Age and growth of the sandbar shark, Carcharhinus plumbeus, in Hawaiian waters through vertebral analysis

Age and growth estimates were determined for the sandbar shark, Carcharhinus plumbeus, from Oahu, Hawaii in the central Pacific Ocean. Age estimates were obtained through vertebral centra analysis of 187 sharks. We verified our age estimates through marginal increment analysis of centra and oxytetracycline marking methods of at liberty sandbar sharks. Sizes of sampled sharks ranged from 46 to 147 cm pre-caudal length. Four growth models were fitted to length-at-age data; two forms of the von Bertalanffy growth model, the Gompertz growth model, and a logistic growth model. Males and females exhibited statistically significant differences in growth, indicating that females grow slower and attain larger sizes than males. Growth parameter estimates revealed slower growth rates than previously estimated (based on captive specimens) for Hawaiian sandbar sharks. The von Bertalanffy growth model using empirical length-at-birth provided the best biological and statistical fit to the data. This model gave parameter estimates of L _∞ = 138.5 cm PCL and k = 0.12 year⁻¹ for males and L _∞ = 152.8 cm PCL, k = 0.10 year⁻¹ for females. Male and female sandbar sharks mature at approximately 8 and 10 years of age, respectively.     

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.     

Age, growth, and sexual maturity of the yellownose skate Dipturus chilensis in the south‐eastern Pacific

The age and growth parameters of Dipturus chilensis were estimated by counting growth rings from thin sections of vertebral centra from 400 fish (246 females and 154 males), ranging from 23 to 124 cm total length (L_T), and backcalculating fish lengths at previous ages. Marginal increment analysis lent support to the hypothesis of annual deposition of band‐pairs, which formed during the winter months. The oldest female D. chilensis aged in this study was 21 years and 117 cm L_T, whereas the oldest male was 18 years and 93 cm L_T. A 4·7% index of average per cent error (I_APE) suggested that this is a precise method for calculating the age of D. chilensis. Observed L_T were lower than backcalculated L_T, which implies the influence of Lee's phenomenon. The von Bertalanffy growth equations, based on mean length‐at‐age data, were estimated as L_t = 128·3 (1 ? e^{?0·112 (t + 0·514)}) for females and L_t = 107·8 (1 ? e^{?0·134 (t + 0·862)}) for males where t is age (years). Growth was significantly different between sexes: females reached a larger adult size. Ages and lengths at 50% maturity were estimated at 14 years of age and 106 cm L_T for females and 11 years of age and 86 cm L_T for males. At c. 14 years, there was a decline in growth rates in females. The factor most likely responsible for this was sexual maturity, which caused a discontinuity in growth of female fish. These results show that this species is slow‐growing, long‐lived, relatively large and of delayed maturity, characteristics that make it vulnerable to exploitation.     

Age,growth and reproductive traits of invasive goby Taenioides cirratus in the Chaohu Lake,China

The eel goby Taenioides cirratus (Blyth, 1,860) is a small fish inhabits muddy bottoms of brackish-water in the Indo-West Pacific. It has invaded many inland freshwater lakes in China, such as the Chaohu Lake, Gaoyou Lake and Nansi Lake, and its population increased rapidly in these freshwater lakes in recent years. The age, growth and reproductive traits of T. cirratus invading the Chaohu Lake were studied. A total of 482 specimens (210 females, 204 males and 68 juveniles) with total length (TL) ranging from 9.4 to 20.6 cm were collected using the benthic fyke nets at monthly intervals from March 2018 to February 2019. The sagittal otolith was used for age determination. Monthly variation of marginal increment ratio indicated that the annual forming of opaque band on sagittal otolith was completed during March and April. For both sexes, only four (from 0+ to 3+ years) age groups were observed and 1+ and 2+ years age individuals dominated the population. Back calculated length at age showed males grew faster than females. Both sexes reached maturity at 1+ year age and the TL at first maturity (TL₅₀) was 12.6 cm for females and 11.9 cm for males. Monthly variation of gonado-somatic index indicated that the spawning occurred from May to August. The fecundity ranged from 967 ova to 5,114 ova, with a mean of 3,205 ova. Our study provides a comprehensive data on the key life history traits of T. cirratus for the first time.     

Life‐history traits of the leatherjacket Meuschenia scaber,a long‐lived monacanthid

The present study describes the age and growth of the leatherjacket Meuschenia scaber, a common Australasian monacanthid and valued by‐catch of the inshore bottom trawl fishery in New Zealand. Age was determined from the sagittal otoliths of 651 individuals collected between July 2014 and March 2016 in the Hauraki Gulf of New Zealand. Otolith sections revealed alternating opaque and translucent zones and edge‐type analysis demonstrated that these are deposited annually. Meuschenia scaber displayed rapid initial growth, with both males and females reaching maturity in 1–2 years and 50% of both sexes matured at 1·5 years. Maximum age differed substantially between the sexes, at 9·8 years for males and 17·1 years for females. Growth rate was similar between sexes, although males reached greater mass at age than females in the early part of the lifespan. The length–mass relationship differed significantly between the sexes, with males displaying negative allometric growth and females isometric growth. Female condition was highest in July, declined in August with the onset of spawning and showed a slight peak in January and February, immediately following the spawning season. This study substantially extends the maximum longevity recorded for monacanthids, although males had much shorter lifespans and higher mortality, than females.     

Age and growth estimates of the bull shark,Carcharhinus leucas,from the northern Gulf of Mexico

Synopsis Length at age and growth rates for 59 bull sharks, Carcharhinus leucas, collected from the northern Gulf of Mexico were estimated from the band patterns formed seasonally in the vertebral centra. The combined age at length data for both sexes were applied to a von Bertalanffy growth model producing parameter estimates of L = 285 cm TL, K = .076, t₀ = –3.0 yr. Lengths at age for males and females were similar except that males did not attain as great a length as females. Growth was apparently slow and varied among individuals, but in general, was estimated to be 15–20 cm yr^–1 for the first five years, 10 cm yr^–1 for years 6–10, 5–7 cm yr^–1 for years 11–16, and less than 4–5 cm yr^–1 thereafter. Males mature at 210–220 cm TL or 14–15 yr of age; females mature at>225 cm TL or 18+ yr of age. The largest male (245 cm TL) was 21.3 yr old; the largest female (268 cm TL) was 24.2 yr old.     

Age and growth of the tiger shark Galeocerdo cuvier off the east coast of Australia

Total lengths (L_T) at age and growth rates for south‐west Pacific Galeocerdo cuvier were estimated from vertebral growth‐band counts of 202 sagitally sectioned centra from 112 females (71–430 cm L_T), 79 males (72–351 cm L_T) and 11 of unknown sex. Captive growth data were also examined to complement vertebral age estimations. The sexes combined modelled growth coefficient (k = 0·08) was smaller than previously reported for G. cuvier populations elsewhere. Split‐band and narrow banding patterns were identified as potential sources of age underestimation in this species.     

Using cross-dating techniques to validate ages of aurora rockfish (<Emphasis Type="Italic">Sebastes aurora</Emphasis>): estimates of age,growth and female maturity

Since fishery management regulations have shifted much of the groundfish trawl effort in the northeastern Pacific from the continental shelf to the slope, fishery impacts on unassessed demersal slope rockfish species like the aurora rockfish (Sebastes aurora) may have increased. Understanding the life history of these species is a critical first step in developing management strategies to protect them from overharvest. In this study we employ cross-dating methods to validate the annual periodicity of growth increments and investigate the age, growth and maturity of aurora rockfish, a species for which life history information is quite limited. Specimens were collected on an opportunistic basis from Oregon commercial landings and from research cruises, over the years 2003–2006. Age was estimated for 438 individuals using otoliths processed via the break-and-burn method. The maximum estimated age was 118 years for females (n = 324) and 81 years for males (n = 114). The von Bertalanffy growth function showed that males grow faster and reach a smaller maximum size than females (males: L _inf = 34, K = 0.09, t ₀ = −1.9; females: L _inf = 37, K = 0.06, t ₀ = −5.5), though both sexes demonstrate relatively slow growth. Visual assessment of ovaries showed that the aurora rockfish is a synchronous spawner with parturition occurring in May and June off Oregon. Female age and length at 50% maturity were calculated at 12.6 years and 26 cm, respectively (n = 307). Maturity and age data provided evidence for a protracted adolescence in this species.     

Length‐ and age‐based demographics of exploited populations of stout whiting,Sillago robusta Stead, 1908

This study assessed variability in the length and age compositions, longevity, length‐at‐age and rates of growth and mortality of the east Australian stout whiting Sillago robusta Stead, 1908 population harvested by demersal trawl fisheries. Sampling was done over 2 years and was spatially stratified across three depth strata between 11 and 90 m at two locations approximately 400 km apart. There were no consistent depth‐related differences in length and age compositions, but the mean and median length and age of the population was greater at the lower latitude location. Age classes 2 and 3 years dominated samples in the north, and 1 and 2 years in the south. Observed longevity was 10 years in the north, and 6 years in the south. Mean length‐at‐age was not consistently different between sexes, years or locations, nor did the von Bertalanffy growth function differ significantly between sexes, even though females had a greater estimated L_∞ (23.45 cm FL) compared to males (22.36 cm FL). Estimated natural mortality (M) ranged between 0.42 and 0.77, using age‐ and length‐based methods. Age‐based catch‐curve analyses identified the instantaneous rate of total mortality (Z) to range between 1.48 and 2.70, with subsequent estimates of fishing mortality (F) ranging between 1.15 and 2.00, being greater than M. Exploitation rates (E) were greater than 0.7, indicating that S. robusta at the study locations was heavily fished. The data provided here can be used as a basis to evaluate future fishery‐ and climate‐related changes in the population demographics of east Australian S. robusta.     

Age,growth and reproduction of the white seabream Diplodus sargus sargus (Linneaus, 1758) off the eastern coast of Algeria

The objective of this study was to determine the basic population‐specific parameters necessary for fish stock assessment in the area and to compare these with data from other Mediterranean regions. White seabream Diplodus sargus sargus (n = 449) were caught along the Algerian coast between December 2005 and December 2006. Total length ranged from 12.2 to 34.6 cm, representing age classes between 0 and 10 years. Validity of the otolith readings for estimating age and growth was supported using the back‐calculation method. The von Bertalanffy growth parameters for all fish were calculated as: TL = 36.3 cm, k = 0.154 year⁻¹, t₀ = −0.488 year. The growth performance index (φ) showed a relatively slow growth of the local population. The length‐weight relationship showed an isometric growth (b = 2.98; r = 0.98). Subspecies Diplodus sargus sargus was characterized as being a proterandric hermaphrodite. Overall ratio of males to females was 1 : 1.4, with males predominant in the smaller size intervals and females in the larger ones. The reproductive season extended from January to May, with a March–April peak in spawning activity. Fifty per cent maturity of the tested cohort was reached at a total length of 20.2 cm for males and 20.0 cm for females.     

Growth and Development in Wild Owl Monkeys (<Emphasis Type="Italic">Aotus azarai</Emphasis>) of Argentina

Life history predicts that in sexually dimorphic species in which males are the larger sex, males should reach sexual maturity later than females (or vice versa if females are the larger sex). The corresponding prediction that in sexually monomorphic species maturational rates will differ little between the sexes has rarely been tested. We report here sex differences in growth and development to adulthood for 70 female and 69 male wild owl monkeys (Aotus azarai). In addition, using evidence from natal dispersal and first reproduction (mean: 74 mo) for 7 individuals of known age, we assigned ages to categories: infant, 0–6 mo; juvenile, 6.1–24 mo; subadult, 24.1–48 mo; adult >48 mo. We compared von Bertalanffy growth curves and growth rates derived from linear piecewise regressions for juvenile and subadult females and males. Growth rates did not differ between the sexes, although juvenile females were slightly longer than males. Females reached maximum maxillary canine height at ca. 2 yr, about a year earlier than males, and females’ maxillary canines were shorter than males’. Thus apart from canine eruption and possibly crown–rump length, the development of Azara’s owl monkeys conforms to the prediction by life history that in monomorphic species the sexes should develop at similar paces.     

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.     

Movements,reproductive seasonality,and fisheries interactions in the whitetip reef shark (<Emphasis Type="Italic">Triaenodon obesus</Emphasis>) from community-contributed photographs

Despite being a common apex-level predator on coral reefs throughout the tropical Indo-Pacific, surprisingly little is known about whitetip reef shark (Triaenodon obesus) movements and biology. This study used photo-identification from community-contributed photographs to reveal patterns in movements, reproductive biology, and fisheries interactions in this species that have not been previously revealed through more traditional methods. At least 178 individual sharks were identified, and 26 movements were observed. These included movement distances of up to 26.4 km, movement rates of up to 3.27 km/day (9.8 km in 3 days), and movements that required the transit of a 140 m deep channel. Other animals showed high philopatry, being re-sighted at the same locality on multiple occasions (up to 13 sightings for one individual) over periods of up to 7 years. Females showed higher philopatry than males and were more likely than males to be found at shallow (<10 m depth) localities throughout the year. The proportion of male sightings at shallow localities was significantly higher in April and May than other months of the year, possibly due to males coming into the shallows to mate with females. A peak in sightings of late-term females followed by an abrupt decline suggests that pupping season is May into early June, and two females were observed pregnant in consecutive years despite evidence that the gestation period is approximately 1 year for this species. Nine percent of animals carried fishing tackle or exhibited jaw injuries associated with fishery interactions, with multiple individuals found dead after being hooked by fishers.     

Biological traits of naturally induced hybrid individuals of two gizzard shads, <Emphasis Type="Italic">Nematalosa come</Emphasis> and <Emphasis Type="Italic">N. japonica</Emphasis>, in coastal waters around Okinawa Island,Ryukyu Archipelago,southwestern Japan

The age, growth, reproductive condition, and occurrence of natural hybrids of two Nematalosa species around Okinawa Island were examined using 128 specimens obtained from April 2003 to June 2004. Standard length (SL) reached approximately 150–210 mm within the first 2 years, and then remained stagnant. The maximum age for both sexes was ca. 5 years old. Maturity sizes and ages were estimated to be at least 173.2 mm SL and 2 years old for females and 192.6 mm SL and 3 years old for males. Spawnable individuals were mainly observed from January to March based on histological observations of gonads. Natural hybrids appeared at all sampling sites except for the Haneji Inlet and were dominant at Makiminato (in south-central Okinawa Island). Their incidence was also quite high (66.9%) in the Makiminato population, when compared with records for other marine fishes around Japan. In Okinawa Island, these shallow areas are rapidly decreasing in size because of recent reclamation and land exploitation. Hybrid production may be caused by not only the reproductive biology and sympatric distributions of the parent species but also recent environmental changes.     

Sex ratio and gender-dependent neighboring effects in <Emphasis Type="Italic">Podocarpus nagi</Emphasis>, a dioecious tree

We analyzed sex ratio, growth rates, and spacing among individuals of Podocarpus nagi, a dioecious tree, on Mt. Mikasa, Nara City, Japan. The sex ratio of reproductive trees ≥ 5 cm in stem diameter at breast height (dbh, 130 cm above ground level) was significantly male-biased. The sex ratio was male-biased in the < 20 cm and ≥ 50 cm size classes, while it did not depart from 1:1 in the 20 ≤ dbh < 50 cm class. Growth rate varied with tree size in males but not in females. The precocity and vigor of males suggests that differences in reproductive costs between sexes induce the biased sex ratio. Random labeling tests on the positions of reproductive trees showed that in the < 30 cm class, males and females were distributed randomly and independently from each other. In the ≥ 30 cm class, males were significantly clumped, whereas females were randomly distributed. Males and females showed significant repulsion, i.e., a spatial segregation of sexes. Both intra- and intersexual effects on the growth rate of crowding by neighbors were significant for females, but not for males. Maximum competitive interference was observed at a distance of 5 m, which corresponded approximately to the radius of clumps of large males and to the significant repulsive distance between large males and females. These results suggest that sexual differences in sensitivity to local crowding are related to the formation of gender-dependent spatial patterns. Formation of female-repulsive male clumps and a male-biased sex ratio may intensify the decreased probability of regeneration near males, as suggested by the limited seed-dispersal range of this species, thereby promoting coexistence with other species.     

Age, growth, maturity, and mortality of the Alaska skate, Bathyraja parmifera, in the eastern Bering Sea

The Alaska skate, Bathyraja parmifera, is the most abundant species of skate on the eastern Bering Sea shelf, accounting for over 90% of total skate biomass. However, little is known regarding the life history of this species despite its common occurrence as bycatch in several Bering Sea fisheries. This is the first study to focus on the age and growth of B. parmifera. From 2003 to 2005, more than one thousand specimens were collected by fisheries observers and on scientific groundfish surveys. Annual banding patterns in more than 500 thin sections of vertebral centra were examined for age determination. Caudal thorns were tested as a potentially non-lethal ageing structure. Annual band pair deposition was verified through edge and marginal increment analyses. A three-parameter von Bertalanffy growth function and a Gompertz growth function were fit to observed length-at-age data. Both models provided significant fits, although the Gompertz function best described the overall pattern of growth in both males and females, based upon statistical criteria and parameter estimates. Age and size at 50% maturity were 9 years and 92 cm TL for males and 10 years and 93 cm TL for females. The maximum observed ages for males and females were 15 years and 17 years, respectively. Estimates of natural mortality (M) ranged from 0.14 to 0.28, and were based on published relationships between M and longevity, age at maturity, and the von Bertalanffy growth coefficient. Due to these life history characteristics and a lack of long-term species-specific stock data, a conservative management approach would be appropriate for B. parmifera.     

Age and growth of ocellated icefish, Chionodraco rastrospinosus DeWitt and Hureau, 1979, from the South Shetland Islands

Age and growth of ocellated icefish, Chionodraco rastrospinosus, were investigated using counts of annual growth increments from sagittal otoliths. Samples were collected during research surveys by benthic trawl carried out around Elephant and South Shetland Islands in January–February 2002 and December 2006–January 2007. A total of 290 specimens were selected for the study, consisting of 120 females and 170 males. The age of fish was estimated by counting annuli on transverse sections obtained by grinding and polishing whole otoliths embedded in epoxy resin. The precision-of-age estimates within and between readers were tested applying both the average percent error (APE) and the coefficient of variation (CV). The estimated age-range was 1–12 for both sexes of C. rastrospinosus. Applying the von Bertalanffy growth function to the age–length data, a growth curve was obtained for each sex. The estimated values of VB growth parameters L _∞ and k were, respectively, 47.9 cm and 0.28 for females and 42.9 cm and 0.36 for males. Compared to other congeneric species, the growth performance of C. rastrospinosus was relatively high, being 2.82 and 2.81 in males and females, respectively. Age at sexual maturity was estimated to be about 4 years in both sexes. C. rastrospinosus captured in the studied area consisted mainly of adult specimens between 3 and 8 years, with few older fish.     

Ecology of the growth of Anolis nebulosus (Squamata: Dactyloidae) in a seasonal tropical environment in the Chamela region,Jalisco, Mexico

Juvenile growth rates are thought to be restricted by available food resources. In animals that grow throughout the year, such as tropical lizards, growth is therefore predicted to be faster during the rainy season. We test this prediction using a population of Anolis nebulosusby describing the growth trajectories of both sexes using nonlinear regression models, and we then correlate the growth rates of individuals with food available in the environment, precipitation, and temperature. The Von Bertalanffy model fits the growth rates of the females better, while the logistic‐by‐length model fits the males better. According to both models, the males grew faster than females, reaching slightly smaller sizes at adulthood. Males reached sexual maturity when 35 mm long, at an age of seven months, and females matured at 37 mm (SVL), taking nine months to reach this size. In 1989, juvenile males and females grew more in both seasons (rainy and dry) than adults; for 1990, there were no differences by season or between age classes. These results are interesting since in the 1989 and 1990 rainy seasons, practically the same orders of prey and the greatest abundance of prey available in the environment were registered. A possible explanation could be that predation was more intense in 1990 than in 1989. There is little evidence that food, temperature, and humidity affect growth rates of A. nebulosus, refuting our predictions. This is mainly due to the low variation in growth observed in 1990. Therefore we think that the growth of this species reflects a complex combination of ecological and genetic factors.     

Length-based growth,maturity and natural mortality of the cockfish <Emphasis Type="Italic">Callorhinchus callorhynchus</Emphasis> (Linnaeus, 1758) off Coquimbo,Chile

The growth rate, reproductive aspects, and natural mortality of chimaeras and ratfish are poorly known. In this study, life-history parameters for cockfish Callorhinchus callorhynchus (Holocephali—Callorhinchidae) are estimated, which is an important fish resource exploited in Chile. Specimens were sampled from the artisanal fishery captures, from November 2006 to November 2007. The standard length (SL) of males fluctuated between 20 and 62 cm, and between 21 and 70 cm for females. Von Bertalanffy growth parameters were estimated through length-frequency data analysis using MULTIFAN. The length-weight relationship and von Bertalanffy growth parameters were significantly different for males and females, as well as the length at 50% maturity. For males a model with 5 age-classes was the best, with asymptotic length L _∞ = 52 cm SL, growth coefficient K = 0.473 yr⁻¹, and age at length zero t ₀ = −0.690 yrs. For females the best model was represented by 10 age-classes (L _∞ = 70.3 cm SL, K = 0.193 yr⁻¹, t ₀ = −1.158 yrs) in the length-frequency data sets. Length at 50% maturity of males was estimated in 43.7 cm SL, and in 50.2 cm SL for females. The natural mortality rate fluctuated between 0.42 and 0.82 yr⁻¹ for males and between 0.12 and 0.37 yr⁻¹ for females, depending upon the method used. It is concluded that C. callorhynchus is a species with life-history parameters significantly different between males and females, and such differences should be taken into account in future population dynamics analysis.     

Sex‐specific compensatory growth in the larvae of the greater wax moth Galleria mellonella

Deficiency of food resources in ontogeny is known to prolong an organism's developmental time and affect body size in adulthood. Yet life‐history traits are plastic: an organism can increase its growth rate to compensate for a period of slow growth, a phenomenon known as ‘compensatory growth’. We tested whether larvae of the greater wax moth Galleria mellonella can accelerate their growth after a fast of 12, 24 or 72 h. We found that a subgroup of female larvae showed compensatory growth when starved for 12 h. Food deficiency lasting more than 12 h resulted in longer development and lower mass gain. Strength of encapsulation reactions against a foreign body inserted in haemocoel was the weakest in females that showed compensatory growth, whereas the strongest encapsulation was recorded in the males and females that fasted for 24 and 72 h. More specifically, we found sex‐biased immune reactions so that females had stronger encapsulation rates than males in one group that fasted for 72 h. Overall, rapidly growing females had a short larval development period and the shortest adult lifespan. These results suggest that highly dynamic trade‐offs between the environment, life‐history traits and sex lead to plasticity in developmental strategies/growth rates in the greater wax moth.