Campbell R. Davies

Spatial and Sex-Specific Variation in Growth of Albacore Tuna (Thunnus alalunga) across the South Pacific Ocean

Spatial variation in growth is a common feature of demersal fish populations which often exist as discrete adult sub-populations linked by a pelagic larval stage. However, it remains unclear whether variation in growth occurs at similar spatial scales for populations of highly migratory pelagic species, such as tuna. We examined spatial variation in growth of albacore Thunnus alalunga across 90° of longitude in the South Pacific Ocean from the east coast of Australia to the Pitcairn Islands. Using length-at-age data from a validated ageing method we found evidence for significant variation in length-at-age and growth parameters (L ∞ and k) between sexes and across longitudes. Growth trajectories were similar between sexes up until four years of age, after which the length-at-age for males was, on average, greater than that for females. Males reached an average maximum size more than 8 cm larger than females. Length-at-age and growth parameters were consistently greater at more easterly longitudes than at westerly longitudes for both females and males. Our results provide strong evidence that finer spatial structure exists within the South Pacific albacore stock and raises the question of whether the scale of their “highly migratory” nature should be re-assessed. Future stock assessment models for South Pacific albacore should consider sex-specific growth curves and spatial variation in growth within the stock.

Reproductive Dynamics and Potential Annual Fecundity of South Pacific Albacore Tuna (Thunnus alalunga)

The reproductive biology of albacore tuna, Thunnus alalunga, in the South Pacific Ocean was investigated with samples collected during broad-scale sampling between 2006 and 2011. Histology was done in a single laboratory according to standard protocols and the data analysed using generalized linear mixed-effects models. The sex ratio of albacore was female biased for fish smaller than approximately 60 cm FL and between 85 and 95 cm, and progressively more male biased above 95 cm FL. Spawning activity was synchronised across the region between 10°S and 25°S during the austral spring and summer where sea surface temperatures were ≥24 °C. The average gonad index varied among regions, with fish in easterly longitudes having heavier gonads for their size than fish in westerly longitudes. Albacore, while capable of spawning daily, on average spawn every 1.3 days during the peak spawning months of October to December. Spawning occurs around midnight and the early hours of the morning. Regional variation in spawning frequency and batch fecundity were not significant. The proportion of active females and the spawning fraction increased with length and age, and mature small and young fish were less active at either end of the spawning season than larger, older fish. Batch fecundity estimates ranged from 0.26 to 2.83 million oocytes with a mean relative batch fecundity of 64.4 oocytes per gram of body weight. Predicted batch fecundity and potential annual fecundity increased with both length and age. This extensive set of reproductive parameter estimates provides many of the first quantitative estimates for this population and will substantially improve the quality of biological inputs to the stock assessment for South Pacific albacore.

Regional patterns in reproductive biology of Lethrinus miniatus on the Great Barrier Reef

Uniformity in fish population biology is a common assumption in many fishery assessments and man- agement arrangements. Although spatial patterns in population biology are often unknown, ignorance of significant variation within a fished stock has profound implications for fishery assessments and management. In the current paper, the reproductive biology of an exploited reef fish, Lethrinus miniatus, was examined for populations in the northern and southern regions of the Great Barrier Reef (GBR). Significant regional variation was observed in some reproductive parameters, but not others. In the northern region, the proportion of spawning females observed during the spawning season was significantly greater and the size at sex change significantly smaller than in the southern region. However, the spawning season, age at sex change and sex ratios did not differ significantly between the northern and southern regions. Size and age at maturity could not be estimated from either of these regions, but an estimate was obtained from the Capricorn-Bunker region at the southern tip of the GBR. The observed regional patterns in reproductive biology of L. miniatus populations have important implications for specific management arrangements such as size limits and seasonal closures, and for fisheries management in general.

Variation in the periodicity and timing of increment formation in red throat emperor (Lethrinus miniatus) otoliths

Marginal increment analysis was used to validate the periodicity of opaque increment formation in red throat emperor (Lethrinus miniatus) otoliths and to test for variation in the periodicity and timing of increment formation across age classes, years and regions of the Great Barrier Reef (GBR). Marginal increment analysis demonstrated that opaque increments were deposited on an annual basis, a result supported by a preliminary experiment on a chemically tagged captive fish, although significant regional and temporal variation in the timing of increment formation was observed. Marginal increment analysis revealed a clear annual periodicity in increment formation for all age classes in the southern region but a more ambiguous pattern for most age classes in the northern region of the species distribution on the GBR. Opaque increments appeared to form 1 month earlier in the southern region than in the northern region and, in the southern region, formed 2 months earlier in 1999 than in 2000. No significant variation was observed in the timing of increment formation between age classes. Based on knowledge of the timing of otolith increment formation and spawning season of L. miniatus, an algorithm was developed for the estimation of age in months, irrespective of the time or location of capture.

Spatial variation in the effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus.

The effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus populations were compared between coral reefs open or closed (no-take marine reserves) to fishing and among four geographic regions of the Great Barrier Reef (GBR), Australia. The specific reproductive metrics investigated were the sex ratio, the proportion of vitellogenic females and the spawning fraction of local populations. Sex ratios became increasingly male biased with length and age, as expected for a protogyne, but were more male biased in southern regions of the GBR (Mackay and Storm Cay) than in northern regions (Lizard Island and Townsville) across all lengths and ages. The proportion of vitellogenic females also increased with length and age. Female P. leopardus were capable of daily spawning during the spawning season, but on average spawned every 4·3 days. Mature females spawned most frequently on Townsville reserve reefs (every 2·3 days) and Lizard Island fished reefs (every 3·2 days). Females on Mackay reefs open to fishing showed no evidence of spawning over 4 years of sampling, while females on reserve reefs spawned only once every 2-3 months. No effect of length on spawning frequency was detected. Spawning frequency increased with age on Lizard Island fished reefs, declined with age on Storm Cay fished reefs, and declined with age on reserve reefs in all regions. It is hypothesized that the variation in P. leopardus sex ratios and spawning frequency among GBR regions is primarily driven by water temperature, while no-take management zones influence spawning frequency depending on the region in which the reserve is located. Male bias and lack of spawning activity on southern GBR, where densities of adult P. leopardus are highest, suggest that recruits may be supplied from central or northern GBR. Significant regional variation in reproductive traits suggests that a regional approach to management of P. leopardus is appropriate and highlights the need for considering spatial variation in reproduction where reserves are used as fishery or conservation management tools.

Demography of a large exploited grouper, Plectropomus laevis: Implications for fisheries management

Species in the coral trout complex Plectropomus spp. are some of the most desired and exploited in the Indo-Pacific, although data are limited for most species. Demographic parameters of blue-spot coral trout, Plectropomus laevis, were estimated on the basis of specimens collected from five regions of the Great Barrier Reef and Torres Strait, Australia, between 1995 and 2005 to define demography and population status over a broad range. In total, 1249 individuals were collected to estimate growth, mortality and reproductive parameters. Growth parameters based on the von Bertalanffy model were K = 0.096 year–1, L∞ = 1159 mm fork length (FL) and the best-fit model suggested non-asymptotic growth over the size and age ranges represented in the sample. Maximum age was 16 years, total mortality was estimated at 0.39 (±0.031), and estimated size and age at first maturity were 299 mm FL (range: 299–872) and at 1 year (range: 1–6.5), respectively, with the size range over which sex change occurred at 460–872 mm FL. Population parameters and available relative abundance estimates indicated that long-term sustainable harvest rates of P. laevis will be lower than those for P. leopardus and species-specific size-based management measures are necessary to ensure the species is not overfished.

Age estimation and validation for South Pacific albacore Thunnus alalunga

Validated estimates of age are presented for albacore Thunnus alalunga, sampled from a large part of the south-western Pacific Ocean, based on counts of annual opaque growth zones from transverse sections of otoliths. Counts of daily increments were used to estimate the location of the first opaque growth zone, which was completed before the first assumed birthday. The periodicity of opaque zones was estimated by marginal increment analysis and an oxytetracycline mark-recapture experiment. Both validation methods indicated that opaque zones formed over the austral summer and were completed by autumn to winter (April to August). The direct comparison of age estimates obtained from otoliths and dorsal-fin spines of the same fish indicated bias, which was assumed to be due to poor increment clarity and resorption of early growth zones in spines, resulting in imprecise age estimates. As such, age estimates from otoliths are considered to be more accurate than those from spines for T. alalunga. This is consistent with results for a growing number of tropical and temperate tuna Thunnini species. It is recommend that validated counts of annual growth zones from sectioned otoliths is used as the preferred method for estimating age-based parameters for assessment and management advice for these important stocks.

Comparative biology of tropical Lethrinus species (Lethrinidae): challenges for multi‐species management

Life-history characteristics of six tropical Lethrinus species sampled from the Great Barrier Reef World Heritage Area were compared. Two species groups were identified based on fork length (LF ): large species with maximum LF > 640 mm (longface emperor Lethrinus olivaceus, yellowlip emperor Lethrinus xanthochilus and spangled emperor Lethrinus nebulosus) and small species with maximum LF < 480 mm (Pacific yellowtail emperor Lethrinus atkinsoni, pink ear emperor Lethrinus lentjan and ornate emperor Lethrinus ornatus). Lifespan was not correlated with LF . Early growth for all species was rapid and similar during the first few years of life, but coefficients of the von Bertalanffy growth function varied considerably among species. Growth also differed between sexes for L. atkinsoni. Reproductive characteristics varied among species, with peak periods of spawning occurring in November to December for L. atkinsoni, July to August for L. nebulous, September to October for L. olivaceus and a protracted season for L. lentjan, although fewer samples were available for the last two species. Sex-specific LF and age distributions and gonad histology of L. lentjan were suggestive of a functional protogynous reproductive pattern, as observed in other lethrinids. Gonad histology indicated non-functional protogynous hermaphroditism for L. atkinsoni and L. nebulosus. The diversity of life histories among these closely related species emphasizes the difficulty in devising single management strategies appropriate for multi-species fisheries and illustrates the importance of understanding species-specific life histories to infer responses to exploitation.

Demographic Structure, Sex Ratio and Growth Rates of Southern Bluefin Tuna (Thunnus maccoyii) on the Spawning Ground

The demographics of the southern bluefin tuna (SBT) Thunnus maccoyii spawning stock were examined through a large-scale monitoring program of the Indonesian longline catch on the spawning ground between 1995 and 2012. The size and age structure of the spawning population has undergone significant changes since monitoring began. There has been a reduction in the relative abundance of larger/older SBT in the catch since the early 2000s, and a corresponding decrease in mean length and age, but there was no evidence of a significant truncation of the age distribution. Pulses of young SBT appear in the catches in the early- and mid-2000s and may be the first evidence of increased recruitment into the spawning stock since 1995. Fish in these two recruitment pulses were spawned around 1991 and 1997. Size-related variations in sex ratio were also observed with female bias for fish less than 170 cm FL and male bias for fish greater than 170 cm FL. This trend of increasing proportion of males with size above 170 cm FL is likely to be related to sexual dimorphism in growth rates as male length-at-age is greater than that for females after age 10 years. Mean length-at-age of fish aged 8–10 years was greater for both males and females on the spawning ground than off the spawning ground, suggesting that size may be the dominant factor determining timing of maturation in SBT. In addition to these direct results, the data and samples from this program have been central to the assessment and management of this internationally harvested stock.

Maturity Ogives for South Pacific Albacore Tuna (Thunnus alalunga) That Account for Spatial and Seasonal Variation in the Distributions of Mature and Immature Fish

Length and age at maturity are important life history parameters for estimating spawning stock biomass and reproductive potential of fish stocks. Bias in estimates of size and age at maturity can arise when disparate distributions of mature and immature fish within a population are not accounted for in the analysis. Here we investigate the spatial and temporal variability in observed size and age at maturity of female albacore tuna, Thunnus alalunga, using samples collected across the South Pacific. Maturity status was identified using consistent histological criteria that were precise enough to allow for mature but regenerating females to be distinguished from immature females during the non-spawning season, permitting year-round sampling for maturity estimation in albacore. Using generalised linear mixed models, we found that the proportion of mature females at length varied significantly with latitude and time of year. Specifically, females at northern latitudes (∼10–20°S, where spawning occurs) were mature at significantly smaller lengths and ages than females at southern latitudes (∼20–40°S), particularly during the spawning season (October–March). This variation was due to different geographic distributions of mature and immature fish during the year. We present a method for estimating an unbiased maturity ogive that takes into account the latitudinal variation in proportion mature at length during a given season (spawning or non-spawning). Applying this method to albacore samples from the western region of the South Pacific gave a predicted length at 50% mature of ∼87 cm fork length (4.5 years).