Observations on the behaviour of the red Kangaroo (Megaleia rufa) in captivity

School of Zoology, University of N e w South Wales, Sydney , Australia

Observations on the behaviour of the red Kangaroo (Megaleia rufa) in captivity

by ELEANOR RUSSELL

With 8 figures

Received: 17. 8. 1968

C o n t e i i t s : Introduction p. 385. - Material and Methods p. 386. - Results: 1. Daily activity pattern p. 387. - 2. Body posture\ when stationary p. 388. - 3. Locomotion p. 389. - 4. Feeding and Drinking p. 390. - 5. Comfort movements - Grooming p. 391. - 6. Miscellaneous p. 391. - 7. Interaction with other individuals p. 393. - (a) Non-sexual interactions between adults p. 393. - (b) MotheriYoung interactions p. 396. - Discussion p. 401. - Summary p. 403. - Zusammenfassung p. 403. - References p. 404

Introduction

The red kangaroo, Megaleia rufa, is widely distributed in the dry inland areas of Australia, generally in areas of less than 15 inches annual rainfall. It is found in all types of habitat except thick scrub, but it is most commonly found in open plain country which has a few scattered trees. An adult cl' weighs up to 75 kg., and stands about 6 feet tall. An adult 9 weighs up to 35 kg., and stands 5 feet tall. 66 are typically rusty red in colour, while most 99 are blue-grey. Both have a paler tail and undersurface. There are conspicuous white and dark brown or black face markings behind the nostrils, and there are obvious individual differences in these.

Despite much popular description of various macropod marsupials, very little detailed information has yet been published on the behaviour of the red kangaroo. From records of the daily activity of a few individuals in the field, CAUGHLEY (1962) found that although they fed and moved by night and rested up during the day, they showed some activity during the day. If they slept a t all, i t was for only short periods.

NEWSOME (1965) found that during severe drought, the kan, Garoos were concentrated along the margins of watercourses and open plain areas (where food persisted) that dissected the mulga (Acacia aneura) woodland. They rested in the shade of the trees during the day, and came out to feed a t night. After good rain had fallen, the distribution was much more scattered, as food was more widely available, but was still closely related to the presence of some shelter. FKITH (1964) showed that when a reasonable amount of food is available, red kangaroos are relatively sedentary-a group marked by an albino Or was observed for 8 months and had a daily range of only 3 miles.

Free-ranging animals tend to be found in small groups, but CAUGHLEY (1962) could find no other evidence of social organization than the relationship between mother and offspring. Groups did not appear to have a constant composition.

Z. f . l'ierpsychol. Ed. 2 7 , Heft 4 2 5

386 ELEANOR RUSSELL

Aspects of reproduction and reproductive behaviour have been described by SHAKMAN and PILTON (1964) and SHAKMAN and CALABY (1964). The 9 has an oestrous cycle of 35 days, and a gestation period of 33 days. The young at birth weighs approximately 800 mg., and crawls unaided from the urogenital opening to the pouch, where it attaches to one of the four teats. The young reinailis in the pouch completely for about 190 days, although from 150 days on the head is often to be seen, and the young may crop grass. From 190 days on, the young leaves the pouch for increasingly longer periods. The young leave the pouch permanently after about 235 days, and mothers which have inated at the previous post-partum oestrus and are carrying a delayed blastocyst give birth to a new young within 24 hours of the previous young leaving the pouch. The first young, now called the young-at-foot, is suckled for a further 130days from outside, using the same teat as when it was in the pouch. The second young is suckled on a second teat i n the pouch. There inzy also be another dorinant blastocyst in the uterus.

Material and Methods Observations of a yard colony of red kangaroos was commenced i n March 1966. The

main aim of thcse observations was to determine thc nature of mother-young interactions, but it was found possible to record other behaviour observed. In the following account, I have attempted to describe the behavioural repcrtoirc of the red kangaroo in captivity, as a basis for comparison with the behaviour of other species of macropod marsupial, as the basis of a study of the development of the behaviour of thc young, and to simplify the problem of field observation of such a fast moving animal.

The animals observed were 29 red kangaroos (Megaleia rufa) of varying ages and histories. This number includes 4 young (SK 10, 17, 21, 23) which at the beginning of the study were pouch young, but which left the pouch in the course of the year, and for the rest of the observation were suckled by their mothers as young-at-foot. 2 animals (SK 30 and SK 26) were young-at-foot at the beginning of the study and were weaned in the course of the year. 6 animals died in the course of the study. The oldest animal was 10 years old, most adults were 2-5 years old.

None of the animals was captured in the wild as an adult. Most were reared in captivity either by their own mothers or from young whose mothers were shot in the wild. Some were animals which had been reared as domestic pets from wild-caught young. They were kept in large yards at the Cowan Field Station of the Zoology Department, University of New South Wales. The first animals were transferred to the Cowan yards on 10, 13, 15 January, 1966. More were added on 4 March, 1966, and observation commenced on 25 March, 1966.

The yards were situated on both sides of a central laneway, in which was an observation tower, from which there was a clear view of yards Aa, A4 and Bi, where the animals observed were kept. Only one yard was under continuous observation at one time. When the animals were first introduced to the yards, there was a dense cover of grass, shrubs and young eucalypts, but in about 6 weeks this was eaten down and the floor of the yards consisted of hare earth. The animals were fed sheep nuts and oats from a self-feeding hopper and lucerne from a rack.

Two groups of kangaroos were observed: Group 1 of 12 individuals, observed from 25/3/66 to 916166, when the group was broken

and individuals moved into other groups in other yards. For the duration of the time :!bbservation, Group 1 was in yard As (area 470 m2).

Growp 2, ultimately of 17 individuals, was observed from 25/3/66 to 18/5/66 in yard A4 (area 470 m2) and from 29/5/66 to 14/12/66 in yard Bi (area 560 m2). There were additions to this group when young left the pouch or when animals were transferred from another yard. Some deaths also occurred.

Each group contained only one d , since, if two 8 8 are kept in the same yard, they may fight and damage each other.

Changes of yards and groups were minimised but were sometimes necessary because the same animals were the subjects of a number of experimental programmes. Normal yard practice was for the kangaroos to be moved to a new yard when they ate their current yard bare. This was done for yards As and A4 after 3 months, but the group in yard Bi was not moved again after they were moved there from A4.

Observations on the behaviour of the red Kangaroo 3 87

Each aninial was ear-marked with a pattern of holes representing its number. With prricticc, individuals can be identified very easily from ear-marks and individual charac- teristics of size, colour, markings, etc. To aid the initial process of familiarisation, sonic animals were niarkcd with different coloured plastic collars, but these later became un- necessary.

126 hours of observation, from April 1966 to 1)ecenibcr 1966 are considered. 6 hours observation per wcck was done when possible, generally on one day, from 9.30 a. m. to 12.30 p. in. and 2.00 p. m. to 5.00 p. m., but occassionally the week’s observations were made on 2 o r 3 days, or not a t all. O n two occasions, observations were made f rom 5.00 p. m. to 9.00 a. m. Observations were spoken into a portable tape recorder and later transcribed with an added time-base of 30 seconds.

All behaviour was recorded as the occurrence of the motor pattern involved. An estimate of the duration of any pattern could be made from the number of 30-second periods over which it extended; quantitative analysis of these obscrvations will be pretented in a later paper. Hcre only a description of the behavioural repertoire is given.

Results 1. Daily Activity Pattern

The daily activity pattern varied with the weather and the time of year. In the yard animals with food freely available, there was an increase in

activity to wards late afternoon. On a still sunny day, they spent a large part of the day resting but not

completely inactive. They lay in the sun for a time, then stood up, perhaps scratched and moved into the shade. Rarely more than 1-2 animals, o r a mother/young pair, in a group were moving a t one time in the middle of the day. Nevertheless, a complete range of activities could occur during this time, but a t much lower frequency than from late afternoon to early morning.

Fig. I : Body postures when stationary (a) Standing erect (b) Alert erect (c) Standing semi-erect (d) Standing crouched (e) Lying down, head up

388 ELEANOR RUSSELL

On overcast days, especially in winter, there was much more activity-general movement around the yards plus feeding. On windy days, the animals were often disturbed and a strong gust of wind in the trees would bring them all to their feet. After a little general movement, most lay down until disturbed again.

In the summer, the animals were generally lying down resting alter- nately in shade and sun (but mostly shade) by 9.30-10.00 am., and they did not become active until 4.30-5.00p.m. In winter, they did not begin “resting” until about 10.30, lying mostly in the sun, but perhaps moving briefly into the shade, and became active again about 4.00-4.30 p.m.

2. Body Postures when Stationary (a) Standing erect. The animal stood with back almost vertical, tail

horizontal. I t was usually seen when the animal was alerted or rather disturbed (fig. 1 [a]).

(b) Alert erect. The animal stood back vertical or even inclined back from the vertical. Head up, ears up, animal poised ready for rapid movement (fig. 1 [b]).

(c) Stunding semi-erect. The back was held almost horizontal, with forelimbs not touching the ground, and head higher than back. In normal circumstances the animal did not stand like this for very long periods-if the animal remained standing it was replaced by (d) or possibly (a), or the animal relaxed and lay down. In yard animals, it was most commonly seen in animals which had been disturbed-by a noise or by someone approaching the yard. After initial rapid movement away, or a period of standing erect or alert, a period of standing semi-erect followed, before the animal lay down or returned to standing crouched (d) (fig. 1 [c]).

(d) Standing crouched. The animal stood with forelimbs on the ground (just touching), back bent, head lower than highest point of back. Animals held this position for considerable periods of time. On a hot day in summer, it was quite common to see an animal stand crouched for an hour or so, in the shade. O n a cool day, they stood in the sun. Standing crouched was commonly interspersed with periods of lying down ([el, [f] and [g]) (fig. 1 [dl) .

(e) Lying down , head up. The animal lay on its side, head held up at right angles to the body. The eyes were open, and the animal was obviously not asleep. This was the commonest of the three lying down positions (fig. 1 [el, fig. 2).

(f) Lying down , head down . The animal lay completely on its side, with the head also on the ground. Eyes may be closed or open, but i t was often very hard to tell.

(8) Lying on buck. The animal lay on its back, with the head to one side, legs apart, and abdomen perhaps turned a bit on one side. The undersurface was thus exposed to the sun on a cool, still day, or to a breeze on a hot day, when the animal was lying in the shade. This position was not generally maintained for longer than about 10 minutes.

When a red kangaroo was “inactive” during the day, it would change its manner of lying, stand for a short time, move from shade to sun or vice versa, or from one patch of shade to another, lie down again. This was interspersed with short periods of grooming, and they also sometimes fed for short

Observations on the behaviour of the red Kangaroo 389

Fig. 2: 9 and young-at-foot pair in typical position, with young standing crouched beside its recumbent mother

periods, stopping by the hay rack or food hopper for a few minutes. Morning and evening feeding occurred in longer “bouts”.

The act of lying down was almost always preceded by digging. The animal stood crouched as in (d), scratched the ground one or more times with its forelimbs, and then lowered one flank sideways into the area dug. The bareness of a yard after about 6 weeks occupation is partly attributable to this excavation. In some areas of a yard, numbers of quite deep “hip-holes’’ can be found after a few weeks’ occupation.

Similar excavations have been seen in the field, by the base of trees and shrubs under which red kangaroos or euros were seen to be lying during the day. In other places, such holes or bare places were also seen, with kangaroo droppings nearby, although the animals were not seen.

3. Locomotion

The hopping gait of the kangaroo is well known. I n yards, rapid hopping locomotion was not usual, unless the kangaroos became disturbed, as for instance when one of their number was being caught for examination. Rapid locomotion was digitigrade, and the tail and forelimbs did not touch the ground. In slow hopping progression, the tail touched the ground, the forelimbs did not.

Particularly when grazing, but also at other times, a slow, almost “walk- ing” gait was seen, which involved the tail, fore- and hind-limbs-each was moved forward while the other two were on the ground.

390 ELEANOR RUSSELL

4. Feeding and Drinking Patterns of feeding as seen were, of course, influenced by the way food

was provided. (a ) Feeding f rom self-feeding hopper

Sheep nuts were provided in a self-feeding hopper. A number of animals could stand and feed from the trough at the same time. Some animals used their forelimbs to scratch round in the trough, to make more pellets fall into the trough. SK 7 invariably did this, and her eating was always accompanied by a lot of noise. Other animals ate very quietly. Food scattered on the ground beside the hoppers was also eaten.

(b) Feeding f rom bay racks Generally, hay was pulled down through the bottom of the racks or

picked up from the ground, when it was often held between the forelimbs while it was eaten. A few larger and/or older animals (SK 4, 8, 12, 16, 19) were able to stand on the tips of their hind limbs, balanced on the end of the tail and supported by forelimbs on the top of the hay rack, and pull hay from the top of the rack.

(c ) Feeding on natural vegetation After about 6 weeks, a yard of area 500m2 with 12 kangaroos became

quite bare. All low-growing shrubs were stripped of leaf and bark, and all grass was eaten. The animals used their forelimbs to dig in the search for grass, or to help them to eat as much as possible of a clump of grass. Low-

Fig. 3: Positions adopted when grooming (a) Use of hindlimb for head and shoulders (b) Tail-forward position for grooming tail and hindlimb (c) Position for grooming pouch and front of abdomen (d)-(e) Usc of forelimb for scratching front of thorax and flank


growing eucalypts were also eaten, and again, some animals raised themselves on hind legs and tail to pull down leaves otherwise out of reach.

After eating, either hay or pellets, animals were frequently seen to stand very erect and make “pumping” or “hiccoughing” movements of the thorax. EALEY (1962) called this “coughing”, and compares this with rumination in ungulates. BARKER, BROWN and CALABY (1963) have reviewed a large number of reported occurrences of this behaviour in Macropods, and conclude that it is not analogous to rumination in ruminants.

5. Comfort Movements - Grooming

The animals were able to reach every part of their body to groom, using either the forelimb or hind limb. The tongue and teeth were also used (fig. 3

Head and Shoulders The hind limb was used only to groom and scratch ears and muzzle,

under the chin, high up on the shoulder and the back of the neck. The animal might be (i) standing crouched, (ii) propped on the tail and one foot (fig. 3 [a]), or (iii) sitting with tail forward between the hind legs, and head lowered (fig. 3 [b]). The inner (syndactylous) claw of the hind limb was used.

[al-[el)*

The forelimbs also were used to reach anywhere in this region.

Tail To groom the tail, the animal sat with the tail stretched out in front

between the hind limbs as in fig. 3 (b). The tail was then scratched along its length with the forelimbs, or the tongue and teeth were used.

Pouch and front of abdomen 09 used the tongue for what is called “pouch cleaning”, holding the

pouch open with the forelimbs, inserting the muzzle into the pouch, and licking. This occurred whether or not the 9 had a pouch young. She might be (i) standing crouched, (ii) sitting with tail forward, or (iii) lying on her back. Pouch cleaning also occurs, with particular intensity, in the 24 hours before a 9 gives birth (SHARMAN & CALABY, 1964). At this time, she usually sits with tail forward, and her back leaning against (e.g.) a tree or a fence. The area round the urogenital opening (pouch or testis) and the front of the abdomen was generally sniffed and licked or groomed with the forelimbs (fig. 3 [c], [d], fig. 4).

Fore and hind limbs The hind limb was usually groomed or scratched when the animal was

sitting with the tail forward (fig. 3 [b]). The fore-paw was licked or rubbed against the mouth, often after eating.

This was also commonly seen in the young-at-foot after it had been suckled by its mother. The animal usually stood semi-erect. Typically, the rest of the body was groomed with the forelimbs only. The animal might be standing in any position, sitting with tail forward, or lying down.

6. Miscellaneous (a) Digging

Before lying down, the kangaroos generally dug with their fore-paws, or at least scratched the surface of the ground in front of them. When they lay down, it was generally the flank which was lowered onto the area dug. Some-

392 ELEANOR RUSSELL

Fig. 4 : Pouch cleaning. Note use of forelimbs to hold pouch open; the sphincter round the mouth of the pouch is also relaxed

times “digging” was a mere perfunctory scrape, but on other occasions, digging lasted for 10-20 seconds, and a considerable quantity of soil was thrown out behind the animal. Certain areas in the shade of trees became dotted with quite deep hollows excavated in this way. Certain larger, older, dominant animals were often to be found in the same hollow, and often chased away another animal who was lying there when the “owner” approached. This excavation must contribute to the destruction of vegetation on any area inhabited by kangaroos. Similar excavations have been seen in the natural habitat of the animal, round the bases of trees and shrubs.

(b ) Licking of Forelimbs In a number of circumstances, kangaroos were to be seen licking their

forelimbs, from shoulder topaw, so that the whole limb waswet and appeared darker than the surrounding fur (fig. 5). Sometimes the same effect was produced by rubbing the limb against the mouth, without licking. Occasion- ally the front of the leg and abdomen was wet in the same way. This behaviour was seen (i) on very hot days, (ii) when the animals had been disturbed-by workmen erecting fences nearby, or when one or more animals in a yard was being caught for examination, transfer to another yard, etc. In animals not used to handling, this may involve some chasing, and even on a cold day, they started to salivate profusely, and to rub their forelimbs against their mouths. (iii) An animal transferred to a new yard with strange animals was seen next day to remain isolated from other animals in the yard, and to lick her forelimbs frequently-, at a time when this behaviour was not evident in other animals in the yards.


Fig . 5 ; Liking of forelimbs. The wet area of the forearm is clearly visible. Abdomen and hindlimb have not been licked. Air temperature: 33" C

(c) Alarm Signals Normally the only sound made by an animal hopping rapidly was a

series of soft thuds. An animal which was alarmed and moved away rapidly made a relatively loud slapping noise with the soles of the hind feet, for the first few hops. This sound was sufficient to alert all nearby animals, including those who were not in a position to see the animal moving fast. If the other animals then detected the disturbance, they also nioved away. If the disturbance was transitory, the animals relaxed from the alert position.

7. Interaction with other Individuals (a ) Non-sexual Interactions between Adults

Interactions between 99 and non-sexual interaction between 0" d and 99 are described i n detail. The behaviour patterns involved f i t the basic pattern of A approaches: B retreats, even if displacement is not involved, and are classified into Non-Agonistic and Agonistic interactions.

Non-Agonistic ApproachlRetrcat Supplanting. A approached B, which moved away, and the approaching

animal A then took the place of B at, for example, the feed bin or in a patch of shade or sun.

Avoiding. A moved out of the way of B moving towards it. This is almost the same as Supplanting, but A did not replace B a t e.g. food.

Sniffing. A approached B slowly, and sniffed at it, generally near the mouth or nose, but possibly also near the pouch or urogenital aperture (fig. 6

3 94 ELEANOR RUSSELL

[a]). B might show no response a t all, or might sniff back a t A, in which case they stood nose to nose. Eventually A or B or both moved away. Sniffing each other in this way was seen sometimes when two animals met. It is possible that it is one means by which the animals recognize each other. It

Fig . 6 : Types of interaction between two individuals (a) Sniffing when two adults or mother and young meet (b) Cuffing-agonistic interaction (c) Mother-young pair lying close together in a typicaI spatial arrangement (d) “Sparring”, a complex interaction between mother and young-at-foot, generally seen

after the young has been sudrling

is perhaps questionable whether this pattern should be classified as Approach/ Retreat or Friendly when the sequence is: A approaches B, sniffs B, A goes away from B; but for convenience all encounters of this type have been grouped together.

Agonistic ApproachlRetreat Snorting. The vocalisation which I have called “Snorting” was really

a loud unvoiced “Ha”. It might occur by itself or with Cuffing. It was frequently heard by itself when one animal approached another, who might be feeding, or lying in the sun or shade, and who snorted loudly, which often deterred further approach.

Cuffing. A struck B, with one or both forelimbs, in the region of the neck and thorax. The action was frequently more as if A took hold of B, at the same time pushing away (fig. 6 [b], fig. 7). If B was moving away, A struck at whatever part was nearest, e.g. butt or tail. Occasionally, B might stand still and strike back at A in the same way, or else both A and B might stand erect holding each other in the region of the neck or thorax and occasionally strike with one forelimb.


Fig . 7: Agonistic Approach/Retreat interaction - Cuffing. The large animal second from right (8) is chasing a smaller individual

Snorting and cuffing might occur together or in sequence, when snorting usually preceded cuffing. In order of increasing intensity, the 3 may be placed: Snorting, Cuffing, Snorting-and-Cuffing.

0“-0‘ agonistic interactions were not observed, as only one 0‘ was kept in each yard, or with a group of 99. SHARMAN & CALABY (1964) describe aggressive displays by one male to another, which may be a prelude to fighting in which the animals grasped their opponents round the neck and kicked each other with their hind limbs while supporting themselves on their tails. Grasping the opponent around the neck as described is similar to that seen in the lower intensity interaction described above under “cuffing”. It is also reasonable to suppose that as well as fights as described by SHARMAN & CALABY, lower intensity interaction such as cuffing and snorting may occur between 60‘ as they do between 60“ and 99 in non-sexual encounters.

Response to Aggression Avoiding Aggression-an animal moved quietly away from a threatening

animal with no other response. The threatened animal moved aside to allow the aggressor to approach food, or moved from a patch of shade or sun, into which the aggressor moved. Fleeing, in the sense of one animal running away from another, usually being chased, was not seen.

Aggression. Less frequently, some resistance to the aggressor, in the form of snorting andlor cuffing was offered. Sequences such as “A snorted a t B, B snorted at A, A cuffed at B, B moved away” were seen. The initiator of the sequence was not necessarily successful.

396 ELEANOR RUSSELL

Non-Agonistic d-9 Interactions 3 patterns were described by SHARMAN & CALABY (1964): (a) Investi-

gation, (b) Following, (c) Mating. Only the first 2 are considered here. (a) Investigation. The d approached a 9, and smelt round the pouch

opening and urogenital opening. dd might also smell the ground where a female had urinated, or the stream of urine itself.

(b) Following. The 6 followed close behind the 9, from time to time grasping a t her tail, and she then moved away. SHARMAN & CALABY say that dd begin to follow 9Q closely some 2 hours before the mating act occurs.

(b) MotherlYoung Interactions MotheriYoung interactions include elements which may be classified as

Approach/Retreat, Agonistic or Non-Agonistic, etc., bu t this has not been done. The association between a 9 and her young has been divided into 5 periods, based on the stages of development of the young; the behaviour of 9 and young is described within each period. 1. Young entirely within the pouch all the time. Birth to ca. 150 days. 2. Young in pouch all the time, head occasionally out of pouch. 150- ca.

190 days. 3. Young out of pouch for short periods. 190-235 days. 4. Young permanently out of pouch, still suckling from mother. 235-365

days. 5. Young no longer suckling from mother. 365 days on.

I . Young entirely within the pouch all the time. Nothing could be seen of the young during this stage, except that move-

ments of the young within the pouch were seen as movements of the ventral wall of the pouch. The mother periodically cleaned in her pouch as already described (fig. 3 (c), fig. 4). It could not be seen whether she licked the young, but this can be inferred from the fact that young in the pouch were generally quite clean, and that urine and faeces from the young were generally removed. The skin of the young was kept moist, probably by a combination of licking by the mother and secretion from the pouch. QQ with pouch young showed a higher frequency of pouch cleaning than did QQ with no young. The figures given by SHARMAN 8r CALABY (1964) are for one only, but the present work confirmed this observation. Some QQ kept their pouch and young much cleaner than did others.

The movements of the mother must, of course, be transmitted to the young, much more than would be the case in Eutherian mammals; especially any grooming which involves touching the part of the abdomen near the pouch. During this time, there is no way of knowing from observations alone whether or not the young is permanently attached to the teat.

2. Young in pouch, head occasionally out of pouch ca. 150 - ca. 190 days. In addition to the types of interaction described above, QQ were often

seen to lick any part of the young which projected from the pouch - head, feet, area near urogenital aperture. From about 150 days on, the young had its head out of the pouch for increasingly long periods, but as the young was rapidly becoming larger, other parts of the body were also likely to project from the pouch when the young changed position. Young a t this stage had


a light covering of fur. They began to crop grass or pick up food from the ground while still entirely in the pouch. They could now see their mother and were seen to sniff a t her head, especially round the muzzle, when she bent her head towards the pouch.

3. Yormg out of the pouch for short periods ca. 190- ca. 235 days All previously described interactions, in addition to new ones which

occurred when the young was leaving, entering, or out of the pouch. a) Leaving the pouch. The young might actively leave the pouch - this

was more common towards the end of pouch life. In this case, they would dive out head-first when the mother was standing up, or climb out when she was lying down on one side, and the mouth of the pouch stretched open.

Young might also fall out of the pouch when the mother was lying down or if they stretched out too far when feeding from the ground. This was less common in older young, which may feed when outside the pouch.

Young were sometimes tipped from the pouch by their mothers. Typic- ally, when this happened, the 9 began to clean in her pouch, holding the mouth of it open with her fore paws. A rapid contraction of the front wall of the pouch was seen, the mouth of the pouch relaxed, and the young was tipped out. In the normal course of pouch cleaning, the position of the 0 made it unlikely that the young would fall out of its own accord. After the young was tipped out, the 9 continued to clean her pouch.

b) Entering the pouch. The mother stood crouched while the young groped for the pouch opening, held the rim with its fore paws while it put its head in, dived in head first and then somersaulted so that the head was near the opening. The mother might incline her hind legs backwards from the vertical, bringing the pouch opening closer to the ground. (A similar description of entering the pouch is given by SHARMAN and CALABY 1964). Entering the pouch seemed always to be initiated by the young. Any sudden stimulus which alarmed the young might initiate the return to the pouch, but on most occasions when the young re-entered it was impossible to identify any external causal factor.

A young returning to the pouch when not alarmed approached its mother slowly from in front, and if she stood crouched properly, it could get into the pouch as described, alone. If the mother was lying down, the young might approach her, nuzzle round the pouch opening, and perhaps scratch round the mouth of the pouch with its fore paws, as i t does when feeling for the pouch opening when the 9 is standing. The mother inight then stand and allow the young to get into the pouch, or she might remain lying down, in which case the young could get into the pouch, and might lie down near her, or go away again. When the young was alarmed, it dashed quickly back to its mother, and got into the pouch very quickly if she was standing. If the mother was not also alarmed she might remain lying down and the young could not get into the pouch, but remained near its mother.

c) When the young is out o f the pouch. For about one week after it first leR the pouch, the young left the pouch for only short periods of a few minutes, and generally did not go more than about 5 feet from its mother. The young was still very shaky on its feet, and did not do very much while it was o u t of the pouch. I t moved about close to its mother, and perhaps sniffed at a few objects - trees, the ground, etc. AS the young became steadier on its feet,

398 ELEANOR RUSSELL

and stayed out of the pouch for longer, it became more adventurous. It generally stayed close to the Q, but made brief excursions away from her for a much greater distance, and then came rushing back. I t investigated more things, and ate leaves, grass, pellets and hay. This was done mostly in company with its mother - where she moved, the young followed. If the young was out of the pouch when the 9 went to the food bin, the young followed, and stood beside or behind her, and might eat also. When the mother was lying down, the young of this age was usually to be seen lying very close to her, in front or behind, with heads only a few feet apart, and bodies probably touching

If something alarmed the animals, so that the mother moved away rapidly when the young was out of the pouch, the young followed. Recog- nition of the mother in these circumstances was not very specific. If there was a group of adults, the young might follow one other than its mother, and try to get into her pouch. The stranger then cuffed and snorted at it, and pushed it away. The young might then stand and produce what can best be described as a distress call - a loud sibilant squeak, which brought its mother towards it. (The mother must recognise her young as i t does not seem able to recognise her.) The mother, in this circumstance, made a clucking noise, rather louder and harsher than that made by dd during sexual following. The young might make the same distress call if it went away from its mother, and could not see her to go back to. The distress call might attract the attention of other 99, which have been seen to approach the young, and smell them. The strange 99. might then attack the young, by pushing them away, or may move rapidly away, as from an alarming stimulus.

Early in this stage, the young was not suckled when it was outside the pouch. Later the young might be suckled when outside the pouch, and presum- ably also when inside. During the later stages of pouch life, and also after the young had left the pouch, suckling was initiated by the young. The young might be seen to approach its mother who was lying down, and “pester” her. The young sniffed around the pouch, scratched round the opening with its fore paws, tried to get its head in, and also sniffed at the head of the mother and pushed a t her head with its fore paws. This might continue for as long as 5 minutes, after which the young might stop and go away, or lie down again beside its mother, or else the mother might stand up, and the young then started to suckle from the pouch. Young which were young enough to get back into the pouch might do so after such behaviour, instead of suckling. In young which were permanently out of the pouch, such behaviour was followed by suckling.

When her young was suckling from the pouch, the female stood crouched, generally with her fore paws beside the young, or on its back. Suckling was not always continuous, especially when the young was older. It might take its head out of the pouch for a few seconds for no obvious reason, or if some noise occurred. The young was not necessarily suckling all the time its head was in the pouch, but this was not measured. After the young took its head out of the pouch, it generally stood beside its mother and licked its fore paws and rubbed them on its mouth and under its chin. Similar grooming movements may be seen a t other times.

A complex interaction between mother and young occurred most commonly after the pouch young or young-at-foot finished suckling, but was also seen a t other times. In its simplest form mother and young sniffed at each

(fig. 2 , fig. 6 [cl).


other in the region of the mouth, sometimes jus t nose-to-nose, at other times each sniffed all round the head of the other (Fig. 6 (a), Fig. 8). In addition, each might spar ‘it the head and shoulders of the other with fore paws; or they might stand fairly erect, while each clasped the shoulders of te other, and heads were rubbed together and moved from side to side (Fig. 6 (d)). The significance of this behaviour is not clear. I have seen it between females and weaned young up to 2 years old, and I have also seen it between two young- at-foot of different mothers.

Fig . 8: Interaction between Q and young-at-foot. Shortly afterwards, the Q stood up, and the young began to suckle from her pouch

4 . Young permanently out of the pouch, still suckling from i ts mother 235-365 days. Except that the young did not get back into the pouch, its behaviour was

not very different from that of the previous stage. The young leaves the pouch permanently about 235 days after birth.

According to SHARMAN and CALABY (1964) the duration of pouch life is not dependent on whether the 9 is carryinga delayed blastocyst which has assumed development SO that she is about to give birth. Nevertheless, at some point on or about day 235, the 0 refuses to allow the young back into the pouch after it has left, or she may tip it out and refuse to allow it back.

The young was seen to be, by now, considerably more independent of its mother. I t went further away from her for longer periods, but returned to her when alarmed and to suckle, and still followed her about for a large part

400 ELEANOR R U ~ S E L L

of the time. If an adult 9 was seen to be lying down with a young animal close beside her, it was almost certainly her own young. Suckling was less frequent, and young ate more hay, grass and pellets.

If young of this age approached too close to older animals in the group, they were chased in the same way as adult animals. Young animals seemed to have to learn not to go too close to other animals, in particular the 6 of a group. I t was rare to see an adult 9 go close enough to a 6 for him to chase her away, but young animals were seen to approach the 6 and sniff a t him, and were quickly chased away.

Young-at-foot were weaned by about 365 days, by which time they weighed about 10 kg. As the young-at-foot reached the age of weaning, it fed from its mother less frequently - feeding was seen only 2 or 3 times in 24 hours. The young-at-foot might approach its mother several times, and “pester” her to stand up, but she ignored it, or moved away, or even pushed the young away when it tried to get its head into the pouch. Finally, the Q would not allow the young to suckle at all. There followed a few days when the young attempted to suckle and the mother would not permit it. After this, the young seemed to stop trying. Times given for weaning, etc. must be treated with some suspicion.

O n one occasion, a young-at-foot was recorded as suckling from its mother, and was recorded as having its head in the 9’s pouch for several minutes on successive days. However, when the pouch was checked, it was found that the 9 had stopped lactating some days earlier, and so could not have been feeding the young-at-foot.

5 . Young no longer suckling from their mother All patterns seen in the previous stage, except suckling from the mother,

were again seen. At the beginning of this stage the young was still to be seen frequently associated with its mother, following her about, lying near her, feeding with her, etc. The amount of time spent with the mother descreased, but 8 months after weaning, young were still seen near their mothers, and occasionally the pattern of sparring was seen. 99 rarely showed any aggression towards their own weaned young, at least within 12 months of weaning. Young were tolerated within a smaller distance than unrelated adults. They might be chased away if they attempted to put their head into their mother’s pouch, but might still feed and drink close to their mothers. Young 9 red kangaroos reach sexual maturity at 15-20 months and dd a t about 2 years (SHARMAN and PILTON, 1964). Whether some kind of relationship between mother and young persists as long as this is not yet known.

Vocalisations 1 . The “soft-clucking” of the d emitted during sexual following and

displays (SHARMAN and CALABY, 1964). 2. The distress call of the young - a high-pitched sibilant squeak, made

when the young was separated from its mother, a t least up to the time of leaving the pouch, and probably longer.

3. The “loud-clucking’’ made by a 9 when calling a young which was calling her. I have also heard this sound from hand-reared joeys in two circumstances before they have reached normal weaning age: when they were disturbed by strangers, and when they were fed, especially if they were


hungry. A vocalisation which sounded identical was sometimes heard from adults when they were caught in the yards and handled for any reason.

4. The aggressive ‘‘snort” described above. These vocalisations were rarely heard in normal circumstances.

Discussion Since behaviour has been described only qualitatively, few conclusions

can be drawn as to the motivational significance of particular elements. But recognition of individual elements permits later quantitative description, and the determination of relationships of these elements with each other and the environment.

The daily pattern of activity in the yards does not differ greatly from that destribed by CAUGHLEY (1964) for red and grey kangaroos. There is the same low-level activity throughout the day, and the animals never seem to be completely asleep. The time of beginning of the period of increased activity is placed by CAUGHLEY as approximately 1 hour before sunset. For the Euro (Macropus robustus) EALEY (1967) places this time as early as 1600 hours in winter, and after sunset in summer. Animals in yards in Sydney generally became more active about 1 hour before sunset in summer and winter. The high daily maximum temperatures of the area in which EALEY worked (Pil- bara district of Western Australia) may help to explain the late resumption of activity in the summer. GRIFFITH and BARKER (1967) note that in S.W. Queensland, in the cooler months, feeding starts at about 1630 hours and goes on till about 2100 hours, while in summer, with shade temperatures of 120’ F, feeding does not start till about 2000 hours.

Of the various postures described for the animals when stationary and generally inactive, those of most interest are the various recumbent positions, and “standing crouched” (i. e. with forelimbs on the ground). The location of these, in sun or shade, their duration and frequency relative to each other and to other activities during the day, and the relation of this picture to the environmental conditions at the time may help to explain the survival of the animal in very hot, dry conditions.

Licking of forelimbs, hindlimbs and abdomen, and profuse salivation have been observed as a response to increased temperature by several authors (ROBINSON 1954; HIGGINBOTHAM 8c KOON 1955; BARTHOLOMEW 1956; MOR- RISON 8c ROBINSON 1957), and suggested by them as a means of temperature regulation. BENTLEY (1960) found that the prevention of licking of fur had no effect on body temperature in the quokka, Setonix brachyurus, and DAWSON (1969) says that while licking occurs a t high temperatures in the brushtail possum (Trichosurus vulpecula), its contribution to temperature regulation is negligible. Any contribution which licking may make to temperature regulation in the red kangaroo has not been measured. HEDIGER (1 958) mentions the behaviour, and quotes heat regulation, nervous excite- ment and displacement activity as its possible significance. The significance of digging behaviour is likewise obscure. EALEY (1967) mentions the occurrence of “scratchings” in shady areas inhabited by Euros, but gives no suggestion as to their function. Initially, it may remove stones and spiky grass from the area, but this soon ceases to be necessary. It may remove warmer surface soil SO that the soil on which the animal lies is relatively cool. In winter this would not be necessary, a t least in Sydney. I t may merely be more comfortable for the animal to lie in a hole than on a flat surface. There may no longer be any functional significance a t all.

Z. f . Tierpsychol. Bd. 2 7 , Heft 4 26

402 ELEANOR RUSSELL

How animals feed and drink in captivity is of doubtful relevance to such activities in the wild, but the daily pattern of such activities may be signific- ant. GRIFFITH and BARKER (1967) found that in a good season, kangaroos were solely grazing and not browsing, but the ability of the red kangaroo to eat all available plant material within reach in yards suggests that when food is scarce in a natural environment, the red kangaroo is able to take far greater advantage of what is available than, for example, a sheep.

CAUGHLEY (1964) suggests that although red and grey kangaroos are found in groups, the size of groups is determined by an essentially random process of animals joining and leaving. He says that both species show few of the attributes of sociality usually found in gregarious animals, although they do form groups. H e recognizes that mother and offspring continue to associate for a time after weaning, and that juveniles may call when tem- porarily separated from their mothers, but could find nothing directly comparable with an alarm call, and did not observe any kind of hierarchy of dominance within groups. In the yard colony, there is certainly an alarm signal, but i t is not a vocalisation. Alarm signals of this nature are not unknown; the thumping of lagomorphs is a similar signal. HEDIGER (1958) also reports the slapping of the hind limb as an alarm signal for kangaroos in captivity. The variety of interactions observed between individuals permits a dominance hierarchy to be distinguished in these artificial groups of animals. This will be treated quantitatively in a subsequent paper, but it can be said here that the dominant animals, usually older and larger animals, have precedence at food, water, shade and shelter from rain. EALEY (1967) says that a social hierarzy based on size exists in the Euro. H e bases this on observations in regard to water at a time when water was scarce. Larger animals displaced smaller animals with a low gutteral hiss, which EALEY terms a “challenging” sound.

There is a long period of association between mother and offspring, first in the pouch, and then as young-at-foot before and after weaning. The period of association aRer weaning recorded by EALEY (1967) and which I have observed in yard groups is substantiated by the numbers of groups of 2 and 3 recorded by CAUGHLEY (1964) and KIRKPATRICK (1966). SHARMAN and CALABY (1 964) described Mother/Young relationships briefly, and a few more observations have been added here, but there is not yet sufficient quantitative information available to enable much valid comparison to be made with eutherian Mother/Young relationships. The mother’s readiness to show maternal behaviour which is never very great, decreases as the young gets older. Nevertheless, after the young leaves the pouch it may be said that it becomes increasingly responsible for maintaining proximity to its mother. This a t least is comparable with Eutherian mammals such as cats (SCHNEIDER, ROSENBLATT and TOBACH 1963) and dogs (RHEINGOLD, 1963) with rather mobile young. Young-at-foot or pouch young out of the pouch for a short time, who try to get into the pouch of a 9 with no young or a small pouch young, are often rejected violently, but young can be transferred to the pouch of a 9 with a young of approximately the same age, and successfully reared. MERCHANT and SHARMAN (1967) report transfers made of young of Megaleia rufa from less than 1 day to 215 days to 99 of the same species, and successful transfers of Megaleia young to grey kangaroo Macropus giganteus foster mothers have also been made. The MotheriYoung relationship does not appear to be very specific.


Acknowledgement This work was i n par t supportcd by a grant from the Aus t rd ian Rcscarch Grants Com-

mittee.

Summary

1. 2 groups of red kangaroos (Megaleia Y U ~ U ) were watched in yards for a total of 126 hours (29 animals), and all behaviour recorded. A qualitative description is given of all but reproductive behaviour.

2 . Activity increased a t dusk and early in the morning. In summer, activity began to increase later in the afternoon and to cease earlier in the morning. During the day, the animals spent most of the time either lying down or standing crouched in the sun (winter) or shade (summer). Nevertheless, a full range of behaviour was seen during the day, but a t lower frequencies.

3. In the course of grooming, the animals could reach every part of their body with fore- and hindlimb, teeth and tongue. 99 clean in their pouches whether or not they have a pouch young, but more frequently when a young is present. The tail-forward posture adopted by the 9 while giving birth is similar to the tail-forward position used when the tail is groomed.

4. A number of Non-agonistic and Agonistic Approach/Retreat interactions between two individuals are described. These were generally seen when one animal chased another from place in the shade or sun or a t the feed-bin, or when one animal approached another already established in sun or shade or a t the feed-bin.

S. The association between a 9 and her young is divided into S stages based on the delopment of the young from birth. These stages are marked by the young's increasing independence of its mother, bu t some association between mother and young can be seen after the young is weaned when about 1 year old.

Zusammenfassung

(1) Zwei Gruppen (29 Tiere) des roten Kanguruhs (Megalcia Y U ~ U ) wur- den im Gehege uber eine Gesaintzeit von 126 Stunden beobachtet. Eine qualitative Beschreibung des Verhaltens mit Ausnahme des Fortpflanzungsver- haltens, wird gegeben.

(2) Die Aktivitat nahm am spaten Nachmittag und in den Morgenstunden zu; im Sommer begann sie am Nachmittag spater und horte fruher am Morgen wieder auf. Die meiste Zeit des Tages verbrachten die Tiere entweder liegend oder kauend, in der Sotine (Winter) oder iin Schatten (Sommer). Dennoch war - wenn auch mit groflen Pausen - w5hrend des Tages das volle Ver- haltensinventar zu beobachten.

( 3 ) Die Tiere putzen jeden Teil ihres Korpers mit den Vordergliedern, Hintergliedern, Zahnen oder mit der Zunge. Die Q9 reinigten auch ihren Beutel, und zwar wenn er ein Junges enthielt, haufiger. Die Nach-Vorn- Schwanzhaltung des 99 wahrend der Geburt ahnelt der Schwanzhaltung beim Putzen.

(4) Eine Anzahl nicht-agonistischer und agonistischer Begegnungen zwischen zwei Tieren mit Annaherung und Ruckzug werden beschrieben, meist, wenn ein Tier sich einem anderen naherte, welches sich bereits in der Sonne, dem Schatteii oder an der Futterkrippe niedergelassen hatte, oder es gar von cinem solchen Platz verdrangte.

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(5) Die Beziehung zwischen dem 0 und seinem Jungen kann in 5 Phasen nach der Entwicklung des Jungen eingeteilt werden. Diese Phasen sind ge- kennzeichnet durch die immer grot3er werdende Unabhangigkeit von der Mutter. Eine gewisse Verbindung zwischen Mutter und Jungem besteht noch nach der Entwohnung, wenn das Junge etwa ein Jahr alt ist.

Li terature cited BARKER, S., G. D. BROWN EL J. H. CALABY (1963) : Food Regurgitation in thc Macropodidae

Aust. J. Sci. 25, 430-432 BARTHOLOMEW, E. A., (1956): Temperature regulation i n the macropod marsupial Setonrx brachyurus. Physiol. 2001. 29, 26-40 * BENTLEY, P. J. (1960): Evapora- tive water loss and temperature regulation in the marsupial Setonix brachyurus. Aust. J. exp. Biol. 38, 301-306 CALABY, J.H. (1958): Studies on marsupial nutrition. 11. The rate of pas- sage of food residues and digestibility of crude fibre and protein by the quokka, Setonix brachyurus (Quoy and Gaimard). Aust. J. biol. Sci. 11, 571-580 CAUGHLEY, G. (1964): Social organization and daily activity of the red kangaroo and the grey kangaroo. J. Mam- mal. 45, 429-436 EALEY, E. H. M. (1967): The biology of the euro, Macropus robustus (Gould), in north-western Australia. 11. Behaviour, movewents and drinking patterns. CSIRO Wildl. Res. 12, 27-51 FRITH, H. J. (1964): Mobility of the red kangaroo, Megaleia rufa. CSIRO Wildl. Res. 9, 1-19 9 GRIFFITHS, M. gi R. BARKER (1967): The plants eaten by sheep and by kangaroos grazing together in a paddock in south-western Queensland. CSIRO Wildl. Res. 11 , 145-168 HEDIGER, H. (1958): Verhalten der Beuteltiere (Marsupialia). Handb. 2001. 8, 10 (9), 1-28 9 HIGGINBOTHAM, A. C. gi W. E. KOON (1955): Temperature regulation in the Virginia opossum. Am. J. Physiol. 181, 69-71 KIRKPATRICK, T. H. (1966): Studies of Macropodiae in Queensland. 4. Social organization of the grey kangaroo (Ma- cropus giganteus). Old. J. Agric. Anim. Sci. 23, 317-322 * MERCHANT, J. c., gi G. B. SHAR- MAN (1966): Observations on the attachment of marsupial pouch young to the teats and on the rearing of pouch young by foster-mothers of the same or different species. Aust. J. Zool. 14, 593-609 MOLLISON, B. C. (1960): Food regurgitation in Bennet’s wallaby, Proternnodon rufogrisea (Desmarest) and the scrub wallaby, Thylogale billardieri (Desmarest). CSIRO Wildl. Res. 11, 145-168 NEWSOME, A. E. (1965): The distribution of red kangaroos, Megaleia ruja (Desmarest) about sources of persistent food and water in Central Australia. Aust. J. 2001. 13, 289-299 RHEINGOLD, H. L. (1963): Maternal behaviour in the dog. In: Maternal Behaviour in Mammals (H. L. Rheingold ed.) New York: John Wiley, 169-202 ROBINSON, K. W. (1954): Heat tolerances of Australian monotremes and marsupials. Aust. J. biol. Sci. 7, 348-360 ROBINSON, K. W. et P. R. MORRISON (1957): The reaction to hot atmospheres of various species of Australian marsupials and placental animals. J. cell. and comp. Physiol. 49, 455-478 SCHNEIRLA, T. C., J. S. ROSENBLATT ~r E. TOBACH (1963): Maternal behaviour in the cat. In: Maternal Behaviour in Mammals (H. L. Rheingold ed.). New York, John Wiley, 169-202 SHARMAN, G. B., ~r J. H. CALABY (1964): Reproductive behaviour in the red kangaroo, Megaleia rufa, in captivity. CSIRO Wildl. Res. 9, 58-85 SHARMAN, G. B. sr P. E. PILTON (1964): Life history and reproduction of the red kangaroo (Megaleia rufa). Proc. 2001. SOC. Lond. 142, 29-48.

Appendix DAWSON, T. S. (1969): Temperature regulation and evaporative water loss in the

brush-tailed possum T~ichosurus vulpecula. Comp. Biochem. Physiol. 28, 401-408.

Author’s address: Eleanor Russell, University of New South Wales, Box 1, Post Office Ken- sington, 2033, School of Zoology, Sydney, Australia.

Observations on the behaviour of the red Kangaroo (Megaleia rufa) in captivity

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