University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh BOOK NUMBER QL6185SM d4 Thes?s. R o o m .. ACCESSION NO. . $ m i 2 3 j.. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh THE SYSTEMATICS OF INDO-PACIFIC SAUR1VA AND TRACHINOCEPHALUS (PISCES: SYNODONTIDAE) AND THE BIOLOGY OF FOUR LOCAL SPECIES. by EBENEZER LARYEA ADJEI, B.Sc. (Hons), Ghana. MARCH, 1984 Thesis submitted for the degree of Doctor of Philosophy Department of Zoology, University of Queensland, St. Lucia, Queensland, Australia. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh ABSTRACT Lizardfish species of the genera Saurida Cuvier and Valenciennes and Traohinoeephalus Gill 1862 in the Indo- Pacific are morphologically cryptic and widely distributed. Their habitats range from shallow water to deeper waters, sand/muddy bottoms or reefs. Many morphological features of lizardfish are variable and often of limited value in establishing taxonomic relationships. Synonymies and especially misidentifications are prevalent. An exploratory electrophoretic analysis confirmed intra-specific variation but also revealed specific differences which reflected species affinities. Accordingly the diagnosis of Indo-Pacific Saurida is revised and eleven species - S. argentea, S. australis, S. elongata, S. filamentosa, S. flamma, S. gracilis, S. isarankurai, S. longimanus, S. nebulosa, S. tumbil and S. undosquamis - are described. 5. australis is removed from the synonymy of S. undosquamis and inter­ relationships between members of the genus are described. S. wanieso and S. micropectoralis both described by Shindo and Yamada (1972) are synonymized under S. filamentosa and S. argentea respectively. S. wanieso is considered as a geographical race of S. filamentosa whereas S. micropectoralis is, by priority, a junior synonym of S. argentea. The biology of lizardfish were studied from collections taken within and without Moreton Bay between August 1979 and December 1981. The analysis of stomach items and morpho­ logical adaptations revealed that the food and feeding strategies of lizardfish consist of adaptability to a wide range of food items, food sizes and environmental conditions instead of specialization to few food items. The most important component of stomach items (by frequency of occurrence) were fish in over 85% of stomachs analysed (i.e. those with food). The other two major components were Crustacea and Cephalopods in about 5-16% and about 7% respectively. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Stomach content analysis indicated that lizardfish fed during both day and night. Mouth and body structures are well suited to their hunting techniques and carnivorous mode of feeding. Lizardfish reproduction within and without Moreton Bay is described with emphasis on the development of the gonads. Four types of eggs based on size and yolk content are described. Eight and six stages of identifiable gonads based on a) types and quantities of ova present and b) texture and macroscopic appearance are described for the ovaries and testes respectively. The species exhibit size polymorphism - the females predominate in sizes greater than male S.L^q whereas the males predominate in sizes about the male S.L.^q and below. The reproductive strategy involves an extended spawning season, early maturation, high fecundity coupled with serial spawning. The ages of S. tumbil, S. undosquamis and T. myops on the one hand and S. filamentosa on the other were determined by the scale and vertebrae methods respectively and were supplemented by the length frequency method. The observed maximum ages in the species were 5, 4, 12 and 3 for S. undosquamis, S. tumbil, S filamentosa and T. myops respectively. Lizardfish growth is asymptotic and alloraetric and was described in terms of Von Bertalanffy and other growth parameters. The overall growth strategy apparently involves a number of features - very rapid linear growth, early maturation, annual and allometric growth. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh The four lizardfish species were parasitized by cestodes, hemiurid trematodes, nematodes, isopods and acanthocephalans. Negative binomial distribution (truncated and non-truncated) analyses in comparison with actual frequency distribution of the most common parasite in Moreton Bay, Callitetrarhynohus gracilis (Rudolphi) 1819, suggested that ’heavily’ infected fish were missing from the samples. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh iv STATEMENT OF RESPONSIBILITY This thesis is an original work and has not been previously submitted to another university. Assistance received and any information not derived from the study are specifically acknowledged. — ^ EBENEZER LARYe4 adji^ University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh V ACKNOWLEDGEMENT I am grateful to Professor J.M. Thomson for his supervision, for translating some articles in Russian and critical reading of the manuscript. Many people have been very helpful during the course of this study and to all I extend my sincere gratitude. In particular I would like to thank - Dr. R.J. McKay and Mr J. Johnson of the Queensland Museum, Brisbane; Dr. J.R. Paxton of the Australian Museum, Sydney (AMS) and Dr. B.C. Russell formerly of AMS but now at the Smithsonian Institute, Washington - for loan of material, for providing some important literature references and laboratory space during the study. I am especially grateful to Mr. Les Wale, skipper of the University research vessel, 'Sea Wanderer', for his pleasant company and considerable efforts he put in in obtaining specimens from Moreton Bay and also for setting up contacts with some professional fishermen who assisted me in many ways, in particular Mr. Bob Sanderson, then skipper of the 'Rex Andria' who provided the deep sea specimens. Messrs S. Cook, D. Webb and Dr. N. Bruce often provided assistance in the field. Messrs S. Cook, C. Keenan, A.P. Bradshaw and W. Thomas of Zoology Department very often collected specimens for me during their field work on Moreton Bay. On the other hand, Messrs Greg Campbell, Queensland Fisheries, Bundaberg; B. Wallis, skipper of 'Curlew' Innisfail; Dr. H. Sweatman, Centre for Environmental and Urban Studies, Macquarie University; the crews of FV Courageous and FRV Soela and Dr. K. Sainsbury all of CSIRO, Cronulla provided specimens from the Great Barrier Reef and the North West Shelf. In addition I would like to thank the following persons or curators of fish for loans of personal or museum collect­ ions - Dr. P.J. Whitehead, British Museum of Natural History, University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh vi London (BMNH); Dr. V.G. Springer, National Museum of Natural History - Smithsonian Institution, Washington; Dr. J.E. Randall, Bernice P. Bishop Museum, Honolulu; Mr. John Paska, Kanudi Fisheries Research Museum, Konedobu, Papua New Guinea; Dr. Y. Tominaga, University of Tokyo Zoology Museum; L.E. Timms, Department of Primary Production of the Northern Territory of Australia, Fisheries Division; Mrs. P.J. Kailola, Fish Taxonomy consultant, Adelaide and Mr. T. Gloerfelt-Tarp of Bali, Indonesia. Summaries of occurrences of lizardfish from North West Cape to Darwin, and off New South Wales were supplied by Dr. Grant West, CSIRO, Cronulla and Mr. R.J. William, New South Wales State Fisheries, Sydney respectively. Dr. J.B. Shaklee, CSIRO, Cleveland and Mr. C. Keenan provided equipment for electrophoretic runs. I would also like to thank Dr. R. Lester, Parasitology Department, University of Queensland for the identification of cestode and nematode parasites and discussions on data of cestode larvae; Dr. D.I. Gibson, BMNH, for the identific­ ation of stomach trematodes of S. undosquamis and Dr. N. Bruce, formerly of Zoology Department, University of Queensland, now with Smithsonian Institution, Washington for the identific­ ation of crustacean parasites. My gratitude to the technicians of Zoology Department for their help with equipment on many occasions - in particular, to Mr. T. Dyer, the Chief Technician, and to Mr. T. Gorringe for his invaluable help and patience in photographing the dental configuration of lizardfish. The staff of the Biological Sciences Library were very helpful in obtaining literature references, especially inter-library loans. Miss Margaret Russet formerly of University of Queensland, and now at Batemans Bay kindly translated several articles in French. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh vii Mr. W. Hutchinson, the Administrative Officer of Zoology Department, was very helpful on administrative matters. My sincere thanks to my friend Miss Lidia C. Meszaros for her support and encouragement, and also for typing the manuscript. The research was funded by University of Queensland Research Grants. The Ghana Government (University of Ghana) provided only for living expenses between July 1979 and March 1983. Finally, I would like to thank my step-father, Mr. F.P.K. Amo-Tenkorang, and especially my mother who had to take out a mortgage to see me through the initial years of secondary school when my father refused to help. And also my brother Mr. W. Adjetey Adjei and sisters Helena and Sabina Adjeley Adjei who have supported me whenever possible in my academic career so far. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh CONTENTS Page ABSTRACT 1 ACKNOWLEDGEMENTS 1v 1. GENERAL INTRODUCTION 1 1.1 Materials and Methods 3 1.1.1 Systematics 3 1.1.2 Biology 4 1.1.2.1 Collection of samples 4 1.2 Moreton Bay 4 1.3 Treatment of samples 5 1.4 Changes in weight and length of fish after fixation in formalin followed by soaking in water. 7 1.5 Data analyses 8 2. MORPHOLOGICAL AND BIOCHEMICAL SYSTEMATICS 9 2.1 Introduction 9 2.1.1 Historical review of the classification of lizardfish 10 2.1.2 The family Synodontidae 12 2.2 Materials and Methods 17 2.2.1 Electrophoretic studies 17 2.2.1.1 Species samples 17 2.2.1.2 Preparation of tissue extracts 18 2.2.2 Morphological/Anatomical studies 21 2.2.2.1 Species samples 21 2.2.2.2 Measurements, counts and definitions 22 2.2.2.3 Abbreviations of Museums/Institutions 24 2.3 Results 25 2.3.1 Electrophoresis (Biochemical Systematics) 25 2.3.1.1 General proteins 25 2.3.1.2 Specific proteins 25 2.3.1.3 Generic differences and similarities 26 2.3.1.4 Inter- and intra-specific variation 26 2.3.1.4.1 Observations in Group A 27 2.3.1.4.2 Observations in Group B 28 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Page 2.3.2 Morphological Systematics 34 2.3.2.1 Generic characters 34 2.3.2.2 Genus Saurida 35 2.3.2.2.1 Important diagnostic characters of Saurida 37 2.3.2.2.1.1 Palatine teeth 37 2.3.2.2.1.1.1 5. undosquamis type 37 2.3.2.2.1.1.2 S. tumbil type 37 2.3.2.2.1.1.3 S. nebulosa type 37 2.3.2.2.1.2 Snout and nasal flap shapes 38 2.3.2.2.1.2.1 S . undosquamis type 38 2.3.2.2.1.2.2 5. tumbil type 38 2.3.2.2.1.2.3 S . nebulosa type 39 2.3.2.2.1.3 Pectoral fin 39 2.3.2.2.1.4 Second and third dorsal fin rays 39 2.3.2.2.1.5 Jaws 40 2.3.2.2.1.6 Colour 40 2.3.2.2.2 Specific descriptions and distributions 40 2.3.2.2.2.1 Saurida argentea Macleay, 1882 40 2.3.2.2.2.2 Saurida australis Castelnau, 1878-79 48 2.3.2.2.2.3 Saurida elongata (Temminck and Schlegel), 1846 52 2.3.2.2.2.4 Saurida filamentosa Ogilby, 1910 54 2.3.2.2.2.5 Saurida flamma Waples, 1982 57 2.3.2.2.2.6 Saurida gracilis Quoy and Gaimard, 1824 59 2.3.2.2.2.7 Saurida isarankurai Shindo and Yamada, 1972 61 2.3.2.2.2.8 Saurida longimanus Norman, 1939 63 2.3.2.2.2.9 Saurida nebulosa Valenciennes, 1849 65 2.3.2.2.2.10 Saurida tumbil (Bloch), 1795 68 2.3.2.2.2.11 Saurida undosquamis (Richardson), 1848 72 2.3.2.3 Genus Trachinocephalus Gill, 1862 75 2.3.2.3.1 Trachinocephalus myops (Bloch and Schneider) , 1801 7 6 2.4 Discussion 3 2 3. FOOD, FEEDING HABITS AND ADAPTATIONS 89 3.1 Introduction 89 3.2 Methods and Materials 94 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Page 3.3 Results 99 3.3.1 Feeding patterns of lizardfish 99 3.3.2 Orientation of item in stomach 106 3.3.3 Variation of feeding and prey abundance 106 3.3.4 Variation of feeding and food com­ position with sex, length and month 106 3.3.5 Daily variation in feeding and activity 107 3.3.6 Morphology of Synodontidae associated with feeding 107 3.3.6.1 Mouth, body cavity and gut 107 3.3.6.2 Teeth 109 3.3.6.3 Sensory organs 110 3.3.6.4 Fins 111 3.3.6.5 Colour 111 3.4 Discussion 112 AGE AND GROWTH 125 4.1 Introduction 125 4.2 Methods and Materials 133 4.3 Results 135 4.3.1 Variation of scale morphology with body site 135 4.3.2 Definition of rings on scale, time of ring formation and age determination 136 4.3.2.1 Definition of rings on scale 136 4.3.2.2 Time of ring formation 140 Age and standard length 140 4.3.3 Scale radius and standard length 141 4.3.4 Back-calculation of standard length at time of ring formation in s. t u m b i l 141 4.3.5 Cassie’s length frequency method for the estimation of age 145 4.3.6 Growth equations and growth curves 148 4.3.6.1 Formulae used in the calculation of Von Bertalanffy and other growth parameters 148 4.3.6.1. 1 Von Bertalanffy growth equations and parameters 149 4.3.6.2 Recruitment, mortality and survival in S. t u m b i l and S. u n d o s q u a m i s 153 CO CM CO University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Page 4 .3.6.3 Relationship between body weight and standard length 154 4.3.6.4 Growth curves 157 4.4 Discussion 158 5. REPRODUCTION OF FOUR LOCAL SPECIES 166 5.1 Introduction 166 5.2 Methods and Materials 168 5.3 Results 171 5.3.1 Ova types 171 5.3.2 Maturity stages 172 5.3.2.1 Stages of maturity of ovaries 173 5.3.2.2 Stages of maturity of testes 174 5.3.3 Length at maturity 178 5.3.4 Spawning 179 5.3.4.1 5 . undosquamis and S. tumbil 179 5.3.4.2 s. filamentosa 180 5.3.4.3 t. myops 180 5.3.5 Fecundity 180 5.3.6 Sex ratio 184 5.3.6.1 S. undosquamis 184 5.3.6.2 S . tumbil 184 5.3.6.3 S. filamentosa and T. myops 187 5.4 Discussion 189 6. THE PARASITES OF FOUR LOCAL LIZARDFISH INCLUDING AN ANALYSIS OF THE FREQUENCY DISTRIBUTION OF THE CESTODE CALLITETRARHYNCHUS GRACILIS (LARVAE) (WITH REFERENCE TO THE NEGATIVE BINOMIAL DISTRIBUTION) 200 6.1 Introduction 200 6.2 Methods and Materials 205 6.3 Results 207 6.3.1 Parasite fauna 207 6.3.1.1 Trematodes 208 6.3.1.2 Cestodes 208 6.3.2 Negative binomial distribution and the frequency distribution of C. gracilis larvae 213 6.4 Discussion 225 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Page 7. GENERAL DISCUSSION 230 REFERENCES 240 APPENDIX 1 284 APPENDIX 2 285 APPENDIX 3 286 APPENDIX 4 - A 287 B 288 C 289 APPENDIX 5 - A 290 B 291 c 292 D 293 APPENDIX 6 294 APPENDIX 7 295 APPENDIX 8 ,q . University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh list of tables Table No. Pa^e No* 1. Effect of formalin on length and weight 7 2.0 Legend for plates and figures 25a Meristic characters of lizardfish (2.1 - 2.4) 2.1 Lateral line scales 42 2.2 Fin rays 43 2.3 Scale counts and Morphometry 44 2.4 Mean and range of morphometric characters continued A. As percentage of standard length 45 B. As percentage of head length 46 3.1 List of stomach items of lizardfish 100 3.2 Relative percentage prey sizes 105 3.3 Feeding indices 108 3.4 Dimensions of prey relative to lizardfish (predator) 116 4.1 A. Definition of rings on scale of female S . t umbi1 137 B. Definition of rings on scale of male S. tumbil 138 4.2 Age and standard length 142 4.3 Relationship between standard length and scale radius 143 4.4 A. Mean ring radius of s. tumbil 144 B. Back-calculated lengths 144 4.5 Growth of lizardfish A. Observed and calculated standard length 146 B. Calculated whole body weight 147 4.6 Von Bertalanffy and other growth parameters 151 4.7 Standard length-weight relationship 155 4.8 Analysis of covariance in length-weight relationship between the sexes 156 4.9 Von Bertalanffy growth parameters and latitude 163 5.1 Ova diameters of Saurida and Trachinocephalus 172 5.2 Length at maturity of S . undosquamis, S. tumbil and T. m y o p s 176 5.3 Length at maturity of S. filamentosa 177 5.4 Relative sizes of mature and immature lizardfish 178 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Table No. Page No. 5.5 A. Constants in fecundity equation 181 B. Fecundities 182 5.6 Chi squared tests of sex ratios 183 5.7 Sex ratio during spawning season 185 5.8 Monthly sex ratio during spawning season of S. filamentosa and T. myops 186 5.9 Percentage ratios of second and third dorsal rays to standard length of S. filamentosa 188 5.10 Relative fecundity and number of spawnings of lizardfish 196 5.11 Annual mortalities in s. undosquamis 198 6.1 Parasites found in lizardfish 207 6.2 Percentage incidence of trematodes in lizard- fish samples 208 6.3 Incidence of c. gracilis in s. tumbil and S. undosquamis 209 6.4 Frequency distribution of c. gracilis with size and sex of lizardfish 210 6.5 Statistics of frequency distribution of C. gracilis in lizardfish 212 6.6 Frequency distribution of c. gracilis by location in lizardfish 214 6.7 Negative Binomial Distribution A. c. gracilis by aorta in female S. tumbil 215 B. c. gracilis in female S. tumbil 216 C. c. gracilis by aorta in male s. tumbil 217 D. c. gracilis in male S. tumbil 218 E. c. gracilis by aorta in female s. undosquamis 219 F. c. gracilis in female S. undosquamis 220 G. c. gracilis by aorta in male 5. undosquamis 221 H. c. gracilis in male 5. undosquamis 222 6.8 Negative Binomial Distribution of c. gracilis larvae by aorta in lizardfish using p = x/k 223 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh LIST OF FIGURES (Figures and legends to these are on unnumbered pages in the body of the thesis. For ease of reference the number of the preceding page is recorded). Figure No. Page No. 1.1 Map of Moreton Bay showing areas trawled 4 2.1 Length measurements (morphometry) 22 2.2 Diagrams of the zymogram patterns of some enzymes (AK, IDH, MPI, AAT) of Saurida and Trachinocephalus 26 2.3 Diagrams of the zymogram patterns of some enzymes (ADH, GAPDH, GDH, PGDH) of Saurida and Trachinocephalus 26 2.4 Diagrams of the zymogram patterns of some enzymes (SDH, SOD, UMB, XDH) of Saurida and Trachinocephalus 2 6 2.5 Diagrams of the zymogram patterns of CK and EST of species of Saurida and Trachinocephalus 27 2.6 Diagrams of the zymogram patterns of GAPDH and G-3- PDH of species of Saurida and Trachinocephalus 27 2.7 A diagram of the zymogram patterns of SOD of species of Saurida and Trachinocephalus 28 2.8 Diagrams of the zymogram patterns of some enzymes (AK, MDH, AAT) of species of Saurida 29 2.9 Diagrams of the zymogram patterns of some enzymes (ADH, AH) of species of Saurida 29 2.10 Diagrams of the zymogram patterns of some enzymes (G-6-PDH, GPI, IDH, MDH) of Saurida species 30 2.11 Diagrams of the zymogram patterns of some muscle enzymes (ME, MPI, PGDH, XDH) of saurida 31 2.12 Ventral view of pelvic fins of Trachinocephalus , Synodus and Saurida 35 2.13 Diagrams of snout, nasal flap and pectoral fin of Saurida species 35 2.14 Diagrams of palatine and vomerine teeth configurations in saurida species 37 2.15 Distribution of saurida argentea Macleay 1882, S. elongata (Temminck and Schlegel) 1846 and S. australis Castelnau 1878-79 48 2. 16 Distribution of Saurida australis Castelnau 1878-79 51 2.17 Distribution of Saurida filamentosa Ogilby 1910, s. longimanus Norman 1939 and s. isarankurai Shindo and Yamada 1972 56 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Figure No. Page No. 2 #i8 Distribution of Saurida nebulosa Valenciennes 1849, S. gracilis Quoy and Gaimard 1824 and 5. flamma Waples 1982 59 2.19 Distribution of Saurida tumbil (Bloch) 1795 71 2.20 Distribution of Saurida undosquamis (Richardson) 1848 75 2.21 Distribution of Traohinocephalus myops (Bloch and Schneider) 1801 81 3.1 Monthly variation of prey abundance and frequency of occurrence in lizardfish stomachs 106 3.2 Daily activity of s. undosquamis 107 3.3 Typical lizardfish, gape and second lower gill arch 109 3.4 Types of teeth, caudal aspect and stereoscopic vision 109 3.5 Regression of width and length of premaxillary of standard length of S. undosquamis 121 4.1 The scale of lizardfish 134 4.2 Scale and body sites 134 4.3 s . t umbi1 : A, B, C, D - Relationship between ring radii and scale radius 141 E, F - Relationship between standard length and scale radius 141 4.4 Relationship between standard length and scale radius of s . undosquamis and T. myops 141 4.5 Size frequency distribution of lizardfish 145 4.6 Cassie curves for - A. 478 female s. tumbil 147 B. 475 male s . tumbil 147 C. 2606 female s. undosquamis 147 D. 2492 male S. undosquamis 147 E. 497 female t . myops 147 F. 537 male r. myops 147 G. 350 female S. filamentosa 147 H. 278 male S. filamentosa 147 4.7 Walford plots of lizardfish A. S. tumbil and S. undosquamis 148 B. S. filamentosa and T . myops 148 C. S. tumbil - back calculated lengths 150 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Figure No. Page No. 4.8 Time of ring formation and mean monthly standard length and weight of lizardfish 153 4.9 Relationship of weight to standard length of - A. s. tumbil 154 B. s. undosquamis 1 5 4 C. s. filamentosa 154 D. T . myops 154 4.10 Von Bertalanffy growth curves of lizardfish - standard length and age 157 4.11 Von Bertalanffy growth curves of lizardfish - weight and age 157 5.1 Probit analysis; estimation of length at maturity of female lizardfish 178 5.2 Probit analysis; estimation of length at maturity of male lizardfish 178 5.3 Monthly mean gonad index of lizardfish A. Females 179 B. Males 179 5.4 Frequencies of stages of maturity of - A. Ovaries of s . undosquamis 179 B. Testes of s . undosq uamis 179 C. Ovaries of s . tumbi1 179 D. Testes of s . tumbi1 179 E. Ovaries of s . f ilamentosa 179 F. Testes of s . f ilamentosa 179 G. Ovaries of T . myops 179 H. Testes of T. myops 179 5.5 Scatter diagram showing the relationship between fecundity and standard length/ weight of A. S . undosquamis 181 B. S. tumbil 181 C. T . myops 181 5.6 S.L^q and Latitude 191 5.7 Mean monthly rainfall, mean monthly minimum and maximum temperatures recorded at Cape Moreton Lighthouse Weather Station by the Bureau of Meteorology between 1958 and 1981 193 6.1 A. Incidence of c. gracilis larvae in population of s. tumbil in Moreton Bay 209 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Figure No. Page No. 6.1 B. Incidence of c. gracilis larvae in popu­ lation of 5. undosquamis in Moreton Bay 209 6.2 Frequency distribution of c. gracilis in Saurida spp. by aorta and in whole fish 213 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh LIST OF PLATES (Plates and legends to these are on unnumbered pages in the body of the thesis. For ease of reference the number of the preceding page is recorded). Plate No. Page No. 1. Comparison of general protein patterns of Saurida (Group B) 25a 2 . Comparison of general protein patterns of Saurida (Group A) and Traohinocephalus 25a 3 - 8 Inter-generic comparison of zymogram patterns of enzymes of Saurida and Traohinocephalus ate Enzyme 3 AK 26 4 MDH 26 5 GAPDH 26 6 PGDH 26 7 EST 26 8 LDH 26 9 - 1 2 Comparison of zymogram patterns of enzymes of species of Saurida and Traohinocephalus Plate Enzyme 9 G-3-PDH 27 10 LDH 27 11 MDH 28 12 SOD 28 13 - 28 Comparison of zymogram patterns of enzymes of five species Of Saurida Plate Enzyme 13 AK 29 14 MDH 29 15 AAT 29 16 EST 29 17 GDH 29 18 IDH 29 19 LDH 30 20 MDH 21 31ME 22 31MPI 23 31PGDH 31 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate No. Page No. 13 - 28 Plate Enzyme Cont'd 24 PGM 32 25 PK 32 26 SDH 32 27 SOD 32 28 XDH 33 29. Teeth configuration in palatine bands and on vomer of S. undosquamis 37 30. Teeth configuration in palatine bands and on vomer of S. tumbil 37 31. Teeth configuration in palatine bands and on vomer of S. nebulosa 37 32. Teeth configuration in palatine bands and on vomer of S. argentea 47 33. Saurida australis Castelnau 1878-79 50 34. Teeth configuration in palatine bands and on vomer of S. fi1amentosa 56 35. Teeth configuration in palatine bands and on vomer of S. longimanus 65 36. S. undosquamis 75 37. Teeth configuration in palatine bands of T . myops 7 9 38. S. tumbil scales with annuli 135 39. A. Callitetrarh ynchus gracilis larvae 209 B. C. gracilis (in situ) by aorta in S.tumbil 209 C. C. gracilis (in situ) by aorta in S.tumbil; expanded view. 209 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 1. GENERAL INTRODUCTION The family Synodontidae is a group of fish character­ ized by lizard-like head, spineless fins and a small adipose fin and are almost circumtropical in distribution. The family, represented on a conservative estimate by 40 species belonging to three genera have some of the most abundant species in the Indo-Pacific region. These species, especially S. undosquamis, S. tumbil and S. elongata form the basis of important commercial fisheries for food or poultry feed in Israel (Ben-Tuvia, 1953; Latif and Shenouda, 1973; Ben-Yami and Glaser, 1974), India (Rao, 1974), Gulf of Thailand (Ruamragsa and Isarankura, 1965; Vadhanakul, Eiamsa ard and Kuantanom, 1975; Sinoda and Intong, 1978; Sinoda, Lim and Tan, 1978), Sunda Strait, northern and north-western shelves of Australia (Liu, Lai and Yeh, 1978; Liu and Lai, 1980) and the seas of China and Japan (Liu and Tung, 1956; Liu and Chen, 1959; Tatara, 1965). In Japan they are the raw material for meat paste known as 'Kamaboko' or 'Himodori (Makinodan and Ikeda, 1971; Akashi and Oono, 1972; Suzuki, Kanna, Okazaki and Morita, 1978; Kurokawa, 1979; Yamazawa, Murase and Shiga, 1980). Although about 25 lizardfish species occur in Australian waters ( See Castelnau, 1878- 79; Macleay, 1882; Ogilby, 1897, 1910a,b; Waite, 1905 - 08; McCulloch, 1920-22, 1927, 1929 - 30, 1934; Norman, 1935; Whitley, 1943, 1948; Marshall, 1964; Grant, 1965, 1972 1975, 1978; Shindo and Yamada, 1972; Allen, Hoese, Paxton, Randall, Russell, Starck, Talbot and Whitley, 1976; Russel and Cressey (1979); Cressey, 1981) they are unacceptable to the Australian market because of their rather insipid nature. Notwithstanding they are a potential resource for export-eeriented fisheries in Australia. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 2. In the development of reliable management strategies, correct identification of the species is paramount. Incorrect or inconsistent identification of fish species form a fundamental block to the scientific management of fisheries by preventing both biological studies and the collection of reliable fisheries statistics. In 1981, Roger Cressey of the Smithsonian Institute, Washington, published a comprehensive review of the species of the genus Synodus of the Indo-West Pacific region. However the current taxonomic literature on Indo-Pacific Saurida is unsatisfactory and is plagued by synonymies, incorrect and inconsistent identifications. With regards to the biology of lizardfish, too often in the past, many authors have been content to accumulate sometimes unrelated facts, mainly anecdotes. The two most studied species are S. undosquamis and S. tumbil, many of these studies over short periods and even some tainted with incorrect identification (i.e. polyspecific samples e.g. Okada and Kyushin, 1955; Liu and Tung, 1959; Rao, 1974) and have thus depreciated the fishery management value of the studies. The present study is divided into two sections, the first dealing with the taxonomy and distribution of Indo- Pacific Saurida and Trachinocephalus as a whole, and the second with the biology of four abundant local species. Under the first section the taxonomy has been revised. The revision provides identification keys as well as descriptions, figures, synonymies and distributions for all species. The problem of inter- and intra-specific variation which has plagued their classical or conventional systematics, as with many other cryptic species, was investigated by exploratory electrophoretic analysis of general and specific (enzyme) proteins. This aspect of the study is reported on under - "Morphological and biochemical systematics of Indo- Pacific lizardfish” . The biology involved the collection and analyses of baseline data on adaptive strategies for or University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 3. 1) food and feeding 2) age and growth 3) reproduction and 4) parasites. The principles and common methods in use for the study of the four topics just mentioned are described and/or discussed in the various introductory sections. The synthesis of the results on the various topics was under­ taken in a general discussion with the view of elucidating the obvious success of the family in the Indo-Pacific region. 1.1 MATERIALS AND METHODS 1.1.1 Systematics Where possible materials held by museums were obtained and examined, especially from the Queensland Museum, Brisbane (QM), Australian Museum Sydney (AMS) and B.P. Bishop Musuem Honolulu, Hawaii (BPBM). Trips were also made to QM and AMS to examine collections which could not be borrowed for various reasons. Data and *fresh specimens of Synodontidae of the North and North-Western shelves of Australia were obtained by crews of FV Courageous and FRV Soela, both of the Commonwealth Scientific and Industrial Research Organis­ ation (CSIRO), Cronulla, New South Wales. Also many **people provided many other specimens from various parts of the Indo-Pacific region. Personal collections were also made within and off Moreton Bay. * Used for electrophoresis. ** Details provided in appropriate sections. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 4. 1.1.2 BIOLOGY 1.1.2.1 Collection of Samples Samples of the four abundant local species - Saurida tumbil (Bloch) 1795, S. undosquamis (Richardson) 1848, S. filamentosa Ogilby 1910 and Traohinooephalus myops (Bloch and Schneider) 1801 - were collected by bottom trawls. S. undosquamis and S.tumbil and a handful of T. myops were obtained by *forty-minute otter trawls using the University’s trawler 'Sea Wanderer’ in Moreton Bay from depths between about 10 to about 40 metres between August 1979 and December 1981. Large specimens (2) of S. undosquamis, the bulk of T. myops and all of S. filamentosa were obtained outside Moreton Bay, between 32-64 kilometres north of Cape Moreton from depths of about 80-200m mainly between May 1980 and February 1982. Any significant observations on condition of fresh fish with regards to reproduction (e.g. running eggs or milt and spent individuals), and parasites (ecto- and sometimes endo-parasites) were noted in the field and those individuals marked. The nature of the bottom of the section of bay trawled (gauged by sediments brought up with catches) was noted. 1.2 MORETON BAY (Fig. 1.1) Figure 1.1 shows the trawled areas or stations in Moreton Bay. Moreton Bay is a subtropical water mass. It is bordered by the Moreton and North Stradbroke Islands but linked to the Pacific Ocean by a number of channels. * Unless otherwise stated. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig.1,1 : Map of Moreton Bay showing areas trawled. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 5. Moreton Bay is also partly a complex estuarine system into which drain a number of rivers - Brisbane, Logan-Albert, Pine, and Caboolture. There are a number of islands in the bay, especially in the Southern Section. Apart from those islands close to estuaries (i.e. Fisherman, Mud and Macleay), all are surrounded by ill-defined fringing reefs extending to depths of 2-4 metres and dominated by Favia speciosa (Dans). (See Stephenson, Williams and Lance, 1970; Stephenson, Williams and Cook, 1974; Stephenson, Cook and Newlands, 1978; Stephenson and Williams, 1981). Fish samples were regularly taken from the following areas: Bramble Bay, off Redcliffe, east of Mud Island, north of Peel Island and from Middle Banks (especially from Tangalooma Point). For safety reasons and also possible damage to gear, trawling was restricted to the above areas since many of the other areas were too shallow, had wrecks or were too confined for safe trawling. 1.3 TREATMENT OF SAMPLES All samples for biological *study were preserved in 5-10% formalin and returned to the laboratory. After three weeks of formalin preservation, formalin was washed off fish four times with fresh tap water and then soaked out in tap water for two days, presumably to remove more formalin trapped in the body cavity. Each fish was given a reference number and the following biological observations were noted after excess water had been drained off and also removed with paper towel:- * Some were also used in conventional systematic studies. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 6. Standard ’►length the perpendicular distance between the snout tip and the crease of the caudal fin (was measured to the nearest 1mm on a measuring board i.e. the posterior margin of the hypural bones). Whole body *weight measured to the nearest 0.1 g. Food and Feeding types of items in stomach, length and weight of some well preserved prey specimens, diurnal feeding activity (gauged by fish numbers and stomach fullness with time), and morphological adaptations associated with food and feeding. Age determined from scales or vertebrae and length frequency analysis. Growth as indicated by changes in weight and length of fish. Sex determined by examining gonads under a dissecting microscope. Reproduction as indicated by changes in weight and texture of gonads. Parasitology types of ecto- and endo-parasites and also the analysis of numbers of the common parasite and its effect on the species. * All references to length and weight unless otherwise stated refer to standard length and whole body weight respectively. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 7. 1.4 CHANGES IN WEIGHT AND LENGTH OF FISH AFTER FIXATION IN FORMALIN FOLLOWED BY SOAKING IN WATER The changes (loss or gain in weight or length varied with the individual fish, most probably its physiological state. The results for 38 S. undosquamis and 20 S. tumbil are summarized in Table 1. Table 1 : Effect of formalin on Length and Weight Percentage changes (+-) in Species Weight Length *Duration After *Duration After Soaking Soaking out in out in 24hrs 3 weeks water 24hrs 3 weeks water range 0.3-6.4 0.7-7.1 0.6-3.4 0.0-4.8 0.0-3.0 0.0-1.4 S. undosquamis ** (121-182nm) Average 2.5 3.5 2.2 1.0 0.8 0.4 range 0.0-7.4 0.4-2.5 0.3-1.9 1.0-4.8 1.8-4.6 1.0-2.8 S. tumbil ** (124-267nm) Average 2.0 1.1 1.6 2.4 3.3 1.9 * of treatment in formalin ** size range. In a study of Pond and Amur wild carps, Amosov (1960) found changes in weight and length to be between 0.1 - 7.3 and 0.20 and 0.93 respectively. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 8. 1.5 DATA ANALYSES Statistical methods followed Sokal and Rohlf (1969) and to a limited extent ’’Statistical Package for the Social Sciences" (SPSS) documented in Nie, Hull, Jenkins, Steinbrenner, Bent (1975), Statistical Package for the Social Sciences Second Edition McGraw-Hill Book Company. Other statistical texts were also consulted and are mentioned in the appropriate sections. Methods - hypotheses or models - employed in the analyses of age and growth were mainly taken from Ricker (1975) and Bagenal (1978). The analysis of parasite data (i.e. distribution of cestode larvae) followed Crofton (1971). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 9. 2. MORPHOLOGICAL AND BIOCHEMICAL SYSTEMATICS 2.1 INTRODUCTION All species are the product of an historic process; speciation: first and foremost the establishment of reproductive isolation and a subsequent divergence of the population. The concomittant adaptations with speciation enable the species to meet their environmental conditions. The survival in, and the effective utilization of an heterongeneous environment by a species might entail the possession or the acquisition of a variety of adaptive responses (i.e. intra-specific variation in responses/ morphology) by members of the population. This variation in species can either be genetically or environmentally induced, or can be the product of a combination of the two. With time groups corresponding to varieties of adaptive responses become new species if reproductively isolated. Proper identification of species is a pre-requisite for most ecological studies, and this identification is dependent upon an adequate classification being available. Unfortun­ ately, more often than not, it is very difficult to distinguish between inter- and intra-specific variation using techniques of classical or conventional taxonomy. However, each change in an organism can ultimately be traced to alterations occurring at the molecular level, at least in theory and with the application of protein analysis, especially the electrophoretic studies of genetic variation, inter- and intra-specific variation of species and also the reproductive strategy practised by species are rather easily circumscribed. In the past thirty years, electrophoretic analysis of proteins have been used to elucidate the evolutionary process and also to disentangle many inter- and intra­ specific variations: University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 10. a) in populations - e.g. Irahof, Leary and Booke (1980) demonstrated the existence of at least four populations of lake whitefish Coregonus olupaeformis (Salmonidae) in northern Lake Michigan. b) in species and/or genera of e.g. Albulidae (Shaklee, Tamaru and Waples, 1982), Mugilidae (Anderson, 1982), Percichthyidae (MacDonald, 1978), Salmonidae (Ferguson, 1974; Ferguson, Himberg and SVardson, 1978), Scorpaenidae (Tsuyuki, Roberts, Lowes Hadaway. and Westrheim, 1968), Synodontidae (Shaklee et al, 1982; Taniguchi, 1969). Shaklee et al (1982) found strong agreement between the estimate of relationships based on genetic distance values and that based on morphological similarity in Synodus ulae, S. variegatus, S. englemani, S. binotatus and Saurida gracilis. Despite these findings some school of taxonomists have been very wary of these ’’biochemical species” . This state of affairs is adequately summed up by Moss (1979) - "Advances in data and technique can provide additional dimensions of knowledge, but they also pose a continual challenge: How does one integrate and reconcile them with existing classi­ fications? Some taxonomists are inclined to believe that only new data sources and new techniques yield crucial insight; others are more inclined to stretch the existing classifications rather than to demolish them1’. 2.1 Historical review of the classification of Lizardfish Lizardfish belong to a group of fish usually referred to as 'iniomous’ fish. *Regan (1911) considered these iniomous fish as ’marine malacopterous physostomes with the pelvic fins abdominal or thoracic in position, and the pelvic bones free from the cleithra; an adipose dorsal fin * Quoted in Harry, 1952. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 11 is typically present, the premaxi 1laries exclude the maxi 1laries from the gape, the gill openings are wide, the pectoral arch is attached to the skull by a forked post­ temporal, there is no mesocorocoid, the lateral centra are co-ossified with the arches, the air bladder is small or absent, and the ovaries are provided with oviducts.' *Prior to Regan's definition, the ordinal level of these fish was variously and correctly or incorrectly referred to as Cyprini (Bonaparte 1832-41, 1846), Physostomi (Mllller, 1944; Gunther, 1864, 1880, 1887), Isospondyli (Gill, 1872; Jordan, 1905), Malacopterygii (Gill, 1893, Goode and Bean, 1895a, b), Teleostei (Boulenger, 1904) and Haplomi sensu Gill (Gregory, 1907). The name 'Iniomi' ('Iniomes') was first proposed by Gill in 1884 to describe Sternoptychidae and its near relations. ^However between 1896 and 1940 'Iniomi' was commonly used to describe the ordinal level (Jordan and Evermann, 1896; Regan, 1909, 1911, Jordan, 1923; Parr, 1928, 1929) and Berg (1940) referred to the Iniomi as Scopeliformes. Most of the above works placed emphasis on osteological characters and sometimes with very little research (Harry, 1952). Harry (op cit) grouped iniomous fish into three orders based on his research on anatomy and classification: 1. Iniomi (with suborders Myctophoidea (Aulopodoidea) and Alepisauridea (Paralepidoidea). 2. Cetunculi. 3. Ateleopodes In recent times iniomous fish (Iniomi) have been referred to as Myctophiformes (Gibbs, 1959; Golvan, 1962; Mead, 1966; Okiyama, 1974; Masuda, Araga and Yoshino, 1975; Sulak, 1977) though in a general classification of teleostean fish using modern concepts of phyletic classification, Greenwood, Rosen Weitzman and Myers (1966) assigned iniomous fish to Salmoni- formes sensu lato. The order Myctophiformes is frequently * See Harry 1952. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 12. divided into two suborders, Myctophoidea/Myctophoidei and Alepisauroidea (Mead, 1966; Okiyama, 1974). Markle, Scott and Kohler (1980) however put the family Synodontidae under order Aulopiformes. The familial classification of lizardfish has not been without a diversity of names either. The *family name of this group of fish has been variously given as or placed under Scopelidae (Bonaparte 1832-41, 1846; Gunther 1864, 1880, 1887; Macleay, 1882; Day, 1878-88; Boulenger, 1904; Gregory, 1907; Goodrich, 1909), Scopelini (Mtlller, 1844), Aulopidae (Cope, 1871), Synodontidae **(Gill, 1862, 1872, 1893; Goode and Bean, 1895a, b; Jordan and Evermann, 1896; Ogilby. 1897; Jordan, 1905, 1923; Regan, 1909, 1911, 1929; Parr, 1929; McCulloch, 1920-22, 1927, 1929-30, 1934; Romer, 1945; Munro, 1961; Greenwood et al, 1966; Nielson, 1973; Linberg, 1974; Masuda, Araga and Yoshino, 1975; Sulak, 1977; Coleman, 1980), Alepidosauridae (Goodrich, 1909), Myctophidae (Parr, 1928), ***Synodidae (Ogilby, 1910a; Berg, 1940; Golvan, 1962; Dutt, 1973; Rao, unpublished), ***Sauridae (Rao, unpublished). 2.1 The Family Synodontidae According to the ^Zoological Code, 'Synodontidae', first proposed by **Gill (1862), is the correct etymological name of the family of lizardfish. Though the type genus of the lizardfish family is Synodus (Gronow) Scopoli, 1763, its family name (Synodontidae) is based on Synodontis Cuvier, 1817, the type genus of the family of African freshwater catfish (now Mochochidae). The discrepancy is due to a once held * See Harry (1952). ** Established as Synodontoidae in 1861 and amended to Synodontidae in 1872. *** Also see Linberg (1974) p. 279. + Article 28, p. 81 of the Bulletin of Zoological Nomenclature Vol. 31, 1974. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 13. belief that the two groups of fish were closely related as reported by Gunther in 1864 - 'These *fishes show a great resemblance in the family characters to the Siluroids; they might be called the marine Siluroids.1 Thus some taxonomists (e.g. Dutt, 1973) have persisted with Synodidae to indicate the difference in stem of the family name: Synodus and Synodidae as against Synodus and Synodontidae. The family Synodontidae appears to have evolved from the Aulopidae (Harry, 1952; Sulak, 1977). Synodontidae, a family of small or moderate size fish with elongate and almost cylindrical bodies with an adipose fin are found in tropical and warm temperate seas in all oceans, some at great depths. The head is lizard-like and the mouth is large and terminal. Teeth numerous, long, sharp, usually recurved and in rows, visible even when mouth is closed. At present there are about forty recognized species in the family Synodontidae. The number of genera however depends upon individual opinion as to the relationship between the genera and the many myctophiform families. Norman (1935) and Sulak (1977) recognize five genera: Harpadon Lesueur, 1825a; Saurida Cuvier and Valenciennes, 1849; Bathysaurus Gunther, 1878; **Synodus Gronow, 1763 and Trachinocephalus Gill, 1861, whereas Harry (1952) and Marshall (1964) exclude Harpadon and Bathysaurus from the family. Synodus and Trachinocephalus are closely allied and rather advanced with respect to the more primitive Saurida, Harpadon or Bathysaurus (See Sulak 1977 pp. 53-60). Sulak (1977) revised Synodontidae as follows: Family Synodontidae Gill, 1862 Subfamily Harpadontidae Jordan, 1923 Genus Harpadon Lesueur, 1825 (Incl. Peltharpadon Fowler, 1934) * i.e. Iniomous fish sensu lato ** Cressey (1981) included Xystodus Ogilby 1910 in Synodus University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 14. Genus Saurida Valenciennes, 1849 Subfamily Bathysauridae Genus Bathysaurus Gunther, 1878 (Incl. Macristium Regan, 1911) Subfamily Synodontinae Gill, 1862 Genus Synodus Gronow, 1763 (Incl. Xystodus Ogilby, 1910) Genus Traohinocephalus Gill, 1862. Whitley (1937) proposed six new subgenera for the genus Synodus: Negotirusy New tons cottia, Austrotirus3 Allouamia, Exotirichthys and Esosynodus, on the basis that 'the key characters given by Norman being sufficiently diagnostic for their definition'. In 1935, Norman undertook a major review of the genera Synodus3 Traohinocephalus and Saurida and reported 16, 1 and 9 species respectively from the warmer parts of the Atlantic and Indo-Pacific Oceans. Since the publication of Norman's (1935) monograph, several new species have been described from the Indo-Pacific region and in one instance Waples (1982) removed Saurida nebulosa from the synonymy of S. gracilis. Several reviews have also been attempted usually on regional basis and very often with only the literature as the source of data (e.g. Matsubara and Iwai, 1951; Gibbs, 1959; Anderson, Gehringer and Berry, 1966; Shindo, 1968; Shindo and Yamada, 1972; Yeh and Liu, 1973; Rao, 1977; Yamada and Ikemoto, 1979). Because of the widespread range of lizard­ fish and their similarities, there have been several instances of misidentifications, synonymies and also nomenclatural errors (e.g. Matsubara and Iwai, 1951; Okada and Kyushin, 1955; Liu and Tung, 1959; Shindo and Yamada, 1972; Rao, 1977). These errors and synonymies have arisen through lack University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 15. of awareness of previously published names (e.g. S. argentea Macleay, 1882 vrs S. micropectoralis Shindo and Yamada, 1972), or through nor>appreciation of the amount of variation existing within lizardfish (e.g. S. filamentosa Ogilby, 1910 vrs S. wanieso Shindo and Yamada, 1972) or sometimes as a result of lack of sufficient specimens. Trachinocephalus has been recognized as a monotypic species (Norman, 1935; Harry, 1952; Sulak, 1977; Cressey, 1981). On the contrary, Saurida consists of several morphologically cryptic species. The classification of Saurida has proved difficult using conventional morphological and anatomical characters - so many of these characters have limited value in establishing taxonomic relationships because of their overlapping ranges among the species. The difficulty is compounded by considerable intra-specific variability; the most similar and misidentified pairs and the triad being - 1. S. undosquamis - S. australis 2. S. tumbil - S. argentea 3. S. filamentosa - S. wanieso 4. S. flamma - S. gracilis - S. nebulosa In a biochemical and morphological review of Hawaiian Saurida, Waples (1982) reported the co-existence of three species where only one species, S. gracilis (Quoy and Gaimard) 1824, was hitherto recognized. The following are currently the recognized Indo-Pacific species of Saurida and Trachino­ cephalus : 1. Saurida tumbil (Bloch) , 1975 2. S. gracilis (Quoy and Gaimard) , 1824 3. S. elongata (Temminck and Schlegel), 1846 4. S. undosquamis (Richardson), 1848 5. S. nebulosa Valenciennes, 1849 6. S. filamentosa Ogilby, 1910 7. S. longimanus Norman, 1939 8. S. isarankurai Shindo and Yamada, 1972 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 16. 9. S. micropectoralis Shindo and Yamada, 1972 10. S. wcmieso Shindo and Yamada, 1972 11. S. flamma Waples, 1982 and 12. Traohinocephalus myops (Bloch and Schneider), 1801. In this study, a review of the genera Saurida and Traohinocephalus is attempted using conventional morphological/ anatomical techniques and exploratory electrophoretic analysis of general and selected proteins. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 17. 2.2 MATERIALS AND METHODS 2 .2.1 Electrophoretic Studies 2.2.1.1 Species Samples Fish samples were obtained from the following areas (number of specimens and length (mm) range in brackets): Remarks a) Moreton Bay (S.E. Queensland) Trawled from 10- S. tumbil (20,180-250) 40m on muddy/sandy S. undosquamis (20,150-220) flats. T. myops (4,130-150) b) Off Moreton Bay (S.E. Queensland) Trawled from 80- S. filamentosa (15,250-400) 200m by Bob T. myops (15,80-190) Sanderson, Skipper of the 'Rex Andria' c) Hervey Bay (Queensland) Taken on a hook by S. argentea (2,169-214) Clive Keenan from estuarine shallow water in a sand bank S. australis (30,230-300) Trawled by Greg Campbell between Lady Elliot and Fairfax Islands, east of Bundaberg d) North-West Shelf of Australia Obtained by the S. argentea (30,200-300) Crewof FRV 'Soela' S. filamentosa (12,200-300) trawling on coral, S. longimanus (2,108-125) muddy and/or sandy T. myops (8,80-160) bottoms; 50-120m. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 18. e) Lizard Island (Great Barrier Reef) Obtained on coral S. nebulosa reef by Dr. Hugh Sweatman. All fish samples were put on ice or were stored in a freezer prior to transportation to the laboratory. In the laboratory, the fish were weighed, measured for length, sealed in polythene bags and then frozen at -20 to -30°C until required. 2.2.1.2 Preparation of tissue extracts Tissue extracts were prepared from the eye, liver and white muscle - the muscle was excised from between the dorso-medial line and above the lateral line but just anterior to the vertical plane of the origin of the dorsal fin. Between l-3g of tissue in an approximately equal volume of grinding *buffer was ground with an Ultra-Turrax tissue grinder. The homogenized tissue was centrifuged at about 5°C at 14250-14500 rpm (i.e. about 25300g**) for 20 minutes in Sorvall Centrifuge. The supernatant was decanted into 10 ml glass tubes and centrifuged for a further 60 minutes. The supernatant was again decanted into smaller glass or plastic tubes and stored in a freezer at -20 to -30°C (at the Zoology Department, University of Queensland) or at -70°C (when working at CSIRO (Cleveland Laboratories)). The enzymes, at these freezing temperature, retained their resolution for at least six months. However samples could not be frozen and thawed more than three times without considerable loss of resolution. * Grinding buffer - 1.21g Tris:0.37g EDTA (Na^):0.00153g NADP per 1 , ph adjusted to 6.8 with HC1. ** Acceleration due to gravity. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 19. The method involving an horizontal electrophoresis on pore gradient polyacrylamide gels and staining with o.2% Coamasie Blue R as described by Gahne, Jenuja and Crolmus (1977) was employed in the analysis of general proteins. Specific enzyme analysis followed Redfield and Salini (1980) and a CSIRO (Cleveland) Electrophoresis Manual (unpublished). A variety of buffers were tried in the preliminary analysis to find the best buffer-enzyme systems with regard to enzyme resolution but concurrently with the analysis of inter-generic differences and similarities using samples of S. tumbil> S. undosquamis and T. myops. In the main analysis, extracts of all the species were examined. However all extracted samples, including the only samples of S. longimanus and 5. nebulosa were lost due to prolonged thawing and a resultant loss of resolution by the enzymes because of power failure over a weekend. Work continued on only readily available species and the following 24 enzymes were surveyed: 1. Aspartate aminotransferase (AAT) 2. Alcohol dehydrogenase (ADH) 3. Aconitate hydratase (AH) 4. Adenylate kinase (AK) 5. Creatine kinase (CK) 6. Esterase (EST) 7. Fructose biphosphate aldolase (ALD) 8. Gylceraldehyde-phosphate-dehydrongenase (GDH) 9. Glutamate dehydrongenase (GDH) 10. Glycerol-3-phosphate dehydrongenase (G-3-PDH) 11. Glucose-6-phosphate dehydrogenase (G-6-PDH) 12. Glucose phosphate isomerase (GPI) 13. Isocitrate dehydrogenase (IDH) 14. Lactate dehydrogenase (LDH) 15. Malate dehydrogenase (MDH) 16. Malate dehydrogenase (NADP)/Malic enzyme (ME) 17. Mannose phosphate isomerase (MPI) 18. Phosphogluconate dehydrogenase (PGDH University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 20. 19. Phosphoglucomutase (PGM) 20. Pyruvate kinase (PK) 21. Sorbitol dehydrogenase (SDH) 22. Superoxide dismutase (SOD) 23. Umbelliferyl esterase (UMB) 24. Xanthine dehydrogenase ( XDH) The following *buffers were employed in gel preparations and electrode set-ups: Buffer Chemical Composition Continuous buffer CAAPM Citric acid H^O Aminopropylmorpholine CAEA Citric acid 1^0 N - (3 aminopropyl) - diethanolamine EBT Tris : Boric Acid : EDTA (Tetrasodium salt) PC-1 Citric acid (monohydrate) : K^HPO^ TC-1 Tris Citric acid H^O TC-2 Tris Citric acid E^O TC-4 Tris Citric acid TECB Tris EDTA : Citric acid Boric acid Discontinuous buffer POULIK - gel; Tris Citric acid electrode; Boric acid NaOH. The approximate voltages and running times of the buffer systems were as follows: Buffer Starting Starting Running Voltage (V) Current (mA) Time (hrs) CAAPM 200 45 4 CAEA 200 27 4 EBT 300 51 4 * See Redfield and Salini (1980) (and CSIRO (Cleveland laboratories Electrophoresis Manual (unpublished)) for details. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 21. PC-1 100 65 5i POULIK 140-250 ** 40 5 TC-1 175 55 4 TC-2 100 58 6 TC-4 150 55 4 TECB 200 60 7 All species were examined in side-by-side comparisons on the gels so as to eliminate variations due to treatment. A number of photographs were taken of each gel as the bands developed. Illustrations of patterns were also drawn and distances travelled by the various bands from the origin noted in many cases. Other data obtained included relative enzyme activity (presence or absence) and comparison between the tissue types. 2.2.2 Morphological/Anatomical Studies 2.2.2.1 Species Samples This study was based on ***specimens collected from various localities or examined on loan from institutions and museums and on the literature. Several specimens, and where possible holotypes or paratypes, of each species were examined. Various characters were studied and where necessary or possible photographs were taken or illustrations drawn. All measure­ ments are straight-line distances made either with wooden or plastic rulers and recorded to the nearest millimeter, or with dial calipers with an accuracy of i().05mm. Excessively distorted specimens were measured when they were the only specimens available. ** Constant Current. *** Details provided under the appropriate species University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 22. 2.2.2.2 Measurements, Counts and definitions (See Fig 2.1) Measurements are given as proportions of a) standard (S.L.) and/or b) head (H.L.) length unless otherwise stated. Standard length, as described previously in the main introduction, was measured by placing the fish on a measuring board with the tip of the snout touching the head block and measuring the perpendicular distance between the snout tip and the crease of the caudal fin (i.e. the posterior margin of tbehypural bones). The crease was located by flexing the caudal fin at right angles to the body’s axis. Head length is the distance between the tip of the snout and the posterior extension of the fleshy margin of the opercle at the level of the upper edge of the pectoral fin base. Snout length stands for the distance between the tip of the snout and the anterior margin of the orbit, whereas eye diameter is the horizontal distance between the anterior and posterior borders of the orbit. The length of a *fin or ray was measured as the distance between the tip and the origin or base. Prepelvic, predorsal, preanal, prepectoral and preadipose lengths denote distances between the tip of the snout and the origin of the designated characters or features. **Scales on the dorso-medial line were counted as follows: predorsal - the number of scales between occiput and origin of dorsal fin (including the anterior-most rounded scale which is almost covered with skin as well as scales in front of dorsal origin. * Dorsal fin measured when laid back. ** Scales bisected by the dorso-medial (straight) line running from occiput to caudal fin. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 11 1. Preorbital/snout 2. Eye diameter 3. Postorbital 4. Head 5. Predorsal 6. Dorsal fin base 7. Dorsal fin 8. Interdorsal S. Peduncle/postadipose 10. Preadipose 11. Standard 12. Premaxillary 13. Pectoral fin 14. Prepelvic 15. Pelvic 16. Dorsal fin to anal fin. 17. Anal fin base 18. Interventral 19. Depth of body University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 23. interdorsal - those scales between the last ray of the dorsal fin and adipose fin. preadipose - scales between the adipose and caudal fins. The number of scales in lateral line denotes the count of all pored scales between operculum and the crease at the base of the caudal fin. Scales after the crease (all elongated and some pored) were ignored. In the transverse scale row count the number of upper rows was counted from the origin of the dorsal fin down to the lateral line; the number of lower rows was counted from the origin of the anal fin, upward to the lateral line. In either transverse row counts, the lateral line scale itself and the modified scales at the base of the dorsal or anal fin were not counted. Scale-pockets/traces when visible were counted where scales were lost. With the fin ray counts, the last ray of the dorsal or anal fin which is divided to its base was counted as one element. Fin rays were counted more easily under a magnifying glass (or a microscope as with very small specimens,. ) The premaxilla length was taken as the straight line from the symphysis to the tip of the snout. All other measurements or counts are as denoted by the names or terms as illustrated in Fig. 2.1. In addition to the number of rows in the palatine teeth, the shape of the teeth bands and the configuration of the teeth in the bands and on vomer were studied in detail - many specimens were decapitated and also cut at only one articulation of the premaxillary to angular. These were dried out gradually in the laboratory to reduce shrinkage and later dried out in the sun after an exploratory confirmation of the configur­ ation of the teeth with 'wet' specimens. The configurations before and after drying were compared and ̂ ecimens with least shrinkage were photographed. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 2 4 . Coloration was recorded from both fresh and preserved specimens in a range of sizes or as reported in the literature. Unless otherwise noted for the species, the ecological information presented is based on personal observation in the field and field notes of the collectors and museums or institutions. Vernacular names were taken from Grant (1972, 1978), Marshall (1964), Munro (1961), Coleman (1980) and Waples (1982). ^Synonymies are, with few exceptions, those mentioned by McCulloch (1929-30), Norman (1935) and Anderson, Gehringer and Berry (1966). 2.2.2.3 Abbreviations of Museums/Institutions AMS - Australian Museum, Sydney BMNH - British Museum (Natural History), London BPBM - Bernice P. Bishop Museum, Honolulu DPPFD - Fisheries Division (Department of Primary Production), Darwin, Northern Territory. KFR - Kanudi Fisheries Research Museum, Konedobu, Papua New Guinea QM - Queensland Museum, Brisbane SI - Smithsonian Institution, Washington, D.C. SOSC - Smithsonian Oceanographic Sorting Centre USNM - United States National Museum ZUMT - University of Tokyo, Zoology Department, Tokyo * Only the original papers or those containing valuable information are quoted when synonymies are extensive. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 25. 2.3 RESULTS 2.3.1 Electrophoresis (Biochemical Systematics). 2.3.1.1 General Proteins (Plates 1,2A, B) General protein patterns were examined in eight species: S. filamentosa, S. australis, S. undosquamis> S. tumbil, S. argentea, S. longimanus, S. nebulosa and Traohinocephalus myops by separation of muscle and liver extracts on polyacrylamide gels. The examination revealed similar but distinct electrophoregrams which are consistent with patterns of morphological similarities and differences between the species. Individual electrophoregrams also exhibited intra-specific variations. 2.3.1.2 Specific Proteins Starch gel electrophoresis was employed for study of muscle, liver and eye enzymes. Data were obtained for relative enzyme activity (i.e. rate of staining and/or intensity of stain, presence or absence in some cases), inter- and intra-species comparisons. The following observations were made: Whereas some tissue specificity of enzymes was noted, the eye extracts showed consistently low activity levels for many of the enzymes examined, probably due to low concentrations. Poor resolution was noted in some enzymes and only extracts giving best resolutions are discussed here - enzymes with poor resolution are given cursory treatment. In some instances achromatic bands occurred with the desired patterns. The achromatic bands were due to dehydrogenase(s), probably ADH or SOD reacting with 'sufficient alcohol present as a contaminant of the chemical used’ - Ferguson (1980). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 25a. TABLE 2.0: Legend for plates and figures. a Saurida australis f - S. filamentosa 1 S. longimanus m Trachinocephalus myops n Saurida nebulosa r S. argentea t S. tumbil u S. undosquamis University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh m u s c l e Plate 1 Comparison of general protein patterns of Saurida (group B). Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh A B Plate 2* Comparison of general protein patterns of Saurida (group A) and Trachinocephalus. Legend follows Table 2.0 (p.25a). *Two different exposures of film same gel. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 26. 2.3.1.3 Generic differences and similarities Twenty *enzyme-buffer systems were surveyed for genetic differences and similarities in mobilities of speci­ mens of **Saurida and ***Traohinocephalus. Specific mobility differences, presumably reflecting generic differences, were found in AK (CAEA-m; Plate 3, Fig. 2.2A), IDH (CAAPH-1; Fig. 2.2B), MPI (PC-1-1,e; Fig. 2.2.C), GPI (TC-2-t), MDH (CAEA-t; Plate 4) and ME (CAEA-t). Similarities in band mobilities in Saurida and Traohinocephalus were found in the following enzyme-buff er systems: AAT (CAEA-t; Fig. 2.2D), ADH (TC-2-m,l; Fig. 2.3A), GAPDH (CAEA-t; Plate 5, Fig. 2.3B), GDH (TC-2-m; Fig. 2.3C), PGDH (CAEA-t; Plate 6, Fig. 2.3D), SDH (PC-l-m; Fig. 2.4A), SOD (POULIK-1, e; Fig. 2.4B), UMB (EBT-t ; Fig. 2.4C), XDH (EBT-m,1; Fig. 2.4D), EST (POULIK-t; Plate 7), G-3-PDH (TC-l-t) , LDH (CAAPM-t; Plate 8), PGM (TC-l-t) and PK (TECB-t). 2.3.1.4 Inter- and intra-specific variation The electrophoresis runs involved two periods of analyses (January - March 1982, July - September 1982) with regards to the availability of the species. To avoid the repetition of names, the species analysed during the two periods are referred to as groups: Group A (January - March 1982) - S. tumbil, S. australis, S. undosquamis, S. filamentosa, S. argentea, S. nebulosa, S. longimanus and T. myops. +Group B (July - September 1982) - all species in Group A except S. nebulosa, S. longimanus and T. myops. * Tissues: m - muscle, 1 - liver, e - eye, t - all three tissues. ** 3 each of S. tumbil and S. undosquamis *** 2 specimens of T. myops + Species available after power failure caused loss of stored extracts. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 3: AK (CAEAL muscle 1 m { PLATE 4: MDH (CAEA), Liver Plates 3 and 4: Inter-generic comparison of zymogram patterns of AK and MDH of saurida and Trachinocephalus, Legend follows Table 2,0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig.2.2 : Diagrams of the zymogram patterns of SOme enzymes Of Saurida ond Traahino- aephaius. Legend follows Table 2.0 (p.25a) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 5: GAPDH (CAEA) ii if I L ~ l i v e r i I t h lml Plates 5 and 6: Inter-generic comparison of zymomgram patterns of GAPDH and PGDH of Saurida and Traohinocephalus. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.3 : Diagrams of the zymogram patterns of some enzymes of saurida and Trachinocephalus. Legend fOllOWS Table 2.0 (p.25a.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.4: Diagrams of the zymogram patterns of some enzymes of saurida. and Traohinocephalus. Legend fOllOWS Table 2.0 (p.25a) * Achromatic bands University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 7: EST (POULIK) Plate 8: LDH (CAAPFI) Plates 7 and 8: Intergeneric comparison of zymogram patterns of EST and LDH of Sccurida and Trachinocephalus. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 27- 2.3.1.4.1 Observations in Group A ALD (EBT) Though the ALD zymogram indicated good separation of species, its resolution was very poor and cannot be discussed any further. CK (EBT). Fig. 2.5A CK stained well in muscle and eye. Each species, except S. tumbil and S. argentea had distinct eye (CK-B2) - muscle (CK-A2) array of bands. EST (POULIK). Fig. 2.5B Liver: Two forms (fast and slow) were detected in the species. S. australis and S. filamentosa were single banded and most anodal, whereas S. nebulosa had the lease anodal form. S. argentea, S . undosquamis and S. longimanus shared identical double banded patterns. EST was not detected in muscle and was poorly resolved in eye. GAPDH (CAEA). Fig. 2.6A Muscle: anodal and cathodal forms were detected in the specimens. Each species appears to exhibit a distinctive array of bands. The most anodal form was detected in S. argentea and the most cathodal in S. tumbil and S. australis. No activity was apparent in S. longimanus. There was poor resolution in both liver and eye extracts. G-3-PDH (TC-1). Plate 9, Fig. 2.6B Muscle: each species exhibited a distinctive array of bands. S. longimanus, S. nebulosa and S. undosquamis had a common (most) anodal band. The least anodal forms were found in S. filamentosa, S. undosquamis, S. australis and T. myops. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.5: Diagrams of the zymogram patterns of CK and EST of species of Saurida Qfld Trachinocephalus. Legend follows Table 2.0 (p.25a) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 9: G-3-PDH) (TC-1) £2.- ELA-qtj Plate 10: LDH (TC-1) £L/[-qx; Plate 9 and 10: Comparison of zymogram patterns of G-3-PDH and LDH of species of Sccuvida and Trachinooephalus. Legend fOllOWS Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 28. Similar patterns were discernible in eye and liver extracts though poorly resolved. LDH (TC-1). Plate 10 LDH was detected in all tissues examined. The mobilities of the species reflected species differences and similarities. MDH (TC-1). Plate 11 MDH was detected in all tissues, each of the eight species exhibiting distinctive phenotypes for the various isozymes. SOD (POULIK). Plate 12 and Fig. 2.7 S. tumbil was found to be polymorphic for liver SOD, an achromatic band pattern characteristic of control by two co-dominant alleles. No polymorphism was detected in the eye isozyme of S. tumbil and neither was any difference found between the mobilities of the two isozymes. SOD was not detected in muscle. As illustrated in Fig. 2.7, the fastest band of 5. tumbil is homologous with bands in S. argentea and S. longimanus. 2.3.1.4.2 Observations in Group B The species were examined by side-by-side comparisons with S. filamentosa interspersed between species as control for enzyme activity - four specimens were scored for each species per gel. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 11: MDH (TC-1) Plate 12: SOD (POULIK) t r m uscie & X - ELK - S'f • 1 Plate 11 and 12: Comparison of zymogram patterns of MDH and SOD of saurida and Traehinocephalus. Legend fOllOWS Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 801 6 0 - 4 0 2 0 - F~ r u a f L n m muscle * K i v e r . eye -» Fig, 2.7 : A diagram of the zymogram patterns of SOD of species of saurida and Traohinocephalus. Legend follows Table 2.0 (p.25a) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 29. The species exhibited similar patterns in AK (muscle; Plate 13, Fig. 2.8A) as well as in MDH (muscle; Plate 14, Fig. 2.8B) They each had three zones of activity - the first zone of AK was broad; S. undosquamis stained faster than the others. Observations on other enzymes (with buffer in brackets) are as follows:- AAT (CAEA) Two forms of AAT were present in the species: anodal and cathodal forms. In both muscle (Fig. 2.8D) and liver (Plate 15, Fig. 2.8C) S. undosquamis shared same identical bands with S. australis. S. tumbil and 5. argentea shared identical anodal and cathodal forms in the liver but different liver forms. ADH (EBT) Fig. 2.9A Muscle: S. undosquamis had a fast band, whereas the other species had similar slow bands. The intensity of the stain was low indicating low enzyme activity. AH (TECB) Muscle (Fig. 2.9B): There were two types of mobilities - fast in S. filamentosa, S. australis and S. undosquamis, and slow in S. tumbil and S. argentea. Liver AH showed up only in S. tumbil (Fig. 2.9C). EST (EBT) Liver (Plate 16): S. australis, S. filamentosa and S. undosquamis shared distinctly different patterns from S. tumbil and S. argentea though resolution was poor. GDH (CAAPM) Muscle (Plate 17): All the five species except S. undosquamis had an identical band mobility. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh P l a t e 13. Plate 14. Plates 13 and 14: Comparison of zymogram patterns of AK and MDH of 5 species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh A k (m u s C L 6) +r^ 2 0 - - f i a i f i_u_i f i__t__l_Lj__r. iJ- Fig, 2,8 : Diagrams of the zymogram patterns of some enzymes of species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 15 Plate 16 Plates 15 and 16: Comparison of zymogram patterns of AAT and EST of 5 species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh +f ra4-*-___-__-__-______________ A D_H ( m u s c l e )mm — —» ** “ ----- .. 1 ..f 1 a Ifl u 1 f 1 t I f I r Ifau t + A H ( m u s c l e ) 4 0 - a a o o a a a o a a a o a a 2 0 - 0 f I a 1 f I u i f i t Fig. 2.9 : Diagrams of the zymogram patterns’ of some enzymes of species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 17 f Plate 18 Plates 17 and 18: Comparison of zymogram patterns of GDH and IDH of 5 species of Saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 30. G-6-PDH (CAEA) Liver (Fig. 2.10A): This enzyme appears as a double­ banded pattern except in S. tumbil (3-banded). S. filamentosa, S. australis and S. argentea had identical patterns. The second band of S. argentea indicated reduced activity. GPI (EBT) Three types of mobility were detected in the muscle (Fig. 2.10B) - fast in S. filamentosa, intermediate in S. filamentosa and S. argentea, and slow common to S. australis, S. undosquamis and S. tumbil. IDH (CAAPM) Muscle (Plates 18, Fig. 2.10C): S. tumbil exhibited two types of mobility - fast and slow. It shared the fast with S. filamentosa, S. australis and S. undosquamis. The slow band was in common with S. argentea. The liver isozyme patterns were different in S. australis, S. undosquamis and S. tumbil but was not detected in S. filamentosa and S. argentea. LDH (CAAPM) Muscle (Plate 19A): The zymograms of the species except S. undosquamis were more or less identical and appeared as a 4 or 5 banded pattern. The zymogram of S. undosquamis was very distinct. This isozyme stained intensely and quickly. Liver (Plate 19B): This isozyme had species specific patterns. These stained less intensely and slowly than the muscle isozyme except in S. tumbil. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh A G -6-P DH ( I iver) 10- - - = =_r------- f 1 a f 1 u 1 f 1 t 1 f 1 r 1 f a u t r B G PT (muscle) 20- --- - --- ' ----- ' ---- n_ f 1 a 1f 1 u 1 f 1 t 1 f 1 r 1 f a u t r| C I DH (muscle) 10- 1 f 1 a 1f 1 u 1 1 t 1 1 r 1 D M DH( I iver) | 20- U" mmmm ' f .1. a . 1f 1 u 1 f 1 t 1 f 1 r 1 f a u t r! Fig. 2.10: Diagrams of the zymogram patterns of some enzymes of Saurida species. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 19A Plate 19B Plate 19: Comparison of zymogram patterns of muscle (A) and liver (B) LDH of 5 species of Saurida. Legend follows Table 2.0 (p,25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 31. MDH (CAEA) Liver (Plate 20 and Fig. 2.10D): S. filamentosa and S. undosquamis, S. australis and S. argentea shared 3 and 4 banded patterns respectively. S. tumbil was polymorphic with 3 and 5 banded forms. ME (CAEA) Muscle (Plate 21 and Fig. 2.11A): Two zones of enzyme activity were detected. One zone was common to all the species except S. undosquamis. Though S. undosquamis shared its first zone with S. filamentosa and S. australis, its second zone of activity consisted of three sub-units. The liver isozyme stained very slowly. S. tumbil had the most anodal form. MPI (PC-1) Muscle (Fig. 2.11B). MPI stained faintly in muscle and three forms of mobility were detected - fast in S. filamentosa, S. undosquamis, S. tumbil and S. argentea; intermediate in S. australis and slow also in 5. undosquamis. Liver: MPI stained intensely, especially in S. tumbil and S. undosquamis. There were three types of mobility - fast in S. filamentosa, intermediate found in all but S. argentea which also had the slow band (Plate 22) . The achromatic bands are probably due to dehydrogenase(s) as explained previously in this chapter. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 20 Plate 21 Plates 20 and 21: Comparison of zymogram patterns of MDH and ME of 5 species of Saurida. Legend follows Table 2.0 (p.25a) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh + C____________________P GDH 20 - F l a l f l u l F l t l f l r l F a u t n Fig. 2.11: Diagrams of the zymogram patterns of some muscle enzymes of saurida species. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 22 Plate 23 Plates 22 and 23: Comparison of zymogram patterns of MPI and PGDH of 5 species of saurida. Legend follows Table (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 32. PGDH (CAEA) Muscle (Fig. 2.11C): The enzyme was detected as a single or double (in S. undosquamis ) banded form. All the species except S. tumbil showed reduced activity (i.e. low intensity or stain) by the enzyme. The liver isozyme was only detected in S. undosquamis and S. tumbil - these two species exhibited a high staining rate indicating high activity of PGDH (Plate 23). PGM (CAAPM) Migration was only cathodal in muscle extracts, with the species exhibiting distinct array of bands. (Plate 24A). The liver isozyme was only detected in 5. undosquamis and S. tumbil (Plate 24B) . PK (CAAPM) Muscle: Plate 25A shows a 3-banded pattern with one major zone of activity. Liver: Isozyme poorly resolved in S. undosquamis and S. tumbil and not detected in the other species (Plate 25B). SDH (PC-1) Liver (Plate 26): Activity zones of this enzyme were achromatic and of three mobility types. The least anodal forms were found in S. australis3 S. tumbil and S. argentea. S. undosquamis had the most anodal form and was well separated from the other species. Resolution of muscle SDH was poor. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 24A Plate 24B Plate 24: Comparison of zymogram patterns of muscle (A) and liver (B) PGM of 5 species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 25A Plate 25B Plate 25: Comparison of zymogram patterns of muscle (A) and liver (B) PK of 5 species of Saurida. Legend follows Table 2.0 (p.25a) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 26 Plate 27 Plates 26 and 27: Comparison of zymogram patterns of SDH and SOD of 5 species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 33. SOD (EBT) Liver (Plate 27): The species exhibited the same patterns as with SDH except that S. argentea was found to be polymorphic for two co-dominant alleles. UMB (EBT) The bands were only visible under ultraviolet light - all the species except S. filamentosa shared a common band. S. filamentosa had the most anodal band. XDH (TECB) Muscle: All the species share a common band except S. undosquamis (Plate 28A, Fig. 2.11D). The liver isozyme (Plate 28B) is poorly resolved and appears as a single or double banded pattern. The achromatic zones in the zymograms are due to some other dehydrogenase(s) probably ADH or SOD. In the main polymorphisms characterized by two co-dominant alleles, were noted in a number of the following monomeric proteins: enzyme locus species AAT liver S. australis S. undosquamis GPI muscle S. filamentosa IDH muscle S. tumbil MPI liver S. australis S. undosquamis MPI muscle S. undosquamis The only dimeric protein found to be polymorphic was SOD in S. tumbil and S. argentea. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh PLATE 28A Plate 28B Plate 28: Comparison of zymogram patterns of muscle (A) and liver (B) XDH of 5 species of saurida. Legend follows Table 2.0 (p.25a). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 34. Though the results suggest considerable intra-specific variability, each individual could be assigned unequivocably to one of the species - the species apparently exhibit distinctive *phenotypes, notably, for CK (M,E), GAPDH (M), G-3-PDH (M ,E), SOD (L,E) and to varying degrees for LDH (M,L ,E), IDH (M ,L), PGM (L,E), AH (L), SDH (L), EST (L) and AAT (M,L,E). S. undosquamis and S. australis shared common electromorphs at AAT (M,L) AK (M), MDH (M) and IDH (M) but differed in ADH (M), ME (M), MDH (L), G-6-PDH (L). PK (M) , MPI (L,M) and XDH (M). Likewise though S. tumbil and S. argentea were identical or similar in AAT (L), AK (M) , MDH (M), ME (M), PK (M), and MPI (L) , they differed in ADH (M), GPI (M), AAT (M) and MDH (L). 2.3.2 Morphological Systematics 2.3.2.1 Generic characters Saurida: Two bands of palatine teeth on each side of palate. Vomerine teeth present or absent (see Plates 29-35 and Fig. 2.14). Nine pelvic fin rays, inner rays of which are subequal or slightly longer than outer rays **(Fig. 2.12C) Polyspecific genus. * M - muscle, E - eye, L - liver. ** Adopted from Anderson, Gehringer and Berry 1966. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 35. Traohinocephalus and Synodus: One band of palatine teeth on each side of palate. Vomerine teeth absent (See Plate 37). Eight pelvic fin rays, innermost rays distinctly longer than outer­ most rays, about two or more time longer *(Fig. 2.12 A ,B). Traohinocephalus: Anal fin base longer than dorsal fin base. Monotypic genus. Synodus - Anal fin base shorter than dorsal fin base. Polyspecific genus. 2.3.2.2 Genus Saurida Saurida: Cuvier and Valenciennes, 1849. 1849 Saurida: Cuv. and Val. Hist. Nat. Poiss. xii, 1849, p. 499; type species Salmo tumbil Bloch 1795, by subsequent designation of Jordan, Tanaka, and Snyder, J. Coll. Sci. Tokyo, 33(1), 1913: 53 fide Fowler, Bull. Amer. Mus. Nat. Hist., 70(2), 1936 : 1218. Body elongate, sub-cylindrical, fins with no spines. Scales cycloid and of moderate size. Lateral line straight except for a slight curvature above pectoral fin base. Head lizard-like, oblong and depressed with an antero- supra-orbital bony ridge just after second nostril. Nostrils approximate, the first fringed by a nasal flap. Eye of moderate size and nearer tip of snout than posterior edge of head. Eye partly covered by an anterior and posterior eyelid. Dorsal fin nearly in the middle of standard length. Adipose fin small and set opposite to posterior * Adopted from Anderson, Gehringer and Berry 1966. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.12: Ventral view of pelvic fins of Traohinocephalus (A), Synodus (B) and Saurida (C). (Adopted from Anderson, Gehringer and Berry 1966) Fig. .13: A,B - Snouts of Saurida species C,D,E - Nasal flaps of Saurida species G ,H - Diagrams showing lengths of pectoral fins of Saurida species. 1 = Saurida longimanus University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 36. half of anal fin. Pelvic fin base well behind pectoral fin base but inserted before dorsal fin. Tail tapering and covered with scales. Caudal fin bifurcate, pectoral falcate to subfalcate. Anal fin short. 3 or 4, and 5 scales row above and below lateral line respectively. Mouth terminal and bordered by very long styliform intermaxillary to which the thin maxillary is closely adher­ ent. Gape wide and more than half length of head. Teeth seriate; more or less regular, unequal in size, depressible, conical to sagittate (see Plate 29-35 and Fig.3,4A). Palatine teeth in two bands; outer is 2-4 rows, inner patch-like and pluriserial (see Fig. 2,14), Size of teeth in both bands increase inwardly. Vomerine teeth present or absent. Numerous fine caninoid teeth on tongue and branchial arches. Jaw lips with about 5-7 rows of inwardly slanting recurved teeth which increase in size and curvature towards mouth cavity. 13-16 branchiostegal rays, gill opening very wide. Opercular apparatus well developed, scaly as well as post­ orbital part of head. Gill membranes free, pseudobranchiae well developed. Fishes of moderate size inhabiting warm seas. Species cryptic and eleven are recognized in this study: 1 . Saurida argentea Macleay, 1882 2 . S. australis Castelnau, 1878-79 3. S. elongata (Temminck andSchlegel) , 1846 4. S. filamentosa Ogilby, 1910 5. S. flamma Waples, 1982 6 . S. gracilis (Quoy and Gaimard) , 1824 7. S. isarankurai Shindo and Yamada, 1972 8 . 5. longimanus Norman, 1939 9. 5. nebulosa Valenciennes, 1849 1 0 . S. tumbil (Bloch) , 1795 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 37. 11. S. undosquamis (Richardson), 1848 2.3.2.2.1 Important diagnostic characters of Saurida 2.3.2.2.1.1 Palatine teeth Three types of teeth configuration in palatine bands and on vomer were established - each epitomized by the *patterns in S. undosquamis, S. tumbil and 5. nebulosa respectively. 2.3.2.2.1.1.1 S. undosquamis type (Plate 29) Other band teeth usually in 2 and occasionally 3 rows anteriorly. Inner band with 5-7 rows of teeth. Bands widely separated. Vomer with or without teeth; if present usually on flanks or wings of vomer. Species - S. undosquamis, S. filamentosa (Plate 34), S. australis, S. longimanus, S. isarankurai (see Fig. 2.14). 2 . 3 . 2 . 2 .1.1. 2 S. tumbil type (Plate 30A) Outer band teeth in three or four rows. Inner band with 5-8 rows of teeth. Bands narrowly separated. Vomerine teeth present and central (see Plate 30B) Species - S. tumbil, S. argentea (Plate 32A, B) and S. elongata (see Fig. 2.14) 2.3.2.2.1.1.3 S. nebulosa type (Plate 31) Outer band with a short neck, distal aspect divergent. Outer band teeth in 2 or 3 poorly arranged rows, first three or four teeth of innermost row in neck of outer bank very long and pointed. Inner band consists of * See description of species for specific palatine patterns. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate Teeth configuration in palatine (p) bands and On VOmer (V) Of 5. undosquamis. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 30A Teeth configuration in palatine(p) bands and on vomer(v) of s. tvmbu. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 30B Teeth configuration in palatine(p) bands and on vomer (v) of s. tumbil University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 31. Teeth configuration in palatine (p) bands and on vomer (v) of s. nebulosa. X = long and pointed teeth. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.14: Diagrams of palatine (p) and vomerine (v) teeth configurations in saurida species A s. undosquamis B s. australis C s. filamentosa D s. longimanus E s. isarankurai F s. tumbil G s. argentea H 5. elongata I s. nebulosa s. flamma J s. gracilis University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 38. 3-4 poorly defined rows. Bands narrowly separated. Vomerine teeth present, set centrally or 2-4 teeth on the wings of vomer (v, see Plate 31) and deceptively * alligned with outer band. Species - S. nebulosa, S. gracilis and S. flamma (see Fig. 2.14) 2.3.2.2.1.2 Snout and nasal flap shapes Shapes of snout and nasal flap are also typified by S. undosquamis, S. tumbil and S. nebulosa. 2.3.2.2.1.2.1 S. undosquamis type (Fig. 2.13) Snout more or less rounded (Fig. 2.13B). Anterior margin of nasal flap small and slightly arched, the posterior margin flabellate, inner half, of which is more or less produced (Fig. 2.13C). Species - S. undosquamis, S. australis, S. filamentosa, S. longimanus and S. isarankurai. 2.3.2.2.1.2.2 S. tumbil type (Fig. 2.13) Snout pointed (Fig. 2.13A). Anterior margin of nasal flap domed or arched, the posterior margin broad and more or less flabellate (Fig. 2.13D). Species - S. tumbil, S. argentea and S. elongata. *Could be verified by slight probing and pushing - especially outward - with a seeker, viewed under a magnifying glass or microscope. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 39. 2.3.2.2.1.2.3 S. nebulosa type (Fig. 2.13) Snout more or less pointed (Fig. 2.13A). Anterior margin of nasal flap very narrow, posterior margin prolonged and spatulate (Fig. 2.13E). Species - S. nebulosa> S. gracilis and S. flamma. 2.3.2.2.1.3 Pectoral fin (Fig. 2.13) The length of pectoral fin is also a good diagnostic feature. In S. longimanus the pectoral fin is long, reaching beyond the origin of the dorsal fin; about 78-103% of H.L. (Fig. 2.13G1) Pectorals of S. undosquamis, S. filamentosa, S. australis, S. nebulosa> S. gracilis, S. flamma and S. isarankurai are also long. Though not as long as in S. longimanus, they reach at least, to the level of pelvic fin base; 46-75% of H.L. (Fig. 2.13G). Pectoral fin is short in S. tumbil> S. argentea and S. elongata. They were not found to reach beyond the pelvic fin base; 44-58% of H.L. (Fig. 2.13H). 2.3.2.2.1.4 Second and Third dorsal fin rays The second and sometimes third dorsal fin rays filamentous in S. filamentosa; the second ray on average being about six times as long as the last ray; range : 4.1-8.3. No other known species of Saurich possesses these filamentous dorsal rays. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 40. 2 .3.2 .2 .1 .5 Jaws The upper jaw is slightly longer than or equal to the lower jaw in all species except S. isarankurai and three Atlantic species. The lower jaw of S. isarankurai is produced beyond the tip of the *upper jaw and is visible from above. 2.3.2.2.1.6 Colour S. nebulosa3 S. flamma and S. gracilis are variously mottled and blotched on body and fins. S. elongata is uniformly brown. The other species are variously brown to olive dorsally and silvery below the lateral line. 2.3.2.2.2 Specific Descriptions and Distributions (See Tables 2.1 - 2.4B) 2. 3. 2. 2. 2.1 Saurida argentea Macleay, 1882 (Silvery saury) Saurida argentea Macleay, Proc. Linn. Soc. N.S. Wales, VI, 1882, p. 220. Endeavour River, North Queensland. Saurida micropectoralis Shindo and Yamada, Uo. Jap. Soc. Ichthyol. No. 11, 1972, p. 11. Prachuap Khiri Khan Province, Gulf of Thailand. * A specimen identical with S. undosquamis but with a longer lower jaw was found in Moreton Bay. It was considered an eberrant specimen of S. undosquamis. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 41. Material examined (32 specimens, size range : 123- 260mm) Syntype: AMS 1.16271-001, 2 specimens; 141, 133mm. Endeavour River, North Queensland. 15° 28’S, 145° 15'E. Other specimens (number examined in brackets) Queensland: AMS 1.15421-014 (3). Magnetic Island; 19° 10'S, 146° 50'E. Hervey Bay (1), collected by Clive Keenan. Northern Territory (of Australia): DPPFD-ME30 (1), Melville Island. North-Western Shelf of Australia (18), collected by CSIRO ('SOELAf). Gulf of Thailand : ZUMT 54349 (1), Prachuap Khiri Khan Province. Hong Kong: SOSC Cat. No. 9454 (2). Sri Lanka : SOSC Ref. No. 553 (3). Indonesia : SI, HS- F242 (1), Halmahera Island. Description Counts: 11-12 dorsal fin rays; 10-11 anal fin rays; 13-15 pectoral fin rays; 53-58 scales in lateral line; 19-23 predorsal scales; 13-16 interdorsal scales and 9-12 scales between adipose and caudal fins. Morphometry: Mean and range (in brackets) of characters calculated as percentages of University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 42. CM CD eg TC—DI LO in oo oo co ^ CM lO lO CO elgO CM CM lO CM eg co LO oo CD co HCO rH LO 00 e g CM CM O CO LO CM / CO CO 00 00 a CO A§? + O* CO K <3 COCO CD 4̂ oco •H 8o 4̂ KC3O o rCs0 la "K « 0) 0) CO I Kco co Co CO* CO Co Co CO Co Co Eh TABLE 2.1 : Meristic Characters of Lizardfish Number of specimens University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 43. 00 (0M0 00 rH 05 LO CD o CO 00 rH CO CM ^ . CiOSl OH O CMO tH trHH ^tH C D C D C O CM t> 00 ^ rH CO CQ CM r0H0 CD rH CT—M1 05 rH £* £ SS CO Hg g O03^ co « c o w 1 ! i •< i ! r i 1 1 1 | ^ CO oi Co' 0̂ to' oj ^ ^ ^ ^ TABLE 2.2 : MERISTIC CHARACTERS OF LIZARDFISH DORSAL PECTORAL ANAL Species 10 11 13 14 11 12 13 14 15 16 10 11 12 13 14 :15 16 17 3 3 1 2 17 8 9 2 4 2 6 42 13 23 6 9 13 21 71 29 42 42 12 53 28 7 21 8 15 13 3 2 1 1 3 32 10 22 18 14 18 72 22 43 7 31 46 18 8 8 6 1 8 2 6 5 49 14 30 4 28 30 17 71 1 26 40 64 27 Number of specimens University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh TABLE 2.4A : Mean and ranRe of morphoroetric characters as percentage of standard length University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 46. '3' m %o 3- CN CO •ti r̂. CN cn *3 i • «* CO rH • CN - | O' rH a *3- rH CC 1 rs CO rs m rs TABLE 2.4B : Mean and range of morphometric characters as percentage of head length S. s. S. S. S. Species auetralie filamentoea tumbil argentea elongata Number of specimens 28 71 17 72 32 71 Eye diameter 17.0-24.0 15.5-23.2 14.6-22.1 20.3-21.0 18.0-24.8 14.2-23.5 16.5-20.0 16.7- 16.8 18.0-22.1 15.7-23,3 17.1-24.1 15.6-25.6 20.0 19.5 19.9 20.6 21.8 18.6 18.5 16.7 20.1 19.5 19.7 19.7 Preorbital L. 16.9-22.9 16.6-23.4 20.9-22.8 20.0-23.7 19.4-23.6 21.0-25.7 19.5- 26.3 21,9-25.5 9.9-14.7 21.1 21 .0 21 .8 22.2 21.6 22.3 23.3 23,8 12.5 Postorbital L. 58.7-62.5 55.9-62.0 54.7-66.4 59.5-59.7 51.9-58.9 51.4-63.8 55.6- 64.0 57.7- 62.1 56.5-57.2 52,5- 59.5 58.9-76.1 60.4 59.4 60.3 59.6 56.1 60.8 59 .0 59.9 58.3 56 .3 67.6 Snout L. 21.5-23.1 20.0-25.9 17.1-25.5 19.8-24.5 18.0-24.6 19.0- 25.2 21.9-27.7 19.9-25.5 21.4-26.1 22.3 21.7 20.0 22.5 22.0 22.1 23.1 22.3 23.6 --------------------------- Premaxillary L. 69.0-71.9 66.1-73.9 66.1-75.9 65.1- 68.8 67.2- 73.3 63.0- 74.3 65.5- 72.9 67.8- 73.4 73.2-78.2 67,0- 79,1 67,0-76.5 61,5- 66,3 70.5 70.2 69.7 66 .8 70.1 66.9 69.2 70 .1 75,2 71.1 73 .1 63.0 Pelvic fin L. 69.6-87.0 67.6-79.9 70.6-87.8 65.3-87.5 70.3-87.7 70.8- 87.3 84.5-90.8 77,1-93.7 89.0-98.7 75.9 75.9 77.9 79.2 79.9 78 .0 87.1 84,2 93.8 78.1-103.4 Pectoral fin L. 57.7- 63.2 58.2-71.8 58.9-74.5 ^6.3-57.5 45.5- 60.1 54.9- 61.2 58.9 66.9 67 .6 88.3 51 .0 52.5 58.7 length University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 47. a) Standard length Head 24.8 (22.2 - 25.5); pectoral fin 12.2 (10.6 - 14.6); pelvic fin 18.9 (17.5 - 20.2); dorsal fin 20.5 (20.9 - 23.4); interdorsal 26.2 (25.0 - 28.8); interventral 35.6 (33.2 - 37.8); predorsal 42.6 (41.2 - 43.6); prepelvic 37.7 (34.6 - 39.2); prepectoral 24.6 (22.9 - 26.0); preanal 75.0 (72.0 - 77.4); dorsal fin base 13.1 (11.6 - 14.2); anal fin base 9.6 (8.2 - 10.5). b) Head length Eye diameter 18.5 (16.6 - 20.0); preorbital 22.3 (21.0 - 25.7); postorbital 59.0 (55.6 - 64.0); snout 22.1 (20.0 - 24.2); premaxillary 69.2 (65.5 - 72.9); pelvic fin 78.0 (70.8 - 87.3; pectoral fin 51.0 (46.3 - 57.5). Body moderately depressed, head flat and tapering to an obtusely pointed snout. Snout bears a mesial ossified protuberance which reduces posteriorly to a shallow fronto- occipital groove. Head profile slightly convex at nostrils Adipose eyelid moderate. Posterior margin of nasal flap flabellate. Lower jaw slightly inferior. Outer palatine band mostly in two rows of teeth posteriorly and usually in three or four rows anteriorly; outer band of a broad ’V' shape. Inner band in the form of a broad patch consisting of about six to seven rows of teeth at widest section. Vomerine teeth present. Teeth sagittate and slender (Plate 32A, B *, Fig. 2.14G) * Syntype (AMS 1.16271, fwet’ specimen) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 32A, Teeth configuration in palatine (p) bands and on vomer (v) of S. argentea. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 32B ('wet specimen'), Teeth configuration in palatine (p) bands and on vomer (v) of S . argentea University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 48. Caudal peduncle moderately depressed and carinate. Pectoral fin does not or just reaches base of pelvic fin. Colour: Bluish-brown, whitish beneath, many scales with silvery centres. Inner and distal outer surface of pectoral dusky. Upper inner half of pelvic slightly dusky, outer surface clear. Tips of caudal rays dusky black; especially lower lobe. Last ventral ray of subcaudal sometimes with 3-4 dark or brown spots. Upper half of rounded adipose fin dark. Tips of dorsal rays black, leading edge of dorsal fin with very faint brown spots. About 7-8 light blotches along the lateral line. Distribution: S. argentea has been reported mainly from the Central Indo-Pacific region (Fig. 2.15). It occurs on coral, muddy and sandybottoms, and in estuarine or shallow water to a recorded depth of about 116m. Size: Maximum reported is 380mm; 200-300mm being the common sizes (Fisher and Whitehead, 1974). Remark: S. argentea was found to be conspecific with S. mioropeotoralis Shindo and Yamada 1972. 2 .3.2.2.2.2 Saurida australis Castelnau 1878-79 Saurida australis Castelnau, Proc. Linn. Soc. N.S.W. Wales, 1878-79, p. 393. Port Jackson, N.S. Wales University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh * ■>CD T D cr CD 00 C t—i •r—« E : • o X - - cn CD ~ CD * O ') ooi cCO C/3 cCdZ “O CN] -od oo E; o00 CJ oCO o o ■N CDCDD =O3 .C/3 CD > a C l _ a .— i 4 - =3 CD CD O 4-» f— « o 4—' a o c= CO CO i _ o •rH a (D CD : C J 4->§ » 4-» s CD CO **— t >— i 0) 1— ■ Hr-»i O ,— i £ g O a a o c CO *— * o § <3 >rHCtj « 4-» 4-> £ ;§ o C •F r-i £ to CD *o 0) CO "O T D _Q CO T D ■»■ 4 CD L_ T D X I 4-> •rH o CO »— « 4-» «f 1 o a oo X I Lr—A| CNJ cn LL_ University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 49. Saurida tumbil McCulloch, 1929-30, p. 77, in part (Australia). Material examined (28 specimens, size range : 207 - 300mm) Neotype: QM I. 20691, 1,263mm; collected from between Lady Elliot and Fairfax Island, East of Bundaberg, Queensland, August 1981. Other specimens (number examined in brackets). QM I 20690 (14); between Lady Elliot and Fairfax Island, Hervey Bay (11), QM 1.12054 (1), Q, 1.3510 (1); Mud Island, Moreton Bay. Description Counts: Range and neotype (in brackets). 10-11 (11) dorsal fin rays; 10 - 11 (11) anal fin rays; 14 - 15 (14) pectoral fin rays: 55 - 57 (55) scales in lateral line; 16 - 21 (17) predorsal scales; 17 - 18 (17) interdorsal scales; 11 - 12 (12) scales between adipose and caudal fins. Morphometry: Mean and range (neotype in brackets) of characters calculated as percentages of a) Standard length Head 23.5, 23 - 23.9 (23.2); pectoral fin 13.8, 13.4 - 14.3 (14.1); pelvic fin 18.2, 17.5 - 18.7 (17.3); dorsal fin 19.2, 18.4 - 19.6 (18.6); interdorsal 27.5, 25.8 - 29.9 (28.4); interventral 38.8, 35.9 - 42.5 (36.9); predorsal 41.1, 40.6 - 41.5 (40.9); prepelvic 35.7, 34.3 - 36.9 (36.2); pre­ pectoral 24.4, 23.9 - 24.8 (23.9); University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 50. preanal 74.2, 72.7 - 75.4 (74.1); dorsal fin base 11.9, 11.4 - 12.4 (12.1); anal fin base 9.9, 8.9 - 12.2 (11.4). b) Head length Eye diameter 20.0,17.0 - 24.0 (21.5); preorbital 21.1, 16.9 - 22.9 (21.4); post­ orbital 60.4, 58.7 - 62.5 (69.2); snout 22.3, 21.5 - 23.1 (20.9); premaxillary 70.5, 69.0 - 71.9 (69.2); pelvic fin 77.4, 75.8 - 81.4 (74.4; pectoral fin 58.9, 57.7 - 63.2 (60.6). Body subfusiform, elongated and depressed. Fronto- occipital groove broad and concave. Slight mesial pro­ tuberance at snout tip. Predorsal profile slightly convex. Eyes large. Adipose eyelid well developed and broad. Nasal flap broad and slightly produced at inner half of posterior margin. Upper jaw slightly superior; more or less equal with lower jaw. Outer palatine band mostly in 2 rows; in the shape of a broad 'V'. Inner band in 6 - 8 rows in the broadest section. 2 - 3 teeth on wings of vomer (Fig. 2.14B). Pectoral fin reaches level of pelvic fin base. Scales in lateral line slightly keeled, especially on the caudal peduncle. Colour (Plate 33 A,B): Generally olive green above lateral line with faint traces of at least two longitudinal silvery-blue lines more or less formed by the silvery centres of the scales. The silvery centres also given an impression of rather diffused or dispersed University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh A B Plate 33: Saurida australis University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 51. dorsal origin, 3 in length of dorsal fin and about 4 between dorsal fin and adipose fin. Sides of head more or less gilt. Four longitudinal narrow and silvery-blue bands alternating with three golden longitudinal bands below lateral line and extending from pectoral fin base to the crease of caudal fin - first silvery-blue band below lateral line the most conspic­ uous and broadest. About 8 dark blotches along the lateral line. Leading edge of caudal and dorsal fins with 5 - 1 0 dark dots. Lobes of caudal fin more or less equal; tips black, especially so in the lower lobe. Caudal fin with a greenish tinge. Anal fin silvery-white. Proximal inner half of pelvic fin sparingly dusky, distal inner half and outer surface white or yellowish. Upper half of pectoral fin surface dusky, bottom half more or less silvery-white. Adipose fin with a terminal dark brown blotch. Distribution (Fibs 2.15 & 2.16): S. australis has been recorded from Hervey and Moreton Bays, and Port Jackson. These disparate localities suggest the occurrence of the species on the eastern seaboard of Australia, delimited by the 24°S and 34°S latitudes. Size: S. australis grows to over 350mm. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 2.16 : Distribution of saurida australis Castelnau,, 1878-79. Solid symbols represent localities of specimens examined; the hatched area is an additional literature record. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 52. 2.3.2.2.2.3 Saurida elongata (Temminck and Schlegel), 1846. Aulopus elongatus Temminck and Schlegel, 1846, in Siebold, Faun. Japon. (Pise.), pp. 233-234 pi. cv. fig. 2 . Saurida altipinnis (part) Gunther , 1864 , Cat. Fish . v. p. 397. Saurida truoulenta Macleay, Proc. Linn. Soc. N.S. Wales, vi 1881, p. 219. Port Jackson, N.S. Wales. Saurida eso Jordan and Herre, 1907, Proc. U.S. Nat. Mus. xxxii, p. 520; Jordan, Tanaka and Snyder, 1913, J. Coll. Sci. Tokyo, xxxiii (1), p. 53; Jordan and Hubbs, 1925, Mem. Carnegie Mus. x, p. 155. Saurida microlepis Wu and Wang, 1931, Contri, Biol. Lab. Sci. Soc. China, Zool. Ser. viii, p. 1. Material examined (3 specimens, size range : 226 - 412mm). (Number examined from each locality in brackets). AMS 1.16270 - 001 (1), Port Jackson, NSW, 33° 50', 151° 15'E; CSIRO 118001 - 38 (1), Dampier, Western Australia; Hervey Bay (1), Queensland. Description Counts: 13 - 15 dorsal fin rays; 11 - 12 anal fin rays; 11 pectoral fin rays; 58 - 60 scales in lateral line; 21 - 25 predorsal scales; 17 - 18 interdorsal scales; 10 - 12 scales between adipose and caudal fins. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 53. Morphometry: Mean length and range (in brackets) of characters calculated as percentages of a) Standard length Head 21.4 (20.7 - 21.7); pectoral fin 10.9 (10.7 - 11.0); pelvic fin 17.7 (15.7 - 19.6); dorsal fin 20.1 (20.0 - 20.2); interdorsal 28.5 (27.9 - 30.0); interventral 36.0 (34.3 - 39.4); predorsal 41.6 (38.6 - 43.5 ) ; pre- pelvic 36.1 (32.2 - 38.4); prepectoral 23.2 (22.2 - 23.9); preanal 73.7 (72.1 - 75.5); dorsal fin base 12.4 (12.3 - 12.6); anal fin base 9.7 (9.8 - 10.0). b) Head length Eye diameter 16.7 (16.7 - 16.8); preorbital 23.3 (19.5 - 26.3); postorbital 59.9 (57.7 - 62.1); snout 22.1 (19.0 - 25.2); premaxillary 70.1 (67.8 - 73.4); pelvic fin 80.4 (72.4 - 87.3); pectoral fin 49.0 (48.9 - 49.0). Body elongate, more or less cylindrical, and slightly depressed. Snout obtusely pointed. Eyes large, adipose eyelids broad. Lower jaw subequal. Caudal preduncle depressed and carinate. Pectoral fin short, not reaching base of pelvic fin. Pectoral axillary long and broad. Outer palatine band consists of three irregular rows of teeth. The inner band has about 8 rows of teeth at its widest section. Vomerine teeth present; in a central patch of about 8 - 1 5 teeth (Fig. 2.14H). Colour: Back and sides of uniform pale brown colour; no blotches or cross-bands. Very faint dusky spots on leading edge of dorsal and caudal fins. Inner surface of pectoral fin uniformly dusky, first dorsal ray tip black. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 54. Distribution (Fig. 2.15): S. elongata occurs in the Sea Japan, South China and Phillipine (Western) Seas. Records of S. elongata from Dampier (Western Australia), Hervey Bay (Queensland) and Port Jackson (New South Wales) suggest a widespread distribution in the Central Indo- Pacific region. Size: Maximum recorded is about 500mm, 250 - 380mm being the common sizes (Fisher and Whitehead, 1974). 2.3.2.2.2.4 Saurida filamentosa Ogilby, 1910 (Thread-fin saury) Saurida filamentosa Ogilby, Proc. Roy. Soc. Queensland xxiii, p. 88. Off Cape Moreton, Queensland. Saurida wanieso Shindo and Yamada, Uo Jap. Soc. Ichthyol. No. 11, 1972, p.8 . East China Sea. Material examined (71 specimens, size range : 163 — 575mm). Isotypes: AMS 1.12571, 176mm; QM 1.481, 163mm; QM 1.8917, 185mm, off Cape Moreton, Queensland Other specimens (number examined in brackets): AMS 1.20319 - 010 (2), 1.20444 - 005 (1), 1.15525 - 002 (1). Off Cape Moreton (50), collected by Bob Sanderson. North-western Australia; Indian Ocean (12), collected by CSIRO ('SOELA1), 1981. East China Sea (3), collected by Shindo and Yamada. *ZUMT 52502 (1), East China Sea. * Examination and measurements done by Y. Tominaga of University of Tokyo Museum on request. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 55. Description Counts: 11 - 12 dorsal fin rays; 10 - 12 anal fin rays; 13 - 16 pectoral fin rays; 51 - 55 scales in lateral line; 15 - 21 predorsal scales; 13 - 20 interdorsal scales; 10 - 12 scales between adipose and caudal fins. Morphometry: Mean length and range (in brackets) of characters calculated as percentages of a) Standard length Head 23.6 (21.3 - 25.5); pectoral fin 15.9 (12.8 - 18.2); pelvic fin 18.2 (16.2 - 19.0); *dorsal fin 34.9 (24.4 - 51.2); interdorsal 26.9 (23.3 - 28.6); interventral 37.2 (32.2 - 42.8); predorsal 40.3 (36.4 - 42.5); prepelvic 35.9 (24.4 - 41.8); prepectoral 24.0 (21.2 - 26.0); preanal 75.0 (68.2 - 80.6); dorsal fin base 13.4 (11,0 - 16.6); anal fin base 10.3 (8.0 - 13.8). b) Head length Eye diameter 19.8 (14.6 - 22.1); preorbital 19.8 (16.6 - 21.7); postorbital 60,2 (54.7 - 66.4); snout 20.0 (17.1 - 25.5); premaxillary 69.0 (66.1 - 75.9); pelvic fin 75.9 (67.6 - 79.9); pectoral fin 67.6 (58.9 - 74.5). Body subfusiform, caudal peduncle without lateral ridge. Predorsal moderately depressed with a gentle convex profile. Fronto-occipital groove narrow and shallow, with a slightly domed mesial ossification at snout tip. * Filamentous second and third rays included. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 56. Eye subdorso-lateral. Adipose eyelid broad. Nasal flap broadest at inner half of posterior margin. Upper jaw a little longer than the lower jaw. Outer palatine band consists of two rows of teeth and bottle-neck shaped. Inner band has about 7 - 8 irregular rows at widest section. Vomer usually with 2 - 3 teeth set on each wing. Teeth strongly sagittate (Plate 34, Fig. 2.14C). Pectoral fin reaches to beyond base of pelvic. Second and sometimes third dorsal ray filamentous; the second longest and sometimes reaching to or beyond the base of the adipose fin when laid back. Colour: Dark brown above with a greenish tinge. 7 - 8 dark blotches along lateral line. Pale yellow ventro-laterally. Upper half of the pectoral fin violet or sometimes dusky black on both surfaces, especially the bottom half. Pelvic and anal fins clear, upper margin of adipose fin dusky black. Caudal fin dusky with black tips. Leading edge of Caudal sometimes with dark spots. Distribution (Fig. 2.17): Central Indo-Pacific. Coasts of India. Deep water species, occurring down to 200m on fine sand or mud. Size: Grows to over 575mm but commonly obtained in the 200 - 350 size range. Remarks: S. filamentosa was found to be conspecific with S. wanieso Shindo and Yamada 1972. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 34 Teeth configuration in palatine (p) bands and On VOUier (V) Of S. filamentosa University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh cCOCD • •EOf EH CO- * QCD ■a OJ CO . ou crn\ 4O— i C—D i— i *\ CCDO i C_D O a •+r-H% » 4 = o— 3* i—I TCD £a D a r *.——<* LCD- cn >- ao *4—- rH ’ - CD TD r— o4 t—l do +d-> r—a« CD *\ o CD oc:O C3 a c= ■ .c O CCDO +-» cOo £oL_ G_ (—cz1 iQ— •r—4 £ s: O CD . TD TD CVO •r<* i_ a rO A§ 3 r—C—O« Ci_D Vg £ _ oo a Ct~j»} 5§ >. aCO•S o CvOjv-̂ CO CD§« ■a oCO CO O ■CO SC aD CD +o Z x= a-» TCDD CO O Q XCD i—i CNJ CD 12 0 ___________160 *__________ 160* 120 ' University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 57. 2. 3. 2. 2. 2. 5 Saurida flamma Waples, 1982 (The orangemouth lizardfish) Saurida flamma Waples. Pacific Science, 35 (3), 1982, pp. 225 - 226. Material examined (8 specimens, size range : 74 - 280mm). Holotype: BPBM 25942 (1,280mm), Kewalo Basin, Oahu, Hawaiian Islands. 21° 17’ 27"N, 157° 51’ 58MW. Sand next to coral ledges. 10m. Other specimens (number examined in brackets). Hawaiian Islands, Oahu: BPBM 25943 (3), BPBM 25944 (1), BPBM 25945 (1), BPBM 29546 (1). French Frigate Shoals: BPBM 25947 (1). Description Counts: Range and holotype (in brackets). 11 dorsal fin rays; 9 - 11 (10) anal fin rays; 13 - 14 (13) pectoral fin rays; 51 - 54 (53) scales in lateral line; 16 - 21 (16) predorsal scales; 15 - 17 (16) interdorsal scales; 10 - 11 scales between adipose and caudal fins; 3 and 4 scale rows above and below the lateral scales respectively. Morphometry: Mean, range (holotype in brackets) of characters calculated as percentages of a) Standard length Head, 25.0, 24.2 - 27.4 (24.9); pectoral fin 14.7, 13.6 - 14.9 (13.7); pelvic fin 23.4, 22.0 - 24.5 (23.5); dorsal fin 23.6, 22.1 - 25.1; interdorsal 26.8, 24.8 - 27.8 (27.8); interventral 39.2, 36.1 - 41.6 (39.9); pre­ dorsal 41.4, 39.8 - 43.4 (41.3); prepelvic 35.1, 33.4 - 36.7 (35.2); prepectoral 25.2, 24.2 - 26.8 (25.7); preanal 75.9, 75.0 - University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 58. 76.6 (76.8); dorsal fin base 15.0, 11.7 - 15.6 (14.5); anal fin base 10.5, 10.0 - 11.6 (10.2). b) Head length Eye diameter 19.7, 17.1 - 24.1 (18.1); preorbital 23.8, 21.9 - 25.5 (25.5); postorbital 56.3, 52.5 - 59.5 (58.1); snout 23.6, 21.4 - 26.1 (24.0); premaxillary 73.1, 67.0 - 76.5 (76.5); pelvic fin 93.8, 89.0 - 98.7 (94.3); pectoral fin 58.7, 54.9 - 61.2 (54.9). Body subfusiform, caudal peduncle slightly carinate. Predorsal profile depressed to rounded. Snout more or less pointed, with a medio-lateral ossified slight protuberance by each pair of nostrils. The inner half of the posterior margin of nasal flap is produced as a spatulate process, the outer half more or less arched Eye large and sub-lateral. Posterior adipose eyelid well developed, anterior narrow and usually absent from upper margin of orbit. Superior border of eye slightly higher than occipital region. Lower jaw slightly shorter. Inner palatine band consists of four irregular rows, with 2 or 3 such rows in the outer band. The first three or four teeth of the innermost row long and pointed. Vomerine teeth are set on the flanks of vomer and deceptively alligned with the outer band (Fig. 2.141, also see Plate 31). Pectoral sub-falcate and reaches beyond level of pelvic fin base. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 59. Colour: Mottled brown above, pale or pale rose below. all fins with spots and bars of dark pigment and a series of 8 - 9 bright orange bands on gums on each side of mouth. Opercle, lower jaw, and base of pectoral, pelvic, and caudal fins also tinged orange or red, particularly in larger specimens (Waples, 1982). Distribution (Fig. 2.18): Hawaii and neighbouring islands. French Frigate Shoals. Reef species occurring over sandy or partly sandy bottoms. 10m Size: Grows to over 280mm. 2. 3. 2. 2.2. 6 Saurida gracilis Quoy and Gaimard, 1824. Saurus gracilis Quoy and Gaimard 1824: P. 224, Hawaii and Mauritius. Saurus minutus Lesueur, 1825b: 118, pi. v, Mauritius Saurida nebulosa Valenciennes in Cuvier and Valenciennes, 1849: p. 504, in part, Mauritius. Saurida gracilis Jenkins, 1904 (1902), p. 433, Hawaii. *Saurida maculata nomen illegitimum Material examined (8 specimens, size range : 175 - 251mm). (Number of specimens examined in brackets). AMS 1.17232 - 008 (1), Lord Howe Island. Queensland: *QM 1.8903 (1); QM 1.12089 (1), North of Swain Reef; QM 1.427 (1), Moreton Bay; QM 1.12112, Myora Bight, Moreton Bay; * Queensland Museum Records. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh s_ CO O \ TD o C a a CD “O a CD COo CO* c= T<> •i—• 3 rQ 4—EE a 0) O R X >> CD . E * CO >vCO a '—' *—' d M— o CD ■=T CD O a ■=T CM E >> CO OO ■—i CO CO rH i—1 • CJ ■ a CD i— CD ■\ *> CL o S Z CO TD CO o A~> CD i_ CD c O * 4 - i_ E lc : e O O CD v_, CD L_ •>—« a CO L_ 4— O CD Csl CD H3 c o o *rH 4-» “O CD ■ a c n -M O CD •—i c ; i—i t H i_ > o a »““H CD O > ■> a 4-» E£ >* CO CJ ■< CD <3 o CD o ■—i i— CO =3 •—« r—t O a CL >—i CO O 4-» a O rQ c a S :0) CO CD o £ r-~̂ | CO •rH a CD 4-» CO o i_ -<—i a or~-i Q . *o CJi CD "Oi_ o -Q - . . 0) ---s Co Co Co CO CD CSJ L_ o o 4— o a . CD O J=> CO «—1 Ez CO c >> o a ‘Io—* CO CD CO —i 4-» TD a CO ZD -Q .—* CD •»—« TDO CD <3 CL CO sz Ss O -M o CO CD 4-> 3 : *»—« XT a . >* Q I— _cz CO -Q o1—oI CNJ CD •lrlH. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 60. French Frigate Shoals: BPBM 27771 (2), Counts: 10 - 11 dorsal fin rays, 9 - 1 0 anal fin rays; 13 - 14 pectoral fin rays; 15 - 16 predorsal scales; 13 - 16 interdorsal scales; 9 - 1 0 scales between adipose and caudal fins; 3 scale rows above lateral line and 4 or 5 scale rows below the lateral line. Morphometry: Mean and range (in brackets) of characters calculated as a percentage of a) Standard length Head 25.1 (24.7 - 25.4); pectoral fin 14.0 (13.6 - 14.5); pelvic fin 21.8 (20.9 - 22.6); dorsal fin 22.4 (21.7 - 22.9); interdorsal 25.8 (25.4 - 26.2); interventral 40.1 (38.4 - 41.8); predorsal 42.0 (41.2 - 42.7); prepelvic 34.7 (33.7 - 35.7); prepectoral 25.9 (25.1 - 26.6); preanal 76.7 (75.7 - 77.8); dorsal fin base 14.8 (14.4 - 15.1); anal fin base 9.8 (8.8 - 10.6). b) Head length Eye diameter 20.1 (18.0 - 22.1); preorbital 21.4 (17.1 - 23.7); postorbital 58.3 (56.5 - 57.2); snout 23.1 (21.9 - 27.7); premaxillary 75.2 (73.2 - 78.2); pelvic fin 87.1 (84.5 - 90.8); pectoral fin 56.1 (54.2 - 58.3). Body elongate, slightly depressed to rounded. Snout pointed, a medio-lateral ossified slight protuberance by each pair of nostrils. The inner half of the posterior margin of nasal flap is produced as a spatulate process, the outer half more or less arched. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 61. Eye subdorso-lateral. Adipose eyelid very narrow, interorbital space slightly concave. Superior border of eye at level of dorsal surface. Jaws more or less equal. Inner and outer palatine teeth in 2 - 3 irregular rows; outer palatine bands converge anteriorly and are bridged by a patch of vomerine teeth. The first three or four teeth of innermost row of outer band long and pointed (Fig. 2.14J). Pectoral fin more or less falcate, reaches to level of the pelvic base. A terminal dark brown spot on the posterior of adipose fin. Caudal peduncle slightly carinate. Colour: Pale brown irregularly variegated with dark cross-bands. Head similarly mottled but darker dorso-laterally; variegation extending to 4 or 5 scales below lateral line. About ten blotches in lateral line, fins with dark cross-bands. Lips of jaw with dark brown blotches interspersed with pale yellow. Distribution (Fig. 2.18): Known from the coast of East Africa, the Red Sea and throughout the Central Indo-Pacific region. Hawaii. Size: Medium size species. 2. 3. 2. 2. 2. 7 Saurida isarankurai Shindo and Yamada, 1972 Saurida isarankurai Shindo and Yamada, Uo, Jap. Soc. Ichthyol. No. 11, 1972, p. 7. Prachuap Khiri Khan Province, Gulf of Thailand. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 62. Material examined (3 specimens, size range : 103 - 111mm) . Holotype ZUMT 52501 (1,111mm). Paratypes ZUMT 54347 (1,106mm), ZUMT 54348 (1,103mm). Prachuap Khiri Khan Province, Gulf of Thailand. Description Counts (*Holotype and paratypes (in brackets)): 11 (11) dorsal fin rays, 11 (11, 12) anal fin rays; 13 ( 12, 13) pectoral fin rays; 49 48**, 49**) scales in lateral line; 20 (19**) predorsal scales. Morphometry (Holotype and paratypes (in brackets)): values of characters calculated as percentages of a) Standard length Head 24.5 (21.5, 22,1); pectoral fin 21.2; pelvic fin 17.3 (18.5, 18.9); dorsal 21.6; interdorsal (23.3, 26.5); interventral (34.0, 35.6); predorsal 43.0 (43.0, 43.2); prepelvic 37.0; prepectoral 24.7 (23.2, 24.6); preanal 71.1 (73.6, 74.3); dorsal fin base 14.4 (11.8, 13.7); anal fin base 11.7 (10.8, 11.8). b) Head length Eye diameter 21.0 (20.3, 20.8); preorbital (20.9, 22.8); postorbital (59.5, 59.7); snout 19.1; premaxillary 65.1 (68.0, 68.8); pelvic fin 70.6 (83.6, 87.8); pectoral fin 86.4 * Measured on request by Y. Tominaga of University of Tokyo Musuem. ** Scales lost but scale pockets counted. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 63. Body cylindrical and elongated. Snout slightly rounded Lower jaw longer than upper, distinctly visible from above when mouth is closed. Palatine teeth in two bands; outer band in two rows anteriorly and posteriorly, and in a single row about midway; inner band in form of a broad patch of three or four rows of teeth. Vomer toothless (Fig. 2.14E). Pectoral fin long and extending beyond base of pelvic fin; reaching to about the origin of dorsal fin. Scales deciduous. Colour: 9 - 1 0 distinct brownish blotches along lateral line. Upper margin of dorsal fin dusky, upper half of inner side of pectoral fin dusky, whole surface of lower lobe of caudal fin dusky. Distribution (Fig. 2.17): Prachuap Khiri Khan Province. Gulf of Thailand. Size: Small size species; largest specimen recorded - 117mm. 2. 3. 2. 2. 2. 8 Saurida longimanus Norman, 1939 Saurida longimanus Norman, in Jordan Murray Expedition (1933-34) Scientific Reports, 7 (1-3) pp. 23-24. Material examined (17 specimens, size range : 72 - 185mm). (Number of specimens examined in brackets). AMS 1.21617 - 004 (1), North-West Shelf, Western Australia. *PJPW 791 (1), Bali, Indonesia. SI Field No. BBC-1681 (4), off mouth of Fly River. Gulf of Papua, Papua New * Dr Peter Whitehead of BMNH1s collection. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 64. Guinea, 08° 45.0’S, 144° 05.8', 7 June 1979. SI 9450 (1), Hong Kong. SI Cat. No. 136050 (10, off Luzon, Phillippines, 13° 23’ 10"N, 123° 45* 19f,E, 4 June 1909. *TGT 3150(1), Bali, Indonesia. Description Counts: 11 - 12 dorsal fin rays; 10 - 12 anal fin rays; 12 - 15 pectoral fin rays; 48 - 51 scales in lateral line; 16 - 20 predorsal scales; 11 - 15 interdorsal scales; 9 - 1 1 scales between adipose and caudal fins. Morphometry: Mean and range (in brackets) of characters calculated as percentages of a) Standard length Head 23.5 (21.9 - 26.3); pectoral fin 20.4 (17.0 - 24.0); pelvic fin 17.0 (16.3 - 19.9); dorsal 24.2 (23.1 - 25.4); interdorsal 23.9 (21.6 - 27.3); interventral 33.4 (27.8 - 37.7); predorsal 43.5 (38.8 - 45.7); prepelvic 38.0 (32.3 - 40.8); prepectoral 25.0 (21.0 - 27.0); preanal 73.1 (64.5 - 77.2); dorsal fin base 14.2 (11.4 - 15.9); anal fin base 11.3 (9.8 - 12.4). b) Head length Eye diameter 21.8 (18.0 - 24.8); preorbital 22.2 (20.0 - 23.7); postorbital 56.1 (51.9 - 58.9); snout 22.5 (19.8 - 24.5); premaxillary 70.1 (67.2 - 73.3); pelvic fin 79.2 (65.3 - 87.5); pectoral fin 88.3 (78.1 - 103.4). * Thomas Gloerfelt-Tarp’s Collection. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 65. Body cylindrical and elongated. Snout broader than long, more or less rounded at tip. Eyes large and prominent. Adipose eyelids moderate. Bony ridges rise from occiput along interorbital edge to about the anterior edge of orbit. Outer bands of palatine teeth, more or less straight, in two bands and set close to the lips of jaw; anterior teeth large and recurved inwards. Inner band set about midway of upper jaw with about 4 rows of teeth in the widest section. Teeth more or less sagittate (Plate 35, Fig. 2.14D). Pectoral fin long, extending beyond origin of dQrsal fin when laid back. Colour: Brownish above, silvery white below; distal parts of dorsal, caudal and pectoral fins blackish. Traces of dark marks sometimes along leading edge of caudal. Distribution (Fig. 2.17): Central Indo-Pacific and Gulf of Oman. These disparate areas suggest a widespread distribution of S. longimanus in the Indo-West Pacific region. S. longimanus occurs down to about 200m. 2. 3. 2. 2. 2. 9 Saurida nebulosa Valenciennes, 1849 Saurida nebulosa Valenciennes in Cuvier and Valenciennes, 1849, p. 504, in part. Mauritius. Synodus sharpi Fowler, 1901, p. 497. Hawaii. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 35 ('wet' specimen) Teeth configuration in palatine (p) bands and on vomer (v) of S. longimanus University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 66. Saurida gracilis Gosline and Brock, I960, p. 99, in part Hawaii. Material examined (49 specimens, size range : 42 - 190mm). (Number examined in brackets). Queensland/Great Barrier Reef: AMS 1.18755 - 014 (3), between Palfrey and Lizard Islands; QM 1.1502 (1), Darnley Is., Torres Strait; QM 1.9576 (7), Heron Is., QM 1.10956 (1), Palm Is., QM 11741 (1), One Tree Is., QM I. 13787 (1), Noosa River; QM 1.15383 (2), Clack Reef; QM 1.15630 (4), Nymph Is., 14° 36 1S , 145° 14'E ; QM 1.15690 (11), Nymph Is., 14° 33'S , 145° 0 6 ’E; QM 1.16016 (1), Tijou Reef, 13° 0 5 ’S, 143° 57'E; QM 1.16201 (1), Decapolis Reef, Cape Flattery, 14° 5 0 'S, 145° 17'E . Papua New Guinea: KFR F01299 (13), Fairfax Harbour; USNM 220015 (1), Ninigo Is.. Hawaii Is: BPBM 27770 (2), Kaneohe Bay, Oahu, 21° 26' 20"N, 157° 45’ 45"W. Description Counts: 9 - 1 2 dorsal fin rays; 9 - 1 1 anal fin rays; 12 - 14 pectoral fin rays; 47 - 50 scales in lateral line; 13 - 17 predorsal scales; 13 - 16 interdorsal scales; 10 - 11 scales between adipose and caudal fins; 3 and 4 scales above and below the lateral line respectively. Morphometry: Mean and range (in brackets) of characters calculated as percentages of University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 67. a) Standard length Head 24.9 (22.0 - 27.9); pectoral fin 13.0 (12.3 - 15.5); pelvic fin 20.7 (19.2 - 23.1); dorsal fin 21.6 (20.2 - 24.1); interdorsal 25.3 (23.7 - 27.7); interventral 39.9 (37.6 - 42.6); predorsal 42.9 (39.3 - 45.6); prepelvic 34.8 (31.4 - 37.7); prepectoral 25.3 (21.0 - 28.6); preanal 76.1 (72.2 - 83.6); dorsal fin base 14.4 (12.7 - 16.5); anal fin base 9.9 (9.0 - 11.7). b) Head length Eye diameter 19.5 (15.7 - 23.3); preorbital 21.9 (18.7 - 24.7); postorbital 58.5 (50.4 - 63.9); snout 22.3 (19.9 - 25.5); premaxillary 71.1 (67.0 - 79.1); pelvic fin 84.2 (77.1 - 93.7); pectoral fin 52.5 (45.5 - 60.1). Body elongate, slightly depressed and more or less slender. Snout pointed, a medio-lateral ossified slight protuberance by each pair of nostrils. The anterior margin of nasal flap is semi-circular, inner half of the posterior margin villi-form to spatulate; the outer half more or less arched. Adipose eyelid very narrow. Superior border of eye at level of dorsal surface, interorbital space sloping gently to nasal area. Lower jaw slightly shorter than upper. Outer band of palatine teeth of 2 or 3 poorly defined rows of teeth; the first three or four teeth of innermost row long and pointed. Inner band consists of about 3 poorly defined rows. Vomerine teeth set on wings of vomer and deceptively aligned with outer band (Plate 31, Fig. 2.141). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 68. Pectoral fin reaches to level of pelvic base. Colour: Dorsal aspect greenish brown, irregularly variegated with dark cross-bands; pale ventro-laterally. Dark blotches also on ventral sides of lower jaw and sometimes extend to near base of pelvic fin. Fins, except anal fin, also with dark cross­ bands. Anal fin usually clear. Distribution (Fig. 2.18): Widely distributed in the Indo-West Pacific region. Hawaii Islands. Size: A small species, common sizes being 80 - 130mm. 2.3.2.2.2.10 Saurida tumbil (Bloch), 1795 (The common or yellow-banded grinner) Salmo tumbil Bloch, 1795, Naturg. Ausl. Fische., ix, p. 112, pi. ccccxxx.; Schneider, 1801, in Bloch, Syst. Ichth, p. 405. Osmerus tumbil Lacepede, 1803, Hist. Nat. Poiss. v. pp. 230, 236. Saurus tumbil Cuvier, 1817, R. Anim. ii p. 169. Saurus badi Cuvier, 1829, R. Anim. ed. 2, ii, p. 314; Cantor, 1850, J. Asiat. Soc. Bengal, xviii (1849), p. 1252. Saurus milii Bory de St. V., 1829, Diet. Class, H.N. xv, p. 189, pi. civ. fig. 2. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 69. Saurus badimottdh Riippel, 1837, N. Wirbelth, Abyssin. (2), p. 77. Laurida tumbel Swainson, 1839, N. H. Fishes etc. ii, p. 288, Saurus argyrophanes Richardson, 1846, 15 Rept. Brit. Assoc. Adv. Sci. (Cambr. 1845), p. 302. Saurida torribil Cuvier and Valenciennes, 1849, Hist. Nat. Poiss, xxii, p. 500; Bleeker, 1851, Nat. Tijdschr. Ned. Ind. ii, p. 214; Bleeker, 1852, Verh. Batav. Gen. xxiv, Chirocentr. p. 20. Saurida turribil3 1859, Nat. Tijdschr. Ned. Ind. xviii, p. 356; Gunther, 1864, Cat. Fish. v. p. 399 (etc. See Norman, 1935). Material examined (72 specimens, size range 52 - 327mm). (Number examined in brackets) Queensland: QM 1.6763 (1), Cape Cleveland; QM 1.8536 (1), Chinamans Bay; QM 1.12111 (1), Moreton Bay; QM 1.16200 (1), Decapolis Reef, Cape Flattery, 14° 50'S, 145° 17’E; QM I. 16263 (1), Torres Strait, 9° 51'S , 142° 49’E; QM 1.18150 (1), 12° 35'S, 143° 25' 7"E. *'Curlew’ Collection: Burkitt Is. (4), Innisfail (4), East of Wilkie Is., between Magpie and Nobody Reefs (4), Cape Melville (4), Brampton Beach (4); Moreton Bay (39); Hervey Bay (4). * Collected by Bryan Wallis of Innisfail, skipper of the 1 Curlew1. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 70. Bali (Indonesia): *PJPW 109(2); *PJPW 721(1); *PJPW 721a(l). Papua New Guinea: KFR F0626(3), South of Ramu River. Description Counts: 11 - 13 dorsal fin rays; 9 - 1 1 anal fin rays; 14 - 15 pectoral fin rays; 49 - 54 scales in lateral line; 18 - 24 predorsal scales; 10 - 16 interdorsal scales; 8 - 1 2 scales between adipose and caudal fins. Morphometry: Mean and range (in brackets) of characters calculated as percentages of a) Standard length Head 23.4 (21.4 - 28.6); pectoral fin 11.4 (10.1 - 12.7); pelvic fin 18.7 (16.8 - 20.6); dorsal fin 21.7 (20.3 - 25.0); interdorsal 26.5 (22.5 - 28.7); interventral 37.1 (33.6 - 41.2); predorsal 42.2 (40.8 - 43.6); pre- pelvic 37.2 (34.8 - 41.5); prepectoral 24.0 (22.9 - 27.4); preanal 74.1 (72.5 - 77.8); dorsal fin base 13.7 (11.8 - 15.1); anal fin base 8.8 (7.2 - 14.2). b) Head length Eye diameter 18.6 (14.2 - 23.5); preorbital 21.6 (19.4 - 23.6); postorbital 60.8 (51.4 - 63.8); snout 22.0 (18.0 - 24.6); premaxillary 66.9 (63.0 - 74.3); pelvic fin 79.9 (70.3 - 87.7); pectoral fin 48.9 (43.7 - 56.4). Body elongate, sub-cylindrical, and anteriorly slightly depressed. Snout conical and short, preorbital profile more or less straight and steep. Interorbital concave. * Dr. Peter Whitehead of BMNH’s collection. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 71. Occipito-caudal profile convex, caudal peduncle slightly depressed and carinate; especially in large sizes. Adipose eyelid moderate. Nasal flap arched on both anterior and posterior margins, the latter being broader. Upper jaw slightly longer than lower jaw. Outer palatine of three irregular rows, 3 - 4 rows anteriorly. Each outer band of a 'stretched-out' sigmoid shape. Inner band set close to outer band and consists of about 6 rows of teeth at the widest section. 3 - 7 vomerine teeth present. Teeth strongly sagittate and stout (Plate 30 A, B, Fig. 2.14F). Pectoral fin short, not or just reaching to level of pelvic fin base. Opercular membrane sometimes covers base of pectoral fin. Colour: Dusky brown with a greenish tinge dorsally. Pale yellow ventrally. Inner surfaces of upper half of pectoral and outer half of pelvic fins dusky black. Anal clear. Distal ends of dorsal and caudal fins dusky black. 7 - 9 dark blotches along lateral line. Traces of three cross bands on dorsal aspect of body; one each at origin and posterior of dorsal fin base, the third posterior to adipose fin base. Adipose fin dusky black. Faint dark spots on leading edge of dorsal fin. Distribution (Fig. 2.19): Widely distributed in the Indo-West Pacific Region. Size: Grows to over 450mm. Common size range - 200 - 300mm. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh TD CD CZ •rH OEE X CD CO CO CZ TJ CD i _ £ O - ■r 4 o m o CD C D CD i— C l *— 1 CO CD i— ■> u - Z J o 4-» _CZ O c_) CO i — o CD CD 1— « -»—4 +-> CQ -M — '— ' o _ o a o c i r-H c •rH 4-J 4-> w C ■r < CD T U CO " O CD a § i _ a Co C l CD CD i— M— 1— a o CO CO c= .— • a o o CD •rH _ o L_ -M EE o Z3 > > _ Q CO TD •r—t CD i— " O .CZ -M •rH o CO •—« -M •i—i O O Q GO -CZ Ci—DI CM CD University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 72. 2.3.2.2.2.11 Saurida undosquamis (Richardson) , 1848 (Large-scaled saury or spotted-tail grinner) Saurus undosquamis Richardson, 1848, Zool. 'Erebus' and ’Terror', Fish. p. 138. pi. 1., figs. 1 - 6 . Saurida undosquamis Gunther. 1864, Cat. Fish, v, p. 400; Regan, 1918, Ann. Durban Mus. ii, p. 76; Gilchrist, 1922, Fish. Mar. Biol. Survey S. Afr. ii, (1921). Spec. Rep. iii, p. 55. Saurida grandisquamis Gunther, 1864, Cat. Fish, v, p. 400; Gunther, 1880, Shore Fishes 'Challenger', p. 50; Weber, 1911, Abh. Senckenb. Naturf. Gesellsch, xxxiv, p. 22; Weber and Beaufort, 1913, Fish. Indo-Austral. Arch, ii, p. 141; Fowler and Bean, 1922, Proc. U.S. Nat. Mus. lxii, Art. 2, p. 3; Fowler, 1928, Mem. B.P. Bishop Mus. x, p. 66; McCulloch, 1929, Mem. Austral. Mus. v. pp. 71, 78; Fowler, 1929, Ann. Natal Mus. vi, p. 249; Hardenberg, 1933, Treubia, xiv, p. 220. Saurida tumbil (in part) Playfair and Gunther, 1866, Fish. Zanzibar, p. 116. Saurida argyrophanes Jordan and Evermann, 1902, Proc. U.S. Nat. Mus. xxv, p. 329; Jordan and Herre, 1907, Proc. U.S. Nat. Mus. xxxii, p. 519; Jordan, Tanaka and Snyder, 1913, J. Coll. Sci. Tokyo, xxxiii (1), p. 53; Jordan and Hubbs, 1925, Mem. Carnegie Mus. x, p. 155. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Saurida tumbil Regan, 1908, Trans. Linn. Soc. London, Zool. xii, p. 219; Gilchrist and Thompson, 1909, Ann. S. Afric. Mus. vi, p. 265; Barnard, 1925, Ann. S. Afric. Mus. xxi, p. 225. Saurida macrolepis Tanaka, 1916, xxix, p. 39? Material examined (42 specimens, size range : 100 - (300mm). (Number examined in brackets). AMS 1.12569 (1), Croker Is., Northern Territory; BMNH 1847.2.30.13 (1); BMNH 1977. 4.22.1 (1), Northern Western Australia; Queensland: QM 1.12422 (1), Mouth of Proserpine R; QM 1.12983 (1), N.W. Cape Moreton; QM I. 13735 (1), Noosa River, Munna Point; QM 1.15070 (1), Torres Strait; QM I. 16170 (1), Hannibal Is., 11° 33'S, 142° 57’E; QM 1.18300 (1), 12 faths, 14° 10.7', 144 2.3'; Hervey Bay (4), collected by Clive Keenan, and Greg Campbell, 1981; Moreton Bay (22); Western Australia: QM 1.13681 (1); Bali (Indonesia): *PJPW 712 (3); **TGT 3212A (1); Papua New Guinea: KFR F0834 (2), Girra Garra Bay, Jones Reef, Portlock Harbour. Description Counts: 11 - 13 dorsal rays; 9 - 1 2 anal fin rays; 12 - 13 pectoral fin rays; 46 - 53 scales in lateral line; 16 - 20 predorsal scales; 13 - 19 interdorsal scales; 9 - 1 2 scales between adipose and caudal fins. Dr. Whitehead of BMNH's collection Thomas Gloerfelt-Tarp collection. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 74. Morphometry: Mean and range (in brackets) of characters calculated as percentages of a) Standard length Head 24.3 (23.1 - 25.5); pectoral fin 15.6 (13.5 - 17.0); pelvic fin 17.7 (16.1 - 20.9); dorsal fin 22.3 (20.1 - 24.0); interdorsal 25.4 ( 20.9 - 27.9); interventral 36.3 (32.7 - 39.4); predorsal 42.8 (40.8 - 44.6); pre- pelvic 37.5 (33.6 - 38.7); prepectoral 25.1 (23.1 - 26.9); preanal 71.9 (72.1 - 77.9); dorsal fin base 13.7 (12.1 - 15.3); anal fin base 9.8 (8.0 - 12.1). b) Head length Eye diameter 19.5 (15.5 - 23.2); preorbital 21.4 (19.8 - 24.7); postorbital 59.4 (55.9 - 62.0); snout 21.7 (20.0 - 25.9); premaxillary 70.2 (66.1 - 73.9); pelvic fin 75.9 (69.6 - 87.0); pectoral fin 66.9 (58.2 - 71.8). Body moderately depressed and predorsal more or less rounded. Snout rounded with a mesial ossified protuberance. Fronto-occipital groove wide and shallow, slightly more concave at occiput. Dorsal profile slightly convex. Adipose eyelid broad. Nasal flap broader and slightly produced at inner half of posterior margin. Jaws more or less equal. Teeth of outer band in 2 rows and occasionally in 3 rows anteriorly. Outer band slightly bottle-necked; each arm more or less straight except at anterior portion. Inner band with 5 - 7 rows of teeth. Vomer with or without teeth; if present 1 or 2 teeth set on each wing of vomer. Teeth short and sagittate (Plate 29, Fig. 2.14A). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 36: s. undosquamis (general body colour), s = rounded snout, d = black spots on leading edges of caudal and dorsal fins. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh ■•CO *N CDZ •C=O « r O EE 00 CXD c00 - /-S CD T CDO c £ OS—oCO c_>"i—O C CJD 1C—D c COO. CDsz J- .o—• >4O— -ZM3 '—^ CCDO a CiD~CO •4»—is •r̂-» r' 4r-H> Cr ■—• —-4 co oo -o' •—o• ocz 4~> ••4 »—' ■§ CaD ~l -->H CCOD "oO 0 CO Ci—Cl CDMO— i_ D a CZ .—CO1 CO •o(-H _o C oD 4-» Q _ZQD E a f-H C >O* TCJ 4i-—» •TrDH -oC DZ CO f—H 4-» oOo xaz CoNJ CvJ o> University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 76. Body slender, head large, both more or less compressed. Interorbital space concave, occipital regions very rugose. Adipose eyelids narrow. Snout blunt and very short; its length always shorter than eye diameter. Lower half of operculum prolonged backward, edge generally covering the base of pectoral fin. Pseudobranchiae well developed. Lower jaw usually extending beyond upper jaw when mouth is closed. Teeth conical, depressible and closely set; a single band of teeth on each side of palate (Plate 37). Vomerine teeth absent. Scales cycloid. Scales on cheeks and opercular. Anal fin base long, about 1£ times dorsal fin base. Pectoral fin extending much beyond pelvic fin base and usually reaching dorsal origin. Pelvic fin large, long and with 8 rays, the first ray the shortest. Caudal fin deeply forked, lobes subequal. Anus nearer to origin of pelvic than to caudal fin. Monotypic genus, widely distributed in the Atlantic, Indian and Pacific Oceans. Closely related to Synodus. 2.3.2.3.1 *Traehinocephalus myops (Bloch and Schneider) 1801 (**Painted saury or grinner) Salmo foetens (non Linnaeus), Bloch, 1794, Naturgesch. Ausl. Fische. vii. p. 118, pi. ccclxxxiv fig. 2; Schneider, 1901, in Bloch, Syst. Ichth. p. 404. * An extensive synonymy and list of references are included in Anderson, Gehringer and Berry (1966). ** Known as snakefish in the United States and Canada (See American Fisheries Society Special Publication, 1980). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 7 7 . Salmo myope Schneider (ex Forster MS), 1801, in Bloch, Syst. Ichth. p. 421. Osmerus lerrmisoatus Lacepede, 1803, Hist. Nat. Poiss. V, p. 230, pi. vi. fig. 1. Saurus trunoatus Spix, 1829, Pise. Brazil, p. 82, pi. xlv Laurida foetens Swainson, 1839, N.H. Fishes, etc. ii, p. 288. Saurus limbatus Eydoux and Souleyet, 1842, in Vaillant, Voyage ’Bonite', Zool. 1, p. 199. Saurus lerrmisoatus Richardson, 1846, 15 Rept. Brit. Assoc (Cambr. 1845), p. 301. Saurus trachinus Temminck and Schlegel, 1846, in Siebold Faun. Japon. (Pise.), p. 231. pi. cvi. fig. 2. etc. Saurus myops Cuvier and Valenciennes, 1849, Hist. Nat. Poiss. xxii, p. 485 etc. Saurus brevirostris, 1861, Poey, Mem. Hist. Nat. Cuba, ii, p. 305. Traohinooephalus my ops Gill, 1861, Proc. Acad. Nat. Sci. Philad., Suppl. (Cat. Fish. East Coast N. Amer.), p. 53; Jordan, 1890, Proc. U.S. Nat. Mus. xiii, p. 314 etc. Synodus myops Bleeker, 1875, Atl. Ichth. vi. p. 153, Saurid, pi. ii, fig. 3; Jordan and Gilbert, 1883, Bull. U.S. Nat. Mus. xvi, p. 281. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 78 . Goodella hypozona Ogilby, Proc. Linn. Soc. N.S. Wales, xxii, 1897, pp. 250 and 253. Traohinooephalus hypozona Ogilby in Whitley (1948), W. Australia Fish. Dept. Fish. Bull. No. 2, p. 13. Material examined (71 specimens, size range : 49 - 233mm). (Number of specimens examined from each locality in brackets). Queensland/Great Barrier Reef: QM 1.12114 (1), Broadhurst Reef; QM 1.16219 (1), Lizard I s . , 14 ° 3 8 ' S, 145° 2 4 ' E; QM 1 .17283 ( 1 ) , Dalrymple Islet, Torres Strait. Moreton Bay (30). QM 1.10268 (2), Exmouth Gulf, Western Australia. BPBM 19855 (6), Red Sea, N.W. Shore of Gulf of Aqaba. BPBM 20553 (1), Madras, India. Hawaiian Is.: BPBM 24049 (8), Molokai,; BPBM 23896 (16), Oahu. KFR F0139 (5), South East of Ramu River. Additional specimens given qualitative but limited quantitative examination. Queensland: Moreton Bay - QM 1.8896/901; 1.13084; 1.13094; 1.12930; 1.9537. Pt Cartwright, 26° 43 ' S , 153° 1 5 ' E - QM 1 .9 9 5 3 ; 1 . 10000. Fraser Is. - QM 1.8; 1.8904. Caloundra - QM 1.9959; 1.9995. QM 1.11370, Maroochydore. QM 1.17307, Mooloolaba. QM 1.3302, Currumbin. Torres Strait - QM 1.16259, Stephen to Bramble Bay; QM 1 .16261 , 9 ° 5 6 ’ S, 142° 2 5 'E . Description Counts: 11 - 14 dorsal fin rays; 14 - 17 anal fin rays; 11 - 13 pectoral fin rays; 51 - 58 scales in lateral line; 13 - 17 predorsal scales; University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 70. 13 - 17 interdorsal scales; 6 - 1 0 scales between adipose and caudal fins; 3, and 5/6 scales above and below the lateral line respectively. Morphometry: Mean and range (in brackets) of characters calculated as percentages of a) Standard length Head 25.4 (23.4 - 28.6); pectoral fin 13.3 (12.0 - 15.1); pelvic fin 29.0 (26.7 - 30.9); dorsal fin 27.1 (22.0 - 32.9); interdorsal 25.9 (20.6 - 29.9); interventral 30.0 (25.4 - 34.2); predorsal 40.3 (36.4 - 44.5); pre- pelvic 33.0 (32.0 - 37.2); prepectoral 25.1 (23.2 - 28.1); preanal 65.6 (58.6 - 70.4); dorsal fin base 17.4 (15.6 - 19.7); anal fin base 25.3 (20.8 - 28.5). b) Head length Eye diameter 19.7 (15.6 - 25.6); preorbital 12.5 (9.9 - 14.7); postorbital 67.6 (58.9 - 76.1); snout 12.9 (11.4 - 14.6); premaxillary 63.0 (61.5 - 66.3); pelvic fin 115.3 (105.2 - 122.4); pectoral fin 52.4 (42.8 - 59.5). Body elongate and sturdy. Head blunt, sloping abruptly downward to preorbital area. Snout shorter than eye. Upper surface and occipital regions of head strongly rugose. Interorbital space deeply concave. Adipose eyelid rudimentary. Jaws subequal, lower jaw slightly longer. Only one band of two rows of teeth on each side of palate (Plate 37). No vomerine teeth. Pectoral fin extending beyond base of pelvic fin, which is longer than head. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 37 Teeth configuration in palatine (p) bands of T. myops. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 80. Colour: An oblique black scapular patch above pectoral fin base. Four blue-grey longitudinal bands distributed on body as follows: - one from just behind origin of pectoral fin but below scapular patch and running below lateral line to the base of the lower lobe of caudal - one running from just behind eye through scapular patch and just above lateral line to mid-portion of caudal lobe, fading towards caudal lobe from about the adipose fin - one originating from about just above the scapular patch to the base of upper caudal lobe - one originating from post-ocular portion of head coalescing with its ^opposite number before and again behind adipose fin. First three longitudinal bands edged dark brown. Pairs of short bands on dorsal aspect - three pairs in predorsal length; third at origin of dorsal fin, two pairs in dorsal fin base. Two pairs and one fused patch before adipose fin, and another fused patch after. Dorsal fin with two prominent golden brown stripes on a silvery blue hue, pelvic fin with a golden brown band running vertically from first ray to the tip of last ray, either side of band violet (bluish). Inner side of pectoral and caudal fins dusky yellow, anal bluish and edged by a golden brown band. Caudal fin tips black. * from other side of body. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 81. Distribution (Fig. 2.21): Almost circumtropical. It has been reported from disparate areas in the Indian, Pacific (and Atlantic) Oceans - however no records are available from the western seaboard of the Americas. T. myops occurs in a wide range of depths, from shallow water to over 100m. Gibbs (1959) found T. myops at over 2000m in or near the Gulf Stream. Size: Grows to over 250mm. Common size range 140 - 220mm. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 0O0 / —s •n i— TD CD CD - o d ■i—i CD E C o x : X o CD o o - CO CO TD c : "O C CD i— o EE o iH o s z o a> c_) CD L_ o C l I—1 co CD CQ L_ — M— Z3 O -«-» CO O Cl, CO i_ O CD CD •rH 4-» £ 4-» f —« CO ---- - 3 ' 1 a ----« CJ o a a . o c*—' o O •rH O 4-» 4-» C CZ •r—« CD •O CO “O a CD o j* i_ E~H Q . CD CD i— l_ o o CO CO c f—1 a o •11 o CD -Q L_ +-> EE o =3 >> •-iQ—« CO "OCD "O X I 4-J t c_) CO 4-» •• ( o o Q CO x : Csi CM O) 120 160* 160 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 82. 2.4 DISCUSSION The family Synodontidae is a large one of very similar species grouped under three genera - Saurida3 Synodus and Traohinocephalus. These genera were separated on account of pelvic rays (9 in Saurida, 8 in Synodus and Traohinocephalus) and number of palatine bands (2 in Sauridaj 1 In Synodus and Traohinocephalus ) as well as the relative lengths of the dorsal and anal fin bases, as have been reported by a number of authors (e.g. Norman, 1935; Chen and Yeh, 1964; Anderson et al 1966). However Cressey (1981) found that SynodusfSaurida and Traohinocephalus could be separated easily on the basis of the presence or absence of scales on the caudal fin - in Saurida, Harpodon, and Bathysaurus each procurrent and principal rays bear a row of scales. In Traohinocephalus only the procurrent rays bear a row of scales. In Synodus there are no scales on the caudal fin rays. Traohinocephalus is monotypic and no apparent differences were found between its specimens studied from various Indo- Pacific areas between the Red Sea and the Hawaiian Is. On the contrary Saurida is polyspecific with 11 cryptic species in the Indo-Pacific region alone. Difficulties in Saurida systematics because of similar morphologies were compounded by their widespread distribution and supposedly unambiguous meristic traits or characters like number of lateral line scales and fin rays falling into overlapping ranges between many of the species. Though general protein analysis indicated considerable inter-specific similarity and also intra-specific variability, specific enzyme analysis revealed differences between some or all species in the following enzymes - CK, GAPDH, G-3-PDH, SOD, LDH, IDH, PGM, AH, SDH, EST and AAT. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 83. Distinctive phenotypes were also recognized in the most cryptic pairs of S. undosquamis and S. a u s t r a l i sand S. tumbil and S. argentea. Similar results were obtained by Waples (1982). Waples (op ait) used electrophoretic analysis to confirm the presence of three species - S. gracilis, S. nebulosa and S. flamma - where only a single species, S. gracilis, had previously been recognized. Three groups of species with similar morphologies with regards to palatine and vomerine teeth configuration, pectoral fin length, shapes of nasal flap and snout and general body colour were evident - S. undosquamis, 5. tumbil and S. nebulosa types as described previously. From the description and analyses of morphological and meristic characters S. gracilis, S. nebulosa and S. flamma closely resemble each other. The most distinctive feature of S. gracilis is the patch of vomerine teeth (central) that bridges the gap between the outer rows of palatine teeth. The striking reddish-orange coloration on the mouth and body of S. flamma and its higher scale count, 50-54, as opposed to 47 - 50 in S. nebulosa identifies 5. flamma (Waples, 1982). In the S. undosquamis group, S. longimanus is readily recognized from any other species by 1) its longer pectoral fin - the longest in all Indo-Pacific and Atlantic Saurida; 2) the narrower inner bands of palatine teeth, outer band set close to lips, and 3) its elongated body. S. isarankurai is the only known Indo-Pacific species with a lower jaw longer than upper jaw and visible from above when mouth is closed. The Atlantic species: S. suspicio Breder 1927, S. caribbaea Breder 1927, S. brasiliensis Norman 1935 all possess a longer lower jaw. However as Shindo and Yamada (1972) also reports 1 S. isarankurai have fewer scales in lateral line, 47 - 50 (mode 59), whereas S. suspicio and S. caribbaea have 51 - 60 scales. S. isarankurai has nine to University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 84. ten marked brownish blotches along the lateral line, while S. brasiliensis has about six faint blotches; and upper half of pectorals dusky in the former, and there are a few spots of pigment on lower portion of pectoral in the latter. In addition to these, S. isarankurai has a somewhat longer lower lobe than the upper of the caudal fin, while in S. bvasiliensis both lobes are nearly of equal length. ' There are slight colour variations amongst S. tumbil and S. undosquamis taken from along the Queensland Coast. The northern specimens of S. tumbil and 5. undosquamis were found to be of lighter coloration than the southerners. Yamada and Ikemoto (1979) found that S. undosquamis which distribute in the waters from the Southern East China Sea down to the warm Thai, Philippine and Hong Kong waters have clear dots on the upper edge of caudal, whereas those found in the northern waters (north of the Central East China Sea, Kii channel and Wakasa Bay) have none. Shindo and Yamada (1972) considered S. wanieso a distinct species from S. filamentosa. However, examination of the holotype and paratypes indicated that S. wanieso is conspecific with S. filamentosa. The only difference apparent between the two seems to be in coloration - the violet coloration of the upper half of the pectoral fin of S. filamentosa - which is not uncommon amongst widely distributed species. The most distinctive character of S. filamentosa is its elongated and filamentous second and sometimes third dorsal ray. In this study, S. australis Castelnau, 1878 - 79 has been removed from the synonymy of S. turribil (see McCulloch,1929-30). S. australis is rather very similar to S. undosquamis3 especially in palatine and vomerine dentition and meristic characters, as Castelnau (1878-79) reports: 'The number of the fin rays seems to unite this species (i.e. S. australis) with undosquamis3 but the pectorals University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 85 . are much shorter, and are far from extending to near the vertical of the origin of the dorsal'. The percentage values of the pectoral length relative to the standard length (mean in brackets) are 13.4 - 14.3 (13.8) and 13.5 - 17.0 (15.6) for S. australis and S. undosquamis respectively. The other point worth of mention is the distinct range of lateral line scales of the two species: 46 - 53 in S. undosquamis and 55 - 57 in S. australis. Richardson (1848) states that 'there are about fifty-seven rows of scales between the gill-opening and caudal fin....' However the examination of the lectotype (most of its scales lost) suggested that Richardson's count probably included scales on caudal fin which were ignored in this study. S. australis also possesses a rather elongated body with a peculiar coloration. In the biochemical regime, S. australis and S. undosquamis were found to exhibit distinctive *phenotypes for ADH (M), ME (M), MDH (L), G-6-PDH (L), PK (M), MPI (L,M) and XDH (M,L) which is indicative of the maintenance of their gene pool integrity though circumstantial evidence suggests they are sympatric at least in/or between Hervey and Moreton Bays. Thus they are not interbreeding naturally and should therefore be considered as two distinct bio­ logical species. The S. tumbil group comprises S. tumbily S. argentea and S. elongata. S. micropectoralis Shindo and Yamada (1972) was found to be conspecific with S. argentea and the assertion by Shindo and Yamada 1972 that the pectoral fin of S. micro- pectoralis (i.e. S. argentea) is 'the shortest in all Indo- Pacific species' is not valid. Short pectorals are common to the S. tumbil group. The percentage values (mean in brackets) relative to standard length for the 72 S. tumbil 32 S. argentea and 3 S. elongata specimens examined are 10.1 - 12.7 (11.4), 10.6 - 14.6 (12.2) and 10.7 - 11.0 (10.9) respectively. S. elongata is readily recognized by the * M - muscle, L - liver. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 86. uniformity of its colour (brown) and higher scale count of over 59. Wu and Wang (1931), Chen and Yeh (1964) and Shindo and Yamada (1972) reported of 59 - 71 lateral line scales in S. elongata. The respective ranges for S. tumbil and S. argentea are 49 - 54 and 53 - 58. Teeth of S. tumbil are stout but slender in S. argentea. The latter species is silvery blue below lateral line, whereas the former silvery white. S. ferox as described by Ramsay in 1883 appears to be a nomen dubium. With regards to the present study and the relevant literature, the taxonomic key for Saurida species is thus revised as follows: 1. a. Outer bands of palatine teeth usually in 2 and occasionally 3 rows anteriorly ................. 2 b. Outer bands of palatine teeth in three of four rows. Vomerine teeth present and central. Pectoral fin short ............................. 9 2. a. Outer palatine band consists of 2 or 3 poorly arranged rows of teeth; anteriormost 3 or 4 teeth of innermost row conical, elongated and pointed. Outer and inner bands narrowly separated. Body and fins variously mottled and blotched. Posterior margin of nasal flap more or less spatulate .............................. 3 b. Outer palatine band consists of well defined rows of teeth. Outer and inner bands well separated. Teeth in each row more or less equal, sagittate and short. Vomerine teeth present (on 'wings' of vomer) or absent. Posterior margin of nasal flap more or less flabellate ................... 5 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 87. 3. a. Outer palatine bands converging anteriorly and bridged by a patch of vomerine teeth. 49 - 52 lateral line scales. S. gracilis b. Outer bands (deceptively) aligned with vomerine teeth on ’wings' of vomer ...................... 4 4. a. 50 - 54 lateral line scales; reddish-orange coloration on the mouth and body. S. flamma b. 47 - 50 lateral line scales; colour generally buff. S. nebulosa 5. a. Pectoral fin long, extending beyond origin of dorsal fin when laid back. Outer palatine band in two rows of teeth and set closely to lips of jaw. Vomerine teeth absent. 48 - 51 lateral line scales. S. longimanus b. Pectoral reaching to at least base of pelvic fin but not extending beyond origin of dorsal fin...................................... 6 6. a. Second and sometimes third dorsal ray filamentous, the second longest and sometimes reaching to or beyond the base of the adipose fin when laid back. 51 - 55 lateral line scales. Vomerine teeth usually present. Teeth strongly sagittate. S. filamentosa b. No dorsal ray filamentous ...................... 7 7. a. Outer palatine band in at least two rows in its entire length. Vomerine teeth present or absent..8 b. Outer palatine band in two rows anteriorly and posteriorly, and in a single row about midway. Vomerine teeth absent. 47 - 50 lateral line scales. S. isarankurai University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 88. 8. a. 55 - 57 lateral line scales. Body with silvery- blue bands alternating with golden brown bands running below the lateral line to the caudal section. Vomerine teeth present. S. australis b. 46 - 53 lateral line scales. Body generally olive dorsally and silvery white below lateral line. Vomerine teeth present or absent. S. undosquamis 9. a. 58 - 71 scales in lateral line. Back and sides uniform, no blotches or cross bands on back and sides. S. elongata b. Scales in lateral line less than 59 ............ 10 10. a. 53 - 58 lateral line scales. Teeth sagittate and slender. Bluish-brown to light brown dorsally, whitish ventro-laterally. S. argentea b. 49 - 54 lateral line scales. Teeth sagittate and stout. Dusky-brown with a greenish tinge dorsally, pale yellow or white ventro-laterally. S. tumbil University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 89. 3. FOOD, FEEDING HABITS AND ADAPTATIONS 3.1 INTRODUCTION The basic functions of an organism - its growth, development, reproduction etc. - all take place at the expense of energy (food) (Nikolsky, 1963). An organism could be described as a conduit of energy in the ecosystem. The organism obtains and disburses energy amongst its life-sustaining activities - as decreed by a compromise between its genotype and the ever- changing environment - for optimal returns on its energy outlays. The developing embryo draws on a limited supply of energy in the form of food reserves, which it receives from the maternal organism (Nikolsky, 1963). After exhausting its energy reserves in the initial stage of growth, the young, and also later as an adult organism, must obtain both materials and energy from the environ­ ment by feeding. The acquisition of food by fish is a process that usually involves searching, detection, capture, and ingestion. A hungry fish performs searching activities that increase the probability that it will discover food (Keenleyside, 1979). In his review of problems and approaches in the analysis of motivation of animal behaviour Colgan (1973) proposed a model which states that "Hunger is presumed to be determined jointly by the metabolic debt (systemic need) of the fish and the amount of food in the stomach .... For a given environmental condition, hunger determines the probability that an encountered item is eaten and the interval between encounters." Fishes are adapted to a wide variety of foods (Lagler, Bardach and Miller, 1962; Nikolsky. 1963) and knowledge of these foods and feeding habits is of prime importance in fishery management and population studies. Every species University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 90. of fish is adapted to feeding on a particular combination of food (Nikolsky, 1963). In varying or uncertain environments, extreme specialization for a few types of food or of the structural attributes are few. This flexibility reflects the environment, for it is advantag­ eous for all species to be able to take advantage of tempor­ arily super-abundant resource (Keast and Webb, 1966). Very few fish are stenophagous and diet may change with growth, size, environment and sometimes with sex. The literature on food and feeding habits of fish dates back to the days of Aristotle (384 - 322BC). In his exhaustive study of the digestive system and feeding habits of about 150 species of fish Suyehiro (1941), also provided an excellent historical review of food studies on fish. The common methods of studying food and feeding habits of fish - occurrence (volume and weight measurements, number, fullness) have evolved from the analyses of stomach contents either qualitatively (usually relying on list of items and their percentages e.g. Suyehiro, 1941; Hynes, 1950; Hiatt and Strasburg, 1960; Kuthalingam, 1959; Maitland, 1965; Keast and Webb, 1966; Thomas, 1966; Godfriaux, 1969, 1970a, 1970b; Chen and Lee, 1980; Collins, 1981) or quantitatively (usually using more complex mathematical models or equations e.g. Darnell and Meierotto, 1962; Keast and Welsh, 1968; Atmar and Stewart, 1972; Moriarty and Moriarty, 1973; Eggers, 1977; Diana, 1979; Jobling, 1981). Many of the determinations of the 24 hour chronology of feeding intensity in natural populations have been based on the analyses of - 1. Frequency of prey species found in stomachs of a sample of fish (e.g. Diana, 1979) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 91. 2. Weight or volume of food found in the stomachs (Keast and Welsh, 1968; Atmar and Stewart, 1972) 3. Daily variation in the level of food in the stomach based on weight of food and the rate of digestion (evacuation) (Darnell and Meierotto, 1962; Moriarty and Moriarty, 1973; Eggers, 1977; Collins, 1981; Jobling, 1981). Hynes (1950) made a critical review of the methods in common use and stated that "For any fish with a generalized diet, provided a large number of specimens is examined, any of the commonly accepted methods of assessing the composition of the diet of fish from gut contents will give substantially the same results. Those food items important in a diet will always stand out clearly from those that are occasional or rare and so unimportant and the variation between the different methods will probably not be greater than that between different samples of fish .... although the volume or weight methods are probably the most satisfactory, the points method is a short cut to the same result and that the simpler occurrence or dominance methods are likely to give the same result as the points method." The majority of fish are carnivorous and generally capture of their food requires elaborate techniques because potential prey organisms have a wide range of behavioural and structural adaptations for avoiding capture (Keenleyside, 1979). Many morphological features of fish have been intimately connected with specific modes of feeding relationships. The feeding habits of carnivorous fish (especially the trapping of prey) is mostly controlled by the structure of the mouth (Yasuda, 1960c); the mouth and its component structures as well as other body structures have evolved adaptively to ensure an efficient procurement of food. Ridewood (1896) and Gregory (1959) have extens­ ively studied the teeth and skull respectively. In many instances the morphologies of teeth and skull are pointers University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 92. to the feeding habit of fish. For example, Keast and Webb (1966), Yazdani (1969), Nikol’skaya and Verigina (1974), have investigated and correlated the functioning of mouth and body structures with the feeding habits of 14 cohabit­ ing fish species, flatfish and flounders respectively. In most of these works, the attempt has been to understand the mechanical nature of evolutionary adaptations by relating to the mechanics to specific functions. Smith (1982) argues that "A more justifiable orientation would be to evaluate the constraints which limit the mechanical efficiency of structures." Nonetheless, the primary task of a theory of feeding strategies is to specify for a given animal that complex of behaviour and morphology best suited to gather food energy in a particular environment (Schoener, 1971). The species of the family Synodontidae take their collective vernacular names - 1) lizardfish - from their reptilian look (more or less flattened lizard-like heads 2) grinners - from their large mouths with rows of sharp, long and depressible teeth (not covered by lips), the teeth even showing when the mouth is closed thus expressing a permanent 'grin' (Marshall, 1964; Wheeler, 1975; Thomson, 1977; Grant, 1978). Lizardfish are euryphagous. They feed on crustaceans, cephalopods and (mainly) on other fish (Okada and Kyushin, 1955; Toriyama, 1958; *Chervinsky, 1959; Hanoaka, Hayashi, Murakami, Takahashi and Yamaguchi, 1959; Tung, 1959; Hayashi, Yamaguchi and Hanoaka, 1960; Bograd-Zismann, 1965; Tatara, 1965; Budnichenko, 1974; Sainsbury and Whitelaw, 1981; Rowling, unpublished). However Hiatt and Strasburg (1960) and Matsumiya, Kinoshita and Oka (1980) found only fish in the stomach of Saurida undosquamis. * Quoted in Ben-Yami and Glaser, 1974. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 93. Lizardfish are voracious (Marshall, 1964; Arnold, 1951; Hanoaka et al, 1959; Tung, 1959; Sweatman, manuscript). Hanoaka et al (1959) and Tatara (1965) discussed selection of prey sizes by different size groups of S. undosquamis. Budnichenko (1974) found variation in the food composition of S. undosquamis and S. tumbil with size of predator, depth, and seasons. Tatara (1965) reported variations in food composition with different areas. Diurnal activity and variations in feeding of S. undosquamis, S. tumbil and S. elongata have been investigated by Toriyama (1958), Tung (1959), Hayashi et al (1960), Tatara (1965), Budnichenko (1974) and Rowling (unpublished). Descriptions of morphological or osteological features of lizardfish per se have been made by many authors (e.g. Norman, 1935; Gregory, 1959; Agarwala, 1968; Shindo and Yamada, 1972; Rao, 1977; Yamada and Ikemoto, 1979; Waples, 1982 and many others). The burying and hunting behaviour of lizardfish have also been described by Breder (1944), Hiatt and Strasburg (1960) and Sweatman (manuscript). To date little or no attempt has been made to investigate any possible correlation between the striking teeth configur­ ation and body structures and the food and feeding behaviour of lizardfish. Also apart from the works of Sweatman (manuscript), Rowling (unpublished) and general knowledge in books (e.g. Marshall, 1964; Carcasson, 1977 and others) there is no available information on the food and feeding habits of any species of Synodontidae from Australian waters. The object of this chapter is first and foremost to investigate the food and feeding habits of four local species, and then to describe and inter-relate morpholog­ ical structure with way of life, food and feeding habits. The study draws upon a three-year study of the food of 8470 specimens of Saurida and Traohinocephalus and the study of the feeding and digestive system of 14 species of the University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 94. aforementioned genera and Synodus. Various aspects of the morphology which have an important bearing on the success and efficiency of the feeding complex (morphology and behaviour) have been investigated and an attempt is made to draw correlations between the components of the feeding complex. 3.2 METHODS AND MATERIALS 5411, 1340, 612 and 1107 specimens of Saurida undosquamis3 S. tumbil, S. filamentosa and T. myops respectively were used in the study of food and feeding habits. S. undosquamis and S. tumbil were collected monthly over a period of 26 months, mainly from Moreton Bay. S. filamentosa and T. myops collections were made in 15 and 17 months respectively off Cape Moreton. All samples were obtained by forty-minute bottom trawls and were preserved in 5% formalin as previously described in Chapter One (i.e. the general introduction). The sampling covered a wide area, from muddy and sandy or silty bottoms of varying depths (10 - 40m) in different parts of the Bay (see Fig. 1.1) to provide reliable data on the basic food and feeding habits of lizardfish. The variation of feeding with prey abundance was also investigated:- The survey ran concurrently with baseline studies on the effects of enlargement of the Brisbane Airport upon trawled biota, by Stephenson, Chant and Cook (manuscript) between November 1979 and March 1981. Samples were collected at intervals of lunar months (28) days. At each site at each month three other sampling variables were involved: port and starboard nets, trawling with and against the tide for fifteen minutes, and sampling at midday and dusk. For each sample, counts were made of the species collected. 60 species were considered. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 95. No incision was made into the body wall before preserv­ ation in formalin because the preliminary survey showed - 1) that the species feed on quite large items which are to a large extent unaffected by digestion during the time lag until formalin actually seeped into the stomach to stop digestion. 2) substantial loss of body fluids (via incision) and consequent loss in whole body weight and distortion of ratios incorporating whole body weight. Diurnal feeding and activity surveys were attempted on the following days off Tangalooma Point (see Fig. 1.1). Summer 17 January 1980 4 December 1980 Autumn 3 March 1980 13 May 1981 Winter 17 June 1980 Spring 10 September 1980 6 November 1980 10 November 1980 21 October 1981 Adequate data were obtained on only 17 June 1980, 10 September 1980 and 21 October 1981. The other surveys were curtailed with little or no data because of bad weather. The samples were collected at approximately hourly intervals unless otherwise stated. Monthly and hourly samples were kept separate. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 96. In the laboratory, the fish were washed free of excess formalin and dried with paper towels as previously described in the general introduction. After whole body weight and standard length have been measured, the alimentary canal of the fish was removed from the abdomen by making an incision of the body wall between the anus and branchiostegals, severing the oesophagus from the pharynx, lifting out the entire alimentary canal and severing the rectum at the anus. Each alimentary canal was kept separately in a 150ml bottle. The sex of each fish was determined by examining its gonad under a dissecting microscope. Digestion was found to be well advanced in the intestine so examination of food items was confined to the stomach. Stomachs were cut open and items were taken out and identified to the lowest systematic level possible. Fish identifications were according to Marshall (1964), Carcasson (1977) and Grant (1978). Invertebrate identifications followed Meglitsch (1972), Barnes (1974) and Grant ( (Penaeidae; prawns) 1978). The isopod genus Anilocra was identified by Dr. Neil Bruce. The stomachs were given points of fullness 0 - empty 5 - with little food 10 - half full 15 - quite full 20 - full and distended 20++ - greatly distended. This system is a modification of the Swynnerton and Worthington (1940) points method. Here each stomach was allotted a number of points of fullness instead of on categories of food in the stomach. This method, though more or less subjective, was considered appropriate for the study. Compared to other methods, it is rapid and easy and requires no special apparatus for measurement. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 97- Large samples and several months of such analysis undoubtedly diminished the subjectivity in the method considerably. The occurrence method was also considered appropriate in assessing the relative importance of prey items. Each stomach scored only once for each type of prey notwith­ standing the numerical strength or size of the prey in the stomach. Thus the composition of food of lizardfish was reduced to percentages of occurrences of prey in stomach. The Feeding Index; F.I., was calculated as - F . I . = Number of fish with items in stomach x 100% Total number of fish The number of species and their numbers examined precluded measurement of stomach contents of each individual fish. Only whole or sometimes large items were measured for length and weight. The orientation of the head or tail of the prey (if discernible) in the stomach (i.e. swallowed head-first or tail-first) was also noted.* It is likely that some food items were taken in the trawl and **Bograd- Zismann (1961-62) regarded this as "as result of panicky indiscriminate attacking in the trawl cod end." Budnichenko (1974) and Rowling (unpublished) discarded stomachs with perfectly preserved prey for the above reason. No stomach was rejected in this study because - 1) the large number of fish with empty stomachs caught almost always with large numbers of prey and potential prey during the preliminary study indicated that "panicky indiscriminate attacking", if it occurred at all, did so at a low frequency. 2) the premise for rejection of some stomachs - perfect preservation of prey, is spurious because - (i) lizardfish swallow prey entire, and * Between February 1981 and February 1982 ** Quoted in Ben-Yami and Glaser, 1974. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 98. (ii) it more or less implies very fast or instant breakdown of prey taken under "natural” conditions; this is contrary to their findings and those in this study. A simplified approach was taken: Every item found in the stomach was regarded as a food item. Prey items found stuck in mouth or pharynx were discarded. The following measurements were also taken of the lizardfish - a) lengths of head and premaxillary b) width of premaxillary at posterior edge of eye and size of eye c) depth and width of body at origin of dorsal fin d) the triangular base of each half of caudal fin and the span of the caudal fin. The Aspect Ratio (A.R.) of the caudal fin is defined as the span of the foil/chord; this is often more conven- 2 iently stated as span /area (Nursall, 1958). Mouth, jaws, lips, teeth, gill rakers, the entire gut, body form, pectoral and pelvic fins, caudal peduncle and fin were examined macroscopically. A number of heads of lizardfish were boiled and the muscles and skin removed. The maxillary, premaxillary, angular and dentary were then cleaned with a hard brush and studied. Several attempts to keep some lizardfish specimens in an aquarium in the laboratory for feeding behaviour studies were unsuccessful. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 99 i 3.3 RESULTS 3.3.1 Feeding Patterns of Lizardfish The feeding patterns of lizardfish were estimated from the percentage occurrence of (potential) prey items in the stomach. About 50% or more of the stomachs of the species looked at were empty: specifically 58.5, 67.9, 50.8 and 48.4 percent of S. undosquamis, S. tumbily S . filamentosa and T. myops respectively (see Appendix 1). The most important component (by frequency of occurrence) were fish; in over 85% of stomachs, Crustacea; 5 - 16% and cephalopods (mainly squids); up to about 7% depending on species of lizardfish (Table 3.1). Prey items were swallowed whole and many stomachs were distended. The frequencies of occurrence of prey items indicate that lizardfish fed mainly on fish and, amongst other, crustaceans and cephalopods. The food composition included items of many divergent shapes and sizes; some were cylindrical and more or less fusiform (e.g. Sillaginidae, Synodontidae), flattened and depressed (e.g. Callionymidae, Platycephalidae, Heterosomata), compressed and deep-bodied (e.g. Apogonidae, Carangidae, Chaetodontidae, Leiognathidae, Macroramphosidae, Priacanthidae, Sparidae, Trachichthyidae), others were rounded (e.g. Engraulidae), or rectangular; box-like (e.g. Tetraodontidae). The prey items were also representatives of families inhabiting equally divergent *habitats; pelagic, benthic and benthopelagic: shallow and inshore/coastal waters (e.g. Apogonidae, Callionymidae, Cepolidae, Clupeidae, * See Marshall, 1964 and Wheeler, 1975. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 100 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 4-+- I Percentage occurrence ( + ) of items in stomach (%) Number of stomachs University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 101. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 102 T a b l e 3-1 C o n t ’d University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 103. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 104. Leiognathidae, Priacanthidae, Synodonthidae Saurida undosquamis), deep water (e.g. Triglidae, Macroramphosidae, Synodontidae (Synodus s p )). Some live or feed in seagrass beds (e.g. Antennaridae, Parapercidae, Tetraodontidae), in or on sandy bottoms/flats (e.g. Platicephalidae, Theraponidae, Mullidae, Scorpaenidae), or in mud bottoms or burrows (e.g. Annelida, Alphaeidae, Opisthognathidae) and some like the Clupeidae and Engraulidae are pelagic. *Locomotory abilities of the items ranged from the sedentary or crawling (e.g. Antennaridae, Triglidae) to the rapid swimming cephalopods (e.g. Loligo ) and the powerful, active and fast swimming fish (e.g. Carangidae). Some of the prey species too were representatives of families that concealed themselves by total camouflage (e.g. Antennaridae, Scorpaenidae) or by burying themselves in the bottom (Platycephalidae). The respective averages and ranges of percentage sizes of prey relative to the four lizardfish species are: S. undosquamis** - 41.5, 23.9 - 78.7% S. tumbil - 46.7, 16.3 - 84.3% S. filamentosa - 42.4, 18.3 - 80.2% T. myops - 38.5, 25.7 - 62.8% Thus the distention of the stomachs, the various forms and types of prey, the divergent shapes and relative size ranges, various habitats and locomotory abilities indicate that lizardfish are voracious, that they search for food and probably select prey by relative prey size than by prey species. * See Marshall, 1964 and Wheeler, 1975 ** Also see Table 3.2. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh lAuî r, j.z : Relative reLcentdKc v/»/ . i w -- ------------- S. undosquamis S. tumbil Prey item S.L % F. S.L % F. L. woretonieneie 189 30.2 10 261 19.5 15 162 35.2 15 265 18.9 15 180 31.7 15 225 22.2 20 200 28.5 20 306 16.3 15 172 31.4 20 172 30.2 15 157 34.4 15 183 27.3 20 187 24.1 20 180 28.9 15 145 30.4 20 173 30.1 20 Apogonidae 212 33 .5 20 215 33.5 15 177 36 .7 20 160 39 .4 15 180 33 .3 15 190 34 .2 20 158 41 .1 20 197 43 .1 20+ S. undosquamis 110 59 .1 20+ 182 22.0 10 165 25 .4 10 215 23.3 10 177 33 9 15 275 61.8 20+ 140 42 9 10 317 43.6 20+ 178 78 7 20+ Loligo sp. 266 56 4 20+ 270 59.3 20+ 147 37 4 15 140 53 6 15 148 60 8 20 81 43 2 15 176 56 8 20 227 30 8 20+ Merogyrrmus sp. 167 47 9 20 Cynoglossus sp. 174 57 5 20 Pseudorhombus sp. 146 41 1 10 Parapercidae 188 23 9 10 170 18.8 10 Parapercidae 170 31 2 10 182 17.6 10 Platyoephalus longispinus 280 43 9 20+ Engraulidae 204 49 0 20 Mullidae 197 45 7 20+ 252 43.7 20 165 55. 8 20+ Pelates quadbrilineatus 172 39.0 20+ Sillago maculata 172 84.3 20+ 255 64.7 20+ Carangidae 317 42.5 20+ Semi-digested fish 157 44. 6 20 243 39.1 20 196 40. 8 20 S.L Standard length F Fullness of stomach University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 106. 3.3.2 Orientation of Item in Stomach The orientation of whole prey in the stomach; headfirst or tail-first, was noted and Chi squared test of the observed *183 cases in S. undosquamis indicated no signif­ icant difference (p>0.05) from the expected 1:1 ratio. The number of cases of S. tumbil, S. filamentosa and T. myops were statistically too small for such analysis. The prey items involved with S. undosquamis were Apogonidae, Leiognathidae, Carangidae, Parapercidae, Mullidae, Callionymidae, Platycephalidae, Synodontidae, Heterosomata, fish larvae, semi-digested fish, Loligo sp. and prawns. 3.3.3 Variation of Feeding and Prey Abundance Apogonidae, Leiognathidae, Loligo sp and Crustacea occurred quite frequently in the stomachs of S. undosquamis and S. tumbil compared to other trawled biota. The population profiles of these items were correlated with percentage occurrences in stomachs (Fig. 3.1 , also see Appendix 2). Fig. 3.1 shows a positive correlation between the abundance of the prey and its percentage occurr­ ence in S. undosquamis and S. tumbil stomachs, especially in the former. This indicates that lizardfish are more or less responsive to variation in numbers of prey item. 3.3.4 Variation of Feeding and Food Composition with Sex, Length and Month There was no trend in the variation of the composition of food items with size (length) or sex in fish. Fish of * Number of headfirst = 97 Number of tailfirst = 86 X2 = 0.6612, l df. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 3. : Monthly variation of prey abundance and frequency of occurrence in lizardfish stomachs. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 107. standard length < 100mm (mostly very young fish) fed on small sized prey - very young S. undosquamis fed on fish, fish larvae, crustaceans (prawns), Loligo sp. and annelids. Very few S. tumbil and T. myops 4 100mm were obtained and their stomachs contained fish larvae and semi-digested fish. No young S. filamentosa were obtained. The feeding indices of the four species; the percentage of stomachs with food to total number of stomachs examined, showed no trend with season or year. However, the female average index was higher than the male average index (Table 3.3). 3.3.5 Daily Variation in Feeding and Activity Using number of trawled fish as a criterion for activity, Fig. 3.2 shows higher percentages between 1000 and 1800 hours. The trend in Fig. 3.2 indicates higher diurnal activity in S. undosquamis. Stomach content analysis indicated that the specimens examined fed during both day and night. Any effect of tide levels on activity of S . undosquamis was not evident (Fig. 3.2, Appendix 3). 3.3.6 Morphology of Synodontidae Associated with Feeding The following general accounts are the results of the study of fresh and preserved specimens of 14 lizardfish types: 11 Indo-Pacific Saurida species, 2 Synodus species and T. myops. 3.3.6.1 Mouth, Body Cavity and Gut The mouth is large, terminal and the buccal cavity is small. The premaxillae are elongated; about 61 - 76% of University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig, 3.2 : Daily activity of s. undosquamis University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh LjS.U Jo j a q u u n u 9 6 e ^ u a o j 9 d University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh TABLE 3.3 : FeedinR Indices S. undo8quamie S. tmbil S. filament0 8a T. my Op 8 Month and Year male female male female male female malefemale November 1979 84.4 66.7 December 1979 9.5 10.0 64.7 55.6 January 1980 18.2 8.3 40.5 22.6 February 1980 70.8 78.5 38.9 26.3 March 1980 58.2 57.2 31.3 50.0 April 1980 26.6 28.2 85.0 88.9 May 1980 35.9 27.3 0.0 0.0 40.0 0.0 June 1980 29.3 37.1 15.8 12.5 76.5 0.0 100.0 0.0 July 1980 36.5 31.6 4.3 0.0 66.7 0.0 August 1980 54.2 35.3 0.0 0.0 36.4 33.3 September 1980 51.0 39.1 79.1 80.0 58.8 33.3 100.0 100.0 October 1980 65.5 55.4 0.0 0.0 0.0 60.0 55.6 42.9 November 1980 29.4 37.1 0.0 6.7 100.0 100.0 63.1 40.0 December 1980 35.5 41.4 0.0 6.0 79.7 63.6 January 1981 5.0 8.3 47.2 27.4 66.0 50.7 95.9 31.3 February 1981 50.0 45.6 50.0 25.0 38.9 20.0 25.0 50.0 March 1981 47.4 42.5 14.3 18.2 44.4 33.8 0.0 100.0 April 1981 35.4 39.6 36.7 34.6 0.0 0.0 20.0 18.2 May 1981 32.5 23.5 41.4 36.8 42.0 62.5 June 1981 37.9 25.9 28.6 41.7 40.0 50.0 July 1981 16.5 25.6 7.1 9.5 66.7 100.0 August 1981 32.2 25.7 8.7 16.7 September 1981 41.8 42.2 26.3 16.7 66.7 50.0 October 1981 30.8 32.5 22.2 30.8 59.2 50.6 November 1981 45.2 22.6 57.1 65.2 December 1981 36.8 61.5 January 1982 February 1982 33.3 100.0 76.1 40.5 Average * 39.1 36.5 29.0 27.5 49.6 41.6 69.8 48.5 Very young fish 57.8 16 0 71. 9 * Very young fish excluded University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 109 . the length of the head (see Table 2.4B) and the premaxillae articulate further behind the orbit with the angular bones. The maxillae are rudimentary and also excluded from the gape (Fig. 3.3B). The gill arches extend far into the mouth, well in advance of the angle of the gape. The first four lower jaw arches each has an additional joint (Fig. 3.3C). The anterior section of the lower jaw is movable (downwards only) about two joints at the base of the first and fourth gill arches. This structural frame further increases the gape of the mouth. The gill rakers are rudimentary; pads of teethlike setae. The body is muscular and slender, rounded and fusiform or slightly depressed. The ratio of the width to depth at dorsal fin origin is about unity. The lizardfish has a rather large body cavity. The oesophagus and intestine are short. The stomach is ""f" shaped in situ3 sensu Suyehiro (1941). The oesophagus and stomach have very thick walls, the inner section with thick longitudinal folds. The oesophagus has two bands of bristles, each overlaid by a thick bundle of muscle fibres. Lizardfish have about 13 - 21 pyloric appendages. Liver lobes are thick, and the left lobe a little longer. They have no swim bladder. The thick wall of the oesophagus and stomach allows the great distensibility of the gut; especially of the stomach. 3.3.6.2 Teeth The teeth of lizardfish are moderately sized, sharp, conical to sagittate (Fig. 3.4A) and depressible. The premaxillary and dentary carry about 3 - 6 bands of teeth. The roof and floor of the mouth bear a number of groups of teeth: roof - the palatine has one (in Synodus and University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh sa g itta te i t t a t e conical (curved) A :.Types of teeth B^Cauclal aspect v 2 A spect span stereoscop ic Ratio Area vision ■ 4h b 3 , the fish becomes disproportionately heavier as it grows larger. Equations 4.1 and 4.2 assume that fish grow towards some theoretical maximum length and weight, and that the closer the length or weight gets to the maximum, the slower the rate of change or size or weight; the corollary being the larger K, the faster W _ or L ^ i s reached. 00 °0 The above classical equations usually describe central tendency or mean value behaviour; they do not acknowledge stochastic variation of the individual fish and provide no explicit construction for moving from observable characteristics of individual fish to an aggregate represent­ ation of biomass (Cohen and Fishman, 1980). Cohen and Fishman (op oit) proposed a model designed to approximate the real world. They ’attempted to shift the emphasis in characterizing of fish population from a pre-occupation with University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 132. 'average1 behaviour to an awareness of distributional considerations in length, weight and biomass through time.” However exactitude in growth models such as that proposed by Cohen and Fishman (1980) is probably more academic than practical. Errors in age determination and the effect on growth estimations have been discussed by. amongst others, Brander (1974), Carlander (1974) and Ricker (1981). Studies on the growth of Synodontidae species as deduced from the available literature have been on only two species: Saurida undosquamis and S. tumbil of the Indo- Pacific region where they are the basis of important commercial fisheries. The scale method has been used to estimate ages in populations of S. undosquamis (Tatara, 1953, 1965; *Bograd- Zismann, 1961-62; Budnichenko and Nor, 1978) and of S. tumbil (Tung and Liu, 1965; Okada and Kyushin, 1955; Yeh, Lai and Liu, 1977; Budnichenko et a l , 1978); all in the temperate region. Sinoda and Intong (1978) and Rao (unpublished) working in the tropics found the scales of S. undosquamis unsuitable for age determination and relied on length-frequency analysis. Sinoda et al (1978) reported a "single" prominent mode which seems to consist of several groups" in their preliminary results on S. undosquamis, whilst Rao (unpublished) noted an indication of a number of broods in the year class of S. tumbil. Sainsbury and Whitelaw (1981) supplemented otolith readings with length-frequency analysis in the estimation of ages of S. undosquamis in the North-West Shelf of Australia. Time of ring formation in scales have been correlated with spawning in S. tumbil in the Yellow and East China * Quoted by Ben-Yami and Glaser, 1974 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 133. Seas, and Taiwan Strait (Okada et a l , 1955; Tung et al, 1965; Yeh et a l , 1977) and in the Arabian Sea (in also S. undosquamis, Budnichenko et al, 1978.) Maximal increase in size have been observed in these species in the early years of life (Budnichenko et al, 1978; Tatara, 1965) and differences in growth due to sex have been reported by (Okada et a l , 1955; Tatara, 1965; Budnichenko et al, 1978). The asymptotic lengths reported after the fitting of data to the Von Bertalanffy equation range from about 640 to 790mm in S. tumbil (Yeh et al, 1977; Tung et a l , 1965; Rao, unpublished) and 400 to 571mm in S. undosquamis from the inner Gulf of Thailand and the North-West Shelf of Australia (Sinoda et al, 1978; Sainsbury and Whitelaw, 1981). The only information available on the age and growth of lizardfish species in Australian waters is the report submitted at CSIRO Divisional Research Seminar by Sainsbury and Whitelaw (op oit). The purpose of this study is to fit observed data on S. tumbil, S. undosquamis3 S. filamentosa and Traohinooephalus myops to the classical growth equations, and relate their features of growth to their life histories. 4.2 METHODS AND MATERIALS Measurements of length, weight and the determination of sex followed the same procedures as described in Chapter One. After having noted the above, scales were taken and cleaned in water. These were then mounted between two microscope slides and examined under a dissecting microscope with transmitted light at a magnification of 60x or 120x. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 134. On the other hand, vertebrae were sectioned with a sharp scalpel and the cut surfaces were examined under the microscope in glycerol against a black background using reflected light. Otoliths were examined whole using transmitted light. The criterion for selection of tissues was the clarity of rings. Scales were chosen for the determination of ages in S. tumbil, S. undosquamis and T. myops, whereas vertebrae were chosen for S. filamentosa. Otoliths were rejected because they were decalcified during preservation which resulted in the blurring of the ring patterns. The scales had in addition to the clear rings (i.e. complete breaks in circuli) cutting or crossing over of crowded circuli within the basi-lateral and lateral areas (see Fig. 4.1). These were ignored. On the other hand, dark zones alternating with translucent zones were seen on the cut surfaces of vertebrae of S. filamentosa. These were assumed to represent annual growth marks. In order to *fulfil Van Oosten's (1929) conditions for validity of age determination techniques - 1) analysis of variation in scale morphology with portion or site of body was undertaken to find site with least variation 2) recognition of rings was based only on complete break(s) in ring circuli which could be traced all round the scale, and 3) periodicity of ring formation was investigated using the formula - ^ _ R ~~ rn x 100% (....equation 4,4) r -r . n n-1 where R is the radius of scale r and r are n n-1 radii of ultimate and penultimate rings respect­ ively. o(, is the monthly change of the growth rate of marginal region of scale. * was very difficult in the case of vertebrae. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4-1 Scale of L i za rd f i sh B Fig. 4-2 : Scale and Body si tes University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 135. Scales were taken from the same row from the left side of each fish. Regenerated or lost scales were made up with scales taken from the same row on the right side of the fish (see Fig 4.2). Scale radius and ring radii were measured (along the line AB, Fig. 4.1) with an ocular micrometer mounted with the right eye-piece. The ocular micrometer had a conversion factor of 1 unit = 0.8mm at a magnification of 60x; calculated from a graduated stage micrometer. Alizarin treatment of scales after having washed scales in 5% sodium hydroxide and generous amounts of water did not significantly enhance readability of rings and was thus discontinued. Samples of ringed scales were photographed. The age-length determination as obtained from ring counts on scales and vertebrae were compared to age- modal mean lengths from the Cassie (1954) Curve. The growth patterns of the four species were described in terms of Von Bertalanffy and other growth parameters. Statistical analysis followed Senter (1969) and Sokal and Rohlf (1969). 4.3 RESULTS 4.3.1 Variation of scale morphology with body site The scales of the four species are cycloid with 3 to 6, but usually 3 radial grooves from the focus which is about a third of the diameter of the scale from the apical margin. The circuli are concentric (see Plate 38). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh A = 3 year old scale B = 2 year old scale University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 136. The regularity of scale morphology with body site was investigated by comparing scale sizes from the various sites (see Fig. 4.2). Coefficients of variation per body site were calculated for groups of six scales’ radii for a number of fish specimens. Site 10 was chosen as site from which scales were taken for study because it had the least coefficients of variation and least probability of regener­ ation (Appendices 4A, 4B, 4C). 4.3.2 Definition of rings on scale, time of ring formation and age determination 4.3.2.1 Definition of rings on scale Regression analyses of *ring radii on scale radii for 4-, 3-, and 2-ringed scales of specimens of S. tumbil showed significant relationships - the degree of relation­ ship ranged from moderate to very high correlation; all significant at p ^ 0.02 (Fig. 4.3, Tables 4.1A, 4.IB). The t test for differences between the mean radii of contiguous rings also indicated significant differences between means in the 3-ringed female, 2-ringed male and the first three rings of the 4-ringed female scales (Tables 4.1A, 4.IB). The difference between means of the third and fourth ring radii of the 4-ringed female was only significant at a higher probability level (i.e. p ^ 0.1). This was probably due to slowing down of growth after the third ring had been laid down and thus a disproportionate little spacing between the third and fourth rings compared to spacing between the first three rings. 3-ringed male scales showed no significant differences between means of * Though similar rings were found on scales of S . undosquamis and T. myops, the smallness of their samples with 2 or more rings precluded satisfactory statistical analyses. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 137. ------ — CO t> CO CO* ✓I"IN o S CM CM tH CD CO 8 -P CM in O CM 0005 8 tH CO + 1 n o CM rH LO O P r->. 05 CD rH II O O CM l> o 05 r—1 CO a CM CD 00 o $ + 1 00 00 o 05 o CO II o CO V -/ in o -p CQ T3 bp a V, +-> 3 05 CO U CQ CQ 9 .155 u CH £ £ P ta fH 8 CO P 1 P Table 4.1A: S. tumbil - Definition of rings on scale FEMALE 4+ 16 Mean scale radius 8.15625 Standard deviation 0.773646 First Second Third Fourth First Second Third Mean radius of ring 5.14375 6.28750 6.99375 7.65625 5.19189 6.24054 6.93784 Standard deviation of ring 0.9280580 0.800729 0.894031 0.880885 0.952009 0.681689 0.689344 Correlation of ring 0.5874 to scale radius 0.9496 0.9543 0.7731 0.8742 0.9221 1.9891 Critical value of t 2.°42 , t 10( University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 138. 8 m LO 05 300 o H I /—N o 8 r'—H✓ tH ! -pCQ CD tHO 00 tH CQ 73 bo a T3 q CD 5 | 8 S§ ■8 3 CM P. m 0 0m5 oCD 5 in o coo CMO 00 00 CcoO CCMO CDCM w •H o CO m m tj mi 00n5 CT—O 0 oo 00 1 00 0005 CM U o 05 05 00 0 i n CO tH in i—I Q& O w SH CO 0 C 3r co •H CO O "§ 8* 0 ft Is CO Co Table 4.3 : Relationship between standard length (SL in mm) and University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 144 CO J 00 T—1 00 o o csi CD »J csi w w w J £1 w -P < • J 2S hfl <£ C w a) pH i—i TJ a) p LO G) oo o ctf s tH CO rH rH CO io 3 o rH rH CD o rH LO LO d J * CO a G) s i rH rH o d CQ ja JC bl) § s rH females, n - 350 H - S. filamentosa j males , n - 278 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh I a s s ( mm) hoz University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 146 co Xi -p bfi a0 C3O CM T3 u fl d -P CO T05 -P d O00 rH O O U0 CO CQ (sM XJ O CM $ ■8 o tH LO Jh 00 CO ■§ rH C+SI ■§ 00 CD O CO toH OtH CD CO tH tH O tH H tH T3 O i—i fH > Js + CD to tH W rO 0G ,Q O H o t0H r0H cc V CO o E Kr* o: CO S •< 1CQ Co Co . Table 4.5B : Growth of lizardfish; calculated #whole body weight (g). Key calculated whole body weights as fran the following Species rl=regression line, von=Bertalanffy equation, *exp=exponential equation Age von von von von von Sex exp exp exp exp exp 76.2 210.9 325.7 423.7 female 243 174.7 257 207.9 270 242.4 280 156.4 193.3 ______ 235.2 273.6 58.6 129.3 188.1 male 165 52.5 204 101.5 234 155.4 48.1 86.7 ______ _____1_36.5 222.4 327.3 female 250 169.8 273 222.3 293 276.0 255.7 361.7 51.5 176.5 279.3 male 155 39.3 200 85.8 237 144.2 33.1 72.1 137.0 656.2 1015.9 1714.9 female 240 148.0 290 268.4 350 385 203.3 410.0 617.3 73.0 232.4 363.8 472.3 561.7 205 90.1 233 134.8 268 209.4 289 265.4 314.6 76.5 117.4 200.5 276.5 353.2 78.8 210.8 321.7 female 193 109.8 218 161.6 239 216.4 103.0 172.2 265,2 31.5 96.7 148.9 125 27.7 155 54.8 179 86.5 25.4 47.1 77.2 Exponential values calculated frcm separate male and female equations. See Appendices 5A-D for observed mean whole body weight. L'Ya 9 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4.6A,B : Cassie curvesfor 478 females (A) and 475 males (B) of 5. tvmbn. Each continuous curve represents the percentage cumulative frequency of each (whole) sample, and the oblique straight lines represent the percentage cumulative frequency in each age group. Arrows(J) indicate points chosen as dividing the successive age groups. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4.6C,D : Cassie curvesfor 2606 females (C) and 2492 males (D) of s. undosquamis. Each continuous curve represents the percentage cumulative frequency of each (whole) sample, and the oblique straight lines represent the percent cumulative frequency in each age group. Arrowstj,) indicate points chosen as dividing the successive age groups. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4.6E,F : Cassie curvesfor 497 females (E) and 537 males (F) of T. my op s. Each continuous curve represents the percentage cumulative frequency of each (whole) sample, and the oblique straight lines represent the percent cumulative frequency in each age group. Arrows(j) indicate points chosen as dividing the successive age groups. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig, 4.6G,H : Cassie curvesfor 350 females (G) and 278 males (H) of s. fiiamentosa. Each continuous curve represents the percentage cumulative frequency of each (whole) sample, and the oblique straight 1inesrepresent the percent cumulative frequency in each age group. Arrows (I) indicate points chosen as dividing the successive age groups. University of Ghana U n i v e r s i t y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh •*« University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 148 . The lengths of contiguous age-groups overlapped considerably (Fig. 4.5, 4.6A-H). The overlaps and the crowding of older fish into the upper part of the percent­ age cumulative curve reduced the accuracy of the Cassie curve values (Table 4.5A). Nonetheless Cassie curve values were concurrent with observed mean values - they were generally within the ranges of the observed values. 4.3.6 Growth equations and growth curves Hirschhorn (1974), showed that the exclusion of older ages from data sets affects the estimation of Bertalanffy parameters and is therefore inadvisable. In his studies of growth parameters of three fishes and a mollusc he noted that "The ranges of ages used may differ through excluding mean lengths based on few fish owing to their * ’unreliability' or through abandoning the difficult outer rings when using back-calculated lengths.” 4.3.6.1 Formalae used in the calculation of Von Bertalanffy and other growth parameters. The parameters were calculated directly or derived from one or a combination of the classical equations (i.e. equations 4.1 to 4.3). L is the X-axis intercept of the regression of Lt + 1 on Lt (i.e. Walford plot, Figs. 4.7A, 4.7B, see Ricker, 1975; Ricker 1977). K = -Ln b where b is the regression coefficient of the Walford plot line. * Quotes mine. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh S tandard l eng th L^(mm) Fig. 4.7A, Walford plots of lizardfish ( •- female, o- male) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Standard Length L . +1(mm) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 149. L - L, t t + A (Ln (-2--- ^)) ( .....equation 4.8) o T W. t )) ( .....equation 4.9) %00 where L is the mean length in the catch (sample) and Lc is the mean length of entry into the catch. (See Gulland 1968; 1977) The annual mortality rate; A = 1 - e and the rate of survival; S = 1 - A (See Ricker 1975). The life span (A pc.) was calculated sensu Taylor (1958): A - t .95 o — g— 4.3.6.1.1 Von Bertalanffy growth equations and parameters Computations of the Von Bertalanffy growth equations and parameters were based on observed age-group mean lengths in lieu of individual lengths at age: S. tumbil -0.1592 (t +3.5619)) Female = 425 ( 1 - e Male Lt = 337 (1 - e"-00,.11823 (t + 3.1784)^ S. undosquamis Female Lt = 450 (1 - e-°-1398 (t +4.6691)^ Male Lt = 407 (1 - e~0 -1957 (t + 1.4551)^ University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 150 . S. filamentosa -0.1574 (t + 1.7847)) Female L̂ _ = 654 (1 - e -0.1927 (t + 1.7815)^ Male Lt = 438 (1 - e T. myops -0.1744 (t + 2.9845). Female L^ = 375 (1 - e ) -0.2331 (t + 1.6120). Male Lt = 280 (1 - e ) The females had larger asymptotic lengths and longer life spans (Table 4.6, Figs. 4.7A, 4.7B) which suggest sexual dimorphism. This is corroborated by the larger mean size per age group and predominance of females in the larger size groups. The K values also indicate that the rates of approach to these asymptotes are slower in the females. Theoretically tQ values suggest that growth in size according to the Von Bertalanffy equation began more or less at birth. t'Q were however larger and suggest that growth in weight started later than growth in linear size. This might be a strategy of ensuring larger young lizardfish; making them less vulnerable to predation and therefore the survival of a higher percentage of the young. L , K and tQ estimated from back-calculated lengths of S . tumbil (female) were considerably lower than those estimated from observed values (Fig. 4.7C). This further substantiates the operation of Lee's phenomenon on the female population. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh fl d Cf * fH LO Ot ^ s o csi CO i—I d) >» K >> ID CO CD co 00 ch w Q) d •H CO H J oo 00 CO CD CO CO CO LO CO lO t > rH rH CSI LO csi [> o CD CSI rH 00 CSI o CD O ^ O 05 05 csi CO 00 CD t > CD bD Jh 05 ̂ 00 ̂ CSI O LO rH CD CSI 00 o o ̂ LO rH txD 05 ^ CD O ID O CD ^ CO 05 t> 0 5 COcsi CO CD(D ĉsi rH lO (M CO ^ ^ U 05 rH LO lO LO CD 0005 oLO C0S00 0 5 5I o LO m o 00 0 rH d e CO (D rQ •H O 5 t-'-i CO CM CD CD q S H -H 00 CM 0 0 lO (Ji 0 0 CO CM CO 0 0 CO q ti CM t > o lO CM rH CM tH rH (D CD S rH P •H rcHi (S) CD CO 8B PU < CO 00o’ O >» ■p •H r rH CH$ tH CMLO 0 0 cd -p o 0 0rH CO -P U N rH tH £ § o tH rH CM CD w CD •H O 0 ft T-̂ CO CO Table 4.6 Von Bertalanffy and other growth parameters. ______ (B)___ ______ ______ ______________ Value does not include very young University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4.7C. Walford plot of back-calculated lengths of 5. twnbii (•- female, o- male) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh if) cr> c S. tumbil3 S. filamentosa and T. myops respectively. ** No such estimation was possible in S . filamentosa and T. myops because of missing monthly samples. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4,8 Time of ring formation (A) and mean monthly standard length and weight (B,C) of lizardfish. Legend for B and C Standard length Weight female y + male o • female + male + very young (sex A © indeterminate) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh S t a n d a r d l e n g t h ( m m ) W e i g h t ( g ) M o n t h and Y e a r University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 154. 4.3.6.3 Relationship between body weight and standard length ♦Plots of mean whole body weight and mean standard length (Appendices 5 A-D) indicated a curvilinear (or exponential) relationship. Each sex was plotted separately (Fig. 4.9 A-D). The curvilinear relationship; W = aL (....equation 4.3), was analysed by plotting its logarithmic transformation: Log W = Log a + b Log L. The exponent ' b' is the slope of the regression line (i.e. regression coefficient) and the constant 'a1 is the intercept of the line on the Y-axis. The regression line was calculated by the least squares method (Table 4.7). Covariance analyses indicated no significant difference at the 5% level of confidence (Table 4.8) thus the scatter or dispersion among the two sexes was well within the limits of variation expected by virtue of peculiar length- weight relationships of the individual fish. The length-weight equations resulting from the pooled data are - S. tumbil W = 6.8368 x 10~6 (L3-1051) S. undosquamis W = 7.7836 x 10-6 (L 3 -0605) S. filamentosa W = 4.7831 x 10-6 (L3,1470) T. myops W = 6.1359 x 10-6 (L3 '1733) ' b T in each species was greater than 3. This indicates that the species became heavier for length as they grew larger. * Weights of individuals with greatly distended stomachs, i.e. very full stomachs were discarded. The whole body weight included weight of gonads. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 4.9A. Relationship of weight to standard length of s. tumbii. (•-female, +- male) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh (6) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 4.9B, Relationship of weight to standard length of s. undosquamis. (•-female, +- male) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh (6) }M & 19 M University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 1500- 1X0- 1100- 900- 700- 500 + • + 300- 100 100 200 5oo 400 500 eoo S t a n d a r d l e n g t h ( m m ) Fig. 4.9C. Relationship of weight to standard length of s. filamentosa (•- female, +- male) W e i g h t ( g ) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 155. 00 CM CO rH CM O o PQ LO LO CO LO o 0) t> 00 LO rH Gi CM CM CO CNJ CO* CM CM CM* CO* CM CD oo> 8 LO rH CO O LO O CD* 0> CD rH LO rH o> o o CO CO CO CO CO CO CO co* CO M h a o 00 «H o 8 LO CM 05CO CD 0) rH CD rH m ll W o lO 00 o o 0005 rCHO o* ✓—■S. /—\ /-*\ /•—N /—■> »—N/̂v u £ '—✓ Wv£—- '—* >—£/ 8 8 8 ►J -po J p o o -P o 8 8 € P3 -p po o►J J •p ►J I r—a X +■ = b sK I oo tH w Q) •H o 0 ft c w 4- s —Dt 6 Table 4.9: Von Bertalanffy growth parameters vrs latitude. S . undosquamis S . tumbi1 Sex female male female male Author Area Latitude Method Parameters South East scale and 450 407 425 377 Adjei Queensland (Moreton 27° 26'S length - 0.1957 0.1592 0.1823 Bay) frequency -4.6691 -1.4551 -3.5619 -3.1784 794.6 Yeh et al Gulf of 0.0965 0.0790 (1977) -1.5430 -1.7934 742.4 686.9 Southern 27° N East 0.1043 0.1177 China Sea -1.4160 -1.3953 644 479 Tung et al Taiwan 23° N 0.3169 0.3486 (1965) Straits -0.2486 -0.1021 North-West 15° S otolith and 571 466 Sainsbury shelf of length — et al Australia frequency 0.39 (1981) 0.2 637 Rao 15° N (unpub­ India frequency 0.249 lished) -0.334 Sinoda et Gulf of al 1978 Thailand 10 - 12 N Length-freq. 400 (UIU) 03 rj University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 164. effect of temperature but stressed that "L^ would decrease but rather slowly with increasing temperature”. In Table 4.9 trends, if any, are obscured probably by the different methods used by the authors and also the sensitivity of L to the slope of the Walford Line, *”K”. The data 00 however suggest larger L in the northern hemisphere; 00 especially in the more northern populations. The numbers of any animal population change considerably in time. In fishes the variability of the annually recruit­ ing year-classes provides the main source of such temporal fluctuations. Recruitment is highly variable in S. undosquamis (Tatara, 1965; Sainsbury et a l , 1981). Recruitment sizes(and age) in S. undosquamis and S. tumbil in Moreton Bay were estimated to be about 90.4 (and 0+) and 95.8mm (and 0+) respectively. The recruits appeared after the spawning season. Okada et al (1955) and Sinoda et al (1978) also reported recruitment after the spawning season. However Sinoda et al (1978) estimated the mean size at recruitment of S. undosquamis to be about 150mm in total length in the Gulf of Thailand. Ben-Yami and Glaser (1974) on the other hand estimated recruitment of this species to be between 200-300mm and 2-3+ along the Mediterranean Coast of Israel from the analysis of Bograd-Zismann1s (1961-62) data. The calculated total mortalities in male and female 5. undosquamis and S . tumbil were 2.1767, 1.1382, 1.1087 and 0.4151 respectively and are probably due to a number of factors. S. undosquamis and S. tumbil were hosts to a number of parasites. The most common parasite was the trypanorhynch tapeworm Callitetrarhynchus gracilis. **In a preliminary analysis, no significant correlation was found * K = -Ln Slope. ** See Chapter on Parasites. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 165. between a number of parasites and length of the lizardfish. However the truncated form of the negative binomal dist r i b ­ ution fitted to the frequency distributions suggested that some heavily infected fish were missing from the samples and most probably due to the adverse effects of the larvae on the general fitness of the fish. The m a l e s were affected by lesser numbers of larvae than the females. This difference probably contributes to or r e flects in the differences in total mor t a l i t y between the sexes. In South East Queensland, l i z a r d f i s h f o r m a large percentage of ’t r a s h’ fish t a ken w i t h p r a w n s a nd o t her resource fisheries (pers. comm., L es Wale, B o b Sanderson, Bryan Wallis, p e r s o n a l o b s e r v a t i o n ) . A nd the l a r g e n u m b e r of boats involved in these fisheries must therefore indicate substantial fishing mortality. Comparing male and female total mortalities, supposing that there were no differences between sexes in fishing mortality, the differences in total mortality between sexes would be due to natural mortality. The larger mean sizes of females probably makes them more susceptible to fishing mortality thus Rosa Lee's phenomenon in females. Tatara (1965) found higher total and natural mortalities in males but the same fishing mortality in both sexes in his investigation of sub-populations of S. undosquamis in the seas of Japan. Sainsbury et al (1981) reported that total mortality in S. undosquamis increased with age. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 166. 5. R E P R O D U C T I O N OF F O U R LO C A L S PECIES 5.1 INTRODUCTION Reproduction is the cornerstone for the continuation of species. As any other life process, reproduction demands three basic elements: energy, materials and time. Availability of the first two elements are constrained within limits imposed by the species genotype and environ­ ment. The strategy adopted by an animal or plant is a compromise allocation of energy to the various aspects of its life-history, each of which contributes to total fit­ ness (Begon and Mortimor, 1981). Fish species live under a great variety of conditions and have of necessity evolved adaptive reproductive strategies that guarantee the perpetuity of their type. These strategies usually relate to the egg and larva (Nikolsky, 1963). The literature on the reproduction of fish is immense. Hickling and Rutenberg (1936) studied the ovary as an indicator of the spawning period of four species. They suggested that measurement of the diameters of eggs may provide information as to the spawning habits of fish. Many researchers (e.g. Radhakrishnan, 1963; Kagwade, 1970- 72; Hislop and Hall, 1974; Morse, 1981) have since used this method to establish spawning periods of many fish. Russell (1976) described the eggs and planktonic stages of fish in the British Isles. Nikolsky (1963) discussed the various aspects of reprod­ uction and development of fish. He noted that fecundity of fish varied with species, size, age, environmental conditions, food and population size. The course of matur­ ation of gonads as manifested in changes in the gonad index (G.I.; the weight of the gonad as a percentage of the weight of the whole body), rises with the approach of spawning, University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 167. peaks and decreases after spawning. Time, duration and the place of spawning are adaptations which ensure the most favourable conditions for the development of the eggs and larvae. Hubbs (1943) and Balon (1975a) considered the terminology of the early stages, whereas ecological groupings or guilds have been proposed by Kryzhanovsky (1949) and Balon (1975a) based on reproductive strategies. Bagenal and Braum (1968) compiled methods of investigation on development and reproduction. The standard references on reproduction have been developed for fishes from the temperate regions and Qasim (1973) discusses these methods with special reference to works from the tropical Indian sub-continent. He suggested that ’studies should not be restricted to sketchy observ­ ations on spawning, but should be planned to develop an understanding of the maturation process of the population in a particular area". With exception of two species, the literature on the reproduction and larvae of lizardfish species consist of anecdotes (Norman, 1935; Delsman, 1938; Gopinath, 1946; Nair, 1952; Gibbs, 1959; Mito, 1961; Mead, 1966, *Tiews, 1972; and Ben-Yami and Glaser, 1974). The two species which have been studied most are Saurida undosquamis and S. tumbil. Budnichenko and Nor (1978) and Budnichenko and Dimitrova (1979) reported an extended spawning season for S. undosquamis in the Arabian Sea. However, Sinoda and Intong (1978) concluded from studies in the Gulf of Thailand that S. undosquamis spawns all through the year. Chervinsky (1959)** and Bograd-Zismann (1961- 62)** reported the occurrence of ripe, nearly ripe and spent females almost throughout the year in the Mediterranean * Quoted in Budnichenko and Dimitrova, 1979. ** Quoted in Ben-Yami and Glaser, 1974. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 168. Sea. Tatara (1965) on the one hand found different extended spawning periods for two sub-populations of S. undosquamis in the East China Sea. S. tumbil has also been found to undergo extended spawning in the Arabian (Annigeri, 1963; Budnichenko and Nor, 1978; Budnichenko and Dimitrova, 1979) and East China Seas (Yamada, 1968a,b; Yamada, Tagawa and Mako, 1968). Significant differences have been found in sex ratios and sizes amongst males and females of S. undosquamis (Tatara, 1965; Latif and Shenouda, 1973; Budnichenko and Nor, 1978; Budnichenko and Dimitrova, 1979) and S. turribil (Yamada, 1968a; Budnichenko and Nor, 1978; Budnichenko and Dimitrova, 1979) especially during the spawning season. Yamada (1968b) reported on diseased ovaries and S. tumbil has been described as sexually dimorphic; the male having a prolongation of the second soft dorsal ray at lengths greater than 250- 300mm by Okada and Kyushin (1955) and Liu and Tung (1959). A number of aspects of the reproductive ecology of four local species were studied and are reported on in this chapter. 5.2 METHODS AND MATERIALS The four species - S. undosquamis, S. tumbil, S. filamentosa and T, myops - were collected by bottom trawls and treated with formalin as previously described in the main introduct­ ion . Each fish was measured for length and then was dissected to determine the sex and state of gonad development. The gonads were weighed as discussed previously. Data on each fish were kept separate under its species, reference number, month and year. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 169. Gonads and ova were examined under a dissecting micro­ scope with reflected and transmitted light respectively. Four types of ova were distinguished: immature, maturing, mature and ripe ova. Fifty of each type of ova were randomly selected from ovaries in each stage of maturity and measured along their diameters on a graduated micrometer. Unlike the female, the stages of development of testes were difficult to establish. In the female, the stages were based on types and quantities of ova present, whereas in the male discrimination depended only on the macroscopic appearance and texture of the testes. Fresh specimens of ovaries and testes at the various stages which were available during the preliminary investigation were labelled and preserved. These were subsequently used as standard references. In the preserved testis the texture was the main discriminating feature between the stages of develop­ ment . The stages of gonad maturity or development recognised were modified from Kesteven's (1960) general classification of maturity stages. In the preliminary survey, Stage II of the ovaries and testes were found to weigh only about 0.2g or less, the accuracy of the weighing machine being - O.lg. Each of these, for the convenience of speed and also for inclusion in the calculation of the gonad index was given an arbitrary weight of 0.2g if found to be less. The estimation of fecundity was complicated by - 1) ova being of four types and sizes 2) fractional spawning by the species. Absolute fecundity - the number of ova contained in the ovary of a fish (Nikolsky, 1963) - was estimated with Stage IV ovaries. This stage of maturity of the ovary had predominantly mature ova with a few tiny immature ova in the interstices. Partial fecundity - defined here as the number of ripe eggs to be released at the next fractional University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 170. spawn - was estimated from the average of the sub-samples taken consistently from 3 sections of the ovary: the distal, middle and proximal regions. All fecundity estimations were by the gravimetric method (see Bagenal and Braum, 1968) using average counts and weights of sub-samples in proportion to weight of ovaries. Ova were counted in a counting chamber under a dissecting micro­ scope with transmitted light. An attempt was made to grow artificially fertilized eggs of S. tumbil. However, whereas many of the ova were damaged by mechanical disturbance and temperature changes during transportation to the laboratory, the few which survived did not develop any further than the first few cell divisions. Gonad index, G.I. or the coefficient of maturity, expresses the weight of the gonad as a percentage weight of the whole body (Nikolsky, 1963). The relative fecundity. R.F., is the ratio of the partial fecundity to the standard length or whole body weight. S. undosquamis, S. tumbil and T. myops were aged by scale reading, whereas S. filamentosa was aged by the vertebrae. Data on water temperatures and rainfall were supplied by the Department of Science, Bureau of Meteorology, Brisbane. Since water temperatures can be expected to lag behind air temperatures, mean monthly minimum and maximum temperatures between 1958 and 1981 at Cape Moreton Lighthouse were used as approximations of those which prevailed during the study. The mean rainfalls at Cape Moreton Lighthouse were also used as an indicator of monthly outwash into Moreton Bay from the Brisbane River watershed. Statistical analysis followed Sokal and Rohlf (1969) Senter (1969) and Finney (1952). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 171. 5.3 R ESULTS 5.3.1 Ova types Four identical types of ova were described for the four species based on size and yolk content. a. Immature ova (type D) - Tiny, angular to ovoid and transparent with a single central distinct nucleus. Ova were yolkless and measured up to 0.16mm in diameter. b. Maturing ova (Type C) - Yolk deposition just began or had already taken place but not complete. Ova centrally opaque, nucleus not visible. The size range was 0.16 - 0.32mm. c. Mature ova (Type B) - Ova densely yolked and opaque. Ova were pale yellowish. d. Ripe ova (Type A) - Fresh ova were transparent and spherical. They floated on seawater. Microscopic examination of preserved ova at 500x magnification showed that - 1. Perivitteline space was small. 2. Yolk was a paste-like semi-fluid with several scattered oil droplets. 3. The egg membrane was sculptured in the form of streaks of short lines in a maze of hexagonal figures. S. filamentosa had the largest ova (Type A) and S. undosquamis the smallest. S. tumbil ova sizes were intermediate between those of S. undosquamis and S. filamentosa. T. myops ripe ova were ciliated and similar to S. undosquamis in diameter (Table 5.1). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 172. Table 5.1 : Ova diameters (in mm) of Saurida and Traohinooephalus Type A Type B Species range mean range mean S. undosquamis 0.62-1.08 0.89 0.36-0.62 0.47 S. tumbil 0.80-1.44 1.20 0.32-0.80 0.57 S. filamentosa 1.08-1.46 1.35 0.54-0.77 0.68 T. myops 0.85-1.23 1.05 0.23-0.69 0.45 5.3.2 Maturity stages The maturity stages were identical in all four species though two genera were involved: Saurida and Traohinooephalus. These stages were based on a) types and quantities of ova present and b) texture and macroscopic appearance, in ovaries and testes respectively. Eight stages of maturity were established for the ovaries and six for the testes. Determination of sex at even 500x magnification was imposs­ ible in some small fish, usually less than 100mm. Their gonads were small and thread-like and were assigned to Stage 1. Immature ova were observed in all stages of ovarian maturity. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 173. 5.3.2 . 1 Stages of maturity of ovaries Stage II: Ovaries long, slender and colourless with tiny blood vessels. Only immature ova were present. Preserved specimens were translucent. Stage Ill: Maturing ovaries. Ovaries were reddish due to ramification of blood capillaries. The main blood vessel was dorsal and extended to about one-third the length of the ovaries from the proximal end. Maturing ova were present. Preserved specimens had tinge of red. Stage IV: Mature ovaries. Ova consisted predominantly of mature types. Ovum angular to ovoid and attached to each other. Pronounced ramification of blood capillaries over ovaries. Ovaries reddish- yellow in fresh specimen but pale in the preserved specimen. Stage V: Ripening ovaries. Ovaries with extensive ramification of blood capillaries. Ripe and mature ova present. Ripe ova were not free from the ovarian tissue and were seen as translucent round spots on a generally orange-yellow gonad. These spots were more or less evenly distributed. Stage VI: Gravid. Ova ripe and transparent but not running. Ovaries were swollen and laid along the entire length of the body cavity. Ova sometimes ran when light pressure was applied to the thoracic region. The ratios of mature to ripe ova were between 3 and 7. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 174. Stage VII: Spawning or running ripe. Ripe and mature ova present. Ripe ova free of ovarian tissue and aggregated inside the ovary or disposed towards the medial and adjacent sections of the two pairs of ovary starting from the proximal end. In the latter, the inner medial sections were more or less trans­ lucent and the outer sections opaque (due to opaque mature ova). Stage VIII: Spent. Ovaries bloodshot, flaccid or shrunken. The mature and ripe ova present were small and not free from ovarian tissue and in a state of resorption. Stage IX: Recovering spent. More or less bloodshot ovaries with immature and maturing ova. Ovaries had several tiny blood vessels. 5.3.2. 2 Stages of maturity of testes Stage II: Immature. Testes grey and longer than broad. Thin blood vessel ramification. Texture more or less ’fibrous' and ’plastic' in preserved specimen. Stage Ill Mature.. IV: Testis was reddish-white. Accumulation of milt slight and proximal and whitened dorsally. Texture 'fibrous' and brittle in preserved specimen. Stage V: Ripening. Testis pinkish distally. Proximal end white and soft as a result of accumulation of milt. This accumulation was usually up to half the length of testis. Drops of milt were exuded when slight pressure was applied to the thoracic University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 175. region. A median dorsal blood vessel gave off branches to each pair of testes. The preserved specimens were paler and of a fluffy texture proximally. Stage VI - Running ripe. V I I : Milt ran freely from fresh testes. Testes wholly white, soft and swollen. Testicular matter more or less fluid. Stage VIII: Spent. Testes bloodshot, more or less empty and flaccid. Stage IX: Recovering spent. Testes reddish with considerable blood vessel ramification. The changes in size and yolk content of ova from the immature to the ripe stage indicated continuous asynchronous maturation. Stage IV contained predominantly mature and some immature ova, a fraction of which ripened through Stages V to VII. Only a few small mature but relatively many immature ova were observed in the spent condition. These suggest - 1) that there is differential or continuous ripening of mature ova. 2) fractional or serial spawning. 3) yolked ova in Stage III and above were all part of the stock released in a spawning season or period. Nematode and cestode parasites were found within and without some mature ovaries. Parasites are discussed in the next chapter. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 176. f csi CO LO CO I csi csi COSi CS] Q) 05 CO c t oCSHI H CO O 05 05 tH rH CO Cu CO CD O COo O CSI 8 05 05 8CO I3 & CO rH CD 05 O CM O Es CO CO 05 00 t> CMCD CSI CS] rH r CHD 00 CD a LO Ph CM CD CO I r CM rH CO LO LO LO 05 05 00 LO 05 05 05 05§ 05 05 0505 05 05 05CO 05 05 CO T LO 05 CM CM CM CM & 05 05 CO CD O O O O O OCO •> 5S rX CS] 05 CSJ CbR o 00 050̂ Co & o csi LO rH CD 00 rH CO tH 0 1—o1 rH 5 CS] CO 0cs0i rHrH CO csi rH o o o o CO a 0L5O CM00 $ 3 CD o o rH oPh 05 w CO - ■O c O) x s p u i p e u o o University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.3B. Monthly mean gonad index of (male) lizardfish. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh C\J C O -Q -z: -o - § Q cn bz j- m 00 x a p u i p e u o o M o n t h and Y e a r University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.4A Frequencies of stages of maturity of ovaries of s. undosquamis obtained between November 1979 and December 1981. (Number of specimens examined per month in brackets.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 100- Dec.'81 (19) 2 3 4 5 6 7 2 3 ,1 5 6 2 8 9 Oct.'81 (156) 2 3 4 S 6 '7 ' 8*91 2 3 4 5r6T7T8 9 2'3 4 I5 ,6,7 ,8 I9' S ta g e of m a t u r i t y P e r c e n t a g e f r e q u e n c y (%) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.4B. Frequencies of stages of maturity of testes of 5. undosquamis obtained between November 1979 and December 1981. (Number of specimens examined per month in brackets.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 100i 100- 100- 50h S e p t'80 NOV. 79 A pr '80 Mar.'81 Aug.'81 6 (115)( ) 50 (177) 50- (106 ) 50- (167) 12 14 16 t— I— I— I--r 40 1-1-1-1-r 13 15 17 100 14 16 12 14 16 12 14 16 100- Dec.'79 Apr '81 50 ( 10 ) May *80 12 14 16 Sw> ev- py tv./ •8 ^1 ■(206 ) (77) Nov.'80 50- ■ ( 64) (62) 12 14 16 0 T-1-1-1-T 12 14 16 100- 14 16 14 16 0 50- 12 14 16Jan/80 . 100 (12 J u n e '80 50- Dec.'80 50 O c t. '81 ) (116) (29) May '81 (280) 12 14 16 (98 )50- 12 14 16 12 14 16 Feb. ’80 1 qq■ 14 16 ( 79) 50- Jan.'81 0 1-1— I— 1— r (60) 100- 14 16 Nov.'81 J u ly ’80 ^ (31) 501 Ju n e ‘81 (187) 0 (116) 00 ̂ ^ — 1— r- "I— I— i— I— r 12 14 16 50- 12 14 16 12 14 16 100- 40 Mar. '80 1— 1— 1— 1— f-12 14 16 F e b .'81 ■I B■ D0(1C3l (187) 100- (79 ) 100i 14 16 0 50 — —1 r50- 12 14 16 Aug. '80 50 J u ly '81 (17) 50- "I I 1— 1— r (176) 12 14 16 ‘141 'l6 1— |— |— |— r 12 14 16 . 1— 1— 1—1— r 12 14 16 Stage of m a t u r i t y P e r c e n t a g e f r e q u e n c y ( University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.4C Frequencies of stages of maturity of ovaries of s. tumbil obtained between November 1979 and October 1981. (Number of specimens examined per month in brackets.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Nov.'79 May 80 ^q_ Dec.'80 J u ly '81 5CH (16) (2 8 ) 0 l— I— I— I— r 2 3 4 5 6 7 8 9 2 ^ 4 5 6 7 8 9 2 34 5 6 Z 8 9 2 3 4 5 6 7 8 9 50 H D ec.'79 | 50- (17) " 50H ■ A u9- 81 ( 23 ) 2 3 4 5 6 7 89 0N? 100- 2 3 4 5 6 7 8 9 2 34 5 6 7 8 9 ̂ J u ly '80 ^ Feb.'81 B J m ■ (23 ) ~ (20) H I 501 S ept. 81 (19 ) 50- R s ' e V s ? 1 , , I m * I n 50-j 23 4 5 6 i 8 9 0|l 2 3 4 5 6 7 8 9 1 1 r 1 1 r i l l - 2 3 4 5 6 7 8 9 - Oct. '81 (9 ) Feb. *80 ° (18) j . . m . m £ 2 3 4 5 6 7 8 9 M a r. '80 (16) 2 3 4 5 6 7 89 2 34 5 6 7 8 9 2 '3 '4 '5 '6 V 8 9 50- Ju n e 81 (21) I— I— i— '— r — ^Xdl— ̂i— r- 2 3 4 5 6 7 8 9 2 3 4 5 6 7 8 9 2 3 4 5' 6' 7' 8 9 Stage of m a t u r i t y P e r c e n t a g e f r e q u e n c y ( University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig, 5.4D Frequencies of stages of maturity of testes of s. tumbil obtained between November 1979 and October 1981. (Number of specimens examined per month in brackets.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Oct. 81 (39) 12 14 16 12 14 16 S tage of m a tu r i t y p e r c e n t a g e f r e q u e n c y (%) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig, 5.4E Frequencies of stages of maturity of ovaries of s. filamentosa obtained between May 1980 and February 1982. (Number of specimens examined per month in brackets; nd = no data.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Aug. 81 (n d ) Sept. 81 (6) — ----1 1 2 3 4 5 6 7 8 9 2 3 4 5 6 7 8 9 2 3 4 5 6 7 8 9 J u ly 80 (nd ) Oct. '81 ( nd ) Mar. 81 o 50- Nov. '81 ( nd ) (117) c (D =3 Dec. *81 (n d ) C T CD 2 3 4 5 6 7 8 9 2 3 4 5 6 7 8 9 50 Jan. '82(nd ) CD cn S ep t.'80 CQ -1—1 (17) c s C-J 1— i— i— i— r~ 100' n— I— — I— I— r c 12 14 16 Mar. '81 12 14 16 CD o J u ly '80 ( n d ) cr (142) Aug. '81 ( nd ) CD Aug. ‘80 ( n d ) 50~ 100H S ept. '80 50- Sept. (3 ) ( 2 ) CD cn CD 50- 12 14 16 -i—' c I CD O A p r '81 (_ Oct. —Si—1('— nt_d ) CD 50- (4 ) CL 0 12 14 16 12 W W Nov. '81 ( nd ) Oct- '80 Dec. '81 ( nd)40- ( 5 ) o' L 12 14 16 50- Feb. '82 0 i t 12 14 16 50-j — ■ M■a->y -'8■1 ( 4 ) 50-«Nov '80 - I (16) ■ 1 . . 1 I I r12 14 16" r i i i 12 14 16 i d. 14 16 Stage of m a t u r i t y University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.4G. Frequencies of stages of maturity of ovaries of t . myops obtained between June 1980 and February 1982. (Number of specimens examined per month in brackets; nd = no data.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 100H May '81 ( nd) June 80 — Dec.'80 (1) (202) June &1(n d) 50- 2 3 4 5 6 7 8 9 - 2 3,4 ,5 '6 ,7 '8 I9' 1 j an .'81 J u ly 81 50- ( 2 ) July 80 50 (3) i— i— i— i— i— i— 1 i , . 2 34 5 6 7 8 9 2 3 4 5 6 7 8 9 Aug. '81 ( n d ) 100 Sept. *81 ( 2 ) . i— i— i— i— i— i— r 2 3 4 5 6 7 89 t --i f-i . ■ i i-T 2 34 5 6 7 89 2 3 4 5 6 JZ. 8 9 D e c . B l ] = ( n d) Apr '81 Feb-'82 J‘ a ni.B82|) ( 10) (43) p _ _ 2 3 4 5 6 7 8 9 0 i □ . ii 2 3 4 5 6 7 8 9 i 2 U3 4i . 5' 6 7 8 9 S t age of m a t u r i t y University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.4H. Frequencies of stages of maturity of testes of t. myops obtained between January 1980 and February 1982, (Number of specimens examined per month in brackets; nd = no data.) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 100- 1001 100- Nov. '81 Jan. '80 Oct. 80 (2 3 ) (7 ) 50- ( 1 ) i i i i i r i— i— i— r 12 14 16 12' 14 16 50- Nov.'80 Feb.-Mar 1980 ■ ( 110) ( nd) o r- 12 14 16 100 40- Dec- 80 12 14 16 J u ly '80 | L ( 1(61677)) M ay-June 1981 “ ( 1 ) — ^ — I— r- ( n d ) ^ 12 14 16 100H Jan. 81 J u l y ‘81 q" (96) (1) 12 14 16 14 16 A ug .- Sept. 81 ( nd ) ^ 14 16 '' 50- m Oct. '81 ( 79 ) Stage of m a t u r i t y Per cent age fre q u e n c University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 180. 5 . 3 . 4 . 2 S. fi lamentosa Two years (1980-82) data were combined in the estimation of the spawning season based on the occurrence of mature, ripe and running fish. G.I. was not used because of many missing monthly data (Figs. 5.3 A,B; Appendix 6). However an overview of pooled results (Figs. 5.4 E,F) about the occurrences of ripe and spawning and running females indicate a prolonged spawning season from January to October. 5. 3 .4 . 3 T. myops Mature fish of this species were observed between October 1980 and February 1982. Spent and recovering stages were obtained in November 1981 (Figs. 5.4 G,H). These are in agreement with the trend in G.I. between this period (pooled data; Figs. 5.3 A,B; Appendix 6). Very small sized fish made 8.1% of the December 1980 sample (Appendix 7). Thus the trend in mean, maximum G.I. and occurrences of ripe and spawning fish indicate an asynchronous maturation of ovaries and a spawning season from October to April. 5.3.5 Fecundity The absolute fecundity was estimated using Stage IV; the mature stage. Partial fecundity was estimated using Stage VI; the ripe ovary. Plots of partial fecundity and length, and fecundity and weight indicated curvilinear relationships governed by the equation F = ax^ where F is fecundity, x = length/ weight, a, a constant and b̂ = an exponent. Each relationship University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 181. was analysed by plotting its logarithmic transformation : log F = log a + b log x (Table 5.5A). Table*5.5A : Constants in Fecundity equation a Species SL WT ST WT S .undosquamis 2168.4 3107.7 0.3152 0.3142 S. tumbil 0.301 451.4 2.0082 0.7151 T. myops 470.4 641.4 0.4894 0.4886 Fecundity increased with increase in weight or length (Figs. 5.5 A-C). The estimated number of spawns per spawning season was calculated as the ratio of the absolute to partial fecundity. The absolute fecundity range calculated for S. filamentosa was 401,855 (345mm) - 1,288,576 (575mm), whereas the partial fecundity range was 30796 (168mm) - 54823 (322mm). The other three species also have high absolute and relative fecundities (Table 5.5B). Considering only specimens from the bay, S. undosquamis was found to be more fecund and also more abundant (Appendix 7) than S. tumbil. * SL = Standard length and WT = whole body weight University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.5A, Scatter diagram showing the relationship between fecundity and standard length (•)/weight ( + ) Of s. undosquamis. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh (C0Lx ) ̂ ipuroaj iei }jed University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.5B, Scatter diagram showing the relationship between fecundity and standard length (•)/weight ( + ) of s. tumbii. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh cn + -C cn Ĉ Ol*) ^ j p u m a j ' lei j jed University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 5.5C. Scatter diagram showing the relationship between fecundity and standard length (•)/Weight (+) Of T. myops. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh (c0 LX) A ^ i p u n o a j i e i ^ j e d University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 182. CO lO -P- •H CD CD U T3 Vl Cl) 0 O a +-> u Gw o W CM 00 LO CO CO CD LO CO<1> s •P g ctf Cl)W (/) w 5T CD I -P rH *H lO 00 C<$ XJ G> 00SI O COiH CD 00 s I G> t> CO3 CM T—I COa. 4p->*"> O CD t> in CM O) 1—I CS] I o o OSi CO £>> T'-i I I c&'g-S CO CO Table 5.5B FECUNDITIES (N ♦Mean Specific Partial fecundity = Mean Partial Fecundity Standard length (SL) or whole body weight (WT) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 183. lO O 00 o t> oo CM CO CO CM oo in CM o X CM lO O in m m Q) CM CD CO rH rH t> in rH CM CM CO o t> o m CD CO in CM CM CM CM oo CO o I c*- m CM CM s CO * +-> X CD CD CD 00 CO CO O c$ P£ o CM w “ AQ* X p 0 CD w CO LO m in m o ^ SH o ’ in O 00* CM rH ctf CM O CD rH CM CD X 00 & CO in m w -+-> rH CM W XJ 0 cCor dU 0 Eh O 3 3 CM CM CD CD o< CM & COCO 8 in CO Co CM 1 ■£1H ^ ^ Co O i l l o o 00 00 C f l ? > ■s o I o rH U oo p _ 00 O 00 O Cfl CD CD CD - 00 ► 00 rH t> ft ĉ ■S". s : £2 ^ o ^ p ti £ • c§* a a; $ 8 T a b l e University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 184. 5.3.6 Sex Ratio Sex reversal was not observed in any of the four species studied. 5 . 3.6 .1 S. undosquamis The male to female monthly ratio varied considerably (Appendix 8). In the general population, chi squared tests indicated significant differences (p > 0.05) from the expected Mendelian ratio of 1:1 in many months. Sex ratio were then calculated by seasons and year to determine possible variation related to migration, natural mortalities or different distribution of sexes during the main spawning and nonspawning seasons. There was little evidence for any of the above. In the spawning season (September 1980 - January 1981), mature males outnumbered mature females at a ratio of 1.58 : 1.00. A significant trend was evident when sex ratios were calculated at 20mm intervals (Table 5.7). Males dominated intervals between 120 - 159mm and were poorly represented in samples greater than 200mm. Females dominated intervals between 160 - 300mm; an indication of sexual dimorphism in sizes. The largest male and female were 300 and 320mm respectively. 5.3.6.2 S . tumbil In the general population, the sex ratios between April to August showed varying trends (Table 5.6). Chi squared tests demonstrated significant distribution of size by sex when sex ratios were calculated at 20mm size class intervals (Table 5.7). Males were essentially absent in samples greater than 260mm in length. Females were more abundant in all class sizes greater than 200mm; the males dominated the University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 185 csi CO TO T—1 tH CO o CQ o o o o o <0 o tH m CM o o o o rH o i csi .°® 0C3r w 1 o T—1 t> T—1 rH o o rH00 o o O1 CO CJ XCSO •rH Ph o CO SS o o o o O SEX RATIO DURING SPAWNING SEASON Fish numbers Chi- Chi squared squared observed observed observed observed Class value valuefemale male expected female male expected sizes (ran) 60 - 79 8 0 - 9 9 100 - 119 14 14 14 2.5 120 - 139 124 89 27.5281 1.0000 140 - 159 171 400 285.5 91.8406 10 18 14 2.2857 160 - 179 161 124.0 22.0806 11 134 72.5 104.3379 180 - 199 106 24 65.0 51.7230 16 139 77.5 96.6064 200 - 219 38 19.5 35.1026 59 15 26.1622 220 - 239 6.40 60 33.5 41.9254 240 - 259 85 43.5 79.1839 260 - 279 68 280 - 299 26 300 - 319 15 320 - 339 340 - 359 0008*1 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 186. 00 $ s CO o o o o T—1 co CM rH a, TC—Do 1 T—1 3 CO 00 00 CD CM « CO o £ •< LO TC—D1 CD t-H CM CM CD SEX RATIO DURING SPAWNING SEASON Month Fish numbers Fish numbers and Year observed observed expected Chi- observed observed expected Chi- female male squared female male squared value value October 1980 18 12.5 4.840 November 1980 110 97 3.485 December 1980 202 182 January 1981 53 73 63 3.175 49 72.5 15.234 February 1981 18 11.5 7.347 March 1981 117 142 129.5 2.413 ------ April 1981 0.667 10 11 10.5 May 1981 42.5 33.047 June 1981 10 July 1981 12 6.5 9.308 September 1981 2.000 Spawning season 287 249 2.694 368 385 376.5 000’I 69 96 96£* ̂ University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 187. range 160-199rmn. Mature females outnumbered mature males at a ratio of 1.10:1.00. The largest male observed was 246mm. The evidence suggests sexual dimorphism in sizes. 5.3.6.3 S. filamentosa and T . myops These two species could not be analysed for yearly or seasonal differences in sex ratios because of the absence of many monthly data. Therefore these findings should be considered preliminary. Chi squared tests of observed monthly sex ratio in the spawning season showed significant differences (p > 0.05) (Table 5.8). The observed sex ratios of the spawning season as a whole did not however show any signif­ icant differences in the two species. In the mature population, males were found to be significantly more abundant in sizes equal to or less than 179mm and 239mm in T. myops (Table 5.2) and S. filamentosa (Table 5.3) respectively. The females dominated sizes equal to or less than 180mm and 240mm, also respectively. This is evidence for sexual dimorphism in the two species. The second and third dorsal rays of S. filamentosa are usually elongated and filamentous (Ogilby, 1910; Norman, 1935; Shindo and Yamada, 1972; and personal observation). The percentage ratios of the length of second and third dorsal rays to the standard length were calculated to determine possible variation related to standard length or sex (Table 5.9). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 188. S. tumbil and T, myops from September to April and probably beyond. Nutrients are also continually released and replen­ ished by wave disturbance of the sediments by strong winds accompanying the period. Thus, though the onset of spawning coincides with the rise in rainfall (Fig. 5.7), the slow development of the ova guarantees the prior availability of plankton before the emergence of larvae. The evidence for the importance of food in spawning habits of T. myops and S. filamentosa within and without Moreton Bay is less convincing because of inadequate data. Very few small and immature T. myops and no S. filamentosa were collected in Moreton Bay. They were obtained off-shore. Further work off-shore is required to determine stimuli to spawning and other factors important to survival of their larvae. Availability and abundance of food might trigger maturation and spawning. For it is probable that the larvae University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Month Fig. 5.7 Mean monthly rainfall (+), mean monthly minimum (•) and maximum (o) temperatures recorded at Cape Moreton Lighthouse weather station by the Bureau of Meteoro­ logy between 1958 and 1981 T e m p e r a t u r e (°C ) Mean m o n t h l y r a i n f a l l ( m m ) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 194 move down the water column to take advantage of relatively great plankton production within the discontinuity layer in summer. In winter, convection and wind mixing extend deep into the water column, thus bringing up nutrients to the surface which would guarantee abundance of plankton to their larvae. The works of Gibbs (1959) and Tabery. Ricciardi and Chambers (1978) give some support to the above hypothesis. Gibbs (1959) collected many Atlantic Synodontidae larvae including T. myops from shallow water in every month of the year in the Western Gulf of Mexico. Some collections, however, were made at considerable distance from any shallow water. He noted that this was especially true of T. myops which was taken on occasion, in or near the Gulf Stream, over more than 2,000 metres! Larvae of Synodus were collected in the estuary of the Hudson River (New York State, U.S.A.) by T abery et al (1978). The western Gulf of Mexico and the coast of New York State are areas of mixing in which productivity of marine life would be expected to be very high or moderately high (Isaacs, 1969). The Sargosso is a semi-tropical marine desert area where even horizontal mixing by surface currents is slight. Here the production rate is very poor, except during the brief winter period when some convectional mixing may take place (Tait, 1981). Thus in the Gulf Steam, T. myops larvae must descend to great depths for their food supplies. The works of several others too indicate a correlation between the onset of spawning, spawning and the occurrence of larvae on the one hand, and the food supplies on the other. Budnichenko and Dimitrova (1979) noted that the spawning in S. undosquamis and S. tumbil in the Arabian Sea reach a maximum in the winter monsoon. Tait (1981) reported that in the north Indian Ocean, upwelling occurred in the northern- University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 195. most part of the Arabian Sea during the winter north-east monsoon, and off the Arabian and Somali coast in summer when the monsoon current developed. This is substantiated materially by Fursa (1969, 1976)* who provided data showing that larger numbers of lizardfish hatchlings occurred in winter than in summer in waters of Western Hindustan (Eastern Arabian Sea). Saurida species spawn between June (onset of spring) and September (onset of autumn) in the Japanese (Tatara, 1965), East China (Yamada, 1968a) and Mediterranean (Bograd- Zismann, 1961-62**; Chervinsky, 1959**) Seas. This period is characterized by favourable combination of temperature, light, nutrient supply (from mixing (Isaacs, 1969)) and stable water column and a subsequent enormous increase in the quantity of plankton (Tait, 1981). In the tropics, Sinoda and Intong (1978), after analysing size frequency distribution of the S. undosquamis from the east coast of the inner Gulf of Thailand, concluded that S. undosquamis spawns all through the year. And in the off-shore waters of the Madras Coast; Bay of Bengal, where there is little or no seasonal variation in sea water temperature (Varadarajan and Subramoniam, 1982), S. tumbil spawns from November to February (Annigeri, 1963); a period of high nutrient content due to upwelling (Muthu, 1956)***. From the foregoing, the Saurida in the Arabian Sea, the Bay of Bengal and Gulf of Thailand, all in the tropics, have various spawning times and durations. Thus, the general conclusion is that whereas in the temperate areas temperature and food availability determines onset and duration of spawning, in the tropics it appears that only the availability of food is critical. * Quoted in Budnichenko and Dimitrova, 1974. ** Quoted in Ben-Yami and Glaser, 1974. *** Quoted in Varadarajan and Subramoniam, 1982. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 196. Intra- and inter-species spawning periods were also reported from the East China Sea. Tatara (1965) found different spawning times in two sub-populations of S. undosquamis: May to August for the eastern sub-population and June to September for the western sub-population. The period of S. tumbil , April to June, overlapped the above two. Here in Moreton Bay the peaks and durations of S. tumbil and S. undosquamis though overlapping, were quite different. S. filamentosa spawned all year in the open sea. Fecundity of lizardfish is quite high, and in a curvi­ linear relationship with either standard length or weight (Figs. 5.5A - 5.5C). Table 5.10 lists relative fecundities. Table 5.10 Relative Fecundity (R.F.) and Number of Spawnings of Lizardfish Number of Species per gram per mm spawns per total weight standard length season S. undosquamis 906 340 2 - 8 S. tumbil 387 296 3 - 6 T. myops 569 420 3 - 5 The high ova production to body weight is maintained by serial spawning as a result of asynchronous maturation. The number of spawnings per season was approximated as the multiple factor of mean partial fecundity in the mean absolute fecundity per size class (Table 5.5B). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 197. Similar results have been found in the Arabian Sea (Budnichenko and Dimitrova, 1979) and in the East China Sea (Yamada, 1968a). However, Latif and Shenouda (1973) reported a straight line relationship between fecundity and any of the following variables: length, weight and age. And also a straight line relationship between length and R.F (S.L) but found R.F (Wt) to be constant. S. tumbil and S. undosquamis spawn in Moreton Bay and circumstantial evidence suggests that the latter probably also occurs and spawns off-shore: One spent male* and one mature female* were collected off-shore from 80 - 160m. Similar results found elsewhere support the above hypothesis. In the Gulf of Thailand, 5. undosquamis greater than 210mm migrate to deeper waters (Sinoda and Intong, 1978). Ben-Yami and Glaser (1974) reported that lizardfish spread over deeper trawling grounds during the warm season in the Levant Basin of the Red Sea. From the Sea of Japan, Tatara (1965) reports that "Distribution of S. undosquamis changed little by little as its age advances, it occurred mainly in the Channel waters in larval and fry stages, and then covered all the Inland Sea in immature stage, range of migration became wider in adult stage. Its distribution in winter was limited to the Channel waters, but its range expanded both in spawning and feeding seasons.” Budnichenko and Dimitrova (1969) found S. undosquamis and S. tumbil in depths down to 500m and 300m respectively in the Arabian Sea. The monthly seasonal and yearly variation in sex ratios of S. undosquamis and S. tumbil suggests differential natural mortalities of sexes assuming equal fishing mortalities. Latif and Shenouda (1973) found an inverse relationship between percentages of males and females in S. undosquamis in the Gulf of Suez in 19 months sampling. Tatara (1965) * The largest obtained during the study. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 198. reported differences between sexes in natural mortalities in two sub-populations in the Sea of Japan - with the males having high natural mortality (Table 5.11). Table 5.11* Annual mortalities in S. undosquamis (M = male, F = female) Sub-population Sex Total Fishing NaturalMortality Mortality Mortality M 0.96 0.87 Eastern 0.69 F 0.87 0.59 M 0.88 0.87 Western 0.45 F 0.72 0.59 The larger class sizes either consisted exclusively of females or were dominated by them. This is the result of differential growth rate between the sexes and greater maximum age of females (Tatara, 1965; Budnichenko and Nor, 1978; Sainsbury and Whitelaw, 1981). Maximal increase is observed in 1+ group; in succeeding year groupings the rate of increase decreases (Budnichenko and Nor, 1978). These differences in growth rate and natural mortality are reflected in the calculated asymptotic lengths of 571mm for females and 466mm for males (Sainsbury and Whitelaw, 1981). The dominance of males in the small size groups (Table 2 and 3) and the differences in S.L^q (Tables 5.4) for males and females appear to maintain a 1:1 sex ratio in the spawning population (Tables 5.6 to 5.8). In S. undosquamis and S. tumbil the sex ratios for mature males and mature females were different, 1.58:1.00 and 1.00:1.10 respectively. Thus, S. undosquamis has a dominance * Adopted from Tatara, 1965. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 199. of males and S. tumbil , females. Yamada (1968a) observed that S. tumbil females were always more numerous and larger than males; fish equal or greater than 400mm were almost all female. On the other hand, Budnichenko and Nor (1978) reported that in spawning populations of younger fish, males outnumbered the females but in older age groups, females outnumbered the males in the Arabian Sea. Thus, it appears both length at maturity and differential sex ratios by length combine to produce equal male and female reproduct­ ive capacities in the spawning stock. S. undosquamis outnumbered S. tumbil in Moreton Bay by a ratio of about 4:1. This is probably due to the relatively high fecundity of S. undosquamis per size (Table 5.5). In the Arabian Sea, the comparative numbers of S. undosquamis were greater than for S. tumbil, on the average 1-2 times as many (Budnichenko and Dimitrova, 1979). Here too Budnichenko and Dimitrova (1979) related the difference to the greater number of ova produced by the former on account of its greater population. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 200 . 6. THE PARASITES OF FOUR LOCAL LIZARDFISH INCLUDING AN ANALYSIS OF THE FREQUENCY DISTRIBUTION OF THE CESTODE C. GRACILIS (LARVAE) (WITH REFERENCE TO THE NEGATIVE BINOMIAL DISTRIBUTION)_____________ 6.1 INTRODUCTION Many organisms are parasitic on fishes. In the marine environment the common parasites are the helminths - trematodes, cestodes, nematodes and acanthocephalans (Lager, Bardach and Miller, 1962; Sindermann, 1966). Usually it is as larvae that the helminths are of greatest signif­ icance. Adults occur in the digestive tract and the larvae are usually found in the flesh or in the viscera (Sindermann, 1966). These parasites, as any others, may act directly or indirectly either to increase host susceptibility to predation or to diseases or to reduce competitive fitness (Anderson, 1982). Parasitism is an ecological relationship that can be expressed quantitatively in terms of populations. One of the few methods of expressing this quantitative relation is the frequency distribution of the parasite in its host (Crofton, 1971). Individuals of any one kind (or species) are seldom, if ever, randomly disposed in space. Most commonly the distribution is overdispersed (i.e. variance> mean), implying a partially determinate pattern of distrib­ ution and thus a reservoir of information about the behaviour of the organisms concerned which could then be described in terms of empirical or fundamental mathematical models (Cassie, 1962). Cassie (op cit) states that "Empirically, it is desirable to condense the sample data, so that any given population may be described by a few parameters, which are readily comparable with the corresponding parameters of another population.... In such an application, the theory behind the model is relatively unimportant, provided it 'fits' the data in hand. The fundamental model, University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 201. on the other hand, is based on some hypothesis of real biological significance. If it fits the data better than other possible models, it provides some justification for the hypothesis concerned.” Cassie (1962) however cautioned that the same distribution may often be generated by different, and even contradictory, set of postulates. Crofton (1971) pioneered and also justified the use of the negative binomial from amongst other similar models to describe the frequency distribution of parasites among hosts. Crofton (op cit) states that the negative binomial distribution can arise as a result of: 1) A series of exposures to infection in which each exposure is random but the chances of infection differs at each exposure or wave of infection. 2) The infective stage not being randomly distributed. 3) Infection increasing the chances of further infections occurring. 4) Infection decreasing the chances of further infection. 5) The variation in host individuals which makes the chances of infection unequal. 6) The chances of infection of individual hosts changing with time. The negative binomial distribution is defined in terms of two parameters: fk' and 1 p'. The mean and variance of the negative binomial distribution can be calculated as follows: mean; X = ^ (.... equation 6.1) variance; V = ^ (.... equation 6.2) P and P = yX (.... equation 6.3) The parameter ’ p ’ must lie between zero and one, and 'k’ may assume any positive value. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 202. The negative binomial distribution is given by the expression (q - p) _ and the number of parasites can take the form pj = (k + j ~ 1} pkq5 (J = °’1 ’2 ’3’4 -• - > Because ( k-iq ) = and pj+1 = ( (k+j) q/(j +1) ) pj (.... equation 6.5) The fitted frequencies (Fj) may be calculated in the following manner: F o = N opkq° and F j. =■ FF,( j.-D where Fq and F^ are frequencies of zero and j parasites (See Pollard, 1977 pp. 114-117) The quantitative relationship between parasite and host is investigated by the application of the non-truncated and the truncated forms of the negative binomial distribution (Crofton, 1971). A good fit of observed data to the non­ truncated form suggests the parasites probably had little effect on host mortality. The truncated form is dependent on the assumption that the parasite can kill their hosts and that the lethal effect is a function of the number of parasites in an individual host. A good fit to the truncated form therefore lends support to the latter hypo­ thesis, in which case the observed data shows an abrupt fall in frequencies at higher infection levels when compared to the theoretical distribution (Crofton, 1971). F, p and k are estimated by successive iteration, usually by computer. In finding the truncation point the usual procedure is to start with the low classes (e.g. 0,1,2, 3 ..,) and determine values of F, p and k. The latter is referred to as truncation at 3. F, p and k can then be determined for 0-4 (truncation at 4), 0-5 and so on. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 203. When the operation has proceeded until the last recorded frequency in the sample has been included the values for F, p and k should be the same as or identical to those obtained by fitting a negative binomial distribution to the whole of the observed data. If no truncation has occurred then the successive values of F, p and k should be constant. There will always be variation because the sample is finite and usually small, but in practice the estimates of F appear to be the best point of reference for quick inspection. If truncation has occurred this can be observed by reference to changes in F as well as in p and k. A precise truncation point is not easy to determine because tests for goodness of fit after iteration present considerable difficulties. In most cases, however, a specific level of truncation is rarely necessary because the values of F, p and k can be expressed in terms of narrow ranges (Crofton, 1971). Pennycuick (1971d) satisfactorily fitted the non­ truncated negative binomial to the frequency distribution of parasites in a population of three-spined sticklebacks, Gasterosteus aouleatus L. On the other hand Lester (1977) found that the best fit of the negative binomial to the trematode frequency distribution in a population of yellow perch, 'Perea flavescens was the truncated form - indicating the ’loss' of heavily infected fish from the sample. Lester (op oit) ascribed the ’loss’ to higher mortality in the heavily infected fish. The parasitic fauna of lizardfish are diverse but in the main the literature on these parasites are taxonomic descript­ ions or records. Little is known about the effect of these parasites on lizardfish populations. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 204. The parasites of lizardfish include several species of protozoa (Narasimhamurti and Kalavati, 1972; Jensen, Moser and Heckmann, 1979), monogenean (Unnithan, 1972) and digenean trematodes, cestodes, acanthocephalans, nematodes (Overstreet, 1968; Rees, 1969; Parukhin, 1974; Jensen and Heckmann, 1977; Jensen, Moser and Heckmann, 1979), and ecto-parasitic crustaceans (Avdeev. 1977; Cressey and Cressey, 1979). Parukhin (1974) recorded 27 species of helminths in populations of five species of lizardfish in the Indian Ocean. Cressey and Cressey (1979) found 18 species of copepod parasites on 14 Indo-West Pacific lizardfish species. The digestive tract and the gills of lizardfish harbour several species of trematodes. Unnithan (1972) reported a monogenetic trematode, Sauricotyle sprostoni on the gills of S. tumbil from the Arabian Sea. Species of Lecithoohirium, Sterrhurus and several other digenean trematode genera have been reported to occur in the stomachs of Synodus foetens, S. lucioceps, S. variegatus, Saurida undosquamis, S. gracilis, S. tumbil and Trachinocephalus myops (Overstreet, 1968; Parukhin, 1974; Jensen, Moser and Heckmann, 1979). Several species of cestodes have also been found in lizardfish. Overstreet (1968) reported the occurrence of tetraphyllidean larvae in the body cavity and viscera of Synodus foetens. The adult pseudophyllidean, Anantrum tortum was found in the intestines of S. foetens (Overstreet, 1968) and S. intermedius (Rees, 1969). S. luoiooeps plays host to adult Anantrum histocephalum, larval tetraphyllideans; Scolex pleuronectis, Phyllobothrium sp. and larval trypanorhynch; Grillotia smarisgora, Lacistorhynchus tenuis and Callitetrarhynchus gracilis (Jensen, Moser and Heckmann, 1979). Trypanorhyncha larvae also occur in Saurida undosquamis, S. gracilis, S. tumbil, Synodus variegatus and Trachino­ oephalus myops (Parukhin, 1974). Larval stages of nematodes (e.g. Anisakis sp) have been found in Synodus foetens (Overstreet, 1968), S. lucioceps (Jensen, Moser and Heckmann, 1979), S. variegatus, Saurida undosquamis, S. gracilis and S. tumbil (Parukhin, 1974). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 205. The only quantification of a relationship between lizardfish and a parasite have been done on Synodus foetens - parasitic fauna from South Florida by Overstreet (1968). However this was limited to estimations of incidence and intensity of infection and associations among parasites. In this study, many members of the parasitic fauna of four local species were identified. The effect of the most common parasite in Moreton Bay lizardfish, C. gracilis (Rudolphi) 1819 on lizardfish population numbers was investigated by fitting the negative binomial to parasite- frequency distributions. 6.2. METHODS AND MATERIALS The parasitic fauna of Saurida tumbil, S. undosquamis, S. filamentosa and T. myops were studied between November 1979 and December 1981. The process of preservation, measure­ ment of length and the determination of sex of lizardfish followed the same procedures as described previously in the main introduction unless otherwise stated. The presence of parasites was determined by direct examination of each fish. The general condition of each fish was noted and the external surface of the body and gills were carefully examined for parasites. Each fish was measured and sexed. The fish was cut open and the body and pericardial cavities and viscera were searched for parasites. The gut was slit along its length and also thoroughly searched for parasites under a binocular micro­ scope - the search of the stomach and gonads were done concurrently with food item and gonad studies. In each case the parasites were removed and identified to the lowest systematic level possible. The position of each parasite in the host was also noted. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 206. The number of species and individuals of lizardfish, and the initial fixation of many of the parasites by formalin (used to preserve lizardfish), precluded specific identification of many parasites. However some samples of lizardfish were periodically frozen in the field and identifications of the 'fresh' parasites made. 'Fresh' helminths (adults) were fixed in Berlund's fixative for specific identification. With regards to cestode larvae, the capsules of the cysts were broken and larvae left in distilled water for about 15+ minutes for the evagination of the hooked tentacles; important taxonomic features. These larvae were then fixed in 10% formalin for about two weeks and then transferred to 70% alcohol. Photo­ graphs were taken of the cestode larvae, some in situ (i.e. by the aorta). Dr. Lester identified the cestodes and nematodes, whereas Dr. D.I. Gibson Museum of Natural History identified trematodes taken from the stomachs of S. undosquamis. Dr. Neil Bruce identified the crustacean parasites. Initially only specific identifications of the parasites were envisaged. However, on account of the frequency of occurrence of the larvae of the cestode C. gracilis in 5. tumbil and S. undosquamis during the preliminary survey, notes of the larvae distribution and location were incorporated into the main survey. Negative binomials were fitted to the frequencies of the larvae after having obtained k, p and q from equations 6.1 to 6.3. k was varied until the least variation between expected and observed distribution was obtained. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 207. 6.3 RESULTS 6.3.1 Parasite Fauna Parasitological findings are summarized in Table 6.1. The most common parasites were trematodes and cestode larvae. Table 6.1 : Parasites found in Lizardfish Parasite Site of Infection TREMATODES Lizardfish S. undo­ S. fila­Species S. tumbil squamis mentosa T. myops Lecithochirium magnus Yamaguti 1938 ST? ST Other digeneans (including Hemiuridae) ST ST ST ST CESTODES Trypanorhych Larval Callitetrarhychus gracilis A0,M,BC,L AO,M,BC, (Rudolphi, 1819) A,MU,0E, L, A,MU, PC 0E,PC Larval Nybelinia BC BC Pseudophyllidea Adult Onchodiscus sauridae I I? ACANTHOCEPHALANS M M NEMATODA BC,ST,0 BC, ST ,0 BC,ST,0 BC,ST,0 Larval TAnisakis Type 2 T M ECTGPARASITIC CRUSTACEANS (ISQPCDA) G G G G Cymoithoidae Nerocila saurida Avdeev 1977 SR Anilocra cavicauda Richardson 1910 SR Gnathidea Gnathia, Larval Praniza G L, liver SR, scapula region AO,by aorta BC, body cavity 0, ovary ST, stcmach G, gills QE, stuck in oesophageal M, mesentary or stcmach wall MU, muscle I, intestines PC, by pyloric caeca University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 208. 6.3.1.1 Trematodes All trematodes were obtained from the stomach. In a few instances up to 100+ trematodes were counted in indiv­ idual stomachs. Specifically 67 trematodes were counted in a specimen of T. myops in December 1980. On the other hand as many as 85, 123 and 146 were counted from three S. undosquamis specimens in December 1979 - these individuals harboured no other parasites. Table 6.2 shows the percent­ age incidence of trematodes in various samples. Table 6.2: Percentage incidence of trematodes in lizardfish samples. Species Percentage (%) Incidence Sample Number S. undosquamis 2.1 5411 S. tumbil 3.7 1343 S. filamentosa 19.1 612 T. myops 43.6 1343 The trematodes obtained from S. filamentosa were different from those obtained from shallow-water lizardfish - these digenean trematodes were considerably bigger; about 2-4mm in length and about 1.5mm wide. 6.3.1.2 Cestodes Larval trypanorhynch, C. gracilis were the most prevalent of the parasites in S. tumbil and S. undosquamis (also see Table 6.1) University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 209. The percentage incidence of C. gracilis larvae in the two populations were 53.4 and 9.9 in S. twnbil and S. mdosquamis respectively; many of the larvae occurring by the aorta (Table 6.3). Table 6.3: Incidence of C. gracilis in S. tumbil and S. undosquamis Percent inci dence in Location S. twnbil S. undosquamis Female Male Female Male Whole fish 55.9 50.9 11.7 8.4 By aorta* 84.6 89.8 77.0 72.4 The trend in monthly percentage incidence indicates high occurrences of the larvae in the S. undosquamis population in Moreton Bay between May and September (Fig. 6.IB). The trend is however less discernible in S. tumbil population (Fig. 6.1A). Intensity of infestation was different in the sexes (Tables 6.4A,B; 6.6). The highest infestations per individ­ ual were found in the females. There was no correlation between number of larvae and length of fish in either of the two lizardfish species: Species Correlation Coefficient S. tumbil Female -0.05634 Male -0.06777 S. undosquamis Female -0.004382 Male -0.003411 * See Plates 39 A,B,C. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh C, gracilis (in situ) by QOftQ in S. tumbil Plates 39A and B University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Plate 39C: c. gracilis (in situ) by aorta in 5. t w ib i i; expanded view University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh h O -cn c 2 oco Q^ &r- ( % ) a e A j e i a o u a p i o u i a 6 e ^ u 3 D j 9 d Month and Y e a r University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Fig. 6,IB. Incidence of c. gracilis larvae in POPUlatiOn Of s. undosquamis in Moreton Bay University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh Mont h and Y e a r University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 210. CM rH CM rH o CO CM 00 CD CM rH CM CM r0H0 LO rH LO CO O CM 00 CM I> T—1 CM rH CD rH O CM LO rH CM o rH O rH CM CM CO LO o CM CO CO CM rH 00 rH CM o 00 CM LO CD CM rH rH CO LO O a> rH O CO CM r0H0 t> a CD rH 1 L/ s o c l / 1 ZoJ?/ ~/ a sCO/ co 'o Table 6.4A: Frequency distribution of C. gracilis with size and sex of lizardfish SEX FEMALE SPECIES S. tumbil 100-119 120-139 140-159 14 23 17 160-179 100 46 12 180-199 10 85 50 21 11 200-219 29 11 11 220-239 49 13 240-259 39 21 12 260-279 23 280-299 12 11 300-319 320-339 340-359 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 211. m rH CO rH LO rH CM CM i-\ rH CD O rH rH rH 0 rH CO CO t> rH CD O m rH O T—1 CM CM O rH O CO rH CO O rH O CM O CO O rH rH CM rH rH O in LO CM W / -CPD A/ n W0* // wI/ Ort/ w 0 ft w / Q) 0$ZJ N rH / ‘H O i---£0-. ■ Table 6.4B : Frequency distribution of C. gracilis with size and sex of lizardfish SEX FEMALE MALE SPECIES S. undosquamis 8 0 - 9 0 1 8 1 3 1 0 0 - 1 1 9 102 , 1 2 6 1 2 0 - 1 3 9 4 3 7 20 j 6 4 2 3 9 2 0 1 4 0 - 1 5 9 8 0 2 4 9 2 2 1 2 4 8 8 2 1 7 1 6 0 - 1 7 9 7 0 3 7 4 2 7 1 9 1 8 0 - 1 9 9 2 8 6 2 8 1 3 3 3 2 0 0 - 2 1 9 j 5 8 2 2 0 - 2 3 9 1 3 2 4 0 - 2 5 9 2 6 0 - 2 7 9 2 8 0 - 2 9 9 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 2 1 2 . m CO CO rH m m rHo CM O O 00 rH 00 C oO CO COO CoO CD CD C O CD m t>O CD O G> CCM CCOO CC M CM t> OD CO 00 G 00d O 00 CD rH C 0O0 CO rHm [> •H O CD t> CD in CO CD CD d rH m m rCHD rCHD m 00 > rH rH 0 > 0CMo d 0 rHrH l> CCO t> rH M CoO o rmH 00 0C O rH CM 0 D o ,CQD <0 AU) OCM m COCD CD t> 00 CM Gi in 00 t> CT) CO x: S ‘tS 'H C c> in o CD3 ^ i> CM CM rH rH CG00 •H G «o O 00 CM t*- O O CD O «Hu o m 00 rH g ^ aq rH• r CHD rH00 00 rH rCHD o rH d CD d ̂ . . • • • • • +-> rH. rH rH o o o o o o Os O dO X£I >> G X2 -H CD (D G cti d •oH 04-> «H >ctfdO o u d & d rQ d d £>O Id-H «OH O O0 OCD G w 0 G T0 X - 3 C0O eastern Atlantic and of the Mediterranean. Unesco, Paris. pp.161-162. Nikol’skaya, N.G., and Structural characteristics of I.A. Verigina (1974) the organs of feeding and digestion in three White Sea flounder species in relation to food consumed. J. Ichthyol 14(2):99-109. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 269. Nikolsky, G.V. (1963) "The ecology of fishes”. Academic Press, London. (Russian trans. L. Brikett). 352pp. Nikolsky, G.V. (1969) Theory of fish population dynamics as the biological background for rational exploitation and management of fishery resources. Oliver and Boyd, Edinburgh. 323pp. Nishikawa, S., and Comparative studies on the K. Sakamoto (1978) chromosomes in Japanese fishes 4. Somatic chromosomes of 2 lizardfishes. J. Shimon- oseki Univ. Fish. 27(1): 113-118. Norman, J.R. (1935) A revision of the lizardfishes of genera Synodus, T-radhinooeph- alus, and Saurida. Proc. Zool. Soc. Lond. (1):99-135. Norman, J.R. (1939) Fishes. John Murray Expedit­ ion 1933-34. Sci. Rep., London 7(1):1-116. Nursall, J.R. (1958) The caudal fin as a hydrofoil. Evolution 12:116-120. Ogilby, J.D. (1897) Some new genera and species of fishes: Synodontidae. Proc. Linn. Soc. N.S.W. 22(2): 245- 251. Ogilby, J.D. (1910a) On some new fishes from the Queensland coast. Endeavour Series No. 1:85-189. Ogilby, J.D. (1910b) On new or insufficiently described species: Synodidae. Proc. R. Soc. Queensl. 23(7): 5-6. Okada, R., and Studies on the stock of the K. Kyushin (1955) lizardfish, Saurida turribil, in the East China and the Yellow Seas (1). Bull. Seikai Reg. Fish. Res. Lab. 7:93-112. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 270. Okiyama, M. (1974) The larval taxonomy of the primitive Myctophiform fishes. In : The Early Life History of Fish (Ed. J.H.S. Blaxter) Springer-Verlag, Berlin, Heidelberg, New York. 765pp. Overstreet, R.M. (1968) Parasites of the inshore lizardfish, Synodus foetens from South Florida, including a description of a new genus of cestoda. Bull. Mar. Sci. 18:444-470. Pannella, G. (1974) Otolith growth patterns: an aid in age determination in temperature and tropical fishes. In : Ageing of Fish, pp. 28-39. (Ed. T.B. Bagenal), Unwin Brothers, Old Woking.234 pp. Parr, A .E . (1928) Deepsea fishes of the order Iniomi from the waters around the Bahama and Bermuda Islands. Bull. Bingham Ocean. Coll. 3(3):1-193 (not seen). Parr, A .E . (1929) A contribution to the osteology and classification of the orders Iniomi and Xenoberyces; with description of a new genus and species of the family Scopelarchidae, from the western coast of Mexico; and some notes on the visceral anatomy of Rondeletla. Occ. Pap. Bingham Ocean. Coll. 2:1-45. (not seen) Parukhin, A.M. (1974) Helmintho-fauna in Synodontidae fishes from the Indian Ocean. Vestn. Zool. 3:42-46. Pennycuick, L. (1971a) Seasonal variations in the parasite infections in a pop­ ulation of three-spined sticklebacks, Gasterosteus aouleatus L. Parasitology 63:373-388. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 271. Pennycuick, L. (1971b) Frequency distributions of parasites in a population of three-spined sticklebacks, Gasterosteus aouleatus L. , with particular reference to the negative binomial distribution. Parasitology 63:389-406. Pennycuick, L. (1971c) Differences in the parasites infections in three-spined sticklebacks (Gasterosteus aouleatus L.) of different sex, age and size. Parasit­ ology 63:407-418. Pennycuick, L. (1971d) Quantitative effects of three species of parasites on a population of three-spined sticklebacks, Gasterosteus aouleatus. J. Zool. Lond. 165: 143-162. Petersen, C.G.J. (1892) Fiskensbiologiske forhold i Holboek Fjord, 1890-91. Beret- ning fra de Danske Biologiske Station for 1890(91), 1:121- 183. (not seen). Pianka, E.R. (1978) Evolutionary ecology, Second Edition. Harper and Row, New York. 397pp. Pitcher, T.J., and Fisheries ecology. Croom Helm., P,J.B.Hart (1982) London. 414pp. Pollard, J.H. (1977) A handbook of numerical and statistical techniques (with examples mainly from the life sciences). Cambridge Univers­ ity Press, Cambridge.349pp. Pollock, B.R. (1981) Age determination and growth of luderick, Girella tricuspidata (Quoy and Gaimard), taken frcm Moreton Bay, Australia. J. Fish Biol. 19:475-485. Qasim, S.Z. (1973) An appraisal of the studies on maturation and spawning in marine teleosts from the Indian waters. Indian J. Fish. 20:166-181. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 272. Quoy, J.R.C., and Voyage autour du monde ... P. Gaimard (1824) ex6cut£ sur les corvettes de S.M. "L'Uranie" et "La Physicienne" pendent les ann6es 1817-20 Poissons. Pillet Am6, Paris. Radhakrishnan, N. (1963) Notes on the maturity and spawning of Opisthopterus tardoore (Cuvier). Indian J. Fish. 10:102-107. Ramsay, E.P. (1883) Description of some new Australian fishes. Proc. Linn. Soc. N.S.W. 8:177-180. Rao, K.V.S. (1974) Studies on some aspects of the biology and fishery of lizard­ fishes {Saurida spp.) and the 1 Ghol1 , Pseudosciaena diacanthus (Lac6pede) from Indian waters. Ph.D. thesis, Banaras Hindu University. 186pp. Unpublished. Rao, K.V.S. (1977) Systematics and comparative osteology of Indian fishes (Saurida spp). Indian J. Fish. 24(1-2):143-171. Re, P. (1978) Contribution to the knowledge of littoral fish fauna of Ilhas Desertas Madeira, Portugal. Bolm Soc. Port. Cienc. Nat. 18(2):47-50. Redfield, J.A., and Techniques of starch-gel J.P. Salini (1980) electrophoresis of penaeid prawn enzymes (Penaeus spp. and Metapenaeus spp.). CSIRO Division of Fisheries and Oceanography Report 116:1-20. Rees, G. (1969) Cestodes from Bermuda fishes and an account of Acompsocephalum tortum (Linton, 1905) gen. nov. from the lizardfish Synodus intermedius (Agassiz). Para­ sitology 59:519-548. Regan, C.T. (1909) The classification of tele- ostean fishes. Ann. Mag. Nat. Hist. 3(8):75-86.(not seen). University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 273. Regan, C.T. (1911) The anatomy and classific­ ation of the teleostean fishes of the order Iniomi. Ann. Mag. Nat. Hist. 7(8): 120-133. (not seen). Regan, C.T. (1929) Article on fishes. Encyclo­ pedia Brittanica, 14th ed. pp.305-328. (not seen). Richardson, J. (1843) Aulopus purpurissatus (Nob. ) , Imperial Aulopus. In : Iconespiscium, or plates of rare fishes. Richard and John Taylor, London. Richardson, J. (1845) On the Ichthyology of the Seas of China and Japan (with a new index to names in Richardson's Report on the Ichthyology of the Seas of China and Japan) by P.J. Whitehead. Reprinted by Antiquariaat Junk Dr R. Schierenberg and Sons N.V. P.O. Box 5 LOCHEM/Nether- lands, 1972. 320pp. Richardson, J. (1848) Saurus undosquamis. In : Zool. ’Erebus1 and ’Terror', Fish pp. 138-139. Ricker, W.E. (1973) Linear regressions in fishery research. J. Fish. Res. Board Can. 30(3):409-434. Ricker, W.E. (1975) Computation and interpret­ ation of biological statistics of fish populations. Bull. Fish. Res. Board Can. 191: 382pp. Ricker, W.E. (1977) The historical development. In : Fish population dynamics, pp. 1-21. (Ed. J.A. Gulland), John Wiley and Sons, New York. 372pp. Ricker, W.E., and On the methods of measuring D. Merriman (1945) fish. Copeia, 1945(4):184-191. Ridewood, W.G. (1896) The teeth of fishes. Natural Science Vol. 8:380-391. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 274. Romer, A.S. (1945) Vertebrate paleontology. 2nd ed. Chicago. 687pp. (not seen). Rosen, D.E. (1973) Inter-relationships of higher euteleostean fishes. Zool. J. Linn. Soc. 53 (Suppl. 1) :397-513. Rowling, K.R. (1975) The preliminary study of the feeding habits of 3 families of coastal fishes from the Townsville region of North Queensland. BSc (Hons) Thesis. Unpublished. Royce, W.F. (1972) Introduction to the fishery sciences. Academic Press, New York.351pp. Ruamragsa S., and An analysis of demersal catches A.P. Isarankura (1965) taken from the experimental trawling operations in the Gulf of Thailand. Department of Fisheries, Bangkok, Thailand. Contribution No. 3 pp. 1-51. Ruello, N .V . (1975) A small beam trawl for sampling surface of demersal and benthic animals. Aust­ ralian Society for Limnology Bulletin No. 6:9-16. Russell, B.C., and Three new species of Indo- R.F. Cressey (1979) West Pacific lizardfish (Synodontidae). Proc. Biol. Soc. Wash. 92(1):166-175. Russell, F.S. (1976) The eggs and planktonic stages of British marine fishes. Academic Press, London.524pp. Sainsbury, K., and Age, growth, mortality and W. Whitelaw (1981) feeding of Nemipterus pevonii, Saurida undosquamis and Lethrinus choerhynchus on the north west shelf of Australia. CSIRO Division of Fisheries Research, Divisional, Research Seminar, pp.63-64. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 275. Sanzo, L. (1915) Contributo alia conoscenza dello sviluppo negli Soopelini Muller (Saurus griseus Lowe, Chlorophthalmus agassizi Bp. ed. Aulopus filamentosus Cuv . ) . Mem. R. Comit. Talasso. Italiano 49:1-21. (not seen) Schoener, T.W. (1971) Theory of feeding strategies. Annu. Rev. Ecol. Syst. 2: 369-404. Senter, R.J. (1969) Analysis of data. Scott, Foresman and Company, Glen­ view. Illinois. 512pp. Shaklee, J .B ., Developmental genetics of M.J. Champion, and teleosts: A biochemical G.S. Whitt (1974) analysis of Lake Chubsucker ontogeny. Dev. Biol. 38:356- 382. Shaklee, J.B. Speciation and evolution of C.S. Tamaru, and marine fishes studied by the R.S. Waples (1982) electrophoretic analysis of proteins. Pac. Sci. 36(2): 141-157. Shimizu, Y., Extractability of proteins S. Karata, and from fish skeletal muscle at F. Nishioka (1976) low ionic strengths. Bull. Jpn. Soc. Sci. Fish. 42(9): 1025-1031. Shindo, S. (1968) On the preparation of an illustrated key for Saurida (lizardfishes) in the Indo- Pacific waters. CSK Working Papers, Honolulu Apr. 29- May 2, 1968. Shindo, S., and Description of three new U. Yamada (1972) species of the lizardfish genus Saurida, with a key to its Indo-Pacific species. Uo Japan Soc. Ichthyol. No.11: 1-13, No. 12:1-14. Sindermann, C.J. (1966) Diseases of marine fishes. Academic Press, London.89pp. Sinoda, M., and S. Intong (1978) Size frequency distribution of lizardfish, Saurida undo­ squamis, in the inner Gulf of Thailand. Bull. Jpn. Soc. Sci. Fish. 44(1):1-6. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 276. Sinoda, M., Preliminary study of trash P.Y. Lim, and fish landed at Kangkar fish S.M . Tan (1978) market in Singapore. Bull. Jpn. Soc. Sci. Fish. 44(6): 595:600. Smith, H .M . (1969) Parapatry: Sympatry or allopatry? Syst. Zool. 18: 254-255. Smith, R.J. (1982) On the mechanical reduction of functional morphology. J. Theor. Biol. 96:99-106. Sokal, R .R., and Biometry. W.H. Freeman and R.J. Rohlf (1969) Company, San Francisco. 776pp. Springer, S. (1946) A collection of fishes from the stomachs of sharks taken off Salerno, Florida. Copeia, 1946:174-175. Stanbury: P.J. (1968-69) Type specimens in the Macleay Museum, University of Sydney. Proc. Linn. Soc. N.S. Wales 93(2):203-210. Steinmetz, B. (1974) Scale reading and back- calculation of bream Abvamis brama (L.) and rudd Soardinius erythrophthalmas. In : Ageing of fish, pp. 148-157 (Ed. T.B. 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(1929) Life history of the lake herring (Leuoiohthys artedii LeSueur ) of Lake Huron as revealed by its scales, with a critique of the scale method. Bull. U.S. Bur. Fish. 44:265-428. (not seen). Varadarajan, S., and Reproduction of the continu­ T .Subramonlara (1982) ously breeding tropical hermit crab Clibanarius clibanarius. Mar. Ecol. Prog. Ser. 8:197- 201. Vaughan, G.E. and Sublethal effects of three D.W. Coble (1975) ecto-parasites on fish. J. Fish Biol. 7:283-294. Von Bertalanffy, L. (1938) - A quantitative theory of organic growth. Hum. Biol. 10:181-213. Waite, E.R. (1905-1908) Notes on fishes from Western Australia. Rec. Aust. Mus. Syd. 6:55-82. Waples, R.S. (1982) A biochemical and morpho­ logical review of the lizard­ fish genus Saurida in Hawaii, with the description of a new species. Pac. Sci. 35(3): 217-235. Weber, M., and The fishes of the Indo- L.F. de Beaufort (1913) Australian Archipelago. II. Malacopterygii, Myctophoidea, Ostariophysi : Siluroidea. Leiden. 404pp. Westrheim, S.J. (1973) Age determination and growth of Pacific Ocean perch (Sebastes alutus) in the north­ east Pacific Ocean. J. Fish. Res. Board Can. 30(2):235-247 - Wheeler, A. (1975) Fishes of the World. Ferndale Editions, London.366pp. Whitfield, P. (1978) The Hunters. Hamlyn, London. 160pp. Whitley, G.P. (1937) The Middleton and Elizabeth Reefs, South Pacific Ocean: Fishes. Aust. Zool. 8(4):214- 231. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 282. Whitley, G.P. (1948) A list of the fishes of Western Australia. Western Australia Fisheries Depart­ ment, Fisheries Bulletin No. 2:1-35. Whitley, G.P. (1943) Icthyological notes and illustrations (Part 2). Aust. Zool. 10(2):167-187- Wootten, R., and Studies on the biology of I.F. Waddell (1976-77) larval nematodes from the musculature of cod and whiting in Scottish waters. J. Cons. Int. Explor. Mer. 37(3):266-273. Wu, H.W., and Four new fishes from Chefoo. 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Fish. 46(2):191-195 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 283. Yasuda, F. (1960a) The feeding mechanism in young fishes. Rec. Oceanogr. Works Jpn. 5(2):132-138. Yasuda, F. (1960b) The relationship of the gill structure and food habits of some coastal fishes in Japan. Rec. Oceanogr. Works Jpn. 5(2):139-152. Yasuda, F. (1960c) The feeding mechanism in some carnivorous fishes. Rec. Oceanogr. Works Jpn. 5(2):153- 160. Yeh, S.Y., Age and growth of lizardfish, H.L. Lai, and Saurida tumbil (Bloch), in the H.C. Liu (1977) East China Sea and the Gulf of Tonkin. ACTA Oceano- graphica Taiwanica. Science Reports of the National Taiwan University No. 7:134-145. Yeh, S .Y ., and Comparative morphometric and H.C. Liu (1973) meristic studies of lizardfish (Saurida tumbil) from the South China Seas. J. Fish. Soc. Taiwan 2(2):59-74. Yazdani, G.M. (1969) Adaptations in the jaws of flatfish (Pleuronectiformes). J. Zool., Lond. 159:181-222. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 284. A P P E N D I X 1 1 il •-- ±. j x. CIIIUL V L̂uiiicicns in total sample Month and Year S.undosquamis S. tumbil S .filamentosa T. myops Nov. 1979 17.9 - Dec. 1979 - 48.1 Jan. 1980 87.0 68.5 Feb. 1980 28.3 67.6 March 1980 45.2 65.5 April 1980 69.4 51.7 May 1980 68.0 - June 1980 58.6 82.1 31.6 July 1980 66.1 - Aug. 1980 53.2 - Sept. 1980 55.0 43.4 40.9 Oct. 1980 41.2 - 46.2 Nov. 1980 61.4 - 50.0 Dec. 1980 62.5 — 28.9 Jan. 1981 47.4 59.6 42.3 54.4 Feb. 1981 53.8 60.0 66.7 March 1981 54.3 85.4 62.9 April 1981 61.7 64.8 81.0 May 1981 72.7 62.6 55.0 June 1981 66.2 67.2 56.3 July 1981 78.9 91.8 Aug. 1981 70.1 87.5 Sept. 1981 59.5 74.4 Oct. 1981 68.2 73.4 45.8 Nov. 1981 64.3 - 40.0 Dec. 1981 52.8 - Jan. 1982 1982 41.1 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 2 6 S. vO O CN •vT O' o rH o o o O o CN o O o O z o O' o CO 00 r—i CJ o o o o *H •VO o 14 r-i r-i CN o W 00 ON rH rH • o o O' r-. o cn m -* 3- vO -J r-» cn -j -a O 'S’ s? 'T r*»ncn ri cn o CrNCN H rH m vO u. -a- 00 -J cn \0 O COO CN O o o o 3 rH' rrHH 'O' —1 orH -a o r--J« \0 o ’ vr vO v£> o o o o o o o O' Or'H o 3- o CN Z N Z K Z H aepyuo8odv 3 B p jq 3 B U 8 o y 3 T de oByioq e n j ^ APPENDIX 2: ABUNDANCE (N ) , AND PERCENTAGE (Z) OCCURRENCE OF PREY ITEMS IN STOMACHS OF S. undoaquamie (S.U) and S. tumbil (s.T) ) Y E A R Year 1981 Month November December Jan. Feb. April May June July Aug. Sept. Oct. Dec. Jan. Feb. March N 905 1565 872 855 591 90 516 157 317 270 379 Z 9.0 15.5 8.6 8.5 5 .8 0 .9 5.1 1.6 3.1 2 .7 3 .7 Z in S.U 12.1 33.3 2.3 3.3 9.1 5.1 3.0 5.9 Z in S.T 8.7 8.3 12.5 513 620 473 597 546 1038 1827 589 1217 516 3 .1 3.7 2 .8 3.6 3.3 6 .2 10.9 3.5 7 .3 3 .1 Z in S.U 0 .6 2.7 18.2 13.4 22.7 Z in S .T 6.7 8 .3 6.7 7.7 11.6 12.5 158 209 519 2020 1255 905 609 211 205 72 194 175 245 515 266 746 1.7 2 .2 5.6 21.7 13.5 9.7 6.5 2 .3 2.2 0 .8 2.1 1.9 2.6 5 .5 2 .9 8.0 Z in S.U 7.9 15.3 26.0 2 .5 1.5 20.0 12.3 Z in S.T 8.3 627 2282 670 6 7 10 1368 182 1220 168 132 139 7.1 25.8 7.6 30.1 30 .! 0 .1 15.5 2.1 13.8 1.9 1.5 1.6 Z in S.U 7.1 5.8 9.1 Z in S.T 20.0 9.3 12.5 0*9 0*01 L'L University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 286. APPENDIX 3. A. DAILY VARIATION IN FEEDING AND ACTIVITY OF S a u r id a u n d o s g u a m is . Day 17 June 1980 10 September 1980 21 October 1981 Time Number Percent Percent Percent (Hours) Number Percent stomach stomach Time Number Percent of of of of with at least (Hours) of ofFish Total Fish Total food half-full Fish Total 1000 9 4.0 33.3 33.3 0940 221 14.4 1100 24 10.3 29.2 20.8 1035 111 7.2 1200 37 12.8 9 4.0 77.7 44.4 1140 297 19.4 1300 67 23.2 5 2.1 0.0 0.0 1200 * - 1400 46 15.9 4 1.7 25.0 25.0 1340 335 21.9 1500 . 45 15.6 5 2.1 0.0 0.0 1540 239 15.6 1600 36 12.5 47 20.2 39.1 17.4 1740 107 7.0 1700 * 43 18.5 46.5 41.8 1840 22 1.4 1800 21 7.3 3 1.3 33.3 33.3 1940 13 0.8 1900 17 5.9 * 2040 15 1.0 2000 8 2.8 5 2.1 100.0 100.0 2140 8 0.5 2100 8 2.8 2 0.8 100.0 100.0 2240 24 1.6 2200 4 1.4 9 4.0 44.4 22.2 2340 10 0.6 2300 0 0.0 3 1.3 66.7 66.7 0040 8 0.5 2400 0 0.0 5 2.1 40.0 20.0 0205 11 0.7 0100 5 2.1 100.0 100.0 0300 10 0.6 0200 1 0.4 0.0 0.0 0340 10 0.6 0300 6 2.6 50.0 16.7 0440 32 2.1 0400 8 3.4 71.4 57.1 0540 60 3.9 0500 3 1.3 66.7 66.7 0600 19 8.2 47.4 31.6 0700 4 1.7 50,0 50.0 0800 4 1.7 20.0 0.0 6 specimens 20 specimens obtained in all; caught between Time Number 15 between 1200 and 2300 Hrs, 1200 and 1600 0940 1140 44 S. 12 of these at 1600 Hrs.tumbil Hrs. 5 of them 1340 - 1540 6 between 1500 and at 2240 2 1600 Hrs. * OTTER BOARDS TANGLED WITH NETS. 2, 3 and 7 r. myops were obtained on 17 June 1980, 10 September 1980 and 21 October 1981 respectively. B. TIDES 16th 17th 18th HOURS METRES HOURS METRES HOURS METRES 0501 0.4 0547 0.4 0048 2.2 1045 2.0 1130 1.8 0734 0.6 1649 0.3 1727 0.4 1321 1.6 2313 2.5 2356 .2.4 1900 0.7 9th 10th 11th HOURS METRES HOURS METRES HOURS METRES 0334 0.3 0403 0.3 0431 0.3 0924 1.9 0957 1.9 1030 2.0 1525 0.3 1600 0.3 1634 0.3 2136 2.2 2207 2.2 2237 2.1 20th 21st 22nd HOURS METRES HOURS METRES HOURS METRES 0236 1.4 0402 1.4 0520 1.5 0830 0.4 0952 0.5 1112 0.5 1518 2.0 1633 2.0 1739 2.0 2214 0.5 2326 0.4 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 288. APPENDIX 4B. Saurida undoequarrrCe : Variation in Beale size with body site (See F ig .4 .2 ) C.V. - Coefficient of Variation University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 289. University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 290. Appendix 5A: Mean standard length (Jj) and mean whole body weight (W) of S. _____________________ Sex Female Male Tofo tasla mpnluemsber 478 475 Size class No, of Mean L Mean W No. of Mean L Mean Wsamples (mm) Cg) samples (mm) Cg) 100 - 109 3 103.0 15.87 1 102.0 10.50 120 - 129 2 125.0 20.65 1 124. 0 21.60 130 - 139 7 135.7 26. 91 14 136.0 28.30 140 - 149 18 144.6 34.23 39 144.6 35.52 150 - 159 32 152.8 40.95 35 152.5 41.66 160 - 169 13 163.6 51.44 49 164.7 53.51 170 - 179 12 174.4 59.87 119 173.5 64.19 180 - 189 8 182.4 73.29 121 183.3 73.26 190 - 199 9 193.6 88.40 53 191.8 84.36 200 - 209 27 204. 3 98.67 17 202.2 96.60 210 - 219 37 213.2 116.73 4 211.5 105.40 220 - 229 41 223.8 132.54 6 222.3 131.38 230 - 239 48 233.3 147.61 4 232.8 149.68 240 - 249 47 243.2 173.09 3 245.0 186.35 250 - 259 43 253.8 189.78 260 - 269 48 263.0 219.29 270 - 279 23 273.6 259.01 280 - 289 20 282.6 282.08 290 - 299 14 292.8 321.75 300 - 309 11 301.9 333.34 310 - 319 3 313.7 421.03 320 - 329 4 322.5 437.38 330 - 339 2 335.00 439.80 340 - 349 2 342.5 549.25 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 291. Appendix 5B: Mean standard length (L) and mean whole body weight (W) of -S. undo^Quami6.__________________ Sex Female Male Total number of samples 2606 2492 Size class No. of Mean L Mean W No. of Mean L Mean W samples (mm) (g) samples (mm) (g) 80 - 89 3 87.3 7.33 3 84.7 6.40 90 - 99 16 94.9 9.92 10 96.6 8. 97 100 - 109 26 104.6 11.10 27 104.0 11.81 110 - 119 79 114.5 14.70 99 114.7 14.81 120 - 129 154 124.3 19.24 253 125.0 19.41 130 - 139 294 134.5 24.72 421 134.6 25.47 140 - 149 355 144.2 29.98 740 144.2 31.71 150 - 159 491 153.9 36.98 582 153.5 38.74 160 - 169 434 164.0 44. 94 244 163.3 46.45 170 - 179 342 173.7 52.97 79 172.9 56.16 180 - 189 209 183.0 62.33 22 182.2 66.06 190 - 199 109 192.9 75.62 9 191.7 79.56 200 - 209 42 203.3 91.53 1 203.0 96.90 210 - 219 22 214.2 103.19 220 - 229 13 224.2 127.63 230 - 239 4 233.3 142.03 1 237.0 144.20 240 - 249 4 244.5 166.35 250 - 259 2 254.0 179.05 260 - 269 3 263.3 214.10 270 - 279 2 275.5 247.00 280 - 289 1 280.0 269.10 300 - 309 1 300. 0 238.80 320 - 329 1 320.0 340.70 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 292. Appendix 5C: Mean standard length (L) and mean whole body weight (W) of S. ____________________ Sex Female Male Total number of samples 350 278 Size Class No. of Mean L Mean W No. of Mean L Mean W samples (mm) (g) samples (mm) (g) 90 - 99 1 96.0 7.10 100 - 109 1 105.0 14.80 110 - 119 4 115.0 12.70 23 114.2 13.58 120 - 129 15 124.7 17.17 23 123.4 16.65 130 - 139 24 133.7 22.45 43 134.6 22.29 140 - 149 20 142.5 26.59 43 142.8 26.89 150 - 159 20 152. 9 32.63 25 153.3 35.14 160 - 169 10 163.5 42.98 10 162.9 41.72 170 - 179 5 173.4 52.30 19 173.7 55.75 180 - 189 6 182. 5 62.17 5 183.0 63.82 190 - 199 7 190.4 73. 91 9 193.6 79.84 200 - 209 5 204.6 95.38 11 204.4 91.37 210 - 219 11 215.3 104.65 17 213.1 104.8 220 - 229 22 224.6 119.32 13 223.1 122.36 230 - 239 25 232. 8 134.62 9 235.4 138.03 240 - 249 22 243. 0 160.67 5 246.4 167.84 250 - 259 27 253.1 184.22 6 252.0 173.02 260 - 269 19 263.9 199.96 1 265. 0 216.00 270 - 279 17 272. 9 232.88 1 276.2 242.24 280 - 289 24 283.5 284.27 290 - 299 15 293.2 323.03 2 295.0 283.20 300 - 309 15 303.1 348.90 310 - 319 8 313.5 371.03 320 - 329 10 322.7 420.71 1 325.0 399.80 330 - 339 8 332.8 484.90 340 - 349 4 342. 0 518.25 1 342.0 353.70 360 - 369 1 361.0 579.90 370 - 379 1 370. 0 576.50 390 - 399 1 390.0 737.60 420 - 429 1 424. 0 872.80 490 - 499 1 490.0 1066.60 570 - 579 1 575. 0 1483.20 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 293. Appendix 5D: Mean standard length CL) and mean whole body weight (W) of T. myop4. ______ __ Sex Female Male Total number of samples 497 537 Size Class No. of Mean L Mean W No. of Mean L Mean W samples (mm) Cg) samples (mm) (g) 80 - 89 6 85.8 8.50 8 85.9 8.50 90 - 99 20 95.0 11.47 13 95.3 11.53 100 - 109 20 102.8 15.39 17 104.5 16.14 110 - 119 27 114.7 20.83 28 114.8 21.07 120 - 129 28 124.1 26.59 46 124.5 27.04 130 - 139 38 134.5 34.46 83 133. 9 33.78 140 - 149 65 143.8 42.83 114 144.2 42.91 150 - 159 59 154.8 54.15 106 153.8 52.53 160 - 169 48 164.3 67.28 62 163.4 64.98 170 - 179 46 173.8 82.33 42 173.2 74.63 180 - 189 29 183.2 98.31 16 182.2 87.64 190 - 199 32 194.2 117.97 2 192.5 100.00 200 - 209 31 203.7 135.11 210 - 219 22 212.4 153.60 220 - 229 16 223.6 179.79 230 - 239 4 234.8 199.23 240 - 249 5 243.0 218.06 250 - 259 1 253.0 267.60 University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 294. APPENDIX 6: GONAD INDICES OF SYNODONTIDAE. T. my of8 S. undoequamie S. twribil S. filartient08a and lale Femalf M[ale Femalc Male Peniale ■lale Fema Le Year nean min mean min mean mpan mpfln TTIPY min Tipnn ... mi n megH max ml n Tnpan TT1PV Min N o v .'79 0.43 0 .57 3.68 0.27 0.97 8.46 0 JO 0 .40 0.50 0.4C 0.57 0 .90 D e c .'79 3.50 0 .60 ).90 1.26 2.02 2.77 0.3! 0 .65 1.85 0.29 2.71 7.84 J a n .'80 0.60 1 .18 2.53 2.09 7.96 17.12 0 . 2( 1 .40 3.88 0.46 8.00 3..41 1 .0 1 .0 1 .0 Feb.'8 0 0.50 1.18 2.60 0 .29 1.01 2.56 0.8( 2.00 3.25 0.30 4 .68 12.90 Mar.’80 3.41 0 .95 3.28 0.29 1.02 13.01 0 .H 1.14 3.29 0.10 1.17 5 .38 Apr.’80 3.36 0 .69 2.38 0 .23 0 .58 5.07 O.lf 0 .82 2.24 0.49 1.02 4 .30 May ’80 3.39 0 .68 L.06 0.20 0.55 1 .34 0.30 0.79 1.54 0.36 0 .90 2.40 0.20 D.35 0.33 0 .20 2.64 6 .X Jun. ' 80 3.23 0 .62 L. 37 0 .21 0.47 1.13 0 . 3 : 0 .43 0 .61 0.24 0 .72 2.03 0.23 0.23 0.23 0 .15 1.80 9.2D 1.90 1.90 1.90 J u l . '80 3.30 0 .65 L.39 0.22 0 .52 1.15 0.3^ 0 .48 0 .75 0.50 0.60 0.79 1.40 1 .67 1.80 1.20 1 .30 1 .40 Aug. ' 80 3.37 0 .69 J..91 0 .28 0.72 2 .41 0 . 3f 0 .47 0 .75 0.51 0 .81 1.01 0 .20 3.60 1.20 0 .30 1 .93 L2.9 1 .40 1 .40 1.40 Sept. '83 0.28 0 .64 ..37 3.30 1.58 12.11 o .4: 0 .77 1.07 0.52 1 .00 1.33 0 .16 0.22 0.31 0 .12 1 .62 L9.08 1 .00 1.00 1.00 1.80 1 .80 1 .80 Oct.'80 0.37 0 .89 ..89 0.43 4.15 10.16 0.8] 1 .25 5 .62 1.73 2.81 5 .01 0 .20 3.52 1.00 0 .10 0 .23 0.4C 0 .36 0 .76 1.21 0.37 2 .35 7.39 Nov.'80 D .ll 0 .71 L. 20 3.34 2.21 34.47 3.1C 2.25 7.50 2.80 4 .76 7.40 0 .30 ) .30 0.30 0 .20 0 .25 0.33 0 .54 1 .88 4.96 0 .39 5 .10 14.77 Dec. ' 80 0.28 0 .73 ..94 3.30 2.75 13.89 0.4C 1.72 3.20 4.21 6.70 10.82 0 .32 1 .81 4.04 0 .39 3 .31 H .8 8 Jan.'81 0.30 0 .57 ..04 3.32 2.32 8 .75 0.34 1 .26 3.60 2.73 7.25 5 .8 4 0 .18 ) .65 2.14 0 .12 1 .32 9.1? 0 .50 2 .10 4.31 0 .57 5 .00 H .5 5 Feb.'81 0.41 1.00 1.77 3.36 0 .92 2.94 1.31 2.06 2.66 2.92 7.84 B .72 0.15 ) .56 1.12 0 .28 2 .42 8.54 1.70 1 .93 2.10 2.40 4 .65 6 .30 ter.'81 ).33 0 .88 2.35 3.26 0 .85 9 .54 0 .4 ! 1 .96 5 .61 1.65 8.71 18.52 0 .18 0 .94 1.6* 0 .15 1 .50 9.56 0.60 0 .60 0 .60 6 .80 6 .80 6 .80 Apr.'81 ). 31 0 .63 2.27 3.25 0 .61 2.86 0.24 0.66 1 .45 0.18 L .12 5 .68 0.10 ) .15 0.20 0 .4 2 .65 4.90 0 .51 1 .23 2.26 0 .00 5 .30 17.11 ■lay '81 3. 34 0.59 2.22 3.24 0.50 1.17 0.18 0.59 1.76 0 .27 3.60 1.47 0.16 0 .36 1.48 0 .10 0 .55 5 .7 : Jun.' 81 ). 31 0 .58 1.43 3.18 0.46 1.50 0 .41 0.55 0 .73 0.41 3.59 1.27 0 .15 0 .40 0.79 0 .16 0 .29 0 .4 ! July'81 ). 26 0.70 2.02 3.21 0.60 1.70 0 .27 0.52 0 .77 0 .23 3.61 1.29 0 .50 0 .50 0.5C 0 .20 1 .70 8.0C 3.7 0 .7 0 .7 0.40 0 .60 0 .80 \ug.'8l ). 30 0.69 2.38 ) .25 0.60 2.20 0 .29 0.72 1 .91 0 .40 3.68 1 .31 >ept.'81 ).29 0.62 L.20 3.29 1.04 5.60 0 .24 3.68 2.56 0 .09 3.65 1.30 0 .20 0 .30 0.40 0.20 1.90 D .10 0.50 1.00 0 .50 0.26 0.70 2.75 3.19 5.22 24.1] 0 .24 0.91 2.17 0 .95 3.02 5.65 3.34 1.12 2 .29 0.00 3.19 L3.2C tov.'81 ). 39 0 .95 2.06 3.39 1.12 3.38 0 .70 3.70 0.70 0 ,60 3.60 0 .60 3.88 1.67 2.56 0.55 4.07 7 .13 )ec.'8l 0,31 0.80 L. 33 ). 25 2.07 9 .72 Jan.'82 ?eb.'82 3.30 0,30 0.30 3.30 3,40 9 .50 L.06 3,14 5.46 4.01 8.86 .5.04 min - minimum max - maximum r—! oo Li University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 295. S.undo^quamU S. tumbril S. {i4JLame.n£oAa T. myopA MONTH AND YEAR *F M V *F M V *F M V *F M V November 1979 32 6 0 31 20 0 December 1979 21 10 0 17 9 0 January 1980 11 12 0 74 62 0 0 1 0 February 1980 106 79 54 18 19 0 March 1980 165 187 6 16 12 0 April 1980 213 177 0 20 9 0 May 1980 64 77 1 16 6 0 5 2 0 June 1980 150 116 2 19 8 3 17 1 0 1 0 0 July 1980 159 187 3 23 18 0 3 3 0 August 1980 24 17 6 30 25 2 11 3 0 0 1 0 September 1980 98 115 3 9 10 2 17 3 0 1 1 0 October 1980 55 56 4 15 17 1 3 5 0 18 7 0 November 1980 34 62 1 55 90 0 2 2 0 84 110 0 December 1980 31 29 5 50 50 0 202 162 32 January 1981 80 60 4 53 62 0 53 73 1 49 96 0 February 1981 56 79 4 20 12 0 18 5 0 4 4 0 March 1981 306 106 11 28 22 0 L17 L42 0 1 1 0 April 1981 144 260 17 58 26 2 2 4 0 10 11 0 May 1981 83 98 0 29 68 0 69 16 0 June 1981 219 116 0 21 36 0 10 6 0 July 1981 170 176 2 28 21 0 12 1 0 2 1 0 August 1981 171 167 1 23 12 12 September 1981 98 64 8 19 12 3 6 2 0 2 0 0 October 1981 156 280 0 9 39 0 76 79 0 November 1981 31 31 2 1 1 0 7 23 0 December 1981 19 13 1 February 1982 3 4 0 43 37 0 * F - Female M Male. V - Very young fish University of Ghana U n i v e r s it y o f G h a n ah t t p : / / uhgttps:p//augcsep.aucge..eudg.ue.dguh.gh 296. Appendix 8: SAUR1VA: MONTHLY SEX RATIOS. MONTH AND S. undoAquam'LA S. tumbril YEAR * fr equency (f) Chi- * fr equency Chi- fo(F) ofM') fe square fo(F) fo(M) UJfe sauare Nov. 1979 32 6 19 17.789 31 20 25.5 2.372 Dec. 1979 21 10 15.5 3.903 17 9 13 2.461 Jan. 1980 11 12 11.5 0.043 74 62 68 1.059 Feb. 1980 106 79 92.5 3.941 18 19 18.5 0.027 March 1980 165 187 176 1.375 16 12 14 0.571 April 1980 213 177 195 3.323 20 9 14.5 4.172 May 1980 64 77 70.5 1.999 16 6 11 4.550 June 1980 150 116 133 4.346 19 8 13.5 4.481 July 1980 159 187 173 2.266 23 18 20.5 0.610 Aug. 1980 24 17 20.5 1.195 30 25 27.5 0.455 Sept. 1980 98 115 106.5 1.357 9 10 9.5 0.53 Oct. 1980 55 56 55.5 0.009 15 17 16 0.125 Nov. 1980 34 62 48 8.167 55 60 57.5 0.217 Dec. 1980 31 29 30 0.067 50 50 50 0 Jan. 1981 80 60 70 2.857 53 62 57.5 0.704 Feb. 1981 56 79 67.5 3.919 20 12 16 2.00 March 1981 306 106 206 97.087 28 22 25 0.720 April 1981 144 260 202 3a 307 58 26 42 12.190 May 1981 83 98 90.5 1.243 29 68 48.5 15.680 June 1981 219 116 167.5 31668 21 36 28.5 3.917 July 1981 170 176 173 0.104 28 21 24.5 1.000 Aug. 1981 171 167 169 0.047 23 12 17.5 3.457 Sept. 1981 98 64 81 7.135 19 12 15.5 1.581 Oct. 1981 156 280 218 3^ 266 9 39 24 18.750 Nov. 1981 31 31 31 0 1 1 1 0 Dec. 1981 19 13 16 1.125 fo(.F) " observed (female). fQ(M) = observed (male). fg = expected.