Age validation, growth modeling, and mortality estimates for striped trumpeter (Latris lineata) from southeastern Australia: making the most of patchy data

Age estimates for striped trumpeter (Latris lineata) from Tasmanian waters were produced by counting annuli on the transverse section of sagittal otoliths and were validated by comparison of growth with known-age individuals and modal progression of a strong recruitment pulse. Estimated ages ranged from one to 43 years; fast growth rates were observed for the first five years. Minimal sexual dimorphism was shown to exist between length, weight, and growth characteristics of striped trumpeter. Seasonal growth variability was strong in individuals up to at least age four, and growth rates peaked approximately one month after the observed peak in sea surface temperature. A modified two-phase von Bertalanffy growth function was fitted to the length-at-age data, and the transition between growth phases was linked to apparent changes in physiological and life history traits, including offshore movement as fish approach maturity. The two-phase curve was found to represent the mean length at age in the data better than the standard von Bertalanffy growth function. Total mortality was estimated by using catch curve analysis based on the standard and two-phase von Bertalanffy growth functions, and estimates of natural mortality were calculated by using two empirical models, one based on longevity and the other based on the parameters L∞ and k from both growth functions. The interactions between an inshore gillnet fishery targeting predominately juveniles and an offshore hook fishery targeting predominately adults highlight the need to use a precautionary approach when developing harvest strategies.

Age, growth and mortality of the toadfish, Halobatrachus didactylus (Schneider, 1801) (Pisces: Batrachoididae), in the Bay of Cádiz (southwestern Spain)

Age, growth and mortality of the toadfish, Halobatrachus didactylus, were determined by examination of the whole sagittal otoliths of fish sampled in the Bay of Cádiz (southwestern Spain) from March 1999 to March 2000. A total of 844 specimens (425 males, 416 females, and 3 of indeterminate sex), ranging from 95 to 470 mm in total length were examined. Eighty-nine percent of the otoliths could be read allowing an age estimation. The opaque zone was formed between April and May coincident with the maximum reproductive peak, while the translucent zone formed mainly in summer-fall (June to December). Maximum ages for males and females were 12 and 10 years, respectively. The samples were dominated by 2- to 6-year-old specimens. Males matured at an age of approximately 2 years and females at 3 years. Fish total length and otolith radius were closely related. The von Bertalanffy growth curve was used to describe growth. The parameters were derived from back-calculated length-at-age. Significant differences in the growth parameters were found between sexes. Although the growth analysis revealed that this species is slow-growing, males reached larger sizes than females. Females appeared to experience higher natural mortality rates than males.

Use of skeletochronological analysis to estimate the age of leatherback sea turtles Dermochelys coriacea in the western North Atlantic

Although growth rate and age data are essential for leatherback management, estimates of these demographic parameters remain speculative due to the cryptic life history of this endangered species. Skeletochronological analysis of scleral ossicles obtained from 8 captive, known-age and 33 wild leatherbacks originating from the western North Atlantic was conducted to characterize the ossicles and the growth marks within them. Ages were accurately estimated for the known-age turtles, and their growth mark attributes were used to calibrate growth mark counts for the ossicles from wild specimens. Due to growth mark compaction and resorption, the number of marks visible at ossicle section tips was consistently and significantly greater than the number visible along the lateral edges, demonstrating that growth mark counts should be performed at the tips so that age is not underestimated. A correction factor protocol that incorporated the trajectory of early growth increments was used to estimate the number of missing marks in those ossicles exhibiting resorption, which was then added to the number of observed marks to obtain an age estimate for each turtle. A generalized smoothing spline model, von Bertalanffy growth curve, and size-at-age function were used to obtain estimates of age at maturity for leatherbacks in the western North Atlantic. Results of these analyses suggest that median age at maturation for leatherbacks in this part of the world may range from 24.5 to 29 yr. These age estimates are much greater than those proposed in previous studies and have significant implications for population management and recovery.

Age determination and growth of the night shark (Carcharhinus signatus) off the northeastern Brazilian coast

Age and growth of the night shark (Carcharhinus signatus) from areas off northeastern Brazil were determined from 317 unstained vertebral sections of 182 males (113–215 cm total length [TL]), 132 females (111.5–234.9 cm) and three individuals of unknown sex (169–242 cm). Although marginal increment (MI) analysis suggests that band formation occurs in the third and fourth trimesters in juveniles, it was inconclusive for adults. Thus, it was assumed that one band is formed annually. Births that occur over a protracted period may be the most important source of bias in MI analysis. An estimated average percent error of 2.4% was found in readings for individuals between two and seventeen years. The von Bertalanffy growth function (VBGF) showed no significant differences between sexes, and the model derived from back-calculated mean length at age best represented growth for the species (L∞=270 cm, K=0.11/yr, t0=–2.71 yr) when compared to the observed mean lengths at age and the Fabens’ method. Length-frequency analysis on 1055 specimens (93–260 cm) was used to verify age determination. Back-calculated size at birth was 66.8 cm and maturity was reached at 180–190 cm (age 8) for males and 200–205 cm (age ten) for females. Age composition, estimated from an age-length key, indicated that juveniles predominate in commercial catches, representing 74.3% of the catch. A growth rate of 25.4 cm/yr was estimated from birth to the first band (i.e. juveniles grow 38% of their birth length during the first year), and a growth rate of 8.55 cm/yr was estimated for eight- to ten-year-old adults.

Effects of density-dependence and sea surface temperature on interannual variation in length-at-age of chub mackerel (Scomber japonicus) in the Kuroshio-Oyashio area during 1970–1997

Annual mean fork length (FL) of the Pacific stock of chub mackerel (Scomber japonicus) was examined for the period of 1970–97. Fork length at age 0 (6 months old) was negatively correlated with year-class strength which fluctuated between 0.2 and 14 billion in number for age-0 fish. Total stock biomass was correlated with FL at age but was not a significant factor. Sea surface temperature (SST) between 38–40°N and 141–143°E during April–June was also negatively correlated with FL at age 0. A modified von Bertalanffy growth model that incorporated the effects of population density and SST on growth was well fitted to the observed FL at ages. The relative FL at age 0 for any given year class was maintained throughout the life span. The variability in size at age in the Pacific stock of chub mackerel is largely attributable to growth during the first six months after hatching.

Age and growth of sailfish (Istiophorus platypterus) in waters off eastern Taiwan

Age and growth of sailfish (Istiophorus platypterus) in waters off eastern Taiwan were examined from counts of growth rings on cross sections of the fourth spine of the first dorsal fin. Length and weight data and the dorsal fin spines were collected monthly at the fishing port of Shinkang (southeast of Taiwan) from July 1998 to August 1999. In total, 1166 dorsal fins were collected, of which 1135 (97%) (699 males and 436 females) were aged successfully. Trends in the monthly mean marginal increment ratio indicated that growth rings are formed once a year. Two methods were used to back-calculate the length of presumed ages, and growth was described by using the standard von Bertalanffy growth function and the Richards function. The most reasonable and conservative description of growth assumes that length-at-age follows the Richards function and that the relationship between spine radius and lower jaw fork length (LJFL) follows a power function. Growth differed significantly between the sexes; females grew faster and reached larger sizes than did males. The maximum sizes in our sample were 232 cm LJFL for female and 221 cm LJFL for male.

Red snapper (Lutjanus campechanus) demographic structure in the northern Gulf of Mexico based on spatial patterns in growth rates and morphometrics

Red snapper (Lutjanus campechanus) in the United States waters of the Gulf of Mexico (GOM) has been considered a single unit stock since management of the species began in 1991. The validity of this assumption is essential to management decisions because measures of growth can differ for nonmixing populations. We examined growth rates, size-at-age, and length and weight information of red snapper collected from the recreational harvests of Alabama (n=2010), Louisiana (n=1905), and Texas (n =1277) from 1999 to 2001. Ages were obtained from 5035 otolith sections and ranged from one to 45 years. Fork length, total weight, and age-frequency distributions differed significantly among all states; Texas, however, had a much higher proportion of smaller, younger fish. All red snapper showed rapid growth until about age 10 years, after which growth slowed considerably. Von Bertalanffy growth models of both mean fork length and mean total weight-at-age predicted significantly smaller fish at age from Texas, whereas no differences were found between Alabama and Louisiana models. Texas red snapper were also shown to differ significantly from both Alabama and Louisiana red snapper in regressions of mean weight at age. Demographic variation in growth rates may indicate the existence of separate management units of red snapper in the GOM. Our data indicate that the red snapper inhabiting the waters off Texas are reaching smaller maximum sizes at a faster rate and have a consistently smaller total weight at age than those collected from Louisiana and Alabama waters. Whether these differences are environmentally induced or are the result of genetic divergence remains to be determined, but they should be considered for future management regulations.

Radiometric validation of age, growth, and longevity for the blackgill rockfish (Sebastes melanostomus)

As nearshore fish populations decline, many commercial fishermen have shifted fishing effort to deeper continental slope habitats to target fishes for which biological information is limited. One such fishery that developed in the northeastern Pacific Ocean in the early 1980s was for the blackgill rockfish (Sebastes melanostomus), a deep-dwelling (300−800 m) species that congregates over rocky pinnacles, mainly from southern California to southern Oregon. Growth zone-derived age estimates from otolith thin sections were compared to ages obtained from the radioactive disequilibria of 210Pb, in relation to its parent, 226Ra, in otolith cores of blackgill rockfish. Age estimates were validated up to 41 years, and a strong pattern of agreement supported a longevity exceeding 90 years. Age and length data fitted to the von Bertalanffy growth function indicated that blackgill rockfish are slow-growing (k= 0.040 females, 0.068 males) and that females grow slower than males, but reach a greater length. Age at 50% maturity, derived from previously published length-at-maturity estimates, was 17 years for males and 21 years for females. The results of this study agree with general life history traits already recognized for many Sebastes species, such as long life, slow growth, and late age at maturation. These traits may undermine the sustainability of blackgill rockfish populations when heavy fishing pressure, such as that which occurred in the 1980s, is applied.

Estimating long-term growth-rate changes of southern bluefin tuna (Thunnus maccoyii) from two periods of tag-return data

Southern bluefin tuna (SBT) (Thunnus maccoyii) growth rates are estimated from tag-return data associated with two time periods, the 1960s and 1980s. The traditional von Bertalanffy growth model (VBG) and a two-phase VBG model were fitted to the data by maximum likelihood. The traditional VBG model did not provide an adequate representation of growth in SBT, and the two-phase VBG yielded a significantly better fit. The results indicated that significant change occurs in the pattern of growth in relation to a VBG curve during the juvenile stages of the SBT life cycle, which may be related to the transition from a tightly schooling fish that spends substantial time in near and surface shore waters to one that is found primarily in more offshore and deeper waters. The results suggest that more complex growth models should be considered for other tunas and for other species that show a marked change in habitat use with age. The likelihood surface for the two-phase VBG model was found to be bimodal and some implications of this are investigated. Significant and substantial differences were found in the growth for fish spawned in the 1960s and in the 1980s, such that after age four there is a difference of about one year in the expected age of a fish of similar length which persists over the size range for which meaningful recapture data are available. This difference may be a density-dependent response as a consequence of the marked reduction in the SBT population. Given the key role that estimates of growth have in most stock assessments, the results indicate that there is a need both for the regular monitoring of growth rates and for provisions for changes in growth over time (possibly related to changes in abundance) in the stock assessment models used for SBT and other species.

Life history of the Atlantic sharpnose shark (Rhizoprionodon terraenovae) (Richardson, 1836) off the southeastern United States

The life history of the Atlantic sharpnose shark (Rhizoprionodon terraenovae) was described from 1093 specimens collected from Virginia to northern Florida between April 1997 and March 1999. Longitudinally sectioned vertebral centra were used to age each specimen, and the periodicity of circuli deposition was verified through marginal increment analysis and focus-to-increment frequency distributions. Rhizoprionodon terraenovae reached a maximum size of 828 mm precaudal length (PCL) and a maximum age of 11+ years. Mean back-calculated lengths-at-age ranged from 445 mm PCL at age one to 785 mm PCL at age ten for females, and 448 mm PCL at age one to 747 mm PCL at age nine for males. Observed lengthat-age data (estimated to 0.1 year) yielded the following von Bertalanffy parameters estimates: L∞= 749 mm PCL (SE=4.60), K = 0.49 (SE=0.020), and t0= –0.94 (SE=0.046) for females; and L∞= 745 mm PCL (SE = 5.93), K = 0.50 (SE=0.024), and t0= –0.91 (SE = 0.052) for males. Sexual maturity was reached at age three and 611 mm PCL for females, and age three and 615 mm PCL for males. Rhizoprionodon terraenovae reproduced annually and had a gestation period of approximately 11 months. Litter size ranged from one to eight (mean=3.85) embyros, and increased with female PCL.

Age validation, growth, mortality, and additional population parameters of the goldband snapper (Pristipomoides multidens) off the Kimberley coast of northwestern Australia

Goldband snapper (Pristipomoides multidens) collected from commercial trap and line fishermen off the Kimberley coast of northwestern Australia were aged by examination of sectioned otoliths (sagittae).A total of 3833 P. multidens, 80–701 mm fork length (98–805 mm total length), were examined from commercial catches from 1995 to 1999. The oldest fish was estimated to be age 30+ years. Validation of age estimates was achieved with marginal increment analysis. The opaque and translucent zones were each formed once per year and are considered valid annual growth increments (the translucent zone was formed once per year between January and May). A strong link between water temperature and translucent zone formation was evident in P. multidens. The von Bertalanffy growth function was used to describe growth from length-at-age data derived from sectioned otoliths.

Life history and population dynamics of the finetooth shark (Carcharhinus isodon) in the northeastern Gulf of Mexico

The life history and population dynamics of the finetooth shark (Carcharhinus isodon) in the north-eastern Gulf of Mexico were studied by determining age, growth, size-at-maturity, natural mortality, productivity, and elasticity of vital rates of the population. The von Bertalanffy growth model was estimated as Lt=1559 mm TL (1–e–0.24 (t+2.07)) for females and Lt = 1337 mm TL (1–e–0.41 (t+1.39)) for males. For comparison, the Fabens growth equation was also fitted separately to observed size-at-age data, and the fits to the data were found to be similar. The oldest aged specimens were 8.0 and 8.1 yr, and theoretical longevity estimates were 14.4 and 8.5 yr for females and males, respectively. Median length at maturity was 1187 and 1230 mm TL, equivalent to 3.9 and 4.3 yr for males and females, respectively. Two scenarios, based on the results of the two equations used to describe growth, were considered for population modeling and the results were similar. Annual rates of survivorship estimated through five methods ranged from 0.850/yr to 0.607/yr for scenario 1 and from 0.840/yr to 0.590/yr for scenario 2. Productivities were 0.041/yr for scenario 1 and 0.038/yr for scenario 2 when the population level that produces maximum sustain-able yield is assumed to occur at an instantaneous total mortality rate (Z) equaling 1.5 M, and were 0.071/yr and 0.067/yr, when Z=2 M for scenario 1 and 2, respectively. Mean generation time was 6.96 yr and 6.34 yr for scenarios 1 and 2, respectively. Elasticities calculated through simulation of Leslie matrices averaged 12.6% (12.1% for scenario 2) for fertility, 47.7% (46.2% for scenario 2) for juvenile survival, and 39.7% (41.6% for scenario 2) for adult survival. In all, the finetooth shark exhibits life-history and population characteristics intermediate to those of sharks in the small coastal complex and those from some large coastal species, such as the blacktip shark (Carcharhinus limbatus).

Age and growth of the estuarine dolphin (Sotalia guianensis) (Cetacea, Delphinidae) on the Paraná Coast, southern Brazil

Teeth of 71 estuarine dolphins (Sotalia guianensis) incidentally caught on the coast of Paraná State, southern Brazil, were used to estimate age. The oldest male and female dolphins were 29 and 30 years, respectively. The mean distance from the neonatal line to the end of the first growth layer group (GLG) was 622.4 ±19.1 μm (n=48). One or two accessory layers were observed between the neonatal line and the end of the first GLG. One of the accessory layers, which was not always present, was located at a mean of 248.9 ±32.6 μm (n=25) from the neonatal line, and its interpretation remains uncertain.The other layer, located at a mean of 419.6 ±44.6 μm (n=54) from the neonatal line, was always present and was first observed between 6.7 and 10.3 months of age. This accessory layer could be a record of weaning in this dolphin. Although no differences in age estimates were observed between teeth sectioned in the anterior-posterior and buccal-lingual planes, we recommend sectioning the teeth in the buccal-lingual plane in order to obtain on-center sections more easily. We also recommend not using teeth from the most anterior part of the mandibles for age estimation. The number of GLGs counted in those teeth was 50% less than the number of GLGs counted in the teeth from the median part of the mandible of the same animal. Although no significant difference (P>0.05) was found between the total lengths of adult male and female estuarine dolphins, we observed that males exhibited a second growth spurt around five years of age. This growth spurt would require that separate growth curves be calculated for the sexes. The asymptotic length (TL∞), k, and t0 obtained by the von Bertalanffy growth model were 177.3 cm, 0.66, and –1.23, respectively, for females and 159.6 cm, 2.02, and –0.38, respectively, for males up to five years, and 186.4 cm, 0.53 and –1.40, respectively, for males older than five years. The total weight (TW)/total length (TL) equations obtained for male and female estuarine dolphins were TW = 3.156 × 10−6 × TL 3.2836 (r=0.96), and TW = 8.974 × 10−5 × TL 2.6182 (r=0.95), respectively.

Age and growth estimates of the winter skate (Leucoraja ocellata) in the western Gulf of Maine

Age and growth estimates for the winter skate (Leucoraja ocellata) were estimated from vertebral band counts on 209 fish ranging in size from 145 to 940 mm total length (TL). An index of average percent error (IAPE) of 5.8% suggests that our aging method represents a precise approach to the age assessment of L. ocellata. Marginal increments were significantly different between months (Kruskal-Wallis P<0.001) and a distinct trend of increasing monthly increment growth began in July. Estimates of von Bertalanffy growth parameters suggest that females attain a slightly larger asymptotic TL (L∞=1374 mm) than males (L∞=1218 mm) and grow more slowly (k=0.059 and 0.074, respectively). The oldest ages obtained for the winter skate were 19 years for males and 18 years for females, which corresponded to total lengths of 932 mm and 940 mm, respectively. The results indicate that the winter skate exhibits the characteristics that have made other elasmobranch populations highly susceptible to exploitation by commercial fisheries.

Modeling red sea urchin (Strongylocentrotus franciscanus) growth using six growth functions

The growth of red sea urchins (Strongylocentrotus franciscanus) was modeled by using tag-recapture data from northern California. Red sea urchins (n=211) ranging in test diameter from 7 to 131 mm were examined for changes in size over one year. We used the function Jt+1 = Jt + f(Jt) to model growth, in which Jt is the jaw size (mm) at tagging, and Jt+1 is the jaw size one year later. The function f(Jt), represents one of six deterministic models: logistic dose response, Gaussian, Tanaka, Ricker, Richards, and von Bertalanffy with 3, 3, 3, 2, 3, and 2 minimization parameters, respectively. We found that three measures of goodness of fi t ranked the models similarly, in the order given. The results from these six models indicate that red sea urchins are slow growing animals (mean of 7.2 ±1.3 years to enter the fishery). We show that poor model selection or data from a limited range of urchin sizes (or both) produces erroneous growth parameter estimates and years-to-fishery estimates. Individual variation in growth dominated spatial variation at shallow and deep sites (F=0.246, n=199, P=0.62). We summarize the six models using a composite growth curve of jaw size, J, as a function of time, t: J = A(B – e–Ct) + Dt, in which each model is distinguished by the constants A, B, C, and D. We suggest that this composite model has the flexibility of the other six models and could be broadly applied. Given the robustness of our results regarding the number of years to enter the fishery, this information could be incorporated into future fishery management plans for red sea urchins in northern California.