Context Within the western and central North Pacific Ocean stock of striped marlin (Kajikia audax), reproductive information is lacking for the central North Pacific. Aims To derive estimates of sex-specific functional maturity ogives, eye-to-fork length at 50% maturity, reproductive phase composition, spawning season and sex ratios by sampling the Hawaii-based pelagic longline fishery. Methods Gonad histology was exclusively used to confirm sex and estimate all reproductive life history traits. Key results Females spawn in May–July; males are capable of spawning year-round. Best length at 50% maturity estimates were based on datasets restricted to the female spawning season (eye–fork length for females 152.2 cm and for males 109.0 cm). Length distributions and sex ratios are seasonally dynamic around Hawaii and differ from those in other Pacific regions. Conclusions Length-at-maturity estimates for the central North Pacific are lower than all sex-specific estimates reported from other Pacific regions. Implications The new maturity estimates will improve determinations of stock resilience and productivity in upcoming international assessments.
Element:Ca ratios in the otolith cores of young-of-the-year (YOY) swordfish, Xiphias gladius, were used as natural tracers to predict the nursery origin of subadult and adult swordfish from three foraging grounds in the North Pacific Ocean (NPO). First, the chemistry of otolith cores (proxy for nursery origin) was used to develop nursery-specific elemental signatures in YOY swordfish. Sagittal otoliths of YOY swordfish were collected from four regional nurseries in the NPO between 2000 and 2005: (1) Central Equatorial North Pacific Ocean (CENPO), (2) Central North Pacific Ocean (CNPO), (3) Eastern Equatorial North Pacific Ocean (EENPO) and (4) Western North Pacific Ocean (WNPO). Calcium (43 Ca), magnesium (24 Mg), strontium (88 Sr) and barium (138 Ba) were quantified in the otolith cores of YOY swordfish using laser ablation inductively coupled plasma mass spectrometry. Univariate tests indicated that three element:Ca ratios (Mg:Ca, Sr:Ca and Ba:Ca) were significantly different among nurseries. Overall classification success of YOY swordfish to their nursery of collection was 72% based on quadratic discriminant analysis. Next, element:Ca ratios in the otolith cores of subadults and adults collected from three foraging grounds where targeted fisheries exist (Hawaii, California and Mexico) were examined to calculate nursery-specific contribution estimates. Mixed-stock analysis indicated that the CENPO nursery contributed the majority of individuals to all three foraging grounds (Hawaii 45.6 ± 13.2%, California 84.6 ± 10.8% and Mexico 64.5 ± 15.9%). The results from this study highlight the importance of the CENPO nursery and provide researchers and fisheries managers with new information on the connectivity of the swordfish population in the NPO.
Hawaiian grouper (Hyporthodus quernus) is endemic to the Hawaiian Islands and is regionally important, yet little is known about its life history. This large species is managed within the Deep 7 bottomfish complex, which includes six snapper species that are assumed to have similar life history traits. Previous age estimates were not validated and suggested a maximum age of 34 years. To evaluate the preliminary study and provide a valid basis for life history parameters, we aged otoliths using bomb radiocarbon (14C) dating. Measured 14C values provided ages for smallest to largest fish that differed from the original study. The fundamental information provided here when evaluating Hawaiian grouper conservation status is longevity (valid to 50 years and estimated to 76 years) — no male sampled was <80 cm total length (TL) and younger than 34 years — and age-at-sexual maturity and age-at-sex change, which were indirectly estimated and compared with prior published estimates for this and other groupers. Updated life history parameters (k = 0.078, L∞ = 95.8 cm TL) should be used to improve future management and conservation assessments.
The most common goatfishes in Hawai'i, Mulloidichthys flavolineatus and M. vanicolensis, comprise a unique resource due to their cultural, ecological and biological significance. These species exhibit pulse-type recruitment to nearshore areas during the summer months. Such pulses of juvenile fishes provide prey for pelagic and nearshore fishes and support a popular directed fishery. However, limited scientific information exists on juvenile stages of these fishes, known locally as oama, despite their contribution to coastal ecology and the extensive nearshore fisheries. Here we resolve growth rates, habitat preferences, hatching dates, size and age structure, as well as fishing catch rates based on new recruits in 2014 and 2015. We sampled 257 M. flavolineatus and 204 M. vanicolensis to compare ecological and fisheries characteristics between species and years. Both show strong habitat segregation, with M. vanicolensis found almost exclusively on hard and M. flavolineatus on soft substrates. Oama recruited in anomalously high numbers in 2014, a trend reflected in a higher catch per unit effort. In contrast, 2015 recruits grew faster, were heavier on average and hatched later than during 2014. Both species have calculated hatch dates in March to July, with M. vanicolensis hatching earlier, recruiting earlier and being consistently larger than M. flavolineatus. This baseline information regarding recruitment and early life-history characteristics can enhance management for other data-limited species that comprise a substantial component of nearshore fisheries in Hawai'i.
We address a critical knowledge gap for the Eightbar Grouper, Hyporthodus octofasciatus, one of the largest groupers targeted in mostly tropical, deep-water fisheries, using genetic analyses of mitochondrial DNA (mtDNA; N = 248) and age and growth information (N = 727) from specimens collected from the eastern Indian Ocean to the western central Pacific Ocean. We additionally used a genotype-by-sequencing approach on a subset of these samples (N = 94) to determine the extent of egg and larval dispersal between tropical and temperate regions along the west coast of Australia (WA). Discontinuity in mtDNA and nuclear genomes, and marked variation in growth trajectories and longevity revealed a distinct, non-interbreeding cryptic lineage in WA. The WA lineage appears to be geographically isolated from the oceanic Indo-West Pacific (IP) lineage and precinctive to a single continuous coastline from tropical to temperate waters (covering > 2200 km and ∼20° of latitude). The Hyporthodus phylogeny reveals a possible allopatric speciation scenario with sister lineages endemic to the east (i.e. H. ergastularius) and west (i.e. WA lineage of H. octofasciatus) coasts of Australia, each diverging from a broader IP distributed ancestor (i.e. IP lineage of H. octofasciatus) in the early Pleistocene Epoch. Sustainable management of these species therefore needs to consider their evolutionary distinction, in addition to aspects of their life-history strategies and limited geographic distribution that infers high vulnerability to fishing at relatively low levels of exploitation.
The crimson jobfish Pristipomoides filamentosus (locally known as ‘opakapaka’ in Hawaii) is a deep-water eteline lutjanid that supports important commercial and recreational fisheries in Hawaii and throughout much of the Indo-Pacific region. It is one of the most commonly caught species of the ‘Deep-7’ bottomfish species complex in the commercial bottomfish fishery of Hawaii. However, there are currently no validated estimates of median body sizes at sexual maturity that can be used in a comprehensive evaluation of the species’ stock status within Hawaiian bottomfish fisheries. Herein we provide size-at-maturity estimates for the species in the main Hawaiian Islands: median length at maturity of 40.7-cm fork length (FL; 95% confidence interval (CI) 40.3–41.2cm) for females and 34.3cm FL (95% CI 33.3–35.3cm) for males, with respective equivalent median weight-at-maturity estimates of ~1.17 and ~0.70kg (2.58 and 1.55 lb). We suggest these newly available data be seriously considered in any future evaluations of minimum size regulations, currently set at ~0.45kg (1 lb) regardless of sex, for the species’ fisheries in Hawaii.
The velvet dogfish (Zameus squamulosus) is a wide-ranging species of shark that is captured as bycatch in bottom and pelagic longline fisheries in the Atlantic, Pacific, and Indian oceans from near the surface to depths as great as 2000 m. Here we provide information on the vertical distribution, diet, and reproduction of the velvet dogfish based on examination of 21 specimens captured in Hawaiian longline fisheries. Only females (576-839 mm in total length [TL]) were captured in waters off Hawaii and this finding may indicate sexual segregation for this species. All individuals were captured in epipelagic and pelagic oceanic waters at estimated target hook depths between 24 and 400 m. Stomach and intestinal contents consisted of squid, fish, and shrimp. Females were immature at 576-729 mm TL and mature at 715-839 mm TL. Mature females contained 6-10 uterine eggs and 4-8 embryos. On the basis of results from a pregnant female (715 mm TL), the size of ovarian ova, and the width of the uteri of slightly larger individuals, female maturity was estimated to occur at 715-730 mm TL. No reproductive seasonality was detected; however, our sample size was small. Reproductive data from published records for size of near-term embryos and smallest free-swimming specimens with umbilical scars indicate that size at birth is 245-270 mm TL.
The longevity of blue marlin (Makaira nigricans) remains unresolved. The use of fin spines and sagittal otoliths for age reading has led to unconfirmed longevity estimates near 20–30 years. Age validation has been elusive because large individuals are uncommonly caught, and a technique that can be applied to structures that provide estimates of age was absent. The use of otolith chemical signatures has been limited by sagittal otoliths that are very small — whole otolith mass of adult blue marlin rarely exceeds 10 mg for the largest fish. Recent advances in the detection limits of radiocarbon (14C) with accelerator mass spectrometry — coupled with recently acquired knowledge of marine bomb 14C signals spanning the tropical Pacific Ocean — have led to an opportunity to age blue marlin from small amounts of otolith material. In this study, otoliths from a recently collected 1245 lb (565 kg) female blue marlin at a measured 146-inch (371 cm) lower jaw fork length were analyzed for 14C. Estimated longevity was either 12–21 or 32–44 years on the basis of bomb 14C dating. Using multiple lines of evidence, it was determined that the young age scenario was most likely, with evidence for an age close to 20 years using a series of deductions in the bomb 14C dating method.
Tropical deep-water snappers (Etelinae) support valuable fisheries across the Indo-Pacific, with stock assessments reliant on age-based information in the absence of reliable catch and effort statistics. These long-lived species have been considered notoriously difficult to age. However, nascent developments in ageing protocols, particularly thinner transverse sections of otoliths (∼180–200 µm), are providing improvements in growth zone clarity, interpretation and repeatability of annuli counts. At a recent international workshop, thin sectioned otoliths from three deep-water snappers were read under reflected light by eight fisheries scientists from across the Indo-Pacific, with various levels of fish-ageing experience. Precision and bias were assessed using traditional ageing precision metrics (index of average percent error, IAPE; and coefficient of variation, CV), and a novel approach using multivariate analyses (metric multidimensional scaling, mMDS) based on Euclidean dissimilarity among readers’ counts and subsequent von Bertalanffy (vB) growth parameter estimates. Annuli counts between the primary reader and all other readers were within two for 80% of estimates, with uniform variation across a wide age range for Etelis carbunculus (intraclass correlation coefficient (ICC) = 0.924, n = 20, 3–25 annuli) and Etelis sp. (ICC = 0.933, n = 15, 2–27 annuli). In contrast, annuli counts for Pristipomoides filamentosus (n = 14, 4–49) were less precise (i.e. ICC = 0.835, 66% of counts within two of primary reader) with a bias toward greater variation in younger, pre-maturational life stages (≤5 annuli). Traditionally accepted ageing precision (IAPE ≤ 5.5%, CV ≤ 7.6%) was achieved for each species, but was commensurate with reader experience. The multivariate mMDS ordination was more informative in identifying both distance (i.e. dissimilarity) and direction (i.e. form) of variations in annuli counts and vB growth parameter estimates among readers. The acceptable level of ageing precision and bias achieved among most readers indicated that deep-water snappers can be aged precisely when appropriate otolith preparation methods are used. This study contributes towards ageing protocols that can be used to (i) interrogate bias and precision of age data for stock assessments, and (ii) standardise comparisons of age-based life history characteristics across a species range.
Bluespine unicornfish (Naso unicornis) from Hawaii were aged to >50 years using cross-sectioned sagittal otoliths. Fish length was a poor indicator of age because of rapid and variable early growth, exemplified by fish aged to be 4 years near maximum length. Growth was deterministic with adult ages decoupled from body length. Otolith mass and thickness were evaluated as proxies for age and both were encouraging; thickness explained more variance but mass was easier to measure. An age estimation protocol was validated through ontogeny using bomb radiocarbon (14C) dating. Use of the postbomb 14C decline period from a regional reference chronology enabled age validation of young fish — a novel approach for the Pacific Ocean. A probabilistic procedure for assigning bomb 14C dates (CALIBomb) was used for the first time to determine fish birth years. The age-reading protocol was generally validated, and it was possible to describe length-at-age despite difficulties in counting otolith annuli beyond 30–40 years. Growth curves differed between the sexes, and a four-parameter generalized von Bertalanffy growth function provided the best fit.
Knowledge of both the life history characteristics and catch and effort statistics of exploited deep-water fishes such as snappers and groupers is typically limited. This contributes to increased uncertainty in stock assessments and, depending on the diligence in governance, is more likely to result in either highly conservative or unsustainable management arrangements. Developments, challenges and advances in the methods for obtaining life history data for these species were discussed at a workshop in May 2015. The key points raised included nascent methods for otolith sectioning and interpretation, the need for standardised international ageing protocols and the issues and challenges in characterising reproductive maturation, including during non-spawning months due to resource limitations. Strategic research directions are identified to address knowledge gaps and thus better inform fisheries management.
Growth characteristics of Pristipomoides filamentosus, a deepwater eteline snapper of major economic importance, are incomplete and inconsistent across its geographical range. Early growth rates have been validated using daily increment and length-frequency analyses, but historical estimates of adult growth rates are variable and longevity is unknown. Studies of P. filamentosus in the Hawaiian Islands have cautioned against unjustified estimates of longevity, but 18 years has at times been uncritically assumed as the maximum age. The present study addresses these age, growth, and longevity issues using lead-radium and bomb radiocarbon dating by providing valid age estimates for adult P. filamentosus. Valid length-at-age estimates ranged from approximately 10 years to more than 40 years. These data, together with robust daily increment data, were used to model a fully validated, long-lived life history for P. filamentosus. This study adds to the few existing studies supporting a view that many tropical fishes, particularly deepwater species, can be longer lived than previously surmised.
Fin spine ageing is the most common technique used to estimate age and growth parameters of large pelagic billfishes from the families Istiophoridae and Xiphiidae. The most suitable methods for processing and interpreting these calcified structures for age estimation have not been clearly defined. Methodological differences between unvalidated ageing studies are of particular concern for highly migratory species because multiple researchers in different regions of the world may conduct age estimates on the same species or stock. This review provides a critical overview of the methods used in previous fin spine ageing studies on billfishes and provides recommendations towards the development of a standardized protocol for estimating the age of striped marlin, Kajikia audax and white marlin, Kajikia albida . Three on-going fin spine ageing studies from Australia, Hawaii, and Florida are used to illustrate some of the considerations and difficulties encountered when developing an ageing protocol for highly migratory fish species. Particular areas of concern that may influence age and growth estimates included differences in fin spine selection, sectioning methods, criteria for identifying and measuring annuli, distinguishing false annuli, validation procedures, identification of the first annulus, and methods used to replace annuli lost due to vascularization of the fin spine core.
We verified the age and growth of swordfish (Xiphias gladius) by comparing ages determined from annuli in fin ray sections with daily growth increments in otoliths. Growth of swordfish of exploitable sizes is described on the basis of annuli present in cross sections of the second ray of the first anal fins of 1292 specimens (60-260 cm eye-to-fork length, EFL) caught in the region of the Hawaii-based pelagic longline fishery. The position of the initial fin ray annulus of swordfish was verified for the first time with the use of scanning electron micrographs of presumed daily growth increments present in the otoliths of juveniles. Fish growth through age 7 was validated by marginal increment analysis. Faster growth of females was confirmed, and the standard von Bertalanffy growth model was identified as the most parsimonious for describing growth in length for fish greater than 60 cm EFL. The observed growth of three fish, a year-old in size when first caught and then recaptured from 364 to 1490 days later, is consistent with modeled growth for fish of this size range. Our novel approach to verifying age and growth should increase confidence in conducting an age-structured stock assessment for swordfish in the North Pacific Ocean.
Istiophorid billfishes are notoriously difficult to identify to species, especially early life history stages. Traditional use of morphology and pigment based characters for species separation of larvae can fail when faced with moderate levels of intra-specific phenotypic plasticity. Unfortunately, as the early life history of marine fishes can play an important role in reproductive success and management strategies, we are potentially missing or confounding valuable data by misidentifying specimens. We recently presented a DNA based method of identification for near real-time processing of ichthyoplankton samples. Using this method we have unambiguously identified seven striped marlin larvae from Hawaiian waters. Additional analysis of DNA sequence data shows that these larvae all have unique mitochondrial haplotypes indicating they each had different mothers and were not the result of a single chance event. This represents the first detection of spawning activity in an area that historically has been regarded as only nursery habitat for this species. This finding has important ramifications for fishery management as striped marlin represent a significant portion of the billfish catch in the Hawaii-based longline fishery.
The trace element composition of young‐of‐the‐year (YOY) juvenile swordfish Xiphias gladius sagittal otoliths were analysed as a preliminary test of the value of otolith elemental fingerprints for determining swordfish nursery ground origins in the central Pacific Ocean. A suite of five elements (Mg, Zn, Sr, Ba and Pb) was assayed with isotope dilution ICP‐MS; all elemental concentrations were roughly comparable to otoliths of other marine fishes. Multivariate analyses of elemental fingerprints based on Ba and Sr revealed differences between sample sites, and the magnitude of the differences increased with latitudinal separation. With more comprehensive sampling of nursery grounds, it should be possible to identify origin of nursery ground for adult swordfish by analysing the YOY juvenile portion of the sagittal otolith.