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70.5 G44

Supplement to The Australian Zoologist,

Vol. 19, Part 1, 1976

THE
AUSTRALIAN
ZOOLOGIST

Se VOLUME 18
1973-1975

ROYAL ZOOLOGICAL SOCIETY OF NEW SOUTH WALES,

c/o TARONGA ZOO, MOSMAN, NEW SOUTH WALES 2088

INDEX TO VOLUME XVIII

Za VASON TAR

Supplement to Aust. Zool. 19(1) f \ Page One

~ £7 aia? :
( reo if 1077
44 ¢
4 s
7
ca
a

1—INDEX TO SUBJECTS

Part Page

PEREINEEDORIFOS MOOD 6S oe ete hae eR ecole MRR USC Sn cect Caer aa tenner are @A) 129
Anuran adaptations to abidity 6.0.04 5.3... Sees eee ee! PER a ee ae an (2) 53
PaeRi rats WEAR OTIORDY (8560.65 oth cc AOE Cs Ee RAC US RE Lge Re (3) 149
Pemerinig trom South Auisteelias oer haa crue Ry AR ee eae aR @ 88
Pestiatian sanuran, adaptations «os asic. eee ade teen Ree eR eae Te tee (2) 53
Bennwionn oO: rnyloeale: Bilardiery 4 oR ee ee en oN (1) 1
Breeding biology of Pseudophryne ........ eves malepeaiogs A ia saree eh eae Sek area (1) 15
TREE: WN GL age CT a a ee ae Oe, MER De 4 NATE Aine CoP A gta alee LN ret Sakai Oh Ca (2) 99
Craterocephalus dalhousiensis n.sp,. ................000000c0eeeee cathe Wie thelial eae am eae ee (2) 88
Gages arom: south-western, Tasmania is. 5. ae a es ee eae ae (3) 197
Biniseiisnes. > Tree TIEW. SPECIES ike a cl, eT ee a eee ea (1) 31
Pmecranas. bispinosus- feeding. mortality im: v.25. ke ne A es (1) 47
Rega We CLAY MSI 8 oe My OR ie ads dhe ae ae EE GB) anats7
pene. KEY dO. 0 oe oleae Wea Ege Ss eng dV ee ea icles 2 (3) La
eertrecis AOREIDINNIS -AVSD oe 2.52565 kk, SO eos) ca tek agama ms (3) HET)
Peer OPe ALE AN a MISH tee ee stew gees ie 0 SN Re a la) 47
Pearsnaearon? south Australian 26.) ce ee eke a De i ea (3) 137
Hytidac im. south-eastern eAusiralia’ 8. vs a eS Re ee 2a (3) 149
Revueles fmpeb or. Ae ear OR a nc ee PAM re vee (3) Za
Ranger SoMth-castehm: AUSthalla oe ves see ie eee eee a DER 4:3 (3) 215
aaa UPC, PCOMIPICXE, oo Ge he a Pe eee (3:): 7
SILOS PORING: 0.:.. Po. Senate hemes, | Oecd O pecalionn Vila Weta ee aiie Sie (2) qe
MR EMAITIAS SPPEETE VE (eos RP oe SE. 3k alot nn See A other ds eect (1) 43
Munriove -comimunity ecology = \)j036 oo ee Mee et I: (2) 99
Moanimals*in= western New: South™ Wales. 00. oho on eee te te (3) 179
New. Holland? Mouse. status ofc. 2) See eee Ne os Ngee ti a eee mE (2) 66
REE MPAZOIT LIN RECTLIINA AVEAPETIUS oS eed Le ia eM Ne SU acct ag ee eS Ae ae a (Oye o se
1(3)) 165

Rete FdE STACI CN LOSTIOTUCUS oh Ost Rey We: MONG cee LIN) or SSN ee eR Sea (3) 197
Pama waliaby, occurrence “and -mecOsmitione ..) (058.4850 ee tse eee eee @) 72
ER PES; “Capilire and lMarkine “ae ee las a ey | re LR oie (2) 133
(ELSE UTP a TTT 6S OVE ON cater ie ie Mate MEP 0S MRA a ee Km er A peg aIDOer TS LA a Cu (3) 165
Pamtsicenirigac, tron) the zPaciic was a ee eae lee Sate le OE ia ead ec (1) S11
Population movements of lizard ........0...0000.... SEN Osis a mens NCe li nthe wa MN a (2) 129
Pserdomiys: novachollandiags Stats Olen es... iso. alse (2) 66
ESI OPULENCE USO eaten ee Niiiiees ohana s a gen ORE A cate eden Aaa, ales (i) 1S
RANeRnCiOn il Wwhiptall -wallaby rte oie. Ce area a ee dn ee aces Gq) 43
Bee Piss COMIN) CCOIOBY, ee Tae Beh cts n Nan salt at iea sede ce en eae tee ents dae (2) 99
South: Australas new especies: of fish’ from 245005 6 ee ee | aS, (2) 88
Teviovaie=pillardient~ DehaviGul. OL seas ea eee (4) 1
BPE COSPRAL RAS EOUUIS ia) 3h ce, «as gree yippee Bes eT a ee Ra Ns i a a neh (3) 219

Page Two Supplement to Aust. Zool. 19(1)

l.—INDEX TO AUTHORS

ALLEN, G. R.
Three new species of deep-dwelling damselfishes from the south-west
ACTACS OCCA Ae, wets eh ene SN UNI a Ayal nO ARON 8 Oh ina aS ale ee
BURTON, T. C.
(See Morton, S. R. & T. C. Burton)
CARRICK, F. N.
(See Grant, T, R. & F. N. Carrick)
‘-COURTICE, G. P. & G. C. GRIGG
A taxonomic revision of the Litoria aurea complex (Anura: Hylidae) in
SOUMM-CASteRI AUISEa May ene ek a eel se ie |S eo ae
DENNY, M. J. S.
Mammals of Sturt National Park, Tibooburra, New South Wales ........
GLOVER, C. J. M. :
(See Ivantsoff, W. & C. J. M. Glover)
GRANT, T. R. & F. N. CARRICK
Capture and marking of the platypus, Ornithorhynchus anatinus, in the
NISC ie PRN CS SU SG ITA ca oye RSE a GAS Oo Mea gteas M U Ga cmH Ae Iara pare 9
GRIGG, G. C.
(See Courtice, G. P. & G. C. Grigg)
‘HUTCHINGS, P. A. & H. F. RECHER
The fauna of Careel Bay with comments on the ecology of mangrove and
SEa-O8FaSS: (COMMUNITIES wie eee ee a i ae SOR oO dest aye

IVANTSOFF, W. & G. J. M. GLOVER
Craterocephalus dalhousiensis n.sp., a sexually dimorphic freshwater teleost
(Atherinidae) trem: South: Australian wie. cee ee ae ee
LAKE, P. S. & K. J. NEWCOMBE
Observations on the ecology of the crayfish Parastacoides tasmanicus
(Decapoda: Parastacidae) from south-western Tasmania ...............00..0......
LAVENBURG, R. J.
(See Paxton, J. R. & R. J. Lavenburg)
LEB, A?’ Ki
(See Seymour, R. S. & A. K. Lee)
MAYNES, G. M.
Aspects of reproduction in the whiptail wallaby, Macropus parryi ........
Occurrence and field recognition of Macropus parma. ......0......ccccccce
MORTON, S. R. & T. C. BURTON
Observations on the behaviour of the macropodid marsupial Thylogale
billardiert, (Wesmarest)i- in Capuyity’ a cae cece Ue ee
NEWCOMBE, K. J.
(See Lake, P. S. & K. J. Newcombe)
PAXTON, J. R. & R. J. LAVENBURG
Feeding mortality in a deep sea angler fish (Diceratias bispinosus) due to
a Macrounia hshy CVeniriyfOSSa: SP). ak ee ge A
PENGILLEY, R.
Breeding biology of some species of Pseudophryne (Anura: Lepto-
dactylidae) of the Southern Highlands, New South Wales .................0......
PONDER, W. F.
The identity of the common keyhole limpet of south-eastern Australia
(Fissurellidae) © 2.0.0.0... SUR eS ea OL coh by Ga nal Oke kuna rales sae va ee

Supplement to Aust. Zool. 19(1)

Part Page
(1) 3]
(3) 149
(3) 179
(2) 133
(2) 99
(2) 88
(3) 197
(1) 43
@) 72
(1) 1
(1) 47
(1) 15
(3) 215
Page Three

II—INDEX TO AUTHORS—cont’d

Part Page

POSAMENTIER, H. & H. F. RECHER

The status of Pseudomys novaehollandiae (the New Holland Mouse) (2) 66
RECHER, H. F.

(See Hutchings, P. A. & H. F. Recher)

(See Posamentier, H. & H. F. Recher)
SEYMOUR, R. S. & A. K. LEE

Physiological adaptations of anuran amphibians to aridity: Australian

1 ESTE © IER cee ROR ae ac SURREAL ta nea I ee Re Nt OPO AS (2) 53
STRAHAN, R.

Eptatretus longipinnis, n.sp., a new hagfish (Family Eptatretidae) from

South Australia, with a key to the 5-7 gilled Eptatretidae ................. Rath C3) 137
STRAHAN, R. & D. E. THOMAS

Courtship of the platypus, Ornithorhynchus anatinus ......00.0.....0000000cceceees (3) 165
THOMAS, D. E.

(See Strahan, R. & D. E. Thomas)
Wriltray, G. P.

Bins fe Geyt Troughton, jC. M.7.S:., F-R:Z.S. (1893-1974). ee Ga, 218
WITTEN, G. J.

Population movements of the agamid lizard, Amphibolurus nobbi. ........ (2) 129

DATES OF PUBLICATION

oe TS o's | ae ear, Uk Hie Pm elena OMI ieee eg UGY ns August, 1973
PAR AMARCS OO 190. 20 ee ae Vora a aae ces a en June, 1974
ene oe DAGCS.. 137-2380. een ee | tur As ee nes August, 1975

Printed and published for the Royal Zoological Society of New South Wales,

c/o Taronga Zoo, Mosman, New South Wales, 2088

Surrey Beatty & Sons, Rickard Road, Chipping Norton, New South Wales, 2170.

Volume 18, Part

August, 1973

Scientific Journal of

The Royal

Price $2.50

IK

ological Society of New South Wales

NOTICE TO AUTHORS

Papers will be considered for publication in The Australian Zoologist
if they make an original contribution to whole animal biology of the Australian
fauna. Papers submitted will be subjected to review and thence to the norm
editorial process, in the course of which authors will receive edited gall

G a

oS

Australian Zoologist”, School of Biological Sciences, Macquarie University,
North Ryde, N.S.W. 2113. They should be typewritten (double spaced
good quality paper. All pages of the manuscript must be numbered consecutivel
including those containing references, tables and figure legends, which sh
all be placed after the text. ) ea ety tae oh:

MANUSCRIPTS (original and one copy) should be sent to the Editor, “The

On the first page of the manuscript should appear the title of the pape
name of the author, the name of the Institution where the work was done
the present postal address if different from that of the Institution. Titles
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head and consisting of not more than 50 letters (including spaces) must
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The abstract (up to 200 words) should state concisely the scope «
work and the principal findings and should be suitable for direct
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and Methods, Results, Discussion, Acknowledgements and References. Pres
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consecutive sections should be avoided. Footnotes should be avoided. =.

REFERENCES are cited in the text by the author and date and are no
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Abbreviations of titles of periodicals should conform to those used i
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A

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TABLES should be numbered with arabic numerals, be accompanied b

aa

ar An
et

THE AUSTRALIAN ZOOLOGIST

VOLUME 18 1973 PART 1

Observations on the Behaviour of the Macropodid
Marsupial Thylogaile billardieri (Desmarest) in Captivity

S. R. MORTON and T. C. BURTON
Department of Zoology, University of Melbourne, Parkville, Victoria 3052.

ABSTRACT

Observations were made on the diurnal behaviour of a captive group

(82 6, 1122) of Thylogale billardieri (the rufous-bellied pademelon). The

daily activity pattern, postures, locomotion, feeding, grooming, vocalisations,

agonistic and sexual behaviour are described, and comparisons made with
the behaviour of other macropodids.

INTRODUCTION

The rufous-bellied pademelon, Thylogale billardieri (Desmarest) is one of the
smaller members of the Macropodinae, adult males standing approximately 70 cm
and adult females approximately 60 cm tall. A mature female weighs about 5.5 kg.
There is wide variation in coat colour, ranging from rufous to dark brown. Some
animals possess a pale hip-stripe or faint facial markings, but these vary between
individuals. The pademelon is restricted to Tasmania and the Bass Strait islands,
although it formerly also occurred in Victoria and south-eastern South Australia
(Calaby, 1971). Information on the biology of T. billardieri is sparse. It has been
described as an “extremely gregarious” species (Troughton, 1965) which lives in
“colonies” (Sharland, 1962). Mollison (1960) summarised its favoured habitats
in Tasmania as mixed forests of lower slopes, bracken or grassland on forest fringes,
and burnt and cutover buttongrass and Poa savanna. Troughton (1965) noted that
runways ate constructed through dense scrub.

The maintenance of a group of rufous-bellied pademelons by the Melbourne
Zoological Gardens provided the opportunity to make behavioural observations on
the species. While captivity undoubtedly results in artificialities and modifications
of normal behaviour, studies of captive animals have nevertheless contributed useful
data. For example, the observations of Packer (1969) on captive quokkas (Setonix
brachyurus) and Russell (1970a) on captive red kangaroos (Megaleia rufa)
gave results consistent with field-derived behavioural data. The present study is a
preliminary assessment of aspects of pademelon behaviour, which may provide a
basis for more definitive studies.

Aust. Zool. 18(1), 1973 1

S. R. MORTON and T. C. BURTON

MATERIALS AND METHODS

The composition of the pademelon group during the study is shown in Table
1: most data in the table are drawn from Zoo records. Females preponderate over
males, due to the Zoo’s policy of removing most male neonates from the pouch
in order to maximise breeding stock. 3

TABLE 1
COMPOSITION OF PADEMELON GROUP

Zoo Date of
Code Sex Leaving Ojfspring Origin Removal From
No. Pouch Group
502 Male 4 Healesville
1967 d. March 30
501 Female ? 505, 504, Healesville
35 1967 d. August 9
505 Female Sept. 1968 42 Zoo |
504 Male Oct. 1969 Zoo Escaped April 29 |
598 Female 1969 43 King Is. 1970
2 Female ? 33 Tasmania
1970 )
5 Female. :>;? Tasmania d. August 20
1970
32 Female 1970 ay Zoo d. August 11
33 Female 1970 Zoo d. May 11
35 Female 1971 Zoo
37 Female 1971 Zoo
42 Male May 1971 Zoo
43 Female April 1971 Zoo
“44 Male u *“May 1971

*44. a mature male, was introduced into the group from a bachelor stock kept elsewhere in
the Zoo.

Three rectangular enclosures housed the pademelons at different times during
the study. The first enclosure (March 18 to April 19, 1971) was about 180 m?
in area and contained a shed, water trough, trees along the northern boundary, and
scattered shrubs elsewhere. The second enclosure (April 20 to May 20) was the
same as the first, but contained several more trees and some scattered boulders
as well as a shed and trough. The third enclosure (May 21 to October 6) was
about 360 m? in area, and contained a shed and trough but fewer trees than the
previous enclosures. In this last enclosure there were seven red-necked wallabies
( Macropus rufogrisus) as well as the pademelons. Hay, bran and carrots were placed
in the sheds as supplementary food.

Observations of behaviour totalling 81 hours were made between March
18 and October 6, 1971. All observations were made during daylight hours from

2 Aust. Zool. 18(1), 1973

BEHAVIOUR OF THYLOGALE BILLARDIERI

outside the enclosure, sometimes with the aid of 6 x 30 binoculars. Individual
animals were readily recognised by combinations of characteristic facial markings,
coat colour and size. In addition, each animal had a Zoo registration number on
an eattag; in this report these numbers are used to identify individuals. The duration
of observation periods varied between 30 minutes and 180 minutes, with the
majority being about 90 minutes. Initially, observations were made at various
times of the day, but as it was found that activity was greatest in the late afternoon,
observations were later concentrated in this period.

RESULTS

(1) Daily Activity Pattern:

Pademelons are said to be crepuscular in habits, emerging from cover during
dusk and early mornings to feed (Troughton, 1965). Limitations of the Zoo study
did not allow this aspect of activity to be analysed. However, there was often
activity — grooming and grazing — in the mid-morning and always in the late
afternoon. During periods of inactivity the animals assumed a resting posture and
often slept for periods of more than an hour. Diurnal activity appeared to vary
with daily weather pattern and season, with greatest activity occurring on cool
and cloudy days.

A preference for shade was normally own. the animals moving into direct
sunlight only when grazing. However this preference altered with season, being
most marked in summer. On two occasions (July 25 and September 24) some
pademelons were observed lying stretched out in the sun.

(2) Body Postures:

When asleep or resting, the pademelons adopted a characteristic pose
(Fig. 1). The tail was thrust forward between the hind legs and the head rested
on the tail. In general this attitude was adopted with the back to a tree or fence,
although depressions in the ground were sometimes used.

The lying-down pose characteristic of the red kangaroo (Russell, 1970a) was
rarely adopted, and then only when lying in the sun in winter.

During rain the pademelons adopted a semi-upright stance, with tik orien
posteriorly and head thrust forward.

-(3) Locomotion:

The hopping gait of the pademelon is basically similar to that of es
small macropodids, but shows some differences. Pademelons hold their forelegs
against the chest, and their gait is not markedly saltatorial. When alarmed they
showed great speed and agility.

The walking gait was employed when grazing or when a male was “inspecting”
females (see Sexual Behaviour). Unlike the red kangaroo (Russell, 1970a), the tail
is not used as a support during walking.

Aust. Zool. 18(1), 1973 3

MORTON and T. C. BURTON

R.

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Zool. 18(1), 1973

Aust.

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BEHAVIOUR OF THYLOGALE BILLARDIERI

(4) Feeding and Drinking:

In the wild, pademelons eat a wide variety of grasses, shoots and leaves
(Mollison, 1960). In captivity all grasses in the pens were grazed, as well as
leaves, bark and supplementary foods. Forepaws were sometimes used for picking
up food and parting thick grass.

Regurgitation in the rufous-bellied pademelon appeared to be comparatively
infrequent, being observed only six times during the period of study. The process
seemed identical to, but less frequent than, that described for the quokka
(Moir, Somers and Waring, 1956). Pademelons re-ate all spilled material after
regurgitation, and chewed the bolus in every case.

Drinking was most commonly seen when the animals began grazing in the
late afternoon.

(5) Grooming:

Grooming was a prominent activity throughout the day. Three general modes
of grooming were observed:—

(a) Licking — Mainly used for pouch and abdominal cleaning. The tongue
was used to clean while the snout nuzzled into the fur. Forepaws were often used
to part fur to facilitate licking (Fig. 2).

(b) Scratching with Forepaws — Used in grooming flanks, tail, back,
abdomen, chest and face. The forepaws scratched vigorously through the fur and
were often licked clean.

(c) Scratching with Hindfeet —- Used in grooming head and shoulders. The
hindfoot has a syndactylous inner claw which Wood Jones (1924) considered to
be an adaptation for combing fur.

Pouch cleaning may be considered as a specialised type of grooming
(Fig. 3). It was observed in females both with and without pouch-young, and was
essentially as described for the red kangaroo (Russell, 1970a).

Pademelons groomed at various times during the day while at rest. They
awoke, stood erect, and groomed for varying periods (1 to 10 minutes), then
lapsed back into the resting posture and returned to sleep. This pattern of waking,
grooming and sleeping often continued for some hours. Finally in the late afternoon
the animals became more active and groomed again for a considerable period
(usually more than 15 minutes) before beginning to graze. Grooming appeared
to increase after rain, and in some observation periods after heavy rain it was
the predominant activity.

Mutual grooming is discussed under mother-young interactions.

(6) Vocalisations:
The following vocalisations were heard:—

(a) Growling — Low intensity sounds made by the dominant individual in
an ageressive interaction. Both males and females used this sound, which was

Aust. Zool. 18(1), 1973 5

S. R. MORTON and T. C. BURTON

ae

NX

Figure 3—Grooming: Pouch

cleaning.

Figure 4—Sexual behaviour: “Inspection” by male

(right) of a female. The third individual
is another female. (Photos: A. A. Martin).
6

Aust. Zool. 18(1), 1973

at

BEHAVIOUR OF THYLOGALE BILLARDIERI

dificult to pinpoint accurately. This vocalisation is probably equivalent to the
“snort” of the red kangaroo (Frith and Calaby, 1969; Russell, 1970a).

(b) Clucking — This was heard in three different situations:

(i) Emitted by a male in sexual following, and presumably analogous to the
“soft clucking” of the red kangaroo (Russell, 1970a).

(ii) Emitted by a male in aggressive interactions, and probably equivalent
to that described for the red kangaroo (Sharman and Calaby, 1964) and Bennett’s
wallaby (LaFollette, 1971).

(iii) Emitted by a female to her young. This seems equivalent to the “loud
clucking” of the red kangaroo (Russell, 1970a). No distinction could be made
between the clucking in each of these situations, but because of the low intensity of
the sound it is possible that differences went unnoticed. Sounds equivalent to the
distress squeak of young red kangaroos (Sharman and Calaby, 1964; Russell, 1970a)
were not heard. However, this may also be due to the difficulty of hearing
vocalisations adequately.

(7) Aggressive Interactions:

(a) Male-Male — Two males (502 and 504) were present in the enclosure
together for a period of only two weeks, but 10 hours of observation during this
period showed 12 aggressive interactions between them. All these resulted in
the younger, smaller 504 retreating. Most interactions occurred when 504 came
within about 2m of 502, and the result was immediate chasing by 502 for a
distance of approximately 5m. The two never came close enough together to
actually fight. On one occasion 504 approached the resting 502 and adopted a
pose face to face with him, his front paws on the ground and his body stretched
horizontally so that his quivering snout was almost touching that of 502. When

FABLE: :'2

AGGRESSIVE INTERACTIONS BETWEEN FEMALES

DOMINANT
505 e508 35 32 33 a7 gone 4a
s | 505 as
U 2 9 ao 1
. 598 17 7 vas
= 35 6 12 7 its. 2 5
D 32 5 8 2 1 aed
I 33 2 1 1 3 0 <2
: 37 5 13 3 12 5 0 au 4
= 5 9 3 3 1 1 0 0 ia
Bie a3 1 5 19* 1 0 0 ie ee

*AIl these interactions occurred after August 20 when weaning occurred.

Aust. Zool. 18(1), 1973 7

S. R. MORTON and T. C. BURTON

502 began to rise after about 15 sec 504 fled. The pose adopted by 504 is
tentatively termed a threat pose. It is not certain that 504 was sexually mature
at this time, but he engaged in sexual following (see Sexual Behaviour) often with
penis erect. .

(b) Female-Female — Table 2 is a summary of the results of aggressive
interactions between females: the data suggest that a near linear hierarchy exists.
The types of interaction observed and included in the table were:

(i) One animal retreating from another with no apparent provoking action
from the dominant.

(ii) One animal chasing another for varying distances.
(iii) One animal cuffing another with the forepaws.

(iv) Fighting, i.e., mutual cuffing or wrestling with forepaws. Hindfeet were
not used in fighting.

The latter three types of interaction often included growling. Between females
the threat pose as described for males was observed 15 times, but quivering of the
muzzle was not always seen. Usually the animal adopting the pose subsequently
cuffed or chased the other.

The level of aggression of females varied throughout the study, and there is
evidence that this was related to the presence or absence of pouch-young. For
example, 60 per cent of the observed aggressive interactions of female 5 occurred
between July 11 and her death on August 20. Pouch inspection on August 12
revealed the presence of a 4cm pouch-young (estimated age about 25 days).
Similarly, 68 per cent of the observed interactions of female 37 occurred between
July 6 and October 6, compared with 32 per cent between March 18 and July 6.
She carried a 5 cm pouch-young (estimated age about 35 days) on August 12.
Again, 67 per cent of the observed interactions of female 505 occurred before May
20, when her young first left the pouch. Of her later observed interactions, 50 per
cent were with females 5 and 37 who were then acting more aggressively. The
increased aggression of females did not affect their hierarchical position. The ability
to dominate in an interaction appeared to be positively correlated with body size

(Table 3).

TABLE 3
RELATION BETWEEN BODY SIZE AND POSITION IN DOMINANCE HIERARCHY
IN FEMALES
Order of size (largest to
smallest by visual assessment) 505 2, 598 5 35 32 Sil 43
equal
Position in hierarchy .............. 1 2 3 4 4 4 7 8
equal
(c) Male-Female — Aggression between a male and a female was noted

several times. During attempts at courtship (described below), cuffing was used by

8 Aust. Zool. 18(1), 1973

BEHAVIOUR OF THYLOGALE BILLARDIERI

a female to break the male’s hold, and the male often retaliated by cuffing or
chasing the female. Males were also cuffed about the face whenever they attempted
to sniff a female’s pouch area, and the males retaliated by cuffing but rarely by
chasing. On one occasion female 5 was chased extensively by male 504 for no
apparent reason. The threat pose described above was observed twice in females
during male-female interactions.

(8) Sexual Behaviour:

A daily occurrence was the “‘inspection” of females carried out by a male,
during which he walked around the enclosure, stopping to sniff the genital area
and/or pouch region of each female encountered (Fig. 4). Often there
was no further interaction, but on each occasion the male attempted to court at
least one female. This involved the male’s standing at full height, sometimes with
head raised, in front of the female. The female either raised her head until it was
just under the male’s, or the male wrestled her or cuffed her under the chin until
she adopted this position. At this stage the male either bent to sniff the female’s
genital area again, or, more often, placed his forearms around the female’s neck,
and, retaining his grip with one arm, moved behind the female. The male then
held the female’s flank with the free arm and slipped the other back until the
female was held by both flanks; then he attempted to mount. The female reacted
to this by either turning to cuff the male, or, more frequently, hopping forward
out of the male’s grasp. The male usually followed, sometimes making a clucking
sound, and repeated the courtship attempt. This pattern of courtship, escape,
pursuit, courtship often continued for some time, and on one occasion male
44 made 28 attempts to court female 42 over a period of 15 minutes. At any stage
of the courtship sequence the male erected his penis, sometimes by licking it. On
two occasions the male was seen to perform a brief copulation-like display in front
of a female (505 and 42), in which, with penis erected, he jerked his pelvis 5 or 6
times.

Only once was copulation observed: between male 44 and female 505 (1630
hours, September 3). After the initial sniffing, the male licked the female’s genital
area. The female pushed him away and adopted the resting posture. Shortly
afterwards, while the female was cleaning her pouch, the male groomed himself
and licked his penis, which was erect. He then groomed the female’s ears by
licking, and pushed his snout in under her chest and licked her genital area. Then
he stood upright with his head raised vertically, and wrestled the female until
she was also upright (but without head raised). He then moved behind the female
as described above, gripping her flanks so tightly that his forearms were hidden
in the folds of skin formed by her hind legs and side. He then mounted and mating
occurred.

Three phases were observed in mating:—

(a) Intromission, during which the female was almost lifted off her hind
legs, and strained forward with forepaws on the ground. Sometimes her head lay
on the ground, sometimes it stretched forward horizontally. On one occasion she

Aust. Zool. 18(1), 1973 9

S. R. MORTON and T. C. BURTON

ate briefly. At times she “walked” sideways on her forepaws so that the pair moved
in a circle as copulation proceeded. No lateral tail movements took place.

(b) A rest phase in which both animals relaxed. The male retained his firm
srip, but his penis relaxed and both animals Se still. Occasionally the male’s
head rested on the female’s back.

(c) A phase in which the male vigorously massaged the female’s flanks. His
penis was relaxed in this phase.

These phases were repeated in seemingly random order during each of the
three mountings. The durations of these mountings were respectively 8, 2 and 8
minutes, and approximately half the time in each was spent in copulation, the rest
of the time being equally shared between rest and massage. Between mountings
the animals groomed themselves and fed briefly. The times between mounts were
4 and 15 minutes respectively. After the third mounting the two animals fed and
groomed side by side. There may have been subsequent mountings but the closure
of the Zoo terminated observations.

Following this copulation an attempt was made to learn the gestation period,
assuming successful fertilisation on September 3. Because the lowest recorded
macropodid gestation period is 22 days (in the boodie, Bettongia lesueuri; Frith and
Calaby, 1969), pouch checking of 505 did not commence until 24 days later. On
this day a young was present in the pouch and from its development Zoo staft
estimated its age as 2 weeks. Either the pademelon has an extremely short gestation
period or the female was pregnant at the time of mating.

Zoo records indicate that pademelons breed throughout the year in captivity,
and that females have only one young per year.

(9) Mother-Young Interactions:

The exact period of pouch life has not been determined, but is estimated to
be about 25 weeks, as in the quokka (Waring, Sharman, Lovat and Kahan, 1955).

When young first left the pouch for short periods they were easily alarmed
and dashed back to the mother on any disturbance. The time spent out of the
pouch increased gradually until about 5 weeks later, when it increased sharply over
about 2 days, so that the young no longer entered the pouch. The young-at-foot
continued to suckle, but with decreasing frequency. About 4 months later weaning
occurred. This appeared to involve a significant amount of aggression, as can be
seen from the interactions of 598 and 43 in Table 2. These interactions mostly
consisted of cuffing, and sometimes chasing, by the mother. Aggression towards
the young tapers off rapidly and in one instance the mother was seen to briefly
suckle the young one month after weaning.

Further interactions involved what is best termed mutual grooming. The
mother and her offspring cleaned and groomed each other’s head and neck, licking
and using forepaws. While not doing so, the two often rested side by side for
periods of over an hour, but the actual grooming rarely persisted for longer than
15 minutes. Such grooming occurred between all mother-offspring pairs in the

10 Aust. Zool. 18(1), 1973

BEHAVIOUR OF THYLOGALE BILLARDIERI

group, but the male offspring appeared less inclined to take part in it than the
females. Male 504 was seen to groom 501 on only two brief occasions, and male
42 (left the pouch in June) began grooming less with his mother after September.

On the other hand, mutual grooming between mother and female offspring
appeared to persist for a longer time, although with gradually decreasing duration.
For example, 505 left 501’s pouch in September 1968 and was seen grooming with
her in April 1971. The strength of this bond is also evidenced by observation of
mutual grooming between a mother and an offspring which was carrying a large
pouch-young, and between a mother and two different offspring simultaneously.
Thus mutual grooming is a persistent phenomenon, although its frequency and
duration slowly decrease.

Sparring between mother and offspring, as described for red kangaroos by
Russell (1970a) was observed only once in the pademelon group.

DISCUSSION

General Behaviour:

The general activity and body postures resemble those of other small
macropodids. Thus the diurnal rhythm of activity is very similar to those described
for the red kangaroo, Megaleia rufa (Caughley, 1964; Russell, 1970a), and grey
kangaroo, Macropus giganteus (Caughley, 1964) and clearly, at least in captivity,
pademelons are not strictly crepuscular. The resting posture seems suited to
conservation of body heat, and may be additionally adaptive in making the animal
less conspicuous to predators. The lying-down posture characteristic of the red
kangaroo was seen on only two days in winter, when animals lay in the sun; this
may also relate to regulation of body warmth. The resting and threat postures are
virtually identical to those recorded for the quokka by Packer (1969).

Social Organisation:

The aggression displayed by female pademelons is more pronounced than that
described for captive groups of boodies (Stodart, 1966), quokkas (Packer, 1969),
Bennett’s wallabies (LaFollette, 1971) or red kangaroos (Russell, 1970a, b). In
captive groups of all these animals, however, some form of dominance between
females was observed. The relevance of this aggression in captive animals to their
behaviour in the wild is questionable. Caughley (1964) and Frith and Calaby
(1969) concluded that red kangaroo mobs showed little social cohesion. Dunnet
(1962) stated that quokkas were non-gregarious and lacked any social organisation,
although Holsworth (1967) demonstrated the existence of group territories within
which individual quokkas inhabited home ranges. Boodies appear to live in social
groups (Finlayson, 1958), but the field behaviour of Bennett’s wallabies is un-
described. Hence the presence of dominance interactions between female
macropodids in captivity is not proof that natural groups have definite social
organisations.

The degree of aggression shown may, however, illuminate differences in
organisation. Stodart (1966) gave some evidence of a hierarchy amongst female

Aust. Zool. 18(1), 1973 11

S. R. MORTON and T. C. BURTON

boodies, while Packer (1969) concluded that no linear hierarchy could be
distinguished in female quokkas. Similarly, interactions between female red
kangaroos (Russell, 1970a, b) appear to be of a much lower intensity than those
seen between boodies, even though Russell (1970b) demonstrated the existence ©
of a hierarchy in groups of female red kangaroos. These differences may well be
due to the differences in social organisation, and because female pademelons appear
to be more aggressive it may be that pademelons have a definite social structure.
Both Troughton (1965) and Sharland (1962) state that pademelons live in
colonies or communities, and the latter writes of their aggressive temper. It is,
therefore, possible that the aggression and dominance hierarchy observed in captive
pademelons form part of their natural behaviour. Artificiality does, however, arise
from the heterogeneity of the group of pademelons studied (see Table 1), the
lack of males, and the limited area of the enclosures.

Sexual Behaviour:

The mating behaviour of the pademelon affords interesting comparisons with
that described for other macropodids: red kangaroo (Sharman and Calaby, 1964),
grey kangaroo (Poole and Pilton, 1964), the boodie (Stodart, 1966), and the
quokka (Packer, 1969). .

(a) Licking — The licking by the male of the urinogenital area of the female
is known in the boodie, but is not described for the other three macropodids. In
the boodie, licking was observed on the day preceding presumed copulation whereas
it was observed immediately before copulation in the pademelon.

(b) Circling during Intromission — The female grey and red kangaroo are
both described as adopting a “submissive posture’ during intromission. The
boodie was observed circling in one of the three matings described by Stodart.

(c) Repeated mating — Not observed in the boodie, observed poly once in
the red kangaroo, but “typical” of the grey kangaroo.
(d) Rest periods — These occur in the other macropodids, but there is no

description for any of them of the vigorous and unmistakeable flank massage
which was seen in each mating in the pademelon.

Breeding in pademelons occurs throughout the year in captivity, but field
behaviour may be different (Guiler, 1960). Quokkas breed continuously in captivity
but seasonally in the field (Shield, 1968).

The anomalous result obtained for gestation period may be explained either
by an anoestrous mating or by the existence of an extremely short gestation period.
While the former is more likely, Poole and Pilton (1964) and Frith and Calaby

(1969) have shown how rare anoestrous mating is in grey and red kangaroos.

Mother-Young Association:

The sequence of mother-young interactions in the pademelon closely parallets
that described for the red kangaroo by Russell (1970a), but mother-offspring
relations appear to persist Jonger in the pademelon. Russell (1970a) showed that
although there was a long period of association between mother and offspring in
red kangaroos, there was no evidence that such associations persisted beyond

a2 Aust. Zool. 18(1), 1973

BEHAVIOUR OF THYLOGALE BILLARDIERI

sexual maturity. Similarly, association of euro (Macropus robustus) mother and
offspring may continue for only two years (Ealey, 1967). There is evidence of
association of young quokkas in their natal territories (Holsworth, 1967), but
there is no evidence of any persistent mother-offspring bond. Association in the
above examples is related to suckling, whereas in the rufous-bellied pademelon there
is stronger association through mutual grooming. No evidence regarding the
adaptive significance of this persistent bond is available.

Weaning may also be marked by differences from other macropodids, as
ageression towards the young-at-foot during weaning is quite evident in pademelons.
In other species weaning appears to involve mere rejection of the young (e.g.,
Russell, 1970a).

ACKNOWLEDGEMENTS

We are indebted to the Director of the Melbourne Zoo, Mr. J. H. Sullivan, for allowing
us the facilities for this study. We are also very grateful to the staff of the Zoo, particularly
Mr. E. Weber, Mr. R. W. Dunn, Mr. G. C. Morris, Mrs. H. da Costa and Mr. G. Cox,
for their assistance. Mr. Bill Atyeo helned with observations in the early part of the study.

Special thanks are due to Dr. A. A. Martin for his helo and encouragement throughout
the study. Dr. A. A. Martin and Mr. G. F. Watson read and criticised the manuscript.

REFERENCES

CALABY, J. H. (1971). The current status of Australian Macropodidae. Aust. Zool. 16,
17-29.

CAUGHLEY, G. (1964). Social organisation and daily activity of the red kangaroo and
grey kangaroo. J. Mammal. 45, 429-36.

DUNNET, G. M. (1962). A population study of the quokka, Setonix brachyurus (Quoy
and Gaimard) (Marsupialia). Il. Habitat, movements, breeding, and growth. C.S.I.R.O.
Wildl. Res. 7, 13-32.

EALEY, E. H. M. (1967). Ecology of the euro, Macropus robustus (Gould), in north-
western Australia. II. Behaviour, movements, and drinking patterns. C.S.I.R.O. Wildl.
Res. 12, 27-51.

FINLAYSON, H. H. (1958). On central Australian mammais. Pt. HI—The Potoroinae.
Rec. S. Aust. Mus. /3, 235-302.

PRO He Jo and “ CALABY, 3:44. (1969)... “Kangaroos”. (Cheshire: . Meloburne):.

GUILER, E. R. (1960). “Marsupials of Tasmania”. (Tasmanian Museum and Art Gallery:
Hobart. )

HOLSWORTH, W. N. (1967). Population dynamics of the quokka, Setonix brachyurus,
on the west end of Rottnest I.. Western Australia. I. Habitat and distribution of the
quokka. Aust. J. Zool. 15, 29-46.

LaFOLLETTE, R. M. (1971). Agonistic behaviour and dominance in confined wallabies,
Wallabia rufogrisea frutica. Anim. Behav. 1/9, 93-101.

MOIR, R. J.,. SOMERS, M. and WARING, H. (1956). Studies on marsupial nutrition.
1. Ruminant-like digestion in a herbivorous marsuvia! (Setonix brachyurus). Aust.
J. Biol. Sci. 9, 293-304.

MOLLISON, B. C. (1960). Progress renort on the ecology and control of marsupials in
the Florentine Valley. Appita. 14, XXI-XXXVI.

PACKER, W. C. (1969). Observations on the behaviour of the marsupial Setonix
brachyurus (Quoy and Gaimard) in an enclosure. J. Mammal. 50, 8-20.

POOLE, W. E. and PILTON, P. E. (1964). Revroduction in the grey kangaroo, Macropus
canguru, in captivity. C.S.L.R.O. Wildl. Res. 9, 218-34.

Aust. Zool. 18(1), 1973 13

S. R. MORTON and T. C. BURTON

RUSSELL, E. M. (1970a). Observations on the behaviour of the red kangaroo (Megaleia
rufa) in captivity. Z. Tierpsychol. 27, 385-404.

RUSSELL. E. M. (1970b). Agonistic interactions in the red kangaroo (Megaleia rufa).
J. Mammal. 5/, 80-88.

SHARLAND, M. (1962). “Tasmanian Wild Life”. (Melbourne University Press:
Melbourne. ) ;

SHARMAN, G. B. and CALABY, J. H. (1964). Reproductive behaviour in the red
kangaroo, Megaleia rufa, in captivity, C.S.I.R.O. Wildl. Res. 9, 58-85.

SHIELD, J. (1968). Reproduction of the quokka, Setonix brachyurus, in captivity.

J. Zool., Lond. /55, 427-44.

STODART, E. (1966). Observations on the behaviour of the marsupial Bettongia lesueuri
(Quoy and Gaimard) in an enclosure. C.S.I.R.O. Wildl. Res. 11, 91-102. -

TROUGHTON, E. (1965). “Furred Animals of Australia”. (Angus and _ Robertson:
Sydney. )

WARING, H., SHARMAN, G. B., LOVAT, D. and KAHAN, M. (1955). Studies on
marsupial reproduction. 1. General features and techniques. Aust. J. Zool. 3, 34-43.

WOOD JONES, F. (1924). “The Mammals of South Australia”. Part 2. (Government
Printer: Adelaide. )

14 Aust. Zool. 18(1), 1973

Breeding Biology of some Species of Pseudophryne
(Anura: Leptodactylidae) of the Southern Highlands,
New South Wales

R. PENGILLEY

School of Biological Sciences, Macquarie University, North Ryde,
New South Wales, 2113.

ABSTRACT

Populations of Pseudophryne corroboree, P. dendyi and P. bibroni in the
highlands and tablelands of south-eastern N.S.W. breed when temperatures
are decreasing. In P. corroboree the onset of breeding is negatively
correlated with altitude. Modal clutch sizes of the three species are as follows:
corroboree (20-29), dendyi (80-89), bibroni (80-99). In general fecundity is
not related to female body length.

INTRODUCTION

Most species of the genus Psewdophryne are terrestrial breeders, laying their
eggs in burrows in soil, under leaf litter, or in moss in areas which are subject
to seasonal inundation. The number of relatively large, heavily yolked eggs laid
varies from 20 in P. australis, to well over 100 in P. bibroni. From the information
available (Table 1), it appears that most species breed during the late summer
to late autumn or early winter. Both P. australis and P. occidentalis are atypical
in this respect as the former species is reported to breed throughout the year
after rain (Harrison, 1922) and the latter ‘“‘breeds after summer rains and
when these fail . . . breeds after the first warm rains of winter’ (Main, 1965).

In P. corroboree, P. dendyi and P. bibroni, the male constructs the burrow
which also serves as a calling site (Pengilley, 1971a). Colefax (1956) reported
that in P. corroboree the female performs this task as he observed in the
laboratory, females making burrows in moss after injection with Bufo marinus
pituitaries. In nature, no female has been observed to make a burrow.

From the literature, species appear to differ in their behaviour after egg
laying. Males of P. douglasi (Main, 1964), P. coriacea (Straughan, 1963, pers.
comm.), P. dendyi (Moore, 1961), P. corroboree (Jacobson, 1963) and females
of P. australis (Harrison, 1922) have been found in burrows containing eggs.
Jacobson (1963) reported that both sexes of P. bibroni remain with the eggs.
This is contrary to my observations on this species, where only males have
been found in nests containing eggs.

Aust. Zool. 18(1), 1973 15

R. PENGILLEY

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18(1), 1973

Aust. Zool.

16

PSEUDOPHRYNE BREEDING BIOLOGY

Because of the paucity of information available on the breeding biology of
species of this genus, a number of populations of P. corroboree, P. dendyi and
P. bibroni in the Southern Highlands and Southern Tablelands were examined
as frequently as possible during the breeding seasons. During the examination of
these areas, particular attention was placed on obtaining fairly accurate information
on the onset and duration of the calling and breeding seasons of the three species.
In the case of the calling season, it was thought that the time when the males were
first heard calling probably indicated that males were beginning to move into the
breeding areas. Known burrows in the breeding areas were periodically checked
to see whether there were any males present. The examination of burrows during
these visits also provided information on fecundity and on the date of commence-
ment of breeding and fecundity.

MATERIALS AND METHODS

Locations of the study areas have been given elsewhere (Pengilley, 1971a).
Most of the areas that were visited more than once during the calling and breeding
seasons are situated in the montane or subalpine zones of Costin (1954) within

south-eastern N.S.W.

The main study area, which is located at Coree Flats (alt. 1036m) on the
Brindabella Range near the north-western corner of the N.S.W.-A.C.T. border,
is a wet heath surrounded by dry-wet sclerophyll forest (Eucalyptus dalrympleana-
E. robertsoni alliance). Within the wet heath, the majority of breeding animals
were concentrated at the northern and southern parts where the vegetation had
been partly modified by man. In these small areas, the herbaceous layer consisting
of a diverse assemblage of snow grass (Poa caespitosa), species of Agrostis, and
various herbs and mosses, (Fig. 1) provided suitable breeding sites for P.
corroboree and P. dendyi. Snowy Flats (alt. 1646 m) is situated on the Brindabella
Range approximately 27 km south of Coree Flats. It is essentially a valley bog
surrounded on the slopes by subalpine woodland (E. pauciflora) underlain by a
well-developed and virtually continuous sward of snow grass (Poa caespitosa).
The bog is an example of the Carex gaudichaudiana-Sphagnum cristatum bog
alliance (Costin et al., 1959) of which a large part consists of active sphagnum
moss showing hollow-hummock development (Fig. 2). Most burrows of P.
corroboree were found in sphagnum bordering pools or in between pools.

The area here called Smiggin Holes is about 3.2 km north of the village of
Smiggin Holes on the Smiggin Holes-Guthega road at an elevation of about
1650 m. It is a small sphagnum bog surrounded by disclimax heath (Costin, et al.,
1959), which was originally subalpine woodland.

Boggy Plain (alt. 1585 m) is located about 5 km south-east of Smiggin
Holes on the same mountain range. As the name suggests, this was once a very
wet sphagnum bog but due to repeated burning off and grazing of sheep, it is
now very dry and there is little actively growing sphagnum.

Round Mountain is 39 km NNW of Smiggin Holes at an elevation of 1585 m.
It is a small bog surrounded by subalpine woodland. Little Thredbo River Flats

Aust. Zool. 18(1), 1973 17

PENGILLEY

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. Zool. 18(1),

Aust

18

PSEUDOPHRYNE BREEDING BIOLOGY

(alt. 1280 m) is located near the junction of the Little Thredbo and Thredbo
Rivers in the Thredbo Valley, approximately 7 km south of Smiggin Holes.
The area consists of a disturbed sphagnum bog surrounded by wet sclerophyll
forest.

The area here called Yarrangobilly is actually 2 km south-east of the village
of Yarrangobilly on Highway 18 at an elevation of approx. 1260 m. It is part
of the western montane zone of Costin (1954) and consists mainly of wet
sclerophyll forest.

Weather data for two highland localities (Smiggin Holes and Yarrangobilly )
and one tableland locality (Canberra) are shown in Fig. 3.

Smiggin Holas fees
m

min. o——a

TEMP. ( °C)

RAINFALL ( cms.)

° be
JAN FEB MAR APR MAY JUN JUL AUG SEPT OCT NOV DEC

Fig. 3. Upper Graph. Mean maximum and minimum monthly screen temperatures of
three selected stations in the Southern Highlands and Tablelands of N.S.W.
Lower Graph. Mean monthly rainfall in centimetres for the same three stations.

Rainfall and temperature data for Coree Flats for the years 1963 to 1965
have also been reported in another publication (Pengilley, 1971b). During the
early part of 1965, drought conditions prevailed over most of south-eastern
N.S.W. Very little rain was recorded at Coree Flats during the months of
January and February, and no rain was recorded in March. Less than 5 cms
of rain fell in each of the four subsequent months.

Aust. Zool. 18(1), 1973 19

R. PENGILLEY

A preliminary statement on the taxonomic status of the three species studied
has been given in an earlier publication (Pengilley, 1971a).

The number of eggs laid by each of the three species was determined (1) by
counting the number of eggs in the body cavity and oviducts of females that had.
ovulated, and (2) by counting eggs, which were of one embryological stage of
development, that were deposited in burrows. The snout-urostyle measurements
refer to those obtained from preserved adults which were collected during the
breeding season.

OBSERVATIONS

Size at Sexual maturity

Snout-urostyle lengths of males P. corroboree found in burrows during the
breeding season and of gravid females are given in Table 2. At any given locality,
males are generally a few millimetres smaller than females. There is a gradual
but parallel increase in the mean size of both males and females with increase
in altitude.

At Coree Flats, P. corroboree probably attains sexual maturity at 3 years.
This rather crude estimate of the length of the pre-reproductive period is based.
on size attained by juveniles at the end of first year of their terrestrial phase,
but in their second year of growth following fertilization. At this stage in the

TABLE 2:

Snout-urostyle lengths of adult male and adult female P. corroboree.
Southern Highlands, N.S.W.

MALES FEMALES
Mean _ Snout- Mean _ Snout-
| } Non inna urostyle length No. in urostyle length
Locality | Sample (mm) +SE? Sample (mm!) °==2SE
Coree Flats* et 22.0 + 0.30 57 24.6 + 0.26
(alt. 1040 m)
Bulls Head* — —- — — 8 25, 2D
(alt. 1220 m)
Snowy Flats* 53 Di 026 5 26.025 20-62
(alt. 1650 m)
Yarrangobilly | 20 IDG = U6 5 25-0 cep th
(alt. 1280 m) |
Smiggin Holest We IO POT od) Dearie 00 12 27.0 + 0.70
(alt. 1650 m) | |
Round Mountain? | 42 24.0 + 0.34 z, 27.6022 (O92
(alt. 1590 m)

* Localities in the Brindabella Range
7+ Localities in the Snowy Mountains
= SE = Standard error

20 Aust. Zool. 18(1), 1973

PSEUDOPHRYNE BREEDING BIOLOGY

life cycle, juveniles ranged from 7.5 to 10.0 mm in snout-urostyle length and
at the end of their first year of terrestrial growth they range from 9.0 to 14.0 mm.

P. dendyi males are also slightly smaller than females, but they tend to
be slightly larger than males of P. corroboree from the same altitude (Fig. 4),
but unlike the situation in P. corroboree, there is no clearly defined trend in
size with altitude. There are, however, some differences between localities differing
greatly in altitude over a short distance. For example, males from Boggy Plain
(alt. 1585 m) are larger than those from Little Thredbo River Flats (alt. 1158 m)
and similarly males from Clyde Mountain (alt. 610 m) on the Coastal Range
are significantly larger than those from near sea level at Moruya.

+ —— SS == =
MALES
Swampy aera
Plains R pee
(427m.)
Little Thredbo R wae N=25
(1158m.)
Boggy Plains N=46
(1585m.) ne
Eucembene R. ae N=44
(1440m.)
tong Plain ee a N=21
(1440m.)
Yarrangobilly ope N=47
(1280m.)
Coree Flats seme N=35
(1040m.)
Clyde Mtn. oe N=21
(610m.)
Moruya
(15m.)

FEMALES

Little Thredbo R.
(1158m.)

Long Plain
(1440m.)

Yarrangobilly
(1,280m.)

Coree Flats
(1040m.)

21 22 23 24 25 26 27 28 29
SNOUT-UROSTYLE LENGTH (mm )

Fig. 4. Snout-urostyle lengths of adult males and females of P. dendyi. Thin vertical
lines represent the means and the thicker horizontal bars represent the means + 2SE.

Both adult male and adult female P. bibroni (Table 3) are approximately
the same size as most of the Southern Highland populations of P. dendyi.
Number of eggs laid

P. corroboree

The clutch size of P. corroboree based on counts of eggs in gravid females
ranged from 16 to 38, the mean clutch sizes for the two samples (Snowy Mts.

Aust. Zool. 18(1), 1973 21

R: PENGILLEY

and Brindabella Range) being 26.4 and 24.0 respectively. The larger mean clutch
size of the Snowy Mountains females is attributable to the larger mean size
of these females (Table 2). However, from the small sample of 24 females.
obtained from the Snowy Mountains there is no relationship between snout-urostyle
length of the female and fecundity, although fecundity is correlated with size
in the sample from the Brindabella Range (Fig. 5).

TABLE 3.
Snout-urostyle lengths (sub) of adult male and female P. bibroni, Southern Tablelands,
N.S.W.
MALES FEMALES
Number in Mean SUL Number in Mean SUL
Locality Sample (mm) + 2SE Sample (mm), = 2SE
11.2 km. south 25 DAS ee Onis 5) 2536) eels
of Yass, N.S.W.
Mountain Creek 11 2328) FEN 0:8 2 24.2
Road, N.S.W.
Uriarra, A.C.T. 19 DAA rst OES 2 25.4
TABLE 4.

Percentage of nests of P. corroboree containing a given number of clutches*.

NGIGEE Percentage of nests containing a
Locality Near eects given number of clutches
examined 1 2 3 4 5
Coree Flats 1963 104 ASE Si Sd 5d 330, SO alae
1964 136 60/0) 34:35.) S30" % 0:6 020
1965 7 TIS 222 200 OO nO
1967 21 33.3-"525.0%) 23.0" 7 9:0 ONO
Snowy Flats 1964 83 ADT NASI O Ped oat et LU,
1965 14 PSA AS85630 OOF a OO
Smiggin Holes 1964 14 za Me A Ni ai (2 Pen OO OND
1965 58 O9n2, 029 Sk SeemneOLO |. nO.U
Round Mtn. 1965 28 OLB 3 852) OOOO a Ole

* For this purpose, a clutch consisted of less than 30 eggs at the same stage of
development. If two batches of eggs were at different stages of development, they were
considered to be two clutches even though each batch may have consisted of less than
15 eggs.

22 Aust. Zool. 18(1), 1973

PSEUDOPHRYNE BREEDING BIOLOGY

Colefax (1956) reported that P. corroboree (Snowy Mountains) lays 12 eggs.
It is evident from the counts of eggs in gravid females and the number of eggs
per nest (Figs. 6 and 7) that this number represents only part of the egg
complement of a female. A small percentage of burrows, generally less than 20%
do contain less than the minimum clutch size recorded. Most, however, contain
one or more clutches of eggs (Figs. 6 and 7 and Table 4) indicating that a
burrow is used by more than one female as an oviposition site.

P. dendyt

The only report in the literature on the number of eggs laid by P. dendyi
is that of Moore (1961) who found 85 eggs in a nest near Jindabyne, N.S.W.
The number of mature eggs contained in the ovaries of 30 gravid females ranged
from 35 to 102. There was no relationship between size of female and fecundity.

In 9 nests of P. dendyi at Coree Flats examined in 1963, the number of
eggs ranged from 45 to 154, the mean being 101.1. Examination of 36 nests
during the 1964 breeding season showed that modal clutch size was within the
range 80-89 (Fig. 8) and the mean was 81.8. In 1965, there was a reduction
in the modal (40-49) and mean (65.0) ‘clutch’ size.

NO. OF EGGS / FEMALE

21 22 23 24 25 26 27 28 29 36

SNOUT-UROSTYLE LENGTH (mms.)

Fig. 5. Plot of the number of eggs found per female against snout-urostyle length.
The open squares represent specimens obtained from Snowy Flats, Brindabella Range.

Aust. Zool. 18(1), 1973 23

R. ‘PENGILLEY

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Aust. Zool.

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24

PSEUDOPHRYNE BREEDING BIOLOGY

P. bibroni

This species is recorded as laying 130 (Harrison, 1922) and 260 eggs
(Jacobson, 1962). From 12 nests at Uriarra, A.C.T., the modal ‘clutch’ size was
between 80-99 (Fig. 8) the mean being 85.3. Since only three gravid females
were dissected, it is not known whether there is any relationship between body
length and egg complement. The clutch size of these females ranged from 82
tonOD: (Pies /9).

Calling and Breeding Seasons

Male P. corroboree have been heard calling from early December until early
April at Snowy Flats and from late December until mid-April at Coree Flats
(Table 5). Thus males are present in the breeding areas 4 to 8 weeks before
breeding commences and some males remain until 2 to 4 weeks following egg

Smiggin Holes

PERCENT FREQUENCY

PERCENT FREQUENCY

0-9 20-29 40-49 60-69 80-89 100-109 120-129
10-19 30-39 50-59 ~- 70-79 90-99 110-119 130-139 °

NO. OF EGGS

0-9 10-19 20-29 30-39 40-49 50-59 60-69 70-79 80-89 90-99
NO. OF EGGS

Fig. 6 Fig; 7

Fig. 6. Frequency distribution of the number of eggs found in nests of P. corroboree
at two localities on the Brindabella Range. The number of nests (N) and the year
during which they were examined are shown on the diagram.

Fig. 7. Frequency distributions of the number of eggs found in nests of P. corroboree
at two localities in the Snowy Mountains region. The number of nests (N) and the
year during which they were examined are shown on the diagram.

Aust. Zool. 18(1), 1973 25

R. PENGILLEY

laying. On the limited and qualitative information available, both migration of
males into and out of the breeding areas appears to be related to altitude.

The onset of breeding as evidenced by the presence in burrows of recently-
laid eggs or of eggs in the early stages of development is more clearly related
to altitude than is the initiation of thie vid iGRvios aialee nntea the breeding area.
Populations of P. corroboree in the sub-alpine zone (>1524 m) generally begin
breeding in the latter half of January, but those at lower altitudes (about
1000 m) do not commence until early March. This negative relation between
altitude and the commencement of egg laying appears to be consistent from
year to year (Fig. 10 and Table 5).

However, in 1965, breeding was much later at all localities except for
Round Mountain and Smiggin Holes in the Snowy Mountains when recently-
laid eggs were found on January 24 and 25. If allowances are made for errors

P. dendyi

a Little Thredbo R.

@ Long Plain

o Yarrangobilly

® Coree Flats

* P. bibroni

NO. OF EGGS / FEMALE

PERCENT FREQUENCY

1964

100-109 120-129 140-149 30
0-9 20- 40-49 60-69 80-89
10-19 ee ag 50-59 70-79 90-99 10-119 130-139 150+ 22 23 24 25 26 27 28 29

NO OF EGGS SNOUT-UROSTYLE LENGTH (mm )

Fig. 8 Fig. 9

Fig. 8. Upper Graph. Frequency distribution of the number of eggs found in nests of
P. dendyi.

Lower Granh. Frequency distribution of eggs found in 12 nests of P. bibroni at
Uriarra, A.C.T. during the 1964 breeding season.

Fig. 9. Plot of the number of eggs found in gravid females of P. dendyi and P. bibroni
against snout-urostyle length.

26 Aust. Zool. 18(1), 1973

PSEUDOPHRYNE BREEDING BIOLOGY

in estimating the date of commencement of breeding, the dates given do not
differ greatly from those of the previous year. At both Snowy and Coree Flats,
however, breeding was two to three weeks later than the previous year.

Colefax (1956) recorded P. corroboree breeding in the latter half of
December at Alpine Hut (alt. approx. 1768 m). If 1964 is considered to be a
normal season for P. corroboree, then from the relationship between altitude
and the onset of breeding, the Alpine Hut population would have started
breeding towards the end of December.

P. dendyi and P. bibroni

The breeding of P. dendyi and P. bibroni tends to follow the same pattern
as of P. corroboree in that the males migrate into the breeding areas some
weeks before breeding commences.

During 1965, males of P. dendyi were observed in the breeding areas towards
the end of January at both Boggy Plain and Little Thredbo River Flats, but
were not heard calling until early March at Coree Flats. By the second week
of April, the majority of males had moved out of the breeding area at Boggy
Plain, but this did not take place until at least early May at Coree Flats as
males were still present on April 26.

The close correspondence seen in breeding schedule and altitude of P.
corroboree was not so apparent in P. dendyi during 1964 and 1965. In 1964
at 1585 m breeding began towards the end of January and at 1219 m eggs were
first noticed towards the end of the second week in February. Delayed breeding
was also observed in 1965 when most populations bred approximately two weeks
later than in the previous year.

Male P. bibroni were first heard calling at 610 m on April 13, 1964, but as
these populations were not visited before this date, calling possibly could have
begun much earlier. Males were last heard calling towards the end of May of the
same year.

1500

1000

ALTITUDE (metres)

19652
Snowy Mins 1964 «
1955

7; 500

JAW FES MAR BPR JAN FEB MAR APR | PAR AFR MAY JUN

Fig. 10. Time of onset of breeding in P. corroboree, P. dendyi and P. bibroni at various
altitudes in the Southern Highlands and Southern Tablelands, N.S.W.

Aust. Zool. 18(1), 1973 ot

R. PENGILLEY

The data available for P. bibroni are too limited to indicate whether breeding
schedule is related to altitude. However, some observations made in 1963
suggest that breeding in this species is partially dependent on latitude. P. bibroni
was found breeding near Pt. Lookout (alt. approx. 1200 m) New England
Tablelands, N.S.W. on January 16, 1963. Three days later (January 19) in
the Blue Mountains, males were found calling in burrows but no eggs were
present. At Canberra, however, males were not heard calling until towards the
end of March.

DISCUSSION

Where P. corroboree and P. dendyi are sympatric, there is some overlap in
breeding seasons, with P. corroboree tending to breed much earlier than
P. dendyi. At Coree Flats, P. dendyi males begin to move into the breeding
area at the time when P. corroboree is just beginning to breed. There is always
the possibility then that P. dendyi males could mate with P. corroboree females.
However, the courtship calls of the two species are quite different (Pengilley,
197la) and this undoubtedly plays some part in enhancing reproductive isolation
between the two species.

Where P. corroboree and P. dendyi are allopatric there is some evidence
which suggests that there is greater overlap in breeding seasons. During 1964,
in the Snowy Mountains, P. dendyi began breeding about one week later
than P. corroboree, and if a correction is made for the difference in altitude between
the two localities, there is very little difference between the times at which they
bred. In 1965, however, the overlap was not as great as the time of breeding of
both species (at the same altitude, but at different localities) differed by three
weeks.

The drought of 1965 affected the breeding of all populations below 1560 m.
in one of two ways: breeding was either restricted to very wet places in the
breeding areas, or it was totally inhibited. No breeding of P. corroboree occurred
in the main study area at the southern end of Coree Flats. A total of seven nests
containing eggs were found, however, in two small areas bordering a small
stream in the northern and western edges of the wet heath. In this area, male
frogs were largely confined to the relatively moist, peripheral, ecotone of
Leptospermum myrtifolium and Epacris breviflora. Only a few males were
located in the central part which, in 1963 and 1964 had provided the most
favourable breeding sites. It was estimated that the area suitable for breeding
in this small part of the flat had been reduced to about 1% or less of the
1964 value.

At Coree Flats, P. dendyi was not as greatly affected by the drought as
P. corroboree because it bred much later in the year when climatic conditions
were less severe.

28 Aust. Zool. 18(1), 1973

PSEUDOPHRYNE BREEDING BIOLOGY

Populations of P. bibroni on the tablelands were also greatly affected by
the drought. Males migrated to the breeding areas at 5 localities but eggs were
found at only one of these.

At localities about 1500 m. on the Brindabella Range, and the Snowy
Mountains, all populations of P. corroboree and P. dendyi bred. However,
the breeding was not entirely successful as large numbers of eggs became desiccated
and died as a result of the sphagnum drying up.

The demonstration of well-defined seasonal breeding in these species of
Pseudophryne suggests that their reproductive activities are controlled by climatic
conditions. Temperature and rainfall are considered as being the two most
important climatic factors affecting reproduction in amphibia (Gallien, 1959;
Noble and Noble, 1923; Smith, 1955). Although experimental evidence is available
only for the effects of temperature (Van Oordt, 1960), there is much evidence
(e.g. Savage, 1961) which suggests that one or both factors are influential in
determining the onset of breeding. In P. corroboree and to a lesser extent in
P. dendyi, it appears that the breeding schedule is determined by temperature
but the precise thermal conditions necessary to induce breeding are not known,
primarily because of the lack of records from different localities extending over a
number of years. However, the rate of decline of temperatures during the period
January to May may be of some importance. At Coree Flats, breeding was much
later in 1965 than in 1964. In 1965, temperatures declined at the mean daily
rate of 0.06°C, whereas in 1964 the rate was 0.12°C. Robinson (1965, pers.
comm.) suggested that the slower rate observed in 1965, may have been responsible
for delayed breeding in the winter-breeding lyre bird, Menura novae-hollandiae at a
locality about 17 km. south of Coree Flats.

The rate of decline of temperatures is probably not the prime factor
responsible for breeding of Pseudophryne in the subalpine zone, because these
species begin breeding when temperatures are just beginning to decrease.

Although rainfall has been implicated as a stimulus for breeding in some species
of Pseudophryne (Fletcher, 1889; Jacobson, 1963; Main, 1965) it seems from
the data obtained on P. corroboree and P. dendyi that there is very little
relationship between the two. Both species breed from late summer to mid-
autumn, a period during which rainfall is generally minimal. The absence of
rain, however, can affect reproduction by making the breeding sites too dry
for egg deposition. In moist areas, at the same localities, breeding will occur.

REFERENCES

COLEFAX, A. N. (1956). New information on the corroboree frog (Pseudophryne
corroboree Moore) Proc. Linn. Soc., N.S.W. 80: 258-266.

FLETCHER, J. J. (1889). Observations on the ovivosition and habits of certain Australian
Batrachtans:: Proc.: Minn.) Soc. <oN:S.W:, 42 (2) * 357-387.

GALLIEN, L. (1959). Endocrine bases for renroductive adaptations in Amphibia.
Symposium on Comparative Endocrinology. A. Gorbman ed. 479-487.

Aust. Zool. 18(1), 1973 29

RR. PENGILLEY

HARRISON, L. (1922). On the breeding habits of some Australian Frogs. Aust. Zool.
3: 17-34.

JACOBSON, C. M. (1962). A study of developmental variation within the genus
Pseudophryne (Fitzinger). M.Sc. Thesis, Univ. of Sydney. 55pp.

LITTLEJOHN, M. J. (1963). Frogs of the Melbourne area. Vict. Nat. 79: 296-304.

MAIN, A. R. (1964). A new species of Pseudophryne (Anura: Leptodactylidae) from
North-Western Australia. West. Aust. Nat. 9: 66-72.

MAIN, A. R. (1965). Frogs of southern Western Australia. West. Aust. Nat. Club,
Perth. 73 pp.

MAIN, A. R., LITTLEJOHN, M. J. and LEE, A. K. (1959). Ecology of Australian
Frogs. In Biogeography and ecology in Australia. A. Keast, R. L. Crocker, and
C. S. Christian (eds.). W. Junk, The Hague. 396-411.

MOORE, J. A. (1962). The frogs of eastern New South Wales. Bull. Amer. Mus. Nat.
Hist. 1/2]: 149-386.

NOBLE, G. K. and NOBLE, R. C. (1923). The Anderson treefrog (Hyla andersoniv
Baird), observations on its habits and life history. Zoologica. 2: 416-455.

PENGILLEY, R. K. (1971la). Calling and associated behaviour of some species of
Pseudophryne (Anura: Leptodactylidae) J. Zool. Lond. 163: 73-92.

PENGILLEY, R. K. (1971b). The food of some Australian anurans. J. Zool, Lond.
163: 93-103.

SAVAGE, R. M. (1961). The Ecology and Life History of the Common Frog (Rana
temporaria temporaria). Pitman, Lond. 221 pp.

VAN OORDT, P. G. W. J. (1960). The influence of internal and external factors
in the regulation of the svermatogenetic cycle in Amphibia. Smyp. Zool. Soc. Lond.,
No. 2. Cyclic activity in Endocrine Systems. 29-51.

30 Aust. Zool. 18(1), 1973

Three New Species of Deep-dwelling Damselfishes
(Pomacentridae) from the South-West Pacific Ocean

GERALD R. ALLEN
Department of Ichthyology, The Australian Museum, Sydney, New South Wales, 2000.

ABSTRACT

Three new species of damselfishes are described which were collected
during 1971-1972 while conducting ichthyological investigations aboard the
research vessel E/ Torito at Madang, New Guinea; Fergusson Island,
D’Entrecasteaux Group; Osprey Reef, Coral Sea; and the Great Barrier Reef.
These species were taken in depths ranging from 31 to 50 metres. Abudefduft
starcki and A. caeruleolineatus appear to have no close relatives. The
remaining species, Pomacentrus nigromarginatus, belongs to the complex
which include P. melanochir, P. melanopterus, P. nigromanus and P.
philippinus. \t is most closely related to the latter species, but differs on the
basis of habitat preference and counts for the gill rakers and tubed lateral
line scales.

INTRODUCTION

The use of SCUBA (self-contained underwater breathing appartus) by marine
scientists has become increasingly popular. However, in spite of its widespread
application, there are few ichthyologists who regularly employ this equipment
below 100 feet, primarily because of the physiological hazards involved. During
the period extending from November 1971 to June 1972, I had the opportunity
to work with Dr. Walter A. Starck II aboard his 65-foot research vessel E/ Torito.
This ship was designed by Dr. Starck for support of deep-diving and is equipped
with both conventional SCUBA and closed circuit, electronically controlled, mixed
gas apparatus. Other equipment includes air compressors, two decompression
chambers, and a host of accessories designed for probing the deeper reefs. Our
itinerary included the following localities: Palau Archipelago, Western Caroline
Islands; Madang, Cape Nelson and Samarai Island, along the north-east coast of
Papua New Guinea; Goodenough, Fergusson and Normanby Islands, D’Entrecas-
teaux Group; Egum Atoll, Solomon Sea; Osprey Reef, Coral Sea; portions of
the Great Barrier Reef off Cairns, Queensland. A significant amount of our
collecting effort was expended between depths of 100 and 250 feet. The collections
were made with rotenone, quinaldine, dipnets and small multi-prong spears. They
contain at least 20 new species including three species of deep-dwelling pomacen-
trids which are described below.

Aust. Zool. 18(1), 1973 31

GERALD R. ALLEN

METHODS OF COUNTING AND MEASURING

The methods of counting and measuring are the same as those described in
Allen (1972) except the length of the dorsal and anal spines are measured proxi-
mally at the base of the spine rather than the point at which the spine emerges
from the scaly sheath. Measurements were made with needlepoint dial calipers to
the nearest one-tenth millimetre (mm). Standard length is abbreviated as SL.
The fraction = which appears in the dorsal and anal fin ray formulae refers to a
bifurcate conditon of the last ray.

The counts and proportions which appear in parentheses under the description
section for each species apply to the paratypes when differing from the holotype. A
summary of counts for the dorsal, anal and pectoral fin rays, gill rakers on the first
gill arch, and tubed lateral line scales are presented in Tables 1 and 2.

Type material has been deposited at the following institutions: Australian
Museum, Sydney (AMS); Bernice P. Bishop Museum, Honolulu (BPBM); British
Museum (Natural History), London (BMNH); Museum National d’Histoire
Naturelle, Paris (MNHN); United States National Museum, Washington, D.C.

(USNM ).

DESCRIPTIONS

Abudefduf starcki, new species
Figure 1

Holotype — AMS I.16477-001, 44.7 mm SL, collected with multi-prong spear on outer
slope of Osprey Reef, Coral Sea (13°58’S, 146°41V’E; U.S.H.O. Chart No. 2002) in 37

metres by G. R. Allen on June 22, 1972.

TABLE, (1

Dorsal and Anal Fin Ray Counts for New Pomacentrids from Papua New Guinea and
the Coral Sea

DORSAL RAYS ANAL RAYS
SPECIES XIII XIV 11 114 12 123 13 1331414315 II 13 13414 14315
Abuacfdus starck) 5% Geren 2 2 2 1 yg
As caeruleolineatus -0. Se 10 1 Sica 10: 22 iO owe
Pomacentrus nigromarginatus 5 De 12 ae
TABLE 2

Pectoral Ray, Gill Raker, and Tubed Lateral Line Scale Counts
for New Pomacentrids from Papua New Guinea and the Coral Sea.

Pecoral Rays Gill Rakers) Tubed Lateral Line Scales
SPECIES 15 1647, ZO: 2 22S 12213214" 153.46

2

Abudefdife Marcki A 1 treet
A: CGCTNCOINEOIUS. Gio Be ee 149 Be yh: At DF ty ad A oa
Pomacentrus nigromarginatus .......... DAN 1 4 1 4

32 Aust. Zool. 18(1), 1973

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GERALD R. ALLEN

Paratype — AMS 1.16478-001, 36.0 mm SL, same collecting data as holotype.

Diagnosis A species of Abudefduf with the following combination of characters:
Dorsal spines 13; horizontal scale rows between middle of lateral line and dorsal fin base
14; scales on upper part of head reaching to nostrils; teeth uniserial; preorbital and suborbital
naked; body mostly blue with mid-dorsal strive, dorsal fin and caudal fin yellow.

Description — The proportional measurements: for the type specimens are expressed
as percentage of the standard length in Table 3.

Dorsal rays XIII,14; anal rays II,14 (11,15); pectoral rays 17 (16 on right side of
paratype); pelvic rays I,5; branched caudal rays 13; gill rakers on the first gill arch 22 (21)
with birfurcate raker at angle of gill arch; tubed lateral line scales 15 or 16 (damaged on
both specimens); vertical scale rows from upper edge of gill opening to base of caudal fin
28: horizontal scale rows from base of dorsal fin to middle of lateral line (exclusive of dorsal
base sheath scales) 12; from lateral line to anal fin origin 9; predorsal scales about 20
extending well forward of anterior edge of orbits, about to levei of nostrils; teeth uniserial,
close-set, somewhat incisform with rounded tips, about 36 to 38 in the lower jaw and about
48 to 5O in the upper.

Body ovate and laterally compressed, the greatest depth 2.2 in the standard length.
Head profile conical, the head length contained 3.4 (3.3) times in the standard length;
snout 4.9 (5.2), eye diameter 2.3 (2.2), interorbital width 3.8 (4.6), least depth of caudal
peduncle 2.1 (2.3), length of caudal peduncle 2.5 (2.7), of pectoral fin 1.0 (0.9), of pelvic
fin 1.0, of middle caudal rays 1.3 (1.4), all in the head length

‘EABBE..3

MORPHOMETRIC PROPORTIONS (IN % OF SL) OF TWO SPECIMENS
OF ABUDEFDUF STARCKI FROM OSPREY REEF, CORAL SEA

Morphometric measurement N Range (% SL) Mean (% SL)
Standard length (SL) 2 36.0-44.7 —
Body depth 2 45.0-46.1 45.5
Head length 2, 29.4-30.4 29.9
Snout length 2 5.8-6.1 a
Eye diameter 2 12.8-13.9 is
Interorbital width DE 6.7-7.9 73
Least depth of caudal peduncle 2 13.1-13.9 13.5
Length of caudal peduncle Z 11.1-11.6 H1e3
Snout to origin of dorsal fin 2 32.5-33.4 32.9
Snout to origin of anal fin 2 62.4-62.8 62.6
Snout to origin of pelvic fin 4 40.0-40.0 40.0
Length of dorsal fin base Zz 60.8-63.0 61.9
Length of anal fin base z 31.1-33.0 32.0
Length of pectoral fin 2 29.5-32.3 30.9
Length of pelvic fin 2 29.1-30.6 29.8
Length of pelvic spine g) 17.5-20.0 18.7
Length of Ist dorsal spine 1 —- 6.6
Length of 6th dorsal spine 2 15.0-16.7 15.8
Length of last dorsal spine p) 15.0-16.4 LY,
Length of longest soft dorsal ray Z, 19.7-20.3 20.0
Length of Ist anal spine 2 7.6-8.1 7.8
Length of 2nd anal spine 2 12.3-13.4 12.8
Length of longest anal ray 2 20.0-22.0 210
Length of middle caudal rays js Deo 2 33 22.8

34 Aust. Zool. 18(1), 1973

NEW SPECIES OF DAMSELFISHES

Single nasal opening on each side of snout, difficult to distinguish from surrounding
sensory pores; mouth oblique, terminally located; lateral line gently arched beneath dorsal
fin, terminating below base of first few soft dorsal rays; preorbital, most of suborbital
(2-3 scales on posterior portion), tip of snout, lips, chin, and isthmus naked, remainder of
head and body scaled; most of head scales cycloid, remainder of scales finely ctenoid;
preopercle with two horizontal scale rows with additional row of smaller scales on inferior
limb; small sheath scales covering about basal 4 of membraneous portions of dorsal, anal
and caudal fins; lower margin of preorbital and suborbital entire; margin of preopercle and
opercle entire.

Origin of dorsal fin at level of second tubed lateral line scale; spines of dorsal fin
gradually increasing in length to sixth spine, remaining spines about equal; length of first
dorsal spine 6.6, of sixth and thirteenth dorsal spines 2.0 (1.9), of longest soft dorsal ray
1.5, of first anal spine 3.9 (3.8), of second anal spine 2.4 (2.3), of longest soft anal ray
1.3 (1.5), all in the head length; caudal fin emarginate; pectoral fins pointed.

Colour of holotype in alcohol: Ground colour of body dark blue grading to blackish
posteriorly; broad pale yellow mid-dorsal stripe extending anteriorly from origin of soft
dorsal fin to forehead, covering most of region above lateral line; head dusky, paler
ventrally; spinous dorsal fin pale with fine black margin; soft dorsal fin blackish basally,
pale distally with fine black margin; caudal fin and upper and lower portions of caudal
peduncle pale; pelvic and anal fins blackish; pectoral fins pale with blackish basal bar.

The paratype is coloured similarly except the interorbital, snout, and ventral portion
of the head are less dusky.

Colour of holotype and paratype in life: Most of body royal blue, broad mid-dersal
stripe of canary yellow extending anteriorly from origin of soft dorsal fin to tip of snout,
also engulfing spinous dorsal fin and distal half of soft dorsal fin; .¥entral
portion of head yellow; most of opercle, posterior portion of preopercle and
suborbital royal blue; basal half of soft dorsal fin, anal fin, and pelvic fins bluish+black;
caudal fin and upper and lower portions of caudal peduncle canary yellow; pectoral fin
pale yellow with blackish basal bar.

Remarks—A. starcki does not appear to have any close relatives. The
combination of characters listed under the diagnosis in addition to the colour pattern
are distinctive. About 15 specimens were encountered in the vicinity of the
type locality in depths ranging from 35 to 52 metres. The type locality consisted
of a steep slope intersected by sand channels. Individuals of A. starcki were seen
around rocky outcrops and near crevices on the reef slope and also around outcrops
in the sand channels. They typically feed on plankton a short distance above the
bottom. The stomach contents of the types consisted of 99 per cent a
with a small quantity of gastropod fragments and polychaetes.

Named starcki in honour of Dr. Walter A. Starck II, who first pointed out
the species to me while diving at Osprey Reef.

Abudefduf ca¢ruleolineatus, new species
Figure 2

Holotype — USNM 20741, 34.9 mm SL, collected with mali and dipnets on
outer reef slope off middle of Kranket Island (5°12’00”S, 145°50°52”E; Aus. Chart No.
648), about 1.5 nautical miles north of Madang, New Guinea in 40 metres by .G.: R. Allen
on April 9, 1972.

Paratypes — AMS I.16480-001, 2 specimens, 36.4 and 37.4 mm SL, collected); with
multi-prong spear on outer slope of Osprey Reef, Coral Sea in 50 metres by W. A.‘Starck II
on June 23, 1972; BMNH 1972.8.14.4-5, 2. snecimens, 30.1,,and 37.4-mm SL, same collecting

Aust. Zool. 18(1), 1973 35

GERALD R. ALLEN

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Aust. Zool. 18(1), 1973

NEW SPECIES OF DAMSELFISHES

data as the holotype; BPBM 13227, 2 specimens, 31.8 and 34.9 mm SL, same collecting
data as the holotype except collected on April 4, 1972; MNHN 1972-87, 2 specimens,
32.5 and 36.3 mm SL, same collecting data as the holotype except collected on April 4,
1972; USNM 207942, 36.3 mm SL, same collecting data as the holotype.

Diagnosis — A species of Abudefduf with the following combination of characters.
Dorsal spines 14; pectoral rays usually 16; tubed lateral line scales 12 to 15; horizontal
scale rows between middle of lateral line and dorsal fin base 12; predorsal scales extending
to mid-interorbital; head scales cycloid; teeth uniserial; greatest body depth 2.3 to
2.7; colour bright yellow with iridescent blue strive above lateral line, extending from anterior
portion of soft dorsal fin to snout.

Description — The proportional measurements for the type specimens are expressed
as percentage of the standard length in Table 4.

Dorsal rays XIV, 122 (XIV, 112 to 13); anal rays II,132 (11,13 to 14); pectoral rays
16 (one paratype with 15); pelvic rays I,5; branched caudal rays 13; gill rakers on the
first gill arch 21 (20 to 23, usually 22); tubed lateral line scale 15 (12 to 15, usually 13
or 14); vertical scale rows from upper edge of gill onening to base of caudal fin 28 (27
to 28); horizontal scale rows from base of dorsal fin to middle of lateral line (exclusive
ef dorsal base sheath scales) 12, from lateral line to anal fin origin 8 (7 to 8); predorsal
scale about 12 (about 10 to 14), extending to about mid-interorbital; teeth uniserial, conical
with rounded tips, about 40 to 44 in each jaw.

TABLE 4

MORPHOMETRIC PROPORTIONS (IN % OF SL) OF TEN SPECIMENS
OF ABUDEFDUF CAERULEOLINEATUS FROM MADANG, NEW GUINEA AND
OSPREY REEF, CORAL SEA

Morphometric measurement N Range (% SL) Mean (% SL)
Standard length (SL) 10 30.1-37.4 _
Body depth 10 37.7-44.0 40.9
Head length 10 29.1-32.9 31.6
Snout iength 10 4.7-7.5 6.3
Eye diameter 10 12.0-14.0 13.1
Interorbital width 10 6.9-8.4 7.6
Least depth of caudal peduncle 10 14.1-16.1 14.9
Length of caudal peduncle 10 2-129 10.2
Snout to origin of dorsal fin 10 36.0-40.0 38.4
Snout to origin of anal fin 10 64.9-71.5 67.2
Snout to origin of pelvic fin 10 37.0-44.8 40.4
Length of dorsal fin base 10 54.8-63.8 59:6
Length of anal fin base 10 24.5-26.3 Ps
Length of pectoral fin 10 28.9-35.8 32.0
Length of pelvic fin 10 219-2534 23.4
Length of pelvic spine 10 14.0-15.2 14.8
Length of ist dorsal spine 10 4.4-6.2 4.7
Length of 7th dorsal spine 10 8.7-13.2 11.6
Length of last dorsal spine 10 10.3-13.2 P2 ek
Length of longest soft dorsal ray 10 17.3-20.4 19.0
Length of Ist anal spine 10 4.7-6.3 a7.
Length of 2nd anal spine 10" 11.4-15.2 13.3
Length of longest anal ray 10 15.0-19.6 179
Length of middle caudal rays 10 20.9-26.7 22.6

Aust. Zool. 18(1), 1973 37

GERALD R. ALLEN

Body relatively elongate and laterally compressed, the greatest depth 2.4 (2.3 to 2.7)
in the standard length. Head profile rounded, the head length contained 3.2 (3.0 to 3.4)
times in the standard length; snout 4.3 (4.6 to 6.9), eye diamteer 2.3 (2.2 to 2.7), inter-
orbital width 3.9 (3.8 to 4.5), least depth of caudal peduncle 2.2 (2.1 to 2.3), length of
caudal peduncle 3.4 (2.3 to 3.1), of pectoral fin 1.0 (0.9 to 1.1), of pelvic fin 1.3 (1.3 to
1.5), of middle caudal rays 1.5 (1.2 to 1.5), all in the head length.

Single nasal opening on each side of snout; mouth oblique, terminally located; lateral
line gently arched beneath dorsal fin, terminating one scale row below base of 13th (11th
to 13th) dorsal spine; preorbital, suborbital, snout, lips, chin, isthmus and anterior portion
of interorbital naked; remainder of head and body scaled: scales of the head region cycloid,
those of the body finely ctenoid except most of scales above lateral line which are cycloid;
preopercle with two horizontal scale rows with additional row of smaller scales on inferior
limb; small sheath scales covering about basal 4 to 73 of membraneous portions of dorsal,
anal and caudal fins; margin of preorbital, suborbital, preopercle and opercle entire.

Origin of dorsal fin at level of fourth tubed lateral line scale; spines of dorsal fin
gradually increasing in length to about seventh svine, remaining spines about equal in
length; length of first dorsal spine 5.0 to 7.4 (paratypes only, holotype damaged), of seventh
dorsal spine 2.6 (2.4 to 3.8), of last dorsal snine 2.7 (2.4 to 3.2), of longest soft dorsal
ray 1.8 (1.5 to 1.9), of first anal spine 5.5 (4.2 to 7.1), of second anal spine 2.1 (2.2 to
2.8), of longest soft anal ray 1.7 (1.6 to 2.0), all in the head length; caudal fin
emarginate; pectoral fins pointed.

Colour of holotype in alcohol: Head and body entirely pale yellowish except black-
edged bluish stripe, about 4 eye diameter in width extending forward from base of
anteriormost soft dorsal rays to tip of snout (passing through dorsal half of eye), and
covering most of dorsal portion of body above lateral line; mid-dorsal region between stripe
of each side including spinous dorsal fin brownish; remainder of fins pale; small black
spot superiorly at pectoral base.

Colour of holotype in life: Head and body entirely yellow except black-edged iridescent
blue stripe above lateral line; mid-dorsal region between stripe of each side dusky; spinuous
dorsal fin bluish, remainder of fins yellow; small black spot superiorly at pectoral base.

Remarks — A. caeruleolineatus does not appear to have any close relatives.
The dorsal spine count of 14 is a rare feature among the Abudefduf. The colour
pattern of A. caeruleolineatus is very similar to that of A. leucopomus Lesson
as figured by Jordan and Seale (1906, colour plate XLIII, fig. 1) except it lacks
the small black spot at the base of the posteriormost soft dorsal rays and the
larger triangular blotch at the base of the caudal fin. A. leucopomus is a very
common species throughout the western tropical Pacific and is restricted to shallow
surge areas.

A. caeruleolineatus was the deepest dwelling member of the genus which
we observed. It lives on outer exposed reef slopes among rubble or small rocky
outcrops which are usually situated in sandy areas at depths ranging from 38 to
at least 62 metres. At Madang, New Guinea, it was moderately common in
such areas. It was sighted, but not collected, at 60 metres depth on the outer
reef off Seymour Bay, Fergusson Island, D’Entrecasteaux Group.

The name caeruleolineatus refers to the distinctive blue stripe on the upper

back.

Pomacentrus nigromarginatus, new species
Figure 3

Holotype — USNM 207937, 57.3 mm SL, collected with aquinaldine and dipnets on
outer reef slope off north end of Kranket Island (5°11’24’S, 145°50’54”E, Aus. Chart No.

38 Aust. Zool. 18(1), 1973

NEW SPECIES OF DAMSELFISHES

648), about two nautical miles north of Madang, New Guinea, in 35 metres by G. R. Allen
On eApril17e1972.

Paratypes — AMS I.16655-001, 57.1 mm SL, collected with multi-prong spear at
Thedford Reef, Great Barrier Reef, off Cairns, Queensland, Australia, in 30 metres by
G. R. Allen on August 15, 1972; BMNH.1972.8.14.3, 60.2 mm SL, same collecting data
as the holotype except collected with multi-prong spear; BPBN 13226, 2 specimens, 49.6
and 50.5 mm SL, collected with multi-prong spear on isolated pinnacle reef about two
nautical miles off south-west side of Seymour Bay, Fergusson Island, D’Entrecasteaux Group
in 35 metres by G. R. Allen on May 29, 1972; USNM 207938, 59.9 mm SL, same collecting
data as the holotype except collected by W. A. Starck II with multi-prong spear in 31
metres on April 4, 1972.

Diagnosis — A species of Pomacentrus with the following combination of characters:
dorsal spines 13; preorbital and suborbital naked; slight notch between preorbital and
suborbital; inferior edge of suborbital entire; teeth biserial; tubed lateral line scales usually
15; colour basically medium-grey with black margin about 3 to % pupil diameter on
posterior edge of dorsal and caudal fins; base and axil of pectoral fin black.

Description — The proportional measurements for the type specimens are expressed
as percentage of the standard length in Table S.

Dorsal rays XIII,15 (XIII,144 to 15); anal rays 1,142 (11,142 to 15); pectoral
rays 16 on right side, 15 on left side (16 to 17); velvic rays, 1,5; branched caudal rays
13; gill rakers on the first gill arch 21 (20 to 21); tubed lateral line scales 14 on right

TABLES

MORPHOMETRIC PROPORTIONS (IN % OF SL) OF FIVE SPECIMENS
OF POMACENTRUS NIGROMARGINATUS FROM MADANG, NEW GUINEA AND
FERGUSSON ISLAND, D’ENTRECASTEAUX GROUP

Morphometric measurement N Range (% SL) Mean (% SL)
Standard length (SL) 5 49.6-60.2 —
Body depth 5 46.8-48.1 47.4
Head length 5 28.5-30.0 29.4
Snout length 5 6.1-6.9 6.5
Eye diameter 5 10.7-11.5 11.0
Interorbital width 5 7.0-8 .4 7.6
Least depth of caudal peduncle 5 13.7-15.0 14.4
Length of caudal peduncle 5 8.5-10.7 9.0
Snout to origin of dorsal! fin 5. 36.2-40.9 37.8
Snout to origin of anal fin 5 65.6-68.0 66.3
Snout to origin of pelvic fin 5 37.8-39.8 . 39.0
Length of dorsal fin base 5 62.0-66.6 63.7
Length of anal fin base 5 28.1-29.7 28.6
Length of pectoral fin 5 31.4-34.5 S865
Length of pelvic. fin 5 29a 31,2
Length of pelvic spine 5 16.1-18.0 16.7
Length of 1st dorsal spine 5 6.3-9.4 7.4
Length of last dorsal spine 5 15.2-16.7 16.1
Length of longest soft dorsal ray 5 20.9-26.0 23.6
Length of Ist anal spine 5 6.3-8.6 7.4
Length of 2nd anal spine 5 15.1-17.4 16.5
Length of longest anal ray 5 19.0-22.4 20.6
Length of middle caudal rays 5 22.2-25.3 23.4

Aust. Zool. 18(1), 1973 39

GERALD R. ALLEN

inéea.

tus, holotype, 57.3 mm SL, from Madang, New Gu

igromargina

Pomacentrus n

Bige 3,

40 Aust. Zool. 18(1), 1973

NEW SPECIES OF DAMSELFISHES

side, 13 on left side (15, excevt 17 on left side of one paratype); vertical scale rows
from upper edge of gill opening to base of caudal fin 28; horizontal scale rows from
base of dorsal fin to middle of lateral line (exclusive of dorsal base sheath scales) 12;
from lateral line to anal fin origin 9; predorsal scales about 20 (about 18 to 22), extending
well forward of anterior edge of orbits, about to level of nostrils; teeth biserial at front
of jaws, remainder uniserial, conical in shape with gently rounded tips, about 32 in the
lower jaw and 42 in the upper.

Body ovate and laterally compressed, the greatest depth 2.1 in the standard length.
Head profile conical, the head length contained 3.4 (3.3 to 3.5) times in the standard
length; snout 4.8 (4.4 to 4.6), eye diameter 2.7 (2.5 to 2.8), interorbital width 3.5
(3.7 to 4.3), least depth of caudal peduncle 2.0 (1.9 to 2.2), length of caudal peduncle
2.8 (3.4 to 35), of pectoral fin 0.9, of pelvic fin 0.9 (0.9 to 1.0), of middle caudal rays 1.2
(1.2 to 1.3), all in the head length.

Single nasal ovening on each side of snout; mouth oblique, terminally located; lateral
line gently arched beneath dorsal fin, terminating 134 scale rows below base of posteriormost
dorsal spines; preorbital, suborbital, tip of snout, lips, chin and isthmus naked; remainder
of head and body scaled; scales finely ctenoid; vreopercle with 2 or 3 transverse scale rows
with additional row of scales on inferior limb; small sheath scales covering about basal 3 to
*%z of membraneous portions of dorsal, anal and caudal fins; margin of preorbital and
suborbital entire except for very slight denticulations on two of the paratypes; posterior
margin of preopercle denticulate; posterior margin of opercle entire.

Origin of dorsal fin at level of third tubed lateral line scale; spines of dorsal fin
gradually increasing in length to last spine; length of first dorsal spine 3.9 (3.2 to 4.7),
of last dorsal spine 1.8 (1.7 to 1.9), of longest soft dorsal ray 1.2 (1.1 to 1.4), of first
anal spine 4.2 (3.5 to 4.5), of second anal spine 1.8 (1.7 to 1.9), of longest soft anal
ray 1.4 (1.3 to 1.5), all in the head length; caudal fin emarginate; pectoral fins pointed.

Colour of holotype in alcohol: Head and body greyish-brown with reddish suffusion,
scale margins and dorsal portion of body slightly darker; spinous dorsal greyish with
translucent submarginal band on outer half of fin; basal half of soft dorsal and caudal
fins greyish-brown, outer half ovale grey to _ tramslucent; dorsal and _ caudal
fins with black margin, about 3 to + pupil diameter, especially prominent on posterior edges
of soft dorsal and caudal fins; pelvic fins pale; pectoral fins pale with prominent black
spot covering entire base and axil of fin.

The paratypes are lighter, nearly whitish on the ventral portion of the head and body,
and their overall colouration lacks the reddish suffusion of the holotype.

Colour of holotype in life: Head and body pale grey, whitish ventrally; scale margins
brownish, but not detracting from the overall pale colouration of the body; prominent black
spot covering pectoral fin base and axil; fins greyish with prominent black margin on
edge of dorsal and caudal fins.

Remarks—P. nigromarginatus belongs to an Indo-Australian Archipelago
species complex which includes P. melanochir Bleeker, P. melanopterus Bleeker,
P. nigromanus Weber, and P. philippinus Evermann and Seale. These species share
similar counts and possess a prominent spot which covers the base and axil of the
pectoral fin. P. nigromarginatus is clearly separable from melanochir, melanopterus
and nigromanus on the basic of colour pattern and number of tubed lateral line
scales. The latter group of species usually have 17 to 19 tubed scales compared
with a modal count of 15 (holotype with 13 to 14) for P. nigromarginatus.
P. melanochir is further separable by the presence of a strongly denticulate sub-
orbital and one or more spinules on the rear margin of the opercle. P. melanopterus
usually has a few weak denticulations on the suborbital and a flattened spinule

Aust. Zool. 18(1), 1973 41

GERALD R. ALLEN

on the upper edge of the opercular margin. The suborbital and opercle are
virtually entire in P. nigromarginatus.

P. nigromarginatus appears to be most closely related to P. philippinus with
regards to morphology and colour pattern. They differ appreciably, however, in
number of tubed lateral line scales and total gill rakers on the first arch.
Furthermore, the suborbital width of philippinus fits about 4.5 to 5.0 in the width
of the bony orbit compared to about 6.0 to 7.0 for nigromarginatus. The suborbital
of philippinus also differs by possessing scales on its posterior half. There appears
to be a great deal of latitude in the colour pattern of philippinus, but the species
is invariably much darker than nigromarginatus. According to Montalban (1927),
specimens from the Philippines are predominantly blackish with pale streaks
on the scales. The caudal fin and posterior portions of the dorsal and anal fins are
yellow. Specimens of philippinus collected by the author at the D’Entrecasteaux
Group, Egum Atoll (Solomon Sea, near Woodlark Island), and Osprey Reef
(Coral Sea) were basically dark brownish with a red-orange suffusion, particularly
noticeable on the ventral surface. The caudal and posterior portion of the dorsal
and anal fins were also red-orange and had blackish margins similar to those of
P. nigromarginatus. Specimens of P. philippinus collected at Pixie Reef, on the
Great Barrier Reef, off Cairns, Queensland, Australia were entirely blackish,
including all the fins except the pectorals, and had pale streaks on the scales.
P. nigromarginatus and P. philippinus are further separable on the basis of
ecology. Both species occur on exposed outer reef slopes. However, P. philippinus
lives in depths ranging from 1.5 to 12 metres, usually around coral outcrops
or in the shadows of overhanging cliffs and ledges. P. nigromarginatus is usually
restricted to depths ranging from 30 to at least 46 metres and frequents rocky
outcrops which are usually situated on sandy slopes. It was common in this
habitat at Madang, Fergusson and Normanby Islands, Egum Atoll, and Osprey Reef.
The stomach contents of one of the paratypes consisted mainly of copepods and algal
fragments.

The name migromarginatus refers to the characteristic black margin on the
dorsal and caudal fins.

ACKNOWLEDGMENTS

I would like to thank Dr. Walter A. Starck II for making possible my participation
on the “El Torito” cruises of 1971-1972. Dr. John E. Randall provided the initial stimulus
for my studies of Indo-Pacific damselfishes under National Science Foundation Grant
GB-8732. Thanks are also due my wife, Connie, who typed the manuscript.

REFERENCES
ALLEN, G. R. (1972). The anemonefishes, their classification and biology. T.F.H.
Publications, Inc., New Jersey, 288 pp., 140 figs.

JORDAN, D. S. and SEALE, A. (1906). The fishes of Samoa. Bull. Bureau of Fisheries
(U:S.); 25, 173-455, 21: plis., 211 text-figs:

MONTALBAN, H. R. (1927). Pomacentridae of the Philippine Islands. Monogr. Bur. Sci.
Manila, P. I., 24, 117 pp., 19 pls.

42 Aust. Zool. 18(1), 1973

Aspects of Reproduction in the Whiptail Wallaby,
Macropus parryi

G. M. MAYNES

School of Biological Sciences, Macquarie University, North Ryde,
New South Wales, 2113.

The whiptail wallaby, Macropus parryi Bennett, 1835, is a large slender
macropodid which is common in Queensland and locally common in northern
New South Wales. Relatively little is known about its biology. Calaby (1966)
briefly reported on the habitat, habits and numbers of this species in the Upper
Richmond and Clarence Rivers area of northern New South Wales. Clark and
Poole (1967) mentioned in passing that the whiptail wallaby resembled the
grey kangaroo, Macropus giganteus, in not having a post-partum oestrus. Calaby
and Poole (1971) quoted unpublished data of Merchant for the length of oestrous
cycles, gestation periods and pouch life. However, no data are available to indicate
whether embryonic diapause occurs in the species.

This paper reports data on reproduction obtained from a small sample of
whiptail wallabies collected in September, 1971 at Coombadjha (29° 21’ S,
152° 31’ 42” E) on the Clarence River about a mile from its junction with the
Mann River. The habitat was partly cleared, dry sclerophyll forest essentially the
same as described and figured by Calaby (1966). The sample of M. parryi was
collected to investigate the mode of inheritance of the enzyme phosphoglycerate
kinase (P.G.K.) and to further studies on X chromosome inactivation in macro-
podids. These results are reported elsewhere (VandeBerg, Cooper and Sharman,
1973). In addition some data were collected on reproduction and are reported here.

Half of each ovary and a piece of uterus were collected from all females
carrying pouch young. These were fixed in Bodian’s solution, embedded in paraffin,
sectioned at 7 or 8 » m and stained in haematoxylin and eosin. A blastocyst
was recovered from female 27 (Table 1) using Sharman’s (1955) method.
Standard body measurements were recorded for all animals and weights and
pouch depth or scrotal size were recorded for all adults and juveniles. A total of
41 adults and juveniles was collected.

18 breeding females ranged in weight from 8.5 kg (molar eruption stage
MII.O, (Maynes, 1972) primiparous) to 12.8 kg (multiparous); while 12 mature
males ranged from 13.4 kg (almost MII.2) to 25 kg.

Table 1 gives the body measurements of the pouch young of 16 females
taken during the period 23-27 September, 1971. Limited data on growth of pouch
young in captivity and comparison with the growth of pouch young of other

Aust. Zool. 18(1), 1973 43

G. M. MAYNES

species (M. parma (Maynes, 1972); M. eugenii (Murphy and Smith, 1970)
and Megaleia rufa (Sharman, Frith and Calaby, 1964) ), suggests a range in
age from about 1 week to about 35 weeks. Thus the whiptail wallaby would
appear to breed at least from January to September. The data do not exclude the
possibility that they breed year round.

TABLES?

Body measurements of pouch young of whintail wallaby (all measurements in mm)

Parent No. Sex of P.Y. Head Ear Arm Leg Foot Body Tail

30 d 9.4 2 Set 3.0 4.1 25.8 7.2
34 2 ibe 5) Site! eek 14.0 10.1 56.1 Dee
54 d 20.0 6.6 B23 15 11.8 65.6 QZ
a2 2 Sad We 6.8 12.9 18.0 12.8 67.6 32.6
16 d 28.9 9.9 18.2 28.1 20.9 93:9 45.4
9 d 34.8 11.4 2223 37.0 28.2 109 68.2
46 2 36.1 Ley PHI 39.0 28.5 106.1 62.5
5 2 36.1 13.4 DBS 43.2 32.4 big 76.0
42 d 38.5 13:3 23.8 40.2 31.2 112.4 69.2
af d 48.1 17.8 31.0 S57 42.9 ef 90
48 2 48.0 18.9 BME 64.5 50.5 145 100.8
7 d 49.5 17.8 S077 3922, 45.2 145 B3
14 2 S221 ZAsd 34.4 Fi bas | 56.5 162 104
23 2 70.6 38.7 57.0 132 107 236 216
19 ) 73:3 375 58.5 130 111 245 211
27 2 91.6 73.6 83.9 196 168 348 393%.

Examination of the ovaries and uteri of those females carrying a pouch
young showed that all except female 27 were anoestrous. The uterine glands
of anoestrous females were small and scattered with little or no lumen present.
They consisted of low columnar epithelial cells with oval nuclei occupying about
half to two-thirds of the cell. The uterine lumen was lined with cuboidal epithelial
cells having mainly oval nuclei which occupied most of the cell. A few females
had large atretic follicles up to 1 mm diameter. Female 19 had a large non-atretic
follicle of 1.5 mm diameter.

Female 27 had a blastocyst, with an external diameter of 257 » m (measured
after fixation), present in the right uterus. Tall pseudostratified columnar epithelial
cells with mainly oval, basally-situated nuclei lined the uterine lumen. The uterine
glands had tall columnar cells with rounded nuclei. The lumina of most glands
were open and had some material present in them (Fig. 1). There was no evidence
of either coagulated semen or sperm in the uterine glands or the uterine lumen.
Neither was there any sign of mitoses in the uterine glands or lining. The right
ovary had a large corpus luteum with large luteal cells possessing rounded nuclei
(Fig. 2). No mitoses were observed in the luteal cells. It is concluded that this
female was either in a quiescent state (Clark and Poole, 1967) or just coming
out of it.

44 Aust. Zool. 18(1), 1973

REPRODUCTION IN MACROPUS PARRY!

These data support the statement by Clark and Poole (1967) that M. parryi
does not have a post-partum oestrus. The presence of a large functional follicle
in female 19 and the blastocyst in female 27, suggest that mating may occur
while females are carrying a large pouch young. Following mating, the embryo
derived from it would become quiescent and would not begin to develop until
towards the end of pouch life of the primary young. During the early period of
pouch life there is direct inhibition of ovarian activity by the suckling young
as shown by females 30-23 (Table 1). Such a reproductive pattern has been
observed in the eastern grey kangaroo, M. giganteus, by both Kirkpatrick (1965)
and Clark and Poole (1967).

Direct inhibition of ovarian activity by a suckling young appears to be the
ancestral form of ovarian inhibition (Sharman and Berger, 1969). It has been
retained to varying degrees by the parma wallaby (M. parma), the whiptail
wallaby (M. parryi) and the grey kangaroos (M. giganteus and M. fuliginosus).
In all other macropodids that have been studied in detail, this primitive form
of ovarian inhibition has been lost completely. In these species, ovarian inhibition
while carrying a pouch young is maintained by the suckling stimulus acting via
the corpus luteum of lactation derived from a pre- or post-partum oestrus (Sharman,
1970). The reproductive patterns shown by M. parma, M. parryi, M. giganteus
and M. fuliginosus are thus regarded as being representative of different stages

Fig. 1. (left): Uterine glands associated with 257 ym blastocyst of the whivtail wallaby.
Uterine lumen lower left.

Fig, 2. (right): Corpus luteum associated with 257 ym blastocyst in the whiptail wallaby.

Aust. Zool. 18(1), 1973 45

G. M. MAYNES

in the evolution of embryonic diapause and ovarian. inhibition as observed in
other macropodids.

The retention of transitional reproductive patterns in some species of the
genus Macropus cannot be regarded alone as evidence for, or an indication of, a
close affinity between these species.

REFERENCES

CALABY, J. H. (1966). Mammals of the upper Richmond and Clarence Rivers N.S.W.,
C.S.I.R.O. Div. Wildl. Res. Tech. Paper No. 10.

CALABY, J. H. and POOLE, W. E. (1971). Keeping Kangaroos in Captivity. Int. Zoo.
Vbe af i.- 5-12.

CLARK, M. J. and POOLE, W. E. (1967). The reproductive system and embryonic
diapause in the female grey kangaroo, Macropus giganteus, Aust. J. Zool. 15, 441-59.

KIRKPATRICK, T. H. (1965). Studies of Macropodidae in Queensland 3. Reproduction
in the grey kangaroo (Macropus major) in southern Queensland. Qld. J. Agric. Anim.
Seno 22; “S19-28:

MAYNES, G M. (1972). Age estimation in the parma wallaby, Macropus parma Waterhouse.
Aust. J. Zool. 20, 107-118.

MURPHY, C. R. and SMITH, J. R. (1970). Age determination of pouch young and
juvenile Kangaroo Island wallabies. Trans. R. Soc. S. Aust. 94, 15-20.

SHARMAN, G. B. (1955). Studies on marsupial revroduction III Normal and delayed
pregnancy in Setonix brachyurus. Aust. J. Zool. 3, 56-70.

SHARMAN, G. B. (1970). Reproductive physiology of marsupials. Science 167, 1221-1228.

SHARMAN, G. B. and BERGER, P. J. (1969). Embryonic diapause in marsupials—
Advances in Reproductive Physiology. 4, 211-240.

SHARMAN, G. B., FRITH, H. J. and CALABY, J. H. (1964). Growth of the pouch
young, tooth eruption and age determination in the red kangaroo, Megaleia rufa.
C.S.LR.O. Wildl. Res. 9, 20-49.

VANDEBERG, J. L., COOPER, D. W. and SHARMAN, G. B. (1973). Phosphoglycerate
kinase A polymorphism in the wallaby, Macropus parryi: activity of both X chromo-
somes in muscle. Nature New Biology: in the press.

46 Aust. Zool. 18(1), 1973

Feeding Mortality in a Deep Sea Angler Fish
(Diceratias bispinosus) due to a Macrourid Fish
(Ventrifossa_ sp.)

JOHN R. PAXTON

Department of Ichthyology, The Australian Museum,
Sydney, New South Wales, 2000.

and

ROBERT J. LAVENBERG

Los Angeles County Museum of Natural History,
Los Angeles, California, U.S.A.

A number of deep sea fishes are known to prey on other fishes which exceed
them in size. Large mouths and highly distensible stomachs characterize such fishes
as the swallowers (Chiasmodontidae), gulpers (Saccopharyngidae), and anglers
(Ceratioidea ); all have been recorded with stomach contents containing fishes either
considerably longer than the abdominal cavity of the predator or larger than the
whole fish. The morphological specilizations enabling some fishes to consume
large prey have been described by Tchernavin (1953). The maximum size of food
organisms can only be inferred from the relatively few known instances where the
predator has succumbed due to the size of the prey attempted. Unequivocal
evidence of such circumstances is usually available only when both animals are
found floating at the surface or washed upon a beach.

Recently a 112 mm diceratiid angler fish, Diceratias bispinosus, was found
floating on the surface in the Bismarck Sea off New Ireland with a 369 + mm
macrourid of the genus Vevtrifossa protruding from its mouth (Fig. 1). The
finding has prompted us to make a survey of the known occurrences of this
phenomenon.

Our specimens were taken floating at the surface south-east of the island
of New Ireland, Papua New Guinea (3° 48’ S, 153° 32’ E) at 1600 hours
on 30 April 1968. Both specimens were dead, although little sign of decay is
present and both are in relatively good condition. The macrourid has been
engulfed to a point just behind the tips of the pectoral fins. The head of the
macrourid has distended the stomach and body wall of the angler posteriorly
and ventrolaterally to such a degree that the caudal fin was no longer the most
posterior portion of the angler’s body. With such a mass filling the entire mouth
and body cavity of the angler, both fishes probably died because they could
not pass enough water over the gills to obtain sufficient oxygen.

Aust. Zool. 18(1), 1973 47

48

PAXTON

and LAVENBERG

Aust.

Zool. 18(1), 1973

th partially engulfed Ventrifossa sp.

ispinosus Wi

Diceratias b

Fig. 1.

FEEDING MORTALITY IN A DEEP SEA ANGLER FISH

The long, depressible teeth of the angler effectively prevented the macrourid
from escaping; some dissection was necessary to remove the latter. With the
exception of the area around the tooth marks on the body and in the midventral
area between the anal and ventral fins, no visible digestion of the macrourid has
occurred, as even the eyes are in good condition. One of us (R.J.L.) has previously
noted that fishes from the stomach of gigantactid angler fishes had the greatest
amount of digestion around the tooth marks. Although the cause could be the
ease with which digestive enzymes of the stomach entered the wounds, possibly
a digestive enzyme is associated with the teeth themselves. The presence of
digested areas associated with the tooth wounds on our macrourid specimen
supports this hypothesis for ceratioid angler fishes.

The difference in size between predator and prey is exaggerated when standard
lengths are considered, due to the elongate and narrow tail of the macrourid. A more
meaningful relationship for these two species is biomass; wet weight of the angler
is 175 grams, that of the macrourid 258 grams.

The following list is a literature compilation of deep sea fishes found floating
at the surface with a large food organism in the stomach and/or mouth;
Saccopharynx ampullaceus body 215 mm long with 230 mm Halargyreus johnsoni
in stomach (Gunther, 1887: 256, Fig. 66); S. ampullaceus body 510 mm long
with unidentified fish 180 mm in circumference presumably in mouth (Gunther,
1887: 256); S. flagellum body 355 mm long with 255 mm unidentified fish in
stomach (Gunther, 1887: 256); 50 mm Omosudis lowei with Argyropelecus sp.
25 mm long and 25 mm high in stomach (Gunther 1887: Fig. 52C’; Marshall,
1954: 142); 95 mm Chiasmodon niger with 145 mm Gonostoma denudata in
stomach (Johnson, 1863: 410); 165 mm C. niger with 265 mm Neoscopelus
macrolepidotus? in stomach (Carte, 1866: 35, Fig. 2); C. niger with unidentified
fish in stomach (Goode and Bean, 1896: 292, Fig. 264); 49 mm Linophryne

Fig. 2. Esca of Diceratias bispinosus; scale equals 2 mm.

Aust. Zool. 18(1), 1973 49

PAXTON and LAVENBERG

lucifer with 70 mm scopeloid fish in stomach (Collett, 1886: 138, Fig. 15);
3 Melanocetus johnsoni 75 to 95 mm, each with a 180 to 205 mm Lampanyctus
crocodilus in stomach (Gunther, 1864: 301, Fig. 25; Regan, 1913: 1097).

A number of deep sea fishes taken in trawls have had extremely large
prey organisms in the stomach (for example see Marshall, 1954: 142), yet
presumably have not been adversely affected. The 12 known surface captures
indicating feeding mortality give little information as to the relative frequency
of the phenomenon. However, it is surprising that all of the previous reports were
made prior to 1900, considering the larger number of oceanographic vessels at
sea today. In the present case the macrourid is very close to the maximum circum-
ference possible for the angler to ingest with extreme jaw expansion. Since the
dental morphology of the angler does not allow a large food organism to be
rejected once swallowed, the trial and error method of learning maximum food
size must be severely limited.

The present angler (Austr. Mus. Sydney, I.15602-001) is the fourth recorded
metamorphosed female of Diceratius bispinosus. Others have been taken from the
Banda Sea (Giinther, 1887), the Indian Ocean off Malabar (Alcock, 1899)
and the western Atlantic off South America (Grey, 1959); all three were taken
with bottom fishing gear (Grey, 1959). Most macrourids, including species of
Ventrifossa, are benthic, and apparently D. bispinosus, while having no obvious
morphologic adaptations for benthic life, lives and feeds close to the bottom.
Bertelsen (1951) recorded two larval females from the upper waters of the
Indian Ocean and Celebes Sea.

The present specimen agrees in almost all respects with that described by
Grey (1959), although there are no illicial papillae on our specimen. The outer
primary appendages of the esca (Fig. 2) are more highly developed than in the
specimen figured by Regan and Trewavas (1932: Fig. 85A). A moderately large
pore is on the right or dorsal side of the middle appendage of the esca. We
cannot ascertain whether the esca is normally held in a lateral or dorsoventral
position over the mouth, but the pore is definitely not on the left side, as in
Giinther’s (1887) specimen. Grey (1959: 226) described a subcutaneous structure
over the dentary which she considered a gland or possibly a tentacle. Pietsch
(1972: 31) called the structure a labial cartilage. In the present specimen the
structure is not cartilagenous, but filled with an opaque substance of jelly-like
consistency after formalin preservation. The skin over the organ is thickened
and darkly pigmented, somewhat as the ‘cavernous luminous tissue’ found in
some whale fishes of the family Cetomimidae (Harry, 1952). However, the
functional significance of the structure is unclear.

The specimen of Ventrifossa (AMS 1.15602-002) does not match the des-
cription of any species described by Gilbert and Hubbs (1920) or Parr (1946).
The fish is partially digested between the anal and ventral fins and the obscurity
of the often diagnostic luminous organs and associated lenses and scaleless areas
precludes formal naming and description.

50 Aust. Zool. 18(1), 1973

FEEDING MORTALITY IN A DEEP SEA ANGLER FISH

We are indebted to the Papua New Guinea Department of Agriculture, Stock
and Fisheries and particularly to W. A. Filewood of the Fisheries Research and Survey
Station, Port Moresby, for making the specimens avilable. Father W. Hager of the
Catholic Mission, Tonga collected the specimens, forwarded them to fisheries authorities,
and made available additional collecting details. C. V. Turner took the photograph.
B. B. Collette, T. W. Pietsch and F. H. Talbot kindly reviewed the manuscript. To all
go our appreciation.

REFERENCES

ALCOCK, A. (1899). A descriptive catalogue of the Indian deep-sea fishes in the Indian
Museum. Calcutta. 211 pp.

BERTELSEN, E. (1951). The ceratioid fishes. Ontogeny, taxonomy, distribution and biology.
Dana Rept., 39, 1-276.

CARTE, A. (1866). Notes on the genus Chiasmodon. Proc. Zool. Soc. London, 1866,
35-39.

COLLETT, R. (1886). On a new pediculate fish from the sea off Madeira. Proc. Zool.
Soc. London, 1886, 138-143.

GILBERT, C. H. and HUBBS, C. L. (1920). The macrouroid fishes of the Philippine
Islands and the East Indies. U.S. Nat. Mus. Bull., 100, J (7), 369-588.

GOODE, G. B. and BEAN, T. H. (1896). Oceania ichthyloogy. Smithsonian Inst. Spec.
Bull., 2; 1-553.

GREY, M. (1959). Descriptions of newly discovered western Atlantic specimens of
Diceratias bispinosus Giinther and Paroneiroides wedli (Peitschmann). Copeia, 1959
(3), 225-228.

GUNTHER, A. (1864). On a new genus of pediculate fish from the sea of Madeira.
Proc. Zool. Soc. London, 1864, 301-303.

. (1887). Report on the deep-sea fishes collected by H.M.S. Challenger during
the years 1873-1876. Challenger Rept. Zool., 22, 1-335.

HARRY, R. R. (1952). Deep sea fishes of the Bermuda oceanographic expeditions.
Families Cetomimidae and Rondeletiidae. Zoologica (N.Y.), 37 (1), 55-72.

JOHNSON, J. Y. (1863). Descriptions of three new genera of marine fishes obtained at
Madeira. Proc. Zool. Soc. London, 1863, 403-410.

MARSHALL, N. B. (1954). Aspects of deep-sea biology. Hutchinson, London.

PARR, A. E. (1946). The Macrouridae of the Western North Atlantic and Central
American seas. Bull. Bingham Oceanogr. Coll., 10 (1), 1-99.

PIETSCH, T. W. (1972). A review of the monotypic deep-sea anglerfish family Centro-
phrynidae: taxonomy, distribution and osteology. Copeia, 1972 (1), 17-47.

REGAN, C. T. (1913). Exhibition of a specimen of the deep-sea angler-fish Melanocetus
johnsonii Giinth. Proc. Zool. Soc. London, 1913, 1096-1097.

and TREWAVAS, E. (1932). Deep-sea angler fishes (Ceratioidea). Dana Rept.,
Ze tes

TCHERNAVIN, V. V. (1953). The feeding mechanisms of a deep sea fish Chauliodus

sloanei Schneider. British Mus. (Nat. Hist.), London.

Aust. Zool. 18(1), 1973 51

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ROYAL ZOOLOGICAL SOCIETY OF NEW SOUTH WALES
Established 1879

Registered under the Companies Act, 1961

Patron:

His Excellency the Governor of New South Wales, Sir Arthur Roden Cutler,
Wi@e Kee MG. K.C:V.O., CBE. KtStJ.,. B-Ec, ELD. - DSc.

COUNCIL, 1972

President:
Ronald Strahan, M.Sc., F.L.S., F.R.Z.S.
M.I. Biol., F.S.I.H.

Vice-Presidents:

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THE AUSTRALIAN ZOOLOGIS

VOR, 1S 7 AUGUST, 1973

Contents

Observations on the behaviour of the macropodid marsupial Ti
billardieri (Desmarest) in captivity. S. R. MORTON and ©
BURTON ceca hase ee ce acta c/o ec

Breeding biology of some species of Preidaphane (Anura: Leptod: “ty
of the Southern Highlands, New South Wales. R. PENGILLE

Three new species of deep-dwelling damselfishes (Pomacentridae)

SOHEU-west «Pacmicn@cean=s rm tre VL IUGINS Sea -eeetie ere
Short papers—

IMLAY NES fr oct en- pce aacineta eee ici ae ea

odin mortality in a deep sea angler fish rec ae bispi

macrourid fish (Ventrifossa sp.). J. R. a & RI La |

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he Royal Zoological Society of |

“NOTICE TO AUTHORS

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THE AUSTRALIAN ZOOLOGIST

VOL. 18 JUNE, 1974 PART 2

Physiological Adaptations of Anuran Amphibians to
Aridity: Australian Prospects

R. S. SEYMOUR and A. K. LEE
Department of Zoology, Monash University, Clayton, Victoria, 3168

ABSTRACT

We attempt to assess research on Australian anurans in the light of recent
discoveries involving species outside Australia and consequently question the
relevance of several studies of frogs in laboratory conditions.

Amphibians generally avoid water loss by burrowing or otherwise selecting
favourable microclimates. Although it has been thought that the evaporative
water loss from anurans is high and uncontrolled, several species from arid
environments greatly reduce water loss under certain conditions. Some arboreal
frogs apparently secrete a lipid-like substance onto the skin. Other fossorial
species produce ‘‘cocoons”’ of dead epidermis.

Uricotelism, once thought excluded from the amphibia as a major form of
nitrogen excretion, has been demonstrated in several arboreal species, one of
which may excrete over 80% of the waste nitrogen as urates. Several Australian
Litoria should be examined because they exploit niches similar to those of the
uricotelic frogs from Argentina and Rhodesia.

Virtually nothing is known about the thermal relations or energy economy of
Australian anurans. Desert species, studied elsewhere, feed intensely throughout
a limited season of activity and show rapid gains in fat reserves correlated
with a high calorigenic effect. Basking in some species may be significant in
taking maximum advantage of a transient food supply typical of arid environments.
Long periods of dormancy characterized by temperature independent reduction
of metabolic rate occur in some desert toads.

INTRODUCTION

Bentley (1966a), Mayhew (1968), Schmid (1969), and Shoemaker (1974)
have thoroughly reviewed the physiology of desert amphibians, and Thorson
(1955), Chew (1961), Heatwole et al. (1969), and Bentley (1971) have
considered specific aspects of their water economy. The intent of this report
is not to offer another review of the literature because that cannot be justified
at this time. Our purpose is to assess research on Australian desert anurans
in the light of a number of interesting discoveries, published primarily since
1969, which have not yet appeared in review. Most of this new information

Aust. Zool. 18(2), 1974 53

RA. S. SEYMOUR: zandiiAy Ke BEE

comes from outside Australia but some of it may ultimately apply to Australian
species. It is tempting to suggest Australian candidates which may be profitably
examined. Our task has been eased by an excellent review of the ecology of
Australian frogs by Main (1968). This report is restricted to the physiological
adaptations of adult anurans. Very little has been added to our knowledge of
the ecology and life histories of the larvae of desert anurans.

WATER ECONOMY

Evaporative Water Loss—Because amphibians generally show little physio-
logical capacity to restrict evaporation from the skin and none is known to
produce hypertonic urine, many investigators have been drawn to study amphibians
in deserts where free water is seldom available and the potential evaporation
is high. The results show that few species are physiologically adapted to arid
conditions to the standards set by higher vertebrates. Most adaptations are
behavioural.

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Fig. 1. Evaporative water loss from seven species of Australian frogs ranked according
to decreasing aridity of habitat: Cyclorana platycephalus (1), Cyclorana alboguttatus
(2), Neobatrachus pictus (3), Litoria caerulea (4), Limnodynastes dumerili (5), Litoria
aurea (6), Mixophyes fasciolatus (7). Data from Clyne (1969).

54 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

Among the majority of amphibians, there have been no large or consistent
differences in evaporative water loss (EWL) between terrestrial species from
mesic and xeric habitats (see Bentley, 1971) (Fig. 1). The lowest rate of
EWL from a North American species is about 50 times the rate in lizards
(Claussen, 1969). It is important to note that most measurements have been
obtained by placing the animals in conditions favouring a rapid water loss
(e.g. over drying agents, in streams of dry air, in wire cages indoors, etc.).
As experiments designed to compare maximal rates of water loss, they have
been successful, but there have been only a few studies which assess evaporation
rates under natural conditions (Packer, 1963; Lee, 1968). If by behavioural
means most frogs naturally avoid these extreme drying conditions, there should
be little selective pressure for physiological mechanisms limiting water loss. On
the other hand, there is apparently great selective advantage in maintaining
the skin as an organ of gas exchange and water uptake. Both of these functions
require a skin permeable to water.

The majority of desert species burrow into the soil where humidities are
high and where water is available to be absorbed through the skin. Examples include
the species of Notaden, Cyclorana, and Neobatrachus from Australian deserts. A few
desert species remain close to permanent water such as Litoria rubella and
L. caerulea, although L. rubella may exist away from permanent water and seek
refuge in moist crevices.

Until recently it has been assumed amphibian skin had no influence on
the rate of evaporation. However, various dormant anurans produce cocoons
of shed skin which restrict water loss. The cocoons may be as many
as 50 cell layers thick, as in the Argentinian toads, Ceratophrys ornatus and
Lepidobatrachus llanensis (McClanahan ef al., 1974), or only one cell layer
as in the Australian burrowing frogs, Cyclorana platycephalus, C. alboguttatus
and possibly Limnodynastes spenceri (Lee and Mercer, 1967). The isolated
cocoon of Cyclorana alboguttatus limits water transfer from a saturated atmos-
phere into a dry atmosphere to 0.65 mg HzO cm-h* at 25°C. The rate of
water loss from the intact animal is some 72 times higher. Shoemaker, Ruibal, and
McClanahan have observed cocoon formation in two species of burrowing ceratophrid
frogs from Argentina. Cocoons form during exposure to room air and are also
produced by buried frogs as the soil dries. The EWL from frogs with cocoons
is about one-tenth of that from frogs without cocoons. Lepidobatrachus llanensis
exhibits a progressive decrease in EWL when exposed to room air (Fig. 2).
After 5 months underground, EWL in dry air was extremely low.

Most other measurements of EWL in burrowing frogs have been made
on animals taken immediately from moist conditions and dehydrated at an
extremely rapid rate. It is not surprising that the EWL was high and death
occurred quickly even in frogs known to produce cocoons. However, under
conditions of long-term, low-level dehydration, such as beneath gradually drying
soil, amphibians may be given enough time to physiologically adjust the rate of

Aust. Zool. 18(2), 1974 55

Re So. SEYMOUR? rand A; (Ki cGEE

10
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1 3 5 7 9 18 25 26 150

Days

Fig. 2. Evaporative water loss of Lepidobatrachus llanensis related to number of days
exposed to room air. Cocoons were visible after three days. X symbolize values
from two frogs after about five months under dried soil. Data from McClanahan
et al. (1974).

water loss by producing a cocoon. Short-term experiments by Warburg (1965, 1967)
suggest that EWL decreases with time in Neobatrachus pictus, N. centralis,
Limnodynastes dumerili, L. ornatus and Cyclorana spp. All of these frogs probably
produce cocoons. However, Packer (1963), found no change in the rate of EWL
from Heleioporus eyrei over a period of six days in dry sand. It is not known
if Heleioporus spp. produce cocoons during dormancy. None of the experiments
on EWL of Australian anurans was directed toward an assessment of the
effect of changes in the skin so none is conclusive. We feel that more work
is needed to determine the role of the cocoon on water loss as well as gas
exchange and water uptake. The cocoons of the Australian frogs which have been
examined are continuous over the eyes and the cloaca, but the nostrils remain
open. Seymour (1973c) has shown that all gas exchange occurs cutaneously in
dormant spadefoot toads (Scaphiopus) without cocoons, but it is possible that
the buildup of shed skin limits respiratory gas exchange to a point necessitating
pulmonary ventilation. The cocoons may also function to limit contact of the
frogs with the soil thus reducing water exchange. In very moist soil, a cocoon
could limit an influx of water which may be harmful to arid-land anurans
(Schmid, 1965). On the other hand, if the soil dries sufficiently to pull moisture
out of a frog by virtue of a high soil moisture tension, a cocoon may effectively
break capillary contact between the soil and the frog’s skin.

56 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

Recently, some arboreal frogs from semi-arid habitats in Argentina (Phyllo-
medusa spp.) and Rhodesia (Chiromantis xerampelina) have attracted attention
because their rates of EWL are characteristic of reptiles (Loveridge, 1970; Shoe-
maker et al., 1972). These moderately sized tree frogs may spend many months
in exposed situations which would quickly kill other anurans. The nature
of the cutaneous barrier to water loss is currently being analyzed by V. H.
Shoemaker, R. Ruibal, and L. McClanahan and their associates. They found
that water beaded when placed on the skin of the Argentinian frog, Phyllomedusa
sauvagii, and detergent washes destroyed the barrier. This implied that a
lipid layer covers the skin, possibly spread by stereotyped wiping movements
whereby each leg wipes the dorsal and ventral surfaces of the body after the
frog emerges from water. Histological sections revealed that the skin contains
glands which react intensely with lipid stains, and thin-layer chromatography of
skin extract showed lipid constituents not present in a closely related frog having
high rates of water loss.

It is even more surprising that both frogs (Chiromantis xerampelina from
Rhodesia and P. sauvagii from Argentina) excrete primarily urates, the salts
of uric acid. Urates were formerly thought to be excreted by amphibians only
in trace amounts (Deyrup, 1964). However, in P. sauvagii, urates account for
over 80% of the nitrogen excreted, even when free water is available (Shoemaker,
pers. comm.). Under conditions of restricted water, urates are stored in the
bladder, and are subsequently voided within a mucous envelope when water is
available. Thus P. sauvagii maintains plasma electrolyte and urea concentrations
at relatively low levels even when the only source of water is in insect food.
Under the same conditions, Hyla pulchella shows a pronounced rise in plasma
urea and electrolytes.

It is interesting to note that uricotelism and low evaporative water loss
occur in a rhacophorid from Africa and in several phyllomedusids from South
America. This may represent convergent evolution in an arboreal and semi-arid
niche. Several Australian Litoria spp. occupy this niche, and should be examined
for these adaptations. L. rubella may be found close to dry gilghis in the semi-
arid areas of western New South Wales where they may spend months without
free water. L. peronii commonly calls from trees some distance from free water,
and L. caerulea resides in crevices though near to water.

Measurements of EWL have been made on some Australian Litoria spp.
by Warburg (1972) and Clyne (1969). The values Warburg obtained for xeric
anurans from Australia and Israel are extremely low and these frogs would seem
superficially better adapted than frogs elsewhere. However, values obtained by
Clyne on some of the same species under identical conditions are roughly 20
times higher and compare favourably with values from other literature. Clyne
showed no consistent relationship between water loss and habitat (Fig. 1).
Furthermore, EWL in L. caerulea was not significantly different (on a surface
specific basis) than that in L. aurea, and the large rain forest frog, Mixophyes

Aust. Zool. 18(2), 1974 57

RLS.) SEYMOUR’ sand:sA: K. CEBE

fasciolatus had significantly the lowest surface specific EWL. There are, however,
no careful measurements of EWL of Australian semi-arid and arid land frogs
under conditions favouring gradual water loss for periods longer than six days.

Cutaneous Uptake of Free Water—Many studies have attempted to define
a correlation between the rate of cutaneous water uptake and independence
of water in amphibians. (Thorson, 1955, McClanahan, 1967, Claussen, 1969,
Fair, 1970). In general, no reasonably consistent relationship has appeared. Bentley
et al. (1958) found a significant correlation in four species of Neobatrachus,
but not in five species of Heleioporus from habitats of varying aridity. Warburg
(1965, 1972) claimed to have demonstrated a relationship in several Australian
anurans, but his results are equivocal, and the degree and rate of dehydration
were not adequately controlled.

All of the published studies compare water uptake based on body weight
or total skin area and thus ignore possible regional differences in skin permeability.
McClanahan and Baldwin (1969) have shown that the water uptake through
the ventral pelvic “seat patch” accounts for about 70% of the total uptake
in dehydrated Bufo punctatus. This regional difference in skin permeability may
not be universal in desert anurans, but it may be considered adaptive in allowing
the animal to take advantage of transient wet surfaces resulting from dew or
rain. Bufo punctatus cannot be distinguished from other North American anurans
when the rate of water uptake is expressed in terms of total surface area
(Claussen, 1969). However, with access to wet surfaces, this toad may rehydrate
more rapidly than other species not possessing a “‘seat patch’’. Stille (1958) and
Johnson (1969) have described various postural adjustments in dehydrated
American and Australian frogs which appear to enhance contact between the
frog and wet surfaces.

It has been generally assumed that a high rate of cutaneous water uptake,
per se, has adaptive value for an anuran in arid conditions. Main (1968) summarized
the rates in 19 species of Western Australian frogs and attributed the significant
differences to selection favouring individuals which dehydrate quickly. Thus these
frogs could begin calling immediately after rain following a severe drought.
Bentley (1971, p. 172) rationalized the failure to find universally higher rates
in desert species by suggesting that the differences between species “‘arose and
peristed in response to ecological stresses on the Amphibia during their evolutionary
history, even though they may not today be precisely related to the life of all
contemporary species”. For several reasons, these conclusions must be viewed
with caution. First, experiments where frogs are immersed in water may not
be relevant because rapid rates of dehydration in the laboratory and regional
differences in water uptake may obscure the natural rate of uptake in the field.
Second, if dehydration occurs gradually over a long period during dormancy
as in American spadefoot toads (Shoemaker e¢ al., 1969), it seems energetically
disadvantageous for frogs to produce short-lived neurohypophysial hormones
continuously in preparation for rehydration. Moreover, the hormones could con-

58 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

ceivably enhance the rate of water loss from the frogs by increasing the permeability
of the skin in the face of a drying soil. It may be significant in this connection
that Jasinski and Gorbman (1967) found only minimal differences in stainable
hypothalamic neurosecretion between hydrated and naturally dehydrated spadefoot
toads in the field. It should be interesting to determine the titres of hormones
and rates of water uptake in dehydrated frogs emerging in response to summer
rain. Finally, the difference in average time for replacement of a 25% water
deficit between the desert toad, Neobatrachus wilsmorei, and its most mesic
relative, Neobatrachus pelobatoides, is only one hour (Bentley e¢ al., 1958). Even
if complete hydration is a prerequisite to chorusing in these frogs, it seems
unlikely that this small difference has much effect on the frogs’ reproductive
success.

Uptake of Water from Soil.—In even the driest deserts, moist soil may occur
close to the surface. Amphibians may be the only tetrapods which can directly
absorb moisture held by soil. The rate at which water moves between an
amphibian and the soil depends basically on (1) the osmotic pressure of the
animal’s body fluids, (2) the permeability of the skin, (3) the osmotic pressure
of the soil water and (4) the affinity of the soil particles for water (soil matric
potential). The effect of the soil matric potential on the water economy of
American spadefoots has been examined by Ruibal ef al. (1969), Shoemaker e¢ al.
(1969) and McClanahan (1972). Soils of fine particle size have a higher affinity
for water than do sandy soils with identical water content. Therefore it is more
difficult for toads to obtain water from silty soil than sandy soil when the water
content is low. When soil dries to a point where the soil matric potential is
greater than the equivalent osmotic pressure of the body fluids, the frogs deaminate
protein, producing urea which is retained along with electrolytes. The effect is to
balance, to a certain extent, the soil matric potential with elevated total osmotic
pressure. This response is similar to that observed in Rana cancrivora and Bufo
viridis which swim in saline water (Gordon et al., 1961; Gordon, 1962).

Aside from one experiment by Packer (1963), there have been no investi-
gations of water uptake from soil in Australian species. It should be interesting
to compare the responses of Neobatrachus spp. which burrow in silty soil
with those Helezoporus spp. which prefer sand (Main, 1968).

Water Storage and Tolerance to Dehydration—Amphibians generally have a
greater tolerance to water loss than most vertebrates and a number of studies
have shown still greater tolerance in desert species (Thorson and Svihla, 1943;
Thorson, 1955; Main and Bentley, 1964; Schmid, 1965). Desert anurans also
have large urinary bladders (Bentley, 1966b) which may serve to store water
to replace evaporative losses (Ruibal, 1962; Shoemaker, 1964; Clyne, 1969).
Thus the production of hyposmotic urine and the active transport of ions through
the bladder wall result in a significant store of very dilute urine. Spencer (1896,
p. 163-164) popularized Cyclorana platycephalus as the “water holding frog”,
familiar to early settlers (and the aborigines before them) along the Diamantina

Aust. Zool. 18(2), 1974 59

R. S. SEYMOUR and A. K. LEE

River for their capacity to hold ‘‘a wine glass full of clear sweet water”. This
species together with Notaden nicholsi and Neobatrachus wilsmorei have capacities
which approximate 50% of the rest of the body weight (Main and Bentley, 1964).
However, Clyne (1969) obtained similar values for Mixophyes fasciolatus, a frog
from wet sclerophyll and rain forest in eastern Australia. Thus large bladders are
not always correlated with arid conditions, and their function in Na’ retention
may complicate this relationship.

Because water losses are replaced by bladder reserves, we expect that large
bladders should be favoured in arboreal frogs which bask away from water. However,
not enough reliable data are available to judge this issue. Litoria moorei has a
bladder capacity between 20 to 30% (Main and Bentley, 1964), or similar to
North American desert toads (Ruibal, 1962; McClanahan, 1967), and this frog
may bask like the related species, L. aurea.

The value of bladder water in an Australian frog under natural conditions
has been demonstrated only once (Lee, 1968). Heleioporus eyrei may evaporate
over 20% of the body weight during one night of foraging. If food is available,
the water loss is regained to an extent by eating. However, it is assumed that
losses from the body are also met by reabsorption from the bladder at night,
and ultimately by reabsorption from the soil during the day.

The “water balance response” whereby dehydration stimulates release of
neurohypophysial hormones, which, among other things, increase the rate of
water uptake from the environment and from the bladder, has been well described
elsewhere (see Bentley, 1971). We therefore wish only to point out a long over-
looked feature of some amphibian kidneys which may be functionally related to
reabsorption of bladder water. Many investigators during the 19th and early
20th centuries described the nephrostomes opening from the kidney into the
coelomic cavity of adult anurans (see Smith, 1961). Sweet (1907) determined that
they also open into the renal veins and associated blood vessels, since carmine in-
jected into the body cavity appeared later in the peritubular blood sinuses, renal
veins, and post cava, but not in the tubules. Sweet suggested that water reabsorbed
from the bladder into the coelom entered the vascular system through the nephro-
stomes and was distributed from there to peripheral regions. There appears to be a
correlation between the number of nephrostomes and arid habitats in Australian
anurans (Table 1). Those species with large bladders have the largest number
of nephrostomal openings, and among the Litoria, it may be significant that
L. aurea basks whereas L. lesueurii apparently does not. We surmise that the
rate of reabsorption of water from the bladder in a basking frog would be higher
than in a non-basking frog under natural conditions but this remains to be
demonstrated.

THERMAL RELATIONS

When amphibians seek shelter, they avoid not only drying conditions but also
stressful temperatures. Accordingly, their absolute temperature tolerance is low

60 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

TABLE 1

NUMBER OF NEPHROSTOMAL OPENINGS PER KIDNEY IN AUSTRALIAN
ANURANS FROM VARIOUS HABITATS (SWEET, 1907).

Species Number Habitat

Crinia signifera “small number” Mesic—ponds and marshes

Litoria lesuerii 300 Mesic—rivers and streams
Neobatrachus pictus 105 Mesic, Xeric—ephemeral ponds

Litoria aurea 150-200 Mesic—ponds, billabongs, rivers (basks)
Cyclorana alboguttatus 210 Xeric—ephemeral ponds

Notaden bennetti 1067 Xeric—ephemeral ponds

compared to other vertebrates (see Bentley, 1966a; Brattstrom, 1970). Most
differences in tolerance among anurans from different habitats can be ascribed

to acclimation and acclimatization within a genetically fixed range (Brattstrom,
197.0):

Body temperatures of desert anurans in the field are generally below 30°C
(see Mayhew, 1968, for references), but a few species have higher temperatures.
The highest body temperatures, 39.2°C, come from Australian Cyclorana spp.
captured in shallow pools (Main, 1968); this temperature is close to maximum
burrow temperatures experienced by spadefoot toads in North America (Ruibal
Cera. L969),

High body temperatures also occur in species which bask. Juvenile Bufo
californicus and Bufo debilis are active diurnally and reach maximum temperatures
of 37°C and 34°C respectively (Cunningham, 1962; Seymour, 1972). We have
no field body temperatures of basking Australian frogs.

ENERGY METABOLISM

In addition to the problems of heat and water, deserts are characterized by
an inconsistent, if not low, supply of food. Further, desert frogs feed only when
surface conditions permit and consequently do not feed for most of the year.
Hence desert frogs must build up substantial energy reserves within a relatively
brief period to survive long periods of dormancy.

North American spadefoot toads (Scaphiopus spp.) show several behavioural
and physiological adaptations which optimize the utilization of transient food
sources during the summer rainy season. Like most anurans, they are opportunistic
predators which acept a large range of prey size (R. S. Seymour, unpubl. data;
see also Heatwole and Heatwole, 1968). They feed during the night, especially
following rains which also promote the emergence of insects, and enter shallow
burrows during the day where humidity and temperature are high (Ruibal e¢ al.,
1969). The high burrow temperatures (27-39°C) doubtless facilitate digestion.
A high calorigenic effect following feeding in S. hammondii suggests that proteins

Aust. Zool. 18(2), 1974 61

A... Ss SEYMOUR? and, A: -K: LEE

are rapidly converted into non-protein products, probably chiefly fat (Seymour,
1973b). The rate of fat deposition appears to be higher in S. couch and S.
hammondii than in mesic anurans, and the relative weight of fat stored is
higher (Seymour, 1973b). These differences are consonant with longer periods of
dormancy at higher ambient temperatures in desert toads.

Rates of oxygen consumption and heart beat decrease during dormancy in
spadefoot toads (Seymour, 1973b, c). Oxygen consumption during dormancy is
about 20% of values from toads resting on the surface in the same season
(Fig. 3). At this low metabolic rate, fat reserves are sufficient for the toads
to remain in dormancy for at least 10 months. When additional resources of energy
in body protein and eggs are considered, it is evident that some individuals in a
population could survive for two or more years without feeding, even with an
average body temperature of 15°C.

The aerobic metabolic scope for activity and the tolerance of exercise are
high in S. hammondii and Bufo cognatus compared to Rana pipiens and R.
catesbeiana (Seymour, 1973a). This may be related to the regular occurrence of

Ol
O

we)
O

rook
O

on

O, Consumption ul g™hr7

Dormant

Gis 10 5 BO oS 30
Body Temperature C

Fig. 3. Regression lines of oxygen consumption of Scaphiopus couchii during dormancy
beneath soil and after arousal from dormancy during the same season. Measurements
from aroused toads were taken under conditions required to produce the accepted
concept of standard metabolic rate (SMR). Data from Seymour (1973b).

62 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

burrowing in the two desert toads. Burrowing demands a sustained high rate
of energy supply whereas the short bursts of activity exhibited by the two ranids
may occur largely anaerobically. These observations have been confirmed in
certain Australian frogs by D. Bromell (pers. comm.). The leptodactylids,
Limnodynastes dumerili and L. tasmaniensis, have higher aerobic scopes than the
hylids, Litoria aurea and L.-verreauxi. Low aerobic scope appears to be complemented
by high anaerobic scope in anurans (Bennett and Licht, 1973).

NEW DIRECTIONS

The mainstream of research into the adaptations of amphibians to arid
conditions has been directed largely at aspects of water economy and particularly
at assessment of tolerances and capacities under laboratory conditions. Recently
emphasis has shifted to studies of water economy in natural environments, and
to energetics. Several attributes new to our knowledge of amphibia have emerged—
uricotelism, cocoons, lipid barriers to water loss, etc. This report suggests Australian
species which may show adaptations similar to these. However, we feel that
it would be most rewarding to search for new mechanisms which are not variations
on a theme. It must be remembered that studies on the performance of desert
amphibia in natural environments have been based largely upon species from
North American deserts characterized by a relatively predictable climate. Therefore
the rospect of discovering new strategies in Australia seems good since
Australian deserts are characterized by frequent yet unpredicable drought.

ACKNOWLEDGEMENTS

We thank V. H. Shoemaker for supplying unpublished material which is represented
in a large portion of this report. The junior author gratefully acknowledges the receipt
of an A.R.G.C. grant (65-15965) funding formerly unpublished research by Clyne (1969).

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BRATTSTROM, B. H. (1970). Amphibia. In: Comparative Physiology of Thermoregulation.
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Aust. Zool. 18(2), 1974 63

R. S. SEYMOUR and A. K. LEE

CHEW. R. M. (1961). Water metabolism of desert-inhabiting vertebrates. Biol. Rev.
36, 1-31.

CLAUSSEN, D. L. (1969). Studies of water-loss and rehydration in anurans. Physiol. Zool.
42, 1-14.

CLYNE, F. (1969). Water and ionic relations of selected Australian frogs. M.Sc. thesis.
Monash University.

CUNNINGHAM, J. D. (1962). Observations on the natural eee of the California toad,
Bufo californicus Camp. Herpetologica 17, 255-260.

DEYRUP, I. J. (1964). Water balance are kidney. In: Physiology of the Amphibia. J. A.
Moore (ed.). Academic Press, London and New York. pp. 251-328.

FAIR, J. W. (1970). Comparative rates of rehydration from soil in two species of toads,
Bufo boreas and Bufo punctatus. Comp. Biochem. Physiol. 34, 281-287.

GORDON, M. S. (1962). Osmotic regulation in the green toad (Bufo viridis). J. Exp.
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GORDON, M. S., K. SCHMIDT-NIELSEN & H. M. KELLY (1961). Osmotic eerie in

the crab- -eating frog (Rana cancrivora). J. Exp. Biol. 38, 659-678.

HEATWOLE, H. & A. HEATWOLE (1968). Motivational aspects of feeding behaviour
in toads. Copeia 1972, 692-698.

HEATWOLE, H., F. TORRES, S. BLASNI DE AUSTIN & A. HEATWOLE (1969).

Studies on anuran water balance. I. Dynamics of evaporative water loss by the
coqui, Eleutherodactylus portoricensis. Comp. Biochem. Physiol. 28, 245-269.

JASINSKI, A. & A. GORBMAN (1967). Hypothalamic neurosecretion in the spadefoot toad,
Scaphiopus hammondi, under different environmental conditions. Copeia 1967, 271-279.

JOHNSON, C. R. (1969). Water absorption response of some Australian anurans. Herpeto-
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LEE, A. K. (1968). Water economy of the burrowing frog, Heleioporus eyrei (Gray).
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LEE, A. K. & E. H. MERCER (1967). Cocoon surrounding desert dwelling frogs. Science
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LOVERIDGE, J. P. (1970). Observations on nitrogen excretion and water relations of
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MAIN, A. R. (1968). Ecology, systematics and evolution of Australian frogs. In: Advances
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MAIN, A. R. & P. J. BENTLEY (1964). Water relations of Australian burrowing frogs
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MAYHEW, W. W. (1968). Biology of desert amphibians and reptiles. Jn: Desert Biology.
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McCLANAHAN, L. Jr (1967). Adaptations of the spadefoot toad, Scaphiopus couchi, to
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McCLANAHAN, L. Jr (1972). Changes in body fluids of burrowed spadefoot toads as a
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McCLANAHAN, L. Jr & R. BALDWIN (1969). Rate of water uptake through the integument
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McCLANAHAN, L. Jr, V. H. SHOEMAKER & R. RUIBAL (1974). Evaporative water
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64 Aust. Zool. 18(2), 1974

PHYSIOLOGICAL ADAPTATIONS OF ANURANS

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RUIBAL, R., L. TEVIS Jr & V. ROIG (1969). The terrestrial ecology of the spadefoot
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SPENCER, B. (1896). Report on the work of the Horn Scientific Expedition to Central
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WARBURG, M. R. (1965). Studies on the water economy of some Australian frogs,
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Aust. Zool. 18(2), 1974 65

The Status of Pseudomys novaehollandiae
(the New Holland Mouse)

H. POSAMENTIER and H. F. RECHER

Department of Environmental Studies
The Australian Museum, Sydney, New South Wales, 2000.

ABSTRACT

The New Holland Mouse, Pseudomys novaehollandiae (Waterhouse), has been
found to have a wide if somewhat patchy distribution from the north coast of
New South Wales to the Kangaroo Valley south of Sydney. It occurs again on
the Mornington Peninsula in Victoria. In many of the places where it has been
found, it is very abundant and can not be considered endangered. Rather, its
ecology and habitat requirements are suggestive of an opportunistic species
which flourishes during certain successional stages after fire.

INTRODUCTION

After a long period when it was considered that it might be extinct,
Pseudomys novaehollandiae (the New Holland Mouse) was rediscovered in 1967.
Reporting on the rediscovery, Mahoney and Marlow (1968) considered that the
apparent rarity of P. novaehollandiae resulted from its being confused with the
abundant and widespread introduced House Mouse (Mus musculus) and from
the limited amount of trapping which has been done on the east coast of Australia
in past years. However, the exact status of P. novaehollandiae has remained
in doubt and both Calaby (1969) and Ride (1970) considered it an ‘endangered
species’. The animal is listed in the JTUCN handbook on rare and endangered
species as endangered.

In the course of studies on the small mammal fauna of coastal heaths in
New South Wales, we trapped P. novaehollandiae at a number of places in
northern New South Wales and found it to be abundant and widely distributed.
Other workers have also taken P. novaehollandiae since 1967 and it seems
appropriate at this time to review its status. Additionally though our work has
been confined largely to heaths it provides information on the features of the
habitat which influence the distribution and abundance of P. novaehollandiae.
Detailed analyses of the relation between habitat and small mammals in coastal
heaths will be presented elsewhere, but are discussed briefly in this paper
with specific reference to the status of P. novaehollandiae.

DISTRIBUTION

Previous to 1967, P. novaehollandiae was known from only four specimens
taken in northern New South Wales near Scone, upper Hunter River, and from

66 Aust. Zool. 18(2), 1974

STATUS OF PSEUDOMYS NOVAEHOLLANDIAE

the vicinity of the Richmond River. In 1967 a single specimen was collected
in Ku-ring-gai Chase National Park near Sydney (Mahoney and Marlow, 1968)
and in 1968 a number of specimens were taken from a locality near Port Stephens
(Keith and Calaby, 1968). Subsequently, specimens have been collected at Smiths
Lake near Forster (Batt et al., 1972) and from south of Sydney in the Royal
National Park (P. Foster, pers. comm.) and the Kangaroo Valley (Mahoney,
pers. comm.). A second specimen was taken in Ku-ring-gai Chase in October
1973 from a place near the original record. We have collected it ourselves at a
number of localities on the north coast of New South Wales from Port Stephens
to the Richmond River (Evans Head).

Though P. xovaehollandiae is likely to occur on the south caast of New
South Wales, the specimen from the Kangaroo Valley is the only record from
that part of the state. We have not trapped it in an extensive trapping programme
in the Nadgee Nature Reserve south of Eden nor has it been taken at Jervis
Bay despite intensive trapping by Douglas (pers. comm). However, in an interesting
extension of its known modern range, Seebeck and Beste (1970) have taken
P. novaehollandiae on the Mornington Peninsula in south-eastern Victoria and
further trapping could well extend the animal’s known range. Mahoney and

‘FABLE 1

NUMBERS OF SMALL MAMMAL SPECIES

No. of individuals

=
3
: S

= &g = =
a 3 8 S = =
zs Vg nc bee ee arg oe a,
Locality Trapnights ies po See Se ee eg
Kingscliff 988 — — 3 ~—~ —~ — I — — — —
Evans Head 3648 25 — — 4 — 18 16 — —
Yamba pas) — — gs’ — — — 5 —_- — ——
Diggers Camp 3648 SS 8 22S Se
Bonny Hills 524 BQ BAD EO a Ag of BS AS eet
Port Stephens 5472 1 Peo 2eos-) = ST 14. ee 5
Pearl Beach 1920 5-9 PS 2 ta ee Sa
Nadgee Nature Reserve 15240 94° 21) — —,; — — — 1 9 =— 2
Total 35692 UPA RAD A Gig ee oSt 57 CL OL fo wD Ged

Aust. Zool. 18(2), 1974 67

H. POSAMENTIER and H. F. RECHER

Marlow (1968) point out that there are subfossil bone deposits from caves in
Tasmania and western Victoria which are remains of P. novaehollandiae.

HABITAT REQUIREMENTS

Previous to our studies, P. novaehollandiae was teported mainly from dry
sclerophyll forest with a sclerophyllic understorey (Keith and Calaby, 1968;
Seebeck and Beste, 1970; Batt et. al., 1972). Keith and Calaby (1968) also
found it between dunes at swamp edges and we have trapped it extensively in
coastal heaths. In these places, P. novaehollandiae is as abundant as other native
and introduced small mammals (Table 1). However, on the north coast of New
South Wales, the distribution of P. novaehollandiae is not uniform and there
appears to be extensive areas where it is absent or rare. The reason for this patchy
distribution is not clear, but is probably in part historical and in the part the
animal’s response to differences between and within habitats.

In our studies of small mammal communities, we attempt to relate the
presence or absence of a species to the structure of the habitat. Plant species
composition, soil firmness, total vegetation density, foliage height density, extent
of bare ground, number of plant species and amount of leaf litter are measured
and the presence or absence of small mammals is determined by trapping. A
brief analysis of the major habitat requirements of P. novaehollandiae is essential
to an understanding of its distribution and abundance.

PABEE OZ
OCCURRENCE OF P. NOVAEHOLLANDIAE AND M. MUSCULUS IN DIFFERENT
HEATHS
Substrate Heath Type No. of P. novaehollandiae M. musculus
Plots A B A B c
Rock headland—dry 26 0 0.0 0 0.0 0
headland—moist 9 0 0.0 1 0.1 0
Sand foredune 5 0 0.0 3 1.6 0
moist 22 6 0.5 iM 331 2
dry 18 ifs 1.6 10 1.6 8
heavily disturbed 8 2 0.6 4 0.9 1
Total 88 Di 29 11
A—Number of plots with animals C—Number of plots with
B—Average number of animals per plot both species

Various workers have commented on the possible importance of the seeds
of legumes (e.g. Acacia, Dillwynia) as food of P. novaehollandiae in forest
habitats (e.g. Keith and Calaby, 1968). Our observations support this suggestion:
the forest habitats in which we have taken this mouse comprise a dense sclero-
phyllic understorey with a high proportion of legumes. However, in coastal heaths,

68 Aust. Zool. 18(2), 1974

STATUS OF PSEUDOMYS NOVAEHOLLANDIAE

the presence of P. novaehollandiae is associated with a small number of habitat
variables, but not obviously with the proportion of legumes.

The heaths sampled in this study have been grouped according to substrate,
substrate moisture and degree of disturbance (Table 2). Moisture relations and
the level of disturbance were ranked and each plot placed in the appropriate
class. All of the heaths sampled have a modern history of fire and grazing. The
heaths at Nadgee on the south coast of New South Wales differ from those
on the north coast in having been protected from fire and grazing for the last
20 years. The north coast heaths showed the effects of more recent disturbances.
P. novaehollandiae was not found on any of the plots at Nadgee and on the
north coast it was most abundant in dry and only moderately disturbed heaths
(Table 2). Numbers were lower in recently and heavily disturbed heaths and
where P. novaehollandiae occurred in wet heaths, it was where such heaths were
associated with and surrounded by dry heaths. The wet heaths sampled have
probably been less affected by fire and grazing than the dry heaths.

The grouping of heaths into rather broad habitat types does not tell us
why P. novaehollandiae occurs in one place and not another. To answer this
question we need to know the features of the habitat which are important to
the animal and how these features differ between plots where the animal is present
and where it is absent. One method of analysis is to use ‘stepwise discriminant
analysis’ (Sokal and Rohlf, 1969) to obtain a measure of how well the respective
environmental parameters which we measured separate the plots in which
the mouse is present from those in which it is absent. The results of this
analysis are summarized in Table 3. Only the foliage density at the 20 cm
height level was used as this variable was highly correlated with foliage densities
at other heights (i.e. 10, 50, 100 and 150 cm) and of all the foliage densities
measured it gave the best association with the presence of P. novaehollandiae.

The analysis presented in Table 2 is based on fifty plots which were
sampled twice in different years. We found that there was no association with the
number of plant species, with soil firmness or with the composition of the
plant community. There was a positive but insignificant association with the

TABLE 3

ASSOCIATION BETWEEN P. NOVAEHOLLANDIAE AND HABITAT VARIABLES

Variable Association F-value d.f. p
Vegetation density at 20 cm positive S225 1.48 0.1
Bare ground positive 16.25 1.48 0.001
Total vegetation negative 4.62 1.48 0.05
Bare ground and vegeation at 20 cm positive 15.15 2.47 0.001

The results have been derived from 50 plots trapped twice.

Aust. Zool. 18(2), 1974 69

H. POSAMENTIER and H. F. RECHER

amount of leaf litter. It is interesting that while there was a positive association
with vegetation density at 20 cm, there was a negative association with total
vegetation. The most important single variable was the amount of bare ground.
Using two variables, the amount of bare ground and vegetation density at 20 cm
increases the separation between plots with and without P. novaehollandiae.
Studies now in progress suggest that these features are characteristic of heaths
regenerating after fire.

DISCUSSION

Our observations suggest that an optimum habitat for P. novaehollandiae
is a dry heath which has been disturbed by fire and is actively regenerating. The
positive association with bare ground, foliage density at 20 cm and leaf litter
and the negative association with total vegetation support this conclusion.
Similarly its abundance in sclerophyll forests with a well developed sclerophyllic
understory and a high proportion of legumes suggest an animal which does
well in the early and: middle successional stages following fire.

Habitats protected from fire are as unsuited for P. novaehollandiae as are
the first stages of succession after fire or habitats in which succession is disrupted
by too frequent burning, heavy grazing or mining. A similar set of habitat
requirements has been reported by Ahlgren (1966) for granivorous rodents in
North America. Baker (1971) also noted that granivores invaded disturbed
areas because seeds and insects became more abundant and readily available
during the early stages of succession. It is particularly interesting that Christensen
(pers. comm.) working in Western Australia, Leonard (1973) in Victoria and
Recher in southeastern New South Wales (unpub. data) have found that Mus
musculus invades recently burned forests and heaths, increases rapidly in abundance
and then becomes rare as succession proceeds and food presumably becomes less
abundant. Based on the evidence available, it is not unreasonable to assume that
P. novaehollandiae has a similar ecology.

One other variable merits discussion. The introduced M. musculus is widely
distributed on the coast of New South Wales and occurs abundantly in habitats
used by P. novaehollandiae (Table 2). As these rodents are very similar in size,
appearance and probably in feeding habits, it is possible that they use similar
resources. If so, the distribution and abundance of the native mouse might be
affected by the presence of the introduced animal. Of the 88 plots sampled
one or more times inthis survey, P. mnovaehollandiae occurred on 21 and
M. musculus on 29 (Table 2). They occurred together on 11 of the plots,
but there is no indication that the abundance or distribution of P. novaehollandiae
has been affected by the introduced M. musculus. Of the 18 plots on which
M. musculus was present without P. novaehollandiae, only two had habitat
characteristics (i.e. extent of bare ground and foliage density at 20 cm) suitable
for P. novaehollandiae. There is a negative association between M. musculus
and P. novaehollandiae which is almost significant (0.1 p 0.5). It is possible,

. 10 Aust. Zool. 18(2), 1974

STATUS OF PSEUDOMYS NOVAEHOLLANDIAE

therefore, that the two species have apportioned the available range of heath
habitats among themselves and that the modern habitat range of P. novaehollandiae
is less than that used prior to the advent of M. musculus. Historically, M. musculus
has been present in Australia since the later part of the 18th century and there
has been adequate time for any competitive interactions between P. novaehollandiae
and M. musculus to have come to an equilibrium with a separation by habitat.

Regardless of any part interactions that may have occurred between M.
musculus and P. novaehollandiae, the latter has a wide range from northern
New South Wales to southern Victoria and occurs in forest as well as heath habitats.
Though its distribution within this range is patchy, it can not be considered
an endangered species. We prefer to consider this mouse an opportunistic or
fugitive species; one which responds quickly to favourable conditions with a
great increase in numbers and then becomes uncommon or rare as the habitat
changes or as other species of small mammals colonize the area.

ACKNOWLEDGEMENTS

We would like to thank Graeme Caughley, Jack Mahoney and Frank Talbot for
their helpful comments during the preparation of this manuscript. We are especially
indebted to Jack Mahoney and Basil Marlow for their continued assistance in identifying
material and to Steve Clark for profitable discussions on ways and means of measuring
mammal habitats and for assistance in the field.

REFERENCES

AHLGREN, C. E. (1966). Small mammals and reforestation following prescribed burning.
_J. Forestry 64, 614-618.

BAKER, R. H. (1971). Nutritional strategy of myomorph rodents in North American
grasslands. J. Mamm. 52, 800-805.

BATT, J., C. GREAGH, P. HUGHES, G. McPHERSON, J. PETERS & J. WADDY
(1972). The New Holland Mouse, Pseudomys novaeholalndiae (Waterhouse), in the
Smith’s Lake Area. In ‘Ecological investigations in the Myall Lakes area’ edited
by F. J. Burrows and K. E. Mount. Biology, Macquarie University.

CALABY, J. H. (1969). Australian Mammals since 1770. Aust. Nat. Hist. pp. 271-275.

KEITH, K. & J. H. CALABY (1968). The New Holland Mouse, Pseudomys novaehollandiae
(Waterhouse) in the Port Stephens District, N.S.W. C.S.LR.O. Wildl. Res. 13, 45-58.

LEONARD, B. (1970). Effects of Control Burning on the Ecology of Small Mammal
Populations. Zn Symposium Papers, Forests Commission, Victoria.

MAHONEY, J. A. & B. J. MARLOW (1968). The Rediscovery of the New Holland
Mouse. Aust. J. Sci. 3/7, 221-223.

RIDE, W. D. L. (1970). A Guide to the Native Mammals of Australia. Oxford University
Press, Melbourne.

SEEBECK, J. H. & H. J. BESTE (1970). First Record of the New Holland Mouse
(Pseudomys novaehollandiae) (Waterhouse, 1843) in Victoria. Vic. Nat. 87, 280-287.

SOKAL, R. R. & F. J. ROHLF (1969). Biometry. W. H. Freeman and Company, San
Francisco and London.

Aust. Zool. 18(2), 1974 71

Occurrence and Field Recognition of
Macropus parma

G. M. MAYNES

‘ School of Biological Sciences, Macquarie University,
North Ryde, New South Wales, 2113.

ABSTRACT

The parma wallaby, Macropus parma, is recorded from the Dorrigo Plateau
for the first time in 40 years. The parmas were found feeding in areas of disturbed
wet sclerophyll forest and appeared to be coming from areas of rainforest
and wet sclerophyll forest with a rainforest understorey. Characteristics of the
species which will aid in field recognition are detailed.

INTRODUCTION

_ The parma wallaby, Macropus parma, was originally discovered in the Illawarra
district of New South Wales by John Gould who listed the name Halmaturus
parma for the species (Gray, 1843). Gould, however, failed to describe the
species and this was done by Waterhouse in 1846.

In the early 1840’s, Gould wrote to John Gilbert, “Three kinds of wallaby
run in the brushes of Illawarra, viz. Halmaturus ualabatus |= Wallabia bicolor],
H. tithys (the common pademellan, a red-necked kind [= Thylogale thetis])
and a nearby allied species called ‘Pama’ by the natives. Of this latter which
is very like Derbyanus, [= Macropus eugenii], I wish as many specimens and
crania as convenient”? (Longman, 1922). Unfortunately, Gilbert was speared
to death by aborigines during the ill-fated Leichhardt expedition to Port Essington
in 1845 before he could collect further specimens for Gould.

Twenty years later, Gould wrote, “In these brushes, it doubtless still exists,
as since my return, other specimens have been sent to me by the late Mr.
Strange. How far its range may extend westwardly [south] to Port Phillip or
eastwardly [north] in the direction of Moreton Bay, I am unable to state”

(Gould, 1863).

Ride, in 1957, could find only 12 specimens in the world museums and
records of 4 others which could no longer be located. He reported the species
as apparently extinct (Ride, 1957).

Prior to the turn of the century, all specimens came from the south coast
region of New South Wales. The last specimen recorded south of Sydney was
taken at Mawarra, Sassafras (approx. 35° 05’ S, 150° 15’ E) in 1889.

72 Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

In 1921, a single specimen was taken at Point Lookout Gorge (30° 30’ S,
152° 24° E) near Ebor by H. Raven who was collecting for the American
Museum of Natural History. This locality may now lie inside the boundary
of the New England National Park.

Two other specimens were taken by P. J. Darlington at Cascade (30° 16’, S,
152° 47-18 E) 22 km. north of Dorrigo in February, 1932. Darlington
recorded taking the male in a pen gum forest (sic) on a ridge and the female
in an open space in a pen gum forest (sic) on the side of a ridge (Darlington,
field notes).

Johnson and Mawson (1939, 1940) described several parasites from the
parma wallaby using material collected by Luke Gallard in the Ourimbah
region (33° 03’ S, 151° 21’ E) in 1909-11. The Gallard collection is of parasites
only and no voucher specimens were entered with the collection. However, in
1965, Mr. E. Worrell of the Australian Reptile Park at Gosford received a
female M. parma with a male pouch young, reportedly from the same general
district. The skin and skull of the female are now lodged in the National Museum
of Victoria register number C 10725. The exact locality of the population
from which this animal came is yet to be determined.

In 1967, a population of the parma wallaby was. reported on Kawau
Island, New Zealand (36° 25’ S, 174° 51’ E) where they had been taken
about 1870 by Sir George Grey (Wodzicki and Flux, 1967).

A field survey to determine the current distribution and status of the
parma wallaby in New South Waes was commenced in 1972. This paper reports
the discovery of a population of M. parma in Moonpar State Forest (30° 12’ S,
152° 41’ 25” E) on the Dorrigo Plateau and details field recognition characters
for this species.

METHODS

A single specimen of M. parma was collected in Moonpar S.F. on the last
night of a nine day field trip to Cascade in March, 1972. A further four weeks
was spent in Moonpar S.F. between June 26 and July 21, 1972 during which
time I attempted to determine recognition characters for separating M. parma
from the two pademelons Thylogale thetis and Thylogale stigmatica.

The survey was conducted from a Land-Rover as outlined by Calaby (1966).
During the day much of the area was inspected on foot.

Ages were estimated using data in Maynes (1972).

RESULTS
(a) Habitat
(i) Topography and Soil
Moonpar State Forest (30° 12’ S, 152° 41’ 25” E) lies on the northern
edge of the Dorrigo Plateau at the junction of the Blicks and Nymboida Rivers and

Aust. Zool. 18(2), 1974 73

G. M. MAYNES

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Aust. Zool.

74

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

covers approximately 3 360 ha. It is an area of undulating hills ranging from 425
m, in the river valleys to a maximum height of 855 m, with the majority of the
forest between 550 m, and 730 m. (Forestry Commission N.S.W. Map. Misc.
F 842).

McArthur (1964) has surveyed the soils of the Dorrigo Plateau and described
a Moonpar Association comprised of red podzolic soils on granite. The surface
layer of soil is a dark brown gritty loam below which is a red gritty clay
at about 0.5 m. (McArthur, 1964). Although McArthur did not survey Moonpar
S.F. itself, one of his sample profiles was 400 m. from the southern entrance to
the forest. It would appear that this association applies throughout most of the
State Forest.

(iit) Climate

The area has a high annual rainfall with the period December-March being
the most reliable. Table 1 gives the mean annual rainfall for the three nearest
stations to Moonpar S.F. These are Bobo (13 km. east), Cloud’s Creek (13 km.
north north west) and Dorrigo (13 km. south west).

Table 2 records the mean temperature data for Bobo and Cloud’s Creek
stations. The average frost-free period at these stations is 241 days and 200
days respectively. The winters are mild with heavy frosts at times, especially
within small openings in the forest.

(111) Vegetation

The dominant forest types have been mapped by the Forestry Commission
of New South Wales. Moonpar S.F. consists mainly of rainforest and wet
sclerophyll forest with some scattered areas of dry sclerophyll forest (Forestry
Commission Map Misc. F 842). The rainforest generally occupies the gullies
while the eucalypt forest is usually on the ridges.

The rainforest is a coachwood (Ceratopetalum apetalum)—Crabapple
(Schizomeria ovata) type in which coachwood is clearly dominant. Associated
species include Hoop Pine (Araucaria cunninghamii), Red Carabeen (Geissois
benthamii), Silver Sycamore (Cryptocarya glaucescens), Sassafras (Doryphora
sassafras), Corkwood (Endiandra sieberi), Lilly Pilly (Acmena smithit) and
Black Myrtle (Backhousia myrtifolia).

The areas of wet sclerophyll forest are dominated by Sydney Blue Gum
(Eucalyptus saligna), Tallowwood (E. microcorys) and Blackbutt (E. pélularis).
There is generally a dense mesophyllic shrub layer of rainforest species.

In the northern section of the forest, control burning and grazing is
maintaining an open woodland habitat dominated by Sydney Blue Gum, Blackbutt,
Tallowwood and New England Blackbutt (E. campanulata) (G. Grey, pers.
comm.). The forest oak (Casuarina torolosa), black she-oak (Casuarina litoralis ),

Aust. Zool; 18(2), 1974 75

G. M. MAYNES

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Aust. Zool.

76

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

black wattle (Acacia irrorata), Sally wattle (Acacia floribunda), bracken fern
(Pteridium sp.), blady grass (Imperata sp.), kangaroo grass (Themeda australis),
Poa (Poa caespittosa), Dogwood (Jacksonia coperia) and other species were
observed in this area.

(b) Field recognition of M. parma

(i) Flesh measurements

Body measurements of the parma wallabies collected in Moonpar S.F. are
given in Table 3.

There is considerable overlap in the size of the hind foot between M. parma
(103-147 mm), the red-necked pademelon (Thylogale thetis) (102-151 mm)
and the red-legged pademelon ( Thylogale stigmatica) (122-134 mm, adults only).
Consequently, attempts to determine specific differences between the foot prints
were abandoned.

TABLE 4
TAIL LENGTH AS A PERCENTAGE OF BODY LENGTH

Species n x + SE. % Range %
M. parma 28 99.91 + .65% 106.2-94.0
T. thetis 23 88.05 + .81% 95.4-81.1

ey 2 90 85.6-73.8

T. stigmatica 6 Jo2

Table 4 gives the value of the tail length expressed as a percentage of
the body length for M. parma and both T. thetis and T. stigmatica. The parma
value is based on animals from Kawau Island, New Zealand, while the pademelon
values are from animals collected in northern New South Wales. ( All measurements
were taken by the author). Values for the five parmas taken in Moonpar S.F.
were 91.1, 95.8, 96.5, 99.2 and 102.3%.

As can be seen from Table 4, parmas have the longest tail relative to the
body size of all three species. This can be observed, under suitable conditions,
in the live animal at some distance.

(ii) Gait

The appearance of M. parma and T. thetis in profile are shown in Figs 1 and 2.
When hopping at medium pace, parmas carry their tail curved upwards in a
shallow U-shape. Pademelons in contrast tend to carry the tail either straight
out behind them or only slightly curved. Pademelons’ tails appear to bob up
and down more obviously and vigorously than do parmas. When travelling at

Aust. Zool. 18(2), 1974 2 77

78

G. M. MAYNES

Profiles of parma _ wallabies.

Fig. 1.

Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS

Aust. Zool. 18(2), 1974

PARMA

Profiles of red-necked pademelons.

Fig. 2.

79

G. M. MAYNES

speed, the tail is held straight out behind in all species. Parmas appear to
remain close to the ground when hopping and maintain an almost horizontal
position. Pademelons seem to “bounce”? more when hopping.

Pademelons generally appear to be somewhat stockier than parmas and
tend to be heavier in the lower abdominal and pelvic regions. From rear view,
parmas generally appear thinner in the hips than pademelons which are more
circular in outline.

(iii) Pelage

In the hand, T. thetis is readily recognized by its prominent rufous-brown
shoulders and the lack of any cheek stripe or nuchal stripe. T. stigmatica has
the back and shoulders a uniform dark grey. The cheeks, a patch above the
eyes and the base of the ears are reddish-brown. There is an indistinct white
cheek-stripe and an indistinct dark nuchal stripe.

M. parma has a uniform brownish grey back with a black stripe down
the spine that ends in the mid-back. There is a well developed white cheek
stripe along the upper lip and the rest of the head is a grizzled grey. The
shoulders appear only slightly lighter than the rest of the back.

In some M. parma, there is a white tip to the tail which is usually 20-40
mm long when present. White tail tips have also been observed in some
T. stigmatica but never in T. thetis.

Although parmas invariably have a white chest and throat, some specimens
of both species of pademelon have been observed with this same condition.
The presence of reddish-brown heels cannot be considered alone as a sufficiently
good character for identifying T. stigmatica in the field. Some T. thetis have
heels almost as red as those observed in T. stigmatica while the M. parma collected
in Moonpar also had very red-brown heels. Faint hip stripes have been observed
in all three species.

(iv) Skull characteristics

M. parma can be readily separated from the two species of pademelons by
the position of the groove on the labial surface of i*. The outer lamina is
about 60% of the length of the whole tooth in M. parma, about 80% in T.
stigmatica and about 95% in T. thetis (Fig. 3). The permanent premolar
is about equal in length to m? in M. parma while it is 20% and 70% larger
than m! in T. thetis and T. stigmatica respectively (Fig. 3).

The differences in the shape of the nasal bones between the three species
is shown in Fig. 3.

In M. parma the suture between the maxilla and palatal bones on the
bridge between the palatal foramina lies about a third from the front. This
suture is in the middle of the bridge in both species of pademelon (Fig. 3).

80 Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

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Aust. Zool. 18(2), 1974

G. M. MAYNES

The incisive foramina (Fig. 3) are long and thin in M. parma (about
6.5 mm long) whereas they are more uniform in outline and smaller in the
pademelons. They are about 4.5 mm long in T. thetis and about 5.4 mm in
T. stigmatica. :

(v) Faecal Pellets

Two distinctive types of small macropodid droppings were found in the
study area. Identification of the species producing the droppings was done by
examining the rectal contents of shot animals.

Parma droppings are flattened, square to slightly rectangular pellets (Figs.
4, 5). Faecal pellets produced by the red-necked pademelon are elongate cylinders
and tend to be less regular in outline (Figs. 5, 6). The few droppings collected
from T. stigmatica resembled those from T. thetis rather than M. parma.

A collection of faecal pellet groups from four separate sites was taken
on the last day of the July field-trip to assess whether the pellet-group count
technique could be applied to a study of the biology of M. parma. No T. stigmatica
were seen or collected at any of these sites during the entire period spent
in Moonpar S.F. so all pademelon droppings were attributed to T. thetis.

TABLE 5

SPECIES COMPOSITION OF FAECAL PELLET GROUPS AT FOUR SITES IN
MOONPAR STATE FOREST

Site n M. parma T. thetis Uncertain Other Species

1 293 50% 3217-9 92% 8.1%
2 260 3.5% 86.1% 4.6% 5.8%
3 112 24.1% 33.0% 10.7% 522%
4 a1 23.0% 32.0% | 3.3% 41.7%

Table 5 gives the percentage species composition of the pellet groups at the
four sites. Specimens of M. parma were collected at all sites except site 2 where
a single parma was observed but not collected. Not all pellet groups could be

CAPTIONS FOR PLATE OPPOSITE

Fig. 4. Upper surface view of two parma wallaby faecal pellet groups. Juvenile on
left, adult on right.

Fig. 5. Lateral view of parma wallaby pellet (upper) and pademelon pellets (lower).
Note the much greater flattening of the parma pellet.

Fig. 6. Upper surface view of three faecal pellet groups from red-necked pademelons. All
figures natural size.

82 se Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

Fig. 4
Fig. 5
OOD LL ce Ee
G ‘ oe 4 . S .& 7 g
Fig. 6

Aust. Zool. 18(2), 1974 ; 83

G. M. MAYNES

identified with certainty as either parma or pademelon and these have been
recorded as uncertain. The other species category is mainly Wallabia bicolor but
includes some Macropus giganteus and Macropus rufogriseus droppings.

There is no apparent difference in the pattern of production of pellet
groups between M. parma and T. thetis (Table 6).

TABLE 6

NUMBER OF PELLETS PER GROUP

Number 1 2 3 4 5 6-8 Total
% % % % % % Groups

M. parma 23.6 40.0 24.1 7.4 3.0 2.0 203

T. thetis 24.4 40.4 20.2 11.4 2.6 0.8 386

(c) Mammal Distribution

The platypus, Ornithorhynchus anatinus, occurred in both the Blicks and
Nymboida Rivers which flank Moonpar S.F. Within Moonpar, were six species
of macropodids besides M. parma. Both species of pademelon, T. thetis and
T. stigmatica, occurred in the rainforest and wet sclerophyll forest areas. T.
stigmatica was observed on only a few occasions and would be considered
one of the less common species of the area. Wallabia bicolor, the swamp wallaby,
was widely distributed throughout the forest in both rainforest and wet sclerophyll
forest areas.

A single wallaroo, Macropus robustus, five eastern grey kangaroos, Macropus
giganteus, and six red-necked wallabies, Macropus rufogriseus, were observed
feeding on Mills Farm and the adjacent areas which had been opened up for
grazing. Both the grey kangaroos and the red-necked wallabies were more
common in the open forest to the west. M. parma, T. thetis and W. bicolor
have all been observed feeding in the same areas at one time or another. These
three species have also been seen feeding in areas grazed by the grey kangaroos
and red-necked wallabies.

Five species of phalangerids were observed in Moonpar. Trichosurus caninus
and Pseudocheirus peregrinus were observed in rainforest areas, while Trichosurus
vulpecula, Petaurus breviceps and Petaurus australis were seen in the dry sclero-
phyll forest and open woodland area in the north.

Several tiger cats, Dasyurus maculatus, were heard and one was observed
crossing the road in a rainforest area. A small mammal which appeared to be an
Antechinus was also observed on the road.

84 Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

Feral cats, Felis catus, were present in the forest as was the dingo, Canis
familiaris dingo.

(d). Biology of M. parma

Examination of the area in which parmas were most common based on
the analysis of faecal pellet groups indicated that kangaroo grass (Themeda
australis) was the preferred food in that area. Patches of kangaroo grass were
grazed to lawn level whereas only small amounts of Poa had been eaten and
the blady grass (Imperata sp.) was virtually untouched.

Evidence was found of predation by both the carpet snake (Morelia spilotes)
and the dingo upon the red-necked pademelon. It must be assumed that these
species would also take parmas if the opportunity arose. The tiger cat and
feral cat are other potential predators but no evidence was found for predation
by either of these species upon macropodids.

Four of the five specimens of M. parma collected were feeding with other
animals, but there was no other evidence that they were part of a social group.
All animals appeared to make for the rainforest and thicker wet sclerophyll
forest areas when disturbed.

The estimated months of birth of the four animals less than two years
old and the two pouch young are February (2), March (2), June and October.
Estimated age at first birth for 73/5P was 70 weeks old which agrees with
the data obtained from yard animals (Maynes, 1973). Both 72/1P and 72/2P
would appear to have begun to mature sexually as judged by their pouch condition.
Thus the weight at sexual maturation is probably between 2.5 and 2.8 Kg in the
female.

DISCUSSION

The habitat of the parma wallaby would appear to be rainforest and wet
sclerophyll forest. This agrees with Gould’s statement that he had seen it in the
brushes (i.e. rainforest) of the Illawarra area (Gould, 1863). Both Darlington’s
specimens and my own were taken in eucalypt forest where they were feeding.
It may be that the parmas feed in the more open eucalypt forest areas and shelter
in or on the edges of the rainforest areas.

One reason for the apparent rarity of the parma wallaby may be the difficulty
in recognising it in the field from the two species of pademelon. Most sightings
are at night, of short duration, and of moving animals as they run into the
brush on the side of the road or as they cross the road. Under such conditions,
some experience is needed in recognising the three species. Parmas are generally
lighter in build than the pademelons. The length of the tail in relation to the
body is considered the most useful character for recognising parmas in the field.
If one observes a pademelon-sized animal with a long white-tipped tail, it is a

Aust. Zool. 18(2), 1974 85

G. M. MAYNES

parma wallaby. The presence of the distinct white cheek stripe can also be used
for identifying them provided one can observe them long enough.

The presence of distinctive parma wallaby pellets in Moonpar S.F. suggests
that this may be a useful means of checking for the presence of the species.
It also holds promise as a means of assessing the relative use of different areas
by.parmas, pademelons and other species.

The parma wallaby appears to be in no immediate danger in Moonpat Salts
provided there is no radical change in current management practices. It is to be
hoped that it will be found to be more widely distributed in the state forests
of the district. Further field survey work is required to assess the distribution,
habitat preference and status of this species in New South Wales.

The discovery of this population of parma wallabies in New South Wales
suggests that there may be no need to repopulate Australia with animals from
Kawau Island as has been contemplated in some quarters (Anon., 1973). It is
strongly recommended that no releases into the wild be attempted until detailed
studies have revealed the ecological consequences of such an action.

ACKNOWLEDGEMENTS

ng am grateful ‘to the National Parks and Wildlife Service for licences to collect
the animals reported on in this paper. Thanks are also due to the Forestry Commission
of New South Walés for permission to work in State Forests and for supplying maps.

I am indebted to Mr. A. Floyd, Research Officer, Forestry Commission N.S.W.,
Coffs: Harbour for Plant identifications in Moonpar. I thank Mr. G. Baur, Forestry
Commission, N.S.W. for climatic data from Bobo and Cloud’s Creek stations. I am also
indebted to Mr. G. Gray, Forester, Dorrigo, Mr. G. King, Assistant Forester, Dorrigo
and to Misses L. Day, A. Dunn, P. Kenway, and Messrs. R. Close, K. Bishop, P. Blanton,
D. Hockley and J. Lindsay all of Macquarie University for field assistance at various
times. Copy of P. J. Darlington’s field notes was supplied by C. W. Mack.

I thank Mr. W. Meldrum for Lieut 1, Miss B. Thorn for figures 2 and 3 and
Mr. P. Jamieson for photographs.

_.. This research was supported by funds from the Macquarie University Reseaten Grant
Fund.

REFERENCES

ANONYMOUS (1973). Australian Wildlife 6 (3), 16.

CALABY, ie GLI Ge): Mammals of the, Upper Richmond and Clarence: Rivers, New
South Wales. CSIRO Division of Wildlife Research, Technical Paper No. 10.

GOULD, jai es (1863). “The Mammals of Australia”. (Gould, London).

GRAY, J. E. (1843). List of the specimens of mammalia in the British Museum.
The British Museum (Nat. Hist.): London, 91.

JOHNSTON, T. H. & P. M. MAWSON (1939). Strongylate nematodes from marsupials
in New South Wales. Proc. Linn. Soc. N.S.W. LXIV, 513-536.

86 Aust. Zool. 18(2), 1974

OCCURRENCE AND RECOGNITION OF MACROPUS PARMA

JOHNSTON, T. H. & P. M. MAWSON (1940). New and known nematodes from Australian
marsupials. Proc. Linn. Soc. N.S.W. LXV, 468-476.

LONGMAN, H. A. (1922). John Gould’s notes for John Gilbert.. Memoirs Qld “Mus.
VII, 291-294.

McARTHUR, W. M. (1964). Soils and Land Use in the Dorrigo-Ebor-Tyringham ‘area,
New South Wales. CSIRO Soils and Land Use Series No. 46.

MAYNES, G. M. (1972). Age Estimation in the parma wallaby, Macropus parma
Waterhouse. Aust. J. Zool. 20, 107-118.

MAYNES, G. M. (1973). Reproduction in the parma wallaby, Macropus parma Water-
house. Aust. J. Zool. 21, 331-351.

WODZICKI, K. A. & J. E. C. FLUX (1967). Rediscovery of the White-throated wallaby,
Macropus parma Waterhouse 1846, on Kawau Island, New Zealand. Aust. J. Sci. 29,
429-430.

Aust. Zool. 18(2), 1974 87

Craterocephalus dalhousiensis n.sp., a Sexually
Dimorphic Freshwater Teleost (Atherinidae) from
South Australia

W. IVANTSOFF

School of Biological Sciences, Macquarie University,
North Ryde, New South Wales, 2113.

C. J. M. GLOVER
South Australian Museum, Adelaide, South Australia, 5000.

ABSTRACT

Craterocephalus dalhousiensis n.sp. is described from specimens collected
in shallow water of artesian springs at Dalhousie Springs, South Australia and
represents the first Australian record of a sexually dimorphic atherinid.
C. dalhousiensis appears to be closely related to C. /acustris and C. ster-
cusmuscarum although meristic and morphometric differences make the new
species readily distinguishable. A number of specimens of C. dalhousiensis
has been examined internally for gonad condition, gut content and presence of
parasites. Water temperature, salinity and pH were measured at the time of
collection and upper and lower thermal tolerances were also determined.

INTRODUCTION

As a result of aquatic surveys conducted by the South Australian Museum
in the Far North of South Australia since 1968, the known distributions of
several species of fishes have been extended and some new forms have been
collected, including the species described here.

Preliminary field observations and experiments on this new species’ environ-
ment, activity and thermal physiology can be compared with the findings of
Brown and Feldmeth (1971) on desert pupfish (Cyprinodon spp.) in southern
California.

(A) TAXONOMY
MATERIALS AND METHODS

The technique used for counts and measurements was the same as used by
Munro (1967) except for the inter-dorsal count where the scales were counted
from the origin of the last spine of the first dorsal to the origin of the second
dorsal. Transverse scale rows count was made diagonally across the body from the

88 Aust. Zool. 18(2), 1974

A NEW. FRESHWATER TELEOST °

origin of the first dorsal to the origin of the ventral fin. Relative positions of
the fins were recorded in numbers of scales in front or behind the origins or tips

of fins.

All measurements were made with dial calipers to the nearest one-tenth
of a millimetre (mm). Body measurements were expressed as proportions of
standard length or as otherwise indicated in Table 1. A mean and range (in
brackets) was also given for the ten male and ten female specimens examined.

Type material has been deposited at the following institutions: Australian
Museum, Sydney; American Museum of Natural History, New York; British
Museum of Natural History, London; Museum National d’Histoire Naturelle, Paris;
Zoologisch Museum, Amsterdam.

List of abbreviations used in Table 1:

DL standard length

B behind

F in front of

OD1 origin of first dorsal fin

OD2 origin of second dorsal fin

OV origin of ventral fin

iF, “Pee: tip of pectoral fin

TV tip of ventral fin
DESCRIPTION

Craterocephalus dalhousiensis, new species

Figure 1

Holotype—SAM F3453 (male), allotype—SAM F3453 (female), type locality, Dalhousie
Springs, Main Spring, South Australia, 26° 25’ S., 135° 30’ E. (Map ref. 1:250,000
Topographic Maps, Series R502, Commonwealth Government: Printer, Canberra. Map Sheet:
Dalhousie SG 53-II. Grid ref.: 345718). Holotype and allotype in South Australian
Museum (SAM). Paratypes in South Australian Museum and The Australian Museum,
Sydney (AMS).

Material examined:

Main Spring, Dalhousie Springs, South Australia, designated holotype SAM _ F3453
(male), 51 mm SL, designated allotype SAM F3453 (female), 63 mm SL, designated
paratypes SAM F3453, AMS I.17756-001; Spring, 2.4 km south-west of Dalhousie Main
Spring, South Australia, SAM F3466.

General description—The material examined ranged from 27 to 63 mm SL.
In both sexes, the lips are thick and fleshy with the skin of the upper jaw
fusing with the skin over lower j jaw about half way along the premaxilla. Premaxilla
never reaches the anterior margin of the. orbit. , pits) ee

Aust. Zool. 18(2), 1974 , 89

W. IVANTSOFF and C. J. M. GLOVER

The lower jaw is massive when compared with other species of. this genus,
the anterior end expanding to form a wide horseshoe-shaped band with many
rows of small but easily visible curved teeth. Upper jaw is similarly toothed. The
test of the mouth is edentelous. Both the preopercle and opercle are covered
with scales. Body scales are cycloid, round in smaller specimens but becoming
elongated vertically in older fish. The ridges on scales do not extend to the
posterior part of the scale in specimens examined. As in all other Australian
hardyheads, the midlateral scales are pierced with pores, but in many specimens
scales on the dorsal surface and on the lateral sides are likewise pierced.

_ Specimens preserved in alcohol are dark brown above the midlateral band
due to presence of many small chromatophores uniformly dispersed along the
back. Occasionally, spots immediately above the midlateral band form a faint dis-
continuous line. A dark band extends from the snout through the eye and opercle,
then continues as a midlateral band formed by large pigment spots in the centre of
each midlateral scale. Scales beneath the midlateral band have smaller pigment spots
usually forming two or three discontinuous lines along the abdomen. Very young
fish are dusky brown with few or no spots below midlateral band.

Fig. 1. Line drawings of holotype male (above) and allotype female (below) showing
external features. Scales (6th along the midlateral band) from both specimens.
Line diagrams of premaxilla, maxilla and dentary are also included. Position of
anus is indicated by arrow. Projected scale, 10 mm.

90 Aust. Zool. 18(2), 1974

A NEW FRESHWATER TELEOST

TABLE 1

BODY MEASUREMENTS AND COUNTS OF TWENTY-TWO SPECIMENS
OF C. DALHOUSIENSIS FROM DALHOUSIE SPRINGS

Mean, range Mean, range

Holotype Allotype of 10 males of 10 females

Head in SL 3.4 51 Bey Es se) 3.3 (3.0-3.5)
Greatest body depth in SL 3.9 3.8 4.0(3.7-4.7) 4.1(3.7-4.5)
Premaxillary process in eye tel ia 1.3(1.1-1.7) 1.2(1.0-1.6)
Eye in the head 3.8 4.4 3.6(3.0-3.9) 3.8(3.3-4.4)
Interorbital in head 2.6 ne! 2.6(2.3-2.9) 2.6(2.4-3.1)
Snout in eye 1a 1.0 11 G0et3) 1.1(0.9-1.2)
Premaxilla in eye 1.0 1.0 1.0(1.0-1.4) 1.0(0.8-1.1)
Least depth of caudal (ae

peduncle in SL 8.7 10.2 9.5(8.7-10.0) 10.0(9.4-10.1)
Midlateral scales 30 Sle 29.6(29-30) 29.9 (29-31)
Transverse scales Ped 75 7.1(6-8) F.36.5-7-5)
Predorsal scales 17 26 15.9 15=17) 16(15-=18) 42
Interdorsal scales 6 8 7(6-8) 7.6(7-8)
Dorsal fin count V,.116 VI,1i6 V-VI, 114-6 IV-VI, li5S-6
Anal fin count lig li7 1i6-7 1i6-8
Pectoral fin count ele Til2 Til 1-13 Vil 1-12
Gill rakers in lower

gill arch 8 7 7.1(7-8) 7.1(7-8)
Vertebral count 32 32 31.6(31-32) 31.2(30-32) .
OD1, in relation to TV

(scales ) B-3:5 [Be 35) F373 2024.5) 9 GB)
OD2, in relation to OA

(scales ) Bl Bl B 1(0.5-1.5) B 1.5(0.5-1.5)
OD1, in relation to T. Pec. ipa eee ghia

(scales) eae B 1.5 B2 B.2:4( 15-320)". -B’2¢1.5-3:0) "=
OV, in relation to T. Pec. directly

(scales) Fl below F 0.6(0-1.5) F 1.1(0-3)
Position of anus to TV

(scales) B 0.5 B 1 B 1(0-2 B) - B0.5(FO0.5-B0.5)

Remarks—C. dalhousiensis is sexually dimorphic, a condition which is exceptional
amongst the Australian hardyheads, except for some sexual differences during
the breeding season (Llewellyn, 1971). It is, however, quite normal for blue eyes
(Pseudomugilinae) and rainbow fish (Melanotaeniidae) their close relatives. Fish
longer than 40 mm SL are easily sexed (see Fig. 1). externally. On the average (in a
sample of 64) the adult male is shorter than the adult female. The dorsal surface
in males tends to be almost horizontal from the snout to the origin of the first
dorsal; the abdomen on the other hand is gently arched from the isthmus to the
tips of ventrals. In the female, the belly is flat, but the head slopes sharply towards
the snout with the interorbital space being flat to concave, depending on maturity.
No other differences can be observed between males and females externally.
Specimens less than 40 mm SL are difficult to sex although ripe gonads were

Aust. Zool. 18(2), 1974 91

W. IVANTSOFF and C. J. M. GLOVER

found in a 34 mm male and a 27 mm female. Immature individuals tend to
resemble adult males, but the abdomen is flat. Gonads in both sexes lie below
and to one side of the intestine. The ovary, a long cylindrical sac is completely
enclosed by black mesovarium. It lies below and to the right side of the
intestine with its duct extending past the rectum to open just posteriorly to the
anus. The testis is a yellow-white mass lying below and to the left side of the
intestine with its duct extending past the anus to open posteriorly as in the female.
Gonads were checked histologically to confirm the relationship between sex and
external morphology. In a 34 mm male collected in November, sperm was
found in great abundance. In a thin section of an ovary from a 37 mm female
collected in August, no ripe oocytes were visible, although the more immature
stages were clearly evident.

Superficial examination of adult females suggests a close affinity with C.
lacustris Trewavas. Apart from significant colour differences, the size range,
vertebral, midlateral scale and gill raker counts are very different. Coloration
and spot pattern suggests a closer similarity to C. stercusmuscarum (Gunther),
but none of the 64 specimens of C. dalhousiensis examined for colour and spot
pattern had more than one row of spots above the midlateral band as seen in C.
stercusmuscarum. However, immature specimens of the two species are easily
confused because of their similar pigmentation. C. dalhousiensis is deep bodied,
C. stercusmuscarum is slender, the midlateral scales, vertebral and gill raker counts
are also different. A comparison with other allied species also shows meristic and
morphometric differences (Table 2).

TABLE 2

MERISTIC AND MORPHOMETRIC DIFFERENCES (EXPRESSED AS MEANS AND
RANGES) OF FOUR CLOSELY ALLIED SPECIES OF CRATEROCEPHALUS

C. lacustris C. stercusmuscarum — C. marjoriae C. dalhousiensis

Greatest body depth

in SL 4.6(3.5-6.0) 4.9(3.5-5.6) 4.1(3.7-4.5) 4.1(3.7-4.7)
Midlateral scales 34.3(32-38) 32.6(30-34) 28.5(24-30) 29.8(29-31)
Transverse scales 7.3(6.0-8.5) 7 6(5.5-7.0) 7.1(6.5-7.5)
Vertebral count 36.5(35-39) 36.8(35-39) 31.4(30-32) 31.4(30-32)
Gill rakers in lower

gill arch 10.7(9-12) 9.1(8-10) 10.1(9-11) 7.1(7-8)

On the basis of body colour, pattern and body shape, C. dalhousiensis appears
to be closely related to C. lacustris and C. stercusmuscarum. Counts and measure-
ments also suggest a close affinity with C. marjoriae Whitley. A comparative
study of the premaxillary bone of five species of Craterocephalus confirms the
relationship with C. lacustris and suggests affinity with C. eyresii (Steindachner )
(Fig. 2). Counts and measurements do not confirm close relationship with
C. eyresii.

92 Aust. Zool. 18(2), 1974

A NEW FRESHWATER TELEOST

Diagnostic features—C. dalhousiensis can be distinguished from its congeners by
the following characters: it is a deep bodied fish which is dark brown above
and with two to three rows of brown spots below the midlateral line, 7 to 8
gill rakers in the first lower arch, 29 to 30 midlateral scales, 62 to 72 transverse
scales; females have a head which slopes sharply towards the snout whilst the
abdomen tends to be flat. Mature males have a sloping abdomen and a relatively
horizontal dorsal surface.

Range—At present, this species is only known from Dalhousie Springs, South
Australia which is about 120 km north of Oodnadatta. It may have a wider range
as has proved to be the case with C. eyresii which has been recorded (unpublished

e

Fig. 2. Comparison of the premaxillary bones of 5 species of Craterocephalus.

(a) C. lacustris

(b) C. dalhousiensis
(c) C. marjoriae

(d) C. eyresii

(e) C. stercusmuscarum

Aust. Zool. 18(2), 1974 93

W. IVANTSOFF and C. J. M. GLOVER

data) from the Namoi, Peel and ee Rivers and the inland lakes of Victoria
and South Australia. ee : | ek ee ee

ieee
ate

The name given to the species is ced dice he name of the type locality,
a major area of mound spring development, embracing about thirty-three artesian
springs of varving size: and activity, scattered over an area- oe Aen Je kor:

(B) a NOTES

MATERIALS AND METHODS

The fish were collected with wire mesh traps (26 gauge wire, 2 mm
mesh) 31 x 15 x 15 cms. Each end of the cage had an aperture with an internal
collar 3.5 cms in diameter. Traps were set both during the day and _ night.
The size and the composition of catches was noted and recorded. Each species
was identified and examined for gut content.

Water temperatures were recorded at five stations over a distance of ap-
proximately 1 km along the course of Dalhousie Main Spring stream. pH and
salinity were also taken at the time of trapping.

Observations of movements of C. dalhousiensis were made at Dalhousie,
Springs (by C.J.M.G.) on two occasions. The observations were followed up by a
series of simple field experiments to determine upper and lower thermal tolerances

of the hardyheads.

(a) In one experiment the fish trapped in a cage. were transferred to a point
5 metres upstream where the temperature recorded was higher. The experi-
ment was repeated later in the day with a metal baffle inserted over the
upstream end of the cage to reduce the force of the water current.

(b) Captive hardyheads and catfish (Neosilurus sp.) from Dalhousie Main Spring
were held in non-heated, non-aerated tanks in which the temperature was
allowed to drop to well below 20°C. |

To determine whether thermal acclimation takes place, freshly retrieved
hardyheads were subjected in batches of four to rising water temperature by
placing the holding tank (one litre polythene beaker) in a large water bath
(an enamel bucket) containing water at 100°C. The fish were taken from
waters of various temperatures to determine whether fish inhabiting the warmer
sections of Dalhousie Main Spring were tolerant to higher temperatures . than
those from cooler regions. Constant agitation of water in the holding tank by
stirring with a hand-held glass bulb thermometer, produced rapid and uniform
transfer of heat, enabling the upper thermal limit of fish to be reached in
about 10 minutes. A total of sixteen specimens from each of the two regions
of the Spring stream were tested for thermal acclimation. :

94 Aust. Zool. 18(2), 1974

A NEW FRESHWATER’ TELEOST

“RESULTS AND. DISCUSSION ©

Trapping—Far greater numbers of hardyheads (C. dalhousiensis) are trapped in
daylight than during equivalent periods at night. In the case of the catfish (Neosilurus
sp.) the reverse applies. A. comparison of day and night catches (Table 3),
made with a wire trap set along the Dalhousie Main Spring stream for five
hour periods, confirms these catch patterns which presumably reflect activity
patterns.

Associated fish—In the Dalhousie Main Spring (Table 3), an extremely large
population of hardyheads (C. dalhousiensis) is found in association with an equally
abundant population of catfish (Neosilurus sp.) together with very much smaller
populations of two other small species, the desert goby (Chlamydogobius eremius )
and the spotted gudgeon (Mogurnda mogurnda). :

The catfish (Neosilurus sp.) appears ‘to be an undescribed form.

_ Similar associations were found in all but one of the other inhabited springs

inspected at Dalhousie; in one large spring approximately 0.8 km west of the
main spring the species collected were C. dalhousiensis, C. fluviatilis McCulloch
and Madigania unicolor.

Diet—Examination of gut contents in all specimens form a series of C. dalhousiensis
yielded a large number of small unidentified gastropods and remnants of plant tissue.
Parasites—No external or internal parasites were noted on or in any of the
C. dalhousiensis specimens examined.

TABLE 3

- NUMBERS OF FISH TRAPPED DURING FIVE HOUR SETTINGS
Temperature and pH data recorded on morning of 19/11/69.

(a) day trapping—1100-1600 hours, 18/11/69.
(b) night trapping—2100-0200 hours, 18-19/11/69.

Surface = Water

Station C. dalhousiensis Neosilurus sp. M.mogurnda C. eremius ; water pH

(a) (Bee). (bea Can. Cb) (aj), (6) temp.
a 309 0 0 —~0 0 0 0 0 43.0°C 2
b 283 0 0 136 3 3 pe 0 35.6°C Te
c 15 (Onde OA 126 1 0 0 0 35:0°C° TZ
d 62 0 1 24 1 0 0 0 23 4G 7.2
e 37 0 0 0 8 2) Once. 0 2162 7.2.
8 2 0

Total 706 Opeets 286 13

Aust. Zool. 18(2), 1974 95

A NEW FRESHWATER TELEOST

Water temperature, pH and salinity—Surface temperatures recorded at five
stations over a distance of approximately 1 km along the course of Dalhousie
Main Spring stream during summer in 1967 varied from 43°C in a narrow channel
feeding into the main body of the spring at the west end, down to 21.6°C in the
shallows of the swamp at the east end into which the spring waters spread (Fig. 3).

low vegetated mound

swamp
shallows

Fig. 3. Diagramatic representation of Dalhousie Spring and stream with collecting
stations indicated.

Water pH (Table 3), measured at each of the above stations was con-
sistent at 7.2, a fairly typical value for swiftly flowing artesian waters of the
Lake Eyre drainage region. The pH of outflowing waters is usually about 7.0;
this often rises along the stream path, minimally in swiftly flowing waters, more
steeply in slowly flowing shallow streams and terminal waters, especially in the
presence of large quantities of algae or other aquatic vegetation when pH may
rise to very high levels.

Water sampled about mid-way along the Dalhousie Main Spring stream
and subsequently analysed (Table 4), indicated relatively low salinity (as total
dissolved solids) compared with other artesian waters of the central Australian
region.

Thermal tolerance—(a) Upper thermal tolerance

Large numbers of C. dalhousiensis have been observed making brief solitary
excursions of up to one minute’s duration from the edge of the main body

96 Aust. Zool. 18(2), 1974

W. IVANTSOFF and C. J. M. GLOVER

TABLE 4

ANALYSIS* OF WATER SAMPLED FROM THE MAIN SPRING AT DALHOUSIE
SPRINGS, SOUTH AUSTRALIA, 6/6/68.

(*Analysis performed by the Australian Mineral Development Laboratories,
South Australia).

pao Hardness
Constituent m composition m
a oe salts a (as Ca COs) ie
Anions
Cl 370
SO, 150
HCO. 140 Ca HCO, 186 Total 245
hag face Ca SO, 41 Temporary 115
Cations
Na DAL Mg SO, 119 Permanent 130
K
Ca 58 Na SO, 38 Due to Ca 145
Mg 24
F Na Cl 610 Due to Mg 100
Total dissolved Total alkalinity
solids 994 as Ca CO; 115

of the Main Spring into the narrow channel of rapidly inflowing shallow water,
originating from the spring’s outlet higher up on an adjacent vegetated mound.

Specimens of C. dalhousiensis and Neosilurus sp. trapped near the entrance
to the channel at 30 cm depth, where the water temperature was 38.8°C, were put
back into the water about 5 metres upstream in depth of 10 cm and temperature
of 41.8°C. It was observed that all the hardyheads collapsed rapidly and died
within 16 minutes of the transfer. The catfish died 2 to 3 minutes later.

When the force of the current was reduced by the insertion of a metal
baffle over the upstream end of the trap, the hardyheads and the catfish survived
approximately 60 and 120 minutes respectively.

Clearly those fish inhabiting the surface waters in the vicinity of the channel’s
outlet into the spring’s large pool are living close to their upper thermal limit
and in fact the hardyheads intrude briefly into water which, if their stay is
prolonged, proves lethally warm.

Although the motivation for these incursions is not clear it is probable
that they are foraging upon blue-green algae that grows abundantly over the
bed of the channel, parallelling the findings of Brown (1971) who reported
that the desert pupfish (Cyprinodon spp.) in southern California feed on
blue-green algae in water of about 42°C which is close to their upper thermal
limit.

Aust. Zool. 18(2), 1974 97

A NEW FRESHWATER: TELEOST ..

(b) Lower thermal tolerance:

Fish held in tanks where temperature was allowed to fall below 20°C
rapidly became comatose. None survived overnight in this situation by which
time the water temperature had dropped to 8.7°C.

In one experiment where water temperature had fallen to 16°C, a comatose
catfish was revived by raising the water temperature above 20°C. Tic experiment
was not repeated with the hardyheads. ?

Thermal acclimation:

The results obtained for thermal acclimation are summarized in Table
The data collected suggest the occurrence of direct acclimation by C. dalhousiensis
to the thermal gradient along the Main Spring stream. Similar results were obtained
with the catfish from the same spring and with desert gobies (Chlamydogobius
eremius) elsewhere (C.J.M.G., unpublished data).

TABLE 5
SUMMARY OF RESULTS FOR THERMAL ACCLIMATION BY C. DALHOUSIENSIS

Surface water Standard Critical Critical
temperature at lengths SL thermal thermal
Station collecting range mean maxima maxima
station (16 specimens) range mean
a 38.8°C 30-52 mm 36 mm 41.8-44.6°C 43.6°C

d 3228 © 37-50 mm 43 mm — = 37.8-42.5°C 40.2°C

These results are similar to those of Brown and Feldmeth (1971) who
found that the upper thermal limit of desert pupfish (Cyprinodon spp.) freshly
collected from thermal springs correlated with the temperatures of their habitats.

ACKNOWLEDGEMENTS

We wish to thank Drs. D. A. Pollard and R. Pengilley and Mr. J. R. Bassett for
their constructive criticism of the manuscript.

REFERENCES

BROWN, J. H. (1971). The desert pupfish. Sci. Am. 225, 104-110.

BROWN, J. H. & C. R. FELDMETH (1971). Evolution in constant and fluctuating
environments; thermal tolerances of desert pupfish (Cyprinodon). Evolution 25,
390-398.

LLEWELLYN, L. C. (1971). Breeding studies on the freshwater forage fish of the
Murray Darling river system. Fisherman, St. Fish. N.S.W. 3(13), 1-12.

MUNRO, I. S. R. (1967). The Fishes of New Guinea. Dept. Ag., Stock, Fish., Port
Moresby, New Guinea, 650 pp., 84 pls.

98 Aust. Zool. 18(2), 1974

The | Fauna of Careel Bay with Comments on the ~
Ecology of Mangrove and Sea-Grass Communities

P. A. HUTCHINGS and H. F. RECHER

| Department. of Marine Invertebrates, and Department of Environmental Studies, ~
| Australian Museum, Sydney, New South Wales, 2000.

ABS LRAGE

Careel Bay, Pittwater, New South Wales can be divided into five zones, salt
marsh, mangroves, Zostera and Posidonia weed beds and sandy beach. The flora
and fauna of each of these zones are described with comments on the seasonal
abundance of animals in the weed beds and on the sandy beach. Finally the
inter-relationship of these zones to each other especially in relation to the detritus
food chain is discussed.

INTRODUCTION

It is widely accepted that salt marsh, mangrove and sea-grass communities
are important components of estuarine ecosystems (Lyford and Plinney, 1968;
Odum, 1961; Odum and de La Cruz, 1967. These communities act as nurseries for
many organisms and produce large quantities of organic matter which forms the base
of estuarine food chains. These communities have been little studied in Australia
(Clarke and Hannon, 1967, 1969, 1970, 1971; Collins, 1921; Goodrick, 1970;
Hamilton, 1919; Kratochvil, Hannon and Clarke, 1972; Macnae, 1966, 1967)
and even basic descriptions of fauna are lacking.

During 1972, the New South Wales Division of the Australian Littoral
Society surveyed the biology of Careel Bay in Pittwater on the Central Coast
of New South Wales (Hattersley, Hutchings and Recher, 1973). The survey
considerably expanded our knowledge of the fauna of estuarine habitats on the
Central Coast, but in view of the limited data available for the fauna of mangrove
and sea-grass communities in Australia, it was decided to extend the survey
for an additional year. The results of the work during 1972 and 1973 are
presented in this paper and although emphasis is on a description of the fauna, its
distribution within Careel Bay and seasonal changes, the role of salt marsh, mangrove
and sea-grass communities in the ecology of the estuary is also considered.

Careel Bay

Careel Bay (33° 37° S; 151° 20° E) is situated on the southeastern shore
of Pittwater. Pittwater is part of the Hawkesbury River-Broken Bay estuary
(Fig. 1): an extensive drowned river valley 32 km north of Sydney. No major

Aust. Zool. 18(2), 1974 99

P. A. HUTCHINGS and H. F. RECHER

river flows into Pittwater and it is unlikely that any significant amount of
freshwater reaches Pittwater from the Hawkesbury River. The volume of fresh
water coming down the Hawkesbury River is irregular and carries a heavy
silt load. At times of flood on the Hawkesbury we have not observed any intrusion
of river water into Pittwater or Careel Bay.

Development on the watershed of Pittwater is primarily residential with
substantial areas along the northern shores reserved as National Park. Careel
Bay is shallow, but well protected from prevailing winds. Surrounding lands
have been developed for residential purposes and water flowing into the Bay

Lise eo

j th
ASS
} Ly
C f
ad [ smeane
/) y
L
es
eo x
WN 7 s
t
a,
“1, BROKEN Bay
ote
oe
“Po PACIPi
+ OcBan
A 4
A
o s
+
v rs
o ae &
677
KURINGAI
WATIONAL
PaRk ae
WATER

H
i
|
|
i

e

Fig. 1. Map of Pittwater, Hawkesbury River and Brisbane Waters, showing the relationship
of Careel Bay to these areas.

100 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

comes primarily as runoff from the storm water drainage system. There is
some seepage from septic systems and runoff from the garbage tip.
Other than minor local effects, water pollution, however, is not a problem. The
Bay opens widely to Pittwater and the tidal range is 2 m. The entrance to
Careel Bay is marked by a sand bar which consists of sediments
transported from the ocean side of Pittwater. The bar prevents sand
from entering the Bay, but does not seem to impair the exchange of water
with Pittwater. Sediment deposition is occurring at the head of Careel Bay
and over the last century, 1.3m of sediment has been deposited (Hattersley,
Hutchings and Recher, 1973). The mangroves and sea grass flats at the head of
the Bay appear to trap and retain most of these sediments. It is likely that these
sediments have been derived from disturbances associated with the residential
development of the foreshores and surrounding areas and the rate of sedimentation
may decline when the area is fully developed.

TABLE 1
AREA OF MAJOR LITTORAL HABITATS

Zone Area (m?°) %
Sandy beach 149.0 15
Salt marsh 32.9 3
Mangroves 82.9 7
Zostera
Sea grass flats 427.5 38
Posidonia
Sea grass flats 410.0 37

The estuarine environment of Careel Bay (Fig 2) can be conveniently divided
into five zones, sandy beach, salt marsh, mangrove (Fig. 3), Zostera grass flats
and Posidonia grass flats. The extent of each zone is given in Table 1. The
zones and their approximate distributions are shown in Fig. 4.

METHODS

Between January 1972 and December 1973 a number of general collecting
trips were made to Careel Bay. These included night collecting. A representative
sample of all the animals collected during this survey has been deposited in the
Australian Museum, Sydney. In addition quantitative sampling was carried out
in February, June and November, 1973. In February only the upper zone of
Zostera was sampled, but in June and November the upper and lower zones
were sampled separately.

During each period of quantitative sampling ten samples were collected from
the Posidonia and Zostera weed beds and sandy beach. Using a spade 9000 cm? of

Aust. Zool. 18(2), 1974 101

P. A. HUTCHINGS and H. F. RECHER

Fig. 2. Aerial photograph of Careel Bay. Photo: Peter Whalan.

Fig. 3. The salt marsh mangrove junction. Numerous small bushes of Avicennia marina
in the foreground, amongst the salt marsh vegetation. Photo: Peter Whalan.

102 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

mud or sand and associated plants (if any) was collected and passed through a 2.0
mm sieve. In taking the samples, only the surface layer of the substrate to a depth of
10 to 20 cm was collected. Fast swimming and deep burrowing animals were
probably missed or underestimated by this method. In the Posidonia zone, a
different method was adopted for the June and November samples. Divers with
SCUBA pulled out plants with their root systems and adhering mud until 9000
cm® of material had been collected. Again 10 samples were collected, but these are
not comparable with the samples taken in January or from the other zones. We
found collecting in the Poszdonia difficult and this zone was not well sampled.

In addition to the volumetric samples, the population density and _ size
distribution of the Sydney cockle Avxadara trapezia was measured using a 0.25m?
quadrat in the lower Zostera. A 0.25m? quadrat was used also to estimate the
biomass of Zostera and Posidonia. The quadrat was placed randomly ten times in
each zone and all plant material was removed from within the quadrat. The plants
were washed, excess moisture and epiphytes removed and the plants weighed.
The samples were then dried at a constant humidity and temperature (21°C)
ad re-weighed until constant weight.

\

\ _Upper
spelen ih \ Zostera

=>

Posidonia ii I

t

t
Beds !

Sandy Beach

GY) Mangroves 800 a
HUI} Sart Maren — = FEET

rig. 4. Careel Bay showing the extent of the 5 zones. The reclaimed areas of the
Municipal Tip and Playing Fields are also shown.

Aust. Zool. 18(2), 1974 103

P. A. HUTCHINGS and H. F. RECHER

FAUNA

The fauna of Careel Bay will be presented in two ways. First we des-
cribe the major groups of animals which occur in Careel Bay. Second we
relate the distribution and abundance of the fauna to each of the five major
habitats (zones) in Careel Bay.

Marine Invertebrates

A list of the marine invertebrates collected during 1972 and 1973 at Careel
Bay is presented in Table 2. This table also shows the distribution of species
by zones and an indication of species abundance on a scale of loge throughout the
year. Absolute values of abundance are not given as we have insufficient quantitative
data for most animals to justify giving precise figures. Also some of the species
are very patchy in their distribution. The Zostera has been considered as a single
zone in this Table. The faunal lists for the five zones present in Careel Bay are not
complete, as even after 2 years of regular collecting, new records for the Bay
continue to be obtained. These probably represent seasonal visitors to the Bay
or species which are sparsely represented in the Bay. The species list for the
mangrove zone is incomplete in that the infauna of the mud was inadequately sam-
pled. For practical reasons it was only possible to sample areas adjacent to the creek.
Other invertebrates occur in Careel Bay such as encrusting organisms on pilings,
moorings or rocky habitats, and these were not sampled during this survey.

TABLE 2

DISTRIBUTION AND ABUNDANCE (WHERE AVAILABLE) ON A SCALE LOG.
OF MARINE INVERTEBRATES IN CAREEL BAY

Salt Sandy
marsh mangroves Zostera Posidonia beach
Coelenterates Edwardsia sp. 1 1
Platyhelminthes Turbellarian 1
Nemerteans Heteronemertean 2 Dp} 1
Polychaetes
F. Polynoidae Paralepidonotus cf.
ampuliferons 1 i
F. Phyllodocidae Phyllodoce
novaehollandiae 1
Phyllodoce sp. 1
F. Syllidae Syllis sp. 1 . 1 1
Syllis sp. 2 1
F. Nereidae Australonereis ehlersi 1 4
Ceratonereis
erythraeensis I 1
Ceratonereis mirabilis 1
Neanthes yaalii 1 1
Nereis (Hediste) diversicolor 1 3 1

104 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

TABLE 2 (continued)

Salt Sandy
marsh mangroves Zostera Posidonia beach
F. Nephtyidae Nephtys australiensis 1 4 1 1
F. Glyceridae Glycera americana 1 2
F. Eunicidae Marphysa sanguinea 1 1
F. Lumbrineridae Lumbrineris latreilli 1 1
F. Orbiniidae Haploscoloplos n. sp. 1 1 1 3
Naineris_ sp. 1
Orbinia sp. 1 1
Phyllo sp. 1 1
Scoloplos sp.
F. Spionidae Boccardia sp. 1
_ Dispio sp. 1
Malacoceros sp. 1
Polydora sp. 2 2 1 1
Prionospio sp. 1 2
F. Magelonidae Magelona sp. 1 1
F. Chaetopteridae Chaetopterus sp. 1 1 1
F. Cirratulidae Cirratulus sp. 1
F. Flabelligeridae Pherusa sp. 1
F. Opheliidac Armandia lanceolata 1 1
F. Capitellidae Barantolla lepte 3 1 1
Capitella capitata 1 1 1
Heteromastus sp. 1
Mediomastus sp. 1 1
Notomastus hemipodus 5 Z
F. Maldanidae —incomplete specimen 1
F. Arenicolidae Arenicola bombayensis 1
F. Oweniidae Owenia fusiformis 3 1 1
F. Ampharetidae Amphicteis sp. 1
Lysippides sp. 1
Samytha sp. 1
F. Terebellidae Lysilla pacifica 2 1
Thelepus setosus 1
Rhinothelepus lobatus 2 1
Pista sp. 1
F. Serpulidae Spirorbis sp. 3 3
Hydroides sp. 1
Phoronids Phoronis sp. 1
Crustaceans
Sub-Class. Cirripedia
F. Balanidae Balanus amphitrite 1
Order Isopoda
F. Anthuridae 1
F. Sphraeromidae Cymodoce coronata 1
F. Ligidae Ligia australiensis x 1
Order Amphipoda
F. Ampithoidae Ampithoe sp. x 2 2
F. Gammaridae Melita sp. 2
F. Liljeborgidae Liljeborgia sp. 1
F. Haustoriidae Urohaustorius sp. 1 1

Aust. Zool. 18(2), 1974 105

Order Decapoda
F.

Penaeidae

Palaemonidae

F. Alpheidae

bono ton

EF.

poMeoe Meola oles!

. Hippolytidae
. Laomediidae
. Callianassinae
. Paguridae
Hymenosomatidae
. Portunidae

. Xanthidae
. Grapsidae

. Ocypodidae

Mictyridae

Molluscs

E:
FF:

a0

ole oBe fies ies aiille sine iss ie silos

Arcidae
Mytilidae

. Laternulidae
. Lodabiidae

Leptonidae

. Ostreidae

Veneridae

Macturidae
Garidae

. Tellinidae
. Teredinidae
. Trochidae

. Liotidae

106

P. A. HUTCHINGS and H. F. RECHER

TABLE 2 (continued)

Salt
marsh

Penaeus plebejus

Penaeus esculentus
Penaeus sp.
Macrobrachium intermedius
Alpheus euphrosyne
Alpheus sp. B

Alpheus sp. C

Hippolyte tenuirostris
Laomedia healyi
Callianassa sp.

Halicarcinus ovatus
Portunus pelagicus
Thalamita sima

Thalamita crenata

Scylla serrata
Heteropanope serratifrons
Sesarma erythrodactyla x
Paragrapsus laevis Xe
Heloecius cordiformis x
Macrophthalmus setosus
Macrophthalmus crassipes
Macrophthalmus cf punctulatus
Macrophthalmus sp. 1
Macrophthalmus sp. 2
Species A

Species B

Australoplax tridentata
Mictyris longicarpus

Anadara trapezia
Modiolus pulex

Mytilus edulis
Xenostrobus securis
Laternula tasmanica
Cavatidens omissa
Mysella sp.
Saccostrea cucullata
Eumarcia fumigata

Tapes watlingi
Notospisula parva producta
Florisarka onuphria
Macoma deltoidalis
Teredo sp.

Austrocochlea obtusa x
Calliostoma australe
Salsipotens meyeri

Liotia sp.

mangroves

~

mm mM OM

~ eh x

Sandy
Zostera Posidonia beach
2 1
i 1
1
2 D
1 1
x
1
1 1 pe
mx
1
1 1
1 1
1 1
2
2
x 1
x 1
x x
2
1 1
x 1
Xi 3
3 x
x
x
1
1
Ds 1 1
1 1
1 1
1 1
2 1 Zz
1
2 2 1
1
1
1

Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

TABLE 2 (continued)

i

Salt Sandy
marsh mangroves Zostera Posidonia beach
wee Eg eh Pe MRSA RS a SOM a Dae Pa gn ae SUA a SAO See Rae Nee ee
F. Acmaeidae Chiazacmea flammea x
F. Neritidae Nerita atrementosa x
F. Littorinidae Littorina scabra x x
Bembicium auratium x x 1
F. Dialiidae Diala sp. 1
F. Tateidae Tatea_ rufilabris x
F. Cerithidae Cacozeliana lacertina 1 2 1
F. Potamiidae Pyrazus ebenensis 1
Velacumantus australis x 1
F. Epitoniidae Epitonium sp. 1
F. Naticidae Conuber sordidum 1 1
Conuber sp. 1
F. Nassariidae Nassarius burchardi x 1 a 1 f,
Nassarius jonasi aK
F. Muricidae Bedeva_ hanleyi 1S LB
F. Pythidae Melosidula zonata x
Ophicardelus sulcatus x x
Ophicardelus ornatus x
Ophicardelus quoyi x x
F. Amphibolidae Salinator solida x
Salinator sp. thea bah
F. Onchidiidae Onchidium damelli x x
F. Akeridae Haminoea_ wallissi 2
F. Aplysidae Aplysia dactylomela x %
Aplysia juliana me x
F. Elystidae Elysia australis x *
F. Dorididae 1
F. Sepiolidae Euprymna_ stenodactyla ys Xx
Echinoderms Astropecten polycanthus x
Ascidian Botrylloides nigrum x

The sea anemone Edwardsia sp. and an unidentified turbellarian were
collected in the upper Zostera zone. A single species of heteronemertean was
collected from the sandy beach, Zostera and Posidonia zones. This species is being
described by Gibson (Liverpool Poly., U.K.).

Forty-six species of polychaetes representing 22 families were collected
including 21 identified to species level. Many of the polychaetes are new records
for New South Wales and some undescribed species are present. These will be
described in a paper by Hutchings (in prep.). No species of polychaete was found
throughout the Bay although Haploscoloplos n. sp. was found in all zones except
the salt marsh (Table 2). Twenty-three species occurred in two or three zones
while 19 species were confined to a single zone. The Zostera weed beds had
the richest fauna of polychaetes many of which are tubiculous including the
abundant nereid Awstralonereis ehlersi. The salt marsh, mangroves and sandy

Aust. Zool. 18(2), 1974 107

P. A. HUTCHINGS and H. F. RECHER

beach had few species present although some in the sandy beach (e.g. Australonereis
ehlersi and Haploscoloplos n. sp.) were abundant. A phoronid was found living
in the empty tubes of the polychaete Owenia fusiformis, in November, 1973.

The majority of the polychaetes found in Careel Bay are detritus feeders.
Some like Arenicola and all the capitellids actively ingest the substrate and
digest the algae and bacteria adhering to the surface of the mud _ particles.
Newell (1965) and Longbottom (1968) have shown how the resulting faecal
material is enriched with nitrogen compounds which are then utilized by the
bacteria and algae in the sediment. Subsequently this sediment is re-ingested and
the cycle is repeated. The terebellids are surface deposit feeders, (Dales, 1955).
Other species of polychaetes present are filter feeders, feeding on suspended
detritus material and phytoplankton, these include Chaetopterus, and the serpulids.
It is likely that Nephtys australiensis and many of the nereids are omnivorous
scavengers although some experimental work has indicated that the polynoid
Paralepidonotus cf. ampuliferons, the glycerid Glycera americana and the syllids
are probably active carnivores feeding on small worms and crustaceans (pers.
obser. ).

Thirty-eight species of crustaceans were recorded from Careel Bay, most of
these (31) occurred in the Zostera and Posidonia zones with only eight species
present on the sandy beach. Three species occurred in the salt marsh and eleven
species among the mangroves. Amphipods, isopods and other smaller crustacea
were poorly sampled.

Many of the larger crustaceans were extremely abundant and show
precise patterns of zonation with reference to high and low water marks. On
the sandy beach a new species of Callianassa has been found and this is being
described by Poore (Fisheries & Wildlife, Victoria, Australia). Three species
of the snapping prawns Alpheus sp. were present in the weed beds and these are
being described by Banner (Hawaii). Two crabs species, A & B, as yet unidentified,
were especially abundant in the lower Zostera, but also occurred in the upper
Zostera zone. Species A (Table 2) may be Camptandrium paludicola a species
recorded by Snelling (1959) from the Brisbane River, Queensland. Three species
of penaeid and carid prawns, Penaeus plebejus and P. esculentus and Macro-
brachium intermedius were very abundant in the weed beds.

Many of the crabs in the mangroves feed on the algae growing on the
bases of the mangrove trees and pneumatophores. Large numbers of crabs were
observed feeding at night during low tide. For example Paragrapsus laevis was
present in numbers ranging from 24-30 individuals /m”, Sesarma erythrodactyla
vatied from 13-88/m?, Heloecius cordiformis 3/m” and Australoplax tridentata
9/m?. Similar measurements made during the day at low tide were in the order
of 10-15 crabs/m?. The amphipods, Callianassa and the alpheids are predominantly
detritus feeders; and the soldier crab Maictyris longicarpus ingests the sand
removing organic material.

108 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

Forty-six species of molluscs were collected during the survey, Gastropods
were the most numerous (31 species) followed by pelecypods with 14 species.
We did not collect any chitons (Placophora), but they occur at Careel Bay along
the rocky foreshores. Two species of cephalopods occur, Octopus cyaneus and Sepia
sp., (Dakin, Bennett and Pope, 1952), but were not collected in this survey.
The opisthobranchs, Aplysia dactylomela and Aplysia juliana, were particularly
abundant during the spring and early summer on the sea grass flats. In contrast
to the polychaetes, many of the molluscs were confined to a single zone. No
species occurs in more than three of the five zones.

The molluscs can be divided into two main feeding categories, filter feeders and
grazers. The bivalves which would represent the greatest biomass amongst the
molluscs are all filter feeders; these are present in all zones except the salt marsh.
In the weed beds, common species are Anadara trapezia, Macoma deltoidalis
and in the mangroves Saccostrea cucullata. The grazers are represented
by the gastropods, which feed either on the film of epibiota on the surface of
the sea grasses or on the surface of the mud in the salt marsh or mangroves.

The only other marine invertebrates collected during the survey were the
burrowing starfish Astropecten polycanthus in the lower Zostera and Posidonia,
and the compound ascidian Botrylloides nigrum. This ascidian was found encrusting
the leaves of the Posidonia in the November samples.

ABLE: 3
SPIDER FAUNA OF CAREEL BAY

Salt
Order. Araneae Marsh Mangroves
F. Clubionidae Clubiona robusta x
Miturga sp. x
F. Pisauridae Dolomedes. facetus x
F. Lycosidae Anoteropis longipes Xx Xx
Lycosa furcillata x
Lycosa sp. xX x
F. Hersiliidae Tama novaehollandiae x
F. Thomisidae Diaea sp. x
F. Therididae _Argyrodes antipodianus Xi
F. Tetragnathidae Tetragnatha bituberculata
Tetragnatha sp. X
F. Argiopidae Aranea_ insuta x Xx
Aranea sp. x
Argiope aethera x
Nephila ornata x
F. Linyphiidae Erigoninae sp. x x
Laetesia sp. x
Linyphiinae sp. x

Many of the spiders identified only to genus are juveniles.

Aust. Zool. 18(2), 1974 109

P. A, HUTCHINGS and H. F. RECHER

Non Marine Invertebrates

A list of the spiders collected during the survey is presented in Table 3.
Many species were collected only once and information on the seasonal occurrence
of species is not available. Many of the spiders were identified to genus only
as they were juveniles. In Appendix JI, a list of insects which have been recorded
from Careel Bay is given, this was compiled by D. McAlpine and G. Holloway of the
Australian Museum from their Departmental records. Two interesting flies of
the tropical genera Merodonta and Pemphigonotus have been found in the
Bay and constitute new southern records for the genera. Chelonus unimaculatus
and C. australiensis have been recorded for the first time since their original
description in 1905. The larvae of the mangrove plume moth Cenoloba obliteratis
which are restricted to the fruit and young shoots of Avicennia marina were
present. Also present is the mangrove fruit fly Euphranta sp. which is restricted
to mangrove habitats as its larvae live in the fruits of A. marina.

TABLE 4

WADING AND SEA BIRDS OF CAREEL BAY
(species which occur regularly)

White-faced Heron Ardea novaehollandiae
Mangrove Heron Butorides striatus

White Egret Egretta alba

White Ibis Threskiornis molucca
Straw-necked Ibis Threskiornis spinocollis
Black Billed Spoonbill Platalea regia

Yellow Billed Spoonbill Platalea flavipes

Little Pied Cormorant Phalacrocorax melanoleucos
Australian Pelican Pelecanus conspicillatus
Black Cormorant Phalacrocorax carbo

Black Duck Anas superciliosa

Dusky Moorhen Gallinula tenebrosa

Grey Teal Anas_ gibberifrons

Spur Winged Plover Vanellus miles novaehollandiae
Bar-tailed Godwit Limosa lapponica

Eastern Curlew Numenius madagascariensis
Whimbrell Numenius phaeopus
Red-necked Stint Calidris ruficollis

Grey tailed Tatler Tringa brevipes

Southern Stone Curlew Burhinus magnirostris
Sacred Kingfisher Halcyon sancta

Azure Kingfisher Alcyone azurea

White Breasted Sea Eagle Haliaetus leucogaster
Whistling Kite Haliastur sphenurus
Silver Gull Larus novaehollandiae
Crested Tern Sterna bergii

White Fronted Tern Sterna striata

110 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL. BAY

Marine Vertebrates

Two groups of marine vertebrates need to be considered; fish and birds.
Both groups are conspicuous and important parts of the Bay and the Hawkesbury
River-Broken Bay estuary. Indeed, in the context of immediate social concern and
economic value, both groups are of special interest. It is difficult to sample
either group quantitatively and our observations are necessarily qualitative. Birds
and fish are highly mobile animals and changes in numbers and species composition
occur with each tide and season.

SANDY SALT
BEACH MARSH MANGROVE| ZOSTERA|POSIDONIA

Australian Pelican

so en Aenean anen amen eee al

Black Cormorant —
Sr mm ne

a

Little Pied Cormorant
White-faced Heron
Mangrove Heron

White Egret

Sel rte. cone ——- YI _

Black—billed Spoonbill se WHY
Yellow-billed Spoonbill ot reer YEE
Eastern Curlew a HTL,
Bar-tailed Godwit rt HY

Southern Stone Curlew

ert Fal eS: eee

Fig. 5. Schematic diagram showing the abundance and feeding zones of the common
Careel Bay Birds.

Uk ane 1-10
aa 10-20 nos. of birds of species always present in
the Bay.
1-10

10-20 nos. of birds of species often present in
Y = zs ,
VDL the Bay

Aust. Zool. 18(2), 1974 444

P. A. HUTCHINGS and H. F. RECHER

a. Fish

Fish were sampled using a 8 m seine with a fine 6 mm mesh. Thus,
small fish and juveniles rather than large fish were taken. The grass flats of
Careel Bay supported large numbers of young fish of commercially important
species. For example, numerous juveniles of trevally, blackfish, bream, tarwhine,
whiting and leather jackets were taken. These fish depend on the shallow waters
of the estuary for their early growth and survival. Other species are resident
on the grass flats and among the mangroves. Some, such as mullet, form important
parts of the estuarine food chain and channel the organic production of the sea
grasses and mangroves into larger and economically important species (Odum
and Heald, 1972). A list of fishes collected in Careel Bay and elsewhere in
the Hawkesbury system are presented in Appendix II; all the species listed are
expected to occur in Careel Bay.

b. Birds

The estuarine food chains which begin with the sea grasses and mangroves
often terminate with birds. At Careel Bay, wading and diving birds are a
conspicuous part of the fauna (Table 4). Wading birds forage on the sea grass
flats and among the mangroves at low tide while others such as terns, cormorants
and pelicans feed when the flats are covered by water. The Zostera zone is the
most important feeding area for the birds at Careel Bay and relatively few _
forage in the mangroves or on the salt marsh. However, the mangroves and
salt marsh are important refuges at high tide and the mangroves provide nesting
sites for herons. The salt marsh at Careel Bay is one of two places on the Central
Coast where the Southern Stone Curlew Burhinus magnirostris nests.

In Fig. 5 we have shown the most common birds recorded at Careel Bay
during 1972 and 1973 and the habitats in which they fed. Birds are highly
mobile animals and their numbers tend to fluctuate with tide, weather and
seasonal conditions. Probably most of the birds which forage on the tidal
areas of Careel Bay range widely throughout the Hawkesbury River-Broken Bay
estuary. In Fig. 5, we have, therefore, given relative numbers based on consistency
of occurrence and absolute numbers.

HABITATS
Sandy Beach

The beach is made up of coarse clean sand. Some sediment and freshwater
is carried onto the beach by a storm water drainage channel which empties
directly on the beach. The beach gradually merges into the Zostera. On the
land side of the beach there is limited development of Juncus maritimus and
Salicornia quinqueflora. Residential development adjoining the beach has limited
the area occupied by these plants.

112 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

Polychaetes were the most abundant and diverse group of invertebrates
found in this zone. The most abundant species were Australonereis ehlersi and
Haploscoloplos n. sp., both of which bred successfully during the year. This was
indicated by their increase in numbers per sample due to the settlement of
juveniles (Table 6). Haploscoloplos n. sp. bred between June and November
and juveniles were abundant in the November sample. There was no evidence
of mass mortality of the adults occurring after breeding. Australonereis ehlersi
bred earlier in the year; in the June sample, numerous newly settled individuals
were present, suggesting breeding had begun in April or May. Breeding
was still occurring in June as many of the females were gravid. Juveniles were
present in November samples. It is not known whether breeding occurred continually
throughout this period or if there were distinct outbursts of spawning, as found
in many nereids (Clark, 1965; Schroeder, 1968). Hartman (1954) suggests
A. ehlersi may be viviparous no evidence of this was found. The numbers
present in November are greater than those found in the preceding February
suggesting that either 1972 was a less successful breeding season than 1973
or that many of the spawned worms will die during the subsequent summer
months. Most species of nereids which have been studied breed once and then
die (Clark, 1961; Dales, 1950).

The soldier crab Mictyris longicarpus was the most abundant crustacean
in the sand, and newly settled individuals were numerous in June and November
samples. The juvenile stages were more easily sampled by our methods than the
adults and appear more abundant in our samples. Nevertheless, Careel Bay has a
large population of adult soldier crabs. All other crustaceans were sparsely
represented in the samples (Table 6).

Notospisula parva producta was the most abundant mollusc present in this
zone and numbers increased in June due to the settlement of juveniles between
February and June. Many of the other molluscs were represented only by one
or two individuals e.g. Eumarcia fumigata, Cacozeliana lacertina and Macoma
deltoidalis. A sirgle juvenile of Salinator sp. was found in June.

Salt Marsh

The salt marsh occupies the area between the mangroves and fully terrestrial
habitats. This zone has been affected by land fill operations and was probably
bordered originally by paperbarks Melaleuca sp. and she-oak Casuarina glauca. The
terrestrial margin of the salt marsh is dominated by the sedges, Cyperus polystachyos
and Juncus maritimus var. australiensis, among which are scattered Casuarina glauca.
Between the sedge zone and the mangroves is a region with bare mud and a
patchy vegetation of Salicornia quinqueflora and Triglochia striata. Towards the
mangroves there is a scattering of stunted mangroves Avicennia marina (Fig. 3).
The patchy distribution of Salicornia, Triglochia and Avicennia in this zone
reflects different levels of soil moisture and soil salinity (Clarke and Hannon,

Aust. Zool. 18(2), 1974 TRS)

P. A. HUTCHINGS and H. F. RECHER

SS

Fig. 6. Inside the mangroves, looking towards the weed beds.
Photo: Dan Lunney.

Uy
Woy);
Hi

Fig. 7. The upper Zostera weed beds. In the foreground are numerous seedlings of
Avicennia marina. Photo: Dan Lunney.

114 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

1967, 1969, 1970, 1971). The mangroves for instance, invariably occupy depressions
in the substrate surface.

The marine invertebrate fauna of the salt marsh was limited but the species
present were usually represented by large numbers of individuals. The gastropods
Salinator solida (1080/m*), Tatea rufilabris (74/m?), Ophicardelus sulcatus
(264/m?) and O. guoyi (217/m’) were the most abundant molluscs present.
These animals were found on the surface of the mud, and were closely associated
with the vegetation. The crabs Sesarma erythrodactyla, Paragrapsus laevis and
Heloecius cordiformis were also common, especially in wetter areas.

Polychaetes appear to be entirely restricted to the mangrove margin of the
salt marsh where residual pools of water remain. Three species were found,
Ceratonereis erythraeensis, Boccardia sp. and Capitella capitata. All these species
were represented by one or two individuals.

Mangroves

The mangrove zone is dominated by the grey mangrove Avicennia marina
var. australasica which is the common mangrove on the Central Coast (Fig. 6).
Adjoining the salt marsh there is a small number of the river mangrove Aegiceras
corniculatum, a species which is more abundant in brackish waters. Despite in-
trusions by the Council garbage tip, mangroves occupy a significant area of Careel
Bay (Table 1) and the stand appears to be colonizing the upper part of the Zostera
zone (Fig. 7). The bases of the mangrove trees and pneumatophores are covered
with a tufted red algae. The species has not been determined.

CANOPY (7200cm )

DOOR & D

MIDDLE STOREY (50-200cm)

FOLIAGE COVER

GROUND STOREY (« 50cm)

Fig 8. Schematic diagram of the Mangrove community in Careel Bay.

QhNON ED

Aust. Zool. 18(2), 1974 115

P. A. HUTCHINGS and H. F. RECHER

The mangrove zone is inundated on each high tide and the size of
an individual tree is closely related to its position in the inter-tidal zone. The best
growth of mangroves at Careel Bay occurs immediately adjacent to the Zostera
flats; that is, just about mid tide level. A transect through the mangroves (repre-
sented schematically in Fig. 8) shows that tree height declines behind the first
line of trees, with trees being noticeably smaller and more widely spaced in the
central portion of the stand and adjacent to the salt marsh. The small size
of the trees and the wide spacing of trees in the central portion of the stand may
indicate that this part of the stand is higher above low water than areas along
the fringe of the stand. Associated with changes in the height of trees and their
spacing are changes in the density of foliage cover of the seedling and sapling
layers. Throughout the mangrove stand is a layer of seedlings persisting at the
two or four leaved stage. The leaves of these seedlings are arranged horizontally
to the ground to intercept a maximum amount of the light diffusing through the
rather dense canopy layer (Fig. 8) (see Horn, 1972 for a discussion of the
adaptive morphology of trees). Whenever there is an opening in the canopy,
the seedlings shoot towards the light and a dense middle storey rapidly develops.

TABLE 5
DENSITY DETERMINATIONS OF MOLLUSCS IN THE MANGROVES

Site Nos.

Species 1 2 3 4 5 6
Saccostrea cucullata 671 171 307 Ey. 1342 167
Bembicium auratum 125 48 51 Te 250 52.
Modiolus pulex Dy 2 2 3 10 1
Chiazacmea flammea 37 12 23 16 74 15
Unknown bivalve 4 1 pi — 8 1
Calliostoma sp. 3 — — a 6 =

Total no. of individuals per m°* 845 234 385 249 1690 236

The marine invertebrate fauna of the mangroves was richer than that found
in the salt marsh both in terms of number of species and individuals. This was
particularly noticeable among the molluscs where 16 species were recorded. The
oyster Saccostrea cucullata was abundant on the lower trunks of mangrove trees
especially those close to the weed beds and creek. Littorina scabra was commonly
found grazing on the epibiota on the surfaces of the mangrove leaves. The densities
of some of the common molluscs found in this zone are given in Table 5.
Though not numerically abundant the pulmonate Onchidium damelli was common
in this zone.

Crabs were common in the mangroves and several species occur, Sesarma
erythrodactyla, Paragrapsus laevis, Heloecius cordiformis and Scylla serrata. Some

116 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

indication of the densities of these crabs observed feeding at night were given
in the preceding section on marine invertebrates. The amphipod Avmpithoe sp.
was also common. Odum and Heald (1972) have shown that amphipods are
important in the shredding of mangrove leaves. The shredding makes the leaves
available to other detritus feeders. A single specimen of Laomedia healyi was
found on the surface of the mud at low tide during night collecting.

The polychaete fauna of the mangroves was virtually restricted to the
channels and creek which flow through the mangroves, Nereis (Hediste) diversicolor
and Nephtys australiensis were the most common species. The nereid Neanthesi
vaalii was found in the oyster communities at the bases of the mangrove trees.
Zostera

Between 0.8m above low water to 1.2m depth contour, the shallows of Careel
Bay are dominated by the marine angiosperm, Zostera capricornis (Fig. 7). Halophila
ovalis was also present, though during the period of our survey it was not
abundant. Among the Zostera were the red algae, Gracilaria verrucosa, and
Hypnea valentiae and the brown alga Dictyota dichotoma. The algae are seasonal
in occurrence and reach a maximum density in late spring and early summer. The
blades of Zostera commonly supported a well developed epibiota community.

The Zostera zone can be conveniently divided into an upper and lower
zone (see Fig. 4). The junction of the upper and lower Zostera occurs at Indian
Spring low water. The lower zone was distingushed by a greater biomass of Zostera
and the presence of the Sydney cockle, Anadara trapezia. The upper zone was
characterized by a sparser plant growth and the occurrence of the Hercules
club-shell or Sydney whelk Pyrazus ebeninus. Biomass measurements were not
made in the upper Zostera. Though the Zostera of the lower zone gave the
impression of a fairly uniform density, biomass measurements indicated a rather
patchy distribution (mean 80.60 gm/m* SD+ 90.32). Such figures are within
the range that McRoy (1970) found. The Zostera in the upper zone was noticeably
patchy with much bare space. In the lower Zostera and in the Posidonia zone
there were scattered circular spots devoid of plant growth and animal life. The
origin of these spots is not known, but may be resting places for rays.

a. Upper Zostera

Far greater numbers of species and individuals were present in this zone
in comparison to the sandy beach. Polychaetes and molluscs dominated the samples
both in terms of numbers of species and individuals.

The dominant polychaete in all samples was Nephtys australiensis but in
the November samples, large numbers of juvenile Haploscoloplos n. sp. were also
present (Table 6). Also in November, large numbers of Nereis (Hediste)
diversicolor were found although this species was previously only found in small
numbers (e.g. February).

Aust. Zool. 18(2), 1974 117

P. A. HUTCHINGS and H. F. RECHER

The most abundant species of molluscs were Macoma deltoidalis and
Nassarius burchardi which was commonly found grazing on the epibiota of the
Zostera. In November, many juveniles of M. deltoidalis were collected. Similarly
numerous juveniles of Notospisula parva producta were present in June and
November samples. Two juveniles of Azadara trapezia were found in February
and June but no adults were found in this zone. Many of the other molluscs
were present in very few numbers e.g. Laternula tasmanica, Eumarcia fumigata,
and Austrocochlea obtusa. |

The crustaceans were represented by more species than in any of the
preceding zones, but many species were collected in small numbers (Table 6).
The amphipod Melita sp. was numerous in two samples collected in November,
but absent from most others. Relatively similar numbers of Macrophthalmus setosus
and M. crassipes were found in both the upper and lower Zostera zones. The
numbers of M. setosus increased greatly in June in both zones. In the June
and November samples, species A and B were collected (see note in marine
invertebrates section). Both these species were more abundant in the lower Zostera.
b. Lower Zostera

Quantitative samples were collected only in June and November. In February
no division of the Zostera into upper and lower zone was made and all the
samples collected in February were from the upper Zostera.

The lower Zostera was the richest of all the zones sampled (Table 6) with
an average of 14-15 species per sample. Polychaetes dominated this zone. The
capitellids Notomastus hemipodus and Barantolla lepte were particularly abundant.
Less common but invariably present were the terebellids, Lysilla pacifica and
Rhinothelepus lobatus. In November large numbers of juvenile Owenia fusiformis
were present indicating successful breeding had occurred between June and
November. The Zostera blades were covered in Spirorbis sp. but no attempt to
quantify them was made. Virtually all the species of polychaetes which were
found in the upper Zostera also occurred in the lower Zostera, but additional
species noticeably the terebellids and spionids were restricted to the lower
Zostera.

The rich mollusc fauna of this zone was characterized by Macoma deltoidalis,
Nassarius burchardi and Anadara trapezia. In June and more notably in November
very small specimens (5-8 mm in length) of A. trapezia were found. The volume
of sediment collected in these quantitative samples 9000cm* was insufficient to
collect adequate numbers of adults for determination of their relative abundance.
Using a quadrat 0.25 m?® in size, the average density of A. trapezia was
8-59/m*, although there was some evidence of clustering. Within a quadrat
anything from 0-6 specimens could be found. The adults range in size from
42-64 mm in height and no obvious year groups could be distinguished (Fig. 9).
While collecting A. trapezia from 40 quadrats, two specimens of the echinoderm
Astropecten polycanthus were found.

118 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

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Aust. Zool. 18(2), 1974

P. A. HUTCHINGS and H. F. RECHER

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Aust. Zool. 18(2), 1974

P. A. HUTCHINGS and H. F. RECHER
In comparison with the molluscs and polychaetes, crustaceans were poorly

represented in this zone. The amphipod Ampithoe sp. and the crabs Macro phina ras
setosus and species A and B were the most abundant species.

300

HEIGHT OF CANOPY (cm)

SALT 60 120 180 _ 240 300 360 420
MARSH
. METRES bapielia
ZOSTERA

Fig. 9. Size distribution of the Sydney Cockle Anadara trapezia at Careel Bay.

Posidonta

The Posidonia flats extend from 1.2m depth contour to a depth of approxi-
mately 11m. The marine angiosperm, Posidonia australis, dominates this zone and
other plant life is absent or epiphytic. Collections of epiphytes on the leaves
of Posidonia were made and sorted, but these have not yet been identified.
Polychaetes and small crustaceans were well represented in these collections.
From 11m, plant growth becomes increasingly sparse and gives way
to a sandy sea floor. The depth to which Posidonia can grow is determined by
water clarity and its occurrence in 10-12 m of water in Careel Bay attest to
the lack of turbidity or of the intrusion of Hawkesbury River water. Biomass
measures in the Posidonia zone showed less variation than those made in the
lower Zostera (287g/m? - SD+57.88.

The Posidonia zone had a large number of species but relatively few
individuals in any one sample in comparison to the lower Zostera zone (Table 6).
This was particularly true of polychaetes where many species were represented
only by one or two individuals e.g. Armandia intermedia, Mediomastus sp.,

122 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

Chaetopterus sp. and Cirratulus sp. This zone had some very large species of
polychaetes present, Glycera americana, Marphysa sanguinea and Chaetopterus sp.
and it is possible that the biomass of polychaetes would be greater in the Posédonia
than in the lower Zostera, even though -far fewer species are present.

Molluscs were poorly represented in this zone and the only numerically
abundant species was Cacozeliana lacertina. Similarly crustaceans were poorly
represented in the quantitative samples but this was probably a reflection on
sampling techniques. As in the lower Zostera the amphipod Ampithoe sp. was
common.

DISCUSSION

Without measuring energy flow, comments on the ecology of Careel Bay
and its importance to the total estuary must necessarily be speculative. However,
to judge by the number of species, the presence of many juvenile fish and the
total standing biomass of plants and animals in Careel Bay, there is a significant
production of organic matter in the salt marsh, mangrove and sea grass habitats.
The invertebrate fauna of the salt marsh, mangrove and sea grass habitats is
dominated by detritus feeding animals. It appears similar to the mangrove-
invertebrate system described in detail for Florida by Odum and Heald (1972).
Organic matter in the form of leaves and fruits enter the estuary where they
are attacked by bacteria, fungi and small invertebrates. These break down the
material to a size which can be ingested by other detritus feeders. Through a
long series of steps in which the original matter of a mangrove leaf may be
reingested many times, the complex molecular structure of the leaf is broken
down and its energy released. The detritus feeders are in turn fed upon by
predators. A later paper will discuss the possible reasons for the zonation of
animals in the Bay and the potential food web operating both within and between
the zones.

Our results have shown that Careel Bay is a very rich area in terms of
numbers of species present. A similar ecological and geographical area to Careel
Bay is Towra Point in Botany Bay which has been sampled by Weate (pers.
comm.) using the same quantitative methods used in Careel Bay. She found
in June, 30 species of animals in the Zostera and 13 species in the Posidonia
weed beds. By comparison in Careel Bay, 51 and 54 species were found re-
spectively in the Zostera and Posidonia weed beds. The reasons for the greater
number of species being present in Careel Bay are not fully understood but parts
of Botany Bay are heavily industrialized and extensive dredging occurs. Dredging
must have deleterious effects on pelagic larvae which pass through the dredge.
The majority of animals found in estuarine situations have pelagic larvae.

We have looked at other areas in the Hawkesbury-Broken Bay system, notably
Patonga Creek (Broken Bay), Spectacle Island (Hawkesbury River) and Riley’s
Island (Brisbane Waters), which have many fewer species compared.to Careel

Aust. Zool. 18(2), 1974 123

P, A. HUTCHINGS. and Ri \F.. RECHER

Bay. However, individual species (e.g. Macrophthalmus setosus, Mictyris longz-
carpus and Callianassa sp. achieve considerably greater population densities on
the tidal flats of Patonga than at Careel Bay.

One of us (Hutchings, 1974) has sampled in Wallis Lake, N.S.W. This is
a coastal lagoon continually open to the sea but at times it receives large volumes
of freshwater. A total of 32 species of polychaetes were found along the eastern
shores of the lake in the weed beds of Zostera and Halophila. All the species
present in Wallis Lake are also present in Careel Bay. Weate (pers. comm.)
has sampled in the adjacent Smiths Lake, which is often closed to the sea. Sampling
in December, using a hand operated corer, a total of 28 animal species were
found. Similarly, surveys conducted by the N.S.W. Division of the Australian
Littoral Society have shown that the numbers of species of animals decreases
progressively up the estuary as the salinity falls. All the areas mentioned above
have some species in common, e.g. Polychaetes, Australonereis ehlersi, Nereis
(Hediste) diversicolor, Marphysa sanguinea, Haploscoloplos sp. and Notomastus
hemipodus; Molluscs, Nassarius jonasi, Pyrazus ebeninus, Macoma deltoidalis;
Crustaceans, Paragrapsus laevis, Macrophthalmus sp. and Halicarcinus ovatus.

It appears that Careel Bay represents the richest estuarine area yet sampled
in N.S.W. This can partially be attributed to its almost totally marine situation,
but other factors may be involved. We hope to elucidate these other factors by
continuing to survey wetland areas on the N.S.W. coast.

It is very difficult to compare our work with studies done overseas as
sampling techniques vary widely, but some comments can be made on geographically
similar temperate areas. Day (1967) working on South African estuaries has
found that estuaries vary in the richness of their fauna and he attributes this to
the amount of freshwater entering the estuary and to the clarity of the water.
The richest area comparable to Careel Bay that he has studied is the Knysna
estuary which has clear water, allowing dense beds of sea grasses to flourish.
Only small quantities of fresh water enter the river throughout the year. Other
estuaries in the same temperate region are subjected to flooding and have
eroding river banks, reducing clarity of water, hence weed growth is diminished
and far fewer species of animals are present. Day (1967) also suggests that a
wide opening to the sea (such as Careel Bay has) is important in maintaining
water exchange and tidal movement. Day has recorded 350 species from Knysna
estuary including, 69 polychaetes, 12 crabs, and 40 fishes. Considering that Careel
Bay in comparison is a very small area and only the larger components of the fauna
have been collected, the number of species of polychaetes is of the same order of
magnitude. The number of species of polychaetes (at present 46) seems certain to
increase, once the epibiota of the Posidonia and Zostera is analysed. Careel Bay
is thus similar in species diversity to temperate South African estuaries and is
the richest wetland area yet sampled in New South Wales.

124 Aust. Zool. 18(2), 1974

FAUNA OF CAREEL BAY

ACKNOWLEDGEMENTS

We should like to thank the New South Wales Division of the Australian Littoral
Society for allowing us to use their preliminary data and for supplying some of the
photographs. We are particularly indebted to Diane Brown, Phil Colman, Rod Fletcher,
Geoff Holloway, Helen Larson, Dan Lunney, Dr. David McAlpine, Dr. John Paxton,
Dr. Gary Poore, Hymo Posamentier, Penny Weate and Peter Whalan for help in collecting,
identifications, providing photographs and information on the insect fauna. Finally we
should like to thank all the marine Curators at the Australian Museum for assisting in
the identifications. Our grateful thanks also to Drs. Clark, Griffin, Hodson, Rainer and
Talbot and Mr. Dunstan for reviewing the manuscript.

REFERENCES

CLARK, R. B. (1961). The origin and formation of the heteronereis. Biol. Rev. 36, 199-227.

CLARK, R. B. (1965). Endocrinology and the Reproductive Biology of Polychaetes.
Oceanogr. mar. Biol. Ann. Rev. 3, 211-255.

CLARKE, L. & N. HANNON (1967). The mangrove swamp and salt marsh communities
of the Sydney district. I. Vegetation, soils and climate. J. Ecol. 55, 753-772.

CLARKE, L. & N. HANNON (1969). The mangrove swamp and salt marsh communities
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CLARKE, L. & N. HANNON (1970). The mangrove swamp and salt marsh communities
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CLARKE, L. & N. HANNON (1971). The mangrove swamp and salt marsh communities
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DAY, J. H. (1967). The Biology of Knysna estuary, South Africa in “Estuaries”,
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HARTMAN, O. (1954). Australian Nereidae. Trans. R. Soc. S. Aust. 77, 1-41.

MLE RSEE Yoko ha Poe Aly HUTCHINGS, & A. Py RECHER, (1973). Careel Bay:
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HORN, H. S. (1971). The Adaptive Geometry of Trees. Monographs in Population Biology.
3. Princeton University Press, 133 pp.

HUTCHINGS, P. A. (1974). The polychaeta of Wallis Lake, N.S.W. Proc. Linn. Soc.
N.S.W. 98, 175-195.

Aust. Zool. 18(2), 1974 125

P. A. HUTCHINGS and H. F. RECHER

KRATOCHVIL, M., N. J. HANNON & L. D. CLARKE (1972). Mangrove swamp and
salt marsh communities in Southern Australia. Proc. Linn. Soc. N.S.W. 97, 262-275.

LONGBOTTOM, M. R. (1968). Nutritional factors affecting the distribution of Arenicola
marina L. Ph.D. Thesis, University of London.

LYFORD, J. & H. PLINNEY (1968). Primary Productivity and community Structure of
an Estuarine Impoundment. Ecology, 49, 854-866.

MACNAE, W. (1966). Mangroves in eastern and southern Australia. Aust. J. Bot. 1/4,
67-104.

MACNAE, W. (1967). Zonation within mangroves associated with estuaries in North
Queensland. In “Estuaries” Amer. Assoc. Advan. Sci. Publ. 83, 432-441.

McROY, C. P. (1970). Standing Stocks and other Features of Eelgrass (Zostera marina)
Populations on the coast of Alaska. J. Fish. Res. Bd. Can. 27, 1811-1821.

NEWELL, R. (1965). The role of detritus in the nutrition of two marine deposit-
feeders, the prosobranch Hydrobia ulvae and the bivalve Macoma balthica. Proc. zool.
Soc. Lond. 144, 25-45.

ODUM, E. P. (1961). The Role of Tidal Marshes in Estuarine Jieeche ernionn: The Con-
servationist June-July 1961, 12-15.

ODUM, E. P. & A. A. DE LA CRUZ (1967). Particulate organic detritus in a Georgia salt
marsh estuarine ecosystem. In “Estuaries”. Amer. Assoc. Adv. Sci. Publ. 83, 383-388.

ODUM, W. E. & E. J. HEALD (1972). Trophic analysis of an estuarine mangrove
community. Bull. Mar. Sci. 22, 671-738.

SCHROEDER, P. C. (1968). On the life history of Nereis grubei (Kinberg), a Polychaete
Annelid from California. Pacif. Sci. 22, 476-481.

SNELLING, B. (1959). The distribution of intertidal crabs in the Brisbane River.
Aust. J. Mar. Freshwat. Res. 10, 67-83.

APPENDIX I
Insects Recorded from Careel Bay, N.S.W.

Diptera
Family:
Tephritidae
Euphranta n. sp.
Dacus (Strumeta) tryoni
Family:
Platystomatidae
Rivellia n. sp.
Family:
Lauxaniidae
Panurgopsis n. sp.
Prochaetops n. sp.
Chilocryptus sp.
Trigonometopsis n. sp.
Trigonometopus sp.
Family:
Carnidae
Australimyza sp.
Family:

Milichiidae
Desmometopa sp.

126 Aust. Zool. 18(2), 1974

FAUNA OF. CAREEL BAY

Family:
Cryptochetidae
Cryptochetum sp.
Family: an ay
Tethinidae
Dasyrhicnoessa sp.
Family:
Canaceidae
Canace sp.
Xanthocanace sp.
Family:
Chloropidae
Merodonta sp.—Southern Record for genera
Pemphigonotus sp.—Southern Record for genera

Lepidoptera

Family:
Oxychirotidae
Cenoloba obliteratis

Hymenoptera
Family:
Ichneumonidae
Echthromorpha_ intricatoria Fabricius
Lissopimpla_ excelsa Costa
Netelia producta Brullé
Eriborus sp. (parasite of Coenoloba sp.)
Sphinctus sp.
Family:
Braconidae
Priaspis sp.
Chelonus unimaculatus Szepligeti—first record since original description
Chelonus australiensis Szepligeti—first record since original description
Phanerotoma australiensis Ashmead
Apanteles ruficrus Haliday
Microgaster sp.
Opius sp. (parasitic on Euphranta sp.)
Family:
Evaniidae
Evania .appendigaster Linné
Family:
Sphecidae
Sphex fumipennis
Sphex globosus
Sceliphron laetum
Isodontia nigella
Isodontia sp.
Larra sp.
Pison sp.
Family:
Tiphiidae
Diamma bicolor
Family:
Vespidae
Polistes tasmaniensis

Aust. Zool. 18(2), 1974

127

P. A. HUTCHINGS and H. F. RECHER

Family:
Apidae
Apis mellifera
Many of these insects which have been identified only to genus, are new species.

APPENDIX II

The following fishes were collected in Careel Bay at various times of the year, and
in the Hawkesbury River estuary on November 23, 1972, at low tide by 20’ seine.
The following are the individual collecting localities: (1) beach at eastern side of Dangar
Island; (2) Fisherman’s Beach, Cowan Creek, Broken Bay; (3) Head of Jerusalem Bay,
Cowan Creek; (4) Careel Bay. The numbers after the species refer to the various localities.

Family Urolophidae-stingrays Family Sparidae—bream, tarwhine, snapper
Urolophus testaceus 2 Acanthopagrus australis 2, 3
Family Clupeidae—sardines, herrings, sprats Rhabdosargus sarba 2, 3, 4
Hyperolophus vittatus 2 Family Mullidae
Family Syngnathidae—pipefishes and seahorses Upeneichthys tragula 4
Hippocampus sp. 4 Family Mugilidae—mullet
Stigmatophora nigra 2, 4 Myxus elongatus 1, 2, 3
Urocampus carinirostris 1, 3 Family Pomatomidae—tailor
Family Hemiramphidae—garfishes Pomatomus saltatrix 2
Hyporhamphus ardelio 2 Family Clinidae
Family Atherinidae—hardyheads and Cristiceps australis 4
blue eyes Family Gobiidae—gobies
Pranesus ogilbyi 2 Arenigobius frenatus 1,' 3, 4
Taeniomembras microstomus 3 Bathygobius kreffti 1, 2, 3, 4
Pseudomugil signifer 3 Favonigobius exquisitus 3, 4
Family Scorpaenidae—fortesques and Lizagobius olorum 3, 4
red perches ~- Nesogobius pulchellus 4
Centropogon australis 1, 2, 3, 4 Redigobius macrostomus 1, 3
Family Centropomidae—glass perches Family Bothidae—left eyed flounders
Velambassis jacksoniensis 1, 2, 3, 4 Pseudorhombus jenynsii 1
Family Platycephalidae—flatheads Family Balistidae
Platycephalus sp. juv. 2 Meuschenia skotlower 4
Family Enoplosidae—old wives Family Aluteridae—leatherjackets
Enoplosus armatus 2 Navodon skottowei 2, 3
Family Theraponidae—grunters Monocanthus macrolepis 4
Pelates sexlineatus 1, 3, 4 Family Tetraodontidae—toados
Femily Carangidae Torquigener hamiltoni-2, 3, 4
Caranx sp. 4. Family Diodontidae—porcupine fishes
Family Gerridae—silver biddies Dicotylichthys myersi 2, 3
Gerres ovatus 3, 4 Diodon punctulatus 4

Family Scorpidae—luderick, sweep
Girella tricuspidata 3, 4

128 ‘ Aust. Zool. 18(2), 1974

Population Movements of the Agamid Lizard
Amphibolurus nobbi

GEOFFREY J. WITTEN
Zoology Department, University of New England, Armidale, N.S.W. 2351*.

Amphibolurus nobbi nobbi Witten is associated with arenaceous soils. It
occurs commonly in the granite belt of southern Queensland, is present in many
of the sandy granite areas of northern N.S.W. and also inhabits sand dunes of
the northern N.S.W. coast (Witten, 1972). During a study of its taxonomic
status and thermoregulation, population movements became evident and are
presented here.

Snout-vent lengths were measured on preserved specimens, and these were
related to the date and area of collection. Specimens of less than 35 mm s-v
length were considered hatchlings, specimens from 35 to 50 mm as juveniles and
specimens over 50 mm as adults. These categories were established on the basis
of (1) minimum size of breeding adults, (2) the condition of the yolk sac scar,
and (3) the size at which yellow and red secondary colouration first appeared.
Only specimens from the northern tablelands of N.S.W. were studied, as breeding
areas outside this region were not so clearly defined topographically.

Two types of area were inhabited, breeding areas, and areas into which
animals dispersed. Three breeding areas are considered in this study. All were
estimated to have an area of between one and two hectares. Each consisted
of a steep north-facing slope. Similar, though less extensive data were obtained
from areas outside the region.

In Autumn and Spring large numbers of adult A. x. nobbi are found in the
breeding areas (Fig. 1) but desert them during the summer months. Adults emerge
from hibernation in the first half of October. Dispersal of adults from the
breeding areas takes place before the second week of December. Adults return
to the breeding areas in autumn, from mid-February to the first week of March.
Adults retreat for hibernation in late March or early April.

Emergence of hatchlings occurs from mid-October until mid-April. There
are peaks in numbers of hatchlings in October and March-April. The emergence
of hatchlings in spring (October) indicates that eggs or young overwinter in
the soil. As it is normal for smaller individuals to emerge earlier in the
spring (Cowles, 1941; Fitch, 1956), emergence of hatchlings later than adults
may indicate that eggs complete their development in the spring and then hatch.

* Current address “Wyndella”, Barraba: N.S.W.. 2347.

Aust. Zool. 18(2), 1974 129

WITTEN

GEOFFREY.” J:

Kew

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SION ‘s][npe pue sojusAn{ ‘s8urpyojey JO sol10d9}e OpIAIP soul] [eJUOZIIOH{ ‘svoie
SUIPIIIG-UOU }BOIPUI So[dIID paso[d ‘svoIe SUIPseIq WOIJ susUIDedS 9s}JeUSISOp so[dIID
usdQ “UOTSEI[0D IW9Yy] JO 33ep 9Y} JsuTese poyjord susudeds Jo syJsuUs, JUSA-JnNOUS “[ ‘sIY

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18(2), 1974

Aust. Zool.

130

POPULATION MOVEMENTS OF AMPHIBOLURUS

Similar observations have been made on several species of turtles (Sexton, 1957;
Gibbons, 1969), but to the author’s knowledge no such observations are available
for lacertilians.

Hatchlings with fresh yolk sac scars were from 25 to about 32 mm in s-v
length. They move out of the breeding area almost immediately; no juvenile
of more than 35 mm s-v was collected within a breeding area. Juveniles
apparently grow to adult size outside the breeding areas, and migrate to such
sites upon attaining sexual maturity. Bradshaw (1971) observed juvenile emigration
from an area dominated by adults in another agamid species, A. ornatus. However,
this species demonstrates territorial behaviour and this movement was explained
by Bradshaw on the basis of adult aggression against juveniles. A. n. nobbi
is apparently not territorial in behaviour, and at least some juveniles vacate the
breeding area while it is unoccupied by adults.

Territorial behaviour has been observed in several species of Amphibolurus
(Carpenter e¢ al., 1970; A. inermis, A. barbatus, A. muricatus). Neither in the
field, nor under confined laboratory conditions was territorial behaviour ever
observed in A. nv. nobbi. The presence of bright colouration (which is largely
lacking in its territorial relatives) is puzzling, but may be involved in courtship
displays; however, none were observed. This hypothesis is supported by the fact
that sexual dichromatism is much more pronounced during the spring months
than at other times of the year.

Migration to oviposition areas has been observed in Iguana iguana (Hirth,
1963; Rand, 1968); in that species only gravid females undertake the migration.
This movement is of short duration, and activity in the oviposition area is
concerned with finding a suitable nest site, nest digging and oviposition. Spent
females return to their usual habitat within a few days after oviposition
(Montgomery ef al., 1973). In A. n. nobbi the entire adult population is
resident in the breeding area for 4 of the 62 month adult activity season.
A. n. nobbi also differs from I, iguana in that the breeding and non-breeding
areas are basically similar, differing in gradient of slope and in the aspect
presented to the sun.

There was no direct observation of individuals moving between two areas.
A number of specimens were collected at a distance of six km. from known breeding
areas in two localities, and one was taken some fifteen km. from the nearest
known breeding area (coastal sand dunes). These observations suggest that the
lizards are capable of moving relatively long distances, but further study is
required to obtain reliable estimates.

ACKNOWLEDGEMENTS

I am indebted to Associate Professor H. Heatwole for advice, assistance and
criticism. Many of the staff and postgraduate students of the Department of Zoology,
University of New England also offered criticism of the manuscript.

Aust. Zool. 18(2), 1974 | 131

GEOFFREY J. WITTEN

REFERENCES
BRADSHAW, S. D. (1971). Growth and mortality in a field population of Amphibolurus
lizards exposed to seasonal cold and aridity. J. Zool., Lond. 165, 1-25.

CARPENTER, C. C., J. A. BADHAM & B. KIMBLE (1970). Behaviour patterns of three
species of Amphibolurus (Agamidae). Copeia 1970, 497-505.

COWLES, R. B. (1941). Observations on the winter activities of desert reptiles. Ecology
22, 125-140.

FITCH, H. S. (1956). Temperature responses in free-living amphibians and reptiles of
north-eastern Kansas. Univ. Kansas Publ. Mus. Nat. Hist. 8, 417-476.

GIBBONS, J. W. (1969). Ecology and population dynamics of the chicken turtle,
Deirochelys reticularia. Copeia 1969, 669-676.

HIRTH, H. F. (1963). Some aspects of the natural history of Iguana iguana on a
tropical strand. Ecology 44, 613-615.

MONTGOMERY, G. G., A. S. RAND & M. E. SUNQUIST (1973). Post-nesting move-
ments of iguanas from a nesting aggregation. Copeia 1973, 620-622.

RAND, A. S. (1968). A nesting aggregation of iguanas. Copeia 1968, 552-561. _
SEXTON, O. J. (1957). Notes concerning turtle hatchlings. Copeia 1957, 229-230.

WITTEN, G. J. (1972). A new species of Amphibolurus from eastern Australia.
Herpetologica 28, 191-195.

132 Aust. Zool. 18(2), 1974

Capture and Marking of the Platypus,
Ornithorhynchus anatinus, in the Wild

School of Zoology, University of New South Wales,
P.O. Box 1, Kensington, N.S.W. 2933.

T. R. GRANT and F. N. CARRICK

We began trapping platypus in early 1971 as part of our investigation of
temperature regulation (T.R.G.) and reproductive physiology (F.N.C.) in this
species. Standard gill nets have been used in the past (Temple-Smith personal
communication) to capture platypus but accidental drowning of some animals
has always been a problem with the use of these nets. Over the last two years
we have been able to modify this technique to minimise the risk of drowning

animals.

As part of our studies we wish to observe animals in the wild to
determine how long platypus remain in water and to investigate the relationship
of this to ambient temperatures. This requires the capture and identification of

LP KS LEELA LL LING

7

< SX ER ees

Fig. 1. Piacement of nets in a stream

a. ropes **2 bi nets

Aust. Zoal. 18(2), 1974

c. direction of water flow

Oo Zu D
eh RAN aN NS ike
7s :

Fan salt ey a XZ

\A

133

T. R. GRANT and F. N. CARRICK

platypus wtih visible, individual marks. With a view to possible longer term
studies these animals are also being marked with leg bands.

The increasing interest in the biology of the platypus in recent years has
induced us to write this short paper to make our experiences available to others.

For capture we normally use two 338-3 ” mesh nets laid in the stream
or lake parallel to the bank (Fig. 1). These nets vary between 50 and 150
feet in length and in deep water nets having a drop of 8 feet are used, while
in shallow water or over weed beds nets of 4 feet drop are necessary.

The standard fishing net has floats on the top and weights on the bottom.
We have modified our nets by painting the floats white and removing the weights.
The white floats allow the nets to be seen easily in the beam of a spotlight to
facilitate checking at regular intervals during the night. The lack of weights
makes it easier for an entangled animal to carry the net to the surface. It is
necessary to use some weights in areas of stronger currents. However, the selection
of the waterway is very important, and areas of rapid water flow, large numbers
of snags, or rapidly changing water levels should be avoided. In such areas the
risk of drowning animals is high and in our experience capture rates are usually
low. In areas where the water temperature is very low, the need to remove
animals from nets as soon as possible after capture is an important consideration.
However, platypus are also very susceptible to heat stress and should be kept
cool after capture and during transport in hot weather.

Snags, fish and other heavy objects caught in the nets account for quite a
number of drownings. To lessen this hazard the nets are lifted every hour, to
release fish and remove inanimate objects. Brief scanning of the nets every 10-15
minutes with a spotlight allows the detection of movement and seems to have no
marked effect on capture rates.

Since we began using unweighted nets in December 1972, we have caught
40 platypus (which have been marked and released except for the small number
of animals taken back to be maintained in the laboratory) without a single
incident of drowning. It must be noted however that the capture of platypus
in the wild is far from simple. It is time consuming, arduous and rates of
capture can be quite low in some areas. The procedures outlined above only
serve to show that these animals cam be caught safely if certain precautions
are taken.

Visible marks consist of adhesive bandage tape, painted with oil-based
fluorescent paint attached to the tail. Plain colours are visible with field glasses
at distances of up to 100 yards, and patterns can be distinguished at shorter
distances. When applied firmly to the animal they will stay attached for around
10 days to a month. These tail marks are satisfactory for individual recognition
of animals for short term observation work. We also intend to use them to
allow visual location of animals to which transmitters have been attached. This

134 Aust. Zool. 18(2), 1974

CAPTURE AND MARKING OF THE PLATYPUS

should remove the necessity to locate an animal by the radio signal and allow
the use of transmitters with only a moderate range.

Leg bands of stainless steel rings 2.0 cm in diameter and 0.5 cm in
width are applied to the shank of the rear foot using pliers to overlap the
ends (Fig. 2). These are inscribed with a number and address. These bands
were tried on captive animals before being used to mark wild ones. No damage
to the legs of two captive platypus occurred and three wild animals caught after
being banded for 7 months also showed no ill effect from the bands. These bands
can be seen when an animal grooms itself in the water, but they are of limited
use for observational purposes. They can, however, be used in mark and recapture
studies.

We wish to thank Stockbrands Pty. Ltd., Mt. Hawthorn, W.A. for manufacturing the
bands to our specifications. Bryan Shadbolt made the initial suggestion to remove all the
weights from one of our nets. Thanks also go to various technicians and friends who
have spent many hours helping to tend nets. The N.S.W. National Parks and Wildlife
Service and N.S.W. State Fisheries Branch are acknowledged for allowing us to trap
platypus in N.S.W. Part of the support for this work was provided by a grant from
the Australian Research Grants Committee.

Fig. 2. Marked animal showing leg band
(photo: Bob ~ McBlain).-

Aust. Zool. 18(2), 1974 135

ROYAL ZOOLOGICAL SOCIETY OF NEW SOUTH WALES
Established 1879
Registered under the Companies Act, 1961

COUNCIL, 1973

President:
Gilbert Percy Whitley, F.R.Z.S.

Vice-Presidents:
Ernest Jeffrey Gadsden, F.R.Z.S.

Gordon Clifford Grigg, B.Sc., Ph.D.
William Steel

Rotalde Strahan. NMUSc. E.LS:. F.R-ZSs., .M.F. Biol., F.S.1.H:
Honorary Secretary: Richard M. Mason, M.B., B.S.
Honorary Treasurer: Ronald Strahan

Honorary Editors: Jack Harvey Prince, F.B.O.A., F.A.A.O., F.Z.S., F.R.M.S.
(“Koolewong”’)

Edward S. Robinson, M.Sc., Ph.D. (“The Australian Zoologist’”)

Members of Council:

Mrs. A. M. Clayton Jonn Moore Smail, L1.B.

Henry John de Suffren Disney, M.A. William Steel

Mrs. X. Forbes Ronald Strahan

UV. Eric Friese, B.Sc. Frank Hamilton Talbot,

Ernest Jeffrey Gadsden Messe. PhD. ES. FReZ-S.
Gordon Clifford Grigg Ellis Le Geyt Troughton,
Francis McCamley CWEZS.. FRUZS:

Richard M. Mason Gilbert Percy Whitley

Jack Harvey Prince Mrs. Olive Wills, F.R.Z.S.
Edward S. Robinson Mrs. Mary Wray

Members of Publications Committee:
U: E. Friese, G. C. Grigg, J. H. Prince and E. S. Robinson

MEMBERSHIP OF THE SOCIETY
- (The Society's year commences on Ist July)

APPLICATION FOR MEMBERSHIP

should be addressed to the Honorary Secretary, Royal Zoological Society of
New South Wales, Taronga Zoo, Mosman, New South Wales, 2088.

Class Amount of Subscription
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THE AUSTRALIAN ZOOLOGIST

VOLUME 18 1974

Contents

Physiological adaptations of anuran amphibians to aridity: Australian: ‘pros
Re S. DEY MOURS and Aakers bicteriatrt tetas ila hs AS: eee

The status of Pseudomys Bical ee bres (the New Holland h
i. POSAMENTIER and H. F. REGHIER 20 see Se

Occurrence and field seeoaiien of ee parma. 1G Me

Gens dalhousiensis n.sp., a “sexually acai Lehane |
(Atherinidae) from South Australia. ave IVANTSOFF and ioe
GLOVER Pee Be cscrtn. es snes nnnnnnnennsnnnntennnnn

The fauna be Cea Bay ‘with comments on the ee a mai

| sea-gtass communities. P. A. UCHINGS and A. F,

=

Short papers—

Population movements of he agamid lizard Amphibolurus nobbi. G
WITTEN rnin bi

Grpeare: and marlin of Ae platypus, Ornitborkyncbus anatinus, ‘in the 1

1 R. GRANT and EN. CARRICK on (aha ane Ae eae

VEN

mn |

Volume 18, Part 3

August, 1975

=

ee EAE ———— ete
2
1 ee
if is)
i
os
i OLE

NOTICE TO AUTHORS

©

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THE AUSTRALIAN ZOOLOGIST

VOL. 18 AUGUST, 1975 PART 3

Eptatretus longipinnis, n. sp., A New Hagfish (Family
Eptatretidae) from South Australia, with a Key to the
5-7 Gilled Eptatretidae.

R. STRAHAN
Australian Museum, College St., Sydney, N.S.W. 2000

ABSTRACT

A new species of hagfish, Eptatretus longipinnis (Family Eptatretidae) based on
a specimen from coastal waters near Robe, South Australia, is described. The
species differs from other members of the genus in having a well developed
ventral fin-fold extending into the branchial region. The genus Paramyxine is
regarded as a junior synonym of Eptatretus. A key is given to the species of
Eptatretus having 5-7 pairs of gills.

INTRODUCTION

Until 1971 no hagfishes had been recorded from Australian waters. Since
then a number of specimens of Eptatretus cirrhatus have been taken from the
Pacific Ocean off the coast of New South Wales in deep trawls (300-700m).
Also in 1971, a quite different hagfish was taken from the south-eastern Indian
Ocean near Robe, South Australia from a depth of approximately 40m. The
hagfish had burrowed into the flesh of the abdomen of a spiny lobster and was
still embedded when the lobster was brought to the surface. The specimen,
preserved in alcohol, was forwarded to the author by Dr. C. J. M. Glover of
the South Australian Museum.

DESCRIPTION OF EPTATREUS LONGIPINNIS, N. SP.

Holotype: A specimen 422mm long from south-eastern Indian Ocean off
Robe, South Australia (149°43’E, 37°10’S) at depth of approximately 40m.
In the South Australian Museum, Adelaide, South Australia (registration number
F4042).

Major dimensions mm % length
Rostrum to first branchial aperture 120 29
Rostrum to last branchial aperture 139 33
Rostrum to origin ventral fin-fold 112 26
Rostrum to posterior border cloaca 387 OZ
Maximum depth of body 18 4

Aust. Zool. 18(3), 1975 137

R. STRAHAN

Mucous glands range mean
Prebranchial series 27-29 28
Branchial series 5-6 6
Abdominal series 64 64
Caudal series 8-9 9
Total 104-108 106

Lingual teeth: inner 2+-5; outer 3-5.

The body is elongate and subcylindrical, slightly deeper than broad, tapering
towards either extremity from the middle. The skin is dusky-brown dorsally,
clearer brown ventrally. A very slight paling of the skin marks the position of the
eyes on the dorsolateral surface of the head, about 18mm behind the rostrum. The
distal thirds of the nasal and anterior oral barbels are yellow, as are the margins
of the caudal fin and ventral fin-fold. There is no change of pigmentation to mark
the borders of the external branchial apertures.

The nasal rostrum is bluntly curved, overlying the nostril which is approxi-
mately three times wider than high. The nasal barbels are 7mm long, slender,
and taper evenly to a point. (The left ventral nasal barbel is bifid at the tip,
possibly the result of injury and regeneration). The anterior oral barbels are
8mm long and similar in shape to the nasal barbels. The posterior oral barbels,
which partially overlie the mouth in the anterior midline, are approximately 4mm
long and 2.5mm wide.

Six pairs of gills communicate with the exterior by short, nearly vertical,
efferent branchial ducts which are approximately equal in length except for the
most anterior, which is approximately twice the length of the others due to
dorsad displacement of the first gill. The branchial apertures are ventral, arranged
on each side of the body in a longitudinal series curving towards the midline as
it passes posteriorly. The aperture of the pharyngocutaneous duct, shared with
the efferent duct of the sixth gill on the left side, is irregular in outline, somewhat
longer than wide.

Bo em

Fig. 1. Eptatretus longipinnis, nm. sp., holotype. (a) lateral view; (b) ventral
view of branchial region showing external branchial apertures, mucous
glands and ventral fin-fold.

138 Aust. Zool. 18(3), 1975

A NEW HAGFISH FROM SOUTH AUSTRALIA

The cloacal aperture is a longitudinal slit surrounded by lateral lips. The caudal
fin, which originates dorsally and ventrally at the level of the posterior end of
the cloacal aperture, is bluntly rounded and has a height equal to the maximum

height of the body.

A prominent mid-ventral fin-fold originates anterior to the branchial region
and extends between the branchial apertures to the anterior border of the cloacal
aperture. It reaches a maximum depth of 4 mm in the branchial region.

The prebranchial series of mucous glands begins about 30 mm behind the
rostrum on the ventral surface and extends to just in front of the first branchial
aperture. The branchial mucous glands are immediately mesial to each branchial
aperture except that of the sixth gill on the left side which lacks an associated
gland. The abdominal series extends from the posterior end of the branchial
region to about 10mm in front of the border of the cloacal aperture. The caudal
series extends from the level of the middle of the cloacal aperture to about half
the length of the tail.

The lingual tooth plates bear eight teeth in the outer row and seven in the
inner. The bases of the anteriormost three teeth of the outer row are fused to-
gether, as are those of the anteriormost two of the inner row.

TAXONOMY OF POLYBRANCHIATE MYXINOIDS

The status of Bdellostoma and Paramyxine

The most recent comprehensive review of the Order Myxinoidea (Holly,
1933) recognises two genera of polybranchiate myxinoids: Bdellostoma and Para-
myxine. There is long-standing disagreement on the proper name of the former
genus, European workers tending to favour Bdellostoma Muller, 1835, while
others favour Eptetretus Cloquet 1819. Supporters of Bdellostoma (e.g. Rauther,
1924; Holly, 1933) argue that Bdellostoma has had greater usage but, while this
maybe true, it cannot be maintained that Eptatreus (or Heptatretus) is a for-
gotten name, resurrected on formal grounds of priority, for it has been in con-
tinued use since its introduction. Apstein (1915) included Bdellostoma in a list
of recommended nomina conservanda but the International Commission on Zoo-
logical Nomenclature has ruled (Opinion No. 74) that it has no power to adopt
this list ex bloc. In the absence of a ruling to the contrary by the International
Commission, the genus is properly called Eptatretus.

The retention of Eptatretus as a subgenus to include the 5-8 gilled poly-
branchiate myxinoids has little value and has been criticised by Bigelow and
Schroeder (1948), Adam and Strahan (1963), and Hubbs (1963). However,
full consideration of this must await a revision of the entire genus, including
those species at present included in the subgenus Polistotrema.

Aust. Zool. 18(3), 1975 139

R. STRAHAN

The diagnosis of the genus Paramyxine Dean, 1904 is based on the branchial
anatomy of the type of P. atami Dean: ‘“Hyperotretes with branchial apertures
ventrad of sacs. Ectal branchial ducts of distinctly unequal length, the most
anterior several times the length of the most posterior. The duct of the most
anterior gill opening at the surface opposite the fourth (or fifth) gill sac. Openings
of the branchial ducts drawn close together and compressed transversely, that of
the ductus oesophagoeus, however, longitudinally, the latter aperture of large
size, its length equalling the sum of the interspaces of several gills. Transverse
constrictor muscles of the branchial region developed as a distinct element in the
region of the branchial sacs”. (Dean 1904, p. 22).

Studies subsequent to Dean’s erection of the genus have called these criteria
into question. (1) Although the branchial apertures of the species assigned to
Paramyxine tend to be closer to the mid-ventral line than in those traditionally
assigned to Eptatretus, the distinction is difficult to assess, and, in the case of E.
longipinnis, not valid. (2) Differences in length of the anterior and posterior ef-
ferent branchial ducts does not separate Paramyxine from Eptatretus: in E. burgeri
the anteriormost efferent branchial duct is 2-3 times longer than the most posterior
(Strahan, 1962). In E. cirrhatus it is approximately twice the length, three ran-
domly chosen specimens having the respective ratios 23:12, 24:11, 2:1. In E.
longipinnis the anteriormost efferent branchial duct is twice the length of the
others but this is due to a dorsad displacement of the first gill rather than a
caudad displacement of its efferent branchial aperture. (3) The first branchial
aperture of P. atami Dean commonly opens at the level of the fourth or fifth gill
but may open at the level of the third gill, as in E. cirrhatus. (4) The shape of
the branchial apertures of P. atami Dean is extremely variable (Matsubara, 1937;
Strahan and Honma, 1961): in P. springeri Bigelow and Schroeder, the apertures
are almost circular and in P. yangi Teng they are semicircular. (5) The common
aperture of the pharyngocutaneous duct and the last gill of the left side is not
normally elongated longitudinally in P. atami Dean but is subject to considerable
variation (Strahan and Honma, 1961): it is almost circular in P. springeri Bigelow
and Schroader and P. yangi Teng: its size does not bear the relationship to the
intervals between the branchial apertures suggested by Dean. (6) The transverse
branchial constrictor muscles of E. burgeri and E. cirrhatus are as well developed
as those of P. atami Dean.

On the above consideraticrs, I am of the opinion that Paramyxine must be
regarded as a junior synonym of Eptatretus.

Taxonomic characters

Regan (1912) diagnosed the polybranchiate myxinoids in terms of the number
of gills, the position of the anteriormost branchial aperture, and a rigidly defined
dental formula. Holly (1933) followed Regan’s general plan but indicated a
range of variation in the dental formula and followed Dean in taking note of the
position of the posterior end of the lingual musculature with respect to the gill
series.

140 Aust. Zool. 18(3), 1975

A NEW HAGFISH FROM SOUTH AUSTRALIA

None of those characters is constant. Large samples of a population of myxi-
noids usually include individuals with more or less than the model number of gills
or gill apertures, and the range of variation in the dental formula is so great that
there is considerable overlap between species. Moreover, these characters are not
readily accessible: the number of gills may be inferred from the number of
branchial apertures but these do not always correspond. In most preserved speci-
mens the lingual teeth are retracted within the buccal cavity and a deep incision
must be made to reveal them. The smaller teeth lie under a fold of buccal epi-
thelium and are easily overlooked. Clearly it would be advantageous if species of
myxinoids could be diagnosed by their external characters.

Studies of variation in the external characters of E. atami and E. burgeri
(Strahan and Honma, 1961; Strahan, 1962) have demonstrated that the position
of the first and last branchial apertures, cloaca, and ventral fin-fold; the depth of
the body; and the number and distribution of mucous glands provide a constella-
tion of individually variable characters which, taken together, are usually sufficient

to define an eptatretid species.

COMPARISON OF EPTATRETUS LONGIPINNIS WITH OTHER 5-7
GIELED EPTATRETIDS

The six-gilled hagfish, E. longipinnis does not fit the description of any of
the six-gilled species of Eptatretus (= Bdellostoma) listed by Holly (1933).
However, in view of known variation in the characters considered by Holly, it is
reasonable to establish that E. longipinnis is not a variant of a species with one
more or one less pair of gills. These are: E. yangi (Teng) 1958; E. springeri (Bige-
low and Schroeder) (1952); E. atami (Dean) 1904; E. profundus Barnard
1923; E. burgeri (Gunther) 1870; E. hexatrema (Muller) 1835; and E. cirrhatus
(Bloch and Schneider) 1801. |

Not all of these species have been described in sufficient detail to compare
them character by character. Where possible, therefore, specimens have been re-
examined and described in a standard manner. This has not been possible with
E. yangi or E. springeri but the relatively recent descriptions of these species
contain all the information necessary for the present purpose.

Eptatretus yangi
A description of the type and seven paratypes has been given by Teng (1958).
The following is a summary.

Length: 198-250 mm.

Major dimensions as percentage of length range mean) =e" S:D:
Rostrum to first branchial aperture 29 — 33 B23
Rostrum to last branchial aperture 31 — 35 3 So pegcamesel (0)
First to last branchial aperture 1— 2 see: TZ
Rostrum to origin ventral fin-fold 39 — §5 AT eri Aey
Rostrum to posterior border cloaca 86 — 88 S6nse w1e8
Maximum depth of body 5 — 8 6, 2E 7 OLF

Aust. Zool. 18(3), 1975 141

R. STRAHAN

Number of mucous glands _ range mean’ == Ss:
Prebranchial series 17 — 20 Sess oO
Branchial series nil
Abdominal series 35 — 40 BY ates ge)
Caudal series 7 — 10 Seu NG
Total 60 — 69 64) (a1 8

Lingual teeth: inner 2+6-8; outer 3-6-7.

Five pairs of branchial apertures in two irregular rows on the ventral surface.
Violet-brown to violet-grey, paler ventrally; white border to ventral fin and to
branchial apertures. South-western coast of Taiwan.

Eptatretus profundus

Only the holotype (SAM 13035) in the South African Museum is known.
The following data are based on a re-examination of this specimen.

Length: 620 mm.

Major dimensions mm % length
Rostrum to first branchial aperture 125 20
Rostrum to last branchial aperture 160 26
Rostrum to origin ventral fin-fold 290 45
Rostrum to posterior border cloaca 505 81
Maximum depth of body 5)5) 9

Number of mucous glands range mean
Prebranchial series 13 13
Branchial series 4 4
Abdominal series 50-51 St
Caudal series 15 15
Total 82-83 83

Lingual teeth: inner 248; outer 2+8.

Five pairs of ventrolateral branchial apertures arranged in regular rows.
Branchial mucous glands dorsal to branchial apertures. Dark brown dorsally and
ventrally. Atlantic Ocean, off Cape Pt., South Africa, 720 m depth.

Eptatretus springert

The summary below is based on the detailed description of the holotype and
two paratypes (Bigelow and Schroeder, 1952).

Length: 338-590 mm.

Major dimensions as percentage of length range mean + S.D.
Rostrum to first branchial aperture 23 — 24 23 ero
Rostrum to last branchial aperture 26 — 28 DT tea 1.0
First to last branchial aperture 2 — 6 4+ 1.4
Rostrum to origin ventral fin-fold Big a) AQAA
Rostrum to posterior border cloaca 82 — 87 Sie see.
Maximum depth of body 8 — 9 8 aE O'S

142 Aust. Zool. 18(3), 1975

A NEW HAGFISH FROM SOUTH AUSTRALIA

Number of mucous glands

Prebranchial series
Branchial series
Abdominal series
Caudal series
Total

Lingual teeth: inner 2+-9-10; outer 3+-10-11.

range
15 — 19
3 — 6
44 — 57
i1 — 14
77 — 92

It Ht + + + It

6

Ae Ue eR

AouaN TD

Six pairs of branchial apertures in two regular rows on the ventral surface.
Greyish-brown dorsally and ventrally; branchial apertures fringed with white.
Gulf of Mexico, 400-500 m; Atlantic Ocean off Florida, 800-900 m; Caribbean,

500 m. |

Eptatretus atami

Variation in the dimensions of 146 specimens has been described by Strahan
and Honma (1961) who diagnosed an eastern and a western form of the species.

The following is a summary.

Eptatretus atami (western form)

Length: 130-583 mm.

Major dimensions as percentage of length
Rostrum to first branchial aperture
Rostrum to last branchial aperture
First to last branchial aperture
Rostrum to origin ventral fin-fold
Rostrum to posterior border cloaca
Maximum depth of body

Number of mucous glands

Prebranchial series
Branchial series
Abdominal series
Caudal series
Total

Lingual teeth: inner 3+6-8; outer 3-5-9,

81
9
68

nil

mean

28
32

It Lt It It Lt H+

It

hues tinea

a

Six (sometimes 5) pairs of branchial apertures arranged in irregular (rarely
regular) rows on the ventral surface. Purplish-brown dorsally, grey ventrally.
Sea of Japan, coastal waters of north-western Honshu, Japan, 60-160 m depth.

Eptatretus atami (eastern fornz)

Length: 318-444 mm.
Major dimensions as percentage of length

Rostrum to first branchial aperture
Rostrum to last branchial aperture
First to last branchial aperture
Rostrum to origin ventral fin-fold
Rostrum to posterior border cloaca
Maximum depth of body

Aust. Zool. 18(3), 1975

range

— 28

143

R. STRAHAN

Number of mucous glands range
Prebranchial series 18 — 18
Branchial series
Abdominal series 44 — 47
Caudal series 8 — 9
Total 68) —— 72

Lingual teeth: inner 3+8-9. outer 348-9.

nil

mean + S.D.
17 div

+

46
9
71

Hel le

NO —

Six pairs of branchial apertures arranged in regular or irregular rows on
the ventral surface. Purplish-brown dorsally, grey ventrally. Pacific Ocean, coastal

waters of north-eastern Honshu, Japan, 50-490 m depth.

Eptatretus burgeri

The summary description below is based upon six specimens in the collection
of the Imperial University of Tokyo and six in the collection of the Enoshima

Aquarium, Japan.
Length: 235-435 mm.

Major dimensions as percentage of length range
Rostrum to first branchial aperture 26 — 30
Rostrum to last branchial aperture 32 — 35
First to last branchial aperture 4 — 6
Rostrum to origin ventral fin 46 — 54
Rostrum to posterior border cloaca 84 — 90
Maximum depth of body 7—8

Number of mucous glands
Prebranchial series 19 — 22
Branchial series 5
Abdominal series 45 — 50
Caudal series hie 4
Total 81 — 89

Lingual teeth: inner 2+ 6-7; outer 3+6-7.

29 Wak
34 + 1.0
a= 106
SO) rte 7220)
ees) a)
ESOS
PAS ate 0.9

5)
AS sks b3
12) 229038
Sdipzewul

Six (rarely 5 or 7) pairs of branchial apertures in regular ventrolateral rows.
Light brown to purplish-brown dorsally, paler ventrally; thin unpigmented mid-
dorsal stripe; conspicuous pale patch of skin (translucent in life) over eye; tips
of barbels pale. Southern Japan, southern Korea, 5-10 m depth.

Eptatretus hexatrema

The summary description below is based on six specimens in the British

Museum and eight in the South African Museum.

Length: 112-720 mm.

Major dimensions as percentage of length range
Rostrum to first branchial aperture 27 — 32
Rostrum to last branchial aperture 32 — 39
First to last branchial aperture 5 — 6
Rostrum to origin ventral fin 47 — 54
Rostrum to posterior border of cloaca 85 — 88
Maximum depth of body 6 — 7

144

mean + S.D.
30% 2E eT
SGN a2 129
6+ 0.4
Sliste 2246
86 EOI
GS x= OS

Aust. Zool. 18(3), 1975

A NEW HAGFISH FROM SOUTH AUSTRALIA

Number of mucous glands

Prebranchial series DE 77) 26; cel)
Branchial series 5 5

Abdominal series 50 — 60 S45 324
Caudal series ie | 0S) ee) OE
Total 91 — 105 98 + 4.9

Lingual teeth: inner 2+8-10; outer 3+ 8-10.

Six (rarely 5) pairs of branchial apertures opening ventrolaterally in regular
rows. Light brown to blackish-brown dorsally, paler ventrally. South Atlantic
Ocean off Cape of Good Hope, 10-45 m depth.

A

Fig. 2. Ventral view of heads of 5-7 gilled species of Eptatretus. (a) E. longi-
pinnis, n. sp., 422 mm long, holotype; (b) E. cirrhatus, 660 mm long;
(c) E. burgeri, 400 mm long; (d) £E. atami,; 350 mm long. (e) E.
springeri, 590 mm long (after Bigelow & Schroeder); (f) E. profundus,
620 mm long; (g) E. hexatrema, 285 mm long; (h) E. yangi, 212 mm
long (after Teng). Note that figures differ in scale: line below each
figure is equivalent to 10 mm in length.

Eptatretus cirrhatus

The following summary description is based upon three specimens in the
British Museum and ten from the National Museum, Wellington, New Zealand.

Dimensions as percentage of total length range mean + S.D.
Rostrum to first branchial aperture 21 — 26 2 eet 8
Rostrum to last branchial aperture 28 — 33 pl ee ah
First to last branchial aperture 6 — 9 eam
Rostrum to origin ventral fin 47 — 55 bY Aare eerie 5
Rostrum to posterior border cloaca 80 — 86 84 + 1.4
Maximum depth body 7—9 8.54se 0.3

Aust. Zool. 18(3), 1975 145

R. STRAHAN

Number of mucous glands range mean + S.D.
Prebranchial series 15 — 18 160S2e..0
Branchial series 6 — 7 G:Sics= TO
Abdominal series 49 — 54 les cn Ae
Caudal series 12 — 14 13. 327,120
Total 83 — 92 oY Minka 25

Lingual teeth: inner 3+6-9; outer 3+6-9.

Seven (rarely 6) pairs of branchial apertures opening ventrally. Blackish-
brown to brown dorsally, paler ventrally; prominent clear area over eye; white
borders to branchial apertures. South Pacific Ocean off both islands of New
Zealand, 8-10 m depth; off south-eastern coast of Australia, 300-700 m depth.

DISCUSSION

It is not difficult to separate E. longipinnis from the other species described
above. In four characters—the anterior position of the ventral fin, the posterior
position of the cloaca, the large number of abdominal mucous glands, and the
small number of lingual teeth—values for E. longipinnis fall outside the combined
range of the other species. In two characters, position of the ventral fin and
position of the cloaca, values for E. longipinnis fall outside three standard deviations
from the mean values for any of the other species. Indeed, in respect of the
characters under consideration, E. longipinnis differs more from each of the
other species or varieties than any two of them differ from each other.

KEY 210. THE -SPECIES OF EPTATRETUS, HAVING 5-7; PAIRS OF BRANCHEAL

APERTURES

la” Oren. of ventral fin-fold. anterior to third wesille aperture 7s ee se longipinnis
ib -Onem- ot ventral fin-told® posterior :to third: ell aperture: ee ee eee

2a First gill aperture to last gill aperture more than 6% of total length ........ cirrhatus
2b First gill aperture to last gill aperture less than 6% of total length. ........................ 3
BA PAlee -tidicl Gd ObSal- SEEUDC 3 62 eee ee Ve wea es ee Pe ae adn em ee burgeri
Sb. Mowmid-dorsal::.Sir wpe: 5 e305. IN oni een lt er OS cag oc a ae an di a Eee eae ee 4
43 essethan, 22 prebranchialiimuicous plands sts) Ao Se ae hexatrema
2b. More “than-22, prebranchial:mucous Slama sys ses sse), Ge ee aes Peseta 5
5a Rostrum to origin ventral fin less than 40% of total length ................ atami (western)
5b. Rostrum to origin ventral fin more than 40% of total length .........0000000 6
Gas Bianca mucous. lands: absent ose ee ee ek aw cine eaten Oe ee Al a ¥
ab: - Branchial. mucous: lands: present) 2.3 Sse ee a ie ae han eh enamel 8
7a First gill aperture to last gill aperture more than 3% of total length ..... atami (eastern)
7b First gill aperture to last gill aperture less than 3% of total length ................ yangi
oa --branchial mucous. glands dorsal to sill: apertures 2] Sh oe ene. profitndus
$b. Branchial. mucous: slands: ventral*ite pill’ apertures: 1 eee ee springeri

ACKNOWLEDGEMENTS

Thanks are due to the South Australian Museum; the South African Museum; the
British Museum; the National Museum of New Zealand; the Imperial University, Tokyo,
and its Marine Research Station, Misaki. Niigata University; the Enoshima Aquarium, and
the N.S.W. State Fisheries, for providing access to their collections.

146 Aust. Zool. 18(3), 1975

A NEW HAGFISH FROM SOUTH AUSTRALIA

ADDENDUM

While this paper was in press, a second specimen, 443 mm long, from
South Australia “off Cape Douglas” (South Australian Museum No. F3611)
came to hand. In this specimen the branchial region is slightly longer (1% total
length) than in the type, and is slightly asymmetrical, the anteriormost branchial
aperture on the right side being anterior (0.2% total length) to that on the left.
The origin of the ventral fin-fold is at the level of the latter. Otherwise, this
specimen corresponds well with the type.

Major dimensions mm % length
Rostrum to first branchial aperture 133-134 30
Rostrum to last branchial aperture 154 35
Rostrum to origin ventral fin-fold 134 30
Rostrum to posterior border cloaca 410 93
Maximum depth of body 21 5

Mucous glands range mean
Prebranchial series 27-28 28
Branchial series 5-6 6
Abdominal series 60-63 62

~ Caudal series 9 9
Total 102-105 104

Lingual teeth: inner 2+5; outer 3-5-6.

REFERENCES
ADAM, H. & R. STRAHAN (1963). Systematics and geographical distribution of myxinoids.
In ‘The Biology of Myxine’, (Ed. A. Brodal & R. Fange) Universitetsforlaget, Oslo, 1-8.

APSTEIN, C. (1915). Nomina conservanda. Sitzber. Ges. naturf. Freunde Berl., No. 5,
119-202.

BARNARD, K. H. (1923). Diagnoses of marine fishes from South African waters. Ann.
S. Afr. Mus. 13, 439-445.

BIGELOW, H. B. & W. C. SCHROEDER (1948). Cyclostomes. In ‘Fishes of the Western
North Atlantic No. 1, Pt. 1, Sears Found. Mar, Res., New Haven, 29-58.

BIGELOW, H. B. & W. C. SCHROEDER (1952). A new species of the cyclostome genus
Paramyxine from the Gulf of Mexico. Brev. Mus. comp. Zool. Harvard No. 8, 1-10.

DEAN, B. (1904). Notes on Japanese myxinoids, a new genus Paramyxine and a new
species Homea okinoseana, reference also to their eggs. J. Coll. Sci. Univ. Tokyo
1D. te 3

HOLLY, M. (1933). Cyclostomata. In ‘Das Tierreich’ Lief. 59 (Ed. F. E. Schultze &
W. Kukenthal) de Gruyter, Berlin & Leipzig, 1-62.

HUBBS, C. L. (1963). Cyclostomata. In ‘Encyclopaedia Britannica’ 6, 941-944.

MATSUBARA, K. (1937). Studies on the deep-sea fishes of Japan, III. On some remarkable
variations found in Paramyxine atami Dean, with special reference to its taxonomy.
J-pamp, Fishes Inst.: Tokyo. 32, 13-15.

RAUTHER, M. (1924). Ciclostomi. In ‘Bronn’s Klassen und Ordnungen des Tier-Reichs’
6, Abt. 1, Buch 1, Lief 39, Akad. Verlag. Leipzig, 250-703.

REGAN, C. T. (1912). A synopsis of the myxinoids of the genus Heptatretus or Bdello-
stoma. Ann, Mag. nat. Hist. (8) 7, 534-536.

Aust. Zool. 18(3), 1975 147

R. STRAHAN

STRAHAN, R. (1962). Variation in Eptatretus burgeri, with a further description of the
species. Copeia, 801-807.

STRAHAN, R. & Y. HONMA (1961). Variation in Paramyxine, with a redescription of
P. springeri Bigelow and Schroeder. Bull, Mus. comp. Zool. Harvard 125, 323-342.

TENG, F. T. (1958). A new species of cyclostome from Taiwan. Chin. aquat. Prod.
(Chin. Fish.) No. 66, 3-6. (In Chinese).

148 Aust. Zool. 18(3), 1975

A Taxonomic Revision of the Litoria aurea Complex
(Anura: Hylidae) in South-Eastern Australia

GILLIAN P. COURTICE and GORDON C. GRIGG
School of Biological Sciences, University of Sydney, N.S.W., 2006

ABSTRACT

Four subspecies of Litoria aurea have been previously recognised in south-
eastern Australia. In the present paper it is proposed that two of these subspecies
(Litoria aurea aurea and Litoria aurea raniformis) be raised to specific status as
Litoria aurea and Litoria raniformis respectively; that an isolated population
from the New England region of N.S.W. be recognised as a new species Litoria
flavipunctata; and that the remaining two subspecies, Litoria aurea ulongae and
Litoria aurea major be synonymised with Litoria aurea and Litoria raniformis
respectively.

This revision in taxonomy is proposed on the basis of morphological evidence
and the apparent lack of hybridisation in areas of sympatry.

INTRODUCTION

Frogs of the Litoria aurea complex, representing a number of taxa, are
extensively distributed in the south-east of Australia, including Tasmania, and in
the south-west of Western Australia.

The group was first described as a single species, Rava aurea (Lesson 1831),
the type locality being the Macquarie River at Bathurst, N.S.W. Giinther (1858)
placed the species in the genus Hyla. Keferstein (1867) described Chirodryas
raniformis from a specimen of unknown locality, but later (1868) sank this
species into the synonymy of Hyla aurea. Parker (1938) recognised two sub-
species of Hyla aurea in eastern Australia, the N.S.W. coastal form, Hyla aurea
aurea and the inland and southern form, Hyla raniformis. In Western
Australia, Parker recognized Hyla aurea raniformis and raised a further sub-
species, Hyla aurea cyclorhynchus (Boulenger 1882) to specific status. Copland
(1957) described the raniformis type from Western Australia as a new species,
Hyla mooret. This was based on cross-fertilisation experiments reported by Moore
(1954). Hyla aurea ulongae (Loveridge 1950) was described from Ulong, N.S.W.
Yet another subspecies was erected when Copland (1957) re-named the Tas-
manian group, previously ascribed to Hyla aurea raniformis, Hyla aurea major.
Subsequently, Tyler (1971) has proposed that all Australo-Papuan species of
Hyla be attributed to the genus Litoria Tschudi (1839) because of differences in
vocal sac musculature compared with hylids from elsewhere.

Aust. Zool. 18(3), 1975 149

GILLIAN P. COURTICE and GORDON C. GRIGG

Thus, two Western Australian species of the Litoria aurea complex are
recognised, Litoria cyclorhynchus (Parker) and Litoria moorei (Copland). Until
now, four subspecies of Litoria aurea have been recognized in south-eastern
Australia, as follows:

Litoria aurea aurea (Lesson), a coastal form in N.S.W. and eastern Victoria,

Litoria aurea raniformis (Keferstein), distributed in southern inland N.S.W.,
Victoria and through the Murray R. Valley into South Australia,

Litoria aurea ulongae (Loveridge), from near Dorrigo, N.s.W.,

Litoria aurea major (Copland), from Tasmania.

There has been uncertainty of the taxonomic status of these south-eastern
forms. Moore (1961) and Littlejohn, Martin & Rawlinson (1963) have suggested
that the first two taxa may deserve full specific status. Litoria aurea ulongae was
described from a single preserved specimen (Aust. Mus. No. R13817) (Loveridge
1950). Litoria aurea major was differentiated from Litoria aurea raniformis only
on the basis of size (Copland 1957).

The present taxonomic revision is based on morphological evidence and
apparent lack of hybridisation in zones of sympatry. We propose:

(i) that Litoria aurea aurea and Litoria aurea raniformis be raised to full
specific status as Litoria aurea and Litoria raniformis,

(ii) that Litoria aurea ulongae and Litoria aurea major be synonymised
with Litoria aurea and Litoria raniformis respectively; and

(iii) that the geographically isolated population of Litoria aurea raniformis
on the New England tableland be recognised as a new species, Litoria
flavipunctata.

MATERIAL AND METHODS

Data on morphology and geographic distribution were obtained from our
collections made throughout Victoria and New South Wales, and from preserved
material in the Australian Museum. In addition, literature records were taken
into consideration. Traverses were made near Canberra, Australian Capital Terri-
tory, and in East Gippsland, Victoria, to investigate the relationship between
Litoria aurea aurea and Litoria aurea raniformis in sympatry.

In morphological studies, the following measurements were taken using
vernier calipers (Peacock); snout-vent length (SV), tibia length (TL), head
length (HL), and head width (HW) as defined by Goin and Netting (1940);
foot length (FL) measured from outside heel to tip of toe; distance between
eye and naris (EN), measured along the canthus rostralis; internarial span, (IN),
diameter of the tympanum (T) and diameter of the eye (E). Using an ocular grid
in a binocular microscope, the width of the third digit of the hand (DW) at the
narrowest part between the digital pad and the next joint, and the width of the
third digital pad (PW) were measured.

150 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

Fig. 1. View of foot and hand of Litoria aurea (top), Litoria raniformis (centre) and
Litoria flavipunctata (bottom).

Aust. Zool. 18(3), 1975 151

GILLIAN P. COURTICE and GORDON C. GRIGG

LITOAIA
AVREA
LITORIA
RANI FORMS
Y LITORIA
fy, FLAVIPUNCTATA

Fig. 2. Map of south-eastern Australia showing distribution of the Litoria aurea complex.
The distribution of L. raniformis in Tasmania is based on Littlejohn (1967).

152 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

COMPARISONS BETWEEN TAXA

To avoid confusion in the ensuing discussion, we will use the proposed names
Litoria aurea, Litoria raniformis and Litoria flavipunctata.

(a) Morphology

(i) Dorsal stripe. Litoria raniformis and Litoria flavipunctata have an
obvious mid-dorsal stripe. This is absent in Litoria aurea (Plate 1).

(ii) Digital pads (Fig. 1). Litoria raniformis and Litoria flavipunctata have
digital pads the same widths as the digits, whereas the pads of Litoria aurea are
wider than the digits.

(iii) Colouration. Litoria flavipunctata is characterised by having large
yellow spots in the groin and on the ventral surfaces of the legs. Smaller yellow
spots are usually evident on the posterior surface of the thigh. Individuals of
Litoria raniformis occasionally have a few small yellow spots on the posterior
surface of the thigh. Litoria aurea never has yellow spots in these regions. In
preservative, these spots are white.

(iv) Webbing (Fig. 1). The extent of foot webbing is consistently different
in each of the three species. The fullest web is seen in Litoria flavipunctata where
webbing extends to the tip of each toe. This webbing is usually covered with
large yellow spots as found on the ventral surfaces of the hind limb. In Litoria
raniformis the webbing is somewhat reduced, whereas in Litoria aurea the foot
is only one half to three quarters webbed.

(b) Zones of Sympatry
The population on the New England tableland (Litoria flavipunctata) is
geographically isolated from related taxa, so no interactions can occur.

Between Litoria aurea and Litoria raniformis however, two limited areas of
sympatry are known. These areas are near Canberra, A.C.T. and on the Gippsland
coast of Victoria near Orbost (Fig. 2). In both these areas, specimens of each
species were collected, often in the same pond and even within less than a metre
of each other. Using the morphological criteria given earlier, each species was
clearly recognisable and no evidence of hybridisation was apparent. Littlejohn,
Martin and Rawlinson (1963), from collections in East Gippsland between
Metung and Orbost, noted that both types occurred in sympatry. without
hybridisation

CAPTION TO COLOUR PLATE

Plate I: A. Litoria aurea.
B. Litoria raniformis.
C_ Litoria flavipunctata.
D. Ventral views of Litoria aurea (left), Litoria raniformis (centre) and Litoria
flavipunctata (right).

Aust. Zool. 18(3), 1975 153

GILLIAN P. COURTICE and GORDON C. GRIGG

Of another collection made in East Gippsland, Littlejohn (1969) writes:
“specimens may be assigned to the subspecies H. aurea aurea with the exception
of a series from Orbost in which there are indications of raniformis characteristics”.
The raniformis type is common to the east of 148°E, well within East Gippsland
as defined by Littlejohn (1969) and yet no further mention of it appears in his
checklist. Therefore we consider his statement indicative of his uncertainty of
the taxonomic relationship between these taxa rather a clear implication that
hybridisation occurs near Orbost. Moore (1961) refers to specimens that he
considered to be intermediates between the two taxa. We have examined some
of these specimens in the Australian Museum (R.9482-9484 from Koonadon near
Leeton, R.7344 from Emu Plains near Urana, R. 5233 from Berridale, Monaro
and the series R.5959-5962, R.6683 from Narracoorte, S.A., two of which
Moore suggested may be intergrades). By our criteria these are all specimens of
Litoria raniformis. Furthermore, these four localities are far distant from any
known area of sympatry so that occurrence of intergrades at these localities is
unlikely. Another specimen mentioned by Moore (1961) as a possible intergrade
is in the British Museum (No. 96.6.17.17). It was collected from the Yarra
River, Victoria. Unfortunately we have been unable to examine it.

In our view, the evidence suggests that Litoria aurea and Litoria raniformis
maintain their distinctiveness in sympatry. The isolating mechanisms which operate
to maintain this situation are, at present, unknown.

(c) Litoria aurea ulongae

This subspecies was described by Loveridge (1950) from a single preserved
specimen from Ulong, N.S.W. No other specimen is known. Loveridge based its
validity as a separate taxon on the lack of a dorso-lateral fold and the presence
of an undivided ridge of vomerine teeth. Collections made in the area by one of us
(G.P.C.) revealed no further specimens. The type specimen in the Australian
Museum (R.13817) was examined carefully and it appears to be a somewhat
unusual specimen of Litoria aurea. Hence we consider that Litoria aurea ulongae
should be placed in the synonymy of Litoria aurea.

(d) Litoria aurea major

Copland (1957) ascribed the Tasmanian frogs of the “raniformis’” type to a
new subspecies, Litoria aurea major, solely on the basis of their supposed larger
size. He wrote that “In large series of raniformis 60 mm was only exceeded in
about 6% of specimens, and 70 mm only once (about 1%), and then only by
1 mm (71 mm). It must be noticed, however, that Keferstein gave the body length
of his type of raniformis at 82 mm, which can only be regarded as very excep-
tional.” (Copland 1957, p. 80.) Moore (1961) questioned the validity of this
sub-species. He reported that several raniformis individuals which he had collected
exceeded 72 mm, the length of the largest major referred to by Copland (1957).

In the present study, 66% of those Litoria raniformis measured exceeded
60 mm (compare 6% recorded by Copland, 1957) and one female caught at Nar-

154 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

randera, N.S.W. was 90 mm in length. It would seem that no basis exists for retain-
ing the Tasmanian frogs in a separate taxon and we propose that Litoria aurea
major should be placed in the synonymy of Litoria raniformis.

SYSTEMATICS

The observations in the previous section support the validity of recognising
three separate species within the Litoria aurea complex in south eastern Australia.
Full descriptions of the three species will now be given.

Litoria aurea (Lesson 1831)

Synonymy:
Rana aurea Lesson 1831 (1830);
Ranoidea jacksoniensis Tschudi, 1839;
Ranoidea resplendens Girard, 1853;
Hyla jacksonii Bibron & Dumeril, 1854;
Hyla castanea Steindachner, 1867 (1869);
Fanchonia elegans Werner, 1893;
Hyla blandsuttoni Proctor, 1924.
Hyla aurea ulongae Loveridge, 1950;
Hyla aurea ulongensis Loveridge, 1950;
Hyla aurea major Copland, 1957.

Type Locality: Macquarie River, Bathurst, N.S.W.

Diagnosis: This species can be recognised in life by the following characteristics:
smooth green band with irregular gold spots, yellow dorsolateral fold and a
brilliant turquoise colour in the groin and on the back of the thigh.

In preservative the green and gold colours of the dorsum become
smoky grey and light brown respectively. The dorsolateral fold becomes
white. The groin colouring is grey.

Description of a female collected from Ourimbah, N.S.W. in October 1972.
Australian Museum No. R42156.

The dorsal surface is smooth and green with irregular gold spots. A yellow
dorsolateral fold extends from the eye to the groin. Another short yellow line
extends from under the eye to above the shoulder.

The belly and ventral surfaces of the thighs are white and coarsely granu-
late. The skin is more finely granulate under the forelimbs and the throat. The
stoin, the posterior surface of the thigh and the ventral surfaces of the tibia are
coloured brilliant turquoise.

The head is obtusely pointed and slightly longer than broad (HL/HW ratio
1.15). The head length is approximately 1/3 of the snout to vent length (HL/SV

Aust. Zooi. 18(3), 1975 155

GILLIAN P. COURTICE and GORDON C. GRIGG

ratio 0.34). The snout is not prominent and does not project conspicuously
beyond the anterior limit of the mandible. The distance between the eye and
the naris is greater than the internarial span. The canthus rostralis is distinct
and straight; the loreal region is concave.

The tympanum is distinct and gold in colour. Its diameter is 0.72 x that
of the eye. The vomerine elevations are large, paired and obliquely convergent
posteriorly; they lie between the choanae. The tongue is broad, free and nicked
posteriorly.

Fingers are long and free from webbing. They are, in order of length,
3 > 4 > 2 = 1. Terminal discs are wider than the digits.

The hind limbs are long and slender (TL/SV ratio 0.50). Toes are, in order
of length, 4 > 3 = 5 > 2 > 1. They are approximately 3/4 webbed (see Fig. 1).

In preservative, the dorsal surface is a greyish green and the ventral surfaces
and the yellow lines become immaculate white.

Dimensions

SV, 76:91m: VL 38.7 mm: BL. 55) mm ls 26. 9--nams), FAW 2320) paar
IN 4.2 mm; E 8.7 mm; T 6.3 mm.

Variability

The snout to vent distances of 24 frogs of this species had a mean of 58.3 mm
and a range of 46.9-76.8 mm. The amount of gold colouring on the dorsal surface
varied greatly; this variation seemed to be unrelated to any geographic pattern.

Specimens examined

Australian Museum, N.S.W.; R18484, Wollongong, N.S.W.; R19424-7,
(1890) Richmond, N.S.W.; R19643-5, (1890) Manly, N.S.W.; R4194-6, 4198-
203, 4205; (1908) “Maroubra, INSAW + -R4452. (1909) Sans“ Souck) NS We
R4665-8, (1910) Woonona, N.S.W.; R3388-9, (1911) North Sydney, N.S.W.;
R5849°°( 1912) Cooks, River, N'S.W.; R7325,; (1921) Capertreey N:'SAWis R76
(1921) Burrawang, N.S.W.; R7446-1, (1922) Pambula, N.S.W.; R7974, (1922)
Upper Colo, N.S.W.; R8454, 8456, 8483, (1924) Sydney, NS.W.; R9424,
R9426-7, (1928) Wentworthville, N.S.W.; R9532, (1928) Botany, N.S.W.;
R9558, (1928) Five Dock, N.S.W.; R9644, Tumut, N.S.W.; R18768, (1957)
Shoalhaven Heads, N.S.W.; R19669-81, (1958) Singleton, N.S.W.; R2455, (1965)
Londonderry, N.S.W.; R27635, (1968) Cobargo, N.S.W.; R29446, (1972) Lake
Lidell, N.S.W.; R29441-3, (1972) French’s Forest, N.S.W.

Author’s collections

Goulburn, N.S.W. (4 specimens); Pearl Beach, N.S.W. (50 spec.); Tomerong,
N.S.W. (3 spec.); nr. Jacqua, N.S.W. (2 spec.); Liddell Lake, N.S.W. (4 spec.);
Cann River, Vic. (4 spec.); 12 m E. of Orbost, Vic. (10 spec.); 1 m E. of Orbost,

156 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

Vic. (4 spec.); 5 m SE of Orbost, Vic. (2 spec.); Marlo, Vic. (6 spec.); Foxlow,
N.S.W. (1 spec.); near Maclean, N.S.W. (2 spec.); Bungonia, N.S.W. (20 spec.);
12 m NE Canberra, N.S.W. (4 spec.); 12 m NE Tarago, N.S.W. (2 spec.).

Literature records
These records do not include material in the Australian Museum which has
been examined by the authors.

Lower Clarence River, N.S.W.; Gurravembi, Nambucca River, N.S.W.;
Warrell Creek, Nambucca River, N.S.W.; Nambucca River, N.S.W.; Botany Bay,
N.S.W.; Bulahdelah, N.S.W.; Sydney, N.S.W.; East Lakes, Sydney, N.S.W.;
Burradoo, N.S.W.; Bodalla, N.S.W.; Eden, N.S.W.; Nowra, N.S.W.; (Moore
1961). Jervis Bay, N.S.W.; (Fletcher 1894, cited in Moore 1961). Razorback
Mt., near Picton, N.S.W.; Bundanoon, N.S.W.; Tallong, N.S.W.; Fitzroy Falls,
N.S.W.; Barrengarry Mt., N.S.W.; Waverley, N.S.W.; (Copland 1957). Thirroul,
N.S.W.; (Harrison 1922). Gipsy Point, Vic.; Orbost, Vic.; Corringle Beach, Vic.;
6 miles N of Lake Entrance, Vic.; Metung, Vic.; Genoa, Vic.; Nowa Nowa, Vic.;
(Littlejohn et al. 1963). Goongerah, Vic.; Fairhaven, Mallacoota Inlet, Vic.;
1 ml N of Marlo, Vic.; 11 miles E of Orbost, Vic.; 8 miles E of Bendoc, Vic.;
(Littlejohn 1969).

Biology :

This species inhabits permanent ponds, farm dams and the still backwaters of
rivers. Their habitat usually can be identified by the presence of beds of tall reeds.
The frogs can be found on these reeds at night, feeding. They call in open water,
and eggs are laid on top of the water but later sink (Littlejohn 1971).

Larvae

The tadpoles are large and dark. The anus is dextral; the spiracle sinistral;
eyes lateral. The dorsal fin extends nearly halfway up the back. The mouth disc
has two rows of upper labial teeth, and three rows in the lower labium. The inner
row in each labium is divided. A narrow row of papillae extend around the lateral
and posterior margins of the mouth.

Litoria raniformis (Keferstein 1867).

Synonymy:
Chirodryas raniformis Keferstein 1867.
Hyla aurea major Copland 1957.

Diagnosis: This species can be recognised by its green and gold warty back with a
green middorsal stripe. The groin and back of thigh are coloured blue, slightly
less brilliant than that of Litoria aurea. There are sometimes small yellow
spots on the back of the thigh.

Aust. Zool. 18(S), 1975 157

GILLIAN P. COURTICE and GORDON C. GRIGG

In preservative the mid-dorsal stripe and the dorsolateral folds stand
out as pale grey markings against a darker grey back, on which longitudinal
rows of darker warts are seen.

Description of a male collected from Narrandera, N.S.W., August, 1972. Australian
Museum No. R42155.

The dorsal surface is green and brown with rows of black spots and a
distinct green mid-dorsal stripe extending from between the eyes to the cloaca.
There is a dorsolateral gold line which starts on the eye and fades out just before
the groin. There is another line from below the tympanum to the shoulder.

The belly and most of the ventral surfaces of the thighs are coarsely
granulated. The region of the vocal sac has a yellow tinge. The groin, the
posterior surface of the thigh and the inner surface of the tibia and foot are
blue in colour.

The head is obtusely pointed and approximately as long as it is broad (HL/
HW ratio 1.02). The head approximately 1/3 of the snout to vent length (HL/
SV ratio 0.37). The snout is not prominent and does not project conspicuously
beyond the anterior limit of the mandible. The distance between the eye and
naris is greater than the internarial span. The canthus rostralis is distinct and
straight, the loreal region is concave.

The tympanum is distinct and gold in colour. Its diameter is 0.59 x that of
the eye.

The vomerine elevations are paired, and are slightly convergent posteriorly.
The anterior ends are directly between the choanae. The tongue is broad, free
and indented posteriorly.

Fingers are long and free from web. Fingers are, in order of length
3 > 4 > 2 = 1. Terminal discs are approximately the same width as the digit.

The hind limbs are long and robust (TL/SV ratio 0.50). Toes are, in order
of length, 4 > 3 = 5 > 2 > 1. They are extensively webbed (see Fig. 1).

In preservative the dorsal surfaces become a dirty green or brown and the
ventral surfaces and the dorsolateral lines become dirty white.

Dimensions

SV 603mm: Th 30:3 ams EL +42°8) mm: Hls..22.5. mm: AW <22.0iemane,
EN 4.5 mm; IN 3.2 mm; E 7.8 mm; T 4.6 mm.

Variability
Members of this species usually have a green dorsum with brown and black
warts arranged in longitudinal rows. Some individuals have a few small yellow

spots on the posterior surfaces of the thighs. Males have a yellow tinge in the
region of the vocal sac.

158 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

Specimens examined

Australian Museum: R3121-3, (1900) Fish River, Tarana, N.S.W.; R5233,
(1911) Berridale, N.S.W.: R7344, (1921) Emu Plains, Utana, N.S.W.; R8421,
(1924) Tumut, N.S.W.; R10661-8, (1932) Yanco, N.S.W.; R10669-88, (1932)
Tubbo Stn., Riverina, N.S.W.; R25852-55, (1966) Bombala, N.S.W.; R26181,
(1966) Jumparcumbay NSW. R27G05, (1967) Mt.. Beauty, Vic. R5999-62.
R6683 (1912) Narracoorte, S.A.

Author’s collection

Bywong, N.S.W. (1 specimen); Boorowa, N.S.W. (2 spec.); Narrandera,
INS OW (GID spec.): ~Morundah, N:S.W: (1 spec.); Wagea, N.S.Wi(S> spec.
Delegate, N.S.W. (2 spec.); 12 m NE Canberra, N.S.W. (15 spec.); near Bun-
sendore, N.S.W. (2 spec.); Booligal, N.S.W. (10 spec.); Saucy Ck., Bombala,
N.S.W. (2 spec.); Maclaughlin R. nr. Nimmitabel, N.S.W. (4 spec.); Swan Hill,
Vic. (2 spec.); Whittlesea, Vic. (3 spec.); 10 m SE Orbost, Vic. (8 spec.).

Literature records

These do not include specimens in the Australian Museum already examined
by the authors.

Albury, N.S.W.; Delegate, N.S.W.; Canberra, A.C.T.; Gunbower, Vic.;
Melbourne, Vic.; Healesville, Vic.; Yarra River, Vic.; Millgrove, Vic. (Moore
1961). Tocumwal, N.S.W.; Gol Gol, N.S.W.; Darlington Point, N.S.W.; Bal-
ranald, N.S.W.; 10 m W of Narrandera, N.S.W.; 8 m S of Bombala, N.S.W.;
Adaminaby, N.S.W.; Bacchus Marsh, Vic.; 13 m SE of Horsham, Vic.; Cobungra,
Vic.; Macedon, Vic.; 18 m W of Bairnsdale, Vic.; Brighton, Vic.; Mordialloc,
Nie.) Vailem bend, tou. (Copland 1957): dock No. -9, Murray R:;’ 3 mE ‘of
Mildura, Vic.; Lake Cullulleraine, Vic.; 6 m SE of Red Cliffs, Vic.; Carwarp, Vic.;
Nangiloc, Vic.; Boundary Bend area, Vic.; Nyah, Vic.; Sea Lake, Vic.; Wyche-
proof, Vic.; 16 m W of Nhill, Vic.; Kaniva, Vic.; Mildura, Vic.; Lake Boga, Vic.;
5 m NW of Dimboola, Vic.; Wail, Vic.; 15 miles W of Kaniva, Vic. (Littlejohn
1966). Flinders Is., Bass Strait; King Island, Bass Strait (Littlejohn and Martin
1965).

Biology :

These frogs occupy a’ similar habitat to Litoria aurea. However, they are
often found during the day on a grassy bank near water, basking in the sun. This
habit is not usually observed in L. aurea.

Larvae
Martin (1965) has described the tadpole of this species.

Litoria flavipunctata sp. nov.

Synonymy:
Hyla aurea raniformis Moore 1961 partim.

Aust. Zool. 18(3), 1975 159

GILLIAN P. COURTICE and GORDON C. GRIGG

Holotype

A male, No. R40676 in the Australian Museum, from a swamp on the
Booralong Creek Road, 12.8 km west of Guyra, N.S.W. (30° 16'S. 151° 33’E).
(Paratypes Aust. Mus. R40677-82).

Derivation of name

The name flavipunctata is from the Latin meaning “yellow spots’’. This refers
to the large yellow spots found in the groin and on the ventral surfaces of the
hind limb of these animals. The presence of these spots is a distinguishing feature
of this species.

Diagnosis: This species can be recognised by its dorsal colouration: green with
small brown and black spots and a green mid-dorsal stripe, together with
large and conspicuous yellow spots in the groin and on the ventral surfaces
of the hind limb. These large yellow spots are not found in Litoria aurea or
Litoria raniformis.

In preservative the dorsum appears very similar to that of preserved
Litoria raniformis with a slightly less obvious mid-dorsal stripe. Spots in
the groin and on the legs are white.

Description of the type specimen

The dorsal surface is green with brown and black spots, giving it a warty
appearance. There is a distinct green mid-dorsal stripe extending from between
the eyes to the cloaca.

The belly and part of the ventral surface of the thigh are white and coarsely
granulate. The skin is more finely granulate under the forelimbs and the throat,
and there is a hint of pale yellow in the region of the vocal sac. In the groin, on
the ventral surface of the thigh and the ventral surface of the tibia and the foot,
there are very large, distinct, yellow spots. Smaller yellow spots are found on the
posterior surfaces of the thighs.

A gold line extends dorsolaterally from behind the eye to behind the shoulder,
where it continues as a series of gold warts almost to the groin. Another short
yellow line extends from under the tympanum to above the shoulder.

The head is obtusely pointed and slightly longer than broad (HL/HW ratio
1.15). The head length is approximately 1/3 of the snout to vent length (HL/SV
ratio 0.27). The snout is not prominent and does not project conspicuously beyond
the anterior limit of the mandible. The distance between the eye and the naris is
greater than the internarial span (IN/EN ratio 0.70). The canthus rostralis is
distinct and straight; the loreal region is concave.

The tympanum is distinct, gold in colour with a green centre. Its diameter
is 0.6 x that of the eye. The vomerine elevations are paired, obliquely convergent

160 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

posteriorly. The anterior ends are directly between the choanae. The tongue is
broad, free and nicked posteriorly.

The fingers are long and free from web. Fingers are, in order of length,
3 > 4 > 2 = 1. Terminal discs are approximately the same width as the
digit.

The hind limbs are long and robust (TL/SV ratio 0.53). Toes are, in order
of length, 4 > 3 = 5 > 2 > 1. They are fully webbed (see Fig. 1).

In preservative the dorsum becomes a dark grey colour with a lighter grey
stripe. The large yellow spots become white.

Dimensions
oy 6473) mms [34.1 ein Gis mins AL 23-9 mm: HW 20s/imim:
BINS > mim:- LN 377mm: EF 6:9 nm: 14.1 mm,

Variation

There is a variable amount of gold on the dorsal surfaces of these frogs, but
the dorsal stripe remains distinct at all times. The ventral yellow spots are some-
times outlined in black. The size of the digital pad in relation to the digit on
the hand and the amount of foot webbing were the same in every frog examined.
A series of 21 adult specimens had a range of SV lengths from 52.6-84.8 mm.

Biology

This species occupies a similar habitat to Litoria aurea and Litoria raniformis.
It is found also in ponds or slow moving streams with overhanging grassy banks
in the absence of reed beds.

It has been found to overwinter in the hollow centres of rotting logs and
in the earth surrounding the roots of uprooted trees.

Specimens examined

Australian Museum, N.S.W. R32183-9, (1971) Guyra, N.S.W.; R32560,
(1971) Llangothlin Lagoon, N.S.W.; R32547, (1971) Llangothlin Lagoon;
R34814-7; (1972) Black Mt. Lagoon, nr Guyra, N.S.W.; R33816, (1972) Sattern
Gully, nr Armidale, N.S.W.; R34937, (1972) Abby Green Stn., nr Guyra, N.S.W.

Author’s collection

Black Mt., near Guyra, N.S.W.; Llangothlin Lagoon, N.S.W.; 4 m S of
Guyra, N.S.W.; Booralong Ck., New England, N.S.W.

Literature records
Booralong Ck., N.S.W. (Moore, 1961), recorded as Hyla aurea raniformis.
To our knowledge, this is the only reference in the literature which specifically
refers to the individuals on the New England tableland.

Aust. Zool. 18(3), 1975 161

GILLIAN P. COURTICE and GORDON C. GRIGG
KEY TO SOUTH-EASTERN SPECIES

1 (a) Mid dorsal stripe present

Digital “pads: same width. as aieits ee ey ee ae ee eee ae 2
(b) Mid dorsal stripe absent

Disital-pads 412° times: width of “digits... 8 See ee ee ees Litoria aurea

2 (a) Large yellow spots (white in preservative) in groin and ventral surfaces of

legs

Hoes completely webbed A ee ee ee eee ees Litoria flavipunctata
(b) No large yellow spots covering groin and ventral surfaces of legs

does Jess: than’completely webbed) ) 53 eee Litoria raniformis

ACKNOWLEDGEMENTS

Financial assistance from a University of Sydney Research Grant is gratefully acknow-
ledged. In addition we wish to express our gratitude to Dr. Harold Cogger at the
Australian Museum for his advice and assistance throughout and for reading and commenting
on the manuscript. The problem crystallised in its early stages from discussions with Mr. John
Barker, and thanks are due to Miss Christine Lehmann for her enthusiasm and assistance
in the fieldwork. The artwork is by David Stanley.

REFERENCES

BIBRON, G. & A. M. C. DUMERIL (1854). Erpétologie générale ou Histoire naturelle
complete des Reptiles. Paris. 9, 409.

BOULENGER, G. A. (1882). Catalogue of the Batrachia salientia s. Ecaudata in the
collection of the British Museum. 2nd ed. London.

COPLAND, S. J. (1957). Presidential address. Australian tree frogs of the genus Ayla.
Proc. Linn. Soc. N.S.W. 82, 9-108.

GIRARD, C. (1853). Descriptions of new species of reptiles collected by the U.S. Exploring
Expedition, under the command of Capt. Charles Wilkes, U.S.N. Second Part. Including
the species of batrachians exotic to North America. Proc. Acad. nat. Sci. Philad. 6,
420-424.

GOIN, C. J. & M. G. NETTING (1940). A new gopher frog from the gulf coast,
with comments upon the Rana areolata group. Ann. Carneg. Mus. 28, 137-168.

GUNTHER, A. (1858). Catalogue of the Batrachia salientia in the collection of the
British Museum, London xvi + 160 pp.

HARRISON, L. (1922). On the breeding habits of some Australian frogs. Aust. Zool. 3,
17-34.

KEFERSTEIN, W. (1867). Ueber einige neue oder seltene Batrachier aus Australien und
dem tropischen Amerika. Nachr. Ges. Wiss. Gottingen 18, 341-361.

KEFERSTEIN, W. (1868). Ueber die Batrachier Australiens. Archiv. fur Naturg. Berlin
34 (1): 253-290.

LESSON, R. P. (1831 (1830)). Zoologie in Duperrey’s ‘Voyage autour du monde, sur la
corvette de la majeste, La coquille’. 2, 59-61.

LITTLEJOHN, M. J. (1966). Amphibians of the Victorian mallee. Proc. R. Soc. Vict. 79
(2), 597-604. Thee

162 Aust. Zool. 18(3), 1975

TAXONOMY OF LITORIA AUREA COMPLEX

LITTLEJOHN, M. J. (1967). Patterns of zoogeography and speciation in south eastern
Australian Amphibia. In ‘Australian Inland Waters and Their Fauna’. (Ed. A. H.
Weatherley). (Australian National University Press: Canberra).

LITTLEJOHN, M. J. (1969). Amphibia of East Gippsland. Proc. R. Soc. Vict. 82 (1),
105-112.

LITTLEJOHN, M. J. (1971). Amphibians and Reptiles of Victoria. Part 1. Victorian Year
Book No. 85.

LITTLEJOHN, M. J. & A. A. MARTIN (1965). The vertebrate fauna of the Bass
Strait Islands: 1. The Amphibia of Flinders and King Islands. Proc. R. Soc. Vict. 79,
247-256.

LITTLEJOHN, M. J., A. A. MARTIN & P. A. RAWLINSON (1963). New records
of frogs in East Gippsland. Victorian Nat. 80, 225-226.

LOVERIDGE, A. (1950). New frogs of the genera Cyclorana and Hyla from south-
eastern Australia. Proc. biol. Soc. Wash. 63, 131-138.

MARTIN, A. A. (1965). Tadpoles of the Melbourne area. Victorian Nat. 8 (5), 139-149.

MOORE, J. A. (1954). Geographic and genetic isolation in Australian amphibia. Amer.
Nat. 88: 65-74.

MOORE, J. A. (1961). The frogs of eastern N.S.W. Bull. Am. Mus. nat. Hist., /2/,
149-386.

PARKER, H. W. (1938). The races of the Australian frog Hyla aurea Lesson. Ann. Mag.
Nat, Hist: Cll): 2: (9): 7302-305.

PROCTER, J. B. (1924). Unrecorded characters seen in living snakes and description of a
new tree frog. Proc. zool. Soc. Lond. 1924 (4), 1125-1129.

STEINDACHNER, F. (1867 (1869)). Amphibien. In ‘Reise der Osterreichen Fregatte
Novara um die Erde in den Jahren 1857-1859’. (Vienna) Zool. Theil 7 (4), 1-70.

TSCHUDI, J. J. (1839). Classification der Batrachier mit Berucksichtigung der fossilen
Thiere dieser Abtheilung der Reptilien. Mém. Soc. Sci. Nat. Neuchatel 2, 30-75.

TYLER, M. J. (1971). The phylogenetic significance of vocal sac structure in Hylid frogs.
Univ. Kansas Publications Museum of Natural History. 19 (4): 319-360.

WERNER, F. (1893). Herpetologische nova. Zool. Anz. 16, 81-84.

Aust. Zool. 18(3), 1975 163

Courtship of the Platypus, Ornithorhynchus anatinus

R. STRAHAN
Australian Museum, Sydney, N.S.W., 2000

D. E. THOMAS
Taronga Zoological Park, P.O. Box 20, Mosman, N.S.W., 2088

ABSTRACT

An account is given of the courtship of a pair of captive platypuses over a
period of 16 weeks and of subsequent incubation by the female over a period
of six weeks.

The female takes the initiative in the courtship. Nine components are described
in the behaviour of the female and five in that of the male. The male marks
objects under water with cloacal secretion.

The findings are compared with the observations of other authors on the sexual
behaviour of wild and captive platypuses.

INTRODUCTION

Spending most of its life in burrows or under water, and being inactive
by day, the platypus is a difficult animal to study in the wild and little is known
of its normal behaviour. Courtship and mating, which in many mammal species
attract attention because of striking differences from normal, year-round, behaviour,
are virtually undescribed for the platypus, and the few existing accounts are at
some variance with each other.

Verraux (1848), who collected platypuses in tributaries of the Derwent
River near New Norfolk, Tasmania, observed an apparent instance of mating
one evening in September. According to him, the male pursued the female on or
near the surface of the water for about an hour, driving her into the rushes at
the edge of the river. There, he seized her with his bill by the nape of the neck,
with his spurs clutching her rump. The female swam about, struggling violently
and emitting cries like those of a piglet. Copulation lasted 5-6 minutes, after which
the animals ‘played’ together for more than an hour.

Burrell (1927) was sceptical of Verraux’s account, asserting that the male’s
bill is so constructed that it could not be used to grasp the female’s neck: “‘. . . he
manifestly did not understand that the extent to which the upper mandible over-
laps the lower would render that impossible, quite apart from the pliable nature
of the lips and the fact that both jawbones are divided at their extremities and

Aust. Zool. 18(3), 1975 165

R. STRAHAN and D. E. THOMAS

are pliable as far back as the secateuring ridges” (p. 168). Burrell also states
that platypuses do not make the sound attributed to the female by Verraux.

Burrell (1927) claims to have seen two matings in the Namoi River, New
South Wales. On 27 August 1909 at 0730, two platypuses came to the surface
and one began to swim in a circle around the other. The second soon followed so
that they swam in a circle. “After about one minute of this circling, one of the
animals (which proved to be the female) submerged its body and tail and floated
perfectly still with its head above the water. The male then came slowly up and
mounted in a leisurely fashion. The whole process offered a very close resemblance
to the early stages of copulation of a drake and duck with the exception that the
male platypus did ot take a grip with its bill. The male then threw himself back
into a sitting posture, partly out of the water, but at this moment there was a
great splash and both animals disappeared’ (p. 168).

On 23 September 1921 at 0700, on the Namoi River three miles from the
site of his 1909 observations, Burrell observed “‘a pair of platypuses coupled in
an extraordinary position. The tail of each was laid flat along the belly of the
other. . . . The precise position of the hindlimbs could not be made out, as no
movement thereabouts was discernible; but it must have been the grip of these
(spurs) that kept the animals together. So closely were they apposed that they
appeared at times like a giant platypus” (p. 169). The conjoined pair, female in
a normal position, male upside-down and _ tail-forward, dived and_ surfaced
repeatedly for a period of three minutes, then separated and swam on the surface.
Since the animals were already coupled before Burrell saw them, the actual period
spent in this posture was not less than three minutes.

Home (1802) proposed a sexual role for the venomous spurs of the foot of
the male platypus. “It is probably by means of these spurs, or hooks, that the
female is kept from withdrawing herself in the act of copulation, since they are
very conveniently placed for laying hold of the body on that particular occasion”
(p. 72). Burrell agrees with Home and suggests that the spurs of the male fit
into corresponding sockets in the foot of the female and proposes a rather complex
manoeuvre by which a male could proceed from the sitting posture that he
observed in 1909 to that observed in 1927.

Fleay (1944) observed the apparent onset of courtship in two platypuses
in captivity at the Sir Colin Mackenzie Sanctuary, Healesville, Victoria. In mid-
September 1943, “Jack seized Jill’s tail in a firm grip with his bill and the two
animals swam slowly in a processional circle” (p. 11). This behaviour was observed
several times over the next 27 days. Mating took place on 11 October but the
details are somewhat obscure. “It is worth noting that during the act when the
animals were fast for nearly ten minutes, no spur grip was noted. A good deal
of splashing and floundering about occurred, and in the first place the male doubled
his body under him while maintaining his grip on the. female’s tail with his bill”
(p. 29). The male was removed from the enclosure on 18 October and the mating

166 Aust. Zool. 18(3), 1975

COURTSHIP OF THE PLATYPUS

was followed by the hatching and successful incubation of one young (the only
such breeding in captivity).

Burrell (1927) and other authors reviewed by him have described the nest
of leaves and grass used by the female as a brooding chamber. Fleay (1944)
described the way in which the female collects wet leaves and carries them to
her burrow, clamped between the ventrally curled tail and the abdomen. On 23
October, twelve days after copulation, the female collected leaves vigorously
and apparently completed her nest. She fed ‘ravenously’ on 25 October, then
retired to her burrow, returning to the water briefly on 31 October and 1 and 3
November, but not eating again until 6 November, 12 days later, when she resumed
feeding at a level much greater than normal. It is reasonable to assume, with
Fleay, that the period of fasting approximates to the period of incubation of the
eges, and that subsequent heavy feeding is related to lactation.

Collins (1973) states that two platypuses in the Bronx Zoo, New York,
displayed tail-holding and circular swimming from 21 June to 23 June, 1953.
Sixteen days later the female displayed nest-building behaviour.

MATERIALS AND METHODS

The Platypus House, Taronga Zoo, Sydney, was designed to the brief of
the senior author with the following major provisions: (i) the animals are exhibited
in subdued light in a long glass-sided tank containing flowing water, and surrounded
by a darkened viewing gallery; (ii) the animals feed in two large tanks of still
water, open to air and daylight; (iii) separate nesting boxes are provided for the
male and female and these are connected with the exhibition tank and feeding tanks
in such a way that the male and female have free and independent access to each
(it being possible to restrict animals to separate feeding tanks if required). The
system is designed so that there is no need to handle the animals and so as to
provide them with sufficient information on day-length to maintain annual
rhythms.

The exhibition tank has glass sides 6.1 m long and 1.3 m high. It contains
approximately 8,000 | of slowly running fresh water and is lit from above by
diffuse indirect sunlight, supplemented by fluorescent lamps when the animals are
exhibited: the viewing gallery is illuminated only by the light from the tank. A
tunnel from the exhibition tank leads to the male and female nesting boxes, and
paired tunnels lead from these to two feeding tanks, each 5.2 m long and each
containing approximately 4,000 | of water. Approximately one month prior to
the observations recorded below, an annex 120 cm long x 46 cm x 46 cm was
connected to the female’s nesting box. This was filled with damp earth and leaves
in the hope that it would be used for an incubation burrow.

Animals are enticed from their nesting boxes, via the tunnels, to the exhibi-
tion tank for about two hours each day (1100-1200, 1400-1500). A small
amount of food (mealworms, woodlice, scarabaeid larvae, small crayfishes) is

Aust. Zool. 18(3), 1975 167

R. STRAHAN and D. E. THOMAS

placed in the exhibition tank and the shutter leading to it opened. The animals
have learned that operation of the shutters is a signal for food and enter the feeding
tank without being handled or driven. Once in the tank, the shutter is closed and
the animals compelled to remain there except on the rare occasions when the female
has persisted in attempts to return to her nest. After an hour, the shutter is opened
and the animals usually retire 5-10 minutes later.

Food is placed in the outside feeding tanks at about 1500 each day and
the considerable debris is removed when the tanks are cleaned and refilled at
about 0800 each day. Although they have free access to the feeding tanks, the
animals seldom enter during daylight.

The behaviour described below was recorded during the two hours of daily
exhibition by one of the authors (D.E.T.), assisted by keepers of the Mammal
Department, Taronga Zoo. !

The male and female platypus were taken from the Kangaroo River, New
South Wales, and introduced directly to the newly-completed Platypus House in
May, 1969. Both animals were sub-adult. At the time of writing (January, 1975),
they were still in good health.

FEMALE. BEHAVIOUR

The courtship behaviour of the female comprises a number of components

that may be performed as isolated acts or in sequences. These are briefly described
below.

Interest in the male

For most of the year, the male and female show no interest in each other,
swimming past without reaction or apparent recognition. In view of the solitary
habits of platypuses in the wild, their relatively distant spacing along river banks,
and reports of combat (Burrell, 1927) one might expect aggressive behaviour
between individuals: lack of such behaviour in these captive animals may be due
to habituation.

In late August 1971, the female began to evince interest in the male. If the
male came near her while she was grooming (out of the water) she ceased this
activity and entered the water, lying passively until the male moved away—when
she would leave the water and resume grooming. This unoriented approach is the
simplest component of the female repertoire.

From September to December, the interest of the female in the male increased
and, as described below, involved specific, oriented, approaches to the male. It
should be noted that sequences of interaction were usually of short duration,
separated by much longer periods in which the animals ignored each other.

168 Aust. Zool. 18(3), 1975

COURTSHIP OF THE PLATYPUS

Fig. 1. Female courtship behaviour. A, frontal approach; B, side-passing; C, ventral-passing;
D, cloacal grooming; E, tinder- -passing.

Frontal Approach

The female approaches the male head-on. As the male rests on the surface
of the water, the female swims to him and rests with her bill touching, or nearly
touching his, the two bodies in a straight line (Fig. 1A). This relationship may be
retained for as long as ten minutes.

Side-passing
The female swims to the male as in a frontal approach, then passes alongside
him, rubbing against his side (Fig. 1B).

Ventral-passing

The female swims to the male as in a frontal approach but rolls over just
before meeting him, so that her ventral surface rubs against his as she glides
beneath him, apes down (Fig. 1C). Often the male responds by grasping her
tail and Glens her (see ‘tail-holding’, p. 171).

Aust. Zool. 18(3), 1975 169

R. STRAHAN and D. E. THOMAS

Q A Ch

Fig. 2. Male courtship behaviour. A, tail-holding. B, neck-holding; C, sitting posture
(possibly copulation); D, stretching. Where the position of the limbs is uncertain,
the outline is incomplete.

Under-passing

The female approaches the male from one side as he rests on the surface,
dives under his body, and surfaces again (Fig. 1E), the male may respond by
tail-holding.

Circling
The female swims in a tight circle (always clockwise in our female) on the
surface of the water. The male may follow and engage in tail-holding.

Grooming of cloacal region

While at the surface of the water, the female curls forward, almost into a
ball, and grooms the cloacal region with her bill for 10-20 seconds (Fig. 1D).
The animal may take up this posture lying on its side.

Regurgitation
The female, lying motionless on the surface, regurgitates a muddy vomit.
As this sinks to the bottom in a cloud, the male swims through it repeatedly

170 Aust. Zool. 18(3), 1975

COURTSHIP OF THE PLATYPUS

and the female follows him. The male grasps her tail and swims with her to the
surface.

Clas ping

When the male grasps the nape of the female’s neck, she rolls onto her
back, presents her ventral surface to his and the pair roll over in the water for
10-20 seconds. It is not clear which animal is clasping, or whether both are
involved. |

Leaf-carrying

Strictly speaking, this is an aspect of maternal behaviour rather than court-
ship. The behaviour is as described by Fleay (1944). The female picks up leaves,
either from the bottom or the surface, by tucking them between the ventrally
curled tail and the abdomen.

MALE BEHAVIOUR

The greater part of the behaviour of the male is in response to stimulation
by the female. Distinguishable components of male courtship are described below.

Following
The male follows the female as she swims in a circle.

Tail-holding

The male seizes the tip of the female’s tail in his bill and is towed behind
her at, or below, the surface of the water (Fig. 2A). The male may assist locomo-
tion with his forelegs or be pulled along passively. As mentioned above, the male
may be stimulated to this behaviour by the female’s circling, ventral-passing,
under-passing, or regurgitation. ,

Continued grasping of the female’s tail leads to loss of hair in two patches
on either side of the midline (possibly from pressure by the tips of the male’s
premaxillae). We have no evidence that these paired bald patches arise, as
suggested by Burrell (1927) from the use of the tail to plug the burrow with
balls of mud, although generalised wear of the hair at the tip of tail in either sex
may well be due to this.

Neck-holding

The male, previously engaged in tail-holding, releases his grip and climbs
upon the female’s back, grasping the fur on the nape of the female’s neck with
his bill (Fig. 2B). She responds by rolling onto her back (see ‘clasping’, above).

Marking
The male swims to the bottom, settles over a stone or other object, and
everts the cloaca, from which is produced a yellow, mucilaginous liquid which

Aust. Zool. 18(3), 1975 171

R. STRAHAN and D. E. THOMAS

forms a cloud, settling over the object. After secretion of the substance, the male
swims forwards, away from it. Although we have no direct proof of its function,

we regard this as marking behaviour. Interestingly, the female has not been
observed to show any interest in these marks.

Fig. 3. Platypus eggs in nest. This photograph was taken on Day 113 (13 November 1971)
while the female was in the exhibition tank. The only disturbance involved was
removal of the lid of the nesting box and the placing of two pieces of card (handled
with surgical gloves and forceps) alongside the eggs. The three compartments of
the nesting box are connected by small apertures to form an S-shaped chamber
with a single entrance (out of sight) at the lower left of the photograph.

172 Aust. Zool. 18(3), 1975

COURTSHIP OF THE PLATYPUS

Stretching

When normally at rest at the surface, the male has its four legs extended
downwards and outwards from the body; the bill and tail are horizontal and half-
submerged. During the period of courtship, the male was seen on a number of
occasions to stretch itself at the surface, raising its bill and tail above the surface
and extending its forelegs backwards and upwards along its sides (Fig. 2d). The
significance, if any, of this behaviour is not known. It has not been observed out-
side the mating season.

COURTSHIP

Daily observations of the behaviour of the two animals was initiated on
August 22, 1971 (Day 1) when it was noticed that the female had been making
pellets of earth (“‘pugs” of Burrell, 1927, p. 129) from the material in the annex
to her nesting box (described above). We thought that she might be about to
seal herself off for oviposition, but this proved not to be the case, and pellets were
not seen again, although the nesting vos and annex were opened every 14 days
thereafter for four months.

To summarise the records of 22 weeks, these are grouped below into five
successive 28-day periods and a final period of 20 days.

Period 1 (Days 1-28, 22 August-19 September)

The female showed interest in the male from Day 3 and would enter the
water when he came near. Tail-holding in response to circling was observed from
Day 5 onwards. Frontal approaches and side-passing were first observed on Day
10, as was the single instance of regurgitation. Ventral passing was first observed
on Day 11 and on this day very small bare patches were seen on the female’s
tail. Under-passing was first observed on Day 13. Marking was first observed on
Day 16, as was stretching.

From Day 1 to Day 23 there was a gradual increase in intensity of female-
male interactions, all of the above-mentioned components of behaviour (except
regurgitation ) increasing in frequency. From Day 24 onwards, the male was notably
less responsive.

There was no obvious succession of the components of female behaviour, but
it seemed that the frontal approach developed into side-passing, and ventral passing.
Under-passing was always less frequent than side-passing.

By Day 28, two distinct bare patches had been worn on the female’s tail.

Period 2 (Days 29-56, 20th September-17th October)

Although all components of behaviour observed during Period 2 were
displayed during Period 1, the frequency and intensity were less, particularly on

Aust. Zool. 18(3), 1975 173

Amount eaten

R. STRAHAN and D. E. THOMAS

the part of the male, who was often unresponsive to repeated approaches. by the
female.

Period 3 (Days 57-84, 18th October-14th November)

Early in this period, i.e. from about Day 60, the intensity of courtship
increased. The male occasionally chased the female for the length of the exhibition
tank. |

Period 4 (Days 85-112, 15th November-12th December)

The general intensity of male-female interactions increased and new com-
ponents were added. On Day 85 the male was first observed in neck-holding:
the female responded almost immediately by rolling over on her back and apposing
her ventral surface to his, but the two did not clasp. Clasping in response to
neck-holding was observed on Days 86 and 105: on the latter occasion the pair
remained clasped together for about 20 seconds, rolling over and over in the
water. On Day 104 the male displayed an element of female behaviour, ventrally
passing the female.

Cloacal grooming, by the female, was first observed on Day 86 and sub-
sequently almost daily until Day 112. On Day 104 the female was observed to
carry leaves from the bottom of the tank to the exit platform of the exhibition
tank.

Period 5 (Days 113-140, 13th December-9th January)

On Day 113, the female entered the exhibition tank as usual at 1100.
Examination of her quarters revealed two eggs (Fig. 3) in the nesting box (not
in the adjoining earth-filled annex). While the female was in the exhibition tank,
the male engaged in tail-holding, which the female accepted normally.

ar 10 20 30. | AO
113 120 130 140 150

Days (upper, from discovery of eggs; lower, from start of observations)

Fig. 4. Feeding behaviour during incubation and subsequently. 0, no feeding; 1, small amount
eaten; 2, normal amount eaten; 3, more than normal amount eaten; 4, much more
than normal amount eaten.

174 Aust. Zool. 18(3), 1975

47
159

COURTSHIP OF THE PLATYPUS

The male was separated from the female at 1200 and the Platypus House
closed indefinitely to the public. The female was given free access to the exhibition
tank and one feeding tank, and close attention was paid to the amount of food
eaten by her each night. The female did not leave the nest between Day 113 and
Day 117 (when she emerged at night and fed normally). She did not feed again
until Day 124. The pattern of feeding during Periods 5 and 6 is illustrated in
Fig. 4.

The female made very few daylight excursions during Period 5, being seen
only on Days 126, 127, 135 and 136. When in the water she was notably shyer
than usual.

Period 6 (Days 141-161, 10th January-30th January)

Over this period the female fed every night, usually eating more than the
normal (non-courting) amount. She made more frequent excursions in daylight
and was less shy.

The nesting box was opened on Day 159 and the eggs were found to be
shrivelled and without any sign of embryonic development. The male and female
wete reintroduced on Day 161 and showed no courtship behaviour or any
obvious interest in each other apart from occasional slight aggression (bill-
snapping) on the part of the female when the male passed near her.

DISCUSSION

We are aware of many shortcomings in the observations recorded above. The
observational methods were not planned in advance and the system of recording
involved several persons with no previous experience. The records are qualitative
and we recognise from the daily notes that behaviour that was regularly repeated
tended not to be logged by some observers: we have made some allowance for
this in the analysis given above and in Table I in which the behaviour is further
summarised.

It could be objected that it is inappropriate to study the behaviour of a
crepuscular-nocturnal species in the middle of the day — and we accept this. It
should be recognised, however, that daytime activity had been “grafted onto’”’ the
normal activity cycle two years previously and had been reinforced daily since
then. It: is likely that the intensity of male-female interactions was much greater
outside the periods of observation and that the behaviour observed in the late
morning and early afternoon did not cover the full span: certainly it seems not
to have included copulation. Nevertheless, the behaviour described has a self-
consistency that indicated ‘naturalness’, and our findings are reasonably in accord
with those of other workers. Comparing our observations with theirs, a number
of points of interest arise.

Aust. Zool. 18(3), 1975 175

R. STRAHAN and D. E. THOMAS

TABLE 1
RELATIVE FREQUENCY OF COMPONENTS OF COURTSHIP BEHAVIOUR

Period 1 2 3 eA 5 6
Day no. 1-28 29-56 57-84 85-112 113-140 141-16]
Female behaviour

frontal approach ++ oils

side-passing cine eae aoe apo ts

ventral-passing + ee eee cea res

under-passing a ae a ane

circling *

cloacal grooming en ae

regurgitation *

clasping ai haieee ey alee On ale

leaf-carrying a

Male behaviour

tail-holding cine ae vase as sirot at Alec -

neck-holding a RE

marking 3 at als sa as Aine ie

stretching . SE eStats a wats

clasping Soci

Note: ++ indicates infrequent (less than daily)

++ indicates relatively frequent (at least daily)
+++ indicates frequent (several times daily)
s indicates observed occurrence on only one day

Duration of courtship

It is possible that a low level of courtship may have passed unnoticed prior
to our institution of regular observations in late August. Thus the period of 110
days between the beginning of observations and the discovery of eggs would
appear to be the minimum duration of courtship in this case. Burrell (1927)
has no direct information to offer on the duration of courtship. Fleay (1944)
observed courtship over a period of 27 days. Courtship in the Bronx Zoo (Collins,
1973) was observed for no longer than 16 days.

At least two.possible reasons come to mind for the great discrepancy between
our observations and those mentioned above. The first is that the situation in
Taronga Zoo was abnormal (although hardly more so than that obtaining in New
York). The second is that observers who were limited to a view of the animals
from above the water surface were able to see much less than we were able to
through the glass walls of the display tank and therefore failed to observe much
of the behaviour. We are inclined to the latter explanation, while not excluding
the possibility of some abnormality in the behaviour of our animals such as the
possible habituation mentioned earlier, and the possibility, mentioned below, that
our records extend over an ovulatory cycle subsequent to the normal one.

176 Aust. Zool. 18(3), 1975

COURTSHIP OF THE PLATYPUS

Time of mating

Burrell’s two observations of mating were made in late August and late
September. Less direct, but convincing evidence for the time of mating comes
from the collection of eggs from nests. He found none in New South Wales
eatlier than 24th August or later than 6th November (Burrell, 1932). Based on
records of egg-collecting elsewhere, Burrell suggests that mating is from July to
August in central Queensland; from August to September in New South Wales;
and from September to October in Victoria. If this is so, oviposition in our
female was three months late.

A possible explanation for this is suggested by the events that led to the
institution of regular observations; unusual behaviour of the female in September.
As mentioned earlier, she had been making balls of earth and it is possible that
this was either an expression of an earlier, failed, mating that took place at
the usual time, or overflow activity arising from a strong but unfulfilled mating
drive. It seems probable that the courtship observed by us was the outcome of
a second oestrus, such as suggested by Burrell (1932), but we cannot tell whether
or not this led to an unusually long courtship.

Role of the female

Our observations clearly demonstrate that the initiative of courtship rests
largely with the female. The components of her stimulatory behaviour proceed
in increasing complexity from initial interest, through the frontal approach, to
side-passing, and ventral passing. Under-passing does not fit neatly into this
sequence and is less frequently observed.

Role of the male and copulation

We agree with the other authors that following the female in a circle and
tail-holding are early components of the male’s responses to the female. Tail-
holding is also released by ventral passing and under-passing on the part of the
female, and by her regurgitation. We have observed transition from tail-holding
to climbing on the back of the female and assert (contrary to Burrell) that the
male can, and does, grip the neck of the female with his bill.

In 1971 we were convinced that copulation took place head to head and
abdomen to abdomen, for such behaviour was observed at its peak 8 days prior
to the discovery of eggs. However, this behaviour lasted only 20 seconds and we
now interpret it as pre-copulatory. In a chance observation during the breeding
season of 1974, the male was seen to move from a grip on the neck of the female
to a sitting posture with his tail wrapped under the side of the female’s tail (Fig.
2c), as described by Burrell in his 1909 observation and by Fleay (1944). For
the position described by Verraux to have been functional, the tails of the two
animals would also have needed to be oriented in this way. This sequence and
final posture seems natural, whereas that observed by Burrell in 1921 does not.
A possible explanation of his observation is that the male became dislodged from

Aust. Zooi. 18(3), 1975 177

R. STRAHAN and D. E. THOMAS

the sitting posture while his barbed penis was still lodged within the female and
that he was towed helplessly behind her, upside-down.

Burrell’s assertion that the spurs of the male are used to lock into sockets
in the feet of the female is based on his supposition that the tail-to-tail, upside
down position of the male is normal for copulation. In the absence of any evidence
that the spurs are so employed, his hypothesis is extremely weak: the copulatory
position of the platypus seems to conform to the general mammalian pattern.

Marking

The platypus has a prominent sternal gland which increases in size in the
male during the breeding season and is used to mark burrows with ‘a foxy odour’
(Burrell, 1927, p. 163). As we have shown, the male platypus also marks under-
water. The nature of the secretion is unknown, but it is not faecal. The two
mucous glands opening on either side of the wall of the cloaca by multiple apertures
(Home, 1802) seem to be suitably situated for this function.

Cessation of feeding during incubation

Our observations are in accord with those of Fleay (1944) that the female
does not feed for some days following oviposition. In the case recorded by Fleay,
the female fasted for 12 days; in our case for 11 days (with one feeding on

the fifth day).

ACKNOWLEDGEMENTS

We gratefully acknowledge the assistance of Mr. A. Carrick, Mr. D. Igglesdon, and
Mr. S. Dnombacs who assisted in compiling the daily observations, and to Mrs. VY.
Broadsmith who recorded some observations in sketches.

REFERENCES

BURRELL, H. (1927). “The Platypus’, Angus and Robertson, Sydney.
BURRELL, H. (1932). Fact and fancies pertaining to platypus. Aust. Zool. 7, 110-115.

COLLINS, L. R. (1973). ‘Monotremes and Marsupials’, Smithsonian Institution Press,
Washington.

FLEAY, D. (1944). Observations on the breeding of piatypus in captivity. Victorian
Naturalist, 67, 8-14, 29-37, 54-57, 74-78.

HOME, E.. (1802). A description of the anatomy of the Ornithorhynchus paradoxus. Phil.
Trans. Roy. Soc. Lond., 1802, 67-84.

VERRAUX, J. (1848). Observations sur l’Ornithorhynche. Rev. Zool. 11, 127-134.

178 ’ Aust. Zool. 18(3), 1975

Mammals of Sturt National Park, Tibooburra,
New South Wales

M. J. S. DENNY
School of Zoology, University of New South Wales, Kensington, N.S.W.

ABSTRACT

A survey of mammals inhabiting a national park in the north-western corner of
N.S.W. was undertaken during 1973 and 1974. Fourteen species were identified,
many of which were new records for this region, Detailed observations were made
of the habitat preferences, distribution and population numbers of several of the
small mammals. It would appear that up to five mammal species can inhabit one
area at the same time, particularly an area subjected to periodic flooding. How-
ever several of these mammals have discrete distributions and there is little
overlap between different species. Numbers of animals changed noticeably during
the study period, particularly those associated with periodic flooding. It is sug-
gested that many of the small mammals living in the region surveyed are not
arid-adapted but are dependant upon those areas scattered throughout the arid
zone which have a habitat similar to that of a more temperate climate.

INTRODUCTION

In 1972 a large area of land in the north-western corner of New South
Wales was set aside for the formation of a national park. This park, called Sturt
National Park, now encompasses an area of approximately 200,000 hectares and
further land acquisition will increase its size. Sturt National Park is situated ap-
proximately 340 km north of Broken Hill and comprises a variety of landforms
typical of an arid and semi-arid environment. At the northern end of the park is
the southern extremity of the Grey Range, with its associated mesa formations
and eroded gullies. The remainder of the Park consists of the occasional stoney hill
and numerous drainage basins which are influenced by Frome and Twelve-mile
creeks (Figure 1).

Three major plant communities are recognized for this area (Beadle, 1948).
These are the Mulga (Acacia aneura) association, the saltbush (Afriplex) associa-
tion and the bluebush (Kochia) association. Because of the unusually good seasons
during 1973-74 many grasses were also present, Mitchell (Astrebla spp.) and Flin-
ders grass (Iseilema membranaceum) the most common. Climatically, Sturt National
Park can be classified as within the arid area of Australia having a mean rainfall
of under 200mm whilst the mean annual temperature is 27.4°C. Maximum tem-
peratures are as high as 49°C. However during the survey period, record quantities
of rain fell (over 760 mm of rain was recorded for the first five months of 1974).

Aust. Zool. 18(3), 1975 179

M. J. S. DENNY

During 1973 and 1974 I was able to live within the National Park and,
in conjunction with other research, I conducted a survey of the mammals in this
relatively little known atea.

MATERIALS AND METHODS

Sampling of the mammal population was mainly achieved by use of Elliott
live animal traps baited with a mixture of rolled oats, peanut butter and fried
bacon. Over 4000 trap-nights were accomplished with an average trapping success
of 14.5% (range 2-50% ). Estimation of population numbers of the House Mouse
(Mus musculus) and the Long-Haired Rat (Rattus villosissimus) was achieved by
use of the Capture-Mark-Release Recapture method (C.M.M.R.). The calculations
used are those outlined by Davis (1963). All animals were marked by using coded
eatholes and sprayed with a coloured dye (Vasco Stock Mark, V.S. Supplies Ltd.)
to ensure rapid identification.

RESULTS AND DISCUSSION

1 EUTHERIANS
Order: Rodentia
Family: Muridae
(a) Forrest’s Mouse (Leggadina forresti). Until recently this animal had not
been recorded in N.S.W. Morton (1974) obtained two specimens from Fowler’s

Mount Sturt

Mount Poole

Fig. 1. Map of the north-western corner of N.S.W., showing principal study sites and the
then western boundary of Sturt National Park. The eastern boundary had not been
precisely delineated at time of writing.

180 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK :

Gap Station, 110 km north of Broken Hill. To date only one specimen has been
found in Sturt National Park. This animal was caught by hand after being dis-
turbed from its nest. The nest consisted of a shallow burrow about 20 cm long
made in the cracking soil underneath a flat rock. The area where the mouse was
found was unusual for this species, being a flat-topped outcrop at the
edge of the Grey Range. This type of outcrop is characterized by a silcrete type
duricrust surface, cracking soil and shallow depressions similar to gilgais. Morton
(1974) caught both specimens on flat, treeless plains whilst Watts (1972) and
Parker (1973) describe the habitat of L. forresti as black soil alluvial flats and
grassy plains. However, the areas at Fowler’s Gap and Sturt National Park are
similar in one respect, both contain cracks in the soil which are probably used for
shelter,

The distribution of L. forresti is not limited to the Grey Range in Sturt
National Park as several fragments of Forrest's Mouse have been found in owl
pellets in an area surrounded by flat grassy plains.

(b) Sandy Inland Mouse (Pseudomys hermannsburgensis). According to
Finlayson (1961) this murid is not commonly found east of Sturt’s Line, however
Watts (1974) recently found P. hermannsburgensis in three localities within West-
ern Queensland. Only one specimen was captured within Sturt National Park, this
animal was trapped in a sandy creek bed within the low hills found at the edge of
the Grey Range. The area comprised of sandy soil with the dominant vegetation
being shrubs (hakea etc.) and trees (gidgee, mulga, corkwood etc.). There was
also extensive grass cover as well as many fallen trees.

(c) House Mouse (Mus musculus). Unlike Morton (1974) and Watts and
Aslin (1974) I have found that the Tibooburra region was experiencing a virtual
plague of these ubiquitous rodents. For instance, an area which yielded a 20%
trapping rate (all animals trapped were M. musculus) was not uncommon during
1973-74. Systematic trapping along 48 km of the Gidgealpa-Sydney Gas Pipeline
route (see Figure 1) covering eight different habitat types ranging from sandy creek
beds to stoney hills, yielded a predominance of house mice in all habitats.

There appear to be two types of M. musculus in the area. One type was
uniformly brown both dorsally and ventrally whilst the other type was fawn
coloured dorsally with white belly fur. These two types have been previously
described by Cleland (1918) and Jones (1923) whilst Johnson (1964) also men-
tions the phenomenon and likens the white form of mouse to the Mediterranean
sub-species M. musculus brevivostris. Of 105 house mice examined at Sturt
National Park, 56 were of the white form. No relationship between the colour
forms of M. musculus and their distribution could be found.

To obtain some estimate of the numbers of house mice C.M.R.R. studies
were carried out in an area where these animals were known to reside. This area
was beside a small creek (Chunky Creek, see Figure 1) about 3 km from Mt. King
homestead. Because of the closeness of semi-permanent water the vegetation was

Aust. Zool. 18(3), 1975 181

M. J. S. DENNY

relatively constant throughout the study period. The dominant plants being a
mixture of sedge, Leptochloa digitata and a small bush (Chenopodium species) with
a scattering of Coolabah trees (Eucalyptus microtheca) and an undergrowth of
native clover (Medicago sp.). Twenty-five traps were laid at 10m intervals during
a week in September 1973, then again, immediately after a large flood in March
1974. During the second period ninety-one traps were used.

Three mathematical methods were used to estimate population numbers:

(i) the Schnabel (Krumholz) formula (Schnabel, 1938).
(ii) the Schumacher-Eschmeyer Procedure (Davis, 1963).
(iii) a graphical method outlined by Hayne (1949).

During September 1973, the three estimates of the mouse population using
the three above methods were 14.7, 11.0 and 13.3. Considering the differences in
the three methods the estimates were comparable and the mean of three estimates
will be used, i.e. 13. The results of the 1974 trapping study were as follows. The
three estimates of population size were; 36.3, 30.5 and 35.7. This gives a mean
estimate of 34.

The difference between the two mean estimates was not entirely due to the
difference in the number of traps used and the resultant larger area covered in 1974.
By relating population numbers to the length of trap line (250mm in 1973, 400mm
in 1974) the density of mice in 1973 was 0.05 mice per metre and in 1974 0.09
mice per metre. Consequently a higher density of mice in 1974 would seem to
indicate a larger population in the trapping area during March 1974. A similar
picture of population fluctuation has been observed by Newsome in his study of
house mice inhabiting a reed bed in South Australia (Newsome, 1969a and b).
Newsome found that population numbers fell during the dry months when soil
suitable for nest-making (by burrowing) was limited. During the wetter months,
particularly if in summer, numbers increased because of a greater availability of
food and particularly nesting sites. It is interesting to note that a soil of similar
characteristics to that found by Newsome (1969a) occurred at that site studied
in the National Park. The ground consisted of a red soil (classified as Ug 5.3 by
Stace ef al., 1968) and due to its high proportion of clay, tends to form large
cracks when dry. During the drier months of 1973 the house mice had a limited
amount of soft cracking soil in which to burrow and make nests. During the early
months of 1974, particularly after the flooding experienced in January, the soil
was ideal for breeding and there was a consequent increase in numbers of mice.
The importance of the cracks in the soil is emphasized by Newsome (1969a), who
points out that the microclimate within the crack provides a cool, moist shelter
during the drier months.

(d) Plague or Long-haired Rat (Rattus villosissimus, Figure 2). This animal
has only been seen occasionally in the area over the last century (Plomley, 1972).
According to Finlayson (1961), its distribution appears to depend on local seasonal
conditions, the animals spreading from western Queensland and the Northern

182 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

Fig. 2. Long-haired Rat (Rattus villosissimus)

Territory during wet periods. However, Watts and Aslin (1974) believe that R.
_villosissimus are always present in small numbers in pockets of favourable habitat
from which they spread out during wet seasons. If this is so, then a pocket of
favourable habitat for the long-haired rats was the area studied at Sturt National
Park (the same area as described for Mus musculus. Here the rodents had estab-
lished two types of burrow system.

The most common type was a shallow hollowing out of the soft, moist ground
which lies beneath the dry, hardened crust of surface soil. This type of burrow
appears to be used as temporary cover for the rats, possibly as protection against
predators. The other type of burrow is more extensive and many of those dug up
contained a nesting chamber of dried grass. This burrow resembled a small rabbit
watren in that several side passages were always present and several separate bur-
row systems were usually found together in an area of soil suitable for burrowing.
Finlayson (1939) describes a similar burrow system for R. villosissimus from the
Lake Eyre Basin and I have found these burrows in the Cooper’s Creek region
(Windorah, Queensland). Nesting chambers would indicate that these rodents

Aust. Zool. 18(3), 1975 3 183

M. J. S. DENNY

were breeding and although no pregnant females were caught several immature
animals were found.

However extensive trapping prior to the 1974 floods yielded no evidence of
R. villosissimus either at the study site or in the surrounding area. If pockets of
long-haired rats did occur in the Tibooburra area during the drier periods of the
year they must have been spread out over a vast area and consequently migrating
rats would have had to move a relatively long distance and from other reports
they appear capable of such movements. There is evidence that R. villosissimus
reached Spencer’s Gulf via the Lake Eyre district in 1887 (Plomley, 1972).

Order: Chiroptera
Suborder: Microchiroptera

So far four species of bat have been found in Sturt National Park. These are
the Little Brown Bat (Eptesicus pumilus), Lesser Long-eared Bat (Nyctophilus
geoffroyi), Gould’s Wattle Bat (Chalinolobus gouldii) and the White-Striped
Mastiff Bat (Tadarida australis). All have been accidental captures either due to
hitting a moving car or a windmill or else captured alive by mist net. Considering
the number of deep caves found within the Grey Range it was expected that bats

Fig. 3. Sminthopsis froggatti

184 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

would be found inhabiting these sites, but after thoroughly inspecting twenty of
these caves no evidence of bat occupancy has been found.

2 MARSUPIALS

Order: Polyprotodonta
Family: Dasyuridae

Three species of this family have been found within Sturt National Park.
The three are:

(1) Swminthopsis crassicaudata
(ii) Sminthopsis froggatti (Figure 3)
(iii) Planigale gilesi (Figure 4)
Studies on P. gilesi and the two Sminthopsis species show an interesting ecological

picture of several little-known species of marsupial. These three species have been
found in similar areas within the National Park, however there is little overlap

in habitat.

Fig. 4. Planigale gilesi

Aust. Zool. 18(3), 1975 185

M. J. S. DENNY

The habitat of Planigale gilesi is perhaps the simplest to describe. This habitat
is as rescribed for that of house mice at the Chunky Creek study site, ie. red
cracking soil, thickly covered by vegetation which is predominantly sedge and
canegress. This area is subject to periodic inundation with water. Other areas
where P. gilesi have been found are the Bulloo Overflow (near Teuriko, see Figure
1) and the Channel country associated with Cooper’s Creek (Windorah, Queens-
land). It is interesting to note that Aitken (1972) described the habitat of P. gilesi
similarly and Troughton (1928) states that the habitat for two species of planigale
(P. ingrami brunneus and P. teniurostris) was beside large rivers in Queens-
land and N.S.W. and Parker (1973) gives a similar habitat for P. ingrami.

Both species of Sminthopsis also occur at the Chunky Creek site but are also
much more widely spread. S. crassicaudata was found throughout the Park mainly
associated with claypans and flood plains. S. froggatti was found primarily near
creeks, recently flooded areas and gilgais as well as in the sandy hills of the Grey
Range. This animal does not appear to be as widespread as S. crassicaudata and
is perhaps restricted to a more specialized habitat which requires either soft ground
for burrows or vegetation for cover. Where the three species were found together
(Chunky Creek) there is a definite separation of the Planigale from the Smin-
thopsis species. Figure 5 shows the area at Chunky Creek where a trap line was
laid and it can be seen that this line cuts through three different types of vege-
tation. Firstly the canegrass-chenopod association close to the creek, then a vege-
tation complex of Bassia quinquecus pis as the dominant species and finally a stand
of grass containing such species as Mitchell, Flinders, Umbrella (Chloris acicularis )
and Ray (Sporobolus actinocladus). The results dom he days trapping in June
1974 are shown in Figure 6, where the species of mammal found in each trap is
given. It can be seen that the range of S. froggatti and S. crassicaudata does not ex
tend into the chenopod complex whilst the planigale is only found within that
patticular habitat. Previous trapping has shown that when larger numbers of P.
gilesi are present the same distribution occurs.

_ A similar situation appears to occur for the marsupial mice caught within the
Park as that found for Mus musculus. Planigale appeared to utilize the cracks in
the soil as cover and/or nesting sites and are possibly dependant on the periodic
wetting of the ground to ensure the correct condition for burrowing. The influence
of favourable conditions upon population numbers was illustrated by the numbers
of Planigale caught at Chunky Creek during 1973-74. In January 1973, when the
creek still contained pools of water, four Planigale were caught, then again in May
another three were captured (all were released). After this, despite several months
trapping, no more Planigale were found until February — ane March 1974 when
eleven were caught. Similarly, no Sminthopsis froggatti were trapped during 1973,
but. sixteen were trapped in June 1974 at Chunky Creek.

| _The fall in numbers of Planigale during 1973 was perhaps due to the changed
conditions during the drier months of the year but was possibly aggravated by the
increasing numbers of house mice that occurred during that year. There is evidence

186 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

BASSIA

CHENOPOD “

GRASS

o

Fig. 5. The area of Chunky Creek used as a study site. Distances in metres.

of competition between Mus musculus and other mammal species not only for food
(Caldwell and Gentry, 1965) but also for a place to live (Briese and Smith, 1973).
In the present study competition for food eaten by both species would not have
been intense as there were large numbers of insects (particularly the Spur-throated
locust, Austracis guttulosa) in the area during 1973. Consequently an inability to
occupy the cracks in the soil by Planigale when facing competition by house mice
could have caused the fall in the number of marsupials. It is feasible that the mar-
supials were forced to move further away from the study site during the drier
months, this bringing with it the greater possibility of predation. Aitken (1972)
points out that the dense covering of vegetation associated with the habitat of P.
gilesi would provide a barrier against aerial and terrestrial predation. Predators of
small mammals in the Sturt National Park include feral cats, foxes, owls (remains
of Sminthopsis sp., L. forresti, Planigale sp. and Mus musculus have been found in
owl pellets) and fork-tailed kites. In appendix I is given a table of flesh measure-
ments of these three marsupials.

Family: Macropodidae
Subfamily: Macropodinae

Aust. Zool. 18(3), 1975 187

M. J. S. DENNY

Three species of the kangaroo family are found within Sturt National Park.
The two most common are the Red Kangaroo (Megaleia rufa) and the Euro
(Macropus robustus erubescens) whilst the Eastern Grey Kangaroo (Macropus
giganteus) is not often encountered. The distribution of these marsupials within
the Park is as would be expected from their known habitat preferences (Frith
and Calaby, 1969). The occurrence of M. giganteus at Sturt National Park is the
most westerly record of this kangaroo in this area of N.S.W. and it would appear
that these animals are gradually moving westwards, their numbers increasing in
areas west of the Darling River.

Some preliminary data is given in appendix II on various measurements taken
from over 100 red kangaroos trapped in Sturt National Park during 1973-74 (the
locations of the traps are shown in Figure 1, numbered 1 to 6). This data is being
incorporated into a larger study on the migratory habits of red kangaroos.

p
dabble
ET cAenropticnon|'sarnm <- lemaes

Fig. 6. Results of a trapping study in June 1974. Explanations of lettering is given in Table 1.

3 MONOTREMES

Although there is indirect evidence to show the existence of the Echidna
(Tachyglossus aculeatus) in Sturt National Park, none have been sighted by the
author. Use by the echidnas of the caves mentioned above appears to have occurred
in the past. In the majority of caves echidna droppings were found. These droppings
vary in length from two to eight cm and have a diameter of about two cm. They
are composed almost entirely of clay mixed with crushed insect cuticle. This cuticle
was found to be from ants, i.e. no termites (the predominant food for echidnas)
were eaten by echidnas in this area. This is possibly due, not to any dietary pre-
ference for ants but to a low incidence of termites in the Tibooburra area (M.
Griffiths, pers. comm.).

In the Cheviot Range in south-western Queensland a similar type of dropping
as well as echidna quills and a nest made of grass (similar to that used by echidnas )
were found in caves of the same structure as those occurring in the Grey Range.

4, FLOODING AND WILDLIFE

It is perhaps relevant here to give some general observations upon the influ-
ence of flooding on wildlife. There appear to be two patterns of flooding occurring

188 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

during heavy rains. The first to be experienced is the “local” flooding formed by
the large quantity of rain falling in a relatively short time, for instance approxi-
mately a year’s rain (150mm) fell in Tibooburra within 24 hours during January,
1974. Local flooding has three major characteristics:

(a) It is quick to begin, does not last long, and the ground rapidly dries
out after inundation.

(b) There is a strong water current not only draining off the slopes and
across the claypans, but also down the various creeks in the area.

(c) Large depositions of silt occur. This can be as deep as 1.5m and is
commonly found near the creeks and on the extensive claypans surrounding the
waterways. Deposition of silt implies removal of soil from other areas. This re-
moval is seen in the higher areas where channels are cut into the top 2m of ground.

Effects of local flooding are superficially disastrous. Any terrestrial animal
unable to reach high ground will risk being swept away in the ensuing water
movement. This, in fact, occurred during the floods in 1974 when large numbers
of Rattus villosissimus were washed away and drowned in the Cooper’s Creek.
A few dead (presumably drowned) kangaroos were also seen in creeks draining
into the Bulloo River Overflow (Narriearra Station, see Figure 1). Not all animals
washed away would die however, many were capable of surviving on trees or
small rises until the water drained away then they were able to repopulate an
area. This appeared to be the case with Sminthopsis crassicaudata, one being
caught just leaving the water after having been washed down Twelve Mile Creek.

Slow flooding occurs after the rains have fallen, sometimes over a fortnight
later, when the larger rivers and lakes in the area gradually fill to capacity then
spread out from their banks. This type of flooding was slow to occur and lasted a
long time. The time factor involved in the slow inundation of the area allowed
most animals to seek safety from the waters, but if local flooding occurred when
the rivers were rising, many animals were trapped and died either from drowning
or starvation. Consequently, slow flooding can be more disastrous than local
flooding.

The effects of flooding upon populations of animals appeared to be influenced
by three factors:

(i) Habitat Selection

(ii) Mobility; and

(iii) Population Numbers.

(i) Habitat Selection. Because the habitat for Planigale gilesi was the
“cracking” soils found near creeks or low flooding areas, these animals were sensi-
tive to the spread out of water that occurred during slow flooding. For example,
when the Bulloo Overflow flooded areas of cane-grass over 16 km from the main
channel, the Planigale in that area were not able to escape the waters and en-
deavoured to exist on the cane-grass left exposed above the water.

Aust. Zooi. 18(3), 1975 189

M. J. S. DENNY

(ii) Mobility. This applied mainly to the larger mammals, particularly the
red kangaroo. These animals, even during small rainstorms, seek the highest point
in the immediate area. Previous observations have shown that red kangaroos were
to be found on the stony undulating hills around Mt. King and Olive Downs during
the wet months and on the flats (using the shade trees in the creeks) during the
drier months. Aerial traverses of flooded areas during early 1974 showed that few
kangaroos were seen on the still wet flood plain or near the creeks. Most kanga-
roos sighted were found in the hills except those trapped by the rising flood waters.

During local flooding kangaroos sought high spots, e.g. sandhills, and stayed
there until the surface water had run off, then spread out to drier areas being able
to travel through mud up to 1m deep. Kangaroos staying on the sandhills and
small rises were in danger of being trapped if the slow flooding due to river risings
isolated their refuge. This was seen in areas flooded by the Bulloo River where
one “island” in the flooded Overflow held 137 kangaroos.

(iit) Population Numbers. Large populations of mammals were more sus-
ceptible to flooding and its after effects than smaller populations. For instance the
portion of long-haired rats killed by drowning in the Windorah (Qld.) area
was far in excess of any other animal observed (over 60 carcases were seen).
Those left would also find it difficult to survive because of the relatively small area
of dry land left. The trapping frequencies found during this survey were high
compared with other surveys carried out in Australia. Calaby (1971) pointed out
that in the United States catches are in the order of 20%, sometimes higher,
whereas in Australia catches average 2 to 5%. Trapping frequencies up to 50%
were found in the present study. Calaby then states that in times of high population
density of particular species, large numbers of animals may be caught. It was this
situation that was found during the survey described here, considering the high
densities of Mus musculus. However small islands may also represent a situation
where high densities of mammals may be found (Calaby, 1971). An island does not
necessarily have to be a piece of land surrounded by water but is better defined as
an area hospitable to certain species surrounded by an area through which move-
ment is limited because of the vastly different environment found there, i.e. an
area of acceptance surrounded by a sea of inhospitality.

Arid and semi-arid land cannot be strictly defined by an overall. description
because closer inspection yields a mosaic of differing habitats. Sometimes these may
only be the size of a fallen log or a small pool of water. In a forest environment
this mosaic is more complicated with numerous microhabitats existing in a small
area (Calaby, 1966; Brereton, 1973). In the drier areas, although not as numerous,
many such microclimates exist and some clearly defined examples are the surrounds
of creeks (whether permanently watered or periodically flooded), stony outcrops
areas of cane-grass found in drainage basins and even rubbish tips (now an estab-
lished feature of a pastoral area).

It is this situation that exists in Sturt National Park. Small mammals, other
than Mus musculus, were not caught in all places but only in those areas that can

190 Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

be defined as “islands”. For instance, the trapping site at Chunky Creek yielded
five different mammal species during a single night’s trapping (Mus musculus,
Rattus villosissimus, Sminthopsis froggatti, S. crassicaudata and Planigale gilesi)
and gave a mean capture rate of 50%. An area away from the creek on the open
ground, yielded only house mice and gave a capture rate of 5%. To further il
lustrate this point several different areas were chosen within the park and trapped
for mammals. The results of this survey are shown in Table 1. It is apparent that
those areas containing adequate ground cover for small mammals were the areas
where the highest trapping frequency and species diversity were found, i.e. at
Chunky Creek, Mulga Hills and gilgai.

Other examples of high densities of mammals in “‘island’’ situations were
found outside the park. Fifteen Planigals gilesi were rescued from the cane-
grass stalks protruding above the water in the flooded Bulloo Overflow. This was
in an area of about 120 hectares and no other small mammals were found during
several journies over the Overflow. Another example was seen on the Cooper’s
Creek where fourteen Sminthopsis (a mixture of froggatti and crassicaudata) were
spotlighted along fourteen miles of road running near the creek.

A single similarity exists between all habitats where mammals were caught,
i.e. the need for shelter from climatic extremes. Whether at Cooper’s Creek, Bulloo
Overflow, Chunky Creek etc., all habitat sites contained ground shelter which ap-
peared to be used by the animals during the day. Consequently, these mammals
are not necessarily adapted to desert living but are animals living within a more
temperate environment within an arid area. Even during extensive dry times the
characteristic shelters still exist and, as long as food supplies hold out, these ani-
mals should be able to survive. The recent geological past was climatically easier on
the animals in the area. During the late Pleistocene the climate was cooler and
more humid with perhaps a higher rainfall. Since the end of the Pleistocene the
climate has become drier with several climatic fluctuations occurring
(Keast, 1968; Jones, 1968). Such a sequence would allow establishment of arid
adapted species in the drier areas. Those species adapted to a more temperate
environment would either move further toward the still moist coastal areas or es-
tablish colonies in the islands of less arid habitat still retained in the arid areas.

Such animals would possibly have been the native mice caught within Sturt Natio-
nal Park.

There is need for further work to be carried out on the situation of small
native mammals within the arid zone, particularly in regards the situation described
here, where periodic flooding occurs. Sheppe (1972) describes a similar situation
in a Zambian floodplain and points out the remarkable adaptability of the small
mammals to such an unstable environment. It would appear that those mammals
associated with periodic flooding in Sturt National Park are equally adaptable.

Aust. Zool. 18(3), 1975 191

M. J. S. DENNY

TABLE

Species diversity and trapping frequency at various sites in Sturt National Park

Site

Mokely Ck.

Mokely Ck.

(upstream )

Mokely Ck.

(branch)

Whittabreena

Creek
Gilgai

Olive Downs

Hills I

Olive Downs

Hills I

Flood Plain

Chunky

Creek

Sand Hill

Trapping
frequency
(mean %)
and date

Habitat

Dense low vegetation beside

creek, cracking soil. |

Sandy creek bed.

Dense vegetation, cracking

soil.

Sparse vegetation, dry creek.

Dense, low vegetation, damp

cracking soil.

Sparse grass, sandy soil.

Tree cover, sandy and stony

soil, logs.

Sparse, low vegetation, soft,

silty soil.

Dense low vegetation beside

creek, cracking soil.

Sparse vegetation, sandy soil.

Dates trapped in parenthesis.

M.m. denotes Mus musculus

R.v. denotes Rattus villosissimus

P.h. denotes Pseudomys hermannsburgensis

S.f. denotes Sminthopsis froggatti

S.c. denotes S. crassicaudata

P.g. denotes Planigale gilesi

192

19
28

20

21

(5/74)
(7/74)

(7/74)

(7/74)
(6/74)
(6/74)
(6/74)
(6/74)
(6/74)
(3/74)

(6/74)
(7/74)

Species
diversity

1 (M.m.)
1 (M.m.)
1 (M.m.)

3 (CMe Rive S&)
1 (M.m.)
SCM mi. Reva. Sates)
1 (M.m.)

3 (Mim Pohs St)

3°. CMim.s Sa S:c2)

ASC MGmS Riv. Sefes Poe)
SMa IRV, 1 Sa Cae ee

1 (M.m.)

Aust. Zool. 18(3), 1975

MAMMALS OF STURT NATIONAL PARK

APPENDIX I

Flesh measurements of three species of marsupials taken from live animals

Head and Body

Species Head Length Ecansh Tail Length Foot length
Planigale DAA Se 02077 6.69 + 0.94 GIF = O14 tit] 007
gilesi (10) (10) (10) (10)
Sminthopsis DAT =e 1.068 6.79 + 0.091 6.5 722-- 0,085 Db O03)
crassicaudata (8) (8) (8) (8)
Sminthopsis 2h Bin 0.056 Liss OL 855 9.59 + 0.199 1.74 + 0.045
froggatti (18) (18) (18) (18)

All measurements in centimetres and given as mean + standard error, number of
animals in parenthesis.

APPENDIX If

Data on red kangaroos (Megaleia rufa) caught in Sturt National Park during 1973 (in
collaboration with G. Wilson, National Parks and Wildlife Service).

Males, $3.7" ==" 2 82 ke (n=53)
(range 9 to 95 kg)

Female: 25.1 -- 0.61 kg (n=74)
(range 9 to 37 kg)

Body weights

Pouch young : Mean age: 106 days (n=31)
(range 2 to 220 days)
Sex ratio: 20¢:; 79

* Mean = standard error.

ACKNOWLEDGEMENTS

I wish to thank the many people who helped me in the field, particularly “Spoggy”
Wyndham and Ray Williams. The work at Sturt National Park was accomplished with the
permission and co-operation of the National Parks and Wildlive Service of N.S.W. The
author was in receipt of a University of N.S.W. Post-doctoral Research Fellowship. Identifi-
cation of specimens was achieved with the kind assistance of Peter Aitken (S.A. Museum)
and Jack Mahoney (University of Sydney). The work was partially funded by a grant from
the Department of the Environment and Conservation.

Aust. Zool. 18(3), 1975 193

M. J. S. DENNY
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and Robertson Pty. Ltd., Sydney.

BRIESE, L. A. & M. H. SMITH (1973). Competition between Mus musculus and Pero-
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CALABY, J. H. (1966). Mammals of the Upper Richmond and Clarence Rivers, New
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CALABY, J. H. (1971). Man, fauna, and climate in Aboriginal Australia. In “Aboriginal
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CALDWELL, L. D. & J. B. GENTRY. (1965). Interactions of Peromyscus and Mus in a one-
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HAYNE, D. W. (1949). Two methods for estimating populations from trapping records.
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JOHNSON, D. H. (1964). Mammals, in “Records of the American-Australian Scientific
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JONES, F. Wood. (1923). The Mammals of South Australia. Pt. 3. The Monodelphia.
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KEAST, A. (1968). Evolution of mammals on Southern continents [V Australian mammals:
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MORTON, S. R. (1974). First record of Forrest’s Mouse Leggadina forresti (Thomas, 1906)
in N.S.W. Vict. Nat. 91, 92-94.

NEWSOME, A. E. (1969a). A population study of house mice temporarily inhabiting a
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NEWSOME, A, E. (1969b). A population study of house mice permanently inhabiting a
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PARKER, S. A. (1973). An annotated checklist of the native land mammals of the
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MAMMALS OF STURT NATIONAL PARK
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STACE, H. C. T., G. D. HUBBLE, R. BREWER, K. H. NORTHCOTE, J. R. SLEEMAN,
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TROUGHTON, E: Le Gay. (1928). A new genus, species and subspecies of Marsupial
Mice (Family Dasyuridae). Rec. Aust. Mus. 16, 281-288.

WATTS, C. H. S. (1972). Handbook of South Australian Rodents and Small Marsupials.
Field Naturalists’ Society of S.A. (Inc.).

WATTS, C. H. S. & H. J. ASLIN. (1974). Notes on the small mammals of north-eastern
South Australia and south-western Queensland. Trans. R. Soc. S. Aust. 98, 61-69.

Aust. Zool. 18(3), 1975 195

.

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Pia cn i

Observations on the Ecology of the Crayfish
Parastacoides tasmanicus (Decapoda; Parastacidae)
from South-Western Tasmania

P. S. LAKE and K. J. NEWCOMBE

Department of Zoology, University of Tasmania, Hobart, Tasmania
Urban Biology Unit, Australian National University, Canberra, A.C.T.

ABSTRACT

A typical habitat of the crayfish, Parastacoides tasmanicus, in south-west Tasmania
is described. Data were collected in 1970 on the rainfall, temperatures, burrow
water properties, soil and vegetation in a study area containing a _ thriving
population of P. tasmanicus. Information is given on the structure of the crayfish
burrows and the distribution of occupied burrows in relation to vegetation. The
breeding cycle of the crayfish was determined and observations were made on
the food of the crayfish. A provisional list of animals dwelling in occupied
crayfish burrows is given.

INTRODUCTION

Parastacoides tasmanicus was first described as Astacus tasmanicus by Erich-
son (1846). Subsequently Haswell (1882) named it Astacopsis tasmanicus. Clark
(1936a) in her taxonomic revision of Australian crayfish formed the genus Parasta-
coides and renamed Astacopsis tasmanicus, Parastacoides tasmanicus. The endemic
Tasmanian genus Parastacoides has been the subject of considerable taxonomic
attention, notably by Clark (1936a, 1939), Riek (1951, 1967, 1969, 1972) and
Sumner (1971). At present six species of Parastacoides are formally recognized,
though the Honours study of Sumner (1971) with its statistical analysis of
morphological characters suggests that there may be only two distinct species,
P. tasmanicus (Clark) and P. inermis (Clark).

Parastacoides tasmanicus is confined in its distribution to the south-western
corner of the State, where it reaches its greatest abundance in areas of wet button
grass (Gymnoschoenus sphaerocephalus) moors. If one accepts the species defini-
tions of Sumner (1971), the distribution of P. tasmzanicus becomes more extensive
with the focus of distribution being still in the south-west of the State, but with
populations extending to the central west coast and to the central north-western
region of Tasmania (Sumner 1971, J. L. Hickman, B. Knott and P. Suter pers.
comm. ).

In contrast to the taxonomic attention, ecological studies of the freshwater
crayfish of south-eastern Australia have been very limited. Prior to the study of

Aust. Zool. 18(3), 1975 197

P. S. LAKE and K. J. NEWCOMBE

Newcombe (1970), no ecological study had been made of crayfish in the genus
Parastacoides. General observations on the biology of Tasmanian parastacid cray-
fish have been made by Clark (1936a), Smith (1912), Smith and Schuster (1913),
Lynch (1969), Williams (1968, 1974) and Riek (1969, 1972).

The present paper describes some aspects of the habitat and life history of
Parastacoides tasmanicus. The species remarkable tolerance to pH has been
described in another paper by Newcombe (In press).

OBSERVATIONS

The major portion of the data on the habitat of Parastacoides tasmanicus,
the structure of the burrows, population structure and food was gained from the
investigation of a population in an area near McPartlan Pass in south-west Tas-
mania (about 146° 13’ west, 42° 52’ south), (320m., a.s.l.). The study area
contained a thriving population of P. tasmanicus and can be regarded as being

C)
z

(&

TEMPERATURE

Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

MONTHS

Nov Dec

Fig. 1. Graph of mean monthly maximum air temperatures (O—O) and mean monthly
minimum air temperatures (@®—®@) at Strathgordon and burrow water temperatures
(®-_®) in the study area in 1970.

198 Aust. Zool. 18(3), 1975

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

typical of the areas in south-western Tasmania populated by P. tasmanicus. An
area, 91.44m (100 yds) by 91.44 m (100 yds), was marked out and field observa-
tions were made within this area, apart from the weather data which were
obtained from nearby weather stations at the Knob and Strathgordon.

The soil of the study area conformed to the definition of moor podzol peat of
Nicolls and Dimmock (1965) except for the frequent presence of a quartzite
gravel bed, 16 to 90 cms from the surface.

Figure 1 is a plot of the mean minimum and maximum monthly air tem-
peratures for 1970 near the area, along with the mean of burrow water temperature
for each field trip. The burrow temperature was measured at a point 10 cms
down from the burrow entrance. The burrow water temperatures as measured
in three separate field trips (February, April, June, 1970) indicated that there
was very little diurnal change in burrow water temperatures.

RAINFALL (cm )
Ie) N Ww (ee)
oO on oO oO

—
oO

10

Dec Jan " Feb* Mar’ Apr" May’ Jun Jul Aug © Sep’ Oct * Nov’ Dec

MONTHS 1970
Fig. 2. Histogram of the rainfall recorded at Strathgordon in 1970.

Aust. Zool. 18(3), 1975 199

P. S. LAKE and K. J. NEWCOMBE

As indicated in Figure 2, the area is subject to a high rainfall (circa 267 cms
for 1970) with the maximum fall in late winter and spring.

Oxygen concentrations of water collected 10-15 cms down from the burrow
entrances were determined by the standard Winkler procedure. As indicated in
Fig. 3 the lowest oxygen concentration (circa 1 p.p.m. or 11% saturation)
occurred in the hottest month of the year (February) and also at the driest time
of the year. The months of February and March appear to be the period of
greatest stress for the crayfish, with high burrow water temperatures and low
concentrations of oxygen.

The pH of the burrow water was acidic and did not fluctuate through the
year. The mean pH of the burrow system water was 4.4 + 0.3. Calcium con-
centrations in the burrow water are indicated in Figure 4. As can be seen they
range from 0.125 p.p.m. (12.5 weq/L) in June to 0.98 p.p.m. (49 weg/L) in
March. An analysis of burrow water from one field trip in November, 1970, gave
mean values for the water of 5 burrows of:

Na ; 173. pea/L
Ke 17.9 peq/ 1.

More 75 peas 1
Cae 39. peq/ i:
pH : 4.4

60

50

20

PERCENTAGE SATURATION OF .OXYGEN IN WATER

Feb Mar Apr May Jun Jul Aug
MONTHS 1970

Fig. 3. Graph of oxygen levels in burrow water determined on each field trip.

Sep Oct Nov

200 Aust. Zool. 18(3), 1975

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

10

CALCIUM (P. p.m.)
an fon} a @ (vo)

$

ee)

Feb Mar Apr May Jun Jul Aug Sep Oct Nov

Fig. 4. Calcium levels in burrow water determined on each field trip in 1970.

These results indicate that the water chemistry is similar to that described
by Buckney and Tyler (1973 a, b) for waters of south-western Tasmania. The
burrow water is of a deep brown colour.

BOTANICAL DATA ON STUDY AREA

Five distinctive types of plant communities were found in the study area.
The names given to the types of plant communities are not accepted definitions.
Briefly the plant communities are:

1. Button Grass Moor. Large clumps of button grass (Gymnoschoenus sphareo-
cephalus) represent the dominant species and claim about 60% of the available
ground. Sprenglia incarnata, Bauera rubioides, Leptospermum nitidum, Boronia
pilosum, Restio oligocephalus, Xyris sp., were common sub-dominants in this
community.

Aust. Zool. 18(3), 1975 201

\

P. S. LAKE and K. J. NEWCOMBE

2. Button Grass Swamp. The button grass in small clumps claims about
20 to 30% of the available ground. With the clumps also occurs Xyris operculata,
Sprenglia incarnata, Bauera rubioides, Leptospermum nitidum, while in the spaces
between the clumps Leptocarpus tenax, Baeckea leptocaulus, Lepyrodia tasmanica
and Lycopodium sp. occut.

3. Marsh. This community is dominated by Leptocarpus tenax and Gleichenia
microcarpa. Dense entwining patches of Hypolaena longissima are present. In the
understorey, Lycopodium occuts.

4. Heath Sedgeland with Emergent Shrubs. The common species in this
community are Bauera rubioides, Leptospermum nitidum and Gymnoschoenus
sphaerocephalus. Other species present include Sprengelia incarnata, Restio oigo-
cephalus and Boronia pilosa.

5. Swamp. This community harbours a similar species range to that of the
marsh community, with the addition of Meloleuca squarma, Restio complanatus and
Sprengelia incarnata. The area consists of small islands of soil scattered over a.
complex of anastomosing channels. The channels have a characteristic white sandy
substrate.

The communities described for the McPartlan Pass study site resemble some
of those described in the Lake Edgar area by Macphail and Shepherd (1973).
The Button Grass moor community resembles their button grass moor community,
while the button grass swamp resembles their sedge swamp (Gymnoschoenus-X yris
association). The marsh community is somewhat similar to their swamp sedgeland
community (Lepidosperma-Leptocarpus-X yris association, ) with the difference being
an abundance of Gleichenia and an absence of Lepidosperma at McPartlan Pass.
The heath sedgeland with emergent shrub community of this study is similar to
that of the same name in MacPhail and Shepherd (1973), while the creek com-
munity of this study shows a similarity to their sedge swamp community.

DISTRIBUTION OF CRAYFISH IN RELATION TO PLANT COMMUNITIES

The various plant communities undoubtedly reflect varying levels of soil
moisture, free water availability, water table levels and extent of soil develop-
ment. Such factors can be expected to be of major importance in determining the
distribution of crayfish. Within the study area, over a period of 25 days in August-
September, 1970, a planned excavation of ten 9.144 m (10 yds) by 9.144 m
(10 yds) plots, chosen at random, was carried out. All the burrows in each plot
were dug up, the position of the burrow recorded and all the animals in the
burrows collected.

As can be seen from Figure 5, the crayfish were largely confined to the
three wetter plant communities; button grass swamp, marsh and swamp. Visual

inspection revealed no burrows in the small area of heath sedgeland with emergent
shrub community. While these data may apply for the area investigated, in many

202 Aust. Zool. 18(3), 1975

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

other areas P. tasmanicus occurs in a considerable diversity of wet plant com-
munities ranging from closed scrub to rain forest gullies.

STRUCTURE OF BURROWS

A knowledge of burrow structure came from the excavation exercise and
from the extensive sampling of P. tasmanicus populations that has been carried
out by members of the Zoology Department in recent years in south-western
Tasmania. A diagrammatic depiction of a typical burrow is presented in Figure 6.

The structure and depth of the burrow appears to be dependent on the
amount of water available and the depth of the water table. In more swampier
areas, with free surface water available, the burrow entrance is often found at
the bottom of a pool, usually tucked underneath overhanging vegetation. Often

Fig. 5. Distribution of plant communities in the study area. The squares represent randomly
selected areas which were excavated. Dots represent the occurrences of. crayfish in
the excavated areas.

Button grass moor.

Button grass swamp. Y Heath sedgeland with emergent shrubs,

Marsh. eS Swamp,

Aust. Zool. 18(3), 1975 203

bs} LJ [24

P. S. LAKE and K. J. NEWCOMBE

iN / AW \\|

i! mM

/
; -

‘
1

‘

NV AN
I\ j ,
i hy ie! ”
r / | W/o ul j
5 LAY f fas :
C7,

1
\

s JRL
oY

| CHL

Fig. 6. Diagrammatic transverse section of a typical burrow system occupied by one adult
Parastacoides tasmanicus.

two or three holes may open to the surface, but these usually meet below the
surface. From the major burrow entrance, a branch may sometimes be found,
leading to a small chamber beneath a large clump of vegetation. On drier soils,
the burrow entrance is frequently found in clumps of vegetation.

The main burrow from the entrance invariably in going deeper takes a
number of twists and turns and finally terminates in all systems, in a bottom
chamber.

Where there is a quartzite gravel layer, the level of the bottom chamber is
fixed, for the crayfish do not penetrate deeper than 1 to 2 cms into the quartzite
layer. In drier areas, the burrows are deeper, because the water table is deeper.
The burrows were from .4 to 1 metre deep in the study area. In the tunnel
system of the study area, the burrows usually open at the base of vegetation clumps
and are not very deep (7.7 to 15 cms). Recent digging activity in burrows may
be indicated by shall chimneys, but these are never as well developed or as large
as those created by species of Engaeus.

In the study area in the August-September excavation 185 burrows were
dug up. The percentage of burrow occupancy was 83.5%. Of the burrows dug up,
only four burrows were found which were interconnected. In each of these cases
the interconnecting passages were full of mud and detritus and may be regarded
as being disused. The relatively large diameter of the burrows suggest that adults

204 Aust. Zool. 18(3), 1975

NUMBER OF SPECIMENS COLLECTED

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

carry out the active digging. Signs of small surface holes indicating possible digging
by young crayfish were very uncommon. Young animals may build their burrows
de novo, but it seems more likely that they occupy vacated adult burrows.

As a tule, there is one crayfish per burrow system, except for the case of
females which have recently shed their young, in which case up to ten young
hatchlings may be found in the burrow system. Adult individuals if kept together
in captivity are very aggressive to one another. Leggett (1971) has described
the agonistic encounters between individuals. If kept in a pair, the larger animal
invariably wins.

15

14

41-5 9-1-10 141-15 171-18 224-23 281-29 301-31 341-35
| CARAPACE LENGTH (mm)

Fig. 7. Histogram of number of crayfish collected in the excavation of ten areas of the
study area against carapace length.

Aust. Zool. 18(3), 1975 205

P. S. LAKE and K. J. NEWCOMBE

POPULATION STRUCTURE AND DENSITY

In the excavation of ten plots in August-September, 1970, 154 crayfish were
dug up. For the study area as a whole, at that time, this gives an upper population
limit of 2,316 and a minimum population limit of 764 at the 95% confidence
level. In terms of density, there is a maximum density of 1 animal per .985 m/?.
A notable feature of the population at that time was the dominance of the
population by large adults (22 to 35 mms carapace length). As the burrow
sampling was very thorough, it is highly unlikely that many small individuals
were missed. At the time of excavation, one could have reasonably expected to
find a considerable number of small juveniles in the population, as a result of
the young that presumably left their mothers in February-March of that year.
This suggests that the population consists of slow growing mature adults (Figure
7) and that there is a high mortality of the juveniles each year with only a small

fraction of them surviving the first year. The sex ratio in the study area was
40% males and 60% females.

BREEDING

From dissections of females collected in the field and on the basis of those
carrying eggs, it appears that females in the study area become sexually mature
at a carpace length of at least 24 mms. It appears that larger individuals with a
carapace length from 32 to 36 mms are the most fertile. However, the percentage
of females judged on the basis of carapace length to be sexually mature but which
did not carry eggs was quite high. Only 40% of the eligible females carried eggs.
Females of the eligible carapace length, dissected in September, contained mature
but unfertilized oocytes. The relatively low level of fertilization in the population
may partly explain the female-biased sex ratio.

The number of eggs carried per female ranged from 38 to 80, and the
number of eggs carried per female increases linearly with increase in carapace
length. Where y = number of eggs and x = carapace length in mms, the
relationship between them is of the form, y = 3.86x — 61.11 with b having
95% confidence limits of — 7.30 and + 14.02.

As can be seen from Table 1, of the total number of eggs carried per female,
the fraction carried per pleopod pair increases toward the tail, with the 2nd, 3rd
and 4th pairs each carrying almost the same fraction of eggs.

Females carry eggs over winter from April through to November. The
hatchlings then remain attached to the pleopods of the females through to
February-March. The hatchlings are attached to the female’s pleopods by their
last two pairs of pereiopods. The young leave the parent individually and not as
a single group. Their carapace length at departure ranges from 3.5 to 3.8 mm.
In captivity in a small container the young are at great risk from being eaten
by the mother and they seek any shelter available. In the burrows, the young leave
the mother and presumably flee from her and seek cover. When one digs up a

206 Aust. Zool. 18(3), 1975

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

burrow containing a female with young leaving her it is rare to find the young
in close proximity of the mother.

TABLE 1

Distribution of eggs in relation to pleopod pair.

First pair Second pair | Third pair Fourth pair |
Observed Observed pay es Total
. numbers = number number number
Animal No. (Decimal (Decimal (Decimal (Decimal number
fraction fraction fraction fraction of eggs
of total) of total) of total) of total)
1 11 18 15 27. 71
(.15) (.25) (21) (.38)
Di 13 15 18 16 62
(21) (.24) (29) €26)
3 10 17 20 15 62
CLG) C27) (.32) (.24)
4. 14 pag 21 17 iis)
(.19) (.29) (.29) (23)
5: 10 17 13 20 60
(.17) (.28) @22) (.33)
6. i 15 17 17 60
(.18) (25) (.28) (.28)
Te 14 14 14 43
(.02) (33) (33) (33)
8. 13 10 10 42
C21.) (31) C24) (.24)
9. 10 14 Ps on 66
(15) (.21) (.32) (.32)
10 8 11 13 15 47
Cig) (22:3) (.28) (32)
Mean fraction (.16) C21) (.28) (.29)

The young appear to remain in the burrow system of the mother until the
level of the water table rises with the rains and decreased evapo-transpiration in
autumn and early winter. At this time of the year young crayfish may be picked
up in hand-nets in small pools, tunnels and depressions. The young may be found
in surface water under debris, stones, etc., or in unoccupied burrows. The repro-
ductive cycle of the females largely agrees with the annual moulting cycle of
P. tasmanicus meticulously determined by Mills (1973). Moulting was observed
in March-April, which would, in females, be prior to oviposition. and after the
young had left. :

Foop

Ten specimens fixed in the field on field trips in March, June, August and
October had the contents of the gastric mills examined. From this examination
a list of food items may be compiled.

Aust. Zool. 18(3), 1975 207

P. S. LAKE and K. J. NEWCOMBE

1. Phylum Arthropoda:
Class Insecta:

O. Hymenoptera,
F. Formicidae.

Pheidole sp. (only in one crayfish).

2. Order Coleoptera. Adults and larvae. (In 3 crayfish).

3. Class Crustacea:

O. Amphipoda,
Family Gammaridae.

Neoniphargus sp. (Only i

4. Phylum Annelida.

n one crayfish).

Class Oligochaeta. Setae regularly found.

5. Plant Material: By far the greatest amount of material in the gastric mills
was plant material and by far the greatest amount of the plant material

consisted of root material.

6. Miscellaneous: All gastric mills contained fine quartzite granules.

Thus, animal material is consumed by the crayfish, when this is available.
Plant material is digested by the animals and forms the major part of their diet.
Root material is the dominant type of plant material. There is good evidence for
a cellulase enzyme being present in the crayfish gut (Newcombe 1970).

ANIMALS LIVING IN OccuPIED PARASTACOIDES TASMANICUS Burrows

For this part, the assistance of Mr. B. Knott and Dr. J. L. Hickman of the
Zoology Department, University of Tasmania, was invaluable. From many field
trips involving the digging up of P. tasmanicus burrows, a list of animal species
in the burrows was compiled. Earthworms (Megascolidae) may be found in
the soil surrounding the burrows, but in the water of the burrows all the macro-

scopic animals so far collected are crustaceans.

Class Crustacea

Order Copepoda
§ub-Order Cyclopoida

i?

One species
mania).

Order Syncarida

Ze

208

Micraspides calmani (Crotty).

. Allanaspides helonomus (South-

West Tasmania).
Allanaspides hickmani
West Tasmania).

(South-West Tas-

(South-

Order Isopoda
Sub-Order Phreatoicidea

a

6:

Family Phreatoicoideae
Undescribed genus. One sp.
(South-West Tasmania).
Colubotelson sp. (Same as large
species on Pedder Beach (Bayly
1973):

Family Amphisopidae
Phreatoicoides sp. 1 (Olga River
Valley).

Aust. Zool. 18(3), 1979

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

8. Phreatoicoides sp. 2 (Crotty). 13. Neoniphargus sp. 3 (blind)
Asellota (South-West Tasmania and West
Family Janiridae Tasmania )

9. Undefined Genus — possibly 14. Neoniphargus sp. 4 (blind)
Heterias (South-West Tasmania ) South-West Tasmania )

10. Pseudasellus sp. Unpigmented 15. Undescribed genus. Blind animals
(West Tasmania e.g. Crotty) very similar to Niphargus pul-
chellus (Crotty )
Order Amphipoda 16. Undescribed genus. Very similar
Family Gammaridae to Gammarus australis (South
11. Neoniphargus sp. 1 (eyed) large West and West Tasmania)
(South-West Tasmania ) Family Eusiridae

12. Neoniphargus sp. 2 (eyed) small 17. Paracalliope sp. (Crotty)
(South-West Tasmania and West 18. Paraleptamphopus sp. (South
Tasmania ) West and West Tasmania)

It is very surprising that no insect adults or larvae occur in the burrows.

DISCUSSION

A brief description of some major features of a typical habitat of P. tasmanicus
has been given. From the distribution of the species in Tasmania it is clear that
it is restricted to areas of high rainfall and high water table level. Its distribution
closely coincides with the distribution of tussock sedgeland and sedge swamp
plant communities, where these communities have buttton grass, Gymnoschoenus
sphaerocephalus as one of the dominants. As opposed to the judgment of Riek
(1972), we do not regard P. tasmanicus as having streams and lakes as part of their
normal habitat. All members of the genus Parastacoides are found in swampy
ground; occasionally young individuals may be found in streams. One population
of mature P. tasmanicus has been found in a small creek near Penguin in northern
Tasmania in early December, 1973 (B. Knott and P. Suter, pers., comm.). Such
a situation is highly unusual.

Previous accounts of parastacid crayfish burrow structure have been few.
The depth of the burrows of P. tasmanicus appears to be primarily determined
by the depth of the water table in the driest period of the occupied burrow’s |
life. Consequently it varies considerably from locality to locality. Parastacoides
tasmanicus has a burrowing ability, as has been often reflected upon by the authors
in wet swamps under inclement conditions. We would not regard, as Riek (1972)
does, Parastacoides as a ‘moderate burrower’, or Geocharax for that matter,
especially in comparison to Riek’s (1972) definition of ‘strong burrowers’ in the
genus Engaeus. In our opinion, burrowing ability as opposed to burrow structure
and choice of burrow site does not distinguish Parastacoides from Engaeus in
Tasmania. The structure of engaeid burrows has been noted by Smith and Schuster
(1913), Clark (1936 a&b), Riek (1969, 1972), Clark (1936b) noted that some

Aust. Zool. 18(3), 1975 209

P. S. LAKE and K. J. NEWCOMBE

species of Engaeus, e.g. E. fossor, have solitary burrows while other species, e.g.
E. victoriensis, form ‘community’ burrows. Riek (1969) describes some engaeids
as living in family groups; with a family consisting of ‘a mature pair of individuals
and juveniles of one or two age groups’. Parastacoides lives in solitary burrows, ©
and as observed by Leggett (1971) in the laboratory, individual Parastacoides
when kept together are very aggressive to one another. Such aggression between
individuals appears to greatly diminish any likelihood of social community or even
grouping. |

For Engaeus fossor in Tasmania it is our impression that this species also
normally lives in solitary burrows and thus does not conform to the pattern of
burrow occupancy ascribed to engaeids by Riek (1972). The same applies for
E. leptorhyncus. Prior to this study very little had been published on the life
history of parastacid crayfish. Clark (1936 a, b) concluded that the breeding
season is in spring, after moulting. Hatching of the young, she concluded, occurred
in December and January. Breeding and development in Euastacus kershawi has
been described by Clark (1937) and aspects of the life history of the Marron,
Cherax tenuimanus, have been described by Shipway (1951 a, b), Morrissy (1970
a&b, 1974). In New Zealand, Hopkins (1966, 1967 a&b) has described growth
and breeding in the stream dwelling Paranephrops planifrons. The domination of
the P. tasmanicus population by mature adults suggests that the population size
is very stable and that both recruitment to the population and adult mortality are
low in relation to the population size. A high mortality of juveniles can be expected.
A situation of density limitation of recruitment is suggested for the marron
(Morrissy 1970 b, 1974), and also for unexploited populations of Astacus astacus,
Abrahmasson (1966).

P. tasmanicus females carry eggs over winter; the eggs hatch in spring (Novem-
ber) and the young leave the parent in February or March. Clark (1936 a&b)
thought that for Tasmanian and Victorian parastacids the breeding season was a
short one from spring to early summer (16 weeks or so). She overlooked the
long over-wintering period of egg incubation. Hopkins (1967 a) found in Parane-
phrops the incubation of eggs extended from April to November and that the
young left the mother in early December. Thus the breeding period for Parane-
phrops is long (25-26 weeks). However, as opposed to the situation in Parane-
phrops and Parastacoides, the marron Cherax tenuimanus appears to have a short
breeding season covering spring and early summer (Morrissy 1970 b). Possibly
the short breeding season may be due to high rate of larval development induced
by relatively high water temperatures (bottom temperatures from 14-17°C in
October and 16-25°C in December, Morrissy 1974), as opposed to the slow rate
of egg and larval development in the low water temperatures experienced by
Parastacoides and Paranephrops.

The majority of northern hemisphere astacid crayfish have a short breeding
season usually occurring over spring and early summer, e.g. Orconectes rusticus
(Langlois 1937), Cambarus longulus (Smart 1962), O. obscurus, O. sanborni,

210 Aust. Zool. 18(3), 1975

ECOLOGY OF CRAYFISH PARASTACOIDES TASMANICUS

O. propinquus (Fielder 1972), O. virilis (Weagle and Ozburn 1972), or for a
short period in late summer and autumn, Procambarus clarkii (Penn 1943), Pro-
cambarus hayi (Payne 1972). A relatively long breeding season involving egg-
carrying over winter and the release of young in late spring has been found for
European species of Astacus and for the North American crayfish, Pacifastacus
leniusculus (Riegel 1959) and P. trowbridgei (Mason 1970). The method of
attachment of the eggs to the female pleopods in parastacids has been described
by Clark (1937) and Hopkins (1967 a). The number of eggs carried per pleopod
pair in P. tasmanicus increases posteriorly.

This differs from the pattern in Paranephrops (Hopkins 1967 a), where the
greatest number of eggs are carried on the second and third pair of pleopods. The
number of eggs carried by P. tasmanicus females varied from 36 to 79, which is
higher than the 20 to 30 noted by Riek (1967). As for other crayfishes (e.g.
Hopkins 1967 a, Abrahamsson 1966, 1971, Weagle and Ozburn 1972), the
nuiber of eggs carried per female increases linearly with increase in length of
the female.

A number of observations have been made on the food of parastacids. Smith
and Schuster (1913) noted that Engaeus species probably had a carnivorous diet.
They found earthworms, insect larvae and ‘probably land crustacea’ in their gastric
mills. Clark (1936 a&b) noted that in captivity species of Ewastacus, Cherax and
Engaeus ate earthworms, raw meat, fish and tadpoles. In their gastric mills Clark
(1936 a) found ‘mud and debris’. Clark (1936 b) also noted the propensity for
cannibalism in captive parastacids. In the marron, Cherax teniumanus, Shipway
(1951 a&b) reported that they are omnivorous, while Morrissy (1970 b, 1974)
found that the gastric mills of marron in farm ponds contained allochthonous plant
material. Shireman (1973) found that marron in captivity preferred crayfish tails,
iresh fish and catfish pellets in that order. They also ate plant material. In astacids,
snere is considerable evidence to support the idea that crayfish eat considerable
amounts of plant material (e.g. Tack 1941, Bovbjerg 1952, 1970, Abrahamsson
1966). In Parastacoides tasmanicus plant material, especially root material, domin-
ates the diet, though oligochaetes, some insects and amphipods may be eaten.
Newcombe (1970) found good evidence for cellulase being present in the gut of
P. tasmanicus. Whether this enzyme is produced by the cells of the crayfish or
by symbionts is not clear. In both amphipods (Wildish and Poole 1970) and
Procambarus clarkii (Yokoe 1960) cellulase has been found, and in both cases
it appears that the cellulase is both of symbiotic and animal cell origin.

From the list of macroscopic animals found in P. tasmanicus burrows it
appears that only crustaceans are found, in spite of the fact that oligochaetes,
insects and molluscs are to be found in the surface pools of areas with high
crayfish populations. The crustaceans found with P. tasmanicus have presumably
evolved means to live a subterranean existence and also to evade or offset crayfish
predation. There must be a strong selection pressure to push the crustaceans in
such an adaptive direction. Possibly such a pressure is the need to survive in

Aust. Zool. 18(3), 1975 211

P. S. LAKE and K. J. NEWCOMBE

dry periods, when in the button grass swamp areas of South-West Tasmania, the
amount of standing surface water is very greatly reduced. At such times insects,
oligochaetes and bivalve molluscs may survive by burrowing into the mud. In
the case of some insects, this period may be spent in the adult phase. However,
the requirement for continual free water by the crustaceans has forced them to
adapt to living in the only standing free water readily available; that in the burrow
systems of Parastacoides tasmanicus.

ACKNOWLEDGEMENTS

We are very grateful to Mr. B. Knott, Dr. J. L. Hickman and D. Coleman of the
Department of Zoology, University of Tasmania, for very useful information on parastacid
taxonomy and biology and for invaluable field observations. We are also grateful to Mrs.
N. Dorney for drawing the figures and Mrs. T. Vasos for typing the manuscript.

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214 Aust. Zool. 18(3), 1975

The Identity of the Common Keyhole Limpet of
South-Eastern Australia (Fissurellidae)

By W. F. PONDER
The Australian Museum, College St., Sydney, N.S.W. 2000.

The common, large species of Diodora from south-eastern Australia has been
known since 1924 as Elegidion audax Iredale (1924). This binomen was intro-
duced by Iredale to replace Fissurella lineata Sowerby, 1835, described from
unknown locality, and its synonym F. imcii Reeve, 1850 which was said to have
been collected from ‘“‘Raine Island, Torres Straits”. Examination of material in
the British Museum (Natural History) has shown that specimens of F. incii were
wrongly localised and agree exactly with shells from New South Wales (compare
figs. 1-6 with 7-15). There is also no known species in the Torres Straits north-
eastern Queensland area that closely resembles F. imciz (i.e. lineata).

Iredale (1924) argued that a new name was warranted on the basis of
locality (in the case of zmcii) and on a point in Sowerby’s (1835) description (in
the case of lineata). Sowerby described the internal margin of the foramen (i.e.
of the surrounding callus) as being truncated posteriorly in F. lineata and Iredale
states that the southern Australian shell does not show this feature. However,
Iredale’s figured syntype (figs. 9-11) (a small specimen), and most small speci-
mens of this species, do show a posterior truncation of the callus. Because Sowerby’s
original figure (which is presumably actual size) is ony 23.3 mm long it would
also be expected to show this feature.

A brief synonymy of F. lineata appears below.

Diodora (Elegidion) lineata (Sowerby, 1835). Plate 1, figs 1-15.

Fissurela lineata Sowerby, 1835: 7, pl. 78, fig. 68; Sowerby, 1866: 195, pl. 241,
fies. 134. 135.

Fissurella-incii Reeve, 1850, pl. 10, figs 69a, 8.
?Fissurella australis; T. Woods, 1877: 44 (non Krauss, 1848).
Glyphis lineata; Hedley, 1900: 95, -pl. 3, fig. 11 (animal).
Fissurella mccoyi (T. Woods MS) Pritchard & Gatliff, 1903: 185 (nomen nudum).
Elegidion audax Iredale, 1924: 220, pl. 35, figs 5, 6.
Eligidion (err.) audax; Macpherson, 1966: 207.
It appears that the types of Sowerby’s (1866) figured specimen

of Eo eda orebably came itrom:, thetisame lot. as i. imen
and both names appear on the labels with the material. None of the

Aust. Zool. 18(3), 1975 215

W. F. PONDER

specimens in the British Museum (Natural History) agree exactly with the
figures given by Sowerby or Reeve, but in the case of Sowerby’s figures these
were often composite. One specimen closely fits the dimensions of one of the
original figures of F. incii and all agree well in all but individual, minor details.
There is little doubt that the specimens examined are part of the original series
of F. incii and can be regarded as syntypes of F. incii. The types «of D. lineata
(Sowerby) appear to be lost. They were 2 specimens from the G. Humphreys
collection and Sowerby (1835) states that they were “now in Mr. Cuming’s (col-
lection)”. No trace of these can be found in the British Museum, but the original
figure is a good one and there can be little doubt as to its specific identity.

| Length of
Dimensions: _ Length Width - Height foramen

Figured syntypes of lineata 44.82 mm 29.60 mm 16.93 mm 3.84 mm
38.22.mm 23:86 mm’; 12.80 min 3.46 mm
Syntypes of E. audax

Iredale
largest 54.60 mm 35.20 mm 23.84 mm 4.62 mm
Iredale’s figured
specimen 18.17. mm 11.06: mm 5.76 mm 1.94 mm

Range: Burleigh Heads, S. Qld. (C. 98620) to Port Phillip Bay (Macpherson,
1966) and Flinders, Vic. (Burn, 1959). T. Woods (1877) recorded the South
African species Fisswrella australis Krauss from Tasmania and this was regarded
by Tate & May (1901) as being a misidentification of lineata. May (1921),
however, rejected the Tasmanian record of lineata.

Elegidion Iredale, 1924 is used as a subgenus of Diodora Gray, 1821 following
Knight et al. (1960). The external appearance of the living animal is described by
Hedley (1900) and Burn (1959).

ACKNOWLEDGEMENTS

I am grateful to the staff of the Mollusc Section, British Museum (Natural History) for
their help during my visit. This work was supported by travel funds provided by the Ian
Potter Foundation, the British Council, the Australian Research Grants Committee, the
Science and Industry Endowment Fund, and the Trustees of the Australian Museum. I am
particularly grateful to my wife for her assistance and for taking the photographs of the
types in the British Museum. The other photographs were taken by Mr. G. Millen of the
Australian Museum.

216 Aust. Zool. 18(3), 1975

THE IDENTITY OF THE COMMON KEYHOLE LIMPET

Plate 1. Diodora (Elegidion) lineata (Sowerby).
Figures 1-6 Probable syntypes of Fissurella incii (British Museum (Natural History
registered number 197546).
7-11 Syntypes of Elegidion audax Iredale Twofold Bay, N.S.W. (C. 99178).*
12-13. Balmoral, Sydney Harbour, N.S.W. (C. 76384).
14-15 Kurnell, Botany Bay, N.S.W. (C. 88654).
* Australian Museum registered number.

Aust. Zool. 18(3), 1975

W. F. PONDER

REFERENCES

BURN, R. (1959). Molluscan field notes. Part 3. Victorian Naturalist, 75, 179-181.
HEDLEY, C. (1900). Studies on Australian Mollusca. Proc. Linn. Soc. N.S.W. 25, 495-513.

IREDALE, T. (1924). Results from Roy Bell’s molluscan collections. Proc. Linn. Soc.
N.S.W., 49, 179-278.

KNIGHT, J. B., L. R. COX, A. M."KEEN, R. L. BATTEN, E. L. YOCHELSON, R.
ROBERTSON (1960). Archaeogastropoda. Treatise on Invertebrate Paleontology /,
169-310. 7

MACPHERSON, J. H. (1966). Port Phillip Survey 1957-1963. Mollusca. Mem. Nat. Mus.
Vict., 27, 201-384.

MAY, W.:<L. (1921). A checklist of the Mollusca of Tasmania, Govt. Printer, Hobart.

PRITCHARD, G. B. & J. H. GATLIFF. (1903). Catalogue of the marine shells of Victoria.
: Part'6) Prog. Re Socs Vict>(vs;) 19. (2); 76-223.

REEVE, L. A. (1850). Monograph of the genus Fissurella. -Conchologia Iconica, 6. London.

SOWERBY, G. B. (2nd), (1835). A catalogue of ine Recent species of Fissurella. The
Conchological Illustrations, London.

SOWERBY, G. B. (lst, 2nd & 3rd), (1862). A monograph of the family Fissurellidae.
. ‘Thesaurus Conchyliorum. Vol. 3. London.

TATE, R. & W. L. MAY. (1901). A est census of the marine Mollusca of Tasmania.
Proc. Linn. Soc. N.S.W., 26, 344-471.

WOODS, J. E. TENISON. (1879). Ganois: with brief descriptions of the marine shells of
Tasmania and the adjacent islands. ‘Pap. Proc. R. Soc. Tasm., 1877, 26-57.

218 Aust. Zool. 18(3), 1975

Memorial Notice and Bibliography
by the late GILBERT P. WHITLEY

BEETS Ley GEYT TROUGHTON, G:M.Z'S. F.R.Z.S. (1893-1974)

The death, at Concord, Sydney, on 30 November 1974 of Mr. E. Le G.
Troughton ended the career of one who had served longer on the scientific staff
of the Australian Museum than any other person in that institution’s long history.
He was the first full-time Curator of Mammals in any museum in Australia, a
staunch conservationist, and a sociable friend of considerable charm. For over
fifty years we were colleagues, working in adjoining rooms, so here I try to record
some recollections of his life and work, albeit inadequately, so that he may be
placed in his deserved niche in the history of zoology in Australia.

Ellis Le Geyt Troughton was born at Oxford, Kellett Street, Darlinghurst,
Sydney on 29 April 1893. His birthplace is unrecognizable now, having been
demolished for the King’s Cross extensions to the Eastern Suburbs Railway.
The Sydney Morning Herald (4 May 1893: 1) announced his birth to the wife
of Donald E. Troughton, police magistrate, of Moree. His mother’s maiden name
was Elizabeth M. Ryrie. His boyhood haunts have also been largely destroyed
by the inundation of the countryside for the Jindabyne Dam, New South Wales.
Troughton was educated at West Bush Public School.

Troughton was one of several young Cadets appointed to the Australian
Museum early in 1908. He was a general assistant at first, helping Hedley to
label mollusca, many of which were mounted on nearly 25,000 tablets at that
time. Next Troughton worked in the Department of Ethnology (the term Anthro-
pology was not used in those days) and then Allan McCulloch trained him in
taking care of the collections of all the vertebrates, in cataloguing literature, and
in field work. Troughton helped McCulloch with the editing of the first part of
the Australian Zoologist sixty years ago. Robert Etheridge was Director of the
Australian Museum when Troughton joined the staff. There was no superannuation
in those days and working conditions were poor, like the pay! Troughton was
told that it was a privilege for him to hold a “‘gentleman’s job” and that he was
lucky to receive a salary of ten shillings (nominally one dollar) a week. Over
the years, Troughton was to fight for the betterment of the underdogs. In 1909
and 1910 he studied zoology at Sydney Technical College and at the University of
Sydney under Professor Thomas Harvey Johnston and passed the required exami-
nations. ““By way of a stimulus” the fees of successful students were refunded;
otherwise cadets had to pay their own way. On 1 January 1913 Troughton was
promoted to the permanent scientific staff as a Second Class Assistant, until he
enlisted in the Army (First A.I.F.) in July 1916. He served as a stretcher-bearer

Aust. Zool. 18(3), 1975 219

GILBERT P. WHITLEY

Ellis Le Geyt Troughton, C.M.Z.S., F.R.Z.S. in 1973.

Photograph by courtesy of the Linnean Society of New South Wales

220 Aust. Zool. 18(3), 1975

MEMORIAL NOTICE and BIBLIOGRAPHY

in the Fourth Australian Field Ambulance on the battlefields of France in 1917-
1918. He visited London when on leave and was hospitably received at the
British Museum (Natural History), where he studied Australian specimens, as he
also did in the museums of Paris, Cologne and Monaco. On his way back to
Sydney, he visited the museums of Cape Town and Durban and those of the
mainland Australian States. He then became the first full-time Curator of mam-
mals in any museum in Australia, when in October 1919, he was placed in charge
of a new Department [of Mammals and Skeletons] in the Australian Museum. A
year or so later he was promoted to First Class Scientific Assistant. He had already
been largely responsible for the fine display of bleached skeletons against a plain
black background in the public gallery.

The Records of the Australian Museum, part 3 of volume 13, published in
December 1920, was facetiously referred to as the ‘“‘Children’s Supplement’’,
because the early efforts of several young men on the staff therein saw the light
of day: these included Troughton’s ‘Notes on Australian Mammals, No. 1”. His
writings are fully listed in the Bibliography below. On the proposal of Professors
T. Thomson Flynn and J. P. Hill, Troughton in 1936 was given the honorary
status of Corresponding Member of the Zoological Society of London (C.M.Z.S.).
Though writing did not come easily to him, Troughton was the author of many
papers and popular articles on mammals. The book Troughton’s Furred Animals
of Australia recently passed its ninth edition and Iredale & Troughton’s Check-list
of the Mammals recorded from Australia (1934) still has not been supplanted.
Troughton discovered numerous unnamed rats (studied in connection with Weil’s
Disease in Queensland and scrub typhus in Papua New Guinea), many bats, some
whales and several marsupials from Australia and the Pacific. He has gone into
historical details regarding the identity of Captain Cook’s Kangaroo, the Grampus,
Koala, New Guinea Dog, and other noteworthy mammals. Troughton revised
Professor W. A. Haswell’s mammal articles for the first edition of the Australian
Encyclopaedia, to which he added one of his own, and he wrote a number of
entries for the second edition. Though he worked principally on rats and mar-
supials, the bats seem to have been his favourite subject.

In 1922, he reported on Australian seals and sealing for the Commonwealth
Institute of Science and Industry, appreciation of which was expressed by the then
Prime Minister (Hon. W. M. Hughes). Another report was on “matters arising
out of the invasion of Lord Howe Island by rats’, at the request of the Lord
Howe Island Board of Control (Chief Secretary’s Department, Sydney) in 1923.
Troughton proposed that owls be introduced to reduce the rats (a course of action
induced by his observations on these birds and the rats on the Nullarbor Plain)
and his recommendations were adopted. His curatorial work at the Museum
involved care of human crania, whales, horses, cattle, sheep and all placental,
marsupial and monotreme specimens, and the skeletons of all vertebrates. He
supervised the setting up of habitat groups and gallery displays and attended
to public enquiries. The latter included identification of human remains for
Police; rats, etc., for the Health Department, and advising the Chief Secretary’s

Aust. Zool. 18(3), 1975 221

GILBERT °-P.. WHITLEY

Department concerning animal protection, open seasons and the issuing of
collecting permits. He was on Government committees concerning Koala Park
and on fur-farming and one formed to advise State and Commonwealth Govern-
ments on the export of animals through H.M. Customs. He was a member of the
Green Committee concerning confiscated birds’ eggs which came to the Australian
Museum, and of the Grant Bird Collection Committee of 1926. He delivered
many lectures and broadcasts.

Troughton joined the Royal Zoological Society of New South Wales in 1920
and was elected a Councillor in 1925. He was President in 1931-1932 and a Fellow
in 1940. He was elected a member of the Linnean Society of New South Wales
in 1921, became a Life Member, was on the Council from 1939 to 1972, President
1943-1944 and Vice-President 1944-1947. In 1928 he was a member of the Aust-
ralian National Research Council. In 1932, as Secretary of Section D (Zoology) of
ANZAAS, he sponsored a resolution that the Federal and State Governments
conduct a biological survey of Australian fauna and flora. At the Auckland Congress
(1937) he was Secretary of the Biological Survey Committee of ANZAAS whose
recommendations about a decade later led to the establishment of the Wildlife
Research Division of CSIRO. Troughton was a foundation member of the Fauna
Protection panel of New South Wales, since replaced by the National Parks and
Wildlife Service, serving on the panel from 1949 to 1963.

During the final phases of the Second World War, Troughton served in New
Guinea with the Tropical Scientific Section, A.I.F., investigating the mammal
reservoirs of scrub typhus, on the recommendation of, and reporting to, the
U.S.A. Typhus Commission. I have not traced a copy of his official report, which
was probably a “restricted” publication, but Mr. Troughton told me that no new
species were named therein. Troughton had received medals for his army services
in two World Wars: General Service and Victory Medals and Pacific Star and
War Medal 1939-45.

“Troughtie”’, as he was known to his many friends, was of a mercurial
disposition, very fond of company, and a member of numerous clubs, societies
and sporting bodies. He was suntanned and wiry and had played much tennis,
enjoyed climbing at Lord Howe Island, and kept fit at gymnasium in his younger
days. He took part in swimming races at the Bondi Diggers’ Club gatherings,
and rode horseback on his field work in Mount Kosciusko National Park. He was
a prominent member of a Lodge of Freemasons.

When so many younger people were away on war work, Troughton returned
from New Guinea to find himself in charge of all the vertebrates again and referred
to himself, with characteristic good humour as the Mammalornithoichthyherpeto-
logist! In those days, curators were officially classed as ‘“‘scientific assistants”, a
disparaging designation and always a handicap. Then we were known as ichthyo-
logists, mammalogists, etc., which always had to be spelt over the telephone. In
1948, mercifully, the term was changed to ‘Curators’.

222 Aust. Zool. 18(3), 1975

MEMORIAL NOTICE and BIBLIOGRAPHY

To give him his due, well before the Piltdown skull was denounced as a
fake in 1953, Troughton had always maintained that the mandible could not
possibly have belonged to that skull.

Troughton was a life-long follower of the theatre and ballet and was fond
of classical music. His memories of ballet went beck to Adeline Genée, Anna
Pavlova, and many later eminent artists and companies. When he was a boy, he
may have seen Sir Henry Irving. Later, Troughton took small parts or was an
extra on stage in various productions, not so much for the pay, but to study great
acting. He could not sing but mimed and danced as a “galloping major” in one
play, and he had a very strong vein of comedy. I remember him in jet doublet
and hose in “Elizabeth and Essex”. He acted in Gregan McMahon’s Sydney
Repertory Theatre Society productions in the 1920s and later in Miss Doris Fitton’s
Independent Theatre, North Sydney. He was a personal friend of Errol Flynn, Sir
Laurence Olivier, Vivien Leigh, Elsa Lanchester, Bette Davis and other actors,
of Sir Robert Helpmann and many ballet performers, as well as the entrepreneur,
Hugh Ward, and his actor-zoologist son, Melbourne Ward. Because of his stage
experience, Troughtie was always in much demand at museum ‘“‘smokos”, when
he alternated serious recitations (such as ‘‘The Hell-Gate of Soissons”) with
excruciatingly funny ones (“‘Mary”’, for example, or the young subaltern addressing
his troops on the uses of the ‘““Wifle”’), when anything was likely to happen—and
did!

Troughton used to work back at the museum on his mammal books and
papers until night-time, when the only sounds to disturb him were the newsboy’s
call and the clop-clop of horses’ hooves on the roads now choked with motor
traffic. On two occasions he was to encounter burglars: on one of these, in 1935,
an intruder was hiding on the roofing of the “Birds of the Admiralty Islets” case—
it was a wonder that he did not crash through the glass. ““We know you’re there,
we've got you covered,” called the police as Troughton crouched behind a pillar.
The man came down (he had stolen some gold) and was arrested. Troughton,
always the loyal public servant, pointed out to him the disgraceful nature of his
offence, emphasizing that a museum is an educational institution. ““Oh,” said the
culprit, ‘““who cares? It’s only the Government.” Monty Hey, the genial policeman
on point duty outside the Museum, moaned afterwards, “Only once in a hundred
years the museum has a burglary and they didn’t send for me!”

Ellis Troughton never married and his only brother predeceased him. Up
to the time of his death, after a heart attack, he was a Research Associate of the
Australian Museum, a Councillor of the Royal Zoological Society of New South
Wales, and an Honorary Fellow of the Museums Association of Australia.

ACKNOWLEDGEMENTS

I am grateful to the Australian Museum for access to relevant material. Most of my
account is based on memories of my long friendship with Ellis Troughton and on his
curricula vitae and manuscript lists of papers. Portraits of him appeared in the Australian

Aust. Zool. 18(3), 1975 223

GILBERT.Ps Whiley

Museum Magazine (2, 1926:413 and 12, 1958:338) and in the newspapers, Sydney
Morning Herald, 10 August 1931 and Aftenposten (Oslo, Norway), 15 October 1964.
Others are held by scientific societies or are in private hands. I am grateful to the Linnean
Society of New South Wales for the photograph of him, taken in 1973, reproduced herewith.

FIELD WORK AND TRAVELS

Mr. Troughton had travelled widely, always with the interests of the
Australian Museum at heart. For example, during a brief stop at Canton Island,
when being flown across the Pacific, he collected a shark for me. He had no
preservatives, so informed a Customs official in Sydney that if he held the shark
much longer, it would “‘declare’’ itself!

He had visited China, flown over the Burma Hump, and had had several
trips to England, Europe and the United States, also visiting Japan and Antarctica.
An argumentative old bushman, debating with him about the mode of reproduction
of kangaroos, once scornfully referred to Troughton as a “Pitt and George Street
naturalist’. Nothing could have been further from the truth, for Troughton spent
considerable periods in the bush, in many parts of Australia remote from those
Sydney thoroughfares, and he also collected in the Santa Cruz Islands, New
Guinea, the Moluccas and elsewhere. He preserved not only mammals but birds,
fishes, insects, etc., for the Australian Museum.

His more important collecting localities, roughly in chronological order, were
as follows.

His first field trip in 1912 at the age of 19 was to the Capricorn Group,
Queensland, with A. R. McCulloch and Professor Thomas Harvey Johnston. The
period from December 1919 to April 1920 was spent in South Australia (Farina,
Eyre’s Peninsula and Kangaroo Island) with C. M. Hoy, a representative of the
United States National Museum, Washington. Troughton then secured 1,416
specimens for the Australian Museum at a total cost of £74. In 1920 and 1921,
he collected at the Myall Lakes, Blue Mountains and the Nepean River, New
South Wales, and made one of what was to be many trips to Lord Howe Island.
From October to December 1921, with J. H. Wright as assistant, Troughton
collected at several stations along the Trans-Australian Railway,across the Nullarbor
Plain to south-western Australia, securing over 1,000 specimens for a total cost
of £105. A long series of bats was collected at Hunter’s Hill, near Sydney, in
1926. Later that year (July-August 1926), he visited Vanikoro and other islands
of the Santa Cruz Group with A. A. Livingstone; these places were important
as being the type-locality of some of Quoy and Gaimard’s species. Good collections
of marine life and ethnographical material were made at the same time, so that the
Museum received rich stores of mammals (notably bats), birds, reptiles, fishes,
shells, corals and insects.

In 1931, Troughton visited the National Park and Minnamurra, New South
Wales, with A. J. Marshall. In 1934, at his own expense, he visited remote parts

224 Aust. Zool. 18(3), 1975

MEMORIAL NOTICE and BIBLIOGRAPHY

of Queensland and the Northern Territory with H. O. Fletcher (Atherton, Mount
Isa, Alexandria Downs, Alroy Downs, the Barkly Tablelands and Avon Downs,
etc., to the Gulf of Carpentaria). After travelling overseas in the 1930s, he had
collected in every State of Australia and had visited New Zealand for ANZAAS.

His movements in New Guinea and environs in 1945 were shrouded in
wartime secrecy, but he investigated mammal carriers of scrub typhus thereabouts
with the Tropical Science Section of the Australian Imperial Forces and the
American Typhus Commission. At about this period Troughton visited: Morotai
(Moluccas, Indonesia), Biak, Schouten Island, and Japen Island (West Irian),
Hollandia [now Sukarnapura] (West Irian), and in Papua New Guinea, the foot
of the Cyclops Ranges and the Mount Lamington district (six years before the
famous eruption), Lae, Port Moresby, Buna, and Dobodura, as well as Kiriwina
in the Trobriands and Torokina, Bougainville, Solomon Islands.

In 1946, Troughton was one of a party to make a zoological reconnaissance
of the Mount Kosciusko region, New South Wales. He spent long service leave
abroad in 1949 and was Australian delegate to the International Technical Con-
ference for the Protection of Nature, under the auspices of UNESCO and the
International Union for the Protection of Nature, held at Lake Success, U.S.A.

He revisited Papua New Guinea with Norman Camps on the Australian
Museum 1954 Expedition, paying his own expenses (£75). I have given elsewhere
(Rec. Austr. Mus., 24, 1956: 23) a map of their route. This was the first expedi-
tion on behalf of any museum in Australia to undertake field work in general
zoology in New Guinea. In 1950-1951 Troughton revisited Mount Irvine, New
South Wales and, after retirement, went abroad and to Japan and Papua New
Guinea for science congresses and to Antarctica on a cruise.

He collected or discovered various new species of mammals, birds, reptiles,
fishes, shells, insects and the external and internal parasites of animals. Some of
these were very appropriately named after him. His own new names for taxa
are listed at the end of his bibliography, below.

The Cave Owl was named Tyto novaehollandiae troughtoni by Cayley (What
bird is that? 1931:32, pl. 5, fig. 4). I named one of his Vanikoro fishes Teuthis
troughtoni, and Iredale a clam (Tridacna). Musgrave named a nycteribiid bat
parasite, Livingstone a star-fish (Parasterina) and Johnston & Mawson, in 1941, a
nematode (Physaloptera troughtoni), and doubtless there are, or will be, other
taxa named in his honour.

Aust. Zool. 18(3), 1975 225

GILBERT P. WHITLEY

BIBLIOGRAPHY OF THE PUBLISHED WRITINGS OF
ELLIS LE GEYT TROUGHTON, C.M.ZS., F.R.ZS.

Books and papers are listed in chronological order, exact dates of publication being

given when possible. Names of joint authors are given in brackets after titles of papers.
Most items were published in Sydney unless otherwise stated.

Nm

226

1920
Notes on Australian mammals: No. 1.
Rec. Austr. Mus., 13 (3), Dec. 4, 1920, pp. 118-122, figs. 1-6.

192K
Exhibition of skull of Bettongia cuniculus.
Absir, “Proc. ‘Linn.--Soe7-N‘S:W ales:')356:: Sept: 2,5, 1924, =p: 2:> Proc Linn> See
N.S.Wales, 46, Nov. 2, 1921, p. 350.
Exhibition of yellow-footed pouched mouse, Phascogale flavipes.
Abstr. Proc. Linn. Soc. N.S.Wales, 356, Sept. 2, 1921, p. 2; Proc., 46, Nov. 2, 1921;
p. 350.

1922
A horse’s hardship.
Austr. Mus. Mag., 1 (5), July 1922, pp. 150-151, fig. [Osteclogical note. ]
Notes on Australian bats and the occurrence of Chalinolobus gouldii, Gray at Norfolk
Island.
Australian Zoologist, 3 (1), Sept. 15, 1922, pp. 39-41.

1923
Exhibition of skins of a rare native rat, Rattus mondraineus.
Proc. Linn. Soc. N.S:Wales, 47, 1922 (Feb. 15, 1923), p. xxiv. [Not in Abstract. ]
Exhibition of skulls of white-necked hair seal, Eumetopias albicollis.
Proc. Linn. Soc. N.S.Wales, 47,.1922 (Feb. 15, 1923), p. xxiv. [Not im Abstract.]
A revision of the rats of the genus Leporillus and the status of Hapalotis personata
Krefit.
Rec. Austr. Mus., 14 (1), Feb. 28, 1923, pp. 23-41, pls. 5-6.
The ‘honey mouse’, Tarsipes spenserae.
Austr. Zool., 3 (4), Aug. 15, 1923, pp. 148-156, pl. 23 and text-figure.

1924
Singular nesting sites of birds of the Nullarbor Plain.
The Fim,» 23.3) Jans: 7, 4924." pp2. 5-247 ples3 3:

. The stick-nest building rats of Australia.

Austr. Mus. Mag., 2 (1), late Jan. 1924, pp. 18-23, 5 figs.

Exhibition of insectivorous bat, Nyctinomus australis Gray, 1838, from Mittagong,
N.S.W.

Abstr. Proc. Linn. Soc. N.S.Wales, 381, June 27, 1924, p. 1; Proc., 49, 1924 (Feb.
A192), KRY

Exhibition of white-backed wren (Malurus leuconotus).

Abstr. Proc. Linn. Soc. N.S.Wales, 381, June 27, 1924, p. 2; Proc., 49, 1924 (Feb. 18,
EOD). ps XKVe

. The honey eating marsupial mice of Australia.

Austr. Mus. Mag., 2 (4),. Oct. 9;. 1924, pp. 127-132, 3° figs:
The fruit bat group.
Austr. Mus. Mag., 2 (4), Oct. 9, 1924, p. 140, fig.

1925

. The fruit bats or ‘flying foxes’ of Australia.

Austr. Mus. Mag., 2 (5), early Jan. 1925, pp. 169-174, 5 figs.

Aust. Zool. 18(3), 1975

17:

18.

19),

20.

25

22

25)
24.
23.

26.

27.

ZS

29:
30.
30
32.
See

34.

33.

36.

MEMORIAL NOTICE and BIBLIOGRAPHY

Captain Cook’s kangaroo. (By Tom Iredale & Troughton.)

Aussie Zo0l 263 08) % janes 42, 19295 pp. 3-3 16..pl.. 44:

Some strange African fruit-bats.

Austr. Mus. Mag., 2 (6), early April 1925, pp. 201-204, 3 figs.

A revision of the genera Taphozous and Saccolaimus (Chiroptera) in Australia and
New Guinea, including a new species, and a note on two Malayan forms.

Rec. Austr. Mus., 14 (4), April 9, 1925, pp. 313-341, pls. 47-48 and text-figs. 1-2.
Exhibition of skull of adult female aboriginal showing remarkable recovery from
injury.

Abstr. Proc. Linn. Soc. N.S.Wales, 394, Oct. 30, 1925, p. 2; Proc., 50, 1925 (Feb. 15,
1926) 5p: xiv:

Exhibition of mounted specimen of Caenolestes fuliginosa from Ecuador.

Abstr. Proc. Linn. Soc. N.S.Wales, 395, Nov. 27, 1925, p. 2; Proc., 50, 1925 (Feb. 15,
1926). pe xivil.

1926
‘Vampire’ bat.
Sunday News (newspaper, Sydney), March 1926, and fig.
Allan Riverstone McCulloch.
Austr. Mus. Mag., 2 (10), March 31, 1926. p. 346.
The mystery of marsupial birth and transference to the pouch.
Austr. Mus. Mag., 2 (11), July 13, 1926, pp. 387-391, frontispiece and 2 figs.
Recent expedition to the Santa Cruz Group.
Austr. Mus. Mag., 2 (12), Oct. 8, 1926, pp. 413-414, fig.
Striped opossums (Dactylopsila).
Austr. Encyclopaedia, 2, Oct. 11, 1926, p. 508. [Troughton also revised Haswell’s
articles on mammals for this encyclopaedia. |
A chapter on the bats of Australia and New Guinea.
In A. S. Le Souef and H. Burrell, Wild Animals of Australasia (London: Harrap),
Oct. 1926, pp. 21-88, illustrated, plates, figs. 2-8 and 8 text-figs.
[Exhibition of the] skull of large bandicoot, Isoodon macrourus, from Clarence
River, New South Wales.
ADSIER ETOGs4 bin. SOC..IN-S: W ales... 403,.:: Oct, 329; 1926... pp: : 1-25. Proc. 51,,.1926
(Feb. 23, 1927). p:. xin.

1927

Fixation of the habitat, and extended description, of Pteropus tuberculatus, Peters.
Rec. Austr. Mus., 15 (5), April 6, 1927, pp. 355-359, fig. 1.
Cruising in the Santa Cruz. (By Troughton & A. A. Livingstone.)
Austr. Mus. Mag., 3 (2), April 11, 1927, pp. 41-48, 9 figs.
Further notes on marsupial birth.

Austr. Mus. Mag., 3 (2), April 11, 1927, pp. 53-56.

The Isles of Santa Cruz. (By Troughton & A. A. Livingstone.)
Austr. Mus. Mag., 3 (3), July 8, 1927, pp. 77-85, 11 figs.

Last days at Santa Cruz. (By Troughton & A. A. Livingstone.)
Austr. Mus. Mag., 3 (4), Oct. 13, 1927, pp. 113-125, 13 figs.
Hints for collectors: mammals.

Austr. Mus. [Circular, 4], Dec. 1927, pp. 1-6.

1928
A new genus, species and sub-species of marsupial mice (Family Dasyuridae).
Rec. Austr. Mus., 16 (6), June 11, 1928, pp. 281-288, pl. 39.

Sea cows. The story of the dugong.
Austr. Mus. Mag., 3 (7), July 13, 1928, pp. 220-228, 7 figs.

Aust. Zool. 18(3), 1975 227

38.

39.

40.

41.

42.

43.

44,

45.

46.

47.

48.

49.
50.
sd ie

D2,

53s

54.

228

GIEBERT cP." WHIMEEY

1929

A new genus and species of bat (Kerivoulinae) from the Solomons, with a review
of the genera of the sub-family.

Rec. Austr. Mus., 17 (2), June 26, 1929, pp. 85-99, fig. 1.

New forms of mosaic-tailed rats (Melomys and Uromys) from Hinchinbrook Island,
Queensland. (By Troughton & A. S. Le Souef.)

Austr. Zool., 6 (1), Aug. 13, 1929, pp. 96-99.

Note on the occurrence of a second species of fruit-bat (Pteropus scapulatus) in
New South Wales.

Austr. Zool., 6-(1), Aug. 13, 1929, pp. 104-106.

A new fruit-bat (Pteropus rayneri Group) from the Solomons.

Rec. Austr. Mus., 17 (4), Sept. 4, 1929, pp. 193-198.

A new species of ring-tailed phalanger (Ps. laniginosus Group) from the Bunya
Mountains, S.E. Queensland. (By Troughton & A. S. Le Souef.)

Rec. Austr. Mus., 17 (6), Nov. 28, 1929, pp. 291-296, coloured pl. 45 and text-fig. 1.

1930

Notes on striped opossums of the genus Dactylopsila. ~

Austr: Zool., 6 (2), Jan: 14, 1930, pp. 169-174:

A new species and sub-species of fruit bats (Pteropus) from the Santa Cruz Group.
Rec. Austr. Mus., 18 (1), Nov. 10, 1930, pp. 1-4.

(930

The occurrence of a male and female Grampus griseus (Delphinidae) at Sydney,
New South Wales.

Proc. Zool. Soc. (London), 1931 (June 29, 1931), pp. 565-569, pl. 1.

Three new bats of the genera Pteropus, Nyctimene and Chaerephon from Melanesia.
Proc. Linn. Soc. N.S.Wales, 46 (3), July 15, 1931, pp. 204-209.

The habits and food of some Australian mammals.

Austr. Zool., 7 (1), Aug. 24, 1931, pp. 77-83. [Chalinolobus gouldi, Dromicia nana &
Perameles nasuta.]

1932

On five new rats of the genus Pseudomys.

Rec. Austr. Mus., 18 (6), April 20, 1932, pp. 287-294.

A new species of fat-tailed marsupial mouse, and the status of Antechinus froggatti
Ramsay.

Rec. Austr. Mus., 18 (6), April 20, 1932, pp. 349-353, fig. 1.

The egg-laying furred animals of Australia.

Austr. Mus. Mag., 4 (10), April 29, 1932, pp. 327-334, frontispiece and 6 figs.
Australian furred animals, their past, present, and future. Presidential address.
Austr.’ Zool. 7 (Be Sept-21554 19382; spprlsa-193..

A revision of the rabbit-bandicoots. Family Peramelidae, Genus Macrotis.

Austr. Zool.,’T (3).:Sept.* 15,1932, pp7219-236, fig. 1.

Australian mammals ... Mr. E. Troughton’s story.

Sydney Morning Herald (newspaper), Oct. 7, 1932, p. 11, col. h.

1933
The egg-laying furred animals of Australia.
Mid-Pacific Magazine (Honolulu), 45 (4), April 1933, pp. 358-364, 1 fig.
The correct generic names for the grampus or killer whale, and the so-called grampus
or Risso’s dolphin. (By Tom Iredale & Troughton.)
Rec. Austr. Mus., 19 (1), Aug. 2, 1933, pp. 28-36, pl. 10.

Aust. Zool. 18(3), 1975

3D:

56.

65.

66.

67.
68.
69.
70.
TKIP

ee

ee

74.

MEMORIAL NOTICE and BIBLIOGRAPHY

1934

Check-list of the mammals recorded from Australia. (By Tom Iredale & Troughton. )
Austr. Mus. Mem., 6, May 4, 1934, pp. i-xii + 1-122. [The status of some new names
in this publication has been (wrongly) questioned; in any case the new genera have
been reinforced in Troughton, Furred Animals of Australia, ed. 2, 1943, nos. 83 & 94,
below. |

Marsupial gliders ocr ‘flying possums’.

Austr. Mus. Mag., 5 (8), Nov. 21, 1934, pp. 257-264, coloured cover and 3 figs.

1935
The largest gliders or ‘flying possums’.
Austr. Mus. Mag., 5 (9), Jan. 16, 1935, pp. 314-319, 2 figs.
Five new rats of the genera Hydromys and Melomys from northern Australia.
Rec. Austr. Mus., 19 (4), Sept. 19, 1935, pp. 251-258, figs. 1-2.
A new genus and species of giant rat from the Solomons.
Kec. Austr. Mus) 19°4)2 Septs.19,: 1935 > pp. 259-262; ply 19:
The southern race of the koala.
Austr. Naturalist, 9 (6), Sept., 1935, pp. 137-140 (reprinted without pagination).

1936

The mammalian fauna of Bougainville Island, Solomon Group.

Rec, Aus. Mas... 19° (5). April.-7,. 1936, pp.).341-353.

Descriptions of new rats and mice from Queensland.

Mem. Qld. Mus. (Brisbane), 11 (1), April 17, 1936, pp. 14-22.

Two new tree kangaroos from Papua, with notes on allied forms. (By Troughton &
A. S. Le Souef.)

Austra Zoal:,- 8 (3)... June 29: +19396,. pps 93-197.

A redescription of Solomys (‘Mus’) salamonis Ramsev.

Proc: Linn, Soc. -N.S.Wales,. 61, Sept. “157 1936; pp:. 128-130.

A new tree-kangaroo from south-eastern Papua. (Ry Troughton & A. S. Le Souef.)
Rec. Austr. Mus., 19 (6), Oct. 7, 1936, pp. 388-390.

A creed for nature lovers.

Austr. Mus. Mag., 6 (4), Nov. 20, 1936, p. 128. Also reprinted in Furred Animals of
Australia, from 2nd edition, 1943, p. xv to later eds., and by Fauna Protection Panel,
no date, one sheet.

1937

Six new bats (Microchiroptera) from the Australasian region.

Austr. Zool., 8 (4), March 12, 1937, pp. 274-281.

On new forms of the eastern swamp rat, and the relationship of Mastacomys.
Austr. Zool., 8 (4), March 12, 1937, pp. 281-286.

The identity of Cook’s kangaroo. (By Tom Iredale & Troughton.)

Rec Ansir. Mus. 20:.(l je May: 15,1937. pp: 67-71.

Descriptions of some New Guinea mammals.

Rec: sAusth. Mus... 20 (2), Aue 27-5 1937, pp. 117-127:

The status of ‘Mus’ novaehollandiae Waterhouse, and allied forms.

Rec. Austr. Mus., 20 (2), Aug. 27, 1937, pp. 185-1990.

New marsupial groups in the Australian Museum.

Austr. Mus. Mag., 6 (8), Oct.-Dec. 1937, pp. 255-259, frontispiece and 5 figs.

1938
The platypus group: life history exhibit in the Australian Museum.
Austr. Mus. Mag., 6 (11), Sept. 30, 1938, pp. 363-367, frontispiece and 2 figs.
Australian mammals: their past and future.
J. Mammalogy (Baltimore, U.S.A.), 19 (4), Nov. 14, 1938, pp. 401-411, + corri-
gendum on p. 408.

Aust. Zool. 18(3), 1975 229

76.

Md:

78.

Fo.
80.
81.

82.

83.

84.

85.

86.

87.

230

GILBERT WP: AWHUREY

Mammals.

Guide to the Australian Museum and its contents, [ed. 1,] 1938, pp. 9-24, 12 figs.
Revised [2nd.] ed., 1941, pp. 10-25, frontispiece and 15 figs. Ed. 3, 1943, reprinted
1944, 1948 & 1952, pp. 10-25, frontispiece and i5 figs. See also no. 144 = Handbook
Austr. Mus., ed. 1, 1957, pp. 99-109, 6 figs. Ed. 2, 1962.

Osteology, or the study of skeletons.

Guide to the Australian Museum and its contents, [ed. 1,] 1938, pp. 25-32, 7 figs.
Revised [2nd.] ed., 1941, pp. 56-61, 4 figs. Ed. 3, 1943, reprinted 1944, 1948 & 1952,
pp. 56-61, 4 figs. See also no. 145 = Handbook Austr. Mus., ed. 1, 1957, p. 109.
Ed. 2, 1962.

1939

Queensland rats of economic importance, and new forms of Rattus and Thetomys.
Rec. Austr. Mus., 20 (4), March 31, 1939, pp. 278-281.

Australian mammals in peril.

The Australian National Review (Canberra: Canberra Publ. Co.), 5 (30), June 1,
1939, pp. 50-61.

1940

The urgency of biological survey in relation to faunal and economic problems.
Austr. J. Sci. (Sydney), 2 (5), April 22, 1940, pp. 147-150.

Ambergris: its uses and identification.

Austr. Mus. Mag., 7 (5), June 5, 1940, pp. 169-173, 2 figs.

Furred aeronauts of night: little bats as friends of man.

Austr. Mus. Mag., 7 (6), Sept. 2, 1940, pp. 206-211, 6 figs.

Vampire bats in fact and legend.

Austr. Mus. Mag., 7 (7), Dec. 28, 1940, pp. 244-248, 3 figs.

1941

Furred animals of Australia (Sydney: Angus & Robertson).
ed. 1, Nov. 20, 1941, pp. i-xxviii + 1-376, col’d. pls. 1-25.
ed. 2, Nov. 1943, pp. i-xxvili + 1-376, col’d. pls. 1-25.
[Repeated as no. 94, below, because this revised edition has new generic and
subspecific names and confirms the status of Iredale & Troughton’s 1934 names.]
ed. 3, Nov. 1946, pp. i-xxx + 1-376, col’d. pls. 1-25.
Reprint of ed. 3 (New York: Charles Scribner’s), 1947, pp. i-xxvili + 1-374, col’d.
pls. 1-25.
[Ed. 4] (Sydney: Angus & Robertson), 1951, pp. i-xxxii + 1-376, col’d. pls. 1-25.
[Bd- 51,1954; be
PEd?6]. 1957, a
fEd- 74; 1962, a
[Ed. 8], 1965, 9
[Ed. 9], 1967, pp. 416, col’d. pls. 1-25.
‘Paperback’ ed., 1973 = Troughton’s furred animals of Australia [ed. 10, 1974],
pp. i-xviii + 1-314, col’d pls. 1-25 and map.
Australian water-rats: their origin and habits.
Austr. Mus. Mag., 7 (11), Dec. 15, 1941, pp. 377-381, fig.
The Solvol animal book: 55 Australian mammals described and illustrated. (Sydney:
Halstead Press), 1941, pp. 1-28, 55 coloured figs.

1942

The striped possums of Australia and New Guinea.

Austr. Mus. Mag., 7 (12), March 16, 1942, pp. 431-434, fig.
The kangaroo family: origin and earliest discoveries.

Austr. Mus. Mag., 8 (1), July 20, 1942, pp. 17-22, 3 figs.

Aust. Zool. 18(3), 1975

88.

89.

90.
ot.
BD

93;

94.

DS:

06,

97.
98.
99:

100.

101.
102.
103.
104.

105.

MEMORIAL NOTICE and BIBLIOGRAPHY

Earliest, hitherto unnoticed, account of the unaided transference of a newly-born
kangaroo to the teat, by A. Collie... 1830... Thylogale eugenii.

Proc. Linn. Soc. N.S.Wales, 67, May 15, 1942, p. xxii; Abstr., 546, Sept. 25, 1942,
p. 2:

The truth about marsupial birth.

Austr. Mus. Mag., 8 (2), Nov. 20, 1942, pp. 40-44, frontispiece and 2 figs.

1943
The hopping or jerboa marsupial-mice.
Austr. Mus. Mag., 8 (5), Sept. 30, 1943, pp. 159-160, fig.
The kangaroo family: rat kangaroos, I.
Austr. Mus. Mag., 8 (5), Sept. 30, 1943, pp. 171-175, 3 figs.
[Exhibition of] photo of a hopping or ‘jerboa’ marsupial-mouse, Antechinomys laniger
Gould, 1856... from . . . Central New South Wales [and note on] A. spenceri.
Proc. Linn. Soc. N.S.Wales, 68, May 15, 1943, pp. xix-xx; Abstr. 555, Oct. 1, 1943,
pp. 1-2.
The third specimen of the false water-rat (Xeromys myoides Thomas 1899) known
to have been preserved.
Proc. Linn. Soc. N.S.Wales, 68, May 15, 1943, p. xxi; Abstr., 557, Nov. 26, 1943,
pz.
[The second edition of Furred Animals of Australia was published in November 1943,
being a revised account of the first edition of 1941 (q.v.); it contains new generic
and subspecific names and confirms Iredale & Troughton’s 1934 names. For other
editions of Furred Animals of Australia, see entries no. 83 and 168. ]

1944
The kangaroo family: rat-kangaroos, II.
Austr. Mus. Mag., 8 (6), Feb. 15, 1944, pp. 204-207, 2 figs.
The imperative need for federal control of post-war protection of nature. Presidential
address.
Abstr. Proc. Linn. Soc. N.S.Wales, 558, March 31, 1944, pp. 2-3. Full text in Proc.,
69, 1944, pp. iv-xvi. (Reviewed as ‘A new order for our fauna’ in Austr. Mus. Mag.,
8 (7), May 1, 1944, pp. 217-218.)
The kangaroo family: hare wallabies.
Austr. Mus. Mag., 8 (7), May 1, 1944, pp. 229-233, 2 figs.
Some aspects of the natural history of New Guinea. (2) the mammals.
Proc. Linn. Soc. N.S.Wales, 69, May 15, 1944, p. xxi; Abstr., 560, June 2, 1944, p. 1.
The kangaroo family: tree wallabies.
Austr. Mus. Mag., 8 (8), Aug. 1, 1944, pp. 277-280, 2 figs.
The kangaroo family: rock wallabies.
Austr. Mus. Mag., 8 (9), Nov. 1, 1944, pp. 295-299, 3 figs.

; 1945
The kangaroo family: the nail-tail wallabies.
Austr. Mus. Mag., 8 (10), Feb. 16, 1945, pp. 346-349, frontispiece and 2 figs.
Conservation of wild life, total protection for al! wallabies in Victoria.
Austr. Mus. Mag., 8 (10), Feb. 16, 1945, p. 349.
Diagnoses of new mammals from the south-west Pacific.
Rec: Austr. Mus.,-21 (6), June 25; 1945, pp: 373-375:
Review: “The story of Shy the platypus’, by Leslie Rees.
Austra Mus? Mag... 8012). Dee. b. 1945, 2p. 427.
Review of field investigations concerning the mammal reservoirs of scrub typhus.
Commonwealth (Restricted) Report, Australian Imperial Forces, Scientific Bureau,
C.S.ILR., Trop. Sci. Sect., 2, 1945, pp. 1-14. [Not seen. Not in Zoological Record.
No new names. Troughton’s researches in New Guinea. ]

Aust. Zool. 18(3), 1975 231

106.

107.

108.

109.

110.

Pt:

[t2-

ii}:

114.
ED;

116.

£17.

113;

£15:

120.
121.

122:

123.

232

GIEBERT PP: sWHITREY

1946

Diagnoses of new rats from the New Guinea area.

Rec. Austr. Mus., 21 (7), June 24, 1946, pp. 406-410.

Report on the mammals.

Report to the Trustees Kosciusko State Park by the joint scientific committee of the
Linnean Society of New South Wales and the Royal Zoological Society of N.S.W.
on a reconnaissance natural history survey of the park. January-February, 1946.
(Sydney: roneo’d), pp. 49-55.

1947

Kangaroo twins — and triplets.

Austr. Mus. Mag., 9 (5), Aug. 30, 1947, pp. 160-164, 4 figs.
1948

The marsupial banded ant-eater or numbat.
Austr. Mus. Mag., 9 (9), Dec. 31, 1948, pp. 298-302, 3 figs.

1949
Notes on the furred animals.
In Axel Poignant, Bush Animals of Australia. (Sydney: Shepherd Press), 1949,
pp. 3-38, 19 figs.
Creatures of the bush. Jn This land of ours: Australia. (Edited by G. M. Farwell &
F. H. Johnston). (Sydney: Angus & Robertson), 1949, pp. 61-65, 3 ‘figs.

The egg-laying platypus — wonder mammal of Australia.
Mangrovite Calendar, 1949, 1 page, coloured plate.
1950

Unique Australian mammals.
Discovery (Norwich, England: Jarrold), 11 (5), May 1950, pp. 148-152, cover

and 9 figs.

Bandicoots — rare and otherwise. Part I.

Aust. Mus. Mag., 10 (3), Sept. 15, 1950, pp. 95-98, 2 figs.
Bandicoots — rare and otherwise. Part II.

Austr. Mus. Mag., 10 (4), Dec. 15, 1950, pp. 113-117, 3 figs.
Review. “The mammals of Victoria’, by C. W. Brazenor.
Austr. Mus, Maeg., \0 (4), Dec. 5, 19509 p. 136.

A951

Book review. “Ihe mammals of Victoria’, by C. W. Brazenor.

Ausit: Zool lt (4). uly Bi. 195. p: 370. [Different from the araceaine: no. 116.]
The marsupials.

Australian Junior Encyclopaedia (Melbourne: Georgian House), 2, Aug. 1951, pp.
820-823, 4 figs. and Sydney edition, 3, 1956.

Kangaroos and wallabies.

Australian Junior Encyclopaedia (Melbourne: Georgian House), 2, Aug. 1951, pp.
837-844, 6 figs. and Sydney edition, 3, 1956.

The kangaroo family — the pademelons or scrub-wallabies — Part I.
Austr. Mus. Mag., 10 (7), ‘Sept. 15, 1951, pp. 218-222, 2. figs.
The kangaroo family — the pademelons or scrub-wallabies — Part II.

Austr. Mus. Mag., 10 (8), Dec. 15, 1951, pp. 261-264, cover and 1 text-fig.
Land of living fossils.
Science Digest (U.S.A.), 29 (2), 1951, pp. 15-19. [Not seen — G.P.W.]

1952
Albert Sherbourne Le Souef.
Proc, Roy. Zool. Soc; 4N.S.Wales,.~V950-51o (March 5;, 1952). . pps 8-10,— portrait.
[Obituary notice. |

Aust. Zool. 18(3), 1975

124.

125.

126.
{27.
128.

129.
130.
Sh.
132.

133.

134.

135;

136.

137:

138.
139.

140.

141.
142.

143.

MEMORIAL NOTICE and BIBLIOGRAPHY

1953

Review: ‘Possums’, by Carl G. Hartman.

Austr. Mus. Mag., 11 (2), June 15, 1953, pp. 65-67.

Our marsupial ‘native-cat’.

Austr. Mus. Mag., 11 (3), Sept. 15, 1953, pp. 90-92, cover-photo and text-fig.

1954

Australian mammal series. The platypus (Ornithorhynchus anatinus).

Austr. Mus. School Leaflet, 6, Jan. 1954, pp. 1-4, 2 figs. Reissued as Austr. Mus.
Leaflet, 50, April 1960, and again, Dec. 1962, pp. 1-4, 2 figs.

Australian mammal series. Possums.

Austr. Mus. School Leaflet, 7, Jan. 1954, pp. 1-4, 3 figs. Reissued as Austr. Mus. Leaflet,
7, revised August 1959, pp. 1-4, 3 figs.

Australian mammal series. No. 2. Spiny ant-eater (Tachyglossus aculeatus).

Austr. Mus. School Leaflet, 8, Jan. 1954, pp. 1-4, 2 figs. Reissued as Austr. Mus.
Leaflet, 8, March 1960, pp. 1-4, 2 figs.

Scientific research and curatorial duties.

Kalori: (Sydney),; 6, Jan. 29; .1954;:-p--3.

The marsupial ‘tiger-cat’.

Austr. Mus. Mag., 11 (5), March 15, 1954, pp. 168-170, cover-photo and 3 figs.
The marsupial ‘tiger-cat’, birth and growth in captivity.

Austr. Mus. Mag., 11 (6), June 15, 1954, pp. 200-202, 4 figs.

Australian mammal series. Koala (Phascolarctos cinereus).

Austr. Mus. School Leaflet, 9, June 1954, pp. 1-4, fig.

Museum work in the New Guinea highlands.

Austr. Mus. Mag., 11 (8), Dec. 15, 1954, p. 256.

1955

The koala.

Austr. Mus. Mag., 11 (12), Dec. 15, 1955, pp. 396-401, cover-photo and text-fig.
1956

Interest in marsupials.

Austr. Mus. Mag., 12. (1), March 15, 1956, p. 10.

Koala — orphan of the gum-trees.

Walkabout (Melbourne: Journ. Austr. Geogr. Soc.), 22 (12), Dec. 1, 1956, pp.
36, 39 & 41, fig. [See also no. 154, below. ]

Retirement of Mr. J. R. Kinghorn.

Austr. Mus. Mag., 12 (4), Dec. 15, 1956, p. 125.

1957
Our front cover.
Austr. Mus. Mag., 12 (5), March 15, 1957, p. 137 and cover-photo [of Cuscus].
Book for Colombo Plan countries.
Austr. Mus. Mag., 12 (5), March 15, 1957, p. 159.
The wombat.
Austr. Mus. Leaflet, 34, March 1957, pp. 1-4, fig. Revised, Sept. 1959, pp. 1-4, fig.,
and later.
A new native dog from the Papuan highlands.
Proc. Roy. Zool. Soc. N.S.Wales, 1955-56 (May 8, 1957), pp. 93-94.
The kangaroo family: brush-wallabies.
Austr. Mus. Mag., 12 (6), June 15, 1957, pp. 186-192, cover-photo and 3 text-figs.
The kangaroo family: kangaroos and wallaroos.
Austr. Mus. Mag., 12 (7), Sept. 15, 1957, pp. 230-237, 4 figs.

Aust. Zool. 18(3), 1975 233

144.

145.

146.

147.

148.

149.

150.

151.

234

GILBERT: P.c WHITLEY.

Mammals.

Austr. Mus. Handbook, ed. 1, 1957, pp. 99-109, 6 figs. Ed. 2, 1962, pp. 105-114,
6 figs. [Derived from no. 75, above. |

Osteology, or the study of skeletons.

Austr. Mus. Handbook, ed. 1,, 1957, p. 109. [Omitted from ed. 2. Derived from no. 76,
above. |

1958

[Twenty-five articles in the Australian Encyclopaedia, all published on June 3, 1958,
as follows:—]

Ant-eater, Vol. 1, p. 189.

Banded ant-eater or numbat, 1, p. 408.
Bandicoots, 1, pp. 409-410, fig.

Bats, 1, pp. 458-461.

Cats, Marsupial, 2, pp. 287-288. Z

Dolphins, 3, pp. 266-267.

Dugong, 3, p. 307.

Kangaroos and Wallabies, 5, pp. 156-162, 4 figs.
Koala, 5, pp. 203-205, pl. 202 A and text-fig.
Mammals, 5, p. 471, coloured pl. 470 A.
Marsupials, 6, pp. 1-2, fig.

Mice, Marsupial, 6, pp. 67-69.

Mole, Marsupial, 6, pp. 115-116.

Monotremes, 6, pp. 139-141.

Platypus, 7, pp. 150-154, pl. 152 A and text-fig.
Possums, 7, pp. 233-239, 4 figs.

Rats and Mice, 7, pp. 389-391.

Rodents, 7, p. 476.

Seals, 8, pp. 54-56.

Spiny ant-eaters, 8, pp. 241-243, fig.

Tasmanian Devil, 8, pp. 437-438, fig.

Thylacine, 8, pp. 495-496, fig.

Water-rats, 9, p. 195.

Whales, 9, pp. 268-273.

Wombats, 9, pp. 345-346.

Kangaroos.

Austr. Mus. Leaflet, 46, Dec. 1958, pp. 1-8, 9 figs. Reissued Aug. 1960.

1959

The thylacine or “Tasmanian wolf’.

Austr. Mus. Leaflet, 49, Aug. 1959 (and reissued July, 1960), pp. 1-4, fig.

The marsupial fauna: its origin and radiation.

In A. Keast (ed.), Biogeography and ecology in Australia. Monogr. Biol. (Den Haag:
Junk), 8, Sept. 1959, pp. 69-88.

1960

Hints for collectors: mammals.
Austr. Mus. Leaflet, 52, Aug. 1960, pp. 1-6. [Compare no. 34, above. ]

1962

The actual identity of Captain Cook’s kangaroo. (By Tom Iredale & Troughton.)
Abstr. .Proc. Linn: Soc. N.S.Wales, 714, July 25. GpublugAug. 1) 1962; pity Proc:
87 (2), 1962 (Jan. 10, 1963), pp. 177-184, figs. 1-2.

Aust. Zool. 18(3), 1975

152.

ESS":

154.

i535:

156.

157:

158.

159.

160.

161.

162.
163.
164.

165.

166.

MEMORIAL NOTICE and BIBLIOGRAPHY

1963
[Exhibition of] series of colour transparencies showing the effect of the slaughter of
the Red Kangaroo in the Western Division.

Abstr. Proc. Linn. Soc. N.S.Wales, 724, Dec. 4, 1963, p. 2; Proc., 88, 1963 (April 10,
1964), p. 404.

1964

[Exhibition of] skins and skulls of the whiptail wallaby and wallaroo taken from
within 12 miles of Cooktown where the Endeavour was beached for repair in 1770.
Abstr. Proc. Linn. Soc. N.S.Wales, 727, June 3, 1964, p. 3; Proc., 89 (3), 1964 (May 7,
1965), p. 382; correction to Abstract 727, 1964.

Koala — orphan of the gum-trees.

In A. Bolton (ed.), Walkabout’s Australia (Sydney: Ure Smith), 1964, pp. 140-148,
fig. [Similar to no. 136, but with a different illustration and with a postscript added. ]
Comments on the proposed stabilization of Macropus Shaw, 1790. (By D. F. McMichael
& Troughton.)

Bull. Zool. Nomencl. (London), 21, 1964, pp. 255-259. [The Zoological Record gives
as joint authors, Finlayson, Kirkpatrick, McMichael, Morrison-Scott and Troughton,
and the pagination as 329-331.] .

The last of the toolaches.

In A. H. Chisholm (ed.), Land of Wonder (Sydney: Angus & Robertson), 1964,
pp. 122-123; also reprinted in 1966 (Sydney), and in paperback ed. (Hong Kong:
Pacific Books); 1971) pp. 122-123. ie

1965
A review of the marsupial genus Sminthopsis (Phascogalinae) and diagnoses of new
forms.
Proc. Linn. Soc. N.S.Wales, 89 (3), 1964 (May 7, 1965), pp. 307-321, 7 figs.
Note from an Australian mammalogist on usage of the common name ‘possum’.
Tuatara (Wellington, New Zealand), 13 (3), Nov. 1965, pp. 192-193.

1966

[Exhibition of] skull of the adult male holotype of the New Guinea Highland Dog,
Canis hallstromi, from the Huri-Duna country, Southern Highlands district of Papua.
Abstr. Proc. Linn. Soc. N.S.Wales, 746, Dec. 5, 1966, pp. 1-2; Proc., 91 (3), March
D2 ASG op: 238:

1967
Author’s comment on book review.
Austr. Nat. Hist., 15 (9), March 15, 1967, p. 298.
Egg-laying mammals — the monotremes.
In D. F. McMichael (ed.) A Treasury of Australian Wildlife (London: Ure Smith),
Nov. 15, 1967, pp. 22-31, 7 figs.
Bandicoots — rare and otherwise.
Ibid., pp. 32-41, 5 figs.
Kangaroos and wallaroos.
Ibid., pp. 42-47 & 50-53, 4 figs.
The koala.
Ibid., pp. 54-60, 2 figs.
Marsupial native and tiger cats.
Ibid., Nov. 15, 1967, pp. 48-49 & 65-71, coloured pls. ve 2 & 4 figs.

1968

Broom’s pygmy possum.
Proc. Roy. Zool. Soc. N.S.Wales, 1966-67 (Jan. 17, 1968), pp. 20-24, pl. 3 and
text-fig. 1.

Aust. Zoo]. 18(3), 1975 Poo

GILBERT...P) WHITLEY

1974

167. The early history and relationships of the New Guinea Highland Dog (Canis
hallstromi).

A.N.Z.A.A.S. 42nd. Congress (Port Moresby, Papua) Programme, 1970, p. 98 (title
only); Proc. Linn. Soc. N.S.Wales, 96 (2), Sept. 15, 1971, pp. 93-98, figs. 1-2.

1973
168. Troughton’s furred animals of Australia.
(Sydney: Angus & Robertson), 8vo., pp. i-xvili + 1-314, col’d. pls. 1-25 and map.
Ninth edition revised and abridged of no. 83, above, 1973. [Virtually 10th ed., 1974. ]
Mr. Troughton also wrote to the Daily Telegraph newspaper, Sydney, protesting
against the building of an airstrip on Lord Howe Island.

LIST OF JOINT AUTHORS

Finlayson, H. H. See item no. 155.
Iredale, T. e459, O09 ea bol.
Kirkpatrick, T. H. 155:
Le Souef, A. S. 38, 41, 63 & 65.
Livingstone, A. A. 30, 32 & 33.

’ McMichael, D. F. 155.

Morrison-Scott, T. C. 15S.

Zoological Record lists also J. T. Woods, E. Mayr, W. D. L. Ride and H. Lemche as
associated with no. 155.

INDEX TO TROUGHTON’S NEW TAXA

‘In the following list, all the new generic, specific and subspecific names proposed by
Troughton (including those in collaboration with other authors) are arranged in alphabetical
order, with references to the papers and pages wherein they were first proposed.

altape, Rattus browni, 70: 122.
Anamygdon, 37: 87.

apex, Rattus culmorum, 77: 280.
apicalis, Potorous tridactylus, 94: 162.
aquilo, Scoteinus orion, 67: 278.
aramia, Rattus gestri, 70: 119.
Austronomus, 55: 100 & 94: 363.
berneyi, Gyomys, 62: 15.

biakensis, Rattus, 106: 409.

bolami, Pseudomys hermannsburgensis, 47: 292.
bougainville, Melomys, 61: 344.
bougainville, Nyctimene, 61: 349.
brazenori, Leggadina hermannsburgensis, 71: 187.
brunneus, Planigale ingrami, 35: 282.
bunae, Rattus gestri, 106: 408.
cambrica, Macrotis lagotis, 51: 230.
cambricus, Rattus lutreolus, 68: 283.
caprenus, Scoteinus balstoni, 67: 279.
decres, Thylogale eugenii, 83: 194.
deltae, Dendrolagus, 63: 195.

desertor, Pseudomys (Gyomys), 47: 293.
dobodurae, Rattus ringens, 106: 407.

236 Aust. Zool. 18(3), 1975

MEMORIAL NOTICE and BIBLIOGRAPHY

eitape, Conoyces hageni, 70: 117.
exilis, Grampidelphis, 54: 32.

exilis, Uromys macropus, 38: 98.
Falsistrellus, 94: 349.

flavescens, Pseudomys minnie, 62: 19.
Fretidelphis, 55: 65.

froggatti, Melomys muscalis, 70: 123.
gawae, Rattus browni, 103: 374.
Grampidelphis, 54: 31.

grandis, Macrotis lagotis, 51: 229.
granulipes, Sminthopsis, 48: 350.
grootensis, Hydromys, 58: 252.
hageni, Melomys, 70: 124.

hageni, Rattus mordax, 70: 120.
hallstromi, Canis, 141: 93.

heffernani, Pteropus tonganus, 43: 3.
howensis, Pteropus, 45: 204.

imbil, Rattus lutreolus, 68: 283.
insulae, Melomys littoralis, 38: 96.
interjecta, Macrotis lagotis, 51: 227.
kohlsi, Petaurus, 103: 373.

kuzira, Grampidelphis, 54: 34.
lamington, Uromys, 70: 126.
lawnensis, Hydromys, 58: 253.
limicauda, Melomys, 58: 255.
lumholtzi, Sminthopsis, 55: 11.
maugeanus, Tursiops, 55: 68.
mediocris, Rattus praetor, 61: 343.
megalotis, Notomys, 55: 84.
Micronomus, 55: 100.

minnie, Pseudomys (Pseudomys), 47: 287.
mixtus, Melomys, 58: 257.

mixtus, Saccolaimus, 19: 322.

moae, Hydromys, 58: 254.
monticola, Sminthopsis, 157: 311.
ooldea, Sminthopsis murina, 157: 313 & 316.
oriens, Hydromys, 70: 127.

orion, Scoteinus, 67: 277.

owiensis, Rattus, 103: 374.

pallidus, Melomys cervinipes, 38: 97.
peccatus, Rattus greyi, 71: 189.
philipi, Echymipera, 103: 373.
Planigale, 35: 282.

ponceleti, Pipistrellus, 61: 351.
ponceleti, Unicomys, 59: 260.
praecelsus, Rattus browni, 70: 121.
profugus, Dendrolagus dorianus, 65: 389.
pumilis, Gyomys, 62: 16.

purdiensis, Rattus, 106: 408.
rawlinnae, Pseudomys (Pseudomys), 47: 289.
reginae, Hipposideros diadema, 67: 275.
reginae, Notomys alexis, 62, 20.
Registrellus, 94: 349.

rennelli, Pteropus, 40: 193.

rennelli, Rattus, 103: 375.
Rhinophyllotus, 55: 92.

Aust. Zool. 18(3), 1975 237

238

GILBERT P. WHITLEY

rubidus, Pseudocheirus, 41: 294.
salebrosus, Solomys, 61: 346.

sanborni, Scoteinus, 67: 280.
sanctacrucis, Nyctimene, 45: 206.
sanctacrucis, Pteropus, 43: 3.

sansapor, Rattus, 106: 409.

Scoteanax, 94: 353.

Scotorepens, 94: 354.

solomonis, Anamygdon, 37: 89.
solomonis, Chaerephon, 45: 207.
spadix, Dendrolagus, 63: 194.

sufflectus, Rattus browni, 70: 122.

tatei, Sminthopsis murina, 157: 313 & 316.
tenuirostris, Planigale, 35: 285.

tibicen, Rattus browni, 70: 123.

trobrius, Hipposideros diadema, 67: 276.
ultra, Thetomys gracilicaudatus, 77: 281.
Unicomys, 59: 259.

Vespadelus, 55: 95 & 94: 351.

victor, Phascolarctos cinereus, 60: 139.
waitei, Pseudomys (Leggadina), 47: 290.
Wombatula, 55: 35 & 94: 150.

Aust. Zool. 18(3), 1975

pe «

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PROb( lls SRR Le oh opr

ROYAL ‘ZOOLOGICAL SOCIETY OF NEW SOUTH WALES
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Patron

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COUNCIL, 1975
me i ; President:
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a phe M.I. Biol. F.S.LH.
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Ni = THE SOCIETY

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VOLUME 18 en ae 1975

Contents

Eptatretus lone ie Spy) NeW. faoceh ( Family Epeterencdeys
South Australia, with a key to the 5-7 gilled See R. STRAH.

A taxonomic revision of the Gi5i aurea complex ( Anura: Hylidaey“s in sou

eastern Australia. GIEELAN PY COURRICE: and GORDON” CE GRIC 1G

Courtship of the platypus, Ornithorkyncbus anatinus. oe STRAHAN :
= hes THOMAS .. oe gta ame seaarenesurrmnsen nines Ree

capoda: Parastacidae) from south-western Tasmania. P. S. LAK]

K. J. NEWCOMBE

Rott paper —

ihe identity of the common keyhole foes of south- -eastern aise
urellidae). W. F. PONDER igor nemo soeneireinc

Memorial notice and Boece —_— os teen
Ellis Le Geyt Troughton, C.M.ZS., FRZS. (1895, 1974). Pik ‘

Sh
: =
Rickard ae
a

5 GO SIFY

THE

AUSTRALIAN
ZOOLOGIST

Volume 19, Part |
September, 1976

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THe AUSTRALIAN: ZOOLOGIST
VOLUME 19 1976 PART |

Studies of an Island Population
of Antechinus minimus (Marsupialia, Dasyuridae)

J. W. WAINER
Department of Zoology, University of Melbourne, Parkville, Victoria, 3052.

ABSTRACT

Field studies including a grid trapping and mark, release, recapture programme,
of an island population of Antechinus minimus indicated that population density
was exceptionally high (80 individuals per hectare). Males were significantly
more mobile than females. Although the sequence of life history events closely
corresponds to that of most other members of the genus, the timing of breeding
is 2-3 months earlier than in the other south-east Australian species. An hypo-
thesis for this reproductive phase difference is put forward.

INTRODUCTION

Of the four south-east Australian species of the dasyurid marsupial genus
Antechinus, A. minimus (the swamp phascogale) is the least common and most
restricted in distribution. The mainland subspecies A. minimus maritimus seems to
be limited to a few scattered coastal localities from Robe in South Australia to
Wilson’s Promontory in Victoria (Wakefield and Warneke, 1963, 1967) and the
Casterton district in western Victoria.

The only member of the genus on which there is much recent literature is
A. stuartii, a common eastern Australian species (Barnett, 1973, 1974; Braithwaite,
1973, 1974; Marlow, 1961; Wood, 1970; Woollard, 1971; Woolley, 1966).
Most work on A. minimus has been taxonomic (Finlayson, 1958; Tate, 1947;
Wakefield and Warneke, 1963) although Green (1972) supplied notes on some
broader aspects, including field observations, of the Tasmanian and Bass Strait
islands nominate form.

The present paper reports on some findings, principally concerning population
density and reproductive biology, of a field study of an insular population of
A. minimus.

Aust. Zool. 19(1), 1976 1

J. W. WAINER

MATERIALS AND METHODS
1. Stupy AREA

Field work was carried out on Great Glennie Island, an island in Bass Strait
about 6 km west of Wilson’s Promontory. The island is elongate (N-S), about
2.5 km long, with an area of approximately 66 hectares (160 acres). The study
area was on a northern slope in the Glennie Cove region at an altitude below 50 m.

The parent material is a grey, coarse-grained granite which is locally weathered
with the development of coarse, sandy and often humus-rich soil.

Being a relatively small unprotected land mass, Glennie Island has a uniform
maritime macroclimate. It is assumed that weather conditions are comparable with
those of the nearest Bureau of Meteorology weather station, the Lighthouse at
South-East Point, Wilson’s Promontory, at an altitude of 100 m (see Bureau of
Meteorology, 1969).

The vegetation of the study area is relatively uniform in character and typical
of vegetation of much of the island. In the more rocky and sheltered situations
there is a closed heath structural formation to a height of about 2 m. The upper
stratum of dominant plants consists of a diversity of shrubs including silver banksia
(Banksia marginata), coast tea tree (Leptospermum laevigatum), white correa
(Correa alba), dusty daisy bush (Olearia phloggopappa) and common boobialla
(Myoporum insulare). There is usually an understorey of Poa tussocks, the density
of which appears to depend upon the intensity of light penetrating the shrub layer.
In exposed areas where soil is often humus-rich and moderately deep, tussock
grassland of the blue tussock grass (Poa poiformis) predominates, often inter-
spersed with small shrubs, especially hoary sunray (Helipterum albicans), and
many low succulent herbs including bower spinach (Tetragonia implexicoma),
saloop (Rhagodia hastata) and rounded noon-flower (Disphyma australe).

2. FIELD TECHNIQUE

Three visits to Glennie Island in 1975 took place in April, May and September.
For the collection of data on movement, home range and population density a
programme of grid live-trapping was undertaken, using aluminium Elliott Type B
traps (32 cm x 9 cm x 9 cm). A rectangular grid of five columns and six rows
(30 stations) was layed out with adjacent trap stations placed 10 m apart. Since
many Rattus fuscipes also entered the traps, three traps were placed at each station.

3. MOVEMENTS AND POPULATION DENSITY

As some individuals from outside the periphery of the grid entered traps it
was necessary to estimate the effective area trapped to calculate abundance of A.

2 Aust. Zool. 19(1), 1976

ISLAND POPULATION OF ANTECHINUS MINIMUS

minimus. The measure used was the average distance moved between successive
captures (Av.D.), which was shown by Brant (1962) to be a good measure of
the width of the boundary strip around the grid from which individuals were
trapped. Av.D. was calculated for each recaptured animal and mean values for
males, females and the population were obtained.

To estimate the number of animals in the effective area trapped a mark,
release, recapture programme was conducted. The grid was trapped on three nights
in May when animals were marked by toe-clipping and released at the site of
capture. 3

RESULTS
1. TRAPPING SUCCESS
(a) GENERAL

A total of 65 individual A. minimus (334, 322) was trapped at Glennie
Island in the 1500 trap-nights of the study. Of these, 15 (104, 52) were
recaptured in a later trapping session. There were 74 recaptures in all with from
one to seven recaptures per individual. With a total of 152 captures the overall
trapping efficiency was 10%/trap-night. Trapping efficiency in May was 23%/
trap-night.

(b) Grip TRAPPING

Over the three nights of grid trapping 34 animals (22¢, 122) were captured,
with a total of 99 captures (746, 252). The average number of captures per
male was 3.36 and per female was 2.08. Therefore the ratio of males to females
captured was 1.62.

2. MOVEMENT

The mean Av.D. values for the 16 males and 7 females trapped more than
once on the grid were 19.3 m and 13.4 m respectively. These values are signifi-

cantly different at the 95% probability level. The Av.D. for the population was
16.4 m.

3. DENSITY ESTIMATE

Table 1 shows grid recapture trapping results. The three Lincoln Index
(Lincoln, 1930) estimates of population size between trapping nights 1 and 2, 1
and 3, and 2 and 3 are 42, 26 and 34 respectively. The population size calculated
using Hayne’s method (Hayne, 1949) is 36, the value used for the density
estimate. Using 15 m as an approximately minimum value of Av.D. and diameter
of the boundary strip, the area of effective trapping was 0.45 hectares (1.1 acres).
The population density was therefore 80 individuals per hectare (32 per acre).

Aust. Zool. 19(1), 1976

iv)

J. W. WAINER

TABLE 1
Recapture trapping results of A. minimus on the grid at Glennie Cove, May, 1975.
Night ‘Number of animals trapped Number of animals recaptured f
3 2 Total
May 24th ...._.... 18 — —
hs) Le a ae 15 iB

May 27th

4. REPRODUCTION

(a) Sex RATIO

The sex ratio of males to females in April was 1.73:1 (194, 112) and in
May was 1.50:1 (246, 162 ). The preponderance of males is probably partly due
to their greater mobility. The total number of each sex trapped (3346, 322)
gives a false idea of the overall sex ratio since the September trapping session
took place after the postmating male die-off.

Male die-off

Young suckled in nest
————————————————

Young in pouch
<———— HH __?

Parturition
SSS

Gestation

Mating

ee oe

Pre-reproductive animals

JAN FES MAR APR MAY INES SUEY AG SEP OCT NOV DEC

Fig. 1. Timing of life history events of A. minimus at Glennie Island.

4 Aust. Zool. 19(1), 1976

ISLAND POPULATION OF ANTECHINUS MINIMUS

(b) Lire CycLe

An estimate of the timing of the important features of reproduction at Glennie
Island is shown in Fig. 1. In late May, mating was occurring. Most parturitions
occurred in July and the neonates remained in the pouch until late August to mid
September. In early September some females had nest-young and others still had
pouch-young. Young were then suckled in the nest until mid or late November;
in early November 1975 no juveniles were found, whereas in early December 1974
they were abundant (Barnett, pers. comm.). Although conclusive evidence is
wanting, the male die-off, which is universal, probably occurs early in July, because
in late July 1973 only females with pouch young were present (Robinson, pers.
comm.). No second-year females were encountered.

(c) LiTTER SIZE

Of the 10 females captured in September three had pouch-young, the litter
sizes being 6, 8 and 8. All females examined possessed 8 teats. The number of
well developed nipples of females with nest-young is a reliable indication of the
number of nest-young (Woolley, pers. comm.), and the five females with nest-
young had 6, 7, 8, 8 and 8 well developed nipples; therefore the average number
of young per mother for the 8 females with young was 7.4. The remaining two
of the 10 animals captured had not bred.

DISCUSSION

TRAPPING SUCCESS

The trapping efficiency values at Glennie Island of 10%/trap-night overall,
and 23% /trap-night during winter, when trapping is most productive, are much
higher than values obtained elsewhere. At the Parker River Inlet, Otway Ranges,
Vic., in November when juveniles were mobile, a trapping efficiency of 3.69% was
obtained (Norris, pers. comm.), and Green (1972) found an efficiency of approxi-
mately 2.5% for Tasmanian populations in the most suitable habitat in winter.

MOVEMENTS AND POPULATION DENSITY

Close trap spacing gives capture points too close to disclose the true home
range (Chitty, 1937; Stickel, 1954). All or nearly all of the animals on the grid
were trapped on the first two nights of the grid trapping programme, suggesting
that the area was saturated by traps. Thus Av.D. was kept at a minimum, giving
underestimates of the effective area of trapping. Nevertheless, the Av.D. values
for males and females, 19 m and 13 m respectively, are 50%-70% those found
by Wood (1970) and closer to those found by Braithwaite (1973), both for
A. stuartit.

Mobility of males was found to be significantly greater than that of females.
possibly because males were actively defending territories at this time.

Aust. Zool. 19(1), 1976 5

J. W, WAINER

REPRODUCTION

Breeding is restricted to winter, conforming with the limited information
available on Tasmanian individuals (Green, 1972). As in the three other
south-east Australian species of Antechinus (Wakefield and Warneke, 1963,
1967; Woolley, 1973), A. minimus appears to be monoestrous. Males and females
are sexually quiescent except during the very limited breeding season. There is a
total postmating mortality of males, a phenomenon which Lee e¢ al. (in press)
have investigated in A. stuartii.

Breeding is 2-3 months earlier than in A. stwartii, in which mating occurs in
late August in southern Victoria (Lee et al., in press). This difference in timing
may be related to their habitats and availability of food. A. stuwartii shows extensive
arboreal activity (Wood, 1970) and probably hunts for much of its food, consist-
ing of invertebrates including adult insects, on the trunks of trees. Thus it synchro-
nises its breeding with the spring flush of insects. On the other hand A. minimus
is terrestrial and a large part of its diet consists of larval insects from the soil
as well as other terrestrial arthropods (Wainer, 1975). In spring and early
summer when A. stuartii females are rearing young, and as a result probably require
relatively more food, adult insects are at a peak of abundance; however A.
minimus appears to synchronise its breeding with the peak of larval insects in
winter.

In the Glennie Island population no parous second year females, which can
be distinguished from virgin females on the basis of development of nipples
(Woolley, 1966), were found. The situation in A. minimus, therefore, may differ
from that in A. stwartii where some females survive to a second breeding season

(Wood, 1970).

ACKNOWLEDGEMENTS

The National Parks Service of Victoria kindly granted permission to conduct a live-
trapping programme at Glennie Island and their co-operation is gratefully acknowledged.
Also I wish to express my gratitude to Dr. Angus Martin and Mr. Stephen Morton for
reading and commenting on the manuscript, and to Dr. Martin for supervising the project.

REFERENCES

BARNETT, J. L. (1973). A stress response in Antechinus stuartii (Macleay). Aust. J. Zool.
21, 501-513;

BARNETT, J. L. (1974). Changes in the hydroxyproline concentration of the skin of
Antechinus stuartii with age and hormonal treatment. Aust. J. Zool. 22, 311-318.

BRAITHWAITE, R. W. (1973). An ecological study of Antechinus stuartii (Marsupialia:
Dasyuridae). M.Sc. thesis, University of Queensland, Brisbane. 143 pp.

BRAITHWAITE, R. W. (1974). Behavioural changes associated with the population cycle
of Antechinus stuartii (Marsupialia). Aust. J. Zool. 22, 45-62.

6 Aust. Zool. 19(1), 1976

ISLAND POPULATION OF ANTECHINUS MINIMUS

BRANT, D. H. (1962). Measures of the movements and population densities of small
rodents. Univ. Calif. Publs. Zool. 62, 105-184.

BUREAU OF METEOROLOGY (1969). “Climatic Averages Australia.”
CHITTY, D. (1937). A ringing technique for small mammals. J. anim. Ecol. 6, 36-53.

FINLAYSON, H. H. (1958). A case of duplex convergent resemblance in Australian
mammals, with a review of some aspects of the morphology of Phascogale (Antechinus)
swainsoni Waterhouse and Phascogale (Antechinus) flavipes Waterhouse. Proc. Roy. Soc.
S. Austea, 141i 5.

GREEN, R. H. (1972). The murids and small dasyurids in Tasmania. Parts 5; 6 and 7.
Rec. Queen Vict. Mus., Launceston, Tas. 46, 1-34.

HAYNE, D. W. (1949). Two methods for estimating population from trapping records. J.
Mammal. 30, 1-18.

LEE, A.K., A. J. BRADLEY & R. W. BRAITHWAITE (in press). Life history and male
mortality in Antechinus stuartii. In: Gilmore, D. and Stonehouse, B. (eds.) The Biology
of Marsupials. Macmillan, London.

LINCOLN, F. C. (1930). Calculating waterfowl abundance on the basis of banding returns.
US. Dept. ‘Agric: Circ. 178, 1-4.

MARLOW, B. J. (1961). Reproductive behaviour of the marsupial mouse, Antechinus
flavipes (Waterhouse) (Marsupialia) and the development of the pouch young. Aust. J.
Zool. 9, 203-218.

STICKEL, L. F. (1954). A comparison of certain methods of measuring ranges of small
mammals, J. Mammal. 35, 1-15.

TATE, G. H. H. (1947). Results of the Archbold Expeditions No. 56. On the anatomy and
classification of the Dasyuridae (Marsupialia). Bull. Amer. Mus. Nat. Hist. 88, 101-155.

WAINER, J. W. (1975). Ecological studies of Antechinus minimus (Geoffroy) (Marsupialia:
Dasyuridae) on Glennie Island, Bass Strait. B.Sc.(Hons.) thesis, University of Melbourne.
47 pp.

WAKEFIELD, N. A. & R. M. WARNEKE (1963). Some revision in Antechinus (Marsu-
pialia) 1. Victorian Nat. 80, 194-219.

WAKEFIELD, N. A. & R. M. WARNEKE (1967). Some revision in Antechinus (Marsu-
pialia) 2. Victorian Nat. 84, 69-99.

WOOD, D. H. (1970). An ecological study of Antechinus stuartii (Marsupialia) in a south-
east Queensland rainforest. Aust. J. Zool. 18, 185-207.

WOOLLARD, P. (1971). Differential mortality of Antechinus stuartii (Macleay): nitrogen
balance and somatic changes, Aust. J. Zool. 19, 347-353.

WOOLLEY, P. (1966). Reproduction in Antechinus spp. and other dasyurid marsupials. [n:
Rowlands, Comparative Biology of Reproduction in Mammals.

WOOLLEY, P. (1973). Breeding patterns, and the breeding and laboratory maintenance of
dasyurid marsupials. Expt. Animals 22 (suppl.), 161-172.

Aust. Zool. 19(1), 1976 é

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Maturational and Seasonal Moult in the New Holland
Mouse, Pseudomys novaehollandiae

CATHERINE M. KEMPER
School of Biological Sciences, Macquarie University, North Ryde, N.S.W. 2113

ABSTRACT

The post-juvenile moult is described in a captive colony of Pseudomys novae-
hollandiae. This moult, which may be found in animals aged 35 to 105 days,
progresses from the ventral to the dorsal surface of the body, ending on the
head and near the base of tail. There appears to be no post-subadult moult in
this species. The occurrence of moulting in a wild population was also studied.
The post-juvenile moult occurs mainly from October to January and a seasonal
moult between January and June.

INTRODUCTION

Moult is the process of losing hair or replacing one pelage with another. Two
major types of moult may be recognised in small rodents, those which are of a
seasonal nature and occur irrespective of age and those which are associated with
the development of the animal and are therefore called maturational moults. Two
maturational moults, post-juvenile and post-subadult, separate three pelage phases
(juvenile, subadult and adult) in almost all species of Cricetidae (Buhlow, 1970;
Ecke and Kinney, 1956; Layne, 1968; McManus and Zurich, 1972), Muridae
(Borum, 1954; Saint-Girons, 1967) and Heteromyidae (Eisenberg, 1963), studied
to date. However, Linzey and Linzey (1967) found only one maturational moult
in the cricetid, Ochrotomys nuttalli. Brief mention has been made of moult in the
Australian genus Pseudomys (Green, 1968; Finlayson, 1944) but not enough
information was presented to say whether one or two maturational moults were
present in the two species studied.

Seasonal moults, in spring and autumn, are known to occur in several species
of small rodents (Buhlow, 1970; Layne, 1968; Linzey and Linzey, 1967; Saint-
Girons, 1967) but Brechner and Kirkpatrick (1970) reported that moulting
occurred throughout the year in Mus musculus.

The New Holland mouse, Pseudomys novaehollandiae is an Australian rodent
inhabiting several localities in New South Wales and Victoria. In the wild it is a
seasonal breeder, giving birth between September and December each year. It is

Aust. Zool. 19(1), 1976 9

CATHERINE M. KEMPER

weaned during the fourth week of age but does not usually appear in trapped
samples until at least five weeks. In the course of studying the growth and
development of P. novaehollandiae information was obtained on the post-juvenile
moult. It was felt that this might prove useful in the age determination of wild
populations so information was gathered on all trapped animals. As a result, it
was found that seasonal moult existed as well.

MATERIALS AND METHODS

The sequenece and timing of the post-juvenile moult was observed in 35 live
P. novaehollandiae from 10 litters. These laboratory-raised animals were progeny
of mice obtained from the Nelson Bay and Smith Lake areas of New South Wales.
They were maintained under the laboratory conditions described in Kemper (1976).

A pelage dye (Durafur Black) was applied to the first litter studied but this
proved detrimental to the health of the animals so all other animals used were
not treated in any way. The new pelage was simply noted to be present beneath
the juvenile pelage by its shorter length and browner colouration. The chin, manus,
pes and tail were not investigated for moulting because of the difficulty in
observing the new fur in these areas. Not all animals were followed through from
before the beginning to the end of moult and observations were made at different
ages for each animal. To facilitate computations these were grouped into five-day
intervals. Table 1 was obtained by totalling the number of animals of a certain
age interval for each moult stage and taking that figure as a percentage of the
total number of observations for that age. This was necessary because of the
different sample sizes and because some of the animals were observed more than
once during an age interval.

The skins of 150 field-trapped animals (from the same localities as the original
laboratory stock) were examined for evidence of moulting. These were obtained
Over a two-year period from June 1972 to May 1974. Most were prepared as
museum round skins but some were made into flat pelts. Moult was noted either
by the emergence of new fur or by recording the position of darkly pigmented
areas on the inside of the skin (pigment is deposited at sites of new hair forma-
tion), or both.

RESULTS

PosT- JUVENILE MOULT

The sequence of the post-juvenile moult was much the same for all animals
observed so it was possible to categorise this progression into a set pattern of
phases which are listed below. An illustration of each phase appears in Fig. 1.
The values in parentheses are ages when most animals were in the phase in question.

10 Aust. Zool. 19(1), 1976

MOULT IN THE NEW HOLLAND. MOUSE

Phase I

Moulting begins in the mid and anterior ventral regions. This may include
the ventral surfaces of the forelimbs (35-50 days).

Phase II

Moulting is occurring over most of the ventral surface, except that of the
hindlimbs, and may have reached the dorsal surfaces of the forelimbs (45-55 days).

Phase III

Moulting has progressed to the sides of the body including the dorsal and
ventral surfaces of both fore- and hindlimbs. The new pelage in the mid-ventral

region has grown enough so that it is indistinguishable from the juvenile pelage
(50-75 days).

Phase IV

The ventral surface has completed moulting, except for the hindlimbs. On
the dorsal surface the moult has spread to the mid-back to form a saddle joining
the sides and is complete on the forelimbs (60-80 days).

Phase V

Moulting is visible all over the dorsal surface except the limbs, head and
base of the tail (55-100 days).

Phase VI

Moulting is only visible on the head and near the base of the tail (65-105
days).

Observations on each animal were later placed into one of the above phases
even though slight variations did exist when different body parts were moulting.
Table 1 presents these data as a percentage of total observations for each five-day
age interval. Two additional categories were included (not started and finished
moulting ) to give some idea of when most animals begin and end the moult. It
can be seen from Table 1 that most animals begin the post-juvenile moult at
35-45 days of age and that all have started by 55 days, and end it by 100 days
but this may occur as early as 70 days and as late as 110 days.

Only 13 of the 35 animals were followed from before the beginning of
moulting to its completion. The average age at commencement was 43-46 days
and at completion was 85-91 days (Table 2). The average duration of the moult
for these animals was 41-50 days. Although the sample size is small, there appears

Aust. Zool. 19(1), 1976 1

CATHERINE M. KEMPER

Stippled areas indicate

Phases of the post-juvenile moult in P. novaehollandiae.

1.

Fig.

to be no obvious difference in timing of the post-juvenile moult between sexes.
Some variation in timing was observed between litters. In all three litters examined.

littermates began the moult within a few

days of each other. The time of completion was more variable than the time of

from beginning to end of moulting,

onset, although this may have been due at least in part to the difficulty in telling
exactly when the moult ended. Individuals of other litters also showed similar

phases of the moult at most times of observation.

19(1), 1976

Aust. Zool.

12

MOULT IN THE NEW HOLLAND MOUSE

parts of the body in the process of fur replacement.

1S

between 100 and 200 days of age, were

examined for evidence of a post-subadult moult. Forty-two percent of these had

b]

her surface as is the case with the post-juvenile moult. It

therefore, that this is not a post-subadult moult but one of the periodic

areas of pigmentation on the inside of the skin. In most this was confined to the
anterior and mid-ventral regions and to the head and none had extensive areas of

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19(1), 1976

Aust. Zool.

CATHERINE M. KEMPER

TABLE 1
Percentage of P. novaehollandiae at moult stages for age intervals from 35-110 days.

Age

Moult stage 35. 402 45° 50) °55.- 60065270 735. SO. 85.90) 295... 100% 105 eis
Not started 80) 446520 ©) 6c. OF eure Oe WO Ore Ose 0 0 0
I 20" S05 8205.13 296" JO OO Os Ost Oem Ono 0 0 0

Il Os 6s 48" 19 328 SiO NO Oe Oe OO rao) 0 0 0

iil Or OS 8592 238 0 2 5 oe Or ea 0 0 0 0

IV Oe 10 Oe? S66 oD AS 8 29382 Os ZOO 0 0 0

Vv Oo Oo 4 SMOG SiS 2 ae OA 26" 77) 30s, ON 22 0 0

VI Os OO 08 6 0S Pe ee 2 265 Sn 50 Ole S TT AUG 0
Finished OS 205 Oe OO ON Oe 120 od 8 9 OY Gy 0 100
Total animals Sp G24 30 NS 2522 6, le Sf) EO 9 I 1

TABLE 2

Age at commencement and completion of the post-juvenile moult in three litters of P. novae-
hollandiae. The sixth member of the first litter was not included because it was develop-
ilar mentally retarded.

Litter size Sex Age at commencement (days) Age at completion (days) Duration
6 3} 3h7/ 67-70 40
} 38-40 78-82 39-44
J 38-40 73-78 33-40
2 38-40 67-70 27-32
3 38-40 83-87 43-49
4 2 52-57 91-99 34-37
6 47-51 91-99 40-52
d 52-57 91-99 37-47
L 47-51 105-110 54-63
4 3g 41-43 92-100 49-59
3 41-43 85-91 42-50
3 41-43 92-100 49-59
Ee 41-43 92-100 49-59
Mean 43-46 85-91 41-50

SEASONAL MOULT

Fig. 2 illustrates the percentage of wild animals moulting during each month
of the year for two age classes (C1 — those less than one year of age; and C2 —
those older than one year). Two different types of moult were found in C1, the
post-juvenile moult and another, which was assumed to be a seasonal one. Although

14 Aust. Zool. 19(1), 1976

MOULT IN THE NEW HOLLAND MOUSE

100

7% MOULTING

ON Ded oF
(4) (10) (17) (16) (9)

MONTH OF CAPTURE

Fig. 2. Percentage of monthly captures of P. novaehollandiae in the process of moulting.
Age class C1, — post-juvenile moult, - - - - seasonal moult; age class C2...-.-
seasonal moult. Numbers in parentheses are sample sizes.

there is a possibility that the second type is a post-subadult moult this was
discounted in view of the laboratory findings and because a seasonal moult was
being experienced by C2 at the same time. Some of the observations of seasonal
moult in C1 in January and February may have been, in fact, post-juvenile moults
because it is difficult to differentiate these two moults in the later stages.

The post-juvenile moult was found from October to January and also in May,
the latter resulting from a trapping period at Smith Lake in 1973 when a small
population was found to be breeding out of season. Seasonal moult was found in
C1 from January to June and in C2 from January to March (Fig. 2).

Aust. Zool. 19(1),.1976 15

CATHERINE M. KEMPER

The seasonal moult in both age groups did not appear to follow the same
pattern as the post-juvenile moult, although this was difficult to tell without
following several animals through. The summer pelage is easily distinguished from
the winter one because of its less dense nature and more grizzled appearance.
As would be expected there seems to be a greater proportion of guard hairs in
the summer pelage thus adding to its uneven texture.

DISCUSSION

The juvenile pelage covers the whole body surface by 13 days of age in
P. novaehollandiae although it is not of maximum length (Kemper, 1976). It is
soft, fine and grey in colour and is thus distinguishable from the coarser, brown
pelage which replaces it. If, as suspected from the results in this study, there is
only one maturational moult in this species, then, the later pelage is the adult one.
This is in contrast to most species of rodents studied to date which have three
basic pelage phases — juvenile, subadult and adult.

Buhlow (1970) has reviewed the literature pertaining to the timing of both
maturational moults in the Muridae and Microtinae. In all species which experience
a second moult this follows within two weeks of the end of the first. The duration
of the post-juvenile moult is generally much shorter (12-31 days) than in P.
novaehollandiae in which it lasts about 45 days. Rattus norvegicus also has a longer
post-juvenile moult of 33-42 days duration (Buhlow, 1970). In Ochrotomys
nuttalli, which has only one maturational moult, it lasts 25 (females) and 29
(males) days (Linzey and Linzey, 1967). In the small sample investigated no sex
differences were found in P. novaehollandiae. Both other species of Pseudomys, in
which moult has been reported, have long post-juvenile moults. Pseudomys albo-
cinereus (= apodemoides) has a first moult (probably the post-juvenile one) at
7-15 weeks old (Finlayson, 1944) and Pseudomys higginsi, at 50-100 days
(Green, 1968).

The progression of the post-juvenile moult follows roughly the same pattern
in all species studied (Borum, 1954; Buhlow, 1970; Ecke and Kinney, 1956;
Layne, 1968; Linzey and Linzey, 1967; McManus and Zurich, 1972; Saint-Girons,
1967). Moult begins on the ventral surface, spreads to the dorsal surface and
ends on the head and at the base of the tail. However, it may be that P. higginsi
and P. albocinereus differ somewhat in that the former begins to moult on the
flanks (Green, 1968) and the latter on the nape of the neck (Finlayson, 1944).

Most mammals in temperate climates have two pelage phases related to
season, the winter one being denser for protection against cold. In P. xovae-
hollandiae the moult which results in the winter pelage takes place between
January and June but there is no obvious new growth of fur in the spring. It

16 Aust. Zool. 19(1), 1976

MOULT IN THE NEW HOLLAND MOUSE

may be that the spring moult is simply a result of thinning of the underfur by
loss of hairs rather than a complete new pelage. This change could be so gradual
that it would not be detected by the methods used in this study.

REFERENCES

BORUM, K. (1954). Hair pattern and hair succession in the albino mouse. Acta pathol.
microbiol. Scandinav. 34, 521-541.

BRECHNER, R. E. & R. D. KIRKPATRICK (1970). Molt in two populations of the
house mouse, Mus musculus. Proc. Ind. Acad, Sci. 79, 449-454.

BUHLOW, E. (1970). Untersuchungen uber den haarwechsel bei Schermausen, Arvicola
terrestris (L., 1978). Zool. Anz. 184, 18-32.

ECKE, D. H. & A. R. KINNEY (1956). Aging meadow mice, Microtus californicus, by
observation of molt progression. J. Mamm. 37, 249-254.

EISENBERG, J. F. (1963). The behaviour of heteromyid rodents. Univ. California Publ.

Zool. 69, 1-100,

FINLAYSON, H. H. (1944). A further account of the murid, Pseudomys (Gyomys)
apodemoides Finlayson. Trans. Roy. Soc. S.A. 68, 210-224.

GREEN, R. H. (1968). The murids and small dasyurids in Tasmania, Parts 3 and 4.
Rec. Queen Victoria Mus. Launceston No. 32, 19 pp.

KEMPER, C. M. (1976). Growth and development of the Australian murid Pseudomys
novaehollandiae. Aust. J. Zool. 24, 27-37.

LAYNE, J. N. (1968). Ontogeny. In: Biology of Peromyscus (Rodentia). Amer. Soc.
Mamm. special publication no. 2. 592 pp.

LINZEY, D. W. & A. V. LINZEY (1967). Maturational and seasonal molts in the golden
mouse, Ochrotomys nuttalli. J. Mamm, 48, 236-241.

McMANUS, J. J. & W. M. ZURICH (1972). Growth, pelage development and maturational!
molts of the Mongolian gerbil, Meriones unguiculatus. Amer. Midland Nat. 87, 264-271.

SAINT-GIRONS, M. C. (1967). Etude du genre Apodemus Kaup, 1892 en France.
Mammalia 3/7, 55-100.

Aust. Zool. 19(1), 1976 17

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Relative Efficiency of Two Types
of Small Mammai Trap in Eastern Australia

BARRY J. FOX

School of Biological Sciences,
Macquarie University, North Ryde, N.S.W. 2113

and

HEIMO G. POSAMENTIER
Australian Museum, Sydney, N.S.W. 2000

ABSTRACT

A comparison of the response of small mammals to snap traps and live traps
was carried out using two methods. Overall captures indicate that at the 5%
level of significance snap traps are more effective than live traps. Subtotals
for each method and for each genus, however, show no significant difference.
Advantages to be gained from the use of both types of trap are discussed. It
is concluded that snap traps should be used for large-scale survey work unless
the quality of the specimens is of prime importance.

INTRODUCTION

There are no published records in Australia on the response by small
mammals to different types of traps. Cowan and Barnett (1975) do discuss new-
object reactions but under laboratory conditions. Work done overseas indicates
that the trap response may be species specific (Edwards, 1952). Some authors
found live traps to be more efficient (Cockrum, 1947; Sealander and James,
1958), others that snap traps were more efficient in trapping small mammals
overall (Smith, 1972; Buckner, 1957; Van Vleck, 1968). Response to traps is
also discussed by Calhoun (1959), Crowcroft and Jeffers (1961), Golley (1961)
and Eberhardt (1969).

The authors of this paper have experience in trapping small mammals on
the East Coast of New South Wales using live and snap traps, and have found
no apparent difficulty in trapping any particular species effectively using either
type of trap. In view of the differential findings by the authors quoted above, and
the often species specific response, we found it necessary to look at trap response
in Australia. With many more groups now studying small mammal field biology
in Australia we hope that other more specific studies will follow this one.

In this paper two types of traps, live and snap traps, were tested for their
relative efficiency in the field. The work was carried out during a survey of the

Aust. Zool. 19(1), 1976 19

BARRY J. FOX and HEIMO G. POSAMENTIER

rainforests on the East Coast of Australia, jointly undertaken by the Australian
Museum, Sydney, and the Queensland Museum, Brisbane. Besides trap response,
the advantages of both trap types are discussed.

METHODS

The trapping program was carried out at six sites in different eastern
Queensland rainforest types between Bundaberg and Mackay. Trap types compared
were: snap trap — household break-back rat trap; live trap — Elliott 33 x 10 x 10
cm collapsible aluminium live trap. All traps were baited with a mixture of
peanut butter, rolled oats and bacon fat. Traps were checked each morning and
rebaited or reset where necessary.

Trap response was compared using two methods:

A. Comparison on adjacent areas

At each site one area was set with six snap trap plots 100 m apart at the
corners of a hexagon, each plot comprised 20 traps equally spaced on a cross with
four 15 m arms. A central plot was set with 32 live traps equally spaced on a
cross with four 16 m arms. The adjacent area was identical except that the
central plot was set with snap traps and the six radial plots with live traps.

B. Comparison by use of paired traplines

At each site one or two index lines of 20 or 25 points was set up with
trapping points 10 m apart. At each point one snap trap and one live trap were
set within 1 m of each other but not immediately adjacent, being placed to catch
small mammals (next to logs, tree trunks, etc.).

Method A was used on the first three sites while method B was used at
all sites. At the first three sites the number of traps set off without a mammal
capture was noted for each site.

TABLE 1
RATTUS AND MELOMYS CAPTURES

Rattus Melomys
Number of Snap Live Snap Live
Site No. Nights Trap Trap Trap Trap
1 5 4 4 1 0
2 5 3 1 2 0
3 5 0 0 19 13
4 4 5 2 1 1
5 4 18 18 0 0
6 4 25 16 0 0
37 55, 41 23 14

20 Aust. Zool. 19(1), 1976

SMALL MAMMAL TRAP EFFICIENCY

RESULTS AND DISCUSSION

Five species were trapped: Rattus fuscipes, R. tunneyi, R. leucopus, Melomys
sp. and Antechinus stuarti. The low capture rate, 136 animals in 5574 trapnights
(2.4%), meant that data could only be analysed at a genus rather than a species
level (Table 1). No significant difference is apparent between trap types when
considered at a genus level. There is no significant difference between types when
each method is considered separately (Table 2). The paired trapline method was
considered to be the better method to use for comparing trap types. The Chi-
square test used was a one sample test with one d.f. testing equal probabilities of
capture (Siegal, 1956).

All species taken in this study entered both types of trap and previous
experience provided similar findings for other east Australian species of small
mammals. The greater overall efficiency of snap traps may be related to the
sreater accessibility of bait on snap traps or to a lesser new object response toward
snap traps compared to live traps. The latter view is also held by Buckner (1957)
and supported by two direct observations by the authors, where animals shied
away from a live trap and readily approached the snap trap nearby. A single
animal was also observed attempting to get at the bait near the back of a live
trap from the rear of the trap and left without finding the open entrance to the
trap. Reduced new object response towards snap traps would mean that they could
effectively sample a given population of small mammals sooner than live traps.
Due to low capture rates this could not be evaluated directly in this study.

The results of this study certainly indicate that differences in trap response
to the two types of traps used are not sufficient to exclude the consideration of
other factors in the choice of traps. These factors depend on the type of study and
data required. If good specimens are required then live traps should be employed
to ensure fresh specimens and avoid skull damage. However, if a large survey or
short-term studies are envisaged the use of snap traps is suggested as handling is
simpler due to their smaller size, lighter weight and reduced cleaning time. These

TABER, 2

ANALYSIS OF TRAP DATA FOR ALL SPECIES

Snap Live
Trapnights Trap Trap Chi-Square
1622 Pamed; Erapune?Captures, ©.2.021./.20. 55 39 Done
3952 Adjacent Area Captures. ...-.:..0:.c20.2..-. 25 17 £52
S514 Total Captures Both Methods ........ 80 56 4,.24*
Traps Sprung Without Mammal
5102 CAPLIO i es rhe de dette Bees aa 473 74 326.00**
*Significant at 5% level **Significant at 0.1% level

“Aust. Zool. 19(1), 1976 21

BARRY J. FOX and HEIMO G. POSAMENTIER

factors together with the much lower cost of snap traps mean that many more
snap traps can be used in any study thus increasing trap density and so over-
coming problems such as having traps sprung without mammal capture and bait
loss both of which occur more frequently with snap traps (Table 2). Extensive
trapping experience in eastern N.S.W. has not shown such loss of effective traps
to be a significant problem (Posamentier and Recher, 1974; Posamentier, 1975).

From the results of this study and the reasons outlined above the authors
suggest the use of snap traps for large-scale general surveys of small mammals.
However, the variability in species trap response (Smith, 1972) indicates the need
for further studies looking at individual species, a variety of habitats and other
trapping techniques to provide more specific information. In addition, a study
comparing the rates of capture by different types of traps is needed to indicate
whether snap traps sample a given population of small mammals in a shorter
period than live traps.

ACKNOWLEDGEMENTS

The authors wish to thank Drs. S. S. Clark, G. M. McKay and H. F. Recher and
M. Archer and B. Marlow for useful discussions in the development of this project. The
project was supported by the Australian Biological Resources Study Interim Council.

REFERENCES

BUCKNER, C. H. (1957). Population studies on small mammals of Southeastern Manitoba.

ea =Mamm. 38,.°87-97:

CALHOUN, J. H. (1959). Revised sampling procedure for the North American census of
small mammals (NACSM). U.S. Dept. Hlth. Educat. Welf. Bethasda, No. 10, 1-12.

COCKRUM, L. E. (1947). Effectiveness of livetraps versus snaptraps. J. Mamm., 28, 186.

CROWCROFT, P. & J. N. P. JEFFERS (1961). Variability in the behaviour of wild House

Mice (Mus musculus L.) towards live traps. Zool. Soc. Lond. 137, 537-82.

EBERHARDT, L. L. (1969). Population estimates from recapture frequencies. J. Wildl.
Mgmt. 33, 28-39.

EDWARDS, R. Y. (1952). How efficient are Sta p in taking small mammals? J. Mamm..,
33, 497-498.

GOLLEY, F. B. (1961). Trap and bait Pearce in Cotton Rats. J. Mamm., 42, 106-7.

POSAMENTIER, H. G. (1975). Habitat requirements of small mammals in coastal heath
lands of New South Wales. M.Sc. thesis. University of Sydney, Sydney, Australia.

POSAMENTIER, H. G. & H. F. RECHER (1974). The status of Pseudomys novae-
hollandiae (The New Holland Mouse). Aust. Zool. 18, 66-71.

SEALANDER, J. A. & D. JAMES (1958). Relative efficiencies of different small mammal
traps. J. Mamm., 39, 215-23.

SIEGAL, S. (1956). Nonparametric statistics for the behavioural sciences. McGraw-Hill,
Sydney.

SMITH, M. H. (1972). Relative efficiencies of four small mammal traps. J. Mamm..,

“TPIS, 808-7 3.

VAN VLECK, D. B. (1968). Movements of Microtus pennsylvanicus in relation to de-
populated areas. J. Mamm., 49, 92-103.

22 Aust. Zool. 19(1), 1976

Mammals of Clouds Creek, North-Eastern New South
Wales, and their Distribution in Pine ana Native Forests

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS
Department of Zoology, La Trobe University, Bundoora, Victoria, 3083

ABSTRACT

A survey was made of mammals in an area of 118 km’ in Clouds Creek State
Forest in north-eastern N.S.W. This area is being converted to Pinus spp.
Twenty-seven species of indigenous mammals and seven introduced species were
recorded. The distribution of these species in native and pine forest indicated that
only Rattus rattus, R. lutreolus and Mus musculus inhabited pine forest (4-7
years old) exclusively, although this association was less obvious in older pines
(25 years old) where they were trapped only on the periphery.

Changes in the habitats resulting from forestry practices influenced the distri-
bution and abundance of mammals in the area. Individuals of Rattus fuscipes
and Macropus rufogriseus were able to survive in pine plantings and their
associated grasslands. No individuals of Trichosurus caninus or T. vulpecula
used pine areas exclusively and they are dependent on adjacent native forest.
Other phalangers were sighted only in native forest, including altered habitat
such as grazed woodland. No populations of either Antechinus_ stuartii or
Melomys cervinipes were found away from native forest and the latter species
was trapped only in moist forest.

Macropods were similarly sensitive to habitat changes. Macropus giganteus was
widespread in both native and pine areas, Thylogale thetis occurred on the pine
periphery and M. parma, T. stigmatica and Aepyprymnus rufescens were restricted
to closed and tall open forests.

INTRODUCTION

Tyndale-Biscoe and Calaby (1975) used the eucalypt forests of southern
Australia to illustrate the dependence of certain mammals on specific forest types.
They stressed the scarcity of information on distribution of Australian fauna with
respect to vegetation type.

Calaby (1966) surveyed a large area of the upper Richmond and Clarence
Rivers and demonstrated that north-eastern N.S.W. had a diverse mammalian
fauna. Much of this area has been included as a target area in various proposals
for a woodchip industry currently being considered for northern N.S.W. In
addition, native forests are being cleared for the planting of exotic pines.

This paper reports on a long, intensive survey of a small area of highly
productive native forest and of pine plantations in northern N.S.W. Emphasis
is made of the distribution of the more abundant species between vegetation types.

Aust. Zool. 19(1), 1976 So e2S

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

STUDY AREA AND METHODS

In 1968 an ecological study of two possum species, Trichosurus caninus and
T. vulpecula, was commenced in an area of forest that was partially being
converted to Pinus spp. (How, 1972). This study was extended in 1975 to
include the small mammals in the area. The study area was visited regularly from

mh

4<
Pre Se Wo!

44

<

|

[1 CLOSED FOREST
= TALL OPEN FOREST
OPEN FOREST

3} WOODLAND
GRASSLAND

[_] eine

to
CHAPMANS
lkm PLAIN 4
7km

Fig. 1. Map of the Clouds Creek area showing the localities mentioned in the text. Solid
circles enclose areas of fixed trap localities.

24 Aust. Zool. 19(1), 1976

MAMMALS OF CLOUDS CREEK

1968, except for a period from January 1972 to August 1974. During the five and
a half years spent in the area, various methods were used to assess the mammal
populations, including live trapping, break-back trapping, spotlighting, the collection
of roadkills and chance sightings.

STUDY AREA

The study area of 118 km? is at Clouds Creek (30° 05° S, 152° 37° E) in
north-eastern N.S.W. (Fig. 1). Clouds Creek is situated on the eastern escarpment
of the New England Tableland, about 120 km north-east of Armidale, and is on
the northern edge of the Dorrigo Plateau, 24 km north-east of Dorrigo. Based
on the classification of Specht (1970), four types of tree-dominated communities
occur in the area. These are closed forest, tall open forest, open forest and
woodland; pine plantation and grassland communities also occur in the study area.

Small areas of closed forest occur to the north and the east of the pine
plantation. Species commonly found in this association are Coachwood (Cerato-
petalum apetalum D. Don) and Sassafras (Doryphora sassafras Endl.) along with
several others of commercial value. A southern area of closed forest was cleared
in early 1970 and planted with pines.

In the transitional zone between closed forest and tall open forest, Brush
Box (Tristania conferta R. Br.) is prominent, while tall open forest in the area
is dominated by Tallowood (Eucalyptus microcorys F. Muell.), Sydney Blue Gum
(E. saligna Sm.), White Mahogany (E. acmenoides Schau), Dunns White Gum
(E. dunni Maiden) and Turpentine (Syncarpia glomulifera Sm.).

There is a large area of open forest to the north-west of the plantation
dominated by Blackbutt (E. pilularis Sm.), while the area immediately to the
south of the Forest Commission Headquarters (F.C.H.; Fig. 1) was New England
Blackbutt (E. campanulata Baker and Smith) and Camerons Stringybark (E.
cameronti Blakely and McKie), as codominants prior to its clearing in 1970.

Grazed woodland to the west of the F.C.H. is mainly Broadleafed Stringybark
(E. caliginosa Blakely and McKie) and Ribbon Gum (E. viminalis Labill.). Along

the creek Water Gum (Tristania laurina R. Br.) forms a dense association.

Grasslands in the region range from the “‘anomolous grassland communities
associated with rainforest on basalt’? (Baur, 1962) to improved pasture with
white clover.

The central feature of the study area was the pine plantation. Pinus elliottii
Engel. and P. taeda L. were first planted in 1950-51. Planting was recommenced in
1965 mainly with P. taeda but other species, P. radiata D. Don. and P. patula
Schl et Cham. were also planted.

The principle climatic features are the cold dry winter and the wet summer
months. The lowest minimum temperature over the 25 years previous to 1972
was —11.1°C and the highest maximum temperature was 39.5°C. Rainfall is

Aust. Zool. 19(1), 1976 25

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

concentrated over the summer months; 55 per cent of the average annual rainfall
of 1443 mm falls between November and March.

LIVE TRAPPING

Live trapping for small mammals was with Elliott folding aluminium traps
(Elliotts Scientific Co., Upwey, Victoria) baited with a mixture of peanut butter
and rolled oats. Medium sized mammals were trapped with three sizes of open-
mesh wire traps. Two sizes of non-collapsible traps (Mascot Wire Works, Enfield,
N.S.W.) 61 x 30 x 30 cm and 76 x 30 x 30 cm, and one size of collapsible trap
(Tomahawk Live Trap Co., Tomahawk, Wisconsin, U.S.A.) 66 x 22 x 22 cm
were used. These were usually baited with apple but on occasions peanut butter
and rolled oats, tinned dog food, bread, and bread and honey were used.

There were 30,500 trap nights with the open-mesh traps of which 500 trap
nights were on survey work, the remainder being in fixed locations. There were
10,621 trap nights with small mammal traps in fixed locations and 2,624 trap
nights on survey work. The majority of the survey work where traps were either
moved daily or every two days to a new location was generally on road edges
separating pine and native vegetation.

Small mammals were individually marked by toe clipping and possums by
ear nicks.

BREAK-BACK TRAPPING

Certain pine areas used as fixed locations for small mammal traps gave <1%
trap success. In February 1976 a trap success of 7.59% was obtained in these
areas from 120 trap nights with break-back rat traps baited with peanut butter
and rolled oats. The only species caught in this way were Rattus rattus and one
juvenile R. fuscipes.

SPOTLIGHTING
There was a total of 38 spotlight hours both on foot and from a moving

vehicle. Spotlighting from a vehicle was in general along the periphery of the
pine plantation and when on foot was generally along the roads associated with

Clouds Creek.

ROADKILLS

During 1975/76 regular journeys were made along the road between Clouds
Creek and Billys Creek, a distance of 13 km. All roadkills were collected,
decapitated and skulls subsequently prepared. Fifteen animals were collected in
this way. Roadkill specimens and animals that died in traps are located at La
Trobe University.

DAYLIGHT SIGHTINGS

During the time spent in the study area any mammals observed during the
day were identified and recorded.

26 Aust. Zool. 19(1), 1976

MAMMALS OF CLOUDS CREEK

TABLE 1

SYSTEMATIC LIST OF MAMMALS OF THE CLOUDS CREEK AREA INDICATING
THEIR OBSERVED ABUNDANCE IN EIGHT HABITATS

1 — closed forest; 2 — tall open forest; 3 — open forest; 4 —- woodland; 5 — central pines:

6 — peripheral pine; 7 — grassland; 8 — creek association

abundant (A) >50; common (c) 10-50; uncommon (U) 5-10; scarce (S) < 5S animals

observed

er ———

Common name

Scientific name

Family Tachyglossidae
1. Echidna

Family Ornithorhynchidae
2. Platypus

Family Dasyuridae

3. Brown phascogale

4. Tiger-cat

Family Peramelidae

_5. Short-nosed bandicoot
6. Long-nosed bandicoot
Family Phalangeridae

7. Brush-tailed possum
8. Mountain possum
Family Petauridae

9. Ringtail possum

10. Sugar glider

11. Yellow-bellied glider
12. Greater glider

Family Burramyidae

13. Feathertail glider
Family Phascolarctidae
14. Koala

Family Macropodidae

15. Grey kangaroo

16. Wallaroo

17. Red-necked wallaby
18. Parma wallaby

19. Swamp wallaby

20. Red-necked pademelon
21. Red-legged pademelon
22. Rufous rat-kangaroo
23. Potoroo

Family Vespertilionidae
24. Bent-winged bat

Family Muridae

25. Bush rat

26. Swamp rat

27. Black rat

28. Mosaic-tailed rat
29. House mouse

Aust. Zool. 19(1), 1976

Order Monotremata
Tachyglossus aculeatus (Shaw)

Ornithorhynchus anatinus (Shaw)

Order Marsupialia

Antechinus stuartii Macleay
Dasyurus maculatus (Kerr)

Jsoodon macrourus (Gould)
Perameles nasuta Geoffroy

Trichosurus vulpecula (Kerr)
Trichosurus caninus (Ogilby)

Pseudocheirus peregrinus (Boddaert)

Petaurus breviceps Waterhouse
Petaurus australis Shaw
Schoinobates volans (Kerr)

Acrobates pygmaeus (Shaw)
Phascolarctos cinereus (Goldfuss)

Macropus giganteus Shaw
Macropus robustus Gould
Macropus rufogriseus (Desmarest )
Macropus parma Waterhouse
Wallabia bicolor (Desmarest)
Thylogale thetis (Lesson)
Thylogale stigmatica Gould
Aepyprymnus rufescens (Gray)
Potorous tridactylus (Kerr)

Order Chiroptera

Miniopterus schreibersii (Kuhl)

Order Rodentia

Rattus fuscipes (Waterhouse)
Rattus lutreolus (Gray)
Rattus rattus (L.)

Melomys cervinipes (Gould)
Mus musculus L.

I

Ceetee

~~

Habitat
Ds CRU TA SS Dn
A Sees
Ss s
Ss Cc U
S S
A c
A U re
Ss
S
Ss
C U
S
Ss
U Cc
Ss
oma ene
U
U Sey 0)
Cc Cc
Ss
Ss
? Ss ?
aerial
c em &
U
Diete
Cc
Cc

ano

8

D

U

27

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

Common name Scientific name Habitat

Order Lagomorpha DE Die Bh. An SG eee
Family Leporidae
30. Rabbit Orvctolagus cuniculus (L.) 5+ -GaNs

Order Carnivora
Family Canidae

31. Dog/dingo Canis familiaris L. GOS) Sov SASHES Saas
32. Fox Vulpes vulpes (L.) s

Family Felidae

33.. Feral cat Felis catus L, C'S "SS 'siits| fsa

Order Artiodactyla
Family Bovidae
34. Cattle Bos taurus CC. A Ac AU AS JAeaen

RESULTS

A systematic list of the species of mammals observed in the study area is
given in Table 1; Rides’ (1970) classification is used. Twenty-seven indigenous
and seven introduced species were recorded.

1. Echidna, Tachyglossus aculeatus

During the entire study period only one echidna was seen; this was in the
grassland area near the Forest Commission Headquarters (F.C.H.).

2. Platypus, Ornithorhynchus anatinus

Platypus were sighted on three occasions in the same area although a
considerable amount of time was spent along creeks at all times of the day and
during all seasons. The sightings were in Clouds Creek near the F.C.H. in a region.
where the water was moving slowly and there were large pools. Both the
monotremes must be considered uncommon.

3. Brown phascogale, Antechinus stuartii

_ A total of 131 individual A. stwartiz were trapped, mainly in 1975/76. They
were found in closed forest and tall open forest near the F.C.H. and to the east
of, Chapmans Plain Road in open forest, and in the pine plantation itself. Of the
nine animals trapped in pine compartments, only two, both males, were trapped
in the central plantation (in August at the time of mating); the others were
caught in areas adjacent to native forest.

4. Tiger-cat, Dasyurus maculatus

Three tiger-cats were trapped during winter months, all in different vegetation
types. One was in pines and another in closed forest near the F.C.H., and the
other in tall open forest to the east of Chapmans Plain Road.

28 Aust. Zool. 19(1), 1976

MAMMALS OF CLOUDS CREEK

5. Short-nosed bandicoot, Isoodon macrourus

Seventy-seven short-nosed bandicoots were trapped in the grazed woodland
adjacent to the Armidale-Grafton Road, in the grassland, tall open forest, open
forest and pines near the F.C.H. One roadkill was collected at Billys Creek and
others were spotlighted in pines, in grassland around the F.C.H. and at the airport.
No short-nosed bandicoots were trapped during November, December and January;
42 of the specimens were trapped between May and August.

6. Long-nosed bandicoot, Perameles nasuta

The long-nosed bandicoot was considerably less common than I. macrourus.
Four specimens were trapped in grassland, tall open forest and grazed woodland,
all in the vicinity of the F.C.H.

7. Brush-tailed possum, Trichosurus vulpecula

This was one of the target species of the ecological study. There was a total
of 593 captures representing 98 individuals. None were trapped in tall open forest
or closed forest and they occurred less frequently in pines than in grazed woodland,
open forest and grassland.

8. Mountain possum, Trichosurus caninus

This species was the other major possum species. There was a total of 2,597
captures representing 193 individuals. They occurred mainly in tall open forest
and closed forest. They were also found in the other vegetation types although
few occurred in grazed woodland. They were widely distributed over the whole
of the study area, and all individuals were of the grey colour phase.

9. Ringtail possum, Pseudocheirus peregrinus

This species was considered rare in the area. One specimen was trapped in
tall open forest near the F.C.H. and two were spotlighted along the creek in
the same area.

10. Sugar glider, Petaurus breviceps

Three sugar gliders were observed during the day emerging from nest holes
in open and tall open forest near the F.C.H. They appeared to have been forced
out by T. caninus and T. vulpecula which climbed the trees after being released
from traps.

11. Yellow-bellied glider, Petaurus australis

This species was not sighted but its characteristic calls were heard several
times in trees along the banks of the creek and in tall open forest near the F.C.H.

12. Greater glider, Schoinobates volans

This species was widely distributed throughout the study area; it was not
observed in pines or grassland.

Aust. Zool. 19(1), 1976 29

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

13. Feathertail glider, Acrobates pygmaeus

Only one dried skin of this species was found in tall open forest near

the F.C.H.

14. Koala, Phascolarctos cinereus

One koala was found dead after a “control burn” in grazed woodland
approximately 2 km to the west of the Armidale-Grafton Road.

15. Grey kangaroo, Macropus giganteus

This species was common and widely distributed in the study area and was
sighted in grassland, grazed woodland, open forest and pines.

16. Wallaroo, Macropus robustus

This species is considerably less common than the grey kangaroo. One skull
was collected opposite a transition of tall open and open forest approximately
5 km south of Clouds Creek. One live specimen was observed in a similar forest
type near the F.C.H.

17. Red-necked wallaby, Macropus rufogriseus

This species was found extensively throughout the pines and in areas of
grazed woodland and grassland.

18. Parma wallaby, Macropus parma

Four roadkills of this species were collected, one near closed forest and two
near tall open forest ca. 3 km to the south towards Billys Creek, and one at
Muck Creek. This species is probably quite common throughout the area.

19. Swamp wallaby, Wallabia bicolor

Two specimens were trapped in pine and along the creek in tall open forest
near the F.C.H. This species was observed mainly during the day in closed forest,
tall open and open forest.

20. Red-necked pademelon, Thylogale thetis

This small macropod was common throughout the whole study area and was
sighted mainly on roads between the pines and native forest. Two roadkills were
collected near grazed woodland.

21. Red-legged pademelon, Thylogale stigmatica

This pademelon was less common than T. thetis and was observed only
along roads near closed forest.

22. Rufous rat-kangaroo, Aepyprymnus rufescens

Only one specimen was collected as a roadkill near tall open forest 10 km
towards Billys Creek.

30 Aust. Zool. 19(1), 1976

MAMMALS OF CLOUDS CREEK

23. Potoroo, Potorous tridactylus

Only one specimen was trapped in grazed woodland near the F.C.H.
However, spotlight sightings gave only a glimpse of many small macropods
throughout the study area which were probably potoroos; this species is probably
more common than indicated by the one capture and lack of positive sightings.

24. Bent-winged bat, Miniopterus schreibersii

The only bat collected was one specimen of this species which was brought
in by a cat. Numerous bats were observed in the area but these remain unidentified.

25. Bush rat, Rattus fuscipes

Eighty-five individual R. fuscipes were trapped in closed forest, tall open
forest, open forest and on the periphery of the pines. They were scattered
throughout the study area, but were nowhere in high numbers; only one individual
was caught in the central pine area. The nipple formula 1 + 3 = 8 and 2 + 3 = 10
occurred in the ratio of 3:1.

26. Swamp rat, Rattus lutreolus

Nine individuals were trapped in February and May 1975. They were all
trapped within 35 m of the edge of the eastern pine compartments.

27. Black rat, Rattus rattus

A total of 16 individuals were trapped in both central and peripheral pines
with live and break-back traps. None were trapped in native forest. Two colour
phases, the black-backed and the brown-backed, occurred in the ratio of 7:1.

28. Mosaic-tailed rat, Melomys cervinipes

Twenty-six individuals were trapped in closed forest and tall open forest
to the east of Chapmans Plain Road. Of these only two were trapped along the
edge of a road.

29. House mouse, Mus musculus

Only eight individual mice were trapped, all in peripheral pines near the
F.C.H. and in the eastern pines. None were trapped in native forest. Others have
been brought in by cats and observed in grassland near the F.C.H.

30. Rabbit, Oryctolagus cuniculus

No rabbits were seen during 1975/76 and they must now be considered
rare or absent from the area. However, in 1971 they were present in the one
to two-year-old pine compartments.

31. Dog/dingo, Canis familiaris
One dog skeleton was found in grassland at Chapman Plain 11 km south-east
of F.C.H., and numerous pads and faeces were observed along the pine periphery.

Aust. Zool. 19(1), 1976 31

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

Three known individuals have been seen together in open forest near the airport,
along Sheep Station Creek Road, and along a road through the pines near the
F.C.H. Dogs probably roam throughout the area.

32. Fox, Vulpes vulpes
Only one roadkill fox was collected 6 km along the Armidale-Grafton Road
towards Billys Creek; this species must be considered rare.

33. Feral cat, Felis catus

Feral cats are common and widely distributed throughout the study area
except for no sightings in closed forest. Specimens have been trapped in both
tall open forest and in pines.

34. Cattle, Bos taurus

This species is included as cattle are allowed to graze in the State Forest.
They are wide-ranging and occur throughout the whole study area in all
vegetation types.

DISCUSSION

The mammal fauna of Clouds Creek State Forest is very diverse. There are
27 species of native mammal and seven introduced species; this list will
undoubtedly be considerably extended when more bats and some of the small
macropods are collected and identified. The faunal diversity compares favourably
with that of the area drained by the upper Richmond and Clarence Rivers (100 km
to the north) where 45 native species (12 of which were bats) and seven
introduced species were recorded (Calaby, 1966) in an area of 2,400 km? (cf.
118 km? in the Clouds Creek study area) and a smaller survey in Moonpar State
Forest 15 km to the south where 17 species were recorded (Maynes, 1974).

The small mammals, however, were not particularly abundant; trap success
with live small mammal traps averaged 4.4% ranging from 0.4% in the central
pines to 8% in native forest to the east of Chapmans Plain Road. This latter area
held three species, A. stuartii, R. fuscipes and M. cervinipes, the two rodents
being present in relatively low numbers.

The distribution of mammals throughout the area showed that only R. rattus,
R. lutreolus and M. musculus were trapped only in pines. This appears dependent
on the age of the pines; in the older pines R. rattus and M. musculus were very
strongly associated with the periphery. Although R. lutreolus were trapped in
pines, this species was a transient in the study area and all captures were again very
strongly associated with the periphery. Little is known of the biology of this
species, but Braithwaite (pers. comm.) believes their distribution is closely
correlated with the penetrability of the soil in which they burrow. High summer
and early autumn rainfall could explain the patchy temporal distribution of this
species.

32 Aust. Zool. 19(1), 1976

MAMMALS OF CLOUDS CREEK

Individual R. fuscipes and M. rufogriseus and possibly some M. giganteus and
I. macrourus also occurred in pines and associated grasslands, but other individuals
of these species occurred solely in other vegetation types. Although a few (nine
out of 131) A. stwartii were trapped in pines, none were recaptured. Two transient
male A. stwartii were trapped in the central pine area during the mating period,
but females appeared absent from this area. Other species were observed in pines
but they appeared dependent on nearby native forest areas; these included D.
maculatus, Trichosurus caninus, T. vulpecula and Thylogale thetis.

The species found solely in association with native forest were S. volans,
Melomys cervinipes, Thylogale stigmatica and probably Perameles nasuta, Pseudo-
cheirus peregrinus, Petaurus breviceps, P. australis, Acrobates pygmaeus, Phasco-
larctos cinereus, Macropus robustus, M. parma, M. dorsalis and Aepyprymnus
rufescens; none of these species were cbserved in anything other than native forest
although several occurred in grazed woodland.

None of the forest in the Clouds Creek area has been left untouched by
man. Disturbance varies from selective logging in closed forest areas to complete
clearfelling of native forest and subsequent planting with Pinus spp. The
remaining closed and tall open forests are probably the least disturbed. Grazing
obviously alters the vegetation association and clearing and replacing with pines
is a serious perturbation,

The replacement of native forest by pines has a range of effects on the fauna
depending upon the age of pines and the distance from native vegetation. Melomys
cervinipes, Schoinobates volans and Thylogale stigmatica are always lost, while
Trichosurus caninus and T. vulpecula utilize the pine margins but do not live
solely in pines (How, 1972). In the central pines no native small mammals occur
and are replaced by R. rattus and Mus musculus.

ACKNOWLEDGEMENTS
This study was supported by grants from the New South Wales Forestry Commission
(1967-72) and the Federal Department of the Environment and Conservation (1975-76).

We wish to thank numerous colleagues who have accompanied and assisted us on our
numerous field trips, and helped in identification of material; Jenny Barnett, Gill Humphreys,
Leigh Ahern, John Blyth, Dave Middleton, Ken Johnson and John Brereton.

REFERENCES

BAUR, G. N. (1962). Forest vegetation in north-eastern N.S.W. For. Comm. N.S.W. Res.
Note No. 8, 1-30. }

CALABY, J. H. (1966). Mammals of the upper Richmond and Clarence Rivers of N.S.W.
C.S.I.R.O. Wild. Res. Paper No. J0, 1-55.

HOW, R. A. (1972). The ecology and management of Trichosurus species (Marsupialia) in
N.S.W. Ph.D. thesis, Department of Zoology, University of New England.

MAYNES, G. M. (1974). Occurrence and field recognition of Macropus parma, Aust, Zool.
18, 72-87.

Aust. Zool. 19(1), 1976 33

34

J. L. BARNETT, R. A. HOW and W. F. HUMPHREYS

RIDE, W. D. L. (1970). A Guide to the Native Mammals of Australia. Oxford University
Press: London.

SPECHT, R. L. (1970). Vegetation, In “The Australian Environment”. 4th Ed. (Ed. G. W.
Leeper), pp. 44-67. (CSIRO-Melbourne University Press: Melbourne. )

TYNDALE-BISCOE, C. H. &. J. H. CALABY (1975). Eucalypt: forests. as refuge ‘for
Wildlife. Aust. For. 38, 117-133.

ADDENDUM

Continued survey of the area since submission of the manuscript has resulted in the
following additions. An individual male of each of Mus musculus and Pseudocheirus

peregrinus was trapped in closed forest and one male Rattus lutreolus was trapped in creek-
associated grassland.

Aust. Zool. 19(1), 1976

Aggressiveness, Body Weight and Injuries in
Long-haired Rats (Rattus villosissimus)

ROBERT J. BEGG
Department of Zoology, Monash University, Clayton, Victoria, 3168*.

*Present address: Wildlife and National Parks Section, Department of the Northern
Territory, Darwin, Northern Territory, Australia, 5790.

ABSTRACT

This paper details methods used to objectively estimate the aggressiveness of
male Rattus villosissimus. The principal technique involves ranking rats in a
series according to their incidence of dominance and subordination in encounters.
A second measure, attack latency, is used to test the established rankings. A
very high correlation of body weight and the incidence of wounding is examined
to show the usefulness of the technique.

INTRODUCTION

During a study of the visual, tactile and auditory signals involved in the
agonistic behaviour of Rattus villosissimus (Begg and Nelson, in press), it was
evident that individual rats differed greatly in aggressiveness, defined here as the
tendency to attack conspecifics.

A prelude to any investigation of the causal factors for individual differences
in aggressiveness, or a correlation of this variable with other factors (e.g. repro-
ductive success), is a reliable technique for the measurement of aggressiveness.

This task has been attempted in several ways, involving the estimation of
one or more of the variables listed below:

(i) the frequency of occurrence of selected behaviour patterns;
(ii) attack latency;
(iii) total duration of attack;
(iv) the general outcome of encounters (i.e. ““win” or “loss’’);
(v) rating scales of aggressiveness (e.g. Bevan ef al., 1958).
Application of the last method has not produced many useful results, “due

to the arbitrary ranking of behaviours that are not highly correlated or not
correlated at all, on a linear scale’ (Scott, 1966).

To be reliable, any method chosen to estimate individual aggressiveness,
should give the same result for each animal tested. Working with Mus musculus,
Catlett (1961), found measures (i), (ii), or (iii), above, to give consistent

Aust. Zool. 19(1), 1976 35

ROBERT J. BEGG

results for the same animal on successive days. However, he was only able to
obtain a correlation of 0.66 between measures.

There is usually a clear-cut dichotomy in the behaviour of two adult R.
villosissimus engaged in an encounter. After an initial period of intense activity
(usually very brief), one rat adopts a “subordinate” role and usually crouches
alone, when undisturbed. Subsequently the opponent initiates all attacks and is
termed “dominant”. The subordinate rat usually attempts to defend itself or flee,
when approached by a dominant opponent. This behaviour is described fully by
Begg and Nelson (in press).

The terms ‘dominant’ and ‘subordinate’, as used in this context, do not
imply any social values found under natural conditions. They are simply used to
describe the different kinds of behaviour that may be observed in two rats involved
in a conflict.

These terms follow Scott’s (1956) definition of a ‘‘dominance-subordination
relationship”, as “‘antagonistic behaviour, where the behaviour of one individual
consists of threat, or actual fighting, while the other individual remains passive
or attempts to escape’”’.

MATERIALS AND METHODS

All rats used were trapped in the wild as adult males and kept as described
previously (Begg, 1975). Encounters were run between pairs of rats in a pine-
board cage measuring 90 x 55 x 45 cm, with a clear Perspex front. A thin metal
slide inserted through a longitudinal slot in the Perspex divided the cage floor
into two equal areas. The cage floor was covered with wood shavings which
were changed after each encounter to minimise the effects of strange odours.

For each encounter one rat was placed in each section of the cage and
allowed five minutes to settle down. Most rats spent this time exploring the area
in which they were placed. The slide was then removed and the rats were
observed for 15 minutes.

The time between the removal of the partition and the initiation of the first
aggressive act by one rat was measured and termed the attack latency for that
encounter. If no aggression occurred during the 15-minute period the encounter
was termed neutral.

Each rat was scored as dominant or subordinate for each encounter in which
it participated. At the end of each encounter each rat was weighed to the nearest
gram and all visible injuries were recorded. This included: the number of cuts
per rat; the areas in which cuts were located; and a subjective estimate of the
severity of each wound.

One hundred and five encounters were run between pairs of rats, randomly
chosen from a stock of 19 animals. No rat was used more than once per day.

36 Aust. Zool. 19(1), 1976

AGGRESSIVENESS IN RATS

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ROBERT J. BEGG

RESULTS
CALCULATION OF AGGRESSIVE RANKING

A ratio of the number of encounters in which the rat was dominant to the
number in which it was subordinate, was calculated for each rat.

The more aggressive a rat is the higher the ratio may be expected to be.
Thus, by comparing the ratios for each rat in a group it was possible to establish
a ranking from the most aggressive rat (listed as number one) to the least
aggressive rat in the group (Table 1). If any pair of rats had the same ratio the
dominant rat in an encounter between the two was given the higher ranking.

All that this ranking is meant to imply is the tendency for a rat to be
dominant under the conditions of the study. It does not imply that a linear
dominance hierarchy (with associated ordering of access to food, water, mates,
etc.) would exist if the animals were grouped. In fact Barnett (1958) showed
that, at least for R. norvegicus, such a system does not exist.

STABILITY OF AGGRESSIVE RANKING

To test the stability of rankings, 10 encounters between pairs of rats,
chosen from a table of random numbers, were repeated four months after the
initial series of trials.

Of the 10 encounters repeated, eight resulted in the same rat being dominant
on both occasions. The other two encounters resulted, on the second occasion, in

LATENCY

ATTACK

logio

Pt 2 8it i Serer 7 Ory 10st ih 12218 ARMS 46

AGGRESSIVE RANK

Fig. 1. logi attack latency vs aggressive rank.

38 Aust. Zool. 19(1), 1976

AGGRESSIVENESS IN RATS

some aggression between pairs of rats that had previously had neutral encounters.
In both cases, however, the dominant rat on the second occasion was of a higher
ageressive rank than its opponent. Thus, although certainly not conclusive, these
results tend to indicate that the relationships between rats are fairly stable over
long periods.

ATTACK LATENCY

A mean attack latency was calculated for each rat over all encounters in
which the rat initiated the first attack (Table 1).

To test the correlation between aggressive ranking and attack latency as
estimates of the aggressiveness of individuals, the logarithm to base 10 of the mean
attack latency of each rat was plotted against the established aggressive rank of that
rat (Fig. 1). Logarithmic values were used for the ordinate as the relationship was
found to be non-linear. A linear regression analysis was used to test the fit between
these two variables. The regression line (y = ao + aix) is plotted in Figure 1
(ao = 0.104, a1 = 0.785, with r = 0.90, df. = 14, and P = 0.01).

AGGRESSIVE RANK AND BODY WEIGHT

To test if body weight is a factor in aggressiveness, the mean body weight of
each rat over the total period of observations was computed (Table 1). The mean

WEIGHT (gms)

BODY

MEAN

ye ener SOOO Ie Note ate eS. 16 Phe
b

AGGRESSIVE RANK
Fig. 2. Mean body weight vs aggressive rank.

Aust. Zool. 19(1), 1976 39

ROBERT J. BEGG

body. weight of each rat involved was then plotted against the aggressive rank
of that animal (Fig. 2) and a linear regression analysis carried out. Taking rankings
one to 19 in male-male encounters resulted in a correlation coeflicient of —0.226,

d.f. = 17, P > 0.05. However, a linear regression analysis on rankings one to 15
produced a correlation coefficient of —0.764. This value is significant at a 0.1%
level (d.f. = 13). This regression line (a) = 292.3, a1 = —6.079) is plotted in
Figure 2.

- Thus, for rats with aggressive rankings from one to 15, there is a highly
significant correlation between aggressive rank and body weight. For animals with
a lower ranking, the relationship is lost.

AGGRESSIVE RANK AND THE INCIDENCE OF WOUNDING

In 105 male-male encounters, 79 cuts resulted. All but one of these were to
subordinate rats. The numbers of cuts delivered or received by rats were plotted
against the aggressive rank of each rat (Figs. 3, 4). For cuts delivered to opponents

a9 — 11.64, a =-— 0°78; with r = --0:583, df= 15, P = 0.02? Thus the
number of wounds caused by an attacker was significantly correlated with the
aggressive rank of the attacking rat. For cuts received, however, ao = 3.57,

DELIVERED

NUMBER OF CUTS

fe a teehee eo 1g tm Io a ee eG, 47

AGGRESSIVE RANK

Fig. 3. Number of cuts delivered by rats vs aggressive rank.

_ 40 Aust. Zool. 19(1), 1976

AGGRESSIVENESS IN RATS

RECEIVED

OF CUTS

NUMBER

OVC Stas sorte Lot Oe OM. TR AD Abs ISAS iE

AGGRESSIVE RANK

Fig. 4. Number of cuts received by rats vs aggressive rank.

a1 = 0.05, with r = 0.055, df. = 15, P > 0.05. Thus the number of wounds
received by a rat in encounters was not significantly correlated with the aggressive
rank of the injured rat.

DISCUSSION

Ranking rats according to their incidence of dominance and subordination in
encounters proved to be an effective method of representing the aggressiveness of
male R. villosissimus. When compared with attack latency, a more commonly
used measure, both methods gave the same estimate of the aggressiveness of
individual rats. Previously Catlett (1961) was only able to obtain a correlation
of 0.66 between different techniques for estimating the aggressiveness of Mus
musculus.

It must again be stressed that the estimates of aggressiveness discussed in
this study were obtained under specific conditions of cage size and construction,
and experimentation, and thereby do not necessarily hold for a field situation.
Aggression is probably heightened under these conditions of study, as caged
animals have no avenue of escape, as discussed by Begg and Nelson (in press).

A series of encounters repeated four months after the original series of trials
indicated that the rankings assigned to rats apply over a long period of time, as

Aust. Zool. 19(1), 1976 41

ROBERT J. BEGG

almost all encounters gave the same results on both occasions. Lerwill and Makings
(1971). found that for golden hamsters (Mesocricetus auratus) “relationship
tends to be the same for any particular pair of hamsters, whenever they meet
in similar circumstances”. These facts tend to indicate that the establishment of
a dominance-subordination relationship between rats probably does not depend on
learning between two animals, as in the first series of encounters each pair only
met once. More likely, some physical characteristics such as body weight, which
was shown to be correlated with aggressiveness, are involved.

There was found to be a highly significant correlation between mean body
weight and positions one to 15 on the aggressive ranking of male rats. Thus
heavier rats tend to be dominant in more encounters. Barnett (1958) found that
for colonies of R. norvegicus “most of the rats which grew well were dominant
over the other males’, and ‘“‘alpha males were always large by comparison with
other members of the colony”. Similarly, Calhoun (1962) found that for R.
norvegicus “for their respective ages, winners have greater than mean weights, and
losers less than mean weights”’.

Thus, positions 16 to 19 on the aggressive ranking represent heavy rats that
for some reason are subordinate in more encounters than may be expected. It is
suggested that these rats are older (and thus heavier) than the other rats in the
series and are not able to assert dominance over younger, stronger rats, and are
thus subordinate in more encounters. Ewer (1971) found that both male and
female wild R. rattus become less belligerent and show a general decline in
activity as they become old. As Ewer says, males “‘become heavy, less agile, and
a top ranking male is distinctly Jazy”. Newsome (1969) also suggested that with
wild Mus musculus living in reed beds, old mice may be unable to maintain social
superiority.

It was not possible to age the rats used in this study accurately as, firstly, they
were needed for further experiments; secondly, one of the major methods used for
aging R. villosissimus by Carstairs (1971) was to weigh the eye lenses of the
animals. The curve for the relationship age/eye lens weight flattens out between
the ages of 300-400 days and is thus inaccurate at the upper limits.

The number of injuries caused by a rat was found to be significantly correlated
with its aggressive rank. That is, more aggressive rats deliver more wounds to
opponents. While this may appear to be a truism, the converse did not hold.
That is, the number of cuts received by a rat was not significantly correlated with
its aggressive rank. Calhoun (1962) in his enclosure study of wild R. norvegicus,
found more cuts and scars on animals of lower social rank. This measure is thus
often used as an index of social status in the field. In R. villosissimus, however,
this correlation probably does not hold. In this case animals toward the centre
of the established aggressive ranking tended to receive the most injuries in
encounters. These individuals may be attacked more by rats higher in the aggressive
ranking, as they are closer to the latter in terms of body weight and behaviour.

42 Aust. Zool. 19(1), 1976

AGGRESSIVENESS IN RATS

REFERENCES
BARNETT, S. A. (1958). An analvsis of social behaviour in wild rats. Proc. zool, Soc.
Lond., 130, 107-152.

BEGG, R. J. (1975). The agonistic vocalisations of Rattus villosissimus. Aust. J. Zool., 23,
597-614.

BEGG, R. J., & J. E. NELSON (in press). The agonistic behaviour of Rattus villosissimus.

BEVAN, J.M., W. BEVAN, & B. F. WILLIAMS (1958). Spontaneous aggressiveness in
young castrate C3H male mice treated with three dose levels of testosterone. Physiol.
Zool., 31, 284-288.

CALHOUN, J. B. (1962). The ecology and sociology of the Norway rat. U.S. Dept. Health.
Education and Welfare.

CARSTAIRS, J. L. (1971). Unpublished Ph.D. thesis.

CATLETT, R. H. (1961). An evaluation of methods for measuring fighting behaviour, with
special reference to Mus musculus. Anim. Behav., 9, 8-10.

EWER, R. F. (1971). The biology and behaviour of a free-living population of Black Rats.
Anim. Behav. Mon. 4, part 3, 127-174.

LERWILL, C. J... & P. MAKINGS (1971). The agonistic behaviour of the Golden Hamster
Mesocricetus auratus (Waterhouse). Anim. Behav. 19, 714-721.

NEWSOME, A. E. (1969). A population study of house-mice permanently inhabiting a
reed-bed in South Australia. J. Anim. Ecol. 38, 361-377.

SCOTT, J. P. (1956). The analysis of social organisation in animals. Ecology, 37, 213-221.

SCOTT, J. P. (1966). Agonistic behaviour of mice and rats: a review. Am. Zool., 6,
683-701.

Aust. Zool. 19(1), 1976 43

sae
TR

li Te

te oa a

More Fish Genera Scrutinized

The late GILBERT P. WHITLEY

Formerly Research Associate, The Australian Museum, College Street, Sydney,
New South Wales, 2000.

ABSTRACT

More than 16,000 generic or subgeneric names have been given to fishes. With
a view to eradicating preoccupied or otherwise invalid names, all of them have
been checked with nomenclators. Each doubtful genus is discussed below in
alphabetical order, the style following the author’s ‘‘Some fish genera scrutinized’’
which appeared in Proc. Roy. Zool. Soc. N.S. Wales, 1964-65, pp. 25-26.

INTRODUCTION

An overall modern classification of fishes in the broadest sense (to include
lancelets, lampreys, elasmobranchs, teleosts, etc., and all recent and extinct families )
down to generic level is still a desideratum.

More than 16,000 generic or subgeneric names (including nomina vana such
as synonyms, homonyms, emended or variant spellings, etc.) have been bestowed
on fishes since 1753 when Artedi listed the 50 genera known in his time. Thus
there are many more names than natural genera. One way of reducing the excess
is to eradicate preoccupied names; another is to dispose of synonyms. All of the
known names published between 1758 (Linnaeus) and the early 1970s have been
personally checked with nomenclators (for references see Whitley, Austr. Zool.,
13, 1966, p. 234). In this paper, certain generic names are purged.

The old name of each genus discussed is set out in alphabetical order below,
following the same style as my earlier paper, ‘““Some fish genera scrutinized”’ (Proc.
Roy. Zool. Soc. N.S.Wales, 1964-65, pp. 25-26).

Sometimes (e.g. Acropholis, Mesocyprinus, Palaeogadus) new generic names
were ascribed to more than one author by nomenclators, yet they were evidently
duplicated publications of the new names, the later not preoccupied by the earlier
ones. The nomina nuda of Agassiz, and Woodward’s old names, restored in
Casier’s papers (especially his Faune ichth. London Clay, 1966) are other cases
in point (examples: Orvikuina, Percostoma, Podocephalus and Rhinocephalus).
These need not be considered further.

Aust. Zool. 19(1), 1976 ; 45

MORE FISH GENERA SCRUTINIZED

REMARKS
Ankistrorhynchus (Family Pristidae):

Ankistrorhynchus Casier (Bull. Inst. r. Sci. nat. Belg., 40(11), 1964, pp.
13-17) should be invalidated by Ancistrorhyncha Ulrich & Cooper (J. Paleont.,
16, 1942, p. 624) in Brachiopoda and preoccupied by Anxcistrorhynchus L’Hardy
(Cah. Biol. Mar., 4, 1963, p. 220), a genus of turbellarian worms.

Ateleopus (Fam. Ateleopidae):

Ateleopus Temminck & Schlegel (Fauna Japonica, Pisces, 1846, p. 255, pl.
112, fig. 1) was a genus caelebs, that is a new genus without a specifically named
type-species. It was published in May 1846 according to Mees (Zool. Verhand.,
54, 1962, p. 79) and validly so. In the same year an amphibian generic name was
emended to Ateleopus by Agassiz (Nomencl. Zool., 1846, Index Univer., p. 39).
Whose name was earlier? The Index Universalis was prefaced December 1845,
with title-page dated 1846. Bowley & H. M. Smith (J. Soc. Bibliogr. Nat. Hist.,
5 (1), 1968, p. 35) do not specify a month in 1846, so we must give the fish
name the benefit of the doubt and regard Agassiz’s amphibian name as preoccupied.
Furthermore, current usage will not be upset.

Temminck & Schlegel’s unnamed species was called Azeleopus schlegelii by
J. van der Hoeven (Handboek Dierkunde, 2, 1855, p. 326; Handbook of Zoology,
1858, p. 125). I have not seen the early editions of van der Hoeven’s Handbook,
but Sherborn (Index Animalium (2) 1, 1922, p. Ixviii of the bibliography) lists
them and they are mentioned in Opinion 487 of the International Commission on
Zoological Nomenclature. Some of van der Hoeven’s generic names were discussed
by Gill (Proc. U.S. Nat. Mus., 28, 1904, p. 119), Jordan (Genera of Fishes,
1919, p. 243) and Whitley (Rec. Austr. Mus., 16, 1928, pp. 294-295). Ateleopus
schlegeliit van der Hoeven, 1855 (if it were of earlier date, it would have been in
Sherborn’s Index) is however a synonym of Afeleopus japonicus Bleeker (Verh.
Bat. Gen., 25, 1853, p. 21) likeswise based on Temminck & Schlegel’s type-species.

A genus related to Ateleopus is Giintherus Osério (Arg. Univ. Lisboa, 4,
1917, p. 117. Guentherus Walters’ (Gopeia; 1963 (3); p.° 456) 1s’ simply am
emended spelling, not a new genus.

Barombia (Fam. Cichlidae):

Barombia Trewavas (Bonn Zool. Beitr., 13, 1962, p. 184), according to Neave
(Nomencl. Zool.) is preoccupied by Puchi Rarsch, 1891, 4n Orthoptera and
by Jacoby, 1903, in Coleoptera.

Bertella (Fam. Oneirodidae ):

Bertella Pietsch (Copeia, 1973 (2), p. 193) is not preoccupied by Bertella
Paetel, 1875, because, according to Neave (Nomencl. Zool., 1, 1939, p. 422) the
latter was an error for Berthella de Blainville, 1842, in Mollusca.

46 Aust. Zool. 19(1), 1976

The late GILBERT P. WHITLEY

Brachypterus (Fam. Scorpaenidae, subfam. Pteroinae ):

Brachypterus Catala (Carnaval sous la mer, 1964, p. 95), with type-species
Brachyrus zebra (Quoy & Gaimard, 1825], was a substitute for Brachirus Swainson,
1839, of which it is a synonym. Neave’s Nomencl. Zool. shows that Brachypterus
is preoccupied by Kugelann, 1794, and by other authors in entomology and
ornithology.

Buritia (Fam. Serrasalmidae):

Buritia Brant, 1972 — Brant & Marzulo (Bol. Mus. Hist. Nat., U.F.M.G.
(Brasil), Zool., 17, p. 1, figs. 1-10) is preoccupied by a genus of Hemiptera,
Buritia Young (Univ. Kansas Sci. Bull., 35, 1952, p. 67).

Byssacanthus (indet. Ichthyodorulite):

Byssacanthus Salter (Mem. Geol. Soc., 1861, p. 244, without type-species )
is preoccupied by Agassiz (Poiss. F.V.G.R., 1845, p. 116) and is an unnamed
ichthyodorulite allied to Chimaeridae according to Fowler (Quart. J. Taiwan Mus.,
23, 1970, p. 152; Cat. World Fishes, 1 (13), p. 656), so Salter’s name may
be allowed to lapse as his genus appears to be unidentifiable.

Coridodax (Fam. Odaciidae):

Coridodax Gunther (Cat. Fish. Brit. Mus., 4, 1862, pp. 69 & 243) is a new
synonym of Coregonoides Richardson (Ann. Mag. Nat. Hist., 11, 1843, p. 426).
L. D. Ritchie, quoted by Doak (Fishes of the New Zealand Region, 1972, p. 97)
shows that the type-species of these two generic names are conspecific; as a
corollary the genera are synonymic and the type-species should be known as
Coregonoides pullus (Bloch & Schneider, 1801), comb. nov.

Cretaspis (Fam. Odontaspidae ):

Cretaspis M. Sokolov (Byull. mosk. Obshch. Ispyt. Prir. Geol., 40 (4),
1965, p. 132) is preoccupied by Cretaspis Joleaud (Ann. Mus. Marseille, 15,
1916, p. 43), a genus of Crustacea.

Doryaspis (Fam. Lyktaspididae):

Doryaspis White (Phil. Trans. Roy. Soc. Lond., (B), 225, 1935, p. 444)
was considered by Denison (J. Linn. Soc. Lond., 47, 1967, pp. 31 & 34) to be
preoccupied, evidently by Doryaspis Dejean (Cat. Coleopt., ed. 2, 1835, p. 301),
but that is a nomen nudum according to Neave’s Nomenclator Zoologicus. Halstead
(Biol. Reviews Cambridge, 48 (3) 1973, p. 326) places the fish genus in the
synonymy of Lyktaspis Heintz, 1968.

Endemichthys (Fam. Dictyopygidae):

_ _Endemichthys Forey & Gardiner (Palaeont. Africana, 15, 1973, p. 29, fig. 1)
is preoccupied by Endemichthys Hopkirk (Diss. Abstr., 29B, 1968, p. 414), a
genus of Cyprinid fishes.

Aust. Zool. 19(1), 1976 47

MORE FISH GENERA SCRUTINIZED

Galeaspis (Ordo Liuaspidinida, nov., Fam. Liuaspidinidae, nov. ):

Galeaspis Liu Yi-hai (Vert. Palasiat., 9 (2), 1965, p. 130), a genus of the
Class Cephalaspides and of a new family named Galeaspidae by Liu, was placed
in the then new Order Galeaspiformes. The generic name is preoccupied by
Galeaspis Ivshin (in Borukaev, Prepaleozoic & Lower Paleozoic N.E. centr.
Kazakhstan. Acad. Sci. Kazakhstan S.S.R. (Inst. Geol. Sci.), Moscow, 1955,
p. 276) a genus of trilobites, recorded in the Zoological Records for 1962 as well
as for 1957.

“Goniophorus”

“Goniophorus” Toombs, Miles & Patterson (Zool. Rec., 104 (15) for 1967,
Pisces, 1970, p. 118; and Ricketts, Zool. Rec., 104 (20) for 1967, index, p. 8) is
a mis-spelling of Goniporus Gross, 1967. It is preoccupied by Goniophorus
Agassiz (Mon. Ech., 1, 1838, p. 30) in Echinodermata.

Humbertia (Ordo Salmoniformes ):

Humbertia Patterson (Bull. Brit. Mus. (Nat. Hist.) Geol., 19 (5), 1970,
p. 207, plates & figs.) is preoccupied by Humbertia de Beaumont (Bull. Soc. vaud.
Sci. nat., 69, 1965, p. 142), a genus of fossil mammals.

Katoporus (Fam. Phlebolepididae ):

Katoporus Gross (Palaeontographica, (A) 127, 1967, pp. 5, 24, etc., pls. 2
& 3) is preoccupied by Katoporus John (Ann. Mus. Congo Belg. (Ser. 8) Sci.
Zool., 51, 1956, p. 384) in Insecta.

Logania (Fam. Phlebolepididae ):

Logania Gross. (Palaecontographica, (A) 12/7, 1967, pp. 5, 31, ete, pe oe
is preoccupied by Logania Distant (Rhopal. Malay., 1884, pp. 197 & 208), a
genus of Lepidoptera.

Nybelinea (Fam. Roachiidae, nov.):

Nybelinia Nielsen (Galathea Rept., 10, 1969, p. 12 etc.) was preoccupied
by Nybelinia Poche (Arch. Naturgesch., 91, 1925, A. 2-3, 1926, Syst. Platodaria,
p. 364) a genus of Cestode worms, and has been renamed Nybelinella in the
family Aphyonidae by Nielsen elsewhere.

Palaeotroctes (Fam. Alepocephalidae):

Palaeotroctes Daniltshenko (Akad. CCCP, 78, 1960, p. 15, pl. 4, fig. 3) is
preoccupied by Palaeotroctes Enderlein (Palaeontographica, 58, 1911, p. 350), a
genus of Insecta (Psocoptera).

Priacanthopsis (Fam. Serranidae):

Priacanthopsis Arambourg (Notes Mém. Moyen-Orient, 8, 1967, p. 103) is
preoccupied by another genus of fishes, Priacanthopsis Fowler (Proc. Acad. Nat.
act? Philad 98:4 906, 1 122)5

48 Aust. Zool. 19(1), 1976

The late GILBERT P. WHITLEY

Pseudorhegma (Fam. Grammistidae ):

Pseudorhegma Schultz (Ichthyologica, April 1966, p. 185).

Not Pseudoregma Doncaster (Entomologist 99, 1966, p. 158), a genus of
Insecta (Aphids).

Quebecaspis ( Arthrodira: Dolichothoraci ) :

Quebecaspis Pageau (Naturaliste canad., 96, 1969, p. 832) is preoccupied
by Quebecaspis Rasetti (J. Paleont., 18, 1944, p. 254), a genus of Trilobita.

Siberiaspis (Subclass Heterostraci ):
Siberiaspis Obruchev (Osn. pal., 11, 1964, p. 68) is preoccupied by Sibiriaspis
Repina (Trudy SN 11 GGIMS, 19, 1960, p. 252) a genus of Trilobita.

Tangia (Fam. Lutjanidae):

Tangia Chan (Hong Kong Fisher. Res. Bull. 1 (F.R.S. Contrib. 17, 1970,
p. 19) is preoccupied by Tangia Stal (Berlin ent. Z., 3, 1859, p. 317), a genus
of Insecta (Hemiptera).

Thysia (Fam. Cichlidae):

Thysia Loiselle & Welcomme (Rev. Zool. Bot. Afr. 85, 1972, pp. 37-58) is
preoccupied by Thysia Thomson (Essai Class. Cerambyc., 1860, p. 96), a genus
of Coleoptera.

Trewavasia (Fam. Cichlidae):

Trewavasia Thys van den Audenaerde, 1967-71 (fide Trewavas, Bull. Brit.
Mus. Nat. Hist. (Zool.), 25 (1), 1973, p. 15 & footnote) is preoccupied by
another genus of fishes in fossil Siluridae (White & Moy-Thomas, Ann. Mag. Nat.
Hist. (11) 7, 1941, p. 400). A new name for the type-species, Tilapia guinasana
Trewavas, is evidently being supplied elsewhere.

Trigonocephalus (Fam. Gobiidae, subfam. Tridentigerinae):

Trigonocephalus Okada (Studies on the freshwater fishes of Japan, 1961,
p. 265) is preoccupied by Trigonocephalus Oppel (Ann. Mus. Hist. Nat. (Paris),
16 (95), 1811, pp. 377 & 388), a genus of reptiles. Okada’s name was a lapsus
calami for, and therefore a synonym of, Tridentiger Gill (Ann. Lyc. Nat. Hist.
New York, 7, 1858, p. 16) according to Hubbs (Copeia, 1962 (1), p. 238).

Ulapiscis (Fam. Lutjanidae):

Ulapiscis Whitley (Rec. Austr. Mus., 19, 1933, p. 78) is a hitherto unreported
synonym of Aphareus Cuvier & Valenciennes (Hist. Nat. Poiss., 6, 1830, p. 485).
The type-species, Ulapiscis kennedyi Whitley from the Ellice Islands, agrees with
Aphareus furcatus (Lacepede, 1802) as described and figured by Schultz (US.
Nat: Mus-Bull=:202; 1953\ip. 539, pl. 48; fie B):

Aust. Zool. 19(1), 1976 4g

MORE FISH GENERA SCRUTINIZED

Yukonaspis (Heterostraci: Fam. Traquairaspididae ):
Yukonaspis Obruchev (Osnovy Paleont., 2, 1964, p. 63) is preoccupied by
Yukonaspis Kobayashi (J. Paleont., 10 (3), 1936, p. 164), a genus of Trilobita.

Zisius (Fam. Xiphiidae):
Zisius Oken (Lehrb. Naturg., 3, Zool. 2, 1816, pp. ii & 151) is a synonym
of Xiphias Linnaeus 1758, the swordfish.

ACKNOWLEDGEMENTS

_. The author acknowledges, with grateful thanks, facilities afforded in the libraries of
the Australian Museum, the CSIRO McMaster Animal Health Laboratory, the Royal Society
of New South Wales, and the Department of Geology and Geophysics at the University
of Sydney.

acne. | EDITOR’S FOOTNOTE

While the original manuscript contained new generic names for most of the synonyms,
the editor has followed reviewer’s suggestions that the new generic names be deleted and
left for future workers.

50 Aust. Zool. 19(1), 1976

A Redescription of Heteroclinus adelaidae Castelnau
(Pisces: Clinidae), With Description of a
Related New Species

DOUGLASS F. HOESE
The Australian Museum, College Street, Sydney, New South Wales, 2000.

ABS ERAGE

Heteroclinus adelaidae, a southern Australian clinid, is redescribed from the
holotype and numerous specimens from Tasmania, Victoria, South Australia and
Western Australia. A new and closely related species is described from Tasmania
and South Australia. The generic classification of the Australian clinids is briefly
discussed.

INTRODUCTION

Clinids are one of the more poorly known groups of Australian marine
fishes. The previous taxonomic studies have been based on limited material,
primarily from one geographical area at a time. Castelnau described several species
from southern Australia in the late 1800s, while Giinther described various species
from eastern and western Australia. McCulloch (1908) was the first Australian
worker to extensively examine the group, although he treated only eight species,
primarily from eastern Australia. Little subsequent work was done on the group
until Whitley (1941, 1945) examined types in Europe. Scott (1955, 1966 and
1967) has treated the known Tasmanian species extensively. Milward (1967)
revised the species known from Western Australia, and McKay (1970) described
an additional Western Australian species.

At present 34 species have been described from Australia. Whitley (1964)
recognized 18 species, two of which had been regarded as uncertain species. Types
of these two species, Heteroclinus adelaidae and Neoblennius fasciatus, were
examined in this study. Previously no thorough attempt has been made to examine
the group on a widespread geographical basis, due to lack of adequate material.
Recent field work in New South Wales, Victoria, Tasmania and South Australia
has alleviated this problem to some extent, with 25 species now being known, but
many of the species are still represented by few specimens. Most of the collecting
has been restricted to depths shallower than five metres. Springer (pers. comm. )
suggested that the ophiclinids are clinids, bringing the total number of Australian
species to over 30, comparable in number to the South African fauna.

Aust. Zool. 19(1), 1976 51.

DESCRIPTIONS OF HETEROCLINUS SPECIES

Many of the Australian species are superficially similar, although many differ
in live colour pattern, particularly of the head. In alcohol the colour fades, and
important species characters are often lost. The body colouration, however, tends
to be highly variable. Inadequate comparative material has led earlier workers to
incorrectly synonymize species and not separate closely related ones. It is apparent
that the range of variation in meristics within a species is narrower than previously
suspected.

With the exception of a species of Springeratus, the Australian species are
confined to temperate regions of eastern, southern, and south-western Australia.
Two of the Australian species occur at Lord Howe Island and one occurs in New
Zealand. The species occur in rocky tidepools, in association with algae around
rocky reefs, or in grass flats over mud or sand bottoms. All the species are
undoubtedly live-bearers.

METHODS

The following abbreviations are used in reference to material examined: AMS,
Australian Museum, Sydney; MNHN, Museum National d’Histoire Naturelle;
NMV, National Museum of Victoria, Melbourne; QM, Queensland Museum,
Brisbane; QVM, Queen Victoria Museum, Launceston; SAM, South Australian
Museum, Adelaide; TM, Tasmanian Museum and Art Gallery, Hobart; WAM,
Western Australian Museum, Perth.

In material examined lists, the number of specimens are given followed by
the size range of standard length in mm. All fish lengths given are standard

lengths (SL).

Counts and measurements follow those given by Hubbs and Lagler (1958).
The last anal ray and last dorsal ray are counted as separate. Vertebral counts
were determined from radiographs and cleared and stained specimens.

GENERIC CLASSIFICATION OF AUSTRALIAN CLINIDS

The generic classification of Australian clinids has not been thoroughly studied.
Five nominal genera (Heteroclinus, Neoblennius, Petraites, Clinus, and Cristiceps)
have been recorded from Australia, and a sixth, Springeratus, has been collected
from the Great Barrier Reef. Cristiceps has long been regarded as distinct. Some
Australian species have been placed in the African genus Climus while other species
have been placed in Petraites, with the two genera separated on the basis of the
dorsal spine count. Recently, Penrith (1969) suggested that Petraites was a
synonym of Clinus, and McKay (1970) has followed Penrith. Shen (1971),
however, has noted that African clinids lack the genital flaps characteristic of
Australian species. While it is apparent that the Australian species, all of which
are undoubtedly live bearers, are related to African genera, no thorough comparison

52 Aust. Zool. 19(1), 1976

DE. HOESE

has been made. Two of the Australian genera, Heteroclinus and Neoblennius, have
been treated as uncertain genera. Examination of the holotype of Heteroclinus
adelaidae Castelnau (1873) reveals it to be a distinct species subsequently
described as Cristiceps phillipi Lucas (1891). Similarly, the syntypes of Neoblennius
fasciatus Castelnau are representatives of the previously described Cristiceps
perspicillatus Cuvier and Valenciennes (1836). At present no attempt is made
to resolve Heteroclinus, Petraites, and Neoblennius problem. However, since
Heteroclinus is the oldest of the three genera, the type species is redescribed and
compared with a new closely related species.

The type species of Petraites, P. heptaeolus, and four closely related species
(P. wilsoni and three undescribed species) differ from H. adelaidae complex in
having three stout pelvic rays, a strongly compressed head, and the last two dorsal
rays widely separated from the preceding. It is unlikely that P. roseus and P.
fosteri also are referable to Heteroclinus. Other Australian species generally have
a more robust head, and many species have biserial head pores. Whether or not
these species belong in other genera is the subject of further study.

The species of the H. adelaidae and H. heptaeolus complexes differ substan-
tially from the African genus Clinus. Of the characters used to define Clinus
(Penrith, 1969), the shape of the orbital tentacle and presence of clusters of cirri
on the dorsal spines are often different between closely related Australian species
and do not seem of generic importance in these species. The species of these two
complexes differ from Clinus in the arrangement of lateral line pores, in having
three or more rows of teeth in the upper jaw, in having a more compressed head
and body, and in always having the anterior scales overlapping. In general body
form Heteroclinus is more similar to the African genus Pavoclinus than Clinus.

Heteroclinus Castelnau

Heteroclinus Castelnau 1873: 68 (type species, Heteroclinus adelaidae
Castelnau, by monotypy )

Since the Australian clinids are heterogeneous and require further study,
only the two species of the Heteroclinus adelaidae complex are characterized below:

Last two dorsal rays evenly spaced, not widely separated from the preceding
ray or spine. Head moderately compressed; body strongly compressed. Snout
pointed to rounded in lateral view. Body elevated along first dorsal margin. Infra-
orbital and preopercular mandibular pores small and uniserial. Nasal tentacle short
to moderately elongate, simple or weakly bilobed. Orbital tentacle short and
rounded, elongate and pointed, or elongate with numerous lateral lobes. Dorsal
origin above a point before preopercular margin to near end of eye. First three
dorsal spines usually separate from rest; membrane of first dorsal connected at
very base of first spine of second dorsal fin or to back before spine, rarely
connected to tip of first spine of second dorsal fin. Segmented caudal rays 11.
Pectoral rays typically 11 or 12. Second dorsal spines typically 29-31. Segmented

Aust. Zool. 19(1), 1976 | 53

DESCRIPTIONS OF HETEROCLINUS SPECIES

dorsal rays 2-5. Branchiostegals 6. Mouth small, ending below eye. Pelvics 1, 2,
two segmented rays slender and elongate reaching to near anal fin; an inner rudi-
mentary ray, visible only by dissection. Vomer with 1-2 rows of teeth forming
an inverted V. Precaudal vertebrae 13 to 15. Anteriorly, lateral line scales over-
lapping, with a posteromedian pore; posteriorly, near end of pectoral fin scales
separated with tubes at each end; behind pectoral fin lateral line scales few and
separate or absent.

Fig. 1. Attachment of last anal ray to caudal peduncle in Heteroclinus adelaidae and
_ H. macrophthalmus.

Considerable confusion has occurred over the identity of Heteroclinus adelaidae
and H. antinectes. Whitley (1945) designated a lectotype of Cristiceps antinectes
Giinther (1861) and indicated that it was a senior synonym of Petraites phillipi
(= H. adelaidae). Although one paralectotype is H. adelaidae, the lectotype and
two paralectotypes are of a different and distinct species. Milward (1967) followed
Whitley in treating Cristiceps phillipi Lucas as a synonym of Petraites antinectes.
However, the species treated by Milward (1967) as P. antinectes is an undescribed
species with biserial circumorbital pores not closely related to H. antinectes.
Recent sampling has revealed a new species closely related to H. adelaidae from
South Australia and Tasmania.

The two species treated here are easily distinguished from other Australian
clinids by the combination of the two slender pelvic rays, the broad connection of
the last anal ray to the caudal peduncle (Fig. 1) and the reduction of the lateral line.

Heteroclinus adelaidae Castelnau

Figures 2, 3a, and 4

-Heteroclinus adelaidae Castelnau 1873: 68 (type locality, Adelaide, South
Australia ).

Cristiceps phillipi Lucas 1891: 11, pl. 3, fig. 2 (type locality, Port Phillip Bay,
Victoria ).

Petraites phillipi — McCulloch 1908: 43, pl. 10, fig. 3 (Western Port, Victoria).
Scott 1966: 109, fig. 1d (Green’s Beach, northern Tasmania). Scott 1967:
194 (in part, Green’s Beach, Tasmania).

54 Aust. Zool. 19(1), 1976

DEF HOESE

‘UdTJA “D Aq o10yd “eruRUIsey, UseYyIOU Wry

appiwjapo snuoodajayY Ul UOHCUA INO[OD

WG a a

55

Aust. Zool. 19(1), 1976

DESCRIPTIONS OF HETEROCLINUS SPECIES

Description based on specimens 30 mm to 76 mm SL. An asterisk indicates
count of holotype.

A

aS

Fig. 3. Intromittant organ of males of Heteroclinus adelaidae (A) and H. macrophthalmus

(B).

Pectoral rays 10> (in 1), 24--Gin 71)*, 12 (in 10), 13 (in 1). Seomentea
caudal rays 11 (in 84)*. Gill rakers usually 2 + 6, total rakers 6 (in 1), 7 (in 6),
8 (in 39), 9 (in 3). Branchiostegal rays 6 (in 17). Dorsal, anal, and lateral line
counts given in Table 1 and 2. Longitudinal scale rows 86 to 118.

Head moderately compressed, width 14.5 to 17.2% of SL. Body strongly
compressed. Snout obtusely pointed, snout length less than eye diameter. Eye
moderate, length 7.5 to 9.0% of SL. Interorbital narrow, about one half of eye
diameter. Mouth short, reaching to a point below pupil; upper jaw length 11.0
to 14.3% of SL in males, shorter in females, 9.7 to 11.7% of SL. Anterior nostril
tubular with a short, slender, posterior, simple or bilobed flap, positioned midway

15mm

Fig. 4. Head of Heteroclinus adelaidae showing head flaps and sensory pores.

56 Aust. Zool. 19(1), 1976

D. F. HOESE

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Aust. Zool.

DESCRIPTIONS OF HETEROCLINUS SPECIES

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19(1), 1976

Aust. Zool.

58

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between upper jaw and posterior nostril. Posterior nostril a pore above anterior
margin of eye. Orbital tentacle (Fig. 4) low and rounded, sometimes elongate,
width varying from equal to length to twice as long as length; margin smooth;

length 1.0 to 3.4% of SL.

First dorsal fin higher than second; first and second spines longest and about
equal in length, third spine slightly shorter; fin lower in females. Second dorsal
spine varying from about 10.5% of SL in young males, less than 50 mm SL to
14-17% of SL in larger males; varying from 8.6 to 15.3% of SL in females.
First dorsal origin over end of preopercular margin, last spine above a point behind
middle of operculum. Membrane of first dorsal usually connected basally to second
dorsal, varying from attaching to body before second dorsal fin to tip of first spine
of second dorsal. Flaps from dorsal spines bound in interspinal membranes, never
free. Second dorsal origin above a point behind pelvic insertion. Anteriorly spines
in second dorsal short, becoming progressively longer posteriorly, last spine 10.0
to 12.5% of SL; first ray longer than last spine. Dorsal rays evenly spaced. Last
dorsal ray connected by membrane to caudal base. Anal origin below 10th or 11th
spine in second dorsal fin. Anal II, 21-25, posterior rays becoming progressively
longer; last ray shorter than preceding and broadly connected to two thirds of
caudal peduncle. Caudal short and rounded, with 11 segmented rays and smaller
procurrent rays above and below. Caudal length 19 to 22% of SL. Pectoral rays
elongate, middle rays largest reaching to a point above or behind anal origin.
Pelvics with a hidden spine, 2 elongate rays and a third rudimentary ray visible
only upon dissection; two developed rays slender and about equal in length or
inner ray slightly longer, reaching approximately to anal origin.

Gill rakers short and simple. Tongue tip broadly rounded, or tapering to a
point. Intromittant organ elongate and curved forward, with bilobed papillae
attached anteriorly at base. (Fig. 3a).

Head pores shown in Figure 4. Circumorbital and preopercular pores uniserial.
Circumorbital pores (exclusive of median pores and pore below anterior nostril)

12 (in 47), 13 (in 9), 14 (in 17), 15 (in 1), number constant over size range
studied.

Head naked, body scales small and cycloid, extending forward to above oper-
culum below first dorsal. Scales overlapping and forming rows anteriorly, becoming
nonimbricate and irregular posteriorly. Anterior lateral line scales overlapping with
a small terminal and median or dorsomedian pore; first scale also with a ventral
pore; posteriorly scales separate with pores at each end, extending in an elevated
arch curving downward near end of pectoral fin; usually no scales along midside
beyond pectoral tip, but occasionally up to 8 widely spaced scales in a line
extending to below middle of second dorsal. '

Upper jaw with outer row of conical teeth slightly enlarged extending over
most of premaxilla. Anteriorly five inner rows of small conical teeth tapering
posteriorly to a single row. Lower jaw with outer row of conical teeth slightly

Aust. Zool. 19(1), 1976 ‘69

DESCRIPTIONS OF HETEROCLINUS SPECIES

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BF H@ESE

enlarged covering all of dentary. Anteriorly three inner rows of smaller conical
teeth tapering posteriorly to a single row. In males smaller than 50 mm a single
row of vomerine teeth in an inverted V. In larger males an incomplete inner row
present. In females, usually only a single row of vomerine teeth. Palatine edentulus.

Colouration in alcohol variable (Fig. 2). Head and body varying from tan

to dark brown. In some (including holotype) an oblique white bar extending from
eye across preoperculum and operculum to pectoral base, continuing onto pectoral
base divided by a dark brown bar. Stripe continues on body from base of pectoral
to end of caudal peduncle. Head often with several dark brown spots, less than
pupil diameter; spots occasionally extending onto body. Spots usually larger and
forming a row along margins of head bar when present. In holotype a second
white stripe present directly below anterior base of dorsal fins. Body often with
5 to 8 dark brown irregular vertical bands, sometimes broken into one upper and
one lower row of larger irregular shaped blotches, with upper blotches darkest.
Some specimens uniform tan or brown. Lateral line scales usually black anteriorly.
Fins varying from uniform tan to mottled to banded. First dorsal fin sometimes
black and darker than second dorsal fin. Banded individuals often with narrow
extensions of bands onto second dorsal and anal fins either continuous with body
bands or as separate blotches. Caudal and pectoral clear, dusky, or narrowly
banded (pectoral banded in holotype). Pelvic fins light tan.

A moderate sized species reaching a size of 80 mm.

Variation.—H. adelaidae is peculiar among Australian clinids in being sexually
dimorphic in jaw length and first dorsal spine height (Table 4). Males have a
larger mouth and a higher dorsal fin than in females.

Lateral scale counts show considerable variation. Counts of 8 specimens from
each of 3 States gives counts of 86 to 99 for South Australia, 95 to 103 for
Victoria and northern Tasmania and 97 to 118 for southern Tasmania.

TABLE 4

SEXUAL DIMORPHISM OF HETEROCLINUS ADELAIDAE IN JAW LENGTH AND
SECOND DORSAL SPINE LENGTH

Measurements expressed as a percentage of standard length, rounded to the nearest per cent.
The size distributions for the two sexes are similar, ranging from 47 to 65 mm SL. Based on
sample from Green’s Beach, Tasmania, January 1968.

Sex Jaw length as a per Second dorsal spine length as a
cent of standard length per cent of standard length
50 YEA b) eae BA a eee BaF TOS ER AB Ta eas AG
Males DNS), OY CS teed Beg Det Gl Ge Pe ead
Females 7 14 #2 — — 2 3113 -4—-— — —

Aust. Zool. 19(1), 1976 61

DESCRIPTIONS OF HETEROCLINUS SPECIES

As noted above colouration is highly variable. Although variation: within a
sample is high, from the limited material available it appears that the variation is
greater between samples from different geographical areas.

Too few specimens are available to determine the extent of geographical
variation. However, the data suggest that there is probably population variation
in the height of the first dorsal fin and in the number of the lateral line scales
along the straight portion of the lateral line. The meristic data suggest that there
may be clinal variation in the number of second dorsal spines and anal rays and
vertebrae in South Australia and Tasmania, with counts tending to be higher in
the southern areas of these States (Tables 1 and 3).

The species ranges throughout Tasmania, Victoria, South Australia and is
known from southern Western Australia.

In general head shape, body form, reduction of the lateral line, the broad
connection of the last anal ray to the caudal peduncle, the presence of two slender
pelvic rays and in the shape of the intermittent organ H. adelaidae is closest to
H. macrophthalmus. H. adelaidae differs from that species in lacking free lobes
from the dorsal spines, supraorbital and tentacle shapes and in segmented dorsal
ray counts.

While most Australian clinids occur in association with algae around rocky
reefs, H. adelaidae was taken in this study only from seagrass beds in shallow
water. One specimen was dredged from 10 to 18 metres.

MATERIAL EXAMINED

‘Holotype. MNHN A.1077, a female 76.0 mm SL.

Victoria: Petersborough, AMS 1.16987-011 8 (35-40). Port Phillip Bay,
AMS 1.2511, 1 (33). Westernport Bay, NMV (uncatalogued, dredged between
Crawfish Point and Eagle Rock), 7 (28-79); AMS IA.1318, 1 (64); AMS 1.7610,
2 (48-69); AMS 1.9006-9008, 3 (60-75).

Tasmania: Green’s Beach, QVM (out of 1972/5/231 (B)), 2 (47-90).
QVM (out of 1972/5/225 (B)), 63 (46-66). Kelso, QVM (out of 1972/5/212
(C)), 10 (46-65); QVM 1972/51377, 11 (51-67). Port Latta, AMS I.17588-001,
2 (60-80). Port Arthur, AMS 1.17550-003, 2 (41-63). Coles Bay, AMS I.17553-
006, 7 (50-74). Wedge Bay, AMS 1.17193-001, 2 (27-33). Bruny Island, TM
B95 cd 8 3))s

South Australia: Port Vincent, AMS IB.7133, 1 (71). Venus Bay, SAM
F.3410, 2 (68-69). Port Lincoln, NMV_ (uncatalogued), dredged 10-18 m
Spencer Gulf off Port Lincoln, taken with holotype of H. macrophthalmus, 1
(64). Marion Bay, Yorke Peninsula, SAM F.3629, 2 (30-55). Kangaroo Island,
SAM F348, 1 (75).

62 Aust. Zool. 19(1), 1976

D2 Fi) H@ESE

~ Western Australia: Emu Point, near Albany, WAM P.21687,-1 (66). Frey-
cinet Harbour, BM (NH) 1974.11.29.3 (formerly part of 1858.12.27.67), male
66.5 mm SL, a~paralectotype of Cristiceps antinectes. Cockburn Sound off
Rockingham, WAM P.25258-002, 10 (38-64).

Heteroclinus macrophthalmus n. sp.
Figures 3b, 5, 6 and 7

Petraites phillipimScott 1967: 194 (in part, ‘abnormal specimen” from Green’s
Beach only).

Based on 39 mm male and 64 to 83 mm females.

Measurements of types given in Table 5. Counts of holotype marked with an
asterisk. Pectoral rays 12* (in 4), 13 (in 1). Segmented caudal rays 11* (in 5).
Gill rakers 2 + 6 (in 1), 3 + 6 (in 1). Branchiostegal rays 6 (in 2). Dorsal ITI,
BOO, 4 ag 1) ET XT 9 * Cin. 4)- Anal-rays TF, 25*' fin: 3); 11; 26 (in-1);
II, 27 (in 1). Lateral line scales — arched and straight part of lateral line
oe he (ice ek? it 26 24 (in Be 20 25%, (in 1) 2a. 19
(in 1). Longitudinal scale rows 127 (in 1), 119 (in 1), 110 (in 1).

TABLE 5

MEASUREMENTS OF HOLOTYPES OF TWO SPECIES OF HETEROCLINUS
(in millimetres)

Measurement H. adelaidae H. macrophthalmus
MNHN A 1077 NMV A.495
Standard length 76.0 64.0
Head length 21.6 18.0
Predorsal length 15.0 11.2
Body depth at anal origin 15.5 13.8
Caudal peduncle depth 3.5 3.0
Caudal peduncle length 6.9 6.2
Upper jaw length 8.3 8.2
Eye length 6.3 6.9
Snout length 4.1 3.4
Pectoral length 14.5 12.8
Pelvic length 15.1 15.0
First dorsal spine length 8.8 8.2
Last dorsal spine length 9.5 7.6
First dorsal ray length 11.4 ot
Caudal length 15.1 14.3
Orbital tentacle length | 1.5 3.1

Aust. Zool. 19(1), 1976 : yabeeen a 63

DESCRIPTIONS OF HETEROCLINUS SPECIES

Head moderately compressed, width 17.0 to 17.3% of SL. Body strongly
compressed. Snout blunt, rounded in dorsal view, much shorter than eye diameter.
Eye large 9.7 to 10.8% of SL. Interorbital narrow, about half eye diameter. Mouth
he teaching to a point below middle to end of pupil. Upper jaw length 11.6 to
12.7% SL in females and 11.79% SL in male. Anterior nostril tubular with a large
posterior branched tentacle, composed of 3 to 6 elongate lobes; anterior nostril
positioned about half way between snout tip and posterior nostril. Posterior nostril
with a raised rim, positioned above anterior margin of eye. Orbital tentacle
(Fig. 7) flattened with 5 to 10 slender lobes extending from base; length 4.9
to 6.1% of SL.

age | RO —

Fig. 5. Female holotype of Heteroclinus macrophthalmus, 64 mm SL, based on preserved
colouration.

First dorsal fin higher than second; first and second dorsal spines about equal
in length, third spine slightly shorter. ceband spine 9.2% SL in 39 mm male and
12.6 and 14.6% SL in large females. First dorsal origin over end of posterior
margin of preoperculum, last spine above a point behind middle of operculum.
Membrane of first dorsal fin separate from second dorsal or connected to second
dorsal at very base. Tips of first 3 dorsal spines with 3-6 elongate free flaps. First
spine in second dorsal fin without flaps. Tip of second spine in second dorsal fin
with 1 or 2 free flaps. Tips of remaining dorsal spines with an elongate free flap.
Second dorsal origin above a point behind pelvic insertion. Anteriorly spines in
second dorsal fin short, becoming progressively longer posteriorly, last spine 10.2
to 11.8% of SL; first ray longer than last dorsal spine. Dorsal rays evenly spaced.
Last dorsal ray connected by membrane to caudal base. Anal origin below 11th
and 12th spine in second dorsal fin. Posterior rays of anal becoming progressively
longer; last ray shorter than preceding and broadly connected to about two thirds
of caudal peduncle. Caudal short 17.9 to 22.4% of SL and rounded, with 11
segmented rays and smaller procurrent rays above and below. Pectoral rays
elongate, middle rays longest. Pelvic rays I, 3, third ray a short rudiment visible

64 Aust. Zool. 19(1), 1976

D. F. HOESE
only upon dissection; 2 slender outer rays about equal in length, reaching approxi-
mately to anal origin.
Gill-raker short and simple. Tongue tip free, tapering to a point.

Intromittent organ elongate and curved forward, with bilobed papillae attached
anteriorly at base (Fig. 3b).

Head pores shown in figure 7. Circumorbital and preopercular pores uniserial.
Circumorbital pores (exclusive of median pore below anterior nostril) 13 (in 3).

Squamation as in H. adelaidae, except that straight part of lateral line more
extensive, with 12-24 scales, reaching well beyond tip of pectoral fin.

: Sept TS ay

: HK i Aa ZA

Fig. 6. Male paratype of Heteroclinus macrophthalmus, 39 mm SL, based on Kodachrome of
fresh specimen.

Live colouration of male paratype from Kodachrome. Head and anterior half
of body dark with brown or maroon; belly black; posterior half of body lighter
brown. A prominent oblique silver bar, less than an eye diameter in width,
extending from posteroventral margin of eye over preoperculum and operculum.
Midside of body with silver stripe anteriorly, broken into irregular shaped silver
blotches, posteriorly. Two faint narrow vertical bars below eye. Two white spots
at angle of preoperculum. a broken black stripe on body below dorsal fin. Second
dorsal fin with 2 narrow reddish brown longitudinal stripes, more prominent
posteriorly. Six broad, faint yellow vertical bands on second dorsal fin. Anal fin
reddish brown, with some yellow along base. Caudal clear with 7 reddish brown
irregular bands. Pectoral clear with several narrow irregular reddish brown bands.
Ventrals maroon. No black along anterior part of lateral line.

Colouration in alcohol of male paratype. Head and body tan. Silver and white
blotches and stripes of fresh specimens tan in alcohol. Dark brown stripe behind
eye continuing on body to near caudal base, upper margin of stripe below dorsal
base sharply defined; ventral margin diffuse. Cheek dark brown. Sides of belly
dark brown; rest of belly tan. First dorsal black anteriorly. Second dorsal dusky
with 2 faint narrow horizontal dark stripes. Caudal clear with 7 irregular dusky
bands. Anal dusky. Pectoral clear with several faint wavy dusky bands. Pelvic fin

Aust. Zool. 19(1), 1976 65

DESCRIPTIONS OF HETEROCLINUS SPECIES

clear. Female colouration. Colour pattern of holotype shown in figure 5. Head and
body tan. A diffuse dark brown, broken stripe on back, from behind eye to caudal
peduncle. Six narrow transverse bands extend dorsally and ventrally from stripe,
some extending on to dorsal and anal fins. Sides of head with dark brown mottling.
Fins clear.

The Tasmanian and the largest South Australian paratype differ from the
South Australian specimens in having a more extensive lateral line and in having
2 (rather than 1) rows of vomerine teeth. Since the extent of the lateral line
raises geographically in H. adelaidae and the number of vomerine teeth increase in
size in H. adelaidae these differences are not regarded as indicating any specific:
differences. Since only one small male has been studied, it is not known if H.
macrophthalmus exhibits the sexual dimorphism characteristic of H. adelaidae.
Although the male paratype has a different colouration from the female, it may not
reflect sexual dimorphism since H. adelaidae, which has a similar colour variation,
may not be sexually dichomatic.

15mm

Fig. 7. Head of Heteroclinus macrophthalmus showing head flaps and sensory pores.

The species is known only from South Australia and northern Tasmania. It
is closest to H. adelaidae (see discussion of that species). H. macrophthalmus differs
from that species in having a larger eye, the free lobes on the dorsal spines, the
branched orbital and nasal tentacles with numerous elongate lobes, a typically
higher number of dorsal, anal and pectoral rays, a different snout shape, and more
numerous lateral line scales.

This species has been taken only from grass flats subtidally and from 10-18
metres, twice in association with H. adelaidae.

66 Aust. Zool. 19(1), 1976

D. F. HOESE

MATERIAL EXAMINED
Holotype — NMV A.495, a 64 mm female, off Port Lincoln South Australia.
Coll. J. Vestch 13 Mar. 1969. Dredged 10-18 m.

Paratypes — AMS 1.17609-001, a 39 mm male, Victor Harbour, South
Australia, West Island Aquatic Reserve; coll. D. F. Hoese and W. Ivantsoff, 20
Dec. 1973. 0-2 m, grass flat and algae, boulders. QVM 1974/5/131, a 75 mm
female, Green’s Beach, Tasmania; coll. R. H. Green, 9 Dec. 1965, grass flat and
rocks. SAM F.1540, an 83 mm female, Spencer or St. Vincents Gulf, 19 Feb.
1931. SAM F.2589, a 73 mm female, Spencer or St. Vincents Gulf.

ACKNOWLEDGEMENTS

I would like to thank W. and H. Ivantsoff, N. and W. Congleton, and D. Brown for
assistance with field work. J. Glover, South Australian Museum, J. Dixon, National Museum
of Victoria, R. H. Green, Queen Victoria Museum, P. Andrews, Tasmanian Museum and
Art Gallery, G. Allen and B. Hutchings, Western Australian Museum, M. L. Bauchot,
Museum National d’Histoire Naturelle, Paris, and G. Palmer, British Museum (Natural
History) kindly made material available for this study. Mrs. H. K. Larson drew the
illustrations.

REFERENCES

CASTELNAU, F. (1873). Contribution to the Ichthyology of Australia. No. IV. Fishes of
South Australia. Proc. Zool. & Acclim. Soc. Vict. 26, 59-82.

CUVIER, G. & A. VALENCIENNES (1836). Histoire naturelle des poissons, //, Paris
506 pp.

GUNTHER, A. (1861). Catalogue of the Acanthopterygian Fishes in the Collection of the
British Museum, Vol. III. British Museum, London 586 pp.

HUBBS, C. L. & K. F. LAGLER (1958). Fishes of the Great Lakes region. Cranbrook
Institute of Science — Bloomfield Hills, Michigan, 213 pp.

LUCAS, A.H. (1891). On the occurrence of certain fish in Victorian Seas, with descriptions
of some new species. Proc. Roy. Soc. Vic. (N.S.), 3, 8-14.

McCULLOCH, A. R. (1908). Studies in Australian Fishes, No. 1. Rec. Austr. Mus. 7, 36-43.

McKAY, R. J. (1970). Additions to the fish fauna of Western Australia — 5. Fisheries
Bull. W.A. 9 (5), 3-24.

MILWARD, N. E. (1967). The Clinidae of Western Australia (Teleostei, Blennioidea).
J. Roy. Soc. W. Austr. 50 (1), 1-9.

PENRITH, M. L. (1969). The systematics of the fishes of the Family Clinidae in South
Africa. Ann. S, Afr. Mus. 55 (1), 1-121.

SCOTT, E. O. G. (1955). Observations on some Tasmanian Fishes. Part VII. Pap. Proc.
Roy. Soc. Tasm. 89, 131-146.

SCOTT, E. O. G. (1966). Observations on some Tasmanian Fishes. Part XIV. Pap. &
Proc. Roy. Soc, Tasm. 100, 93-115.

SCOTT, E. O. G. (1967). Observations on some Tasmanian Fishes: Part XV. Pap. & Proc.
Roy. Soc. Tasm. 101, 189-220,

SHEN, S. (1971). A new genus of clinid fishes from the Indo-West-Pacific, with a redescrip-
tion of Clinus nematopterus. Copeia (1), 697-707.

WHITLEY, G. P. (1941). Ichthyological notes and illustrations. Austr. Zool. /0, 1-50.

WHITLEY, G. P. (1945). New sharks and fishes from Western Australia. Part 2. Austr.
Zool. Il, 1-42.

WHITLEY, G. P. (1964). Presidential Address. A survey of Australian Ichthyology. Proc.
Linn. Soc. N.S.W. 89 (1), 11-127.

Aust. Zool. 19(1), 1976 67°

Mosquitoes Feeding on Ectothermic Vertebrates:
A Review and New Data

HAROLD HEATWOLE and RICHARD SHINE’
Department of Zoology, University of New England, Armidale, N.S.W. 2351,
Australia

ABSTRACT

The literature on feeding by mosquitoes on ectothermic vertebrates is reviewed.
Original observations of the feeding of Uranotaenia argyrotarsis on Rana daemelii
on Cape York Peninsula are presented.

LITERATURE REVIEW

Gilett (1971) in a review of mosquito biology lists lizards, snakes and frogs
as hosts of filarial parasites transmitted by Anopheles quadrimaculatus and Aédes
taeniorhynchus, mentions a trematode lung fluke of frogs which is transmitted from
snails via larvae of Anopheles maculipennis to frogs (mosquito perhaps eaten by
frog), and indicates that lizards and frogs are known to have non-human
Plasmodium malarias which are presumably transmitted by mosquitoes. He also
mentions that Culex tarsalis feeds on snakes and cites his personal observations of
Coquillettidia fuscopennata feeding on a variety of vertebrates including frogs and
toads. Some species of Culex feed exclusively on ectotherms (Woke, 1937a).

Aédes aegypti has frequently been observed to feed on ectotherms. Yuill
(1964) developed the ingenious technique of using individuals of this species as
micro-syringes for withdrawing small blood samples from Thailand geckos (Hemz-
dactylus frenatus and Platyurus platyurus) for viraemia determinations of Japanese
encephalitis; in the laboratory this same species fed on two species of Puerto
Rican lizards, Anolis cristatellus and Anolis pulchellus. (Fox. and Bayona, 1964),
and on the Oriental lizards Gecko verticillata and Calotes sp. (Toumanofl, 1949);
the latter reported that Aédes albopictus also feed on G. verticillata and Calotes.
Woke (1937a, b) records A. aegypti feeding on a turtle (Terrapene carolina) and
on a frog (Rana clamitans). Gordon and Lumsden (1939) induced this species
to feed on frogs (Rana sphenocephala) but could not get Anopheles maculipennis

1. Present address: Dept. Biology, University of Utah, Salt Lake City, Utah, U.S.A.

Aust. Zool. 19(1), 1976 69

HAROLD HEATWOLE and RICHARD SHINE

Fig. 1. Uranotaenia argyrotarsis feeding on Rana daemelii in the field. Note blood-engorged

mosquito piercing upper eyelid and the swollen appearance of the eyelid.

79 Aust. Zool. 19(1), 1976

MOSQUITOES FEEDING ON FROGS

or Culex molestus to do so. Although A. aegypti obviously has wide food
preferences, it does not feed on all species of ectothermic vertebrates, as in their
literature review of this species, Fox and Bayona (1964) mention reports of failure
to induce it to bite the lizard Urocentron azureum and some Brazilian geckos. Woke
(1937a) also reports this species rejecting a lizard.

There is disagreement as to the relative nutritive value of the blood of
ectotherms and homeotherms for this species. Woke (1937a,b) found that
ectotherm blood was equal or superior to that of some homeotherms whereas
Toumanoft (1949) found that A. aegypti feeding on lizards matured fewer eggs
than those feeding on humans.

Thomas and Ekland (1960) found that garter snakes (Thamnophis sirtalis
parietalis and T. ordinoides vagrans) experimentally infected with western equine
encephalitis virus could transmit it to a mosquito vector of the disease (Culex
tarsalis) even after having overwintered under artificial conditions subsequent to
infection with the virus. Gebhardt e¢ al. (1964) further showed that at least four
species of snakes (Thamnophis elegans vagrans, T. sirtalis, Pituophis catenifer and
Coluber constrictor) served as natural overwintering hosts for the virus. Under
certain climatic conditions (especially high per cent of available sunshine in late
spring or early summer), favourable both for large mosquito hatches and viraemia
in infected snakes, epidemics resulting from transmission to warm-blooded hosts
might result.

Craighead et al. (1962) suggest that lizards (Ameiva, Cnemidophorus,
Basiliscus, Iguana) may play a role in the natural history of eastern equine
encephalitis in Panama. A number of papers have recently indicated that a number
of snakes, lizards and turtles can be experimentally infected by Japanese encepha-
litis virus or by western equine encephalitis virus and that natural reptilian
populations often have high levels of antibodies to such viruses (see review by
Shortridge et al. 1975). Hoff and Trainer (1973) in a review list 7 species
of Aédes, 6 species of Culex, 2 of Culiseta, and Mansonia perturbans and
Anopheles quadrimaculatus having been reported to feed on lizards, turtles and
snakes. To the above records Tempelis and Galindo (1975) recently added 4
species of Culex that feed predominantly on reptiles and mention that some
Deinocerites feed preferentially on ectotherms. Pinger and Rowley (1975) addi-
tionally record Aédes trivittatus as sometimes feeding on amphibians and reptiles.

The genus Uranotaenia seems to have a predilection for amphibian hosts.
Remington (1945) records Uranotaenia lowii feeding on the frog Rana spheno-
cephala, the tree frog Hyla cinerea, and the toad Bufo valliceps in the field, and
upon H. cinerea, B. valliceps, B. woodhousii fowleri and the salamander Desmog-
nathus fuscus auriculatus in the laboratory; only the salamander reacted to the
bites. In laboratory tests, this mosquito rejected box turtles, two species of lizards,
and humans; it seems to be largely restricted to amphibians.

Aust. Zool. 19(1), 1976 71

HAROLD HEATWOLE and RICHARD SHINE

Fig. 2. Uranotaenia argyrotarsis piercing the eyelid of Rana daemelii in the field. Note the
whitish swelling on the frog’s right eyelid,

72 Aust. Zool. 19(1), 1976

MOSQUITOES FEEDING ON FROGS ©

Marks (1960) records Uranotaenia albescens and Culex (Lophoceraomyia) sp.
biting the hylid frog Litoria caerulea; this frog harbors a hemogregarine parasite
in its blood and Marks suggested these mosquitoes may act as the invertebrate host.
Her observations represent the only previous record of mosquitos feeding on
ectothermic vertebrates in the Australian region.

OBSERVATIONS AND DISCUSSION

The present paper reports still another species of Uranotaenia feeding on
amphibians, again from Australia. While collecting frogs at night along a small
stream 16 km north-west of Iron Range, Queensland, on 6 September 1972, we
noticed that a number of Rana daemelii were being bitten by mosquitoes. Many
of the mosquitoes were observed to be blood-filled. Nine mosquitoes were collected
and subsequently proved to be female Uranotaenia argyrotarsis Leceister. No other
species was observed on the frogs. We subsequently approached the frogs cautiously
using flashlights. The frogs remained motionless as long as the light was kept upon
them and they could be closely observed. Of the 12 specimens captured and about
three times that number observed but unmolested, nearly all were being bitten by at
least one mosquito (Fig. 1) and some had as many as six biting simultaneously.
The site of biting was always on the head, especially the upper eyelids, while no
mosquitoes were observed biting the body or legs although these areas were
exposed. The bites resulted in raised whitish swellings (Fig. 2). On no occasion
did any frog give evidence of being irritated by the insects or in fact to be even
aware of their presence.

We do not know how host-specific these mosquitoes are. However, we did
not observe them biting any other species of frog in the area and they did not
attempt to bite us although we were in close proximity to them for the several
hours we observed the frogs. It is certain that they are not restricted to R. daemelii
however, as the mosquito’s geographic range is much wider than that of the frog.
U. argyrotarsis occurs in Malaya, the Moluccas, the Philippines, New Guinea, the
Solomon Islands and in Australia in the Northern Territory and north Queensland
(Marks, pers. comm.). R. daemelii is found only on Cape York Peninsula, Aust-
ralia. Some herpetologists consider R. daemelii as a subspecies of Rana papua.
Even the range of R. papua in this broad sense is much smaller than the range of
the mosquito, encompassing only New Guinea and Cape York.

We had the opportunity to collect R. daemelii at a different locality. On 7
September 1972, 24 km east of Iron Range, we obtained frogs from around small
ponds and found that frogs are attacked in this area also. The swelling caused by
mosquito bites are ephemeral except on the eyelids where they persist for some
time, even in preservative. The preserved collections from the two sites were

Aust. Zool. 19(1), 1976 73

HAROLD HEATWOLE and RICHARD SHINE

examined in November 1975. On the 12 frogs collected on the 6th, 5 (45%) had
one or both eyelids swollen; the corresponding value for the 40 collected by a
pond on the 7th was 18 (45%). Thus, the incidence of biting seems to be
similar in both localities. However, at the time of collection, no mosquitoes were
seen on the frogs in the second area despite the fact that the collection was made
at approximately the same time of evening as before. Weather conditions were
probably responsible. The 6th was a calm night and no wind was noticeable. The
7th was windy and this may have inhibited mosquito activity.

Our observations in conjunction with similar ones from the literature suggest
that feeding of mosquitoes on ectothermic vertebrates may be more common than
generally realised. There may be economic and/or medical implications. A possible
benefit which might accrue from examination of mosquito-transmitted diseases to
amphibians could be the selective control of introduced amphibian pests in
Australia such as the cane toad (Bufo marinus).

Studies are badly needed which survey (1) the range of hosts attacked by
different species of mosquitoes, (2) the range of mosquito species biting particular
species of amphibians and reptiles, (3) the diseases and/or parasites transmissible
between mosquitoes and ectotherms, and (4) the environmental factors related to
such transmission. Also studies evaluating ectotherms as potential reservoirs of
viral or other mosquito-borne diseases attacking man, domestic animals or wildlife
would be extremely valuable.

ACKNOWLEDGEMENTS
We are grateful to the Department of Zoology, University of New England, for financial
support of part of the Cape York Expedition during which these observations were made,
to Miguel Heatwole and Alana Young for assistance, to Dr. E. N. Marks for identification
of the mosquitoes, to Professor A. F. O’Farrell and J. Parmenter for criticism and sugges-
tions, and to Audry Heatwole for aid in preparation of the manuscript.

REFERENCES
CRAIGHEAD, J.E., A. SHELOKOV & P. H. PERALTA (1962). The lizard: a possible
host for eastern equine encephalitis virus in Panama. Amer. J. Hygiene 76, 82-87.
FOX, I. & I. G. BAYONA (1964). Aédes aegypti feeds on lizards in Puerto Rico. J. Econ.
Ent. 57, 417-418.

GEBHARDT, L: P.;:G: J: SEANEON, D: W. BIEL &) GC) COLLET (1964). Natal
overwintering hosts of the virus of western equine encephalitis. New England J. Med.
ZINA IL AG Te

GILETT, J. D. (1971). Mosquitoes. Weidenfeld & Nicholson, London, 274 pp.

GORDON, R. M. & W. H. R. LUMSDEN (1939). A study of the behaviour of the
mouth-parts of mosquitoes when taking up blood from living tissue; together with

some observations on the ingestion of microfilariae. Ann. Trop. Med. Parasit. 33,
259-278.

74 Aust. Zool. 19(1), 1976

MOSQUITOES FEEDING ON FROGS

HOFF, G. & D. O. TRAINER (1973). Arboviruses in reptiles: isolation of a Bunyammera
group virus from a naturally infected turtle. J. Herpetology 7, 55-62.

MARKS, E. N. (1960). Mosquitoes biting frogs. Aust. J. Sci, 23, 89.

PINGER, R. R. & W. A. ROWLEY (1975). Hest preference of Aédes trivittatus (Diptera:
Culicidae) in central Iowa. Amer. J. Trop. Med. Hygiene 24, 889-893.

REMINGTON, C. L. (1945). The feeding habits of Uranotaenia lowii Theobald (Diptera:
Culicidae). Ent. News 56, 32-68.

TEMPELIS, C. H. & P. GALINDO (1975). Host-feeding patterns of Culex (Melano
convon) and Culex (Aedinus) mosquitoes collected in Panama. J. Med. Ent. /2,
205-209.

TOUMANOFF, C. (1949). L’hemophagie varieé et l’activité reproductrice chez Aédes
aegypti L. et Aédes albopictus Skuse. Bull. Soc. Pathol. Exotique 42, 466-470.

THOMAS, L.A. & C. M. EKLUND (1960). Overwintering of western equine encephalo-
myelitis virus in experimentally infected garter snakes and transmission to mosquitoes.
Proc. Soc. .Bxp.: Biol. Med:.,105, 52-55.

YUILL, T. M. (1969), Mosquitoes for drawing blood from small reptiles. Trans. Roy. Soc.
Trop. Med. Hyg. 63, 407-408.

WOKE, P. A. (1937a). Cold-blooded vertebrates as hosts for Aédes aegypti Linn. J.
Parasitol. 23, 310-311.

WOKE, P.A. (1937b). Comparative studies of the blood of different species of vertebrates
on egg production of Aédes aegypti Linn. Amer. J. Trop, Med. 17, 729-745.

Aust. Zool. 19(1), 1976 75

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Biology of Neophyllaphis brimblecombei Carver
(Homoptera: Aphididae) in the Sydney Region

DINAH F. HALES
School of Biological Sciences, Macquarie University, North Ryde, N.S.W. 2113

The annual cycle of Neophyllaphis brimblecombei Carver in Sydney is described.
The aphids generally occur only on young shoots of their host plants, Podocarpus
elatus and P. spinulosus. Fundatrices hatch and colonise new shoots on P. elatus
in late July or August. Winged oviparous females are present by the end of
September. Alatae viviparae and males are uncommon. The shoots mature in
November, and the aphid population then disappears. The eggs laid by the
oviparae diapause until the following July or August. The cycle is similar on
P. spinulosus, but is correlated with the variable commencement of shoot
development in the host plant. Shoots produced on P. elatus in response to
pruning are colonised by WN. brimblecombei, as are naturally occurring autumn
shoots on P. spinulosus. Annual cycles of other Neophyllaphis spp. on Podocarpus
are compared with that of N. brimblecombei and possible factors controlling
production of sexuales are discussed. The relatively low success of aphids in
Australia is discussed in relationship to factors controlling production of sexual
forms.

INTRODUCTION

Only 130 species of Aphidoidea are known from Australia, including 17
species probably restricted to native plants and known only from Australia (Carver,
pers. comm.). A large proportion of the species now found here are thought to
have been introduced accidentally on their host plants. As the total number of
described aphid species exceeds 3600 (Eastop, 1970), the Australian aphid fauna
is disproportionately small.

The biology of aphids occurring primarily on native plants in Australia is
virtually unknown, the only exception being Sensoriaphis furcifera (Carver and
Hales, 1974). Yet the subject is of considerable interest, particularly in view of
the small representation of aphids in the Australian fauna. It appears that the
Australian environment has been unfavourable for radiation of this group.

The genus Neophyllaphis has many primitive characteristics and is associated
with the ancient plant genus Podocarpus, which originated in the Gondwanaland
flora and has since spread into Asia and Africa. (Some species of Neophyllaphis
occur on host plants belonging to other genera, e.g. N. rappardi Hille Ris Lambers

Aust. Zool. 19(1), 1976 77

DINAH F. HALES

on Agathis in West Irian, N. michelbacheri (Essig) on Pilgerodendron in South
America, N. araucariae Takahashi on Araucaria in Australia, New Guinea,
Hawaii, Mauritius.) Neophyllaphis brimblecombei Carver occurs on Podocarpus
elatus R. Br. ex Endl. in coastal regions of New South Wales and Queensland.
It is very similar morphologically to N. podocarpi Takahashi on Podocarpus spp.
in Japan and Taiwan. The present paper describes its annual cycle in the Sydney
region. We have already described its unusual adhesive vesicles (White and
Carver,. 1972) and its flattened eggs and oviposition behaviour (Carver and
Hales, 1974).

STUDY SITES

Neophyllaphis brimblecombei Carver was studied on four trees of Podocarpus
elatus R. Br. ex Endl. growing in a street in the Sydney suburb of Mosman. The
trees were approximately 10 m tall; collections were taken from the lowest branches
at weekly or fortnightly intervals for one full season, and less frequently at other
times, between 1971 and 1975. Aphids occurring on Podocarpus spinulosus (Sm.)
R. Br. ex Mirb. are considered by Dr. Carver to belong to Neophyllaphis brimble-
combei. P. spinulosus is a small scrambling shrub and occurs naturally in the
bush around Sydney Harbour; some observations were made on populations on
this host, also in Mosman. When found on P. spinulosus, N. brimblecombei is
covered in a coating of long closely packed strands of wax wool, whereas on
P. elatus it is only slightly pulverulent.

RESULTS
OBSERVATIONS ON THE HOST PLANTS

In the mature trees of Podocarpus elatus which were studied, shoots were
produced almost synchronously in August, with only slight differences between
the trees studied. The new leaves, at first pinkish and soft, became pale green
and eventually reached a size of about 9 x 1 cm. Thereafter they began to darken
in colour and became hard and stiff. Hardening and darkening of the leaves was
evident, but incomplete, in mid-October; the leaves were not fully mature and
hardened until late November.

In untouched trees, no further shoots were normally produced until the
following August. Two of the trees studied were periodically pruned to prevent
them touching powerlines and fresh shoots were present on these for long periods.
Accidental damage also induced formation of new shoots out of season.

Although P. elatus is native to the Sydney region (as defined by Beadle e¢ al.,
1972) most specimens in Sydney itself are cultivated as street or garden plants,
and therefore are subject to pruning.

Podocarpus spinulosus commences shooting between late July and late
September, the onset varying between plants in different situations. The leaves

73 Aust. Zool. 19(1), 1976

BIOLOGY OF NEOPHYLLAPHIS BRIMBLECOMBEI

reach a maximum length of 6 cm before hardening. Hardening is completed in
about 10 weeks from the beginning of shooting. Occasional new shoots are
produced in natural conditions in April (mid-autumn). P. spinulosus is not
cultivated and its normal growth cycle is not affected by human activities.
ANNUAL CYCLE

The annual cycle of N. brimblecombei in the study area can be summarised
as follows: On Podocarpus elatus, the fundatrices hatch in late July or early
August; they may survive briefly on old leaves if the new shoots have not
developed, but transfer to new shoots before maturity. Fundatrices are characterised
by a deep purple pigmentation and relatively short antennae. They give birth to
apterae viviparae on the new shoots in the second half of August. Fundatrigeniae
and later apterous generations are cream in colour with patches of purple pigment
around the siphunculi. The population increases rapidly, reaching a peak in late
September or early October. At this stage, before mid-spring, oviparae (which
are always alate) become the dominant morph. Alate viviparae and alate males
were never observed to be numerous but were usually collected in small numbers
in October and November. Eggs were present on maturing leaves from October
onwards and could be collected at any time until hatching. No signs of embryonic
development were noted until shortly before hatching. The aphid population
waned during late October and November, when only isolated specimens were
found. Table 1 shows the occurrence of different morphs on a month-by-month
basis.

TABLE 1
RECORDS OF MORPHS OF N. BRIMBLECOMBEI ON P. ELATUS

al = alate viviparae ap = apterous viviparae
f= tondatrix — = no survey
1971-2 1972-3 1973-4 1974-5 1975-6
SPRING Sept. ap f, ap, 2 ap, 9 — f,ap, al, ¢, 2
Oct. — ape. Co ap, al By 6 -ap, ial (Oud
Nov ape al 9 3 ap — ap, al, 2
(few)
SUMMER Dec. nil few larvae nil ap
only
Jan. — nil nil ==
Feb. — nil —_ nil
AUTUMN Mar. — nil — nil
Apr. _ nil — nil
May nil nil oo —
WINTER June so few ap — nil
July — = — —
Aug. —_— f — f

Aust. Zool. 19(1), 1976 79

DINAH F. HALES

When new shoots were produced on P. elatus out of season in response to
pruning or damage, they were usually colonised by N. brimblecombei. The
succession of morphs in these colonies has not been studied, but in the early stages
of their development they contain apterae and apteriform nymphs. Neither
fundatrices nor alatae have been collected in these situations. However, the
colonies are presumably founded by migrants from locally more favourable host
plants.

The collections from P. spinulosus indicate a similar annual cycle, with some
differences. While fundatrix nymphs were collected in August 1975, the initiation
of colonies sometimes appears to be delayed until October (no specimens were
found in August and September 1973) and this may be correlated with the
variable commencement of shooting of the host plant (see Table 2). The peak
population size and production of sexuales also occurs later than on P. elatus,
so that the aphids are still numerous in November and even December. Small
colonies recur in April or May but do not persist. The aphids are absent in mid-
summer. Specimens on this host retain their deep purple colouration in all genera-
tions, but it is only apparent when the dense wax wool coating is removed.

TABLE 2

RECORDS OF MORPHS OF WN. BRIMBLECOMBEI ON P. SPINULOSUS
(Abbreviations as in Table 1)

1971-2 1972-3 1973-4 1974-5 1975-6
SPRING Sept. — — nil — f, ap, al
Oct. — — — — ap, al, 2
Nov. ap, al — — ap, al, 2, ¢
SUMMER Dec. ap, al, 9 — — g
Jan. — — — —_
Feb. — — — ap
AUTUMN Mar. — — — nil
Apr. a larvae only ao larvae
May ap nil ~ — nil
WINTER June — — — —
July — — — —
Aug. — nil — f

Neophyllaphis brimblecombei is preyed on by Micromus tasmaniae Walker
and Drepanacra humilis McLachlan (Neuroptera: Hemerobiidae), by Les conformis
(Boisduval) and Coelophora inaequalis (F.) (Coleoptera: Coccinellidae) and by
Melangyna viridiceps (Macquart), Simosyrphus grandicornis (Macquart), Beta-
syrphus serarius Wiedemann (Diptera: Syrphidae) and another syrphid which has
not been successfully reared. The syrphid identifications are tentative as the group
is in need of revision. No parasitised Neophyllaphis brimblecombei have been
observed.

80 : Aust. Zool. 19(1), 1976

BIOLOGY OF NEOPHYLLAPHIS BRIMBLECOMBEI

DISCUSSION

ANNUAL CYCLE OF Neophyllaphis brimblecombei COMPARED WITH CYCLES OF
OTHER Neophyllaphis spp.

The annual cycle of Neophyllaphis brimblecombei in the Sydney region is
basically a very short one, with only one or at the most two generations inter-
vening between fundatrix and the first ovipara. Facultative parthenogenetic repro-
duction can continue, however, if suitable young growth is available on the host
plants, as happens occasionally in natural conditions and frequently on cultivated
plants. The fundatrix hatches late in winter (July-August) and from October to
July the major part of the population is in the form of eggs.

Takahashi (1920, 1923) has given brief notes on the biology of Neophyllaphis
podocarpi Takahashi in Japan and Taiwan. As mentioned above, N. podocarpi is
very similar morphologically to N. brimblecombei. In Japan, N. podocarpi produces
sexuales in late spring and they occur together with apterous and alate viviparae
until the end of autumn. In Taiwan, on the other hand, N. podocarpi is
continuously viviparous throughout the year and sexuales are rare. It is evident
that the host plants (P. macrophylla and P. nageia) remain suitable for extended
periods in Japan and continuously in Taiwan. The aphids are said to attack the
young leaves, but it is not stated whether they can survive on mature leaves.
Each apterous sexupara can produce both sexuales and apterous viviparae

(Takahashi, 1923).

Neophyllaphis gingerensis Carver occurs on P. alpina R. Br. in alpine condi-
tions in the Australian Capital Territory (Carver, 1959). It is mostly confined
to young shoots of its host plant, and its season is again a short one. The first
aphids are observed in December, with sexuales and viviparae both present from
January onwards. By May no aphids remain.

The New Zealand species Neophyllaphis totarae Cottier produces sexuales in
spring on P. tofara, but can apparently continue reproducing parthenogenetically
all the year round (Cottier, 1953, p. 24). The present author has collected both
viviparae and sexuales in February in several South Island localities on P. totara,

P. nivalis and P. hallit.

No published information is available on the annual cycles of the other
Podocarpus-feeding species of Neophyllaphis.

Certain common features emerge from the annual cycles outlined above. Most
species produce sexuales soon after the fundatrix generation. Sexuales and partheno-
genetic forms can be produced concurrently, thus allowing facultative viviparous
reproduction to occur in suitable conditions, while still ensuring early deposition
of diapausing eggs. In some cases the aphids are present only for a short time;
in N. brimblecombei the limiting factor appears to be the availability of young
shoots, while in some species climatic factors may be more important.

Aust. Zool. 19(1), 1976 81

DINAH F. HALES

The absence or rarity of sexuales in N. podocarpi in Taiwan is an exception
to the pattern found elsewhere. The loss of ability to produce sexuales after a
long period of parthenogenetic reproduction is well known in the most advanced
aphid sub-family, the Aphidinae (see Lees, 1966), and it is interesting to note a
similar phenomenon in a primitive aphid such as N. podocarpi.

MECHANISM OF INDUCTION OF SEXUALES

Sexual forms in most aphids that have been studied experimentally are
induced by short photoperiods and low temperatures (see Lees, 1966 for
references ). There is usually an obligatory delay or ‘interval timer” which prevents
production of sexuales until a certain time has elapsed from the fundatrix genera-
tion. This has been shown, for example, in Aphis chloris (Wilson, 1938),
Brevicoryne brassicae, Myzus persicae and Sappaphis plantaginea (Bonnemaison,
1951) and Megoura viciae (Lees, 1961), all members of the sub-family Aphidinae.

Dixon (1971, 1972) has demonstrated a similar phenomenon in Drepano-
siphum platanoides and Eucallipteris tiliae, both of which belong to the sub-family
Drepanosiphinae, as does Neophyllaphis. In D. platanoides the obligate delay was
much shorter than in the Aphidinae listed above, and could be overcome completely
by very short experimental photoperiods. With the passage of time from the
fundatrix generation, the critical photoperiod became longer, i.e. the threshold
of response became lower. In E. ftiliae even the fundatrix produces some oviparae
in natural conditions.

The observations reported here on N. brimblecombei are consistent with a
short obligatory delay and a low threshold of photoperiodic response. On the
other hand, nutritional factors have been shown to induce the production of
sexuales in some aphids, e.g. Dysaphis devecta (Forrest, 1970) and also in the
closely related Pemphigidae, e.g. Tetraneura ulmi (Schwartz, 1932), Pemphigus
bursarius (Dunn 1959, 1974) and Eriosoma pyricola (Sethi and Swenson, 1967).
The hypothesis that host plant factors are involved in production of sexuales in
N. brimblecombei is attractive, as it would account for the close correlation
between aphid and host plant cycles.

ADAPTIVE VALUE OF THE Neophyllaphis ANNUAL CYCLE

Aphids have radiated and diversified most strongly in the north temperate
regions, and the number of species endemic to Australia is small. It is rather
surprising that so few north-temperate species have become established in Australia
since the time of European settlement. One might seek the explanation in the
annual cycles of northern aphids, which are specialised to avoid the harsh winter.
A long obligate delay between fundatrix and sexuales, such as occurs in most
holocyclic Aphidinae, forces species to rely on continuous parthenogenetic repro-
duction in summer. Long hot, dry periods, common in most parts of the Australian
mainland in summer, reduce the suitability of host plants and put aphid popula-
tions at risk. The more primitive Neophyllaphis is better adapted to avoid the

82 Aust. Zool. 19(1), 1976

BIOLOGY OF NEOPHYLLAPHIS BRIMBLECOMBEI

rigours of both summer and winter by producing diapausing eggs in the spring.
Other factors related to the control of diapause may also limit the success of
holocyclic strains of northern aphids in Australasia (Eastop, 1966, Lowe, 1973).

Early production of sexuales is characteristic of Sensoriaphis (Drepano-
siphinae) (Carver and Hales, 1974) and Schoutedenia (Greenideinae) (Hales and
Carver, 1976), two other genera associated with Australian native plants. The
apparently native species belonging to the Aphidinae are anholocyclic as far as
is known, and tend to be uncommon compared with Neophyllaphis, Schoutedenia
and Sensoriaphis. The Neophyllaphis type of annual cycle thus appears to have
a high adaptive value in Australian conditions.

ACKNOWLEDGEMENTS

I thank Dr. D. H. Colless, Dr. E. F. Riek and Dr. E. B. Britton of C.S.I.R.O. Division
of Entomology for identifying the predators mentioned in this paper, and Dr. Mary Carver
of the Waite Institute for critical reading of an earlier draft of the manuscript.

REFERENCES

BEADLE, N. C. W., O. D. EVANS & R. C. CAROLIN (1972). Flora of the Sydney
Region. A. H. and A. W. Reed. Sydney.

BONNEMAISON, L. (1951). Contribution a l’étude des facteurs provoquant l’apparition
des formes ailées et sexuées chez les Aphidinae. Annls Epiphyt. 2, 1-380.

CARVER, M. (1959). Two new and possibly indigenous aphids from Australia (Homoptera:
Aphidoidea). Proc. R. ent. Soc. Lond. (B) 28, 19-27.

CARVER, M. & D. HALES (1974). A new species of Sensoriaphis Cottier, 1953 (Homop-
tera: Aphididae) from New South Wales. J. Ent. (B) 42, 113-125.

COTTIER, W. (1953). Aphids of New Zealand. Bull. N.Z. Dept. Scient. Ind. Res., 106 pp.

DIXON, A. F. G. (1971). The “interval timer” and photoperiod in the determination

of parthenogenetic and sexual morphs in the aphid, Drepanosiphum platanoides.
J. Insect Physiol. 17, 251-260,

DIXON, A. F. G. (1972). The “interval timer,” photoperiod and temperature in the seasonal
development of parthenogenetic and sexual morphs in the lime aphid, Eucallipterus
tiliae L. Oecologia (Berl.) 9, 301-310.

DUNN, J. A. (1959). The biology of the lettuce root aphid. Ann. Appl. Biol. 47, 475-491.

DUNN, J.-A. (1974). The influence of the host plant on the production of sexuparae in
the aphid Pemphigus bursarius. Entomologia exp, appl. 17, 445-447.

EASTOP, V. F. (1966). A taxonomic study of Australian Aphidoidea (Homoptera).
Aust. J. Zool. 1/4, 399-592.

EASTOP, V. F. (1970). In The Insects of Australia (C.S.1.R.O., Melbourne University
Press).

FORREST, J. M. S. (1970). The effect of maternal and larval experience on morph
determination in Dysaphis devecta. J. Insect Physiol. 16, 2281-2292.

HALES, D. & M. CARVER (1976). A study of Schoutedenia lutea (van der Goot, 1917)
(Homoptera: Aphididae). Aust. Zool. 19, 84-94.

Aust. Zool. 19(1), 1976 83

DINAH F. HALES

LEES, A. D. (1961). The role of photoperiod and temperature in the determination of
parthenogenetic and sexual forms in the aphid Megoura viciae Buckton II. The
operation of the interval timer in young clones. J. Insect Physiol. 4, 154-175.

LEES, A. D. (1966). The control of polymorphism in aphids. Adv. Insect Physiol. 3,
207-277.

LOWE, A. D. (1973). Aphid biology in New Zealand. In Perspectives in Aphid Biology.
Ent. Soc. N.Z., Bull. No, 2.

SCHWARTZ, H. (1932). Der Chromosomenzyklus von Tetraneura ulmi De Geer. Z.
Zellforsch. 15, 645-687.

SETHI, S. L. & K. G. SWENSON (1967). Formation of sexuparae in the aphid, Eriosoma
pyricola Baker and Davidson, on pear. Entomologia exp. appl. 10, 97-102.

TAKAHASHI, R. (1920). A new genus and species of aphid from Japan. (Hem.). Canad.
Ent. 52, 19-20:

TAKAHASHI, R. (1923). Aphididae of Formosa Part IJ. Rep. Govt. Res. Inst. Formosa 4,
1-173.

WHITE, D. & M. CARVER (1972). Adhesive vesicles in some species of Neophyllaphis
Takahashi, 1920 (Homoptera: Aphididae), J. Aust. ent. Soc. 10, 281-284.

WILSON, F. (1938). Some experiments on the influence of environment upon the forms
of Aphis chloris Koch. (Aphididae). Trans. R. ent. Soc. Lond. 87, 165-180.

84 Aust. Zool. 19(1), 1976

A Study of Schoutedenia lutea (van der Goot, 1917)
(Homoptera: Aphididae)

DINAH F. HALES
School of Biological Sciences, Macquarie University, North Ryde, N.S.W. 2113

and

MARY CARVER
Waite Agricultural Research Institute, University of Adelaide, Glen Osmond, S.A. 5064

ABSTRACT

Observations on a population of the aphid Schoutedenia lutea have been made
over a period of several years. The annual cycle is described. Sexuales occur
concurrently with parthenogenetic females throughout summer and autumn. Some
morphological and taxonomic notes are provided on the fundatrix, a green
viviparous form, the sexuales and the eggs. Schoutedenia viridis is confirmed
as a synonym of S. /Jutea. A number of insect parasites, hyperparasites and
predators of S. /Jutea have been reared and are listed.

INTRODUCTION

Schoutedenia lutea (van der Goot) is a member of a small group (Schou-
tedeniini) of eight known genera in the subfamily Greenideinae, which is
distributed in South America, Africa, India, Australia and in eastern Asia, north-
ward to Japan, and includes two indigenous Australian genera, Avomalaphis Baker
and Meringosiphon Carver.

S. lutea itself occurs naturally in Australia, S.E. Asia and India and probably
Africa and China also, but its true range will not be determinable until the species
composition of Schoutedenia is elucidated. The following species and subspecies of
Schoutedenia have been described: S. ralumensis Rubsaamen, 1905; S. lutea (van
der Goot, 1917); S. viridis (van der Goot, 1917); S. bougainvilleae (Theobald,
1920); S. formosanus (Takahashi, 1929); S. emblica (Patel and Kulkarni, 1952);
S. emblica andhraka David and Hille Ris Lambers, 1956. However, some
synonymy is suspected (Ghosh e¢ al., 1972; Eastop, 1966; Tao, 1962) and it is
likely that the genus actually contains only one or two species.

Schoutedenia has been recorded from woody plants of the family Euphor-
biaceae. The only known host plant of S. Jutea in Australia is Breynia oblongifolia
J. Muell., a small bush growing to 2 m high in or near rainforest and in sandstone

bushland in New South Wales and Queensland, northward to Cape York.

Aust. Zool. 19(1), 1976 85

DINAH F. HALES and MARY CARVER

MATERIALS AND METHODS

All collections were made in sandstone bushland in the Sydney suburb of
Mosman. Observations were made from 1972 to 1975 inclusive, during spring.
summer and autumn. At each inspection, notes were made of the morphs present.
Predators and parasites were collected and reared for identification.

For taxonomic studies, microscopic mounts were made according to the
method recommended by Stroyan (van Emden, 1972). Dissections were carried
out in saline solution and serial sections were prepared from material fixed in
alcoholic Bouin’s solution containing 0.5% trichloracetic acid and stained with
Mayer’s haemalum and celestin blue, or with Regaud’s iron haematoxylin and
phloxin or fast green.

ANNUAL CYCLE

The following account is a composite summary based on three seasons’
observations. Table 1 gives a full month-by-month summary of morphs present.
Fundatrices hatched in mid-September, although fresh shoots were present on the
host plant from the beginning of August. Production of viviparae commenced at
the end of September and, by late October, the first males were observed. There-
after, apterae viviparae, alatae viviparae, oviparae and males were present in the
population until May, except in 1974, when no aphids were observed after
February. Occasional monthly samples after October lacked males and/or alatae
viviparae.

TABEB Ss
COLLECTIONS OF SCHOUTEDENIA LUTEA (VAN DER GOOT)

1972/1973 1973/1974 1974/1975
SPRING Sept. —— Fundatrices —
Oct. _ apt; 6 ¢ nil
Nov. apt; al; 99; ¢¢ apt; al; 2 2 ==
SUMMER Dec. apt; al; 99; 36 apt; al; 2 9 ==
Jan. apt; al; 99; 32 apt; 22; ¢6¢ apt; al; 99; 26
Feb. apt; 2 9 apt; 22; 46 apt; al; 0 Os vere
AUTUMN Mar. apt; al; 9 9 nil apt; als 9 Ossdse
Apr. 10) Gr) OCA) nil apt; al; 99: 464
May apt; al; 2 Q nil apt; al; 9 9
WINTER June — me apt; al
July = — nil
Aug. nil — nil
apt. — apterae viviparae 292 = oviparae
al. = alatae viviparae 66 = males
nil = no live aphids found — = no survey

86 Aust. Zool. 19(1), 1976

A STUDY OF SCHOUTEDENIA LUTEA

Eggs were laid continuously from November to May but presumably did
not hatch until the following spring, as fundatrices were found only in spring.

The aphid was not confined to fresh shoots and could be found on young
leaves, older leaves and on stems of B. oblongifolia. Although the host plant was
common in the area, few individual plants were colonised by S. lutea. On
colonised plants, however, aphid numbers rose to very high levels; over 80
individuals per leaf were counted, corresponding to a density of about 40 cm®. In
late December or early January, alatae viviparae initiated secondary colonies on
neighbouring bushes. These however did not usually reach the size of the primary
colonies. Large populations of aphids appeared to damage the plants; the leaves
turned yellow and were shed.

MORPHOLOGICAL AND TAXONOMIC NOTES
EGe

When freshly laid, the egg is light orange in colour and has a white papilla
at the apex of the posterior (wider) end. Elongated but not flattened and
somewhat irregular in cross section, it might best be described as banana-shaped
(Fig. 1). Length of 20 eggs of S. lutea laid in the laboratory: 0.62-0.73 mm
(mean = 0.68); width: 0.18-0.24 mm (mean = 0.21). Field-laid eggs may be
a little larger. Within a day or so of being laid the eggs turned black. They were
generally Jaid in leaf axils or in cracks in the bark, and were curved to fit the
contours of the substratum. Eggs within macerated oviparae lack the papilla and
instead appear truncate or faintly concave in this area. The papilla is presumably
lost during maceration. Macerated eggs of S. bougainvilleae are similarly truncated
at the posterior end (Eastop, 1961).

—— FF,
0-1 mm x 8
Fig. 1. Egg of Schoutedenia lutea (van der Goot).

Aust. Zool. 19(1), 1976 87

DINAH F. HALES and MARY CARVER

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19(1), 1976

Aust. Zool.

a}eIPSWJ9}UI a)ew ejeje BIeCIAO BIPCIAIA e1ojde xl}yepuny
(OOD Y3qd NVA) VILNT VINJGFLNOHOS 4O SHdHOW 4O NOSIYVdNOD

¢ HTEaVL

A STUDY OF SCHOUTEDENIA LUTEA

Oviposition was observed but had no unusual features. The ovipara stretched
the abdomen posteriorly at the commencement of laying and gradually drew the
tip forwards as the egg was extruded. The ovipara then moved away without
giving further attention to the egg.

FUNDATRIX

Readily identifiable in field; deeper yellow than other apterae viviparae and
with patches of brown on head and dorsum of thorax and abdomen. Antennae
shorter than those of other morphs and only 4-segmented, segment III being
proportionately longer than in other morphs; without secondary rhinaria; flagellar
hairs about 9 »m in length, short, blunt, pale, sparse, i.e. usually no more than
one hair near each rhinarium (in addition to apical hairs). Body pale; dorsal body
hairs 5-8 um long; a little shorter and sparser than in other apterae viviparae;
siphuncular hairs 5-13 »m long; hairs on spinal processi 13 »m long; tergite VII
with two pleural hairs, lateral to spinal processi; tergite VIII with two hairs,
7-19 wm long. Legs proportionately shorter than those of other morphs. Subanal
plate apical, small, pale, spinosely imbricated, with 8-12 short and fine to long
and stout hairs, symmetrically arranged. Subgenital plate pale, spinosely imbricated,
with 4 anteriorly placed hairs and 5-8 posterior hairs. Gonapophyses consist of
4 groups each of 2-4 minute hairs. For further comparison with other morphs,
see Table 2.

APTERA VIVIPARA

Apterae viviparae of S. lutea are usually a clear, uniform lemon-yellow in
colour. Green viviparous specimens were found from time to time intermingled
with yellow ones; on closer inspection, the green specimens proved to be alatoid
individuals with varying degrees of development of wings, thorax, ocelli and
compound eyes. Yellow alatoid apterae also occurred and once (2.11.75), a
collection was made which contained only green apterae viviparae (and darker,
olive-green oviparae). These specimens possessed no alatoid characteristics and
gave birth in the laboratory to the usual yellow apterae. The field colony
contained only yellow specimens when next checked (20.iii.75). A comparison
of these green Schoutedenia with yellow apterae viviparae collected on the same
day from a locality a few kilometres distant showed no significant differences
except perhaps one of size (see Table 2).

OvIPAROUS FEMALE

Apterous. Living specimens olive-green in colour, in contrast to the lemon-
yellow or lighter green colour of the apterae viviparae; orange coloured eggs often
visible through the body wall. Antennae 5-segmented, without secondary rhinaria;
flagellar hairs as in aptera vivipara, 8-12 um long; segment III with only 5 hairs,
segment IV with only 2, near rhinarium; segment V with only 1, near rhinarium
(in addition to apical hairs). Legs with 9-20 pseudosensoria on the hind femur;

Aust. Zool. 19(1), 1976 89

DINAH F. HALES and MARY CARVER

these are difficult to distinguish from the pronounced gland-like ornamentation of
the femora. Body hairs as in aptera vivipara, sparse; on anterior abdominal tergites,
with a maximum of 2 spinal hairs, 8-9 um long, and 2 submarginal hairs on each
side; tergite VII with 2 pleural hairs lateral to spinal processi; hairs on spinal
processi approximately 13 um long; tergite VIII with 2 hairs, 16-43 »m long.
Posterior abdominal segments somewhat attenuated. Subanal plate apical, large,
pale, spinosely imbricated, with 80-90 hairs varying in size from small and fine
to long and stout (cf. that of vivipara with only 10-12 hairs): subgenital plate
large but not obviously demarcated; presence indicated by 2 transverse groups of
variously sized, fine, pale hairs; each group with 30-40 hairs (cf. that of aptera
vivipara, with 2 rows each of usually 6 hairs). Gonapophyses borne on 3
pronounced tubercles, the outer tubercles, 42-64 »m long, both inclined postero-
laterally and each with 4-6 hairs; inner tubercle shorter, 35-50 »m long, some-
times appearing faintly bifid at apex; with 6-8 hairs (cf. viviparae whose gono-
pophyses consist of 4 groups each of 2-5 hairs, which are very rarely borne on

3 very small, flat, pale tubercles). For further comparison with other morphs,
see Table 2.

An internal structure, which persists after maceration, is very conspicuous
in the abdomen of some oviparae. This structure consists of two large, broad,
irregularly shaped, greenish arms which are united posteriorly by a narrow
isthmus. It is also present in oviparoid nymphs, but the arms are not united.
Dissection and serial sections showed that the structure corresponded to the
accessory glands of the reproductive system. The sections also showed that the
accessory glands contain a refractile green substance which is not removed by
ordinary histological solvents. The precise nature and function of this substance
are not known but presumably it forms a protective or adhesive layer on the
surface of the eggs as they are deposited. Dissection of several oviparae showed
four ovarioles on each side of the body cavity. Each ovariole contained one or

Fig. 2. Reproductive system of S. Jutea (ovipara). Abbreviations: s.—spermatheca;
a.g.—accessory gland,

90 Aust. Zool. 19(1), 1976

A STUDY OF SCHOUTEDENIA LUTEA

two large eggs. A diagrammatic representation of the reproductive system is given
in Figure 2.

The presence of pseudosensoria on the hind femora is very unusual, the only
other published instances being S. bougainvilleae and the related Eonaphis
euphorbiae (Eastop, 1961; Quednau, 1964). These observations conflict with the
references to the occurrence of pseudesensoria on the tibiae of oviparae of
S. lutea and S. emblica andhraka (Ghosh eft al., 1972: David and Hille Ris
Lambers, 1956).

MALE

Alate. Antennae proportionately longer than those of other morphs; 5-
segmented, with 47-59 circular secondary rhinaria on antennal segment III, 13-24
on segment IV and 0-2, usually 0, on segment V (compared with 27-37 secondary
rhinaria on segment III and 0-7 on segment IV in alata vivipara); concentrated on
ventral surface. Tibiae and spinal processi spinosely imbricated, as in alata
vivipara. Genitalia comparatively simple, not heavily sclerotized; claspers roughly
trapezoidal in shape and medially acutely triangular; armature not extensive; penis
short, sparsely haired; about 19 hairs at base of genitalia, which are anterior to

or are a continuation of subanal! plate. For further comparison with other morphs,
see Table 2.

On at least two occasions ( 30.iv.73, 8.1.75), a few small, apparently functional
males were collected, possessing varying combinations of apteroid and alatoid
characteristics. The genitalia appeared to be normal, and compound eyes and
ocelli were present but the specimens possessed abortive wings, a variously
developed thorax and a varying and intermediate number of secondary rhinaria
on the oddly proportioned antennal segments. In addition, they were noticeably
smaller than specimens of any other morph studied.

Ghosh e¢ al. (1972) have briefly described the male of S. lutea and David
and Hille Ris Lambers (1956) the apteroid male of S. emblica andhraka.

The males, alatae viviparae and alatiform nymphs were green in colour.

PREDATORS AND PARASITES

The populations of S. lutea were subject to attack by a range of other insects.
Where possible, the predators, parasites and hyperparasites were reared and

identified.

A. Predators

1. Diptera: Syrphidae. Larvae of Simeosyrphus grandicornis (Macquart) and a
second species, probably Melangyna viridiceps (Macquart). Predation by
larvae of these species was heavy, particularly in spring. Diplazon laetatorius
(F.) (Hymenoptera: Ichneumonidae) was reared from S. grandicornis.

Aust. Zool. 19(1), 1976 91

od,

DINAH F. HALES and MARY CARVER

Diptera: Chamaemyiidae. Leucopis sp. (probably undescribed). This species
was very common, the larvae consuming large numbers of S. /utea throughout
the summer. Their passage through a colony was marked by the presence of
dead, brown aphids hanging by their stylets.

Two specimens of Euryischia sp. (Encyrtidae: Eriaporinae) were reared from
two puparia of this predator. Species of this genus are hyperparasites through
parasitic Diptera (Riek, pers. comm. ).

Coleoptera: Coccinellidae. Coelophora inaequalis (F.), Amidellus ementitor
(Blackburn), Lezs conformis (Boisduval). Adults and larvae of these were
observed from time to time. They did not appear to affect well-established
colonies greatly. On one occasion, L. conformis was found to be parasitised
by dipterous larvae, Phalacrotophora sp. (Phoridae: Metopinae) probably
undescribed. Four parasite larvae emerged from the remains of the host pupa,
and pupated separately, alongside their host.

Neuroptera: Hemerobiidae. Micromus tasmaniae Walker. Not common. Sur-
prisingly, no Chrysopidae were observed.

B. Parasites

1. Hymenoptera: Chalcidoidea: Aphelininae. Aphelinus gossypii Timberlake.
Aphelinus gossypii is at times a common parasite, in N.S.W. and South
Australia at least, of many species of Aphidinae. While parasitism of S. lutea
by this species sometimes reached moderate levels (the brownish gut contents
of the parasite larva could be seen through the body wall of the living host),
few mummified aphids and adult Aphelinus were obtained.

HYPERPARASITES PARASITES PREDATORS PARASITES and

HYPERPARASITES
COLEOPTERA - DIPTERA -
COCCINELLIDAE PHORIDAE
Leis conformis Phalacrotophora
/ | Coelophora inaequalis
Amidellus ementitor

HYMENOPTERA — DIPTERA - DIPTERA — HYMENOPTERA —

PLATYGASTRINAE CECIDOMYIIDAE / _ SYRPHIDAE ICHNEUMONIDAE

Trichacis WS Pseudendaphis Simosyrphus grardicornis << diptazon laetatorius

Melangyna “viridiceps”
DIPTERA — HYMENOPTERA —
CHAMAEMYIIDAE ENCYRTIDAE
Schoutedenia Leucopis Euryischia
lutea
HYMENOPTERA — NEUROPTERA -
APHELININAE HEMEROBIIDAE
Aphelinus gossypii Micromus tasmaniae
Fig. 3. Parasites and predators of Schoutedenia lutea.
92 Aust. Zool. 19(1), 1976

A STUDY OF SCHOUTEDENIA LUTEA

2. Diptera: Cecidomyiidae. Pseudendaphis sp. This may be an undescribed
species but its status will be uncertain until there has been a thorough
revision of cecidomyiid endoparasites of aphids (Harris, pers. comm.). The
species was extremely common from January to May as an endoparasite of
S. lutea. The cecidomyiid larvae were in turn parasitised by a minute wasp,
Trichacis sp. (Proctotrupoidea: Platygastrinae). Members of the Platy-
gastrinae are usually parasites of gall-forming Cecidomyiidae. Cecidomyiids
have not previously been recorded as aphid parasites in Australia.

C. Relationships with ants

At least four species of ants were observed from time to time collecting
honeydew from colonies of S. lutea.

The relationships of S. lutea with other insects are shown diagrammatically
in Fig. 3. With the exception of Pseudendaphis sp. specimens of those insects not
identified to species have been deposited in the Australian National Insect Collec-

tion, C/S. RO) Canberra: AiG. FT,

DISCUSSION

The annual cycle of Schoutedenia lutea in the Sydney region can be sum-
marised as follows: fundatrices hatch in early spring (mid-September) and produce
a generation of apterous fundatrigeniae. Adult males were first noted in late
October, and oviparae in mid-November. It is therefore probable that the third
generation can include males, and the following generation oviparae. After this
time, alate and apterous viviparae as well as sexuales occur concurrently until the
onset of the following winter. There is no evidence in the literature to indicate
whether or not the life cycle of S. lutea has similar unusual features in other
parts of its geographical range.

Other species producing sexuales early in the season either as the final morphs
of an abbreviated life cycle or concurrently with viviparae include some Japanese
species of Greenidea, Paratrichosiphum and Eutrichosiphum (Takahashi and Sorin,
1959; Takahashi, 1962), some species of the unrelated Neophyllaphis in Australia,
New Zealand and Japan (see Hales, 1976), and Sensoriaphis furcifera in Australia
(Carver and Hales, 1974).

Eastop (1961) describes the apterae of S. bougainvilleae as “yellow or green”
and according to Hille Ris Lambers (pers. comm.), Schoutedenia collected in Java
were “repeatedly yellow apterae, sometimes yellow apterae with green alatae”’.

The present study has established that S. lutea occurs in two distinct colour
forms, yellow and green. Ghosh ef al. (1972) are of the opinion that S. lutea,
S. viridis and also S. bougainvilleae are synonymous. Tao (1962) lists S. bougain-
villeae and S. formosanus as synonyms of S. viridis and Eastop (1966) says that
S. bougainvilleae may be a synonym of S. viridis. Van der Goot (1917) described

Aust. Zool. 19(1), 1976 93

DINAH F. HALES and MARY CARVER

(as Setaphis) the apterous and alate morphs of two species of Schoutedenia viz.
S. lutea, a yellow aphid, and S. viridis, a green aphid, which were additionally
distinguishable from one another, in both morphs, by a longer third antennal
segment in S. /utea relative to the processus terminalis (4:1 in S. lutea, 3:1 in
S. viridis).

It can be seen from Table 2 that this distinction does not hold for our
specimens and further comparison of our colour forms and apterous and alate
morphs with van der Goot’s two descriptions leads to the conclusion that S. lutea
and S. viridis are synonymous.

ACKNOWLEDGEMENTS

We thank the following people for identification of insects mentioned in this paper:
E. B. Britton and S. Misko (Coccinellidae), E. F. Riek (Hemerobiidae and Hymenoptera,
except Aphelinus), D. H. Colless (Syrphidae, Chamaemyiidae, Phoridae), K. M. Harris
(Pseudendaphis). Miss Betty Thorn drew the illustrations. Mary Carver wishes to acknowledge
the support of the Division of Entomology, C.S.I.R.O., Canberra.

REFERENCES

CARVER, M. & D. HALES (1974). A new species of Sensoriaphis Cottier, 1953 (Homop-
tera: Aphididae) from New South Wales. J. Ent. (B) 42, 113-125,

DAVID, S. K. & D. HILLE RIS LAMBERS (1956). Notes on South Indian aphids. Indian
J. Ent, 18, 41-44.

EASTOP, V. F. (1961). “A Study of the Aphididae (Homoptera) of West Africa”, (British
Museum (Natural History): London).

EASTOP, V. F. (1966). A taxonomic study of Australian Aphidoidea (Homoptera). Aust.
J. Zool. 14, 399-592.

van EMDEN, H. F. ed. (1972). “Aphid Technology” (Academic Press: London).

GHOSH, A. K., M. R. GHOSH & D. N. RAYCHAUDHURI (1972). Studies on the
aphids (Homoptera: Aphididae) from eastern India. XI. Descriptions of hitherto unknown
or newly recorded sexual morphs of some species from West Bengal. Oriental Insects
6, 333-342.

van der GOOT, P. (1917). Zur Kenntnis der Blattlause Java’s. Contr, Faune Indes Néerl.
y CERT ONES

HALES, D. F. (1976). Biology of Neophyllaphis brimblecombei Carver in the Sydney region
(Homoptera: Aphididae), Aust. Zool. 19, 77-83.

QUEDNAU, F. W. (1964). Further notes on the aphid fauna of South Africa (Homoptera,
Aphidoidea). S. Afr. J. Agric. Sci, 7, 659-672.

TAKAHASHI, R. (1923). Aphididae of Formosa—2. Rep. Govt. Res. Inst. Dep. Agric.
Formosa 4, 129 and 153,

TAKAHASHI, R. (1962). Key to genera and species of Greenideini of Japan with descriptions
of a new genus and three species (Homoptera: Aphididae). Trans, Shikoku ent. Soc. 7,
65-73.

TAKAHASHI, R. & M. SORIN (1959). Biology of the aphid, Paratrichosiphum kashicola
(Kurisaki). (Aphididae, Homoptera). Ecology of Insects, Tokyo 7, 129-132.

TAO, C. CHIA-CHU (1962), Aphid Fauna of China. Science Year Book of Taiwan Museum
V, 33-80.

94 Aust. Zool. 19(1), 1976

Observations on Bdellasimilis barwicki, A Marine
Triclad from Australian Freshwaters
(Platyhelminthes: Turbellaria)

IAN R. BALL

Department of Entomology and Invertebrate Zoology, Royal Ontario Museum,
Toronto, Ontario, Canada

ABSTRACT

Bdellasimilis barwicki, an aquatic planarian ectoconsortic on turtles from Aus-
tralian freshwaters, is re-described from type material. Corrective remarks are
made and new taxonomic data provided. The species is here regarded as a
member of the Maricola that has invaded freshwaters and it is assigned to the
family Procerodidae.

INTRODUCTION

Bdellasimilis barwicki is a probursal planarian originally described as an ecto-
consort from the limb-pits of turtles from Lake Wyangan, near Griffith, New South
Wales (Richardson, 1968). Subsequently further specimens were obtained from a
turtle on the flood-plain of the Lower Clarence Valley, Northern Rivers Region,
New South Wales, and there is a possibility that a second species occurs in Queens-
land (Richardson, 1970). There are many unusual morphological features of this
species that made its taxonomic assignment difficult and pointed to the possibility
that the genus may require a new suborder within the Tricladida (Richardson,
1970). Through the kindness of Professor L. R. Richardson I have been able to
examine two paratypes of the species and from my detailed study of these I am
able to correct some aspects of the original description and to provide new data
that enhance our understanding of the systematic position of this animal.

Originally the species was assigned by Richardson (1968) to the Paludicola,
the freshwater planarians, “provisionally and with much reservation”. In my
opinion the species is a true maricole that has invaded freshwaters (Ball, 1974a:
343) a phenomenon that has occurred in some other marine planarians (Ball,
1974b: 29). Kenk (1974) has also expressed doubt as to the paludicolan affinities
of this form.

Aust. Zool. 19(1), 1976 95

IAN R. BALL

MATERIALS AND METHODS

The two paratypes from Professor Richardson were received preserved in
glycerine. They were transferred to alcohol and then embedded in paraffin in the
usual way. Sagittal and transverse serial sections were cut, at a thickness of 8
microns, and the sections were mounted on 2 in x 3 in glass microslides. The
ribbons were arranged in vertical rows with the first section in the upper left
corner when the slide label is to the right. The sections were stained in Mallory-
Heidenhain stain (Gurr, 1963) but the fixation technique used by the original
collector proved not to be entirely compatible with this stain and so the sections
were only weakly coloured.

The two paratypes are now in the collections of the Department of Ento-
mology and Invertebrate Zoology, Royal Ontario Museum, as follows: sagittal
sections on four slides (ROM C418a) and transverse sections on three slides
(ROM C418b). The original type material is a whole-mount in the Australian
Museum (W417c).

SY ST EMATIC SECTION

Infraorder MARICOLA
Family PROCERODIDAE Diesing

Genus Bdellasimilis Richardson, 1968

The assignment of the genus to the Procerodidae may prove to be a temporary
measure. The classification of the Maricola is in a state of flux (Holmquist and
Karling, 1972; Mitchell and Kawakatsu, 1972) and it is more than likely that
our concept of the Procerodidae, a family defined only by primitive characters,
will undergo considerable revision in the near future (e.g. Ball, 1973, 1975).

Bdellasimilis barwicki Richardson, 1968

CORRECTIVE REMARKS

The original description (Richardson, 1968) and subsequent biological notes
(Richardson, 1970) are here supplemented and corrected by my own observations
on the two paratypes.

The eyes, which may be of taxonomic importance in both the Paludicola and
the Maricola (Ball, 1974b, 1975), consist of a single-celled pigment cup containing

about three large retinal cells. Across the opening of the pigment cup there is a
thick corneal structure like that described in some other Procerodidae and in some

96 Aust. Zool. 19(1), 1976

REVISION OF BDELLASIMILIS BARWICKI

Bdellouridae (Bohmig, 1906). I am unable to determine if this is a true lens in
the sense of Lehmensick (1937).

Richardson (1968: 93) found the testes to be a paired tubular system. I
find them to be represented by numerous small round follicles extending from near
the anterior end of the animal, and well anterior to the eyes and ovaries, to the
level of the pharyngeal pore. They are principally ventral in position though
anteriorly they extend over the ovaries.

be gl’ gl” gl”

100 pm

od vd cvd pb ma pp go

Fig. 1. Bdellasimilis barwicki, paratype, ROM C418a. Sagittal section of the copulatory
apparatus, viewed from the left side. bc, bursa copulatrix; cvd, common vas deferens:
gl, atrial glands: go, gonopore; ma, male atrium: od, left oviduct; pb, penis bulb;
pp, penis papilla; vd, left vas deferens.

A sagittal view of the copulatory apparatus was not provided by Richardson;
this is done here as Fig. 1. My interpretation of the sections differs in some
respects from that of the original author (cf. Fig. 1 with Richardson, 1968: Fig. 2).

The penis consists of an elongate weakly muscular bulb, and a long slender
papilla projecting into the atrium. The vasa deferentia unite to a stout common
duct, with thick walls, anteriorly to the penis bulb and it is this duct that enters
the frontal face of the bulb. Within the bulb it forms a small vesicle that narrows
posteriorly to form an ejaculatory duct opening terminally.

Aust. Zool. 19(1), 1976 o7

IAN R. BALL

The bursa copulatrix is a large sacciform organ lying anteriorly to the penis
as above the common vas deferens. The wide and thick-walled bursal canal
passes over the penis bulb and opens into the atrium in an unusually anterior
position. The thin musculature of the bursal canal, which overlies a nucleate
lining epithelium, continues the sequence of layers found in the atrium and the
bodywall; there is no reversal of the circular and longitudinal layers. The two
oviducts, which were not seen by Richardson, are most unusual in that they enter
the lateral walls of the bursa copulatrix.

The single, ventral, gonopore lies between the two posterior adhesive discs.
It opens into a large chamber into the walls of which numerous and extensive fine
glands open. These glands appear as fine yellow-brown granules that extend into
much of the mesenchyme (Fig. 1 gl’). Entally to these the atrium narrows to
form a tubular duct and into this zone there open numerous coarser glands which
stain purple (Fig. 1 gl”). The ental part of the atrium, surrounding the penis, is
larger once more and its walls are pierced by very coarse and very extensive
eosinophil glands (Fig. 1 gl’). It is my belief that Richardson (1968) mistook
this glandular complex for muscle tissue, which also may be eosinophil in nature,
and thus erred in his interpretation of the penis bulb and atrium.

These corrections to the original description make necessary a new diagnosis
of the genus.

DIAGNOSIS OF THE GENUS Bdellasimilis

Translucent marine planarians, ectoconsortic on turtles, with long pre-ocellar
region and broad posterior end with two adhesive discs. Eyes two, with single-
celled pigment cup with lens, and three retinal cells. Bursa copulatrix anterior;
oviducts enter the lateral walls of the bursa. Ovaries behind the eyes. Testes
numerous, and ventral, and extending posteriorly to the level of the single
gonopore. Penis with a weak bulb and slender papilla extending into a spacious
and highly glandular atrium. Common vas deferens enters the penis bulb. Cocoons
slender, ovoid or cylindroid, and with a slender stalk. Type and only known species:
Bdellasimilis barwicki Richardson, 1968.

COMPARATIVE DISCUSSION

The eye-structure of Bdellasimilis is quite unlike that of the Dugesiidae (Ball,
1974b, 1974c), the sole family to which the paludicolan planarians of Australasia
belong, for in this family the multi-cellular pigment cup embraces numerous small
retinal cells. In contrast the single-celled pigment cup, containing three large
retinal cells, of Bdellasimilis is similar to that of many of the Maricola. This,
together with the facts that there is a long, diverticulated pre-ocellar duct of the
anterior ramus of the intestine, a primitive feature found only in the Maricola

98 Aust. Zool. 19(1), 1976

REVISION OF BDELLASIMILIS BARWICKI

(Meixner, 1928), and many testes anterior to the ovaries, strongly suggests that
B. barwicki cannot be a paludicolan triclad, notwithstanding the fact that it is
probursal. In fact, the small size and large number of the testes are reminiscent
of the condition found in species of the maricolan families Bdellouridae and
Micropharyngidae (Bohmig, 1906; Wilhelmi, 1909) and they are unlike those of
the Australasian Dugesiidae that I have examined. The probursal condition is
unusual because in most Maricola the bursa is behind the penis, but there is one
marine family, the Probursidae of Hyman (1944), in which the probursal condition
is present and presumably this is an independent acquisition in the two groups.

Most of the unusual structures of Bdellasimilis are paralleled by similar features
in other marine planarians; whether the similarities are homologous or analogous
cannot yet be stated. The extremely long atrium is also found in Micropharynx
parasitica and Nexilis epichitonius, and Nexilis also possess an unusually ental
opening of the bursal canal into the atrium (Ball, 1975). Interestingly enough,
both these species, like B. barwicki, live in close association with higher aquatic
organisms for N. epichitonius is usually ectocommensal on chitons and M. parasitica
is ectoparasitic on skates in the North Atlantic Ocean ( Jagerskiold, 1896; Jennings,
1971; Ball and Khan, 1976). M. parasitica shows additional similarities with B.
barwicki because it, too, has a long common vas deferens and the oviducts may
open into a small vesicle that may be homologous with the bursa copulatrix of
other Maricola (Ball and Khan, 1976). Apart from Bdellasimilis, and to the best
of my knowledge, Micaplana misae, a marine planarian from Japan, is the only
other aquatic planarian in which there is a well developed bursa into which the
oviducts open (Kato, 1937).

The extreme glandular nature of the atrium of B. barwicki is without parallel
in the Maricola. In the Paludicola, however, there seems to be a similar glandular
complex within species of the genus Cura, particularly in the North American
species C. foremanii, as described by Kenk (1935), and in the Australasian species
C. pinguis (Weiss, 1910). In both these species the bursal canal opens into the
atrium far more anteriorly than is usual in freshwater planarians and its terminal
part, beneath the oviducal openings, is expanded to form a wide female atrium
(Weiss, 1910: Fig. 32; Kenk, 1935: Fig. 15). The extensive shell glands open
into this chamber. The posterior part of the atrium, leading to the gonopore, is
quite extensive and its walls receive the openings of large numbers of glands. In
C. foremanii Kenk recognised two sorts of glands, whereas Weiss (1910) refers
simply to eosinophil glands in C. pinguis. But from my own slides of these two
species I can confirm that in both there are two types of atrial glands. Those
surrounding the gonopore stain brownish-yellow in Mallory-Heidenhain, and the
more ental ones are coarser and eosinophil in reaction. The similarity of this

Aust. Zool. 19(1), 1976 99

IAN R. BALL

arrangement to that of B. barwicki is quite striking for if the ental atrial glands
(el’) of B. barwicki are homologous with the shell glands of the Cura species,
then the remaining two sets of glands (gl, gl’) are identical with respect to
position and structure with the two types of atrial glands in the Cura species.
The origins of the freshwater planarians from their marine ancestors are obscure
(Bail, 1974a) but it is interesting to note that Cura is generally considered to be
one of the most primitive genera (Ball, 1974b) and is therefore most likely to
show retention of maricole characters.

For the above-discussed reasons there can be little doubt that Bdellasimilis is
a true marine planarian that has invaded freshwaters. A final analysis of its
phylogenetic relationships within the Maricola must await a much-needed revision
of the entire suborder something that has not been undertaken since the work

of Wilhelmi (1909).

ACKNOWLEDGEMENTS

It is a pleasure to thank Professor L. R. Richardson, Grafton, N.S.W., for his generosity
in donating two paratypes to me and for his interest in this study. I am also grateful to
Maria Tran Thi Vinh-Hao for drawing Figure 1.

REFERENCES

BALL, |. R. (1973). On the anatomy and affinities of Nesion arcticum Hyman, a marine
triclad from Alaska (Platyhelminthes: Turbellaria). Can. J. Zool. 57, 1299-1305.

BALL, I. R. (1974a). A contribution to the phylogeny and biogeography of the freshwater
triclads (Platyhelminthes: Turbellaria). Jn ‘Biology of the Turbellaria’. (Eds. N. W.
Riser and M. P. Morse), pp. 339-401. (McGraw-Hill Book Company: New York.)

BALL, I. R. (1974b). A new genus of freshwater triclad from Tasmania with reviews of
the related genera Cura and Neppia (Turbellaria, Tricladida), Life Sci. Contr., R.
Ont. Mus. 99, 1-48.

BALL, I. R. (1974c). A new genus and species of freshwater planarian from Australia
(Platyhelminthes: Turbellaria). J. Zool., Lond. 174, 149-158.

BALL, I. R. (1975). Contributions to a revision of the marine triclads of North America:
the monotypic genera Nevilis, Nesion, and Foviella (Turbellaria: Tricladida). Can. J.
Zool, 53, 395-407.

BALL, I. R. & R. A. KHAN (1976). On Micropharynx parasitica Jagerskidld, a marine
planarian ectoparasitic on thorny skate, Raja radiata Donovan, from the North Atlantic
Ocean. J. Fish. Biol. 8 (in press).

BOHMIG, L. (1906). Tricladenstudien I. Tricladida Maricola. Z. wiss. Zool. 81, 344-504.

GURR, G. T. (1963). ‘Biological Staining Methods.’ (G. T. Gurr Ltd.: London.)

HOLMQUIST, C. & T. G. KARLING (1972). Two new species of interstitial marine triclads
from the North American Pacific coast, with comments on evolutionary trends and
systematics in Tricladida (Turbellaria). Zool. Scr. J, 175-184.

JAGERSKIOLD, L. A. (1896). Uber Micropharynx parasitica n.g., n. sp. Eine ectopara-
sitische Triclade. Ofv. Vet. Akad. Forhand. Stockholm 53(10), 707-714.

100 Aust. Zool. 19(1), 1976

REVISION OF BDELLASIMILIS BARWICK]

JENNINGS, J. B. (1971). Parasitism and Commensalism in Turbellaria. Adv. Parasitol. 9,
1-32.

KATO, K. (1937). A new marine triclad from Japan. Annotnes Zool. Japon. /6, 28-33.

KENK, R. (1935). Studies on Virginian triclads. J. Elisha Mitchell Scient. Soc, 51, 79-133.

KENK, R. (1974). Index of the genera and species of the freshwater triclads (Turbellaria)
of the world. Smithsonian Contr. Zool. /83, 1-90.

LEHMENSICK, R. (1937). Morphologie und Histologie einer neuen Meerestriclade
(Procerodes Harmsi n. sp.) mit Linsenaugen. Z. wiss. Zool. 149, 131-160.

MEIXNER, J. (1928). Der Genital apparat der Tricladen und seine Beziehungen zu ihrer
allgemeinen Morphologie, Phylogenie. Okologie und Verbreitung. Z. Morphol. Oekol.
Tiere. 11, 570-612.

MITCHELL, R. W. & M. KAWAKATSU (1972). A new family. genus, and species of
cave-adapted planarian from Mexico (Turbellaria. Tricladida. Maricola). Occ. Pap.
Mus. Tex. Tech. Univ. 8, 1-16.

RICHARDSON, L. R. (1968). A new bdellourid-like triclad turbellarian ectoconsortic on
Murray River Chelonia. Proc. Linn. Soc. N.S.W. 93, 90-97,

RICHARDSON, L. R. (1970). A note on Bdellasimilis barwicki and an indication of a
second species (Turbellaria: Tricladida). Aust. Zool. 15, 400-402.

WEISS, A. (1910). Beitrage zur Kenntniss der australischen Turbellarien I. Tricladen. Z.
wiss. Zool. 94, 541-604.

WILHELMI. J. (1909). Tricladen. Fauna und Flora des Golfes Neapel 32. (R. Friedlander:
Berlin. )

Aust. Zool. 19(1), 1976 101

psy A

Bibs Rien nae

Small Mammals Trapped in Dorrigo National Park,
New South Wales

A. B. ROSE

National Parks and Wildlife Service, 24 Fisher Avenue, Wahroonga,
New South Wales, 2076

Small mammal trapping was carried out in Dorrigo National Park for the
first time in April 1975. Elliott live traps and a few wire cage traps were baited
with a mixture of oats, peanut butter, and bacon grease. Three distinct habitats
were sampled:

A. Cool temperate rainforest regrowth, altitude 731 m.
B. Dense canopied subtropical rainforest, altitude 609 m.
C. Alluvial subtropical palm forest, altitude 76 m.

Some specimens were taken for the preparation of study skins and skulls
and are deposited in the reference collection of the N.S.W. National Parks and
Wildlife Service, Mount Colah Depot.

TABLE 1

Mammal captures. Animals indicated with an asterisk (*) were captured in wire cage traps.
All others were captured in Elliott live traps.

Trap Melomys Rattus Ratus Mus Antechinus Trichosurus Felis
Site nights cervinipes fuscipes rattus musculus — stuartii caninus catus
A 294 8 10 — 1 7 — —
B 253 5 39 1 — 8 2 i=
C 144 4 15 1 1 i — —
Total 691 17 64 2 2 16 fu 1

The results are shown in Table 1. Captures of Rattus fuscipes in habitat A
(0.034 captures per trap night) were significantly, at the 1% level, lower than
in habitat C (0.104 captures per trap night) and habitat B (0.154).

Identification of Melomys cervinipes on external characteristics was difficult
due to general similarities with R. fuscipes. However, female M. cervinipes could
be distinguished on the basis of nipple count (Table 2), and the body weight of

Aust. Zool. 19(1), 1976 103

A. B. ROSE

TABLE 2
Comparative data from Melomys cervinipes and Rattus fuscipes.
Body wt (gm) Mammary formula
Species Area (mean = s.d.) (thoracic & inguinal )
M. cervinipes Dorrigo VS ei/ Seas ebay 8) (Ent == SU) (= —
R. fuscipes Dorrigo 13433) = 23.9 (1 = 59) 5:28
R. fuscipes Ku-ring-gai 93935 == 28.6 (m= 51) 2-3

this species was much lower than that of R. fuscipes trapped in the same area.
However, as shown in Table 2, this relationship might not hold throughout the
range of these species, as the mean body weight of bush rats from Ku-ring-gai
(Sydney region) is closer to that obtained for M. cervinipes in this study.
Members of the genus Melomys are frequently called ‘mosaic tailed rats’, but this
character was not readily apparent in the Dorrigo population. The scales on the
tail appeared to be arranged in rings (similar to the bush rat), and only upon
close examination of the base of the tail of prepared study skins could a clear
mosaic pattern be seen. There was some difference in the arrangement of hairs on
the tail. M. cervinipes appeared to have a naked tail on first sight, but close
examination disclosed short appressed hairs no longer than the scales. R. fuscipes
had longer hairs emerging from under the scale rings. Further details for individuals
of these two species collected at Dorrigo are given in Table 3. Colour is omitted,
as there was considerable overlap in general coat colour. Most Dorrigo R. fuscipes
did however have white patches of fur on the ventral surface. M. cervinipes

did not.

TABLE 3
Measurements taken prior to preparation of study skins from Dorrigo rodents.
M. cervinipes R. fuscipes
Sex EAS ue Re ee Re RE F M Juv. F M Juv.
Weight (gm) A ok en SM 60 71 40 150 138 50
Head and: body (Gmim).. 3...- 2. 122 128 106 162 169 123
Tail Eee Aus Pleiae ALTRI ta 152 166 126 my 152 13
Hindfoot without claw vee en 27 DiS 26 Sled 203 30.9

Ear PRvEtT. .Rateeen Cote awe cee 20.4 20 17 Doeao eos 19.6

A further characteristic that might be useful in distinguishing between these
two rodent species in the field is the ready inclination of M. cervinipes to climb.
A typical animal, when released from the trap, climbed 12 m up a sapling and

104 ‘Aust. Zool. 19(1), 1976

MAMMALS TRAPPED IN DORRIGO NATIONAL PARK

disappeared. R. fuscipes on the other hand could not be induced to climb,
preferring in all cases observed to jump to the ground and run off.

ACKNOWLEDGEMENTS

I wish to thank J. Mahoney (Sydney University) for help in identifying the specimens
collected, and the Dorrigo Park Trust for allowing Chief Ranger N, Fenton and Ranger I.
Archibald to assist; their help in setting the trap lines was much appreciated. This paper
would not have been prepared without the encouragement and assistance of Dr. M. Augee
(University of N.S.W.) to whom I am extremely grateful.

Aust. Zool. 19(1), 1976 105

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An Occurrence of Albino Frog Tadpoles in
New South Wales

NORMAN GRADWELL
7 Centennial Avenue, Lane Cove, N.S.W. 2066.

In a review of albinism in amphibians, Noble (1931) states: ‘The albino
axoltl is known to differ from the normal colored phase merely by a single
Mendelian factor (Haecker, 1908), although there appear to be various degrees
of albinism in this species which may be due to other genetic factors.” More
recently, Browder (1972) has discussed the literature relevant to the genetics of
albinism in the Anura. However, there appear to have been no publications on
the occurrence in nature of albinism among Australian anurans. The present report
is on the finding and laboratory development of tadpoles of Limnodynastes, of
which about half were albinos. Although no genetic data are presented, some
references on this aspect of albinism are discussed.

On 7 October, 1974, a small population of both albino and normally pigmented
frog tadpoles (Fig. 1A, B, C) were found near the golf links at La Perouse, ca.
8 km south of Sydney, N.S.W. A few normally pigmented adult Limnodynastes
frogs were seen, but there were no albino adults. Normally pigmented tadpoles
of Ranidella signifera were plentiful. The habitat consisted of muddy pools (ca.
0.5 m maximum depth) surrounded by patches of grass and wattle. Scattered
refuse such as motor car chassis, tyres, and corrugated iron roofing, was also
present. A random collection with dip nets yielded 27 albinos and 18 normally
pigmented tadpoles of Limnodynastes. Both normal and albino tadpoles were of
about the same size at corresponding stages of development (Table 1).

TABLE 1

Length in mm

Tadpole Entire Snout-to-vent Stages of Gosner, 1960
Melanated a7. I BUY ects (352 27—35
Albino 44,5: 12:3 19.2 + 4.0 26—35

Aust. Zool. 19(1), 1976 | 107

NORMAN GRADWELL

In dissection, the albinos were found to contain no melanin in their blood
vessels, in their retinas, or anywhere else. Other slight anatomical differences
between the two kinds of tadpole were also found. For example, the albinos had
a wider median gap in the posterior tooth row of the upper jaw, than the normal
pigmented tadpoles (Fig. 1D, E). These differences were enhanced during meta-
morphosis (Fig. 1C) by the appearance of patterns of distinctive non-melanated
pigment. However, the beaks and teeth of the albinos were black like those of
normally pigmented tadpoles (Fig. 1D, E). Both kinds of tadpole were identified
by their mouth parts as belonging to the genus Limnodynastes.

The bioreaction whereby melanin is produced has been summarised by Bell.
Davidson, and Scarborough (1950). In the skin of several pigmented Amphibia,
as in many other animal tissues, this reaction is catalysed by tyrosinase, but as
Smith-Gill, Richards and Nace (1972) point out, albinism is not dependent on
tyrosinase function alone. As this enzyme, like those associated with certain other
genes, may become active only at specific stages of ontogeny, it was decided to
determine whether albinism is restricted to the tadpole stage of the life history.
In addition, Smith-Gill, Richards and Nace report on the failure of certain albino
tadpoles of Rana pipiens to metamorphose.

The above findings encouraged the laboratory rearing and metamorphosis of
the La Perouse tadpoles. Ten melanated and 10 albino tadpoles were kept in
aerated aquaria containing Sydney municipal tap water at 20°C, and fed on boiled
spinach for three to four weeks. They were illuminated with subdued incandescent
light. To investigate the possibility of water-borne chemical interactions between
the two kinds of tadpole, five albino and five melanated tadpoles were placed in
one aquarium, and five tadpoles of each kind were placed separately in two other
aquaria.

The albino tadpoles all became frogs of Limnodynastes tasmaniensis and the
melanated tadpoles all became frogs of Limnodynastes peroni (Fig. 1F, G). How-
ever, from stage 42 onward (Fig. 1C, G), yellow-orange and white pigment
appeared on the dorsum of only the albino animals. These pigments occurred in
areas corresponding to the brown-orange and pale areas respectively of normally
pigmented frogs of L. tasmaniensis.

Fig. 1. A. Albino tadpole of Limnodynastes tasmaniensis. B. Normal pigmented tadpole of
L. peroni. C. Metamorphosing tadpole of L. tasmaniensis. D. Mouth parts of an
albino tadpole of L. tasmaniensis. E. Mouth parts of a melanated tadpole of L. peroni.
F. Young albino adult of L. tasmaniensis; slightly over-exposed photograph to
emphasise the area of yellow-orange pigment bordered by white. G. Young normal
pigmented adult of L. peroni.

108 _ Aust. Zool. 19(1), 1976

OCCURRENCE OF ALBINO FROG TADPOLES

el

Aust. Zool. 19(1), 1976 109

NORMAN GRADWELL

The present research has shown that under laboratory conditions the natural
occurrence of albinism in certain tadpoles of La Perouse persists into the frog
stage of the life history. However, the immaturity of the newly metamorphosed
frogs precluded sexing them by dissection and it could not be established if albinism
is a sex-linked character in these forms. Another finding concerns the fact that
keratin itself is not black, even in darkly pigmented mammals, and therefore it is
of interest to note that the keratinised beaks and teeth of all the albino tadpoles
were black. In addition, there appears to be no melanin-influencing chemical
interaction between the normally pigmented and the albino tadpoles during late
larval growth and metamorphosis. Neither did melanin appear in the young albino
frogs fed for three to five days on insects. However, as Browder (1972) points
out, amelanotic melanophores may still be present in albinos. Finally, the failure
to find albino frogs at La Perouse need not indicate a lethal genetic association
between albinism and the assumption of life as a frog. It is more likely that poor
eyesight in the albinos and their conspicuousness to predators, reduces viability. In
this respect, Browder reports that even in laboratory conditions free from predators,
the albinos of Rana pipiens are less viable than their melanated siblings.

Apart from melanation, no significant taxonomic differences were found
between albino and normally pigmented tadpoles of L. tasmaniensis (from Sydney
and Oberon districts ).

A representative collection of the two kinds of tadpole and the frogs obtained
by laboratory metamorphosis has been lodged at the Australian Museum, Sydney
(registration numbers R47257 to R47278). The collection consists of 12 albino
tadpoles, 10 melanated tadpoles, 11 albino frogs, and 8 melated frogs.

ACKNOWLEDGEMENTS

1 am grateful to Mrs. G. Goidsack for drawing my attention to her chance discovery
of the tadpoles. I also thank Dr. H. Cogger for his assistance and Mr. M. J. Tyler for
reading the manuscript and identifying and attempting to sex the frogs. This research has
been supported by a grant from the Australian Biological Resources Study Interim Council.

REFERENCES

BELL, G. H., J. N. DAVIDSON & H. SCARBOROUGH (1961). Textbook of physiology
and biochemistry. E, & S. Livingstone Ltd., London.

BROWDER, L. W. (1972). Genetic and embryological studies of albinism in Rana pipiens.
J. Exp. Zool. 180, 149-156.

GOSNER, K. L. (1960). A simplified table for staging anuran embryos and larvae with
notes on identification, Herpetologica 16, 180-190.

HAECKER, V. (1908). Ueber axolotlkreuzungen; II. Mitteilung (zur Kenntnis des partiellen
Albisimus). Verh. deutsch. zool. Ges. 18. Vers., 194-205. (Quoted from Noble, 1931.)

NOBLE, G. K. (1931). The Biology of the Amphibia. McGraw-Hill, N.Y. Reprinted 1954.
Dover Publ., N.Y.

SMITH-GILL, S. J., C. M. RICHARDS & G. W. NACE (1972). Genetic and metabolic
bases of two “albino” phenotypes in the leopard frog, Rana pipiens. J. Exp. Zool. 180,
157-168.

110 Aust. Zool. 19(1), 1976

Memorial Notice

GILBERT PERCY WHITLEY, F.R.Z.S. (1903-1975)
Photograph by courtesy of Dr. J. R. Paxton

Mr. Gilbert Whitley died in Sydney after a short illness in July, 1975.
Details of various facets of his life and work have been published elsewhere and
need not be repeated here. The opportunity remains, however, for this journal
to record the extent of the contributions made by Mr. Whitley to the Royal
Zoological Society of New South Wales.

A brief review of Mr. Whitley’s association with the Society presents a
picture of extraordinary service extending over half a century. He was a Member
of the Society from 1926 until his death, a Life Member from 1938 and a long-
time Fellow. He served as a member of Council for many years and was
President or Vice-President on several occasions. A gentle, knowledgeable,
industrious mah with a charming sense of humour, he was highly esteemed as
a Council Member whose views always deserved consideration.

Aust. Zool. 19(1), 1976 111

MEMORIAL NOTICE

The contributions made by Gilbert Whitley to the publications of the
Society are unlikely to be equalled. His first paper in The Australian Zoologist
(on parasitism in lower animals) appeared in 1925 in Volume 3, Part 8, and
almost every issue since then (including the present one) contains one or more
of his articles. The name of G. P. Whitley first appears in The Proceedings of the
Royal Zoological Society of N.S.W. in 1936 and again he was a regular contributor
to subsequent volumes. Many of his original scientific papers deal with fishes,
but the remainder cover a wide variety of animal groups, as one would expect
from a naturalist with such broad zoological interests. Other articles, such as
book reviews, biographies and historical notes also appeared frequently.

In 1947 Gilbert Whitley became editor of both The Zoologist and The
Proceedings. He edited all issues of The Zoologist from Volume 11, Part 2, to
Volume 16, Part 1 (except for 13(2)) and all volumes of The Proceedings until
1970. In all these issues, a painstaking attention to detail and an editorial
precision of professional standard is clearly evident. His editorial as well as his
literary talents are also seen in the Handbooks he produced on scientific and
historical subjects.

The Society has lost one of its most distinguished supporters whose level
of scholarship and productivity were quite exceptional. Many members have felt
the loss of a colleague and a friend.

ES Re

112 Aust. Zool. 19(1), 1976

Erratum

Vol. 18(2), 1974. In the article by P. A. Hutchings and M. F. Recher
entitled “The fauna of Careel Bay with comments on the ecology of mangrove and
sea-grass communities”, Figure 8 was split into Figures 8 and 9 and the correct
Figure 9 was not included.

Figure 8 should be a composite of the published Figure 8 (top) and the
published Figure 9 (bottom).

Figure 9 should be the following:

NUMBER OF INDIVIDUALS
=

42 44 46 48 50 52 54 56 58 60 62 64 66 68 70

SIZE (mm)

Fig. 9. Size distribution of the Sydney Cockle Anadara trapezia at Careel Bay.

Aust. Zool. 19(1), 1976 113

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ROYAL ZOOLOGICAL SOCIETY OF NEW SOUTH WALES
Established 1879
Registered under the Companies Act, 1961

Patron

His Excellency the Governor of New South Wales, Sir Arthur Roden Cutler,
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-COUNCIL, 1976

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Honorary Secretary: Thomas J. Bergin, B.Sc., B.V.Sc., M.R.C.V.S.
Asst. Hon. Sec.: Mrs. O. Wills, F.R.Z.S.
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Honorary Editors: Edward Stanley Robinson, M.Sc., Ph.D
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MEMBERS OF COUNCIL

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F.Z.S., F.R.M.S.

Members of Publications Committee: G. C. Grigg, R. Strahan and E. S. Robinson

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should be addressed to the Honorary Secretary, Royal Zoological Society of
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THE AUSTRALIAN ZOOLOGIST

VOLUME 19 1976 PART |
Contents
| Page
Studies of an island population of Antechinus minimus (Marsupialia,
Dasyuridae): J.-M. WATNER fo eine cpceortectectee citrate ol 1
Maturational and seasonal moult in the New Holland mouse, Pseudomys
navaehollandiae. CG. Mi. KE MPIER Oijesccccscciecscssscossstonrscosseseticasecneeesesscee 9
Relative efficiency of two types of small mammal trap in eastern Australia.
B. J. POXand TG, POSAMEN TEER (octet ctisteaie an i

Mammals of Clouds Creek, north-eastern New South Wales, and their
distribution in pine and native forests. J. L. BARNETT, R. A. HOW

and W2-F. "HUMPHREY Sia. iistsccssccdieoscediedeoten ncccinettmaneh tee 23
Aggressiveness, body weight and injuries in long-haired rats (Rattus

villosissimus ). RE JRBEGG aecctithe tate RB acca iene 35
More fish geneta scrutinized. The late Gi2Ps- WERITEEY 2..tissticscccemmereeneons 45
A redescription of Heteroclinus adelaidae Castelnau (Pisces: Clinidae),

with desctiption of a related new species. D. F. HOESE .......oyaeyaueaae 51
Mosquitoes feeding on ectothermic vertebrates: a review and new data.

HH HEAT WOLE. afd’ R. SHUENE cit cide thcrat tenga tes tee 69
Biology of Neophyllaphis brimblecombei Carver (Homoptera: Aphididae)

im the Sydney region, De FP AES erie isetienceencteneiere Fé
A study of Schoutedenia lutea (van der Goot, 1917) (Homoptera:

Aphididae). D. F, HALES and M, CARVER i.ccccgtore-en:ene i 85
Observations on Bdellasimilis barwicki, a marine triclad from Australian

freshwaters (Platyhelminthes: Turbellaria). IT. R. BAL wn ccessesnesnsen 95

Short papers — .

Small mammals trapped in Dorrigo National Park, New South Wales.
PB ROSE ie ce cee Sancta co scree ne eee ca et 103

An occurrence of albino frog tadpoles in New South Wales. N. GRADWELL 107

Memorial notice —
Gilbert Percy Whitley, F.R.Z.S. (1903-1975 ) -esccsssccnsesnesreracenesenceranetinterarsnnesanesntes 111

FerGtyge dinars hehe eects ec

SURREY BEATTY & SONS
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THE
AUSTRALIAN
ZOOLOGIST

Volume 19, Part 2
September, 1977

Scientific Journal of

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Price $5.00

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THE AUSTRALIAN ZOOLOGIST

VOLUME 19 1977 PART 2

Maintenance of the Platypus (Ornithorhynchus anatinus)
in Captivity Under Laboratory Conditions

T. R. GRANT, R. WILLIAMS and F. N. CARRICK

School of Zoology, University of New South Wales, P.O. Box 1 Kensington,
New South Wales, 2033

ABSTRACT

Platypus were kept in captivity during studies of their temperature regulation
and reproductive biology. They were housed in a complex consisting of a tank,
a tunnel and three nest boxes, and were fed on various live and dead invertebrate
and vertebrate species. Details of the dimensions of the housing facilities and
descriptions of the diet provided are outlined, and general considerations on the
maintenance of this species for research purposes are discussed.

INTRODUCTION

Burrell (1927) described the attempts of the early naturalists at keeping
the platypus in captivity. All of these attempts were notable failures due to a
lack of understanding of the animals’ requirements. Bread, milk, and rice were
common in the diets offered to the animals which were kept as house pets, in
the absence of water! Burrell himself had little success with the species in captivity,
although he made quite an effort to provide his captives with suitable food
(worms, fresh water shrimps, insect larvae and pond snails) and living conditions.
He maintained that “the platypus can be kept alive in captivity; and it is very
probable that with sufficient pain and interest, it can be satisfactorily exhibited”.

Sufficient “pain and interest” have been taken and the platypus has been
successfully kept and exhibited in captivity since early this century (Table 1,
modified from Collins, 1973). At present the Sir Colin MacKenzie Sanctuary in
Victoria, David Fleay’s Fauna Sanctuary in Queensland, Taronga Zoological Park
in Sydney and the Australian Reptile Park at Gosford, N.S.W. all display
Ornithorhynchus anatinus.

These animals, however, have been kept mainly for exhibition purposes and
until the initiation of studies of the thermal biology and reproduction of the
platypus at the University of New South Wales, the difficulties of handling the
species in a laboratory situation were thought to be insurmountable. In 1972
the Director of Targona Park Zoo, Mr. Ron Strahan, conveyed his opinion of the
impossibility of laboratory studies on the platypus. Smyth (1970) tried to maintain

Aust. Zool. 19(2), 1977 117

T. R. GRANT, R. WILLIAMS and F. N. CARRICK

TABLE 1

LONGEVITY DATA FOR EIGHT ORNITHORHYNCHUS ANATINUS IN
CAPTIVITY (MODIFIED FROM COLLINS, 1973)

Sex Dates Captivity Institution
l M 1939-1956 17y Om Sir Colin Mackenzie
Sanctuary
2 F 1943-1955 12y Om
(escaped )
3 F 1938-1947 9y Om Sh
4 M 1933-1937 4y 1m :
5 M 1946-1957 lly 4m ie
and Bronx Zoo
6 F 1946-1957 bby? 3m Bronx Zoo
(escaped )
fi F — 4y lim Melbourne Zoo
8 — 1913-1917 4y 3m Budapest Zoo

platypus in the animal house at the Department of Zoology at the Australian
National University, but eventually in his experiments he had to use animals
taken immediately from the wild, after losing four within 7-10 days after bringing
them into captivity.

During the present studies, commenced in 1972, three platypus died in
captivity. Seven animals were kept and used in experiments for periods of time
ranging from three’ months to slightly under a year. Six of these seven animals
were subsequently released. Hopefully the experience gained during the
maintenance of these animals will be of benefit to other workers, and more
particularly be of benefit to Ornithorhynchus anatinus in captivity.

MATERIALS AND METHODS
HOUSING

The first successful enclosures used for platypus by Eadie (1934) and
Fleay (1944) were quite large; attempts being made to imitate a “natural”
situation. Fleay’s platypusary was 90 feet long and included an artificial bank,
into which the animals could burrow. More recently constructed platypus displays,
such as that at the Targona Zoological Park have also proved to be successful,
despite their less elaborate design. Concrete maintenance tanks, wooden sleeping
boxes and glass sided display tanks have increased the practicability of keeping the
species in an artificial environment. However, even this type of facility was beyond
the scope of the resources available and the size of the installation is not feasible
in most research institutions (the feeding tank alone at the Taronga Zoo measured
5.2 metres in length, Strahan and Thomas, 1975).

The facilities finally set up at the University of New South Wales were largely
governed by space available, although valuable suggestions on minimum dimensions

118 Aust. Zool. 19(2), 1977

THE PLATYPUS IN CAPTIVITY

were received from Mr. D. Thomas of Taronga Zoo. The first tank built (Fig. 1)
was constructed from 1.1 cm waterproof plywood with the dimensions of 2.4 m
by 1.2m by 0.6m. The tank was filled to approximately 0.3 m in depth, which
put the water level just below a shelf placed at one end. From this shelf a tunnel
was built along the outside of the entire length of the tank, ending in a unit of
three sleeping boxes. These could be closed individually.

After the failure of the original coating (polyurethane) the joints of the
tank were filleted with fibreglass cloth, and the whole complex was recoated with
a tar epoxy resin (Epirez 304)*. The resin was reapplied after nearly a year
because of abrasion of the surface by ice used in experiments in the tank, and
this second coating remained waterproof after 2 years of further continual use.
A second tank was built and because of the slight odour left with the tar epoxy,
a ketimine cured epoxy was used as a coating (Epirez 235)*. Waterproof plywood
was replaced by marine plywood in the construction of this second tank.

Figs. 1 & 2. Specifications of platypus enclosures.

1 RESTING 2
CHAMBERS TANK
a 8

HINGED LID

DRAIN

YEWWIIA{EHA’AV’IVWI MA AV AAVVCECE
PIPE NN

N
N
x

ANNANAAAANAAAAAANAAAANAAAAAAAAAAARAAAARAAAAAAAARAAARRAD SAV
NY

; ‘ ip Hi
SIDE VIEW Pa
HINGED LID SUPPORTS
i ah ae
SHELF |4 REMOVABLE LID
7
61 TUNNEL
REMOVABLE LIDS alee bce 1. Large Tank (plywood).
TOP_VIEW 2. Small Tank (fibro-cement).

The whole complex was mounted on a table and sloped slightly from one
end. This allowed drainage through a 5.1 cm plug and waste. Water from the fur
of animals was able to escape through drain holes in the floor of the tunnel. Collins
(1973) mentions the use of rubber “squeegees” to remove excess water from
animals moving along the tunnels of enclosures. All of the platypus which lived

* Indelab Pty. Ltd., Granville, N.S.W.

Aust. Zool. 19(2), 1977 119

T. FR. GRANT, R. WILLIAMS and F. N. CARRICK

in captivity groomed and dried themselves before entering the sleeping chambers
without the aid of these devices.

Sleeping chambers were lined with soft dry grass and condensation was dried
off the roof and walls daily, when the animals were weighed and grass was
checked for dryness and/or replaced. Initially soil was used under the grass but
this proved a hinderance to the closing of doors in the tunnel and nest unit
and became very soggy when new animals did bring water into the sleeping
chambers. Platypus were at first carried and housed in hessian bags but this
practice was quickly abandoned as it produced severe abrasions to the animals’
feet.

During an experiment by one of the authors a female platypus was maintained
in a small circular tank (0.6m x 1m in diameter, Fig. 2) with a simply
constructed tunnel and sleeping box complex, for 3 months. However, a male
animal had to be returned to the large tank when it did not settle in the small
one, and another female kept in it for a short time died soon after its return
to the large tank. In this latter instance the large tank had been vacated to enable
replacement of the original coating. The smaller complex was more convenient
and proved satisfactory in the instance mentioned, but in view of experience, the
larger model has proved to be the most satisfactory enclosure for housing platypus
in an animal house situation for any length of time.

In all instances tanks were drained, cleaned and refilled daily and were
scrubbed with running fresh water every 4 or 5 days to stop the accumulation of
algal growth.

FOOD

Platypus are known to eat a variety of food items in captivity but the
provision of an ample supply of acceptable food has always hindered their
maintenance in zoos and sanctuaries. Reports in the literature suggest that the
weight of food taken by a platypus in one day can amount to over half its own
body weight. Burrell (1927) recorded one of his captives devouring 70 earthworms,
10 ground grubs, and 600 salt water prawns in 72 hours and Fleay (1944)
lists the food eaten by a lactating female as 400 earthworms, 338 grubs and 38
live yabbies during one 24 hour period.

Table 2 compares the daily diets given in two other institutions with the
food supplied to the animals during the present studies. These lists can be noted
as little more than guidelines because food intake by an individual is never static,
being modified by factors such as activity, ambient temperature, animal size and
individual diet preferences. In dealing with the platypus experience always played
a great part in determining diet offered, and amounts and types of food given
varied from week to week and from animal to animal. Most animals took some
time to begin eating prawns. Some never ate them in any quantity, and in
these instances many more earthworms and cockroaches had to be supplied.
Mosquito fish (Gambusia sp.) were eaten by some of the captive platypus,

120 Aust. Zool. 19(2), 1977

THE PLATYPUS IN CAPTIVITY

TABLE 2
DAILY DIETS GIVEN TO PLATYPUS IN CAPTIVITY
A. Bronx Zoo, New York*

Items Details Quantities

1. Earthworms 8 454g
2. Crayfish as 24
3. Frogs — 2,
4. Eggs steamed 2
5. Cockroaches and

Mealworms — 1 handful
B. Taronga Zoo, Sydney *+
1. Earthworms oa 454g
2. Prawns — 2272
3. Egg yolks hard boiled 2
4. Mealworms _—
5. Woodlice oes ie

s ll quantities
6. Scarabaeid larvae — } ge
7. Small crayfish — ee
C. School of Zoology, University of N.S.W.
1. Earthworms given with some soil (live) 160g
2. Prawns heads removed 340-510¢g
3. Cockroaches killed 12-24
4. Mealworms live 12-36
5. Mosquito fish only to some up to 100 in place of 3 & 4.
6. Small crayfish animals on occasions 6-12
-e Collins (1973 ):
+ Strahan and Thomas (1975).
TABLE 3

ACCEPTANCE OF VARIOUS FOOD ITEMS BY THE EIGHT PLATYPUS KEPT
IN CAPTIVITY

Food Items 21 Q2 Q5 231 248 38 3 20 G2
Earthworms A A A A A A A A
Mealworms — A == A A A A S
Cockreaches — == — A A — A Ss
Mosquito fish == == = A A U S N
Insect larvae oe — — A A A A A
School prawns N A N U S A U U
_ Yabbies — A — A N — —
Eggs — N — N _ N — N
Squid — N — — N — —_
Mussels (marine) — N — -— — N — —
A = always S:°= seldom — = not offered
U = usually N = never

Aust. Zool. 19(2), 1977 121

T. R. GRANT, R. WILLIAMS and F. N. CARRICK

while others showed little or no interest when these fish were released live into
the tank. Data showing diet preferences of the individual animals are summarised

in Table 3.

Earthworms were the most preferred food item and large worm cultures
were established to cope with the demand. These were in pits, 0.5m to 0.6m
in depth and of various sizes, the bottoms and sides being lined with galvanised
iron. Soil was placed in the pits and over a period of several weeks, organic
matter in the form of grass clippings and leaves was worked into the soil.
Earthworms of various species were introduced into these beds, which were kept
moist (but not sodden) and supplied with more organic matter, including horse
manure, at regular intervals for several months.

Lumbricus terrestris in Britain takes up to a year to reach maturity after
10 weeks of incubation in the cocoon. Environmental factors such as food supply,
temperature and soil moisture are known to influence rates of production in
this and other species (Edwards and Lofty, 1972) but even if optimal conditions
could be supplied in the worm beds, turnover rates are still likely to be quite
slow. For this reason beds were kept unused for as long as possible and when
harvested, only the smaller worms were removed. In times of very high demand
breeding worms were taken, but where possible these were replaced by new
stock at a later date. One set of beds were established in a lawned area and
migration of worms from the lawn into the beds helped to supplement numbers.

Slowing of growth and a reduction in reproductive capacity in the winters
proved to be a problem but the establishment of beds in a temperature-controlled
glass house permitted ample supply of worms during the final winter of the
study. An attempt was made to heat the outside cultures using trenches with
a mixture of grass clippings and slaked lime, kept moist to produce heat. No
noticeable effect on worm populations was observed.

Cultures were always covered with a layer of organic material and hessian
bags which were kept moist. The organic material was mixed into the soil and
replaced every time worms were removed. One large bed was kept entirely for
“red” and “tiger” worms (probably Eisenia sp.), being supplied with rotting
fruit, as well as leaves and grass. These worms seemed to have a higher turnover
rate. Platypus usually only ate these in the first few weeks of captivity but their
utilisation of them at this time of maximum demand eased the burden on the
other cultures.

Various methods of extraction of the worms from the soil were tried (electrical
current, chemicals and heat) but forking through the soil was the most successful
and also served to mix in fresh organic material.

Because of their slow rate of reproduction, earthworms are not the most
ideal food for captive platypus being maintained in a laboratory situation. It was
fortunate that the animals could be kept on the mixed diet given in Table 2.
Mealworms were the only other food item cultured during this study. Other

122 Aust. Zool. 19(2), 1977

THE PLATYPUS IN CAPTIVITY

food was either purchased or supplied from cultures existing in other parts of the
School of Zoology.

ANIMALS

Platypus were captured using the methods detailed in Grant and Carrick
(1974). They were weighed daily, or once every two days after their capture.
This was done in an attempt to keep a check on their condition and to slowly
adapt them to being handled.

RESULTS AND CONCLUSIONS

Weight changes in eight of the animals kept are summarised in Table 4.
All animals showed an initial weight loss before beginning to regain weight
and stabilise. During experimentation, which was not commenced until animals
had regained at least 80% of their original weight, some fluctuations of body
weight normally occurred. If a steady decline commenced, experiments were
terminated until this trend ceased.

Platypus released after being maintained in captivity were, with one exception
above their original weight at capture. Male number 8 reached 91% of its
original weight while telemetry experiments were in progress but lost weight
during attempts at metabolic determinations. These experiments were terminated
and after a recovery period the animal was released. Throughout its captivity
this male retained substantial tail fat deposits.

Initial attempts to maintain platypus in the laboratory for a prolonged
period were not successful. Both the platypus which died within a month of
their capture did not regain lost weight (Table 4). Female number 2 was kept
successfully for several months but died after a rapid decline in its condition,
whilst housed in a small tank during emergency repairs to the main tank. In

TABLE 4
WEIGHT CHANGES IN, THE. FIGHT: PLATYPUS KEPT IN. CAPTIVIPY

ANIMALS on Oe oe) $8 O51 3 20 $22 248
Capture Wt. (g) 764 790 950 2350 1000 1348 1430 992
Date GO D212 6:62; D2 2 S21 Ghe | Ono oO 142 aL. Val Se fed,
Min. Wt. 618 604 665 1843 822 ra? 1344 950
Date A Pega Pai) MOM Se a 2278 a Ocoee ea Wh Sead. so Ouee
Max. Wt. 764 956 950 2350 1002 1680 1708 1078
Date DOG 2 UE 9T2 PO AZ 12-3273 a1 S14 201274 4S: = esas
Death Wt. 618 628 665 a — = — —
Date ATA 2 Sl lea a Oy ley Sa — — — _
Release Wt. _ — — 1982 1002 1680 1634 1046
Date = = Se Or ay OLA ae Noe ae Some Loe
Duration (months) 38) 5 1 11 Bh) 6 3:33 3

Aust. Zool. 19(2), 1977 123

T. R. GRANT, R. WILLIAMS and F. N. CARRICK

these three instances 20-30% of their original body weight was lost by these
animals. Post mortem examination suggested that death was caused by septicaemic
pneumoenteritis, possibly caused by bacterial proliferation associated with stress-
induced lowering of immunological mechanisms. Similar symptoms were noted in
a platypus which died at the Australian Reptile Park, Gosford, N.S.W. (Mr. George

Wilson, personal communication ).

Platypus were kept in a laboratory situation during this project with some
success. It must be noted that any researchers considering the maintenance of
this species in captivity should make provision for the considerable commitment
of time required for the routine animal husbandry and culturing of the live-food
component of the diet. As Burrell (1927) indicated, the platypus “demands
a varied diet, and will starve to death in the presence of food which no longer
pleases it. It must have clean, clear water, and sweet, dry sleeping-quarters. It
is impatient of observation, and resents being handled. It is easily killed by too
much excitement”. Perhaps the most disturbing aspect of keeping this species in
captivity is the realisation that, in spite of all the care and precaution taken, some
of the animals may die.

REFERENCES

BURRELL, H. (1927). The Platypus, Angus and Robertson, Sydney.

COLLINS, L. R. (1973). Monotremes and Marsupials: A _ reference for zoological
institutions, Smithsonian Institution Press, Washington.

EADIE, R. (1934). The platypus in captivity. Vic. Nat. 5/, 3-11.

EDWARDS, C.:.A. & J. R. LOFTY (1972). The Biology of Earthworms. Chapman and
Hall, London.

FLEAY, D. (1944). We Breed the Platypus. Robertson and Mullens, Melbourne.

GRANT, T. R. & F. N. CARRICK (1974). Capture and marking of the platypus,
Ornithorhynchus anatinus, in the wild, Aust. Zool, 18, 133-135.

SMYTH, D. M. (1970). Thermoregulation in the platypus, Ornithorhynchus anatinus (Shaw).
Honours Thesis, Dept. of Zoology, Australian National University, Canberra.

STRAHAN, R. & D. E. THOMAS (1975). Courtship of the platypus, Ornithorhynchus
anatinus. Aust. Zool. 18, 165-178.

ACKNOWLEDGEMENTS

The N.S.W. National Parks and Wildlife Service and the N.S.W. State Fisheries Branch
are acknowledged for allowing us to trap platypus in the rivers of N.S.W. Part of the
support for this work was provided by a grant from the Australian Research Grants
Committee. George Wilson of the N.S.W. National Parks and Wildlife Service is also
acknowledged for his personal communication.

124 Aust. Zool. 19(2), 1977

The Need for Joining Illawarra Wilderness Areas

N. H. ROBINSON
33b Gipps Road, Keiraville, N.S.W. 2500

ABS PRACT

An examination of proposed and existing reserves between Royal National Park
and the Shoalhaven River shows that each is ecologically valuable and contributes
towards the preservation of existing flora and fauna. Due to the altering pattern
of land usage and wide road construction, it has become apparent that a number
of mammal species at present inhabiting the reserves may cease to do so when
these reserves become small isolated wilderness areas.

A system of corridors is suggested to aid in making possible mammal movements
between all reserves and other areas of significant biological value.

INTRODUCTION

A mammal survey of the whole area was conducted between 1966 and
1970 inclusive with work carried out since in selected areas. Every nature reserve
and national park has been surveyed or check listed as well as all sections of
bushland of significant size. The methods and results of this survey are contained
in “Mammals of the Illawarra and Surrounding District’? (Robinson, in press).

The major wilderness areas in the region are —

(1) Catchments of the Metropolitan Water Sewerage and Drainage Board;
(2) Morton National Park; (3) Royal National Park — Heathcote National Park;
(4) Illawarra Escarpment which includes land donated by A.I.S. Pty. Ltd. to
the National Parks and Wildlife Service.

The smaller national parks and nature reserves are —

(1) Macquarie Pass National Park; (2) Seven Mile Beach National Park;
(3) Barren Grounds Nature Reserve; (4) Red Rocks Nature Reserve; (5) Black
Ash Nature Reserve; (6) Rodway Nature Reserve; (7) Devils Glen Nature
Reserve.

RESULTS AND? DISCUSSION

EFFECTIVE POPULATION NUMBER AND RESERVE SIZE:
Tyndale-Biscoe and Calaby (1975) stated: ‘Effective population number
is the population size that will retain the original genetic diversity of the species,
or a large fraction of it, in perpetuity and provide the genetic means for continued
evolution. It must take account of natural fluctuations and be large enough to

Aust. Zool. 19(2), 1977 125

126

40

35

30

NUMBER

25

N. H. ROBINSON

20
15

OF

10

SPECIES

‘YN ualy
S|1Aaq

‘dN Aempoy

dN
ysy 9F1g

dN
syaoy pay

dN yoeag
aj|A| uaAaS

dN ssed
auenhoesy

WN spunolg
uaueg

yied ‘diedsy
pasodolg

9}094}e3}
pue jeAoy

d'N volo

paAaAins
paie jenjoe
‘aasMmW
s}uawya}e)

1.064 664 647 89 83 40

1,962

107,027 17,163 8,000

110,000

app.

app.

AREA in HA.

Illawarra wilderness areas and mammal species within them.

1.

Fig.

Aust. Zool. 19(2),

1977

ILLAWARRA WILDERNESS AREAS

withstand the vicissitudes of fire, disease and drought, it is the lowest number
that the population can fall to under these circumstances.”

After examining the work of Crow and Kimura (1970) and Main and Yadav
(1971) they point out that the relatively abundant species such as the greater
glider, Schoinobates volans, needs a reserve of no less than 6,000 ha. Species
dependent on forest and less abundant, would need a proportionately increased
area to accommodate a minimum effective population. Main and Yadav (1971)
suggest selected areas of more than 20,000 ha which are likely to retain their
characteristic biotic diversity.

Tyndale-Biscoe and Calaby (1975) also state: “The evidence at present
available indicates that species with high attachment to their home sites do not
move to other areas because these are already occupied and secondly, the number
of animals that the small reserve can support in isolation is quite inadequate for
the species long term survival.”’

The author has shown this to be so during studies on this district’s reserves.
The commonest large native mammal in Illawarra is the swamp wallaby, Wallabia
bicolor, which inhabits forest or natural areas containing thick undergrowth.
Edwards and Ealey (1975) studied the home range and territory of this species
in Victoria. They stated: “The averages of the areas enclosed by the most widely
separated records was 4.5 ha (males) and 7.2 ha (females). In any case these
values are minimum estimates of the real dimensions.’’ The second most commonly
observed mammal is the wombat, Vombatus ursinus. Mclllroy (1973) found a
density of approximately one wombat per 4% ha. in tall open forest near Tumut.
With an effective population of 1,000 individuals of a species appearing to be near
to the minimum for the continuance of genetic variability, Tyndale-Biscoe and
Calaby consider 5,000 as a safe number, as this figure is the total population,
including non-breeding juveniles, etc.

From this it can be seen that swamp wallabies and wombats need a large
area to ensure their ultimate survival. In the Illawarra where some species are,
overall, thinly distributed, as in the case of the wallarco, Macropus robustus,
very large areas are necessary. Only the larger mammals are given as examples
because the smaller species such as Antechinus and Rattus, utilize smaller home
ranges and these should survive if areas of sufficient size for large species remain.

ILLAWARRA RESERVES AND SPECIES:

Figure 1 shows the reserve size and number of recorded mammal species
excepting bats and marine mammals. The numbers might well be increased with
additional work. The following points should be noted in relation to Fig. 1.

(a) The Catchments surveyed were Cataract, Cordeaux, Avon, Nepean, Woronora

and O’Hares.

(b) Extensive additions have been made to Morton National Park since the
completion of the field work which led to the compiling ot the check list.

Aust. Zool. 19(2), 1977 AQT

N. H. ROBINSON

(c) Seven Mile Beach National Park has long been isolated by rural development.
The largest terrestrial native mammal recorded was the long-nosed bandicoot,
Perameles nasuta.

(d) Field work is still progressing on Red Rocks and Devils Glen Nature Reserves,
the boundaries of which are largely undefined. Devils Glen is predominantly
a deep, steep sided gully, difficult of access.

NOTES ON RESERVES:

The Metropolitan Water Sewerage and Drainage Board Water Catchments
and the Illawarra Escarpment are as one whole at present with Royal National
Park joined by existing forest to the [lawarra Escarpment. Royal National Park
and Heathcote National Park have effectively been separated by an expressway
and the urban spread.

Morton National Park and Red Rocks Nature Reserve are joined at present
by narrow natural or near natural belts of vegetation. Rodway Nature Reserve
is isolated to a considerable degree but only a short distance separates it from
the Barren Grounds Nature Reserve which is joined to Morton National Park
via the proposed Buderoo National Park and belts of natural forest. Black Ash
and Devils Glen Nature Reserves are almost joined together but are isolated at
present from any large wilderness area.

It can be seen from Fig. 1. that only the three largest wilderness areas
contain sufficient biotic diversity to support a normal range of species typical of
this part of Australia. A number of the species currently inhabiting most of our
reserves will cease to do so if these reserves are isolated by the changing pattern
of land usage

One example is the Barren Grounds Nature Reserve of 1,962 ha. It was
surveyed during the years of 1966-1970. Twenty-five (25) species excluding bats
were recorded. Of these, five were seen on no more than one occasion each,
because of low numbers e.g. red-necked wallaby, Macropus rufogriseus, or cryptic
behaviour, e.g. pigmy glider, Acrobates pygmaeus.

At present the reserve is part of a large wilderness area. If it was to become
an “island”? due to surrounding development, it is doubtful whether any large
native mammal species would ultimately survive because much of the area is
unsuitable for them. They utilize the surrounding wilderness as well at this time.
Even Royal National Park, covering 14,912 ha., will probably lose a number of
its large mammal species unless it is rejoined by corridors to Heathcote National
Park and the proposed Illawarra Escarpment Park. Examples are the wombat,
Vombatus hirsutus, and the wallaroo, Macropus robustus.

Only the catchments of the M.W.S.D.B. and Morton National Park appear
large enough in isolation to retain their present species. Even on the catchments,
there is a definite threat to some larger species through the effects of proposed
very wide roads or expressways unless provisions are made in the design and
construction of such roads to allow mammals to cross.

128 Aust. Zool. 19(2), 1977

ILLAWARRA WILDERNESS AREAS

Forest Areas of Illawarra and Suggested Corridors

t 2 5 1Okm oo

Bowral

“tc EGE ND:

Z 1. Royal

2. Heathcote

3. Macquarie Pass

4. Seven Mile Beach
5a. Barangary (Morton)
5 Morton

Nature Reserves
Barren Grounds
Rodway

Devils Glen
Black Ash

Red Rocks

MOom@p

Others

C.A. MW.S.&D.B Catchment Areas
(only forested part shown)

F. Holsworthy Field Firing Range

P.B. Proposed Budderoo National
N.P. Park

P.E.P Proposed Escarpment Park

W.S.F Wedderburn State Forest
Corridors

©xxXxx (CI to ClO)

! at Forest Areas
Vb ve ; SOEs
\n el
Lice Se

| MELBOURNE Critical Factors
Mountain Roads

Aust. Zool. 19(2), 1977 129

N. H. ROBINSON

To try to keep maximum ecological values in all the National Parks, Nature
Reserves and large wilderness areas, a system of corridors is suggested to make
possible faunal movements between them. Corridors linking these would
contribute towards one whole rather than having three major wilderness areas
and a number of smaller ones, each of which is very valuable in many ways and
which contribute towards the preservation of existing flora and fauna.

Once a framework is established, it might be possible to enlarge upon it
if these minimum sized corridors are found to be of insufficient width and habitat
variety.

CRITERIA USED IN CORRIDOR SELECTION:

The distance between areas to be linked is important. Where possible short
corridors have been chosen. Vegetation types are crucial and eucalypt forest and
rainforest together in this district form a near ideal link, with the eucalypts on
the exposed sections and escarpment top, while the rainforest usually grows in
the sheltered parts such as along gullies or escarpment sides. Unfortunately there
are vegetation gaps in several corridors. Regrowth however, would eventually grow
to form a corridor.

The composition of species and their habits, where known, can have a big
bearing. Wombats and red-necked wallabies in this district generally keep to the
escarpment top, although around the Shoalhaven River area, red neck wallabies

do descend.

At the top of Macquarie Pass, the belt of existing vegetation is crucial
because wombats do not move below the escarpment at this particular place.

Existing land usage has to be considered. Steep hillsides on water catchments
such as on the rim of the Kangaroo Valley, or elsewhere, are best vegetated by
natural forest to assist soil conservation.

PROPOSED CORRIDORS:

The corridors suggested are shown on the accompanying map (Fig. 2). The
actual boundaries, description of geology and vegetation, mammals currently
inhabiting each, immediate threat and other relevant comments have been
submitted to the National Parks and Wildlife Service. For the purpose of this
paper, only the mammals inhabiting each corridor are shown. The map shows
the existing forested areas.

It is hoped that in future satisfactory crossings can be provided between
both sides of the expressway and Princes Highway between Bulli and Engadine.

CORRIDOR 1 would link the Royal National Park to the proposed Illawarra
Escarpment Park and M.W.S.D.B. Catchments.

CORRIDOR 2 would link the proposed ITlawarra Escarpment Park and
M.W.S.D.B. Catchments to Macquarie Pass National Park and
the Barren Grounds Nature Reserve.

130 Aust. Zool. 19(2), .1977

ILLAWARRA WILDERNESS AREAS

TABLE 1
REFERENCE CHECK LIST OF MAMMALS INHABITING THE PROPOSED
CORRIDORS
CORRIDORS

SPECIE CUES ED ER ESKSRARSEE
Tachyglossus aculeatus ~
Ornithorhynchus anatinus (Oe AeA ey lapel ites | eee VA
Antechinus stuartii we Y
pasar verins = Fe || || Lae fa [oe [ow Jw
Dasyurus maculatus Wr v
a a ed es We ea BeBe
Vombatus hirsutus i
Acrobates pygmaeus Pap fb be Ve
Cercartetus nanus ee pecak\ | ee
pens awrats = | AL AL PP | pe
Petaurus norfolcensis ae re ee ee a Ee
Petaurus breviceps ay eA CARO AE a4 Va
Trichosurus vulpecula an ah yg 4 WH
Trichosurus caninus Ree ees oa ee
Pseudocheirus peregrinus Se
Schoinobates volans Fae wd Pau Ea ve
Phascolarctos cinereus meee Foe eee ee a as
Potorous tridactylus ot tt 4
Petrogale penicillata (| re cid pee | ae Ped a ees aa ee
Pla thet om Cr ea a a
Macropus rufogriseus Fe Gay ere a ea £
Macropus robustus Fo a Li
Macropus giganteus eae ee RS Ss ee a a | we
WolbiaBetr wid civ lele| |e
te fap ape bia aw seabed Cane ee
Hydromys chrysogaster Alem eta alt Melee
Oryctolagus cuniculus Roe he PE ere ee Oi
Fels cn ay lu todwlvivis be
Cons font ee eee ae
aps ae 22 oa re aa tee
Cervus timoriensis Pa bp pototo ~
ae on a ae tas We ed Ae

*No recent record .
Aust. Zool. 19(2), 1977 131

N. H. ROBINSON

CORRIDOR 3 would link the Barren Grounds and Rodway Nature Reserve.

CORRIDOR 4 would link Rodway to Black Ash and Devil’s Glen Nature Reserves.
CORRIDOR 5 would link Black Ash and Devils Glen eserves to Good Dog
CORRIDOR 6 would link Good Dog Mountain to Red Rocks Nature Reserve.
CORRIDOR 7 would link Morton National Park to Red Rocks Nature Reserve.

CORRIDOR 8 would link the Barren Grounds through the proposed Buderoo
National Park to Barangary Park, an isolated part of Morton.

CORRIDOR 9 would link Barangary Park to Morton National Park.

CORRIDOR 10 would link Bengalee Creek to Bugong Creek’s edges westwards
to Morton National Park.

The long term planning of Region II, embracing the whole of the Illawarra
and more, is being carried out. Unless provision is made to include mammal
corridors, only the largest wilderness areas will retain maximum wildlife value.
Some species at present widespread will eventually contract in range to only the
largest parks or reserves.

It is suggested that similar corridor schemes be instituted where possible
in other parts of Australia. Unless this is done, the future of a number of species
is indeed bleak in many areas.

ACKNOWLEDGEMENTS

The author wishes to thank the National Parks and Wildlife Service for the necessary
permits to work on the various National Parks and Nature Reserves; the Metropolitan
Water, Sewerage and Drainage Board for permission to enter their catchments; to various
employees of both organisations for their assistance; to a number of land owners who gave
permiszion to enter their properties and to Mr. P. Hendricks for his assistance on a number
of field trips.

REFERENCES
DIAMOND, J. M. (1975). The Island Dilemma: Lessons of Modern Biogeographic Studies
for the Design of Natural Reserves. Biol. Conserv. 7, 129-146.

EDWARDS, G. P. & E. H. M. Ealey, (1975). Aspects of the ecology of the swamp
wallaby Wallabia bicolor (Marsupialia : Macropodidae). Aust. Mammal. J 307-317.

MAIN, A. R. & M. YADAV, (1971). Conservation of Macropods in Reserves in Western
Australia Biol. Conserv. 3, 123-133.

ROBINSON, N. H. (In press). Mammals of the Illawarra and Surrounding Districts.

ROBINSON, N. H. (1976). Mammals and Expressways (Parks and Wildlife, 7, 173).

TYNDALE-BISCOE, C. H. & J. H. CALABY, (1975). Eucalypt Forests as Refuge for
Wildlife. Aust. Forestry 38, 117-133.

132 Aust. Zool. 19(2), 1977

A New Species of Python from Arnhem Land

G. F. GOW
Museums and Art Galleries of the Northern Territory, P.O. Box 4646, Darwin, N.T. 5794

ABSTRACT

A new species of Python (Family Boidae) is described from two male specimens
from the Arnhem Land escarpment of northern Australia. The new species is
considered to belong to the reticulatus group of the genus Python and may be
distinguished from all other species by its high ventral and subcaudal scale
counts. An account is given of meristic variation of the dorsal scale rows
of the new species and comparison made with other species.

INTRODUCTION

About ten species of pythons belonging to four genera are known from
Australia (Cogger, 1975). The most recent critical work concerning the Pythoninae
is that of McDowell (1975) whose generic definitions are followed here. McDowell
has divided the genus Python into the molurus and reticulatus groups; the latter
possessing infralabial pits set in a deep groove, slit-like supralabial pits set
diagonally, hemipenis with proximally directed chevron-like flounces, and the
upper lip below the eye light-coloured and similar in colour to the rest of the
upper lip. The reticulatus group includes Python amethistinus, P. boeleni,
P. spilotus, P. reticulatus and, possibly, P. timorensis. Two specimens of a boid
snake from Arnhem Land in northern Australia appear to belong to this group
also and are described herein as a new species. The specific name is derived
from Oenpelli, the settlement nearest to the type locality.

Python oenpelliensis n. sp.

Holotype: 6; 6.5 km S.W. of Oenpelli, Northern Territory, Australia (12° 21’
S., 133°01’E.), hereby designated type locality. Coll. Mr. B. Jukes, 17.V1I.1975.
Specimen in Museums and Art Galleries of the Northern Territory, Darwin
(Reg. No. RO840). Measurements (in mm): Total length 3560; snout to vent
3037; tail 523; head length 69; snout to eye 25.4; interorbital distance 21.7;
snout to nostril 4.7; orbit to nostril 17.3; internasal distance 11.3.

Paratype: é; Little Nourlangie Rock, N.T., Australia, (12°51’S., 132°487E.).
Coll. Dr. R. Begg, 9.VI.1976. Specimen in Australian Museum, Sydney (Reg. No.
R55009). Measurements (mm): Total length 3593; snout to vent 3043; tail 550;
head length 71.5; snout to eye 25; interorbital distance 22.2; snout to nostril 4.6;
orbit to nostril 18.6; internasal distance 10.7.

Aust. Zool. 19(2), 1977 133

G. F. GOW

DESCRIPTION OF HOLOTYPE

A large boid snake with a strongly prehensile tail. Scales without apical pits
and without granular gular scales bordering the mental groove.

Rostral visible from above, slightly broader than long, with a pair of deep
slit-like sensory pits. Supralabials 15, the seventh and eighth on the left side and
the seventh, eighth and ninth on the right entering the eye. First three supralabials
with deep sensory pits; pit of first supralabial a long diagonal slit broadening
dorsally and contacting ventral edge of the nasal. The slit of the second
supralabial almost contacts ventral edge of nasal; third supralabial slit is distinct
but shorter and broader than first two. Fourth supralabial without a distinct
slit but an obscure elongate mark is present. Infralabials 24, six of the more
posterior infralabials with deep rectangular pits. The row of infralabial pits
begins beneath the eye and is sunk in a longitudinal groove. The row is preceded
by an infralabial with a shallower pit.

One pair of prefrontals is present, each almost twice as long as broad.
Internasals preceded by a pair of small scutes. The posterior prefrontal area,
extending to the anterior frontals and supraoculars, consists of 22 subequal scales.

Uy y
Whirijjj
jj Y
Vij

Fig. 1. Python oenpelliensis n. sp., holotype, in life.

134 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PYTHON

Figs. 2 & 3, Python
oenpelliensis n. Ssp.,
holotype,

2, dorso-lateral
view of head.

3, dorsal aspect of
head.

135

Aust. Zool. 19(2), 1977

Gok. "GoW

Frontal plate divided into three; an equal anterior pair followed by a single
plate. Parietal scutes absent, the top of the head covered with numerous subequal
scales. Nasal partially divided; nostril directed upwards and outward; fissure
extending from nostril to anterior loreal scutes. Loreal scutes, including small
granules, 30 on right and 26 on left side. Preoculars 4 right, 3 left; postoculars
6 right, 5 left. Supraocular entire on right side (with partial division), divided
on left with anterior portion the largest.

Mental groove relatively long, bordered anteriorly by first infralabials; two
scales lie in rear of groove. Lateral border of groove of 9 and 10 scales similar
in size to other gulars.

Dorsal scales smooth without apical pits; the first row large but less than
half as broad as ventrals. At one head length behind head scales in 53 rows; at
midbody, 70 rows; one head length anterior to the cloaca, 35 rows. About
50% of dorsal scale rows doubled (see discussion).

Ventrals 429° Subcaudals 155, mostly divided) (7 16, 19, 34. 35,° 56) 39.
118, 119 are entire). Anal single. Cloacal spurs present.

One pair of premaxillary teeth. Maxillary teeth; 11 on right maxilla (estimated
6 missing), 14 on left (est. —3); estimated maxillary total 17. Palatine teeth;
10 on right (est. —5), 15 on left (intact series). Dentary teeth; 15 on right
(est. —3), 14 on left (est. —4); estimated dentary total 18.

Hemipenis forked extending for about 10 subcaudals forked about 3
subcaudals proximal to this. Chevron like flounces of organ, 7.

DESCRIPTION OF PARATYPE

As for holotype with the following exceptions (figures in brackets refer to
holotype).

supfalabials’ 17. °(15)= 8," 9° meet eye (7-8); om. leit: 8, 9: 110) meer ere
(7, 8, 9) on right. Infralabials 24 right, 22 left (24); seven on left pitted.
Posterior prefrontal area 25 (22). Loreals 45 (30) right, 43 (26) left. Postoculars
5 (6) right, 7 (5) left. Supraoculars right divided, left entire (opposite
arrangement in holotype). Ventrals 445 (429). Subcaudals 163 (155), mostly
divided; 15 single subcaudals (9).

COLOUR. NOTES

Pale fawn above merging into a lighter putty grey on the lower sides. A series
of dark grey-brown blotches on the dorsal surface is irregularly arranged in four
to five longitudinal rows. These dark blotches and streaks are largest in the
vertebral series and much smaller laterally. Many of the dorsal scales carry an
indistinct dark brown or black spot at the apex. The dorsal pattern becomes more
obscure posteriorly. The dorsal scales of the tail are dark-edged producing a
reticulated effect. The top of the head is pale fawn intermixed to a lesser degree with
dark brown. Several of the head shields have the centre or the edge coloured dark

136 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PYTHON

brown. A dark brown supraocular streak about two scales wide extends from
the supraoculars down each side of the head to the nape. A similar streak,
indistinct anteriorly, runs from the nostril through the eye and is bordered above
and below by equal areas of pale fawn. The ocular and supraocular streaks are
joined by a diffuse band of dark spots in the post-temporal region. The supralabials
and infralabials are fawn, the latter partially edged with dark brown below.
The sublingual and gular area is white and the ventral surface of the body off white
to pale yellow. |

Holotype and paratype do not differ markedly in colour.

LOCALITY AND COLLECTING NOTES

Both specimens of P. oenpelliensis were collected near sandstone rock
outcrops; the holotype specimen was collected crossing a road bordered by
sandstone. More data is available for the locality from which the paratype was
collected. The area is Myrtle-Pandanus savannah and mixed scrub. The specimen
was collected a short distance from a sandstone outcrop (Dr. R. Fox, pers. comm.).

180-4

alk

i]
=
= a
140 re c
2 =
®
: 2
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ep)
Ww
a
oO S
W100 =
2a) | =
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=) \2
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Oo
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=
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w $ a v= a
=) 2 2 2.2 =
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150 210 270 230 390 450

No. VENTRAL SCALES
Fig. 4. Ranges of ventral and subcaudal scale counts for eleven species of the genus

: Python. Data from Boulenger (1893 & 1912), Cogger (1975). Fitzsimons (1962),
Gow (1976), McDowell (1975) and Taylor (1965).

Aust. Zool. 19(2), 1977 137

Gi FeGOw

It was noted by both collectors and subsequent handlers that both specimens
were docile during and after capture. Anecdotal evidence suggests that individuals
of this new species may be longer than the specimens recorded herein.

DISCUSSION

Python oenpelliensis n. sp. adds a further species to the diversity of
Pythoninae known from the Australasian region (see McDowell, 1975 p.3). The
new species may be distinguished from other species of the genus Python by
the high ventral and subcaudal scale counts, no other species possessing more
than 346 ventrals or 122 subcaudals (maximum counts recorded for Python
amethistinus; MacDowell loc. cit.). The distribution of numbers of ventrals
and subcaudals for eleven species of Python are shown in Fig. 4. These
characteristics are also adequate to separate the new species from all other
Australian boid snakes.

TABLE 1
DOUBLE DORSAL SCALE ROWS OF 9 SPECIES OF PYTHONS

wie OE | ee ee |
ventrals scale rows dorsal rows
Python oenpelliensis 429 221 SiS Holotype
7 445 223 50.1 Paratype
P. amethistinus 343 16 4.7 Queensland
P. boeleni 314 57 18.2 Sepik R., P.N.G.
P_ spilotus 296 76 plate Oenpelli, N.T.
- 282 98 34.8 Central Australia
291 109 37.5 | Central Australia
99 283 90 31.8 Melville Is., N.T.
Liasis olivaceus | 353 107 30.3 | Adelaide R., N.T.
L. childreni | 261 65 24.9 Central Australia
L. boa | 267 56 20.9 New Ireland
L. mackloti * 275 83 S02 Katherine, N.T.
A spidites
melanocephalus 329 124 37.7 Katherine, N.T.

2S BA ROE SOR es A SSR rk | nt rac Ge |S i Me eM GS
* Liasis fuscus Peters, 1847 is a synonym of Liasis mackloti Dumeril and Bibron according
to McDowell (1975, 6.34 et. seq.).

138 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PYTHON

The dorsal scalation also distinguishes P. oenpelliensis from other species
of Pythoninae. The number of dorsal scale rows of the new species is not the
same as the number of ventral scales. About 50% of the ventral scales correspond
to double, not single, rows of dorsal scales. It is believed that this is the first
time that this meristic variation has been reported in the Pythoninae. The degree
of doubling of dorsal scale rows in relation to the number of ventral scales is
shown in Table 1.

It is apparent, from the limited data available, that the doubling of dorsal
scale rows is very much greater in P. oenpelliensis than in the other species
investigated. It is not apparent, by inspection at least, whether any pattern is
present in the distribution of double dorsal scale rows either in a specific or an
individual sense. Such analysis must await more data but it is suggested that the
meristic variation of dorsal doubling in pythons is worth further investigation.

ACKNOWLEDGEMENTS

Thanks are due to Mr. B. Jukes and Dr. R. Begg who collected the specimens. Many
people have helped in many ways and grateful acknowledgement is made to Mr. A. J.
Dartnall, Dr. R. Fox, Mr. M. Gillam, Dr. R. Pengilley and Mr. R. Wells. I am also
grateful to Dr, H. Cogger, of the Australian Museum, Sydney, who made the paratype
specimen available to me after preservation.

REFERENCES

BOULENGER, G. A. (1893). Catalogue of Snakes in the British Museum (Natural
History). British Museum (N.H.), London, (1961 reprint, Cramer, Weinheim; 382 pp.).

BOULENGER, G. A. (1912). A Vertebrate Fauna of the Malay Peninsula : Reptilia and
Batrachia. Taylor and Francis, London; 294 pp.

COGGER, H. G. (1975). Reptiles and Amphibians of Australia. A. H. and A. W. Reed.
Sydney; 584 pp.

FITZSIMONS, V. F. M. (1962). Snakes of Southern Africa. Macdonald, London; 423 pp.

GOW, G. F. (1976). Snakes of Australia. Angus and Robertson, Sydney; 88 pp.

McDOWELL, S. B. (1975). A Catalogue of the Snakes of New Guinea and the Solomons.
with Special Reference to Those in the Bernice P. Bishop Museum. Pt. II Anilioidea
and Pythoninae. J. Herpetol. 9 (1): 1-79.

ROOIJ, NELLY DE. (1917). The Reptiles of the Indo-Australian Archipelago, II. Ophidia.
Brill, Leiden; 334 pp.

TAYLOR, E. H. (1965). The Serpents of Thailand and Adjacent Waters. Univ. Kansas Sci.
Bull. XLV (9), 609-1096.

Aust. Zool. 19(2), 1977 139

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Tail Regeneration and Other Observations in a
Species of Agamid Lizard

C. J. HARDY and C. M. HARDY
Cranbrook School, Bellevue Hill, N.S.W. 2023

ABSTRACT

Evidence of tail regeneration is reported for the first time in an agamid lizard
contrary to the long held belief that lizards in this family are unable to regenerate
damaged tails. This evidence was obtained in a study of the common Australian
agamid lizard, Physignathus /esueurii lesueurii. Observations are also reported on
mass-length relationships, food, and a lower temperature limit for activity for
this lizard.

INTRODUCTION

The family Agamidae (dragon lizards) contains about three hundred species
which are found in many of the warmer areas of the world including Australasia,
and forty of these species occur in Australia. Agamid lizards have up to the
present been considered to be unable to regenerate the tail once it is broken,
compared with many other families of lizards, e.g. Geckonidae ( geckos ), Pygopodidae
(legless lizards), and Scincidae (skinks ).

During a study of the eastern water dragon (Physignathus lesueurii
lesueurii) in suburban and country habitats several specimens showed evidence
of new growth on damaged tails, and a few specimens were selected for detailed
study in captivity. The resulting observations on tail regeneration, mass-length
relationships, food, and a possible lower temperature limit for activity, are
reported as being of interest to the study of this and other species of the Agamidae
family in Australia and in other parts of the world.

DESCRIPTION AND HABITATS OF THE EASTERN WATER DRAGON

The type locality of the eastern water dragon is given by Worrell (1963)
as Queensland, but it is also found close to water courses east of the Great
Dividing Range from southern New South Wales to Cape York and it has also
been reported in New Guinea. The lizard is brownish-grey to olive-green on the
upper surface with black bands on each side of the ventral line and a broad black
stripe from the eye to the temporal region on both sides. The belly in both sexes
in mature specimens may be bright red. The lizard is reported to grow to a
length of 1 metre. It has a large angular head, long strong hindlimbs, and a

‘Present Postal Address: 12 Brassie St., N. Bondi, N.S.W. 2026

Aust. Zool. 19(2), 1977 141

C. J. HARDY and C. M. HARDY

1: Specimen III (Table 2). a0

Figs. | & 2. Eastern water dragon with regenerated tail. 1:
e up of regenerated tail.

Sept. 1974 photographed on 100 mm square grid. 2: Clos

442 Aust. Zool. 19(2), 1977

TAIL REGENERATION IN AN AGAMID LIZARD

bicarinated (doubly-keeled) tail which may be up to 2.5 times the body length,
and which is compressed laterally for swimming. It is a shy animal and if disturbed
takes refuge under rock ledges or in water, in which it can remain submerged
for long periods. Worrell (1958) described the behaviour of specimens of this
lizard along river banks and observed young lizards sunning themselves outside
the nesting burrows in which they hatched from eggs after an incubation period
of about three months.

A small colony of eastern water dragons was observed regularly for three
years in a small reserve in the densely populated suburb of Vaucluse in Sydney.
In this habitat a small stream passes over a waterfall in a cliff face in which
there are many rock ledges on which the lizards bask. The colony has suffered
considerable disturbance during this period due to workmen clearing the rough
native undergrowth and large areas of alien lantana (Lantana spp.) to landscape
the slopes with large native plants. The removal of many of the small plants
has reduced the plant food available to the colony and has exposed the lizards
to predators, particularly large birds such as kookaburras (Dacelo gigas). However,
the colony was still breeding in 1975 and four very young lizards were seen,
although two large mature specimens seen in 1974 had apparently disappeared.
The maximum number of specimens seen at any one time over the three years
was ten.

Specimens were also observed and photographed at a large colony at Jenolan
Caves, about 150 km west of Sydney in New South Wales. This colony is distributed
over a wide area along a river through the caves and State Park and some of the
lizards have grown accustomed to the public approaching them to within a few
metres.

RESULTS AND DISCUSSION
MASS-LENGTH RELATIONSHIPS

During the three years eleven specimens were captured, weighed, measured,
and photographed in close-up for reference in case of recapture. Five of these
had perfect tails, and two had apparently had the tip broken off. Three specimens
had distinct changes in tail thickness and appearance part way along the tail
when captured, with evidence of regenerated growth of 38, 52, and 75 mm.
One specimen in the colony at Jenolan Caves had two tails, a perfect main tail
399 mm long and an accessory tail 67 mm long growing from a point at the side
of the main tail, 108 mm from the anus. The mass-length data are given in Table 1
together with data for five newly hatched dragon lizards reported by Longley
(1947), but for which no masses were given.

The total length per g for perfect specimens decreases markedly with total
mass, and therefore with age, e.g., from 18-26mm.g* at 9-1lg to 1.2-1.4
mm.g? at 446-665 g. The ratio of tail length to body length for five specimens
with perfect tails was in the range 1.6-2.6, whereas the ratio for five specimens
with single regenerated tails was in the range 1.1-1.6.

Aust. Zool. 19(2), 1977 143

C. J. HARDY and C. M. HARDY

TABLE 1
MASS-LENGTH DATA FOR SPECIMENS IN NATURAL HABITAT

Length of Total
Total Body regenerated length
Mass length length tail per g es
g mm mm mm mm.g™'
9 234 65 — 26 Perfect tail
1] 200 Te — 18 Perfect tail
4] 300 100 —_ 7.3 Perfect tail
102 285 135 52 — Specimen I (Table 2)
136 380 156 == — Specimen II (Table 2)
156 556 157 67 = Double tail
270 480 185 aS —= Specimen III (Table 2)
446 613 213 — 1.4 Perfect tail
596 558 236 — — Tip of tail broken
660 2570 250 38 —
665 790 235 a 12 Perfect tail
_ 127 38 a= = 5 specimens at birth:

Longley (1947)

t For perfect specimens

The body length (BL, mm), defined as the length from the tip of the nose
to the anus, correlates very well with the total body mass (M,g). The data points
for the 11 specimens are fitted by a curve:

BL = M°? + 34
with a standard deviation of 5 mm. Seven of the calculated values of BL from this
equation are within + one standard deviation of the measured values, and the
maximum deviation of a calculated from a measured value is two standard
deviations.

There is a considerable degree of scatter in the correlation of total length
(TL, mm) with mass, but the data can be fitted by a curve:
OE ese ILS
with a standard deviation of 74mm. The much larger degree of scatter compared
with the correlation of BL and M is due to the variability of tail length which
in turn arises from the different extent of tail loss and extent of regeneration for
different specimens.

TAIL REGENERATION

A specimen captured for study in May, 1973 broke off a piece of its tail
a month later. This specimen has now been kept in captivity for over three
years in a large outdoor cage containing large slabs of rock, a pool, and areas
of native plants, and its tail growth and behaviour with respect to temperature

444 Aust. Zool. 19(2), 1977

TAIL REGENERATION IN AN AGAMID LIZARD

a) 10mm

Figs. 3-5. X-radiographs of regenerated tails. Arrows mark start of regenerated growth.
Conditions: 0.06sec. exposure at 75mA and 60KV with stationary grid to
collimate beam. 3: Specimen I (Table 2). 4: Specimen III (Table 2). 5: Double-
tailed specimen (Table 1).

Aust. Zool. 19(2), 1977 145

C. J. HARDY and C. M. HARDY

variations studied periodically. After the tail broke the specimen had a mass of
136 g and a length of 380 mm (Table 1) and several months later it showed new
growth at the end of the tail. After one year the new tail section was 65 mm
(Table 2). The new tail had smaller scales than those on the old tail and was
a uniform dark brown colour, not striped with lighter coloured bands as on the
old tail, or on the whole length of perfect tails of other specimens. Clear differences
between the appearance of regenerated and original tails of lizards in the families
of geckos, skinks, and legless lizards are well established (Bustard, 1970).

TABLE 2
RATE OF TAIL REGENERATION FOR THREE SPECIMENS IN CAPTIVITY
Length of Rate of increase
Specimen Mass Date regenerated of length of
(Table 1) g tail regenerated tail
mm mm.y~’
I 102 Delius yd =
106 8.2.76 112 280
Il 136 1529573 0 se
156 23314. 65 78
256 198 75 79 17
DO, PAG 93 13
Ill 270 22.8.74 152 ra
267 22 AVS 102 eas
Boil, T2276 109 34

* See Figure 1 for photograph of tail.

The rates of tail regeneration for three specimens studied in captivity are
given in Table 2. Specimen I gained only 4g in mass (4%) in ten weeks in
captivity, but it increased in total length by 69 mm (24%) and in tail length
by 64mm (43%). Most of this tail growth was in the regenerated section of
the tail, which increased by 60 mm (115%) compared with 4 mm on the original
tail (4%). This very large rate of growth may be due to this specimen having
broken its tail much closer to the body than where the breaks occurred with the
two other specimens and therefore it having a greater area of blood supply to
the new tail.

Specimens II and III (Table 2) did not increase smoothly in mass with
time. Specimen II increased in mass rapidly in the first eighteen months in
captivity and its tail grew rapidly in the first year. It decreased in mass by 11%
in the last year but increased in length in this time and added 14mm of new
growth to its tail. Specimen III showed tail growth during the first year in
captivity even though its mass remained constant. These observations indicate
that these lizards were able to redistribute mass from the body to new tail

146 Aust. Zool. 19(2), 1977

TAIL REGENERATION IN AN AGAMID LIZARD

growth during periods when they may have had insufficient food, or a poor variety
of food, available to maintain a steady increase in mass.

The specimen with a double tail (Table 1, 156g) probably developed the
accessory tail following damage sufficiently deep to expose the spinal cord. This
possible mechanism has been discussed briefly for other species of lizards by Goss
(1969) who stated that lizards regenerate an unsegmented cartiliginous tube
within the tail instead of vertebrae.

Accordingly, X-radiographs were taken of three of the lizards with regenerated
tails, and prints of these are shown in Figs. 3-5. The regenerated parts of the
tails can be clearly seen to contain a cartiliginous tube, as compared with the
original tails in which normal vertebrae can be seen. The X-radiograph of the
double-tailed specimen shows that the vertebra from which the accessory tail
emerged has a central diameter of 2.5mm compared with 2.0mm for the
vertebrae on either side and has the same density on the negative print as the
cartiliginous tube. It appears that a sheath of cartilige formed around the vertebra
and extended into the accessory tail which is therefore strongly attached to the
main tail. This thickening of the cartiliginous tube around the vertebra at the
point of breakage can also be seen in the X-radiographs of the two other specimens.

LOWER TEMPERATURE. LIMIT FOR ACTIVITY

One drazon lizard (Specimen II) was observed in relation to its appearance
from, and return to, hiding on sunny and dull days. The temperature was recorded
at hourly intervals over a period of two weeks in May, 1973 and the time
noted when it emerged from and returned to its overnight hiding place. The
lizard basked on the rocks during the day whether the sun was shining or the
sky was overcast provided that the temperature was above about 17°C.
Observations on the specimens in captivity and in the small colony in Vaucluse
indicate that the lizards hibernate in this area in the winter months because the
temperature is usually below 17°C.

In summer months the temperature at night is usually over 17°C on the
east coast of Australia. The lizard then shows the normal daytime behaviour of many
species of lizard of basking in the sun, and when the ambient temperature is high,
in order to raise its body temperature before foraging for food, and then returning
to shade or to its hole to prevent overheating or to escape from predators.
Bustard (1967) reported that the variegated gecko (Gehyra variegata) often
stopped foraging for food when the temperature dropped below 18°C, and
studies on other species may show that they also have a similar temperature limit
for activity.

OBSERVATIONS ON FOOD

The authors of many general books on Australian lizards have stated that
the eastern water dragon takes a wide variety of food. Davey (1970) quoted
mice, young rats, snails, grubs, worms, crustaceans, native flowers and berries.
Observations of food taken in the wild are very difficult with this lizard because

Aust. Zool: 19(2), 1977 147

C. J. HARDY and C. M. HARDY

of its shy nature, and because it forages for food in crevices in rocks and within
vegetation.

Samples of faeces freshly deposited by large lizards in the wild were therefore
collected and analysed. These were usually: firm cylindrical pieces about 5 mm
diameter and 10-15mm long. After they were dried the components could be
separated with tweezers and grouped into animal and plant remains. A large
part consisted of the hard indigestible parts of beetles and other insects, e.g. legs
and wing cases. There were also many small hard seeds, a large number of which
were from lantana plants. Pieces of fern and other small plants were found, as
well as powdery white lumps of dried salts. An unusual discovery in one sample
was a 200 mm length of nylon thread. Young dragons observed near their nesting
burrows spend most of the day picking up small flies and other small insects
within easy reach, but have not been seen to take plant food.

In captivity these lizards will eat a variety of food, small pieces of meat,
live insects (beetles, flies, cicadas), snails, worms, and whole or chopped fruit
such as tomatoes and strawberries, of which the latter appear to be very attractive.

BODY COLOURATION

The eastern water dragon develops a bright red belly colouration in early
summer after it has shed the old skin which is usually a dull-red-brown in that
region. Both sexes are said to develop the colour. However, it is not possible to
differentiate between the sexes in the field, and this can only be done reliably by
a careful examination of the anal slit. Generally the whole of the underside from
between the forelegs to between the hindlegs is uniformly coloured red. One of
the specimens captured, a large mature dragon (660g), had a ring of deep
red-brown skin around the belly with the centre the normal grey-green colour.

It is not clear, from the few specimens studied, at what age or mass the
red colour develops. The 106g Specimen I (Table 2) in early February, 1976
was a deep red colour over the whole belly region, with the colour suffused into
the legs and under the throat. The larger specimen (156g, Table 1) with the
double tail had only a slight tinge of red at the same date, and the much
larger Specimens II and III (227 and 327g, Table 2) both had distinctly red
coloured bellies, but no colour on legs or throats.

ACKNOWLEDGEMENT
We are grateful to Dr. A. D. Tucker for taking the X-radiographs of the specimens.

REFERENCES

BUSTARD, H. R. (1967). Activity Cycle and Thermoregulation in the Australian Gecko
Gehyra variegata. Copeia 1967, 753.

BUSTARD, H. R. (1970). Australian Lizards, Collins, Sydney.

DAVEY, K. (1970). Australian Lizards, Lansdowne Press, Melbourne.

GOSS, R. J. (1969). Principles of Regeneration, Academic Press, London, 214.

LONGLEY, G. (1947). Notes on the Hatching of Eggs of the Water Dragon. Proc. Roy.
200. SOC. NS WEY, -.25-

WORRELL, E. (1958). Song of the Snake, Angus and Robertson, Sydney.

WORRELL, E. (1970). Reptiles of Australia, Angus and Robertson, Sydney.

148 Aust. Zool. 19(2), 1977

Electrophoretic Evidence for the Specific Status of the
Lizards Lampropholis guichenoti (Dumeril & Bibron,
1839) and L. delicata (de Vis, 1887)

BELINDA F. HARRIS and P. G. JOHNSTON*
School of Biological Sciences, Macquarie University, North Ryde, New South Wales, 2113

ABSTRACT

Electrophoretic differences in two of the five enzymes studied indicate that
sympatric populations of Lampropholis guichenoti and L. delicata are maintaining
separate gene pools and should be recognised as separate species. Coloration
was found to be the most useful feature for distinguishing the two species in
the field.

INTRODUCTION

Lampropholis guichenoti (Dumeril & Bibron, 1839) and L. delicata (de Vis,
1887) are small ground lizards of the family Scincidae. In his original description
of L. delicata, de Vis (1887) stated that the differences between it and L. guichenoti
were “enlarged preanals . . . slender form, feebler limbs and completely different
physiognomy”’.

Loveridge (1934) grouped the two together as subspecies L.g. guichenoti
and L.g. delicata. Worrell (1963) supported this conclusion but applied the
subspecies name delicata only to the type specimen from Warra, Queensland and
to some specimens from Groote Eylandt.

Cogger (1975) listed them as separate species distinguishable by their
coloration. He noted that they had similar scalation, size and habits and that .
their distributions overlapped considerably.

Clark (1965) attempted to differentiate between the two species using scale
counts, body proportions and coloration. She found that the ranges of scale
counts and body proportions were similar and decided that only coloration and
the number of supraciliaries could be used to separate the two species.

On morphological grounds it is clearly difficult to distinguish between these
putative species. Coloration appears to be the most reliable character but even
this shows considerable variation.

Electrophoretic methods have been used successfully to investigate taxonomic
problems in a wide range of species (see Avise, 1974). It was therefore decided

*Reprint requests to Dr. P. G. Johnston

Aust. Zool. 19(2), 1977 149

BELINDA F. HARRIS and P. G. JOHNSTON

to employ electrophoresis as well as morphology to investigate sympatric
populations of these lizards with the aim of clarifying their taxonomy.

MATERIALS AND METHODS

Sixty adult lizards were collected in a suburban garden in Epping, N.S.W.
They were divided into two groups according to their coloration. One group
classified as L. delicata had a definite pale stripe above the dark lateral stripe
with a pale stripe below the lateral stripe less obvious or missing. The other
eroup classified as L. gwichenoti had a pale stripe below the dark lateral stripe
which was at least as obvious as the pale stripe above the lateral stripe. Using
these criteria 33 specimens were classified as L. guichenoti and 23 as L. delicata.
It was not possible to classify four individuals using these criteria.

MORPHOLOGY

The supraciliaries and 4th toe lamellae were counted and snout-vent length
measured. The amount of overlap of adpressed limbs and the size of the preanals
were also examined on some specimens. The presence or absence of a dark
vertebral stripe, pale stripes above or below the dark lateral stripe and small
pale and dark spots on the back was noted.

ELECTROPHORESIS

Livers were removed and homogenised in an equal volume of distilled water,
using a micro mortar and pestle. The homogenates were then applied to starch
gels or cellulose acetate ‘“‘cellogel” (Chemetron, Milan). Electrophoresis conditions,
buffers and stains for lactate dehydrogenase (LDH), malate dehydrogenase (MDH)
and phosphoglucomutase (PGM) are described in Johnston (1973). The methods
of Shaw and Prasad (1970) were used for detecting esterases and a glycerophosphate
dehydrogenase (a GPDH).

TABLE 1

Length and scale counts of L. guichenoti and L. delicata

n x s range
Supraciliaries L. guichenoti 35 6.58 0.91 5-9
L. delicata #68 6.41 0.84 5-8

4th toe lamellae L. guichenoti Be Ail 1.34 18-24
L. delicata DA 24.83 1.15 22-26

Snout vent length (mm) L. guichenoti 27 40.30 3.30 34-46
L. delicata 20 37.28 2.69 31-41

150 Aust. Zool. 19(2), 1977

ELECTROPHORETIC DIFFERENCES IN LIZARDS

PGM

Pee a ilgiuicke. akg:

Fig. 1. Phosphoglucomutase patterns of L. guichenoti (L.g.) and L. delicata (L.d.) after
cellogel electrophoresis.

RESULTS

MORPHOLOGY

Table 1 gives the mean, standard deviation and range of values for the
characteristics measured. Student’s ‘“‘t” test showed that the number of 4th toe
lamellae and snout-vent length differed significantly between the two groups at
the 5% level of significance. The difference in the number of supraciliaries was
not significant.

No difference was observed in the size of the preanals or amount of limb
overlap between the two groups.

COLOUR

Table 2 gives the percentage of lizards in each group with each of the
coloration features studied. All L. delicata had a pale stripe above the dark
lateral stripe, but no dark vertebral stripe or pale spots on the back. L. guichenoti
was more variable. The majority of them had a dark vertebral stripe, pale stripes
above and below the dark lateral stripe and pale spots on the back. However,
none of these features was possessed by all the lizards in the group.

Aust. Zool. 19(2), 1977 151

BELINDA F. HARRIS and P. G. JOHNSTON

TABLE 2

Percentage of lizards showing coloration features

L. guichenoti (n=37)

. delicata (n=23)

Dark vertebral) stripes tee ee 86%
Pale stripe above lateral’ 3... ee oe 83%
Pale>stripe: below slaterali- 95%
Pale, and dark spots onbacke. 28.0. 92%
ELECTROPHORESIS

0%
100%
17%
0%

Phosphoglucomutase: L. delicata and L. guichenoti showed different electrophoretic
patterns for PGM. In all cases the band of L. delicata was faster migrating and
was easily distinguished from the slower migrating band of L. guichenoti (Fig. 1).
Esterases: The electrophoretic patterns observed were complex. However each
species had a distinct pattern and could be easily classified (Fig. 2).

ESTERASES

oa

iG GE 510 Pian al ie A, sa Lgl Ok

Fig. 2. Esterase patterns of L: guichenoti and L. delicata after starch gel electrophoresis,

152

Eo

Aust. Zool. 19(2), 1977

ELECTROPHORETIC DIFFERENCES IN LIZARDS

Malate dehydrogenase, lactate dehydrogenase and a glycerophosphate dehydro-
genase: These enzymes were monomorphic and had identical electrophoretic
mobilities in both species. The four lizards which could not be classified on the
basis of their colour patterns all had PGM and esterase patterns typical of
L. guichenoti. These lizards were treated as L. guichenoti for the morphological
studies.

DISCUSSION

The morphological data showed no significant difference between groups in
the number of supraciliaries. The two characters that showed significant differences
between groups were snout length and the number of 4th toe lamellae. These
results contrast with those of Clark (1965) who found a significant difference
between species in the number of supraciliaries but not in the number of 4th toe
lamellae. It is thus apparent that these characters show considerable intra and
interpopulation variation in both groups and are therefore not satisfactory for
differentiating the two groups.

As has been found by other workers coloration is the most useful method
for identification in the field. However, like the other morphological characters
that have been used, there is some overlap and it was not possible to classify all
the specimens on their colour patterns. Electrophoresis, on the other hand, clearly
separates the lizards into two groups with no overlap. The PGM and esterase
patterns were distinctive for each group. It has been established that PGM is
a monomer in a wide variety of species (see Manwell and Baker 1970). Double
banded phenotypes representing heterozygotes would therefore be expected if
gene exchange was taking place between these groups. No double banded pheno-
types were observed in any of the lizards examined.

The interpretation of the esterase variation is not so straightforward because
of the unknown number of loci involved, but as with the PGM results, there is
no evidence of heterozygotes being formed between the two groups. The electro-
phoretic results indicate that two groups are maintaining separate gene pools and
that reproductive isolation exists between them. These reults support the con-
clusion of Clark (1965) and Cogger (1975) that L. guichenoti and L. delicata are
separate species.

REFERENCES
AVISE, J. C. (1974). Systematic value of electrophoretic data. Syst. Zool. 23, 465-481.

CLARK, C. J. (1965). A comparison between some Australian five-fingered lizards of the
genus Leiolopisma Dumeril & Bibron (Lacertilia: Scincidae). Aust. J. Zool 135. 51d oD

COGGER, H. G. (1975). Reptiles and Amphibians of Australia, A. H. and A. W. Reed.

Sydney.
DE VIS, C. W. (1887). A contribution to the herpetology of Queensland. Proc. Linn. Soc.
N.S.W. 2, 820.

JOHNSTON, P. G. (1973). Variation in island and mainland populations of Potorous
tridactylus and Macropus rufogriseus (Marsupialia). Ph.D. thesis, University of N.S.W.

Aust. Zool. 19(2), 1977 153

BELINDA F. HARRIS and P. G. JOHNSTON

LOVERIDGE. A. (1934). Australian Reptiles, Bull. Mus. Comp. Zool. Harvard 77, 243-383.

MANWELL, C. & C. M. A. BAKER (1970). Molecular Biology and the Origin of Species:
Heterosis, Protein Polymorphism and Animal Breeding. Sidgwick and Jackson, London.

SHAW, L. R. & R. PRASAD (1970). Starch gel electrophoresis of enzymes — a compilation
of recipes. Biochem. Genet. 4, 297-320.

WORRELL, E. (1963). Reptiles of Australia. Angus and Roberson, Sydney.

154 Aust. Zool. 19(2), 1977

Voluntary Submergence Time and Breathing Rhythm in
the Homalopsine Snake, Cerberus rhynchops

HAROLD HEATWOLE
Department of Zoology, University of New England, Armidale, N.S.W., 2351, Australia

Characteristics of breathing and submergence were analyzed for captive Cerberus
rhynchops, a homalopsine aquatic snake. Some aspects of diving behavior
differed markedly from that of sea snakes. Inactive snakes either hid in tubes
on the bottom of the aquarium or rested with the head at the surface and the
nostrils in air. Activity resulted in a reduction of submergence time. Within a
given activity category, no diel difference in submergence time was observed,
differences among individuals were large, and oxygen and CO, content of the
water had only a weak effect on submergence time. Within the range 20°-30° C
submergence time was only weakly temperature-dependent. Number of breaths
per breathing episode showed great differences among individuals but little
dependence upon time of day or gaseous content of the water. Individuals sub-
merged for long periods tended to take a greater number of breaths per
breathing episode.

Snakes at the surface for prolonged periods breathed regularly with the interval
between breaths being longer than when they were alternatively surfacing and
diving. Interbreath interval was independent of O, and CO, content of the water
or of temperature between 20° and 33°C.

INTRODUCTION

Except for a few species scattered throughout various other taxa, the truly
aquatic snakes belong to 5 major groups in 4 families. These are the sea kraits'
(family Laticaudidae), the sea snakes’ (family Hydrophiidae), the file snakes
(family Acrochordidae) and two subfamilies of the Colubridae, the Homalopsinae
and the Natricinae. Each of these five groups has probably developed their aquatic
habits independently of the others and consequently it is of evolutionary signifi-
cance to compare their diving adaptations.

Various aspects of the diving physiology and behavior of representatives of
four of these groups have been investigated (Johansen 1959, Murdaugh and

1. The two families Laticaudidae and Hydrophiidae are sometimes lumped with their terres-
trial relatives into the single family Elapidae (McDowell 1969) but were historically treated
as subfamilies of the family Hydrophiidae (Smith 1926). It is now clear that they represent
two independent marine radiations from the elapids and treatment as two separate families,
as recently done by Burger and Natsuno (1974) seems the most appropriate procedure.

Aust. Zoo!. 19(2), 1977 155

HAROLD HEATWOLE

Jackson 1962, Pough 1973, Standaert and Johansen 1974, Graham 1974, Graham
et al. 1975, Heatwole and Seymour 1975a, 1975b, 1976, Heatwole 1975, 1977,
Heatwole e¢ al. in press, Seymour 1974, 1976, Seymour and Webster 1975, Jacob
and McDonald 1976, Irvine and Prange 1976). The only previously uninvestigated
group is the Homalopsinae. The present paper broadens the base of comparison by
providing information on several aspects of diving and breathing in a homalopsine
species, Cerberus rhynchops.

All of the 10 genera and 36 species of homalopsines are aquatic and have
anterodorsally located nostrils which are lunate in shape and are valvular. Most
are restricted to freshwater whereas others occur in brackish or marine situations.
Cerberus rhynchops frequents tidal rivers and estuaries, both in the upper reaches
where the water is fresh and in salt waters along coasts. It is widely distributed
in suitable habitat from India throughout southeast Asia, Sunda Islands, Philip-
pines, and into the Pacific to Palau (Gyi 1970). The animals used in the present
study were collected in August-September 1975 in mangrove swamps at the United
States Naval Base at Subic Bay, Republic of the Philippines, and at Jagoliao Island,
Bohol, Republic of the Philippines. At the former locality, snakes were active at
night in water 5 to 20 cm deep. The same area had been previously searched by
day and no snakes observed.

MATERIALS AND METHODS

Except for an experiment on the effect of temperature on diving time, which
was carried out in October, 1975, in Armidale, Australia, and using a snake that
was transported there from the Philippines, all experiments and observations were
carried out aboard the RV Alpha Helix in Philippine waters in September 1975.

Several limitations were imposed on the study by the animals and by the time
available: (1) Individual snakes varied significantly from each other in respect
to a number of characteristics and hence data from different animals could not be
grouped for comparing experimental procedures. (2) Not all snakes engaged in
all the types of behavior studied, and consequently some types of comparisons
had to be omitted for some animals. (3) The submergence times were long and
accumulation of a sufficient number for valid comparisons was time-consuming;
available time was limited. Thus, some sample sizes are smaller than desirable.

The snakes were housed in 60-litre aquaria containing 20-35 cm of sea water
maintained at a temperature of 25-28° C. Opaque polyvinyl tubes 3 cm in inside
diameter and 40 cm long were provided as shelter for the snakes in the bottom
of the aquaria. Lights in the laboratory were turned on at 0700 hrs and off at 1900
hrs each day. However, at night light from an adjacent laboratory entered through
a large window and was of sufficient intensity that observations could be made.
The snakes ranged in body weight from 93 to 222 g.

A total of 187 submergences on 6 individuals under a variety of different
conditions were timed; when the snakes were not submerged data was recorded

156 Aust. Zool. 19(2), 1977

SUBMERGENCE AND BREATHING IN A SNAKE

on the number and frequency of breaths. Two experiments were conducted to
ascertain the possible effect of dissolved gases on the submergence time and breath-
ing rhythm. In one of these, pure nitrogen was bubbled through the water of the
aquarium until the oxygen level was lowered. O» level prior to bubbling was 5.2
ppm (measured by a YSI Model 57 oxygen meter) but dropped to 2.8 ppm after
1 hr 15 min of bubbling. Bubbling was discontinued during timed submergences
but was maintained between observation periods, in some cases an entire day. O>
levels in the aquarium were between 1.3 and 2.8 ppm during timed submergences;
at the completion of the experiment O. was 1.4 ppm. By the following morning
it had risen only to 1.6 ppm in the absence of bubbling.

A second experiment involved the same procedure except instead of nitrogen
a gas mixture of 600 ppm CQ, in nitrogen was bubbled through the water in
order to raise the CO, level as well as to deplete oxygen. O» levels during timed
submergences ranged from 1.4 to 1.6 ppm and after bubbling was discontinued
had risen only to 3.9 ppm overnight.

The room in which these experiments were carried out was temperature-
controlled and ventilated. There were no decreases in O> levels of the air above the
water greater than 0.1 ppm and usually values were the same there as elsewhere
in the room. Thus, any effect on submergence time or breathing rhythm would be
a result of the change in dissolved gases rather than of alteration of composition
of inspired air.

The temperature experiment carried out in Australia involved keeping a snake
in an aquarium as described above but periodically altering the water temperature
and then recording submergence times and breathing characteristics.

Time is expressed in minutes and tenths of minutes (rather than minutes
and seconds) to facilitate calculation of means and standard errors.

RESULTS

GENERAL BEHAVIOR

When snakes were placed in the aquaria, they swam actively for several
hours in what appeared to be escape behavior. However, by the end of the first
day, all had found the tubes and entered them, alternating periods of swimming
with periods of retreat into the tubes. Usually the head was just at the entrance
of the tube and facing out, or the head and neck protruded from the pipe by a
distance of up to 15 cm. Initially, disturbance such as sudden movements or sub-
strate vibrations would result in the snake jerking backwards into the tube with
a tapidity unexpected from the slow, deliberate movements generally characterizing
this species. The first night one animal escaped from the aquarium and moved
rapidly and as agilely as a terrestrial snake over a rather smooth sink top when
the investigator entered the room. By the third day, overt fright responses were
seldom observed (and then only in response to some unusual stimulus) and the
snakes seemed accustomed to movement of people in the laboratory.

Aust. Zoo!. 19(2), 1977 157

HAROLD HEATWOLE

BREATHING

When C. rhynchops breathes, it first exhales, then inhales and finally aia
exhales. There is then a brief, or a prolonged period of apnea (depending on
circumstances) before the next breath. Breathing movements are usually easily
detectable by observing expansion and contraction of girth of the anterior part of
the body. On some occasions when breathing was shallow it was difficult to detect
individual breaths and they could not be counted.

The initial exhalent phase seems to be the active one; it is more rapid than
the subsequent inhalent and second exhalent phase and sometimes the snake’s
body sways with the effort.

When a snake emerges it remains a short time with the nostrils out of the
water before it takes a breath (mean 0.32 min; range 0.10-0.67 min). Subse-
quently, it breathes at short intervals, maintaining its nostrils continuously above
water for the entire series of breaths. This series of breaths is called a breathing
episode and the time elapsing between two consecutive breaths in that
episode, the inter-breath interval. After the last breath of the episode, the animal
almost immediately submerges (the time elapsing between the last breath and
submergence averaging 0.06 min; range 0.03-0.27 min). The time between breaths
of an inactive snake remaining eentinionl: at the surface is also referred to as
the inter-breath interval.

ACTIVITY AND INACTIVITY

) During active periods, the snakes slowly swam along the bottom of the
aquarium, poking the nose along the crevices at the sides of the plastic tubes or
along the edges of the aquarium, or swam at the surface with occasional forays
to the bottom and back. Movements were slow and deliberate. Activity took place
chiefly at night. Snakes interspersed prolonged periods of inactivity with activity.
There was considerable individual variation in level of activity, one animal seldom
engaging in more than momentary periods of activity during the study; others
were active much of the night with only occasional periods of rest.

There were two distinct behavioral patterns adopted by inactive snakes. One
was to lie motionless in the tube except for periodic trips to the surface to breathe.
Even during breathing the snake would not completely leave the tube but would
extend the head and neck just far enough to protrude the nostrils and eyes above
the surface. Upon termination of breathing, the snake backed slowly into the tube
and again became motionless. While the head was on the surface, the body and
neck stretched toward the surface either nearly vertically or at an angle of about
45°. The body was held straight without conspicuous bends or curves.

The second posture adopted during inactivity was to remain motionless at
the surface with the nostrils and eyes emergent. The body extended to the bottom
of the aquarium and the posterior part and the tail trailed on the bottom in a
sinuous curve; in some cases the tail and part of the body were in a tube. Occasion-
ally a snake would lower the head into the water and look around or even sub-

158 Aust. Zool. 19(2), 1977

SUBMERGENCE AND BREATHING IN A SNAKE

merge briefly so that the head and anterior part of the body pointed upward from
the bottom of the aquarium at about an angle of 45°, with the head at about
mid-water. Bouts of actvity were interspersed with this type of inactivity more
frequently than with inactivity in shelter, and emergent inactvity is probably an
indication of an alert state bordering on activity. Individual snakes differed greatly
in their preference for inactivity at the surface as opposed to inactivity in shelter.
Some snakes never were observed at the surface except for brief periods of
breathing; others spent most of their inactive time with their heads at the surface.

VOLUNTARY SUBMERGENCE TIME

During the first two days in the aquarium, mean submergence times for a
given activity category were low (31-37% those of the same individuals after
3 or more days) and no values even of inactive snakes, exceeded 28 mins.

Underwater time of individual active snakes averaged 0.7-3.5 min, or 3.3
— 26% of the inactive values for the same snakes under comparable conditions.
There was one exception in which active values were 84.4% of inactive ones:
this exception occurred, however, during the second day in the experimental
container when inactive values were abnormally low, probably because of the
unfamiliar conditions. Pairing active and inactive means by animal, date and
experimental conditions for all periods in which both inactivity and activity
occurred, a Wilcoxon Matched Pairs Signed Ranks test revealed that the above
differences were highly significant (P < 0.005, one-tailed; 8 pairs of means).
Thus, actvity greatly affects submergence time.

There was very little difference in submergence time between day and night
for a given activity category and set of conditions. It was predicted that since
the species is nocturnal, submergence times would be shorter at night. However,
pairing day and night mean values by animal, activity category, date and type of
experiment, a Wilcoxon Matched Pairs Signed Ranks test indicated no statistical
diel differences (P > > 0.05, one-tailed, 9 pairs of means). Consequently values
for night and day were lumped for further analyses but including only data from
inactive snakes after the initial 3-day period of becoming accustomed to the new
environment.

TABLE 1. RELATION OF SUBMERGENCE TIME OF CERBERUS RHYNCHOPS
- TO GASEOUS CONTENT OF THE WATER

Mean Submergence Time in Minutes (= s.e.)

Water
Individual Water %p Bubbled %
Snake Untreated Bubbled _—__ Re- with CO, _ Re-
Designation N Water N with N, duction N in nitrogen duction
R Py 9 Soe t7 2 7 36.01 9 | 4 33.06 16

L 10 29,12+5.16 6 22.01 24 | ie ee. 2

Aust. Zool. 19(2), 1977 159

HAROLD HEATWOLE

ro -—_+-

VLVG ON

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J0OSId3 ONIHLV3SY¥8 / SHIV3SYNE JO ON

(NIW)

VLVG ON

SJWIL JONASYSAWENS

SPECIMEN CODE NUMBER

Aust. Zool. 19(2), 1977

160

SUBMERGENCE AND BREATHING IN A SNAKE

Under equivalent conditions, different individuals had markedly and signifi-
cantly different mean submergence times (Fig. 1). These differences bore no
consistent relation to body weight, e.g., the animal with the longest submergence
times was the smallest individual and the one with the shortest submergence times
was the second smallest snake. There did seem to be a general relationship with
tendency toward activity. Qualitatively it was observed that the snakes with
short submergences during inactive periods tended to be those which were most
frequently active.

Mean submergence time decreased in oxygen depleted water in both individ-
uals tested (Table 1). The differences were not statistically significant, however
(Mann-Whitney U-test), as P values were just slightly greater than 0.05 in both
cases. In water that was oxygen poor and COb rich, there was a further (but not
significant; P > 0.05) decrease in submergence times. Some of the differences
were near the level of significance and all were in the direction predicted. It is
possible that failure to achieve significance was largely a reflection of the small
sample sizes. At present, however, it can only be concluded that the differences
may have been due to chance. In any event, the reductions in submergence time
possibly arising from bubbling N: and/or CO: were of less magnitude than
individual differences and it can be concluded that if gaseous content of the water
does influence submergence time, the effect is not great.

Voluntary submergence time is surprisingly independent of temperature
within the range 20°-30° C. For the single individual tested at several tempera-
tures, mean submergence time did not differ significantly between 20° and 23.5° C
(Fig. 2) and even the value of the animal at 30° C was not greatly reduced from
that at the other two temperatures (differences between 23.5° and 30°C not
significant; Mann-Whitney U-test, P slightly greater than 0.05, one-tailed). The
shortest submergences timed were practically identical at all temperatures (Fig. 2).
However, the maximum values progressively decreased with increasing tempera-
ture. it seems that the snakes do not greatly alter their submergence times within
the range of temperatures tested but that there may be a tendency to dispense
with the exceptionally long ones at higher temperatures. Differences in submergence
times at different temperatures for a given individual were less than differences
among individuals at the same temperature (Fig. 2).

THE NUMBER OF BREATHS PER BREATHING EPISODE

Number of breaths taken when a snake surfaced to breathe varied from 1 to
17. There was considerable and significant variation among individuals, inter-
individual means varying almost 2-fold for a given set of environmental conditions

Fig. 1. Submergence times and number of breaths per breathing episode in individuals
R, L, J, B, and M of Cerberus rhynchops. Numbers above figures indicate number
of submergences or breathing episodes involved. Vertical line indicates range,
horizontal line the mean, and rectangles, 2 standard errors on either side of the mean.

Aust. Zool. 19(2), 1977 161

HAROLD HEATWOLE

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(NIN) AWIL J3ONS9YSWENS

162 Aust. Zool. 19(2),

TEMPERATURE

1977

SUBMERGENCE AND BREATHING IN A SNAKE

(Fig. 1). In contrast to submergence times, the number of breaths per breathing
episode did not show consistent differences between the periods before and after
the snakes became familiar with their new environment, nor did it vary signifi-
cantly between night and day (half of the day values were higher than night ones
and half were lower) or dissolved gaseous content of the water (Mann-Whitney
U-test, P > 0.10 in all comparisons). Insufficient data on this characteristic were
available for active snakes for evaluation to be made.

It appears that for a given individual, various environmental factors have
little effect on number of breaths per episode but that each individual has its own
characteristic in this regard and individuals differ. Inspection of the two sets of
figures in figure 1 indicates that the individuals that submerge the longest tend to
take a larger number of breaths than those which have shorter submergence times,
suggesting that there may be a relation between these two breathing characteristics.
However, when the number of breaths taken were related to the corresponding
previous submergence time for each individual snake, no significant correlations
were obtained (Spearman Rank Correlation Test; P >> 0.10 for each snake).
Thus it seems that the number of breaths taken does not depend upon the length
of submergence immediately preceding a given episode even though snakes that
submerge long times tend also to take a large number of breaths per breathing
episode.

INTER-BREATH INTERVAL

The intervals between the breaths of a given breathing episode (inter-breath
intervals) were very short; means were less than half a minute in all snakes and
averaged from 0.4% to 3.0% of the submergence times for given individuals. As
was true of other breathing characteristics, there were significant differences among
individuals in inter-breath interval (Fig. 3); difference among individuals varied
as much as 3-fold.

Those individuals that had long submergence times and high numbers of
breaths per episode tended to be those with shorter inter-breath intervals (compare
ites. dl “and 3h).

For a given individual, inter-breath interval was remarkably similar under a
variety of conditions. For example, bubbling nitrogen or nitrogen and carbon
dioxide through the water did not affect it (Fig. 3), nor did temperature (Fig. 2).

Inter-breath intervals were greatly different when inactive snakes remained
continuously on the surface as compared to the brief breathing episodes between

Fig. 2. Relation of submergence times and inter-breath interval of Cerberus rhynchops to
water temperature. Narrow figures as in Figure 1; all are of individual A. The
wide rectangle indicates the range in means of snakes R, L, J, B and M over the

_temperature range indicated by the width of the box. The dotted lines extending
the rectangle indicate the range of values.

Aust. Zool. 19(2), 1977 163

164

(MIN )

INTERVAL

INTERBREATH

6.0

HAROLD HEATWOLE
N 32

l2

R L J B

SPECIMEN CODE NUMBER

Aust. Zool. 19(2), 1977

SUBMERGENCE AND BREATHING IN A SNAKE

submergences (Fig. 3). For given individuals the inter-breath intervals between
the two situations varied by 2-3 fold, more rapid and deeper breathing occurring
in the latter case. As in other characteristics, inter-breath intervals of snakes
remaining at the surface showed significant variation among individuals and was
not affected by bubbling nitrogen through the water (Fig. 3).

The change between these two patterns of inter-breath interval when a snake
emerges and then remains on the surface for a long time, is rather abrupt. For
example, snake ““L” emerged and had deep breaths at intervals of 0.20 to 0.32
minutes (mean 0.26). It then abruptly began to engage in shallow breathing with
inter-breath intervals of 0.42-0.65 min (mean 0.51). The difference in inter-breath
interval between these two periods was significant (Mann-Whitney U-test,
P < 0.002). This observation also suggests that breathing frequency is related
to tidal volume.

Prolonged apnea in Cerberus is related to submergence. When snakes are at
the surface inter-breath intervals are short, yet there are some occasions when
snakes at the surface did suspend breathing for rather long periods (usually 1-4
mins although some longer ones were observed). These apneic periods were usually
coincident with some unusual disturbance in the laboratory and were probably
fright responses. They were not included in the calculations of mean inter-breath
intervals. Eleven such apneic periods occurred in snakes with the nostrils emergent
during the study.

DISCUSSION

Breathing rhythm of sea snakes has been studied by Heatwole and Seymour
(1975a, b, 1976) and Heatwole (1975). Breathing rhythm in Cerberus differs
from that of sea snakes in several important ways. (1) Sea snakes undergo pro-
longed periods of apnea not only when submerged but also when they are at the
surface, whereas Cerberus does so only when submerged (or when frightened)
and {2) sea snakes usually take only 1-3 breaths per breathing episode whereas
Cerberus takes a much larger number. The breathing rhythm of Cerberus when
emergent resembles that of land snakes in air and it would appear to be somewhat
intermediate in breathing rhythm between land snakes and sea snakes, or more
likely, has retained the land rhythm and can switch between it and one more
appropriate to aquatic animals depending upon circumstances. Sea snakes on the
other hand seem to have lost the characteristics of the land rhythm and have
exaggerated the apneic interval under all circumstances.

Fig. 3. Inter-breath interval of individuals of Cerberus rhynchops during breathing episodes
between submergences (solid figures), and when remaining on the surface for
eytended periods in normal water (open figures) and when nitrogen was bubbled
through water (cross-hatched figure). N indicates that nitrogen was bubbled through
the water, and C indicates that CO, in nitrogen was bubbled through the water.
Numbers above figures indicate number of submergences. R, L, J, B, and M refer
to code numbers of individual snakes; other symbols as in Figure 1.

Aust. Zool. 19(2), 1977 165

HAROLD HEATWOLE

The relatively slight (if any) effect dissolved gases have on submergence times
of Cerberus may be related to other respiratory features of this species. Sea snakes
are capable of cutaneous respiration to a greater extent than are land snakes. (Gra-
ham 1974, Standaert and Johansen 1974, Heatwole and Seymour 1975a, b, 1976).
Such a capability would seem to be advantageous to any aquatic snake which
remained submerged for extended periods. However, the relatively slight (if any)
effect dissolved gases have on submergence times in Cerberus suggests that its
aerial breathing rhythm is not greatly influenced by a capacity for underwater
respiration. Heatwole and Seymour (in prep.) have found that this species has
much lower cutaneous oxygen uptake than did four sea snakes tested under the
same conditions. The waters of mangrove swamps (an habitat of Cerberus rhyn-
chops) are often relatively depleted of oxygen and are rich in carbon dioxide. There
would be little selection for cutaneous respiration in such an environment and the
insensitivity of Cerberus to dissolved gases may be a reflection of its particular
ecology.

ACKNOWLEDGMENTS

This work was supported by grants from the Australian Research Grants Committee,
the Internal Research Funds of the University of New England and by National Science
Foundation (USA) grants DES 74-24129 to Pennsylvania State University and OFS 74-02888
and OFS 74-01830 to the Scripps Institution of Oceanography for the operation of the Alpha
Helix program (Visayan Sea Expedition). I am grateful to William Dunson, Sherman
Minton, and Renato Garcia for assistance in procuring specimens, to William Dunson, Roger
Seymour, and George Pickwell for criticism of the manuscript, to Susan Murdock for
assistance in the laboratory and tc Elizabeth Cameron, Viola Watt and Neva Walden for
aid in preparation of the manuscript.

LITERATURE CITED

BURGER, W. L. & T. NATSUNO (1974). A new genus for the Arafura smooth seasnake
and redefinitions of other seasnake genera. The Snake 6, 61-75.

GRAHAM, J. B. (1974). Aquatic respiration in the sea snake Pelamis platurus. Respir.
Physiol. 21, 1-7.

GRAHAM, J. B., J. H. GEE & F. S. ROBISON (1975). Hydrostatic and gas exchange
functions of the lung of the sea snake Pelamis platurus. Comp. Biochem. Physiol. 50A,
477-482.

GYI, K. K. (1970). A revision of colubrid snakes of the subfamily Homalopsinae. Univ.
Kansas Publ. Mus. Nat. Hist. 20, 47-223.

HEATWOLE, H. (1975). Voluntary submergence times of marine snakes. Marine Biology
32, 205-213,

HEATWOLE, H. (1977). Heart rate during breathing and apnea in marine snakes. J.
Herpetology, 11, 67-76.

HEATWOLE, H. & R. SEYMOUR (1975a). Pulmonary and cutaneous oxygen uptake in
sea snakes and a file snake. Comp. Biochem. Physiol. 51A, 399-405.

HEATWOLE, H. & R. SEYMOUR (1975b). Diving Physiology. Chapter 15 (pp. 289-327)
in: The Biology of Sea Snakes (ed. W. A. Dunson), University Park Press, Baltimore,
530 pp.

HEATWOLE, H. & R. SEYMOUR (1976). Respiration in marine snakes. pp. 375-389 in:
Respiration of Amphibious Vertebrates (ed. G. M. Hughes), Academic Press, London,
402 pp.

166 Aust. Zool. 19(2), 1977

SUBMERGENCE AND BREATHING IN A SNAKE

HEATWOLE, H., S. A. MINTON Jr., R. TAYLOR & V. TAYLOR. Underwater observa-
tions on sea snake behaviour. Aust. Mus. Rec., in press.

IRVINE, A, B. & H. D. PRANGE (1976). Dive and breath hold metabolism of the brown
water snake, Natrix taxispilota. Comp. Biochem. Physiol. 554A, 61-67.

JACOB, J. S. & H. S. McDONALD (1976). Diving bradycardia in four species of North
American aquatic snakes. Comp. Biochem, Physiol. 53A, 69-72.

JOHANSEN, K. (1959). Heart activity during experimental diving of snakes, Amer. J.
Physiol. 197, 604-606.

McDOWELL, S. B. (1969). Notes on the Australian sea-snake Ephalophis greyi M, Smith
(Serpentes: Elapidae, Hydrophiinae) and the origin and classification of sea-snakes.
Zool. J. Linn. Soc, 48, 333-349.

MURDAUGH, H. V. Jr. & J. E. JACKSON (1962). Heart rate and blood lactic acid
concentration during experimental diving of water snakes. Amer. J. Physiol. 202, 1163-
£165.

POUGH, F. H. (1973). Heart rate, breathing and voluntary diving of the elephant trunk
snake Acrochordus javanicus. Comp, Biochem. Physiol. 44A, 183-189.

SEYMOUR, R. S. (1974). How sea snakes may avoid the bends. Nature 250, 489-490.

SEYMOUR, R. S. (1976). Blood respiratory properties in a sea snake and a land snake.
Aust. J. Zool, 24, 313-320.

SEYMOUR, R. S. & M. E. D. WEBSTER (1975). Gas transport and blood acid-base balance
in diving sea snakes. J. Exp. Zool. 197, 169-181,

SMITH, M. A. (1926), Monograph of the sea-snakes (Hydrophiidae). Verlag J. Crammer,
Weinheim, 130 pp. (1964 reprint).

STANDAERT, T. & K. JOHANSEN (1974). Cutaneous gas exchange in snakes. J. Comp.
Physiol. 89, 313-320.

Aust. Zool. 19(2), 1977 167

ex, g.¥ oe a wet =) Ee,
ee

5 Hw

The Labrid Fish Genus Pseudojuloides, with Description
of a New Species

A. M. AYLING' and B. C. RUSSELL?

‘Department of Zoology, University of Auckland, Auckland, New Zealand.
“The Australian Museum, 6-8 College Street, Sydney, N.S.W. 2000*

ABSTRACT

Pseudojuloides elongatus n.sp. is described from specimens collected in New
Zealand, Norfolk Island, New South Wales and Western Australia, and Japan. The
genus Pseudojuloides Fowler and the type and only other species in the genus,
P. cerasinus (Snyder), are redescribed.

INTRODUCTION

The genus Pseudojuloides was created by Fowler (1949) to separate
Pseudojulis cerasina Snyder (1904) from other species of the genus Pseudojulis
Bleeker (1862). Fowler differentiated Pseudojuloides by its much larger thoracic
scales, shorter pectoral fins, greater number of scale rows on the caudal fin base,
and differences in colouration. These characters alone seem insufficient to warrant
separate generic status and probably for this reason there has been some confusion
as to the validity of the genus, Randall (1973) for example, has referred
Pseudojuloides cerasinus to the genus Leptojulis.

The discovery of a new species of Pseudojuloides in northern New Zealand
waters led us to closely re-examine the status of the genus. Comparing our
specimens and those of P. cerasinus with Fowler’s description revealed several
major discrepancies in the original description of Pseudojuloides. These necessitate
a review of the genus.

In this paper we redescribe Pseudojuloides and the type species P. cerasinus.
The new species of Pseudojuloides collected from northern New Zealand and
subsequently found also in eastern and western Australia, at Norfolk Island, and
in southern Japan, is described.

METHODS

Measurements were made with vernier calipers to the nearest half millimeter.
Standard length is abbreviated as SL. In the description, counts and proportions
follow Randall (1972). For the new species, those for the holotype are given

*Present address: School of Biological Sciences, Macquarie University, North Ryde,
NES We 2Ats:

Aust. Zool. 19(2), 1977 169

A. M. AYLING and B. C. RUSSELL

first while those for the paratypes, when different from the holotype, appear in
parentheses.

_ Type material has been deposited in the following institutions: Australian
Museum, Sydney (AMS); Bernice P. Bishop Museum, Honolulu (BPBM);
National Museum of New Zealand, Wellington (NMNZ); Tanaka Memorial
Biological Station, Tokyo (TMBS); Western Australian Museum, Perth (WAM).

DESCRIPTIONS

Pseudojuloides Fowler

Pseudojuloides Fowler, 1949: p. 119 (type species: Pseudojulis cerasina Snyder,
by original designation).

DESCRIPTION:

Dorsal’ ‘rays: 1X, 11-12; anal rays Vie" 492 pectoral rays: 1) (ae
pelvic rays 1,5; principal caudal rays 14. Lateral line continuous, abruptly bent
downward beneath soft portion of dorsal fin, 27 scales in lateral line, an additional
enlarged scale beyond caudal base. Lateral line pores simple, unbranched. Scale
rows above lateral line beneath origin of dorsal fin 22, below lateral line to
origin of anal fin 72. Gill rakers small, 13-16 on first arch. Branchiostegal rays
6. Vertebrae 10+15.

Body elongate, moderately compressed, snout pointed. Interorbital convex,
low; eye small. Mouth terminal, lips moderately broad and well developed; lower
lip with prominent downward projecting flap on each side, inner surface of upper
lip plicate. Single pair of well-developed, forwardly projecting canines in front of
jaws, upper pair splayed apart, those of lower jaw fitting between them; teeth
on sides of jaws small and laterally compressed, restricted to anterior part of
jaw. Larger individuals with a posterior canine in the angle of jaw on each side.
Lower pharyngeal plate broadly Y-shaped, posterior row of teeth laterally
compressed and elongate, those in centre somewhat enlarged and asymmetrically
conical, remainder molariform with 4-5 small conical teeth extending in single
row onto anterior shank of bone. Upper pharyngeals with teeth in triangular
patch, anterior row of which are asymmetrically conical, remainder molariform

(Figs. 1-6).

Preoperculum entire with free lower edge, about 1.5 times longer than
posterior free edge; gill membranes broadly attached to isthmus with very narrow
free fold posteriorly. Nostrils small, anterior nostril in short tube, posterior nostril
with small dermal flap on anterior margin.

Head naked except for patch of small scales on nape or on side of head
above operculum. Thoracic scales smaller than those on rest of body. No scales
on fins except for caudal which has about 4 rows of small scales on base.

Caudal fin rounded. Dorsal fin long, its origin slightly forward of vertical
through upper pectoral base. Anal fin elongate, its origin below last dorsal spine.

170 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PSEUDOJULOIDES

Pectoral fins rounded, first soft ray longest. Pelvic fins pointed, first two rays
longest.

Sexually dimorphic; males brightly coloured, females drab.

REMARKS:

Fowler (1949) appears to have erred in his original description of
Pseudojuloides. His description of colouration as “little contrasted, largely uniform”’
applies only to female specimens. A more important discrepancy is in his
description of the teeth as “‘uniserial, largest in front of jaws, gradually smaller to
last or posterior, all simple, pointed, conical.” In all the specimens we have
examined, the teeth in the sides of the jaw are laterally compressed. John R.
Paxton, who has examined the type of P. cerasinus for us in the U.S. National
Museum, reports that the teeth in that specimen also are small and laterally
compressed.

Figs. 1-6. Pharyngeal bones of two species of Pseudojuloides.
1-3 P. cerasinus 63mm SL. Fig. 1. Oblique lateral view of lower pharyngeal
bone. Fig. 2. Dorsal view of lower pharyngeal bone. Fig. 3. Ventral view of
upper pharyngeal bones.
4-6 P. elongatus 54mm SL, same sequence as above.

Aust. Zool. 19(2), 1977 4 171

A. M. AYLING and B. C. RUSSELL

The relationships of Pseudojuloides to the related genera Pseudojulis,
Leptojulis and Halichoeres are not clear. All are characterised by small thoracic
scales, naked head, absence of scales on dorsal and anal bases and 9 pungent dorsal
spines. The type of Pseudojulis (P. girardi Bleeker) appears to be a juvenile
Halichoeres and the status of this genus is doubtful. Separation of the other
genera has been mainly on the basis of differences in jaw dentition. Leptojulis
(type L. cyanopleura Bleeker) differs from both Pseudojuloides and Halichoeres
in possessing two pairs of large canine teeth at the front of the jaws, although in
small specimens of L. cyanopleura (less than 45 mm SL) the second pair are not
well developed. In Halichoeres (type H. bimaculatus Ruppell) the jaw teeth are
small and conical and clearly distinct from those of Pseudojuloides. The presence of
more laterally compressed jaw teeth in H. biocellatus Schultz, although less widely
separated than in Pseudojuloides, however, suggests that the structure of the jaw
teeth in Halichoeres may be somewhat variable. Similarly, the presence of a
posterior canine in the angle of the jaw, used as a diagnostic character by some
authors, appears to be variable among species of Halichoeres which we have
examined. The posterior canine in Pseudojuloides elongatus is developed only in
specimens greater than about 100 mm SL. For the present we regard Pseudojuloides
as a valid genus distinct from Halichoeres by its more elongate body and
compressed, widely separate jaw teeth.

Pseudojuloides cerasinus (Snyder )
Bigs) 1-3ic7

Pseudojulis cerasina Snyder, 1904: p.528 (type locality, Honolulu, Hawaii).
Pseudojuloides cerasinus — Fowler, 1949: p.119 (Hawaiian Is.).
Leptojulis cerasinus — Randall, 1973: p.197 (Society Is.).

MATERIAL EXAMINED:

AMES 71.1-74702004,. °( 10273 mar. SE. Uvea sAtolle: (20° 237S2* toa ei
Loyalty Islands, 10-25 m, speared by G. R. Allen and W. A. Starck 18 June,
1973. AMS J.18094-002, (1) 78.5 mm SL., One Tree Island (23°30’S, 152°05’E),
Great Barrier Reef, 20 m, speared by A. M. Ayling and B. C. Russell 16 September,
1974. AMS 1.18366-001, (2) 60-71 mm SL., off Lahelahe Point (21°27.5’N,
158°13’E), Oahu, Hawaii, 28 m, speared by J. E. Randall 13 July, 1968.

DIAGNOSIS:

A species of Pseudojuloides with the following combination of characters:
Dorsal rays IX,11; pectoral rays i, 11; lateral jaw teeth 6-7; triangular patch
of small predorsal scales on nape; male dark green with deep blue midlateral line,
below which is light green line, belly blue; lower part of head and cheeks blue
with distinctive blue band passing from back of eye across operculum down to
pectoral base, distinctive blue band on caudal fin separating dark outer half;
female uniform reddish-brown.

172 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PSEUDOJULOIDES

DESCRIPTION:
Dorsal rays (X, 11; anal rays LIT, 12; pectoral rays 1,°11> pelvic rays 1,5;
principal caudal rays 14; gill rakers on first arch small, 7+6.

Body elongate, depth in front of anal fin 4.1-4.2 in SL, width 1.9-2.1 in
depth; head pointed, length 2.9-3.2 in SL; snout including lips 3.1-3.2 in head;
eye diameter 1.4-1.9 in snout; interorbital space convex, bony width 4.9-5.4 in
head; least depth of caudal peduncle 2.7-3.1 in head; length of caudal peduncle
1.0-1.1 in least depth of peduncle.

Upper jaw nearly reaching a point vertically below anterior nostril; single
pair of pointed, well-developed, forward projecting canines at front of jaws;
6-7 small and laterally flattened teeth, well-spaced, in anterior part of jaws along
each side. Pharyngeal teeth (Fig. 1): upper bones with 20-22 teeth forming
triangular patch, anterior row in each patch asymmetrically conical, remainder
molariform; lower bone broadly Y-shaped, row of 10-11 laterally compressed
elongated teeth, centre three teeth somewhat enlarged and asymmetrically conical,
remainder molariform; 4-5 conical teeth extend uniserially on to anterior shank of

the bone.

Caudal fin length 1.5-1.8 in head length; first dorsal spine 1.7-2.1 in snout
length; second dorsal spine about one and a quarter times as long as first; ninth
dorsal spine about one and a half times as long as first; length of longest dorsal
ray 3.2-3.3 in head length; first anal spine 2.3-2.7 in first dorsal spine; third anal
spine about three times as long as first; longest anal ray about equal to longest
dorsal ray. Pectoral fins 1.8-2.1 in head length; origin of pelvic fins below lower
base of pectoral fins; pelvic fins 2.1-2.7 in head length, first pelvic ray longest,
reaching almost to vent.

Head naked except for a triangular patch of small predorsal scales on nape.

coLouR: (from colour transparencies )
Male — body colour dark green, a deep blue row of scales midlaterally,
below which is a row of light green or yellowish scales. Scattered indistinct

A

S &)
"

Fig. 7. Pseudojuloides cerasinus 6 75mm SL.

Aust. Zool. 19(2), 1977 173

A. M. AYLING and B. C. RUSSELL

blue spots on belly giving an overall blue diffusion. Head dark green above, a
diffuse broad blue band passing from upper lip to just behind eye. A narrower
dark blue band beginning behind eye, crossing preopercule, bending downwards
across operculum and ending just above pectoral base. Lower part of head and
cheeks blue. Iris green with a blue rim. Dorsal and anal fins dark green with
blue margin, a second narrower blue line medially along fin. Caudal dark green
at base, a narrow deep blue band separating the outer half which is black. Pectorals
hyaline; pelvics dark green with blue markings. Colours in preservative are
largely faded but in the male the dark blue midlateral line remains as a dusky
line, the light green band below it somewhat lighter than the rest of the body.
The dark outer margin and band on the caudal fin remain distinct.

Female — uniform reddish brown, fins translucent reddish brown.

DISTRIBUTION:

Hawaii, Loyalty Islands, Great Barrier Reef. John W. Shepard reports that
this species also occurs in southern Japan and the Ryuku Islands.

Pseudojuloides elongatus n. sp.
Figs. 4-6, 8-10; Table 1
Leptojulis sp. Masuda et al, 1975: p. 304 (Izu Oceanic Park, southern Japan)

HOLOTYPE:

NMNZ 6153, 123mm SL, Nursery Cove (35°28.5’S, 174°44E), Poor
Knights Islands, New Zealand, 12 m, speared by A. M. Ayling 4 December, 1974.

PARATYPES:

AMS: 1,17033-036, (1) 49.5mm Sb» Northo Head (33°49°S. (151° te Be
Sydney Harbour, 7m, collected with rotenone by G. R. Allen, D. F. Hoese,
G. McPherson, J. Paxton, D. Pollard, B. C. Russell, 6 April, 1974. AMS
1.17735-008, (1) 58 mm SL., Camp Cove (33°50’S, 151°16’E), Sydney Harbour,
5m, hand-netted by R. Kuiter 28 April, 1974. AMS 1.17743-004, (3) 56.5-60 mm
SL., Watsons Bay (33°50’S, 151°16’E), Sydney Harbour, 5m, hand-netted by
R. Kuiter 13-14 April, 1974. AMS I.17767-001, (6) 47.5-67 mm SL., between
Watsons Bay and Parsley Bay (33°49.5’S, 151°16’E), Sydney Harbour, 5m,
hand-netted by R. Kuiter 5 May, 1974. AMS I.17800-001, (3) 66.5-75 mm SL.,
Sugarloaf Point (31°26’S, 152°32’E), Seal Rocks, 8 m, hand-netted by R. Kuiter
13 May, 1974. AMS 1.18772-001, (1) 66mm SL., Phillip Island (29°07’S,
167°56.5'E) Norfolk Island, 15m, speared by B. C. Russell and A. Piper
20 September, 1975. BPBM 18022, (1) 67mm SL., Balmoral Beach (33°49’S,
151°15°E), Sydney Harbour, 5 m, hand-netted by R. Kuiter 3 May, 1974. NMNZ
6155, (1) 106 mm SL., Nursery Cove, Poor Knights Islands, New Zealand, 15 m,
speared by A. M. Ayling 18 April, 1974. NMNZ 6157, (1) 95 mm SL., Sandager’s
Reef (35°28.5°S, 174°44’E), Poor Knights Islands, New Zealand, 10 m, speared

174 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PSEUDOJULOIDES

by A. M. Ayling 20 February, 1974. NMNZ 6156, (1) 126.5 mm SL., same data
as holotype. NMNZ 6154, (1) 121mm SL., Sandager’s Reef, Poor Knights
Islands, New Zealand, 10 m, speared by A. M. Ayling 2 February, 1975. TMBS
750819-1, (1) 115mm SL., Igaya Bay (34°05’N, 124°10’E), Miyake-jima, Izu
Islands, Japan, 14m, screen netted by K. Meyer and J. T. Moyer. WAM
P25110-001, (3) 88-97 mm SL., Kendrew Island (20°28.5’S, 116°32’E), Dampier
Archipelago, Western Australia, 10 m, speared by G. R. Allen 2 November, 1974.
WAM P25318-007, (1) 106.5 mm SL., Batavia wreck site (28°30’S, 113°44’E),
Abrolhos Island, Western Australia, 8-10 m, speared by G. R. Allen 22 May,
1975. WAM P25318-008, (1) 112 mm SL., same data as previous specimen.

DIAGNOSIS:

A species of Pseudojuloides with the following combination of characters:
dorsal rays IX, 12; pectoral rays i, 10; lateral jaw teeth 3-4 (upper jaw), 4-5
(lower jaw); predorsal scales absent; male olive green, back dusky, head

8

zs IEE EXE a7
ANE A <a 4
ie AS p g AL
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. Pee Sine sey fafa (SUNS Va NANG [2 ‘
, peg tet aes s HA ths) . PIAS) me
=a 2 : ee = 7s aa PPK tig
—< D 22 a, in r EN ep N es 6 --
, = ad ay A 9 NG AS 4), 8 ERG [ND
_=s 7) 2 m OF Sr ad A ig ne PES ras oo
. Cwra4 a } > *y a RY Soi een bd 7 "4
x or 32 hams Le TIS Say Cals Ne RE CANS A =
CERES hac ay’ = ot * Sy: AN PAX Wwe Wis
en Oa pte 8 Sag hooey las fh very” i Cet SPARES AMD =
a fea o eee 7 4,7 —F a 7, ~ E PZ, entee
= ’ et ie ibs’ ~* “fs 4 a fe Ore so An
é = . 3 KOS He, Petar (EE Lif SE he AY Soa
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7

Figs. 8-10. Pseudojuloides elongatus. Fig. 8. Juvenile 28mm SL. Fig. 9. @ 72mm SL.
Fic. 10: 4 78mm SL.

Aust. Zool. 19(2), 1977. 175

A. M. AYLING and B. C. RUSSELL

TABLE 1

MORPHOMETRIC PROPORTIONS OF HOLOTYPE AND 5 PARATYPES OF
PSEUDOJULOIDES ELONGATUS

Proportions are as a percentage of the standard length

Registration number NMNZ NMNZ NMNZ AMS WAM TMBS
6153 6156 6154 1.17800-001 P25318-007 750819-1

Sex re) 3 2 ey 2 )
Standard length (mm) 123.0 126.5 121.0 Fo 106.5 115.0
Depth of body 22.8 fadindl 23 19.3 19.7 ZT
Width of body 12.6 12.3 14.9 10.7 8.5 13.0
Head length 29'3 30.4 29.3 30.0 30.0 294
Snout length 11.4 123 11.6 liaise’ 10:3 ri
Eye diameter 4.3 4.2 a 5.0 5.6 52
Bony interorbital width 6.1 59) 6.0 6.0 52 6.5
Length of upper jaw Wa Ws 6.6 Deh 6.1 Sz
Least depth of

caudal peduncle 11.4 10.7 10.7 LO. 10.3 it3
Length of caudal

peduncle 9.4 8.7 9.5 Sav. 122 ii7
Snout to origin of

dorsal fin 28.1 29.3 29.8 28.7 28.6 27.8
Snout to origin of

anal fin 52.0 50.6 52:5 48.7 49.8 49.6
Snout to origin of

ventral fin 3215 34.4 31.4 32.0 30.0 30.9
Length of caudal fin 19.9 20.6 19.0 20.0 19.7 20.9
Length of pectoral fin 14.2 14.6 14.9 14.7 | ESS: 13.0
Length of ventral fin A 11:5 112 12.7 12.2 12:1
Length of first

dorsal spine aed. 4.0 3.9 3.3 3.8 43
Length of second

dorsal spine 4.7 5.9 5:6 a1 5.2 7
Length of last

dorsal spine 8.1 6.9 9.1 8.7 8.0 8.7
Length of longest

dorsal ray 10.6 10.7 10.7 ia Wes 10.3 D1 .7
Length of first anal spine 3.1 2.8 25 3.3 23 2.6
Length of second

anal spine 4.3 4.6 552 5.0 4.7 4.3
Length of third

anal spine 5.9 6.5 TA 6.7 5.6 6.5
Length of longest

anal ray 10.2 oy oo 10.7 a 10.8
Length of dorsal fin base 58.1 58.1 58.7 56.0 59.2 60.4
Length of anal fin base 41.3 41.9 39°F 39.3 39.4 43.9

176 Aust. Zool. 19(2), 1977

A NEW SPECIES OF PSEUDOJULOIDES

brownish with four distinctive blue lines on each side, a dusky patch behind the
pectoral fin base; female a uniform olive green.

DESCRIPTION:

Dorsal rays IX, 12; anal rays III, 12; pectoral rays i, 10; pelvic rays I, 5;
principal caudal rays 14; gill rakers on first gill arch small, 9 + 7.

Body very elongate, depth in front of anal fin 4.4 (3.2-5.6) in SL, width
1.8 (1.5-2.5) in depth; head pointed, length 3.4 (3-3.4) in SL; snout including
lips 2.5 (2.5-3.1) in head; eye diameter 2.7 (1.5-2.9) in snout; interorbital space
convex, bony width 4.8 (4.5-5.8) in head; least depth of caudal peduncle 2.6
(2.7-3.3) in head; length of caudal peduncle 0.8 (0.7-1) in least depth of peduncle.

Upper jaw nearly reaching a point vertically below anterior nostril; single
pair of pointed, well-developed, forward projecting canines at front of jaws;
upper jaw with 3-4 small laterally flattened teeth, well-spaced in anterior part
of jaw; lower jaw with 4-5 teeth similarly shaped and arranged. Pharyngeal teeth
(Fig. 1): upper bones with 19-20 teeth forming triangular patch, anterior row
in each patch asymmetrically conical, remainder molariform; lower bone broadly
Y-shaped, posterior row of 9 laterally compressed elongate teeth, centre three
somewhat enlarged and asymmetrically conical, remainder molariform; 4-5 small
conical teeth extend uniserially onto anterior shank of bone.

Caudal fin length 1.5 (1.4-1.7) in head length; first dorsal spine 3.1 (2.2-4.4)
in snout length; second dorsal spine about one and a half times as long as first;
ninth dorsal spine about twice as long as first; length of longest dorsal ray 2.8
(2.6-3.4) in head length; first anal spine 1.2 (0.7-2.2) in first dorsal spine; third
anal spine about twice as long as first; longest anal ray about equal to the longest
dorsal ray. Pectoral fins 2.1 (1.3-2.6) in head length; pelvic fins 2.6 (2.3-3.2)
in head length, first pelvic ray longest, reaching about two thirds of way from
fin base to vent. See Table 1.

Head naked except for 4-5 diagonal rows of small scales on each side of
head in forward projecting V-shaped patch above operculum; no midline predorsal
scales.

COLOUR:

Male (from holotype) — Body colour olive green with lines of irregularly
shaped turquoise blue spots and scattered irregular orange-brown patches on the
- dorsal scales; head brown above, orange-brown on the sides and green beneath
chin with four blue lines on each side; the upper line originating above the eye
and passing back to the upper origin of the opercular flap; the second beginning
dorsally on the snout and passing back through the eye before slanting upwards
to run around the upper edge of the opercular flap; the third line originating on
the upper lip and immediately breaking into two, one part passing down around
the lips, the other back through the eye and slanting abruptly downwards towards
the isthmus; the fourth arching from the lower lip up past the lower rim of the

Aust. Zool. 19(2), 1977 . 177

A. M. AYLING and B. C. RUSSELL

eye and down to the preopercular margin; on the operculum between the second
and third lines is another short diagonal blue line that combined with the
posterior portion of the second line separates a dusky patch on the opercular flap.
Iris orange with a blue rim. A dusky patch behind pectoral fin base and an orange,
blue bordered patch on the body behind this. Dorsal fin pale olive with red-brown
diagonal lines and a narrow blue margin; anal fin red-brown with a prominent
blue margin and a series of irregular diagonal blue lines. Caudal pale olive with
a faint red rim, a blue line on upper and lower margins and blue spots at the
base of the fin rays. Pectoral fins hyaline; pelvics blue with red-brown streaks.
Colours in preservative are faded, body colour pale green, the blue lines on the
male becoming dusky, but the dark back and dusky patch behind pectoral fin
remain prominent.

Female — Uniform olive green with pale brown fins.

REMARKS:

This species may be distinguished from Pseudojuloides cerasinus by its extra
dorsal soft ray; one fewer pectoral ray; slightly more elongate body; fewer
jaw teeth; absence of predorsal scales, head scales being limited to a small patch
on either side of the head above operculum; and differences in colour.

Male specimens from Western Australia and from Japan that we have
examined are much more darkly pigmented on the back, some almost black
compared with those from eastern Australia and New Zealand.

Named elongatus in reference to the elongate body form.

DISTRIBUTION:

North-eastern New Zealand, Norfolk Island, New South Wales, Western
Australia and southern Japan.

ACKNOWLEDGEMENTS

We wish to thank Rudi H. Kuiter for his drawings of Pseudojuloides and Dr. John R.
Paxton for his constructive criticism of the manuscript. Dr. Gerald R. Allen, Rudi H. Kuiter
and Dr. John W. Shepard provided additional information and specimens of P. elongatus.
Collecting trips in New Zealand were supported by the New Zealand University Grants
Committee.

REFERENCES

BLEEKER, P. (1862). Comspectus generum Labroideorum analyticus. Proc. Zool. Soc.
Lond., XXVII (1861), 408-418.
FOWLER, H. W. (1949). The fishes of Oceania — Supplement 3. Mem. Bernice P. Bishop

Museum XII (2), 3-186.
MASUDA, H., ARAGA, C. & T. YOSHINO (1975). Coastal fishes of southern Japan.
Tokyo: Tokai University Press. 379 pp.
RANDALL, J. E. (1972). A revision of the labrid genus Anampses. Micronesica 8, 151-190.
RANDALL, J. E. (1973). Tahitian fish names and-.a preliminary checklist of the fishes
of the Society Islands. Occ. Pap. Bernice P. Bishop Museum XXIV, 167-214.
SNYDER, J. O. (1904). A catalogue of the shore fishes collected by the steamer Albatross
about the Hawaiian Islands in 1902. Bull. U.S. Fish Commission XXII, 513-538.

178 Aust. Zool. 19(2), 1977

A Juvenile Gempylid Fish, Nealotus tripes, from
Eastern Australia

IZUMI NAKAMURA! and JOHN R. PAXTON
Department of Ichthyology, The Australian Museum, Sydney, New South Wales, 2000

ABSTRACT

The gempylid fish Nea/otus tripes Johnson is newly recorded from eastern
Australia, based on one juvenile specimen 49.8mm standard length taken off
Sydney. The juvenile has relatively larger, serrated fin spines than 24 larger
specimens from Lord Howe Island. The species is also known from Papua-
New Guinea, on the basis of a single specimen previously identified as
Promethichthys prometheus.

INTRODUCTION

The fishes of the family Gempylidae are widely distributed in tropical and
temperate waters of both the Atlantic and Indo-Pacific Oceans. The family consists
of 14 genera and 15 species (Parin and Bekker, 1972) considered to be predaceous
and mostly inhabiting deep waters. However, the systematics of the family is
not understood comprehensively. There is also a paucity of life history information
on these fishes.

Around Australia very few species of gempylid fishes have been recorded.
Munro (1958a) reported only four species in his ‘Handbook of Australian
fishes”, Leionura atun (=Thyrsites atun), Rexea solandri, Lepidocybium
flavobrunneum and Ruvettus tydemani (=Ruvettus pretiosus). Of these, both
Thyrsites atun and Rexea solandri are important commercially. The other two
are rather rare. Five more species, which are all uncommon, are recognised
around Australia (Nakamura, unpublished data; Parin and Bekker, 1972). They
are Epinnula orientalis, Gempylus serpens, Rexea prometheoides, Nealotus tripes
and Promethichthys prometheus.

Nealotus tripes Johnson, a cosmopolitan species, has been recorded from
Lord Howe Island (Ogilby, 1899; Allen e¢ al., 1976) and off north western
Australia (Parin and Bekker, 1972). Recently, a juvenile specimen of the species
was obtained off Sydney by F. R. V. Kapala, and is a new record of occurrence
from the waters of the east Australian continental shelf. This paper compares the
juvenile specimen with the specimens collected from Lord Howe Island and
deposited in The Australian Museum (AMS, 23 specimens) and the Queensland
Museum (QM, 1 specimen, Ogilby’s type of Machaerope latispinus).

Present address: Fisheries Research Station, Kyoto University, Maizuru, Kyoto 625, Japan.

Aust. Zool. 19(2), 1977 179

IZUMI NAKAMURA and JOHN R. PAXTON

MATERIALS AND METHODS

A juvenile specimen of Nealotus tripes (AMS I. 17887-006, 49.8 mm standard
length) on which this description is based, was obtained from off Sydney, N.S.W.
(34°13’S, 150°22’E) in a midwater trawl fishing from 0-145 m over a bottom
of 165m on May 6, 1974 by F. R. V. Kapala of New South Wales Fisheries.
Twenty-four speciments collected from the beach at Lord Howe Island were
used for the comparison (AMS JIA. 1398-1404, I. 4121-4123, 1.4312, 1.5189,
1.5590-5595, 1.5597, 1.5598, 1.7858, 1.7859, 1.10700; 130.5-236.1 mm in standard
length and QM 11/74 137.5mm in standard length). The methods of
measurement follow Hubbs and Lagler (1949). The drawings were made with
the aid of a binocular microscope and vertebral counts from X-rays.

DESCRIPTION OF STE EON RINNE TS

Counts, measurements and description of the juvenile specimen (1) are
followed by those of the Lord Howe Island specimens (24) in parentheses.
Dorsal. fin rays XX, 1, 182. (X1X-XX] 4,.°16192-2)- “anal rays 41 = eee
(II, 15-19+2); pectoral rays 14 (12-14); pelvic rays I,1(1,1); gill-rakers on
first arch 54+1+11 (5-8+1+10-17); branchiostegals 7 (7); intramuscular bones
— (31-34); vertebrae including urostyle 37 (20+17=37 or 21+16=37); pyloric

caeca — (8).

The following measurements are expressed as percent standard length: head
length 28.1 (25.6-28.3); snout to insertion of first dorsal 24.7 (24.0-26.4); snout
to insertion of second dorsal 75.7 (73.1-75.9); snout to insertion of pectoral 26.7
(26.3-28.5); snout to insertion of pelvic 26.3 (27.3-31.4); snout to anus 72.5
(67.2-71.7); snout to insertion of anal 73.4 (66.9-76.2). The following
measurements are expressed as percent of head length: body depth 41.4
(43.0-56,2);. body... width. 17.9 (12.0-31,2);. snout. lensth 35.0: (3624295.
upper jaw length 42.0 (43.6-48.2); orbit length 25.0 (19.4-23.4); interorbital
width 15.0 (17.1-20.6); pectoral fin length 43.6 (37.9-60.5); pelvic spine length
37.1 (7.8-15.9); lensth of longest first dorsal ray (spine) 30.7 (27.7-33:3);
length of longest second dorsal ray (soft ray) 31.5 (25.8-34.0); length of first

Fig. 1. Juvenile of Nealotus tripes (AMS I. 17887-006) obtained from off Sydney. Ventral
broken. Line shows the position of anus. Scale equals 5 mm.

180 Aust. Zool. 19(2), 1977

GEMPYLID FISH FROM AUSTRALIA

Fig. 2. Inside of upper (left) and lower (right) jaws of the juvenile. YO: vomer, PAL:
palatine, GH: glossohyal. Scale equals 1 mm.

Fig. 3. Outer surface of first gill arch (left side) of the juvenile. Scale equals 1 mm.

Fig. 4. Spines in anterior view of the juvenile. From left to right: first dorsal spine, 20th
dorsal spine, pelvic spine of left side, first anal spine and second anal spine. Each
scale indicates 1 mm respectively.

anal spine 15.0 (5.4-14.2); length of longest anal ray (soft ray) 29.2 (24.5-35.4);
caudal peduncle depth 14.3 (14.3-20.6).

Body elongate and compressed (Fig. 1). Greatest body width less than
greatest body depth. Upper profile of head slightly convex from tip of snout to
insertion of first dorsal fin (nearly straight in Lord How Island specimens). Snout
less than twice as long as eye (slightly more than twice in larger specimens).
Jaws bluntly conical; lower jaw projecting forward beyond tip of snout. Mouth
large, maxillary scaleless, extending backward slightly beyond front edge of pupil.
Three fangs on each side of upper jaw near tip of snout. A pair of canines near
symphysis of lower jaw. A pair of minute teeth on vomer (vomer edentulous in
larger specimens). A single series of small teeth on palatines (Fig. 2). Eye large,
almost round. Infraorbital space very narrow, its least width less than diameter
of pupil. Interorbital space narrower than eye. Lower part of preopercle armed
with 2 spines. Several spines on hind part of opercle and subopercle (spines
inconspicuous on hind part of opercle and subopercle in larger specimens). Lateral
line single from near upper margin of opercle almost straight backward and

Aust. Zool. 19(2), 1977 181

IZUMI NAKAMURA and JOHN R. PAXTON

gradually downward from middle of first dorsal fin to base of caudal fin. Scales
small, cycloid, imbricate in regular rows, but rather deciduous. Dorsal fin inserted
above upper margin of opercle, base of first dorsal fin about three times as long
as base of second dorsal fin. Soft rays of second dorsal fin preceded by a small
spine, almost the same as soft rays of anal fin in shape and size. A flattened,
dagger-shaped, serrated. spine followed by a small, embedded, serrated spine
between anus and insertion of soft anal (a dagger-shaped, smooth spine followed
by a small, embedded, smooth spine in larger specimens). Soft rays of second anal
fin inserted under third soft ray of second dorsal fin. Finlets 2 in dorsal and
anal fins. Pectoral fin rather long, extending to vertical from base of seventh
dorsal spine. Pelvic fin consists of a long flattened spine with serration followed
by a soft ray (pelvic fin reduced to a small, single, smooth spine followed by a
minute soft ray in larger specimens), inserted below origin of pectoral fin. All fin
spines (20 first dorsal, 1 second dorsal, 1 pelvic and 2 anal) flattened and
serrated more or less as shown in Fig. 3 (all fin spines smooth, elongated, dagger-
shaped in larger specimens). Gill-raker at angle of first arch long, Y-shaped,
other rakers short and acute (Fig, 4).

Colour of juvenile specimen fixed by formalin and preserved in ethanol,
head and dorsal side of body dark brown; ventral side of body pale brown;
pectoral fin dark brown; other fins pale brown. All fin membranes pale.

DISCUSSION

There have been few records of Nealotus tripes from waters around
Australia. Some twenty specimens were recorded from Lord Howe Island (Ogilby,
1899; Allen et al., 1976). Several specimens were recorded from off north western
Australia (Parin and Bekker, 1972). There is no record from around New Zealand
(Whitley, 1968), although four specimens were known from Kermadec Island
(Waite, 1910). There is hitherto no record from around Papua-New Guinea
(Munro, 1958b, 1967; Kailola, 1971, 1973, 1974). However, Promethichthys
prometheus (Cuvier), recorded from Papua-New Guinea on the basis of two
specimens by Munro (1958b) as species number 1128, represents two species.
The Laughlan Island specimen (C.S.I.R.O. B.620) is Nealotus tripes, while the
Collingwood Bay specimen (C.S.I.R.O. C.1616) is Promethichthys prometheus.
The juvenile described here is a new record of Nealotus tripes to the waters of
the east Australian continental shelf. This species is rather rare and distributed
in chiefly circum-tropical and sub-tropical zones throughout the world.

The pelvic fin consists of a spine and a soft ray in both the juvenile fish
and specimens from Lord Howe Island, although Matsubara and Iwai (1952)
described the ventral reduced to a single smooth spine. The senior author
re-examined the materials of Matsubara and Iwai deposited in Kyoto University
and found that they missed a minute soft ray because of the bad condition of
the materials. Strasburg (1964) gave a spine and 0-2 (mostly 2) soft rays as
the pelvic fin elements in postlarval stages. The relative length of the pelvic spine
(expressed as percent of standard length) decreases with growth (Fig. 5).

182 Aust. Zool. 19(2), 1977

GEMPYLID FISH FROM AUSTRALIA

LE)

100

LENGTH x

=
o

PELVIC SPINE LENGTH 7 STANDARD
wr

50 100 150 200 250
STANDARD LENGTH (mm)

Fig. 5. Relative length of pelvic spine in Nealotus tripes. Pelvic spine lengths are plotted
against the standard length. Squares: after Strasburg (1964); triangle: AMS IA.1424,
from Lord Howe Island (smallest specimen of the Australian Museum collection);
open circle: juvenile from off Sydney; closed circles: Lord Howe Island specimens.

The second anal spine is embedded along the ventral margin of the body,
following the erected longer first anal. spine in both juvenile and fishes from
Lord Howe Island, although both the first and second anal spines are erected in
Fig. 11 of Matsubara and Iwai (1952). The second embedded anal spine is
thought to be one of the specific characteristics (adult phase) of Nealotus tripes.
According to the figures shown by Strasburg (1964), both the first and second
anal spines are erect in the postlarval stage.

Comparing our juvenile with a juvenile of Nealotus tripes (41.5 mm in
standard length) described by Strasburg (1964) from southwest of Hawaii, both

Aust. Zool. 19(2), 1977 183

IZUMI NAKAMURA and JOHN R. PAXTON

are similar in external appearance. But they show some differences, such as head
contour (‘almost straight” of the former to ‘“‘round” of the latter), spines on
opercular and subopercular region (several spines on opercle and subopercle of
the former to two opercular spines only of the latter) and the condition of the
second anal spine (embedded of the former to erect of the latter).

ACKNOWLEDGEMENTS

We acknowledge with thanks the help of the captain and crew of F.R.V. Kapala
of New South Wales State Fisheries for obtaining the juvenile specimen. We are very
grateful to Roland J. McKay of the Queensland Museum and Ian S. R. Munro of C.S.I.R.O.,
Division of Fisheries and Oceanography, Cronulla for their permission to examine specimens
deposited there. We wish also to thank Reiko K. Nakamura for her help.

REFERENCES

ALLEN, G.,° D> HOESE,. J. PAXTON, J: RANDALL, Bo. RUSSEEL. W. STARCHES
F. TALBOT, & G. WHITLEY (1976). Annotated checklist of the fishes of Lord
Howe Island. Rec. Aust. Mus. 30, 365-454.

HUBBS, C. L. & K. F. LAGLER (1949). Fishes of the Great Lakes Region. Cranbrook
ins Sci, Bull) 26. 1-186;

KAILOLA, P. (1971). New records of fish from Papua. Papua New Guinea Agric. J. 22,
116-133.

. (1973). Additions to the fish fauna of Papua New Guinea, I. Papua New

Guinea Agric. J. 24, 1-15.
. (1974). Additions to the fish fauna of Papua New Guinea. III. Depart.

Agric., Stock Fish., Port Moresby, Res. Bull, 12, 54-89.

MATSUBARA, K. & T. IWAI (1952). Studies on some Japanese fishes of the family
Gempylidae. Pac. Sci. 6, 193-212.

MUNRO, I. S. R. (1958a). Handbook of Australian fishes. (Aust.) Fish. Newsletter 17,
113-116.

. (1958b). The fishes of the New Guinea region. A check-list of the fishes

of New Guinea incorporating records of species collected by the Fisheries Survey

Vessel “Fairwind” during the years 1948 to 1950. Papua New Guinea Agric. J. 10,

97-369.

. (1967). The fishes of New Guinea, Department of Agriculture, Stock and
Fisheries, Port Moresby, xxxvii + 651 pp. 78 pls., 6 color pls.

OGILBY, J. D. (1899). Additions to the fauna of Lord Howe Island. Proc. Linn. Soc.
New Scuth Wales 23, 730-745.

PARIN, N. V. & V. E. BEKKER (1972). Classification and distribution data of some
trichiurid fishes (Pisces Trichiuroidea; Scombrolabracidae, Gempylidae, Trichiuridae).
Trudy Inst. Okean. SSSR Akad. Nauk 93, 110-204. (In Russian).

STRASBURG, D. W. (1964), Postlarval scombroid fishes of the genera Acanthocybium,
Nealotus, and Diplospinus from the Central Pacific Ocean. Pac. Sci. 18, 174-185.
WAITE, E. R. (1910). A list of the known fishes of Kermadec and Norfolk Islands.

Trans. New Zealand Inst. (1909) 42, 370-383.

WHITLEY, G. P. (1968). A check-list of the fishes recorded from the New Zealand

region. Aust. Zool. 15, 1-102.

184 Aust. Zool. 19(2), 1977

Sex Change in the Wrasse Pseudolabrus gymnogenis
(Labridae)

G. R. McPHERSON'
School of Biological Sciences, Macquarie University, North Ryde, N.S.W. 2113

ABSTRACT

In Pseudolabrus gymnogenis there is a sexually undifferentiated primary germ
cell, here referred to as a protogonium that divides mitotically to produce a
slightly smaller yet similar deuterogonium cell. At this stage the germ cells
differentiate, prospective oocytes enter meiotic prophase and _ prospective
spermatogenic cells begin a phase of chromatin condensation prior to meiosis.
Meiotic stages and maturation stages of oocytes and sperm cells are described.
The only evidence of intersexuality in females was seen from two specimens
where small cysts of primary spermatogonia were located in the lamellar
epithelium although this was not regarded as evidence for an impending sex
inversion. A few meiotic stage oocytes were found in males and many males
displayed remnants of degenerate mature oocytes.

INTRODUCTION

The origin of primordial germ cells from the extra embryonic endoderm or
mesoderm and the early development of the teleost gonad has been investigated
by many authors (D’Ancona, 1945; Okkleberg, 1921). It has been well established
that primordial germ cells undergo a period of rapid mitotic increase when they
reach the gonadal primordium. However, the terminology of the early derivatives
of primordial germ cells is confusing.

In the teleost Oryzias latipes, Satoh and Egami (1972) reported mitotic
divisions of the primordial germ cells in sixteen day old fry. D’Ancona (1945)
believed the primordial germ cells (which he called protogonia) multiplied for a
number of generations, giving rise to smaller daughter cells or deuterogonia. As
the deuterogonia (as defined by D’Ancona, 1945) were believed to divide
mitotically to produce oogonia or spermatogonia, then this term is equivalent to
primary germ cell used by other workers, such as Barr (1963), Henderson (1962)
and Ahsan (1966).

In the lamprey, Lampetra planeri, Hardisty (1965) recognised three forms
of undifferentiated gonial cells, a primordial germ cell, a protogerium and a
deutergonium with total diameters of 22.5, 12.9 and 10.2 um respectively.
Although the dimensions of the three gonia differed, Hardisty (1965) indicated

1Present Address: Northern Fisheries Station, Queensland Fisheries Service, C/o Post Office.
Gairas, Qld) 4670

Aust. Zool. 19(2), 1977 185

G. R. McPHERSON

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19(2), 1977

Aust. Zool.

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SEX CHANGE IN THE WRASSE

similar histological characteristics such as a prominent nucleolus, a nuclear
membrane and a relatively chromatin-free reticulum. He showed that the primordial
germ cells divide to form the small protogonia which in turn keep dividing to
produce either more protogonia or the smaller deuterogonia. The deuterogonia
then enter meiotic phophase in definitive females, although they still retain the
capacity to divide mitotically.

Pseudolabrus gymnogenis is the most common species of labrid found along
the rocky foreshores of the N.S.W. Central Coast. However little is known of its
reproductive biology. This paper describes only the development of germ cells
in P. gymnogenis.

MATERIALS AND METHODS

Thirty-two collecting stations were made between 29th December, 1972, and
3rd October, 1973 from Long Reef, Sydney Harbour, Cronulla, Palm Beach,
Jervis Bay, Ulladulla and Batemans Bay, N.S.W. (Table 1).

Most specimens were collected by spearfishing. A fine mesh drop trap and
a wire trap 4 feet long, 2 feet in diameter and enclosed in one inch wire mesh
were also used. Badly damaged specimens were rejected.

Since fixation was not always possible immediately after death, post mortem
changes were inevitable in some instances.

All fish were preserved in 10% formal saline. The right lobe of two
mature ovaries, and the entire gonad of 10 juveniles were fixed in sea water
Bouin’s solution. The middle portion of one lobe from each gonad was embedded
in paraffin wax. Testes and immature females were sectioned at 5 um, mature
oocytes filled with large yolky masses crumbled if sectioned at less than 10 um.

Seven slides were prepared from each gonad, five were stained in Delafield’s
haematoxylin and alcoholic eosin and were used for the standard identification
of oocyte and sperm maturation stages. The remaining two slides were later
selected for more specific staining techniques. Feulgen proved to be successful
for the identification of mitotic and meiotic division stages. Azan was useful for
the differentiation of oocyte stages and spermatogenic stages. Periodic Acid Schiff
(PAS) was used to test for mucopolysaccharides in oocytes. Nile Blue A localised
lipofuscin pigments in testis tissue.

Sections of formalin fixed tissue from 10 males and 5 females were embedded
in gelatin, and sectioned at 8 um (usually 10 um for mature ovaries) on a freezing
microtome. Two slides were prepared, one was stained with Sudan Black B, the
other was subjected to the Schultz test for cholesterol (Lillie, 1965).

RE SUES
UNDIFFERENTIATED GERM CELLS IN P. gymnogenis

In Pseudolabrus gymnogenis protogonia and deuterogonia (as defined by
Hardisty) have been found in females and males. The interphase protogonium of

Aust. Zool. 19(2), 1977 187

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19(2), 1977.

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PROTOGONIUM

Mitotic Division

INTERPHASE
DEUTEROGONTIUM

PRIMARY
SPERMATOGONIUM LEPTOTENE

Mitotic Division

SECONDARY ES
SPERMATOGONIUM GS:

‘ ¥ ZYGOTENE
PRIMARY
SPERMATOCYTE

1st Meiotic Division

PACHYTENE
® SECONDARY

bh SPERMATOCYTE
2nd Meiotic Division

& SPERMATID

LATE
PACHYTENE -
EARLY
DIPLOTENE

r SPERMATOZOOQN

Fig. 1. Diagrammatic representation of the germ cell stages of P. gymnogenis.

Mature
oocyte stages are not represented.

190 Aust. Zool. 19(2), 1977

SEX CHANGE IN THE WRASSE

P. gymnogenis is a large cell with an average nuclear diameter of 5.5-5.6 um,
total diameter of 7.4-7.5um, and large nucleolus of 1.4 um and a well defined
nuclear membrane (Table 2 and 3). The reticulum is free of chromatin except
for a number of chromatin bridges extending from the nucleolus to the nuclear
membrane. Small clumps of chromatin may be seen around the nuclear periphery.
Prior to their mitotic division the protogonia loose the nucleolus and _ the
chromatin bridges disappear as the reticulum develops a high number of chromatin
granules. During mitotic phophase the nucleus elongates slightly and condensed
chromosomes are occasionally seen within the cell membrane, (Figure 2).

Deuterogonia result from the mitotic division of protogonia and may be
distinguished from the latter by the higher cytoplasm/nucleus ratio (of 1.6).
The average nuclear diameter ranges from 3.9-4.1 um and the total diameter is
6.59-7.19 um. Interphase deuterogonia with a distinct nucleolus and clear nuclear
reticulum are not often seen.

At this stage the germ cells differentiate, prospective oocytes enter meiotic
prophase, and prospective spermatogenic cells begin a phase of chromatin
condensation prior to meiosis (Figure 1).

OocyTE DEVELOPMENT

Meiotic Stages

Although meiotic prophase oocytes were not recorded for counts of ovarian
maturity, details of their morphology are necessary for the identification of oocyte
remnants in testes. The following description of P. gymnogenis was made from
sea water Bouin fixed material.

Although protogonia are common in juvenile females (Figure 3) interphase
deuterogonia are not often seen as they rapidly enter meiotic prophase. In what
may be described as the leptotene stage the nucleolus rapidly decreases in size,
there is an increase of coarse peripheral chromatin and fine chromatin threads
appear across the reticulum. The outline of the nuclear membrane tends to become
less distinct.

In zygotene, the chromosome threads progressively lengthen, thicken and stain
intensely. The progression from leptotene to early zygotene is seen as a sudden
concentration of chromatin to form a large mass of nuclear material in the centre
of the nucleus (Figure 4). The nuclear membrane is extremely faint. As the
nuclear diameter increases, the chromosomes spread out across the reticulum and
may be resolved as long thin strands. In the European Lamprey, in many zygotene
oocytes, a number of chomosomes at one pole of the nucleus condense into a
small cresentic mass, called a synapsis figure by Hardisty (1965). In the
other stages of zygotene, the chromosomes condense and orientate towards the
faintly defined nuclear periphery. Later stages develop a variable ovoid
configuration which gains a more extreme expression in pachytene.

While the leptotene nucleus shows little increase in size over that of the
interphase deuterogonia (3.9um to 4.2um), this is a 19% increase at the

Aust. Zool. 19(2), 1977 191

G. R. McPHERSON

Aust. Zool. 19(2), 1977

192

SEX CHANGE IN THE WRASSE

zygotene stage ().0um). The cytoplasm of the zygotene oocyte does not stain
well with haematoxylin and the presence of many nuclei in very close proximity
would suggest that the nucleus has expanded at a faster rate than the total cell
diameter.

The most sharply defined nuclear configuration is described as pachytene
(nuclear diameter 5.9um, total diameter 8.0um) which is marked by the
condensation of chromatin around the nuclear periphery.

The chromosomal threads contract and are predominantly restricted to the
periphery of the nucleus. At the points where the chromosomal threads intersect
or are in contact with the inner surface of the nuclear membrane, there are
often distinct chromatin granules (Figure 5). The nucleus at pachytene shows
20% increase over the zygotene stage, while the cell diameter has only increased
by 17% over the leptotene stage.

Hardisty (1965) regarded the reappearance of the nucleolus as marking the
transition to the pachytene stage. In P. gymnogenis the nucleolus is not evident
until late pachytene when it appears as a small round structure near the nuclear
periphery. The transition from late pachytene to early diplotene is difficult to
distinguish, although the most obvious change is the acceleration of cytoplasmic
growth (36% over pachytene, 18.0-10.9 um) and its rapid increase in basophilia.
The nucleus is 6.4 um in diameter. :

The final stage in oocyte proliferation is the development of diplotene oocytes.
This stage is marked by a further rapid increase in nuclear diameter to 15-30 um.
The cytoplasm also increases at a comparable rate, although the cell shape is variable.
The nuclear reticulum is more finely granular than in late pachytene — early
diplotene stage oocytes, and a very large basophilic nucleolus is present (Figure 6).

Nuclei up to the pachytene stage are always present in cysts scattered along
the lamellar epithelium. Late pachytene — early diplotene oocytes are invariably
located around the interior periphery of these epithelial cysts while the late
diplotene oocytes are always free of the lamellar epithelium. They are scattered
by a surrounding layer of connective tissue, which contributes to the formation
of the granulosa layer in mature oocytes.

Fig. 2. A mitotic prophase protogonium (P) in a testis.
Fig. 3. Protogonium (P) in an ovary. A late stage deutergonium (De) and a Leptotene
nucleus (L) are also shown.

Fig. 4. Early zygotene (EZ) and typical zygotene (Z) nuclei. In zygotene, synapis figures
are often shown.
Fig. 5. Pachytene nuclei (PT) showing a marked peripheral condensation of chromatin.
Fig. 6. Late pachytene/early diplotene (PT/ED) occyte. The nucleolus (n) is reappearing.
Fig. 7. Early yolk vesicle cocyte (YU). Perinucleolar oocyte (PN).
(Bigs) 2-67 x9 500; Fie. 7. xa 370)

Aust. Zool. 19(2), 1977 193

G. R. McPHERSON

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SEX CHANGE IN THE WRASSE

Maturation Stages
(a) Perinucleolar Stage (Figure 7)

This stage is rarely seen in juveniles and represents the first stage of oocyte
maturation, the nuclear diameter ranges from 30-40 um. The cell shape is far
more regular than that of diplotene oocytes and the nucleoli become closely
associated with the nuclear periphery and change from a spherical to a
hemispherical or elliptical configuration.

(b) Yolk Vesicle Stage (Figures 7 and 8)

Oocytes at this stage usually have a nuclear diameter range of 40-50 um,
a total diameter range of 120-160 um and are characterised by the appearance
of PAS positive yolk vesicles in the outer half of the cytoplasm. The vesicles first
appear as small, scattered vacuoles (often containing basophilic granules) which
become organised into a relatively distinct ring around the oocyte periphery, the
largest vesicle reaching about 7 um in diameter. Almost immediately after the
appearance of the yolk vesicles in the outer cytoplasm, small fatty droplets
develop in the inner half of the cytoplasm. The fatty droplets increase in size
and coalesce with others to form a ring of droplets around the nucleus, with
individual droplets up to 10 um.

(c) Primary Yolk Stage (Figure 9)

Oocytes with a nuclear and total diameter of 70-80 um and 210-260 um
respectively, develop small eosinophilic yolk globules between the yolk vesicles.
These globules are first noticed with a diameter of 3-5 um and they rapidly
increase to 8-10 um while still restricted to the outer half of the cytoplasm.

(d) Secondary Yolk Stage (Figure 10)

During this stage the yolk globules rapidly increase in size to a maximum
of 15 um. The nucleus diameter and total diameter increases to 80-90 um and
330-360 um respectively. As the globules increase in size they occupy all the
remaining free cytoplasm. The largest fatty droplet increases to a maximum of
25 um, The expansion of the yolk and fat within the cytoplasm restricts the yolk
vesicles to a narrow band around the periphery of the cytoplasm, the largest
vesicle being reduced to about 6 um.

Fig. 8. Yolk vesicle stage oocyte (YU) with peripheral yolk vesicles (YU).

Fig. 9. Primary yolk stage oocytes with yolk vesicles (yv) and yolk globues (yg). Fatty
droplets (fd) appear around the nucleus.

Fig. 10. Secondary yolk stage oocyte with fatty droplets (fd) and yolk globues (yg). In
tertiary yolk stage, yolk globules begin to rupture.

Fig. 11. Interphase deuterogonium (De).

Fig. 12. Spermatogenic maturation stages. Primary spermatogonia (SPGI), Secondary
spermatogonia (SPGII), primary spermatocytes (SPC I), secondary spermatocytes
(SPC II), spermatids (ST) and spermatozoa (SZ).

Fig. 13. Perinucleolar oocyte (PN) and atretic bodies (AB) in a testis.

(Pigs.8, 95°10, 13. x 370; Pig. 11 x-3,300; Fig. 12 x 700)

Aust. Zool. 19(2), 1977 195

G. R. McPHERSON

(e) Tertiary Yolk Stage (Figure 10)
Oocytes at this stage are of similar dimensions to secondary yolk oocytes.

The yolk globules rupture, releasing their finely granular basophilic contents into
the cytoplasm.

(£) Mature Oocytes

The final stage in oocyte maturation occurs when the external membrane of
the tertiary yolk oocyte rapidly expands to a diameter of about 600 um. The
increase in cytoplasmic volume decreases the density of the free yolk material.
The result of this is a large translucent oocyte with a single large fat droplet up
to 150 um. These mature oocytes are released into the ovarian lumen.

SPERMATOGENESIS
(a) Protogonia, Deuterogonia and Primary Spermatogonia

Protogonia are seen in the interstitial areas and are similar to those seen
in ovaries. These cells divide to form an interphase deuterogonia which in
testicular tissue rapidly become basophilic, although the nucleolus and the
distinctive cytoplasm/nucleus relationship (ratio of 1:6) are evident (Figure 11).
It is an increase in basophilia that marks the male orientation of deuterogonia. In
this study, this basophilic stage is referred to as the primary spermatogonium stage.
In the later stages of primary spermatogonium chromatin condensation, there is a
further increase in basophilia as well as a reduction in the nucleus diameter from
4.1um to 3.4um, and the total cell diameter from 7.1 to 5.1 um. The presence
of a primary spermatogonial stage prior to the regular, the intensely basophilic
spermatogonial phase, reported by most workers, has been described in only two
teleosts, Couesius plumbeus (Ahsan, 1966) and Salvelinus fontinelis (Henderson,
1962). In both these teleosts, in the primary germ cell, there is a prominent
nucleolus and a relatively clear nuclear reticulum.

Occasional mitotic divisions are seen among cysts of late primary spermatagonia
to produce secondary spermatogonia.

(b) Secondary Spermatogonia

Secondary spermatogonia represent a marked increase in haematoxylin
affinity over the latter stages of the primary spermatogonial chromatin condensation
(Figure 12). The nucleus is slightly ovoid with a mean diameter of 2.8 um and
a total cell diameter of 3.9um. The condensed chromatin occupied the entire
nucleus and the presence of a nucleolus is only evident with Feulgen staining which
is not specific for RNA.

As reported by Ahsan (1966) the divisions of the primary spermatogonia
are slow or infrequent and the large cysts of secondary spermatogonia are produced
by numerous mitotic divisions within the cyst.

(c) Primary Spermatocytes

Within many cysts of secondary spermatogonia a few nuclei show a further
decrease in nuclear diameter to the primary spermatocyte stage. They have a

196 Aust: Zool. 19(2), 1977

SEX CHANGE IN THE WRASSE

nuclear and total diameter of 2.3 um and 3.6 um respectively. No nuclear detail
is evident (Figure 12). Apart from the spermatozoa, these cells are the most
commonly seen in the testis.

Meiotic figures are often seen within primary spermatocyte cysts. Leptotene
is characterised by a slight increase in nuclear diameter from the primary
spermatocyte average of 2.3 um to 2.6 um. If a nucleolus is seen it is very small.
Zygotene figures are not common. They lack a nucleolus, return to the diameter of
interphase spermatocytes, and demonstrate fine and clearly demarkated chromatin
threads that extend across the nucleus. The chromosomes at the pachytene stage
are difficult to resolve. They appear in 5-6 large clumps which are radially
arranged in a star-like pattern. Metaphase spreads are common.

(d) Secondary Spermatocytes

Secondary Spermatocytes have a nuclear diameter of 1.9 um (Figure 12). The
chromatin is hightly condensed and is often located near meiotic prophase figures.
These cells divide (Meiosis II) to form spermatids.

(e) Spermatids

Spermatids have a nuclear diameter of 1.4um (Figure 12). They are
distinguished from secondary spermatocytes by their smaller nuclear diameter and
crimson staining with Azan (in contrast to yellow-orange) and from spermatoza
by their !arger nucleus, and absence of a flagellum.

({) Spermatozoa

Spermatozoa have a nuclear diameter of 1.0 um and a fine long flagellum.
Often they appear within the spermatogenic lobules (or tubules) in “parachute
shaped” clumps (Pollard, 1972), although they are usually distributed with
varying densities within the lobules (Figure 12),

INTERSEXUALITY

Spermatogenic Cells in Ovaries

The only evidence of intersexuality in females was seen in two specimens
(169 and 196 mm) from the April sample in which a few small cysts of primary
spermatogonia were found. They were always located in the lamellar epithelium
and were sometimes adjacent to groups of early meiotic prophase oocytes. The
cysts were rare and there was no evidence to suggest an impending sex inversion
as perinucleolar oocytes were developing and the colour pattern was quite normal.
It is more likely that these cells were formed during the period of oocyte
proliferation that appears to occur around April when deuterogonia normally
differentiated to leptotene oocytes. However, a few entered the male germ cell line.

Oocyte Remnants in Testes

Early meiotic stage oocytes were only found in one male of 206 mm SL with a
prominent anal bar, no red on the sides and 15% area of primary spermatogonia.

Aust. Zool. 19(2), 1977 197

G. R. McPHERSON

Figure 13 shows an intact diplotene or perinucleolar oocyte surrounded by
testicular tissue. The empty vaculoes located around this oocyte were probably
formed around oocytes which have since been resorbed. In this male, the vaculoes
are surrounded by connective tissue; in established males similar structures are
usually filled with spermatogenic stages.

The only other oocyte remnants were of collapsed mature oocytes and most
commonly, atretic bodies (or atretic oocytes) that remain throughout the
testicular phase of the gonad (Figure 13). Atretic bodies are formed only from
oocytes in the yolk globule stage of development. Upon atresia, the granulosa cells
that surround the mature oocyte enter through the ruptured membrane and
envelop and resorb the yolk and fat material. In haematoxylin and eosin stained
sections, the bodies are irregularly shaped masses of faintly eosinophilic cytoplasm
with small densely basophilic nuclei. No comparable structure appears in ovaries,
yet frozen sections of testes stained with Sudan Black B showed that atretic bodies
had numerous black and dark green staining masses, identical to the black
staining fatty droplets and the dark green yolk globules of similarly stained yolky
oocytes. Intact yolk globules may sometimes be identified in haematoxylin and
eosin stained sections viewed under bright blue florescent microscope (with a
Schott BG 12 blue filter and a Wratten G 15 yellow filter). In some bodies large
round globules fluoresce with a bright green colour (as do the many small nuclei),
like the yolk globules of mature oocytes.

The remains of mature oocytes are more easily identified. They never contain
connective tissue, but have a basophilic granular cytoplasm, give a positive lipid
reaction with Sudan Black B and retain an intact, though grossly misshapen outer
membrane.

DISCUSSION

Reinboth (1962) presented the only description of gametogenesis in sex
reversing species within the sub-order Labroidei. He failed to provide a description
of the stem germ cells in Coris julis, although he recognised their similarity in
females and males and called them deuterogonia following the terminology of
D’Ancona (1945)). Reinboth believed that deuterogonia merely transformed into
oogonia or spermatogonia and found great difficulty in differentiating between
early forms of the two, especially in females where both were sometimes present.
In P. gymnogenis, the stem germ cells (protogonia) divide mitotically to form
interphase deuterogonia, a transient cell type that rapidly differentiates as a
leptotene oocyte or a primary spermatogonium.

In two descriptions of natural sex reversal in labrids, in Coris julis (Reinboth,
1962) and Halichoeres poecilopterus (Okada, 1964) rapid spermatogenic
proliferation was concomitant with ovarian degeneration. In Coris julis the
testicular tissue was restricted to the periphery of the gonad, the ovarian lamellae
collapsing completely. In contrast, Okada reported that the germ cell proliferation
originated from stem germ cells located within the lamellae. Okada’s photo-

198 Aust. Zool. 19(2), 1977

SEX CHANGE IN THE WRASSE

micrographs demonstrate that there was not a marked reduction in the size of
the gonad with sex inversion; this was probably because H. peocilopterus usually
transforms at the end of the breeding season when the gonads are free of large
yolk stages and mature oocytes. A marked reduction in the gonad size was a
feature of sex inversion in P. gymmnogenis as the gonads of the transforming
females were often distended with large maturing oocytes.

On a morphological basis there appears to be no difference between the
stem germ cells in females and males of P. gymmnogenis, yet they divide and
differentiate into either female or male germ cells. In the crustaceans Charniau
Cotton (1965) has shown that the titre of hormone produced by the androgenic
gland decides whether the gonia enter oogenesis or spermatogenesis. The reviews
of Pickford and Atz (1957), Hoar (1965) and Barr (1968) have shown that
teleosts present a more complicated picture; however, it is generally accepted
(Reinboth, 1970) that undifferentiated gonia are sexually bipotent and may be
directly influenced by heterologous sex hormones.

In one male P. gymnogenis the remnants of a few small meiotic stage oocytes
were seen in the central zone of a lamella, and not around the lamellar wall
where meiotic oocytes are normally found in functional temales. Although the
nuclei were smaller than those of normal meiotic oocytes, they have been shown
to be oocytes, and not degenerate lobule boundary cells. Okada (1964) believed
that in transforming H. poecilopterus, oocytes were formed for some time, but
gradually decreased in total diameter from 25 um to 5um. As Okada (1964)
makes no reference to normal meiotic stage oocytes it is likely that the 25 um
cells were diplotene oocytes which had been present prior to the sex inversion.
These small early meiotic prophase stages may have differentiated during the sex
inversion period if low concentrations of oestrogen were still present, as was the

case in the natural sex inversion of the protogynous synbranchid eel, Monopterus
albus (Chan and Phillips, 1969).

ACKNOWLEDGEMENTS

I wish to thank Mr. V. Ivantsoff of Macquarie University for encouragement and the
supervision of the work and also for proof corrections. My thanks also go to Dr. G. Alcorn
and Mr. R. Oldfield from Macquarie University for advice on the preparation of high
resolution photomicrographs, and to Mr. G. French for producing the photomicrographs.

REFERENCES

AHSAN, S. N. (1966). Cyclical changes in the testicular acuy ity of Lake Chub, Couesins
plumbeus (Agassiz). Can. J. Zool. 44, 149-160.
BARR, W. A. (1963). The Endocrine control of the Sexual Cycle in the Plaice, Pleuronectes

platessa (L.) Il. The Endocrine Control of Spermatogenesis. Gen. comp. Endocrinol.
3, 216-225.

BARR, W. A. (1968), Patterns of Ovarian Activity. In: Perspectives of Endocrinology
pp 164-238, E. J. W. Barrington and C. Barker-Jorgensen, eds Academic Press, London.

CHAN, S. T. H. and J. G. PHILLIPS (1967). The structure of the gonad during natural
sex reversal in Monopterus albus. J. Zool., Lond. 151, 129-141.

Aust. Zool. 19(2), 1977 199

G. R. McPHERSON

CHARNIAUX-COTTON, H. (1965). Hormonal control of sex differentiation in invertebrates.
In: Organogenesis. pp. 701-40. R. L. de Haan and H. Ursprung. Holt, Rinehard and
Winston, New York.

D’ANCONA, U. (1945). Sexual Differentiation of the Gonad and the Sexualisation of the
Germ Cells in Teleosts. Nature, Lond. /56, 603-4.

HARDISTY, M. W. (1965). Sex differentiation and gonadogenesis in Lampreys. Part 1. The
ammocoete gonads of the brook lamprey, Lampetra planeri. J. Zool. 146, 305-345.
HENDERSON, N. (1962). The annual cycle in the testis of the Eastern Brook Trout,

Salvelinus fontinalis. Can. J. Zool. 40, 631-641.

HOAR, W. S. (1965). Comparative physiology: Hormones and reproduction in Fishes.
Ann. Rev. Physiol. 27, 51-70.

LILLIE, R. D. (1965). Histopathologic Technic and Practical Histochemistry. McGraw-Hill
Co., New York.

OKADA, Y. K. (1965). Bisexuality in sparid fishes. II Origin of bisexual gonads in
Mylio macrocephalus. Proc, Joys. Acad. 41, 295-9.

OKKLEBERG, P. (1914). Hermaphroditism in the Brook Lamprey. Science 39, 478.

PICKFORD, G. E. and J. W. ATZ. (1957). The Physiology of the Pituitary Gland in
Fishes. N.Y. Zool. Soc., New York.

POLLARD, D. A. (1972). The biology of a landlocked form of the normally catadromous
salmoniform fish Galaxias maculatus (Jenyns) III. Structure of the Gonads Aust. J.
mar. Freshwat. Res. 23, 17-38.

REINBOTH, R. (1962). Morphologische and funktionelle Zweigeschlechtlichlceit bei marinen
Teleostiern (Serraridae, Centracanthidae, Labridae). Zool. Jb. (Allg. Zool. Physiol.
Tiere). 69, 405-480.

REINBOTH, R. (1970). Intersexuality in Fishes. In: Hormones and the Environment.
Mem. Soc. Endocrinol. /8, 516-543.

SATOH, N. and N, EGAMI, (1972). Sex differentiation of germ cells in the teleost,
Oryzias latipes, during the normal embryonic development. J. Embryol. exp. Morph.
28, 385-395.

200 Aust. Zool 19(2), 1977

A new Genus and Two New Species in the Family
Mithrodiidae (Echinodermata : Asteroidea) with
Comments on the Status of the Species of
Mithrodia Gray 1840

ELIZABETH C. POPE
8 Loorana Street, East Roseville, N.S.W. 2069. Australia.

and

F. W. E. ROWE
The Australian Museum, 6-8 College Street, Sydney, N.S.W. 2900, Australia.

ABSTRACT

A new genus and two new species of mithrodiid asteroids from the Pacific and '
Indian Oceans are described. Mithrodia gigas Mortensen, from South Africa,
is referred to the new genus. The status of the species of Mithrodia is discussed.
Mithrodia fisheri Holly and Mithrodia clavigera (Lamarck) are retained as valid
species but Mithrodia victoriae Bell and Mithrodia bradleyi Verrill are provisionally
referred to the synonymy of M. clavigera.

INTRODUCTION -

During a visit to the Aquarium of Nouméa, New Caledonia, in 1969, the
senior author was brought a giant asteroid, taken locally by the Aquarium’s
scuba divers. It clearly represented an undescribed form of the family Mithrodiidae.
For several years this specimen was thought to be unique. Subsequently a specimen
was sent from Guam by Dr. L. G. Eldredge in 1971 and a juvenile from the
Ogasawara Islands, taken in 1974, was forwarded to us by Dr. M. Yamaguchi.
Finally two further specimens were taken in 1975 by M. Labout and Mr. A.
Birtles, near the original locality, in the lagoon off Nouméa. Photographic evidence
also exists of an undoubted specimen of this species from the Philippine Islands,
recorded on the cover of a Japanese periodical, “Marine Diving” 1972, No. 15
(cover picture).

Two specimens from the Seychelles Islands, Indian Ocean, forwarded to us
by Miss A. M. Clark of the British Museum (Natural History), proved to be
closely allied to but specifically distinct from the Pacific specimens.

Research of the literature indicated that both of these forms are not only
closely related to Mithrodia gigas Mortensen from South Africa, but also to the
“peculiar specimens of Mithrodia’’ described and illustrated by Fisher (1906).
It seemed to both of us, however, that their features were sufficiently distinctive

Aust. Zool. 19(2), 1977 201

ELIZABETH C. POPE and Eo W. Ex ROWE

to separate them generically from other species included in the genus Muthrodia
Gray by Engel, Dilwyn John and Cherbonnier in their 1948 revision of that genus.
We also concluded that some of these authors’ findings were incorrect and we
therefore felt it appropriate that, while describing and justifying the new taxa,

we should include our comments on the status of species within the genus
Mithrodia.

DESCRIPTIONS

Order: Spinulosida.
Family: Mithrodiidae.
Genus Mithrodia Gray, 1840 p. 288.

DIAGNOSIS.

Slender-armed forms, usually 5-rayed (4- to 6-rayed), with R (centre of
disc to arm tip) up to about 300 mm, r (centre to edge of disc) up to about
20mm, so that R:r lies between 7.5: 1 and 15:1 (usually about 10-12= 1):
Arms tapering to a fairly acute tip. Skeleton open or compact, reticulate, with
plates and trabeculae bearing tubercles. The whole (body and spines) covered
by a thick skin beset with scales or thorn-like granules.

Large, stout spines occur actinally and abactinally, forming up to 9
longitudinal rows (including the rows of subambulacral spines). Tubercles or
spines absent from the papular areas. Marginal plates inconspicuous but their
position indicated by two of the rows of stout spines. Infero- and supero-marginal
rows separated, connected by trabeculae. Actinal intermediate areas narrow, with
only one row of plates. Abactinal spines and spines of the first actinal intermediate
row usually longer than the inner, subambulacral spines and first actinal spines
occurring irregularly, one adjacent to every 1 to 7 of the subambulacral spines.
Furrow spines finger like, 10 to 12 in webbed fans. Pedicellariae multi-valved,
typical of the family, present actinally and abactinally.

TYPE SPECIES:
| Asterias clavigera Lamarck, 1816
= Mithrodia spinulosa Gray, 1840. Type species by
monotypy: a synonym of M. clavigera fide Perrier,
1875; de Loriol, 1885 and subsequent authors.

OTHER SPECIES INCLUDED:
Mithrodia fisheri Holly, 1932; (M. bradleyi Verril,
1867; M. victoriae, Bell, 1882 are provisionally
referred to synonymy of M. clavigera)

Genus Thromidia gen. nov.
DIAGNOSIS:
Large, obese, 5-rayed forms with R up to 350mm, r up to 8) mm. R:r as
4:1 up to 7:1. Arms with more or less parallel. sides, tapering only distally

202 Aust. Zool. 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

to a blunt, rounded tip. Skeleton is open or compact, reticulate. Skeletal plates
bear small tubercles, the whole being covered by a thick skin with scale- or
thorn-like granules. The larger, stout spines are restricted to a subambulacral plus
one (rarely two) actinal rows. Tubercles occur in the papular areas. Marginal
plates inconspicuous and without spines. Actinal intermediate areas wide, with
several rows of plates. Spines of the first actinal intermediate row usually
shorter than the inner, subambulacral spines and forming a regularly arranged
line, one to each 2 (proximally) — 4 (distally) subambulacral spines. Furrow
spines number up to 12, in webbed fans. Multi-valved pedicellariae, typical of
the family, present only actinally and usually restricted to areas adjacent to and
between the spines of the first actinal and subambulacral rows.

The only exception to the above diagnosed arrangement of spines is seen
in a juvenile specimen (R = 58 mm) bearing an even distribution of small spines
over the whole body (see p. 204, figs. 5-6 and p. 205).

ETYMOLOGY: Anagram of Mithrodia; gender, feminine.
TYPE SPECIES: T. catalai sp. nov.

OTHER SPECIES INCLUDED: T. seychellesensis sp. nov., Mithrodia gigas, Mortensen,
L935:

REMARKS:

This genus is separated from Mithrodia because of its much greater bulk, the
shape of its arms, the restriction of the stout spines to the subambulacral and
first (rarely the second) actinal rows, the regularity of the actinal row of spines,
the wide actinal intermediate area and the presence of tubercles in the papular
areas.

Thromidia catalai sp. nov.
Figs. 1-6 and 9

Mithrodia sp., “peculiar specimen” Fisher 1906, p. 1069, Pl. XXXVII, figs. 2-3.
Mithrodia fisheri, Engel, Dilwyn John and Cherbonnier, 1948, p. 27 part.
(non Mithrodia fisheri Holly, 1932).

MATERIAL EXAMINED-

Holotype, R = 352mm (in life), 283 mm after preservation in alcohol
(Australian Museum No. J. 7792), near base of Tabu Reef, off Ilot Amedée, New
Caledonia, circa 10m depth, on sandy bottom with boulders, 14.1X.1969,
collected by M. B. Conseil and M. G. Bargibant for the Aquarium of Nouméa. Two
paratypes: (Australian Museum No. J. 9926 R = 238mm) and British Museum
(Natural History) No. 1976.10.14.1, R = 225-250 mm after preservation and
drying), both taken in the same locality as the holotype, on 11.VII.1975, collected
by M. M.P. Labout and Mr. A. Birtles in a depth of c. 10 m; one paratype, (R =
245 mm preserved dry) from the vicinity of Guam Island, taken by scuba divers

Aust. Zool. 19(2), 1977 203

ELIZABETH G.. POPE and” Fo W.&,

Figs. 1-6. Thromidia catalai n. sp. 1, holotype, A.M. No. J7792, abactinal view, R = 283 mm.
2, paratype, B.P.B.M. No. W2507, actinal view of one arm, R= 245 mm. 3-4,
juvenile, A.M. No. J10134, whole abactinal view (3); actinal view of one arm (4),
R = 125mm. 5-6, juvenile, U.S.N.M. No. 32271, abactinal (5) and actinal (6)
views, R = 58 mm.

Aust. Zool 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

(no other data supplied), deposited at the Bernice P. Bishop Museum, Hawaii
(W2507); one specimen (juvenile), R = 125 mm, preserved dry (Australian
Museum No. J.10134) from Miyano-hama, Chichi-jima, Ogasawara Islands, 10 m
depth, collected by Mr. T. Matuura, 24.IJI.1974; one specimen (juvenile) R =
58 mm (United States National Museum No. 32271) ‘‘Albatross” Station 4158,
off the Bird Islands, Hawaii, 38-45 m, on coral and coralline bottom; one specimen
(size not recorded; colour print only), Hansa Bay, 200 km north of Madang,
Papua New Guinea, print sent by Dr. M. Jangoux, December, 1976.

DESCRIPTION.

A large obese species. In life. adults range from R = 355 — 384mm,
r = 82 — 90mm, R/r = 4 — 4.6; after preservation R = 220 — 296 mm,
r = 50 — 66mm and R/r = 4 — 5.2. The arms are slightly constricted at their
origin from the disc, which is relatively small. The five arms are cylindroid with a
taper which becomes marked only from approximately 2/3 R. The madreporite is
single, small and situated halfway between the centre of the disc and the interradial
margin (Fig. 1).

The skeleton of plates and radiating trabeculae form an open meshwork
enclosing large papular areas (up to 10-11 mm in adults) in which finer, minute
ossicles are present. Tubercles (about 1 mm wide by 1.5 mm high) are extremely
numerous and crowded over the whole surface, occurring on the skeletal elements.
Smaller tubercles occur in the papular areas. On the distal 1/7 of the arm
only the larger tubercles are present, becoming more widely spaced. There are
no stout spines abactinally (Fig. 9).

The whole surface, including all the tubercles and actinal spines (even
including a few of the furrow spines) is covered by a thick skin in which are
scale- or thorn-like granules. These granules become enlarged and pointed towards
the apical regions of the spines and tubercles. Marginal plates are inconspicuous
and do not bear spines.

Actinal intermediate areas are broad, with 5-6 rows of plates between the
adambulacral and ventral lateral (inferomarginal) margin (Fig. 2). Adambulacral
plates each bear a webbed fan of 6-7 (sometimes 5-8) finger-like furrow spines,
backed by a single, stout, scale-covered subambulacral spine, up to 8 mm high
and 2mm in breadth. The inner side of the subambulacral spine is often rubbed
bare. A regular row of spines which are slightly shorter than the subambulacrals
(6mm high and 3 mm in breadth) are found on the first actinal row of plates.
Each actinal spine occurs adjacent to every second subambulacral spine. A second
tow of spines, intermediate in size between those of the first actinal row and the
tubercles covering the rest of the actinal surface, is evident particularly on the
distal part of the arms (Fig. 2).

Pedicellariae are multi-valved, occurring only actinally between the first
actinal and subambulacral rows of spines. Tube feet are large, with a double
ampulla and large terminal disc.

Aust. Zool. 19(2), 1977 205

ELIZABETH ‘C:.’POPE (and FW. En ROWE

A dried juvenile R = 125 mm and r = 30mm, R/r = 4.5 from Ogasawara
Islands, which judging from its photograph in life has become very shrunken,
has tubercles which are restricted to the plates and do not occur on the trabeculae
(Fig. 3). Usually only one or two tubercles occur in its papular areas. The spines
of the second row of the actinal intermediate plates are more prominent,
particularly distally, than in the adult specimens. There are only three rows of
actinal intermediate plates (Fig. 4).

The larger of the two “peculiar specimens’’ recorded from Hawaii by Fisher
(1906) is only about half the size of the holotype but his description and
figures of it (plate XXXVII, figs. 2 and 3) do not indicate any major differences
from the one we examined from the Ogasawara Islands. We have, unfortunately,
been unable to locate the whereabouts of Fisher’s larger specimen. The smallest
specimen we examined (USNM No. 32271), labelled “M. bradleyi Verrill’,
coming also from the vicinity of Bird Islands, Hawaii, but from ‘‘Albatross”’
Station 4158, has proved something of an enigma (Figs 5-6). We believe it to
be the smaller of the two “peculiar specimens” of Mithrodia recorded by Fisher
(1906, p. 1096). Its Station number and locality data agree with Fisher’s record
but there is a discrepancy in the measurement he gives for R (= 38mm
according to Fisher). In all other respects it agrees with his short description.
We found R = 58mm, r = 12mm, R/r = 4.8. Unfortunately Fisher did not
figure this specimen so we cannot be absolutely certain that it is the specimen
to which he refers. However, a simple typographical error (mistaking a 5 for a 3)
would account for the difference in the measurements. There is no doubt, however,
that this is a juvenile Thromidia. It exhibits characters which are not visible in
the larger juvenile from Ogasawara Islands. This at first led us to believe it
might represent a fourth species of Thromidia since it has small spines scattered
abactinally. These are only slightly smaller than the subambulacral spines but
are similar to those of the first actinal row. It is not unknown, however, for
juvenile forms in this family to have abactinal spines which later disappear with
growth (Engel et al. 1948, pp. 7-10). The position of the marginal plates is
determinable by the spines carried on them. This is a feature not shown by the
adults. The papular areas vary in diameter between 2-2.5mm. In our present
state of knowledge we agree with Fisher, that this is a still earlier stage of
his larger “peculiar specimen”. However, we conclude further that it should also
be regarded as a juvenile of the giant Pacific species T. catalai. The colour note
given by Fisher adds convincingly to the argument for considering this specimen
a juvenile of T. catalai.

COLOUR.

In life (adults), uniformly pink abactinally, deeper pink actinally, with arm
tips cinnamon-brown; actinal and subambulacral spines rose-red with their
flattened tips cream; furrow spines cream; tube feet whitish with the terminal
disc muddy-brown. A colour print of the live juvenile from Ogasawara Islands
shows it to be rose red, with arm tips a darker red. Fisher recorded the life
colour of the larger of the ‘“‘peculiar specimens” as “dull, light cinnamon-pink and

206 Aust. Zool. 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

maroon at end of the arms, the cinnamon mottled in places with buff. Actinal
surface is light pinkish buff or vinaceous, darkest on tubercles. Ambulacral feet
raw sienna’.

HABITS-

In the field the starfish have been observed lying in the open, on the bottom,
in depths of 10m or more. In one instance one was observed with its stomach
everted over a grey, encrusting sponge, growing on a coral boulder (A. Birtles,
personal communication). The holotype and the larger of the two specimens
captured by Labout and Birtles were kept alive over periods of up to 10 months
in the Aquarium of Nouméa and Mme. Catala reported that no solid food (flesh
of fish, crustacea and molluscs) such as was fed to the fish and other inhabitants
of their tank was taken by Thromidia, but it survived by feeding on encrusting
growths on the boulders and side walls of the tank. The animals are more active
by night and are capable of moving on their tube feet at speeds of up to 25 mm
per second. The specimen now selected as the holotype A.M. J7792_ lived
unmolested for several months in the largest tank at the Aquarium which it shared
with a number of species of fish, molluscs and stingrays. However, a newly
introduced fish (a balliste) immediately attacked the starfish, tearing a hole in
one arm. The attack was not repeated but the starfish responded to the injury
by emitting a stream of whitish eggs upon which the fish fed. At this stage the
Thromidia was removed from the tank for preservation.

When Thromidia lay on the bottom it was frequently observed to become
covered abactinally by pebbles and shellgrit which rained down on it as a result
of the feeding activities of certain species of fish. It seemed incapable of cleaning
itself while flat on the bottom but the rubbish fell off when the starfish climbed
vertical surfaces. The starfish could move on its tube feet with equal facility over
a loose or solid substratum.

ETYMOLOGY-

The species is named in honour of Mme. Stucki and Dr. René Catala of
the Aquarium of Nouméa who sponsored the collection of the Nouméa specimens
and also kept Thromidia alive in their Aquarium making notes and observations
on its behaviour.

Thromidia seychellesensis sp. nov.
Figs. 7-8 and 11

Mithrodia fisheri, Jangoux, 1974, p. 791 (non M. fisheri Holly, 1932).

MATERIAL EXAMINED:

Holotype R = 123 — 124mm (British Museum [Natural History] No.
1974.9.25.18); Recife d’Anse a la Mouche, Mahé Isle, Seychelles, Indian Ocean,
collected by Dr. M. Jangoux, 1972; Paratype R = 135mm (B.M. [N.H.] No.

Aust. Zool. 19(2), 1977 207

ELIZABETH C. POPE and F. W. E. ROWE

208 Aust. Zool 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

1976.10.14.2) Seychelles, Cousin Id, 12m, October, 1972, collected by R. A.
Birtles.

DESCRIPTION-

A species with five cylindroid arms which have parallel sides along the
greater part of the length, tapering only slightly to a rounded, blunt tip. R =
ioe oi, ==) 22 23mm, R/r = 5.5 (Holotype); R = 135 mm
r = 27 — 28mm, R/r = 4.9 (Paratype). The madreporite is rather insignificant
and positioned half way between the centre of the disc and the interradial margin
Chis. 7).

The skeletal plates and radiating trabeculae form a compact meshwork
enclosing small papular areas (up to 3.5 mm diameter). Tubercles (about 0.6 mm
wide by 0.75 mm high) are numerous and crowded over the actinal and abactinal
surfaces, occurring on the skeletal elements. A smaller tubercle occurs in each
of the papular areas (Fig. 11). On the distal 1/5 — 1/4 of the arms the plates
are rounded convex and are closely packed together, giving the tip of the arms
the appearance similar to that of a cobbled pavement (Fig. 11). The distal papular
areas are so small that they contian only one or two papulae. Occasionally the
smaller tubercles persist on these papular areas. Trabeculae connecting the larger
plates are very reduced. There are no stout spines abactinally.

The whole surface, as in T. catalai, is covered with a thick skin beset with
scale- or thorn-like granules. These granules become slightly enlarged towards
the apical region of the spines and tubercles but the distal arm plates are
characterised by the even size of the granules covering them (Fig. 11). The
marginal plates are inconspicuous.

Actinal-intermediate areas are broad, with 3-4 rows of plates between the
adambulacral plates and the ventro-lateral (inferomarginal) margin (Fig. 8).
Adambulacral plates each bear a webbed fan of 6-8 furrow spines backed by a
single, stout subambulacral spine up to 4-5 mm high and 1.5mm in breadth.
The inner side of the subambalacral spine is often rubbed bare. A regular row
of spines, shorter than the subambulacrals (3.5 — 4.5mm by 1.5 — 2.0 mm) is
found on the first actinal row of plates. The actinal spines are spaced, proximally,
adjacent to every second subambulacral spine, but more distally (beyond 1/2R),
particularly on the holotype, adjacent to every 3rd or 4th subambulacral spine.
A second row of spines, intermediate in size between those on the first actinal

Figs. 7-8. Thromidia seychellesensis n. sp., holotype, B.M. (N.H.) No. 1974. 9.25. 18,
whole abactinal view (7); actinal view of one arm (8), R = 123-124mm
Fig. 9. T. catalai n. sp., paratype, B.P.B.M. No. W2507, abactinal view of arm, R = 245 mm.
Fig. 10. 7. gigas (Mortensen), holotype, S.A.M. No. A 22561, abactinal view of arm,
R = 330-347 mm.
Fig. 11. T. seychellesensis n. sp., holotype, B.M. (N.H.) No. 1974.9. 25.18, abactinal]
view of arm-tip, R = 123-124mm

Aust. Zool. 19(2), 1977 209

ELIZABETH C. POPE and F. W. E. ROWE

row and the tubercles covering the rest of the actinal surface, is evident particularly
on the distal parts of the arms (Fig. 8).

Pedicellariae are typical of the family and distributed as in T. catalai. It is
possible that the specimens of seychellesensis described above represent a young
stage of the species.

COLOUR:

The dried holotype is museum colour with darker tips to the arms. The
dried paratype is generally museum colour, with some trace of pink abactinally
and actinally while the arm tips are a darker museum brown. A colour transparency
sent to the authors by Dr. Jangoux (1976, pers. comm.) shows the species to
be uniformly light orange abactinally, with arm tips darker.

HABITS.

This species is relatively common in the Seychelles Archipelago (Jangoux,
1976, pers. comm.) where Jangoux (1974) records it frequently as host to the
fish parasite, Carapus homei. The fish occurs within the body cavity of the starfish.

ETYMOLOGY: Named after the area where it was taken.

Thromidia gigas (Mortensen)
Fig. 10

Mithrodia gigas Mortensen, 1935, p. 1, fig. 1, plate 1; Cherbonnier, 1975, p.639,
pls. I-II.

MATERIAL EXAMINED-

Holotype (South African Museum No. A 22561) off Point Morgan, East
London, South Africa, in a depth 45-54m, on 28 January, 1934, collected by
Mr. Bell Marley. The starfish was taken on a fish-hook, after taking the bait.

DESCRIPTION:

There is little to add to the detailed description given by Mortensen (1935).
There are, however, some slight discrepancies in his measurements and in the
number of furrow spines Mortensen recorded. We found R = 330 — 347 mm,
r = 45mm, R/r = 7.3 — 7.7, and the furrow spines number from 5 — 6
generally and sometimes up to 8 per plate. We have included a new photograph
of an arm tip of the holotype (Fig 10) for direct comparison with those of
T. catalai and T. seychellesensis.

Cherbonnier (1975) has recorded a specimen off the west coast of Madagascar
thus extending the range of the species. His specimen is about two thirds of the
size of the holotype and rather contorted. Whether the measurements given by
Cherbonnier correspond to the conventional way of measuring R and r is unclear.

-210 Aust. Zool. 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

We think that by “le diametre due disque . . .” Cherbonnier may refer, in fact,
to r giving his specimen a R/r ratio of 6.6 — 7.6, and this conforms with his
illustration and our diagnosis of the genus.

COLOUR:

In life the holotype was purplish-pink abactinally with the tips of the arms
more cinnamon red, actinally it was pale yellowish, with the tube-feet white.

COMPARISON OR f CAIALAL 1” SEYCHELLESENSIS. AND £.° GIGAS

T. catalai differs markedly from T. seychellesensis (comparing two specimens
of similar size, one from each species), in the arrangement of the skeletal reticulum,
size of the papular areas and arrangement of plates and tubercles near the arm
tips. Both of these new species are distinguished from T. gigas in their arm
proportions, which are shorter and stouter than in the South African species,
and in the lack of the very prominent, globular bosses on the arm tips.

Kee 1©® SPECIES OF THROMIDIA
ie Rh /t =65-/.6/1; atm tips with larse
globular bosses (5 mm diameter) (Fig. 10);
S.E. Africa (East London) — west coast

Ip Lea eiCiaS Cie nce Renamed Behe Neaaca Seat ive sla Ree ROP EO T. gigas (Mortensen )
lt’ R/r < 6/1, arm tips without large
SUIS SUTIN: | Gy OSS A ee ee eae ive al ee ec fa ora Oreo hn NSE 2

2 OR /r 949 55/1 arm tips with closely

packed rounded, slightly convex plates,

covered with even granules, giving cobbled

appearance (Fig. 11); Seychelles (Western Indian

RO) Cay ee ee echoes T. seychellesensis sp. nov.
2 RR /t == 4252/1, arm. tip with spaced plates

bearing small tubercles (Fig. 9); Pacific Ocean

(Hawaii, Japan, Philippines, New

Grileclnraicten Nien Gri Sek es ooo ccccadeccescscncnteasschecocaccansrucnsveetettne T. catalai sp. nov.

DISCUSSION

The first two representatives of Thromidia were collected by the USS.
Fisheries ship “Albatross” off the Bird Islands, Hawaii in 1902. These were
the juvenile specimens referred to by Fisher (1906, p. 1069) as “A peculiar
specimen of the genus Mithrodia’. They differed somewhat from other specimens
of the species collected by the “Albatross” which Fisher identified as Mithrodia
bradleyi Verrill. He did not, however, propose a new name for the peculiar
specimens. We have not been able to locate the whereabouts of the larger of the
two “‘peculiar specimens” but what we believe to be the smaller one is discussed
above (p. 205).

- Aust. Zool. 19(2), 1977 oti

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Oo.

ELIZABETH C.

NEW SPECIES OF MITHRODIIDAE

Holly (1932) described a new Mithrodia from Hawaii, M. fisheri, from a
damaged, obviously juvenile specimen. He wrongly synonymised Fisher’s “peculiar
specimens’”’ with his new species and failed to appreciate that it was the specimens
identified and figured by Fisher (1906) in Plate XXXVI and Plate XXXVII
Fig. 1, as M. bradleyi that in reality represented an adult of his M. fisheri
(see below).

Mortensen (1935), in describing M. gigas, compared it with M. bradleyi
sensu Fisher (1906), non Verrill (1868) and he failed to recognise in M. gigas
differences considered by us to be of generic significance. He listed M. bradleyi
as a valid species, not realising that Fisher’s Hawaiian record in reality referred
to Holly’s species which had been undescribed at the time of Fisher’s report.
The validity of M. bradleyi Verrill, from the west coast of America is dealt with
below (p. 214).

Engel et al. (1948) also did not realise the significance of Fisher’s “peculiar
specimens” but did realise Fisher’s blunder in the identification of M. bradleyi.
They referred both M. bradleyi sensu Fisher and his “peculiar specimens” to
the synonymy of M. fisheri Holly.

There are no further records of the capture of specimens of Thromidia until
the senior author encountered the first of the very large specimens, now the
holotype, of T. catalai, from the lagoon off Noumea, New Caledonia in 1969.
All subsequent known records are set out above.

COMMENTS“ ON THE’ STATUS OF THE SPECIES OF MITHRODIA

In a major review of the genus Mithrodia, sensu lata, Engel, John and
Cherbonnier (1948) listed five species as follows: M. clavigera (Lamarck, 1816);
M. victoriae Bell, 1882; M. bradleyi Verrill, 1867; M. fisheri Holly, 1932 and
M. gigas Mortensen, 1935. Of these M. gigas has been referred in this paper
to the new genus, Thromidia, for the reasons stated. Of the remaining four, we
believe that M. fisheri is clearly defined and that the relationship between
M. clavigera, M. bradleyi and M. victoriae still needs clarification.

Holly correctly recognised a new species, M. fisheri for a Hawaiian form
(Figs. 16-17) even though his description was based on a damaged specimen. We
do not agree with him, however, in identifying Fisher’s “peculiar specimens’? with

Figs. 12-13. Mithrodia clavigera (Lamarck), A.M. No. J5845, abactinal (12) and actinal
(13) views, R = 200mm. (Great Barrier Reef [Capricorn Group]). One
arm broken off.

Figs. 14-15. M. clavigera (Lamarck) (identified as M. bradleyi Verrill), Allan Hancock
ING= 245.5 (2. abactinal (14) and. actinal (15)~ views,” R/ = 165mm.
(California [Mexico]).

Figs. 16-17. M. fisheri Holly, Allan Hancock No. 726 — 1, abactinal (16) and actinal
(17) views, R = 152.5mm (Hawaii).

Aust. Zool. 19(2), 1977 213

ELIZABETH OC; POPE*-and”Fe-W? ‘E- ,ROWE

his new species as we have stated above (p. 213). In fact, part of the confusion
in the recognition of M. bradley: has arisen from Fisher’s (1906) identification
of that species from Hawaii. It appears he had examined at least one of Verrill’s
specimens from La Paz and confused the identity of his Hawaiian specimens
with what he thought to be Verrill’s species. Fisher’s material was clearly in a
good state of preservation when examined and photographed. He probably did
not directly compare his specimens against Verrill’s description (1867) and
(1914) of the holotype of bradleyi but rather with one of the poorly preserved
La Paz specimens, this being inferred in a letter which he subsequently wrote
to, and was quoted by, Engel et al (1948, p. 18) admitting his mistake. We
therefore concur with Engel et al in recognising the validity of fisheri for the
reasons set out by them.

The status of M. victoriae is doubtful. Miss A. M. Clark examined the two
syntypes in the British Museum (Natural History) (No. 1879.8.19.97) and
concluded that she could find no difference between them and similar sized
specimens of M. clavigera from Macclesfield Bank, South China Sea (personal
communication to senior author, 1972) and that there seemed little doubt as to
the origin of the specimens from Victoria Bank, Brazil (personal communication
to the junior author, 1976). The junior author examined Bell’s specimens of
M. victoriae in 1975 and was unable to draw any different conclusions from
those of Miss Clark. Engel et al were certainly very doubtful of both the validity
of this species and its actual origin from Brazil, together with a second similar
specimen also labelled ‘Brazil’, held in the Leiden Museum, Netherlands. The
authors therefore believe that M. victoriae should be considered a synonym of
M. clavigera.

That M. clavigera is a valid species is without doubt. However, on referring
to Verrill’s description of the holotype of M. bradleyi (1867) and his subsequent
photographic illustration (1914, pl. CXII) of the type-specimen, we can see
little justification for the separation of this form from clavigera. We have compared
a specimen from Lower California (Allan Hancock Foundation Cat. No. 275.12),
labelled M. bradleyi (Figs. 14-15) with a comparably sized specimen of M. clavigera
from North Reef, Capricorn Group of Islands, Queensland (Australian Museum
No. J5845) (Figs. 12-13) and with a whole range of specimens in the Australian
Museum from localities across the Pacific and from Mauritius, Indian Ocean.
Taking point for point the characters discussed by Engel et al in distinguishing
M. bradleyi, considering the wide variation in the characters of M. clavigera
admitted by those authors, we find it difficult to detect any character which would
justify the recognition of a specific distinction between eastern and western
Pacific forms. The only two features of distinction appear to be the stouter arms
and shorter spines of bradleyi, both features being present also in a large
specimen (R = 225mm, r = 32 mm; A.M. No. J9927) collected from New Cale-
donia. If these features are considered of specific importance then M. bradleyi and
M. clavigera are sympatric (at least in the central Pacific area). It is unfortunate
that the type specimens of M. clavigera and bradleyi cannot be directly compared

214 Aust. Zool. 19(2), 1977

NEW SPECIES OF MITHRODIIDAE

and that the two specimens from La Paz identified by Verrill as bradleyi are in
such a poor state of preservation. Interestingly, Perrier (1875, p. 117) in a
footnote commented that M. bradley: was ‘‘peut-étre identique elle aussi a l’espéce
de l’He de France (M. clavigera)”, though Engel et al did not agree. We conclude
that until a greater amount of comparative material has been examined (Engel
et al compared only 4 specimens including at least 2 poorly preserved specimens
of bradleyi against 47 specimens of clavigera), firm conclusions on the relationship
between these two species are difficult to draw, but we feel that bradleyi is little
more than an eastern Pacific form of the widely distributed and variable
M. clavigera.

DISTRIBUTION

The distribution of the genus Mithrodia, like that of Thromidia, is Indo-
Pacific. The Atlantic records for Mithrodia victoriae (synonym of M. clavigera)
may be doubtful (Engel et al. 1948). Mithrodia clavigera, omitting the above
doubtful record, is therefore distributed from Mauritius and the Red Sea through
the Pacific to Lower California and Panama. Mithrodia fisheri occurs around
Hawaii. It is also reported from Arica, Peru by H. L. Clark (1910) under the
name of M. bradleyi (sensu Fisher) and while it is obvious from his figured
specimen that he was dealing with M. fisheri his suspicions concerning its locality
of origin might be well founded and should be taken into account. Engel and his
co-authors (1948) report a specimen of fisherz from New Ireland, Bismarck
Archipelago (originally identified as M. clavigera by Sluiter 1895) and one from
the Philippine Islands. We have not examined these specimens.

ACKNOWLEDGEMENTS

The authors wish to thank Dr. René and Madame Stucki Catala of the Aquarium of
Nouméa for their generous help in donating specimens of Thromidia to the Australian
Museum and for allowing the senior author to use the facilities of their Aquarium during
the course of this investigation. We also thank them for their continued interest in keeping
two of the specimens alive and under observation for long periods, recording habits and
food preferences.

We also thank George Bargibant and Bernard Conseil (Collectors of the holotype)
and M. Labout of Nouméa and Alistair Birtles of Townsville, Queensland (collectors of
the paratypes) who supplied data as well as specimens. We are also grateful to Professor
L. G. Eldredge of Guam University for allowing us to examine a beautifully preserved
specimen from that island; to Dr. M, Yamaguchi, at present stationed at James Cook
University of North Queensland, Townsville for presenting an important juvenile specimen
from the Ogasawara Islands, accompanied by a colour print of the live animal, to the
Australian Museum thus make it available to us for study; to Dr. M. Jangoux of
Université Libre de Bruxelles for supplying data and colour photographs of T. catalai and
T. seychellesensis; and to Dr. Maria E. Caso for allowing us to examine Se eke of a
Mithrodia from the New Hebrides.

The authors are indebted to a number of institutions and fellow echinodermologists
for the loan of specimens and for supplying data as follows:— to Dr. D. L. Pawson and
Miss Maureen Downey of the U.S. National Museum (Smithsonian Institution) Washington,

Aust. Zool. 19(2), 1977 215

ELIZABETH GUoPOPE rand hav? Ey ROWE

D.C.; to Miss Ailsa M. Clark of the British Museum (Natural History) London, U.K.;
to Dr. D. Devaney of the Bernice P. Bishop Museum, Hawaii; to Dr. T. H. Barry of the
South African Museum, Capetown; to Dr. Janet Haig of the Allan Hancock Foundation,
University of Southern California, Los Angeles and to Dr. E. Kritscher, of the Vienna
Museum for supplying kodachrome photographs of Holly’s type of Mithrodia _fisheri.
Finally, we thank Mr. G. Millen, Photographic Department, The Australian Museum, for
photegraphing specimens illustrated in this paper.

REFERENCES

BELL, F. J. (1882). Descriptions of new or rare species of Asteroidea in the collections
of the British Museum. Proc. Zool. Soc. Lond. 1882, 121-124, 1 pl.

CHERBONNIER, G. (1975). Sur la presence, a Madagascar de l’Astérie Mithrodia gigas
Mortensen. Bull. Mus. natn. Hist. nat. Paris (Zool.) (3) 210 (300), 639-646, pls. I-II.

CLARK, A. M. & F. W. E. ROWE (1971). ‘Monograph of shallow-water Indo-west-Pacific
echinoderms’. London, vii+238 pp., 100 figs., 31pls.

CLARK, H. L. (1910). The echinoderms of Peru. Bull. Mus. comp, Zool. Harv. 52 (17),
321-358, 13 pls.

ENGEL, H.; D. D. JOHN & G. CHERBONNIER (1948). The genus Mithrodia Gray, 1840.
Zeol. Verh. Leiden 2, 1-39, 12 figs., 8 pls.

FISHER, W. K. (1906). The starfishes of the Hawaiian Islands. Bull. U.S. Fish. Comm.
23 (3), 987-1130, 49 pls.

GRAY, J. E. (1840). A synopsis of the genera and species of the class Hypostoma
(Asterias Linnaeus). Ann. Mag. nat. Hist. (1) 6, 175-184, 275-290.

HOLLY, M. (1932). Echinodermata from Pearl and Hermes Reef. Occ. Pap. Bernice
P. Bishop Mus. 10 (1), 1-9, 2 figs.

JANGOUX, M. (1974). Sur I’ “association” entre certaines Asteries (Echinodermata) et des
poissons Carapidae. Rev. Zool. Afr. 88 (4), 789-796.

LAMARCK, J.B.P.A. de (1816). ‘Histoire naturelle des animaux sans vertébres’. Paris.
Bdiot2 2, 522-568.

LORIOL, P. de (1885). Catalogue raissonné des Echinodermes recueillis par M. V.
Robillard a l’Ile Maurice. II Stellérides. Mem. Sos. Phys. Hist. nat. Geneve 29 (4),
1-84, pls 7-22.

MORTENSEN, T. (1935). A new giant sea-star, Mithrodia gigas n. sp., from South Africa.
Ann. S. Afr. Mus. 32 (1), 1-4.

PERRIER, E. (1875). ‘Revision de la collection de Stellérides du Museum d’Histoire
Naturelle de Paris’. Paris, 1-384, Also published in Arch. Zool. exp. gén. 4 (1875),
263-449; 5 (1876): 1-104, 209-304.

SLUITER, C. P. (1895). Die Asteriden Sammlung des Museums zu Amsterdam. Bidjr.
Dierk, 17, 49-64.

VERRILL, A. E. (1867). Notes on the Radiata in the Museum of Yale College, with
descriptions of new genera and species, 2. Notes on the echinoderms of Panama and
West Coast of America, with descriptions of new genera and new species. Trans. Cenn.
Acad; -Aris: Seno, 2) ip 254-322.

VERRILL, A. E. (1914). ‘Monograph of the shallow-water starfishes of the North Pacific
coast from the Arctic to California’, Harriman Alaska Ser. 14 (2 vols). xii + 408 pp..
110 pls.

216 Aust Zool. 19(2), 1977

A Redescription of Periclimenes aesopius (Bate, 1863)
(Crustacea : Decapoda) with Remarks on
Related Species

A. J. BRUCE
Heron Island Research Station, Gladstone, Queensland, 4680

ABSTRACT

The pontoniine shrimp Periclimenes aesopius (Baie, 1863) is redescribed in
detail. It is considered to be endemic to southern Australia. Other records of
this species are referred to P. indicus (Kemp) or P. holthuisi Bruce, species
which may be found in association with actiniarians or other coelenterates, and
which also occur in Australian waters. It is considered probable that P. aesopius
is involved in a similar association.

Periclimenes aesopius is remarkable for the great enlargement of postero-dorsal
lobe of the third abdominal segment. Several closely related species have
conspicuous colour markings on this segment and are also involved in fish-
cleaning symbioses. It is also considered likely that the development of this
process has the result of increasing the effectiveness of this signal and that
P, aesopius may also be a fish-cleaning species.

Periclimenes aesopius is considered to be a member of a widely distributed
group of closely related species, generally associates of actiniarians, including
the Indo-West Pacific species: P. indicus (Kemp), P. tosaensis (Kubo,
P. holthuisi Bruce; the east Atlantic-Mediterranean PP. scriptus (Risso),
P, amethysteus (Risso); and the tropical western Atlantic P. yucatanicus (lves),
P,. pedersoni Chace, P. magnus Holthuis and P. anthophilus Holthuis & Eibl-
Eibesfeldt. The group is not represented in the eastern Pacific region.

INTRODUCTION

The pontoniine shrimp Periclimenes aesopius was first described by C. Spence
Bate as Anchistia aesopia in 1863. It is of particular interest in that it was the
first pontoniine shrimp to be recorded from Australian waters and also that
it has not subsequently been reported from any other localities and is still known
only from St. Vincent Gulf, South Australia. Other records from the Indo-West
Pacific region have proved to be erroneous, and Bate’s species appears to be
endemic to South Australia.

DESCRIPTIONS

Periclimenes aesopius (Bate) (Figs. 1-6).

Anchistia @sopia Bate, 1863, Proc. zool. Soc. Lond., 1863: 502-503, pl. 41, fig. 5.
Anchistia eopia—Haswell, 1882, Catal. Aust. Crust.: 194.

Aust. Zool. 19(2), 1977 QUT.

A. J. BRUCE

Fig. 1. Periclimenes aesopius, ovigerous female. Graduated scale = 3 mm.

Periclimenes aesopius— Borradaile, 1898, Ann. Mag. nat. Hist., (7)2:382.

Urocaris @sopius —Borrodaile, 1917, Trans. Linn. Soc. Lond. Zool., (2) 17:354
(key), 354.

Periclimenes (Periclimenes) aesopius— Kemp, 1922, Rec. Indian Mus., 24:140.
Periclemenes aesopius— Kemp, 1922, Rec. Indian Mus., 24:142-143, fig. 12 —
Hale, 1927, Crust, S. Aust. 1 9G, me 50; 1908 Rees. Aust. Musso a ae
Bruce, 1969, Zool. Meded., Leiden, 43(20):258.

nec Periclimenes (Periclimenes) aesopitus Holthuis, 1952; Siboga Exped. Mon.,
39 a’:34-37, figs. 5-6. — Bruce, 1966, Crustaceana, 10(1): 19, 21, fig. 3b;
19 2A. 8,3, ACE:

nec Periclimenes (Periclimenes) aesopius Johnson, 1961, Bull. Mus. Nat. Singapore,
BO D8 61. i] 5S taboayls

nec Periclimenes aesopius Bruce, 1967, Zool. Verhand., Leiden, 87:51-53.

218 Aust. Zocl. 719(2). 1977

A REDESCRIPTION OF PERICLIMENES AESOPIUS

MATERIAL EXAMINED:

(i) 16,192, Gulf of St. Vincent, South Australia, British Museum (Natural
History), registration No. 68-81, Types. (ii) 2 ovig. °, St. Vincent Gulf, South
Australia, South Australian Museum, Cat. Nos. C626, C628. (iii) 1 ovig. 2, 5
miles off Semaphore Beach, South Australia, South Australian Museum, Cat.
No: C1063:

DESCRIPTION:

A relatively large and slenderly built species of Periclimenes, with a smooth
carapace and abdomen.

The rostrum reaches almost to the end of the antennular peduncle and is
feebly to moderately curved, directed slightly upwards in relation to the
longitudinal axis of the carapace (from the posterior orbital notch to the median
postero-dorsal point of the carapace, in lateral view). The lamina tapers feebly,
with poorly developed lateral carinae, situated nearer to the ventral margin,
which is straight or feebly concave, than to the dorsal margin. The dorsal margin
is armed with from 9 to 11 acute teeth, of which 3 or 4 are situated on the
carapace posterior to the level of the orbital margin. The first tooth is situated
at a level of half the post-orbital carapace length or more from the orbital margin.
The teeth on the carapace are larger and more widely spaced than those on the
dorsal lamina, which gradually decrease in size towards the tip. The ventral
border of the lamina bears 2 or 3 teeth on the distal fourth. The orbit is
obsolescent and there is no supra-orbital spine. The inferior orbital angle is
acutely produced, with a short medial ventral flange, and extends far beyond the
tip of the antennal spine. The antennal spine is slender, acute, marginal and
situated closely below the inferior orbital angle. The hepatic spine is well
developed, larger than the antennal spine and situated more posteriorly at a
distinctly lower level. The antero-lateral angle of the carapace is bluntly rounded.

The third abdominal segment is strongly produced postero-dorsally as a
compressed rounded lobe projecting backwards, from the posterior half of the
segment. The fifth segment is about 0.4 of the length of the sixth, which is 2.3
times longer than wide. The postero-lateral angle is acutely produced and the
postero-ventral angle blunt and setose. In the male, the pleura are normal, all
broadly rounded posteriorly. In the non-ovigerous female they are similar but
larger. In the ovigerous female, the first three are greatly expanded, with the
first and third broadly notched and the second deeply notched antero-ventrally.
The pleuron of the first abdominal segment is strongly produced antero-ventrally
and the lower part separated by a deep notch from the upper part of the pleuron,
which is also produced anteriorly.

The telson is about 0.8 of the length of the sixth abdominal segment. It
is about 3.5 times longer than the anterior width, tapering with straight sides
to half that width posteriorly. The two pairs of dorsal spines are situated at
0.5 and 0.75 of the telson length and are equal to about 0.6 of the telson length.
The posterior margin is moderately produced, with a slender acute median point.

Aust. Zool. 19(2); 1977 219

A. J. BRUCE

19(2), 1977

Aust. Zool.

220

A REDESCRIPTION OF PERICLIMENES AESOPIUS

The lateral pair of posterior spines is similar to the dorsal spines. The intermediate
spines are long and robust, equal to 0.2 of the telson length. The submediam spines
are also robust, equal to 0.4 of the intermediate spines and finely setulose. A pair
of simple setae are also present dorsally.

The cornea is globular, of slightly greater diameter than the distal end of
the eyestalk, which is subcylindrical, about 2.3 times longer than wide at the
base. An accessory pigment spot is distinct. The ophthalmic somite bears an
acute median, anteriorly directed hook-like process.

The antennular peduncle slightly exceeds the tip of the rostrum but not the
spine of the scaphocerite. The proximal segment is about 2.7 times longer than
wide at half its length, with the antero-lateral lobe strongly produced with
numerous marginal setae, and medially inclined to reach the level of the proximal
end of the distal peduncular segment. The lateral margin is straight, with a strong
distal tooth at a level just distal to the proximal end of the intermediate segment
and far exceeded by the antero-lateral lobe. The ventro-medial margin bears a
short acute tooth. The stylocerite is well developed, slender and acute, reaching
to the level of the middle of the medial border. The statocyst is normal with a
regular oval statolith. The intermediate and distal segments are together equal to
half the length of the medial margin of the proximal segment, with the former
segment equal to two thirds of the length of the latter and with a small setose
antero-lateral lobe. The lower flagellum is biramous, with the rami fused for the
twelve proximal segments. The shorter free ramus, in the dissected female specimen,
consists of seven segments. The larger ramus is filiform and has about 20 segments.
About 15 groups of aesthetascs are present.

The basicerite of the antenna is robust, with a strong lateral tooth. The
carpocerite exceeds the tip of the stylocerite, and is about twice as long as broad
and compressed. The scaphocerite is almost three times longer than broad, with
the greatest width distally at the level of the disto-lateral spine. The anterior
margin of the lamella is broadly rounded, slightly medially orientated and
extending far beyond the disto-lateral spine, which is at the end of the straight
lateral margin. The flagellum is long and slender.

The mouthparts are typical of the genus Periclimenes. The mandible is
without a palp. The corpus is stout and the incisor process is robust, tapering
distally to three stout terminal teeth. The molar process bears several large
blunt teeth distally. The maxillula has a slender bilobed palp, the lower lobe
having a simple hook-like seta disto-ventrally. The upper lacinia is broad, with
7-8 finely denticulate spines and numerous setae distally. The lower lacinia is

OO ae

Figs. 2-14. Periclimenes aesopius. 2-4, rostral variation in ovigerous females. 5, inferior
orbital angle. 6, pleuron of first abdominal segment, left lateral. 7, antennule.
8, antenna. 9, first pereiopod. 10, second pereiopod. 11, dactyl or fifth pereiopod.
12, uropod. 13, telson. 14, posterior telson spines.

Aust. Zool. 19(2), 1977 221

A. J. BRUCE

87-97 ‘TT IT
| wwo-t ]

19(2), 1977

Aust. Zool.

222

A REDESCRIPTION OF PERICLIMENES AESOPIUS

slender and setose distally. The maxillula has a slender, non-setiferous palp. The
basal endite is slender and tapering, with slender blunt distal lobes. The upper
lobe is distinctly longer than the lower and both bear a small group of slender
simple terminal setae. The coxal endite is absent and the margin is broadly
rounded. The scaphognathite is well developed, long and slender, about 3.3 times
longer than broad at the level of the base of the palp. The anterior lobe is slender,
with the medial margin strongly concave. The first maxilliped has a small slender
palp, non-setiferous, falling far short of the anterior margins of the basal endite
or caridean lobe. The basal endite is large, broad, rounded with numerous setae
along the antero-medial margins. The coxal endite is distinct, but small and
rounded, with a few short, simple setae and a single long plumose seta. The
caridean lobe is well developed, elongated and narrow. The epipod is small and
feebly bilobed. The endopod of the second maxilliped is typical. The dactylar
segment bears numerous stout denticulate spines along its medial margin. Longer
spines are present on the disto-medial angle of the propod. The coxa is produced
as a rounded medial lobe, with a few simple setae, and a small sub-rectangular
epipod, without a podobranch, is present laterally. The third maxilliped is slender
and exceeds the carpocerite by the length of the terminal segment. The
antepenultimate segment consists of the ischio-merus and basis, which are
completely ankylosed, with the medial border of the latter slightly produced,
broadly rounded and setose. The segment is strongly bowed, about 5 times
longer than broad, with a small spine at the distal end of the lateral margin and
the medial border is sparsely setose. The penultimate segment is slightly shorter
than the antepenultimate, about 5 times longer than wide, with numerous groups
of finely denticulate setae. The terminal segment is about 0.9 of the length of
the penultimate, about five times longer than wide, tapering distally, with more
numerous groups of denticulate setae. The coxa is not produced medially, but a
well developed rounded epipod is present laterally with also a small multilamellar
arthrobranch. All three maxillipeds have the flagella of the exopods well developed,
with about six large plumose setae distally and several shorter ones.

The fourth thoracic sternite is without a median process.

The first pereiopod is moderately stout, exceeding the carpocerite by the
length of the fingers. The palm of the chela is about twice as long as wide,
slightly compressed and scarcely tapering. The fingers are subequal to the palm
length, compressed, with feebly hooked tips and entire cutting edges. The carpus
is 0.75 times the length of the chela, 3.5 times longer than the distal width,

Figs. 15-29. Periclimenes aesopius. 15, mandible. 16, maxillula. 17. maxilla. 18, first
maxilliped. 19, second maxilliped. 20, third maxilliped. Types 21,22, carapace
and rostrum. 23, tip of rostrum, 24, inferior orbital angle, lateral aspect.
25, inferior orbital angle, ventral aspect. 26, 27, third abdominal segment.
lateral aspect. 28, first three abdomina! segments, dorsal aspect. 29, chela of
second pereiopod. 21, 24, 25, 26, 28, 29, allotype. 22, 23, 27, holotype.

Aust. Zool. 19(2), 1977 223

A, J. BRUCE

twice as wide distally as proximally. The merus is slender, 7.5 times longer than
wide and slightly shorter than the chela. The ischium is short, 0.4 of the length
of the merus and feebly setose along the medial border. The basis is 1.1 times
the length of the ischium and is also sparsely setose medially. The coxa is more
robust, with a small setose ventro-medial process.

The second pereiopods are subequal and similar. The carpocerite is exceeded
by the carpus and chela. The palm of the chela is about 2.5 times longer than
wide, feebly compressed and slightly tapering distally. The fingers are slightly
bowed and a little shorter than the palm length. The tips are hooked and the
cutting edges straight and entire. The carpus is short and stout, 0.6 of the palm
length, moderately expanded distally and unarmed. The merus and ischium are
also unarmed, the merus subequal to the palm length and 5.5 times longer than
wide, the ischium more slender, subequal in length to the merus and 6.0 times
longer than wide.

The ambulatory pereiopods are slender, the third exceeding the basicerite by
dactyl, propod and carpus. The dactylus is slender and curved, about 4.0 times
longer than the width at the base. The unguis is slender, equal to 0.6 of the
length of the corpus, which bears a stout distal accessory spine. Sensory setae
are present disto-laterally. The propod is about 10 times longer than wide, with
5-6 ventral spines. The carpus is 0.6 of the propod length and the merus is
subequal; both are without spines. The fourth and fifth pereiopods are similar
but longer, the ratios of the segments being, in the dissected female specimen:

Po P4 P5

Propod 5S) 39 45
Carpus 23 23 25
Merus - BD 43 52

The pleopods are normal. The appendices internae of the ovigerous female
are very well developed. The appendix masculina bears numerous long spiniform
setae.

The uropods are normal. The protopodite is without a postero-lateral spine.
The exopod is broad, 2.6 times longer than wide, with the lateral margin straight,
setose, terminating in an acute tooth distally, with a strong mobile spine medially.
The endopod is about 3.4 times longer than broad and falls short of the end
of the exopod.

TYPES.

Bate’s specimens are preserved in the collections of the British Museum
(Natural History). His description and figure 5 are based on the female specimen,
with a rostral dentition of 9/2. This specimen is now in a fragmented state. The
carapace is detached from the cephalothorax, which is also separated from the
abdomen. The mouthparts, except the third right maxilliped, the eyes and the
antennae are detached, but all pereiopods are still attached to the cephalothorax.

224 Aust. Zool. 19(2), 1977

A REDESCRIPTION OF PERICLIMENES AESOPIUS

The male specimen, with a rostral dentition of 11/2, is almost intact, and lacks
only the right third pereiopod. The female specimen is now designated as the
holotype and the male as allotype.

COLORATION:

Bate has reported that preserved specimens were spotted with red on
rostrum, antennae and pereiopods. Hale (1927, 1928) has also reported a similar
coloration in his specimens and illustrated the colour pattern.

MEASUREMENTS-

Post-orbital carapace length, ¢é (allotype) 3-4mm; 2, (holotype), non-
ovigerous, 5.0mm; ovigerous, 4.6-5.6 mm. Length of ovum, 0.5 mm.

Periclimenes holthuisi Bruce (Fig. 7).
Urocaris longicaudata Pearson, 1905, Rep. Ceylon Pearl Oyster Fish., 4:78
Sb a Sag
Periclimenes (Periclimenes) aesopius— Holthius, 1952, Siboga Exped. Mon.,
39a": 34-37, figs. 5-6.
Periclimenes aesopius— Bruce, 1966, Crustaceana, 10(1):21, fig. 1b, 2ef.
Periclimenes holthuisi Bruce, 1969, Zool. Meded., Leiden, 43(26):258-259.
—Monod, 1969, Cahiers Pacifique, 13:216-220, figs. 69-73. —Bruce, 1972,
Crustaceana, 23(2):300-302; 1973, Crustaceana, 24(1):99; in press.

>

MATERIAL EXAMINED.

4 spms., Bowen, Queensland, Coll. E. M. Rainford. British Museum (Natural
History) registration no. 1923.8.22.8-10.

DESCRIPTION:

The specimens agree well with the previously available information (Holthius,
1953; Bruce, 1969).-The rostral dentition is 8-9/1-2. The dorsal lobe on the third
abdominal segment is distinctly smaller than in P. aesopius. The inferior orbital
angle is particularly long and acutely produced. All specimens are in a good state
of preservation.

HOST:
The collector noted that the specimens were obtained from an anemone.

REMARKS.

The type material, from Hong Kong, was also found in association with an
anemone. The species has also been found in association with the jellyfish
Cassiopea andromeda Forsskal in Zanzibar (Bruce, 1972) and with the coral
Fungia actiniformis (Quoy & Gaimard). It has also been recorded from
Peloris Island, Queensland (Bruce, in press).

Aust. Zool. 19(2), 1977 295

A. J. BRUCE

DISTRIBUTION:

Type locality, Hong Kong. Also recorded from Zanzibar; Maldive Islands;
Ceylon; Indonesia, New Guinea; South China Sea; Japan; New Caledonia and
Queensland, Australia.

Periclimenes indicus (Kemp) (Fig. 8)

Urocaris indica Kemp, 1915, Mem. Indian Mus., 5:275, fig. 26, pl. 13 fie. -9.
=—Borradaile, 1917: Trans. Linn. Soc. Lond: Zool (2)7 1-523:

Periclimenes (Periclimenes) indicus— Kemp, 1922, Rec. Indian Mus., 24:104
(key), 144, 145, 146, fig. 13; 1925, Rec. Indian Mus., 27:323. —Holthius,
1952, Siboga Exped. Mon., 39a?°:9, 32, 33, 39-40, fig. 8.

Periclimenes indicus— Kemp, 1922, Rec. Indian Mus., 42:115. —Fujino &
Wiyake, 1970, J..Fac. Agnc., Kyushu: Univ. 16(5) > 253,

Periclimenes indica— Panikkar & Aiyar, 1939, Proc. Indian Acad. Sci., 98:253.
Periclimenes (Periclimenes) aesopius— Johnson, 1961, Bull. Mus. Nat., Singapore,
BODO. 468 9 79 tape le

MATERIAL EXAMINED:

(1) 31 “spms., (8 ovis.’ 9°, 4 juv.) Siglap, Simeapore, 6 ins: below tees
25 February 1952, coll. D. S. Johnson, British Museum (Natural History )
registration no. 1954.11.5.11-15.

(ii) 32 (2 ovig.), 4 juv., Myora, North Stradbroke Is., Queensland,
Australia, 31 January 1968, coll. A.J.B., (no. 836).

(iii) 46,149 (13 ovig.), Myora, North Stradbroke Is., as above) (no. 837).

(iv) 2 ovig. ¢, Peel Island, Moreton Bay, Queensland, Australia, 24
September, 1968> coll? ACB. (noe: 963):

(v) 26, 2 ovig. 2, Chilka Lake, no date, Zool. Survey India. (no. 952).

DESCRIPTION:

This species has been well described by Kemp (1922) and the Australian
specimens agree closely with his description and with the specimens from the
type locality, Chilka Lake, Orissa, India.

The specimens réported by Johnson (1961) as common in the Exhalus beds
at Singapore, and referred then to P. aesopius, have been deposited in the
collections of the British Museum (Natural History). These have been re-examined
and found to belong to P. indicus (Kemp). Of six ovigerous females with
undamaged rostra the dentition was 9/2 (4); 9/3 (1) and 10/2 (1), and the
proximal part of the dorsal crest is distinctly elevated. The epigastric spine is
situated well behind the posterior orbital margin. The inferior orbital angle is
produced, but short and distally blunt, with a small ventral flange. The third
abdominal segment is only slightly produced in the postero-dorsal midline and
is not elevated to form a hump.

226 Aust. Zool. 19(2), 1977

A REDESCRIPTION OF PERICLIMENES AESOPIUS

Figs. 30-33. Periclimenes spp. 30, P. aesopius ophthalmic somite, dorso-lateral view.
31, P. holthuisi Bruce, ovigerous female, Bowen, carapace and rostrum.
32, P. indicus (Kemp), ovigerous female, Siglap, Singapore, carapace and
rostrum. 33, same, third abdominal segment.

COLORATION:

Generally transparent but mottled with red-brown; especially along ventral
aspect of body, and dorsally over posterior margins of third and fourth abdominal
segments, tip of scaphocariate, ventral eyestalk, coxae of pereiopods and tip of
caudal fan. Scattered small white dots over branchiostegite and pleura. Ovary and
ova bright green.

HOST:

Macrodactylus aspera Haddon & Shackleton (Actiniaria). All 18 specimens
(no. 837) were obtained from the single host anemone.

REMARKS.

This species has not been previously recorded from Australia and these
records represent a considerable extension of the known range of this species.
The association of this species with an actinarian host has also not been
previously reported and is of interest. It is thought probable that the other
Australian specimens, as well as those from Singapore, may have been also
associated with anemones but that on disturbance these hosts had withdrawn
into the soft substrate leaving the shrimps among the surrounding phanerogams
to be caught by the collector’s net.

Aust. Zool. 19(2), 1977 227

A. J. BRUCE

DISTRIBUTION-

Type locality, Chilka Lake, Orissa, India. Also reported from the Gulf of
Manaar; and Madras, India; the Nicobar Islands, and Sumbawa and Celebes,
Indonesia. Now also from Singapore and Queensland, Australia.

DISCUSSION

Periclimenes aesopius (Bate) is not known outside the restricted region of
South Australia and appears to be an endemic relict species confined to that
area. It belongs to a group of widely distributed species that are characterised
particularly by the posterior production of the dorsal margin of the third abdominal
segment and a produced inferior orbital angle with a ventral flange. Most of the
species have a biunguiculate dactylus on the ambulatory pereiopods but this is
lacking in some species. In the Indo-West Pacific region this group is represented
by P. aesopius, P. holthuisi, P. indicus and also P. tosaensis Kubo, which differs
from the first three species in the lack of an accessory spine on the dactyls of
the ambulatory pereiopods (Kubo, 1951). In the Eastern Atlantic and
Mediterranean Seas the group is represented by P. amethysteus (Risso) and
P. scriptus (Risso), while in the Western Atlantic and Caribbean region there
are P. anthophilus Holthuis & Eibl-Eibesfeldt, P. pedersoni Chace, P. yucatanicus

Ti —S eS ee ee SS eS ee SS

40 60° 80° 100 120° 140° 160 180° 160 140° 120

Fig. 34. Distribution of Periclimenes aesopius (Bate) >, and related Indo-West Pacific species:
P. holthuisi Bruce A, P. indicus Kemp @, P. tosaensis Kubo ®.

228 Aust. Zool. 19(2), 1977

A REDESCRIPTION OF PERICLIMENES AESOPIUS

Ives and P. magnus Holthuis, the latter species also differing from the others in

the lack of an accessory spine on the dactyls of the ambulatory pereiopods
(Holthuis, 1952).

In the Pontoniinae there are no established examples of free-living species
that have biunguiculate dactyls on the walking legs. Of the species mentioned
above P. holthuisi, P. indicus, P. anthophilus, P. pedersoni and P. yucatanicus
are all known associates of sea anemones, (Holthuis & Eibl-Eibesfeldt, 1964;
Limbaugh et al, 1961). In the Mediterranean Sea P. amethysteus and P. scriptus
are also associates of actiniarians (Svoboda, per. comm.) and other coelenterates.
The hosts of P. aesopius, P. tosaensis and P. magnus have not yet been identified
but it seems probable that they will also be associates of sea anemones, or other
coelenterates.

The four American species all have the maxilla with a simple distal endite.

This contrasts with all the other species, East Atlantic and Indo-West Pacific,
~ in which this endite is normally distinctly bifid, although in P. amethysteus it may
be simple or bifid (Holthuis, 1952), suggesting that the two groups have originated
by radiation from different ancestral species. Of the species known at present
P. indicus is the least morphologically specialized but P. scriptus and P. amythsteus
also show little indication of specialization. In P. tosaensis and P. magnus the
simple dactylus of the ambulatory pereiopod is due to the secondary loss of the
accessory tooth.

Periclimenes aesopius may be distinguished from all the related species by
the remarkable hump-like process developed dorsally on the third abdominal
segment, and by the presence of three or four post-rostral teeth on the dorsum
of the carapace, as well as the biunguiculate dactylus of the ambulatory pereiopods.
In the other species usually only a single post-rostral tooth is present but in
P. holthuisi and P. anthophilus there may be 1 or 2, and P. yucatanicus generally
has two. The abdominal hump is less well developed in the other species but is
distinct in P. pedersoni, P. yucatanicus and P. magnus, as well as P. holthuisz.
In P. scriptus, P. amethysteus and P. indicus, it is very feebly developed, but the
postero-dorsal margin of the third abdominal segment is produced backwards but
not elevated, as it is also in P. anthophilus. In P. tosaensis a feebly elevated
hump is present. The antero-ventral emargination of the pleura of the ovigerous
female is also apparently unique in the Pontoniinae. With respect to these features,
P. aesopius appears to be the most highly specialized species.

The function of the dorsal abdominal hump is obscure but this region is
often conspicuously marked with bright colours in related species, (Chace, 1958;
Limbaugh, et al 1961), often fish cleaners, and so it is probably to increase the
visibility of the displayed signals.

Hale (1927) has described and illustrated the appearance of P. aesopius in
life. The body of the shrimp is largely transparent but the dorsum of the first
three abdominal segments, including the hump of the third segment, is white
outlined in red. This must present a conspicuous signal. This region is similarly

Aust. Zool. 19(2), 1977 229

A. J. BRUCE

conspicuously coloured in the cleaner shrimp P. pedersoni. A large tan coloured
saddle shaped spot, ringed with white is present on the third abdominal segment
in P. yucatinicus, another cleaner shrimp. Conspicuous red and white patches are
also present on the third abdominal segment dorsally in Leandrites cyrtorhynchus
Fujino & Miyake. In this palaemonine shrimp the third abdominal segment is
posteriorly produced and it is also reported to be a fish cleaning species (Holzberg
1971). The markings in this shrimp are also red and white (Bruce, 1975).

REFERENCES

BATE, C.S. (1863). On some new Australian Species of Crustacea. Proc. zool. Soc. Lond..,
1863, 498-505, pls. 40-41.

BORRADAILE, L. A. (1898). A Revision of the Pontoniidae. Ann. Mag. nat. Hist., (7) 2,
376-391.

1g DG AE ae a a ee 2 (1917). On the Pontiniinae, The Percy Sladen Trust Expedition to
the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner. Trans. Linn.
Soc. Eond:, (2) 17, 323-396) als, 52-50:

BRUCE, A. J. (1966). Notes on some Indo-Pacific Pontoniinae, I. Periclimenes tosaensis
Kubo. Crustaceana, /0 (1), 15-22, figs. 1-4.

are y sea ey ENE RS (1967). Notes on some Indo-Pacific Pontoniinae, HI-IX. Descriptions
of some new genera and species from the western Indian Ocean and South China Sea.
Zool. Verhand., Leiden, 87, 1-73, figs. 1-29.

Fgh Pibee Lc Maia ee ro ee ee (1969). Preliminary descriptions of sixteen new species of the
genus Periclimenes Costa, 1844, (Crustacea, Decapoda Natantia, Pontoniinae). Zool.
Meded., Leiden, 43 (20), 253-278.

PT SE DAS OI ud aM Sa aes (1972). An association between a pontoniinid shrimp and a
rhizostomatous scyphozoan, Crustaceana, 23 (3), 300-302.

Spe th ata oe 2 Dis AR (1973). Notes on some Indo-Pacific Pontoniiae, XXII. Pliopontonia furtiva

gen. nov., sp. nov., a new shrimp associated with a corallimorph zoantharian.
Crustaceana, 24 (1), 97-109, figs. 1-5; pl. 4.

BASEN hake AY t Une (1975). Coral reef shrimps and their colour patterns. Endeavour,
39, 23-2). f198:, 1-1 6,

LAT Rp a oat lea: ke ON In press. A report on a small collection of pontoniinid shrimps
from Queensland, Australia. Crustaceana.

CHACE, F. A, Jr. (1958). A new shrimp of the genus Periclimenes from the West Indies.
Proc. biol. Soc. Washington, 77, 125-132, figs. 1-17.

FUJINO, T. & S. MIYAKE (1970). Caridean and Stenopodidean Shrimps from the East
China and the Yellow Seas (Crustacea, Decapoda, Natantia). J. Fac. Agric., Kyushu
Wniv..<165 (3); 239-312) shes, 1-25:

HALE, H. M. (1927). The Crustacea of South Australia, 7, 1-201, figs. 1-202.

So ae pa a eae coe aM Lee ee (1928). Some Australian Decapod Crustacea, Rec. S. Aust. Mus.,
4, 91-104, figs. 19-27.

HASWELL, W. A. (1882). Catalogue of Australian stalk- and _ sessile-eyed Crustacea,
i-xxiv, 1-324, pls. 1-4.

HOLTHUIS, L. B. (1951). The Subfamilies Euryrhynchinae and Pontoniinae. A general
Revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas. I.
Allen Hancock Found. Publ., Occ. Pap., 1/1, 1-332, pls. 1-63.

ee USES AN, AOL One a NaaE ra (1952). The Decapoda of the Siboga Expedition. Part XI. The
Palaemonidae collected by the Siboga and Snellius Expeditions, with remarks on other
species. II. Subfamily Pontoniinae. Siboga Exped, Mon., 39a’, 1-252, figs. 1-110, tab. 1.

230 Aust. Zool. 19(2), 1977

A REDESCRIPTION OF PERICLIMENES AESOPIUS

HOLTHUIS, L. B. & I. EIBL-EIBESFELDT (1964). A new species of the genus Periclimenes
from Bermuda (Crustacea, Decapoda, Palaemonidae). Senck. biol., 45, 185-192, figs. 1-4.

HOLZBERG, S. (1971. Beobachtungen einer Putzsymbiose zwischen der Garnele Leandrites
cyrtorhynchus und Riffbarschen. Helgolander wiss. Meeres unters., 22, 362-.

JOHNSON, D. S. (1961). A synopsis of the Decapod Caridea and Stenopodidea of
Singapore with notes on their distribution and a key to the genera of Caridea occurring
in Malayan waters. Bull. nat. Mus. Singapore, 30, 44-79, pl. 1.

KEMP, S. (1915). Crustacea Decapoda. Fauna of the Chilka Lake. Mem. Indian Mus.,
5, 199-325, figs. 1-38, pls. 12-13.

She tne = i eR (1922). Notes on Crustacea Decapoda in the Indian Museum. XV.
Pontoniinae, Rec. Indian Mus., 24, 113-288, figs. 1-105, pls. 3-9.

Pee net a es Eee (1925). Notes on Crustacea Decapoda in the Indian Museum.
XVII, On various Caridea. Rec. Indian Mus., 27, 249-343, figs. 1-24.

KUBO, I. (1951). Some macrurous decapod Crustacea found in Japanese waters, with
descriptions of four new species. J. Tokyo Univ. Fish., 38 (2), 259-289, figs. 1-16.

~ LIMBAUGH, C., H. PEDERSEN & F. A. CHACE Jr. Shrimps that clean fishes. Bull. mar.
Sci. Gulteand \Caribbean, 11, 237-257, figs. 1-9.-.

MONOD, Th. (1969). Sur quatre crevettes de Noumea (Nouvelle Caleconie). Cahiers
Pacific, 13, 191-222, figs. 1-73.

PANIKKAR, N, K. & R. G. ATYAR (1939). Observations on Breeding in Brackish-water
Animals of Madras, Proc. Indian. Acad. Sci., 9B, 343-364.

PEARSON, J. (1905). Report on the Macrura collected by Professor Herdman, at Ceylon
in 1902. In: Herdman, W.A., Report to the Government of Ceylon on the Pearl Oyster
Fisheries of the Gulf of Manaar, 4, 65-92, pls. 1-2.

Aust. Zool: 19(2), 1977 231

Sdn
Nbc BRD kk

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Diop aay

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Ne uy ee ‘nye ety ey

A System of Intersomital Dorsoventral Muscles
in the Posterior Body Somites of Land-leeches
(Hirudinea : Haemadipsoidea)

LAURENCE R. RICHARDSON
4 Bacon Street, Grafton, N.S.W. 2460

ABSERAGT

Six pairs of dorsoventral muscles have their origins in somites xxii to xxvii,
and insertions in xix to xxiii. The system is known only in Haemadipsoidea.

INTRODUCTION

Somatic dorsoventral muscles are known in Hirudinea as extending from
the ventral to the dorsal body wall with the origin and insertion of a strand
essentially in the same transverse plane.

There is described here a system of six pairs of strap-like dorsoventral
muscles having their origin in the vicinity of ganglia xxii to xxvii and the posterior
ganglionic mass, ascending to insertions on the dorsal body wall in somites xix to
xxiii, and accordingly intersomital. Dissections of some land-leeches also show
paired strap-like vertical muscles originating on the ventral body wall in the
middle of somites xxii and xxiii, and inserted onto the intestine. These and the
intersomital system are known to me only in land-leeches.

I have observed the intersomital system in land-leeches in the Haemadipsidae
s.s., and in all four subfamilies of the Domanibdellidae. The system is detailed
here as seen in Chtonobdella limbata Grube 1866, with brief comparative notes
on the system in three other species of the Chtonobdellinae.

The anatomy of the intersomital system indicates function in such actions as:
the maintenance of the extended body elevated at angles up to the vertical; torsion
on the longitudinal axis of the elevated extended body; the lateral swing through
an arc up to 160° of the body extended in the near horizontal plane; and the
positioning of the relatively large and heavy posterior sucker adjacent to the
anterior sucker during rapid dicotylal locomotion.

Excepting longitudinal undulation which I have not seen in land-leeches, these
and. other actions are performed with precision, slowly to rapidly, and this without
the advantage of relief from gravitational and other mechanical stresses such as
the surrounding medium provides for aquatic leeches.

It is proposed that the system be named the mm. Habenae (habena, -ae: the
thing holding a thing in place; the reins of a bridle; etc.).

Aust. Zool. 19(2), 1977 233

LAURENCE R. RICHARDSON

The mm. habenae are assessable as dorsoventral muscles modified along with
the reduction on the venter of the posterior body somites, a combination
representing an early specialization in land-leeches for terrestrial locomotion.

RESULTS AND DISCUSSION

GENERAL ANATOMY OF THE POSTERIOR BODY SOMITES AND DORSOVENTRAL
MUSCLES IN EUTHYLAEMATOUS LEECHES:

In aquatic euthylaematous sanguivorous and macrophagous leeches, somites
xxiii and xxiv are fully formed on all aspects of the body wall, as also xxv which
may be shortened on the venter by a reduction in the length or number of annuli.
Somite xxv is commonly the most posterior somite formed on the venter, in
some xxvi may be represented by a very short thin ridge. Somite xxvii is formed
only on the dorsum. In these leeches, somital ganglia xxiii to xxvii are spaced
apart, and xxvii narrowly spaced from the posterior ganglionic mass. The posterior
end of the body cavity tapers gradually to the narrow base for the posterior sucker.

In land-leeches, somite xxiii (xxiv in nearly all Domanibdellinae) is the
most posterior somite formed on the venter, and reduced in length on this aspect.
Somites xxiv (xxv) to xxvii are transversely abbreviated, progressively reduced
in length on the dorsum and margins, and not represented on the venter. The
reduction of the venter is reflected in the central nervous system in which somital
ganglia xxii and xxiii are well spaced apart; xxiii spaced narrowly from xxiv; and
xxiv to xxvii are in intimate contact, with xxvii in contact with the posterior
ganglionic mass (Richardson, 1975).

The posterior end of the body cavity in land-leeches has the form of a broad
shallow basin, the dorsal and dorsolateral walls formed by xxiv to xxvii, and the
wide floor by the base for the posterior sucker.

The dorsoventral muscles in euthylaematous leeches are arranged in four
longitudinal rows extending from the pharyngeal region into the intestinal region.
A pair of paramedian rows define a median splanchnic chamber between them.
The body cavity on each side of this is divided by a lateral row into a paramedian
and a lateral splanchnic chamber (Richardson, 1969).

The four rows are complete palisades in macrophagous leeches. In the
sanguivorous leeches, the paramedian rows are represented along their length by
intersomital clusters of strands, embracing and dividing the crop into compartments
which extend into the paramedian chambers.

In aquatic sanguivores, the paramedian rows are represented by intersomital
clusters in the intestinal region at xix/xx to xxv/xxvi.

In land-leeches, the paramedian rows have the typical sanguivorous form,
with clusters in the intestinal region at xix/xx to xxii/xxiii, some lacking clusters
at xxi/xxil.

234 Aust. Zool. 19(2), ‘977

INTERSOMITAL MUSCLES OF LEECHES

Dissection: An incision through the margin of the body wall at xviii/xix is
continued across the dorsum into the opposite margin. From this, an incision is
made along the midline of the dorsal median field to the anus. This displays the
paramedian dorsoventral muscles, and the intestine and the rectum in the median
splanchnic chamber. The intestine is transected at the junction to the rectum,
and these organs reflected.

Dissection on each side of the ventral nerve cord exposes the origins of the
mm. habenae, and the course and insertions of the individual muscles can be
followed. The somital levels of the origins can be assessed from the somital ganglia:
the insertions, from the annulation of the dorsal body wall.

ANATOMY OF THE MM. HABENAE IN Chtonobdella Limbata GRUBE 1866.
CEies- 1) to"S))

Bilaterally symmetrical, elongate, thin, strap-like muscles originating close to
the midline on the floor of the median splanchnic chamber in xxii and the
following preanal somites; ascending anteriorly, laterally to transversely wide
spaced insertions on the body wall roofing the paramedian and lateral splanchnic
chambers in somites xix to xxiii.

In terms of their origins, the mm. habenae are divisible into:
a) anterior habenae, two pairs originating in the adjacent halves of xxii and xxiii;

b) middle habenae, two pairs originating adjacent to ganglia xxiv to xxvii and
the posterior ganglionic mass; and

c) posterior habenae, two pairs originating posterior to the middle pair.

The anterior and middle habenae have their origins in the ventral body wall,
and the posterior pairs essentially on the base for the posterior sucker.

The two pairs at each level differ in their origins, courses and insertions.
The most suitable anatomical description is provided in the following groupings: —

GROUP A.

Three pairs of habenae, anterior (A!), middle (A*), and posterior (A*),
the origins close to the midventral line, entering and ascending in the paramedian
splanchnic chambers to insertions on the body wall roofing these chambers.

A’: origins in the contiguous halves of xxii and xxiii; ascending in the
paramedian chamber medial to the postcaeca, to insertions in xix.

A?: origins lateral to ganglia xxiv to xxvii and the posterior ganglionic mass;
course as Al; insertions on the anterior half of xxi.

A®: origins posterior to the posterior ganglionic mass; ascending in the
paramedian chambers posterior to the postcaeca; insertions on the anterior half of
Moxa

Aust. Zool. 19(2), 1977 235

LAURENCE R. RICHARDSON

Figs. 1-3. The mm, habenae in Chtonobdella limbata Grube 1866. 1, Dissection of somites
xix to xxvii from the dorsal aspect to show the origins, courses and insertions of the mm.
habenae: Group A, on the left; on the right, A*, the origins of A®* and A®*, and Group B.
Only two pairs of dorsoventral clusters of the paramedian row are shown, others and the
lateral palisades omitted for clarity. 2 and 3, semidiagrammatic representations of the dorsal
and lateral views showing the general relationships of the habenae in the entire extended
specimen.

Somital limits indicated by broken lines; somites and somital ganglia, by roman figures:
annuli, as, etc.; somital ganglia represented at relative size. Scales, mm.

Abbreviations: an., anus; int., intestine;l].mm., longitudinal muscular layer in the body
wall; l.o., lambertian organ; pc., postcaecum; p.g.m., posterior ganglionic mass; pm. dvm.,
paramedian dorsoventral muscle; re., rectum,

236 Aust. Zool. 19(2), 1977

INTERSOMITAL MUSCLES OF LEECHES

GROUP B.

Three pairs of habenae, anterior (B1), middle (B*), and posterior (B*), the
origins immediately lateral to the origins of the corresponding habenae of Group
A, entering and crossing the floor of the paramedian chamber, to enter and ascend

in the lateral splanchnic chambers to insertions on the body wall roofing these
chambers.

B*: origins in the contiguous halves of xxii and xxiii; crossing the paramedian
chamber ventral to the postcaeca, ascending lateral to the postcaeca, to insertions
on the contiguous halves of xx and xxi.

B*: origins lateral to ganglia xxiv to xxvii and the posterior ganglionic mass;
course, as B'; insertions on the posterior half of xxii.

B®: origins posterior to the posterior ganglionic mass; course posterior to the
postcaeca; insertions on the posterior half of xxiii.

The above provides a longitudinal alternation of the insertions of the habenae
of Groups A and B, with the individual habenae of the B group having their
insertions posterior to the corresponding habenae of the A group.

COMPARATIVE ANATOMY OF THE MM. HABENAE IN SOME CHTONOBDELLINAE:

The Chtonobdellinae contains the 5-annulate 2-jawed land-leeches of continental
Australia east of the Great Divide (Richardson, 1975). The species considered
here are C. limbata, central New South Wales, zoogeographically central eastern
Bassian; QOuwaesitobdella bilineata Richardson 1975, central to northern New
South Wales, central and northern eastern Bassian; Jaabdella whitmani (Lambert,
1899), southern and central Queensland, southern eastern Torresian; and
Amicibdella nigra Richardson 1975, northern Queensland, northern eastern
Torresian.

The annulation of the posterior body somites, the form of the posterior end
of the body cavity, and the general anatomy of the mm. habenae are similar
in the four species. One specimen of J. whitmani had a fourth pair of dorsoventral
muscles originating posterior to the posterior ganglionic mass and inserted in the
posterior half of xxiii. These were not seen in other specimens.

The summary given below shows the origins of A* and B* as the same in
all four species. Groupings based on the differences in the origins and/or insertions
of other pairs, show some peculiar to one species: e.g. the origin of A* more
posterior in J. whitmani; the insertion of A‘ and the origin of B’ more posterior
in A. nigra; the insertion of B® more posterior in C. limbata; etc.

A major difference separates the bassian and torresian species: the origins of
A® and B? are posterior to the posterior ganglionic mass in C. limbata, Q. bilineata;
lateral to ganglia xxiv to xxvii and the posterior ganglionic mass in J whitmant,
A. nigra.

A!: origins in the contiguous halves of xxii and xxiii in C. limbata, Q. bilineata,
A. nigra, the posterior half of xxiii in J. whitmani; insertions in the middle halt

Aust. Zool. 19(2), 1977 237

LAURENCE R. RICHARDSON

of xix in C. limbata, the posteror half of xix in Q. bilineata, J]. whitmani, the
anterior half of xx in A. nigra.

A®: origins lateral to ganglia xxiv to xxvii and the posterior ganglionic mass
in all four species; insertions in the anterior half of xxi in C. limbata, J. whitmani,
the posterior half of xxi in Q. bilineata, A. nigra.

~~

A®: origins posterior to the posterior ganglionic mass in C. limbata, Q.
bilineata, \ateral to ganglia xxiv to xxvii and the posterior ganglionic mass, common
with A? in J. whitmani, A. nigra; insertions, the anterior half of xxiii in C. limbata,
contiguous halves of xxii and xxiii in J]. whitmani, posterior half of xxii in
QO. bilineata, A. nigra.

B': origins in the contiguous halves of xxii and xxiii in C. limbata, Q. bilineata,
]. whitmani, in the posterior half of xxiii in A. migra; insertions in the contiguous
halves of xx and xxi in C. limbata, J]. whitmani, A. nigra, posterior in xx in
QO. bilineata.

B?: origins lateral to ganglia xxiv to xxvii and the posterior ganglionic mass
in all four species; insertions in the posterior half of xxii in C. limbata, the
contiguous halves of xxi and xxii in Q. bilineata, J. whitmani, the anterior half
of xxii in A. nigra.

B®: distinct separate origins posterior to the posterior ganglionic mass in
C. limbata, Q. bilineata, a common origin with B? in J. whitmani, A nigra;

insertions in the posterior half of xxiii in C. limbata, the contiguous halves of
xxii and xxiii in Q. Dilineata, J. whitmani, A. nigra.

ACKNOWLEDGEMENTS

This study has been conducted under an award from the Australian Research Grants
Committee for studies on the zoology of Australian freshwater and terrestrial leeches.

REFERENCES

RICHARDSON, L. R. (1969). A contribution to the systematics of the hirudinid feeches.
with description of new families, genera and species. Acta. Zool. Hung. J5 (1-2),
97-149.

RICHARDSON, L. R. (1975). A contribution to the general zoology of land-leeches
(Hirudinea : Haemadipsoidea Superfam, nov.) Acta Zool. Hung. 2/ (1-2), 119-152.

238 Aust. Zool. 19(2), 1977

Short Paper:

Artificial Hybridization Techniques For
Anuran Amphibians

G. F. WATSON
Department of Zoology, University of Melbourne, Parkville, Victoria, 3052

The general characteristics of most anurans (for example, their availability,
external fertilization, lack of opaque covering on the egg, and ease of laboratory
maintenance of embryos and larvae) make them particularly suitable for
experimentation involving in vitro hybridization tests. This approach has been used
both in studies of postmating isolation in closely-related species groups, and in
establishing phylogenetic relationships in larger taxonomic groupings. For example,
both applications have been used extensively in studies of the genus Bufo (see
review by Blair, 1972).

Females for crosses should be freshly-collected, naturally ovulated
individuals which are used within six hours of collection. Males can be stored for
up to three months in cool cabinets (5-8°C) without any detectable effect on
production of sperm. All animals should be killed by pithing immediately prior
to use.

The following instruments have been found useful in performing artificial
crosses. The Lawton Instrument Company catalogue number is given in parentheses
to aid in instrument selection.

1 pair straight iris scissors (05-1450)

2 pairs small thumb dressing forceps, serrated (23-0560 )

1 pair large angular forceps, serrated (09-0196)

All instruments should be clean and kept free of chemical contamination.

In addition, the following materials should be prepared before beginning the
experiment:

- 500-1000 ml of distilled water (cold)

500-1000 ml of distilled water (boiling )

For each male in the experiment, a new sterilized, disposable plastic petri
dish (e.g. 9.0 cm petri dish from Sterimed, Melbourne) should be marked on the
lid and base with a permanent code to identify each cross, filled with about 10 ml
of pond water [half strength Holtfreter’s solution (Rugh, 1962) can be used
instead of pond water] and placed so that the base is tilted and supported on one

side by the lid.

Aust. Zool. 19(2), 1977 239

G. F. WATSON

The following techniques are modifications of the basic methods outlined
in Rugh (1962) and have proved highly successful in crosses within all Australian
genera of anurans which have been studied to date, viz. Crinia, Geocrinia,
Limnodynastes, Litoria, Neobatrachus, Pseudophryne, and Ranidella (Watson, 1974,
unpub. obs.).

A sperm suspension is prepared by dissecting testes from a male and macerating
them (angular forceps facilitate this step) in water in a petri dish. Not more than
50 eges should be used in each cross, so depending on the number of eggs available
several ‘males can be-crossed to one female (for example, four to five males can
be crossed to females of Ranidella, while 12 or more may be used with females
of Limnodynastes ).

After allowing 5 to 10 minutes for the sperm to become active, the female is
killed and oviducal eggs dissected out and distributed among the sperm suspensions
in such a way that each suspension receives eggs from different regions of both
oviducts. After adding eggs to the sperm suspension the petri dish should be
shaken gently to spread the eggs into a monolayer.

During all the above procedures it is important to prevent cross contamination
of sperm suspensions. This is achieved by placing the dissecting instruments into
boiling water and then into cold water between each operation.

For all artificial hybridization experiments it is essential to perform a partial
control for each experimental series by fertilizing some of the eggs from the female
with conspecific sperm.

Techniques for maintenance of embryos will vary according to the life history
characteristics of the species involved. :

In species where embryonic development is wholly aquatic, the petri dishes
are filled with pond water after the eggs have rotated (rotation, resulting in the
animal pole of each egg being uppermost, indicates successful fertilization).
Organic debris, and damaged and infertile eggs should be removed. The egg
clumps, which normally adhere to the bottom and sides of the petri dishes should
be freed so as to float in the water, thus maximizing the exposed surface area of
the clumps to ensure adequate gaseous exchange for the developing embryos. The
dishes should then be covered.

In groups where embryonic development normally occurs on land in a
saturated atmosphere (e.g. Geocrinia, Pseudophryne) the following procedure
is followed. After rotation of the eggs and hydration of the egg capsules, excess
water, organic debris, and damaged and infertile eggs are removed. The developing
eggs should be arranged in a monolayer such that the eggs are in contact in order
to provide an adequate exposed surface for gaseous exchange, while reducing
to a minimum the surface area through which water loss from the egg capsules
can take place. A wad of water-soaked paper tissue is placed in each dish out
of contact with the egg capsules, and the dishes are covered. The egg capsules
should be maintained in a fully hydrated state prior to hatching of the embryos

240 Aust. Zool. 19(2), %977

HYBRIDIZATION TECHNIQUES FOR AMPHIBIANS

by daily addition of drops of distilled water directly on to the eggs and by keeping
the tissue wad saturated.

All developing embryos should be examined regularly for the presence of
developmental abnormalities (see examples in Rugh, 1962). All abnormal embryos
should be removed from the cultures.

After hatching, the young larvae should be transferred immediately to
larger aquaria. Clean, plastic, glass or enamelled metal dishes containing 2 to 3
litres of pond water or Holtfreter’s solution (Rugh, 1962) are suitable for this
purpose.

Conditions for larval culture vary with species. Most species seem to grow
well when fed par-boiled lettuce (clean lettuce leaves boiled for 5 to 6 minutes;
a large quantity can be prepared and deep frozen for future use). However,
some taxa, in particular members of the Myobatrachinae, require an additional
high protein supplement for successful larval development. This can be provided
by using high protein breakfast cereal, or by short duration addition of liver, or
by making a suspension of fine-ground, dried pet food (e.g. Dog-Chow) in agar
gel which when set can be cut up into small cubes and added to the aquarium.
Particular care must be taken, especially when using high protein food sources,
to keep the aquarium water clean and free of large-scale bacterial contamination.

When larvae reach stage 42 (Limbaugh and Volpe, 1957; i.e. when the
fore-limbs are free) they should be removed from the culture dishes and placed
in containers with tight-fitting lids. A small amount of pond water is added
to each container and the containers should be kept in a tilted position so that
the froglets can climb out of the water when metamorphosis is complete.

For most experimental purposes, raising hybrids to metamorphosis is
sufficient. If adult hybrid progeny are required, the froglets should be transferred
to terraria and feed live food appropriate to the size of each individual.

REFERENCES

BLAIR, W. F. (1972). Evolution in the Genus Bufo. University of Texas Press, Austin
and London.

LIMBAUGH, B. A. & E, P. VOLPE (1957). Early development of the Gulf Coast Toad,
Bufo valliceps Wiegmann, Amer. Mus. Novit. 1642: 1-32.

RUGH, R. (1962). Experimental Embryology. 3rd Ed. Burgess, Minneapolis.

WATSON, G. F. (1974). The evolutionary significance of postmating isolation in anuran
amphibians. Unpublished Ph.D. thesis, University of Melbourne.

Aust. Zool. 19(2), 1977 244

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NOTICE

The Royal Zoological Society of N.S.W. is sponsoring a symposium
on Monotreme Biology, to be held at the University of N.S.W. on
15th-16th May, 1978.

Prof. G. B. Sharman (Macquarie University) is the symposium
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Papers are invited dealing with unpublished material concerning
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All persons wishing to attend the symposium, whether speaking
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Authors presenting papers to the symposium may have them
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(price $5.00) in advance.

Aust. Zool. 19(2), 1977 243

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j ast

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COUNCIL, 1976

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THE AUSTRALIAN ZOOLOGIST

VOLUME 19 1977 PART II

CONTENTS

Maintenance of the platypus (Ornithorhynchus anatinus) in captivity under
laboratory conditions. T. R. GRANT, R. WILLIAMS and F. N.

The need for joining Illawarra wilderness areas. N. M. ROBINSON
A new species of Python from Armhem Land. G. PF. GOW ..............n0eae

Tail regeneration and other observations in a species of agamid lizard. C. J.
HARDY and C) M. HARON 6) yen eA le

Electrophoretic Evidence for the Specific Status of the Lizards Lampropholis

guichenoti (Dumeril and Bibron, 1839) and L. delicata (de Vis, 1887)
Bo B, HARRIS. and PP. G. JOHNSION 22... eae

Sex change in the Wrasse Pseudolabris gymnogenis (Labridae). G. R.
McPHERSGN i ee a ee

Voluntary submergence time and breathing rhythm in the homalopsine snake,
Cerberus rhynchops A HEATWORE fo.i60 4 dscns oe

The labrid fish genus Pseudojuloides, with a description of a new species.
A, MU AYEING and BG RUSSELL \.. 2.0.4 al. rr

A juvenile gempylid fish, Nealotus tripes, from eastern Australia. I.
NAKAMURA, and Jo ik, PAXTON RS) 20.0.0.0 25x... re aa

A new genus and two new species in the family Mithrodiidae (Echinoder-

mata: Asteroidea) with comments on the status of the species of
Mithrodia Gray 1840. ©. CC. POPE. and FF. W. E. ROWE :.. ae

A redescription of Periclimenes aesopius (Bate, 1863) (Crustacea:
Decapoda) with remarks on related species. A. J. BRUCE... es

A system of intersomital dorsoventral muscles in the posterior body somites

of land-leeches (Hirudinea: Haemadipsoidea). L. R. RICHARDSON

Short paper—.
Artificial a fr techniques for anuran amphibians. G. F. WATSON

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THE
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ZOOLOGIST |

Volume 19, Part 3
August, 1978

Scientific Journal of

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THE AUSTRALIAN ZOOLOGIST
VOLUME 19 1978 PART III

A Survey of
Central Australian Ichthyology

C. J. M. GLOVER & T. C. SIM
South Australian Museum, Adelaide, South Australia, 5000

ABSTRACT .

The history of Central Australian ichthyology is reviewed from its inception in
the 1840’s to the present. Hampered by geographical isolation, an initial period
of slow development preceeded the start of concerted scientific exploration
towards the close of the nineteenth century. Occurrences and distributions of
the Centre’s fishes are now well documented and a provisional check-list is herein
given. Taxonomic studies are well underway and increasing interest is now evident
in the ecological field.

INTRODUCTION

For the present purpose “Central Australia” is regarded as virtually all of
arid Australia, thereby embracing the drainage divisions of the Western Plateau and
Lake Eyre as defined by Map 5 of the Review of Australia’s Water Resources 1975
(1976), but excluding the Bulloo-Internal and Murray-Darling divisions (see
Pigure [);

Except for one isolated occurrence (see later, Basedow 1918), no fishes were,
until very recently, recorded from the mainly ephemeral waters of the Western
Plateau. Most of the following commentary will therefore be almost entirely
restricted to the Lake Eyre division.

Good general accounts of the country within the Central Australian region
are given in Scenic Wonders of Australia (1976), whilst the Review of Australia’s
Water Resources 1975 (1976), provides concise descriptions and summarised data
of the drainage divisions themselves.

The Lake Eyre drainage division embraces an arid area of 1.17 million km’.
Major rivers periodically flow in the relatively high-rainfall country of the north-
east sector, in times of heavy seasonal precipitation, towards the interior and Lake
Eyre itself. Flows reach the latter, and other major salt pans, irregularly, following

Aust. Zool. 19(3), 1978 245

C. J. M. GLOVER and T. C. SIM

TABLE ONE

A PROVISIONAL CHECK-LIST OF CENTRAL AUSTRALIAN FISHES

All species listed, except Carassius auratus, have been recorded from the Lake Eyre
Drainage. Those marked with a dagger (7) have also been recorded from the Western
Plateau Drainage. Those with an asterisk (*) appear to be endemic to the Central Australian
region.

Class TELEOSTOMI
Sub-Class ACTINOPTERYGII

Family Clupeidae
+Nematalosa erebi (Ginther, 1868)

Family Retropinnidae
Retropinna semoni (Weber, 1895)

Family Plotosidae
Neosilurus hyrtlii (Steindachner, 1867)
+* Neosilurus argenteus (Zietz, 1896)
* Neosilurus sp, nov. (Ms., Feinberg & Nelson)
Neosilurus spp. noy. (3?)

Family Poeciliidae
tGambusia affinis (Baird & Girard, 1853)
Gambusia dominicensis Regan, 1913

Family Melanotaeniidae
{Melanotaenia tatei (Zietz, 1896)

Family Atherinidae
Craterocephalus stercusmuscarum (Ginther, 1867) (= C. fluviatilis McCulloch, 1913)
Craterocephalus eyresii (Steindachner, 1884)
*Craterocephalus dalhousiensis Ivantsoff & Glover, 1974
Craterocephalus sp. (nov.?)

Family Centropomidae

+Ambassis castelnaui (Macleay, 1881)
Denariusa bandata (Whitley, 1948)

Family Serranidae
Plectroplites ambiguus (Richardson, 1845)

Family Teraponidae

|Leiopotherapon unicolor (Gunther, 1859)
Amniataba percoides (Gunther, 1864)
* Hephaestus sp. (nov.?)
*Bidyanus welchi (McCulloch & Waite, 1917)
Scortum hilli (Castelnau, 1877)
*Scortum barcoo (McCulloch & Waite, 1917)

Family Gobtidae

Glossogobius giurus (Hamilton-Buchanan, 1822)
Mogurnda mogurnda (Richardson, 1844)
*Chlamydogobius eremius (Zietz, 1896)

Hypseleotris spp. nov. (2?)

Family Cyprinidae
7+Carassius auratus (Linnaeus, 1758)

246 Aust. Zool. 19(3), 1978

CENTRAL AUSTRALIAN ICHTHYOLOGY

%

Lake Eyre Division 2%

0, *00 6 eoeeg8

Fig. 1. The drainage divisions of Central Australia. Partly after Map 5 in Review of
Australia’s Water Resources 1975 (1976).

periods of exceptional rainfall and flooding in the north-east. Excluding the latter
region, most of the freshwater of the Lake Eyre system is ephemeral, consisting
mainly of scattered waterholes and rivers and creeks, which only fill with seasonal
rains. Apart from the present phenomenal and prolonged flooding of Lake Eyre,
permanent aquatic habitats in the arid interior are only represented by some large
permanent waterholes, man-made dams and reservoirs, and surface waters associated
with scattered artesian springs and bores.

The Western Plateau division, an immense area of 2.5 million km? of unco-
ordinated drainage, is, on the other hand, a region of very few permanent aquatic
habitats. Creeks and rivers only flow for brief periods, following sporadic heavy
rains, before the surface water is rapidly lost by high evaporation and absorption
into the porous ground. The absence of fish throughout most of this division is
therefore understandable.

Despite paucity of permanent water, a variety of fish, about twenty-nine
species to date (some undescribed), are known to inhabit the Lake Eyre division,
six of which (together with Carassius auratus) are also known from the Western
Plateau division. These thirty species therefore constitute the total known fish

Aust. Zool. 19(3), 1978 247

C. J. M. GLOVER and T. C. SIM

fauna of Central. Australia (see Table 1). Often found in small populations, espe-
cially in ephemeral waterways, many of these fishes in the Centre also maintain
themselves in great abundance in some of the more permanent waters, including
those of artesian origin. ,

In view of the many taxonomic problems remaining, either under current
investigation or awaiting attention, it must be emphasised that the list given in
Table 1 is provisional and subject to revision. For example, although Nelson and
Rothman (1973) tentatively referred the clupiid of the Lake Eyre drainage to
Nematalosa erebi, recent examination of larger collections than were -available to
Nelson and Rothman leave this designation open to doubt (C.J.M.G., unpublished
data). Furthermore, although Vari’s (1978) recent world revision of the Tera-
ponidae advocated a number of changes to the Centre’s teraponids, the region’s
plotosids remain very poorly known.

The following resumé traces the more noteworthy discoveries and ee
tions pertaining to Central Australian fishes. Many of the early observations were
clearly incidental to the then main task of geographical exploration. It was not
until the last decade of the nineteenth century that the impetus of scientifically
oriented exploration was evident. Whitley (1964) strangely omitted any mention
(other than a scant and indirect reference) of this aspect of Australian ichthyology
in his otherwise full and comprehensive survey.

Early references to specific, but otherwise formally undescribed fishes, are
accompanied, where possible, by suggested probable current species names. Limited
though these early reports may be, they do enable some comparisons to be made
with existing occurrences of fish in Central Australia: changes accompanying Euro-
pean settlement have been thus deduced (see Glover & Sim, 1978).

Detailed route maps of major expeditions are, in some instances, appended to
relevant reports and journals, and the routes of some expeditions are conveniently
mapped together in certain publications such as those of Landsborough (1862),
Calvert (1895) and Robinson (1927).

RESULTS

EarLy DISCOVERIES, 1840-1900

Documentation of the fish fauna of Central Australia commenced with the
earliest European exploration of the region. Eyre (1845) certainly recognised that
fish were a major food item of the Central aboriginal peoples, whilst Sturt (1849)
observed ‘‘immense numbers of fish in reservoirs’ adjacent to the desert later to
bear his name and queried how they gained access to such an isolated place. Sturt
(1849) appears to have provided the first description of a Central Australian fish,
which seemed “‘to be identical with the silver perch of the Murray . . . but deeper
and thinner” (Scortum barcoo or Bidyanus welchi? ).

Babbage e¢ al (1858) noted small fish at Gregory Creek and Finniss Springs
west of Marree, whilst Stuart (1861, 1865) frequently sighted fish during his
travels, including his epic 1861-62 journey across the continent. Stuart’s (1865)

248 Aust. Zool. 19(3), 1978

CENTRAL AUSTRALIAN ICHTHYOLOGY

account of stranded dead fish “resembling bream” (Nematolosa erebi?), at Lake
Eyre and elsewhere, appears to be the first record of a phenomenon later noted
in The Report of the Lake Eyre Committee (1955) following the Lake’s flooding
of 1949-50, and in more recent years by Ruello (1976) and Dulhunty and Merrick
(1976a, 1976b). Waterhouse (1863), who accompanied Stuart’s 1861-62 expedi-
tion, reported small fish “resembling gudgeon” (Chlamydogobius eremius?) at
Strangways Springs. Although Waterhouse made collections of these and other
natural history material (evidently the first to do so in the Central Australian
region), many of the more delicate specimens, including the fish, did not survive
the return journey. Moreover, as pointed out by Hale (1956), unpublished por-
tions of Stuart’s diary suggest that Stuart himself was little interested in Water-
house’s work, being as he was, preoccupied with crossing the Continent, and thus
adverse to allowing adequate time for collecting.

Burke and Wills et al (1861) frequently referred to fish obtained from
aborigines or that they themselves caught during their ill fated expedition. These
included a variety “‘nine to ten inches (229-254 mm) long and two to three inches
(51-76 mm) deep” (a Teraponid?); another termed by the aborigines “cupi’”—
“five to six inches (127-152 mm) long and not broader than an eel” (Neosilurus
sp.?); one termed a “peru” (see later, Gason 1879 re “paroo”, and Horne and
Aiston 1924)—a “common one with large coarse scales’? (Nematalosa erebi? );
“calwichi’”—“‘a delicious fish up to two pound” (0.9kg); and “silver perch”
(Bidyanus welchi? ).

McKinlay (1862) observed fish in Lake Blanche and elsewhere, Lands-
borough (1862) speculated on the origins of fish in waterholes on the southern
side of the Barkly Tablelands, and Giles (1894-95) reported ‘‘a number of small
fish of a bright brown color’ at Dalhousie Springs (Craterocephalus dal-
housiensis?). Giles (1875a, 1875b, 1889) sighted great concentrations of
fish in waterholes south-west of Alice Springs, including “a species up
to three pounds” (1.3kg) weight that “had a great resemblance to Mur-
ray Cod”, and many small fish in the Finke river. Gosse (1874), on
the other hand, although he inspected numerous permanent and ephemeral waters
in the Centre, made only one direct reference to “a hole of water with fish in it”
south-west of Alice Springs. Gason (1879), writing of the Dieri tribe of the lower
Cooper, states, in relation to food, that ‘fish and other freshwater habitants . . .
are few and unimportant, being caught in the waterholes and lakelets, which can
only be called creeks or rivers when the floods come down; the last of which
occurred in 1864”. Gason (1879) listed three fish species—‘“‘paroo”’, “‘a small
bony flat fish” (Nematalosa erebi?) (see Burke & Wills e¢ al 1861 above re “peru”
and Horne & Aiston 1924); “multhoomulthoo”—“a fish weighing from 3 to
3% Ibs” (1.4-1.6 kg); and ‘“‘moodlakoopa’’—“‘a fish averaging 2 lbs” (0.9 kg).

The Lake Eyre Expedition; under the leadership of Lewis (1875), collected
what was to constitute the type material of Craterocephalus eyresii, the first fish
from the Centre to be formally described (Steindachner, 1884).

Aust. Zool. 19(3), 1978 249

C. J. M. GLOVER and T. C. SIM

It is interesting to note here, in the South Australian Museum Annual Report
of 1886, that “we have only three species (of fish) from Lake Eyre...... any
small fish from creeks or waterholes in the interior, would, even if not new, at least
add to the knowledge of the distribution of species”. As indicated by South
Australian Museum Annual Reports, and that Museum’s fish register, acquisitions
of small collections of fish from the Centre were received intermittently from
individual donors in subsequent years. Lucas (1894) reported two fish from near
the MacDonnell Ranges, namely Chatoessus erebi (= Nematolosa erebi) and
Therapon fasciata (= Amniataba percoides).

To this point in time very little effective collecting had in fact been accom-
plished, the number of recognisable fishes reported from the Centre amounting to
no more than six species, only three of which could be considered as adequately
described forms. These six species represented the families Clupeidae (1),
Teraponidae (2), Atherinidae (1), Plectroplitidae (1) and Gobiidae (1). The
absence of Central Australian localities from Macleay’s (1881) Catalogue of
Australian Fishes, and other checklists of the period, e.g. Tennison-Woods (1883)
and Ogilby (1886), is indicative of the then sparse knowledge of the Centre’s fish
fauna. Indeed, Lucas (1894) prophetically wrote ... “It is to be hoped that the
(then underway) Horn Expedition will bring back abundant material by means
of which further light may be thrown on the distribution of Australian fresh-water

Hist e

The Horn Scientific Expedition to Central Australia (1896) in fact heralded
a major break-throuigh in the region’s ichthyology. From a substantial collection
of fish, eight species representing five genera were recognised, six of which com-
prised new records for the Centre and all (at that stage) evidently new forms
(Zietz 1896a, 1896b). The number of known fishes in the Centre now amounted
to nine, but three of the above ‘‘new”’ species later proved to be synonymous with
species described beforehand from elsewhere. In lieu of an apparent absence of
affinity with Murray River forms, likely northern origins were now suggested for
these fish in the above Horn Expedition’s report. Furthermore, a lack of evidence
to support aestivation, the probable role of floodwaters in dispersal and a suggestion
of aerial transport of fish eggs by attachment to the feet of birds were also noted
and commented upon in the expedition’s report. In addition to the general narrative
in Part I of this report, and Zietz’s formal descriptions in Part II, Spencer & Gillen
(1912) added further data and comment to the discoveries of fish by this expedi-
tion. Thus, at one stroke, the Centre’s known described fish fauna had been
increased from three to nine species, and basic but significant observations made
relating to their ecology.

DISCOVERIES SINCE 1900
A scurry of scientific exploration, including biological collecting, now followed.

First, Basedow’s (1914) journal of the 1903 Government North-West (of
South Australia) Expedition reported what, in the context of the time, were

250 Aust. Zool. 19(3), 1978

CENTRAL AUSTRALIAN ICHTHYOLOGY

particularly interesting observations, i.e. that there were “countless small fish in
pools, along the overflow . . . of the highly mineralised Oodnadatta bore water”
and that “the temperature of the water in which they were swimming, without
the least sign of discomfort, was as high as 45°C”. Basedow reported that this
phenomenon, “induced the local folks to suppose the fish came up with the water
from below’. Having “disproved” that theory “by tying a net over the outlet of
the bore for the greater part of the day without catching a fish”, Basedow com-
mented that “it is more probable that the fish came there as eggs, through the
agency of the water fowl that occasionally frequent the water”. Basedow further
reported that he had also seen fish in hot springs at the Douglas River in the
Northern Territory.

Next White’s expedition (White et al, 1914) noted fish at a number of bore
sites, and collected Leiopotherapon unicolor elsewhere, in the Centre.

A South Australian Museum expedition to Strzelecki and Cooper Creeks in
1916 resulted in a further two species being described: Bidyanus welchi and Scortum
barcoo (McCulloch & Waite, 1917).

Basedow (1918) next reported Leiopotherapon unicolor as having been collect-
ed in the Patterson Ranges of North-Western Australia. Interestingly, this last
record was, until recently, the one and only fish record from the Western Plateau
drainage division, despite earlier and subsequent explorations of the region by War-
burton (1875), the Elder Expedition party (Lindsay, 1893; Stirling & Zietz,
1893; Helms, 1896), Slater and Lindgren (1955), the Western Australian Museum
(Western Australian Museum Annual Reports 1963-64, 1964-65, 1969-70), the
Australian Museum (Australian Museum Annual Reports 1951-52, 1952-53) and
the Western Australian Wildlife Research Centre (Burbidge et al, 1976). Allen
(1977, in press) certainly did not record any species from the Western Plateau in
his checklist of Western Australian freshwater fish. However, in October 1977,
a South Australian Museum Expedition collected a number of species in the north
east sector of the Western Plateau drainage (Wiso and Barkly basins) i.e. Nemma-
talosa erebi, Neosilurus argenteus, Melanotaenia tatei, Ambassis castelnaui and
Leiopotherapon unicolor. In 1978 Carassius auratus and Gambusia affinis were
reported and confirmed present (personal communication from P. Burdon) in
a dam at Woomera in the south-east sector of the Western Plateau drainage
(Gairdner basin): this solitary occurrence of Carassius auratus is almost certainly
the result of human introduction (both wild and domestic colour varieties being
present), whilst the poeciliids (both here and in the Lake Eyre drainage)
have probably also been similarly introduced (see Glover & Sim, 1978).
Furthermore there are reports (personal communication from R. J. McKay)
of fish having been found by railway fettlers on the Nullabor Plain
during the period 1917-22, and of “blind fish” in cave pools on the Nullabor Plain
close to the railway line. In 1968 the South Australian Museum received a report
of “blind fish” having been sighted in a cave pool north of the main road between
Nullabor Homestead and Yalata Mission Station.

Aust. Zool. 19(3), 1978 251

C. J. M. GLOVER and T. C. SIM

By now McCulloch (1929-30) listed but nine species as being specifically
from ‘“‘Central Australia’.

Fletcher (1937) reported Leiopotherapon unicolor in Waroona Creek near
Camooweal (Queensland), whilst the Simpson Desert Expedition of 1939 obtained
Ambassis castelnaui, a new record, together with Leiopotherapon unicolor and
Bidyanus welchi, all from temporary waters (Whitley, 1945).

The early 1950’s saw a series of South Australian Museum trips to the then
flooded Lakes Eyre and Callabonna, resulting in a variety of fish being collected
from both (South Australian Museum Annual Reports 1952-53, 1954-55). In the
late 1960’s the Arid Zone Research Institute (Alice Springs) collected extensively
in the Northern Territory, thereby establishing new locality records for a number
of fishes in the north-western sector of the Lake Eyre drainage division (personal
communication from S.A. Parker).

Following on from an embryological study of Retropinna semoni by Milward
(1966), there then appeared a spate of breeding and physiologically oriented
works. Although not based directly on Central Australian stock, many of these
studies were upon species or closely allied forms known from that area, and thus
are of direct relevance. Lake (1967a, 1967b) investigated the breeding and dev-
elopment of Plectroplites ambiguus, Bidyanus bidyanus and Carassiops klunzingeri,
whilst Llewellyn (1968, 1971, 1973) examined aspects of the ecology, including
thermal tolerance and breeding, of Lezoptherapon unicolor and other species. McKay
(1973) later examined aspects of the reproductive cycle of Plectroplites ambiguus,
whilst more recently Beumer (1976) has further examined thermal (and salinity )
tolerance, and aestivation, in Lezopotherapon unicolor.

In 1967 a South Australian Museum investigation of the taxonomy and
ecology of Chlamydogobius eremius, a fish known to inhabit artesian springs and
bores, was initiated by one of the present authors (C.J.M.G.), and reported by
Glover and Inglis (1971). This study revealed interesting ecological adaptations
to an unusual habitat, in addition to many new locality records in the far north
of South Australia, and provided further insight into fish dispersal mechanisms in
the Centre (Glover 1971, 1973).

By this time Lake (1971) had listed seventeen species from nine families
from the Lake Eyre division, but none from the Western Plateau.

A party from the Australian Museum collected a series of fishes from Cooper
Creek, near Innamincka, including an evidently undescribed neosilurid, in 1971
(personal communication from J. R. Paxton). A joint party from the Australian
Museum and New South Wales State Fisheries, in 1975, collected in the same
place and from Lake Callabonna and other localities in the vicinity (personal
communication from D. F. Hoese).

Following a visit to Dalhousie Springs, as reported by Souter (1974a, 1974b),
Ivantsoff and Glover (1974) described a new atherinid from that locality and
reported ecological observations made there on previous occasions.

252 Aust. Zool. 19(3), 1978

CENTRAL AUSTRALIAN ICHTHYOLOGY

Stemming from earlier South Australian Museum studies, the present authors,
in 1975, embarked upon a field survey of waters throughout the entire Lake
Eyre division. Initial results of this project, augmented with complementary tax-
onomic/ecological studies, were reported by Glover (1975-78, 1977, in press)
and Glover & Sim (1978).

CONCLUSION

It is evident that Central Australian ichthyology has experienced a checkered
career, but that the last twenty-five years have seen an increasing impetus in
fundamental field surveys and collecting, and more recently in taxonomic and
ecological studies.

Suffice to say at this stage that adequate field observations and collections
probably now exist to provide a nearly complete picture of the overall occurrence
and incidence of the Centre’s fishes, though not all forms are yet described or their
relationships yet clear. These collections should engage systematists for some time
yet in unravelling the taxonomic complexities of the respective groups, particularly
of such lesser known ones like the notoriously obscure neosilurids. Other workers
known to be at present examining Central Australian fishes include D. F. Hoese
(Australian Museum : Gobiidae), W. Ivantsoff (Macquarie University : Atherini-
dae), J. R. Merrick (Sydney University : Teraponidae) and M. N. Feinberg & G. J.
Nelson (American Museum of Natural History : Plotosidae).

Although little is known of the general biology of most of the Centre’s fishes,
it is this avenue of research, upon specific species, that probably offers especially
promising and interesting results, particularly relating to their environmental
adaptations.

ACKNOWLEDGEMENTS

The authors thank Miss S. Hanson and Mr R. O. Ruehle for preparing the illustration,
Dr D. F. Hoese and Messrs G. F. Gross and N. Pledge for reviewing the manuscript, and
Miss J. Casaretto and Mrs J. Murphy for typing same. A number of colleagues kindly
provided records which enabled the provisional check-list to be compiled, namely M. N.
Feinberg, D. F. Hoese. W. Ivantsoff, J. S. Lake, R. J. McKay, J. R. Merrick and J. R. Paxton.

Preparation of this paper was supported by a grant awarded (1974-77) by the Austral-
ian Biological Resources Study Interim Council.

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256 Aust. Zool. 19(3), 1978

Observations on some Buprestidae (Coleoptera) from
the Blue Mountains, N.S.W.

T. J. HAWKESWOOD
Department of Botany, University of New England, Armidale, N.S.W. 2351

ABSTRACT

Forty-two species of Buprestidae in nine genera were collected and observed in
the Glenbrook area of the Blue Mountains N.S.W. during 1975-77. Specimens
of three of these species were also collected from Wentworth Falls and Medlow
Bath, in the higher Blue Mountains, during early 1977. A list of the jewel beetles,
their occurrence, food plants and dates of collection, is included. Field observ-
ations on flight behaviour, feeding, defence and escape mechanisms, grooming
and pollination are recorded and discussed. All of these aspects were not
observed in every species mentioned. Stigmodera macularia (Don.), S. variabilis
(Don.) and Cisseis leucosticta (Kirby) are figured.

INTRODUCTION

The Buprestidae commonly referred to as “jewel beetles” are well represented
in Australia, with over 800 described species (Britton, 1970). When conditions
are suitable e.g. high temperatures and profuse flowering and nectar-bearing by
food plants, some species may be found in abundance, especially those of the
dominant nectar-feeding genus Stigmodera. Individuals of foliage feeding genera
e.g. Cisseis, Ethon, Germarica and Paracephala may be found during the summer
months on their food plants, e.g. Acacia spp. (Mimosaceae) or Casuarina spp.
(Casuarinaceae) (Froggatt, 1907; Tillyard, 1926; McKeown, 1945).

Some species have a widespread distribution, e.g. Stigmodera erythroptera
Boisd. found in all states (Carter, 1931a), but others have somewhat limited dis-
tribution, e.g. Stigmodera magnetica Cart., from Western Australia (Glauert,
1948; Barker et al., 1956; Barker ef al., 1960; Barker and Edward, 1963).

Despite the wide occurrence of some species, the great number of species and
their abundance, very little is known about their biology (Hughes, 1975). This
may be due, in part, to the difficulty of locating many species, as it appears from
data at hand that numbers of individuals fluctuate from season to season; one
species may be rare or absent one year in a particular area but be common the next
season (Whitlock, 1947; Williams, 1977; and the present work).

Aust. Zool. 19(3), 1978 257

T. J. HAWKESWOOD

Since very little has been written about any aspect of the biology of Australian
Buprestidae, this study was undertaken in order to discover, within a relatively
small area in the Blue Mountains, N.S.W., the species present and aspects of their
behaviour in their natural habitat.

MATERIALS AND METHODS

(a) Stupy AREAS

The majority of observations were made on n living insects in an area of radius
approximately 6 km from the centre of Glenbrook, N.S.W. This area includes the
townships of Lapstone, Blaxland and Warrimoo, 2 km S.E., 3 km N.W. and 6 km
N.W. respectively, from the Glenbrook Post Office. Specimens of three species of
buprestid were also collected from Wentworth Falls and Medlow Bath, in the
higher Blue Mountains.

The township of Glenbrook is situated about 70 km (43 miles) by road,
west of Sydney, at an altitude of 163m (535 ft) above sea-level. Much of the
natural bushland in the immediate vicinity of Glenbrook and other areas in the
Blue Mountains has made way for residential development and no doubt the flora
and fauna have been affected in various ways by this disturbance.

(b) COLLECTION TIMES AND MAIN FLOWERING SPECIES

Most field data and specimens were collected between December 1975 and
February 1976 when Angophora floribunda (Sm.) Sweet, A. bakeri C. Hall, and
Leptospermum phylicoides (A. Cunn. ex Schau.) Cheel, major food plants for
Stigmodera, flowered profusely and between December 1976 and February 1977,
when Leptospermum flavescens Sm. was a dominant flowering plant in the areas
studied. Specimens of foliage-feeding species of Cissezs were collected during both
seasons but Germarica, Paracephala and Astraeus were only collected during the
1976/77 season. Specimens of Cyria imperialis. Fabr. were only collected during
1975/76 when its main food plant in the area, Banksia spinolosa Sm. was in flower.

Specimens were collected by hand and/or net, released after examination, or
kept for later examination and identification.

The majority of observations were made during periods of one to three hours,
between 1200 and 1700 hrs (Eastern Standard Time).

(c) WEATHER CONDITIONS

The Glenbrook area receives an average annual rainfall of about 80cm (32
inches) and temperatures range broadly from 1°C to 40°C. Weather conditions
during the two seasons were variable with daily temperatures ranging from 15°C
to 39°C. Good summer rains in excess of 203 mm (8 inches) fell during both
seasons. During days when observations took place, conditions were mainly fine
and clear with temperatures usually varying from 21°C to 39°C. The hottest day

258 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

recorded during the two-year period of observation occurred during the summer of
1976/77, when 41.5°C was recorded at Glenbrook on the 5th January, 1977.

(d) NOMENCLATURE

The nomenclature used for plant species follows that of Beadle et a, (£972):
The nomenclature used for Buprestidae are as follows: for Stigmodera (Carter,
1916; 1929; 1931a; 1931b), for Germarica, Paracephala, Curis, Cyria and Torre-
sita (Carter, 1929), for Melobasis (Carter, 1923b; 1929), for Cisseis (Carter,
1923a, 1929) and for Astraeus (Barker, 1975)

VEGETATION

The present vegetation of the Glenbrook area is one of dry sclerophyll forest
dominated by scribbly gum, Eucalyptus haemastoma var. sclerophylla Blakely, the
yellow bloodwood, E. eximia Schau., the red bloodwood, E. gummifera (Gaertn. )
Hochr., and the Sydney peppermint, E. piperita Sm., as well as the angophoras, i.e.
Angophora costata (Gaertn.) Druce, A. floribunda (Sm.) Sweet and A. bakeri
C. Hall. The understorey vegetation is largely composed of species of Acacia
(Mimosaceae); Persoonia, Hakea, Banksia and Grevillea (Proteaceae):

Pultenaea, Bossiaea, Dillwynia, Oxylobium, Gompholobium, Daviesia and Phyllota
(Fabaceae); Leptospermum (Myrtaceae); and Casuarina (Casuarinaceae). Other
small plants belonging to various plant families, including the grasses (Poaceae)
occupy the ground zone strata. In deep gullies and gorges numerous ferns and
rainforest tree species may be found, e.g. Syncarpia glomulifera (Sm.) Niedenzu
(Myrtaceae). Many of the plants mentioned above, e.g. Angophora, Leptospermum
and Acacia are food-plants for members of the Buprestidae (see Tables 1 and 2).

OBSERVATIONS AND DISCUSSION
(a) List of species, occurrence, food plants and dates of collection

This information is summarised in Table 1. The occurrence of buprestid
species was rated on a per season basis similar to that of Williams (1977). The
term “rare” indicates less than three specimens, “few” three to 10 specimens,
and “common” more than 10 specimens captured and/or observed during each
season of collection. Since this system was used by Williams (1977), comparisons
can be made with his findings.

One fact that has emerged as a result of the two seasons of observations is
the marked synchronization between the flowering of the food plants and the
appearance of the nectar feeding species of Stigmodera and Cisseis upon the
blossoms of these plants. Williams (1977) noted that he collected the most species
and the highest number of individuals during the final flowering phase of the
food plant (Leptospermum flavescens Sm.) at East Minto, New South Wales,
during 1972-1975. This observation generally agreed with those at Glenbrook.

Aust. Zool. 19(3), 1978 299

T. J. HAWKESWOOD

TABLE 1
List_of species of Buprestidae collected in the Blue Mountains, 1975-1977,

SEASON
SPECIES 1975/76 1976/77

Food Dates of Food Dates of

k Occurrence Locality Plant(s) Collection Occurrence Locality Plant(s) Collection

Subfamily Buprestinae
Tribe Buprestini

1. Astraeus pygmaeus * — — — few G 1,2 7-20 Jan.
van de Poll

2. Melobasis costata Macleay % — — — rare G 4(?) 10 Jan.

3. Melobasis cupriceps . — — — rare G 5, 6 28 Aug., 5 Dec
(Kirby) (?)

4. Torresita cuprifera rare G 10 8 Dec. few G 9, 10 5, 22 Dec.
(Kirby) :
Tribe Stigmoderini

5. Stigmodera macularia common G 1C 2 Dec.- rare G 9 1-2 Dec.
(Don.) (Fig. 1A and B) 15 Jan.

6. Stigmodera variabilis few G 10, 11 3-15 Jan. % — — —
(Don.) (Fig. 2A) 13

7. Stigmodera andersoni few G 11 3-8 Jan. = — — —

C. & G.

8. Stigmodera bella rare G 12 18 Dec.- few G, B 9 2-8 Dec.
Saund. 16 Jan

9. Stigmodera bicincta Boisd. rare G 12 5 Jan. rare B 9 6, 8 Dec.

10. Stigmodera brutella rare G 11 4 Jan “2 — =
Thoms.

11. Stigmodera crenata rare G 10 15, 18 common G. B 9, 10 1-11 Dec.
(Don.) Dec.

12. Stigmodera cruenta few G 10, 12, 10-15 Dec., rare G 9 18 Dec.
L. & G. 14 22 Jan.,

18 Feb.

13. Stigmodera octospilota few G 10 12-15 Dec. few GB; 9,10 1-18 Dec.,
L. & G. 6 Jan. MB 20 Dec.

14. See erythroptera few G 10 8-15 Dec. common G,B,W_ 9,10 1-18 Dec.

oisd.

15. Stigmodera (?) flavopicta = — — rare G 15 18, 29 Dec.
Boisd.

16. Stigmodera kerremansi rare G 12 5-6 Dec. rare Gr B 9 1 Dee!
Blackburn

17. Stigmodera kirbyi Guer. rare G 12 8 Dec. rare WW 9 19 Jan.

18. Stigmodera luteipennis rare G 10 15, 28 Dec. . — os —

L. & G.

19. Stigmodera nasuta Saund. rare G 10 12 Dec. ze — — —

20. Stigmodera decemmaculata few G 10 6-22 Dec., . — — —
(Kirby) 5 Jan.

21. Stigmodera puella Saund. “3 — — — rare B 9 9 Dec.

22. Stigmodera rufipennis few G 10 5-23 Dec. common G,B 9, 10 1-10 Dec.
(Kirby)

23. Stigmodera scalaris Boisd. rare G 12 12 Dec. common G,B 9,16 2-9 Dec.

24. Stigmodera spilota L.&G. rare G 5(?) 8 Feb. 3 — == =

25. Stigmodera spinolae rare G 10 12 Dec. - — — =
L. & G.

26. Stigmodera subpura Blkb. * — few B 9 2-10 Dec.

27. Stigmodera undulata few G 10, 11 3-12 Dec., few GAB 9 2, 8, 10 Dec.
(Don.) 28 Jan.

28. Stigmodera vigilans rare G 11 18 Dec. (?) rare Gas 9512 6-11 Dec.,
Kerr. (?) 3 Jan.

29. Curis caloptera (Boisd.) rare G 11 15 Dec. = — — —
Tribe Agrilini

30. Paracephala cyaneipennis - — — — rare G 2 5 Jan.
Blkb.

31. Germarica lilliputana es = — — common G, MB Dee 9, 17-19 Jan.
(Thoms. )

32. Cisseis acuducta (Kirby) rare G 6 14 Dec. rare G, L 6,17 18, 28 Dec.

33. Cisseis atroviolacea Thoms. ce —_ — — few B, W 9 1-12 Dec:

34. Cisseis cupripennis Ae —_ — — few G 6 22, 26 Dec.,
Guer. 10, 20 Jan.,

25 Feb.

35. Cisseis leucosticta common G 6 25, 28 Janrare G 7 4 Jan.
(Kirby) (Fig. 2B) 3-5 Feb.

36. Cisseis maculata L. &G. Be = — — rare B 9 5, 12 Dec.

37. Cisseis marmorata L.&G. rare G 8 16 Dec. few if TESS 14, 15, 20 Feb.

38. Cisseis notulata Germ. rare G 5 5 Jan. common G,B 5, 18 ne De

-15 Jan.

39. Cisseis pygmaea Blkb. few G 21 16-24 Jan., few G 21 3-18 Jan.

14-16 Feb.
40. Cisseis roseo-cuprea Hope “7 — — — few G 19 29 Aug.77.
41. Cisseis vicina Kerr. - _ — — few GaBeWwer 910 1-8 Dec.,
12 Feb.
Subfamily Chalcophorinae
Tribe Chalcophorini
42. Cyria imperialis (Fabr.) few G 20 22-31 Dec., * — —
1-3 Jan.
No. of species present 29 32

260 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

Key to Table 1.

Occurrence
* = no specimens observed or collected
rare= = 3 specimens collected
few = 3-10 specimens collected
common = > 10 specimens collected.

Locality

G = Glenbrook;
B = Blaxland;
l=) leapstomne:
W = Warrimoo;

WW = Wentworth Falls;
MB = Medlow Bath.

Food Plants
Casuarinaceae
1. Casuarina torulosa Ait.; 2. C. littoralis Salisb.;
3. C. nana Sieb. ex Spreng.

Poaceae (Gramineae )
4. Themeda australis (R.Br.) Stapf.

Mimosaceae
5. Acacia linifolia (Vent.) Willd.; 6. A. longiflora (Andrews) Willd.;
7. A. decurrens (Wendl.) Willd.; 8. A. parramattensis Tindale

Myrtaceae er

9. Leptospermum flavescens Sm.; 10. L. phylicoides (A. Cunn. ex Schau.) Cheel;
11. Angophora floribunda (Sm.) Sweet; 12. A. bakeri C. Hall;

13. Eucalyptus piperita Sm.

Pittosporaceae
14. Bursaria spinosa (Cav.) Druce.

Asteraceae (Compositae )
15. Cassinia compacta F, Muell.; 16. C. wncata A. Cunn. ex DC.

Fabaceae ( Papilionaceae )
17. Jacksonia scoparia R. Br.; 18. Dillwynia retorta (Wendl.) Druce var. retorta;
19. D. floribunda var. teretifolia Blakely.

Proteaceae
20. Banksia spinulosa Sm.

Sapindaceae
21. Dodonaea triquetra Wendl.

Aust. Zool. 19(3), 1978 261

T. J. HAWKESWOOD

TABLE 2

Numbers of buprestid species collected from dominant flowering plants during
each season in the study area

Number of Species

Heod ean 1975/76 1976/77
Lepiasnermum, flavescens. “SM oe ee ee ae es 0 19
Lophyticoides (A. Cunnséx, Schau) Cheel 32 ee ee 13 6
“incopnora.jlorivunda: (SWS) SWeEEU. se idecncc eee ot eee 6 0
ree DTICCRE AC LAND coe eg ee See ca SM ec te ee ae pee 6 1

During the 1975/76 season the dominant flowering species upon which
beetles were observed were Angophora bakeri, A. floribunda and Leptospermum
phylicoides. Flowering of A. bakeri in the lower Blue Mountains occurred through-
out most of December and into the first half of January, while A. floribunda
flowered later in December and extended into late January. Six species of Stig-
modera were collected from A. bakeri (see Table 2), mainly during early to
middle January, while five species of Stigmodera and Curis caloptera were collected
from the latter plant species, mostly during early January (Tables 1, 2). Generally,
buprestid species were not very common and often long waiting for beetles to
arrive at blossoms and intensive searching failed to locate any specimens.

Leptospermum phylicoides yielded 12 species of Stigmodera and Torresita
cuprifera. Flowering of this plant began in the last week of November and reached
a peak flowering period during middle to late December. Many plants were still
flowering by mid-January, 1976. Stigmodera macularia was the only buprestid
collected from this plant during the second week of January.

No buprestids were observed before December, but specimens may have
been present on early flowering plants of L. phylicoides in other localities where
collection did not occur.

Most species of Stigmodera showed preference for one or two food plants,
and only S. variabilis and S. cruenta were found on three different plant species
during the 1975/76 season (see Table 1).

During 1976/77 the angophoras, eucalypts and L. phylicoides flowered poorly
in the Glenbrook area, while the yellow tea-tree L. flavescens flowered profusely,
yielding 15 species of Stigmodera, three species of Czsseis and Torresita cuprifera
(see Tables 1, 2). Flowering of this species at Glenbrook began in the last week
of November, reached a peak during 8-10 December and had ceased by the 14
December, by which time petals had fallen off or blown off and fruit develop-
ment was well advanced. No buprestids were collected after the 18 December
from L. flavescens at Glenbrook. This represents a marked synchronization between
the flowering of the food plant and the appearance of adult beetles.

One specimen of Stigmodera kirbyi was collected on the 19 Jan. 1977 from
a late-flowering plant of L. flavescens, 5.5 km S.E. of Wentworth Falls, and a

262 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

single example of S. octospilota was collected from a poorly flowering L. phylicoides
near Medlow Bath, in the higher Blue Mountains on the 20 Dec. 1976. These
were the only Stigmodera collected from Leptospermum after the 18 Dec. 1976.

Williams (1977) collected 16 species of Stigmodera from L. flavescens at
East Minto, N.S.W., during 1972-75, which is 50 km S.E. of Glenbrook; of
these only six species were collected during 1976/77 at Glenbrook, while
Stigmodera bicincta, S. crenata, S. octospilota, S. erythroptera, S. kerremansi,
S. scalaris, S. subpura and S. vigilans (?) were not collected in any season by
Williams, but most of the above were present both seasons at Glenbrook. Williams
(1977) records the interesting observation of Neocuris guerini, Curis caloptera
and Cyria imperialis on L. flavescens; none of these species were observed on
L. flavescens at Glenbrook (N. guerini was not collected at all from any plants);
on the other hand Torresita cuprifera, Cisseis atroviolacea, C. maculata and C.
vicina also found on L. flavescens at Glenbrook were not recorded by Williams.
Williams (1977) noted considerable seasonal variation both in number of species
and numbers of individuals present during his three years of observation. This
observation was also noted during this study (Tables 1, 2).

It appears that the presence of many feeding beetle species is strongly
associated with the species of plant flowering, and rainfall and temperature
appeared to have little effect on their occurrence. One example is Cyria imperialis,
which usually shows specificity to Banksia spp. (Froggatt, 1907; Tillyard, 1926;
McKeown 1945), but was recorded on Leptospermum flavescens by Williams
(1977). At Glenbrook it was only found during late December and early January
1975/76 on Banksia spinulosa Sm., which flowered well during this season. This
plant did not flower during 1976/77 and extensive searching amongst Banksia
bushes during the summer failed to find any of these buprestids.

Other examples of nectar-feeding buprestids showing apparent specificity
for one species of plant and appearing only during one season include Stigmodera
andersoni, S. brutella, S. flavopicta, S. luteipennis, S. nasuta, S. decemmaculata,
S. spinolae, S. subpura, Curis caloptera, Cisseis atroviolacea and C. maculata. It is
felt, however, that with further observations many of these species may be found
to frequent more than one species of plant.

(b) Flight behaviour

Observations on flight were obtained during the 1976/77 season. Periods
spent in flight varied from one to greater than 70 seconds, depending on the genus
and size of species (Table 3). Generally, the larger species, such as Stigmodera
rufipennis and S. undulata, were more sedentary than the smaller more active
species, e.g., S. scalaris and S. crenata. The very active Cisseis atroviolacea, C.
vicina and C. maculata, observed during the 1976/77 season, spent most of their
time amongst the flowers of Leptospermum flavescens, and usually flew when
disturbed or closely approached. Observations showed that standing or movement

Aust. Zool. 19(3), 1978 263

T. J. HAWKESWOOD

of the author at distances of 15 to 50 cm from the feeding beetles aroused them,
causing them to take immediate flight or to display the “free-fall and flight-escape
mechanism” (see later). Beetles, when disturbed while feeding, usually ceased
movement for about one second, quickly opened their elytra outwards and flew
‘apidly upwards or slightly horizontally.

One small specimen of S. macularia, the largest species examined, observed
yn the 2 Dec. 1976 at Blaxland, was very inactive, remaining motionless for more
han five minutes before being collected, examined and later released (Table 3).

TABLE 3

Data on flight, feeding, defence and escape mechanisms for some species of
Buprestidae observed in the study area during 1975-1977

Periods spent Periods in Feeding Defience
Species feeding flight heights and escape
(secs) (secs) (m) mechanisms +
A. Small species 5-13 mm
(total length)
1. Stigmodera crenata (Don.) .... 3-15 10->25 1.2-2.0 1... 2-2
2. Stigmodera erythroptera
BOISGS ie core ng Lo Hen od 2-25 5-30(?) 122-322, 2S 5
3. Stigmodera kerremansi Blkb. .... 5-15 S507) 0.7-2.0, 6.0 1 5-4
4. Stigmodera scalaris Boisd. ...... 3-20 5-25(?) 1.2-2.2 1 bey: |
S$. Cisseis atroviolacea Thoms, .... >120 pc Ge) ie1=2.5 1 3
6. Cisseis: maculata ..& G. .:....2 >90 * 15-20 1 3
7. Cisseis pygmaea Bikb.. ............ * 1-10 0.5-1.3 1
Si CISSEIS, VICING INCL es cnc as 120 ** 1.5-2.0 1 3
B. Large species >13 mm
(total length)
9. Stigmodera macularia (Don.) 20->300 15-30 0.8-2.5 Des
10. Stigmodera rufipennis (Kirby) 25-195 15-70(?) 1.4-2.8 3 5
11. Stigmodera undulata (Don.) .... 25-125 3-20(?) 1.0-2.0 2 014
12. Stigmodera_ variabilis (Don.) >180 >5(?) 3.2, 6.0-9.0 4
KEY

(2?) Specimens of these species were observed to fly away from food plants into surrounding
bushland or out of sight. Hence the larger time(s) given is the time from departure of
the food-plant to the time when the beetle(s) disappeared from sight.

* — No data available or not observed.

+ Defence and escape mechanisms

Free-fall and flight

Free-fall and death feign

Upward flight

Bright colour(s) on dark background

Batesian mimic.

A BRWN eR

When in flight, all species of Stigmodera flew with abdomen pointed down-
wards, elytra positioned horizontally, and head and pronotum pointed slightly
forward. They made circular flights around the largest bushes of L. flavescens,
which had the most flowers open. They often reversed direction and circled back

264 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

and forth over and around the tops of the bushes before landing. Slight disturb
ances, causing branches to move suddenly, caused individuals to take flight.

Barker (1975) first recorded the presence of a spring mechanism, involving
the release of the elytra from the closed position, in the genus Astraeus. When
the beetle releases the spring, the elytra flick open with considerable force, enabling
the insect to be flung upwards. Barker (1975) recorded the height achieved by
some species as ‘‘several metres’; observations on A. pygmaeus at Glenbrook,
on the 7 Jan. 1977, showed the height obtained before flying in other directions
to be approximately 0.3-0.6 m. Beetles observed were quite active and when
approached would quickly flick to another branchlet, often turning in one direction,
reversing and flying (?) or flicking off again. Occasionally individuals were observed
to fall quickly to another Casuarina branchlet, [species of Astraeus are mainly
found on Casuarina plants, Barker, (1975) ], either resting there or flicking off
to a higher altitude (up to 2.5 m). From the observations of these uncommon
insects at Glenbrook it appears that individuals fly rarely and mainly remain on
branchlets of their food plant Casuarina torulosa Ait; it appears that the spring
mechanism is used to propel the insect upwards or downwards, and the wings
used only to reach a convenient resting post, which was unable to be reached
by the spring mechanism alone.

Germarica lilliputana, observed during January, 1977, at Glenbrook and
Medlow Bath, tended to remain on Casuarina torulosa Ait., or C. nana Sieb. ex.
Spreng. branchlets, and flight was not observed.

Cyria imperialis, a strong flier producing a loud whirring noise while in the
air, was observed during January, 1976, at Glenbrook. The species appears to be
a wary insect, and flies far and fast when disturbed or approached; three specimens,

observed 2.2 km N.E. of Glenbrook, displayed the ‘free-fall and death feign”’

escape mechanism (see later).

Flight was not observed in Melobasis, Curis, Paracephala or Torresita.

(c) Feeding biology

The Leptospermum, Angophora and Eucalyptus flowers (Family Myrtaceae),
upon which many species of Stzgmodera were observed and/or collected during
1975/77, are constructed so as to give easy access to the nectar supply for sugar-
requiring species of Buprestidae and other species of insects. (Fig. 1 A and B, and
Fic. 2 A and BY

The small species of buprestid, upon landing on a blossom, proceed to move
through or over the series of stamens surrounding the ovary of the flower and
to the areas where the numerous secretory cells are situated. The nectar collects
in a hollow or groove formed by the edge of the ovary summit and the inner
surface of the floral tube, but may spread over the top of the ovary in flowers
with a considerable nectar supply. The nectar of these plant genera is sweet-

Aust. Zool. 19(3), 1978 265

T. J. HAWKESWOOD

Stigmodera macularia on flowers of Angophora bakeri.

S. macularia on Leptospermum flavescens.
(Note deep pits on elytra and puncturing on pronotum).

Aust.

Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

smelling and the sugar content appears to be composed, in Leptospermum at least,
of glucose and fructose (Dr. H. A. Ford, pers. comm.), which is the necessary
energy source for these active beetles. The buprestid, with head extended or
placed downwards into the flower, is able to draw up through the rostrum (present
in Stigmodera) the sugary liquid.

Small species of Stigmodera examined, e.g. S. scalaris, S. bella, S. bicincta,
S. erythroptera, 8. kerremansi, S. nasuta and S. subpura, alight on a flower (Lepto-
spermum) and, aided by quick movements of their legs and quivering movements
of the antennae, proceed to the nectar-bearing areas and position themselves head
down into the flower. The elytra remain closed, and extend above the stamens,
while the legs are used for balance during feeding.

Time spent feeding on Leptospermum flavescens varied from two seconds to
approximately five minutes, most probably depending on the amount of nectar
present in the flowers and the activity of the species (Table 3). The larger and
slower moving species, e.g. S. rufipennis, spent more time feeding on the flowers
of the tea-tree and less in flight than did the smaller species. The latter moved
quickly and readily from flower to flower (although they often remained several
minutes feeding within one flower).

Of the beetles observed feeding, either on Leptospermum phylicoides during
1975/76, or on L. flavescens during 1976/77 (both at Glenbrook), most species
of Stigmodera, Cisseis atroviolacea, C. vicina and C. maculata fed singly and but
for a few occasions remained isolated and relatively distant (15 cm or more
away) from members of the same species, and also individuals of other species of
buprestid. On the 5 Dec. 1976 a pair of S. rufipennis was observed feeding from
1 to 8 cm away from each other. However, since most species feed singly, this
suggests that a spacing mechanism is involved, possibly enabling individuals to
feed undisturbed and to gain a more equal share of the available nectar. However,
the latter suggestion would be difficult to prove under field or laboratory conditions.

Most specimens of Stigmodera and Cisseis which were observed on Lepto-
spermum visited flowers which were more than 1 m above ground level. The
smaller bushes of L. flavescens, less than 1 m in height, were seldom visited by
any buprestid. One specimen of S. macularia, collected on the 1 Dec. 1976
frequented a group of flowers of L. flavescens situated 0.8 m above ground level.
Specimens of Cisseis pygmaea, observed during Jan.-Feb., 1976 and 1977, fre-
quented young plants of Dodonaea triquetra Wendl., less than 1.3 m in height
(Table 3). For Leptospermum flavescens, the height range for flower visitation
was > 1 and <3.5 m. Stigmodera flew at heights also within this range and
seldom flew higher than 4 m. One individual of S. rufipennis flew approximately
7 m into the air after being disturbed on the 3 Dec. 1976. Stigmodera variabilis
preferred greater heights when feeding. Specimens of this species were observed
or collected only during Jan., 1976, feeding on the flowers of Angophora bakeri
and Eucalyptus piperita at heights ranging between 6-9 m, and on the 6 Jan.

Aust. Zool. 19(3), 1978 267

T. J. HAWKESWOOD

FIG. 2.—A. Stigmodera variabilis on flowers of Angophora bakeri.

B. Cisseis leucosticta feeding on a stem of Acacia decurrens.

268 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

1976 one individual was collected on a large Leptospermum pbhylicoides plant at
a height of about 3.2 m. Curis caloptera, Stigmodera kerremansi and other species
of Stigmodera collected during January, 1976, at Glenbrook, fed upon flowers of
Angophora floribunda at heights of over 6 m. Most species of Stigmodera found
feeding on Leptospermum phylicoides during 1975/76 frequented flowers at
heights of > 1 m and <2 m. It appears, then, that nectar-frequenting species
of Buprestidae have a height preference for flowers (depending on the species
of food plant); those flowers less than 1 m above ground level are seldom visited.
Those flowers visited appear to be mainly the top flowers, or flowers at tips of
branches. It may be that the top flowers contain more nectar than the bottom
flowers, and so these attract more insects. However, only superficial examination
of the nectar supply of bottom and top flowers by the naked eye could be
undertaken. From this examination it appeared that the bottom flowers were
similar in condition to those higher up on the plant.

There appears to be no height preference at Glenbrook for species which
are foliage eating, i.e. Cisseis. Melobasis, Paracephala and Germarica.

(d) Flower pollination by buprestids

Because of the habit of nectar-feeding Stigmodera, their movements over
flowers and the structures of the Leptospermum, Angophora or Eucalyptus flowers
themselves, it would appear that buprestid beetles as a whole are important
pollinators of these plants. Many Stigmodera examined during Dec.-Jan., 1975/76
and 1976/77, had pollen attached to the sides of the pronotum and head region,
as well as on the tarsi, other leg segments and the elytra.

Some species of Stigmodera have deep pitting on the elytra (Fig. 1B). These
pits may act as reservoirs for fallen pollen, which may be carried by the beetle
~ to other flowers. S. macularia is one such species; observations so far on this
species, however, have not shown that the deep pits of the elytra are used for
the transfer of pollen.

Several specimens of Cisseis atroviolacea and C. vicina, collected on the 4
Dec. 1976, near Blaxland, were copiously covered in pollen from Leptospermum
flavescens.

(e) Defence and escape mechanisms

(i) Bright colours—many species of jewel-beetle observed were brightly
coloured with dominant colours of red, yellow, orange, green and/or blue, Le.
bright colours on a dark background (see Tables 3, 4).

(ii) Dull colours—most species of Cisseis, Germarica lilliputana, Paracephala
cyaneipennis and Melobasis collected were dull in colour—browns, dull greens,
blues and dark shades. The green tint of a vivid species of Melobasis (Table 4)
appeared to match the colour of Acacia longiflora (Andrews) Willd. leaves upon

Aust. Zool. 19(3), 1978 269

T. J. HAWKESWOOD

which the insect was resting; this cryptic colouration may aid the species to escape
from would-be predators. Small size and colouration of Germarica lilliputana are
two factors which may aid in the protection of this species.

TABLE 4

Heights at which beetles were collected on host plants and defence and escape
mechanisms observed for some species of Buprestidae from the Blue Mountains,
N.S.W. (1975-1977)

Heights Defence and escape
(m) mechanisms -+--
A. Small species < 16 mm
(total length)
tS SGISSCLS ICHCOSTIGIO: CIRILDY)) ~~ soe. ae ees ee 0.7-1.2 Ds 4
DeACISSCLS HINOANMOL ALA Wen Gran eae eee oe 0.6-1.2 4
See CISSCLS NOMLGLG AGEL 0 ee a he el i 0.9-1.5 1
Aa: Cisseis: TOSCO-CUPTED MAOPe <2. see Oh uaa 0.4-0.6 2
5. GErMmarico. Wiliputana = GlnOms.) oi. eee 0.5-3.5 5
6.) Melobasis cupriceps. CKitby)) Cri ee 1.0-2.0 5
i= ‘Paracephala-cyaneipennis BIKO.. 2 aie 0.3 *
8. Stigmodera (?). flavopicta Boisd.. f).cc:..0.s:0s0s000 1.2-1.5 1 3 6
B:. Large species > 16 mm
(total length)
Oe Cyrigi tm perlas: CF abrw,ceete. aetna sia eee 1.1-1.5 2 6
KEY

+ Defence and escape mechanisms

= Free-fall and flight

= Free-fall and death feign

= Upward flight

= Defence secretions

= Cryptic colouration

= Bright colour(s) on a dark background
Not observed or no data available

? = The identification of this species is uncertain.

|| An RWNR

(iii) Batesian mimicry—this phenomenon has been noted in Stigmodera
(Nicholson, 1927; Britton, 1970). Batesian mimics in the author’s collection were
S. rufipennis, S. nasuta, S. erythroptera, S. subpura and S. spinolae. Nicholson
(1927) and Britton (1970) illustrate in colour other mimics from the beetle
families Cerambycidae, Curculionidae (Belidae), Buprestidae, Cantharidae and
Oedemeridae. These beetles mimic the distasteful betles of the genus Metriorrhyn-
chus Lycidae (or Lampyridae of older works). The most common of these beetles
is M. rhipidius Macleay. It is interesting to note that this and other species of
Metriorrhynchus were also present on Leptospermum flavescens, together with
most species of buprestid mentioned above. (See also list of insects recorded
visiting L. flavescens, apart from members of the Buprestidae, Table 5).

(iv) Free-fall and flight—several species of Cisseis (including C. pygmaea
and C. notulata) and Stigmodera crenata, S. scalaris, S. karremansi and S. (?)

270 Aust. Zool. 19(3), 1978

F

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

flavopicta, were observed to fall from a branch or flower when disturbed or
approached, and then to fly away to a safer location immediately before hitting
the ground (Tables 3, 4). This is no doubt a successful escape mechanism when
attacked from above by a predator such a bird.

(v)Free-fall and death fetgn—Stigmodera crenata, S. macularia, S, erythrop-
tera, S. bella, 8. undulata, Cisseis leucosticta, C. roseo-cuprea and Cyria imperialis,
all exhibited this escape mechanism when disturbed (Tables 3, 4). Hawkeswood
(1977) has made a brief comment about this escape mechanism exhibited by

TABLE 5

The occurrence of identified insect species, other than the Buprestidae, visiting the

leaves or flowers of LEPTOSPERMUM FLAVESCENS S™m.,

collected at Glenbrook and Blaxland, N.S.W., during December 1-12, 1976
Order Family Genera and. Species Occurrence
Hymenoptera Apidae Apis mellifera L. common
Sphecidae Exeirus lateritius Shuck. rare
Psammocharidae Cryptochielus fulvidorsalis Turn. rare
(Pompilidae )
Formicidae Iridomyrmex sp. few
Polyrhachis ammon Fabr. rare
Lepidoptera Amatidae
(Syntomidae) Eressa paurospila Turn. (?) common
Hemiptera Reduviidae Poecilosphrodrus sp, (?) few
Diptera Asilidae Chrysopogon crabroniformis Roder. rare
Coleoptera Mordellidae Mordella leucosticta Germ. rare
Mordellisterna spp. (2) common
Scarabaeidae Phyllotocus macleayi Fisher common
Phyllotocus sp. common
Polystigma punctata (Don.) rare
Liparetrus discipennis Gter. few
Repsimus manicatus manicatus (Swartz) rare
Glycyphana brunnipes (Kirby) few
Lycidae Metriorrhynchus rhipidius Macleay common
M. irregularis Waterhouse few
M. heterodoxus Lea few
M. eremitus Fabricius few
Cantharidae Chaulignathus pulchellus (Macleay) rare
Heteromastix anticus Blackburn common
H. simplex Lea common
Selenurus sydneyensis Blackburn few
Chrysomelidae Paropsis spp. (2) rare
Cleridae Trogodendron fasciculatum Schreib. rare
Scrobiger sp. few
Aulicus sp. : few
Opilo congruus Newman common
Elateridae Anilicus sp. common
KEY .
Tare = < 3 specimens observed; few = 3-10 specimens observed: common = > 10
specimens observed; ? = identification of these species is uncertain.
Aust. Zool. 19(3), 1978 Pai

observed and/or

T. J. HAWKESWOOD

some Australian members of the Chrysomelidae (Coleoptera), but, from the
literature available, little has been noted on this escape mechanism in the Bupres-
tidae. Whitlock (1947), noting observations in the field on some Western
Australian buprestids, records “‘jewel-beetles I am acquainted with have the habit
of dropping down in an inert condition” when he approached or disturbed feeding
beetles. The buprestid beetles observed at Glenbrook exhibited an immediate fall
to the ground from heights ranging from 0.6 m to 2.0 m. On the ground they
would lie positioned upside-down with a uniform coloured undersurface facing
upwards. Between small pebbles, small rocks, sand, or amongst grass and other
plants, the beetles appeared well camouflaged.

(vi) Upward flight—most species of Stigmodera immediately flew off in a
vertical or angular direction if violently disturbed by a gust of strong wind causing
a branch to move suddenly where an individual was feeding.

(vii) Defence secretions—Defence secretions have been noted to be exuded
by Torresita cuprifera, Cisseis leucosticta and C. marmorata.

One specimen of T. cuprifera, captured on the 13 Dec. 1976 on Leptospermum
flavescens exuded, from the mouth, a non-odorous crimson-coloured liquid when
handled. A pale yellow non-odorous liquid was also exuded from the mouth in
Cisseis leucosticta and C. marmorata, which stained the skin of the author’s fingers.

(f) Tarsal and antennal cleaning

These aspects of grooming in Buprestidae were only observed in Stigmodera
undulata, S. rufipennis and S. macularia in this study.

Individuals observed stroked the tarsal segments of the forelegs through the
mouth region, starting at tarsal segment I and continuing to tarsal segment IV.
Having completed one to three strokes in this manner for one tarsus, individual
beetles would then repeat this action for the tarsus of the other foreleg. The
exchange from one foreleg to the other was occasionally repeated twice or more,
but usually one series of cleaning for each foreleg was administered. After the
completion of tarsal cleaning, the antennae would be wiped several times by the
previously cleaned tarsi of the forelegs. This is most probably an effective method
in removing dust and pollen grains, which may impede the sensory function of
the antennae. Tarsi of legs II and III were not observed to be cleaned; this is
most probably because the beetles are unable to reach them with their mouthparts.
However, these tarsi may be cleaned by other tarsi, although this was not observed.

(g) Predators and relationships with other insects

Table 5 lists the main insect species collected and observed feeding or visiting
flowers of Leptospermum flavescens, excluding the Buprestidae. The list excludes
unidentified members of the Diptera (Tabanidae, Muscidae and Syrphidae) and
Hymenoptera (Sphecidae, Ichneumonidae). Several spiders of the family Argiopi-
dae were also observed living amongst the branches of the tea-tree.

272 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

Predation of insect species by other insects was observed in Apis mellifera L.,
which was predated upon by the red and black assassin-bug Poecilosphrodus sp.
(2), a resident insect amongst the leaves of the tea-tree. Araneus spiders, resident
amongst the foliage of the tea-tree, had caught numerous specimens of Selenurus
sydneyensis Blkb., Metriorrhynchus spp., and Chaulignathus pulchellus (Macleay),
but careful examination of webs failed to find any captured buprestids.

Observations also showed no physical contact between buprestids and other
species of insects.

On the 4 Dec. 1976 a 4 eastern spinebill (Acanthorhynchus tenuiostris )
spent about 10 minutes feeding on insects frequenting a stand of L. flavescens
1.7 km E. of Blaxland; however, due to quick movements of the spinebill through
and over the tea-tree shrubs, the author was unable to determine if buprestids
were included in the bird’s diet.

SUMMARY

(a) There appears to be a marked synchronization between flowering of
food plants and the appearance of adults of nectar-feeding species of Stigmodera,
Cisseis and Torresita. Generally, most individuals and species were found just
before and during the final flowering phases of the food plants when most flowers
were open and contained large quantities of nectar. No nectar-feeding buprestids
were found on plants either before or after flowering had occurred.

(b) Most species of Stigmodera showed preference for one or two food
plants and only S. variabilis and S. cruenta were collected on three different
plant species during 1975/76. Many species of Stigmodera were collected only
during one of the two seasons, while some, e.g. S. macularia, were collected during
both seasons. Since a strong degree of specificity for a particular food plant appears
to exist with certain nectar-feeding buprestids, the absence of beetles may have
been due to the plants’ failure to flower, or flower poorly.

A dead specimen of Melobasis costata was collected on a grass stalk during
January, 1976, while a slow-moving specimen of Stigmodera spilota was collected
from a non-flowering plant of Acacia linifolia (Vent.) Willd. during February,
1976, so it is unlikely that these are the food plants for these buprestids. For the
remaining 40 species of buprestid collected, data on food plants have been given.

(c) Generally, larger species of Stigmodera were more sedentary than the
smaller, more active species of Stigmodera and Cisseis.

Specimens of three species of Cisseis observed on Leptospermum flavescens
during 1976/77 flew upwards or exhibited the “free-fall and flight” escape
mechanism when approached within 15-50 cm from where beetles were feeding.
This appears to indicate an acute awareness of predators and efficient methods
of escape.

Aust. Zool. 19(3), 1978 273

T. J. HAWKESWOOD

Many Stigwodera made circular flights and often flew back and forth over
and around the tops of Leptospermum bushes before landing to feed. It appears
that there is some type of spacing mechanism which exists between nectar-feeding
buprestids, enabling beetles to feed undisturbd and to procure an adequate supply
of nectar. More observations, however, are needed on this behaviour.

Barker (1975) was the first to record the presence of a spring mechanism
in the genus Astraeus. Observations on A. pygmaeus at Glenbrook showed that
beetles were quite active and used the spring mechanism to propel themselves
upwards or downwards to reach other resting posts. Heights obtained were found
to be about 0.3-0.6 m.

(d) Larger and slower moving species of Stigmodera spent more time upon
and feeding within flowers between flights than did the smaller species, which
usually spent only several seconds feeding within one flower before flying to
another flower.

There appears to be a height preference for nectar-feeding species of Stig-
modera and Cisseis. For Leptospermum flavescens height visitation was mostly
between 1 and 3.5 m, for L. phylicoides 1 to 2 m, Angophora floribunda > 6 m,
and for A. bakeri 6 to 9 m. The blossoms visited appeared to be the topmost
flowers, or flowers at tips of branches. Superficial examination showed that the
lower flowers were in similar condition to those above; therefore quantity of nectar
did not appear to determine the height preferred for feeding.

There appears to be no height preference for buprestids which are foliage
feeders, e.g. Paracephala, Germarica and Cisseis which were found at various heights
on different food plants.

(e) Many specimens of Stzgmodera and nectar-feeding Cisseis were observed
to carry pollen of their food plants. It appears that these beetles may be important
pollinators of these plants.

(f) Several escape mechanisms have been observed, two of which have been
descriptively termed in this paper, the “free-fall and flight” and the ‘‘free-fall and
death feign” escape mechanisms. They appear to be efficient methods of escape (as
verified by human disturbance), but there have been no observations on escape
from natural predators.

The defence secretions occurring in three species of buprestid appear to
represent the first record of such secretions in the Australian Buprestidae. The
usefulness of these secretions in defence against natural predators has not been
observed.

(g) Observations on tarsal and antennal cleaning by three species of Stig-
modera have been recorded.

(h) A large number of insect species belonging to numerous families were
found visiting blossoms of Leptospermum flavescens at Glenbrook and near

274 Aust. Zool. 19(3), 1978

OBSERVATION OF BUPRESTIDAE (COLEOPTERA)

Blaxland, N.S.W., during December, 1976. Spiders (Araneus sp). had captured
numerous beetles, but careful examination of the webs showed that no buprestids
had been captured.

ACKNOWLEDGEMENTS

I would like to thank Professor A, F. O'Farrell of the Zoology Department and Dr N.
Prakash of the Botany Department, University of New England, Armidale, N.S.W., for
examining and commenting on the manuscript, for the use of the University of New
England Insect Collection for assistance in species identification, for the use of the Macleay
Museum, University of Sydney, and to Mr Allan Davies for assistance while I was at the
Museum. Lastly, but not least, I would like to thank my brother Brian for accompanying
me on most of my collecting trips, assisting in collection and data recording, and for
photographing the species figured in this paper.

i a

REFERENCES

BARKER, S. (1975). Revision of the genus Astraeus Laporte & Gory (Coleoptera:
Buprestidae): Trans, R. Soc. S.A. 99, 105-141, text figs, 1-25.

BARKER, S. & D. H. EDWARD (1963). Corrections to type localities of three species of
Western Australian Stigmodera (Buprestidae, Coleoptera). West. Aust. Nat. 8, 169-171.

BARKER, S., D. H. EDWARD & J. A. L. WATSON (1960). The distribution of the jewel-
beetle Stigmodera (Castiarina) magnetica Cart. West. Aust. Nat. 7, 168.

BARKER, S., R. P. McMILLAN & J. A. L, WATSON (1956). A jewel beetle, Stigmodera
(Castiarina) magnetica Cart. West. Aust. Nat. 5, 143-144, plate 1.

BEADLE, N. C. W., O. D. EVANS & R. C. CAROLIN (1972). Flora of the Sydney Region.
A. H. & A. W. Reed, Sydney, 724 pp.

BRITTON, E. B. (1970). Jewel beetles (Buprestidae: Coleoptera). Chapter 30, In: The
Insects of Australia. C.S.7.R.O., Melb. Uni. Press, Carlton, Victoria, 1029 pp.

CARTER, H. J. (1916). Revision of the genus Stigmodera and descriptions of some new
species of Buprestidae (Order Coleoptera). Trans & Proc. R. Soc. S.A. 40, 78-144,
plates IX & X.

CARTER, H. J. (1923a). Revision of the genera Ethon, Cisseis and their allies (Buprestidae),
Proc. Linn. Soc. N.S.W. 48, 159-176.

CARTER, H. J. (1923b). A revision of the Australian species of the genus Melobasis (Family
Buprestidae, Order Coleoptera), with notes on allied genera. Trans, Ent. Soc. Lond.
71, 64-105, plates I, II.

CARTER, H. J. (1929). A check list of the Australian Buprestidae. Aust. Zool. 5, 265-301.

omnes J. (1931a). Check list of the Australian Buprestidae, Corrigenda. Aust, Zool. 6,

CARTER, H. J. (1931b). Notes on the genus Stigmodera (Family Buprestidae), (together
with descriptions of a new species of and a retabulation of the subgenus Casfiarina).
Aust. Zool. 6, 337-367,

FROGGATT, W. W. (1907). Australian Insects. W. Brooks & Co., Sydney, 449 pp.

GLAUERT, L. (1948). A rare jewel beetle. West. Aust. Nat. J, 129-130.

HAWKESWOOD, T. J. (1977). Those mighty plant defoliators—the Chrysomelidae. Wildlife,
Aust. 14, 11-14,

HUGHES, R. D. (1975). Living Insects, Collins, Sydney, 304 pp.

McKEOWN, K. C. (1945). Australian Insects, Royal Zoological Society of New South
Wales, Sydney, 303 pp.

NICHOLSON, A. J. (1927). A new theory of mimicry in insects, Aust. Zool. 5, 10-104,
text figs, 1-3, plates I-XIV.

TILLYARD, R. J. (1926). The Insects of Australia and New Zealand. Angus & Robertson,
Sydney, 560 pp.

WHITLOCK, F. L. (1947). West Australian jewel beetles. West. Aust. Nat. 1, 9-13.

WILLIAMS, G. A. (1977). A list of the Buprestidae (Coleoptera) collected from Lepto-
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Aust. Zool. 19(3), 1978 275

Gait of the Brush-tailed Possum
(Trichosurus vulpecula)

ANGELA J. GOLDFINCH
Department of Physiotherapy, St. Vincent’s Hospital,
Victoria Parade, Fitzroy, Victoria

R. E. MOLNAR
School of Anatomy, University of New South Wales,
P.O. Box 1, Kensington, N.S.W.

ABSTRACT

Locomotion of Trichosurus vulpecula was studied on the treadmill and under
quasi-natural conditions in a wire enclosure. On the treadmill Trichosurus used
a walk and a half-bound. In the enclosure a third gait, the bound, was also
observed. The walk was used at low speed and during climbing, and the bound
on steeply inclined surfaces. Modified gaits were also observed.

INTRODUCTION

Analysis of locomotion has usually been conducted on the larger forms of
both marsupials and placentals. To date there has been no detailed work on the
locomotion of Trichosurus vulpecula (the brush-tailed possum), or, indeed, on
most non-macropod marsupials. T. vulpecula has often been used in physiological
experiments (e.g. Anderson, 1937; Burke, 1954; Dawson, 1969; Gilmore, 1969,
etc.), and for these reasons description of the normal locomotion is of interest.
Such study would help to characterize the normal behaviour of T. vulpecula, as
well as to add to our knowledge of the locomotion of small mammals.

Trichosurus is an arboreal, nocturnal, mainly herbivorous creature. Their
numbers are not being reduced by the expansion of civilisation in Australia, and
they have successfully colonized New Zealand. Further information about the
biology of T. vulpecula may be found in Troughton (1941) and Tyndale-Biscoe
(1973').

To help elucidate the locomotion of T. vulpecula we undertook to study
the locomotion both under laboratory and quasi-natural circumstances.

Aust. Zool. 19(3), 1978 277

TRICHOSURUS GAIT

METHODS

The gaits of Trichosurus vulpecula were studied under two conditions, on a
treadmill and in a wire enclosure (10 feet square). In the enclosure were placed
a number of cut tree branches, from about 5 to about 30 cm in diameter. These
branches were placed in orientations ranging from nearly horizontal to vertical.
Locomotion under both conditions was recorded using high-speed cinematography.

Cinematography of the two animals on the treadmill was conducted by
Dr. R. Baudinette of the Department of Zoology, Flinders University, who kindly
made his films available to us. The treadmill formed the floor of a small cage
with transparent sides. The animals were trained to move at speeds of 2.8, 4.2,
5.5 and 6.8 km hr'!. A mirror inclined at 45 degrees to the back wall, and placed
in front of the animal, permitted filming of the anterior and lateral aspects
simultaneously. The film was exposed at 128 frames per second. These films were
analysed using the methods of Hildebrand (1966; 1977) and Dagg and de Vos
(1968a; 1968b). Consecutive frames of the film were individually inspected during
projection upon a screen. The gait diagram was then constructed by noting which
limbs were in contact with the substrate in each frame, and representing this
contact as a horizontal line in the gait diagram (e.g. Fig. 1).

RF re oy aE ES TES EER PEL Man Re a
LF
RH
LH

RF —_——
LF
a
LH

LF
RH
LH

RF
LF
RH
LH ql

WALK ii ee eee sO frames

FIG. 1.—Gait diagram of five walking strides of Trichosurus vyulpecula (on a treadmill).
The diagram is “read” from upper left to lower right. Each vertical bar marks
the beginning of a stride. Further explanation is given in the text. Ten frames
of the film represent 0.08 seconds. The individual feet are indicated on the
vertical line at the left, RF indicating the right fore-foot, LF the left fore-foot.,
RH the right hind-foot, and LH the left hind-foot.

278 Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

Hildebrand has used two sets of variables (one set for each type) to
characterise symmetrical and asymmetrical gaits. These variables are determined
from the inspection of gait diagrams. Suitable combinations of these variables are
then plotted as graphs, each point representing a stride. The fields of these graphs
may be divided into areas each representative of a specific type of gait.

In the enclosure films were made after allowing several days for the animals
to adjust to the enclosure. The films were exposed at 72 frames per second and
analysed, as were the treadmill films. Due to the problems of filming free-ranging
animals, relatively few film sequences were adequate for analysis, but it was
possible to construct gait diagrams of at least one full stride of each gait reported.

60

18)
oO

LF lag after LH: %S.I.

50 60 70

Hind contact [ave.|: %S.lI.

FIG. 2.—Results of analysis, after the method of Hildebrand (1966), of walk by fT.
vulpecula (on a treadmill). Each point represents one stride, and the average
is indicated by the star. The ordinate presents the period by which the left fore
footfall lags after the left hind footfall taken as a percentage of the stride interval.
and the abcissa, the average duration of substrate contact of one or both hind
feet taken as a percentage of the stride interval. Further explanation is given in

the text. LF: left fore footfall; LH: left hind footfall; SI: stride interval (duration
of the stride).

Aust. Zool. 19(3), 1978 279

TRICHOSURUS GAIT

RESUEVS

Two gaits were observed to be used by Trichosurus vulpecula on the treadmill.
For speeds less than 5.5 km hr’! a walk was observed, whilst at greater speed a
half-bound was used.

The gait diagram of T. vulpecula walking on the treadmill is presented in
rig. 1. The vertical axis indicates each of the four feet, and the contact of each
foot with the substrate is represented by a horizontal bar. The whole sequence
is “read”, like a printed page, from left to right, and the sequences follow one
ynother from top to bottom. The horizontal dimension represents time, and the
vertical bars indicate the beginning of each stride, the first stride (but not
necessarily any subsequent strides) beginning with the vertical line at the left
of the diagram. Each stride commences with the initiation of substrate contact

by the left hind foot.

The results of the analysis are presented in Fig. 2. Hildebrand (1966) has
shown that the strides of symmetrical gaits may be represented as points in a
two-dimensional space. This space is characterised by two variables, in turn

HALF BOUND yee ee (Oiftrames

FIG. 3.—Gait diagram of JT. vulpecula half-bounding (on a treadmill). Explanation of gait
diagrams is given in caption of Fig. 1 and in text. Gaps, not occupied by any
horizontal lines, indicate periods of suspension referred to in text. A indicates
the first period of flexed suspension (present at the end of each stride), whilst B
indicates one (the only one present in the diagram) of the periods of extended
suspension. Ten frames of the film represent 0.08 seconds. Other abbreviations
as in Fig. 1.

280 Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

55

45

: %S.1.

Midtime lag
w
un

nN
on

5
30 40 50 60
Hind contact [total]: %S.I.

FIG. 4.—Results of analysis, after the method of Hildebrand (1977), of half-bound by
T. vulpecula (on a treadmill). As in Fig. 2, each point represents one stride and
the average is represented by the star. The ordinate represents the midtime lag,
which is the period by which the middle of the period of substrate contact of the
forelimbs follows the middle of the same period of the hind limbs, The abcissa
represents the period during which one or both hind feet are on the substrate
during the stride. Further explanation is given in the text, SI: stride interval.

represented as the two axes of a graph (Fig. 2). Indicated on the ordinate is the
period of time by which each fore footfall follows the corresponding hind footfall
expressed as a percentage of the stride interval; i.e. the duration of the stride.
We found that this period (lag) was not usually equal for both left and right sets
of limbs, and arbitrarily chose to use that of the left set of limbs. The second
variable, indicated on the abcissa, is the period of time for which each foot is in
contact with the substrate, again expressed as a percentage of the stride interval.
In T. vulpecula these periods were not equal for each of the four feet, and we have

Aust. Zool. 19(3), 1978 281

TRICHOSURUS GAIT

arbitrarily chosen to use the average of the two hind contact durations, keeping
in mind that Hildebrand (1977) found this variable of use in analysing asym-
metrical gaits. The inequality between the duration of the left and right substrate
contacts was due both to variations between individuals and between successive
strides of one individual. The right members of a pair of limbs contact the ground
about 50 per cent of the stride after the left footfall and vice versa. Hind contact
is 55 to 70 per cent of the stride interval, and forelimb lag ranges from 40 to 60
per cent.

FIG. 5.—T. vulpecula climbing a flat vertical surface, using the walk.

282: Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

Fig. 3 shows the gait diagram for the half-bound on the treadmill. Unlike
the walk, there is a period, termed a suspension, at the end of each stride, during
which there is no substrate contact. J. vulpecula shows a relatively long period
of flexed suspension (during which the vertebral column is flexed, the forelimbs
retracted and the hind limbs protracted) between loss of forelimb contact with
the ground and hind footfall. On occasion a short period of extended suspension
(during which the vertebral column is extended, the forelimbs protracted and
the hind limbs retracted) was observed prior to fore footfall.

Fig. 4 presents the results of Hildebrand’s (1977) method of analysis of
asymmetrical gaits applied to T. vulpecula. Hildebrand (1977) shows that at least

FIG. 6.—T. vulpecula half-bounding between branches and on to a log. Two successive
strides are shown.

Aust. Zool. 19(3), 1978 283

TRICHOSURUS GAIT

}— ||

BOUND A BOUND B

FIG. 7.—Gait diagram of one stride each of the two types of bounds used by T. vulpecula.
Explanation of gait diagrams is given in caption of Fig. 1 and in text. A period
of suspension may be seen in bound A that is not found in bound B, Abbreviations
as in Fig. 1. Each stride represented is 0.35 seconds in duration.

lateral

US)
i=)

lateral sequence

S
i=]

couplets

oa
t=)

°o
| {
q ] i
: uck: Pos
| 5 rey
i 5
Bay ene Xy

lateral
couplets

OP!S QWRS UO PUY SMO]]OJ 4jej2300; YUOsy 34} BPIsys yo 4,

100,
1

% of stride that each foot is on the ground

FIG. 8.—The symmetrical gait of JT. vulpecula (in the two lightly stippled envelopes)
compared with those of the mammals studied by Hildebrand (1966) (densely
stippled). The descriptive terms labelling the upper and right borders are those
of Hildebrand. It may be seen that 7. vulpecula largely uses a moderate walk
with diagonal couplets. Although points for ZT. vulpecula fall within the area
designated ‘trot’? by Hildebrand, this gait was not observed. Further discussion is
given in the text, (Diagram modified from Hildebrand, 1966).

284 Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

five variables are necessary to characterise asymmetrical gaits. Five variables require
9 graphs for their graphic expression. Rather than work with all 9, Hildebrand
chose to represent points on a graph analogous to that used for the analysis of
symmetrical gaits. On this diagram the ordinate indicates the duration (midtime
lag) by which the middle of the period of substrate contact of the forelimbs
follows the middle of the period of contact of the hindlimbs represented as a
percentage of the stride interval. The abcissa represents the period during which
one or both hind feet: are in contact with the substrate, again expressed as a
percentage of the stride interval.

Three gaits were observed to be used by T. vulpecula in the enclosure. The
walk was frequently used both on the ground and along inclined branches, as
well as in climbing vertical surfaces (Fig. 5). In the latter case, as with movement
on inclined branches, a slow diagonal sequence with two- and three-limb support
was used.

The second gait observed was the half-bound, used often in movement

along horizontal and inclined branches and in jumping from one branch to another
(Fig. 6).

The third gait observed, which was never seen on the treadmill, was the
bound. Simultaneous movement of both fore- and hindlimbs characterises this
gait (Dagg, 1973). Two types of bounds were observed in the enclosure (Fig. 7).
The first type, “A”, agrees with Dagg’s description. The second type, ‘“B”, also
showing simultaneous movement by both pairs of limbs, is slower: two-limb
support alternates with four-limb support. Speed appears to be the only criterion
determining the choice between these two gaits. Bounding was observed mainly
on vertical or near-vertical surfaces.

DISCUSSION

Hildebrand found that the points on his graph (Hildebrand, 1966, Fig. 10)
for all mammalian symmetrical gaits observed fell within a restricted area of the
total graph. Points in proximity indicate similar gaits. Fig. 8 compares walking by
Trichosurus vulpecula with those gaits studied by Hildebrand. T. vulpecula shows
a moderate to fast walk using diagonal couplets, to use the terminology of

Hildebrand.

The portion occupied by gaits of T. vulpecula in Fig. 8 extends into the
region designated by Hildebrand as the trot. It is thus important to point out
that T. vulpecula was never observed to trot. This discrepancy may have arisen
from the modification necessary to accommodate our data to Hildebrand’s method
of analysis. The most important of these modifications (arising from the inequality
of the contact duration for different limbs) is probably the replacement of Hilde-
brand’s variable, ‘per cent of stride that each foot is on the ground”, by our
variable, “per cent of stride of average hind contact”. Gambaryan (1974) avers
that gaits are not discrete, but fade one into another, forming a continuum over

Aust. Zool. 19(3), 1978 285

TRICHOSURUS GAIT
different speeds. These points on our graph fall at the gradation (under his

interpretation) between the trot and the two walks with diagonal couplets, and
provide support for his contention.

100
80

60

% S.1.

lag:

40

Midtime

20

O 20 40 60 80 100 120
Hind support: %SI.

FIG. 9.—The asymmetrical gaits of 7. vulpecula (coarsely stippled envelope) compared
with those of the mammals studied by Hildebrand (1977) (finely stippled).
Hildebrand presents this graph with the origin at upper right, and we have
rotated it to put the origin at lower left as is the convention. The field of the
graph is divided into four quadrants: i, extended suspension; ii, no suspension;
ill, flexed suspension; and iv, flexed and extended suspension. 7. vulpecula uses
gaits with one (flexed) or two periods of suspension per stride. SI: stride interval.
(Diagram modified from Hildebrand, 1977).

Hildebrand found that, as for symmetrical gaits, all observed asymmetrical
gaits fall within a restricted area of the graph (Hildebrand, 1977, Fig. 9).
Hildebrand included asymmetrical gaits of 79 genera of mammals, including five
genera of Marsupialia, one of Chiroptera, two of Insectivora, 13 of Primates, one
of Lagomorpha, 11 of Rodentia, 15 of Carnivora, two of Pinnipedia (which he
considers separate from Carnivora), one of Hyracoidea, three of Perissodactyla
and 25 of Artiodactyla. Fig. 9 compares half-bounding by T. vulpecula on the
treadmill with the gaits studied by Hildebrand. Trichosurus’ half-bound may be
described as a medium speed gait with one (flexed) or two periods of suspension.

286 Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

In the half-bound the hindlimbs strike the ground together, but the forelimbs
do not. The forelimb striking second is the leading forelimb, as it is placed in
front of the other fore-limb. While half-bounding T. vulpecula was able to change
the leading forelimb without breaking the rhythm of movement. This may reduce
fatigue. However, because of this change, use of Hildebrand’s method of analysis
proved difficult. Midtime lag was impossible to calculate, as the delays in contact
by either forelimb were unrelated to loss of contact by the same limb. Variability
was frequently observed between strides of a single individual.

*S.1.

Y
50
i y
7
.
/
30 Y
_
| |
| g
DL
aE TL | o
a= po = i Eo i 10
1

COMBINATION

FIG. 10.—Possible combinations of support as percent of stride interval (S.1.), for the walk
(shaded) and the half-bound (unshaded) of T. vulpecula on the treadmill.

1: single hindlimb; 6: both forelimbs;

2: single forelimb; 7: both hind- and one forelimb;
3: diagonal limbs; 8: both fore- and one hindlimb;
4: ipsilateral limbs; 9: all limbs;

5: both hindlimbs; 10: no limbs.

(i.e. suspension).

Dagg and de Vos (1968a; 1968b) have considered combinations of support
limbs. Histograms (Fig. 10), indicating each possible combination of support as
a percentage of the stride interval, allow direct comparison of the two gaits used
on the treadmill. Diagonal limbs support the body for more than half the stride
interval whilst walking. Support by one (fore) or three (two hind and one fore)

Aust. Zool. 19(3), 1978 287

TRICHOSURUS GAIT

limbs allows transference between both pairs of diagonal limbs. These three
support patterns account for nearly the entire stride during walking.

The walk is a slow gait without suspension and with support usually by
diagonal pairs of limbs. Gray (1944) demonstrated that this pattern is the most
stable for slow-moving tetrapods. The half-bound is a medium to high speed gait
often used to clear obstacles by mammals with short limbs and a flexible vertebral
column. Howell (1944) suggested that it is a more efficient gait than the bound,
in which more energy may be expended in overcoming the deceleration of landing.
In the half-bound (as compared with the walk) stability is compromised for
speed. During approximately 13 per cent of each stride of the half-bound T.
vulpecula is not in contact with the substrate. Such suspension allows the animal
to increase its stride length, although each stride is of shorter duration than
walking (cf. Figs. 1 and 6). Suspension usually precedes landing on both hind-
limbs.

Rarely, the half-bound is modified by the introduction of a short hind lead,
never comprising more than three per cent of the stride interval. The fore lead
is long, to 22 per cent of the stride interval, which equals the duration of support
by both forelimbs. A second modified gait was seen during walking, when the
footfalls became almost simultaneous and the gait approached the trot (as defined
by Dagg, 1973). As stated previously, the trot itself was not observed. Such
examples demonstrate that consideration of any gait as a single entity may be
misleading, and support the contention of Gambaryan (1974) that all gaits
should be considered to form a continum over different speeds. Gaits appeared
more discrete upon the treadmill than in the enclosure, and we suggest that
this arises from the discrete speeds at which the treadmill was run.

Under quasi-natural conditions in the enclosure the presence of four-limb
(and absence of single-limb) contact during the walk indicates that walking is
normally undertaken at a slower speed than on the treadmill (cf. Dagg, 1973).
The diagonal support pattern seen in walking is quite stable (Howell, 1944; Gray,
1944) and may reflect adaptation to locomotion over an irregular substrate.

The half-bound was used in the enclosure for jumping from branch to branch
and for clearing obstacles. Both forelead and the period of flexed suspension are
reduced from those exhibited on the treadmill. Extended suspension was not
observed, although all limbs were extended immediately prior to forelimb contact.
The tail was apparently used as an anchor, being wound around the branch during
jumping, as in Pseudocheirus herbertensis and P. archeri (Breeden and Breeden,
1970),

CONCLUSIONS

In the enclosure, and hence presumably under natural conditions, Trichosurus
vulpecula exhibited three gaits: the walk, half-bound and bound. Only the first
two of these were exhibited upon the treadmill.

288 Aust. Zool. 19(3), 1978

A. J. GOLDFINCH and R. E. MOLNAR

The walk was used to move along horizontal and up inclined surfaces. Half-
bounds were used to jump from branch to branch, to move along ground with
grass and other low cover and to move up strongly inclined surfaces. The bound
was used on steeply inclined and vertical surfaces. T. vulpecula avoided moving
down steeply inclined branches, preferring to use more nearly horizontal branches
or to jump down. Modified gaits were also seen.

Study of animals on a treadmill is useful if considered as an introduction
to some (not all) of the gaits used in the wild. This supports Hildebrand’s (1977)
contention that gaits exhibited on a treadmill may not be characteristic of those
exhibited in the wild.

ACKNOWLEDGEMENTS

Dr R. Baudinette very kindly made available for study films of two possums, Fred and
TV 33, on the treadmill. Likewise Dr Milton Hildebrand generously made available for our
use a manuscript, at the time unpublished, and we. wish to express out gratitude to both.
We would also like to thank Mr M. Oakey for his assistance in filming, the School of
Zoology, University of New South Wales, for the use of their enclosure, and the Department
of Medical Illustration, University of New South Wales, for their aid in preparation of the
illustrations.

REFERENCES

ANDERSON, D. (1937). Studies on the opossum (Trichosurus vulpecula). Austral. Journ.
Exp. Biol. - Med Sci... 15: 17-32.

BREEDEN, S. & K. BREEDEN (1970). Tropical Queensland. Sydney: William Collins Ltd.,
262 pp.

BURKE, J. D. (1954). Blood volume in mammals, Physiol. Zool., 27: 1-21.
DAGG, A. I. (1973). Gaits in mammals. Mammal Rev., 3: 135-154.
DAGG, A. I. (1974). The locomotion of the camel, Journ. Zool., 174: 67-78.

DAGG, A. I. & A. DE VOS (1968a). The walking gaits of some species of Pecora. Journ.
Zool., 155: 103-110.

DAGG, A. I. & A. DE VOS (1968b). Fast gaits of Pecoran species. Journ. Zool., 155: 499-506,

DAWSON, T. J. (1969). Temperature regulation and evaporative water loss in the brush-tail
possum, Trichosurus vulpecula. Comp. Biochem. Physiol., 28: 401-407.

GAMBARYAN, P. P. (1974). How Mammals Run. New York: Halstead Press. 367 pp.

GILMORE, D. P. (1969). Seasonal reproductive periodicity in the male Australian brush-
tailed possum (Trichosurus vulpecula). Journ. Zool., 157: 75-98.

GRAY, J. (1944). Studies in the mechanics of the tetrapod skeleton. Journ. Exp. Biol.,
20: 88-116.

HILDEBRAND, M. (1966). Analysis of symmetrical gaits of tetrapods. Folia Biotheor.,
67 9-22;

HILDEBRAND, N. (1977). Analysis of asymmetrical gaits. Journ, Mammal., 58: 131-156.

HOWELL, A. B. (1944). Speed in Animals. Chicago: University of Chicago Press, 270 pp.

TROUGHTON, E. le G. (1941). Furred Animals of Australia. Sydney: Halstead Press,
375° pp:

TYNDALE-BISCOE., H. (1973). Life of Marsupials. London: Edward Arnold, 254 pp.

289

Preliminary Assessment of Defecation Patterns
for the Eastern Grey Kangaroo (Macropus giganteus)

Gey. ES AILE
Geography Department, University of Queensland, St. Lucia, 4067

As faecal pellets of the eastern grey kangaroo (Macropus giganteus), wallaroo
(M. robustus) and red-necked wallaby (M. rufogriseus) differ in shape and size,
each animal can be identified from its faeces.

Number of pellets per defecation for grey kangaroos was found to vary markedly
between different habitats. It is therefore suggested that individual pellets are
more suitable than pellet groups as the foundation for survey work based on
pellet counts.

A four-day trial monitored daily defecation rate for five kangaroos. Average for
the group was 412 pellets per animal per day (co-efficient of variation 16 percent).

The defecation rate for night was approximately twice that for day periods,
there being a strong association between defecation and feeding activity.

INTRODUCTION

An appraisal has been made of the faecal pellet count technique as a base
for research into the ecology and distribution of the grey kangaroo (Macropus
giganteus) within Queensland. Successful application of pellet counts, particularly
if census work is envisaged, must be based on tedious preliminary investigations.
The primary concern in any feasibility study is the defecation patterns of the
target species. Depending upon the stability of pellet form, defecation rate and
defecation in relation to activity cycles, the animal will vary in its suitability for
study by this method. In accordance with these problems, a programme has been
initiated to monitor defecation patterns of the grey kangaroo. Details are given
of trials conducted with a captive population at Lone Pine Koala Sanctuary,
Brisbane. Free ranging kangaroos were studied in the Durikai State Forest and
an adjoining property “Eddington”, 40 km west of Warwick, southern Queens-

land.

METHODS
SPECIES IDENTIFICATION
The usefulness of pellet surveys revolves around the assumption that an
animal can be accurately identified from its faeces. In most cases pellet shape is

Aust. Zool. 19(3), 1978 291

Goodies nie

the most reliable guide for identification purposes (Riney, 1957). At Durikai,
grey kangaroos live in association with sizable populations of wallaroos (Macropus
robustus) and red-necked wallabies (M. rufogriseus). Comparisons were made
between the faeces of these three species. At Lone Pine pellets were collected
from the enclosures of caged animals while at Durikai; observation of feeding
animals allowed collection of fresh pellets under natural conditions.
DEFECATION RATE

A preliminary trial at Lone Pine into defecation rate for grey kangaroos was
conducted over four consecutive days during June, 1977. Five animals (two adult
males, two adult females and one juvenile male) were placed in a 0.7 ha section
of the grey kangaroo enclosure. The area was well grassed and the animals had
access to supplementary feed. The enclosure was cleared of pellets twice daily.

Results were tabulated as pellet totals and pellet group totals. A group was
classed as any assemblage of one or more pellets judged to be separate from
surrounding pellets on the basis of distance, pellet size, shape and age. No
minimum distance rule was adopted for separating adjacent groups, and the
results probably reflect “‘splitting’ rather than “lumping” of pellet groups (Neff,
1968> 607):

For any pellet survey or assessment of defecation rate the problem of what
is to be counted must be resolved. The choice lies between individual pellets,
pellet groups, presence or absence of pellets and mass of pellets. Research into
kangaroos has made use of presence or absence of pellets (Caughley, 1964a) and
mass of pellets (Warren, 1971). However, from the viewpoint of conducting a
census or extracting maximum quantifiable data from the survey, the use of
individual pellets or pellet groups is more appropriate. The pellet count technique
was developed around deer surveys and has traditionally been based on pellet
group totals (Bennett et al, 1940). For deer, the number of defecations per day
(pellet groups) varies much less than total number of faecal pellets (Riney, 1957;
Smith, 1964). As deer defecate less frequently than kangaroos, but pass a higher
number of faecal pellets per day, it does not necessarily follow that pellet groups
are the most suitable base for kangaroo surveys.

If it can be assumed that grey kangaroos from different areas have similar
defecation rates, the problem is to determine whether pellet group totals or
individual pellet totals best reflect this constancy.

Work with pellet groups from Durikai and Lone Pine suggested that the
average number of pellets per defecation was quite different for the two
populations. Data were therefore recorded to quantify this observation.

For Lone Pine, early morning counts were made of fresh, undisturbed
groups of pellets deposited by 25 grey kangaroos housed in a 1.4 ha enclosure.

292 Aust. Zool. 19(3), 1978

DEFACATION PATTERNS OF THE GREY KANGAROO

A systematic sampling pattern was completed for each count, although no attempt
was made to cover the entire area.

At Durikai, pellet groups judged to be less than one month old were recorded
from 0.001 ha circular plots. These were positioned at regular intervals along
randomly located transects straddling varying habitat types. Twenty-eight days
prior to this survey 120 fresh pellets from Lone Pine and Durikai were placed
on 60 permanent plots (three rows of 20 pegs). This provided a control group
of pellets of known age for use in the survey. As less than one per cent of the
control group (one pellet from 120) disappeared during the 28 days, results of
the survey were not biased significantly through pellet losses.

DEFECATION AND DaILy ACTIVITY

For the trial study of defecation rates the enclosure was cleared twice daily
to correspond roughly with the late afternoon to early morning feeding session
(4 p.m. to 8 a.m.) and the day resting session (8 a.m. to 4 p.m.). A separate
defecation rate for these two periods could therefore be calculated. Classification
of these activity periods was based on published material dealing with behavioural

GREY KANGAROO WALLAROO

° 10 20 30 40 50 60 70
‘rt — rt

MILLIMETRES

Fig.|
Comparative shape and size
of faecal pellets of the grey

kangaroo, wallaroo, and red
-necked wallaby.

RED-NECKED WALLABY

Aust. Zool. 19(3), 1978 293

Giov.9E. JAE

patterns of grey kangaroos (Caughley, 1964b & Kaufman, 1975) and on personal
observation.

RESULTS
SPECIES IDENTIFICATION

As reported by Caughley (1964a) and Grant (1974), grey kangaroo pellets
have a distinctive shape. Opposite sides of the pellet are approximately equal in
size and are usually rectangular in shape. Pellets often resemble rectangular prisms
with rounded edges (Fig. 1). During the defecation process pellets are normally
passed separately and pellet groups are therefore a collection of individual pellets.

The faeces of the wallaroo are of comparable size to those of the grey
kangaroo. However, the pellets are more elongated and flattened and show a
tendency to taper towards one end (Fig. 1). Unlike the grey kangaroo, wallaroos
may pass a string of pellets compressed together.

Pellets of the red-necked wallaby are generally smaller than those of the
two larger macropods. The shape is usually rounded, although tapered pellets are
not uncommon (Fig. 1).

In Fig. 1 pellets were selected to illustrate the range that commonly occurs
in pellet shape for each species. While variations in shape, that could result in
wrong identification, have been noted, errors of this nature are not likely to be
significant. Most variations of this type occurred with the Lone Pine animals,
and these were attributed to reliance on a concentrate diet. This has been observed
to cause similar problems in other herbivores (Smith, 1964).

Several weeks of sampling work at Durikai have confirmed that recognition
of pellets from each species can be achieved quickly and confidently in the field.
DEFECATION RATE

TABLE 1
Daily defecation rate for five kangaroos over a four day period.
Co-efficient
Range Mean of variation
Pellets per animal 319-465 412.2 16%
Groups per animal 53-79 68.4 16.5%

Although the trial was too short to be given great significance, it does provide
a rough guide line for comparing census estimates from pellet surveys with other
survey methods. The conversion figure allows unwieldy population estimates of
pellet totals to be reduced to a more easily interpreted form.

Monitoring of defecation rate for the Lone Pine kangaroos suggested that
for this population individual pellet totals and pellet group totals were equally
suited as a survey base. Co-efficients of variation for the two are similar (Table 1).

294 Aust. Zool. 19(3), 1978

DEFACATION PATTERNS OF THE GREY KANGAROO

However, comparison of the frequency distributions of pellets per group for
captive and wild kangaroos (Fig. 2a & b) indicates that other factors must be
taken into consideration. Average number of pellets per group (6.2 and 3.7
respectively ) and the skewness of the two distributions differ markedly (significant
difference, 0.1 per cent level, Kolmogorov-Smirnov two-tailed test).

Unless the Lone Pine kangaroos pass approximately twice as many faecal
pellets per day as the wild kangaroos, pellet group totals cannot be a reliable
indicator of defecation rate for grey kangaroos.

Observation of feeding kangaroos implies that the number of pellets deposited
in any one group depends on animal movement patterns.

For the captive kangaroos in a confined space, and with ample food, move-
ment was minimised and the number of pellets passed at one situation large. At

20

Isp (a) DURIKAI (wild) 300
(n = 1912)

200

Percentage of total no. of groups
fo)
No. of groups

keane OS HBS er = : ia: S
r 5 10 15 20 25 °
Pellets per group

300

(b) LONE PINE (captive)
(n=1749)

8
°
No. of groups

100

Percentage of total no. of groups

Pellets per group

Fig.2. Distribution of pellets per group.

Aust. Zool. 19(3), 1978 295

Gi Ey EE

Durikai, on the other hand, where pasture conditions were variable, feeding
animals moved more often than their Lone Pine counterparts. Here defecation
was characterised by fewer pellets per group.

During feeding kangaroos defecate frequently, both while stationary and
when moving. For any short time-span a certain number of pellets will be
deposited. Whether these form one large group or several small groups depends
largely on the density of the pasture being utilised. This hypothesis is supported
by comparison of the frequency distributions of pellet groups collected from well-
grassed and sparsely-grassed plots located within the one paddock on Eddington
Station at Durikai (Fig. 3a & b).

25

alae (a) WELL GRASSED
: PLOTS

(n=585)

fe)
°

Percentage of total no. of groups
No. of groups

Pellets per group

(b) SPARSELY GRASSED
PLOTS

(n=276)

Percentage of total no. of groups
No. of groups

1 a0 15 20 25 30
Pellets per group

Fig. 3. Distribution of pellets per group at Eddington.

296 Aust. Zool. 19(3), 1978

DEFACATION PATTERNS OF THE GREY KANGAROO

The two distributions are significantly different (0.1 per cent level,
Kolmogorov-Smirnov two-tailed test). Where food is plentiful, the number of
pellets per group is higher, as kangaroos stay in the one position for longer.

Comparisons between separate surveys in the spatial or temporal context could
lead to serious interpretation errors if pellet groups are used. For example, should
relative abundance of two kangaroo populations be compared by a pellet group
survey, estimates will be positively biased toward the population utilising the
more widely dispersed forage resource.

The pellet survey for Durikai described earlier in this paper provides data
to illustrate the point. During this survey a 90 km? block of forested country had
been sampled by 515 circular plots of area 0.001 ha. Each plot was searched for
pellets judged to be less than four weeks of age. Averages were 1.22 pellets
per plot with 0.3 pellet groups per plot. These estimates were converted to
kangaroos/km? using the formula:

y x 100,000
D Saree tea ar ah De
Poa GIB &
where D == kangaroos/km2
y = mean per plot
(pellets or pellet groups)
100,000 = number of 0.001 ha plots/km2
28 = survey period in days
r = daily defecation rate

(412 pellets or 68 pellet groups)

Result using pellet totals was 10.6 kangaroos/km?, while pellet groups
returned an estimate of 15.8 kangaroos/km*. Although the accuracy of these
estimates cannot be checked on the wild population, a density of 15.8
kangaroos/km? does appear very high for the country involved. On the other hand,
10.6 kangaroos/km? corresponds reasonably well with the density estimates of
Fox for similar habitats (AREA, 19) close by across the New South Wales border
(Pox; P9742 997):

As a common yardstick for comparing different populations, pellet totals
appear to be the most suitable base for surveys.

DEFECATION AND DaILy ACTIVITY

Any use of pellet surveys as a guide to habitat usage must be finely attuned
to known behaviour of the animal involved. Correlations between pellet density
and habitat type cannot take for granted that density of pellets is related to time
spent in a particular area or is indicative of animal preferences.

Caughley (1964a: 242) states: “As most faeces are dropped at night when
the animals are feeding, the dispersion of faeces gives an indication of how
kangaroos utilise their habitat over a period of time’.

Aust. Zool. 19(3), 1978 297

Gi 2E. GHIEE

Observations at Lone Pine and Durikai both substantiate Caughley’s findings,
although several points need to be clarified. For the Lone Pine study on defecation
rates, pellet production per hour for the night period (4 p.m. to 8 a.m.) was
higher than that of the day period.

TABLE 2
DEFECATION RATES FOR DAY AND NIGHT
Time Period Weather Pellets per Roo per Hr.
Tuesday fine 112
Tuesday night fine DAEG
‘Wednesday fine yl
Wednesday night fine DOT
Thursday overcast 2,
Thursday night rain 13.0
Friday clearing 13.9

For the last day rain appears to have reduced overall defecation rate and
evened out the differences. A similar phenomenon for rabbits is described by
Taylor et al (1956).

When examining the data presented in table 2 it should be noted that these
virtually domesticated animals actively foraged in their preferred feeding areas
sporadically during the day. Wild kangaroos rarely follow this behaviour pattern.
The major portion of daylight hours is spent in habitat types that offer conceal-
ment but little in the way of forage. In this case then, the data presented ue
overestimate the pellet contribution for day periods.

One point gleaned from field work at Durikai is that heavy defecation is not
usually associated with the daily bedding sites in the scrub. Resting kangaroos at
Lone Pine have only been observed to defecate when they get up to shift position.

These observations and the data from table 2 indicate that defecation is
strongly associated with feeding. This activity is most common during the period
from late afternoon to early morning. In this context it must also be stated that
pellet counts will not necessarily offer an indication of overall habitat usage by grey
kangaroos. Day time shelter zones for example may exhibit very low pellet densities
unless good ground cover is present that will keep kangaroos in the area during the
feeding cycle.

DISCUSSION
Prolonged monitoring of average defecation rates is necessary before accurate
population estimates can be made through pellet counts. However, this does not
detract from the value of the technique in providing indices of kangaroo abundance.
Greatest potential exists for areas where other survey methods prove unsatisfactory.

298 Aust. Zool. 19(3), 1978

DEFACATION PATTERNS OF THE GREY KANGAROO

Heavily timbered country can be placed in this category. As only a small propor-
tion of the kangaroo population is visible to observers, census estimates are not
accurate. For habitats of this type, the reliability of aerial or ground counts of
kangaroos is also doubtful because of seasonal movement patterns of the animals.
To test this assumption an aerial survey programme has begun at Durikai to
compare successive kangaroo counts with seasonal changes in pellet density distri-
butions.

Pellet counts also offer considerable scope in providing distributional arr aE
ecological and management significance. Habitat preference and seasonal movement
pattern studies appear well suited in this regard. These factors are rele-
vant in appraising issues such as competition with domestic stock, seasonal
vulnerability to hunting and likely impact of land management programmes.

A factor favouring the use of pellet counts for research into grey kangaroos
is the high defecation rate in terms of groups passed per day. Faecal material is
therefore distributed widely across usage zones. In areas with a moderate density
of animals, counts based on short deposition periods should return pellet densities
high enough for surveys to be completed with success. Success can be defined in
terms of workload required for acceptable statistical manipulation of data.

Defecation patterns of the target animal are only one of several aspects that
must be examined when assessing the usefulness or feasibility of pellet counts.
Determining pellet age, pellet decay rates and pellet visibility poses its own set
of problems, as does the design of the survey itself. Success or failure however
relies in the main on defecation patterns. The study outlined in this paper suggests
that the defecation patterns of the grey kangaroo make it a suitable subject for
research by the pellet count method.

ACKNOWLEDGEMENTS
The management of Lone Pine Koala Sanctuary, Brisbane has allowed access to their

kangaroos and wallabies over_an extended period of time.

Permission to Traverse state forestry reserves was granted by the Queensland Forestry
Department and Mr Tom Thornton has permitted research to be carried out on his property
Eddington.

Without the co-operation of the above the project would not have been possible.

Mr E. E. Savage produced the graphs and Mr H. Stewart-Kellick sketched the faecal
pellets.

A commonwealth government, post-graduate research award cibeoried this project.

REFERENCES

BENNETT, L. =., ENGLISH. P. F. & McCAIN. R. (1940). “A Study of Deer Populations
by use of Pellet Group Counts”. J. Wildl. Magmt. 4(4): 398-403.

CAUGHLEY, G. J. (1964a). “Density and Dispersion of Two Species of Kangaroo in
Relation to Habitat”. Aust. J. Zool. 12: 239-249.

CAUGHLEY. G. J. (1964b). “Social Organisation and Daily Activity of the Red Kangaroo
and the Grey Kangaroo”. J. Mammal, 45(3): 429-436.

Aust. Zool. 19(3), 1978 299

God. EAE

FOX, A. M. (1974). Kangaroos in N.S.W., N.S.W., N.P.W.S., Sydney.

GRANT, T. R. (1974). “The Shapes of Faecal Pellets of Red and Grey Kangaroos”. Aust.
Mammal. 1(2): 261-263.

KAUFMAN, J. H. (1975). “Field Observations of the Social Behaviour of the Eastern Grey
Kangaroo, Macropus giganteus”. Anim. Behav. 23(1): 214-221.

NEFF, D. J. (1968). “The Pellet-Group Count Technique for Big Game Trend, Census and
Distribution: A Review”. J. Wildl. Magmt. 32(3): 597-614.

SMITH, A. D. (1964). “Defecation Rates of Mule Deer”. J. Wildl. Magmt. 28(3): 435-444.

TAYLOR, R. H. & WILLIAMS, R. M. (1956). “The Use of Pellet Counts for Estimating

the Density of Populations of the Wild Rabbit, Oryctolagus cuniculus (L)”. N.Z. J. Sci.
Technol. B. 38: 236-256.

WARREN, J. F. (1971). “Estimating Grazing Trends in Western N.S.W. by Dung Sampling”.
J. Soil Cons. N.S.W. 27(3): 182-186.

300 Aust. Zool. 19(3), 1978

Eye Closure During Agonistic Behaviour
of Rattus villosissimus

ROBERT J. BEGG*

* Department of Zoology, Monash University, Clayton, Victoria, Australia.

Present Address: Territory Parks and Wildlife COTMSSISES P.O. Box 38496,
Winnellie, N.T. 5789

Several groups of workers have noted the eye closure that may occur in a
rat involved in certain acts or postures of flight or defence (Chance, 1962; Grant
and Mackintosh, 1963; Barnett and Evans, 1965). Various hypotheses have been
advanced to explain the significance of this response.

Chance (1962) proposed that closing of the eyes, turning away of the head,
and similar responses when they occur in an agonistic context act as ‘“‘cut-off”
postures. They result in a reduced stimulus input from an attacking opponent and
reduce the drive for flight in the actor. Thus, the rat under attack may remain in
the proximity of its opponent. According to Chance, the advantage to the rat under
attack is inherent in the idea that the formation of dominance-subordination rela-
tionships involves a fluctuating balance of attack and defence. Then, ‘‘cut-off”’
postures would allow the lability necessary for a fleeing rat to return to social
contact.

Grant and Mackintosh (1963) working on a series of rodents including
laboratory rats and mice, stated that “‘one of the general features of defensive and
submissive postures is the withdrawal of attention from the aggressive animal”’.
This they claim, reduces the tendency for the defender to retreat, thus concurring
with the findings of Chance (1962). Barnett and Evans (1965) add the possibility
that eye closure in agonistic situations, may be a product of autonomic activation.
Secondarily, this response may change “the visual stimulus provided by the
defender, and so reduces the likelihood of further assault”. Thus, closing of the
eyes may affect the subsequent behaviour of both the actor and reactor.

During a study of the agonistic behaviour of Rattus villosissimus (Begg and
Nelson, 1977), rats were found to close one or both eyes partly or wholly, during
many different acts or postures observed in agonistic situations. It occurred spon-
taneously (with no obvious preceding change in the behaviour of an opponent),
or as a result of some direct action by another rat. All situations in which R.
villosissimus were seen to display this response, are listed in Table 1. A full

Aust. Zool. 19(3), 1978 301

Ry J. BEGG

FIG. 1.—Variability of eye closure in one bout of OFFENSIVE (right) and DEFENSIVE
(left) UPRIGHT posture in Rattus villosissimus.

302 Aust. Zool. 19(3), 1978

EYE CLOSURE IN RATTUS VILLOSISSIMUS

description of the behaviour involved is given in Begg and Nelson (1977). No
measures of frequency are given as it is hard to quantify this response due to rapid
movements, variable orientation of rats involved, and differences in degree of eye
closure.

TABLE 1
Postures and Acts Involving Eye Closure in Rattus villosissimus

Behaviour of Behaviour of Rat with Eye,

Dominant Rat Subordinate Rat or Eyes, Closed
LUNGING CROUCHING D
APPROACH LUNGING . D and/or S
CROUCHING during a lull LUNGING S
in a ROLLING WRESTLING
FIGHT
CROUCHING during a lull ESCAPE LEAP. D
in a ROLLING WRESTLING
FIGHT

ATTACK LEAP
(a) as D makes contact

with mouth and forelimbs DEFENSIVE SIDEWAYS POSTURE D
(b) D releases grip with

teeth and BITES again DEFENSIVE SIDEWAYS POSTURE D
OFFENSIVE UPRIGHT POSTURE DEFENSIVE UPRIGHT POSTURE D and/or §
OFFENSIVE SIDEWAYS DEFENSIVE SIDEWAYS D and/or §
POSTURE POSTURE
STANDING OVER “SUBMISSIVE” POSTURE D. of.S
STANDING OVER SUBMISSIVE CROUCH D.or S$
CROUCHING SUBMISSIVE CROUCH S
GROOMING SELF SUBMISSIVE CROUCH S
SUBMISSIVE CROUCH SUBMISSIVE CROUCH D
INVESTIGATE CROUCHING S
GROOMING SELF CROUCHING S
CROUCHING GROOMING D D

D = dominant rat S = subordinate rat

Eye closure occurred during acts and postures associated with offence or
defence. It also occurred simultaneously in both attacking and defending rats (Fig.
1 shows a single instance of OFFENSIVE and DEFENSIVE UPRIGHT postures
where eye closure occurred first in neither rat, then in the attacker, then in both
rats). During postures which were maintained for several seconds or more, often
only the eye nearer the opponent was closed. However rats closed both eyes while
carrying out rapid acts (such as ATTACK LEAP, LUNGING, BITING), or fol-
lowing the performance of such rapid acts by an opponent. This was observed
frequently when an animal made an ATTACK LEAP. Just before making contact
with its forelimbs and mouth on an opponent, the attacking rat often briefly closed
both eyes. Some rats have also been observed, during an ATTACK LEAP, to release

Aust. Zool. 19(3), 1978 303

R. J. BEGG

a grip with their teeth, and take a second bite further over the back of the defender,
closing their eyes as they do so. In situations such as this, eye closure does not
result in an obvious reduction of an attack drive as a theory such as that of Chance
(1962) would propose. The animals still press home the attack.

Situations involving eye closure do however have some features in common.
Most importantly they almost always involve some tactile contact between rats.
Thus the eyes of each rat, which are particularly vulnerable due to their large size
and degree of protrusion from the orbit, are placed in a situation of potential
danger from the nearness of the opponent‘s teeth. Thus, in the situations outlined
above, eye closure (which does, as detailed, often only involve the eye nearer to
the opponent) may serve to protect the eyes from possible damage by an opponent.
Vine (1970) suggests that a similar mechanism operates in many gulls and terns
where the head-turning that occurs in many agonistic encounters may be a defensive
reaction, removing vulnerable aspects of the face from the proximity of an
opponent’s bill.

Eye closure during sudden movement by either the actor or reactor then
probably serves as a protective mechanism. Landis and Hunt (1936) demonstrated
that a sudden loud noise may produce eye closure in man. Andrew (1963) lists
a series of “‘protective responses” found in mammals, one of which is closing of
the eyes. These responses may be evoked by an intense stimulus contrast, such as
a sudden loud noise or a blow towards the face. They serve to protect the major
sense organs against possible noxious effects from the source of contrast.

Thus, rather than forming some sort of “‘cut-off” mechanism, in R. villosissimus
at least, eye closure in agonistic situations is probably a defensive reaction, evoked
by the proximity of an opponent, or sudden movement by an opponent, or during
sudden movement towards an opponent.

REFERENCES

ANDREW, R. J. (1963). The origin and evolution of the calls and facial expressions of
the primates. Behaviour 20, 1-109.

BARNETT, S. A. and EVANS, C. S. (1965). Questions on the social dynamics of rodents.
Symp. Zool. Soc. Lond. 14, 233-248.

BEGG, R. J. and NELSON, J. E. (1977). The agonistic behaviour of Rattus villosissimus.
Aust. J. Zool. 25(3), 291-327.

CHANCE, M. R. A. (1962). An interpretation of some agonistic postures; the role of “cut-
off” acts and postures. Symp. zool. Soc. Lond. 8, 71-89.

GRANT, E. C. and MACKINTOSH, J. H. (1963). A comparison of the social postures of
some common laboratory rodents, Behaviour 2/, 246-259.

LANDIS, E. and HUNT, W. A. (1936). Studies of the startle pattern: LHI. Facial Pattern.
J. Psychol. 2, 215-219.

VINE, I. (1970). Communication by facial-visual signals. In “Social Behaviour in Birds
and Mammals”. Ed. J. H. Crook, pp. 279-354. (Academic Press: London),

304 Aust. Zool. 19(3), 1978

A Triploid Male Individual Amphibolurus nobbi nobbi
(Witten (Lacertilia: Agamidae)

GEOFFREY J. WITTEN
Department of Anatomy,
University of Sydney,
Sydney, 2006.

INTRODUCTION

Triploidy has been observed in representatives of three lizard families ( Aga-
midae, Geckonidae, Teiidae; see Gorman, 1973, for review). Most of these are
parthenogenetic species (Hall, 1970; Lowe et al, 1970b). The only previous
report of triploidy within the Agamidae is an apparently parthenogenetic popula-
tion of Leiolepis belliana (Hall, 1970).

MATERIALS AND METHODS

Live males of Amphibolurus nobbi in breeding condition were injected intra-
peritoneally with 0.2% solution of colchicine at the rate of 0.05 mls/g body
weight within 2-3 days of collection. After 5 hours at room temperature (24-28°C)
the animal was sacrificed. A testis was chopped finely, macerated in 1% sodium
citrate solution for 10 minutes, then centrifuged. The cell button was fixed in
three changes of freshly prepared acetic acid: methanol (1:3), and resuspended
after the third change of fixative. Three drops of the suspension were spread on a
slide, ignited, allowed to burn out, and the residue flung off. The preparation was
then stained for 15-20 minutes in Giemza and air-dried before mounting.

RESUETS

Of three Amphibolurus nobbi karyotyped, one was triploid. The most com-
mon karyotype for Australian agamids consisting of 12 macrochromosomes and 20
microchromosomes (2n = 32) (Witten, unpubl.) was recorded in the other two
individuals. A comparison with a normal diploid figure demonstrates that triploidy
has resulted from an extra haploid complement, with no observable difference in
any of the macrochromosomes (Figs. 1, 2). The triploid specimen (Austr. Mus.
Field tag 11392) appeared to be normal.

Aust. Zool. 19(3), 1978 305

G. J. WITTEN

Bis. t

Fig. 3 Fig. 4

FIG, 1 — Cut-out representation of karyotype of diploid (left) and triploid A. nobbi.
FIG. 2 — Photomicrograph of triploid mitotic figure. Bar represents 10u.
FIG. 3 — Diakinesis of triploid individual. Most figures of this stage showed even less

organisation. ; Fie :
FIG. 4 — Metaphase II of meiosis in triploid A. nobbi. c.f. Fig 2 for appearance of

chromosomes.

306 Aust. Zool. 19(3), 1978

A TRIPLOID INDIVIDUAL AGAMID

_ It was apparent that diakinesis did not always result in the organisation of
chromosomes into triplets (Fig. 3). Many cells representing this stage were
observed, but no consistent pattern emerged.

Cells representing the second meiotic metaphase are distinguishable in normal
testis preparations by the thinner appearance of the arms, the non-alignment of
these arms away from the centromere, and by their haploid number. Using the first
two of these criteria, second meiotic metaphase cells were identified in the triploid
individual, but all such cells had a triploid complement (Fig 4). From this observ-
ation it was assumed that no reduction of chromosome number cccurred in meiosis

i

DISCUSSION

Triploidy has been associated with parthenogenesis in all three families where
it has been reported (Teiidae; Pennock, 1965; Geckonidae; Kluge and Eckardt,
1969: Agamidae; Hall, 1970). In the teiid genus Cuemidophorus, Lowe et al
(1970b) have suggested that the triploid condition in at least some parthenogenetic
species has arisen through hybridisation between diploid males and females of
diploid parthenogenetic species. It is improbable that the present example of tri-
ploidy is due to hybridisation: the animal in question is sympatric with only two
other agamid species. One is Lophognathus gilberti, which has a distinctively
different karyotype (Witten, unpubl.). The other is Amzphibolurus barbatus which
is about 50 times the weight of A. zobbi and therefore is unlikely to produce a
hybrid under natural conditions.

The presumed failure of meiosis I in this study is in contrast with the con-
clusions but not the results of Lowe e¢ al (1970a). In an allotetraploid hybrid they
noted an “infrequency of cells exhibiting meiosis II”, and concluded that this was
due to a “delayed testicular cycle’. It is interesting to speculate that the failure to
divide in meiosis I observed in this study could be due to polyploidy. If this is the
case, then ‘accidental’ triploid females would produce triploid gametes. It has long
been established in the amphibia that haploid individuals are less viable than
diploids or polyploids (Fankhauser, 1945). If this holds true for reptiles, then
gynogenetically or parthenogenetically developing triploid ova are more likely to be
viable than haploid ova, and the strong correlation between triploidy and partheno-
genesis could in part be explained.

ACKNOWLEDGEMENTS
I would like to thank Dr L. A. Moffat for her criticism and suggestions.

REFERENCES

FANKHAUSER, G. (1945). The effects of changes in chromosome number on amphibian
development. Q, Rev. Biol. 20: 20-78.

Aust. Zool. 19(3), 1978 307

G. J. WITTEN

GORMAN, G. C. (1973). The chromosomes of the reptilia, a cytotaxonomic interpretation.
pp. 349-424. In A. B. Chiarelli & E. Capanna (eds.) Cytotaxonomy and Vertebrate
Evolution. Academic Press, London.

HALL, W. P. (1970). Three probable cases of parthenogenesis in reptiles (Agamidae,
Chamaeleontidae, Gekkonidae).

Experientia 26: 1271-1273.

KLUGE, A. G. and M. J. ECKARDT (1969). Hemidactylus garnotii Dumeril and Bibron,
a triploid all-female species of gekkonid lizard. Copeia 1969: 651-664.

LOWE, C. H., J. W. WRIGHT, C, J. COLE and R. L. BEZY (1970a). Natural hybridiza-
tion between the teiid lizards Cnemidophorus sonorae (parthenogenetic and Cnemido-
phorus tigris (bisexual). Syst. Zool. 19: 114-127.

—_____________ (1970b). Chromosomes and evolution of the species groups
of Cnemidophorus (Reptilia: Teiidae). Syst. Zool. 19: 128-141.

PENNOCK, L. A. (1965). Triploidy in parthenogenetic species of the teiid lizard, genus
Cnemidophorus, Science N.Y. 149: 539-540.

308 Aust. Zool. 19(3), 1978

Sexual Dimorphism in an Australian Galaxiid
(Pisces: Galaxiidae)

R. M. McDOWALL
Fisheries Research Division, Ministry of Agriculture and Fisheries, Private Bag,
Christchurch, New Zealand
(Fisheries Research Publication No. 328)

ABSTRACT

Sexual dimorphism in the diminutive Australian galaxiid Galaxiella pusilla (Mack)
is described; this is the only galaxiid in which sexual dimorphism has been
reported; its significance is discussed.

INTRODUCTION

The family Galaxiidae contains about 34 species, and is widely distributed in
the southern cold temperate zone — Australia, New Zealand, South America, South
Africa, and smaller islands near these areas. The species have been described in
considerable detail (McDowall, 1968, 1970, 1971, 1973a; McDowall and Franken-
berg, In Prep.), but in no instance was sexual dimorphism described. Massola
(1938), however, in a popular article for aquarists, described the colouration of
the tiny Australian species Galaxiella pusilla (Mack) (Fig. 1). He noted that there
is a bright orange longitudinal stripe in males but not in females, a colour differ-
ence that has eluded recognition by later workers. Scott (1971) and Andrews
(1976) described colouration in G. pusilla in considerable detail but made no
mention of sexual dimorphism; it was not recognised by McDowall (1973b), nor
by McDowall (1978) when describing the genus Galaxiella to contain G. pusilla
(Mack), and two similar and related species from Western Australia — G. nigros-
triata (Shipway) and G. munda McDowall.

The observation of live material, by the author, at the Arthur Rylah Institute
for Environmental Research, Melbourne, and the collection of a substantial further
sample of live material, showed that sexual dimorphism does occur in G. pusilla.

MATERIAL EXAMINED

Description of dimorphism in G. pusilla is based primarily on a sample of 43
specimens (22 males, 21 females) collected from a small, overgrown farm drain,

Aust. Zool. 19(3), 1978 309

R. M. McDOWALL

FIG. 1 — Galaxiella pusilla (Mack), 28.5 mm T.L., Narracan Creek, Victoria; female.

flowing intermittently, and entering Narracan Creek, a tributary of the La Trobe
River, near Yallourn, southern Victoria (30 August, 1977); the sample is in the
author’s collection of galaxiid fishes.

A series of 26 body measurements was taken, and counts of the number of fin
rays, gill rakers and vertebrae, following techniques described elsewhere (Mc-
Dowall, 1970; McDowall and Frankenberg, In Prep.). Scanning of data from five
specimens of each sex suggested that there are differences in three body proportions
— body depth, pectoral-pelvic length, and pelvic-anal length; these dimensions were
measured for the entire sample.

FIG. 2 — Galaxiella pusilla (Mack), Narracan Creek, Victoria; above — female 36mm
T.L.; below — male 25 mm T.L.

310 Aust. Zool. 19(3), 1978

SEXUAL DIMORPHISM IN A GALAXIID

RESULTS

DIMORPHIC CHARACTERS
1. Size: Males are much smaller than females (Table 1; Fig. 2).

2. Body proportions: Ripe males are more slender-bodied than ripe females (Table
1). This is primarily a reflection of the swollen abdomens of the ripe females and
the difference has little significance. The pectoral-pelvic and pelvic-anal intervals are
less in males than in females (Table 1). Examination of plots of pectoral-pelvic
length/standard length and pelvic-anal length/standard length suggested that there
is allometric growth in this fish; the higher ratios in females are produced by
allometry common to the two sexes, but which continues further in females than
in males because the females grow to a greater size (Fig. 3).

TABLE 1

Sexual dimorphism in size and body proportions in Galaxiella pusilla (22 males, 21 females,
Narracan Creek, Victoria).

Range Mean S.D.

Size (total length — mm) male 24.5 - 29.5 DENS 1.24
female 31.0 - 39.0 34.20 1.86

Body depth/standard length (%) male 16.8 - 21.5 18.70 127
female 19.0 - 23.3 21.09 0.99

Pectoral-pelvic length/standard male 23.7 = 2931 26.06 1.44
length (%) female 26.7 - 34.7 30.79 1.87
Pelvic-anal length/standard length (%) male 15.9 = 19.3 17.40 0.83
female 17.0 - 22.6 19.00 1.74

3. Ventral keel and genital papilla: In both sexes there is a fleshy keel along the
ventral abdomen, beginning at about the pelvic fin bases, and extending back as far
as the vent. The relative depth of the keel is slightly greater in males than in
females. In the male the vent is followed by a low, fleshy genital papilla that pro-
jects slightly more than the vent; in the female the genital papilla is a distinct
fleshy mound.

4. Colour: When alive the male G. pusilla is a dull olive-amber on the back and
upper sides, with three black, longitudinal stripes along the trunk (Fig. 1); the
upper stripe is weakest and merges medio-dorsally with darker pigment on the back.
The middle stripe is much thinner and very distinct, and is positioned at about
the lateral line; it is above and well-separated from a characteristic and very bright
orange stripe along the abdomen, but converges with it on the caudal peduncle. The
lower dark stripe is strongly developed and forms the lower margin of the orange
stripe. Ventrally there is a pair of black lines, originating together in the isthmus,
diverging, and then running parallel along the belly to the pelvic fin bases; the
pelvic-anal interval is more or less unpigmented but there are black lines along the

_ Aust. Zool. 19(3), 1978 317

R. M. McDOWALL

e e
6

e
bd Females

Pelvic-anal length (mm)

_
On

Females

Pectoral-pelvic length (mm)

20 25 30 35
Total length (mm)

40

FIG. 3 — Relationship between pelvic-anal length and pectoral-pelvic length, and total
length in Galaxiella pusilla.

anal fin base and the ventral margin of the caudal peduncle, becoming more or less
confluent with the lowermost dark lateral stripe. The belly is silvery-white, and the

eyes are silvery-gold, with small areas of bright orange as a continuation of the
lateral orange stripe.

The female differs from the male primarily in lacking the bright orange stripe.
The longitudinal dark lines occur, but these are a little less bold, and there is a

lateral band of silvery iridescence comparable in position to the orange stripe in the
male.

312 Aust. Zool. 19(3), 1978

SEXUAL DIMORPHISM IN A GALAXIID

DISCUSSION

Little is known about reproductive patterns in galaxiid fishes, particularly
about their spawning behaviour. Breeding has been reported in a few galaxiids —
Galaxias olidus Ginther (Walford, 1940 — as G. coxit Macleay); G. divergens
Stokell (Hopkins, 1971); G. vulgaris Stokell (Benzie, 1968; Cadwallader, 1976);
Neochanna apoda Ginther (Eldon, 1971, 1978); N. burrowsius (Phillipps) (Cad-
wallader, 1975). However apart from the report by Massola (1938) of spawning
in Galaxiella pusilla, only in Galaxias maculatus (Jenyns) has spawning actually
been observed (Hayes, in Hefford 1931a, b, 1932; Pollard, 1971) — small to large
shoals of fish were observed to spawn together, releasing the eggs haphazardly
over the substrate. In none of these species has sexual dimorphism, like that des-

cribed for G. pusilla, been observed.

Galaxias maculatus is one of very few species of galaxiid that is known to live
in shoals as an adult, and it is likely that its spawning habits are distinctive in the
family. Galaxiella pusilla was reported by Massola (1938) to spawn in pairs, the
female depositing its eggs singly on aquatic vegetation and the male subsequently
fertilising them. Massola’s report agrees with more recent observations of spawning
in G. pusilla by R. W. Vanner and G. D. Backhouse, of Victoria, Australia, both
of whom have seen the fish spawning in indoor aquaria (pers. comm.). Unlike
most galaxiid fish, G. pusilla is a mid-water, free-swimming species. This be-
havioural distinction and the formation of pairs for spawning seem likely to be
related to the occurrence of sexual dimorphism.

G. pusilla has recently (McDowall, 1978) been placed, with G. migrostriata
(Shipway) and G. munda McDowall, in the genus Galaxiella on account of
similarities between these obviously closely related species, and differences from
the remainder of the family Galaxiidae, in particular differences from the genera
Galaxias Cuvier and Brachygalaxias Eigenmann.

There are not yet sufficient data available to indicate whether dimorphism
occurs in the two Western Australian species of Galaxiella. G. nigrostriata very
closely resembles G. pusilla (it has been regarded as a subspecies by some workers
—Shipway, 1953; Munro, 1957), and sexual dimorphism seems likely in this
species. G. munda less resembles the other Galaxiella species; it lacks the brilliant
orange stripe and has instead a dull brick-orange stripe. Colour photographs of
some specimens of G. munda show silvery iridescence along the lower sides,
_ perhaps indicating that these are females comparable with the females of G. pusilla.

If the two Western Australian species are sexually dimorphic, the occurrence
of such dimorphism in this small, distinctive radiation of galaxiid fishes would add
to the array of characters setting Galaxiella apart from other galaxiid fishes, and
thus add weight to the recognition of the genus as distinct.

ACKNOWLEDGEMENTS

I am particularly grateful to Mr G. N. Backhouse, Arthur Rylah Institute for Environ-
mental Research, Heidelberg, wagons: for his assistance in collecting the fish, and to Mr

Aust. Zool. 19(3), 1978 313

R. M. McDOWALL

J. F. C. Wharton, Director, Fisheries and Wildlife Division, Ministry for Conservation,
Victoria, for making the collections possible. Financial support was received from the Aus-
tralian Biological Resources Study.

REFERENCES

ANDREWS, A. P. (1976). A revision of the family Galaxiidae (Pisces) in Tasmania. Aust.
JI mar, Freshwat. Res. 27(2): 297-349.

BENZIE, V. (1968). The life of history of Galaxias vulgaris Stokell with a comparison with
G. maculatus attenuatus. N.Z. JI mar. Freshwat. Res. 2(4): 628-653.

CADWALLADER, P. L. (1975). Distribution and ecology of the Canterbury mudfish
Neochanna burrowsius (Phillipps) (Salmoniformes: Galaxiidae). JJ R. Soc. N.Z. 5(1):
21-30.

CADWALLADER, P. L. (1976). Breeding biology of a non-diadromous galaxiid, Galaxias
vulgaris Stokell, in a New Zealand river. J] Fish. Biol. 8: 157-177.

ELDON, G. A. (1971). Suspected terrestrial deposition of eggs by Neochanna apoda
Giinther (Pisces: Salmoniformes: Galaxiidae). J/ Fish. Biol. 3: 247-249.

ELDON, G. A. (1978). Life history of Neochanna apoda Giinther (Pisces: Galaxiidae).
Fish. Res. Bull. N.Z. 19: 1-44.

HEFFORD, A. E. (1931a). Whitebait, Rep. Fish, N.Z. 1930: 15-16, 21-23.

HEFFORD, A. E. (1931b). Whitebait. Rep. Fish. N.Z. 1931: 14-17.

HEFFORD, A. E. (1932). Whitebait investigation. Rep. Fish. N.Z. 1932: 14-15.

HOPKINS, C. L. (1971). Life history of Galaxias divergens (Salmonoidei: Galaxiidae).
N.Z. J1 mar. Freshwat. Res. 5(1): 41-57.

McDOWALL, R. M. (1968). The status of Nesogalaxias neocaledonicus (Weber and de
Beaufort) (Pisces: Galaxiidae). Breviora 286: 1-8.

McDOWALL, R. M. (1970). The galaxiid fishes of New Zealand. Bull. Mus. Comp Zool.
Harv. 139(7): 341-431.

ee z M. (1971). The galaxiid fishes of South America. JJ Linn. Soc. Zool.
O(1): -73.

McDOWALL, R. M. (1973a). The status of the South African galaxiid. Ann. Cape Prov.
Mus. (Nat. Hist.) 9(5): 91-101.

McDOWALL, R. M. (1973b). Limitation of the genus Brachygalaxias Eigenmann, 1928
(Pisces: Galaxiidae). JJ R. Soc. N.Z. 3(2): 193-197.

McDOWALL, R. M. (1978). A new genus and species of galaxiid fish from Australia. JJ R.
SOC MINA WSC1) =) 1152124:

McDOWALL, R. M. and FRANKENBERG, R. S. In Prep. The galaxiid fishes of Australia.
- Rec. Aust. Mus,

MASSOLA, A. (1938). Description of a new species of Galaxia. Aquar. JI San Francisco
11(10): 129-130.

MUNRO, I. S. R. (1957). Handbook of Australian fishes, No. 8. Aust. Fish. Newslett.
16(2): 33-36.

POLLARD, D. A. (1971). The biology of a landlocked form of the normally catadromous

salmoniform fish Galaxias maculatus (Jenyns). I. Life cycle and origin. Aust. JI mar.
Freshwat. Res. 22(2): 91-123.

SCOTT, E. O. G. (1971). On the occurrence in Tasmania and on Flinders Island of Brachy-
galaxias Eigenmann, 1928 (Pisces: Galaxiidae) with descriptions of two new sub-species.
Rec. Q. Vict. Mus. 37: 1-14.

SHIPWAY, B. (1953). Additional records of fishes occurring in the freshwaters of Western
Australia. W. Aust. Nat. 3: 173-177.

ee F. (1940). The mountain minnow — some additional notes. Aust. Mus, Mag.
: 234-237.

314 Aust. Zool. 19(3), 1978

Comments on the Technique of Acid Dissolution of
Coral Rock to Extract Endo-Cryptolithic Fauna

P. A. HUTCHINGS and P. B. WEATE
Australian Museum, 6-8 College Street, Sydney, N.S.W. 2000.

ABSTRACT

Comparisons of fauna recovered by hammer and chisel technique and acid
dissolution technique are made. A modified technique for acid dissolution of
coral rock is described in which complete identifiable animals are recovered.

INTRODUCTION

Brock and Brock (1977) described a technique for dissolving coral rock in
dilute acid to extract the infauna. The term infauna according to Hutchings (1974)
includes the true borers and the opportunistic species which cannot themselves
bore but utilise the burrows made by the true borers. As the term infauna is
normally used for describing infaunal communities in soft sediments, we propose
that the term infauna be replaced with the term endo-cryptolithic fauna indicating
its two components, the endolithic fauna the true borers and the cryptolithic fauna
the opportunistic species.

Previously two main methods of extracting this endo-cryptolithic fauna have
been used. Clausade (1970) put coral samples into freshwater for several hours to
force the mobile cryptolithic fauna out. Similarly Leviten (1976) allowed coral
samples to stand in seawater for several hours and as the water became anoxic
the mobile cryptolithic fauna evacuated the sample. Subsequently both these
workers broke up the coral to extract the cessile endolithic fauna. However, as
the samples are not preserved immediately many of the smaller animals may de-
compose either in or out of the coral rock and the results are thus only semi-
quantitative (Brock and Brock, 1977). The other main method used by McCloskey
(1970), Grassle (1973), Kohn and Lloyd (1973) and Hutchings (1974, 1978)
is to crush the coral with a hammer and chisel after rapid fixation and then to sort
the resulting residue under a microscope. These latter methods, although tedious,
give quantitative results and are extensively quoted when discussing the biomass
or numbers of individuals of endo-cryptolithic fauna in coral rocks. It is presumed
however, that the acid bath technique of Brock and Brock (1977) will be used in

Aust. Zool. 19(3), 1978 315

P. A. HUTCHINGS and P. B. WEATE

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Aust. Zool. 19(3), 1978

316

EXTRACTION OF ENDO-CRYPTOLITHIC FAUNA

future studies as it is rapid and efficient. Brock and Brock suggest that the biomass
and numbers of individuals of the endo-cryptolithic fauna extracted by other
methods are dependent on the patience and ability of the worker, which makes
absolute quantification and comparative studies between these methods and the
acid bath technique difficult. Brock and Brock did not attempt to quantify the
percentage of endo-cryptolithic fauna overlooked by the hammer and chisel extrac-
tion method. Such quantification is essential to see whether all the earlier work
using this extraction technique is valid and can therefore be compared with any
future work using the acid bath technique. Therefore the coral residue or chips
remaining after the extraction of the endo-cryptolithic fauna by the hammer and
chisel method were dissolved using the acid bath technique to determine the per-
centage of fauna which had been missed. As it was our intention to process all our
future samples using acid dissolution, the technique of Brock and Brock (1977)
was modified to obtain faster dissolution and to ensure that the extracted fauna was
intact and readily identifiable.

Finally glacial acetic acid was tested as a dissolving agent for the dead coral
since it had been suggested as a potentially suitable acid.

METHOD

The residue of 3 samples which had been processed by breaking them up
with a hammer and chisel to extract the fauna were selected at random from over
100 samples. The fauna of these samples is currently being analysed. The 3 samples
had an original dry weight of 687.9, 652.1 and 618.3 ¢ respectively before being
used as part of an experimental series to determine the rate of colonisation by endo-
cryptolithic fauna of coral rock (Hutchings and Weate, in press). The residue of
each sample which consisted of small coral chips (10%-10-20 mm, 30%-5-10 mm,
40%-2-5 mm and 20%-1-2 mm, in diameter), was dissolved in an acid solution
(5% formalin acidified with nitric acid to 4% by volume) using the technique
described by Brock and Brock (1977). The rate of acid dissolution was increased
by using six plastic ice-cream buckets (21., 15 cm x 15cm x 10cm) the sides and
bottom of which were replaced with 0.3 mm polyester mesh. They sat on a re-
movable tray in a bath in which the volume of acid was increased to 451-, thus
necessitating less frequent replacement. The solution was agitated by 2 magnetic
stirrers. The acid solution however, was still replaced as it became neutralised
every 2-3 days. The residual pulp remaining after the coral chips were dissolved,
was sorted under a microscope and the fauna such as molluscs, crustaceans, sipun-
culans and polychaetes were extracted, weighed and counted.

In order to ensure that the majority of the endo-cryptolithic fauna is removed
intact and well preserved from dead coral samples using the acid dissolution tech-
nique, the following procedure is recommended. After a sample has been collected
-and put into a polythene bag and sealed, it should then be left for approximately
an hour sitting in cool water to allow many of the mobile cryptolithic organisms

Aust. Zool. 19(3), 1978 317

P. A. HUTCHINGS and P. B. WEATE

to evacuate the sample of their own accord. Care should be taken to prevent the
sample becoming too anoxic. This fauna is then removed and fixed separately and
the rock sample is then broken into small pieces and all additional visible fauna
is removed. The remaining coral chips and seperated faunal samples are fixed in
7% formalin. Several days later, the coral chips are dissolved in the acid solution
as outlined above to extract the smaller elements of the fauna.

Experiments were also carried out using glacial acetic acid to dissolve the
coral block after having been previously fixed in gluteraldehyde.

RESULTS

The percentage of faunal biomass recovered in the pulp in comparison to
the total, previously extracted by the hammer and chisel method is 8.8, 3.3 and
3.8% respectively for the 3 samples. Therefore the majority of the endo-cryptolithic
fauna (91.2, 96.7, 96.2% respectively) is extracted by the hammer and chisel
method. In Table I, a breakdown of the biomass in terms of number and size of
individuals extracted by the hammer and chisel technique and from the resultant
coral chips by acid dissolution is given. The animals recovered from the coral
chips, i.e. the fauna missed by the hammer and chisel technique, were predomin-
antly polychaetes and were mainly small thin worms. The relatively high figure of
8.8% for one sample is mainly due to the presence of one gastropod whose weight
minus the shell was 0.04 g, representing 4.8% of the fauna missed. Most of the
fauna including the sipunculans were well preserved and easily identifiable. The
vast majority of molluscs whose shells would dissolve in the acid had been removed
during the hammer and chisel extraction. Trial runs with ophiuroids which are
relatively rare (none were in the 3 samples dissolved) showed that their skeleton
is partially dissolved in acid but they are generally still identifiable.

The rest of the pulp remaining after 4 to 6 days of acid dissolution, consisted
mainly of fragments of sponges, bryozoans and algae. These fragments are difficult
to identify and separate into species, and this technique is therefore not suitable for
determining the biomass of these colonial animals or algae. MacGeachy (1977)
has described a more suitable technique using ultra thin sections to determine the
biomass of boring sponges and this could probably be adapted for the bryozoans,
ascidians and algae.

Dissolving coral rock in glacial acetic acid to extract the endo-cryptolithic
fauna is not recommended as many of the animals are partially or completely
dissolved.

DISCUSSION

Our results show that the majority of the endo-cryptolithic fauna is extracted
by the hammer and chisel method and therefore the figures quoted by workers
such as McCloskey (1970), Kohn and Lloyd (1973), Hutchings (1974) and Hut-

318 Aust. Zool. 19(3), 1978

EXTRACTION OF ENDO-CRYPTOLITHIC FAUNA

chings and Weate (1977) are comparable to those obtained by Brock and Brock
(1977). Kohn and Lloyd’s (1973) figures are probably more accurate than those
of Hutchings (1974) as they pulverised their coral chips to powder. However,
they may have rendered a few species unidentifiable by such harsh treatment (pers.
comm). Although we recommend the acid dissolution technique for rapid extrac-
tion of the endo-cryptolithic fauna, we have modified it to ensure that the majority
of animals are removed intact and well preserved. This is important since much of
the fauna is new or represents new records.

Direct comparisons between biomass, numbers of individuals and_ species
composition of different blocks of dead coral rock are still difficult. For the physical
and biological nature of the coral rock such as surface area, percentage cover of
epifauna and flora, density of the coral skeleton and the percentage of boring in
the coral rock profoundly influence the biomass, numbers and species composition
of the endo-cryptolithic community (Hutchings and Weate, 1977). Many workers
have shown that live coral has a poor endo-cryptolithic fauna and that the fauna
does not develop until the coral dies. However, when sampling in the field, one
has no idea of how long that block of coral has been dead and therefore how old
the community is. Also the community has a finite existence as eventually the
endolithic fauna will destroy the block of coral completely. This probably explains
much of the between sample variation reported by various workers, and perhaps
more emphasis in the future should be put on determining the rates of colonisation
of newly dead coral and the rates of boring by the endolithic fauna to determine
the age of the community.

ACKNOWLEDGEMENTS

We gratefully acknowledge support from the Australian Research Grants Committee who
pays the salary and travel costs of one of us (PBW).

REFERENCES
BROCK, R. E. and J. H. BROCK (1977). A method for quantitatively assessing the in-

| faunal community in coral rock. Limnol. Oceanogr. 22: 948-951.

CLAUSADE, M. (1970). Répartition qualitative at quantitative des polychétes vivant dans
les alvéoles des constructions organogénes épirécifales de la portion septentrionale du
Grand Récif de Tulear. Rec. Trav. Sta. mar. Endoume 1/0: 259-270.

GRASSLE, J. F. (1973). Variety in Coral Reef Communities, p. 247-270. In: O. A. Jones
and R. Endean (eds.), The Biology and Geology of Coral Reefs vol. 2, Biology 1.
Academic Press.

HUTCHINGS, P. A. (1974). A preliminary report on the density and distribution of inver-
tebrates living on coral reefs. Proc, (2nd) Int. Coral Reef Symp. (Brisbane) 7: 285-296.

HUTCHINGS, P. A. (1978), Non-colonial cryptofauna p. 251-263. In: D. R. Stoddart and
R. E. Johannes (eds.). Coral reefs: research methods. Monograph on oceanographic
methodology 5. UNESCO, 581 pp.

HUTCHINGS, P. A. and P. B. WEATE (1977). Distribution and abundance of cryptofauna
from Lizard Island, Great Barrier Reef. Mar. Res. Indonesia 17: 99-113.

HUTCHINGS, P. A. and P. B. WEATE (in press). Experimental recruitment of endo-cryp-

tolithic communities at Lizard Island, Great Barrier Reef: Preliminary Results. New
Zealand Inst. Ocean. Mem.

Aust. Zool. 19(3), 1978 319

P. A. HUTCHINGS and P. B. WEATE

KOHN, A. J. and M. C. LLOYD (1973). Polychaetes of truncated reef limestone substrates
on Eastern Indian Ocean coral reefs: Diversity, abundance and taxonomy. Int. Rev.
Gesamten Hydrobiol. 58: 369-399.

LEVITEN, P. J. (1976). The foraging strategy of vermivorous conid gastropods. Ecol.
Monogr, 46: 157-178.

MacGEACHY, J. K. (1977). Factors controlling sponge boring in Barbados reef corals
Proc. (3rd) Int. Coral Reef Symp. (Miami) 2: 477-485.

McCLOSKY, L. R. (1970). The dynamics of the community associated with a marine
scleractinian coral. Int. Rev. Gesamten Hydrobiol, 55: 13-81.

320 Aust. Zool. 19(3), 1978

The Introduction into Western Australia of the Frog
Limnodynastes tasmaniensis Gunther

A. A. MARTIN
Department of Zoology, University of Melbourne,
Parkville, Vic., 3052,

and

MW. Je YLER
Department of Zoology, University of Adelaide,
Adelaide, S.A., 5001

INTRODUCTION

There is a number of records of the establishment of anurans in geographically
extralimital areas as a result of intentional or accidental releases. The intentional
introductions are usually well documented, such as the initial release of Bufo
marinus in Australia (Mungomery, 1935), and the repeated releases of three
Australian species of Litoria in New Zealand (McCann, 1961). However, we are
unaware of any documented accounts of the establishment of an Australian species
following an unintentional release.

In 1977, in the course of field studies of frogs in the East Kimberley region
of north-western Australia, we found a small population of the frog Limnodynastes
tasmaniensis Gunther at Kununurra. Formerly the species was known from eastern
and south-eastern Australia 1,800 km distant; thus it was clearly evident that the
Kimberley isolate had been introduced.

Further observations at Kununurra in 1978 revealed a significant expansion
of the population there, leading us to deduce that the original introduction had
occurred only a few years previously.

Here we document details of the Kununurra population and discuss the
likely manner of its introduction.

EcoLoGIcaAL NOTES

Limnodynastes tasmaniensis is a moderate-sized frog (40-50 mm snout to
vent length) inhabiting an extremely wide variety of environmental niches. It

Aust. Zool. 19(3), 1978 321

A. A. MARTIN and M. J. TYLER

occurs commonly amongst vegetation in damp situations near static or flowing
water, but is adept at avoiding periods of temporary drought. For example, in
the arid north-east of South Australia the frogs spend the day within fissures in
the soil, emerging at night to feed. Throughout its range the species tends to be
an opportunistic breeder, spawning most commonly in static water. In south-
eastern Australia three call races of L. tasmaniensis are recognised (Loftus-Hills,
1973; Littlejohn and Roberts, 1975; Roberts, 1976).

GEOGRAPHIC DISTRIBUTION

Limnodynastes tasmaniensis occurs in an arc from the Eyre Peninsula of
South Australia to northern Queensland (Fig. 1). Moore (1961) listed a specimen
from Somerset at the northern extremity of Cape York Peninsula, but Cogger
(1975) indicates the northern limit to be near Townsville, and we have adopted
that modification here.

FIG. 1.—The distribution of Limnodynastes tasmaniensis in Australia. The distribution is
based on Cogger (1975), Roberts (1976) and unpublished data; the position of
the Kununurra isolate is shown by an arrow,

322 Aust. Zool. 19(3), 1978

INTRODUCTION OF A FROG TO WESTERN AUSTRALIA

THE KUNUNURRA ISOLATE

On 18 and 23 February 1977 we heard L. tasmaniensis calling from an area
of flooded land in Coolibah Drive, Kununurra. Subsequently we recorded the
calls of four individuals, and obtained a reference sample of four specimens now

deposited in the Western Australian Museum (WAM R58830-33).

Observations in the surrounding district established that the species was
confined to the north-western periphery of the Kununurra township, extending
from the junction of Coolibah Drive with the Parry Creek Road to a point 1.8 km
distant on the Parry Creek Road.

In 1978 we returned to the site, and on January 25 we plotted the population
in more detail by making road traverses and noting the presence or absence of
calling male frogs at intervals of 300 m. The extension of range beyond the
1977 limit was considerable, involving a continuous northern extension of 6.7
km, and a total range (in road km) of 8.5 km. However, no expansion of range
east or west of the road had occurred; the population was confined to a roadside
zone no more than 20 m wide. A single reference specimen has been deposited in

the South Australian Museum (SAM R16919).

The population is currently restricted to roadside vegetation, where cover is
provided by a rich growth of mixed grasses to a height of 1 m. The road runs
through flat pastures dissected by irrigation channels. The frogs breed in a
continuous, shallow, flooded depression on the western side of the road, between
the road and a parallel irrigation channel. In the same area we collected Limno-
dynastes convexiusculus, L. ornatus, Cyclorana australis, C. cultripes, Cyclorana sp.,
Litoria nasuta and L. rothi.

CALL CHARACTERISTICS

The call is a short, staccato rattle consisting of 5-7 notes. One individual
gave calls with five or six notes; calls of the second all had six notes; and calls
of the other two individuals all had seven notes. Note duration ranged from
12-16 msec and call duration from 150-228 msec. The dominant frequency of the
calls of all individuals lay at 1,900-2,000 Hz.

Water temperatures at the calling sites of the four recorded individuals ranged
fom 29/1 to 3822-C.

PossIBLE MANNER AND DATE OF INTRODUCTION

We examined the vicinity of Coolibah Drive, Kununurra, seeking information
about possible modes of introduction of the frogs. Several of the nearby houses
were pre-constructed transportable units, and enquiries revealed that these had
been imported from South Australia. The manufacturer is Atlas Industrial Housing
Pty Ltd, whose construction plant is at Pooraka, 12 km north of Adelaide.
Enquiries directed to the manufacturer showed that the company had provided
the transportable modular accommodation at the Lake Argyle construction camp

Aust. Zool. 19(3), 1978 323

A. A. MARTIN and M. J. TYLER

(now the Lake Argyle Tourist Village), and several hundred transportable homes
throughout Kununurra. Most of the latter homes had been despatched from
Adelaide between 1969 and 1971.

Resting directly on the soil at Pooraka prior to transportation, the home
foundations would have provided ideal refuges for L. tasmaniensis. The frogs often
form aggregations of a dozen or more during dry conditions (our unpublished
observations); this could result in a number adequate for colonisation being
transported together. We suggest that L. tasmaniensis was introduced into W.A.
via one or more of the units.

The substantial expansion of the range of the population observed between
February, 1977, and January, 1978, is best accounted for in terms of an extremely
recent introduction. For this reason we suggest 1971, this being the last possible
date for entry via the transportable homes.

DISCUSSION

Roberts (1976) showed that only one call race of L. tasmaniensis, the
Western, occurs west of the Murray River in South Australia. Hence our sup-
position that the Kununurra population of L. tasmaniensis originated in the
Adelaide area can be tested by comparing the call structure of the Kununurra isolate
with that of the Western Call Race.

Unfortunately the temperatures at which our recordings were made consider-
ably exceed those in Roberts’ study (6.0-25.0°C); thus direct comparisons are
not valid. However, the most striking call difference between the three races is
in the number of notes per call; values are generally 1 in the Southern Race, 2-4
in the Northern Race and 3-8 in the Western Race (Roberts, 1976). In this
characteristic the Kununurra population coincides with the Western Call Race,
and the other aspects of call structure analysed, except dominant frequency, also
fall within the range of this race. Dominant frequency of the Kununurra population
lies above the range recorded by Roberts; possibly this is a temperature effect.

Thus the call data are consistent with: the hypothesis of an Adelaide origin
of the Kununurra population of L. tasmaniensis.

ACKNOWLEDGEMENTS

The 1977 field studies were supported by grants from the Australian Research Grants
Committee to M.J.T. and the Ian Potter Foundation to A.A.M.; the Utah Foundation
sponsored our 1978 visit.

John Bishop and Margaret Davies of the University of Adelaide provided field assistance
during both visits, and Bill Duellman (University of Kansas) accompanied us in 1978.
John Caratti of Kununurra provided us with transport in 1977 and helped us in many
other ways.

REFERENCES

COGGER, H. G. (1975). Reptiles and amphibians of Australia. A. H. & A. W. Reed,
Sydney. 584 pp.

324 Aust. Zool. 19(3), 1978

INTRODUCTION OF A FROG TO WESTERN AUSTRALIA

LITTLEJOHN, M. J. & J. D. ROBERTS (1975). Acoustic analysis of an intergrade
between two call races of the Limnodynastes tasmaniensis complex (Anura: Lepto-
dactylidae) in south-eastern Australia, Aust. J. Zool. 23, 113-122.

LOFTUS-HILLS, J. J. (1973). Comparative aspects of auditory function in Australian
anurans. Aust. J. Zool. 21, 353-367.

McCANN, C. (1961). The introduced frogs of New Zealand. Tuatara 8(3), 107-120.

MOORE, J. A. (1961). Frogs of eastern New South Wales. Bull. Amer. Mus. Nat. Hist.
121, 149-386.

MUNGOMERY, R. W. (1935). The giant American toad (Bufo marinus). Cane Growers
Bull oly 19355,°2 1-27.

ROBERTS. J. D. (1976). Call. differentiation in the Limnodynastes tasmaniensis complex
(Anura: Leptodactylidae), Ph.D. Thesis, University of Adelaide.

Aust. Zool. 19(3), 1978 325

kes

Ne Rhea e I

2

i

Insulin Receptor Proteins in the Erythrocytes of
Monotremes and other Vertebrates

ARTHUR WHITE
School of Biological Sciences, University of Sydney, Sydney, N.S.W.

ABSTRACT

The membranes of mature red blood cells (RBC’s) from a variety of vertebrate
species were examined for insulin binding activity. A strong correlation was
found between the presence or absence of a nucleus in the mature cell and
the presence or absence of insulin binding activity in the erythrocyte membrane.
An hypothesis is presented that the nucleus is inyolved in the production of
an insulin receptor protein on the surface of the RBC. The RBC’s of monotremes
and marsupials are of particular interest, as these taxa represent a phylogenetic
dividing line in the evolution of the mammalian non-nucleated erythrocyte.
feageney ol the course of evolution of the enucleate form of the RBC are
discussed.

INTRODUCTION

Mature erythrocytes of mammals are enucleate, although they are derived
from nucleated precursors (e.g. reticulocytes). The erythrocytes of other vertebrate
groups remain nucleated throughout their life span. The distinction between animals
whose erythocytes do not have nuclei and those whose erythocytes have nuclei is
complicated by the fact that some mammals have relatively high reticulocyte
numbers. The camel, for example, has a high reticulocyte count which has been
related to the ability of the red blood cells to withstand osmotic stresses of
dehydration and rehydration (Banjaree e¢ al 1962; Perk 1963, 1966).

Enucleation may be related to changes in the life span of erythrocytes. As
shown in Table 1, mean life span for erythrocytes in the circulation is. much
greater in reptiles and amphibians (both groups possess nucleated erythocytes )
than it is in the mammals. However, birds have nucleated erythrocytes, and a
mean erythrocyte life span similar to, and in some cases shorter, than mammals.
Short erythrocyte life span might therefore be related to the high body temperature
and high metabolic rate of mmals and birds. Basal heat production (Rodnan e¢ al
1957) and metabolic rate (sp. Altland and Brace, 1962) have been correlated with
erythrocyte turnover. That being the case, the enucleate condition of mammalian
erythrocytes, as opposed to the nucleate erythrocyte of birds, must have an

Aust. Zool. 19(3), 1978 327

A. WHITE

TABLE 1

COMPARISON OF THE TURNOVER RATES AND LIFE SPANS
OF A VARIETY OF VERTEBRATE RBC.

Mean RBC
Mean RBC Turnover
Species Life Span (% RBC vol. References
Es (Days) replaced/
0
7
Eutheria Mouse 5 Burwell et al. 1953
a = Marmot : Brace 1953
aX Rabbit Sutherland et al 1959
<Z Cat : Kaneko et al 1966
a Dog 108 P Weissman et al 1960
oS Auodad* 170 Cornelius et al. 1959
5O Horse Cornelius et al. 1960
ZA.
Z Pro- and
3 Metatheria
O Aves Pigeon : Marvin and Lucy 1957
= Chicken : Hevesy and Ottesen 1945
= Duck : Brace and Altland 1956
A<
ais Reptilia Turtle ; Altland and Brace 1962
<2 Alligator Cline and Waldmann 1962a
A
3 Amphibia Cane Toad c1000 = Altland and Brace 1962
Fa Frog c 200 Cline and Waldmann 1962b

*In the Auodad sheep two populations of enucleate red-blood cells were found.

evolutionary basis. Information about monotremes and marsupials might, on
phylogenetic grounds, be expected to shed some light on the question, but as seen
in Table 1 there are no comparative data on erythrocyte life span or turnover from
those mammals. Data are needed from as many monotreme and marsupial species
as possible, since evolutionary trends may be difficult to sort out from differences
of longevity and body weight in various species.

In this study the metabolic significance of enucleate vs. nucleate erythrocytes
was approached from a different angle. Instead of measuring erythrocyte life-span,
red-blood cell membranes were prepared and analysed for insulin binding activity,
which should be a more direct measure of the metabolic activity of the cell. The
relatively long lived, nucleated cells should have a greater variety of enzymes and
a greater metabolic capacity than enucleate cells. Along with this there should
also be differences in insulin binding activity and an examination of a wide range
of vertebrate species might shed light on the phylogenetic adaptations of erythrocyte
function.

328 Aust. Zool. 19(3), 1978

RBC INSULIN RECEPTORS

MATERTAUS “AND (METHODS

Blood was obtained from 19 vertebrate species. The red cells were separated
from the white cells and plasma by centrifugation. To each ml of compacted red
cells an equal volume of distilled water was added and then mixed to ensure
complete hemolysis. Erythrocyte membranes were collected and washed repeatedly
(until colourless) following the method of Dodge et al (1963). Compacted
membranes were stored at —20°C.

To test for insulin binding activity, 0.25 ml of compacted membranes was
suspended in 0.75 ml cold (4°C) phospho-saline buffer (40 mM) containing
1.5% bovine serum albumin and adjusted to pH 7.4. All further steps were
carried out at 4°C. |

Each 1 ml volume of suspended membranes was subdivided into 10 100 ul
parts, to each of which was added 100 pl of I'* labelled human insulin (98%
T.C.A. precipitable; prepared by the Endocrine Unit, University of Sydney Medical

TABLE 2

PERCENTAGE OF SPECIFIC COUNTS OF BOUND INSULIN
OVER POUR INCUBATIONAL PERIODS: AT 4°C.

Time of Incubation at 4°C

TAXA Speci

ee + hr. 1 hr. Ze. oes

Reptilia Egernia cunninghami 2.681 1.913 <M.E. <M.E.
Tiliqua scincoides 1.884 1.675 <M.E. <M.E.

Crocodylus porosus 3.044 2.806 Ey Gly; <M.E.

Aves* Chicken 2.531 eA EL 1.856 <M.E.
Galah 2.922 2.016 2.002 <M.E.

Monotremata Platypus <M.E. <M.E. <M.E. <M.E.
Echidna <M.E. <M.E. <M.E. <M.E:

Marsupialia Macropus giganteus <M.E. <M.E. <M.E. <M.E.
Macropus robustus erub. <M.E. <ONIEE <M.E. <M.E.

Macropus fuliginosus f. <M.E. <M.E. <M.E. <M.E.

Macropus eugenii <M.E. <M.E. <M.E. <M.E.

Macropus rufus <M.E. <M.E. <M.E. <M.E.

Potorous tridactylus <M.E. <M.E. <M.E. <M.E.

Dasyuroides byrnei =M-E- <M.E. <M.E. <M.E.

Dasyurus maculatus <M.E. <M.E. <M.E. <M.E.

Trichosurus vulpecula 1.643 <M.E. <M.E. <M.E.

Placentalia Rat <M.E. <M.E. <M.E. <M.E.
Rabbit <M.E. <M.E. <M.E. <M.E.

Mouse <M.E. <M.E. <(MiES. ) <GMCB:

The calculated maximum error (M.E.) for the experimental design was 1.590%.

*In 1977 Ginsberg et al. demonstrated the presence of insulin receptors on turkey eryth-
rocytes.

Aust. Zool. 19(3), 1978 329

A. WHITE

School). One of the 10 aliquots was set aside for the determination of total
protein (Lowry e¢ al 1951) to ensure that the total membrane content of each
sample was approximately equal.

In order to determine the non-specific binding component, 100 pl samples
of porcine insulin (20 ug/ml) were added to three of the tubes for each sample.
These three tubes, with remaining six, were treated together as follows:

Binding trials were carried out over 0.5, 1, 2 and 4 hours at 4°C. At the
completion of that incubation period the vials were spun for 20 minutes at 6,000 g.
The supernatant was discarded, and 1 ml chilled buffer was added to each vial
before being centrifuged for 20 minutes at 6,000 g. The supernatant was discarded
and the activity of the bound labelled insulin was determined with a gamma
counter.

The maximum error of the system was determined by processing 12 vials
of labelled insulin through the incubational procedure and scoring any differences
in the counts obtained.

TABLE 3

COMPARISON OF INSULIN RECEPTOR ACTIVITY AGAINST THE
PRESENCE OR ABSENCE OF A NUCLEUS IN ERYTHROCYTES.

Condition of

Presence or

TAXA Species Absence of Insulin
Erythrocytes Receptor Activity
Reptilia Egernia cunninghami Nucleated Present
Tiliqua scincoides Nucleated Present
Crocodylus porosus Nucleated Present
Aves Chicken Nucleated Present
Galah Nucleated Present
Monotremata Platypus Non-nucleated Absent
Echidna Non-nucleated Absent
Marsupialia Macropus giganteus Non-nucleated Absent
Macropus robustus erub, Non-nucleated Absent
Macropus fuliginosus ful. Non-nucleated Absent
Macropus eugenii Non-nucleated Absent
Macropus rufus Non-nucleated Absent
Potorous tridactylus Non-nucleated Absent
Dasyuroides byrnei Non-nucleated Weak
Dasyurus maculatus Non-nucleated Absent
Trichosurus vulpecula Non-nucleated Weak
Placentalia Rat Non-nucleated Absent
Rabbit Non-nucleated Absent
Mouse Non-nucleated Absent
330 Aust. Zool. 19(3), 1978

RBC INSULIN RECEPTORS

RESULTS

Table 2 sets out the results of insulin binding trials on a number of vertebrate
species expressed as “‘percentages of specific counts bound”, calculated as:

% specific counts bound = Bound counts — nonspecific counts

total counts

The presence or absence of insulin receptor activity for each species indicated
by the detectable specific binding of radioactive insulin, or lack of such binding,
is compared with the erythrocyte condition in Table 3. It is clear from this Table
that nucleated erythrocytes possess insulin receptors while enucleated erythrocytes
do not.

DISCUSSION

In terms of erythrocyte longevity there is no significant difference between
birds and mammals (Table 1). However, in terms of RBC insulin receptor activity,
there is a marked difference (Table 2), and monotremes and marsupials are the
same as other mammals in lacking detectable receptor activity and are clearly
different from birds. Monotremes and marsupials are also like the eutherians in
erythrocyte morphology. The echidna (Bolliger and Backhouse 1960a, Parer and
Metcalfe 1967a, Lewis et al 1968), and the platypus (Parer and Metcalfe 1967b),
possess non-nucleated erythrocytes and have low reticulocyte numbers. Parsons
et al (1970) have likewise shown 18 species of marsupials to have mainly enucleate
RBCs. Therefore the presence of enucleate erythrocytes coincides with lack of
detectable insulin receptor activity, suggesting that the nucleus is essential for
the production of the insulin receptor on the erythrocyte membrane.

Two marsupials, Trichosurus vulpecula and Dasyuriodes byrnei, seem to be
an exception to the above, since they both have some (although weak) receptor
binding activity. However both species have unusually high reticulocyte numbers
(1.4% and 1.2% of the total RBCs respectively), and the low binding activity
shown by T. vulpecula and D. byrnei could be explained by circulating reticulocytes
at that level. Some other species of marsupials which were not available for the
present study have also been shown to have high reticulocyte counts; koala
(Bolliger and Backhouse 1960b), hairy-nosed wombat (Parsons e¢ al 1970), and
opossum (Giacometti et al 1972).

Another experimental result which appears to be an exception to the lack
of mammalian erythrocyte insulin binding reported above is the demonstration
of insulin binding sites on human erythrocytes by Gambhir e¢ al (1977). However,
they used a much more sensitive technique, and the interpretation of their results
is difficult. It seems likely that the binding found by Gambhir ef al (1977) is
due to remnant receptor protein in the cell membrane. Certainly such receptors
- could have no physiological significance in mature enucleate erythrocytes. The

Aust. Zool. 19(3), 1978 331

A. WHITE

fact that receptor proteins still exist beyond the reticulocyte stage indicates that
the apparatus which normally degrades these molecules has itself become non-
functional after enucleation.

Further evidence of the essential role of the RBC nucleus in production of
protein within the RBC is provided by the work of Silber et al (1961). They
related the nucleus to the production of RBC surface antigens. Enucleation was
correlated with loss of transplantation antigens (and thus could explain the
survival of transfused erythrocytes ).

Therefore the RBC nucleus has important functions throughout its life span
in most vertebrates and in reticulocytes in mammals. The loss of the nucleus
in both ontogeny and phylogeny of mammals has led to a loss of synthetic ability
of the erythrocyte, loss of receptor proteins and decreased erythrocyte life span.
Clearly two strategies exist in the vertebrates: 1. The production of nucleated
erythrocytes with a complete metabolic repertoire and capable of supporting them-
selves in a functional state in the circulation for relatively long periods; 2. The
production of non-nucleated erythrocytes with reduced metabolic capacity and
with reduced functional circulating life span; i.e. RBC turnover rates are relatively
high. The adaptive advantage of one strategy over the other must depend on the
energetic cost of producing a small number of cells capable of supporting them-
selves for a long time as opposed to producing a large number of cells which
have a relatively short lifespan, but which utilise very little of the blood energy
substrate. :

This study has shed some light on the phylogeny of RBC enucleation to
the extent that it has shown that all mammals, including monotremes, lack
detectable insulin binding on the erythrocytes in marked contrast to other
vertebrates.

A promising area for further studies of the functional significance of the
RBC nucleus in mammals would be studies on the blood of pouch young of
monotremes and marsupials. Marsupial pouch young have high circulating stem
erythrocyte counts. Richardson and Russell (1969), and Yadav (1972), have
found large numbers of eosinophilic megaloblasts in newly born pouch young of
kangaroos. The embryonic young of monotremes and marsupials are, of course,
much more readily available for study than eutherian young of similar development.

ACKNOWLEDGEMENTS

This project was made possible by a number of people who allowed me to gather
enough blood from the variety of species sampled to complete this study; Mr R. Whitford,
Dr P. Johnson, Dr E. Skadhauge, Mr S. Donnellan and Dr G. Grigg. Also I thank
Mr P. Williams of R.P.A.H. for his ideas and inspiration for the study.

REFERENCES

ALTLAND, P. D. & K. C. BRACE (1962). Red-cell lifespan in the turtle and the toad.
Am. J. Physiol, 203, 1188-1190.

332 | Aust. Zool. 19(3), 1978

RBC INSULIN RECEPTORS

BANJAREE, J., R. C. BHATTACHARJEE & T. A. I. SINGH (1962). Haematological
studies on the normal adult Indian camel (Camelus dromedarius). Am. J. Physiol. 203,
ee

BOLLIGER, A, & T. C. BACKHOUSE (1960a). Blood studies of the echidna (Tachyglossus
aculeatus). Proc. Zool. Soc, Lond, 135, 91-97,

BOLLIGER, A. & T. C. BACKHOUSE (1960b). The blood of the koala (Phascolarctos
cinereus). Aust. J. Zool. &, 363-370.

BRACE, K. C, (1953), Life span of the marmot erythrocyte. Blood 8, 648-650.

BRACE, K. C. & P. D. ALTLAND (1956). Life span of the duck and chicken erytluocyte
as determined with C14, Proc. Soc. Exp. Biol. Med. 2, 615-617.

BURWELL, E, L., B. A. BRICKLEY & C. A. FINCH (1953). Erythrocyte lifespan in
small animals; a comparison of two methods using radio iron. Am. J. Physiol. 172,
718-724

CLINE, M. J. & T. A. WALDMANN (1962a). The effect of temperature on red-cell survival
in the alligator. Proc. Soc. exp. Biol. Med, 3, 716-718.

CLINE, M. J. & T. A. WALDMANN (1962b). Effects of temperature on erythropoesis
and red-cell survival in the frog, Am. J. Physiol. 203, 401-404.

CORNELIUS, C, E., J. J. Kaneko & D. C. BENSON (1959). Erythrocyte survival studies
in the mule deer, aoudad sheep and sprinkbok antelope using glycine-2-C14, Am. J.
vet. Res. 20, 917-920.

CORNELIUS, C. E., J. J. KANEKO & D. C. BENSON (1960). Erythrocyte survival
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DODGE, J. T., C. MITCHELL & D. J. HANAHAN (1963). Preparation and chemical
characterisation of haemoglobin-free ghosts of human erythrocytes. Arch, Biochim.
Biophys. 100, 119-130.

GAMBHIR, K. K., J, A. ARCHER & L. CARTER (1977). Insulin radioreceptor assay
for human erythrocytes. Clin, Chem. 23, 1590-1595.

GIACOMETTI, L., A. K. BERNTZEN & M. L. BLISS (1972). Haematologic parameters
of the opossum (Didelphis virginiana). Comp, Biochem, Physiol. 43A, 287-292.
GINSBERG, B. H., C. R. KAHN & J. J. ROTH (1977). The insulin receptor of the turkey

erythrocyte. Endo. 100, 82-90.

HEVESY, G. & J. OTTESEN (1945), Life cycle of the red corpuscle of the hen. Nature
156, 534-536.

KANEKO, J. J.. R. A. GREEN & S. S. MIA (1966). Erythrocyte survival in the cat as
determined by glycine-2-C!+. Proc. Soc. exp. Biol. Med, 1/23, 783-784.

LEWIS, J. H., L. L. PHILLIPS & C. HAHN (1968). Coagulation and haematological
studies in primitive Australian mammals. Comp. Biochem. Physiol. 25, 1129-1135,

LOWRY, H. O., N. J. ROSEBROUGH, A. L. FARR & R. J. RANDALL (1951). Protein
measurement with Folin phenol reagent. J, Biol. Chem. 193, 263-268.

MARVIN, H. N. & D. D. LUCY (1957). The survival of radio-chromium tagged erythrocytes
in pigeons, ducks and rabbits. Acta Haematol. 18, 239-245,

PARER, J. T. & J. METCALFE (1967a). Respiratory studies of monotremes. I, Blood of
the platypus. Resp. Physiol. 3, 136-142.

PARER, J. T. & J. METCALFE (1967b). Respiratory studies on monotremes. II, Blood
of the echidna. Resp. Physiol. 3, 143-150.

PARSONS, R. S., J. ATWOOD, E. R. GUILER & R, W. L. HEDDLE (1970). Comparative
studies on the blood of monotremes and marsupials. I. Haematology. Comp. Biochem.
Physiol. 39B, 203-208.

PERK, K, (1963). The camel erythrocyte. Nature 200, 272-273.

PERK, K. (1966). Osmotic fragility of the camel’s erythrocyte. I. A. Microcinematographic
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RICHARDSON, B. J. & E. M. RUSSELL (1969), Changes with age in the proportion of
red-blood cell types and in the types of haemoglobin in kangaroo pouch young, Aust.
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Aust. Zool. 19(3), 1978 333

A. WHITE

RODNAN, G. P., F. G. EBAUGH & M. R. FOX (1957). The life span of the red-blood
cell and blood-cell volume in the chicken, pigeon and duck as estimated by the use of
Na.Cr®!10,. Blood 12, 355-366.

SILBER, R., S. E. HEDBERG, J. H. AKEROYD & D. FELDMAN (1961), The transfusion
behaviour of avian erythrocytes: the lack of functional transplantation antigens in
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SUTHERLAND, D. A., P. MINTON & H. LANZ (1959). The life span of the rabbit
erythrocyte. Acta. Haematol. 21, 36-42.

WEISSMAN, S. M., T. A. WALDMANN & N. I. BERLIN (1960). Quantitative measure-
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YADAV, M. (1972), Characteristics of blood in the pouch young of the marsupial Setonix
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334 Aust. Zool. 19(3), 1978

ROYAL ZOOLOGICAL SOCIETY OF NEW SOUTH WALES
Established 1879
Registered under the Companies Act, 1961

His Excellency the Governor of New South Wales, Sir Arthur Roden Cutler,
ma K.CMG. ECVO., CBE. KiSti.,. B.Ec., (Hon); LLD. -(Hon),-D-Se.
COUNCIL, 1978

President:
Gordon Clifford Grigg, B.Sc., Ph.D.

Vice-Presidents:

Ronald Strahan, M.Sc., F.L.S., F.R.
M.I. Biol., F.S.1.H.

William S, Steel, O.B.E., O.G.D.S.
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Leighton C. Llewellyn, B.Sc., Ph.D:

Honorary Secretary: Thomas J. Bergin, B.Sc., B.V.Sc., M.R.C.V.S.
Asst. Hon. Sec.: Mrs M. Wray.
Honorary Treasurer: Mrs D. S. Forbes.

Honorary Editors: Leighton C. Llewellyn (Australian Zoologist),
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Publicity Officer/Librarian: Mrs M. Wray.

MEMBERS OF COUNCIL

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Members of Publications Committee: G. C. Grigg, R. Strahan, L. C. Llewellyn, E. Wingfield.

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THE AUSTRALIAN ZOOLOGIST

VOLUME 19 1978 PART Ill

CONTENTS

A survey of Central Australian Ichthyology. C. J. M. GLOVER and T. C. SIM

Observations on some Buprestidae (Coleoptera) from the Blue Mountains,

INS. We. Dae J. LAWS ep rcerr ce serra

Gait of the brush-tailed possum (Trichosurus vulpecula). A. J. GOLDFINCH
crete i ae Se eh a eee

Preliminary assessment of defecation patterns for the eastern grey kangaroo

(Macropus giganteus). G.. J. TE. FTV sect ccoes ence

Short papers —
Eye closure during agonistic behaviour of Rattus villosissimus. R. J. BEGG

A triploid male individual amphibolurus nobbi nobbi Witten (Lacertilia:

Aoamidae)., (a. J. WIT TEIN | ccissssseniosamsssaneeinsuicumoccenbeaideparsieecnenics lela

Sexual dimorphism in an Australian galaxiid (Pisces: Galaxiidae). R. M.

DSTO ALLL sscvcceasinesbiessernennhiicsnisbce nnrseomenutchesgaccna tsar eerste ga

Comments on the technique of acid dissolution of coral rock to extract endo-

cryptolithic fauna. P. A. HUTCHINGS and P. B. WEATE 6. csccsssssscssessnseeene

The introduction into Western Australia of the frog Limnodynastes tasmani-
ensis Gunther. A. A. MARTIN and Ms J. TYGER nc. nccscumueee

Insulin Receptor Proteins in the Erythrocytes of Monotremes and Other
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