Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

山地麻蜥个体发育过程中头部两性异形和食性的变化

研究了山地麻蜥(Eremias brenchleyi)个体发育过程中头部两性异形和食性的变化．成体个体大小(SVL)无显著的两性差异,但雄体具有较大的头部(头长和头宽)．头部两性异形在孵出幼体就已存在,成体头部两性异形比幼体(包括孵出幼体)更为显著,雄性较大的头部与其头部随SVL的增长速率大于雌性有关．两性头部总体上随SVL呈异速增长,表现为个体发育过程中头长和头宽与SVL的线性回归方程斜率有显著的变化．孵出幼体有相对较大的头部,这种形态特征是胚胎优先保证生态学意义更为显著的头部生长的结果,有利于孵出幼体的早期生存和生长．相对头部大小在个体发育过程中有显著的变化．不同性别和大小的山地麻蜥摄入食物的种类及各种食物在摄入食物中所占的比例有一定程度的差别,食物生态位宽度和重叠度因此有一定的差别．然而,没有直接的证据表明头部两性异形能导致两性食物生态位的明显分离,并有利于减缓两性个体对食物资源的竞争。     

Sexual dimorphism of body and relative head sizes in neonatal common garter snakes

Morphological and behavioral differences between sexes are commonplace throughout the animal kingdom. Body size is one of the most obvious sex differences frequently found in snakes. However, the developmental origins of size differences in many species, including snakes, are not well known. We examined post-natal variation in sexual size dimorphism in garter snakes Thamnophis sirtalis . The weights, body and tail lengths, and head sizes of male and female neonates born to mothers collected from ecologically dissimilar habitats on Beaver Island, Lake Michigan were compared. Sexual size dimorphism was prominent. Overall, males had significantly longer bodies and tails than females. Females were significantly heavier and had larger heads than male snakes. Maternal site affected head but not body measurements, perhaps due to differences in prey availability. The body condition of maternal females predicted neonatal body length. Significant litter variation suggests heritable variation in morphological traits possibly correlated with feeding success and survival.     

Sexual divergence in diets and morphology in Fijian sea snakes Laticauda colubrina (Laticaudinae)

In the Fiji Islands, female yellow‐lipped sea kraits (Laticauda colubrina) grow much larger than males, and have longer and wider heads than do conspecific males of the same body length. This morphological divergence is accompanied by (and may be adaptive to) a marked sex divergence in dietary habits. Adult female sea kraits feed primarily on large conger eels, and take only a single prey item per foraging bout. In contrast, adult males feed upon smaller moray eels, and frequently take multiple prey items. Prey size increases with snake body size in both males and females, but the sexes follow different trajectories in this respect. Female sea kraits consume larger eels relative to predator head size and body length than do males. Thus, the larger relative head size of female sea kraits is interpreted as an adaptation to consuming larger prey items. Our results are similar to those of previous studies on American water snakes (natricines) and Australian file snakes (acrochordids), indicating that similar patterns of sex divergence in dietary habits and feeding structures have evolved convergently in at least three separate lineages of aquatic snakes.     

Sexual dimorphism, reproductive biology, and food habits of two species of African filesnakes (Mehelya, Colubridae)

The ecology and general biology of African snakes remains virtually unstudied, even in highly distinctive species such as the filesnakes (genera Mehelya and Gonionotophis ). Our measurements and dissections of preserved specimens provided information on body sizes, sexual dimorphism in size and bodily proportions, clutch sizes, and food habits of two Mehelya species. In both M. capensis and M. nyassae , females attain sexual maturity at the same size as conspecific males, but grow to much larger sizes. Mehelya capensis displays extreme differences in body shape between males and females at the same body length: females have longer and wider heads, thicker bodies, and larger eyes (relative to both head length and head width) than do conspecific males. Dimorphism in body proportions is less marked in M. nyassae. Female reproductive cycles are seasonal in M. capensis , and clutch sizes are larger in this species than in its smaller congener (5-11 eggs in M. capensis , 2-6 eggs in M. nyassae ).
Contrary to popular wisdom, Mehelya are not specialized ophiophages. Mehelya nyassae feeds primarily upon lygosomatine skinks, including many fossorial taxa. Mehelya capensis has a broader diet, feeding on a wide variety of terrestrial lizards (especially agamids and gerrhosaurids) and snakes. Toads are also common prey items. The diversity of prey types taken by M. capensis suggests that these snakes may use ambush predation as well as active foraging. Mehelya is strongly convergent with Asian elapids of the genus Bungarus in its morphology (triangular body shape; powerful jaws; visible interstitial skin), behaviour (nocturnality; reluctance to bite when harassed), and diet (feeding on elongate reptiles, including snakes). Observations of preyhandling and ingestion by captive snakes are needed to clarify possible selective forces for the evolution of the unusual traits shared by these taxa.     

Relationships between sexual dimorphism and niche partitioning within a clade of sea-snakes (Laticaudinae)

Previous studies in Fiji have shown that females of the amphibious sea-krait Laticauda colubrina are much larger than males, and have larger heads relative to body size. The dimorphism has been interpreted in terms of adaptation to a sex divergence in prey-size: females primarily eat large (conger) eels rather than smaller (moray) eels. The hypothesis that dimorphism is affected by niche divergence predicts that the degree of sex dimorphism will shift when such a species invades a habitat with a different range of potential prey sizes. On the island of Efate in Vanuatu, L. colubrina and a regionally endemic sibling species (L. frontalis) both consume smaller eels (in absolute terms, and relative to the snake's body size) than do the previously-studied Fijian snakes. Patterns of morphology and sexual dimorphism have shifted also. Both Vanuatu taxa are slender-bodied, and frontalis is smaller and less dimorphic than L. colubrina. Females grow larger than males in all taxa, and have larger heads (relative to body length), but the degree of sexual divergence is lower in Vanuatu (especially in frontalis). Dietary overlap (in prey species as well as size) is high between adult frontalis and juvenile colubrina, but the two taxa differ in prey size/predator size relationships. In particular, male frontalis eat very small prey and have very short heads. Our results are consistent with the hypothesis that sex differences in the mean adult body sizes and relative head sizes of laticaudine snakes are linked to sex differences in feeding biology.     

Effects of season, sex and body size on the feeding ecology of turtle-headed sea snakes (Emydocephalus annulatus) on IndoPacific inshore coral reefs

In terrestrial snakes, many cases of intraspecific shifts in dietary habits as a function of predator sex and body size are driven by gape limitation and hence are most common in species that feed on relatively large prey and exhibit a wide body-size range. Our data on sea snakes reveal an alternative mechanism for intraspecific niche partitioning, based on sex-specific seasonal anorexia induced by reproductive activities. Turtle-headed sea snakes (Emydocephalus annulatus) on coral reefs in the New Caledonian Lagoon feed entirely on the eggs of demersal-spawning fishes. DNA sequence data (cytochrome b gene) on eggs that we palpated from stomachs of 37 snakes showed that despite this ontogenetic stage specialization, the prey comes from a taxonomically diverse array of species including damselfish (41 % of samples, at least 5 species), blennies (41 %, 4 species) and gobies (19 %, 5 species). The composition of snake diets shifted seasonally (with damselfish dominating in winter but not summer), presumably reflecting seasonality of fish reproduction. That seasonal shift affects male and female snakes differently, because reproduction is incompatible with foraging. Adult female sea snakes ceased feeding when they became heavily distended with developing embryos in late summer, and males ceased feeding while they were mate searching in winter. The sex divergence in foraging habits may be amplified by sexual size dimorphism; females grow larger than males, and larger snakes (of both sexes) feed more on damselfish (which often lay their eggs in exposed sites) than on blennies and gobies (whose eggs are hidden within narrow crevices). Specific features of reproductive biology of coral reef fish (seasonality and nest type) have generated intraspecific niche partitioning in these sea snakes, by mechanisms different from those that apply to terrestrial snakes.     

Rejection of non-adaptive hypotheses for intraspecific variation in trophic morphology in gape-limited predators

Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evidence for adaptation of the trophic apparatus in gape-limited predators to local prey size. However, such variation might also arise from non-adaptive processes (such as allometry, correlated response, genetic drift, or non-adaptive phenotypic plasticity). We test predictions from these alternative hypotheses using data on the allometric relationship between head size and body size in two wide-ranging snake species: eight populations of adders ( Vipera berus ) and 30 populations of common gartersnakes ( Thamnophis sirtalis ). Our data enable strong rejection of the alternative (non-adaptive) hypotheses, because the relationship between head and body size differed significantly among populations, the geographic distance separating pairs of populations explained less than 1.5% of their divergence in allometric coefficients, and the within-population allometric coefficients were higher than the among-population coefficients in each species. In addition, the geographical variability of allometric coefficients in females did not parallel that in males, suggesting that allometric coefficients have evolved independently in the two sexes. Phenotypic plasticity also cannot explain the data, because laboratory studies show that the allometric relationship between head size and body size is relatively insensitive to differing growth rates. We conclude that the intraspecific head size divergence in these snakes is better explained by spatially heterogeneous selection to optimize prey handling ability, than by non-adaptive processes.     

Sexual differences in morphology and niche utilization in an aquatic snake,Acrochordus arafurae

Filesnakes (Acrochordus arafurae) are large (to 2 m), heavy-bodied snakes of tropical Australia. Sexual dimorphism is evident in adult body sizes, weight/length ratios, and body proportions (relative head and tail lengths). Dimorphism is present even in neonates. Two hypotheses for the evolution of such dimorphism are (1) sexual selection or (2) adaptation of the sexes to different ecological niches. The hypothesis of sexual selection is consistent with general trends of sexually dimorphic body sizes in snakes, and accurately predicts, for A. arafurae, that the larger sex (female) is the one in which reproductive success increases most strongly with increasing body size. However, the sexual dimorphism in relative head sizes is not explicable by sexual selection.The hypothesis of adaptation to sex-specific niches predicts differences in habitats and/or prey. I observed major differences between male and female A. arafurae in prey types, prey sizes and habitat utilization (shallow versus deep water). Hence, the sexual dimorphism in relative head sizes is attributed to ecological causes rather than sexual selection. Nonetheless, competition between the sexes need not be invoked as the selective advantage of this character divergence. It is more parsimonious to interpret these differences as independent adaptations of each sex to increase foraging success, given pre-existing sexually-selected differences in size, habitat or behavior. Data for three other aquatic snake species, from phylogenetically distant taxa, suggest that sexual dimorphism in food habits, foraging sites and feeding morphology, is widespread in snakes.     

Evolution of Sexual Size Dimorphism in a Frog Obeys the Inverse of Rensch’s Rule

Rensch’s rule refers to a pattern in sexual size dimorphism (SSD) in which SSD increases with body size when males are the larger sex and decreases with body size when females are the larger sex. Using data on body size from 40 populations and age from 31 populations of the rice frog Rana limnochari with female-biased size dimorphism, I tested the consistency of allometric relationships between males and females with Rensch’s rule and evaluated the hypothesis that SSD was largely a function of age differences between the sexes. Statistical comparisons of body sizes between the sexes showed the evidence for the inverse of Rensch’s rule, indicating the level of SSD increased with increasing mean body size. One of the explanations for the occurrence of the inverse of Rensch’s rule may be the fecundity selection hypothesis assuming increased reproductive output in large females. However, differences in age between males and females among populations could explain mildly the variation in SSD.     

Reproductive output,costs of reproduction,and ecology of the smooth snake,Coronella austriaca,in the eastern Italian Alps

A 5-year mark-recapture study of smooth snakes (Coronella austriaca) in the Carnic Alps (1100 m above sea level) of north-eastern Italy provided extensive information on the biology and life-history of these small viviparous snakes. Offspring were relatively large (mean=15 cm total length, 2.9 g) when they were born in late summer, and females grew to maturity (44 cm, 50 g) in approximately 4 years. Larger neonates retained their size advantage for at least 12 months, but did not have a higher probability of survival. Although sexual size dimorphism (at birth and at mean adult body sizes) was minor, the sexes differed significantly in several respects. Females grew faster than males during juvenile life, and adult females diverged in dietary habits from the rest of the population. Whereas juveniles (of both sexes) and adult males fed primarily on lizards, larger females shifted to feeding less frequently, but taking larger prey (mammals and snakes). Reproductive output increased strongly with maternal body size: larger females reproduced more frequently, produced larger litters of larger neonates, had higher relative clutch masses (RCMs), and had a lower proportion of stillborn off-spring. Most females produced a litter every 2nd or 3rd year. We did not detect significant year-to-year variation in reproductive traits over the 5 years of our study. Females were consistent from one litter to the next in several traits (e.g., litter sizes, offspring sizes and shapes, proportions of stillborn neonates, RCMs), but this consistency was due to differences in body size among females rather than to size-independent maternal effects. Overall litter sex ratios averaged 50/50, but sex ratios tended to be more male-biased in litters that were unusually large relative to maternal body size, and in litters containing a high proportion of stillborn offspring. Costs of reproduction appear to be high in this population, in terms of both energy allocation and risk. Reproduction reduced growth rates, and females that recovered condition more quickly in the year after reproduction were able to reproduce again after a briefer delay. Mortality was highest in reproducing females with high RCMs, and in females that were very emaciated after parturition. The marked increase in reproductive output with increasing maternal body size in C. austriaca may reflect a reduction in costs as females grow larger, and the dietary shift to larger prey may enhance the rate that females can accumulate energy for reproduction.     

蓝尾石龙子的头部两性异形和食性

通过测量头、体大小和胃检研究浙江泰顺产蓝尾石龙子 (Eumeceselegans)个体发育过程中两性异形和食性的变化。蓝尾石龙子成体个体大小和头部大小的两性差异显著 ,雄性大于雌性。不同发育阶段雌性头长与SVL的线性回归斜率无显著差异 ,头宽与SVL线性回归斜率的差异显著 ,成体和SVL <5 0mm幼体头宽随SVL的增长速率显著小于SVL为 5 0 - 6 9mm的幼体。雄性头部相对于SVL呈加速式异速生长。两性比较发现 :雌雄幼体头长和头宽随SVL的增长速率无显著差异 ,SVL <5 0mm幼体特定SVL的头长和头宽无显著的两性差异 ,但SVL为 5 0 - 6 9mm的雄性幼体头长和头宽大于SVL相同的雌性幼体 ;雄性成体头长和头宽随SVL的增长速率显著大于雌性。SVL <5 0mm的雌性幼体头部相对小于SVL为 5 0 - 6 9mm的同性幼体 ,性成熟雌体头部相对小于SVL为 5 0 - 6 9mm的同性幼体。雌性幼体、雄性幼体、雌性成体和雄性成体食物生态位宽度分别为 12 3、 12 5、 4 8和 14 4。雌雄幼体食物生态位重叠度最高 ,雌雄成体食物生态位重叠度次之 ,成体与幼体食物生态位重叠度较小。成体摄入食饵的大小 (用胃内完整食物长度的平均值表示 )和变化范围大于幼体。两性成、幼体摄入的食饵大小差异显著。两性个体摄入的食饵大小均与其SVL呈正相关 ,表明较大     

The evolution of sexual size dimorphism in the house finch. IV. Population divergence in ontogeny

Differences among taxa in sexual size dimorphism of adults can be produced by changes in distinct developmental processes and thus may reflect different evolutionary histories. Here we examine whether divergence in sexual dimorphism of adults between recently established Montana and Alabama populations of the house finch (Carpodacus mexicanus) can be attributed to population differences in growth of males and females. In both populations, males and females were similar at hatching, but as a result of sex-specific growth attained sexual size dimorphism by the time of independence. Timing and extent of growth varied between the sexes: Females maintained maximum rates of growth for a longer time than males, whereas males had higher initial growth rates and achieved maximum growth earlier and at smaller sizes than females. Ontogeny of sexual dimorphism differed between populations, but in each population, sexual dimorphism in growth parameters and sexual dimorphism at the time of nest leaving were similar to sexual dimorphism of adults. Variation in growth of females contributed more to population divergence than did growth of males. In each population, we found close correspondence between patterns of sexual dimorphism in growth and population divergence in morphology of adults: Traits that were the most sexually dimorphic in growth in each population contributed the most to population divergence in both sexes. We suggest that sex-specific expression of phenotypic and genetic variation throughout the ontogeny of house finches can result in different responses to selection between males and females of the same age, and thus produce fast population divergence in the sexual size dimorphism.     

灰椋鸟身体大小和内脏器官形态的两性差异

鸟类性二态现象广泛存在,比如身体大小、羽色等,性二态很可能是自然选择和性选择共同作用的结果.为了探索和更好地了解雀形目鸟类身体大小性二态的进化,在2019年繁殖季节早期研究了灰椋鸟(Sturnus cineraceus)野外种群身体大小和内脏器官形态的两性差异.结果表明,除嘴宽外,其他身体特征参数均雄性显著大于雌性,表...     

Seasonal dimorphism in the horny bills of sparrows

Bill size is often viewed as a species‐specific adaptation for feeding, but it sometimes varies between sexes, suggesting that sexual selection or intersexual competition may also be important. Hypotheses to explain sexual dimorphism in avian bill size include divergence in feeding niche or thermoregulatory demands, intrasexual selection based on increased competition among males, or female preference. Birds also show seasonal changes in bill size due to shifts in the balance between growth rate and wear, which may be due to diet or endogenous rhythms in growth. Insight into the function of dimorphism can be gained using the novel approach of digital x‐ray imaging of museum skins to examine the degree to which the skeletal core or the rhamphotheca contribute to overall dimorphism. The rhamphotheca is ever‐growing and ever‐wearing, varying in size throughout life; whereas the skeletal core shows determinant growth. Because tidal marsh sparrows are more dimorphic in bill size than related taxa, we selected two marsh taxa to investigate dimorphism and seasonality in the size of the overall bill, the skeletal core, and the rhamphotheca. Bill size varied by sex and season, with males having larger bills than females, and bill size increasing from nonbreeding to breeding season more in males. Skeletal bill size varied with season, but not sex. The rhamphotheca varied primarily with sex; males had a larger rhamphotheca (corrected for skeletal bill size), which showed a greater seasonal increase than females. The rhamphotheca, rather than the skeletal bill, was responsible for sexual dimorphism in overall bill size, which was particularly well developed in the breeding season. The size of the rhamphotheca may be a condition‐based character that is shaped by sexual selection. These results are consistent with the evidence that bill size is influenced by sexual selection as well as trophic ecology.     

Variation in the diet of the viperine snake Natvix mama in relation to prey availability

The diet of the viperine snake was compared with food availability in the Ebro Delta, a wetland largely occupied by rice fields, in 1990 and 1991. Snake selection of prey type and size was studied seasonally and by snake group: males, females and immature snakes. Overall, feeding activity (percentage of individuals with prey and number of prey per stomach) increased with food availability. Diet analysis showed that viperine snakes mainly foraged on the green frog Rana perezi (adults and tadpoles) and the carp Cyprinus earpio. Conversely, viperine snakes rejected the mosquito fish Gambusia holbroki which is the most abundant species in autumn, when Natrix maura has a low feeding activity. Statistical comparisons between viperine snake diet and prey availability showed that males selected small carp, immature snakes selected tadpoles and, in spring, females selected frogs. The selection of small carp by males may reflect a sexual divergence of trophic niche related to sexual size dimorphism, as females are larger than males. As tadpoles are presumably easier to catch than fish, tadpole selection by immature individuals may reflect variance in capture abilities. In spring, the selection of frogs by females overlapped with vitellogenesis, suggesting that females compensate for the cost of reproduction by selecting green frogs, which have a greater biomass and higher energy content than fish. Carps eaten in spring were smaller than in summer. Moreover, in summer viperine snakes selected smaller carp than the available mean size. This divergent tendency between carp size selection and carp size availability reveals how seasonal diet shifts in prey size selection may be a response to an increase in prey size.     

Reproductive versus ecological advantages to larger body size in female snakes, Vipera aspis

Body size can influence an organism's microevolutionary fitness either via ecological factors (ecological selection) or changes in reproductive output (sexual or fecundity selection). Published studies on sexual dimorphism in reptiles have generally focussed on sexual-selective forces on males, under the implicit assumption that the intensity of fecundity selection in females (and hence, overall selection on female body size) is likely to be relatively consistent among lineages. In this paper, we explore the degree to which larger body size enhances ecological attributes (e.g., food intake, growth, survival) and reproductive output (reproductive frequency, litter size, offspring size, offspring viability) in free-ranging female aspic vipers, Vipera aspis . The less-than-annual reproductive frequency of these animals allows us to make a direct comparison between females in years during which they concentrate on "ecological" challenges (especially building energy reserves) versus reproductive challenges (producing a litter). Because female snakes have limited abdominal space to hold the clutch (litter), we expect that fecundity should depend on body size. However, our data show that larger body size had a more consistent effect on ecological attributes (such as feeding rates and "costs of reproduction") than on reproductive output per se. Indeed, total reproductive output was maximised at intermediate body sizes. These results suggest that variation in female body size among and within species (and hence, in the degree of sexual dimorphism) may be driven by the ecological as well as reproductive consequences of body size variation in both sexes.     

Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size

Abstract.  1. Sexual differences in body size are expected to evolve when selection on female and male sizes favours different optima.
2. Insects have typically female-biased size dimorphism that is usually explained by the strong fecundity advantage of larger size in females. However, numerous exceptions to this general pattern have led to the search for selective pressures favouring larger size in males.
3. In this study, the benefits of large size were investigated in males of four species of ichneumonine wasps, a species-rich group of parasitoids, many representatives of which exhibit male-biased size dimorphism.
4. Mating behaviour of all ichneumonine wasps are characterised by pre-copulatory struggles, in the course of which males attempt to override female reluctance to mate. A series of laboratory trials was conducted to study the determinants of male mating success.
5. A tendency was found for larger males as well as those in better condition to be more successful in achieving copulations. Size dimorphism of the species studied, mostly male-biased in hind tibia length but female-biased in body weight, indicates that sexual selection in males favours longer bodies and appendages rather than larger weight.
6. The qualitative similarity of the mating patterns suggests that sexual selection cannot completely explain the considerable among-species differences in sexual size dimorphism.
7. The present study cautions against using various size indices as equivalents for calculating sexual size dimorphism.
8. It is suggested that female reluctance in ichneumonine wasps functions as a mechanism of female mate assessment.     

Sexual dimorphism and allometry in two seed beetles (Coleoptera: Bruchidae)

Male Callosobruchus chinensis (Coleoptera: Bruchidae) have elaborated, pectinate antennae, which are absent from conspecific females and both sexes of a congener, Callosobruchus maculatus. To begin to unravel the mechanisms producing this striking dimorphism, we examined which morphological traits best explain body size variation in bruchid beetles and quantified sexual dimorphism of antenna size through allometric analyses. Using principal component analyses, we found that elytron length and pronotum width were significantly correlated with the first principal component, which was interpreted as explaining variation in body size. Regressions of log‐transformed body size measures on log‐transformed antenna length revealed that males of both species had longer antennae than conspecific females for any given body size, although most of this effect was attributable to higher intercepts, rather than increased allometry, in males. Comparisons among heterospecific males revealed that C. maculatus males have noticeably longer antennae than C. chinensis males at large body sizes. Callosobruchus chinensis males, thus, appear to have increased the receptive area of their antennae by adding to the width of, rather than further elongating, their antennae. Finally, we found evidence for positive allometry between log‐transformed antenna length and log‐transformed antenna width in C. chinensis males. We discuss our results in the context of evidence supporting the presence of an additional, and potentially unique, sex pheromone in C. chinensis females.     

Dietary habits of the fanray Platyrhina tangi (Batoidea: Platyrhinidae) in Ariake Bay, Japan

The dietary habits of the fanray Platyrhina tangi were investigated by analyzing the stomach contents of specimens collected in Ariake Bay, Japan. Of 334 stomach specimens, 324 contained food and 10 (3.0%) were empty. The mean percentage weight of stomach contents per unit of body weight was 0.59%. Thirty-seven taxonomic levels of prey were identified. The most common prey was shrimp, followed by fish and mysids. There were no differences in the composition of the diet between sexes, but an ontogenetic dietary shift was observed. Shrimps were the most common prey in all size classes. In addition, smaller individuals frequently ate mysids, and larger individuals often consumed fish. Dietary breadth values increased with size. Trophic level analysis revealed that trophic level increased with size; however, this species is consistently a secondary consumer. Dental sexual dimorphism was also observed. Specifically, mature males had much longer and sharper cusps than females and immature males. Since males and females had similar diets, dental sexual dimorphism may be related to their reproductive behaviour.