The effect of food on laying date and clutch-size in Tengmalm's Owl Aegolius funereus

The breeding of Tengmalm's Owl Aegolius funereus was studied at Umeå, Sweden, during the 1984–85. Mean clutch-size was one egg larger in 1984 than in 1985 despite the later laying in 1984. The difference in clutch-size was related to a better food supply in 1984. Daily weight increase of females during the prelaying period showed a high negative correlation with laying date in 1985, and a high positive correlation with clutch-size independently of laying date in 1984–85. This suggested that food eaten before and during laying had a great and direct influence on both laying date and clutch-size. Many females increased in weight during laying and most others decreased only moderately (relative to egg weight), suggesting that body reserves were not a main source for egg production.
Late breeding females were provided with extra food during the prelaying and laying periods in 1985. Fed females weighed more, bred eight days earlier and laid one more egg than controls. At the same laying dates in mid season, and after heavy snow-fall, clutch-size and female weight were larger in the fed birds than in controls, but this was not so near the end of the laying season. Although the earliest of the fed late breeders weighed more, and probably were less restricted by food availability just before or during laying, they did not lay more eggs than did early breeders. This result suggested some limitation on clutch-size that could not be overcome by the supplementary feeding. Weights of females during laying did not show any consistent relationship with clutch-size during successive laying date intervals, suggesting that clutch-size was not directly related to body condition.     

Clutch size and breeding success in relation to nest-box size in Tengmalm's owl Aegolius funereus

Clutch size and breeding success was related to nest-box size in 215 nests of Tengmalm's owl in the Kauhava region, western Finland in 1966–82. In vole peak years, but not in other years, mean clutch size was positively correlated with the bottom area of the nest-box, increasing on average by 0.005 eggs cm⁻². In successful nests there were fewer nestlings in small and medium-sized than in large boxes. Breeding was most successful in thick-walled boxes with a small entrance diameter. The optimal box for Tengmalm's owl has the following dimensions: bottom area 20 × 20 cm, entrance diameter 8 cm, and wall thickness at least 3 cm.
Hypotheses suggested to explain the "area effect" based on breeding data of hole-nesting passerines do not hold good for Tengmalm's owl. Since there were more stored prey animals in medium-sized and large boxes than in small boxes, I suggest that the "area effect" is related to food availability, which may determine how sensitive the species is to different bottom areas of the boxes, at least in areas with cyclic food (e.g. northern Europe).     

Timing of breeding of Tengmalm's Owl Aegolius funereus in relation to vole dynamics in western Finland

Timing of breeding in Tengmalm's Owl was studied in western Finland for 13 years. During 1973-85, half of the females started laying before 4 April, near the seasonal low of main food abundance (voles), and earlier than in southern Finland or as early as in southeastern Norway. The reason for this latitudinal trend is the shallow snow cover of the study area. The annual variation in the median laying dates was one month and was negatively correlated with the spring abundance of Microtus voles. The mean clutch size was related to the start of laying with early clutches being larger than late ones. These findings accord with the 'food limitation hypothesis', which states that laying begins as soon as the female can accumulate enough energy stores for forming eggs. Early breeding is adaptive, since juveniles of early clutches probably survive better during their first winter. In adults, early nesting improves the chances of rearing two broods per year, allows them to moult after breeding and gives more time to accumulate fat reserves to survive the winter. Tengmalm's Owl is one of the earliest breeders among North European birds. This is possible because of its hole-nesting and resident habits, small body-size in relation to the main prey and the greatest sexual size dimorphism among European owls.     

Contrasting phenotypic correlations in food provision of male Tengmalm's owls (Aegolius funereus) in a temporally heterogenous environment

Summary We examined the food provision rate of male Tengmalm's owls,Aegolius funereus, during one 3 year vole cycle consisting of consecutive low, increase and peake vole years. The data were collected in the midnestling period when males provisioned the whole family. In the low vole year, males with a low loading index (g/cm²) of flying area fed their offspring more often than did males with a high loading index, whereas in the peak vole year the opposite trend was evident. Similar relationships were found in the food mass provisioned to the nest. In the increase vole year, male body size had no effect on feeding efficiency. In the peak vole year, when large voles are abundant, heavy males preyed on larger voles than were generally available in their territories, indicating that largeness may increase strike power in hunting attempts. In the low vole year, when breeding is costly due to food scarcity and extensive hunting area, small males are more economical fliers and efficient hunters than large males. The contrasting trends in correlations between male size and feeding efficiency in years of vole abundance versus scarcity suggest that no fixed phenotype may most efficiently cope with variable food supply.     

Genetic similarity among Eurasian subspecies of boreal owls Aegolius funereus

Boreal owls Aegolius funereus (referred to as Tengmalm's owls in Europe) breed in boreal forests throughout the Holarctic region and in high-elevation subalpine forests further south. They are currently classified as seven subspecies; six found throughout Eurasia, and one in North America. The geographic distribution of boreal owls in North America and Eurasia is similar, as are their patterns of dispersal and irruption. Because a recent genetic study of boreal owls in North America found very little genetic differentiation among widely disparate locations, we expected that boreal owls in Eurasia similarly would have very little genetic differentiation. Using seven microsatellite markers, we analyzed genetic samples from 275 boreal owls in North America, 36 in Norway, and five in eastern Russia. We found no detectable genetic differentiation between Norwegian and Russian owls, but notable differentiation between North American and Eurasian owls. Low intra-continental genetic differentiation likely results from high rates of long-distance dispersal among subpopulations of boreal owls. In light of these results, we recommend further genetic sampling of boreal owls throughout Eurasia in order to determine whether six separate subspecies here are warranted.     

Differential Movement Patterns of Juvenile Tengmalms Owls (Aegolius funereus) during the Post-Fledging Dependence Period in Two Years with Contrasting Prey Abundance

Fledgling behaviour and movement patterns throughout the post-fledging dependence period (PFDP), especially in relation to changing environmental conditions, have been rarely studied, despite the fact that this period is recognized as of crucial significance in terms of high mortality of juveniles. The PFDP can extend over quite a protracted period, particularly in birds of prey, and a knowledge of the movement patterns of individuals is fundamental for understanding mechanisms underlying survival, habitat use and dispersion. We radiotracked 39 fledglings of the Tengmalm’s owl (Aegolius funereus) in two years with different availability of prey: 2010 (n = 29) and 2011 (n = 10) and obtained 1455 daily locations. Fledglings reached independence on average in 45 days after fledging in 2010 (n = 22) and 57 days in 2011 (n = 6). Within years, the most important measures influencing the distance moved from the nest box were age of fledglings and number of surviving siblings present. Individual home range size and duration of PFDP in particular were dependent on maximal number of siblings seen outside the nest box. In the season with low prey availability fledglings were observed at greater distances from the nest box than in the year with higher prey availability (mean distance: 350 m in 2010 and 650 m in 2011) and occupied larger home ranges (mean: 30.3 ha in 2010 and 57.7 ha in 2011). The main factor causing these differences between years was probably the different availability of prey in these two years, affecting breeding success and post-fledging survivorship of the Tengmalm’s owls.     

Mating order-dependent female mate choice in the polygynandrous common lizard Lacerta vivipara

Recent studies indicate that directional female mate choice and order-dependent female mate choice importantly contribute to non-random mating patterns. In species where females prefer larger sized males, disentangling different hypotheses leading to non-random mating patterns is especially difficult, given that male size usually correlates with behaviours that may lead to non-random mating (e.g. size-dependent emergence from hibernation, male fighting ability). Here we investigate female mate choice and order-dependent female mate choice in the polygynandrous common lizard (Lacerta vivipara). By sequentially presenting males in random order to females, we exclude non-random mating patterns potentially arising due to intra-sexual selection (e.g. male–male competition), trait-dependent encounter probabilities, trait-dependent conspicuousness, or trait-dependent emergence from hibernation. To test for order-dependent female mate choice we investigate whether the previous mating history affects female choice. We show that body size and body condition of the male with which a female mated for the first time were bigger and better, respectively, than the average body size and body condition of the rejected males. There was a negative correlation between body sizes of first and second copulating males. This indicates that female mate choice is dependent on the previous mating history and it shows that the female’s choice criteria are non-static, i.e. non-directional. Our study therefore suggests that context-dependent female mate choice may not only arise due to genotype-environment interactions, but also due to other female mating strategies, i.e. order-dependent mate choice. Thus context-dependent female mate choice might be more frequent than previously thought.     

Mating patterns,mate choice,and birth season heterosexual relationships in free-ranging rhesus macaques

Birth season adult heterosexual nonkin relationships of 50 free-ranging female rhesus macaques (Macaca mulatta) in two social groups at Cayo Santiago, Puerto Rico were examined using focal follow (289 hr) and ad lib data. Eighty-eight percent of subjects had at least one relationship characterized by particularly high frequencies of spatial proximity, grooming, or both. These were designated “friendships.” Males intervened in aggressive interactions more frequently on behalf of Friends than non-Friends. Female aggressive support of males was extremely rare. Higher-ranking males experienced more friendships than lower-ranking males. High-ranking females had higher-ranking Friends than low-ranking females. Older females had higher-ranking Friends than younger females. Females groomed high-ranking Friends more than they were groomed by them, whereas they groomed low-ranking Friends less than they were groomed by them. In one social group, high-ranking females were more likely than low-ranking females to groom their Friends more than they were groomed by them. Males were more responsible than females for spatial proximity maintenance in 9 of 14 Friend dyads for which sufficient data were available. Neither male nor female dominance rank affected responsibility for proximity maintenance in Friend dyads. Eight of 24 females had friendships with males with whom they had completed copulations during their conception peri-ovulatory period of the preceding mating season. Two of 19 females completed peri-ovulatory copulations with Friends during the following mating season. Friendship was not correlated with either of two demonstrated female mate choice indicators: (1) proximity maintenance during estrus; or (2) cooperation with male “hip-grasp” courtship attempts. Males directed “muzzle-up” courtship signals at lower rates toward Friends than toward non-Friends. These and other investigators' results indicate that (1) protection from aggression is the primary benefit to female rhesus macaques of birth season heterosexual relationships; (2) the most effective protectors are in greatest demand as Friends; and (3) friendship has no effect or an inhibitory effect on mate choice in this species. Benefits to males of friendships were not apparent from this study but may include coalitional support against lower-ranking males.     

Sexual selection and mate choice

The past two decades have seen extensive growth of sexual selection research. Theoretical and empirical work has clarified many components of pre- and postcopulatory sexual selection, such as aggressive competition, mate choice, sperm utilization and sexual conflict. Genetic mechanisms of mate choice evolution have been less amenable to empirical testing, but molecular genetic analyses can now be used for incisive experimentation. Here, we highlight some of the currently debated areas in pre- and postcopulatory sexual selection. We identify where new techniques can help estimate the relative roles of the various selection mechanisms that might work together in the evolution of mating preferences and attractive traits, and in sperm-egg interactions.     

Stress hormones and mate choice

A few recent studies have suggested that glucocorticoid stress hormones can play a role in sexual selection. In terms of mate choice, these studies have shown that individuals can exhibit preferences for mates with either low baseline or peak glucocorticoid levels. This appears to occur because stress hormones can be key mediators of many condition-dependent, sexually selected traits that serve as honest signals of mate quality. We suggest that our understanding of how sexual selection can influence the glucocorticoid stress response will be expanded by studying the interactions among glucocorticoid stress hormones, sex steroids and body condition.     

甘肃省莲花山鬼鸮繁殖巢址记述

鬼鸮在世界上呈环北极分布,其甘肃亚种自1928年在甘肃西北部的天堂寺首次发现后,80多年来在甘肃一直未见报道,其它分布点亦非常零散。1999～2003年,作者在甘肃省莲花山区发现鬼分布,并记录鬼鸮繁殖巢址3个。鬼鸮营巢于针叶树的树洞中,雌雄共同育雏。     

Sex roles and mutual mate choice matter during mate sampling

The roles of females and males in mating competition and mate choice have lately proven more variable, between and within species, than previously thought. In nature, mating competition occurs during mate search and is expected to be regulated by the numbers of potential mates and same-sex competitors. Here, we present the first study to test how a temporal change in sex roles affects mating competition and mate choice during mate sampling. Our model system (the marine fish Gobiusculus flavescens) is uniquely suitable because of its change in sex roles, from conventional to reversed, over the breeding season. As predicted from sex role theory, courtship was typically initiated by males and terminated by females early in the breeding season. The opposite pattern was observed late in the season, at which time several females often simultaneously courted the same male. Mate-searching females visited more males early than late in the breeding season. Our study shows that mutual mate choice and mating competition can have profound effects on female and male behavior. Future work needs to consider the dynamic nature of mating competition and mate choice if we aim to fully understand sexual selection in the wild.     

Romantic love: a mammalian brain system for mate choice

Mammals and birds regularly express mate preferences and make mate choices. Data on mate choice among mammals suggest that this behavioural 'attraction system' is associated with dopaminergic reward pathways in the brain. It has been proposed that intense romantic love, a human cross-cultural universal, is a developed form of this attraction system. To begin to determine the neural mechanisms associated with romantic attraction in humans, we used functional magnetic resonance imaging (fMRI) to study 17 people who were intensely 'in love'. Activation specific to the beloved occurred in the brainstem right ventral tegmental area and right postero-dorsal body of the caudate nucleus. These and other results suggest that dopaminergic reward and motivation pathways contribute to aspects of romantic love. We also used fMRI to study 15 men and women who had just been rejected in love. Preliminary analysis showed activity specific to the beloved in related regions of the reward system associated with monetary gambling for uncertain large gains and losses, and in regions of the lateral orbitofrontal cortex associated with theory of mind, obsessive/compulsive behaviours and controlling anger. These data contribute to our view that romantic love is one of the three primary brain systems that evolved in avian and mammalian species to direct reproduction. The sex drive evolved to motivate individuals to seek a range of mating partners; attraction evolved to motivate individuals to prefer and pursue specific partners; and attachment evolved to motivate individuals to remain together long enough to complete species-specific parenting duties. These three behavioural repertoires appear to be based on brain systems that are largely distinct yet interrelated, and they interact in specific ways to orchestrate reproduction, using both hormones and monoamines. Romantic attraction in humans and its antecedent in other mammalian species play a primary role: this neural mechanism motivates individuals to focus their courtship energy on specific others, thereby conserving valuable time and metabolic energy, and facilitating mate choice.     

鬼鸮甘肃亚种繁殖期叫声研究

利用叫声回放和声谱分析对莲花山自然保护区鬼甘肃亚种(Aegolius funereus beickianus)繁殖期的叫声进行了研究。共记录了领域叫声、尖叫声等6种叫声,分析了各种叫声的特征及与行为的联系。发现不同地点录制的领域叫声存在差异,但同一个体的叫声也有变化。甘肃亚种的领域叫声和欧洲的指名亚种A.f.fu-nereus及北美的亚种A.f.richardsoni相比较,单音数量少,单音长度小,频率高,但差异不大。     

Alternative hypotheses linking the immune system and mate choice for good genes

Why do males often have extravagant morphological and behavioural traits, and why do females prefer to mate with such males? The answers have been the focus of considerable debate since Darwin''s The descent of man, and selection in relation to sex appeared in 1871. Recently the broadening of investigation to include fields outside evolutionary biology has shed new light on mate choice and sexual selection. Here, we focus on a specific set of hypotheses relating the biology of resisting disease-causing organisms with the production of condition-dependent sexual signals (advertisements). We present a framework that distinguishes three different hypotheses about trade-offs within the immune system that affect general condition. Hamilton and Zuk''s original hypothesis suggests that hosts fight off disease through resistance to particular pathogens, which consequently lowers resistance to other pathogens. Changes in pathogens over evolutionary time in turn favours changes in which genes confer the best resistance. Alternatively, the immunocompetence hypotheses suggests that the energetic costs of mounting a response to any pathogen compete for resources with other things, such as producing or maintaining advertisements. Finally, improving resistance to pathogens could increase the negative impacts of the immune system on the host, via immunopathologies such as allergies or autoimmune diseases. If both disease and immunopathology affect condition, then sexual advertisements could signal a balance between the two. Studies of hypothesized links between genes, condition, the immune system and advertisements likely will require careful consideration of which hypothesis is being considered, and may necessitate different measures of immune system responses and different experimental protocols.     

Indirect mate choice, direct mate choice and species recognition in a bower-building cichlid fish lek

Sexual selection arising through female mate choice typically favours males with larger, brighter and louder signals. A critical challenge in sexual selection research is to determine the degree to which this pattern results from direct mate choice, where females select individual males based on variation in signalling traits, or indirect mate choice, where male competition governs access to reproductively active females. We investigated female mate choice in a lekking Lake Malawi cichlid fish, Hemitilapia oxyrhynchus, in which males build and aggressively defend sand 'bowers'. Similar to previous studies, we found that male reproductive success was positively associated with bower height and centrality on the lek. However, this pattern resulted from males holding these territories encountering more females, and thus their greater success was due to indirect mate choice. Following initial male courtship, an increase in the relative mating success of some males was observed, but this relative increase was unrelated to bower size or position. Crucially, experimentally manipulating bowers to resemble those of a co-occurring species had no appreciable effect on direct choice by females or male spawning success. Together, these results suggest indirect mate choice is the dominant force determining male-mating success in this species, and that bowers are not signals used in direct mate choice by females. We propose that, in this species, bowers have a primary function in intraspecific male competition, with the most competitive males maintaining larger and more central bowers that are favoured by sexual selection due to higher female encounter rates.     

Alternative mechanisms of nonindependent mate choice

Aspects of the environment, including the social environment, can contribute to intrapopulation variation in mating preferences. One example of the effect of social environment on mate preferences is mate choice copying; however, other types of socially influenced (nonindependent) choice might exist. We develop a list of such alternatives based on possible physiological or psychological mechanisms, evaluate the evidence distinguishing one from another and clarify some controversial aspects of mate choice copying. This framework reveals many ways in which one female's mate choice can influence that of another, and suggests a broader array of hypotheses about the selective forces acting on such mechanisms. Because nonindependent choice can occur in a variety of ways, it could be more important for understanding patterns of mate choice than current theory suggests. Copyright 2000 The Association for the Study of Animal Behaviour.