Body Condition Indices Predict Reproductive Success but Not Survival in a Sedentary,Tropical Bird

Body condition may predict individual fitness because those in better condition have more resources to allocate towards improving their fitness. However, the hypothesis that condition indices are meaningful proxies for fitness has been questioned. Here, we ask if intraspecific variation in condition indices predicts annual reproductive success and survival. We monitored a population of Neochmia phaeton (crimson finch), a sedentary, tropical passerine, for reproductive success and survival over four breeding seasons, and sampled them for commonly used condition indices: mass adjusted for body size, muscle and fat scores, packed cell volume, hemoglobin concentration, total plasma protein, and heterophil to lymphocyte ratio. Our study population is well suited for this research because individuals forage in common areas and do not hold territories such that variation in condition between individuals is not confounded by differences in habitat quality. Furthermore, we controlled for factors that are known to impact condition indices in our study population (e.g., breeding stage) such that we assessed individual condition relative to others in the same context. Condition indices that reflect energy reserves predicted both the probability of an individual fledging young and the number of young produced that survived to independence, but only during some years. Those that were relatively heavy for their body size produced about three times more independent young compared to light individuals. That energy reserves are a meaningful predictor of reproductive success in a sedentary passerine supports the idea that energy reserves are at least sometimes predictors of fitness. However, hematological indices failed to predict reproductive success and none of the indices predicted survival. Therefore, some but not all condition indices may be informative, but because we found that most indices did not predict any component of fitness, we question the ubiquitous interpretation of condition indices as surrogates for individual quality and fitness.     

Lifetime Reproductive Success in the Solitary Endoparasitoid, Venturia canescens

Parasitoid wasps have long been considered excellent organisms in studies examining the evolution of reproductive and life-history strategies. In examining the lifetime reproductive success of parasitoids in the laboratory, most investigations have provided the insects with excess hosts and food, where they exist in a relatively constraint-free environment. Importantly, these conditions may not accurately reflect the true heterogeneity of natural systems, where suitable hosts and food sources are likely to be limiting. This study examines the influence of differences in host and food availability on reproductive and life-history parameters in an asexual strain of the solitary endoparasitoid, Venturia canescens (Hymenoptera: Ichneumonidae). Lifetime reproductive success in V. canescens was measured in response to temporal variations in host and food (honey solution) access. Cohorts of parasitoids were provided with 200 fifth-instar larvae of the Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae), and food for variable periods daily after eclosion. V. canescens is synovigenic, and host-deprived wasps continued to mature eggs over the first few days after eclosion until the egg storage capacity was reached in the oviducts. When these wasps were subsequently provided with hosts, oogenesis resumed and continued until later in adult life. Constantly fed wasps lived longer and produced more progeny than wasps from cohorts which were alternately fed and starved or were starved from eclosion. Moreover, wasps with constant host and food access produced most progeny early in life and usually experienced prolonged periods of postreproductive survival. In contrast, the reproductive period of wasps with limited host access was more evenly distributed throughout the adult life. Consequently, the cumulative progeny production by V. canescens with constant food access was fairly uniform irrespective of host availability. Longevity and fecundity in V. canescens were positively correlated with adult size. However, variable host access had little effect on the longevity of wasps which were constantly supplied with honey. Over the first 2 days of adult life, variation in food access also had no effect on progeny production by V. canescens. We argue that manipulating temporal host and food access to parasitoids in the laboratory more closely approximates natural conditions, where these resources are likely to be spatially separated. Moreover, our findings suggest that many highly synovigenic parasitoids like V. canescens, which produce microtype (=hydropic) eggs, have a considerably higher reproductive potential than ovary dissections have revealed. Our findings are discussed in relation to life-history evolution in the parasitic Hymenoptera.     

Boldness and Reproductive Fitness Correlates in the Eastern Mosquitofish, Gambusia holbrooki

Animals are known to exhibit 'personality'; that is, individual differences in behaviour that are consistent across time and/or situations. One axis of personality of particular importance for behavioural ecology is boldness, which can be defined as the tendency of an individual to take risks. The relationship between individual personality and fitness correlates, particularly those involved in reproduction, remains largely unexplored. In the current study, we used female Eastern mosquitofish ( Gambusia holbrooki ) to ascertain whether certain reproductive traits (e.g. stage of pregnancy, fecundity) are correlated with individual personality in two wild populations in New South Wales, Australia. To quantitatively assess this relationship, we tested individual fish for their level of boldness, as measured by their latency to exit a refuge and tendency to shoal in a novel environment. We also quantified individual differences in general activity and tendency to swim near the water surface and substratum. For both populations taken together, bolder individuals tended to be smaller, relatively less fecund (when taking body size into account), and spent more time near the water surface than near the substratum compared with timid individuals. Individual boldness was not correlated with either general activity or stage of pregnancy. To our knowledge, our study characterizes for the first time a relationship between an individual personality trait (boldness) and a reproductive fitness correlate (fecundity) in fishes.     

Wealth, Status, and Reproductive Success among the Mukogodo of Kenya

The evolutionary biological hypothesis that culturally defined values and goals are proximate means of enhancing reproductive success is tested on data from the Mukogodo, a small group of Maa-speaking pastoralists in north-central Kenya who value the accumulation of livestock. The results support the prediction that, at least among males, livestock wealth should correlate with reproductive success. This correlation appears to be due mainly to greater polygyny among wealthier men. Lower age at first marriage among wealthier men may also contribute to the correlation between livestock wealth and reproductive success. The association between livestock wealth and reproductive success does not appear to be due to the productivity of wives and children, to bridewealths obtained when daughters marry, or to the effects of wealth on the reproductive success of men's wives.     

Psychophysiological Correlates of Athletic Success in Athletes Training for the Olympics

Long-term studies related to comprehensive analysis of psychophysiological correlates of the athletic potential were performed on the basis of innovative technologies of molecular genetic and biological energy analyses using modern automated software-hardware complexes. The results were used to select indices for personalized diagnosis of the athletic potential, to analyze it comparatively at the team level, and to calculate an athletic rating correlating with athletic success. These indices make it possible to detect preclinical health disturbances, development of energy deficit, and overtraining. The approaches may be used to analyze the general well-being of a population.__________Translated from Fiziologiya Cheloveka, Vol. 31, No. 3, 2005, pp. 84–92.Original Russian Text Copyright (c) 2005 by Bundzen, K. Korotkov, A. Korotkova, Mukhin, Priyatkin.     

Effect of Breed,Cage Type,and Reproductive Phase on Fecal Corticosterone Levels in Doe Rabbits

Fecal corticosterone concentration (FCC) is increasingly being used as a noninvasive indicator of stress in assessment of nonhuman animal welfare. The aim of this study was to evaluate effects of breed, cage type, reproductive phase, and their interactions on FCC levels in doe rabbits. A total of 252 doe rabbits were randomly assigned to 2 groups. Does were individually housed in either standard dimension cages (SC) or in cages with a volume more than double that of the SC. Bigger cages (BC) were equipped with a plastic foot mat. Breed, cage type, and reproductive phase significantly affected FCC. New Zealand hybrids showed higher FCCs (p < .001) when compared witho the autochthonous breed (27.77 ± 0.47 vs. 24.61 ± 0.36 pg g^?1, respectively). Increased cage size coupled with a plastic foot mat resulted in a significant decrease in doe FCC. The highest FCCs were detected at partum (BC: 30.40 ± 0.81 pg g^?1; SC: 33.36 ± 0.86 pg g^?1; p ≤ .05), followed by postweaning (BC: 25.09 ± 0.95 pg g^?1; SC: 27.63 ± 0.95 pg g^?1; p ≤ .05). These results support the hypothesis that measurement of FCC provides a useful indicator of chronic stress in doe rabbits.     

Positive Carotenoid Balance Correlates with Greater Reproductive Performance in a Wild Bird

Background

Carotenoids can confer somatic and reproductive benefits, but most evidence is from captive animal experimentation or single time-point sampling. Another perhaps more informative means by which to assess physiological contributions to animal performance is by tracking an individual''s ability to increase or sustain carotenoids or other health-related molecules over time, as these are likely to be temporally variable.

Methodology/Principal Findings

In a field study of North American barn swallows (Hirundo rustica erythrogaster), we analyzed within-individual changes in carotenoid concentrations by repeatedly sampling the carotenoid profiles of individuals over the course of the breeding season. Our results demonstrate that carotenoid concentrations of individuals are temporally dynamic and that season-long balance of these molecules, rather than single time-point samples, predict reproductive performance. This was true even when controlling for two important variables associated with reproductive outcomes: (1) timing of breeding and (2) sexually selected plumage coloration, which is itself positively correlated with and concomitantly changes with circulating carotenoid concentrations.

Conclusions/Significance

While reproduction itself is purported to impose health stress on organisms, these data suggest that free-ranging, high-quality individuals can mitigate such costs, by one or several genetic, environmental (diet), or physiological mechanisms. Moreover, the temporal variations in both health-linked physiological measures and morphological traits we uncover here merit further examination in other species, especially when goals include the estimation of signal information content or the costs of trait expression.     

Reproductive Success and Body Size in the Cricket Gryllus firmus

Male body size influences mate choice and sexual selection in many animal species. Here we investigate the role of male body size in the reproductive success of the field cricket Gryllus firmus. This species hybridizes with a close smaller relative, G. pennsylvanicus, and it is thought that this size difference may affect reproductive isolation between these species. We paired large and small G. firmus males with a single G. firmus female and genotyped the resulting offspring. Overall, larger males sired a greater proportion of offspring and in a majority of the crosses the larger male sired all of the offspring. For crosses in which both males sired offspring, there was no difference in the proportion of offspring sired by small and large males. Intrasexual competition, female choice, and differences in ejaculates between males could all influence the patterns we observe. We discuss the implications of our findings within the context of reproductive isolation between G. firmus and G. pennsylvanicus.     

Physically Challenging Song Traits,Male Quality,and Reproductive Success in House Wrens

Physically challenging signals are likely to honestly indicate signaler quality. In trilled bird song two physically challenging parameters are vocal deviation (the speed of sound frequency modulation) and trill consistency (how precisely syllables are repeated). As predicted, in several species, they correlate with male quality, are preferred by females, and/or function in male-male signaling. Species may experience different selective pressures on their songs, however; for instance, there may be opposing selection between song complexity and song performance difficulty, such that in species where song complexity is strongly selected, there may not be strong selection on performance-based traits. I tested whether vocal deviation and trill consistency are signals of male quality in house wrens (Troglodytes aedon), a species with complex song structure. Males’ singing ability did not correlate with male quality, except that older males sang with higher trill consistency, and males with more consistent trills responded more aggressively to playback (although a previous study found no effect of stimulus trill consistency on males’ responses to playback). Males singing more challenging songs did not gain in polygyny, extra-pair paternity, or annual reproductive success. Moreover, none of the standard male quality measures I investigated correlated with mating or reproductive success. I conclude that vocal deviation and trill consistency do not signal male quality in this species.     

Somatic Donor Cell Type Correlates with Embryonic,but Not Extra-Embryonic,Gene Expression in Postimplantation Cloned Embryos

The great majority of embryos generated by somatic cell nuclear transfer (SCNT) display defined abnormal phenotypes after implantation, such as an increased likelihood of death and abnormal placentation. To gain better insight into the underlying mechanisms, we analyzed genome-wide gene expression profiles of day 6.5 postimplantation mouse embryos cloned from three different cell types (cumulus cells, neonatal Sertoli cells and fibroblasts). The embryos retrieved from the uteri were separated into embryonic (epiblast) and extraembryonic (extraembryonic ectoderm and ectoplacental cone) tissues and were subjected to gene microarray analysis. Genotype- and sex-matched embryos produced by in vitro fertilization were used as controls. Principal component analysis revealed that whereas the gene expression patterns in the embryonic tissues varied according to the donor cell type, those in extraembryonic tissues were relatively consistent across all groups. Within each group, the embryonic tissues had more differentially expressed genes (DEGs) (>2-fold vs. controls) than did the extraembryonic tissues (P<1.0×10^–26). In the embryonic tissues, one of the common abnormalities was upregulation of Dlk1, a paternally imprinted gene. This might be a potential cause of the occasional placenta-only conceptuses seen in SCNT-generated mouse embryos (1–5% per embryos transferred in our laboratory), because dysregulation of the same gene is known to cause developmental failure of embryos derived from induced pluripotent stem cells. There were also some DEGs in the extraembryonic tissues, which might explain the poor development of SCNT-derived placentas at early stages. These findings suggest that SCNT affects the embryonic and extraembryonic development differentially and might cause further deterioration in the embryonic lineage in a donor cell-specific manner. This could explain donor cell-dependent variations in cloning efficiency using SCNT.     

Stages of Age-Related Reproductive Success in Birds: Simultaneous Effects of Age, Pair-Bond Duration and Reproductive Experience

SYNOPSIS. Most organisms reproduce episodically rather thancontinuously. Forms of discontinuity may include seasonal breedingin annual cycles, and delayed breeding and/or senescence overthe lifespan of individuals. In addition, age can have profoundeffects on the annual reproductive success (RS) of a wide varietyof animals. Explanations of the effects of age on RS can beconfounded, however, by the acquired breeding experience ofindividuals, and the duration of pair-bonds, both of which haveeffects on RS similar to those of age itself. However, moststudies of the effects of age on RS prior to senescence havenot accounted for the effects of pair-bond duration, and manyhave treated age and experience as synonyms. In this paper Isummarize the effects of all three factors, discuss the simultaneousand interrelated nature of their effects, and suggest ways toseparate the relative contributions of each.     

Lek Social Organization and Reproductive Success in the Greater Prairie Chicken

Between 1964 and 1969, studies were made of Prairie Chickenreproductive behavior on a lek with a stable social organization.The social organization of this lek was altered by the removalof dominant males; three in 1970 and two in 1971. Social organizationand reproductive behavior were observed on this lek prior toand following removal of the dominant males. Removal of dominantmales disrupted the stable social organization on the lek. Totalnumber of aggressive interactions between males increased followingremoval of dominant males.No change was noted in the total numberof females visiting the lek in 1970 compared with the previous6-year period. Likewise, no change was noted in the number ofattempted copulations observed in 1970 compared with previousyears. However, the number of successful copulations was drasticallyreduced (90% fewer) during 1970 because of increased male-maleaggressive interactions on the lek. During 1971, fewer territorialmales were on the lek compared to previous years, and fewerfemales visited the lek and fewer successful copulations wererecorded than in previous years.Late nests of Prairie Chickensare less successful and contain fewer eggs compared with nestsbegun earlier. In some instances, the female social hierarchymay serve to delay onset of nesting, therefore reducing totalreproduction for the year.     

Hurricane Disturbance,Plant-Animal Interactions,and the Reproductive Success of a Tropical Shrub1

Hurricane disturbance may have strong effects on plant-animal interactions important in plant reproductive success. Components of reproductive success (flowering, pollination, seed predation) in the tropical shrub Ardisia escallonioides (Myrsinaceae) were examined from 1991–1994 in four southern Florida populations. Hurricane Andrew struck three of the four populations on 24 August 1992. Hurricane Andrew delayed flowering by two months in 1992. In 1993 and 1994, the three hurricane-damaged populations had increased flowering and inflorescence production compared to 1991 and 1992, while the undamaged population had no flowering. Hurricane disturbance had different effects on generalist versus specialist plant-animal interactions. Species composition and relative abundance of the generalist pollinator community that visits A. escallonioides was similar before and after the hurricane, indicating little effect of the disturbance on this interaction. In contrast, populations of a specialist flower galling moth (Periploca sp., Cosmopterigidae) declined in 1992 following Hurricane Andrew. Although moth populations increased at two of the three sites in 1993, the relative impact of moth predation on seed production was low due to extensive flower production. One moth population suffered local population extirpation for two years, reestablishing itself in November 1994. Hurricane disturbance resulted in a window of opportunity for massive seed production of Ardisia escallonioides in south Florida. Total seed production in 1993 increased twelve to seventy-three times the 1992 levels. Total seed production declined in 1994, but remained high compared to prehurricane levels.     

Male Incubation and its Effect on Reproductive Success in the Black Swan,Cygnus atratus

Male incubation is very unusual among Anatidae. In the black swan, males spend more time incubating the clutch than females do. We investigated the incubation patterns of males and females in a breeding colony in Vienna over a one-year period. Breeding occurred year round, with a peak in winter. Inattentive periods during incubation were shorter in winter than at other times of the year, probably because the heat loss of eggs is then higher. We tested the importance of the costs of incubation by prolonging the breeding period experimentally. Eggs were exchanged for dummies, which were readily accepted and incubated for up to 342 days. The higher the incubation effort of females, measured as the product of their proportion of incubation and the total length of the incubation phase, the longer was the interval between the end of incubation and laying of a subsequent clutch. This suggests that the incubation effort of females is limiting the reproductive output of a pair. By taking charge of the greater part of incubation, males may increase the productivity of their female partners and thereby their own reproductive success. This amplification of female fecundity is a factor favouring a high share in male incubation which is discussed in polyandrous species (e.g. Jenni 1974) but generally overlooked in monogamous systems. The ultimate cause for male incubation in black swans may lie in the combination of a monogamous mating system and the capability of this species to breed whenever the environmental conditions turn favourable, at any time of the year.     

Evidence for the Cost of Reproduction in Humans: High Lifetime Reproductive Effort Is Associated with Greater Oxidative Stress in Post-Menopausal Women

Life history theory predicts trade-offs between reproductive effort and maternal survivorship in energy-restricted environments. However, empirical evidence for the positive association between maternal mortality and reproductive effort from energetically challenged human populations are mixed and physiological mechanisms that may underlie this association are poorly understood. We hypothesized that increases in aerobic metabolism during repeated periods of pregnancy and lactation result in increased oxidative stress that may contribute to somatic deterioration, vulnerability to illness, and accelerated aging. We therefore predicted that lifetime gravidity and parity would be related to levels of biomarkers of oxidative stress, as well as antioxidative defence enzymes in post-menopausal women. Our hypothesis was supported by positive linear associations between levels of 8-OHdG, a biomarker of DNA oxidative damage (β = 0.21, p<0.05), levels of antioxidative defence enzyme Cu-Zn SOD (β = 0.25, p<0.05), and number of lifetime pregnancies. Furthermore, independent of age and health status, post-menopausal women with higher gravidity and parity (> = 4 pregnancies per lifetime) had 20% higher levels of 8-OHdG and 60% higher levels of Cu-Zn SOD compared to women with lower gravidity and parity (<4 pregnancies per lifetime). Our results present the first evidence for oxidative stress as a possible cost of reproductive effort in humans.     

Body Size Correlates with Fertilization Success but not Gonad Size in Grass Goby Territorial Males

In fish species with alternative male mating tactics, sperm competition typically occurs when small males that are unsuccessful in direct contests steal fertilization opportunities from large dominant males. In the grass goby Zosterisessor ophiocephalus, large territorial males defend and court females from nest sites, while small sneaker males obtain matings by sneaking into nests. Parentage assignment of 688 eggs from 8 different nests sampled in the 2003–2004 breeding season revealed a high level of sperm competition. Fertilization success of territorial males was very high but in all nests sneakers also contributed to the progeny. In territorial males, fertilization success correlated positively with male body size. Gonadal investment was explored in a sample of 126 grass gobies collected during the period 1995–1996 in the same area (61 territorial males and 65 sneakers). Correlation between body weight and testis weight was positive and significant for sneaker males, while correlation was virtually equal to zero in territorial males. That body size in territorial males is correlated with fertilization success but not gonad size suggests that males allocate much more energy into growth and relatively little into sperm production once the needed size to become territorial is attained. The increased paternity of larger territorial males might be due to a more effective defense of the nest in comparison with smaller territorial males.     

When Money Is Not Enough: Awareness,Success, and Variability in Motor Learning

When performing a skill such as throwing a dart, many different combinations of joint motions suffice to hit the target. The motor system adapts rapidly to reduce bias in the desired outcome (i.e., the first-order moment of the error); however, the essence of skill is to produce movements with less variability (i.e., to reduce the second-order moment). It is easy to see how feedback about success or failure could sculpt performance to achieve this aim. However, it is unclear whether the dimensions responsible for success or failure need to be known explicitly by the subjects, or whether learning can proceed without explicit awareness of the movement parameters that need to change. Here, we designed a redundant, two-dimensional reaching task in which we could selectively manipulate task success and the variability of action outcomes, whilst also manipulating awareness of the dimension along which performance could be improved. Variability was manipulated either by amplifying natural errors, leaving the correlation between the executed movement and the visual feedback intact, or by adding extrinsic noise, decorrelating movement and feedback. We found that explicit, binary, feedback about success or failure was only sufficient for learning when participants were aware of the dimension along which motor behavior had to change. Without such awareness, learning was only present when extrinsic noise was added to the feedback, but not when task success or variability was manipulated in isolation; learning was also much slower. Our results highlight the importance of conscious awareness of the relevant dimension during motor learning, and suggest that higher-order moments of outcome signals are likely to play a significant role in skill learning in complex tasks.     

Limitations to Reproductive Success in the Dioecious Tree Rhamnus davurica

The reproductive success of a female plant in a dioecious species may be affected by pollen limitation and resource limitation. This study presents evidence that the reproductive success of the dioecious understorey tree species, Rhamnus davurica, is affected by the distance to the nearest male. The sex ratios were female-biased, although showing fluctuations in the three years of conducting the study. The mortality rate of females was higher than that of males indicating a trade-off between reproduction and survival. Altogether 49 females, designated as “focal females”, were randomly selected for monitoring their reproductive status between April and October in 2010. But successful reproduction (meaning that the flowering female trees had fruit in the fruiting season) was observed only in 28 females in 2011 and 16 females in 2012. The method of path analysis was applied to determine the effect of topography, local competition and proximity to the nearest male on the fruit set of the females. In the three years of the study, elevation, competition and female size had no significant effect on the fruit set. The distance to the nearest male, however, had a significant effect on fruit set. Number of fruits and fruit set were decreased with increasing distance to the nearest male. It was possible to estimate maximum fruit set, based on the comparatively large dataset. The number of fruits and the fruit set are exponentially related to the distance to the nearest male and the relationships are described by an exponential model. The results of this study support the importance of pollen limitation on the reproductive success in Rhamnus davurica.     

Eggshell Appearance Does Not Signal Maternal Corticosterone Exposure in Japanese Quail: An Experimental Study with Brown-Spotted Eggs

Reproduction is a critical period for birds as they have to cope with many stressful events. One consequence of an acute exposure to stress is the release of corticosterone, the avian stress hormone. Prolonged stress can have negative impacts on the immune system, resulting in, for example, increased oxidative stress. Through maternal effects, females are known to modulate their investment in eggs content according to their own physiological condition. Less is known about maternal investment in eggshells, especially in pigments. The two main eggshell pigments may possess opposite antioxidant properties: protoporphyrin (brown) is a pro-oxidant, whereas biliverdin (blue-green) is an antioxidant. In Japanese quail, we know that the deposition of both pigments is related to female body condition. Thus, a chronic stress response may be reflected in eggshell coloration. Using female Japanese quails that lay brown-spotted eggs, we explored whether physiological exposure to corticosterone induces a change in female basal stress and antioxidant factors, and eggshell pigment concentration, spectrophotometric reflectance, and maculation coverage. We supplemented adult females over a 2 week period with either peanut oil (control) or corticosterone (treatment). We collected pre- and post-supplementation eggs and analysed the effect of corticosterone treatment on female physiology and eggshell appearance parameters. Except for corticosterone-fed birds which laid eggs with brighter spots, supplementation had no significant effect on female physiology or eggshell pigment concentration, reflectance and maculation. The change in eggshell spot brightness was not detected by a photoreceptor noise-limited color opponent model of avian visual perception. Our data confirms that eggshell reflectance in spotted eggs varies over the laying sequence, and spot reflectance may be a key factor that is affected by females CORT exposure, even if the changes are not detected by an avian visual model.     

Reproductive Success Associated with Territoriality, Sneaking, and Grouping in Male Rose Bitterlings, Rhodeus ocellatus (Pisces: Cyprinidae)

The spawning success of male rose bitterlings, Rhodeus ocellatus, adopting an alternative reproductive style, was estimated through behavioural data and electrophoretic paternal analyses in field observations and experiments. Three mating patterns were observed: territoriality, sneaking, and grouping. Mating patterns depended on a male's relative size and on local male density (the number of males around a spawning spot: a mussel). Spawning patterns (pair spawning, pair spawning with sneaker, and group spawning) varied with local male density. Time-budget data of the territorial males indicated a trade-off between chasing and courtship behaviour as local male density changed. Females deposited appoximately only 1 egg per egg-laying into the mussels. As a result of isozyme analysis, a minimum of 12% (two out of 17) of the offspring in the sample were found to have been fathered by sneaker males in pair spawning with sneaker. I scored through behavioural data the mating success per spawning for each pattern, on an individual basis. The average reproductive success per spawning for each pattern was: territorial (0.61), sneaking (0.31) and grouping (0.11), and thus the successes of the patterns were not equal. Accordingly, the alternative reproductive styles of male rose bitterlings are best interpreted as alternative phenotypes in a conditional behaviour.