• Walker, L.A. & Holwell, G.I. (in press) Static allometries do not reflect evolutionary allometry in exaggerated weaponry of male New Zealand sheet-web spiders (Cambridgea spp.) Journal of Evolutionary Biology
  • LeGrice, R. J., & Holwell, G. I. (2022). Size variation, allometry and mating success in Aotearoa| New Zealand kelp flies (Coelopidae). Evolutionary Ecology, 1-18 Link
  • Saunders, T. E., Avila, G. A., & Holwell, G. I. (2022). Retrospective host-specificity testing shows Trissolcus basalis (Wollaston) and the native Trissolcus oenone (Dodd)(Hymenoptera: Scelionidae) have overlapping physiological host ranges in New Zealand. Biological Control, 170, 104926. Link
  • LeGrice, R. J., Ward, D. F., & Holwell, G. I. (2022). Coastal Diptera species and communities and their geographic distribution in Aotearoa| New Zealand. New Zealand Journal of Zoology, 1-20 Link
  • McCambridge, J. E., Painting, C. J., Walker, L. A., & Holwell, G. I. (2022). Contests between male New Zealand sheet-web spiders, Cambridgea plagiata (Araneae: Desidae). New Zealand Journal of Zoology49(1), 53-66 Link


  • Mark, C.J., Painting, C.J., O’Hanlon, J.C. and Holwell, G.I., 2021. Lichen moths do not benefit from ‘element imitation’masquerade in the absence of a matching background. Evolutionary Ecology35(3), pp.401-412. Link
  • Powell, E. C., Willmott, N. J., Selleck, C. J., Painting, C. J., Hickey, A. J., Machado, G., & Holwell, G. I. (2021). No risk to scrambling? Mating tactic does not affect the frequency of leg autotomy in a New Zealand harvestman. Animal Behaviour177, 99-106 Link
  • Pollo, P., Burke, N. W., & Holwell, G. I. (2021). Effects of male and female personality on sexual cannibalism in the Springbok mantis. Animal Behaviour182, 1-7 Link
  • Burke, N. W., & Holwell, G. I. (2021). Increased male mating success in the presence of prey and rivals in a sexually cannibalistic mantis. Behavioral Ecology32(4), 574-579 Link
  • Burke, N. W., & Holwell, G. I. (2021). Male coercion and female injury in a sexually cannibalistic mantis. Biology letters17(1), 20200811. Link
  • Fisher, A. M., Le Page, S., Manser, A., Lewis, D. R., Holwell, G. I., Wigby, S., & Price, T. A. (2021). Relatedness modulates density‐dependent cannibalism rates in Drosophila. Functional Ecology35(12), 2707-2716 Link
  • Powell, E. C., Painting, C. J., Hickey, A. J., Machado, G., & Holwell, G. I. (2021). Diet, predators, and defensive behaviors of New Zealand harvestmen (Opiliones: Neopilionidae). The Journal of Arachnology49(1), 122-140. Link
  • Giribet, G., Sheridan, K., Baker, C. M., Painting, C. J., Holwell, G. I., Sirvid, P. J., & Hormiga, G. (2021). A molecular phylogeny of the circum-Antarctic Opiliones family Neopilionidae. Invertebrate Systematics35(8), 827-849 Link
  • Horrocks, K. J., Avila, G. A., Holwell, G. I., & Suckling, D. M. (2021). Irradiation-induced sterility in an egg parasitoid and possible implications for the use of biological control in insect eradication. Scientific Reports11(1), 1-12 Link
  • Saunders, T. E., Avila, G. A., & Holwell, G. I. (2021). Pre‐emptive host‐specificity testing of Trissolcus japonicus (Ashmead)(Hymenoptera: Scelionidae) reveals high parasitism levels against the endemic New Zealand alpine shield bug in laboratory no‐choice tests. Austral Entomology60(2) Link


  • Fea, M., Boisseau, R. P., Emlen, D. J., & Holwell, G. I. (2020). Cybernetic combatants support the importance of duels in the evolution of extreme weapons. Proceedings of the Royal Society B287(1928), 20200254. Link
  • Powell, E. C., Painting, C. J., Hickey, A. J., & Holwell, G. I. (2020). Defining an intrasexual male weapon polymorphism in a New Zealand harvestman (Opiliones: Neopilionidae) using traditional and geometric morphometrics. Biological Journal of the Linnean Society130(2), 395-409. Link
  • Griffin, M. J., Holwell, G. I., & Symonds, M. R. (2020). From quiet-night-in to party animal: Sex ratio and density affect male/female aggregations in a ‘harem’polygynous insect. Animal Behaviour170, 111-118. Link
  • Bennik, R. M., Hoare, R. J., & Holwell, G. I. (2020). Seasonal variation in body size and male mating success within lichen tuft moths Izatha (Lepidoptera: Xyloryctidae). Austral Entomology. Link
  • Fisher, A. M., Holwell, G. I., & Price, T. A. (2020). Behavioural correlations and aggression in praying mantids. Behavioral Ecology and Sociobiology74, 1-10. Link
  • Fisher, A. M., Cornell, S. J., Holwell, G. I., & Price, T. A. (2020). Mate‐finding Allee effects can be exacerbated or relieved by sexual cannibalism. Journal of Animal Ecology. Link
  • Walker, L. A., Vink, C. J., Holwell, G. I., & Buckley, T. R. (2020). A preliminary molecular phylogeny for New Zealand sheet-web spiders (Cambridgea) and comparison of web-building behaviour. New Zealand Journal of Zoology47(3), 187-205. Link
  • Griffin, M. J., Holwell, G. I., & Symonds, M. R. (2020). Sex ratio and female allocation to harems in a polygynous bark beetle. Austral Entomology59(1), 149-155. Link
  • Horrocks, K. J., Avila, G. A., Holwell, G. I., & Suckling, D. M. (2020). Integrating sterile insect technique with the release of sterile classical biocontrol agents for eradication: is the Kamikaze Wasp Technique feasible?. BioControl, 1-15. Link


  • O’Brien, D.M., Boisseau, R.P., Duell, M., McCullough, E., Powell, E.C., Somjee, U. Solie, S., Hickey, A.J. Holwell, G.I. Painting, C.J. Emlen, D.J. Muscle mass drives cost in sexually selected arthropod weapons. Proc. R. Soc. B 286 (1905), 20191063 Link
  • Le Grice, R. Holwell, G.I. & Painting, C.J. (2019) Directional selection on body size but no apparent survival cost to being large in wild New Zealand giraffe weevils Evolution 73 (4), 762-776 Link
  • McCambridge, J. E., Painting, C. J., Walker, L. A., & Holwell, G. I. (2019). Weapon allometry and phenotypic correlation in the New Zealand sheetweb spider Cambridgea plagiata. Biological Journal of the Linnean Society126(2), 349-359.Link
  • Langton-Myers, S. S., Holwell, G. I. , & Buckley, T. R. (2019) Weak premating isolation between Clitarchus stick insect species despite divergent male and female genital morphology. Journal of Evolutionary Biology 32 (5), 398-411 Link
  • McCambridge, J.E., Walker, L.A. & Holwell, G.I. (2019) Natural history and ecology of the New Zealand sheet-web spiders Cambridgea plagiata and C. foliata (Araneae: Desidae). 53 (19-20), 1153-1167 Link
  • Griffin, M.J., Holwell, G.I. & Symonds, M.R.E. (2019) Insect harem polygyny: when is a harem not a harem? Behavioral Ecology & Sociobiology 73 (4), 40 Link
  • Fea, M. P., Mark, C. J., & Holwell, G. I. (2019). Sexually dimorphic antennal structures of New Zealand Cave Wētā (Orthoptera: Rhaphidophoridae). New Zealand Journal of Zoology, 1-25. Link
  • Seldon, D. S., & Holwell, G. I. (2019). Monteremita gen. nov.(Coleoptera: Carabidae), a new genus of ground beetle from New Caledonia. Austral Entomology58(4), 724-728. Link


  • Fea, M., & Holwell, G. I. (2018). Exaggerated male legs increase mating success by reducing disturbance to females in the cave wētā Pachyrhamma waitomoensis. Proc. R. Soc. B, 285(1880), 20180401. Link
  • Walker, L. A., & Holwell, G. I. (2018). The role of exaggerated male chelicerae in male–male contests in New Zealand sheet-web spiders. Animal Behaviour, 139, 29-36. Link
  • Fea, M., & Holwell, G. (2018). Combat in a cave-dwelling wētā (Orthoptera: Rhaphidophoridae) with exaggerated weaponry. Animal Behaviour, 138, 85-92. Link
  • Toth, C. A., Santure, A. W., Holwell, G. I., Pattemore, D. E., & Parsons, S. (2018). Courtship behaviour and display-site sharing appears conditional on body size in a lekking bat. Animal Behaviour, 136, 13-19. Link
  • Fisher, A. M., Cornell, S. J., Holwell, G. I., & Price, T. A. (2018). Sexual cannibalism and population viability. Ecology and Evolution, 8(13), 6663-6670. Link
  • Walker, L. A., & Holwell, G. I. (2018). Bad tenants: female sheet-web spiders, Cambridgea foliata (Araneae: Desidae), lose feeding opportunities when cohabiting with males. The Journal of Arachnology, 46(3), 391-398. Link
  • Mark, C. J., Parsons, S., & Holwell, G. I. (2018). Antennal morphology and micro‐sensory architecture of the New Zealand magpie moth, Nyctemera annulata (Lepidoptera: Erebidae): diversity, distribution and dimorphism. Austral Entomology, 57(3), 303-323. Link


  • ME Herberstein, CJ Painting & GI Holwell. (2017) Scramble competition polygyny in terrestrial arthropods.  Advances in the Study of Behavior 49, 237-295 Link
  • SS Myers, GI Holwell, & TR Buckley (2017) Biogeography underpins alternative consequences of secondary contact in the New Zealand stick insect genus Clitarchus. Journal of Biogeography 44 (9) 2069–2081 Link
  • CJ Painting, SS Myers, GI Holwell, & TR Buckley (2017) Phylogeography of the New Zealand giraffe weevil Lasiorhynchus barbicornis (Coleoptera: Brentidae): A comparison of biogeographic boundaries Biological Journal of the Linnean Society 122 (1) Pages 13–28 Link 
  • GA Avila, TM Withers & GI Holwell (2017). Courtship and mating behaviour in the parasitoid wasp Cotesia urabae (Hymenoptera: Braconidae): mate location and the influence of competition and body size on male mating success. Bulletin of Entomological Research, 107 (4) 439-447 Link
  • S Lamont, D Seldon, C Vink & GI Holwell (2017). Spider diversity and composition in native broadleaf-podocarp forest fragments of northern Hawke’s Bay, New Zealand. New Zealand Journal of Zoology. 44 (2) 129-143  Link
  • C Mark, S Parsons & GI Holwell (online). Antennal morphology and micro-sensory architecture of the New Zealand magpie moth, Nyctemera annulata (Arctiinae, Erebidae): diversity, distribution, and dimorphism. Austral Entomology Link


  • SS Myers, TR Buckley GI Holwell (2016). Male genital claspers influence female mate acceptance in the stick insect Clitarchus hookeri. Behavioral Ecology and Sociobiology, 70(9), 1547-1556. Link
  • GI Holwell, PJD Allen, F Goudie, PE Duckett, CJ Painting (2016) Male density influences mate searching speed and copulation duration in millipedes (Polydesmida: Gigantowales chisholmi). Behavioral Ecology and Sociobiology, 1-8. Link
  • RM Bennik, TR Buckley, RJB Hoare, GI Holwell (2016) Molecular phylogeny reveals the repeated evolution of complex male genital traits in the New Zealand moth genus Izatha (Lepidoptera: Xyloryctidae). Systematic Entomology 41 (2), 309-322. Link
  • Avila, GA, TM Withers & GI Holwell (2016). Retrospective risk assessment reveals likelihood of potential non-target attack and parasitism by Cotesia urabae (Hymenoptera: Braconidae): A comparison between laboratory and field-cage testing results. Biological Control, 103, 108-118. Link
  • M Marinov, C Amaya-Perilla, GI Holwell, A Varsani, K Van Bysterveldt, S Kraberger, D Stainton, A Dayaram, N Curtis, RH Cruickshank, A Paterson (2016) Geometric morphometrics and molecular systematics of Xanthocnemis sobrina (McLachlan, 1873)(Odonata: Coenagrionidae) and comparison to its congeners. Zootaxa 4078 (1), 84-120. Link
  • GA Avila, TM Withers, GI Holwell (2016) Olfactory cues used in host‐habitat location and host location by the parasitoid Cotesia urabae. Entomologia Experimentalis et Applicata 158 (2), 202-209. Link
  • GA Avila, TM Withers, GI Holwell (2016) Laboratory odour-specificity testing of Cotesia urabae to assess potential risks to non-target species. BioControl, 1-13. Link


  • CJ Painting, AF Probert, DJ Townsend, GI Holwell (2015) Multiple exaggerated weapon morphs: a novel form of male polymorphism in harvestmen. Scientific Reports 5. Link
  • LA Walker, GI Holwell (2016) Sexual cannibalism in a facultative parthenogen: the springbok mantis (Miomantis caffra). Behavioral Ecology, 27, 851 – 856.  Link
  • JC O’Hanlon, ME Herberstein, GI Holwell (2015) Habitat selection in a deceptive predator: maximizing resource availability and signal efficacy. Behavioral Ecology 26 (1), 194-199. Link
  • MC Saavedra, GA Avila, TM Withers, GI Holwell (2015) The potential global distribution of the Bronze bug Thaumastocoris peregrinus Carpintero and Dellapé (Hemiptera: Thaumastocoridae). Agricultural and Forest Entomology 17 (4), 375-388. Link
  • CJ Painting, GI Holwell (2015) Temporal variation in body size and weapon allometry in the New Zealand giraffe weevil. Ecological Entomology 40 (4), 486-489. Link
  • GA Avila, TM Withers, GI Holwell (2015) Host testing of the parasitoid Cotesia urabae (Austin & Allen, 1989)(Hymenoptera: Braconidae) to assess the risk posed to the New Zealand nolid moth Celama parvitis (Howes, 1917)(Lepidoptera: Nolidae): do host deprivation and experience influence acceptance of non‐target hosts? Austral Entomology 54 (3), 270-277. Link
  • SS Myers, TR Buckley, GI Holwell (2015) Mate detection and seasonal variation in stick insect mating behaviour (Phamatodea: Clitarchus hookeri). Behaviour 152 (10), 1325-1348. Link
  • GI Holwell, O Kazakova, F Evans, JC O’Hanlon, KL Barry (2015) The Functional Significance of Chiral Genitalia: Patterns of Asymmetry, Functional Morphology and Mating Success in the Praying Mantis Ciulfina baldersoni. PloS One 10 (6), e0128755. Link
  • MC Saavedra, TM Withers, GI Holwell (2015) Susceptibility of four Eucalyptus host species for the development of Thaumastocoris peregrinus Carpintero and Dellapé (Hemiptera: Thaumastocoridae). Forest Ecology and Management 336, 210-216. Link


  • CJ Painting, TR Buckley, GI Holwell (2014) Weapon allometry varies with latitude in the New Zealand giraffe weevil. Journal of Evolutionary Biology 27 (12), 2864-2870. Link
  • CJ Painting, GI Holwell (2014) Flexible alternative mating tactics by New Zealand giraffe weevils. Behavioral Ecology 25 (6), 1409-1416. Link
  • JC O’Hanlon, GI Holwell, ME Herberstein (2014) Pollinator deception in the orchid mantis. The American Naturalist 183 (1), 126-132. Link
  • CJ Painting, TR Buckley, GI Holwell (2014) Male‐biased sexual size dimorphism and sex ratio in the New Zealand Giraffe Weevil, Lasiorhynchus barbicornis (Fabricius, 1775)(Coleoptera: Brentidae). Austral Entomology 53 (3), 317-327. Link
  • CJ Painting, GI Holwell (2014) Exaggerated rostra as weapons and the competitive assessment strategy of male giraffe weevils. Behavioral Ecology 25 (5), 1223-1232. Link
  • GI Holwell (2014) Four new species of Ciulfina Giglio-Tos, 1915 (Mantodea: Liturgusidae, Liturgusinae) from the Northern Territory, Australia. Zootaxa 3797 (1), 029-038. Link
  • CJ Painting, GI Holwell (2014) Observations on the ecology and behaviour of the New Zealand giraffe weevil (Lasiorhynchus barbicornis). New Zealand Journal of Zoology 41 (2), 147-153. Link
  • LE Allen, KL Barry, GI Holwell (2014) Different paths to sexual size dimorphism in two praying mantids, Pseudomantis albofimbriata and Hierodula majuscula. Insect science 21 (2), 227-233. Link
  • JC O’Hanlon, GI Holwell, ME Herberstein (2014) Predatory pollinator deception: Does the orchid mantis resemble a model species? Current Zoology 60 (1), 90-103. Link
  • PJ Lester, SDJ Brown, ED Edwards, GI Holwell, SM Pawson, DF Ward, CH Watts (2014) Critical issues facing New Zealand entomology. New Zealand Entomologist 37 (1), 1-13. Link
  • MP Fea, MC Stanley, GI Holwell (2014) Cannibalistic siblicide in praying mantis nymphs (Miomantis caffra). Journal of Ethology 32 (1), 43-51. Link


  • MP Fea, MC Stanley, GI Holwell (2013) Fatal attraction: sexually cannibalistic invaders attract naive native mantids. Biology Letters 9 (6), 20130746. Link
  • CJ Painting, GI Holwell (2013) Exaggerated trait allometry, compensation and trade-offs in the New Zealand giraffe weevil (Lasiorhynchus barbicornis). PLoS One 8 (11), e82467. Link
  • GA Avila, LA Berndt, GI Holwell (2013) Dispersal behavior of the parasitic wasp Cotesia urabae (Hymenoptera: Braconidae): A recently introduced biocontrol agent for the control of Uraba lugens (Lepidoptera: Nolidae) in New Zealand. Biological Control 66 (3), 166-172. Link
  • GA Avila, LA Berndt, GI Holwell (2013) First releases and monitoring of the biological control agent Cotesia urabae Austin and Allen (Hymenoptera: Braconidae). New Zealand Entomologist 36 (2), 65-72. Link
  • KDL Umbers, NJ Tatarnic, GI Holwell, ME Herberstein (2013) Bright turquoise as an intraspecific signal in the chameleon grasshopper (Kosciuscola tristis). Behavioral Ecology and Sociobiology 67 (3), 439-447. Link


  • KDL Umbers, NJ Tatarnic, GI Holwell, ME Herberstein (2012) Ferocious fighting between male grasshoppers. PLoS One 7 (11), e49600. Link
  • LE Allen, KL Barry, GI Holwell (2012) Mate location and antennal morphology in the praying mantid Hierodula majuscula. Australian Journal of Entomology 51 (2), 133-140. Link


  • JP Evans, C Gasparini, GI Holwell, IW Ramnarine, TE Pitcher, A Pilastro (2011) Intraspecific evidence from guppies for correlated patterns of male and female genital trait diversification. Proceedings of the Royal Society of London B: Biological Sciences 278, 2611-2620 Link
  • LE Allen, KL Barry, GI Holwell, ME Herberstein (2011)  Perceived risk of sperm competition affects juvenile development and ejaculate expenditure in male praying mantids. Animal Behaviour 82 (5), 1201-1206. Link
  • KDL Umbers, GI Holwell, AJ Stow, ME Herberstein (2011) Molecular evidence for variation in polyandry among praying mantids (Mantodea: Ciulfina). Journal of Zoology 284 (1), 40-45. Link
  • KL Barry, GI Holwell, ME Herberstein (2011) A paternity advantage for speedy males? Sperm precedence patterns and female re-mating frequencies in a sexually cannibalistic praying mantid. Evolutionary Ecology 25 (1), 107-119. Link
  • JC O’Hanlon, GI Holwell (2011) The influence of abrupt forest edges on praying mantid populations. Insect Conservation and Diversity 4 (2), 107-114. Link


  • GI Holwell, C Winnick, T Tregenza, ME Herberstein (2010) Genital shape correlates with sperm transfer success in the praying mantis Ciulfina klassi (Insecta: Mantodea). Behavioral Ecology and Sociobiology 64 (4), 617-625. Link
  • GI Holwell, ME Herberstein (2010) Chirally dimorphic male genitalia in praying mantids (Ciulfina: Liturgusidae). Journal of Morphology 271 (10), 1176-1184. Link
  • KL Barry, GI Holwell, ME Herberstein (2010) Multimodal mate assessment by male praying mantids in a sexually cannibalistic mating system. Animal Behaviour 79 (5), 1165-1172. Link


  • CRM Attard, GI Holwell, TS Schwartz, KDL Umbers, A Stow, ME Herberstein, LB Beheregaray (2009) Microsatellite markers for the praying mantid Ciulfina rentzi (Liturgusidae). Molecular Ecology Resources 9 (6), 1480-1482. Link
  • KL Barry, GI Holwell, ME Herberstein (2009) Male mating behaviour reduces the risk of sexual cannibalism in an Australian praying mantid. Journal of Ethology 27 (3), 377-383. Link
  • CG Winnick, GI Holwell, ME Herberstein (2009) Internal reproductive anatomy of the praying mantid Ciulfina klassi (Mantodea: Liturgusidae). Arthropod Structure & Development 38 (1), 60-69. Link
  • J O’Hanlon, GI Holwell (2009) Patterns of praying mantid population density following cyclone disturbance in far north Queensland, Australia. Australian Journal of Zoology 57 (2), 85-88. Link


  • GI Holwell (2008) Geographic variation in genital morphology of Ciulfina praying mantids. Journal of Zoology 276 (1), 108-114. Link
  • KL Barry, GI Holwell, ME Herberstein (2008) Female praying mantids use sexual cannibalism as a foraging strategy to increase fecundity. Behavioral Ecology 19 (4), 710-715. Link
  • GI Holwell (2008) Male control of copulation duration in Ciulfina praying mantids (Insecta Mantodea). Ethology Ecology & Evolution 20 (1), 61-70. Link

2007 and earlier

  • GI Holwell, SG Ginn, ME Herberstein (2007) Three new species of Ciulfina Giglio-Tos (Mantodea: Liturgusidae) from north-eastern Australia. Zootaxa 1583, 23-35. Link
  • GI Holwell, KL Barry, ME Herberstein (2007) Mate location, antennal morphology, and ecology in two praying mantids (Insecta: Mantodea). Biological Journal of the Linnean Society 91 (2), 307-313. Link
  • GI Holwell (2007) Spermatophore feeding and mating behaviour in praying mantids (Mantodea: Liturgusidae). Journal of Zoology 271 (3), 255-260. Link
  • PJB Hill, GI Holwell, A Göth, ME Herberstein (2004) Preference for habitats with low structural complexity in the praying mantid Ciulfina sp.(Mantidae). Acta Oecologica 26 (1), 1-7. Link
  • BV Griffiths, GI Holwell, ME Herberstein, MA Elgar (2003) Frequency, composition and variation in external food stores constructed by orb-web spiders: Nephila edulis and Nephila plumipes (Araneae: Araneoidea). Australian Journal of Zoology 51 (2), 119-128. Link


A Stow, N MacLean, GI Holwell (2014)  Austral Ark – The State of Wildlife in Australia & New Zealand. Cambridge University Press. Buy It