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Perspectives on over-marking: is it good to be on top?

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Abstract

What we refer to as over-marking occurs when one individual places its scent mark on top of, touching, or adjacent to the scent mark of another individual, usually a conspecific. Over-marking frequently occurs among mammals that share common paths, trails, and runways. Despite its ubiquity among terrestrial mammals, we know little about how individuals respond to over-marks and the function(s) of over-marking. Studies on voles and golden hamsters indicate that after exploring an over-mark, individuals respond selectively to the mark of the top-scent donor relative to that of the bottom-scent donor. Thus, individuals may be able to focus their attention on a particular scent mark relevant at a particular time and in a particular context, neglecting other scent marks that are present. The function(s) of over-marking are examined within the framework of ten hypotheses. Several hypotheses are plausible. However, the bulk of the literature is consistent with hypotheses stating that over-marking serving a role in olfactory communication between opposite and same-sex conspecifics. Lastly, we postulate the costs and benefits that may be garnered by the top-scent donor of an over-mark.

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References

  • Begg CM, Begg KS, Du Toit JT, Mills MGL (2003) Scent-marking behaviour of the honey badger, Mellivora capensis (Mustelidae), in the southern Kalahari. Anim Behav 66:917–929

    Article  Google Scholar 

  • Bel MC, Couton J, Sreng L, Allaine D, Bagneres AG, Clement JL (1999) Social signals involved in scent-marking behavior by cheek-rubbing in Alpine marmots (Marmota marmota). J Chem Ecol 25:2267–2283

    Article  CAS  Google Scholar 

  • Biben M (1980) Over-marking of alien conspecific odors by Mongolian gerbils. Biol Behav 5:139–145

    Google Scholar 

  • Boonstra R, Xia X, Pavone L (1993) Mating system of the meadow vole, Microtus pennsylvanicus. Behav Ecol 4:83–89

    Article  Google Scholar 

  • Brashares JS, Arcese P (1999) Scent marking in territorial African antelope: II the economics of marking with faeces. Anim Behav 57:11–17

    Article  PubMed  Google Scholar 

  • Brown RE, Macdonald DW (eds) (1985) Social odours in mammals, vol 1 and 2. Oxford University Press, Oxford

  • Carter CS, Getz LL (1993) Monogamy and the prairie vole. Sci Am 268:100–106

    Article  PubMed  CAS  Google Scholar 

  • Cohen AB, Johnston RE, Kwon A (2001) How golden hamsters (Mesocricetus auratus) discriminate top from bottom flank scents in over-marks. J Comp Psychol 115:241–247

    Article  PubMed  CAS  Google Scholar 

  • Daly M (1977) Some experiments on the functional significance of scent marking in gerbils (Meriones unguiculatus). J Physiol Comp Psychol 91:1082–1094

    Article  Google Scholar 

  • Dewsbury DA (1990) Individual attributes generate contrasting degrees of sociality in voles. In: Tamarin RH, Ostfeld RS, Pugh SR, Bujalska G (eds) Social systems and population cycles in voles. Birkhauser, Basel, pp 1–9

    Google Scholar 

  • Drea CM, Sacha NV, Kim HS, Weldele ML, Glickman SE (2002) Responses to olfactory stimuli in spotted hyenas (Crocuta crocuta) II. Discrimination of conspecific scent. J Comp Psychol 116:342–349

    Article  PubMed  Google Scholar 

  • Ferkin MH (1999a) Meadow voles (Microtus pennsylvanicus, Arvicolidae), over-mark and adjacent mark the scent marks of same-sex conspecifics. Ethology 105:825–837

    Article  Google Scholar 

  • Ferkin MH (1999b) Over-marking and adjacent-marking may be used as competitive tactics during odor communication in voles. In: Johnston RE, Muller-Schwarze D, Sorenson PW (eds) Advances in chemical signals in vertebrates, number 8. Plenum Press, New York, pp 239–246

  • Ferkin MH (2001) The response of individuals to over-marks of conspecifics differs between two species of microtine rodents. In: Marchlewska-Koj A, Lepri JJ, Muller-Schwarze D (eds) Chemical signals in vertebrates, vol 9. Plenum Press, New York, pp 343–346

  • Ferkin MH, Dunsavage J, Johnston RE (1999) What kind of information do meadow voles, Microtus pennsylvanicus, use to distinguish between the odors of the top and bottom-scent donors of an over-mark? J Comp Psychol 113:43–51

    Article  Google Scholar 

  • Ferkin MH, Mech SG, Paz-y-Mino CG (2001a) Scent marking in meadow voles and prairie voles: a test of three hypotheses. Behaviour 138:1319–1336

    Article  Google Scholar 

  • Ferkin MH, Leonard ST, Bartos K, Schmick MK (2001b) Meadow voles and prairie voles differ in the length of time they prefer the top-scent donor of an over-mark. Ethology 107:1099–1014

    Article  Google Scholar 

  • Ferkin MH, Lee DN, Leonard ST (2004a) The reproductive state of female voles affects their scent marking behavior and the responses of male conspecifics to such marks. Ethology 110:257–272

    Article  Google Scholar 

  • Ferkin MH, Li HZ, Leonard ST (2004b) Meadow voles and prairie voles differ in the percentage of conspecific marks that they over-mark. Acta Ethol 7:1–7

    Article  Google Scholar 

  • Ferkin MH, Pierce AA, Sealand RO, delBarco-Trillo J (2005) Meadow voles, Microtus pennsylvanicus, can distinguish more over-marks from fewer over-marks. Anim Cogn 8:82–89

    Article  Google Scholar 

  • Getz LL, Carter CS (1996) Prairie vole partnerships. Am Sci 84:56–62

    Google Scholar 

  • Gosling LM (1982) A reassessment of the function of scent marking in territories. Z Tierpsychol 60:89-118

    Google Scholar 

  • Gosling LM, Roberts SC (2001) Scent marking in male mammals: cheat-proof signals to competitors and mates. Adv Study Behav 30:169–217

    Article  Google Scholar 

  • Hayes RA, Richardson BJ, Wyllie SG (2002) Semiochemicals and social signaling in the wild European rabbit in Australia: I. Scent profiles of chin gland secretion from the field. J Chem Ecol 28:363–384

    Article  PubMed  CAS  Google Scholar 

  • Heymann E (1998) Sex differences in olfactory communication in a primate, the moustached tamarin, Saginus mystax (Callitrichinae). Behav Ecol Sociobiol 43:37–45

    Article  Google Scholar 

  • Huck UW, Lisk RD, Gore AC (1985) Scent marking and mate choice in the golden hamster. Physiol Behav 35:389–393

    Article  PubMed  CAS  Google Scholar 

  • Humphries RE, Robertson DH, Beynon R J, Hurst JL (1999) Unravelling the chemical basis of competitive scent marking in mice. Anim Behav 58:1177–1190

    Article  PubMed  Google Scholar 

  • Hurst JL (1990a) Urine marking in populations of wild house mice, Mus domesticus Rutty I. Communication between males. Anim Behav 40:209–222

    Article  Google Scholar 

  • Hurst JL (1990b) Urine marking in populations of wild house mice, Mus domesticus Rutty II. Communication between females. Anim Behav 40:223–232

    Article  Google Scholar 

  • Hurst JL (1990c) Urine marking in populations of wild house mice, Mus domesticus Rutty III. Communication between the sexes. Anim Behav 40:233–243

    Article  Google Scholar 

  • Johnson RP (1973) Scent marking in mammals. Anim Behav 21:521-535

    Article  Google Scholar 

  • Johnston RE (1999) Scent over-marking: how do hamsters know whose scent is on top and why should it matter. In: Johnston RE, Muller-Schwarze D, Sorenson PW (eds) Advances in chemical signals in vertebrates, vol 8. Plenum Press, New York, pp 227–238

  • Johnston RE (2001) Scent over-marking: a sexually selected trait with specialized mechanisms for scent deposition and perception. Presentation at the 81st American Society of Mammalogist Meeting

  • Johnston RE (2003) Chemical communication in rodents: from pheromones to individual recognition. J Mammal 84:1141–1162

    Article  Google Scholar 

  • Johnston RE, Bhorade A (1998) Perception of scent over-marks by golden hamsters (Mesocricetus auratus). Novel mechanisms for determining which individual’s mark are on top. J Comp Psychol 112:230–243

    Article  PubMed  CAS  Google Scholar 

  • Johnson CN, Johnson KA (1983) Behaviour of the bilby, Macrotis lagostis (Reid), Marsupiala: Thylacomyidae) in captivity. Aust Wildl Res 10:77–87

    Article  Google Scholar 

  • Johnston RE, Chiang G, Tung C (1994) The information in scent over-marks of golden hamsters. Anim Behav 48:323–330

    Article  Google Scholar 

  • Johnston RE, Munver R, Tung C (1995) Scent counter marks: selective memory for the top scent by golden hamsters. Anim Behav 49:1435–1442

    Article  Google Scholar 

  • Johnston RE, Sorokin ES, Ferkin MH (1997a) Scent counter-marking by male meadow voles: females prefer the top-scent male. Ethology 103:443–453

    Article  Google Scholar 

  • Johnston RE, Sorokin ES, Ferkin MH (1997b) Female voles discriminate males’ over-marks and prefer top-scent males. Anim Behav 54:679–690

    Article  Google Scholar 

  • Kappeler PM (1998) To whom it may concern: the transmission and function of chemical signals in Lemur catta. Behav Ecol Sociobiol 42:411–421

    Article  Google Scholar 

  • Kohli KL, Ferkin MH (1999) Over-marking and adjacent marking are influenced by sibship in prairie voles, Microtus ochrogaster. Ethology 105:1–11

    Google Scholar 

  • Kruuk H (1995) Wild otters: predation and populations. Oxford University Press, Oxford

    Google Scholar 

  • Lazaro-Perea C, Snowdon CT, Fatima Arruda M (1999) Scent-marking behavior in wild groups of common marmosets (Callithrix jacchus). Behav Ecol Sociobiol 46:313–324

    Article  Google Scholar 

  • Leonard ST, Ferkin MH, Johnson MM (2001) The response of meadow voles to an over-mark in which the two donors differ in gonadal hormone status. Anim Behav 62:1171–1177

    Article  Google Scholar 

  • Lewis RJ (2005) Sex differences in scent-marking in sifaka: mating conflict or male services? Am J Phys Anthropol 128:389–398

    Article  PubMed  Google Scholar 

  • Macdonald DW (1980) Patterns of scent marking with urine and feces among carnivore communities. Symp Zool Soc Lond 45:107–139

    Google Scholar 

  • Madison DM (1980) An integrated view of the social biology of meadow voles, Microtus pennsylvanicus. Biologist 62:20–33

    Google Scholar 

  • McClintock MK (2002) Pheromones, Odors, and Vasanas: the neuroendocrinology of social chemosignals in humans and animals. In: Pffaf D (ed) Hormones, Brain, and Behavior. Elsevier, Amsterdam, pp 797–870

  • Mertl-Millhollen AS, Goodmann PA, Klinghammer E (1986) Wolf scent-marking with raised-leg urination. Zool Biol 5:7–20

    Article  Google Scholar 

  • Nevison CM, Barnard CJ, Beynon RJ, Hurst JL (2000) The consequences of inbreeding for recognizing competitors. Proc Biol Sci 267:687–694

    Article  PubMed  CAS  Google Scholar 

  • Palagi E, Telara S, Borgognini Tarli SM (2004) Reproductive strategies in Lemur catta: balance among sending, receiving, and countermarking scent signals. Int J Primatol 25:1019–1031

    Article  Google Scholar 

  • Penzhorn BJ (1984) A long-term study of social organization and behaviour of Cape Mountain zebras (Equs zebra zebra). Zeit Tierpsychol 64:97–146

    Google Scholar 

  • Peterson SC (1988) Chemical trail following by caterpillars of Malacosoma neustria. J Chem Ecol 14:815–824

    Article  CAS  Google Scholar 

  • Rich TJ, Hurst JL (1998) Scent marks as reliable signals of competitive ability of mates. Anim Behav 56:727–735

    Article  PubMed  Google Scholar 

  • Rich TJ, Hurst JL (1999) The competing counter-marks hypothesis: reliable assessment of competitive ability by potential mates. Anim Behav 58:1027–1037

    Article  PubMed  Google Scholar 

  • Roberts SC, Dunbar RIM (2000) Female territoriality and the function of scent-marking in a monogamous antelope (Oreotragus oreotragus). Behav Ecol Sociobiol 47:417–423

    Article  Google Scholar 

  • Rosell F, Bjorkoyli T (2002) A test of the dear enemy phenomenon in the Eurasian beaver. Anim Behav 63:1073–1078

    Article  Google Scholar 

  • Rosell F, Johnson G, Parker H (2000) Eurasian beavers (Castor fiber) behavioral response to simulated territorial intruders. Can J Zool 78:931–935

    Article  Google Scholar 

  • Rostain RR, Ben-David M, Groves P, Randall JA (2004) Why do river otters scent mark? An experimental test of several hypotheses. Anim Behav 68:703–711

    Article  Google Scholar 

  • Rozenfeld FM, Rasmont R (1991) Odour cue recognition by dominant male bank voles, Clethrionomys glareolus. Anim Behav 41:839–850

    Article  Google Scholar 

  • Rozenfeld FM, LeBoulenge E, Rasmont R (1987) Urine marking by male bank voles (Clethrionomys glareolus Schreber 1780; Microtidae, Rodentia) in relation to social rank. Can J Zool 65:2549–2601

    Article  Google Scholar 

  • Sillero-Zubiri C, Macdonald DW (1998) Scent-marking and territorial behaviour of Ethiopian wolves Canis simensis. J Zool 245:351–361

    Article  Google Scholar 

  • Sliwa A, Richardson PRK (1998) Responses of aardwolves, Proteles cristatus, Sparrman 1783, to translocated scent marks. Anim Behav 56:137–146

    Article  PubMed  Google Scholar 

  • Smith TE, Abbott DH (1999) Behavioral discrimination between circumgenital odor from peri-ovulatory dominant and anovulatory female common marmosets (Callithrix jacchus). Am J Primatol 46:265–284

    Article  Google Scholar 

  • Smith TE, Gordon SJ (2002) Sex differences in olfactory communication in Saguinus labiatus. Int J Primatol 23:429–441

    Article  Google Scholar 

  • Stralendorf FV (1986) Urinary signaling pheromone and specific behavioral response in tree shrews (Tupaia belangeri) I. Basic investigations for a bioassay. J Chem Ecol 12:1573–1561

    Article  Google Scholar 

  • Thiessen DD, Rice M (1976) Mammalian scent gland marking and social behavior. Psychol Bull 83:505–539

    Article  PubMed  CAS  Google Scholar 

  • Thom MD, Hurst JL (2004) Individual recognition by scent. Ann Zool Fenn 41:765–787

    Google Scholar 

  • Thomas SA (2002) Scent marking and mate choice in the prairie vole, Microtus ochrogaster. Anim Behav 63:1121–1127

    Article  Google Scholar 

  • Thomas SA, Wolff JO (2002) Scent marking in voles: a reassessment of over marking, counter marking, and self-advertisement. Ethology 108:51–62

    Article  Google Scholar 

  • Wilcox RM, Johnston RE (1995) Scent-counter marks: specialized mechanisms of perception and response to individual odors in golden hamsters (Mesocricetus auratus). J Comp Psychol 109:349–356

    Article  PubMed  CAS  Google Scholar 

  • Wolff JO (2003) Laboratory studies with rodents: facts or artifacts? Bioscience 53:421–427

    Article  Google Scholar 

  • Wolff JO, Mech SG, Thomas SA (2002) Scent marking in female prairie voles: a test of alternative hypotheses. Ethology 108:483–494

    Article  Google Scholar 

  • Woodward Jr RL, Schmick MK, Ferkin MH (1999) Response of prairie voles, Microtus ochrogaster (Rodentia, Arvicolidae), to scent over-marks of two same-sex conspecifics: a test of the scent masking hypothesis. Ethology 105:1009–1017

    Article  Google Scholar 

  • Woodward Jr RL, Bartos K, Ferkin MH (2000) Meadow voles (Microtus pennsylvanicus) and prairie voles (M. ochrogaster) differ in their responses to over-marks from opposite- and same-sex conspecifics. Ethology 106:979–992

    Article  Google Scholar 

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Acknowledgments

We thank L. Drickamer, J. delBarco-Trillo, A. Vaughn, and L. LaDage for commenting on earlier drafts of this manuscript. This work was supported by National Science Foundation grant IOB 0444553 and NIH grant AG 16594 and HD 049525.

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  1. Biology Department, The University of Memphis, Ellington Hall, Memphis, TN, 38152, USA

    Michael H. Ferkin & Andrew A. Pierce

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  1. Michael H. Ferkin
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  2. Andrew A. Pierce
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Correspondence to Michael H. Ferkin.

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Ferkin, M.H., Pierce, A.A. Perspectives on over-marking: is it good to be on top?. J Ethol 25, 107–116 (2007). https://doi.org/10.1007/s10164-006-0012-1

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