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COUNCIL 1970-1971

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Hon. Editor

B. J. Grieve, M.Sc., Ph.D., D.I.C., F.L.S.

G. M. Storr, B.Sc., Ph.D.

R. M. Berndt, M.A., Dip. Anth. (Sydney),
Ph.D. (London), F.R.A.I., F.F.A.A.A.

P. E. Playford, B.Sc., Ph.D.

B. Ingram, B.Sc. (Hons.)

P. G. Wilson, M.Sc.

R. N. Hilton, M.A.

Ariadna Neumann, B.A.

A. S. George, B.A.

S. D. Bradshaw, B.Sc. (Hons.), Ph.D.

S. J. Curry, B.Sc.

A. B. Hatch, M.Sc., Dip. For.

J. H. Lord, B.Sc.

D. C. Lowry, M.Sc.

A. J. McComb, M.Sc., Ph.D.

D. Merrilees, B.Sc., Ph.D.

R. T. Prider, B.Sc., Ph.D., M.Aust.I.M.M., F.G.S.

5. — The Western Australian psolid holothurian Ceto cuvieria (Cuvier)

By DAVID L. PAWSON*
Communicated by E. P. Hodgkin

Manuscript received 28 July 1970; accepted 18 August 1970

Abstract

A unique species of psolid holothurian form-
erly known as Stolinus cataphractus Selenka,
with 15 tentacles, five large radially placed oral
valves, extremely numerous ventral tube feet
and more than 25 Polian vesicles, is redescribed
here. Its apparent identity with a species
illustrated by Cuvier in 1817 necessitates a
change of name to Ceto cuvieria (Cuvier).
Definite locality data are given for the first
time; the species is known from three localities
in south-west Australia. On the basis of gross
morphological characters, Ceto cuvieria is
sharply distinguished from all other members
in the family Psolidae.

Introduction

Holothurians of the family Psolidae are not
a conspicuous element in the Australian fauna.
Clark (1946) recorded five species from Aus-
tralia, while more than 75 species are known
from world seas. Included in the Australian
fauna is a species known only from Western
Australia, which is unique in having 15 ten-
tacles, and in having the strongly arched dorsal
surface of the body covered by a thick leathery
skin, which overlies the dorsal scales. Clark
(1946) referred to the species as Stolinus
cataphractus Selenka, and noted that its locality
data are inexact at best.

Through the courtesy of Drs. W. D. L. Ride and
R, George of the Western Australian Museum I
received for study a large collection of dendro-
chirotid holothurians from Western Australia,
and included in the collection were six speci-
mens of “Stolinus cataphractus”. Study of
earlier literature shows that at present the
species is incorrectly named, and that a psolid
holothurian figured by Cuvier (1817) is undoubt-
edly conspecific with it. The opportunity is taken
here to redescribe the species, and to make
the necessary name changes.

For access to materal, I am particularly grate-
ful to Drs. W. D. L. Ride and R. George of the
Western Australian Museum and Miss Ailsa
M. Clark of the British Museum. I thank Miss
Clark, Dr. D. J. G. Griffin (Australian Museum)
and colleagues at the Smithsonian Institution
for helpful discussions on the status of some
holothurian names.

The correct name for “Stolinus cataphractus”:
Cuvier (1817, 1830) illustrated a holothurian
on PI. XV, fig. 9 (reproduced in fig. 1, here)
in his “Regne Animal”, and called the species
Holothuria cuvieria. On page 239, Cuvier makes
the following statement (original in French) :

* Department of Invertebrate Zoology, Smithsonian
Institution, Washington, D.C. 20560, U.S.A.

squaviata, Mull., Zool. Dan., X, 1,2,3.
A small species inhabiting European
seas; those of hot climates produce
larger ones (1)”

The (1) refers to the footnote which reads
as follows:

“Those which Peron calls the Cuvieries”
It is evident from Cuvier’s illustration of
Holothuria cuvieria that this species is the
psolid from Western Australia (compare figs.
1 and 2 here). Both Bell (1882) and Lampert
(1885) were aware that this was so, but Bell
rejected the early species name, while Lampert
referred to the species as Psolus cuvieria
Jager.

In the terms of the International Code of
Zoological Nomenclature the species Holothuria
cuvieria Cuvier is valid, but as Bell (1882) has
pointed out, Blainville (1821) and Brandt (1835)
were incorrect in citing Cuvier (or Peron in
Cuvier) as the author of the genus-name
Cuvieria , using Cuvier’s footnote as a basis for
their decision.

Figure 1. — Original illustration of Holothuria cuvieria.
From Cuvier (1830); pi. XV fig. 9.

Jager (1833) was the first to diagnose the
genus Cuvieria for the echinoderms, and he in-
cluded two species, which he referred to (p. 20)
as “Bp. 1 Squaviata Muller . . and “Sp. 2
Cuvieria . . . Cuvier ...” The species cuvieria
is the type-species of the genus Cuvieria by
absolute tautonymy, according to Article 68 (d)
of the Code.

Cuvieria Jager is preoccupied (by Cuvieria
Lesueur and Petit, 1807). Nevertheless the
name persisted in use as a senior synonym in
the holothurians for many years (e.g. Studer,
1877; Bell, 1881). Gistel (1848) was apparently
aware of the precarious position of the

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

33

echinoderm genus-name Cuvieria, and <p. viii)
proposed a replacement name, Callisto, as fol-
lows :

“ Cuvieria (Agass. prodr. echin.): Cal-

listo, N.”

Unfortunately Callisto is also preoccupied (by
Callisto Stephens, 1834), and thus is not avail-
able for echinoderms. Later in the same work
(p. 174 h Gistel (1848) proposed another new
name for Cuvieria . as follows:

‘ Cuvieria (H squamata, Miill.): Ceto

(mihi)

As the above was included in his section
“Echinodermata” (p. 175), and as the name
H. squamata Muller is obviously used in refer-
ence to the holothurian of that name, it is
obvious that Gistel was referring to the
echinoderm genus-name Cuvieria , and not to
any other hymonyms. There is also little doubt
in my mind that Ceto is in fact being proposed
here as a replacement for Cuvieria. Gistel may
or may not have been aware that Callisto was
preoccupied, but this now has no bearing
on the case.

It may be argued that it was Gistel 's inten-
tion to provide the name Ceto for Holothuria
squamata only, and that the species name
cuvieria should not be construed as belonging
in the genus Ceto, but it would seem more
logical to assume that Ceto was proposed simply
to replace the genus name Cuvieria , and as the
species cuvieria is ipso facto the type-species
of Cuvieria then the combination Ceto cuvieria
is valid.

Later authors have referred to the Western
Australian species by various names (see synony-
mies below), and it is indeed remarkable that
a species which is known from so few specimens
(approximately six specimens, not including the
collection described here) has been referred to
under 6 generic names, and no less than 11
binomial combinations.

It has been suggested by some colleagues that
the name Stolinus cataphr actus Selenka, 1868
should be applied to this species; it is referred
to as such by Clark (1946). If this step were
taken, it would require application to the Inter-
national Commission on Zoological Nomenclature
for suppression in the interest of stability of the
names Holothuria cuvieria Cuvier and Ceto
Gistel. Clark (1946) did not have occasion to
study specimens of this species, and did not
present a case for maintaining the name Stolinus
cataphractus. I feel that the interests of stab-
ility would best be served by retention of the
name Ceto cuvieria, because it definitely fixes
the identity of the species named Holothuria
cuvieria by Cuvier, because Ceto is, as far as I
can determine, a perfectly valid generic name,
and because the troublesome names Cuvieria,
Callisto, Stolinus, Hypopsolus and Theelia are
thereby reduced to the status of junior synonyms.
The use of the incorrect name Stolinus cata-
phractus by Clark (1946) does not appear to be
sufficient justification for retention of that name.

Order Dendrochirotida Grube, 1840 (restricted
Pawson and Fell, 1965)

Family Psolidae Perrier, 1902
Ceto Gistel, 1848

Cuvieria Jager, 1833, p. 20 (preoccupied by
Cuvieria Lesueur and Petit, 1807); type-
species Holothuria cuvieria Cuvier, by ab-
solute tautonymy.

Callisto Gistel, 1848^p. viii; replacement name
for Cuvieria Jager (preoccupied by Callisto
Stephens, 1834).

Stolinus Selenka, 1868, p. 110; type-species cata-
phractus Selenka, by monotypy.

Psolus < Hypopsolus ) Bell, 1882, p. 648; type-
species ambulator Bell, by monotypy.
Theelia Ludwig, 1892, p. 350 (preoccupied by
Theelia Schlumberger, 1891).

Type-species: Holothuria cuvieria Cuvier. (By
original tautonymy ) .

Diagnosis: Psolids with five conspicuous radially
placed oral valves. Anus with five or more in-
conspicuous valves. Sole with extremely numer-
ous tube feet in all three radii, and scattered in
two interradii. Large dorsal scales covered with
leathery skin. Dorsal surface with scattered
pores which contain tube feet, and which pass
through or between scales. Tentacles 15. More
than 25 Polian vesicles.

Content of the genus Ceto and status of
Cuvieria porifera Studer :

Ceto is here regarded as monotypic. Studer
(1877) described Cuvieria porifera from Royal
Sound, Kerguelen Island, 65 fathoms. He diag-
nosed the species as follows:

“Dorsum medium alepidotum, sulcis de-
cussatis corrugatum, interstitus 1-2 porosis,
granulis raris munitis, margine squamis 1-2
porosis tectum. Facies ventralis mollis,
ambulacris marginalibus, mediano nullo.
Color fuscus. Hab. Kerguelen.”

Theel (1886, p. 130) noted that the description
is “. . . very summary and unsatisfactory. The
species is possibly identical with Bell’s Psolus
ambulator ” (Psolus ambulator- Ceto cuvieria:
see synonymy below). Ludwig (1892) essentially
repeated Theel’s suggestion, and in a later paper
(1898) synonymized the two “species” under the
name Theelia porifera. Subsequently, some
authors have come to regard porifera as a syn-
onym of the species here referred to as cuvieria.
Others, of whom the most recent is Clark (1946),
have maintained that they are distinct.

The presence of very numerous tube feet in the
midventral radius of Ceto cuvieria immediately
separates that species from porifera. in which
Studer (1877. p. 452) notes that the midventral
radius is naked. I have not been able to
examine material of Cuvieria porifera, and until
more material becomes available, the status of
the species is problematical. It is possible, how-
ever, that Psolidium incertum (Theel) from the
vicinity of Kerguelen Island is synonymous with
Cuvieria porifera. In his original description,
Theel (1886, p. 86) notes that P. incertum has
a naked midventral radius and that the dorsal

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

34

B

D

Figure 2. Ceto cuvieria (Cuvier). A. Lateral view, 71mm specimen from locality 2. B. Oral view, 63 mm speci-

men from locality 1. C. Dorsal view, 63 mm specimen from locality 1; note dorsal pores. D. Ventral view, 71
mm specimen from locality 2. E. Lateral view, 63 mm specimen from locality 1, showing conspicuous dorsal

pores.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

35

G

100m

■ 50m

Figure 3 . — Ceto cuvieria (Cuvier). A. Cup-shaped deposits from dorsal surface of body. B. Simpler types of cup-
shaped deposits from sole. C. Open meshwork deposits from dorsal surface of body. D. Perforated rods from
stems of tentacles. E. Deposits from tube feet. F. Smaller plates from branches of tentacles. G. One-to two-
layered deposits from stems of tentacles.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

36

scales carry one, two or three small pores. As
both species are from the same general locality,
and as both share the morphological features
discussed above, the evidence for uniting them
is quite strong. At least, the species poriferus
should be referred to the genus Psolidium . The
synonymy of the species is given here.

Psolidium poriferum (Studer)

Cuvieria porifera Studer, 1877, p. 452; 1879. p.
p. 123.

Psolus porijerus: Lampert, 1886, p. 122; Theel,
1886, p. 130; Lampert, 1889, p. 854.
Theelia porifera (part) : Ludwig. 1898, p. 52.
Theelia ambulatrix: Erwe, 1913, p. 358.

The synonymy of Ceto cuvieria is now T as
follows :

Ceto cuvieria (Cuvier)

Figures 1-3

Holothuria cuvieria: Cuvier. 1817, pi. 15 fig. 9;

Blainville, 1821, p. 315; 1834, p. 192.

Cuvieria cuvieria: Jager, 1833, p. 20.

Stolinns cataphractus: Selenka, 1868, p. 110, pi.

8 figs. 1,2; Clark, 1946. p. 413.

Psolus cuvierius: Semper, 1868, p. 63, 240.
Psolus (Hypopsolus) ambulator: Bell, 1882,

p. 648, pi. 48 fig. 2.

Psolus ambulator: Lampert, 1885, p. 118; Theel,
1886. p. 131.

Psolus cuvieria: Lampert, 1885, p. 121.

Psolus cataphractus: Theel, 1886, p. 131.
Theelia ambulatrix: Ludwig, 1892, p. 350; 1894,
p. 136, Erwe, 1913, p. 358,

Theelia cataphracta: Ludwfig, 1892, p. 350.
Theelia porifera (part) : Ludwig, 1898, p. 52.

Diagnosis: As for the genus (see above).
Remarks: Only approximately six specimens of
this species have been recorded. Selenka (1868)
had access to two specimens, and the remaining
authors (Cuvier, 1817; Bell, 1882; Theel, 1886;
Erwe, 1913) apparently each had one specimen
at his disposal.

Material Examined: Six specimens from three
localities, as follows:

1. Smith Rock, Recherche Archipelago,
Southwest Australia, 15-37 fathoms (27-66
metres), 2 Feb., I960, coll. R. W. George on
"Bluefin”, 2 specimens.

2. Four miles off Dunsborough, Geographe
Bay, Western Australia, 55 feet (16.5 metres),
18 April, 1960, coll. B. R. Wilson, 3 specimens.

3. Three miles off Dunsborough, Western
Australia, 50 feet (15 metres), 20 Dec., 1961,
coll. B. R. Wilson and R. Slack-Smith, 1 speci-
men.

Description

External features: Total length of sole 58-71
mm. All specimens but one (locality 3) with
tentacles retracted. Body elongate, oval, with
well-defined flat sole; dorsal surface arched,
mouth anterior dorsal, surrounded by five con-
spicuous radially placed triangular oral valves;

less conspicuous anus posterior dorsal sur-
rounded by five or more very small valves
(Fig. 2c). Expanded specimen with 15 richly
branched tentacles, all of approximately equal
size. Colour in alcohol variable; specimens
from localities 2 and 3 dark browm to black
dorsally, with scattered grey-brown patches
dorsally and laterally. Sole uniformly dark
grey to grey-brow r n or black. Specimens from
locality 1 mottled and variegated dark grey or
greyish white dorsally; sole dark grey to
greyish-white. In expanded specimen from
locality 3, tentacles and introvert uniformly
black.

Dorsal surface of body extremely thick
(2-3 mm), consisting of several large scales up to
1 cm in diameter, overlain by thick leathery
rugose skin filled with calcareous deposits.
Scattered on dorsal surface but tending to be
concentrated in areas of dorsal radii are pores,
through W'hich pass small tube feet, w T hich are
often retracted. Pores either single, or in pairs,
sometimes superficially resembling echinoid
diplopores. In greyish coloured specimen from
locality 1, pores conspicuous, black, approxi-
mately 0.5 mm in diameter. In other specimens
pores less conspicuous against darker coloured
background. On dorsal surface of greyish speci-
men are 27 pores; of these 8 are located close
to anal aperture. This total of 27 includes a
single pore on each of the large oral valves;
in some specimens, the pores are more fre-
quently double than single; this is especially
noticeable on the oral valves. Dissection show's
that w'eli developed tube feet pass either be-
tween large dorsal plates, or through them.
The holes which pass through the plates, often
take a meandering course.

Dorsal surface fringed toward ventral surface
by numerous smaller plates; these form w T ell-
defined edge to dorsal surface, delimiting it
from sole.

Sole thick, leathery, with extremely numerous
tube feet, which are closely aggregated at an-
terior and posterior ends of sole and along
lateral ventral radii. They are less numerous
near centre of sole, but are nevertheless present
in large numbers on midventral radius and
scattered all over interradial areas. Tubefeet
highly retractile, and can be retracted deep into
sole, leaving conspicuous holes. Scattered around
sharp ventral edge of dorsal surface are small
tubefeet, w r hich are presumably sensory.

Internal features: Dissection rendered extremely
difficult by thickness of dorsal body w T all and
sole. Internal anatomy presents many unusual
features, which will form subject of later paper.
One specimen (female) dissected; briefly, major
features are as follow's: —

Gut typical, with single coil. Respiratory
trees well developed, short, less than i body
length with numerous branches. S’tone canal
short, terminating in nodular madreporite
located on right side of dorsal mesentery. Polian
vesicles numerous, 28 being regularly scattered
around posterior edge of calcareous ring.
Vesicles bulbous, large, up to 10 mm in length.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

37

Retractor muscles short, well developed,
attaching to radial longitudinal muscles by
means of a muscular membrane at a level
slightly posterior to oral valve complex. Radial
longitudinal muscles thin straps. Circular
muscles well developed, thrown into ridges about
2-4 mm apart.

Gonad a bunch of unbranched caeca up to
approximately 20 mm long, originating anteriorly
at level of posterior edge of oral valves. Caeca
contain eggs approximately 0.8 mm in diameter.
Genital duct short, opening to exterior mid-
dorsally within crown of tentacles.

Calcareous deposits: Scales on dorsal surface
rounded, smooth, extremely thick (1.8 mm),
joined to each other by flexible leathery integu-
ment. Overlying massive dorsal plates are two
kinds of deposits. First type are longer than
broad, consisting of an open meshwork of cal-
cite (Fig. 3C> somewhat reminiscent of the
hourglass-shaped deposits in some species of
Lissothuria (see Pawson, 1967). These are ex-
tremely numerous, averaging 93 p in length.
Also present and also extremely numerous are
second type, dense deposits, approximately cup-
shaped, with one end rounded and the other
truncate (Fig. 3A). Cup consists of a dense and
intricate network of calcite. Truncate surface
carries numerous rounded projections. Average
length of cups 105 average greatest width,
across truncate portion, 90 /x.

Sole contains “cups” which show some simil-
arities to those of dorsal surface, except that
cup portion is generally shorter, and sole spic-
ules tend to be somewhat flatter than those
from dorsal surface. However, as in cups of
dorsal surface, one side of cup is rounded, while
other carries numerous short projections, and
entire spicule is composed of complex network
of calcite (Fig. 3B>. Average diameter across
greatest width of cups is 96 /<♦ No “hourglass-
shaped” deposits found in sole.

Tubefeet of sole with well-developed 1-2
layered end plates approximately 800 fx in
diameter. These are surrounded by smaller per-
forated plates which may be simple and one-
layered or two-layered (Fig. 3E), and vary
greatly in size. Dorsal tube feet apparently lack
end plates but do contain numerous one-layered
perforated plates, polygonal to circular, 136 fx
in average length.

Branches of tentacles armored with massive
curved plates with numerous perforations, the
plates variable in shape, up to 1.1 mm long.
Numerous smaller, simpler plates (Fig. 3F),
greatly variable in shape and size, also present.
Stems of tentacles contain elongate perforated
rods (Fig. 3D) and also perforated one-to two-
layered “buttons” with marginal and surface
projections (Fig. 3G). All obviously derived from
a four-holed form, in which two holes are larger
than others.

Additional material: British Museum (Natural
History) specimen No. 63.9.23.32, (holotype of
Psolus f Hypopsolus) ambulator Bell, 1882).
Specimen virtually identical to those described
above. Cups in sole slightly smaller (average
diameter 88 u > than those in Western Australian
Museum specimens.

Distribution : Until now, no definite locality data
have been available for this species. Jager
(1833), Selenka (1868), Bell (1882), Theel (1886)
and Ludwig (1898) merely cite “Australia” as
the locality for their material. Erwe (1913) des-
cribed a single specimen from the collection of
the museum at Perth, Western Australia. The
present records definitely establish the fact that
the species occurs off South West Australia, in
depths of 15-66 metres.

Remarks

The major morphological features of the pre-
sent specimens agree in large part with those
described by Selenka (1868). Bell (1882) and
Theel (1886), None of these authors, however,
called attention to a most conspicuous feature
of this species, and that is that the oral valves
are radial in position, while in all other psolids
with large oral valves they are interradial.
Further, although both Bell (1882) and Theel
'1886) described the pores in the dorsal surface
cf the body, neither noted that the pores contain
well-developed tube feet. The feet can be com-
pletely retracted within the pores, and usually
they are retracted so that they are not visible
without dissection. The most remarkable feature
of the internal anatomy is the presence of a
large number of Polian vesicles. Selenka (1868)
noted that a “large number” (p. Ill) of Polian
vesicles are present, while in the specimen dis-
sected here, there are 28 vesicles.

The relationships of Ceto cuvieria are not
clear. Within the Psolidae, the cosmopolitan
genus Psolidium Ludwig and the New World
tropics genus Lissothuria Verrill also carry tube
feet on the dorsal surface of the body, and the
latter genus has hourglass-shaped deposits (see
Pawson, 1967) which show some similarities to
those of the present species. But the presence in
Ceto cuvieria of radially placed oral valves, a
large number of Polian vesicles, 15 tentacles, and
a thick leathery skin overlying the scales sharply
distinguish the species from all others in the
family Psolidae.

References

Bell, F. J. (1881). — Account of the zoological collections
made during the survey of H.M.S. "Alert"
in the Straits of Magellan and on the coast
of Patagonia. IX. Echinodermata. Proc.
Zool. Soc. London . 87-101, pis. 8-9.

(1882). — Studies in the Holothuroidea.— I.

On the genus Psolus and the forms allied
thereto. Ibid.,; 641-650. pi. 48.

Blainville, (1821). — Holothuries. In; Dictionaire

des Sciences Naturelles 21.* 310-319. Paris.

(1834). — Manuel d’ Actinologie ou Zoophy-

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(1830). — Ibid., nouvelle edition, revue et

augmentee.

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(1894). --The Holothurioidea. (Report of

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(1898). — Holothurien. Ergebn. Hamburger

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years 1873-76. Part II. “Challenger” Sci.
Results — Zoology 4, pt, 39: 290 pp., 16 pis.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

39

6. — Two new genera and five new species of Percophidid fishes (Pisces:

Percophididae) from Western Australia

by R. J. McKAY*

Manuscript received 16 June 1970; accepted 14 December 1970

Abstract

Two new genera, Branchiopsaron, Liopsaron,
and three new species belonging to the genera
Bembrops and Chrionema are described from
Western Australia.

Introduction

A collection of fishes trawled in 15 to 202
fathoms off the north Western Australian coast
has been donated to the Western Australian
Museum by Captain K. Ozawa of the Japanese
Research Vessel Umitaka Maru. The collection
contains many fishes unrecorded from Western
Australia.

The family Percophididae is predominantly
Indo-Pacific in distribution; Percophis is only
found in the tropical Atlantic, and the genera
Bembrops and Cliriomystax have representatives
in the Atlantic Ocean as well as the Indo-Pacific.

Only the genus Acanthaphritis , from the
Arafura Sea, has been recorded from the eastern
Indian Ocean. This family has been previously
recorded in eastern Australian waters by one
species, Enigmapercis reducta Whitley, from
New South Wales.

A brief key to the percophidid genera is given
in Schultz (1960; 273-277) and a key to some
of the genera is provided by Okamura and
Kishida (1963; 47-48).

B Space between mandibular rami

scaled .... .... .... .... .... Cliriomystax

Ginsburg

1955

4. A Lower jaw projecting, noticeably
longer than upper jaw; dorsal
spines 8 or 9; branched caudal
rays 11 Percophis

Quoy and

Gaimard

1824

B Lower jaws not projecting; dorsal
spines 6 or less; branched caudal

rays 7 or 8

5

5. A

Outer edge of nasal
ringed with inwardly
cirri

openings

projecting

Cirrinasus

Schultz

1960

B

Outer edge of nasal
without cirri

openings

6

6. A

Dorsal spines 2 or 3;
15-18

dorsal rays

7

B

Dorsal spines 4 to 6;
20-26

dorsal rays

8

7. A

Branchiostegal rays 7

Enigmapercis

Whitley

1936

B

Branchiostegal rays 5

Matsubaraea

Takai 1953

Key to the genera of Percophididae

1. A Opercle with two conspicuous

spines; subopercle with single
spine; scales on body ctenoid, the
anterior-most scales of the lateral
line armed with a keel; branched

caudal rays 11 2

B Opercle with a single spine, a
ridge, or unarmed; subopercle
without a spine; scales on body
cycloid or weakly ctenoid, the
anterior-most lateral line scales
without a keel; branched caudal

rays 11 or less 4

C Opercle with a single spine;
subopercle with a strong spine;
no scales on body, lateral line
absent; branched caudal rays 8 Liopsaron new

genus

2. A A dermal tentacle or flap on the

posterior end of the maxillary Bembrops

Steindach-
ner 1876

B No dermal tentacle or flap on
maxillary 3

3. A Space between mandibular rami

naked Chrionema

Gilbert 1905

* Western Australian Museum, Perth, Western Australia

8. A Subopercle with a posterior flap
that extends on to the lower part
of the pectoral base; scales on
body weakly ctenoid; teeth on
vomer

B Subopercle without a posterior
flap; scales on body cycloid; no
teeth on vomer

11

9. A Ceratohyal with an inwardly

directed process; dorsal fin rays
24-26; anal rays 29-31; branchio-

stegal rays 7 Branchiopsaron

new genus

B Ceratohyal without an inwardly
directed process; dorsal fin rays
20-23: anal rays 24-26; branchio-
stegal rays 6 io

10. A Cleft of mouth oblique; opercle

unarmed; no barbel on upper

jaw Acanthaphritis

Gunther

1880

B Cleft of mouth almost horizontal;
opercle with a ridge along the
dorsal margin: a barbel at the
symphysis of the upper jaw ... Spinapsaron

Okamura
and Kishida
1963

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

40

11. A Inner edge of preopercle with a
row of cirri that interdigitate
with gill rakers on the first gill
arch; anal rays 27; fin rays of
first dorsal and anal elongate .... Pterapsaron

Jordan and
Snyder 1902

B Inner edge of preopercle without
cirri; anal rays 22; fin rays not

elongate Osopsaron

Jordan and
Starks 1904

Okamura and Kishida (1963: 46-47) have
made an attempt at grouping some genera. Fol-
lowing a survey of the literature I have included
all the genera of Percophididae in the grouping
below. Many genera are mono ty pic and a revision
of the family, although perhaps still premature,
may reduce the number of genera considered
below.

Group 1 contains Bevibrovs , Chrionema and
Chrioviystax. The opercle has two spines
and the subopercle one spine in this closely
related group.

Group 2 contains Acanthaphritis, Branchiop-
saron and Spinapsaron } these genera
possessmg preorbital spines, a subopercular
flap (also found in Matsubaraeat and 4-5
dorsal spines.

Group 3 contains Pteropsaron and Osopsaron;
these genera lack the preorbital spmes and
subopercular flap.

Group 4 contains Matsuharaea and Enigmaper-
cis. These two genera seem to be morpho-
logically similar in having teeth on the pal-
atines, 2-3 dorsal spines, 18-19 dorsal rays,
reduced gill rakers, almost cycloid body
scales and no preorbital spines. The
branchiostegal ray count differs, and Whit-
ley (1944 p. 254, text fig.) shows the opercles
as approaching or overlapping the pectoral
fin base, whereas in Matsubaraea there is a
distinct subopercular flap. This group is
similar to groups 2 and 3.

The three remaining genera of the family,
Cirrinasus, Percophis and Liopsaron are quite
distinct from the genera listed above. Cirrinasus
approaches group 4 in fin formula; Percophis is
very different in having a large number of dor-
sal and anal fin elements (D.VIII-IX, 31: A.i
38-40 ) ; Liopsaron is most aberrant in lacking
scales, and has a single opercular and suboper-
cular spine. In general morphology this latter
genus is rather similar to groups 2, 3 and 4 but
shows relationships with group 1 in having a
subopercular spine.

Branchiopsaron McKay, new genus
Type species; Branchiopsaron ozawai McKay,
new species.

Body slender and subcylindrical, covered with
large thin scales with weakly ctenoid ragged
edge Opercles and cheeks with cycloid scales;
snout naked. Lateral line along mediolateral
axis of body, scales differentiated. Eyes large,
mostly dorsal in position and separated by veiy
narrow interorbital ridge. Snout depressed, with
well developed preorbital spines not projecting

beyond outer margin of upper jaw. Upper jaw
slightly longer than lower; mouth slightly
oblique; maxillary expanded posteriorly, without
flap. Upper and lower jaws with about three
rows of small depressable teeth; vomer with two
well-separated, raised mounds bearing strong
teeth; palatines without teeth. Tongue without
teeth, narrow posteriorly between vomerine
patches, but dilated anteriorly into spoon-
shaped process, lying well anterior to vomer,
Ceratohyal with terminal w ? ell-developed in-
wardly-projecting process. Opercle with single
spine; subopercle with membranous posterior
flap covering low’er half of pectoral fin base;
posterior-dorsal edge smooth. Gill memberanes
free from isthmus. Twn dorsal fins, first fin low
with five slender spines; second fin with twenty
four to twenty six rays. Anal fin rays twenty
nine to thirty one. Branchiostegal rays seven.

This new T genus is similar to the genera Spin-
apsaron Okamura and Kishida, Pteropsaron
Jordan and Snyder, Osopsaron Jordan and
Starks, and Acanthaphritis Gunther, but can be
separated by the high dorsal and anal fin counts,
and the presence of a w'ell developed, flattened
hook-like process on the terminal end of the
ceratohyal: this process projects into the bran-
chial chamber, and has not been reported in the
above genera. The high branchiostegal ray count
of 7 is also diagnostic.

Branchiopsaron is most similar to Spinapsaron
in possessing a flap on the subopercle, preorb-
ital spines, and edentulous palatines, but differs
in having a high ray count, no barbel on the
premaxillary, a process on the ceratohyal, seven
branchiostegal rays instead of six, and in lacking
the high posterior rays of the soft dorsal fin. The
posterior-dorsal margin of the subopercular flap
is smooth in Branchiopsaron instead of serrated
as in Spinapsaron . Acanthaphritis , the only
other genus with what appears to be a developed
subopercular flap (Gunther, 1880, pi. XVIII, fig.
A), has the cleft of the mouth oblique, unarmed
opercles, a low^er number of dorsal and anal
rays, and six branchiostegal rays.

Derivation: Branchiopsaron comes from the
Greek branchia meaning gills and psaron mean-
ing little fish. Gender masculine.

Branchiopsaron ozawai, McKay, new T species
(Figures 1, la, lb)

Description: Based on the holotype; counts and
proportions of 6 paratypes given within paren-
theses.

Dorsal V, 26 (V, 24-26), anal 31 % (29-310,
pectoral 21 (21), pelvic I, 5 (I, 5).

Lateral line scales unlike those of remainder
of body in possessing a sculptured margin (see
figure lb) 40 in longitudinal series (39-43);
3 above, 3 below T lateral line.

Head 31.3 (29.4-31.1), depth 12.3 (10.9-11.8),
snout 4.2 (4.0-4.7), eye 9.9 (10.1-10.9), inter-
orbital ridge 0.7 (0.6), head width 14.9, length
of maxilla 14.8, snout tip (between preorbital
spines) to first dorsal origin 31.2 (32.1-33.2),
snout tip to second dorsal origin 15.7, snout
tip to vent 36.3. Second dorsal fin base 51.8,

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

41

Figure 1 . — Branchiopsaron ozawai sp. nov. lateral view; la. dorsal view of head; lb. lateral line scale.

anal fin base 60.0, length of pectoral fin 20.3,
length of pelvic fin 13.9, length of caudal fin
18.9. length of first dorsal spine 9.7, length
of 3rd dorsal ray 9.5, length of 4th anal ray
7.2, length of preorbital spine 2.2. Depth of
caudal peduncle 5.0; all expressed as percent
of standard length taken from between the
preorbital spines to the commencement of the
caudal fin.

Head somewhat flattened above, widest at
preopercle margin. Mouth wide with upper jaw
slightly projecting beyond lower jaw. Nostrils
situated close to anterior margin of eye. Space
between preorbital spines concave; spines reach-
ing anterior margin of premaxillae. Maxilla
extending little beyond middle of orbits, its
posterior end expanded and truncate, without
flap. Teeth small and conical in 2-3 rows in
both jaws. Gill rakers consisting of low patches
of teeth situated along gill arch, one patch
above angle, nine below. No row of cirri on
inner preopercle edge. Hind margin of pre-
opercle free and quite thin. Subopercle with
flap of skin extending over basal half of pec-
toral fin. Pectoral fin extending to 4th dorsal
ray. Pelvics commencing well before pectorals;
their bases widely separated. Second dorsal
fin originating above 4th anal ray. Caudal fin
with eleven rays; seven rays branched.

Colour i7 1 formalin: Body with dorsal scale
pockets possessing dark margins, sides with a
diffuse longitudinal band ( broken into dusky
spots in one paratype), lower sides white. Head
greyish, with grey opercles. First dorsal fin
base pale, remainder of fin intensely black.
Second dorsal, anal, pectoral, pelvic and caudal
fins hyaline.

Colour in life: With about seven bright

lemon-yellow spots, about one scale in diameter
situated dor sally to the faint longitudinal grey
band on sides.

Holotype: WAM P 19153, 105.8 mm in stan-
dard length, trawled by Umitaka Maru
in 350 metres, Station UMPT 6905, 17°17'0"S'.
119 D 57'0"E, December 20, 1969.

Paratypes: WAM P 19152, s.l. 108.5 mm;
WAM P 19154, s.l. 86.5 mm; WAM P 19155,
s.l. 93 mm; WAM P 19156, s.l. 70 mm; WAM
P 19157, s.l. 65.5 mm; WAM P 19158, s.l. 57
mm; Station UMPT 6905, data as above.

Derivation: Named in honour of Captain K.
Ozawa of the Umitaka Maru .

Bembrops aethalea McKay, new species
(Figure 2)

Description: Described from the holotype;
proportions and counts of 9 paratypes given
within parentheses.

Dorsal fin VI 14 (VI, 14-15); anal 1 6h (16£);
pectoral 27 (27); pelvic I, 5 (I, 5). Gill rakers
2/13.

Lateral line scales 54 (51-56). Transverse
scales below rayed dorsal fin origin 6 above, 4i
below lateral line; 3 scales between lateral line
and first dorsal fin.

Head 38.3 (36.0-39.4), depth 13.7 (12.7-15.3),
snout 10.3 (9.6-11.1), eye 10.7 (9. 0-9.8), inter-
orbital 2.3 (1. 1-2.2) snout to first dorsal origin
38.1 (36.9-38.2), depth of caudal peduncle 6.5.
pectoral length 21.5, pelvic length 17.5, caudal-
fin length 19.8. second dorsal base 33.9, anal
base 35.1, greatest width of head 19.5, length
of maxilla 14.7, all expressed as percent of
standard length.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

42

Head flattened dorso-ventrally, widest at
opercles. Mouth wide with lower jaw project-
ing beyond upper jaw. No preorbital spines.
Maxilla extends little beyond anterior margin
of eye; posterior end truncate with well de-
veloped flap of skin extending to middle of
eye. Opercular and subopercular spines pre-
sent and equidistant; opercular flap reaching
sixth lateral-line scales. Small spine directed
posteriorly above origin of opercle. Anterior
four lateral -line scales with median raised keel.
Tongue without teeth; tip slightly expanded.
Teeth on vomer and palatines. Fine conical
teeth on both jaws, those on upper jaw extend-
ing outside mouth. Gill rakers 2/13 (2/13-
4/12) well developed with a row of small spines
on inner surface. Gill membranes free from
isthmus. First dorsal spine not filiform.

Colour in formalin: Body with about 13 smoky
cross-bars on lower .sides, cross-bars varying
between one and 3 scales in width. Edge of
dorsal scales dark forming a network pattern
on the back. Upper and lower caudal rays
dusky (a caudal spot present on the base of
the upper caudal rays in some paratypes, the
spots fading with an increase in body length).
First dorsal membranes greyish; second dorsal
fin membranes with dark basal streaks. Anal
and pelvic fins hyaline. Pectoral fin faintly
dusky with a vague basal streak. In life, bright
lemon cross-bars present on the body.

Holotype : WAM P 19144, 177 mm standard
length, 250 metres, 13°45'S, 123° 30' 5"E, Sta-
tion UMPT 6909, December 23, 1969.

Paratypes: WAM P 19142, s.i. 179 mm; WAM
P 19143, s.l. 180 mm; WAM P 19145, s.l. 183
mm: WAM P 19146, s.l. 138 mm; WAM P
19147. s.l. 143.7 mm; Station UMPT 6909.
WAM P 19148, s.l. 179 mm; WAM P 19149,
s.l. 175 mm: WAM P 19150. s.l. 170 mm;

WAM P 19151, s.l. 121 mm; Station UMPT
6911, depth 242 metres, 13 C 47'0"S, 123 C 18'0"E,
December 23, 1969.

Two other Indo -Pacific species of Bembrops
lack a filamentous first dorsal spine, B. caudi-
maculata Steindachner and B. adenensis
Norman. From B. caudimaculata this new
species differs in coloration (membranes of the
first dorsal not black; no vague spots along
body), and the pectoral fin ray count is 27
instead of 23-24. The lateral line count is 51
to 56 instead of 42-48. The snout of B. aetha-
lea is scaled not naked as in B. caudimacu-
lata.

From B. adenensis the higher lateral line count
(46-48 in B. adenensis ), and the presence of
smoky bars on the lower sides rather than 3
or 4 dark blotches (Norman 1939: 69-70) are
diagnostic.

Derivation: aetlialea from the Greek aethalos
meaning smoke.

Bembrops indica McKay, new species

Description: Described from the holotype;
proportions and counts of 5 paratypes given
within parentheses.

Dorsal fin VI, 15 (VI, 15 >; anal 17 (17), pec-
toral 28 (28), pelvic I, 5 (I, 5). Gill rakers 3/15.

Lateral line scales 66 (64-65). Transverse

scales below rayed dorsal fin origin 7 to lateral
line, 6 below.

Head 37.8 (36.4-39.3), depth 11.3, snout 11.4
(10.3-12.1), eye 8.2 (7.8-8.5), interorbital 1.1
(0.95-1.0), depth of caudal peduncle 5.0, pectoral
length 20.2, pelvic length 15.2, caudal length
18.6, second dorsal base 34.0, anal base 34.7.
greatest width of head 14.2, length of maxilla
10.5, all expressed as percent of standard length.

Head flattened dorso-ventrally, widest at
opercles. Mouth wide, with lower jaw projecting
beyond upper jaw. No preorbital spines.
Maxilla extending little beyond anterior margin
of eye; posterior margin truncate with flap of
skin extending almost to middle of eye. Opercular
and subopercular spines present; middle spine
closer to upper opercular spine than to sub-
opercular spine. Opercular flap reaching eighth
lateral line scale. Small spine directed poster-
iorly above origin of opercle. Anterior five
lateral line scales with median raised keel.
Tongue without teeth; tip slightly expanded.
Teeth on vomer and palatines. Fine conical
teeth on both jaws, those on upper jaw extend-
ing outside mouth. Gill rakers 4/14, well de-
veloped with a row of small spines on inner
surface. Gill membranes free from isthmus.
First dorsal spine filiform, reaching second
dorsal ray when spine depressed.

Colour in formalin: Body with about 12

diffuse dark spots along lateral line extending
2-3 scales above and below the line. A series
of about 9 faint dark spots longitudinally above
lateral line, the first 2 of this series on the
anterior part of the lateral line. Dorsal mid-
line with about 12 faint dusky spots, those along
the second dorsal fin situated at the base of the
2nd. 4th, 6th, 8th, 11 -12th and 14th rays. A

Figure 2 . — Bembrops aethalea sp. nov. lateral view.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

43

Figure 3 . — Bembrops indica sp. nov. lateral view.

dark spot present at upper base of caudal fin.
Fins pale to dusky, the membrane between the
first three dorsal spines dark. A small dark spot
at the base of the humeral spines. (One para-
type P 19137 has well defined round spots in
2 to 3 series on sides.) Second dorsal with a
submarginal black band; tip of caudal with a
dark bar.

Holotype: WAM P 19138, 161.5 mm standard
length, collected in 350 metres, 17°17'0"S,
1 19°57’0"E. Station UMPT 6905, December 20,
1969.

Paratypes : WAM P 19137, s.l. 158 mm; WAM
P 19139, s.l. 79.6, Station UMPT 6905; WAM P
19140, s.l. 192 mm; WAM P 19141, s.l. 119 mm.
Station UMPT 6913, 62 metres, 13°43'5"S,
128°38'6"E, December 26, 1969; WAM P 19162,
s.l. 99 mm; WAM P 19163, s.l. 172 mm; Station
UMPT 6910, 370 metres, 13°28'3"S, 123°09'7"E,
December 23, 1969.

From B. filifera Gilbert this new species dif-
fers in having 17 instead of 18 anal rays, a short
snout, shorter maxillary and different coloration
(Gilbert, 1905 : 643-644, pi. 84). From all other
species with an elongate first dorsal spine B.
indica differs in having 64-66 lateral scales. The
Atlantic B. anatirostris Ginsburg is morpholog-
ically similar but has the second dorsal spine
produced and has only one series of lateral spots
on the body (Ginsburg, 1955 : 635-637).

It is noteworthy that two specimens, from
Station UMPT 6913, were taken in shallow depth
<62 metres) whereas the other five all came from
much greater depths, between 350 and 370
metres.

Derivation: indica in relation to the Indian
Ocean.

Chrionema chloro taenia McKay, new species
(Figure 4)

Description : Described from the holotype;
proportions and counts of 7 paratypes given
within parentheses.

Dorsal fin VI, 164 (VI, 16-17), anal 25 (25-
264), pectoral 24, pelvic I, 5. Gill rakers 5/15.
Lateral line scales 79 (80-85 ). Transverse scales
below origin of second dorsal fin 13 above lateral
line, 6£ below.

Head 33.5 (33.2-35.9), depth 13.4, snout 8.8
(8. 3-8.9), eye 8.1 (8. 3-9.9), interorbital space 1.3
(0.84-1.4), length of maxilla 13.4 (12.8-14.4),

snout tip to first dorsal origin 33.5 (33.6-34.9),
greatest width of head 17.7, depth of caudal
peduncle 5.2, length of pectoral 17.5, length of
pelvic 15.6, length of caudal 16.9, length of sec-
ond dorsal base 36.2, length of anal base 43.4,
longest dorsal ray 11.9, longest anal ray 7.7.
length of second dorsal spine 12.0, all expressed
as percent of standard length.

Head slightly flattened dorso-ventrally, widest
at opercles. Mouth wide with the lower jaw pro-
jecting beyond upper jaw anteriorly, but wide
upper jaw extending beyond lower jaw laterally.
No preorbital spines. Most of snout naked. Max-
illa extending to front third of eye; posterior
margin truncate, without a flap of skin. Oper-
cular and subopercular spines present; middle
spine closer to upper opercular spine than to
subopercular spine. Opercular flap reaching
eighth lateral line scale. Low spine present at
commencement of lateral line. Anterior five lat-
eral line scales with median raised keel. Tongue
without teeth, tapered anteriorly with rounded
tip. Teeth on vomer and palatines. Fine conical
teeth on jaws, those of upper jaw extending out-
side mouth. Gill membranes free from isthmus.
Angle of preopercle with three very small points.

Colour in alcohol: Body with about 11 vague
cross-bars variable in width; sides of body with
three large and 4 small alternating dusky areas.
Head dusky with vague, slightly oblique bars on
the opercles. Scales with darker edges forming a
network pattern on cheeks and back.

Premaxillae dark, lower caudal rays dark,
membrane between first and second dorsal spines
slightly darker than remainder of fin, second
dorsal fin hyaline, anal fin with dark margin,
pectoral fins slightly dusky, pelvic fins with inner
rays dark.

Colour in life: Lemon-yellow cross-bars pre-
sent on the body; two large lemon spots on
opercle flap, smaller lemon spots before eyes.

Holotype: WAM P 19136. 194 mm in standard
length. 350 metres, 17° 17' 0" S, 119 c 57' 0" E,
Station UMPT 6905, December 20, 1969.

Journal of the Royal Society of Western Australia. Vol. 54 Part 2. July, 1971

44

Figure 4 . — CKHonema chlorotaenia sp. nov. lateral view.

Paratypes: WAM P 19129. s.l. 187; WAM P
19130, s.l. 187 mm; WAM P 19131, s.l. 175 mm;
WAM P 19132, s.l. 193 mm; WAM P 19133, s.l.
169 mm; WAM P 19134, s.l. 156 mm; WAM P
19135, s.l. 131 mm; Station UMPT 6905, Decem-
ber 20, 1969.

Very similar to Chrionevia chryseres Gilbert,
the only other species in the genus, but differs
in the higher lateral line count (79-85 instead
of 72 or 76), the absence of a well developed
preopercular spine, shorter snout, 25-26 anal
ravs (C. chryseres 24) and the shorter predorsal
length (33.5-34.9 percent instead of 38 percent).

Derivation; chlorotaenia from the Greek
chloros meaning greenish-yellow and taenia
meaning banded.

Liopsaron McKay, new genus
Type species Liopsaron insolitum McKay,
new species.

Body slender, without scales, head naked.
Lateral line absent, no pores on body, a conspic-
uous nerve present below the skin immediately
behind the pectoral fin; this nerve continuing
along the middle of the sides, with branches or-
iginating near the upper pectoral fin origin and
about midway along the pectoral fin length.

Eyes large, dorso-lateral, and separated by a
very narrow interorbital space. Snout short,
without preorbital spines. Upper jaw longer
than lower; mouth almost horizontal, maxillaiy
without a flap. Upper and lower jaws with two
to three rows of small depressable teeth; vomer
with numerous very small teeth; palatines with-
out teeth. Tongue without teeth.

Opercle with a single strong spine projecting
a little beyond the opercular flap; upper origin
of opercle with a conspicuous, small, tube-like
pore Subonercle with a strong spine extending
beyond the level of the opercular spine but
not extending beyond the subopercular flap
which is somewhat, pointed and forms a hori-
zontal floor to the posterior branchial cham-
ber. Ceratohyal without a process. Gill mem-
branes united to isthmus and attached Pos-
teriorly between the pectoral and pelvic fin
origins. Pectoral fin with a fleshy base sepaiate
from the body anteriorly for a short distance
and bound to the body posteriorly by almost
transparent skin. Pelvic fins originating before
pectorals, their bases widely separate. Anal
fin commencing below the third to fourth
dorsal ray. Branchiostegal rays six.

This new genus differs from all other genera
within the family in lacking scales. Scale
pockets are absent and no lateral line pores
were found. The gill membranes are horizont-
ally disposed between the subopercular spine
and the isthmus to which they are joined. The
closest relative is unknown at present.

Derivation: Liopsaron from the Greek lios
meaning smooth and psaron meaning little fish.
Gender masculine.

Liopsaron insolitum McKay, new species
(Figures 5, 5a)

Description: Based on the holotype; counts
and proportions of 2 paratypes given within
parentheses.

Dorsal III, 14 (III, 14), anal 13£ (13-13£),
pectoral rays 26 (26), pelvic I, 5 (I, 5), branched
caudal rays 4 + 4.

Head to tip of opercular spine 28 (28.7, 27.3),
head to tip of subopercular spine 30.1 (31.0,
29.9), horizonal diameter of eye 11.8 (12.6,

11.3), length of snout 4.3 (4.6, 4.6), interorbital
space 1.1 (1.1, 0.7), length of upper jaw 7.5
(7.5, 8.4), snout tip to first dorsal fin origin
30.1 (29.9, 31.8), snout tip to origin of second
dorsal fin 43.0 (43.1, 43.6), snout tip to anal
fin origin 50.5 (52.2, 50.7), snout tip to pectoral
fin origin 30.1 (29.4, 30.1 ), snout tip to pelvic
fin origin 25.8 (24.7, 23.2), length of second
dorsal fin base 47.3 (48.2, 48.7), length of anal
fin base 39.3 (40.2, 39.4), length of pectoral fin
22.6 (23.0, 21.6), length of pelvic fin 22.6 (24.1,

25.8) , length of longest dorsal ray 17.2 (17.2,

29.5) , length of longest anal ray 11.3 (10.7,

11.6) , length of second dorsal spine 9.0 (8.5,

9.8) , greatest width of head 17.2 (16.7, 16.9),

greatest depth of body 14.5 (15.0, 14.8), depth
of caudal peduncle 5.9 (5.8, 6.1), width of

caudal peduncle 2.2 (2.3, 2.1), height of fleshy
pectoral base 8.0 <8.3, 7.6), distance between
tip of opercular spine and tip of subopercular
spine 7.5 (8.0, 8.4); all expressed as percent of
standard length taken from the tip of the
snout to the commencement of the caudal fin.

Head slightly depressed; eyes large and
separated by narrow interorbital space; mouth
protrusible; snout slightly concave anteriorly
but lacking spines; opercular and subopercular
spines diverging posteriorly, both with sharp
tips, opercular spine ending at about level of

Journal of the Royal Society of Western Australia, Vol. 54 Part 2. July, 1971

45

Figure 5. — Liopsaron insolitum sp. nov. lateral view; a. dorsal view of head.

upper pectoral origin, subopercular spine ter-
minating almost at lower pectoral origin; pec-
toral fin rounded posteriorly, a free fleshy base
quite conspicuous; pelvic fin with inner rays
longest, spine much reduced and extending
posteriorly to tip of subopercular spine; caudal
fin almost truncate.

Colour in formalin : Body and head pale,
almost completely faded; some faint dark
blotches above opercle, before spinous dorsal
fin, and one small spot on middle of pectoral
base. Behind tip of pectoral fin a group of 3
somewhat variable almost vertical stripes be-
coming wider at base. A faint blotch present
before caudal fin. All fins hyaline with oc-
casional dorsal rays with a very fine, darker
anterior margin. Eyes black above and silvery
below pupil.

Colour in life: Rather pink with some lemon
yellow spots on body, pectoral base yellowish and
hind border of orbit orange. Much of the
colour faded rapidly in formalin fixative; the
colour in life is believed to be rather attractive.

Holotype: WAM P 19164 standard length 93
mm, total length 115 mm, trawled by Umitaka
Maru in 350 metres, Station UMPT 6905,
17°17'0"S, 119°57'0"E, December 20, 1969.

Paratypes : WAM P 19165, s.l. 87 mm, t.l. 107
mm; WAM P 19166, s.l. 71 mm, t.l. 88 mm;
Station UMPT 6905, data as above.

This new genus and species was taken in
association with Branchiopsaron ozawai, Bern -
brops indica and Chrionema chlorotaenia ; thus
all the genera known from the north west of
Western Australia were taken in a 30 minute,
1 mile trawl. The general construction of the
gear employed is as follows; Lower-net, side-net

and upper-net have meshes of 60 mm, becoming
meshes of 45 mm in the cod-end. The otter
board is 2.54 x 0.9 m in size; the bridle wires
from the board to the towing warp are 20 m
long with a diameter of 16 mm; the wire con-
necting the boards to the net is 5 m long;
the tickler chain is 25 m long and 8 mm in
diameter.

The bottom temperature ai Station UMPT
6905 was 11.4°C.

Derivation: insolitum from the Latin insolitus
meaning uncommon.

Acknowledgements

I am indebted to Dr R. W. George and Dr
G. M. Storr of the Western Australian Museum
for advice and criticism given throughout this
study.

I also wish to thank Miss Jeanne-Marie
Thomasz for the carefully executed figures in
the text.

References

Gilbert, C. H. (1905). — The deep-sea fishes of the
Hawaiian Islands. Bull. U.S. Fish. Comm.
23: 575-713.

Ginsburg, I. (1955). — Fishes of the family Percophididae
from the coasts of eastern United States
and the West Indies, with descriptions of
four new snecies. Proc. U.S. natn. Mus. 104 :
623-639.

Norman, J. R. (1939). — Fishes. In: The John Murray
Expedition 1933-34 Scientific Reports, 7 (1):
1-116.

Okamura, O. and Kishida, S. (1963). — A new genus and
species of the bembroid fish collected from
the Bungo Channel, Japan. Bull. Misaki
mar. Viol. Inst. No. 4: 43-48.

Schultz, L. P. (1960). — Family Trichonotidae. In: Fishes
of the Marshall and Marianas Islands. Bull.
U.S. natn. Mus. no. 202, 2: 273-281.
Whitley, G. P. (1944). — Rare Fishes Aust. Mus. Mag. 8
(8): 254.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

46

7 . — Habeobdella stagni, a new genus and species from South-Western
Australia (Hirudinoidea: Richardsonianidae)'

By Laurence R. Richardson* *

Manuscript received 20 October 1970; accepted 16 March 1971.

Abstract

Leeches from Herdsmans Lake, Lake Monger,
Serpentine (and possibly Greenbushes) are a
new species assigned to a new genus distinct
in having xxv 5/5- or 4/5-annulate, and xxvi
incomplete 3-annulate. A possible second genus
is indicated in leeches from near Bullsbrook
and from the Donnelly River.

Introduction

This paper gives the first account of the
general morphology of an aquatic, jawed,
sanguivorous leech from Western Australia, a
leech with somite xxv essentially 5 -annulate, as
shown by Johansson (1911) who had five speci-
mens from Western Australia, and briefly de-
scribed the colour, pattern, and the annulation
of the posterior somites of the body. Johansson
assigned these leeches to Hirudo australis
Bosisto 1859, a species recognized at that time
and until recently as the only 5-banded aquatic
jawed sanguivore in the Australian Region.
H . australis (Richardson, 1968) had been trans-
ferred to the genus Limnobdella Blanchard 1893.
Johansson did not consider Limnobdella separ-
able from the genus Hirudo. He followed
Moquin-Tandon and Blanchard in accepting a
very wide range of variation in pattern and
colour in the hirudiniform species, and the
1 Hirudo australis * in his 1911 paper can now be
seen as including leeches belonging to three
distinct genera (Richardson, 1971 1 ’).

It has been accepted since Whitman (1886)
that the complete 5 -annulate somite occurs in
hirudiniform leeches only on the nephric somites,
viii to xxiv; rarely on all; commonly on ix to
xxiv with xxv 4 -annulate or on ix to xxiii with
xxiv 4 -annulate and xxv 3-annulate, and with
xxvi 2 -annulate in both these groups.

In terms of this ‘rule’, the novelty shown by
Johansson has been open to doubt as a possible
artefact, the more so since he showed xxvi as
incomplete 3-annulate, and did not consider the
novelty as a sufficient basis for separation of
the leeches from Western Australia from
australis known until then only in eastern
Australia.

There are other divergences from the ‘rule’
(Richardson, 1969; 1971 a ). These can be

recognized now as indications for distinct
systematic status.

1 Prepared during the course of studies on the Austra-
lian leeches assisted by a grant from the Nuffield
Foundation.

* 4 Bacon St., Grafton, New South Wales 2460.

The annulation described by Johansson is
confirmed in specimens among 23 aquatic jawed
sanguivores in collections from 11 localities in
Western Australia, housed in the Western Aus-
tralian Museum. The collections are limited.
The specimens vary in condition. From among
them, it is possible to provide a necessary new
genus based on a type from a convenient locality.
It is hoped that studies will be undertaken on
ample material to demonstrate the nature of the
variation in annulation.

Habeobdella Richardson, gen. nov.

Derivation of generic name: habere, to be in
possession of; bdella, a leech. Gender:
feminine.

Monostichodont; ix to xxiv complete 5-annu-
late (total 16); xxv more or less 5 -annulate
above and below; xxvi incomplete 3-annulate;
somital sense organs small, circular, obscure;
jaws small; teeth small, about 58 to 60; no sali-
vary gland papillae; dorsal salivary glands,
compact masses with right and left columns
of aggregated ducts; radial muscles an obvious
extrinsic system: mouth and lumen of pharynx
narrow, the lumen tubular, tapering; pharynx
with 6 internal muscular ridges joining as
dorsomedian and ventrolateral pairs to enter
the corresponding jaws, none ending inde-
pendently between the bases of the jaws:
pharynx terminating in ix; crop with a short
compartment in ix, x to xviii, the compart-
ments each with anterior small and posterior
larger pairs of caeca, the anterior in xix form-
ing the postcaeca extending to xxvi; genital
pores xi and xii bo/be; testes normally 10 pairs;
epididymis in the posterior half of xii and in
xiii, posterior to the simple fusiform ejaculatory
bulb in the contiguous halves of xi and xii;
median regions bimyomeric, mesomorphic; penis
sheath cylindrical, reflected in xii; oviducts,
short; common oviduct, longer; vagina fusi-
form, on recurrent limb of primary loop;
vaginal caecum enlarged, of the width of the
vagina and as long as wide; vaginal duct
longer than the vagina.

Size, medium. Pattern, a medium dorsal
and two pairs of uninterrupted median longi-
tudinal dark bands. Aquatic. Sanguivorous
South Western Australia.

Type species: Habeobdella stagni sp. nov.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

47

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

48

Habeobdella stagni Richardson, sp. nov.

(Figure 1.)

Type specimen: Western Australian Museum,
Coll, No. 49-69, taken in mud, south end
of Herdsmans Lake, Perth, W.A., 5 Nov. 1965.
Length 43.0 mm, dissected, xx/xxi to xxvi
opened along left side, dorsomedian and
right ventrolateral jaws mounted separ-
ately. The type has xxv complete 5-annu-
late.

With this, one paratype, 42.0 mm long,
xxv with (ba+bfi) on dorsum, the furrow
bs/b* marginal and complete across the
venter; dissected.

The description is taken from the type.
General form

Slightly contracted, preserved, a leech of
medium size; elongate, depressed, the dorsum
low convex, the venter nearly flat, the margins
broadly rounded, the posterior sucker large.
The velum thick with rounded margins; behind
this, the margins subparallel, diverging very
gradually from iv/v to x br,/b«, slightly wider
as the clitellum to xiii a 2 , then almost sub-
parallel diverging only slightly to the maximum
width at about xx to xxii; converging behind
this in the postnephric region to form the base,
which is about half the width of the sucker,
and the sucker slightly wider than the maxi-
mum width of the body.

The total length 43.0 mm; 2.0 mm wide at
iii/iv; 3.0 mm wide at v/vi; the width 4.5 mm
and the depth 2.0 mm at the male pore,
9.0 mm from the tip of the velum; maximum
width 5.0 mm and depth 3.5 mm at 30 mm
from the tip of the velum; the basis of the
posterior sucker 3.0 mm and the sucker 6.0 mm
in diameter.

Colour

Preserved: generally darkish grey; the dorsum
with five black longitudinal bands, a narrow
medial and wider inner and outsr paired bands,
the inner pair subdued along most of their
length by much pale greyish clouding to such a
degree and in such a manner as to appear as
two narrow separate longitudinal bands; the
dark bands separated by narrow pale yellow
stripes; lateral to the outer paired bands, pale,
faintly yellowish to brownish grey wide marginal
bands sharply marked off in the submarginal
field from the light grey immaculate venter.
The sucker with a large wide black patch con-
taining a central pale yellow patch, otherwise,
laterally and anteriorly, the sucker pale, of the
colour of the venter.

The colour in life possibly brownish black,
divided by bright yellow, even golden, stripes;
venter, ashen grey.

Pattern Figure 1 A,G,F.

The first four pairs of eyes marginal on a
wide ocular patch in ii to vi a,, divided by the
narrow inner pair of light stripes which define
the median band from ii/iii, just behind the
1st pair of eyes, to the posterior border of xxvii.
The median band generally about two thirds of
the width of the median field and narrower
than the inner pair of light stripes on either
side of it which extend along the paramedian
lines and include sense organs of this series.
The inner pair of dark bands is defined between
the inner and outer paired stripes between vi
ai/a> and xxv/xxvi, posteriorly joining a dark
patch extending across the paramedian and
intermediate fields of xxvi and xxvii, divided
by the inner paired stripes and median band.
The inner paired bands are wide, occupy about
the medial two thirds of the paramedian field,
widening as this field widens along the body,
narrowing in and posterior to xxiv; clouded
liberally from the middle of viii to xxiii with
paler grey to brownish grey, appearing then
more as black margins to a pale stripe as wide
or wider than the combined widths of the
margins; but the whole constitutes one dark
band.

The outer paired light stripes are lateral in
the paramedian field from vi ai/a 2 to xxvi aia-i/a*
wider than the inner paired stripes, and gen-
erally uniform in width along the greater part
of the body.

The outer paired dark bands separate from the
ocular band at vi ai/a 2 and extend to xxvi
aia 2 /a. { , pass immediately lateral to the 5th pair
of eyes and the intermediate organs of vii, but
behind this, the sense organs of the intermediate
series lie on or just within the inner margin of
these bands which also occupy the greater part
of the intermediate field, being narrowly separ-
ated from the supramarginal line of sense organs
which lie in the marginal band extending across
the supramarginal and submarginal fields as pale
bands continuous from the velum to xxvii and
onto the sucker, and are sharply distinct from
the paler venter. The outer pair of dark bands
show minor clouding behind viii a 2 .

The right and left dark patches on xxvii are
continuous with the dark patch on the sucker
which occupies the inner half or just less of
the intermediate fields, the paramedian and
median fields except for the central light ovoid

Figure 1 . — Habeobdella stagni Richardson, gen. et sp. nov. A. B.— Somital annulation and pattern, dorsal aspect—
A, somites i to x; B, somites xxiv to xxvii and sucker. C.— Dorsomedian jaw and dental ridge (arrow indicates
medial end of ridge). D.— Anterior half of pharynx opened along the midventral line to show internal mus-
cular ridges; jaws. E.— Crop and caecation. somites xviii and xix; Intestine; rectum. F.— Ventral aspect, somites
xi and xii, showing genital pores, nephropores, etc. G — Anterior region of male paired ducts, male median
region, and female reproductive system (median aspect of the organs on the male paired ducts shown; the dorsal
aspects lateral in the figure.)

All figures prepared from the type. Somites and somital ganglia indicated by Roman numerals. Scales in
millimetres.

Abbreviations: ce., caecum; c.od., common oviduct; ej.b., ejaculatory bulb; ej.d., ejaculatory duct; epd.,
epididymis; nep., nephropore; ov., ovary; pe.s., penis sheath; pr., prostate; te., testis; va., vagina; va.d., vaginal
duct; v.d., vas deferens.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

49

patch which is separated more narrowly from
the posterior edge of xxvii than from the
posterior edge of the sucker.

Annulation Figure 1A, B.

Somital sense organs are small, rounded, de-
tectable often with difficulty on the dorsum
excepting the intermediates, not always as con-
tinuous series, and much smaller and quite
obscure on the venter. Secondary sensillae are
most difficult to detect. The nephropores are
obvious in the postclitellar region where most
are raised on low papillae; elsewhere concealed.
Interannular and intersomital furrows essen-
tially equivalent, the annuli not arranged in
obvious couplets or triplets, and somital limits
without definition along most of the body. Gen-
erally the annuli are richly divided into longi-
tudinal rectangles by fine lines which are lack-
ing on well-extended somites, and seem to have
no morphological value.

The velum consists of i, ii and iii, carries
the 1st and 2nd pairs of eyes, shows no defined
furrows and the margin is entire, without in-
cisions. Somite iv is marked off anteriorly by
iii/iv which extends across the median and
intermediate fields; a longer furrow dividing
iv into aia 2 /a s , with aia 2 , including the 3rd
pair of eyes and first detectable paramedian
sense organs >a 3 ; the furrow iv/v not quite
reaching the margin of the sucker so that the
dorsolateral lobe of the margin of the sucker
is poorly defined; v 2-annulate above, the 4th
pair of eyes in a,a 2 a :i , the furrow a^/a*
extending into the submarginal field so that
aia 2 forms the lateral portion of the margin
of the sucker and uniannulate v, the ventral
portion: vi 3 -annulate above, ai<a 2 <a 3 , the 5th
pair of eyes in a 2 , the furrow a,/a 2 extending
to the ventral intermediate line, and then vi
2-annulate below with aia- 2 >a :i in the para-
median and median fields; vii 3 -annulate above

and below, ai<a 2 <a ;} (- viii at) ( a« with

a very fine furrow extending between the
supramarginal sense organs dividing a :{ appar-
ently into b.-,<b <; which would suggest it
may not be morphological); viii 4-annulate,
ai>a 2 >bo>b, 5 the first nephropores close

to the posterior border and just medial
to the ventral intermediate lines on

a, ; ix to xxiv complete 5-annulate (total 16 >;
from ix to xxiii the somites variously contracted
and the relative lengths of the annuli cannot
be determined with full confidence; in general
it would seem that the somital annulation for
ix to xviii is bi=b 2 <a 2 b 5 >b (i ; for xix to
xxi, bi b ; ?<a 2 <bs>br,; for xxii to xxiv,

b. bj>a 2 >b., b t; ; the last nephropore on xxiv
b>; xxv complete 5-annulate, bi<b 2 =a 2 >b 5 >b«,
the furrow b*'b fi strongly defined and deep on
both the dorsum and venter, with a 2 . b.-, and b 6
reduced in length on the venter where
b, b 2 >a o b 3 >b«; xxvi 2-annulate above,
aia$>a. s , the somital sense organs posterior in
a,a 2 , and at the margin a shallow w r ide groove
in a a* which deepens into a definite furrow
across the venter where although ata 2 is re-
duced in length it is weakly but definitely

divided into a t and a 2 , and a 2 is the last
annulus complete across the venter, a 3 extend-
ing onto the dorsum of the sucker as also xxvii
which is uniannulate above. The anus at the
posterior border of xxvii. The dorsum of the
sucker shows 4 paramedian sense organs in
each line, and three in each of the intermediate
and supramarginal series; some 7 or 8 concentric
rows of tesselations show on the posterior half
of the dorsum.

Alimentary tract Figure 1C, D, E.

The jaws are small, tall, compressed, the
dorsomedian in profile at the median end,
about 0.5 mm high and slightly narrower at
the base: the dorsomedian housed in an open
groove, the ventrolaterals in pits with such
poorly defined margins as to be nonmorphologi-
cal; the dental margin long, so highly convex
as to be almost semicircular; the teeth about
60 on the dorsomedian, the largest about 0.05
mm high, the row reducing gradually and the
teeth in the middle of the row about half as
tall as those at the median end, the row pro-
gressively reducing to the last few teeth which
are low, rounded, almost granular; about 56
teeth on the ventrolateral.

The pharynx is relatively thin -walled with
3 pairs of internal muscular ridges, each pair
joining to enter the base of a jaw, none ending
independently between the bases of the jaws;
mouth narrow 7 , little wider than the base of
the dorsomedian jaw; lumen of the pharynx nar-
row, tubular, tapering; radial muscles sparse,
but an obvious extrinsic system extending back
into x; dorsal salivary glands compact right
and left masses extending back into x, each
drained by a poorly defined column of aggre-
gated ducts, the columns continuing indepen-
dently into the dorsomedian jaw.

The pharynx terminates in ix a 2 , followed by
a short acaecate compartment in ix; the crop
in x to xviii fully compartmented, each com-
partment with a pair of simple anterior and
a pair of simple posterior caeca, the tw T o pairs
nearly equivalent in x and xi: then the pos-
terior pair progressively longer, and from xv
back extending in the paramedian chamber
posteriorly into the following somite where
they are lateral to the anterior caeca, but all
as seen w T ere simple and not folded; the post-
caeca originate from the lateral aspect of the
anterior end of the compartment in xix, behind
this the compartment tubular, with the lining
epithelium in longitudinal rugae as elsewhere
in the crop, tapering briefly to enter the intes-
tine sub terminally in the anterior annuli of
xx ; the postcaeca extend to xxv /xxvi.

The intestinal epithelium is raised in trans-
verse rugae. The intestine commences with a
caecate tubular segment in xx, another simple
segment in xxi, followed by a tapering tubular
length entering terminally into the rectum which
tapers to the anus.

Reproductive system Figure 1G.

The relationship of structures on the an-
terior portion of the male paired ducts is
richardsonianoid; the median regions bimyo-
meric. mesomorphic; the vagina caecate formed

Journal of the Royal Society of Western Australia, Vol. 54 Part 2. July, 1971

50

essentially on the recurrent limb with a vaginal
duct on the procurrent limb. The specimen is
adult, but does not appear to be in a condition
of sexual maturity.

The genital pores at xi b»/b 6 and xii bs/b«;
the penis-sheath elongate, cylindroid.

The first testes are at xiii/xiv; the last in the
posterior portion of xxii; behind xix, the testes
are in the paramedian longitudinal chamber of
the body -cavity; anterior to this, in the median
chamber, all connecting by short vasa efferentia
to the vas deferens in the paramedian chamber
which reduces in diameter in the posterior annuli
of xiii before expanding into the initial narrowly
tubular portion of the epididymis which is thin-
walled, tortuous and much folded on itself and
becomes more widely tubular before joining the
ejaculatory bulb; the whole epididymis in the
contiguous annuli of xii and xiii and not readily
separable into two masses. The bulb is mus-
cular, tapering elongate fusiform, connected by
a short ejaculatory duct to the male atrium
situated above g.xi. The atrium continues in
the strongly muscular penis sheath, slightly
folded on itself, and reflected at the level of g.
xii as two nearly equal limbs.

The simple saccular ovaries are posterior in
xii, connect by short oviducts, neither longer
than the ovary, to a distinct atrium from which
the thin-walled common oviduct of large lumen
extends tortuously without close relationship to
the vaginal duct, to join the ventral face of the
vagina more remote than usual from the caecate
end, about two fifths along the body of the
vagina so that the caecum is quite long, the
length of the width of the vagina, and the
caecum unusually large; the common oviduct
opens into the vagina on the tip of a low papilla
at the level of the point of attachment; an
epithelium with longitudinal rugae continues
from the vagina into the caecum.

The caecum and vagina are in the posterior
annuli of xii and in somite xiii. The muscular-
ized body of the vagina is stoutly tubular, com-
pletes the recurrent limb of the primary loop
and tapers abruptly on the initial portion of the
procurrent limb into the strongly muscular
vaginal duct which has a reduced lumen, is
distinctly longer than the body of the vagina,
and is folded closely on itself on both sides of
the ventral nerve cord.

The prostate glands form an elongate taper-
ing subovoid cap on the atrium and extending
briefly along the penis sheath. The albumin
glands invest the atrium and the length of the
common oviduct.

Paratype

Similar in colour and anteriorly in pattern to
the type; posteriorly as in the type excepting
that the narrow paramedian light stripes extend
onto the dorsum of the sucker, but only so much
as to closely encircle the anus, and there is a
small light patch central on the posterior margin
of the sucker. The paramedian bands along the
body fill half or slightly more of the paramedian
fields, each with a wide dark inner margin about
half the width of the whole band, and the rest
clouded so that there is no dark outer margin.

Annulation differs from the type: iv aia«/a»,
the first furrow extends only to the intermediate
line, as also iv/v; the clitellar annuli, xbs to
xiiia 2 , each with a distinct deep furrow between
the submarginals so that these annuli are all
strongly subdivided on the ventral surface; xxv
4 -annulate above, with bi - b 2 <a 2 <a 3 , but the
furrow br,/bs commences in the marginal fields
and continues across the venter where b r , and
b « are distinctly represented and bi ~ b 2 >a 2
b.-,>b H ; xx vi as in the type, aia 2 on the dorsum
but divided on the venter into ai and a 2 , with
a 2 the last annulus complete on the venter; xxvi
a* and xxvii continue onto the dorsum of the
sucker.

Alimentary tract as in the type.

Clitellar glands well developed. Ejaculatory
bulbs shorter, more stoutly fusiform; penis
sheath with the recurrent and procurrent limbs
of the same length; oviducts, both very short;
vaginal caecum at least as long as wide, of the
width of the vagina and about a quarter of its
length.

Other material

Leeches identifiable as H. stagni were present
in the following collections:

1. W.A.M. 9034. Hirudo australis Bosisto.
Serpentine, W.A.B., K.A.B., & I.J.D, No
other data.

Four specimens, 23.0 to 33,5 mm long;
faded; pattern as in type; xxv bi<b 2 <a 2

b 5 >bn above and below, with b« the last
annulus across the venter; xxvi a,a 2 /a.-t
above, ai/a 2 in the submarginal field;
vaginal caecum as in type.

2. W.A.M. 52-69. Lake Monger, Oct. 1952.

Five specimens, 14.0 to 33.0 mm. Stripes,
yellow; pattern and annulation as in type;
pattern fully established in the 14.0 mm
specimen; vaginal caecum as in type. One
specimen retained, deposited the Austra-
lian Museum, Coll. No. W 4269.

3. W.A.M. 50-69. Greenbushes swimming
pool (in abundance), coll. H. Williams. 26
Dec., 1962.

Two specimens, 32.0 and 49.0 mm. Stripes
yellow; pattern and annulation as in type
excepting xxv incomplete 5-armulats below
with a 2 the last annulus across the venter;
enlarged vaginal caecum, of the width of
the vagina, but not as long as wide. Note:
with a full knowledge of variation in
stagni, this might prove distinct.

Discussion

The genus Habeobdella conforms in morpho-
logy to the Richardsonianidae of the Austra-
lian Region.

Lacking salivary gland papillae on the jaws,
with 16 complete 5 -annulate somites (ix to
xxiv), bimyomeric mesomorphic median regions
on the reproductive systems, simple fusiform
ejaculatory bulbs and vagina, a vaginal duct,
and a pattern of 5 uninterrupted dark longi-
tudinal dorsal bands, Habeobdella is similar

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

51

(Richardson, 1970) to the genus Richardsoni -
anus Soos 1968 based on Hirudo australis
Bosisto 1859.

Habeobdella differs from Richardsonianus,
as also from the other 16 5-annulate genera
so far known in Australia, in the essentially
5-annulate condition of xxv, the 3 -annulate
condition of xxvi and the very large vaginal
caecum which has not been seen in other Aus-
tralian leeches.

In the collections there is another leech pos-
sibly representative of a second genus; but the
material is not adequate for a complete descrip-
tion.

A specimen from n:ar Bullsbrook (W.A.M.
56-69), 65.0 mm long, and two specimens from
the Donnelly River (W.A.M. 54-69) 16.0

and 28.0 mm long, differ from H. stagni in
having elongate almost subcylindrical ejacu-
latory bulbs resembling the bulbs in bassianob-
dellid leeches; the vaginal caecum small,
short, the vagina simple fusiform with a
vaginal duct.

Both have xxv 5-annulate above, with a 2 the
last annulus across the venter in the Bulls-
brook specimen, and b 5 the last such annulus
in the Donnelly River specimens. Somite xxvi
is 2 -annulate above in all three.

There are about 50 teeth on the dorsomedian
jaw of the Eullsbrook specimen. These are
distinctly small, the tallest at the median end
being 0.03 mm high, and the row diminishes so
gradually as to appear almost uniform in height
along the greater part of its length.

They differ from H. stagni in pattern. The
larger of the Donnelly River leeches lacks the
outer paired light stripes lateral in the para-
median fields; the smaller has these stripes,
but they are interrupted and extend only be-
tween x a 2 (right) and xi a 2 (left) to xxii/xxiii.
In both, a dark marginal band commences in
the lateral half of the intermediate field and
extends to just include the marginal line of
sense organs, where it is then sharply separated
from the pale venter which has a few spaced
large maculae.

In the Bullsbrook leech there are poorly de-
fined narrow outer paired light stripes which
extend along the line of the intermediate somi-
tal sense organs. There are dark marginal
bands which are continuous with the immacu-
late venter.

The differences in the topography of pattern
in the Bullsbrook and Donnelly River leeches
are indicative of possible distinct species
(Richardson, 1971 b ) ; but as I cannot de-
termine the relative lengths of the annuli in
the midnephric group of somites, the matter
cannot be taken further on these specimens.

Acknowledgements

I am indebted to the Director of the Western
Australian Museum and to Dr. R. W. George
for the privilege of examining leeches in the
collections of the Museum; to Miss E. Pope, the
Australian Museum, Sydney, for assistence in
many ways; to Professor Marvin C. Meyer, the
University of Maine, for assistance in the mat-
ter of difficult literature; the Librarian, the
University of New England, for help with other
literature; and to the Science and Industry
Endowment Fund for the loan of microscopic
equipment.

References

Johansson, L. (1911). — Hirudinea. in: Fauna Siidwest-
Australiens. Bd. iii. L. 12: 409-431.
Richardson, L. R. (1963).— An annotated list of Austra-
lian leeches. Proc. Linn. Soc. N.S.W. 92 (3):
227-245

(1969). — A contribution to the systematics

of the hirudinid leeches, with description
of new families, genera and species. Acta
Zool. Hung. 15 (1-2): 97-149.

(1970 ). — Bassianobdella victoriae gen. et

sp. nov. ( Hirudinoidea : Richardsonianidae) ,
Mem. Nat. Mus. Vic. 31: 41-50.

(1971 a ). — A new genus and species of

Sudan leech formerly confused with Lim-
natis nilotica (Hirudinidae s.l.: Hirudinea).
Bull. Br. Mus. nat. Hist. (Zool.) 21, 7.

( 1971 b ) . — Bassianobdella ingrami sp. nov.

from Tasmania (Hirudinoidea: Richard-

sonianidae). Pap. Proc. Roy. Soc. Tasm. 105
Whitman, C. O. (1886). — The leeches of Japan. Quart.
J. Micr. Sci. (N.S.) 26; 317-416.

Journal of the Royal Society of Western Australia. Vol. 54 Part 2, July, 1971

52

8. — Remains of Potorous platyops (Marsupialia, Macropodidae) and other
mammals from Bremer Bay, Western Australia

by W. H. Butler f and D. Merrilees*

Manuscript received 14 December 1970; accepted 16 March 1970.

Abstract

Mammal remains, probably derived from
disintegrated pellets regurgitated by an owl, are
described from sand dunes at Bremer Bay.
Murid bones from this sample have been dated
at 1190 ± 80 radiocarbon years B.P. (GaK-2887).

The sample includes remains of the small rat
kangaroo Potorous platyops, and other remains
of this little known species are discussed. A
radiocarbon date of 620 ± 90 years B.P. (GaK-
2888) on dead wood from the site, taken in
conjunction with the bone date and with the
present condition of the site suggests that
devegetation of the area, presumably by wind
blown sand, has extended over more than a
thousand years.

Introduction

In March 1970, W. H. Butler collected a small
sample of bones from what appeared to be the
dispersed remains of owl pellets at Bremer Bay,
and presented them to the Western Australian
Museum. Remains of two little known marsu-
pials, Potorous platyops and Antechinus apicalis,
were recognized in this sample, and both authors
visited the site later in the same month in
order to make further collections and field ob-
servations.

We follow the nomenclature of Ride (1970),
including his use of Potorous tridactylus for the
populations formerly known as P. gilberti. The
catalogue numbers quoted represent specimens
in the Western Australian Museum collection.

The Bremer Bay site

About 44 km north east of the old telegraph
station at Bremer Bay, the channel of the
Hunter River is blocked by sand drift about 1
km inland from the coast. Extensive dunes of
mobile sand lie south west of this blocked river
channel, and these mobile dunes at present ap-
pear to be encroaching on low vegetated dunes
west of the channel of the Hunter River. About
central in the mobile dunes, bones lay unevenly
scattered in an area about 200 m long in a north
to south direction and about 250 m long east
to west. The site is at 239753 on the Bremer
Bay sheet (SI 50-12) of the Australia 1: 250,000
series of maps, lat. 34°22'S, long. 119°24'E.

Bones representing more than 650 animals
were recovered from the 200 x 250 m zone,
lying on bare sandy surfaces. Only about 10
animals were represented by bones collected on
similar surfaces of the remainder of the mobile
dune area, about 1 km x 1 km in extent.

tWanneroo Wildflower Nursery, Wanneroo, Western
* Western Australian Museum, Perth, Western Australia.

Within the 200 x 250 m zone relatively rich
in bone, local concentrations were evident, five
such local concentrations accounting for more
than 600 of the animals represented. Our col-
lection from the principal one of these local
concentrations, about 5 x 20 m in extent, repre-
sented nearly 500 animals. We did not recover
every piece of bone exposed at the surface of
the 200 x 250 m area, and our recovery rate
varied from place to place, being higher in the
local concentrations. We estimate that more
than 700 but less than 1,000 animals of various
sizes were represented by bony remains within
the 200 x 250 m bone rich zone.

Remains of dead trees and shrubs were repre-
sented at all the sites of bone concentration
mentioned above, some of them upright in the
position of growth, but more lying scattered on
the sandy surface. The largest of these plant
remains is shown in Fig. 1. This represents a
fallen tree in which roots and branches were
clearly distinguishable, showing that the tree
extended at least 7 m above the ground. A
comparison was made of wood from this fallen
tree with wood specimens from peppermint and
eucalypt trees growing nearby. This comparison,
kindly made for us by Mr H. D. Ingle of the
C.S.I.R.O. Division of Forest Products, Mel-
bourne, showed that the fallen tree was pepper-
mint ( Agonis cf. A. flexuosa ) rather than
eucalypt.

Fig. 1 also shows (top left) a tree stump in the
growth position: not more than 1 m thickness
of sand or soil has been removed by deflation
from round the roots of this tree. The principal
bone concentration, representing nearly 500 ani-
mals, lay to the left of the large fallen tree
shown in Fig. 1.

Many bones and most tree trunks showed signs
of sand blasting or corrosion, but most bones,
especially of the smaller mammals (of about the
size of a rabbit or less), were unbroken, save
that skulls usually had come apart at the
sutures.

Thickets of swamp yate (Eucalyptus Occident -
alis), peppermint (Agonis flexuosa) and other
small trees occupied the banks of the Hunter
River, but the vegetation otherwise consisted
primarily of shrubby plants.

Weather data are available for the nearby
township of Bremer Bay, which has an average
annual rainfall of about 620 mm, with a marked
maximum in winter (June average 91 mm, Jan-
uary average 14 mm). The mean minimum tem-
perature for July, the coldest month of the year

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

53

Figure 1. — Fallen peppermint tree near principal local accumulation of bone, Bremer Bay, The bow saw is
propped against a branch of this tree, partly buried by sand; the roots of the tree are in the foreground.
The tree has fallen uphill away from the sea, presumably in the direction of the strongest winds; it lived
over 600 years ago. Note stump in growth position on sky line, top left.

is about 8°C, and the mean maximum temper-
ature for February, the hottest month, is about
24 c C (Director of Meteorology 1962). Winds are
frequent and strong, predominantly from the
south west, but often with an easterly compon-
ent in summer.

Animals represented

Most of the animals represented in our bone
collection — about 540 in a total of about 650 —
were murids. A few other eutherian species,
dasyurid, peramelid and phalangeroid marsup-
ials, reptiles, amphibians and fish were repre-
sented by bones, while shells of the land snail
Bothriembryon were prominent among inverte-
brate remains.

The following mammal species were present:
Murids

Rattus fuscipes — e.g. 70.3.40

Pseudomys shortridgei — e.g. 70.4.61

Pseudomys albocinereus — e.g. 70.4.114

Notomys cf. N . mitchellii — e.g. 70.4.10

Other eutherians

Bat, not specifically identified —

70.4.48

Oryctolagus cuniculus

— e.g.

70.4.43

Ovis aries

—

70.4.155

Canis familiaris

—

70.3.59

Vulpes vulpes

—

70.4.167

Marsupials

Dasyurus geoffroii

—e.g.

70.3.58

Antechinus apicalis

—e.g.

70.4.98

Antechinus flavipes

—e.g.

70.4.103

Sminthopsis murina

—e.g.

70.4.106

Isoodon obesulus

—e.g.

70.4.91

Peravieles cf. P. bougainville

—e.g.

70.3.15

Pseudocheirus peregrinus

—e.g.

70.3.29

Potorous platyops

—e.g.

70.4.66

Bettongia penicillata

—e.g.

70.4.1

Macropus eugenii

—e.g.

70.3.61

Macropus irnia

— e.g.

70.3.28

Macropus (probably

—e.g.

70.3.60

fuliginosus)

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

54

The whole collection from this locality is in-
cluded under the catalogue numbers 70.3.12 to
70.3.21, 70.3.28 to 70.3.70 and 70.4.1 to 70.4.197.

Potorous platyops

According to Glauert (1950), one or more
specimens of this species were obtained from
a locality not recorded in the south west of
Western Australia by L. Preiss about 1839, but
the name Hypsiprymnus platyops was conferred
by Gould (1844) on a specimen obtained by
John Gilbert probably in the Goomalling district
(Calaby 1954).

According to Thomas (1888), Gilbert also ob-
tained a specimen in the Albany district, and
according to Glauert (1950), G. Masters obtained
four specimens in the King George’s Sound to
Pallinup (Salt) River district between 1866 and
1869. Five specimens from unknown localities
in Western Australia reached the National
Museum of Victoria in 1874 and 1875 (Ride
1970). Glauert (1933) recorded a specimen
from the Margaret River district in the extreme
south west of Western Australia, but this record
is believed to be erroneous (Ride 1970), and
Glauert did not repeat it in his later paper
(Glauert 1950).

Figure 2.— Occurrences of Potorous platyops in geologically Recent (including historic) time.

X Historic records
• Prehistoric

(Few animals have been taken in historic time, and for many of these, locality records are vague. Thus his-
toric localities other than those shown may be involved. The prehistoric locality on North West Cape is queried
because it is based only on one molar tooth, and that near Naracoorte, South Australia, because it may repre-
sent a time earlier than Recent.)

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

55

There are no specimens of Potorous platyops
in the Western Australian Museum collection of
modern mammals, but the collection of fossil
mammals contains fragmentary remains from
Bremer Bay ( discussed herein), and from several
caves in the Moore River-Dongara coastal region
(61.7.32, 62.1.15, 64.10.35, 68.2.25-26 from Hast-
ings Cave, 65.12.422-423, 70.6.66, 70.6.77 from
Wedge's Cave, 69.5.12 from Smithy’s Cave, and
70.4.203 from Weelawadji Cave). A single lower
molar tooth (68.7.102) from a cave in North
West Cape also appears to represent P. platyops;
this record represents a great extension of the
known range of the species, and because it is
based only on an isolated tooth, is queried on
the distribution map (Fig. 2). The geological age
of these specimens is not known, but all come
from surface litter or shallow excavations, and
are likely to be of late Quaternary if not late
Recent age.

Lundelius (1957. 1960, 1963) records fossil
specimens, probably of Recent age, in Hastings
and Wedge’s Caves in the Moore River-Dongara
legion, and also from Webbs Cave in the south
central Nullarbor region. Neither the South
Australian Museum (N.S Pledge, personal
communication) nor the Western Australian
Museum collections of fossil mammals contains
any specimen of P. platyops from the Nullarbor
legion. However, small numbers of P. platyops
specimens have been found in cave deposits
near Madura, Eucla and Koonalda, southern
Nullarbor region, excavated by the Australian
Institute of Aboriginal Studies Nullarbor
Plains Archaeological Survey; all appear to be
of late Recent age (P. Thompson, personal
communication > .

Finlayson (1938) based a separate species
Potorous morgani on bony remains of two
animals from a cave in the south west of
Kangaroo Island. South Australia, This species
is now generally included in Potorous platyops
<e.g. Ride 1970). Finlayson’s specimens have
been examined by one of us (D.M.) and found
closely to resemble the Western Australian
Museum fossil specimens listed above, ascribed
to P. platyops. Other specimens, representing
up to seven individuals, have been found in
surface litter in a cave near the type locality
of P. morgani (P. F. Aitken and C. R. Tidemann,
personal communications), and Finlayson
(1959) records the finding of another specimen
in the south east of Kangaroo Island.

Hale and Tindale (1930) record Potorous
remains from three levels in an archaeological
excavation at Devon Downs, on the bank of the
lower Murray River, and Finlayson (1959) re-
cords these specimens as P. morgani. In
archaeological excavations in Shelter 2 at the
nearby Fromm’s Landing site, two levels yielded
remains of P. morgani (Wakefield, in Mulvaney,
Lawton and Twidale 1964). Remains of
several individuals attributed to P. morgani
have been found recently m an excavation in
Victoria Cave in the Naracoorte district, South
Australia (M. J Smith and R. T. Wells, personal
communication).

Thus, the species appears to have been wide
ranging but not abundant. There appears to
have been no record of it alive for nearly a
century (Ride 1970), and it may be extinct.
The Bremer Bay specimens (70.3.21, 70.3.43-46
70.4.4-5, 70.4.11, 70.4.19, 70.4.55, 70.4.66-88!

70.4.134 and 70.4.177) are fragmentary and
represent only a few animals (probably about
five), but even so are a welcome addition to the
few 7 known remains of Potorous platyops.

The Western Australian specimens cited
above, which include tooth-bearing fragments
of skulls and mandibles and some post-cranial
elements, show that in these respects Potorous
platyops closely resembled Gilbert’s potoroo
(Potorous tridactylus from the extreme south
west of Western Australia) except in being
considerably smaller. Thus Potorous platyops
femur 70.4.68 has a length of about 55 mm,
whereas P. tridactylus femur 65.6.73, of com-
parable stage of growth, is 68 mm long. For
comparison, rabbit femur M 6935 is 78 mm
long. The length of the lower cheek tooth row
in adult P. platyops is about 15 mm (14.3 mm
in 70.4.134, 15.3 mm in 70.4.66) wdiereas the
lower cheek tooth row is about 21 mm long
(20.5 mm in 65.9.49) in adult P. tridactylus.
The length of the lower permanent premolar
in P. platyops ranges from 3.5 to 4.1 mm in 9
measurable specimens ( including one of
P. morgani from Kangaroo Island) and the
W’idth from 1.4 to 2.1 mm in the same specimens;
the lower permanent premolar in P, tridactylus
specimen 65.9.49 is 5.2 x 2.6 mm.

Interpretation of the site

The preponderance of murids, the scarcity
of animals larger than rabbit size, the pre-
ponderance of mammals over other vertebrates,
the predominantly unbroken character of indi-
vidual bones, and the occurrence of local con-
centrations of bone all point to a large night-
hunting predatory bird as being responsible for
the bone concentrations. The Barking Owd
iNinox connivens ) or the Masked Ow T l (Tyto
novaehollandiae ) appear to be the species most
likely involved. We suggest that each of the
local bone concentrations described above repre-
sents the feeding place of a Barking or Masked
Owl.

These owls normally kill their prey at first
attack, transport the body to a feeding site,
and either swallow 7 very small animals whole
or tear off lumps of meat and open the body
cavity and then swallow 7 larger mammals (of
about the size of a rat). The carcases remain
in the modified upper portion of the alimentary
canal during digestion of soft tissues. The re-
maining hair, bone and other indigestible tis-
sue from each meal is w r orked by muscular
contractions into a compact, coherent pellet
which is then regurgitated. Breakdown of such
pellets and removal of hair by insect and other
scavengers, and physical weathering, would then
leave just such bone concentrations as we
found.

Journal of the Royal Society of Western Australia. Vol. 54 Part 2, July, 1971

56

An owl pellet origin satisfactorily accounts
lor almost all the smaller animals, not only
mammals, but also the very small numbers
(about 25) of small birds, lizards and frogs
represented. However, such an origin for

animals as large as a dog or sheep, or as well
armed as an adult Dasyurus , is most improb-
able, unless they represent carrion. Nor can
fish, large or small, reasonably be ascribed to
the activities of owls. We suggest that the
basic owl pellet deposit has been overlaid at
various times by other bone from various
sources. Fox droppings with chewed bone
(e.g. 70.4.167) would decay to leave fragmented
remains of large as well as small vertebrates,
and other carnivores such as Sarcophilus
might also be involved in this way. Fish re-
mains could result from scavenging by foxes
on the beach nearby, or perhaps more likely,
scavenging by birds such as ravens, ospreys,
or whistling eagles (all of which were observed
during our visit, carrying fish in the first tw T o
cases). Remains of sheep might well be due

to animals dying naturally on the site; Macro-
pus irma specimen 70.3.28, consisting of most
of the bones of a single animal found in close
association, almost certainly represents such
a natural death.

It is most unlikely that the 200 x 250 m zone
of bone concentration would attract owls to
roost in its present state, completely devoid of
living plants, and almost devoid of perching
places above the ground. But there are areas
a few hundred metres further inland which
might attract owls. Such an area is shown
in Fig. 3, in which “islands” of living trees and
shrubs occur among the mobile dunes, distant
only a few hundred metres from continuously
vegetated areas. It is conceivable that owls
would hunt in the vegetated areas, but take
their prey to an “island” of vegetation from
which the approach of enemies would be readily
visible.

We suggest, thsn, that at the time of accumu-
lation of the bones, each site of local bone
concentration within the 200 x 250 m bone-rich

Figure 3. — Stands of living peppermint trees and shrubs isolated among mobile sand dunes, Bremer Bay. Such
stands, if not too far from continuously vegetated areas of profitable hunting, conceivably could serve as owl

roosts.

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

57

zone was an isolated clump or “island” of vege-
tation somewhat resembling those shown in Fig.
3, in an area in which the general plant cover
had already begun to deteriorate, and the sandy
soil to shift. In such a deteriorating area, owl
pellets might be buried before they disintegrated,
thus protecting the enclosed bones from direct
exposure to the weather, which would have
resulted in a more cracked and split appear-
ance than they have in fact.

Age of the deposit

Radiocarbon dates on wood from the fallen
peppermint tree shown in Fig. 1 and on murid
bones and teeth from the main bone accumula-
tion near this tree have been obtained. They
are 620 ± 90 years B.P. on the wood (GaK-2888)
and 1190 ±: 80 years B.P. on the bones and
teeth <GaK-2887).

Discussion

The 250 x 200 m bone rich zone mentioned
above is now completely devoid of living vegeta-
tion, but presumably at the time of accumula-
tion (about A.D. 760) supported clumps of
trees like those shown in Fig. 3, if not more
continuous vegetation. Nearly six centuries
later (about A.D. 1330) it still supported at
least one large peppermint tree, very much
larger than those shown in Fig. 3. At the pre-
sent time, isolated clumps of peppermint trees
still grow a few hundred metres away, within
the same complex of mobile dunes. Thus it
would appear that devegetation of the dune
area by (or accompanying) drifting sand has
been a very slow process, still not complete
after more than a thousand years.

Early in this history of slow devegetation, it
would appear that one or more owls inhabited
the area. Assuming that a single owl catches
mammals of about rat size at an average rate
of one mammal per night, the total bone col-
lection made by us represents only about two
years' residence by one owl.

The native mammal species listed above fit
the concept of their having been drawn by an
owl from a territory consisting partly of heath,
partly of gallery forest on the banks of a stream,
as the area is now, and under a similar climate.
Similar associations of species are known from
other deposits of Recent geological age cur-
rently under study.

For example, in Hastings Cave in the Jurien
Bay district, all four murid species listed above
(and others) occur with the marsupial species
A. apicalis, S . murina , I. obesulus , Perameles
cf. P. bougainville , P . platyops and others
(A. Baynes, personal communication). In an
archaeological deposit in the Wanneroo district
known informally as Orchestra Shell Cave,
A. apicalis , A. flavipes and S. murina occur
together with other marsupial and eutherian
species CM. Archer, personal communication).

Similar associations of species are also known
from historic time. For example, in the Cheyne
Beach district, in which A. apicalis was recently
found still to persist (Ride 1970), A. flavipes
and S. murina also have been found living.
According to Glauert (1950), the collector

George Masters obtained specimens of Pera-
meles myosurus (now generally regarded as
conspecific with P. bougainville, e.g. by Ride
1970) as well as Perameles obesula (now gener-
ally known as Isoodon obesulus /, along with
Hypsiprymnus platyops ( Potorous platyops )>
Antechinus apicalis and other marsupial species
in the King George's Sound-Pallinup (Salt)
River region in 1868-69.

Acknowledgements

We are grateful to Messrs A. Baynes (Uni-
versity of Western Australia) and M. Archer
(Western Australian Museum) for identifying
murid and small dasyurid remains respectively,
to Dr W. D. L. Ride (Director, Western Aus-
tralian Museum) for arranging to have our
wood and bone samples dated, to those persons
named in the text as having supplied informa-
tion for publication, and to Mr P. F. Aitken
(South Australian Museum) for his criticisms
of the text.

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Ride, W. D. L. (1970). — “A guide to the native mam-
mals of Australia.” (Oxford University
Press, Melbourne, 249 pages.)

Thomas. O. (1888). — “Catalogue of the Marsupialia and
Monotremata in the collection of the
British Museum (Natural History).” (Trus-
tees, British Museum (Natural History)
London, 401 pages.)

Journal of the Royal Society of Western Australia, Vol. 54 Part 2, July, 1971

58

INSTRUCTIONS TO AUTHORS

Contributions to this Journal should be sent to The Honorary Editor,
Royal Society of Western Australia, Western Australian Museum, Perth.
Papers are received only from, or by communication through, Members of the
Society. The Council decides whether any contribution will be accepted for
publication. All papers accepted must be read either in full or in abstract
or be tabled at an ordinary meeting before publication.

Papers should be accompanied by a table of contents, on a separate
sheet, showing clearly the status of all headings; this will not necessarily
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importance to be considered for publication. The Abstract (which will
probably be read more than any other part of the paper) should not be an
expanded title, but should include the main substance of the paper in a con-
densed form.

Typescripts should be double-spaced on opaque white foolscap paper; the
original and one carbon copy should be sent. All Tables, and captions
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issues of the Journal as a guide to the general format of their papers, includ-
ing the preparation of references; journal titles in references may be given in
full or may follow any widely used conventional system of abbreviation.

Note that all illustrations are Figures, which are numbered in a single
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each of these should be designated by letter (e.g. Figure 13B). Illustrations
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trations, which are usually best prepared between and 2 times the required
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mittee of the Geological Society of Australia.

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Authors are solely responsible for the accuracy of all information in their
papers, and for any opinion they express.

Journal

of the

Royal Society of Western Australia

Volume 54

Part 2

1971

Contents

5. The Western Australian psolid holothurian Ceto cuvieria (Cuvier). By D.
L. Pawson.

6. Two new genera and five new species of Percophidid fishes (Pisces: Perco-
phididae) from Western Australia, By R. J. McKay.

7. Habeobaella stagni, a new genus and species from South-Western Austra-
lia (Hirudinoidea: Richardsonianidae). By L. R. Richardson.

8. Remains of Potorous platyops (Marsupialia, Macropodidae) and other
mammals from Bremer Bay, Western Australia. By W. H. Butler and D.
Merrilees.

Editor: A. S. George
Assistant Editor: N. G. Marchant

The Royal Society of Western Australia, Western Australian Museum, Perth

87901/4/71-625

WILLIAM C. BROWN, Government Printer, Western Australia