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The Taxonomy and Phylogeny of Three Gnathiid Isopod Species Parasitising Elasmobranchs from the Great Barrier Reef, Australia

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The Taxonomy and Phylogeny of Three Gnathiid Isopod Species Parasitising Elasmobranchs from the Great Barrier Reef, Australia THE TAXONOMY AND PHYLOGENY OF THREE GNATHIID ISOPOD SPECIES PARASITISING ELASMOBRANCHS FROM THE GREAT BARRIER REEF, AUSTRALIA. BY MARYKE LOUISE COETZEE DISSERTATION SUBMITTED IN FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE MAGISTER SCIENTIAE IN ZOOLOGY IN THE FACULTY OF SCIENCE AT THE UNIVERSITY OF JOHANNESBURG SUPERVISOR: DR. N. J. SMIT MAY 2006 TABLE OF CONTENTS 1. LIST OF TABLES. 1 2. LIST OF FIGURES. 2 3. SUMMARY. 8 4. OPSOMMING. 10 5. CHAPTER 1: GENERAL INTRODUCTION. 12 1.1. History of gnathiid research 14 1.2. Basic gnathiid morphology 16 1.3. Gnathiid life cycles 18 1.4. Gnathiid distribution 21 1.5. Parasitic gnathiid larvae 24 1.6. Gnathiid larvae as potential vectors 25 1.7. Gnathiids parasitising elasmobranchs 25 1.8. Project objectives 26 6. CHAPTER 2: TAXONOMIC DESCRIPTIONS OF THREE NEW GNATHIID SPECIES FROM THE GREAT BARRIER REEF. 27 2.1. Introduction 28 2.2. Materials and Methods 28 2.3. Results 30 2.4. Taxonomic Descriptions 35 2.4.1. Gnathia trimaculata sp. n. 35 2.4.2. Gnathia nigrograndilaris sp. n. 58 2.4.3. Gnathia australis sp. n. 75 TABLE OF CONTENTS 7. CHAPTER 3: PHYLOGENETIC ANALYSIS AND GEOGRAPHIC DISTRIBUTION OF THE FAMILY GNATHIIDAE. 96 3.1. Introduction 97 3.2. Phylogenetic Methods 102 3.3. Results and Discussion 105 3.3.1. Family Gnathiidae 106 3.3.2. Genus Gnathia 107 3.3.3. Geographical Distribution 107 8. CHAPTER 4: MULTIVARIATE STATISTICS. 117 4.1. Introduction 118 4.2. Methods 118 4.3. Results 118 4.4. Discussion 119 9. CHAPTER 5: GENERAL DISCUSSION, RECOMMENDATIONS AND SCOPE FOR FUTURE RESEARCH. 124 10. ACKNOWLEDGEMENTS. 129 11. REFERENCES. 130 List of Tables 1 List of Tables Table 2.1. Families and species of elasmobranchs collected from various localities and examined for gnathiid isopod larvae. Abbreviations: N – number, ML – mean length, SD – standard deviation, P – prevalence, MB – mean burden. Table 3.1. Morphological characters of adult males of the eleven genera of the family Gnathiidae Harger, 1810. Table 3.2. Character transformations used in the Phylogenetic analysis of the family Gnathiidae according to Cohen and Poore (1994). The plesiomorphic state is listed first and separated by a semi-colon from the apomorphic states. Table 3.3. Species-character matrix of species of the family Gnathiidae, with outgroups Cirolanidae and Protognathidae. Unknown characters are stated as "?". Plesiomorphic states are indicated by a 0 and apomorphic states by a 1 or more in the data matrix. List of Figures 2 List of Figures Figure 1.1. Line drawing of a male gnathiid (dorsal view) showing main anatonomical features (Redrawn from Cohen and Poore, 1994). Abbreviations: P – pereonites. Figure 1.2. Line drawing of a female gnathiid (dorsal view) showing main anatonomical features. Figure 1.3. Line drawing of gnathiid praniza larva (dorsal view) showing main anatonomical features. Figure 1.4. Schematic representation of the life cycle of a gnathiid (Gnathia africana Barnard, 1914). Redrawn from Smit et al., (2003). Abbreviations: P – praniza stage, Z – zuphea stage. Figure 1.5. Representation of the eight marine zoogeographical regions where gnathiids are commonly found according to Wye,(1991). Figure 2A. Data collection field trips undertaken to Lizard and Heron Island Research Stations Fig. 2A.1. Lizard Island notice board stating that it is a facility of the Australian Museum. Fig. 2A.2. Aerial photograph of Lizard Island. Fig. 2A.3. Boats used for shark collection. Fig. 2A.4. Working in the microscope room.Fig. 2A.5. Aerial photographs of Heron Island. Fig. 2A.6. Heron Island Research Station. Figure 2B. Maps representing sampling localities. Fig. 2B.1. Heron Island (Website 1)*. Fig. 2B.2. Shark Bay represented by C1 (Website 2)*. Fig. 2B.3. Moreton Bay (Website 3)*. Fig. 2B.4. Lizard Island (Website 4)*. * See Websites 1-4 in reference list, section websites Figure 2C. Line drawings of the elasmobranchs hosts collected at Lizard Island, Heron Island (Shark Bay) and Moreton Bay. Fig. 2C.1. Carcharinus amblyrhynchos Bleeker, 1856. List of Figures 3 Fig. 2B.2. Triaenodon obesus Rüppell, 1837. Fig. 2C.3. Urogymnus asperrimus Bloch & Schneider, 1801. Fig. 2C.4. Himantura fai Jordan & Seale, 1906. Fig. 2C.5. Pastinachus sephen Forsskål, 1775. Fig. 2C.6. Taeniura lymma Forsskål, 1775. Fig. 2C.7. Rhynchobatus djiddensis Forsskål, 1775. Redrawn from Smith and Heemstra (2003) and Website 5 (See Website 5 in Reference list, section websites). Figures 2.1 – 2.7. Gnathia trimaculata sp. n., male. Fig. 2.1. Full length dorsal view. Fig. 2.2. Frontal border and mandibles. Fig. 2.3. Lateral view of cephalosome. Fig. 2.4. First antenna. Fig. 2.5. Second antenna. Fig. 2.6. Left pleopod 2 with appendix masculina. Fig. 2.7. Pleotelson and uropods. Scale bars: Figs. 2.4 – 2.7 = 100µm; Figs. 2.2, 2.3 = 500µm; Fig. 2.1 = 1mm. Figures 2.8 – 2.10. Cephalosome appendages of a male Gnathia trimaculata sp. n. Fig. 2.8. Pylopod. Fig. 2.9. Article 2 and 3 of pylopod. Fig. 2.10. Maxilliped. Scale bars = 100µm. Figure 2.11. Pereopods 2 to 6 (P2-P6) of a male Gnathia trimaculata sp. n. Scale bar = 200µm. Figures 2.12 – 2.17. Gnathia trimaculata sp. n., female. Fig. 2.12. Full length dorsal view. Fig. 2.13. Cephalosome and frontal border. Fig. 2.14. First antenna. Fig. 2.15. Second antenna. Fig. 2.16. Left pleopod 2. Fig. 2.17. Pleotelson and uropods. Scale bars: Fig. 2.13 = 10µm; Figs. 2.14 – 2.17 = 100µm; Fig. 2.12 = 1mm. Figures 2.18, 2.19. Cephalosome appendages of a female Gnathia trimaculata sp. n. Fig. 2.18. Left pylopod. Fig. 2.19. Right maxilliped. Scale bar = 100µm. Figure 2.20. Pereopods 2 to 6 (P2-P6) of a female Gnathia trimaculata sp. n. Scale bar = 200µm. Figures 2.21 – 2.26. Gnathia trimaculata sp. n., praniza larva. Fig. 2.21. Full length dorsal view. Fig. 2.22. Dorsal cephalosome with labrum and antennae. Fig. 2.23. First antenna. Fig. 2.24. Second antenna. Fig. 2.25. Right pleopod 2. Fig. 2.26. Pleotelson and uropods. Scale bars: Figs. 2.23, 2.24 = 100µm; Figs. 2.25, 2.26 = 200µm; Fig. 2.22 = 500µm; Fig. 2.21 = 1mm. List of Figures 4 Figures 2.27 – 2.31. Cephalosme appendages of a praniza larva of a Gnathia trimaculata sp. n. Fig. 2.27. Gnathopod. Fig. 2.28. Mandible. Fig. 2.29. Maxilliped. Fig. 2.30. Maxillule. Fig. 2. 31. Paragnath. Scale bars: Fig. 2.27 = 10µm; Figs. 2.28 - 2.31 = 100µm. Figure 2.32. Pereopods 2 to 6 (P2-P6) of a praniza larva of a Gnathia trimaculata sp. n. Scale bar = 200µm. Figures 2.33 – 2.41. Figures 2.33 – 2.41. Scanning electron micrographs of Gnathia trimaculata sp. n., male (Figs. 2.33 – 2.36), female (Figs. 2.37, 2.38) and larva (Figs. 2.39 – 2.41). Fig. 2.33. Lateral view of male cephalosome. Fig. 2.34. Ventral view of male frontal border. Fig. 2.35. Lateral view of male pleon. Fig. 2.36. Dorsal view of male pleotelson and uropods. Fig. 2.37. Dorsal view of female cephalosome. Fig. 2.38. Ventral view of female cephalosome. Fig. 2.39. Dorsal view of praniza larva cephalosome. Fig. 2.40. Ventral view of praniza larva cephalosome. Fig. 2.41. Dorsal view of praniza larva pleotelson and uropods. Scale bars: Fig. 2.34 = 50µm; Figs. 2.35 – 2.41 = 100µm; Fig. 2.33 = 200µm. Figures 2.42 – 2.45. Figures 2.42 – 2.45. Light microscope photographs of Gnathia trimaculata sp. n., male (Fig. 2.42), female (Fig. 2.43) and larvae (Figs. 2.44, 2.45). Fig. 2.42. Dorsal view of male. Fig. 2.43. Dorsal view of female. Fig. 2.44. Dorsal view of praniza larvae. Fig. 2.45. Dorso-lateral view of praniza larvae. Scale bars = 1mm. Figures 2.46 – 2.52. Gnathia nigrograndilaris sp. n., male. Fig. 2.46. Full length dorsal view. Fig. 2.47. Second antenna. Fig. 2.48. Frontal border and mandibles. Fig. 2.49. Lateral view of cephalosome. Fig. 2.50. Pleotelson and uropods. Fig. 2.51. First antenna. Fig. 2.52. Left pleopod 2 with appendix masculina. Scale bars: Figs. 2.48, 2.50 - 2.52 = 100μm; Figs. 2.47, 2.49. = 500μm; Fig. 2.46 = 2mm. Figures 2.53 – 2.55. Cephalosome appendages of a male Gnathia nigrograndilaris sp. n. Fig. 2.53. Pylopod. Fig. 2.54. Article 2 and 3 of pylopod. Fig. 2.55. Maxilliped. Scale bars = 100μm. Figure 2.56. Pereopods 2 to 6 (P2 - P6) of a male Gnathia nigrograndilaris sp. n. Scale bar = 200μm. List of Figures 5 Figures 2.57 – 2.62. Gnathia nigrograndilaris sp. n., praniza larva. Fig. 2.57. Full length dorsal view. Fig. 2.58. Dorsal cephalosome with labrum and antennae. Fig. 2.59. Second antenna. Fig. 2.60. First antenna. Fig. 2.61. Left pleopod 2. Fig. 2.62. Pleotelson and uropods. Scale bars: Figs. 2.59 – 2.62 = 100μm; Fig. 2.58 = 500μm; Fig. 2.57 = 2mm. Figures 2.63 – 2.67. Cephalosome appendages of a praniza larva of a Gnathia nigrograndilaris sp. n. Fig. 2.63. Maxilliped. Fig. 2.64. Gnathopod. Fig. 2.65. Mandible. Fig. 2.66. Maxillule. Fig. 2.67. Paragnath. Scale bars: Fig. 2.64 = 10µm; Figs. 2.63, 2.65 – 2.67 = 100µm. Figure 2.68. Pereopods 2 to 6 (P2 - P6) of a praniza larvae of a Gnathia nigrograndilaris sp. n. Scale bar = 200μm. Figures 2.69 – 2.77. Scanning electron micrographs of Gnathia nigrograndilaris sp. n., male (Figs. 2.69 – 2.73) and larva (Figs. 2.74 – 2.77).
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