Greater degree of body size plasticity in males than females of the rhinoceros beetle Trypoxylus dichotomus

  • Wataru KojimaEmail author
Original Research Paper


Female body size is more sensitive to the environmental conditions during development than male body size in many insect species with female-biased sexual size dimorphism (SSD) (i.e., females are larger than males). However, the sexual difference in body size plasticity is largely unknown in species with male-biased SSD. Here, I conducted a laboratory experiment using low- and high-quality diets to examine sexual differences in body size plasticity in the male-larger rhinoceros beetle Trypoxylus dichotomus Linnaeus, 1771 (Coleoptera: Scarabaeidae). I found that male body size was much more greatly affected by nutritional status than female body size. The stronger condition dependency in male body size is likely associated with sexual selection for male body size. Furthermore, in wild-caught beetles, males showed larger body size differences between years within populations, while the body size of females was more constant. Within populations and years, male body size also showed greater variation than female body size. In addition, SSD of field-caught beetles was usually much smaller than that of lab-reared beetles of corresponding populations. Thus, the stronger condition dependency of male body size in this species probably has profound effects on the variation of observed body size and SSD among and within populations.


Coleoptera Intrasexual selection Life-history traits Phenotypic plasticity Scarabaeidae Sexual size dimorphism (SSD) 



I would like to thank Dr. Takuma Takanashi, Hiroshi Makihara and Aika Kawachi for their help in collecting beetles and morphological measurement. I am very grateful to the two anonymous reviewers and the associate editor for valuable comments on the manuscript. This study was supported by JSPS Grant-in-Aid for Research Activity Start-up, Grant number 17H06901.


  1. Alcock J (1984) Long-term maintenance of size variation in populations of Centris pallida (Hymenoptera: Anthophoridae). Evolution 38:220–223CrossRefGoogle Scholar
  2. Bonduriansky R (2007) The evolution of condition-dependent sexual dimorphism. Am Nat 169:9–19Google Scholar
  3. Bordalo MD, Vieira HC, Rodrigues AC, Rosa R, Soares AM, Pestana JL (2018) Combined effects of predation risk and food quality on freshwater detritivore insects. Mar Freshw Res 69:74–81CrossRefGoogle Scholar
  4. Bowden JJ, Eskildsen A, Hansen RR, Olsen K, Kurle CM, Høye TT (2015) High-Arctic butterflies become smaller with rising temperatures. Biol Lett 11:20150574CrossRefGoogle Scholar
  5. Chown SL, Gaston KJ (2010) Body size variation in insects: a macroecological perspective. Biol Rev 85:139–169CrossRefGoogle Scholar
  6. DeBlock M, Stoks R (2008) Short-term larval food stress and associated compensatory growth reduce adult immune function in a damselfly. Ecol Entomol 33:796–801Google Scholar
  7. DeBlock M, McPeek MA, Stoks R (2008) Life history plasticity to combined time and biotic constraints in Lestes damselflies from vernal and temporary ponds. Oikos 117:908–916CrossRefGoogle Scholar
  8. Emlen DJ (1997) Diet alters male horn allometry in the beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Proc Roy Soc B 264:567–574CrossRefGoogle Scholar
  9. Emlen DJ, Warren IA, Johns A, Dworkin I, Lavine LC (2012) A mechanism of extreme growth and reliable signaling in sexually selected ornaments and weapons. Science 337:860–864CrossRefGoogle Scholar
  10. Evans EW (2000) Morphology of invasion: body size patterns associated with establishment of Coccinella septempunctata (Coleoptera: Coccinellidae) in western North America. Eur J Entomol 97:469–474CrossRefGoogle Scholar
  11. Fairbairn DJ (2005) Allometry for sexual size dimorphism: testing two hypotheses for Rensch’s rule in the water strider Aquarius remigis. Am Nat 166:69–84Google Scholar
  12. Fernández-Montraveta C, Moya-Laraño J (2007) Sex-specific plasticity of growth and maturation size in a spider: implications for sexual size dimorphism. J Evol Biol 20:1689–1699CrossRefGoogle Scholar
  13. Gotthard K (2000) Increased risk of predation as a cost of high growth rate: an experimental test in a butterfly. J Anim Ecol 69:896–902CrossRefGoogle Scholar
  14. Gotthard K, Nylin S, Wiklund C (1994) Adaptive variation in growth rate: life history costs and consequences in the speckled wood butterfly, Pararge aegeria. Oecologia 99:281–289CrossRefGoogle Scholar
  15. Hirst AG, Horne CR, Atkinson D (2015) Equal temperature–size responses of the sexes are widespread within arthropod species. Proc R Soc B 282:20152475CrossRefGoogle Scholar
  16. Honěk A (1993) Intraspecific variation in body size and fecundity in insects: a general relationship. Oikos 66:483–492CrossRefGoogle Scholar
  17. Hongo Y (2007) Evolution of male dimorphic allometry in a population of the Japanese horned beetle Trypoxylus dichotomus septentrionalis. Behav Ecol Sociobiol 62:245–253CrossRefGoogle Scholar
  18. Johns A, Gotoh H, McCullough EL, Emlen DJ, Lavine LC (2014) Heightened condition-dependent growth of sexually selected weapons in the rhinoceros beetle, Trypoxylus dichotomus (Coleoptera: Scarabaeidae). Integr Comp Biol 54:614–621CrossRefGoogle Scholar
  19. Kari JS, Huey RB (2000) Size and seasonal temperature in free-ranging Drosophila subobscura. J Therm Biol 25:267–272CrossRefGoogle Scholar
  20. Karino K, Seki N, Chiba M (2004) Larval nutritional environment determines adult size in Japanese horned beetles Allomyrina dichotoma. Ecol Res 19:663–668CrossRefGoogle Scholar
  21. Kojima W (2015a) Variation in body size in the giant rhinoceros beetle Trypoxylus dichotomus is mediated by maternal effects on egg size. Ecol Entomol 40:420–427CrossRefGoogle Scholar
  22. Kojima W (2015b) Attraction to carbon dioxide from feeding resources and conspecific neighbours in larvae of the rhinoceros beetle Trypoxylus dichotomus. PLoS One 10:e0141733CrossRefGoogle Scholar
  23. Kojima W, Sugiura S, Makihara H, Ishikawa Y, Takanashi T (2014) Rhinoceros beetles suffer male-biased predation by mammalian and avian predators. Zool Sci 31:109–115CrossRefGoogle Scholar
  24. Lovich JE, Gibbons JW (1992) A review of techniques for quantifying sexual size dimorphism. Growth Dev Aging 56:269–269Google Scholar
  25. Maekawa M (2005) A preliminary study on movement of beetles living on sappy trees in a secondary oak forest in rural landscape, central Japan. Bull Inst Nat Educ Shiga Heights 42:13–16 (in Japanese) Google Scholar
  26. McCullough EL, Emlen DJ (2013) Evaluating the costs of a sexually selected weapon: big horns at a small price. Anim Behav 86:977–985CrossRefGoogle Scholar
  27. Moczek AP (2003) The behavioral ecology of threshold evolution in a polyphenic beetle. Behav Ecol 14:841–854CrossRefGoogle Scholar
  28. Nylin S, Svärd L (1991) Latitudinal patterns in the size of European butterflies. Ecography 14:192–202CrossRefGoogle Scholar
  29. Plaistow SJ, Tsuchida K, Tsubaki Y, Setsuda K (2005) The effect of a seasonal time constraint on development time, body size, condition, and morph determination in the horned beetle Allomyrina dichotoma L. (Coleoptera: Scarabaeidae). Ecol Entomol 30:692–699CrossRefGoogle Scholar
  30. Puniamoorthy N, Schäfer MA, Blanckenhorn WU (2012) Sexual selection accounts for the geographic reversal of sexual size dimorphism in the dung fly, Sepsis punctum (Diptera: Sepsidae). Evolution 66:2117–2126CrossRefGoogle Scholar
  31. Rodrigues D, Moreira GR (2004) Seasonal variation in larval host plants and consequences for Heliconius erato (Lepidoptera: Nymphalidae) adult body size. Austral Ecol 29:437–445CrossRefGoogle Scholar
  32. Rohner PT, Blanckenhorn WU, Puniamoorthy N (2016) Sexual selection on male size drives the evolution of male-biased sexual size dimorphism via the prolongation of male development. Evolution 70:1189–1199CrossRefGoogle Scholar
  33. Rohner PT, Teder T, Esperk T, Lüpold S, Blanckenhorn WU (2018) The evolution of male-biased sexual size dimorphism is associated with increased body size plasticity in males. Funct Ecol 32:581–591CrossRefGoogle Scholar
  34. Rosa E, Saastamoinen M (2017) Sex-dependent effects of larval food stress on adult performance under semi-natural conditions: only a matter of size? Oecologia 184:633–642CrossRefGoogle Scholar
  35. Rowe L, Houle D (1996) The lek paradox and the capture of genetic variance by condition dependent traits. Proc R Soc B 263:1415–1421CrossRefGoogle Scholar
  36. Rust R (2006) Latitudinal variation in the size and developmental parameters of the alkali bee, Nomia melanderi (Hymenoptera: Halictidae). J Kansas Entomol Soc 79:239–249CrossRefGoogle Scholar
  37. Siva-Jothy M (1987) Mate securing tactics and the cost of fighting in the Japanese horned beetle, Allomyrina dichotoma L. (Scarabaeidae). J Ethol 5:165–172CrossRefGoogle Scholar
  38. Smith SM, Nager RG, Costantini D (2016) Meta-analysis indicates that oxidative stress is both a constraint on and a cost of growth. Ecol Evol 6:2833–2842CrossRefGoogle Scholar
  39. Stillwell RC, Morse GE, Fox CW (2007) Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle. Am Nat 170:358–369Google Scholar
  40. Stillwell RC, Blanckenhorn WU, Teder T, Davidowitz G, Fox CW (2010) Sex differences in phenotypic plasticity affect variation in sexual size dimorphism in insects: from physiology to evolution. Annu Rev Entomol 55:227–245CrossRefGoogle Scholar
  41. Stoks R, Block MD, McPeek MA (2006) Physiological costs of compensatory growth in a damselfly. Ecology 87:1566–1574CrossRefGoogle Scholar
  42. Teder T, Tammaru T (2005) Sexual size dimorphism within species increases with body size in insects. Oikos 108:321–334CrossRefGoogle Scholar

Copyright information

© The Japanese Society of Applied Entomology and Zoology 2019

Authors and Affiliations

  1. 1.Graduate School of Sciences and Technology for InnovationYamaguchi UniversityYamaguchiJapan

Personalised recommendations