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.     

Persistent Unequal Sex Ratio in a Population of Grayling (Salmonidae) and Possible Role of Temperature Increase

In some fishes, water chemistry or temperature affects sex determination or creates sex‐specific selection pressures. The resulting population sex ratios are hard to predict from laboratory studies if the environmental triggers interact with other factors, whereas in field studies, singular observations of unusual sex ratios may be particularly prone to selective reporting. Long‐term monitoring largely avoids these problems. We studied a population of grayling (Thymallus thymallus) in Lake Thun, Switzerland, that has been monitored since 1948. Samples of spawning fish have been caught about 3 times/week around spawning season, and water temperature at the spawning site has been continuously recorded since 1970. We used scale samples collected in different years to determine the average age of spawners (for life‐stage specific analyses) and to identify the cohort born in 2003 (an extraordinarily warm year). Recent tissue samples were genotyped on microsatellite markers to test for genetic bottlenecks in the past and to estimate the genetically effective population size (N_e). Operational sex ratios changed from approximately 65% males before 1993 to approximately 85% males from 1993 to 2011. Sex ratios correlated with the water temperatures the fish experienced in their first year of life. Sex ratios were best explained by the average temperature juvenile fish experienced during their first summer. Grayling abundance is declining, but we found no evidence of a strong genetic bottleneck that would explain the apparent lack of evolutionary response to the unequal sex ratio. Results of other studies show no evidence of endocrine disruptors in the study area. Our findings suggest temperature affects population sex ratio and thereby contributes to population decline. Persistencia de Proporción de Sexos Desigual en una Población de Tímalos (Salmonidae) y el Posible Papel del Incremento de la Temperatura     

水体低氧的早期暴露对青鳉(Oryzias latipes)后期的生长、性别比和繁殖能力的影响

水体低氧已是全球性生态问题,常以季节性、偶发性和昼夜间等不同形式存在于不同的水体中。长期低氧可影响鱼类正常的生长和繁殖,但鱼类早期生活阶段暴露于不同形式的低氧后,后期的生长和繁殖是否会受到不利影响,目前研究甚少。本研究在实验室模拟了连续低氧(2.8 mg·L~(-1)DO)(H1)、昼夜低氧(H2)和发生在胚胎器官形成时期的偶发性低氧(H3)等3种情景对青鳉胚胎的发育影响,评估了这一早期暴露对青鳉后期的生长、存活和繁殖的影响。我们发现,3种低氧方式都可以显著延长青鳉胚胎的孵化周期,引起胚胎卵黄囊吸收和鱼鳔发育异常;暴露结束120 d后,H1组青鳉成鱼的畸形率显著升高、存活率和生长速度都显著下降;H1、H2和H3组中成鱼的雌雄比都发生了改变,鱼群中以雄鱼为主,且产卵量和受精率都显著下降。结果表明,鱼类早期胚胎发育阶段所受到的低氧暴露可对后期生长和繁殖产生不利影响,对子代补充和种群稳定产生重要影响;鱼类关键发育期所经历的低氧事件,以及昼夜低氧事件所产生的生态后果不容忽视。     

Primary sex ratios in zebra finches: no evidence for adaptive manipulation in wild and semi-domesticated populations.

When breeding diet is restricted, domesticated zebra finches,Taeniopygia guttata, produce male-biased primary and secondary sex ratios, but unexpectedly produce unbiased ratios when food is unrestricted. We investigated the primary sex ratios (at laying) of wild zebra finches in southeastern Australia in response to food supplementation and environmental factors predicted to enhance female breeding condition and to bias the primary sex ratio towards daughters. Molecular sexing of all nestlings in 54 complete broods where every egg hatched, failed to show any significant biases from random. Time of egg laying (month, season) and environmental conditions (rainfall, temperature) did not significantly predict variation in the primary sex ratio, but time of breeding did affect clutch size. Wild zebra finches at our colony did not bias their sex allocation as there were no differences in the primary sex ratio and no differences in the numbers and mass of sons and daughters at the end of parental care (day 35–40 post-hatch). Biases in primary sex ratio of our wild population are probably weak or non-existent possibly due to the unpredictable environment and/or multiple contrary selective forces acting on sex ratios. We also investigated the effects of photoperiod, biases in the adult sex ratio, and parental attractiveness on primary sex ratios of semi-domesticated, laboratory zebra finches. Molecular sexing of three-day old embryos from complete clutches, failed to reveal significant biases from random. In contrast to previous studies, sex of eggs did not correlate with laying order and egg mass declined with order, rather than increased. Domestication may be responsible for these differences.     

Reproductive plasticity,oviposition site selection,and maternal effects in fragmented landscapes

Traditionally, evolutionary ecology and conservation biology have primarily been concerned with how environmental changes affect population size and genetic diversity. Recently, however, there has been a growing realization that phenotypic plasticity can have important consequences for the probability of population persistence, population growth, and evolution during rapid environmental change. Habitat fragmentation due to human activities is dramatically changing the ecological conditions of life for many organisms. In this review, we use examples from the literature to demonstrate that habitat fragmentation has important consequences on oviposition site selection in insects, with carryover effects on offspring survival and, therefore, population dynamics. We argue that plasticity in oviposition site selection and maternal effects on offspring phenotypes may be an important, yet underexplored, mechanism by which environmental conditions have consequences across generations. Without considering the impact of habitat fragmentation on oviposition site selection, it will be difficult to assess the effect of fragmentation on offspring fitness, and ultimately to understand the impact of anthropogenic-induced environmental change on population viability.     

Unprecedented Low Levels of Genetic Variation and Inbreeding Depression in an Island Population of the Black-Footed Rock-Wallaby

Abstract: It has been argued that demographic and environmental factors will cause small, isolated populations to become extinct before genetic factors have a significant negative impact. Islands provide an ideal opportunity to test this hypothesis because they often support small, isolated populations that are highly vulnerable to extinction. To assess the potential negative impact of isolation and small population size, we compared levels of genetic variation and fitness in island and mainland populations of the black-footed rock-wallaby ( Petrogale lateralis [Marsupialia: Macropodidae]). Our results indicate that the Barrow Island population of P. lateralis has unprecedented low levels of genetic variation (  H _e = 0.053, from 10 microsatellite loci) and suffers from inbreeding depression (reduced female fecundity, skewed sex ratio, increased levels of fluctuating asymmetry). Despite a long period of isolation ( ∼ 1600 generations) and small effective population size (  N _e ∼ 15), demographic and environmental factors have not yet driven this population to extinction. Nevertheless, it has been affected significantly by genetic factors. It has lost most of its genetic variation and become highly inbred (  F _e = 0.91), and it exhibits reduced fitness. Because several other island populations of P. lateralis also exhibit exceptionally low levels of genetic variation, this phenomenon may be widespread. Inbreeding in these populations is at a level associated with high rates of extinction in populations of domestic and laboratory species. Genetic factors cannot then be excluded as contributing to the extinction proneness of small, isolated populations.     

Erosion of Heterozygosity in Fluctuating Populations

Abstract: Demographic, environmental, and genetic stochasticity threaten the persistence of isolated populations. The relative importance of these intertwining factors remains unresolved, but a common view is that random demographic and environmental events will usually drive small populations to the brink of extinction before genetic deterioration poses a serious threat. To evaluate the potential importance of genetic factors, we analyzed a model linking demographic and environmental conditions to the loss of genetic diversity in isolated populations undergoing natural levels of fluctuation. Nongenetic processes—environmental stochasticity and population demography—were modeled according to a bounded diffusion process. Genetic processes were modeled by quantifying the rate of drift according to the effective population size, which was predicted from the same parameters used to describe the nongenetic processes. We combined these models to predict the heterozygosity remaining at the time of extinction, as predicted by the nongenetic portion of the model. Our model predicts that many populations will lose most or all of their neutral genetic diversity before nongenetic random events lead to extinction. Given the abundant evidence for inbreeding depression and recent evidence for elevated extinction rates of inbred populations, our findings suggest that inbreeding may be a greater general threat to population persistence than is generally recognized. Therefore, conservation biologists should not ignore the genetic component of extinction risk when assessing species endangerment and developing recovery plans.     

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.     

Effects of temperature variation on TSD in turtle (C. picta) populations

We formulate a two-sex model of temperature-dependent sex determination (TSD) for a freshwater turtle (C. picta) population. The aim is to understand how environmental temperature variations and nest heat conduction properties affect the long term dynamics of the population. This is a key to understanding how global temperature changes may affect their survival. With stochastic inputs of ambient temperature and solar radiation, the model uses the heat equation to determine the temperature in the egg layer in the nest; in turn, this determines the sex ratio in the egg clutch using a variable degree-day model. Finally, a nonlinear Leslie type, stage-based, two-sex model, is used to determine the long term male and female populations. A two-sex model is required because of different development rates for males and females. The model is flexible enough to enable other researchers to examine the effects of temperature variation variations on other species with TSD, e.g., crocodilians, reptilians, as well as other turtle species. It can be adapted to study effects of nest location, soil type, rain events, different incubation periods, and density effects, for example, the dependence of the mating function on the ratio of males to females and each’s contribution to the sex of hatchlings. Modifications can be easily made to fit a specific life history traits. The model is a beginning step in understanding the long term, high fitness shown by many reptile species with TSD, and it may suggest to experimentalists what data may be relevant to these issues; it can also be useful to wildlife managers in developing strategies for intervention if needed. Among the principal findings are that temperature variability and detailed nest heat conduction properties may buffer projected negative effects on a population.     

Phylogenetic Comparative Methods Strengthen Evidence for Reduced Genetic Diversity among Endangered Tetrapods

Abstract: The fitness of species with little genetic diversity is expected to be affected by inbreeding and an inability to respond to environmental change. Conservation theory suggests that endangered species will generally demonstrate lower genetic diversity than taxa that are not threatened. This hypothesis has been challenged because the time frame of anthropogenic extinction may be too fast to expect genetic factors to significantly contribute. I conducted a meta‐analysis to examine how genetic diversity in 894 tetrapods correlates with extinction threat level. Because species are not evolutionarily independent, I used a phylogenetic regression framework to address this issue. Mean genetic diversity of tetrapods, as assessed by protein heterozygosity, was 29.7–31.5% lower on average in threatened species than in their nonthreatened relatives, a highly significant reduction. Within amphibians as diversity decreased extinction risk increased in phylogenetic models, but not in nonphylogenetic regressions. The effects of threatened status on diversity also remained significant after accounting for body size in mammals. These results support the hypothesis that genetic effects on population fitness are important in the extinction process.     

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.     

Climate and maternal effects modify sex ratios in a weakly dimorphic marsupial

There is growing evidence that the sex ratios of wild vertebrate populations are determined by mechanisms that are directly influenced by environmental characteristics. The Trivers–Willard (TWH) and extrinsic modification (EMH) hypotheses postulate differing determinants of mammalian offspring sex ratios. TWH states that mothers allocate resources according to their current condition and sex-specific offspring costs. EMH states that environmental forces that affect maternal condition determine offspring sex ratios, independently of maternal tactics of sex-biased allocation. We statistically assessed support for each of these hypotheses using long-term life histories of the allied rock-wallaby, Petrogale assimilis; a continuously breeding, polygynous, weakly dimorphic marsupial. We showed that birth sex ratios were equal and independent of maternal and environmental conditions. However, secondary sex ratios were male-biased under good environmental conditions and for high quality mothers or mothers in good condition. Sex differences in offspring survival contributed to these biases: (1) environmental conditions strongly influenced survival to pouch emergence (in support of EMH) and (2) maternal quality affected survival to the end of maternal care (in support of TWH). Environmental effects on survival were more important than maternal factors over the entire period of maternal care and contributed most to male-biased sex ratios at pouch emergence. In contrast, maternal mass was the best predictor of sex ratios at the end of maternal care—the life history stage where offspring body mass differed between the sexes.     

What stops the ‘diploid male vortex’?—A simulation study for species with single locus complementary sex determination

The survival of hymenopteran populations especially that of ecologically and economically important pollinators and parasitoids, has become a major topic for empirical and theoretical studies. Complementary sex determination (CSD) in Hymenoptera may impose a substantial genetic load through the production of inviable or sterile diploid males. Modelling and laboratory studies have indicated that this genetic load may trigger a ‘diploid male vortex’ leading to rapid extinction of populations.Here we take a broader theoretical approach to analyze why populations exhibiting CSD persist in nature, even if they contain large proportions of homozygous diploid individuals. Using an individual-based model of spatially structured hymenopteran populations, we show that (i) inviability or reduced fertility of homozygous individuals, (ii) female-biased sex ratio, and (iii) strong intra-specific competition can mitigate the negative influence of CSD on population persistence and that (iv) already extremely low dispersal rates will result in long term survival. These findings underline the importance of life history traits for population survival and demonstrate that rather specific conditions must be met to initiate the ‘diploid male vortex’.     

The impact of behavioral and physiological maternal effects on offspring sex ratio in the common snapping turtle,<Emphasis Type="Italic"> Chelydra serpentina</Emphasis>

Theory suggests that maternal effects are especially important in organisms with environmentally-sensitive sex-determining mechanisms. However, there is no substantive body of empirical evidence to confirm this conjecture. We integrated field and laboratory studies to jointly evaluate the significance of behavioral (nest-site choice) and physiological (yolk hormone allocation) maternal effects on offspring sex ratio in the common snapping turtle (Chelydra serpentina), a species with temperature-dependent sex determination (TSD). Of the 16 microhabitat variables measured, only three (south, east, and total overstory vegetation cover) were significantly correlated with nest temperature: cooler nests were located under more vegetation cover. In turn, these microhabitat predictors of nest temperature, and nest temperature itself, may influence nest sex ratio: shadier, cooler nests were more likely to produce a higher proportion of male offspring than less shady, warmer nests. Analysis of eggs from these same nests incubated in a common garden design in the laboratory revealed that clutch sex ratio was unaffected by levels of yolk estradiol, yolk testosterone, or their interaction. Examination of both behavioral and physiological maternal effects revealed no concordant impact on offspring sex ratio. However, eggs from nests that produced male-biased sex ratios in the field yielded higher proportions of males under constant-temperature conditions in the laboratory. Our study confirms the importance of behavioral maternal effects in nature on offspring sex ratios in species with TSD, while also revealing the potential presence of a predisposition for sex-ratio production underlying TSD in this system.Communicated by S. Krackow     

Prioritizing Pacific Salmon Stocks for Conservation

Over 300 native stocks of Pacific salmon, steelhead, and coastal cutthroat trout (Oncorhynchus spp.) are at risk of extinction in the Pacific Northwest. With only limited resources available for conservation and recovery, prioritization of these stocks may become necessary if meaningful measures are to be implemented. We propose criteria by which prioritization may be guided. First, we rank stocks for risk of extinction, either by population viability analysis or by a set of surrogate measures. Then we rank stocks for biological consequences of extinction, using sets of questions designed to establish the genetic and evolutionary consequences and the ecological consequences if a stock were to become extinct. Together, these rankings allow stocks to be prioritized for a range of possible actions, with those stocks at highest risk and bearing the greatest biological consequences of extinction receiving attention first. Application of the prioritization process to 20 Pacific anadromous salmonid stocks worked as intended, although data limitations are considerable. The process is most likely to work successfully when applied to many stocks on which data exist, when several experts carry out the prioritization, and when the results are peer reviewed.     

环境内分泌干扰物对鱼类性别分化的影响

鱼类性别分化是在性别决定机制的控制下,鱼类未分化的性腺向卵巢或精巢发育,并出现第2性征的过程,受自身内分泌系统的精确调控。环境内分泌干扰物能通过干扰内源激素的合成、释放、转运、代谢、结合和作用等过程影响鱼类的内分泌机能,从而对鱼类的性别分化产生影响。介绍了鱼类性别分化的模式和过程以及内分泌系统对鱼类性别分化的调控作用,综述了环境内分泌干扰物对鱼类性别比例、配子发生、性腺发育和第2性征的影响,从性类固醇激素的合成以及性别分化相关基因的表达两个方面探讨了环境内分泌干扰物影响鱼类性别分化的作用机制,并展望了该领域未来的研究方向。     

Karyotype and sex determination in <Emphasis Type="Italic">Dinophilus gyrociliatus</Emphasis> (Polychaeta: Dinophilidae)

The study of the male and female karyotypes of the polychaete Dinophilus gyrociliatus was performed with the aim of ascertaining the diploid number of chromosomes and confirming the existence of karyological differences between the two sexes. Our results evidenced that the female karyotype of D. gyrociliatus consists of 2n=24 chromosomes, while the male karyotype has 2n=23 chromosomes. X chromosomes are subtelocentric and much bigger than the autosomes. The size of the D. gyrociliatus chromosomes, which in the female embryos vary in length from little more than 2.8 wm to less than 0.7 wm, are relatively small by comparison with those of other polychaetes. The existence of maternal and environmental factors able to influence the sex ratio of D. gyrociliatus and chromosomal differences between the sexes confirms that: (1) sex determination is chromosomal and syngamic of the XX-X0 type and (2) control of the sex ratio is progamic and depends on genetic and environmental factors.     

Effects of Social Structure and Prey Dynamics on Extinction Risk in Gray Wolves

Extinction models based on diffusion theory generally fail to incorporate two important aspects of population biology—social structure and prey dynamics. We include these aspects in an individual-based extinction model for small, isolated populations of the gray wolf (Canis lupus). Our model predicts mean times to extinction significantly longer than those predicted by more general (diffusion) models. According to our model, an isolated population of 50 wolves has a 95% chance of surviving just 9 years and only a 30% chance of surviving beyond 100 years. Reflecting the influence of social structure, a wolf population initially comprising 50 individuals is expected to persist only a few years longer, on average (71 years), than is a population initially comprising just a single reproductive pair (62 years). In contrast, substantially greater average prey abundance leads to dramatically longer expected persistence times. Autocorrelated prey dynamics result in a more complex distribution of extinction times than predicted by many extinction models. We contend that demographic stochasticity may pose the greatest threat to small, isolated wolf populations, although environmental stochasticity and genetic effects may compound this threat. Our work highlights the importance of considering social structure and resource dynamics in the development of population viability analyses.     

Possible Extinction Vortex for a Population of Iberian Lynx on the Verge of Extirpation

Abstract: Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female‐biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co‐occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.     

Temperature,age of mating and starvation determine the role of maternal effects on sex allocation in the mealybug <Emphasis Type="Italic">Planococcus citri</Emphasis>

Environmental effects on sex allocation are common, yet the evolutionary significance of these effects remains poorly understood. Environmental effects might influence parents, such that their condition directly influences sex allocation by altering the relative benefits of producing sons versus daughters. Alternatively, the environment might influence the offspring themselves, such that the conditions they find themselves in influence their contribution to parental fitness. In both cases, parents might be selected to bias their sex ratio according to the prevailing environmental conditions. Here, we consider sex allocation in the citrus mealybug Planococcus citri, a species with an unusual genetic system in which paternal genes are lost from the germline in males. We test environmental factors that may influence either female condition directly (rearing temperature and food restriction) or that may be used as cues of the future environment (age at mating). Using cytological techniques to obtain primary sex ratios, we show that high temperature, older age at mating and starvation all affect sex allocation, resulting in female-biased sex ratios. However, the effect of temperature is rather weak, and food restriction appears to be strongly associated with reduced longevity and a truncation of the usual schedule of male and offspring production across a female’s reproductive lifetime. Instead, facultative sex allocation seems most convincingly affected by age at mating, supporting previous work that suggests that social interactions experienced by adult P. citri females are used when allocating sex. Our results highlight that, even within one species, different aspects of the environment may have conflicting effects on sex allocation.