Contrasting lifetime patterns of territorial success in the nymphalid butterflies Hypolimnas bolina and Melanitis leda: a question of flight physiology?
Darrell J. KempSchool of Life Sciences, Arizona State University, Tempe, AZ 85287-4501, USA; and School of Tropical Biology, James Cook University, Cairns, Qld 4870, Australia. Email: darrell.kemp@jcu.edu.au
Australian Journal of Zoology 53(6) 361-367 https://doi.org/10.1071/ZO05036
Submitted: 6 July 2005 Accepted: 8 November 2005 Published: 6 January 2006
Abstract
Territorial behaviour is widespread among insects and serves as an important constituent of male reproductive success. In butterflies, competition for perching sites is mediated through aerial persistence duels in which the mechanisms of resolution have remained obscure. There is evidence in some species for an effect of age upon contest outcome, which could arise due to individuals adopting lifetime strategies of reproductive risk taking, or as a consequence of lifetime changes in resource-holding potential (RHP). RHP in territorial insects is often equated to the ability for sustained or high-performance flight, and if the morphological and/or energetic determinants of flight performance vary with age then this may explain putative age effects. I addressed this possibility by charting the correlates of residency in two nymphalid species, Hypolimnas bolina L. and Melanitis leda L., at popular perching sites in tropical Australia. Among samples of 55 male H. bolina (20 residents paired to 35 non-residents) and 36 male M. leda (13 residents paired to 23 non-residents), I found moderately sized relationships (effect sizes, d = 0.46–0.79) between residency and the studied biophysical variables (body size, absolute and relative lipid stores, and relative flight musculature). However, differences in age were always much larger (effect sizes, d = 1.23–1.52), with old male H. bolina and young male M. leda favoured for residency. The most parsimonious logistic model of residency in each case proved to be the model containing age only; hence, this study supports the notion of primarily age-based competitive strategies in these two butterflies.
Acknowledgments
I thank Nicole de la Perrelle for assistance with the field component of this investigation, and Andrew Krockenberger for access to equipment.
Alcock, J. (1988). The mating system of three territorial butterflies in Costa Rica. Journal of Research on the Lepidoptera 26, 89–97.
Candolin, U. (1998). Reproduction under predation risk and the trade-off between current and future reproduction in the threespine stickleback. Proceedings of the Royal Society of London. Series B. Biological Sciences 265, 1171–1175.
| Crossref | GoogleScholarGoogle Scholar |
Convey, P. (1989). Influences on the choice between territorial and satellite behavior in male Libellula quadrimaculata Linn. (Odonata: Libellulidae). Behaviour 109, 125–141.
![](../../media/share/OpenURL_Image.gif)
Eickwort, G. C. (1977). Male territorial behaviour in the mason bee Hoplitis anthocopides (Hymenoptera: Megachilidae). Animal Behaviour 25, 542–554.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Hastings, J. M. (1989). The influence of size, age, and residency status on territory defense in male western cicada killer wasps (Sphecius grandis, Hymenoptera: Sphecidae). Journal of the Kansas Entomological Society 62, 363–373.
![](../../media/share/OpenURL_Image.gif)
Hernández, M. I. M. , and Benson, W. W. (1998). Small-male advantage in the territorial butterfly Heliconius sara (Nymphalidae): a paradoxical strategy? Animal Behaviour 56, 533–540.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
![](../../media/share/OpenURL_Image.gif)
Karlsson, B. (1994). Feeding habits and change of body composition with age in three nymphalid butterfly species. Oikos 69, 224–230.
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2000). Contest behavior in territorial male butterflies: does size matter? Behavioral Ecology 11, 591–596.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2001). Age-related site fidelity in the territorial butterfly Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae). Australian Journal of Entomology 40, 65–68.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2002a). Butterfly contests and flight physiology: why do older males fight harder? Behavioral Ecology 13, 456–461.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2002b). Sexual selection constrained by life history in a butterfly. Proceedings of the Royal Society of London. Series B. Biological Sciences 269, 1341–1346.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2002c). Visual mate searching behavior in the evening brown butterfly, Melanitis leda (L.) (Lepidoptera: Nymphalidae). Australian Journal of Entomology 41, 300–305.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. (2003). Twilight fighting in the evening brown butterfly, Melanitis leda (L.) (Nymphalidae): residency and age effects. Behavioral Ecology and Sociobiology 54, 7–13.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. , and Alcock, J. (2003). Lifetime resource utilization, flight physiology and the evolution of contest competition in territorial insects. American Naturalist 162, 290–301.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. , and Rutowski, R. L. (2001). Spatial and temporal patterns of territorial mate locating behaviour in Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae). Journal of Natural History 35, 1399–1411.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. , and Wiklund, C. (2001). Fighting without weaponry: a review of male–male contest competition in butterflies. Behavioral Ecology and Sociobiology 49, 429–442.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Kemp, D. J. , Wiklund, C. , and Van Dyck, H. (in press). Contest behavior in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance. Behavioral Ecology and Sociobiology ,
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Leimar, O. , and Enquist, M. (1984). The effect of asymmetries in owner–intruder interactions. Journal of Theoretical Biology 111, 475–491.
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. (1987). Maximum lift production during takeoff in flying animals. Journal of Experimental Biology 130, 235–258.
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. (1989a). Bodybuilding dragonflies: costs and benefits of maximizing flight muscle. Physiological Zoology 62, 505–521.
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. (1989b). Effects of load-lifting constraints on the mating system of a dance fly. Ecology 70, 496–502.
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. (2000). Variability in the size, composition, and function of insect flight muscles. Annual Review of Physiology 62, 157–178.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. , and Chai, P. (1991). Aerial predation and butterfly design: how palatability, mimicry, and the need for evasive flight constrain mass allocation. American Naturalist 138, 15–36.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. , and Rollins, R. A. (1994). Assessment of energy reserves by damselflies engaged in aerial contests for mating territories. Animal Behaviour 48, 1023–1030.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Marden, J. H. , and Waage, J. K. (1990). Escalated damselfly territorial contests are energetic wars of attrition. Animal Behaviour 39, 954–959.
![](../../media/share/OpenURL_Image.gif)
Petersson, E. (1989). Age-associated male mating success in three swarming caddis fly species (Trichoptera: Leptoceridae). Ecological Entomology 14, 335–340.
![](../../media/share/OpenURL_Image.gif)
Plaistow, S. , and Siva-Jothy, M. T. (1996). Energetic constraints and male mate-securing tactics in the damselfly Calopteryx splendens xanthostoma (Charpentier). Proceedings of the Royal Society of London. Series B. Biological Sciences 263, 1233–1238.
![](../../media/share/OpenURL_Image.gif)
Plaistow, S. , and Tsubaki, Y. (2000). A selective trade-off for territoriality and non-territoriality in the polymorphic damselfly Mnais costalis. Proceedings of the Royal Society of London. Series B. Biological Sciences 267, 969–975.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Poizat, G. , Rosecchi, E. , and Crivelli, A. J. (1999). Empirical evidence of a trade-off between reproductive effort and expectation of future reproduction in female three-spined sticklebacks. Proceedings of the Royal Society of London. Series B. Biological Sciences 266, 1543–1548.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Rutowski, R. L. (1992). Male mate-locating behavior in the common eggfly, Hypolimnas bolina (Nymphalidae). Journal of the Lepidopterist’s Society 46, 24–38.
![](../../media/share/OpenURL_Image.gif)
Sartori, M. , Keller, L. , Thomas, A. G. B. , and Passera, L. (1992). Flight energetics in relation to sexual differences in the mating behavior of a mayfly, Siphlonurus aestivalis. Oecologia 92, 172–176.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Shreeve, T. G. (1987). The mate locating behaviour of the male speckled wood butterfly, Pararge aegeria, and the effect of phenotypic differences in hind-wing spotting. Animal Behaviour 35, 682–690.
![](../../media/share/OpenURL_Image.gif)
Srygley, R. B. (1994). Locomotor mimicry in butterflies? The associations of positions of centres of mass among groups of mimetic, unprofitable prey. Philosophical Transactions of the Royal Society of London (B) 343, 145–155.
![](../../media/share/OpenURL_Image.gif)
Stjernholm, F. , and Karlsson, B. (2000). Nuptial gifts and the use of body resources for reproduction in the green-veined white butterfly Pieris napi. Proceedings of the Royal Society of London. Series B. Biological Sciences 267, 807–811.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Wickman, P.-O. (1985). Territorial defence and mating success in males of the small heath butterfly, Coenonympha pamphilus L. (Lepidoptera: Satyridae). Animal Behaviour 33, 1162–1168.
![](../../media/share/OpenURL_Image.gif)
Wickman, P.-O. (1992). Sexual selection and butterfly design – a comparative study. Evolution 46, 1525–1536.
![](../../media/share/OpenURL_Image.gif)
Williams, G. C. (1957). Pleiotropy, natural-selection, and the evolution of senescence. Evolution; International Journal of Organic Evolution 11, 398–411.
![](../../media/share/OpenURL_Image.gif)
Windig, J. J. , and Nylin, S. (1999). Adaptive wing asymmetry in males of the speckled wood butterfly (Pararge aegeria)? Proceedings of the Royal Society of London. Series B. Biological Sciences 266, 1413–1418.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)
Yuval, B. , Kaspi, R. , Shloush, S. , and Warburg, M. S. (1998). Nutritional reserves regulate male participation in Mediterranean fruit fly leks. Ecological Entomology 23, 211–215.
| Crossref | GoogleScholarGoogle Scholar |
![](../../media/share/OpenURL_Image.gif)