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The Growth and Reproduction of Patella Granularis (Mollusca: Patellogastropoda) on the South- East Coast of South Africa

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The Growth and Reproduction of Patella Granularis (Mollusca: Patellogastropoda) on the South- East Coast of South Africa View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by South East Academic Libraries System (SEALS) THE GROWTH AND REPRODUCTION OF PATELLA GRANULARIS (MOLLUSCA: PATELLOGASTROPODA) ON THE SOUTH- EAST COAST OF SOUTH AFRICA A thesis submitted in fulfilment of the requirements for the degree of DOCTOR of PHILOSOPHY of Rhodes University by LAURA SUZANNE VAT January 2000 ii ABSTRACT Aspects of the biology and ecology of Patella granularis were investigated along a 130km stretch of the south-east coast of South Africa. Distribution, biomass, density and population structure were investigated at seven localities. In addition, a more detailed study of the growth rate and reproductive biology of populations inhabiting three different substrata (aeolianite, quartzitic sandstone and mussel shells) was conducted. The genetic relationships between these three populations was also examined, as was the foraging behaviour of the limpets inhabiting an aeolianite and a quartzitic sandstone shore. Finally, differences in food availability on the different substrata were studied. On the south-east coast, P. granularis has a wide intertidal distribution, occurring from the upper Balanoid zone through to the Cochlear zone, where it is a common inhabitant of mussel shells. The mean shell length of P. granularis was found to decrease down the shore. The largest limpets (46.6 mm shell length) were found on an offshore island in Algoa Bay. At most localities investigated, the sex ratio deviated from a 1:1 ratio with more males than females being recorded on five shores. Both limpet density and biomass were lower on the south-east coast when compared to data published for west coast populations. On the south-east coast, both density and dry biomass were highest in the lower Balanoid zone. Allozyme electrophoresis indicated that P. granularis inhabiting aeolianite, quartzitic sandstone and mussel shells are all part of a single population. Extremely high genetic identity values (0.998), low levels of heterozygosities (0.035 - 0.061), low levels of polymorphisms (25% - 31%) and low FST values (0.021) all suggest that the three populations of P. granularis form a common breeding group, despite the high levels of phenotypic plasticity observed. On all shores, P. granularis was found to grow allometrically, increasing in shell height more rapidly than shell length. Estimation of the growth rate (determined by the Von Bertalanffy growth model) of P. granularis suggested that limpets inhabiting the mussel shells grew more slowly, and attained a smaller maximum size, than those inhabiting both the aeolianite and the quartzite (K = 0.25, 0.32 and 0.33 respectively; 27.12 mm, 31.89 mm and 32.96 mm respectively). Previous work has shown that west coast P. granularis grow more quickly (K = 0.7) and reach a greater size (. 40 mm). Translocation of limpets among sites suggested that limpet size in the mussel beds was spatially constrained. Shell microgrowth bands were deposited tidally, but could not be used for aging limpets due to shell erosion. iii Limpets from the aeolianite had the greatest reproductive fitness, producing more eggs (.366 000/limpet) than those inhabiting quartzite (.119 500/limpet) or mussel shell limpets (.85 800/limpet). Aeolianite limpets also spawned throughout the year, whereas those from the quartzite and mussel shells spawned twice a year (once in winter and once in summer) although a great deal of interannual variability was observed. The onset of sexual maturity occurred at a similar age in all limpets (1 - 2 years) and is probably genetically entrenched. P. granularis inhabiting both an aeolianite and a quartzitic shore were active during nocturnal low tides. All limpets returned to a home scar after foraging. Whilst foraging, limpets inhabiting the aeolianite shore moved shorter distances (.17 cm) at a slower rate than those from quartzite (.30 cm). Limpets that were translocated from one substratum to the other initially moved similar distances to the source group, but after a maximum period of one week, moved distances that mirrored those moved by the resident limpets. Neither season nor tidal phase influenced the distances foraged. No directionality in foraging was found. Wear of radula teeth, particularly the pluricuspid tooth, was greater in limpets from the quartzite. It is hypothesised that the observed differences in life-history parameters and foraging behaviour of limpets both within the south-east coast and between the west and south-east coasts are related to food abundance. Chlorophyll-a, and hence microalgal biomass, was consistently higher on aeolianite (.2.5 times) than on both quartzite and mussel shells. Estimates of chlorophyll-a were higher (although not significantly) in winter. Previous studies determined that primary productivity is also higher along the west coast. Finally, the lack of evidence for migration of limpets from the low-shore to high-shore in south-east coast P. granularis is discussed. It is suggested that this species settles opportunistically within its physiological tolerances and responds morphologically to localised environmental conditions. iv ACKNOWLEDGEMENTS I am deeply indebted to my supervisor, Professor A.N. Hodgson, who has guided and encouraged my academic career since my undergraduate studies. He must also be thanked for his abundant patience and support throughout the preparation of this thesis, as well as his financial assistance throughout the course of this project. Thanks must also be extended to my parents, John and Phyllis Moxham, who always supported my interest in things biological. They are also thanked for their comments on earlier drafts of this work. I am extremely grateful to the many friends without whom this project would not have been possible. They enthusiastically provided ideas for, and comments on, the work included in this thesis. In addition, they were often called on to provide “extra hands” on field trips that inevitably took place in less than ideal conditions. Special mention must thus be made of Drs. Dave Gray, Greg Foster and Sven Kaehler, Janine Gray, Carol Simon, Greg Tinney, Craig Whittington-Jones, Lynn Randell, Claire Jackson, Leigh Gurney and Jennifer Gush. Extra thanks go to Mark Robertson and Tammy Smith, who not only provided accommodation for overnight field trips, but were frequently to be found on the rocky shores at night observing limpets. I also wish to thank Dr. Sarah Radloff (Dept. Statistics) and Dr. Martin Villet for their help with statistical analyses of the data; the staff of the Electron Microscopy Unit of Rhodes University, particularly Rob Cross and Shirley Pinchuck for their help with related techniques and procedures; Dr. Barbara Stewart (and the students working under her at the electrophoresis laboratory at the South African Museum) firstly, for allowing me to invade the laboratory and secondly, for patiently explaining allozyme electrophoresis to me. Finally, I am deeply indebted to my husband, Nick, who from the beginning of this journey of discovery, has provided immeasurable help and support. He is especially thanked for his constantly enthusiastic company on many field trips, his extensive help with computer related problems, his support during the frustrating times and his continued faith in me. He must also be thanked for providing constructive comments on earlier drafts of this thesis and for spending much time formatting and printing the final document. The financial assistance of the National Research Foundation (NRF, South Africa) and the Rhodes University Graduate Assistance programme is hereby acknowledged. Opinions expressed and conclusions arrived at, are those of the author and are not necessarily to be attributed to the National Research Foundation. v TABLE OF CONTENTS Chapter 1 GENERAL INTRODUCTION ............................................... Page 1 Chapter 2 POPULATION STRUCTURE, BIOMASS AND DENSITY OF PATELLA GRANULARIS ON THE SOUTH-EAST COAST OF SOUTH AFRICA ................................... Page 8 2.1: INTRODUCTION: ........................................................ Page 9 2.2: MATERIALS AND METHODS: ............................................. Page 11 2.2.1: Study localities: ............................................... Page 11 2.2.2: Sampling procedure: ........................................... Page 11 2.2.3: Statistical analyses: ............................................ Page 12 2.3: RESULTS: ............................................................ Page 16 2.3.1: Distribution: ................................................. Page 16 2.3.2: Density: .................................................... Page 18 2.3.3: Biomass: .................................................... Page 22 2.3.4: Population structure: ........................................... Page 25 2.3.5: Sex ratios and size at first sexual maturity: .......................... Page 39 2.4: DISCUSSION: ......................................................... Page 43 Chapter 3 ALLOZYME ELECTROPHORESIS OF THREE POPULATIONS OF PATELLA GRANULARIS FROM THE SOUTH-EAST COAST OF SOUTH AFRICA ........................ Page 51 3.1: INTRODUCTION: ....................................................... Page 52 3.2: MATERIALS AND METHODS: ............................................. Page 54 3.2.1: Collection of samples: .......................................... Page 54 3.2.2: Electrophoresis: .............................................. Page 54 3.2.3: Data analysis: ...............................................
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