Inverse of Rensch’s Rule of Allometry for Sexual Size Dimorphism in Bufo andrewsi

Difference in body size between males and females(sexual size dimorphism:SSD)and its variation are a common phenomenon in animal kingdom.Rensch’s rule predicts that the degree of SSD variation increases with the enlarged mean body size when males are larger than females and decreases when females are larger than males.Here,whether the patterns of variations in SSD in the Andrew’s toad(Bufo andrewsi)follow Rensch’s rule was tested using unpublished data from 14 populations and published data from 17 populations.Results show the reduced major axis regression of log10(male size)on log10(female size)across all populations displayed a significant hyperallometric relationship,which was consistent with inverse Rensch’s rule(the degree of SSD increased with enlarged mean body size).SSD could also be explained by sexual age difference(SAD)due to a positive SSD–SAD relationship among all populations.The findings suggest that the occurrence of inverse Rensch’s rule in B.andrewsi is likely to be a result of fecundity selection on increased reproductive investments in larger females.

Significant Male Biased Sexual Size Dimorphism in Leptobrachium leishanensis

Sexual size dimorphism（SSD） is a widespread phenomenon among animals, and whose evolution and maintenance has been a central topic in evolutionary biology since Darwin＇s time. SSD varies in direction among the major taxonomic groups of animals and even within the same groups. In anurans, female biased SSD is the rule in many lineages, whereas male biased SSD is a rare phenomenon. In this paper, we analyze whether SSD exists inLeptobrachium leishanensis by comparing morphological characteristics between the sexes. Our results show that all six morphological characteristics measured are significantly different between the sexes. Males are significantly larger than females, indicating that the male biased SSD of this species is apparent. The size of the nuptial spines, a special secondary sex trait of males, is significantly and positively correlated with body size. We suggest that the resource defense polygyny mating system and parental care behavior may be explanations for the evolution of male biased SSD and nuptial spine development in this species.

Sex Ratio and Sexual Size Dimorphism in a Toad-headed Lizard, Phrynocephalus guinanensis

Phrynocephalus guinanensis has sexual dimorphism in abdominal coloration, but its ontogenetic development of sexual size dimorphism （SSD） is unknown. Using mark-recapture data during four days each year from August from 2014 to 2016, we investigated the development of sex ratios, SSD, sex-specific survivorship and growth rates in a population of P guinanensis. Our results indicated that the sex ratio of males to females was 1：2.8. Males had a lower survival rate （6%） than females （14%） across the age range from hatchling to adult, which supported the discovered female-biased sex ratio potentially associated with the low survival rate of males between hatchlings and juveniles. Male-biased SSD in tail length and head width existed in adults rather than in hatchling or juvenile lizards. The growth rates in body dimensions were undistinguishable between the sexes during the age from hatchling to juvenile, but the growth rate in head length from juvenile to adult was significantly larger in males than females. Average growth rate of all morphological measurements from hatchling to juvenile were larger compared with corresponding measurements from juvenile to adult, but only being significant in tail length, head width, abdomen length in females and snout-vent length in males. We provided a case study to strengthen our understanding of the important life history traits on how a viviparous lizard population can survive and develop their morphology in cold climates.

Sexual size dimorphism and female reproduction in the white-striped grass lizard Takydromus wolteri

Sexual size dimorphism （SSD） has long attracted the attention of biologists, and life-history variation is thought to play an important role in the evolution of SSD. Here we quantified SSD and female reproductive traits to identify potential asso- ciations between SSD and female reproduction in the white-striped grass lizard Takydromus wolteri. In a population from Chuzhou, China, the largest male and female were 53.0 mm and 57.5 mm in snout-vent length （SVL）, respectively. Females were larger in SVL and abdomen length, whereas males were larger in head size and tail length. Females produced up to five clutches of eggs during the breeding season, with large females producing more clutches and more eggs per clutch than small ones. As a result, large females had a higher annual fecundity and reproductive output. Egg size was positively correlated with matemal SVL in the first clutch, but not in subsequent clutches. These results suggest that T. wolteri is a species with female-biased SSD, and that fectmdity selection, in which large females have higher fecundity due to their higher capacity for laying eggs, is likely correlated with the evolution of SSD in this species

Sexual size dimorphism in lizards:Rensch’s rule,reproductive mode,clutch size,and line fitting method effects

Rensch’s rule relates to a pattern whereby sexual size dimorphism is more female-biased in small-sized species and more male-biased in large-sized ones.We collected literature and museum data on the body size of males and females belonging to 4032 lizard species,as well as data on their reproductive modes and clutch sizes.We used phylogenetic comparative analyses,and general linear mixed models,to test Rensch’s rule and examined how re-productive mode and clutch size affect sexual size dimorphism.Sexual size dimorphism was independent of clutch size in lizard species with variable clutch sizes and in oviparous lizards.Large litters were associated with female-biased sexual dimorphism in viviparous and in scincomorph lizards.Inference regarding Rensch’s rule depended on the analytical method used to identify it.The widely used,but less conservative,reduced major axis regression usually support Rensch’s rule while ordinary least squares regressions mostly show isometric relationships.The rule tended to apply more to oviparous than to viviparous lizards.We infer that Rensch’s rule is,at best,a weak pattern in lizards.This is especially true in viviparous lineages where females reproduce infrequently and therefore evolve large sizes to maximise fecundity,resulting in female-biased dimorphism.

Sexual Dimorphism of Head Size in Phrynocephalus przewalskii: Testing the Food Niche Divergence Hypothesis

Sexual size dimorphism （SSD） is a general phenomenon in lizards, and can evolve through sexual selection or natural selection. But natural selection, which was thought to operate mainly through reducing the competition be- tween the two sexes （niche divergence hypothesis）, gave rise to a lot of controversy. We tested the niche divergence hypothesis in the toad-headed lizard Phrynocephalus przewalskii by comparing diet composition and prey sizes between males and females. The species was found to be sexual dimorphic, with males having relatively larger snout-vent length, head width, head length, and tail length, while females have relatively larger abdomen length. Based on analysis of 93 studied stomachs, a total of 1359 prey items were identified. The most common prey items were formicid, lygaeid and tenebrionid. The two sexes did not differ in the relative proportions of prey size categories they consumed and the dietary overlap based on prey species was high （O = 0.989）. In addition, the meal size, the volume or any maximal dimension of the largest prey item in the stomach was not explained by the sexes. According to our results, food niche divergence might not play an important role in the SSD evolution ofP. przewalskii.

Ontogenetic mechanisms underlying sexual size dimorphism in Urodele amphibians： An across-species approach

Why do two sexes of the same species differ in body size holds a long-standing question of evolutionary biology. While many across-species comparisons have focused on ultimate causes behind sexual size dimorphism （SSD）, only have a few been directed toward elucidating its ontogenetic basis. Urodeles are an amphibian group in which the direction and degree of SSD vary greatly among species. Using demographic data yielded by skeletochronology for 33 urodele species, the current study re- veals a positive across-species correlation between SSD and the sex difference in mean age of adult animals, and the latter in- creases with the corresponding difference in age at maturity; annual growth rate does not differ between the sexes. We conclude that extended longevities in one sex, which is mediated by delayed maturation, would allow it to grow for longer and get larger, with growth rate making a weak contribution to body size. The sex-specific divergence in ontogenetic trajectory might be ex- plained by potentially high growth costs of reproduction to females in association with stronger fecundity selection, and to males that are expected to experience stronger sexual selection [Current Zoology 59 （1）： 142-150, 2013].

Variation and Sexual Dimorphism of Body Size in the Plateau Brown Frog along an Altitudinal Gradient

Variation in body size and sexual size dimorphism（SSD） can have important consequences for animal ecology, behavior, population dynamics and the evolution of life-history traits. Organisms are expected to be larger in colder climate（i.e., Bergmann＇s rule） and SSD varies with body size（i.e., Rensch＇s rule）. However, the underlying mechanisms are still elusive. The plateau brown frog（Rana kukunoris）, a medium-sized anuran species with femalebiased SSD, is endemic to the Qinghai-Tibetan Plateau（QTP）. From 1797 m（Maoxiang＇ping） to 3453 m（Heihe＇qiao） in the eastern margin of the QTP, we surveyed 10 populations of R. kukunoris and collected phalanges and snout vent length（SVL） data for 258 adult individuals（199 males versus 59 females）. Based on these data, we explored how body size and SSD varying along the altitudinal gradient and examined the corresponding effects of temperature. We found body size to be larger at higher altitude for males but not for females, with likely effects from the temperature on the variation in male body size. Sex differences in growth rates may be the main cause of the variation in SSD. Our results suggested that only males follow the Bergmann＇s rule and variation in SSD of R. kukunoris do not support the Rensch＇s rule and its inverse. Therefore, the variations of body size can be different between sexes and the applicability of both Bergmann＇s rule and Rensch＇s rule should depend on species and environment where they live.

Different paths to sexual size dimorphism in two praying mantids, Pseudomantis albofimbriata and Hierodula majuscula

Sexual size dimorphism （SSD） is widespread among diverse animal taxa and has attracted the attention of evolutionary biologists for over a century. SSD is likely to be adaptive and the result of divergent selection on different size optima for males and females, given their different roles in reproduction. The developmental trajectory leading to SSD may help us to understand how selection acts on male and female size. Here, we describe the growth and development of two Australian praying mantids, Pseudomantis albofimbriata and Hierodula majuscula including the number of moults, time to adulthood, size at each moult, and the degree of SSD. While both species exhibit the common pattern of female-biased SSD, the number of moults required for individuals to reach adulthood differed between males and females and between species. Despite their larger adult size, P. albofimbriata females require fewer moults and less time than males to reach adulthood, but are significantly larger than males from the second instar onwards. In contrast, H. majuscula males reached adulthood in fewer moults, and less time than females, however males and females did not differ in size until females went through their final moult into adulthood. H. majuscula also required more time and more moults to reach adulthood than 17. albofimbriata. We discuss these different developmental pathways in light of the existing knowledge of reproductive biology for each species. We also suggest that these differences may relate to the different phenologies that occur in strongly seasonal temperate environments compared with those in the tropics. This study provides evidence that SSD can result from two different patterns of growth and development in closely related species.

Subtle sexual plumage color dimorphism and size dimorphism in a South American colonial breeder, the Monk Parakeet(Myiopsitta monachus)

Background:Parrots(Psittacidae Family)are one of the most colorful groups of birds in the world,their colors pro-duced both structurally and via unusual pigments(psittacofulvins).Most species are considered to be monogamous,and many have been viewed historically as sexually monomorphic and monochromatic.However,studies using morphometric analysis and spectrophotometric techniques have revealed sexual size dimorphism and also sexual plumage color dimorphism among some species.The Monk Parakeet(Myiopsitta monachus),a native parrot of South America,is an interesting species for the study of plumage coloration and size since it is considered sexually mono-chromatic and monomorphic.Furthermore,recent studies show that the Monk Parakeet has extra-bond paternity behavior and even breeding trios,which suggests that sexual selection may play an important role in this species,and that it might have sexually dimorphic plumage(albeit imperceptible by humans)and be dimorphic in size.Methods:For the determination of plumage color we used spectrophotometry in the range of avian vision(300-700 nm)and performed a morphological analysis.Results:Our spectrophotometric results indicate that the Monk Parakeet shows subtle sexual plumage color dimor-phism in three(crown,nape and wing)out of twelve body regions.Similarly,our morphometric analysis showed that there are subtle sex differences in body size(bill and weight).Conclusions:Although the Monk Parakeet shows extra-bond paternity and breeding trio behaviors which could increase sexual dimorphism,these behaviors occur among highly related individuals;perhaps the high rate of inbreeding is responsible for the attenuation of sexual plumage color dimorphism and sex differences in body size observed.

Thermoregulatory function and sexual dimorphism of the throat sack in Helmeted Guineafowl(Numida meleagris)across Africa

The responses of ground-dwelling birds to heat and cold stress encompass a variety of behavioural,physiological and even morphological mechanisms.However,the role of glabrous skin in this respect has been marginally addressed so far.The Helmeted Guineafowl(Numida meleagris)is a landfowl distributed across Sub-Saharan Africa with eight traditionally recognised extant subspecies.Among the most prominent morphological traits underlying intraspecific variability are size and pigmentation of the bare throat skin(or sack),which might be related to the different habitats and environmental conditions across its wide range.In order to explore the Helmeted Guineafowl range-wide sack variation and pigmentation in relation to thermoregulation and sexual signalling,we collected morphometric and environmental information for N.m.coronata integrating field data with the inspection of photographic material encompassing seven subspecies and environmental information from their habitats.Field data evidenced that sack size was significantly correlated with ambient temperature,thus pointing to a likely involvement of the throat sack in thermoregulation.When the pictorial data from all subspecies were pooled,sack size correlated negatively with biomass,rainfall and humidity,while a positive correlation was found with annual solar irradiation.Sack size correlated positively with monthly temperature variation among the bluethroated subspecies from southern Africa as opposed to the black-throated subspecies ranging north to Zambia and Mozambique.Still,in this latter group the sack was often larger during winter months,possibly to maximise solar radiation absorbance.Noteworthy,sack size was related to sex dimorphism in two subspecies.Sack morphology and colour in the Helmeted Guineafowl likely modulate body temperature by evaporative cooling or heating upon needs,but in some subspecies it is also seemingly related to sexual signalling.Additional studies are needed to fully understand the multifunctionality of this important morphological feature in this species.

Seasonal change in sexual differences in nestling size and survival： a framework to evaluate sex-dependent environmental sensitivity in the wild

Background: Sexual size dimorphism(SSD) occurs in a wide range of species in birds and other animals, but the magnitude of SSD often varies with environmental conditions. In general, in the developmental stages, the larger sex is more vulnerable to adverse environmental conditions because the larger sex requires more energy than the smaller sex. However, this may not hold true for birds with large brood sizes; the larger sex can acquire more food by suppressing the smaller sex. In addition, most previous studies have been experimental, such as by manipulating clutch size and ectoparasites, which may not reflect natural conditions.Methods: In the present study, we propose a general framework to assess sexual differences in environmental sensitivity in natural populations. Because environmental conditions change throughout the breeding season, seasonal changes of nestling SSD and sex ratio should reflect sexual differences in environmental sensitivity. We applied this approach to a large dataset(1555 nestlings over 5 years) of Japanese Tits(Parus minor). In this population, the male nestling is generally larger than the female(5% SSD in body weight).Results: We found that the magnitude of SSD(weight, tarsus, wing) and fledgling sex ratio increased both in the beginning and the end of the breeding season.Conclusion: Our study suggested that female nestlings are more valuable to poor environmental conditions in the relatively fecund species. This study underscores the importance of brood size on sexual differences in environmental stochasticity and our framework encourages comparative analysis among different bird species.

Bare-throated spurfowl(Pternistis spp.)males across Africa impress females with bright throat colours during courtship

The role of bare body parts in sexual signalling in birds has received relatively little attention.I describe how the bare-throated spurfowl males saturate the colours of their throats to attract females.Of the 23 Afrotropical spurfowl species,the bare-throated subgroup includes Yellow-necked Spurfowl(Pternistis leucosceptus),Rednecked Spurfowl(P.afer),Grey-breasted Spurfowl(P.rufopictus) and Swainson’s Spurfowl(P.swainsonii).The rest of the species include fully feathered throated spurfowls.Throat colour intensity of bare throats was scored using an extensive online digital photographic archive encompassing the four species across the year’s seasons.Each throat(n=836) was assigned to one of four colour-intensity categories to explore the relationship between colour intensities,breeding cycles,and environmental variation.Except for Swainson’s Spurfowl male saturation of throat colours correlated with monthly rainfall,which peaks one or two months before egg laying.Swainson’s Spurfowl peaks during egg laying.Yellow-necked Spurfowl has the largest bare throat.Bare-throated spurfowl males perform an elevated courtship display posture above the female to feature their throat colour.No such displays occur in feather-throated spurfowl.Males with low throat colour saturation harbour more ectoparasites on their bare throats than birds with saturated throats.Male Red-necked Spurfowls have significantly larger bare throats than females.The primary function of bare throats probably assists in thermoregulation,particularly in arid regions.The bare throat may have evolved a secondary role in mating.Yellow-necked,Red-necked,and Greybreasted Spurfowls use their saturated throat colours as ornaments to court females during the breeding season.Unobtrusive female throat colours(unsaturated) may discourage male interlopers and predation during egg laying.Saturation appears to be carotenoid-food based.The different colours among the bare-throated species may serve as prezygotic mechanisms that inhibit cross-breeding and explain why females also have coloured throats.

Three new species of Parandrexis(Coleoptera:Parandrexidae) from the Middle Jurassic Jiulongshan Formation of Inner Mongolia,China

Three well-preserved fossil species ofParandrexis longicornis Lu, Shih ＆ Ren sp. nov., P. agilis Lu, Shih ＆ Ren sp. nov. and P. oblongis Lu, Shih ＆ Ren sp. nov. are described and illustrated from the late Middle Jurassic Jiulongshan Formation of Daohugou in Inner Mongolia Autonomous Region, China. These new species are distinguished from other described species of Parandrexis by body size, head dimension （ratio of length to width）, pronotum shape, and antennal characters. Based on new morphological data, we emend the generic diagnosis. These findings support the sexual size dimorphism of parandrexids and suggest a gymnosperm sap feeding habit by Parandrexis.

A Review for Life-history Traits Variation in Frogs Especially for Anurans in China

Environmental variation can promote differentiation in life-history traits in species of anurans. Increased environmental stress usually results in larger age at sexual maturity, older mean age, longer longevity, slower growth, larger body size, and a shift in reproductive allocation from offspring quantity to quality, and a stronger trade-off between offspring size and number. However, previous studies have suggested that there are inconsistent geographical variations in life-history traits among anuran species in China. Hence, we here review the intraspecific patterns and differences in life-history traits（i.e., egg size, clutch size, testes size, sperm length, age at sexual maturity, longevity, body size and sexual size dimorphism） among different populations within species along geographical gradients for anurans in China in recent years. We also provide future directions for studying difference in sperm performance between longer and shorter sperm within a species through transplant experiments and the relationships between metabolic rate and brain size and life-history.

Skeletochronological Study on Age Structure of a Chinese Endemic Frog(Rana omeimontis)

We used skeletochronology to estimate age structure of the Omei Wood Frog, Rana omeimontis, from southwestern China.Average age differed significantly between males and females.Age at sexual maturity in both males and females was 1 year. Longevity was 6 and 7 years for males and females, respectively. Average body size differed significantly between the sexes, with females being larger than males. A non-significant correlation between age and body size was found within each sex in this population. The ANCOVA analysis revealed that females also had larger body size than males when the effect of age was controlled.

Essential roles of stat5.1/stat5b in controlling fish somatic growth

Signal transducer and activator of transcription 5b（STAT5b） has been identified as a key downstream mediator of growth hormone（GH） signaling in somatic growth of mammalian. However, the corresponding homologue gene of Stat5b is unknown in fish species. In this study, we generated loss-offunction mutants in stat5.1 and stat5.2, two stat5 homologues existing in zebrafish. In stat5.1-deficient zebrafish, a significant reduction of body length and body weight was detected in the embryos/larvae and adults compared with the wild-type control fish, and sexual size dimorphism in adult zebrafish was also eliminated. However, the stat5.2-deficient zebrafish displayed a normal developmental phenotype during all lifespan. Chromatin immunoprecipitation combined with deep sequencing（Ch IP-seq） method was adopted to further investigate the potential transcriptional targets of Stat5 protein and cast much light upon the biological function of Stat5. We identified more than 800 genes as transcriptional targets of Stat5 during zebrafish embryogenesis. KEGG analysis indicated that the Stat5 target gene network is predominantly linked to the metabolic pathways, neuroactive ligand-receptor interaction and JAK-STAT signaling pathways. Further validation studies suggested that Stat5.1 protein could directly regulate the expression of gh1, and stat5.1-mutated zebrafish showed a reduction of gh1 mRNA level. In the present study, stat5.1 was revealed as the corresponding homologue gene of Stat5b in fish species. Additionally,we found a novel molecular interaction between Stat5.1/Stat5b and GH, and unraveled a positive feedback loop Stat5.1-GH-Stat5.1 which is necessary for somatic growth and body development in zebrafish.

Lack of support for Rensch＇s rule in an intraspecific test using red flour beetle （Tribo/ium castaneum） populations

Rensch＇s rule proposes a universal allometric scaling phenomenon across species where sexual size dimorphism （SSD） has evolved： in taxa with male-biased dimorphism, degree of SSD should increase with overall body size, and in taxa with female-biased dimorphism, degree of SSD should decrease with increasing average body size. Rensch＇s rule appears to hold widely across taxa where SSD is male-biased, but not consistently when SSD is female-biased. Furthermore, studies addressing this question within species are rare, so it remains unclear whether this rule applies at the intraspecific level. We assess body size and SSD within Tribolium castaneum （Herbst）, a species where females are larger than males, using 21 populations derived from separate locations across the world, and maintained in isolated laboratory culture for at least 20 years. Body size, and hence SSD patterns, are highly susceptible to variations in temperature, diet quality and other environmental factors. Crucially, here we nullify interference of such confounds as all populations were maintained under identical conditions （similar densities, standard diet and exposed to identical temperature, relative humidity and photoperiod）. We measured thirty beetles of each sex for all populations, and found body size variation across populations, and （as expected） female-biased SSD in all populations. We test whether Rensch＇s rule holds for our populations, but find isometry, i.e. no allometry for SSD. Our results thus show that Rensch＇s rule does not hold across populations within this species. Our intraspecific test matches previous interspecific studies showing that Rensch＇s rule fails in species with female-biased SSD.