https://theses.gla.ac.uk/ Theses Digitisation: https://www.gla.ac.uk/myglasgow/research/enlighten/theses/digitisation/ This is a digitised version of the original print thesis. Copyright and moral rights for this work are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This work cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Enlighten: Theses https://theses.gla.ac.uk/ [email protected] ASPECTS OF THE BIOLOGY OF THE SQUAT LOBSTER, MUNIDA RUGOSA (FABRICIUS, 1775). Khadija Abdulla Yousuf Zainal, BSc. (Cairo). A thesis submitted for the degree of Doctor of Philosophy to the Faculty of Science at the University of Glasgow. August 1990 Department of Zoology, University of Glasgow, Glasgow, G12 8QQ. University Marine Biological Station, Millport, Isle of Cumbrae, Scotland KA28 OEG. ProQuest Number: 11007559 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 11007559 Published by ProQuest LLC(2018). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 To Ahmed, Tarneem and Vinous DECLARATION. I hereby declare that this thesis represents, except where a note is made to the contrary, work carried out by myself. It has not been previously submitted for any degree. Khadiia A Y. Zainal August 1990 ACKNOWLEDGMENTS This work was carried out at the Zoology department, University of Glasgow and at University Marine Biological station, Millport, Isle of Cumbrae. The project was financed by a grant from Qaboos University, Sultanate of Oman- Muscat. I wish to thank my supervisors, Dr Alan Taylor and Dr Jim Atkinson for their invaluable advice, constructive criticism and generous guidance throughout the project. I gratefully acknowledge the facilities provided by Professor R.S. Phillips and the members of the Zoology department and by Professor J.A. Allen and members of the University Marine Biological Station at Millport. Many thanks to the crews of the research vessels for obtaining experimental animals; to Dr Mark Davies for catching small size Munida rugosa, and to Dr A.C. Rice (Institute of Oceanographic Sciences) for his help in identifying Munida sarsi. I am grateful to Cathy McLagan and June Freel for their help in technical laboratory work; to Liz Denton for her kind assistance in the use of the Apple Macintosh plus Computer and to Dave Henning for his help with the graphs. I thank Phil Smith for his help with Minitab for statistical analyses, Dr Chrachri, Coneilia Leibrock and Mett Thomsen for translating French, German and Danish references. Thanks to P. Rickus for his help with the photography, and to the SEMU for their help in specimen preparations for scanning electron microscopy. Many thanks go to my husband Hussein Al-Mosawi for his friendship, support, and help throughout. I am also grateful to my mother, who established my early interest in marine animals, and to my sister Fawzia Zainal for her encouragement. I am indebted to the members of the YWCA (Young Women Christian Association) who helped with the care of my children. Finally, I am especially grateful to my children Ahmed, Tarneem, and Vinous who gave moral support and made the past few years a happy time. II TABLE OF CONTENTS: Section Page No. Declaration I Acknowledgments II Contents III Abstract VI CHAPTER 1 . INTRODUCTION General context 1.1 1 Galatheid biology 1.2 2 Galatheid fisheries 1.3 4 The species studied 1.4 6 Aim and structure of thesis 1.5 9 CHAPTER 2. ECOLOGY, RELATIVE GROWTH AND FEEDING BIOLOGY INTRODUCTION 2.1 11 MATERIALS AND METHODS 2.2 14 Ecology 2.2.1 14 Relative growth; heterochely 2.2.2 14 Aspects of reproduction 2.2.3 16 Diet 2.2.4 17 Feeding behaviour 2.2.5 18 Functional morphology of the mouth parts and stomach 2.2.6 19 Munida sarsi 2.2.7 20 RESULTS 2.3 20 Ecology 2.3.1 20 Relative growth; heterochely 2.3.2 21 Aspects of reproduction 2.3.3 25 Diet 2.3.4 26 Feeding behaviour 2.3.5 26 Morphology of the mouth parts and pereiopods 2.3.6 31 Morphology of the stomach 2.3.7 36 Munida sarsi 2.3.8 39 DISCUSSION 2.4 42 Ecology 2.4.1 42 Relative growth; heterochely 2.4.2 43 Aspects of reproduction 2.4.3 47 Dietary analysis 2.4.4 50 Functional morphology of the mouth parts and chelipeds 2.4.5 51 Functional morphology of the stomach 2.4.6 54 III CHAPTER a. BRANCHIAL MORPHOLOGY, VENTILATION AND CARDIAC ACTIVITY INTRODUCTION 3.1 60 MATERIALS AND METHODS 3.2 64 Preparation of gills for light microscopy 3.2.1 64 Preparation of gills for scanning electron microscopy 3.2.2 65 Preparation of gills for transmission electron microscopy 3.2.3 65 Gill area measurements 3.2.4 66 Branchial ventilation and scaphognathite activity 3.2.5 66 Recordings of ventilatory activity under normoxic conditions 3.2.6 68 Effects of disturbance on ventilatory and cardiac activity 3.2.7 68 RESULTS 3.3 69 Branchial morphology 3.3.1 69 Thickness of the lamellar cuticle 3.3.2 71 Gill area measurements 3.3.3 72 Ventilation 3.3.4 73 The effect of disturbance on cardiac and ventilatory activity 3.3.5 76 Body weight relationships for heart rate and scaphognathite rate 3.3.6 76 DISCUSSION 3.4 77 Branchial chamber morphology 3.4.1 77 Ventilation and cardiac activity 3.4.2 82 CHAPTER 4. RESPIRATION INTRODUCTION 4.1 88 MATERIALS AND METHODS 4.2 89 Oxygen consumption rate under normoxic conditions 4.2.1 90 Effect of temperature on rates of oxygen consumption 4.2.2 92 The effect of feeding on rates of oxygen consumption 4.2.3 93 Effects of hypoxia on oxygen consumption rate 4.2.4 94 Survival experiments under anoxic condition 4.2.5 94 RESULTS 4.3 96 The relationship between M 02 and body weight . 4.3.1 97 Effect of temperature on M 02 IV heart rate and scaphognathite rate 4.3.2 99 Effects of feeding on metabolic rate 4.3.3 101 Effect of hypoxia on the rates of oxygen consumption, and on heart and scaphognathite rate 4.3.4 104 Anoxia tolerance 4.3.5 103 Oxyygen consumption, heart and scaphognathite rate during recovery 4.3.6 103 DISCUSSION . 4.4 104 Relationship between M 02 and body size . 4.4.1 105 Effect 01 temperature on Mo? 4.4.2 107 Effect of feeding on M 02 4.4.3 108 Effect of declimng oxygen tension 4.4.4 110 Oxygen debt 4.4.5 113 CHAPTER 5. RESPIRATORY PROPERTIES OF THE BLOOD INTRODUCTION 5.1 116 MATERIALS AND METHODS 5.2 117 Ionic composition of the blood 5.2.1 117 Lactate concentration of the blood 5.2.2 118 In vivo P02 and pH of the blood 5.2.3 118 Oxygen carrying capacity of the blood 5.2.4 119 Oxygen affinity of the haemocyanin 5.2.5 122 Carbon dioxide equilibrium curves 5.2.6 123 RESULTS 5.3 126 Ionic composition of the blood 5.3.1 126 P02, pH, and oxygen carrying capacity of the blood 5.3.2 126 O9 dissociation curves 5.33 127 CO2 equilibrium curves 5.3.4 129 DISCUSSION 5.4 130 Ionic composition of the blood 5.4.1 130 Oxygen carrying capacity of the blood 5.4.2 131 In vivo blood pH and P 02 5.4.3 133 O2 affinity of the blood 5.4.4 134 CO2 transport and acid-base balance 5.4.5 139 CHAPTER £. GENERAL DISCUSSION 142 REFERENCES 147 V ABSTRACT. The thesis presents a comparative study of respiratory physiology of the galatheids Munida rugosa (Fabricius, 1775) and M. sarsi Huus, 1935. Both species were obtained from the Firth of Clyde, M. rugosa from a depth range of 8-115m and M. sarsi from 95-115m depth. To provide background information for this study, some aspects of the general biology of both species were studied, with greatest emphasis given to M. rugosa. Both species were collected mainly from sandy muds using creels and trawls. Studies of relative growth were carried out and indicated that sexual differences were particularly apparent in cheliped lengths (positive allometry in males) and abdomen widths (positive allometry in females). Heterochely was observed in both species and, although there was much variation, it was more commonly seen in large males. Extrapolation of relative growth regression relationships for males and females suggested that, in M. rugosa, sexual maturity occurs at approximately 17mm carapace length and at approximately 10mm carapace length in M sarsi. Preliminary studies on the reproductive cycle of M. rugosa indicated that ovary development occurred between spring and autumn.