Behavioral Ecology and Sociobiology

, Volume 56, Issue 1, pp 34–41 | Cite as

Male-male interference competition decreases spawning rate in the European bitterling (Rhodeus sericeus)

  • Martin Reichard
  • Pavel Jurajda
  • Carl Smith
Original Article


We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.


Alternative mating tactics Density dependence Operational sex ratio Sperm competition Reproduction 



We are grateful to Tamsin Burland, Ulrika Candolin, and Lars Chittka for their comments. This research was supported by a Royal Society (NATO/Royal Society fellowship awarded to M.R. and Royal Society Joint Project Grant awarded to C.S. and P.J.), and Czech Academy of Sciences Internal Grant awarded to M.R. M.R. holds a license for conducting experimental work on vertebrates in accordance with Czech legal requirements, and experiments complied with current Czech laws.


  1. Addicot JF, Aho JM, Antolin MF, Padilla DK, Richardson JS, Soluk DK (1987) Ecological neighbourhoods: scaling environmental patterns. Oikos 49:340–346Google Scholar
  2. Alonso-Pimentel H, Papaj DR (1996) Operational sex ratio versus gender density as determinant of copulation duration in the walnut fly Rhagoletis juglandis (Diptera: Tephritidae). Behav Ecol Sociobiol 39:171–180Google Scholar
  3. Alonzo SH, Warner RR (2000) Allocation to mate guarding or increased sperm production in a Mediterranean wrasse. Am Nat 156:266–275CrossRefGoogle Scholar
  4. Anholt BR (1997) How should we test for the role of behaviour in population dynamics? Evol Ecol 11:633–640CrossRefGoogle Scholar
  5. Birkhead TR, Møller AP (1998) Sperm competition and sexual selection. Academic, LondonGoogle Scholar
  6. Butchart SHM, Seddon N, Ekstrom JMM (1999) Polyandry and competition for territories in bronze-winged jacanas. J Anim Ecol 68:928–939CrossRefGoogle Scholar
  7. Candolin U, Reynolds JD (2002) Adjustments of ejaculation rates in response to risk of sperm competition in a fish the bitterling (Rhodeus sericeus). Proc R Soc Lond B 269:1549–1553CrossRefPubMedGoogle Scholar
  8. Clutton-Brock TH, Rose KE, Guinness FE (1997) Density-related changes in sexual selection in red deer. Proc R Soc Lond B 264:1509–1516CrossRefPubMedGoogle Scholar
  9. Ekman J, Eggers S, Griesser M, Tegelstrom H (2001) Queuing for preferred territories: delayed dispersal of Siberian jays. J Anim Ecol 70:317–324CrossRefGoogle Scholar
  10. Emlen ST (1976) Lek organization and mating strategies in the bullfrog. Behav Ecol Sociobiol 1:283–313Google Scholar
  11. Enders MM (1993) The effect of male size and operational sex ratio on male mating success in the common spider mite Tetranychus urticae Koch (Acari, Tetranychidae). Anim Behav 46:835–846CrossRefGoogle Scholar
  12. Gross MR (1996) Alternative mating strategies and tactics: diversity within sexes. Trends Ecol Evol 11:92–98CrossRefGoogle Scholar
  13. Hoenig JM, Heisey DM (2001) The abuse of power: the pervasive fallacy of power calculations for power analysis. Am Stat 55:19–24CrossRefGoogle Scholar
  14. Jennions MD, Møller AP (2003) A survey of the statistical power of research in behavioral ecology and animal behavior. Behav Ecol 14:438–445CrossRefGoogle Scholar
  15. Jirotkul M (1999) Population density influences male-male competition in guppies. Anim Behav 58:1169–1175PubMedGoogle Scholar
  16. Kanoh Y (1996) Pre-oviposition ejaculation in externally fertilizing fish: how sneaker male rose bitterlings contrive to mate. Ethology 102:883–899Google Scholar
  17. Kvarnemo C, Ahnesjö I (2002) Operational sex ratios and mating competition. In: Hardy ICW (ed) Sex ratios: concepts and research methods. Cambridge University Press, Cambridge, pp 366–382Google Scholar
  18. Le Boef BJ, Peterson RS (1969) Social status and mating activity in elephant seals. Science 163:91–93Google Scholar
  19. McNamara JM (2001) The effect of adaptive behaviour on the stability of population dynamics. Ann Zool Fenn 38:25–36Google Scholar
  20. Meiri I, Gothilf Y, Zohar Y, Elizur A (2002) Physiological changes in the spawning gilthead bream Sparus aurata succeeding the removal of males. J Exp Zool 292:555–564CrossRefPubMedGoogle Scholar
  21. Mills SC, Reynolds JD (2003) Operational sex ratio and alternative reproductive behaviours in the European bitterling, Rhodeus sericeus. Behav Ecol Sociobiol 54:98–104Google Scholar
  22. Mysterud A, Coulson T, Stenseth NC (2002) The role of males in the dynamics of ungulate populations. J Anim Ecol 71:907–915CrossRefGoogle Scholar
  23. Nagata Y (1985) The experimental approaches for analyzing the causes of spawning efficiency of the bitterling, Rhodeus ocellatus (Kner). Mem Osaka Kyoiku Univ 34:81–101Google Scholar
  24. Nilsson SG (1987) Limitation and regulation of population density in the nuthatch Sitta europea (Aves) breeding in natural cavities. J Anim Ecol 56:921–937Google Scholar
  25. Oliveira RF, Almada VC, Forsgren E, Goncalves EJ (1999) Temporal variation in male traits nesting aggregations and mating success in the peacock blenny. J Fish Biol 54:499–512CrossRefGoogle Scholar
  26. Parker GA (1970) Sperm competition and its evolutionary consequences in the insects. Biol Rev 45:525–567Google Scholar
  27. Parker GA, Ball MA, Stockley P, Gage MJG (1996) Sperm competition games: individual assessment of sperm competition intensity by group spawners. Proc R Soc Lond B 263:1291–1297Google Scholar
  28. Poizat G, Ponton D (1996) Multi-scale approach to species-habitat relationships: juvenile fish in a large river section. Freshwater Biol 36 611–622Google Scholar
  29. Przybylski M (1996) The diel feeding pattern of bitterling Rhodeus sericeus amarus (Bloch) in the Wieprz-Krzna canal Poland. Pol Arch Hydrobiol 43:203–212Google Scholar
  30. Ray C, Hastings A (1996) Density dependence: are we searching at the wrong spatial scale? J Anim Ecol 65:556–566Google Scholar
  31. Reichard M, Jurajda P, Ondračková M (2002) Interannual variability in seasonal dynamics and species composition of drifting young-of-the-year fishes in two European lowland rivers. J Fish Biol 60:87–101CrossRefGoogle Scholar
  32. Reichard M, Smith C, Jordan WC (2004) Genetic evidence reveals density-dependent mediated success of alternative mating tactics in the European bitterling (Rhodeus sericeus). Mol Ecol (in press)Google Scholar
  33. Rose SM (1959) Population control in guppies. Am Midl Nat 62:474–481Google Scholar
  34. Smith C, Reynolds JD, Sutherland WJ (2000a) The population consequences of reproductive decisions. Proc R Soc Lond B 267:1327–1334CrossRefPubMedGoogle Scholar
  35. Smith C, Reynolds JD, Sutherland WJ, Jurajda P (2000b) Adaptive host choice and avoidance of superparasitism in the spawning decisions of bitterling (Rhodeus sericeus). Behav Ecol Sociobiol 48:29–35CrossRefGoogle Scholar
  36. Smith C, Douglas A, Jurajda P (2002) Sexual conflict, sexual selection and sperm competition in the spawning decisions of bitterling (Rhodeus sericeus). Behav Ecol Sociobiol 51:433–439CrossRefGoogle Scholar
  37. Smith C, Reichard M, Jurajda P (2003) Assessment of sperm competition by bitterling (Rhodeus sericeus). Behav Ecol Sociobiol 53:206–213CrossRefGoogle Scholar
  38. Smith C, Reichard M, Jurajda P, Przybylski M (2004) The reproductive ecology of the European bitterling (Rhodeus sericeus). J Zool 262:107–124CrossRefGoogle Scholar
  39. Spencer M, Blaustein L, Cohen JE (2002) Oviposition habitat selection by mosquitoes (Culiseta longiareolata) and consequences for population size. Ecology 83:669–679Google Scholar
  40. Sutherland WJ (1996) From individual behaviour to population ecology. Oxford University Press, OxfordGoogle Scholar
  41. Taborsky M (1994) Sneakers satellites and helpers: parasitic and cooperative behaviour in fish reproduction. Adv Stud Behav 23:1–100Google Scholar
  42. Tyler CR, Sumpter JP (1996) Oocyte growth and development in teleosts. Rev Fish Biol Fish 6:287–318Google Scholar
  43. Tyler CR, Sumpter JP, Witthames PR (1990) The dynamics of oocytes growth during vitellogenesis in the rainbow trout Salmo gairdneri. Biol Reprod 43:202–209PubMedGoogle Scholar
  44. Village A (1983) The role of nest site availability and territorial behavior in limiting the breeding density of kestrels. J Anim Ecol 52:635–645Google Scholar
  45. Warner RR, Hoffman SG (1980) Population density and the economics of territorial defence in a coral reef fish. Ecology 61:772–780Google Scholar
  46. Wiens JA, Rotenberry JT, Vanhorne B (1987) Habitat occupancy patterns of North-American shrubsteppe birds—the effects of spatial scale. Oikos 48:132–147Google Scholar
  47. Wiepkema PR (1961) An ethological analysis of the reproductive behaviour of the bitterling (Rhodeus amarus Bloch). Arch Neerl Zool 14:103–199Google Scholar
  48. Wootton RJ (1998) Ecology of teleost fishes. Kluwer, DordrechtGoogle Scholar

Copyright information

© Springer-Verlag 2004

Authors and Affiliations

  1. 1.School of Biological Sciences, Queen MaryUniversity of LondonLondon UK
  2. 2.Department of BiologyUniversity of LeicesterLeicesterUK
  3. 3.Institute of Vertebrate BiologyAcademy of Sciences of the Czech RepublicBrnoCzech Republic
  4. 4.Institute of Vertebrate BiologyAcademy of Sciences of the Czech RepublicBrnoCzech Republic

Personalised recommendations