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1.
Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life‐history traits are known to correlate with latitude, but sex‐specific responses are not well understood. We analyzed life‐history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex‐dependent latitudinal variation in life‐history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female‐biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large‐scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female‐biased species. Our study highlights the importance of sex‐specific differences in life‐history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.  相似文献   

2.
The present study investigated altitudinal variation in sexual size dimorphism of a Tibetan frog Nanorana parkeri. Size dimorphism was female‐biased in all populations, although this bias became less at higher altitudes because of a steeper altitudinal decrease in female size than male size. Operational sex ratios, an indicator of the opportunity for sexual selection on larger males, changed independently of altitude. Clutch volume, an indicator of the strength of fecundity selection on larger females, was positively with female size, and tended to decrease approaching high altitudes. Females lived longer and grew more slowly than males, and the mean age in both sexes increased and growth rate decreased altitudinally, although the changes were more rapid in females than males. These results suggest that, relative to males, females (i.e. the sex that typically bears greater reproductive costs and experiences stronger directional selection for larger size to take fecundity advantages) should be more sensitive to environments, attaining a larger size via enhancing growth under favourable lower‐latitude conditions but a smaller size as a result of retarding growth when conditions become harsher at higher altitudes. This supports the condition‐dependence hypothesis with respect to intraspecific variation in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 558–565.  相似文献   

3.
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.  相似文献   

4.
Carotenoids are a large group of biochemicals, with similar properties, synthesised by bacteria, fungi, algae and plants. Vertebrates obtain these biologically active pigments through the diet, and they are a disproportionately common component of animal colour signals and play important roles in immune functions and as antioxidants. Carotenoids are believed to be a limited resource and because of the trade‐off between allocation of carotenoids to signals and to other functions, carotenoid based signals are often thought to be handicap signals. The purpose of this study was to investigate the signalling potential of carotenoid‐based tissue coloration in the great black‐backed gull Larus marinus. The intensity of carotenoid‐based coloration in bill, gape and eye‐ring coloration was investigated in relation to body condition, reproductive parameters, levels of immune activity, and sexual dimorphism. In males there was a positive relationship between colour intensity and body condition, but in females no such relationship was found. However, females with high colour intensity had larger eggs and clutches. Additionally, females with high red scores tended to have high density of circulating lymphocytes. There was no sexual dimorphism in coloration and there was a negative relationship between colour intensity and sampling time, which indicates that this coloration is most intensely expressed early in the breeding season. The results in this study suggest that carotenoid‐based coloration in great black‐backed gull are partly condition dependent and reveal information about individual quality in both males an females. Hence, it might have evolved as an important signal for assessing the quality of potential mates.  相似文献   

5.
Female‐biased sexual dimorphism in size at maturity is a common pattern observed in freshwater fishes with indeterminate growth, yet can vary in magnitude among populations for reasons that are not well understood. According to sex‐specific optimization models, female‐biased sexual size dimorphism can evolve due to sexual selection favouring earlier maturation by males, even when sexes are otherwise similar in their growth and mortality regimes. The magnitude of sexual size dimorphism is expected to depend on mortality rate. When mortality rates are low, both males and females are expected to mature at older ages and larger sizes, with size determined by the von Bertalanffy growth equation. The difference between size at maturity in males and females becomes reduced when maturing at older ages, closer to asymptotic size. This phenomenon is called von Bertalanffy buffering. The predicted relationship between the magnitude of female‐biased sexual dimorphism in age and size at maturity and mortality rate was tested in a comparative analysis of lake whitefish Coregonus clupeaformis from 26 populations across a broad latitudinal range in North America. Most C. clupeaformis populations displayed female‐biased sexual dimorphism in size and age at 50% maturity. As predicted, female‐biased sexual size dimorphism was less extreme among lower mortality, high‐latitude populations.  相似文献   

6.
Sex steroid inhibitors were used to characterize the effects of 17beta-estradiol (E2) and testosterone (T) on the sexual growth dimorphism of Eurasian perch juveniles. In experiment 1, growth responses to different doses of either E2 (25, 50, 75, and 100 mgkg(diet)-1) or fadrozole (Fa; 50 and 100 mgkg(diet)-1) were compared in triplicate tanks of 30 fish each during 85 days. In experiment 2, five diets containing (50 mgkg(diet)-1) Tamoxifen (Ta), Flutamide (Flu), Fa, E2, and T were tested in triplicate tanks of 20 fish each during 90 days. Steroid supplementation or inhibition increased or decreased E2 and T plasma levels. Moreover, E2 treatment induced a higher plasma vitellogenin level but decreased triidothyronine levels. Brain aromatase activity (AA) was lower in Fa-treated fish than in other groups. In experiment 1, E2 supplementation did not promote growth, but high doses had negative effects as did Fa. In experiment 2, a greater growth response was observed only in E2-treated females in relation to higher food intake (FI) not feeding efficiency. Fa also promoted growth and FI both in females and males during the last month of the experiment. Other treatments did not affect growth, but T treatment decreased FI in males. In conclusion, the results did not provide clear evidence for E2 action on sexual growth dimorphism, but showed that testosterone may decrease growth in males by decreasing food intake in Eurasian perch. Therefore, the acceleration of male-to-female growth differences with age may not be a result of promotion of growth in females by estrogens, but a consequence of a reduction in growth by increased secretion of androgens in males.  相似文献   

7.
For 8 weeks, perch (22–25 g) were reared in a water recirculated system (WRS) and in floating cages (FC), and fed with an artificial food for trout. These two culture systems resulted in similar specific growth rates (1.1–1.4%/day). However, at 4% feeding rate, WRS ensured higher survival rates (90–100 %) vs. FC (70–80%), better food conversion indices (3.0 vs 4.1) and lower growth variability. WRS appeared more adapted to raise perch than FC. Nevertheless, sexual growth dimorphism in favour of females existed in WRS with restricted feeding rate (2 % body weight per day). Likewise, this preliminary study suggested an effect of the rearing system on the perch body composition.  相似文献   

8.
The water spider Argyroneta aquatica is the only spider spending its whole life under water, and one of the few spider species in which males are larger than females. Previous studies indicated that males can cannibalize females, which is uncommon among spiders. Here we aimed to further test for a potential influence of sexual selection on male body size. We examined the importance of female choice by testing whether females prefer the larger of two simultaneously presented males as mating partners. Further, we examined the influence of male–male competition by comparing the fighting behaviour between large and small males when alone or when together with a female, and we determined the outcome of fights. We found that females approach and choose large males as mating partners, despite the risk of male cannibalism. Additionally, males intensively compete for females, and large males clearly win against smaller ones. Hence sexual selection seems to be important for the evolution of the peculiar sexual size dimorphism of water spiders, as large size is beneficial for males in both the intra‐ and intersexual context. Previous studies have suggested an important role of natural selection in the sex‐specific body size of water spiders, but natural and sexual selection mechanisms apparently work in the same direction, favouring large male size.  相似文献   

9.
The expression of sexual dimorphism is expected to be influenced by the acquisition of resources available to allocate to trait growth, combined with sex‐specific patterns of resource allocation. Resource acquisition in the wild may be mediated by a variety of ecological factors, such as the density of interspecific competitors. Allocation may in turn depend on social contexts, such as sex ratio, that alter the pay‐off for investment in sexual traits. How these factors interact to promote or constrain the expression and evolution of sexual dimorphism is poorly understood. We manipulated sex ratio and interspecific resource competition over the growing season of red‐spotted newts (Notophthalmus viridescens) in artificial ponds. Fish competitors had a stronger effect on female than male growth, which effectively eliminated the expression of sexual size dimorphism. In addition, newt sex ratio influenced fish growth, leading to reduction in fish mass with an increase in female newt frequency. Fish also reduced the expression of male tail height, a sexually selected trait, but only in tanks with a female‐biased sex ratio. This suggests males alter their resource allocation pattern in response to the strength of sexual selection. Our results demonstrate that ecologically and socially mediated interactions between sex‐specific resource acquisition and allocation can contribute to variation in the expression of sexual dimorphism.  相似文献   

10.
Coloration plays an important role in sexual and social communication, and in many avian species both males and females maintain elaborate colours. Recent research has provided strong support for the hypothesis that elaborate female traits can be maintained by sexual or social selection; however, most research on female ornamentation has focused on pigment‐based colours, and less is known about how structural colours are maintained. Both sexes of the turquoise‐browed motmot (Eumomota superciliosa) have a blue‐green racket‐tipped tail, and it remains unknown if tail coloration serves as a sexual or social signal in one or both sexes. Here, we describe sexual dichromatism in the blue‐green portion of the tail racket, and we test for a relationship between coloration and condition, as indicated by growth bars. Tail colour of both sexes has a similar spectral shape, and there is significant, although moderate, sexual dichromatism: males are brighter than females, and males have marginally greater blue‐green saturation than females. The length of feather grown per day is positively related to overall feather brightness, but this relationship is only present in males. The relationship between male coloration and condition suggests that tail colour has the potential to convey information about individual quality during mate choice or contest competition. The lack of a similar relationship in females suggests that female tail colour does not convey the same condition‐dependent information that we suggest may be reflected by male colour. Female tail colour may therefore reflect other aspects of condition, be involved in other (non‐condition‐dependent) forms of communication, or be expressed as a non‐functional byproduct of genetic correlation between the sexes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 673–681.  相似文献   

11.
Whether premating isolation is achieved by male‐specific, female‐specific or sex‐independent assortative preferences often depends on the underlying evolutionary processes. Here we test mate preferences of males presented with females of different allopatric colour variants of the cichlid fish Tropheus sp., a Lake Tanganyika endemic with rich geographical colour pattern variation, in which the strength of sexual isolation varies between populations. We conducted two‐way mate choice experiments to compare behaviour of males of a red‐bodied morph (population Moliro) towards females from their own population with behaviour towards females from four allopatric populations at different stages of phylogenetic and phenotypic divergence. Males courted same‐population females significantly more intensely than females of other populations, and reduced their heteromorphic courtship efforts both with increasing genetic and increasing phenotypic distinctness of the females. In particular, females of a closely related red‐bodied population received significantly more courtship than either genetically distinct, similarly coloured females (‘Kirschfleck’ morph) or genetically related, differently coloured females (‘yellow‐blotch’ morph), both of which were courted similarly. Genetically and phenotypically distinct females (Tropheus polli) were not courted at all. Consistent with previous female‐choice experiments, female courtship activity also decreased with increasing genetic distance from the males’ population. Given successful experimental and natural introgression between colour morphs and the pervasive allopatry of related variants, we consider it unlikely that assortative preferences of both sexes were driven by direct selection during periods of secondary contact or, in turn, drove colour pattern differentiation in allopatry. Rather, we suggest that sexual isolation evolved as by‐product of allopatric divergence.  相似文献   

12.
While male mate choice behaviour has been reported in many taxa, little is known about its plasticity and evolutionary consequences. In the damselfly Ischnura senegalensis, females exhibit colour dimorphism (gynomorph and andromorph). The body colour of gynomorphs changed ontogenetically in accordance with sexual maturation, while little change occurred in andromorphs. To test the male mate choice between sexually immature and mature females of both morphs, binary choice experiments were conducted. Virgin males that were reared separately from females after emergence did not show significant preference between sexually immature and mature females for both morphs, indicating that virgin males were unable to discriminate female reproductive status. On the other hand, males that had experienced copulation with gynomorphs preferred sexually mature gynomorphs to sexually immature ones. However, males that had experienced copulation with andromorphs could not discriminate between sexually immature and mature andromorphs, probably due to the absence of significant ontogenetic change in their thoracic colour. Therefore, female body colour is an important cue for males in discriminating between sexual maturation stages. Learned mate discrimination depending on copulation experience might help males to detect potential mates effectively and avoid sexually unreceptive immature female. We finally discuss the adaptive significance of the ontogenetic colour change in females.  相似文献   

13.
Theory predicts marked sexual dimorphism in terms of body size and body structures used as weapons (e.g. chelipeds) in gonochoric species with intense male sexual competition for receptive females and reduced or no sexual dimorphism in species where competition among males is trivial. We tested this hypothesis using a pair of closely‐related species of symbiotic porcelain crabs as a model. In one species that inhabits sea anemones solitarily, competition among males for receptive females is unimportant. In a second species that dwells as dense aggregations on sea urchins, male–male competition for sexual partners is recurrent. We expected considerable sexual dimorphism in body size and weaponry in the urchin‐dwelling crab and reduced sexual dimorphism in the anemone‐dwelling crab. In agreement with expectations, in the urchin‐dwelling crab, male body size was, on average, larger than that of females and males invested considerably more to cheliped length than females. Also supporting theoretical considerations, in the anemone‐dwelling crab, sexual dimorphism in terms of body size was not detected and differences between the sexes in investment to cheliped length were minor. Interestingly, chelipeds were more developed both in males and females of the anemone‐dwelling crab than in the urchin‐dwelling crab as a result of the importance of these structures for monopolization of their naturally scarce anemone hosts. Another difference between the studied species was the existence of two clearly distinguishable ontogenetic phases in males of the urchin‐dwelling crab but not in males of the anemone‐dwelling crab. Whether the two different male morphs display different male reproductive strategies in the urchin‐dwelling crab remains to be addressed. Other conditions that might additionally explain the observed differences in sexual dimorphism (e.g. female mate choice) between the studied species remain to be explored. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 548–558.  相似文献   

14.
Energy investment in reproduction and somatic growth was investigated for summer spawners of the Argentinean shortfin squid Illex argentinus in the southwest Atlantic Ocean. Sampled squids were examined for morphometry and intensity of feeding behavior associated with reproductive maturation. Residuals generated from length‐weight relationships were analyzed to determine patterns of energy allocation between somatic and reproductive growth. Both females and males showed similar rates of increase for eviscerated body mass and digestive gland mass relative to mantle length, but the rate of increase for total reproductive organ weight relative to mantle length in females was three times that of males. For females, condition of somatic tissues deteriorated until the mature stage, but somatic condition improved after the onset of maturity. In males, there was no correlation between somatic condition and phases of reproductive maturity. Reproductive investment decreased as sexual maturation progressed for both females and males, with the lowest investment occurring at the functionally mature stage. Residual analysis indicated that female reproductive development was at the expense of body muscle growth during the immature and maturing stages, but energy invested in reproduction after onset of maturity was probably met by food intake. However, in males both reproductive maturation and somatic growth proceeded concurrently so that energy allocated to reproduction was related to food intake throughout the process of maturation. For both males and females, there was little evidence of trade‐offs between the digestive gland and reproductive growth, as no significant correlation was found between dorsal mantle length‐digestive gland weight residuals. The role of the digestive gland as an energy reserve for gonadal growth should be reconsidered. Additionally, feeding intensity by both males and females decreased after the onset of sexual maturity, but feeding never stopped completely, even during spawning.  相似文献   

15.
Theoretical and empirical observations generally support Darwin's view that sexual dimorphism evolves due to sexual selection on, and deviation in, exaggerated male traits. Wallace presented a radical alternative, which is largely untested, that sexual dimorphism results from naturally selected deviation in protective female coloration. This leads to the prediction that deviation in female rather than male phenotype causes sexual dimorphism. Here I test Wallace's model of sexual dimorphism by tracing the evolutionary history of Batesian mimicry-an example of naturally selected protective coloration-on a molecular phylogeny of Papilio butterflies. I show that sexual dimorphism in Papilio is significantly correlated with both female-limited Batesian mimicry, where females are mimetic and males are non-mimetic, and with the deviation of female wing colour patterns from the ancestral patterns conserved in males. Thus, Wallace's model largely explains sexual dimorphism in Papilio. This finding, along with indirect support from recent studies on birds and lizards, suggests that Wallace's model may be more widely useful in explaining sexual dimorphism. These results also highlight the contribution of naturally selected female traits in driving phenotypic divergence between species, instead of merely facilitating the divergence in male sexual traits as described by Darwin's model.  相似文献   

16.
Carotenoids are a large group of biochemicals, with similar properties, synthesised by bacteria, fungi, algae and plants. Vertebrates obtain these biologically active pigments through the diet, and they are a disproportionately common component of animal colour signals and play important roles in immune functions and as antioxidants. Carotenoids are believed to be a limited resource and because of the trade-off between allocation of carotenoids to signals and to other functions, carotenoid based signals are often thought to be handicap signals. The purpose of this study was to investigate the signalling potential of carotenoid-based tissue coloration in the great black-backed gull Larus marinus . The intensity of carotenoid-based coloration in bill, gape and eye-ring coloration was investigated in relation to body condition, reproductive parameters, levels of immune activity, and sexual dimorphism. In males there was a positive relationship between colour intensity and body condition, but in females no such relationship was found. However, females with high colour intensity had larger eggs and clutches. Additionally, females with high red scores tended to have high density of circulating lymphocytes. There was no sexual dimorphism in coloration and there was a negative relationship between colour intensity and sampling time, which indicates that this coloration is most intensely expressed early in the breeding season. The results in this study suggest that carotenoid-based coloration in great black-backed gull are partly condition dependent and reveal information about individual quality in both males an females. Hence, it might have evolved as an important signal for assessing the quality of potential mates.  相似文献   

17.
Males and females differ in body size in many animals, but the direction and extent of this sexual size dimorphism (SSD) varies widely. Males are larger than females in most lizards of the iguanian clade, which includes dragon lizards (Agamidae). I tested whether the male larger pattern of SSD in the peninsula dragon lizard, Ctenophorus fionni, is a result of sexual selection for large male size or relatively higher mortality among females. Data on growth and survivorship were collected from wild lizards during 1991–1994. The likelihood of differential predation between males and females was assessed by exposing pairs of male and female lizards to a predator in captivity, and by comparing the frequency of tail damage in wild‐caught males and females. Male and female C. fionni grew at the same rate, but males grew for longer than females and reached a larger asymptotic size (87 mm vs. 78 mm). Large males were under‐represented in the population because they suffered higher mortality than females. Predation may account for some of this male‐biased mortality. The male‐biased SSD in C. fionni resulted from differences in growth pattern between the sexes. The male‐biased SSD was not the result of proximate factors reducing female body size. Indeed SSD in this species remained male‐biased despite high mortality among large males. SSD in C. fionni is consistent with the ultimate explanation of sexual selection for large body size in males.  相似文献   

18.
Monitor lizards belong to the largest and the most sexually dimorphic lizards in terms of size, making this group an ideal model for studies analyzing ontogenetic causes of sexual dimorphism. Understanding of these ontogenetic factors is essential to the current discussion concerning patterns of sexual dimorphism in animals. We examined the ontogenetic trajectories of body weight and snout-vent length to analyze the emergence of sexual size dimorphism. Experimental animals were 22 males and 13 females of mangrove-dwelling monitors (Varanus indicus) hatched at the Prague Zoo. They were regularly weighed and measured up to the age of 33-40 months, and subsequently sexed by ultrasonographic imaging. The logistic growth equation was used to describe and analyze the observed growth patterns. Our results confirm considerable sexual size dimorphism in the mangrove monitor. The mean asymptotic body weight of males was nearly three times higher than that of females. As the body size of male and female hatchlings is almost equal, and the growth rate parameter (K) of the logistic growth equation as well as the absolute growth rate up to the age of 12 months do not differ between the sexes, size differences between fully grown males and females should be attributed to timing of the postnatal growth. Males continue to grow several months after they reach the age when the growth of females is already reduced. Therefore, the sexual size dimorphism emerges and sharply increases at this period.  相似文献   

19.
Sexual dimorphism is a major component of morphological variation across the tree of life, but the mechanisms underlying phenotypic differences between sexes of a single species are poorly understood. We examined the population genomics and biogeography of the common palmfly Elymnias hypermnestra, a dual mimic in which female wing colour patterns are either dark brown (melanic) or bright orange, mimicking toxic Euploea and Danaus species, respectively. As males always have a melanic wing colour pattern, this makes E. hypermnestra a fascinating model organism in which populations vary in sexual dimorphism. Population structure analysis revealed that there were three genetically distinct E. hypermnestra populations, which we further validated by creating a phylogenomic species tree and inferring historical barriers to gene flow. This species tree demonstrated that multiple lineages with orange females do not form a monophyletic group, and the same is true of clades with melanic females. We identified two single nucleotide polymorphisms (SNPs) near the colour patterning gene WntA that were significantly associated with the female colour pattern polymorphism, suggesting that this gene affects sexual dimorphism. Given WntA''s role in colour patterning across Nymphalidae, E. hypermnestra females demonstrate the repeatability of the evolution of sexual dimorphism.  相似文献   

20.
Despite the common occurrence of ontogenetic niche shifts, their consequences for morphological adaptations have been little studied. To address this question, we studied morphological adaptations related to ontogenetic niche shifts in Eurasian perch (Perca fluviatilis) in eight lakes that varied in density of benthic resources and planktivorous fish biomass. Perch start to feed on pelagic zooplankton, then shift to benthic resources at intermediate sizes, and finally, when large enough, mainly feed on fish. These three functional niches over ontogeny are expected to set constraints on the morphology and size-specific growth of perch. The growth of perch was negatively related to planktivorous fish biomass in the zooplanktivorous niche, but positively related to planktivorous fish biomass in the piscivorous niche. The number of gill rakers of perch was negatively related to the biomass of planktivorous fish, providing evidence for the occurrence of character displacement as a consequence of competition in the zooplanktivorous niche. Perch in lakes with low densities of predator-sensitive macroinvertebrates had greater body height measurements and a larger mouth early during ontogeny. This pattern is suggested to be a result of a selection for increased efficiency in the benthic niche when the availability of benthic resources is low. Perch in lakes with a high biomass of planktivorous fishes had fusiform body morphology, a thicker tail and a larger mouth then the average piscivorous perch. The different responses of perch morphology to variation in the availability of benthic resources compared to variation in planktivore biomass are suggested to be partly because the availability of the former resource to a larger extent is set by abiotic conditions (humic content). We suggest that the key factors affecting size-specific growth and body morphology of perch in the system studied are the availability of resources in the benthivorous and piscivorous niches. We also provide evidence for morphological trade-offs, especially between the benthivorous and the piscivorous ontogenetic niches. Received: 6 July 1999 / Accepted: 8 September 1999  相似文献   

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