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.     

Investigations of Δ 14C, δ 13C, and δ 15N in vertebrae of white shark (Carcharodon carcharias) from the eastern North Pacific Ocean

The white shark, Carcharodon carcharias, has a complex life history that is characterized by large scale movements and a highly variable diet. Estimates of age and growth for the white shark from the eastern North Pacific Ocean indicate they have a slow growth rate and a relatively high longevity. Age, growth, and longevity estimates useful for stock assessment and fishery models, however, require some form of validation. By counting vertebral growth band pairs, ages can be estimated, but because not all sharks deposit annual growth bands and many are not easily discernable, it is necessary to validate growth band periodicity with an independent method. Radiocarbon (¹⁴C) age validation uses the discrete ¹⁴C signal produced from thermonuclear testing in the 1950s and 1960s that is retained in skeletal structures as a time-specific marker. Growth band pairs in vertebrae, estimated as annual and spanning the 1930s to 1990s, were analyzed for Δ¹⁴C and stable carbon and nitrogen isotopes (δ¹³C and δ¹⁵N). The aim of this study was to evaluate the utility of ¹⁴C age validation for a wide-ranging species with a complex life history and to use stable isotope measurements in vertebrae as a means of resolving complexity introduced into the ¹⁴C chronology by ontogenetic shifts in diet and habitat. Stable isotopes provided useful trophic position information; however, validation of age estimates was confounded by what may have been some combination of the dietary source of carbon to the vertebrae, large-scale movement patterns, and steep ¹⁴C gradients with depth in the eastern North Pacific Ocean.     

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.     

Validated age and growth estimates for the shortfin mako, Isurus oxyrinchus, in the North Atlantic Ocean

Age and growth estimates for the shortfin mako, Isurus oxyrinchus, derived from vertebral centra of 258 specimens (118 males, 140 females), ranging in size from 64 to 340 cm fork length (FL) were compared with data from 22 tag–recaptured individuals (74–193 cm FL) and length–frequency data from 1822 individuals (1035 males, 787 females; 65–215 cm FL). Annual band-pair deposition, confirmed by a concurrent bomb radiocarbon validation study, was used as the basis for band interpretation. Validation was further confirmed with a tetracycline-injected male shortfin mako recaptured after being at liberty off South Africa for 1 year and aged at 18 years. Growth rates from tag–recapture analysis (GROTAG) were higher than those derived from vertebral annuli and were only available from sharks up to 193 cm FL at recapture. Modal length–frequency data were used to verify the first four age classes. Growth curves were fit using both von Bertalanffy and Gompertz models. The 3-parameter version of the von Bertalanffy growth function produced the most biologically reasonable values for males, based on observed data (L _∞  = 253 cm FL, K = 0.125 year^?1 (estimated longevity = 21 year), and L ₀ = 72 cm). The 3-parameter version of the Gompertz growth function produced the most biologically reasonable estimates, for females (L _∞  = 366 cm FL, K = 0.087 year^?1 (estimated longevity = 38 year) and L ₀ = 88 cm. Males and females were aged to 29 (260 cm FL) and 32 years (335 cm FL), respectively. Both sexes grew similarly to age 11 (207 cm FL, 212 cm FL for males and females, respectively) when the curve leveled in males and continued to rise in females. Age at 50% maturity was estimated at 8 years for males (185 cm FL) and 18 years for females (275 cm FL). The species grows slower, matures later and has a longer life span than previously reported in North Atlantic waters.     

Age and growth of the bonnethead shark, Sphyrna tiburo, from northwest Florida, with comments on clinal variation

Age and growth rates of the bonnethead shark, Sphyrna tiburo, from northwest Florida were estimated from vertebrae collected between October 1992 and October 1995. The von Bertalanffy growth equation was fit to male and female vertebral age data. Initial growth was rapid (≈ 200 mm TL) for both sexes from age 0–1. At age 2 growth slowed for males but continued for females. Similar to many species of sharks, females grew slower than males (K = 0.28 and K = 0.69, respectively) but attained a larger maximum size (L_∞=1226 and L_∞=897). Maximum age was estimated in males and females to be 8+ and 12+ years, respectively. Growth of young-of-year sharks was 21 to 30 mm TL per month determined by three different methods. A comparison of age and growth estimates from populations at more southerly latitudes suggest that clinal variation in total length may be evident among bonnethead sharks in the Gulf of Mexico with females reaching larger sizes in northern areas as compared to south Florida. This revised version was published online in July 2006 with corrections to the Cover Date.     

Age and growth of the tropical oviparous shark,Chiloscyllium punctatum in Indonesian waters

The brown-banded bamboo shark, Chiloscyllium punctatum, is the most common shark caught in coastal commercial fisheries throughout Southeast Asia, yet there is a lack of the life-history information necessary for reliable stock assessments. The authors estimated growth rates and age at maturity using analysis of growth bands in vertebral centra. They trialled four different techniques to enhance the visibility and improve identification of the putative annual growth bands necessary for age estimation. The authors found that the burn method on whole vertebral centra provided the most readable and consistent results for age analysis. The logistic model was chosen as the best-fit growth model for age estimation of 330 individual C. punctatum from Indonesia. Several age verification methods, including marginal increment ratio and length-frequency analysis, were performed with the support of age validation through the use of calcein-labelled vertebrae from two sharks maintained in captivity. This study found that C. punctatum from Indonesian waters is a fast-growing species that can grow up to 18 cm year⁻¹, reach an estimated maximum total length of 1 m, mature at c. 6.5 years and live for up to 14 years.     

Application of bomb radiocarbon chronologies to shortfin mako (Isurus oxyrinchus) age validation

Age estimation is an issue for the shortfin mako, Isurus oxyrinchus, because of disagreement on vertebral band-pair deposition periodicity. In the 1950s–1960s, thermonuclear testing released large amounts of radiocarbon into the atmosphere, which diffused into the ocean through gas exchange. This influx created a time-specific marker that can be used in age validation. Annual band-pair deposition in the porbeagle, Lamna nasus, was validated in a previous study and indicated preliminary annual deposition in the shortfin mako, using four samples from one vertebra. In the present study, age estimates from 54 shortfin mako vertebrae collected in 1950–1984 ranged 1–31 years. Ageing error between readers was consistent, with 76% of the estimates ranging within 2 years. Twenty-one Δ¹⁴C values from eight shortfin mako vertebrae (collected in the western North Atlantic in 1963–1984) ranged −154.8‰ to 86.8‰. The resulting conformity with the Δ¹⁴C timeline for the porbeagle supported annual band-pair deposition in vertebrae of the shortfin mako.     

Bomb dating and age validation using the spines of spiny dogfish (Squalus acanthias)

Bomb radiocarbon has previously been used to validate the age of large pelagic sharks based on incorporation into vertebrae. However, not all sharks produce interpretable vertebral growth bands. Here we report the first application of bomb radiocarbon as an age validation method based on date-specific incorporation into spine enamel. Our results indicate that the dorsal spines of spiny dogfish, Squalus acanthias, recorded and preserved a bomb radiocarbon pulse in growth bands formed during the 1960s with a timing which was very similar to that of marine carbonates. Using radiocarbon assays of spine growth bands known to have formed in the 1960s and 1970s as a dated marker, we confirm the validity of spine enamel growth band counts as accurate annual age indicators to an age of at least 45 year. Radiocarbon incorporation into northeast Atlantic dogfish spines occurred in similar years as those in the northwest Atlantic and northeast Pacific, although the amount of radiocarbon differed in keeping with the radiocarbon content of the different water masses. Published reports suggesting that Pacific dogfish are longer lived and slower growing than their Atlantic counterparts appear to be correct, and are not due to errors in interpreting the spine growth bands. Radiocarbon assays of fin spine enamel appears to be well suited to the age validation of sharks with fin spines which inhabit the upper 200 m of the ocean.     

Aspects of the reproductive biology of two pelagic sharks in the eastern Atlantic Ocean

This study used data provided by the Chinese Longline Fishery Scientific Observer Programme from the tropical eastern Atlantic Ocean to estimate the reproductive parameters of the blue shark (Prionace glauca) and crocodile shark (Pseudocarcharias kamoharai). Sizes ranged from 80 to 298 cm fork length (FL) for blue sharks and from 48 to 99 cm FL for crocodile sharks. Sexual segregation was observed during different months for both sharks. The sex ratio for blue sharks was 1.38 F:1 M, and 1 F:2.79 M for crocodile sharks. The size of adult blue sharks ranged from 144 to 280 cm for males and from 174 to 298 cm for females; and that of crocodile sharks from 63 to 97 cm for males and 78–99 cm for females. The size at 50% of maturity for blue sharks was estimated at 191.7 cm FL for females and 197.5 cm FL for males, and that of crocodile sharks was assessed at 84.9 cm FL for females and 78.5 cm FL for males. Most sexually matured females were pregnant; their means were 207.2 ± 16.4 cm FL for blue sharks and 89.4 ± 4.3 cm FL for crocodile sharks. Mature sizes for both species were significantly different among months. Embryonic sizes also varied widely among months for crocodile sharks, but a slight change was recorded for those of blue sharks. The observed mean size at birth and litter size were 34.5 cm FL and 37 ± 12 for the blue sharks, and that of the crocodile sharks, 39.5 cm FL and a dominant four embryos in the uterus. Due to the observed increasing catch trend of blue sharks and the slow reproductive cycle of crocodile sharks, this study presents the need of implementing conservation measures to ensure the sustainability of both species in their habitat.     

Growth and Maximum Size of Tiger Sharks (Galeocerdo cuvier) in Hawaii

Tiger sharks (Galecerdo cuvier) are apex predators characterized by their broad diet, large size and rapid growth. Tiger shark maximum size is typically between 380 & 450 cm Total Length (TL), with a few individuals reaching 550 cm TL, but the maximum size of tiger sharks in Hawaii waters remains uncertain. A previous study suggested tiger sharks grow rather slowly in Hawaii compared to other regions, but this may have been an artifact of the method used to estimate growth (unvalidated vertebral ring counts) compounded by small sample size and narrow size range. Since 1993, the University of Hawaii has conducted a research program aimed at elucidating tiger shark biology, and to date 420 tiger sharks have been tagged and 50 recaptured. All recaptures were from Hawaii except a single shark recaptured off Isla Jacques Cousteau (24°13′17″N 109°52′14″W), in the southern Gulf of California (minimum distance between tag and recapture sites  =  approximately 5,000 km), after 366 days at liberty (DAL). We used these empirical mark-recapture data to estimate growth rates and maximum size for tiger sharks in Hawaii. We found that tiger sharks in Hawaii grow twice as fast as previously thought, on average reaching 340 cm TL by age 5, and attaining a maximum size of 403 cm TL. Our model indicates the fastest growing individuals attain 400 cm TL by age 5, and the largest reach a maximum size of 444 cm TL. The largest shark captured during our study was 464 cm TL but individuals >450 cm TL were extremely rare (0.005% of sharks captured). We conclude that tiger shark growth rates and maximum sizes in Hawaii are generally consistent with those in other regions, and hypothesize that a broad diet may help them to achieve this rapid growth by maximizing prey consumption rates.     

Age and growth of Atlantic bluefin tuna, Thunnus thynnus (Osteichthyes: Thunnidae), in the Mediterranean Sea

The objective of the study was to describe the biometry of Mediterranean bluefin tuna, Thunnus thynnus, the biology of which is not yet well understood. A total of 504 specimens was collected from 1998 to 2005 in the central part of the Mediterranean basin. They were sexed and measured; fork lengths (FL) ranged from 51.0 to 255.0 cm while body weights (W) ranged from 2.6 to 247.0 kg. The first spiniform ray (spine) of the first dorsal fin was removed and cross‐sectioned near the condyle base in order to count annuli for age estimation. The regression coefficient (b) of the female FL–W relationship was significantly higher than that of the male, and both sexes displayed a negatively allometric growth (b < 3); male regression equation: ln W = ?2.942 + 2.730 ln FL; female regression equation: ln W = ?3.660 + 2.878 ln FL. Based on counts of the translucent zones in the sections of the first ray of the first dorsal fin, estimated ages ranged from 1 to 15 years for males and 1 to 14 years for females. The correlation between the spine ray (R) and FL fit the allometric model best; the R–FL regression equations of the two sexes did not differ significantly and the overall equation was: ln FL = 3.721 + 0.851 ln R. Due to the R–FL allometric correlation, estimates of fork lengths at previous ages, FL_i, were back‐calculated with a body proportional hypothesis. Von Bertalanffy growth equations were derived from both observed and back‐calculated FLs‐at‐age, which did not differ significantly. Moreover, no significant difference was found between the growth equations of the two sexes; the overall equation was FL_t = 373.08 [1?e^{?0.07(t + 1.76)}]. Weight‐at‐age values were derived from the von Bertalanffy predicted FLs‐at‐age by the FL–W correlation equations for males and females. The paper represents the first comprehensive study on the biometry, including age and growth, of bluefin tuna captured in the Mediterranean Sea.     

Age,growth and maturity of tub gurnard (Chelidonichthys lucerna Linnaeus 1758; Triglidae) in the inshore coastal waters of Northwest Wales,UK

The tub gurnard Chelidonichthys lucerna has been identified by ICES as a potential commercial species in the northeast Atlantic with recommendations made to monitor landings and discards and to derive information on population biology for stock assessment purposes, however, data are lacking for the species in the northeast Atlantic. Therefore, aims of this study were to provide data on the size/age‐structure and patterns of growth, maturity and mortality of C. lucerna in Northwest Wales, UK, and in doing so to provide data on the biological characteristics of the most northerly population studied to date for comparison with the existing data for southerly Mediterranean populations. Data on the age, growth and maturity of C. lucerna were collected by otter trawling (73 mm cod‐end stretched mesh size) in the coastal waters of Northwest Wales, UK in October (2000–2011, excluding 2006). Total length (TL) of fish sampled ranged between 10.5–41.0 cm (males) and 10.4–57.5 cm (females). The majority of the female fish were between 20–30 cm TL (60.2%) and the majority of the male fish between 20–30 cm TL (58.3%) respectively. TL/weight (W) relations for male and female fish were similar and the combined data was described by W = 0.0067 TL^3.10. Age of fish ranged between 1–7 years old for female fish and 1–5 years old for male fish respectively with the majority of female fish 3 years old (40%) and the majority of male fish 3 years old (37%). The age structures of female and male tub gurnards were not significantly different with the older age classes consisting predominantly of female fish. Both males and females exhibited similar asymptotic growth patterns and the combined von Bertalanffy growth function was TL_t = 51.6 (1 ? e ^{[?0.25(t + 0.41)]}). Instantaneous rates of total mortality were calculated as 1.04 year^?1 for males and 1.11 year^?1 for females. The size (L₅₀) and age at first maturity (A₅₀) were estimated to be 29.1 cm TL and 2.8 years for males, 27.7 cm TL and 2.7 years for females and 28.0 cm TL and 2.8 years for both sexes combined. The results of this study provide the first information on the biology and population dynamics of C. lucerna in the Irish Sea, the first data collected in the northeast Atlantic since 1985 and the most northerly population studied to date.     

Reassessment of spiny dogfish Squalus acanthias age and growth using vertebrae and dorsal-fin spines

Male and female spiny dogfish Squalus acanthias were collected in the western North Atlantic Ocean in the Gulf of Maine between July 2006 and June 2009. Squalus acanthias ranged from 25 to 102 cm stretch total length and were caught during all months of the year except January. Age estimates derived from banding patterns visible in both the vertebrae and second dorsal-fin spines were compared. Vertebral growth increments were visualized using a modified histological staining technique, which was verified as appropriate for obtaining age estimates. Marginal increment analysis of vertebrae verified the increment periodicity, suggesting annual band deposition. Based on increased precision and accuracy of age estimates, as well as more biologically realistic parameters generated in growth models, the current study found that vertebrae provided a more reliable and accurate means of estimating age in S. acanthias than the second dorsal-fin spine. Age estimates obtained from vertebrae ranged from <1 year-old to 17 years for male and 24 years for female S. acanthias. The two-parameter von Bertalanffy growth model fit to vertebrae-derived age estimates produced parameters of L∞ = 94·23 cm and k = 0·11 for males and L∞ = 100·76 cm and k = 0·12 for females. While these growth parameters differed from those previously reported for S. acanthias in the western North Atlantic Ocean, the causes of such differences were beyond the scope of the current study and remain to be determined.     

A re-examination of the age and growth of sand tiger sharks, Carcharias taurus, in the western North Atlantic: the importance of ageing protocols and use of multiple back-calculation techniques

Age and growth estimates for sand tiger sharks, Carcharias taurus, in the western North Atlantic were derived from 96 vertebral centra collected from sharks ranging from 94 to 277 cm total length (TL), and compared to previously published age and growth data. The oldest female and male sand tiger sharks aged in this study were 17 and 15 years of age, respectively. von Bertalanffy growth parameters derived from vertebral length-at-age data are L _∞ = 295.8 cm TL, k = 0.11 year⁻¹, and t ₀ = −4.2 years for females, and L _∞ = 249.5 cm TL, k = 0.16 year⁻¹, and t ₀ = −3.4 years for males. Sexual maturity is estimated to be 9–10 years for females and 6–7 years for males. Weight-to-length relationships determined for female and male sand tiger sharks in the western North Atlantic are; W = 1.3 × 10⁻⁴ × L ^2.4 (r ² = 0.84, n = 55) and W = 9.0 × 10⁻⁵ × L ^2.5 (r ² = 0.84, n = 47), respectively, and 7.9 × 10⁻⁵ × L ^2.5 (r ² = 0.84) for the sexes combined. Our results show sand tigers possess a slower rate of growth than previously thought. This information is crucial for accurately assessing this population’s ability to recover, and further justifies the need for this species to be fully protected.     

Insights into vertebral band pair deposition rate in the juvenile common thresher shark (Alopias vulpinus) in the northeastern Pacific Ocean

Validation of band pair deposition rates in elasmobranch vertebrae is essential for accurate age estimation using band pair counting techniques. We present a validation study of the vertebral band pair deposition rate for juvenile common thresher sharks Alopias vulpinus in the northeastern Pacific Ocean (NEPO) using tag and recapture with oxytetracycline (OTC) injection. A total of 14 juvenile A. vulpinus marked with OTC from 1998 through 2013 were recaptured with times at liberty ranging from 1.08 to 3.81 years with an average of 2.14 years (±0.97 years standard deviation, SD). Shark size ranged from 80 to 128 cm fork length (LF) at the time of OTC injection and from 112 to 168 cm LF for those measured at recapture. The slopes of the relationships between band pairs post OTC and years at liberty for each reader ranged from 0.84 to 0.95, slightly lower than the 1.0 slope expected from annual band pair formation. These findings preliminarily support previous age and growth assumptions based on a one band pair per year deposition rate. However, high variation in band pair deposition rates between samples, coupled with regression slopes falling just under one band pair per year, indicates that further investigation is needed to refine band pair deposition rate estimates.     

Slow growth of the overexploited milk shark Rhizoprionodon acutus affects its sustainability in West Africa

Age and growth of Rhizoprionodon acutus were estimated from vertebrae age bands. From December 2009 to November 2010, 423 R. acutus between 37 and 112 cm total length (L_T) were sampled along the Senegalese coast. Marginal increment ratio was used to check annual band deposition. Three growth models were adjusted to the length at age and compared using Akaike's information criterion. The Gompertz growth model with estimated size at birth appeared to be the best and resulted in growth parameters of L_∞ = 139·55 (L_T) and K = 0·17 year^?1 for females and L_∞ = 126·52 (L_T) and K = 0·18 year^?1 for males. The largest female and male examined were 8 and 9 years old, but the majority was between 1 and 3 years old. Ages at maturity estimated were 5·8 and 4·8 years for females and males, respectively. These results suggest that R. acutus is a slow‐growing species, which render the species particularly vulnerable to heavy fishery exploitation. The growth parameters estimated in this study are crucial for stock assessments and for demographic analyses to evaluate the sustainability of commercial harvests.     

Age and growth of the blacknose shark, <Emphasis Type="Italic">Carcharhinus acronotus</Emphasis>, in the western North Atlantic Ocean with comments on regional variation in growth rates

We examined the age and growth of the blacknose shark, Carcharhinus acronotus, in the western North Atlantic Ocean by obtaining direct age estimates using vertebral centra. We verified annual deposition of growth increments with marginal increment analysis and validated it by analyzing vertebrae marked with oxytetracycline from a female blacknose shark held in captivity. Von Bertalanffy growth parameters indicated that female blacknose sharks have a lower growth constant (k), a larger theortical maximum size (L), and are longer lived than males. We compared these growth parameters for blacknose sharks in the western North Atlantic Ocean to growth parameters for blacknose sharks collected in the eastern Gulf of Mexico to test for differences between regions. Females in the western North Atlantic Ocean have a significantly lower L, lower k, and a higher theoretical longevity than females in the Gulf of Mexico. Males in the western North Atlantic Ocean have a higher L<>, lower k, and higher theoretical longevity than males in the Gulf of Mexico. The significant differences between these life history parameters for blacknose sharks suggest that, when possible, future management initiatives concerning blacknose sharks should consider managing the populations in the western North Atlantic and the Gulf of Mexico as separate stocks.     

Ageing culturally significant relic trees in southeast Queensland to support bushfire management strategies

Appropriate fire management strategies are needed to protect forests and large old ecologically and culturally significant trees in natural landscapes. The aim of this study was to determine the age of large old and relic trees of cultural significance that included Cypress Pine (Callitris columellaris F. Muell.), a species that is sensitive to crown scorching fires in a fire‐prone landscape, and to calibrate a tree‐growth‐rate method for estimating tree age. Twelve large trees were dated using radiocarbon (¹⁴C) dating. The trees are located on North Stradbroke Island (Indigenous name: Minjerribah), southeast Queensland (Australia) in a fire‐prone landscape where recent wildfires have destroyed many large trees. The median tree ages ranged from 155 to 369 years. These results suggest an important role of past Indigenous land management practices in protecting Cypress Pine from crown scorching fires. The tree‐growth‐rate‐based method for estimating tree age generally overestimated the age derived from radiocarbon dating. Bias correction factors were developed for correcting various measures of periodic growth rates. This study provides evidence that appropriate low‐intensity fire strategies have the potential to contribute to the survival of forests and conserve large old trees.