Maternal Condition and Nest Site Choice: An Alternative for the Maintenance of Environmental Sex Determination?

SYNOPSIS. Egg size and nest site selection are two potentialeffects that can have a persistent influence on the phenotypeof offspring. In this paper, I develop the maternal condition-dependentchoice hypothesis for the maintenance of environmental sex determination.The hypothesis stipulates three conditions: 1) there must bevariation in the maternal effect, 2) the variation in the maternaleffect must influence fitness of the offspring differently betweenthe sexes, and 3) female reproductive behavior is determinedby her condition or how her condition will influence her offspring'sfitness. Females with the ability to recognize environmentsthat have a higher probability of producing the sex that wouldbenefit the most from maternal condition will have an advantage.Using egg size as a maternal effect, I test this hypothesisin the diamondback terrapin, an emydid turtle with temperature-dependentsex determination. Terrapins have large variation in egg sizeamong clutches and little variation within clutches. Egg massis the primary determinant of hatchling mass and can resultin as much as a three year difference in reaching minimum sizeof first reproduction in females, but may not affect age orsize of first reproduction in males. Finally, terrapins selectopen nesting sites with warmer incubation conditions and placelarger eggs there. Females place smaller eggs in cooler sites.Terrapin reproduction is consistent with the prediction of thematernal condition-dependent nest site choice hypothesis. Themodel and supporting data demonstrate how maternal effects canbe an important factor to consider in studies of environmentalsex determination.     

Social and Thermal Cues Influence Nest‐site Selection in a Nocturnal Gecko,Oedura lesueurii

In oviparous species lacking parental care, successful reproduction depends on females selecting nest sites that facilitate embryonic development. Such sites may be limited in the environment, which can lead to multiple females using the same nest site simultaneously. However, there are several alternative explanations for communal nesting, including natal homing, predator satiation, and adaptive benefits to offspring. We used laboratory experiments to evaluate three hypotheses about nest‐site selection in velvet geckos (Oedura lesueurii), which often nest communally. We investigated whether the trend to nest communally is influenced by the following: (1) evidence of previous nesting (hatched eggshells); (2) body size; and/or (3) thermal regimes. When given the choice, females laid their eggs in shelters containing hatched eggshells rather than in empty shelters, and this was not influenced by body size. Females selected nest sites that were cooler than their own mean selected body temperatures, suggesting that thermal requirements of their developing embryos could outweigh their own thermoregulatory preferences. Field observations of natal homing and high predation rates on gravid females suggest that imprinting on nest sites and/or predator swamping also play roles in communal nesting. Collectively, our results suggest that female velvet geckos use multiple cues to select appropriate nest sites, and hence that multiple mechanisms result in communal nesting behavior in this species.     

Body size-specific maternal effects on the offspring environment shape juvenile phenotypes in Atlantic salmon

Positive associations between maternal investment per offspring and maternal body size have been explained as adaptive responses by females to predictable, body size-specific maternal influences on the offspring’s environment. As a larger per-offspring investment increases maternal fitness when the quality of the offspring environment is low, optimal egg size may increase with maternal body size if larger mothers create relatively poor environments for their eggs or offspring. Here, we manipulate egg size and rearing environments (gravel size, nest depth) of Atlantic salmon (Salmo salar) in a 2 × 2 × 2 factorial experiment. We find that the incubation environment typical of large and small mothers can exert predictable effects on offspring phenotypes, but the nature of these effects provides little support to the prediction that smaller eggs are better suited to nest environments created by smaller females (and vice versa). Our data indicate that the magnitude and direction of phenotypic differences between small and large offspring vary among maternal nest environments, underscoring the point that removal of offspring from the environmental context in which they are provisioned in the wild can bias experimentally derived associations between offspring size and metrics of offspring fitness. The present study also contributes to a growing literature which suggests that the fitness consequences of egg size variation are often more pronounced during the early juvenile stage, as opposed to the egg or larval stage.     

Nest taphonomy of common terns (Sterna hirundo) on Poplar Island,Chesapeake Bay,Maryland

Mesozoic non-avian theropod dinosaurs displayed a diverse range of egg types and clutch forms, suggesting a variety of nesting behaviours, some of which may be shared with birds. More accurate inferences of these behaviours require taphonomic studies of modern nesting sites. Here, we document common tern (Sterna hirundo) nesting sites on Poplar Island in Chesapeake Bay, Maryland. Nests were surveyed on multiple occasions, documenting nest composition, density and distribution, as well as eggshell concentration and orientation. Three colonies yielded 79 tern nests with 193 eggs. Twelve nests hatched, 7 were predated, 30 failed and the fates of 30 remain unknown. Abundant eggshells occurred at the centre of the nests. Concave-up eggshell in or on the nest surface characterised hatched and predated nests, whereas eggshell forced into the subsurface seemed to favour concave-down orientation and may reflect post-hatching extended use of the nest. Eggs buried deep within the nest materials and/or substrate suggest adult abandonment. Quantitative data on eggshell orientation and observations regarding controls on nest distribution, egg predation and incorporation of intact eggs into the substrate at modern nesting sites provide physical guidelines for the improved interpretation of fossil nesting localities.     

Why do female lizards lay their eggs in communal nests?

1. In many reptile species, females oviposit communally (i.e. many clutches are laid within the same nest). This behaviour might result from constraint (scarcity of nest-sites offering suitable incubation conditions) or adaptation (direct fitness benefits accruing from the proximity of other eggs). 2. To test between these alternatives, we gathered field and laboratory data on montane scincid lizards Bassiana duperreyi from south-eastern Australia. Our data support the adaptationist hypothesis. 3. In the field, communal vs. solitary clutches were laid in similar sites, and the relative frequency of communal nesting was not predictable from nest-site availability. Thermal regimes for incubation did not differ between communal vs. solitary nests, nor between eggs at the core vs. periphery of a communal nest. In the laboratory, females selectively oviposited beside existing eggs rather than in otherwise identical potential nesting sites. 4. From cycling-temperature incubation in the laboratory, eggs incubated within a cluster of other eggs took up less water, but produced hatchlings that were larger and faster-running than were hatchlings from eggs incubated alone. 5. Hydric modifications of incubation conditions within a cluster of tightly packed eggs thus may provide a direct fitness benefit to communal oviposition.     

Adaptive egg size plasticity for larval competition and its limits in the seed beetle Callosobruchus chinensis

Life‐history theory predicts that females who experienced stressful conditions, such as larval competition or malnutrition, should increase their investment in individual offspring to increase offspring fitness (the adaptive parental hypothesis). In contrast, it has been shown that when females were reared under stressful conditions, they become smaller, which consequently decreases egg size (the parental stress hypothesis). To test whether females adjust their egg volume depending on larval competition, independent of maternal body mass constraint, we used a pest species of stored adzuki beans, Callosobruchus chinensis (L.) (Coleoptera: Chrysomelidae: Bruchinae). The eggs of females reared with competitors were smaller than those of females reared alone, supporting the parental stress hypothesis; however, correcting for female body size, females reared with competitors produced larger eggs than those reared in the absence of competition, supporting the adaptive parental hypothesis, as predicted. The phenotypic plasticity in females' investment in each offspring in stressful environments counteracts the constraint of body size on egg size.     

Don'T put all your eggs in one nest: spread them and cut time at risk

Abstract In many egg-laying animals, some females spread their clutch among several nests. The fitness effects of this reproductive tactic are obscure. Using mathematical modeling and field observations, we analyze an unexplored benefit of egg spreading in brood parasitic and other breeding systems: reduced time at risk for offspring. If a clutch takes many days to lay until incubation and embryo development starts after the last egg, by spreading her eggs a parasitic female can reduce offspring time in the vulnerable nest at risk of predation or other destruction. The model suggests that she can achieve much of this benefit by spreading her eggs among a few nests, even if her total clutch is large. Field data from goldeneye ducks Bucephala clangula show that egg spreading enables a fecund female to lay a clutch that is much larger than average without increasing offspring time at risk in a nest. This advantage increases with female condition (fecundity) and can markedly raise female reproductive success. These results help explain the puzzle of nesting parasites in some precocial birds, which lay eggs in the nests of other females before laying eggs in their own nest. Risk reduction by egg spreading may also play a role in the evolution of other breeding systems and taxa-for instance, polyandry with male parental care in some birds and fishes.     

Dispersed versus gregarious nesting strategies in the mason bee Chalicodoma siculum

Chalicodoma siculum (Rossi) is a mason bee that usually nests in dispersed sites, constructing spherical mud nests on twigs or hemispherical nests on hard surfaces. Under favourable conditions it may nest in aggregations of several hundred bees. At the nesting aggregation, nest usurpation often occurs, either by the laying of eggs in finished but unsealed cells, by opening a sealed cell and replacing its egg, or by taking over an unfinished cell. Mutual construction and provisioning of a single cell by several females was also often observed. Because participant bees display extreme aggression in prolonged disputes over cell ownership, these interactions can be regarded as competitive rather than cooperative. Moreover, since the total investment in such cells is not different from that in cells located in dispersed sites, the female that actually manages to lay an egg gains by participating in a cell under construction.
We discuss the advantages of nesting in aggregations vs. the risks of being usurped, as selective forces underlying nesting behaviour of bees.     

A new Upper Cretaceous titanosaur nesting site from La Rioja (NW Argentina), with implications for titanosaur nesting strategies

Cretaceous titanosaur nesting sites are currently known only from Europe, Asia and South America. In the latter, only the Auca Mahuevo and Sanagasta nesting sites have been confidently assigned to this clade of sauropod dinosaurs. Here we report the discovery of the first eggs and egg clutches found at Tama, a new Upper Cretaceous fossiliferous locality in the Los Llanos Formation, Sierra de Los Llanos (La Rioja, NW Argentina). At least five egg clutches, several partially preserved, isolated eggs and many eggshell fragments were discovered in a single outcrop of a sandstone horizon which represents a cumulative palaeosol profile. Although the mechanical and digital preparation of eggs did not reveal any embryonic remains in ovo, the morphology of the eggs and eggshells closely matches that of titanosaur eggs and eggshells found worldwide. The morphology and spatial grouping of the titanosaur eggs from Tama, along with geological observations support a burrow‐nesting strategy for these dinosaurs. Although the Sanagasta and Tama eggs were found in the same stratigraphical unit and share several morphological characters, they clearly differ in shell thickness and egg size. This, coupled with the interpretation of different sedimentary contexts for these nesting sites, strongly suggests that at least two different titanosaur species nested in La Rioja during the Late Cretaceous, using different nesting strategies. The occurrence of this new titanosaur nesting site in a semiarid palaeoenvironment represents an interesting case study for the reproductive biology of the titanosaur dinosaurs, particularly their labile nesting behaviour.     

Allocation of offspring size and sex by female black ratsnakes

How females allocate resources to each offspring and how they allocate the sex of their offspring are two powerful potential avenues by which mothers can affect offspring fitness. Previous research has focussed extensively on mean offspring size, with much less attention given to variance in offspring size. Here we focussed on variation in offspring size in black ratsnakes, Elaphe obsoleta . We collected and hatched 105 clutches (1283 eggs) over 9 years. We predicted that females should lay larger eggs, or more variable eggs, when the environment is less predictable. We also predicted that females laying early or laying larger eggs should produce mostly sons because adult males are larger than adult female ratsnakes. The largest hatchling was more than twice the length and almost four times the mass of the smallest hatchling. Variation in offspring size was itself highly variable, with CVs in offspring mass among clutches ranging from 1% to 25%. With one exception, the variables we expected should influence variation in offspring size had little effect. We found that clutch size increased with maternal size and that egg size decreased with clutch size, but we found no evidence that variance in egg size among clutches increased as the season progressed or that females increased the mean size of their offspring the later in the season they laid their eggs. Females in better condition after they finish laying their eggs did produce larger eggs. There was no relationship between within-clutch variation in egg size and laying date or mean egg size. Finally, sex ratio did not vary with mean egg size or hatching date. Given evidence that offspring size in snakes affects survival, selection should reduce variation in offspring size unless that variance enhances maternal fitness and yet we found little support for hypothesized advantages of varying offspring size.     

Loggerhead turtle eggshells as a source of maternal nuclear genomic DNA for population genetic studies

Tagging studies on nesting beaches are commonly used to estimate nesting frequency, remigration interval and nesting population size for marine turtle rookeries. Estimates of these demographic parameters from tagging projects may be biased because of the small scale of tagging efforts relative to female nest site fidelity and the logistical difficulty of intercepting all nesting females. Therefore, alternative and supplemental means of individual identification of nesting females are required. We demonstrate that maternal nuclear microsatellite DNA can be isolated from unincubated eggshells of the loggerhead sea turtle (Caretta caretta) through comparison of DNA extracted from 59 eggs collected within 15 h of oviposition and DNA derived from skin samples from respective nesting females. Scorable microsatellite genotypes were produced in 897 of 994 (90.2%) single-locus egg amplifications attempted. Among eggs from known females, 730 of 748 (97.6%) single-locus, egg-derived genotypes matched the respective skin-derived genotypes. Allelic dropout was the most common type of error, followed by the presence of nonmaternal, presumably paternal, alleles. Genotypes derived from unincubated eggshells permit individual assignment of nests and therefore demographic parameter estimates for loggerhead turtle nesting populations, despite genotyping errors that require further optimization. Although sampling unincubated eggs is destructive, this technique is noninvasive to nesting females and is applicable in marine turtle population genetics studies when individual resolution is required but direct interception of nesting females is undesirable or logistically infeasible.     

Effect of soil hardness on aggression in the solitary wasp Mellinus arvensis

1. Two alternative nesting strategies are exhibited by soil‐nesting Mellinus arvensis females, digging a new nest (diggers) and searching for an old unoccupied burrow (searchers). Wasps appear unable to distinguish between occupied and unoccupied nests, and aggressive interactions between searchers and nest owners at nest entrances are frequent. 2. In aggressive encounters, there is an advantage in size and residency status. 3. The costs associated with the two nesting strategies vary across geographically separated populations: nest digging incurs costs in terms of time, and these vary according to the hardness of the soil substrate; nest searching is variably costly in terms of risk of injury in aggressive encounters with nest‐owning females. 4. Individual female wasps can switch between nesting strategies, and thus soil hardness, by affecting the cost of nest construction, affects the relative frequencies of the two nesting strategies within a population, favouring an increase in the searching strategy. This, in turn, affects the frequency and intensity of aggression between females at a nesting aggregation. 5. Female body size is correlated with soil hardness. As aggressive encounters are more frequent in sites with hard soil substrates, there is increased selective advantage in having large body size at these sites. 6. Body size is determined primarily by the availability of food resources during larval development, which is, to a degree, a function of the size of the adult female. There is a trade‐off between provisioning a few cells with many provisions in each, leading to the development of few but large adults, as opposed to many cells with few provisions, leading to many small offspring. The relative advantage of these two provisioning strategies is, at least in part, a function of the hardness of the soil substrate.     

Brood parasitism,female condition and clutch reduction in the common eider Somateria mollisima

Conspecific brood parasitism (CBP) is an alternative reproductive tactic found in many animals with parental care. Parasitizing females lay eggs in the nests of other females (hosts) of the same species, which incubate and raise both their own and the foreign offspring. The causes and consequences of CBP are debated. Using albumen fingerprinting of eggs for accurately detecting parasitism, we here analyse its relation to female condition and clutch size in High Arctic common eiders Somateria mollissima borealis. Among 166 clutches in a Svalbard colony, 31 (19%) contained eggs from more than one female, and 40 of 670 eggs (6%) were parasitic. In 6 cases an active nest with egg(s) was taken over by another female. Many suitable nest sites were unoccupied, indicating that CBP and nest takeover are reproductive tactics, not only consequences of nest site shortage. Similarity in body mass between female categories suggests that condition does not determine whether a nesting female becomes parasitised. There was no evidence of low condition in parasites: egg size was similar in hosts and parasites, and parasitism was equally frequent early and late in the laying season. Meta‐analysis of this and 3 other eider studies shows that there is a cost of being parasitised in this precocial species: host females laid on average 7% fewer eggs than other females.     

Egg survival is related to the colour matching of eggs to nest background in Black-tailed Gulls

A long-standing hypothesis posits that, for species with exposed nests, a close match between the colour of the eggs and that of the nest in which they are laid should enhance egg survival, but this has rarely been tested in a rigorous manner. Here, we demonstrate the effects of egg–nest colour matching on egg survival in Black-tailed Gulls (Larus crassirostris) on Hongdo Island, Korea. We quantified the ground colour of eggshells and that of the nest background using a digital camera and computerized RGB and greyscale colour systems. We show that a close match of eggshell ground colour and nest background colour was associated with increased chances of eggs surviving through to hatching. In particular, there were strong survival advantages for eggs matching the nest colour in sites with poor concealment, whereas there was no effect of eggshell ground colour in nests that were more concealed by vegetation. Our findings support the hypothesis that egg colour functions to make eggs cryptic and that egg colouration may be a significant factor affecting egg loss.     

Cues for communal egg‐laying in lizards (Bassiana duperreyi,Scincidae)

Animals may aggregate either because the presence of conspecifics provides information about habitat suitability, or because the presence of conspecifics directly enhances individual viability. For a female lizard, the advantage of laying her eggs in a communal nest may entail either information transfer (hatched eggshells show that the site has been successful in previous seasons) or direct physiological benefits (recently‐laid eggs can enhance water availability to other eggs). We tested the relative importance of these two mechanisms in the three‐lined alpine skink (Bassiana duperreyi Gray, 1838) by offering gravid females a choice between sites with hatched eggshells versus freshly‐laid eggs. Females selectively oviposited beside fresh eggs. In this species, early‐nesting females use information transfer (i.e. the presence of old eggshells) as a nest‐site criterion, whereas later nesters switch to a reliance on direct benefits of conspecific presence (i.e. the presence of freshly‐laid eggs). © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 839–842.     

Nesting in a thermally challenging environment: nest‐site selection in a rock‐dwelling gecko,Oedura lesueurii (Reptilia: Gekkonidae)

In egg‐laying species, maternal oviposition choice can influence egg survival and offspring phenotypes. According to the maternal‐preference offspring‐performance hypothesis, females should choose oviposition sites that are optimal for offspring fitness. However, in thermally challenging environments, maternal oviposition behaviour may be constrained by the limited availability of suitable oviposition sites. We investigated nest‐site selection in a nocturnal lizard [velvet gecko Oedura lesueurii (Duméril and Bibron)] that inhabits a thermally challenging environment in south‐eastern Australia. The viability of these gecko populations is critical for the persistence of an endangered snake species (Hoplocephalus bungaroides Wagler) that feeds heavily on velvet geckos. Female geckos chose nest sites nonrandomly, with 87% of nests (N = 30) being laid in deep crevices. By contrast, only 13% of clutches were laid under rocks, which were the most readily available potential nest sites. Nest success in crevices was high (100%), but no eggs hatched from nests under rocks. Temperatures in nest crevices remained relatively low and constant throughout the incubation period (mean = 22.7 °C, range 21.0–24.5 °C), whereas thermal regimes under rocks showed large diurnal fluctuations. Geckos selected crevices that were deeper, had less canopy cover, and were warmer than most available crevices; in 85% of cases, such crevices were used simultaneously by more than one female. The thermally distinctive attributes of nest sites, and their frequent communal use, suggest that nest sites are a scarce resource for female velvet geckos, and that the shading of rock outcrops through vegetation encroachment may influence nest success in this species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 250–259.     

Individual Variation in Trophic Egg Production: Evidence for Maternal Manipulation in Response to Resource and Competition Levels

Females of the subsocial shield bug Parastrachia japonensis (Heteroptera: Parastrachiidae) incorporate trophic eggs (nutritive eggs) into their egg mass. Considerable variation occurs among females in trophic egg number and the proportion of an egg mass that is composed of trophic eggs. Because trophic eggs are essential to the development and survival of young, this variation could significantly impact female fitness. We tested the hypothesis that trophic egg abundance is induced by maternal phenotype (weight, body size) and resource exposure. We predicted that resource limitations would cause females to produce fewer fertile eggs and more trophic eggs and that larger and heavier females would produce more of each egg type. Females ovipositing early in the season are exposed to different resource conditions than those that oviposit late. Thus, we compared egg production patterns between these two groups and several other factors related to nesting. No correlation was seen between body size and trophic egg abundance, or, indeed, egg production, overall; however, heavier females produced heavier egg masses. Counter to our prediction, late females, which had greater access to food, produced significantly more total eggs, fewer fertile eggs, and more trophic eggs than early females. A binomial generalized linear model analysis indicated that the factors most correlated with the percentage of an egg mass destined to become trophic eggs were resource abundance, resulting from early or late oviposition, and distance of the nest from the host tree, with closer females producing more trophic eggs. The findings support our hypothesis that resource availability and, to a lesser extent, maternal phenotype affect trophic egg abundance.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Natural beaches confer fitness benefits to nesting marine turtles

Coastal ecosystems provide vital linkages between aquatic and terrestrial habitats and thus support extremely high levels of biodiversity. However, coastlines also contain the highest densities of human development anywhere on the planet and are favoured destinations for tourists, creating a situation where the potential for negative effects on coastal species is extremely high. I gathered data on marine turtle reproductive output from the literature to determine whether coastal development negatively influences offspring production. Female loggerhead (Caretta caretta) and green turtles (Chelonia mydas) nesting on natural beaches (as opposed to beaches with permanent development) produce significantly more hatchling turtles per nest; all else being equal, females that successfully produce more offspring will have higher fitness than conspecifics producing fewer offspring. Thus, female marine turtles nesting on natural beaches probably have higher fitness than turtles nesting on developed beaches. Consequently, populations nesting on natural beaches may be able to recover more quickly from the historic population declines that have plagued marine turtles, and some species may recover more quickly than others.     

Reproductive ecology of the jacky dragon (Amphibolurus muricatus): an agamid lizard with temperature‐dependent sex determination

Abstract The jacky dragon, Amphibolurus muricatus (White, ex Shaw 1790) is a medium sized agamid lizard from the southeast of Australia. Laboratory incubation trials show that this species possesses temperature‐dependent sex determination. Both high and low incubation temperatures produced all female offspring, while varying proportions of males hatched at intermediate temperatures. Females may lay several clutches containing from three to nine eggs during the spring and summer. We report the first field nest temperature recordings for a squamate reptile with temperature‐dependent sex determination. Hatchling sex is determined by nest temperatures that are due to the combination of daily and seasonal weather conditions, together with maternal nest site selection. Over the prolonged egg‐laying season, mean nest temperatures steadily increase. This suggests that hatchling sex is best predicted by the date of egg laying, and that sex ratios from field nests will vary over the course of the breeding season. Lizards hatching from eggs laid in the spring (October) experience a longer growing season and should reach a larger body size by the beginning of their first reproductive season, compared to lizards from eggs laid in late summer (February). Adult male A. muricatus attain a greater maximum body size and have relatively larger heads than females, possibly as a consequence of sexual selection due to male‐male competition for territories and mates. If reproductive success in males increases with larger body size, then early hatching males may obtain a greater fitness benefit as adults, compared to males that hatch in late summer. We hypothesize that early season nests should produce male‐biased sex ratios, and that this provides an adaptive explanation for temperature‐dependent sex determination in A. muricatus.