in & 9 than
jn the nominate.

Astrapia mayeri Ribbon-tailed Astrapia
(Table 24)

Astrapia mayeri Stonor. 1939, Bulletin of the
British Ornithologists’ Club 59: 57. "Eighty to a
hundred miles west of Mt Hagen’ [Station] = Mt
Hagen, PNG.

A markedly smaller than Ad, MWL and MW
by 12 & 10% respectively. MTL 62 & 97% of
MWLin Ad and AG respectively. MTL of A9
and Id longerthan A d by 37 & 46% respectively.
MTL of A9, Id and Ad 50, 46 & 88% shorter
than MTCL respectively, Thus, MTL of Ad de-
creases considerably in length with age (hy 32%),
whereas MTCL increase dramatically (198%
longer), MLL of A? 4% shorter than in Ad, but
as å proportion of MWL negligibly (2%) longer.
MBL similar in both sexes, but as a proportion of
MWL negligibly (29%) longer in AN.

Astrapia stephaniae Stephanie's Astrapia
(Table 25)

AN on average slightly smaller than Ad,
MWL and MW hy 9% & 12% respectively. MTL
only 85% MWL in Ad, but in A? 25% longer.
MTLof Ad shorter than A? and Id by 25 & 27%
respectively. MTL of A?, Id and Ad 40, 43 &
77% shorter than MTCL respectively, Thus, MTL
of Ad decreases considerably with agé (27%
shorter), whereas MTCL increases (8566 longer).
In A d , rectrices other than the central pair are not
proportionately greatly reduced in length as in A.
mayeri, but progressively decrease in length with
age synchronously with grossly increasing cen-
tral rectrices length. MLL of A 3 4% shorter than
in AG, but asa proportion of MWL similar in both
sexes, MBL similar in both sexes, but as a pro-
portion of MWL slightly (3%) longer in A? than
in Ad, MBL of both sexes much longer (13%),
and also longer (3%) as proportion of MWL, than
in the closely related A. mayeri.

MEMOIRS OF THE QUEENSLAND MUSEUM

1) A, s. srephaniae (Finsch & Meyer, 1885).
Zeitschrift für die gesammte Ornithologie 2; 378.
Hufeisengebirge.

2) A. s. feminina Neumann, 1922, Ver-
handlungen der Ormthologishchen Gesellschaft
in Bayern 15: 236. Schraderberg. MWL of Ad
slightly shorter (3%) than stephaniae but the
same as in ducalis whereas MTL the same ås
stephaniae but larger than ducalis, but samples
too small (n 22) for meaningful comparisons.

3) A. 5. ducalis Mayr, 1931. Mitteilungen aus
dem Zoologischen Museum, Berlin 17: 711,
Dawong, Herzog Mts. Like nominate bul less
contrast between colour of crown/nape and the
back, and the crown and nape more bluish-black-
On average smaller than the nominate in all sig-
nificant characters. Confusion over the supposed
range of feminina (sensu stricta) exists:
Gyldenstalpe (1955) and Coates (1990) included
the Wahgi-Sepik Divide, which is considered by
others to be occupied hy ducalis (Mayr &
Gilliard, 1952; Gilliard, 1969). We note
Cracraft's (1992) observations and tentatively
agree with him in relegating ducalis to synonomy
with feminina, while noting their general overlap
in Measurements,

Astrapia rothschildi Huon Astrapia
(Table 26)

A, rothschildi Foerster, 1906. Two New Birds
of Paradise; 2. Rawlinson Mis, north-eastern
New Guinea. Monotypic, AY markedly smaller
than Ad, MWL and MW (small sample) by 124
& 23% respectively. MTL as a proportion of
MWL 87 & 33% longer in Ad and A9 respec-
tively. MTL of Ad shorter than those of A? and
Id by 38 & 36% respectively. MTL of AF, Id
and Ag 14, 12 & 2156 shorter than MTCLrespec-
tively. Thus, å tail length increases considerably
with age, MTL & MTCL by 56 & 74% respec-
tively. Unlike other Astrapia species, except A.
nigra, rectrices other than the central pair do not
decrease in length with age but become popes
sively longer as does the central pair. MLL of A Ẹ
4% shorter than in Ad, but as a proportion to
MWL negligibly (2%) longer. MBL similar in
both sexes, but as a proportion to MWL slightly
(3%) longer in A? than Ad ,

Lophorina superba Superb Bird of Paradise
(Table 27)

AN markedly smaller than Ad, MWLand MW
by 12 & 23% respectively, A 9 also smaller than
IG, MWL, MTL and MW by 6, 3, & 12% respec-

BIOMETRICS OF THE BIRDS OF PARADISE

tively. MTCL negligibly (1-2mm) longer than the
outer rectrices in birds of all ages and sexes, and
this is true for all subspecies. MTL shorter (796)
in A 9 , but as a proportion of MWL longer (3%)
thanin A d . MTL increases slightly (4%) with age
in ĝ, but data vary with subspecies. MLL of A?
7% shorter than in Ad, but as a proportion to
MWL negligibly (2%) longer. MBL similar in
both sexes, but as a proportion of MWL slightly
(3%) longer in A? than Ac.

It is noteworthy that bill measurements of this
species clearly illustrate a serious potential prob-
lem in measuring exposed culmen as opposed to
bill length. Culmen measurements presented by
Gilliard (1969) and Cooper & Forshaw (1977)
suggest that the bill is longer in females than in
males in this species. This is, however, an artifact
of sexual dimorphism in the extent of forehead
feathers and structure; in functional bill length
per se there is no sexual dimorphism, while in
proportionate bill length females are larger.

Complex subspeciation has long been debated,
with various authors accepting different numbers
of subspecies (Gilliard, 1969; Diamond, 1972;
Coates, 1990; Cracraft, 1992). We list all subspe-
cies below, compare their biometrical data and
note which have been, or we consider should be,
synonomised.

1) L. s. superba (Pennant, 1781). Specium
Faunula Indica, in Forster's Indian Zoology, p. 40
(based on Daubenton, Planches Enluminées, pl.
632). New Guinea, restricted to Arfak Mts. A9
blackish-brown on head and nape, except the chin
and throat, with a short line of tiny whitish spots
as a sub-obsolete, post-ocular, supercilium stripe.
Mantle to upper tail coverts and wings dark red-
brown. Upper tail fuscous with pale chestnut
outer feather edging. Chin, throat and underparts
pale buff to darker buff, uniformly and narrowly
barred brownish-black. MTL 73 and 78% of
MWL in Ad and A? respectively. MWL and
MTL of both sexes notably longer than in most
other subspecies.

2) L. s. minor Ramsay, 1885. Proceedings of the
Linnean Society of New South Wales 10: 242.
Astolabe Mts, Mekeo, south-east New Guinea, A
9 blacker plumaged than other subspecies, with
head and throat blackish-brown and upperparts
rich dark chestnut. No supercilium stripe in front
of and above eye and sub-obsolete behind it. No
or little pale nape marking. MTL 67 & 70% of
MWL in Ad and A 9 respectively. MWL of Ad
and A9 shorter (6 & 8%) than in the respective
sexes of the nominate, as is MTL (14 & 16%).
MWL of Ad shorter than all other subspecies.

173

3) L. s. latipennis Rothschild, 1907. Bulletin of
the British Ornithologists’ Club 19: 92. Rawlin-
son Mts, Huon Peninsula. A? head dark brown,
chin and throat whitish, with broad whitish super-
cilium stripe, white streaking on forehead, crown
and nape; upperparts variably olive-brownish.
MTL 70 & 71% of MWL in AG and A 9 respec-
tively. Overall smaller than nominate, being most
similar in size to connectens and addenda with
which Cracraft (1992) has synonomised it. MTL
of Ad almost identical, but MTL longer than in
connectens (4%) and addenda (7%).

4) L. s. feminina Ogilvie-Grant, 1915. Ibis,
Jubilee Supplement 2: 27. Utakwa R, Nassau Ra.
AN similar in general appearance to latipennis,
the broad supercilium stripes joining narrowly
across the nape. MTL 63% & 66% of MWL in
Ad and A9 respectively. Thus, wings are pro-
protionately much longer relative to tail length
than in other subspecies.

5) L. s. niedda Mayr, 1930. Onithologische
Monatsberichte 38: 179. Wondiwoi, and
Wandammen Mts. Ad like nominate in appear-
ance, but 9 plumage with distinctly darker un-
derparts being more ochraceous. MTL 73% of
MWL in Ad, but in A9 this figure much larger
(8296), as in the nominate, but samples small. In
general and proportionate measurements most
like the nominate.

6) L. s. connectens Mayr, 1930. Onithologische
Monatsberichte 38: 180. Dawong, Herzog Mts.
MTL 66 & 70% of MWL in Ad and A 9 respec-
tively. In general and proportionate measure-
ments most like addenda, and similar to
latipennis. Female plumage like latipennis. Con-
sidered virtually indistinguishable from
latipennnis by Diamond (1972), who considered
it a synonym, and was followed by Cracraft
(1992). We concur with Cracraft (1992) that con-
nectens and addenda be synomised with
latipennis (see below).

7) L. s. sphinx Neumann, 1932. Onithologische
Monatsberichte 40: 121. One (type) ?-plumaged
specimen only for which the locality is unknown.
Gilliard (1969) recognised this as a distinct sub-
species with ‘More reddish brown on upperparts
than minor. Eyestripe less extensive and forehead
and neck without white flanks’. Wing and tail
lengths are markedly longer than those of minor
and more like those of latipennis. Cracraft (1992)
synonomised sphinx with minor, an action as
acceptable as any. We note that the presence of L.
superba on the Hunstein Ra is indicated by the
hybrid L. superba x P. carolae previously known
as L. s. pseudoparotia (Frith & Frith, 1996b). It is

therefore passihle the type specimen of spånix is
from the Hunstein Ra (ils measurements are in
fact closer to those of geographically closest ad-
denda).

8) L. s. pseudoparotia Stresemann, 1934.
Onithologische Monatsberichte 42; 144,
Hunsteinspitze, middle Sepik. Until our visit to
the Zoologisch Museum, Berlin, this subspecies
was still recognised (Gilliard, 1969; Diamond,
1972; Cooper & Forshaw, 1977; Cracraft, 1992),
however, the unique type specimen is, in fact, an
individual resulting from the hybridization of
Lophorina superba and Parotia carolae in the
Hunstein Ra (Frith & Frith, 19965). No other
specimen of (normal) L. superba is known from
this mountain range. The brometrical data of
pseudoparotia are therefore excluded from Table
27.

9) L. s. addenda Iredale, 1948. Australian Zo-
ologist 11: 162. Mt Hagen district. A? plumage
like feminina and latipennis but with some differ-
ences (Gilliard, 1969). MTL 65 & 70% of MWL
in Ad and A9 respectively, Similarly size to
latipennis, although MTL in AG 6% shorter. In
general and proportionate measurements it is
most like connectens, however. Placed in the
synonomy L, s, latipennis hy Cracraft (1992) on
plumage morphology.

10) L. s, subspecies. We measured four individ-
uals, one Ad , one SAG (with 15% Ad plumage
but otherwise in female plumage) and two AY,
from Wanuma, Adelbert Mts, Madang district,
We, like Pratt (1982), could not assign these to å
subspecies and they are much larger than any
other laxon measured. Wing, tail, tarsal, bill
lengths and bill width measured 144, 95, 33.5,
29,6 & 5.9mm in the Ad and averaged 124, 84,
29.7, 29.5 & 6.35mm in A (n — 2) respectively.
As in other subspecies, the tail centralsof Ad and
A9 are slightly longer (5 & 3mm respectively)
than the remainder of the tail, Ad wing and tail
lengths are notably long, but the sample too small
for meaningful comparison.

With regard to the validity of taxa in this species
those accepted by Cracraft (1992) broadly agree
with some or most doubts expressed by Gilliard
(1969), Diamond (1972) and Coates 1990), Until
such time as a comprehensive re-evaluation of its
systematics indicates otherwise, we accept the
following subspecies ` L s, superba, niedda,
minor (sphinx), latipennis (connectens, ad-
denda), feminina, with synonyms given in paren-
theses. L. s. pseudoparotia is invalid as detailed
above.

MEMOIRS OF THE QUEENSLAND MUSEUM

Parotia wahnesi Wahnes' Parotia
(Table 28)

P. wahnesi Rothschild, in Foerster & Roth-
schild, 1906. Two New Birds of Paradise: 2
Rawlinson Mts. Monotypic. AN smaller than A d
in MWL (almost exclusively so) and MW (sam-
ple small), by 7 & 15% respectively. MTL as a
proportion of MWL 16 & 3% longer in Ad and
AN respectively, unlike all other Paroria species.
MTL. considerably shorter than MTCL, hy 8%
shorter in both AG & Id and 12% in AG. MTL
and MTCL in AN shorter (18 & 22% respec-
uvely) than Ad, the central pair exclusively so,
MTL of Id longer (5%) than A 9, but shorter
(18%) than Ad, Thus, d tail length increases
with age, MTCL more so (2296) than MTL (16%),
MLL of A9 9% shorter than in Ad, but as à
proportion of MWL similar in both sexes, MI.L,
as in all Parotia species, proportionately long for
a typical bird of paradise, averaging 32% of
MWL in Ad. MBL similar in both sexes, but as
a proportion of MWL neghgibly (2%) longer in
A9 than Ad. MOPL of Ad average 138 t 4mm
(n2 18). METL of Ad longer (25 * 4mm, n— 15)
than Id (15 + tmm, n 2 7) and A? (15 £ 2mm.
n=17%

Parotia sefilata Western Parola
(Table 29)

P. sefilata (Pennant, 1781), Specium Faunula
Indica, in Forster's Indian Zoology, p 40. New
Guinea, restricted to Arfak Mts, Monotypic. A 9
smallerthan Ad in MWL (almost exclusively so)
and MW, by 8 & 10% respectively. MTL 77 &
8455 of MWL in Ad and A respectively. MTCL
negligibly (1-3mm) longer than MTL in birds of
all ages and sexes. MTL and MTCL of AG and
A similar, but 13 5% longer than adults of both
sexes, IG 5% longer than adults of both sexes.
Thus, å tail length decreases slightly (4%) with
age. MLL of A? 10% shorter than in Ad , but as
a proportion of MWL similar in both sexes, aver-
aging 32%, MBL similar in both sexes, but as a
proportion of MWL negligibly (2%) longer in
AN than Ad. Occipital plumes of Ad average
175 +3 mm (n=29). METL of Ad longer (30+
3mm, n = 28) than Id. (16 + imm, n = 16) and
A9 (15+ Imm, n 2 24).

Parotia carolae Carola's Parotia
(Table 30)

AN smaller (7%) than Ad in MWL (almost
exclusively sa) and considerably lighter (37%).

BIOMETRICS OF TME BIRDS OF PARADISE

suggesting a far greater sexual dimorphism in
weight, but samples too small for meaningful
conclusions. MTL. 50 & 64% of MWL in AG and
AN respectively, MTL and MTCL similar in
length in birds of all ages and sexes. MTL of A?
and Id 198 25% longer than in Ad respectively.
Thus, d tail length decreases considerably (20%)
with age. MLL of A 9 8% shorter than in Ad, but
as a proportion of MWL similar in both sexes and
averages 3396. MBL 656 longer in A? than in
Ad, unlike other Parotia species, and as a pro-
portion of MWL slightly (3%) longer. MOPL of
Ad 122 t 7mm (n = 28). METL of Ad longer
(22+ 2mm, n = 11) than Id (12 = Imm, n = 7)
and A? (12+ Imm, n 2 42).

This species presents a complex, and one ol the
most interesting, problems of intraspecific varia-
tion, which requires considerably more collecting
over its range before a satisfactory study can he
undertaken. The subspecific biometries do not
help greatly as sample sizes are small, the char-
acters dO not vary greatly in size between the
subspecies, and relative proportions of MTL,
MTL and MBL to MWL are remarkably similar
throughout the species

1) P e. carolae Meyer, 1894, Bulletin of the
British Ornithologists Club 4: 6, Amberno R (but
apparently in fact Weyland Mts). MOPL 117 +
3mm (n= 11).

2) FP. c. berlepschi Kleinschmidt, 1897.
Onithologische Monatsberichte 5; 46. New
Guinea, The Van Rees Mts of Irian Jaya have been
suggested as the unknown home of this distinct
subspecies (Mayr, 1962) as has the Foja Mts, Irian
Jaya (Diamond, 1985). Only four specimens are
known, Like nominate in size and appearance but
upper neck, nape and mantle heavily bronzed and
with throat black and spatulate tips to occipital
plumes relatively very small; MOPL 126 € Imm
(n 2 2),

3) P. c. meeki Rothschild, 1910. Bulletin of the
British Ornithologists’ Club 27: 35. Setekwa R.
Durch New Guinea, Like nominate in size and
appearance but chin and sides of throat blackish
and d bill longer (near exclusively so, but sample
small). MOPL 119 + 10mm (n — 4).

4) P. c. chalcothorax Stresemann, 1934.
Onithologische Monatsberichte 42: 145
Doormanpaad, upper Mamderano (Idenburg R).
Like nominate but upperparts with bright coppery
sheen, underparts more coppery and long loral
feathering less intense black, being slightly
brownish. MWL of Ad exclusively longer than
Ihe nominate, hut samples small, Occipital
plumes long, averaging 131 +4mm (n = 2).

175

5) P. c. chrysenia Stresemann, 1934.
Onithologische Monatsberichte 42: 147,
Londberg, Sepik Mis. Traditionally said to differ
from nominate hy having the Jong black koral
feathering with a coppery sheen (like eye ring hut
darker), but several specimens lack this character,
their lores being pure black. MWL and MTL of
Ad longer than the nominate, but samples tou
small for meaningful comparisons, Occipijal
plumes long, averaging 131 X 8 mm (n —2).

6) H c, clelandiae Gilliard, 1961. American
Museum Novitates 2031: 5. Telefolmin SQUON.
(1524m), Victor Emanuel Mts, Mandated Terri-
tory of New Guinea, Like nominate but up-
perparts darker, more jet-black, less brown and
on average larger, and AG bill slightly shorter,
Occipital plumes long, averaging 126+ 7mm (n

Z

Parotia lawesii Lawes’ Parntia
(Table 31)

AT slightly smaller than Ad, MWL and MW
by 5 & 14% respectively, MTL as a proportion of
MWL 66 & 52% in Ad and A9 respectively.
MTL and MTCL similar in length in birds of ull
ages and sexes. MTL of A9? and Id 23 & 24%
larger than Ad, Thus, å tail length decreases in
length slightly with age. MLL and MBL ot A?
are 7 & 2% shorter than in Ad respectively, but
as à proportion of MWL similar in both sexes and
averages 31%. MBL proportionately similar in
both sexes. MOPL of Ad 162 + 7mm (n= 58}
METLof Ad longer (30+4mm, n = 32) than 1d
(14 + 2mm.n =26) and AG (I5 X Imm,n=27).

The subspecific biometrics do not help greatly
as the characters do not vary much in size be-
tween the subspecies and the relative proportions
of MTL, MLL and MBL to MWL are remarkably
similar throughout.

DEL lawesi Ramsay, 1885. Linnaean Society
of New South Wales LO: 243. Astrolabe Mis,
MOPLof Ag 160+ 5mm (n = 21).

2) P. L helenae De Vis, 1897. Ibis p. 390,
Neneba, upper Mambare R, north of MI
Srratehley. Differs [rom nominate im Ad
forecrown being bronzed-brown, not silver-
white, and in Y being slightly less red. All mea-
surements of both sexes near-identical Lo
nominate. MOPL of Ad 161 + 7mm (n= 10)
Originally described as a distinet species and
subsequently long considered a subspecies of P
lawestt until Schodde & McKean (1973) and
Cracraft (1992) resurrected it as å species. We
concur with Gilliard (1969), Diamond (19724,

176

Bechler & Finch (1985), Beehler et al. (1986) and
Coates (1990), however, in treating it as P L
helenae.

3) P. I. fuscior Greenway, 1934. Proceedings of
the New England Zoological Club 14: 2, Mt
Missim, Morobe district (Schodde & McKean,
1973; Cracraft, 1992). Ad like nominate but 9
suid to be duller, less-chestnut brown above and
with a darker head than lawesil (Gilliard, 1969),
when there is in fact much variation within the
species. Measurements are entirely compatible
with those of nominate, MOPL of Ad 169 +
IDmm (n = 7). We concur with Schodde &
McKean (1973) and Cracraft (1992) that this
subspecies is invalid and is a synonym of the
nominate.

4) P. I exhibita Iredale, 1948, Australian Zool-
ogist 11: 162. Mt Hagen district. MOPL of
Ad 163 > 7mm (n = 10), Ad said to be like
nominate but with darker sides to head and ven-
trally more uniformly barred (Gilliard, 1969) but
vanatión within the species makes this diagnosis
inadequate, Measurements are entirely compati-
ble with those of nominate and, thus, it should be
synonomised with it, in accord with Diamond
(1972), Schodde & McKean (1973), Coates
(1990) and Cracraft (1992).

Pteridiphora alberti King of Saxony Bird of
Paradise (Table 32)

A 9 slightly smaller than Ad, MWL and MW
by 9 & 11% respectively. MTL negligibly (2%)
shorter in A , but as a proportion of MWL sex-
ually dimorphic, 73% in A9? and 68% in Ad.
MTL of IG negligibly (2%) longer than in Ad.
Thus, d tail length decreases slightly with age.
MLL of A9 5% shorter than in Ad, bur as a
proportion of MWL. similar in both sexes. MBL
similar in both sexes, but as a proportion of MWL
negligibly (2%) longerin AS. MOPLof Ad 410
t 42mm (n 2 37). METLof A9 & Id 17 + 2mm
(n — 46).

1) P a. alberti Meyer, 1894, Bulletin of the
British Ornithologists’ Club 4: 11. ‘Mountains on
the Amberno R`, hut apparently from Weyland
Mis.

2) P a. burgersi Rothschild, 1931. Novitates
Zoologicae 36: 253, Schraderberg, Sepik Mts.
occipital plumes of Ad notably larger than in the
other two subspecies, MOPL 452 + 43mm (n =
3), but sample small for meaningful comparison.

3) P a. hallsiromi Mayr E Gilliard, 1951,
American Museum Novitates 1524: 12, Forests
above Tomba, south slope of Mt Hagen, Central

MEMOIRS DF THE QUEENSLAND MUSEUM

Highlands, Mandated Territury of New Guinei
Birds have on average longer wings (4%, n = 68)
and tail (5%; n = 69) than the two other subspe-
cies, which are almost identical in size, (n 2 76),
but overlap in ranges of measurements 18 consid-
erahle.

We concur with comments of Gilliard &
LeCroy (1968), Diamond (1972), Coates (1990)
and Cracraft (1992) concerning the difficulties of
adequately differentiating the supposed subspe-
cies and therefore consider P. alberti å monotypic
species.

Cicinnurus regius King Bird of Paradise
(Table 33)

AN average the same size as Ad, their MWL
being similar but MW 4% lighter. MTL of AG
considerably longer (75%) than in Ad and as å
proportion of MWL 25% longer. MTL of Id
similar to AY. MTL 4 & 7% shorter than MTEL
in AN and Id respectively, but 80% shorter in
Ad, Thus, å acquire progressively shorter outer
tail rectrices with age, while simultaneously gain-
ing a progressively grossly longer central pair.
MBL and MLL and their proportion of MWL
similar in both sexes.

Central rectrices of SAd show immense van-
ation, Some ?-plumaged birds, with only traces
of Ad plumage, bad normal 9-length central
rectrices or far longer narrowly-pointed ones,
often with slightly curved tips. In other SAd,
with a greater proportion of Ad plumage, cen-
trals were more wire-like with spatulate tips.
which were slightly, half or distinctly curled.
Some individuals even had two very different
central rectrices: one long 9 -shaped one, which
may or may be curved, and a much-longer wire-
like spalulate-tipped one. Near-adult plumaged
SAd often had one central retrice tipped with a
fully coiled green disc and the other hardly coiled
with no green.

The biometrics do not help systematic assess-
ments as characters do not vary greatly in size
between the subspecies, and the relative propor-
üons of MTL, MLL and MBL to MWL are re-
markably similar throughout the species. The
systematic treatment of intraspecies variation has
been the subject of recent debate. Six subspecies
had been long established (Mayr 1941, 1962),
Gilliard ulso (1969) maintained six subspecies
but pointed out that they are all very similar.

1) C c regius (Linnaeus, 1758). Systema Natt-
rae, ed. 10, p, 110. East Indies (= Aru Is — of
Berlepsch 1911 Abhandlungen Senkenberg-

BIOMETRICS OF THE BIRDS OF PARADISE

ischen Naturforschender Gesellschaft 34: 59).
Only known from the Aru Is. MWL, MTL and
MBL of Ad and A9 longest in this form, but
there is considerable overlap in size ranges with
other subspecies. Ad central rectrices are also
long, and in this respect are most like rex and
gymnorhynchus.

2) C. r. rex (Scopoli, 1786). Deliciæ Florae et
Faunae Insubricae (Ticini) pt. 2, 88 (based on
Sonnerat 1776, Voyage Nouvelle Guinea, p. 156,
pl. 95). ‘New Guinea’ = Sorong District,
Vogelkop. Now known from Misol and Salawati
Is and New Guinea except between Geelvink Bay
and the Huon Gulf. Frequently reported to occur
on Batanta I but probably erroneously (Mees,
1982). Slightly smaller than the otherwise similar
nominate in MWL, MTL and MBL. Most similar
in size to gymnorhynchus.

3) C. r. coccineifrons Rothschild, 1896.
Novitates Zoologicae 3: 10. Jobi I (= Yapen I).
Only known from Yapen I. In most measurements
similar to birds of adjacent mainland
(cryptorhynchus). MBL smaller (8%) than the
nominate, being most like similis and
cryptorhynchus.

4) C. r. similis Stresemann, 1922. Journal für
Ornithologie 70: 405. Stephansort, Astrolabe
Bay. Occurs only from Astrolabe Bay and the
upper Ramu R west to at least Humboldt Bay and
upper Memberamo R. This form generally
smaller than all other subspecies in MWL, MTL,
MBL and MLL being most like, but slightly
smaller than, cryptorhynchus in sizes. MTCL
l Imm shorter than nominate.

5) C. r. cryptorhynchus Stresemann, 1922.
Journal für Ornithologie 70: 405. Taua, lower
Mameramo R. Now known from eastern coast of
Geelvink Bay and at north New Guinea near the
Memberamo R mouth only. Most similar in sizes
to similis. MTCL 6mm shorter than nominate.

6) C. r. gymnorhynchus Stresemann, 1922.
Journal für Ornithologie 70: 405. Heldsbachküste
near Finschhafen. Now known from north-east
coastal Huon Gulf. On average generally smaller
and most similar in size to rex overall.

Although birds on the Aru Is. (C. r. regius) are
on average slightly larger than birds of southern
New Guinea (C. r rex), variation is great and
slight differences in plumage morphology are
inadequate to justify an Aru Is subspecies (Mees,
1964; Cracraft, 1992). Gilliard (1969), Diamond
(1972), Cooper & Forshaw (1977) and Cracraft
(1992) also pointed out the weak morphological
differentiation of most populations. Mees (1964,
1965, 1982) suggested that only two subspecies,

177

C. r. regius (including rex and gymnoryhnchus)
and C. c. coccineifrons (including
cryptorhynchus and similis) be retained and that
the roundish spot of green feathers above the eye
of the former and the more elongate one in the
latter be a ‘tolerably constant character’.

More recently Cracraft (1992) recognised four
subspecies (species in his view) but we find
Cracraft's criterea inconsistent (e.g., he syn-
onomises gymnorhynchus into regius stating it is
‘nearly 100% diagnosably distinct’ but retains
three taxa of the coccineifrons group because he
viewed them as ‘nearly 100% diagnosable’). We
do not find Cracraft’s (1992) tentative plumage
characters in support of retaining cryptorhynchus
as distinct convincing. Furthermore all measure-
ments of that population are entirely compatable
with those of Yapen I. (coccineifrons) and north-
ern New Guinea (similis) populations. In fact
almost all measurements of the three populations,
coccineifrons, cryptorhynchus and similis, are
clinal from larger to slightly smaller, west to east.
The more easterly population of gymnorhynchus
does not fit this pattern, however, having a larger
bill and therefore being more like birds of south-
ern New Guinea (C. r. regius).

Thus, we feel Mees (1964) should be followed
in combining these taxa into two subspecies C. r.
regius and C. r. coccineifrons. By so doing, Ad
and A 9 ofthe two subspecies are almost identical
in size, except in tail measurements. There still
exists considerable overlap in size ranges of all
parameters between the two subspecies, however.
MWL of A å of the enlarged C. r. regius (includ-
ing rex and gymnoryhnchus) is a negligible 2%
longer than the enlarged C. r. coccineifrons (in-
cluding cryptorhynchus and similis) and MTL are
the same. MTCL and MBL in Ad C. r. regius,
however, 6 & 7% longer respectively than in C.
r. coccineifrons, and 5 & 2% larger as a proportion
of respective MWL.

Cicinnurus magnificus Magnificent Bird of
Paradise (Table 34)

AN average slightly smaller than AG, MWL
and MW by 4 and 15% respectively. MTL of AN
considerably longer (5196) than in Ad and as a
proportion of MWL 20% longer. MTL of Id
similar to A9. MTL only 2 & 5% shorter than
MTCL in AN and Iĝ respectively, but 86% in
Ad. Thus, å acquire progressively shorter rec-
trices with age, while simultaneously gaining a
progressively grossly longer central pair. MLL of
AN 5% shorter than in Ad, but as a proportion

178

of MWL similar in both sexes. MBL similar in
both sexes and as a proportion of MWL.

The central rectrices of SAG show great vari-
ation, relative to the degree of Ad plumage ac-
quired (as described for C. regius) The
biometrics do not help much as the various char-
acters vary little in size between the subspecies
and the relative proportions of MTL, MLL, and
MBL to MWL are remarkably similar throughout
the species.

1) C. m. magnificus (Pennant, 1781). Specimen
Faunulae Indicae, in Forster's Indian Zoology, p.
40 (based on Daubenton, Planches Enluminées,
pl. 631). Arfak Mts, Vogelkop.

2) C. m. chrysopterus (Elliot, 1873). Mono-
graph of the Birds of Paradise, p. 13. Jobi I (=
Yapen I, Irian Jaya). In all measurements of both
sexes chrysopterus almost identical to nominate,
except for longer (9%) MTCL of Ad. Also differs
from nominate in its secondary coverts and outer
edges of flight feathers being more orange, less
yellow, and the crown darker.

3) C. m. hunsteini (Meyer, 1885). Zeitschrift für
die gesammte Ornithologie 2: 389, pl 21.
Hufeisengebirge, southeastern New Guinea (=
near Astrolabe Mts). Like nominate but paler on
head and back (Gilliard, 1969), and orange on
secondary coverts and outer edge of flight feath-
ers brighter and deeper. All measurements of both
sexes nearly identical to other subspecies, al-
though the MWL in Ad and A 9 slightly shorter
by 3 & 4% respectively.

4) C. m. intermedius (Hartert, 1930). Novitates
Zoologicae 36: 36. Snow Mts (= Upper Setakwa
R, Irian Jaya). Like nominate but the scapulars
and inner secondaries brighter, more dull orange
than brown (Gilliard, 1969). In view of all mea-
surements of both sexes of intermedius being
extremely similar to chrysopterus, and of com-
ments by Cracraft (1992), this subspecies should
be synonomized with the former. MTCL of Ad,
however, shorter (13%) than chrysopterus.

Cicinnurus respublica Wilson's Bird of
Paradise (Table 35)

C. respublica (Bonaparte, 1850). Compte
Rendu des séances de l' Academie des Science,
Paris 30, 131-291. ‘New Guinea’, restricted to
Waigeu I. Monotypic. A 9 average the same size
or slightly smaller than Ad, MWL similar but
MW 8% lighter. MTL of A ? considerably longer
(36%) than in Ad and as a proportion of MWL
15% longer. MTL of Id similar to A 9. MTL and
MTCL similar in AG and Id, but in AG MTL

MEMOIRS OF THE QUEENSLAND MUSEUM

72% shorter than MTCL. Thus, å acquire pro-
gressively shorter rectrices with age, while simul-
taneously gaining a progressively grossly longer
central pair. MLL of A@ negligibly (2%) shorter
than in Ad, but as a proportion of MWL is similar
in both sexes. MBL is similar in both sexes and
as a proportion of MWL.

The central rectrices of SAG show great varia-
tion relative to the degree of Ad plumage ac-
quired (as described for C. regius).

Paradisaea raggiana Raggiana Bird of
Paradise (Table 36)

A9 smaller than Ad, MWL and MW by 13 &
36% respectively. MTL of A 9 also shorter (11%)
than Ad, but as a proportion of MWL negligibly
(2%) longer. MTL of A and Id 7% longer than
MTCL, but in Ad 71% shorter. å do not acquire
significantly longer outer rectrices with age, Id
only 4% shorter than Ad, but do gain a progres-
sively grossly longer central pair. MLL of A9
11% shorter than in Ad, but as a proportion of
MWL similar in both sexes. MBL of A9 496
shorter, but as a proportion of MWL negligibly
(2%) longer. MFPL of Ad averages 214+ 57mm
(n = 96).

The central rectrices of SAG of all Paradisaea
species show immense variation. Some 9 -plum-
aged 3, with only traces of adult plumage, have
normal-length 9-like central rectrices or far
longer narrow and pointed ones, often slightly
curved at their tips. Other SAG, showing a
greater degree of Ad plumage, have thin long
feathery central rectrices, often with feathery
spatulate tips, or adult-like wires with spatulate
tips.

The systematics of this species has been some-
what confused as it was long thought to represent
a subspecies of the Greater Bird of Paradise,
Paradisaea apoda. The establishment of com-
mon names for what earlier ornithologists consid-
ered species, but are now considered subspecies
of P. raggiana, compounded the confusion. With
respect to the (nominal) species, the mean mea-
surements are so similar overall as to be of little
diagnostic use, particularly as samples of the
nominate are small. Variation in plumage mark-
ings and colouration is far more helpful.

1) P. r. raggiana Sclater, 1873. Proceedings of
the Zoological Society of London, p. 559. Or-
angerie Bay. Now known from south-eastern
New Guinea (Orangerie Bay to Milne Bay). The
fore-back is yellow, fading as it overlays sepia of
the mantle. Flank plumes of Ad scarlet, MFPL

BIOMETRICS OF THE BIRDS OF PARADISE

258 + 74mm (n = 8). The average tai] length of
15em by Cooper & Forshaw (1977) for the nom-
inate form is excessively long.

2) P. r. augustaevictoriae Cabanis, 1888. Jour-
nal für Ornithologie 36: 119, “Kaiser Wilhelm's
Land’, restricted to Finschhafen. Now known
from the coast of Huon Gulf, the Markham R, and
the headwaters of the Ramu R (at the Uria R).
This subspecies averages overall c. 4% smaller
than nominate, but there is considerable overlap
in size ranges. Flank plumes of Ad orange,
MFPL 204 + 5| mm (n = 33), and c. 20% shorter
than in nominate. Like intermedia but flank
plumes orange, abdomen/belly paler, and yellow
throat collar narrow.

3) P. r intermedia De Vis, 1894, Annual Report
on British New Guinea, 1893-94, p. 105. Kumusi
R, Now known from Collingwood Bay to
Holnicote Bay, Kumusi R, and the lower
Mambare R. Like salvadorii but both sexes with
yellow mantle/back and yellow streaking down
lo upper tail coverts. MWL and MTL of Ad
shorter than nominate and most like au-
gustaevictoriae in size, Flank plumes of Ad scar-
let and long, MFPL 239 + 44mm (n = I8).

4) P. e granti North, 1906, Victorian Naturalist
22: 156. ‘German New Guinea’. Exact range
unknown but somewhere between the Matnbare
R and Salamaua (= Morobe area). An examina-
tion of the majority of pertinent specimens
proved confusing owing to considerable variation
and lack of localities. Given the now better-de-
fined southeastern-most distribution of au
eustaevictoriae and the northwesten-mosl
intermedia, we concur entirely with Rothschild
(1930) and Mayr (1941, 1962) that granti is a
changing, intergrading, population between the
twa (contra Cracraft, 1992). Only four Ad mea-
sured as many specimens lack a location. MWL,
MTL, MTCL, MLL, MBL and MBW 187, 131,
357, 40.4, 37.1 & 10.6mm respectively. Flank
plumes of Ad vary between the scarlet of inter-
media and orange of augustaevictoriae, MFPL
238 + 47mm (these biometric data are excluded
from Table 36).

5) Px salvadorii Mayr & Rand, 1935. Amer-
ican Museum Novitates 814: 11. Vanumai, Cen-
tral Division, Papua, south-east New Guinea,
Now known from south New Guinea from near
the Papuan- West Irian border (Tarara); upper Fly
and Purari valley (including the Wahgi). Like
nominate but sepia mantle of both sexes lacking
yellow, and scarlet of flank plumes less deep.
Cracraft (1992) found evidence of clinal variation
in characters of salvadorii. Similar in mean mea:

179

surements to nominate, but MTCL of Ad 16mm
longer and MFPL slightly shorter (201 + 58mm,
n= 37). The subspecies may be invalid. Further
studies are required.

Paradisaea apoda Greater Bird of Paradise
(Table 37)

AN smaller than Ad, MWL by 17%. Few
weights available for this widely known species
and none fof adults. MTL of AG also shorter
(14%) than Ad, but as a proportion of MWL
slightly (3%) longer. MTL similar to MTOL in
13, but 6% longer in A 9 and 7596 shorter in A d.
d do not acquire longer outer rectrices with ape,
1d only 9% shorter than Ad, but gain a progres-
sively grossly longer central pair. MLL of A?
13% shorter than in Ad , but as a Fontem of
MWL similar in both sexes. MBL of A ? also 555
shorter, but as a proportion of MWL 3% longer.
MFPL 196 + 56mm (n= 31).

Central tail rectrices of SA d show great varia-
tion, ås in P. raggiana.

Gilliard's (1969) average Ad tarsal length of
57mm for the species 15 presumably erroneous as
it 15 exclusively longer than ours, and Cooper &
Forshaws' (1977) tarsal lengths for nominate are
extremely long, being exclusively longer than
ours for AG.

Considerable confusion exists in the synonomy
of this species and that of P. raggrana because the
latter was long treated as a subspecies of P. apoda.
The considerable differences in size, 9 plumage
and head: and back and flank plume markings and
colouration in Ad leave few doubts that the pres-
ent treatment is. correct, particularly as P. apoda
does not intergrade with £ raggiana, but merely
produces odd hybrid individuals where they
meet. Two subspecies of P. apoda have long been
established and widely accepted, but Cracrafl
(1992) suggested that the two forms are not diag-
nosticålly disünct with respect to the supposed
paler and more maroon upper breast of P a.
novdeguinea, He apparently did not take the
clearly different sizes of the two populations into
account, the mainland birds being much smaller
thal those of the Aru Is. with no or little overlap
in size (LeCroy, 1981). We accept both subspe-
cies.

LI P a. apoda Linnaeus, 1758. Systema Natii-
rae, ed, 10, 1; 110, ‘India’ (= Aru Is, Irian Jaya).
Only known from the Aru Is, MFPL of Ad 210+
7mm (n= 14),

2 P. à, novaeguinea D' Alberus & Salvadori,
1879. Annals Museo Civico Genova 14: 96, Mid-

180

dle Fly R (300-450 miles upstream). This main-
land subspecies has been long accepted on the
basis of resembling the nominate but with the
upper breast paler and more maroon and being
smaller. MWL and MTL of Ad and AG 11 &
14% and 12 & 16% shorter than nominate respec-
tively. MLL and MBL of Ad and AN also less,
by 11 & 16% and 8 & 9% respectively. Moreover,
MWL of AN as a percentage of Ad in the nom-
inate is 88% and MTL 93%, whereas in d
novaeguinea the respective proportions are 85 &
88%. MFPL of Ad 185 X 40mm (n = 18).

Cracraft (1992) considered supposed plumage
characters not diagnosably distinct. We found
mainland A 9 to be (almost exclusively) smaller
than nominate A 9 in wing and tail length and to
average a good deal smaller than the allopatric
island nominate population in other measure-
ments, We therefore retain the subspecies
novaeguinea.

Paradisaea minor Lesser Bird of Paradise
(Table 38)

A9 smaller than AG, MWL and MW by 15 &
35% respectively. MTL of A 9 also shorter (12%)
than in Ad , but as a proportion of MWL negligi-
bly (2%) longer. MTL of A9 and Id 8 and 596
longer than MTCL respectively, but in Ad 73%
shorter. d do not acquire significantly longer
outer rectrices with age, Id only 4% shorter than
Ad, but gain a progressively grossly longer cen-
tral pair. MLLof A 9 11% shorter than in Ad, but
as a proportion of MWL similar in both sexes.
MBL of A9 4% shorter, but as a proportion of
MWL 3% longer. MFPL 216 + 53mm (n = 59).

Central rectrices of SAG show immense vari-
ation, as in P. raggiana.

Four allopatric subspecies have been widely
accepted (Mayr, 1941; Gilliard 1969). Mean mea-
surements are so similar overall for the subspe-
cies minor, pulchra and finschi as to be of little
diagnostic use. Variation in plumage and coloura-
tion is far more helpful.

1) P. m. minor Shaw, 1809. General Zoology -
Aves, 7, pt. 2, p. 486 ‘New Guinea’, restricted to
Dorey, Vogelkop. Now known from west New
Guinea eastward in the north to Humboldt Bay
and in the south to Etna Bay. MFPL of Ad 215+
60mm (n = 23).

2) P. m. finschi Meyer, 1885. Zeitschrift für die
gesammte Ornithologie 2, 383. Karan, between
Aitape and the mouth of the Sepik, at longitude
142°30°E. Now known from northern New Guinea
from the Aitape region and Sepik Valley eastward

MEMOIRS OF THE QUEENSLAND MUSEUM

to Astrolabe Bay and the Upper Ramu R. Yellow
shoulder marking less extensive than in other
subspecies. On average similarly sized to nomi-
nate but flank plumes brighter orange-yellow and
MFPL slightly shorter (203 + 40mm, n = 25).

3) P. m. jobiensis Rothschild, 1897. Bulletin of
the British Ornithologists' Club 6, 46. Jobi I (=
Yapen I). Only known from Yapen I. Generally
larger than nominate, MWL, MTL and MLL of
Ad 6, 8 & 8% longer respectively. Flank plume
colour as nominate and MFPL slightly longer
(250 + 55mm, n= 11).

4) P. m. pulchra Mayr & de Schauensee, 1939.
Proceedings of the Academy of Natural Sciences
of Philadelphia 91: 151. Tip, Misol I. Only known
from Misool I. Like nominate but brown of plum-
age said to be more purplish, less reddish
(Gilliard, 1969). MWL, MTL and MLL of A å
very slightly larger than nominate, but samples
too small for meaningful comparisons, and there
is considerable overlap in all size ranges between
the two forms.

The availability of 11 specimens from Misool I
allowed Mees (1965) to re-assess the validity of
P. m. pulchra, originally described from only two
birds (Mayr & de Schauensee, 1939). Mees con-
cluded that Misool I birds did not differ consis-
tently from birds of the adjacent mainland (P. m.
minor). LeCroy (1981) subsequently presented
measurements suggesting that, while Misool I
birds are a little larger than adjacent mainland
birds, there is considerable overlap. Our measure-
ments show far less difference in size between the
two forms, birds of Misool I being no larger than
those of the Vogelkop and certainly not even close
to ‘large as in jobiensis’ (Gilliard, 1969). In any
event it appears that this subspecies should be
synonomised with minor:

Paradisaea decora Goldie's Bird of Paradise
(Table 39)

Paradisea decora Salvin & Godman, 1883. Ibis
1883, p. 131. Fergusson I. Only known from
D'Entrecasteaux Archipelago (Fergusson and
Normanby Is). Monotypic. A9 slightly smaller
than Ad, MWL by 11%, but only one Ad MW
known. MTL of A 9 also shorter (7%) than Ad,
but as a proportion of MWL 4% longer. MTL of
A9 and Id 17 & 15% longer than MTCL respec-
tively, but in Ad they are 68% shorter. å do not
acquire longer outer rectrices with age, those of
Id similar to Ad, but gain a progressively grossly
longer central pair. MLLof A 9 1096 shorter than
in Ad butas a proportion of MWL similar in both

BIOMETRICS OF THE BIRDS OF PARADISE

sexes. MBL of A negligibly (2%) shorter but as
a proportion of MWL negligibly (2%) longer.
MFPL of Ad 203 + 42mm (n = 14).

Central tail rectrices of SAG show great varia-
tion, as in P. raggiana.

Paradisaea rubra Red Bird of Paradise
(Table 40)

Paradisea rubra Daudin, 1800. Traité
d'Ornithologie (Lesson). 2: 271. 'New Guinea' in
error for Waigeu I. Monotypic. A9 slightly
smaller than Ad, MWL and MW by 10 & 21%
respectively. MTL of A also shorter (4%) than
Ad, butas a proportion of MWL 4% longer. MTL
similar to MTCL in I , but 8% longer in A 9 and
79% shorterin Ad . å do not acquire longer outer
rectrices with age, I å as AG, but gain a progres-
sively grossly longer central pair. MLL of A 9 796
shorter than in Ad, but as a proportion of MWL
similar in both sexes. MBL of A9 also 396
shorter, but as a proportion of MWL only negli-
gibly (2%) longer. MFPL 94 +15mm (n = 21).

Central tail rectrices of SAd show immense
variation, as in P. raggiana.

Paradisaea guilielmi Emperor Bird of
Paradise (Table 41)

Paradisea guilielmi Cabanis, 1888. Journal für
Ornitholgie 36: 119. 'Kaiser Wilhelm's Land'
(restricted to the Sattelberg, Huon Peninsula,
PNG). Monotypic. A9 smaller than AG, MWL
by 11% smaller, but no weights available for
comparisons. MTL of A also shorter (796) than
in Ad, but as a proportion of MWL slightly (396)
longer. MTLof A Ẹ and IG very slightly (2 & 396)
longer than MTCL respectively, but in AG 80%
shorter. 3 acquire a progressively grossly longer
central pair with slightly shorter (3%) outer rec-
trices but our samples of Id too small for mean-
ingful comparisons. MLL of A 9 9% shorter than
in AG, but as a proportion of MWL similar in both
sexes. MBL of A9 negligibly (2%) shorter, but
as a proportion of MWL slightly (2%) longer.
MFPL of Ad 77 X 38mm (n = 20).

Paradisaea rudolphi Blue Bird of Paradise
(Table 42)

A9 smaller than Ad, MWL and MW by 5 &
13% respectively. MTL of A? 13% longer than
in A d and as a proportion of MWL 10% longer.
MTL of AG and Id similar to MTCL, unlike
congeners, but in Ad 80% shorter. å acquire
shorter outer tail rectrices with age, Id 16%

181

longer than Ad, while simultaneously gaining a
progressively longer central pair, contrary to con-
geners, with the possible exception of P. guiliemi.
MLLand MBLof A 9 4 & 2% shorter than in Ad ,
but as a proportion of MWL similar in both sexes.
MFPL 88 + 16mm (n = 26).

Central rectrices of SAG show great variation,
as in P. raggiana.

Three subspecies are recognised. Mean mea-
surements are so similar overall as to be of little
diagnostic use and samples for two subspecies
(ampla and margaritae) are small. Variation in
plumage markings and colouration are more helpful.

1) P. r rudolphi (Finsch, 1885). Zeitschrift für
die gesammte Ornithologie 2: 385, pl. 20.
Hufeisengebirge, south-east New Guinea. MFPL
of Ad 84+ 13mm (n =17).

2) P. r ampla Greenway, 1934. Proceedings of
the New England Zoological Club 14: 1. Mt
Missim, Morobe district. Similar to nominate but
head lighter, more brownish, less blackish in Ad
(Gilliard, 1969), Some measurements of ampla
are on average only slightly smaller than nomi-
nate but with much overlap. MFPL of Ad 90 +
25mm (n = 3). In view of comments by Gilliard
(1969), Cooper & Forshaw (1977) and Cracraft
(1992) regarding the questionable validity of P. r.
ampla, it seems that merging this subspecies into
nominate P. r. rudolphi is overdue.

3) P. r. margaritae Mayr & Gilliard, 1951.
American Museum Novitates 1524: 11. Kimil R,
20 miles west-northwest of Nondugl, Wahgi Val-
ley, Central Highlands, Mandated Territory of
New Guinea. Like nominate but 9 with under-
parts uniformly and narrowly barred blackish.
MTL and MLLof Ad and A 9 on average shorter
than in the nominate, but samples too small for
meaningful conclusions. MFPL 101 + 10.3mm (n
— 4). Only one or two birds of the Tari Valley area
have been collected, and it is not known to which
subspecies they should be allocated. We did not
examine them.

PARADISAEA SPECIES. We found our figures
for the various Paradisaea taxa in close agree-
ment with those of Lecroy (1981), considering
the often gross differences in sample sizes. We
cannot compare our ‘total bill length’ with ‘ex-
posed bill length' (see Methods). Small differ-
ences in measuring techniques between ourselves
and LeCroy (1981) are reflected by negligibly to
slightly longer tail lengths and slightly (2-5mm)
larger tarsus measurements than ours. In calculat-
ing the MTL as a proportion of MWL, these
differences might, therefore, become significant.

182

Differences between LeCroy's measurements of
'unplumed males’ and ours for ‘immature males’
suggest that the former may have included males
with signs of A d plumage, other than flank plumes,
Our ‘immature males’ class included only 2-plum-
aged birds showing absolutely no sign of å plumage.

SYSTEMATIC CONCLUSIONS

The following systematic list includes taxa ac-
cepted by the above review. Genera and their
sequence are those of Beehler & Finch (1985),
with the addition of thosetaxa extralimital to New
Guinea. Species and subspecies, the latter pre-
sented chronologically, are those of Gilliard
(1969); those marked with an asterisk have been
described since Gilliard (1969). Names in paren-
thesis are those of subspecies included in Gilliard
(1969) that were invalid (L. s. psetdoparotia) ot
are not accepted herein. For consistency herein
and ease of reference with that work only spelling
of genus and specific names are as in Gilliard
(1969). It should be noted, however, that a few of
of these have been subsequently corrected.
PARADISAEIDAE
CNEMOPHILINAE
Cnemophilus:

C. m. macgregorii, C. m. (kuboriensis)
sanguineus, C. l lortae, C, l amethystina, C. 1.
inexpectata

Loboparadisea:

L; s. SETTE PU, L. 5. aurora

PARADISAEINAE

Macgreégoria:

M. p. pulchra, M. p. carolinae

Lycocorax:

L. p. pyrrhopterus, L. p.moroteinsis, L. p. obiensis
Manucodia:

M. a. ater, M. a. alter, M. à. subalter, M.
chalybata, M. c. comrii, M. c. trobriandi, M.
(rubiensis) jobiensis, M. Kk. keraudrenii, M. k.
gouldi, M. k. jamesii, M. k. hunsteini, M. k. (may-
eri) purpureoviolaceus, M. E neumanni, M. k.
adelberti, M. k. aruensis*, M. k. diamondi"
Paradigalla:

P. (intermedia) carunculata, P. brevicauda
Priloris:

P. paradiseus, P. victoriae, P. m. magnificus, P. m.
intercedens, P. m. alberti

Semioprera:

S. w wallacei, S. w. halmaherae

Seleucidis:

S. m. melanoleuca, S. m. auripennis
Epimachus:

MEMOIRS OF THE QUEENSLAND MUSEUM

E. a. (geisleri, inversus) albertisi, E. a. cervini-
cauda, E. bruijnii, E. f. fastuosus, E. f. (stresemanni)
atrains, E. f. ultimus", E. m. meyeri E. m.
(megarhynchus) albicans, E. m. bloodi

Astrapia:

A. nigra, A. s. splendidissima, A. s. (elliottsmithi)
helios, A. mayeri, A. 8, stephaniae, A. s. (ducalis)

feminina, A. rothschildi

Lophorina:

L s. superba, L. s. niedda, L. s. (sphinx) minor
(L. s. pseudoparotia), L. s, (connectens, addenda)
latipennis, L. s. feminina

Parotia:

P. wahnesi, P. sefilata, P. e. carolae, P. c.
berlepschi, P. c. meeki, P. c. chalcothorax, P. £
chrysenia, P. c. clelandiae, P. I. (fuscior, exhibita)
lawesii, P. L helenae

Preridophora:

P. ( burgersi, hallstromi) alberti

Cicinnurus:

C. r. (rex, gymnorhynehus) regius, C. r. (similis,
cryptorhynchus) coccineifrons, C. m. magnificus,
C. m. (intermedius) chrysopterus, C. m. hunsteini,
C. respublica

Paradisaea,

P. r. raggiana, P. r. augustaevictoriae, P. r. mier:
media, (P. © granti), P. r. salvadori, P. a. apila,
D a. novaeguinea, P. m. (pulchra) minor. P. m.

finschi, P. m. jobiensis, P. decora, P. rubra, P.

guilielmi, P. r. (ampla) rudolphi, P. r. margaritae

ACKNOWLEDGEMENTS

For much appreciated permission to access and
help with the study of ornithological collections
in their care we sincerely thank the following
persons and institutions: Steve Van Dyck, Glen
Ingram and Carden Wallace, Queensland Mu-
seum. Brisbane; Richard Schodde, fan Mason
and John Wombey, Australian National Wildlite
Collection, CSIRO, Canberra; Walter Boles and
Wayne Longmore, Australian Museum, Sydney;
Les Christidis and Rory Q'Brien, Museum of
Victoria, Melbourne; Philippa Horton, South
Australian Museum, Adelaide; Glen Storr and
Ron Johnstone, Western Australian Museum,
Perth; Barry Baker, Belinda Deumann, Austra-
lian Bird and Bat Banding Scheme, Canberra;
Dean Amadon, Chris Blake, Walter Bock, Joel
Cracraft, Mary LeCroy, Manny Levine and Les-
ter Short, American Museum of Natural History,
New York; Robert Prys-Jones, Michael Wallers
and Peter Coulston, Ornithology, The Natural
History Museum, London; Ned Johnson, Mu-
seum of Vertebrate Zoology, University of Cali-

BIOMETRICS OF THE BIRDS OF PARADISE

fornia, Berkeley; David Willard, The Field Mu-
scum, Chicago; Raymond A, Paynter Jr., Mu-
seum of Comparative Zoology, Cambridge; Allen
Allison and Carla Kishinami, Bishop Museum,
Honolulu; James Dick and Brad Millen, Roya!
Ontario Museum, Toronto; Frank Gill, Academy
of Natural Sciences, Philadelphia; James M.
Loughlin, The Carnegie Museum of Natural His-
tory, Pittsburgh; Fred C. Sibley, Peabody Mu-
seum., Yale University, New Haven; Gene K.
Hess, Delaware Museum of Natural History, Wil-
mington; Mohammad Amir and Darjono, Mu-
seum Zoologicum Bogoriense, Bogor, Frank
Bonaccorso, Haiah Bigilale and Paul Wanga, Na-
tional Museum and Art Gallery of Papua New
Guinea, Port Moresby; Clem Fisher, National
Museums & Galleries, Liverpool; Josefina
Barreiro, Museo Nacional de Ciencias Naturales,
Madrid; Giuliano Dona, Museo Civico, Genova;
Rene Dekker and Peter Van Dam, Nationaal
Natuurhistorisch Museum, Leiden; Per Ericson,
Gunnar Johansson and Göran Frisk, Swedish Mu-
seum of Natural History, Stockholm; Jon Fjeldsá,
Zoologisk Museum Københavns Universitet,
Københaven: Eric Pasquet, Museum National
d'Histoire Naturelle, Paris; Siegfried Eck,
Staatliches Museum für Tierkunde, Dresden;
Josef H. Reichholl, Ornithalogy, Zoologische
Staatssammlung, Mönchen; Burkhard Stephan,
Zoologisch Museum, Berlin; R. van den Elzen,
Museum Alexander Koenig. Bonn, for kindly
forwarding skins to Frankfurt. Claus König,
Staatliches Museum für Naturkunde, Stuttgart,
for kind hospitality in addition to acces to collec-
tions; D. Stefan Peters, Joachim Steinbacher,
Martina Küsters and Karin Böhm,
Forschungsinstitut und Naturmuseum
Senckenberg, Frankfurt; H. Hoerschelmann, Zoolog -
isches Institut und Zoologishes Museum, Harnburg.

People who &indly provided hospitality, inter-
est and help in other ways include Joan Airey,
Manon Buchanan, Brian and Del Coates, Jeff and
Barbara Davies, Paul and Luisa Frith, Peter and
Daphne Fullagar, Andrew and Trish Gillison and
Peter and Janet Marsack, Sincere thanks to the
Chapman Fund and the Fund Committe for the
support enabling us to examine the remarkable
and fine collections of the American Museum of
Natural History. We sincerely thank Walter Boles
and Ralph E. Molnar for helpful constructive
comment and editorial work,

Dedicated to the memory of Charles Walter De
Vis (Rev.), (1829-1915), Curator at the Queens-
land Museum, who described several birds of
paradise.

183

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184

1992. Standardwing Bird of Paradise Semioptera
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In press. Huia (Heteralocha acutirostris:
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FRITH, C.B. & COOPER, W.T. 1996. Courtship dis-
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FRITH, C.B. & FRITH, D.W. 1990. Discovery of the
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1996a. Description of the unique Parotia lawesii x
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1996b. The unique type specimen of the Bird of
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In press. The taxonomic status of the bird of paradise
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1969. 'Birds of Paradise and Bowerbirds'.
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GILLIARD, E.T. & LECROY, M. 1961. Birds of the
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MEMOIRS OF THE QUEENSLAND MUSEUM

1968. Birds of the Schrader Mountain region, New
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GYLDENSTOLPE, N. 1955. Notes on the collection of
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JOHNSGARD, P.A. 1994, ‘Arena Birds — Sexual Se-
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LAMBERT, F.R. 1994, Notes on the avifauna of Bacan,
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LECROY, M. 1981. The genus Pardisaea — display and
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MACGILLIVRAY, W. 1918. Ornithologist in North
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MAJNEP, LS. & BULMER, R. 1977. 'Birds of my
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MAYR, E. 1936. New subspecies of birds from the New
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1941. ‘List of New Guinea Birds-a systematic and
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1962. ‘Family Ptilonorhynchidae and Para-
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MAYR, E. & de SCHAUENSEE, R. 1939. Zoological
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56.

BIOMETRICS OF THE BIRDS OF PARADISE 185

Bower-Birds'. Part 1 (Southeran: London).

TABLE 1. Measurements (mm) and weights (g) of specimens of

Cnemophilus macgregorii

Wing

Tail

Tarsus

length length length head length width

Males: adult
Mean

subadult
Mean

SD

Min

Max

n
immature
Mean

Females: adult
Mean

SD

Min

Max

n

Males: adult
Mean

SD

Min

Max

n
subadult
Mean

SD

Min

Max

n
immature
Mean

SD

92
2.48

2.79

C. m. macgregorii

40.8
1.82
378
45.8

23

C. m. sanguineus

41.2
146
38.0
44.3

26

SIBLEY, C.G. & AHLQUIST, J. E. 1990. ‘Phylogeny
and Classification of Birds. A Study in Molecular
Evolution'. (Yale University Press: New Haven).

STRESEMANN, E. 1923. Dr Burger's ornithologische
Ausbeute in Stromgebiet des Sepik. Ein Beitrag
zur Kenntnis der Vogelwelt Neuguineas. Archiv
für Naturgeshichte 89 no. 7: 1-96; no. 8: 1-92.

1930. Welche Paradiesvogelarten der literatur sind
hybriden ursprungs? Novitates Zoologicae 36: 6-15.

1934. Vier neue unterarten von paradiesvogeln. Or-
nithologische Monatsberichte 42: 144-146.

WALLACE, A.R. 1869. ‘The Malay Archipelago’.
(Macmillan: London),

APPENDIX 1
Min 10 8 389 52 260 50
Max 19 97 433 534 297 66
n 16 15 16 2 16 16 1
Females: adult
Mean m 89 402 498 260 561 93
SD 361 377 154 007 149 033 1302
Min wm 80 376 497 230 48 7
Total Bil Bill Weight Max 15 97 427 498 280 64 125
n 2 2 2 SS: 4p 9 9
length
C. m. kuboriensis
Males: adult
Mean 12 — 88 416 296 53 94
549 291 57 —98 SD 096 096 092 136 026
149 155 037 481 Min nm pp 406 280 50
S32 241 49 %4 Max 13 89 427 309 56
591 320 65 104 " h å á K 4 i
17 24 21 4 immature
Mean 112 8 395 272 56 97
544 304 58 hi 1 1 1 1 1 1
105 044 049 Females: adult
533 297 853 Mean 109 — 83 390 256 52
554 307 64 SD 283 283 0.49 000 0.14
3 4 4 Min 10 81 386 256 Si
Max mi 85 393 256 53
532 298 55 8 5 2 2 2 2 2
1.15 133 036
519 279 50 all subspecies
541 313 59 Males: adult
3 6 7 1 Mean M4 o Ai 547 288 57 100
SD 259 267 162 164 140 043 937
521 271 56 89 Min 107 86 378 526 241 48 o
091 141 044 Max 118 o 458 591 320 65 120
509 250 50 n 6 eo 54 2 5 50 M
533 294 63 subadult
7 10 10 1 Mean 113 90 414 540 296 58 86
sD 284 366 188 096 095 039
Min 107 84 383 532 277 53
Max 18 95 438 554 307 64
528 283 57 105 Mg n vw mn 5 10 10 1
025 1.13 047 1156 ee
526 258 48 94 Mean 13 91 412 530 282 56 Dë
531 300 64 120 SD 290 379 129 099 148 042 808
3 22 24 5 Min 107 | 8 389 519 260 50 8
Max 19 103 433 541 313 66 o
533 291 59 8 n 24 2 24 5 23 24 3
0.14 089 035 Females: adult
532 277 53 Mean mp 90 399 516 263 56 o
534 301 63 SD 368 450 174 130 149 036 11,77
2 6 6 1 Min 10 80 357 497 230 48 7
Max 115 101 427 533 294 64 125
527 27 56 87 n 34 33 33 9-— d "äm
099 11 047

TABLE 2. Measurements (mm) and weights (g) of specimens of

Cnemophilus loriae

Wing Tail Tarsus Total Bill Bil Weight
length length length head length width
length

C. I. loriae
Males: adult
Mean 103 75 373 533 266 6.5 83
SD 215 255 160 139 137 037 7.41
Min 99 7) 343 511 214 56 76
Max 107 81 42 555 294 74 94
n 29 29 28 15 29 28 7
subadult
Mean 100 74 36 25.8 6.4
n 1 1 1 ] 1
immature
Mean 102 77 370 512 258 62 89
SD 120 293 125 071 148 022 985
Min 101 71 352 505 224 6 81
Max 104 81 383 521 272 6.6 100
n 9 9 9 4 9 9 3
Females: adult
Mean 102 77 36.6 515 26.1 6.4 86
SD 239 371 156 097 082 040 621
Min 98 72 342 496 243 57 78
Max 106 84 404 534 275 ZA 96
n 18 18 18 12 18 18 8

C. I. amethystina
Males: adult
Mean 105 77 370 525 261 6.3 93
SD 2.14 208 149 113 065 034 8.95
Min 101 73 323 507 247 56 80
Max 109 81 391 538 275 68 101
n 25 25 25 7 25 25 5
subadult
Mean 106 80 372 533 265 6.3 98
SD 198 311 089 187 084 036
Min 102 75 359 513 249 5.7
Max 108 85 387 55 279 69
n 12 13 13 3 13 13 ]
immature
Mean 103 80 369 532 264 6.5 82
SD 252 273 1.11 057 069 043 1461
Min 100 75 348 525 249 5.7 66
Max 107 86 389 541 276 7,3 100
n 14 14 14 7 13 14 6
Females: adult
Mean 103 80 369 515 259 63 786
SD 294 367 149 137 076 0.35 11.18
Min 98 73 34 496 243 6 69
Max 109 88 399 538 272 72 95
n 20 20 20 7 19 20 7

CL inexpectata
Males: adult
Mean 103 71 375 526 264 6.2 79
SD 272 261 140 097 093 O31 435
Min 97 66 347 51.1 252 55 75
Max 107 77 400 538 287 6.7 85
n 25 25 24 6 24 24 4
subadult
Mean 103 7 373 529 262 6.4
SD 266 278 097 000 08) 023
Min 99 6 364 529 253 6
Max 107 76 389 529 276 6.8
n 8 8 8 3 8 8
immature
Mean 104 76 383 530 270 6.1 87
SD 304 325 073 095 093 047

MEMOIRS OF THE QUEENSLAND MUSEUM

Min 100 70 369 522 25.1 52
Max 109 81 392 546 283 7
n 13 13 12 5 13 13
Females: adult

Mean 102 73 364 513 260 6.3
ES 234 263 117 092 092 042
Min 97 69 344 497 240 5.1
Max 108 79 386 525 280 L2
n 27 28 28 10 28 28

all subspecies

Males: adult

Mean 104 74 37.3 530 264 6.3
SD 244 353 150 127 1.05 0.37
Min 97 66 323 507 214 55
Max 109 81 42 555 294 74
n 79 79 77 28 78 77
subadult

Mean 104 76 37.2 531 26.4 6.3
SD 264 539 091 120 081 031
Min 99 66 359 513 249 57
Max 108 85 389 550 279 69
n 21 22 22 6 22 22
immature

Mean 103 78 37.4 526 265 6.3
SD 253 348 121 109 110 042
Min 100 70 348 505 224 52
Max 109 86 392 546 283 73
n 36 36 35 16 35 36
Females: adult

Mean 103 76 366 51.4 26.0 6.3
SD 256 440 137 103 084 0.39
Min 97 69 34 496 240 5.1
Max 109 88.1 404 538 280 72
n 65 66 66 29 65 66

TABLE 3. Measurements (mm) and weights (g) of specimens of

Loboparadisea sericea

Wing Tail Tarsus Total Bill Bill
length length length head length width
length
L. s. sericea

Males: adult
Mean 94 55 31.1 446 20.4 6.1
SD 232 242 108 100 104 056
Min 90 52 288 432 188 5.2
Max 98 59 335 465 230 72
n 17 16 16 9 16 15
immature
Mean 97 60 318 436 204 6.3
SD 115 432 083 070 0.19
Min 96 56 307 19.8 6
Max 98 66 327 214 64
n 3 4 4 1 4 4
Females: adult
Mean 98 58 319 472 214 74
SD 183 271 083 007 058 049
Min 96 55 31 471 208 6.5
Max 101 63 333 472 222 76
n 6 6 6 2 6 4
immature
Mean 97 60 314 457 20.5 6.5
SD 227 196 079 046 046 0.30
Min 92 56 304 454 19.7 61
Max 99 63 326 46.2 214 69
n 10 10 10 3 9 9

9.08

101

Weight

BIOMETRICS OF THE BIRDS OF PARADISE 187

L. s. aurora
Males: adult
Mean 97 59 313 455 212 65 62
ES 191 145 091 1,08 101 040 887
Min 93 56 294 432 198 5.8 50
Max 100 61 326 472 242 7.3 75
n 16 16 16 n 16 14 12
immature
Mean 96 63 306 46) 21.4 6.2 70
sD 07) 163 219 028 028 007 520
Min 95 62 290 459 212 EN 63
Max 96 64 321 463 216 62 75
n 2 2 2 2 2 2 4
Females: adull
Mean 97 63 309 470 227 6.6 73
SD 513 061 151 055 040 021 283
Min 9] 62 298 465 221 64 70
Max 101 63 326 476 233 68 75
n 3 3 3 3 3 3 2
immature
Mean 99 63 318 462 213 72 70
SD 324 337 088 144 053 028 881
Min 93 59 303 436 202 67 e)
Max 102 6? 328 478 216 76 77
n 7 7 $ 6 7 7 3
all subspecies
Males: adult
Mean 95 57 312 451 208 6.3 64
ES 268 274 098 111 109 051 894
Min 90 52 288 432 188 52 50
Max 100 61 335 472 242 73 75
n 33 32 32 20 32 29 14
immature
Mean 96 61 314 453 207 62 68
5D li0 3279 134 1456 077 016 543
Min 95 56 290 434 198 50 63
Max 98 66 327 463 216 64 75
n 5 6 6 3 6 6 5
Females; adult
Mean 7 59 315 471 21,8 6.9 73
SD 300 326 111 040 085 043 283
Min 9] 55 298 465 208 64 n
Max i01 63 333 476 233 76 75
n 9 9 9 5 9 7 2
immature
Mean 98 él 316 46.0 20.9 68 72
SD 294 323 082 119 060 044 7.10
Min 92 % 302 434 197 6.1 Ka
Max 102 5 328 478 216 å 77
n 17 17 17 9 16 16 5

TABLE 4. Measurements (mm) and weights (g) of specimens of

Mucgregoria pulchra

Wing Tail Tarsus Total Bill Bil Weight
length length length head length width

length

M. p. pulchra
Males: adult
Mean 202 161 613 700 394 6.1 256
5D 578 315 224 265 157 026 13.37
Min 187 156 546 658 369 55 242
Max 211 168 646 745 424 55 274
n 19 18 19 10 18 18 å
Females: adull
Mean 183 147 Béi 685 378 59 198
sD 435 306 092 222 029 862
Min 175 143 851 350 56 190
Max 187 152 577 417 564 207
n 7 7 6 1 H H 3

M. p. carolinae
Males: adult
Mean 192 134 644 737 416 6,0 349
SD 527 395 127 205 214 023 1202
Min 184 128 620 708 373 $4 34
Max 200 141 659 765 440 61 357
n 10 10 10 5 10 8 2
Females: adult
Mean 172 124 583 682 378 5.6 230
5D 413 463 278 118 167 02
Min 164 120 529 603 356 5
Max 179 133 613 693 403 6
n 10 10 10 5 10 10 1

all subspecies
Males: adult
Mean 199 1531 624 712 402 6,1 279
5D 728 1361 246 301 206 026 4452
Min 184 128 546 658 369 54 242
Max 211 168 059 765 440 65 357
n 29 28 29 15 28 26 8
Females: adult
Mean 77 134 574 682 378 57 206
ER 718 1227 247 107 185 034 17.63
Min 164 120 529 663 350 50 190
Max 187 152 613 693 417 bå 230

n 17 17 16 å 17 17 4

188

TABLE 5. Measurements (mm) and weights (g) of specimens of
Lycoecrax pyrrhopterus

Wing Tai Tonus Total Bill Bil Weight
length length. length head length width
length

L p. pyrrhopterus
Males: adult
Mean 190 143 420 776 466 MA 269
ES 8,66 841 12) O64 1465 O74 2205
Mir 178 127 387 769 48 96 242
Max 206 160 440 753 491 132 304
n 28 28 27 5 27 28 D
Females: adult
Mean 186 l39 417 756 446 109 250
ED Sin &B9 147 172 193 074 1922
Min 176 131 389 731 408 94 218
Max 197 183 447 772 488 120 276
n 27 27 27 7 27 27 7

L. p. morolensis
Males: adult
Meon 219 1568 492 555 1248
ER 387 436 09 073 097
Min 214 152 481 545 114
Max 223 161 505 562 133
n 4 4 4 4 4
Females; adult
Mean 209 145 465 529 116
5D 058 200 223 058 Q15
Min 209 143 448 526 11,4
Max 210 147 a 536 117
n 3 3 3 3 3

L. p. obiensis
Males: adult
Mean 204 145 455 819 523 1209 332
ES 759 549 157 237 251 001 2554
Min 190 135 430 785 490 n 300
Mox 224 186 493 867 562 13 370
n 26 26 26 13 26 26 9
Females: adult
Mean 198 141 451 796 $503 118 291
ES 591 636 171 151 214 062 2747
Min 186 130 422 777 449 104 280
Max 208 162 477 820 544 126 316
n 19 19 19 5 19 19 5

all subspecies
Males: adult
Mean 198 145 441 807 498 11.79 302
sD 2794 19.19 545 1760 653 653 8499
Min 175 127 387 769 418 96 242
Max 224 161 505 867 562 133 370
n 58 58 Eu 18 57 58 17
Females: adult
Mean 192 140 433 775 3473 13 267
so 925 609 241 261 367 080 3058
Min 176 130 389 731 408 94 218
Max 210 153 490 820 544 126 316
n 49 a 49 13 ae 49 12

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6.
Manucidia atra
Wing fail Tarsus Total
length length length heod
length
M. a. atra
Males: adult
Mean 181 150 384 680
ER 999 1098 189 274
Min 160 133 345 615
Max 204 170 42! 727
n 25 25 25 14
immature
Mean 173 144 387 689
5D 586 1002 161 1.48
Min 164 123 361 674
Max 185 158 408 71.4
n 1) n n 6
Females: adult
Meon 175 143 381 664
5D 949 1159 155 258
Min 159 127 354 62
Max 194 168 414 722
n 27 27 27 19
immature
Mean 167 137 369 634
SD 773 940 196 187
Min 156 125 318 61]
Max 182 155 397 67]
n 17 17 17 9
M. a. alter
Males: adull
Mean 204 172 452 787
5D 313 616 126 158
Min 198 163 429 759
Max 208 182 467 804
n 7 7 7 5
immature
Mean 195 161 444 799
n 1 1 1 1
Females; adult
Mean 191 160 426 748
5D 737 762 10d 159
Min 182 152 209 nN8
Max 201 172 242 764
n Wu n n 6
M. a. subalter
Males: adult
Mean 195 165 412 725
5D 8.76 1055 213 225
Min 175 144 347 673
Max 21 183 447 765
n 29 29 28 23
immature
Mean 184 155 404 712
so 918 1064 174 384
Mm 172 140 374 658
Max 199 168 420 749
n 7 7 7 5
Females; adult
Mean 185 154 397 689
SD 842 794 152 1.95
Min 173 138 366 651
Max 205 168 433 729
n 26 26 26 ?0

Bill Bill
length width
38.9 8.9
235 065
34.3 7.8
446 10,7
25 25
39.1 SA
203 058
36.8 83
427 1085
11 1
37.5 8.9
203 044
34.1 75
426 10
27 27
36.1 8.7
243 075
327 72
417 10
7 y
46.3 96
037 066
458 84
468 104
7 7
449 9,8
1 I
429 93
086 059
40.8 Bé
442 103
H H
41.8 9.2
159 058
386 82
441 104
29 29
39.6 92
188 D58
376 B4
427 102
7 7
39.1 90
122 0.80
369 72
411 102
25 26

Measurements (mm) and weights (g) of specimens of

Waight

224
30.82
170
315
17

BIOMETRICS OF THE BIRDS OF PARADISE

immature
Mean 188 160 416 711 399 97
sD 84| 599 100 280 196 033
Min 177 153 403 672 369 93
Max 200 168 428 755 430 101
n 6 6 6 6 & 6

oll subspecies
Males; adult
Mean 190 160 405 717 412 9.1 238
5D 1204 1311 289 404 296 064 3688
Min 160 133 345 615 343 78 170
Mox 211 183 467 804 468 107 315
n 51 ål oo 42 6) ål 23
immature
Mean 178 149 397 708 396 94 225
ES 965 1152 2.11 400 227 057
Min 164 123 361 658 368 BA
Mos 199 168 444 799 449 105
n 19 19 19 12 19 19 i
Females; adult
Mean 182 151 395 686 391 90 208
SD 1041 1162 214 347 248 O71 2192
Min 189 127 354 620 341 72 (155
Max 205 172 442 764 442 103 252
n Ki bA åå 45 63 bA 24
immature
Mean 172 143 381 645 371 B9 177
ES 1195 13.39 274 446 283 079 2354
Min 186 125 38 611 327 72 153
Max 200 168 428 755 430 101 200
n 23 23 23 15 23 23 3

TABLE 7. Measurements (mm) and weights (g) of. specimens of

Manucodia chalybata

Wing

Tail

Tarsus

length length length

Males; adult

Mean 173
5D 5.42
Min 161
Max 183
n 37
immature

Mean 169
SD 5.58
Min 157
Max 179
n 23
Females: adult
Mean 167
SD 662
Min 153
Max 182
n 27
immature

Mean 161
5D 5,34
Min 150
Max 178

Total — Bill Bill
head length width
length

Weight

14]
703
128
152
37

133
502
120
144

70.1 404 9.1
230 174 053
645 376 80
757 444 100

25 36 37
700 402 8.9
246 238 058
674 339 77
752 441 102

10 23 23
662 383 8.7
175 186 056
518 344 77
683 425 99

15 26 27
66.0 370 8.7
275 204 047
602 340 80

71 418 95

189

TABLE 8. Measurements (mm) and weights (g) of specimens of

Manucodia comrii

Wing Tail Torus Total Bill Bil Weight
length length length head length width
lenath

M. c. comni
Males: aduli
Mean 244 173 508 903 576 115 448
SD 658 574 122 183 186 059
Min 229 162 484 858 532 104
Mox 257 182 8527 939 610 127
n 23 22 23 19 23 23 1
immature
Mean 233 164 505 518 115
n 1 i 1 1 1
Females: adult
Mean 231 165 493 850 532 iil! 418
ER 499 394 171 281 179 0680
Min 223 158 463 820 508 102
Max 240 170 S814 90! 569 123
n 13 13 13 10 13 13 1
immature
Mean 219 163 483 838 514 112
SD 709 841 072 240 224 024
Min 207 149 472 821 496 109
Mox 224 Wi 490 855 547 115
n 5 5 5 2 5 5

M. c. frobriandi
Males: adult
Mean 226 158 495 855 548 IL
5D 7,333 631 133 284 181 084
Min 213 148 467 806 516 101
Max 235. 169 516 880 565 123
n 14 14 14 7 14 14
Females: adult
Mean 217 154 484 837 528 108
SD 459 760 177 235 2% 035
Min 209 147 463 812 499 102
Max 223 172 520 878 571 ma
n 9 9 9 4 9 9
immature
Mean 204 149 454 804 497 99
n 1 1 1 1 1 H

all subspecies
Males: aduli
Mean 237 167 503 891 Á 566 114 448
SD 119 966 139 30) 231 Déd
Min 213 148 467 806 516 101
Max 257 182 527 939 610 127
n 37 36 37 26 a7 37 1
immature
Mean 233 164 50.5 5.8 ILS
n 1 1 1 1 1
Femoles: adult
Mean 225 160 489 845 531 110 48
SD 863 792 175 266 200 054
Mn 209 147 463 B12 499 102
Max 240 172 520 wi 571 123
n 22 22 22 16 2 22 1
immature
Mean 217 160 478 827 511 11.0
SD 894 9535 135 260 212 056
Min 204 149 454 804 490 99
Max 224 i3 490 855 547 115
n 6 6 5 3 6 6

190

TABLE 9. Measurements (mm) and weights (g) of specimens of

Manucadia jobiensis

Wing Tol Tarsus Total Bill Bill Weight
length length length head length width
length
M. j .jobiensis
Males; adult
Mean 184 140 376 739 40.4 97 245
SD 613 616 131 039 073 1697
Min 178 132 361 aD 89 233
Max 192 147 393 409 106 257
n 4 4 4 1 4 å 2
Femoles: adult
Mean 174 130 368 657 372 Kä 205
50 141 283 064 007 071 057
Min 173 198 363 656 367 a7
Max 175 132 372 657 377 95
n 2 2 2 2 2 2 Ì
M. i rubiensis
Males: adult
Mean 175 132 371 685 387 92 222
ES oëd 704 174 291 226 070 823
Min 1å1 119 34 631 346 76 212
Max 194 146 405 734 424 105 232
n 26 25 25 21 26 26 6
immature
Meon 170 128 366 664 384 90 197
5D 410 424 078 103 125 082
Min 167 123 356 655 374 8,3
Max 174 132 33 675 38 78
n 4 4 4 3 3 4 1
Females: adult
Meon 188 127 361 656 370 92 174
ER 494 437 123 256 134 066 2496
Min 157 13 338 612 4346 78 150
Max V7 14) 392 (695 391 IWI 200
n 24 24 24 19 24 24 4
immature
Mean 166 119 353 622 355 88
5D 424 212 014 318 148 049
Min 169 120 352 599 344 BA
Max 167 130 354 644 365 DÅ
n 2 2 på 2 2 2
all subspecies
Males: adult
Mean 174 133 3721 687 390 93 228
ES 724 74] 170 307 220 07! 1430
Min 161 19 340 6431 34.4 764 212
Max 194 147 405 739 424 106 28
n 30 29 29 22 30 30 8
immature
Maon 170 128 2366 664 38.4 90 197
5D 310 424 078 103 125 0682
Min 167 123 356 655 374 8.3
Max 174 132 373 675 398 9B
n å 4 4 3 3 4 ]
Females: adult
Mean 169 127 23561 £56 320 92 180
ER 501 620 120 243 130 064 2574
Min 187 113 335 612 346 78 150
Max 177 l4 392 695 391 101 205
n 26 26 26 21 26 26 5
immature
Maan 166 19 353 622 355 88
E 424 212 Old 316 148 049
Min 1b 17 382 899 34 84
Max Wé) 120 354 644 365 9)
n 2 2 2 2 H H

TABLE 10. Measurements (mm) and weights (g) of. specimens of

MEMOIRS OF THE QUEENSLAND MUSEUM

Manucodia keraudrenii

Wing Tal Tarsus Total Bil Bil Weight
length length length head length width
length

Mk keraudrenii
Males: oduli
Mean 160 124 340 625 33.5 8.0 149
sD 484 547 155 154 131 070 1406
Min 144 MO 302 601 31.1 66 130
Max 164 133 370 647 363 98 175
n 24 24 24 là 24 24 7
immature
Mean 155 125 348 627 332 7.7
SD 606 245 121 059 09 040
Min 148 122 329 618 32] 69
Max 185 129 367 63) 350 82
n B 8 B 4 B å
Females: adult
Mean 152 117 329 598 316 77 136
5D 516 546 102 152 162 045 476
Min. 139 108 312 575 288 7;0 1320
Max 158 125 347 62] 36 85 140
n 15 15 15 7 15 15 7
immature
Mean 145 17 313 598 329 7 126
n T 1 1 1 ! 1 ]

M. k. gouldi
Males: adult
Meon 165 135 368 626 337 70 161
SD 464 380 131 147 104 036 1281
Min 155 — 124 338 603 318 62 150
Max 177 142 392 657 357 77 184
n 35 35 35 17 32 33 b
immature
Mean 155 137 375 637 352 6.7
n ] 1 1 1 1 1
Females: adult
Mean 158 127 350 601 37 74 138
sD 379 233 115 158 104 D47
Min 181 122 328 566 301 53
Max 166 130 377 614 338 EK
n lår 16 15 B ló 16 1
immature
Mean 147 124 33.66 591 31.9 68
SD 513 230 1,42 180 108 D.74
Min 142 122 314 8723 305 60
Max 154 127 355 509 332 7.6
n 5 6 5 3 5 5

M. k. jamesii
Males: adult
Mean ló4 128 358 641 343 82 187
EN A432 4657 149 265 171 053 495
Min 155 19 325 871 297 74 183
Max 171 138 387 682 354 98 190
n 26 26 26 14 2 26 2
immature
Mean 143 114 333 595 319 49 175
5D 071
Min 174
Mox 175
n ! 1 ! 1 1 D 2
Females: adult
Mean 160 125 356 617 333 79 160
$b 523 534 188 142 162 054 1768
Min 148 119 318 £02 310 73 147
Mox 166 135 387 63.4 371 93 172
n 13 13 3 4 13 13 2

BIOMETRICS OF THE BIRDS OF PARADISE

immature
Mean 151 123 340 61.4 326 7.5
SD 6.15 346 132 181 145 063
Min 142 118 320 587 310 6.6
Max 159 128 359 638 357 8.5
n 9 9 9 8 9 9

. k. hunsteini
Males: adult
Mean 188 139 392 67.3 366 8.5
SD 371 307 08! 107 080 059
Min 179 133 371 658 352 73
Max 194 143 406 686 376 9.7
n 13 13 13 8 13 13
immature
Mean 176 133 400 660 359 8.2
SD 404 058 049 014 050 057
Min 174 133 397 659 354 77
Max 181 134 406 66.1 36,4 8.8
n 3 3 3 2 3 3
Females: adult
Mean 182 135 379 629 348 8.3
SD 260 3.64 162 167 122 0.36
Min 179 132 350 614 325 79
Max 186 141 397 647 368 89
n 8 8 8 3 8 8
immature
Mean 172 131 2375 62.6 337 8.4
SD 294 419 146 085 066 058
Min 169 127 2363 617 329 7.7
Max 175 137 395 634 343 9.1
n 4 4 4 3 4 4

M. k. purpureoviolaceus

Males: adult
Mean 168 127 370 660 35.4 8.0 190
SD 394 504 1.14 172 154 051 2561
Min 157 15 2344 615 310 70 159
Max 174 138 388 689 384 93 240
n 35 35 35 26 34 35 10
immature
Mean 166 126 378 647 353 7.6 182
SD 624 171 073 049 083 0.37
Min 159 124 372 643 344 73
Max 174 128 386 65 364 8
n 4 4 4 2 4 4 ]
Females: adult
Mean 162 123 358 633 33.6 7.6 162
SD 628 471 165 157 107 035 18.13
Min 148 113 315 603 316 67 140
Max 180 133 382 671 365 82 182
n 29 29 29 21 29 29 7
immature
Mean 157 125 354 627 331 7.9 152
SD 379 115 176 153 112 095
Min 154 124 343 615 323 70
Max 161 126 374 64.4 34.4 89
n 3 3 3 3 3 3 1

Lk neumanni
Males: adult
Mean 156 124 33.1 61.7 329 7.7 157
SD 3.53 351 069 279 182 037 421
Min 150 119 320 587 302 70 152
Max 162 129 343 658 364 82 162
n n 1 n fe} 10 n 5
immature
Mean 145 17 339 60 318 8.2 136
SD 424 424 078 028 028 007
Min 142 114 333 598 316 8.1
Max 148 120 344 602 320 8.2
n 2 2 2 2 2 2 1

Females: adult

Mean 149 117 332 589 31 29
SD 321 459 192 062 05) 042
Min 142 110 300 582 304 73
Max 152 123 348 597 321 8.6
n 8 8 8 4 8 8
M. k. mayri
Males: adult
Mean 169 132 36.7 664 359 7.8
SD 365 407 096 092 061 023
Min 164 128 350 650 349 74
Max 175 138 382 673 368 8.1
n 7 6 7 5 6 7
immature
Mean 160 125 365 622 323 7.4
n 1 1 1 1 1 1
Females: adult
Mean 160 126 360 635 333 7.5
SD 638 616 153 072 077 033
Min 151 120 343 627 324 71
Max 166 135 382 645 343 79
n 5 5 5 5 5 5
immature
Mean 155 124 354 61.7 323 7.4
ES 551 5.13 010 173 181 058
Min 151 118 353 602 310 7
Max 161 128 355 63.6 344 8.1
n 3 3 3 3 3 3
M. k. adelberti
Males: adult
Mean 163 130 339 33.0 79
SD 3.89 5.42 276 121 062
Min 156 122 278 30.8 7.
Max 168 139 370 348 92
n 10 10 9 9 10
Females: adult
Mean 157 126 325 32.1 7.5
n 1 1 1 1 1
immature
Mean 146 121 3445 55.1 30.85 7.45
SD D 141 064 219 0.21
Min 146 120 340 293 73
Max 146 122 349 32.4 76
n 2 2 2 1 2 2
M. k. aruensis
Males: adult
Mean 168 136 355 635 336 7.6
SD 206 263 173 057 044 0.37
Min 166 132 342 631 330 73
Max 170 138 380 639 339 B.1
n 4 4 4 2 4 4
immature
Mean 158 134 372 624 332 8.0
SD 400 702 066 130 082 049
Min 154 127 366 61.1 325 74
Max 162 141 379 637 341 8.3
n 3 3 3 3 3 3
M. k. diamondi
Males: adult
Mean 171 129 374 652 35.6 7.8
SD 582 487 170 085 148 051
Min 156 121 342 637 334 73
Max 176 134 400 659 380 90
n 1 n 9 6 n n

191

MEMOIRS OF THE QUEENSLAND MUSEUM

192
immature
Mean 184 12 377 341 68 174
n T 1 1 1 i i
Females: adult
Mean 182 123 352 606 327 7,4 172
ER 383 150 206 219 167 087 ON
Min 159 18) 322 882 309 67 71
Max 167 124 367 625 348 83 172
n 4 4 4 3 4 4 2
immature
Mean 172 128 362 33,6 7.7
n i 1 1 | I

all subspecies
Males; adull
Mean 167 ]30 361 644 344 7.8 171
SD 832 639 1210 248 172 067 1979
Min 144 110 278 571 297 og 130
Max 194 143 406 689 384 98 240
n 174 175 173 100 169 174 ay
immature
Mean 159 127 366 628 338 77 168
SD 1009 6.41 207 201 149 054 1831
Min 142 114 329 595 316 67 136
Max 181 141 406 66) 364 88 182
n 24 24 24 16 24 24 5
Females: adult
Mean 160 124 351 618 328 74 185)
5D 923 653 206 219 163 053 1477
Min 139 108 300 566 288 63 13
Mux 186 141 397 67,1 37.1 7.3 182
n Kei 99 98 $5 b D 36
immature
Mean 155 125 348 611 324 7.5 141
SD 992 439 184 226 140 074 1608
Min 142 n na 58] 293 60 126
Max 175 137 395 f44 357 9.1 163
n 28 28 28 22 28 28 5

TABLE 11. Measurements (mm) and weights (g) af
Paradigalla carunculata

Wing
length

Males: adult

Mean 186
50 505
Min 180
Max 201
n 10
immature

Mean 71
SD 735
Min 140
Max 179
n 7
Females: adult
Mean 165
D 6.66
Min 157
Max 175
n n

Tail

Bill

length central length head length

Tal Tarsus Total

length length
160 49.0
1061 077
132 479
170 802
10 9

133 479 696

929 224 071

121 444 69]

148 498 701

7 H 2

132 456 679
47) 292
15 la
138 502

9 11 1

specimens of

Bill
width

Weight

170

TABLE 12, Measurements (mm) and weights (g) of specimens of
Paradigalla brevicauda

Wing "oi Toil Tarsus Total — Bil Bil Weight
length length central length head length width
length length

Moles: adult
Mean 158 53 Sl 440 716 442 5.6 173
SD 384 952 763 106 O78 165 04 12,12
Min 151 44 42 415 694 396 Ag 160
Max 168 88 73 456 744 489 62 184
n 25 25 24 25 8 23 25 à
immature
Mean 159 78 78 432 696 44.0 5.8 169
ER 366 1604 1400 159 238 212 032 586
Min 148 54 53 398 671 41 52 160
Max 164 107 106 463 738 480 6,6 175
n 30 29 26 30 14 27 30 4
Females: adult
Mean 150 68 ai 419 709 444 59 163
ER 296 1550 1278 110 142 149 035 1061
Min 144 457 53 391 673 421 52 155
Max 154 96 9) 439 723 478 65 170
n 26 26 23 26 12 25 25 2
immature
Mean 153 89 78 424 70 447 63
5D 071 566 1556 085 0.78
Min 152 85 67. 418 44,1
Max 153 93 89 430 452
n 2 2 2 2 i 2 2

TABLE 13. Measurements (mm) and weights (g) of specimens of

Ptiloriy paradixeux

Wing Toil Tal Tarsus
length length central length
length
Males: adult
Mean 160 98 89 33.3
sD 275 217 220 134
Min 153 92 84 301
Mas 165 — 103 95 362
n Ai 37 37 35
subadult
Mean 154 96 94 339
3D 244 170 162 097
Min 151 94 9) 31019
Max 158 100 97 352
n 13 12 12 13
immature
Mean 148 94 91 32.9
SD 635 377 286 119
Min 139 ag 88 314
Max 154 98 93 345
n 5 4 3 5
Females: adult
Mean 144 91 87 323
5 507 345 337 14)
Min 137 86 83 295
Max 156 99 96 356
n 26 26 23 24

Total Bill Bill
head length width
length

80. 54] 64
2.19 248 042
750 493 56
844 616 72
21 34 36
80.3 $41 65
150 113 034
775 527 58
818 567 69
7 12 13
805 54.5 6.0
458 348 033
7713 819 55
853 606 64
4 5 5
829 585 6.1
223 297 038
768 8M 52
858 658 70
15 25 26

Weiaht

112

BIOMETRICS OF THE BIRDS OF PARADISE

TABLE 14. Measurements (mm) and weights (g) of specimens of

Puilaris victoriae

Wing Tall Tal Torsus Total Bil
length length central length head length
length length

Moles: adult
Mean 139 DN 77 321 686 432
5D 305 775 211 100 132 124
Min 133 77 72 298 655 403
Max 148. B5 Bi 446 717 467
n al 5l 51 5I 40 EN
subadull
Mean 134 83 8] 324 674 429
Es] 363 177 214 101133 183
Min 125 8) 79 307 5 400
Max 138 BG Bò 342 694 457
n n 10 9 1i 7 n
immature
Mean 129 81 79 323 674 427
ER 39] 229 200 139 085 1.35
Min 121 75 74 297 662 403
Max 134 85 83 345 686 458
n 25 23 20 24 " 24
Females: adult
Mean 125 78 74 303 689 447
SD 500 359 284 155 185 22
Min 115 72 72 280 642 401
Mox 136 85 83 331 723 49
n 34 33 31 31 27 33

TABLE 15. Measurements (mm) and weights (g) of
Ptilorix magnificus

Wing Tail Tol Tomus Tote! Bil
length length central length head length
lenath length

i P. m. magnificus
Males: adult
Mean 192 102 97 396 BBA 597
ES 454 407 454 109 206 237
Min 182 92 88 372 847 542
Max 200 10 08 423 912 635
n al Ki 29 29 IO ål
subadull
Mean 184 109 108 401 861 597
sp 722 283 682 113 368 300
Min 7 105 97 379 835 56
Max 196 ma 116 405 6887 656
n 9 9 7 3 2 9
immature
Meon 177 109 0 399 899 589
ER 407 378 277 1314 134 263
Min 169 — 102 106 359 882 504
Max 183 118 0" 42 97 620
n 20 20 18 20 å 20
Females: adull
Mean 157 99 99 355 808 526
SD 522 480 541 O90 215 278
Min 147 HI 90 331 776 Mo
Max 185 la 115 359 838 569
n 25 25 25 24 ü 25

Bil Weight
wath
6.0 105
035 713
EX 9!
69 119
51 22
60 105
0.34
55
65
[E i]
59 93
035 423)
53 82
65 104
25 22
59 86
030 587
54 7
65 96
32 25

specimens of

Bil Weight
width
69 207
049 1361
58 180
80 230
I W
73 182
0,58
63
7,8
B I
72 180
038 828
66 168
82 190
20 9
70 142
0,38 19.58
62 120
79 185
25 8

193
P. m. alberti
Males: adult
Mean 181 99 91 379 823 550 60 140
5D 319 284 314 101 138 151 029 1204
Min 174 Då Bj 356 775 %99 56 143
Max 187 305 98 al 85 573 58 1
n a 38 35 39 19 34 Ki å
subadull
Mean 171 102 102 378 6811 547 59 137
5D 575 50 525 133 391 155 053
Min 160 94 932 351 71" 505 53
Max ez gg 107 395 837 56,3 75
n 14 14 8 14 a 14 14 1
immature
Mean 161 lat 102 383 B05 541 60 131
5n 712 an 26! 080 319 174 029
Min 181 7 98 3/75 74i 50.8 56
Max io 104 ID 393 823 56 6,3
n H 5 å 5 4 å 5 1
Females; adull
Mean 149 96 93 35] 7&6 506 58 104
Si 764 610 565 306 377 296 O25 0907
Min 139 BB 844 911 718 458 54 Qå
Mox 185 108 107 442 824 557 63 112
n 24 zl 16 24 18 23 24 3
P. m. intercedens
Males: adull
Mean 193 101 96 404 841 558 70 — 184
SD 303 292 373 1.44 204 201 039 1820
Min 188 % 90 348 804 8524 &2 155
Max 199 106 105 427 881 619 77 214
n 25 25 25 23 17 24 25 12
subadull
Mean 191 109 102 401 851 550 72 166
ES 436 203 522 148 114 1851 040 4285
Min 1864 107 98 382 838 S20 45 120
Max 197 H3 109424 HBó6 572 78 2
n 9 9 5 7 ò 9 y 3
immature
Mean 1755 108 106 405 832 548 71 176
ER 594 402 3.9) nn 134 153 054 9409
Min 162 103 100 384 815 513 57 168
Max 187 U "3 2425 87 525 82 189
n 20 20 19 19 y 19 20 7
Females; odult
Mean 152 9B 97 352 750 48.0 68 126
5D 446 326 306 1,21 154 131 034 1310
Min 145 92 9| 326 720 453 63 102
Max 162 103 05 381 775 507 je 147
n ?7 27 24 27 20 27 27 15
oll subspecias
Males: adult
Meon 188 101 95 39.) 843 568 64 194
SD 713 339 446 157 293 289 OM 2135
Min 174 92 B3 356 7095 509 56 143
Ma» 200 10 100 427 7312 635 80 23
n 96 74 Di 9) 45 B9 94 36
subadull
Mean 181 106 104 391 832 552 67 163
5D oes 494 589 177 365 30) 084 3432
Min 160 9a 92 351 719 805 53 120
Max 197 "4 16 44 887 6055 78 205
n 32 ER 20 a2 16 32 31 5
immature
Mean 174 10 107 400 843 565 70 175
SD 727 437 418 127 357 298 O56 1421
Min 181 37 98 389 761 506 56 131
Max 187 118 17 425 917 626 82 190
n at 45 ES 44 27 45 45 17
Females; adult
Mean 153 9n 97 353 764 503 6.6 128
5D Déi 486 833 192 337 308 062 1867
Min 139 88 Bhd ALI 718 2446 54 94
Max 165 13 H5 442 838 569 79 185
n 76 73 Ki 78 46 75 76 26

194

TABLE 16. Measurements (mm) and weights (g) of. specimens of

Semioptera wallace

Wing Toil Tail Tarsus Total ` Du Bil Weight
length length central length heed length wiath

jength length

5, w. wallacei
Males: adult
Mean 156 78 66 419 730 451 &6 163
ED 256 461 417 108 137 163 039 1102
Min 152 n oO 394 WA 412 57 152
Max 142 90 al M3 747 475 73 174
n 29 28 27 28 IW 26 25 3
subadult
Mean 156 85 79 422 714 455 6.7
50 243 327 909 064 184 De 0.34
Min 153 79 656 ALI 70,1 44.8 63
Max 159 97 9? 429 727 abd 72
n 6 b 6 é 2 & 5
immature
Mean 150 87 88 413 720 449 67 153
SD 462 158 200 218 152 098 044 354
Min at 85 84 350 702 436 &2 180
Max 156 89 90 435 74) 472 73 155
n 1 n H 1 7 1 11 2
Femalas: adult
Meon 145 B4 B5 399 707 438 67 135
SD 445 349 404 149 095 102 042 8854
Min 135 7a 76 371 691 AE 60 126
Mox 154 94 % 413 722 465 77 145
n 26 25 24 25 17 24 25 3

S, w. halmaherae
Males: adult
Mean 157 85 74 417 722 442 6,7
SD 254 340 627 120 136 179 038
Min 151 m ai 390 703 399 6.1
Max 162 93 91 439 740 478 76
n 32 3l 30 3) 9 4) 30
subadull
Mean 154 Bé 82 419 710 436 6,5
sD 404 192 698 DSS 29) 267 O46
Min 149 BA 74 414 878 415 59
Max 140 89 Wi 426 735 482 49
n 5 5 5 T 3 5 5
immature
Mean 154 b 91 4L5 701 439 72
sD 292 318 328 095 104 148 039
Min 147 84 85 375 688 423 56
Max 187 94 96 424 713 471 78
n B B 8 a 4 7 H
Females: adult
Mean 144 Bé 87 386 700 43 69
SD 330 230 257 100 146 146 042
Min 138 D 84 362 680 406 EN
Max 18] Kai "| 403 /20 457 75
n 25 25 2! 25 13 24 25

oll subspecies
Moles: adult
Mean 157 B2 70 418 727 446 Ai 163
5D 202 536 655 114 148 174 039 1102
Min 152 71 oO 390 703 399 57 182
Max 162 93 93 445 747 478 76 174
n o 5 57 5B 23 57 55 3
subadull
Maan 155 85 80 421 712 446 46
so 323 276 798 059 227 206 040
Min 149 79 66 411) 678 415 59
Max 160 Bo 9) 429 735 482 72
n n n 1 1 5 10 19

immature

Mean 152
SD 427
Min 141
Max 187
n 19
Females

Mean 144
SD 391
Min 135
Max 154
n 51

MEMOIRS OF THE QUEENSLAND MUSEUM

89 414 713
296 173 162
84 359 ORB
% 435 74)
19 19 11
86 393 704
358 140 122
76 362 680
% 433 722
45 EU 30

44.5 69 153
135 047 354
EEN ER 150
472 78 155

a 19 2
43.3 68 135
133 042 854
40,6 60 126
46,5 a 143

48 50 3

TABLE 17. Measurements (mm) and weights (g) of specimens of

Seleucidis melanoleuca

Wing Tail Tol Torsus Total Bill Bl Weight
length length central length head length width

length length

S. m. melanoleuca
Males: adult
Meon 177 n 70 410 1020 710 74 205
sp 427 422 354 219 233 222 Dé 1115
Min 167 62 63 306 954 652 50 190
Max 185 Bá 78 4à5 1059 748 8&3 27
n 34 34 34 33 18 32 Ki 4
subadult
Mean 179 97 100 4|! 1001 705 75 196
SD 532 827 GAB 103 228 180 04
Min 187 D 78 380 950 649 EN
Max 187 "2 12 430 129 731 87
n 22 22 EA 22 9 22 22 1
immoture
Meon 177 110 108 414 1003 701 78 202
SD 458 Alà 474 130 240 252 O47 191
Min 163 102 101 377 963 636 6.4 182
Max 184 125 12] 433 1039 731 88 220
n 22 21 21 22 8 22 22 3
Females: adult
Mean 165 108 108 380 928 645 75 187
SO 3.17 322 400 081 282 253 O88 14)
Min 158 102 101 366 843 5467 64 186
Max 171 13 dh 399 967 686 BY 188
n 21 19 19 20 16 21 20 2

S. m. aunpennis
Males: adult
Mean 168 65 é 393 955 653 68 189
sD 239 264 215 106 21) 180 074 1058
Min 163 6) 62 370 921 625 $7 179
Max 173 70 7 AF 984 682 84 202
n 25 25 25 25 12 25 25 12
subadult
Mean 168 93 92 396 952 662 7.0 188
Ee 359 565 844 108 189 221 048 1579
Miti 164 BS 75 371 928 47 EN 153
Max 179 107 IM 42 v7 715 8,6 200
n 16 16 15 lò ó s 16 9
immature
Mean 166 100 97 391 937 639 69 175
5D 233 424 302 087 319 267 08) 11.88
Mn 161 95 9 367 899 600 EN 158
Max Mo 12 163 40] 988 690 8.4 188
n 15 15 12 15 B 14 14 5
Females: adult
Meon 158 99 98 365 888 605 HI 14)
50 446 445 590 151 102 225 060 173
Min 152 23 9 343 870 555 42 160
Max Té? 111 12 399 898 641 42 163
n 19 19 18 19 6 18 19 3

Malas: adult

Females: adult
Mean

BIOMETRICS OF THE BIRDS OF PARADISE

all subspecies

69 68 403 994
456 370 197 392
öl 62 306 921
B6 7B 435 1059
59 58 58 3
95 97 405 982
743 995 126 325
EU 75 371 928
12 112 430 1029
a8 35 38 15
106 104 404 970
656 660 161 437
95 92 367 899
120 121 433 1039
36 33 37 16
103 103 373 917
573 687 142 305
93 % 343 843
H3 118 399 967

38 37 39 22

68.5 7.1 193
353 073 1257
625 57 170
74.8 84 217

58.7 73 189
289 0.49 1508
637 6.1 153
74.1 87 20

38 38 10
67,7 75 185
395 084 1971
600 81 158
73.1 88 220

36 36 8
627 T3 nm
3,10 0.66 1431
555 62 160
bió B9 188

TABLE 18. Measurements (mm) and weights (g) of specimens of

Epimachus albertist

Males; adult
Mean

Females: adull
Mean

SD

Min.

Males: adult
Mean

SD

Min

Wing Toll Toil Tarsus Total
length length central length heod
length length
E. a. albertisi
155 134 135 341 1010
281 36 332 076 172
151 125 129 328 979
161 140 14) 356 1036
19 19 19 18 B
151 130 133 3475 1027
495 141 212 064
147 129 131 34.3
154 131 134 352
2 2 2 2 1
150 132 133 344 1017
43 529 436 149 317
145 126 128 3313 987
153 136 136 — 36] 105
3 3 3 3 3
152 130 131 348 1073
249 230 263 123 457
150 127 127 335 1044
156 133 133 365 1126
5 5 4 5 3
E. a. cervinicauda
152 126 125 346 976
310 321 360 094 242
145 127 116 322 903
159 13) 130 357 1017

29 25 28 21

Bill Bill Welght
length width

783 562 12

205 06 829

747 43 105

826 bA 125

286 7.78
775 50 105
83.1 52 M6

3 3 2
84.6 61 122
470 025 2333
792 58 105
910 65 138

5 5 2
749 536 12
199 055 638
692 43 103
796 65 120

subadull

Mean 181
50 089
Min 150
Max 152
n 5
immature

Meon 150
n ]
Females; adull
Mean 147
SD 241
Min. 143
Max 153
n 25
Males: adull

Mean 152
SD 354
Min 149
Max 154
n 2
immature

Mean 150
SD. 545
Min 145
Mox 187
n å
Females: adult
Mean 145
Eu 0.58
Min 145
Max 145
n 3
Males: adult

Mean 159
n H
immature

Mean 153
ES 7.81
Mn 144
Max 156
n 3
Females: adult
Mean 151
n H
Males: adull

Mean 153
50 3.45
Min 145
Max 161
n 51
subodult

Mean 181
5D 2.19
Min 147
Max 154
n 7
immature

Mean 181
ES 522
Min 144
Max 158
n 1
Females: adult
Mean 148
sn 299
Min 143
Max 156
n KI

129

123 3438
517 085
116. 333
130 3&3
5 5.
129 342
i |
122 339
445 073
ns 324
132 351
2] 28
E. a. geísleri
128 353
078
347
128 358
1 2
13) 357
Bl) 126
120 343
37 372
4 å
128 339
424 115
125 326
131 348
2 A
E. a. inversus
34.0
|
133 343
115 Lë
122 332
144 359
H 3
128 342
1 i
all subspecies
129 | 344
609 089
118 322
14] 35,8
45 49
126 3449
626 076
114 333
134 353
? 7
132 348
505 133
10 332
137 32
n n
124 340
521 087
115 323
134 365
28 34

100.3
4.18
913

1066

21

5.50

195

no

12
6.87
103
125
12

196

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 19. Measurements (mm) and weights (g) of. specimens of

Epimuchus bruijnit

Wing — Tall Jal Tarsus Total Bill Bl Welgnt
length length central length head length width
length length

Males: adult
Mean 180 110 107 330 1032 790 627 16l
SD 307 329 375 09 264 221 054 189
Min 153 103 100 308 984 735 53 160
Mox lé (ie 115 348 1076 826 73 164
n 28 26 23 26 17 26 27 å
subadult
Mean 161 12 109 331 1035 78,4 63 165
5D 206 329 287 073 O86 142 067 6.36
Min 156 104 103 317 1020 748 56 160
Max 163 | 5 113 341! 1044 805 EN 1689
n 18 18 14 18 6 ló 16 2
Immature
Mean 1537 H5 115 320 1010 750 72
SD 610 45| $65? 094 29N 477 081
Min 1464 109 108 304 959 624 60
Max Io 128 133 335 1054 404 EN
n 12 12 n 12 B 2 12
females: adult
Mean 155 nt 110 32] 98,1 75] 6.8 146
SD 478 291 250 121 319 316 OM 265
Min 148 107 107 302 93 683 54 144
Max 162 (ip 15 339 1024 79] B5 14
n 14 14 12 14 é 12 13 a

TABLE 20. Measurements (mm) and weights (g) of specimens of

Epimachus fustuosus

Total
head
length

Weight

E, f. fastuosus

Wing [ail loi Tarsus
length length central length
length
Males: adult
Mean 197 392 710 $502
SD 424 3765 5705 1,44
Min 188 340 593 465
Max 205 539 790 535
n 24 23 23 23
subadult
Mean 191 274 332 50.67
Es 306 2053 26]! 025
Min 188 257 327 504
Mak 194 297 339 509
n 3 3 3 3
immature
Mean 177 242 304 510
5D 477 1088 1452 122
Min 167 226 283 488
Max 182 287 321 825
n B 8 7 H
Females: adull
Mean 166 229 291 470
"D 1003 1248 1715 217
Min 182 204 270 432
Max 187 249 321 ló
n 12 11 12 2

105.3
2,82
1002
nos
13

Bill Bill
length width
765 | 66
206 049
725 59
80.2 76
22 23
764 69
148 023
75.1 58
78 72
å a
74.5 64
497 037
679 57
1A 6.8
6 B
723 &4
407 055
654 55
77.4 75
10 12

E, t. atratus
Males: adult
Mean 199 39) 712 501 1080 778 672 268
so 765 3223 6302 190 434 392 065 1204
Min 188 342 640 479 1023 714 56 255
Max 214 451 856 547 1148 838 Bü 280
n 23 21 18 23 14 19 20 5
subadult
Mean 197 287 46| 507 1090 78.6 67 250
SD 9.40 4250 10819 293 550 426 073
Mir 187 254 384 467 1019 734 60
Max 208 3s 537 535 1138 825 HI
n 4 4 2 a 4 H A i
immature
Meon 180 250 308 804 73.8 6.6
ER 1228 590 2188 149 693 045
Min 164 242 273 484 64.8 55
Max 193 205 333 823 819 71
n D 5 5 5 5 5
Females: adult
Mean 170 215 278 468 1044 766 68 185
sD 845 2511 3196 263 298 251 058 2500
Min 154 171 214 427 NE 722 58 160
Max 187 257 361 534 1112 815 EN 210
n 25 25 17 25 16 25 25 3
E. f. stresemanni
Males: adult
Mean 216 458 847 538 1131) 826 742 315
ES 71) 3975 S463 142 221 192 052 252
Min 206 421 790 510 1106 #00 66 313
Max 232 537 946 554 1159 858 82 318
n 12 10 9 12 5 12 12 å
subadult
Mean 185 365 737 516 79.4 66
n ! 1 1 1 T 1
immature
Mean 196 267 357 522 1109 803 78 286
Es 843 3499 5164 318 396 270 052 1556
Min 179 207 275 458 1047 746 68 275
Max 205 236 424 8560 1146 834 82 297
9 9 H 9 6 y 9 2
Females: adult
Mean 182 242 322 $00 1090 794 7.5 237
ES 544 157] 3187 255 146 45! O69 26016
Min 176 230 292 473 1046 716 6.0 218
Max 193 280 38 550 1101 885 83 255
n 1 9 10 H 4 10 H 2
E. [, ullimus
Males; adull
Mean 196 | 380 725 504 1019 717 703 278
50 208 768 5208 141 07 136 034 3M
Min 193 374 665 49] 1014 879 66 275
Max 198 391 76) 524 1024 73.) 73 280
n 4 4 3 4 2 å 4 2
immature
Mean 183 264 347 507 983 703 65 239
SD 557 993 2254 107 594 316 027 11.51
Min 172 254 320 49l 941 657 EA 223
Max 186 282 382 520 1028 737 68 252
n 6 6 6 & 2 å È 6
Females: adult
Mean 165 235 320 456 992 709 65 190
5D 253 745 1523 (084 176 113 037 Béi
Min 161 225 307 442 VD éa 60 184
Max 168 248 341 468 1007 728 7.0 207
n 10 9 8 10 4 10 10 å

BIOMETRICS OF THE BIRDS OF PARADISE 197

all subspecies immature
Males: adult Mean 168 204 327 477 1107 855 6.4
Mean 201 402 735 509 107.4 77.9 683 277 SD 343 20.55 3218 503 491 5.14 048
SD 970 4282 7698 214 442 396 062 2111 Min 162 171 280 379 1059 773 58
Min 188 340 593 465 1002 699 56 250 Max 171 232 370 518 1157 895 7
Max 232 539 946 554 1159 858 8.2 318 n 6 6 5 ó 3 5 6
n 63 58 53 62 34 57 59 17 Females: adult
subadult Mean 153 193 275 449 111.2 83.1 6.3 160
Mean 193 292 443 508 1090 779 6.8 250 SD 678 2488 3922 210 596 519 052 1450
SD 7.61 4247 16466 195 550 3.16 051 Min 142 151 213 390 956 69.1 57 145
Min 185 254 327 467 1019 734 6.0 Max 170 225 331 470 11772 884 74 175
Max 208 365 737 535 1138 825 74 n l6 15 9 16 13 15 16 4
n 8 8 6 8 4 8 8 1
immature E. m. megarhynchus
Mean 179 256 331 51.2 1062 754 6.9 237 Males: immature WE
SD 3667 24.10 3946 2.11 670 5.70 0.76 2552 Mean 166 229 250 50.7 82.2 6.3
Min 0 207 273 458 941 64.8 5.5 205 n 1 1 ] 1 1 1
Mox 205 306 424 566 1146 834 82 297 Femolés: adult
n 28 28 26 28 12 26 28 15 Mean 154 221 304 442 1103 809 6.8
Females: adult SD 000 3889 63.64 0.07 255 078
Mean 171 226 298 472 1036 753 6.8 195 Min 154 193 259 44] 79.1 62
SD 9.52 21.98 3205 268 417 428 068 2426 Max 154 248 349 442 1103 827 73
Min 152 171 214 427 956 664 55 160 n 2 2 2 2 1 2 2
Max 193 280 387 550 1112 885 8.3 255
n 58 54 47 58 29 55 58 18 E. m. bloodi
Males: adult
Mean 177 237 670 48.7 1129 844 6.57 187
SD 292 10.58 5106 120 143 193 068 45.69
Min 171 218 595 463 110.1 81.7 5.5 144
Mox 181 264 766 505 1144 885 75 230
n 16 16 16 16 8 13 15 6
subadult
Mean 172 215 437 49.1 86.9 6.4 239
SD 520 700 255 121 076 5.06
TABLE 21.:asurements (mm) and weights (g) of specimens of Min 169 210 46,5 85.8 56 235
Epimachus meyert Max 178 223 516 88.2 71 25
n 3 3 1 3 3 3 2
immature
Wing Tal fail Tomus Total Bil Bil Weight Menn E "ape AA- WEE DINE BO hae
length length central length head length width sp SW, RÉI 18:00 Nr DEE GIK 4049 TOP
length length Min 151 194 306 457 957 684 5.4 190
Max 169 235 348 496 1093 869 7. 200
Tom-meyen n 9 9 7 9 2 8 9 2
Males: adult Females: adult
Mean 185 261 748 511 1149 843 67 281 Mean 19x WAP ge 208, 44. INE CADE. Be” Vien
sp 595 1342 5657 156 301 316 0.50 2037 SD 4.16 12.89 2322 193 3.45 249 050 2716
Min 77 235 631 484 109.8 784 59 253 Min 144 171 262 413 103] 78.2 5.3 140
Max 204 282 822 540 1199 904 29 310 Max 165 226 341 498 1158 887 73 202
n 30 29 23 30 22 25 28 7 n 23 20 18 23 1 23 23 4
subadult -
Mean 182 248 469 510 11596 8655 71 245 all subspecies
3D oL NAA Doi SEA Xen DAN inet Di 182 252 706 502 1146 84.4 6.68 234
ek: p Fag 3 åå are En Sé SD 669 1679 7566 178 279 272 0.59 5802
y 6 6 4 6 5 6 6 1 Min 171 218 522 463 1098 784 55 144
redo, Max 206 282 822 540 1199 90.4 79 310
Mean 173 217 335 510 1122 840 7.0 236 d Ane g s Ae SS 33 de ^ de
ise BEP Hae OA a aod, E ADR 00 BTS Mean 178 237 463 50.39 1160 867 69 241
Min 150 180 265 448 102.1 76.7 58 186
Mox 184 260 372 539 1218 917 29 260 SD 7,18 26.42 168.28 169 362 263 052 529
a 19 18 18 19 9 17 17 4 Min 169 205 308 465 1121 81.4 56 235
Max 186 276 741 519 1205 889 73 245
Females: adult A 9 9 5 9 5 9 9 3
Mean 163 208 306 467 1112 810 6.9 173
SD 1169 1941 3054 356 684 597 066 10.47 immature
Min 147 178 257 420 955 656 59 162 Mean 169 215 329 497 1105 83.1 6.7 222
Max 185 250 359 53.2 1203 905 8.3 188 sD 8.43 19,57 30.18 319 688 505 062 33.68
n 24 24 21 24 16 22 21 6 Min 150 171 250 379 957 68.4 54 186
Max 184 260 372 539 1218 91.7 79 260
E. m. albicans n 35 34 31 35 14 31 33 6
Males: adult Females: adult
Mean 177 247 574 49.5 1168 85.2 7.02 189 Mean 157 201 294 455 1103 818 6.5 166
SD 657 14.40 7186 079 182 237 076 SD 947 2048 3237 277 576 463 064 1730
Min 171 234 522 483 1148 823 62 Min 142 1581 213 390 955 656 53 140
Max 188 265 656 50,1 1184 87.2 79 Max 185 250 359 532 1203 905 83 202
n 5 5 3 4 3 4 5 1 n 65 61 50 65 EN 62 62 14
Max HO. WI UE ee A Ee ns AE eu sD 477 2460 5694 1.38 173 190 U2 1.25
n nu 1! ë n T cM 8 Min 162 142 343 393 630 342 50 13
immature Max 180 231 519 442 0686 43 61 158
Mean 162 203 352 422 653 386 5.6 164 19 19 17 19 n 19 16 5

en £02 59657 184A 117 1AA 143 n aA

198

TABLE 22. Measurements (mm) and weights (g) of specimens of
Astrapi nigra

Wing Tail Tal Torus Total Bil Bil Weight
length length central length head length width
length length

Males; adult
Maan 185 399 569 417 6894 414 64
5p 120 14.16 4573 092 076 036
Min 179 30 518 39.0 40.1 59
Max 193 437 756 433 424 70
n 25 24 23 25 1 22 24
subadult
Meon 182 29 38 422 695 47 $5
Eu 204 2847 2302 147 118 039
Min 179 273 335 398 ad 61
Max 185 347 398 438 43.8 72
n 6 5 5 6 1 4 å
immature
Mean 177 266 327 2417 701 42 &5 190
SD 264 2279 2944 063 191 Lei 03
Min 173 239 299 407 690 383 60
Maz 180 303 37 425 723 431 69
n å å ó 4 4 å 6 1
Females: adult
Mean 170 à 258 310 400 676 4412 6.8
SD 835 1191 1469 202 167 099 030
Min 157; 233 290 372 659 392 EA
Max 182 274 332 434 699 427 72
n 17 18 l4 y 4 17 17

TT SS kw

TABLE 23, Measurements (mm) and weights (g) of specimens of
Astrapia splenddidissime

ese
Wing Tail Tall

Torsus Total ` Bill DI Weight

length length central length head length width
length lenath
A. 5. splendidissima.

Males: adult
Mean 134 161 203 372 655 397 55
SD 175 6407 522 077 072 135 (08
Min 181 148 193 353 6447 377 56
Max 187 175 23 385 ei 424 73
n lå 18 i lė 1 Vi 16
subadult
Mean 135 159 20) 379 640 382 &2
sO 212 2404 849 035 014 0,85
Min 133 142 195 36 38.1 55
Max We 6 207 38] 38.3 åB
n 2 2 2 2 i 2 2
immature
Mean 135 174 206 365 643 396 65
SD 276 1020 i861 094 290 101 025
Min 132 162 181 352 622 389 oi
Max 139 192 283 3756 663 48 59
n 7 å 5 7 2 7 7
Females: adult
Mean 132 176 204 363 644 392 68 120
SD 243 1234 1245 115 124 136 O40
Min 128 166187 327 022 345 59
Max 136 — 204 232 388 660 412 75
n 21 19 20 2| 7 21 21 1

MEMOIRS OF THE QUEENSLAND MUSEUM

A. s. helios
Malas: adult
Mean 138 178 218 377
SD 294 2145 1406 1.46
Min 133 131 198 Gad
Max 145 200 243 309
n 22 18 18 22
subodult
Mean 140 188 216 386
EN 179 557 1793 153
Min 137 182 205 370
Max 142 195 237 402
n 5 å 3 5
immature
Mean 143 200 231 380
EN 269 1402 1316 139
Min 133 165 213 352
Max 147 222 25) 406
n 2å 23 18 23
Females: adult
Mean 137 189 212 368
SD 321 888 903 091
Min 130 173 208 347
Max 145 208 Á 236 388
n 31 29 24 31

A. s. elliottsmithi
Males: adult
Mean 139 199 235 38.4
"D 243 689 778 140
Min 136 189 225 357
Max taz 207 243 398
n 8 5 7 8
immature
Mean 139 197 226 387
ED 330 1089 1068 098
Min 136 182 213 379
Max 143 206 237 401
n 4 4 4 4
Females: adult
Mean 136 200 224 380
sD 298 1201 1439 0.88
Min 132 187 29 367
Max 140 224 249 395
n 10 10 10 10

all subspecies
Malas: adult
Mean 137 174 2] 377
SD A2] 1961 1550 129
Min 131 131 193 344
Max 145 207 243 309
n 46 38 36 a
subadull
Mean 138 178 210 384
50 A09 1873 1579 13
Min 133 142 195370
Max i42 195 287 402
n 7 å & H
immature
Mean 141 195 225 378
SD 4]0 1581 1638 La
Min 132 162 181 352
Max 147 222 251 404
n 34 33 27 KI
Females: adull
Mean 135 187 211 368
SD 356 1334 1347 114
Min 128 186 199 327
Max 145 224 249 395
n 52 58 54 he

68.6

0.45

BIOMETRICS OF THE BIRDS OF PARADISE

TABLE 24. Measurements (mm) and weights (g) of specimens of

Astrapia mayert

Wing Toll Toll Torus Toro! ` Di Bill
length length central length head length width

Weld

length length
Males: adult
Meon 179 111 893 409 611 331 47 M7
E» 274 455 8176 173 137 149 029 1052
Mir 173 97 657 315 588 285 Al 134
Max 185 126 1017 432 636 350 52 WA
n 30 2) 26 29 14 25 23 12
subadull
Mean 172 155 377 414 617 338 50 155
E» 477 1527 6180 165 066 11 029 707
Min 163 118 295 367 608 313 45 15
Max 181 176 580 433 622 369 55 160
n 20 19 19 20 5 19 19 2
immature
Maon 165 162 300 408 604 332 51 148
ER 570 1666 7215 139 126 109 Dän 422
Min 156 134 153 394 892 314 47 143
Max 175 175 380 431 425 348 55 19
n Du B 10 10 H 10 9 5
Females: adult
Mean 156 152 309 394 609 330 50 132
SD 48) 1452 2460 120 107 077 027 19%
Min 150 1722 200 354 588 214 44 102
Max 163 178 374 b 621 349 54 157
n 27 24 24 27 1 25 25 7

TABLE 25. Measurements (mm) and weights (g) of specimens of

Astrapia stephaniue

Wing Tail Tall Tarsus Total ` Di DR Weight
length length central length head length width
length length
As slephaniae

Males; adult
Maan 171 154 640 419 666 38.6 54 160
sb 502 1185 3415 140 Le 174 029 1138
Min 160 132 574 YO 640 338 46 144
Max 182 182 693 454 694 414 58 169
n 32 27 20 w 1B 26 26 4
subodull
Mean 171 199 396 425 658 382 55
Es 474 1932 4865 134 187 1171 029
Min 163 147 343 398 634 357 51
Max IBO 217 502 442 686 413 61
n II Hn 10 n 7 " B
immature
Meon 182 203 352 422 653 386 56 164
$D 593 2557 1836 117 164 143 036
Min 153 14% 316 402 635 359 49
Max o 247 380 2439 6073 423 62
n 1? 18 16 19 B 18 18 1
Females: adull
Mean 156 201 328 404 653 383 57 148
5D 438 1490 2054 149 135 122 032 8%
Min 145 170 279 363 630 Md 49 139
Mar 165 230 368 431 87h 419 hå 159
n 34 30 30 34 22 34 32 4

199
A. s. leminina
Males: adult
Mean 166 154 727 40.95 35 51 150
SD 495 4455 0,54 184 021
Min 182 122 405 337 49
Max M9 185 DÉI 36,3 52
n 2 2 i 2 2 2 1
subadull
Mean 166 191 398 428 379 54 147
5D 212 212 3182 163 078 ooo 1131
Min 164 189 Á 375 416 37,3 54 139
Max lb; 192 40 439 384 54 155
n 2 2 2 2 2 2 2
immature
Mean 162 180 370 406 634 358 54 160
SD 250 2152 4679 141 156 1M 029 35
Miri 158 148 Á 3423 375 6823 333 50 158
Mox 165 201 487 An 645 369 58 164
n v 7 7 9 2 9 9 3
Females: adull
Mean 152 190 Hå 400 640 MS 57 135
5D 616 1954 3145 128 130 088 027 424
Min tdd 151 274 382 629 348 54 132
Mox 165 38 357 426 Gei 39 63 138
n y 7 7 9 5 9 9 2
A. s. ducalis
Males: odult
Mean 166 129 628 416 643 366 53 150
5D 3.89 1422 3776 1,64 127 157 034 450
Mir 166 107 565 376 625 341 47 145
Max 173 158 69) 438 664 400 58 156
n 2 ES 20 24 12 24 23 a
subadull
Mean 166 178 436 415 642 353 52 153
ER 339 3273 7749 133 101 076 020 Aen
Min 162 182 357 33 0630 342 50 18
Mas mm 24 s 43 652 361 55 158
n 5 å 5 å å 6 å A
immature
Mean 158 196 322 418 644 367 53 144
sp 325 3879 2727 169 131 152 029 II
Min 182 1588 24 393 616 332 AB 130
Max 163 225 351 465 664 39 59 157
n 16 16 i3 14 12 16 % 7
Females: adult
Mead 152 185 317 394 650 371 5&6 134
5D 281 1145 2114 135 204 150 037 755
Min 144 157 268 370 S99 34] 47 123
Max 189 21 362 419 674 4D! 65 148
n 30 26 26 n] 17 20 4 13
oll subspecies
Males; adult
Mean 169 144 637 ANB 657 375 63 155
SD 517 1897 3845 149 184 196 031 918
Min 156.107 565 376 625 37 46 144
Mor 182 185 727 454 60B 44 5,8 187
n 5B 49 4l 56 30 52 E 9
subadult
Mean 168 192 408 422 652 372 54 150
sb 477 2460 5694 138 173 190 028 723
Min 62 142 341 303 630 342 SD 13
Max 180 23] AIG 442 084 43 6.1 158
n 19 19 17 19 il 19 16 5
immature
Mean 160 196 345 MLB 646 373 55 150
sp 493 2280 3359 152 15] 180 035 123
Min 152 146 Á 246 375 616 332 48 130
Mox 179 242 467 455 673 423 52 164
n 44 4] 36 dd 22 43 ERI H
Females: adult
Mean 154 193 322 400 651 376 &7 137
sD 468 1589 2262 147 1.66 Län 034 947
Min 1440 1510 268 363 599 341 47 123
Max 168 230 362 431 875 49 65 159
n 7à x] 43 72 44 H 70 19

P aur me wre WE: ew Tre TT
Moles: adult n 2 na 8 2 39 2 27 6
Mean 185 399 549 47 674 Ald 64 subadull
5D 320 1616 4573 092 076 036 Mean M0 18 216 386 652 398 62 138
Min 179 aw 98 30 401 59 sp 179 557 1793 153 028 039 049 17.68
200 MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 26. Measurements (mm) and weights (g) of specimens of
Astrapia rothschildi
L s. minor
Wing Tol Tol Torus Total Bil Bil Weight Males: adult
length length central length head length width Mean 132 85 88 315 555 300 52 83
sengih length so 343 296 333 102 156 130 031 508
Min 126 80 B2 281 534 277 4b 77
Max MD 9 94 333 585 335 62 o
Moles: og n a 5 33 2 90 nm 8
Mean 188 352 443 418 OI 392 67 207 AG
sp 335 3002 3319 138 162 104 040 Ga Mei PPT A
Min 182 271 367 380 650 372 55 184 BA 290 Aine varen "uix "inp «ux
Mak 194 387 486 44] 718 408 72 225 ME US i ål BB Ss *08å BD
d mi *- GB NE wa 134 — 8 o 336 558 321 A
Doc: n v mm 2 9$ ?7 9? Ww i
Mean 178 237 270 423 658 380 ai gegen
ai DON i44 AN 1085 086 d77 Mean vz 8 8) 33 551 296 53 72
Min 178 27 25 417 850 367 e d
Mes da. 2v gd ADM lad» MO 8b 453 434 490 109 05 043 028 793
Min "m D T i 542 WI 48 e
VEER 2-2, MÉI. KE E ow» Max Im o o 326 558 305 58 a
3 ?
Mean là? 225 255 414 648 391 73 197 GH? Zoch bk i ETTE PM sa ht? E
3 BEL 15:08 PTATO ME) Dm UE. ZE "weg ma am mn 287 536 292 53 o
Min 158 202 23 392 38] 67 Io
Mos eae) Sa Å ord Sch KT reg sD 385 381 Am 137 134 118 O24 4.4
M p Å P : j å i Min Mo nm 73 266 509 275 Ai =
Fences daji Max 1% 88 o 321 855 313 57 e
Mean 164 220 256 401 852 393 an 159 P ad is Esa MP ee 3
sD a B n iz 128 128 030 1797 La ipeo
Min 152 208 24 386 624 365 EN 143 Males: adull $^
Se Å :
= m m E Ki iei ME de: = Mean 186 o 95 323 556 23 53 8
sD 284 332 324 093 OFF 09 034 265
Min 132 85 om 304 549 278 49 85
Max M5 oe 102 346 56 37 oi o
n a mm 5 6 2 2 9
subadult
Mean 134 90 a 320 565 295 54 87
SD O71 424 566 O14 156 057
Min 133 87-87 ns 284 — 50
TABLE 27. Measurements (mm) and weights (g) of. specimens of ui E“ K: E: ks i = ne :
Lapharina superba Immature
Mean 129 87 88 304 542 283 52 80
Es 3.61 173 252 072 042 017 02
Wing Tod Tal Torus Total Bil al Weight Min 15 85 e 298 539 281 50
length length cential length head length width Max 12 8 om 312 545 284 53
length length n 3 3 3 3 2 3 2 1
Females; odult
L s. superba Mean n7 82 B3 296 543 29] 53 o
Males: adull D 236 230 257 105 204 107 023 420
Mean M0 — 99 102 312 565 302 52 o Mh ina 78 78 275 522 279 49 S
sp 357 240 317 106 083 097 O46 440 Max 124 8 8 310 5B4 307 57 7
Mn 13 om om 280 553 282 44 8 n Wm B 14 8 14 M 7
Mos 149 104 109 324 8579 325 62 o
n 24 23 20 23 12 2 23 H L s, feminina
subadult Males: adult
Mean 139 102 104 314 308 58 Mean 139 87 88 324 562 307 54 a
5D 153 200 212 049 036 010 sp 295 347 339 11) Dap 090 Dä 1315
Min 138 (0 o 311 304 57 Min 134 — 82 au 297 545 287 48 o
Max äi 104 105 320 311 89 Mox lb 9 w Ma Si 325 5B o
n 3 3 2 3 3 3 n 2 7 »= 3 2 B 2 5
immature subadull
Mean 134 Im 104 312 567 297 55 Meon 139 84 85 331 310 58
sp 230 212 214 047 092 054 057 so 216 245 163 128 026 0.14
Min 129 98 101 304 580 289 Ap Min ur mn 8 320 305 57
Max 135 10 107 317 873 304 43 Max M? Bb pp MS 312. 60
n 7 2 7 7 2 7 5 n 4 4 4 4 4 4
Females: adult immature
Mean 126 — 96 om 300 555 302 56 74 Mean 131 — 85 — 84 319 561 304 57
sD 317 336 427 111 096 139 047 712 sp Am 586 482 125 108 O76 Dän
Min 122 90 o 283 544 22 49 o Min 10 779 786 302 547 289 50
Max 133 wm up 320 570 321 68 85 Mai Im o o 340 576 31 al
n b 1 ib 15 7 i 15 5 n no w å nu 5 "n W

Females: adull

BIOMETRICS OF THE BIRDS OF PARADISE

Mean 122 80 805 299 550 31 57 63
sp 326 328 399 OF] 203 112 044 1202
Min US 73 72 286 523 287 49 54
Max 127 87 B8B 316 584 329 6,6 H
n 18 18 18 i8 8 18 17 2
L s. niedda
Males: adult
Mean 139 100 101 304 582 309 54 103
5D 222 289 330 159 049 03) 040 354
Min 137 97 9H 282 SB W5 50 100
Max 142 102 15 3i7 585 2 ER) 105
n 4 4 4 4 2 4 3 2
Females: adult
Mean 124 100 102 304 559 306 5.5 71
5D 071 354 424 042 021 021 007 07)
Min 123 97 99 301 557 304 54 70
Max 124 102 105 307 560 307 55 n
n 2 2 på 2 2 2 2 2
L s. connectens
Males: adult
Mean 137 88 9 319 559 299 51 87
SD A65 227 286 066 058 086 040 496
Min 128 85 AS 308 553 29.0 46 79
Max 144 D % 330 567 309 60 99
n 12 12 12 12 6 12 12 20
subadult
Mean 133 92 88 3.4 556 295 52 7
ED 071 9,19 283 007 057 007 07
Min 132 85 Bó 313 29.1 EN 76
Max 133 9B NW 314 299 52 7T
n 2 2 H 2 ] 2 2 2
immature
Mean 127 86 8&7 317 552 294 52 77
SD i21 153 153 117 092 095 032 850
Min 123 84 BG 304 548 288 48 bå
Mox 129 87 88 326 558 303 54 83
n 3 3 3 3 2 3 3 H
Females: adult
Mean na 8) 82 302 542 299 54 66
SD 425 364 404 188 138 09 036 3.92
Min 110 77 76 285 517 286 50 61
Max 126 86 88 353 569 320 62 rr
n 12 12 2 n n 12 12 15
L. s. sphinx
Males: immature
Mean 137 9i 96 317 28.2 50
n 1 1 1 1 1 ]
L 3. addenda
Males: adull
Mean 136 87 89 316 557 301 53 87
EN 420 279 341 13) 078 199 033 6413
Min 127 61 BLY 291 550 284 46 78
Max 145 93 96 335 565 327 59 95
n 26 26 25 26 3 26 26 10
subadull
Mean 136 91 92 3215 546 300 EN 79
SD 7JB 424 495 035 156 120 000 636
Mir 130 88 88 319 555 29] 51 74
Max 141 94 9 324 577 308 51 83
n 2 2 2 2 2 2 2 2
immature
Mean 128 86 87 319 549 296 53 78
ES 412 300 299 1.29 1,78 104 032 529
Min 119 80 82 288 $525 278 47 70
Max 135 a 93 347 573 317 59 Bå
[ 19 19 18 19 7 18 18 B

Females: adult

Meun 121 8l 8) 300 546 0294 54 Eu
50 46 232 281 144 150 098 021 542
Mit! Hå 76 75 278 524 277 4B 58
Max 131 85 #8 365 575 314 58 7B
n a 29 29 29 10 29 29 13
all subspecies
Males; adull
Menn 136 90 92 318 561 301 5,3 87
sD 452 576 612 115 i146 111) O35 676
Min 126 80 819 280 534 277 44 Ku
Max 149 104 107 346 585 335 62 105
n 148 — 147 138 145 53 la — 144 62
subadull
Mean 134 88 89 319 554 299 55 78
sD 452 664 561 1.15 128 101 O85 571
Mir 125 80 8) 288 526 284 50 72
Max 142 104 105 349 577 321 6.1 87
n 24 76 24 26 1 26 2 à
immature
Mean 128 87 B9 316 553 297 54 782
SD 546 (32 696 115 124 089 037 6.58
Min 117 7 766 288 526 278 åJ od
Max 137 104 107 347 8576 317 63 85
n 58 E 5 54 26 56 52 17
Females: adult
Mean 120 83 B4 297 544 297 55 67
SD 496 650 733 143 160 109 036 B77
Min 110 H 72 266 509 262 47 54
Max 133 103 110 365 584 329 6B 85
n 17 105 113 15 oa 17 115 52

TABLE 28. Measurements (mm) and weights (g) of specimens of

Purutia walmesi

Wing fail Tal Tarsus Total Bill DI Weight
length length central length head length width
length lenath

Males: adult
Mean 161 187 213 509 596 308 66 171
5D 266 893 69) 141 208 113 044 1,16
Min 187 159 20 485 504 288 56 179
Max 166 198 2235 532 020 327 73 172
n 21 19 jò 2) 9 18 19 3
subadull
Meon 160 161 182 521 597 31.4 6.7
5D HOD 1193 200 070 185 0%
Min 159 148 180 514 29,3 67
Max 161 171 184 528 32,8 68
n 3 3 3 3 ] 3 3
immature
Mean 157 161 175 489 615 3019 6.7 174
5D 287 1034 629 306 173 110 053
Min 152 140 168 428 593 307 2
Max 161 172 184 $517 635 329 75
n B 7 7 B 4 4 4 1
Females: adult
Mean 149 153 166 463 607 341 7.1 148
5p 383 580 360 24 10 123 052 350
Min 142 140 ji 4358 591 276 A 144
Max 157 164 75 521 624 333 78 154
n 20 19 17 20 n 19 17 7

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 29. Measurements (mm) and weights (g) of specimens of

Parotia setilata subodult
Mean 152 92 85 4925 61.1 330 6.8
5D 283 566 035 O64 014
Wing Tail Tai} Tanus Total ` DU Bil Welght Min 90 8l 49 325 ER
length length central length head length width Max 04 B9 495 $3.4 69
length length n 1 2 2 2 2 2
P, c. meeki
Malas; adull Males: adult
Mean 186 128 130 528 871 353 57 192 Mean 153 76 75 508 379 67
ES 2,41 284 327 163 1.34 193 027 1030 5D 150 216 126 066 148 030
Min 141 123 125 499 640 304 50 175 Min 151 73 73 500 36,8 EI
Max 179 133 137 866 689 395 64 205 Max 154 78 76 5185 38.9 7.)
n 32 3ü 29 31 14 31 24 12 n 4 4 4 4 2 4
subadult subadult
Mean 165 131 133 822 672 361 5.1 202 Mean 150 B4 D 503 632 380 69
SD 175 349 441 278 145 088 044 76 ES 405 476 480 127 039 342 DAI
Min 162 126 125 439 656 342 54 198 Min 142 75 75 486 628 341 63
Max 168 139 142 551 696 375 5&9 210 Max 157 ag BG 521 636 42.2 74
n 18 15 14 15 7 13 13 3 n 9 9 ü 9 5 H
immature immature
Mean 162 133 136 529 67.1 365 6.0 183 Mean 148 91 48.1 7
5D 359 403 478 1.61 115 1,16 043 14,8 5D 071 495 2.12
Min 154 123 126 ($00 658 345 54 160 Min 147 87 46.6
Max 147 138 147 554 685 380 68 200 Max 148 94 49.6
n 17 Ww 1? 17 4 16 15 å n 2 2 2 i
Females: adull Females: adult
Mean 152 128 129 475 652 350 6.1 172 Mean 141 91 90 463 622 376 74
5D 4là 317 303 235 124 166 O49 1286 SD 283 261 277 114 143 045
Min 141 121 122 444 629 303 54 140 Min 136 86 B5 446 605 346 65
Max 183 135 134 541 675 370 75 185 Max 146 96 95 479 639 386 81
n 26 25 24 24 18 25 25 14 n 12 12 11 1 7 12
P. c. chalcothorax
Males: odult
Mean 159 77 76 517 anl 336 7,3
sb 141 283 283 OF
, T . , Min mm 75 74 a
TABLE 30, Measurements (mm) and weights (g) of. specimens of Max 160 79 7B 524
Parotia carolae n 2 2 ? 2 1 !
subadull
Mean 157 92 85 513 610 343 77
Wing Tal Tail Tarsus Total BI — Bü Weight ` "9 100 954 50 08 202 02
length length central length head length width DA IS H 58. 505 67 328, 15
length length Max 158 98 90 522 621 358 79
n 3 3 3 3 2 3
P. c. carolae Females: adull
Males: adult Mean 146 93 92 452 589 351 75
Mean 154 75 76 501 615 349 7.2 SD ld 1795 183 142 085 160 094
sp 135 282 156 125 072 223 03 Mn Vë 90 GH NS Va — $5
Min 152 73 73 483 0609 305 57 Max 148 95 94 465 595 362 BS
Mox | — 82 78 526 622 376 8&1 n SWEET "Ze T og
n 12 12 12 12 4 10 12
subadult
Mean 153 85 81 8500 626 373 70 P. c. chrysenia
SD 257 526 616 184 101 175 0.46 Males: adult
Min 148 74 75 481 615 345 62 Mean 159 82 82 526 615 358 74
Max 157 E 95 526 635 401 79 sD o7! 07 071 057 0,57
n 12 12 9 d 3 R ë R Min 158 81 8l 522 7
Females: adult Max 159 82 82 530 78
Mean 143 a o 454. 616 367 73 " 2 2 2 2 | 2
sD 34| 293 281 107 163 172 029 subodull
Min 133 87 87 434 591 337 og Mean 155 94 92 499 601 345 74
Max 146 96 95 47] 634 39] 79 so 208 549 1.53 1.01 162 045
n no 18 m 13 7 R 12 Min 153 88 om 490 588 326 70
Max 157 Kéi 94 50 608 355 79
P. c. berlepschi n 3 3 3 3 3 a
Males: adull immature
Mean 156 77 75 473 579 342 6.8 Mean 153 99 99 495 632 339 7.6 210
SD 000 000 014 2246 064 5D 212 141 07] 205 134 014
Min 156 LE) 75 a2 326 63 Min. 151 98 98 4B 329 75
Max 156 7 75 47.4 35.8 72 Max 154 100 99 509 348 77
n 2 2 2 2 1 2 2 n 2 2 2 2 2 2 1

BIOMETRICS OF THE BIRDS OF PARADISE

Females; adult

Mean 147 95 95 470 616 342 B2 130

sp 465 173 265 090 098 258 035 2113

Min 138 9 Wi 451 601 294 76 110

Max 154 97 99 482 628 370 8.6 182

n 9 9 9 9 7 5 9 3
P. c. clalandiae

Males: adull

Mean 157 80 B0 503 625 319 74 205

5D 450 314 264 168 083 072 053

Min 1497 77 77 486 616 WS 67

Max lol Bò ü3 529 636 327 EN

n 8 8 6 7 4 6 8 i

subadult

Mean 154 B9 87 52 364 69 211

5D 557 751 919 120 Q71 042

Min 148 84 Bo 512 35,9 64

Max 159 98 D 529 369 72

n 3 3 2 2 2 3 ]

immature

Mean 182 98 98 518 636 370 72

5D 058 404 404 1.98 017 010

Min 151 94 94 AQT 36.8 7.1

Mox 152 10 102 536 37,1 73

n 3 3 3 3 1 3 3

Females: adult

Mean 147 94 94 486 623 362 79 163

ER 464 230 179 267 103 137 O46

Min 141 yt 93 459 612 352 71

Max 153 97 9] 57 632 385 8,3

n 5 5 5 5 3 5 5 H
all subspecies

Males: adull

Meon 155 77 7] 93 615 343 72 205

SD 321 33) 280 159 143 245 046

Min 149 73 73 472 8579 305 6,3

Max 161 86 B3 530 636 389 EN

n 30 20 28 29 n 22 29 1

subodull

Mean 153 87 83 503 618 365 70 m

sD 367 646 625 137 148 251 O46

Min 142 74 75 48.1 588 325 ER

Max 159 99 % 529 635 422 79

n 31 32 27 ap 14 26 31 1

immature

Mean 151 96 98 501 634 357 73 210

ER 243 493 292 239 028 186 026

Min 147 87 94 466 632 329 70

Max 154 102 102 536 636 371 77

n 7 7 5 7 2 5 6 l

Females: adult

Mean 144 92 92 463 615 362 76 138

5D 429 289 317 168 152 205 056 240

Min 133 Bé 85 43 579 294 65 110

Max 154 97 D 57 639 391 aa 163

n 44 ad 40 43 26 34 43 4

203

TABLE 31. Measurements (mm) and weights (g) of specimens of

Paroria lawesii

Wing Tail

Bill

length length central length head length width

Males: adult
Mean

subadull
Mean

ES

Min

Max

n
immature
Mean

Females: adult
Mean

Males: adult
Mean

5D

Min

Mox

Females: adult
Mean

ES

Min

Max

n

Moles: adult
Mean

152
365
144
157
13

150
552
144
154
n

150
277
147
154
9

148
371
143
156
14

Tal Tarsus Total
lengt length

P. I, lawesii
80 79 497 636
233 175 186 125
7à 76 457 Gin
84 83 823 671
25 16 23 20
89 88 502 630
661 622 156 0.49
80 76 473 620
100 97 523 637
17 16 16 12
98 98 494 629
332 388 238 180
9] 92 445 605
103 104 3528 660
13 i 12 7
99 9; 478 625
304 346 330 097
94 92 439 613
102 101 $28 643
n 10 LE 10

P.I helenae

80 80 494 645
229 106 169 1.48
76 7B 457 619
84 82 531 688
23 12 22 17
91 79 497 655
1.39 707 114 090
75 74 2485 636
108 Bä 515 665
H 2 6 6
98 | 100 495 63.9
371 252 140 142
92 97 469 614
1026 102 SIO 654
9 3 B 8
99 98 467 651
317 342 178 054
95 95 449 642
107 108 511 659
14 H 14 13

P.I fuscior
80 80 491 628
209 000 123 082
76 80 476 621
82 80 8510 639
7 2 7 6
B4 497 639
1 1 H
96 95 512 621
1 1 1 j

Déi

0.51

0.35

Weight

141

164
14

204 MEMOIRS OF THE QUEENSLAND MUSEUM

Females: adult TABLE 32. Measurements (mm) and weights (g) of specimens of
Mean 147 98 94 440 618 312 5.1 143
sp 477 280 224 101 199 04 837 Pteridophora alberti
Min 141 92 403 605 260 46 122
Max 157 1024 489 628 337 62 169
n 13 13 I 13 å 13 13 55 Wing Tai Tarsus Total Bil Bil Weight
length length length head length width
PL exhibila length
Males: adult
Mean 157 79 80 498 624 331 5,2 175
ES 271 277 173 1.60 125 195 0.68 Males: adult
Min 152 75 7? 470 ol 299 40 Mean 124 B5 322 494 246 52 84
Max 161 B3 82 528 64) 353 55 SD 246 343 139 102 O97 037 378
n 12 12 8 12 H 12 1 1 Min 120 7) 295 479 227 46 BI
subadult Max 129 97 38] 513 265 EN 89
Mean 157 92 90 SLI 32.1 56 179 n 26 26 25 II 25 25 5
SD 283 141 141 D.64 0.42 subadult
Min 155 MI 89 506 318 Mean 124 B7 325 481 234 48
Max 159 93 HI 515 324 n 1 1 1 1 1 1
n 2 2 2 2 2 1 1 immature
immature Mean 120 Bé 325 494 244 53 78
Mean 153 100 100 515 624 326 53 173 Ee 374 353 106 091 iia 032 506
SD 230 334 277 122 202 289 04 638 Min 112 78 304 484 214 48 72
Min 150 94 97 502 595 270 48 168 Max 125 o 344 509 257 59 84
Max 157 104 104 5235 653 35] 59 180 n 12 12 10 5 12 1 4
n 7 7 5 7 5 H H 4 Females: adult
Femalas: adult Mean 113 82 306 48.) 240 55 75
Mean M45 ai % 456 622 318 55 154 sD 195 259 109 112 097 033 493
ES 187 290 239 088 104 228 DA? 1115 Min 108 77 298 456 224 48 68
Min 143 93 92 444 612 27)! 50 139 Mox 17 87 328 496 262 DN ER
Max 147 103 98 469 636 344 65 166 n 25 25 25 13 25 24 8
n 10 10 4 10 4 10 10 4
P. a. burgersi
Males: adult
Mean 125 85 326 485 243 5,3
EN 557 252 168 223 032
all subspecies Min 19 82 215 219 49
Mies; org Max 130 87 345 263 55
Meon 155 80 80 496 636 335 53 167 > 3 3 3 1 3 3
sD 323 236 148 147 147 205 041 755 immolure
Min 148 73 7 457 601 289 40 153 Mean n9 86 312 472 231 53 93
Max 163 D — 83 ai 688 385 bF Im a 556 347 189 106 032 054
n CHE P ME ME I ME MET, Im B 295 462 228 49
Sal Max 124 89 334 483 234 ål
Mean 155 o pg 501 639 334 53 1799 4 FME va^ 23 1
SD 286 750 643 140 133 181 047 58 Femoles: adult
Min 149 75 74 473 620 310 47 174 Mean 114 83 298 479 248 54 73
Max 160 103 97 8523 665 366 EM 185 sD 200 274 Dën 361 05 023
n w 26 D 235 V V U 3 min Mm 79 293 453 234 82
immatura Max là 86 304 504 257 58
Mean 152 99 99 50.0 631 330 5.4 166 "n 5 5 5 2 5 5 1
ES 327 337 349 201 174 188 045 B45
Min 144 o o 445 595 270 44 151 P. ar hallskomi
Max 157 104 104 535 850 354 686 180 males: adult
n m PP a I 2 ZS Å 70 Mem 127 B8 321 505 243 50
Females: gi 5b 338 23 075 162 V11 031 434
Meon 148 98 — 97 460 634 330 56 14 Mn 12 — 8 308 41 2015 44
ER 440 297 314 258 161 236 059 892 Max 133 94 335 525 264 57 95
Min Iw 92 92 403 AH 260 An 12 q 24 25 5 M % 723 8
Mox 159 107 105 528 659 363 73 1 subadull
n m Ap GAS pe MM 04) 01557 Moon 122 ^89 321 502 244 49 e
so 577 354 104 028 088 043
Min 114 B3 306 500 236 43
Max 128 92 333 504 258 55
n 8 5 5 2 5 5 1
immature
Mean 123 321 484 241 5.1 80

Females; adult

BIOMETRICS OF THE BIRDS OF PARADISE

Mean 118 86 310 485 240 57
Sp. 488 34) 73 167 094 0.42
Min 109 80 281 458 231 ER
Max 128 91 340 EA 258 64
n 15 15 15 7 14 14
all subspecies
Males: adull
Mean 126 B6 322 500 244 51
5D 336 326 113 147 110 035
Min 119 79 295 47) 216 4,4
Max 132 97 361 525 266 ól
n 53 54 54 25 52 51
subadult
Mean 123 89 322 49.5 243 49
EN 520 326 095 123 089 039
Min 114 83 306 48] 234 ER
Max 128 92 333 504 258 55
n å & & å H é
immature
Mean 122 88 321 484 241 52
SD 423 359 140 168 103 042
Mini 12 78 284 452 2,4 Ab
Mox 129 95 344 511 26,6 EN
n 40 40 Ai 20 39 3
Females
Mean ns 84 306 482 241 55
ER 393 331 117 148 0947 036
Min 108 77 281 453 224 47
Max 128 9] 34 513 262 64
n 45 45 45 22 44 43

12

6.41

TABLE 33. :asurements (mm) and weights (g) of specimens of

Cicinnuruy reylux

Wing Toil Tall Tarsus Total Bill Bill
lenatn length central length head length width
length length
C. t, regius

Males: adult
Mean 105 35 161 273 503 281 52
5 242 148 757 Dei 125 135 048
Min Kb 3i 181 255 464 259 43
Max 110 38 178 288 54 312 [30]
n 26 26 2 25 24 25 26
subadult
Mean 107 ål 99 271 511 284 56
50 191 1284 3440 069 072 122 030
Min 105 35 59 263 502 270 81
Max 110 68 138 279 520 308 6.1
n 7 7 7 7 7 7 7
immature
Mean 104 60 6| 262 505 275 56
SD 335 219 246 120 069 OFF 034
Min Kë 56 57 229 492 206 EI
Max 110 64 64 275 5|) 283 61
n " 11 II dno 100 100 11
Females: adult
Mean 102 59 60 269 496 277 56
SD 191 183 207 068 112 087 034
Min 98 56 57 259 481 269 50
Max 105 62 63 281 514 302 6,1
n 18 15 14 15 14 15 18

Weight

Males: adult
Mean

SD

Min

Max

n
subadull
Meon

SD

Min

Max

n
immature
Mean

Females; adull
Mean

so

Min

Max

n

Males: adult
Mean

SD

Min

Mox

n

immature
Mean

so

Min.

Max

n

Females: adult
Mean

E

Min.

May

n

Moles: adull
Mean

sb

Min

Mox

n
subadult
Mean

5D

Min

May

n
immature
Mean

SD

Min

Mox

n

Females: adult
Meon

EN

Min

Max

n

205
C. r. rex
100 31 163 261 489 266 49 53
280 ]49 1068 102 119 087 038 530
96 28 142 222 465 252 42 43
i06 35 182 275 508 287 58 65
30 29 26 Ka 22 29 23 32
103 47 78 263 492 26,4 51 53
355 1052 3770 110 114 077 04 40
97 30 5| 240 479 253 44 48
109 585 155 277 512 283 58 60
18 15 10 15 9 14 14 7
102 54 58 269 484 265 52 50
am 194 261 065 072 073 O49 520
96 50 52 255 474 249 4,4 42
107 59 ó2 280 496 279 59 53
EA 20 15 20 v 19 20 4
100 56 57 260 495 270 5.4 49
307 259 297 077 094 127 048 $75
94 50 5| 247 472 247 Ai 38
107 D 61 275 509 304 64 58
28 28 23 28 23 28 28 10
C. r, coccineifrons
102 34 158 265 482 259 5.1 54
137 175 1056 067 157 060 028 331
100 32 136 249 465 251 az 51
105 37 170 zh5 502 273 59 57
14 14 12 14 4 13 14 7
104 58 ál 261 472 249 50 55
Ion 173 141 Den 156 025
102 E 59 255 233 47
106 a w 27] 264 53
4 4 4 4 ] 3 4 1
102 59 ei 269 25.6 5.1
071 O71 00 042 007
101 58 e) 253 5
102 59 60 259 DI
2 2 2 i} 2 2
C. r. similis
98 3l 150 260 476 248 49 53
239 146 607 103 150 3120 032 34
93 2H IA 240 449 226 EN Ab
102 34 "ei 280 504 276 56 Ki
36 a5 32 35 19 35 35 40
99 49 94 266 481 252 51 ál
153 740 4017 075 086 097 019 354
96 32 $4 249 462 235 47 58
102 55 154 277 498 272 54 63
16 14 a 16 n 15 14 2
100 54 59 262 472 248 50 54
252 219 223 073 065 O68 029 29
96 50 Zo 245 462 235 44 50
106 58 63 274 479 263 54 a
2i 21 17 21 10 20 20 13
97 55 58 260 484 253 54 52
208 205 209 Dô 175 129 058 374
93 491 529 250 444 226 44 ag
103 574 ai 278 511 277 E 57
23 73 a 24 19 23 22 5

206 MEMOIRS OF THE QUEENSLAND MUSEUM

C. r. erypforhynehus TABLE 34. Measurements (mm) and weights (g) of. specimens of
Males; adult Cicinnurus magnificus
Mean 100 32 155 265 475 253 52
sp 165 187 957 069 154 093 038
Min 9) W 19 252 254 20 47 Wing Tol Tall Torus Total Bil ` Bil Weight
Max 104 26 IW 23 <48,7 266 58 length length central length head length width
n w 0 7 100 4 à o length dee
subadult 7
Mean 102 44 106 264 480 247 53 COD
5D 071 1344 5364 044 0.14 DJd Males: adult
Min 101 34 ól 259 246 52 Mean là 39 269 319 54675 289 70 97
Max o — 53 181 268 248 -54 5D 251 205 2303 109 103 112 044 582
n 2 2 2 2 ! 2 2 Min m 35 21 289 532 262 62 B
immature Max 121 43 319 339 565 313 79 no
Mean 99 a 59 265 495 254 46 h o 3 o 8 12 25 5 Ü
n 1 1 | | l 1 1 subadull
Females: adult Mean M5 55 157 308 561 292 74
Mean 99 56 Si 258 486 259 56 5D 228 724 10947 059 Don 034
a 178 184 378 03 235 190 022 Min We) wat mob 219. $8
Min We NE REIST E ET mv Sa Max "m 5) 2 315 299 — 76
Max m — 5 o 22 52 23 59 å T. d 5 i 4 5
n å 5 5 å 3 6 6 WC Se
Mean 17 o aw 314 552 293 70 a
C. t gymnorhynehus sD 474 122 266 142 148 055 O48 4.84
Males; adult Min l4 am o 26 541 285 64 82
Mean 99 30 IW 25 4&] 246 32 Max 124 o e 335 562 299 76 o
5D 222 174 1307 074 137 071 DA a $a te. P Eius 5 A 6 2
Min 9 28 134 248 460 253 45 Ferier Kant
Mox 108 35 177 281 498 277 58 Mean na 6 o 304 542 293 70 — 79
n 235. 34 w 4 8 02 2 sO 274 279 411 149 095 128 039 6.26
subadull Min JD. 56 57 276 532 275 on 73
Mean 100 — 45 143 261 488 249 53 Max TW 84 o 33A 558 327 77 8
sD 071 283 2687 om 17] 127 049 å (à må M ab Kc «^q 7
Min 9 24 124 35] 475 26 49
Mox 100 — a8 162 28] 500 278 55 C. m, chrysoplenis
n 2 4$ 2 2 2 2 2 Males: adult
Femoles: adull Mean 16 39 294 314 557 297 70 102
Mean 95 53 MH 258 49 28 57 sp 188 217 1964 106 124 105 025 630
sD 158 302 158 063 172 203 042 Mii [i "um Ae uaa 515 97 Ae 920
Min % 80 5 254 480 250 6 Mox po 4 32) 30 ai 319 76 1190
Max MO 573 mm 269 514 303 42 5 bn xc o dp oi 7 c 2 3
n 5 5 F 6 3 5 5 Vrimatüra
- Mean "å o aw » % nm 7 o
oll subspecies sp 247 327 354 065 089 083 038 4667
Males: adult Min Im an 5 300 546 285 682 780
Mean 100 32 158 264 488 263 51 53 Max Im 6 72 322 564 322 79 1080
2 334 221 1049 09 168 153 041 483 e TG cig emer n9 VIS Dod OS
Min 93 28 1M 222 449 226 Ai 43 Forster dent
Max ho æ 182 288 540 312 61 65 Maan n2 6 62 32 554 294 71 a
n H e Aa dde GNSS 18 W åp A24 195 428 113 112 106 033 793
subadult Min w 8 5 279 539 124 64 BIO
Mean 102 å 48 9A 266 492 262 52 5 ha før. ett (ëmge eng Sove Saline
sp 3,9 954 3959 088 149 148 037 502 p mm dà wo 3 om D a
Min % 30 a 240 457 25 44 48
Max MO op 162 279 52 308 ai a =n beh
n 2 2 2 2 3 D 3 9 ` Moles eau
immature Mean "3 39 272 312 563 307 66 909
Mean 102 55 359 265 488 20 52 53 gp 242 183 1988 104 088 070 035 321
sp 320 316 270 088 139 127 044 3/9 mi 105 35 225 288 548 289 59 745
Min % D 52 229 462 233 44 420 Max 17 — 43 303 331 584 327 7a 105
Max 110 64 64 280 511 283 ol &00 n 38 38 32 35 24 37 38 32
n 58 8 S S8 35 5 5 18 about
Females: adult Mean i2 54 A 307 566 302 69 o
Maan 99 — 56 5 262 492 26 55 Å SD 515 Am 194 199 136 170 055
SD 296 285 244 077 143 157 047 598 Min 104 4? 58 283 551 249 Did
Min 93 291 Al 247 444 226 41 38 Max 120 58 t4. 35] 582 38 78
Max 107 62 03 281 5]4 30,4 59 58 n 8 a b 8 5 B 8 1
" T» 38 70 78 & V 78 6 mmaurå
Mean mn 59 o an 558 302 68 — B7
sp 299 273 298 087 086 088 114 Bad
Min NÉ — 54 97 301 538 298 05 72

Females: adult

BIOMETRICS OF THE BIRDS OF PARADISE

Mean 108 59 60 300 554 304 69 77
5D 209 235 315 105 089 077 033 879
Min 104 539 533 265 539 286 63 62
Max 115 64 66 320 S571 318 78 94
n 45 EI 31 45 22 45 4A 37
C. m. intermedius
Males: adult
Mean 115 38 258 320 561 308 7,1
sD 171 258 1702 O85 141 160 0,30
Min 112 35 220 302 545 287 65
Max 118 44 287 331 877 345 73
n 16 16 10 lė 4 15 14
subadult
Mean 119 55 31.5 30.4 72
SD 354 07 0,78 092 028
Min 116 54 30.9 297 70
Mox 121 55 32.0 310 74
n 2 2 2 2 2
immature
Mean 116 59 62 318 561 313 72
SD 240 205 206 089 130 068 0.38
Min 111 55 581 295 543 296 63
Max 120 63 65 331 581 321 78
n 18 18 17 18 8 18 17
Females: adult
Mean 109 59 59 297 55] 30.7 72
SD 242 260 288 087 061 087 032
Min 104 54 53 276 540 291 6.6
Max 116 63 64 310 56] 327 79
n 27 27 24 27 12 26 26
all subspecies
Moles: adult
Mean 115 39 275 315 558 300 69 97
SD 263 210 2317 107 121 131 041 868
Min 105 33 213 286 532 262 59 745
Max 121 4A 321 339 584 345 79 119
n 104 101 82 99 47 99 109 82
subadult
Mean 114 54 992 314 565 299 71 85
ER 450 486 80.31 1.51 123 142 O50 849
Min 104 42 58 283 551 26.9 60 79
Max 121 59 256 351 582 318 78 91
n 15 15 10 15 5 14 15 2
immature
Mean 114 60 63 317 558 302 6.8 92
SD 328 269 297 09 096 095 083 9.15
Min 105 54 57 295 538 285 D5 720
Max 124 69 72 337 58.1 32.2 79 1080
n 76 76 74 76 33 76 75 5
Females: adult
Mean 110 59 60 300 551 301 71 82
5D 3256 252 370 1.11 O96 109 035 10.48
Min 104 539 53 265 532 264 60 620
Max 125 66 å? 334 673 327 79 1130
n 108 107 B5 108 50 107 106 67

207

TABLE 35. Measurements (mm) and weights (g) of specimens of

Cicinnurus respublica

Wing

Tail

Tail

Tarsus

length length central length

Total Bill Bill
head length width

length length
Males: adult
Mean 98 39 140 27.1 490 254 4.1
5D 149 140 641 055 062 100 032
Min 94 35 121 259 480 233 34
Max 101 4l 150 28 502 277 50
n 28 28 22 27 13 27 27
subadult
Mean 100 48 81 274 506 265 43
5D 133 475 3822 118 297 127 0.34
Min 98 Al 53 246 485 250 a7
Max 102 53 146 285 527 287 4b
n g 9 B 8 2 8 7
immature
Mean 101 52 52 278 498 261 4.5
ER 134 255 152 106 066 080 007
Min 99 50 5] 269 490 251 44
Max 102 56 54 296 506 272 4.6
n 5 5 4 5 4 5 5
Females: adult
Mean 97 53 52 265 497 255 48
sp 250 229 224 105 1,18 142 021
Min 93 49 49 247 474 225 45
Max 101 58 59 287 516 276 51
n 16 16 14 15 9 14 13

Weight

208

TABLE 36. Measurements (mm) and weights (g) of specimens of

MEMOIRS OF THE QUEENSLAND MUSEUM

Paradisaga raggiana

Wing Tail Tal Torus Tota Bill Bil Welght
length: length central length heod ength width
fength length
P rraggiana
Males: adult
Mean 189 136 447 421 698 396 101 272
5D 353 464 453] 225 093 111 Déi 1647
Min 185 128 358 386 684 yo 93 238
Max 197 E) 43 456 709 414 Wa 285
n 9 9 9 8 7 9 9 H
subadull
Mean 190 139 291 422 69.00 3870 10.23 272
SD 61 61| 205.48 238 1123 105 021 1202
Min 186 132 M5 305 682 377 100 263
Max 193 14a 437 430 098 398 104 280
n 3 3 2 3 2 3 3 2
immature
Msan 186 132 126 436 69.7 404 96 233
sD 354 49 14) 071 078 120 085 10.4
Min 183 128 125431 691 395 90 2
Max 188 135 1287 44]. 702 412 102 240
2 2 2 2 2 2 2 2
Females: adull
Mean 164 121 114 383 669 365 93 184
D 6594 439 787 157 103 138 048 1782
Min 157 115 103 367 657 352 86 170
Max 172 125 25 406 à 682 383 102 215
n 5 5 5 5 4 4 5 5
P. t augustaevictoriae
Males: adull
Mean 184 RI 45) 419 664 375 112 276
50 535 3i4 3012 147 177 113 074 2407
Min WO 124 3 394 622 454 100 WA
Max 194 136 523 455 703 9398 30 300
n 37 37 E 37 EU 36 37 là
subadull
Mean 186 131 157 426 667 379 Iå
50 390 357 4314 130 136 085 077
Min 179127 19408 646 30 103
Mox 192 139 227 446 881 896 128
n H 9 9 9 6 9 9
immature
Mean 177 129 119 424 6862 376 108 223
SD 7W 636 766 140 1658 112 058 4099
Min 163 120106 393 0635 361 97 199
Max 191 146 137 45 684 2398 114 270
n 15 15 18 15 i5 15 15 3
Females: adull
Mean 161 n7 108 381 642 362 101 159
ED BOs 49) 524 254 135 088 049 2127
Min 180 108 100 339 626 348 92 13
Max T8) | 126 120 44 678 378 109 195
n 2 27 2å 26 ib 27 27 10
P. r. intermedia
Males; odult
Mean 185 132 448 424 682 3971 10,5 257
ER 402 385 3443 200 263 113 040 2082
Min 178 127 384 W 645 354 90 244
Max 19) 141 505 457 712 406 115 27
n 18 18 15 18 7 15 16 3
subadull
Mean ]B6 13) 325 418 690 397 104
SD 4d) — 329 125.82 38 O66 159 054
Min 180 127 130 396 684 375 27
Mux 199 (Dë 40 431 697 2422 014
n 5 5 5 5 å 5 5

immatura
Meon 175 128 121 430 644 386 102 225
sD 403 377 548 122 148 1,07
Min 192 124 1] 414 370 93
Mas 181 133 129 dai 396 11.5
n 4 å 4 4 1 3 3 1
Females: adult
Mean 162 121 114 376 653 373 9.7 192
ER &08 585 617 250 17) 149 072 4031
Min 148 Nå 14 354. 620 3565 Bó 163
Max 178 | 128 126 442 684 393 109 220
n 13 18 13 12 10 13 13 2
P. 1. salvadorii
Males: adull
Mean 188 135 463 430 699 389 98 266
50 447 553 3106 135 131 117 048 1525
Min 180 125 410 2388 670 365 90 240
Max 198 184 527 456 725 411 10.8 295
n 4] 42 38 42 23 42 42 21
subadull
Mean 186 133 284 435 690 388 98 249
sp 530 589 18612 152 098 109 053 2624
Min 178 Á 1722 15 403 626 2357 &8 210
Max 197 144 517 463 710 405 107 288
n x 20 17 19 13 20 18 9
immature
Mean 197 127 120 2427 691 387 97 25
sD 78| 607 719 199 181 140 0489 23454
Min ted 115 106 354 663 364 8.4 189
Max 190 136 131 451 716 421 112 255
n 23 23 22 22 1 23 23 8
Females: adull
Mean 164 120 nh 325 663 374 96 175
SD 642 515 666 139 1,48 104 046 1759
Min 180 110 jiu 34] 639 343 8,7 135
Max 182 132 128 403 698 404 104 210
"n ål 3! 29 30 20 31 3! 24
oll subspecies
Moles: adult
Meen 186 133 455 424 681 385 104 270
sD 490 483 3285 165 20 1-46. 088 1938
Min 160 124 358 386 622 354 90 234
Max 198 154 522 3457 725 414 130 300
n 105 — 106 98 105 68 102 47
subadult
Mean 186 133 256 429 684 387 10.3 2535
SD 442 53) 14688 1.60 145 125 088 2540
Min 178 122 115 305 ób 37 88 210
Mox 197 144 NP 463 710 422 128 288
n Ai 37 33 36 24 3? 35 H
immature
Mean 177 1285 120 428 675 384 101 225
SD 710 588 707 1469 226 142 087 2399
Min 180 115 1064 3584 635 361 Be 189
Mas 191 146 137 45] 716 421 115 270
n A4 44 43 43 29 43 4) 14
Females: adult
Mean 182 19 III 378 655 359 9.8 173
sn 726 527 644 205 173 157 O58 2079
Min 148 108 100 339 620 348 B6 133
Max 182 132 128 442 698 404 109 220
n 76 76 73 73 49 75 76 4)

BIOMETRICS OF THE BIRDS OF PARADISE

TABLE 37. Measurements (mm) and weights (g) of specimens of

Paradisaea apoda

Wing Tail Tail Tarsus Total Bill Bil Weight
length length central length head length width
length length
P. a.apoda
Males: adult
Mean 232 165 637 526 773 439 9.6
ES 461 511 10472 242 305 277 055
Min 225 156 500 450 715 377 81
Max 240 175 854 548 80 476 104
n 16 16 14 17 6 16 17
subadult
Mean 228 161 361 523 760 444 9.65
SD 525 533 20560 1.44 155 158 027
Min 219 152 150 496 747 426 9.2
Max 239 170 659 546 779 480 10,2
n 12 12 1 12 4 12 12
immature
Mean 209 151 146 512 759 440 9.6
SD 907 568 861 271 398 223 04
Min 198 141 132 450 725 414 89
Max 224 161 160 534 803 48 103
n 8 8 8 8 3 8 8
Females: adult
Mean 202 152 144 486 737 426 9.5
SD 673 576 723 377 236 184 045
Min 194 141 132 449 714 401 89
Max 215 158 153 532 777 451 102
n 8 8 8 8 7 8 8
P. a. novaeguinea
Males: adult
Mean 206 146 591 467 725 404 9.4
SD 305 395 5760 120 127 125 032
Min 200 140 480 446 700 376 90
Max 213 154 663 489 747 426 103
n 24 22 17 24 9 22 23
subadult
Mean 203 143 356 46.17 709 401 9.27
SD 4.44 3.66 217.40 1.02 164 099 040
Min 190 135 134 448 688 38.8 8.6
Max 210 148 662 489 733 430 101
n 27 26 19 26 7 24 26
immature
Mean 191 137 135 470 710 401! 93 224
SD 795 390 677 121 227 09 035 212
Min 180 130 125 448 655 38.1 87 22
Max 209 143 146 493 740 416 101 225
n 25 24 21 25 14 22 25 2
Females: adult
Mean 173 128 119 409 680 2390 89 172
SD 6.29 676 684 188 104 109 036 212
Min 159 118 105 388 666 37.1 82 170
Mox 188 142 132 458 705 407 9.6 173
n 26 26 25 26 15 25 25 2
all subspecies
Males: adult
Mean 217 154 612 491 744 419 9.5
SD 13.39 10.44 84.10 343 320 265 043
Min 200 140 480 446 700 376 8.1
Max 240 175 854 548 800 476 104
n 40 38 31 E 15 38 40
subadult
Mean 211 148 358 48.092 7276 41.51 9.39
SD 1250 956 2096 310 300 239 041
Min 190 135 134 448 688 388 8.6
Max 239 170 662 546 779 480 102
n 39 38 30 38 n 36 38

immature

Mean 196
SD 11.19
Min 180
Max 224
n 33
Females: adult
Mean 180
SD 13.92
Min 159
Mox 215
n 34

140 138 480
771 856 246
130 125 448
161 160 534
32 29 33
133 125 427
1223 1252 41
118 105 388
158 153 532
34 33 34

71.9
3.15
655
80.3

17

69.8
3.11
66.6
777

22

41.2 9.3
223 041
38.1 87
480 103

30 33
39.9 9.1
201 046
371 8.2
45] 102

33 33

209

TABLE 38. Measurements (mm) and weights (g) of specimens of

Paradisaea minor

Wing Tail Tail Tarsus Total Bill Bil Weight
length length central length head length width

length length

P. m. minor
Males: adult
Mean 188 127 503 447 695 390 105 239
SD 471 443 4876 165 219 170 045 2372
Min 180 116 420 394 653 340 92 185
Max 196 133 641 478 725 425 114 285
n 27 27 23 27 19 27 27 12
subadult
Mean 184 125 318 441 687 378 104
SD 562 457 15564 172 197 168 043
Min 172 115 108 395 649 345 97
Max 195 132 498 465 715 408 11.2
n 21 20 16 19 14 21 21
immature
Mean 179 123 116 438 683 38.1 103 216
SD 800 627 757 220 196 157 065 1991
Min 162 105 98 392 648 360 9.3 186
Max 194 130 127 469 727 414 121 242
n 22 22 18 21 21 22 22 7
Females: adult
Mean 161 112 103 397 658 370 100 158
SD 6.18 606 650 230 170 142 068 10.84
Min 152 103 93 371 628 347 90 145
Max 174 120 118 452 690 400 117 170
n 23 22 19 23 21 23 22 6

P. m. finschi
Males: adult
Mean 190 131 469 444 700 383 104 256
SD 547 427 2369 232 176 117 078 2909
Min 180 124 430 392 668 357 8.5 183
Max 199 141 537 492 738 406 119 300
n 27 26 24 26 22 27 26 21
subadult
Mean 190 130 357 446 688 382 105 230
SD 462 495 13306 1.14 151 109 0.76
Min 182 124 133 419 668 361 97
Max 201 141 481 46 712 403 118
n 15 15 14 15 13 15 15 ]
immature
Mean 179 124 121 439 678 373 98 205
SD 8.26 534 710 223 205 167 068 29.42
Min 165 110 108 406 628 322 8.7 151
Max 193 131 131 467 708 39.1 108 268
n 18 16 14 16 1 16 15 14
Females: adult
Mean 164 115 109 41.4 669 37.4 10.2 164
SD 645 473 857 297 178 131 058 18.46
Min 152 104 97 365 643 345 9.0 141
Max 175 123 129 474 713 407 114 210
n 26 26 24 25 22 26 26 16

210

MEMOIRS OF THE QUEENSLAND MUSEUM

P. m. jobiensis
Males: adult
Mean 200 137 495 482 710 394 108 298
SD 755 378 0264 158 209 131 052 404
Min 183 130 353 455 674 358 96 293
Max 210 144 S&A 404 736 2414 15 30
n 12 i2 9 11 7 10 12 3
subadull
Mean 196 135 349 483 712 398 105 240
SD 575 650 14688 209 200 148 1,00
Min 184 122 162 439 685 375 92
Max 203 144 482 50] 737 42 125
n 9 9 7 9 å 9 9 1
immature
Mean 189 131 122 473 712 397 109 230
EN 1219 | 453 640 197 126 077 O71 2928
Min 163 124 H4 430 693 388 101 210
Max 200 138 10 490 728 4807 122 250
n B B 8 8 7 B 8 2
Females: adult
Meon 172 12] 112 422 36.4 SA 15
sD 20.5)
Min 152
Max 189
n 1 1 | ! | ! 3
P. m. pulchra
Males: subadult
Mean 190 130 435 451 668 383 108
sD 465 403 2974 195 Dou 028
Min 184 124 203 437 369 105
Max 194 133 462 473 390 nl
n 4 4 3 3 1 å 4
immature
Mean 187 128 121 449 717 405 103
n 1 | I 1 1 1 d
Females: adult
Moon 160 1»1 101 391 88]! 392 102 186
E» 24 096 432 029 035 OF 052
Min 157 mW 9 389 678 384 94
Max 162 112 105 395 683 40 105
n 4 4 4 4 2 4 4 i
all subspecies
Males: odull
Mean 191 130 487 452 700 388 105 254
SD 691 549 4493 232 200 147 062 3013
Min 180 116 353 392 653 340 B5 183
Max 210 144 64) 494 728 425 119 30
n 66 65 56 64 A8 Ki 65 36
subadult
Mean 189 129 346 451 691 383 105 235
5D 670 602 13979 228 201 156 006 707
Min 72 118 108 395 649 345 92 230
Max 203 143 498 501 737 420 125 240
n Eu 48 E 46 34 ay 49 2
immature
Mean 181 125 119 444 687 381 102 210
SD 952 633 730 245 223 169 07) 2682
Min 162 — 105 98 392 628 322 87 151
Mox 200 138 181 490 728 24 122 268
n E 50 ad E) 42 EU 49 23
Females: adult
Mean 163 114 106 404 663 32 104 165
EN 636 549 796 267 183 144 061 1747
Min 152 103 93 365 628 345 90 14]
Max 1795 126 0329 474 713 407 017 210
n 56 55 Eu 55 AJ E? 55 26

TABLE 39. Measurements (mm) and weights (g) of specimens of

Puradisaea decora

Wig Toil Toi Totrsus Total Bål Bil Weight
length length central lengin head length widin
length lenath

Males; adult
Mean 181 137 43] 411 676 367 89 237
sD 262 347 4170 D% 213 103 047
Min 177 128 368 390 641 352 82
Max 188 143 536 425 709 400 98
n 22 22 16 21 7 ?0 20 1
subodult
Mean 180 139 290 41.4 669 368 89
D 232 197 9449 059 077 09% 08
Min 176 136 177 404 657 345 8.2
Max 155 142 448 423 681 383 99
n 17 17 ló 16 7 17 16
immature
Mean 172 138 120 (412 659 366 91
ES 511 379 273 074 104 023
Min 164 134 iló 402 35.1 Bä
Max 178 144 123 422 378 94
n 7 7 b 7 1 6 7
Fernales: adult
Mean 160 128 109 372 645 359 89
SD 256 282 227 079 040 3| O48
Min 157 123 108 364 oi 344 79
Max lé 131 u2 385 649 387 96
n H 8 8 8 3 8 B

TABLE 40. Measurements (mim) and weights (g) of. specimens of
Parudisaea rubra

Wing Tail Tai Tarsus Total Bil Bill Weight
length length central length head length width
length length

Males: adult
Mean 176 19 563 419 639 355 86 201
sp 372 272 6089 141 128 186 048 2562
Min 169 114 478 378 608 309 70 158
Max 186 125 765 437 657 392 97 224
n 27 27 23 26 18 26 27 7
subadult
Mean 174 121 243 417 631 340 &2 201
so 307 300 10747 079 115 100 038
Min 168 115 112 405 612 2320 75
Max 178 125 457 434 645 362 90
n 17 17 16 17 7 y 16 i
immature
Meon UO 119 17 423 634 352 B6 195
5D 443 419 522 128 156 152 049
Min 162 12 109 403 610 333 76 156
Max 178 126 127 447 658 38,4 92 212
n 10 10 9 10 7 10 10 8
Females; adult
Mean 159 114 108 388 631 346 85 158
5D 626 419 388 184 161 128 062 2683
Min 181 105 102 352 602 328 71 ns
Max 174 121 17 435 664 385 98 208

BIOMETRICS OF THE BIRDS OF PARADISE

TABLE 41. Measurements (mm) and weights (g) of specimens of

Puradisuea guilielmi

Wing

Tail

Tail

Tarsus

length length central length

Males: adull
Mean 97
sp 3.69
Min 172
Max 188
n 27
subadult
Mean 177
5D 248
Min 173
Max 182
n 10
immature
Mean 172
SD 64)
Min 163
Max 179
4
Females: adult
Mean 158
E 5,82
Min 181
Max 171
n 15

lengih

75,41

Bill Bil Weight
length width

40.9 B9 256

165 043 674

366 Bl 250

433 93 265

41.8 93 250
LI7 063
40.4 86
435 107
10 y 1
43.3 95 219
161 028 438
423 93 188
457 99 280
4 4 2
39.9 9.0
109 052
376 50
42 100
15 15

TABLE 42. Measurements (mm) and weights (g) of specimens of

Paradisaeu rudolphi

Wing

Males: adult

Mean 157
SD 272
Min 153
Mox 163
n 2
subadull

Mean 160
5D 4,24
Min 157
Max 163
n 2
immature

Mean 155
sD 3.30
Min 180
Mas 157
n å
Females: adult
Mean 150
50 571
Min 142
Max 158

Tail Tol Tarsus
length length central length
length

P. t. rudolphi
83 409 401
406 2419 130
76 34) 365
91 452 418
?0 y 19
87 102 39
454 O71 099
84 10] 383
B9 102 397
2 2 2
93 96 400
316 450 114
90 9| 388
97 102 42
4 a 4
95 95 384
424 346 168
Kal 90 355
122 101 410
18 14 15

Bill Bil Weight
length width
41.6 Bé 177
156 057 212
39.4 76 175
45.6 26 178
19 20 2
423 8.45
071 049
418 81
428 88
2 2
41.3 &4 185
041 019
40.9 ER
418 87
4 4 U
40.8 86 157
118 047
38.4 80
429 97
14 15 t

211
P. r. ompla
Males: adult
Mean 154 B2 409 381 684 397 B2 178
sD 327 321 1204 170 088 068 039 556
Min 151 78 | 38 361 675 390 76 174
Max 159 B6 422 40.1 697 404 85 189
n 5 5 3 5 5 5 5 6
immature
Mean 152 93 94 384 687 403 84 153
SD 515 308 266 155 118 085 O46 1054
Min 142 Ka 9 357 669 392 77 14
Max 157 b 98 406 +70 418 KN 174
n 7 7 å 7 6 7 H 4
Female
Mean 147 91 94 363 653 380 8.2
n 1 1 1 i 1 1 1
P. r. margaritae
Males: adult
Mean 156 78 451 378 694 413 8.8 158
sD. 295 187 1270 082 1.6) 135 0854
Min 151 76 436 366 680 393 81
Max 158 81 458 389 710 423 95
n 5 5 3 5 3 4 E] 1
immature
Mean 156 98 97 391 692 401 8.4 169
SD 327] 239 182 l4 170 142 039 424
Min 151 95 95 377 680 386 79 166
Max 161 101 100 404 704 42 89 172
n 5 5 5 4 2 5 5 2
Females: adult
Mean 148 89 92 374 681 403 8.8 152
ES 522 270 344 138 149 124 029 2397
Min 143 85 88 357 66 384 85 124
Max 159 94 96 395 70 | 419 92 164
n 7 7 å 7 H H 7 3
all subspecies
Males; adult
Mean 157 82 A4 394 701 412 Bé 176
SD 306 395 2567 165 162 156 05 799
Min 151 76 34 361 675 390 75 158
Max 163 9] 458 418 738 456 96 187
n 30 30 23 29 24 28 30 9
subadult
Mean 160 87 102 39 709 423 LÉI
SD 424 354 071 099 078 071 0.4
Min 157 B4 101 383 703 418 8.1
Mox 163 89 102 397 714 428 88
n 2 2 2 2 2 2 2
immature
Mean 154 95 96 390 69.1 40.5 BA 165
SD 436 352 331 141 120 104 034 81
Min 142 90 90 357 669 386 7.7 149
Max 161 101 12 412 704 420 9] 174
n 16 16 15 15 10 16 16 7
Females; adult
Mean 149 93 94 380 68.7 405 8.7 153
sD §39 452 364 163 180 129 043 1975
Min 142 86 88 355 653 380 80 124
Mox 159 102 101 410 720 429 97 166
n 23 23 2) 23 20 22 23 4

212

NEW DISTRIBUTION AND HABITAT DATA FOR
THE PYGOPODID, PARADELMA ORIENTALIS
(GÜNTHER, 1876). Memoirs of the Queensland Museum
42(1): 212. 1997:- Four specimens of Paradelma orientalis
(QM transparency NR221-3) collected recently (4 Sept.,
1996) in Eena State Forest (28*19'S 150*50' E) 20km WNW
of Inglewood, SCQ extend the knowledge of the distribution
and habitat preferences of this species. This locality is approx-
imately 1 10km south of Dunmore State Forest, near Cecil
Plains, SCQ (26°24’S, 151?01' E), the former southernmost
known collection locality for the species (S. Wilson, QM,
pers. comm.). The specimens were unremarkable morpholog-
ically, closely resembling the photographs in Cogger (1996)
and Wilson & Knowles (1988). Snout-vent lengths were 154,
155, 156 and 172mm.

In Eena State Forest, three specimens were collected in
Eucalyptus crebra and E. microcarpa open forest with a dense
subcanopy dominated by Callitris columellaris and Al-
locasuarina luehmannii on loose sandy clay substrate at
28?19'S 150?50'E. The site had been silviculturally treated
on three occasions between 1937 and 1973 and sections
logged in 1971 (Inglewood DPI Forestry records). The re-
maining specimen was found in A. luehamannii closed forest
with widely scattered E. crebra emergents on a similar sub-
strate at 28?21'S 150°50’E, This site had been silviculturally
treated in the 1930s and 1950s and partially logged in 1984
(Inglewood DPI Forestry records). In both sites, all individu-
als were found sheltering under thick slabs of E. crebra bark
that had fallen from standing ringbarked trees.

P. orientalis is one of several species of reptiles from
Queensland's Brigalow Biogeographic Region with special
conservation ‘rating’, It is treated as ‘vulnerable’ by McDon-
ald et al., 1991; Cogger et al., 1993 and Queensland Nature
Conservation Regulation 1994. Little is known about the
species' biology and habitat preferences (Kluge, 1974;
Jenkins, 1979; Shea, 1987; Coggeretal., 1993; Cogger, 1996).
Previous published records suggested the species was con-
fined to remnant Acacia harpophylla forest and eucalypt
woodland with an understorey of A. harpophylla or sparse
tussock grass ground vegetation on grey cracking soils (e.g.,
Shea, 1987; Cogger et al., 1993).

This range extension combined with the knowledge that P.
orientalis appears to adapt to several soil and vegetation types
indicate that the species is more widespread than supposed.
Further, these records combined with recent observations in
Acacia falciformis woodland on Boyne Island (23*50'S
150?24' E) (M. Fitzgerald, pers. comm.); on a sandstone rise
in dry sclerophyll forest in Dunmore State Forest (S. Wilson,
pers. comm.); in Corymbia maculata and E. crebra dominated
open forest in Barakula State Forest (26?17'S 150%52'E) (D.

MEMOIRS OF THE QUEENSLAND MUSEUM

Hannah, pers. comm.); and in mixed species open woodland
with Triodia mitchelli dominated ground layer in the Chester-
ton Range, Charleville district (26?09'S 147?14'E) (C. Doll-
ery, pers. comm.) suggest it may be more abundant than the
published records indicate.

Whether the conservation status of P. orientalis should be
modified requires further investigation.

Acknowledgements

Thanks are due to Maritza de Oliveira for assisting with
fieldwork; Dave Hannah for providing details on the Barakula
record; Col Dollery for providing details on the Chesterton
Range records; Steve Wilson for providing details on his
southernmost localities for the species; and Mark Fitzgerald
for providing valuable comments about the species and taking
photographs that are lodged with the QM. Jeanette Cov-
acevich, Mark Fitzgerald and Geoff Smith commented on
early drafts of this note. The work was conducted as part of
the Dept of Natural Resources research into the distribution
and habitat requirements of fauna occurring in production
forests in Queensland.

Literature Cited

COGGER, H.G, 1996, ‘Reptiles and Amphibians in
Australia’. Rev, Ed. (Reed: Sydney). 796 pp.

COGGER, H.G., CAMERON, E., SADLIER, R. & EGGLER, P.
1993. "The Action Plan for Australian Reptiles'. (Aust.
Nature Conserv. Agency: Canberra). 254 pp.

JENKINS, R.W.G. 1979. The status of endangered Australian
reptiles. Pp. 169-176. In Tyler, M.J. (ed) ‘The status of
endangered Australasian wildlife’. (R. Zool. Soc. S.
Aust.: Adelaide). 210 pp.

KLUGE, A.G. 1974. A taxonomic revision of the lizard
family Pygopodidae. Miscellaneous Publications of the
Museum of Zoology, University of Michigan 147: 1-
221.

McDONALD, K.R., COVACEVICH, J.A., INGRAM, G.J. &
COUPER, P.J. 1991. The status of frogs and reptiles.
Pp. 338-345. In Ingram, G.J. & Raven, R.J. (eds) ‘An
atlas of Queensland's frogs, reptiles, birds and
mammals'. (Queensland Museum: Brisbane). 391 pp.

SHEA, G.M. 1987. Notes on the biology of Paradelma ori-
entalis. Herpetofauna (1) 17: 5-6.

WILSON, S.K. & KNOWLES, D.G. 1988. Australia's rep-
tiles; a photographic reference to the terrestrial reptiles
of Australia. (Collins: Sydney). 447 pp.

M. Schulz & T.J. Eyre, Forest Wildlife, Resource Sciences

Centre, Department of Natural Resources, P.O. Box 631,

Indooroopilly, Queensland 4068, Australia; 22 January 1997.

NOTES ON THE GENUS THORELLIOLA STRAND, 1942 (ARANEAE: SALTICIDAE)

JOANNA GARDZINSKA AND BARBARA PATOLETA

Gardzinska, J. & Patoleta, B. 1997 06 30: Notes on the genus Thorelliola Strand, 1942
(Araneae: Salticidae). Memoirs of the Queensland Museum 42(1): 213-222. Brisbane. ISSN

0079-8835.

Genus Thorelliola Thorell, 1887 is studied. Five species are diagnosed, described and
ilustrated. Four of them, T. biapophysis, T. glabra, T. javaensis, T. truncilonga, are described
as the new. Remarks on relationships and distribution are presented. [ ] Thorelliola, Indone-

sia, Papua New Guinea, Tahiti.

Gardzinska, J. & Patoleta, B., Zaklad Zoologii WSR-P, Prusa 12, 08-110 Siedlce, Poland;

18 February 1997.

Genus Thorelliola was established by Strand
(1942) for Thorellia ensifera (Thorell). The type
species was originally described as Plexippus ensi-
fer Thorell, 1877 and then transferred into
Thorellia by Keyserling (1882). The latter ge-
neric name was, however, preoccupied, thus
Thorelliola is the present valid name. Simon
(1901) included the genus (Thorellia at that time)
into a group Spilargeae — together with Eupalia
and Spilargis. Petrunkevich (1928) added six
other genera, none of them related. According to
Berry, Beatty & Pröszyfiski (in prep.), body shape
and genitalia structures of Thorelliola resemble
Euophrys and related genera.

Males of Thorelliola are very distinctive for
their ‘horns’ arising either from circular sockets
on the clypeus or from a tip of a robust truncus.
Such strange and extravagant structures are also
found in genera Padilla Simon, Allococalodes
Wanless and Cocalodes Pocock. According to
Wanless & Lubin (1986) the structures are prob-
ably used during intra-specific 3-3 interactions.
The same authors suggest that the horns in
Thorelliola have evolved from the lower of two
setae, that form part of a triad of stiff setae on the
clypeus in many salticids. The function of the
horns in Thorelliola seems still unknown. Since
there are no pores on any part of them, they cannot
be used for dispersing pheromones, moreover,
there is no evidence of any spider being injured
by horns of another spider (Jackson & White-
house, 1982), so these structures cannot be the
weapons. Zabka (1988) suggests that the protrud-
ing spines may work as stridulatory organs to-
gether with the corresponding palpal spines.

MATERIAL AND METHODS

Material from Indonesia and Tahiti, from Dr
C.L. Deeleman-Reinhold's collection and speci-

mens collected by D.J. Court from Papua New
Guinea are deposited in the Queensland Museum,
Brisbane (QM). Comparative material from Sin-
gapore and Sulawesi (Indonesia) was borrowed
from the Naturhistoriska Riksmuseet, Stockholm
(NRS). Measurements are given in millimetres.
The drawings were made using a grid system.
Dissected epigynes were digested in lactic acid.
Abbreviations used are: AEW = anterior eye row
width, ag — accessory gland, AL = abdominal
length, CH = cephalothorax height, CL = cepha-
lothorax length, CW = cephalothorax width, e =
embolus, EFL = eye field length, fd = fertilisation
duct, h = horn, id = insemination duct, la = lateral
appendix, PEW = posterior eye row width, pta =
prolateral tibial apophysis, rta = retrolateral tibial
apophysis, s = spermatheca, sd = seminal duct,
t = truncus.

Order ARANEAE
Family SALTICIDAE

Thorelliola Strand, 1942

Thorellia Keyserling, 1882: 1353.

Thorelliola Strand, 1942: 400; Pröszyfiski, 1984: 148;
Zabka, 1988: 470-471; Jackson & Whitehouse,
1989: 1-16; Koh, 1989: 118.

DIAGNOSIS. Small spiders (3.5-5.0) with rela-
tively high cephalothorax. Abdomen nearly
round; legs rather robust and long (IV the long-
est). 3 d. with horns or a truncus on the clypeus,
in 9 ? two smaller and more delicate setae.

DESCRIPTION. Cephalothorax longer than
wide. Clypeus low. Cephalothorax usually yel-
low-beige to brown, posteriorly darker. Abdomen
oval, light with grey-black pattern, covered with
shiny scales. Chelicerae with two prolateral teeth
and one retrolateral fissidentate tooth, with mul-

214

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Thorelliola ensifera, 3 3. A-E, from Bali. F-J, comparative material from Singapore. A-B, F-G, general
views. C, H, cheliceral teeth. D-E, I-J, palpal organs (lateral and ventral views).

tiple cusps. External margins of d chelicerae
with a lateral cusp. Maxillae and labium not dis-
tinctive, sternum heart-shaped. Legs relatively
robust and long, yellow-brown with darker mark-
ings, especially near the joints. Leg formula: IV,
III, I, II. Palpal organ simple, embolus counter-
clockwise, weakly sclerotized. Epigyne, trans-
verse oval, spermathecae more or less spherical;
insemination ducts rather short, without loops.

DISTRIBUTION. Malaysia, Indonesia, Polyne-
sia, Hawaii, Tahiti, Melanesia, Singapore, Papua
New Guinea.

Thorelliola ensifera (Thorell, 1877)
(Figs 1-3)

Plexippus ensifer Thorell, 1877: 606.

Thorellia ensifera Keyserling, 1882: 1353.

Thorelliola ensifera Strand, 1942: 400; Pröszyfiski,
1984: 148; Wanless & Lubin, 1986: 1211, 1214;
Zabka, 1988: 470-471; Jackson & Whitehouse,
1989: 1-16; Koh, 1989: 118.

MATERIAL EXAMINED. QMS35296, 3, Bali: Sanur,
5.08.1992., C.L. Deeleman-Reinhold; QMS35297, 9,
North Sulawesi: Dumoga Bone National Park, near
Doloduo, 27-30.07.1982; QMS35298, 9 , Tahiti, 1000m,
15.02.1985, C.L. Deeleman-Reinhold. COMPARATIVE

NOTES ON THORELLIOLA 215

0.25

FIG. 2. Thorelliola ensifera, 9 9. A-C, from Sulawesi. D-F, from Tahiti. G-I, comparative material from
Sulawesi. J-K, comparative material from Singapore. A-B, D-E, G-H, J, general views. C, F, I, K, cheliceral

teeth.

MATERIAL, NRS267/1806b, d, 9, Singapore
(Workman ded.), Thorell; NRS267/1806a, 9, Sulawesi
(Doria ded.), Thorell.

COMPARISON. T. ensifera differs from the other
species by the presence of two central horns

protruding on the clypeus, and by the shape of
embolus and spermathecae, Abdomen with three
white posterior spots.

DESCRIPTION. Male. Cephalothorax (Fig. 1A-
B) moderately tall, yellow-brown, covered with

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. Thorelliola ensifera, epigynes and its internal structures. A-B, from Sulawesi; C-D, from Tahiti; E-F,

comparative material from Sulawesi.

dark hairs. Surroundings of eyes black with some
dark, long and stiff hairs. Abdomen (Fig. 1A)
oval, shiny and yellow, with pattern of black and
gray markings and three whitish spots in front of
spinnerets, Anterior part of abdomen with scat-
tered protruding, long hairs. Spinnerets light yel-
low, with fine, dark hairs. Clypeus (Fig. 1B) low,
pale brown with two large, distinctive setae

(horns) in the centre. Chelicerae (Fig. 1C), max-
illae and labium yellow-beige. Sternum and
coxae pale yellow, Legs yellow-grey, covered
with numerous dark hairs and some spines. Ven-
ter light yellow with some black markings, espe-
cially on the lateral parts.

Palpal organ (Fig. I D-E). Embolus thinner than
in other species, seminal duct meandering. Retro-

NOTES ON THORELLIOLA 217

FIG. 4. Thorelliola biapophysis sp. nov., d d. A-C,
from Ambon; D-F, from Banda Islands. A-B, D-E
general views. C, cheliceral teeth.

+

lateral tibial apophysis rather slender, not furced,
Prolateral tibial apophysis with robust spine.
Dimensions. CL 2.28, CW 1.80, CH 1.44, AL
2.16, AW 1.80, AEW 1.56, PEW 1.49, EFL 1.02.
Female. Cephalothorax (Fig. 2A-B, C-D) mod-
erately tall, orange-brown, darker posteriorly,
covered with fine, whitish hairs. Eyes surround-
ings black with fringes of fine, white hairs and
long, brown bristles, especially above anterior
eyes. Abdomen oval, shiny, anteriorly whitish-
beige, posteriorly black-grey. Pattern of markings
shows some variability (Fig. 2A, D), but three
white spots on the posterior dorsal abdomen are
always conspicuous. Spinnerets thin, whitish-
grey, with paler tips. Clypeus (Fig. 2B, E) low,

FIG, 5. Tharelliala biapaphysis sp. nov., palpal organs,
lateral and ventral views. A-B, from Ambon; C-D,
from Banda Islands,

beige to brown, with two protruding setae —
more delicate than in the 3, Chelicerae (Fig. 2C,
F) orange-brown, without lateral cusps. Maxillae
and labium orange with paler tips. Sternum yel-
low, heart-shaped. Legs yellow-orange, darker
distally, with some black markings. Venter whit-
ish with å few black or whitish dots. Palps yellow.
Epigyne (Fig. 3A-D) transverse oval, in compar-
ison to the other species spermathecae more
spherical.

Dimensions. CL 1.92, CW 1.53, CH 1.11, AL
2.16, AW 2.28, AEW 1.56, PEW 1.44, EFL 1.04.

DISTRIBUTION, Malaysia, Indonesia, Polyne-
sia, Melanesia, Hawaii, Tahiti, Singapore.

218

FIG. 6. Thorelliola glabra sp. nov., & from Banda
Islands. A-B, cephalothorax, dorsal and lateral views
(horns on the tip of truncus reconstructed). C, chelic-
eral teeth. D-E, palpal organ, lateral and ventral views
(pta with the reconstructured spine).

Thorelliola biapophysis
sp. nov. (Figs 4-5)

MATERIAL EXAMINED. HOLOTYPE. QMS35299,
d, Ambon: Hila, forest, from bark with ants, 29-
31.01.95, C.L. Deeleman-Reinhold. PARATYPE.
QMS35300, d, Banda Islands: Lonthoir, Namulu,
21.01.1995, C.L. Deeleman-Reinhold.

DIAGNOSIS. d with the trunk on the clypeus;
in comparison to T. glabra. and T. truncilonga trun-
cus of the middle length. Posterior part of abdo-
men dark with one white spot in front of
spinnerets.

DESCRIPTION. Male. Cephalothorax (Fig. 4A-
B, D, F) moderately tall, brown-beige to brown,
with scattered brown hairs. Surroundings of eyes
black with fine whitish or brown hairs. Abdomen
(Fig. 4A, E) nearly oval, covered with reflecting
scales, yellow-whitish-black, with pattern of
some dark spots and markings. One whitish spot
in front of yellow-grey spinnerets. Clypeus low,

MEMOIRS OF THE QUEENSLAND MUSEUM

brown-grey to brown with trunk ended by two
protruding horns. Chelicerae (Fig. 4C), maxillae
and labium brown. Sternum heart-shaped, yel-
low, covered with fine, brown hairs. Legs yellow-
brown, tarsi and metatarsi paler. Coxae I and II
darker than the others. Venter pale grey.

Palpal organ (Fig. 5A-D). Embolus rather short
and robust; retrolateral tibial apophysis wide, api-
cally forked; two protruding setae near the retro-
lateral tibial apophysis. Prolateral tibial
apophysis slightly dislocated dorso-laterally.

Dimensions. CL 1.85, CW 1.35, CH 1.06, AL
1.45, AW 1.20, AEW 1.44, PEW 1.27, EFL 0.98.

Female unknown.

DISTRIBUTION. Ambon, Banda Islands.

ETYMOLOGY. From the structure of retrolateral tib-
ial apophysis, which is divided apically.

Thorelliola glabra sp. nov.
(Fig. 6)

MATERIAL EXAMINED. HOLOTYPE. QMS35301,
d, Banda Islands, Moluccas, Neira, 02.1995, C.L.
Deeleman-Reinhold.

COMPARISON. Differs from T. biapophysis by
the course of seminal duct, trunk shorter.

DESCRIPTION. Male. Cephalothorax (Fig. 6A-
B) tall, brown; scarce hairs only around the ante-
rior eyes; eyes surroundings black. Abdomen
missing. Clypeus low, brown; trunk rather
short. Chelicerae (Fig. 6C), maxillae and la-
bium brown.

Palpal organ (Fig. 6D-E). The course of seminal
duct similar to the upturned figure 3.

Dimensions. CL 1.80, CW 1.38, CH 1.08, AEW
1.40, PEW 1.25, EFL 0.96.

Female unknown.

DISTRIBUTION. Banda Islands.

ETYMOLOGY. From the Latin word glabrus =
smooth.

Thorelliola javaensis sp. nov.
(Figs 7-8)

MATERIAL EXAMINED. HOLOTYPE. QMS35302,
3, Java: Cibodas National Park, 1450m, from leaves,
10.12.1986, Suharto Djojosudharmo. ALLOTYPE.
QMS35303, 9, same data. PARATYPES.
QMS35304, 3,7 9, same data.

DIAGNOSIS. Dark brown stripes on the cepha-
lothorax. Embolus in palpal organ (Fig. 7F-G)
longer and thin, seminal duct not meandering; no

NOTES ON THORELLIOLA

FIG. 7. Thorelliola javaensis sp. nov., å from Java. A-C, general views. D,
truncus. E, cheliceral teeth. F-G, palpal organ, ventral and lateral views,

bristles near the retrolateral tibial apophysis, the
latter is slender and not furced apically. Sper-

mathecae of different shape (Fig. 8E-F). ducts (Fig. 8F),

nerets dark grey, with lighter
tips. Clypeus (Fig. 7B) very
low, brown, with the trunk (Fig.
7D) ended by two long horns,
Chelicerae (Fig. 7E) brown;
maxillae and labium orange
with paler tips. Sternum heart-
shaped with brown margins
and covered with brown, short
hairs. Legs I brown, darker and
more robust than the others;
tarsi yellow. Venter grey, with
no spots.

Palpal organ (Fig. 7F-G).
Embolus rather thin; seminal
duct non-meandering; retro-
lateral tibial apophysis rather
slender and slightly bended
apically; prolateral tibial
apophysis short, with the ro-
bust spine.

Dimensions. CL 2.04, CW
1.68, CH 1.32, AL 2.28, AW
1.44, AEW 1.68, PEW 1.50,
EFL 1.08.

Female. Cephalothorax (Fig,
8A-C) tall, yellow-beige, with
two distinctive, brown stripes,
generally paler than that in the
3. Surroundings of eyes dark
brown. Eye field covered with
the fine, white hairs. Abdomen
near oval, of the similar colour
pattern as in d, but å little
darker. Spinnerets yellow-
grey. Clypeus (Fig. 8C) rather
low, orange, with two protrud-
ing central setae. Chelicerae
(Fig. 8D), maxillae and labium
orange. Sternum pale yellow,
covered with fine, brown,
hairs. Legs yellow, with dark
grey markings, especially near
the joints. Palps yellow-beige.
Venter pale grey.

Epigyne (Fig. 8E-F), Sper-
mathecae less spherical than in

other species, with different shape of fertilization

Dimensions. CL 1.94, CW 1.45, CH 1.14, AL
DESCRIPTION. Male. Cephalothorax (Fig. 7À- 2.33, AW 1.78, AEW 1.51, PEW 1.36, EFL 0.90.

C) tall, brown-beige in the eye field and yellow

around fovea, with two thoracic stripes. Sur- DISTRIBUTION. Java: Cibodas N.P.

roundings of eyes black. Abdomen (Fig. 7A)

grey-beige with some darker spots and dots. Spin- © ETYMOLOGY. From the locality,

220

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Thorelliola javaensis sp. nov., Ẹ from Java. A-C, general views. D, cheliceral teeth. E-F, epigyne and

its internal structures.

Thorelliola truncilonga sp. nov.
(Figs 9-10)

MATERIAL EXAMINED. HOLOTYPE. QMS29963,
d, Papua New Guinea, Central Province, National
Capital District, House Pomims, (possibly escaped
from Yule Islands' litter), left palp missing,
20.08.1985, D.J. Court. ALLOTYPE. QMS29964, 9,
Port Moresby, Botanic Garden, on foliage, 28.06.1988,
D.J. Court.

DIAGNOSIS. Male truncus long and robust,
ended by two short setae. Prolateral tibial apoph-
ysis short and with no spines. Shape of sper-
mathecae and course of insemination ducts
different than in other species.

DESCRIPTION. Male. Cephalothorax (Fig. 9A-
B) tall, orange-brown, darker posteriorly. Eyes
surroundings black, covered with brown hairs.
Abdomen oval, orange-black, with a pattern of
dark brown and grey spots. One white spot in
front of the spinnerets. Spinnerets yellow, with
brown markings laterally, Clypeus (Fig. 9B)
rather low, brown, with sparse, brown bristles and
a massive trunk ended by two protruding, but
short horns. Chelicerae (Fig. 9C), maxillae and
labium brown with lighter tips. Sternum broad
and yellow. Legs brown-orange, covered with
brown hairs, more numerous distally. Venter yel-
low-grey with some small greyish lateral spots.
Palpal organ (Fig. 9D-F). Embolus more robust
than in the other species; prolateral tibial apoph-

NOTES ON THORELLIOLA 221

FIG. 9. Thorelliola truncilonga sp. nov., d from Papua New Guinea. A-B, general views. C, cheliceral teeth.
D-F, palpal organ, ventral, dorsal and lateral views.

ra
r2
ra

ysis short, with no spines; near
the retrolateral tibjal apophysis
— an appendix with two long
spines.

Dimensions. CL 2.40, CW
2.10, CH 1.45 AL 2.05, AW 1.70,
AEW 1.80, PEW 1.70, EFL 1,02.

Female. Cephalothorax (Fig.
10A-B) tall, yellow-beige with
dark brown markings. Sur-
roundings of eyes black, cov-
ered with sparse, stiff brown
hairs. Abdomen (Fig. 10A)
oval, yellow-beige, with the
pattern of grey and black trian-
gle-like markings and with
scattered brown hairs. Spinner-
ets yellow-grey. Clypeus pale
yellow with sparse amber bris-
tles. Chelicerae (Fig, LOC) yel- ,
low orange. Maxillae and labium p |
similar in colour, with darker 4
tips. Stemum pale yellow. Legs
yellow with brown markings,
especially around joints. Ven-
ter pale yellow, with small dark
brown spots laterally.

Epigyne (Fig. 10D-E).

Dimensions. CL 1.80, CW
1.40, CH 1.16, AL 1.60, AW 1.45,
PEW 1.25, AEW 1.35, EFL 0.90.

DISTRIBUTION. Papua New Guinea: Port
Moresby.

ETYMOLOGY. From the structure of the trunk.

ACKNOLEWLEDGEMENTS

We wish to thank Dr C. Deeleman-Reinhold
(Ossendrecht) and Mr D.J. Court (Singapore) for
providing specimens to this study. We are also
grateful to Dr M. Zabka and Prof. Prószynski for
critical review ol the typescript.

LITERATURE CITED

BERRY, J., BEATTY. J. & PRÓSZYNSKI, J. (submit-
ted), Salticidae (Araneae) of the Pacific Islands.

JACKSON, R.R. & WHITEHOUSE, M.E.A. 1989.
Display and mating behaviour of Thorellia en-
sifera à jumping spider (Araneae: Salticidae) from
Singapore. New Zealand Journal of Zoology 16:
1-16.

KEYSERLING, E 1882. Die Arachniden Australiens,
nach der Natur beschrieben und abgebildet.
Nürnberg 2: 1325-1420.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 10. Thorelliola truncilonga sp. nov., 9 from Papua New Guinea. A-B,
genera] views. C, cheliceral teeth. D-E, epigyne and its internal structures.

KOH, J.K.H. 1989. A Guide to common Singapore
spiders. Singapore 1-160.

PETRUNKEVITCH, A. 1928. Systema Aranearum,
Transactions of the Connecticut Academy of Ans

. and Sciences 29: 1-270.

PROSZYNSKI, J. 1984. Diagnostic drawings of less
known Salticidae (Araneae) — an atlas. Zeszyty
Naukowe Wyzszej Szkoly Rolniczo-
Pedagogicznej w Siedlcach |: 1-177.

THORELL, T. 1877. Studi sui Ragni Malesi et Papuani.
I. Ragni di Selebes raccolti nel 1874 dal Dott. O.
Beccari. Annales del Museo Civico di Storia Na-
turale di Genova, 10: 341-634.

SIMON, E. 1901. Histoire Naturelle des Araignées.
Paris, Roret; Libraire Encyclopedique. 2 (3): 762-

67

WANLESS, ER. & LUBIN, Y.D. 1986. Diolenius
minotaurus sp. nov., a remarkable horned jumping
spider from Papua New Guinea (Araneae:
Salticidae). Journal of Natural History, 20; 1211-

1220.

ZABKA, M. 1988. Salticidae (Araneae) of oriental,
Australian and Pacific Regions, Ill. Annales
zoologici 41: 421-479.

1991, Studium taksonomiczno-zoogeograficzne
nad Salticidae (Arachnida; Araneae) Australii,
Zeszyty Naukowe Wyzszej Szkoly Rolniczo-
Pedagogicznej w Siedicach. Rozprana naukowa 32.

CRETACEOUS FRESHWATER BIVALVES FROM QUEENSLAND

SCOTT A. HOCKNULL

Hocknull, Scott A. 1997 06 30: Cretaceous freshwater bivalves from Queensland. Memoirs
of the Queensland Museum. 42(1): 223-226. Brisbane. ISSN 0079-8835.

Three new freshwater unioids are described from the Griman Creek and Winton Formations.
Protovirgus wintonensis sp. nov. is a medium-sized elongate unioid with the anterior
adductor muscles raised on a platform. Prohyria macmichaeli sp. nov. is charactertised by
its more anteriorly placed umbones and its less robust form in comparison to other members
ofthe genus. Velesunio goondiwindiensis sp. nov. is small, elongate-ovoid with finer growth
lines than most members of the genus. All three taxa range from middle Albian to Cenoman-
ian. [ ] Unionida, Prohyria, Protovirgus, Velesunio, Winton Fm., Griman Creek Fm.

Scott A. Hocknull, Queensland Museum PO Box 3300 South Brisbane, Queensland Australia

4101; 2 January 1997.

Collections made from the middle Cretaceous
Winton and early Cretaceous Griman Creek For-
mations have yielded three new taxa; Protovirgus
wintonensis sp. nov., Prohyria macmichaeli sp.
nov. and Velesunio goondiwindiensis sp. nov. The
Griman Creek Fm. (Exon & Senior, 1976) is a
non-marine to brackish unit which outcrops in the
Surat Basin. It is considered to be of middle
Albian age by Burger (1986, 1995) who placed
the unit within the Coptospora paradoxa Zone.
The younger Winton Fm. out-crops extensively
within the central and southwestern Eromanga
Basin being of latest Albian to Cenomanian age
and is, except for its extremities, a bare and ex-
clusively freshwater deposit, comprising lacus-
trine and fluviatile siliciclastic sediments.
McLoughlin et al. (1995) described the Cen-
omaniam flora from the Winton Fm. Localities are
prefixed QML and are detailed in the appendix.

Newton (1915) erected Unio whitecliffsensis
and Unio jaqueti respectively from the Coreena
Fm. of White Cliffs and the Griman Creek Fm. of
Lightning Ridge. McMichael (1956) revised this
material and assigned U. jaqueti to Velesunio and
U. whitecliffsensis to Hyridella. The two speci-
mens figured by Dettman et al. (1992) are here
assigned to Velesunio goondiwindiensis
(Dettmann et al., 1992: 244, Fig 19(j)) and Pro-
tovirgus wintonensis (Dettmann et al., 1992: 244,
Fig 19(i)).

List of Cited Localities. QML229: Meuller
Range, west of Cork Station, near Winton, CWQ,
Winton Formation, Cenomanian. QML379:
Franklin Station, "Trevor Cluff's' locality in pad-
dock west of Mt Gideon below small tabletops
and west on flats towards hills, near Winton,
CWQ. Winton Formation, Cenomanian.
QML570: Te Apiti Stn, NW of Goondiwindi,

SEQ, Dam site. Griman Creek Formation. Late
Albian.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA
Class BIVALVIA
Order UNIONOIDA Stoliczka, 1871
Family MUTELIDAE Gray, 1847

Protovirgus McMichael, 1956

Protovirgus wintonensis sp. nov.
(Fig 1. A-E)

Unioid, Dettmann et al. 1992: 244, fig. 19(i).
ETYMOLOGY. For the town of Winton.

MATERIAL EXAMINED. HOLOTYPE: QMF-
34635, L379; Franklin Stn. Trevor Cluff s locality, Mt
Gideon. PARATYPES: QMF5681-5682, QMF34645,
QMF34646 from QML570; QMF34634, QMF34644,
QMF34647, QMF34648 from QML379.

DIAGNOSIS. Small to medium-sized, equi-
valved, elongate unioid with slightly inflated
umbones and fine, comarginal ornament. Hinge
straight; anterior adductor muscle raised on plat-
form. Tapering fairly strongly with well-rounded
posterior margin.

DESCRIPTION. Maximum height ranges from
27-34mm, width 19-30mm and length 70-80mm
(Table 1). Shell elongate, hinge straight, ligament
thin, extending approximately 2/3 of shell length.
Fine comarginal ornament of growth lines.
Rounded posterior margin. Beak unsculptured.
Shell thin. Anterior adductor muscle positioned
behind umbo, raised on platform. Posterior mus-
cle scar unknown. Dentition unknown.

224

REMARKS. The tapering
form, sharp ventral margins and
strongly anterior umbones are
typical of Protovirgus. The
type species, P. dunstani
(Ethridge Jr 1888, described by
McMichael, 1956; 232, fig. 8)
from the Triassic of NSW is
approximately half the length
and much more linguiform
than P. wintonensis. P. flemingi
McMichael (1956: 232-233,
fig. 1-3) from the Cretaceous of
New Zealand 1s approximately
two-thirds the size, and the
umbones are placed more pos-
teriorly than in P. wintonensis,
P. jaenschi Ludbrook, 1961
from the Triassic of SA is
slightly larger at 80mm long,
31mm high, and has a less de-
veloped muscle scar and is
more ovate than P. wintonensis.
P. jaenschi has an inflexion in
the posterior 1/3 which is not
seen in P. wintonensis. P. coatsi
Ludbrook, 1961 from the
Triassic of Leigh's Creek is
smaller (two-thirds the length)
with more concave ventral and

MEMOIRS OF THE QUEENSLAND MUSEUM

convex dorsal margins. P
clellandi Hocknull, 1994 from
the Triassic of Ipswich, south-
east Queensland is much

FIG. 1. A-E, Protovirgus wintonenis sp. nov. A-C, Holotype QMF34635, x
1.2 (A, right valve; B, left valve; C, dorsal view). D, E, Paratype
QMF34634, internal mold x 1.2 (D, dorsal view; E, left valve). F-H,
Prohyria macmichaeli sp. nov. Paratype QMF34638 x 1.2 (F, right valve;

larger, the holotype being
123mm long and 39mm high,
and is sharper posteriorly with greater inflation of
the umbones.

Prohyria McMichael, 1956

Prohyria macmichaeli sp. nov.
(Fig. 1. F-H, Fig. 2. A-C)

ETYMOLOGY. For Donald F. McMichael for his
studies of freshwater bivalves.

MATERIAL EXAMINED. HOLOTYPE: QMF-
34636, OML 375; Franklin Stn. Trevor Cluff s locality,
Mt. Gideon. PARATYPES: QMF5677 from QML570
and QMF34637 from QML379, QMF34638 from
QML229.

DIAGNOSIS. Medium-sized, equivalved unioid,
ovale, rugose ornament, umbones anterior and
inflated, beak slightly sculptured, shell thick.

G, left valve; H. dorsal view).

DESCRIPTION. Maximum height 32mm, width
27mm and in length 55mm (Table 2). Elongate-
ovale, umbones anterior and inflated. Anterior
margin convex, ornament rugose, beak slightly
TABLE 1. Morphometric data for specimens of Pro-

tovirgus wintonensis sp. nov. from the Winton and
Griman Creek Formations. Measurements in mm.

CRETACEOUS FRESHWATER BIVALVES FROM QUEENSLAND

sculptured. Hinge straight,
short. Shell thick, ligament
short, and thick. Escutcheon
relatively broad, anterior ad-
ductor muscle small, raised
slightly and orientated an-
teroventrally. Dentition un-
known.

REMARKS. The anteriorly
placed, inflated umbones, ru-
gose ornament, elongate-ovoid
shape ally the present material
to Prohyria. The type species,
P. johnstoni (Ethridge Jr, 1881)
of McMichael (1956: 227-228,
figs 6&7) from the Oligocene
of Tasmania is much larger, up
to 126mm long, wider (up to
55mm) and the dorsal margin is
sharper. P. eyrensis (Etheridge,
1892) of McMichael (1956:
228-230, figs 8-12) from the
Triassic of Leigh's Creek,
South Australia is also longer
(up to 102mm), wider (up to
50mm), and umbones are
placed more posteriorly than P.
macmichaeli.

Velesunio Iredale, 1934

225

Velesunio goondiwindiensis
Sp. nov.
(Fig. 2. D-J.)

FIG. 2. A-C, Prohyria macmichaeli sp. nov. Holotype QMF34637, x 1.2 (A,
right valve; B, left valve; C, dorsal view). D-J, Velesuniuo goondiwindiensis
sp. nov. D-F, Paratype QMF34639, x 1.2 (D, right valve; E, left valve; F,
dorsal view). G-I, Holotype QMF5684, x 1.2 (G, right valve; H, left valve;

Unioid, Dettmannetal.1992 : fig. 19j.

ETYMOLOGY. For the town of Goondiwindi.

MATERIAL EXAMINED. HOLOTYPE: QMF5684,
from QML570; Dam at ‘Te Apiti’ Stn., near
Goondiwindi, SEQ. PARATYPES: QMF5683, QMF
5685, QMF5686, QMF34639-34641 from QML570.

DIAGNOSIS. Small, elongate-ovoid, fine com-
arginal ornament and thin shell. Umbones ante-
rior. Anterior adductor muscles small, elongate,
raised slightly. Shell expanded posteroventrally.

DESCRIPTION. Small, equivalved unioid. Elon-
gate-ovoid, compressed with fine comarginal or-
nament. Maximum height ranges 33-35mm,
width 16-18mm and length 35-38mm (Table 3).
Umbones placed anteriorly, at anterior 1/3 of total
length. Beak sculptured but small. Dorsal and

I, dorsal view). J, Paratype QMF34640, x 1.2, dorsal view.

ventral margins run subparallel, forming an ex-
panded posterior. Hinge 1/2 the length of total.
Ligament thin, escutcheon narrow. Maximum
height posterior to umbones. Adductor muscle
scars small, elongate and orientated antero-ven-
trally. Dentition unknown.

REMARKS. The elongate-ovoid, anteriorly
placed umbones with subparallel dorsal and ven-

TABLE 2. Morphometric data for the specimens of
Prohyria macmichaeli sp. nov. from the Winton and
Griman Creek Formations.

tu
LA
a

TABLE 3. Morphometric data for (he specimens of
Velesunio goondiwindiensis sp. nov. from the Winton
and Griman Creek Formations. Measurements in mm.

iral borders expanding posteriorly indicate
Velesunio Iredale, 1934, V, jagueti (Newton,
1915) of McMichael (1956: 240) from the Gri-
man Creek Formation of Lightning Ridge is
larger, more elongate (37-42mm long) and nar-
row (12-15mm) with umbones placed more pos-
teriorly than V goondiwindiensis. All recent
species of Velesunio are much larger, more robust
and have å more pronounced winged posterior
(McMichael & Hiscock, 1958) than V
goondiwindiensis.

ACKNOWLEDGEMENTS

I would like to thank Alex Cook, Natalie
Camilleri and Paul Tierney for their help and
encouragement. The described material was col-
lected by Mary Wade, H.J. McMilland and Ralph
E. Molnar.

LITERATURE CITED

BURGER, D. 1970. Early Cretaceous angiosperm pol-
len grains from Queensland. Australian Bureau of
Minerals and Resources, Geology and Geophys-
ics, Bulletin 166: 1-15.

1986. Palynology, cyclic sedimentation and palaeo-
environments in the Late Mesozoic of the
Eromanga Basin. Geological Society of Aus-
tralia: Special Publication 12: 53-70,

1995. Timescales. 9, Cretaceous. AGSO Record
1995/38:1-35.

DETTMANN, ME, MOLNAR, RE, DOUGLAS,
J.G., BURGER, D., FIELDING, C,, CLIFFORD,

MEMOIRS OF THE QUEENSLAND MUSEUM

H.T., FRANCIS, J., JELL, P, RICH, T., WADE,
M.. RICH, P.V., PLEDGE, N.. KEMP, A. &
ROZEFELDS, A. 1992. Australian Cretaceous
terrestrial faunas and floras biostratigraphic and
biogeographie implications, Cretaceous Research
13: 207-262.

DETTMANN, M.E. & PLAYFORD, G. 1969. Palynol-
ogy of the Australian Cretaceous: A review. Pp.
174-21D. In Campbell, KS W (ed.) ‘Stratigraphy
and Palaeontology. Essays in honour of Dorothy
Hill,’ (Australian National University Press: Can-
berra).

EXON, N.F. & SENIOR, B.R, 1976, The Cretaceous
geology of the Eromanga and Surat Basins. BMR
Journal 1: 33-50.

HAAS, F. 1969. Superfamily Unionacea. Pp. N411-468.
In Moore, R.C. (ed), "Treatise on Invertebrate
Palaeontology’ Part N. Vol. 2, Mollusca 6,
Bivalvia. (Geological Society of America and
University of Kansas; Lawernce, Kansas).

HELBY, R., MORGAN, R. & PARTRIDGE, A.D.
1987. A palynological zonation of the Australian
Mesozoic: Association of Australian Palaeo-
ntologists Memoir 4 1-94.

HOCKNULL, S.A. 1994. A new freshwater bivalve
from the Triassic of southeastern Queensland
Memoirs of the Queensland Museum. 37(1): 146.

IREDALE, T. 1934 The freshwater mussels of Aus-
tralia. Australian Zoologist 8: 57-78.

LUDBROOK, N.H. 1961. Mesozoic non-marine
Mollusca (Pelecypoda: Unionidae) from the north
of South Australia. Transactions of the Royal So-
ciety of South Australia 84: 139-148.

McMICHAEL, D.F. 1956. A review of the fossil fresh-
water mussels (Mollusca, Pelecypoda) of Austral-
asia. Proceedings of the Linnean Society of New
South Wales 81:223-244,

McMICHAEL, D.F. & HISCOCK D. 1958. A mono-
Eraph of the freshwater mussels (Mollusca:
Pelecypoda) of the Australian region. Australian
Joumal of Marine and Freshwater Research 9:
372-508.

MORGAN, R. 1980. Palynostraugraphy of the Austra-
lian early and middle Cretaceous Geological Sur-
vey of New South Wales Palaeontological
Memoir 18: 1-153.

PSEUDANTHESSIUS NEWMANAE, NEW SPECIES (COPEPODA:
POECILOSTOMATOIDA: PSEUDANTHESSIIDAE) FROM MARINE
TURBELLARIANS IN AUSTRALIA

ARTHUR G. HUMES

Humes, A.G. 1997 06 30: Pseudanthessius newmanae, new species (Copepoda:
Poecilostomatoida: Pseudanthessiidae) from marine turbellarians in Australia. Memoirs of
the Queensland Museum 42(1): 227-231. Brisbane. ISSN 0079-8835.

Pseudanthessius newmanae, a new species of poecilostomatoid copepod, is associated in
Australia with the marine turbellarians Tyrthosoceros lizardensis Newman & Cannon (in
press) and two undescribed species of Pseudobiceros. The female of the new copepod may
be distinguished from its 37 congeners by a combination of characters: the length of the body,
the length to width ratio of the caudal ramus, and the shape of the genital double-somite.
This is the second species of Pseudanthessius recorded as associated with Turbellaria. [ ]
Pseudanthessius, Copepoda, Poecilostomatoida, associates, Turbellaria, Australia.

Arthur G. Humes, Boston University Marine Program, Marine Biological Laboratory,
Woods Hole, Massachusetts 02543, U.S.A.; 21 March 1997.

Most of the 37 species in the genus
Pseudanthessius are associated with marine in-
vertebrates (polychaetes, bivalves, asteroids,
echinoids, ophiuroids, crinoids, and holothuri-
ans). Pseudanthessius nemertophilus Gallien,
1936, lives with a nemertean, Lineus longissimus
Sowerby, on the Atlantic coast of France. Only
one species, Pseudanthessius latus Illg, 1949, is
associated with Turbellaria (with Cryptophallus
magnus Freeman, now known as Kaburakia ex-
celsa Bock, in Washington and California).

This paper contains the description of a second
species of Pseudanthessius living with marine
turbellarians, this time in Australia.

MATERIALS AND METHODS

The copepods, collected by Dr Leslie J. New-
man, were preserved in 70% ethanol. They were
measured and studied in lactic acid, according to
the method described by Humes & Gooding
(1964). All figures were drawn with the aid of a
camera lucida. The letter after the explanation of
each figure refers to the scale at which the figure
was drawn.

Order Poecilostomatoida Thorell, 1859
Family Pseudanthessiidae
Humes & Stock, 1972
Genus Pseudanthessius Claus, 1889

Pseudanthessius newmanae, sp. nov.
(Figs la-g, 2a-k, 3a-j)

MATERIAL EXAMINED. HOLOTYPE. QMW21831,
ovigerous 9, from the polyclad turbellarian

Pseudobiceros, undescribed species, under rubble, in a
depth of approximately 5m, between South Island and
Palfrey Island, Lizard Island lagoon, Queensland, Aus-
tralia, 14?40' S, 145?28"E, L. Newman and A. Flowers
collectors, 5 April 1995. ALLOTYPE: d
(QMW21830), from Pseudobioceros, undescribed
species, sublittoral, under rubble, South Passage, Coral
Bay, Eel Bommie, Western Australia Australia, L.
Newman and A. Flowers collectors, 5 May 1996.
PARATYPES. nonovigerous 9, QMW21349, same
data as for holotype. 2 9 9, (1 in QM W21348, other
dissected, in the collection of the author), from
Tytthosoceros lizardensis Newman & Cannon, in
press, low water mark, South Island, Lizard Island
lagoon, Queensland, Australia, 14?40' S, 145?28'E, L.
Newman and A. Flowers collectors, 31 March 1995. 1
9, QMW21398, from Tytthosoceros lizardensis, under
rubble, sublittoral, in a depth of 6m, South Passage,
Coral Bay, ‘Eel Gardens’, Western Australia, Aus-
tralia, L. Newman and A. Flowers collectors. 4 May
1996. 2 99, QMW21399, 1 å (QMW21399), from
Pseudobioceros, undescribed species, sublittoral,
under rubble, South Passage, Coral Bay, Eel Bommie,
Western Australia Australia, L. Newman and A. Flow-
ers collectors, 5 May 1996.

DESCRIPTION. Female (description based on
specimens from Tytthosoceros lizardensis at Liz-
ard Island): Body (Fig. la) elongate. Average
length (not including setae on caudal rami)
1.44mm (1.34-1.54mm) and greatest width
0.66mm (0.60-0.72mm), based on 4 specimens.
Greatest dorsoventral thickness 0.35mm. Somite
bearing leg I separated from cephalosome by
dorsal transverse furrow. Epimera of metasomal
somites posteriorly rounded. Ratio of length to
width of prosome 1.58:1. Ratio of length of pro-
some to that of urosome 1.46:1.

be
LA
Dei

Somite bearing leg 5 (Fig.
Ib-d) 70 x 172um. Genital
double-somite in dorsal view
190 x 172um, ratio 1.05:1, ;
somewhat swollen laterally in — / \
anterior two-thirds, but with — .—
sides subparallel in posterior `
third (width 115m), Genital
areas located dorsolaterally
near middle of double-somite,
each area bearing 2 very un-
equal setae Alum and 14j.m
and minute process (Fig. 1d).
Three postgenital somites from
anterior to posterior 88 x 102,
73 x 94, and 81 x 82jum.
Caudal ramus (Fig, le) 112 x
391m, ratio 2.87:1. Outer lat-
eral seta 62jum, dorsal seta
27m, both smooth. Outer-
most terminal seta 96jum, in-
nermost terminal seta 140j.m,
and 2 median terminal setae
308um (outer) and 462pm
(inner), all with lateral setules.
Body surface smooth, but
few minute refractile points on x
caudal ramus (Fig. 1e). e (5. )

Egg sac (Fig. 1f) elongate, d PR
muluseriate, 995 x 297pm, ^y
ratio 3.35:1. Eggs 68jum in
average diameter (range 62- a ENS
70pm). Ss

Rostral area subquadrate
(Fig. 1g). Antennule (Fig. 2a)
320um long. Lengths of its 7
segments (measured along
their posterior nonsetiferous
margins): 16 (47 um along an-
terior margin), 101, 26, 52, 49,
31, and 18m, respectively.
Formula for armature: 4, 13, 6,
3, 4 + 1 aesthetasc, 2 + I
aesthetasc, and 7 + 1 aesthetasc. Several setae
unusually long, all smooth. Antenna (Fig. 2b)
4-segmented, with armature 1, 1, 3, and II + 4.
Third segment 27m along outer side, 22um
along inner side; fourth segment 35jum along
outer side and 43um along inner side. Two claws
slender, 40um and 53jum. All elements smooth,

Labrum (Fig. 2c) with 2 elongate linguiform
lobes. Mandible (Fig. 2d) with constricted prox-
imal area followed on concave side by transverse
row of long spinules, and on convex side hy 2
small spiniform processes and minute spinules.

MEMOIRS OF THE QUEENSLAND MUSEUM

0.2 mm

Di mm

FIG. 1. Pseudanthessius newmanae, new species. Female. a, body, dorsal
(scale A); b, urosome, dorsal (B); c, urosome, ventral (B); d, urosome,
lateral (B); e, anal somite and caudal ramus, dorsal (C); f, egg sac, ventral
(D); g, rostral area, ventral (B). Al = antennule, A2 = antenna,

Lash tapered with very small lateral spinules.
Paragnath small lobe. Maxillule (Fig. 2e) with 4
setae. Maxilla (Fig. 2f) with unarmed proximal
segment; distal segment with outer small smooth
seta and long inner seta bearing unilateral setules.
Lash long with outer row of graduated spines.
Maxilliped (Fig. 2g) 3-segmented. First segment
unarmed, swollen second segment with 2 setae,
and small third segment with 2 minute setae and
terminating in pointed tip.

Ventral area between maxillipeds and first pair
of legs (Fig. 2h) slightly protuberant.

NEW AUSTRALIAN MARINE TURBELLARIAN 229

Legs 1-4 (Figs 2i-k, 3a) with
3-segmented rami except l-seg-
mented endopod in leg 4. Table
I shows the formula for arma-
ture (Roman numerals indicat-
ing spines, Arabic numerals
representing setae).

Third segment of endopod of
leg 1 with outermost seta rather
spinelike (Fig. 2i). Leg 4 (Fig.
3a) with exopod 180um. Endo-
pod 114 x 36jum, ratio 3.17:1,
its 2 terminal fringed spines
75um (outer) and 101um
(inner). Anterior surface of en-
dopod with slight incomplete
suggestion of division. Both
sides of endopod with long lat-
eral hairlike setules.

Leg 5 (Fig. ld) without free
segment, consisting of slight
ridge bearing 2 setae 52j.m and
491m, and adjacent seta
50jum. All setae smooth.

Leg 6 represented by 2 setae,
3lum and l4jum, on genital
area (Fig. ld).

Color of living specimens
unknown.

Male (description based on
specimens from Pseudo-
biceros, undescribed species,
in Western Australia): Body
(Fig. 3b) with prosome more
slender than in female. Length
1.12mm (1.07-1.19 mm) and
greatest width 0.32mm (0.29-
0.36mm), based on 3 speci-
mens. Greatest dorsoventral
thickness 0.25mm. Ratio of
length to width of prosome
1.79:1. Ratio of length of pro-
some to that of urosome 1.31:1.

Somite bearing leg 5 (Fig.
3c) 36 x 86um. Genital somite
elongate, 130 x 102jum, longer

FIG. 2. Pseudanthessius newmanae, new species. Female. a, antennule,
dorsal (scale B); b, antenna, posterior (E); c, labrum, ventral (C); d,
mandible, dorsal (C); e, maxillule, posterior (C); f, maxilla, inner (C); g,
maxilliped, postero-inner (C); h, area between maxillipeds and first pair of
legs, ventral (B); i, leg 1 and intercoxal plate, anterior (B); j, leg 2 and
intercoxal plate, anterior (B); k, leg 3 and intercoxal plate, anterior (B).
MXPD = maxilliped. P1 = leg 1.

than wide; in lateral view (Fig. 3d) 133 x 101m. TABLE 1. Pseudanthessius newmanae, new species.
Four postgenital somites from anterior to poste- Formula for armature.

rior 57 x 63, 60 x 57, 52 x 52, and 52 x 52um.

Caudal ramus (Fig. 3c) resembling that of fe-
male but smaller, 73 x 25jum, ratio 2.92:1.

Rostrum, antennule, antenna, labrum, mandi-
ble, maxillule, and maxilla similar to those in
female. Maxilliped (Fig. 3e) with second segment

coxa | basis endopod

0-1; HLL4
02; ILL3
0-2; ILI2

bearing 2 setae and 2 rows of spinules. Claw (Fig.

230

3f) 165jum long, bearing prox-
imally 2 unequal setae, longer
seta with few setules on inner
side, and having large terminal
lamella and small subterminal
tooth.

Ventral area between maxil-
lipeds and first pair of legs as in
female.

Legs 1-4 segmented and
armed as in female, except for
sexual dimorphism in endopod
of leg 1, with clawlike process
between spine and reduced first
seta (Fig. 3g, h).

Leg 5 (Fig. 3d) as in female.

Leg 6 (Fig. 3d, i) postero-
ventral flap on genital somite
bearing 2 setae.

Spermatophore (Fig. 3j),
seen only inside genital somite
of male, elongate, 135 x 44u.m.

ETYMOLOGY. The new species
is named for Leslie J. Newman,
University of Queensland, who
sent the copepods to me for study.

REMARKS. Only one species
of Pseudanthessius has been
recorded as associated with
marine flatworms. Pseud-
anthessius latus Illg, 1950,
liveson a polyclad turbellarian,
Kaburakia excelsa Bock (see
Faubel, 1983) (2 Crwpto-
phallus magnus Freeman), on
the coasts of Washington and
California (Illg, 1950). As Illg
(1950) pointed out, Wilson's
(1935) specimens of
Pseudanthessius obscurus (A.
Scott, 1909), from large gray
flatworms in California, are P.
latus.

Pseudanthessius newmanae
differs from P. latus as follows:
the average length of the female is 1.44mm (ver-
sus 1.82mm in P. latus), the second segment of
the antennule is relatively short (versus exceed-
ing the combined lengths of the five terminal
segments in P. latus), the ratio of the length to the
width of the caudal ramus is 2.87:1 (versus ap-
proximately 6:1 in Illg's fig. 1k), and the endopod
of leg 4 is elongate (versus inflated in Illg's fig.
li).

MEMOIRS OF THE QUEENSLAND MUSEUM

0.05 mm
r——.

FIG. 3. Pseudanthessius newmanae, new species. Female. a, leg 4 and intercoxal
plate, anterior (scale B). Male. b, body, dorsal (F); c, urosome, dorsal (B);
d, somite bearing leg 5, genital somite, and first postgenital somite, lateral
(E); e, maxilliped, postero-inner (C); f, claw of maxilliped, antero-outer
(G); g, endopod of leg 1, anterior (C); h, third segment of endopod of leg
1, anterior (H); i, genital somite and first postgenital somite, showing leg
6, ventral (E); j, spermatophore, inside body of male, lateral (E).

A combination of characters of the female will
serve to separate the new species from its 37
congeners. In 16 species of Pseudanthessius the
length of the body is less than 1 mm. In 12 species
the ratio of length to width of the caudal ramus is
3.5:1 or more. In 5 species the ratio of the caudal
ramus is less than 2:1. Three species cannot be
distinguished from the new species by these cri-
teria, but show other characters by which they
may be separated from P. newmanae. In

NEW AUSTRALIAN MARINE TURBELLARIAN

Pseudanthessius sauvagei Canu, 1891, the geni-
tal double-somite of the fernale is slender, about
1.7:1, not laterally expanded, and the fourth seg-
ment of the antennule is elongate. In P spinifer
Lindberg, 1946, the genital double-somite of the
female in dorsal view has pointed lateral pro-
cesses and the mandible has à long slendér blade.
in P. vinnulus Humes, 1977, the genital double-
somite of the female is quadrate and the longest
seta of leg 5 is longer than the genital somite. In
one species, Pseudanthessius faouzii Steuer,
1940: 21, the female is unknown. The male of P.
faouzii differs from the male of the new species
in that the length of the body is 0.574 mm, and
the caudal ramus is approximately 2:1.

The nature of the sexual dimorphism in the
third segment of the endopod of leg | of the male,
with a clawlike process between the spine and the
reduced first seta, is unique among the 22 species
of Pseudanthessius Where males are known and
where the endopod of leg 1 has been described or
illustrated.

Pseudanthessius newmanae occurs on three
species of pseudocerotid worms (two of them
undescribed) on both eastern and western coasts
of Australia. The nature of the association of the
copepods with the flatworms is not known, The
copepods move actively on the host, and may
perhaps feed on the mucus of the worms. It is of
interest that Tytthosoceros lizardensis is very
toxic (L.J. Newman, in correspondence).

ACKNOWLEDGEMENTS

I thank Leslie J. Newman for her special effort
in collecting the copepods and allowing me to
study them.

LITERATURE CITED

Canu, E. 1891. Les copépodes marins du Boulonnais.
V. Les semi-parasites. Bulletin Scientifique de la
France et de la Belgique 23: 467-487.

231

Faubel, A. 1983. The Polycladida, Turbellaria. Proposal
and establishment of a new system. Part |. The
Acotylea, Mitteilungen aus dem hamburgischen
zoologischen Museum und Institut 80: 17-121.

Galhen, L. 1936. Pseudanthessius nemertophilus nov.
Sp. copépode commensal de Lineus longissimus
Sowerby. Bulletin de la Société Zoologique de
France 60: 451-459.

Humes, A.G. 1977. Pseudanthessiid copepods
(Cyclopoida) associated with crinoids and echi-
noids (Echinodermata) in the tropical western Pa-
cific Ocean. Smithsonian Contributions to
Zoology 243; 1-43,

Humes, AO. & Gooding, RU. 1964. A method for
studying the external anatomy of copepods,
Crustaceana 6: 238-240.

lg, P.L. 1950. A new copépod, Pseudanthessuts latus
(Cyclopoida; Lichomolgidae), commensal with a
marine flatworm, Journal of the Washington
Academy of Sciences 40: 129-133,

Lindherg, K. 1946. Un nouveau copépode poecilostome
de l'inde de là famille de Lichomolgidae;
Pseudanthessius spinifer, n. sp. Bulletin de la
Société Zoologique de France 70: 81-84.

Newman, L.J. & Cannon, L.R.G. (In press.) Bulaceras,
new genus and Tymhosoceros, new genus
(Platyhelminthes: Polycladida) from the Great
Barrier Reef, Australia, and Papua New Guinea.
Raffles Bulletin of Zoology.

Sars, G.O. 1917. An account of the Crustacea of Nor-
way. Vol. V Copepoda Cyclopoida Parts XI and
XII Clausidiidae, Lichomolgidae (pari), Bergen
Museum, Bergen, Norway.

Scott, A. 1909, The Copepoda of the Siboga Expedition.
Part I. Free-swimming, littoral and semi-parasitic
Copepoda. Siboga Expedition 29a; 1-323.

Steuer, A. 1940. Uber einige Copepoda Cyclopoida der
Mediterranen Amphioxussande. Notizen des
Deutsch-Italienischen Institutes für
Meeresbiologie in Rovigno d'Istria 11: 1-27.

Wilson, C.B. 1935. Parasitic copepods from the Pacific
coast. American Midland Naturalist 16: 776-797.

NEW INFORMATION ON PSEUDECHIS PAPUANUS
(THE PAPUAN BLACK SNAKE), A MEDICALLY SIG-
NIFICANT ADDITION TO AUSTRALIA'S REPTILES.
Memoirs of the Queensland Museum 42(1) 232. 1997:-
Pseudechis papuanus Pewrs & Doria, 1879 (the Papuan Black
Snake) is known from `. along the south coast of New
Guinea, from Prince Frederik Hendrik Island in Irian Jaya,
ihe Western Province (TransFly and Lake Murray), and the
coast af the Central Province and Yule Island" (Parker, 19821.
Iris rare in Papua New Guinea (Whitaker et al., 1982, O'Shea,
1996), and it has been suggested by Parker (1982), Whitaker
et al. (1982) and O'Shea (1996) (hat its apparent rarity may
be attributed to attempted predation on Bufo marinus (the
introduced Cane or Marine Toad), although these reports ure
not based on field studies of P. papuanus. Published data on
its feeding habits differ slightly. Whitaker et al. (1982) regard
P. papuanus as a frog eater, while O'Shea (1996) states it
feeds "primarily on frogs, bupalso small mammals, lizards and
possibly ground nesting birds’.

P. papugnus has a poten venom and is n porentially
dangerous snake to humans (Campbell, 19673, 1967b; Parker,
1982: Whitaker, et al.. 1982; and O'Shea, 1996). Campbell &
Chesterman (1972) examined the action of its venom and
showed it lo be ‘anticoagulant, fibrinogenolytic and
caseinalytic'. Parker (1982) reported that P. papuanus caused
many fatalities in southern coastal Papua New Guinea and
CH Shea (1996) regards it as ‘the second most venomous land
suake in PNG after The laipan'.

In October 1996, E. Vanderduys, A. McManus and I were
conducting a reptile survey of the Torres Strait islands, Mr
Manual Namoa, Acting Principal of the Saibai Island State
School, gave me 3 snake which had been killed in the local
school grounds, It had been held for some time in preservative
and no date of collection was recorded.

The snake has been photographed (Fig 1.) and added ro the
Queensland Museum reference collection (QM 62555). In the
following standard identification characteristics, QMJ62555
from Saibai Island (9°23'S, 142°40'E) is readily referred to
P. papuanus as defined by Golay (1985) and O'Shea (1996):
Midbody scales 19; Ventral scales 221; Subcaudal scales 51,
anterior 26 enüre. The specimen (in spirit) is immaculate
black above and dark grey below, with a pink ventral flush on
the chin, posterior body and tail. Ic is å female with à total
length of 158 cm and a tail length of 21cm. The gut contained
3 spirurid nematodes (Physaloprera sp.) but no prey items,

Saihai Is. is only 4km south of the Papua New Guinea
zoast. ft is thus geographically New Guinean but, politically
Australian, lt is low-lying and mangrove-fringed with pockets
of monsoon forest. lts most conspicuous topographic feature
isan exensive mozaic of freshwater and saline swamps, Parker
11982) and O'Shea (1996) note that P. papuanus has a pref-
erence for swampy areas ard its occurrence on Saibai Is. is
consistent with their observations. A large freshwater swamp
lies only about 100 metres from the school and during the "wet
season’ it undoubtedly expands to be very close to rhe school
prounds.

The people of Sabai Is. are familiar with P. papuanus,
know it as the 'Papuan Black Snake" and regard itas danger-
pus and abundant, There are no records of B, marinus on the
island and, given the apparent decline of P, papuanus in Papua
New Guinea, Saibai Is, may prove to be å stronghold for this
species.

The discovery of P pppuanis on Saibai Is, adds to the
Australian fauna another species of elapid snake known to
have inflicted fatal bites or believed capable of doing so, lis
congener, P, australis, occurs in Papua New Guinea and on
Cape York Peninsula, and has recently been recorded on

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Pseudechis papuanus (QM) 62555) from Saibai Is.

Prince Of Wales Island (QMJ62560) between them. To treat
biles from both species, Black Snake Antivenom is avitilable.
It should be stocked at least in the Thursday Island Hospaal
and, probably, in the clinic on Saihai Is.

Acknowedgements

[ thank Mr Manual Namoa, Acting Principal of Saibai
Island State School, for donating the specimen; Mr Eric
Vanderduys and Ms Joan Whittier for rhe opportun to
participate in their Torres Strait field survey: Department of
Agriculture, Hawaii, for funding the field work; Ms Anne
McManus for assistance in the field; Mr Jetf Wright (Queens-
land Museum) for photographing the specimen; Ms Tracy
Brown for developing the photograph: Dr Lester Cannon
(Queensland Museum) for identifying the nematodes; and Ms
Jeannette Covacevich (Queensland Museum) for assisting me
in compiling this note.

Literature Cited

CAMPBELL, C.H. 19673. Antivenene in the treaunent of
Australian and Papuan snake bite. Medical Journal of
Australia 2: 106-110.

1967h. The Papuan Black Snake (Psendechis papuanus)
and the effect of its bite, Papua New Guinea Medical
Journal 10; 117-121.

CAMPBELL, CH & CHESTERMAN, C.N. 1972. The ef-
fect of the venom of the Papuan Black Snake
(Pseudechis papuanus) on blood coagulation, Papua
New Guinea Medical Journal 15: 149- 154.

GOLA Y, P: 1985. Checklist and keys to rhe terrestrial pro-
teroglyphs of the world: Elapidae-Hydrophiidae,
Elapsoidea. Geneva.

O'SHEA, M. 1996, Å guide to the snakes of Papua New
Guinea, (Independent Publishing Group: Port
Moresby).

PARKER, F. 1982. Snakes of the Western Province. Wildlife
in Papua New Guinea No 82/1, (Hebamo Press:
Boroko),

WHITAKER, R,, WHITAKER, J, & MILLS, D. 1982 Rep
"les of Papua New Guinea, Wildlife in Papua New
Guinea No 82/2. (Hebarno Press: Borsko).

Stephen K. Wilson. Queensland Muscum, P. O. Bex 3300

South Brisbane, Queensland 4101.: 3 April 1997,

SOME NEW AND PREVIOUSLY KNOWN EARTHWORM SPECIES FROM CAPE

YORK PENINSULA (ANNELIDA; OLIGOCHAETA: MEGASCOLECIDAE)
B.G.M. JAMIESON

Jamieson, B.G.M. 1997 06 30; Some new and previously known species of earthworms from
Cape York Peninsula (Annelida: Oligochaeta: Megascolecidae), Memoirs of the Queensland
Museum 42( 1): 233-270. Brisbane. ISSN 0079-8835.

Ten new species, in five genera, are described from the Cape York Peninsula, Queensland:
Diplotrema attenuata sp. nov,; D, scheltingai sp. nov.; Neodiplotrema mcdonaldi sp. nov.,
and Terrisswalkerius meihvraithi sp. nov., from the Mcilwraith Range; Diplotrema
acropetra Sp. nov., Neodiplotrema altanmoui sp. nov., N. paripunciata sp. nov., and
Kayarmacia adelphicus gen. et sp. nov, from Cape Melville National Park; Terrisswalkerius
carbinensis sp, nov, from MI Carbine Tableland, and T. miseriae sp. nov., from Mt Misery.
Kayarmacia (=Rhododrilus) queenslandicus Michaelsen, 1916, is redescribed from the
Alice-Mitehell R. National Park, near the type-locality; it is clearly the sister-taxon of K.
adelphicus. The phylogenetic and biogeographic affinities of these species are briefly
discussed. [ ] Megascolecidae, phylogeny, biogeography, Oliguchaeta, Cape York Penin-

sula.

B.G.M. Jamieson, Zoology Department, University of Queensland, Brisbane 4072, Ans-

tralia; 21 March 1997.

The oligochaete fauna of the Cape Melville
National Park and of the MclIlwraith Range, in
North Queensland, has not previously been de-
scribed. As a result of the assiduous collecting of
Mr K.R. McDonald, new megascolecid species
from these areas, from Mt Carbine Tableland and
from Mt Misery, in the wet rropics, are described.
A new genus Kayarmacia is erected for one of
the new species from Cape Melville and for
Rhododrilus queenslandicus Michaelsen, 1916,
from the Alice-Mitchell River National Park
which is redescribed from new material.

Subfamily Acanthodrilinae

Genus Diplotrema Spencer, 1900. Emend.
Jamieson & Dyne, 1976

Diplotrema acropetra sp. nov. (Figs 1-4; 39)

TYPE LOCALITY, 14°43°54"S 144°46'45"E, Rocky
Peak, Cape Melville National Park, altitude 450m, on
sandstone plateau, base of trees where pigs had been
digging, edge of Banksia robur swamp, K.R. McDon-
ald, P.J. Lethbridge, 4 Apr 1995.

MATERIAL EXAMINED. HOLOTYPE. QMG212027
(includes microscope slide of right genital seta of IX).
PARATYPES. P1-9 QMG212028-212036. OTHER
MATERIAL. Several not designated types.
QMG213391.

DESCRIPTION, Length 38-55mm (H, Pi-9),
43mm (H). Width (midclitellar) 1.7-2.4mm (H,
P1-9), 2.3mm (H). Segments (longest and short-

est specimen P3 and 9) 163-169, H 166. Pigment-
less buff in ethanol. Prostomium large, epilobous
1/2, closed, the lateral margins of the dorsal
tongue strongly convergent and concave (H, P1,
P3-5) but in P2 prolobous with longitudinal
grooves on the peristomium giving an impression
of an epilobous condition. Peristomium longer
than segment I; neither it nor the prostomium
bisected ventrally. Dorsal pores poorly visible,
commencing on or behind the clitellum (H, P1-5).
Setae 8 per segment, commencing on II; in XII,
aa; ab; be: cd: dd = 7,2: 1.0: 6,6: 0,7: 29.2; or
13.6: 1.9: 12.5: 1.3: 55.1% (Hy, ventral setal
couples of XVIII absent; those of XVII and XIX
modified as enlarged penial setae; ventrål setae
of X forming genital setae. Nephropores not vis-
ible, Clitellum (poorly developed in the holo-
type); in XIII-XVTI, annular, but interrupted
ventrally in XVII by the prostatic porophores
(P1). Male pores not visible. Prostatic porophores
2 pairs, in XVII and XIX, each an elliptical pa-
pilla centred approximately in ah lines; penial
setae protuberant from one or both pairs (H, P1-
5), seminal grooves not apparent (H) or a faint
outwardly eurved (parenthetic) groove connect-
ing the prostate pores on each side (e.g, P4).
Genital tumescences: unilateral in X, (H, P1), left
(P1, 2); also paired in XI (P2); or absent in X and
left only in XI (P3). Genital markings: an un-
paired midventral ellipical to rounded oblong
pad between the ventral setal couples (in aa) in
each of intersegmental furrows 13/14-15/16,
16/17, 17/18, 20/21-23/24; those in 16/17 and

234

spermathecal
pore 1

prostatic
porophore 1

prostatic
porophore 2

markings

XXIV

FIG. 1. Diplotrema acropetra sp. nov. Holotype,
QMG212027. Ventral view of forebody and clitellar
region (clitellum drawn from Paratype 1). Note that
in this species the spermathecal pores are transposed
one segment posterior to the normal megascolecid
location.

17/18 small; a suggestion of a pad in 18/19 (H).
Female pores not visible. Spermathecal pores 2
pairs, in 8/9 and 9/10 (H); this unusual location
being confirmed in P1-9.

Septa 8/9 and 9/10 the thickest, fairly strongly
thickened. Dorsal blood vessel single, continuous

MEMOIRS OF THE QUEENSLAND MUSEUM

prostomium

A

vas deferentia

prostate duct

spermathecal
diverticulum

spermathecal ampulla

FIG. 2. Diplotrema acropetra sp. nov. Holotype,
QMG212027. A, dorsal view in region of prosto-
mium. B, right prostates, with penial setae omitted;
only the anterior prostate is shaded C, right sperma-
theca of IX.

EARTHWORMS FROM CAPE YORK PENINSULA

FIG. 3. Diplotrema acropetra sp. nov. Holotype, QMG212027. A and B, two right penial setae of XIX. C and
D, different views of the tip of seta show in A. E-G, successively more posterior to near basal regions of same.

onto the pharynx. Last hearts in XIII; in XI-XIII
stout, laterooesophageal, the connectives to the
supraoesophageal vessel being wide, those to the
dorsal vessel scarcely apparent; commissurals in
X anteriorly slender, with dorsal connectives
only. Gizzard very large, an elongate, glossy,

muscular cylinder about three times longer than
wide, in V, but its posterior end at the level of
segment X. Oesophagus segmentally dilated, and
longitudinally compressed, in each of XIV-XVI;
the walls in these segments with numerous
closely situated, deep, internal lamellae but

236 MEMOIRS OF THE QUEENSLAND MUSEUM

lumen not constricted off from that of the oesoph-
agus. Intestine commencing with abrupt expan-
sion in XVIII. Dorsal typhlosole very well
developed from about XXIX posteriad.
Holonephric throughout; at least two pairs of
nephridia anterior to the gizzard are convoluted,
not tufted, and each nephridium sends a duct
anteriorly to the pharynx. Typical nephridia each
with preseptal funnel and slender, avesiculate
duct discharging presetally in the vicinity of the
dorsal setal couples (cd). Metandric; large semi-
nal funnels, with spermatozoal iridescence, in XI
only. Seminal vesicles racemose, in XII only.
Flattened ovaries, with many egg strings, in XIII.
Two pairs of thickly tubular prostates, with ectal
ducts discharging in XVII and XIX. The anterior
pair is very much the larger and winds from XVII
to XXII; the tortuous muscular duct is joined near
its ectal end by the thick, vas deferens, the width
of which, greatly exceeding that of a normal vas,
suggests that it is a sperm reservoir. Posterior
prostates winding from XIX to XXIII; the ectal
duct shorter, though still long, less tortuous and
poorly muscularized. Both pairs of prostate ducts
overlain by penisetal follicles. Penial setae
curved through a right angle or slightly less; the
tip widened reflexed 'dorsally' for a short dis-
tance before curving 'ventrally' to a V-shaped tip,
the tip thus having the appearance of a poised
cobra; in frontal view this may give the spurious
appearance of a ladle-shape; sculpturing in the
midregion consisting of palisades of pointed
scales in incomplete circlets, the circlets being
spaced longitudinally at approximately 10j.m in-
tervals; the scales sparser basally; some groups of
scales continuing to, but not including, the mod-
ified tip; alternatively the scales may form small
obliquely arranged groups, each group consisting
of a small protuberant semicircle of pointed teeth
which overlie a depression, so that the seta resem-
bles a grater; length of a well developed right
penial seta 1mm (H). Genital setae present at the
tumescences in X or X and XI, each with the usual
diplotreman appearance, a stout seta with four
opposed longitudinal series of long notches and
the tip slightly expanded below the terminal
point; the seta gently curved; the longitudinal
notches confined to approximately the ectal third;
length right seta of IX (measured in a straight line
from tip to base) = 0.9mm; greatest width, near
base, 60j.m. Spermathecae two pairs, transposed

FIG. 4. Diplotrema acropetra sp. nov. Holotype,
QMG212027. A-D, Tip and successively more pos-
terior regions of a penial seta of XVII.

EARTHWORMS FROM CAPE YORK PENINSULA

posteriorly one segment, relative to the usual
megascolecid condition, so that the posterior pair
is in X, the anterior pair in IX. The posterior pair
of spermathecae, in X, very much larger than the
anterior pair; each posterior spermatheca with a
sacciform ampulla, a short narrow duct; and a
large multiloculate and apically lobed diverticu-
lum, containing sperm bundles, which joins the
junction of ampulla and duct and is elongated at
approximately a right angle to the duct. Length
right spermatheca of X = 1.5mm; length ampulla
= 1.0mm; ratio length spermatheca: length duct
(including base of diverticulum) = 3.5; length of
diverticulum lateral of duct = 0.6mm, The
diverticula of the anterior spermathecae are small
‘rosettes’ of loculi.

ETYMOLOGY. From the Greek acros, peak, and
petros, å rock.

REMARKS. Transposition of the spermathecae,
from the usual position in VIII and IX to IX and
X, is unknown elsewhere in Diplotrema. This
correlates with the unusual slender condition of
the commissural blood vessels, and suppression
of testes and funnels in X, giving the metandric
condition. As the prostatic porophores are in their
normal segments of XVII and XIX, it is clearly
not due to interpolation of a segment, às occurs
in some species of the closely relàted genus
Acanthodrilus, in New Caledonia (Jamieson &
Bennett, 1979),

Rocky Peak is an isolated upland plateau of
Battle Camp sandstone, adjacent to the Deighton
Tableland, The swampy habitat is the most north-
erly location of a habitat type dominated by Bank-
sia robur.

Diplotrema attenuata sp. nov. (Figs 5-8; 39)

TYPE LOCALITY. Peach Creek, McIlwraith Range,
13744 17"S. 143720' 15 £5"E, altitude 500-520m in
bank of stream in notophyll vine forest, on Kintore
adamallite granite, K.R. McDonald, À.J. Stewart, W.E.
Martin, 26 Sep 1995 and, KR Mc., A.J. S., 23 Sep
1996.

MATERIAL EXAMINED, HOLOTY PE, QMG21 2000,
(Includes microscope slides. of left and right anterior
penial seta and left genital setae of VIII).
PARATYPES. 1-8 QMG212038-45; P9 QMG-
212046; P10-12 QMG212019-212021; P13-17
QMG213386-213390; P18-20 QMG211972, P21-23
QMG213402-213404, OTHER MATERIAL. Several
not designaled types QMG211980, 212022.

DESCRIPTION. Length 106-163 (a live speci-
men 230) mm, H 115mm. Width (midclitellar)

237

KU

XIV

prostatic
porophore 1

Suz?

prostatic
porophore 2

FIG. 5. Diplotrema attenuata sp. nov. Holotype,
QMG212000. Ventral view of forebody and clitellar
region.

2.0- (H) 2.5mm. Segments 140-256, H 155. Uni-
formly circular in cross-section throughout.
Pigmentless buff in alcohol, clitellum, when
tumid, pale pink; in life, blood red throughout.
Prostomium small, prolobous, slightly indenting
the peristomium which is about as long as seg-
ment II. Dorsal pores absent. Setae 8 per segment,
commencing on II; ventral setal couples absent
from XVIII; those of XVII and XIX modified as
enlarged penial setae; genital setae present in
VIII, with the usual diplotreman ornamentation
consisting of notches. aa: ab: bc: cd: dd = 4.8:
1.0: 4.3: 0.8: 11.6; = 16.8: 3.5: 15.1: 17.9: 40.5%.
Nephropores not visible. Clitellum well-devel-
oped, saddle-shaped, extending over 1/2XII-
XVII, with a midventral gap that extends to, or
slightly beyond, b lines. Male pores not visible.
Prostatic pores 2 pairs, in XVII and XIX, each
pore on a minute oval papilla which is equatorial
and slightly lateral of b lines relative to adjacent
segments; the papillae of a side linked by a
weakly developed slightly parenthetic seminal
groove; the entire male genital area forming a
raised, almost square area, slightly longer than
wide, with rounded vertices. Genital markings: a
broad, unpaired midventral pad extending later-
ally beyond P lines, intersegmental in 10/11 (P1,
P3, 4, 6-9), 11/12 (P8), 12/13 (H, P2), 13/14 (H,
P2), 14/15 (P2) but extending almost to the setal
arcs of each adjacent segment; each pad bearing
a pair of approximately hemispheroidal papillae
median of a lines; highly diagnostic is a similar
pair of papillae, on a less distinct transversely
oval area, segmentally situated on XVIII (all
specimens); a transverse segmental pad fre-
quently present in XX (P1, 3, 6-9). Further mark-
ings a midventral postsetal triangular slight
tumescence in VII and VIII (H, P2, 6, 7, 9). An
unpaired oval-rectangular genital tumescence on
the left side straddling and extending beyond ab,
in the anterior half of VIII, with two punctuations
representing genital setae. Female pores small
transverse slits presetally in a (P1) or b (H) lines.
Spermathecal pores 2 pairs, in 7/8 and 8/9,
slightly lateral of setae b; inconspicuous but def-
inite orifices visible by parting the intersegment
(H, P1-9).

Septa 5/6-8/9 strongly thickened, the last two
the thickest. Dorsal blood vessel single, continu-
ous onto the pharynx. Last hearts in XIII; those
in X-XIII, latero-oesophageal, with connectives
to the dorsal and supraoesophageal vessels; com-
missurals in IX anteriorly slender, not heart-like,
though in IX, at least, possibly latero-oesopha-
geal. Gizzard small, compressible though with

MEMOIRS OF THE QUEENSLAND MUSEUM

prostomium
peristomium

prostatic pore 1
100%

prostatic pore 2
60%

FIG. 6. Diplotrema attenuata sp. nov. Diagram show-
ing distribution of genital markings in the holotype
and 9 paratypes,

some muscular sheen, almost vestigial, in V. Oe-
sophagus suppressed in VI by backward exten-
sion of septum 5/6; segmentally slightly swollen,
and vascularized in VII to IX; a simple tube from
X(?), XI to 1/2XIX in which the intestine com-
mences with abrupt expansion; typhlosole ab-
sent. Ventrolateral masses in XIII give the
spurious impression of calciferous glands but are
not connected to the oesophagus and disintegrate
on manipulation. Holonephric throughout;
nephridia not seen in a few anterior segments and
no tufting detected; nephridia with preseptal fun-
nel near nerve cord and slender duct discharging
in line with the dorsal setal couples (cd); caudally
the body of the nephridium has the appearance of
a convoluted sac but bladders are absent. Holan-
dric; large sperm masses, and very large irides-
cent sperm funnels free in X and XI. Two pairs

EARTHWORMS FROM CAPE YORK PENINSULA

prostate
duct

prostate duot

diverticulum

spermatheva!
ampulla

C = 1mm P D

FIG. 7. Diplotrema avenuata sp. nov. Holotype,
QMG212000. A, dorsal view in region of prosto-
mium. B, right prostates, with penial setae omitted, C
and D, ventral and dorsal views of the left sperma-
theca of IX.

239

of large, racemose seminal vesicles which fill the
length of their segments, in XI and. XII. Moder-
ately large, flattened bushy ovaries, and funnels,
in XIII; small, multichambered, berry-like
masses on each side of the oesophagus in XIV are
presumably ovisacs. Prostates two pairs, with
ducts discharging in XVII and XIX; those in XVII
considerably the larger and with the form of tor-
tuous thick tubes which end in XVIII; those in
XIX flattened and strap-like in their midregion,
and approximately S-shaped; each prostate with
ashort slender duct which is obscured by the large
penisetal follicles. Male genital field of XVII-
XIX represented internally by diffuse glandular
modification of the body wall but no bursae pres-
ent. Paired, conjoined vasa deferentia with
closely adpressed bends, and spermatozoal iri-
descence, traced on each side to immediately
behind the ectal end of the duct of the anterior
prostate, but termination not seen. Pemal setae
strongly curved throughout their lengths; the tip
not widened, reflected dorsally or not and ending
bluntly in a rounded swelling with a pitted sur-
face; sculpturing (best developed in the midre-
gion) consisting of pointed scales, contiguous
side by side, in incomplete circlets; the circlets
spaced longitudinally at approximately 154m in-
tervals; the scales stopping short of the modified
tip near which they are in small groups rather than
circlets; length of a well developed right penial
seta 1.3mm (P10). Genital setae present at the
tumescences in VIII; each with the usual diplotre-
man appearance, a stoul seta with four opposed
longitudinal series of long notches and the tip
slightly expanded below the terminal point;
length left seta 20.5mm (incomplete?); greatest
width, near base, = 25jum. Spermathecae 2 pairs,
the posterior somewhat the larger, each with a
large irregularly ovoid ampulla and straight
broad, well demarcated muscular duct of about
half its length; a multiloculate diverticulum. with
a form reminiscent of a clenched fist, sessile
dorsally on the duct at the junction of the latter
with the ampulla, the few main loculi being lat-
erally situated and filled with innumerable very
small, iridescent sperm balls; length left sperma-
theca of IX = 2.3mm; length ampulla = 1.4mm;
ratio length spermatheca; length duct= 2.6; great-
est dimension of diverticulum (transversely) —
0.8mm (H).

ETYMOLOGY. Named for its unusually attenuated
form.

REMARKS. The genital field, particularly the
two knob-like penital markings median to the

240 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Diplotrema aitenuata sp. nov. Paratype 10, QMG212019. Scanning electron micrographs. A, a right
penial seta of XVII. B and C, a right penial seta of XIX. D-G, appearance of seta shown in A from the tip to
near the base. H-K, same for seta shown in B and C.

EARTHWORMS FROM CAPE YORK PENINSULA

seminal grooves on segment XVIII, is diagnostic
of D. attenuata, The male genital field of this
species, when fixed, is emphasized by the fact
that the body is almost always dorsally arched
anteriorly and posteriorly to it, The body is also
contorted elsewhere.

The absence of dorsal pores is altributable to an
amphibious existence im earthworms. It is un-
known in other published species of Diplotrema
with the exception of D cornigravei
(Michaelsen, 1907) and possibly D. macleayi
(Fletcher, 1890), both inadequately described
species from Western and North Western Aus-
tralia, respectively.

Diplotrema scheltingai sp. nov.
(Figs 9-11; 39)

TYPE LOCALITY. 13*44'17"S 143920" 15"E, Peach
Creek, Mellwraith Range, altitude 500-520m. In moist
upper root horizon down to about 25cm, notoph yll vine
forest with fan and feather palms, stream bank, sandy
loam, on Kintore adamallite granite. K.R. McDonald,
A.J, Stewart, 25-27 Sep 1996.

MATERIAL EXAMINED. HOLOTYPE. QMG212001
(includes microscope slide of left genital seta of VIII).
PARATYPES. P] & 2 QM G212003-212004; P3 QM
G211981; P4 QM G213401; P5 QM G213406.

DESCRIPTION. Length 74 (H)-104 (P3)mm.
Width (midclitellar) 5mm (H, P3). Segments 188
(P3)-204 (H). Uniformly circular in cross-section
throughout, pigmentless buff in ethanol, includ-
ing clitellum. Prostomium prolobous but deeply
indenting the penstomium dorsally; peristomium
langer than segment II; first dorsal pore in 11/12
but not definitely perforate until 17/18. Setae 8
per segment, commencing on II; in XII, aa: ab:
be: cd: dd 2 5.0: 1.0: 4.8: 1.0: 21.4; or 12.5; 2.5:
12.1: 2.4: 53.5% ; ventral setal couples of XVIII
absent; those of X VII and XIX modified as en-
larged penal setae; setae a and b of VIII forming
genital setae. Nephropores visible in the
posiclitellar body, a pair in each segment, each
pore a faint spot anterior to each seta b. Clitellum
well-developed, extending over XI-XVII, inter-
rupted ventrally by the male field in XVII, with
doubtful midventral development anterior to the
field (H), or interrupted between the ventral setal
couples throughout, i.e., saddle-shaped (P1-3).
Male pores not visible. Prostatic pores 2 pairs, in
XVII and XIX, each pore on à minute oval papilla
which is in line with the ventral setal couple (ab)
of adjacent segments; the papillae of a side linked
by a parenthetic seminal groove; the pore accom-
panied by one or more minute punctuations pre-

241

B imm

FIG. 9. Diplotrema scheltingai sp. nov. Holotype.
QMG212001. A, dorsal view in region of prosto-
mium. B, ventral view of forebody and clitellar re-
gion,

sumably representing penial setae, but the latter
not protuberant; the posterior end of each seminal
proove continuous onto a small transversely el-
liptical papilla which is centred lateral to b lines
at the posterior limit of XIX; the male genital
area depressed relative to the anteriorly bounding
clitellum. Genital markings: a pair of rounded
papillae in ab of XVIII, filling the segment lon-
gitudinally, the summit of each with a pore-like
marking which from internal examination appear
to be the male pore. The two papillae connected

by å prominent transverse bår
which is widened midventrally so

as lo impinge strongly on segments
XVII and XIX, the widened region
bearing à median ellipucal protu-
berance (H, P1-3). Further genital
markings a small transversely el-
liptical pad anterior to the prostate A
pores of XVII (H, Pl) and asimilar
pad in XX with tumid lateral exten-
sions which include the ventral
setal couples (H, Pl, P3). A
strongly protuberant genital tu-
mescence in VII fills, and expands
the segment longitudinally, and
takes in seta c on each side (H,
P1-3); genital setae present. Fe-
male pores a pair of small trans-
verse slits, each with narraw
border, presetall y slightly lateral of
a lines of XIV (H). Spermathecal
pores 2 pairs, in 7/8 and 8/9, in or
very slightly lateral of setae a; in-
conspicuous but definite orifices,
closely apposed to the genital tu- B
mescence,

Septa 7/8-12/13 strongly thick-
ened; 9/10-11/12 slightly thick-
ened. Dorsal blood vessel single,
continuous onto the pharynx, Last
hearts in XII; those in X-XIII,
latero-oesophageal, with connectives. to the dor-
sal and supraoesophageal vessels, Gizzard very
large, an elongate, glossy, muscular, cylinder; in
V. Oesophagus lacking calciferous glands; but
very wide and vascular, without intersegimental
constriction, in XVI and XVII (H, P1). Intestine
commencing, with abrupt expansion. in XX; a
very large dorsal typhlosole commencing in
XNXIT-XXI, consisting of two distinct laminae
(H, P1). Halonephric, a large nephridium present
on each side throughout but in caudal segments
several longitudinal zigzagged ducts are present
on each side running from one segment to the
next; tulting absent. Holandric; testes, large
sperm masses, and very large iridescent sperm
funnels free in X and XI. Seminal vesicles race-
mose, in IX and XU; similar in size in the two
segments (H, P1). Small ovaries, with few egg
strings, in XIII. Somewhat flattened
tubuloracemose prostates, two pairs, in X VII and
XIX, restricted to these segments; those in XIX
considerably larger than those in XVII (H), or the
anterior pair slightly larger (P1), each gland
folded on itself at least twice, and, especially

prastate duct

Spermathecal diverticulum

MEMOIRS OF THE QUEENSLAND MUSEUM

prostate gland

spermathécal
diverticulum

FIG. 10. Diplatrema scheltingai sp. nov. Holotype, QMG212001. A,
right prostate of XIX. B, approximately dorsal and ventral views of
right spermatheca of IX,

ectally, incised, so as to appear racemose; a nar-
row central lumen present but the surrounding
glandular tissue very thick; the duct moderately
long, and slender, medianly directed, and accom-
panied by two large follicles of penial setae (H,
P1). The duct of each anterior prostate (examined
closely in the holotype) discharges at a large
bursa median to which is as smaller bursa associ-
ated with the penisetal follicles, The larger bursa
overlies a more posteromedian smaller bursa, in
XVIII, into which discharge the two thick, con-
joined vasa deferentia of its side, these curving
medianly in a wide arc. Thé posterior prostate
ducts do not appear to terminate at bursae but
there are internal protrusions of the body wall
near the point of entry of the penisetal follicles
into tbe body wall (H). Penial setae curved
through from 30° to 180°, the up tapering to a
smooth, simple point and bent to å varying extent;
sculpturing consisting of many unevenly spaced

FIG. 11. Diplotrema scheltingai sp. nov. Holotype,
QMG212001. Scanning electron micrographs. A-C,
iwo right penial setae of XIX. D, Tip of C, E-H,
appearance of seta shown in Å and B from the tip to
near the base,

EARTHWORMS FROM CAPE YORK PENINSULA 243

groups each of several contiguous pointed teeth;
near the tips the groups are extensive cir-
cumferentially at fairly regular longitudinal inter-
vals of approximately 1. Zum (H). Genital setae,
in VIII, gently curved but a short basal region
more strongly curved in the same direction; more
slender relative to length than usual for
Diplotrema but with the usual longitudinal notch-
ing and expansion of the tip below the terminal
point; length (straightened) = 1.8mm; greatest
width, near base, = 51um (H). Spermathecae 2
pairs, in VIII, and IX, the posterior pair slightly
thelarger; each with an ovoid ampulla and longer,
slender fairly well demarcated duct which is
joined near its swollen ectal end by a large, ellip-
soidal, multiloculate diverticulum containing
many minute iridescent balls of sperm; length left
spermatheca of VIII = 2.6mm; length ampulla =
1.1mm; ratio length spermatheca: length duct =
1.7; greatest dimension of diverticulum = 0.9mm.

ETYMOLOGY. Named for my colleague and friend,
David Scheltinga.

REMARKS. The tripartite transverse bar on seg-
ment XVIII is distinctive of D. scheltingai. The
fact that the nephridia, though apparently only
one pair per segment (preservation being inade-
quate for certain determination of their condi-
tion), have multiple longitudinal ducts possibly
merits placement of this species in
Neodiplotrema. The validity of separating
Neodiplotrema trom Diplotrema is uncertain as it
is possible that the meronephric condition of the
latter taxon has evolved more than once from the
holonephric Diplotrema condition.

Genus Neodiplotrema Dyne, 1996

The definition of Neodiplotrema as recently
defined by Dyne (1996) differs from that of
Diplotrema only in the description of the excre-
tory system: *Meronephric, avesiculate; anterior
tufted nephridia present’. It is shown below that
tufted nephridia may be absent.

The three new species, Neodiplotrema al-
tanmoui, N. mcdonaldi and N. paripunctata, rep-
resent a significant extension of the known range
of the genus to the east of the northern continua-
tion of the Great Dividing Range. Previously the
genus was known only from monsoonal semi-
deciduous vine forests in the Lockerbie, Iron
Range and Weipa areas, and Thursday Island
(Dyne, 1996), that is western or northern flowing
catchments at the tip of the Cape York Peninsula.

MEMOIRS OF THE QUEENSLAND MUSEUM

spermathecal
pore 1

genital
marking

female pore

genital
markings

prostatic
pore 1
seminal
groove
prostatic
pore 2

FIG. 12. Neodiplotrema altanmoui sp. nov. Holotype,
QMG211986. Ventral view of forebody and clitellar
region,

In the newly demonstrated localities the genus is
sympatric with Diplotrema.

Neodiplotrema altanmoui sp. nov.
(Figs 12-16; 39)

TYPE-LOCALITY. 14?33'31"S 144?38'08"E, Al-
tanmoui Section, Cape Melville National Park, alt.
560-570m, vine forest on sandstone escarpment, under
logs and rocks on forest floor, K. McDonald, P.J.
Lethbridge, 8 Apr 1995. Formalin fixation.

EARTHWORMS FROM CAPE YORK PENINSULA

genital
marking

genital
markings

prostatic
porophore 1
seminal

porophore 2

FIG. 13. Neodiplotremaaltanmoui sp. nov. Paratype 3,
QMG211987. Ventral view of forebody and clitellar
region.

MATERIALEXAMINED. HOLOTYPE. QMG21 1986.
PARATYPES. PI and 2 QMG212023.

Same data, in mesophyll vine forest, under
rock, Paratype 3 QMG211987,

DESCRIPTION. Length 140mm. Width (mid-
clitellar) 7.6mm. Segments 258 (H), Circular in
cross-section, pigmentless in alcohol. Prosto-
mium wide and prolobous but a suggestion of a
narrow continuation onto the much fissured peri-
stomium, a middorsal groove being wider than
the others (H, P3). First dorsal pore 12/13 (H, P3).
Setae 8 per segment, in regular longitudinal rows
throughout; ventral sera couples of XVIII pres-
ent; those of XVII and XIX modified as enlarged
penial setae; no genital setae (H, P3); in XII, aa:

245
prostate duct

penisetal
follicle

prostate

gland

A 1mm
j spermathecal
diverticulum

spermathecal
ampulla

FIG, 14. Neodiplorrema altanmoui sp. nov. Paratype I,
QMG212023. A, right prostate of XIX. B, right sper-
matheca of IX,

ab; bc: cd: dd = 6.3: 1.0: 7,1: 1.0: 48,3; or 8.7:
1.4: 9,8: 1.4: 66.4% (H). Nephropores sporadi-
cally visible as a transverse series of minute ovals
anteriorly in their segments. Clitellum saddle-
shaped, protuberant, over XHI-XVIII; ventral
margins well above b lines (H, P3). Prostatic
pores 2 pairs, in XVII and XIX. coincident with
the penial seta orifices, in ab, on small subeircular

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 15. Neodiplotrema altanmoui sp. nov. Holotype, QMG21 1986. Scanning electron micrographs. A, right
penial setae of XVII. B, ectal region of a penial seta. C, detail of sculpturing of same. D, penial setae of XIX.
E. ectal region of a penial seta of XIX. F, detail of sculpturing of same.

porophores; seminal grooves, joining the pros-
tatic porophores of a side, forming parentheses
with their midpoint, and presumably the male
pores, lateral of b lines a the equator of XVIII (H,
P3), the grooves deeply incised in P3, faint
(owing to maceration?) in H; the anterior prostate
pores 2.3mm, ca. 0.1 body circumference apart
(H). Genital markings indefinite in the macerated
holotype. whitish somewhat keratinized-looking
transverse unpaired midventral ridges in interseg-
ments 13/14, 14/15 and 15/16(the most pro-
nounced), XX and XXI, including the ventral
setal couples; clearly visible in the better pre-
served P3 in which they consist of a narrow
ventral band or pad posterior in each of VIII and
X and extending well laterally of & lines, with
their anterior borders at but not including the setal

arc; that in VIII posteriorly abuts closely on the
spermathecal pores of 8/9; other markings two
much larger but transversely less extensive pads
one at each of intersegments 14/15 and 15/16
extending shortly lateral of b lines, each abutting
the ventral setal couples of the previous segment
but posteriorly not reaching the following setal
arc. Female pores minute transverse slits, each
with a crescentic anterior border, immediately
anterior to the ventral setal couples, almost in b
lines (H). Spermathecal pores 2 pairs, in 7/8 and
8/9, in ab, definite pores filled with coagulum and
with anterior and/or posterior crescentic lips;
those of each pair conjoined by a narrow band
medianly,

Septa 5/6-10/11, especially 8/9 and 9/10,
strongly thickened. Dorsal blood vessel single,

EARTHWORMS FROM CAPE YORK PENINSULA

continuous onto the pharynx; last hearts in XIII;
those in X-XII latero-oesophageal, with well
developed connectives to the supraoesophageal
vessel and much thinner connectives to the dorsal
vessel; commissurals in IX and anteriorly slender
with dorsal connectives only. Gizzard large,
stoutly cylindrical, firm and muscular in V. Oe-
sophagus lacking calciferous glands; transition
from oesophagus to intestine indefinite in terms
of appearance of the gut wall and presence of fine,
silty contents, but not widening until XX (P2, 3);
a low, narrow dorsal typhlosole commencing be-
hind the prostates (P1, 3), Meronephric through-
out, nephridia commencing in Il; very
conspicuous, profusely divided tufts present in II
and IV, very large and restricted to IV in P3, a
thick duct of each tuft running anteriorly, to the
wall of the buccal cavity. In the midbody, shortly
behind the prostates, approximately 8 astomate
micromeronephridia on each side and a median,
not especially enlarged but more convoluted,
meronephridium for which a preseptal funnel was
demonstrated sporadically. Caudally (P2). the
medianmost nephridium is greatly enlarged rela-
tive to those lying more laterally and has a con-
spicuous preseptal funnel. Holandric; seminal
funnels largein X and XI, only those in XI with
spermatazoal masses and iridescence and accom-
panied on Ihe anlerior septum by extensive
masses of rounded testicular follicles; in X, fun-
nels lacking spermatozoal iridescence and testes
not recognizable (H, P1) but in P2 and P3 there
are sperm masses, and funnels are iridescent, in
X and XI. Racemose seminal vesicles in IX and
XII, those in XII the larger and visible dorsally
(H, P3). Bushy ovaries with few oocytes, in XIII
(P2) or large, undulating laminae with many oo-
cytes (P3), Prostatic glands two pairs of simple,
much convoluted tubular organs; both pairs ex-
tending back into XXV; the anterior pair some-
what longer than the posterior pair but both pairs
well developed; or the anterior pair in XVII and
XVIII, the posterior pair in XIX and XX (P3);
each with a short slender, slightly muscular duct
(P1); or the glands restricted to segments XVII
and XIX (H). Penial setae with two posteriorly
joined muscular follicles on each side, each con-
taining two setae, presumably corresponding to d
and P follicles, at each prostate duct; length of à
well developed setae (measured in a straight line
from base to tip) 2 1.9mm. The setae bowed

FIG. 16. Neodiplotrema allanmoui sp. nov. Right an-
terior penial seta of paratype 3, QMG211987. A, tip
of seta. B-D, progressively more posterior regions of
the anterior ornamentation,

D)

248 MEMOIRS OF THE QUEENSLAND MUSEUM

through varying degrees, maximally about 90°;
the ectal region of the shaft ornamented with
circumferential rows of single- or multiple-
pointed ectally directed teeth; the tooth rows
closely spaced along the seta, approximately
IOjum apart; cross section of the seta circular
excepting at the tip which forms an elongate
spatula, only slightly concave, with thickened
margins, and a knoblike, pointed tip; scattered
teeth continuing into the spatulate region; the
latter sometimes bent ‘dorsoventrally’, possibly
because of compression during copulation. No
notable difference in form observed between
setae of XVII and XIX (P2); setae of P3 closely
similar bit with more protuberant thorn-like
scales. Spermathecae 2 pairs, in VII and IX, each
organ consisting of a rounded diverticulum ses-
sile on the body wall and obscuring beneath it all
but a small ental region of the wide duct and a
large saclike ampulla which in the holotype has a
series of encircling annuli or pliae; the margins of
the diverticulum irregularly and indistinctly lobed
owing to many internal sperm chambers (H, P1, 3);
in IX, length right spermatheca = 3.3mm; length
ampulla 2.0mm; ratio length spermatheca: length
duct (with superior diverticulum) = 2.5 (P1).

ETYMOLOGY. After the type-locality.

REMARKS. The N. altanmoui specimens.were
located in moist mesophyll vine forest on the
Jurassic Dalrymple sandstone and moist soil
where the sandstone escarpment boundary met
the Permian Altanmoui granites. It occurs at the
headwaters of Wakooka Creek, an eastwards
flowing coastal stream arising in the Altanmoui
Range. The feather palm forest along drainage
lines was dominated by Archontophoenix sp. and
at an altitude of 540 metres is the highest of this
vegetation type in the Laura basin (Cooktown to
Princess Charlotte Bay) by a large margin (K.R.
McDonald citing J.P. Stanton pers. comm.).

N. altanmoui resembles the sympatric N.
paripunctata and N. exigua Dyne, from
Lockerbie East, and differs from all other known
species of the genus in having seminal vesicles in
segments IX (in addition to those in XII). It differs
from N. exigua in the lobed multiloculate sperma-
thecal diverticulum whereas the latter species has
a sacciform, blunt diverticulum and in the more
anterior origin of the intestine. The male genital
field was not developed in N. exigua. N. al-

FIG. 17. Neodiplotrema mcdonaldi sp. nov. Holotype,
QMG212005. Ventral view of forebody and clitellar
region.

marking

XIII

XIV female pore

prostatic
porophore 1

porophore 2

genital
marking

1 mm

EARTHWORMS FROM CAPE YORK PENINSULA

tanmoui further resembles N. paripunctata in the
form of the spermathecae, with a rosette-like ar-
rangement of loculi of the spermathecal diverticu-
lum on the short spermathecal duct. Differences
from the latter species are numerous and include the
distinctive setal ratios, the strong denticulation of
the penial setae, and the absence of the paired,
apposed genital markings posteriorly in XV.

Neodiplotrema medonaldi sp. nov.
(Figs 17-19; 39)

TYPE LOCALITY. 13°44°17"S 143920'15"E +5",
Peach Creek, Mcllwraith Range, alt. 500-520m. In
creek bank, in pig diggings and/or in moist upper root
horizon, Notophyll vine forest with fan and feather
palms, on Kintore adamallite granite, K.R. McDonald,
A.J. Stewart, 23-27 Sep 1996.

MATERIAL EXAMINED. HOLOTYPE. QMG21 2005.
PARATYPES. P1-4 QMG212006-212009; P5
QMG212010 (includes microscope slide of left poste-
rior penal seta); P6 & 7 QMG21201 1-212012; PR QM
212013: P9-12 QMG212014-17; P13 QMG?212018.

DESCRIPTION. Length 91-134mm, H 112mm.
Width (midclitellar) 4.6-5.0mm, H 4.9mm. Seg-
ments 139-264, H 210. Circular in cross-section
throughout. Pigmentless pale brownish grey in
ethanol. Prostomium more than half the width of
the peristomium; prolobous; first dorsal pore in
10/11 (H)or 12/13 (P2); peristomium shorter than
segment II. Setae 8 per segment, commencing on
Il, in XII, aa: ab: bc: cd: dd = 8.3: 1.0: 9.5: 1.2:
22.6; or 15.2: 1.8: 17.5: 2.3: 41.6% (H). Ventral
setal couples absent from XVIII; those of XVH
and XIX modified as enlarged penial setae; gen-
ital setàe not demonstrable. Nephropores not rec-
ognizable. Clitellum well-developed,
saddle-shaped, extending over XIII-XIX, with a
mid-ventral gap that extends to, or slightly lateral
of, ab lines. Male pores not visible. Prostatic
pores 2 pairs, in XVII and XIX, each pore on å
strongly protuberant oval papilla which is equa-
torial in line with the ventral setal couples (ab)
relative to adjacent segments; the papillae of å
side linked by a clearly developed narrow semi-
nal groove which is bent medially (not forming
parentheses) and is transversely incised by inter-
segmental furrows 17.18 and 18/19 and an in-
trasegmental groove of XVIII; the entire male
genital area wider than long and not depressed.
Genital markings: a broad, unpaired midventral
pad extending laterally well beyond b lines, inter-
segmental in 10/11 and 14/15 and 21/22 but ex-
tending almost to the setal arcs of each adjacent
segment; each pad traversed by a groove corre-

prostomium

~~ spermalhacal
M diveniculum

FIG. 18. Neodiplotrema mcdonaldi sp. nov. Holotype,
QMG212005, A, dorsal view in region of prosto-
mium. B, right prostate of XVII, with penial setae
omitted. C. right spermatheca of IX.

sponding with the intersegmental furrow. Further
markings a suggestion of a small papillae
posteromedian io the prostatic porophores of
XVII. Genital tumescences absent. Female pores,
with elliptical borders, a pair near the anterior
margin of XIV, in ab. Spermathecal pores 2 pairs,
in 7/8 and 8/9, in or slightly lateral of setae b; each
on à conspicuous elliptical papilla which is wider
than à setal couple.

Septa 6/7-9/10 strongly thickened. Dorsal
blood vessel single, continuous onto the pharynx.

250 MEMOIRS OF THE QUEENSLAND MUSEUM

Last hearts in XIII; those in XI-XIII large, with
conspicuous supraoesophageal and smaller dor-
sal connectives; those in X slender, though still
apparently laterooesophageal; in EX anteriorly
slender, with dorsal connective only. Gizzard
moderately large and firmly muscular, in V, wid-
ening anteriorly. Oesophagus with a pair of lateral
blood red dilatations, with many closely situated
circumferential vascular striae which correspond
with low internal lamellae, in each of 1/2XII-XV;
not pinched off from the oesophagus but true,
sessile, calciferous glands as indicated by white
granular contents which effervesce vigorously in
dilute acid whereas the contents of the adjacent
oesophagus and the intestine do not (H), Intestine
commencing in XVII; a very deep bilaminar
typhlosole commencing in XX (P5). No anterior
tufted nephridia present. At least four
micromeronephridia on each side in the intestinal
region; no funnels seen. Caudally with three or
four meronephndia on each side, of which at least
the median one has à preseptal funnel. Holandric;
large seminal funnels with spermatozoal irides-
cence in X and XI. Large racemose seminal ves-
icles a pair in each of XI and XII, each vesicle a
compucl mass, with a single connection to the
anterior septum of its segment, divided into many
small iridescent loculi. Dendritic ovaries with
many egg strings in XIV. Prostate glands flat-
tened very tortuous wide tubes investing the in-
testine, a pair 1n each of XVII and XIX; with à
narrow S-shaped duct the ental fourth of which is
transitional to the structure of the gland. Penisetal
follicles immensely long and conspicuous, the
posterior pair extending posterolaterally to join
the body wall in XXIV. Penial setae exceedingly
thin and filiform; a long ectal region ornamented
with many long slender sinuous scales directed
obliquely towards the tip and standing well up
from the surface; each scale itself consisting of
three or more parallel pointed strands; length of
a well developed right posterior penial setà
(straightened) =6.7mm, Genital setae not demon-
strable. Spermathecae two pairs, in VIII and IX,
the diverticulum (so identified by presence of
sperm chambers) a longue-shaped mass so large
as to be visible on opening the specimen dorsally,
resembling a seminal vesicle, its outer and termi-
nal aspect with numerous small protuberant

FIG. 19, Neodiplotrema mcdonaldi sp. nov. Holotype,
QMG212005. Scanning electron micrographs of a
right penial seta of XIX. A, seta, excepting base. B,
higher magnification, C-G, appearance of seta from
the tip to near the base.

EARTHWORMS FROM CAPE YORK PENINSULA

chambers containing sperm balls; the ampulla a
slightly smaller pointed sac appearing to be an
appendage of the diverticulum; the duct a short
narrow ectal continuation from the diverticulum,
Length right spermatheca of IX, trom pore to tip
of diverticulum = 3mm; ratio length spermatheca:
length duct = 17:1; length ampulla = 2mm.

ETYMOLOGY, Named for Keith McDonald who first
collected this species.

REMARKS. Neodiplotrema mcdonaldi is distin-
guished from all other species of the genus by the
large mammillated spermathecal diverticulum
which exceeds the length of the ampulla and by
the sinuous hair-like penial setae. A similar type
of spermatheca occurs in Diplotrema ridei and,
though less similar, in D. shandi. Close relation-
ship of N. mcdonaldi to D. ridei, from Cape York
and Melville Island, deserves consideration in
view of considerable similarities of their genital
fields.

Neodiplotrema paripunctata sp. nov.
(Figs 20-23; 39)

TYPE LOCALITY. 14°33°27"S 144*38'08"E, Al-
tanmoui Section, Cape Melville National Park, altitude
52010 m, in mesophyll vine forest, with feather palms,
along creek, located from pig diggings, K. McDonald,
P.J. Lethbridge, 7 & 10 Apr 1995.

MATERIAL EXAMINED, HOLOTY PE. QMG213360.
PARATYPE. P1 QMG213361.

DESCRIPTION. Length 150mm. Width (mid-
clitellar) 3.8mm, Segments 171 (H). Colour
brown in ethanol with a pale grey-hrown
clitellum. Prostomium prolobous. Peristomium
with anterior parallel grooves; approximately as
long as segment Il; not bisected ventrally. First
open dorsal pore 18/19, but occluded pores pres-
ent on the remaining, more anterior, segments of
the clitellum (H). Setae 8 per segment, in regular
longitudinal rows. throughout; ventral setal cou-
ples of XVIII present immediately median to the
seminal ridges; those of XVII and XIX modified
as enlarged penial setae; genital setae absent; in
XII, aa: ab: bc: ed: dd = 3.6: 1.0: 4.2: 1.0: 18.0;
or 10.6: 2,9: 12.4; 2.9: 52.9% (H). Nephropores
not visible. Clitellum saddle-shaped, protuberant,
XIIEXIX; ventral margins well above b lines.
Prostatic pores 2 pairs, in XVII and XIX, coinci-
dent with the protuberant penial setae, in line with
the ventral setal couples (ab) of other segments:
the pores on each side on small porophores which
are not distinguished from a tumid parenthetic
ridge which bears a simple seminal groove. A

markings

1mm

FIG. 20. Neodiploirema paripunctata sp. nov, Holo-
type, QMG213360, Ventral view of forebody and
clitellar region.

deep midventral depression present between the
seminal grooves. Male pares a pair of minute
orifices, on the seminal ridges, at the level ol

252

spermathecal
diverticulum

spermathecal [^ `
ampulla

FIG. 21. Neodiplotrema paripunctata sp. nov. Holo-
type, QMG213360. A, dorsal view in region of pro-
stomium. B, left prostates; penial setae omitted. C,
right spermatheca of IX, B and C same scale.

17/18 and shortly lateral of b lines. Female pores
in small oval fields anterior to the ventral setal
couples (ab) of XIV. Spermathecal pores 2 pairs,
a short distance posterior (relaxation artefact?) to
7/8 and 8/9, in b lines. Genital markings paired or

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 22. Neodiplotrema paripunctata sp. nov. Holo-
type. QMG213360. Scanning electron micrographs
of a ventral seta of VIII. A, seta, showing node typical
of somatic setae. B-D, appearance of seta from the tip
to near the base.

unilateral pale non-protuberant oval patches, bor-
dered by brown pigment, lateral of or just includ-
ing setae b, in IX, XI and XII (H, P1) and in X

EARTHWORMS FROM CAPE YORK PENINSULA

(P1); a pair of medianly contiguous papillae mid-
ventrally in 15/16 (H, P1); a midventral trans-
verse postsetal pad in XX (H, P1); a small
indistinct midventral oval pad in the posterior
half of XXI (H).

Septa 7/8-11/12 fairly strongly thickened. Dorsal
blood vessel single, continuous onto the pharynx;
last hearts in XIII; those in X-XIII latero-oesoph-
ageal, with supra-oesophageal connective larger
than the dorsal connective; commissurals in IX,
and anteriorly, slender and with dorsal connec-
tives only. Gizzard very large, cylindrical,
slightly widening anteriad, moderately firm and
muscular, in VI(?), i.e., anterior to the diaphanous
septum 6/7. Oesophagus lacking calciferous
glands. Intestine commencing in XIX, not reaching
full width until XXIV. A pair of moderate-sized,
conical anterolateral diverticula on the intestine
in XXVIII. No typhlosole found. Large tufted
nephridia in II and IV send composite ducts to the
wall of the buccal cavity and are accompanied
laterally by many micromeronephridia; in the
oesophageal region at least 12 micromeronephri-
dia on each side, apparently all astomate and
with no enlargement of the median pair; cau-
dally with 8 meronephridia on each side, the
medianmost one enlarged as a mega-
meronephridium with a preseptal funnel; the lat-
eral nephridia apparently lacking funnels (H).
Holandric; iridescent seminal funnels in X and
XI. Seminal vesicles in IX and XII; sacciform,
those in IX subdivided into two or more portions,
and accompanied by a row of small vesicles;
those in XII each forming an exceptionally large
lobulated sac. Ovaries not detectable. Two sub-
equal pairs of thickly tubular very tortuous pros-
tates in XVII and XIX, limited to their segments,
their surfaces minutely mammillated; the ectal
duct with a stout, muscular ectal region and a
shorter less muscular region, bent relative to this,
which is narrow at its origin from the gland. Vasa
deferentia not traceable. Penisetal follicles over-
lying the prostate ducts. Penial setae curved in an
arc, the tip strongly reflexed on itself or with a
scroll-like expansion; ornamentation, on the
shaft, consisting of weakly developed jagged
transverse lines, each encircling less than one
fourth of the circumference, and spaced longitu-
dinally at approximately lOjum intervals or
these jagged lines very few and scarcely visi-
ble; length of a well developed left penial seta
of XVII=2.5mm; greatest width approximately
35ju.m (H). Ventral setal couples of VIII and IX
unmodified. Spermathecae two pairs, in VIII
and IX; each with a large ovoid ampulla and a

253

short narrower but poorly demarcated duct
which is hidden except for its ectal end by a
rosette-shaped multiloculate diverticulum
which is incised slightly or deeply into ap-
proximately 8 large loculi in which many min-
ute iridescent sperm balls are visible. Length
right spermatheca of IX = 8.2mm; length am-
pulla 2 2.9; ration of length spermatheca:
length duct = 5.3; diameter of diverticulum
1.6mm (H).

ETYMOLOGY. paripunctata, referring to the paired
anterior oval patches.

REMARKS. The pair of closely apposed genital
markings in 15/16 is distinctive of N.
paripunctata. This and the absence of large den-
ticulations of the penial setae, is among features
distinguishing N. paripunctata from the sympat-
ric N. altanmoui. Intestinal diverticula are known
in pheretimoids, in the Megascolecinae, but have
not previously been observed in the
Acanthodrilinae.

Kayarmacia gen. nov.

DIAGNOSIS. A pair of combined male and pros-
tatic pores, associated with penial setae, on XVII.
Spermathecal pores 1 pair, in intersegmental fur-
row 7/8. Gizzard in V. Oesophagus lacking cal-
ciferous glands. Holonephric; nephridia
avesiculate. Holandric. Prostate a single pair of
tortuous tubes, each with an exceptionally long
tortuous muscular duct which opens in common
with the equally thick and muscular ectal end of
the vas deferens (ejaculatory duct). Sper-
mathecae a pair in VII; diverticulum, with nacre-
ous sperm masses.

DESCRIPTION. Small worms, less than 75mm
long. Dorsal pores (always?) present. Setae 8 per
segment, closely paired; the lateral couple (cd)
narrower than the ventral couple (ab). Clitellum
anterior to the male pores; annular. A pair of
combined male and prostatic pores, associated
with penial setae, on XVII. Genital markings
present in the vicinity of the male pores. Genital
tumescence and modified ventral setae present or
absent in the spermathecal region. Female pores
presetal in XIV. Spermathecal pores 1 pair, in
intersegmental furrow 7/8.

Dorsal blood vessel single. Last hearts in XIII,
those in X-XIII large, heart-like and latero-
oesophageal, the connective to the supra-
oesophageal vessel larger than that to the dorsal
vessel; commissurals in IX anteriorly slender,
with dorsal connective only. Gizzard large,

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 23. Neodiplotrema paripunctata sp. nov. Holotype, QMG213360. Scanning electron micrographs of a left
penial seta of XVII. A, seta. B-E, appearance of seta from the tip to near the base. F and G, ornamentation of

the posterior regions of a left penial seta of XIX.

sirongly muscular in V. Oesophagus lacking cal-
ciferous glands. Intestine commencing in XVII.
Holonephric; avesiculate ducts apparently dis-
charging in mid bc or c lines; true anterior tufting
not developed. Holandric. Racemose seminal
vesicles in IX and XII. Ovaries in XIII. Prostate
glands extending through several segments; a

single pair of tortuous tubes, each with an excep-
tionally long tortuous muscular duct which winds
forwards to open at the male pore in common with
the equally thick and muscular ectal end of the
vas deferens (ejaculatory duct). Spermathecae a
pair in VIII, opening at the anterior border; diver-
ticulum, with nacreous sperm masses.

EARTHWORMS FROM CAPE YORK PENINSULA

ETYMOLOGY. Kayarmacia, based on the name of the
volleciar. KR. McDonald. Gender masculine.

TYPE SPECIES. Kayarmacia adelphicus sp, nov.

OTHER SPECIES. Kayarmacia queenslandicus
(Michaelsen, 1916).

DISTRIBUTION. Cape York Peninsula.

REMARKS. K. adelphicus is chosen as the Type
species for Kayarmacia as identification ol K.
queenslandicus with Michaelsen's species,
though made with confidence, is subjective and
later changes lo its synonymy, though unlikely,
would cause confusion if were selected as the
lype species.

Michaelsen (1916) placed the Alice River spec-
imens in Ahedodrilus, features of this genus
being of location of male pores in segment XVII,
presence of a gizzard, and absence of nephridial
bladders. However, these are insufficient reasons
for considering R. gueenslandicus to be conge-
neric with the New Zealand type species of the
genus, Rhododrilus minutus Beddard 1889, as the
latter species has four pairs of spermathecal pores
and, more significantly, has the male and prostatic
pores located separately on X VIL. The mere pres-
ence of male and prostatic pores on XVII whether
combined or separate is not unique to one genus,
as 1L is seen inter alia in the acanthodriline genus
Microscolex and is the typical condition in the
Ocnerodrilinae and Eudrilidae. Nevertheless,
it is likely that location on XVII represents a
microscolecin reduction from an acanthodrilin
arrangement of male pores (male pores on
XVIII, prostatic pores on XVI and XIX) and
that Kayarmacia is referable to the subfamily
Acanthodrilinae and not to the Mega-
scolecinae.

The thickening of the distal ends of the vasa
deferentia and their opening in common with the
unusually Jong prostate ducts distinguishes
Kavarmacia from other megascolecid genera, In-
clusion in the Ocernodrilidac, å family in which
male and prostate ducts may be swollen, is pre-
cluded by several features, including the non-oc-
nerodriline condition of the oesophagus. Native
Ocernodrilidae are unknown from Australia,

Rhododrilus glandifera Jamieson, 1995, is ex-
cluded trom Kayarmacia by location of the sper-
mathecal pores in intersegmental furrow 8/9, not
7/8, and, more significantly, the absence of thick-
ening of the vasa deferentia, and the minute size
of the apparently separate prostate ducts. That R.
glandifera is congeneric with Rhododrilus is,
nevertheless, doubtful and placement in that

Ls
r

genus Must be regarded only ås provisional. 14
may well have closer affinities with Diplorrema
than does Kayarmacia,

Kavarmacia adelphicus sp. nov.
(Figs 24-27; 39)

TYPE LOCALITY. (1) 14*16'54"S 144727 ME,
Cape Melville National Park, along creek bank, cluset
canopy, altitude ca. | Om, KR McDonald, L.A. Jack-
son, 24 Feb 1995, (21 Same, 14°16'54"S 14427 MY E,
18 Feb 1995,

MATERIAL EXAMINED. HOLOTYPE, (1) QMG-
213378 ex21 1497 (part) (includes microscope slide of
male ducts), PARATYPES. (1) P1-5 QMG21 1497
(with several not designated types): (2) P6 & 7
QMG21 1495,

DESCRIPTION. Length 25-39mm, 28mm (HJ.
Width (midelitellar) 1.8-(H)2.6mm. Segments
102 (H). Pigmentless greyish buff in ethanol.
Pråstomium proepilobous (H) or epilobous 1/2,
Penstomium approximately as long as segment
IT. Dorsal pares sporadically visible from shortly
anterior to the clitellum (verified by exudition of
alcohol when specimen is placed in water). Sctac
closely paired; in XII, aa: ab: bo ed: dd = 82:
1.0: 7.7: 0.9: 18.1; or 18.1: 2.2: 16.9: 1.9: 40 (5.
Clitellun in XIII-anterior X VII; annular hut ven-
trally interrupted from the anterior third of XVI
posteriorly, to about mid be hy the male held;
setae and intersegmental furrows present on il. A
pair of large, globose but anteriorly pointed
strongly protuberant papillae each bearing the
combined male and prostatic pore on XVII in ab;
the walls of the papillae diaphanous and revealing
internally the large penial setae which support its
anterior protrusion (H), or the penial setae
strongly protuberant (P1, 2). Genital markings a
pair of indisünct swellings, al setae ab of XVI and
ANIL å midventral elliptical papillac postsetally
in XYII ar (P6) in XVL Genital tumescence and
modified ventral setae developed on the right side
in VIT CH) or paired in IX (P1, 2). Female pores
minute, immediately anterior to setae a of XIV.
Spermithecal pores 1 pair, in intersegmental fur-
row 7/8. shortly lateral of b; each a lateral slit in
an oval papilla.

Septa 8/9 and 9/10 the thickest, moderately
strongly thickened, Dorsal blood vessel single,
Last hearts in XII, those in X-XIII large, heart-
like and laterooesophageal, the connective tu the
supraoesophageal vessel larger than that to the
dorsal vessel; commussurals in IX anteriorly slen-
der, with dorsal connective only, Gizzard very
large, strongly muscular but compressible, ellip-

256

XIII

xiv £

prostatic pore

penial seta
(internal)

genital
marking

1mm

FIG. 24. Kayarmacia adelphicus gen. et sp. nov. Cape
Melville National Park, holotype, QMG213378. Ven-
tral view of forebody and clitellar region.

soidal though widening anteriorly, in V. Oesoph-
agus lacking calciferous glands. Intestine com-
mencing, with abrupt expansion, in XVII (H, P2).
Holonephric; nephridial bodies commencing in
IL avesiculate ducts apparently discharging in
mid bc; anterior tufting not developed. Holandric;
iridescent sperm funnels in X and XI. Large,
several lobed racemose seminal vesicles in IX
and XII (H, P2). Small ovaries, with few oocytes,
in XIII. Prostate glands visible externally through
the body wall in the holotype extending from
XVII to XXVIII; a single pair of tortuous tubes,
each with an exceptionally long tortuous muscu-

MEMOIRS OF THE QUEENSLAND MUSEUM

XIII

penial
setae

combined male
and prostatic
porophore

XXVIII

1mm

FIG. 25. Kayarmacia adelphicus gen. et sp. nov. Cape
Melville National Park, holotype, QMG213378. Lat-
eral view of male genital region, showing prostate
gland seen through body wall.

lar duct which winds forwards to open at the male
pore in common with the equally thick and mus-
cular ectal end of the vas deferens (ejaculatory
duct) (H, P2); the prostate duct and ejaculatory
duct pass separately but in contact to the tip of the
male porophore (confirmed in cleared mount).
Penial setae, only 1 on each side; stout, ectally
curved to form a hook; length 1.2mm, greatest,
basal, width ca. 70m; ornamentation consists of
droplet-like or many wart-like elliptical promi-
nences on the ‘dorsal’ and ‘ventral’ surfaces of the
curvature, the sides of the latter being smooth; on
the ventral surface, however, the prominences are
reduced to minute scattered but densely situated
points or cicatrices. Genital setae present on right

EARTHWORMS FROM CAPE YORK PENINSULA

spermathecal F^
diveniculum—p— =

spermathecal
ampulla

spermathecal
diverticulum

uaa)

widening of duct

FIG, 26. Kayarmacia adelphicus gen. et sp. nov. Cape Melville National

prostomium

B 1mm

sparmathecal
ampulla

t2
un
-

= 1.7mm; length ampulla =
0,88mm; ratio length sperma-
theca: length duct — 2.1; length
diverticulum = 1.0mm (H). In
P2, the swollen part of the duct
is approximately as large as the
ampulla.

ETYMOLOGY. adelphicus, from
the Greek Adelphos, brother, re-
ferring to its putative adelpho-
group relationship to K
queenslandicus.

REMARKS, Kayarmacia ad-
elphicus is morphologically
very close to the prior K.
queenslaridicus (g.v.) but geo-
graphical separation by the
Great Dividing Range suggests
that the presence in it of genital
setae, whereas these appear al-
ways to be absent from K.
queenslandicus, | indicates
vicariation of the two taxa from
a common ancestor. Strong
support for recognizing the two
as separate species is seen in
the very different ornamenta-
tion of the penial setae. It con-
sists of scattered droplet-like
prominences in K. adelphicus
contrasting with tooth rows
which ‘ventrally’ are united in
extensive transverse rows in K,

Park, holotype, QMG213378. A, left prostate and male duct, with penisetal
follicle. B, dorsal view in region of prostomium. C and D, two views of left
spermatheca. E, Paratype 2, QM G211497, left spermatheca. A. C-E same

queenslandicus, The distinc-
uve structure of the male and

scale.

in VII (H) or bilateral in VIII (P2); length of left
genital seta of VII, 0.76mm; shaft gently curved
but å short basal region strongly curved; tip
pointed, very slightly expanded beneath the
point; ornamentation four (?) longitudinal series,
each of about five large gouges or notches. Sper-
mathecae a pair in VIII, opening at the anterior
border; each with a large ovoid ampulla;the duct
of which consists of three regions in longitudinal
succession: an ental third which is approximately
tubular and is well demarcated from the ampulla,
a swollen intermediate region which receives the
diverticulum, and an ectal third which forms a
wide, short, muscular, tubular duct, Diverticulum
elongate ellipsoid, filled with nacreous sperm
masses but uniloculate, with à narrow connection
tothe spermathecal duct. Length left spermatheca

prostatic ducts seen in both
species is here regarded as a
generic character. The genital
setae, with longitudinal series of large gouges or
notches, are of the type seen also in Diplotrema
and the New Caledonian genus Acanthodrilus
and indicate relationship with these genera.

This species was found in humic sandy soil
under closed canopy on a creek bank near the
edge of the Cape Melville Altanmoui granite
boulder fields.

Kayarmacia queenslandicus (Michaelsen,
1916), New combination (Figs 28-30; 39)

Rhododrilus queenslandicus Michaelsen, 1916: 4-6,
Pl. 1. fig, 9-11.

MATERIAL EXAMINED. SYNTYPES. ca. 15°S.,
Cape York, banks of Alice River, a single much dis-

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 27. Kayarmacia adelphicus gen. et sp. nov. Cape Melville National Park. A, paratype 2, QMG211497, left
genital seta of VIII. B andC, tip and shaft of same. D, Holotype, QMG213378, right penial seta. E-G, appearance

of seta shown in D from the tip to near the base.

sected specimen labelled ‘Kap York, Mjöberg, Alice
River, Hamburg Museum V8487. NEW MATERIAL.
15?27'45"S 142?10' 12"E, Emu Lagoon, Alice-Mitch-
ell Rivers National Park, altitude 222m, top-soil on
sandy levee bank, open woodland, K.R. McDonald, 9
Feb 1993. ex QMAcc#2344, specimens 1-6 QM
G213385.

DESCRIPTION. Length 49-75mm. Width (mid-
clitellar) 1.2-1.9mm; greatest width (forebody)
2.3mm. Segments 124-143 (shortest and longest
specimens, S3 and 5). Pigmentless in ethanol;
clitellum greyish buff. Prostomium epilobous, ca.
1/4; small, broad and open. Peristomium approx-
imately as long as segment II. Dorsal pores spo-

EARTHWORMS FROM CAPE YORK PENINSULA

female pore

XVIII

XIX [-

i mm

FIG. 28. Kayarmacia queenslandicus (Michaelsen,
1916). Emu Lagoon, Alice-Mitchell Rivers National
Park, specimen QM Acc#2344, QMG213385. Ven-
tral view of forebody and clitellar region.

A tam

spermathecal
N diverticulum

spermathecal
ampulla

B imm

FIG. 29. Kayarmacia queenslandicus (Michaelsen,
1916). Emu Lagoon, Alice-Mitchell Rivers National
Park, specimen QM Acc#2344, QMG213385. A, dor-
sal view in region of prostomium. B, right sperma-
theca,

radically visible from shortly anterior to the
clitellum (verified by exudation of alcohol when
specimen is placed in water). Setae closely
paired; in XII, aa: ab: bc: cd: dd = 6.6: 1.0: 6.1:
0.7: 26.1; or 13.6: 2.1: 12.5: 1.5: 54.2. Clitellum
annular, XIII-XVT, setae normally developed.
Male genital field: a deep transversely elliptical
depression in XVII, preceded posteriorly in XVI
by a midventral, transverse glandular elevation,
and extending somewhat onto XVIII median to a
pair of strongly protuberant combined male and
prostatic porophores, the median borders of
which project ventromedially over the depres-
sion, the lateral borders of which are poorly dis-
tinguished from the lateral region of the segment;
from the summit of each papilla, in setal line a,
projects a penial seta; etc. An indistinct
hemispheroidal midventral genital markings

260

present presetally in XVIII; a larger indefinite
tumescence presetally in XIX (S1), or a
hemispheroidal midventral genital marking pres-
ent post-setally in XVIII. Female pores a pair of
deep punctuations very shortly anterior median to
seta a of XIV. Spermathecal pores 1 pair, in
intersegmental furrow 7/8, in or shortly lateral of
b; each with a large approximately ellipsoidal
epidermal elevation surrounding it which extends
well laterally of b and to or median of a and is
bounded anteriorly by a crescentic groove.
Septa 5/6 (S2), 6/7 (S1)-8/9 the strongest, mod-
erately strongly thickened. Dorsal blood vessel
single, continuous on to the pharynx. Last hearts
in XMI; those in X-XIII large and heart-like with
large connective to the supraoesophageal vessel
and slender connective to the dorsal vessel; those
in IX anteriorly slender, with dorsal connective
only. Gizzard large, strong, glossy, muscular
though compressible. Oesophagus lacking calcif-
erous glands; segmentally slightly swollen and
vascularized in VIII-XVI, especially vascular in
X-XIV (S1) or XI-XVI(S2). Intestine commenc-
ing in XVII but not sharply demarcated from the
oesophagus (S1, 2). Typhlosole absent.
Holonephric. Nephridia of II, III and especially
IV enlarged and much lobed though not truly
tufted; connection to the gut not detected (S2).
Other nephridia with avesiculate ducts discharg-
ing anterior to the lateral setal couples, specific-
ally in c (S1). Large male funnels in X and XI (S1,
2), with spermatozoal iridescence in S1. Seminal
vesicles in IX and XII, racemose, much divided,
those in XII the larger (S1); in IX only (immature)
in S2. Paddle-shaped ovaries in XIII. Prostates
tubular, extending from XVII to XXIV (SI, 2);
zigzagged in S1, almost straight and very slender
owing to immaturity in S2; at maturity (S1) the
duct extending from XVII- 1/2XIX, with an
abrupt bend in XIX; the short portion ental to the
bend not noticeably muscular, the remaining,
more ectal portion forming a wide, muscular,
glossy tube extending forwards to the male
porophore; vasa deferentia in XVI and XVII
forming a muscular tube of similar appearance
but slightly wider (ejaculatory duct), a short re-
gion ental to this, also in XVI, is more slender and
represents the posterior portion of the unmodified
more anterior region of the conjoined vasa defer-
entia. Penial seta ca. 0.96mm long ; only one on
each side; the tip curved in a hook as in K.
adelphicus (S2), extending from XVII into ante-
rior XIX (S1, 2); the curved apical portion or-
namented ‘dorsally’ and ‘ventrally’ by extensive
finely toothed transverse rows but the sides

MEMOIRS OF THE QUEENSLAND MUSEUM

smooth; the ventral tooth rows linked up trans-
versely so as to straddle much or the whole of the
ventral surface. Spermathecae a pair in VIII,
opening at the anterior border, not fully mature in
S2; in S1 each with a large ovoid ampulla;the duct
of which consists of three regions in longitudinal
succession: an ental third which is approximately
tubular and is well demarcated from the ampulla,
a swollen intermediate region which receives the
diverticulum, and an ectal third which forms a
wide, short, muscular, tubular duct. Diverticulum
elongate ellipsoid, with numerous nacreous
sperm masses which possibly represent non-pro-
tuberant sperm chambers, and with a narrow con-
nection to the spermathecal duct. Length left
spermatheca = 1.5mm; length ampulla = 0.9mm;
ratio length spermatheca: length duct = 2.5;
length diverticulum = 0.7mm.

REMARKS. The type locality is the Alice River,
which Michaelsen (1916) wrongly thought to be
a tributary of the Coleman River. Michaelsen’s
material was collected from the banks of the river
but he recognized its at least potentially
"limnische Lebensweise’. The stated absence of
dorsal pores would correlate with an amphibious
existence. The water table is at the surface and in
the rainy season most of the area, excepting the
levee banks, is covered by water (K. R. McDon-
ald, pers. comm.). However, in the new material
dorsal pores are sporadically demonstrable
though possibly vestigial.

Identification of the new material as
Rhododrilus (now Kayarmacia) queenslandicus
is based on close morphological similarity and
collection near the type locality. The identifica-
tion appears to be confirmed by the unusual or-
namentation of the penial setae in which
transverse rows of scales are present dorsally and
ventrally on the apical curvature but the sides are
smooth. The above account is derived solely from
the new material. Michaelsen's description of
Rhododrilus queenslandicus accords closely but
some points of disagreement require mention.
The clitellum is said to occupy XIII-XVII, in XIII
and posterior XVII being less well developed,
whereas in the Emu Lagoon specimens it ends
with segment XVI. No accessory genital mark-
ings were present. The thickened end of the male
duct was shown to open into the prostate duct
whereas in the Emu Lagoon material these two
ducts were shown to be closely apposed at the
male pore but actual fusion has not be demon-
strated. Two penial setae were seen in each folli-
cle whereas one only is present on each side in

EARTHWORMS FROM CAPE YORK PENINSULA 261

FIG. 30. Kayarmacia queenslandicus (Michaelsen, 1916). Emu Lagoon, Alice-Mitchell Rivers National Park,
specimen QM Acc#2344, QMG213385. Scanning electron micrographs of specimen 6, QMG213385. A, ventral
view. B, region of spermathecal pores. C, a right spermathecal pore. D, penial setae in situ. E, right penial seta
in situ. F, ventral ornamentation of penial seta. G, same, higher magnification. H, penial seta of specimen 2. I,
dorsal ornamentation of a penial seta of same. J, dorsal ornamentation of a penial seta of specimen 6.

the new material. Although the description of the
spermathecae given by Michaelsen differs in
some regards relative to that given here for the
Emu Lagoon material, the differences are doubt-
fully significant and his illustration agrees well
with that for the latter; the very wide duct
illustrated for the type corresponds with the wide
middle region of the duct in the new material. In
view of the close similarities of the new material
with the types, including the ventral ornamenta-
tion and smooth sides of the setal curvature, these
differences of description do not appear to merit
specific separation.

The close similarity of the material here identi-
fied as Michaelsen's Rhododrilus queenslandicus
and the new species Kavarmacia adelphicus is
discussed in the account of the latter, The speci-
men of K. queenslandicus in which the genital
marking in XVIII is postsetal, as in K. adelphicus,
further narrows the gap between the two taxa. If
genital setae were found in K. queenslandicus
retention of adelphicus às a distinct species
would still be supported by the difference in
urnamentation of the penial setae.

Subfamily MEGASCOLECINAE
Genus Terrisswalkerius Jamieson, 1995

Terrisswalkerius carbinensis sp. nov.
(Figs 31 & 32; 39)

TYPE LOCALITY. 16°30'30"S 145*16'30"E,, Mt
Carbine Tableland, 980m, Notophyll vine forest on
granite, K.R. McDonald and C. Hoskins, 22 Jan 1996.

MATERIAL EXAMINED. HOLOTYPE.
QMG212025. PARATYPES. P1, 3-9, QMG213392-
213399; P2 QMG212026; P10 QMG213400.

DESCRIPTION. Length 25-51mm; H 46mm.
Width 2- (H) 3mm. Segments 69-145; H 147.
Pigmentless buff in ethanol, clitellum pale brown.
Prostomium broad and prolobous but the peri-
stomium describing a wide V posterior to it, Dor-
sal canalicula absent. Peristomium slightly
shorter than segment II; not bifid ventrally. First
dorsal pore 4/5. Setae 49 in XII; 54 caudally (12
segments from posterior end); aa: ab: bc: cd: yz:
zz— 2.1: 1.0: 0,8: 0,6: 0,6; 0,8: thus ventral break
(aa) pronounced; dorsal break (zz), except ante-
riorly, hardly perceptible, all lines approximately
straight. Nephropores clearly visible on the
clitellum in å straight lateral series, in setal lines
12, anteriorly in their segments. Clitellum annu-
lar, XIV-XVII; intersegmental furrows weakly
represented; dorsal pores almost occluded; setae
present, male pores minute, each with a narrow

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 31. Terrisswalkerius carbinensis sp, noy, Holo-
type, QMG212025. Ventral view of forebody and
clitellar region.

pale border immediately lateral to setal line b, on
a conspicuous oval papilla which extends from
median to a to approximately c line; the papillae
is an evaginated structure representing the termi-
nation of the unusually muscular prostate duct,
Female pores a pair, presetally in a lines, ina pale
field at the anterior border of the clitellum, in
XIV, Spermathecal pores not visible but their
sites indicated by a short transverse glandular
ridge; approximately in c lines, immediately be-
hind intersegmental furrows 4/5, 5/6 and 6/7.

EARTHWORMS FROM CAPE YORK PENINSULA

prostate
gland

Å prostate

een fiar duct - x
B mm C mm
di spermathecal
spermathecal & diverticulum
pore f
spermathesal
ampulla
D imm
diverticulum éctal duct-at
eee it
E i mm nephrapore

FIG, 32. Terrisswalkerius carbinensis sp. nov. Holo-
type, QMG212025. A, dorsal view in region of pro-
stomium. B. left prostate. C, duct of same. D, right
spermatheca of VIL E, left nephridium in anterior
intestinal region.

Septa 7/8-12/13 strongly thickened. Dorsal
blood vessel single, continuous onto the pharynx.
Last hearts in XII. Gizzard a very large, glossy,
firmly muscular anteriorly slightly widening cyl-
inder; anterior to septum 6/7; its posterior end al
the level of segment LX; septum 5/6 attached to it
near its anterior limit but apparently not investing
il, Large pharyngeal glands in two or more pre-
ceding segments to not envelope the gizzard.
Oesophagus greatly dilated and obliquely striated
in XVI; the striations coincidmg with numerous
deep internal lamellae projecting inta the lumen
of what is here considered an unpaired calcifer-
ous gland. Intestine commencing in XIX in which
itis constricted by the prostates. Nephridia simple
stomate, vesiculate, holonephridia; the first blad-

ders anteriorly in II elongate pyriform, ad-
iverticulate; anteriormost nephridia forming å
thick coiled tube but not tufted; bladders in the
intestinal region with å small expansion where
the duct approaches the nephropore and large
lateral digitiform portion constituting a divertic-
ulum. Compacted sperm masses and well devel-
oped nacreous seminal funnels paired in each of
X and XI; seminal vesicles a pair of large smooth
sacs in IX and å pair of racemose, deeply dis-
sected masses in XII: no pseudovesicles in XIII.
Large laminar ovaries with many strings of large
oocytes, and compact oviducal funnels, in XIT;
ovisacs absent. Prostates large, tubuloracemose,
with very thick adpressed sections in a zigzagged
configuration which are closely adpressed so às
to give the entire gland a racemose appearance;
both embracing the oesophagus and intestine at
their juncture and meeting dorsally above the gul:
each with along much coiled muscular, anteriorly
strongly widening duct; vasa deferentia joining
the duct near its ental end. Spermathecae 3 pairs.
in V, VI and VII, each with an ovoid strongly
flattened ampulla and a short stout duct of about
one third ils length which is joined at the pore by
asingle, elongate clavate diverticulum, often bent
through a large angle near its slender base, with
Spermatozoal indescence; length right posterior
spermatheca (straightened) = 1, 5mm; ratio total
length: length duct = 4.2; ratio length: length
diverticulum (straightened) = 1.8.

ETYMOLOGY. Named for the type locality.

REMARKS. T. carbinensis shares with T:
montislewisi (Jamieson, 1976), T. barronensis
(Fletcher, 1887) and T. raveni (Jamieson, 1976)
the presence of three pairs of spermathecal pares
in 4/5, 5/6 and 6/7. The last two species differ
from it in their long and tortuous spermathecal
diverticula and location ofthe spermathecal pares
in the vicinity of setal lines d-e. T. montislewisi is
similar to T. carbinensis in having a short, clavate
diverticulum uniting with the spermathecal duci
at the body wall but the pores are more ventral, in
line with the ventral setal couples (ab), and its
prostate ducts are not as tortuous. The rnale
porophores in T. carbinensis differ from those of
T. montislewisi in appearing to be partly formed
by evagination of the muscular prostate ducts.

Terrisswalkerius mcilwraithi sp. nov
(Figs 33-36; 39)

TYPE LOCALITY. Peach Creek, McIlwraith Range,
13°44°17"S 143°20°15"E, alt. 500-520m, notophyll

264

FIG. 33. Terrisswalkerius mcilwraithi sp. nov. Holo-
type, QMG211995, Ventral view of forebody and
clitellar region.

vine forest with fan and feather palms, bank of creek,
in loose upper horizon, on Kintore adamallite granite,
K.R. McDonald, 26 & 27 Sep 1996.

MATERIAL EXAMINED, HOLOTYPE. (1)
QMG211995. PARATYPES, (1) Pl, used for SEM (+
immature non-type), QMG211996; P2-4
QMG211997-211999; P5, QMG212024 (used for
DNA study); P6, QMG212037.

DESCRIPTION. Length 25 (P5), 31 (P3), 34mm
(P2; holotype and paratype I are posterior ampu-
tees). Width (midclitellar) 2.0-2.3mm, wider in
the forebody. Segments 125 (P4), 137 (P3), 139
(P2). Colour in life chocolate brown, darker an-
terior to the clitellum; pale brown ventrally;
clitellum pinkish brown; in ethanol reddish
brown, clitellum and ventral surface whitish.

MEMOIRS OF THE QUEENSLAND MUSEUM

prostomium

dorsal
canalicula

vasa deferentla

prostate duct

B
spermathecal diverticulum
spermathecal duct
C spermathecal ampulla
nephropore
diverticulum of
nephridial bladder
D

imm

FIG. 34. Terrisswalkerius mcilwraithi sp. nov. Holo-
type, QMG211995, A, dorsal view in region of pro-
stomium. B, left prostate, C, left spermatheca. D, left
nephridial bladder of XX, A-D same scale.

Behaviour: saltatory by rapid ‘flicking’ move-
ments when disturbed. Prostomium epilobous
1/3, broad, transversely elliptical; in the holotype
giving the impression of being almost tanylobous
owing to transverse plications of the peristomium
posterior to it but this feature less obvious or
absent in paratypes. A deep, wide dorsal can-
alicula or gauge bisects the peristomium dorsally
and impinges on the posterior extremity of the
prostomium; this groove is present segmentally

EARTHWORMS FROM CAPE YORK PENINSULA

in H and HI (but not intersegmentally) and a
middorsal darker pigmented but not incised line
is present throughout the postclitellar region,
Peristomium not shortened, longer than segment
Il, with V-shaped midventral anterior indentation
continuous with a midventral groove. First dorsal
pore 4/5, Setae in XII 28 (H, P3), 36 (P2, P4); in
XX 28 (P2), 30 (H), 32 (P3), 36 (P4); caudally
(12 segments anterior to the posterior extremity)
36 (P4), 40 (H, P2), 42 (P3). The ventral break
(aa) only slightly wider than thc adjacent setal
couple (ab); the dorsal break (zz) slightly less than
iwice the adjacent couple (yz); ab not signifi-
cantly narrower than other intervals of the same
side; selae a and b absent in XVII; a line fairly
straight throughout; z lines check. The setal sur-
faces are cicatriced as is common in earthworms,
Nephropores visible in a straight series on the
clitellum, lateral, in setal lines 10, anteriorly in
their segments. Clitellum annular, XII-XVI
(paralypes 2 and 3) or weak in SI and absent in

VII (H. P1); intersegmental furrows and setae
retained; dorsal pores occluded. Male porophores
a pair of scarcely protuberant whitish ellipses in
XVIII centred slightly lateral of b lines (H, P1-3);
à central male pore visible on each only in P2; the
pores 0.5mm apart. Accessory genital markings
absent. Female pores on XIV, from internal inves-
tigation, but not visible externally. Spermathecal
pores one pair, in 7/8, slightly lateral of b lines
inconspicuous slits demonstrable only with diffi-
culty by parting the intersegmental furrow, very
approximately 0.7mm apart.

Septa 12/13 and 13/14 , sljghtly thickened, the
remainder delicate. Last hearts in XIL Gizzard a
lurge, glossy, firmly muscular, anteriorly slightly
widening cylinder; anterior to septum 7/8 bul
segmental position indeterminable owing to ex-
treme attenuation of seplå by posterior projection
of the gizzard, Very large pharyngeal glands,
forming a lobe on each side, envelope the anterior
region of the gizzard. Oesophagus vascular in
X-XV; enlarged in XIV and XV in which it has a
pair of large circumferential vessels connecting
to the supraoesophageal vessel but no true calcif-
erous glands present. Intestine commencing, with
abrupt expansion, in XVII. Nephridia stomate
vesiculate, exonephric holonephridia commenc-
ing in I; those in H tufted, but not large, and
attached to the buccal cavity though enteronephry
not proven; those in TI] only slightly tufted and
like the remaining, simple nephridia, exonephric;
hladders not demonstrable anterior to IV; those in
ihe intestinal region conspicuous by virtue of an
obliquely vertical, outwardly leaning sac on each

spermalhecal
pore

FIG. 35. Terrisswalkerius meilwraithi 8p. nov. Para-
type 2, QMG211937. Ventral view of lorehody and
clitellar region

side which is the lateral diverticulum of each
bladder and often has a brownish, chloragogen-
ous appearance. Very large compacted sperm
masses and well developed nacreous seminal fun-
nels a pair in each of X and XI; seminal vesicles
a pair of large elongate lobulated sacs in XII and
a small pair in IX; no pseudo-vesicles in ZIL
Small ovaries with few egg strings, and large
oviducal funnels, in XIU; ovisacs absent. Pros-
tates tubuloracemose, confined to XVII, the
gland a flattened band coiled one on itself in a
horizontal plane, with the wide ental end anterior;
duct moderately long, muscular, bént once and
strongly widened ectall y; vasa deferentia joming
the gland shortly before its junction with the duct;

FIG. 36. Terrisswalkerius mcilwraithi sp. nov. Para-
type 1, QMG211996. Scanning electron micrographs.
A, a seta. B, sculpturing of same.

penial setae absent. Spermathecae one pair, with
large, subspheroidal ampulla and clearly demar-
cated, stout, fusiform muscular duct; the comma-
shaped uniloculate diverticulum, with
spermatozoal iridescence, joining the junction of
ampulla and duct; length left spermatheca =
0.8mm; ratio total length: length duct = 2.7; ratio
length: length diverticulum (straightened) = 1.6 (H).

ETYMOLOGY. Named after the type-locality.

REMARKS. Terrisswalkerius mcilwraithi is
closer morphologically to T: blounti than to other
members of the genus and shares with it the
otherwise unique restriction of the spermathecal
pores to intersegment 7/8 (Table 1). In view of the
close similarity of the two entities, and their rel-
atively close geographical proximity, albeit sep-
arated by 3? of latitude, this location of the
spermathecal pores is deduced to have developed
in a common ancestor and is a synapomorphy of
the two species. Of these, often subtle differ-
ences, the more significant, supporting specific
separation of mcilwraithi from blounti appear to
be the lack of closer coupling of the ventral setae

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1. Differences between T. mcilwraithi and T.
blounti.

Terrisswalkerius blounti
(Jamieson, 1976)
Type Locality: 16°34’S

Prostomium bisected by
middorsal groove

Terrisswalkerius mcilwraithi
sp. nov.

Type Locality: 13?44'17"S
Prostomium with middorsal
groove only posterior!

Setae: ab significantly narrower
than other intervals of the same
side

Setae: ab not narrower than `
other intervals of the same side

gaonly slightly wider than the

adjacent setal couple aa 72-4 ab (widest caudally)

zz slightly less than twice the
adjacent couple

Nephropores in setal lines 10

Male porophores 0.5mm apart [Male pores Imm apart

Spermathecal pores, slightly
lateral of b lines, very
approximately 0.7mm ap

zz smaller than adjacent couple

Nephropores in setal lines 5

Spermathecal pores between

setal lines 3 and 4, 1.7-2.0mm

art apart

Septa 12/13 and 13/14 , slightl ;

thickened, Gerais 7 d rione
icate

Nephridia with diverticulate

bladders Nephridia lacking bladders

Prostates long, flattened and
irregularly tubular .
(tubuloracemose), meandering
with adpressed coils in XVII,
XVIII-XXIV, XXII

Prostates tubuloracemose,
confined to XVIII, the gland a
flattened band coiled one on
itself in a horizontal plane

Spermathecae with stout,

2 Spermathetae with long,
fusiform muscular duct s

ender and twisted duct

Ratio total length spermatheca:
length duct = 1.7

Ratio length: length Ratio length: length
diverticulum (straightened) 21.6 | diverticulum = 2.

Ratio total length spermatheca:
length duct = 2.7

a and b; location of nephropores in setal lines 10
(not 5, though lateral in both species); the closer
approximation of the spermathecal and male
pores; the presence of nephridial bladders; and
the restriction of the prostates to XVIII and their
coiled, not tortuous, form.

The type-locality of this species is almost 2° of
latitude north of previous records for the genus.
Its habitat is notophyll vine forest with fan and
feather palms (Licuala ramsayi, Ptychosperma
elegans, Calamus australis, and Archonto-
phoenix spp.) along drainage lines with moist,
humic soils, on Kintore Adamallite granite.

It is found sympatrically with Diplotrema
attenuata, D. scheltingai, Neodiplotrema
mcdonaldi, and species yet to be identified.

Terrisswalkerius miseriae sp. nov.
(Figs 37-39)

TYPE LOCALITY. 15°52’S 145°13’E (I'x1" grid),
Mt Misery, S.W. slope, Timber Reserve 165 Monk-
house, underlogs, notophyll vine forest, alt. 600-750m.
K.R. McDonald, C. Schneider, J. Sumner, 11 Nov
1994.

EARTHWORMS FROM CAPE YORK PENINSULA

male pore

FIG. 37. Terrisswalkerius miseriae sp. nov. Holotype,
QMG?1 1964. Ventral view of forebody and clitellar
region.

MATERIAL EXAMINED. HOLOTYPE, QMG21 1964.
PARATYPES. PI and 2, QMG211965-66. OTHER
MATERIAL. 3 immature specimens not designated
types.

fr vasa
A deferentin

prosials gland
prostate duct median

chambar

tmn
nephtogore

nephndial J

bladder —7—

spermathenal
ampulla

spermathecal
nerve cord diverheulum

FIG. 38. Terrisswalkerius miseriae sp. nov. Holotype,
QMG211964, A, dorsal view in region of prosto-
mium. B, left prostate, C, spermathecae of IX, in situ.
D, right nephridial bladder of VILL B-D same scale.

DESCRIPTION. Length 48-70mm; H 54mm.
Width (midclitellar) 3-(H)5mm (H). Segments
97-111; H 94. Colour in ethanol purplish brown,
pale ventrally; clitellum buff. Prostomium
epilobous 1/2, dorsal tongue open, sides parallel
or slightly concave; a wide, deep, middorsal
groove bisecting the peristomium ànd continuing
posteriorly as a narrow groove (canalicula) which
more distinct in the postclitellar region. Peristom-
ium bifid ventrally, First dorsal pore 6/7. Setae 31
in XII; 40 caudally (12 segments from posterior
end); all rows approximately straight. In XII, aa:
ab: be: cd: yz; zz = 1.0: 1.0: 1.5: 1.1: 3.0: 4.5; no
ventral break; dorsal break ca. 1 to 3 times the
adjacent interval. Nephropores conspicuous from
V posteriorly, near the anterior borders of their

268

Kayarmacta adelphicus
Termisswalkenus rneilwraihi

Diplotreme acropetra

Tarriswalkerius carbinensis

nyarimacia Quesnslandicus
Tarrisswalkerius miserae

a
Dipletrama schenmgar

Neodiplotrama modanaldi

Meoxiplatrems allanmoul

Neodiplotrema partounctata

Diplotremas attenuata

aS Å cm

FIG. 39. Relative sizes of the oligochaete species de-
seribed in this paper.

segments, in a slightly dorsolateral straight line
on each side, with rare irregularity in single seg-
menis; much less conspicuous behind the
clitellum; those on the clitellum in setal lines 10
from the ventrum. Clitellum annular, XIV-XVII;
setae, nephropores, dorsal pores and inter-
segmental furrows retained. Male pore, in XVIII,
in a midventral transverse slit on very small trans-
versely elliptical whitish papilla centred shortly
anterior to the setal arc and not extending poste-
riorly to this, surrounded by à narrow, rounded
oblong, low pad which is subdivided into two
anterior and two posterior quarters. Genital mark-
ings absent. Female pore single, midventral, mid-
way between the setal arc and anterior margin of XIV.

MEMOIRS OF THE QUEENSLAND MUSEUM

Spermathecal pores unpaired, midventral, in 7/8
and 8/9, inconspicuous, concealed in the inter-
segmental furrow but succeeded by an indistinct
whitish spot at the anterior border of the succeed-
ing segment.

Septa 10/11-13/14 the thickest, but not strongly
thickened. Dorsal blood vessel single, continuous
onto the pharynx, Last hearts in XII. Gizzard very
large, an elongate, glossy. muscular, anteriorly
widening cylinder; in V but septum 5/6 very
delicate and displaced, funnel-like, posteriorly,
the posterior end of the gizzard being at the level
of segment IX. Oesophagus in VII-XI segmen-
tally swollen and whitish, increasing in width
throughout these segments; broadly tubular in
XII-XVII; calciferous glands absent. Intestine
thin-walled, commencing with abrupt expansion
in XVIII(?). Nephridia stomate, vesiculate,
holonephridia, with large presepial funnel and
simple, elongate pyriform bladder. Fan-like testes
and large iridescent sperm funnels two pairs, tree
in X and XI. Ovaries large, palmate, and funnels
in XM; ovisacs not seen. Prostates a pair, in
XVIH-XXI, each unusually coiled; the ectal half
almost råcemose, the ental half thickly tubular; å
flaccid, thin-walled duct, lacking muscular
sheen, passing medianly to join the lateral aspect
of a small midventral rounded chamber, not form-
ing a noticeably muscular bursa, at the pore;
conjoined vasa deferentia joining the duct at
about its ectal fourth. Spermathecae 2 pairs, each
with ovoid ampulla and slender duct of approxi-
mately the same length which is joined at approx-
imately its ental third by a digitiform
diverticulum about as long as the duct; the sper-
mathecal duct entering the body wall anterior to
setae a but presumably fusing with its fellow
before opening at the common pore.

ETYMOLOGY. For the type locality.

REMARKS. Terrisswalkerius miseriae com-
pletes the trend to midventral approximation of
the spermathecal and male pores which is evident
in several other species of Terrisswalkerius. Total
fusion of the pores. seen in this species, has
hitherto been a diagnostic character of
Fletcherodrilus. However, Fletcherodrilus re-
mains distinct in having unpaired spermathecae,
each with a pair of diverticula which are reason-
ably deduced to be retained from the single diver-
ticulum of each member of an original pair of
spermathecae. In T. miseriae the spermathecae,
although opening at a single pore, are paired. There
are three or five spermathecal pores in Fletchero-
drilus, contrasting with two in T. miseriae.

EARTHWORMS FROM CAPE YORK PENINSULA

Fletcher (1889), describing the type specimens
af Fletcheradrilus, the type-species of its genus,
claimed to have observed a stage in reduction of
paired spermathecae, each of the pair with a sin-
gle diverticulum, and with one of the ampullac
rudimentary, to the unpaired condition seen in all
species of the genus. Despite this evidence for
origin of Fletcheredrilus from a form with paired
spermathecae, an origin which would be pre-
sumed, as above, even in the absence of the
exceptional individuals, the view is here maim-
tained that Terrisswalkerius miseriae is not more
closely related to Fletcherodrilus than it i$ to
pther species of Terrisswalkerius within what has
been shown ¢ladistically (Jamieson, 1994) to be
D monophyletic Fletcherodrilus-Terrisswalkerius
assemblage, A further difference of T, miseriae
trom Fletcherodrilus is absence of calcilerous
glands, It also lacks the pleated anterior genital
markings which are a novelty of some species of
Fleicherodrilus (F menurus and E sigillatus)
though these markings possibly were not present
in the stem form of Flercherodrilus, being un-
Known in the type-species, E unicus, or in F
affinis and F fasciatus, Relationships of T.
miseriae are the subject of a cladistic study of
mitochondrial DNA which is in progress.

DISCUSSION

In the above account, ten new species, in four
genera, have been deseribed from the Cape York
Peninsula, Queensland: Diploirema attenuata sp,
nov; D. scheltingai sp. noy.; Neodiplatrema
micdonaldi sp. nov; and Terrisswalkerius
meilwraithi sp, nov.. from the McTlwraith Range;
Diplotrema acropetra sp. nov., Neodiplotrema
altanmont sp. NOV, N, paripunctata sp. nov., and
Kayarmacia adelphicus gen. et sp. nov, from
Cape Melville National Park; Terrisswalkerius
carbinensis sp. nov. trom Mt Carbine Tableland,
and T. miseriae sp. nov., from Mt. Misery, near
the headwaters af the Normanby River.
Kayarmacia queenslandicus (Michaelsen, 1916)
is redescribed from the Ahce-Mitchell R. Na-
tional Park, near the type-locality; itis clearly the
sister-taxon of K, adelphicus.

These and other oligochaete species do not
show congruence with the biogeographical affin-
ives of various areas of Australia proposed by
Crisp et al. (1995) in a cladistic analysis nf the
distribution of 11 angiosperm groups, Thus, of
some 25 or more Australian oligochaete penera,
Begemius Easton (1982) ts the only indigenous
genus which is also found in New Guinea, au far

as is known, whereas those authors recognize a
close rélationship between southern New Guinea
and the Australian mainland. The gcnus
Diplotrema, of which species are described in the
present paper, has its closest relative,
Acanthodrilus, in New Caledonia (Jamieson &
Bennett. 1979) and these are in turn very closely
related to eastern North American
acanthodrilines, some of which may be referable
to Diplotrema (see James, 1990). It appears that
the Australian Diplotrema represent a
Gondwanan and possibly Pangaean acanthodrile
stock. The other Australian acanthodriles de-
scribed would previously have been attributed to
the genus Ahododrilus, following Michaelsen
(1916), but that they are congeneric with the
predominantly New Zealand species, and the
type-species, R, minutus Beddard (1889), is not
here supported and the new genus Kayarmacia is
erected to receive them. No acanthodriles are
known from New Guinea,

The genus Neodiplotrema, erected by Dune
(this volume) for species in the north-west of
Cape York Peninsula and Thursday Island, is here
shown tooccur sympatrically with Diplotrema on
the eastern side of the peninsula, at Cape Melville
and in the McIlwraith Range. It is distinguished
from Diplotrema by the meronephric condition
and is reasonably considered by Dyne to have
been derived by the supervention of this condi-
tion on the holonephry of Diplotrema. On this
basis erection of Neadiplorrema would leave a
paraphyletic Diplorrema unless, as seems im-
probable, the twa genera are sister-taxa, There
are, indeed, some indications that Lhe
Neodiplorrema condition was derived locally
from that of Diplotrema, as is seen in the similar-
ity of N, medonaldi 10 D. ridei Jamieson & Dyne
(1976), from Cape York and Melville Island.
However, there is no particular reason to believe
that acquisition of meronephry in Neodiplorrema
wasa monophyletic event and the genus may well
be polvphyletic from within Diplotrema, On
sirict cladistic principles Diplorrema and
Neodiplotreme appear to form å monophylum
and splitting of this into two genera thus is prob-
ably artificial, though convenient.

There remains, however, a possibility that ori-
gin of meronephry in Neodiplorrema was not
temporally confined to Australia but thal is may
have been a more ancient event which occurred
in a larger, Gondwanan or even Pangaean urea.
The similarity of Neodiplorremi to the species of
Dichogaster, itself a congeries, in Africa, central
America and Asia might, if this were the case, he

370

an indication of close relationship of
Neodiplotrema and the acanthodriles with those
Dichogastrini which, like Dichogaster (though
not the sexprostatic Fijian type-species, D,
damonis Beddard), have the acanthodriline ar-
rangement of male pores. The occurrence of true
calciferous glands in Neodiplatrema mcdonaldi
adds to the remarkable similarity of this species
to Dichogaster, notwithstanding the diplotreman
affinities mentioned above. Although at present
there is no firm reason to doubt local derivation
of Neodiplotrema from Diplorrema these altema-
tive affinities must be considered in further stud-
ies. Molecular studies which may shed light on
the phylogenetic relationships of these and other
genera are in progress.

Three species are here added to Terrisswalkerius.
This genus occurs in and near the Wet Tropics of
north Queensland, though there are grounds for
considering relationship to Indian species (see
Jamieson, 1977) at present placed in the plesio-
morphic congeries known as Diporochaera. Ter-
risswalkerius shows no close affinity to Australian
species of Diporachaeta or to the type-species of
that genus, and its congeners, in the New Zealand
(see Jamieson, 1994). There is noindicalion in the
distribution of Terrisswalkerius in the Wet Trop-
ics of Queensland tå suggest that the Atherton
Tableland has closer biogeographic affinities
with more southerly areas of E Australia, postu-

lated for angiosperms by Crisp et al. (1995), than

with other arcas of the wet tropics. Nevertheless,
the genus Fletcheredrilus which forms a
monophylum with Terrisswalkerius (see Jamic-
son, 1994) has a distribution which is both sym-
patric with Terrisswalkerius and includes eastern
Australia as Far south as N New South Wales.

ACKNOWLEDGEMENTS

The author is grateful qo those who collected or
aided in collection of material descrihed in this
paper; L.A. Jackson, PI Lethbridge. K.R. Me-
Donald, AJ. Stewart and WE Martin. It is im-
possible to adequately thank Keith McDonald for
his incomparable collections and for the encour-
agement which he has given the author. David
Scheltinga and Lina Daddow gave valued techni-
cal assistance. All illustrations are by the author.
This study was made possible by ABRS and ARC
grants to the author.

LITERATURE CITED

BEDDARD, F E. 1889. On the oligochactous fauna of
New Zealand with preliminary descriptions of

MEMOIRS OF THE QUEENSLAND MUSEUM

new species. Proceedings of the Zoological Soci-
ety of London 1889: 377.382.

CRISP, M.D., LINDER, H.P. & WESTON, P.H. 1995.
Cladistic biogeography of plants in Australia and
New Guinea: congruent pattem reveals two en-
demic Gëss tracks. Systematic Biology 44(4):
457-473

DYNE, GR. 1997, Two new genera of Acanthodrilinac
(Megascolecidae, Oligochaeta) from Cape York
Peninsula and the Torres Strait. Memoirs of the
Queensland Museum (this volume):

FLETCHER, J.J. 1889. Notes on Australian carth-
worms-Part V. Proceedings of the Linnean Society
of New South Wales 3(2); 1521-1558.

1890, Notes on Australian earthworms-Part VI. Pro-
ceedings of the Linnean Society of New South
Wales 4(2): 987-1019.

JAMES, S.W. 1990. Diplotrema murchiei and D.
papillata, new earthworms (Oligochaeta:
Megascolecidae) from Mexico. Acta Zoologica
Mexicana Nueva Serie 38: 18-27,

JAMIESON, B.G.M. 1976. The genus Diporochaeta
(Megascolecidae; Oligochaeta) in Queensland.
Zoologische Verhandelingen 149: 1-57.

1977. Preliminary description of Indian earthworms
(Megascolecidae: Oligochaeta) from the Piot
Hills. Bulletin du Muséum national d'Histoire
Naturelle, Paris, 3° Série, no. 450, Zoologie 313:
471-502,

1994. Some earthworms from the Wet Tropics and
from Bunya mountains, Queensland
(Meyascolecidae: Oligochaeta). Memoirs of the
Queensland Museum 37(1): 157-181.

1995, New species and a new genus of earthworms
in the collections of the Queensland Museum
(Megascolecidae; Oligochaeta). Memoirs of the
Queensland Museum 38(2): 575-596.

JAMIESON, B.G.M. & BENNETT J,D, 1979, New
species of Acanthodrilinae and à new genus of
Perionychini (Oligochaeta. Megascolecidae),
from New Caledonia, their phylogeny and zooge-
ography. Bulletin du Muséum national d'Histoire
Naturelle, 4* Série 1: section A, 353-403.

JAMIESON, B.G.M. & DYNE, G.R. 1976, The
acanthodriline earthworm genus Micraseolex
(Diplotrema) (Megascolecidae: Oligochaeta) in
the Northem Territory of Australia. Australian
Joumal of Zoology 24 : 445-476.

MICHAELSEN, W. 1907. Oligochaeta, in: Dic Fauna
Sudwesl-Australiens 1(2); 117-232. Gustav Fi-
scher: Jena.

1916. Results of Dr E. Mjóberg's Swedish Scientific
Expedition to Australia 1910-1913.
Oligochaeten. Kungliga Svenska Vetenska-
pakademiens Handlingar 5X 13): 3-74.

SPENCER, W.B, 1900, Further descriptions of Austra-
lian earthworms, Part I. Proceedings of the Royal
Society of Victoria 13 (n.s.)( 1): 29-67.

NEW SPECIES OF PTERIIDAE, PINNIDAE, VENERIDAE, EUCIROIDAE AND
CORBULIDAE FROM AUSTRALIA (MOLLUSCA: BIVALVIA: VENEROIDA).

KEVIN L. LAMPRELL AND JOHN M. HEALY

Lamprell, K.L. & Healy, J.M. 1997 06 30: New species of Pteriidae, Pinnidae, Veneridae,
Euciroidae and Corbulidae from Australia (Mollusca: Bivalvia: Veneroida). Memoirs of the
Queensland Museum 42(1): 271-281, Brisbane, ISSN 0079-8835,

Seven new species of bivalves from Australia are figured and described: Pieria howensis sp.
noy and P cooki sp. nov. (Pteriidae), Atrina (Servatrina) palmensis sp. nov. (Pinnidae);
Dosinia carpentariana sp. nov. (Veneridae); Euciroa queenslandica sp. nov. (Euciroidae);
Corbula (Serracorbula) moretonensis sp.nov and C. (Anisocorbula) stephensoni sp. nov.
(Corbulidae), All species are from Queensland except Preria howensis, which is known only
trom Lord Howe Island. [ ] Mollusca, Bivalvia, Veneroida, Australia,

Kevin L. Lamprell, Malacology Section, Queensland Museum, PO Box 3300 South Brisbane,
Queensland, 4101, Australia; Jonh M. Healy, Department of Zoalogy, University of Queens-

land, Queensland, 4072, Australia; I May 1997.

During the preparation of the second volume of
‘Bivalves of Australia’ (Lamprell & Healy, in
prep.) å number of apparently unnamed species
were uncovered 1n the collections of the Austra-
lian Museum (Sydney), the Queensland Museum
(Brisbane) and that of one of the authors
(Lamprell Collection), These species include rep-
resentatives of the following genera/subgenera
and families: Pteria (Pteriidae), Atrina
(Servatrina) (Pinnidae), Dosinia (Veneridae),
Euciroa (Euciroidae) and Corbula (Anisocorbula
and Notocorbula (Corbulidae). After searching
the relevant literature, especially that pertaining
to the Australian region (including Iredale, 1939;
Lamy, 1941; Hynd, 1954; Takemura & Okutam,
1958; Rosewater, 1961: Fischer-Piette & Delmas,
1967, Healy & Lamprell, 1992; Lamprell &
Whitehead, 1992; Lamprell & Stanisic, 1996;
Poutiers & Bernard, 1995) and after comparisons
with type specimens held in the Natural History
Museum, London, the Muséum national
d'Histoire Naturelle, Paris and the Department of
Malacology, Zoological Museum of the Univer-
sity of Amsterdam, we were satisfied that our
species should be described as new.

MATERIAL AND METHODS

All measurements were made using vernier dial
calipers. Shell height given is from the umbones
lo the ventral margin on the left valve vertical to
the hinge line. Shell length given is the greatest
distance from anterior to posterior margin. Shell
depth given is the greatest distance between the
surfaces of the left and right valves. Shell size
given is for the largest specimen examined by the
authors unless otherwise stated.

Abbreviations used: AMSC = Australian Mu-
seum, Sydney; KL = Lamprell Collection;
QMMO = Queensland Museum, Brisbane; WAM
= Western Australian Museum, Perth; lv = left
valve; rv =right valve; pv = paired or conjoined
valves; NSW = New South Wales; NT = Northern
Territory; Qld = Queensland.

SYSTEMATICS

The systematic arrangement used for each fam-
ily herein follows that of various authors (Cox &
Hertlein, 1969; Hertlein & Cox, 1969; Keen,
19694,b) in the Treatise on Invertebrate Paleon-
tology with the exception of the Euciroidae which
follaws Poutiers & Bernard (1995) and thc
Pinnidae which follows Rosewater (1961).

Family PTERIIDAE Gray, 1847

Pteria Scopoli,1777

TYPESPECIES. Myrtilus hirundo Linnaeus, 1758; SD-
Kennard, Salisbury & Woodward, 1931.

Pteria howensis sp, nov.
(Fig. 1A-H)

ETYMOLOGY. For Lord Howe Island.

DESCRIPTION. Shell length to 40.0mm; mod-
erately solid, obliquely ovate, dorsal margin
straight, umbones extend above the dorsal mar-
gin; both valves convex, left much more so ånd
encompassing the right; a well-defined, rounded
fold extends from the umbones to the postero-
ventral margin. Anterior wing short, moderately
deep from the hinge line to the narrow byssal

t3
I
to

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Pteria howensis sp. nov. A-B, holotype; C-H, paratypes. A, external view of lv; B, external view of rv;
C, internal view of Iv; D, external view of lv; E, internal view rv; F, external view of rv; G, external view of lv;
H, external view of lv.

NEW MARINE BIVALVES

notch; posterior wing narrow, varying from short
to very long; anterior margin convex, posterior
margin concave. Sculpture of concentric growth
striae; penostracum of widely spaced, imbricated
concentric lamellae. Colour externally light
bronze with irregular, darker brown zigzag con-
centric patterns and concentric lines and an irreg-
ular black stripe on the umbonal fold; internally
Silvery nacreous, medially with a narrow pale
brown margin.

MATERIAL EXAMINED. HOLOTYPE: AMSC-
70584, Lpv, off Ball's Pyramid, Lord Howe I, 55m,
MelIntyre, 22.11,1960. Length 40.0mm, height
18.5mm, depth of conjoined valves 6.5mm,
PARATYPES: AMSC203159, Spv, Ilv, same data as
holotype. Length of largest paratype 27.8mm, height
19.6mm, depth of conjoined valves 7.8mm.

HABITAT AND DISTRIBUTION, Attached to
soft corals in shallow water; known only from
Lord Howe Island,

REMARKS. Preria howensis sp. nov. is similar
to P. peasei (Dunker, 1872), P. sibogae Prashad,
1932, P. loveni (Dunker, 1872) and P cypsellus
(Dunker, 1872) in outline, but does nor attain the
large size of these species, Additionally none of
the latter species have the consistent bronze col-
our evident in D howensis. P. penguin (Röding,
1798) and P. faleata (Lamarck, 1819), while sim-
ilar in shape, are much heavier-shelled species
than P. howensis and also attain à much larger
size. P. howensis is similar in shape to juvenile
specimens of P. penguin and P falcata, but differs
in colour, Both these species retain the same
colour to adulthood. P howensis in both juvenile
and adult has a consistent light bronze back-
ground shell colour, (black in P penguin and
hone-green in P. falcata) with brown, irregular
zigzag concentric patterns and irregular concen-
tric lines over the surface and an irregular black
stripe on the umbonal fald.

Pteria cooki sp. nov.
(Fig. 2A-D)

ETYMOLOGY. For Stephen Cook.

DESCRIPTION. Shell length to 52.5mm; fragile,
translucent, elongate, obliquely ovate, dorsal
margin straight, umbones extending above mar-
gin line; both valves convex, left much more than
Ihe right. Anterior wing more or less an extension
of the dorsal margin with a strong oblique depres-
sion or fold separating it from the main body;
posterior wing moderately short with an almost

273

obsolete fold separating it from the mam body of
the shell. Sculpture of concentric microscopic
striae; periosracum of widely spaced, narrow
radial processes arranged in concentric rows, par-
allel to the shell lip. Colour externally off-white
tn pearl-white with a strong radial brown ray
extending from the umbones to the posteroventral
margin, stronger on the lv. weaker and sometimes
obsolete on the right. with a narrow brown stain
extending the full length of the dorsal margin;
internally nacreous medially with a wide trans-
luscent white margin.

MATERIAL EXAMINED. HOLOTYPE:
QMM059356, Lpv, Shelburne Bay, north Qld,
LITS, 142^ 55 E, 18m, S. Cook, May 1992. Length
54. 2mm. height 20,0mm, depth of conjoined valves
8.Gmm.PARATYPES: AMSC203157, 3pw,
WAMA6,97, Ipv, same data as holotype;
AMSC320793, Ipv, 11°13.2'S, 143725.8'E,
Shelburne Bay, north Qld, 22m, S. Cook, Apr 1993.
Dimensions of paratypes: AMSC203157, Jength
52.5mm, height 23.5mm, depth of pv 9.5mm;
WAM46.97, length 51.3mm, height 24.5mim, depth of
pv 10.0mm, OTHER MATERIAL: KL, Ap, same datu
as holotype; QMMO59357, 1pv, Arafura Sea, stn 51 1,
allached (o sea anemone, Bureau of Fisheries, K, Col-
gan, 1989; QMMOS9358, I | pv, Arafura Sea, stn 524,
attached to sea anemone, Bureau of Fisheries, K. Col-
gan, Noy 1989; QMMOS9359, Ipy, Arafura Sea, sm
541, attached 10 sea anemone, Bureau of Fisheries, K.
Colgan, Nov 1989; QMMO59360, 2pv, Arafura Sea,
sm 505, attached to sea anemone, Bureau of Fisheries,
K. Colgan, Nov 1989; QMMO59361, Ipv, Arafura
Sea, stn 403, attached to sea anemone, Bureau of
Fisheries, K. Colgan, Nov 1989; QMMOS9362, Ipv,
Arafura Sea, stn 531, attached to sea anemone, Bureau
of Fisheries, K. Colgan, Nov 1989,

HABITAT AND DISTRIBUTION. Attached to
sea anetnones and soft corals to 22m; Preria cooki
appears restricted to north Qld and the Northern
Territory, presently known from Shelburne Bay
om the east coast of Cape York and the Arafura Sea.

REMARKS. P. cooki sp. nov. differs trom P.
levitara Iredale, 1939 principally in colour: P
cooki being pearly white with a dark brown zig-
zag, pattem and oblique rays whereas Plevirata
has a dark brown, heavily mottled shell with
obscure oblique rays and with no trace of macre-
ous lustre. P. levitata is also more solid, attains a
larger size and has more pronounced hinge teeth.
P. cooki occurs sympatically with P. brevialata
(Dunker, 1872) but can be distinguished from that
species by having a lighter shell, weaker sculp-
ture and shorter wings. The brown dorsal margin
and ray. constant in P cooki is absent in P
brevialata.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-D, Preria cooki sp. nov. A-B, holotype; C-D, paratypes. A, external view of lv; B, internal view of lv;
C, external view of lv; D, view of pv shown principally from rv aspect. E-G, Atrina (S.) palmensis sp. nov, E,
external view of rv of holotype; F, external view of lv of paratype; G, internal view of lv of paratype; H, growth
series A. (S.) palmensis (left to right: lv’s paratype, holotype, paratype); I, Atrina (S.) pectinata: three juvenile
specimens (dredged from same locality as A. (S.) palmensis).

NEW MARINE BIVALVES 2

Family PINNIDAE Leach,1819

Atrina Gray, 1842

TYPE SPECIES. Arrina vexillum (Born, 1778); SM-
Gray, 1847,

A. (Servatrina) Iredale, 1939.
TYPE SPECIES. Atrina assimilis (Reeve, 1858).

Atrina (Servatrina) palmensis sp. nov.
(Fig. 2E-H)

ETYMOLOGY. For Palm Island.

DESCRIPTION, Shell to 51.6mm in height; thin,
relatively small for family. Valves equivalve and
inequilateral, profiles triangular anteriorly and
quadrate posteriorly. Both valves well inflated in
proportion to the shell size. Dorsal margin of each
valve conspicuously convex; ventral margin
straight posteriorly, concave anteriorly; posterior
margin truncate, forming a right angle with the
posteroventral margin; ligament extending ap-
proximately half the length along the dorsal edge
from the umbones. Sculpture of well-defined,
narrow ribs (10 ribs/cm measuring from the
ventroposterior margin), curving strongly to-
wards the ventral edge, ornamented with fine
erect scales which are sparse on the dorsal sur-
lace, stronger and more numerous on the ventral
surface and obsolete towards the umbones; inter-
nally ribs are impressed as deep grooves. Colour
externally horn with purple-black urnbanall y; in-
ternally the nacreous surface covers the anterior
half of the shell.

MATERIAL EXAMINED. HOLOTYPE: AMSC-
203160, I pv, between Palm and Curacoa Islands, north
Nid, dredged. 9m, K. Lamprell & P. Spoor, Nov 1990.
Height 47.5mm, width from dorsal to ventral margins
15.1mm, width of pv 8.9mm, PARATYPES:
AMSC203161, ipv, same data as holotype. Height
33.5mm, width from ventral to dorsal margins
12.6mm, width of py 7.2mm; QMMO59355, llv, |
broken tv, off Palm I, north Qld, dredged, K. Lamprell
& P. Spoor, 1989, Height 51.6mm, width from dorsal
to ventral margins 16.5mm, depth of lv 4.5mm.

HABITAT AND DISTRIBUTION. Dredged to
Om in sandy, weed-covered areas ; between Palm
and Curacoa Islands, north Qld.

REMARKS. Arrina (S.) palmensis sp. nov. ap-
pears to live shallowly anchored to a sandy, weed-
covered bottom in the Palm-Curacoa Island
passage. Freshly collected specimens occasion-
ally showed traces of byssal threads (Lamprell

3
Si

pers. obs.), although these threads are not pre-
served in any of the specimens from the type
series.

Juvenile as well as large specimens of A (S.)
pectinata were also dredged from the same local
ity sometimes occurring in the same dredge haul
as specimens of A. (5. ) palmensis. However even
as very small juveniles, these two species can be
readily separated on the basis of valve profile and
external sculpture. Whereas Å. (5.) pectinata ex-
hibits marked variation in the number of rihs and
degree of ornamentation ( Fig. I ), A. (5.) palmen-
sts is remarkably constant in its sculpture. Atrina
(S.) palmensis compared with the holotype of Å,
(S.) penna (Reeve, 1858) held in the Natural
History Museum, London (BMNH
1252,8,29.41.3) from the Indonesia-Philippines
area, which exhibits a similar narrow valve pro-
file, differs in the dorsal edge (hinge) and ribs
being curved towards the posteroventral margin
(not curved in A. (S.) penna) while the ribs in A.
(S.) palmensis are densely ornamented with low
scales, the ribs in A. (S.) penna have strong, raised
scales medially to the dorsal edge and obsolete 10
the ventral edge.

Family VENERIDAE Rafinesque, 1815

Dosinia Scopoli, 1777

TYPE SPECIES. Venus concentrica Born, 1778; fide
Fischer Piette 1942; M,

Dosinia carpentariana sp. nov.
(Fig. 3A-E)

ETYMOLOGY, For the Gulf of Carpentaria.

DESCRIPTION. Shell length to 38.5mm; ovate,
equivalve, moderately inflated, solid; umbones
prosogyrate, lunule heart-shaped, raised cen-
trally, strongly impressed peripherally, sur-
rounded by the lamellate ends of the concemnc ,
principal ribs continue on the lunule as lamellas,
escutcheon long, narrow, defined by a raised,
spinose ridge: ligament impressed; anterodorsal
margin short, convex, sloping, widely rounded at
ihe anterior margin; posterodorsal margin con-
vex, sloping, becoming widely rounded at the
posterior margin; ventral margin evenly and
widely convex, Sculpture of numerous, strong.
widely spaced, concentric ridges which become
lamellose at the antero- and posterodorsal mar-
gins, interstices with 4-6 minor concentric ridges.
Ligament long narrow, brown in colour, Hinge
plate broad. Hinge of lv with anterior lateral tooth

276 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-E, Dosinia carpentariana sp. nov. A, B, external and internal views of rv of holotype; C, D, external
and internal views of lv of holotype; E, detail of primary and secondary concentric growth ridges. F-H, Euciroa
queenslandica sp. nov. F, G, external and internal views of lv of holotype; H, dorsal view of pv showing
umbones, lunule and extent of valve inflation.

NEW MARINE BIVALVES

small, knob-like, bifid; anterior cardinal blade-
shaped, raised, slightly oblique; median cardinal
thin, posteriorly oblique; posterior cardinal thin,
oblique, parallel to median cardinal. Hinge of rv
with socket to accommodate anterior lateral of lv:
anterior cardinal short, joined at apex with mod-
erately solid, oblique, raised, median cardinal:
postérior cardinal thin, raised, oblique. Muscie
attachment scars well defined, anterior adductor
scar narrow, elongate; posterior adductor scar
fear-drop shaped, Pallial line fine. Pallial sinus
deep, slightly angulate terminally, ascending, ex-
tending beyond shell median. Colour internally
and externally chalk-white; periostracum grey.

MATERIAL EXAMINED. HOLOTYPE; QMMO-
59363. tpv, Gulf of Carpentaria, 10"30' S, 137° 12.] E,
S, Cook, 50m, 19 Nov 1990. Length 27.0mm, height
26.4mm, width of pv 13.3mm. PARATYPES:
AMSC107105, 3lv, Gulf of Carpentaria, 12^42.5' S,
141991.7'E, 18m; AMSCI23934, 11v, 14°30'S,
141?20,5" E. NW of Edward River, Gulf of Carpentaria,
Coll, LLoch, 14,6m, 1976; AMSC304562, lly, NW of
Edward River, Gulf of Carpentaria, 14°46'S,
141?20,7 E, ëm, I. Loch, 1976; AMSC107040, 241v,
26rv, off Albert River mouth, Gulf of Carpentaria,
17°24'S, 139*47 E, D.F. MeMichael & J.C. Yaldwyn,
1963; AMSC107014, 8lv, lOrv, off Albert River
mouth, Gulf of Carpentaria, 1 7^24' S, 139°47'E, D.F.
McMichael & J.C, Yaldwyn, 4-9m, 1963;
AMSCI23102, Irv, W of Nassau River, E Gulf of
Carpentaria, 13*51'S, 141?2]'E, CSIRO, 7.3m. Di-
mensions of largest paratype (AMSC107105); Length
38.5mm, height 39.6mm, width 10.2mm.

HABITAT AND DISTRIBUTION, Trawled and
dredged to 50m in sandy mud: known only from
the Gulf of Carpentaria,

REMARKS. Dosinia carpentariana sp. nov. can-
not be confused with any other known Indo-Pa-
cific Dosinia because of its distinctive sculpture
of numerous, strong, widely spaced, concentric
ridges, whose interstices have 4-6 minor concen-
iric ridges, Other species occuring in northern
Australia åre D. amphidesmoides (Reeve, 1850),
D, histrio (Gmelin, 1790), D, sculpta (Hanley,
1845) D. amina tredale, 1930, D. tumida (Gray.
1838), D. mira Smith, 1885, D. exasperata
(Philippi, 1847), D. lochi Healy & Lamprell,
1992, D. queenslandica Healy E Lamprell, 1992
and D, laminata (Reeve, 1850), all of which have
narrow, concentric interstices, without interstitial
riblets; D. kaspiewi Fischer-Piette & Delmas.
1967 which has very fine concentric lirae with
faint, central radial threads; D. a/tenai Fischer-Pi-
ette & Delmas, 1967, D, jivenilis (Gmelin, 1791),
D, scalaris (Menke, 1843) and D contusa

LA
~~
~

(Reeve, 1850) which have wide concentric ridges
and narrow interstices and D incisa (Reeve,
1850) which has coarse, concentric ridges with
obscure radial ndges medially,

Family EUCIROIDAE Dall, 1895

Euciroa Dall, 1881

TYPE SPECIES. Verrirerdia Megantisvina Dall,
188 1-M.

Euciroa queenslandica sp, nov,
(Fig. 3F-H)

ETYMOLOGY, For Queensland

DESCRIPTION, Shell to 42.3mm in length; light
weight but moderately solid; moderately inflated,
equivalve, with an obscure, oblique ridge poste-
riorly, preceded by a shallow oblique flexure,
slightly gaping posteriorl y. Anterior margin short,
almost straight, widely rounded terminally; pos-
terior dorsa] margin long, straight, sloping, pos-
terior margin narrowly rounded; ventral margin
well rounded, concave at the llexure, Sculpture
of numerous, small, radial spinose ribs, strongest
at the posterior third, less so at the anterior third
and almost obsolete at the medial third, more
noticable ventrally. Colour white, internally na-
creous,

MATERIAL EXAMINED. HOLOTYPE; AMSC-
120128, IL pv, NE of Lady Musgrave L, off central Qld.
23"35,8'S, 152*45.5' E, 365m. Globigerina mud and
siliceous sponge, WP Ponder, I, Loch & P. Terrill,
Dec 1977. Length 34.1 mm, height 29.2mm, width
19,5mm, PARATYPES: AMSC321047, Zrv, same
data as holotype. Length of largest rv 27.8mm, height
21.6, width rv 8.5mm; QM MOS9367, Ip, Iv, Capric-
om Channel, off central Qld, trawled, 128m. Length of
pv, 42.3mm, height 35.3mm, width 24.8mm; length of
Iv, 38.1mm, height 32.0mm, width 11.7mm.

HABITAT AND DISTRIBUTION. Dredged in
Globigerine mud and siliceous sand to 365m:
central Qld

REMARKS. Euciroa queenslandica sp. nov. is
similar in size to E. galarheae (Dell, 1956). How-
ever E, queenslandica differs in having sloped
anterior and posterior dorsal margins (ixlmeast
straight in E. palatheae) and being deeper in the
posterior truncation, additionally it lacks the
strong radial, medial sculpture of E. galatheae. E.
queenslandica differs from Acreuciroa restrara
Thiele & Jaeckel, 1931 in lacking the extended,
narrowly attenuate posterior and the strong me-

278 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. A-F, Corbula (À.) moretonensis sp. nov. A, external view of rv; B, external view of lv; C, D, sculpture
of rv showing radial pustules; E, F, sculpture showing concentric ridges and massed pustules of posterior
margins.

NEW MARINE BIVALVES

dial radial sculpture of that species. E
queenslandica attains a far greater size than the
Western Australian E, granifera (Cotton, 1931)
and the Indo-Pacific (Northern Territory) E
crassa Thiele & Jaeckel, 1931, (42.3mm in
E.gueenslandica, 19.2mm in E. crassa and
6.8mm in E. granifera). E. queenslandica is more
attenuate posteriorly than the Indo-Pacific E.
eburnea Wood-Mason & Alcock, 1891 (roundly
ovate in E. eburnea) while E ,eburnea has denser
radial sculpture. E. gueenslandica lacks the trans-
versely elongate shape of the large Indo-Pacific
E. trapezia Pouners, 1982 and has stronger radial
sculpture posteriorly and ventrally.

Family CORBULIDAE Lamarck, 1818

Corbula Bruguiére, 1797

TY PE SPECIES, Corbula sulcata Lamarck, 1801; SD-
Schmidt, i818

Corbula (Anisocorbula) Iredale, 1930
TYPE SPECIES. Corbula macgillivrayi Smith, 1885 - OD

Corbula (Anisocorbula) moretonensis sp. nov.
(Fig. 4A-F)

ETYMOLOGY. For Moreton Bay.

DESCRIPTION. Shell length to 3.0mm, minute,
solid, moderately inflated, inequivalve; ry more
inflated, encompassing the slightly smaller lv,
widely truncate, with a well-defined fold extend-
ing from the umbone to the posteroventral mar-
gin. Anterior margin steeply sloping, narrowly
rounded terminally; postenor dorsal margin mod-
erately sloping, angulate; ventral margin convex,
concave anterior to the posterior ridge. Sculpture
of strong. well-spaced, rounded, concentric ridges
and wider interstices crossed by radial rows of
minute pustules on both valves. Colour white.

MATERIAL EXAMINED. HOLOTYPE: QMMO-
59365, Ipy, off Middle Banks, Moreton Bay, south-
east Qld, W. Stephenson, between Sept 1972-June
1974, Length 3.0mm, height 2.2mm, width 1.7mm.
PARATYPES: AMSC203162, 19pv, same data as ho-
lotype: AMSC203139, 17pv, same data ås holotype;
AMSC36301, many, Albany Passage, north Old, 24m,
C. Hedley.

HABITAT AND DISTRIBUTION, Dredged in
sund to 10m; Moreton Bay, Qld.

REMARKS. Although Corbula (Anisecorbula)
moretonensis sp. nuv. is minute like C.

(Notocorbula) monilis Hinds, 1843 and Corbula
(Varicorbula) rotalis Hinds, 1843. It can be
readily separated from these species in having
well defined, narrowly rounded, concentric
ridges and wider interstices crossed uniquely hy
radial pustules on both valves while the sculpture
in C. (N.) monilis consists of widely rounded
concentric ridges, stronger at the ventral margin
and obsolete at the umbones and without regular
radial lines. C. (V.) rotalis has widely rounded
concentric ribs on the rv while the lv is almost
devoid of concentric sculpture but has well-de-
fined radial ridges, absent on the rv. In addition
the rv in C. (A.) moretonensis just moderately
encompasses the Iv while in C. (N.) monilis the
rv is much larger than the Iv, and in C. ( V.) rotalis
the lv is even smaller and more deeply encom-
passed. Very similar sculpture to that of C. (A.)
moretonensis also occurs in C. (A) macgilliveavi
Smith, 1885 and C. (A.) taheitensis Lamarck,
1818, but these are much larger species (16-
32mm) and more quadrate in shape than C. (A.)
mortonensis,

Corbula (Notocorbula) Iredale,1930
TYPE SPECIES, Notocorbula vicaria Iredale, 1930-0D

Corbula (Notocorbula) stephensoni sp, nov.
(Fig. 5A-E)

ETYMOLOGY. For the late William Stephenson.

DESCRIPTION. Shell length to 2.0mm, minute;
moderately solid; inflated, inequivalve; rv in-
flated, widely truncate posteriorly; encompassing
the much smaller lv. Anterior margin steeply
sloping, widely rounded terminally; posterior
dorsal margin long, moderately sloping; posterior
margin angulate; ventral margin rounded, con-
cave al the obscure ridge, umbones to postero-
ventral angle. Sculpture of rv medially with
obscure concentric ridges, stronger ventrally wath
microscopic radial lirae; lv globose, smonth
medially with obscure concentric striae stronger
ventrally, Colour white,

MATERIAL EXAMINED. HOLOTYPE: QMM()-
59366. Ipv, Cleveland Bay, north Qld, 19715'S,
146°58'E, io 29m, Sept 1990. Length 2.6mm, height
1.54mm. PARATYPE: AMC203163, (py, same data
as holotype.

HABITAT AND DISTRIBUTION. Dredged in
sand to 29m; Cleveland Bay, north Qld.

REMARKS. Corbula (N, ) stepliensoni sp. nov. is
similar to C. (N.) monilis Hinds, 1833, C, (V)

280 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A-E, Corbula (N.) stephensoni sp. nov. A, external view of rv; B, external view of lv; C, external view
of rv sculpture; D, ventral sculpture in rv; E, posterior margin of lv showing detail of periostracal layers.

NEW MARINE BIVALVES

rotalis Hinds, 1843 and C. (A.) moretonensis in
size. However C. (V) rotalis and C. (A.)
moretonensis have well-defined concentric
sculpture with prominent interstitial radial lirae
(both valves in C. (A.) moreronensis, lv in C. (V.
rotalis) while C. (N.) stephensoni has ventral
concentric striae (obsolete medially) and micro-
scopic radial lines on rv only. C. (N.) stephensoni
is much more produced posteriorly than C. (V)
monilis and has a much narrower umibones
(widely rounded in C. (V.) monilis),

ACKNOWLEDGEMENTS

We are grateful to Ms Kathy Way, Malacology
Section of the British Museum (Natural History)
tor forwarding the type specimens of Arrina (5. )
penna (Reeve) for this study; to Mr Ian Loch,
Malacology Section, Australian Museum, Syd-
ney for allowing access to the collection and
providing loan material; and to Dr John Stanisic
and reviewers for helpful criticism of the manu-
script. Mr Philip Spoor assisted KL in dredging
material of A. (S.) palmensis off Palm Is and
Curacoa Is during November 1990, Thanks are
also due to Kylie Stumkat for the scanning elec-
tron micrographs of the corbulid specimens. Fi-
nancial support for this project was provided by
à Keith Sutherland Award to KL. An Australian
Research Fellowship and Senior Research Fel-
lowship from the Australian Research Council
made possible the participation of JH.

LITERATURE CITED

FISCHER-PIETTE, E. & DELMAS, D, 1967. Revision
des lamellibranches du genre Dosinia Scopoli.
Mémoires du Muséum national d'Histoire
Naturelle, Paris 47: 1-91.

HEALY, J. & LAMPRELL, K, 1992. New species of
Veneridae, Cardiidae, Crassatellidae, Tellinidae
and Mactridae from Australia (Veneroida,
Bivalvia, Mollusca) Journal of the Malacological
Society of Australia 13: 75-97.

281

HERTLEIN L.G. & COX, L.R. 1969, Pieriidae. Pp.
302-306. In Moore, R.C. (ed.), "Treatise on inver-
tebraté paleontology'. Part N, Mollusca 6,
Bivalvia 1-2. (Geological Society ot America and
the University of Kansas: Lawrence, Kansas).

HYND, J.S. 1954. A revison of the Australian pearl-
shells, genus Pinctada (Lamellibranchia). Austra-
lian Journal Marine and Freshwater Research 6:
98-137.

IREDALE, T. 1939. Mollusca. Pt 1. Great Barrier Reef
Expedition, 1928-29, Scientific Repons 5: 209-
425.

KEEN, M. 19693. Veneridae. Pp. 671-688. In Moore,
R.C. (ed.), "Treatise on invertebrate paleontology”.
Part N, Mollusca 6, Bivalvia 1-2. (Geological
Society of America and the University of Kansas:
Lawrence, Kansas).

1969b, Corbulidae. Pp. 692-698. In Moore, R.C.
(ed,).1969, ‘Treatise on invertebrate
paleontology’. Part N, Mollusca 6, Bivalvia 1-2.
(Geological Society of America and the Univer-
sity of Kansas: Lawrence, Kansas).

LAMPRELL, K. & WHITEHEAD, T. 1992, "Bivålves
of Australia’. (Crawford House Press Pty Ltd:
Bathurst, NSW).

LAMY, E, 1941. Révision des Corbulidae vivants du
Muséum national d'Histoire Narurelle de Paris.
Journal of Conchology 84: 1-33, 121-144, 211-
254.

POUTIERS, I.M. 1976. Mollusques: Bivalves,
Resultats Des Campagnes Musorstom, 1-Philip-
pines, Mollusques: Bivalves, Mémoires OR-
STOM, Paris 91: 325-356.

POUTIERS, J.M. & BERNARD, ER 1995, Carnivo-
rous bivalve molluses (Anomalodesmata) from
the tropical Western Pacific Ocean, with a pro-
posed classification and a catalogue of recent spe-
cies, Resultats Des Campagnes Musorstom, Vol.
14. Mémoires du Muséum national d'Histoire
Naturelle Paris 167: 107-187.

ROSEWATER, I. 1961. The family Pinmdae in the
Indo-Pacific. [ndo-Pacific Mollusca 104): 175-
226.

TAKEMURA, Y. & OKUTANI, T. 1958, On the iden-
tificaton of species of Pinctada found attached to
Pinctada maxima (Jarneson) in the Arafura Sea.
Bulletin of the Tokai Regional Fisheries Research
Laboratory 20: 47-59.

282 MEMOIRS OF THE QUEENSLAND MUSEUM

ERRATUM

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.

In the above paper the photograph on p. 307
was inadvertently printed upside down causing
the caption to be misleading. Fig. 1 and its cap-
tion are reproduced correctly here.

FIG. 1. Sectioned hollow stem of Canthium odoratum show-
ing ant workers and larvae (Podomyrma sp.) on the right and
an adult female of Torarchus endocanthium on the left.

PITAR RÓMER FROM AUSTRALIAN AND ADJACENT WATERS, WITH
DESCRIPTIONS OF FOUR NEW SPECIES (MOLLUSCA: BIVALVIA: VENERIDAE)

KEVIN L. LAMPRELL AND JOHN M. HEALY

Lamprell, K.L. & Healy, J.M. 1997 06 30: Pitar Rómer from Australian and adjacent waters,
with descriptions of four new species (Mollusca: Bivalvia: Veneridae). Memoirs of the
Queensland Museum 42(1): 283-306. Brisbane. ISSN 0079-8835.

Twenty seven species of Pitar (Veneroidea, Veneridae) occur in Australian waters. All are
figured and described, including four new species, P. (Hyphantosoma) caperi sp. nov., P.
(Pitarina) queenslandica sp. nov., P. (P.) curnowae sp. nov. and P. (P.) thornleyae sp. nov.
and one new record P. (H.) intricata (Dautzenberg, 1907). P. (H.) caperi sp. nov. is also
recorded from New Caledonian waters. P. (Hyphantosoma) is recorded from Australia for
the first time, with five species assignable including three formerly included in P. (Pitarina)
viz. P. (H.) spoori Lamprell & Whitehead, 1990, P. (H.) nancyae Lamprell & Whitehead,
Bang and P. (H.) limatula (Sowerby, 1853). [ ] Mollusca, Bivalvia, Veneridae, Australia,
Pacific.

Kevin L. Lamprell, Malacology Section, Queensland Museum, PO Box 3300, South Bris-
bane, Queensland 4101, Australia; John M. Healy, Department of Zoology, University of
Queensland, Queensland 4072, Australia; 1 May 1997.

Pitar Rómer, 1857 occurs worldwide but is
especially prolific and speciose in the Indo-West
Pacific region and constitutes the largest group of
venerids present in the Australian fauna. AI-
though most species prefer shallow-water sandy-
mud substrates, some occur among coral sand in
reef lagoons or among shell debris in sandy mud
to moderate depths (up to or occasionally exceed-
ing 146 metres). Previous studies of Australian
Pitar s.l. include Lamprell & Whitehead (1990),
Healy & Lamprell (1992) and Lamprell &
Stanisic (1996) resulting in a total fauna of 16
species. Harte (1993) recorded six fossil and five
extant species in Pitar (Hyphantosoma). She
noted only one living species from the Western
Pacific notably P(H.) intricata (Dautzenberg,
1907) but four from the eastern Pacific — P. (H.)
aletes Hertlein & Strong,1948, P. (H.) hertleini
Olsson,1961, P. (H.) pollicaris (Carpenter, 1864)
including one from Tahiti i.e. P. (H.) festoui
Harte. Harte (1993) emphasized the presence of
the fine zigzag sculpture in Hyphantosoma (ab-
sent in most other pitarines) which prompted a
re-evaluation of the subgeneric assignments of
the Australian Pitar. The present study
demonstrates that in addition to P. (H.) intricata,
Australia has four other species referable to
Hyphantosoma, three of which were previously
included in the subgenus Pitarina viz. (P. (H.)
spoori Lamprell & Whitehead, 1990, P. (H.)
nancyae Lamprell & Whitehead, 1990 and P. (H.)
limatula (Sowerby,1853). Zigzag sculpture has
also been observed in specimens of P. (Pitarina)
healyi Lamprell & Stanisic, 1996 from New Cal-

edonia, suggesting perhaps that this species may
be more correctly placed within the P.
(Hyphantosoma) rather than P. (Pitarina). How-
ever Harte (1993), while not dismissive of zigzag
sculpture as an important feature of the subgenus
Hyphantosoma, also raised the possibility that
such sculpture could have been developed in
more than one line of pitarines. In this account
Pitar is reviewed and an additional five Austra-
lian species are described. The subgeneric status
of all the species is reassessed in the light of
Harte's (1993) review of P. (Hyphantosoma). In
addition, specimens of Australian species occur-
ring in adjacent areas (Papua New Guinea, New
Caledonia and Fiji) are recorded for the first time.

MATERIALS AND METHODS

Material for this study derives from unsorted or
part-sorted lots from the Queensland Museum,
Brisbane (OMMO); Australian Museum, Sydney
(AMSC); Museum of Victoria (MVF); Western
Australian Museum (WAM); South Australian
Museum (SAMD); Natural History Museum,
London (BMNH); Museum national D' Histoire
Naturelle, Paris (MNHN); National Science Mu-
seum, Tokyo (NSMT); and fromthe private col-
lections of G. Curnow, Adelaide, South Australia;
P. Spoor, Townsville, Queensland (PS) and one of
the authors (KL).

Abbreviations used: Iv = left valve; rv = right
valve; pv = paired valves; AMS = Australian
Museum, Sydney; GBR = Great Barrier Reef; KL
= Lamprell Collection, Brisbane; MV = Museum

284

of Victoria; NEQ = northeastern Queensland;
NSW = New South Wales; NT = Northern Teri-
tory; PNG = Papua New Guinea: Qld = Queens-
land; WA = Western Australia;

Shell height given is from the umbe to the
ventral margin on the lv. Shell length given is the
greatest distance [rom anterior to posterior mar-
gin. Shell width given is the greatest distance
helween the surfaces of the conjoined left and
right valves. Shell size given is for the largest
specimen examined by the authors unless other-
wise stated.

Subgenenc divisions of Pitar as recognized by
Keen (1969) in the Treatise on Invertebrate Pale-
ontology, are used in the present study. All dimen-
sions given in figure captions are shell length.

SYSTEMATICS

Family VENERIDAE
Subfamily PITARINAE

Pitar Römer, 1857

DIAGNOSIS, Sheil ovate to subtrigonal and
moderately inflated: umbones situated anteriorly;
shell surface smooth or with obsolete concentric
sculpture sometimes developed into zigzag pat-
terns! lunule Well defined: escutcheon obsolete.

Pitar (Hyphantosoma) Dall, 1902

TYPE SPECIES. Cytherea carbasea Guppy, 1866, by
original designation.

DIAGNOSIS. Sculpture of zigzag chiseled
sculpture on at least part of the external surface
of the shell,

Pitar (Hyphantosoma) intricata
(Dautzenberg, 1907) (Fig. LAB; 8A)

Merernrix (Pitar) intricata Dautzenberg, 1907: 333,
pl.6, fig. |; Harte, 1993: 347, fig 3d,e: Lamprell A
Stanisic, 1996; n. 47, fig. Geh,

DIAGNOSIS. Shell large, length to 53mm; solid,
inflated. Sculpture of dense concentric ridges
crossed by zigzag grooves, weaker medially. Col-
our light brown-white externally, internally
White,

MATERIAL EXAMINED. HOLOTYPE: MNHN,
label marked: Coll. types J. de Conch. V 1907 p.333
pl VI EI. OTHER MATERIAL. uv AMSC142102,
Samarang, Indonesia, 6°58'S, 110°25'E; Irv,
AMSC096751, Poindimié, New Caledonia, 20756'S.
165?20'E; (pv. MNHN, New Caledonia, Noumea, 9m:

MEMOIRS OF THEQUEENSLAND MUSEUM

Irv, AMSCIS6551, Port Mathurin, Rodrigues I, Mas-
carene Islands, 199418, 63°25°E, mud and coralline
dredgings; 11v, AMSC123340, South I, Lizard I, GBR,
14°42'S, 145*27'E, outer side of reef; 1pv, PS, western
side of Great Palm I, NEQ. 4m. in sand among coral,
lv, AMSC306568, Cairns Reef lagoon, NEO,
15742'S, 142?30' E, 9-18m; ilv, AMSC142103, Mac-
tan I, Philippines, 10?5'N, 124"0'E, 7-30m, coral; Irv,
AMSC306573, Micronesia, Pohnpei, Caroline 1,
6?55'N. 158?15"E, on rocks and broken coral, outer
reef; Iv, AMSC306668, Eel Reel, GBR, KS
143°22'E, 4-8m; 11v, 1tv, AMSCIS3285, No Name
Reef, GBR, 14940'8, 145?39'E, rubble at bommie
base, Dec 1984.

HABITAT AND DISTRIBUTION. Among
coral-sand in reef areas, to 18m; ranging from
Mascarene Islands, Indonesia, Philippines, Caro-
line Islands, New Caledonia to northeastern Aus-
tralia,

REMARKS, P. (Hyphantosoma) — intricata
(Dautzenberg, 1907) is easily recognised by its
large size and fine zigzag sculpture, It is similar
to P. (H.) spoort (Lamprell & Whitehead, 1990)
in colour but larger, more inflated with stronger
concentric sculpture and less quadrate posteriorly.

Pitar (Hyphantosoma) spoori Lamprell &
Whitehead, 1990 (Fig. 1C-F; 8B)

Pitar ( Pitarina) spoori Lamprell & Whitehead, 1990:
42, 3341; Lamprell & Whitehead, 1992; pl, 67, sp.
519; Lamprell & Stanisic, 1996: p, 44, lig. 5e-l.

DIAGNOSIS. Shell length to 35mm; quadrate,
thin. Sculpture of dense concentric ridges crossed
by zigzag grooves, weaker medially. Colour
white, with sparse interrupted brown radial rays;
wide brown radial ray covering posterior; esculch-
con marked by a few, spaced brown lines; a deep
brown spot often occuring beneath the umbones,

MATERIAL EXAMINED. HOLOTYPE; Ipv,
AMSC133572, 3-6m, in gutters at top of drop off, N.
West tip of reef, N of No.5 Bank Reef, NEO, 13°40°S,
144*09'E. PARATYPES: AMSC142104, Broadhurst
Reef, Eof Townsville, NEQ, subtidal, 20 Jul 1975; 21v,
219, QMMO019860, DENT NEO, subtidal, Jul 1985;
Lv, Zrv, AMSC155847, Michaelmas Cay, off Cairns,
NEO, GBR Boring Exp, May-Jun 1926; llv, 2rv,
AMSC148161, Mystery Reef, Swain Reefs, NEQ, 4-
6m around bommie in lagoon, 12 Jan 1985; Ipv,
AMSC 155850, Wheeler Reef, NEO, subtidal; 2rv,
AMSC!I55849, Lizard I, NEO, 6- 17m. in sandy rubble,
10 Dec 1974; lly, AMSC155848, Lizard I, NEQ, al
base of reef, 9-12m, 11 Dec 1974. OTHER MATE-
RIAL: llv, Irv, AMSCIS3322, Reef 14-151, GBR,
NEO, 14955'8, 145*41' E, 8-1 3m, among sand in coral
heads, Dec 1984; 11v, AMSC30666, Osprey Reet,

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

Coral Sea, 13*53'S, 146°31'E, 13-17m, Dec 1984;
Ion, AMSC30665, Reef 21-182, Swain Reefs, Qld,
21°22'S, 151*4T'E, 10m, Jan 1995; Ipv, KL, Orpheus
I, NEO, in littoral sand near Octopus hole; ipv, KL,
Palm I side of Fantome I, low tide; Irv, KL, Rudder
Reef. NEQ. low tide.

HABITAT AND DISTRIBUTION. Among coral
sand and rubble to 17m; ranging from Mascarene
Islands, Mauritius, Indonesia, Philippines, Caro-
line Islands, New Caledonia, Papua New Guinea
to northern Queensland.

REMARKS, £ (Hyphantosoma) spoori Lamprell
& Whitehead, 199( is quadrate posteriorly com-
pared with P. (Pitarina) pellucidus (Lamarck,
1818) and P. (P.) subpellucidus (Sowerby, 1851)
which are convex, thin and of similar size. P. (H.)
spoori has a wide coloured posterior ray (no
posterior radial ray in P. (PI pellucidus), ob-
scurely coloured on the median and anterior sur-
face (strongly coloured in P. (P.) subpellucidus)
and lacks the purple spots under the umbones that
occur in P. (P.) pellucidus. Recent collecting in
New Caledonia has produced the first known
live-collected specimen of P. (H.) spoori (see
Lamprell & Stanisic, 1996). Previously, this spe-
cies was only known from dead shells taken in the
littora! zone of northern Queensland (Lamprell &
Whitehead, 1990),

Pitar (Hyphantosoma) caperi sp. nov
(Fig. 1G,H; 8D)

ETYMOLOGY. For the vessel ‘Caper’.

DIAGNOSIS. Shell length 10 26.2mm quadrate,
thin, inequilateral, anterior end of shell less than
one third of maximum length. moderately in-
flated, light-weight; umbones oblique, approxi-
mate, lunule wel] defined, lanceolate, striate,
surrounded by an impressed line; anterodorsal
margin short, shghily convex, sloping, narrowly
rounded terminally; posterodorsal margin
slightly convex, gently sloping, broadly truncate
posteriorly; ventral margin widely convex. Shell
smooth with microscopic concentric ridges
crossed anteriorly and posteriorly by obscure zig-
zag grooves; periostracum, thin, white, situated
posteriorly and at the ventral margin. Ligament
narrow, impressed, Hinge of Iv with anterior lat-
eral tooth long, peg-like; anterior cardinal thin,
acline, separated from median cardinal by in-
verted v-shaped socket; median cardinal solid,
triangular; posterior cardinal free, thin, elongate
and oblique; posterior lateral thin, parallel 10
nymph. Hinge of ry with peg-like antenor lateral

285

(woth; antenor and median cardinals peg-like,
nurrowly parallel, separate; posterior cardinal
oblique, long, bifid; posterior lateral, parallel to
nymph. Muscle attachment scars ill-defined, an-
terior adductor scar teardrop-shaped, posterior
adductor scar large, somewhat ovate. Pallial line
thin; pallial sinus extending approximately two-
fifths towards the shel] centre, wide, rounded
terminally. Colour white, with interrupted tan-ur-
ange zigzag patterning over the whole surface
escutcheon and Iunule crossed hy tan lines; um-
bones with two broad, radial, well defined, white
rays extending medially, internally white.

MATERIAL EXAMINED. HOLOTYPE: Ipv,
AMSC202973, 6m, Little Upolo Cay, NEO. P. Spoor,
Oct 1983: length of shell 22.8mm, height 18.7mm,
width of conjoined valves 11.7mm. PARATYPE: pv,
MNHN, Anse Vila, Noumea, New Caledonia, 3m, Sep
1994: length of shell 26.2mm, height 20.2mm, width
of conjomed valves 12,2mm, OTHER MATERIAL:
Ipv. PS, same data as holotype; Irv, N end of Fantome
1, Palm Island Group, 1986; Ipv, KL, same dila as
paratype,

HABITAT AND DISTRIBUTION. Among caral
sand to 6m; ranging from northeastem Queens-
land. Australia to New Caledonia,

REMARKS. P. (Hypfanrosoma) caperi sp. nov.
is similar to P (H.) spoori, P. (PI pellucidus
(Lamarck, 1818) and P. (Pl subpellucidus
(Sowerby, 1851) in shape but can be readily sep-
arated from these species by its zigzag sculpture
on the extremal surface (concentric striae only on
P. {P} pellucidus and P. (P) subpellucidus). P.
(H.) caperi lacksthe wide, brown, radial posterior
ray and wider pallial sinus of P. (H. ) spoori, while
PH.) spoori lacks the two white radial rays of 4
[H.] caperi. P. (H.) caperi appears to be a rare
species. Examination of the extensive Australian
Museum collection has failed to establish any
further specimens other than those examined in
this study.

Pitar (Hyphantosoma) nancyae Lamprell &
Whitehead, 1990 (Fig. H-L; 8F)

Pitar (Pitarine) nancyie Lamprell & Whitehead,
1990: 45, 3e-h: Lamprell & Whitehead 1992: pl. 67,
sp. 526; Lamprell & Stanisic, 1996; p.45, fig. Dn,

DIAGNOSIS. Shell length to 36mm. moderately
thin. Sculpture of microscopic concentric ridges
crossed anteriorl y and posteriorly by obseure zig-
zag grooves, Colour white, with wide, broken
rays of brown; Iunule and escutcheon sparsely
marked with brown zigzag pattern; umbones pur-

286 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A, B, Pitar (Hypantosoma) intricata (Dautzenberg). A, AMSC123340, Lizard Is, GBR, external lv
(enlargement note zigzag sculpture on posterior third of valve), 52.7mm; B, AMSC156551, Mascarene Islands,
internal rv (enlargement showing adductor scars and pallial sinus), 37mm. C-F, P.(H.) spoori Lamprell &
Whitehead. C, D, holotype, AMSC133572, Bank Reef, NEO. 31mm. C, external lv; D, internal rv. E, F, KL,
Orpheus Is, 33mm. E, external lv; F, internal rv. G, H, P.(H) caperi sp. nov., holotype, AMSC202973, Little
Upolo Cay, NEQ, 22.8mm. G, external lv; H, internal rv. I-L, P.(H.) nancyae Lamprell & Whitehead; I, J,
holotype, QMMO22851, Boyne Is, MEQ, 31 mm. I, external lv; J, internal rv. K, L, KL, trawled Palm Is, NEQ,
28.8mm. K,external lv; L, internal rv. M, N, P.(H.) limatula (Sowerby), KL, Dingo Beach, central Qld, 24.2mm.
M, external lv; N, internal rv.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

ple tipped; interior white, with a small, intense
purple spot inside the unibones.

MATERIAL EXAMINED. HOLOTYPE: (pw, QMMO-
22851, Boyne I, Qld, on beach near mouth of river, at
low nde, Feb 1988. PARATYPES: (pv,
AMSC160472, same data as holotype: Ipv,
AMSC 160473, between Palm 1 and Curacoa 1, NEQ,
LOm, sand and rubble, ipy, WAM940-89_ between
Palm Land Curacoa I, NEO. 10m, sand and rubble: Ipv.
MVF57680, Boyne I, Qld, in sand among shell debris
alter strong winds, on beach near mouth of river at low
tide, Jan 1988. OTHER MATERIAL: 2rv,
AMSCO36338, off Murray I, Torres Strait, 9?56'S,
14474 E, 9-15m, Aug 1907; 1lv, AMSC306543, Dunk
1, Qld, 17757'S, 146°10°E, Aug 1901; 2pv, lv, KL,
Palm L NEO. trawled 10m, Aug 1994; Ipv. KL, Boyne
L Qld, beach, low tide, 1994; 8pv, Lv, 2rv, KL, Palm
l.trawled in Curacoa Channel to 10m: 1 pv, KL, Rodds
Bay, Turkey Beach, central Qld; 2pv, KL. off
Caloundra, Qld, 79m.

HABITAT AND DISTRIBUTION. In sand and
rubble to 79m; ranging from central Queensland
to the Northern Territory, Australia and New Cal-
edonia.

REMARKS. D (Hyphantosoma) | nancyae
Lampreil & Whitehead, 1990 has previously been
recorded only from the coasts of central to north-
ern Queensland and New Caledonia. In Australia
this species usually occurs in depths ranging from
9-30m, however the specimens collected from
New Caledonia ranged from 33-60m. P. (H.)
nancyae is similar to P. (P.) sophiae (Angas.
1877) but åttains å much larger size and is more
solid. This species has been included in
P(Hyphantosoma) on the basis of the zigzag
sculpture on the anterior and posterior ends.

Pitar (Hyphantosoma) limatula (Sowerby,
1853) (Fig, IM,N; 8C)

Cyiherea limatula Sowerby, 1853; 640. sp. 98. pl, 136,
figs 200-201; Pirar (Hyphanrosoma) limarülum
(Sowerby) Habe, 1977: 257, Pirar (Pitarina)
limaiula Lamprell & Whitehead, 1992; pl. 67, sp.

5

A2.

DIAGNOSIS. Shell length to 35mm, solid.
Sculpture of fine concentric growth lines crossed
by obscure zigzag grooves anteriorly and (rarely)
posteriorly. Colour yellow, sometimes lightly
flecked or rayed with chestnut-brown; escutch-
con and lunule dark chestnut-brown.

MATERIAL EXAMINED. SYNTYPES: 2pv,
AMNH1912.6.4.9-10, locality unknown; Ipv,
AMSC306569, Wonga Beach, N of Mossman, NEQ.
16?22'S, |45?26' E, among rocks on sand and gravel:

287

Spv, 2rv, KL, Dingo Beach, central Qld, washed ashore

alter cyclone Aivu; Ipv. KL, Cape Upstart, NEQ,

bench, 1987; Ipv, KL, Goold 1. NEO, low tide, Nov
95.

HABITAT AND DISTRIBUTION. Among litto-
ral sand and gravel; ranging from Japan to central
and northern Queensland, Australia.

REMARKS. P. (Hyphantosoma) ` limatula
(Sowerby, 1853) is readily distinguished from the
ather Australian Pitar species by the conspicuous
dark staining on the lunule and escutcheon.

Pitar (Pitarina) Jukes-Brown, 1913

TYPE SPECIES, Cytherea cirina Lamarck, 1818 hy
onginal designation.

DIAGNOSIS. Posterior cardinal ohlique, not at-
tached to median cardinal, pallial sinus short.
Sculpture of concentric sculpture only. (Diagno-
sis here emended).

Pitar (Pitarina) citrina (Lamarck, 1818)
(Fig. 2A-C; 8B)

Cytherea citrina Lamarck, 1818: 306; Chione striata
Gray, 1838: 306; Dione citrina Reeve, 1863: pl.10,
fig.43: Dione ustulata Reeve, 1863: pl, 11, fig. 49,
Pitar (Pitarina) citrinus Lamprell & Whitehead,
1992: pl. 67, sp, 520; Pitar (Pitarina) citrina
Lamprell & Stanisic, 1996: p. 38. fig. 4c.d.

DIAGNOSIS. Shell length to 40mm: solid, in-
flated, subtrigonal to ovate. Sculpture of fine
concentric lirae. Colour orange, yellow or some-
times white, with purple posteriorly; interior
white with purple centrally and posteriorly.

MATERIAL EXAMINED, HOLOTYPE: Museur de
Genéva, Lamarck collection, No.24. OTHER MATE-
RIAL, 7pv, KL, Shark Bay, northern WA, 1987; 4pv,
KL, Back Beach, Dampier, northern WA, 1987; 2pv,
KL, Broome, northern WA; MNHN, 3pv. Nouméa,
New Caledonia: 41v, MNHN, Platier Quen Toro, New
Caledonia, 2pv, MNHN, Presqu'ile Nou Anse Kuendit,
New Caledonia.

HABITAT AND DISTRIBUTION. In shallaw,
loose sand and eel grass; ranging from Japan
(Habe, 1977 as striata) to north Western Aus-
Wale Northern Territory, Australia and New Cal-
eadonia.

REMARKS. P (Pitarina) citrina (Lamarck,
1818) is å well known northern Western Austra-
lian species where it 15 found abundantly in loose
sand and eel grass in shallow water. Typically it
can be identified by its yellow external colour

288 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-C, Pitar (Pitarina) citrina (Lamarck). A, B, KL, Shark Bay, northern WA, 45.4mm. A, external lv; B,
internal rv; C, KL, Broome, northern WA, external lv showing coloured rays, 38.5mm. D-G, P.(P.) marrowae
Healy & Lamprell. D,E, holotype, QMMO32900, Dingo Beach, central Qld, 38.1 mm. D, external lv; E, internal
rv. F, G, KL, Goold Is, NEQ, 43mm. F, external lv; G, internal rv. H-L, P.(P.) curnowae sp. nov. H, I, holotype,
AMSC202974, Madang, PNG, 42.7mm. H, external lv; I, internal rv. J, K, KL, Magnetic Is, NEQ. J, external
lv; K, internal rv, 50mm (senile specimen). L, paratype, QMM054081, Little Hope I, NEO, external lv, 40mm.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

(occasionally orange or white) and deep purple
internal posterior, Specimens identifiable as
Chione striata Gray, 1838 and Dione ustulata
Reeve, [863 occur sympatrically with PIP)
citrina in northern Western Australia with inter-
grading shells which show without doubt that
they are only forms of P. (PF) citrina

Pitar (Pitarina) marrowae Healy & Lamprell,
1992 (Fig. 2D-G; 8J)

Pitar (Pitarina) marrowae Healy & Lamprell, 1992:
76; Pitar (Pitaring) cf. osmunda (Iredale, 1936)
sensu Lamptell & Whitehead, 1992, pl. 67, sp. 521.

DIAGNOSIS, Shell length to 40mm; thin. in-
flated, subtrigonal to ovate. Sculpture of fine
concentric lirae. Colour cream or fawn, with pur-
ple posteriorly; internally cream, with a purple
stain at pallial sinus and posterior adductor
scar,

MATERIAL EXAMINED. HOLOTYPE: Ipv.
QMMO32900, on sandy mud flats at low tide, Dingo
Beach, central Qld, 20?05'S, 148730'E. 1985, KL-
PARATYPES. 3pv, QMMO3290], Palm I, 18"40'S,
146*35'E, NEO. subridal, 1990; 3pv, AMSC166905,
MVP60432, WAM559-91, Palm I, NEQ, 18°40'S,
146°35’E, subtidal. 1990, KL. OTHER MATERIAL:
Am, KL, Palm J, NEQ, 18*40'S, 146°35°E, subtidal.
1990; Ipv, KL. Slashers Reef No. 1, GBR, 1993; ipv-
Ilv, KL, Goold 1, NEQ, Nov 1955; Hy, Irv,
AMSC306539, E of Moreton Bay, southern Qld,
26°54'S, 153°32'E, 115-176 m; Irv, AMSC306541,
Smith Point, Cobourg Peninsula, NT, 117078,
132"08'E, Jun 1976.

HABITAT AND DISTRIBUTION. Buried in
sand and rubble; ranging from New South Wales
to northeastern Queensland, Australia, Papua
New Guinea, Singapore, Bali, Samoa and the
Philippines.

REMARKS. P. (Pitarina) marrowae Healy &
Lamprell. 1992 differs from P. (P.) citrina
(Lamarck, 1818) in being à much thinner shell,
with a more convex posterior margin. This study
has extends the known range to Papua New
Guinea, Singapore, Bali, Philippines and Samoa.

Pitar (Pitarina) curnowae sp. nov.
(Fig. 2H-L: 8H)

ETYMOLOGY, For Mrs G, Curnow,
DESCRIPTION. Shell subtrigonal, equivalve,
inequilateral with the umbones situated al the

anterior third, solid: umbones prosogyrate; lunule
well developed, flat, striate, defined by a faint

289

impressed line; anterodorsal margin short, almost
Straight, steeply sloping, narrowly convex termi-
nally: posterodorsal margin, long, slightly con-
vex, steeply sloping, convex Lo truncile
posteriorly; ventral margin convex anteriorly,
sinuate posteriorly, more pronounced in mature
specimens. Shell sculpture of strong. irregular
concentric striae and irregular growth pauses.
Periostracum, thick, grey, usually with
aglutinated sand adhering to the posterior of the
shell. Ligament, impressed, olive-brown in col-
our. Hinge of lv with anterior Jateral tooth well
developed. peg-like; anterior cardinal thin, joined
1o thick median cardinal forming an inverted
v-shape: posterior cardinal long, ridge-like, free,
Hinge of rv with pit for the reception of the ly
lateral tooth; antenor cardinal short, narrow; me-
dian cardinal thick, straight, posterior cardinal
well developed, bifid, oblique. Muscle attach-
ment scars well defined; anterior adductor scar
ieardrop-shaped; postenor adductor scar larger
and more round. Pallial sinus moderately shor,
ascending, rounded terminally. Shell colouration
white or cream-white externally, escutcheon yel-
low-orange; internally white with yellow colour
centrally extending to the posterior muscle scar.

MATERIAL EXAMINED. HOLOTYPE: 1PV.
AMSC202974, purchased in market, Madang, PNG.
Length of shell 42.7mm, height 35.3mm, width
25.1mm. PARATYPES. Ipv, SAMD18976, in linoral
sand, Little Hope 1, NEO. 30 Jun 1988. lpw
QMMOS9380 in littoral sand, Little Hope I. NEO, 30
Jun 1988; 4lv, 2rv, AMSC105230, 7pv, 21%, løv,
AMSC87941, Madang, PNG, 5" 12.58, 145"47.5'E,
swamp behind Nui I, among mangroves, 28 May 1970.
OTHER MATERIAL: [py, KL, in littoral sand, Mag-
netic I, NEO, O. Rippingalc-

HABITAT AND DISTRIBUTION. In linoral
sand and mangrove swamps; ranging from north-
erm Queensland to Papua New Guinea.

REMARKS. P. (Pitarina) curnowae sp.nov is
quite variable in shape, Younger specimens have
an evenly convex ventral margin while senile
specimens develop a concave (sinuate) ventral
margin posteriorly and become more obese. P
(P) curnowae ts most similar to P. (P.) cirina
(Lamarck,1818) in having a similar solid, sub-
trigonal inequilateral shell but differing in having
a sinuate margin posteriorly (evenly convex in £
(P.) eitrina) and completely lacking any purple
posterior colour. P. (P.) curnowae lacks the ovate
shape of P. (P.) affinis (Gmelin,1791) and the
truncate posterior and anterior angulate margin of
P. (P) prora (Conrad, 1837) and does nol altain

290 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-C, P. (Pitarina) osmunda (Iredale), Sydney Harbour, NSW. A, AMSC320466, paralectotype, external
lv, 39mm; B, AMSC60607, lectotype, internal rv, 38.5mm; C, AMSC303944, external lv, 36.5mm. D, E, P.
(P.) affinis (Gmelin), KL, Shark Bay, northern WA, 56mm. D, external lv; E, internal rv. F-I, P.(P.) prora
(Conrad); F, G, KL, North West I, 56.3mm. F, external lv; G, internal rv. H, I, AMSC306496, Mascarene Islands.
H, external lv; I, internal rv, 37.2mm.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

the large size of these species. P. (P.) curnowae
lacks the wide convex posterior margin and light
zigzag markings of P (P) potteri Healy &
Lamprell, 1992 which is similar in size and valve
inflation, but lacks a sinuate ventral margin.
P. (P.) curnowae differs from P. (P.) marrowae in
having a more solid shell and in lacking any
purple colour posteriorly,

Pitar (Pitarina) osmunda (Iredale, 1936)
(Fig. 3A-C; 8G)

Pitarina osmunda Tredale, 1936: 277.

DIAGNOSIS. Shell length ta 39mm; solid. sub-
trigonal. Sculpture of dense, fine concentric striac
and irregular coarser growth lines. Colour dirty
white internally and externally.

MATERIAL EXAMINED. LECTOTYPE: Irv,

AMSC60607, Sydney Harbour, NSW. PARALECTO-

TYPE. liv, AMSC320466, same data as lectotype.

OTHER MATERIAL: llv, AMSC303944, "Triton

Dredge’. eras Harbour, NSW, 33°50'-33°52'S,
" "IGE

HABITAT AND DISTRIBUTION. Dredged in
mud and littoral sand; known only from the type
locality, Sydney Harbour, New South Wales

REMARKS. P. (Pitarina) osmunda (Iredale,
1936) is similar to P. (P) citrina and F (P)
marrowae in shape, but lacks the external yellow
colour and purple internally and external poste-
rior of both these species. P. (Pl osmunda is not
as solid as P. (P. citrina and has a more extended
pallial sinus.

Pitar (Pitarina) affinis (Gmelin, 1791)
(Fig. 3D,E; 9A)

Venus affinis Gmelin, 1791: 3278; Pitar (Pitarina)
affinis (Gmelin) Prashad, 1932; 222; Pitar
(Pitarina) affine Habe, 1977: 257; Lamprell &
Whitehead 1992: pl. 67, sp. 514; Lamprell &
Stánisic, 1996: p. 42, fig. Sa-b.

DIAGNOSIS. Shell length to 68mm, solid,
glossy. Sculpture of fine concentric growth striae.
Colour white or cream with tan-brown blotches
and rays centrally and towards the umbones, pur-
ple spots sometimes occurring beneath umbones.

MATERIAL EXAMINED. 2pv, KL, Port Hedland,
northern WA, in littoral sand, 1989; Lpv, KL, Monkey
Mia, Shark Bay, northern WA, littoral sand, 1987; 1 pv.
KL, Capricorn Channel, central Qld, trawled; 4pv, KL,
trawled Taylor Reef, NEO, 10-12m, Dec 1995; Ipv.
KL, Palm I, NEQ, 10m, Aug 1994; Spv, KL, Curacoa

29i

Passage, Palm 1. NEO, Nov 1990; Ipv, KL, Dinge
Beach, central Qld, in littoral sand; 1pv, KL, Fiji, om
beach: løv, KL. PNG,

HABITAT AND DISTRIBUTION. In littora!
sand and eel grass; ranging from Japan, Philip-
pines, northeast and north Western Australia to
New Caledonia.

REMARKS. F (Pitarina) affinis (Gmelin, 1791)
is the largest of the P. (Pirarina) species found in
littoral sand, Examination of the shell middens on
Stradbroke Island shows that the species was acd
ås a major food source by the Aboriginal people

of the island. Cytherea inflata Sowerby, 185
appears to be a synonym of this well Ted
species.

Pitar (Pitarina) prora (Conrad, 1837)
(Fig. 3F-1; 81)

Venus prora Conrad, 1837: 253; Cytherea lacia
Lamarck, 1819: 307, Cytherea obliquara Hanley,
I 844: 355 (app); Piar (Pitarina) obliquatur (Han
ley) Habe, 1977: 257; Pitar (Pitarina) prora (Can-
rad) Lamprell & Whitebead, 1992: pl. 66, sp. 513;
Lampreil & Stanissc, 1996: 44, fig, 5c-u.

DIAGNOSIS. Shell length to 65mm; solid; lu-
nule large. Sculpture of even, fine, concentric
siue and growth lines. Colour usually white or
cream, internally and externally; some specimens
develop à fine tan zigzag or chevron pattern over
the external surface.

MATERIAL EXAMINED. liv, AMSC306669, Port
Mathurin, Rodrigues 1. Mascarene Islands, 19°41'S,
63°25'E, mud and coralline dredgings; Ipv, 3rv, 21v,
AMSC306496, AMSC306497, Anse aux Anglais,
Rodrigues {, Mascarene Islands, 19743'S, 63°25°E,
mud flats, May 1995; 1pv, AMSC306664, Mystery
Reef, Swain Reefs, GBR, 21°23'S, 152"01'E, silt in
lagoon, 10m, 13 Jan 1985; lpv, KL, Feather Reef,
NEO, in lagoon; Ipv, KL, Suva, Fiji, in coral sand on
reef. m. v, KL, Apia, West Samoa, on reef Nat; Ipv, KL,
North West I, Bunker Group, in lagoon, 1991

HABITAT AND DISTRIBUTION. Among coral
sand to 10m and on muddy sand flats at low tide;
ranging from Arabia, Tanzania, Madagascar,
Japan. Cocos-Keeling Islands, Hawaiian Islands,
New Caledonia to the northeast coast of Aus-
tralia.

REMARKS. F (Pirarina) prora (Conrad, 1837)
is the largest and heaviest shell in P. (Pitarina).
P. (P.) prora is not å common species throughout
its ien $ and is easily separated from other mem-

the genus by its almost pointed anterior

292 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. A-E, P. (Pitarina) pellucidus (Lamarck). A-C, AMSC141215, Broome, WA, 34.4mm. A, external lv; B,
internal rv; C, dorsal view of pv showing lunule and escutcheon. D, E, KL, Dingo Beach, central Qld. D, external
lv, 33.7mm; E, external lv, 30.5mm. F-J, P.(P.) potteri Healy E Lamprell. F,G, holotype , QMMO32902, Palm
Is, NEQ, 32.5mm; F, external lv; G, internal rv. H- J, AMSC086251, New Caledonia, 37.8mm. H, external lv;
I, internal rv; J, external lv (length 34.5mm).

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

margin and subquadrate posterior, Specimens of
P. (B) prora have been examined from multiple
lots which show obscure to strong tan radial pal-
terns in some specimens while others have an all
white surface, Cytherea laeta Lamarck, 1819 is
lodged in the Museum de Genéva (Collection
Lamarck, No. 24, type lot 1084/27) and appears
to be a synonym of this species, however the
pallial sinus is deeper while another, smaller
specimen from Lamarck's collection with the
same registration number (1084/27) has a shal-
lower pallial sinus similar to P. (F.) prora. Exam-
ination of the type of Cytherea obliguata Hanley,
1844: 355 (app) in the BMNH showed this to be
conspecific with P. (8) prora.

Pitar (Pitarina) pellucidus (Lamarck, 1818)
(Fig. 4A-E; 9B

Cytherea pellucida Lamarck, 1818: 313; Dione aus-
tralica Reeve, 1863; sp, 47, pl, 11; Pitar (Pitarina)
pellucidus Lamprell & Whitehead, 1992: pl. 67, sp.
516; Lamprell & Stanisic, 1996; 42, lig. del,

DIAGNOSIS. Shell length tå 45min, thin, mod-
erately inflated. Colour ranging from pure white
to cream, usually with tan-coloured zigzag pat-
terning, urnbones often tinged light-brown with à
purple spot beneath them,

MATERIAL EX AMINED, | pv, BMNH1912,6,18.14,
specimen figured by Sowerby (1853); Ipv, BMNH,
holotype of Cytherea ausrralica Reeve, 1863 from the
Torres Strait. OTHER MATERIAL: Arv,
AMSC075783, Bathurst Head, Princess Charlotte Bay,
NEO 14°16'S, 144^12"E ; Irv, AMSC142125, North
Keppel I, Keppel Bay, central Qld, 23°04'S, | 50°54°E,
Aug 1946; Ipy, AMSC141215, Broome, WA,
17°58'S, 122? 1 4E i Irv, Liv, AMSC 102766, Daintree
River beach, NEQ, 16717'S, 145*27'E, Nov 1928;
Spv, KL, trawled between Palfrey and Lizard Is, NEQ;
9pv, KL, Dingo Beach, central Qld, in littoral sand;
Ipv, trv, KL, trawled to 10m, Little Trunk Reef, NEQ,
Dec 1995; 5pv, KL, Suva, Fiji; Ipv, KL. Bundaberg,
NEQ, littoral sand,

HABITAT AND DISTRIBUTION. Buried in
shallow littoral sand; ranging from Japan, Philip-
pines, New Caledonia, Solomon Islands, Fiji.
Papua New Guinea, Australia from Queensland
to northern Western Australia.

REMARKS. P (Pitarina) pellucidus (Lamarck,
1818) is similar in shape to P. (P.) subpellucidus
(Sowerby, 1851), P. (P) spoori Lamprell &
Whitehead, 1990 and P. (P) nancvae but lacks the
strong, broken radial rays of these species while
the wide, shallow pallial sinus separates tL [rom
these and other Australian Fitar species.

293

Pitar (Pitarina) potteri Healy &Lamprell,
1992 (Fig. 4F-J; 9C)

Pitar (Pifarina) sp. Lamprell & Whitehead 1992: på.
67, sp. 515 (as Pitar spå; Pitar (Pitarina) potter
Healy & Lamprell, 1992: 79; fig. lech Lamprell &
Stanisic, 1996: 45, fig. Get

DIAGNOSIS. Shell length to 38mm, sold, in-
flated. Sculpture of microscopic concentric
growth striae. Colour cream-white with faint tan
zigzag patterns near umbones; interior white.

MATERIAL EXAMINED. HOLOTYPE. ipm.
QMMO32902, Palm I, NEQ, 18?40'S, 146*35'E,
trawled in 9-12m in sand and rubble, Nov 1990,
PARATYPES: Ipv, AMCIA2109, Broadhurst Reef
east of Townsville, NEQ, 18?57' S, 147°47'E, subtidal
27-28 Sept 1975; Ipv, AMSC105246, Watsons Bay,
Lizard 1, NEO, 14*40"S5, 145*27'E, 10.5m on sandy
bottom, Dec 1975; Ipv, AMSCIA2108, Rudder Reel,
NE Port Douglas, NEO, 16°13°S, 145?40' E, intertidal,
1977; tpv. AMSCOS621, Lizard I, NEQ, 14°40°S
]45"28'E; 2pv, WAM560-91, MVF60433, Palm I,
NEO. 18°40'S, 146?35'E, subtidal, Nov 1990.
OTHER MATERIAL, 4pv, Irv, KL, dredged Taylor
Reef, NEQ, 10m, coral sand, 1995; Ipv, KL, dredged
in lagoon, Slashers Reef No. 1, GBR, Nov 1993; 4pv.
Irv, AMSC086261, Noumea, New Caledonia,
22°18'S, L66°29"E, among corals and rocks; Irv,
AMSC1(13582, Palfrey I, Lizard 1, 14°41°S, 145°27°E,
NEQ, in shell sand, 3 Dec 1974; Zrv, liv,
AMSOMISTL, Lizard I, GBR, 14740'S, 145"28' E, 18
Jul 1916; Lv, AMSC1 26664, Eagle I, beach W side,
Lizard I, GBR, 14°42'S, 145923'E, 12 Dec 1974.

HABITAT AND DISTRIBUTION, Reef areas, in
shell sand and rubble to 12m; ranging from north-
ern Queensiand to New Caledonia.

REMARKS. F (Pitarina) potteri Lamprell &
Whitehead, 1992 is similar to P. (P.) affinis
(Gmelin, 1791) but does not attain the size of that
species, is more inflated in proportion and has a
more rounded and ascending pallial sinus.

Pitar (Pitarina) trevori Lamprell &
Whitehead, 1990 (Fig, 5A-G; 9D)

Pitar (Pirarina) trevori Lamprell & Whitehead, 1990:
47. 4a-d; Lamprell & Whitehead, 1992: pl. 67, sp
517,

DIAGNOSIS. Shell length to 30mm, moderately
inflated; lunule incised. Sculpture of fine concen-
irie siriae, periodically crowded as growth
pauses, Colour all white; periostracum, pale
straw coloured, usually with sand adhering to the
external surface.

294 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A-G, P. (Pitarina) trevori Lamprell & Whitehead. A, B, holotype, QMMO22850, Dingo Beach, central
Qld, 30mm. A, external lv; B, internal rv. C, D, KL, Turtle Is, Fiji, 30mm. C, external lv; D, internal rv. E, F,
AMSC107878, Prince of Wales Is, north Qld, 29.3mm. E, external lv; F, internal rv. G, AMSC070200, Dingo
Beach, central Qld, with siphon sheath attached, external pv, 31.3mm (including sheath). H-L, P. (P.)
queenslandica sp. nov., holotype, AMSC306560, Black Is, NEQ, 21.1mm. H, external lv; I, internal rv; J,.
internal rv, enlargement; K, L, dorsal view of pv showing lunule and escutcheon. M-O, P.(P.) subpellucidus
(Sowerby) KL, St Crispins Reef, NEQ, 33mm. M, external lv, N, internal rv; O, variation showing broad
coloured rays, and with siphon sheath attached, external pv (length 43.9mm including sheath).

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

MATERIAL EXAMINED. HOLOTYPE: Ipv,
0MM022850, on sand flats at low tide, Dingo Beach,
central Qld, low tide, Jul 1987. PARATYPES, Ipy,
MVF57679, same data as holotype; lpv,
AMSC160471,same data as holotype: Lpv, WAM939-
89, on sand flats at low tide, Gove, NT, KL, Aug 1987,
OTHER MATERIAL: Ipv, AMSC41611, Annan
River near Cooktown, NEQ, Aug 1916; 1pv, 3rv, 1v,
AMSCIOT878, N of Terry Beach, W side of Prince of
Wales 1, Torres Strait, 10742' S, 142^ T'E, Jun-Jul 1976;
Arv. 21v, AMSC306542, Lindeman 1, central Qld,
20°27°S, 1497"02'E, Nov 1934; 3rv, 2lv,
AMSC306537, between Lindeman and Shaw Is, Ken-
nedy Sound, central Qld, 20^25'8, 14901 'E ; 2pv,
AMSC070200, Cape Gloucester, Dingo Beach,
20°05'S, 148°30°E, central Qld; 4pv, KL, Kurritnine
Beach, NEQ: ipv. AMSCI41215, Broome, northern
WA, 17°58'S, 1227 14'E ; 2pv, KL, Turtle I, Fiji.

HABITAT AND DISTRIBUTION. Buried in
shallow littoral sand; ranging in Australia from
central Queensland to the Northern Territory.
northern Western Australia to Fiji.

REMARKS. Pitar (Pirarina) rrevori Lamprell &
Whitehead. 1990 occurs sympatrically with P
(LÉI pellucidus (Lamarck, 1818) but that species
differs in being more inflated, having an incised
lunule and completely lacking ihe umbonal eol-
ouring and umbonal spot of that species.

Pitar (Pitarina) queenslandica sp. nov.
(Fig. SH-L; 9E)

ETYMOLOGY. For Queensland,

DESCRIPTION. Shell length to 22.3mm, sub-
irigonal, equivalve, inequilateral (the umbones
situated forward of centre), thin; umbones pro-
sogyrate: lunule well developed, heart-shaped,
striate, raised centrally, defined by à faint im-
pressed line; anterodorsal margin shorr, almost
straight, steeply sloping, convex terminally:
posterudorsal margin. long, slightly convex, stee-
ply sloping, convex ta slightly truncate posteri-
Orly; ventral margin evenly convex. Shell
sculpture of well defined concentric striae, obso-
lete umbonally, stronger marginally. Periostra-
cum, thick, grey with aglutinated sand adhering
ta che shell margins. Ligament, impressed. Hinge
of Iv with anterior lateral tooth well developed,
peg-like: anterior cardinal thin, joined to thick
median cardinal forming an inverted v-shape;
posterior cardinal long, thin, ridge-like, free.
Hinge of rv with hinge line expanded ventrally at
pit for the reception of the Iv lateral tooth; anterior
cardinal, narrow; median cardinal free, stighily
oblique; posterior cardinal bifid, oblique. Muscle

295

attachment scars well defined; anterior adductor
scar teardrop-shaped, posterior adductor scar
larger and more rounded. Pallial sinus extends
medially, one third ofthe shell length, moderalely
wide, rounded terminally, Shell colour internally
and externally glossy white.

MATERIAL EXAMINED. HOLOTYPE: (pv,
AMSC306560, Black I, Langford Reef, central Qi,
20"05'8, 148"54'E, dredged, 18m, Nov 1969. Length
of shell 21.1 mm. height 18.4mm, width 1365mm,
PARATYPES. 4pv, 2lv, Irv, AMSC96704, cá
off Lindeman I, central Qld, 1935; 2lv, 6rv,
AMSC306534, SE of Sarina, central Qld, 21°47'S,
150?34'E. dredged 59m, Dec 1977; AMSCS58758 (pt),
dredged off Lindeman I, central Qld, Aug 1935; Ipv,
QMM059380. Gulf of Carpentaria, 1272Y'S,
141*35.2'E, 13m. OTHER MATERIAL: trv,
AMSC306556, off Caloundra, southern Qld, 26"47'5,
153°3F E, 128m, Mar 1969; 3lv, 2rv, AMSCSS758
(pt), off Lindeman I, central Qld, Aug 1935;
AMSC306533, E of Sarina, central Qld, 21°28'S,
150"08.5'B, sandy mud, Nov 1977; llv,
AMSC306557, Caloundra, southern Qld, 26*49'S,
153*9"E, beach, Oct 1963; Ziv, Ar, AMSC306562.
Caloundra, southern Qld, 26°49'S, 153? 10 E, beach;
Oly, 6rv, AMSC306540, E of Broad Sound, southerts
Qld, 21°58.5'S, 150^45" E, 57m. in coarse to fine sand
and mud, Nov 1977; many juveniles. AMSCO29671,
Cairns Reef, NEO, 15°42’S, 142? MY E, 9-18m, Aug
1906; 2rv, lly, AMSC306559, off Cairns, NEO.
L6*51.6S, 146^1.2'E, 33-34m, sandy-mud and shell,
Oa 1981; 21v, Ire, AMSC306532, E of Mackay, een-
teal Qld, 209525, 149°29°E, 35m 1977; (px,
AMSC093522, Point Cartwright, southem Qld,
26?41'5, 153?DR'E, 1963: Ap, Ipv. KL, Llewellyn
Bay, Armstrong Beach. Sarina, central Qld, after
storm, Apr 1995; KL, Gulf of Carpentaria, NEQ,
11730 2'8, 139°03.2'E, 523m.

HABITAT AND DISTRIBUTION, Dredged in
silty sand to 128m; appears to be restricted to
Queensland.

REMARKS, P (Pirarina) queenslandica sp. nov.
is a small species, the largest specimen sighted,
AMSC306556 is 22.3mm. It is most like A/F}
rrevori Healy & Lamprell, 1992 which is also
glossy white with a subtrigonal shell, However P.
(P) queenslandica is readily separated by ils
much smaller size, ventrally expanded hinge line
(straight in P (P.) trevori) and much deeper pallial
sinus. Compared with P (P ) pellucidus (Lamarck,
1818), P (P) subpellucidus (Sowerby, 1851), R
(BI lineolatus (Sowerby, 1854) and £F
(Hyphaniosoma) limarula (Sowerby, 1853), D
(P.) queenslandica does not artain the size of these
species, has a more extended pallial sinus and
does not exhibit a colour pattern (uniformly

296 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6. A, B, P. (Pitarina) japonicus Kuroda & Kawamoto, AMSC095267, south of Yeppoon, Qld, 37.6mm. A,
external lv; B, internal rv. C, D, P. (P.) noguchii Habe, AMSC121360, Roebuck Bay, Broome, northern WA,
37.3mm. C, external lv; D, internal rv. E, F, P. (P.) nipponica Kuroda & Habe, AMSC306563, Shelley Beach,
Townsville, north Qld, 27.3mm. E, external lv; F, internal rv. G, H, P. (P.) sophiae (Angas) , KL, off Caloundra,
Qld, 28.9mm. G, external lv; H, internal rv. I-K, P. (P.) thornleyae sp. nov., holotype , AMSC113277, Karumba,
Gulf of Carpentaria, north Qld, 46.8mm. I, external lv; J, internal rv (broken ventro-posterior margin); K, dorsal
view pv showing lunule and escutcheon.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

glossy white in P (P) queenslandica). Compared
with P. (E) coxeni (Smith, 1885) and P. (P)
sophiae (Angas, 1877) which are also small in
size and approximately similar in shape, P (P.)
queenslandica does not have the colour pattern
usually evident in P. (P) sophiae while the pallial
sinus is deeper in P. (FA) queenslandica than in
both P. (P) sophiae and P(P.) coxeni (sinus angu-
late in P. (P.) coxeni),

Pitar (Pitarina) subpellucidus
(Sowerby, 1851) (Fig. 5M-O; 9F)

Cyrherea subpellucidus Sowerby, 1851: 639. pl. 133,
f. 136; Diune tarresica Reeve, 1863: pl. 11, spå 51;
Pitar (Pitarina) subpellucidus (Sowerby) Habe,
1977: 257, Lamprell & Whitehead, 1992; pl, 67, sp.
517.

DIAGNOSIS. Shell length to 40mm, elongate
ovate, moderately inflated. Sculpture of fine con-
centric striae. Colour cream with dense brown
radial rays, purple posteriorly; umbones brown
with purple spots beneath; interior tinged purple
posteriorly.

MATERIAL EXAMINED, KL, 1pv, Barber I, NE
Palm I, NEQ; KL, Ipv, St Crispins Reef, NEQ; KL,
L py, Low I, NEQ; KL, Ipv, Cobham Reef, near Gould
Reef, central Qld; KL, Ipv, Palm I, NEQ, dredged in
10m; KL, 4pv, Lizard I, between Palfrey and Eagle I,
NEQ, 29m.

HABITAT AND DISTRIBUTION. Dredged to
1 Om in coral and littoral sand, coastal and GBR;
ranging from Japan to Queensland, Australia,

REMARKS, This is not a common species but
appears lo be widely distributed, further collect-
ing should increase the known range of the spe-
cies, P, (P) subpellucidus (Sowerby, 1851) is
similar in shape to P. (P) pellucidus (Lamarck,
1818) bul is readily distinguished by its more
highly coloured shell.

Pitar (Pitarina) japonicus Kuroda &
Kawamoto,1956 (Fig. 6A,B: 9G}

Pitar (Agriapoma) japonicum Kuroda & Kawamoto,
1956: 89, figs 1-4; Pitar(Pitarina)japonicum Habe,
1977: 257; Lamprell & Whitehead 1992: pl. 67, sp.
522; Lamprell & Stanisic, 1996: 44, fig, Sg,h.

DIAGNOSIS, Shell length to 35mm; thin, ovate-
trigonal, umbones prosogyrous. Sculpture of fine
concentric growth lines. Colour cream-white,
with pale orange or pink umbones and Interior,

297

MATERIALEXAMINED. 2pv, AMSC 106921, South
Great Keppel I, central Qld, 23°12'S, 150*58' E.; 2rv,
21v. AMSC093524, Keppel Bay, central Qld, 23*25'8,
150°5S'E; Ipv, AMSC095267, Zilzie Point, central
Qld, 23°17°S, 150*50' E, sheltered mud fløts; 1pv, KL,
Lammermoor Beach, central Qld, after storm; 1pv, KL,
Boyne Is Beach, central Qld, low tide; Ipv, KL, Dingo
Beach, central Qld, low tide; Irv, AMSC142126, Pen-
guin Channel, GBR, 16°15°S, 145?31'E, 18-28.5m;
lpv, Irv, KL, Shelburne Bay, NEO, 12°41 3'5,
14042,5'E, Jan 1993; ipv, AMSC092622, Groote
Eylandt, Gulf of Carpentaria, 14*00' 8, 136?25'E, Oct
1973; Irv, AMSC306565, 32km off Point Charles,
Darwin, NT, 12710'S, 130°22°E, 237m, Jul 1938.

HABITAT AND DISTRIBUTION. Buried in
tidal sand and mud; rangifig from Japan, New
Caledonia, central Queensland to northern Aus-
tralia.

REMARKS. P. (Pitarina) japonicus Kuroda &
Kawamoro,1956 is reasonably common through-
out its range. Recent records from New Caledonia
mark the southern limit of its distribution.

Pitar (Pitarina) noguchii
Habe, 1958 (Fig. 6C,D; 9H)

Pitar (Agnopoma) noguchii Habe, 1958- 34; Pitar
(Pitarina) noguchii Habe, 1964: pl. 58, fig. 6; Habe,
1977: 257; Lamprell & Stanisic, 1996; 42, fig. 4g-h.

DIAGNOSIS. Shell length to 37.3mm, thin, in-
flated. Sculpture of fine concentric ridges and
iregular blue growth pauses, with small puncture
marks over surface. Colour white internally and
externill y.

MATERIAL EXAMINED. In, AMSC121360, Roc-
buck Bay, Broome, northern WA, 18°00'S, 122^15'E.;
Ipv, AMSC13871, North Keppel Island, Keppel Bay,
central Qld, 23°04'S, 150*54'S, 1970: Ipv,
AMSC308648, Phili pineg, purchased from Sowerby
and Fulton, presented y C. Hedley,

HABITAT AND DISTRIBUTION. Dredged in
sand to 150m; ranging from Japan to New Cale-
donia and Australia [rom Queensland to north
Western Australia.

REMARKS. Hitherto only recorded from Japan,
recent collecting has extended the known distri-
bution to New Caledonia (Lamprell & Stanisic,
1996), These appear to be the first records of the
species from Australia which is a significant
range extension,

Pitar (Pitarina) nipponica Kuroda & Habe.
1971 (Fig. 6 EF; 91)
Pitar (Pitarina) nippenica Kuroda & Habe, 1971;

Habe, 1977: 257; Lamprell & Whitehead, 1992: pl.
61. sp. 531.

DIAGNOSIS. Shell length to 23mm; subtri gonal.
Sculpture of fine concentric growth striae; sur-
face covered with small, random puncture marks.
Colour white, externally; interior white, yellow
centrally.

MATERIAL EXAMINED, HOLOTYPE: lpv,
NSMT, MOR9601, Amadai-ba [bank], or off Hayama,
Sagami Bay, [Kanagawa, Japan] 1971. OTHER MA-
TERIAL: 3lv, AMSC093520, South West Racks, Trial
Bay, central NSW, 30°53'S, 15304'E, beach, Aug
1960; Ip, AMSC306545, off Tweed Heads, north-
eastern NSW, 28"08'-28"12'5, 153"40'-153*45'E,
54m, Oct 1936; 4rv, 21v, AMSC054657, Caloundra
beach, southeastem Qld, 26°49°S, 153*1Q'E, May
1926; Lpv, KL, Pallarenda Beach, Townsville, NEQ,
low tide; 3py, KL, AMSC306563, AMSC142127,
Shelley Beach, Townsville, NEQ, |9?1]'S, 146^45'E,
low tide, Aug 1970; Ipv, KL, Exmouth Gulf, WA; Lpv,
KL, Arafura Sea, Bureau of Rural Resources, 1990,

HABITAT AND DISTRIBUTION. Buried in lit-
loral sand and to 54m; ranging from Japan to
Queensland and north Western Australia.

REMARKS. Pf(Pitarina) nipponica Kuroda &
Habe, 1971 is similar to P. (P. ) japonicum Kuroda
& Kawamoto, 1956 hut is smaller, has more
lightly coloured umbones and random puncture
marks on the external surface,

Pitar (Pitarina) sophiae (Angas, 1877)
(Fig. 6G-H; 10A)

Cytherea sophiae Angas, 1877: 176, pl.26, £.23; Pitar
(Pitarina) sophiae (Angas) Lamprell & Whitehead,
1992: pl. 68, sp. 531; Lamprell & Stanisic, 1996; 45,
fig. 6c-d.

DIAGNOSIS. Shell length to 17mm, thin, in-
Hated. Sculpture of concentric growth striae, sur-
face smooth, glossy. Colour white, with prey
concentric bands, sometimes with faint brown
broken rays: umbones usually tinged with red
anteriorly, visible internally,

MATERIAL EXAMINED. 2rv, AMSC306495,
Manly Beach, Sydney, NSW, 33?48'S, 151*17'E ;
Ipv, AMSC306544, ENE of Point Danger, northern
NSW, 27°04’-27°59'S, 15350"'E, 137-146m; Ipv,
AMSC007545, inner North Head, Port Jackson, Syd-
ney, NSW, 33?49'S, 151"17'E,15m, in sand; Ir,

MEMOIRS OF THE QUEENSLAND MUSEUM

AMSC306571, Christmas 1, off NE point, 10725' S,
105*42'E, ! 83m.

HABITAT AND DISTRIBUTION. Dredged to
183m in sand; ranging from New South Wales,
central to northern Queensland, Australia, Christ-
mas I. to New Caledonia.

REMARKS. Smith (1885) considered D
(Pitarina) sophiae (Angas, 1877) to be a syn-
onym of Cytherea hebraea Lamarck, 1818. How-
ever comparison of specimens of P (P.) sophiae
in the Australian Museum and the holotype of P
(P) hebraea in the Museum de Geneve shows
that P (P) sophiae is longer in proportion to
height than £ (P) hebraea, is more widely
rounded at the anterior and posterior margins,
does not attain the size of that species and has à
pink flush on the inner side of the valves of many
specimens. Additionally, the pallial sinus in £ ( 7.)
hebraea is wider, deeper and less ascending than
in P. (P.) sophiae.

Pitar (Pitarina) thornleyae sp. nov.
(Fig. 6I-K; 10B)

ETYMOLOGY. For Ms Gertie Thomley.

DIAGNOSIS. Shell length to 46.8mm, elongate
óvate, equivalve, inequilateral, the anterior eid of
shell being 1/3 of the shell length; moderately
inflated, thin; umbones approximate, lunule nar
rowly heart-shaped, striated, surrounded by an
impressed line; anterodorsal margin moderately
convex, sharply sloping, narrowly rounded termi-
nally; posterodorsal margin almost straight,
gently sloping, posterior margin narrowly
rounded; ventral margin convex, Sculpture of
course, irregular concentric ridges and growth
pauses; periostracum grey, with sand adhering to
most of shell surface. Ligament, narrow, partially
external, Hinge of ly with anterior lateral tooth
narrow, peg-like, moderately thick; anterior car-
dinal, slightly oblique; median cardinal solid,
broad, raised; posterior cardinal long, oblique;
posterior lateral, long, thin, parallel to the nymph.
Hinge of rv with paired anterior laterals. sepa-
rated by à deep pit; anterior cardinal thin, raised;
median cardinal thin, not joined, parallel to ante-
nor cardinal, separated by à narrow pit; posterior
cardinal oblique and bifid; posterior lateral long,
solid, parallel to the nymph, Muscle adductor
scars well defined, anterior adductor scar elon-
gate-ovate, posterior adductor scar teardrop-
shaped, Pallial line ragged. Pallial sinus
extending one third medially, triangular, Colpura-
tion off-white externally, internally white.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS 299

FIG. 7. A-E, P. (Pitarina) lineolatus (Sowerby). A, B, KL, Torres Strait, 24mm. A, external lv; B, internal rv. C,
D, KL, Palm Is, NEQ, juvenile, 17mm. C, external lv; D, internal rv. E, KL, Torres Strait, external lv, 24mm.
F, G, P. (P.) bullatus (Sowerby), AMSC141214, Broome, northern WA, 36.2mm. F, external lv; G, internal rv.
H, I, P. (P.) sulfureum Pilsbry, KL, Kurrimine Beach, NEQ, 28.9mm. H, external lv; I, internal rv. J, K, P. (P.)
coxeni (Smith), KL, Shelburne Bay, NEQ, 13.5mm. J, external lv; K, internal rv. L, M, P. (P.) inconstans
(Hedley), AMSC073218, Madang, PNG, 22.1 mm. L, external lv; M, internal rv. N, O, P. (P.) regularis (Smith),
AMSC306561, Magnetic Is, NEQ, 15.5mm. N, external lv; O, internal rv.

300

MATERIAL EXAMINED, HOLOTYPE: Ipv,
AMSCI 13277, off Karumba, SE Gulf of Carpentaria,
northern Qld, 17°30'S, 140°00-141°00'E, 1963-4.
Length of shell 46.8mm, height 36.4mm, width of
conjoined valves 26.8mm. PARATYPE: rv, liv,
AMSC308642, Gulf of Carpentaria, northern Qld,
1980; Irv, AMSC74865, Gulf of Papua, PNG, Feb
1969; Ipv, AMSC14952, Karumba, northern Old,
17°29°S, 140*50'E, beach, Jun 1903, 2pv,
AMSC107015, AMSC107028, off Albert River, Gulf
of Carpentaria, northern Qld, 17°24'S, 139?47' E, Dec
1963; Ipv, AMSC107296, W of Topsy Creek, E Gulf
of Carpentaria, northern Qld, 15°28'S, 141°28.7°E,
Dev 1876; Ipv, AMSCII5732, 7km from Norman
River mouth, Karumba, northern Qld, 17^29'S,
140°50' E, washed from river bank; AMSC 1 17072, ot!
Karumba, SE Gulf of Carpentaria, northern. Qld,
17930'8, 140*00"-141*00"E.

HABITAT AND DISTRIBUTION. Buried in
sandy mud near mouths of rivers; ranging from
northern Queensland to Papua New Guinea.

REMARKS, Pitar (Pitarina) thornleyae sp, nov
is most similar to P. (PF) rrevori Healy &
Lamprell, 1990, P. (P) pellucidus (Lamarck,
1818) and F (FP) subpellucidus (Sowerby, 1851)
in being elongate-ovate and moderately inflated
but is readily separated by its larger size, much
stronger concentric ridges, longer pasterodorsal
margin, triangular shaped pallial sinus (wide and
short in P. (Pl pellucidus and P. (P.) sub-
pellucidus, short and ascending in P. ( P.) rrevori),
There are no other known Australian Pitar spe-
cies which have the strong concentric ridges of P
(P.) rhornleyae.

Pitar (Pitarina) lineolatus (Sowerby, 1854)
(Fig. 7A-E; LOC)

Cytherea lineolatus Sowerby, 1854: pl, 168, figs 214-
5; Pitar (Pitarina) lineolatus Habe, 1977: 257;
Lamprell & Whitehead 1992: pl. 67, sp. 524.
DIAGNOSIS, Shell length to 24mm, ovate.

Sculpture of fine concentric striae. Colour cream

usually with zigzag markings on the escutcheon,

prae, and sometimes faintly on the rest of the
shell,

MATERIAL EXAMINED. lpv, KL, Torres Strait,
northern Qld, dredged to 15m; 2pv, KL, Palm I, NEQ,
dredged in Curacoa Channel to 15m; liv, KL, Little
Trunk Reef, NEQ, dredged to 10m, Dec 1995.

HABITAT AND DISTRIBUTION. Dredged in
sand to 16m; ranging from Japan, to northern
Queensland and the Torres Strait, Australia,

MEMOIRS OF THE QUEENSLAND MUSEUM

REMARKS. Piar — (Pitarina) ` lineolatus
(Sowerby, 1854) is similar to P. (PI limatula
(Sowerby, 1853) but readily separated by the
cream-white colour and tan zigzag pattern on the
lunule and dorsal margin (yellow coloured with
black staining on lunule and escutcheon on P(A.)
limatula),

Pitar (Pitarina) bullatus (Sowerby, 1851)
(Fig. 7F,G; 10D)

Cytherea bullata Sowerby, 1851: sg 96, nl. 126,
fig. 192; Lamprell & Whilehead 1992; pl, 67, sp.
331.

DIAGNOSIS. Shell length ta 36.2mm, solid,
broadly subtrigonal, inflated. with large inflated
umbones, Sculpture of fine concentric striae and
growth lines. Colour chalky while externally;
interior white.

MATERIAL EXAMINED. 17pv, KL, Turkey Beach,
central Qld, on sand/mud flats low tide; | pv, KL, Dinga
Beach, central Qld, sand flats low tide, | pv, KL, Glad-
stone, southern Qld, on sand flats, Jan 1976, | P KL,
Dampier, northem WA, 1987, 1pv, AMSC 141214,
Broome, northern WA, 17958'5, |22?14'E; 4pv,
AMSC051544, AMSC093530, Roebuck Bay,
Broome, northern WA, 18*00'8, 122° 1 5E; 1pv, KL,
Black Ledge, Broome, northern WA, on mud/sand
flats, 1994; 4pv, KL, Broome, northern WA, on
sand/mud fats.

HABITAT AND DISTRIBUTION. Buried in
shallow littoral sand/mud flats; ranging in Aus-
tralia from Queensland, Northern Territory to
northern Western Australia,

REMARKS. Pitar (Pitarina) bullatus (Sowerby,
1851) can be readily separated from other Austra-
lian Pirar species by its solid, chalky, elongate-
ovate shell and straight ventral margin,

Pitar (Pitarina) sulfureum Pilsbry, 1904
(Fig. 7H,I; 10E)

Pitar (Pitarina) sulfureum Pilsbry; 1904: 553. pl, 39;
Lampretl & Whitehead 1992: pl. 68, sp. 532,

DIAGNOSIS. Shell to length 30mm, light, in-
flated; almost chalky; escutcheon ill-defined but
protrusive. Sculpture smooth, indistinct, line
striae. Colour yellow-cream; umbones yellow,
sometimes tinged pink; interior cream-white,
light-yellow centrally.

MATERIAL EXAMINED, 2ly, AMSC306570, SW of
Losuia, Kiriwinna I, Trobriand Group, PNG, 8°33"S,
151*04'E, on sand flats, Jun 1970; 2pv, AMSC306552,
SE of Exmouth Homestead, Exmouth Gulf, northern

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

FIG. 8. Comparative hinge plate and teeth of rv. A, P. (Hyphantosoma) intricata (Dautzenberg), AMSC123340,
52.5mm; B, P.(H.) spoori Lamprell & Whitehead, holotype, AMSC133572, Bank Reef NEO, 31mm; C, P. (H.)
limatula (Sowerby), KL, Dingo Beach, central Qld, 24.2mm; D, P. (H.) caperi sp. nov., holotype,
AMSC202973, Upolu Cay, NEQ, 22.8mm; E, P. (P.) citrina (Lamarck), KL, Shark Bay, northern WA,
45.4mm; F, P. (H.) nancyae Lamprell & Whitehead, off Caloundra, Qld, 28.6mm; G, P. (P.) osmunda (Iredale)
AMSC303944, Sydney Harbour, 36.5mm; H, P. (P.) curnowae sp. nov., holotype, AMSC202974, Madang,
PNG, 42.6mm; I, P. (P.) prora (Conrad) KL, North West Is, Qld, 63mm; J, P. (P.) marrowae Healy & Lamprell,

QMMOJ32900, Dingo Beach, central Qld, 38.1 mm.

302

WA, 22?24' S, 114*08' E, on muddy sand flats; 2pv.
KL, Kurrimine Beach, NEQ, low tide on sand.

HABITAT AND DISTRIBUTION. Buried in
loose shallow muddy sand flats; ranging from
Japan to Papua New Guinea and Queensland.

REMARKS. Pitar (Pitarina) sulfureum Pilsbry,
1904 is recognised hy its rough surface and dense,
fine concentric striae and chalky yellow-cream
appearance. In these respects it is most similar to
P. (P.) bullatus (Sowerby, 1851) but differs by
having a convex ventral margin (almost straight
in P. (FP) bullatus). P. (P.) bullarus also has å rough
surface and dense, fine concentric striae and
chalky white appearance, However P. (P.) sul-
fureum differs from £. (P.) bullatus by having a
convex ventral margin (almost straight in P. (P.)
bullatus). P. (P.) sulfureum appears 10 be a rare
species in Australia with few specimens either in
the Muséum or private collections.

Pitar (Pitarina) coxeni (Smith, 1885)
(Fig. 71, K; 10F)

Cytherea (Caryatis) coxeni Smith, 1885: 139, pl. 1, figs
7-Te; Pitar (Pitarina) coxeni Lamprell & White-
head, 1992: pl, 68, sp. 530.

DIAGNOSIS. Shell length to | 3mm, thin, ovate,
moderately obese. Sculpture of fine, close, dis-
tinct concentric striae, often polished. Colour
white, internally and externally; obscure brown
or red lines and dots or pale growth lines some-
limes occur.

MATERIAL EXAMINED. Løv, KL, Whitsunday Pas-
sage, central Old, trawled to 10m; 2pv, KL, Shelburne
Bay, NEQ, 12°39.4'S, 141°09°E, 41m. Jan 1993,

HABITAT AND DISTRIBUTION. Dredged in
sand ta 25m; ranging from central Queensland to
the Northern Territory and Papua New Guinea.

REMARKS. Pitar (Pitarina) coxeni (Smith,
1885) is very fragile and rare even in Museum
collections. P. (P.) coxeni is most similar to P. (P.)
sophiae ( Angas, 1877) in size, shape and fragility.
but difters in having a more attenuate posterior
margin and angulate pallial sinus (posterior mar-
gin widely convex and pallial sinus widely
rounded in P. (P.) sophiae),

Pitar (Pitarina) inconstans (Hedley, 1923)
(Fig. 7L,M; 10H}

Pitar (Pitarina) inconstans Hedley, 1923: 304:
Lainprell & Whitehead, 1992: pl. 68, sp. 529.

MEMOIRS OF THE QUEENSLAND MUSEUM

DIAGNOSIS, Shell length tọ 20mm, inflated,
subtrigonal, posteriorly attenuate, Sculpture fine,
almost polished, concentric growth lines. Colour
white, sometimes with grey growth bands; inte-
nor white.

MATERIAL EXAMINED. ipv, AMSCO73218,
Madang, PNG, 5*02.5'S, 145"7'E, 3pv, KL, Buffalo
Creek, Darwin, NT, 1984; many, KL, Nanum Beach,
Weipa, NEO, Jul 1992, N, Trevor: 3pv, KL,
Numbulwar, Gulf of Carpentaria, NT, 1984.

HABITAT AND DISTRIBUTION. In shallaw littoral
sand; ranging in Australia from north Queensland and
the Northern Territory to Papua New Guinea and Ma:
laysia.

REMARKS. Pitar (Pírarina) inconstans
(Hedley, 1923) is quite variable in shape and
remarkably similar to the mactrid Notespisula
trigonella (Lamarck, 1819) which is equally vari-
able in shape but is readily separated from that
species by the hinge teeth.

Pitar (Pitarina) regularis (Smith, 1885)
(Fig. 7N,O; 10G)

Cytherea ( Caryatix) regularis Smith, 1885. 140, pl. 1,
figs 8-8b; Pitar (Pitarina) regularis Lamprell &
Whitehead, 1992: pl. 68, sp. 528.

DIAGNOSIS. Shell length to 19mm, com-
pressed. Sculpture of close-set concentric ridges,
slightly lamellate posteriorly. Colour off-white
with rust-brown concentric bands.

MATERIAL EXAMINED. Ipv, AMSC306561,
Horseshoe Bay, Townsville, NEO, 19°06.8'S,
146751.8'E, Sep 1980; 2pv, 21v, 2rv, AMSC306335,
off Broad Sound, central Qld, 22"06'S, 150749 E.
53m, among coarse shell, Dec 1977; lv, AMSC306531,
off Cairns, NEO, 16?55'S, 146707' E, 37-40m, sandy
shell and mud; 2pv (juvenile), AMSC306530, Arafura
Sea, NE of Croker I, NT, 10°36'S, 132*56,5' E, 62m;
ipv, KL, Boyne L Qld, beach; | pv, KL, Gulf of Car-
pentaria, NT, Bureau of Rural Resources, 1990.

HABITAT AND DISTRIBUTION. In littoral
sand and dredged in sandy mud and coarse shell
to 62m; ranging from Hong Kong, West Malaysia
to Australia, north Queensland, Torres Strait,
Northern Territory and north Western Australia.

REMARKS. Pitar (Pitarina) regularis (Smith,
1885) ts most similar to P. (P.) bullatus (Sowerby,
1851) in being solid, having a rough, chalky
surface, fine concentric growth lines and white
colour. P (F.) regularis differs in the ventral mar-
gin being convex (straight in P. (P) bullatus),

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS 303

FIG. 9. Comparative hinge plate and teeth of rv. A, P. (Pitarina) affinis (Gmelin) KL, Shark Bay, northern WA,
55.8mm; B, P. (P.) pellucidus (Lamarck) KL, Dingo Beach, central Qld, 34.4mm; C, P. (P.) potteri Healy &
Lamprell, holotype, QMMO32902, Shelly Beach, NEO, 38mm; D, P. (P.) trevori Lamprell & Whitehead,
holotype, QMMO22850, Dingo Beach, central Qld, 30mm; E, P. (P.). queenslandica sp. nov., holotype,
AMSC306560, Black Is, NEQ, 21 mm; F, P. (P.). subpellucidus (Sowerby) KL, St Crispins Reef, NEQ, 33mm;
G, P. (P.) japonicus Kuroda & Kawamoto, AMSC095267, south of Yeppoon, Qld, 37.6mm; H, P.(P.) noguchii
Habe, AMSC121360, Roebuck Bay, Broome, WA; I, P. (P.) nipponica Kuroda & Habe, AMSC306563, Shelley
Beach, Townsville, NEQ, 27.3mm.

304 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 10. Comparative hinge plate and teeth of rv. A, P. (Pitarina) sophiae (Angas) , AMSC007545, Port Jackson,
NSW, 23.2mm; B, P. (P.) thornleyae sp. nov., holotype, AMSC113277, Karumba, Gulf of Carpentaria, NEQ,
46.8mm; C, P. (P.) lineolatus (Sowerby), KL, Torres Strait, 24mm; D, P. (P.) bullatus (Sowerby),
AMSC141214, Broome, northern WA, 36.2mm; E, P. (P.) sulfureum Pilsbry, KL, Kurrimine Beach, NEQ,
length 28.9mm; F, P. (P.) coxeni (Smith), KL, Shelburne Bay, NEQ,13.5mm; G, P. (P.) regularis (Smith),
AMSC306561, Magnetic Is, NEO. 15.5mm. H, P. (P.) inconstans (Hedley), AMSC073218, Madang, PNG,
22.1mm.

NEW VENERIDAE FROM AUSTRALIAN AND ADJACENT WATERS

having coarser concentric lines and is less in-
flated than that species.

ACKNOWLEDGEMENTS

The authors extend their thanks to Mr I. Loch
(Australian Museum) for allowing us to examine
and borrow material for this project and to Dr H.
Saito (National Science Museum, Tokyo) for or-
ganising loan of the holotype of the holotype of
P. (P.) nipponica from the Imperial Collection.
Mrs G. Curnow (South Australia) and Mr P. Spoor
(Townsville) are thanked for donating or lending
specimens for study. Special thanks to Dr John
Stanisic for reading the manuscript and valuable
advice. This work was supported by a Keith Suth-
erland Award to KL. An Australian Research
Fellowship and a Senior Research Fellowship
from the Australian Research Council made pos-
sible the participation of JH.

LITERATURE CITED

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twenty-five species of marine shells from New
South Wales. Proceedings of the Zoological Soci-
ety London 1877: 171-177.

CONRAD, T.A. 1837. Descriptions of new marine
shells from upper California collected by Thomas
Nuttal, Esq. Journal of Academy of Natural Sci-
ences, Philadelphia 7: 227-268.

DALL, W.H. 1902. Synopsis of the family Veneridae
and of the North American recent species. Pro-
ceedings of the U.S. National Museum 26: 335-
411.

DAUTZENBERG, P. 1907. Description de coquilles
nouvelles de diverses provenances et de quelques
cas tératologiques. Journal de Conchyliologie 55:
327-341.

GMELIN, J.F. 1791. Caroli a Linne, Systema Naturae,
edition 13 (6): 3244-3289.

GRAY, J.E. 1838, Catalogue of the species of the genus
Cytherea of Lamarck, with the description of
some new genera and species. The Analyst, Lon-
don 8: 302-309.

GUPPY, R.J.L. 1866. On the Tertiary Mollusca of Ja-
maica. Quarterly Journal of the Geological Soci-
ety of London 22: 281-295.

HABE, T. 1958. Report on the Mollusca chiefly col-
lected by the S.S. Soyo-Maru of the Imperial
Fisheries Experimental Station on the continental
shelf bordering Japan during the years 1922-1930.
Part 4, Lamellibranchiata 2. Publications of the
Seto Marine Biological Laboratory 7: 19-52.

1964. ‘Shells of the Western Pacific in colour’.
Vol.2. (Hoikusha : Osaka). [English edition].

305

1977. Systematics of Mollusca in Japan. Bivalvia
and Scaphopoda. (Zukan-no-Hokuryukan :
Tokyo).

HANLEY, S. 1842-1856. An illustrated and descriptive
catalogue of Recent bivalve shells. (Williams &
Norgate: London).

HARTE. M.E. 1993. A review of Pitar (Hypantosoma)
Dall, 1902. (Veneridae: Pitarinae) and a descrip-
tion of Pitar (H.) festoui sp. nov. The Veliger
36(4); 343-350. .

HEALY, J. & LAMPRELL, K. 1992. New species of
Veneridae, Cardiidae, Crassatellidae, Tellinidae
and Mactridae from Australia (Veneroida,
Bivalvia, Mollusca), Journal Malacological Soci-
ety of Australia 13: 75-97.

IREDALE, T. 1936. Australian molluscan notes No.2,
Records of the Australian Museum 19: 267-340.

JUKES-BROWNE, A.J. 1913. On Callista, Amiantis
and Pitaria. Proceedings of the Malacological
Society of London 10 (6): 335-347.

KEEN, M. 1969. The Veneridae. Pp. 670-689. In R.C.
Moore (ed.) "The treatise on invertebrate paleon-
tology, Part N, Vol. 2 (of 3), Mollusca, 6, Bivalvia'.
(Geological Society of America and University of
Kansas: Lawrence, Kansas).

KURODA, T. & HABE, T. 1971. "The seashells of
Sagamai Bay'. (Maruzen: Tokyo).

LAMARCK, J.B. 1818-22. Histoire naturelle des an-
imaux sans vertebres. Vols. 5, 6a, 6b. (Paris:
Lamarck).

LAMPRELL, K. & STANISIC, J. 1996. Callista,
Lioconcha and Pitar in New Caledonia and adja-
cent waters (Mollusca, Veneridae). Molluscan Re-
search 17: 27-48.

LAMPRELL, K. & WHITEHEAD, T. 1990. Eight new
marine bivalves from Australia (Mollusca,
Bivalvia). Journal of the Malacological Society of
Australia 11: 33-52.

1992. 'Bivalves of Australia”. (Crawford House
Press Pty Ltd: Bathurst, NSW).

PILSBRY, H.A. 1904a. New Japanese Marine
Mollusca. Pelecypoda. Proceedings of the Acad-
emy of Natural Sciences, Philadelphia 53: 550-
561.

PRASHAD, B.1932. The Lamellibranchia of the
Siboga Expedition. Systematics Part 2.
Pelecypoda. Siboga Expedition. Livr. 118. 34:
1-353.

REEVE, L.A.1843-78. 'Conchologia Iconica: or
illustrations of the shells of molluscous animals'.
vols 1-20. (L. Reeve : London).

SMITH, E.A. 1885. Report on the Lamellibranchiata
collected by H.M.S. Challenger during the years
1873-76. Reports on the scientific results of the
voyage of the H.M.S. Challenger (Zoology) 13:
1-341.

SOWERBY, G.B.(II. 1851-1853. ‘Thesaurus con-
chyliorum, or monographs of genera of shells’.
(G.B. Sowerby: London).

A NEW STREAM-DWELLING £/TORIA FROM THE MELVILLE RANGE,
QUEENSLAND, AUSTRALIA

KEITH R. MCDONALD

McDonald, KR 1997 06 30: A new stream-dwelling Liloria from the Melville Range,
Queensland, Australia. Memoirs of the Queensland Museum 42(1); 307-309. Brishane. ISSN

0079-8835.

Litoria andiirrmalin sp. nov. (Anura: Hylidae) is described from streams amongst boulders
in Cape Melville National Park, Queensland. [t is a large, mottled brown species with
unwebbed fingers, extensively webbed toes and distinct supratympanic fold: females have
pigmented eggs, The species is found in boulder areas around riffles in streams. Morpholog-
ically it cannot be referred to any recognised Australian hylid species group. C] Frog,
Hylidae, Litaria, Queensland, Litoria andiirrmalin.

Conservation Strategy Branch, Department of Environment, PO Box 834, Atherton, Queens-

land 4883, Australia; 6 March 1997.

During botanical surveys in Cape Melville Na-
tional Park on Cape York Peninsula, Queensland
in 1993, å specimen of unidentified hylid frog
was caught along a rocky, rainforest stream by
J.P, Stanton, Examination of the juvenile speci-
men indicated it was possibly an undescribed
species, In May 1994 the opportunity to re-visit
the collection site was made possible during a
botanical survey led by D,G. Fell. Adult speci-
mens were collected and the species was con-
firmed as undescribed. The species is described
in this paper. It is a hylid frog of the genus Litoria
having expanded finger and toe discs with a dis-
linet notch, horizontal pupil and no palpebral
venation on the lower eyelids.

Methods of measurement follow Tyler (1968).
Measurements were made using dral calipers
reading to an accuracy of 0.1mm. Measurements
were: S-V. body length from snout to vent; TL,
tibia length; HL, head length; HW, head width;
E-N, eye to naris length; IN, internarial span; E,
eye diameter; T, tympanum diameter. Specimens
are lodged in the Queensland Museum, Brisbane
(QMS), Australian Museum, Sydney (AMR) and
the South Australian Museum, Adelaide
(SAMR).

Litoria andiirrmalin sp. nov.
Literia species, Frith and Frith, 1995: 233.

MATERIAL EXAMINED. HOLOTYPE: QMJ59000
adult 9 , collected by KR. McDonald, J.A. Ledger and
D.G. Fell at 280m, 2 May 1994 at Temple Ck on the
eastern slopes of the Melville Range, Cape Melville
National Park (144°31°E, 14?16'S). PARATYPES:
QMJS9001-008, AMR144391, SAMR44463 same
data as holotype, QMJ59009 collected hy J.P. Stanton
and D.G. Fell, 21July 1993 at type locality.

DIAGNOSIS, A large species (preserved speci-
mens 93- | 02mm N=5; field measurements of live
animals; ? 9 92.3-109.7mm, N= 6; dd 66.1-
75.6mm, Nz9, S-V) characterised by widely ex-
panded finger and toe discs; moderately long
hindlimbs; unwebbed fingers; extensively
webbed toes; absence of dermal glands; strong
supratympanic fold; in life mottled brown with
irregular cream markings and pale dorsolateral
stripe from eye bulge to groin,

DESCRIPTION OF HOLOTYPE. Head flat-
tened, slightly broader than long (HL/HW 0.90);
head length about 1/3 snout to vent length (HI /S-
V 0.35).

Snout not prominent, slightly rounded in pro-
file, canthus rostralis distinct, slightly curved,
Nostrils lateral, their distance to snout slightlv
less than distance between eye and naris (E-N/IN
1.46).

Eye prominent, slightly larger than eye to naris
distance. Tympanum large with narrow annulus,
diameter 0.6 width of eye. Vomerine teeth on two
large elevations between posterior margin of
choanae. Tongue broadly oval.

Fingers moderately long, slender and un-
webbed; finger lengths 3>4>2>1; terminal discs
large.

Hindlimbs moderately long (TL/S-V 0.57); toe
lengths 4>3>5>2>1, Web reaches base of termi-
nal disc on all toes except 4th; reaches top of
subarticular tubercle at base of penultimate pha-
lanx on 4th toe; prominent inner metatarsal Luber-
cle; no outer metatarsal tubercle.

Skin af dorsum smooth. Strong supratympanic
fold extends beyond superior margin of tyrnpa-
num. Ventral surface coarsely granular. In life
dorsum mottled brown with irregular cream

308

markings and an indistinct pale dorsolateral stripe
Inn eye bulge to groin, posterior ventral surface
cream, throat faintly greyish Lo junction ol arms.

Dimensions of the holotype in mm: S-V 99.44;
HL. 35.14; HW 39.14; TL 56.38; E-N 10.46; IN
7.2; E 10.48; T 6.24.

VARIATION. The overall proportions of the 11
paratypes are similar (HL/HW 0.90-1.11; HL/S-
V 0340.40; E-N/IN 1.19-1.79). Hand and toe
webbing has ne detectable variation: In life low
tubercles on the hack, eye bulge and around the
nosirils ure evident in some animals,

The juvenile frog (QMJS9009) has a disrinctl y
dark throar with a pale line on the mid-throat. The
dark throat pales to a light grey in adults, In life
à dorsolateral stripe extends from the eye bulge
Io groin, becoming less distinct in larger frogs.
Juveniles are light cream. with dark markings
laterally. These fade with increasing size.

Males possess dark brown, glandular nuptial
pads and lack å vocal sac.

ETYMOLOGY) The specific name, andiirrmalin, 18
frøm the Barrow Point Aboriginal language,
Gambulmugu, tor the frog (Roger Hart pers, comm.
Aboriginal elder and cultural informant). Mr Hart ex-
plains that in local tradition the frogs are people who
have been transformed as punishment for breaching
certain miles, The species and habital are believed to be
culturally sensitive by Aboriginal people of the area.

COMPARISON WITH OTHER SPECIES. The
combination of large size, no webbing on the
slender fingers, full webbing on the toes, strong
supratympanic fold and colouration is not shared
by any species groups defined by Tyler & Davies
(1978). The general appearance of body shape
when first observed is similar to Litoria caerulea.
However L. andiirrmalin lacks the large parotoid
glands and finger webbing of L. caerulea. Mem-
bers of the Litoria cirropa species group have
moderately long, slender ünwebbed fingers like
L. andiirrmalin but do not possess full toe wèb-
bing ora well-defined supratympanic fold and are
much smaller.

NATURAL HISTORY

Litoria andiirmalin was observed near cas-
cades and riffles hur not pools, in perennial
sireams above 60m in the Melville Range, The
streamside vegetation is mesophyll vine forest
with Melaleuca emergents occurring as gallery
forest. Large granite boulders and bedrock of the
Permian Altonmoui Granites are a feature of the
stream habitat.

MEMOIRS OF THE QUEENSLAND MUSEUM

The frog was ohserved on rocks, vines and
Wigs adjacent 10 broken water, U did not jump
into the stream to escape when disturbed but
preferred to climb under boulders or into vegeta-
tion. When placed in running water or pools the
frog immediately swam to the bank, and jumped
into the adjacent habitat. It did not attempt ro
escape into the siream like stream frogs of the Wet
Tropics Biogeographic Region (ep. Liroria
nannotis, Taudactylus acutirostris, Unpubl, abs.)

A 9 (QMJ59002) had small (Imm) pigmented
eggs in May 1994, Other 22 observed in the
held in October, 1994 and November, 1995 also
had eggs. ? 9 examined in mid to late February
1995 had no eggs suggesting breeding takes place
between early December to mid February cuin-
ciding with the onset of the monsoonal wet sea-
son. ? 9 carry eggs at least from May to at least
the onset of the monsoon season. Similar carrying.
of eggs outside the breeding season has been
documented ( McDonald & Davies 1990).

Males were calling in late November, 1995.
The call sounds like tapping with a stick on à
piece of bamboo, a rapid, gentle ‘toc toc toc toc
toc’. No calling was heard in other months

Notadpoles of any species were observed in the
stream or pools in May, October, November ar
February. An undescribed species of Cophixalus
was located in the boulders adjacent to the stream
habitat of L. andiirrmalin and adult Rana daemeli
and Litoria infrafrenata were observed in the
same stream habitat. In February 1995 L in-
Jrafrenata, L. caerulea and R, daemelti were call-
ing near and adjacent to the stream whilst
juveniles of L. andiirrmalin and R. daemeli were
present in May and October also indicating pos-
sible monsoonal-season breeding.

Examination of contents of two faccal pellets
deposited overnight disclosed a leaf, an unident-
fiable small frog, freshwaler prawn legs
(Macrobrachium sp.) and beetle remains.
Flushed stomach contents from five frogs re-
vealed a litter skink Lygisaurus sp. coleopteran
larva, beetle and cockroach remains.

DISTRIBUTION AND CONSERVATION
STATUS

The species is restricted to streams in the Mel-
ville Range which is part of the greater Laura
Basin, and an area of higher rainfall than the
adjacent areas of Cape York, This is especially so
on the coast which is influenced by the orientation
of the coastal ranges and prevailing south east
winds. The Melville Range in Cape Melville Na-

A NEW STREAM-DWELLING LITØRTA

tional Park is a distinct geological feature of
Altonmoui granites supporting à unique combi-
nation of rainforest and sclerophyll forest com-
munities with affinities closer to the MeTwraith
Range than to the Wet Tropics biogeographic
region (J.P. Stanton, pers. comm.). The greater
Laura Basin, including the eastern escarpments
and isolated ranges largely in Starke Pastoral
Holding and the dune system of Cape Flattery,
includes endemic species of skinks found on
sandstone, granite or silica sands — Lerista in-
grami, Ctenotus quinkan, Ctentous rawlinsoni,
Crentotus nullum, Carlia dogare and
Cryptoblepharus fuhni and a boulder-inhabiting
frog. Cophixalus sp. nov.

The geological formations, vegetation types
and climatic conditions are atypical for Cape
York and Wet Tropics biogeographic regions
(Stanton & Morgan 1977) indicating that the area
should be recognised as a distinct biogeographic
region, asreflected in the unique assemblages and
endemic species of fauna and flora.

The habitat of Litoria andiirrmalin is restricted
Within a conservation land tenure which is not
threatened at this stage. The conservation status
would be 2R (rare and restricted 10 less than
100km) using the criteria of McDonald et. al.
(1991). Other species endemic to the Melville
Range are the skink Cryptoblepharus fuhni
(Roberts 1994), an undescribed frog Cophixalus
sp., the Foxtail Palm ( Wodverta bifurcata) and the
tree Acmenosperma pringlei.

ACKNOWLEDGEMENTS

I am indebted to Peter Stanton who drew my
attention to frogs in the Cape Melville area and
made it possible to re-visit the site of his original
collection. Dave Fell, Juliana Ledger, Clifford
Frith, Rob Worrall and Les Jackson assisted in the
field and in various ways. Priscilla Gibson, Peter
Sutton and Roger Hart gave helpful guidance

with names used by the aboriginal people in the
Cape Melville area for frogs. Roger Hart also
gave permission for the the name and the aborig-
inal story to be told. Jeanette Covacevich, Dr
Marg Davies, Suzan Horder, and Steve Richards
provided helpful advice. Professor Barrie Jamie-
son and Associate Professor Craig Moritz, Centre
for Conservation Biology, University of Queens-
land provided facilities to complete this work
while I was on the Queensland Government In-
terchange Programme. This assistance is most
gratefully appreciated.

LITERATURE CITED

FRITH, D.W. & FRITH, C.B. 1995. Cape York Penin-
sula: a natural history. (Reed Books: Chatswood,
N.S.W.).

MCDONALD, K.R., COVACEVICH, LA, INGRAM,
GJ. & COUPER, P.J. 1991. The status ol Irogs
and reptiles, Pp 338-345. In Ingram, GJ, & Raven,
RI (eds) ‘An Atlas of Queensland's Frogs, Rep-
tiles, Birds and Mammals’. (Queensland Mu-
seum; Brisbane).

MCDONALD, KR. å DAVIES, M, 1990, Morphol-
ogy and biology of the Australian tree frog Litoria
pearsoniana (Copland) (Anura: Hylidae). Trans-
actions of the Royal Society of South Australia,
11403 & 4); 145-156.

ROBERTS, L, 1994, New data on Cryproblepharus
Juhni, a poorly known skink from Queensland.
Memoirs of the Queensland Museum 35(1): 234.

STANTON, LP & MORGAN, G. 1977. The rapid
selection and appraisal of key and endangered
sites, The Queensland case study. University of
New England, School of Nalural Resources Re-
port, No. PRA.

TYLER, M.J. 1968. Papuan hyhd frogs of the genus
Hyla. Zoologische Verhandelingen, No. 96: 1-
203.

TYLER, M.J. & DAVIES, M. 1978. Species-groups
within the Australopapuan Hylid [rog genus Lito-
ria Tschudi, Australian Journal of Zoology Sup-
plementary Series, No. 63: 1-79

310

A RANGE EXTENSION FOR LITORIA BREVI-
PALMATA (ANURA: HYLIDAE), Memoirs of the Queens-
land Museum 42(1): 310. 1997:- Until 1995 the distribution
of Litoria brevipalmata was believed to have extended from
the SEQ Border Ranges and Darling Downs (McDonald, 1974
collécted specimens from Crows Nest and Ravensbourne
National Parks) to Brisbane's metropolitan area (Nattrass and
Ingram, 1993, recorded species at Woogaroo Ck near Wacol,
Brisbane), through the extreme coastal lowlands, Glass House
Mountains (Hannah & Smith, 1995; Gynther pers. comm.)
and Conondale Ranges/Jimna area, Kilcoy Shire (McEvoy et
al 1979 and Czechura 1978). In 1996 a new locality for L.
brevipalmata was recorded in the. Brooweena area (near
Maryborough, SEQ) constituting å northerly range extension
(Nattrass pers. comm.). This paper documents a new locality
at Yarraman State Forest for L. brevipalmata, (he most west-
ern known locality for this species (approximately 40 km west
of Brooweena). Two adults and one juvenile were recorded at
Yarraman State Forest 289 (Qld Forestry Map number 9244-
2), southeast Queensland. The juvenile was positively identi-
fied by G. Ingram (Qld Museum, pers.comm.).

Two adults were observed during a survey conducted on
18 November, 1995, along Rocky Creek (SF 289), after
51.8mm of rain had fallen two days previously, Rocky creek
is buffered trom surrounding Araucaria cunninghamii plan-
tations by dry scleraphyll forest dominated by Eucalyprus and
Angophora spp. with a ground cover predominantly of exotic
herbs, grasses and Lantana camara. The first adult was
observed at a dam (grid reference Forestry 9244-2; 936 314),
approximately 200m from Rocky Creek, located about 2 m
from the water's edge. The second was observed on an ex-
posed branch of a fallen Casuarina overhanging the water
upstream of Rock Creek, beside another dam (grid reference
Forestry 9244-2; 941 317). Numerous trees, mainly
Casuarina and Eucalyptus spp., were found on the dam wall
with a ground cover similar to that at the first dam.

Other species of frogs sighted or heard vocalising here
included: Crinia signifera, Limnodynastes terraereginae,

MEMOIRS OF THE QUEENSLAND MUSEUM

Litoria caerulea, L, chloris, L, dentata, L, fallax, L. nasuta, L.
latopalmata, L, lesueuri, Mixophyes fasciolatus and Bufo
marinus. At no time was L. brevipalmata heard.

A subsequent survey was conducted on 28 November,
1995, after 59.8mm of rain had fallen between 20 and 23
November. A juvenile was collected 30 cm from the water's
edge at the first dam (grid reference Forestry 9244-2; 936
314). It is interesting that no frogs were observed during
surveys on I October, 1995, or 9 November, 1995, at the dams
along Rocky Creek when no rain had fallen previously. lt
appears that L. brevipalmata only becomes active after sulfi-
cient rainfall, concurring with observations made R. Nattrass
(pers. comm). A final survey conducted on the night of 29
November failed to locate L, brevipalmata, although all other
frog species present during the previous fortnight's surveys
were still found, in low abundance.

Literature cited

CZECHURA, G.C, 1978. A new locality for Litoria
brevipalmata (Anura: Pelodryadidae) from South-
Eastern Queensland. Victorian Naturalist 95: 150-151.

HANNAH Dn & SMITH, G.C. 1995. The effects of pre-
scribed burning on herptiles in southeastern Queens-
land. Memoirs of the Queensland Museum 38(2):
529-511.

McDONALD, K.R. 1974. Litoria brevipalmata, An addition
to the Queensland amphibian list. Herpetofauna 7( 1):
2-4

McEVOY,, J.S., MCDONALD, K.R. & SEARLE, A.K, 1979.
Mammals, birds, reptiles and amphibians of the Kilcoy
Shire, Queensland Journal of Agriculture and Animal
Sciences 36; 167-180.

NATTRASS, A.E.O. & INGRAM, G.J. 1993, New records
of the rare Green-thighed frog. Memoirs of the Queens-
land Museum 33: 348,

Jason J. Aridis, Queensland Forest Research Institute, DPI

Forest Service, 80 Meiers Rd, Indooroopilly, Queensland

4068, Australia; 17 June 1996.

A NEW SEMI-TERRESTRIAL ACOTYLEAN FLATWORM, MYORAMYXA PARDALOTA

GEN. ET SP. NOV. (PLEHNIIDAE POLYCLADIDA) FROM SOUTHEAST
QUEENSLAND, AUSTRALIA

LESLIE J. NEWMAN AND LESTER R.G. CANNON

Newman, L.J. & Cannon, L.R.G. 1997 06 30. A new semi-terrestrial acotylean flatworm,
Myoramyxa pardalota gen. et sp. nov. (Plehniidae, Polycladida) from southeast Queensland,
Australia. Memoirs of the Queensland Museum 42(1): 311-314. Brisbane. ISSN 0079-8835,

Anew acotylean flatworm, Myoramyxapardaloia gen. et sp. nov., is described from the high
intertidal zone, southeast Queensland, Australia. The family Plehniidae Bock, 1913 is
emended to contain this new monotypic genus which is characterised by possessing eyes
around the anterior margin and retractile nachal tentacles with eyes, but no frontal eyes, no
seminal vesicle, and ejaculatory ducts that enter at the side of the large prostate; Lang’s
vesicle is lacking. The False Water-ràt, Xeromys myoides, was observed to prey on these
worms: this is the first record of a mammal feeding on a polyclad flatworm. [ ] Polycladida,
Acotylea, Plehniidae, flatworm, taxonomy.

LJ Newman, Zoology Dept, University of Queensland, Queensland. 4072; Lester R.G.
Cannon, Queensland Museum, P.O, Bax 3300, South Brisbane, Queensland. 4101; 26

March 1997.

Polyclad flatworms are, with one freshwater
exception (Limnostylochus from Borneo), marine
animals (Prudhoe, 1985) where they are well
known as predators of invertebrates such as small
crustaceans, molluscs and ascidians (Prudhoe,
1985; Cannon, 1986; Newman & Cannon, 1994).
Some are even well-known pests of commercial
bivalves including rock oysters (Steade, 1907;
Jennings & Newman, 1996), pearl oysters and
giant clams (Newman et al., 1993) and may pose
a threat to these industries. None are known to
invade terrestrial habitats.

Though active predators themselves, only three
studies report predators of polyclads; these in-
clude a polychaete (Riser, 1974), and putferfish
(Poulter, 1975; Jennings & Newman, 1996), That
polyclads are unlikely to be attacked or eaten is
attributed to their being distasteful according to
Prudhoe (1985), presumably because of their tox-
icity. Studies have shown that some acotylean
Species contain highly toxic substances such as
tetrodotoxin (Jeon et al., 1986; Miyazawa era,
1987). This may also explain why pufferfish can
consume these worms (Flowers, pers. comm.).

This account is of a new monotypic genus of
polyclads found in the semi-terrestrial supralitto-
fal zone of Moreton Bay where it is exposed to
predation by å mammal, the False Water-rat,
Xeromys myoides,

METHODS

Worms were hand collected from under fallen
logs, driftwood and large rocks from the high

intertidal zone from: Myora, North Stradbroke
Island (27°27’40"S; 153°25°40"E), Caloundra
(26^48'S5; 153"08'E) and Pebble Beach, near
Bribie Island (27705'S; 153?08' E), southeast
Queensland. Animals were photographed in the
laboratory, fixed on frozen polyclad fixative and
preserved 1n 70% ethanol for histological prepa
rations (Newman & Cannon, 1995). Whole
mounts were stained with Mayer's Hacmalum,
dehydrated in graded alcohols and then mounted
in Canada balsam. Longitudinal serial sections of
the reproductive region were obtained from spec-
imens embedded in Paraplast (56°C), sectioned
at 5-7um, and then stained with haematoxylin
and eosin,

Measurements of the body were taken from live
animals in a relaxed state and are given as length
mm x width mm. Measurements of the reproduc-
tive organs are taken from the paralypes. Recon-
struction of the reproductive system is
diagrammatic and derived from the sections with
minimal interpretation. Drawings were made
with the aid of a camera lucida by L.J.N. Speci-
mens were collected and photographed by Bruce
Cowell. This material is lodged at the Queensland
Museum (QM) as wholemounts (WM), serial
sections (LS) and wet specimens (S).

PLEHNIIDAE Back, 1913
Myoramyxa gen. nov.

DIAGNOSIS. Plehniidae with marginal eyes in 3
to 4 rows anteriorly, becoming less numerous

about 1/3 of the way posteriorly along the margin.
Frontal eyes absent. Posessing nuchal tentacles.

ETYMOLOGY. Named Myera for its type location
and myxa = shme (L. fem.) for its ability lo produce
copious amounts of mucus.

TAXONOMIC REMARKS. As presently diag-
nosed (dreproductive system without pro-
statoids, ejaculatory duct enters the side of the
prostate, no seminal vesicle: ? system with duc-
tus vaginalis with large ruffled antrum)
Plehniidae contains worms Jacking nuchal tenta-
cles (Prudhoe, 1985; Cannon, 1986). The rather
unusual and distinctive insertion of the ejacula-
tory duct into the side of the prostate distinguishes
the Plehniidae from all other polyclads.
Myoramyxa clearly has this character. We believe
it is appropriate 10 emend the family diagnosis to
include worms with or without nuchal tentacles.
It should be noted that nuchal tentacles may not
always be seen in poorly fixed material and may
have been overlooked previously.

Within the Plehniidae this genus is similar to
Discacelides Bergendal, 1893, however,
Myaramyxa lacks a Lang's vesicle.

Myoramyxa pardalota sp. nov.
(Figs |, 2A-C)

MATERIAL EXAMINED. HOLOTYPE, Myora.
Stradbroke ls., southeast Qld, under logs and rocks,
high intertidal, 04.05.92, S. Van Dyke WM,
QMG210875. PARATYPES. Same data, S.
QM210878; 12.05.92, LS, QMG210874; 19.07.92,
LS, QMG210876; S, QMG210808; WM, QMG-
210877; Pumistone Passage, Caloundra, SE Qld, under
log. high intertidal. 07.10.93, P. Davie, LS,
QMG210873; Pebble Beach, Bribie Is., SE Qld,
05.1 | 92, L, Newman & Z. Khalil, LS, QM G210872.

DESCRIPTION. Dorsal surface light brown with
dark brown leopard spots (composed of micro-
dors) closely packed in å dense pattern, darker
brown medially forming a narrow, longitudinal
line (Fig. 1). Ventral surface light orange-brown,
darker medially with some dark brown pigment
surrounding the mouth and gonopores.

Body round oval, thick and fleshy. Nuchal ten-
tacles short, conical, retractile and relatively
small, with about 40-50 eyes inside each tentacle,
eyes absent aL the base (Fig. 2A.B). Marginal eyes
in 3 (o4 rows anteriorly, becoming less numerous
(to 2 rows) about 1/3 of the way along the margin,
About 150-200 cerebral eyes between the tenta-
cles, no frontal eyes. Pharynx oval, central, mouth
posterior, extending approx. 1/3 the length of the
body. Gonopores posterior to the pharynx. d pore

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Live Myoramyxa pardalora, gen. ei sp. nov,
holotype (QMG210875): dorsal (top) and posterior
ventral views. (Photo by B. Cowell)

small, anterior and close to the ? pore; ductus
vaginalis well separated from gonopores, large
ruffled antrum. Size range 30mm x 25mnt (ma-
iure) to 35mm x 23mm (mature).

Large prostate (1.8 mm), seminal vesical lack-
ing, spermaducal bulbs joining ejaculatory duct
which enters laterally into the prostate, Penis
papillae wide, without stylet. å antrum narrow
and deep. 9 antrum deep and ruffled. Vagina
leads dorsoanteriorly. then is highly coiled poste-
riorly until it leads into à voluminous, muscular,
swollen ductus vaginalis.

DIAGNOSIS. With characters of the genus and a
dorsal surface with dark pattern of brown spots
like a leopard with a narrow, dark median line.

ETYMOLOGY. Named pardalara = spotted like a
leopard (L. fem.) for its spotted dorsal colour pattern.

HABITAT & DISTRIBUTION. Common inside or
under hardwood logs, driftwood or rocks in the high
intertidal (supralittoral) zone particularly with adjacent
inangroves from Moreton Bay. SE Queensland.

DISCUSSION. The Plehniidae are somewhat
aberrent members of the Stylochotdea (sensu
Prudhoe, 1985) in that they frequently have few
or no eyes anteriorly, The distinctive manner in
which the ejaculatory duct joins at the side of the

A NEW SEMI-TERRESTRIAL FLATWORM 313

FIG. 2. Myoramyxa pardalota, gen. et sp. nov.; A, holotype (QMG210875), wholemount of the ventral side and
B, eye arrangement from the dorsal side; C, paratype (QMG210876), diagrammatic representation of the
reproductive anatomy. Scales: A,B, 5mm; C, Imm. (Legend: ce = cerebral eyes, cg = cement glands, dv = ductus
vaginalis, ed = ejaculatory duct, fp = 9 pore, i - intestine, m = mouth, me = marginal eyes, mp = d pore, nt =
nuchal tentacles, p = prostate, ph = pharynx, pp = penis papilla, sb 2 spermaductal bulb, v = vagina).

prostate, however, would seem to be a syn-
apomorphy for the group. Characters such as the
presence of nuchal tentacles, which could so eas-
ily be overlooked, we believe should not be over-

emphasised. Consequently, we consider the
emendation of the family Plehniidae to be justi-
fied and a more conservative approach than erec-
tion of a new family at this time.

314

These worms were brought to our attention by
Steve Van Dyck who had found them being eaten,
with apparent relish, by the False Water-rat,
Xeromys myoides (see figure p. 36-7) (Van Dyck,
1994). The rat forages at night in the supralittoral
and littoral at the upper influence of the tide.
During the day worms are found in sheltered
locations quite out of water such as the moist
inside or underside of hardwood logs thrown up
as driftwood in the suprlittoral zone; as many as
14 worms have been found clustering together.

In the laboratory worms actively retreat from
water, if placed in it and given the opportunity to
crawl out. They produce copious amounts of clear
mucus which surrounds the resting animal like a
jelly coat. In the field their presence is revealed
by slime trails similar to those produced by slugs
and snails. Presumably these animals emerge at
night to forage on small invertebrates. They are
clearly semi-terrestrial which has evidently ex-
posed them to predation.

Many polyclads are known to be highly toxic:
it remains to be seen if Myoramyxa is toxic par-
ticularly since the rat evidently finds them
delectible.

ACKNOWLEDGMENTS

We wish to thank S. Van Dyke for finding this
unusual worm and P. Davie and Z. Khalil for
collecting additional specimens. Specimens were
prepared for histology by Z. Khalil and curated
by K. Sewell. B. Cowell kindly photographed live
animals. Financial support was generously pro-
vided to L.J.N. by the Australian Biological
Resource Study, Canberra. We would also like to
thank the Queensland Museum and the Zoology
Department, University of Queensland.

MEMOIRS OF THE QUEENSLAND MUSEUM

LITERATURE CITED

CANNON, L.R.G. 1986. Turbellaria of the World- A
Guide to Families and Genera. Queensland Mu-
seum, Brisbane. 136 pp.

JENNINGS, K.A. & NEWMAN, L.J. 1996 Four new
stylochid flatworms associated with oysters from
southeast Queensland, Australia. Raffles Bulletin
of Zoology. 44:493-508

JEON, K. J., MIYAZAWA, K., NOGUCHI, T., NAR-
ITA, H., MATSUBARA, S., NARA, M., ITO K.,
& HASHIMOTO, K. 1986. Occurence of Para-
lytic Toxicity in Marine Flatworms. Bulletin of the
Japanese Society of Scientific Fisheries 52(6):
1065-1069.

MIYAZAWA, K., JEON, J. K., NOGUCHI, T., ITO, K.
& HASHIMOTO, K. 1987. Distribution of
Tetrodotoxin in the Tissues of the Flatworm
Planocera multitentaculata (Platyhelminthes).
Toxicon 25(9): 975-980.

NEWMAN, L.J. & CANNON, L.R.G. 1994.
Pseudoceros and Pseudobiceros (Polycladida,
Pseudocerotidae) from Eastern Australia and
Papua New Guinea. Memoirs of the Queensland
Museum 37(1): 205-266.

1995. The importance of the fixation of colour,
pattem and form in tropical Pseudocerotidae
(Platyhelminthes, Polycladida). Hydrobiologia
305: 141-143.

NEWMAN, L.J., CANNON, L.R.G. & GOVAN, H.
1993. Stylochus (Imogene) matatasi n. sp.
(Platyhelminthes, Polycladida): Pest of cultured
giant clams and pearl oysters from Solomon Is-
lands. Hydrobiologia 257: 185-189

POULTER, J.L. 1975. Hawaiian polyclad flatworms:
prosthiostomids. Pacific Science 29: 317-339.

PRUDHOE, S. 1985. A Monograph on Polyclad
Turbellaria. (Oxford University Press: Oxford).

RISER, N.W. 1974 Epilogue. Pp. 517-524. In Riser,
N.W. & Morse, M.P. (eds) ‘Biology of the
Turbellaria’ (McGraw-Hill: New York).

STEAD, D.G. 1907. Preliminary note on the wafer
(Leptoplana australis), a species of dendrocoe-
lous turbellarian worm, destructive to oysters. 6 p.
(Department of Fisheries, NSW: Sydney).

VAN DYCK, S. 1994 The rats at Neptune’s table.
Australian Natural History 24(8): 30-37.

OSTEOLOGY OF IMMATURE DARK SHOULDER MINKE WHALES BALAENOPTERA
ACUTOROSTRATA FROM SOUTHERN QUEENSLAND

R.A. PATERSON, HA JANETZKI AND S.C. WILLIAMS

Paterson, R.A., Janetzki, H.A, & Williams, S.C. 1997 06 30; Osteology of immature dark
shoulder minke whales Balaenoptera acutorostrata from southern Queensland, Memoirs. of
the Queensland Museum 42( 1); 315-325. Brisbane. ISSN 0079-8835,

Osteology of a physically immature 6.02m long dark shoulder minke whale Balaenoprerd
aculorostrata is described as well as its baleen, Comparison is made with the skulls of three
other less mature dark shoulder forms in the Queensland Museum collection and problems
associated with reliance on osteological features of non-adull specimens are discussed. [ ]
Minke whale, Balaenoptera avutorastrata, osteology.

F.A. Paterson, H. A. Janeizki and S.C. Williams, Queensland Museum, PO Box 3300 Soul
Brisbane, Queensland 4101, Australia; received 6 February 1997

On 27 August 1995 å minke whale cårcass was
found drifting in the Great Sandy Strait near
Boonooroo (25?40'S, 152?54' E). It was retrieved
the following day and flensed by å Queensland
Museum team on 29 August. By then, this 6.02m
long whale had decomposed and its skin had
extensively sloughed. Identification as a dark
shoulder or Antarctic form, described as Type 1
ar 2 by Best (1985), was made on baleen appear-
ances and not from external features. The entire
skeleton, with the exception of the pelvic bones.
was recovered together with the baleen and lar-
ynx. The specimen is registered JM 10961 in the
mammal collection of the Queensland Museum.

BALEEN DESCRIPTION

The baleen of JM10961 is illustrated from the
buccal aspect in Fig. 1. There were 282 plates on
the right and 280 on rhe left, à count within the
norma] range for the species worldwide (Hor-
wood, 1990) and the southern hemisphere in par-
ticular (Best, 1955). All-white anterior plates
numbered 119 (42%) and 74 (26%) on the right
and left respectively. The largest plates measured
27cm in length and 10cm in width. The propor-
jional width of the dark outer border of those
plates was 35%. These features, viz. asymmetri-
cal [R>L] anterior white colouration, proportion
of all-white plates, length of largest plates and
width of the dark outer border, conform with the
description by Best (1985) of the dark shoulder
(Antarctic) or Type 1 and 2 forms. In contrast, the
largest plates of the diminutive (Type 3) form
called the dwarf form by Arnold et al. (1987) do
not exceed 20cm in length and the plates are
either all-white or a large proportion are white.

SKULL DESCRIPTION

Measurements, after Omura (1975) and Arnold
et al. (1987). of the skull, mandible, basihyoid and
stylohyals of JM 10961 are contained in Table |
and the structures are illustrated in Figs 2-5. The
skulls of three other dark shoulder minke whales
considered to be more immature than JM 10961,
in the Queensland Museum collection, are shown
from their dorsal aspects in Fig. 6. Details of those
specimens are contained in Table 2. (There are
damaged skulls of two other immature dark
shoulder forms in the collection as well as intact
skulls of two immature dwarf forms.)

Allowing for the immaturity of JM10961, ils
skull morphology is not appreciably different
from the two adult dark shoulder (Antarctic)
specimens described by Omura (1975), Exelu-
sion of the parictals from the vertex is evident in
JM10961 and the other Queensland dark shoulder
forms. The hamular processes of the pterygoids
are blunt and rounded (Fig. 7). Omura (1975)
considered these two cranial features to be im-
portant osteological differences between Antano-
uc and North Pacific minke whales. In the latter
the parietals are included in the vertex, å leature
mis-labelled by Horwood (1990: 11), and the
hamular processes åre elongate, å feature also
seen (Fig. 8) in a North Atlantic specimen
(D'Alton, 1827).

However, there are some differences between
the Queensland dark shoulder specimens and
adult Antarctic specimens. (Omura, 1975). The
anterior concavity of the nasals is less pro-
nounced and this may reflect immaturity but the
retro-position of the ascending processes of the
maxillaries relative to the nasals and premaxillar-
ies (Figs 2 & 6), a feature of North Pacific spec-
imens (Omura, 1975), may not be explicable

316

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. I. Baleen from buccal aspect of JM10961. The right baleen row is on the right of the figure.

solely on the basis of immaturity. Whilst further
growth of the Queensland dark shoulder whales
and those from the North Pacific (which were also
1mmature) may have resulted in maxillary re-po-
sitioning, this feature is subject to revision and
will require inspection of a larger series of adult
specimens from the northern and southern hemi-
spheres. Arnold et al. (1987), however, when
describing the features of an adult Type 3 skull
from northern Queensland, considered that more
immature Type 3 specimens exhibited similar
retro-position of the maxillaries. That feature is
also evident in the illustration of the skull of an
immature 4.1m long Type 3 specimen (JM8808)

in ihe Queensland Museum collection (Paterson,
1994),

The supraoccipitals in the Queensland dark
shoulder specimens (Figs 2 & 6) differ from
Antarctic and North Pacific specimens (Omura,
1975), a North Atlantic specimen illustrated by
D'Alton (1827) and Type 3 specimens (Arnold et
al., 1987). They are narrow anteriorly, particu-
larly in JM8513, and in JM5434 there is associ-
ated depression which may reflect individual
variation. The narrowing may be a manifestation
of the developing cranium as it 18 less pronounced
in IM10961 which is the most mature of the
Queensland dark shoulder specimens.

OSTEOLOGY OF IMMATURE MINKE WHALES 317

FIG. 2. Skull from dorsal aspect (left) and ventral aspect (right) of JM10961. (Scale in cm)

FIG. 3. Skull from lateral aspect of JM10961. (Scale in cm)

318

FIG. 4. Mandible from dorsal aspect of JM10961.
(Scale in cm)

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Basihyoid (below) and stylohyals (above) of
JM10961. (Scale in cm)

POST CRANIAL DESCRIPTION

VERTEBRAL COLUMN INCLUDING CHEV-
RONS. The vertebral formula (C7, D11, L12,
Ca18=48) conforms with the range of 47-50 for
the species worldwide and 48-50 from the Ant-
arctic in particular (Ohsumi et al., 1970; Omura,
1957 & 1975; Tomilin, 1967; True, 1904). The
caudal region was carefully preserved and dis-
sected and it can be confidently stated that no
small caudals were lost. The central epiphyses
were unfused with the exception of Cl and Ca 18.

FIG. 6. Skulls from dorsal aspect of J21708 (left); JM5434 (centre); JM8513 (right). (Scales in cm)

OSTEOLOGY OF IMMATURE MINKE WHALES

FIG. 7. Close-up view of hamular processes of
JM10961. (Scale in cm)

The C7 parapophysis is lacking, a feature consis-
tent with other dark shoulder forms (Omura,
1975). The vertebrae are illustrated from their
lateral aspects in Fig. 9. Their measurements are
contained in Table 3 including mean vertebral
length. This value is derived from the formula
(aXbXc) ^ where a, bande represent the breadth,
height and length respectively of the centra
(Omura, 1971). Comparison between the mean
vertebral lengths of two immature North Pacific
specimens (5.4 and 6.6m in length), two adult
Antarctic specimens (8.5 and 9.8m) and
JM10961 (6.02m) is shown in Fig. 10. The values
are consistent with the varied maturity of the five
specimens.

There were twelve chevrons and they are
illustrated in Fig. 11 and their measurements are
contained in Table 3, The laminae of the first two
and second last were unfused and the last repre-
sented by a solitary lamina.

RIBS AND STERNUM. There were eleven pairs
of ribs and they are illustrated together with the
sternum in Fig. 12. Their measurements are con-
tained in Table 4.

SCAPULAE AND FLIPPER BONES. The scap-
ulae, humeri, radii and ulnae are illustrated in Fig.
13. The phalangeal formula (including the meta-
carpals) is I3-4, Hs, Is-7, IV4, compared with Iss,
To-x, II7-8, IV4 from two adult Antarctic speci-
mens (Omura, 1975), Measurements are con-
tained in Table 5.

319

FIG. 8. Skull from ventral aspect of a North Atlantic
minke whale (from D' Alton, 1827).

Tomilin (1967) noted that, in northern hemi-
sphere minke whales, the proportional breadth
and height of the scapulae increase in larger
(older) whales and breadth increases more than
height. That observation 1s confirmed in Table 6
where the relevant scapular dimensions of
JM10961 are compared with two adult Antarctic
specimens (Omura, 1975).

320

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. Vertebral column from lateral aspect of JM10961. Top, cervical and dorsal vertebrae; centre, lumbar

vertebrae; bottom, caudal vertebrae. (Scale in cm)

DISCUSSION

The evidence presented here with regard to the
dark shoulder form of minke whale, which is
tentatively regarded as Balaenoptera
acutorostrata bonaerensis (Burmeister, 1867) by
Horwood (1990), highlights the problem that the
number of specimens available for comparison is
limited and they are often immature with resul-
tant difficulties in interpretation of osteological
characters. This problem has been considered in
detail by Arnold et al. (1987) in relation to the

dwarf (Type 3) form. A future osteological study,
in which all adult specimens (particularly those
of known Type) in Australasian collections were
assessed, may further clarify the taxonomy of
Balaenoptera acutorostrata.

ACKNOWLEDGMENTS

Vic Hislop retrieved the whale carcass from the
Great Sandy Strait and shore transport was co-or-
dinated by Geoff Brittingham of the Department
of Environment. Paul Avern, Patricia Paterson,

OSTEOLOGY OF IMMATURE MINKE WHALES 321

250

Antarctic | Å

Mean Vertebral Length (mm)

Antarctic

Ve | EN
— ——N Pacific BN

50
| — —N Pacific

+ + am x + + +
T e un nm N ka o o e N W Å kel a e N p. pr Kl « o N wo Ka g
OP a CIO NV 5 328 $ 3 BS SS SS 3
o oO o o o o

Vertebrae

FIG. 10. Comparison between mean vertebral lengths of JM10961, North Pacific and Antarctic minke whales
(from Omura, 1975).

FIG. 11. Chevrons of JM10961. (Proximal at upper left and distal at lower right.) (Scale in cm)

322 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 13. Scapulae (left); humeri, radii and ulnae (right) of JM10961. (Scale in cm)

Steve Van Dyck and Owen Walker assisted with dwarf form. Scientific Reports of the Whales Re-
flensing. Bruce Cowell took the photographs. search Institute, Tokyo 38: 1-46.

BEST, P.B. 1985. External characters of southern minke

LITERATURE CITED whales and the existence of a diminutive form.

de h Insti-
ARNOLD, P, MARSH, H. & HEINSOHN, G. 1987. Scientific Reports of the Whales Research Insti

The occurrence of two forms of minke whales in tute, Tokyo 36: 1-33. — sch
east Australian waters with a description of exter- BURMEISTER, H. 1867. Preliminary description of a

nal characters and skeleton of the diminutive or new species of finner whale (Balaenoptera

OSTEOLOGY OF IMMATURE MINKE WHALES

323

TABLE 1. Skull, mandibular and hyoid measurements (in mm) of JM10961.

Breadth of skull, squamosal

Breadth of skull, frontal 670

Breadth of skull, maxillaries

Length of orbit, frontal, right
Length of orbit, frontal, left
Breadth of occipital bone

156
622

Breadth across occipital condyles (to base of

spongy bone) 166

Height of occipital condyle right

Height of occipital condyle left

Breadth of foramen magnum aperture
Height of foramen magnum aperture

Length from foramen magnum to vertex
(measured at posterior parietals)
Tip of premaxillary to anterior vomer, median |

423

144
950

| Tip of premaxillary to anterior palatine, median |

bonaérensis). Proceedings Zoological Society of
London. 707-713.

D'ALTON, E. 1827. 'Die Skelete der Cetacean,
abgebildet und beschrieben.' (Eduard Weber:
Bonn).

HORWOOD, J. 1990. ‘Biology and exploitation of the
minke whale.' (CRC Press: Boca Raton).

OHSUMI, S., MASAKI, Y. & KAWAMURA, A. 1970.
Stock of the Antarctic minke whale. Scientific
Reports of the Whales Research Institute, Tokyo.
22: 75-125.

OMURA, H. 1957. Osteological study ofthe little piked
whale from the coast of Japan. Scientific Reports
of the Whales Research Institute, Tokyo 12: 1-21.

1971. A comparison of the size of vertebrae among
some species of the baleen whales with special
reference to whale movements. Scientific Re-

Measurement Measurement
Condylo-premaxillary length 1552 Tip of premaxillary to posterior palatine, median
Length of premaxillary, right 1053 Tip of premaxillary to posterior pterygoid
Length of premaxillary, left 1047 Breadth across hamular process of ptergyoid
Length of maxillary, superior right 1066 Length of mandible, straight, right
Length of maxillary, superior left 1058 Length of mandible, straight, left
Tip of premaxillary to vertex 1013 Length of mandible, right, outside curve
Tip of premaxillary to nasals 943 inside curve
Length of nasals, median 152 Length of mandible, left, outside curve
Breadth of nasals, anterior (between 93 inside curve

remaxillaries at anterior end of nasals) ` ; nem
oer Se Height of mandible at coronoid, right
ngriorrosmm Height of mandible at coronoid, left
Breadth of rostrum at middle 307 ` i g
Height of mandible at condyle, right
Breadth of rostrum at base 471 e r
Height of mandible at condyle, left
Breadth across maxillaries at vertex 162 r n
—À à SS Tympanic bulla, length, right
readth of frontal across naga s Tympanic bulla, length, left
Breadth between maxillaries at nares 158 a

Tympanic bulla, greatest breadth, left
Tympanic bulla, thickness at middle, right

Tympanic bulla, thickness at middle, left
Malar length, right

Malar length, left

Malar breadth, right
Malar breadth, left

Lachrymal length, right 155
Lachrymal length, left
Lachrymal breadth, right 68

Lachrymal breadth, left

Length (mm)
Basihyoid 80
Stylohyal, right | 42 120
Stylohyal, left | o 121

ports of the Whales Research Institute, Tokyo 23:

1975. Osteological study of the minke whale from
the Antarctic. Scientific Reports of the Whales
Research Institute, Tokyo 27: 1-36.

PATERSON, R.A. 1994. An annotated list of recent
additions to the cetacean collection in the Queens-
land Museum. Memoirs of the Queensland Mu-
seum 35(1): 217-223.

TOMILIN, A.G. 1967. ‘Cetacea. Mammals of the
U.S.S.R. and adjacent countries”. (Israel Program
for Scientific Translations: Jerusalem).

TRUE, F.W. 1904. The whalebone whales of the west-
ern north Atlantic compared with those occurring
in European waters with some observations on the
species of the north Pacific. Smithsonian Contri-
butions to Knowledge. XXXIII, 332 pp.

324

TABLE 2. Other dark shoulder minke whales in the Queensland Museum.

MEMOIRS OF THE QUEENSLAND MUSEUM

| Registration No Sex | Locality Collection Date Full Length | Skull Length
J21708 9 | Currimundi, Caloundra, 26°46’S 153°08’E 25 August 1971 - 865mm
IM5434 | 9 | Granite Bay, Noosa, 26°23’S 153°06’E st1986 | 3.84m | 930mm
| 18513 | 9 | Woralie Creek, Fraser Island, 25*08'S 153°06'E | —16November1990 | Ain | 990mm
TABLE 3. Vertebral and chevron measurements (in mm) of JM10961.
Q e
WEEK P a |73 VEI KJE NA VEV
© o Ka E ks] B So e 9 5 po Kg 5 So
Zia] @/s |z 14 |3 2 | | aja EZ 14 | ex
Tg] EE OB |e bes B EI gl e VE LE [ee
ef R glg :.:.39||l 8|3]|83l2.l 8. B. 8?
> 5 56 |d | de/og|s8 > 5 KSE AECH
CI 272 218 193 125 19 TI Cal 363 343 159 128 142 142
2 332 188 167 90 40 84 2 305 344 158 128 144 143
3 296 159 128 90 24 65 3 291 305 153 128 146 142
4 291 159 122 92 22 63 4 248 278 151 125 142 | 139
5 293 186 119 95 23 64 5 208 246 149 123 138 136
6 119 97 24 65 6 163 222 143 122 135 133
7 121 99 27 69 7 144 200 143 120 128 130
D1 210 123 98 32 73 8 125 175 124 121 127 124
2 222 115 97 42 78 9 112 148 112 123 100 111
3 331 243 121 94 56 86 | | 10 102 112 101 90 69 86
4 375 263 119 94 65 90 11 92 88 89 86 57 76
5 424 279 120 94 74 94 12 83 75 80 74 52 68
6 451 286 118 93 78 95 13 73 66 63 60 49 57
7 459 296 119 94 88 99 14 62 57 53 51 45 50
8 476 [ 302 129 94 91 103 15 54 45 46 4l 38 42
9 483 301 129 94 F 97 106 l 16 43 32 34 29 31 | 31
10 497 95 98 108 17 30 23 23 18 25 | 22
11 478 99 97 110 18
L1 473 102 101 114 Chevron
2 481 103 104 116 1 22, 28 43, 44
3 479 106 107 118 3 94, 99
4 480 108 109 ‘| 120 3 151
5 478 110 113 122 4 151
6 467 112 114 124 5 134
7 456 114 117 126 6 116
8 451 114 119 127 7 100
9 429 117 124 131 8 77
10 415 119 128 133 9 60
11 404 124 134 137 10 44 35
12 380 127 139 141 11 22.21 17, 16
12 12, - 9,-

OSTEOLOGY OF IMMATURE MINKE WHALES 325

TABLE 4. Rib and sternal measurements (in mm) of JM 10961.

] Straight Length A i

Rib

637
482
Width 195

œ [ca |b j [r2 pee
KI
oc
Ka

Breadth 181

Scapula Humerus, Radius, Ulna
Breadth

Greatest breadth

Ratio of breadth to height

Length of acromion-inferior
Breadth of acromion, distal end
Length of coracoid, inferior

Breadth of coracoid, distal end
Length of glenoid fossa

Breadth of glenoid fossa

Length of Se

Phalanx Right Left
I II DI

EY

* damaged ** possibly missing

TABLE 6. Comparison between scapulae of adult and immature dark shoulder minke whales (measurements in mm).

JM10961

287

7152793 (Omura, 1975)
Breadth Height
812 | aos | 449 | 452
2115

71J2883 (Omura, 1975)
Breadth Height
874

Scapula

Condylo-
Dimensions as % of condylo-
remaxillary length of skull

premaxillary length 1552

30.0 | 30.0 | 18.5 18.7

326

BRYDE'S WHALE BALAENOPTERA EDENI ANDER-
SON, 1878 STRANDED NEAR ROSEDALE, QUEENS-
LAND IN 1931. Memoirs of the Queensland Museum 42(1):
326. 1997;- Bryde's whale B. edeni was the last balaenopterid
to be described and its osteological distinction from the sei
whale B. borealis Lesson, 1828 was not resolved until the
studies of Junge (1950) and Omura (1959).

The Queensland Museum Archives contain correspon-
dence between Heber A. Longman, a former Director of the
Museum, and Rosedale residents concerning the stranding of
a large balaenopterid ~ 6km south of Agnes Water (24? 10'S,
151°53’E) in 1931. Longman wrote to Constable E.M. Riley
of Rosedale Police Station on 26 November 1931: ‘J am
greatly indebted to you and Mrs Riley for the information
telephoned last week regarding the remains of a whale found
near Rosedale. Mr D. Hoare kindly gave me this morning an
excellent series of photographs, and it is quite evident that the
remains represent a Hump-back Whale, which is technically
known as Megaptera nodosa ... Although we have a specimen
of this whale in the Museum, which was presented by Mr
Thomas Welsby, 1 should very much like to obtain the com-
plete set of bones if they could be secured without much
expense’. Riley replied and discussed potential problems as-
sociated with possible retrieval — factors no less relevant 65
years later. He recommended that the assistance of Mr F.G.
Collins of Rosedale Cattle Station be sought: *... The distance
and position where the skeleton [later noted to be ~ 15m long]
is situated render it a very difficult undertaking ... It would
take about 20 men and 4 or 5 motor lorries to accomplish such
a task on å weekend..." . Riley suggested a possible alternative
to road transport viz. that a boat tow, on a very calm day, a
lografton which the bones were loaded to Agnes Water where
motor vehicles had ready access to the water’s edge. Longman
wrote to Collins on 14 December who replied on 29 December
making no reference to the water transport proposal: *... One
of my men who knows the location well advises that it would
be very difficult to convey these large bones from where they
are buried in the sand to where it is possible to fit a conveyance
to bring them to the nearest Railway Station, a distance not
less than 40 miles. There is practically
no road from where the whale remains
are for about 4 miles and the road from
there to Rosedale is mostly bad and
rough. I am of the opinion that the
expense of recovering these bones and
conveying them to the nearest railway
would be far too heavy to warrant the
outlay ...". In Longman's hand-writing
there is an annotation to the letter: 'Mr
Collins visited the Museum on Dec.
Ziler. As far as is known the matter
rested there — as did the skeleton.

Eleven photographs were sent to
Longman. Two are shown here (Fig. 1).
It is likely, from the cleanness of the
skull and mandible as well as the state
of the soft tissues still attached to the
post-cranial skeleton [in photographs
not shown here], that the whale had
been stranded for some months, [ts
identification by Longman as a large
adult balaenopterid is clearly correct
but his suggestion that it was a hump-
back whale is considered erroneous.

The nasal bones are well demon-
strated in the photographs and their
anterior margins are concave and in-
clined forwards on their outer aspects,

` and mandible.
features seen among balaenopterids in

MEMOIRS OF THE QUEENSLAND MUSEUM

B. edeni and the dark shoulder form of the minke whale B.
acutorostrata only (Omura, 1959 & 1975). The large size of
the skull and mandible of the Rosedale whale excludes B.
acutorostrata. There are now two specimens of B. edeni in the
Queensland Museum, J21713 and JM4386, collected from
Tin Can Bay and the Great Sandy Strait in 1965 and 1983
respectively. Their diagnostic nasal bone characteristics have
been confirmed by Paterson & Van Dyck (1988). The skull of
the humpback whale donated by Welsby and referred to by
Longman is registered J3343 in the cetacean collection. Its
nasals (and those of three other humpback whale specimens
more recently collected) are antero-medially peaked and do
not resemble the configuration of those of the Rosedale whale
or the two B. edeni specimens in the Queensland Museum.
The anterior nasal margins of the Rosedale whale are well
posterior to the posterior maxillary concavity. Junge (1950)
considered this an important characteristic of B. edeni com-
pared with B. borealis in which the anterior nasal margins are
at a similar level to the maxillary concavity.

The preserved archival material illustrates perennial prob-
lems associated with the retrieval of large cetaceans and has
enabled a retrospective identification of B. edeni.

Literature Cited

JUNGE, G.C.A. 1950. On a specimen of the rare fin whale
Balaenoptera edeni Anderson, stranded on Puli Sugi
near Singapore. Zoologische Verhandelingen 9: 1-26.

OMURA, H. 1959. Bryde's whale from the coast of Japan.
Scientific Reports of the Whales Research Institute,
Tokyo 14: 1-33,

1975. Osteological study of the minke whale from the
Antarctic. Scientific Reports of the Whales Research
Institute, Tokyo 27: 1-36.

PATERSON, R.A. & VAN DYCK, S. 1988. Bryde's whale
in the coastal waters of eastern Australia. Scientific
Reports of the Whales Research Institute, Tokyo 39:
21-29,

R.A. Paterson, Queensland Museum, PO Box 3300, South

Brisbane, Queensland, Australia 4101; 11 March 1997.

FIG. 1. A, Dorsal view of skull of the Rosedale whale which was estimated to be Jam
in length. The pre-maxillaries are missing and the left maxillary is damaged. B, Skull

RE-EVALUATION OF EMYDURA LAVARACKORUM: IDENTIFICATION OF
A LIVING FOSSIL

SCOTT THOMSON, ARTHUR WHITE AND ARTHUR GEORGES

Thomson, S.A., White, Å. & Georges, A. 1997 06 30: Re-evaluation of Emydura lav-
arackorum: Identification of a living fossil. Memoirs of the Queensland Museum 42(1):
327-336. Brisbane. ISSN 0079-8835.

Post-cranial osteological characters can be used to diagnose Australian short-necked chelid
turtles to genus. Morphological examination of the Pleistocene fossil Emydura lav-
arackorum, from Riversleigh, shows that it is aligned with the genus Elseya not Emydura
and should be referred to as Elseya lavarackorum (White & Archer, 1994). Furthermore, the
fossil specimen is not distinguishable from an undescribed extant form of Elseya from the
Nicholson drainage, with which it shares one unique feature so this name should apply also
to this extant form, identified to date only from electrophoretic data, It is Australia's first
living fossil turtle, an extant population of a Pleistocene taxon. [] Chelonia, Chelidae;
Pleistocene, fossil, turtle.

Scott Thomson & Arthur Georges, Applied Ecology Research Group and CRC for Fresh-
warer Ecology, University of Canberra, P.O. Box 1, Belconnen ACT 2616, Australia; Arthur
White, School of Biological Sciences, University of New South Wales, P,O., Box I, Kensington

NSW 2033, Australia; 11 April 1997.

The taxonomy of Australian chelid turtles is
poorly known and in dire need of review (Cogger
ct al., 1983). Recent electrophoretic surveys
(Georges & Adams, 1992; 1996) have revealed
that in some instances, currently accepted species
boundaries are difficult to jusufy and in others,
what are currently regarded as single species are
in fact two or more, The detailed morphological
analyses required to verify these findings have
not been conducted (but see Thomson & Georges,
1996), and until recently it was not possible to
distinguish even between extant short-necked
genera on the basis of osteological characters
(Gaffney, 1977). This paucity of osteological data
suitable for distinguishing the extant genera
makes the identification of fossil forms, most of
which are incomplete specimens, difficult. In
many instances, chelid fossils have been assigned
Io either Chelodina or Emydura, with little or no
evidence presented to eliminate the possibility
that the short-necked forms among them may be
Elseya, Rheodytes or Elusor.

In 1994 a partial carapace and associated plas-
iron trom Riversleigh was described as å new
species, Emydura lavarackarum, by White A
Archer (1954). The fossil specimen was from
Terrace Site, a fuviatile site on the Gregory
River. These authors interpreted the sediments as
heing Pleistocene in age because of the presence
of remains of Diprotodon optatum. (White &
Archer, 1994), The holotype consists of the ante-
rior halt of the carapace with some antenor pe-
ripherals and an essentially complete plastran

with some pelvic material present. The length of
the plastron is 390mm (White & Archer, 1994)
which corresponds to a carapace length of ap-
proximately 420mm. Two other plastra from the
same site were also collected but not described,

White and Archer (1994) assigned the speci-
mer to Emydura on the mode of the insertion or
the anterior bridge into the ventral surface of the
carapace. They found that in the derived state, the
anterior bridge is angled steeply backwards to-
wards the rib/gomophosisis (called transverse
process in White & Archer, 1994), whereas in all
other chelids the anterior bridge was found to
form a continuous line with the rib/gomophosisis.

In this paper, we reassess the generic assign-
ment of the fossil by comparing the fossil material
with post-cranial character states we have found
useful 1n separating extant genera of Australian
short-necked chelid turtles. We also propose that
the fossil taxon is extant, å distinctive, undescribed
form closely aligned with Elseya dentata.

MATERIALS AND METHODS

Specimens of each of the short-necked species
identified using electrophoresis by Georges and
Adams (1996) were obtained from museums, the
Conservation Commission of the Northem Terri
tory and the University of Canberra. Where forms
have not been included in published keys or de-
scriplions, the specimens were selected from
those lodged as vouchers to accompany rhe elec-

328 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Ventral view of the anterior carapace of short-necked turtles showing the bridge-carapace suture (BCS) the
rib/gomophosisis (R) on pleural | (PI) and their relationship to the vertebral column (V) and the peripherals (Pe). A-B,
Pseudemydura (UC0178). C-D Elseya sp. aff. E. latisternum (Manning) (QM59289); E-F, Elusormacrurus (UC0184).

IDENTIFICATION OF ALIVING FOSSIL

Irophoretic data. The specimen collection was
supplemented by limited field sampling.

Each specimen was skeletonised by removing
excess soft tissue and feeding the remaining car-
case to dermestid beetles. The skeletal material
was bleached in 5% sodium hypochlorate solu-
lion, and the process stopped by immersion in
100% ethanol. Plastra were separated from cara-
paces by disarticulating the plastral-carapacial
sulure between the hyo- and hypoplasira of the
plastron and the lateral peripherals of the cara-
pace, This was done by the carefully heating the
carapace until the sutures become mobile and the
plastron was then gently prised off. This also
required disarticulation of the pelvis from the
carapace, Characters potentially diagnostic at the
generic level were examined to establish their
consistency across a range of specimens within
the polytypic genera Elseya and Emydura, and
across å range of specimens within each species,

The fossil specimens of Emydura lav-
arackorum were examined to determine the pres-
ence of character states which are generically
diagnostic in extant taxa, The fossil specimen was
then assigned to genus.

Throughout this paper, we refer to å generic
group as à group of species that are sufficiently
distinct collectively to warrant recognition at the
level of genus, though this has not yet been for-
mally established. These groups were first iden-
iified by Legler (1981), have a foundation in
electrophoretic studies (Georges & Adams,
1996), and have been referred io since several
limes in the literature. In contrast, å species com-
plex is å group of species, all but one of which are
undescribed, which together presumably repre-
sent a distinct clade but which are not considered
distinctive enough to warrant recognition at the
level of genus.

We refer to the Elseya dentata species complex
as comprising the distinctive forms of Elseya
from coastal Queensland currently assigned to
Elseya dentata, and the Northern Territory forms
including Elseya dentata (sensu stricto) and
Elseya sp. aff, E, dentata from the Alligator Riv-
ers region (Georges år Adams, 1996). The Elseya
dentata generic group (sensu Legler, 1981) com-
prises the Elseya dentata species complex plus
Elseya novaeguineae and Elseya branderhorsti
from New Guinea, The Elseya latisternum ge-
neric group comprises Elseya latisternum (sensu
stricto), a related form from the headwaters of the
Darling River drainage and a sibling species pair
from coastal New South Wales (Georges &
Adams, 1996; Thomson & Georges, 1996), The

329

later three are currently undescribed. It is not the
purpose of this paper to describe new genera, so
for consistency, we use the nomenclature of
Georges & Adams (1992) and Legler (1981) and
recognise six groups of Australian short-necked
chelid al generic level: Elusor, Emydura,
Rheodwes, — Pseudemydura, the — Elseya
latisternum generic group and the Elseya dentata
generic group.

Throughout this paper, names of the bony ele-
ments of the shell and the overlying scutes follow
those of Zangerl (1969). A complete list of the
Specimens examined in this study will be found
in Appendix A.

RESULTS

Five characters were identified as diagnostic at
generic level, Where polarity is indicated, it was
determined hy comparison with South American
chelids and African pelomedusids in à cladistic
analysis (Thomson & Georges, unpublished
data). Only those characters relevant io the iden-
tification of the fossil specimen are presented.

ANTERIOR BRIDGE STRUTS. Character A.
Contact with Pleural |.

AU. In the primitive state, the posterior edge of
the bridge-carapace suture runs parallel and adja-
cent to the rib/gomophosisis of pleural 1 (Fig. 1 A-F).

Al. In the derived state, the posterior edge of
this sulure contacts the rib/gomophosisis at its
anterior end, but is set at a forward divergent
angle of between 15 and 50°. This angle is most
pronounced in Emydura, least in Rheodytes (Figs
2A-F, 3A-D).

Character B. Bridge suture shape.

Bl. The anterior and posterior edges of the
bridge-carapace suture diverge from their point
of congruence closest to the vertebral column,
The widest extent of the sulure is distal lo the
vertebral column and there is no medial constric-
tion (Fig. LA-F)

B2. The anterior and posterior edges of the
bridge-carapace suture are parallel or closely so
with a prominent suture surface between them.
There is no medial constriction (Figs 2A-B, E-F
3A-B]J.

B3. The bridge-carapace suture is expanded for
its full length, but more so at extremes, there
being an obvious medial constrietion (Fig. 2B).

B4. The bridge-carapace suture narrows from
ils widest point proximal to the vertebral column,
and constricts completely to form a ridge conflu-
ent with the edge formed by the ventral suture uf
the peripheral bones (Fig. 3C-D).

330 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Ventral view of the anterior carapace of short-necked turtles showing the bridge-carapace suture (BCS) the
rib/gomophosisis (R) on pleural 1 (PI) and their relationship to the vertebral column (V) and the peripherals (Pe). A-B,
Rheadytes leukops (UC0173). C-D, Elseya dentata (QM59277). E-F, Elseya lavarackorum (extant) (QM46284).

IDENTIFICATION OF A LIVING FOSSIL

FIG. 3. Ventral view of the anterior carapace of short-necked turtles showing the bridge-carapace suture (BCS)
the rib/gomophosisis (R) on pleural I (PI) and their relationship to the vertebral column (V) and the peripherals
(Pe). A-B,Elseya lavarackorum (fossil) (QM24121). C-D, Emydura subglobosa (UCO172).

RIB/GOMOPHOSISIS OF PLEURAL 1. Char-
acter C. Rotation of the Rib/Gomophosisis.

CO. The ventral surface of the distal extent of
the rib/gomophosisis is rotated obliquely, to face
ventrally but with posterior inflection (Figs 1 A-F,
2A-B).

CI. The rib/gomophosisis shows no such tor-
sion distally (Figs 2C-F, 3A-D).

DORSAL CHARACTERS. Character D. Rela-
tive width of Vertebral 1.

D1. Ist 3 vertebral scutes equal or subequal in
width (Figs 4A-D, 5B).

D2. 1st vertebral scute wider than 2nd and 3rd
(Figs 4E-F, 5A).

Character E: Cervical Scute.
EQ. Cervical scute typically present (Fig. 5B).

332 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Dorsal view ofthe anterior carapace of short-necked turtles showing the relative size between the vertebral
scutes (V) and the presence or absence of the cervical scute (N) their relationship to the costal scutes (C) and
marginals (M). Note the indentation at the anterior of some taxa. A, Pseudemydura (UCO178). B, Elseya sp.
aff. E. latisternum (Manning) (QM59289). C, Elusor macrurus (UC0344). D, Rheodytes leukops (UC0173). E,
Elseya dentata (QM59277). F, Elseya lavarackorum (extant) (QM46284).

IDENTIFICATION OF A LIVING FOSSIL 333

FIG. 5. A-B, Dorsal view of the anterior carapace of short-necked turtles showing the relative size between the
vertebral scutes (V) and the presence or absence of the cervical scute (N) their relationship to the costal scutes
(C) and marginals (M). Note the indentation at the anterior of some taxa, A, Elseya lavarackorum (fossil)
(QM24121). B, Emydura subglobosa (UCO172). C-E, Ventral view of the plastrons showing the arrangement of
the sulci between the humeral (H) and pectoral (P) scutes, also shown are the gular scutes (G) and the intergular
(D. C, Elseya lavarackorum (extant) (QM46284); D, Elseya lavarackorum (fossil) (QM24121). E, Elseya

dentata (QM59277).

E1. Cervical scute typically absent (Figs 4E-F,
5A).

The distribution of the character states for each
taxon is provided in Table 1. The holotype
Emydura lavarackorum had a combination of a
widely divergent angle (45?) between the anterior

bridge suture and the rib/gomophosisis of pleural
one; parallel anterior and posterior edges of the
bridge-carapace suture throughout their length,
widely spaced, with no medial constriction; no
distal rotation of the gomophosisis of pleural one;

334

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE |, Character matrix, Distribution of the key character states among taxa. Abbreviations (s =mi, of species

examined in group, n — no, of specimens),

olymorphic characters shown: Pseud, Psexdemydura; Elat, Elseva

latisternum group; Elus, Elusor; Rheo, Rheodyres; Else, Elseya dentata group; Elno, Elseya novaeguineae;
EQld, Queensland Elseya group; Elirv, Elseya luvarackorum (holotype); Emyd, Emydura.

Elus (s=

"t. (s=2) | Elno(s=1)
gen (n=2)

EQld (s23
(n=10)

Elav (821)
{n=1)

Enyd (s241

Pseud(s=1) | Hat ise)
(n=2) (020) (sl SN
CwxeA| o [| a | o |

Charu] 1 Ja Ta |

ET [ o.

Character D HeH
Cmecek| 0 | m | o |

a first vertebral scute that was markedly wider
than vertebrals 2 and 3; and no cervical scute.

A significant feature of Emydura lav-
arackorum, though difficult to quantify, was an

indentation of the carapace margin in the area of

the cervical cleft and first marginal scutes. This
feature is held in common with Lurtles in the
Elseya latisternum group and Pseudemydura, is
variable among the Queensland forms of Elseya
dentata, and never present in the Northern Terri-
tory and New Guinea forms of Elseya denrata nor
in Elusor, Rheodytes and Emydura. Although not
considered a useful character at generic level, we
will use it in combination with other similarities
to establish a close relationship between the fossil
Emydura lavarackorum und an extant form of
Elseya from the Nicholson River.

DISCUSSION

The bridge carapace suture runs parallel and
adjacent to the rib/gomophosisis in species of the
Elseya latisternunt group, Pseudemydura and
Elusar and so can be clearly distinguished from
the fossil Emydura lavarackorum (Table 1). Ro-
tation of the rib/gomophosisis of Pleural 1 elimi-
nates Rheodytes às à possible identification lor
the fossil, leaving only the Elseya dentata generic
group and Emydura as possibilities.

Two sub-groups within the Elseya dentata ge-
nenc group can be distinguished. The first com-
prises Elseya denjata (sensu stricto), Elseya
novaeguineae, Elseya branderhorsti, and Elseya
sp. (Vogelkopf Region, PNG; Anders Rhodin,
pers. comm) and Elseya sp. (South Alligator
River, NT; Georges & Adams, 1996), The second
sub-group is restricted to Queensland (Queens-
land Elseya demata sub-group) and comprises
Elseya sp. (Nicholson), Elseya sp. (Johnstone),
and Elseya sp, (Burnett) (Georges & Adams,
1996). Generic recognition of these sub-groups is
not suggested,

1) | Rhea (sel)
el!

Emydura lavarackorum possesses all charac-
ters [hal are consistent across species ol the
Elseya dentata generic group (Table 1) and, more
significantly, all characters uniquely possessed
by the Queensland Elseya dentata sub-group
(Table 1). Of those characters which separate
Emydura from the Elseya denrara generic group,
the fossil consistently possessed character stales
which distinguished it from Emydura. Therefore,
we assign Emydura lavarackorum |o the genus
Elseya as Elseya lavarackorum (White & Archer,
1994).

Since the description of Elseya lavarackorum,
specimens of the extant Elseya sp. (Nicholson
drainage, Georges & Adams, 1996) have becorne
available, The two forms are indistinguishable in
every diagnostic character, including the indenta-
tion of the anterior margin of the carapace, A
unique feature of the Nicholson population, when
only extant forms are considered, is the sigmoidal
shape of the sulcus between the humerals and
pectorals on the plastron (Fig. 5C), this sulcus is
straight in all other species of the Elseya dentata
generic group (Fig. 5E). This feature is present in
the holotype of Elseya lavarackorum (White &
Archer, 1994) and in one (QM30818) of the ad-
ditional fossil specimens now available (Fig. 5D).
The anterior plastron is absent from the third
fossil specimen (QM30817),

In contrast, the fossil has strong] d embossed,
rounded peripherals i in the region adjacent To the
bridge, a feature not present in the 15 specimens
from the Nicholson population, This is a similar
condition to that found in aged, adult individuals
in a number of species, i.e., individuals which are
large for their species, such as Elusor macrurus
(specimens over 400mm), Elseva sp. all, E
dentata from the Burnett River (specimens over
380mm) and Emydura subglobosa from the
Gregory and Reynolds Rivers (specimens over
250mm). We consider this trait to be essentially
a feature of large aged specimens in a range of

IDENTIFICATION OF A LIVING FOSSIL

chelid turtles. None of the turtles examined from
the Nicholson drainage had carapace lengths in
excess of 320mm, well below the maximum size
for species in the Elseya dentata generic group.

In species level taxonomy, the onus ison differ-
ential diagnosis. The shell of the fossil holotype
15 adequately preserved for diagnostic purposes.
We therefore propose that, in the absence of any
diagnosable difference and the relatively young
age of the fossil material, Elseya lavarackorum
and the Nicholson Elseya sp. aff. E. denrara be
regarded as a single species, It is Australia's first
living fossil freshwater turtle, an extant popula-
tion of a Pleistocene taxon. We do not propose
That allochronic subspecies be recognised,

ACKNOWLEDGEMENTS

The authors would like to thank Patrick Couper
(Queensland Museum), Ross Sadlier (Australian
Museum), Paul Homer (Museum and Art Galler-
ies of the Northern Terntory) and Laurie Smith
{Museum of Western Australia) for the loans of
fossil and skeletal material and for access to the
museum turtle collections; Peter Whitehead for
å large donation of material stored at the Conser-
valion Commission of the Northern Territory:
John Cann, John Legler and Gerald Kuchling for
the loan of material from their private collections
and the Department of Environment and Heritage
in Queensland for permits to collect in the Nich-
olson Drainage of north west Queensland. We
would especially like to thank Simone Reynolds
of the University of Canberra før the illustrations
in this paper and Peter Ogilvie for the photography.

APPENDIX A

Specimens Examined; All names used [or un-
described species are from Georges & Adams (1992,
1996), Abbreviations used: AM, Australian Museum;
NTM. Museum and Art Galleries of the Northern Ter-
ritory; QM, Queensland Museum; WAM, Western Aus-
tralian Museum; UC, University of Canberra; UM,
University of Michigan Field Series: UU, University
of Utah,

Elusor macrurus: UCOT84-93, 0225-29, 0344,
UU19488, 19508; Elseva dentata: NTM13319, 13521,
16330, QM59265, 59277-80, UCO0307-18; Elseya
Jatisternum; AM123037, 123039, 125474-75,
QM48054-55; Elseya novaeguineae: AMA2662,
125038; Elseya lavarackorum: QMF24121, F30817-
18 (fossil), QM31939, 31942. 51944, 31946-47,
31949-50, 31952, 46284, 47908, 47911, 48544, 48547,
60255, UCO201 (extant); Emydura macquarii
QM48016, 48034, 48050-51. 59275-76, UCO175-76,

LITERATURE CITED

CANN, J. & LEGLER, J.M. 1994, The Mary River
lurtarse: å new genus and species of short-necked
chelil from Queensland, Australia (Testudines:
Pleurodira}. Chelonian Conservation and Biology
1(2); 81-96

COGGER, H.G., CAMERON, EE & COGGER, IH, M.
1983, Zoological Catalogue of Australia, Volume
I, Amphibia and Reptilia (Australian Govern:
ment Printing Service: Canberra). 313pp.

GAFFNEY, ES 1977. The side-necked turtle family
Chelidae: a theory of relationships using shared
derived characters. American Museum Novilates
2620: 1-28.

GEORGES, A. & ADAMS, M. 1992. A phylogeny of
Australian chelid turtles based on allozyme elec-
trophoresis. Australian Journal of Zoology 40:
453-476.

1996, Electrophoretic delineation of species bound-
aries within the short-necked freshwater turtles of
Australia (Testudines: Chelidae). Journal of the
Linnean Society 118:241-260.

LEGLER, J.M. 1981. The taxonarny, distribution, and
ecology of Australian freshwater turtles. (Tes-
tudines: Pleurodira: Chelidae). National Geo-
graphie Society Research Repons 13: 391-404.

THOMSON, S & GEORGES, A. 1996, Neural bones
in Australian chelid turtles. Chelonian Conserva:
tion and Biology 2(1): 82-86.

WHITE, A. & ARCHER, M. 1994, Emydura lav-
arackorum, à new Pleistocene (urtle (Pleurodira:
Chelidae) from fluviatile deposits al Riversleigh,
Northwestern Queensland. Records of the South
Australian Museum 27: 159-167,

ZANGERL, R. 1969. The ture shell. Pp, 311-240. In
Gans, C., Bellairs, D. d'A. & Parsons, T.A. (eds)
Biology of the Reptilia. Vol T, Morphology ^.
(Academic Press: London).

0303; Emydura subglobusa: NTM5028, 8206, 13428,
13433, 16332, UC0171-72, 0177: Emydura victoriar:
NTM13513-14, 32917, 32976, UC 0165; Elseya sp
alf. E. dentata (Burnet): UC 0305-6, QM2966, 28449,
36036, 36039, 36041-42, 36044-47, 37933, 38533,
59269-71; Elseya sp. alf. E. dentata (Johnstone):
QM22694, 23175, 23299, 23300, 23322, 24938,
28449, 48060, 48068; Elseya sp. aff. E. dentata (South
Alligator): AMI28002, 123004, QM59285-89,
NTM35097, 13512, 13985, UC0304; Elseya sp. aff. E
latisternum. (Gwyder): AM123028-29, QM 48028,
48038. Elseya sp. aff. E. lansternum (Bellingen):
AMI38387-88, UM02016-17; Elseva sp, aff. E.
latisternum (Manning): AM123040, 123042, QM-
59289-90; Emydura sp. all, E, victoriae (Daly Mission)
AMI25470-71, 125491, NTMS8211, 8213, 17339;
Pseudemydura ` umbrina: UCO] 78, WAM?29337;
Rheodytes leukaps; CO173.

XEROMYS MYOIDES THOMAS, 1889 (RODENTIA: MURIDAE) IN MANGROVE

COMMUNITIES OF NORTH STRADBROKE ISLAND, SOUTHEAST QUEENSLAND

STEVE VAN DYCK

Van Dyck, 5, 1996 06 30; Xeromys myoides Thomas. 1889 (Rodentia: Muridae) in mangrove
communities of North Stradbroke Island, southeast Queensland. Memoirs of the Queensland
Museum 42(1): 337-366. Brisbane. ISSN 0079-8835.

Seven mangrove communities spanning the north-south length of western North Stradbroke
Island were surveyed between 1991 and 1993 for Xeromys nryoides. It was recorded at all
sites, a total of 111 individuals was captured and home range estimates and ecological data
were recorded for nine individuals. The life history of this small, spotted, specialised,
non-aquatic, non-arboreal, nocturnal rodent revolved around the island's mangrove and
adjacent sedgeland communities. It nested in either large termitarium-like mounds or simple
tunnels in the supralittoral bank. Nests contained up to eight individuals of all age groups
und either sex, with one adult male present in the resident group. From their nests, rats
followed the receding tide out through sedgeland and into the mangroves where they loraged
over å home range estimated at approximately 0.8ha for males and 0.6ha for females. Home
ranges overlapped slightly, but core areas, estimated at around 0.2ha, did not. Home range
size Was thought to be determined by the complexity of the mangrove community as reflected
in the structural diversity of the mangrove substrate, The diet of X. myoides consisted of
small crustaceans (mainly the crabs Parasesarma erythrodactyla and Helice leachi), marine
polyclads, marine pulmonates and marine bivalves. The species was recorded in breeding
condition throughout most of the year. It is proposed that the unusual life-style of this rodent
might be linked to problems associated with the need to dissipate heat rather than the necessity
Lo retain it. Caution should be exercised in attributing its locally abundant status on North
Stradbroke [sland to populations on the mainland which await assessment. [] Xeromys, False
Waler-rat, rodents, mangroves, North Stradbroke Island.

Steve Van Dyck, Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101,

Ausiralia; 20 December 1996.

The False Water-rat Xeromys myoides Thomas,
1889 (Fig. 1) is a small, poorly known Australian
hydromyine rodent. Fourteen museum specimens
ure known (Van Dyck & Durbidge, 1992). These
and other specimens liberated after capture, have
been collected from mangrove forests, freshwater
lagoons, swamps or sedged lakes close to
foredunes in the Northern Territory (Parker,
1973; Redhead & McKean, 1975; Magnusson
et al., 1976), coastal central Queensland
(Thomas, 1839; McDougall, 1944) and coastal
southeast Queensland (Dwyer et al., 1979; Van
Dyck et al., 1979). It is not known whether the
paucity of this record reflects genuine rarity or
the inadequacy of searching effort in the appro-
priate habitat, however Ihe animal is currently
classified under the Commonwealth Endangered
Species Protection Act 1992 as "vulnerable".

Xeromys myoides has been known from North
Stradbroke Island, Queensland since 1978 when
an adult female was hand-caught by students at
Myora Springs (Van Dyck et al., 1979), This
specimen represented the most southerly record
lor the species. In 1992, Ellie Durbidge and I
recorded the capture of another adult female

irapped near one of à number of reed-covered
peat mounds in sedgelands of the Myora wetlands
(Van Dyck, 1992, 1994; Van Dyck & Durhidge.
1992). These reports commented on the similarity
of the large, termitarium-like nesting mounds to
that described by Magnusson et al. (1976) from
Melville Island but, in addition, noted the pres-
ence of relatively simple tunnels built into the
supralitloral bank al the marine-sedgeland/
freshwater-wallum (Banksia tree-heath on humus
podzols) ecotonc. In tentatively attributing these
simple tunnels to X. myoides the possibility was
raised that the absence of large nesting mounds
in appropriate habitat may not be å reliable gross
indicator of the absence of X. mvoides.

In an attempt 10 clarify the distribution of X.
myoides 3n the mangrove communities of North
Stradbroke Island, nest recording and trapping
surveys were conducted from near Amrty on the
island's north-westerly tip, to Jumpinpin, the
island's southern extremity 36km south (Figs 2-
6). Radiotelemetry of animals trapped and re-
leased was expected lo generate some
preliminary information for the species on dict,

338

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Xeromys myoides, adult female from Rainbow Channel, North Stradbroke Island (Bruce Cowell).

home range, foraging behaviour and aquatic ad-
aptation.

METHODS

FIELD SURVEYS. Field observations and surveys
associated with this study were made between 3
Nov 1991 and 18 Nov 1993. Fieldwork prior to
Sept 1992 concentrated on nest appraisal, capture
and radiotelemetry of individuals at and near the
Rainbow Channel (site 2, Fig. 3), while the latter
part documented the presence of X. myoides in six
other areas of mangrove habitat along the western
side of the island (Figs 2-6).

CAPTURE TECHNIQUES. Elliott (Elliott Sci-
entific, Upwey, Victoria) size A aluminium traps
(300x100x90mm) were used. Up until Sept 1992
trapping was conducted only at or near nesting
mounds, none of which was found constructed
inside the mangrove zone but in sedges of the
supralittoral zone. Thereafter, given the initial
results of direct observation through radio-
telemetry, traps were laid along the floor of the
mangrove forest. A white cotton string-line guide
was laid simultaneously. Traps, baited with

chunks of chopped mullet or gar, were placed
among mangrove roots, beside trunks with base
hollows, among pneumatophores or inside hol-
low logs. Traps were not set up in trees.

Normal terrestrial trapping protocol was ad-
justed to accommodate threats of drowning im-
plicit in the method. Live-trapping on the
mangrove floor required local knowledge of both
the upper inundation height of night high tides
and the rates of incoming tides as traps required
checking and tripping prior to their inundation. A
rising tide combined with too many traps set over
a long mangrove transect (e.g., 75 traps over
Ikm) proved a drowning hazard to trapped rats
and a formidable task for one person in a night.
Looping of mangrove traplines (25 traps per loop)
north and south of a ‘central’ land-based camp
gave adequate coverage of mangrove forest, and
quicker access to both traps and the station asso-
ciated with processing captures. Trapping was
carried out during the following periods: Nov
199] (8 trap nights); Feb 1992 (13 trap nights);
May 1992 (249 trap nights); June 1992 (29 trap
nights); July 1992 (196 trap nights); August 1992
(188 trap nights); September 1992 (263 trap
nights); October 1992 (50 trap nights); November

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

MORETON
BAY

oc, {Bro Mile

BRISBANE `
u | NORTH

| /

|
i
4 }

27-40! m

i i |
Tv Stockyard
AN BE

oe

ØChigail.Chiggil
® Rainbow Channél
fl Myora Springs; k

® Deanbila /
!

rå OA | STRADBROKE
3 " | ISLAND /

114, 8 Canalpin Creek

condition and individual traits
(tail kinks, albinism in tail-tip,
ear damage etc.). Initial diffi-
culty in determining sex in ju-
veniles and sub-adults was
resolved using a measure of the
distance between the anus and
the urogenital opening (‘anal-
genital distance’) which was
significantly less in females
] (Table 2). X. myoides rarely
struggled, and gave the impres-
sion of being almost incapaci-
tated by fear. However, when
handled with ungloved hands
they inflicted deep bites, so
were restrained for measuring
through a cloth collecting bag
rolled back to give access to the
animals. Dorsal spotting pat-
terns were recorded (Table 2),
and each animal was assigned
to an age category by weight: J
(juvenile) «28g; SA (sub-
adult) 28-35g; A (adult) 235g.
Each animal was individually
tagged. Initially, animals were
tagged on one ear with a num-
bered brass tag (Hauptner-
Ohramarken #73850) or a "fish
& small mammal tag” (No.

/

SOUTH
PACIFIC
OCEAN

FIG. 2. Sites (1-7) surveyed for Xeromys myoides on North Stradbroke Island.

1992 (50 trap nights); December 1992 (100 trap
nights); Jan 1993 (50 trap nights); March 1993
(70 trap nights); May 1993 (50 trap nights); July
1993 (40 trap nights); October 1993 (40 trap
nights); November 1993 (36 trap nights), Trap
nights totalled 1480 and covered sites at Chiggil
Chiggil, Rainbow Channel, Myora Springs, Two
Mile, Deanbilla, Canalpin Ck and Stockyard (see
Table 1, Figs 2-6).

In open Avicennia marina woodland such as at
Canalpin Creek, where old trees were widely
spaced and the substrate simple, it was possible
to spotlight for X. myoides among pneumatophores
and capture them hy hand.

No attempt was made to dig individuals from
nests and no nesting mounds were breached.

HANDLING AND RADIO-TRACKING PRO-
CEDURES. Captured individuals were weighed,
measured, sexed, and assessed [or reproductive

153"30'

1005-1, size 1, National Band
and Tag Co, Newport, KY).
Both tag types proved unsatis-
factory and were found ta have
been ripped from ears after a
few days. Finally, å system of ear nicks was used.
A simple 2mm cut was made in the edge of the
pinna with small surgical scissors dipped in tattoo
ink. Using well-defined areas of the upper and
lower folds of the pinnae. a wide range of iden-
tificatory combinations was available. Ear cuts
healed and rejoined quickly, leaving a fine black
line readily discernable when torch-illuminated
from behind the pinna.

Trapped animals were measured (Table 2) and
released before dawn al their point of capture, or
their nest (where known). In cases where a nest
site was not known for an individual and the
incoming tide had covered the capture site, the
animal was deposited nearest the trap site in a low
mangrove hollow that provided cover and easy
access to the ground. Animals responded pas-
sively but poorly to being held over for measure-
ment during the day following their capture, and

340 MEMOIRS OF THE QUEENSLAND MUSEUM

ET.

Amity Point.

Ce

^»
Drm

f
Ee ki
e ni, Cafe,
IM

NORTH STRADBROKE

[3] Avicennia marina var australasica Tall open-shrubland and Tall shrubland

[4] Avicennia marina var australasica Open-scrub and Closed scrub
[5] Avicennia marina var australasica Low open-woodland, Low woodland and Woodland

[7] Avicennia marina var australasica Low open-forest and Open forest
| [18] Rhizophora stylosa Tall shrubland, Open-scrub and Closed scrub
[=] Land subject to tidal inundation comprising salt water couch and or samphire flats or marine clays

| fe] Land subject to permanent or seasonal inundation

FIG. 3. Sites and vegetation types surveyed for Xeromys myoides on North Stradbroke Island at (1) Chiggil
Chiggil and (2) Rainbow Channel (after Dowling, 1986).

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND 341

The Bluff

NORTH
STRADBROKE

ISLAND .

[4] Avicennia marina var australasica Open-scrub and Closed scrub
[5] Avicennia marina var australasica Low open-woodland, Low woodland and Woodland

[7] Avicennia marina var australasica Low open-forest and Open forest

[18] Rhizophora stylosa Tall shrubland, Open-scrub and Closed scrub

EI Land subject to tidal inundation comprising salt water couch and or samphire flats or marine clays
E

] Land subject to permanent or seasonal inundation

Scale 1: 25,000.

1 METRES. 4 KILOMETRES.

FIG. 4. Sites and vegetation types surveyed for Xeromys myoides on North Stradbroke Island at (3) Myora Springs,
(4) Two Mile and (5) Deanbilla (after Dowling, 1986).

342 MEMOIRS OF THE QUEENSLAND MUSEUM

Pelican

"Seagrass

Black Snake

Lagoon

Seagrass

Seagrass

e

4

^.
"2
iss

oR - ENG \

[=] Land subject to tidal inundation ert salt water couch and or samphire flats or marine clays
| fe] Land subject to permanent or seasonal inundation i |

Scale |: 28,000
MET! `
| RES ü A 2 KILOMETRES q

FIG. 5. Sites and vegetation types surveyed for Xeromys myoides on North Stradbroke Island at (6) Canalpin Ck
(after Dowling, 1986).

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND 343

METRES. V UN MLOMETRES,

[3] Avícennia marina var australasica Tall open-shrubland and Tall shrubland
| [4] Avicennia marina var australasica Open-scrub and Closed scrub
[5] Avicennia marina var australasica Low apen-woodland, Low woodland and Woodland
| [7] Avicennia marina var australasica Low open-forest and Open forest
| [8] Aegiceras comiculatum Low open-shrubland, Low shrubland and Open-heath
[11] Aegiceras comiculatum Tall shrubland, Open-scrub and Closed-scrub
| [15] Ceriops tagal var australis Tall open-shrubland, Tall shrubland, Open-scrub and Closed-scrub
| [=] Land subject to tidal inundation comprising salt water couch and or-samphire flats or marine clays
| [4] Land subject to permanent or seasonal inundation

FIG. 6. Sites and vegetation types surveyed for Xeromys myoides on North Stradbroke Island at (7) Stockyard
(after Dowling, 1986).

344

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE I. North Stradbroke Island sites surveyed, vegetation types and trapping results for Xeromys myoides.

Site
1 Chiggil Chiggil 27725'00-10"S 153?26'13-19"E

Latitude/Longitude block

Recaptures | Individuals

Vegetation type(s)

2 Rainbow Channel

27°27°06"-28'00"S 153?25'22-49"E

3 Myora Springs

27*28'10-21"S 153925" 18-26"E

3,5 64
2,3,5,7,18 1153

4,5 25

4 Two Mile

27728'43-53"S 153°24°31-49"E

5 Deanbilla
6 Canalpin Ck

27°30'48"-31'09"S 153?24'41-47"E
27?36'08-44" S 153724'22-42"E

7 Stockyard

27*43'28-46"S 153°24'17-30"E

= jo j= |o |o

Totals

51 I

in some cases behaved in an ataxic manner when
released the following evening. Rats used in radio
tracking were fitted with a Titley (Ballina, NSW)
*Microlite', two-stage transmitter either attached
to a cable-tie collar with a protruding 15cm whip
antenna or epoxy-potted into a small rectangular
tablet (length 17.7mm, breadth 9.3mm, height
6.8mm) with a protruding 18.5cm antenna. The
ventral surface of the epoxy disc transmitter was
glued to a 20mm x 10mm piece of cloth which
was in turn glued to the animal's rump (Selley's
Supa Glue) after a square of fur approximating
the dimensions of the cloth had first been cropped
close to the rat's skin. The short hair that remained
provided a bonding surface for the glue on the
transmitter cloth and long hair surrounding the
positioned disc was then glued over the dorsal
surface of the cloth and transmitter. Given the
small size of the animal and the saline nature of
its environment, battery size (hence longevity and
range) had to be traded against weight of the
waterproofing agent protecting the unit. Total
weights of transmitter packages were approx-
imately 1.8g (collar) and 1.3g (glue-on). As re-
capture of individual rats was unreliable, collars
were abandoned early in the study in favour of
glue-on units that could be removed more easily
by the rodents, A drawback associated with glue-
on units was the ability of rats to twist around and
bite the transmitter package. Any intrusion of
saltwater into the unit resulted in electrolysis that
rendered it inoperable. Glue-on units usually only
lasted on an individual for two nights. A 2000
channel *Regal 2000" (Titley) portable receiver
operating on the 150.000-151.999mHz band, and
a 2 element hand-held H-frame antenna were
used to locate animals.

HOME RANGE DETERMINATION. Home
range estimates were determined between May
1992 and March 1993 for four adult females, four
adult males and one sub-adult male (Table 3).
Because visits to the area were irregular and

transmitter life was often unpredictable, animals
were tracked soon after capture and thereafter for
one or two nights. Length of tracking period was
always dependent on the time of capture mea-
sured against height and time of the night high
tide. As animals usually stayed in their nest until
the tide receded, up to four hours of telemetry
might be lost in a night to a high tide. All X.
myoides studied on North Stradbroke were noctur-
nal. Tracking was continuous and direct observa-
tion was used to maximise return of ecological
information. Places visited on a rat's traversed
route were marked with flagging tape complete
with time of recording and subsequently sur-
veyed during the day.

The pattern of habitat use by X. myoides was
not uniform throughout its entire home range and
a percentage usage pattern defined by the 95th
and 65th percent probability isopleth proved a
most desirable method for estimating home ranges
and core areas. Swihart & Slade (1985) have
commented that direct observation through con-
tinuous tracking may lead to an underestimation
of home range size through loss of independence
of fixes. To test this, ten-minute fixes were ex-
tracted from examination of each of the nine
routes plotted through time. Home range esti-
mates arrived at through this method greatly un-
derestimated observed patterns of usage and
failed to demonstrate a picture of overlap. Fixes
taken for each animal followed are outlined in
Table 3. Estimates of home range area were gen-
erated using the harmonic mean distance mini-
mum (HM, see Dixon & Chapman, 1980) and the
minimum convex polygon (MCP: Mohr, 1974).
The utility and appropriateness of the HM for
describing home ranges of small, fast-moving
mammals was demonstrated by Quin et al. (1992).
Estimates were generated with the RANGES 4m
computer package.

SPOTLIGHTING, DIET ASSESSMENT AND
NEST ENDOSCOPY. Radio-tagged animals

Head-Bod

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

Mean

Length

86.4-123.3

83.1-119.8

8.99

sub-adult
(28-35g)

83.4-99.0
76.0-103.5

juvenile
«28g

70.2-94.1

62.3-94.8

Hind Foot Length (su)

Tail-Vent Length
adult 43M 81.8 69.3-90.0 6.50
26F 80.0 74.4-87.5
sub-adult 7M 76.0 67.8-91.0
23F 73.8 67.7-84.2
juvenile 5M 63.8 59.0-70.8

7F 59.7-74.1

adult

sub-adult

juvenile

sub-adult

juvenile

Ear (from it

43M 21.6-24.6 0.75

3.23

21.0-24.1

220-240

23F 21.7 20.9-22.8 0.54

210 19.7-21.7 074 |

10.5-14.0 0.72
10.7-14.4 0.79

11.2-12.3 0.37
10.1-13.8 0.87

104-13.1

100-128 | 096 | 813 |

Anal-Genital Distance

8.72

42M 17.3 10.8-23.2
23F 7.0 4.1-9.7
dub-adult 7M 14.1 12.5-17.3
22F 5.8 3.5-7,0
juvenile 5M 11.2 8.0-16.2
JF | 61 5.4-6.4
Weight (g)
adult 42M 46.6 36.0-59.0
26F 42.0 36.0-54.0

sub-adult

28.0-35.0
28.0-35.0

juvenile

| Spotting

18.0-26.0
| 18.0270 |

adult

sub-adult

Juvenile

345

TABLE 2. Measurements, and incidence of dorsal
spotting for Xeromys myoides from North Stradbroke
Is. N=number, OR=observed range, SD=standard de-
viation, CV=coefficient of variation. Lengths in mm.

were observed with ‘Mag-Lite’ (Mag Instru-
ments, Ontario, Calif) torches. The expanding
beam of these torches greatly facilitated detection
of rats moving among mangrove roots. A total of
57 hours was spent observing wild X. myoides
(Table 3). Rats were more difficult to locate inside
densely vegetated communities (e.g., Rainbow
Channel), but in areas of low vegetation complex-
ity (e.g., Canalpin Creek, Chiggil Chiggil) the
white bellies of running rats were conspicuous.
Animals were generally silent and in torchlight
the eyes reflected poorly (dull mauve).The diet
was assessed by directly observing (with binocu-
lars) radio-tagged rats while they fed, and by the
analysis of middens attributed to the rats. Rats
were interrupted soon after they began to eat soft
prey and the food remnants were pirated for iden-
tification. Crustacean prey were identified from
skeletal fragments collected after the meal. Fae-
cal pellets were sampled from many animals, but
apart from parasites, their contents consisted
mostly of minute fragments of unidentifiable
crustacean shell and integument.

Non-intrusive observation inside nesting-
mound tunnels, nest chambers and of nestling
young was facilitated by the use of an endoscope
(Olympus GIF type P2) and cold light supply
(Olympus CLE-3) powered by a portable Honda
240V generator. Extensive convolution of tunnels
in nest mounds made it impossible to reach nest
chambers if the endoscope was inserted through
a natural entrance. However, it was possible to
reach nest chambers using a 15mm-wide hollow
coring rod (fibreglass fishing rod) pushed and
twisted gently into the roof of the mound. When
a lack of resistance in the passage of the rod was
detected, the coring rod was extracted, cleaned
out, reinserted, and the flexible endoscope tube
then introduced and an examination of the area
made.

STUDY SITE AND HABITAT. At each site sur-
veyed for X. myoides, all mangrove species were
noted and the vegetation was classified visually
according to density and height into one of the
community types of Dowling (1986) described
below.

North Stradbroke Island, which forms the
major part of the eastern boundary of Moreton
Bay, lies 40km east of Brisbane. The island is
triangular in shape, 36km long from north to

346

south and about 285km" in area (Fig. 2). Physi-
ography, gealogy, soils, climate, hydrology and

lant communities of this humid subtropical is-
and are broadly outlined in Stevens & Monroe
(1975), Clifford & Specht (1979), Covacevich &
Durbidge (1981) and Coleman ct al. (1984).
North Stradbroke Island's western mangrove
communities facing the shelter of Mureton Bay
are characterised by three ubiquitous and domi-
nanl plant species (Avicennia marina,
Rhizophora stylosa and Aruguiera gymnorrhiza)
of wide ecological amplitude, Le., tolerant of a
wide range of Salinity and temperature regimes
(Bunt etal.. 1982). The drainage basin supporting
the mangrove communities is jn a relatively low-
rainfall range (Dunwich annual precipitation
L6O2 mim) but with an intermediate to low run-off
coefficient (ratio of run-off to rainfall, 0.2-0.5)
which, given the island's extensive swamps,
overflow basins, dense vegetation and sandy sub-
strate, allows considerable retention of water with
more regulated and sustained release to the drain-
age systems (Hutchings & Saenger, 1987). How-
ever, in backing onto Melaleuca quinquenervia
swamps and other poorly drained areas, as com-
monly is the case, the full development of many
mangroves zones is impeded in comparison to
those zones abutting better drained and drier
areas (Dowling, 1979).

The mangrove communities most commonly
associated with this study (Table 1) comprised
Avicennia marina var. australasica, Rhizaphora
stylosa and Bruguiera gymnorrhiza, Two other
species, Aegiceray corniculatum and Ceriops
tagal var. australis were more commonly repre-
sented toward the southern end of the island.
Species descriptions and community composi-
tions for this group in Moreton Bay can be found
in Dowling (1986), from which only broad char-
acteristics relevant to this study are summarised
helow.

Avicennia marina var. australasice, the most
common and widespread species found in
Moreton Bay, occurs in any pari of the intertidal
zone, either as å pure stand or mixed with other
species, hut usually as an emergenti, Ic is also the
most variable species in Moreton Bay, being rep-
resented as large. well-developed trees from 10-
120 (or mare) tall, or as small spreading shrubs
only 0.5m tall when fully developed. Its growth
is rallest and most vigorous when this species
stands at the extreme seaward edge in well.
drained soils that are inundated by all high tides.
At the upper tidal limits its growth is less vigorous
and it usually occurs as å small shrub. In areas of

MEMOIRS OF THE QUEENSLAND MUSEUM

poor drainage, bur frequent tidal inundation, Å.
marine forms å low spindly tree. Where Ireshwa-
ter secpape occurs this species tends t0 form
stunted, round, leafy bushes. Death of Avicenna
marina occurs with waterlogging, ponding of
waler and siltation leading to the covering of
pneumatophores, The structural forms of Avi-
cennid marina encountered during this study
ranged from Low open-shrubland, Low shrub-
land and Open-heath (mapping unit 2), Tall open-
shrubland and Tall shrubland (mapping unit 3),
Low open-woodland, Low woodland and Wood-
land (mapping unit 5) and Low open-forest and
Open-forest (mapping unit 7) (Fig. 7; see Dowl-
ing, 1979, 1986 for definitions and descriptions).

Throughout Moreton Bay, Rhizophora stylosa
occurs either as solitary trees, in clumps of three
1o five, or in pure stands of densely-packed trees,
It may also occur in association with other man-
grove species as a large, single-stemmed shrub or
small tree toward the lower tidal limits or veca-
sionally in a very narrow zone behind the seaward
fringe of A. marina. Freshwater from Melaleuca
swamps is thought to stunt the development of
this species (Dowling, 1986). In the study area R.
stylosa grew in both isolated clumps and in thick
belts. Trees growing in clumps were represented
by individuals of different heights (1-4m) and
age. Belts of this species were found hehind the
seaward fringe of Avicenttia marina and extended
to the land ward edge, Here the trunks were more
erect and unbranched, the prop and still roots
tangled to 1m above the ground and the branches
did not spread laterally. Structural forms of R.
stylosa encountered during this study consisted of
Tall shrubland, Open-serub and Closed-scruh
(Fig. 3; mapping unit 18. Dowling, 1986),

Bruguiera gymnorrhiza occurs in Moreton Bay
as scattered plants or small clumps of 10-15 trees
on the most landward fringes, Tt is found on hard
consolidated muds or peaty clays and is most
common in areas of freshwater seepage from the
supralittoral zone. In the study area it did not
occur in communities, but as solitary trees up to
4m or small clumps (two to five trees).

Aegiceras corniculatum (2m tall) occurred wid-
jacent lo the reed zone on the landward side of the
Canalpin Creek and Stockyard sites. This species
grows on firm muds which have a shallow laver
of soft mud on top. It grows on poorly-drained
soils that are shallowly inundated by most high
tides, Structural forms encountered included
Low open:shrubland ånd Open-heath (mapping
unir 9),

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND 347

FIG, 8. Rhyzophora stylosa open-scrub, Rainbow Channel (Bruce Cowell).

348

Ceriops tagal var. australis
occurred only at the upper tidal
limit ofthe Stockyard site, Nor-
mally, Ceriops communities
are inundated only by a few tides
each month, and they occur on
firm, well-drained clays,
clayey muds or sandy clays.
The structural forms encoun-
tered included Tall open-shrub-
land, Tall shrubland, Open-scrub
and Closed-scrub (mapping
unit 15).

The floor of the mangrove §
community provided an array
of microhabitats including tidal
pools, channels, crab holes,
pneumatophores, crevices in
bark and around roots, hollows j
in standing and fallen timber,
suspended drifts of twigs and
leaves, driftwood, detritus and
tide-washed garbage, The
complexity of microhabitats
vared according to the struc-
ture of the mangrove commu-
nity, Highest structural floor
diversity occurred in the spe-
cies-rich communities at Rain-
bow Channel (Fig. 8) followed
by those at Stockyard. Lowest
structural floor diversity oc-
curred in the Avicennia wood-
lands of Canalpin Ck and
Chiggil Chiggil.

On the landward side, man-
groves were abutted by either a
discrete zone of sedgeland,
Melaleuca quinquenervia
swamp/Eucalyptus robusta
forest alliance, or heathland
(Fig. 9). The sedgeland band
(1.2m high) in which salinity is
reduced by drainage from the
mainland, comprised almost exclusively Juncus
kraussii, Baumea juncea, B. rubiginosa and
Fimbristilis ferruginea, In many places this zone
was up to 30m wide and was sharply delineated
from both the upper reaches of the mangrove
woodland tract and the lowest extent of the
wallum (dominated by Melaleuca quinguenervia,
Eucalyptus intermedia, E. robusta, Leptospermum
liversidgei, Gahnia sieberiana and Caustis blakei).

Supralittoral banks, formed by the combined
effects of spring tides, wind setup and storm

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. Aerial view of Rainbow Channel study site showing zonation from
marine meadow (bottom) through mangrove community and sedge zone to
freshwater swamp/wallum woodland (top) (Bruce Cowell).

surge, occurred at the sedgeland/wallum or man-
grove/freshwater marsh ecotones, The Canalpin
supralittoral bank, at 95cm high, was the tallest
bank encountered. In some areas (e.g., Deanbilla)
it was poorly defined. In most areas where this
bank aburted wallum, the groundsel Baccharis
halimifolia was a dominant species along the
bank. After periods of rain, freshwater percolated
through the supralittoral banks and the integrity
of the bank depended on its consolidation by the
roots of salt-tolerant species such as M. guin-

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

349

TABLE 3. Home Range estimates, linear distances covered and allocation of time in X. myoides. HM 957229555
isopleth of the Harmonic Mean Measure, MCP=Minimum Convex Polygon, Dist(m)-linear distance covered

during observation rime (Obs. time).

quenervia and Casuarina glauca. Where these
species were absent (e.g, Canalpin), the supra-
littoral bank experienced dramatic scouring and
profile changes.

RESULTS

DISTRIBUTION AND TRAPPABILITY. Verger
myoides was recorded from all island mangrove
communities surveyed between Chiggil Chiggil
and Stockyard (see Table I).

The species was apparently not trap-shy, A total
of 11] X. myoides individuals were trapped and
24 individuals were retrapped (Table 5). Trapping
success for new captures was higher in mangrove
transects compared to ‘barricade’ wrapping where
up to eight traps Were set in å ring around a nesting
mound in sedgeland (total traps set in mangroves
= 606. trapping success for first captures 11.7%;
total traps set around sedgeland nest = 710, suc-
cess for first captures = 3.9%). Overall success
(first captures and recaptures) from mangrove vs.
nest settings was slightly higher in mangroves
(mangrove trap success 11.4%, nest success
8.495). The total number of sub-adult males
trapped (seven) was low. If traps were left open
through the day high tide, bait was usually eaten
by toadfish (Tetractenos hamiltoni, Marilyna
pleurostricta) or traps were upturned by Mud
Crabs (Scylla serrata).

Capture of taxa other than X. mvoides was rare.
Five Melomys burtoni were caught (Capembah
Creek, I & 4 May 1992 at the mangrove/dry
woodland ecotone, two individuals; and
Deanbilla, 25 September 1992 in mangroves;
three individuals); M. cervinipes was trapped at
Deanbilla. 26 November 1992 in mangroves (1
individual); and two Ratrus lutreolus were caught

Puce ox | nel uem [ue | mr am] Ex] 197
ME 0.46 | ooe [ osb f NEN HN NN
TT
10.12.92 1063 tr Lam 924

Came | mr | s | om | owon | oss | sos | SwOmi | 673 |

ams | ra | ce | oss | emos | om | 200 | mex | os

077 024.029 0.74 127 | ?hrásmin 95.8
0.47-1.06 342 Shr 1 Imin n
us

at nest-site G of X. myoides at Rainbow Channel,
1 May 1992 and 22 August 1992 at the sedge-
land/dry woodland ecotone. Cane Toads (Hufo
marinus) were frequently seen or caught in
sedgeland traps in the Rainbow Channel site and
in mangroves of the Canalpin Creek site. One
Lewin's Rail Rallus pectoralis was trapped over-
night in sedges at nest-site B, Rainbow Channel,
Suprisingly, no Hydromys chrysogaster were
seen or trapped during the study, although the
dismembered remains of a large Mud Crab
(Scylla serrata) were noted at Myora Springs.
The presence of the Water Rat was suspected ial
the Rainbow Channel site where numerous traps
were found tripped, their baits removed and eaten
close by.

During the study only one Rattus ratius wits
observed (running along the prop roots of Ø.
stylosa trees). but numerous middens and food
caches attributed to that species were recorded jn
trees. Measurements of Xeromys myeides from
North Stradbroke Island appear in Table 2.

NESTING. Xeromys myoides nested in å wide
variety of sites, and nest forms varied consider-
ably, ranging from large termitarium-like mounds
up to 60cm high built in the marine sedgeland
(Fig. 10) to inconspicuous tunnels excavated in
the supralittoral bank at the marine-sedge-
land/freshwater-wallum ecotone Six large
mounds were recorded from within the Rainbow
Channel site and one from Stockyard. Adjacent
nest mounds were separated by distances ol be-
tween 80 and 410m. Mounds were constructed of
a mixture of excavated peat, sedge straw, mud and
sand. They were from 20cm to 60cm high with
basal circumferences from 1.6m to 4.8m, From
one 10 Three entrance holes were found al each

350

TABLE 4. Residency in nest mounds presumed to be
used by more than one animal. Individuals trapped at
nest. * Individuals re-trapped in mangroves directly

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLES. Recaptures of Xeromys myoides (by sex, age
and individual number) at Rainbow Channel.

in front of nest. # Transmitter day-fix. Age Indiv. | p, | Weight FOR | Indiv. Date | Weight
No. | (g) | No. (g)
i Female SA 19 |17.792| 28
e [Mom] age | sex || Rene | ys ama GE
H 1 A M | 26.292 i 2592 | 43 | 22.8.92| 28
27 A M 21.8.92* 18.6.92 | 46 | J 11 2.5.92 27
26 A F E nm a | 4 [2592| 54 | 17692] 26
28 SA F - 18692| 41 |Male
Er SA F - A | 20 [17792] 36 | a | 16 [16792] 43
3 I M 7 17.792| 36 | 17.792| 43
18,19,27,28.7; 18.7.92| 36 18.792] 43
å I" P 10.12.92* | 19.792| 36 | 19792| 44
3 A F 2.5:17.6.92 A | 21 [17792] 40 | 44
34 | SA M 22.8:10.9.92 17.7.92| 40 | 28.7.92 | 44
7 SA | M 18.792 z21892| 40 | a | 27 | 6892
17 SA F 27.1.93* A | 26 [6892] 37 | 21.892| 41
12 SA F | 2592 |3.5:16.7;17.6.92 6.8.92 | 37 | A | 33 |20892| 48
25 SA F |28792 - A | 31 20892 | 39 | 228.92| 48
1 J F | 2592 | 3.517.692 22892| 39 | A | 36 |21892| 43
B 2 A F 20.11.91 - A | S6 |19.9.92 22.892| 43
ai A F 2.5.92 17.6.92 209.95] 40 | A | 38 |21892| 41
18 SA M | 17792 - A | S9 |21.9.92| 47 21.8.92| 41
19 SA F |17792 | 27.7;228.92 21.1092| 42 | A | 82 |27.1192| 50
D 24 A M | 27.7.92 - A | 66 |22892| 37 28.11.92| 50
20 A F 117792 | 18.7;19.7.92 22892| 37 | A | 93 |24593| 53
SA | 12 25.5.93
mound, but others may have been hidden under
the labyrinth of exposed tree roots into which
mounds were often incorporated. The number of
entrance holes at any one nest varied throughout 21.8.92
the year, with disused holes quickly filling in. 22.8.92
Regularly used holes were oval in shape (approx- SA | 17 |17792
imately 60mm by 40mm) and positioned any- 18.792| 31
where from the top to the bottom of mounds; 19.792| 31 J
holes were from 30cm to 2m apart. Although 27.7.92| 23 |

lower holes were flooded at high tide, the mounds
themselves were never seen to have been entirely
inundated (Fig 11).

Evidence of external mound ‘workings’ con-
sisted of fresh roof ‘plasterings’ in which a thin
black paste of mud mixed with peat, dried leaves,
sedge stems, or crab shells had been daubed over
the top of the mound and worked between sedges
(usually Juncus kraussii). Areas ‘plastered’ in this
way measured as much as 0.12m* and in many
cases a semi-tubular path of fresh plastering cov-
ered a track from the nest entrance hole up or
around to the nest top. The absence of fresh
plastering did not indicate that the nest was unoc-
cupied, as plastering was an infrequent event.
Nest-site G was first noticed on 15 Nov 1991 as

a reed-covered hole in the supralittoral bank ac-
companied by traces of plaster in the reed bases.
By March 1993, plastering had built the mound
up by 6cm and the entrance hole had been moved
to the top.

Three mounds were built on slightly elevated
islands supporting small patches of stunted
Melaleuca quinquenervia and Casuarina glauca.
One of these nests incorporated the stem of a
living Vitex trifolia var. bicolor (Figs 10, 11).

Tunnels in the supralittoral bank (shaped and
maintained by spring tides) were inconspicuous
and generally only found by radio-tracking indi-
viduals trapped and released in the mangroves.

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND 351

FIG. 11. Nesting mound (B) moated by high tide, Rainbow Channel.

las
U^
IA

FIG. 12. Xeromys myoides despatching a Red-fingered Crab Parasesarma
eryihradactyla, the most commonly eaten crustacean. (Bruce Cowell).

Four were recorded in the Rainbow Channel site
and confirmed by radiotelemetry and two were
recorded and confirmed from Canalpin Ck, The
face of the bank, and hence the location of en-
trance holes, changed dramatically at Canalpin
Ck with erosion of the bank caused by spring tides
and freshwater runoff,

Several indications from radio-tagged individ-
uals suggested that Xeromys myoides built, or at
least used, extensive and complex burrow net-
works inside and above the supralittoral bank.
One male (AM33) covered approximately 20m
underground after returning to its nest when a
high tide restricted foraging in the mangrove
zone, One female was recorded using a system of
burrows through Gahnia-dominated freshwater
swamp 10m behind the supralittoral bank at Can-
alpin Ck. The peat substrate of both these areas
may facilitate a significant subterranean facet of
life history as yet unsuspected in this species.
Alternatively, Xeromys may have been using tun-
nels constructed by Rattus lutreolus, an ex-
tremely common species in these areas.

Although large termitarium-like nesting
mounds are known to be occupied and main-
tained by groups of rats, the rodents’ proclivity
for unpretentious nesting in supralittoral banks
suggests thal some large nesting mounds may
represent physical, historic evidence of the past
position of the supralittoral shoreline. The rat's
demonsirated ability to consolidate and add to

MEMOIRS OF THE QUEENSLAND MUSEUM

a mound, as well as the likeli-
we hood of a mound overgrowing
4 with sedges and small trees,
both increase the chances of a
nest-site remaining when the
rest of the bank erodes through
į spring tides and heavy wave
action. The maintenance and
extension of a nesting mound
E may represent a response by
the occupants to a wet nesting
chamber during high tides,
Over the course of the study,
mound nests were found to
contain animals of all three age
categories (adult, sub-adult and
juvenile) and of both sexes
(Table 4). However, no more
than one adult male was re-
corded concurrently in the
same mound (other sexes and
age categories were recorded).
Adult males were observed vis-
iting neighbouring nests during
the night, so a capture at a nest site did not
necessarily imply occupancy. Adult females were
noted to avoid entering a co-mounding female's
burrow and if released into a 'wrong' tunnel
would quickly turn around, exit the burrow and
run across the mound to their ‘own’ tunnel. En-
doscopic examination of one nesting mound (A)
revealed extensive tunnel systems through the
mound, with two nesting chambers within the top
16cm. Each nest was lined with the dried leaves of
Melaleuca quinguenervia, which also littered
areas around the mound. Mounds and bank tun-
nels provided humid protection not only during
the day, but during night high tides when most X.
myoides returned to their nests (presumably to
avoid the water). One instance was noted of a

FIG. 13. Unnamed marine pol yclad (30mm long), fea-
turing in the diet of Xeromys. (Bruce Cowell),

XEROMYS MYOIDES OF NORTH STRADHROKE ISLAND 35

FIG. [4. Marine pulmonate Ophicardelus quoyi showing breaches made by

Xeromys (Bruce Cowell),

male remaining in a hollow of a mangrove trunk
during å night high tide.

DIET. The following are food items recorded by
direct observation of wild X. myoides.
Crustacea. The small crab Parasesarma
erythrodactyla was the most commonly recorded
prey item taken inside the mangrove zone.
Xeromys myoides ate both adults and minute ju-
veniles (see sizes documented below). The man-
ner of dispatching à crab was as described by
Redhead & McKean (1975); crabs were first dis-
armed of claws, legs were bitten off, and the body
turned over and breached from the softer ventral
side (Fig. 12). The meat was scoured from the
inside of the carapace. In some instances rats
were observed to bite adult crabs between thc
eyes then drop and leave them, but return after a
few minutes to consume the catch under suitable
cover. Carapaces and basal leg segments were left
uneaten, but claws were sometimes hollowed out.
Parasesarma erythrodactyla was observed being
consumed by four X. myoides (AF4, AF31,
AM34, AM89) in the months of March, May and
August.

Helice leachi (one specimen, carapace width
20mm) and Australoplax tridentata (one speci-
men, width 15mm) were recorded in the diet of
AMB9 at Canalpin Ck on 20 March 1993.
Marine polyclads. An undescribed marine poly-
vlad (Fig. 13) was recorded in the diet of adult
female (AF4) at Rainbow Channel, 3 May 1992.
Two instances of polyclad predation were re-
corded on the same night. Under studio condi-
tions, another adult female was photographed
feeding on the same species. Polyclads were reg-
ularly recorded sheltering in rotting driftwood
washed to the dry supralittoral zone, As many as

pe

14 were once recorded from in-
side the honeycombed core of
a small log (45cm long x 10cm
diameter). There have been no
ütherrecorded predators of ma-
rine polyclads (L. Cannon pers.
comm. ),

Marine pulmonates, Op-
hicardelus quoyi (Fig. 14) was
recorded being eaten by an
adult female (AF4) at Rainbow
Channel on 11 May 1992 (and
were readily eaten by animals
held in captivity), These small
pulmonates gathered under
driftwood in the middle of the
sedgeland, Female AF4 was
heard cracking O. quoyt under
a large plank, On examination, a midden of
opened shells was found in a concavity clipped
from reeds flattened under the plank, All shells
had been breached through the body of the shell,
and not through the aperture.

Salinator solida (Fig. 15), a small (8mm) pen-
winkle-like mollusc was recorded from the diet
of an adult male (AM89) at Canalpin Ck on 19-20
March 1993, and was present in numbers in à
Chiggil Chiggil midden. Entry to the body of the
pulmonate appeared to be through the aperture as
no retrieved specimens were breached,

Onchidina australis, a large (30mm), shell-less

slug was eaten by an adult male (AM89) at Can-
alpin Ck on 20 March 1993, Parts of the gut were
left uneaten,
Marine bivalves, Glauconome sp. (Fig. 15), a
small (11mm) sea-green mussel was eaten by an
adult male (AM89) at Canalpin Ck on 20 March
1993. Opened halves of this bivalve were com-
monly observed across the mangrove floor, but
none was recorded in Xeromys middens,

The following Crustacea are food items re-
corded from middens which were attributed to X
myoides:

Parasesarma erythrodactyla was the most
common component of mangrove middens. One
midden inside the base of a hollow Avicennia al
Chiggil Chiggil contained 14 carapaces ol this
crab (average width 12mm, range 10-16mm) and
fourof Helice leachi (average width 17mm, range
i5-20mm), Largest carapaces of P. erythro-
dactyla believed to have been consumed by X.
myoides measured 20mm in width. Helice leachi
was the most commonly recorded component of
supralittoral middens, A sample ol carapaces
taken from à Chiggil Chiggil reed zone midden

354

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 15. Prey taken by Xeromys on North Stradbroke Island, (anti-clockwise from crab) Red-fingered Crab
Parasesarma erythrodactyla, Sea-green Mussels Glauconome sp., marine pulmonates Ophicardelus quoyi and
Salinator solida, unnamed marine polyclad (Bruce Cowell).

known to be used by an adult female (AF55)
contained 36 carapaces (average width 17mm,
range 13-21mm) and three of Parasesarma
erythrodactyla (average width 13mm, range 11-
14mm). Peragrapsis laevis (four specimens, av-
erage width 17mm, range 13-23mm) was
recorded from a hollow Avicennia marina ground
level spout known to be frequented by an adult
female (AF31) at Rainbow Channel. Skeletal re-
mains of the mud lobster Laomedia healyi were
collected on 20 Aug. 1992 and 16 Sept. 1992 from
a hollow Avicennia marina ground level spout
known to be frequented by an adult female (AF31).
Diet in Captivity. Captive specimens of X.
myoides consumed relatively large quantities of
food. During the period 20 Feb. 1995 — 4 March
1996 the average weight of raw prawns con-
sumed daily by an adult female (of average
weight during that period 38g) was 22g represent-
ing an average daily intake of 57.8% body weight
(at ambient temperature 24°C). For a period of
eight weeks (18 Sept. 1995 — 13 Nov. 1995) this

animal maintained body weight (average weight
38.0g, 35-40g range) on a raw prawn diet with no
access to free water.

FORAGING BEHAVIOUR. Observed radio-
tagged individuals of Xeromys myoides spent
their foraging hours dashing over the substrate
between trees, ‘snaking’ through pneu-
matophores and investigating hollows in man-
grove trunks. Their nocturnal behaviour was
generally frenetic although animals were re-
corded to rest inside a favoured mangrove hollow
for periods of up to three hours. Linear distances
covered and times spent foraging are shown in
Table 3.

Generally rats left their nests and followed the
receding tide out through the sedgeland and into
the mangroves where they foraged. In so doing
they passed through a ‘common’ corridor be-
tween the nest and the foraging area. In some
cases this precipitated meetings (usually brief and
violent) with other rats or an inspection of

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

355

FIG. 16. Typical resting hollow in an Avicennia marina base, Rainbow Channel (Bruce Cowell).

neighbouring nests. Animals scoured their home
range during the course of an evening.

Animals visited and revisited favoured haunts
such as hollows in Avicennia trunks, Rhizophora
root tangles and hollow fallen logs. Large food
items such as adult crabs were taken to favoured
hollows (Fig. 16) where the item was consumed
under cover. With time, discarded food items
formed recognisable middens (Figs 17, 18). All
nocturnal resting took place in such hollows.
Animals climbed up inside gently sloping trunk
hollows to about 1 m, but no arboreal activity was
ever recorded. The favoured 'pathways' of X.
myoides were those that ran closely alongside the
low, long prop roots of Rhizophora stylosa, under
the canopy of twigs and leaves deposited on the
tops of prop roots by the fallen tide, or alongside
fallen logs. In open areas of Avicennia forest
animals ran quickly between trunks and logs in
order to regain cover.

HOME RANGE DETERMINATION, Home
range estimates were made for eight animals
(four adult females, three adult males, one sub-
adult male) from the Rainbow Channel area and
I adult male from Canalpin Creek, Results of
home range analysis and plots of nests appear in

Figures 19, 20, 21 and Table 3. There was a high
degree of correlation between HM ànd MCP es-
timates. The 9596 isopleth of the harmonic mean
distance minimum (HM) gave an average home
range area of 0.637ha for the seven adults radio-
tracked at Rainbow Channel. The minimum con-
vex polygon (MCP) average for the same seven
animals was 0.614ha. The results suggested a
greater home range area for males than females
(male HM average 0.78 ha, MCP 0.77ha vs fe-
male HM average 0.59ha, MCP 0.53ha), but this
may have been influenced by the smaller number
of fixes available for females. These results
should be taken as preliminary and minimal as
some plots indicate that the curves had not quite
asymptoted (see Quin, et al., 1992; Goldingay,
1992). The home range of the Rainbow Channel
sub-adult male (SAM34) was smaller than all
other males and less than the female average. The
adult male (AM89) from Canalpin Creek had a
home range estimated at between 7.35ha (HM
95%) and 3.42ha (MCP). Given the physio-
graphic barriers associated with the Canalpin
Creek site (salt water, dry forest and a very nar-
row band of mangroves, Fig. 21), the MCP esti-
mate represents the most acceptable estimate of
home range size. Core areas (averaged from the

LA
WA
c

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 18. Xeromys myoides midden from Rainbow Channel showing mainly crab shell fragments.

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND 357
=> — —
i E 3 8 se^ S
LU d RD B) x E Kov ge^
g^ E S Lak Wie m^ a
MARINE el FT
MEADOW < ES
vs `
Doro d MANGROVE

WALLUM

FIG. 19. Home range estimates of eight Xeromys myoides individuals from Rainbow Channel (Site 2). For home
ranges, solid black lines show 9546 isopleths of Harmonic Mean measures, hatched areas represent core areas
(65% isopleths). Sub-adult male 34 is from nest A, as is adult male 16, adult female 4 is from nest B, adult
female 69 from nest C, adult female 21 from nest D, adult male 68 from nest E, adult female 31 from nest F
and adult male 33 from nest G. All nests except B are at the supralittoral ecotone between marine sedgeland

and wallum woodland. The arrow points north.

HM 60-65% isopleths) for Rainbow Channel
adults averaged around 0.20ha, with adult male
core area being estimated at around 0.19ha and
adult females at approx. 0.20ha. The Canalpin
Creek male had a core area estimated at around
0.76ha. Core areas of the eight Rainbow Channel
animals did not overlap, but animals moved in
and out of their neighbouring territorial peripher-
ies.

REPRODUCTION. Scrotal males were recorded
in the population in all months except January,
February and April (few traps were set during
February, none in April, and one young adult male
was trapped in January). Females, presumed to be
pregnant by their swollen appearance and in-
creased weights, were recorded in the months of
January, May, August, September and October.
Females with enlarged nipples were recorded
from July through to December. Juveniles were

recorded in the months of May, July, August and
November, Little is known of litter size and noth-
ing of the rate of growth of young X. myoides on
North Stradbroke Island. Endoscopic examina-
tion ofa nesting chamber in nest site 2 on the night
of 26 October 1993 revealed a litter of four dark-
skinned but hairless young. On detection of the
tip of the endoscope the adult removed each pup
separately to another location in the nesting
mound.

In the absence of more information it would
appear that X. myoides is capable of breeding
throughout the year.

PARASITES. Nematodes were retrieved from
the faeces of one adult female X. myoides (AF2)
and from the stomachs of three subadults (OM
JM9478, JM9479, JM9728) and one adult male
(QM JM9480). These were assigned to the genus
Tikusnema Hasegawa, Shiraishi & Rumpus 1992

358

MEMOIRS OF THE QUEENSLAND MUSEUM

7 A Å å
Kä

MARINE ÁS
MEADOW (7^ ^—

MANGROVE

WALLUM

|

FIG. 20. Home range estimates (Minimum Convex Polygons) and actual trackways recorded for the eight
Xeromys myoides individuals of Fig. 19, from Rainbow Channel (Site 2). The arrow points north.

as T. vandycki Smales, 1995. The implications of
the presence of acuariid nematodes, normally
found in birds, in X. myoides and other small
mammalian hosts is discussed by Smales (1995).

Mites, Mesolaelaps australiensis (Hirst, 1926),
were retrieved from the body and ears of AF3,
AF17, AM41 and AM15. This species has been
recorded from a wide range of mammals, birds
and reptiles in Australia and the South West Pa-
cific (R. Domrow pers. comm.)

Ticks Ixodes tasmani were removed from the
body of AM41 and from between the toes of the
hind foot of AM7.

PREDATORS AND COMPETITORS. Carpet
Pythons Python spilotes were recorded in sedge-
land (Nov 1992) and inside mangrove forest (Nov
1992). A2m specimen draped across Rhizophora
prop roots approximately 10cm from the floor of
the forest at low tide was probably hunting rats.
Two Rough-scaled Snakes Tropidechis carinatus
were recorded in the sedgelands at low tide (25
Feb. 1992, 10 Nov. 1992); one was investigating
nest site 2 but was not observed entering any

burrows. Tawny Frogmouths Podargus
strigoides commonly occurred in woodland adja-
cent to the mangrove zone but no owls were
observed during the study. One frogmouth was
observed at Canalpin Creek watching over man-
grove flats. A three-syllable Ninox-like call was
recorded in the Myora mangroves on 26 Oct.
1993 but the identity of the caller was not deter-
mined. Dissection of Cane Toads Bufo marinus,
found feeding in the mangroves of Canalpin Ck
revealed a diet consisting largely of crabs, marine
pulmonates and amphipods. The crabs were of a
similar size to those eaten by X. myoides.

It is possible that Xeromys may represent a
potential prey item for Hydromys. Woollard et al.
(1978) recorded young Hydromys in the diet of
adults of that species (particularly males). If this
behaviour occurs also in Xeromys it may provide
one possible explanation (along with early expul-
sion or killing without ingestion) for the low
record of sub-adult (dispersing) males captured
during the survey (seven sub-adult males: 23
sub-adult females).

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

DISCUSSION

Xeromys myoides on North Stradbroke Island
is a small, spotted, specialised, non-aquatic, non-
arboreal, nocturnal muroid rodent whose life his-
tory revolves around the island's western
mangrove and adjacent sedgeland communities.
The species appears to be carnivorous.

DIET. Truly carnivorous species of rodents which
ingest more than 5096-8046 animal food are very
rare (Dieterlen & Statzner, 1981). African exam-
ples include the dendromurine rodents Deomys
furrugineus and Leimacomys buttneri (Dieterlen,
1976); murine rodents of the genus Lophuromys,
Zelotomys (Dieterlen, 1976) and Colomys
goslingi (Dieterlen & Statzner, 1981). Neotropi-
cal examples include hesperomyine cricetids
such as Ichthyomys, Anatomys, Daptomys, Neu-
stictomys, Rheomys (Starrett & Fisler, 1970;
Voss, 1988) and Nectomys squamipes (Ernest &
Mares, 1986). Examples from Australia and New
Guinea include the hydromyines Hydromys
(Woollard et al., 1978, Menzies & Dennis, 1979),
Crossomys monktoni, Parahydromys asper
(Menzies & Dennis, 1979), Leptomys elegans
(Dwyer, 1984), Mayermys ellermani,
Neohydromys fuscus, Pseudohydromys murina
and P. occidentalis (Jackson & Woolley, 1993)
and probably also the genera Paraleptomys and
Microhydromys (Menzies & Dennis, 1979;
Flannery, 1990).

Carnivore digestive systems are noted for their
simple stomachs, short small intestines, short
large intestines and short or absent caeca (Hume,
1982). Barry (1977) commented that for most
rodents, relative gut lengths are inaccurate indi-
cators of relative absorptive surface area, given
that a large proportion of the total absorptive
surface of the hindgut may be found in the small
intestine although this might account for a rela-
tively small proportion of the total length. How-
ever, Barry also noted that for more carnivorous
forms, relative lengths more accurately reflect
relative absorptive surface areas because the cae-
cum and colon are greatly reduced in relation to
the small intestine. Gut morphology suggests that
the diet of Xeromys myoides should not be totally
carnivorous. Relative alimentary canal dimen-
sions for five New Guinean hydromyines are
presented by Jackson & Woolley (1993).The rel-
ative length of the small intestine of Xeromys
myoides (8596 of the total gut) is slightly shorter
than that recorded for the four New Guinean
hydromyines Mayermys ellermani,
Neohydromys fuscus, Pseudohydromys murinus

359

and P. occidentalis (Jackson & Woolley, 1993),
while the relative length of the large intestine
(14%) is longer. However, relative caecum length
in X. myoides (296) is shorter than in Mayermys
ellermani, Neohydromys | fuscus and
Pseudohydromys occidentalis and approaches
the relatively short caecum length found in P.
murinus (196). Jackson & Woolley (1993) sug-
gested that, on the basis of gut morphology and
dietary analysis, Pseudohydromys murinus
should not be considered an obligate carnivore.
However, they noted that the plant material pres-
ent in the guts of 22 of 24 sampled specimens
could have been contained in the guts of their
insect prey.

Relative lengths of the large intestine and cae-
cum in Xeromys myoides closely approximate
those of Hydromys chrysogaster (pers. obs.)
which, although recorded eating a wide variety of
vertebrate and invertebrate prey (Brazenor, 1936;
Troughton, 1941; Barrett, 1955; McNally, 1960;
Fleay, 1964; Woollard et al.,1978; Harris, 1978)
has rarely been reported consuming vegetation.
Brazenor (1936: 66) noted them to eat 'coarse
grass or rushes' and Barrett (1955: 119) com-
mented that they were 'partly vegetarian'.
Woollard et al. (1978) concluded that plants were
consumed as trace items but increased in import-
ance in winter when resources were low. Harris
(1978) recorded crustaceans as major prey items
of Hydromys chrysogaster at three sites in south-
eastern Queensland but did not observe animals
feeding on vegetation, nor detect vegetation in
scats. At two of these sites mammalian prey ac-
counted for up to 9% of prey and 596 of the
volume of scats.

Plant foods have, to date, not featured promi-
nently in the diet of Xeromys myoides. The com-
ponents of scats (from five individuals) and entire
gut contents (three individuals) have reflected
Observed feeding preferences (recorded from 9
wild individuals) and included gross items such
as mollusc viscera, shell and operculae, crusta-
cean shell and gills, sand and hair (X. myoides).
Microscopic examination of gut contents has re-
vealed plant material in the form of xylum ele-
ments, parenchyma and fibre, probably from
mangrove propagules ingested by (probably)
crustacean prey (J. Bertram pers. comm. ).

HOME RANGE. Individuals of Xeromys tracked
throughout the year in this study at Rainbow
Channel had home ranges (95% isopleths) which
abutted or overlapped outlying areas of those of
their neighbours (Figs 19, 20). However core

360

areas (65% isopleths) did not overlap. Predict-
ably, adult males had larger home ranges (average
MCP 0.77ha) than females (0.53ha) and it was
assumed that this reflected an investment of fe-
male defence and male repulsion that took the
territorial limits beyond the male's dietary re-
quirements. Females had smaller home ranges
which, presumably, provided them with quick
access to resources as close to their nests (and
young) as possible. Xeromys myoides stands to
gain much from the microhabitat specialisation
of the mangrove crab population. At low tide the
rats can exploit common species such as
Parasesarma erythrodactyla out toward the sea-
ward limit of the mangroves. During high tides,
when access is blocked to the mangrove's most
productive zones, those crabs which extend into
the high intertidal or terrestrial zones (e.g., Helice
leachi) can be exploited. Given that large body
size does not increase an animals capacity to
acquire food, a small size of around 40g (as is the
case of X. myoides) could be interpreted as the
evolutionary optimum for harvesting the com-
monest of mangrove crabs without resorting to a
semiaquatic life style such as that seen in the
much larger Hydromys chrysogaster (Water
Rat).

X. myoides appeared to respond more quickly
to scent than any other stimulus offered by its
prey, and it was suspected that given the daily
flushing of home ranges, scent laid down inside
erect hollow trunks of living trees might represent
a cost effective method of home range
familiarisation and territorial defence.

Schoener (1969) predicted that declines of prey
abundance might be offset by expansions in either
prey types or home ranges. In Hydromys, home
range size has been shown to be smaller in areas
of high substrate complexity, notably a high inci-
dence throughout the home range of aquatic veg-
etation, stumps, sunken logs, eroded
embankments and exposed roots (Harris,
1978).The same appears to be true for Xeromys
on North Stradbroke Is. At Rainbow Channel the
mangrove community is wide (200m from the
marine meadow to the sedgeland) and diverse,
with dense forests of Rhizophora stylosa and
Avicennia marina. The substrate is complex with
abundant trunk and root hollows, labyrinths of
prop roots, flotsam rafts of twigs, propagules,
leaves and eel-grass suspended on prop roots, a
profusion of fallen logs and timber flotsam, and
an abundance of tidal pools. Average home range
size for adult males (N23) from Rainbow Chan-
nel was 0.78ha (HM), 0.75ha (MCP). At Canalpin

MEMOIRS OF THE QUEENSLAND MUSEUM

Ck (Site 5), an area of Avicennia parkland, the
mangrove zone was mostly narrow (maximum
20m), with widely spaced (average 5.2m) mature
trees and very low substrate complexity. The
home range of the only adult male (A89) tracked
at Canalpin Ck was larger at 7.35ha (HM), 3.42ha
(MCP). From its nest in the supralittoral bank,
this male emerged to follow the line of the supra-
littoral bank for 300m to an area of greater
microhabitat diversity and broader mangrove
width where it foraged until returning to its nest
at dawn (Fig. 21).

Structural diversity of the mangrove commu-
nity and substrate microhabitat complexity prob-
ably determine centres of reproduction and
dispersal on the island. In this respect, the com-
munities of the Rainbow Channel and Stockyard
represent the most important conservation zones
for the species on, and close to, North Stradbroke.

Harris (1978) recorded lower weights in
Hydromys from freshwater compared to estuarine
habitats, and suggested that this reflected a dif-
ference in the quality of the two habitats.
Although False Water-rats have been recorded in
both habitat types throughout its east Australian
range, no significant foraging took place inside
the freshwater marshes and paperbark swamps
immediately accessible to all radio-tagged indi-
viduals from Rainbow Channel or Canalpin
Creek. Similarly, the outer (marine meadow) ex-
tremity of the mangrove zone, dominated by
more widely separated, more mature Avicennia,
was avoided. This study suggested that at these
two localities, the most productive zone lay
somewhere between Xeromys nests at the supra-
littoral bank and the first 100 metres into the
mangroves.

AQUATIC ADAPTATION. There has been much
speculation on the level of aquatic adaptation
achieved in Xeromys. Thomas (1889) treated it as
a terrestrial rat, Troughton (1954: 273) regarded
it as ‘an aquatic-feeding land rat’, Redhead &
McKean (1975: 352) contended that it was
‘clearly ... aquatic’, and Magnusson et al. (1976:
157) concluded that ‘X. myoides is basically a
terrestrial rat that feeds on hard-shelled aquatic
fauna".

Results of this study indicate that X. myoides
(on subtropical North Stradbroke Island at least)
cannot be described as aquatic. Indeed, on a broad
scale, the species scarcely qualifies as semi-
aquatic. Voss (1988), comparing a range of small
(14-202g) semiaquatic-carnivorous lipotyphlans
and Old World muroid rodents against closely

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

MARINE MEADOW

— ec TS em

¢
MANGROVE `

f LZ
T A
b
*

FIG. 21. Home range estimates (Harmonic Mean Measure — 95% isopleth [solid-line ‘ellipse’ ] and 65% isopleth core
area [broken-line 'circle'], Minimum Convex Polygon and actual trackway) recorded for adult male 89, Canalpin
Creek (Site 5). Fine dotting denotes extensive area of shallow, brackish, stagnant lagoon. N denotes nest of AM89

in supralittoral bank, The arrow points north.

related terrestrial taxa, found semiaquatic species
to converge morphologically in respect of the
following suite of external and cranial attributes:
l, stiffer, more abundantly distributed mystacial
vibrissae supported by thick mystacial pads; 2,
reduced, incomplete or absent philtrum; 3, re-
duced and more densely furred pinnae; 4,
hindfeet fringed with stiff, silvery hairs in plantar
margins; 5, digits of hindfoot wholly or partly
webbed; 6, round tails (in cross section) with
longer, more dense ventral hairing; 7, metatarsal
configuration approaching IV>III>I1=V>I; 8,
large infraorbital foramen; and 9, large foramen
magnum.

In Xeromys myoides the mystacial vibrissae are
no more abundant and only marginally stiffer
than in terrestrial species, but the mystacial pads
are conspicuously thick, imparting a breadth to
the muzzle belied (in the extracted skull) by a
rostrum of average width. The philtrum is absent,

but the pinnae are not greàtly reduced (average
length 12.5mm), being clearly visible above the
head, nor are they thickly furred. The hindfeet
have no fringing hairs and they lack webbing, the
tail is thinly and evenly furred dorsally and ven-
trally, and the metatarsal configuration
(II>IV>II>V>I) is that of terrestrial muroids
(terrestrial soricids and Microgale). The foramen
magnum in Xeromiys is not especially enlarged
(there is considerable cranial reduction in the
vicinity of the cerebellum), and the same applies
lo the infraorbital foramen. However, the molar
row is reduced to two per quarter, the eyes are
small, the posture couchant, the fur water-repel-
lent, and the upper incisors, as in ichthyomyines,
approach an orthodont condition.

The modest degree of aquatic specialisation
exhibited by X. myoides is remarkably conver-
gent on that of the 54-61g African Brook Rat
Colomys goslingi, which consumes. mainly tri-

LA
E
n2

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 22. Wild adult Xeromys shown in typical ‘puddling’ posture among pneumatophores. (Bruce Cowell).

chopteran larvae and terrestrial invertebrates in
and adjacent to rainforest streams (Dieterlen,
1983). The reluctance of Colomys to enter deep
water or to scale trees (Dieterlen, 1983) is also
reminiscent of Xeromys. Other aspects of behavi-
ourand ecology observed in the North Stradbroke
study are consistent with the contention that
Xeromys is an aquatic ‘puddler’ (Fig. 22) and not
adapted to a truly semiaquatic lifestyle.

It is difficult to imagine what additional semi-
aquatic specialisation the littoral and supralittoral
habitats might demand of Xeromys. Swimming
adaptations, such as webbing of digits, fringing
of lateral pes margins, elongation of lateral-most
metatarsal proportions and development of a stiff
ventral caudal brush, are clearly superfluous in à
very small species to which deeper expanses of
water must represent more of a risk of predation
than an untapped resource (see Magnusson et al.,
1976 for the record of Xeromys from a crocodile
stomach). Similarly, stiffer, more abundantly dis-
tributed mystacial vibrissae are probably unnec-
essary given the species' observed preference for
terrestrially acquired prey.

Observations of high daily food intake in cap-
tivity, rapid loss of body weight and ataxic
behaviour associated with food deprivation, and

frenetic behaviour in the wild suggest a high
metabolic level in the species. The very small size
of Xeromys suggests that it might rapidly lose
body heat through its relatively large surface area
(compared to the volume of heat-producing tis-
sue). This might be interpreted as particularly
problematical for an animal continually in con-
tact with water, especially as that small (vulnera-
ble) body size may have already imposed a
nocturnal life-style on the species.

However, the necessity to dissipate heat rather
than the need to retain it may be the problem faced
by Xeromys. Fanning & Dawson (1980) and
Dawson & Fanning (1981) discuss the suggestion
that many semiaquatic mammals have higher
metabolic rates than those of similar-sized terres-
trial mammals, their increased heat production
compensating for losses to the aquatic environ-
ment. They conclude for the hypothermia-prone
H. chrysogaster (Water Rat), that fur insulation
(not tissue) is the major component of the rat's
total insulation and that the ability of the fur to
maintain an air layer between the skin and the
water results in a significant retardation of the rate
of heat loss.

There has been no published research to date on
the insulative properties of the fur of Xeromys

XEROMYS MYOIDES OF NORTH STRADBROKE ISLAND

although its water-repellent nature has heen com-
mented on by Watts & Aslin (1981) and Redhead
& McKean (1975: 350) +... on leaving the warer,
X. myoides was dry, compared fo M. littoralis
which resembled the proverbial drowned rat.
The hair of Xeromys has been described as
‘Pseudomys-like and unlike that of H. chrysogas-
tet. år lacked ‘shields’ characteristics of other
aquatic mammals and exhibited no structure
known to impart water repellency’ (B. Triggs
pers. comm,.). Despite its close associatton with
an extremely wet, muddy habitat, and its ten-
dency to sit, half-submerged, in small puddles for
extended periods (Fig. 22), no examples of dirty,
bedraggled or ‘soggy’ Xeromys were observed
during the present study.

Dawson & Fanning's (1981) comments on the
potential problems of heat loss faced by amphib-
ious Mammals when on land may have relevance
to Xeromys if the water-repellent nature of its fur
brings with it the associated cost of extra insula-
tion. Given its avoidance of swimming, Xeromys
may deal with the problem of heat dissipation by
occupying saturated mud/peat/sand nesting
mounds during the day and by ‘puddling* while
foraging at night (Fig. 22). In this respect it is
difficult to deny an initial impression that the
sparsely-haired tail and sofl, near-naked, highly
vascular pads of the hind feet of X. myoides
represent surfaces evolved to dissipate heat, If
this conjecture is correct, the southerly extent of
distribution for this species might be determined
not so much by thermoregulatory problems asso-
ciated with decreasing ambient temperature and
body heat loss, but by the decreasing abundance
and diversity of suitable prey in less complex
temperate mangrove forests (Clough, 1982;
Hutchings & Saenger, 1987).

Xeromys myoides is a highly specialised terres-
trial muroid rodent especially adapied to the lit-
toral zone, one of the harshest of environments
available to mammals, [ts level of adapiation
should not be regarded as an early stage of
specialisation toward a more jruly semiaquatic
form. but å successful end product,

STATUS. Xeromys myoides has been described
as “one of the rarest of our native mammals
(Ride, 1970), "vulnerable to changes caused by
mam (Wans & Aslin, 1981), ‘rare, scattered
(probably at risk in some habitats and extinct in
vihers) (Redhead, 1983) and ‘vulnerable’ (Ken-
nedy, 1990: Van Dyck, 1991), More recently it
has been listed in the Rodent Action Plan with

363

nine other Australian rodents as ‘rare år insuffi-
ciently known’ (Lee, 1995).

This study has demonstrated that no special
skill is required in capturing X. myeides, and it
suggests that mangrove suryeving elsewhere for
Xeromys has probably been inadequate, How-
ever, the other possibility that North Stradbroke
Island's fecund X. myoides population might rep-
resent the makings of an island remnant popula-
tion such às that seen in Leporillus conditor,
Lagostrophus fasciatus, Setonix brachyurus or
Bettongia lesueur, should not be ignored, Given
the proximity (200m) of areas such as Stockyard
(southem North Stradbroke Island) to the com-
plex mosaic of mangrove communities associ-
ated with lower Moreton Bay and the mainland,
it seems likely that at the very least, the lower
Moreton Bay islands should support populations
of X. myoides similar to that of North Stradbroke.
The number of mainland records also suggests
that the rat ts widespread and of reasonably broad
habitat tolerance. However, until the North
Stradbroke Island population is put into a broader
mainland perspective the conservation status of
Xeromys myoides will remain in question.

ACKNOWLEDGEMENTS

This study is dedicated to Ellie Durbidge who
inspired and encouraged ihe research. I am most
indebted to Judy Conrad and Barney Hines for all
their help with the field and laboratory work
associated with this project. On a voluntary hasis
Judy tirelessly assisted with translating thousands
of flagging tape coordinates from mangrove trees
10 home range figures. Ellie and Tony Durbidge
(Nth Stradbroké Is) provided accommodation,
food, support and baby-sitting. Peter Bostock
(Queensland Herbarium) provided plant 1dentifi-
cations. John Bertram ( nsland University)
identifed injested plant material, Peter Davie and
John Stanisic (both of Queensland Museum)
identified crustacean and mollusc food scraps.
and Bob Domrow (Queensland Museum) and
David Kemp (CSIRO, Long Pocket) identified
arachnid parasites. Consolidated Rutile Limited
and Peter Foote provided access to Canalpin Ck.
Field assistance or advice was made freely avail-
able by lan Gynther (Queensland Department of
Environment), Matthew Shaw and Heather
Janerzki (Queensland Museum), Sally Towniey
(Southern Cross University), Anne-Marie Walt
(Maleny), Bruce and Carolyn Cowell (Queens-
land Museum), Barbara Triggs (Genoa), David
Titley (Titley Electronics, Ballina), Belinda Dett-

man and Lindsay Hardy (Australian Nature Con-
servation Agency). The endoscope and light
source were generously supplied by medical spe-
cialists John Quayle and John Nye {both of
Wickham Tte, Brisbane), Bruce Cowell (Queens-
land Museum) endured many sleepless nights for
the sake of photography associated with this
study, I gratefully acknowledge the help of all
those listed above. I thank Gordon Guymer (Di-
rector, Queensland Herbarium, Dept of Environ-
ment) for permission lo reproduce the maps of
Dowling (1986). Ian Gynther (Queensland Dept
of Environment), John Woinarski (Parks & Wild-
hfe Commission of the Northern Territory) and
John Seebeck (Dept of Conservation and Natural
Resources, Victoria) are especially thanked for
their critical input into drafts of this paper.

The Endangered Species Program (Australian
Nature Conservation Agency) is thanked for pro-
viding much of the funding associated with this
research.

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THE NEW HOLLAND MOUSE PSEUDOMYS NOVAEHOLLANDIAE (RODENTIA:
MURIDAE), AN ADDITION TO THE MAMMAL FAUNA OF QUEENSLAND

STEVE VAN DYCK AND BRUCE LAWRIE

Van Dyck, S, & Lawric, B. 1997 06 30: The New Holland Mouse Pseudomys
navaehollandiae (Rodentia: Muridae), an addition to the mammal fauna of Queensland.
Memoirs of the Queensland Museum 42(1): 367-3776. Brisbane. ISSN 0079-8835.

The New Holland Mouse Pseudemys novachollandiae (Waterhouse, ] 843) is reported for
the first time from Queensland where an adult male was collected at Crows Nest (near
Toowoomba) in open forest. The capture site contrasted markedly with published records of
southern P. nevaehollandiae habitat in its relatively high altitude (560m), distance from the
coast (100km), total lack of a dense shrub layer, and advanced seral stage. However,
examination of unpublished trapping records from northern New South Wales suggests that
at the northem limit of its range these environmental features may not be exceptional. [ ]
Rodentia, Muridae, Pseudomys novaehollandiae, Queensland.

Steve Van Dyck, Queensland Museum, P.O, Box 3300, South Brisbane, Queensland 4101,
Australia: Bruce Lawrie, Queensland Department of Environment, P.O. Box 731,

Toowoomba, Queensland 4350, Australia: 30 January 1997,

Thirty years ago the New Holland Mouse
Pseudomys novaehollandiae (Waterhouse, 1843)
was believed extinct or extremely rare in Aus-
tralia. At that time, given its remarkably obscure
status since description in 1843, few would have
cared, However, in 1967 it was rediscovered alive
at Port Stephens (Keith & Calaby, 1968) and at
Ku-ring-gai Chase National Park in New South
Wales (Mahoney & Marlow, 1968). Three years
låter it was recorded from Victoria (Seebeck &
Beste, 1970) and then, in 1975, discovered in
Tasmania (Hocking, 1980). Since then ithas been
recorded from widely scattered east-coastal lo-
calities between Evans Head (29°07°32"S
1537?26/31"E) in New South Wales (Posamentier
& Recher, 1974), and Friendly Beaches (42°00'S
148*17'E). Tasmania (Hocking, 1980). In New
South Wales its legal status is ‘protected’, but the
species is now the object of considerable conser-
yation concern in Victoria (Wilson, 1993, 1996),
where it is regarded as “endangered’ and listed
under the Fauna and Flora Guarantee Act 1988.

In New South Wales P novaehollandiae has
been recorded in the literature from dry sclero-
phyll forest associated with a dense shrub layer
on dune sands (Keith & Calaby, 1968; Batt et al.,
1972), from around the edges of sedged freshwa-
ter swamps between dunes (Keith & Calaby,
1968), and from coastal heaths (Posamentier &
Recher, 1974; Fox & Fox, 1978). Its occurrence
in heath regenerating after sand mining is docu-
mented by Fox & Fox (1984), In Victoria it has
been trapped in open dry sclerophyll forest
(Seebeck & Beste, 1970), woodland and low-
open forest with a heathy understorey (Kentish,

1981; Wilson, 1994, |996), low dense heath (Gil -
more, 1977; Braithwaite & Gullan, 1978; Norris
ei ål, 1983; Opie, 1983; Wilson, 1991, 1996), and
primary sand dunes with tussocks and sedges or
dune scrub with sedges and low shrubs (Quin,
1994; Menkhorst, 1995; Wilson, 1996).
Menkhorst (1995) considered optimum Victorian
habitat to be seral stages of dry heath. In Tasmania
P, novaehollandiae has been found in the wood-
land/heath mosaics of the coastal northeast, The
generally accepted importance of early succes-
sional vegetation (3-4 years) to this species is
discussed hy Wilson (1994, 1996).

Relatively recent trapping records from north-
erm New South Wales suggest that the habitat
prescription usually associated with P
novaehollandiae needs modification, Our report
here, of the discovery of the species in Queens-
land, 100km from the coast and at over 500m
elevation, is one of a number of records suggest-
ing this is not atypical for the New Holland
Mouse in the north.

THE DISCOVERY

In late August 1996, 12 small Elliott mammal
traps baited with bread and jam were sel (B.L.) in
open forest adjacent to the family home overlook-
ing the township of Crows Nest, 32km north of
Toowoomba, southeast Queensland (Fig. |), The
simple trapping exercise was part of field experi-
ence for a girls’ group staying with the family
overnight, Next moming the traps were found to
contain 3 House Mice and 2 unfamiliar mice
suspected to he Delicate Mice Pseudomys

368 MEMOIRS OF THE

152°
27°]

+ Crows Nest

9 Crows Nest
National Park

@ Big Rooster’

28°

Tenterfield EE

Grafton ES

30?

312

152° 1

QUEENSLAND MUSEUM

153° 154°

27"

28?

Ballina
A i 29°
Queensland Specimens

A + QM JM11409
e Ow! Pellet Material

NSW Specimens
€ D.Read records
* A.Martin record
A NSW NPWS records

A S.Townley records

30°

31°

53° 154°

FIG. 1. Collection sites of live Pseudomys novaehollandiae and owl pellet material from south-east Queensland,
with recent trapping records for northern New South Wales.

delicatulus. One was sent live to the Queensland tity to this specimen, it was compared in particu-
Museum and the other was released at the capture lar against type material representing P. pilligaen-
site. The animal was photographed (Fig. 2) and sis (see Fox E Briscoe, 1980). The relatively
registered in the Queensland Museum mammal broader rostrum, broader interorbital width, nar-
collection as QM JM11409. In assigning aniden- rower palate and incisive foramen, and relatively

THE NEW HOLLAND MOUSE

smaller bullae of the Crows Nest specimen col-
lectively suggested its affinities lay with P.
novaehollandiae and not with P. pilligaensis.
Analysis of hair samples (B. Triggs pers. comm.)
and features of external morphology (B. Wilson
pers. comm.) also confirmed the identity of
JM11409 as P. novaehollandiae.

Since capture of JM11409, trapping (B.L.) at
Perseverance section of Crows Nest National
Park 453 (150 trap nights), Bungaree section of
Crows Nest National Park 453 (150 trap nights),
and at, or adjacent to, the original capture site
(305 trap nights) has failed to secure more exam-
ples of this species.

DESCRIPTION OF CAPTURE SITE

JM11409 (and the released specimen) were
caught in tall open forest (27?15'09"S
152?03'22"E), Ikm north of Crows Nest, The
capture site (Figs 3 & 4) at 560-570m above
(mean) sea level (a.s.1.) abuts the town's cemetery.

TOPOGRAPHY AND SOIL. Crows Nest,
100km northwest of Brisbane, is situated on the
eastern edge of the Great Dividing Range virtu-
ally overlooking Esk in the Brisbane Valley. The
Crows Nest Shire's northern boundary is formed
by the northeast-southwest-aligned Blackbutt
Range which joins the Great Dividing Range at
the north-west boundary. Average elevation
throughout the Shire is 575m a.s.l., the lowest
point in the Shire is Maronghi Creek at around
150m, and the highest, Mt Perseverance at 807m.
Topography throughout is mainly undulating;
rolling hills to the west and steeply sloping ranges
and escarpments to the east. Geology of the area
has been mapped by Cranfield & Schwartzbock
(1973). Soils throughout are mainly hard-setting
loams to clay loams overlying yellowish-grey
clay subsoils (solodized solonetz/solodics), deep
sands (siliceous sands), reddish-brown clay
loams overlying red and brown clays (red-brown
earths and yellow earths) and shallow, stony
sands and loams (lithosols). Surface soil texture
is coarse or clay-loam, and sand is commonly
present on the surface. Coarse-grained sediments
of the Marburg Formation, Woogaroo Subgroup
and Tarong Beds form the parent material. Soils
of the general capture area are massive and cross-
bedded siliceous sandstone, some conglomerate,
minor siltstone and shale. Surface soil in the open
forest of the immediate capture area was dark
friable clay-loam without sand.

369

CLIMATE. Annual rainfall in the Shire ranges
from 650mm to 1200mm. The collection site
receives average annual rainfall of between 750
and 950mm distributed unevenly throughout the
year. The dry season months (between April and
September) receive average monthly falls of around
50mm whereas the summer months of January and
February average around 125mm. Temperature and
humidity records are similar to those for
Toowoomba (32km south) where daily mean max-
ima are around 27°C for January/February and
179C for June/July, with daily mean minima of
17°C for January/February, 6°C for June/July. On
average, temperatures exceed 32°C on ten days
of the year and frosts occur between May and
September. Average humidity is highest February
to July (68%) and lowest in September (54%).

VEGETATION. Vegetation at the capture site
was tall open-forest. Dominant species of the
upper stratum (18-19m with 60% projected fo-
liage cover) were Angophora leiocarpa (common
[C]. brown bloodwood Corymbia trachyphloia
(C), yellow stringy bark Eucalyptus acmenoides
(C), Helidon ironbark E. taurina (sub-dominant
and uncommon [U]) and grey gum E. major (C),
all to a maximum circumference at breast-height
(CBH) of 2m. A very sparse midstorey (6-7m
with 876 cover) was composed of curracabah
Acacia concurrens (C), lightwood A. implexa
(U), Queensland silver wattle A. podalyriifolia to
2m (U), rose sheoak Allocasuarina torulosa (C),
kurrajong Brachychiton populneus subsp. trilo-
bus to 3m (rare [R]) and Lophostemon suaveolens
(U). A low, open understorey (75% cover) con-
sisted of the following: Paspalidium sp. to 40cm
(C), Patersonia sericea to 50cm (C), white root
Lobelia purpurascens (C), Hardenbergia vio-
lacea (U), Laxmannia gracilis to 25cm (R), cof-
fee bush Breynia oblongifolia (R), wiry panic
Entolasia stricta to 30cm (Abundant [A ]), many
flowered matrush Lomandra multiflora (R),
matrush L. filiformis to 30cm (A), blue flax lily
Dianella caerulea (R), tall spear grass Aus-
trostipa pubescens in patches up to 5m diameter
to 1.5m high, variable sword edge Lepidosperma
laterale to 60cm (C), blady grass Imperata cyl-
indrica in patches, to 50cm, barbed wire grass
Cymbopogon refractus to 1m (when seeding)
(A), Solanum nemophilum to 50cm (A), bower
plant Pandorea jasminoides (U), monkey rope
Parsonsia straminea climbing to 8m (R),
Acrotriche aggregata to 2m (A), lady's slipper
Hybanthus monopetalus to 30cm (U), Calotis
cuneifolius (R), C. cannifolia to 20cm (R), Jack-

370 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3, Capture site of QMJM11409, Crows Nest, showing low, open understorey.

THE NEW HOLLAND MOUSE

JO
Hr

Å

sonia scoparia to 3m (C), Leucopogon muticus to
1.5m (U), Melichrus urceolatus to 30cm (U),
Goodenia rotundifolia (A), Canthium buxifolium
(R), Maytenus silvestris to 50cm (C), Pomax um-
bellata to 10cm (C), Desmodium rhytidophyllum
to 30cm (U), Grevillea sp. to 2m (R), and uniden-
tified grasses. It was thickly littered with dead
leaves, sticks, fallen trunks and branches.
Approximately 200m east of the collection site,
sandstone outcrops and boulders along an ephem-

371

eral creek became conspicuous and marked å
dramatic change in the botanical composition of
the understorey and ground layer (Fig. 5). Dom-
inant species of the upper stratum (13m with 50%
projected foliage cover) were Angophora sp. (A),
brown bloodwood Corymbia trachyphloia (U)
and yellow stringy bark Eucalyptus acmenoides
(A) to a maximum CBH of 3.35m.

The understorey changed to à dense species-
rich heath dominated by Xanthorrhoea johnsonii

372

to 2.3m (A). Around the creek, foliage cover
reached 80-90%, in other areas this reduced ta
10% with virtual monocultures of X. johnsonii.
Other heath understorey species included
Leptospermum polvgalifolium to 2.5m (C), Bank-
sia spinulosa var. collina to 1.2m (C},
Leucopogon muticus ta 2m (U), and Aakea
eriantha to. 1.8m in palches. The shrub layer
consisted of Persoonia sericiu to Sien (C),
Notelaea linearis along thé heath edge to 2m (C),
Jacksonia se paria to 2m (U), Mirbelia speciosa
subsp. ringrosei to 80em (C), Hibberria srricta.
Srylidium laricifolium 10 60cm in parches,
Melichrus wrceolams to 40cm (U), Acrotriche
aggregata to 80em (U), Hibbertia linearis var.
obtusifolium (UV). Acacia podalyriifolia along
heath edge to 2.1 m (U), Pomax umbellata Vcr
(C), shorthair plumegrass Dichelachne
micrantha Vo 30cm (C), Lepidosperma laterale to
40cm (C), Austrostipa pubescens in patches to
1.5m, rock fern Cheilanthes sieberi (R), Entola-
via stricta to BOem (A), Paspalidium ` ol.
bovillosum to 80cm (C), Ozothamnus
diosmifolius (R), Poranthera micraph ylla to 8em
(RO, Pimelea linifolia to 50cm (U), wiry panic
Entolasia stricta to 80cm (A), and many uniden-
tified grasses. A thick ground layer consisted of
lullen sticks, leaf litter, bark shreds, rocks, logs
and an abundance of Angophora sp. seed capsules.
Apart from the occasional stray, the collection
site is not grazed hy cattle. and fire may have been
absent from the area for 30 years or more. Un-
burnt skirts on 2m-high Xanthorrhoea individu-
als were noted almost touching the ground.

THE ANIMAL

Adult male H novaehollandiae JM 11409 (Fig.
2) is compared in Table | with measurements
from a wide range of adult males collected in the
Port Stephens area by Keith & Calaby (1968).
While the Port Stephens specimens weighed 13-
18g (collected in February) and IM11409
weighed slighily less at 12g g (collected August)
most measurements suggest t that, in comparison,
the Queensland specimen 15 a small animal. How-
ever, the more commensurate molar measures,
and relative lack of molar wear, suggest that the
Crows Nest mouse may still have been growing.

DISCUSSION

The suspicion that living P novaehollandiae
might occur in Queensland was first aroused in

1993 when Veronica Hinman (University ot

Queensland) analysed unregistered regurgitated

MEMOIRS OF THE QUEENSLAND MUSEUM

ow! pellet material stored in the Queensland Mu-
seum, She attributed fragmentary remains from
three southeast Queensland sites to P
novaehollandiae (Hinman, 1993). These sites
were I, 'Big Rooster Cave’ near Gatton (material
excavated by H. Godthelp in June 1982); 2, Per-
severance section of Crows Nest National Park
(material collected by David Read in May. June
L988); and 3, Cania Gorge (material collected by
Mary Wade in 1978). Confirmation of the identity
of the Gatton ånd Cania Gorge fragments was
sought from Alex Baynes (Western Australian
Museum) who attributed the nine Cania Gorge
dentaries to an unknown pebble-mound mouse
(now known to be Pseudomys patrius, S.N.D.
unpublished data). But he was reluctant to assign
an identity to the maxillary fragments from the
Gatton individual (QM JM10306), its broad,
more rounded molars and the unusual relation-
ship between adjacent cusps on M^ and M^ being
unlike his comparative material of £
novaehollandiae from Smith's Lake, New South
Wales. Given this finding, the more incomplete
Crows Nest National Park fragments (dentaries
and maxillary fragments of one individual QM
JMIO3(4) were temporarily shelved.

More recent comparison with Victorian speci-
mens (Museum of Victoria C26607, C2227,
C1034&, C16047) now suggests that the Galton
specimen is, after all, attributable to a large, old
individual of P. novaehollandiae, ànd reappraisal
of the Crows Nest National Park fragments con-
firms Hinman's assessment that Read's ow! pel-
lets contained a New Holland Mouse. Read's
effort of 600 trap nights between 24 May-~| June
1988 in the Crows Nest National Park resulted in
no mammal captures (Read, 198823) but his owl
pellet material contained examples of Rattus
lutreolus, R. tunneyi, P. gracilicaudatus,
Sminthopsis murina, Perameles nasuta, Petaurus
breviceps and Pseudocheirus peregrinus all of
which are registered it the Queensland Museum
mammal collection.

"Big Rooster’ (south of Gatton) and Crows Nest
National Park are approximately 65km 5 and
Skm E, respectively, of the capture site of
JM11409. In Crows Nest National Park, Read
collected the owl pellets from three caves in Ihe
rocky cliffs (around 27?19'S 152°07'E) over-
looking Perseverance Creek (Read, 1988a).
Godthelp collected the 'Big Rooster’ material
from a rock shelter in eucalypt forest surrounded
by open woodland and degraded grazing country
(the exact coordinates of ‘Big Rooster’ cave have
noL yet been established but are thought to be

THE NEW HOLLAND MOUSE

FIG. 5. Heath adjacent to Crows Nest capture site.

around 27°38'S 152^06'E). Trapping (unsuccess-
ful) for living P. novaehollandiae has since been
conducted in Perseverance section of Crows Nest
National Park, but other sites are yet to be sur-
veyed.

Initially, the collection site of 1M11409 ap-
peared atypical for P. novaehollandiae in its rel-
atively high altitude. distance from the coast, lack
of a dense shrub layer, and advanced seral stage.
It seemed as far removed from the optimum hab-

itat proposed by Posamentier & Recher (1974)
(heath, actively regenerating from fire) as it was
from those habitats occupied in Victoria and New
South Wales (sce Wilson, 1994). However, the
distance from Crows Nest to the coast ( LOOkmi) is
actually exceeded by the distance (166km) from
the type locality (Yarrandi, near Scone, New
South Wales), to the coast, Nearby collection sites
at Belltrees (31*59'S 151?08'E, Kemper, 1977)
and further east near Barrington Tops (1976 record

374

TABLE 1. Comparison of Queensland (Crows Nest)
specimen QM JM11409 against a Port Stephens
(NSW) series of males (213g) (from Keith & Calaby,
1968). Weight expressed in grams, all other measure-
ments in millimetres. For method of measurement see
Taylor et al. (1982).

Maasinersent QMJM Port Stephens series
11409 No. Range Mean
2 | 1 | as | 14 |
Head-bod 78.7 19 84 |
Tail 76.5 17 | 86-101 93
187 | 19 | 205-22| 21
Ear 14.4 19 1|15.5-17| 16
Skull (occipitonasal) | 22.8 18 |231-2251| 242
Condylobasal length 20.7 19 21.9
Basal length ` 18.6 19 |19.1-20.8] 200
Zygomatic width. 11.5 19 11.5-12.7| 119
Interorbital width 3.6 19 3.5-4.1 3.8
Interparietal length 25 | 18 [2938] 34
Interparietal width 7.2 19 7.3-8.3 7.7
| Braincase width 11.0 19 11.1-12.0} 11.5
Mastoid width 9.3 19 |93-108| 99
Nasal length 84 18 8.6
Nasals width 22 17 2.2-2.6 24
Palatal length 11.0 19 11.6-129| 12.3
L. palatal foramen length | 4.9 19 |4554| 49
Palatal foramina width 1.6 19 1.2-1.8 1.5
Inside M'' width 22 19 |2327| 25
Outside MIT width 4.6 19 | 48-53] 50
Bulla length 50 19 | 42-49] 45
Crowns M'? length 34 19 |3538| 3.6
Alveolar M'? length 3.8 19 |3740| 38
Crowns M!” length 2.9 19 25-28 | 27

from New South Wales National Parks and Wild-
life database) are additional examples of P.
novaehollandiae habitat situated far from the
coast. Although Yarrandi, Belltrees and Barring-
ton Tops are approximately 520km south of
Crows Nest, all three collection sites are structur-
ally reminiscent of the Crows Nest site (B. Fox
pers. comm.). Furthermore, examination of un-
published reports and relatively recent trapping
records suggest that around the northern limits of
its range, high altitude and low longitude are
regularly encountered features of the P.
novaehollandiae profile.

Read (1988b) trapped one specimen at
30?05' 12"S 152?28'59"E in Marengo State For-
est (60km southwest of Grafton, NSW) in 1987,
atan altitude of 650m. In August 1992 he trapped
asingle specimen in the Carai State Forest (inland
from Port Macquarie) (Read, 1995), at

MEMOIRS OF THE QUEENSLAND MUSEUM

30°55’39"S 152?05'19"E and around 860m (D.
Read pers. comm.). In September 1993, Sally
Townley trapped six individuals on Carai Plateau,
at 30952'59"S 152?14'14"E, and an altitude of
900m. Here, the site was open forest of mainly E.
laevopinea with an understorey dominated by
Lomandra sp. (S. Townley pers. comm.). Both
Read and Townley have trapped P
novaehollandiae in Chaelundi State Forest. In
March 1991 Read caught an individual at
29°55’S 152?30'E on grass-covered alluvium
with an E. tereticornis overstorey (Read, 1993).
In October 1993, Townley caught several animals
at 30?01'21"S 152?29'46"E at an altitude of
840m. The forest type was New England black-
butt E campanulata, grey gum E. propinqua,
yellow stringy bark E. acmenoides, blue gum E.
saligna and tallow-wood E. microcorys with an
understorey of shrubs, grasses and small herbs
(Townley, 1993). More recently, over 20 individ-
uals of P. novaehollandiae were trapped by
Townley at a number of sites in Oxley Wild
Rivers National Park (southeast of Armidale)
around 30°55-56’S 152?04-12' E, along an altitu-
dinal gradient from about 400-1000m. Trapping
sites supported various types of open forest with
overstorey trees including New England black-
butt, grey box, Blakely’s red gum E. blakelyi, blue
gum and grey gum. Some sites had grassy un-
derstoreys and others had a dense heath layer (S.
Townley, pers. comm.).

Closer to the Queensland border, one specimen
has been trapped at around 900m a.s.l. on
Timbarra Plateau, 29°07’38"S 152*18'22"E, east
of Tenterfield, in forest dominated by New En-
gland blackbutt and E. olida, with a heathy un-
derstorey (Martin, 1995). New South Wales
National Parks and Wildlife database records
document a 1987 record of P. novaehollandiae as
far north as 28°41’S 152?16'E (southeast of
Lindesay View) at around 828m. This, and other
most northerly records from that database appear
with the Read, Townley and Queensland records
in Fig. 1.

All these records confirm both the regular oc-
currence of P. novaehollandiae at high altitudes,
and the use of tall open-forest at low latitudes.
Some of the records confirm its presence in
grassy understoreys. However, the proximity of
grassy understorey to heath (or other forms of
dense understorey) should be examined before
attributing grass as a preferred habitat to the spe-
cies. For example, the Crows Nest capture site
was 182m away from an adjacent area of species-
rich, densely shrubbed (but dry) heath. While

THE NEW HOLLAND MOUSE 37

Kemper (1977) reported an order of habitat pref-
erence in P. novaehollandiae from heath to open
forest regenerating after recent burning, Fox &
McKay (1981) showed that the habitat require-
ments of the species were most likely contained
completely within the shrub layer of heath-type
species with the presence or absence of trees
unimportant. More recently Fox (1996), re-con-
firmed P. novaehollandiae as a species selecting
hare ground and vegetation open to a height of
20cm (typical of the early seral stages). But, he
showed that although in open forest its maximum
abundance occurs around 1.5 years after burning
(followed by decreased abundance), it re-peaked
in abundance around 16 years after the initial
burn, by which time the understorey Was open 10
around 2m above the ground. In heath, similar
peaks occurred at 3 and 13 years following fire,

Male JM11409 (weighing 12g at capture in
August) may have beeen relatively young at the
time of capture. Kemper (1980) recorded sexu-
ally mature and immature males from mid-
coastal New South Wales àt mean weights of
14.3g and 12.9g respectively. One third of males
in her study achieved sexual maturity in the
breeding season of their birth (August to January,
und occasionally to March). That the Crows Nest
male JM11409 was still growing, is made more
plausible by the recent extension of the P
novaehollandiae breeding season (in New South
Wales) to late July (Fox, Higgs & Luo, 1993)
Although trapped in an open grassy understorey,
male JM 11409, if still relatively young, might not
have been residing in the area, but dispersing
through il,

The contemporaneous coastal occurrence of
the New Holland Mouse at low latitude near
Evans Head (29?07'32"S 153°26'31"E)
demonstrates that in the north, habitat parameters
are much broader than those previously ascribed
to the species. Indeed, there seems little reason Lo
doubt that it should also accur in coastal Queens-
land. Habitat resembling that utilised by P
novaehollandiae in Victoria and New South
Wales would appear to occur in coastal and off-
shore southeastern Queensland heaths and
sedged dunes, particularly in localities such as
Coolangatta, North and South Stradbruke Is-
lands, Moreton Is and the Great Sandy Region,

Ironically, the following comment made hy
Keith and Calahy soon after P. novaehollandiae
was rediscovered alive in New South Wales has
not yet outlived iis pertinence 30 years on: This
discovery of å populous colony af the New Hol-
jand panse in am ansa clase ro weban seilemenr

n

and industrial development is an indication of
our lack of knowledge of the abundance and
distribution of at least some members of the in-
digenous fauna! (Keith & Calaby, 1968: 58).

ACKNOWLEDGEMENTS

We thank Rick Galbraith (Crows Nest Shire
Council) for supplying physical feature descrip-
tions of the Shire, Rodney Hancock and Errol
Ryan (Queensland Department of Environment)
for advice on vegetation community distribution
and field trapping sites in Crows Nest National
Park. Heather Janetzki (Queensland Museum)
prepared the map, and along with Marion van der
Donk (Queensland Museum), assisted with cul-
lection and preparation of botanical specimens.
We pratefully acknowledge the help of Alexander
Baynes (Western Australian Museum), Barhara
Trigss (Genoa) and Barbara Wilson (Deakin Uni-
versity), and thank Lina Frigo (Museum of Vic-
toria) and Tish Ennis (Australian Museum) for
lending us specimens in their care, Tim Kingston
(Queen Victoria Museum, Launceston) and
Deyarne Plowman (New South Wales National
Parks and Wildlife Service) kindly supplied their
data records. Bruce Cowell (Queensland Mu-
seum) photographed the animal and habitat fea-
tured in this paper. We particularly acknowledge
Gordon Guymer, Rebecca Francis and Rod
Fensham (Queensland Department of Environ-
ment, Queensland Herbarium) for identifying bo-
tanical specimens, Sally Townley (Southern
Cross University) and David Read (Mount King
Ecological Surveys) for allowing us to usce their
unpublished trapping records, and lan Gynther
(Queensland. Departmen of Environment) and
Barry Fox (University of New South Wales) for
many helpful comments and discussions during
preparation of this manuscript.

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OBITUARY: CHARLES TANNER, HERPETOL-
OGIST, Memoirs of the Queensland Museum 42(1): 377.
1997:- Conversations with herpetologists suggest that some
of us are ‘born’ and some are "made". Charles Tanner (born
19 January, 1911, Brighton, England; died 23 December,
1996, Calms, Ausmalin) was definitely one of the former.
Amongst his earliest memories were those of days spent
"haunting! the reptile house of London Zoo, Wherever he was
— the Unsted Kingdom, Iraq (Abadan), Palestine; Cyprus, the
United Stales of America, Papua New Guinea, or Australia
i (rom Tasmania 10 Cape York and Byron Bay to Carnarvon)
— he observed, admired, photographed and collected reptiles,
and Wrote and talked about them They were a life-long
obsession,

Charles Tanner's contributions to knowledge of
Australia's reptiles were substantial. He had long affiliations
withthe Museum af Victoria, Melbourne, and the Queensland
Museum, Brisbane He was an Honorary Assoclate of the
former from 1953 until his death, His collections for that
museum included 522 frog and 573 reptile specimens, Many
of these were collected on his annual leave. Each year, for
many years, he dnd one of us (AJC) travelled to herpetologi-
cally unknown or poorly known sites, Australia-wide, His
knowledge of, and enthusiasm for finding frogs and reptiles
was a never-ending source of astonishment In 1955, he and
Charles Brazenor (then Assistant Director of the MOV), col-
lected the first specimens of Piera frost Spencer 1901 (a
rare frog, narrowly confined to Mt Baw Baw, Victoria) seen
since collection of the holotype of the species. On New Year
Island, Bass Strait, in 1953 and 1954, he photographed Black
Tiger Snake, Norecliis ater, and Shearwater, Puffinus ren-
uirosteis interactions, Only one of these photographs has heen
published (Worrell, 1963). The rest are now part of his estate,
hopetully tå be lodged in å muscum for posterity.

His association with the Queensland Museum began in
1968, His last donations were made in 1990, Between 1958
and 1990, Charles Tanner donated 302 frog, 548 reptile and
42 mammal specimens to the Queensland Museum. Amongst
(hem are type Specimens oa which desenptions of many new
species were based: Cophixalus xaxatilis Zweilel & Parker,
1977. Carlia dogure Coyacevich & Ingram, 1975; C
Jarnoldae Covacevieh & Ingram, 1975; C. seirtetis Ingram &
Covacevich, 1980; Crenorus astare Czechura, 1986; C.
nullian Ingram & Czechura, 1990; Cryptoblepharus Juhani
Ingram) & Coyacevich, 1978; Lerisra mgrami Storr, (991; and
Leggadina lakedownensis Watts, 1976 (possibly å junior syn-
onym of L forrest), Tanners special interests lay not in
deseription and nomenclature, which he was content ro leave
to others, but discovering new taxa. Recognition ol à new

FIG, |, Charles Tuner (fet) capanne à specimen at O.
microlepidolus on the Morey Plain near W'indorah, SWQ
The sequence was filmed by V ic Martin, 1574. (Palo JAC)

MEMOIRS OF THE QUEENSLAND MUSEUM

species, volluenan of specimens, Meir careful preservariom
and Judgement in a museum were the parts of a bist s
job that he relished, Through this, he encouraged the work
und careers of several taxonomists,

Elapids dominated Tanner's research interests and, 16 å
cennin erent, his life, For many yeurs he kept å wide-range

of species as treasured "pets". Between the carly 1960s and

1985, he ^milked' specimens in his collection daily on behalf
of the Commonwealth Serum Laboratories (now CSL Ltd),
Melbourne, the sole producer of antivenoms in Australia
From his dried venom supplies was made much of the ante
venont produced in Australia. Many of the sarvivors of poten-
tially life-threatening envenomations in Tanner's ‘production
time’ owe their lives, in no small part, to work conducted at
his snake lFarm-Jaboralory near Cooktown. His venons were
always of the highest quality and were used also in mans
research projects. His collaborations with Allen Broad of C81,
were especially rewarding. The most exciting venom studd
was (hat of Cxyuranus miciolepidoras. In. 1979 this venom
was shown lo be the most toxic snake venom n rhe World
Further investigations showed that it was neutralised eflec.
lively by existing Taipan (Oxvu ramus scitellanis) anii venom
(Sutherland et al. 1978; Broad et al, 19798, h). Venans cal
lected by Charles Tanner now form å valuable part of the
National Collection of Venoms held by the Australian Venom
Research Unit of the Department of Pharmacology, The Uni
versity of Melbourne.

Many of his captive snakes had long and very productive
lives in and, sometimes, on! his hands, Tanner survived many
life-threatening bites, Aboul then he was reluctant to talk,
unlike many victims of snakebile (Pear, 1990; Pear er al,
1993), He invariably regarded herpetologists bitten as silly,
mol brave. ‘Aggression is nor å word I would apply m am
snake..." he would say. In 1979, following massive enveno-
Won by å Taipan (COxyaranas scurellatus), Tanner was
fortene to receive the full premedication recommended hy
CSL, The infusion of antivenom was uneventful and Un
patient described as miraculous the rerum of strength o his
limbs and the disappearance of his severe headache. ‘Lite
mother’s milk’ was his description of his therapy. Steraid
therapy followed for the next four days, because of his sex ral
vulnerability to delayed serum sickness. Tanner recovered
uneventfully.

Charles Tannec's expert elapid husbandry involved the
design und development ot several "safe" methods and tools
of trade, all of which are still in use by those who follow hin,
Amongst these are Div potato-masher" jigger and the hoop
bags of plastic und calico for ‘tailing’ specimens of several
species, including Taipans; hide boxes with trap doors; wad
the use of strong, opaque plastic bag from which the largest
elapid could be milked with relative safety, AU minimised
discomfort for tlie snake and maximised safety for the handler.

Tanner figured in the immunological erature as a result
of his Work With rhe large elapids. Following many bites, be
had become highly allergic to CSL amivenom. Forthis reason,
Dr Saul Wiener undertook active immunisation of Tanner
with Tiger Snake (Noreehis seuraus) venom in 1959, Wiener
(1960) successfully immunised Charles Tanner through 24
injections of Tiger Snake venom over 13 months. This protec-
tion was transitory. Thereafter, Tanner rightly became appre
hensive about antivenom therapy.

Al elapids intrigued Tanner, buc F'aipans, Oxyurenus spp.
intrigued him most. Perhaps his greatest excitement and core
tribution was catching the first live specimen seen by re-
searchers of the snake now known as the Western Taipan,
Oxyuranus microlepidolus (McCoy, 1879) Soon after his
close association with rhe Museum pf Victoria began, he read
the desenpiian (1879) of Diemenia au radetééiai and exam.
ned the two type specimens of ihis species in he (ben?

OBITUARY: CHARLES TANNER

FIG. 2. The first ‘milking’ of a specimen of O. microlepidotus,
by Charles Tanner, 1974. (Photo JAC).

National Museum of Victoria. Many years later, following
receipt of a preserved head/tail of this species at the Queens-
land Museum, he and one of us (JAC) relocated this long-lost
species in south western Queensland. Charles Tanner caught
his and the world's first live specimen in Spring, 1974, near
Windorah, SWQ (Fig. 1.). In one week he collected 13 large,
healthy specimens which formed the nucleus of a collection
on which an extensive research programme was based. From
study of those specimens and their progeny, a series of papers
was published on O. microlepidotus. Over some 20 years, the
least well-known elapid species in Australia became probably
its best known (Covacevich & Wombey, 1976; Sutherland et
al., 1978; Covacevich et al., 1981; Shine & Covacevich, 1983;
Covacevich & Tanner, 1983; Broad et al., 1979b; Morrison et
Ga E Covacevich, 1987; Covacevich, 1990; Covacevich,
1994).

In the Australian Venom Research Unit, Department of
Pharmacology, the University of Melbourne, a study to exam-
ine the antibodies to snake venom components and anti-
venoms in the sera of herpetologists is underway. Charles
Tanner had a particular interest in this project. He had not only
two types of antibodies to eight different venom proteins, but
also antibodies to the principle protein in antivenom. Thus, as
his and the other sera continue to be explored, his contribu-
tions to medical research will continue! (Fig. 2.).

Two species have been named to recognise his many
contributions to herpetology: Pseudonaja affinis tanneri
(Worrell, 1961), Lygisaurus tanneri Ingram & Covacevich,
1988.

In addition to his contributions to toxinology and herpeto-
logical taxonomy, Charles Tanner will be remembered for his
wisdom and for the generous way he shared his knowledge
with both amateur and professional herpetologists. He will
also be remembered for his dry sense of humour. Nothing
encapulates his wit and humour so well as a conversation he
had with one of us (JAC) in 1993, It went: JAC: *Charles, how
are you? CT: Fine thanks, except that my memory, especially
my short-term memory, is shot to pieces. JAC: Hell. l'msorry.
That must be difficult for you. CT: It would be, if I could
remember anything to worry about". The words on a memorial
erected on his former farm near Cooktown summarize: ‘Pas-
sionate about the natural world (especially reptiles!), he
contributed significantly in the fields of taxonomy and toxino-
logy, and to knowledge of the natural history of turtles,
crocodiles and elapid snakes. Adventurous, generous of spirit,
witty, intelligent, and ruggedly individualistic, he was ad-
mired and respected by many. Remembered well by his friends
and colleagues.

Alis volat propriis

(He flies on his own wings).

378

Literature Cited

Broad, A.J., Sutherland, S.K. & Coulter, A.R. 1979a. The
lethality in mice of the dangerous Australian snake
venoms and certain exotic snake venoms. Toxicon 17:
661-4.

Broad, A.J., Sutherland, S.K., Tanner, C. & Covacevich, J.
1979b. Electrophoretic, enzyme and preliminary toxic-
ity studies of the venom of the Small-scaled Snake,
Parademansia microlepidota (Serpentes: Elapidae),
with additional data on its distribution. Memoirs of the
Queensland Museum 19(3): 319-329 pls.1,2.

Covacevich, J. 1987. Two Taipans! in Covacevich, J., Davie,
P. & Pearn, J. (eds) "Toxic Plants and Animals: A Guide
for Australia’. (Queensland Museum: Brisbane).

1990. Phangs and Physic: 40,000 years of Risky Business
in Pearn, J. & Cobcroft, M. (eds) ‘Fevers and Frontiers’.
(Amphion Press: Brisbane).

1994. Dandarabilla and Gunjiwuru. The discovery of the
Taipans — the world’s most dangerous snakes. In Pearn,
J.H. (ed) ‘Milestones of Australian Medicine’. (Amph-
ion Press: Brisbane).

Covacevich, J. & Tanner, C. 1983. Come from nowhere ...
then just disappear. Australian Natural History 21(1):
11-13.

Covacevich, J. & Wombey, J. 1976. Recognition of
Parademansia microlepidotus (McCoy), Elapidae, a
dangerous Australian snake. Proceedings of the Royal
Society of Queensland 87: 29-32, pls. 1-2.

Covacevich, J., McDowell, S.B., Tanner, C. & Mengden,
G.A. 1981. The relationship of the Taipan, Oxyuranus
scutellatus, and the Small-scaled Snake, Oxyuranus
microlepidotus Pp. 160-168. In Banks, C.B. & Martin,
A.A. (eds) Proceedings of the Melbourne Herpetologi-
cal Symposium 1980. (The Melbourne Zoological
Board: Parkville).

Covacevich, J., Pearn, J. & White, J. 1988. The world's most
venomous snake. In Pearn, J. & Covacevich, J. (eds)
Venoms and Victims. (Amphion Press and Queensland
Museum: Brisbane).

Morrison, J., Pearn, J., Covacevich, J., Tanner, C. & Coulter,
A. 1984. Studies of the venom of Oxyuranus
micrlepidotus. Clinical Toxicology 21(3); 373-385.

Pearn, J.H. 1990. Herpetologists and snakebite. Memoirs of
the Queensland Museum 29(2): 406.

Pearn, J.H., Covacevich, J., Charles, N. & Richardson, P.
1994, Snakebite in herpetologists. Medical Journal of
Australia 161(5/19 December, 1994): 706-708.

Shine, R. & Covacevich, J. 1983. Ecology of highly venomous
snakes: the Australian genus Oxyuranus (Elapidae).
Journal of Herpetology 17(1): 60-69.

Sutherland, S.K., Broad, A.J., Tanner, C. & Covacevich, J.
1978. The Small-scaled or Fierce Snake (Parademane-
sia microlepidotus) — its recognition as Australia's po-
tentially most venomous snake. Medical Journal of
Australia 1: 288-9.

Wiener, S. 1960. Active immunization of man against the
venom of the Australian Tiger Snake (Notechis
scutatus). American Journal of Tropical Medicine and
Hygiene 9: 284-92.

Worrell, E. 1963. Reptiles of Australia. (Angus & Robertson:
Sydney). 207pp.

J.A. Covacevich, Queensland Museum, Box 3300, South Bris-

bane, Queensland 4101, Australia; S.K. Sutherland, Austra-

lian Venom Research Unit, Department of Pharmacology,

The University of Melbourne, Parkville Victoria 3052, Aus-

tralia; A.J. Coventry, Museum of Victoria, Box 666E, Mel-

bourne, Victoria 3001, Australia; and J. Cann, 27 Yarra Road,

Phillip Bay New South Wales 2036, Australia; 11 March 1997.

CONTENTS (continued)

JAMIESON, B.G.M.
Some new and previously known species of earthworms from Cape York Peninsula ( Annelida:
Ohgochaeta: Meguscolecidae `... 233
LAMPRELL, K.L. & HEALY, IM.
New species of Pteriidae, Pinnidae, Veneridae, Euciroidae and Corbulidae (rom Australia
(Mollusca; Bivalvia: Veneroidà) |... e ie 271
LAMPRELL, K.L. & HEALY, J.M.
Pitur Rómer from Australian and adjacent waters, with descriptians of four new species

(Mollusca: Bivalvia: Veneridae) `... 283
McDONALD, K.R.
A new stream-dwelling Liroria from the Melville Range, Queensland, Australia ............ 307

NEWMAN, L.J. & CANNON, L.R.G.
A new semi-terrestrial acotylcan flatworm, Myoramyxa pardalota gen. ct Sp, nov.
(Plehniidae, Polycladida) from southeast Queensland, Australia `... sg CA? 311
PATERSON, R.A., JANETZKI, H.A. & WILLIAMS, S.C.
Osteology of immature dark shoulder minke whales Balaenoptera acutorostrata from
southern Queensland `... AW Ro De eA MERERI RR Le 315
THOMSON, S.A., WHITE, A & GEORGES, A.
Re-evaluation of Emydura lavarackorum: Identification of a living fossil. ....--.-....-....-
VAN DYCK, S.
Xeromys myoides Thomas, 1889 (Rodentia: Muridae) in mangrove communities of North
Stradbroke Island, southeast Queensland `... AA
VAN DYCK. S. & LAWRIE, B.
The New Holland Mouse Pseudomys novaehollundiae (Rodentia: Muridae), an addition to the

327

mammal fauna of Queensland `... Au MN WV 367

NOTES
COUPER, P.J.

Synonymy of Crenatus monticola Stor, 1984 and Crenoms hypatia Ingram and Czechüra, 1990... . 12
COVACEV ICH, J.A.. COUPER, P.J. & ROBERTS, L.

New data on Lerista ingrami, å rare skink from southern Cape York Peninsula, Australia `... 24
COOK, A.G.

Probable predation scar on a Devonian Brachiopod ..,... ab eese dip Ch bet oh re ce E 50
TURNER, S & COOK, A.G.

Ptyctodont jaw from the Broken River Province, NEQ `. 80
HANNAH, D., AGNEW, G., HAMLEY, B. & HOGAN,L.

New information on the narrowly restricted skink, Nangura spinosa. i... 90
McRAE, PD & COVACEVICH, J.A.

Combat and copulation in Oxvuranus microlepidotus (Elapidae)., e, LOS
DE OLIVEIRA, M.C. & SCHULZ, M.

Echolocation and roost selection in Semon's Leafnosed Bat Hipposideros semoni a.i... 158

SCHULZ, M. & EYRE, T.J.
New distribution and habitat data for the Pygopodid, Paradelma orientalis (Günther, 1876)... 212
WILSON, S.K.
New information on Pseudechis papuanus (the Papuan Black Snake), a medically significant
addition to Australia’s reptiles |... Im mme 232
ARIDIS, J.J.

A range extension for Litoria brevipalmata (Anura: Hylidae) ©... 20200 oe eee 310
PATERSON, R.A.
Bryde's Whale Balaenoptera edeni Anderson, 1878 stranded near Rosedale, Queensland

in1931 ..... OPA ENDE), det ce ON Bee peda 326
COVACEVICH, LA.
Obituary: Charles Tanner, herpetologist 5... eie s 377

ERRATUM

GULLAN, P.J. & STEWART, A.C. 1996 07 20: Å new genus and species of ant-associated cocvid
(Hemiptera: Coccidae: Myzolecantinae) from Canthium Lam. (Rubiaceae)... .+ oa sv viv ann 282

CONTENTS
ARNOLD, P.W. & COOK, P.L.
Some Recent species of the genus Anaskopora Wass, 1975 (Bryozoa: Cribriomorpha)
ME Ws m SET isi deg OE NE 1
BRUCE, A.J.
A new genus of hippolytid shrimp for Thor maldivensis Borradaile ........... lesse 13
CHOY, S. & MARSHALL, J.
Two new species of freshwater atyid shrimps (Crustacea: Decapoda: Atyidae) from northern
Queensland and the distributional ecology of the Caridina typus species-group in Australia .. 25

COOK, A.G.

Gastropods from the Burdekin Formation, Middle Devonian, north Queensland ............. 37
COCK, A.G.

A review of the gastropod'Burdikinia Knight 1937 and Amphelissa Etheridge, 1921 .......... 51
COOK, A.G. & CAMILLERI, N.

Middle Devonian gastropods from the Broken River Province, north Queensland ............ 55
COOK, A.G. & WADE, M.

Symbiotic stromatoporoid-nautiloid association, Middle Devonian, north Queensland ......... 81

COUPER, P.J., SCHNEIDER, C.J. & COVACEVICH, J.A.
A new species of Saltuarius (Lacertilia: Gekkonidae) from granite-based, open forests of
custom FEE RE OE NE EE oe ee eur T kta pale en REO 91
DAVIE, P.J.F. & HUMPHERYS, A.
New species of rhizopine crabs (Crustacea: Decapoda: Pilumnidae) from northern Australia... . 97
DAWBIN, W.H.
Temporal segregation of humpback whales during migration in southern hemisphere waters .. 105
DYNE, G.R.
Two new genera of Acanthodrilinae (Megascolecidae, Oligochaeta) from Cape York
Peninsula aud thé Torres Strait. os ek ey at ree rh rrr ene ee ee pense etre p E 139
FRITH, C.B. & FRITH, D.W.
Biometrics of the birds of paradise (Aves: Paradisaeidae): with observations on variation

and sexual deniowplstE -.-0005 DS ARG Pe Reece Ae Kee beret pere paa dere NIE 159
GARDZINSKA, J. & PATOLETA, B.
Notes on the genus Thorelliola Strand, 1942 (Araneae: Salticidae) ..........0....00 cece eee 213
HOCKNULL, S.A.
Cretaceous freshwater bivalves from Queensland .........uereesnreersesrerresererrorere 223
HUMES, A.G.
Pseudanthessius newmanae, new species (Copepoda: Poecilostomatoida: Pseudanthessiidae)
from marine turbellarians in Australia . 2.2.20... cc cece cee cnet e een eeee 227

(continued inside cover)