Maternal investment in eggs is affected by male feet colour and breeding conditions in the blue-footed booby, <Emphasis Type="Italic">Sula nebouxii</Emphasis>

Females are expected to vary investment in offspring according to variables that may influence the offspring fitness in a way that optimises her inclusive fitness for a particular context. Thus, when sexual ornaments signal the quality of the male, females might invest in reproduction as a function of the attractiveness of their mate. We tested whether breeding conditions and male feet colour influence reproductive decisions of blue-footed booby females. In the blue-footed booby, male feet colour is a dynamic condition-dependent sexually selected trait that is related to paternal effort. During two consecutive years, an El Niño year (poor breeding conditions) and a year with good breeding conditions, we experimentally reduced male attractiveness by modifying their feet colour after the first egg was laid and recorded female investment in the second egg. We found that, relative to the first egg in the clutch, females laid heavier second eggs during the poor year than during the good year. Females paired with males with duller feet colour reduced second-egg mass and volume and delayed the laying of the second egg, independently of the year. Absolute yolk androstenedione (A4) concentration (but not testosterone, T) in second eggs was higher during a poor year than during a good year. Only during a year with poor breeding conditions, females paired with experimental males decreased the relative A4 concentration (but not T) in the second egg compared to control females. Thus, blue-footed booby females probably favour brood reduction by decreasing egg quality and increasing size asymmetry between chicks when the breeding and the mate conditions are poor.     

The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish

The occurrence of male pregnancy in the family Syngnathidae (seahorses, pipefishes, and sea dragons) provides an exceptionally fertile system in which to investigate issues related to the evolution of parental care. Here, we take advantage of this unique reproductive system to study the influence of maternal body size on embryo survivorship in the brood pouches of pregnant males of the broad-nosed pipefish, Syngnathus typhle. Males were mated with either two large females, two small females, a large then a small female, or a small then a large female. Our results show that offspring survivorship depends on an interaction between female body size and the number of eggs transferred by the female. Eggs of larger females deposited in large numbers are more likely to result in viable offspring than eggs of smaller females laid in large numbers. However, when females deposited smaller numbers of eggs, the eggs from smaller females were more likely to produce viable offspring compared to those from larger females. We found no evidence that this result was based on mating order, the relative sizes of competing females, or egg characteristics such as dry weight of eggs. Additionally, male body size did not significantly influence the survivorship of offspring during brooding. Our results suggest that the factors underlying offspring survivorship in pipefish may be more complex than previously believed, with multiple factors interacting to determine the fitness of individual offspring within the broods of pregnant males.     

Food supplementation affects egg albumen content and body size asymmetry among yellow-legged gull siblings

Parental decisions can determine offspring experience of environmental conditions. Such ‘maternal’ effects act both before and after hatching via, e.g., egg quality or the social milieu predisposed by parents. Resource availability may constrain the expression of adaptive maternal effects, and the specific pattern of allocation of these effects among offspring depending on their sex or birth order can result in different fitness payoffs to parents. Declining egg mass with laying order observed in several bird species may constitute an adaptive strategy of parental favouritism towards early hatching offspring with larger reproductive value but may also result from nutritional constraints on laying effort. A previous study has shown that the small size of the third, last laid (c-)egg in yellow-legged gull (Larus michahellis) clutches depends on food availability and that food-supplemented mothers increase the size of their female but not male c-eggs. Here, we show that increased mass of c-eggs laid by females supplemented with food after clutch initiation depends on increased albumen mass, which, in turn, enhances the size of daughters at hatching. Because asynchronous hatching results in a competitive disadvantage of c-chicks, present results suggest that mothers relieved from nutritional constraints enhance the size of daughters to compensate for their larger susceptibility to hatching last. The study also confirms the role of egg albumen content in determining hatchling size, previously experimentally detected only in one species in the wild. The effect of increased egg mass on offspring size persisted at least until day 8 after hatching, when, however, it did not vary with sex, suggesting intense negative selection on small female c-chicks in control broods. Hence, maternal effects mediated by egg albumen content had persistent effects on offspring size.     

Supplemental food affects egg size but not hatching asynchrony in rollers

Maternal effects can function as a mechanism of transgenerational plasticity by which the environment experienced by parents is translated into the offspring phenotype and fitness. In birds, parents may affect the competitive ability of their offspring, and hence their fitness, by modifying their hatching pattern and/or egg size. However, little is known about how mothers can modify offspring phenotypes and their fitness in response to a sudden change in environmental conditions during egg-laying. Here, we studied the effect of supplemental food during egg-laying on hatching asynchrony and egg size in the Eurasian roller (Coracias garrulus), a species with marked hatching asynchrony. We also explored the effects of maternal investment on offspring fitness. Food supplementation did not affect hatching asynchrony. However, females in food-supplemented nests laid eggs that increased in size with laying order except for an ultimate small egg. Meanwhile, size of eggs laid by females in control nests did not change with laying order. Supplemental food positively affected hatchability of the egg laid just before the last one and negatively affected hatchability of the last laid egg, which seemed to be a side effect of egg size. Consequently, food-supplemented nests produced fewer fledglings and had higher probabilities of suffering brood reduction than control nests. We conclude that egg size in rollers is a plastic trait, sensitive to short-term changes in food conditions. Furthermore, our results show that maternal investment in egg size may potentially affect offspring fitness.     

Male sexual attractiveness affects the investment of maternal resources into the eggs in peafowl (<Emphasis Type="Italic">Pavo cristatus</Emphasis>)

According to the differential investment hypothesis, females paired with attractive mates are expected to invest more in the current reproduction relative to females paired with unattractive males. We experimentally tested this hypothesis in the peafowl (Pavo cristatus) by providing females with males that differed in sexual attractiveness. In agreement with the differential allocation hypothesis, females paired with more ornamented males laid larger eggs, and deposited higher amounts of testosterone into the egg yolk, independently of the sex of the embryo. These results show that the association between paternal phenotype and offspring quality could arise via a differential maternal investment. They also suggest that, if ornamented males do transmit good genes to the progeny, the maternal differential investment can amplify the effect of such good genes on the offspring fitness.     

Sexual dimorphism in house sparrow eggs

Recent evidence has revealed an apparently high degree of control by female birds over the physiological aspects of their reproduction and offspring sex allocation, consistent with adaptive hypotheses of sex allocation and differential investment in their offspring. In the house sparrow, we investigated possible mechanisms that may be used by females to enhance the fitness returns from a reproductive effort. Using molecular techniques, we demonstrate that house sparrow eggs containing male embryos are significantly larger than those containing female embryos. We also found that male embryos were laid randomly with respect to laying order. We speculate that this sexual dimorphism of eggs is adaptive, because male house sparrows show greater variance in condition-dependent reproductive success than females. More important, the result provides further evidence of the ability of females to detect or control ovulation of either male or female ova and to differentially invest in one sex over the other. Received: 19 January 2000 / Revised: 29 June 2000 / Accepted 20 July 2000     

Egg size and laying order in relation to offspring sex in the extreme sexually size dimorphic brown songlark,<Emphasis Type="Italic"> Cinclorhamphus cruralis</Emphasis>

In some bird species, mothers can advantage the offspring of one sex either by elevating them in the laying order to promote earlier hatching or by allocating greater resources to eggs of the preferred sex. In size dimorphic species, the predictions as to which sex should benefit most from such pre-laying adjustments are ambiguous. The smaller sex would benefit from an initial size advantage to help compensate for the faster growth rate of the larger sex. However, an early advantage to offspring of the larger sex might have a greater effect on their lifetime reproductive success than an equivalent advantage to offspring of the smaller sex. We investigated these hypotheses in the polygynous brown songlark, Cinclorhamphus cruralis, which is one of the most sexually size dimorphic birds known. We conducted within-clutch comparisons and found that females hatched from larger eggs and were initially heavier (but not structurally larger) than their brothers. This may afford females an early competitive advantage, as egg volume remained correlated with chick mass until at least 5 days of age. Similarly, we found that hatch order was still positively associated with nestling mass and size when the brood was 10 days of age, but there was no clear relationship between offspring sex and hatching order. During this study, food was plentiful and there were few obvious cases of nestling starvation. When food is limited, we suggest that the greater nutrient reserves of female hatchlings could not only help compensate for their slower growth, but could also give them a survival advantage over their brothers early in the nestling period. Consequently, egg size dimorphism may be an adaptation that facilitates an early shift in brood sex-ratio towards cheaper daughters in conditions of low food availability.     

Sex ratio varies with egg investment in the red-necked phalarope (Phalaropus lobatus)

Fisher’s sex ratio theory predicts that on average parents should allocate resources equally to the production of males and females. However, when the cost/benefit ratio for producing males versus females differs, the theory predicts that parents may bias production, typically through underproduction of the sex with greater variation in fitness. We tested theoretical predictions in the red-necked phalarope, a polyandrous shorebird with sex-role reversal. Since females are larger and therefore potentially more expensive to produce and may have greater variation in reproductive success, we predicted from Fisher’s hypothesis a male bias in population embryonic sex ratio, and from sex allocation theory, female biases in the clutches of females allocating more resources to reproduction. We measured eggs and chicks and sexed 535 offspring from 163 clutches laid over 6 years at two sites in Alaska. The embryonic sex ratio of 51.1 M:48.9 F did not vary from parity. Clutch sex ratio (% male) was positively correlated with clutch mean egg size, opposite to our prediction. Within clutches, however, egg size did not differ by sex. Male phalarope fitness may be more variable than previously thought, and/or differential investment in eggs may affect the within-sex fitness of males more than females. Eggs producing males were less dense than those producing females, possibly indicating they contained more yolk relative to albumen. Albumen contributes to chick structural size, while yolk supports survivorship after hatch. Sex-specific chick growth strategies may affect egg size and allocation patterns by female phalaropes and other birds.     

Influence of Forest Fragmentation on Community Structure of Frogs and Lizards in Northeastern Costa Rica

Abstract:  To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1–7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.     

Food supply during prelaying period modifies the sex-dependent investment in eggs of Eurasian kestrels

The theory of sex allocation suggests that if the reproductive value and the cost of producing/rearing offspring differ between male and female offspring, parents should invest differently in sexes depending on environmental conditions. Female parents could allocate more resources to eggs of one sex to compensate potential sex-dependent constraints later during the nestling period. In this study, we tested the influence of environmental conditions on sexual dimorphism in eggs of Eurasian kestrels (Falco tinnunculus) by experimentally manipulating food availability before laying. We found that an increase in food abundance before laying did not increase egg mass but changed sex-dependent resource distribution in eggs. In food-supplemented pairs, but not in control pairs, egg mass and hatchling mass were similar between males and females. In addition, we found, in the food-supplemented group, that the latest hatched females showed shorter hatching times than in the control group. In control pairs, female eggs, hatchlings and nestlings were heavier than males. In addition, male fledglings in the food-supplemented group gained less mass than those in the control group. As that food abundance was only increased until the onset of laying, female kestrels were expected to invest in eggs taking food abundance before egg formation as a predictor of future conditions during brood rearing. Our study shows that environmental conditions before laying promote a subtle adjustment of the resources invested in both sexes of offspring rather than in other breeding parameters. This adjustment resulted in a shortening of hatching time of the last hatched females that possibly gives them advantages in their competitive capacity with respect to male nest-mates.     

A field test of the intraspecific brood parasitism hypothesis in the golden egg bug (<Emphasis Type="Italic"> Phyllomorpha laciniata</Emphasis>)

In natural populations of golden egg bugs ( Phyllomorpha laciniata), females lay eggs on plants where they develop unattended, or on conspecifics, where they remain firmly glued until the nymphs hatch and start an independent life. Mortality rates among eggs laid on plants are higher than among eggs carried by adults. Because females cannot lay eggs on themselves, in order to improve offspring survival, they have to lay eggs on other individuals. Two hypotheses have been proposed to explain egg carrying: (1) the mating pair intraspecific brood parasitism hypothesis suggests that females dump eggs on copulating pairs, and (2) the paternal care hypothesis suggests that the system is driven mainly by males accepting eggs to improve the survival rates of their own offspring. Our data from the field show that 77% of the eggs are carried by males, because more males than females carry eggs, and because males carry a greater number of eggs. In addition, we show that mating males carry more recently laid eggs than single males. These results support the view that egg carrying is performed predominantly by males and that eggs are laid on males by their current mating partner, probably between repeated copulations. Males are likely to accept eggs, despite intermediate levels of paternity, because they cannot discriminate in favour of their own eggs, because rejected eggs will face 97% mortality rates on plants, and because they do not suffer mating costs when they carry eggs. However, females carry 23% of the eggs, but no differences in egg carrying have been found between mating and single females, suggesting that this is not the result of egg dumping while females are copulating. Egg carrying by females could reflect low levels of intraspecific parasitism, which is likely to reflect the low rate of successful attempts by egg-laying females who try to oviposit on other conspecifics rather indiscriminately, in an effort to improve the survival of their offspring.     

Female egg investment in relation to male sexual traits and the potential for transgenerational effects in sexual selection

Life-history theory predicts that individuals should increase their reproductive effort when the fitness return from reproduction is high. Females mated with high-quality males are therefore expected to have higher investment than females mated with low-quality males, which could bias estimates of paternal effects. Investigating the traits females use in their allocation decisions and the aspects of reproduction that are altered is essential for understanding how sexual selection is affected. We studied the potential for differential female allocation in a captive population of a precocial bird, the Chinese quail, Coturnix chinensis. Females paired with males with large sexual ornaments laid larger, but not more, eggs than females paired with males with small sexual ornaments. Furthermore, female egg mass was also significantly positively affected by male testis size, probably via some unknown effect of testis size on male phenotype. Testis size and ornament size were not correlated. Thus, both primary and secondary male sexual traits could be important components of female allocation decisions. Experimental manipulation of hormone levels during embryonic development showed that both male and female traits influencing female egg size were sensitive to early hormone exposure. Differences in prenatal hormone exposure as a result of maternal steroid allocation to eggs may explain some of the variation in reproductive success among individuals, with important implications for non-genetic transgenerational effects in sexual selection.Communicated by C. Brown     

Adjustment of female reproductive investment according to male carotenoid-based ornamentation in a gallinaceous bird

Carotenoid-based ornaments (many yellow–orange–red colourations) may signal the genetic or parental quality of the bearer. Thus, their expression could influence the amount of resources/energy that the mate will invest in the production of offspring, thereby optimising its reproductive fitness. The differential allocation hypothesis (DAH) predicts that females mated with more attractive males should lay more and better eggs. This has been explored only in few bird species with carotenoid-based traits. We tested this hypothesis in the red-legged partridge (Alectoris rufa), a gallinacean with very variable laying capacity. Both sexes display carotenoid-based ornamentation that gradually fades throughout the laying period. Here, the redness of beak and eye rings of captive males was intensified after mating by means of paint. The proportion of females that laid eggs did not differ between treatments. Amongst laying females, those mated with colour-enhanced males (experimental females) tended to lay earlier and produced significantly more eggs than controls, but of similar quality (egg mass and composition). We additionally investigated whether male attractiveness influenced egg components depending on the clutch size and laying sequence. The testosterone level in eggs from experimental females was positively related to the laying order, whereas control eggs did not show any trend. Our results provided mixed support for the DAH, but nevertheless revealed that female red-legged partridges may adjust their breeding investment according to male carotenoid-based ornamentation.     

Long-lasting consequences of elevated yolk testosterone levels on female reproduction

Maternal yolk androgens in bird eggs represent an important pathway along which offspring phenotype is shaped. Most of the hormone-mediated maternal effects are highly important in the context of sibling competition. However, there is also increasing evidence for long-lasting effects far beyond the nestling period, and these effects may have important consequences on the reproductive success of the offspring. Here, we investigated the effects of experimentally elevated yolk testosterone concentrations on growth and reproduction in female canaries. Elevated yolk testosterone concentrations enhanced the post-natal growth rate, but not the asymptotic mass, and reduced the survival probability. The latter may be a consequence of the higher growth rate, which may have rendered females hatching from testosterone-treated eggs (T-females) more vulnerable to harsh environmental conditions. Adult T-females made a larger investment in their clutch by laying more but not heavier eggs than females hatching from control-treated eggs. Our results suggest that the observed long-lasting effect on clutch size relates to changes in the growth trajectory rather than being a direct consequence of testosterone, since studies manipulating early growth conditions obtained similar results. Clearly, further studies are now required in order to investigate the intriguing relationship between yolk testosterone, elevated growth rates, and clutch size.     

Costs and benefits of surplus offspring in the lesser kestrel (Falco naumanni )

Lesser kestrels (Falco naumanni) lay clutches which appear excessive as only 3% of them yield as many young as eggs laid. Four hypotheses may explain the adaptive value of producing surplus eggs: (1) the bet-hedging hypothesis assumes that the environment varies unpredictably and surplus eggs serve to track uncertain resources; (2) the ice-box hypothesis suggests that surplus offspring serve as a reserve food during a period of shortage; (3) the progeny choice hypothesis says that parents produce surplus offspring in order to choose these with higher fitness; and (4) the insurance-egg hypothesis proposes that extra eggs are an insurance against the failure of any egg. To test the significance of this strategy in the lesser kestrel, an experiment manipu-lating brood size at hatching was carried out over 2 years, with good and bad feeding conditions. The experiment consisted of adding a chick to experimental broods where one egg failed to hatch or removing a randomly selected chick from experimental broods where all eggs had hatched. Independently of annual food availability, pairs with brood sizes reduced by one chick fledged more nestlings than pairs with brood size equalling their clutch sizes. Body condition of young was also better in the former group, but only in 1993 (a high-food year). Independently of year, mean local survival of parents with complete broods at hatching was lower than for parents raising reduced broods. These results supported only the insurance-egg hypothesis which says that surplus eggs may be an insurance against the failure of any egg, but parents may suffer reproductive costs when all eggs hatch. Received: 17 January 1997 / Accepted after revision: 27 April 1997     

A Spatially Explicit Decision Support Model for Restoration of Forest Bird Habitat

Abstract:  The historical area of bottomland hardwood forest in the Mississippi Alluvial Valley has been reduced by >75%. Agricultural production was the primary motivator for deforestation; hence, clearing deliberately targeted higher and drier sites. Remaining forests are highly fragmented and hydrologically altered, with larger forest fragments subject to greater inundation, which has negatively affected many forest bird populations. We developed a spatially explicit decision support model, based on a Partners in Flight plan for forest bird conservation, that prioritizes forest restoration to reduce forest fragmentation and increase the area of forest core (interior forest >1 km from "hostile" edge). Our primary objective was to increase the number of forest patches that harbor >2000 ha of forest core, but we also sought to increase the number and area of forest cores >5000 ha. Concurrently, we targeted restoration within local (320 km²) landscapes to achieve ≥60% forest cover. Finally, we emphasized restoration of higher-elevation bottomland hardwood forests in areas where restoration would not increase forest fragmentation. Reforestation of 10% of restorable land in the Mississippi Alluvial Valley (approximately 880,000 ha) targeted at priorities established by this decision support model resulted in approximately 824,000 ha of new forest core. This is more than 32 times the amount of core forest added through reforestation of randomly located fields (approximately 25,000 ha). The total area of forest core (1.6 million ha) that resulted from targeted restoration exceeded habitat objectives identified in the Partners in Flight Bird Conservation Plan and approached the area of forest core present in the 1950s.     

Female choice contributes to offspring fitness in the field cricket,Gryllus bimaculatus (De Geer)

Summary Females of the field cricket, Gryllus bimaculatus, are known to mate non-randomly with respect to male size. The reproductive output of females allowed to choose their mates was compared with that of females allocated mates. Females allowed to choose their mates laid a greater proportion of their available eggs, thereby investing more heavily in reproduction. Female choice did not appear to influence hatching success, suggesting that no short term benefits are derived from non-random mating. However, the offspring of females allowed to choose their mates developed more rapidly and began their own reproductive output before those of females who had been allocated mates. Furthermore, the faster developing offspring of females who chose their mates tended to have a consistantly higher survival rate as well as a lower variance in survival (i.e. less risk of mortality) at adult eclosion. Female choice may therefore contribute to offspring fitness and thus to long term reproductive success. Females allocated large males as mates did not equal those of females allowed to choose their mates, in terms of reproductive output and offspring fitness. This suggests that females may choose their mates with regard to a mixture of characters of which male size is only one.     

Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Possible mutualism between females of the subsocial membracid Polyglypta dispar (Homoptera)

Summary Female Polyglypta dispar membracids facultatively guard egg masses or oviposit into masses guarded by other females. Defending females repel at least some enemies of both eggs and nymphs. Eggs of guarding females may be partially protected from parasitism by the presence of additional eggs laid by other females. Females sometimes desert egg masses or groups of nymphs, and undefended masses are sometimes adopted by other females. Some individuals remain on or near the plant where they grew up, and littermates sometimes oviposit into the same egg mass. Some copulating pairs are littermates, while others are not. Females with larger numbers of mature and nearly mature eggs in their ovaries are less likely to defend egg masses, and females desert smaller groups of nymphs more often.