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Population Genetics of Selected Whitebait Species: Galaxias Maculatus (Jenyns) and Lovettia Sealii (Johnston)

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Population Genetics of Selected Whitebait Species: Galaxias Maculatus (Jenyns) and Lovettia Sealii (Johnston) POPULATION GENETICS OF SELECTED WIITTEBAIT SPECIES: GALAX/AS MACULATUS (JENYNS) AND LOVETT/A SEALII (JOHNSTON). N.C. PAVUK BSc (Tas) Being a thesis submitted in fulfilment of the degree of Master of Science at the University of Tasmania June 1994 Declaration I hereby declare that this thesis contains no material which has been accepted for the award of any degree or diploma in any university and that, to the best of my knowledge, this thesis contains no copy of previously published material except where due reference is made in the text. N.C. Pavuk . ' Acknowledgments Many thanks and much love to my husband, Andrew. Thank you for all the guidance and support from my Supervisors, Dr Robert White and Mr Wayne Fulton together with all the research staff and inspectors of the Tasmanian Inland Fisheries Commission especially Stuart Chilcott and Brett Davies. Mr Les and Arthur Simms for their help in sample collection and insight into the fishery's history. Jenny Ovenden and Wayne Kelly for their help and good humour in the laboratory. Fisheries staff in New Zealand, Western Australia and Victoria who kindly provided specimens of Galaxias maculatus. Dr Mark Adams many thanks for your help and hints in setting up the electrophoretic studies and in particular for making available early drafts of your text on electrophoresis. UNSW staff for the G-test package and use of the computer. Lynne Woodburn from Latrobe University for help in analysis and for sharing the Genestats package. To all that provided hints and help along the way, Many Thanks. u.. Summary Population Genetics of Selected Whitebait Species: Galaxias maculatus (Jenyns) and Lovettia sealii (Johnston). Populations of diadromous G. maculatus and anadromous L. sealii fishes have been studied for genetic variation by cellulose acetate electrophoresis. L. sealii is a small member of the family Aplochitonidae endemic to Tasmania. In contrast, G. maculatus a salmoniform fish of the Family Galaxiidae, is found throughout the southern temperate zone (Australia, New Zealand and South America). Together these species are the predominant components of Tasmanian whitebait. A significant commercial fishery for Tasmanian whitebait began in 1941 and rapidly declined after 1947 probably due to overfishing (Blackburn, 1950), until its closure in 1974. Encouraging runs of whitebait in the early 1980s prompted a major study to investigate the potential yield and present status of this resource. The lack of data regarding the genetic structure of the species prompted the following study. Eight populations of G. maculatus extending from Western Australia to New Zealand were investigated. Twenty seven samples of L. sealii representing thirteen different river populations from Tasmania were included to capture genetic data both within and between three successive spawning seasons. An analysis of the gene frequency data using Nei's genetic distance (D) and identity (I) , G-test and F-statistics indicated population substructuring in the two species. Morphometric and meristic analysis of L. sealii was also undertaken and supported the genetic data. Within G. maculatus, the Western Australian population was genetically distinct primarily based on allele frequencies observed at the ADH locus. Three genetic pools of G. maculatus were identified which have zoo-geographic implications for this species. In particular they appear to confirm the potential of marine larval dispersal between Australia and New Zealand. A clear genetic discontinuity was observed between north and south Tasmanian L. sealii primarily based on the allele frequency data at the PEPD locus. These fmdings were consistent with those of Blackburn (1950) based on differences in growth rate and pigmentation. However, many southern rivers were found to carry unique stocks. A major contributing factor to this genetic diversity is thought to be the prevailing local hydrographic conditions. The results of this study have implications for the management of the recreational whitebait fishery which has re-opened in Tasmania and possibly the commercial whitebait fishery in New Zealand which is dominated by G. maculatus. iii Table of Contents Page Declaration Summary Acknow ledgrnents iii 1. 0 Introduction 1 1. 1 General Aims 1 1.2 Lovettia sealii 1 1.2.1 Taxonomic Background 1 1. 2. 2 Life History 2 1. 3 Gala.xias maculatus 4 1. 3. 1 Distribution 4 1.3.2 Life History 4 1. 3. 3 An Evolutionary Perspective of the Galaxiidae 5 1.4 History of the Whitebait Fishery 6 1.4.1 Species composition of Whitebait 7 1.5 Stock Identity and Population Genetics 8 1. 5 .1 Protein Electrophoresis 8 1.5 .2 Population Models 9 1. 5. 3. Stock Concept 10 2.0 Materials and Methods 11 2.1 Pilot Study 11 2.2 Main Study 13 2.2.1 Field Collection 13 2.2.2 Collection Method 14 2.2.3 Sample Storage and Preparation 15 2.2.4 Cellulose Acetate Gel Electrophoresis and Histochemical Staining 15 2.2.5 Double Staining 16 2. 3 Analytical Procedures 16 2. 3 .1 Genetic Nomenclature 16 2.3.2 Genetic Data Analysis 16 2.4 Morphometric and Meristic Data Analysis 17 2.4.1 Canonical variance Analysis 18 3.0 Results Population genetics of L. sealii 19 3 .1 Pilot study 19 3. 2 Main study 20 3.2.1 Allele Frequencies 20 3.2.2 Allele Frequency Analysis 20 3.2.3 Analysis of Random Mating 20 3.2.4 F-statistics 29 3.2.5 G81 Analysis 29 3.2.5.1 GST analysis of pooled data 29 3.2.5.2 G-statistics between samples 29 3.2.6 Nei's genetic distance and identity 44 3.2.7 Morphometric and Meristic Analysis 44 3.2.7 .1 Meristics 44 3.2.7.2 Canonical variance analysis 44 Population Genetics of G. maculatus 3. 3 Pilot study 48 3.4.Main study 49 3.4.1 Tests for random mating 49 3.4.2 G-tests 49 3.4.2.1 Comparisons between individual samples 64 3.4.2.2 Gst analysis of pooled data 67 3.4.3 Nei's genetic distance and identity 67 4.0 Discussion 4.1 Genetic differentiation among populations 71 4. 2 Galaxias maculatus 71 4. 3 Lovettia sealii 77 4.4 Management Implications 82 4.5 Conclusions 83 5. 0 Appendices 2a Homogenising buffer 84 2b Electrophorectic running buffers 84 2c Optimal electrphoretic running conditions 84 2d Histochemical stains 87 3a L. sealii chi-square test with pooling 88 3b F-statistics values for individual loci in the L. sealii samples 108 6. 0 References 116 Chapter 1 1.0 Introduction 1.1 General Aims Large spring migrations of whitebait were observed in the 1980s by members of the Tasmanian public and, in particular, by retired whitebait fishermen CVV. Fulton pers. comm.). The current study was initiated in response to mounting public and political pressure to re­ open the whitebait fishery. This study was undertaken as part of a Fishing Industry Research Trust Account (F.l.R.T.A.) project (85/52) to determine if the present whitebait populations in Tasmanian rivers could support any level of commercial or recreational fishing. If a sustainable yield could be identified, guidelines for management of the fishery consistent with species and stock conservation would be established. Also included was the possibility of implementing measures to protect populations and, if possible to assist the recovery of the fishery. The two most important components of Tasmanian whitebait are the species Lovettia sealii (Johnston) and Galaxias maculatus (Jenyns). Other less important members of the whitebait runs include Galaxias truttaceus Valenciennes, Galaxias brevipinnis Gunther, Galaxias cleaveri Scott and Retropinna tasmanica McCulloch (Fulton, 1990). The extent to which G. maculatus and L. sealii occur as ecologically and geographically differentiated stocks in Tasmania is a matter of practical interest to fisheries managers, particularly when contemplating the re-opening of the commercial fishery. The identification of distinct genetic populations within the area would establish the need for stock conservation and possibly provide a tool for stock identification. Conversely, if regional populations could not be distinguished by genetic methods, there might be less need for managers to take population structure into account allowing for a broad scale management plan of the species. This research had three major goals: 1. to establish the optimal conditions for the electrophoretic analysis of stocks of Lovettia sealii and Galaxias maculatus, 2. to obtain frozen tissue samples of each species from various localities throughout the species range and, analyze these samples for evidence of stock heterogeneity; and 3. if the existence of two or more genetically differentiated stocks was detected, to determine if possible the genetic characteristics and geographic boundaries of each stock. 1.2 Lovettia sealii 1.2.1 Taxononac Background Lovettia sealii (Johnston) was originally described as a species of Aplochiton (Johnston, 1883). McCulloch (1915) recognised its generic distinction from Aplochiton and established Lovettia. The genus was named in acknowledgement of the specimens 1 captured in the Derwent River, Hobart and preserved by Mr E. F. Lovett (McCulloch, 1915). Jenyns (1842) placed Aplochiton in the family Salmonidae. In a comprehensive taxonomic review, McDowall (1971) considered Lovettia a member of the family Aplochitonidae which contained two genera: Aplochiton Jenyns with 2 species found in South America and Lovettia a monotypic genus found in Tasmania, Australia (McDowall, 1971). The species of Aplochiton recognised by McDowall (1971) were A. zebra Jenyns and A. taeniatus Jenyns. Both are found in Chile, Argentina, Tierra del Fuego and the Falkland Islands. Features of the Aplochitonidae include: small to medium size fishes (up to 240 mm), urinogenital apertures on a papilla set in, a post-anal depression (Aplochiton) or protruding (Lovettia), sexes similar (Aplochiton) or dimorphic (Lovettia), lacking in scales but with a lateral line.
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