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4-25-2017 Growth of Orbicella faveolata in La Parguera, Puerto Rico Darren B. Marshall Nova Southeastern University, [email protected]

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NOVA SOUTHEASTERN UNIVERSITY HALMOS COLLEGE OF NATURAL SCIENCE AND OCEANOGRAPHY DEPARTMENT OF MARINE AND ENVIRONMENTAL SCIENCE

Growth of Orbicella faveolata in La Parguera, Puerto Rico

By

Darren B. Marshall

Submitted to the Faculty of Nova Southeastern University Halmos College of Natural Science and Oceanography Department of Marine and Environmental Science In partial fulfillment of the requirements for the degree of Master of Science with a specialty in:

Marine Biology

Nova Southeastern University April, 2017

Growth of Orbicella faveolata in La Parguera, Puerto Rico

Submitted in Partial Fulfillment of the Requirements for the Degree of

Masters of Science:

Marine Biology

Nova Southeastern University Halmos College of Natural Science and Oceanography Department of Marine and Environmental Science

April 2017

Approved:

Thesis Committee

Major Professor: ______Richard E. Dodge, Ph.D.

Committee Member: ______Andia C. Fonnegra, Ph.D.

Committee Member: ______Ryan P. Moyer, Ph.D.

Acknowledgements

This thesis project has been a journey and an education that has only been possible thanks to you:

To my supervisor Richard Dodge, for giving me the opportunity and support to work in his lab and help me see coral in a new way.

To Terry Quinn, for providing the coral samples that allowed this whole project to get off the ground.

To Kevin Helmle, for introducing me to the world of coral sclerochronology, and showing me the techniques to get the most out of the coral.

To Andia Chaves Fonnegra, for being a guide and a teacher, and for her unending patience in helping me through to the end.

To Ryan Moyer, for collaborating and helping guide to me through this project.

To my family and friends, your patience and understanding through the rough times have made the light at the end of the tunnel that much brighter. Especially Dad, Janet, Tia, Andia, Corinne, Pricilla, Jimmy, and Alex. As well as all of my friends in the cycling community that have helped me burn off lots of stress.

Table of Contents

ABSTRACT:...... 1 KEY WORDS: ...... 1 1. INTRODUCTION: ...... 2 1.1 Background ...... 2 1.2 Objectives ...... 4 2. METHODS AND MATERIALS:...... 5 2.1 Study Area – La Parguera, Puerto Rico ...... 5 2.2 Core sampling and X-radiography ...... 6 2.3 Growth Measurement Methods (Annual and Subannual) ...... 10 2.4 Coral Time Series Methods: Chronology Construction and Comparisons ...... 11 2.5 Temperature Time Series ...... 13 3. RESULTS: ...... 15 3.1 Determination and comparison of average skeletal growth parameters ...... 15 3.2 Annual and Subannual Time Series of Coral Growth Parameters and Correlation Relationships Between Individual Coral Chronologies and Summary Chronologies ...... 21 3.3 Relationships Between the Extension, Density, and Calcification Parameters by Correlation ...... 39 3.4 Correlation Relationship of Time Series of SST to Coral Master Chronology Time Series Over 1973-2004 ...... 44 4. DISCUSSION: ...... 46 4.1 Growth Parameters ...... 46 4.2 Climate Change Relationships with Coral Growth ...... 48 5. CONCLUSIONS: ...... 50 6. REFERENCES: ...... 51

ABSTRACT:

Reef-building corals are subject to high amounts of stress, including pollution and rising sea surface temperatures due to climate change. These factors can affect the ability of corals to produce their calcium carbonate skeletons. Evaluation of the effects of climate change may be facilitated by evaluation of records of coral skeletal growth over a long period of time. The aim of this study was to evaluate skeletal growth of the coral Orbicella faveolata in La Parguera, Puerto Rico over a 32-year period. For this, 14 Orbicella faveolata core samples were collected from corals at two reefs (1.2 km apart) in southwestern Puerto Rico. Coral cores were used to obtain skeletal growth data. Average skeletal extension, density, and calcification was determined for subannual and annual periods, and compared between sites. Time series and growth master chronologies were constructed and compared between corals at the two reef sites. In addition, sea surface temperature (SST) data was obtained and summarized into time series, and correlated with coral growth chronologies for the 32-year period. Results showed that two Orbicella faveolata, growth parameters (extension and calcification) were similar between Turrumote and Pinnacles reefs, while density was non-significantly greater on Pinnacles Reef between 1973 and 2004. SST had a weak, and non-significant correlation to growth parameters over time.

KEY WORDS: coral growth, extension, density, calcification, sclerochronology, densitometry, climate change, sea surface temperatures

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1. INTRODUCTION:

1.1 Background

Marine ecosystems throughout the world are being affected by natural and anthropogenic factors, such as climate change, which includes warmer sea surface temperatures (SST) and ocean acidification (OA) for reef ecosystems (Doney et al. 2009). Reef-building corals in near-shore benthic habitats are one example of a system that is susceptible to a myriad of ongoing pressures (Van Woesik and Jordan-Garza 2011; Logan et al. 2014; Gattuso et al. 2015; Ainsworth et al. 2016; Neal et al. 2017; Perry and Morgan 2017). Environmental changes such as OA and increasing SST have been suggested to negatively impact coral growth parameters (extension rate, skeletal bulk density, and calcification rate) (De’ath et al. 2009; Helmle et al. 2011).

Corals are animals belonging to the phylum Cnidaria, that build calcium carbonate (aragonite) skeletons by depositing annually layered growth bands, similar to many species of trees which have annual growth rings (Buddemeier 1974). The annual density bands consist of two subannual bands (high density and low density bands) and record the chronological growth history of the coral (Whitfield 1898; Knutson et al. 1972; Buddemeier et al. 1974; Dodge and Thomson 1974; Moore and Krishnaswami 1974; Hudson et al. 1976; Barnes and Lough 1996; Helmle and Dodge 2011). The growth data that can be recovered from density bands are defined as annual extension rate (cm yr-1), skeletal bulk density (g cm-3), and annual calcification rate (g cm-2 yr-1). The annual extension rate is multiplied by the bulk density to obtain the annual calcification rate (Helmle and Dodge 2011). Coral growth studies, measuring extension, density, and calcification, are typically short term decadal-scale analyses (Wellington and Glynn 1983; Dodge and Brass 1984; Lough and Barnes 1992; Scoffin et al. 1992; Carricart- Ganivet et al. 2000; Carricart-Ganivet and Merino 2001; Worum et al. 2007). Longer century-scale analyses have usually been limited to only extension rates (Hudson et al. 1976, 1994; Dodge 1981; Hudson 1981; Dodge and Lange 1983). However, there have

2 been some century-scale studies that included all three growth parameters (Lough and Barnes 1997; Bessat and Buigues 2001; Helmle et al. 2011).

Early studies in the 1900s were able to ascertain the relationship between coral growth, environmental impacts, and reef development by measuring the growth rates through external observations of the coral (Finckh 1904; Wood Jones 1908; Mayor 1918). The measure of growth band thickness and distance between the bands was initially used to understand how SST affected coral growth (Ma 1933, 1934a, b, 1937). The alternating growth patterns of Madrepora palmata (Acropora palmata, Lamarck 1816) were connected to seasonal variations in water temperatures in the late 1800s (Whitfield 1898). The discovery of trends brought on by seasonal and environmental changes were also detected on the skeletal structures of Paleozoic and modern corals (Ma 1933). Later, the use of X-radiography was implemented to verify earlier work on coral growth, its growth history, and the environmental conditions that could influence coral growth within the density bands of the coral (Knutson et al. 1972). The utilization of X- radiographs is an extremely useful tool as it allows easily visualizing and retrieving of skeletal growth records. Along with measurable growth data, the bands can also reveal anomalous changes in growth, which are brought by fluctuations in the environment, or other stressors, and make them excellent sources of proxy environmental records (Lough and Barnes 1990).

Coral growth rates are affected by extreme changes in sea surface temperatures (SST). According to Kleypas et al. (1999b), the mean annual reef temperature worldwide is approximately 27°C, which includes spatial and temporal variability of ±1-6°C depending on the location. As SST increase, the period prior to the peak maximum temperature threshold stimulates the coral and increases the calcification rate (Lough and Barnes 2000; McNeil et al. 2004; De’ath et al. 2009). If the temperature remains at least 1°C over the maximum tolerable level for a sustained period of time, coral symbiotic algae (Symbiodinium) will leave the host colony, producing coral bleaching (Donner 2005; Carilli 2010). However, these increased calcification rates are not sustainable, and with time will decrease in corals affected by thermal stress, even before reaching the

3 bleaching point (Marshall et al. 2004; Cooper 2008; Cantin et al. 2010). For example, declines in calcification and extension rates in South Thailand have been found in Porites lutea, due to an increase in SST of 0.161°C per decade, within the temperature range between 28 and 30°C (Tanzil et al. 2009). Massive Porites coral cores from the Great Barrier Reef, which contain over 100 years of skeletal records showed a significant decrease in measured calcification (14.2%) and extension rates (13.3%), OA and the variable SST anomalies proved to drive calcification even lower (De’ath 2009). In comparison, the coral Orbicella faveolata from the Florida Keys, showed no significant abnormal effects in relation to SST, in a study where extension and calcification rates were analyzed over a 60-year period (Helmle et al. 2011). These and other studies have been conducted on the effect of the environment on coral growth in Mexico (Carricart- Ganivet 2004), and the Florida Keys (Hudson 1981; Helmle et al. 2011; Manzello et al. 2015). However, there is a lack of information on long term coral growth records in other parts of the Caribbean Sea and the Western Atlantic region.

Montastraea and Orbicella spp. have been investigated for many environmental factors that affect growth including: radiant light (Graus and Macintyre 1982; Bosscher and Meesters 1993), sedimentation and turbidity (Dodge 1981; Hudson 1981; Dodge and Lang 1983; Dodge and Brass 1984; Tomascik and Sander 1985), bleaching (Dodge 1981; Knowlton et al. 1992; Carilli et al. 2009), disease (Aeby and Santavy 2006), increased SST (Carricart-Ganivet 2004; Sammarco et al. 2006; Cantin et al. 2010; Carilli et al. 2010), dissolved nutrients (Bruno et al. 2003), and environmental stress (Druffel 1982; Weil and Knowlton 1994; Kleypas et al. 1999a; Cruz-Pinon and Carricart-Ganivet 2003;

Helmle et al. 2011; Manzello et al. 2015).

1.2 Objectives

Considering the potential effects of future climate change on coral growth and the lack of long term information on coral growth for some reef areas in the Caribbean, the aim of this study was to evaluate skeletal growth between two reefs and over time of the coral Orbicella faveolata from two reefs in La Parguera, Puerto Rico. The specific

4 objectives were: (1) to determine how average growth varied between two reef sites in a small geographic area, and over a common period of 32 years (1973 to 2004); (2) how linear extension, skeletal bulk density, and calcification rate compared; (3) how time series of parameters of subannual and annual coral skeletal growth were related between individual coral chronologies and summary chronologies; and (4) how time series of SST were related to coral growth time series over the period 1973-2004.

2. METHODS AND MATERIALS:

2.1 Study Area – La Parguera, Puerto Rico

Specimens of Orbicella faveolata were collected by Kilbourne et al. (2008) at two sites off La Parguera, Puerto Rico: Turrumote reef and the Pinnacles reef (17° 56.074’N; 67° 0.074’W and 17° 55.964’N; 67° 0.751’W) respectively (Figure 1). La Parguera is located in southwestern Puerto Rico and sits on an insular shelf that extends south for approximately 8 km, before dropping off into the deep ocean. Turrumote reef is a mid- shelf reef 1.8 km in length and runs east-west along a mixed silt-sand sedimentary benthic environment ranging from approximately 3-20 m in depth (Hubbard et al. 1997). Pinnacles is an adjacent, similarly submerged reef platform approximately 1.2 km southeast of Turrumote reef, and similar depth range. Both Turrumote and Pinnacles reefs are environmentally diverse with abundant fish life and benthic coral cover along the southern shelf, including Millepora spp., Acropora palmata (sp), turf algae, a variety of sponges, as well as massive reef building corals: Dendrogyra cylindricus, Diploria spp., Orbicella spp., and Siderastrea spp. (Garcia-Sais et al. 2008). In comparison to the other reefs within La Parguera, Turrumote reef and the Pinnacles reef are the furthest offshore and are approximately 3-5 km from shore (Garcia-Sais et al. 2008).

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Figure 1. Location of the Study sites in La Parguera, Puerto Rico.

2.2 Core sampling and X-radiography

Kilbourne et al. (2008) collected 19 O. faveolata coral skeletal core samples for chemical analysis. Skeletal core samples were extracted from corals on Turrumote and Pinnacles reefs by using an underwater coring drill in August of 2004. The cores were previously cut longitudinally down the growth axis into parallel sided slabs approximately 0.6-0.8 cm thick, with initial X-radiographs being taken at the time (Kilbourne et al. 2008).

I conducted further processing of the slabs. The thickness of each slab was measured with a Mitutoyo deep throat thickness gauge at various points to map out the thickness of the slab. A precision surface grinder (Central Machine) with a 6x12 in. vacuum stage was then used to remove any thickness variations remaining to within 0.001 cm. Coral slabs were thoroughly rinsed with fresh water and allowed to dry for at

6 least 24 hours, then re-measured at 1 cm intervals throughout the slab for the intended growth measurement transects.

X-radiographs of each core were taken with a dental X-ray machine (X-ray Manufacturing Company of America) operating at 70kV and 15 mA, positioned 125 cm above each coral sample. The slab was placed on top of a 20x25 cm Kodak Industrex AA400 X-ray film packet which was positioned on top of a lead vinyl sheet, which in turn, was on top of a 0.655 cm aluminum plate to reduce backscatter. An aluminum wedge (Figure 3) was included next to the coral slab to calibrate the skeletal density via optical densitometry (Buddemeier 1974; Dodge and Kohler 1985; Chalker et al. 1985; Helmle et al. 2002). Exposure times varied for each X-ray due to the thickness of the individual slabs.

The X-radiographs were manually developed based on resolution guidelines from Kodak. That consisted of 4 minutes in a manual developer solution, 1 minute in a Kodak indicator stop bath, 4 minutes in a rapid fixer solution, and 60 minutes in a fresh water wash bath, followed by 30 seconds in Kodak Photoflo Solution. They were then rinsed and hung to dry.

The negatives were digitized using a high-resolution medical X-ray scanner (RDI Cobrascan CX-312T). The resolution and image capture settings were 357 dpi, using an image resolution of 256 shades of gray and linear gamma image correction, with a maximum gray-scale saturation value of 255. The X-radiographs were horizontally scanned to provide the necessary orientation for transect collection. Additionally, two transparent rulers, positioned vertically and horizontally, were scanned to provide x- and y-axis scaling calibration during the transect measurement process.

The X-radiography of each coral was inspected for clarity and quality of growth bands. Of the 19 original specimens, only 14 had growth bands that were distinguishable to provide several decades of sequential growth data (Table 1). Of the 14 O. faveolata

7 samples from La Parguera, Puerto Rico, 9 were from Turrumote reef, and 5 were from Pinnacles reef.

Table 1. Collection data from 14 La Parguera coral specimens.

Age Depth Core Labels Chronology Records Collected (years) (m) Turrumote Reef (LPT) 04LPT-A 2004-1794 211 4.5 04LPT-B 2004-1695 310 4.5 04LPT-C 2004-1943 62 4.5 04LPT-D 2004-1918 87 4.5 04LPT-E 2004-1939 66 4.5 04LPT-F 2004-1865 140 4.5 04LPT-G 2004-1966 39 4.5 04LPT-H 2004-1918 86 4.5 04LPT-I 2004-1748 257 4.5 Pinnacles Reef (LPP) 04LPP-A 2004-1973 32 6.1 04LPP-B 2004-1901 104 6.1 04LPP-C 2004-1825 181 6.1 04LPP-D 2004-1839 166 6.1 04LPP-I 2004-1970 35 7.6

Transect areas for digital analysis of growth data were then selected on each of the digitized X-radiographs perpendicular to growth bands to provide a quantitative measure of linear extension and optic density. Optic density was converted to skeletal density using a digitized transect of density change along the long axis of the digitized calibration wedge (Dodge and Brass 1984; Chalker et al. 1985).

Data was then processed using the computer program CoralXDS+ (Helmle et al. 2002, 2011, http://cnso.nova.edu/coralxds/index.html) to determine boundaries of coral skeletal annual bands. For each band parameter data was determined on the high density portion, the low density portion, and the entire annual portion.

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Figure 2. Annual and subannual growth measurement output (Coral XDS+).

Figure 3. Coral XDS+ measurement transect layered on an X-ray image, next to the aluminum wedge.

Parameter data was linear extension (cm yr-1), bulk density (g cm-3), and calcification (g cm-2 yr-1) for band and band portion. Calcification for each year was calculated by extension multiplied by bulk density.

Extension (cm yr-1) x Density (g cm-3) = Calcification (g cm-2 yr-1)

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2.3 Growth Measurement Methods (Annual and Subannual)

Data of each coral for each parameter of extension, density, and calcification for the annual and subannual band portions over the 32-year (representing given the time period 1973-2004) common period were averaged for each coral, and standard deviation and coefficient of variation were calculated. Table 2 represents the measurements for all of the 14 corals, while Table 3 is a summary of the measurements for Turrumote reef and Pinnacles reef, as well as the total values for all 14 corals. The mean was averaged and taken for each coral for each parameter to get the grand average of each site. The standard deviation was derived by squaring the differences of the mean with the equation below:

푁 1 휎 = √ ∑( 휒 + µ)2 푁 푖 푖=1

The coefficient of variation was found by dividing the standard deviation over the mean.

To evaluate if coral growth rates differed between Turrumote and Pinnacles reefs, annual and subannual extension, density, and calcification over the common period were compared with an independent sample t-test between locations. Prior to all three analyses, the assumption of homogeneity for variance was assessed using the Levene’s Test for Equality of Variances, with one such test per dependent variable (growth parameter). The results of the Levene’s Test were significant, suggesting that the data in each group were distributed with variances significantly different from one another, and violating the assumption of homogeneity of variance in all three cases (p < .05 for all). Because of these violations to equality of variance, the Welch t statistic was used, as it does not require that the assumption of equality of variance (Stevens 2009). The Welch t-test is also known as the unequal variances t-test (derived from the Student’s t-test), and proves to be more accurate when the sample sizes are unequal. A two sample location test, or

10 two-tailed test was chosen, and normality was also assumed, even though Welch’s t-test was made for unequal variances. The Welch’s t-test is defined by the following formula:

x̅ −x̅ 푡 = 1 2 s2 S2 √ 1 + 2 N1 N2

2 where x̅1, 푠1 , and N1 are the first sample mean, sample variance, and sample size, respectively.

2.4 Coral Time Series Methods: Chronology Construction and Comparisons

Each density band is formed within an annual time period. The HD band formation begins around the middle of July and ends around September (Hudson et al. 1976). The LD band, therefore, begins around October and finishes around the middle of July in the following year.

The 14 individual coral growth parameter chronologies of this study ranged in duration from 1695-2004 to 1973-2004. Detailed analysis was concentrated on the common period of 1973-2004.

Time series of raw data for each of the parameters of extension, density, and calcification for annual and subannual bands were converted into standard anomaly data (STDA) chronologies. The purpose was to reduce variability caused by differing average growth (Fritts 1976). STDA chronologies were accomplished by subtracting the annual mean of each parameter over the common period of 32 years (1973-2004) from each yearly value and dividing by the standard deviation of the common 32-year period. The formula is:

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STDA = Average Growth over Annual the Common Period Growth - of 32 Years

Standard Deviation of Average Growth over the Common Period

Standardized chronologies were compiled into site and grand master chronologies by averaging standardized data by year for each site and for all corals. The purpose was to evaluate the degree to which master chronologies filtered out individual coral variability and represented common patterns.

To detect common variations on skeletal growth factors for all corals over time, the STDA data were used to construct and compare the master chronologies of the Turrumote reef, the Pinnacles reef, and all corals combined.

Product-moment correlation coefficients were calculated between STDA annual and subannual parameter chronologies of extension, density, and calcification to help quantify the relationships and trends found within the two sites and between the site and master growth chronologies. Pearson’s r correlation coefficient was calculated by the following formula:

Ʃxy r = √Ʃ푥2Ʃ푦2

This equation determines the significance of the relationship for Pearson’s r value, the number sits between zero and ±1 to gauge the relationship strength of the variables being tested. Scatter plot graphs were constructed and correlation coefficients were calculated between STDA annual and subannual parameter master chronologies of

12 extension, density, and calcification to help demonstrate and quantify the relationships between the growth parameters.

Growth parameters obtained in this study were compared to other coral growth studies from the greater Caribbean region. The species of interest for all of the studies was the common reef builder, Orbicella spp. (formerly Montastraea spp.).

2.5 Temperature Time Series

Standardized growth master chronologies of each parameter were compared to annual and subannual water temperatures obtained from the 1°x1° gridded HADISST1.1 dataset focused on 17.50° N, -66.50° W, and 18.50° N, -67.50° W. The water temperature data set consisted of a combination of average monthly in situ observations and satellite AVHRR (UK Meteorological Office, Hadley Centre. HadISST 1.1-Global sea-ice coverage and Sea Surface Temperature, 1870-Present, British Atmospheric Data Center, 2006, http://catalogue.ceda.ac.uk/uuid/facafa2ae494597166217a9121a62d3c), from 1973 to 2004 to match the same common period of time analyzed for coral growth (Rayner et al. 2003).

The 12-month annual period for the SST chronology (SST-A) was chosen to begin in July to correspond with the start of the annual band formation. A seasonal 3- month period of July, August, and Sept (SST-H) was chosen for the high density band formation, and a 9-month period of October through June (SST-L) was chosen for the low density band formation comparison period. The minimum SST (SST-MN), and the maximum SST (SST-MX) were taken by selecting the lowest and highest temperature months per year from the data set of multiple monthly temperature measurements.

A 12-month annual period (July-June), SST time series from the Atlantic Multidecadal Oscillation (AMO) was also correlated against the grand master annual and subannual growth parameters. The AMO was a monthly unsmoothed index of North Atlantic SST that are calculated from the monthly Kaplan SST dataset (Enfield et al.

13

2001), https://www.esrl.noaa.gov/psd/data/timeseries/AMO/. A 12-month annual period (July-June), SST dataset of the North Atlantic Oscillation (NAO) was also used in the correlation (https://climatedataguide.ucar.edu/climate-data/hurrell-north-atlantic- oscillation-nao-index-station-based). The AMO and NAO both used the common 32-year time period (1973-2004).

Pearson’s r correlation coefficients were again used to calculate between time series of annual and subannual SST and the annual and subannual extension, density, and calcification parameters for that year for the grand master chronologies.

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3. RESULTS:

3.1 Determination and comparison of average skeletal growth parameters

Average annual skeletal growth of all corals (n=14) was 0.71 cm yr-1 (extension), 0.97 g cm-3 (density) and 0.68 g cm2 yr-1 over the common period of n=32 years (1973- 2004) for both Turrumote and Pinnacles (Table 2). A summary of growth parameters for each reef including mean, standard deviation (SD), and the coefficient of variation (CV) are also included (Table 3).

Table 2. Mean, standard deviation (SD), and coefficient of variation (CV) (n=32 years) for each of the nine corals at Turrumote reef (T), and each of the five corals at Pinnacles reef (P). The average of all 14 coral means is shown in the last column. All 14 T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I Corals Extension Mean: 0.77 0.76 0.65 0.84 0.83 0.67 0.64 0.59 0.64 0.67 0.74 0.76 0.68 0.69 0.71 cm/yr SD: 0.18 0.15 0.13 0.24 0.19 0.12 0.08 0.10 0.11 0.18 0.17 0.18 0.11 0.21 0.15 CV: 23% 20% 20% 28% 23% 17% 13% 17% 17% 27% 24% 23% 16% 30% 22% Density Mean: 0.90 0.90 0.94 0.86 0.90 0.98 0.91 0.96 0.99 1.06 1.01 1.04 0.99 1.07 0.97 g/cm3 SD: 0.08 0.07 0.07 0.13 0.11 0.06 0.06 0.08 0.10 0.08 0.10 0.10 0.07 0.13 0.09 CV: 9% 8% 8% 15% 13% 6% 7% 9% 10% 7% 10% 10% 7% 12% 9% Calcification Mean: 0.69 0.68 0.61 0.70 0.74 0.66 0.58 0.56 0.63 0.70 0.74 0.78 0.67 0.72 0.68 g/cm2/yr SD: 0.15 0.12 0.11 0.14 0.14 0.12 0.07 0.06 0.08 0.16 0.18 0.14 0.10 0.21 0.13 CV: 22% 17% 18% 21% 19% 18% 12% 11% 13% 23% 24% 17% 15% 29% 20% HD Ext Mean: 0.23 0.22 0.20 0.21 0.20 0.23 0.21 0.20 0.23 0.22 0.24 0.20 0.23 0.18 0.21 cm/yr SD: 0.09 0.10 0.07 0.07 0.08 0.10 0.07 0.07 0.08 0.08 0.08 0.09 0.06 0.07 0.08 CV: 38% 46% 34% 34% 40% 42% 32% 34% 37% 35% 34% 44% 26% 40% 37% HD Den Mean: 1.01 1.03 1.06 0.98 1.05 1.09 1.03 1.08 1.11 1.15 1.09 1.18 1.17 1.17 1.09 g/cm3 SD: 0.09 0.08 0.07 0.14 0.11 0.09 0.06 0.09 0.10 0.09 0.15 0.09 0.07 0.12 0.10 CV: 9% 8% 7% 14% 11% 8% 6% 9% 9% 8% 14% 8% 6% 10% 9% HD Cal Mean: 0.23 0.23 0.21 0.21 0.21 0.25 0.22 0.21 0.26 0.25 0.26 0.23 0.26 0.21 0.23 g/cm2/yr SD: 0.08 0.10 0.07 0.08 0.07 0.10 0.06 0.07 0.10 0.09 0.09 0.11 0.06 0.09 0.09 CV: 37% 45% 34% 38% 35% 41% 29% 33% 38% 35% 34% 48% 25% 42% 37% LD Ext Mean: 0.54 0.54 0.45 0.63 0.63 0.44 0.43 0.39 0.41 0.45 0.50 0.56 0.46 0.51 0.50 cm/yr SD: 0.18 0.17 0.15 0.26 0.20 0.16 0.09 0.13 0.13 0.19 0.19 0.22 0.12 0.20 0.17 CV: 33% 31% 32% 41% 32% 37% 22% 32% 33% 43% 38% 39% 25% 39% 35% LD Den Mean: 0.86 0.85 0.89 0.81 0.85 0.91 0.85 0.89 0.91 1.02 0.98 0.99 0.89 1.04 0.93 g/cm3 SD: 0.09 0.09 0.07 0.11 0.13 0.08 0.08 0.09 0.11 0.08 0.08 0.11 0.08 0.13 0.09 CV: 10% 11% 8% 14% 15% 8% 9% 10% 12% 8% 8% 11% 9% 13% 11% LD Cal Mean: 0.46 0.45 0.40 0.48 0.53 0.41 0.36 0.35 0.37 0.45 0.48 0.55 0.41 0.51 0.44 g/cm2/yr SD: 0.15 0.15 0.13 0.17 0.17 0.16 0.09 0.10 0.12 0.18 0.19 0.19 0.11 0.19 0.15 CV: 33% 33% 33% 35% 32% 39% 24% 29% 32% 41% 39% 34% 28% 38% 34%

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Table 3. Summary of average, standard deviation (SD), and coefficient of variation (CV) for Turrumote reef (T) n=9 coral means, Pinnacles reef (P) n=5 coral means and total of both combined n=14 coral means. All 14 T P Corals Extension Mean: 0.71 0.71 0.71 cm/yr SD: 0.14 0.17 0.15 CV: 20% 24% 22% Density Mean: 0.93 1.04 0.97 g/cm3 SD: 0.09 0.10 0.09 CV: 9% 9% 9% Calcification Mean: 0.65 0.73 0.68 g/cm2/yr SD: 0.11 0.16 0.13 CV: 17% 22% 20% HD Ext Mean: 0.21 0.21 0.21 cm/yr SD: 0.08 0.07 0.08 CV: 38% 36% 37% HD Den Mean: 1.05 1.15 1.09 g/cm3 SD: 0.09 0.10 0.10 CV: 9% 9% 9% HD Cal Mean: 0.22 0.24 0.23 g/cm2/yr SD: 0.08 0.09 0.09 CV: 37% 37% 37% LD Ext Mean: 0.50 0.50 0.50 cm/yr SD: 0.16 0.18 0.17 CV: 32% 37% 35% LD Den Mean: 0.87 0.98 0.93 g/cm3 SD: 0.09 0.09 0.09 CV: 11% 10% 11% LD Cal Mean: 0.42 0.48 0.44 g/cm2/yr SD: 0.14 0.17 0.15 CV: 32% 36% 34%

Differences between Turrumote and Pinnacles reefs that were calculated with Welch’s t-test found no significant differences for the annual growth parameters. Extension, density, and calcification values between Turrumote and Pinnacles corals were found (Welch’s t = 0.01, p = 1.00 and t = 2.04, p = 0.08, and, t = 1.00, p = 0.36 respectively) (Table 4). Pinnacles reef corals had a higher density than Turrumote reef (Figure 4).

The HD and LD means were also compared, and significant differences were not found for these parameters either. HD differences for extension, density, and calcification showed Welch’s t = 0.01, p = 1.00, t = 1.86, p = 0.11, and t = 0.41, p = 0.69, respectively

16

(Table 4). Similarly, LD extension, density and calcification showed no significant differences between reefs (Welch’s t = 0.01, p = 1.00, t = 2.19, p = 0.06, and t = 0.67, p = 0.52 respectively).

Annual Average Coral Extension, Density, and Calcification Growth for Turrumote and Pinnacles Reefs

1.2

0.8 Average Average Growth

0.4 T Ext P Ext T Den P Den T Cal P Cal

HD Average Coral Extension, Density, and Calcification Growth for Turrumote and Pinnacles Reefs 1.5

1

Average Average Growth 0.5

0 T Ext P Ext T Den P Den T Cal P Cal

LD Average Coral Extension, Density, and Calcification Growth for Turrumote and Pinnacles Reefs 1.2

1

0.8 Growth 0.6

Average 0.4

0.2 T Ext P Ext T Den P Den T Cal P Cal

Figure 4. Annual, HD, and LD average extension, density, and calcification growth for Turrumote reef (T) (n=9) and Pinnacles reef (P) (n=5). Each individual coral mean was calculated over the 32-year common period 1973-2004. The values shown are the averages of the coral means over the number of corals at each site. Error bars show ± 1 standard deviation. The y-axes of each graph show extension in cm yr-1, density in g cm-3, and calcification in g cm-2yr-1. 17

Annual, HD, and LD parameters were turned into box and whisker plot graphs to explore the statistical distribution of the growth data (Figure 5). The difference in the quartile distribution for annual extension shows that they have a similar median and quartile range, but the minimum and maximum range (the ends of the whiskers) for Pinnacles is slightly greater. Annual density and calcification for Pinnacles reef, shows a slightly greater distribution range than Turrumote reef.

The HD parameters showed a lower range of distribution for extension and calcification, with similar values when comparing both reefs. However, HD density had a much larger range, with Pinnacles reef greater than Turrumote reef. LD parameters also showing similar extension and calcification distribution, with increased density values, and Pinnacles reef had a greater distribution range.

18

Figure 5. Box plot graphs showing annual, HD, and LD average extension, density, and calcification growth for Turrumote reef (T) (n=9) and Pinnacles reef (P) (n=5). Average means for each parameter were grouped into quartiles over the 32-year common period 1973-2004. The y-axes of each graph show extension in cm yr-1, density in g cm-3, and calcification in g cm-2 yr-1.

19

Table 4. Independent t-test between average annual, HD, and LD growth of Turrumote reef (n=9) and Pinnacles reef (n=5).

Turrumote (n=9) Pinnacles (n=5) Source t df p Mean SD Mean SD Ann Ext 0.00 12.0 1.00 0.71 0.14 0.71 0.17 Ann Den 2.04 12.0 0.08 0.93 0.09 1.04 0.10 Ann Cal 1.00 12.0 0.36 0.65 0.11 0.73 0.16 HD Ext 0.00 12.0 1.00 0.21 0.08 0.21 0.07 HD Den 1.86 12.0 0.11 1.05 0.09 1.15 0.10 HD Cal 0.41 12.0 0.69 0.22 0.08 0.24 0.09 LD Ext 0.00 12.0 1.00 0.50 0.16 0.50 0.18 LD Den 2.19 12.0 0.06 0.87 0.09 0.98 0.09 LD Cal 0.67 12.0 0.52 0.42 0.14 0.48 0.17

Average extension and calcification of the corals from both Turrumote and Pinnacles reefs at La Parguera were within range of other studies conducted in the Caribbean Ocean region (Table 5). However, average density was lower. Corals from Puerto Rico tend to have a calcification rate (0.68 g cm-2 yr-1) similar to corals from Discovery Bay, Jamaica (0.65 g cm-2 yr-1). Depth profiles for the sample collections are in the same general shallow to moderate range, making a more level assessment of the growth values.

20

Table 5. Growth studies comparing skeletal extension, density, and calcification of Orbicella annularis (spp.) in several locations of the Caribbean including this study. Minimum values for the variables are included in the parenthesis. * = calculated values with skeletal extension and density means. From Carricart-Ganivet et al., 2000.

Coral Mean Mean Mean Depth Years Author Location Samples Extension Density Calcification (m) measured (n) (cm yr-1) (g cm-3) (g cm-2 yr-1) Baker and Weber 0.91 1.73 1.57 St. Croix, Virgin Islands 4.5-18 60 - 1975 (0.65-1.04) (1.60-1.82) (1.18-1.76)

0.38 1.71 Dustan 1975 Discovery Bay, Jamaica 8-24 189 1 0.65* (0.17-0.67) (1.39-1.94)

0.94 Dodge 1981 Vieques, Puerto Rico 3-5 85 8 - - (0.79-1.06)

Graus & Macintyre 0.76 1.80 Carrie Bow Cay, Belize 1-25 - 8 1.35* 1982 (0.63-0.93) (1.55-1.99)

Dodge & Brass Buck Island, St. Croix, 0.98 1.28 1.23 3-8 607 10 1984 Virgin Islands (0.61-1.44) (0.78-1.63) (0.77-1.58)

Carricart-Ganivet et 0.87 1.74 1.5 Mexican Caribbean 1.5-10 12 10-11 al. 2000 (0.55-1.54) (1.30-2.10) (0.91-2.80) Carilli et al. 2010 Mesoamerican Caribbean 2.5-13 92 57 0.76 - - Helmle et al. 2011 Florida Keys, USA 4-20 7 60 0.79 1.18 0.91 La Parguera, Current Study 4.5-8 14 32 0.71 0.97 0.68 Puerto Rico

3.2 Annual and Subannual Time Series of Coral Growth Parameters and Correlation Relationships Between Individual Coral Chronologies and Summary Chronologies

The individual chronologies constructed with raw data and the converted chronologies to standardized anomaly data (STDA) showed occasional anomalies in some growth parameters (Figures 6 to 14). This is the case of annual extension, annual calcification, LD extension and LD calcification (Figures 6, 8, 12, and 14 respectively). For the year 1984, it appears that one coral core (T-H) skewed the data with this anomaly. However, reductions in variability is generally suggested for the STDA data (Figures 7, 8, 10, 12, 13, and 14). In the figures, the STDA was used to present the master chronologies of each site in the upper graph and the grand master chronology in the lower graph).

21

Raw Growth for Annual Extension 1.6

1.4

1.2

1

0.8

0.6

Extension Extension (cm/yr) 0.4

0.2

0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for Annual Extension 4 3 2 1

0 STDA -1 -2 -3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 -4 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 6. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for annual extension of all 14 collected corals during the 32-year common period (1973-2004).

22

Raw Growth for Annual Density

1.4 ) 3 1.2

1 Density Density (g/cm 0.8

0.6 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for Annual Density

4 3 2 1

0 STDA -1 -2 -3 -4 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 7. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for annual density of all 14 collected corals during the 32-year common period (1973-2004).

23

Raw Growth for Annual Calcification

1.4

1.2

/yr) 2 1

0.8

0.6

Calcification Calcification (g/cm 0.4

0.2 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for Annual Calcification 4 3 2 1

STDA 0 -1 -2 -3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 8. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for annual calcification of all 14 collected corals during the 32-year common period (1973-2004).

24

Raw Growth for HD Extension 0.6

0.5

0.4

0.3

0.2 HD Extension HD Extension (cm/yr) 0.1

0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for HD Extension 4

3

2

1

STDA 0

-1

-2

-3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 9. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for HD extension of all 14 collected corals during the 32-year common period (1973-2004).

25

Raw Growth for HD Density 1.6

1.4

) 3 1.2

1

0.8 HD Density HD Density (g/cm 0.6

0.4 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for HD Density 4 3 2 1 0

-1 STDA -2 -3 -4 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 -5 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 10. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for HD density of all 14 collected corals during the 32-year common period (1973-2004).

26

Raw Growth for HD Calcification 0.7

0.6

/yr) 2 0.5

0.4

0.3

0.2

HD Calcification HD Calcification (g/cm 0.1

0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years Standardized Anomaly Data for HD Calcification 4 3 2 1

STDA 0 -1 -2 -3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 11. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for HD calcification of all 14 collected corals during the 32-year common period (1973-2004).

27

Raw Growth for LD Extension 1.2

1

0.8

0.6

0.4 LD LD Extension (cm/yr) 0.2

0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years Standardized Anomaly Data for LD Extension 4

3

2

1

STDA 0

-1

-2

-3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 12. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for LD extension of all 14 collected corals during the 32-year common period (1973-2004).

28

Raw Growth for LD Density 1.4

1.3 )

3 1.2

1.1

1

0.9

LD Density LD Density (g/cm 0.8

0.7

0.6 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for LD Density 4 3 2 1

0 STDA -1 -2 -3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 -4 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 13. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for LD extension of all 14 collected corals during the 32-year common period (1973-2004).

29

Raw Growth for LD Calcification 1.2

1

/yr) 2

0.8

0.6

0.4

0.2 LD LD Calcification (g/cm

0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

Standardized Anomaly Data for LD Calcification

3

2

1

0 STDA -1

-2

-3 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years T-A T-B T-C T-D T-E T-F T-G T-H T-I P-A P-B P-C P-D P-I

Figure 14. Upper graph shows raw data chronologies, and lower graph shows standardized anomaly data chronologies for LD calcification of all 14 collected corals during the 32-year common period (1973-2004).

30

The STDA master chronologies from Turrumote reef, Pinnacles reef, and the grand master chronology of each parameter were also plotted together to show their similarities (Figures 15 - 24). Correlations to the grand master chronology were expected since a portion of each site master is included in the grand master.

The site chronologies were similar in their variability throughout the parameters. Site chronologies were not always highly correlated to the grand master chronology showing some variability. For example, annual extension and annual calcification (Figures 15 and 17, respectively), as well as, HD extension and HD calcification (Figures 18 and 20, respectively). Density showed good uniform traits for all three chronologies in the annual, HD, and LD bands (Figures 16, 19, and 22, respectively). This proves that while much of the individual coral variability has been filtered out, common variability is still present.

When statistically comparing the two site masters to each other (Turrumote and Pinnacles), and to the grand master, a strong correlation was found between annual extension and annual calcification (r = 0.910, p < 0.001), LD calcification (r = 0.810, p < 0.001) and LD extension (0.873, p < 0.001), (Table 9). Annual density had a strong correlation with LD density (r = 0.913, p < 0.001). Both sites are similarly positive and negative in line with their correlations. The growth parameters between the sites and the grand master coefficients showed a high degree correlation throughout the matrix (Figures 15-23, and Tables 7-9, highlighted in yellow).

31

Annual Extension of Grand Master and Site Comparisons

2 1.5 1 0.5 0

-0.5 STDA -1 -1.5 -2 -2.5 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years GM T P

Figure 15. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

Annual Density of Grand Master and Site Comparisons 3.5 3 2.5 2 1.5 1

STDA 0.5 0 -0.5 -1 -1.5 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 16. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

32

Annual Calcification of Grand Master and Site Comparisons 1.5 1 0.5 0

STDA -0.5 -1 -1.5 -2 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 17. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

HD Extension of Grand Master and Site Comparisons 1 0.8 0.6 0.4 0.2 0

STDA -0.2 -0.4 -0.6 -0.8 -1 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 18. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

33

HD Density of Grand Master and Site Comparisons 2 1.5 1 0.5 0

STDA -0.5 -1 -1.5 -2 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years GM T P

Figure 19. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

HD Calcification of Grand Master and Site Comparisons 0.8 0.6 0.4 0.2 0

-0.2 STDA -0.4 -0.6 -0.8 -1 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 20. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

34

LD Extension of Grand Master and Site Comparisons 1.5 1 0.5 0

STDA -0.5 -1 -1.5 -2 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 21. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

LD Density of Grand Master and Site Comparisons 3 2.5 2 1.5 1

0.5 STDA 0 -0.5 -1 -1.5 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 22. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14.

35

LD Calcification of Grand Master and Site Comparisons 1.5 1 0.5 0

-0.5 STDA -1 -1.5 -2 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 Years

GM T P

Figure 23. STDA site master chronologies for Turrumote reef (T) n=9, Pinnacles reef (P) n=5, and the grand master chronology n=14

Table 6. Correlation coefficient (r) of the standardized growth parameters for the Turrumote reef site master chronology (n=9), during the common 32-year period (1973-2004). Significant correlations are highlighted in yellow, and bold values are significant at p < 0.05.

r = Ann Ext Ann Den Ann Cal HD Ext HD Den HD Cal LD Ext LD Den LD Cal Ann Ext

Ann Den -0.713 p=0.00 1 Ann Cal 0.924 -0.415 p=0.00 1 p=0.018 HD Ext 0.050 -0.222 -0.054 p=0.785 p=0.222 p=0.771 HD Den -0.337 0.533 -0.193 -0.422 p=0.059 p=0.002 p=0.290 p=0.016 HD Cal -0.037 -0.042 -0.087 0.944 -0.118 p=0.839 p=0.821 p=0.637 p=0.001 p=0.521 LD Ext 0.915 -0.597 0.880 -0.332 -0.151 -0.392 p=0.00 1 p=0.001 p=0.001 p=0.063 p=0.409 p=0.026 LD Den -0.650 0.957 -0.349 -0.271 0.378 -0.147 -0.524 p=0.00 1 p=0.001 p=0.050 p=0.133 p=0.033 p=0.423 p=0.002 LD Cal 0.800 -0.339 0.888 -0.462 -0.090 -0.505 0.940 -0.236 p=0.00 1 p=0.058 p=0.001 p=0.008 p=0.625 p=0.003 p=0.001 p=0.193

36

Table 7. Correlation coefficient (r) of the standardized growth parameters for the Pinnacles reef site master chronology (n=5), during the common 32-year period (1973-2004). Significant correlations are highlighted in yellow, and bold values are significant at p < 0.05.

Ann Ext Ann Den Ann Cal HD Ext HD Den HD Cal LD Ext LD Den LD Cal Ann Ext

Ann Den -0.582 p=0.001 Ann Cal 0.948 -0.309 p=0.00 1 p=0.086 HD Ext -0.231 -0.029 -0.267 p=0.203 p=0.874 p=0.139 HD Den -0.326 0.788 -0.093 -0.249 p=0.068 p=0.001 p=0.614 p=0.169 HD Cal -0.346 0.333 -0.271 0.867 0.231 p=0.052 p=0.063 p=0.133 p=0.001 p=0.203 LD Ext 0.975 -0.532 0.935 -0.435 -0.244 -0.514 p=0.00 1 p=0.002 p=0.001 p=0.013 p=0.179 p=0.003 LD Den -0.430 0.870 -0.170 -0.056 0.493 0.116 -0.382 p=0.014 p=0.001 p=0.353 p=0.762 p=0.004 p=0.527 p=0.031 LD Cal 0.932 -0.353 0.961 -0.477 -0.127 -0.513 0.971 -0.173 p=0.00 1 p=0.047 p=0.001 p=0.006 p=0.487 p=0.003 p=0.001 p=0.345

Table 8. Correlation coefficient (r) of the standardized growth parameters for the Turrumote reef site master chronology (n=9) correlated against the Pinnacles reef site master chronology (n=5), during the common 32-year period (1973-2004). Significant correlations are highlighted in yellow, and bold values are significant at p < 0.05.

T Ann T Ann T Ann T HD T HD T HD T LD T LD T LD r = Ext Den Cal Ext Den Cal Ext Den Cal P Ann Ext 0.451 -0.177 0.540 -0.069 -0.149 -0.089 0.469 -0.141 0.522 p=0.010 p=0.333 p=0.001 p=0.708 p=0.414 p=0.629 p=0.007 p=0.442 p=0.002 P Ann Den -0.421 0.575 -0.264 -0.260 0.334 -0.172 -0.331 0.537 -0.180 p=0.016 p=0.001 p=0.145 p=0.151 0.062 p=0.346 p=0.064 p=0.002 p=0.324 P Ann Cal 0.333 0.013 0.491 -0.166 -0.041 -0.156 0.385 0.037 0.501 p=0.063 p=0.943 p=0.004 p=0.364 p=0.826 p=0.395 p=0.030 p=0.842 p=0.003 P HD Ext -0.141 0.056 -0.185 0.303 0.323 0.410 -0.248 0.029 -0.324 p=0.440 p=0.760 p=0.311 p=0.092 p=0.072 p=0.020 p=0.171 p=0.877 p=0.071 P HD Den -0.164 0.359 -0.035 -0.492 0.452 -0.388 0.021 0.287 0.126 p=0.369 p=0.043 p=0.849 p=0.004 p=0.009 p=0.028 p=0.909 p=0.111 p=0.492 P HD Cal -0.180 0.201 -0.178 0.050 0.576 0.222 -0.198 0.117 -0.248 p=0.325 p=0.270 p=0.330 p=0.788 p=0.001 p=0.222 p=0.277 p=0.524 p=0.172 P LD Ext 0.445 -0.172 0.540 -0.134 -0.207 -0.174 0.488 -0.134 0.556 p=0.011 p=0.347 p=0.001 p=0.464 p=0.257 p=0.340 p=0.005 p=0.465 p=0.001 P LD Den -0.489 0.647 -0.285 -0.114 0.107 -0.082 -0.450 0.675 -0.234 p=0.005 p=0.001 p=0.114 p=0.533 p=0.559 p=0.657 p=0.010 p=0.001 p=0.197 P LD Cal 0.364 -0.060 0.505 -0.169 -0.192 -0.206 0.419 -0.020 0.536 p=0.041 p=0.743 p=0.003 p=0.356 p=0.292 p=0.259 p=0.017 p=0.913 p=0.002

37

Table 9. Correlation coefficient (r) of the standardized growth parameters for the Turrumote reef (n=9) and Pinnacles reef (n=5) site master chronologies correlated against the grand master chronology (n=14), during the common 32- year period (1973-2004). Significant correlations are highlighted in yellow, and bold values are significant at p < 0.05.

GM Ann GM Ann GM Ann GM HD GM HD GM HD GM LD GM LD GM LD r = Ext Den Cal Ext Den Cal Ext Den Cal T Ann Ext 0.929 -0.688 0.812 -0.014 -0.320 -0.105 0.839 -0.647 0.709 p=0.00 1 p=0.001 p=0.001 p=0.938 p=0.074 p=0.567 p=0.001 p=0.001 p=0.001 T Ann Den -0.602 0.959 -0.297 -0.158 0.545 0.049 -0.495 0.928 -0.258 p=0.00 1 p=0.001 p=0.098 p=0.387 p=0.001 p=0.791 p=0.004 p=0.001 p=0.155 T Ann Cal 0.910 -0.407 0.933 -0.115 -0.159 -0.145 0.858 -0.355 0.833 p=0.00 1 p=0.021 p=0.001 p=0.529 p=0.384 p=0.429 p=0.001 p=0.046 p=0.001 T HD Ext 0.009 -0.259 -0.108 0.929 -0.515 0.798 -0.292 -0.240 -0.391 p=0.963 p=0.153 p=0.557 p=0.001 p=0.003 p=0.001 p=0.105 p=0.186 p=0.027 T HD Den -0.312 0.521 -0.158 -0.217 0.935 0.141 -0.199 0.318 -0.148 p=0.082 p=0.002 p=0.389 p=0.233 p=0.001 p=0.441 p=0.275 p=0.076 p=0.417 T HD Cal -0.065 -0.091 -0.128 0.925 -0.243 0.915 -0.353 -0.137 -0.437 p=0.725 p=0.620 p=0.486 p=0.001 p=0.179 p=0.001 p=0.048 p=0.456 p=0.012 T LD Ext 0.873 -0.569 0.801 -0.365 -0.105 -0.405 0.919 -0.540 0.830 p=0.00 1 p=0.001 p=0.001 p=0.040 p=0.566 p=0.022 p=0.001 p=0.001 p=0.001 T LD Den -0.540 0.913 -0.240 -0.209 0.400 -0.073 -0.426 0.970 -0.170 p=0.001 p=0.001 p=0.187 p=0.251 p=0.023 p=0.693 p=0.015 p=0.001 p=0.353 T LD Cal 0.810 -0.320 0.856 -0.500 -0.017 -0.518 0.908 -0.254 0.923 p=0.00 1 p=0.074 p=0.001 p=0.004 p=0.925 p=0.002 p=0.001 p=0.161 p=0.001 P Ann Ext 0.749 -0.336 0.786 -0.146 -0.243 -0.216 0.769 -0.247 0.779 p=0.00 1 p=0.061 p=0.001 p=0.426 p=0.180 p=0.235 p=0.001 p=0.172 p=0.001 P Ann Den -0.554 0.782 -0.320 -0.222 0.566 -0.004 -0.472 0.689 -0.283 p=0.001 p=0.001 p=0.074 p=0.222 p=0.001 p=0.981 p=0.006 p=0.001 p=0.117 P Ann Cal 0.640 -0.097 0.772 -0.239 -0.068 -0.240 0.691 -0.029 0.778 p=0.00 1 p=0.599 p=0.001 p=0.189 p=0.713 p=0.185 p=0.001 p=0.875 p=0.001 P HD Ext -0.201 0.033 -0.246 0.635 0.144 0.696 -0.370 0.003 -0.437 p=0.270 p=0.859 p=0.175 p=0.001 p=0.432 p=0.001 p=0.037 p=0.988 p=0.012 P HD Den -0.257 0.545 -0.064 -0.496 0.740 -0.224 -0.096 0.377 0.028 p=0.155 p=0.001 p=0.728 p=0.004 p=0.001 p=0.218 p=0.603 p=0.033 p=0.880 P HD Cal -0.277 0.268 -0.242 0.377 0.527 0.596 -0.371 0.126 -0.401 p=0.125 p=0.138 p=0.182 p=0.033 p=0.002 p=0.001 p=0.037 p=0.493 p=0.023 P LD Ext 0.735 -0.315 0.781 -0.278 -0.253 -0.356 0.793 -0.227 0.820 p=0.00 1 p=0.080 p=0.001 p=0.123 p=0.162 p=0.046 p=0.001 p=0.213 p=0.001 P LD Den -0.541 0.793 -0.278 -0.114 0.278 -0.019 -0.487 0.835 -0.238 p=0.001 p=0.001 p=0.123 p=0.533 p=0.124 p=0.916 p=0.005 p=0.001 p=0.190 P LD Cal 0.657 -0.168 0.766 -0.322 -0.196 -0.381 0.732 -0.072 0.819 p=0.00 1 p=0.359 p=0.001 p=0.072 p=0.283 p=0.031 p=0.001 p=0.694 p=0.001

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3.3 Relationships Between the Extension, Density, and Calcification Parameters by Correlation

The results of the correlation analysis (Table 10) show the relationships between the growth parameters; annual extension has a significant negative correlation relationship to bulk-density (r = -0.649, p < 0.001, n = 448) as well as a very strong and significant positive relationship to calcification with significant differences (r = 0.929, p < 0.001, n = 448). Annual bulk density exhibits a non-significant negative relationship to calcification (r = -0.337, p = 0.059, n = 448).

Annual extension was not significantly correlated to HD extension (r = -0.071, p < 0.699, Table 10), but was significantly correlated to LD extension (r = 0.942, p < 0.001). Annual calcification had a significant positive correlation with LD extension (r = 0.913, p < 0.001), and LD calcification (r = 0.930, p < 0.001).

Annual density showed a significant positive correlation with HD density (r = 0.610, p < 0.001), and LD density (r = 0.944, p < 0.001), also with Pearson’s r correlation coefficient. HD density (r = -0.362, p < 0.042), and HD calcification (r = 0.918, p < 0.001), both had a strong positive correlation with HD extension. The LD band typically has a stronger correlation with the annual band, due to it being the larger of the two subannual bands.

39

Table 10. Correlation coefficient matrix for growth parameters from 1973 to 2004, using 14 corals for 32 years per coral grand master chronology. Significant correlations are highlighted in yellow, and bold values are significant at p < 0.05.

r = Ext Den Cal HD Ext HD Den HD Cal LD Ext LD Den LD Cal Extension

Density -0.649 p=0.00 1 Calcification 0.929 -0.337 p=0.00 1 p=0.059 HD Ext -0.071 -0.197 -0.183 p=0.699 p=0.280 p=0.316 HD Den -0.338 0.610 -0.144 -0.362 p=0.058 p=0.001 p=0.431 p=0.042 HD Cal -0.168 0.036 -0.205 0.918 0.017 p=0.359 p=0.847 p=0.260 p=0.001 p=0.926 LD Ext 0.942 -0.539 0.913 -0.381 -0.188 -0.444 p=0.00 1 p=0.001 p=0.001 p=0.032 p=0.303 p=0.011 LD Den -0.582 0.944 -0.271 -0.194 0.390 -0.061 -0.479 p=0.00 1 p=0.001 p=0.134 p=0.288 p=0.027 p=0.742 p=0.006 LD Cal 0.849 -0.294 0.930 -0.487 -0.101 -0.526 0.950 -0.205 p=0.00 1 p=0.103 p=0.001 p=0.005 p=0.583 p=0.002 p=0.001 p=0.259

The correlation results for the annual parameters show that density has a non- significant negative relationship with calcification (r = -0.3368, p = 0.059, Figure 24, A) extension has a negative relationship with density also with significant differences (r = - 0.6492, p < 0.001, Figure 24, B) and extension showing a strong positive relationship to calcification with significant differences (r = 0.9289, p < 0.001, Figure 24, C).

The results for the HD parameters show that HD density has no significant relationship with HD calcification (r = 0.017, p = 0.93, Figure 25, A). HD extension has a negative relationship with HD density also with significant differences (r = -0.3616, p = 0.042, Figure 25, B), and HD extension showing a strong positive relationship to HD calcification with significant differences (r = 0.917, p < 0.001, Figure 25, C).

The results for the LD parameters show that LD density has a weak negative relationship with LD calcification (r = -0.205, p = 0.259, Figure 26, A). LD extension has a negative relationship with LD density as well as significant differences (r = -0.479, p =

40

0.005, Figure 26, B) and LD extension showing a strong positive relationship to LD calcification with significant differences (r = 0.950, p < 0.001, Figure 26, C).

A Annual Calcification vs Annual Density

2

) 1

- 1.5 r = -0.3368 yr

2 p = 0.059 - 1

cm 0.5 0 -0.5 -1

-1.5 Calcification Calcification (g -2 -1 -0.5 0 0.5 1 1.5 2 2.5 Density (g cm-3)

B Annual Density vs Annual Extension 3

2

)

3 -

1 cm 0 -1

Density Density (g r = -0.6492 -2 p = 0.001 -3 -2 -1.5 -1 -0.5 0 0.5 1 Extension (cm yr-1)

C Annual Calcification vs Annual Extension

1.5

) 1

- 1 r = 0.9289 yr

2 p = 0.001

- 0.5 cm 0 -0.5 -1

-1.5 Calcification Calcification (g -2 -2 -1.5 -1 -0.5 0 0.5 1 Extension (cm yr-1) Figure 24. Scatter plot graphs of annual growth parameters for master chronologies of n= 32 years (1973-2004), for each parameter. The correlation coefficient r quantitatively measures the strength and direction of the linear relationship. Significance is shown for each interaction, and a red star indicates a significant correlation.

41

A HD Calcification vs HD Density

0.6

)

1 -

yr 0.4

2 - 0.2 0 -0.2 -0.4 r = 0.017 -0.6 p = 0.930

HD Calcification HD Calcification (g cm -0.8 -1.5 -1 -0.5 0 0.5 1 1.5 HD Density (g cm-3)

B HD Density vs HD Extension 1.5

) 1

3 - 0.5

0

-0.5

HD Density HD Density cm (g -1 r = -0.3616 p = 0.042 -1.5 -1.0 -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1.0 HD Extension (cm yr-1)

C HD Calcification vs HD Extension

0.6

) 1 - r = 0.917

yr 0.4

2 p = 0.001 - 0.2 0 -0.2 -0.4 -0.6

HD Calcification HD Calcification (g cm -0.8 -0.8 -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1 HD Extension (cm yr-1)

Figure 25. Scatter plot graphs of HD growth parameters for 32 years (1973-2004), for each parameter. The correlation coefficient r quantitatively measures the strength and direction of the linear relationship. Significance is shown for each interaction, and a red star indicates a significant correlation.

42

A LD Calcification vs LD Density

1.5

) 1 - r = -0.205

yr 1 2

- p = 0.259 0.5 0 -0.5 -1 -1.5

LD LD Calcification cm (g -2 -1 -0.5 0 0.5 1 1.5 2 2.5 3 LD Density (g cm-3)

B LD Density vs LD Extension 2 r = -0.479 1.5

) p = 0.005

3 - 1 0.5 0 -0.5

LD Density LD Density cm (g -1 -1.5 -1.0 -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1.0 LD Extension (cm yr-1)

C LD Calcification vs LD Extension

1

) 1 - r = 0.950 yr 0.5

2 p = 0.001 - 0

-0.5

-1

-1.5

LD LD Calcification cm (g -2 -2 -1.5 -1 -0.5 0 0.5 1 LD Extension (cm yr-1)

Figure 26. Scatter plot graphs of LD growth parameters for 32 years (1973-2004), for each parameter. The correlation coefficient r quantitatively measures the strength and direction of the linear relationship. Significance is shown for each interaction, and a red star indicates a significant correlation.

43

3.4 Correlation Relationship of Time Series of SST to Coral Master Chronology Time Series Over 1973-2004

SST correlations for growth factors with a 3-month average (SST-H), 9-month average (SST-L), 12-month average (SST-A), minimum (SST-MN), maximum (SST- MX), AMO (SST-AMO), and NAO (SST-NAO), for the 32-year common period (1973- 2004), indicated weak and non-significant relationships between all growth parameters and SST variables (Table 11). However, when SST parameters were correlated against the each other, there was strong significant correlations with the majority of the parameters for the common 32-year period (Table 11). The only parameters that didn’t correlate were, SST-NAO with SST-AMO (r = -0.299, p < 0.096), and SST-NAO with SST-MX (r = -0.219, p < 0.230).

SST Parameters 30.0

29.0

28.0

C) ° 27.0

SST( 26.0

25.0

24.0 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004

SST-H SST-L SST-A SST-MN SST-MX

Figure 27. Shows SST parameters SST-H, SST-L, SST-A, SST-MN, and SST-MX.

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Table 11. Correlation of STDA grand master growth parameters with 3-month average (SST-H), 9-month average (SST-L), 12-month average (SST-A), min (SST-MN), max (SST-MX), AMO (SST-AMO), and NAO (SST-NAO), for the 32-year common period (1973-2004), bold and highlighted values significant at p < 0.05, n=32.

r = SST-H SST-L SST-A SST-MN SST-MX SST-AMO SST-NAO Annual Extension -0.216 0.013 -0.071 0.076 -0.074 -0.176 -0.123 p=0.248 p=0.873 p=0.700 p=0.679 p=0.688 p=0.335 p=0.502 Annual Density 0.139 0.011 0.093 0.004 0.092 0.215 0.118 p=0.329 p=0.853 p=0.611 p=0.981 p=0.615 p=0.236 p=0.519 Annual Calcification -0.149 0.067 0.008 0.115 0.018 -0.044 -0.108 p=0.531 p=0.727 p=0.967 p=0.530 p=0.922 p=0.812 p=0.555 HD Extension 0.018 0.054 0.007 0.150 -0.039 -0.023 -0.313 p=0.978 p=0.739 p=0.969 p=0.411 p=0.834 p=0.902 p=0.081 HD Density 0.005 -0.230 -0.126 -0.242 -0.021 0.087 0.210 p=0.737 p=0.152 p=0.492 p=0.183 p=0.909 p=0.637 p=0.250 HD Calcification -0.008 -0.043 -0.057 0.068 -0.082 -0.024 -0.271 p=0.935 p=0.798 p=0.758 p=0.713 p=0.656 p=0.897 p=0.133 LD Extension -0.155 0.019 -0.034 0.030 -0.006 -0.122 -0.022 p=0.442 p=0.855 p=0.852 p=0.869 p=0.973 p=0.508 p=0.904 LD Density 0.196 0.102 0.170 0.101 0.170 0.283 0.110 p=0.231 p=0.732 p=0.353 p=0.583 p=0.351 p=0.116 p=0.549 LD Calcification -0.067 0.093 0.061 0.076 0.099 0.020 -0.004 p=0.838 p=0.611 p=0.742 p=0.681 p=0.592 p=0.913 p=0.983 SST-H

SST-L 0.802 p=0.001 SST-A 0.883 0.976 p=0.001 p=0.001 SST-MN 0.478 0.795 0.746 p=0.006 p=0.001 p=0.001 SST-MX 0.896 0.780 0.851 0.417 p=0.001 p=0.001 p=0.001 p=0.018 SST-AMO 0.850 0.757 0.821 0.506 0.817 p=0.001 p=0.001 p=0.001 p=0.003 p=0.001 SST-NAO -0.396 -0.354 -0.369 -0.400 -0.219 -0.299 p=0.025 p=0.047 p=0.038 p=0.023 p=0.230 p=0.096

45

4. DISCUSSION:

4.1 Growth Parameters

This study found that growth (in terms of extension, density, and calcification) of the coral Orbicella faveolata was similar between Turrumote and Pinnacles reefs between years 1973 and 2004. However, the skeletal bulk density was greater on Pinnacles reef compared to Turrumote reef. Corals on Turrumote reef were less dense than those on Pinnacles reef, and had lower calcification, though the difference was not significant.

The subannual growth showed a similar trend with HD and LD density showing non-significant differences in growth on Pinnacles reef over Turrumote reef. Similar growth was measured with HD extension and HD calcification, while LD extension and LD calcification were slightly (non-significantly) larger on Pinnacles than on Turrumote.

Growth differences between reefs are likely related to their specific environmental conditions. Turrumote is slightly larger, while Pinnacles, due to its location on the shelf- edge and proximity to the watershed is relatively more exposed to natural and anthropogenic stress, such as currents, storms, boating activities (Garcia-Sais et al. 2008). Due to an east to west flowing current in the area of La Parguera, Pinnacles reef potentially had some low level exposure to toxins within sediment samples that were found in greater amounts and frequency toward the western side La Parguera, away from the study sites (Burke and Maidens 2004).

Puerto Rico, and the greater Caribbean have been experiencing disease and bleaching events since the early 1970’s and monitoring of these events increase when disease outbreaks occur. Turrumote reef has been affected multiple times by disease and bleaching events, and as much as 30-40% of live coral cover has been affected. Winter et al. (1989) conducted a 30-year study from 1966-1995, to find a correlation between elevated SST and major bleaching events in La Parguera during 1969, 1987, 1990, and

46

1995. Number of days at extremely elevated SST did correspond to the bleaching events, but it wasn’t enough to predict future events. This bleaching record may help to explain the large data anomaly observed in this study, for example during the year 1991 high coral density level were registered (Figure 16). This high density in 1991 was a characteristic shown on all 14 samples in both reefs, and was first seen in the X- radiographs as a possible environmental anomaly (Carilli et al. 2009).

The linear extension (cm yr-1) and calcification rate (g cm-2 yr-1) of the corals from Turrumote and Pinnacles reefs (La Parguera, Puerto Rico), were within the ranges of other studies in the region. However, the linear regression of calcification (g cm-2 yr-1) was comparatively lower than other similar long-term studies (Table 12). This could be possible to limiting factors at each specific reef and region. The smaller reefs tend to have an acute response to environmental signals, however they often get filtered out when measuring regional growth factors (Carricart-Ganivet 2004. The comparison between reefs also suggests that mean density values are decreasing in every new study, while mean calcification appears to vary randomly. More studies comparing trends over time and at different spatial scales are in need to test this hypothesis.

The amount of variability for linear extension (24%), bulk density (9%), and calcification (22%) growth of all 14 Orbicella faveolata corals, from both studied reefs in Puerto Rico, were slightly lower than previous studies in the Great Barrier Reef (31%, 9%, and 30% from 38 Porites cores) (Cooper et al. 2008). However, variability of both linear extension and calcification were slightly higher in Puerto Rico than in the Florida Keys (20%, 11%, and 17%, seven Orbicella corals) (Helmle et al. 2011), the Caribbean Sea (9%, 20%, and 13%, 46 Orbicella corals), and the Gulf of Mexico (5%, 13%, and 8%, 20 Orbicella corals) (Carricart-Ganivet 2004). Thus, coral examined from La Parguera, Puerto Rico tend to have similar variability in the density than other corals in the Caribbean Sea, but variability in linear extension and calcification are higher. The possible reasons for this can be attributed to a more variable environment. It is possible that La Parguera Puerto Rico, has been exposed to anthropogenic factors that are affecting the growth of the corals to a higher degree.

47

The analysis performed on the corals from Turrumote reef and Pinnacles reef, show that their growth patterns are similar. Regardless of the environmental variables that these corals are exposed to (from storms, to disease, to pollution and runoff, to boat traffic and over fishing), these two reefs appear to be affected by similar amounts of stress and pressure exerted upon their growth conditions, and the 1.2 km distance between them is non-significant factor.

4.2 Climate Change Relationships with Coral Growth

The results of this study indicate that SST measured in the Caribbean Ocean and the greater Atlantic Ocean, showed little to no significant correlation with the coral growth in Puerto Rico over a 32-year period (1973-2004). Specifically, correlation coefficients comparing SST to growth parameters showed almost no significant correlations for the master chronology of the 14 corals measured. Results from a previous study in the upper Florida Keys, USA, in which growth rates of Orbicella faveolata, over a period of 60 years (1937-1996), exhibited a significant increase in extension rates, significant decrease in skeletal density, and no significant change in calcification rates in response to SST (Helmle et al. 2011).

However, this pattern does not apply for the Great Barrier Reef, Australia, in which SST correlated with a decline in both calcification (14%) and linear extension (13%), from 1990 to 2008 in the massive Porites corals (De’ath et al. 2009). This shows that higher SSTs may affect coral growth differently and that regional or local environmental stressors may also have an additional effect.

Growth studies of massive Porites by Bessat and Buigues (2001) in Tahiti, as well as Lough and Barnes (1997) on the GBR, showed a positive relationship to calcification with the SST anomaly when compared to time. Calcification rates would increase 0.45 g/cm2/yr in Tahiti, 0.16 g/cm2/yr on the GBR, 0.08 g/cm2/yr in the Florida Keys (Helmle et al. 2011). Our study does not showed correlations between growth factors and SST anomaly, and suggest calcification rates increase of 0.96 g/cm2/yr in Puerto Rico, for an

48 increase in the annual SST by 1°C (Table 21). Showing that SST more accurately explains calcification rates in Tahiti and the GBR, and a 1°C increase in temperature will have a greater impact to calcification rates in the Florida Keys. It is possible that different species of corals react differently to higher SST.

Table 12. Linear regression of annual calcification and SST anomaly over time. Significant p values are highlighted.

Reference Location Regression Equation r value r2 value p value Years n Corals Helmle et al. 2011 Florida Keys y = 0.0834x + 0.9083 0.2666 0.0711 0.039 1937 - 1996 60 7 Bessat and Buigues 2001 Tahiti y = 0.4503x + 1.2214 0.5700 0.3249 < 0.001 1958 - 1990 33 1 Lough and Barnes 1997 GBR y = 0.1594x + 1.5233 0.5382 0.2897 < 0.001 1906 - 1982 77 10

This Study 2017 Puerto Rico y = -0.8929x + 0.0324 0.0161 0.0003 < 0.930 1973-2004 32 14

SST have been shown to have a negative effect on coral growth, however with a lack of correlation to SST with this study, other potential negative influences could have had an effect. Converging currents throughout the Caribbean and Southern Atlantic Ocean create a variability in the nutrient content that changes on an interannual to multidecadal time scale (Kilbourne et al. 2007). Nutrient and pollutant effects from freshwater runoff (Lough and Barnes 2000; Moyer and Grottoli 2011). Another option could be a possible correlation with ocean acidification. Utilizing ocean acidification proxy data could help to understand stresses facing coral growth in that area (Lough and Barnes 1997; Doney et al. 2009; Manzello et al. 2015).

This study explored long-term growth records in an area that has not previously looked at multi-decadal studies against the effects of sea surface temperatures, and it helps to fill in gaps in knowledge of the overall health of coral throughout the Caribbean. Future studies should further explore areas in the Caribbean that can compare sea surface temperature effects, and ocean acidification, to gauge the anthropogenic status of coral reef health.

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5. CONCLUSIONS:

Coral growth measurements of 14 Orbicella faveolata core samples, from La Parguera, Puerto Rico showed that linear extension and calcification rates were not significantly greater for Turrumote reef that for Pinnacles reef, while bulk density was non-significantly greater for Pinnacles reef. Density was the most conservative of the growth factors CV = 9%, while extension was CV = 22%, and calcification was CV = 20%. Extension had a negative relationship to density, density was slightly negative to calcification, and extension had a very positive relationship with calcification.

Annual and subannual growth data for each parameter were assembled into master chronologies from Turrumote reef, Pinnacles reef, and then further assembled into a grand master chronology. In general, the master chronologies of Turrumote reef were significantly correlated to those of Pinnacles reef for each parameter.

The time series chronologies showed that extension was significantly negatively correlated for the annual, HD, and LD bands. There was a negative correlation between density and calcification for the annual and LD bands, no relationship with the HD band. Extension was significantly highly correlated for the annual, HD, and LD bands.

No significant correlations were found with growth parameters and SST for the 32-year time series from 1973-2004.

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