Seasonal changes in the physiology of brook trout, Salvelinus fontinalis (Mitchill), in a sub-Arctic river system

Wild herring larvae were sampled from the Firth of Clyde, Scotland in 1980 and 1981 to assess the suitability of otolith ageing methods for herring larval studies. The ages and growth rates were estimated directly from otolith ring counts, by comparison of ring counts with the results of validation studies on reared larvae of known age, and by back-calculation. Growth rates based on otolith data were compared with indirect calculations based on length at capture. The assumption of daily ring deposition, even allowing for a period of lag before the initiation of daily ring deposition, led to incorrect predictions for age at yolk-sac absorption, and back-calculations led to overestimated growth rates, indicating that ring deposition does not begin at hatching and that light microscope counts give a rate different from one ring per day in wild herring larvae.     

Age determination, bomb-radiocarbon validation and growth of Atlantic halibut (Hippoglossus hippoglossus) from the Northwest Atlantic

Atlantic halibut (Hippoglossus hippoglossus) is the largest and one of the most widely-ranging and commercially-valuable groundfish in the Atlantic Ocean. Although presumed to be long-lived, their age and growth has not been validated. Ages were estimated by counting growth increments from approximately 2400 thin-sectioned sagittal otoliths collected from the Scotian Shelf and southern Grand Banks off eastern Canada. The accuracy of age estimates made from otolith thin sections was validated using bomb-radiocarbon assays of 13 otolith cores whose year of formation ranged from 1949 to 1975, encompassing the timeframe of the global radiocarbon pulse. Known-age juvenile halibut from a culture facility were used to identify the approximate location of the first annulus. Growth rate for males and females was similar up to about 70 cm (~5 years), after which point male growth slowed, while female growth continued to an age of up to 38 years and a maximum observed size of 232 cm. Males grew to an observed maximum length of about 175 cm and a maximum age of 50 years. A comparison of age estimates for otoliths collected in a ‘historic’ time frame (1963 to 1974) with those from recent years (1997 to 2007) shows that growth rate has not changed appreciably between the two time periods. Small but significant growth differences were observed between the Scotian Shelf and southern Grand Banks for both sexes, while large differences in length at age were observed between halibut caught with longline compared to otter trawl due to differences in length-based gear selectivity. Age interpretations based on sectioned otoliths tended to be 10–15% different than those based on break and burn, although the age comparison was confounded by other variables and must be considered provisional. Atlantic halibut is a long-lived fish, living up to at least 50 years, an important consideration for the management of the fishery.     

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.     

Is otolith science transformative? New views on fish migration

Publications focused on fish otolith applications grew exponentially from 1981–2000 but have subsequently stabilized, suggesting maturation of otolith applications. Over the past 30 years >3,500 primary journal publications were identified prompting the question, how have otoliths fundamentally changed our understanding of fishes and their environment? Has otolith science in this way been transformative? I use Harden Jones’ 1968 Fish Migration as a benchmark of fish migration concepts prior to Pannella’s 1971 breakthrough paper on otolith microstructure. For case study species I highlight how otolith science has informed migration concepts including, (1) parent stream theory (Atlantic bluefin tuna), (2) adoptive homing (Atlantic herring), and (3) partial migration (European eel). Harden Jones’ overall conclusion that life cycle closure leads to population structure (also known as the migration triangle) is a first principle in fisheries science, but in recent years has been challenged by otolith science. In particular, a transformative discovery attributable to otoliths is that life cycles vary substantially within populations. New avenues of otolith science are now exploring the causes and consequences of this life cycle diversity, and large advances are expected through integration of otolith approaches with electronic tagging, genetics, field manipulations, and modeling that will permit migration concepts to be tested at multiple ecological scales.     

Suitability of glycerin-preserved otoliths for age validation using bomb radiocarbon

The effect of glycerin storage on the bomb radiocarbon content of otoliths was determined experimentally. Storage in either pre- or post-bomb glycerin had no detectable effect on the bomb radiocarbon content of either pre- or post-bomb otoliths. Therefore bomb-dated age validation studies need not be restricted to freshly-collected samples or dry otoliths, implying that the large numbers of glycerin-archived otoliths around the world are suitable for age validation studies using bomb radiocarbon.     

Using otolith weight to age fish

The problem of determining and verifying ages of fish, from populations having a considerable variation in size at age, has been investigated using the relationship between otolith size and fish size, which has been shown by several authors to be influenced by growth rate. In such a population of Sardinella aurita Val. an index of age can be obtained for individual fish by calculating the equivalent otolith weight at a particular fish length, using the otolith weight–fish length relationship determined for each age group. This statistic not only permits a much greater proportion of fish to be assigned ages than is possible with otolith reading alone, but also enables the age groups to be verified as year classes. However, it is concluded that, although appropriate models based on otolith-fish size relationships can predict age for groups of fish in which growth rates are known or can be assumed to be consistent, such techniques have a limited application in ageing fish from wild populations with highly variable growth rates.     

Do otolith increments allow correct inferences about age and growth of coral reef fishes?

Otolith increment structure is widely used to estimate age and growth of marine fishes. Here, I test the accuracy of the long-term otolith increment analysis of the lemon damselfish Pomacentrus moluccensis to describe age and growth characteristics. I compare the number of putative annual otolith increments (as a proxy for actual age) and widths of these increments (as proxies for somatic growth) with actual tagged fish-length data, based on a 6-year dataset, the longest time course for a coral reef fish. Estimated age from otoliths corresponded closely with actual age in all cases, confirming annual increment formation. However, otolith increment widths were poor proxies for actual growth in length [linear regression r ² = 0.44–0.90, n = 6 fish] and were clearly of limited value in estimating annual growth. Up to 60 % of the annual growth variation was missed using otolith increments, suggesting the long-term back calculations of otolith growth characteristics of reef fish populations should be interpreted with caution.     

Accounting for Age Uncertainty in Growth Modeling,the Case Study of Yellowfin Tuna (Thunnus albacares) of the Indian Ocean

Age estimates, typically determined by counting periodic growth increments in calcified structures of vertebrates, are the basis of population dynamics models used for managing exploited or threatened species. In fisheries research, the use of otolith growth rings as an indicator of fish age has increased considerably in recent decades. However, otolith readings include various sources of uncertainty. Current ageing methods, which converts an average count of rings into age, only provide periodic age estimates in which the range of uncertainty is fully ignored. In this study, we describe a hierarchical model for estimating individual ages from repeated otolith readings. The model was developed within a Bayesian framework to explicitly represent the sources of uncertainty associated with age estimation, to allow for individual variations and to include knowledge on parameters from expertise. The performance of the proposed model was examined through simulations, and then it was coupled to a two-stanza somatic growth model to evaluate the impact of the age estimation method on the age composition of commercial fisheries catches. We illustrate our approach using the saggital otoliths of yellowfin tuna of the Indian Ocean collected through large-scale mark-recapture experiments. The simulation performance suggested that the ageing error model was able to estimate the ageing biases and provide accurate age estimates, regardless of the age of the fish. Coupled with the growth model, this approach appeared suitable for modeling the growth of Indian Ocean yellowfin and is consistent with findings of previous studies. The simulations showed that the choice of the ageing method can strongly affect growth estimates with subsequent implications for age-structured data used as inputs for population models. Finally, our modeling approach revealed particularly useful to reflect uncertainty around age estimates into the process of growth estimation and it can be applied to any study relying on age estimation.     

Using otolith weight to estimate the age of haddock (Melanogrammus aeglefinus): a tree model application

In this study, we investigated whether otolith weight can be used to estimate fish age with the same level of accuracy as that of the traditional annuli counting technique in a commercially important species such as haddock (Melanogrammus aeglefinus). Results indicate that this method is highly effective for young fish (around 97% correct classification), whereas its powers of prediction decrease with the increasing age of the fish. For older fish, the otolith weight cannot be an accurate estimator of fish age if the weight overlap between the different age classes is too large. Nevertheless, the otolith weight technique is strictly dependent on correct age determination through the counting of annuli of those individuals used in the calibration. Hence, an increase in the accuracy of ageing obviously determines an increase of the power of otolith weight to estimate fish age. Therefore, we suggest that otolith weight could represent a routine technique for determination of the age structure of haddock populations. This technique has the merit to be objective, fast, 100% repeatable and has the same level of accuracy as that of annuli counting.     

Otolith variation in Pacific herring (<Emphasis Type="Italic">Clupea pallasii</Emphasis>) reflects mitogenomic variation rather than the subspecies classification

Pacific herring (Clupea pallasii) is divided into three subspecies: two in northeast Europe and one in the north Pacific Ocean. Genetic studies have indicated that the populations in northeast Europe have derived from the northwest Pacific herring recently, or during the last 10–15 kyr, and that they are distinct from the population in the northeast Pacific. In addition, hybridization between the Pacific herring and the Atlantic herring has been documented. Otolith variation has been considered to be largely affected by environmental variation, but here we evaluate whether the genetic differentiation is reflected in otolith shape differences. A clear difference in otolith shape was observed between the genetically differentiated herring species Clupea harengus from the Atlantic and C. pallasii. The otolith shape of C. p. suworowi in the Barents Sea was different from the shape of C. pallasii in northern Norway and C. p. pallasii from the Pacific. Populations of C. p. pallasii, sampled east and west of the Alaska Peninsula, which belong to two genetically different clades of the C. p. pallasii in the Pacific Ocean, show a clear difference in otolith shape. C. p. suworowi and the local C. pallasii peripheral population in Balsfjord in northern Norway are more similar to the northwest Pacific herring (C. p. pallasii) than to the northeast Pacific herring (C. p. pallasii), both genetically and in otolith shape. The Balsfjord population, known to be influenced by introgression of mtDNA from the Atlantic herring, does not show any sign of admixture in otolith shape between the two species. A revised classification, considering the observed genetic and morphological evidence, should rather group the northwest Pacific herring in the Bering Sea together with the European populations of C. pallasii than with the northeast Pacific herring in the Gulf of Alaska.     

Validation of yellowedge grouper, <Emphasis Type="Italic">Epinephelus flavolimbatus</Emphasis>, age using nuclear bomb-produced radiocarbon

Age validation and estimates of longevity of yellowedge grouper (Epinephelus flavolimbatus) from the Gulf of Mexico (GOM) are needed to inform fishery management decisions. Yellowedge grouper sagittal otoliths (n = 100) were collected, aged using conventional means, and cores were submitted for radiocarbon (¹⁴C) measurement. Radiocarbon values of yellowedge grouper otoliths were compared to established radiocarbon chronologies in the region to validate the age and ageing methodology of this species. The yellowedge grouper chronology displayed a similar sigmoidal trend as previously published chronologies. In addition to the core analysis, multiple areas on otolith sections from eight specimens were analyzed for Δ¹⁴C to validate age estimates for fish born prior to the ¹⁴C increase. Our results indicate that yellowedge grouper live longer than previously reported (minimum of 40 years based on radiocarbon measurements). The validated ageing methodology supported an estimated maximum longevity of 85 years and established that yellowedge grouper have the longest lifespan currently known for any species of grouper in the GOM. Results also indicate a depth-age interaction in that material extracted from adult otolith sections assigned to post-bomb dates exhibited lower Δ¹⁴C values than cores (juvenile material) assigned to the same post-bomb dates. This finding is likely explained by lower ¹⁴C levels reported from water masses at deeper depths (>100 m) which are inhabited by adults.     

Seeking the true time: Exploring otolith chemistry as an age-determination tool

Fish otoliths' chronometric properties make them useful for age and growth rate estimation in fisheries management. For the Eastern Baltic Sea cod stock (Gadus morhua), unclear seasonal growth zones in otoliths have resulted in unreliable age and growth information. Here, a new age estimation method based on seasonal patterns in trace elemental otolith incorporation was tested for the first time and compared with the traditional method of visually counting growth zones, using otoliths from the Baltic and North seas. Various trace elemental ratios, linked to fish metabolic activity (higher in summer) or external environment (migration to colder, deeper habitats with higher salinity in winter), were tested for age estimation based on assessing their seasonal variations in concentration. Mg:Ca and P:Ca, both proxies for growth and metabolic activity, showed greatest seasonality and therefore have the best potential to be used as chemical clocks. Otolith image readability was significantly lower in the Baltic than in the North Sea. The chemical (novel) method had an overall greater precision and percentage agreement among readers (11.2%, 74.0%) than the visual (traditional) method (23.1%, 51.0%). Visual readers generally selected more highly contrasting zones as annuli whereas the chemical readers identified brighter regions within the first two annuli and darker zones thereafter. Visual estimates produced significantly higher, more variable ages than did the chemical ones. Based on the analyses in our study, we suggest that otolith microchemistry is a promising alternative ageing method for fish populations difficult to age, such as the Eastern Baltic cod.     

A comparison of different structures and methods for estimating age of northern-form Dolly Varden <Emphasis Type="Italic">Salvelinus malma malma</Emphasis> from the Canadian Arctic

Assessment of anadromous northern-form Dolly Varden Salvelinus malma malma in the Western Canadian Arctic requires reliable methods for estimating ages. Additionally, conservation efforts warrant determining whether fin rays provide a non-lethal alternative to otoliths. Precision and bias of whole and sectioned otoliths, and sectioned pectoral and pelvic fin rays were examined. Two age readers with different levels of experience ageing this species read each structure three times. Coefficient of variation (CV) was calculated to measure precision, and age bias plots were created for each method of preparation/structure within and between readers. The experienced reader demonstrated the highest precision with sectioned otoliths (CV = 1.6 %) followed by whole otoliths (CV = 4.2 %) while pectoral and pelvic fins were the lowest, CV = 7.7 % and 7.5 %, respectively. The age bias plot showed little difference between whole and sectioned otoliths, although greater imprecision/bias was evident for whole otoliths at age ≥9. Compared to otoliths, fin rays produced younger age estimates starting at 5 years; however, pelvic fins were more biased towards younger estimates than pectoral fins. The less experienced reader had greater inconsistencies, tending to overage younger and underage older samples for all methods compared to the more experienced reader, underscoring the importance of experience when estimating age for this species. We conclude that both types of fin rays are a poor non-lethal alternative to otoliths for fish ≥5 years and recommend an experienced ager could use whole otoliths up to age 8 and sectioned otoliths for fish ≥9 years (>500 mm fork length).     

Strontium:cakium concentration ratios in otoliths of herring larvae as indicators of environmental histories

Synopsis Elemental analyses, using wave-length dispersive electron microprobe techniques on otoliths from reared Atlantic herring larvae, Clupea harengus, showed trace quantities of strontium relative to that of calcium, and an inverse relationship between Sr/Ca concentration ratios and rearing temperature. These data are consistent with those for coral aragonite, in that there appears to be an inverse temperature effect on physiological incorporation of strontium in the otolith aragonite. Our determinations of Sr/Ca concentration ratios of lab-reared herring larvae showed that the deposition of strontium relative to calcium and the rearing temperature were related, where: T (° C) = −2.955 [Sr/Ca] × 1000 ± 19.172. This principle thus makes it possible to use Sr/Ca concentration ratios in fish otoliths to delineate past temperatures experienced by an individual. Further, combining electron microprobe analyses with scanning electron microscope (SEM) examinations of daily increments in the same otolith makes it possible to reconstruct the temperature history for an individual fish on a time scale of days. An example of the application of the technique to an approximately six-month-old field-caught herring larva is given, and the limitations of the technique are discussed.     

Pacific halibut chronology of bomb radiocarbon in otoliths from 1944 to 1981 and a validation of ageing methods

Pacific halibut Hippoglossus stenolepis of known age were used to create a reference chronology of radiocarbon levels for the years 1944–1981. Levels of radiocarbon in Pacific halibut otoliths pre‐1958 were among the lowest reported to date, but radiocarbon levels increased dramatically post‐1960. Subsequently, this reference chronology was used to validate ages determined using the standard break and burn and surface ageing technique. These older fish were collected at a later date but were born during this period of rapidly increasing radiocarbon levels. Otolith cores were compared to the reference chronology based upon the presumed birth year determined from annulus counts and year of capture. The ages derived from both break and burn and surface ageing methods were determined to be accurate.     

Validation of first annulus formation in red snapper otoliths with the use of an alizarin complexone fluorescent marker

The red snapper (Lutjanus campechanus) is among the most studied reef fish species in the Gulf of Mexico. Several studies have used counts of annuli in sectioned sagittal otoliths to age red snapper. However, interpretation of the putative first annulus has been a major source of debate among otolith readers throughout the Gulf of Mexico. Our objective was to use the chemical marker alizarin complexone to validate the periodicity of first opaque annulus formation in red snapper otolith sections. Juvenile red snapper were immersed in 100 mg alizarin complexone per L seawater solution in November 2005 and then reared in 6000 l circular tanks until July 2006. Otoliths were then removed from the fish and thin sectioned. All experimental otolith sections displayed a distinct fluorescent mark ranging from 0.62 to 0.96 mm from the core when viewed under the microscope with a rhodamine filter. The diffuse opaque annulus was located distally to the alizarin mark in all specimens (ranging from 0.88 to 1.51 mm). The distal position of the presumptive first annulus relative to the alizarin mark in all specimens indicates that this diffuse opaque annulus in red snapper sectioned otoliths forms during the first winter after hatching. Translucent marginal edges of all otolith sections indicate that first opaque annulus formation is completed by mid-July.     

Screening for Viral Hemorrhagic Septicemia Virus in Marine Fish along the Norwegian Coastal Line

Viral hemorrhagic septicemia virus (VHSV) infects a wide range of marine fish species. To study the occurrence of VHSV in wild marine fish populations in Norwegian coastal waters and fjord systems a total of 1927 fish from 39 different species were sampled through 5 research cruises conducted in 2009 to 2011. In total, VHSV was detected by rRT-PCR in twelve samples originating from Atlantic herring (Clupea harengus), haddock (Melanogrammus aeglefinus), whiting (Merlangius merlangus) and silvery pout (Gadiculus argenteus). All fish tested positive in gills while four herring and one silvery pout also tested positive in internal organs. Successful virus isolation in cell culture was only obtained from one pooled Atlantic herring sample which shows that today''s PCR methodology have a much higher sensitivity than cell culture for detection of VHSV. Sequencing revealed that the positive samples belonged to VHSV genotype Ib and phylogenetic analysis shows that the isolate from Atlantic herring and silvery pout are closely related. All positive fish were sampled in the same area in the northern county of Finnmark. This is the first detection of VHSV in Atlantic herring this far north, and to our knowledge the first detection of VHSV in silvery pout. However, low prevalence of VHSV genotype Ib in Atlantic herring and other wild marine fish are well known in other parts of Europe. Earlier there have been a few reports of disease outbreaks in farmed rainbow trout with VHSV of genotype Ib, and our results show that there is a possibility of transfer of VHSV from wild to farmed fish along the Norwegian coast line. The impact of VHSV on wild fish is not well documented.     

Recruitment dynamics and first year growth of the coral reef surgeonfish Ctenochaetus striatus,with implications for acanthurid growth models

Newly recruited Ctenochaetus striatus were monitored over a 16-month period in American Samoa, 2002–2003. During this period, a mass recruitment of age-0 C. striatus occurred in March 2002 with numbers reaching 22.9 recruits m^?2. This program provided an invaluable opportunity to (1) analyze the dynamics of a mass recruitment episode and to assess its significance with respect to more typical patterns of recruitment and (2) establish the pattern of recruit growth during their first year of life. Age-based analysis indicated that the mass recruitment generated about 90 % of annual recruitment, but recruit mortality was high; thus, most recruitment was provided by continuous settlement throughout the year. The mass event appeared to be a short-lived pulse with recruits residing on the reef an average of 14.1 d compared with 161.1 d for other recruits. Recruits grew rapidly, achieving 90 % of their adult size during their first year, and they formed their first otolith annulus after 1 yr, thereby providing a firm basis for otolith interpretation of fish ages during the early life history phase of this species. The extensive age-based documentation of their first year growth in this study validates the distinctive “square” growth pattern exhibited by acanthurids as described in the literature (i.e., long life span with rapid initial growth that quickly reaches an asymptotic size), and it demonstrates the impact that the presence of age-0 fish has when generating growth parameters for populations exhibiting square growth. We found that the parameters from the re-parameterized von Bertalanffy growth function have preferred characteristics when modeling square growth in fish and that fixing age-at-length zero to pelagic larval duration is a preferable method to constrain growth models when lacking age-0 fish.     

Searching for the Oldest Baobab of Madagascar: Radiocarbon Investigation of Large Adansonia rubrostipa Trees

We extended our research on the architecture, growth and age of trees belonging to the genus Adansonia, by starting to investigate large individuals of the most widespread Malagasy species. Our research also intends to identify the oldest baobabs of Madagascar. Here we present results of the radiocarbon investigation of the two most representative Adansonia rubrostipa (fony baobab) specimens, which are located in south-western Madagascar, in the Tsimanampetsotse National Park. We found that the fony baobab called “Grandmother” consists of 3 perfectly fused stems of different ages. The radiocarbon date of the oldest sample was found to be 1136 ± 16 BP. We estimated that the oldest part of this tree, which is mainly hollow, has an age close to 1,600 yr. This value is comparable to the age of the oldest Adansonia digitata (African baobab) specimens. By its age, the Grandmother is a major candidate for the oldest baobab of Madagascar. The second investigated specimen, called the “polygamous baobab”, consists of 6 partially fused stems of different ages. According to dating results, this fony baobab is 1,000 yr old. This research is the first investigation of the structure and age of Malagasy baobabs.