Prof Pat Backwell

Professor
Research School of Biology

I received my formal education in South Africa. During my graduate years I worked on the breeding behaviour of several species of frogs. After my PhD I switched to studying fiddler crabs and completed three post-docs. I then did a year of lecturing in South Africa before moving to Panama to work for the Smithsonian for the next six years, still studying fiddlers. I took up my present post as lecturer at ANU at the start of 2002. All of this experience has forged me into the truly brilliant scientist that I am.

My current research is entirely field-based and I have spent many hours sitting on sunny beaches staring at amazing little crabs. Most of my fieldwork has been carried out in Mozambique, Japan, South Africa and Central America. I am now working on tropical Australian fiddlers. My main study site is in Darwin, NT.

Research interests

I work on the behavioural ecology of fiddler crabs. There are over 100 species of fiddlers world wide, and about 20 of them occur in Australia. They are often the most commonly seen inter-tidal animal on tropical mudflats, and are ecologically important as a major food source for shore birds. Although a lot is known about them as prey items, surprisingly little is known about their behaviour. They are ideal study animals because they occur in huge numbers and are exceptionally easy to catch, mark, observe and manipulate. They are sexually dimorphic, males having a single enlarged claw which is used both as a weapon and waved to attract mate-searching females.

  • Darnell, M, Backwell, P & Munguia, P 2020, 'Frequency and latency of autotomy of a sexually selected fiddler crab appendage', Journal of Experimental Marine Biology and Ecology, vol. 523, pp. 4pp.
  • Perez, D & Backwell, P 2020, 'The functions of multiple visual signals in a fiddler crab', Ethology, vol. 126, no. 4, pp. 455-462.
  • Gruber, J, Kahn, A & Backwell, P 2019, 'Risks and rewards: balancing costs and benefits of predator avoidance in a fiddler crab', Animal Behaviour, vol. 158, pp. 9-13.
  • Young, C, Cain, K, Svedin, N et al 2019, 'Breeding biology of Crimson Finches (Neochmia phaeton) in the eastern Kimberley, Western Australia', Emu - Austral Ornithology, vol. 119, no. 2, pp. 106-115.
  • Backwell, P 2019, 'Synchronous waving in fiddler crabs: a review', Current Zoology, vol. 65, no. 1, pp. 83-88.
  • Perez, D & Backwell, P 2019, 'Selection for conspicuous visual signals in a fiddler crab', Behavioral Ecology and Sociobiology, vol. 73, no. 61, pp. 1-8.
  • Darnell, M, Backwell, P, Stockbridge, J et al 2019, 'Thermal and desiccation constraints drive territory preference in fiddler crabs', Journal of Experimental Marine Biology and Ecology, vol. 518, no. 151173, pp. 1-6.
  • Perez, D & Backwell, P 2019, 'Male spacing and female choice in a fiddler crab', Behavioral Ecology, vol. 30, no. 6, pp. 1769-1774.
  • Chou, C, Head, M & Backwell, P 2019, 'Effects of temperature on reproductive timing and hatching success in a tropical fiddler crab', Biological Journal of the Linnean Society, vol. 128, no. 4, pp. 817-827.
  • Chou, C, Perez, D, Johns, S et al 2019, 'Staying cool: the importance of shade availability for tropical ectotherms', Behavioral Ecology and Sociobiology, vol. 73, no. 8, pp. 12pp.
  • Mowles, S, Jennions, M & Backwell, P 2018, 'Robotic crabs reveal that female fiddler crabs are sensitive to changes in male display rate', Biology Letters, vol. 14, no. 1, pp. 1-4.
  • Heatwole, S, Christy, J & Backwell, P 2018, 'Taking a risk: how far will male fiddler crabs go?', Behavioral Ecology and Sociobiology, vol. 72, no. 82, pp. 1-6pp.
  • Sanches, F, Costa, T, Barreto, R et al 2018, 'The cost of living in mixed species populations: A fiddler crab example', Journal of Experimental Marine Biology and Ecology, vol. 500, pp. 30-33.
  • Bourdiol, J, Chou, C, Perez, D et al 2018, 'Investigating the role of a mud structure in a fiddler crab: do semidomes have a reproductive function?', Behavioral Ecology and Sociobiology, vol. 72, no. 141, pp. 7pp.
  • Perez, D & Backwell, P 2017, 'Female preferences for conspecific and heterospecific wave patterns in a fiddler crab', Journal of Experimental Marine Biology and Ecology, vol. 486, pp. 155-159.
  • Munguia, P, Backwell, P & Darnell, M 2017, 'Thermal constraints on microhabitat selection and mating opportunities', Animal Behaviour, vol. 123, pp. 259-265.
  • Sanches, F, Costa, T, Barreto, R et al 2017, 'Faster male displays and less complex choice are more attractive to female fiddler crabs as they reduce search costs', Animal Behaviour, vol. 124, no. 1, pp. 119-123pp.
  • Clark, H & Backwell, P 2017, 'Territorial battles between fiddler crab species', Royal Society Open Science, vol. 4, no. 1, pp. 160621-160621.
  • Young, C, Cain, K, Svedin, N et al 2017, 'Predictors of aggressive response towards simulated intruders depend on context and sex in Crimson Finches (Neochmia phaeton)', Behavioural Processes, vol. 138, pp. 41-48pp.
  • Mowles, S, Jennions, M & Backwell, P 2017, 'Multimodal communication in courting fiddler crabs reveals male performance capacities', Royal Society Open Science, vol. 4, no. 3, pp. 161093-161093.
  • Vega Trejo, R & Backwell, P 2017, 'Testing female preferences under more natural conditions: a case study on a fiddler crab', Behavioral Ecology and Sociobiology, vol. 71, no. 5, pp. 1-7.
  • Young, C, Cain, K, Svedin, N et al 2017, 'Nesting Success in Crimson Finches: Chance or Choice?', Ethology, vol. 123, no. 1, pp. 41-50.
  • Clark, H & Backwell, P 2016, 'Male mating success in a fiddler crab: a lesson in sample sizes', Journal of Ethology, vol. 34, no. 2, pp. 119-126.
  • Clark, H & Backwell, P 2016, 'Assortative mating in a fiddler crab', Behaviour, vol. 153, no. 2, pp. 175-185.
  • Young, C, Cain, K, Svedin, N et al 2016, 'The role of pigment based plumage traits in resolving conflicts', Journal of Avian Biology, vol. 47, no. 2, pp. 167-175.
  • Hayes, C, Callander, S, Booksmythe, I et al 2016, 'Mate choice and the operational sex ratio: An experimental test with robotic crabs', Journal of Evolutionary Biology, vol. 29, no. 7, pp. 1455-1461.
  • Painting, C, Splinter, W, Callander, S et al 2016, 'Ladies First: Coerced Mating in a Fiddler Crab', PLOS ONE (Public Library of Science), vol. 11, no. 6, pp. e0155707-e0155707.
  • Backwell, P & Chou, C 2016, 'Why do ovigerous females approach courting males? Female preferences and sensory biases in a fiddler crab', Ecology and Evolution, vol. 6, no. 15, pp. 5473-5478.
  • Perez, D, Christy, J & Backwell, P 2016, 'Choosing a mate in a high predation environment: Female preference in the fiddler crab Uca terpsichores', Ecology and Evolution, vol. 6, no. 20, pp. 7443-7450.
  • Clark, H & Backwell, P 2016, 'Micro-climate and incubation in a fiddler crab species', Journal of Experimental Marine Biology and Ecology, vol. 485, pp. 18-23.
  • Peso, M, Curran, E & Backwell, P 2016, 'Not what it looks like: mate-searching behaviour, mate preferences and clutch production in wandering and territory-holding female fiddler crabs', Royal Society Open Science, vol. 3, no. 8, pp. 160339-160339.
  • Dyson, M & Backwell, P 2016, 'Alternative mating tactics and male mating success in two species of fiddler crab', Behaviour, vol. 153, no. 12, pp. 1403-1418.
  • Perez, D, Heatwole, S, Morrell, L et al 2015, 'Handedness in fiddler crab fights', Animal Behaviour, vol. 110, pp. 99-104pp.
  • Backwell, P & Clark, H 2015, 'Temporal and spatial variation in female mating preferences in a fiddler crab', Behavioral Ecology and Sociobiology, vol. 69, no. 11, pp. 1779-1784.
  • Peso, M, Telford, L & Backwell, P 2014, 'Comparison shopping: detectability and mate preferences in a fiddler crab', Animal Behaviour, vol. 88, pp. 107-111.
  • Holman, L, Kahn, A & Backwell, P 2014, 'Fiddlers on the roof: Elevation muddles mate choice in fiddler crabs', Behavioral Ecology, vol. 25, no. 2, pp. 271-275.
  • Binning, S, Barnes, J, Davies, J et al 2014, 'Ectoparasites modify escape behaviour, but not performance, in a coral reef fish', Animal Behaviour, vol. 93, pp. 1-7.
  • Beponis, L, O'Dea, R, Ohl, V et al 2014, 'Cleaning Up After a Meal: The Consequences of Prey Disposal for Pit-Building Antlion Larvae', Ethology, vol. 120, no. 9, pp. 873-880.
  • Kahn, A, Holman, L & Backwell, P 2014, 'Female preferences for timing in a fiddler crab with synchronous courtship waving displays', Animal Behaviour, vol. 98, pp. 35-39.
  • Gardner, J.L., Amano, T., Backwell, P.R.Y., Ikin, K., Sutherland, W.J., Peters, A., 2014. Temporal patterns of avian body size reflect linear size responses to broadscale environmental change over the last 50 years. Journal of Avian Biology 45, 529–535. Editor’s Choice.
  • Callander, S, Kahn, A, Hunt, J et al 2013, 'The effect of competitors on calling effort and life span in male field crickets', Behavioral Ecology, vol. 24, no. 5, pp. 1251-1259.
  • Booksmythe, I, Backwell, P & Jennions, M 2013, 'Competitor size, male mating success and mate choice in eastern mosquitofish, Gambusia holbrooki', Animal Behaviour, vol. 85, no. 2, pp. 371-375.
  • Hayes, C, Booksmythe, I, Jennions, M et al 2013, 'Does male reproductive effort increase with age? Courtship in fiddler crabs', Biology Letters, vol. 9, no. 2, pp. 20121078-20121078.
  • Callander, S, Hayes, C, Jennions, M et al 2013, 'Experimental evidence that immediate neighbors affect male attractiveness', Behavioral Ecology, vol. 24, no. 3, pp. 730-733.
  • Johnson, L, Mantle, B, Gardner, J et al 2013, 'Morphometric measurements of dragonfly wings: The accuracy of pinned, scanned and detached measurement methods', ZooKeys, vol. 276, no. 276, pp. 77-84.
  • Bolton, J, Backwell, P & Jennions, M 2013, 'Density dependence and fighting in species with indeterminate growth: A test in a fiddler crab', Animal Behaviour, vol. 85, no. 6, pp. 1367-1376.
  • Callander, S, Kahn, A, Maricic, T et al 2013, 'Weapons or mating signals? Claw shape and mate choice in a fiddler crab', Behavioral Ecology and Sociobiology, vol. 67, no. 7, pp. 1163-1167.
  • Kahn, A, Jennions, M, Backwell, P et al 2013, 'Strategic male courtship effort varies in concert with adaptive shifts in female mating preferences', Behavioural Ecology, vol. 24, no. 4, pp. 906-913.
  • Slatyer, R, Jennions, M & Backwell, P 2012, 'Polyandry occurs because females initially trade sex for protection', Animal Behaviour, vol. 83, no. 5, pp. 1203-1206.
  • Callander, S, Backwell, P & Jennions, M 2012, 'Context-dependent male mate choice: the effects of competitor presence and competitor size', Behavioral Ecology, vol. 23, no. 2, pp. 355-360.
  • Callander, S, Jennions, M & Backwell, P 2012, 'The effect of claw size and wave rate on female choice in a fiddler crab', Journal of Ethology, vol. 30, no. 1, pp. 151-155.
  • Reaney, L, Maurer, G, Backwell, P et al 2012, 'Paternity analysis of two male mating tactics in the fiddler crab, Uca mjoebergi', Behavioral Ecology and Sociobiology, vol. 66, no. 7, pp. 1017-1024.
  • Callander, S, Bolton, J, Jennions, M et al 2012, 'A farewell to arms: males with regenerated claws fight harder over resources', Animal Behaviour, vol. 84, no. 3, pp. 619-622.
  • Osborne, L, Umbers, K, Backwell, P et al 2012, 'Male tawny dragons use throat patterns to recognize rivals', Naturwissenschaften, vol. 99, no. 10, pp. 869-872.
  • Slatyer, R, Mautz, B, Backwell, P et al 2012, 'Estimating genetic benefits of polyandry from experimental studies: A meta-analysis', Biological Reviews, vol. 87, no. 1, pp. 1-33.
  • Booksmythe, I, Hayes, C, Jennions, M et al 2012, 'The effects of neighbor familiarity and size on cooperative defense of fiddler crab territories', Behavioral Ecology, vol. 23, no. 2, pp. 285-289.
  • Milner, R, Jennions, M & Backwell, P 2012, 'Keeping up appearances: male fiddler crabs wave faster in a crowd', Biology Letters, vol. 8, no. 2, pp. 176-178.
  • Bolton, J, Callander, S, Jennions, M et al 2011, 'Even Weak Males Help Their Neighbours: Defence Coalitions in a Fiddler Crab', Ethology, vol. 117, no. 11, pp. 1027-1030.
  • Milner, R, Jennions, M & Backwell, P 2011, 'Non-independent mate choice in a fiddler crab: a case of stimulus enhancement', Behavioral Ecology and Sociobiology, vol. 65, no. 7, pp. 1419-1424.
  • Booksmythe, I, Jennions, M & Backwell, P 2011, 'Male fiddler crabs prefer conspecific females during simultaneous, but not sequential, mate choice', Animal Behaviour, vol. 81, no. 4, pp. 775-778.
  • Mokhlesi, A, Kamrani, E, Backwell, P et al 2011, 'Study on the behaviour of two fiddler crabs, Uca sindensis and Uca annulipes (Decapoda: Ocypodidae), in Bandar Abbas, Iran', Journal of the Marine Biological Association of the United Kingdom, vol. 91, no. 1, pp. 245-249.
  • Mautz, B, Detto, T, Wong, B et al 2011, 'Male fiddler crabs defend multiple burrows to attract additional females', Behavioral Ecology, vol. 22, no. 2, pp. 261-267.
  • Callander, S; Jennions MD and Backwell, P 2011, 'Female choice over short and long distances: neighbour effects', Behavioral Ecology and Sociobiology, vol 65, pp. 2071-2078.
  • Milner, R, Jennions, M & Backwell, P 2011, 'Know thine enemy's neighbor: neighbor size
    affects floaters' choice of whom to fight', Behavioral Ecology, vol. 22, pp. 947-950.
  • Backwell, P & Jennions, M 2004, 'Animal behaviour: coalition among male fiddler crabs.', Nature, vol. 430, no. 6998, pp. 417-417.
  • Backwell, P, Christy, J, Telford, S et al 2000, 'Dishonest signalling in a fiddler crab', Proceedings of the Royal Society of London Series B: Biological Sciences, vol. 267, no. 1444, pp. 719-724.
  • Backwell, P, Jennions, M, Christy, J et al 1999, 'Female choice in the synchronously waving fiddler crab Uca annulipes', Ethology, vol. 105, no. 5, pp. 415-421.
  • Backwell, P, O'Hara, P & Christy, J 1998, 'Prey availability and selective foraging in shorebirds', Animal Behaviour, vol. 55, no. 6, pp. 1659-1667.
  • Jennions, M, Backwell, P & Backwell, P 1998, 'Variation in courtship rate in the fiddler crab Uca annulipes: Is it related to male attractiveness?', Behavioral Ecology, vol. 9, no. 6, pp. 605-611.
  • Booksmythe, I, Jennions, M & Backwell, P 2010, 'Interspecific assistance: fiddler crabs help heterospecific neighbours in territory defence', Biology Letters, vol. 6, no. 6, pp. 748-750.
  • Milner, R, Booksmythe, I, Jennions, M et al 2010, 'The battle of the sexes? Territory acquisition and defence in male and female fiddler crabs', Animal Behaviour, vol. 79, no. 3, pp. 735-738.
  • Milner, R, Jennions, M & Backwell, P 2010, 'Safe sex: male-female coalitions and pre-copulatory mate-guarding in a fiddler crab', Biology Letters, vol. 6, no. 2, pp. 180-182.
  • Booksmythe, I, Milner, R, Jennions, M et al 2010, 'How do weaponless male fiddler crabs avoid aggression?', Behavioral Ecology and Sociobiology, vol. 64, no. 3, pp. 485-491.
  • Milner, R, Detto, T, Jennions, M et al 2010, 'Experimental evidence for a seasonal shift in the strength of a female mating preference', Behavioral Ecology, vol. 21, no. 2, pp. 311-316.
  • Milner, R, Detto, T, Jennions, M et al 2010, 'Hunting and predation in a fiddler crab', Journal of Ethology, vol. 28, no. 1, pp. 171-173.
  • Booksmythe, I, Jennions, M & Backwell, P 2010, 'Investigating the 'dear enemy' phenomenon in the territory defence of the fiddler crab, Uca mjoebergi', Animal Behaviour, vol. 79, no. 2, pp. 419-423.
  • Detto, T, Jennions, M & Backwell, P 2010, 'When and Why Do Territorial Coalitions Occur? Experimental Evidence from a Fiddler Crab', The American Naturalist, vol. 175, no. 5, pp. E119-E125.
  • Milner, R, Jennions, M & Backwell, P 2010, 'Eavesdropping in crabs: an agency for lady detection', Biology Letters, vol. 6, no. 6, pp. 755-757.
  • Detto, T & Backwell, P 2009, 'The fiddler crab Uca mjoebergi uses ultraviolet cues in mate choice but not aggressive interactions', Animal Behaviour, vol. 78, pp. 407-411.
  • Detto, T & Backwell, P 2009, 'Social monogamy in a fiddler crab, Uca capricornis', Journal of Crustacean Biology, vol. 29, no. 3, pp. 283-289.
  • Kinnear, M, Smith, L, Maurer, G et al 2009, 'Polymorphic microsatellite loci for paternity analysis in the fiddler crab Uca mjoebergi', Journal of Crustacean Biology, vol. 29, no. 2, pp. 273-274.
  • Murai, M, Backwell, P & Jennions, M 2009, 'The cost of reliable signaling: experimental evidence for predictable variation among males in a cost-benefit trade-off between sexually selected traits', Evolution, vol. 63, no. 9, pp. 2363-2371.
  • Lailvaux, S, Reaney, L & Backwell, P 2009, 'Dishonest signalling of fighting ability and multiple performance traits in the fiddler crab Uca mjoebergi', Functional Ecology, vol. 23, pp. 359-366.
  • Slatyer, R, Fok, E, Hocking, R et al 2008, 'Why do fiddler crabs build chimneys?', Biology Letters, vol. 4, pp. 616-618.
  • Detto, T, Hemmi, J & Backwell, P 2008, 'Colouration and Colour Changes of the Fiddler Crab, Uca capricornis: A descriptive study', PLoS ONE (Public Library of Science), vol. 3, no. 2, pp. 1-10.
  • Fayed, S, Jennions, M & Backwell, P 2008, 'What factors contribute to an ownership advantage?', Biology Letters, vol. 4, pp. 143-145.
  • Reaney, L, Sims, R, Sims, S et al 2008, 'Experiments with robots explain synchronized courtship in fiddler crabs', Current Biology, vol. 18, no. 2, pp. R62-R63.
  • Reaney, L, Milner, R, Detto, T et al 2008, 'The effects of claw regeneration on territory ownership and mating success in the fiddler crab Uca mjoebergi', Animal Behaviour, vol. 75, pp. 1473-1478.
  • Milner, R, Jennions, M & Backwell, P 2008, 'Does the environmental context of a signalling male influence his attractiveness?', Animal Behaviour, vol. 76, pp. 1565-1570.
  • Booksmythe, I, Detto, T & Backwell, P 2008, 'Female fiddler crabs settle for less: the travel costs of mate choice', Animal Behaviour, vol. 76, pp. 1775-1781.
  • Reaney, L & Backwell, P 2007, 'Risk-taking behavior predicts aggression and mating success in a fiddler crab', Behavioral Ecology, vol. 18, pp. 521-525.
  • Backwell, P, Matsumasa, M, Double, M et al 2007, 'What are the consequences of being left-clawed in a predominantly right-clawed fiddler crab?', Proceedings of the Royal Society of London Series B: Biological Sciences, vol. 274, pp. 2723-2729.
  • Reading, K & Backwell, P 2007, 'Can beggars be choosers? Male mate choice in a fiddler crab', Animal Behaviour, vol. 74, pp. 867-872.
  • Reaney, L & Backwell, P 2007, 'Temporal constraints and female preference for burrow width in the fiddler crab, Uca mjoebergi', Behavioral Ecology and Sociobiology, vol. 61, pp. 1515-1521.
  • Murai, M & Backwell, P 2006, 'A conspicuous courtship signal in the fiddler crab Uca perplexa: female choice based on display structure', Behavioral Ecology and Sociobiology, vol. 60, pp. 736-741.
  • Backwell, P, Jennions, M, Wada, K et al 2006, 'Synchronous waving in two species of fiddler crabs', Acta Ethologica, vol. 9, pp. 22-25.
  • Zeil, J, Hemmi, J & Backwell, P 2006, 'Fiddler crabs', Current Biology, vol. 16, no. 2, pp. R40-R41.
  • Detto, T, Backwell, P, Hemmi, J et al 2006, 'Visually mediated species and neighbour recognition in fiddler crabs (Uca mjoebergi and Uca capricornis)', Proceedings of the Royal Society of London Series B: Biological Sciences, vol. 273, pp. 1661-1666.
  • Christy, J & Backwell, P 2006, 'No preference for exaggerated courtship signals in a sensory trap', Animal Behaviour, vol. 71, pp. 1239-1246.
  • Morrell, L, Backwell, P & Metcalfe, N 2005, 'Fighting in fiddler crabs Uca mjoebergi: what determines duration?', Animal Behaviour, vol. 70, pp. 653-662.
  • Murai, M & Backwell, P 2005, 'More signalling for earlier mating: conspicuous male claw waving in the fiddler crab, Uca perplexa', Animal Behaviour, vol. 70, pp. 1093-1097.
  • Backwell, P & Jennions, M 2004, 'Coalition among male fiddler crabs', Nature, vol. 430, p. 417.
  • Backwell, P 2004, 'Cheats Prosper', in Livio Dobrez, Jan Lloyd Jones, Patricia Dobrez (ed.), An ABC of Lying: Taking Stock in Interesting Times, Australian Scholarly Publishing Pty Ltd, Melbourne, Australia, pp. 227-232.
  • Christy, J, Backwell, P & Schober, U 2003, 'Interspecific attractiveness of structures built by courting male fiddler crabs: experimental evidence of a sensory trap', Behavioral Ecology and Sociobiology, vol. 53, pp. 84-91.
  • deRivera, C, Backwell, P, Christy, J et al 2003, 'Density affects female and male mate searching in the fiddler crab, Uca beebei', Behavioral Ecology and Sociobiology, vol. 53, pp. 72-83.
  • Jennions, M, Backwell, P, Murai, M et al 2003, 'Hiding behaviour in fiddler crabs: how long should prey hide in response to a potential predator?', Animal Behaviour, vol. 66, pp. 251-257.
  • Christy, J, Baum, J & Backwell, P 2003, 'Attractiveness of sand hoods built by courting male fiddler crabs, Uca musica: test of a sensory trap hypothesis', Animal Behaviour, vol. 66, pp. 89-94.
  • Christy, J, Backwell, P, Goshima, S et al 2002, 'Sexual selection for structure building by courting male fiddler crabs: an experimental study of behavioral mechanisms', Behavioral Ecology, vol. 13, no. 3, pp. 366-374.
  • Koga, T, Backwell, P, Christy, J et al 2001, 'Male-biased predation of a fiddler crab', Animal Behaviour, vol. 62, pp. 201-207.
  • Christy, J, Backwell, P & Goshima, S 2001, 'The design and production of a sexual signal: hoods and hood building by male fiddler crabs Uca musica', Behaviour, vol. 138, pp. 1065-1083.