Preference of pubescent females for dominant vs subordinate males in root voles 3

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1 49 (3) : A cta Zoologica S inica 3 ( ) ( ) ( ) : (1) ; (2) Y ; (3) 24 h ; [ 49 (3) : ] Preference of pubescent females for dominant vs subordinate males in root voles 3 ZHAO Ya2J un SUN Ru2Yong FAN G Ji2Ming L I Bao2Ming ZHAO Xin2Quan ( Mi nist ry of A gricult ure Key L aboratory f or A gro2biological Envi ronmental Engi neeri ng Chi na A gricult ural U niversity Beiji ng Chi na) ( Instit ute of Ecology Beiji ng Normal U niversity Beiji ng Chi na) ( Northwest Plateau Instit ute of Biology Chi nese Academy of Sciences Xi ni ng Chi na) Abstract A series of animal behavioral experiments were designed to investigate morphological and behavioral factors af2 fecting mate choice in the polygynous root vole ( Microtus oeconom us). Two laboratory experiments on the effect of male dominance on the mate choice of pubescent female root voles were conducted. In an experiment of dyadic interaction of male full siblings the heavier sibling was significantly more dominant than the other so the heavier sibling was regarded as the dominant. In a Y2shape maze experiment female root voles undergoing their first estrus spent more time visiting socializing and being amicable to dominant males than subordinate ones. After the test the three voles were housed to2 gether in a cage for 24 hours the dominant exhibited less latency of intromission than that of the subordinate and the dominant male s copulation with the female was not disrupted by the subordinate male. Combined with the female prefer2 ence for familiar males established in previous experiments ( Zhao et al ) the female preference for dominant males suggests that male social rank could be regarded as such a good indicator of mate choice in virgin female root voles. We hypothesise that interspecific variation in male dominance is consistent with the observed variation in body mass dimor2 phism among monogamous polygynous and promiscuous vole species. Therefore female mate choice for larger males can determine how polygamous vole species differ from monogamous and promiscuous ones [ Acta Zoologica Sinica 49 (3) : (No No ) [ This research was funded by grants from the National Natural Science Foundation of China (No and No ) and the Haibei Alpine Ecosystem Research Station Chinese Academy of Sciences] 33 : net ν 2003 Acta Zoologica Sinica

2 ]. Key words Root vole ( Microtus oeconom us) Pubescent females Mate choice Dyadic interaction of make full sib2 lings Dominance Polygyny F2 F4 ( Zhao et al. 1999) ( Drews 1993 ) Drews (1993) (1982) : 5114 d 4814 d Facemire et al. 42 d d 70 d (20 cm 6 cm 6 cm) 24 h (Dewsbury ) 112 TPX2CP24 ( 46 cm 31 cm 20 cm) ( Hoffmey2 - er 1982 ; Huck et al. 1982a b ; Shapiro et al. 1986a ; Drickmer 1992 ; Horne et al ; Kruczek 1997 ) ( Shapiro et al. 1986b ; 1 Zhang et al ; Zhao et al ; 2002a) ( 2002b) (1983) Y ( Zhao et al ; a b) ( Pierce et al ; Salo et al. 1995) (Dyadic habitation) ( 1 ) 14 5 d 5 d ( ) 2 2 d 1 (? ) 2? 48 h ( M icrot us oecono2 2 3 h m us Pallas) 15 min 2 2

3 3 : Wilcoxon 2 h 2 1 h Chi2square test McNemar Sign test Y 20 min min 30 min 211 : ( ) 5 d ( ) ( ) ( ) [ 1 = g 2 = g n ( = = n = 14 P > 0105) 24 h 3 ( P < 0101) : 1 1 h 1 24 h 2 ( = g = 2016 ( g n = 14 P < ) - - ( = g = 2016 ) ( 015 g n = 14 P < ) ( 1) ) min ( ) 114 SPSS/ PC + 2 = 14 P < 0101) ( 1) The Ob2 ( P < 01001) server ( ) ( ( Student s t ) Fig. 1 Difference in mass body ( Mean S E) between pairs of male siblings and sexual dimorphism in males of similar age to female root voles (Student s t2test 3333 P < n = 14)

4 ( Wilcoxon ) Fig. 2 Frequency in 15 min of dyadic interaction between lighter ( ) and heavier ( ) male sibling root voles and their courtship response to a conspecif ic female housed in a small cage of wire mesh within the test box (Wilcoxon test 33 P < P < n = 14) 015) ( P < 0101) ; 2 1 ; ( P < 0101) ; ; ( ) ( ) ( ) ; ( 2) ( 2) ( ) 1 Table 1 Mating preference of pubescent female root voles for dominant vs subordinate males Behavior Measure Subordinate male (Mean S E) Dominant male (Mean S E) Wilcoxon Test 22tailed P (Visiting) S (Social investigation) F S (Approach) F (Staring) F (Nose vs body) (Naso2genital) F S F S (Coquetting) S (Rejection) S (Aggression) F (Amicableness) F F S 30 min ( ) 14 ( The two males in each trial were full2siblings ; F S : frequency and duration of behaviors in seconds during 30 min tests ; n = 14)

5 3 : h 12 h ( P = Chi2 square test) 10 ( Trivers 1972) 1 ( P = McNe2 mar Sign test) ; 24 h 14 ( P Chi2square test) 13 ( P = McNemar Sign test ) 2 ; ( P = McNemar Sign test) 3 : 1) ( Clutton2Brock et al. 1992) 2 ( Horne et al. 1996) 2 ; 2) 2-2 ( Gilbert 1995) ; 3) (Lambin et al. 2 ( Rozenfeld et al. 1991) ( Tast ( 1966 ; 1982 ; Lambin et al ; Jo2 ) hannesen et al. 1996) ( ) (Lambin et al ; Johannesen et al. 1996) ( 1982 ; Lambin et al. 1992) ( ) ( ( Rozenfeld et al ; Horne et al ; Kruczek 1997) ( ( Potentilla f ruticosa) (Emlen et al. 1977) (Andersson 1994 ; Horne et al. 1996) 1992) 1982) ) (30 min) ( 1982) h 2

6 Capitantio J. P.? 3 ( Capitantio 1991 ; Dewsbury ) ( Andersson 1994) ( Devries et al ; Moore et al ; Plavcan et al ; Post et al. 1999) ( Heske et al. 1990) Emlen S. T. and L. W. Oring 1977 Ecology sexual selection and the evolution of mating system. Science 197 : ( Heske et al ; Ostfeld et al. 1993) Facemire C. F. and G. O. Batzli 1983 Suppression of growth and reproduction by social factors in Microtine rodents : tests of two hy2 : potheses. J. = > > (Boonst ra et al. 1993) 891. ( Tast Heske E. J. and R. S. Ostfeld 1990 Sexual dimorphism in size 1966 ; 1982 ; Lambin et al ; relative size of testes and mating systems in north American voles. Heske et al ; Zhao et al ; J. Mam m. 71 (4) : a b) ( References) Andersson M. Univ. Press Sexual Selection. Princeton NJ : Princeton Boonstra R. B. S. Gilbert and C. J. Krebs 1993 Mating system and sexual dimorphism in mass in microtines. J. Mam m. 74 (1) : Levels of integration and the inheritance of dominance. A ni m. Behav. 42 : Clutton2Brock T. H. O. F. Price and A. D. C. MaColl 1992 Mate retention harassment and the evolution of ungulate leks. Behav. Devries A. Ecol. 3 : C. M. B. Devries S. E. Taymans and C. S. Carter 1996 The effects of stress on social preferences are sexually di2 morphic in prairie voles. Proc. Natl. Acad. Sci. USA 93 : Dewsbury D. A Dominace rank copulatory behavior and differential reproduction. Q. Rev. Biol. 47 : Dewsbury D. A Individual attributes generate contrasting degrees of sociality in voles. In : Tamarin R. H. R. S. Ostfeld S. R. Pugh and G. Bujalska ed. Social Systems and Population Cycle in Voles. Boston : Birkhaser Verlag 1 9. Dewsbury D. A Genes influence behaviour. A ni m. Be2 hav. 42 : Drews C The concept and definition of dominance in animal behavior. Behaviour 125 (3-4) : Drickamer L. C Oestrous female house mice discriminate dominant from subordinate males and sons of dominant from sons of subordinate males by odour cues. A ni m. Behav. 32 : Mam m. 64 : Gilbert A. N Tenacious nipple attachment in rodents : the sibling competition hypothesis. A ni m. Behav. 50 ( 4) : 881 Hoffmeyer I Responses of female bank vole ( Clethrionomys glareolus) to dominant vs subordinate conspecific male and to urine odors from dominant vs subordinate males. Behav. Neural Biol. 36 : Horne T. J. and H. Ylonen 1996 Female bank voles ( Clethriono2 mys glareolus ) prefer dominant males ; but what if there is no choice? Behav. Ecol. Sociobiol. 3 : Huck U. W. and E. M. Banks 1982a Differential attraction of fe2 males to dominant males : olfactory discrimination and mating pref2 erence in the brown lemming ( Lem m us t ri m ucronat us). Behav. Ecol. Sociobiol. 11 : Huck U. W. and E. M. Banks 1982b Male dominance status fe2 male choice and mating success in the brown lemming Lem m us t ri m ucronat us. A ni m. Behav. 30 : Johannesen E. and R. A. Ims 1996 Modeling survival rates : habi2 tat fragmentation and destruction in root vole experimental popula2 tions. Ecology 77 (4) : Kruczek M Male rank and female choice in the bank vole Clethrionomys glareol us. Behavioural Processes 40 :

7 3 : 309 Lambin X. C. J. Kreb and B. Scott 1992 Spacing system of the Liang J. tundra vole ( Microt us oeconom us) during the breeding season in Canada s western Arctic. Can. J. Zool. 70 : R. Z. W. Wang J. X. Zeng and Y. C. Hang 1985 The growth and development in root vole Microt us oeconom us. Acta Biologica Plateau Si nica 1 : [ (2) : ] 1 : ] Moore A. J. P. A. Gowaty W. G. Wallin and P. J. Moore 2001 Sexual conflict and the evolution of female mate choice and male so2 cial dominance. Proc. R. Soc. L ond. B. 268 (1466) : Ostfeld R. S. and E. J. Heske 1993 Sexual dimorphism and mat2 ing system in voles. J. Mam m. 74 (1) : Pierce J. D. J r. and D. A. Dewsbury 1991 Female preferences for unmated versus mated males in two species of voles ( Microt us ochrogaster and M. montanus). J. Com p. Psychol. 105 (2) : Plavcan J. M. and C. P. van Schaik 1997 Intrasexual competition and body weight dimorphism in anthropoid primates. A m. J. Phys. A nthropol. 103 (1) : Post E. R. Langvatn M. C. Forchhamme and N. C. Stenseth 1999 Environmental variation shapes sexual dimorphism in red deer. Proc. N atl. Acad. Sci. USA 96 : Roszenfeld F. M. and R. Rasmont 1991 Odor cue recognition by dominant male bank voles Clethrionomys glareolus. A ni m. Be2 hav. 41 : Salo A. L. and D. A. Dewsbury 1995 Three experiments on mate choice in meadow voles ( Microt us pennsylvanicus). J. Com p. Psychol. 109 (1) : Shapiro L. E. and D. A. Dewsbury 1986a Male dominance fe2 male choice and male copulatory behavior in two species of voles ( Microt us ochrogaster and M. montanus). Behav. Ecol. Socio2 biol. 18 : Shapiro L. E. D. Austin S. E. Ward and D. A. Dewsbury 1986b Familiarity and female mate choice in two species of voles Microt us ochrogaster and M. montanus. A ni m. Behav. 34 : Sun R. Y. and S. W. Zheng 1982 Study on home range of root voles Microt us oeconom us. Acta Theriol. Si n. 2 (2) : [ Tast J The root vole Microt us oeconom us ( Pallas) as an in2 habitant of seasonally flooded land. A nn. Zool. Fenn. 3 : Trivers R. L Parent2offspring conflict. A merican Zoologist 14 : Zhang L. R. Y. Sun and J. M. Fang 2002 Odor preferences of adult male Brandt s vole ( Microt us brandti) female individual scents. Acta Zool. discrimination of Si n. 48 (1) : [ (1) : ] Zhao Y. J. F. D. Tai T. Z. Wang X. Q. Zhao and B. M. Li 2002a Effects of the familiarity on mate choice and mate recogni2 tion in Microt us mandarinus and M. oeconom us. Acta Zool. Si n. 48 (2) : [ 2002a. 48 (2) : ] Zhao Y. J. J. M. Fang and R. Y. Sun 1999 Familiarity and mate choice of female and male root voles ( Microt us oeconom us) in female natural estrus. Acta Theriol. Zhao Y. J. J. M. Fang and R. Si n. 19 (4) : Y. Sun 2000 Study paradigms of mating systems in voles. Acta Theriol. Si n. 20 (1) : [ (1) : ] Zhao Y. J. X. Q. Zhao B. M. Li F. D. Tai and T. Z. Wang 2002b Kin recognition and mate choice of estrous females in root voles ( Microt us oeconom us). Acta Zool. Si n. 48 ( 4) : [ 2002b. 48 (4) : ]

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