Parentage and the evolution of parental behavior

Parentage is the proportion of juveniles in a brood that areoffspring of potential care givers. We analyzed how reductionsin parentage affect the evolution of parental behavior usinga static optimization model. The main benefit of parental effortwas an increase in the survival of offspring, and the main costswere reduced opportunities to seek additional matings or toparasitize neighbors and or reduced survival. Both the costsand benefits included terms for relatedness to young. The effectof parentage depended on (1) whether parents responded in ecologicaltime (facultative response) or in evolutionary time (nonfacultativeresponse), (2) whether the cues enabling assessment of parentagepermitted discrimination among offspring, and (3) whether parentagewas the same among different groups of juveniles (unrestricted)or varied between them (restricted). When parents did not knowtheir own parentage and mean parentage was the same for allmatings, reduced parentage affected the costs and benefits equally,so, as in several previous models, there was no effect on theoptimal level of parental effort. Parentage did affect optimalparental effort when mean parentage to the present brood differedfrom that to young from alternative or future matings. Loweredparentage reduced optimal parental effort when the cost of parentingwas missed opportunities for extrapair copulations or broodparasitism or when parentage was consistently higher in alternativeor future matings. Nonlinear changes in parentage with age gavecomplex trajectories of parental care, with individuals of differentages having similar parentage but exhibiting different levelsof parental effort. Correlations between parentage and othervariables in the model (such as opportunities for additionalmatings) sometimes masked, but never eliminated, the effectsof parentage. When parents could discriminate their own youngin a brood, overall parental effort was reduced, but nepotismwas increased. When parents could not discriminate their ownoffspring but had general cues about average parentage to thebrood, effects varied depending on the costs and benefits ofparental behavior. When parental behavior was costly to caregivers, parentage had more effect than when parenting was notcostly. Likewise, parentage had less effect when care greatlyincreased offspring survival than when care was less necessary.Our analyses reconcile conflicting results from previous modelsand suggest a general framework for analyzing parental behaviorwithin populations and among higher taxonomic groups.  

Extrapair paternity in hooded warblers

We examined the role of extrapair fertilizations (EPFs) in themating system of the hooded warbler (Wilsonia citrina), a monogamoussongbird. DNA fingerprinting revealed that 8 of 17 (47%) femaleshad extrapair young in their first or second brood, and 23 of78 (29%) nestlings were the result of EPFs. Extrapair youngwere signifkandy more likely to occur in first broods than insecond broods. The proportion of EPFs within a brood was stronglybirnodal among broods: nests had 50% or more extrapair youngor none. In seven of eight broods where EPFs occurred, an adjacentmale neighbor was identified as the actual father. Male-likecoloration in females did not reduce the likelihood of havingextrapair young. Females with extrapair young did not receiveless parental care from their mates. All males who obtainedEPFs were mated to fertile females or were feeding offspringat the time they most likely mated with the extrapair female.Our results are consistent with the female control hypothesis,which predicts that females benefit from extrapair copulations(EPCs) and have some control over which males, if any, obtainEPCs. However, we could not reject the alternative hypothesisthat some male neighbors are particularly dominant and aggressiveduring EPC attempts, so females accept these EPCs to minimizecosts.  

Breeding synchrony and extrapair fertilizations in two populations of red-winged blackbirds

We tested the relationship between synchrony of breeding andthe frequency of extrapair fertilizations (EPFs) in two populationsof red-winged blackbirds known to differ in female extrapairbehavior. We found no association between the number of simultaneouslyfertilizable females (temporal neighbors) and EPF rate in eitherpopulation, although a significant difference between populationsin the direction of this relationship (positive where femalesinitiated extrapair copulations and negative where males initiatedthem) suggested a modest difference in the influence of synchrony.Males losing offspring to EPFs tended to have more fertilizablefemales at that time than the actual sires in some analysesbut not in others. We also tested several assumptions underlyingtwo competing hypotheses for the effects of synchrony. We foundno evidence that females pursued extrapair copulations moreoften when other females were synchronous. Rather, females weremore likely to gain EFFs with exirapatr males whose social mateswere not yet building their nests. Synchrony also did not consistentlyaffect male pursuit of exirapair copulations or achievementof EPFs. These results suggest that timing of breeding has someeffects on extrapair activity, but that those effects are bothrelatively weak and influenced by other factors that vary betweenyears or populations.  

海南坡鹿雄鹿发情期的吼叫行为及其在繁殖中的意义

1998年 3～ 5月期间 ,于海南省大田国家级自然保护区的养鹿场内共记录到 7只半野生海南坡鹿(Cervuseldihainanus)雄鹿发情期 4 6次吼叫 ,但未见到雌鹿发出吼叫的现象。记录到的所有吼叫均由年龄大于4 5岁的雄鹿发出 ,其中 1只雄鹿的吼叫达 2 2次 ,占 4 7 8%。雄鹿的吼叫由连续的 3～ 13声组成 ,整个吼叫过程持续 9～ 2 3s ,吼叫频率为 [3 4 74± 6 90 ( 2 5 ) ]声 /min。雄鹿每次的吼叫声数及持续时间存在显著的个体差异。雄鹿的吼叫在白天和夜间均能听见 ,但主要发生于 8:0 0～ 9:5 9及 17:0 0～ 17:5 9两个时间段。吼叫是雄鹿繁殖活动的一个表现指标 ,它与雄鹿的蹭角、泥浴、争斗胜利、交配成功或追逐发情雌鹿等优势行为显著正相关。因此 ,我们认为吼叫是雄鹿发情行为中的一种优势炫耀行为 ,具有明显的繁殖意义。  

Why female crested tits copulate repeatedly with the same partner: evidence for the mate assessment hypothesis

That repeated copulation with the same partner within a singlefertile period is beneficial to the male is generally accepted,but why it should be adaptive to the female is controversialand clear evidence supporting any hypothesis is lacking. Hunteret al. (1993) presented seven hypotheses explaining repeatedmating from the female perspective. Four of them are consistentwith the occurrence of male refusal to copulate: females mighttrade copulations for (1) immediate and or (2) future materialbenefits, or use mating as a mechanism for (3) mate-guardingand or (4) mate-assessment. To test these hypotheses in a populationof crested tits Parus cristatus, we collected data on variationin female solicitation rate, proportion of male refusal, andextra-pair paternity. We found that (1) female solicitationrate was independent of male condition, (2) the proportion ofmale refusal was higher in poor-condition males and (3) femalespaired to poor-condition males sought extra pair paternity.These findings agree with predictions stemming from the mateassessment hypothesis. Therefore, it is suggested that, in crestedtits, male response to female copulation solicitation reflectsmale condition and is used by females to assess male quality  

Patterns of extrapair mating in relation to male dominance status and female nest placement in black-capped chickadees

In sexually promiscuous animals, females may benefit by nestingclose to the edge of their partner's territory to facilitateextrapair copulations. In the present study, we describe theextrapair mating system of black-capped chickadees, Poecileatricapillus, and test whether nest locations are influencedby conspecific attraction to extrapair partners. We conducteda spatial analysis of female mating strategies by using microsatellitepaternity analysis in conjunction with geographic informationsystem (GIS) analysis of nest and territory locations. Extrapairoffspring comprised 52 of 351 offspring (14.8%) and were presentin 19 of 57 broods (33.3%). Females paired to males with lowdominance status in the previous winter's flock hierarchy weremore likely to engage in a mixed reproductive strategy thanwere females paired to males with high dominance status. Femaleshad extrapair copulations and extrapair fertilizations withhigh-ranking males more often than with low-ranking males. Notall extrapair copulations resulted in extrapair fertilizations.Females constructed their nests within 16.8 ± 1.0 m ofthe edge of their partner's territory, significantly closerto the edge of their nearest neighbor's territory than to thecenter of their own partner's territory. Extrapair males usuallyshared territory boundaries with cuckolded males. Females pairedto low-ranking males constructed nests near the territory edgesof neighboring high-ranking males. However, females did nothave extrapair copulations with the neighbor nearest to theirnest or even with the high-ranking neighbor nearest to theirnest. We conclude that conspecific attraction to neighbors mayinfluence nesting location in black-capped chickadees; however,it does not operate by facilitating extrapair copulations.  

Postmating sexual selection: the effects of male body size and recovery period on paternity and egg production rate in a water strider

The role of male body size in postmating sexual selection wasexplored in a semiaquatic insect, the water strider Gerris lateralis.To separate effects of male size per se from those due to numericsperm competition, male recovery period (shown here to be proportionalto ejaculate size) was manipulated independently of body sizein a factorial experiment where virgin females were mated firstwith sterile males and then with focal males. Both relativemale fertilization success and female reproductive rate were measured.The number of sperm transferred increased with male recoveryperiod, an effect that was mediated by longer copulation duration,but there were no effects of body size on ejaculate size. Neithermale size nor recovery period had any significant direct effectson male fertilization success. However, copulation durationinfluenced relative fertilization success, suggesting that malesable to transfer more sperm also achieved higher fertilizationsuccess. Females exercised cryptic female choice by modulatingtheir reproductive rate in a manner favoring large males andmales that were successful in terms of achieving high relativefertilization success. Thus, successful males gained a twofoldadvantage in postmating sexual selection. This study has important implicationsfor previous estimates of sexual selection in this group of insectsbecause pre- and postmating sexual selection will be antagonisticdue to limitations in male sperm production: males mating frequently(high mating success) will on average transfer fewer sperm ineach mating and will hence tend to fertilize fewer eggs permating (low fertilization success).  

中华蜜蜂交配和产卵行为生态学研究

研究了中华蜜蜂(Apis cerana cerana Fab.)蜂王与雄蜂交配行为生态以及蜂王产卵行为生态。结果表明，温度对中华蜜蜂蜂王和雄蜂封盖子期影响，导致蜂王初生重、性成熟时间差异显著(P<0．05)，蜂王和雄蜂认巢飞行次数分别为1．23～1．31和1．08～1．13，持续时间分别为0．12～0．13和0．16～0．20h；蜂王和雄蜂交配飞行次数分别为1．10～1．12和1．01～1．05，持续时间分别为0．22～0．23和0．18～0．23h；蜂王与雄蜂交配最适宜温度为20～28℃，蜂王交配飞行一次侧输卵管的精子数为3．37×10^6～4．15×10^6，自然交配产卵蜂王受精囊精子数为3．55×10^6～3．62×10^6蜂王初生重与产卵量之间呈正相关，周年蜂王产卵量受气候和蜜粉源影响明显。  

Copulation behaviour in mammals: evidence that sperm competition is widespread

We review information on copulation behaviour and sperm competition in mammals using data primarily from the literature. Female mammals of many species regularly copulate with more than one male during each oestrous period. Such multi-male copulations are reported more often in social, compared with solitary species. In addition, the mates of males of polygynous species experience multi-male copulations as often as the mates of males or monogamous species. Male mammals attempt to increase their certainty of paternity through a number of reproductive tactics. Copulation frequency is higher in specks with multi-male copulations compared with other species. Consortships, which can br regarded as a form of mate guarding, are often absent from species in which more than one male copulates with each female during her oestrous period. Most solitary burrow-living small mammal species copulate in the open, whereas social species often copulate inside their burrows. Insurance copulations may occur in many species since high copulation rates occur in four circumstances: (i) when mates are reunited, (ii) when a new male takes over a female, (iii) when a strange male steals a copulation, and (iv) when an audience of males is present.  

Effects of reproductive state and host resource experience on mating decisions in a walnut fly

Prior experience with conspecifics or essential resources, aswell as physiological condition, can have important influenceson an animal's reproductive behavior. While effects of experienceand physiological state (such as reproductive condition) aregenerally treated separately in theoretical discussions, theyoften interact. No previous study has attempted to distinguisheffects of experience on physiological state from other effectsof experience in the context of mating behavior. In a studyof a walnut-infesting tephritid fly (Rhagoletis juglandis),we examined the effects of host fruit experience on mating behavior.We manipulated physiological state in terms of egg load (definedas the number of mature oocytes in a female's ovaries) independentlyof fruit experience to distinguish the effects of these variables.We found that females with high egg loads were significantlymore likely to copulate than low–egg load females; thelevel of fruit experience had no effect on propensity to copulate,except via effects on egg load. In contrast, females with priorexposure to fruit copulated for a significantly shorter durationthan control females, while egg load had no effect on copulationduration. These results suggest that female reproductive conditionand exposure to essential resources can have important, albeitdiverse effects on mating behavior. We discuss how distinguishingdifferent types of variables may provide insight into sexualconflict over mating decisions, as well as which sex controlsspecific aspects of behavior.