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The Ecology of Patch Reef Fishes in a Subtropical Pacific Atoll

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The Ecology of Patch Reef Fishes in a Subtropical Pacific Atoll THE ECOLOGY OF PATCH REEF FISHES IN A SUBTROPICAL PACIFIC ATOLL: RECRUITl~lT VARIABILITY, COMMUNITY STRUCTURE AND EFFECTS OF FISHING PREDATORS A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILUtENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY MAY 1989 By Robert E. Schroeder Dissertation Committee: J maes D. Parr ish, Chairraan Leonard A, Freed Ernst $. Reese Stephen V. Ralston Leighton R. Taylor Kent: W. Bridges We certify that ve have read this dissertation and that, in our opinion, it is satisfactory in scope and quality as a dissertation for the degree of Doctor of Philosophy in Zoology. Dissertation Committee: onar ree Ernst S. Reese ACKNOWLEDGMENTS I sincerely thank the members of my committee for their helpful comments, which greatly improved the quality of this dissertation. Dr. Parrish deserves special recognition for conceiving the idea for the predation experiment, serving as the principal investigator on the Sea Grant projects which largely supported this work, and for going out of his way repeatedly, to support me personally and the proper completion of this research effort. The field work required for this dissertation was labor intensive and physically demanding. I thank all the people who painstakingly contributed their assistance many in a major way! to the long days in the water, often under severe conditions, and late evenings of data transcription and field preparation. Jim Norris deserves special thanks in this regard, having done an excellent job in serving as my main dive buddy during most of the Midway cruises. I am indeed grateful to the many students who contributed to the necessary laboratory work and assisted with data and manuscript preparation. I am also thankful to my wife, Maryjane, for her enduring patience and support, and to my parents, Vernon and Mary Schroeder, for their faith and support throughout my graduate career. This work is a result of research "Effects of Fishing on Predation and the Structure of the Fish Resource Community", Projects MR/R-20 and, NI/R-4! sponsored by the Univ, of Hawaii Sea Grant College Program under Institutional Grant Nos. NA8lAA-D-00070 and NA79AA-D- 0085 from NOAA Office of Sea Grant, Dept. of Commerce. Additional financial and logistic support were provided by the Hawaii Cooperative Fishery Research Unit sponsored by the Hawaii Dept. of Land and Natural Resources, the University of Hawaii, and the U.S. Fish and Vildlife Service, Dept. of Interior!, the American Fishing Tackle Manufacturers Association in the form of the Andrew J. Boehm Fellowship Award!, Sigma Xi, The Scientific Research Society in the form of a Grant-in-Aid of Research!, the 14th U.S. Coast Guard District, and the Midway- Naval Air Facility U.S. Navy! and its contractor, Base Services Inc. The ecology of patch reef fishes was studied to quantify the main factors that affect tha natural variability of t' he fish community and to determine the effects produced on the community by experimental removal of predators. Initially, a year-long baseline description was completed of the physical, biological and ecological characteristics of 8 pristine patch reefs at Midway lagoon. For over 3 subsequent years, piscivorous predators were spearfished at least monthly, often for days at a time, on 4 of the 8 reefs. Fish populations were visually censusad throughout the experiment. In all seasons and years of the project, daily recruitment rate of postlarval fishes to natural patch reefs was compared to that measured on standardized, artificial xeefs of various sizes and degrees of intax'-xeaf isolation. Finally, all baseline measurements were replicated and complete collections wexa made of all fishes, to validate the visual census method. Visual censusing was found to be of adequate precision and accuracy for most resident, non-cryptic species highest for small patch reefs! . Fishes could be assigned to size classes underwater by visual estimate with high accuracy. Rotenone collections wexe highly effective in quantifying many species commonly missed or underestimated in visual censuses. Only a faw species composed the bulk of all recruits, while most species were rare or not seen at all. Variation between species was related to life history strategies or behavioral requirements. High temporal variability was found at the following scales: 1! Annually- where variability increased with the magnitude of recruitment, and different species recruited heavily in different years, suggesting that species specific factors in the plankton are more important than general oceanographic conditions; 2! Seasonally- pulsing strongly in summer, and occasionally late fall, when favorable environmental conditions may maximize growth and survival; and 3! Daily- with 1 or 2 strong peaks each only a few days long! ovex a period of several weeks of low, variable recruitment. Small-scale spatial variability between replicate attractoxs standardized artificial reefs! and between attractor types coral and wire! were both high for a few species recruiting abundantly, although most recruits are probably substrate generalists. Rigorous visual fish censuses can adequately document moderate- to long-term temporal variation in the abundances of x'ecently recruited juveniles on patch reefs i.e., based on similar temporal patterns assessed by daily attractors!. Daily total recruitment rate increased, although at diminishing densities, with attractor! reef size, and with degree of inter-reef isolation. Abundances of recently recruited fish censused on neighboring, natural patch reefs much largex' than attx'actors! increased with reef size. The effect of isolation on these natural reefs was confounded by the stronger effect of reef size. These x'esults suggest that if optimum size and spacing of reefs is provided, either by proper design of artificial reefs or selection of marine vi reserves, managers may enhance fish recruitment and ultimately improve local fisheries. Of the 13S fishes censused an the patch reefs studied, only 6 species together accounted for 70% of the total number of all fish, mainly due to heavy seasonal recruitment pulses. Strong seasonal and annual variability in recruitment was responsible for most of the temporal variation in fish abundance, The structure of patch reef fish communities at Midway was characterized by high unpredictability e.g., great seasonal and/or annua1 variability in recruitment by common species, recruitment limitation for most species, and a high turnover rate detected by frequent sampling!. Some predictions of the theory of island biogeography were also met by these fish communities e.g., species richness coxrelated strongly with patch reef area, volume and relief!. and total fish abundance. Soma populations also exhibited a degree of long-term stability. Species diversity [H'j was similar among different size reefs. The experimental fishing an piscivores produced a catch composed. mainly of lizardfish, due largely to immigration following the removal of other, competitively superior, highly resident piscivares. Scorpionfish and moray eels were also dominant predators. The expected decreases in catch-per-unit-effort were not realized, except for a quantitatively insignificant family hawkfish!. Conversely, the catch of the highly migratory lizardfish actually increased as fishing progressed. Changes in the catch composition fox' other piscivores vii related mainly to major changes in reef size or to patterns of large, inter-year recruitment fluctuations. Census data confirmed the major trends indicted by catch results. Sharks and jacks were attracted to the experimental reefs by spearfishing; the study was unable to determine whether their piscivorous effect was different between reef treatments. Patch reef fish communities at Midway were relatively resilient to long-tern, intense fishing pressure on piscivores. However, enhanced survival of a large, annual, summer recruitment pulse of a common cardinalfish, synchronized with a temporary but significant reduction of lizardfish the most prevalent piscivore! by fishing, suggested that an effect of predation on reef fish populations is experimentally detectable and considerable. However, temporal and spatial variability in recruitment, and reef size differences and changes in size were the primary factors responsible for the observed temporal patterns in fish abundance. Community analysis involves numerous confounding effects and requires the most careful interpretation for valid conclusions. TABLE OF CONTENTS Page ACKNOWLEDGMENTS. ABSTRACT LIST OF TABLES LIST OF FIGURES. xiii GENERAL INTRODUCTION CHAPTER 1. VALIDATION OF THE TOTAL-COUNT VISUAL CENSUS METHOD FOR ASSESSING CORAL PATCH REEF FISH COMMUNITIES. CHAPTER 2. RECRUITMRlT VARIfiBILITY BY CORAL REEF FISHES IN A SUBTROPICAL PACIFIC ATOLL. 61 CHAPTER 3. EFFECTS OF PATCH REEF SIZE AND !SOLAT10N ON CORAL REEF FISH RECRUITMENT. 126 CHAPTER 4. THE ECOLOGY OF PATCH REEF FISH COMMUNITIES IN A SUBTROPICAL PACIFIC ATOLL. 160 CHAPTER 5. EFFECTS OP EXPERIMENTALLY FISHING PREDATORS ON CORAL PATCH REEF FISH COMMUNITIES. 236 GENERAL CONCLUSIONS. 317 LIST OF TABLES Table Page 1.1. Summary of patch reef sizes area and volume! . 35 1,2. Coefficients of variation ! for census replicates for taxa mostly species! present on all control reefs in July-August 1985 36 1.3. Relationship between actual fish standard length SL in cm! classes and length classes estimated visually underwater. 38 1.4. Summery of fish collected from rotenone stations
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