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.     

Sex ratio in lesser black-backed gull in relation to environmental pollutants

In birds, there is ample evidence that the mother can manipulate the sex of the young and produce more of the sex, which gives the highest fitness return. This has previously been documented in gulls, Laridae. Gulls are sexually size dimorphic with males larger than females, and there is good evidence that parents in poor body condition switch their investment to the smallest sex. In the present study, we examined the primary sex ratio and the survival of male and female chicks of lesser black-backed gull (Larus fuscus fuscus) in relation to their blood levels of organochlorines (OCs), perfluorinated compounds (PFCs) and polybrominated diphenyl ethers (BDE-47). We show that females with high levels of OCs (but not PFCs and BDE-47) are likely to skew their sex ratio at hatching towards female offspring. Few females had very high levels of OCs, and the many females with low levels of OCs overproduced sons resulting in a male skew at hatching (59%). The survival of female offspring was lower than the survival of male offspring, causing an even stronger male skew in sex ratio (71%). There is evidence to conclude that circulating levels of OCs in the blood of females may have detrimental effect on the sex allocation strategy and could be of serious threat to the population. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Parental condition, brood sex ratio and differential young survival: an experimental study in gulls (Larus fuscus)

Empirical evidence is growing that the offspring sex ratio in birds can be biased in relation to the body condition of parents during breeding. The sex ratio bias may come about because (1) the actual production of the two sexes may be skewed and/or (2) there may be a sex bias in early nestling mortality contingent on parental condition. By manipulating parental condition and giving them a control brood to rear, thereby eliminating effects operating via the eggs, we examined the extent to which parental condition influences the post-hatching survival of male and female lesser black-backed gulls, Larus fuscus. We found that the pre-fledging survival of male chicks was strongly reduced in all-male broods reared by parents in poor condition. Pre-fledging survival of female chicks was, however, unaffected by parental condition or brood sex composition. Thus, independently of any production biases, sex differences in nestling mortality alone can bias the offspring sex ratio at fledging in relation to the prevailing rearing conditions. In other studies on gulls we have, however, also shown that females in poor condition at laying preferentially produce female eggs. Clearly a bias in fledging sex ratio can occur within the same species due to a combination of differential production and differential post-laying mortality; the latter can involve a differential effect of poor egg quality on male and female offspring, differential effects of brood sex composition on their survival and a difference in the capacity of parents to rear males and females. All of these processes need to be taken into account in attempting to understand offspring sex ratios. Received: 15 February 2000 / Revised: 7 August 2000 / Accepted: 26 August 2000     

Sex-specific maternal effect on egg mass,laying order,and sibling competition in the Bengalese finch (<Emphasis Type="Italic">Lonchura striata</Emphasis> var. <Emphasis Type="Italic">domestica</Emphasis>)

Maternal effects, such as investment in eggs, have profound effects on offspring fitness. Mothers are expected to skew their investment depending on the laying order and sex when unequal sibling competition occurs within a brood because of sex-specific vulnerability and age hierarchy caused by asynchronous hatching. The Bengalese finch hatches asynchronously and shows a moderate reversed sexual size dimorphism. However, contrary to commonly accepted assumptions of size-dependent vulnerability, the smaller sex (male) is more vulnerable to developmental stress caused by sibling competition. We investigated whether maternal investment would be biased by the position in laying order and the sex of eggs, and also explored the possible differences in growth patterns depending on sex, laying order, and age hierarchy by observing chicks fostered to experimentally manipulated broods where brood composition was controlled and age hierarchy was more enhanced than in natural breeding conditions. We found that overall patterns of maternal investment favored the disadvantageous sectors of sibling competition, i.e., eggs of later laying order and sons over those of early laying order and daughters. We also examined the effect of laying order on adult body size and sex differences in growth patterns. When reared in the subordinate age hierarchy, females could compensate for the deficit of decreased growth rate by taking longer to mature, whereas males could not. We suggest that this sex-specific growth pattern could be the cause of sex differences in vulnerability to early developmental stress.     

Effects of elevated yolk testosterone levels on survival, growth and immunity of male and female yellow-legged gull chicks

Androgen hormones of maternal origin contained in the eggs of avian species are considered to have positive effects on offspring characteristics and performance. However, negative consequences have also been reported, suggesting that mothers may experience a trade-off between beneficial and detrimental effects of egg androgens to offspring fitness. We studied the effects of elevated yolk testosterone (T) concentration on survival, development and phenotype of male and female yellow-legged gull (Larus michahellis) chicks by injecting egg yolks with physiological doses of the hormone. Elevated yolk T resulted in a male-biased post-hatching sex ratio, T-treated clutches producing a greater proportion of males compared to control ones at day 4 post-hatching, likely resulting from a reduction of female embryonic survival, whereas no effect of hormone treatment on hatching success or short-term chick survival was observed. In addition, T depressed post-hatching body mass in both sexes but had no effects on the intensity of the cell-mediated immune response or skeletal growth. No sex differences in egg characteristics or chick phenotype were detected. Time to hatching was not affected by T, but females originating from first laid eggs hatched earlier than males of the same laying order, independently of hormone treatment. However, the implications of sex differences in hatching times are unclear in the study species. Taken together, our results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.     

Small Populations and Offspring Sex-Ratio Deviations in Eagles

Abstract:  Stochastic variation of sex ratio has long been appreciated as a potential factor driving small populations to extinction, but it is not the only source of sex-ratio bias in small populations. We examined whether some consequences of sex allocation could affect extinction risk in small populations of size-dimorphic birds such as eagles. We report variations in sex ratio at fledging from a long-term study of a declining population of Spanish Imperial Eagles ( Aquila adalberti ). Nestling sex-ratio deviation apparently was mediated by age of breeders, whereas territory quality had no obvious effect. Adult–adult pairs produced the same proportion of both sexes in high- or low-density situations, but pairs with at least one member in nonadult plumage class produced more males. As the population declined over a period of years, the proportion of breeders with immature plumage increased; consequently, the proportion of fledgling males increased. However, when population density was high, the proportion of breeders with immature plumage decreased and more female offspring were produced. This relationship between population density, composition of breeder age, and fledgling sex ratios allowed us to make predictions of extinction risk due to nonstochastic deviations of sex ratio in small, declining populations. In the study population, on the basis of the Vortex simulation results, an estimated reduction of 42.5% in predicted mean time to extinction was attributed solely to biased sex ratio.     

The energetic cost of protogynous versus protandrous sex change in the bi-directional sex-changing fish Gobiodon histrio

To investigate the relative cost of protogynous versus protandrous sex change we induced sex change in each direction in Gobiodon histrio (Gobiidae) and then compared growth, body condition and biochemical condition between sex-changed and non-sex-changed fish in treatment and control groups. Sex change in each direction was induced by establishing pairs of adult males and pairs of adult females on isolated coral colonies. Heterosexual pairs were used as controls. For both males and females, growth and body condition did not vary between fish that changed sex and those that did not. The relatively low cost of sex change, compared to the likely costs of searching for a mate of the correct sex, appears to explain the evolution of bi-directional sex change in coral-dwelling gobies. Lipid concentrations in the liver of males and females that changed sex were reduced by similar amounts (34% and 41%, respectively) compared to male and female controls that did not change sex. Therefore, changes in biochemical condition were approximately equal for fish that sex change in each direction and cannot explain the unequal frequency of sex change in each direction in natural populations of G. histrio.     

Sex-specific effects of albumen removal and nest environment manipulation on Barn Swallow nestlings

In avian species, maternal provisioning to the eggs is predicted to be more valuable for the offspring under adverse environmental conditions and intense sibling competition. However, studies manipulating both the amount of maternal pre-hatching resources and the harshness of post-hatching environment have seldom been performed to date. In this experimental study of Barn Swallow (Hirundo rustica) nestlings, we tested the consequences of a reduction in the albumen content of the eggs for fitness-related offspring traits, while performing an unbalanced partial cross-fostering soon after hatching, either increasing or decreasing brood size by one nestling. By molecular sexing of the chicks, we additionally tested for sex-specific sensitivity of individual nestlings to experimental treatments and to sex ratio variation in nestmates. We predicted that chicks hatching from albumen-deprived eggs should suffer more than control chicks from the harsher rearing conditions of enlarged broods. However, although albumen removal depressed chick body mass, chicks hatching from control eggs did not fare better than those hatching from eggs with reduced albumen content in enlarged vs. reduced broods. Albumen removal had sex-specific effects on immunity, with males, but not females, hatching from eggs with reduced albumen content showing a lower T-cell-mediated immune response than controls, suggesting that the two sexes were differentially susceptible to resource deprivation during early ontogeny. In addition, both immune response and chick body mass at age 7 days, when maximum growth rate is attained, declined with an increasing proportion of male nestmates. The effect of brood size manipulation on chick body mass at age 12 days, when peak body mass is attained, was also found to depend on brood sex composition, in that an increase in the proportion of male nestmates depressed offspring body mass in reduced broods, while the reverse was true in enlarged broods. On the whole, these findings suggest that sex differences may exist in environmental sensitivity and patterns of resource allocation among different body functions, and that brood size variation and sex composition may affect offspring fitness-related traits.     

Biased sperm use by polyandrous queens of the ant<Emphasis Type="Italic"> Proformica longiseta</Emphasis>

The occurrence and genetic effects of polyandry were studied in the ant Proformica longiseta using three microsatellite markers. The average queen mating frequency (QMF) estimated from the sperm dissected from the spermathecae of 61 queens was 2.4 with 69% of the queens being multiply mated. QMF estimated from worker offspring in a subsample of eight monogynous colonies was 3.5, but the effective paternity (m_e,p) was only 1.23. The difference between these values reflected unequal sperm use by the queens. Most colonies of P. longiseta were polygynous and the average relatedness among workers was 0.35. Polyandry thus added only marginally to the genetic diversity of colonies, and our results gave little support to the genetic-variability hypothesis for explaining polyandry. Diploid male load was low, as only 1% of males were diploid. A large majority (92%) of nests produced one sex only, with males produced in colonies that had higher than average worker relatedness. This contradicted the predictions derived from worker control of sex ratios. Males produced enough sperm to fill the spermathecae of several queens. Thus, the results indicated that diploid male load, sperm limitation and sex ratio conflict are also unlikely explanations of polyandry. Plausible hypotheses for polyandry include mating by convenience, as the sex ratio is male biased and the mating costs to a female can be low because the females are wingless and have no mating flight. The observed unequal sperm use furthermore points to sperm choice and sperm competition as important factors in the evolution of polyandry.     

Studies on the growth,reproduction, and life cycle of the supralittoral isopod Ligia pallasii

A two-year study on a field population of Ligia pallasii Brandt has shown that the isopods live for 1.5 to 2 years. Breeding occurs in the spring and early summer, with some females carrying winter broods of eggs. The mean length of breeding females is 22.5±2.2 mm (standard deviation) and the mean brood size is 48±11 eggs. Mature males are larger than mature females (900 and 300 mg live weight, respectively), and are disproportionately broader (31×17 mm and 22.5×9 mm, respectively). The larger size and breadth of the males is an adaptation for copulation, and may be atributed in some measure to slower growth of the female due to the extra energy demands of reproduction. The overall 1:1 ratio of males to females in the population represents the balance between an equal sex ratio in the immature stages, more females in the 18 to 24 mm length category, and more males in the larger length categories. This condition is attributed mainly to the faster growth of the males.     

An adaptive annual rhythm in the sex of first pigeon eggs

When the reproductive value of male and female offspring varies differentially, parents are predicted to adjust the sex ratio of their offspring to maximize their fitness (Trivers and Willard, Science 179:90–92, 1973). Two factors have been repeatedly linked to skews in avian offspring sex ratio. First, laying date can affect offspring sex ratio when the sexes differ in age of first reproduction, such that the more slowly maturing sex is overproduced early in the season. Second, position of the egg in the laying sequence of a clutch may affect sex ratio bias since manipulating the sex of the first eggs may be least costly to the mother. We studied both factors in two non-domesticated pigeon species. Both the Wood pigeon (Columba palumbus) and the Rock pigeon (Columba livia) have long breeding seasons and lay two-egg clutches. In the field, we determined the sex of Wood pigeon nestlings. In Rock pigeons, housed in captivity outdoors, we determined embryo sex after 3 days of incubation. On the basis of their sex-specific age of first reproduction, we predicted that males, maturing at older age than females, should be produced in majority early and females later in the year. This was confirmed for both species. The bias was restricted to first eggs. Rock pigeons produced clutches throughout the year and show that the sex of the first egg followed an annual cycle. To our knowledge, this study presents the first evidence of a full annual rhythm in adaptive sex allocation in birds. We suggest that this reflects an endogenous seasonal program in primary sex ratio controlled by a preovulatory mechanism.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Female cannibalism and male courtship tactics in threespine sticklebacks

Summary Female threespine sticklebacks (Gasterosteus aculeatus) frequently raid male nests and eat all the eggs therein. We tested the hypothesis of Vickery et al. (1988) that females prefer to raid nests containing large numbers of eggs than ones with smaller numbers of eggs. This hypothesis is based on the finding that females spawning in nests containing many eggs will have reduced hatching success because of egg crowding. By consuming the male's eggs and forcing him to rebuild his nest, raiding females might obtain a new opportunity to spawn under better conditions. Our results were consistent with the first prediction of this hypothesis that females were more likely to spawn in nests containing fewer eggs than in nests with many eggs. However, this may be the result of males becoming less receptive to females as the number of eggs in their nests increases. Prediction 2 was that females should raid those nests containing the most eggs. Contrary to this prediction, males defending only one clutch were as likely to have their nests raided by groups of females as males defending several clutches of eggs. Female cannibalism is therefore unlikely to have evolved as a means of gaining access to a male defending a small number of eggs. We also examined the tactics used by males to counter female raids. Most raids occur when the male is courting, and nests are more vulnerable to shoals of females than to single females. Therefore, we hypothesized that males with eggs preferentially court a single female rather than large groups of females, and that males without eggs court both groups indiscriminately. We also predicted that males restrict the number of females they mate with when risk of having their nest raided is high. Our results indicate that: (1) both males with eggs and those without eggs minimize the risk of female cannibalism by courting solitary females rather than groups of females and (2) males limit the number of females that lay eggs in their nest when several potentially raiding females are present. Offprint requests to: G.J. FitzGerald     

Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae)

Summary Colonies of Anelosimus eximius in Panama had an average sex ratio of 0.15±sd 0.09, i.e. about five females for each male. The sex ratio in egg sacs reared was even lower (0.08±0.01), as was that of immatures in newly founded colonies (0.12±0.05). The possible mechanisms responsible are discussed. Mature colonies had an average ratio of 17 females and 2 males for each egg sac present (range: 2–91 females, 0.2–8.2 males) and contained a large proportion of females which were not inseminated but which presumably help. Since both sexes are diploid, arrhenotoky can be ruled out and it is assumed that some females do not come to reproduction, the proportion depending on the availability of resources. This mechanism may enable entire colonies to survive lean times.     

Reproductive consequences of male body mass and aggressiveness depend on females’ behavioral types

Relatively few investigations explicitly test for concordant versus conflicting selection pressures from intrasexual versus intersexual selection. Here, we examine the effects of male body mass and behavioral type (BT) on reproductive success in the spider Anelosimus studiosus, with emphasis placed on the potential interaction between intrasexual and intersexual selection influences. Female A. studiosus exhibit either an aggressive-active or docile-passive BT, both of which co-occur in multifemale colonies. Males, in contrast, exhibit a more continuous distribution of behavioral tendencies. We investigated the male traits favored by females in five trial types: one docile female, one aggressive female, four docile females, four aggressive females, and two docile and two aggressive females. Male reproductive success was estimated by the number eggs produced by females following staged mating trials. In previous work, it was established that large aggressive males are favored in male–male contests, an intrasexual effect. However, large aggressive males were not universally favored here. We failed to detect an effect of male body mass or aggressiveness on reproductive success in trials with all docile females; however, in situations involving aggressive females, large aggressive males experienced diminished reproductive success relative to small docile males. Large, aggressive males were also more likely to be attacked and killed by aggressive females in the first 20 min of staged encounters and were more likely to be found dead after 72 h of unobserved interactions. Taken together, our data suggest that the reproductive consequences of male traits differ based on (1) the aspect of sexual selection being considered (intrasexual versus intersexual) and (2) the BT of their prospective mates: large aggressive males enjoy advantages in intrasexual selection and when courting docile females and small docile males experience reduced risk of cannibalism and increased reproductive success with aggressive females.     

Lifetime reproductive success in the spotted sandpiper (Actitis macularia) : sex differences and variance components

Summary Lifetime reproductive success (LRS) of male (N=103) and female (N= 66) spotted sandpipers (Actitis macularia) was studied for 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. There was no sex difference in longevity, but females had significantly more mates, eggs, chicks, fledged young, and young returning in subsequent years than did males. Variance in LRS was partitioned into five life-history components: longevity (L), mates per year (M), eggs per mate (E), proportion eggs hatched (H), and proportion of chicks fledged (F). For both sexes, F accounted for the greatest proportion of variance in LRS (males, 43%; females, 47%), followed by L (males, 26%; females, 43%) and H (males, 21%; females, 28%). Positive covariance between H and F was consistent with predator-caused clutch and brood loss. Contrary to our expectations, males had a higher coefficient of variation in reproductive success than did females. This was because males were relatively more likely than females to produce no young.Offprint requests to: L.W. Oring at the current address     

Brood sex ratios, female harem status and resources for nestling provisioning in the great reed warbler (Acrocephalus arundinaceus)

The theory of parental investment and brood sex ratio manipulation predicts that parents should invest in the more costly sex during conditions when resources are abundant. In the polygynous great reed warbler, Acrocephalus arundinaceus, females of primary harem status have more resources for nestling provisioning than secondary females, because polygynous males predominantly assist the primary female whereas the secondary female has to feed her young alone. Sons weigh significantly more than daughters, and are hence likely to be the more costly sex. In the present study, we measured the brood sex ratio when the chicks were 9 days old, i.e. the fledging sex ratio. As expected from theory, we found that female great reed warblers of primary status had a higher proportion of sons in their broods than females of lower (secondary) harem status. This pattern is in accordance with the results from two other species of marsh-nesting polygynous birds, the oriental reed warbler, Acrocephalus orientalis, and the yellow-headed blackbird Xanthocephalus xanthocephalus. As in the oriental reed warbler, we found that great reed warbler males increased their share of parental care as the proportion of sons in the brood increased. We did not find any difference in fitness of sons and daughters raised in primary and secondary nests. The occurrence of adaptive sex ratio manipulations in birds has been questioned, and it is therefore important that three studies of polygynous bird species, including our own, have demonstrated the same pattern of a male-biased offspring sex ratio in primary compared with secondary nests. Received: 1 June 1999 / Received in revised form: 10 January 2000 / Accepted: 12 February 2000     

Secondary sex ratios do not support maternal manipulation: extensive data from laboratory colonies of spiny mice (Muridae: Acomys)

Spiny mice of the genus Acomys (Muridae) represent a very suitable mammalian model for studying factors influencing the secondary sex ratio (SSR). The maternal effort in these rodents is extremely biased in favour of the prenatal period and, therefore, maternal manipulation of the SSR is potentially more advantageous. We studied the SSR in four populations/species of spiny mice kept in family groups consisting of two closely related females, one non-relative male and their descendants. The groups were established from founding animals aged about 3 months (maturing age) and were allowed to breed freely for several months. Each litter was sexed after birth, and relevant data were thoroughly recorded. Altogether, data were collected on 1684 litters: 189 of Acomys sp. from Iran, 203 of A. cilicicus, 875 of A. cahirinus, and 417 of A. dimidiatus. We recorded the sex of 4048 newborns of which 1995 were males and 2053 were females. The overall sex ratio was close to 1:1 (49.2%). Generalized linear mixed models and/or generalized linear models were constructed to evaluate the effect of four life history and eight social variables on the sex ratio. No consistent effects of these variables on the sex ratio were found and, interestingly, none of the variables associated with maternal life history had any effect on the sex ratio. Three factors associated with group composition (i.e. the number of immature males, the number of immature females and the number of breeding females) did have significant effects on the sex ratio, but these effects were not consistent across the studied species. In conclusion, our evaluation of this large dataset revealed that the sex ratio in spiny mice is surprisingly stable.     

Host choice and offspring sex allocation in a solitary parasitic wasp

Summary Females of the parasitic wasp Antrocephalus pandens can detect differences in the quality of their hosts (pupae of Corcyra cephalonica, a stored-product moth) and allocate offspring of either sex accordingly. Larger and younger hosts are accepted more often in both dead and live hosts; more female offspring emerge from the perceived better hosts, while more males emerge from the smaller, older ones. These patterns are consistent with a sex allocation strategy by the mother, since females from a given size host tend to be larger than males and larger females produce more eggs. However, when wasps lay their eggs in groups of hosts of different size and age rather than encountering them one at a time, no difference in number or sex ratio of offspring is detected between groups. This result and evidence from the change in offspring sex ratio with female age and with numbers of females foraging on a group of hosts are interpreted and discussed in the context of sex allocation (Charnov 1979) and local mate competition (LMC, Hamilton 1967) theories.     

Studies on portunid crabs from the Eastern Pacific. II. Significance of the unusual distribution of Euphylax dovii

Euphylax dovii Stimpson (Brachyura: Portunidae: Podophthalminae), a tropical Eastern Pacific swimming crab, has distinctive morphological adaptations for pelagic existence. Crabs in collections from the open ocean had a sex ratio approximating 1:1, with no crabs bearing eggs. Samples from the continental shelf of Colombia contained thousands of females, mostly ovigerous, but no males. Egg attachment has posed a major problem in the evolution of decapod crustaceans, and the two genera of portunid crabs thus far observed cannot attach eggs unless females can bury partly in soft sediments. This suggests that mated E. dovii females must migrate into shallow shelf waters to encounter sediments necessary for spawning. The high energetic cost of swimming while carrying eggs and the presence of abundant food for larvae are factors favoring residence of females in shelf waters until hatching is complete.