Activity, Behaviour and Interactions ofParrot Species at a Peruvian Clay Lick

Research Project Unit 61BL4161 (60 credit route) Activity, Behaviour and Interactions of Parrot Species at a Peruvian Clay Lick Elisabeth Mary Shaw Student Number: 05977369 Submitted by Elisabeth M Shaw, in part fulfilment of the Master of Science degree in Animal Behaviour, awarded by Manchester Metropolitan University, through the Division of Biology in the School of Biology, Chemistry and Health Sciences Submitted February 2008 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw With the exception of any statements to the contrary, all the data presented in this report are the results of my own efforts and have not previously been submitted in candidature for any other degree or diploma In addition, no parts of this report have been copied from other sources I understand that any evidence of plagiarism and/or the use of unacknowledged third party data will be dealt with as a very serious matter Signed ………………………………………………………… ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw ABSTRACT There has been much investigation in recent years into the reasons for and factors affecting geophagy behaviour by parrots and macaws at clay licks There has been much less focus on the behaviour of individual birds at these sites, despite the unique opportunity they present for behavioural investigation, and the potential such research may have to contribute to our understanding of lick use and of parrot behaviour in general This study presents some of the first detailed empirical data on the behaviours of eight parrot species at a Peruvian lick Focal animal samples performed in the early dry season of 2007 allowed the construction of an ethogram and of time budgets for birds both on the lick and in the surrounding trees and vegetation Birds in the trees around the lick spent most of their time vigilant, with the rest engaged mainly in a variety of preening and social behaviours Vigilance did not depend on flock size or body size Behaviour patterns changed little with time of day, but did change significantly during periods of lick activity There were also differences between species, particularly in Chestnut-fronted Macaws, which may have had fledglings present Birds often ate clay away from the lick face, a previously unreported behaviour that may relate to predation, competition, or the mechanics of clay processing Individual birds spent a mean of 2.35 minutes on the lick in any one feeding bout, taking a mean of 13 bites from the clay, and in a third of cases taking clay away from the lick at the end of the bout Some preference was seen for feeding from a vine rather than perched on the clay Most time on the lick was spent chewing clay, and associated behaviours such as gagging and head shaking suggested some difficulties with this Agonistic interaction rates were five times higher on the lick than in the surrounding trees, with a dominance hierarchy based on body size Most aggression occurred between conspecifics, as did most displacement in the trees, whereas most displacement on the lick itself was between heterospecifics and probably due to competition for space ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw INTRODUCTION Geophagy, the deliberate consumption of soil, has been reported from a range of mammal and bird species, as well as reptiles, lepidopterans, and even humans (Abrahams & Parsons 1996, Klaus et al 1998, Klaus & Schmid 1998, Krishnamani & Mahaney 2000, Mahaney et al 1999, Marlow & Tollestrup 1982, Smedley & Eisner 1996, Wink et al 1993) However, it is currently of particular interest in psittaciformes (parrots and macaws), many of which congregate daily at exposed riverbank sites or ‘clay licks’ to feed on the clay soil (Brightsmith 2002a, 2003, 2004a,b, Brightsmith & Aramburú 2004, Burger & Gochfeld 2003, Diamond et al 1999, Duffie 2003, Emmons & Stark 1979, Gilardi et al 1999, Mee et al 2005, Symes et al 2006, Symes & Marsden 2003) The best known are in south-east Peru, where there may be up to 100 or so licks of various sizes (C Munn unpubl data), visited by at least 17 species of psittacids, as well as various pigeons and caracids, and mammals including monkeys and squirrels (Brightsmith 2004a,b, Brightsmith & Aramburú 2004, Brightsmith & Figari 2003, Brightsmith & Houtan 2000, Hammer 2001, Hammer & Tatum-Hume 2003, Munn 1988, Tatum-Hume et al 2003, 2005) The licks are important tourist attractions, and bring in much foreign revenue every year (Brightsmith 2002a, Munn 1992, 1998) There are various theories to explain geophagy, though it may perform different functions in different species, or even multiple functions in some species (Abrahams & Parsons 1996, Diamond et al 1999, Gilardi et al 1999, Wilson 2003) Some reasons cited for soil consumption - obtaining grit for the mechanical breakdown of food, buffering of gastric pH, and treatment for diarrhoea - are largely discounted for parrots (Brightsmith & Aramburú 2004, Diamond et al 1999, Gilardi et al 1999) Instead, parrots may consume soil to obtain sodium and / or for the adsorption of toxins from the seeds and unripe fruits common in their diets (Brightsmith 2002a, 2003, 2004a,b, Brightsmith & Aramburú 2004, Diamond 1999, Diamond et al 1999, Emmons & Stark 1979, Gilardi et al 1999, Munn 1988, 1998, Symes et al 2006) Clay may also stimulate mucus production in the gut, forming a protective layer that further defends against chemical insult (Brightsmith 2002a, Gilardi et al 1999) Such protection against toxins would allow parrots to exploit a wider range of food resources, many of which are unavailable to other animals (Diamond 1999, Gilardi et al 1999) ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw Despite advances in our understanding of parrot geophagy, there are still many unanswered questions, particularly in relation to the importance of clay licks to parrot populations and distribution (Diamond 1999, Diamond et al 1999, Mee et al 2005, Symes et al 2006) Around 28% of parrot species are threatened in the wild (Snyder et al 2000) Parrots and macaws are ideal flagship species, and a better understanding of all aspects of their ecology and behaviour – including a better understanding of geophagy – may help conserve not only them but also the many other species which share their forest habitats (Munn 1998, Snyder et al 2000) Most research at parrot clay licks has focused on the reasons for and factors affecting lick use, such as weather, season, and human disturbance (Brightsmith 2002a, 2004a,b, Brightsmith & Figari 2003, Brightsmith & Houtan 2000, Hammer & Tatum-Hume 2003, Tatum-Hume et al 2003, 2005, 2006) There has so far been little focus on the behaviour of individual birds Most Neotropical parrots live in dense, closed-canopy forest, making them difficult to observe and study (Beissinger & Snyder, 1992, Gilardi & Munn 1998), and few studies have looked at their wild behaviour (Seibert 2006, Pitter & Christiansen 1997); open licks, where birds are highly visible, therefore provide unique opportunities to study them in their natural habitats A few investigations have looked at certain aspects, such as aggression and competition, responses to boats and other disturbances, and behaviours of birds waiting to descend to the lick (Burger & Gochfeld 2003, H Clegg unpubl data, Hammer & Tatum-Hume 2003, Tatum-Hume et al 2003, 2005, 2006) However, there has been a lack of detailed empirical studies Understanding behaviour at the lick may help us further understand lick use It has been suggested that one reason parrots visit is to socialise and find a mate (e.g., Brightsmith 2002a), but there are few data on social or courtship behaviours More detailed information on how birds spend their time at the lick, particularly when this can be related to lick activity, may also help us better understand anti-predator behaviour, responses to disturbance, and factors influencing feeding, as well as provide additional information on behaviours such as feeding of clay to young There has to date been no detailed study of feeding behaviours of individual birds on the lick itself, which would seem important for determining, for example, how much clay is eaten, how birds select which soil to eat, how they process it, and factors affecting these More detailed study of interactions between birds on and around the lick would also be useful in determining the extent to which competition affects lick use ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw The aims of this study were to categorise and describe behaviours shown by parrot species both on and in the trees and vegetation immediately surrounding a clay lick, to quantify patterns of behaviour and examine factors influencing it, and to assess levels of intra- and interspecific competition Some baseline lick monitoring (recording parrot numbers, species and timing of lick use) was also performed, to provide a context within which to interpret behavioural data METHODS Study area The study was performed at a clay lick on the south bank of the Tambopata River, southeastern Peru (S 12°49’23”, W 69°17’25”), at the edge of the Tambopata National Reserve The site is 30km from the town of Puerto Maldonado and 1.6km from Explorer’s Inn tourist lodge, from which it was reached either by boat or on foot along trails from the lodge The lick itself is a northwest-facing cliff approximately 8m long, 3m high, and comprising three main sections of roughly equal width (Figure 1) It is set back 30m from the river, which is approximately 200m wide at this point Behind the site is primary lowland floodplain forest, and dense understorey vegetation overhangs the top of the lick, though the lick face itself is almost totally bare The area between the lick and the river is less densely vegetated, and includes several cecropia trees (Cecropia sp.) of 15-20m high in which birds often perch before descending to feed The lick is regularly visited by up to nine species of parrots, parakeets and macaws, and squirrels and monkeys also occasionally feed and are regularly seen in the area Most species visit in the early morning and spend some time in the surrounding trees and vegetation before descending to feed, although feeding does not always occur for every species every day Tourist groups from Explorer’s Inn and Inotawa lodge visit most days during the high season (April October), the birds being viewed from two small hides 20m downstream, on the same bank as the lick itself Groups arrive in the early morning, before the birds, and usually leave an hour or two later when feeding activity is over These groups are clearly visible to the birds when arriving and leaving by boat Previous study here has found no evidence that tourist presence ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw affects bird numbers or feeding, though it may make the birds more wary (H Clegg unpubl data) Both local and tourist boats regularly pass on the river Figure 1: Photograph of the clay lick, as viewed from the Explorer’s Inn hide, showing birds using the first lick section (nearest the hide) Second section is set back from the others (centre); third section is on the right, and includes a hollow out of sight on the far right Vegetation immediately in front of the hide had been cut back half-way through the study by a member of the lodge staff (Photo: EM Shaw 2007.) Monitoring of daily lick activity Behavioural data were collected alongside monitoring of lick activity as part of ongoing work by the Tambopata Macaw Project Monitoring data were collected by between one and three observers over 30 days between 21 April and 31 May 2007 (early dry season), with most data collection taking place in the early morning between 05:45 and 08:35 Observers usually stayed until all birds had left the area and showed no signs of returning One full day (to 15:50), one late morning and one afternoon monitoring session were also performed to check whether any birds fed later in the day Heavy rain and flooding prevented access to the site on several days during the study period ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw All monitoring was performed from the Explorer’s Inn hide, with observations made with the naked eye or using 10x25 binoculars Observers recorded the start and end time of monitoring, species present, the time each arrived and the time each first landed on the lick, and the maximum number on and in the area around the lick, as visible from the hide; this was defined as the area between the lick and the river, as well as the trees and vegetation immediately above and to each side of the lick, and the trees immediately above the hide Birds out of sight behind the lick were excluded, as were those flying over and any on the opposite side of the river The number of individuals of each species on the lick itself was recorded at five-minute intervals from the time the first bird landed on the clay, and the maximum number on the lick throughout the observation period noted Video recordings were made of lick activity on most days, using a JVC GR-D370EK MiniDV Digital Video Camera at up to 32x optical zoom; on days when the amount of activity and number of behavioural samples being performed made lick counts difficult, these were taken from the recordings Recordings were made from the same position in the hide each time and usually continued for as long as birds remained on the lick Behavioural sampling Behaviour samples were recorded alongside daily monitoring and performed from the Explorer’s Inn hide; all were performed by a single observer (the author), thus avoiding interobserver bias After a few days of preliminary observations, behaviours of all species were divided into categories and an ethogram produced Any additional behaviours observed during the study, or on subsequent playback of video recordings, were added to this ethogram as and when they were identified Behaviours were sampled using focal animal sampling and continuous recording; two sets of samples were performed, the first for birds in the trees and vegetation in the area around the lick (‘behaviours around the lick’), and the second for birds on the lick itself (‘behaviours on the lick face’) All behaviours were treated as states (having duration) and given a minimum duration of one second, except Displacement, which was considered an event, and was recorded as a frequency ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw Behaviours around the lick Focal samples for birds around the lick were performed using 10x25 binoculars, and behaviours recorded into an Olympus DS-50 Digital Voice Recorder The same recorder was used to play back samples, with behaviours transcribed by hand Where possible, focal birds were chosen at random, though samples were limited to species present Samples were performed on an ad hoc basis throughout the monitoring period, with up to 35 samples in any one morning For each sample, the observer recorded the date, time of day, species, and whether any birds used the lick during the sample Samples often had to be fitted between five-minute lick counts; sample length was therefore limited to one minute If the focal bird was lost from sight or left the area during a sample, the sample was ended, unless the bird could be re-sighted within ten seconds of being lost In this case, the time spent out of sight was excluded from the overall sample time Incomplete samples were included in the analysis to avoid biasing against behaviours shown just before birds left the area; however, the sampling time was ended from the moment the bird took off, and time spent flying as it left was not included If a bird flew to the lick during a sample, it was followed on the lick, and the sample resumed if it returned to the trees before the end of the sixty seconds; flying to and from the lick was included, but behaviour while on the lick itself was not To quantify competition, the number of agonistic interactions (Displacement and Aggression) in each sample was noted, along with the species with which the focal bird was interacting, and which bird initiated the interaction In cases where the focal bird interacted with more than one other (for example, displacing two others), the number of interactions was recorded as the number of birds with which it interacted Behaviours on the lick face It proved impossible to sample birds on the lick at the same time as other behaviour samples and general monitoring; these samples were therefore taken from video footage recorded during monitoring sessions Video recordings were replayed on Windows Media® Player 11, with the timing of each behaviour taken directly from the time displayed on the screen and behaviours ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw transcribed by hand Focal birds were chosen at random, though again sampling was limited to species present and to individuals clearly visible on the video Sampling more than one bird during the same time period was avoided, though in a few cases some overlap of samples did occur Samples were started from the moment the focal bird landed on the lick face (or on a vine or branch crossing the lick face), and behaviours recorded until the moment it left the lick (flew away and out of view of the camera) If the bird was lost from sight (for example, behind vegetation) and could not be re-sighted before it left the lick, the sample was ended from the moment it was lost If the bird could be re-sighted the sample was resumed, and any time out of sight was recorded as such The reason the bird left the lick at the end of the sample was noted, and divided into the following categories: unknown (no clear reason for leaving), lost (sample ended before bird had left, either because it was lost, or, on at least one occasion, because the video cut off), flush (some or all of the birds, including the focal individual, suddenly fly from the lick), displacement (focal bird is displaced by another), or leaving with clay (no other clear reason for leaving, but focal bird can be seen holding clay in its beak as it flies away, or is assumed with some certainty to be holding clay, for example because it flies immediately after biting the lick) The observer also recorded the species, time and date, and which lick section was used In addition, the position of the bird (perched on the clay itself, on a vine or branch, or with one foot on each) was recorded throughout the sample Agonistic interactions were recorded as before, and included Displacement occurring as the bird landed at the start of the sample and Displacement of the focal bird when this caused the bird to leave at the end of the sample Data analysis Non-parametric statistical tests were used as most of the data were not normally distributed Statistical analyses were performed using Minitab® 14 Monitoring data were used to calculate mean flock sizes, mean maximum number of birds on the lick on days species fed, and the mean time each species spent on the lick, as well as lag 10 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw The mean bout duration of 2.35 minutes was, however, almost exactly the same as that reported elsewhere for large macaws (2.37 mins; C Munn, unpubl data) The number of birds on the lick did not significantly affect the time an individual spent on it, but less time was spent when there were more birds in the area; perhaps larger flocks encouraged individuals to return to the trees with clay, since they would then be less likely to be alone in the trees Most departures from the lick were not due to displacements or flushes, but to birds leaving with clay or for no apparent reason, perhaps simply having eaten their fill Studies generally only report flushes as terminating feeding bouts; these data suggest these account for only around a fifth of departures There was some preference for feeding from a vine; this was a single piece of vegetation covering only a tiny fraction of the lick, yet birds spent a quarter of their time perched on it, perhaps because they could more easily turn outwards from the lick to watch for predators from this position The clay itself appeared difficult to process, with much time spent chewing and with birds often performing ‘gagging’ motions or head-shaking as if trying to dislodge or discard clay from the mouth, though recent rainfall (which might be expected to affect clay texture) had no effect on these That clay takes much time and effort to eat supports the idea that its consumption must be important Agonistic interactions Only small amounts of time were spent on Aggression, and agonistic interactions were only recorded in 7% of birds around the lick, at a rate of around one interaction per bird every ten minutes Though interaction rates have been reported for some psittacids in captivity (e.g., Seibert & Crowell-Davis 2001), to my knowledge this is the first quantitative data for a lick Interaction rates on the lick itself were almost five times higher than in the trees, which would be expected as birds crowd together and compete for access to clay, and agrees with other studies that have found higher agonistic behaviour when parakeets congregate to feed or roost (Hardy 1965, Power 1966) 25 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw Despite this, birds rarely interacted in situations where they might be expected to, often feeding side-by-side and even piling on top of each other without interacting Lick use may be relatively low at this time of year (Brightsmith 2002a, 2004a, 2006, Brightsmith & Figari 2003, TatumHume et al 2006), meaning fewer birds and less competition for space Parrots are also reported to generally be non-territorial in relation to food resources (Renton 2004), and many report low levels of aggression and competition at feeding sites, both amongst parrots and tropical frugivores in general, possibly as they feed on superabundant resources (Fleming 1979, Gilardi & Munn 1998, Pitter & Christiansen 1995, Seibert & Crowell-Davis 2001), though it remains to be shown whether clay licks constitute such a resource Most Aggression was between conspecifics, though this varied between species This agrees with findings at another Peruvian lick (Burger & Gochfeld 2003) and in Blue-and-yellow Macaws, Ara ararauna, at nesting sites (Renton 2004), though others have found higher interspecific agonistic interactions in congregations of species such as passerine migrants (Salewski et al 2007) Most Displacement in the trees was also between conspecifics, but on the lick was mostly between heterospecifics, suggesting that in the trees it probably involves intraspecific displays of dominance, whereas on the lick it is more to with competition for space As found by others (Burger & Gochfeld 2003, Diamond et al 1999, Duffie 2003), there was a dominance hierarchy based on size, with larger species dominating smaller ones (Table 6) This has also been found in passerine migrants (Salewski et al 2007), mixed-species tit and gull flocks (Alatalo & Moreno 1987, Bellebaum 2005), foraging groups of Australian birds (Shelley et al 2004) and African frugivores (French & Smith 2005), though others suggest competition is highest between similarly sized species (Leyequién et al 2007) Body mass was also important; for example, Chestnut-fronted Macaws, though longer than Mealy Parrots and with a higher Dominance Index, are much lighter, and it was Mealy Parrots that initiated more agonistic interactions, including against the macaws Hierarchies may also be non-linear (e.g., Seibert & Crowell-Davis 2001, Shelley et al 2004); for example, Mealy and Yellow-crowned Parrots initiated the same number of interactions against each other Smaller species also had a tendency towards higher interaction rates, more intraspecific interactions and more Displacements (Table 5), though the reasons for this are unclear, and not appear to relate for example to flock size Pitter and Christiansen (1997) suggest that 26 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw aggression within groups is more common among smaller parrot species, though they offer no explanation for this Burger and Gochfeld (2003) report that Blue-headed Parrots, though relatively small, dominated others at a lick by being most numerous This has also been found for example between gull species (Burger and Gochfeld 1984) However, there was no relation between flock size and dominance here Studies in other bird species have also found no evidence for numerical dominance by smaller species (Bellebaum 2005, Shelley et al 2004) Subordinate species may be relegated to poorer foraging sites or be forced to feed at different times or in areas of higher predation risk (Alatalo & Moreno 1987, Bellebaum 2005, Shelley et al 2004) Burger and Gochfeld (2003) claim more aggressive species usually had primary access to the best places on a lick, and that smaller parakeets either abandoned feeding or moved to different parts of the lick when larger birds arrived Competition may partly explain why Dusky-headed Parakeets returned to the lick later in the day, though further study would be needed to determine whether competition reduces feeding at this lick, and the ways less dominant species respond to it General conclusions This study gives one of the first detailed, quantitative analyses of behaviours of individual birds at a clay lick, and in particular of feeding behaviours on the lick itself There were certain limitations, such as a relatively short study period and possible sampling biases, with some individuals and species more easily sampled than others and possibly more sampling at ‘quieter’ times when birds were easier to observe The behaviour categories could be further refined, but served the purposes of this study However, it was sometimes difficult to determine exactly when one behaviour ended and another began, and just because a behaviour takes a short time does not mean it is less important It also proved difficult to record and interpret behaviours 27 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw without some assumptions about their functions, which may be misleading if the assumed functions are incorrect Despite these limitations, the results suggest behavioural studies can be useful in revealing more about parrot geophagy As well as giving a clearer picture of how birds spend their time when not feeding - which is a first step in discovering why they often spend time at the lick without descending to feed - detailed study of birds on the lick has given estimates of individual clay consumption that may prove more accurate than those currently in common use The study has also started to reveal more about anti-predator behaviour, in particular with the first records of clay being taken to the trees for consumption, and the suggestion that clay takes considerable time and effort to eat supports the idea that its consumption must be important to these species The finding that larger species dominate raises questions over what impacts competition may have on lick use by subordinate species, and so also what knock-on effects changes in abundance of larger species (for example, due to habitat loss) may have There are many possible extensions For example, behaviours could be compared between licks and under different conditions (such as flock sizes, weather conditions, times of year, and human disturbance) to tell us more about the factors influencing behaviour and lick activity Recording how birds distribute themselves would give more information on social structure, and studying behaviours away from the lick site would provide a useful comparison Properly describing and quantifying behaviour is important in being able to objectively perform such comparisons Parrots are notoriously difficult to study in the wild, and there have been few behavioural studies in their natural habitats (Beissinger & Snyder 1992, Gilardi & Munn 1998, Seibert 2006, Pitter & Christiansen 1997) This study presents one of the first attempts at producing an ethogram for a range of wild psittacines, and demonstrates the huge potential that clay licks provide not only for better understanding geophagy but also, by virtue of good visibility and relatively large numbers of a range of species, for performing a variety of investigations into many other aspects of parrot behaviour 28 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw ACKNOWLEDGEMENTS I would like to thank all the staff of Explorer’s Inn / Peruvian Safaris, and in particular Dr Max Gunther, for allowing me to stay at Explorer’s Inn as a Resident Naturalist while performing this research, and for all their help and support I am also particularly grateful to Andy and Karen Wilkinson, Sara Prado, Martha Vazquez, Patricia Escalante, Eugenio Sola, and others for help with data collection, and to the boat drivers at Explorer’s Inn for providing transport to and from the study site Finally, I would also like to thank Chris Kirkby, Alan Lee, Karen Tailby, Les May, Martin Jones and Stuart Marsden for valuable advice and support This work was performed with the permission of INRENA, Peru, and I am grateful to the Tambopata Macaw Project, and in particular Alan Lee, for their help in arranging permits 29 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw REFERENCES Abrahams, P W & Parsons, J A 1996 Geophagy in the tropics: a literature review The Geographical Journal 162 (1): 63-72 Alatalo, R V & Moreno, J 1987 Body size, interspecific interactions, and use of foraging sites in tits (Paridae) Ecology 68 (6):1773-1777 Beissinger, S R & Snyder, N F R 1992 New World Parrots In Crisis Smithsonian Institution Press, Washington, D.C Bellebaum, J 2005 Between the Herring Gull Larus argentatus and the bulldozer: Black-headed Gull Larus ridibundus feeding sites on a refuse dump Ornis Fennica 82: 166-171 Brightsmith, D 2002a The clay licks of 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The Condor 106: 395-400 Smedley, S R & Eisner, T 1996 Sodium: A male moth’s gift to its offspring Proc Natl Acad Sci USA 93: 809813 Snyder, N., McGowan, P., Gilardi, J & Grajal, A (eds.) 2000 Parrots Status Survey and Conservation Action Plan 2000 – 2004 IUCN, Gland, Switzerland and Cambridge, UK x+180pp Symes, C T., Hughes, J C., Mack, A L & Marsden, S J 2006 Geophagy in birds of Crater Mountain Wildlife Management Area, Papua New Guinea Journal of Zoology 268: 87-96 Symes, C T & Marsden, S 2003 Geophagy and parrots in Papua New Guinea PsittaScene 15 (3): 12-13 Tatum-Hume, E., Lee, A., Fothergill, C & Hammer, M 2005 Surveying monkeys, macaws and other wildlife of the Peru Amazon Biosphere Expeditions: Expedition Report 2005 Available from (Accessed January 2007) Tatum-Hume, E., Lee, A., Moore, D & Hammer, M 2006 Surveying mammals, macaws and other wildlife of the Peru Amazon Biosphere Expeditions: Expedition Report 2006 Available from (Accessed January 2007) Tatum-Hume, E., Müller, M M., Schmidt, K S & Hammer, M L A 2003 Surveying monkeys, macaws and other wildlife of the Peru Amazon Biosphere Expeditions: Expedition Report 2003 Available from (Accessed January 2007) Valone, T J & Lima, S L 1987 Carrying food items to cover for consumption: the behavior of ten bird species feeding under the risk of predation Oecologia 71 (2): 286-294 Ward, P & Zahavi, A 1973 The importance of certain assemblages of birds as “information-centres” for foodfinding Ibis 115: 517-534 Westcott, D A & Cockburn, A 1988 Flock size and vigilance in parrots Australian Journal of Zoology 36 (3): 335-349 Wilson, M J 2003 Clay mineralogical and related characteristics of geophagic materials Journal of Chemical Ecology 29 (7): 1525-1547 Wink, M., Hofer, A., Bilfinger, M., Englert, E., Martin, M., & Schneider, D 1993 Geese and dietary allelochemicals: food palatability and geophagy Chemoecology 4: 93-107 33 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw APPENDIX - Ethogram of behaviours shown by parrots on and around the clay lick The following list of behaviour categories is not exhaustive, and does not include behaviours not observed at the lick (for example, sleeping, feeding (other than on clay), copulation); others such as defaecation were also not included Categories are not necessarily mutually exclusive (for example, a bird can Preen while being Allopreened), & some categories may encompass more than one distinct behaviour, grouped together for convenience Behaviour codes, as used in tables, are given in parentheses Where a behaviour is specific to the lick face or to the trees / vegetation around the lick, this has been noted Aggression (AGGR) – aggressive behaviour directed towards and / or received from one or more other birds (conspecifics or heterospecifics); usually involves extending head towards other bird with beak open as if attempting to bite, and may be accompanied by vocalisations and flapping of the wings May or may not make contact with the other bird (such as biting or pecking), and may chase it (pursue it while displaying aggressive behaviour towards it) Other bird may show submissive behaviour such as ‘crouching’, move away, or retaliate with aggressive behaviour of its own Excludes interactions where the intention was not obviously aggressive (such as stretching beak towards another without the apparent intention to bite and without vocalisations or postures normally associated with aggression), and causing the other bird to move away simply by moving towards it (Displacement) Aggressive postures and movements (e.g., ‘gape’, ‘peck’, ‘lunge’, ‘bill-fencing’) have been described elsewhere by Hardy (1965) and Power (1966) Allofeed (ALLFE) – transfers food to another by regurgitating crop contents into the beak of the other, or receives food regurgitated to it by another in this way The birds face each other with heads tilted in opposite directions and press open beaks together, then move their heads up and down as the donor bird regurgitates the food and passes it into the beak of the other Only clay was seen fed in this way during the study (confirmed by the sight of clay in the beak of the recipients), and it appeared to be fed to juveniles (the behaviour was observed between four individuals, two of which fed both the others, suggesting a family group of two parents feeding two offspring) 34 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw Allopreen (ALLPR) – preens feathers or body of another individual (of the same species), and / or another preens the focal bird (runs their feathers through its beak; see Preen) The behaviour was recorded from the moment the beak of the individual doing the preening touched the feathers of the one being preened, to be followed by preening behaviour Focal bird can be engaged in another behaviour (such as Vigilant, Call or Preen) at the same time as being allopreened by another; in this case the behaviour was still recorded as Allopreen Excludes aggressive pecking or biting of the body of another; the birds usually stand close together and no aggressive postures or vocalizations are shown Bite (BT) – ON LICK – bird faces the lick; head moved towards the lick face and beak stretched towards it with the apparent intention of touching it with the beak When beak contacts the lick face, it is opened and closed against it (‘biting’) to remove portions of clay Includes all attempts at biting the clay even if the bird is unsuccessful in removing any or is unable to contact the lick face (for example because it is too far away); excludes biting of any other substrate or food item The behaviour was recorded from the moment the beak first contacted the lick face (or when the beak was first stretched towards it if the bird was unable to reach it), until the moment the head was raised and the beak did not re-contact the clay for at least one second, unless the beak was still stretched towards and being moved across the surface of the lick in this time (as if about to bite again) and the bird was not chewing Call (CALL) – vocalises (opens beak briefly and emits a sound); if the vocalisation could not be heard by the observer (due to distance or general noise levels), calling was inferred from the beak movement, and sometimes also slight movements of the throat, the chest or sometimes the tail that accompanied it; feathers on back of neck also sometimes raised In some species, occasionally accompanied by brief raising and extension of the wings above the back Includes various types of vocalisation (for example, alarm calls, contact calls), grouped together for convenience and because determining function was beyond the scope of this investigation Excludes vocalisations accompanying Aggression, Wing Flip and Hang-Play, and opening of the beak for any other reason (such as yawning or biting), not accompanied by a sound Usually accompanied by Vigilant behaviour, but recorded as Call for as long as there was one second or less between vocalisations Displacement (DISP) – moves towards (flies towards, lands beside, or walks or climbs towards or past) another bird or birds (conspecific or heterospecific) and in doing so causes the other bird(s) to fly or move away, or focal bird itself flies or moves away as another moves towards it in this way; the displaced bird does not usually contest or defend its position May be a deliberate attempt to take the perch occupied by the displaced bird, or the displaced bird may simply move out of the way in an 35 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw attempt to avoid another; there may or may not be contact between them Also includes cases where a bird is physically knocked from the lick face by another hitting it and causing it to lose its grip Excludes a bird moving or flying away in response to Aggression (aggressive postures or behaviour) Eat Clay (CLAY) – TREES / VEGETATION AROUND LICK – clay is held in the beak or grasped in a foot; the bird takes bites from the clay held in the foot (opens and closes beak against the clay to remove portions and take them into the mouth) and / or chews clay in the beak (beak movements can be seen) ‘Gagging’ motions may also be seen (neck repeatedly extended with beak open or being opened and closed) Behaviour ends when beak movements stop and no more bites are taken, even if clay is still being held in the foot May be Vigilant while chewing, but recorded as Eat Clay for as long as beak movements continue Swallowing of the clay was assumed to occur as clay was rarely seen dropped from the beak ON LICK, this behaviour was divided into: Clay From Foot (CFF) – clay grasped in one foot; foot is brought to beak and / or beak is lowered to foot, and bird takes bites from the clay held in the foot Also includes the time taken to initially transfer clay from the beak to the foot Behaviour ends when the head is lifted away from the foot (the foot may or may not be lowered) and no more bites are taken for at least one second Vigilant-Chew (VGCH) – head is raised away from the lick face (beak not extended towards or in contact with it), though the bird may or may not still be facing the lick; constant beak movements can be seen (‘chewing’) The behaviour ends when beak movements stop; if the beak could not be seen (for example because the bird was facing away from the observer), chewing was assumed if the bird had just bitten from the lick face and could not be seen to be performing any other behaviour ‘Gagging’ motions may also be seen Termed Vigilant-Chew as bird often turns whole body to face away from the lick and moves head around as if ‘looking around’ (vigilant) Again, swallowing of the clay was assumed as clay was rarely seen dropped from the beak Includes chewing of clay after lifting the head having taken a bite from clay held in the foot Fly (FL) – airborne, wings extended (flapping or gliding) and body not in contact with a perch or the ground Excludes flapping of the wings while perched or hanging from a perch, or being briefly airborne 36 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw when jumping between perches (possibly with the aid of the wings, but without the wings holding the bird in the air) Grip (GRIP) – ON LICK – usually on landing, flaps wings, and moves feet (and sometimes also beak) against lick face in an apparent attempt to gain purchase on the clay, or to regain balance and grip after slipping (losing purchase on the lick face) Excludes time for which bird is airborne (Fly), lifting or flapping of the wings while perched on the lick face and otherwise more or less stationary with both feet gripping the clay, and extending the wings as an aid to balance while stretching towards the lick face from a vine or branch Behaviour ends when bird succeeds in gripping lick with both feet to end up in a stable, stationary position, or else lets go of the lick entirely and flies Hang (HA) – TREES / VEGETATION AROUND LICK (also occurred on lick, but not recorded for lick samples) – body held beneath perch such that body weight is suspended from one or both feet and / or from beak, and this body part or parts are above the rest of the body This position was recorded separately to other behaviours; that is, behaviour during Hang was recorded as that accompanying it (for example, Vigilant, Manipulate Vegetation, Hang-Play), and the time spent in Hang analysed separately (with the bird assumed to be upright on a perch for all other time except that spent flying) Hang-Play (PLAY) – in Hang position, and interacting with a conspecific that is usually, though not always, also hanging; the birds bite and kick at each other, with wings occasionally spread or flapped Often accompanied by vocalisations Neither bird makes an immediate attempt to flee and interaction does not appear to be aggressive in nature, with bites not apparently intended to cause actual physical harm The behaviour was termed ‘play’ for convenience, as it most closely resembles play-like behaviour and has no other apparent function; this is not meant to imply, however, that this function is certain Also observed in Chestnut-fronted Macaws away from the lick site Head Shake (HSH) – head rapidly flicked / shaken from side to side, up to several times in quick succession; may be repeated Beak may be held open during shaking, sometimes widely, and head may be held so that bird is facing downwards and head is below body, angled towards the ground Behaviour ends when no shaking has occurred for at least one second, unless the beak remains open and / or pointed downwards during shakes, in which case it ends when this position has ended and no more shaking occurs Assumed to be associated with clay eating as it only occurred during chewing of clay Though not recorded in birds in the trees, this was later noted from video footage of birds eating clay in 37 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw the trees Manipulate Vegetation (VEG) – branch, leaf, flower or other plant parts bitten at, chewed in situ on the plant, pulled at with the beak, or removed from plant and chewed in beak (beak movements can be seen); may also be held in foot and bitten at Probably includes several behaviours, grouped together here because their distinctions were unclear May have involved object play in some cases (for example, holding a dead leaf in the beak while hanging upside down), though unclear whether it was a feeding behaviour in others; leaves, for example, were sometimes held in the foot and bitten at but then dropped, and branches were often chewed without bark or other material being removed from them In other cases, bark appeared to be chewed in the beak, though it was not possible to tell whether it was swallowed May be particularly associated with juveniles; for example, fledgling keas (Nestor notabilis) spend more time than adults manipulating inedible objects (Diamond & Bond 1991) Excludes scraping beak against branch without biting motions (beak being opened and closed against it); see Scrape Beak Move (MV) – uses feet and sometimes beak to move the body along a perch, through vegetation, across the lick face, or from the lick face to adjacent vegetation (walking or climbing) Also includes jumping short distances between perches, possibly using the wings to aid balance, but not to hold the bird in the air (that is, not Fly) Behaviour continues as long as the bird remains in motion, with no stationary pauses (feet no longer moving, and the bird not still in the process of using the beak to pull the body upwards) of more than a second Excludes turning on the spot to face the opposite direction on a perch, when the bird does not move more than one step in a horizontal (or vertical) direction in the process Other Interaction (INTER) – any interaction with at least one other conspecific or heterospecific (that is, one bird approaching and / or directing a behaviour towards another, or in some way reacting to the behaviour of another) that does not fall into any other category of interaction or communication, or an interaction that is unidentified or difficult to classify Examples include ‘touching beaks’ (when this is not part of Aggression), ‘ducking’ in response to another landing above on the lick, or pressing the body against a conspecific in an apparent attempt to solicit Allopreening Preen (PR) – focal bird runs its own feathers through its beak, or runs the beak across its own legs or feet (possibly appearing to nibble at the skin) Also includes scratching (foot passed rapidly and repeatedly against head or body), stretching the wings (both wings slowly lifted and brought together behind the back, or one wing extended downwards and to the side whilst simultaneously extending the 38 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw leg on the same side), a momentary raising (‘fluffing’) of all the feathers, usually accompanied by a brief shake of the whole body, or rubbing the head against any part of the body Excludes preening any part of another individual (see Allopreen) Scrape Beak (SCRAPE) – beak pressed against and passed backwards and forwards against a branch or perch, once or a few times in quick succession Beak is not opened and closed around the perch (that is, it is not being bitten or chewed), and no material (such as bark) is being removed from it Excludes the beak being pressed against the lick face in any way Probably functions to clean the mandibles of food or other items (Power 1966); could have been included in Preen Unidentified Behaviour (UN) – any behaviour not included under any other category, or that cannot be identified, either because its identity or purpose is unclear or because the bird is partially or fully obscured from view such that its behaviour cannot be accurately determined (but it has not left the study area, or the lick face for lick samples) Excludes unidentified interactions (see Other Interaction) Vigilant (VG) – TREES / VEGETATION AROUND LICK – engaged in no other obvious behaviour; body held relatively still, though head may be moved as if bird is looking around, or the bird may turn around or adjust its position slightly (without moving more than one step in a horizontal or vertical direction in the process) Eyes are open and head is not held under the wing (that is, bird is not sleeping) Often accompanies other behaviours (such as Call, being Allopreened) – behaviour was recorded as Vigilant only when no other behaviour was being performed at the same time Note that the name Vigilant has been chosen for convenience and is not meant to imply that vigilance is definitely the function of the behaviour; indeed, it may encompass several functions, including resting ON THE LICK – as above Head may be raised and whole body turned so the bird is facing away from the lick face No beak movements (not chewing); if clay is held in the beak it is not being chewed Wing Flip (WF) – feathers on back of neck raised, head may be lowered and may be moved up and down in a slow ‘bobbing’ motion; one or both wings briefly lifted away from the body and then dropped (may be repeated, usually with a few seconds in-between) Wing motions are slight, and wings are not fully extended as if flapping May be accompanied by soft vocalisations Wing motions may be performed alone, without any of the accompanying motions, postures or vocalisations May be performed during another behaviour such as Vigilant or Allopreen, but recorded as Wing Flip for as long as the body posture continues or there is a wing motion at least once a second Appeared to be directed 39 ... (Peru) clay lick: temporal patterns, associations, and antipredator responses Acta Ethol 6: 23-34 30 ACTIVITY, BEHAVIOUR AND INTERACTIONS OF PARROT SPECIES AT A PERUVIAN CLAY LICK Elisabeth M Shaw... Chestnut-fronted Macaw (Ara severa), Dusky-headed Parakeet (Aratinga weddellii), Mealy Parrot (Amazona farinosa), Orange-cheeked Parrot (Pionopsitta barrabandi), Red -and- green Macaw (Ara chloroptera), White-eyed... comparison with behavioural data at these times For focal samples, the total percentage of time spent on each behaviour in each sample was calculated, and a mean percentage calculated over all samples