SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of “diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Contributions to Anthropology
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In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world.

Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover.

Robert McC. Adams

Secretary

Smithsonian Institution

SMITHONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 32

Hydrographic and Meteorological Studies
of a Caribbean Fringing Reef
at Punta Galeta, Panamá:
Hourly and Daily Variations for 1977–1985

John D. Cubit, Ricardo C. Thompson,
Hugh M. Caffey, and Donald M. Windsor

SMITHSONIAN INSTITUTION PRESS
Washington, D.C
1988

ABSTRACT

Cubit, John D., Ricardo C. Thompson, Hugh M. Caffey, and Donald M. Windsor. Hydrographic and Meteorological Studies of a Caribbean Fringing Reef at Punta Galeta, Panama: Hourly and Daily Variations for 1977-1985. Smithsonian Contributions to the Marine Sciences, number 32, 220 pages, 1 figure, 59 tables, 1988.—This report describes hourly and daily conditions of hydrographic and meteorological factors monitored on a fringing coral reef on the Caribbean coast of the Republic of Panama from January 1977 through December 1985. The methods of monitoring are described in detail, including the performance of various types of equipment under the harsh physical conditions at this site. The data include the following variables: mean hourly wind speed and direction; maximum hourly wind speed and direction; daily maximum and minimum air temperature; hourly air temperature; hourly solar radiation;hourly rainfall; hourly water level; hourly upstream and downstream sea temperatures; and daily salinity. These values are listed directly, with statistical summaries for time of day, days, and months.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution′s annual report, Smithsonian Year. SERES COVER DESIGN: Seascape along the Atlantic coast of eastern North America.

Library of Congress Cataloging-in-PublicationData

Hydrographic and meteorological studies of a Caribbean fringing reef at Punta Galeta, Panamá

(Smithsonian contributions to the marine sciences ; no. 32)

Bibliography: p.

Supt. of Docs. SI 1.41:32

1. Meteorology—Panama—Galeta Point—Observations. 2. Oceanography—Caribbean Sea—Observations. 3 Coral reefs and islands—Panama—Galeta Point—Observations. I. Cubit, John D. II. Series.

QC986.P32G354    1987    551.6′57287    87-600218

Contents

Introduction

Acknowledgments

Methods of Monitoring

Wind Speed and Direction

Rainfall

Solar Radiation

Air Temperature

Water Temperature

Water Level

Salinity

Data Validation Procedures

Bibliography

Tables 1-59

Hydrographic and Meteorological Studies
of a Caribbean Fringing Reef
at Punta Galeta, Panamá:
Hourly and Daily Variations for 1977–1985

John D. Cubit, Ricardo C. Thompson,
Hugh M. Caffey, and Donald M. Windsor

John D. Cubit, Ricardo C. Thompson, Hugh M. Caffey, and Donald U. Windsor, Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Panamá.

Introduction

Hydrography and meteorology are often invoked to explain ecological patterns and processes on tropical reefs; however, for such environments, long-term measurements of physical conditions are rare (Moore, 1972; Potts and Swart, 1984; Porter, 1985). The fringing reef at Punta Galeta, on the Caribbean coast of Panamá is an exception. Charles Birkeland, David Meyer, and Gordon Hendler developed environmental monitoring projects at this site in the early 1970s (see Meyer and Birkeland, 1974; Meyer et al., 1975; Birkeland et al., 1976; Hendler, 1976). The projects were continued as part of the Smithsonian Institution’s International Environmental Sciences Program, which was designed to support long-term studies of organisms and their environment. These investigations are part of the more than 120 published studies describing the biota, physical environment, and ecology of the reef, including more than 70 studies that attribute variations of distribution, abundance, reproduction, behavior, and other aspects of the reef species to influences of physical conditions (see Bibliography).

Although Punta Galeta is less than ten degrees from the equator, analyses of the long-term monitoring data show strong seasonal patterns in the physical environment (Cubit et al., 1986; Cubit et al., in prep.). The present report focuses on the variations of the physical environment that occur over shorter periods of time; i.e., on scales of hours, days, weeks, and months. The data were collected in the years 1977 through 1985, extending the records published from earlier studies (Meyer and Birkeland, 1974; Meyer et al., 1975; Birkeland et al., 1976; Hendler, 1976), as well as providing information for factors not previously monitored. More than 15 years of data now exist for some factors, providing a continuity of information that is unique in its duration and detail for a marine site in the tropics. The hourly and daily data in this report are intended to serve three purposes. First, they are the basic documentation for the analyses and summary statistics presented in the other published studies from Punta Galeta. Second, they demonstrate the erratic variations of the physical environment that occur in the short term, but which are evened out and not apparent in the analyses of long-term patterns. Third, the short-term summary statistics and unreduced data can be applied by other investigators to their own purposes, allowing them to draw their own conclusions regarding variability, durations of extremes, frequencies of conditions, and other aspects of the data whose meaning depends on the questions, organisms, and processes under consideration. Broader overviews, such as seasonal patterns and frequencies of variation, as well as interactions among physical factors and implications of the data for understanding physical and biological processes on the reef, have been described elsewhere (see the Bibliography; for recent analyses, see Cubit et al., 1986; Cubit et al., in prep.).

ACKNOWLEDGMENTS.—This work was supported by the Smithsonian Institution Environmental Sciences Program. Many of the monitoring procedures were originated by C. Birkeland, G. Hendler, and D. Meyer. We thank J. Burgett for programming assistance, E. Leigh and I. Rubinoff for helpful discussion, and J. Horn for careful editing.

Methods of Monitoring

In this project we monitored the following factors: wind speed and direction, rainfall, solar radiation, air temperature, upstream and downstream water temperatures, water level, and salinity. Our choices of monitoring instruments and procedures were based on the level of resolution needed, cost-effectiveness, ability of equipment to withstand the harsh environment, and serviceability of the equipment in this remote location. Air-borne salts, wave-driven debris, suspended sediment, high humidities, high temperatures, intense solar radiation, heavy rainfall, strong winds, perching birds, nesting wasps, excavating animals, fouling organisms, crevice corrosion, electrolysis, unstable electrical service, and lightning are examples of factors that destroyed equipment or interfered with measurements. We discuss some of these problems and their solutions. Factors, such as nutrient concentrations, current velocities, water turbidity, wave height, and underwater light intensities were not continuously monitored because we could not get acceptable measurements within the limits of personnel time and budgets. Such measurements, however, were made as part of shorter term projects. In the following description of methods we also include comments about the installation and performance of equipment that may be useful to other investigators.

Instruments were checked about 5 days per week. For instruments recording on chart paper, times were marked on the charts and the charts were rescaled before reading, if necessary. Chart recordings were read by hand or using a digitizer. Where possible, readings were double-checked against independent measurements. All temperatures were checked at least once weekly with a precision mercury thermometer; more frequent checks were made if problems were suspected. Water levels were compared with hand measurements made with a meter stick. Wind speeds were checked against a totalizing anemometer, and wind directions were checked with a magnetic compass. Rainfall was compared with the data from simple collector gauges. The salinometer was calibrated with distilled water. Solar radiation was checked against expected values for full sun at midday and darkness at night; this was the only factor that we were unable to check on a regular basis against a precise standard.

In 1983 data logging began with Datapod portable digital recorders (Omnidata International Inc., Logan, Utah). These battery-operated instruments record data on nonvolatile, solid-state, memories (EPROMs). The EPROMs were read with apparatus connected directly to a microcomputer, using a program that converted the data into the appropriate units and stored them on magnetic disks, thus bypassing the labor-intensive, time-consuming, and error-prone steps of reading charts by hand or with a digitizer. Because the Datapods were sealed in plastic boxes, self-contained, and isolated from the laboratory, fewer problems were posed by environmental conditions such as corrosion, power failures, and lightning. Housings constructed of insulated ice chests with aluminum roofs and ventilation ports protected the Datapods in the field. The locations of the instruments are illustrated in Figure 1. Specific details for each factor monitored are listed below.

WIND SPEED AND DIRECTION (Tables 1–22).—Anemometers at the seaward edge of the reef flat recorded wind speed and direction 3 m above mean low water level. All data before 15 September 1983 were recorded with an MRI Mechanical Weather Station model 1072 (Meteorological Research, Inc., Altadena, California), which recorded wind runs and direction on pressure-sensitive chart paper. Despite special covers and frequent maintenance, the clockwork mechanism and metal construction of this instrument required frequent repair; two instruments were alternately used to reduce the time without data. After 15 September 1983, wind data were collected with a Datapod 214 set to record hourly mean wind speed, hourly mean wind direction, hourly peak wind speed, and the direction of peak speed. (Means are of 12 readings per hour.) The Datapod sensors and lightning arrestor were grounded to a 1.27 cm diameter copper bar driven into the reef. The anemometer and wind direction sensors were made of plastic, which endured well in this environment; however, salt crystals accumulated between the anemometer cup assembly and the transducer housing. This required disassembly and cleaning of the apparatus about every three months.

FIGURE 1.—Map of the reef at Punta Galeta, showing locations of monitoring stations: 1 (laboratory roof), measured solar radiation and rain 2, measured air temperature, upstream water temperature, and water level measured wind speed and direction; 4, measured solar radiation downstream water temperature; 5, measured maximum-minimum air temperature and salinity.

RAINFALL (Tables 23–31).—The rainfall data reported here were measured with a Stevens Type A water level recorder (Leupold and Stevens Inc., Beaverton, Oregon, USA). This instrument recorded the water level in a cistern, which received rainfall from a 30.5 cm diameter collector mounted on the roof of the laboratory. (The water level recorder is further described under “Water Level,” below.)

SOLAR RADIATION (Tables 32–38).—Wide-spectrum solar radiation data are reported here. Until 15 May 1981 these data were recorded from a Moll-Gorczynski-type pyranometer having a 300–2500 nm wavelength range (Kipp and Zonen Co., Delft, Holland). This pyranometer was mounted on the highest point of the laboratory roof, about 5 m above mean low water. A twisted-pair cable with grounded shield connected the sensor to a Mk V digital volt-time integrator with a printed paper tape output (Lintronic Ltd., London, England) inside the laboratory. This apparatus and other instruments near it in the laboratory were destroyed by lightning on 15 May 1981. On 18 May 1984 we began using a Datapod 211 to record from a LI200 pyranometer sensor having a 400–1100 nm wavelength range (Li-Cor, Inc., Lincoln, Nebraska). The Li-Cor sensor was mounted 2 m above the reef flat where it was isolated from the laboratory. The aluminum mounting plate for the sensor was grounded through a 0.98 cm brass rod connected to a 1.27 cm diameter copper rod driven into the reef. Birds (kingfishers and terns) occasionally used the sensor as a foraging perch, a problem that was reduced on 8 April 1985 by using thin wires to hinder access to the perch. (The ends of the wires were bent to avoid injuring the birds and were arranged to prevent shading the sensor.) Data were recorded as the average millivolt output for one hour intervals and converted to watts/m^2 for analysis.

AIR TEMPERATURE (Tables 39–41).—For 1977 only, maximum and minimum air temperatures were read from the chart recordings of a Hydrolab model RT-125 Marine Thermometer (Hydrolab Corp., Austin, Texas), the same instrument used to measure water temperature (see below). From October 1978 to May 1984, air temperatures were measured with a maximum-minimum thermometer (No. 5458, Taylor Instrument, Arden, North Carolina) that was read and reset once daily, about five times per week. The thermometer was mounted in the shade beneath the laboratory dock until 26 August 1982, when it was moved to a ventilated housing above the dock. On 6 July 1983, the housing was repositioned on the dock. The maximum-minimum values in the tables are for the period since the previous reading.

Beginning 11 April 1984, we recorded hourly mean temperatures with a Datapod 212 connected to a sensor mounted about 1.5 m above mean low water level on the midreef flat. The temperature sensor was mounted in a shelter aspirated by a wind-driven turbine atop a 3 m convection stack made of PVC pipe with an inside diameter of 10.5 cm. The stack was painted black to increase convection at low wind speeds and was wrapped with grounded copper wire to drain static charges. The shelter was open at the bottom, but the sensor was shielded from rainfall, salt spray, and direct solar radiation by being recessed within a two-layer bell housing. To prevent heating, the space between the layers of the housing was also aspirated.

WATER TEMPERATURE (Tables 42–49).—Until 21 September 1980, seawater temperature was measured with a Hydrolab RT-125 thermometer which recorded on a continuous paper chart (see “Air Temperature,” above). The temperature sensor was mounted about 15 cm above the reef surface in a depression about 1.5 m in diameter on the midreef flat. The depression did not drain at low water levels, and the sensor was about 20 cm below the surface of the pool during the lowest water levels, which was equivalent to 30 cm below mean low water. The sensor was adjacent to the tide gauge, about 36 m downstream from the seaward edge of the reef flat and the breaking waves. Beginning 11 April 1984, water temperature was recorded with a Datapod 212 in the same location. Water leaking into the the sensors destroyed them in less than a year; this problem was corrected by sealing the probes in a close-fitting copper sleeve with CRC Marine Soft Seal (CRC Chemicals USA, Warminster, Pennsylvania). Beginning 16 May 1984, a second sensor was installed about 91m directly downstream from the midreef sensor. The second sensor was also permanently immersed in a depression about 1.5 m in diameter and 0.25 m deep at lowest low water. Recording at this location was on a Datapod 211. Perforated PVC pipe shaded both sensors. The instruments were set to record hourly mean temperatures. In the data reported here, the midreef location is referred to as “upstream” and the second location as “downstream.”

Self-contained thermographs (Ryan Instruments, Inc., Seattle, Washington) were used to make limited measurements of water temperature in subtidal habitats (Hendler, 1977a). However, in this environment, the stainless steel temperature probes on these instruments were prone to crevice corrosion and other problems, which resulted in aberrant readings before total failure. Because of the expense of repairs and unreliability of the data, we discontinued use of these instruments.

WATER LEVEL (Tables 50–58).—A Stevens Type A water level recorder (Leupold and Stevens Inc., Beaverton, Oregon, USA) measured water levels above the reef flat. A 21 cm diameter float and counter weight, operating through a 5:1 gear reduction, directly drove die pen of this mechanical chart recorder. Water levels were recorded from a 25 cm diameter still well; the entrance to the still well was a PVC pipe 2.54 cm in diameter and about 38 cm long. Bronze wire mesh (coarse scouring pad material) in the entrance tube inhibited the growth of fouling organisms, excluded sediment, and damped short-period wave oscillations. The recorder was housed in a wooden shelter mounted on top of the still well. The base of the well was set in cast concrete reinforced with steel bars driven into the hard substratum of the reef; this was in a depression about 36 m from the seaward edge of the reef flat. Rates of chart movement and accuracy of water level measurement were manually checked about 5 times per week. This is the same apparatus used for the earlier (1973–1976) measurements reported in Meyer and Birkeland (1974), Meyer et al. (1975), and Hendler (1976, 1977a); it was kept in use because its accuracy, precision, resolution, and reliability were better than any electronic method we could find. Conversion of water level to a precise, high resolution electrical signal is difficult with apparatus that can work reliably in this environment. Data on water level are expressed in reference to an arbitrary datum level at the base of the tide gauge; the datum level is about 30 cm below mean low water (see Cubit et al., 1986, for more details).

SALINITY (Table 59).—Water samples for salinity measurements were taken from the upper 15 cm of water flowing across the reef and past the end of the laboratory dock. Samples were made once daily, about 5 times per week. Salinity was measured with a temperature-compensated, hand-held refractometer (model number 10419, American Optical Co., Buffalo, New York). Attempts to record salinity continuously with electronic instruments were unsuccessful. A Hydrolab Corporation (Austin, Texas) conductivity meter with exposed electrodes was repeatedly damaged by electrical potentials during the frequent lightning storms of the rainy season. A Beckman Instruments conductivity meter (Series SMS-900, Cedar Grove, New Jersey) with an induction sensor also failed to operate continuously.

Data Validation Procedures

The tables of data presented include basic statistical summaries for time of day, day, and month. We used various methods to check the data at stages from the actual monitoring through data compilation, analysis, and to the final printout. When the field check of an instrument indicated values outside acceptable limits, we rejected all readings back to the date of the previous check. Our criteria for acceptable data were based on our requirements for precision and accuracy, particularly for the purposes of other studies on Galeta Reef. For example, wind speed and direction were used mainly as indications of seasonal change and wave action, for which precision was more important than accuracy. Comparisons with other anemometers indicated that wind speeds were accurate to within ±5 km/hr, with a precision of 1–2 km/hr. More accuracy was required of temperatures: the tolerance limits of all continuous measurements of temperature (both air and water) were ±0.5°C of the reading on a precision mercury thermometer. Separate, short-term, measurements of photosynthetically active radiation (PAR) were used in the concurrent biological studies, and the purpose of the pyranometer was to provide a relative measure of incoming solar radiation to be related to seasonal changes in other factors.

Data entered into computers were rechecked against spot readings in the field and random checks of the chart recordings, if applicable. Data were scrutinized more closely when errors were more probable; e.g., following repairs or changes of instruments. Because of their biological significance, we carefully traced all extreme values back to the original instrument recordings and calibration procedures. Extremes did occur: their severity, frequency, and consequences are discussed in Cubit et al. (in prep.). If any errors remain in these data, they will be ones similar to adjacent readings; i.e., data mat did not depart from a range of expected values. To avoid human error as much as possible, the following tables have been printed directly from the computer files, and photographically reproduced.

Bibliography

The following references (except those with an asterisk following the publication date) deal specifically with the reefs, seagrass beds, and mangroves at Punta Galeta.

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1978. The Adaptive Significance of the Turf Growth Form: An Enumeration of the Costs and Benefits of Coloniality in Marine Macroalgae. Journal of Phycology, 14(supplement):28.

1979. Exclusion of Sand-plain Algal Assemblages from a Tropical Fringing Reef: The Importance of Fish Grazing. Journal of Phycology, 15(supplement):13.

1980. Algal Ecology on a Caribbean Fringing Reef. Doctoral dissertation, University of California, Irvine.

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1984c. Predictable Spatial Escapes from Herbivory: How Do These Affect the Evolution of Herbivore Resistance in Tropical Marine Communities? Oecologia (Berlin), 64:396–407.

1985. Spatial Patterns of Herbivore Impact and Their Importance in Maintaining Algal Species Richness. In Proceedings of the Fifth International Coral Reef Congress, Tahiti, 4:29–34.

1986. The Functional Geometry of Seaweeds: Ecological Consequences of Thallus Layering and Shape in Contrasting Light Environments. In T.J. Givnish, editor, On the Economy of Plant Form and Function, pages 635–666. Cambridge, Massachusetts: Cambridge University Press.

Hay, M.E., T. Colburn, and D.Downing

1983. Spatial and Temporal Patterns in Herbivory on a Caribbean Fringing Reef: The Effects on Plant Distribution. Oecologia, 58:299–308.

Hay, M.E., and S.D. Gaines

1984. Geographic Differences in Herbivore Impact: Do Pacific Herbivores Prevent Caribbean Seaweeds from Colonizing via the Panamá Canal? Biotropica, 16:24–30.

Hay, M.E., and J.N. Norris

1984. Seasonal Reproduction and Abundance of Six Sympatric Species of Gracilaria Grev. (Gracilariaceae; Rhodophyta) on a Caribbean Subtidal Sand Plain. Hydrobiologia, 1167117:63–72.

Heck, K.L., Jr.

1976. Comparative Community Organization in Tropical and Temperate Seagrass (Thalassia tesludinum) Meadows. Doctoral dissertation, Florida State University, Tallahassee.

1977. Comparative Species Richness, Composition, and Abundance of Invertebrates in Caribbean Seagrass (Thalassia tesludinum) Meadows (Panamá). Marine Biology, 41:335–348.

1979. Some Determinants of the Composition and Abundance of Motile Macro-invertebrate Species in Tropical and Temperate Turtlegrass (Thalassia testudinum) Meadows. Journal of Biogeography, 6:183–200.

Heck, K.L., Jr., and G.S. Wetstone

1977. Habitat Complexity and Invertebrate Species Richness and Abundance in Tropical Seagrass Meadows. Journal of Biogeography, 4:135–142.

Hendler, G.L.

1976. Environmental Sciences Program Marine Studies: Galeta Point In D.M. Windsor, editor, 1975 Environmental Monitoring and Baseline Data, pages 131–252. Washington, D.C.: Smithsonian Institution Environmental Sciences Program, Tropical Studies.

1977a. Environmental Sciences Program Marine Studies—Galeta Point In D.M. Windsor, editor, 1976 Environmental Monitoring and Baseline Data, pages 139–267. Washington, D.C.: Smithsonian Institution Environmental Sciences Program, Tropical Studies.

1977b. The Differential Effects of Seasonal Stress and Predation on the Stability of Reef-flat Echinoid Populations. In Proceedings of the Third International Coral Reef Symposium, Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida, pages 217–223. Florida: University of Miami Press.

1979a. Sex-reversal and Viviparity in Ophiolepis kieri, n.sp., with Notes on Viviparous Brittlestars from the Caribbean (Echinodermata: Ophiuroidea). Proceedings of the Biological Society of Washington, 9:783–795.

1979b. Reproductive Periodicity of Ophiuroids (Echinodermata: Ophiuroidea) on the Atlantic and Pacific Coasts of Panamá In S. Stancyk, editor. Reproductive Ecology of Marine Invertebrates, pages 145–156. Columbia: University of South Carolina Press.

1980. Reproductive Periodicity of Ophiuroids (Echinodermata: Ophiuroidea) on the Atlantic and Pacific Coasts of Panama. Washington, D.C.: Smithsonian Oceanographic Sorting Center.

1984. Brittlestar Color-change and Phototaxis (Echinodermata: Ophiuroidea: Ophiocomidae). Marine Ecology, 5:379–401.

Hendler, G.L., and D.L. Meyer

1982. An Association of a Polychaete, Branchiosyllis exilis with an Ophiuroid, Ophiocoma echinala, in Panamá. Bulletin of Marine Science, 32:736–744.

Jackson, J.B.C.

1973. The Ecology of Molluscs of Thalassia Communities, Jamaica, West Indies, I: Distribution, Environmental Physiology, and Ecology of Common Shallow-water Species. Bulletin of Marine Science, 23:313–350.

Kilar, J.A.

1985. Interactions between the Red Alga Acanthophora spicifera and the Decorator Crab Microphrys bicornutus. Doctoral dissertation, Dalhousie University, Halifax, Nova Scotia, Canada.

Kilar, J.A., and R.M. Lou

1984. Ecological and Behavioral Studies of the Decorator Crab, Microphrys bicornutus Latreille (Decapoda: Brachyura): A Test of Optimum Foraging Theory. Journal of Experimental Marine Biology and Ecology, 74:157–167.

Kilar, J.A., and J. McLachlan

1986. Branching Morphology as an Indicator of Environmental Disturbance: Testing the Vegetative Fragmentation of Acanthophora spicifera and the Turf Morphology of Lawencia papillosa. Aquatic Botany, 24:115–130.

Koehl, M.A.R.

1977. Water Flow and the Morphology of Zoanthid Colonies. In Proceedings of the Third International Coral Reef Symposium, Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida, pages 437–444. Florida: University of Miami Press.

Kramer, D.L.

1983. Aquatic Surface Respiration in the Fishes of Panama: Distribution in Relation to Risk of Hypoxia. Environmental Biology of Fishes, 8:49–53.

LaBarbera.M.

1985. Mechanisms of Spatial Competition of Discinisca strigata (Inarticulata: Brachiopoda) in the Intertidal of Panama. Biological Bulletin, 168:91–105.

Laguna, J.

1985. Systematics, Ecology and Distribution of Barnacles (Cirripedia; Thoracia) of Panama. Master′s thesis. University of California, San Diego.

Lasker, H.R.

1979. Light Dependent Activity Patterns among Reef Corals: Monlastrea cavernosa. Biological Bulletin, 156:196–211.

1980. Sediment Rejection by Reef Corals: The Roles of Behavior and Morphology in Montastrea cavernosa (Linnaeus). Journal of Marine Biology and Ecology, 47:77–87.

1981. Phenotypic Variations in the Coral Montastrea cavernosa and Its Effect on Colony Energetics. Biological Bulletin 160:292–302.

Lawrence, J.M.

1976. On the Reversal of the Covering Response in Lytechinus variegates (Abstract). Florida Naturalist, 39:2.

1976. On the Role of the Tube Feet and Spines in the Righting Response of Sea Urchins (Echinodermata: Echinoidea). American Zoologist, 16:228.

1976. Covering Response in Sea Urchins. Nature, 262:490–491.

Lehman, J.T., and J.W. Porter

1973. Chemical Activation of Feeding in the Caribbean Reef-building Coral Monlastrea cavernosa. Biological Bulletin, 145:140–149.

Lessios, H.A.

1979. Use of Panamanian Sea Urchins to Test the Molecular Clock. Nature, 280:599–601.

1984. Possible Prezygotic Reproductive Isolation in Sea Urchins Separated by the Isthmus of Panamá. Evolution, 38:1144–1148.

Lessios, H.A., J.D. Cubit, D.R. Robertson, M.J. Shulman, M.R. Parker, S.D. Garrity, and S.L. Levings

1984. Mass Mortality of Diadema antillarum on the Caribbean Coast of Panamá. Coral Reefs, 3:173–182.

Lessios, H.A., D.R. Robertson, and J.D. Cubit

1984. The Spread of Diadema Mass Mortality through the Caribbean. Science, 226:335–337.

Levings, S.C., and E.S. Adams

1984. Intra- and Interspecific Territoriality in Nasutitermes (Isoptera: Termitidae) in a Panamanian Mangrove Forest. Journal of Animal Ecology, 53:705–714.

Lighty, R.G., I.G. Macintyre, and R. Stuckenrath

1982. Acropora palmata Reef Framework: A Reliable Indicator of Sea Level in the Western Atlantic for the Past 10,000 Years. Coral Reefs, 1:125–130.

Macintyre, I.G.

1977. Distribution of Submarine Cements in a Modern Caribbean Fringing Reef, Galeta Point, Panamá. Journal of Sedimentary Petrology, 47:503–516.

1978. Reply: Distribution of Submarine Cements in a Modern Caribbean Fringing Reef, Galeta Point, Panamá. Journal of Sedimentary Petrology, Discussions, 48:669–670.

Macintyre, I.G., and P.W. Glynn

1974. Internal Structure and Developmental Stages of a Modern Caribbean Fringing Reef, Galeta Point, Panama. In 7th Caribbean Geological Conference, Guadeloupe, 14 pages.

1976. Evolution of Modern Caribbean Fringing Reef, Galeta Point, Panamá. The American Association of Petroleum Geologists Bulletin, 60:1054–1071.

Mendoza, R.

1977. Inventario de la flora algal de las costas de la Provincia de Colon. Conciencia, 4:13.

Meyer, D.L.

1973. Feeding Behavior and Ecology of Shallow-water Unstalked Crinoids (Echinodermata) in the Caribbean Sea. Marine Biology, 22:105–129.

Meyer, D.L., and C. Birkeland

1974. Environmental Sciences Program Marine Studies, Galeta Point. In R.W. Rubinoff, editor, 1973 Environmental Monitoring and Baseline Data, pages 129–253. Washington, D.C.: Smithsonian Institution Environmental Sciences Program, Tropical Studies.

Meyer, D.L., C. Birkeland, and G. Hendler

1975. Environmental Sciences Program Marine Studies, Galeta Point. In D.M. Windsor, editor, 1974 Environmental Monitoring and Baseline Data, pages 223–409. Washington, D.C.: Smithsonian Institution Environmental Sciences Program, Tropical Studies.

Meyer, K.B.

1977. Dorid Nudibranchs of the Caribbean Coast of the Panama Canal Zone. Bulletin of Marine Science, 27:299–307.

Millar, R.H.

1974. A Note on the Breeding Season of Three Ascidians on Coral Reefs at Galeta in the Caribbean Sea. Marine Biology, 28:127–129.

Montgomery, E., and R.L. Caldwell

1984. Aggressive Brood Defense by Females in the Stomatopod Gonodactylus bredini. Behavioral Ecology and Sociobiology, 14:247–251.

Moore, H.B.

1972.* Aspects of Stress in the Tropical Marine Environment. Marine Biology, 10:217–269.

Palmer, A.R.

1979. Fish Predation and the Evolution of Gastropod Shell Sculpture. Evolution, 33:697–713.

Porter, J.W.

1972. Ecology and Species Diversity of Coral Reefs on Opposite Sides of the Isthmus of Panamá. Bulletin of the Biological Society of Washington, 2:89–116.

1985* The Maritime Weather of Jamaica: Its Effects on Annual Carbon Budgets of the Massive Reef-building Coral Montastrea annularis. In Proceedings of the Fifth International Coral Reef Congress, Tahiti, 6:363–379.

Potts, D.C., and P.K. Swart

1984.* Water Temperature as an Indicator of Environmental Variability on a Coral Reef. Limnology and Oceanography, 29:504–516.

Powell, N.A.

1971. The Marine Bryozoa near the Panama Canal. Bulletin of Marine Science, 21:766–778.

Rabinowitz, D.

1978a. Dispersal Properties of Mangrove Propagules. Biotropica, 10:47–57.

1978b. Early Growth of Mangrove Seedlings in Panama, and an Hypothesis Concerning the Relationship of Dispersal and Zonation. Journal of Biogeography, 5:113–133.

Reimer, A.A.

1975. Effects of Crude Oil on Corals. Marine Pollution Bulletin, 6:39–43.

1975. Effects of Crude Oil on the Feeding Behaviour of the Zoanthid Palythoa variabilis. Environmental Physiology and Biochemistry, 5:248–266.

Reimer, R.D., and A.A. Reimer

1975. Chemical Control of Feeding in Four Species of Tropical Ophiuroids of the Genus Ophioderma. Comparative Biochemistry and Physiology, 51A:915–927.

Robertson, D.R., and G.R. Allen

1981. Stegasles mellis (Emery et Burgess, 1974), le juvenile de la Demoiselle caraibe Stegasles diencaeus (Jordan et Rutter, 1898). Revue Francaise Aquariologie, 7:109–112.

Robertson, D.R., and R.R. Warner

1978. Sexual Patterns in the Labroid Fishes of the Western Caribbean, II; The Parrotfishes (Scaridae). Smithsonian Contributions to Zoology, 255:1–26.

Rubinoff, R.W., and I. Rubinoff

1969. Observations on the Migration of a Marine Goby through the Panama Canal. Copeia, 2:395–397.

Rutzler, L., and W. Sterrer

1970. Oil Pollution: Damage Observed in Tropical Communities along the Atlantic Seaboard of Panamá. BioScience, 20:222–224.

Schemske, D.W.

1980. Evolution of Floral Display in the Orchid Brassavola nodosa. Evolution, 34:489–493.

Schneider, D.C.

1985. Predation on the Urchin Echinometra lucunter (Linnaeus) by Migratory Shorebirds on a Tropical Reef Flat Journal of Experimental Marine Biology and Ecology, 92:19–27.

Sebens, K.P.

1976. The Ecology of Caribbean Sea Anemones in Panamá: Utilization of Space on a Coral Reef. In G.O. Mackie, editor, Coelenterale Ecology and Behavior, pages 67–77. New York: Plenum Publishing Corporation.

1977. Autotrophic and Heterotrophic Nutrition of Coral Reef Zoanthids. Proceedings of the Third International Coral Reef Symposium, Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida, pages 397–404. Florida: University of Miami Press.

1982. Intertidal Distribution of Zoanthids on the Caribbean Coast of Panamá: Effects of Predation and Desiccation. Bulletin of Marine Science, 32:316–335.

Sebens, K.P., and K. DeRiemer

1977. Diel Cycles of Expansion and Contraction in Coral Reef Anthozoans. Marine Biology, 43–247–256.

Smith, W.L.

1973. Record of a Fish Associated with a Caribbean Sea Anemone. Copeia, 3:597–598.

1977. Beneficial Behavior of a Symbiotic Shrimp to Its Host Anemone, Bulletin of Marine Science, 27:343–346.

Steger, R.

In press. Effects of Spatial Refuges and Postlarval Recruitment on the Abundance of Gonodactylid Stomatopods, a Guild of Mobile Prey. Ecology.

Steger, R., and R.L. Caldwell

1983. Intraspecific Deception by Bluffing: A Defense Strategy of Newly Molted Stomatopods (Arthropoda: Crustacea). Science, 221:558–560.

Vasquez-Montoya, R.

1979. Peuplements des herbiers de Thalassia testudinum et d’Halodule sp. de la côte Caraibe de Panama. Doctoral dissertation, Université d’Aix-Marseille II, Marseille, France.

Weinstein, M.P., and K.L. Heck, Jr.

1977. Comments on the Life History and Host-Parasite Relationships of Cymothoa excisa (Isopoda, Cymothoidae) and Three Species of Snappers (Lutjanidae) in the Caribbean Sea, Panama. Fishery Bulletin, 75:875–877.

1979. Ichthyofauna of Seagrass Meadows along the Caribbean Coast of Panamá and in the Gulf of Mexico: Composition, Structure and Community Ecology. Marine Biology, 50:97–107.

Winston, J.E.

1978. Polypide Morphology and Feeding Behavior in Marine Ectoprocts. Bulletin of Marine Science, 28:1–31.

Tables 1–59

TABLE 1.—1977 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 2.—1978 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 3.—1979 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 4.—1980 hourly mean wind speed (km/r) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 5.—1981 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 6.—1982 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 7.—1983 hourly mean wind speed (km/hr) on the reef flat (mcMithly values do not include data from days with missing values; dash indicates no data).

TABLE 8.—1984 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 9.—1985 hourly mean wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 10.—1977 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 11.—1978 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 12.—1979 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 13.—1980 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 14.—1981 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 15.—1982 hourly mean wind direction (octants) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 16.—1983 hourly mean wind direction (Jan-Sep in octants, Oct-Dec in degrees) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 17.—1984 hourly mean wind direction (degrees) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 18.—1985 hourly mean wind direction (degrees) 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 19.—September 1983 through December 1985 maximum hourly wind speed (km/hr) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 20.—1983 wind direction (degrees) at maximum hourly wind speed 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 21.—1984 wind direction (degrees) at maximum hourly wind speed 3 m above the reef flat (month values do not include data from days with missing values; dash indicates no data).

TABLE 22.—1985 wind direction (degrees) at maximum hourly wind speed 3 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 23.—1977 hourly rainfall (mm) on the reef flat.

TABLE 24.—1978 hourly rainfall (mm) on the reef flat.

TABLE 25.—1979 hourly rainfall (mm) on the reef flat.

TABLE 26.—1980 hourly rainfall (mm) on the reef flat.

TABLE 27.—1981 hourly rainfall (mm) on the reef flat.

TABLE 28.—1982 hourly rainfall (mm) on the reef flat.

TABLE 29.—1983 hourly rainfall (mm) on the reef flat.

TABLE 30.—1984 hourly rainfall (mm) on the reef flat.

TABLE 31.—1985 hourly rainfall (mm) on the reef flat.

TABLE 32.—1977 solar radiation (watts/m^2) for January-April and September-December (monthly values d not include data from days with missing values; dash indicates no data).

TABLE 33.—1978 solar radiation (watts/m^2) (monthly values do not include data from days with missing values dash indicates no data).

TABLE 34.—1979 solar radiation (watts/m^2) (monthly values do not include data from days with missing values dash indicates no data).

TABLE 35.—1980 solar radiation (watts/m^2) (monthly values do not include data from days with missing value dash indicates no data).

TABLE 36.—1981 solar radiation (watts/m^2) for January-August (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 37.—1984 solar radiation (watts/m^2) for May-December (monthly values do not include data from day with missing values; dash indicates no data).

TABLE 38.—1985 solar radiation (watts/m^2) (monthly values do not include data from days with missing value dash indicates no data).

TABLE 39.—1981—1984 daily maximum and minimum air temperatures (°C) above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 40.—April-December 1984 hourly mean air temperature (°C) approximately 1.5 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 41.—1985 hourly mean air temperature (°C) approximately 1.5 m above the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 42.—1977 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 43.—1978 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 44.—1979 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 45.—January-September 1980 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with any missing values; dash indicates no data).

TABLE 46.—April-December 1984 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 47.—1985 hourly mean upstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 48.—May-December 1984 hourly mean downstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 49.—1985 hourly mean downstream water temperatures (°C) on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 50.—1977 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 51.—1978 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 52.—1979 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 53.—1980 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 54.—1981 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 55.—1982 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 56.—1983 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 57.—1984 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 58.—1985 hourly mean water level (cm) above Galeta datum on the reef flat (monthly values do not include data from days with missing values; dash indicates no data).

TABLE 59.—1977–1985 daily salinity readings (ppt) at Galeta dock (monthly values do not include data from days with missing values; dash indicates no data).

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