The effect of mating on female reproduction across hermaphroditic freshwater snails

Sexual conflicts often arise between mating partners because each sex tries to maximize its own reproductive success. One major male strategy to influence a partner's resource allocation is the transfer of accessory gland proteins. This has been shown to occur in simultaneous hermaphrodites as well as in organisms with separate sexes. Although accessory gland proteins affect the investment of resources in both male and female function, we here specifically focus on female investment. In the great pond snail, Lymnaea stagnalis, previous studies found that the accessory gland protein ovipostatin reduced female fecundity by suppressing egg laying in the partner in the short term (days). To investigate whether this reduction in egg laying is a commonly found effect of mating in freshwater snails, we compared egg output for evidence of suppression in isolated and paired snails of eight pulmonate species. Furthermore, we determined whether the suppression of egg laying caused a shift in resource allocation to the eggs. We found that in five of the eight species egg laying was suppressed, with fewer and lighter egg masses being laid when they had access to a mating partner. In mated pairs of L. stagnalis and Biomphalaria alexandrina, allocation of resources to the eggs was altered in opposite ways: individuals of L. stagnalis laid fewer but larger and heavier eggs; individuals of B. alexandrina laid smaller and lighter eggs, with no change in egg numbers. Such changes in the female function are most likely the result of combined effects of receiving accessory gland proteins, and the cost of mating in both male and female roles. Thus, effects of the maternal environment, including the receipt of accessory gland proteins, on offspring investment are not restricted to species with separate sexes.     

Laying order and offspring sex in blue tits Parus caeruleus

According to sex allocation theory, females may benefit from a differential investment in sons and daughters, where fitness returns from male and female offspring vary. One way in which investment may be differentiated is by assigning male and female offspring to eggs of different sizes, and/or eggs laid early or late in the laying order. Here, we investigate whether such sex-specific adjustments occur in blue tits Parus caeruleus . We found that the proportion of males changed non-linearly with laying order showing the increase in early-laid eggs. Egg size increased with laying order, but this pattern differed between female and male eggs. Female eggs increased initially in size and became smaller later in the laying sequence, while male eggs exhibited constant increase in mass. However, egg size did not differ between genders. Egg size and laying order was found to interact in moulding sex of the embryo. Thus, females seem to simultaneously adjust the sex of the offspring in relation to their investments into eggs and the position of the egg in the laying sequence.     

Maternal investment during egg laying and offspring sex: an experimental study of zebra finches

To study whether offspring sex is related to the amount of resources invested in eggs we performed an experiment on zebra finches, Taeniopygia guttata. By manipulating their food supply, we forced two groups of females either to increase or to decrease investment in subsequent eggs. Since zebra finches are sexually dimorphic and the reproductive value of the sexes may vary with maternal nutritional status, we predicted that females would adjust the sex of their offspring to egg quality. Females that received poor-quality food for 7 weeks before egg laying, then food supplemented with proteins after they laid the first egg, significantly increased the mass of subsequent eggs. An increase of egg mass with laying sequence was less pronounced in females that received high-quality food before laying and experienced food deterioration after starting to lay. The proportion of sons in subsequent eggs tended to increase in the latter group (although this was marginally significant) but was not related to laying sequence in the other group: these patterns differed significantly between the groups. Offspring sex was not related to egg mass, but newly hatched male chicks were heavier than female chicks. Furthermore, the hatching success of male eggs was lower than that of female eggs. We suggest that differential hatching success of the sexes and sex differences in mass at hatching may constitute important factors shaping brood sex ratios. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sexual dimorphism of egg size in the European Blackbird Turdus merula

Recently, a number of studies have shown that female birds are able to control the sex of their progeny at the stage of the gamete. There is also some evidence that females adjust their investment in offspring depending on the sex of the embryo during egg formation. Differential maternal investment to the eggs depending on their sex is usually interpreted as an adaptive strategy, by which females can increase competitive abilities of the smaller sex, or preferentially invest towards the sex with the potentially higher fitness returns. Here, we studied variation in egg size in relation to embryo sex and laying order in the European Blackbird Turdus merula. We found male and female eggs to differ in size, with larger eggs containing male embryos, as well as a significant interaction between embryo sex and laying order. This interaction resulted from the fact that egg size increased with the laying sequence among eggs bearing females but did not change with laying order among eggs bearing males. There was no relationship between offspring sex and the laying sequence within a clutch. We suggest that sexual dimorphism in egg size recorded in the European Blackbird may reflect favouritism of the sex which may give higher fitness returns.     

Factors associated with the mass of eggs laid by tree swallows Tachycineta bicolor varied yearly between 2008 and 2016 in Michigan

The initial investment a female bird makes in an offspring is her production of its egg. The size of this investment can have important evolutionary consequences for both the female and her offspring. Between 2008 and 2016, we weighed 1977 tree swallow eggs freshly laid in nest boxes in west Michigan to investigate factors associated with egg mass. Breeding female age class was not associated with egg mass. Overall, eggs that hatched weighed significantly more than those that did not. An egg's mass was not associated with clutch size but increased with its position in the laying sequence. Females that bred multiple times showed high repeatability of egg mass. Mother–daughter narrow sense heritability of egg mass was low. There were statistically significant associations, both positive and negative, between female morphological characteristics and egg mass in some years but not others. Similarly, relationships between egg mass and weather conditions during egg formation were statistically significant in some years but not others. In summary, female characteristics and weather conditions were associated with tree swallow egg mass, but their influences varied yearly. These results reinforce the importance of long‐term studies for identifying the causes of selection that shape life‐history characteristics.     

Harem size and oviposition behaviour in a polygynous bark beetle

Abstract. 1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within‐harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis. 2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean (±SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females. 3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly. 4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egg‐laying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.     

The role of female investment in a sexual arms race

Forced copulations are common among waterfowl species (Anatidae), a group with relatively large eggs and very precocial young. They can be viewed as the outcome of an overt sexual conflict, and an evolutionary arms race between the sexes. We examined the female choice hypothesis suggested, but not properly tested by Briskie and Montgomerie (1997) and Montgomerie and Briskie (2007), that penis size in male ducks might correlate with female investment in eggs, predicting that in species where females lay larger eggs, penis size might be larger, because females would be more reluctant to abandon their eggs if forced to copulate. A larger data set than in earlier studies enabled us to test that hypothesis in a comparative way. Our results compelled us to reject the female choice hypothesis since egg size is negatively correlated with penis length and the number of vaginal spirals, both being seen as adaptations to frequent forced copulations. The apparent trade-off between egg size and morphological defences (vaginal spirals) is strong particularly among monogamous species. Overall, we conclude that factors that set a lower limit for egg size constrain the morphological defences of females and the arms race between the sexes in waterfowl.     

Individual quality and food availability determine yolk and egg mass and egg composition in tree swallows Tachycineta bicolor

Variation in egg size and composition can have important consequences for the quality of offspring. We investigated the factors influencing the yolk mass and egg mass of tree swallows breeding in Ithaca, NY. Using a nondestructive technique to estimate yolk mass via standardized digital-candler photographs, we compared yolk size and egg size of tree swallows Tachycineta bicolor in response to variation in food availability and individual quality. Insect availability one to three days prior to laying, but not four to six days, predicted yolk mass, while insect availability two to three days prior to laying predicted total egg mass. This suggests that, while eggs are formed over longer periods, food availability closer to time of laying has the greatest influence on egg size. These results were supported by collected eggs, as yolk rings revealed that tree swallow eggs are formed over 5–6 days. There was an influence of female quality as well, with early laying birds, independent of food availability, laying larger eggs. Eggs laid later in the laying sequence had larger yolks and greater egg mass. Overall, variation in egg quality appears to be due to a combination of environmental conditions, reflected in food resources, individual quality, and allocation tradeoffs during the laying period.     

Two eggs,two different constraints: a potential explanation for the puzzling intraclutch egg size dimorphism in Eudyptes penguins

Phenotypic plasticity and phenotypic stability are major components of the adaptive evolution of organisms to environmental variation. The invariant two-egg clutch size of Eudyptes penguins has recently been proposed to be a unique example of a maladaptive phenotypic stability, while their egg mass is a plastic trait. We tested whether this phenotypic plasticity during reproduction might result from constraints imposed by migration (migratory carry-over effect) and breeding (due to the depletion of female body reserves). For the first time, we examined whether these constraints differ between eggs within clutches and between egg components (yolk and albumen). The interval between colony return and clutch initiation positively influenced the yolk mass, the albumen mass, and the subsequent total egg mass of first-laid eggs. This time interval had only a slight negative influence on the yolk mass of second-laid eggs and no influence on their albumen and subsequent total masses. For both eggs, female body mass at laying positively influenced albumen and total egg masses. Female investment into the entire clutch was not related to the time in the colony before laying but increased with female body mass. These novel results suggest that the unique intraclutch egg size dimorphism exhibited in Eudyptes penguins, with first-laid eggs being consistently smaller than second-laid eggs, might be due to a combination of constraints: a migratory carry-over effect on the first-laid egg and a body reserve depletion effect on the second-laid egg. Both these constraints might explain why the timing of reproduction, especially egg formation, is narrow in migratory capital breeders.     

Calculating egg volume when shape differs: when are equations appropriate?

ABSTRACT Egg volume is often calculated from length and breadth, assuming little variation in egg shape within species. However, egg shape may vary among females, over time, or within clutches. If this variation is not predictable, volume should be estimated using another method. For example, in some species, egg shape changes consistently with laying order and volume can be estimated using laying‐order specific equations. We measured the length, breadth, volume, and mass of 249 Magellanic Penguin (Spheniscus magellanicus) eggs and regressed volume on length and breadth for first and second eggs separately. Differences between measured and calculated volume averaged <1% for both eggs. Using linear measurements, we calculated egg volume for 7085 clutches from 1983 to 2006. First and second eggs had similar calculated volumes (P > 0.05) in 19 of 24 yr; when they differed, the second egg was larger. In contrast, using single equations typically employed to compute egg volume, we erroneously concluded that first eggs were usually (20 of 24 yr) significantly larger than second eggs. Our regression equations should be applicable to other Spheniscus penguins because ratios of elongation coefficients (length divided by breadth) of first and second eggs are similar among these species. In other species of birds where egg shape changes with laying order, the volume of a sample of eggs should be measured to develop regression equations specific to the species and laying order. Slight variation in egg volume can have important evolutionary and ecological implications that would be erroneously interpreted without field tests to measure volume.     

A comparative study of egg mass and clutch size in the Anseriformes

The factors explaining interspecific differences in clutch investment in precocial birds are poorly understood. We investigated how variations in clutch characteristics are related to environmental factors in a comparative study of 151 extant species of ducks, geese and swans (Anseriformes). Egg mass was negatively related to clutch size in a phylogenetic regression, a relationship that was much stronger when controlling for female mass. Nest placement was related to both egg size and clutch size, with cavity-nesting species laying more but smaller eggs. Egg size was positively correlated with incubation period and with female mass, and also with sexual size dimorphism (i.e. male mass relative to that of the female). Clutch size was not related to female mass. Species with long term pair bonds laid smaller clutches and larger eggs. The size of the breeding range was strongly positively correlated with clutch size and clutch mass, and its inclusion in multivariate models made other biogeographical variables (hemisphere, breeding latitude or insularity) non-significant. The small clutches in insular species appear to be a product of small range size rather than insularity per se. Our results suggest there is an evolutionary trade-off between clutch and egg size, and lend support to Lack’s resource-limitation hypothesis for the waterfowl.     

Effect of population bottlenecks on the egg morphology of introduced birds in New Zealand

Hatching failure is inversely correlated with population bottleneck size among exotic birds introduced to New Zealand, but the mechanism for this is unclear. We assess whether the bottlenecks these species experienced during their introduction have affected hatching failure through deleterious changes in egg morphology. We measured egg size and shape of 13 species that passed through bottlenecks of 11–800 individuals during their establishment in the 19th century. Eggs were also measured in the source populations ('pre-bottleneck') of each species to compare pre- and post-bottleneck egg morphology directly. Significant changes in egg volume were found in six of 13 species, with most laying smaller eggs in New Zealand. Egg shape changed in four of 13 species but there was no directional bias; two species developed more elongated eggs and two species broader eggs. There was no relationship between bottleneck size and change in egg volume, but species passing through severe bottlenecks had greater variability in egg volume and were more likely to have eggs that deviated in shape from their source populations. There was no relationship between changes in either egg volume or shape and rates of hatching failure. Further work is needed to assess whether changes in egg morphology have negative consequences on offspring fitness and whether the observed changes are the result of differing environmental conditions in the introduced range.     

Reproductive variation and adult size in two co-occurring grasshopper species

ABSTRACT. 1. Egg sizes and clutch sizes of the grasshoppers Chorthippus brunneus (Thunb.) and Myrmeleotettix maculatus (Thunb.) were compared among three years and among three sites less than 1.3 km apart. Relationships between these reproductive traits and date of egg laying, body size and body condition were sought.
2. M.maculatus , the smaller species, laid fewer but larger eggs; and only the eggs of this species showed significant differences between sites and years.
3. A negative correlation between egg size and number per clutch was evident between species and years, but generally not among sites and among individuals of a population.
4. However, a hidden negative correlation between egg size and number was uncovered within populations when the relationship was examined for females of a given mature weight.
5. Variation in the number of eggs per clutch was explained statistically by a positive relationship between female body weight and egg number. Also, both interpopulation and intrapopulation comparisons revealed that for M.maculatus , but not for C.brunneus , females with long hind femurs laid large eggs.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

产卵次序对山麻雀卵色、卵大小和重量的影响

产卵次序可对鸟类卵黄中的营养成分、卵壳色素以及卵壳厚度等产生影响。一些鸟类可通过产卵次序对不同卵的资源进行分配,从而实现繁殖成效的最大化。本研究对不同产卵次序下山麻雀(Passer cinnamomeus)的卵色、卵大小和卵重量进行分析,结果表明,山麻雀的卵重和卵大小随着产卵次序并无显著变化,卵色与卵的大小和重量均无显著相关性,但卵背景的色度随产卵次序有明显下降趋势,而卵斑点密度则呈相反的变化。卵背景的色度随产卵次序下降可能是山麻雀有限的色素在卵中不均匀分配的结果,而卵斑点密度的增加则可能是对后期卵壳厚度变薄的一种补偿。     

Maturation increases early reproductive investment in Adélie Penguins Pygoscelis adeliae

Correlations between female investment in egg production and age, breeding experience and laying date have been reported in several seabird species. In general, clutch and egg sizes increase with female age and breeding experience but decrease with laying date. Positive correlations of clutch and egg size with age and breeding experience can be caused by an increase in reproductive investment with maturation or they may be an artefact of lower survival rates for individuals with poor-quality phenotypes. Negative correlations of clutch and egg size with laying date might signal an adaptive reduction in egg production or be due in part to variation among individuals. We examined the interactions of female age, breeding experience, laying date and clutch and egg size in Adélie Penguins Pygoscelis adeliae . Breeding experience strongly affected clutch size with 87.3% of all one-egg clutches laid by first-time breeders. In addition, increasing age had a positive influence on egg size and was associated with earlier laying dates. However, there was little evidence to suggest that either clutch or egg sizes are influenced by laying date. Laying dates and clutch and egg sizes did not affect a female's probability of returning to breed in the following year, indicating that increased investment is a product of maturation and not of the loss of poor-quality breeders from the population. Our results suggest that as female Adélie Penguins gain foraging and breeding experience they are able to initiate breeding earlier, to lay complete clutches of two eggs and to lay larger eggs.     

Geographical variation in reproductive traits and trade‐offs between size and number of eggs in the king ratsnake,Elaphe carinata

We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.     

Do Male Pied Flycatchers (Ficedula hypoleuca) Adjust Their Feeding Effort According to Egg Colour?

The colour of blue‐green bird eggs has been hypothesized to signal female quality to attending males, who may adjust their level of investment in the brood accordingly. The hypothesis has gained support in studies of Spanish pied flycatchers Ficedula hypoleuca. We performed a cross‐fostering experiment in a Norwegian population of pied flycatchers to provide an independent test of the sexually selected egg colour hypothesis in this species. Egg colour was not significantly correlated with estimates of female quality (clutch size, average egg volume, first egg laying date and feeding rate). There was a significant decrease in chroma and increase in brightness and egg volume during the laying sequence, with some marked differences between six‐egg and seven‐egg clutches that might reflect differences in female quality. However, we found no significant influence of egg colour (foster or original clutch) on male feeding rate or offspring viability (hatching success, average mass and fledging success) and no significant difference in feeding rate for males with six‐egg and seven‐egg foster or original clutches. We conclude that Norwegian male pied flycatchers do not use egg colour as a cue to female quality, thus questioning the generality of previous support for the sexually selected egg colour hypothesis from this species.     

SEXUAL SIZE DIMORPHISM AND EGG-SIZE ALLOMETRY IN BIRDS

We tested the hypothesis that egg size should evolve in sexually dimorphic birds to reduce costs associated with more rapid growth by nestlings of the larger sex. Consistent with this hypothesis, we found that in species in which males were larger, females laid proportionately larger eggs as sexual size dimorphism increased. However, this result was also consistent with the hypothesis that egg size varied allometrically with both male and female body size. Furthermore we found that in species in which females were larger, relative egg size decreased as size dimorphism increased, which is consistent with the “allometry hypothesis” but not the “cost-reduction hypothesis. That male body size contributes to the allometric relationship between egg size and body size suggests that the basis for the allometric relationship is not wholly a mechanical one stemming from the physical requirements of developing, transporting, and laying an egg of a particular size. Rather, the relationship seems likely to be tied more directly to body size itself the tact that male body size influences a female trait suggests that egg size–body size relationships otter some scope for investigating the basis for allometric relationships in general.     

Egg development time in the Zebra Finch Taeniopygia guttata varies with laying order and clutch size

As female birds are able to lay no more than a single egg each day, in those species producing larger clutches the first laid eggs may get a developmental head‐start over later eggs in the clutch. All other things being equal, the differential pattern of development across the clutch may contribute to hatching asynchrony and subsequent inequity in the competition between brood mates, and ultimately increase variance in the quality and fitness of first‐ and last‐laid offspring. It has been suggested that females might allocate resources differently across the laying sequence to moderate the developmental rate and hatching time of different embryos. We tested this theory in the Zebra Finch Taeniopygia guttata, a common model species for investigating maternal effects in birds. We removed 758 eggs from 160 nests shortly after they were laid and used artificial incubators to control for parental effects and monitor hatching times. Eggs from larger clutches consistently hatched sooner than those from average‐sized clutches, demonstrating that the intrinsic properties of an egg can alter the developmental time of embryos. There were also differences in the development time of eggs across the laying sequence, but these patterns were weaker, inconsistent and unrelated to sequential investment across the laying sequence in a straightforward way. This study indicates that maternal resource allocation to eggs across the laying sequence and across clutch sizes can influence development times and play a potentially important role in determining the competitive dynamics of broods.