Mate detection and seasonal variation in stick insect mating behaviour (Phamatodea: Clitarchus hookeri)

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?


For animals that exhibit a scramble competition mating system, sexual selection pressures on mate searching ability are expected to be strong. Scramble competition mating systems evolve when populations provide females with equal accessibility to all male competitors, yet sex ratio and population density influences mating systems and varies seasonally. The stick insect species, Clitarchus hookeri, is frequently found in copula, yet very little is known about it’s mating behaviour. We preformed behavioural tests and assayed antennal sensory morphology to determine whether males used chemosensory cues to detect females. Through natural field observations we found populations to be significantly male-biased earlier in the season, while later, populations began to display equal sex ratios. With increasing female availability mating pair proportions steadily increased, while copulation duration declined. These results support C. hookeri as a scramble competitor, and demonstrate males may alter their behaviour in response to the seasonal variation in female density.

Mate detection and seasonal variation in stick insect mating behaviour (Phamatodea: Clitarchus hookeri)

in Behaviour



AbleD.J. (1999). Scramble competition selects for greater tail fin size in male red-spotted newts (Amphibia: Salamandridae). — Behav. Ecol. Sociobiol. 46: 423-428.

AlcockJ. (1994). Postinsemination associations between males and females in insects: the mate-guarding hypothesis. — Annu. Rev. Entomol. 39: 1-21.

AllenL.E.BarryK.L.HolwellG.I. (2012). Mate location and antennal morphology in the praying mantid Hierodula majuscula. — Aust. J. Entomol. 51: 133-140.

AllenL.E.BarryK.L.HolwellG.I.HerbersteinM.E. (2011). Perceived risk of sperm competition affects juvenile development and ejaculate expenditure in male praying mantids. — Anim. Behav. 82: 1201-1206.

Alonso-PimentelH.PapajD.R. (1996). Operational sex ratio versus gender density as determinants of copulation duration in the walnut fly, Rhagoletis juglandis (Diptera: Tephritidae). — Behav. Ecol. Sociobiol. 39: 171-180.

AnderssonM. (1994). Sexual selection. — Princeton University PressPrinceton, NJ.

BarryK.L.HolwellG.I.HerbersteinM.E. (2008). Female praying mantids use sexual cannibalism as a foraging strategy to increase fecundity. — Behav. Ecol. 19: 710-715.

BarryK.L.HolwellG.I.HerbersteinM.E. (2010a). A paternity advantage for speedy males? Sperm precedence patterns and female re-mating frequencies in a sexually cannibalistic praying mantid. — J. Evol. Ecol. 25: 107-119.

BarryK.L.HolwellG.I.HerbersteinM.E. (2010b). Multimodal mate assessment by male praying mantids in a sexually cannibalistic mating system. — Anim. Behav. 79: 1165-1172.

Berger-TalR.LubinY. (2011). High male mate search costs and a female biased sex ratio shape the male mating strategy in a desert spider. — Anim. Behav. 82: 853-859.

BondurianskyR. (2001). The evolution of male mate choice in insects: a synthesis of ideas and evidence. — Bio. Rev. 76: 305-339.

BondurianskyR.BrooksR.J. (1998). Copulation and oviposition behaviour of Protopiophila litigata (Diptera: Piophilidae). — Can. Entomol. 130: 399-405.

BradlerS. (2009). Phylogeny of the stick and leaf insects (Insecta: Phasmatodea). — Spec. Phylogenet. Evol. 2: 3-139.

BuckleyT.R.AttanayakeD.NylandarJ.A.A.BradlerS. (2010a). The phylogenetic placement and biogeographical origins of the New Zealand stick insects (Phasmatodea). — Syst. Entomol. 35: 207-225.

BuckleyT.R.MarskeK.AttanayakeD. (2010b). Phylogeography and ecological niche modelling of the New Zealand stick insect Clitarchus hookeri (White) support survival in multiple coastal refugia. — J. Biogeogr. 37: 682-695.

BuckleyT.R.MyersS.S.BradlerS. (2014). Revision of the stick insect genus Clitarchus Stål (Phasmatodea: Phasmatidae): new synonymies and two new species from northern New Zealand. — Zootaxa 3900: 451-482.

Danielson-FrançoisA.A.FettererC.A.SmallwoodP.D.JournalS. (2002). American arachnological society body condition and mate choice in Tetragnatha elongata (Araneae, Tetragnathidae). — J. Arachnol. 30: 20-30.

DelisleJ.McNeilJ.N. (1987). Calling behaviour and pheromone titre of the true armyworm Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae) under different temperature and photoperiodic conditions. — J. Insect Physiol. 33: 315-324.

DewsburyD.A. (1982). Ejaculate cost and male choice. — Am. Nat. 119: 601-610.

EmlenS.T.OringL.W. (1997). Ecology, sexual selection, and the evolution of mating systems. — Science 197: 215-223.

EngqvistL.ReinholdK. (2005). Pitfalls in experiments testing predictions from sperm competition theory. — J. Evol. Biol. 18: 116-123.

FlyF.TryoniD.FletcherS. (1969). The structure and function of the sex pheromone glands of the male queensland fruit fly, Dacus tryoni. — J. Insect Behav. 15: 1309-1322.

GunningG.E. (1987). Behavioral observations of the walking stick, Anisomorpha buprestoides (Phasmatodea: Phasmatidae). — Fla. Entomol. 70: 406-408.

GwynneD.T. (1991). Sexual competition among females: what causes courtship-role reversal?Trends Ecol. Evol. 6: 4-7.

HawJ.M.CloutM.N.PowleslandR.G. (2001). Diet of moreporks (Ninox novaeseelandiae) in Pureora Forest determined from prey remains in regurgitated pellets. — N.Z. J. Ecol. 25: 61-67.

HigginsP.J. (ed.) (1996). Handbook of Australian New Zealand and Antarctic birdsVol. 4. — Oxford University PressMelbourne, VIC.

HoeflerC. (2007). Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus. — Anim. Behav. 73: 943-954.

HolwellG.I.BarryK.L.HerbersteinM.E. (2007). Mate location, antennal morphology, and ecology in two praying mantids (Insecta: Mantodea). — Bio. J. Linn. Soc. 91: 307-313.

KellyC.D. (2014a). Male-biased sex ratios and plasticity in post-insemination behaviour in the New Zealand stick insect Micrarchus hystriculeus. — Behaviour 152: 653-666.

KellyC.D. (2014b). Sexual selection, phenotypic variation, and allometry in genitalic and non-genitalic traits in the sexually size-dimorphic stick insect Micrarchus hystriculeus. — Biol. J. Linn. Soc. 113: 471-484.

KellyC.D.BussièreL.F.GwynneD.T. (2008). Sexual selection for male mobility in a giant insect with female-biased size dimorphism. — Am. Nat. 172: 417-423.

KellyC.D.BussiereL.F.GwynneD.T. (2010). Pairing and insemination patterns in a giant weta (Deinacrida rugosa: Orthoptera; Anostostomatidae). — J. Ethol. 28: 483-489.

KokkoH.RankinD.J. (2006). Lonely hearts or sex in the city? Density-dependent effects in mating systems. — Phil. Trans. Roy. Soc. Lond. B: Biol. Sci. 361: 319-334.

KokkoH.WongB. (2007). What determines sex roles in mate searching?Evolution 61: 1162-1175.

LiangD.SchalC. (1990). Circadian rhythmicity and development of the behavioural response to sex pheromone in male brown-banded cockroaches, Supella longipalpa. — Physiol. Entomol. 15: 355-361.

LinnC.E.CampbellM.G.RoelofsW.L. (1988). Temperature modulation of behavioural thresholds controlling male moth sex pheromone response specificity. — Physiol. Entomol. 13: 59-67.

MathewsL. (2002). Tests of the mate-guarding hypothesis for social monogamy: does population density, sex ratio, or female synchrony affect behavior of male snapping shrimp (Alpheus angulatus)?Behav. Ecol. Sociobiol. 51: 426-432.

Morgan-RichardsM.TrewickS.A.StringerI.N. (2010). Geographic parthenogenesis and the common tea-tree stick insect of New Zealand. — Mol. Ecol. 19: 1227-1238.

Moya-LarañoJ.HalajJ.WiseD.H. (2002). Climbing to reach females: Romeo should be small. — Evolution 56: 420-425.

PaintingC.J.HolwellG.I. (2014a). Exaggerated rostra as weapons and the competitive assessment strategy of male giraffe weevils. — Behav. Ecol. 25: 1223-1232.

PaintingC.J.HolwellG.I. (2014b). Observations on the ecology and behaviour of the New Zealand giraffe weevil (Lasiorhynchus barbicornis). — N.Z. J. Zool. 41: 147-153.

ParkerG.A.BallM.A.StockleyP.GageM.J.G. (1996). Sperm competition games: individual assessment of sperm competition intensity by group spawners. — Proc. Roy. Soc. Lond. B: Biol. Sci. 263: 1291-1297.

ParkerG.A.BallM.A.StockleyP.GageM.J.G. (1997). Sperm competition games: a prospective analysis of risk assessment. — Proc. Roy. Soc. Lond. B: Biol. Sci. 264: 1793-1802.

PitafiK.D.SimpsonR.DayT.H. (1995). Male mate choice for fecund females in seaweed flies. — Pakistan J. Zool. 27: 233-240.

R Core Development Team (2012). R: a language and environment for statistical computing. — The R Project for Statistical ComputingVienna.

ReinholdK.KurtzJ.EngqvistL. (2002). Cryptic male choice: sperm allocation strategies when female quality varies. — J. Evol. Biol. 15: 201-209.

RosénW. (2002). Endogenous control of circadian rhythms of pheromone production in the turnip moth, Agrotis segetum. — Arch. Insect Biochem. Physiol. 50: 21-30.

SimmonsL.W. (2001). Sperm competition and its evolutionary consequences in the insects. — Princeton University PressPrinceton, NJ.

SivinskiJ. (1979). Intrasexual aggression in the stick insects Diapheromera veliei and D. covilleae and sexual dimorphism in the Phasmatodea. — Psyche 85: 395-405.

SivinskiJ. (1980). The effects of mating on predation in the stick insect Diapheromera veliei Walsh (Phasmatodea: Heteronemiidae). — Entomol. Soc. Am. 73: 553-556.

SowerL.L.VickK.W.BallK.A. (1974). Perception of olfactory stimuli that inhibit the responses of male phycitid moths to sex pheromones. — Environ. Entomol. 3: 227-279.

StringerI. (1967). Aspects of reproduction and development in Clitarchus hookeri White. — The University of Auckland Auckland.

StringerI. (1970). The nymphal and imaginal stages of the bisexual stick insect Clitarchus hookeri (Phasmidae: Phasminae). — N.Z. Entomol. 4: 85-95.

SymondsM.R.JohnsonT.L.ElgarM.A. (2012). Pheromone production, male abundance, body size, and the evolution of elaborate antennae in moths. — Ecol. Evol. 2: 227-246.

TakahashiY.WatanabeM. (2011). Male mate choice based on ontogenetic colour changes of females in the damselfly Ischnura senegalensis. — J. Ethol. 29: 293-299.

ThornhillR.AlcockJ. (1983). The evolution of insect mating systems. —, NE.

VepsalainenK.SavolainenR. (1995). Operational sex ratios and mating conflict between the sexes in the water strider Gerris lacustris. — Am. Nat. 146: 869-880.

Wearing-WildeJ. (1996). Mate choice and competition in the barklouse Lepinotus patruelis (Psocoptera: Trogiidae): the effect of diet quality and sex ratio. — J. Insect Behav. 9: 599-612.

WedellN.GageM.J.G.ParkerG.A. (2002). Sperm competition, male prudence and sperm-limited females. — Trends Ecol. Evol. 17: 313-320.

WeirL.K.GrantJ.W.HutchingsJ.A. (2011). The influence of operational sex ratio on the intensity of competition for mates. — Am. Nat. 177: 167-176.

WilligM.R.GarrisonR.W.BaumanA.J. (1986). Population dynamics and natural history of a neotropical walking stick, Lamponius protoricensis rhen (Phasmatodea: Phasmatidae). — Texas J. Sci. 38: 121-137.

ZacharukR.Y. (1985). Antennae and sensilla. — In: Comprehensive insect physiology biochemistry and pharmacologyVol. VI: nervous system: sensory ( KerkutG.A.GilbertL.I. eds). Pergamon PressNew York, NY p.  1-69.


  • View in gallery

    Scanning electron micrographs displaying comparative antennal segment morphology between male (A) and female (B) C. hookeri. The images show; (I) Scapus and pedicel, (II) higher magnification of male basiconic, and female trichoid sensilla, from pedicel, (III) close up of chemosensilla from male antennal segment 9, female 10 and (IV) male antennal segment 9 and female 10.

  • View in gallery

    (a) Season 1. Variation in total abundance of adult male (black) and female (light grey) and the total number of mating pairs (dark gray) of C. hookeri observed over 21 days starting from 23 February 2011 (note day one in season 1 is the equivalent of day 73 in season 2). (b) Season 2 variation of total abundance of C. hookeri over 129 days starting 11 December 2011 of adult male (black) and female (light gray) and mating pairs (dark gray) of C. hookeri observed.

  • View in gallery

    Variation in operational sex ratio (black line) and proportion of individuals mating (dotted line).

  • View in gallery

    Relationship between the total abundance of individuals and the total number of mating pairs found in season 1 (grey, solid line) and season 2 (white dots, large dashed line) for (a) females and (b) males. The line of small dashes is a 1:1 fit to demonstrate how far from a 1:1 ratio this relationship is.

  • View in gallery

    Time (in h) mating pairs were found guarding during documented weeks, where guarding duration is recorded as the time mating pairs were found together until they were seen to separate or could no longer be found together.

  • View in gallery

    The relationship between the body size of females and the number of night male partners remained guarding them for (top) female mass and (bottom) female length.

  • View in gallery

    The relationship between the body size of females and the number of males partners they were found to be mating with. Female body size measurements include mass in grams (a and b), length in mm (c and d), and prothorax length in mm (e and f).

Index Card

Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 12 12 9
Full Text Views 18 18 18
PDF Downloads 3 3 3
EPUB Downloads 0 0 0