Space,density and extra-pair matings have opposing impacts on male and female reproductive success

Many territorial species have a mating system characterized by males establishing home ranges in the breeding grounds prior to females, resulting in males competing for territories and females choosing a mate upon their arrival. It remains unknown, however, how the outcomes of decisions surrounding territory establishment and mate choice are influenced by the spatial configuration of the breeding grounds. We use a spatially explicit, individual-based model to investigate the sex-specific effects of these decisions on reproductive success. In our model, males that arrive earlier obtain higher quality territories and improve their chances for extra-pair copulations. Females can choose their mate to maximize the quality of the male or to attempt to minimize the density of other females near their nesting site to avoid competition. Females therefore face a tradeoff between high-density regions around high-quality males and low-quality males in areas of low competition. Our model predicts a negative correlation between male and female reproductive success under a wide range of conditions when the majority of the territories are on the margins of the breeding area. Most notably, this sexual conflict arises as an edge effect suggesting that fragmentation of breeding habitats could impact the consequences of mate choice in many species with territorial breeding habits.     

Exposure to developing females induces polyuria, polydipsia, and altered urinary levels of creatinine, 17β-estradiol, and testosterone in adult male mice (Mus musculus)

Novel male mice can accelerate reproductive maturation in proximal developing females, an effect mediated by the chemistry of the males' urine. Exogenous estrogens can similarly accelerate female sexual development. In Experiment 1, adult male mice were housed across wire grid from either empty compartments or those containing post-weanling females. Proximity of females caused males to urinate more, progressively over days of exposure, with most urination directed towards females' compartments. Male urine collected after 5 days in these conditions was analyzed by enzyme immunoassay for 17β-estradiol, testosterone, and creatinine. Urinary creatinine of isolated males significantly exceeded that of female-exposed males. Unadjusted urinary steroids also trended toward higher levels in isolates, but creatinine-adjusted estradiol and testosterone of female-exposed males significantly exceeded that of isolated males. In Experiment 2, measurement of water consumption indicated significantly greater drinking by female-exposed as opposed to isolated males. In Experiment 3, males were housed in isolation or beside post-weanling intact (sham-operated) females, ovariectomized females, or intact (sham-operated) males. Male water consumption was elevated in all conditions involving social contact. Urinary creatinine was significantly lower in female-exposed males compared to isolated controls, while unadjusted testosterone was significantly lower in males in all social conditions. Again, creatinine-adjusted estradiol in female-exposed males significantly exceeded that of isolates. These data indicate that adult males drink and urinate more, have more dilute urine, and have a higher ratio of estradiol to creatinine when they are near developing females. These dynamics increase females' exposure to urinary steroids and other urinary constituents that can hasten sexual maturity.     

Does part reproductive history predict competitive interactions and male mating success in pupfish?

Two experiments were performed to determine the effect of past territoriality and mating success on subsequent dominance and ability to attract females. First, in 14 staged, pair-wise encounters, the development of nuptial coloration and agonistic behaviour was examined in two types of males: previously territorial (PT) and previously non-territorial (PNT) males. Previously territorial males developed a more intense nuptial coloration and won all contests. Second, to test the males' ability to attract females, other factors were controlled that could have affected female mate choice, such as male size, quality of breeding substrate and male-male interactions. In 40 trials, PT males were more active, developed a more intense nuptial coloration, courted females more vigorously, spawned sooner and had a higher mating success than PNT males. Thus, in pupfish, Cyprinodon pecosensis, previous reproductive history is a good indicator of both dominance and mating success. Only males in good physical condition achieve high mating success, and both intra- and inter-sexual selection are important in maintaining the close correlation between expression of nuptial coloration, a second sexual trait, and other male attributes, such as physical condition and vigour, that allow a male to secure and defend a territory against rivals and to attract and spawn with females.     

Factors limiting polygyny in the willow ptarmigan

Willow ptarmigan (Lagopus lagopus) usually pair monogamously and males invest more in vigilance and parental care than do males of other grouse species. I tested whether polygyny is rare because male parental care is necessary for successful reproduction. By continuous removal of males I skewed the operational sex ratio, induced polygyny, and then compared breeding success and survival of females that shared a mate and those that did not. Both groups had similar clutch sizes, laid eggs at the same time, produced young of the same weight, and fledged similar numbers of juveniles. Hens that shared a mate suffered higher losses of nests during incubation, and fewer returned to the breeding range in subsequent years. Thus, although male parental care was not essential, it did improve female reproductive success and survival. I suggest that willow ptarmigan are not polygynous because territorial females are able to prevent potential secondary hens from settling, thus securing unshared access to male investment and territorial resources.     

Stress-dependent trade-off Between Immunological Condition and Reproductive Performance in the Polygynous Red Bishop (<Emphasis Type="Italic">Euplectes orix</Emphasis>)

According to life-history theory, a condition-dependent trade-off between reproductive performance and immune function might be expected, with only superior individuals being able to allocate many resources into reproduction without having detrimental effects on their immune system. However, little attention has been paid to the possibility that annual variations in environmentally and socially induced stress levels modify this trade-off. We analysed haematological parameters and investigated the relationship between male immunological condition and reproductive performance as measured by reproductive effort and success in a colony of the highly polygynous red bishop (Euplectes orix) for five consecutive breeding seasons. Haematological parameters can be used to assess an individuals’ stress level, and especially the heterophile/lymphocyte ratio (H/L ratio) is known to increase in response to a wide variety of stressors. Relative basophile counts, heterophile counts, lymphocyte counts, absolute leukocyte counts and H/L ratios of territorial males varied significantly among seasons. Both relative heterophile counts and H/L ratios were significantly positive related to overall breeding activity within the colony as measured in terms of total number of nests accepted and total number of eggs laid in the study colony within a breeding season. The relationship between immunological condition (as assessed by relative lymphocyte and heterophil counts and the H/L ratio) and both male reproductive effort (as measured by the number of nests built by the territorial males) and reproductive success (as measured by the number of nests accepted) varied considerably among seasons, ranging from significant negative to significant positive correlations. Across seasons, we found a significant negative relationship between the within-season correlation coefficients of individual H/L ratios against reproductive performance and the average H/L ratio of all territorial males in that season. Thus, a positive association between the immunological condition of a male and his reproductive performance (indicated by a negative correlation between H/L ratio and reproductive performance) exists only in seasons with high average H/L ratios (i.e. high average stress levels). We conclude that the trade-off between immunological condition and male reproductive performance might be important only in seasons with an overall reduced level of male immunological condition due to environmentally and/or socially induced stress and immunosuppressive conditions.Co-ordinating editor: L.D. Hurst     

Social suppression of ovarian cyclicity in captive and wild colonies of naked mole-rats, Heterocephalus glaber

To investigate the endocrine cause of reproductive suppression in nonbreeding female naked mole-rats, animals from 35 colonies were studied in captivity. Urinary and plasma progesterone concentrations were elevated in pregnant females (urine: 10.0-148.4 ng/mg Cr, 27 samples from 8 females; plasma: 3.6-30.0 ng/ml, 5 samples from 5 females; Days 21-40 of pregnancy) and cyclic breeding females (urine: 0.5-97.8 ng/mg Cr, 146 samples from 7 females; plasma: less than 1.0-35.4 ng/ml, 25 samples from 7 females). The latter group showed cyclic patterns of urinary progesterone, indicating a mean ovarian cycle length of 34.4 +/- 1.6 days (mean +/- s.e.m.) with a follicular phase of 6.0 +/- 0.6 days and a luteal phase of 27.5 +/- 1.3 days (19 cycles from 9 breeding females). In non-breeding females urinary and plasma progesterone values were undetectable (urine: less than 0.5 ng/mg Cr, 232 samples from 64 females; plasma: less than 1.0 ng/ml, 7 samples from 6 females). Breeding females had higher (P less than 0.001) plasma LH concentrations (3.0 +/- 0.2 mi.u./ml, 73 samples from 24 females) than did non-breeding females (1.6 +/- 0.1 mi.u./ml, 57 samples from 44 females). Urinary and plasma progesterone concentrations in non-breeding females from wild colonies situated near Mtito Andei, Kenya, were either below the assay sensitivity limit (urine: less than 0.5 ng/mg Cr, 11 females from 2 colonies; plasma: less than 1.0 ng/ml, 25 females from 4 colonies), or very low (plasma: 1.6 +/- 0.6 ng/ml, 15 females from 4 colonies). In captivity, non-breeding females removed from their colonies (i.e. the dominant breeding female) and either paired directly with a non-breeding male (N = 2), or removed and housed singly for 6 weeks before pairing with a non-breeding male (N = 5) may develop a perforate vagina for the first time in as little as 7 days. Urinary progesterone concentrations rose above 2.0 ng/mg Cr (indicative of a luteal phase) for the first time 8.0 +/- 1.9 days after being separated. These results suggest that ovulation is suppressed in subordinate non-breeding female naked mole-rats in captive and wild colonies, and show that plasma LH concentrations are significantly lower in these non-breeding females. This reproductive block in non-breeding females is readily reversible if the social factors suppressing reproduction are removed.     

Effects of age and experience on the responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

The ability of sexually mature non‐territorial floaters to sire offspring affects the success of floating as a breeding strategy. Red‐winged Blackbirds (Agelaius phoeniceus) have second‐year (SY) and after‐second‐year (ASY) floater males, and genetic studies suggest that floaters may gain paternity. Despite these studies, we still know little about the fitness costs and benefits of floating in this species. By presenting taxidermic models of females in soliciting, precopulatory postures in territories of experienced (previously attracted at least one mate in the study area) and inexperienced (did not previously defend a territory in the study area) males, I was able to examine the copulation behavior and success of floater male Red‐winged Blackbirds as well as the effect of experience for territorial males. Floaters trespassed during 66.1% of presentations and 85.4% of trespassers were SY males. Experienced territorial males (92.5%) and neighbors (87.5%) were most successful in attempts to copulate with models, inexperienced territorial males (62.5%) and ASY floaters (50.0%) had intermediate success, and SY floaters (6.9%) were least successful. Experienced territorial males were more likely to approach models than inexperienced males, and floaters were more likely to approach models in territories of experienced than inexperienced males. These results provide further evidence that floaters trespass frequently, suggest that floaters sire offspring, and demonstrate that prior breeding experience affects the behavior and reproductive success of territorial male Red‐winged Blackbirds. Floating appears to be a conditional strategy for ASY male Red‐winged Blackbirds, but, because it is still not known if SY floaters sire offspring, these males may be trespassing to gain information or experience.     

Male Seminal Fluid Substances Affect Sperm Competition Success and Female Reproductive Behavior in a Seed Beetle

Male seminal fluid proteins are known to affect female reproductive behavior and physiology by reducing mating receptivity and by increasing egg production rates. Such substances are also though to increase the competitive fertilization success of males, but the empirical foundation for this tenet is restricted. Here, we examined the effects of injections of size-fractioned protein extracts from male reproductive organs on both male competitive fertilization success (i.e., P2 in double mating experiments) and female reproduction in the seed beetle Callosobruchus maculatus. We found that extracts of male seminal vesicles and ejaculatory ducts increased competitive fertilization success when males mated with females 1 day after the females’ initial mating, while extracts from accessory glands and testes increased competitive fertilization success when males mated with females 2 days after the females’ initial mating. Moreover, different size fractions of seminal fluid proteins had distinct and partly antagonistic effects on male competitive fertilization success. Collectively, our experiments show that several different seminal fluid proteins, deriving from different parts in the male reproductive tract and of different molecular weight, affect male competitive fertilization success in C. maculatus. Our results highlight the diverse effects of seminal fluid proteins and show that the function of such proteins can be contingent upon female mating status. We also document effects of different size fractions on female mating receptivity and egg laying rates, which can serve as a basis for future efforts to identify the molecular identity of seminal fluid proteins and their function in this model species.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Mate and territory availability affect breeding dispersal and divorce in a resident Southern House Wren Troglodytes aedon musculus population

Breeding dispersal is the movement of an individual between breeding attempts and is usually associated with the disruption of the social pair bond, although mates may disperse together as a social unit. In monogamous territorial species, the decision to disperse may be affected by individual attributes such as sex, age and condition of the disperser. However, environmental and social contexts may also play a crucial role in the decision to disperse. We analysed capture‐resighting data collected over 9 years to study breeding dispersal and divorce rates of a Southern House Wren Troglodytes aedon musculus population in South Temperate Argentina. Between‐season dispersal was more frequent than within‐season dispersal, with females dispersing more often than males, both between and within seasons. Both within‐season and between‐season breeding dispersal probability was affected by territory availability, but not by previous breeding success. When the adult sex ratio (ASR) was more skewed towards males, male between‐season dispersal was also affected by mating status, with widowed and single males dispersing more often than paired males. Within‐season divorce increased the reproductive success of females but not males, and was affected by the availability of social partners (with increasingly male‐skewed ASR). Our results suggest that territorial vacancies and mating opportunities affect dispersal and divorce rates in resident Southern House Wrens, highlighting the importance of social and environmental contexts for dispersal behaviour and the stability of social pair bonds.     

Differential grooming rate and tick load of territorial male and female impala, Aepyceros melampus

During the breeding season in Zimbabwe territorial male impalawere found to engage in much less self oral grooming and allogroomingthan females, presumably as a reflection of the need to remainvigilant in herding females and repulsing challenging bachelormales. Territorial males spent an average of 11 min engagedin all types of grooming during a 12-h day, compared with 40min grooming by females. Rutting activity and time spent scanningpeaked in May, while self oral grooming and feeding by territorialmales was lowest at this time. The decrement in grooming bymales (relative to females) represented half of all time devotedto rutting behaviors. Territorial males appeared to sacrificefeeding and grooming time in exchange for more time devotedto vigilant activities essential to mating success. In a comparableregion of Zimbabwe, territorial males were also found to harborabout six times as many adult ticks as the females on the sameterritories during the breeding season. Because grooming isunderstood to be effective in removing ticks, the higher tickload of territorial males was attributed to their reduced groomingbehavior. However, testosterone and adrenal cortical steroids,which are elevated in territorial male impala, are known todepress the immune system, and so may be important in controllingparasite infections. Sexually active males of many species aregenerally found to harbor more parasites than females. The differencein tick load between territorial male and female impala mayreflect both behavioral and hormonal parameters.     

Polygyny in the Willow Warbler Phylloscopus trochilus in Swedish Lapland

The breeding sex ratio of Willow Warblers Phylloscopus trochilus in a subalpine birch forest was strongly female biased. Very few territorial males remained unmated and most became polygynous. Bigyny was the rule, but some males probably had three females. Male assistance at secondary nests varied and was probably dependent on temporal distance between the females' nests. There was no significant reduction in reproductive output for these females. Variation in the sex ratio of breeding birds can be explained, at least partly, by variation in the timing of male settlement.     

Reproductive behavior of females and its relation to the mating success of two male forms inMnais pruinosa (Zygoptera: Calopterygidae)

The behavior of females of the damselflyMnais pruinosa was observed in the breeding season. Males consisted of both territorial (esakii) and non-territorial (strigata) types in the study area. Females sometimes arrived at a stream for mating and oviposition. The staying time of females in the stream was 1–3 hours, while that of males was 5–8 hours. The proportion of the males that copulated with the females did not differ between the two male forms present in the stream. In the Calopterygidae, of which the females usually perform multiple copulation, oviposition without subsequent recopulation is considered to be advantageous for a mated male in order to avoid the risk of sperm displacement. From such a viewpoint, the time spent for oviposition was measured for females that arrived at the stream after copulation with different male forms. The ratio was 69.1: 11.5–30.9 betweenesakii andstrigata males. Assuming that the oviposition time is proportional to the number of fertilized eggs laid, this ratio would represent the relative reproductive success of the two male forms. In fact, the relative abundance of the two male forms was also biased in favor ofesakii males (61.7:38.3) in this population. The mechanism of coexistence of the two male forms is discussed in relation to their reproductive success.     

Investment in territorial defence relates to recent reproductive success in common loons Gavia immer

As the value of a limited resource such as a territory increases, animals should invest more in the defence of that resource. Because reproductive success often depends on the quality of a breeding territory, reproductive success or failure may alter the perceived value of territory and affect an animal's investment in territorial defence. We used common loons Gavia immer to test the hypothesis that animals with recent breeding success would show stronger territorial defence than those with no recent breeding success. Surprisingly, successful loons responded less, not more, to a simulated intrusion. However, birds with success in the previous season also increased their territorial response as the breeding season progressed. In conjunction with past data showing that recently successful loons experience an increase in conspecific intrusions on their territories, we interpret our data to suggest that loons with recent success offset the cost of increased intrusions by adopting a more efficient strategy for territorial defence (e.g. limiting investment in resource defence until the time of the season when it is most critical).     

Indicators of habitat quality and the reproductive output of a forest songbird in small and large fragments

The distribution of individuals among habitats that vary in quality (i.e. resource availability) may affect reproductive output at a population level. I compared indicators of habitat quality including the breeding experience ratios (inexperienced:experienced birds), turnover rates, pairing success, and densities of a forest songbird, the Eastern Yellow Robin Eopsaltria australis , in two small, food-poor forest fragments, with those in two large, food-rich fragments. I then evaluated the likelihood that the breeding experience ratio affected the reproductive output of populations. Inexperienced males occurred in small fragments eight times as often as in large fragments. Male turnover rates were 1.5 times higher in the small than large fragments, and 20% (10/50) of the male population were unpaired in the small fragments compared to 0% (0/25) in the large. None of these measures differed significantly for females. Experienced birds of both sexes produced almost all of the offspring compared with inexperienced birds. Despite these findings, reproductive output did not vary with fragment size for robins. Thus, while breeding experience clearly influenced the reproductive success of individuals, there were no obvious population consequences of having disproportionately more inexperienced males in the small fragments. I conclude that while male traits may be good indicators of resource levels within fragments (specifically, food availability), they may not be adequate predictors of population performance. Thus, the reproductive output of populations must be measured directly before conclusions concerning population performance can be made. Interestingly, breeding densities were not accurate indicators of either resource levels or population performance. Densities were two times higher in the smaller fragments, and I suggest that this result reflects problems with male dispersal among fragments caused by isolation.     

Territory Area as a Determinant of Mating Systems

Territoriality is an integral part of breeding behavior in manyanimals. Because the reproductive success of territorial malesis often limited by access to females, breeding males shouldbehave as "area maximizers" when the basis of female choiceis either the abundance of resources within the territory orterritory area per se. Being reproductively more limited byenergy, territorial females should be "energy maximizers." Aseries of simple analytical models of territory area for suchforagers is developed to explore how changes in local food productionand/or local competitor density affect both the probabilityof a territorial male securing more than one mate (polygyny)and the probability of his and his mates' reproductive successincreasing. Two cases are modeled (only males territorial vs.both sexes territorial), each for various sets of assumptionsregarding interactions between food production, feeding efficiency,and competitor density. Concurrent responses in territory area,territory food reserves, net energy gain, and time budgetingprovide testable sets of predictions for each scenario. Whereonly males are territorial (Case I), changes in food productioncan have different (indeed, opposite) effects upon an individualmale's probability of becoming polygynous, depending upon whetherthe basis of female choice is the abundance of food within theterritory or another factor positively correlated with territoryarea. Increases in competitor density usually decrease the probabilityof polygyny regardless of the basis of female choice. Whereboth sexes are territorial and territories overlap intersexually(Case II), the mating system becomes a function of the numberof female territories within each male's territory, which varieswith the ratio of male to female territory areas. In this case,the probability of polygyny occurring will increase if foodproduction for both sexes increases without concurrent increasesin competitor density, and will decrease if competitor densityfor both sexes increases without concurrent increases in feedingefficiency. Few data are presently available to test eitherthese general predictions or numerous sets of secondary predictionstabulated in the text. Available evidence is largely consistentwith the models, but mostly circumstantial. This is becausethe predictions of these and other models of territory areaare strongly assumption dependent, and few published studieshave investigated these assumptions. These analyses demonstratethat to accurately assess the mechanisms by which environmentalfactors affect territory area, and thus mating systems, testsof the underlying assumptions of models are essential.     

Male reproductive strategy and the importance of maternal status in the Antarctic fur seal Arctocephalus gazella

Abstract. Although mammalian mating systems are classically characterized in terms of male competition and polygyny, it is becoming increasingly apparent that alternative male strategies and female choice may play important roles. For example, females who mate with males from a dominant dynasty risk producing inbred offspring. Many pinnipeds are highly polygynous, but in some species alternative male strategies such as aquatic mating appear to be important, even when behavioral observations suggest strong polygyny. Here, we analyze male reproductive success in the Antarctic fur seal Arctocephalus gazella , an otariid described behaviorally as being highly polygynous, by combining a microsatellite paternity analysis spanning seven consecutive breeding seasons with detailed behavioral data on both sexes. Territorial males fathered 59% of 660 pups analyzed from our study colony. Male reproductive skew was considerable, with a quarter of all paternities assigned to just 12 top individuals on a beach where mean annual pup production was 635. Most males were successful for only a single season, but those able to return over successive years enjoyed rapidly increasing success with each additional season of tenure. We found no evidence of alternative male reproductive tactics such as aquatic or sneaky terrestrial mating. However, paternity was strongly influenced by maternal status. Females observed on the beach without a pup were significantly less likely to conceive to a sampled territorial male than equivalent females that did pup. In addition, their pups carried combinations of paternal alleles that were less likely to be found on the study beach and exhibited lower levels of shared paternity. Thus, from a territorial male's perspective, not all females offer equal opportunities for fertilization.     

Modulation of the hypothalamic-pituitary-adrenal axis of an Arctic-breeding polygynandrous songbird,the Smith's longspur,Calcarius pictus

To successfully reproduce in the Arctic, birds must modulate their neuroendocrine and behavioural systems. These adjustments include an attenuation of the stress responsiveness of the hypothalamic-pituitary-adrenal (HPA) axis to external stimuli and a behavioural insensitivity to high corticosterone (B) levels. The HPA axis was examined in free-living territorial polygynandrous Smith's longspurs (Calcarius pictus) that migrate to breed on the Arctic tundra. Basal and stress-induced B levels were measured through the breeding season and were found to be significantly lower in females compared with males. This was not a consequence of adrenal insensitivity, because intrajugular injections of adrenocorticotrophin hormone (ACTH) enhanced B release in incubating females. In males the adrenocortical response to stress was significantly attenuated during the parental phase compared with arrival at the breeding ground. In contrast to temperate passerines, there was no significant decrease in male territorial aggressive behaviour when B was experimentally elevated, suggesting a behavioural insensitivity to glucocorticoids. This mechanism is hypothesized to increase reproductive success by preventing interruptions to parental care during transient deleterious environmental perturbations, which are often experienced in the short Arctic breeding season. Modulation of the HPA axis in this species in relation to life-history stage, lifetime reproductive success and the polygynandrous mating system is discussed.     

Compatibility drives female preference and reproductive success in the monogamous California mouse (Peromyscus californicus) more strongly than male testosterone measures

Female assessment of male attractiveness and how preferred qualities impact reproductive success is central to the study of mate choice. Male attractiveness may depend on traits beneficial to the reproductive success (RS) of any female, termed ‘universal quality’, and/or on behavioral and biological interactions between potential mates that reflect ‘compatibility’. The steroid hormone testosterone (T) often underlies male attractiveness in rodents and is associated with enhanced paternal care in the monogamous and biparental California mouse (Peromyscus californicus). We hypothesized that (1) T-characteristics are universally attractive to female California mice and that (2) if reproductive success is higher for females mated with preferred males, then females mated with males preferred by other females will also have higher reproductive success. Alternatively, we speculated that pair compatibility, based on emergent pair qualities, is important for a species with coordinated offspring care. We assessed individual T-characteristics in three ways: (1) T-response to GnRH challenges (2) baseline T-level and (3) T-response to a female. Testosterone-response did not predict female preference, but females spent more time investigating males with higher baseline T (accounting for only 9.6% of the variation in investigation time). None of the T-measures was associated with RS. Females paired with males they preferred produced litters more quickly and had higher RS than females paired with their non-preferred males. Naïve females who did not undergo preference tests had equivalent RS regardless of whether their mate was preferred or non-preferred by another female. These data suggest that higher male T elicits investigation, but female preference in the California mouse is more strongly linked with compatibility because individual preference was a better predictor of RS than any T measure.     

The effects of the minimum threshold condition for breeding on offspring sex-ratio adjustment in the lesser kestrel

We propose a model for sex-ratio adjustment complementary to that of Trivers and Willard. In addition to the three basic assumptions of the Trivers-Willard model, our model assumes that the sex with more variable reproductive success (normally male) is also the sex less constrained for reproduction. This assumption seems realistic, because several studies have demonstrated that poor-condition males may adopt alternative mating strategies and sire some offspring, whereas females have physiological constraints for gestation or egg production that cannot be avoided. Thus, under these circumstances, sons of both poor and good condition would be more valuable for parents than daughters, whereas daughters would be relatively more valuable than sons at intermediate condition. This model predicts, therefore, a U-shaped relationship between parental condition and offspring sex ratio. We present a case study for the monogamous lesser kestrel (Falco naumanni) that fulfills the assumptions and predictions of the model. The minimum body condition for breeding, measured as pectoral thickness, was lower for sons than for daughters. Below this minimum, males had a higher chance of breeding than females. Above this minimum, however, the lifetime reproductive success was condition dependent in males but not in females. Thus, males in better body condition attain, on average, higher reproductive success than females. Offspring sex ratio varied with the size of the father's ornaments and mother condition according to the U-shaped pattern predicted by the model.