Seasonal changes in components of male and female reproductive success in Raphanus sativus L. (Brassicaceae)

Summary To document seasonal changes in the reproductive behavior of the perfect-flowered, self-incompatible mustard, Raphanus sativus L., we monitored individual survival, flower and fruit production among 58 individuals in a California population over six census dates (cohorts). Population size declined dramatically and mean individual levels of fruit set changed significantly between cohorts. The frequency distribution of flower and fruit production became increasingly skewed over the first four cohorts. The phenotypic maleness of individuals, a standardized measure of phenotypic gender, oscillated during the reproductive season, peaking in the third and fourth cohorts. We calculated a simple estimate of expected male reproductive success of each plant (the number of fruits sired on conspecifics); this estimate was a function of an individual's flower production and the fruit production of its potential mates in our sampled population. Mean expected male success did not differ significantly among cohorts; expected male success per flower did, however, change significantly among cohorts. Among individuals within each cohort, maternal fruit production and expected male success were both positively correlated with flower production throughout the season. Spearman rank correlation coefficients indicate that the strength of these associations, however, changed during the season. Linear regressions of transformed variables indicated that the shape of several fitness functions also changed over time. In addition, the amount of variation in maternal or expected paternal success explained by flower production declined over the first four cohorts. If typical of wild populations, these temporal changes in these functions suggest that measurements of the intensity of phenotypic selection on flower production will depend on when and how fitness is measured in natural populations.     

Density and reproductive success in wild populations of Diplotaxis erucoides (Brassicaceae)

Summary One possible consequence of low population density, particularly in self-incompatible plants, is reproductive failure. I surveyed seed set per flower in two populations of the self-incompatible annual Diplotaxis erucoides (Brassicaceae) in Jerusalem, Israel. Widely spaced plants had lower fruit set and fewer seeds per filled silique than did plants growing close to conspecific neighbors. Such density-dependent reproductive success could help explain the maintanence of spatial patchiness in plant populations, and could also have implications for population dynamics of rare species.     

Inter-locality variation in reproductive success of the citril finch Serinus citrinella

Habitat quality is generally thought to affect breeding success. We tested this effect comparing differences in clutch size and reproductive success between citril finch Serinus citrinella sub-populations closely located (<5 km) but differing in habitat quality, within the Port del Comte mountain, in the Catalonian Pre-Pyrenees. We found that birds in the low quality area (Bofia) showed significantly lower hatching, breeding and nesting success than finches in the high quality area (Vansa). These differences in reproductive success fit well with recently found differences in citril finch body mass, fat score, diet, survival rate and speed of moult between these two localities.     

Limitations on reproductive success in endemic <Emphasis Type="Italic">Aquilegia viscosa</Emphasis> (Ranunculaceae) relative to its widespread congener <Emphasis Type="Italic">Aquilegia vulgaris</Emphasis>: the interplay of herbivory and pollination

Plant reproduction can be strongly affected by herbivory and different features of pollination ecology, such as pollinator visitation rates and capacity for self-pollination. The purpose of this study is to compare the relative impact of herbivory and pollination on maternal reproductive success in endemic Aquilegia viscosa and its widespread congener Aquilegia vulgaris. We conducted herbivore exclusion experiments in two populations of each species in 2 different years and showed that the maternal fertility of A. viscosa was significantly more limited by floral predation and pre-dispersal seed predation than its widespread congener. In the absence of herbivory, A. viscosa retained significantly lower maternal fertility than A. vulgaris. Experimental pollinations in an insect-free glasshouse showed that the two species have an equal seed/ovule ratio both in the absence of pollinators and in the presence of non-limiting outcross pollination. Pollinator visitation rates were significantly higher in populations of A. vulgaris than in populations of A. viscosa. In addition, path analyses showed that spur length, an important trait for pollinator attraction in Aquilegia, and, indirectly sepal and petal width, contribute positively to the seed/ovule ratio in A. vulgaris, but not in A. viscosa. These results indicate that maternal fertility of endemic A. viscosa is strongly reduced by flower and seed predation despite low rates of pollinator visitation, and that pollen or resource limitation in the wild may further reduce maternal fertility. Finally, floral trait variation appears to be decoupled from fertility variation in endemic A. viscosa, which possibly constrains the evolution of reproductive traits in this species.     

Age-related and individual differences of reproductive success in male and female barbary macaques (Macaca sylvanus)

Age-related and individual differences in longterm reproductive success were analyzed in two social groups of free-ranging Barbary macaques. Maternity data were obtained from continuous birth records and paternity was determined with oligonucleotide-fingerprinting. The fathers of 246 of 286 investigated individuals could be identified. They were born during a 14-year period and represented 73 and 34% of all known offspring from the females of the study groups B/F and C, respectively. Only these infants were considered when comparing male reproductive success with that of females. The necessary adjustment of the female data resulted in small deviations from the true values in one group, but substantially increased individual differences in female fertility in the second group. Subadult males, 4.5 – 6.5 yrs old, had a much lower reproductive success than adult males (7.5 – 25 yrs old) and same-aged females. Reproductive success of adult males was not significantly affected by age, while females invariably ceased reproduction during the first half of the third decade of life. Males were more likely than females to leave no offspring, unless they survived 9 – 10 yrs of age. The number of years with breeding opportunities was important for male reproductive success but less significant than that for females. Reproductive success of several males during the 14-year study period was similar to or even exceeded that possible for a female in her whole lifetime. Variance of male reproductive success significantly exceeded that of females in both study groups.     

A 15-year study of the association between dominance rank and reproductive success of male rhesus macaques

The reproductive success (RS) of 32 males in a captive group of rhesus macaques (Macaca mulatta) between 1978 and 1992 was determined using paternity exclusion analysis. Dominance rank of each male over age 4 was assessed at the end of each breeding season based on agonistic dyadic interactions. The dominance rank and RS of these males were strongly correlated whether or not subadult males were included. The high reproductive success of males that eventually reached alpha rank is primarily responsible for this outcome. These results support the theory that social dominance has evolved in genusMacaca by sexual selection but some changes in male dominance rank and RS during the 15-year period suggest that priority of access is not the sole focus for such selection.     

Experimentally induced clutch size enlargements affect reproductive success in the Pied Flycatcher

We performed a food provisioning experiment in a population of Pied Flycatchers Ficedula hypoleuca breeding at high altitude in central Spain to test if food availability before and during laying determines clutch size. Food was provided to one of two pairs with the same date of initiation of nest-building (15 dyads of subsequently reproducing pairs were thus created). Food provisioning began on the day of initiation of nest-building and ended on the day after the last egg was laid. Although laying date was unaffected by the experiment, clutch size in the experimental treatment was significantly larger. This result could indicate that food availability at laying (1) proximately constrained clutch size or (2) that females evaluated future conditions for incubating eggs and feeding nestlings based on food availability at laying. Reproductive success (proportion of eggs that resulted in fledged young) was significantly reduced in the experimental treatment. This effect suggest that supplemented females were tricked by the experiment into laying more eggs than the number of eggs they were able to incubate with success and the number of nestlings they were able to feed, a source of error in clutch size adjustment which could be common in non-experimental situations.     

Malarial parasites decrease reproductive success: an experimental study in a passerine bird

Malarial parasites are supposed to have strong negative fitness consequences for their hosts, but relatively little evidence supports this claim due to the difficulty of experimentally testing this. We experimentally reduced levels of infection with the blood parasite Haemoproteus prognei in its host the house martin Delichon urbica, by randomly treating adults with primaquine or a control treatment. Treated birds had significantly fewer parasites than controls. The primaquine treatment increased clutch size by 18%; hatching was 39% higher and fledging 42% higher. There were no effects of treatment on quality of offspring, measured in terms of tarsus length, body mass, haematocrit or T-cell-mediated immune response. These findings demonstrate that malarial parasites can have dramatic effects on clutch size and other demographic variables, potentially influencing the evolution of clutch size, but also the population dynamics of heavily infected populations of birds.     

Plant size,breeding system,and limits to reproductive success in two sister species of Ferocactus (Cactaceae)

Plant reproductive output can be limited by a variety of factors, bothintrinsic and extrinsic. I investigated the reproductive biologies of twospecies of unbranched short-columnar cacti, Ferocactuscylindraceus and F. wislizeni. I recordedfemalereproductive output (flowers produced, fruit set, seeds per fruit and seedmass), plant size and growth, and used hand-pollination experiments todeterminebreeding systems and pollen limitation. In both species, the ability to selfvaried among individuals, but self-pollination resulted in very few seeds,suggesting strong inbreeding depression. Neither species was pollen-limited.Numbers of flowers produced increases with plant size for both species, andseeds per fruit may also be related to plant size, although the relationship isunclear. Seed mass is not correlated with plant size. Flower production wassimilar in both species, but F. cylindraceus producedfewerseeds per fruit than F. wislizeni, and its seeds weighedless. Fruit set by F. cylindraceus was heavily impacted bya florivorous lepidopteran. Fruit set was very high (94 to 96%) inF. wislizeni, suggesting that architectural constraints(e.g., meristem limitation) are more limiting than resource levels or the levelof pollinator services. In F. cylindraceus, numbers ofseeds per fruit was positively correlated with seed mass, whereas inF. wislizeni, the relationship was negative (tradeoff).Thegrowth rates of F. wislizeni are affected by rainfall theprevious season, and growth rates increase as the plant ages.Ferocactus cylindraceus and F.wislizeni are thought to be sister species, meaning that observeddifferences between them are more likely to be the result of recentevolutionaryprocesses in their lineages rather than differing phylogenetic histories.     

Reproductive success and individual variation in feeding frequency of House Sparrows (Passer domesticus)

Parental care is assumed to be closely associated with individual differences in reproductive success. We investigated how feeding frequencies varied among parents and how this affected the subsequent reproductive success in insular populations of House Sparrows Passer domesticus in northern Norway. Female parents fed their offspring more than male parents did, and the feeding rates were positively related to the feeding rates of the partner. A positive relationship between feeding rates and bill depth was present in females. In males, the feeding rates were negatively related to total badge size and positively related to visible badge size, after the effect of other variables had been taken into account. A non-linear convex relationship between feeding frequency and hatch day was present in males, which could reflect either the seasonal change in weather conditions or the seasonal variation in food availability. For both sexes, feeding frequencies increased with increasing brood size, but at the same time the average feeding rate per nestling decreased with increased brood sizes. Finally, our results indicate that the amount of parental investment, measured as feeding rates during the nestling stage, may have a positive long-term influence both on the number of fledglings that recruit as well as the probability that fledglings survive until recruitment.     

The relationship of the female reproductive success of Eucalyptus globulus to the endogenous properties of the flower

Low capsule and seed set is a major factor limiting seed production in Eucalyptus globulus seed orchards. Controlled pollination studies showed that the reproductive success (number of seeds produced per flower pollinated) was primarily determined by the female. We aimed to identify the factors contributing to the differences in reproductive success between female genotypes in terms of the physical and anatomical properties of the flower. We studied pairs of genotypes of high and low reproductive success from each of three races (Furneaux Group, Strzelecki Ranges and Western Otways) growing in a seed orchard. Controlled pollinations were performed on six females and along with flower physical measurements, pollen tube growth and seed set were assessed. Overall tree reproductive success was positively correlated with flower size, ovule numbers, style size, cross-sectional area of conductive tissue within the style (all of which were inter-correlated) and the proportion of pollen tubes reaching the bottom of the style. Significant positive correlations of reproductive success and flower physical properties between different ramets of the same genotypes across seasons suggests a genetic basis to the variation observed. The majority of pollen tube attrition occurred within the first millimetre of the cut style and appeared to be associated with differences in style physiology. When examined as pairs within races the difference in reproductive success for the Western Otways pair was simply explained by differences in flower size and the number of ovules per flower. Physical features did not differ significantly for the Strzelecki Ranges pair, but the proportion of pollen tubes reaching the bottom of the style was lower in the less reproductively successful genotype, suggesting an endogenous physiological constraint to pollen tube growth. The difference in reproductive success between the females from the Furneaux Group was associated with a combination of these factors.     

A trematode infection with no effect on reproductive success of a sand-bubbler crab

The infection effects of the parasitic digenean trematode on the body weight and reproductive success of the sand-bubbler crab were examined. Gynaecotyla squatarolae (Trematoda: Microphallidae) infects the body cavity of Scopimera globosa (Decapoda: Scopimeridae) and uses the crab as its second intermediate host. The parasites infected all reproductive crabs examined to varying degrees. Larger crabs of both sexes had more parasites than smaller ones, probably because body size reflects age, and older crabs had a longer period of exposure to infection. Males had more parasites than females, probably because of sexual difference in acting time on the surface. Ovigerous females stay in closed burrows and do not act on the surface during incubation, and so have less chance of infection than males. The quantity of infecting parasites did not explain variations in either body weight or reproductive success of individual crabs in a field experiment. The life history of this parasite, relative body size of the crabs, and cost and the possible benefit of manipulation for the parasite may explain these results.     

Male dominance rank and reproductive success in primate groups

The relationship between male dominance rank and reproductive success has been a topic of interest since the beginning of primatology. From a theoretical point of view the existence or absence of this relationship has great implications with respect to the meaning of dominance rank and more general of the social relationships between individuals within social groups. Until fairly recently mating behaviour has been used as an indicator of reproductive success, but these two variables need not be correlated. The relations between mating success, reproductive success, and dominance rank indicate whether selective mating is involved (different mating partners at different phases of fertility, for example through male contest or female choice). With the development of genetic techniques to determine reproductive success directly, it has now become possible to investigate these relations. In our study on wild long-tailed macaques (Macaca fascicularis) we find a relatively strong correlation between rank and reproductive success which is attributable to selective mating by the alpha male during fertile periods of the females. In most previous studies no such clear results have been obtained, and we discuss the differences in outcome in relation to the study conditions, group sizes, and possible differences between species in terms of reproductive strategies.     

Molecular phylogeny ofForsythia (Oleaceae) based on chloroplast DNA variation

Phylogenetic relationships of ten wild species and several cultivars ofForsythia were reconstructed based on the chloroplast (cp) DNA variation. A total of 216 cpDNA variants, 44 of which were potentially phylogenetically informative, was detected using 24 restriction endonucleases. Phylogenetic analysis usingFontanesia andAbeliophyllum as outgroups revealed four well defined species groups in the genus: 1)F. suspensa, 2)F. europaea — F. giraldiana, 3)F. ovata — F. japonica — F. viridissima, and 4)F. koreana — F. manshurica — F. saxatilis. The amount of support for each monophyletic group was evaluated by various methods including character number, decay analysis, parsimony bootstrapping, Neighbour-Joining (NJ) — bootstrapping, NJ-jackknifing, and the topology-dependent permutation tail probability (T-PTP) test. The data do not support the hybrid origin ofF. intermedia fromF. suspensa andF. viridissima. The disjunctly distributed European species,F. europaea, was identified as a sister species of the ChineseF. giraldiana and it was probably derived through recent long distance dispersal.     

Consequences of variation in flowering phenology for seed head herbivory and reproductive success in Erigeron glaucus (Compositae)

Summary We examined the relationship between flowering phenology, reproductive success (seed production only), and seed head herbivory for 20 similarly sized clones of Erigeron glaucus growing at Bodega Bay Reserve, northern California, USA. Although clones tended to reach peak flowering on the same date, they differed in the proportion of their total flowers produced around that date (flowering synchrony). Clones also differed in the number and density of flower heads presented at any one time to pollinators and herbivores (floral display). Both of these characteristics had consequences for herbivory and plant reproductive success. The proportion of flower heads damaged by insect herbivores was greater for clones that concentrated flowering activity during the main flowering period for the population as a whole (high synchrony) compared to clones that spread flowering out temporally. The primary reason for this result was that clones with low flowering synchrony produced a significant proportion of their flower heads during the fall and therefore, escaped attack by the tephritid fly, Tephritis ovatipennis. Clones with intermediate synchrony had lower seed success (total number of viable seeds produced over the year) than clones with either low or high synchrony. The proportion of flower heads damaged by insect herbivores and number of tephritid flies reared from flower heads were both negatively correlated to floral display while seed head mass and germination rates were positively related to display. Thus, clones which produced dense floral displays were favored both in terms of reduced herbivory and increased successful seed production.     

The influence of body size on immediate reproductive success in Dugesia gonocephala (Tricladida,Paludicola)

The individual reproductive output of the stream-dwelling flatworm Dugesia gonocephala was investigated. Various measures of reproductive success were related to body size. (I) For the first 30 days in the laboratory small individuals produced no cocoons, individuals of intermediate size produced unfertilized cocoons and large individuals usually produced fertilized cocoons. (II) In individuals that produced a cocoon, no correlation was found between the number of cocoons produced in one month and body size. (III) Large individuals, however, produced larger cocoons. This was not due to the fact that unfertilized cocoons were smaller. (IV) Large cocoons tended to contain more young. (V) The average size of young hatching from large cocoons was larger. (VI) Large individuals produced their first cocoon soon after their arrival in the laboratory and seemed to have a higher chance of producing a fertilized first cocoon. (VII) A trade-off existed between producing many small versus few large young.     

Male reproductive success in the domestic cat (Felis catus L.): A case history

The domestic cat shows a great variability in different life history traits like its social organization, ranging from a solitary life to living in large social groups depending on environmental conditions. Until now, the mating system has not been shown to vary between populations and was described as promiscuous. Here we present data on the reproductive success of a male, which clearly show that monopolization of females by males is possible in this species.     

Male dominance and genetically determined reproductive success in the mandrill (Mandrillus sphinx)

Darwin referred to the adult male mandrill (Mandrillus sphinx) as the most brightly coloured of all mammals, citing the brilliant red and blue pigmentation of the face, rump, and genitalia as extreme examples of evolution by sexual selection. Considerable controversy exists concerning possible effects of sexually selected phenotypes via intermale competition on reproductive success. Behavioural and genetic studies of a large, semi-free ranging mandrill colony in Gabon have now demonstrated that clear-cut relationships exist between male secondary sexual development, social dominance, copulatory behaviour, and reproductive success in this primate species. Two morphological variants of adult male were identified; “fatted” males, with maximum secondary sexual coloration, which occupied dominant positions in the social group, and “non-fatted” males, with muted secondary sexual adornments, smaller testes and lower plasma testosterone levels, which lived as peripheral/solitary individuals. DNA fingerprinting analyses on infants born over five successive years showed that only the two most dominant, fatted males in the group had fathered offspring. Throughout the annual mating season these males attempted to mate-guard and copulate with females during periods of maximal sexual skin tumescence. Male rank and mating success were strongly positively related and the alpha male sired 80 – 100% of the resulting offspring during three consecutive years. Non-fatted adult males and group associated subadult males engaged in infrequent, opportunistic matings and did not guard females. Loss of alpha status resulted in a fall in reproductive success, but the effect was gradual; the deposed alpha male continued to father 67% and 25% of infants born during the next two years. Thus, whilst claims that male dominance determines mating success and paternity in primates have caused considerable debate, these results on mandrills provide unequivocal evidence for the existence of such effects.     

Determinants of male mating success in a large group of Barbary macaques (Macaca sylvanus) at Affenberg Salem

Male mating activities in relation to the likelihood of ovulation and conception were studied in a large group of semifree-ranging Barbary macaques (Macaca sylvanus) during two successive mating seasons. In both mating seasons, adult males attained a significantly higher mating success than subadult males, and they monopolized high ranking females more effectively than low ranking females during the period when conception was most probable. Also, in both mating seasons male rank was significantly correlated with male mating success if all sexually mature males were included. Nevertheless, mating success was not a linear function of age or rank. In both mating seasons mating success of 5-year-old males was much higher than that of dominant but peripheralized 6- and 7-year-old males. Moreover, a significant correlation between rank and measures of mating success among adult males was found in the second but not in the first mating season. The results indicate that mating and, most probably, reproductive success of male Barbary macaques is dependent on the male's social position in the group, which is defined not only by the outcome of dyadic agonistic encounters but also by the ability to get a central position in the group, and on the stability of rank relations.     

The presence of heterospecific pollen on stigmas of nectariferous and nectarless orchids and its consequences for their reproductive success

Summary The widespread occurrence of nonorchid, heterospecific pollen grains on the stigmatic surfaces of a range of nectariferous and nectarless European orchids (Dactylorhiza. Orchis, Goodyera, andGymnadenia species) is reported for the first time, and the impact of heterospecific pollination on orchid reproductive success is experimentally investigated. There are three main components of stigmatic contamination by heterospecific pollen: the frequency of contamination, the diversity of foreign species present on the stigma, and the amount of pollen deposited. Six out of seven of the species examined have greater than 85% of stigmas contaminated with wind and insect-dispersed pollen. From one to nine insect-dispersed foreign pollen species are present per stigma, including pollen of members of the families Apiaceae, Asteraceae, Caryohpyllaceae, Ericaceae, and Primulaceae. Average loads per stigma vary from 13 to 176 grains, with individual stigma loads ranging from one to 909. Whether or not the orchid provides nectar has a major impact on these three components. Nectarless orchids have the greatest frequencies of contamination, diversity of species, and average load per stigma. Insect-dispersed pollen is deposited both by pollinators and visitors but, in spite of low levels of pollination, nectarless orchids still exhibit higher frequencies of heterospecific pollen contamination. The effect of the presence of heterospecific pollen on the reproductive success of orchids is tested in this study for the first time. Average-to-high, naturally occurring loads of heterospecific pollen derived from a mixture ofArmeria maritima,Caltha palustris,Cochlearia officinalis,Cytisus scoparius, andPrimula vulgaris and consisting of 50–250 grains per load are placed onto stigmas ofDactylorhiza purpurella which are subsequently self-pollinated with half of a pollinium. All pollinations produce capsules indicating that heterospecific pollen does not affect fruit set. Although experimental and control fruits are similar in size, they differ in total seed weight and composition. Total seed weight is reduced and the proportion of seeds with normal embryos decreased while the proportion of unfertilised ovules increased following pollination with heterospecific pollen, indicating a detrimental effect on fertilisation. Lower reproductive success caused by fertilisation failure is likely to be most severe in nectarless species because of their generally higher levels of contaminated stigmas. As nectarless orchids are known to have lower fruit set compared with nectariferous ones, this finding suggests that the reproductive success of nectarless orchids may be even lower than previously realised.Abbreviations RS reproductive success