Professor Ken Cheng

Studies in Animal Behaviour


Ken Cheng
Department of Biological Sciences
Macquarie University
Sydney NSW 2109 Australia
phone: 612 9850-8613
FAX: 612 9850-8245

GoogleScholar page:

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My research crosses mechanistic, functional and evolutionary questions in the study of animal behaviour. A central theme of my research concerns how animals process information. Dealing with information is crucial for many important behaviours in an animal's life, including choosing a mate, avoiding predators, and finding food. The range of species I have studied include humans, rats, pigeons, chickadees, Clark's nutcrackers, desert ants, and honeybees. Recently, thanks to a student, my group is also examining the urban ethology of flying foxes on the east coast of New South Wales, Australia. A large part of my research has concentrated on how animals deal with space and time. I have collaborations with a number of researchers around the world.

Macquarie University funds postgraduate students from anywhere in the world with scholarships. I am currently looking for students to study the behaviour of ants in Australia. We are studying one species of desert ant, the highly thermophilic the red honey ant Melophorus bagoti, that lives in cluttered semi-arid habitats. We are also studying of bull ants found on campus here. The latest venture is to provide reconstructed reality for ants, replicas of their natural scenes so that we can better control and manipulate the cues. I welcome enquiries from those interested.

Research links

For prospective Masters and PhD students



I have published a introductory book on animal cognition suitable for high-school and first-year university students, called How Animals Think and Feel (2016), published by ABC-CLIO. The link to the publisher's site for the book is here.

The first half of the book covers various topics in animal cognition, including a chapter on emotions. The second half of the book parades selected groups of animals, ending with thoughts about our own species.

Current research



Starting this century, my team has been studying an Australian desert ant located in Central Australia. Known as the red honey ant (pictured above), Melophorus bagoti shares many characteristics with Cataglyphis. It is long-legged, active in the heat of the day, and moves fast. The work on Australian desert ants started in a collaboration with Ruediger Wehner of the Brain Research Instiitute, University of Zurich. We found the ants in Central Australia together. Wehner has established and run a field research station in Tunisia for over 30 years. The subject of his research has been various species of the desert ant, genus Cataglyphis. Much about the navigational behaviour of these ants is known as a result of the very fruitful work of Wehner and his collaborators. But of course much more remains to be researched.

But the visual ecology of M. bagoti differs from that of hte much studied Cataglyphis fortis. Unlike the salt pans on which C. fortis live, the semi-deserts of Central Australia are richer in plant life, making the visual world of M. bagoti cluttered with landmarks in the form of grass tussocks, bushes, and trees. In comparison with C. fortis, M. bagoti relies more on the terrestrial visual panorama. We have two reviews on the work (Cheng et al., 2009, Behavioural Processes; Cheng et al., 2014, Behavioural Processes).

We have also done some research on a species of Melophorus ants (yet to be named) living on the salt pans of South Australia. This environment is barren and devoid of landmarks. We found the animal in December 2011. Two papers featuring this species have been published (Schultheiss et al., 2012, Australian Journal of Zoology; Scultheiss et al., 2016, Behavioral Ecology and Sociobiology).

In the latest venture, I am collaborating with a team (Jochen Zeil, Ajay Narendra, Andy Barron, and including Ruediger Wehner) to delve into the neurobiology of navigation in ants. Australian bull ants, genus Myrmecia, features in this research.

Myrmecia midas, Australian bull ant Myrmecia midas

Cody Freas has been studying the night-active bull ant we have on campus, Myrmecia midas, and his work describes how this ant navigates. These nocturnal bull ants sport big eyes and powerful jaws; they pack a mighty sting as well. The ants come out foraging at sunset, and return to their nests throughout the night. They nest at Eucalyptus trees, and a portion of the foraging force travel up their nest tree to forage. The surrounding panorama plays a large role in their navigation, both on the ground and on trees.

Myrmecia midas travelling on a tree  Myrmecia midas scanning on a tree Myrmecia midas travelling on trees

When travelling on a tree, we have observed that these ants often try to assume a horizontal head position, from which stance they turn their head to scan the surrounding panorama. Beyond some qualitative descriptions, we have yet to quantify such behaviours in detail. But they do navigate on trees. With the help of the visual panorama, they move themslves to the side where their nest is well before reaching the ground.

Cody has submitted his thesis in January of 2018.

Some publications on M. midas:

Freas, C. A., Narendra, A., & Cheng, K. (2017).  Compass cues used by a nocturnal bull ant, Myrmecia midasJournal of Experimental Biology, 220, 1678-1585.

Freas, C. A., Narendra, A., Lemesle, C., & Cheng, K. (2017).  Polarized light use in the nocturnal bull ant, Myrmecia midasRoyal Society Open Science, 4, 170598.

Freas, C. A., Wystrach, A., Narendra, A., & Cheng, K. (2018).  The view from the trees: Nocturnal bull ants, Myrmecia midas, use the surrounding panorama while descending from trees.  Frontiers in Psychology, 9, 16.

Cataglyphis in Tunisia

Melophorus in Central Australia

Saltpan Melophorus ants in South Australia

Grey-headed flying-fox mother and pup  flying-fox mother with pup2 Grey-headed flying-fox mothers with pup

Urban ethology of flying foxes

Tim Pearson is completing a thesis on the ecological acoustics of flying foxes, and has led me into the world of urban ethology. Grey-headed flying foxes, Pteropus poliocephalus, are found in the Sydney area. They are often found in urban settings, near people's homes, as the following photos show.

Flying foxes in urban areas 1  Flying foxes in urban areas 2 Grey-headed flying foxes in urban areas near homes

The flying foxes in these two photos are in the Ku-ring-gai Flying-fox Reserve, located in a leafy suburb of Sydney called Gordon. The Reserve, managed by Ku-ring-gai Council, includes a variety of wildlife habitats and is home to a variety of threatened species, of which the grey-headed flying fox is one. The population of flying-foxes in the Reserve varies considerably in numbers over the year. Records exist for monthly population counts over the last 22 years. On a year-to-year basis as well, the number of bats in the Reserve varies considerably. Generally, however, highest numbers are found in late maternity season (January - March) and lowest numbers over the southern winter (June - August) when in some years the camp is deserted.

Male grey-headed flying foxes sometimes engage in various male-male contests:

Male grey-headed flying foxes in contest Two male flying foxes contesting at the Gordon camp.

Male flying foxes, of course, also court females. This male (top) has been rebuffed by the female (bottom):

male and female grey-headed flying foxes Male grey-headed flying fox (top) approaching a female grey-headed flying fox (bottom)

Besides the grey-headed flying fox, the region also boasts a rarer black-headed flying fox, Pteropus alecto. These ones are found in northern New South Wales, at a town called Byron Bay:

black-headed flying foxes at Byron Bay Black-headed flying foxes, Pteropus alecto, at Byron Bay, New South Wales

Tim Pearson has networked with the Ku-ring-gai Bat Conservation Society, Ku-ring-gai Council, and other helpful organisations that facilitate research. Both the Ku-ring-gai Bat Conservation Society and the Ku-ring-gai Council encourage and welcome non-invasive research at the Ku-ring-gai Flying-fox Reserve. Tim has also set up recording equipment to capture the behaviour of bats at the Reserve. Thanks to Tim Pearson, we have connections as well as excellent research equipment. The research on the urban ethology of flying foxes, however, has only just begun.We also have numerous lines of research that could be pursued. Interested students are welcome.

Some old stories

  memory in honeybees

My experiments on honeybees have investigated how the foragers find a rewarding place. Past work focussed on the theme of the mechanisms by which honeybees find a place. Current research focuses on the topic of how the worker retrieves the correct memory of a place and whether and how multiple memories are integrated. Catherine Prabhu recently completed a thesis on how honeybees deal with conflicting evidence.

research on free flying honeybees

gallery of bee pictures


geometry and navigation

This is a topic that I started to study in my graduate school years. It has quite taken off. I haven't done any more empirical work on it, but have contributed theoretically. The unkind might say that I have milked other people's work. The kinder might say that theoretically contributions can be important.

Geometry is the layout of surfaces in the environment. What any animal learns about geometric cues is a matter of some debate. The link has more information.

geometry and navigation


Clark's nutcrackers

The Clark's nutcracker has a prolific spatial memory. The bird lives in the Rocky Mountains of North America, and stores a lot of food, mostly pine seeds. It relies on its stored caches for sustenance over winter. It can remember thousands of caches for months. Alan Kamil and Russell Balda have done many studies on this bird. We now know that they use landmarks to remember and retrieve their caches. I collaborated with Debbie Kelly, now at the University of Manitoba, Russ Balda, and Alan Kamil in figuring out the cues used by these birds to remember cache locations in the lab.

Some publications:

Kelly, D. M., Cheng, K., Balda, R., & Kamil, A. C. (in press). The effects of sun compass error on spatial search by Clark’s nutcrackers. Integrative Zoology. [The data in this study are from quite a few years ago.]

Kelly, D. M., Kamil, A. C., & Cheng, K. (2010).  Landmark use by Clark’s nutcrackers (Nucifraga columbiana): influence of disorientation and cue rotation on distance and direction estimates.  Animal Cognition, 13, 175-188.


crab spiders and bees

I collaborated with Astrid Heiling and Marie Herberstein of the Department of Biological Sciences, Macquarie University, on the study of signal interactions between crab spiders, flowers, and bees, which are potential prey for spiders. Crab spiders may lure bees deceptively with their body coloration, to the detriment of the bees.

crab spiders and bees

Spatial generalisation in bees

In spatial generalisation, a worker is trained to find food in a container at one location. After sufficient training, she is presented a container at various locations, including the training location. The question at stake is: how should the animal 'bet' on whether the container at each location has food or not. See the link for some answers.


Self control in bees

In experiments on self control, the forager is presented with two choices of rewards. One is immediately available but is small. The other reward is larger, but the forager has to wait some time for it. Waiting for a larger reward is technically called self control, lack of which is often a nemesis in human behaviour. Bees show a good deal of self control.

learning and face perception in humans

I have an ongoing collaboration with Marcia Spetch of the Department of Psychology, University of Alberta. Students and colleagues have been collaborators, including Colin Clifford of the School of Psychology, University of Sydney. We investigated a number of topics in spatial and temporal cognition in pigeons and humans. A story on spatial cognition is linked below.

We found both peak shift and range effects in human face identification.


Spatial cognition in humans

Marcia Spetch and I have published a substantial story on spatial cognition in university students, investigating spatial generalisation and peak shift. Students were presented marked locations on a computer screen, and had to bet whether it was the rewarding 'hot' spot. Our latest work, published in 2010 online, tries to provide functional explanations for all range effects in human learning.


Perception of bilateral symmetry

Two colleagues at our University, Chris Evans and Peter Wenderoth, both now deceased, collaborated with me in studying the perception of bilateral symmetry in complex stimuli in humans. Bilateral symmetry means mirror symmetry. It is often an attractive property in mate selection. We studied in human subjects the perception of symmetry in complex, naturalistic objects.


Virtual ecology of bee-flower interactions

Chris Evans and I have been exploring this topic 'on the back burner' for a number of years. The idea is to present virtual flowers, generated on computer to real bees. Each flower contains the same reward, but the bees get to choose which flower to land on and get their sugar water from. The virtual aspect makes it possible to manipulate key parameters in evolution, such as costs in producing flowers. It also speeds up the 'evolutionary' process, making generations go by in days. We think that this makes a great project for an interested graduate student.

Publications and curriculum vitae

selected publications and abstracts

curriculum vitae and publication list (some of which can be downloaded)

PhD Students

Cody Freas: studying navigation in ants

Tim Pearson: studying auditory communication in flying foxes

Muzahidul Islam: studying learning and navigation in bull ants

Sudhakar Deeti: studying learning walks in desert ants


I teach BIOL122 Biological Basis of Behaviour. A brief description:

The Greatest Show on the Planet. BIOL122 is a suitable introductory science course for all students. It offers an integrative approach to the amazing world of behaviour. Basic mechanisms are covered, together with function and evolution. Lecture topics include micro- and macro-evolution, evolutionary origins of behaviour, basic neuroscience, perception, learning, brain and behaviour, and topics in animal behaviour. Lectures culminate with some reflections on the lives of humans in our modern world and the role of culture in human evolution.

For prospective Masters and PhD students