Flexible timing of reproductive effort as an alternative mating tactic in black grouse (Lyrurus tetrix) males

in Behaviour
Restricted Access
Get Access to Full Text
Rent on DeepDyve

Have an Access Token?

Enter your access token to activate and access content online.

Please login and go to your personal user account to enter your access token.


Have Institutional Access?

Access content through your institution. Any other coaching guidance?


Alternative reproductive tactics often take the form of dichotomous behavioural phenotypes. Focusing attention on such obvious dichotomy means that flexible patterns of behaviour within tactics is largely ignored. Using a long-term dataset of black grouse (Lyrurus tetrix) lek behaviours, we tested whether there were fine-scale differences in reproductive effort (lek attendance, fighting rates) and whether these were related to age and phenotype. Yearling males increased their lek attendance and fighting rate to a peak when adult male effort was declining. Adults and yearlings allocated reproductive effort according to their body mass but this was unrelated to differences in timing of effort. In adult males, different patterns of lek attendance were associated with different costs of reproduction, measured by mass loss or gain. Overall, our work demonstrates that individuals can use flexible patterns of reproductive effort both in terms of their own condition, their age and the likely costs of behaviours.

Flexible timing of reproductive effort as an alternative mating tactic in black grouse (Lyrurus tetrix) males

in Behaviour



AlataloR.V.HöglundJ.LundbergA. (1991). Lekking in the black grouse — a test of male viability. — Nature 352: 155-156.

AlonsoJ.C.MagañaM.PalacínC.MartínC.A. (2010). Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort. — Behav. Ecol. Sociobiol. 64: 1589-1600.

AlonzoS.H.SinervoB. (2001). Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana. — Behav. Ecol. Sociobiol. 49: 176-186.

BachmanG.WidemoF. (1999). Relationships between body composition, body size and alternative reproductive tactics in a lekking sandpiper, the ruff (Philomachus pugnax). — Funct. Ecol. 13: 411-416.

BrockmannH.J. (2001). The evolution of alternative strategies and tactics. — Adv. Stud. Behav. 30: 1-51.

Clutton-BrockT.H.AlbonS.D.GibsonR.M.GuinnessF.E. (1979). The logical stag: adaptive aspects of fighting in red deer (Cervus elaphus L.). — Anim. Behav. 27: 211-225.

ColtmanD.W.Festa-BianchetM.JorgensonJ.T.StrobeckC. (2001). Age-dependent sexual selection in bighorn rams. — Proc. Roy. Soc. Lond. B: Biol. Sci. 269: 165-172.

CowlesS.A.GibsonR.M. (2014). Displaying to females may lower male foraging time and vigilance in a lekking bird. — Auk 132: 82-91.

DeutschC.J.HaleyM.P.Le BoeufB. (1990). Reproductive effort of male northern elephant seals: estimates from mass loss. — Can. J. Zool. 68: 2580-2593.

DeutschC.J.CrockerD.E.CostaD.P.Le BoeufB.J. (1994). Sex- and age-related variation in reproductive effort of northern elephant seals. — In: Elephant seals: population ecology behavior and physiology ( Le BeoufB.J.LawsR.M. eds). University of California PressBerkeley, CA p.  169-210.

EggertC.GuyétantR. (2003). Reproductive behaviour of spadefoot toads (Pelobates fuscus): daily sex ratios and males’ tactics, ages, and physical condition. — Can. J. Zool. 81: 46-51.

FitzpatrickS. (1998). Birds’ tails as signaling devices: markings, shape, length, and feather quality. — Am. Nat. 151: 157-173.

ForsythA.MontgomerieR.D. (1987). Alternative reproductive tactics in the territorial damselfly Calopteryx maculata: sneaking by older males. — Behav. Ecol. Sociobiol. 21: 73-81.

GrossM.R. (1996). Alternative reproductive strategies and tactics: diversity within sexes. — Trends Ecol. Evol. 11: 92-98.

HämäläinenA.AlataloR.V.LebigreC.SiitariH.SoulsburyC.D. (2012). Fighting behaviour as a correlate of male mating success in black grouse Tetrao tetrix. — Behav. Ecol. Sociobiol. 66: 1577-1586.

HaywardA.D.MooradJ.ReganC.E.BerenosC.PilkingtonJ.G.PembertonJ.M.NusseyD.H. (2015). Asynchrony of senescence among phenotypic traits in a wild mammal population. — Exp. Gerontol. 71: 56-68.

HelminenM. (1963). Composition of the Finnish populations of capercaillie, Tetrao urogallus, and black grouse, Lyrulus tetrix, in the autumns of 1952–1961, as revealed by a study of wings. — Riistatiet Julk 8: 142-149.

HewsD.K.KnappR.MooreM.C. (1994). Early exposure to androgens affects adult expression of alternative male types in tree lizards. — Horm. Behav. 28: 96-115.

HoggJ.T. (1984). Mating in bighorn sheep: multiple creative male strategies. — Science 225: 526-529.

HöglundJ.AlataloR.V.LundbergA. (1992). The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix). — Behav. Ecol. Sociobiol. 30: 71-76.

HöglundJ.AlataloR.V.LundbergA.RättiO. (1994). Context-dependent effects of tail-ornament damage on mating success in black grouse. — Behav. Ecol. 5: 182-187.

HöglundJ.JohanssonT.PelabonC. (1997). Behaviourally mediated sexual selection: characteristics of successful male black grouse. — Anim. Behav. 54: 255-264.

HoviM.AlataloR.V.HöglundJ.LundbergA.RintamäkiP.T. (1994). Lek centre attracts black grouse females. — Proc. Roy. Soc. Lond. B: Biol. Sci. 258: 303-305.

JenningsD.J.CarlinC.M.HaydenT.J.GammellM.P. (2010). Investment in fighting in relation to body condition, age and dominance rank in the male fallow deer, Dama dama. — Anim. Behav. 79: 1293-1300.

JohnL.K. (1993). Alternative reproductive tactics in male eastern gray squirrels: “making the best of a bad job”. — Behav. Ecol. 4: 165-171.

KervinenM.AlataloR.V.LebigreC.SiitariH.SoulsburyC.D. (2012). Determinants of yearling male lekking effort and mating success in black grouse (Tetrao tetrix). — Behav. Ecol 23: 1209-1217.

KervinenM.LebigreC.AlataloR.V.SiitariH.SoulsburyC.D. (2015). Life-history differences in age-dependent expressions of multiple ornaments and behaviors in a lekking bird. — Am. Nat. 185: 13-27.

KervinenM.LebigreC.SoulsburyC.D. (2016). Simultaneous age-dependent and age-independent sexual selection in the lekking black grouse (Lyrurus tetrix). — J. Anim. Ecol. 85: 715-725.

KokkoH.LindströmJ.AlataloR.V.RintamäkiP.T. (1998). Queuing for territory positions in the lekking black grouse (Tetrao tetrix). — Behav. Ecol. 9: 376-383.

KuznetsovaA.BrockhoffP.B.ChristensenR.H.B. (2014). Package ‘lmerTest’: tests for random and fixed effects for linear mixed effect models (lmer objects of lme4 package). — R package version 2.0-11. R Foundation for Statistical Computing Vienna.

LankD.B.SmithC.M.HanotteO.BurkeT.CookeF. (1995). Genetic polymorphism for alternative mating behaviour in lekking male ruff Philomachus pugnax. — Nature 378: 59-62.

LearyC.J.FoxD.J.ShepardD.B.GarciaA.M. (2005). Body size, age, growth and alternative mating tactics in toads: satellite males are smaller but not younger than calling males. — Anim. Behav. 70: 663-671.

LebigreC.AlataloR.V.SiitariH. (2013). Physiological costs enforce the honesty of lek display in the black grouse (Tetrao tetrix). — Oecology 172: 983-993.

LidgardD.C.BonessD.J.BowenW.D.McMillanJ.I. (2005). State-dependent male mating tactics in the grey seal: the importance of body size. — Behav. Ecol. 16: 541-549.

LudwigG.X.AlataloR.V.HelleP.LindénH.LindströmJ.SiitariH. (2006). Short- and long-term population dynamical consequences of asymmetric climate change in black grouse. — Proc. Roy. Soc. Lond. B: Biol. Sci. 273: 2009-2016.

MasonT.H.E.StephensP.A.WillisS.G.ChirichellaR.ApollonioM.RichardsS.A. (2012). Intraseasonal variation in reproductive effort: young males finish last. — Am. Nat. 180: 823-830.

McElligottA.G.NaultyF.ClarkeW.V.HaydenT.J. (2003). The somatic cost of reproduction: what determines reproductive effort in prime-aged fallow bucks?Evol. Ecol. Res. 5: 1239-1250.

MoczekA.P.EmlenD.J. (1999). Proximate determination of male horn dimorphism in the beetle Ontophagus taurus (Coleoptera: Scarabaeidae). — J. Evol. Biol. 12: 27-37.

MysterudA.BonenfantC.LoeL.E.LangvatnR.YoccozN.G.StensethN.C. (2008). The timing of male reproductive effort relative to female ovulation in a capital breeder. — J. Anim. Ecol. 77: 469-477.

OliveiraR.F.TaborskyM.BrockmannH.J. (2008). Alternative reproductive tactics: an integrative approach. — Cambridge University PressCambridge.

PiankaE.R.ParkerW.S. (1975). Age-specific reproductive tactics. — Am. Nat. 109: 453-464.

PitcherB.J.BrieferE.F.VannoniE.McElligottA.G. (2014). Fallow bucks attend to vocal cues of motivation and fatigue. — Behav. Ecol. 25: 392-401.

R Core Team (2013). R: a language and environment for statistical computing. — R Foundation for Statistical ComputingVienna.

SaundersF.C.McElligottA.G.SafiK.HaydenT.J. (2005). Mating tactics of male feral goats (Capra hircus): risks and benefits. — Acta Ethol. 8: 103-110.

SavalliU.M. (1994). Tail length affects territory ownership in the yellow-shouldered widowbird. — Anim. Behav. 48: 105-111.

SiitariH.AlataloR.V.HalmeP.BuchananK.L.KilpimaaJ. (2007). Color signals in the black grouse (Tetrao tetrix): signal properties and their condition dependency. — Am. Nat. 169: 81-92.

SinervoB.LivelyC.M. (1996). The rock-paper-scissors game and the evolution of alternative male strategies. — Nature 380: 240-243.

TaborskyM.OliveiraR.F.BrockmannH.J. (2008). The evolution of alternative reproductive tactics: concepts and questions. — In: Alternative reproductive tactics: an integrative approach ( OliveiraR.F.TaborskyM.BrockmannH.J. eds). Cambridge University PressCambridge p.  1-21.

TennenhouseE.M.WeladjiR.B.HolandØ.NieminenM. (2012). Timing of reproductive effort differs between young and old dominant male reindeer. — Ann. Zool. Fennici 49: 152-160.

VannoniE.McElligottA.G. (2009). Fallow bucks get hoarse: vocal fatigue as a possible signal to conspecifics. — Anim. Behav. 78: 3-10.

VehrencampS.L.BradburyJ.W.GibsonR.M. (1989). The energetic cost of display in male sage grouse. — Anim. Behav. 38: 885-896.

YuvalB.KaspiR.ShloushS.WarburgM.S. (1998). Nutritional reserves regulate male participation in Mediterranean fruit fly leks. — Ecol. Entomol. 23: 211-215.


  • View in gallery

    Mean ± SE individual daily lek attendance (a) and daily fighting rates (b) of males across the lekking season. Filled squares represent adult males and open squares yearling males. Days have been scaled so that on day 0, ⩾50% of all copulations have occurred.

  • View in gallery

    Surface plot showing the individual daily lek attendance in relation to the day and body mass of (a) adult males and (c) yearling males and the individual daily fighting rate in relation to the day and body mass (b) adult males and (d) yearling males. Days have been scaled so that on day 0, ⩾50% of all copulations have occurred. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

  • View in gallery

    Surface plot showing the interaction between individual daily lek attendance, day and the mass loss between pre- and post-lekking for adult males (N=15 males, 148 observations). Mass loss is presented so that negative values indicate large mass loss and positive values indicate mass gain. Days have been scaled so that on day 0, ⩾50% of all copulations have occurred. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.


Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 23 23 11
Full Text Views 91 91 59
PDF Downloads 5 5 1
EPUB Downloads 0 0 0