Patterns of growth and sexual size dimorphism in two species of box turtles with environmental sex determination

An adaptive explanation for environmental sex determination is that it promotes sexual size dimorphism when larger size benefits one sex more than the other. That is, if growth rates are determined by environment during development, then it is beneficial to match developmental environment to the sex that benefits more from larger size. However, larger size may also be a consequence of larger size at hatching or growing for a longer time, i.e., delayed age at first reproduction. Therefore, the adaptive significance of sexual size dimorphism and environmental sex determination can only be interpreted within the context of both growth and maturation. In addition, in those animals that continue to grow after maturation, sexual size dimorphism at age of first reproduction could differ from sexual size dimorphism at later ages as growth competes for energy with reproduction and maintenance. I compared growth using annuli on carapace scales in two species of box turtles (Terrapene carolina and T. ornata) that have similar patterns of environmental sex determination but, reportedly, have different patterns of sexual size dimorphism. In the populations I studied, sexual size dimorphism was in the same direction in both species; adult females were, on average, larger than adult males. This was due in part to males maturing earlier and therefore at smaller sizes than females. In spite of similar patterns of environmental sex determination, patterns of growth differed between the species. In T. carolina, males grew faster than females as juveniles but females had the larger asymptotic size. In T. ornata, males and females grew at similar rates and had similar asymptotic sizes. Sexual size dimorphism was greatest at maturation because, although males matured younger and smaller, they grew more as adults. There was, therefore, no consistent pattern of faster growth for females that may be ascribed to developmental temperature. Received: 20 March 1996 / Accepted: 10 March 1998     

Sexual size dimorphism,growth, and maturity of the fluvial eight-barbel loach in the Kako River,Japan

The maturation and growth pattern of the fluvial eight-barbel loach Lefua sp. (Japanese name: nagare-hotoke-dojo), an endangered species, was investigated using an individual identification-recapture method from 1995 to 1998 in an upper reach of a headwater tributary of the Kako River, Hyogo Prefecture, Japan. Based on observations of the gonads through the abdominal skin, the loach was estimated to breed mostly from May to July. All the males matured by age 1⁺, and all the females matured by age 2⁺. Gamete release in all individuals of both males and females was predicted from recaptured loaches during each breeding season. The standard length of mature females was significantly larger than that of males, showing sexual size dimorphism (SSD). The maximum sizes recorded were 75.4 mm SL for females and 61.2 mm SL for males. Both males and females of immature specimens grew mainly from May to November, including the breeding season, with no significant differences in growth rates between them. After sexual maturity, both males and females grew mainly from July to October (or November), after the breeding season, and the females exhibited higher growth rates than males. Therefore, SSD of the species seems to be attributable to the different growth rates after maturity. The longevity of the loach was estimated to exceed ten years based on individual growth patterns of various sizes during the survey period. It is likely that the loach has an iteroparous life history, breeding every year, and moderate growth rates after maturity.     

Influence of age at mating on the reproductive performance of Parthenium beetle,Zygogramma bicolorata (Coleoptera: Chrysomelidae)

Abstract Age-specific mating incidence, sexual maturation and effect of age at mating on reproductive performance of the Parthenium beetle, Zygogramma bicolorata Pallister, was studied. Based on 50% mating incidence the calculated age of sexual maturation of males and females was 10.5 and 11.1 days, respectively, which was not statistically significant. However, on the basis of age at first mating, that is, sexual maturity, females matured 2 days earlier than males. Fecundity, pre-oviposition, oviposition and post-oviposition period and female longevity appear to be influenced by female age at mating with reproductive performance peaking at 30 days. On the other hand, egg viability was influenced by male age and was highest when males mated at the age of 40 days. To summarise, egg production and timing of egg deposition was female age-dependent, whereas egg fertility was male age-dependent. It was also observed that females mated at a later age and laid a higher number of eggs immediately after mating than did earlier mated females. This was ostensibly in a bid to increase fitness by maximizing reproductive output in the reduced life span available. This is the first investigation on the effect of age of females at mating on reproduction in this beetle.     

Modulation of behavior in zebrafish,Danio rerio,according to female reproductive status and visual and chemical cues

Zebrafish, Danio rerio, is one of the most studied vertebrate models. However, there are still many aspects of its reproductive behavior to be elucidated. The aim of this work was to clarify whether males and females of this species display different behaviors according to sexual status, and the kind of cues they were exposed to (visual and/or chemical). Females in two different sexual status, pre- (PreS) or post-spawning (PostS), were exposed to single males cues (visual or chemical) or complementary cues (visual and chemical together). PreS females spent more time near the male’s compartment than PostS when they were exposed to complementary cues, but no differences were found when only one cue was present. Males showed a higher proximity to females when they were exposed to visual, and both cues together, but no difference was found according to females’ reproductive status. When no barrier was present, males and PreS females spent more time swimming together than males with PostS females. These results showed that females’ reproductive status, and the kind of cues to which females and males are exposed to, can modulate behaviors related to reproduction. Particularly, both cues together (visual and chemical) are necessary to trigger different behaviors in females according to their reproductive status.     

Female assessment of male functional fertility during mate choice in a promiscuous fish

Sexual selection should favour females that can assess the functional fertility of available sexual partners and avoid mating with recently mated, sperm‐depleted males. Our current understanding of the sensory mechanism(s) underlying female assessment of males based on their functional fertility and avoidance of sperm‐depleted males is incomplete. Female Trinidadian guppies (Poecilia reticulata) are known to avoid mating with males that they had previously observed mating with other females. Here, we investigated experimentally the proximate sensory cues that they use to distinguish between paired size‐ and colour‐matched mated and unmated males in the absence of visual public information on their prior mating histories. When only water‐borne chemical cues from the males were available, females avoided the previously mated male and preferred the unmated one, but they chose randomly when only male visual cues (and no chemical cues) were available. They also preferred unmated over mated males when freely swimming with them in a more sensorially complex environment with multiple male cues (i.e., visual, chemical and mechanical cues) concurrently available. Females exhibited no preference for either stimulus males when both were unmated, irrespectively of the sensory environment. These novel results suggest that, in the absence of prior visual public information on the recent mating histories of males, female guppies use olfactory cues putatively emitted by mated males to avoid mating with them. The source and nature of the implicated olfactory cues and the fitness benefits gained by female guppies in sexually preferring males that have not recently mated remain unknown and warrant further research.     

Chemical and visual communication during mate searching in rock shrimp

Mate searching in crustaceans depends on different communicational cues, of which chemical and visual cues are most important. Herein we examined the role of chemical and visual communication during mate searching and assessment in the rock shrimp Rhynchocinetes typus. Adult male rock shrimp experience major ontogenetic changes. The terminal molt stages (named "robustus") are dominant and capable of monopolizing females during the mating process. Previous studies had shown that most females preferably mate with robustus males, but how these dominant males and receptive females find each other is uncertain, and is the question we examined herein. In a Y-maze designed to test for the importance of waterborne chemical cues, we observed that females approached the robustus male significantly more often than the typus male. Robustus males, however, were unable to locate receptive females via chemical signals. Using an experimental set-up that allowed testing for the importance of visual cues, we demonstrated that receptive females do not use visual cues to select robustus males, but robustus males use visual cues to find receptive females. Visual cues used by the robustus males were the tumults created by agitated aggregations of subordinate typus males around the receptive females. These results indicate a strong link between sexual communication and the mating system of rock shrimp in which dominant males monopolize receptive females. We found that females and males use different (sex-specific) communicational cues during mate searching and assessment, and that the sexual communication of rock shrimp is similar to that of the American lobster, where females are first attracted to the dominant males by chemical cues emitted by these males. A brief comparison between these two species shows that female behaviors during sexual communication contribute strongly to the outcome of mate searching and assessment.     

Age at maturity in wild baboons: genetic, environmental and demographic influences

The timing of early life-history events, such as sexual maturation and first reproduction, can greatly influence variation in individual fitness. In this study, we analysed possible sources of variation underlying different measures of age at social and physical maturation in wild baboons in the Amboseli basin, Kenya. The Amboseli baboons are a natural population primarily comprised of yellow baboons (Papio cynocephalus) that occasionally hybridize with anubis baboons (Papio anubis) from outside the basin. We found that males and females differed in the extent to which various factors influenced their maturation. Surprisingly, we found that male maturation was most strongly related to the proportion of anubis ancestry revealed by their microsatellite genotypes: hybrid males matured earlier than yellow males. In contrast, although hybrid females reached menarche slightly earlier than yellow females, maternal rank and the presence of maternal relatives had the largest effects on female maturation, followed by more modest effects of group size and rainfall. Our results indicate that a complex combination of demographic, genetic, environmental, and maternal effects contribute to variation in the timing of these life-history milestones.     

Flirtation reduces males’ fecundity but not longevity

Theory predicts that due to limited resources males should strategically adjust their investment in reproduction and survival. Based on different conceptual framework, experimental designs, and study species, many studies support while others contradict this general prediction. Using a moth Ephestia kuehniella whose adults do not feed and thus have fixed resources for their lifetime fitness, we investigated whether males adjusted their investment in various life activities under dynamic socio‐sexual environment. We allowed focal males to perceive rivals or additional females without physical contact. We show that males do not adjust the number of sperm they transfer to mates in a given copulation at different immediate or both immediate and mean sperm competition levels. Contradictory to general predictions, our results demonstrate that cues from additional females increase males’ investment in courtship and reduce their lifetime number of copulations and sperm ejaculated, whereas cues from rivals have no effect on these parameters. Males have similar longevity in all treatments. We suggest that the sex pheromone produced by multiple females overstimulate males, increasing males’ costly flirtations, and reducing their lifetime copulation frequency and fecundity. This finding offers a novel explanation for the success of mating disruption strategy using sex pheromones in pest management.     

Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies

The evolution of sexual cannibalism has been modelled as both an adaptive and nonadaptive female strategy. Recent evidence from several species suggests a connection between female foraging and sexual cannibalism, but the precise benefits for females have remained obscure. Here, we investigate the difference between cannibalistic and noncannibalistic female Nephila plumipes by removing the potential nutritional benefit of cannibalism. Courting and mating males that were killed by a female were immediately removed so that the female could not consume them. Nevertheless, cannibalistic females gained more mass from maturation to oviposition and produced larger first clutches than noncannibalistic females, although cannibalistic females matured at a smaller size and mass than noncannibalistic females. In juvenile instars, mass gain was generally smaller in females that moulted in a good condition but intermoult intervals were shorter. However, the time from maturity to oviposition was not shorter in females that matured in a good condition. Male behaviour did not differ according to the risk of cannibalism. We suggest that sexual cannibalism in N. plumipes is a side‐effect of an increased foraging vigour of females that matured at a smaller size and body mass. Selection pressure on males to avoid cannibalism may be weak because of limited mating opportunities.     

Seasonal effects on sexual maturation of male bank voles (Clethrionomys glareolus)

The weights of the testes and accessory glands, and the degree of morphological and functional development of the seminiferous tubules, were used as indicators of sexual maturation in male bank voles. Males reared under constant laboratory conditions showed a significant relationship between the seasons and sexual maturation. Young males reached maturity most rapidly in the reproduction season (mid-April to mid-October), while adult males matured earlier, in the spring season (mid-January to mid-April). The influence of season on animals reared in outdoor cages was much more pronounced. One of the important factors was the photoperiod: animals reared from 3 to 12 weeks of age in a short photoperiod (8L:16D) matured less rapidly than did those reared in 16L:8D.     

Adult diet and male-female contact effects on female reproductive potential in Mexican fruit fly (Anastrepha ludens Loew) (Diptera tephritidae)

Wild strains of fruit flies (Diptera: Tephritidae) placed into laboratory rearing conditions are subjected to selection pressures caused by the diet, cages, density of flies, and other factors. Selection that changes mating behavior of the strain may result in less effective males released in sterile insect programs. Tests were performed to examine the effects of protein in diet and adult interactions on egg production and mating during sexual maturation of the Mexican fruit fly (Anastrepha ludens Loew) in laboratory cages. Flies were offspring of wild flies collected from Chiapas or Nuevo Leon, Mexico, and reared on Valencia oranges. Experiments demonstrated effects of yeast hydrolysate protein in adult diet and pairing with males on production of mature and immature eggs, numbers of females producing eggs, and mating with females aged 15 d. Addition of protein to 4% fructose in the adult diet approximately tripled mature egg production in females maintained for the total maturation period with an equal number of males. Females that matured without males produced approximately 33% more-mature eggs when fed protein than those fed no protein. Total egg production of females matured without males and fed sugar only or sugar with protein was more than twice that of females matured with males. Tests to examine the effects of male and female diet separately on female egg production showed slightly higher egg production in females fed protein, or females paired with males fed protein, but these differences were not significant. The most definitive effects were that combining wild strain females and males in cages during maturation reduced egg production. This effect was greatest when flies were not fed protein.     

Social influences on reproductive development and fertility in female Djungarian hamsters (Phodopus campbelli).

Social influences on the sexual maturation of female Djungarian hamsters were investigated in two experiments. In the first experiment females were housed from weaning with an adult male, by themselves, or with a weanling sister. Maturation was accelerated in females housed with males as indicated by younger age at first ovulation, increased rates of ovarian and uterine growth, and lower LH levels at some ages. Maturation was delayed in females housed with sisters compared to those housed alone as measured by time of first ovulation and by lower estradiol levels at some ages. The most marked differences between groups occurred 8 to 12 days after weaning, suggesting that events during this period are particularly important in the social mediation of sexual maturation. In the second experiment the effects of reproductive suppression (caused by living with a sister) on the subsequent fertility of females housed with males were examined. If male-female pairs were housed in clean cages, no effects were observed; however, pairs housed in cages previously soiled by the female and her sister had fewer young surviving until 1 week of age despite no differences in the age of pregnancy onset or in the initial litter size. Thus, even cues present in unrenewed soiled bedding may have subtle but long lasting effects on reproductive function.     

Maturation costs affect maturation timing: sexual reproduction in a heterogonic hydra

If maturation is more costly for females, they may need more distinct environmental cues to induce sexual reproduction than males. We verified this hypothesis by comparing the indirect costs of maturation to males and females of the heterogonic Hydra oligactis, reproducing both asexually and sexually. The laboratory experiments revealed that males mature 2 weeks earlier than the first females at falling temperatures simulating the natural conditions that precede sexual reproduction. The difference between the energy costs of maturation for males versus females has been considered a likely factor responsible for the observed difference in maturation time. Available food supply positively affected the percentage of sexually mature females, indicating that females are more sensitive to food limitation than males. The number of gonads was correlated positively with the size of mature hydra for both males and females. However, males produced twice as many testes as ovaries produced by females. We postulate that females are induced later than males in order to prevent gonadal development after an unseasonable drop in temperature. As sexual reproduction in H. oligactis interferes with asexual budding, under favorable conditions for asexual proliferation unnecessary gonadal development decreases an individual’s fitness through reduction of the number of produced offspring.     

Influence of Male Presence on Sexual Maturation in Female Caribbean Fruit Fly, Anastrepha suspensa (Diptera: Tephritidae)

The presence of males was shown to affect the rate of female ovarian development in the Caribbean fruit fly, Anastrepha suspensa (Loew). Virgin females were maintained either in the absence or presence of males. Those sharing a common space with males had either visual contact with the opposite sex or no visual contact. Three strains of Caribbean fruit fly were tested: mass reared strain (flies in colony for more than 20 years); semi-wild strain, flies recently adapted to the laboratory conditions (ca. 12 months); and a wild strain collected in the field. We found that: (1) Mass reared, semi-wild, and wild strains had different female maturation rates, as measured by the presence of mature oocytes, regardless of male presence. (2) Male presence accelerated maturation in wild females and to a lesser extent in semi-wild flies, but had no effect on the long-domesticated mass reared strain. (3) A barrier between female and male cages removed any possibility of visual communication but had no effect on the males’ effect on female maturation. We discuss the adaptive significance of facultative ovarian maturation and the use of male-produced cues to regulate sexual development, and comment on the rapid rate of selection on female maturation under mass-rearing conditions.     

How do sex and sexual maturation influence the response of Monochamus galloprovincialis to host odours?

Finding the right host plant for feeding and reproduction is crucial for herbivorous insects. Usually they need both visual and olfactory cues to locate and colonize their host plants, but the respective role of the two types of stimuli is difficult to disentangle. Model plants mimicking the shape and colour of natural hosts, combined with odours dispensers, are relevant tools for this purpose. We used dummy pines and cut branches of natural pines to investigate host attraction in Monochamus galloprovincialis, the insect vector of the pine wood nematode in Europe. We compared the behavioural response of males and females, sexually mature or immature, in a large climatic chamber and in an olfactometer. Immature females were significantly attracted by the sole odours released by pines. Immature males and mature males and females needed a combination of visual and olfactory cues to find the host tree. These findings suggest that beetles use both visual and chemical cues to locate pine trees but that relevant stimuli likely differ between beetles of different sex and sexual maturation status. Our results question the implementation of precautionary clear‐cuts around infested trees for limiting the spread of PWN.     

Growth differences and dimorphic expression of growth hormone (GH) in female and male Cynoglossus semilaevis after male sexual maturation

Half-smooth tongue sole, Cynoglossus semilaevis, is an ideal model to investigate the regulatory mechanisms of sexual growth dimorphism in fish species. The aim of the study was to investigate the effect of differential age of sexual maturity for females and males on growth and GH mRNA expression in C. semilaevis. The body weight differences between the sexes were not significant in C. semilaevis at age 5 months when females and males were all immature. Significant differences in body weight between the sexes were found after early sexual maturation of males at the age of 9 months. The body weight of 21-month-old females (621.4 ± 86.4g), still not immature, was even 3.28 times higher than that of the males (189.7 ± 14.4g). The cDNAs encoding GH in C. semilaevis was cloned. The GH gene is 2924bp long and consists of six exons and five introns. The results of qRT-PCR showed that GH mRNA levels of the immature females were not significantly different from that of immature males at age 5 months. However, GH mRNA levels of the immature females were significantly higher compared with those of the mature males at age 9 months (P<0.05). At age 11 months, GH mRNA levels of females were even 6.4-fold higher than that of males. In conclusion, for the first time we show that early sexual maturity of males is the main cause of sexual growth dimorphism in C. semilaevis and exert significant effect on GH mRNA expression.     

Maturation of walleye by age, size and surplus energy

The probability of annual sexual maturation by male and female walleye Stizostedion vitreum was related to age, size and an index of condition, I _VF=[arcsine(visceral fat)^0·5(body mass)^−0·5]. Most males first matured at ages 2 and 3 years; size explained first maturation, but condition explained later maturation. In contrast, most females first matured at ages 4 and 5 years; maturity of females was more dependent upon condition. Maturity of females at ages 4 and 5 years was significantly correlated with average I _VF of the population ( PI _VF). The size reached by age 2 years (early growth) was correlated with the PI _VF. Growing degree-days, Secchi depth, latitude and lake morphology were not correlated with the PI _VF. Annual variations in female spawning stock size were related to the condition of the females, presumably reflecting the net acquisition of energy in the preceding growing season. Annual variations within lakes in the net acquisition of energy may exceed the variations in energy availability between lakes, dictated by lake morphology and geography. Thus, assessment of condition could be used to predict annual potential spawning stock size and egg production.     

Differences in growth,size and sexual dimorphism in skulls of East Greenland and Svalbard polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>)

Size, growth and sexual dimorphism of nine skull traits was studied in 300 East Greenland and 391 Svalbard polar bears (Ursus maritimus). Two traits were significantly larger in bears from East Greenland compared to Svalbard bears, and trait size was smaller after 1960 in five traits. For both localities and both age groups (subadult, adult), mean trait size values were higher in males than females (all: P < 0.05). Gompertz growth models showed trait size increasing with age in seven traits. Depending on the trait, males reached 95% asymptotic trait size at age 3–10, females at age 2–6. The females of both localities matured at approximately the same age, whereas the Svalbard males generally matured years later than their East Greenland peers. The differences found in the present study between the two polar bear subpopulations support the notion that East Greenland and Svalbard polar bears probably should be managed as separate units.     

Sex steroids modulate changes in social and sexual preference during juvenile development in zebra finches

Zebra finches, like many other animals, have close social relationships mainly with the family at young ages but begin to express interest in opposite-sex extra-family animals as they enter the late juvenile period and sexual maturity. This experiment tested a set of hypotheses that sex steroids are involved in this developmental transition. At 25-30 days, subjects were implanted subcutaneously with Silastic tubes that were empty (controls), filled with testosterone propionate, filled with estradiol benzoate, or filled with a combination of ATD (an aromatization inhibitor) and flutamide (an anti-androgen). Once a week between ages 30 and 90 days, they were given three-choice tests where the three stimulus types were the family members, unpaired males, or unpaired females. The preferred category was defined as the one adjacent to the proximity zone in which the subject spent the most time. Control males were more likely to prefer females and less likely to prefer the family as they got older, and control females were increasingly likely to prefer males. Males treated with testosterone or estradiol showed a premature increase in preferences for females. Females treated with ATD plus flutamide failed to show the normal increase in preferences for males shown by controls. These results indicate an involvement of sex steroids in the maturation of sexual preferences in a socially monogamous species that relies on visual and auditory, rather than olfactory, cues for sexual or other social behavior.     

Sex steroids modulate changes in social and sexual preference during juvenile development in zebra finches

Zebra finches, like many other animals, have close social relationships mainly with the family at young ages but begin to express interest in opposite-sex extra-family animals as they enter the late juvenile period and sexual maturity. This experiment tested a set of hypotheses that sex steroids are involved in this developmental transition. At 25–30 days, subjects were implanted subcutaneously with Silastic tubes that were empty (controls), filled with testosterone propionate, filled with estradiol benzoate, or filled with a combination of ATD (an aromatization inhibitor) and flutamide (an anti-androgen). Once a week between ages 30 and 90 days, they were given three-choice tests where the three stimulus types were the family members, unpaired males, or unpaired females. The preferred category was defined as the one adjacent to the proximity zone in which the subject spent the most time. Control males were more likely to prefer females and less likely to prefer the family as they got older, and control females were increasingly likely to prefer males. Males treated with testosterone or estradiol showed a premature increase in preferences for females. Females treated with ATD plus flutamide failed to show the normal increase in preferences for males shown by controls. These results indicate an involvement of sex steroids in the maturation of sexual preferences in a socially monogamous species that relies on visual and auditory, rather than olfactory, cues for sexual or other social behavior.