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Heterospecific Aggression and Dispersion in Two Species

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Heterospecific Aggression and Dispersion in Two Species HETEROSPECIFIC AGGRESSION AND DISPERSION IN TWO SPECIES OF SEA ANEMONES IN THE FLORIDA KEYS by Edward Mark Barham A Thesis Submitted to the Faculty of the College of Science in Partial Fulfillment of the Requirements for the Degree of Master of Science Florida Atlantic University Boca Raton, Florida August 1991 HETEROSPECIFIC AGGRESSION AND DISPERSION IN TWO SPECIES OF SEA ANEMONES IN THE FLORIDA KEYS by Edward Mark Barham This thesis was prepared under the direction of the candidate's thesis advisor, Dr. Godfrey R. Bourne, Department of Biological Sciences, and has been approved by the members of his supervisory committee. It was submitted to the faculty of the College of Science and was accepted in partial fulfillment of the requirements for the degree of Master of Science. SUPERVISORY COMMITTEE: K23~ . Bourne, Chairman ~~-~~~aiPh rcJfiaffiS 2/£'~ -z;u~:;__:_ Dr. Sheldon Dobkin Dr. Michael S Sheila Mahoney Qflte I Acting Dean, Graduate ii ACKNOWLEDGEMENTS I sincerely thank the members of my thesis committee - Dr. Ralph M. Adams and Dr. Sheldon Dobkin for their critical reading of the manuscript. I acknowledge Dr. w. R. Brooks for helping me get started. I especially thank Dr. Godfrey R. Bourne for assuming the role of thesis chair at a critical time. He was also generous with access to his laboratory, computer, and most importantly with time he spent sharpening my research focus, and reviewing and improving my thesis. I also thank Eileen Garcia for help in slide preparation; Cathy Owen for critically reading a draft, and Ann Broadwell, JoAnne Hansen, MaryBeth Mihalik and Jeff Rathgeb for their assistance in the field. Two Marine Biology Research Grants through the Department of Biological Sciences, and Dr. Dobkin's financial assistance during my last year at FAU made this thesis possible. Finally, I am grateful to my parents for their love, patience and support. iii ABSTRACT Author: Edward Mark Barham Title: Heterospecific aggression and dispersion in two species of sea anemones in the Florida Keys. Institution: Florida Atlantic University Advisor: Dr. Godfrey R. Bourne Degree: Master of Science Year: 1991 The relationship between heterospecific aggression and dispersion in sea anemones is poorly understood. This relationship was elucidated for Bartholomea annulata Leseur and Aiptasia pallida Verrill at a quarry in the Florida Keys. Laboratory experiments indicated that B. annulata was the aggressor. Individuals of both species moved to avoid contact under both laboratory and field conditions. Field assessment of dispersion revealed aggregated patterns, as well as vertical segregation between the species. Bartholomea annulata was most abundant at a depth of 0.75 m, while ~- pallida dominated at the surface. Heterospecific aggression may be just one but probably an important one of several factors mediating the spatial distribution of these two sea anemones. iv TABLE OF CONTENTS ACKNOWLEDGEMENTS . • . • • . iii ABSTRACT . .................•..........................•...... i v LIST OF TABLES . ............................................. vi LIST OF FIGURES . ........................................... vii INTRODUCTION . .. 1 MATERIALS AND METHODS ...•.................................... 4 Natural history of study animals ......................... 4 Study site ............................................... 5 Collection and maintenance of study animals .............. 7 Laboratory experiments on aggressive interactions ........ 7 Laboratory experiments on spatial dispersion .....••••.... 9 Field observations of spatial dispersion ................ 10 Field transplant experiments ............................ 11 RESULTS . .................................................... 12 Laboratory aggressive interactions .......•.............. 12 Laboratory dispersion patterns ...•......•.....•••....... 12 Spatial distribution of anemones in the field ........... 15 Aggressive interactions in the field ...•••••••••.••••..• 26 DISCUSS ION . ................................................. 2 9 Aggression . ............................................. 29 Aggression and dispersion patterns ...................... 31 Aggression and vertical zonation ........................ 33 LITERATURE CITED ............................................ 37 v LIST OF TABLES I. Nearest-neighbor analysis of dispersion for sympatric ~· annulata and ~· pallida in an experimental aquarium .......................... 13 II. Nearest-neighbor analysis of dispersion for ~· annulata and ~· pallida in separate experimentalaquariums ............................... 14 III. Analysis of vertical distributions for B. annulata and ~· pallida from transect samples at the Craw 1 Key quarry .............................. 16 IV. Analysis of vertical distributions for ~· annulata at each of seven depths from transect samples at the Crawl Key quarry ............. 17 v. Analysis of vertical distributions for ~· pallida at each of seven depths from transect samples at the Crawl Key quarry ...................... 18 VI. One-way ANOVA for the difference in abundance between ~· annulata and ~· pallida at the Crawl Key quarry . ................................... 19 VII. One-way ANOVA for differences in~· annulata's abundance at different depths at the Crawl Key quarry . .............................................. 2 0 VIII. One- way ANOVA for differences in~· pallida's abundance at different depths at the Crawl Key quarry . .............................................. 2 0 IX. Comparisons of paired abundance means for ~· annulata at different depths using the post hoc Tukey test . ...................................... 21 X. Comparisons of paired abundance means for ~· pallida at different depths using the post hoc Tukey test ....................................... 23 vi LIST OF FIGURES 1. Inset map of southern Florida showing the location of Crawl Key. Enlarged map of the Crawl Key area of Monroe County, Florida, indicating the location of the quarry (X) where field work was conducted ....................... 6 2. Scatter plots and regressions of the abundances of anemones as a function of depth (a)~· pallida and (b)~· annulata ............. 27 vii INTRODUCTION An important goal of evolutionary ecology is to discover how specific processes affect patterns observed in nature. The process of dispersion is usually the result of behavioral interactions among individuals. Analysis of the patterns and behavioral interactions of individuals is important to comprehending the behavioral ecology of these organisms (Brown & Downhower, 1988). Furthermore, the spatial arrangement of individuals may be central to understanding the structural and functional relationships of population density, persistence and gene flow in ecological communities. Competition among con- and heterospecifics may be a major factor determining spatial distributions. Aggressive behavior (i.e., any hostile act or threat made to protect territory, the family group or offspring, or to establish dominance [Lincoln et al. 1982]) is a primary facet of interference competition. Connell (1961) pioneered studies that examined the relationship between heterospecific competition and dispersion under field conditions. He found that the lower limits of the intertidal barnacle Chthamalus stellatus were restricted by competition for space with another barnacle, Balanus balanoides, which occurs lower in the intertidal zone. Aggression and its effect on dispersion has also been studied 1 2 in many anthozoans (Lang, 1973; Sheppard, 1979; Cope, 1981; Bradbury & Young, 1983; Logan, 1984; Chadwick, 1991). However, most of the studies with anemones examined conspecific interactions (Francis, 1973a, b, 1976, 1988; Purcell & Ki tting, 1982; Quicke & Brace, 1983; Dorsett & Turner, 1986). Few studies have addressed heterospecific interactions and their relationships to dispersion in sea anemones. Chao (1975) reported that the dispersion of Anthopleura elegantissima was limited, in part, by heterospecific competition with Metridium senile and Corynactis californica. Likewise, Sebens (1976) found a hierarchy of species aggressiveness in laboratory experiments. This correlated with dispersion in the field; the more aggressive anemones occupied stable areas of the reef. Less aggressive anemones occupied unstable substrates such as areas of rapid coral growth and collapse, thereby avoiding competition with anemones and other organisms. Additional work is needed in order to elucidate life history diversity, and to better understand the relationships between heterospecific competition and spatial distribution in sea anemones living in different environments, and therefore, under different selective forces. 3 I employed a combination of field and laboratory observations and experiments to define and describe the local spatial distribution of two sympatric sea anemones, Bartholomea annulata Leseur and Aiptasia pall ida Verrill, found in the Florida Keys. These data were used to test the following a priori hypotheses as suggested by the earlier findings outlined above, that: (1) B. annulata is more aggressive than A. pallida; (2) heterospecific individuals capture space and minimize contact through aggressive interactions; ( 3) dispersion patterns for both species are clumped; and (4) ~· annulata and~· pallida exhibit vertical zonation. MATERIALS AND METHODS Natural history of study animals Bartholomea annulata is distributed from Florida to Texas, and from the Bahamas southwards through the West Indies (Carlgren, 1951; Amos & Amos, 1988). This species
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