Five new species of dicyemid mesozoans (Dicyemida: Dicyemidae) from two Australian cuttlefish species, with comments on dicyemid fauna composition

Five new species of dicyemid mesozoans in two genera are described from two Australian cuttlefish species, Sepia apama Gray (giant Australian cuttlefish) and S. novaehollandiae Hoyle (nova cuttlefish): Dicyema coffinense n. sp. from S. apama collected from Coffin Bay, South Australia (SA), Australia; D. koinonum n. sp. from S. apama and S. novaehollandiae collected from Gulf St Vincent (GSV) and Spencer Gulf (SG), SA, Australia; D. multimegalum n. sp. from S. apama collected from Cronulla and North Bondi, New South Wales, Australia; D. vincentense n. sp. from S. novaehollandiae collected from GSV, SA, Australia; and Dicyemennea spencerense n. sp. from S. novaehollandiae and S. apama collected from SG, SA, Australia. Totals of 51 S. apama and 27 S. novaehollandiae individuals were examined, of which all except for four S. apama were infected by at least one dicyemid species. Dicyemid parasites were also observed in host individuals that were held in tanks for 2–3 months prior to examination, including nematogen-exclusive infections, leading to questions about persistence of dicyemids after host death and the mechanism responsible for the switch between a nematogen phase and a rhombogen phase. Variations in host size, calotte shape and collection locality are explored as predictors of differences in observed composition of the parasite fauna. In particular, dicyemid parasite fauna varied with host collection locality. As these parasites are highly host-species specific, their use as biological tags to assess cephalopod population structure using a combined morphological and molecular approach is discussed. This study increases the number of dicyemid species described from Australian cephalopods from five to ten, and from 117 to 122 species described worldwide.     

Mechanisms of Population Structuring in Giant Australian Cuttlefish Sepia apama

While a suite of approaches have been developed to describe the scale, rate and spatial structure of exchange among populations, a lack of mechanistic understanding will invariably compromise predictions of population-level responses to ecosystem modification. In this study, we measured the energetics and sustained swimming capacity of giant Australian cuttlefish Sepia apama and combined these data with information on the life-history strategy, behaviour and circulation patterns experienced by the species to predict scales of connectivity throughout parts of their range. The swimming capacity of adult and juvenile S. apama was poor compared to most other cephalopods, with most individuals incapable of maintaining swimming above 15 cm s⁻¹. Our estimate of optimal swimming speed (6–7 cm s⁻¹) and dispersal potential were consistent with the observed fine-scale population structure of the species. By comparing observed and predicted population connectivity, we identified several mechanisms that are likely to have driven fine-scale population structure in this species, which will assist in the interpretation of future population declines.     

Osteohistological insight into the early stages of growth in Mussaurus patagonicus (Dinosauria,Sauropodomorpha)

Here, we describe the bone histology of juvenile specimens of the basal sauropodomorph Mussaurus patagonicus and interpret its significance in terms of the early growth dynamics of this taxon. Thin sections from three juvenile specimens (femur length, 111–120 mm) of Mussaurus were analysed. The sampled bones consist of multiple postcranial elements collected from the Late Triassic Laguna Colorada Formation (El Tranquilo Group, Patagonia). The cortical bone is composed of fibrolamellar bone tissue. Vascularisation is commonly laminar or plexiform in the long bones. Growth marks are absent in all the examined samples. The ‘epiphyses’ of long bones are all formed by well-developed hypertrophied calcified cartilage. The predominance of woven-fibred bone matrix in cortical bones indicates a fast growth rate in the individuals examined. Moreover, given the existence of growth marks in adult specimens of Mussaurus, as in other sauropodomorphs, and assuming that the first lines of arrested growth was formed during the first year of life, the absence of growth marks in all the bones suggest that the specimens died before reaching their first year of life. Compared with the African taxon Massospondylus carinatus (another basal sauropodomorph for which the bone histology has been previously studied), it appears that Mussaurus had a higher early growth rate than Massospondylus.     

The secret life of the giant Australian cuttlefish Sepia apama (Cephalopoda): Behaviour and energetics in nature revealed through radio acoustic positioning and telemetry (RAPT)

Sepia apama were tagged with acoustic transmitters and monitored on their native House Reef, Boston Bay, South Australia, with a radio acoustic positioning telemetry (RAPT) system. Cuttlefish were tagged with position-only and intra-mantle jet pressure transmitters. New data analyses were developed to handle problem data that arise with an uneven reef environment. Maximum range for the cuttlefish varied from 90 m to 550 m. Cuttlefish home range was between 5300 m² and 23,700 m². S. apama were found to be diurnal as average distance travelled was higher in the day than at night, and cuttlefish were active for 32 days, but only 18 nights. After the cuttlefish settled into reef crevices, activity spectrum and positioning analysis showed foraging behaviour at only 3.7% per day and 2.1% per night. Cuttlefish were found to spend more than 95% of the day resting, which suggests that their bioenergetics are more akin to those of octopus than of squid. The cuttlefish combination of predator avoidance, efficient foraging and quiescent lifestyle allows energy to be channelled into growth and fulfillment of the live-fast-die-young cephalopod philosophy.     

A biological sampling problem illustrated by the population structure and growth patterns of Sardinella aurita at Tripoli,Libya

Synopsis Sardinella aurita were sampled from catches of the lampara fishery at Tripoli, Libya during October 1979 through to September 1980. Landings consisted mainly of adult fish with large (mode 21–26 cm total length), relatively fast growing 3–5-group individuals in winter and spring and smaller (mode 14–20 cm total length), slow growing 2- and 3-group fish in summer. Gonad growth commenced in April, when the fish were feeding on zooplankton, and continued at the expense of mesenteric fat until July. It is suggested that the older and bigger fish spawn first, probably after having moved westwards, and that this results in spatial and temporal differences in length at age within the population. There was no consistent difference in growth rate between the sexes, but because first maturity in males tends to occur earlier than in females, there were length distribution differences between the sexes in the youngest (2+) age group as sampled.     

Life-History Traits of the Miocene Hipparion concudense (Spain) Inferred from Bone Histological Structure

Histological analyses of fossil bones have provided clues on the growth patterns and life history traits of several extinct vertebrates that would be unavailable for classical morphological studies. We analyzed the bone histology of Hipparion to infer features of its life history traits and growth pattern. Microscope analysis of thin sections of a large sample of humeri, femora, tibiae and metapodials of Hipparion concudense from the upper Miocene site of Los Valles de Fuentidueña (Segovia, Spain) has shown that the number of growth marks is similar among the different limb bones, suggesting that equivalent skeletochronological inferences for this Hipparion population might be achieved by means of any of the elements studied. Considering their abundance, we conducted a skeletechronological study based on the large sample of third metapodials from Los Valles de Fuentidueña together with another large sample from the Upper Miocene locality of Concud (Teruel, Spain). The data obtained enabled us to distinguish four age groups in both samples and to determine that Hipparion concudense tended to reach skeletal maturity during its third year of life. Integration of bone microstructure and skeletochronological data allowed us to identify ontogenetic changes in bone structure and growth rate and to distinguish three histologic ontogenetic stages corresponding to immature, subadult and adult individuals. Data on secondary osteon density revealed an increase in bone remodeling throughout the ontogenetic stages and a lesser degree thereof in the Concud population, which indicates different biomechanical stresses in the two populations, likely due to environmental differences. Several individuals showed atypical growth patterns in the Concud sample, which may also reflect environmental differences between the two localities. Finally, classification of the specimens’ age within groups enabled us to characterize the age structure of both samples, which is typical of attritional assemblages.     

Size-selective mortality of Sea of Okhotsk pink salmon in the ocean in the winter and spring

The mortality of Sea of Okhotsk pink salmon in the winter and spring varies significantly from year to year, which complicates forecasts of its arrival in the following year based on data on the downstream migration of fry and surveys in the fall. The size-selective mortality of pink salmon was studied and the possibility of using the size and weight parameters of juveniles for predicting their return was evaluated through measurements of scale increments in juvenile pink salmon that were caught in the southern Sea of Okhotsk in the fall of 2007 and 2008 and in fish of these year classes that came back to spawn. In the 2007 year class, which had a low overwinter survival level in the ocean, the average scale increments for the first year of life were considerably smaller than those in adult fish that returned to the spawning grounds. In the pink salmon of 2008, which had a very high level of overwinter survival, the values of scale increments in juveniles and adults were similar. This confirms the hypothesis of a critical size and a critical period, according to which slowly growing juveniles that do not accumulate enough energy reserves for summer are eliminated in the winter to a greater extent as compared to fast-growing fish. Correlation analysis revealed a significant negative relationship between the size and weight of juvenile pink salmon and their mortality in the ocean. After conducting further and more extensive studies this will allow using the size parameters of juvenile pink salmon as one of predictors of its return for the year following the fall surveys. These results emphasize how important it is to take the size and growth rate of juvenile salmon into account when forecasting their return.     

Growth and survival strategy of the Antarctic mite Alaskozetes antarcticus

Growth and mortality rates of three juvenile instars and adults of the oribatid mite Alaskozetes antarcticus were measured over a twelve month period using field enclosures m Its maritime Antarctic habitat A pattern was found of rapid growth during the short summer period (0 2-0 4 μg dry weight increment per day) followed by gradual weight loss overwinter Most individuals completed one instar per year, reaching a premoult resting phase by late summer, with the moult being delayed and relatively synchronised early the next summer A small proportion of proto- and deutonymphs completed two moults within one summer period After the final moult adults overwintered again before commencing oviposition, suggesting that the life cycle (egg-egg) will take at least five years Mortality was not biased towards any instar or time of year Highest individual growth rates were obtained at a constant temperature of 7°C in the laboratory However expenmentai increase of mean field temperature from 2 7 to 4 3°C by use of a plastic cloche led to decreased growth, although no change in moulting frequency or mortality rate Laboratory survival was greatest at 2°C and decreased with increasing temperature
Alaskozetes antarcticus may be descnbed as 'a' (adversity) selected, showing an exceptionally long life cycle of five to six years and extensive physiological investment in survival adaptations Moult synchronisation in early summer, involving entering winter in the inactive premoult phase, will increase overwinter survival by reducing the chance of inoculative freezing from gut contents, and may be advantageous in maximising the time available for feeding during the subsequent summer, or ensuring proximity of the sexes after the final moult     

Postcranial morphology and growth of the pachypleurosaur Anarosaurus heterodontus (Sauropterygia) from the Lower Muschelkalk of Winterswijk, The Netherlands

Sauropterygia from the Muschelkalk are only found in lag deposits known as bone beds, and most of the material consists of isolated bones. Alpha taxonomy of Sauropterygia from the Germanic Basin which include Pachypleurosauria is thus based mainly on skull morphology of a few specimens. Articulated or associated postcranial material of pachypleurosaurs, associated with diagnostic skull material, is very rare in the Germanic Basin and currently occurs in larger numbers only in the Lower Muschelkalk of Winterswijk (Gelderland Province, The Netherlands), which continuously produces new material. For the first time, the morphology of several partially articulated skeletons of the pachypleurosaur Anarosaurus heterodontus is described and compared. Some of those specimens have skull material attached; others were identified as pachypleurosaurs on the basis of their long bone histology. The current study revealed that postcranial bones of A.?heterodontus feature a diverse morphology reflecting differences during ontogeny. Thus, A.?heterodontus specimens could be assigned to size classes (I?CIII). However, on the basis of morphology, histology, and maximal known size of isolated skulls and humeri, none of these specimens represent fully grown individuals. Growth mark counts of midshaft-femur samples, morphologically assigned to size class?III, document that this size class was reached within the first year of life. Size class?III continued into the second year of life, and then afterwards skeletal maturity was reached. Thus, a juvenile A.?heterodontus grew very fast, which is also indicated by its bone tissue type, composed of a high number of radial vascular canals and a fast-deposited bone matrix. The assignment of isolated bones from Lower to Middle Muschelkalk localities to A.?heterodontus is now possible with an extensive amended diagnosis of this taxon. This largely contributes to the understanding of taxonomical diversity and distribution. Morphological comparison of the postcranial skeleton of A.?heterodontus with that of the two other valid pachypleurosaurs from the Germanic Basin, Anarosaurus pumilio and Dactylosaurus, supports their close phylogenetic relationship. Furthermore, the skeleton of A.?heterodontus has no morphological or histological aquatic adaptation such as pachyostosis or pachyosteosclerosis and thus represents the least degree of aquatic adaptation within Pachypleurosauria.     

THE GROWTH AND BREEDING CYCLE OF LITTORINA OBTUSATA (GASTROPODA : PROSOBRANCHIATA) FROM CARDIGAN BAY

The population structure of Littorina obtusata was for mostof the year bimodal, with a stable adult component and an activelygrowing juvenile component. Growth rate of the juvenile componentwas similar at different beach levels and there was a retardationof growth as the snails approached sexual maturity. There was little variation in the histology and external appearanceof the reproductive organs throughout the year and the breedingcycle has been characterized by the amount of spawn laid andthe presence of mature eggs in the oviduct of the female. ^*Present address: Portway School, Penpole Lane, Shirehampton,Bristol. (Received 7 September 1977;     

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.     

Bone histology as an approach to providing data on certain key life history traits in mammals: Implications for conservation biology

The knowledge of the life histories of wild mammals is of crucial importance in the field of conservation management. The endangered status of many species calls for faster data collection that can be used in risk assessment and, ultimately, for designing conservation policies. This study is pioneering the potential of bone histology to provide data on life history traits crucial for conservation biology in long-lived mammals. Long bone cross-sections show pronounced annual cycles of growth arrest allowing application of skeletochronology (counts of lines of arrested growth ‘LAGs’). Consequently, the number of LAGs within the primary fast-growing bone tissue up to the outer cortical slow-growing bone tissue corresponds to the age at first reproduction; whereas the age at death can be estimated by the total number of rest lines throughout the whole of bone cross-section. Furthermore, the diameters of successive growth rings as well as the osteocyte lacuna density may shed light on growth rates. We use the endangered desert dwelling antelope Addax nasomaculatus as a case study. By analyzing different ontogenetic stages in five Addax individuals (three captive and two wild specimens) from a museum collection, we show that bone histology may be a reliable tool for determining certain key life history traits. In our sample, the wild Addax female attained reproductive maturity at three years, whereas the male specimens, both the captive and the wild ones, reached maturity at four years. This is congruent with data from other large antelopes with male-biased size dimorphism, but differs slightly from data on sexual maturity previously published for wild Addax. Moreover, quantification of osteocyte lacunae in both adult males provides a higher cell density in the captive one than in the wild one suggesting the strong effect of constant resources supply in individuals from zoos on growth rates. While age at first reproduction and longevity are essentials parameters to carry out demographic models, growth rates may allow evaluation of the health status of wild populations. This approach may provide useful data on life history traits when applied to bones collected in the wild.     

Reproduction of the sandpaper skate,Bathyraja kincaidii (Garman 1908) in the eastern North Pacific

The sandpaper skate, Bathyraja kincaidii, is commonly taken as bycatch in trawl fisheries off central California. The impact of fisheries exploitation on this population is unknown, however little is known about its biology, including its reproductive life history. To determine reproductive aspects of B. kincaidii, 506 individuals were collected from commercial trawls, museum collections, and monthly trawl and longline surveys conducted by the National Marine Fisheries Service. Males ranged from 112 to 635 mm TL, whereas females ranged from 113 to 610 mm TL. Males reach first maturity at 440 mm TL and at ~3 years, whereas females reach first maturity at 450 mm TL, approximately 1 year later. Size and estimated age at 50 % maturity were similar between sexes: 492 mm TL and 7.5 years for males and 467 mm TL and 7.1 years for females. Gonadosomatic index (GSI) and hepatosomatic index (HSI) values indicate a continuous reproductive cycle, with a seasonal peak in the summer and fall, and a resting period following egg case deposition.     

Interpopulation variation in growth and life-history traits of the introduced sunfish, pumpkinseed Lepomis gibbosus, in southern England

We examined attributes of growth and reproduction in 19 populations of pumpkinseed (Lepomis gibbosus) introduced into southern England in order to: (i) assess variability of these traits in a northern European climate; (ii) assess inter‐relationships among these variables; and (iii) compare these attributes with populations from other parts of Europe where pumpkinseeds have been introduced. Growth rates varied considerably among populations, but juvenile growth rates and adult body sizes were generally among the lowest in Europe. Mean age at maturity ranged from 2.0 to 3.9, and was strongly predicted by the juvenile growth rate (earlier maturity with faster juvenile growth). Other population parameters that also displayed significant negative associations with mean age at maturity were gonadosomatic index, body condition, and adult body size (total length, TL at age 5). Mean TL at maturity and the adult growth increment showed no significant associations with any of the other growth or life‐history variables. Pumpkinseed populations in England matured significantly later than those introduced into warmer, more southerly areas of the continental Europe. All of these data suggest that a combination of cool summer temperatures and resource limitation is the cause of slow growth, small adult body size and delayed maturity relative to introduced populations on the European mainland.     

Age structure and body size of the Chuanxi Tree Frog Hyla annectans chuanxiensis from two different elevations in Sichuan (China)

Age, body size, and growth patterns in the subtropical anuran Hyla annectans chuanxiensis from high (Dengchigou Protection Station) and low (Lingguan Town) elevations in Baoxing County of Sichuan province (China) were described using skeletochronology. Females were significantly older than males at the low-elevation site, but there was no significant difference between the sexes at the high-elevation site. Age at sexual maturity of both males and females was 2 years at the high-elevation site, whereas males matured at 1 year and females at 2 years at the low-elevation site. Males and females from the low-elevation population reached a maximum age of 3 and 4 years, respectively, whereas males and females from the high-elevation population reached a maximum age of 4 and 5 years, respectively. At both sites, females were significantly larger than males. Females and males from the high-elevation population were larger than individuals from the low-elevation population. When the effect of age was controlled, the differences in body size of the two populations were significant only for females. Von Bertalanffy growth curves indicated that the growth rates in males was greater than in females in both populations. They also showed that the growth of both sexes slowed at an earlier age in the low-elevation population than in the high-elevation population. The findings suggest that age is a major factor underlying body size patterns for both sexes, but that the elevation of the locality affects the body size of females.     

A fish‐eye view of cuttlefish camouflage using in situ spectrometry

Cuttlefish are colour blind yet they appear to produce colour‐coordinated patterns for camouflage. Under natural in situ lighting conditions in southern Australia, we took point‐by‐point spectrometry measurements of camouflaged cuttlefish, Sepia apama, and various natural objects in the immediate visual surrounds to quantify the degree of chromatic resemblance between cuttlefish and backgrounds to potential fish predators. Luminance contrast was also calculated to determine the effectiveness of cuttlefish camouflage to this information channel both for animals with or without colour vision. Uniform body patterns on a homogeneous background of algae showed close resemblance in colour and luminance; a Uniform pattern on a partially heterogeneous background showed mixed levels of resemblance to certain background features. A Mottle pattern with some disruptive components on a heterogeneous background showed general background resemblance to some benthic objects nearest the cuttlefish. A noteworthy observation for a Disruptive body pattern on a heterogeneous background was the wide range in spectral contrasts compared to Uniform and Mottle patterns. This suggests a shift in camouflage tactic from background resemblance (which hinders detection by the predator) to more specific object resemblance and disruptive camouflage (which retards recognition). © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 535–551.     

Experimental and demographic analyses of growth rate and sexual size dimorphism in a lizard, Sceloporus undulatus

Sexual size dimorphism (SSD) is a common phenomenon caused by a variety of environmental and genetic mechanisms in animals. In the current study, we investigate the demography of a population of eastern fence lizards ( Sceloporus undulatus ) to compare age structure and survivorship between the sexes, and we examine growth rates of juveniles under both natural and controlled laboratory conditions to elucidate causes of SSD in this species. Furthermore, using our laboratory growth data, we examine the heritability of juvenile growth rates. Our results show that SSD develops in the field before the end of the first year of age (before sexual maturity) because juvenile females grow more rapidly than juvenile males. In the laboratory environment, however, we observed no sexual difference in growth rates for lizards up to the size of maturity in the field. Thus, sexual differences in growth rate and subsequent development of SSD in this population are highly plastic and subject to strong proximal control. We found high levels of additive genetic variance for juvenile growth, indicating a strong potential for selection to operate on juvenile growth rates. Our results indicate that selection on juvenile growth rate could account for differences in growth among populations but would not necessarily contribute to SSD within our population due to the high plasticity in growth rate.     

Growth and tissue composition as a function of feeding history in juvenile cephalopods

We present the results of a series of experiments that examined the effect of feeding history on the growth and tissue composition of juveniles of two tropical cephalopods; the squid Sepioteuthis lessoniana and the cuttlefish Sepia elliptica. Juveniles were reared in individual containers for between 35 and 42 days at different ration levels, three ration levels for the squid and two levels for the cuttlefish. Although differences in ration were sufficient to cause different growth rates, both in body length and mass, the effects on tissue composition were less definitive. Sepioteuthis juveniles on the highest rations had higher concentrations of water, but no difference in lipid, carbohydrate or protein when compared with their lower ration siblings. In the case of juvenile cuttlefish no difference in tissue composition was detected between the two ration levels. RNA:protein ratios were also determined for the juveniles to provide an estimate of instantaneous growth. A significant correlation was found between body size and RNA:protein ratio in the squid; those juveniles that ate more had higher RNA:protein ratios than lower ration individuals. Significantly, the juvenile cuttlefish showed no relationship between growth rate and RNA:protein ratios, which means that we are unable to use this measure to estimate the growth rates of wild individuals. In conclusion, ration level did affect growth rates and food availability is an important factor in modifying growth rates of wild individuals. However, we could not find, at the individual level, an index or measure that could be used to explain the variability of observed differences in growth rates as a function of nutritional history.     

池塘中斑索蟾的种群结构、个体生长与存活率

在澳大利亚新南威尔士南部沿海,作者搜集了一个池塘繁殖的斑索蟾(Crinia signifera)的种群统计资料。通过捕捉进出池塘的1 612只个体,获得种群大小、结构、生长率、性成熟时的大小和年龄、死亡率及寿命资料。迁移高峰从6月持续到11月,蛙的最高、最低遇见数量分别出现在春季和秋季。但第2年,该池塘蛙的数量明显减少,可能是由于补充到种群中的幼体数量很少的缘故。6个月后个体的重捕率很低;但距第一次捕获18个月以后,仍有个别个体再次被捕获。性成熟时,雌性比雄性的身体大一些。生长曲线显示,雌性比雄性的生长更快,所以更早地达到性成熟。研究种群的数量、结构和死亡率趋势等与已知的其它Crinia signifera种群基本一致。但研究种群迁移活动的高峰出现较晚,并且夏季的活动水平明显很高。这种长的活动时间可能会导致存活率的下降,同时有利于选择迅速性成熟的雌性[动物学报51 (3) : 393 -400 , 2005]。     

Age estimation,growth and maturity of the Argentine hake (Merluccius hubbsi Marini, 1933) along the northernmost limit of its distribution in the south-western Atlantic

Age, growth and length-at-maturity of the Argentine hake (Merluccius hubbsi) were studied in the northernmost limit of the species distribution in the south-western Atlantic. A total of 351 otoliths and information from 1610 specimens sampled from the industrial double-rig trawl landings between May 2013 and April 2014 were used. Age and growth were estimated by counting and measuring increments in sectioned sagittae otoliths, and length at maturity was estimated based on macroscopic gonadal analysis. For both sexes, hepatosomatic index and condition index increased mainly during spring, reaching a maximum at the end of summer before the subsequent spawning season began. Gonadosomatic index was highest in April, believed to correspond with peak spawning. The annual periodicity of alternate opaque and translucent zones was validated by marginal increment analysis. Growth curves were fitted to back-calculated size at age by fitting the three-parameters von Bertalanffy growth function. The maximum age was 5 years in fish of either sex. Females attained larger sizes than males. The parameters of the von Bertalanffy growth equations were: L∞?=?533?mm, k?=?0.231 year^?1 and t₀?=??0.935 year for females; L∞?=?394?mm, k?=?0.405 year^?1 and t₀?=??0.463 year for males. The mean length and age at first maturity was 273?mm at 1.9 years for males and 274?mm at 2.0 years for females.