Contents. Page 1. Introduction Why choose the ringtailed lemur? Background information on ringtailed lemurs. 5

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2 Contents Page 1. Introduction... 3 Contents of the pack, objectives 2. Why choose the ringtailed lemur? 4 3. Background information on ringtailed lemurs Methods of recording the behaviour of ringtailed lemurs. 10 Preparation for the zoo visit, continuous and instantaneous methods of recording observations 5. Descriptions of the behaviours that might be displayed 11 Feeding, resting, moving, grooming 6. Analysis of results 14 Continuous and instantaneous recording 7. Discussion of results Research synopsis.. 15 Time budgets, feeding, grooming, moving, resting 9. Acknowledgements Bibl iography Useful addresses Student worksheets.. 18

3 Studying Animal Behaviour in Zoos Recording and Scoring Selected Behaviours of Ringtailed Lemurs 1. Introduction What better way to stimulate and enthuse students about the study of animal behaviour than a day visit to a zoo! There are 61 centres (1991) in the Federation of Zoological Gardens of Great Britain and Ireland, so schools and colleges will not be far from one. Increasingly, zoos welcome organised groups of students as their research, particularly when set up in co-operation with zoo staff, can add to the knowledge and understanding of the species in the zoo collection. This ASAB project is designed to provide teachers with sufficient information to confidently guide students through the scientific process involved in collecting reliable data, analysing the outcomes and evaluating it in terms of previous research. Materials available in this pack background information on ringtailed lemurs two suggested methods for gathering data by observation, with instructions for students recording sheets for each of the methods suggestions for graphically representing and analysing the data selected topics from research papers to extend these studies. Materials available on loan (from the ASAB Education Officer address on page 15) a ten minute video, illustrating the types of behaviour displayed by ringtailed lemurs a selection of relevant articles from biological and psychological journals. Objectives In carrying out this study of animal behaviour, students should be able to: gather some information about the life history of the animals being studied recognise different categories of behavioural activity learn how ethograms (descriptions of the behaviours a species exhibits) are constructed carry out a focal sample to observe the behaviour of individuals record their observations by continuous and instantaneous methods present their results in a graphical form and statistically analyse them, where appropriate discuss their results and relate them to previous research.

4 The materials presented here could be incorporated into the 2001 Advanced Level Biology courses (in particular the Biological Basis of Behaviour module in the NEAB modular course). The new specifications for A level Biology produced by AQA and OCR retain an optional module incorporating behaviour studies in the second year of the course, so this project would be eminently suitable for coursework. In the EDEXCEL specification, the opportunity to undertake animal behaviour studies could be utilised to complete the individual practical study. Indeed, some papers in Animal Behaviour and other (carefully selected) journals could be understood by able sixth form students and used as references when writing up their work. The proposed study is also suitable for supporting topics in the Scottish Advanced Higher Biology Animal Behaviour module. A level Psychology students could also use these materials to support one of their two pieces of coursework, for example, to illustrate aspects of sociality, or the signalling systems used in non-human animals. The new specifications for AS and A2 level Psychology include topics such as allogrooming, stress in captive animals, social learning and animal communication, all topics which could be incorporated into coursework by using the methods described in this project. 2. Why choose the ringtailed lemur? The animals that are the focus for this study are ringtailed lemurs (Lemur catta). The reasons for this choice are: they are hardy primates and found in most zoos in sufficiently large group sizes for study they are instantly recognisable to students as a species (although differences between individuals in a group are quite difficult to spot) they are diurnal creatures, so they will be evident when students visit the zoo whatever the ringtailed lemurs are doing at the time of observation, data can be generated, i.e. the animals are always behaving there is quite a body of documented research into their behaviour, both in the wild and in captivity (some of this research is available with this pack of materials) they are highly active animals they live in social groups, so interactions between individuals can be studied as a species, they are appealing to humans there is the possibility, in some zoos, for humans to interact with the animals in open exhibits there is the possibility, in some zoos, to study interactions between species (in Blackpool Zoo, three species of lemur occupy a shared area).

5 3 Background information on ringtailed lemurs (Lemur catta) This information could be used by a teacher to become an instant expert on the animals being studied, or as a resource for students to use in preparation for their study. Natural habitat Lemurs are confined to the island of Madagascar, which has been separated from the southeast coast of Africa for about 120 million years. These lower primates or prosimians were able to evolve on the island continent in isolation from higher primates, such as monkeys and apes, because they were not subject to the same levels of competition and predation suffered by mainland primates elsewhere. Until about two thousand years ago, when humans colonised the island, there were over 40 species of Malagasy Lemuriformes, ranging from some as small as shrews to others the size of goats. The destruction of forested areas, hunting and the introduction of domestic species resulted in the extinction of at least 14 species of lemurs. The 32 species that survive today are still threatened by the continuing loss of their habitats, chiefly forest. Classification Kingdom Phylum Class Order Family Genus Species Animalia Chordata Mammalia Primate Lemuridae Lemur catta The primates are divided into two main groups, the prosimians, which includes lemurs, and the anthropoids, which include monkeys, apes and man. Lemurs are easily distinguished from true monkeys by their fox-like muzzles and their very large, but inexpressive eyes. Adult ringtailed lemurs have a mass of kg and they are m in length, including their long, ringed tail. Their forelimbs are longer than their hind limbs and their fur is greyish, soft and woolly.

6 Day-time creatures with night-time origins Most lemurs are adapted for night vision by having a reflective layer (tapetum lucidum) behind the retina, ensuring that the maximum amount of light is detected. This feature, along with their tendency to call loudly at night, resulted in their name lemur meaning ghost. Lemurs have become less nocturnal and more active during the day, presumably due to the lack of predators (other than man) on their island habitat. The ringtailed lemur is diurnal. Communication Unlike higher primates, prosimians have a highly developed sense of smell, which plays an important part in communication as well as being used to find food. Lemurs have genital scent glands, which they use to mark branches of their territories. Males also have a functional secretory gland on the flexor surface of each forelimb, which they use to smear their tails to wave at potential rivals or to indicate sexual arousal. Tail flicking, in which the tail is moved from a vertical position forward over the head of the individual, is an effective method of distributing scent. Male tail displays are thought to act as stimulants for the neuro-endocrine mechanisms controlling ovulation. Ringtailed lemurs emit a wide range of vocalisations including purring (like cats) and yapping (like terriers). Feeding The ringtailed lemur is the only lemur, which spends most of its time on the ground. This enables the ringtailed lemur to exploit a wider range of dietary items, including leaves, fruit and flowers and to hunt for insects in dead wood and termite mounds. In contrast, other lemurs are herbivorous. Unlike higher primates, lemurs rarely use their paws to manipulate food items, though leaf-bearing branches may be pulled closer to their mouths to feed from directly. Moving Most lemurs have a long, bushy tail, which is used as a counter balance when leaping from branch to branch in their arboreal environment. The ringtailed lemur is a semi-terrestrial quadruped, spending most of its time on the ground, unlike other species of lemur, which spend most of their time in the trees. The palms of the paws and soles of the feet are leathery, providing effective grip on slippery rocks. It moves through the trees by climbing up large oblique branches and leaping down to others, rather than using narrow terminal branches at the tops of trees and leaping to adjacent trees as seen in other species of lemur.

7 Grooming Unlike higher primates, prosimians use their teeth, including their special, comb-like lower incisors, for grooming as shown on the skull below 1. Allogrooming, where one animal grooms another, ensures that parts of the body not accessible during self-grooming are reached to keep the animal s coat free from parasites and dirt. However, it also serves a social function; it is the commonest form of interaction between members of a group. In many species, other females in the group regularly groom mothers of newborn infants, perhaps enabling access to the infant. Grooming often occurs in bouts; one animal grooms another, then stops and the roles are reversed the groomee becoming the groomer and so on. Affiliation (social bonding) and appeasement (submission to a dominant individual in order to reduce aggression) are both reinforced by social grooming in many species of primates. Resting To conserve heat at night, ringtailed lemurs curl up so that none of their underside is exposed. Huddling, in which the tail is wrapped around the body, reduces the surface area available for heat loss. 1 Photograph kindly provided by Dr. M. Hansell

8 Resting activity during the day may be related to daylength and temperature; sunning (in which the animal exposes its underside to the sun) may be an important thermo-regulatory mechanism. During the day ringtailed lemurs spend about five times as long resting as engaged in any other activity. Presumably, this is linked to their diet; other lemurs spend less of their time resting. Social organisation The ringtailed lemur lives in social groups of about twelve individuals, but this can range from five to thirty. Unusually for primates, females are dominant over males, although the group does not have a single leader. Hierarchy among the males is maintained separately by frequent threats and displays between individuals and occasional fighting. Subordinate males may be pushed to the periphery of the group where, in an arid environment with sparse food, exclusion of excess males could be an adaptive advantage for the whole group. At the top of the hierarchy, successful males are able to retain their aggressiveness and thus provide better protection for a large group of individuals moving on the ground over a fairly large territory (6 23 ha). Females and their female descendants form the social core of the group, only leaving in order to start a new group. Females define and defend the territory; males move between different groups, thus preventing inbreeding and effecting gene flow throughout the population.

9 Reproduction All species of lemurs are seasonal breeders (August November) in the wild (in zoos, breeding groups can be expected to produce offspring in April or May). Female ringtailed lemurs reach sexual maturity at months, give birth once a year and the gestation period is days. Multiple births are rare. The mating season lasts less than two weeks and within this time a single female is receptive to males on only one day. Oestrous is marked by swelling and pinkening of the external genitalia and by repeated presentation of the anogenital region to the male. Synchronisation of menstrual cycles ensures that all adult females in a number of adjacent groups become receptive at the same time. This synchrony is probably achieved by olfactory communication as ringtailed lemurs use their scent glands lavishly at this time. Care of the young The young are born at a relatively early stage of development. For the first 3 days the infant remains attached to its mother s breast, then it may ride on its mother s back. The infant is cared for by the whole group, being suckled by any lactating female for about 5 months. This altruistic behaviour is a common feature of the complex social behaviour of higher primates, enabling other females to learn the skills of parenthood. Group support increases the survival chances of the young and ensures that their alleles are successfully passed on to the next generation.

10 4. Methods of recording the behaviour of ringtailed lemurs Preparation for the zoo visit Students should be encouraged to take their cameras with them as it is relatively easy to photograph at least some of these behaviours whilst the lemurs are in the outdoor enclosure. A video camera would be a particularly valuable piece of equipment to take on the visit, provided students have had previous training or experience in its use. One difficulty students encounter when they first attempt to study animal behaviour is the difficulty in identifying individuals within a group. In some zoos the animals may be tagged with coloured or numbered discs to aid identification. Another difficulty students face when conducting observations of animal behaviour is defining the behaviour in such a way that it can be unambiguously recognised by all observers. Many hours of preliminary observations are required to categorise and define descriptions of behaviours and then construct an ethogram describing these behaviours so that other observers can use them accurately. It would take students the whole of their visit to reach this stage so these materials will enable students to concentrate on the gathering of reliable observational data, using ethograms already prepared. In this study, following preliminary observations, it was decided to focus on four easily distinguishable aspects of a ringtailed lemur s behaviour: feeding, moving, grooming and resting. Definitions of these behaviours are provided (see section 5) and could be discussed with students prior to the zoo visit. A ten-minute video recording of ringtailed lemurs engaged in the four activities is available on loan from the ASAB Education officer (the address is provided on page 15). Close examination of the animals behaviour reveals the complexity of each activity and illustrates how ethograms can be constructed. Observations and recording of animal behaviour On a day visit to a zoo, bearing in mind that other exhibits may wish to be visited, the time available for observations of animal behaviour is likely to be limited. This material is designed to give students the opportunity to experience two different types of recording method in the space of about an hour. 1 Continuous recording The continuous method will enable students to focus first of all on the four easily distinguishable activities listed above. They should focus their attention on the behaviour of a single individual for a period of about thirty minutes and simply record the amount of time spent on each activity on the recording sheet (1a) provided. If the individual being observed should go out of sight, the observation period ceases 9though it is hoped that students will be fortunate enough to observe one individual for minutes). During this period of observation they should notice that each activity can be further defined and this could lead to a discussion of the formation of ethograms.

11 2 Instantaneous recording Students can then experience instantaneous recording of observations using prepared ethograms. The detailed recording sheets (2F, 2G, 2M and 2R) for this part of the study have been designed as simple tick boxes for ease of completion by the students. In this case, students record in as much detail as they can (by completing the appropriate tick boxes on a horizontal line on one of the recording sheets) exactly what their individual lemur is doing at each thirty-second interval for up to about thirty minutes. This will provide a wealth of valuable data and illustrate that apparently simple methods of recording can yield highly useful data. One difficulty, which students working in groups may encounter, is timing the instant of recording accurately. This could be overcome by recording bleeps at 30 second intervals for 30 minutes onto an audio cassette tape. One member of the group could replay the tape on a personal cassette player, releasing both hands for recording the information, whilst relaying the instant for recording to the rest of the group. 5. Descriptions of the behaviours that might be displayed So that all the students can unambiguously complete the tick boxes on the instantaneous recording sheets, detailed descriptions of all the terms used on the sheets are provided here. Individual ringtailed lemurs under observation should be identified as adult or juvenile (on the basis of size). Students are unlikely to observe differences between the sexes. However, in some zoos, male and female ringtailed lemurs are housed in separate exhibits, as breeding of these hardy primates in captivity has become too successful. The number of lemurs located in close proximity (less than 2 m away) to the lemur being observed should be recorded as the group size. Any other observations that might affect the behaviour of the observed lemur, e.g. sudden noise, presence of a keeper, interaction with other lemurs or humans, should be recorded in the notes section. The location of the lemur being observed should be categorised as follows: Inside in the indoor enclosure Outside in the outdoor enclosure Ground either inside or outside, at ground level Trees either inside or outside, in trees or branches/logs provided as an aboveground environment Furniture either inside or outside, on zoo furniture i.e. artificial platforms provided, such as feeding tables, visitor walk-ways, play equipment etc.

12 Activity Terminology Description/Definition Feeding obtaining food by grasping by paw or eating directly from the ground or branches Diet Water Plant Leaf Non-leaf Non-plant Body part used Mouth Paw Right Left liquids and solids consumed drinking from pools, water dispensers and other receptacles in the enclosure herbivorous diet tree leaves other vegetable/fruit/nut materials insects foraged within the enclosure the part of the body used to initially contact the food collecting food directly with the mouth using forelimbs to grasp/hold the food identification of limb used as above; both limbs could be used Activity Terminology Description/Definition Resting remain inactive; not engaged in any other activity that can be identified Posture All fours Haunches Forelimbs apart Forelimbs together Huddle the resting position adopted by the observed lemur inactive, lying horizontally with all four limbs on the ground adopts an upright sitting position; front paws may or may not be on the ground forelimbs abducted, usually when facing the sun forelimbs not abducted rest in a curled position either on its own or in a group Eyes Open Closed Vigilant pupils visible eyelids cover eyes although resting, the observed lemur s head is raise and/or ears are cocked; it is ready to react any sudden stimulus

13 Activity Terminology Description/Definition Moving locomotion of any description Mode of travel Walk Scamper Climb Leap the method used to move move slowly on all fours on the ground move horizontally on all fours quickly, on the ground or in the trees, play chase/charge/flee from other lemurs move vertically, up trees, branches or zoo furniture hop or jump on the ground; jumping between adjacent horizontal branches in the tree Tail Up Down tail held vertically, waved forward or over the head or swung from side to side in the air tail inactive Activity Terminology Description/Definition Grooming repeatedly stroke the fur using the dental comb and/or tongue Groomee Self Other Adult Juvenile Reciprocated Body part Head Torso Front Back Forelimb Hind limb Right Left Tail the lemur being groomed the lemur grooms itself the lemur being observed grooms another individual the groomee is full size the groomee is smaller than an adult mutual grooming which occurs simultaneously or in close alternation with a partner the part of the body being groomed front and back, including ears, face and muzzle chest and abdominal regions, including urinogenital areas anterior surfaces of the body posterior surfaces of the body upper limbs lower limbs identification of limb used as above any part of the tail

14 6. Analysis of Results 1 Continuous recording These results can be used to generate the percentage time that individuals spend in each activity. All the students should pool their results to complete the Summary table of Time Budgets (Sheet 1B) to generate a mean percentage time for each activity. These results could be presented as pie charts. It must be emphasised that the time of day when the observations take place is highly relevant and especially important for captive animals in indoor enclosures, as they may be fed at the same time each day. A comparison of time budgets for different individuals (male/female, adult/juvenile) could be made and an inferential statistical test could be used, if appropriate, to see if there is any significant difference within the data. 2 Instantaneous recording These results can be used to generate the frequency of occurrence of each of the behaviours under observation, which could also be presented as a pie chart for each individual. To see whether the frequency of one activity is engaged in significantly more frequently than another, the Chi-squared test could be used. Diagrammatic representation of data might be possible too. A level Psychology students may be familiar with the work of Jourard (1966) on human body accessibility by other humans. This technique could be adapted using a lemur-body outline (provided as a student worksheet see page 16) to illustrate the body parts that are most frequently groomed by the lemurs, and, perhaps, to which sex and age groups the groomer and groomee belong. 7. Discussion of Results Whilst completing the continuous method of recording observations, students may notice that each of the behavioural activities can be described in much greater detail, leading to the notion of constructing ethograms. The ethogram provided could be evaluated in terms of its effectiveness in covering the range of activities they observed. The wealth of data on the instantaneous recording sheets should prompt a wide-ranging discussion on the complex behaviour of ringtailed lemurs. Indeed the very location of the lemurs could trigger consideration of the proportion of time spent inside or out, and whether this is related to the time of day or year. Evidence for the terrestrial nature of ringtailed lemurs may be supported by comparing the relative amount of time spent in the trees and on the ground.

15 There has been a considerable amount of published research into the behaviour of these animals, both in the wild and in captivity. Selections from this research, relevant to the four activities studied in this project, are provided to stimulate further debate. Students can evaluate their results in the light of this research and suggest further studies that could develop from this work. 8. Research synopsis Time budgets Sussman (1964) completed individual activity records for ringtailed lemurs in the wild. He found that they spent much of the afternoon in unshaded areas of the forest, sunning to help maintain body temperature. In this way they could utilise habitats with large ranges of ambient temperatures. He also noted that they fed in two bouts during the day. Increasingly, zoos are working to provide enclosures in captivity which enable the animals to behave in as natural a manner as possible. Evidence from time budgets can be useful in judging their success. Feeding A study by Sussman (1974) of ringtailed (L. catta) and brown (L. fulvus) lemurs coexisting in the wild, showed that L. catta spent only 43% of its feeding time devouring leaves compared to 89% for L. fulvus. This difference could be related to the different habitats occupied by the 2 species. L. catta has a greater opportunity to exploit a wider range of foods as it moves through all the forest layers, whereas L. fulvus has a more restricted range. In Primate Societies, Richard refers to the work of Sheine who compared the efficiency of leaf digestion in L. catta and L. fulvus in captivity. The relative inefficiency of L. fulvus in digesting leaf material could be explained by it being outcompeted in the past by now-extinct species. Manning and Chamberlain (1990) investigated the left-side cradling preference in great apes. Their results indicate that this behaviour probably originated about 6-8 million years ago, in the common ancestor of African apes and humans. Observations of this behaviour in Old World monkeys could help to determine the origin of this behaviour. Students might observe preference in handedness in lemurs whilst they are feeding. Grooming In a study of a group of rhesus monkeys by Sade in 1965 described by Chalmers (1979), the frequency of grooming between related individuals was 4 times greater than that of non-relatives.

16 O Brien (1993) has reported that allogrooming in female wedge-capped capuchin monkeys has an affiliative function. Grooming sessions tend to be more frequent and longer in duration with equal balance between partners than in unreciprocated and appeasement grooming interactions. Moving Comparisons of the feet of 2 species of lemur (L. catta and L. fulvus rufus) were reported by Sussman in Prosimian Behaviour. In L. f. rufus the heel is covered in hair, whereas hair is absent from the heel in L. catta. The foot of L. catta closely resembles that of monkeys. Further evidence of the link between these morphological differences and the habitat preferences (ground and trees) of the 2 species could be obtained from prepared casts of the feet and analysis of their movement. In Social Behaviour in Primates, Chalmers describes the relative lack of independent mobility in the digits of lemurs. This supports the view that in evolution from the prosimians towards man, the digits become gradually more independent and more mobile. Resting Evans and Goy (1968) have reported that the occurrence of huddling in ringtailed lemurs could perhaps be linked to the strength of the affiliative bonds in the group. In the same paper they have reported that synchronisation of oestrous occurs in females stimulated by male odours distributed in tail displays. This is a distinct advantage as mating is restricted to a brief period with a maximum of 3 ovulations per season. 9. Acknowledgements I would like to thank the Association for the Study of Animal Behaviour for funding the project via a Teacher Researcher Fellowship. Michael Dockery (ASAB Education Officer) has provided constant support throughout the development of these materials and I am indebted to him for his many constructive suggestions. John Baumber (Headteacher), colleagues and students at Rivington and Blackrod High School have all assisted in one way or another to enable me to work on the project. My thanks are also due to Dr. Mike Hansell (University of Glasgow and Secretary of the ASAB Education Committee) for his encouragement and provision of the lemur skull photographs and Nan Swanney (Education Officer at Blackpool Zoo) for support and access to the lemur populations. Tom Mather (Lytham St. Anne s High School) trialled the project with his Psychology Students, Judy Evans produced the illustrations, Mick Hoult (MMU) provided the cover photograph and Miranda Cahill and Lauren Kavanagh assisted in the initial data collection.

17 10. Bibliography Chalmers, N. (1979) Social Behaviour in Primates: London: Edward Arnold. Dockery, M. and Reiss, M. (1996) Animl Behaviour Practical work and data response exercises for sixth form students: London: ASAB * Evans, C. S. and Goy, R. W. (1968) Social behaviour and reproductive cycles in captive ringtailed lemurs. J. Zool., Lond. 156: * Jourard, S. M. (1966) An exploratory study of body accessibility. Brit. J. Soial and Clinical Psychol. 5: * Manning, J. T. and Chamberlain, A. T. (1990) The left-side cradling preference in great apes. Anim. Behav. 39: * O Brien, T. G. (1993) Allogrooming behaviour among adult female wedge-capped capuchin monkeys. Anim. Behav. 46: Pereira, M. E. and Kappeler, P. M. (1997) Divergent systems of agonistic behaviour in lemurid primates. Behaviour 134: Richard, A. F. (1986) Malagasy prosimians: female dominance. In Primate Societies: Smuts (ed.) Univ of Chicago Pr. Sussman, R. W. (1974) Ecological distinctions in sympatric species of lemur. In Prosimian Behaviour: Martin, R. D. (ed.) London: Duckworth. * Wilshaw, J. (1998) Time budgets: advice for A level students carrying out this type of study. ASAB Feedback 13: * indicates those papers which are available on loan from the ASAB Education Officer. 11. Useful addresses Dr. Michael Dockery, Education officer, ASAB Dept. of Biological Sciences, Loxford Tower, Manchester Metropolitan University, Higher Chatham Street, Manchester M15 6HA Tel: m.dockery@mmu.ac.uk The Federation of Zoological Gardens of Great Britain and Ireland Zoological Gardens, Regent s Park, London NW1 4RY Tel: ; Fax: fedzoo@zsi.org

18 Using the Altavista search engine on the world wide web, over 9,000 links are provided for ringtailed lemurs! Useful starting points include: and if you want an amusing half hour: Student Worksheets Instructions for students provides details of the two recording methods. Recording Sheer 1A can be used by students to record continually the time spent by a particular animal in each of the four activities. For each individual animal the percentage time spent on each activity can be calculated. Recording Sheet 1B can be used to pool the data collected by a group of observers. Recording Sheets 2F, 2G,2M and 2R should be provided to a group of students observing a particular animal to record exactly what the individual is doing at each thirty second interval during the observation period. The sheets for each activity could be photocopied back to back for ease of recording up to thirty minutes of observation. Summary boxes at the end of each activity sheet will enable students to generate frequencies for each of the four activities. Lemur body outline (after Jourard (1966)) provides an alternative method for recording the parts of the body being groomed. Tally marks could be used to record the frequency of grooming for different parts of the body.

19 Instructions for students 1. Continuous recording Working in groups of two (an observer/timer and a recorder), you require a clip board, a Continuous Recording Sheet 1A and a stop-clock. What to do: Select an individual to observe. Identify your individual as an adult or a juvenile (and male or female, if possible). Observe which of the four activities the individual is engaged in. Start the clock. Observe the animal closely and note the time when the individual changes to a different activity. Record the time spent in this activity in the first row on the sheet. Continue to watch the same individual for thirty minutes (or until it goes out of sight), recording the time in a new row for each change of activity. Note any other observations that might have affected the behaviour of your individual, such as sudden noise, presence of a keeper or interactions with other lemurs. Also record the date and time of day when the observation took place. At the bottom of the recording sheet, convert all times to seconds and calculate the % time spent on each activity. 2. Instantaneous recording Working in groups of four (a timer/recorder for feeding; a recorder for moving; a recorder for resting and a recorder for grooming), you require 4 clipboards, 4 Instantaneous Recording Sheets (2F, 2M, 2G and 2R) and a stop-clock. What to do: Familiarise yourself with the descriptions used on the recording sheets. Select an individual to observe. Identify your individual as an adult or juvenile (and male or female, if possible). Note precisely where in the enclosure your individual is situated. Observe which of the four activities the individual is doing. Start the clock. At time zero, place a tick in the appropriate boxes on one of the four recording sheets to describe as accurately as you can the location and the activity of the observed animal. Repeat at 30 second intervals for up to thirty minutes (or until your individual goes out of sight). Note any other observations that might have affected the behaviour of your individual, such as sudden noise, presence of a keeper or interactions with other lemurs. Also record the date and time of day when the observation took place. At the bottom of each sheet, record the total number of observations (y) you made (the maximum is 60 if you observed for 30 minutes). On each recording sheet, count the number of instants of each type of activity (f, m, g, r) and use to calculate the frequency of each activity.

20 CONTINUOUS SAMPLING RECORDING SHEET 1A Time Budget for individual behavioural activities Species: Individual: Date: Time of Day: Time engaged in each activity (min.sec) Feeding Grooming Resting Moving Other Notes Time Spent on each Activity (sec) Total Observation Time (sec) = % Time Spent on each Activity

21 SUMMARY OF CONTINUOUS SAMPLING RECORDING SHEET 1B Species: Summary of Time Budgets for a number of individuals %Time Spent Individual Time of Day Feeding Grooming Resting Moving Other Notes Mean % time engaged in each activity SUMMARY

22 INSTANTANEOUS SAMPLING RECORDING SHEET 2F Detailed Observations of Feeding Behaviour Species: Individual: Date: Time: Location Feeding / Diet Body part used Group Water Plant Non-plant Mouth Hand Size Time (min) Inside Outside Ground Trees Furniture Leaf Non-leaf Right Left Notes 1 FEEDING

23 INSTANTANEOUS SAMPLING Detailed Observations of Feeding Behaviour Species: Individual: Date: Time: Location Feeding / Diet Body part used Group Water Plant Non-plant Mouth Hand Size Time (min) Inside Outside Ground Trees Furniture Leaf Non-leaf Right Left Total RECORDING SHEET 2F Notes Total no. of instances of Total no. of occurrences Frequency of Feeding observation (y) = of Feeding (f) = Behaviour (f/y) = 2 FEEDING

24 INSTANTANEOUS SAMPLING RECORDING SHEET 2M Detailed Observations of Moving Behaviour Species: Individual: Date: Time: Location Moving / Mode of Travel Tail Time (min) Inside Outside Ground Trees Furniture Walk Scamper Climb Leap Up Down Group Size Notes 1 MOVING

25 INSTANTANEOUS SAMPLING RECORDING SHEET 2M Detailed Observations of Moving Behaviour Species: Individual: Date: Time: Time (min) Inside Outside Ground Trees Furniture Total Location Moving / Mode of Travel Tail Walk Scamper Climb Leap Up Down Group Size Notes Total no. of instances of Total no. of occurrences Frequency of Moving observation (y) = of Moving (m) = Behaviour (m/y) = 2 MOVING

26 INSTANTANEOUS SAMPLING Detailed Observations of Resting Behaviour Species: Individual: Date: Time: RECORDING SHEET 2R Location Time (min) Inside Outside Ground Trees Furniture Resting / Posture Group All Fours Haunches; forelimbs Huddle Eyes Vigilant Size Apart Together Open Closed Notes 1 RESTING

27 INSTANTANEOUS SAMPLING Detailed Observations of Resting Behaviour Species: Individual: Date: Time: RECORDING SHEET 2R Time (min) Total Location Inside Outside Ground Trees Furniture Resting / Posture Group All Fours Haunches; forelimbs Huddle Eyes Vigilant Size Apart Together Open Closed Notes Total no. of instances of Total no. of occurrences Frequency of Resting observation (y) = of Resting (r) = Behaviour (r/y) = 2 RESTING

28 INSTANTANEOUS SAMPLING RECORDING SHEET 2G Detailed Observations of Grooming Behaviour Species: Individual: Date: Time: Location Time (min) Inside Outside Ground Trees Furniture Grooming / Groomee Part of Body Group Self Other Recip- Head Torso Fore Limb Hind Limb Tail Size Adult Juvenile rocated Front Back R L R L Notes 1 GROOMING

29 INSTANTANEOUS SAMPLING RECORDING SHEET 2G Detailed Observations of Grooming Behaviour Species: Individual: Date: Time: Location Grooming / Groomee Part of Body Group Self Other Recip- Head Torso Fore Limb Hind Limb Tail Size Time (min) Inside Outside Ground Trees Furniture Adult Juvenile rocated Front Back R L R L Total Notes Total no. of instances of Total no. of occurrences Frequency of Grooming observation (y) = of Grooming (g) = Behaviour (g/y) = 2 GROOMING

30