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Download The FACTORS AFFECTING THE DISTRIBUTION AND ABUNDANCE OF TWO SPECIES OF BEACH CRAB, HEMIGRAPSUS OREGONENSIS AND HEMIGRAPSUS NUDUS by CHARLES JAMES LOW B.S.A., University of Guelph, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of ZOOLOGY V/e accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA November, 1970 In presenting this thesis in partial fu1filment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia Vancouver 8, Canada Date /- 7c ii ABSTRACT Hemigrapsus oregonensis and Hemigrapsus nudus, two beach crabs common on the Pacific Coast of North America, show con• siderable variation in numbers, and species dominance from place to place. To determine why this should be so, a number of experiments were performed., and observations made to de• termine what are the morphological, and physiological differ• ences between the two species, and how the crabs would be affected by the different physical conditions prevailing in different places. In general, it appeared that H. nudus could not tolerate muddiness of the substrate, while H. oregonensis tended to be eliminated from clean areas by predators. The abundance of the crabs appears to be correlated with the amount of cover available. iii TABLE OF CONTENTS PAGE ABSTRACT ii LIST OF TABLES iv LIST OF FIGURES vii ACKNOWLEDGEMENTS viii INTRODUCTION 1 NATURAL HISTORY 4 BEHAVIOR 7 EXPERIMENTAL RESULTS 10 OBSERVATIONS ON DISTRIBUTION 11 EXPERIMENTS 13 FACTORS AFFECTION DISTRIBUTION: OXYGEN 13 FACTORS AFFECTING DISTRIBUTION: MUD 17 FACTORS AFFECTING DISTRIBUTION: SETTLEMENT ... .21 FACTORS AFFECTING DISTRIBUTION: COMPETITION FOR COVER 22 FACTORS AFFECTING DISTRIBUTION: PREDATION. .24 FACTORS AFFECTING DISTRIBUTION: SALINITY 26 FACTORS AFFECTING DISTRIBUTION: DESICCATION. .26 FACTORS CONTROLLING ABUNDANCE 28 DISCUSSION 30 CONCLUSION 34 BIBLIOGRAPHY 70 iv LIST OF TABLES TABLE PAGE I Reducing power of substrate samples, classified by the species of crab which dominated in the census sample 35 II Counts of the number of crabs reaching the surface of the water per hour. 37 III Counts of the number of crabs reaching the surface of the water per hour 38 IV Number of each species dead, presumably of suffocation, relative to their choice to remain under cover, or to respond to a horizontal oxygen gradient 39 V Time to death of each member of pairs of crabs, matched for size, in water where most of the oxygen was removed by bubbling nitrogen, in flasks which were corked to exclude air after the addition of the crabs. Al VI (A) Comparison of survival time within matched pairs of H. oregonensis and H. nudus in corked one liter flasks, without mud 42 (B) Comparison of survival time of matched pairs of H. oregonensis and H. nudus in corked one liter flasks, with mud A3 (C) Comparison of survival times of H. nudus, with and without mud, in corked one liter flasks kh V TABLE PAGE VI (D) Comparison of survival times of H. oregonensis in corked one liter flasks with and without mud 45 VII Survival of crabs caged in the field, in areas with different substrate particle size distributions ..... ....46 VIII Total number of crabs on different substrates, counted 13 times for 7 days 48 IX Proportion of all crabs on beaches made up of those with a carapace width less than 7 mm., relative to the dominance of the beach by the adults of the two species .49 X (A) Number of crabs out from under limited cover. 200 H. oregonensis present 50 (B) Number of crabs out from under cover. 200 H. nudus present 50 (C) Number of crabs out from under cover. 100 of each species present 51 (D) Comparison of number of crabs out from under cover by species 51 XI Crab taken first when pairs of crabs, not matched for size, were thrown into a tank with an assortment of crab-eating fish 53 XII Crab predators. Species of birds and fish examined for crabs eaten 54 vi TABLE PAGE XIII Tolerance of the two species for low salinity 57 XIV Time to death of the two species in fresh (tap) water. 58 XV Time to death, in hours, of the two species when kept in dry dishpans 59 XVI Regression on crab census data, with the number of crabs as the dependent variable, and as independent variables; the reducing power of the substrate (0), the percent cover (R), the surf (S), the available food (F), the height of the sample on the beach (H), and the salinity (N) 60 XVII Comparison of the number of crabs found in plots 1, 2 and 3, before and after rocks were removed from plot 1 and added to plot 2. .... .61 vii LIST OF FIGURES FIGURE PAGE 1 Graph of % H. nudus against reducing power of the substrate 62 2 Change in distribution of H. nudus and H. oregonensis over time at different oxygen concentrations 63 3 Change in distribution totals of the two species with oxygen tension 65 A Size distribution of H. nudus from San Juan Island, June, 1969 66 5 Size distribution of H. oregonensis from Spanish Banks, February, 1968 ... .67 6 Number of crabs left alive after addition of 30 of each species to a tank with predatory fish and cover 68 MAP 69 viii ACKNOWLEDGEMENTS I wish to thank Dr. J. R. E. Harger for his advice, criticism and patience, without which this work could not have been carried out. I also would like to express my appreciation for the interest shown in this work by Miss 0. Johannson, Miss S. Behrens, Mr. P. Breen and Mr. J. Stimpson. This work was supported in part by a National Research Council bursary, and by a University of British Columbia fellowship. 1 INTRODUCTION Hemigrapsus nudus and Hemigrapsus oregonensis are small intertidal crabs which are widely distributed on the Pacific North American Coast and occur, usually together, on beaches in the Puget Sound-Strait of Georgia area. The range of H. nudus. according to Schmidt (1921), is from Sitka, Alaska, to the Gulf of California. H. oregonensis ranges from Prince Wil• liam Sound, Alaska, to the Gulf of California. These crabs are found in abundance on beaches covered with loose rock ranging in size from cobbles to boulders. H. nudus is found alone, in small numbers, on solid rock faces, where there are crevices for the crabs to shelter in. H. oregonensis is found alone on sheltered mud and sand flats, where the sub• strate contains enough fine material to allow the crabs to dig holes. In both habitats, the crabs are found in low numbers, usually on the order of 0.1 crabs per square meter. On beaches with loose rock, there can be as many as 500 crabs per square meter. Beaches where H. nudus is dominant tend to have little or• ganic and fine material in the substrate, while where H. ore• gonensis is the more numerous the beach is usually muddy, with much organic material. The water above these muddy beaches is usually quite turbid, even though there is usually little surf. Above the clean, sand and shell beaches where H., nudus is domi• nant, the water is usually clear, even though there may be considerable surf. The muddy beaches dominated by H. oregonensis 2 tend to have a lot of reduced material in the substrate. Under the surface, the muck is black, and usually smells strongly of hydrogen sulphide. Where H. nudus is the more common crab, as a rule there is less black material, and that is usually a lot deeper than is the case on H. oregonensis dominated beaches. Dominance of a beach by one or the other of the two species is unmistakeable. It is usual for one of the two species to make up 80% or more of all the crabs present. However, in some instances it is possible that one species will be replaced by the other in a very short distance. Both species are found almost exclusively between high and low tide levels. Either species can occupy the whole inter• tidal, but where they are found together, the center of abun• dance of H. nudus will be above that of H. oregonensis. Where H. nudus is dominant, H. oregonensis will be confined to a narrow strip along the bottom of the beach, while where H. ore• gonensis is dominant, H. nudus will be confined to a narrow strip along the top of the beach. In the narrow strips to which the subordinate species is confined there are almost al• ways many of the dominant species as well. In these areas where a few of the rarer species exist, at the top, or bottom of the beach, as the case may be, the number of crabs per unit area is much less than in the center of the beach, where the dominant crab will maintain a high population density. Thus, while both species are capable of living from the top to the bottom of the intertidal, they both prefer, in the sense that the greatest populations are maintained, to live 3 near the center of the intertidal, on beaches with a good cover ing of loose rock. This study was undertaken to attempt to understand the reasons for the observed distributions and varying abundances shown by the crabs.
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