Ecology and Biology of the Banded Wrasse, Notolabrus Fucicola
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ECOLOGY AND BIOLOGY OF TH~ BANDED WRASSE, NOTOLABRUS FUCICOLA (PISCES:LABRIDAE) AROUND KAI KOU RA, NEW ZEALAND. A thesis submitted in the partial fulfillment of the requirements for the Degree of Master of Science in Zoology at the University of Canterbury New Zealand By Christopher Michael Denny UNIVERSITY OF CANTERBURY 1998 Table of Contents. iable of Contents List of Figures iv List of Tables viii List of Plates x Acknowledgements xiii Abstract xiv CHAPTER ONE. GENERAL INTRODUCTION 1.1 GENERAL INTRODUCTION 1 1.2 STUDY AREA 9 CHAPTER TWO. DEMOGRAPHY AND DISTRIBUTION 2.1 INTRODUCTION 10 2.3 METHODS . 15 2.3.1 STUDY SITES 15 2.3.2 SAMPLING DESIGN 16 2.4 RESULTS 19 2.5 DISCUSSION 27 CHAPTER THREE. LIFE HISTORY 3.1 INTRODUCTION 32 3.1.1 REPRODUCTIVE BIOLOGY 32 3.1.2 REPRODUCTIVE SEASONALITY 38 3.1.3 AGE AND GROWTH 41 Table of Contents. ii 3.2 MATERIAL AND METHODS 42 3.2.1 GENERAL METHODS 42 3.2.2 HISTOLOGY 43 3.2.3 AGE AND GROWTH 46 3.3 RESULTS 48 3.3.1 RELATIONSHIP BETWEEN COLOUR PHASE, SIZE AND SEX 48 3.3.2 REPRODUCTIVE SEASONALITY 50 3.3.3 HISTOLOGY 56 3.3.4 AGE AND GROWTH 64 3.4 DISCUSSION 67 3.4.1 RELATIONSHIP BETWEEN COLOUR PHASE, SIZE AND SEX 67 3.4.2 REPRODUCTIVE SEASONALITY 68 3.4.2 REPRODUCTIVE BIOLOGY 70 3.4.3 AGE AND GROWTH 75 CHAPTER FOUR. DIET 4.1 INTRODUCTION 78 4.2 MATERIAL AND METHODS 81 4.3 RESULTS 83 4.3.1 PREY ITEMS 83 4.3.2 ONTOGENETIC CHANGES 86 4.3.3 SEASONAL PATIERNS 89 4.3.4 GUT FULLNESS AND CONDITION FACTOR ,, 92 4.4 DISCUSSION 94 Table of Contents. iii CHAPTER FIVE. BEHAVIOUR 5.1 INTRODUCTION 98 5.2 METHODS 101 5.3 RESULTS 102 5.4 DISCUSSION 105 CHAPTER SIX. CONCLUSIONS 6.1 CONCLUSIONS 108 REFERENCES 112 APPENDIX ONE. DIETARY ITEMS 135 List of Figures. iv LIST OF FIGURES CHAPTER ONE Figure 1.1 Map of the Kaikoura Peninsula showing study sites with an 9 inset depicting its location in New Zealand. CHAPTER TWO 2 Figure 2.1 Mean number of species observed per transect (150 m ) (+ 20 s.e.) in each of the nine sites at three depths (0-6 m, 7-11 m and 12-18 m) from Dec 1996 to Dec 1997 (65 transects per site). Figure 2.2 Mean number per site of eleven species of reef fish observed 21 2 per transect (150 m ) (n = 65) over the study period (Dec 1996 - Dec 1997) at nine sites at three depths. A: Shallow sites 0-6m, B: Intermediate sites 7-11 m, C: Deep sites 12-1 Bm (+ s.e.). See Table 2.2 for species codes. Figure 2.3 Mean number of fish per transect (n = 65) for four common 24 reef fish at nine sites around Kaikoura from Dec 1996 to Dec 1997 ( +/ s.e.). See Table 2.2 for species codes. Figure 2.4 Mean size (cm) (+ s.e.) of Notolabrus fucicola at nine sites at 25 three depths, shallow (0-6 m), intermediate (7-11 m) and deep (12-18 m), around Kaikoura from Dec 1996 to Dec 1997 (65 transects per site). Figure 2.5 Mean size (cm) (+/- s.e.) of Notolabrus fucico/a per transect 26 2 (150 m ) around Kaikoura at A: Shallow sites (0-6 m), B: Intermediate sites (7-11 m) and C: Deep sites (12-18 m) from Dec 1996 to Dec 1997. List of Figures. v CHAPTER THREE Figure 3.1 Size frequency distribution of male (n = 172) and female (n = 50 281) Notolabrus fucicola samples collected around Kaikoura from Dec 1996 to Feb 1998. Figure 3.2 Size frequency distribution of initial (n = 224), transitional (n = 50 30) and terminal phase (n = 199) Notolabrus fucicola from samples collected around Kaikoura from Dec 1996 to Feb 1998. Figure 3.3 Mean monthly gonosomatic index(+/- s.e.) of male and female 53 Notolabrus fucicola sampled around Kaikoura from Dec 1996 to Feb 1998. Figure 3.4 Mean monthly gonosomatic index (+/- s.e.) of initial and 54 terminal phase male (A) and female (B) Notolabrus fucicola sampled around the Kaikoura Peninsula from Dec 1996 to Feb 1998. Figure 3.5 Scattergram showing the gonosomatic index versus standard 55 length (mm) of initial and terminal phase male and female Notolabrus fucicola over the spawning period, July to Dec 1997. Correlations given for each class. Figure 3.6 Standard length (mm) versus age (+/- s.e) of Notolabrus 65 fucicola using scales (n = 452) and otoliths (n = 31 ). Dotted lines are linear trendlines. When no error bars are present, the value is based on one or two fish. Figure 3.7 Relationship between standard length and age (+/- s.e) for 65 male and female Notolabrus fucico/a (from scales) (n = 452). Figure 3.8 Marginal increment analysis (mm) (+ s.e.) of 200 - 21 O mm 66 Notolabrus fucicola sampled from Dec 1996 to Feb 1998. List of Figures. vi CHAPTER FOUR Figure 4.1 Frequency of occurrence of prey items in the gut of all 84 Notolabrus fucicola pooled across the study period (Dec 1996 to Feb 1998). Figure 4.2 Mean percentage volume (+ s.e.) of nine major prey items 84 found in the gut of all Notolabrus fucicola sampled from Dec 1996 to Feb 1998. Figure 4.3 Percentage of occurrence of seven major prey categories 87 pooled across all seasons in all Notolabrus fucico/a divided into 1O mm size classes. Figure 4.4 Mean percentage volume (+/- s.e.) of seven major prey 88 · categories pooled across all seasons in all Notolabrus fucico/a divided into 1O mm size classes. Figure 4.5 Mean percentage volume (+ s.e.) of A: Hard prey and B: Soft 91 prey in the guts of Notolabrus fucicola sampled around Kaikoura from Dec 1996 to Feb 1998. Note the different y-axes. Figure 4.6 Mean fullness index of male and female Notolabrus fucicola 93 sampled around Kaikoura from Dec 1996 to Feb 1998 (+/- s.e.). Figure 4.7 Condition factor (K) (+ s.e.) for male and female Notolabrus 93 fucico/a sampled around Kaikoura from Dec 1996 to Feb 1998. List of Figures. vii CHAPTER FIVE Figure 5.1 Mean percentage of time (+ s.e.) spent on nine behaviour 103 categories for Notolabrus fucicola in four standard length classes observed from Sep 1997 to Feb 1998 (See Table 3.3 for behaviour category codes). List of Tables. viii LIST OF TABLES CHAPTER TWO Table 2.1 Summary of characteristics of the nine sites sampled over the 16 study (SW= seaweed). Table 2.2 Summary of species, their common name and code, observed 19 over the study period. Table 2.3 Analysis of variance on the effects of time, depth and sites on 22 the numbers of Notolabrus fucicola; depth nested in site, sites treated as random factors: data log(x+ 1) transformed but remained heterogeneous after transformation (Cochrans C = 0.028). A critical significance level of 0.01 was used. Table 2.4 Analysis of variance on the effects of time, depth and sites on 22 the numbers of Notolabrus celidotus; depth nested in site, sites treated as random factors: data log(x+ 1) transformed but remained heterogeneous after transformation. A critical significance level of 0.01 was used. Table 2.5 Analysis of variance on the effects of time, depth and sites on 23 the numbers of Odax pullus; depth nested in site, sites treated as random factors: data log(x+ 1) transformed to remove heterogeneity of variance (Cochrans C = 0.98). List of Tables. ·ix CHAPTER THREE Table 3.1 Summary of the stages of oocyte development for Notolabrus 45 fucicola. Table 3.2 Summary of the stages of Notolabrus fucicola spermatogenic 46 development. Table 3.3 Number and percentage (in brackets) of male and female 51 Notolabrus fucicola in each colour phase (initial, transitional and terminal) sampled around Kaikoura from Dec 1996 to Feb 1998 (n = 453). Table 3.4 Analysis of variance on the gonosomatic index (GSI) of male 52 and female Notolabrus fucicola over time and between sexes. CHAPTER FOUR Table 4.1 A-I Analysis of variance showing differences in prey items in 89 the gut of Notolabrus fucicola over.time. CHAPTER FIVE Table 5.1 Behaviour categories, codes and the mean percentage of time 102 (+/- s.e.) spent on each activity for all Notolabrus fucicola observed from September 1997 to February 1998 (n = 464). List of Plates. x LIST OF PLATES CHAPTER ONE Plate 1.1 Photo of an initial phase Notolabrus fucicola (reproduced from 6 Francis, 1988). Plate 1.2 Photo of a terminal phase Notolabrus fucicola (reproduced from 6 Francis, 1988). CHAPTER THREE. Plate 3.1 Post spawned female collected in January with characteristic 58 atretic oocytes (AC} and immature (pre-vitellogenic) oocytes (10). Note the nucleus (N) in the chromatin nucleolus oocytes and the trabeculae (TR). Scale bar is 0.1 mm. Plate 3.2 A resting ovary from a female collected in February with a 58 dominance of immature (pre-vitellogenic) oocytes (10) and a thickened gonad wall (GW). Note the nucleolus in the nucleus (N) and the trabeculae (TR). Scale bar is 0.1 mm. Plate 3.3 Female undergoing stage one vitellogenesis collected in July. 59 The oocytes have increased in size and lipid droplets (LP) have appeared. The germinal vesicle (GV) and the chorion have appeared. Note the gonad wall (GW), nucleus (N), chorion (CH) and immature oocytes (10). Scale bar is 0.05 mm.