Effects of Roads on the Structure of Freshwater Turtle Populations

Abstract:  Road mortality has the potential to alter the structure of turtle populations because turtle populations are highly sensitive to additive sources of adult mortality. To address the issue, we captured painted turtles (   Chrysemys picta ; n = 174) and snapping turtles (   Chelydra serpentina ; n = 56) in 18 wetlands surrounded by low road density (≤1.5 km roads/km² of landscape) and 17 wetlands surrounded by high road density (>1.5 km/km²) in central New York in 2002. High road density was associated with male-biased sex ratios in painted turtles (74% vs. 54% males; p = 0.01) and snapping turtles (95% vs. 74% males; p = 0.08), whereas turtle morphology and abundance were not associated with road density. Disproportionate road mortality of females on nesting migrations is the most likely cause of skewed sex ratios.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Sex ratio of juvenile loggerhead turtles in the Mediterranean Sea: is it really 1:1?

Sex ratios are a crucial parameter for evaluating population viability. In species with complex life history patterns and temperature sex determination mechanisms, such as the loggerhead turtle (Caretta caretta), sex ratios may vary within a population and among populations. In the Mediterranean, juvenile sex ratios appear to not differ significantly from 1:1, although estimates for hatchling sex ratios are highly female biased. The immigration of males from the Atlantic has been suggested as a possible cause of such variation. Here, we present results of a multi-year investigation (2000–2011) on the sex ratios of loggerhead turtles foraging along the south Tyrrhenian coast, Western Mediterranean, with the aim of providing a better understanding of the potentially underlying forces that drive regional and age-dependent differences in sex ratios. Sex was determined through visual examination of the gonads in 271 dead turtles (curved carapace length range 29.5–89 cm). A fragment of the mitochondrial DNA control region was sequenced from 61 specimens to characterise the demographic composition of this foraging assemblage by applying a many-to-many mixed stock analysis approach. No significant association was found between sex ratios and years or size classes, although the largest size was male biased. Juvenile sex ratio was 1.56:1, which was different from an even sex ratio but still less female biased than hatchling sex ratios from Mediterranean beaches. Results of the mixed stock analysis indicate that juvenile sex ratios in the Mediterranean are largely unaffected by immigration of Atlantic individuals into the basin, as previously suggested. Continued long-term monitoring of juvenile sex ratios is necessary to detect biologically significant sex ratio shifts in the Mediterranean loggerhead turtle population.     

Deforestation: risk of sex ratio distortion in hawksbill sea turtles.

Phenotypic sex in sea turtles is determined by nest incubation temperatures, with warmer temperatures producing females and cooler temperatures producing males. The common finding of highly skewed female-biased hatchling sex ratios in sea turtle populations could have serious repercussions for the long-term survival of these species and prompted us to examine the thermal profile of a relatively pristine hawksbill nesting beach in Guadeloupe, French West Indies. Data loggers placed at nest depth revealed that temperatures in the forested areas were significantly cooler than temperatures in the more open, deforested areas. Using these temperatures as a predictor of sex ratio, we were able to assess the relative contributions of the different beach zones to the primary sex ratio: significantly more males were likely to be produced in the forested areas. Coastal forests are therefore important male-producing areas for the hawksbill sea turtle, and this has urgent conservation implications. On Guadeloupe, as on many Caribbean islands, deforestation rates are high and show few signs of slowing, as there is continual pressure to develop beachfront areas. The destruction of coastal forest could have serious consequences both in terms of local nesting behavior and of regional demography through the effects on population sex ratios. Human alterations to nesting habitat in other reptile taxa have been shown to modify the thermal properties of nest sites in ways that can disrupt their ecology by allowing parasite transmission, increasing vulnerability to climate change, or rendering existing habitat unsuitable.     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

A Contemporary,Sex‐Limited Change in Body Size of an Estuarine Turtle in Response to Commercial Fishing

Abstract: Juvenile growth rate and adult body size are important components of life‐history strategies because of their direct impact on fitness. The diamondback terrapin (Malaclemys terrapin) is a sexually dimorphic, long‐lived turtle inhabiting brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of its range, terrapins face anthropogenically imposed mortality: juveniles of both sexes inadvertently enter commercial crab traps and drown. For adult females, the carapace eventually grows large enough that they cannot enter traps, whereas males almost never reach that critical size. We compared age structure, carapace dimensions, growth curves, and indices of sexual dimorphism for a Chesapeake Bay population of terrapins (where mortality of turtles is high due to crab traps) with contemporary terrapins from Long Island Sound and museum specimens from Chesapeake Bay (neither group subject to commercial crab traps). We also calculated the allochronic and synchronic rates of evolutionary change (haldanes) for males and females to measure the rate of trait change in a population or between populations, respectively. We found a dramatic shift to a younger male age structure, a decrease in the length of time to terminal female carapace size, a 15% increase in female carapace width, and an increase in sexual dimorphism in Chesapeake Bay. In a new twist, our results implicate a fishery in the selective increase in size of a reptilian bycatch species. These sex‐specific changes in life history and demography have implications for population viability that need to be considered when addressing conservation of this threatened turtle.     

Estimating freshwater turtle mortality rates and population declines following hook ingestion

Freshwater turtle populations are susceptible to declines following small increases in the mortality of adults, making it essential to identify and understand potential threats. Freshwater turtles ingest fish hooks associated with recreational angling, and this is likely a problem because hook ingestion is a source of additive mortality for sea turtles. We used a Bayesian‐modeling framework, observed rates of hook ingestion by freshwater turtles, and mortality of sea turtles from hook ingestion to examine the probability that a freshwater turtle in a given population ingests a hook and subsequently dies from it. We used the results of these analyses and previously published life‐history data to simulate the effects of hook ingestion on population growth for 3 species of freshwater turtle. In our simulation, the probability that an individual turtle ingests a hook and dies as a result was 1.2–11%. Our simulation results suggest that this rate of mortality from hook ingestion is sufficient to cause population declines. We believe we have identified fish‐hook ingestion as a serious yet generally overlooked threat to the viability of freshwater turtle populations.     

Migratory and reproductive movements of male leatherback turtles (<Emphasis Type="Italic">Dermochelys coriacea</Emphasis>)

The biology of the endangered leatherback turtle (Dermochelys coriacea) at sea is poorly understood. As research has been almost exclusively limited to studies of nesting females, the movements of male turtles and the timing and location of mating activity have remained unknown. We report on the first deployments of satellite tags on male leatherbacks. Male migration to and residency in waters adjacent low-latitude nesting beaches in the western Atlantic suggest that this is where mating occurs, and return migration to these areas reveals male fidelity for breeding sites. Rate of travel decreased markedly after arriving in coastal breeding areas, where males remained for up to 96 days before assuming northward migrations. The initiation of these northward migrations coincided with peak nesting activity in adjacent nesting colonies. Data from satellite-linked time-depth recorders attached to two males revealed diel dive patterns in breeding areas and marked differences in diving behaviour between migratory and breeding periods in one turtle. When male turtles were in waters adjacent nesting colonies, their movements differed from those reported for nesting females, with females ranging farther from shore. Our results suggest that male leatherbacks may be vulnerable to entanglement in coastal fishing gear in waters adjacent nesting beaches.Communicated by R.J. Thompson, St. Johns     

Using Expert Opinion Surveys to Rank Threats to Endangered Species: A Case Study with Sea Turtles

Abstract: Little is known about how specific anthropogenic hazards affect the biology of organisms. Quantifying the effect of regional hazards is particularly challenging for species such as sea turtles because they are migratory, difficult to study, long lived, and face multiple anthropogenic threats. Expert elicitation, a technique used to synthesize opinions of experts while assessing uncertainty around those views, has been in use for several decades in the social science and risk assessment sectors. We conducted an internet‐based survey to quantify expert opinion on the relative magnitude of anthropogenic hazards to sea turtle populations at the regional level. Fisheries bycatch and coastal development were most often ranked as the top hazards to sea turtle species in a geographic region. Nest predation and direct take followed as the second and third greatest threats, respectively. Survey results suggest most experts believe sea turtles are threatened by multiple factors, including substantial at‐sea threats such as fisheries bycatch. Resources invested by the sea turtle community, however, appear biased toward terrestrial‐based impacts. Results from the survey are useful for conservation planning because they provide estimates of relative impacts of hazards on sea turtles and a measure of consensus on the magnitude of those impacts among researchers and practitioners. Our survey results also revealed patterns of expert bias, which we controlled for in our analysis. Respondents with no experience with respect to a sea turtle species tended to rank hazards affecting that sea turtle species higher than respondents with experience. A more‐striking pattern was with hazard‐based expertise: the more experience a respondent had with a specific hazard, the higher the respondent scored the impact of that hazard on sea turtle populations. Bias‐controlled expert opinion surveys focused on threatened species and their hazards can help guide and expedite species recovery plans.     

Diamondback Terrapin (Malaclemys terrapin) Mortality in Crab Pots

The entrapment of diamondback terrapins (  Malaclemys terrapin) in crab pots frequently results in drowning and death of the trapped turtles. We determined the rate of capture, size, sex, and age of terrapins captured, and the potential impact crab pot mortality has on local terrapin populations. We estimated terrapin capture rates of 0.17 terrapins/pot/day in shallow water areas of Chesapeake Bay (Maryland, USA). The sex ratio of terrapins caught in crab pots was 3:2 male biased because female terrapins become too large to enter crab pots by the time they reach 8 years of age. Males, however, remain vulnerable to entrapment throughout their life. Our estimates of capture rates and local population size suggest that 15–78% of a local population may be captured in a single year. As a consequence, crab pots may be the major reason terrapins are extirpated in coastal, shallow water areas with heavy crab pot fisheries. Additionally, the selective removal of males may also contribute to female-biased sex ratios observed in this diamondback terrapin population. We developed and tested a modified crab pot that increases terrapin survival and does not reduce the number of crabs caught. Our modified crab pot maintained permanent access to air and prevented the drowning of terrapins. Additionally, our modified crab pot caught more crabs than standard commercial crab pots, suggesting that the modified crab pot could be a viable alternative to standard traps that result in terrapin mortality.     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Sex ratios of loggerhead sea turtles Caretta caretta during the juvenile pelagic stage

Sex ratios are a fundamental trait for species reproduction. In species with temperature-dependent sex determination (TSD), sex ratios are not necessarily even, which has important demographic consequences. We examined the sex ratio of juvenile pelagic stage loggerhead turtles Caretta caretta offshore Madeira Island, North Eastern Atlantic, using laparoscopy and histology. The overall sex ratio was 2:1 (F:M), significantly different from an even sex ratio. Although there was no apparent temporal variation, sex ratios among size classes were significantly different. The sex ratio of juveniles was compared with known sex ratios for the putative source rookery and found to be similar to the subadults’ sex ratio, but significantly less female-biased than the hatchlings sex ratio. This suggests overestimation of hatchlings sex ratios and/or, less likely, differential mortality of females during the first months of life. Alternatively, the Madeira Island aggregation may be recruiting males from other geographical sources such as the Mediterranean and the Cape Verde.     

Effects of urbanization on the population structure of freshwater turtles across the United States

Landscape‐scale alterations that accompany urbanization may negatively affect the population structure of wildlife species such as freshwater turtles. Changes to nesting sites and higher mortality rates due to vehicular collisions and increased predator populations may particularly affect immature turtles and mature female turtles. We hypothesized that the proportions of adult female and immature turtles in a population will negatively correlate with landscape urbanization. As a collaborative effort of the Ecological Research as Education Network (EREN), we sampled freshwater turtle populations in 11 states across the central and eastern United States. Contrary to expectations, we found a significant positive relationship between proportions of mature female painted turtles (Chrysemys picta) and urbanization. We did not detect a relationship between urbanization and proportions of immature turtles. Urbanization may alter the thermal environment of nesting sites such that more females are produced as urbanization increases. Our approach of creating a collaborative network of scientists and students at undergraduate institutions proved valuable in terms of testing our hypothesis over a large spatial scale while also allowing students to gain hands‐on experience in conservation science.     

Facultative sex ratio manipulation in American kestrels

Summary For animals that are sexually dimorphic in size, the larger sex is expected to be more costly to raise to independence. Manipulating offspring sex ratios may thus be one means by which parents can fine-tune their reproductive effort to resource availability. Parents in poor physical condition or during poor food years should produce more of the cheaper (smaller) sex. We examined the sex ratios of 259 broods of American kestrels (Falco sparverius) between 1988 and 1990 in relation to food abundance (small mammals) and various attributes to the parents. The proportion of males at hatching increased as the food supply declined, and both male and female parents in poor physical condition were more likely to have male-biased broods than those in good condition. The mortality of eggs and young did not appear to be responsible for the biased sex ratios. The sex ratio was independent of the laying date; however, it was correlated with female body size. Small females produced more sons, perhaps because small size is more detrimental for females than males. Offprint requests to: G.R. Bortolotti     

Sex allocation and local mate competition in Old World non-pollinating fig wasps

The populations of many species are structured such that mating is not random and occurs between members of local patches. When patches are founded by a single female and all matings occur between siblings, brothers may compete with each other for matings with their sisters. This local mate competition (LMC) selects for a female-biased sex ratio, especially in species where females have control over offspring sex, as in the parasitic Hymenoptera. Two factors are predicted to decrease the degree of female bias: (1) an increase in the number of foundress females in the patch and (2) an increase in the fraction of individuals mating after dispersal from the natal patch. Pollinating fig wasps are well known as classic examples of species where all matings occur in the local patch. We studied non-pollinating fig wasps, which are more diverse than the pollinating fig wasps and also provide natural experimental groups of species with different male morphologies that are linked to different mating structures. In this group of wasps, species with wingless males mate in the local patch (i.e. the fig fruit) while winged male species mate after dispersal. Species with both kinds of male have a mixture of local and non-local mating. Data from 44 species show that sex ratios (defined as the proportion of males) are in accordance with theoretical predictions: wingless male species<wing-dimorphic male species<winged male species. These results are also supported by a formal comparative analysis that controls for phylogeny. The foundress number is difficult to estimate directly for non-pollinating fig wasps but a robust indirect method leads to the prediction that foundress number, and hence sex ratio, should increase with the proportion of patches occupied in a crop. This result is supported strongly across 19 species with wingless males, but not across 8 species with winged males. The mean sex ratios for species with winged males are not significantly different from 0.5, and the absence of the correlation observed across species with wingless males may reflect weak selection to adjust the sex ratio in species whose population mating structure tends not to be subdivided. The same relationship is also predicted to occur within species if individual females adjust their sex ratios facultatively. This final prediction was not supported by data from a wingless male species, a male wing-dimorphic species or a winged male species. Received: 27 July 1998 / Received in revised form: 11 January 1999 / Accepted: 16 January 1999     

Intra-Mediterranean migrations of loggerhead sea turtles (Caretta caretta) monitored by satellite telemetry

The movements of four Mediterranean loggerhead sea turtles (Caretta caretta; three females, one male) were tracked via satellite telemetry for between 108 and 457 days. Total length of the routes traveled by the turtles varied between 2554 and 7098 km, and the average travel rate was 1.2 km h^-1. Long-distance movement between the western and eastern Mediterranean basins followed a seasonal pattern and seemed to be triggered by temperature and food availability. In the autumn/winter months turtles moved from west to east in search of warmer waters, and returned to the western basin in spring, where food resources are generally more plentiful. Three (two females, one male) of the four turtles migrated eastward through the Straits of Messina, which is characterized by high fishing pressure and intense boat traffic. Information about turtle migration patterns and routes will serve to plan effective conservation strategies.     

Putting Longline Bycatch of Sea Turtles into Perspective

Abstract:  Although some sea turtle populations are showing encouraging signs of recovery, others continue to decline. Reversing population declines requires an understanding of the primary factor(s) that underlie this persistent demographic trend. The list of putative factors includes direct turtle and egg harvest, egg predation, loss or degradation of nesting beach habitat, fisheries bycatch, pollution, and large-scale changes in oceanographic conditions and nutrient availability. Recently, fisheries bycatch, in particular bycatch from longline fisheries, has received increased attention and has been proposed as a primary source of turtle mortality. We reviewed the existing data on the relative impact of longline bycatch on sea turtle populations. Although bycatch rates from individual longline vessels are extremely low, the amount of gear deployed by longline vessels suggests that cumulative bycatch of turtles from older age classes is substantial. Current estimates suggest that even if pelagic longlines are not the largest single source of fisheries-related mortality, longline bycatch is high enough to warrant management actions in all fleets that encounter sea turtles. Nevertheless, preliminary data also suggest that bycatch from gillnets and trawl fisheries is equally high or higher than longline bycatch with far higher mortality rates. Until gillnet and trawl fisheries are subject to the same level of scrutiny given to pelagic longlines, our understanding of the overall impact of fisheries bycatch on vulnerable sea turtle populations will be incomplete.     

Demographic Evidence of Illegal Harvesting of an Endangered Asian Turtle

Harvesting pressure on Asian freshwater turtles is severe, and dramatic population declines of these turtles are being driven by unsustainable collection for food markets, pet trade, and traditional Chinese medicine. Populations of big‐headed turtle (Platysternon megacephalum) have declined substantially across its distribution, particularly in China, because of overcollection. To understand the effects of chronic harvesting pressure on big‐headed turtle populations, we examined the effects of illegal harvesting on the demography of populations in Hong Kong, where some populations still exist. We used mark‐recapture methods to compare demographic characteristics between sites with harvesting histories and one site in a fully protected area. Sites with a history of illegal turtle harvesting were characterized by the absence of large adults and skewed ratios of juveniles to adults, which may have negative implications for the long‐term viability of populations. These sites also had lower densities of adults and smaller adult body sizes than the protected site. Given that populations throughout most of the species’ range are heavily harvested and individuals are increasingly difficult to find in mainland China, the illegal collection of turtles from populations in Hong Kong may increase over time. Long‐term monitoring of populations is essential to track effects of illegal collection, and increased patrolling is needed to help control illegal harvesting of populations, particularly in national parks. Because few, if any, other completely protected populations remain in the region, our data on an unharvested population of big‐headed turtles serve as an important reference for assessing the negative consequences of harvesting on populations of stream turtles. Evidencia Demográfica de la Captura Ilegal de una Tortuga Asiática en Peligro     

Experimentally manipulated brood sex ratios: growth and survival in the black-headed gull (Larus ridibundus), a sexually dimorphic species

In sexually size dimorphic species, individuals of the larger sex often suffer from enhanced mortality during the nestling period. This has been attributed to higher nutritional requirements of the larger sex, which may render this sex more vulnerable to adverse food conditions. However, sex-biased mortality might not exclusively depend on the differences in food demand but also on other phenotypic differences, e.g., in competitiveness. Interference competition between the sexes and position in the laying sequence in particular may be essential components contributing to biased mortality.By creating synchronously-hatched unisex broods in the sexually size dimorphic black-headed gull, we specifically tested the effect of sex-specific food demand by excluding interference competition between the sexes as well as hatching asynchrony. To test the effect of egg quality, which varies with the position in the laying sequence, we composed each nest of chicks from eggs of all different positions in the laying sequence.All-male nests showed significantly enhanced mortality compared to all-female nests from the beginning of the development of the sexual size dimorphism onwards. This underlines the role of a higher food demand in biased mortality of the larger sex.In males but not females, asymptotic body mass and skeletal size were negatively associated with position in the laying sequence, while survival was not affected by position. As a consequence, sexual size dimorphism at the end of the nestling period was less pronounced compared to the natural situation. These data show that, although male growth is more sensitive to a decrease in egg quality, the higher mortality of last hatched chicks in natural nests is mainly due to hatching asynchrony and egg size but not egg content.     

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