Ken Cheng: Studies in animal behaviour

Ken Cheng: Studies in animal behaviour

Research

Our research group crosses mechanistic, functional, and evolutionary questions in the study of animal behaviour. We mostly use experimental techniques implemented in the natural habitats of animals to investigate how they process information. A large part of our research concentrates on how animals negotiate space.

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Current projects and Research stories over the years


People

portrait ChengKen Cheng

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

email: ken.cheng@mq.edu.au
Google Scholar page: https://scholar.google.com.au/citations?user=e2wS-jQAAAAJ&hl=en&oi=ao

Photo by Tia Curnoe-Cheng

Ken Cheng has studied animal behaviour for some 40 years. A large part of the research has been and continues to be on spatial behaviour. Over his career, he has studied humans, monkeys, rats, a number of species of birds, honeybees, desert ants, and bull ants. Starting this century, Cheng’s team has been studying navigation and learning in an Australian desert ant located in Central Australia. Known as the red honey ant (the left two photos in the banner above), Melophorus bagoti shares many characteristics with the much studied North African desert ants of the genus Cataglyphis. It is long-legged, active in the heat of the day, moves fast, and deploys a suite of navigational strategies. Cheng is the author of three books, a reference book for a general audience on animal cognition, a textbook on the biological basis of behaviour, and a small, short guide on scientific writing, aimed at postgraduate students or early-career scientists.

How animals think and feel (2016)

This book, published by ABC-CLIO, Santa Barbara, USA, is a introductory book on animal cognition suitable for high-school and first-year university students. 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.

Biological basis of behaviour (2018)

This is an e-textbook published by Top Hat, Toronto, Canada, explicitly written for the first-year students that Cheng teaches, in a course bearing the name of the textbook. The e-textbook is a good deal cheaper than printed textbooks, and contains lots of colour figures as well as movies.

Expository scientific writing: A short guide (2019)

This little self-published book is meant for those writing scientific papers who might need some help, postgraduate students or early-career scientists or those for whom English is a foreign language. It is offered here free of charge.

Muzahidul (Muzahid) Islam

PhD student

Department of Biological Sciences
Macquarie University
Sydney NSW 2109

Australia

Email: muzahid.masum1@gmail.com

MSc (Entomology): University of Dhaka, Bangladesh

BSc (Honors): Department of Zoology, University of Dhaka, Bangladesh

Muzahid Islam is interested in how behaviour is generated by the interaction of brain, body, and environment. In nature, central-place insect foragers are efficient navigators between the nest and their foraging site, guided by a combination of innate strategies and learnt information from the surroundings. Muzahid’s research focuses on learning and visual navigation in the Australian nocturnal bull ant, Myrmecia midas. Currently, he is investigating learning and visual scanning in both familiar and unfamiliar environments, and what this learning and scanning contributes to navigation.

Sudhakar DeetiSudkakar Deeti

PhD student

Department of Biological Sciences
Macquarie University
Sydney NSW 2109

Australia

Email: sudhadeeti@gmail.com

Sudhakar Deeti is studying both the red honey ant, Melophorus bagoti, and the night-active bull ant, Myrmecia midas. He has studied learning walks in M. bagoti. These are small excursions around the nest performed by would-be foragers before they head off for the serious and risky business of foraging. Scanning the environment forms a part of learning walks, and Deeti is also studying scanning behaviours in M. midas.

Tim Pearson

PhD student

Department of Biological Sciences
Macquarie University
Sydney NSW 2109

Australia

Email: p.alecto@gmail.com

Tim Pearson trained initially as a mechanical engineer, and then progressed to a long career in the world of corporate IT. A lifelong interest in the environment and wildlife (particularly bats) led to his spending his spare time volunteering in conservation and wildlife rescue work; and when an opportunity for a career change presented itself he retrained as a wildlife ecologist. This quickly became an all-consuming passion, and he now combines the skills learned over the course of three careers in the pursuit of a PhD studying communication in Australian flying-foxes—trying to work out just what it is they are saying to each other.

Pearson is usually found either giving talks educating the public about bats, or in the midst of a flying-fox camp struggling with an array of cameras and microphones, capturing the lives, loves, and conversations of flying-foxes. Given his constant involvement in activities related to the conservation of bats, it’s inevitable that he’s often referred to as “Batman”, but in spite of this he steadfastly refuses to wear either a cape, or his underwear on the outside.

Tim Pearson is studying the grey-headed flying-fox, Pteropus poliocephalus, which is found in the Sydney area. His thesis concerns the ecological acoustics of this threatened flying mammal. The species is often found in urban settings, near people's homes. The sounds that flying-foxes make can impact humans.


Current projects

A number of our current projects focus on learning and navigation, in an enterprise we have called the experimental ethology of learning to navigate. A former lab member, Dr. Cody Freas, blazed the trail on this work with studies on the red honey ant Melophorus bagoti (Freas & Cheng, 2017; Freas, Whyte, & Cheng, 2017; Freas and Cheng, 2018a, 2018b; Freas, Fleischmann, & Cheng, 2019; see also Wystrach, Schwarz, Graham, & Cheng, 2019). We are continuing to examine the step-by-step and trial-by-trial process by which ants learn about their visual panorama. Research on the behaviour of the red honey ant has proceeded in earnest at our field sites for some 15 years now. We have learned much about the ant's foraging ecology and its navigational prowess. See our Research Stories page for a taste of the field work.

Photo: Desert ants Melophorus bagoti, known as red honey ants. The big red one with wings is a queen. The black winged ants are males, and the rest are workers. Photo by Patrick Schultheiss.

Learning walks in desert ants M. bagoti

Sudhakar Deeti

In the desert ants M. bagoti, we are examining the nature of their learning walks, a project carried out by Sudhakar Deeti. These are small forays that would-be foragers perform around their nest before they head off on the serious business of foraging. The word “perform” strikes a pun in these walks as dance-like movements are part of parcel of the act; these include voltes, really tight little turns, and pirouettes, sharp, saccade-like turns of the whole body on one spot. This lingo borrowed from dancing is now the formal terminology. How do these ants figure out which direction is which in their learning walks? What do they learn in their learning walks?

Photo: The habitat where red honey ants Melophorus bagoti live, here on the grounds of the Centre for Appropriate Technology just south of the town centre of Alice Springs, Northern Territory. The scene is visually rich, with tussocks, bushes, trees, and distant mountains filling the panorama. We often make grids to plot the paths travelled by ants. Photo by Sudhakar Deeti.


The nocturnal bull ants Myrmecia midas, as well as other bull ants, also perform turn-and-scan routines, often when they are released from a tube on an experimental test. A project carried out by Muzahid Islam and Sudhakar Deeti is examining these scans at the start of journeys.

Detour learning in the nocturnal bull ant, Myrmecia midas on their foraging tree

Muzahid Islam

In their foraging life, ants can learn idiosyncratic foraging routes. In another study of learning, we challenged Myrmecia midas ants with a physical barrier along the vertical part of their navigational route, on the tree on which they forage. Myrmecia midas forages on nearby eucalyptus trees from evening twilight to morning twilight searching for and collecting food. The study is on individuals that climb up the tree near which their nest is located, called nest-tree foragers, during ascending navigation. The objectives of this detour learning study are:

1. to examine the navigational behaviour of the tree-foraging nocturnal bull ant when it encounters an obstacle on its foraging tree;

2. to learn how the individual foragers deal with the obstacle over repeated runs.

barrier on treeobstacle on groundPhotos: Left, bull ants are confronted with a barrier extending 180 degrees around the tree on which they forage each night. This barrier was on the nest tree, the tree near which the nest was found, and it was centred at the nest location, hence on the side on which all of the foragers on this tree ascend. Right, in their natural foraging life, these bull ants encounter debris on the ground that dot their landscape with natural obstacles. Photos by Muzahid Islam.

Learning in nocturnal bull ants: Visual scanning behaviour in familiar and unfamiliar environments

Muzahid Islam

Social insects such as ants are efficient navigators, guided by a combination of innate strategies and learnt information from the surroundings. At the early stage, naïve ants perform learning walks around the nest entrance to acquire visual information. This walk contains repeated turns with distinct stopping phases of the body. Experienced foragers also perform turning with stopping phases to scan the environment when they have difficulty pinpointing the goal. Foragers stop and scan the surrounding environment by turning on the spot before departing in their chosen direction. But when the visual environment is unfamiliar, foragers perform more scanning.

Our aim are:

1. to characterise the details of visual scanning in Myrmecia midas in both natural and experimentally altered conditions;

2. to investigate how the bull ants learn to navigate in an unfamiliar environment over multiple trips.


Urban ethology of flying foxes

Tim Pearson

flying foxesblackheaded flying foxTim Pearson is completing a thesis on the ecological acoustics of flying-foxes, taking the lab into the world of urban ethology. Grey-headed flying-foxes, Pteropus poliocephalus, are found in the Sydney area. Far less commonly, black-headed flying-foxes, Pteropus alecto, may also be found. Flying-foxes are often found in urban settings, near people's homes.

Photos: Grey-headed flying-foxes (left) and black-headed flying-foxes (right), in their typical poses hanging from tree branches. Photos by Tim Pearson.

urban settingThe 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.

Photo: In this photo taken from the balcony of a suburban home, the flying-fox camp is literally just beyond the back yard. Photo by Tim Pearson.

flying fox in urban setting male flying foxesTim 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.

Photos: Left, grey-headed flying-foxes at the Gordon Reserve in suburban Sydney. Right, male grey-headed flying foxes bickering (in a combative interaction). Photos by Tim Pearson.


Reconstructed reality in an antarium

photo of antariumIn a different line of projects across multiple labs, virtual reality, or rather, what we call reconstructed reality plays the leading role. Ants are tested in a rig we call an antarium. The antarium consists of a dome that projects LED lights inside emitting green, blue, and ultraviolet wavelengths, in the range of the electromagnetic spectrum that ants’ visual systems are sensitive to. The tested ant would stand on a trackball floating on air, held on the ball by a lead attached to its head. The lead is loose on the ant’s head, although the lead, and hence the ant, can hardly move horizontally (or vertically for that matter) in physical space. Instead, as the ant holding its own weight on the ball walks, the ball spins under it.

It took an engineer by training, Zoltan Kosci at the Australian National University, a hefty effort to put the antarium together and make it work. Kosci is pursuing a PhD in biological sciences under the supervision of Professor Jochen Zeil.

Photo: This is a photo of the antarium, a dome of panels emitting green, blue, and ultraviolet light from LEDs, designed to present a panoramic view to an ant inside it standing on a track ball. Photo by Trevor Murray.

cartoon of antariumWe could create any visual scene to present to the ants, but the main idea is to take ants living just outside a lab, and re-make the visual world it would see were it say, at its nest. The software allows the visual world to change as it would outside in the real world, as the ant walks. This is why the system is called reconstructed reality, as contrasted with virtual reality. Of course, the antarium screen could also present altered reality. We can chop down trees and move mountains with software.

Photo: This drawing shows the inside of the antarium, showing a scene projected to an ant. In the set up, the animal walks on a track ball, with the ant supporting its own weight. As it moves, the scenery changes in response to the path taken by the ant. The inset shows a bull ant, of the genus Myrmecia, on a track ball. In operation, the track ball floats on air. The ant has to support its own weight on the track ball; it is not tethered to the rod above it. A short piece of string between the ant and the rod makes the ant support its own weight. This track-ball set up reproduces much of the natural conditions for walking, and the system can also be used in the natural world outside of the antarium. Photo by Jochen Zeil.


Publications

Books

Cheng, K. (2016).  How animals think and feel. ABC-CLIO, Santa Barbara.

Cheng, K. (2018).  Biological basis of behaviour [e-textbook].  Top Hat https://tophat.com/, Toronto.

Cheng, K. (2019). Expository scientific writing: A short guide. Online book, copyright 2019.

Refereed Journal Papers

Kroger, R.O., Cheng, K., & Leong, I. (1979).  Are the rules of address universal?  A test of Chinese usage.  Journal of Cross-Cultural Psychology, 10, 395–414.

Cheng, K. (1986).  A purely geometric module in the rat’s spatial representation.  Cognition, 23, 149-178.

Cheng, K., Collett, T.S., & Wehner, R. (1986). Honeybees learn the colours of landmarks.  Journal of Comparative Physiology A, 159, 69–73.

Cheng, K., Collett, T.S., Pickhard, A., & Wehner, R. (1987).  The use of visual landmarks by honeybees: Bees weight landmarks according to their distance from the goal.  Journal of Comparative Physiology A, 161, 469–475.

Cheng, K. (1988).  Some psychophysics of the pigeon’s use of landmarks.  Journal of Comparative Physiology A, 162, 815–826.

Cheng, K. (1989).  The vector sum model of pigeon landmark use.  Journal of Experimental Psychology: Animal Behavior Processes, 15, 366–375.

Cheng, K., & Roberts, W.A. (1989).  Timing multimodal events in pigeons.  Journal of the Experimental Analysis of Behavior, 52, 363–376.

Roberts, W.A., Cheng, K., & Cohen, J.S. (1989). Timing light and tone signals in pigeons.  Journal of Experimental Psychology: Animal Behavior Processes, 15, 23–35.

Cheng, K. (1990).  More psychophysics of the pigeon’s use of landmarks.  Journal of Comparative Physiology A, 166, 857–863.

Cheng, K., & Roberts, W. (1991).  Three psychophysical principles of timing in pigeons.  Learning and Motivation, 22, 112–128.

Cheng, K. (1992).  The form of timing distributions in pigeons under penalties for responding early.  Animal Learning & Behavior, 20, 112–120.

Spetch, M.L., Cheng, K., & Mondloch, M.V. (1992). Landmark use by pigeons in a touch-screen spatial search task.  Animal Learning & Behavior, 20, 281–292.

Cheng, K., & Sherry, D.F. (1992).  Landmark–based spatial memory in birds: The use of edges and distances to represent spatial positions.  Journal of Comparative Psychology, 106, 331–341.

Cheng, K., & Westwood, R. (1993).  Analysis of single trials in pigeons’ timing performance.  Journal of Experimental Psychology: Animal Behavior Processes, 19, 56–67.

Cheng, K., Westwood R., & Crystal, J.D. (1993). Memory variance in the peak procedure of timing in pigeons.  Journal of Experimental Psychology: Animal Behavior Processes, 19, 68–76.

Cheng, K. (1994).  The determination of direction in landmark-based spatial search in pigeons: A further test of the vector sum model.  Animal Learning & Behavior, 22, 291–301.

Cheng, K. & Spetch, M.L. (1995).  Stimulus control in the use of landmarks by pigeons in a touch-screen task. Journal of the Experimental Analysis of Behavior, 63, 187–201.

Cheng, K., & Miceli, P. (1996).  Modelling timing performance on the peak procedure.  Behavioural Processes, 37, 137–156.

Cheng, K., Spetch, M.L. & Miceli, P. (1996).  Averaging temporal duration and spatial position.  Journal of Experimental Psychology: Animal Behavior Processes, 22, 175–182.

Spetch, M.L., Cheng, K., & MacDonald, S.E. (1996).  Learning the configuration of a landmark array I: Touch‑screen studies with pigeons and humans.  Journal of Comparative Psychology, 110, 55–68.

Cheng, K., Spetch. M.L., & Johnston, M. (1997).  Spatial peak shift and generalization in pigeons. Journal of Experimental Psychology: Animal Behavior Processes, 23, 469–481.

Spetch, M.L., Cheng, K., MacDonald, S.E., Linkenhoker, B.A., Kelly, D.M., & Doerkson, S.R. (1997).  Learning the configuration of a landmark array in humans and pigeons II: Generality across search tasks.  Journal of Comparative Psychology, 111, 14–24.

Cheng, K. (1998).  Distances and directions are computed separately by honeybees in landmark-based search.  Animal Learning & Behavior, 26, 455–468.

Cheng, K. (1998).  Honeybees (Apis mellifera) remember two near-target landmark constellations. Learning and Motivation, 29, 435–443.

Brodbeck, D.R., Hampton, R.R., & Cheng, K. (1998).  Timing behavior of black-capped chickadees.  Behavioural Processes, 44, 183–195.

Spetch, M.L., & Cheng, K. (1998).  A step function in pigeons’ temporal generalization in the peak shift task. Animal Learning & Behavior, 26, 103–118.

Cheng, K. (1999).  Landmark-based spatial search in honeybees: I. Use of elements and interlandmark angles.  Animal Cognition, 2, 73–78.

Cheng, K. (1999).  Landmark-based spatial search in honeybees: II. Using gaps and blocks. Animal Cognition, 2, 79–90.

Cheng, K. (1999).  Spatial generalization in honeybees confirms Shepard’s law. Behavioural Processes, 44, 309–316.

Cheng, K., Srinivasan, M.V., & Zhang, S.W. (1999).  Error is proportional to distance measured by honeybees: Weber’s law in the odometer.  Animal Cognition, 2, 11–16.

Srinivasan, M.V., Zhang, S.W., Berry, J., Cheng, K., & Zhu, H. (1999). Honeybee navigation: linear perception of short distances travelled.  Journal of Comparative Physiology A, 185, 239–245.

Cheng, K. (2000).  How honeybees find a place: Lessons from a simple mind.  Animal Learning & Behavior, 28, 1–15.

Cheng, K. (2000).  Shepard’s universal law supported by honeybees in spatial generalization.  Psychological Science, 11, 403–408.

Evans, C.S., Wenderoth, P.M., & Cheng, K. (2000).  Detection of bilateral symmetry in complex biological images.  Perception, 29, 31–42.

Cheng, K., & Spetch, M.L. (2001).  Blocking in landmark-based search in honeybees.  Animal Learning & Behavior, 29, 1–9.

Kamil, A.C., & Cheng, K. (2001).  Way-finding and landmarks: The multiple-bearings hypothesis.  Journal of Experimental Biology, 204, 103–113.

Kamil, A.C., Goodyear, A.J., & Cheng, K. (2001).  The use of landmarks by Clark’s nutcrackers: first tests of a new model.  The Journal of Navigation, 54, 429–435.

Cheng, K. (2002).  Generalisation: Mechanistic and functional explanations.  Animal Cognition, 5, 33–40.

Cheng, K., Peña, J., Porter, M.A., & Irwin, J.D. (2002).  Self control in honeybees.  Psychonomic Bulletin & Review, 9, 259–263.

Cheng, K., & Spetch, M.L. (2002).  Spatial generalization and peak shift in humans.  Learning and Motivation, 33, 358–389.

Cheng, K., & Wehner, R. (2002).  Navigating desert ants (Cataglyphis fortis) learn to alter their search patterns on their homebound journey.  Physiological Entomology, 27, 285–290.

Heiling, A.M., Cheng, K., &. Herberstein, M.E. (2004).  Exploitation of floral signals by crab spiders (Thomisus spectabilis, Thomisidae).  Behavioral Ecology, 15, 321–326.

MacDonald, S.E., Spetch, M.L., Kelly, D.M., & Cheng, K. (2004).  Strategies in landmark use by children, adults, and marmoset monkeys.  Learning and Motivation, 35, 322–347.

Spetch, M.L., Cheng, K., & Clifford, C.W.G. (2004).  Peak shift but not range effects in face recognition.  Learning and Motivation, 35, 221–241.

Bingman, V.P., & Cheng, K. (2005). Mechanisms of animal global navigation: Comparative perspectives and enduring challenges.  Ethology Ecology & Evolution, 17, 295–318.

Cheng K. (2005).  Reflections on geometry and navigation.  Connection Science, 17, 5–21 (special issue on navigation edited by K.J. Jeffery).

Cheng, K. (2005).  Goldfish matching geometric and featural cues: A re-interpretation of some of the data of Vargas López, Salas, and Thinus-Blanc (2004).  Journal of Comparative Psychology, 119, 455–457.

Cheng, K. (2005). Context cues eliminate retroactive interference effects in honeybees (Apis mellifera).  Journal of Experimental Biology, 208, 1019–1024.

Cheng, K., & Galllistel, C.R. (2005). Shape parameters explain data from spatial transformations: Comment on Pearce et al. (2004) and Tommasi and Polli (2004).  Journal of Experimental Psychology: Animal Behavior Processes, 31, 254–259.

Cheng, K., & Newcombe, N.S. (2005).  Is there a geometric module for spatial orientation? Squaring theory and evidence.  Psychonomic Bulletin & Review, 12, 1–23.

Heiling, A.M., Cheng, K., Chittka, L., Goeth, A., & Herberstein, M.E. (2005).  The role of UV in crab spider signals: effects on perception by prey and predators. Journal of Experimental Biology, 208, 3925–3931.

Heiling, A.M., Chittka, L., Cheng, K., &. Herberstein, M.E. (2005). Colouration in crab spiders: substrate choice and prey attraction.  Journal of Experimental Biology, 208, 1785–1792.

Oaten, M., & Cheng, K. (2005).  Academic stress impairs self-control.  Journal of Social and Clinical Psychology, 24, 254–279.

Cheng, K., Narendra, A, & Wehner, R. (2006).  Behavioral ecology of odometric memories in desert ants: Acquisition, retention, and integration.  Behavioral Ecology, 17, 227–235.

Cheng, K., Spetch, M.L., Kelly, D.M., & Bingman, V.P. (2006).  Small-scale spatial cognition in pigeons.  Behavioural Processes, 72, 115–127.

Cheng, K., & Wignall, A.E. (2006). Honeybees (Apis mellifera) holding on to memories: Response competition causes retroactive interference effects.  Animal Cognition, 9, 141–150.

Heiling, A.M., Cheng, K., & Herberstein, M.E. (2006). Picking the right spot: crab spiders position themselves on flowers to maximize prey attraction.  Behaviour, 143, 957–968.

Oaten, M., & Cheng, K. (2006). Improved self-control: The benefits of a regular program of academic study.  Basic and Applied Social Psychology, 28, 1–16.

Oaten, M., & Cheng, K. (2006).  Longitudinal gains in self-regulation from regular physical exercise.  British Journal of Health Psychology, 11, 717–733.

Verbeek, E.L., Spetch, M.L., Cheng, K., & Clifford, C.W.G. (2006).  Determinants of range  effects in face recognition.  Learning & Behavior, 34, 229–240.

Wignall, A.E., Heiling, A.M., Cheng, K., & Herberstein, M.E. (2006).  Flower symmetry preferences in honeybees and their crab spider predators.  Ethology, 112, 510–518.

Cheng, K., Shettleworth, S.J., Huttenlocher, J., & Rieser, J.J. (2007).  Bayesian integration of spatial information.  Psychological Bulletin, 133, 625–637.

Narendra, A., Cheng, K., & Wehner, R. (2007).  Acquiring, retaining and integrating memories of the outbound distance in the Australian desert ant Melophorus bagoti.  Journal of Experimental Biology, 210, 570–577.

Narendra, A., Si, A., Sulikowski, D, & Cheng, K. (2007).  Learning, retention and coding of nest-associated visual cues by the Australian desert ant, Melophorus bagoti.  Behavioral Ecology and Sociobiology, 61, 1543–1553.

Oaten, M., & Cheng, K. (2007).  Improvements in self-control from financial monitoring.  Journal of Economic Psychology, 28, 487–501.

Rao, D., Cheng, K., & Herberstein, M.E. (2007).  A natural history of web decorations in the St. Andrew’s Cross spider (Argiope keyserlingi).  Australian Journal of Zoology, 55, 9–14.

Cheng, K. (2008). Whither geometry? Troubles of the geometric module.  TRENDS in Cognitive Sciences, 12, 355–361.

Cheng, K., Simpson, S.J., & Raubenheimer, D. (2008).  A geometry of regulatory scaling.  American Naturalist, 172, 681–693.

Cheung, A., Stürzl, W., Zeil, J., & Cheng, K. (2008).  Information content of panoramic images: II. View-based navigation in nonrectangular experimental arenas. Journal of Experimental Psychology: Animal Behavior Processes, 34, 15–30.

Narendra, A., Cheng, K., Sulikowski, D., & Wehner, R. (2008).  Search strategies of ants in landmark-rich habitats.  Journal of Comparative Physiology A, 194, 929–938.

Prabhu, C., & Cheng, K. (2008).  One day is all it takes: circadian modulation of the retrieval of colour memories in honeybees.  Behavioral Ecology and Sociobiology, 63, 11–22.

Prabhu, C., & Cheng, K. (2008).  Recency preference of odour memory retrieval in honeybees.  Behavioral Ecology and Sociobiology, 63, 22–32.

Rao, D., Cheng, K., & Herberstein, M.E. (2008).  Stingless bee response to spider webs is dependent on the context of encounter.  Behavioral Ecology and Sociobiology, 63, 209–216.

Stürzl, W., Cheung, A., Cheng, K., & Zeil, J. (2008).  Information content of panoramic images: I. Rotational errors and the similarity of views in rectangular arenas.  Journal of Experimental Psychology: Animal Behavior Processes, 34, 1–14.

Cheng, K., Narendra, A., Sommer, S., & Wehner, R. (2009).  Traveling in clutter: Navigation in the Central Australian desert ant Melophorus bagoti.  Behavioural Processes, 80, 261–268.

Graham, P., & Cheng, K. (2009).  Ants use the panoramic skyline as a visual cue during navigation.  Current Biology, 19, R935–R937.

Graham, P., & Cheng, K. (2009). Which portion of the natural panorama is used for view based navigation in the Australian desert ant?  Journal of Comparative Physiology A, 195, 681–689.

Herberstein, M.E., Heiling, A., & Cheng, K. (2009).  Evidence for UV-based sensory exploitation in Australian but not European crab spiders. Evolutionary Ecology, 23, 621–634.

Cheng, K., Spetch, M. L., & Hoan, A. (2010).  Categories and range effects in human spatial memory.  Frontiers in Psychology, 1, 231. Doi: 10.3899/fpsyg.2010.00231

Kartiko, I., Kavakli, M., & Cheng, K. (2010).  Learning science in a virtual reality application: The impacts of animated-virtual actors’ visual complexity.  Computers & Education, 55, 881–891.

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.

Legge, E. L. G., Spetch, M. L., & Cheng, K. (2010).  Not using the obvious: desert ants, Melophorus bagoti, learn local vectors but not beacons in an arena.  Animal Cognition, 13, 849-860, DOI 10.1007/s10071-010-0333-x.

Schwarz, S., & Cheng, K. (2010).  Visual associative learning in two desert ant species.  Behavioral Ecology and Sociobiology, 64, 2033–2041, DOI 10.1007/s00265-010-1016-y.

Bühlmann, C., Cheng, K., & Wehner, R. (2011).  Vector-based and landmark-guided navigation in desert ants inhabiting landmark-free and landmark-rich environments.  Journal of Experimental Biology, 214, 2845–2853.

Philippides, A., Baddeley, B., Cheng, K., & Graham, P. (2011).  How might ants use panoramic views for route navigation?  Journal of Experimental Biology, 214, 445–451.

Schultheiss, P., & Cheng, K. (2011).  Finding the nest: inbound searching behaviour in the Australian desert ant Melophorus bagoti.  Animal Behaviour, 81, 1031–1038.

Schwarz, S., Albert, L., Wystrach, A., & Cheng, K. (2011).  Ocelli contribute to the encoding of celestial compass information in the Australian desert ant Melophorus bagoti.  Journal of Experimental Biology, 214, 901–906. Doi: 10.1242/jeb.049262

Schwarz, S., & Cheng, K. (2011).  Visual discrimination, sequential learning and memory retrieval in the Australian desert ant Melophorus bagoti.  Animal Cognition, 14, 861–870. DOI 10.1007/s10071-011-0419-0

Schwarz, S., Wystrach, A., & Cheng, K. (2011).  A new navigational mechanism mediated by ant ocelli.  Biology Letters, 7, 856–858, doi:10.1098/rsbl.2011.0489

Wystrach, A., Beugnon, G., & Cheng, K. (2011). Landmarks or panoramas: what do navigating ants attend to for guidance?  Frontiers in Zoology, 8, 21.

Wystrach, A., Cheng, K., Sosa, S., & Beugnon, G. (2011).  Geometry, features, and panoramic views: Ants in rectangular arenas. Journal of Experimental Psychology: Animal Behavior Processes, 37, 420–435. DOI: 10.1037/a0023886

Wystrach, A., Schwarz, S., Schultheiss, P., Beugnon, G., & Cheng, K. (2011).  Views, landmarks, and routes: how do desert ants negotiate an obstacle course?  Journal of Comparative Physiology A, 197, 167–179, DOI 10.1007/s00359-010-0597-2.

Cheng, K. (2012). How to navigate without maps: The power of taxon-like navigation in ants.  Comparative Cognition & Behavior Reviews, 7, 1–22.

Cheng, K., Middleton, E. J. T., & Wehner, R. (2012). Vector-based and landmark-guided navigation in desert ants of the same species inhabiting landmark-free and landmark-rich environments. Journal of Experimental Biology, 215, 3169–3174.

Schultheiss, P., Schwarz, S., Cheng, K., & Wehner, R. (2012). Foraging ecology of an Australian salt-pan desert ant (genus Melophorus).  Australian Journal of Zoology, 60, 311–319.

Schwarz, S., Schultheiss, P., & Cheng, K. (2012).  Visual cue learning and odometry in guiding the search behavior of desert ants Melophorus bagoti in artificial channels.  Behavioural Processes, 91, 298–303.

Wystrach, A., Beugnon, G., & Cheng, K. (2012). Ants might use different view-matching strategies on and off the route.  Journal of Experimental Biology, 215, 44–55.

Cheng, K., Huttenlocher, J., & Newcombe, N. S. (2013). 25 years of research on the use of geometry in spatial reorientation: A current theoretical perspective.  Psychonomic Bulletin & Review, 20, 1033–1054.

Perry, C. J., Barron, A. B., & Cheng, K. (2013).  Invertebrate learning and cognition: relating phenomena to neural substrate.  WIREs Cognitive Science, doi: 10.1002/wcs.1248.

Reynolds, A. M., Schultheiss, P., & Cheng, K. (2013).  Are Lévy flight patterns derived from the Weber–Fechner law in distance estimation?  Behavioral Ecology and Sociobiology, 67, 1219-1226 (DOI 10.1007/s00265-013-1549-y).

Schultheiss, P., & Cheng, K. (2013).  Finding food: outbound searcing behaviour in the Australian desert ant Melophorus bagoti.  Behavioral Ecology, 24, 128–135.

Schultheiss, P., Wystrach, A., Legge, E. L. G., & Cheng, K. (2013).  Information content of visual scenes influences systematic search of desert ants.  Journal of Experimental Biology, 216, 742–749.  doi: 10.1242/jeb.075077.

Wystrach, A., Schwarz, S., Baniel, A., & Cheng, K. (2013).  Backtracking behaviour in lost ants: an additional strategy in their navigational toolkit.  Proceedings of the Royal Society B: Biological Sciences, 280, 20131677.

Cheng, K., Schultheiss, P., Schwarz, S., Wystrach, A., & Wehner, R. (2014).  Beginnings of a synthetic approach to desert ant navigation.  Behavioural Processes, 102, 51–61 http://dx.doi.org/10.1016/j.beproc.2013.10.001.

Reynolds, A. M., Schultheiss, P., & Cheng, K. (2014).  Does the Australian desert ant Melophorus bagoti approximate a Lévy search by an intrinsic bi-modal walk?  Journal of Theoretical Biology, 340, 17–22.

Kelaiah, I., Kavakli, M., & Cheng, K. (2014).  Associations between simulator sickness and visual complexity of a virtual scene.  Frontiers in Psychological and Behavioral Science, 3, 27–35.

Wystrach, A., Schwarz, S., Schultheiss, P., Baniel, A., & Cheng, K. (2014).  Multiple sources of celestial compass information in the Central Australian desert ant Melophorus bagoti.  Journal of Comparative Physiology A, 200, 591–601.

Julle-Daniere, E., Schultheiss, P., Wystrach, A., Schwarz, S., Nooten, S. S., Bibost, A.-L., & Cheng, K. (2014).  Visual matching in the orientation of desert ants (Melophorus bagoti): The effect of changing skyline height.  Ethology, 120, 783–792.

Wystrach, A., Philippides, A., Aurejac, A., Cheng, K., & Graham, P. (2014).  Visual scanning behaviours and their role in the navigation of the Australian desert ant Melophorus bagoti.  Journal of Comparative Physiology A, 200, 615–626.

Andrew, S. C., Perry, C. J., Barron, A. B., Berthon, K., Peralta, V., & Cheng, K. (2014).  Peak shift in honey bee olfactory learning.  Animal Cognition, 17, 1177–1186.

Schwarz, S., Julle-Daniere, E., Morin, L., Schultheiss, P., Wystrach, A., Ives, J., & Cheng, K. (2014). Desert ants (Melophorus bagoti) navigating with robustness to distortions of the natural panorama.  Insectes Sociaux, 61, 371–383. DOI 10.1007/s00040-014-0364-4

Legge, E. L. G., Wystrach, A., Spetch, M. L., & Cheng, K. (2014). Combining sky and Earth: Desert ants (Melophorus bagoti) show weighted integration of celestial and terrestrial cues.  Journal of Experimental Biology, 217, 4159–4166.

Cheng, K., & Freas, C. A. (2015).  Path integration, views, search, and matched filters: The contributions of Rüdiger Wehner to the study of orientation and navigation.  Journal of Comparative Physiology A, 201, 517–532.

Schultheiss, P., Reynolds, A. M., & Cheng, K. (2015).  Searching behavior in social Hymenoptera.  Learning and Motivation, 50, 59–67.

Chang, L.-H., Barron, A. B., & Cheng, K. (2015).  Effects of the juvenile hormone analogue methoprene on rate of behavioural development, foraging performance and navigation in honey bees (Apis mellifera).  Journal of Experimental Biology, 218, 1715–1724.

Ushitani, T., Perry, C. J., Cheng, K., & Barron, A. B. (2016). Accelerated behavioural development changes fine-scale search behaviour and spatial memory in honey bees (Apis mellifera L.).   Journal of Experimental Biology, 219, 412–418.

Schultheiss, P., Wystrach, A., Schwarz, S., Tack, A., Delor, J., Nooten, S. S., Bibost, A.-L., Freas, C. A., & Cheng, K. (2016). Crucial role of ultraviolet light for desert ants in determining direction from the terrestrial panorama.  Animal Behaviour, 115, 19–28.

Schultheiss, P., Stannard, T., Pereira, S., Reynolds, A. M., Wehner, R., & Cheng, K. (2016). Similarities and differences in path integration and search in two species of desert ants inhabiting a visually rich and a visually barren habitat.  Behavioral Ecology and Sociobiology, 70, 1319–1329.

Wehner, R., Hoinville, T., Cruse, H., & Cheng, K. (2016).  Steering intermediate courses: desert ants combine information from various navigational routines.  Journal of Comparative Physiology A, 202, 459–472.

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

Freas, C. A., Whyte, C., & Cheng, K. (2017).  Skyline retention and retroactive interference in the navigating Australian desert ant, Melophorus bagoti.  Journal of Comparative Physiology A, 203, 353–367.

Freas, C. A., & Cheng, K. (2017). Learning and time-dependent cue choice in the desert ant, Melophorus bagoti.  Ethology, 123, 503–515.

Freas, C. A., Narendra, A., Lemesle, C., & Cheng, K. (2017).  Polarized light use in the nocturnal bull ant, Myrmecia midas.  Royal Society Open Science, 4, 170598.  http://rsos.royalsocietypublishing.org/content/4/8/170598

Schwarz, S., Wystrach, A., & Cheng, K. (2017).  Ants’ navigation in an unfamiliar environment is influenced by their experience of a familiar route.  Scientific Reports, 7, 14161.  https://www.nature.com/articles/s41598-017-14036-1

Cheng, K. (2018).  Cognition beyond representation: Varieties of situated cognition in animals.  Comparative Cognition & Behavior Reviews, 13, 1–20.

Cheng, K. (2018).  More situated cognition in animals: Reply to commentators. Comparative Cognition & Behavior Reviews, 13, 49–54.

Freas, C. A., & Cheng, K. (2018). Limits of vector calibration in the Australian desert ant, Melophorus bagoti.  Insectes Sociaux, 65, 141–152.

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. https://www.frontiersin.org/articles/10.3389/fpsyg.2018.00016/full

Ferdous, Mst J., Reynolds, A. M., & Cheng, K. (2018).  Distinguishing between aapparent and actual randomness: A preliminary examination with Australian ants.  Behavioral Ecology and Sociobiology, 72, 113. https://doi.org/10.1007/s00265-018-2527-1

Cheng, K., & Byrne, R. W. (2018). Why human environments enhance animal capacities to use objects: Evidence from keas (Nestor notabilis) and apes (Gorilla gorilla, Pan paniscus, Pongo abelii, Pongo pygmaeus).  Journal of Comparative Psychology, 132, 419–426.

Freas, C. A., & Cheng, K. (2018). Landmark learning, cue conflict, and outbound view sequence in navigating desert ants.  Journal of Experimental Psychology: Animal Learning and Cognition, 44, 409–421.

Freas, C. A., Fleischmann, P. N., & Cheng, K. (2019).  Experimental ethology of learning in desert ants: Becoming expert navigators.  Behavioural Processes, 158, 181–191.

Wystrach, A., Schwarz, S., Graham, P., & Cheng, K. (2019).  Running paths to nowhere: Repetition of routes shows how desert ants modulate online the weights accorded to cues.  Animal Cognition, 22, 213-222.

Kelly, D. M., Cheng, K., Balda, R., & Kamil, A. C. (2019).  The effects of sun compass error on spatial search in Clark’s nutcrackers.  Integrative Zoology, 14, 172-181.

Freas, C. A., & Cheng, K. (2019).  Panorama similarity and navigational knowledge in the nocturnal bull ant Myrmecia midas.  Journal of Experimental Biology, in press.

Reviews and Short Communications

Gallistel, C.R., & Cheng, K. (1985).  A modular sense of place?  (A comment on J.A. Fodor’s precis of The Modularity of Mind.)  The Behavioral and Brain Sciences, 8, 11–12.

Cheng, K. (1998).  Review of W.A. Roberts, The Principles of Animal Cognition (McGraw-Hill, 1998). Animal Behaviour, 55, 1409–1410.

Cheng, K. (2001).  Object identity and Tinbergen’s four whys. Cahiers de Psychologie Cognitive, 20, 193–199.

Cheng, K. (2001).  Generalization and Tinbergen’s four whys.  Behavioral and Brain Sciences, 24, 660–661.

Cheng, K. (2004).  Review of K.J. Jeffery (Ed.), The neurobiology of spatial behaviour.  Animal Cognition, 7, 199–200.

Cheng, K. (2004).  What makes us tick: clocks in the brain.  Review of W.H. Meck (Ed.), Functional and neural mechanisms of interval timing. Animal Cognition, 7, 267–268.

Cheng, K. (2005).  Review of Evolutionary Psychology: An Introduction, by L. Workman & W. Reader, 2004.  Educational Psychology, 25, 439–440.

Cheng, K. (2005).  Promoting a Tinbergian approach to animal behavior.  Review of The Behavior of Animals: Mechanisms, Function, and Evolution, Johan J. Bolhuis and Luc-Alain Giraldeau (Eds.) (Malden, MA, Oxford, U.K., Carlton, Australia: Blackwell Publishing, 2004, 515 pages).  Canadian Journal of Experimental Psychology, 59, 287–289.

Cheng, K. (2010).  Review of Cognitive Ecology II, edited by R. Dukas & J. M. Ratcliffe. Chicago: The University of Chicago Press (2009).  Animal Behaviour, 79, 1401–1402.

Cheng, K. (2010).  Q & A.  Current Biology, 20, R463–464.

Kelly, D. M., & Cheng, K. (2014).  Editorial: A synthetic aproach to comparative cognition. Behavioural Processes, 102, 1–2.

Cheng, K., & Ronacher, B. (2015).  Editorial: A champion of organismal biology.  Journal of Comparative Physiology A, 201, 513–515.

Cheng, K. (2015).  Answer (in part) blowing in the wind
Comment on “Liberating Lévy walk research from the shackles of optimal foraging” by A. Reynolds. Physics of Life Reviews, 14, 90–93.

Cheng, K. (2016).  A new direction for grid cells.  Learning & Behavior, 44, 2–3.

Cheng, K. (2018).  Grid-like units help deep learning agent to navigate.  Learning & Behavior, online, https://doi.org/10.3758/s13420-018-0329-y

Chapters in Books

Cheng, K., & Gallistel, C.R. (1984).  Testing the geometric power of a spatial representation. In H.L. Roitblat, H.S. Terrace, & T.G. Bever (Eds.), Animal Cognition (pp. 409–423).  Hillsdale, NJ: Erlbaum.

Cheng, K. (1987).  Rats use the geometry of surfaces for navigation.  In P. Ellen & C. Thinus-Blanc (Eds.), Cognitive Processes and Spatial Orientation in Animal and Man.  Volume I: Experimental Animal Psychology and Ethology (pp. 153–159).  Dordrecht: Nijhoff.

Cheng, K. (1992).  Three psychophysical principles in the processing of spatial and temporal information.  In W.K. Honig & J.G. Fetterman (Eds.), Cognitive Aspects of Stimulus Control (pp. 69–88).  Hillsdale, NJ: Erlbaum.

Cheng, K.  (1995). Landmark‑based spatial memory in the pigeon.  In D.L. Medin (Ed.), The Psychology of Learning and Motivation, Vol. 33 (pp. 1–21).  New York: Academic Press.

Cheng, K., & Spetch, M.L. (1998).  Mechanisms of landmark use in mammals and birds.  In S.D. Healy (Ed.) Spatial Representation in Animals (pp. 1–17).  Oxford: Oxford University Press.

Cheng, K. (1999).  Rappresentazione della spatio.  In P.P.G. Bateson & E. Alleva (Eds.), The Biology of Behaviour (219–229).  Volume 4 of the series Frontiere della Vita, from Enciclopedia Italiana.

Cheng K., & Spetch, M.L. (2001). Landmark-based spatial memory in pigeons.  Cyberchapter in R.G. Cook (ed.) Avian Visual Cognition. http://www.pigeon.psy.tufts.edu/avc/cheng/default.htm

Cheng, K. (2006).  Arthropod navigation: Ants, bees, crabs, spiders finding their way. In E.A. Wasserman & T.R. Zentall (Eds.), Comparative cognition: Experimental explorations of animal intelligence (pp. 189–209). Oxford: Oxford University Press.

Cheng, K., & Newcombe, N.S. (2006).  Geometry, features, and orientation in vertebrate animals: A pictorial review.  In M.F. Brown & R.G. Cook (eds.), Comparative Spatial Cognition.  Cyberbook: http://www.pigeon.psy.tufts.edu/asc/.

Cheng, K. (2008).  Geometry and navigation.  In M.E. Jefferies & W.K. Yeap (eds.), Robotics and cognitive approaches to spatial mapping (pp. 145–161).  Berlin, Heidelberg: Springer Verlag.

Cheng, K., & Crystal, J.D. (2008). Learning to time intervals.  In R. Menzel (ed.), Learning theory and behavior (pp. 341–364).  Volume 1 of Learning and memory: A comprehensive reference, 4 volumes (J. Byrne ed.).  Oxford: Elsevier.

Cheng, K. (2010).  Common principles shared by spatial and other kinds of cognition.  In F. L. Dolins & R. W. Mitchell (Eds.), Spatial cognition, spatial perception: mapping the self and space (pp. 54–74).  Cambridge, New York: Cambridge University Press.

Cheng, K. (2010).  Behavioral ecology of insect cognition: universals and particulars. In W. Zeng & H. Liu (Eds.), Behavioral and chemical ecology (pp. 173–193). New York: Nova Science Publishers.

Wiener, J., Shettleworth, S., Bingman, V. P., Cheng, K., Healy, S., Jacobs, L. F., Jeffery, K. J., Mallot, H. A., Menzel, R., & Newcombe, N. S. (2011). Animal navigation: A synthesis. In R. Menzel & J. Fischer (Eds.), Animal thinking: Contemporary issues in comparative cognition (pp. 51–76).  Cambridge, MA, London: MIT Press.

Cheng, K. (2012).  Arthropod navigation: Ants, bees, crabs, spiders finding their way.  In T. R. Zentall and E. A. Wasserman (Eds.), The Oxford handbook of comparative cognition (pp. 347–365).  New York: Oxford University Press.

Cheng K. (2012).  Testing mathematical laws of behavior in the honey bee.  In C. G. Galizia, D. Eisenhardt, & M. Giurfa (Eds.), Honeybee neurobiology and behavior: A tribute to Randolf Menzel (pp. 457–470).  Dordrecht, Heidelberg, London, New York: Springer.

Cheng, K., & Graham, P. (2013).  Spatial memory: Place learning, piloting, and route knowledge.  In D. Waller & L. Nadel (Eds.), Handbook of spatial cognition (pp. 137–153). Washington, DC: American Psychological Association.

Wehner, R., Cheng, K., & Cruse, H. (2014). Visual navigation strategies in insects: lessons from desert ants. In J.S. Werner & L.M. Chalupa (Eds.), New Visual Neurosciences (pp. 1153–1164).  Cambridge MA: MIT Press.

Cheng, K., & Jeffery, K. J.(2017). Spatial cognition. In J. Call (Ed.) APA handbook of comparative psychology, vol. 2: Perception, Learning, and Cognition (pp. 463–483). Washington, DC: American Psychological Association.

Cheng, K., & Crystal, J.D. (2017). Learning to time intervals.  In R. Menzel (Ed.), Learning theory and behavior, 2nd edition (pp. 203–225).  Volume 1 of Learning and memory: A comprehensive reference, 4 volumes (J. Byrne ed.).  Oxford: Elsevier.

Cheng, K. (2018).  Cognitive map.  In J. Vonk & T. Shackleford (Eds.), Encyclopedia of animal cognition and behavior.  Cham: Springer. DOI: https://doi.org/10.1007/978-3-319-47829-6

Refereed Full Conference Papers

Kartiko, I., Kavakli, M., & Cheng, K. (2009).  The impacts of animated-virtual actors’ visual complexity and simulator sickness in virtual reality applications. In B. Werner (Ed.), Proceedings of the 2009 6th International Conference on Computer Graphics, Imaging and Visualization (pp. 147–152). IEEE Computer Society.

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