Large Female Body Size Research Articles

Testing ‘bigger is better’ and maternal effects hypotheses in hatchlings of the sexually dimorphic spiny softshell turtle (Apalone spinifera)

Abstract The ‘bigger is better’ hypothesis (BIBH) predicts that fitness increases with body size. Eastern spiny softshell turtles (Apalone spinifera) exhibit sexual size dimorphism (SSD) at maturity, and the much larger female body size is attributed to selection favouring greater reproductive output. Hatchlings exhibit sexually dimorphic markings, but hatchling SSD has not been investigated. The BIBH predicts that hatchlings will not exhibit SSD because reproductive benefits to larger, mature females drive size dimorphism at maturity, but selection on hatchlings relates only to survival. We tested this prediction with A. spinifera hatchlings (N = 2630) incubated in controlled conditions to avoid confounding environmental effects on phenotype. Hatchling mass was similar between sexes, suggesting similar nutritional allocation to male and female offspring. Hatchling size (carapace length) was also similar between sexes. Hatchling mass and body size varied among clutches and were positively related to maternal size, suggesting morphological constraints on egg size. Surprisingly, clutch size was related only weakly to maternal size. Taken together, our results suggest that hatchling size in A. spinifera is consistent with the BIBH and that offspring quality increases with maternal age, but that maternal effects drive a large proportion of the variation in clutch size and offspring quality.

Larger is not better: no mate preference by European common frog (Rana temporaria) males

Abstract In explosive breeding frogs, high intrasexual competition between males leads to a sexual coercion ruled mating system, where males presumably evolved preferences for specific female traits. We tested these preferences in the European Common Frog by excluding intrasexual competition. We hypothesized that all males show preferences towards larger female body size, due to higher fecundity. Our results did not show any preference considering female body size, neither in the attempt to amplex a female nor during the formation of pairs. Additionally, we witnessed a high failure rate of male mating attempts, which hints at high mating costs and offers an explanation for the lack of preferences in males. Nonetheless, we observed a non-random mating pattern in successfully formed pairs, where in the absence of size dimorphism females were on average larger than males. This indicates a different mechanism for selection which is independent from male mating preference or scramble competition.

Morphological variability in Lophyra flexuosa (Fabricius, 1787) (Coleoptera, Cicindelidae) in desert countries is affected by sexual dimorphism and geographic aspect.

Lophyra flexuosa, a eurytopic tiger beetle characterized by long phenological activity, wide geographic and altitudinal distribution, and occurring in the highest number of habitats among all Cicindelidae known from North Africa, was chosen to study its geographic variation in morphology and sexual dimorphism. Here, we found significant sexual dimorphism exhibited in larger body size of females and longer mandibles in males, which can be explained by different roles of particular sexes in courtship. Moreover, we recorded significant differences in body sizes between western and eastern Maghreb populations which could suggest genetic isolation between these populations. As the species is related to habitats placed close to the water reservoirs, which in the desert countries are under significant human pressure (including climate change), we expect a reduction of habitat occupied by this taxon. Therefore, the geographic morphological variability that we observe today in the tiger beetle Lophyra flexuosa in the future could lead to speciation.

Differential outcomes of novel plant-herbivore associations between an invading planthopper and native and invasive Spartina cordgrass species

Non-native plants may benefit, briefly or permanently, from natural enemy release in their invaded range, or may form novel interactions with native enemy species. Likewise, newly arrived herbivores may develop novel associations with native plants or, where their hosts have arrived ahead of them, re-establish interactions that existed previously in their ancestral ranges. Predicting outcomes from this diversity of novel and re-established interactions between plants and their herbivores presents a major challenge for invasion biology. We report on interactions between the recently arrived invasive planthopper Prokelisia marginata, and the multi-ploidy Spartina complex of four native and introduced species in Britain, each representing a different level of shared evolutionary history with the herbivore. As predicted, S. alterniflora, the ancestral host, was least impacted by planthopper herbivory, with the previously unexposed native S. maritima, a nationally threatened species, suffering the greatest impacts on leaf length gain, new leaf growth and relative water content. Contrary to expectations, glasshouse trials showed P. marginata to preferentially oviposit on the invasive allododecaploid S. anglica, on which it achieved earlier egg hatch, faster nymphal development, larger female body size and greatest final population size. We suggest P. marginata is in the process of rapid adaptation to maximise its performance on what is now the most abundant and widespread host in Britain. The diversity of novel and re-established interactions of the herbivore with this multi-ploidy complex makes this a highly valuable system for the study of the evolutionary ecology of plant–insect interactions and their influence on invasion dynamics.

Reproduction in the pitviper Bothrops jararacussu: large females increase their reproductive output while small males increase their potential to mate

Reproductive traits vary widely within and among snake species, and are influenced by a range of factors. However, additional studies are needed on several species, especially on tropical snake faunas, to fully understand the patterns of reproductive variation in snakes. Here, we characterized the reproductive biology of B. jararacussu from southeastern and southern Brazil. We combined macroscopic and microscopic examinations of the reproductive system of museum specimens with observations of free-ranging snakes to characterize size at sexual maturity, sexual size dimorphism (SSD), reproductive output, and male and female reproductive cycles. We compared our data with published literature and discuss the factors that may play a role in shaping the reproductive patterns in the species and the genus. Bothrops jararacussu shares several characteristics with its congeners such as autumn mating season, obligatory sperm storage in the female reproductive tract, seasonal timing of parturition (summer-autumn), female-biased SSD, maturity at larger body sizes in females, and a positive relationship between body size and litter size. These characteristics seem phylogenetically conserved in Bothrops. On the other hand, B. jararacussu exhibits some unique characteristics such as a high degree of SSD (one of the highest values recorded in snakes), a large female body size, and a large litter and offspring size, which are among the largest recorded in the genus. Moreover, larger females reproduce more frequently than smaller conspecifics. These characteristics may be collectively interpreted as the result of a strong selection for increased fecundity. Other peculiarities of the species include an asynchrony between spermiogenesis (summer-autumn) and the peak of SSK hypertrophy (autumn to spring) and a prolonged production of SSK granules. Because SSK hypertrophy and mating are androgen-dependent in snakes, the prolonged SSK hypertrophy suggests that male B. jararacussu may prolong their potential to mate (compared with its congeners), which may increase their reproductive success. Our results and previous literature collectively suggest that, in Bothrops, the evolution of SSD is driven by fecundity selection, variation in reproductive output is influenced by variation in female body size, and the timing of spermiogenesis is influenced by other factors in addition to temperature. We also suggest that male Bothrops have undergone multiple evolutionary shifts in the timing of spermiogenesis.

Reduced sexual size dimorphism in a pipefish population where males do not prefer larger females.

Within a species' distribution, populations are often exposed to diverse environments and may thus experience different sources of both natural and sexual selection. These differences are likely to impact the balance between costs and benefits to individuals seeking reproduction, thus entailing evolutionary repercussions. Here, we look into an unusual population (Baltic Sea) of the broadnosed pipefish, Syngnathus typhle, where males do not seem to select females based on size and hypothesize that this pattern may derive from a reduction in direct benefits to the male. We further hypothesize that if larger females do not persistently secure a higher reproductive success, either through pre‐ or postcopulatory sexual selection, a decrease in sexual size dimorphism in the Baltic population should be apparent, especially when contrasted with a well‐studied population, inhabiting similar latitudes (Swedish west coast), where males prefer larger females. We found that, in the Baltic population, variation in female quality is low. We were unable to find differences in abortion rates or protein concentration in oocytes produced by females of contrasting sizes. Direct benefits from mating with large partners seem, thus, reduced in the Baltic population. We also found no evidence of any postcopulatory mechanism that could favor larger mothers as embryo development was unrelated to female size. While female size can still be selected through intrasexual competition or fecundity selection, the pressure for large female body size seems to be lower in the Baltic. Accordingly, we found a noticeable decrease in sexual size dimorphism in the Baltic population. We conclude that, although far from negating the significance of other selective processes, sexual selection seems to have a decisive role in supporting pipefish sexual size asymmetries.

Gopher Tortoise Demographic Responses to a Novel Disturbance Regime

ABSTRACTThe long‐term viability of gopher tortoise (Gopherus polyphemus) populations is jeopardized by increased urbanization and habitat degradation owing to fire suppression. Because the species' remaining natural habitats in the southeastern United States exist within a mosaic of anthropogenic land uses, it is important to understand demographic responses to contrasting land uses and habitat management regimes. We examined differences in demographic parameters among fire‐suppressed sandhill, restored sandhill, and former sandhill (i.e., ruderal) land use‐land cover (LULC) types at Archbold Biological Station in south‐central Florida, USA. Using Program MARK, we estimated population size, and sex‐specific and LULC‐specific survivorship based on 6 years of mark‐recapture data. We also analyzed individual growth trajectories and clutch sizes to determine whether growth rates or reproductive output differed among LULC types. Tortoises in an open, ruderal field occurred at a higher density (7.79/ha) than in adjacent restored (1.43/ha) or fire‐suppressed (0.40/ha) sandhill. Despite this higher density, both adult survivorship and body size were significantly higher in the ruderal field. Furthermore, the larger female body size in the ruderal field likely contributed to increased annual survivorship and slightly larger average clutch sizes. We did not detect offsetting negative demographic effects; in particular, we did not find significant biological or statistical differences in body condition, asymptotic body size, or growth rate among the 3 LULC types. Our results suggest that anthropogenic, grass‐dominated land‐cover types may be important components of the habitat mosaic currently available to this at‐risk species. © 2019 The Wildlife Society.

Body size in tiger beetles (Coleoptera: Cicindelidae) in the south-eastern European biodiversity hotspot: sexual dimorphism and patterns of co-occurrence

In south-eastern Europe, especially in lowland areas located close to the sea coast, the diversity of Cicindelidae is one of the highest in the entire Palaearctic realm. This is both because of geological history of the region and the presence of large mosaic of habitats attractive for these beetles. As a result in the region about 25% of all Palaearctic species of tiger beetles can be found, including endemics. Many of them can be characterized by narrow or very narrow habitat preferences occurring only in one or two types of macrohabitats. In the present study sexual dimorphism of six species of tiger beetles (Calomera littoralis, C. fischeri, Cephalota circumdata, C. chiloleuca, Cylindera trisignata trisignata, Myriochila melancholica) collected along the sea coast of the Black and Mediterranean Seas was studied based on eight body parameters (right mandible length, length of head, width of head, length of pronotum, maximum pronotum width, length of elytra, maximum elytra width, and total body length.). Moreover, patterns of species co-occurrence on the basis of body size were investigated. As results we found significant sexual dimorphism indicated by larger body size of females and longer mandibles in males of the studied Cicindelidae species, what can be explained by different roles of the particular sexes in courtship (females invest much more energy in the reproduction process than males, e.g. to produce eggs or to find a good place to deposit them, as a result bigger size is much more beneficial for this sex). Moreover, we discovered that in the studied area tiger beetle species characterized by similar body size avoid each other and do not occur in the same areas.

Lophyra flexuosa (Fabricius, 1787) (Coleoptera: Cicindelidae) in desert countries: morphological variability in geographic aspect as potential beginning of speciation?

Lophyra flexuosa is one of only several eurytopic tiger beetles species known from Palearctic realm. Its geographical distribution shows several populations that are spread from the Iberian Peninsula and Morocco, across some regions of south-western Europe and northern Africa to Israel and Syria. The species is characterized by long phenological activity, wide altitudinal distribution, and occurs in the highest number of habitats among all Cicindelidae known from Maghreb region. In the present study the geographical variation in morphology and sexual dimorphism in north African populations of L. flexuosa was studied. In total 52 samples with over 700 specimens were collected including 20 samples in Morocco and 32 in Tunisia. To test the variation in morphometric traits measurements of eight body parameters were taken from all males (383) and females (352) including right mandible length, length of head, width of head, length of pronotum, maximum pronotum width, length of elytra, maximum elytra width, and total body length. We discovered significant sexual dimorphism expressed by larger body size of females and longer mandibles in males, what can be explained by different roles of particular sexes in courtship. Moreover, we recorded significant differences in body sizes between western and eastern Maghreb populations which can suggest genetic isolation between these populations. As the species is related to habitats placed close to water reservoirs, which in the desert countries are under significant human pressure (including climate change), we expect an reduction of habitats occupied by this taxon. Therefore, the geographic morphological variability that we observe today in the tiger beetle Lophyra flexuosa may lead to speciation and creation of separate species in the future.

Ontogeny of sexual size dimorphism revisited: Females grow for a longer time and also faster.

Sex-specific mechanisms of the determination of insect body sizes are insufficiently understood. Here we use the common heath moth, Ematurga atomaria (Lepidoptera: Geometridae) to examine how larval growth trajectories differ between males and females. We monitored the development of 1379 larvae in controlled laboratory conditions. Sexually dimorphic development times during the first four instars were associated with sexual size dimorphism (SSD) in the beginning of the fifth (last) instar, when females were on average 15% heavier than males. Similarly, the duration of the last instar was about 13% longer in females. Further, we specifically focussed on the estimates of differential (instantaneous) growth rates of the larvae based on 24h mass increments of the 2nd, 3rd, 4th and 5th day in the beginning of the last instar. We calculated ‘allometric’ differential growth rates as the per-day increase in cube-root-transformed mass of the larvae. We found that allometric growth rates were slightly but significantly larger in females than in males. As this measure of growth rate (in contrast to the relative growth rate, based on the ratio of masses recorded at consecutive measurements) did not depend on body size, it allows an unambiguous separation of the effects of sex and size. We conclude that in accordance with an emerging general pattern, larger female body size in E. atomaria is achieved primarily by means of a longer growth period. Furthermore, our study shows that the differential growth rate can also be sexually dimorphic and contribute to SSD. This contribution, however, is lower than that of the development time by an order of magnitude. In addition to development periods and growth rates, other parameters of the non-linear growth curves of insect larvae also need to be considered in the context of SSD determination. In particular, weight loss prior to pupation was shown to be considerably larger in females than in males.

Assortative Mating by Size in the American Rubyspot Damselfly (Hetaerina americana)

Assortative mating refers to the non-random nature of mating patterns between certain males and females. Thus, males and females may associate negative- or positively, based on different traits. Amongst these associations, assortative mating by size is one of the most common patterns found in natural populations of animals. Two main hypotheses have been proposed to account for the occurrence of assortative mating by size. First, it may be the result of mechanical, temporal, or physiological constraints. Second, it may occur in response to direct or indirect selection on mating preferences. Here we investigate whether the American rubyspot damselfly exhibits true assortative mating by size. Males of this species exhibit high levels of male-male competition, as they compete over territories, to which females are attracted for copulation. There is a documented large male body size advantage: the largest males are better able to hold their territories and thus secure more copulations. Our major results show that i) mated males are more likely to be larger than unmated males, whereas mated and unmated females tend to have similar body sizes; ii) H. americana exhibits true assortative mating by size; as such, this pattern is not driven by seasonal changes in the body sizes of males and females. We suggest that this mating pattern occurs in this species given the advantages of large male size, and the advantages of large female body size (i.e. higher fecundity). We believe that males may be able to evaluate a female’s reproductive value and exert mate choice.

Random size‐assortative mating despite size‐dependent fecundity in a Neotropical amphibian with explosive reproduction

AbstractSexual selection theory predicts that, when body size is correlated with fecundity, there should be fitness advantages for mate choice of the largest females. Moreover, because larger males are expected to monopolise the largest females, this should result in an assortative mating based on body size. Although such patterns could be expected in both explosive and prolonged breeders, non‐assortative mating should be more widespread in species under time constraints. However, patterns of sexual selection are largely unexplored in explosive breeding species, and contrasting patterns have been found previously. We expect that the active choice of partners may be particularly risky when the time period during which sexual partners are available is severely limited. Therefore, to avoid missing an entire reproductive act, males and females should pair irrespective of traits, such as body size. We tested this hypothesis by investigating the mating patterns of the Pacific horned toad, Ceratophrys stolzmanni, a short‐lived fossorial species inhabiting Neotropical dry forests. This species is particularly adequate to test our prediction because it reproduces explosively over the course of a single night per year. Although the number of eggs laid was proportional to the size of females, and individuals of both sexes showed variation in body size, there was no assortative mating based either on size, body condition or age of mates. Egg size was not influenced by either female size or clutch size. The larger body size of females compared to males is likely due to fecundity selection, that is, the selective pressure that enhances reproductive output. Although we cannot dismiss the possibility that individuals could select their partners based on other criteria than those related to size or age, the results fit well our prediction, showing that the explosive breeding makes improbable an active choice of partners in both sexes and therefore favours a random mating pattern.

Sex-specific differences in growth, mortality and migration support population resilience in the heavily exploited migratory marine teleost Mugil cephalus (Linnaeus 1758)

Understanding mechanisms supporting the resilience of exploited fish populations is fundamental to sustainable management. Herein we identify sex-specific differences in life history traits that confer resilience in the heavily exploited population of Mugil cephalus along eastern Australia. M. cephalus in this region emigrate from estuaries and undergo an annual northerly spawning migration that is dominated by males. Males mature, on average, at younger ages than females and were most abundant in the spawning migration at age-4, whereas females were most abundant at age-5. Females grew significantly faster and larger than males, with both sexes being aged up to 14 years. These sex-specific differences relate directly to the population stability of M. cephalus under the reproductive strategy of a spawning migration. Males participate in the spawning migration at younger ages and in greater numbers than females to ensure a sufficient number of males for successful spawning. Females partition more energy to growth than males, maturing and participating in the spawning migration at older ages and larger sizes. The larger body size of females is related to increased fecundity, increased survivorship and more efficient swimming ability. These traits, combined with population strategies of both delayed and skipped spawning, convey substantial resilience.

Comparative development and reproduction ofPlanococcus ficusandPlanococcus citri(Hemiptera:Pseudococcidae) on grapevine under field conditions

AbstractMealybugs are major pests in grape‐growing areas worldwide, causing direct and indirect crop damage. The vine mealybugPlanococcus ficus(Signoret) is a key pest in most of grape‐producing countries, whereas the citrus mealybugPlanococcus citri(Risso) is reported as being destructive inBrazilian andSpanish vineyards.We examined the adaptation of the citrus mealybug to grapevine by investigating its development, reproduction and life‐history parameters underMediterranean field conditions in comparison with those ofP. ficus.Both mealybug species developed and reproduced successfully on grapevine. However,P. ficusshowed a shorter development time, larger female body size, and higher fecundity, fertility and survival thanP. citri.The life‐history parameters further highlighted the different growth potential of vine and citrus mealybug populations becauseP. ficusexhibited a net reproductive rate and an intrinsic rate of increase two‐ to four‐fold higher than that ofP. citri. Furthermore, the vine mealybug population doubled its number in approximately half time with respect to the citrus mealybug.Overall, the results of the present study show a better development and reproductive performance and a higher population growth potential ofP. ficuscompared withP. citri, indicating a higher capacity of the vine mealybug to develop on grapevine inMediterranean vineyards.

A uni- and multivariate analysis approach to reveal sexual size dimorphism in Iranian populations of Hypera postica (Coleoptera: Curculionidae)

Abstract Body size dimorphism between genders is a commonly observed phenomenon in insects, usually manifested in larger female body size. Sexual Size Dimorphism (SSD) varies from species to species, the degree and direction influenced by certain evolutionary pressures. Intraspecific variation in SSD may also occur between populations. The Hypera postica (Gyllenhal, 1813) is a well-known alfalfa plant pest that shows a degree of morphological divergence in its populations. The female alfalfa weevils are very fecund and have a larger body size compared to males. To improve our knowledge on magnitude and direction of SSD in alfalfa weevil, we studied 200 specimens of H. postica from four Iranian populations (Karaj1, Karaj2, Tuyserkan and Jovein). 10 morphological variables from three external anatomic parts (pronotum, elytra and rostrum) and 45 ratio characters were statistically analyzed in order to determine the amount of SSD in Iranian populations. In addition we investigated for morphological divergence pattern in mentioned populations. The results of this study show that a low degree of morphological divergence occurs in Iranian populations. Measured variables indicate that the SSD pattern of H. postica is compatible with the Rensch’s rule, and is related to high fecundity of females and a lack of strong sexual selection. Also it is mentioned that the larger rostrum in females may correspond to its unique role in egg laying.

The evolution of fecundity is associated with female body size but not female-biased sexual size dimorphism among frogs.

Sexual size dimorphism (SSD) is one of the most common ways in which males and females differ. Male-biased SSD (when males are larger) is often attributed to sexual selection favouring large males. When females are larger (female-biased SSD), it is often argued that natural selection favouring increased fecundity (i.e. larger clutches or eggs) has coevolved with larger female body size. Using comparative phylogenetic and multispecies regression model selection approaches, we test the hypothesis that among-species variation in female fecundity is associated with the evolution of female-biased SSD. We also ask whether the hypothesized relationship between SSD and fecundity is relaxed upon the evolution of parental care. Our results suggest a strong relationship between the evolution of fecundity and body size, but we find no significant relationship between fecundity and SSD. Similarly, there does not appear to be a relationship between fecundity and the presence or absence of parental care among species. Thus, although female body size and fecundity coevolve, selection for increased fecundity as an explanation for female-biased SSD is inconsistent with our analyses. We caution that a relationship between female body size and fecundity is insufficient evidence for fecundity selection driving the evolution of female-biased SSD.

Andrew meets Rensch: sexual size dimorphism and the inverse of Rensch's rule in Andrew's toad (Bufo andrewsi).

Variation in sexual size dimorphism (SSD) is a widespread phenomenon and is commonly attributed to variation in sex-specific patterns of selection. According to Rensch's rule, SSD increases with increasing body size when males are the larger sex, and decreases when females are the larger sex. Using data from 17 populations of Andrew's toad (Bufo andrewsi), we tested whether the patterns of SSD conform to Rensch's rule. Using field experiments, we also evaluated the hypothesis that sexual selection favours large male body size and that fecundity selection favours large female body size. The results revealed that the degree of SSD increased with increasing mean size in females, consistent with the inverse of Rensch's rule. Although experiments revealed evidence for a large-male mating advantage, selection for large male size was weak at best, and hence unlikely to be an important source of variation in SSD. However, fecundity selection favouring large females was evident, and likely to explain the observed inverse of Rensch's rule. After correcting male and female body size for age differences, the patterns of SSD remained the same, suggesting that the intra- and interpopulational variation in SSD is not driven by sex differences in age structure. Hence, these findings suggest that the strong fecundity selection favouring large females drives the evolution of female-biased SSD in B. andrewsi, providing an explanation for the inverse of Rensch's rule. As such, the study provides an important addition to the small body of literature that uses an intraspecific approach to demonstrate the inverse of Rensch's rule.

Covariation and repeatability of male mating effort and mating preferences in a promiscuous fish

Although mate choice by males does occur in nature, our understanding of its importance in driving evolutionary change remains limited compared with that for female mate choice. Recent theoretical models have shown that the evolution of male mate choice is more likely when individual variation in male mating effort and mating preferences exist and positively covary within populations. However, relatively little is known about the nature of such variation and its maintenance within natural populations. Here, using the Trinidadian guppy (Poecilia reticulata) as a model study system, we report that mating effort and mating preferences in males, based on female body length (a strong correlate of fecundity), positively covary and are significantly variable among subjects. Individual males are thus consistent, but not unanimous, in their mate choice. Both individual mating effort (including courtship effort) and mating preference were significantly repeatable. These novel findings support the assumptions and predictions of recent evolutionary models of male mate choice, and are consistent with the presence of additive genetic variation for male mate choice based on female size in our study population and thus with the opportunity for selection and further evolution of large female body size through male mate choice.

When the Seasons Don't Fit: Speedy Molt as a Routine Carry-Over Cost of Reproduction

The failure of animals to fit all life-cycle stages into an annual cycle could reduce the chances of successful breeding. In some cases, non-optimal strategies will be adopted in order to maintain the life-cycle within the scope of one year. We studied trade-offs made by a High Arctic migrant shorebird, the red knot Calidris canutus islandica, between reproduction and wing feather molt carried out in the non-breeding period in the Dutch Wadden Sea. We compared primary molt duration between birds undertaking the full migratory and breeding schedule with birds that forego breeding because they are young or are maintained in captivity. Molt duration was ca. 71 days in breeding adults, which was achieved by an accelerated feather replacement strategy. Second-year birds and captive adults took ca. 22% and 27% longer, respectively. Second-year birds start molt in late June, more than four weeks before captive adults, and almost seven weeks before adults that return from breeding in late July–August. Adults finish molt in October when steeply increasing thermostatic costs and reductions in food availability occur. Primary molt duration was longer in female than in male knots (all ages), which was accordance with the somewhat larger body size of females. Since fast growth leads to lower quality feathers, the speedy wing molt shown by Arctic-breeding birds may represent a time constraint that is an unavoidable and routine cost of reproduction. So far it was hypothesized that only birds over 1 kg would have difficulty fitting molt within a year. Here we show that in birds an order of magnitude smaller, temporal imperatives may impose the adoption of non-optimal life-cycle routines in the entire actively breeding population.

Species composition, reproduction, and body size of mud crabs, Scylla spp., caught in Urado Bay, Japan

Species composition, reproduction and body size of mud crabs, genus Scylla de Haan, 1833, were investigated using gill nets from October 2008 to October 2009 in Urado Bay, Japan. Three mud crab species were identified in the area, with S. paramamosain Estampador, 1949 being the dominant species (74% of the catch), followed by S. serrata (Forskal, 1775) (23%) and S. olivacea (Herbst, 1796) (3%). We found temporal changes in species composition with abundances of S. serrata increasing towards the summer season. Ovigerous females of S. paramamosain and S. olivacea occur between January and October, peaking in the warm (May-July) and rainy (June-July) seasons, but we found no berried females of S. serrata. The female-biased sex ratios of S. paramamosain and S. serrata, and the larger body size of females during the period from autumn (November) to early spring (April), suggest that large gravid females may have migrated offshore before the spawning season. The mean body size of females and males of S. paramamosain and S. serrata tended to increase and the mating activity was high between May and October, showing the moulting and growth season.