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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
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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:
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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
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(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
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Figure 3—Grooming: Pouch
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Figure 4—Sexual behaviour: “Inspection” by male
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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,