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Age, Growth and Early Life History of Carp (Cyprinus Carpio L.) in The

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Age, Growth and Early Life History of Carp (Cyprinus Carpio L.) in The à¡ \''ts o.\. '{lÍ.; Age, Growth and Early Life History C) of Carp (Cyprinus cørpio L.) in the River Murray, South Australia Lorenzo Y ilizzi, B. Sc. (Hons) River Murray Laboratory Department of Zoology University of Adelaide Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy 1997 -June - Angell went on to suggest a fascinating and plausible explanation for the origin of the fox terrier simile (no excuse, of course, for its cloning). Fox terriers were bred "to dig out foxes from their burrows, when a fox had gone to earth during a traditional British hunt". Apparently, generations of fox-hunting gentlemen selected fox terriers not only for their functional role in the hunt but also under a breeder's advice to make them look as much like horses as possible. Angell continues, "The dogs rode up on the saddle during the hunt, and it was a pretty conceit for the owner-horseman to appear to put down a little simulacrum of a horse when the pack ofhounds and the pink-coated throng had arrived at an earth where the animal was to do his work". He also pointed out that fox terriers tend to develop varied patches of color on a basically white coat and that a "saddle" along the back is "considered desirable and handsome". Thus, Angell proposed his solution: "Wouldn't it seem possible that some early horse geologist, in casting about for the right size animal to fit his cliché-to-be, might have settled, quite unconsciously, on a breed of dog that fitted the specifications in looks as well as sizel". S. J. Gould (1988) To my belooed wire-høired fox terrier Blichi TABLE OF CONTBNTS ACKNOWLEDGMENTS......... ....... xiii l.INTRODUCTION 1 1.1 C¡.np AND rrs ADVENT To AusrRALrA 1 1.2 KNowN AND LINKNowN IMPACTS: DATA-GATHERING oR CoNTEMPLATIoN? JI 1.3 Pnpvrous BroI.ocrcAl. sruDrES oF cARp rN AusrRnLrA ..... 6 1.4 OerscrrvEs AND oRGANrsATroN....... 8 2. AGE AND GRO\ryTH: VALIDATION OF AGE ASSESSMENT 11 2.1 INrnooucrroN 11 2.2 MerpnrALS AND METHoDS.., I2 2.2.I Terminology 12 2.2.2 Sampling and preparation 12 2. 2. 3 Annulus identification... l3 2.2.3.I Scales l3 2.2.3.2 Opercular bones ...........13 2.2.4 Data analysis . ... 16 2.2.4.2 Edge interpretation and ETA.. t7 TABLE OF CONTENTS 11 t7 2.3 Rpsulrs I7 2.3.1 Marginøl increme'nt ratio analysis 22 2. 3. 2 Edge interpretation ..... ".'... 23 2. 3. j Edge type analysis'..."'...'.... 23 2.3.4 Scale outer circuli counts.... 23 2. 3. 5 Annulus distance .'.....'...'... 27 2.4 Dtscusstt-lt'1. 30 2.5 CoNct usloN 3l 3. AGE AND GROWTH: CONSISTENCY OF' AGE INTERPRETATIONS "" 31 3.1 INrnooucrloN 32 3.2 MerHoPs 32 3.2.I TerminologY 32 j. 2. 2 Experimental laYout . "..'. 33 3,2.2,lSfagel.ComparisonofstructuresandconsistencybylnterpreterA 33 3.2.2.2 Slage 2. Consistency by Interpreters A and E ,JJ 3.2.2.3 STage 3. Precision by Interpreters B, C and D 34 3. 2. 3 Dota analYsis..... .35 3.3 R¡sulrs... 35 3.3.1 Stage l. comparisonof structures; bias andreproducibility.......'....'.'.'.." 37 3.3.2 Stage l. Comparison of structures: bias"""""' """ 38 3.3.2.1 Opercular bones and scales".," """" " 38 3.3.2.2 Whole otoliths and scales """""""" " 38 3.3.2.3 Sectioned otoliths and scales """"""" 38 3,3.2.4 Whole otoliths and opercular bones""""""" """"""' 38 3.3.2.5 Sectioned otoliths and opercular bones"""" " " "" 38 3.3.2.6 Sectioned otoliths and whole otoliths """""' """ """ TABLE OF CONTENTS lll 3.3.3 Stage l. Consistency by Interpreter A. 42 3.3.4 Stage 2. Consistency by Interpreters A and E. 50 3.3.5 Stage 3. Precision by Interpreters B, C and D 52 3.3.6 Comparison of opercular bones andwhole otoliths by categories.............. 52 3.4 DrscussroN........ 53 4. AGE AND GROWTH: EVALUATION OF MODELS 59 4.1 INrnooucrroN 59 4.2 Meruoos 60 4.2.1 Terminologt 60 4.2.2 Sample measurement .... 60 4.2.3 Designation of a birth-date and ages adjustment 60 4.2.4 Data analysis 62 4.3 Rpsulrs 64 4. 3. I Comparison of length-at-age functions 64 4.3.2 Comparison of growth between sexes 66 4.3.3 Growth in body depth and index of obesity 67 4. 3. 4 Otolith growth..... 73 4.4 DrscussroN l5 5. AGE AND GROWTH: THE CARP IN LAKE CRESCENT, TASMANIA....81 5.1 INrnooucrroN 81 5.2 MersnrALS AND Mpruoos... 82 5.2.1 Samples and preparation 82 5.2.2 Validation and ages adjustment 82 lv TABLE OF CONTENTS 5.2.3 Data analysis 84 5.3 R¡sulrs 84 5. 3. I Length-frequency analysis 84 5. 3. 2 Interpretability of otoliths 86 5.3"3 Validation of annulus counts 86 5.3.3.1 Marginal increment ratio analysis 86 5.3.3.2 Edge type analysis 8ó 5.3.3.4 Annulus distance 89 5.3.4 Somatic growth models 89 5.4 DrscussroN 9l 5.5 CoNcr-usroN 6. AGE AND GROWTH: A CENTURY OF STUDIES ON CARP AGEING ....97 6.1 Ovenvrsw 97 6.2 Ev truATroN oF STRUCTURES 6.2.I Scales 99 6.2.2 Opercular bones 102 6.2.3 Spines andfin roys..... 102 6.2.4 Vertebrae and other bones 103 6.2.5 The eye lens 103 6.2.6 Otoliths 104 7. AGE AND GROWTH: "A MOST SUCCESSFUL COLONIZER"...............105 7.1 INrnooucrroN 105 7.2 MnrenrALS AND Meruoos... 108 TABLE OF CONTENTS V 7.2.1 Data collection and criteriafor selection 108 7.2.2 Data analysis 108 7.3 R¡sulrs .111 7.4 DrscusstoN 111 7.5 Ln¿rrnrroNs oF THE sruDY 118 8. EARLY LIFE HISTORY: THE ONSET OF THE JUVENILE PERIOD ....125 8.1 lNrnooucrroN t2s 8.2 Ksv sruDrES t27 8.2.1 Criteriafor the 'larva-juvenile transition' in carp 127 8.2.2 Relative growth 128 8. 2. 3 Functional morphology.. 129 8.2.4 Social relations and 'shooting' 130 8.2.5 Habitat use 131 8.3 DrscussroN 131 9. EARLY LIFE HISTORY: AGE, GRO\ilTH AND COHORT ANALYSIS.. 135 9.1 INrnooucrroN. 13s 9.2 M¿.renrALS AND Msrnoos . 136 9.2.1 Sampling and preparation 136 9.2.2 Otolith processing and examination 138 9.2.3 Validation of microincrement counts 140 9.2.4 Identification of developmental steps and cohort analysis 141 9.2.5 Data analysis 142 9.3 Rpsulrs 143 vr TABLE OF CONTENTS 9.3.1 Validation of microincrement counts . 143 9.3.2 Microincrement anolysis of un-marked otoliths 144 9.3.3 Cohort analysis and estimation of hatching dates 145 9.3.4 Growth models 148 9. 3. 5 Shooting analysis 149 9.4 DrscussroN r54 9. 4. I Otolith microinuement analysis r54 9.4.2 Cohort analysis 156 9.4.3 Growth models 157 9.4.4 Shooting 158 10. EARLY LIFE HISTORY: OBSERVATIONS ON DIET 159 10.1 INrnoDUCTroN ls9 10.2 MerHoos 160 10.3 R¡sulrs 161 10.4 DrscussroN 11. CONCLUSION AND PROSPECTUS......... ................ 165 REFERENCES t69 PAPERS IN SUPPORT 217 SYNOPSIS Since their escape into the Murray-Darling Basin (MDB) in the mid-1960s carp (Cyprinus carpio L,) have attained high densities and are now widely held responsible for detrimental effects on freshwater communities. Given the extensive literature from other countries little is known of the basic biology of this exotic invasive species in Australia. In particular, the ability to assess the age profiles of wild populations and a better knowledge of early life history events are fundamental for studies on population dynamics, particularly recruitment and Year-Class-Strength (YCS), and for the evaluation of control methods like intensive fishing, water drawdown and immunocontraception. The objectives of this research project were therefore (1) to determine a reliable method of age determination, (2) to evaluate models of growth in wild populations, (3) to assess and speculate on growth patterns, (4) to describe the onset of the juvenile period, (5) to monitor the early life history of a wild population, and (6) to review the literature on carp ecology. Interpreting the ages of individual carp is difficult, but possible, as a comparative analysis of the relative value of scales, opercular bones and otoliths (asterisci) from 603 specimens caught in the lower Murray in 1994-95 and an evaluation study of the bias and reproducibility of age interpretations have demonstrated. Validation of age estimates by marginal increment analysis has been achieved for carp 1-7 years old, although there is evidence that annulus patterns in opercular bones and otoliths can be used to interpret the age of fish up to 15 years old. Six models, including the Von Bertalanfry Growth Function (VBGF) and polynomial curves with combinations of log-transformed and decimal variables, were used to describe growth in length and weight of lower Murray carp. A log-linear quadratic function, by virtue of its precision, and the VBGF, with wider applicability and more biological realism, were chosen to describe growth in length, which was found to differ significantly between males and females; similar results were obtained upon comparison of growth in weight between the sexes. Otolith growth was also modelled, although the weight of these calcified structures could not be used as an objective criterion for age estimation. These methodologies, together with length-frequency analysis, were also employed to assess the vlll SYNOPSIS age composition and growth of a recently discovered population in Lakes Crescent and Sorell in Tasmania, based on a sample of 333 specimens.
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