この報告書は、ニューサウスウェールズ州の第一次産業省から Sydney Institute of Marine Science が資金援助を受けたプロジェクト調査の最終報告書という位置づけのもの。
https://www.dpi.nsw.gov.au/fishing/recreational/recreational-fishing-fee/licence-fees-at-work/research-on-fish-and-recreational-fishing
第11回インド―太平洋魚類研究集会（NZ: オークランド）への出席が縁で、著者の一人である Miskiewicz 博士より小西博士に提供されたもので、同報告書の公開についてはMiskiewicz 博士より許可が得られている。
形態に基づく同定と DNA barcoding による分析結果が比較されていて、大変興味深いもので、日本の仔稚魚の同定にも役立ちそうな情報（共通種27種の仔稚魚の画像）が含まれている。
その１：報告書テキスト（英文）https://ameblo.jp/husakasago/entry-12831430972.html
その２：表２と表３（英文）https://ameblo.jp/husakasago/entry-12831436183.html
その３：仔稚魚の写真（日本産種との共通種）https://ameblo.jp/husakasago/entry-12831443049.html
その４：仔稚魚の写真（つづき）https://ameblo.jp/husakasago/entry-12831469885.html
その５：「はじめに」～「考察」の仮和訳
https://ameblo.jp/husakasago/entry-12831566396.html
■参照
■ ■ ■
Larval fish identification of recreationally important species. Final Report to NSW Rec Fishing Trust
Project number SS083 –
Identification of the larvae of recreationally important fish species for long-term monitoring
Final report, October 2023
Prepared for the NSW Recreational Fishing Trust; NSW Department of Primary Industries
Iain M Suthers a,b,*, Noah Baylis a,b, Tony Miskiewicz c, Indiana Riley c, Sharon A. Appleyard d
a Sydney Institute of Marine Science, Mosman, NSW 2088, Australia
b Centre for Marine Science and Innovation, School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington, NSW 2052, Australia
c Australian Museum Research Institute, Australian Museum, Sydney, NSW 2010, Australia.
d CSIRO Australian National Fish Collection, National Research Collections Australia, Hobart, TAS, 7000, Australia
* Corresponding author I.Suthers@unsw.edu.au, Version 6
Frontispiece image of the dusky flathead, Platycephalus fuscus (Neira et al. 1988)
■EXECUTIVE SUMMARY
This report outlines the project outcomes for SS083 – Identification of the larvae of recreationally important fish species for long-term monitoring.
The project had three main objectives:
1) Photograph and/or draw those larvae we have morphologically identified from Port Hacking monitoring station;
2) Genetically identify and/or confirm the species identification of fish larvae of over 60 recreationally important species (including look-alikes that are not recreational), at ~4 different sizes of each, using DNA barcoding of fish larvae from off Port Hacking (at 100 m and 50 m isobaths);
3) Prepare a writen report as a modern supplement to Neira et al. (1998) which aligns the original diagrams with the new photographs and confirmed identifications, with specimens properly stored at the Australian Museum.
The overarching project aim was to align traditional morphology and photographs of larval fish with modern genetics to identify larvae of many recreationally important NSW fish species to aid in revealing spawning habitats and changes in distributions.
This was achieved by integrating DNA barcoding of the COI gene of larval fish with known museum databases, followed by detailing the larval morphological and pigmentation characters.
We completed four plates (n = 95 larvae per plate) for COI barcoding (totalling 380 samples) and all were successfully photographed (Appendix 1), and many specimens remain intact as we only genetically analysed an eyeball.
Aims 1 and 2 were successfully completed within the project; Aim 3 is this report and the Appendix 1 (to support publication in the scientific literature).
Platycephalidae larvae in Plates 1 and 2 formed part of an honour’s thesis by Noah Baylis, 2022, 1st class honours UNSW (found in Appendix 2 as a case study), and is currently being planned for publication.
Other Families identified in these two plates included Family Bregmacerotidae, F. Carangidae, F. Cheilodactylidae, C. Coryphaenidae, F. Gempylidae, F. Monacanthidae, F. Mugillidae and F. Scombridae.
In Plate 1, 85 of the 95 samples (which consisted of small fish larvae or eyeballs from fish larvae – to keep the larval specimen intact for morphology) were genetically barcoded and identified, and 91 samples in Plate 2 were successfully barcoded.
The species level diagnoses for the platycephalid and carangid families are of particular recreational and taxonomic interest, as many of the species identified were previously undescribed at the larval stage.
In Plate 3, 64 of 95 samples were successfully barcoded.
Thirty-one samples had very low input DNA and did not amplify therefore could not be barcoded.
In samples that could be barcoded representatives from 10 Families were identified, with 32 individuals from Family Carangidae represented in the plate
– including Caranx sexfasciatus, Decapterus russelli, Pseudocaranx georgianus, Trachurus declivis and T. novaezelandiae.
Plate 3 included putative larval scad (Carangidae), redfish (Centroberyx affinis), morwong (Chelodactylidae); leatherjackets (Monacanthidae),
and including some confounding taxa of snapper (Lutjanidae), sea bass (Serranidae), a mullet (Gracimugil argenteus), redbait (Emmelichthys nitidus) and ocean perch (Helicolenus percoides).
In Plate 4, there was very good sequencing for 94/95 samples (99%), but for 4 larvae, the morphological identification to family levels was not supported by the barcoding results.
Representatives from 25 Families were identified, including larvae of silver dory (Cyttus australis), blue warehou (Seriolella brama), blue groper (Acherodus viridis) and leatherjacket (Acanthaluteres spilomelanurus).
Plate 4 included flathead (Platycephalidae), flatfishes (flounders, tongue soles and bothids), pomfrets (Brama pauciradiata) and scads (Carangidae); plus some confounding taxa including pomacentrids, nomeids and clingfishes (Gobiesocidae).
Following image quality control and selection of various image views, the COI results from the four plates will be uploaded to the Barcode of Life Data System (BOLD) along with metadata and paired individual species images.
For Aim 3 plates of images for the 90 taxa, mainly to species, that were identified by COI barcoding, are provided in Appendix 1, providing the basis for more refined species identification of larvae future publications on descriptions of larval development series and larval fish based ecological studies.
■INTRODUCTION
Australia has a very diverse fish fauna with over 4,500 species with more than 1,000 species occurring in southern Australian waters.
The majority of larval stages of adult fish species in Australia can only be identified to the taxonomic level of family or genus using morphological characters.
For example, we are confident of identifying larval stages of sardine, anchovy, slimy mackerel, kingfish, yellowtail scad and dusky flathead using our “bible” by Neira et al. (1998).
Larval stages of 124 species, comprising 116 marine and 8 freshwater species from temperate Australian water were identified in Neira et al. (1998) based on morphological characters and pigmentation.
The larvae of many diverse families were not considered at all or only some species within a family in Neira et al. (1998), due to difficulty in separation of larvae of different species within a genus or family that were morphologically similar.
In addition, for example, although described in Neira et al. (1998) it can be difficult to discriminate larval jack mackerel from larval yellow tail scad (Trachurus declivis v T. novaezelandiae) using morphology, especially small or damaged larvae and these are important species for fisheries ecosystems which will likely be impacted by climate change.
Similarly, we are mostly confident we can identify dusky flathead in our coastal and offshore samples, but there are other species, such as tiger flathead, long-spine flathead, blue-spot flathead that have morphologically similar larvae and are currently undescribed.
Ichthyoplankton have been surveyed in Australia since the early 20th century, with a 1910 survey of three species in Port Phillip Bay, and surveys in the 1930–50’s of larvae and eggs of sardine (Sardinops sagax) and anchovy (Engraulis australis).
Descriptions of larvae for various species occurred from the 1950s onwards and in the 1980s surveys of larval fish assemblages began in earnest in Australian temperate marine waters (Smithら2018).
Between late 2014-2022 larval fishes were routinely collected and larval fishes identified in samples collected at five national reference stations (NRS) around Australia including the Port Hacking National Reference Station (NRS).
One challenge with identifying larval fish data is that few scientists have the ability to identify many taxa to species level and given the progressive loss of taxonomic expertise, there is increasing uncertainty as to whether this expertise will extend to a new generation of fisheries scientists.
This suggests that the taxonomic resolution of future surveys of larval fishes could decline without some guidance.
DNA barcoding methods can greatly enhance the identification of ichthyoplankton (Neira et al. 2015), and potentially reduce the reliance on taxonomic experts; but for generating larval fish count data from large surveys, DNA methods are complementary to morphological identification methods.
These days, many fish museums around the world genetically confirm and identify fishes or sharks with DNA barcoding alongside morphological or alpha taxonomy and imaging of the barcoded specimen for future reference.
For fishes, the mitochondrial gene COI, and a world-wide database called BOLD (Barcode of Life System) (Herbertら 2003) is used for genetic identification.
Integrated taxonomy using COI barcoding is now a routine process for taxonomists.
Locally, target species that are recreationally important, which could occur at the Port Hacking IMOS National Reference Station and would benefit from genetic identification as larval stages include representatives from:
Family Platycephalidae – at least six flathead species are known in the area although larval individuals have not been identified from:
• Platycephalus fuscus (Dusky flathead) – identified in Neira et al. (1998), but not genetically confirmed at the NRS.
• Platycephalus caeruleopunctatus (Bluespot flathead) – undescribed larvae for NSW.
• Platycephalus grandispinis (Longspine flathead) – undescribed larvae for NSW.
• Platycephalus richardsoni (Tiger flathead) – undescribed larvae for NSW.
• Platycephalus marmoratus (marbled flathead) – undescribed larvae for NSW.
• Ambiserrula jugosa (Mud flathead) – undescribed larvae for NSW.
• Flathead larvae that are not Platycephalus fuscus are regularly found in Port Hacking NRS samples.
• Platycephalus bassenis (Sand flathead) – larvae described from Tasmanian waters by Jordan (2001).
Family Carangidae – a complex family, and yellowtail kingfish larvae may look like amberjack
• Trachurus novaezelandiae - identified in Neira et al. (1998), but larvae not genetically confirmed.
• Trachurus declivis - identified in Neira et al. (1998), and their eggs and larvae occur off Sydney during winter months (Neira et al., 2015).
• Seriola sp. – only identified to genus in Neira et al. (1998) (yellowtail kingfish S. lalandi, samsonfish S. hippos or amberjack S. dumerili).
• Caranx sp.– undescribed larvae for Australia.
• Decapturus spp. – undescribed larvae for Australia.
• Pseudocaranx georgianus (trevally) - identified in Neira et al. (1998), but larvae not genetically confirmed.
Family Cheilodactylidae - recognised larvae as morwongs, however larval forms have not been identified to species in NSW.
• multiple species in Port Hacking samples assumed to be red morwong (Cheilodactylus fuscus), blue morwong (Nemadactylus douglasii) or jackass morwong (Nemadactylus macroptera-) identified in Neira et al. (1998), but larvae not genetically confirmed.
Family Monocanthidae – recognised larvae as leatherjacket but larval forms unknown to species.
Family Labridae - recognised larvae as wrasses, but larval forms unknown to species
• Blue groper (Achoerodus viridis) and many other wrasses.
Family Scorpaenidae - recognised larvae as scorpionfish but larval forms of majority of scorpenid larvae unknown to species.
• Larvae of ocean perch (Helicolenus percoides) identified in Neira et al. (1998), but larvae not genetically confirmed.
Family Epinephilidae - recognised larvae as grouper and rockcod, but larval forms unknown to species.
Family Scombridae
• Scomber australicus – blue mackerel (slimy mackeral) identified in Neira et al. (1998)
• other species of mackerel or tuna in samples especially at North Stradbroke Island NRS.
Family Mugilidae
• possibly multiple species of mullet in samples (sea mullet (Mugil cephalus), sand (Myxus elongatus), flat tail (Gracilimugil argentea).
The aims of this study were to:
1) Photograph and/or draw those larvae we have morphologically identified from the Port Hacking monitoring station;
2) Genetically identify and/or confirm the species identification of fish larvae of over 60 recreationally important species (including look-alikes that are not recreational), at ~4 different sizes of each, using DNA barcoding of fish larvae from off Port Hacking (at 100 m and 50 m isobaths);
3) Prepare a writen report as a modern supplement to Neira et al. (1998) which aligns the original diagrams with the new photographs and confirmed identifications, with specimens properly stored at the Australian Museum.
■METHODS
■■Selection of larvae for barcoding
The primary requirement for selection of fish larvae for COI barcoding is that the larvae were initially fixed in the field in 95% ethanol.
Other criteria for selection were the condition of each larva and the availability of a size range of individuals of different taxa.
The focus of the study was samples from the IMOS-National Reference Station (NRS) sampling station at Port Hacking station.
However, many recreationally and commercially important species occur in NSW are distributed along the east coast of Australia.
Therefore, to increase the range of taxa that were barcoded and the size range of individual taxa of interest, ethanol fixed larval fish were selected from samples collected at the North Stradbroke Island and Maria Island NRS (Table 1, sampling details below) and from larval fish samples collected in Port Phillip Bay (Forbertら 2019) and samples collected in northern NSW waters on a RV Investigator (Garciaら2022).
In the laboratory, samples were sorted with the aid of a dissecting microscope and larvae were transferred from initial ethanol or formalin solutions to 70% ethanol in the case of initially formalin-fixed specimens, while ethanol-fixed samples were transferred to 80% ethanol solution and stored at –20°C to facilitate DNA barcoding (Appleyardら 2021).
Larvae selected for barcoding were photographed using a Leica z-stacking microscope at the Australian Museum prior to extraction of one eyeball for barcoding.
■■Study region and IMOS sampling program
The Integrated Marine Observing System (IMOS) is a national ocean observation program that monitors oceanographic, climatic, and biological data at several National Reference Stations (NRS) around Australia (Lynchら 2014).
The larval fish used in this study were sourced from three NRS sampling sites under the IMOS Larval Fish Monitoring Program (IMOSLFMP):
North Stradbroke Island near Brisbane, Queensland (27.35°S, 153.56°E), Port Hacking near Sydney, New South Wales (34.12°S, 151.23°E) and Maria Island near Hobart, Tasmania (42.56°S, 148.22°E)
The region is dominated by the poleward flow of the East Australian Current (EAC), transporting warmer, oligotrophic waters from tropical regions such as the Coral Sea to southern waters (Okeら 2019).
■■Ichthyoplankton sampling at three east coast NRS
Samples were collected at approximately monthly intervals at the three NRS for 7 years from September 2014–December 2021.
Two towed samples were collected at each NRS per month and preserved in either ethanol or formalin.
Tows were taken using either a ring net with an 85–cm diameter and 500 μm mesh, or a 75–by–75 cm square net 500 μm mesh from 2014–2018.
From 2019, these nets were replaced with a 60 cm diameter bongo net (500 μm mesh), which has a similar total mouth area, and with the same mesh.
Nets were towed for ~12 minutes at ~3 knots (1.5 metres per second) between 25 metres depth to the surface.
Nets were equipped with a General Oceanics or a TSK flow meter to determine the volume of water filtered and subsequently enable standardisation of larval abundances, and depth of the net was monitored where possible with a HOBO depth logger.
Samples were collected from the nets and fixed in 95% ethanol for COI barcoding or 4% (neutral buffered) formalin.
■■DNA extraction and COI barcoding
For most of the larvae, an eyeball was removed and placed in an individual 96-well microplate well.
Where larvae were too small for eyeball extraction, the entire specimen was placed in the plate wells.
Microplates were sealed with strip caps and sent to the CSIRO marine laboratories in Hobart where the plates were frozen at -20°C until DNA extraction.
DNA extraction was undertaken using a modified small volume Wizard® SV Genomic DNA Purification system (Promega, Australia) which was based on 1/5 volume reduction from that used for adult fish tissues (see Appleyardら 2018; 2021) with digestions and extractions undertaken in the 96-well plates, where total volume capacity of each well was 200 μl.
Plates were always capped with strip caps.
DNAs were stored at 4°C overnight, with a sub-aliquot of each DNA quantified on a Nanodrop 8000 (Thermofisher, USA).
Aliquots of DNA from the plates are also archived at -80°C in the CSIRO marine laboratories.
Aliquots of DNA were sent to the Ramacioti Centre for Genomics (University of New Wales, Sydney) where PCR amplifications and COI Sanger sequencing (using the BCL and BCH primers) (Baldwinら, 2009) were undertaken.
The resulting forward and reverse COI sequences for the extracted samples were sent to CSIRO where the forward and reverse sequences were trimmed, de novo assembled and assemblies were manually checked by eye for base pair calling accuracy and converted into consensus sequences using Geneious Prime® 2021.2.2 (Biomaters Ltd, New Zealand).
The consensus sequences were then compared to the sequences available on the international Barcode of Life Data System (BOLD) (Centre for Biodiversity Genomics, Canada).
The results were tabulated and denoted with the correct species match at a pairwise identity level of >98%, based on the similarity of the observed and known sequence.
Species occurrence was also confirmed in the Australian Faunal Directory.
Where the identity of multiple larval samples per species and or genera were required, genetic UPGMA trees (see Figure 1 for an example) were also used for taxa confirmation.
Based on outcomes, genetic species identifications were assigned to each sample and genetic identifications were checked against the morphology of each specimen.
Successfully sequenced samples, larval images and their metadata will be submited to BOLD into the publicly accessible project FIMOS IMOS Australian Larval Fish Monitoring.
Figure 1 An example of the larval trevally, scad (Family Carangidae) consensus tree diagram showing 3 groupings of different genera and species from our larval unknowns (NSW Rec. Fishing Trust well # of the 95 well plate), in relation to known mitochondrial COI sequences of adult Trachurus novaezelandiae, T. declivis, and Decapturus russelli from BOLD.
■RESULTS
COI barcoding results across each of the four plates in shown in Table 2.
Sequencing success in Plates 1, 2 and 4 were high, while two-thirds of samples in Plate 3 were successfully sequenced.
Overall larvae of 39 families and 90 taxa, majority to species, were identified using COI barcoding (Table 3).
The diverse families in the four plates were larvae of Platycephalidae (9 species), Carangidae (11) and Monacanthidae (9).
Plates 1 & 2: Prior to DNA barcoding, the 190 larvae in the plates were morphologically identified with the level of identification varying from Family to Species (e.g., Ammodytidae; Platycephalus caeruleopunctatus).
DNA barcoding identified Ambiserrula jugosa, Onigocia pedimacula, Platycephalus bassensis, Platycephalus caeruleopunctatus, Platycephalus endrachtensis, Platycephalus fuscus, Platycephalus grandispinis, Rogadius mcgroutheri, Sorsogona tuberculata and Ratabulus diversidens (Table 1).
Table 1: Summary of larval flathead taxa identified at the 3 NRS stations, showing total abundance and mean concentration per 100 m3 (within brackets).
Other Families identified in these two plates included Family Bregmacerotidae, F. Carangidae, F. Cheilodactylidae, C. Coryphaenidae, F. Gempylidae, F. Monacanthidae, F. Mugillidae and F. Scombridae.
In Plate 1, 85 of the 95 samples (which consisted of small whole larval fish or eyeballs from fish larvae) were genetically barcoded and identified, and 91 samples in Plate 2 were successfully barcoded.
Plate 3: 64 of 95 samples were successfully barcoded.
Thirty-one samples had very low input DNA and did not amplify therefore could not be barcoded.
Representatives from ten Families were identified, with 32 individuals from Family Carangidae represented in the plate – including from Caranx sexfasciatus, Decapterus russelli, Pseudocaranx georgianus, Trachurus declivis and T. novaezelandiae.
In Plate 4, there was very good sequencing for 94/95 samples (99%), but for 4 larvae, the morphological identification to the family level of Gobieosocidae was not supported by the barcoding results.
Representatives from 25 Families were identified, including larvae of silver dory (Cyttus australis), blue warehou (Seriolella brama), blue groper (Acherodus viridis) and leatherjacket (Acanthaluteres spilomelanurus).
■DISCUSSION
Previously only ~10% of the ~1000 species of fish (of the larval stages) in southern Australia could be identified to species with a microscope (i.e. morphologically); the other ~90% could only be classified to family or genus (Neira et al. 1998).
Furthermore, for even those known species there were uncertainties as there were other members of the family or genus that may have been confused, especially for the newly hatched larvae.
Therefore, without deploying accurate identification tools, we are unable to precisely understand spawning distributions and the effects of coastal currents, winds and climate regimes on the supply of young fish (Schillingら 2022).
Using a combination of traditional larval morphology and DNA barcoding, this study nearly doubled the number of Australian species whose larvae can be identified and confirms our current (morphological) understanding is sound (Neira et al. 1998).
In particular, this study resolves the taxonomic problems of three important families of fish for our region – the trevallies and scads (Carangidae); the flatheads (Platycephalidae) and the leatherjackets (Monacanthidae). The species level diagnoses for the platycephalid and carangid families are particularly of recreational and taxonomic interest, as many of the species identified were undescribed at the larval stage.
For flatheads, the seasonality of reproduction and spatial distribution off eastern Australia was revealed.
Features of larval A. jugosa, O. pedimacula, P. caeruleopunctatus, P. endrachtensis, P. grandispinis, P. richardsoni, R. mcgroutheri, S. tuberculata and representatives of genus Ratabulus are detailed for the first time to accompany existing descriptions of P. bassensis by Jordan (2001) and P. fuscus by Neira et al. (1998).
From a total of 2563 larval platycephalids, 98% of these were successfully identified to species, with most of the remainder identified as genus Ratabulus spp.
Several differences in spawning habits and seasonality of flathead were observed, and the differences in larval abundance were largely related to seasonal variations in water temperatures.
Larval flathead species diversity decreased with increasing latitude, from 7 species and 6 genera present off North Stradbroke Island, to 5 species and 2 genera at Port Hacking, to 2 species of Platycephalus at Maria Island.
This identification of flathead larvae will enable more targeted species-specific stock assessment and management options, by providing inputs into the management and or remediation of the impacts of climate change and overfishing.
Our genetic (COI) barcoding has separated out morphologically similar scads and trevally (Family Carangidae), such as Trachurus, Decapterus and Pseudocaranx larvae to genus and species levels.
Many of these larvae had been initially mis-classified based on morphological traits.
Four Carangidae species were identified from Port Hacking – the Jack mackerel (Trachurus declivis), Yellowtail horse mackerel (Trachurus novaezelandiae), Silver Trevally (Pseudocaranx georgianus), and Yellowtail kingfish (Seriola lalandi).
Both mackerel species and the Yellowtail kingfish were also identified at North Stradbroke Island NRS, as well as the Indian scad (Decapterus russelli) and the Shortfin scad (Decapterus macrosoma).
Both these Decapterus species were undescribed in their larval stages and initial morphological identifications were otien confused with Trachurus.
This study complements our work on the high concentrations of tuna larvae found off North Stradbroke Island, which we resolved by morphology and COI barcoding with the majority of larvae being Auxis rochei and small numbers of A. thazard (bullet tuna and frigate mackerel) (Caoら in preparation).
We also confirmed the identification and presence off eastern Australia identified Acanthocybium solandri (wahoo), Allothunnus fallai (slender tuna), Auxis rochei (bullet tuna), Auxis thazard (frigate mackerel), Euthynnus affinis (mackerel tuna), Katsuwonus pelamis (skipjack tuna), Sarda australis (Australian bonito), Thunnus albacares (yellowfin tuna) and Thunnus tonggol (longtail tuna).
This project is exciting as with this knowledge we can examine seasonal and long-term changes in reproduction by many more temperate Australian fish.
We could now re-examine archived samples of larval fish at the Australian Museum, including those that were collected by IMOS at the five National Reference Stations around Australia from 2014-2022.
This project is also exciting, as it underpins the future of monitoring fish reproduction using DNA barcoding.
The increasing information in the Barcode of Life Database (BOLD) supports modern identification and has transformed the crisis in modern taxonomy, with the retirement of so much expertise.
It is now technically possible to examine the ethanol preservative solution of a plankton tow, for the presence of unique sequences of particular genes such as the mitochondrial COI (e.g. Goldら 2021, 2023; Appleyard pers. comm.).
Furthermore, we can now obtain COI sequences from formalin preserved larvae, up to 6 months in routine formalin preservation (Appleyardら 2021), overcoming a significant technical obstacle.
■ACKNOWLEDGEMENTS
We thank the NSW Recreational Fishing Trust for funding this investigation; and the Integrated Marine Observing System (IMOS) for enabling the sampling of larval fishes.
We thank honours students Clare Cao and Noah Baylis for their contributions to the project.
We also thank the CSIRO Australian National Fish Collection (ANFC), for their extensive effort in adult fish curation and COI barcoding; without which our larval barcoding and integrated identification processes would not be possible.
■REFERENCES
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Oke, P. R., M. Roughan, P. Cetina-Heredia, G. S. Piloa, K. R. Ridgway, T. Rykova, M. R. Archer, R. C. Coleman, C. G. Kerry, C. Rocha, A. Schaeffer, and E. Vitarelli (2019), Revisiting the circulation of the East Australian Current: Its path, separation, and eddy field. Prog. Oceanogr. 176:102139.
Smith, J. A., Miskiewicz, A. G., Beckley, L.E., Everet, J. D., Garcia, V., Gray, C. A., Holliday, D., Jordan, A.R., Keane, J., Lara-Lopez, A. and Leis, J.M. (2018), A database of marine larval fish assemblages in Australian temperate and subtropical waters. Scientific Data 5: 180207.
Schilling, H. T., Hinchliffe, C., Gillson, J. Miskiewicz, A. G., and Suthers, I. M. (2022), Coastal winds and larval fish abundance indicate a recruitment mechanism for southeast Australian estuarine fisheries. Fisheries Oceanography 31: 40-55
https://onlinelibrary.wiley.com/doi/abs/10.1111/fog.12561
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文献リストに載っていない「Neira et al., 2015 」はこの論文かもしれない。
Francisco J. Neira, Robert A. Perry, Christopher P. Burridge, Jeremy M. Lyle, and John P. Keane (2015). Molecular discrimination of shelf-spawned eggs of two co-occurring Trachurus spp. (Carangidae) in southeastern Australia: a key step to future egg-based biomass estimates. ICES Journal of Marine Science, 72(2), 614–624.
