Growth and Survival of Acropora Cervicornis · Acropora cervicornis fragments were transplanted seasonally (every 3 months) near Key Largo, Florida (patch reef-Admiral Reef and fore

GROWTH AND SURVIVAL OF TRANSPLANTED ACROPORA CERVICORNIS IN

RELATION TO CORAL REEF RESTORATION

by

GEOFFREY CLAYTON CHILCOAT

(Under the Direction of William K. Fitt)

ABSTRACT

Acropora cervicornis fragments were transplanted seasonally (every 3 months) near Key Largo,

Florida (patch reef-Admiral Reef and fore reef-Little Grecian Reef) and Lee Stocking Island,

Bahamas (patch reef) in order to determine the relationship between the size of the fragment,

seasonal growth rate, and survival for the possibility of restoration of this species through

transplantation. Mass per unit length per day (g/mm/d), the total mass accretion per length

extension (g/mm), and the linear extension per day and buoyant weight per day was calculated

for each fragment. The number of branches generated from each fragment was recorded. These

parameters will be compared between the two sites in Florida and between the Bahamas site and

Florida sites. The recovery rates of scientifically produced scars or lesions were investigated in

the Caribbean reef coral Montastrea faveolata. Artificial lesions on Montastrea faveolata filled

with epoxy took approximately twice the recovery time as those allowed to recovery without the

use of filler compounds; however differences in growth rates were only seen in the first three

months.

INDEX WORDS: Acropora cervicornis, Acropora prolifera, Growth rates, Survival,

Transplantation, Regeneration, Calcification, Recovery, Montastrea faveoloata



by


Bachelors of Forest Resources, The University of Georgia, 1997

A Thesis Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment

of the Requirements for the Degree

MASTER OF SCIENCE

ATHENS, GEORGIA

2004

© 2004

Geoffrey Clayton Chilcoat

All Rights Reserved



by


Major Professor: William K. Fitt Committee: James Porter

Karen Porter

Electronic Version Approved:

Maureen Grasso Dean of the Graduate School The University of Georgia August 2004

ACKNOWLEDGEMENTS

The work was funded by the National Undersea Research Program (UNCW in Key

Largo, Florida and the Caribbean Marine Research Center on Lee Stocking Island in the

Bahamas) and the National Science Foundation (9203327,9702032,9906976). We would like to

thank Dr. Steve Miller, Otto Rutten, Mike Birns and others, for continued logistical support from

the NURC/UNCW Day-boat Program in Key Largo. Drs. John Marr, Tom Bailey, Steve Jury, as

well as Brian Kakuk, Jeremy, Ester, both Kerleens, and Craig and Tara Dahlgren of CMRC Lee

Stocking Island Bahamas. Thanks to Dan Thornhill, Mark Warner, Todd LaJuenesse, Tom

Shannon, Nathan Jess, and Peter Anziano for field assistance. Permitting in the United States

thanks to Harold Hudson, John Hallace and others from NOAA Florida Keys.

iv

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS........................................................................................................... iv

LIST OF TABLES........................................................................................................................ vii

LIST OF FIGURES ..................................................................................................................... viii

CHAPTER

1 INTRODUCTION .........................................................................................................1

References .................................................................................................................5

2 SEASONAL BRANCHING PATTERNS IN TRANSPLANTED REEF-BUILDING

CORALS ACROPORA CERVICORNIS AND ACROPORA PROLIFERA ..............7

Abstract .....................................................................................................................8

Introduction ...............................................................................................................9

Methods ...................................................................................................................10

Results .....................................................................................................................12

Discussion ...............................................................................................................15

Acknowlegdments ...................................................................................................17

References ...............................................................................................................18

3 EFFECT OF SEASON AND CORAL SIZE ON THE GROWTH RATE OF

TRANSPLANTED ACROPORA CERVICORNIS..................................................43

Abstract ...................................................................................................................44

Introduction .............................................................................................................45

Methods ...................................................................................................................48

Results .....................................................................................................................50

v

Discussion ...............................................................................................................52

References ...............................................................................................................56

4 EFFECT OF SEASON AND CORAL SIZE ON THE SURVIVAL OF

TRANSPLANTED ACROPORA CERVICORNIS..................................................71

Abstract ...................................................................................................................72

Introduction .............................................................................................................73

Methods ...................................................................................................................74

Results .....................................................................................................................76

Discussion ...............................................................................................................77

References ...............................................................................................................79

5 SCIENTIFICALLY PRODUCED LESIONS IN REEF CORALS: SCARRED FOR

LIFE?.......................................................................................................................86

Abstract ...................................................................................................................87

Introduction .............................................................................................................88

Methods ...................................................................................................................89

Results .....................................................................................................................90

Discussion ...............................................................................................................91

References ...............................................................................................................94

vi

LIST OF TABLES

Page

Table 3.1: Original Transplant Size v. growth Length (mm) p=.05 significance..........................69

Table 3.2: Original Transplant Size v. Weight Growth (grams) p=.05 significance .....................69

Table 5.1: Percent recovery of the epoxy-filled lesion from the initial damage (Bahamas). ........95

Table 5.2.: Percent recovery of the control lesions from the initial damage (Bahamas). ..............96

vii

LIST OF FIGURES

Page

Figure 2.1a: All Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reef),

A cervicornis, Average new branches added since previous sampling period..............19

Figure 2.1b: All Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reef),

A. cervicornis, Total branches since initial transplantation...........................................19

Figure 2.2: All Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reefs)

A. cervicornis, A. prolifera Average new branches per day since previous sampling

period.............................................................................................................................20

Figure 2.3: Total Branches Since Initial Transplantation Bahamas (LSI) (A. cervicornis,

A.prolifera) Little Grecian Reef (LG) and Admiral Reef (ADM), Florida (A.

cervicornis)....................................................................................................................20

Figure 2.4(a-i): ADM AUG 1999, A. cervicornis Original transplant size (length) v new branches

since previous sampling period .....................................................................................21

Figure 2.5(a-i): ADM AUG 1999, A. cervicornis Original transplant size v total branches .........22

Figure 2.6(a-h): ADM Nov 1999, A. cervicornis Original transplant size (length) v new branches


Figure 2.7(a-h): ADM Nov 1999, A. cervicornis Original transplant size (length) v. total

branches.........................................................................................................................24

Figure 2.8(a-g): ADM March 2000, A. cervicornis Original transplant size (length) v new

branches since previous sampling period......................................................................25

Figure 2.9(a-g): ADM March 2000, A. cervicornis Original transplant size (length) v total

branches.........................................................................................................................26

viii

Figure 2.10(a-i): LG June 1999, A. cervicornis Original transplant size (length) v total branches


Figure 2.11(a-i): LG june 1999, A. cervicornis Original transplant size (length) v average new


Figure 2.12(a-i): LG Nov 1999, A. cervicornis Original transplant size (length) v total branches29

Fig. 2.13(a-g): LG Nov 1999 A. cervicornis Original transplant size (length) v average new


Fig. 2.14(a-h): LG Jan 1999 A. cervicornis Original transplant size (length) v average new


Fig. 2.15(a-h): LSI Jan 1999 A. cervicornis Original transplant size (length) v total branches ....32

Fig. 2.16(a-h): LSI May 1999 A. cervicornis Original transplant size (length) v average new


Fig. 2.17(a-h): LSI May 1999 A. cervicornis Original transplant size (length) v total branches ..34

Fig 2.18(a-i): LSI Feb 99 A. prolifera Original transplant size (length) v. total branches ............35

Fig. 2.19(a-i): LSI Feb 1999 A. cervicornis Original transplant size (length) v average new


Fig. 2.20a: ADM Aug99 A. cervicornis New branches/day v. percent growth length/day, percent

growth weight/day (95%CI) ..........................................................................................37

Fig. 2.20b: ADM Nov99 A. cervicornis New branches/day v. percent growth length/day, percent

growth weight/day (95%CI) ..........................................................................................37

Fig. 2.20c: ADM March00 A. cervicornis New branches/day v. percent growth length/day,

percent growth weight/day (95%CI) .............................................................................38

ix

Fig. 2.21a: LG Jun99 A. cervicornis New branches/day v. percent growth length/day and percent

growth weight /day........................................................................................................38

Fig. 2.21b: LG Nov99 A. cervicornis New branches/day v. percent growth length/day and

percent growth weight /day ...........................................................................................39

Fig. 2.21c: LG may00 A. cervicornis New branches/day v. percent growth length/day and

percent growth weight /day ...........................................................................................39

Fig. 2.22a: LSI, Bahamas Jan99 A. cervicornis New branches/day v percent growth length,

percent growth weight ...................................................................................................40

Fig. 2.22b: LSI, Bahamas May99 A. cervicornis New branches/day v percent growth length,


Fig. 2.22c: LSI, Bahamas Aug99 A. cervicornis New branches/day v percent growth length,


Fig. 2.23: All Sites (A. cervicornis, A. prolifera) Total grams/total mm.......................................41

Fig. 2.24: Little Grecian Reef June 1999 Transplants, New branches/day, total grams/mm ........42

Fig. 2.25: Little Grecian Reef, A. cervicornis total grams/total length (g/mm) ............................42

Fig. 3.1a: Admiral Reef, A. cervicornis Percent growth (length, mm/day) of original size for each

sampling period .............................................................................................................59

Fig. 3.1b: Admiral Reef , A. cervicornis Growth length (mm/day) for each sampling period......59

Fig. 3.1c: Admiral Reef, A. cervicornis Percent growth (weight, g/d) of original size for each

sampling period .............................................................................................................60

Fig. 3.1d: Admiral Reef, A. cervicornis Growth weight (g/d) for each sampling period..............60

Fig. 3.2a Admiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length

(mm/d) for Aug 1999 transplants ..................................................................................61

x

Fig. 3.2b: Admiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length

(mm/d) for Nov 1999 transplants ..................................................................................61

Fig. 3.2c: Admiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length

(mm/d) for March 2000 transplants...............................................................................62

Fig. 3.2d: Admiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length

(mm/d) for May 2000 transplants..................................................................................62

Fig. 3.2e: Admiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length

(mm/d) for Aug 2000 transplants ..................................................................................63

Fig. 3.3a: Little Grecian Reef Percent growth length (mm/d) of original size for each sampling

period.............................................................................................................................63

Fig. 3.3b: Little Grecian Reef Total Growth (length, mm/day) for each sampling period............64

Fig. 3.3c: Little Grecian Reef Percent growth weight (grams/day) of original size for each

sampling period .............................................................................................................64

Fig. 3.3d: Little Grecian Reef, A. cervicornis Total growth weight (grams,day) for each sampling

period.............................................................................................................................65

Fig. 3.4a: Lee Stocking Island, A. cervicornis Percent growth (length, mm/day) of original size

for each sampling period ...............................................................................................65

Fig. 3.4b: Lee Stocking Island, A. cervicornis Growth length (mm/day) for each sampling

period.............................................................................................................................66

Fig. 3.4c: Lee Stocking Island, A. cervicornis Percent growth (weight, g/day) of original size for

each sampling period.....................................................................................................66

Fig. 3.4d: Lee Stocking Island, A. cervicornis Growth weight (g/day) for each sampling

period............................................................................................................................ 67

xi

Fig. 3.5: All Sites (A. cervicornis, A. prolifera) Total grams/total mm.........................................67

Fig. 3.6: Lee Stocking Island, Normans Pond (daily averages) 4/20/96-7/11/02 *no data for Jan

99-Oct 99 data from <http://www.cmrc.org/lsi_seawater_temp.htm> .........................68

Fig. 3.7: Molasses Reef Florida 1/98-9/02 (Daily Average Sea Temp) C-Man Station (CRAMP)

data from <http://www.coral.noaa.gov/seakeys/hist_data.shtml> ................................68

Fig. 4.1: Tranplant Survival (Acropora cervicornis) Little Grecian Reef, FLA (LG), Lee

Stocking Island, Bahamas (LSI), Admiral Reef , FLA (Adm)......................................81

Fig. 4.2: LSI percent survival vs size for all transplants (Jan 1999-May 2001) ............................81

Fig. 4.3a: Survival of corals in relation to initial size transplanted Little Grecian Reef June 1999

Tranplants ......................................................................................................................82

Fig 4.3b Survival of corals in relation to initial size transplanted Little Grecian Reef Nov 1999

Tranplants ......................................................................................................................82

Fig 4.3c Survival of corals in relation to initial size transplanted Little Grecian Reef May 2000

Tranplants ......................................................................................................................83

Fig 4.3d Survival of corals in relation to initial size transplanted Little Grecian Reef AUG 2000

Tranplants ......................................................................................................................83

Fig 4.3e Survival of corals in relation to initial size transplanted Little Grecian Reef Nov 2000

Tranplants ......................................................................................................................84

Fig. 4.4a Survival at first sampling period LSI, Bahamas .............................................................84

Fig. 4.4b Florida (Admiral) Survival at first sampling period (Number of days in period in

parenthesis)....................................................................................................................85

Fig. 5.1a Recovery of sampling scars Montastrea faveolata LSI, Bahamas Aug 2000 (initial

lesion) - June 2000 ........................................................................................................97

xii

Fig. 5.1b Recovery of sampling scars Montastrea faveolata with linear regression LSI, Bahamas

Aug 2000(initial lesion) - June 2000.............................................................................98

Fig. 5.2 Admiral Reef, Florida, Montastrea faveolata MF1, MF2, MF4 March 2001 (initial

damage) - Aug 2001 ......................................................................................................99

Fig. 5.3 Monthly growth rates for each season Montastrea faveolata, LSI, Bahamas................100

xiii

CHAPTER 1

INTRODUCTION

1

Percent cover of the branching coral Acropora cervicornis (Lamarck) has declined in the

Caribbean. This decline is has not been recorded in recent history and some evidence shows that

it is unprecedented in the Pleistocene and Holocene eras (Greenstein, 1996, Greenstein, 1998).

Aronson and Precht recently documented an event where A. cervicornis died in a lagunal area

and was completely replaced with another genus; although this switch in species was short lived,

it had not occurred in the past 3,500 years (Aronson, 1997, Aronson, 1998).

Many factors have been implicated in the rapid decline of the Acroporids in the Caribbean

region. Increased hurricanes, decreased herbivore densities including Diadema antillarium die-off

(Lessios et al, 1984) leading to less larval recruitment, increased prevalence of diseases on acroporids

such as white band disease (Ritchie, 1998) and the newly discovered common fecal enterobacterium

of humans, Serratia marcescens (Patterson et al, 2002), as well as increased anthropogenic stressors

such as urban and non-urban run off, untreated sewage, and use of pesticides have been implicated in

the demise of reef corals.

Global warming and increased incidence of El Nino events may also contribute to

physiological stress of reef corals. El Nino conditions can also elevate surface sea temperatures

(SST’S) and have been implicated in the global decline of coral reefs via world-wide bleaching

events involving the loss of symbiotic algae. Woodley (1992) suggests that in Jamaica the lack of

hurricanes associated with long-term weather patterns between 1944 and 1980 resulted in stands of

Acropora cervicornis that were atypical and at one extreme of a variable condition.

2

Changes in major reef organisms may have some impact on the survival of corals. The recent

demise of the herbivorous sea urchin Diadema antillarum in the Caribbean has promoted algal

overgrowth and may hinder recruitment of juvenile corals. Although correlations of algae cover and

urchin densities seem easily tested, no studies have correlated increased algae and decreased

recruitment of hard corals. Others hypothesize that land based pollution, sediments and runoff have

been a primary cause of coral mortality, while Shinn et al (2000) believes African dust brings

antigens and more sediments contributing to the demise of reefs..

Corallivory due to two gastropods, Coralliophila abbreviate and C. caribea, as well as the fire

worm Hermodice carunculata, has also been shown to produce significant mortality of A.

cervicornis. However, since these animals have been present for a long time it has been assumed that

if the coral is healthy, rapid tissue growth of these species can keep pace with the corallivores.

The loss of A. cervicornis has been so great in the Florida Keys that the species has been

recommended for a spot on the endangered species list in 1999, but so far has not been listed (Diaz-

Soltero 1999).

In response to the demise of acroporids in the Caribbean, several questions can be asked in

regards to basic physiology of A. cervicornis. The answers to these research questions may directly

influence the restoration techniques used to successfully transplant and restore A. cervicornis to

damaged areas. Although some research on skeletal growth in A. cervicornis shows diel variation

(Chalker and Taylor 1978) and monthly measurements of growth indicate variation throughout the

year (Gladfelter, 1983, Gladfelter, 1984), several questions about growth patterns remain

unanswered. No studies have investigated the survivorship of transplants with the respect to the time

of year of the initial transplant, and few studies have attempted to correlate calcification and

extension rate to the time of year.

3

In this thesis, experiments were conducted to see if A. cervicornis can be transplanted for

restoration purpose. The first chapter is an introduction and gives a back round of current and

historical literature and problems related to the genus of this species. The second of four chapters

looks at branching patterns of transplanted A. cervicornis and A. prolifera. The third chapter consists

of data on the linear growth and buoyant weight in relation to seasonality of growth. By making

quarterly measurements of buoyant weigh and length, correlations of calcification and extension to

different seasons were made. Differences in growth rates in the Bahamas and the Florida Keys are

explored. Chapter 3 also investigates the best size of coral to transplant to achieve the highest

growth. The fourth chapter contains information on the survival of A. cervicornis transplants. This

chapter relates the time of the year and size of coral fragment to overall survival. The fifth and final

chapter contains information on the recovery rate of artificially produced scars on the boulder coral

Montastrea faveolata in Florida and Bahamas.

4

REFERENCES

Aronson, R. B. and W. F. Precht (1997). “Stasis, biological disturbance, and community

structure of a Holocene coral reef.” Paleobiology 23(3): 326-346.

Aronson, R. B., W. F. Precht, et al. (1998). “Extrinsic control of species replacement on a

Holocene reef in Belize: The role of coral disease.” Coral Reefs 17(3): 223-230.

Chalker, B. E. and D. L. Taylor (1978). “Rhythmic Variations in Calcification and

Photosynthesis Associated with Coral Acropora-Cervicornis-(Lamarck).” Proceedings of the

Royal Society of London Series B-Biological Sciences 201(1143): 179-189.

Diaz-Soltero, H. (1999). “Endangered and threatened species; a revision of candidate

species list under the Endangered Species Act.” Fed Regist 64(210): 33466-33468.

Gladfelter, E. H. (1983). “Spatial and Temporal Patterns of Mitosis in the Cells of the

Axial Polyp of the Reef Coral Acropora-Cervicornis.” Biological Bulletin 165(3): 811-815.

Gladfelter, E. H. (1984). “Skeletal Development in Acropora-Cervicornis 3. A

Comparison of Monthly Rates of Linear Extension and Calcium Carbonate Accretion Measured

over a Year.” Coral Reefs 3(1): 51-57.

Greenstein, B. J., H. A. Curran, et al. (1998). “Shifting ecological baselines and the

demise of Acropora cervicornis in the western North Atlantic and Caribbean Province: a Pleistocene perspective.” Coral Reefs 17(3): 249-261.

Greenstein, B. J. and H. A. Moffat (1996). “Comparative taphonomy of modern and

Pleistocene corals, San Salvador, Bahamas.” Palaios 11(1): 57-63.

Hayes, J. A. (1990). “Distribution, movement and impact of the corallivorous gastropod

Coralliophila abbreviata (Lamarck) on a Panamanian patch reef.” Journal of Experimental

Marine Biology and Ecology 142(1-2): 25-42.

5

Lessios, H. A., M. J. Garrido, et al. (2001). “Demographic history of Diadema antillarum,

a keystone herbivore on Caribbean reefs.” Proceedings of the Royal Society of London Series B-

Biological Sciences 268(1483): 2347-2353.

McClanahan, T. R. and N. A. Muthiga (1998). “An ecological shift in a remote coral atoll

of Belize over 25 years.” Environmental Conservation 25(2): 122-130.

Patterson, K. L., J. W. Porter, et al. (2002). “The etiology of white pox, a lethal disease of

the Caribbean elkhorn coral, Acropora palmata.” Proceedings of the National Academy of

Sciences of the United States of America 99(13): 8725-8730.

Rinkevich, B (2002). "The Branching coral Stylophora pistallata: Contibution of genetics

in shaping colony landscape." Israel Journal of Zoology 48 (1): 71-82

Ritchie, K. B. and G. W. Smith (1998). “Type II white band disease.” Revista de Biologia

Tropical 46(Suppl. 5): 199-203.

Shinn, E. A., G. W. Smith, et al. (2000). “African dust and the demise of Caribbean coral

reefs.” Geophysical Research Letters 27(19): 3029-3032. Woodley, J. D. (1992). “The Incidence of Hurricanes on the North Coast of Jamaica since

1870 - Are the Classic Reef Descriptions Atypical.” Hydrobiologia 247(1-3): 133-138.

6

CHAPTER 2

SEASONAL BRANCHING PATTERNS IN TRANSPLANTED REEF-BUILDING CORALS

ACROPORA CERVICORNIS AND ACROPORA PROLIFERA1

1Chilcoat, G.C., and W.K. Fitt. To be submitted to Coral Reefs.

7

ABSTRACT

Branching patterns of the staghorn coral Acropora cervicornis and related Acropora prolifera,

were monitored quarterly for three years in the Florida Keys and the Bahamas. Highest rates of

branching were observed during the late spring/early summer when growth rates were highest.

Branching, as well as linear growth and accretion, slowed and often completely stopped during

the late summer and early fall when seasonal temperatures were highest. Branching occurred

earlier and more frequently in larger (> 6 cm) pieces of coral, compared to smaller pieces.

Acropora prolifera had higher extension and branching rates compared to A. cervicornis;

however, ratios of linear extension to accretion (g/mm) between species were not statistically

different.

Key Words: Acropora, branching, staghorn coral, restoration

8

INTRODUCTION

There have been few studies on branching of cnidarians, especially corals. In contrast,

the literature on branching patterns in plants is quite extensive. In plants branching patterns are

primarily controlled by genetics, but can be influenced by environmental parameters such as

wind and light. Plant literature states that “All trees of a given species look alike because they

are all conforming to a given set of branching ‘rules’, but each individual will have a unique

location and history.” Thus, a plant of a certain species will look like that species, but local

environmental influences can alter the growth form to a certain extent (Bell 1997). This seems

to be true in branching cnidarians, where species look alike but vary slightly according to depth,

flow and light availability. For instance, Porites divercata almost always ends in tips that have

two branches where as A. prolifera and A. cervicornis seem to have a random single points

ending with a single apical polyp. Different approaches to analyzing colonial organization

amongst invertebrate growth forms have revealed an intrinsic order to branched forms, but there

is no model system for the study of the genetic impact on the architecture of branching modular

organisms (Rinkevich 2002).

The main hypothesis on how factors other than genetic control modulate branching

patterns in cnidarians involves the simple minimum availability of nutrient resources; branching

occurs when a threshold is met. In fact reactive oxygen species (ROS) have recently been found

near the branch sites of hydroids, which are associated with high mitochondrial density and

activity, and are thought to be involved in initiating branching in some species (Blackstone, Neil

2003). Another set of hypotheses holds that external factors control branching, including certain

bacterial types growing on the outside of the colony (Suobodas pers.comm.).

9

METHODS

Transplantation (small pieces artificially fragmented from a parent colony) experiments

were conducted on a patch reef due north of Lee Stocking Island (LSI), Bahamas and at two sites

off Key Largo, Florida; Admiral Reef (ADM) (N 25° 02.732’ W 080° 23.675’) a 1-3m deep

inshore patch reef and Little Grecian Reef (LG) (N 25°07.164’ W 080° 18.107’) a 3m deep

offshore reef crest. Artificially broken fragments of Acropora cervicornis were transplanted four

times corresponding to the seasons and monitored quarterly between 1999 and 2001.

In the Bahamas, coral colonies were transplanted January, May, and August of 1999 and

March of 2000. Florida coral colonies were transplanted first at Little Grecian Reef on June 17,

1999, then both Little Grecian and Admiral Reef in August and November 1999, just Admiral

reef on March 12, 2000 (Little Grecian had 6-8ft seas), then both sites in May and August 2000

and finally just Little Grecian Reef on November 29, 2000. Due to its scarcity, Acropora

prolifera was transplanted only once in the Bahamas on February 21, 1999 and monitored

quarterly until May 2001.

Fifteen A. cervicornis and A. prolifera fragments, ranging in size from 25 mm to 200

mm, were glued to PVC pipe and attached to cement bricks adjacent to the reef they were

collected. Transplants were glued into a 2″ section of ½ ″ PVC with Oately® epoxy putty. The

weight of the epoxy and PVC pipe were measured so that the initial weight of the transplant

could be determined.

Fragments were weighed to the nearest .001 gram using the Buoyant Weight method in

reef water. Temperature (25oC) and salinity (36 ppt) were held constant in the weighing

apparatus for every season. Length was measured to the nearest millimeter using vernier

10

calipers. Branching was analyzed by either calculating the total number of branches (total)

arising since the original transplant date, or the number of branches added (new) since the

previous sampling date. Initially almost all fragments (ca. 75 per site) had a single axial corallite

(no radiating branches from a single stick) and new branches produced from this single stick

were tallied over time. The only exceptions were one transplant from LG 6/99 which started

with three branches (one main stick with two radiating branches), three transplants from LG

11/99 which started with two branches, two transplants from ADM 3/00 which started with two

branches and two that started with three branches. Growth was measured from the top of the

PVC or where the live tissue starts at the base. Any dead parts, such as tips or areas around the

base were predation is often seen, were subtracted from the total length and not calculated in the

results.

Graphs were generated in Excel, with 95% confidence intervals. For ease of comparison,

the initial data point for total cumulative branch data was plotted as one branch, such that the

first branch growing out from the transplanted stick was designated as two total branches. New

branch data included only branches originating since the previous sample time. To facilitate

comparison of data, not all transplant times were plotted on all graphs, and error bars were left

off some graphs.

11

RESULTS

The number of new branches of Acropora cervicornis added since the previous sampling

period was typically highest during the spring, and dramatically decreased by the late summer and

fall, with some branching occurring in the winter (Figure 2.1a). This seasonal pattern resulted in a

step-wise accumulation of total branches in A. cervicornis that was similar at all sites in the Florida

Keys and the Bahamas (Figure 2.1b). The number of new branches added per day was typically

higher in every season in A. prolifera as compared to A. cervicornis (Figure 2.2). This was also

true for the total number of branches (Figure 2.3).

Branching rates were generally higher in larger-sized transplants compared to smaller

pieces of coral, but this phenomenon varied with transplant time, site and species (Figures 2.4

(a-i) - 2.13(a-g)). The basic pattern occurred at each site, and for both Acropora cervicornis and

Acropora prolifera: number of new branches and total cumulative branches were positively

correlated with transplant size during the initial sampling period(s), but only if the initial

transplant was made in the period November through May, the seasonal period of rapid growth

and branching (Figures 2.4-2.13). It should be noted that while over 50% of these correlations

were significant (p< 0.05), the data is quite variable. In fact, the positive correlation tended to

break down with time as other variables, some obvious such as predation and storm breakage,

influenced the data. Transplants begun just prior to or during the warmest seasonal seawater

temperatures (May – October) almost always showed low or no branching during the initial

warm season, but usually exhibited positive branching correlations with size the following

spring-early summer (March – July)(Figure 2.4a-i). Transplants made during August 1999 in the

12

Bahamas and August 2000 in the Florida Keys resulted in 90% mortality from high water

temperatures (Chapter 4) and branching data are not included here.

Branching, in terms of new branches produced since the last measurement, as well as

total cumulative branches, was generally related to growth of the coral; higher rates of branching

occurred during the spring and early summer (March-July) when growth was highest (Figure

2.15a-i). Branching in Acropora prolifera exhibited a similar relationship with growth (Figure

2.17a).

The weight per unit length data (Figure 2.16a) gives an indication of the average

thickness of branches, an attribute that has been correlated with wave exposure in branching

corals (Bottjer, D. J. 1980). All transplants from the Bahamas site exhibited higher weight to

length ratios than transplants from the more protected inshore patch reef site from the Florida

Keys (Figure 2.16a). The highest ratios were measured from the earliest transplants taken from

the reef-crest-reef portion of Little Grecian Reef (2-3 m deep) off Key Largo; the second set of

transplants taken from the reef-flat portion of Little Grecian Reef had ratios similar to the

Bahamas corals, while all subsequent transplants from Little Grecian Reef were taken from the

adjacent channel (3-4m deep) that experiences less wave exposure. Several of the data sets show

the expected decreased weight/length ratio during the spring-time season of rapid branching, as

small new branches are not as thick and heavy as mature branches (e.g. Figure 2.16b). Higher

branching rates in

transplanted Acropora cervicornis were observed at sites with lower wave exposure (Admiral

Reef compared to Bahamas site, later transplants at Little Grecian reef compared to the earliest

two transplant times as detailed above) (Figure 2.1a).

13

Branching rates of Acropora prolifera were initially about an order of magnitude greater

than those of Acropora cervicornis, with similar but less pronounced seasonal peaks in early

summer (Figure 2.17a-e). However, the relationship of branching with growth in A. prolifera

appears to be quite different that with A. cervicornis. Maximum growth typically occurs during

periods of low branching, suggesting a trade-off between the use of resources for extension and

accretion (Figure 2.17a-e). In addition, branches of A. prolifera appear to be thicker

(grams/mm/day) during periods of low branching (Figure 2.17a-e). The resulting cumulative

growth of A. prolifera is a seasonal sinusoidal increase in both total number of branches as well

as coral weight and length (Figure 2.17a-e).

Late summer-early fall decreases in branching rates were more severe in 1999 compared

to 2000, as seen in both A. cervicornis from Florida and in A. cervicornis and A. prolifera from

the Bahamas in the November data point (August-November period, Figure 2.3a). Temperature

data from various parts of the Caribbean confirm higher mean seasonal seawater temperatures in

1999 compared to 2000, with sensors at Molasses Reef in Florida and Bahamas reaching about

310C September 1999 compared to seasonal maximums of about 300C in September of 2000

(Figure 2.18-2.19).

Preliminary genetic analysis showed that Acropora cervicornis populations collected

from the channel environments adjacent to Little Grecian Reef off Key Largo are one clone with

an introgressed genotype, distinct from A. cervicornis from nearby Alligator Reef (S. Vollmer,

pers com). In contrast, the thicket of Acropora cervicornis off Norman’s Pond Cay, Bahamas,

contained at least two distinct genotypes (S. Vollmer, pers comm.), but it was not possible to tell

those types apart by morphology, growth form, or location on the reef. No genetic analysis of

the A. cervicornis growing on Admiral Reef off Key Largo has been completed to date.

14

DISCUSSION

Branching rates of Acropora cervicornis exhibited predictable seasonal variation, with

maximum average rates occurring in the spring and early summer of most years at each site, and

minimum branching rates occurring during the late summer-early fall (Figure 2.4). Maximum

branching rates correlate with maximum growth documented for A. cervicornis (27-29oC)

(Figure 2.4). Minimum branching rates correlated with minimum growth rates (Figure 2.4),

observed in this study in the late summer-early fall (August – November) when seawater

temperatures exceeded 30oC. Branching and growth rebounded in the fall of each year, but were

still low in the winter months when cold fronts often brought seawater temperatures down to 18-

23°C (cf. Hudson 1966), before peaking again in the spring or early summer (Figures 2.18 and

2.19).

If branching rates are strictly under the control of temperature, then winter reductions in

branching rate would theoretically be less in the Bahamas where minimum temperatures average

24-25oC, compared to Key Largo in the Florida Keys, where minimum temperatures are often

less than 20oC for a month or more each winter (Figure 2.7). However, the data in this study

(Fig 2.1) show few differences between sites during the winter; virtually all winter branching

rates are above zero (0.2-1.2 new branches per colony). There were also few differences in

branching rates between sites during the warmest part of the year (August-November sample

times); however, branching data during this season was at or near zero in 1999, but averaged 0.4-

1.0 new branches per colony in 2000 (Figure 2.2). Clearly there was something happening

during the late summer and fall of 1999 that virtually stopped all branching of A. cervicornis at

all site in the Bahamas and Florida, as well as reducing branching in A. prolifera in the Bahamas

15

(Figure 2.3). This change could be due to the higher temperatures during 1999 (31°C) compared

to temperatures during summer 2000 (30°C).

Acropora prolifera (Fused Staghorn Coral) has been described as a hybrid (F1), having

traits similar to both A. palmata and A. cervicornis (Vollmer and Palumbi 2002). Its distribution

lies between A. cervicornis and A. palmata, generally between the high-energy reef crest habitats

of A. palmata and the somewhat deeper and less energetic fore-reef environments inhabited by A.

cervicornis. A. prolifera has a more “bushy” appearance than its counterpart A. cervicornis.

Higher branching rates of A. prolifera, compared to A. cervicornis as shown in Figure 2.3, result

in thin blade-like branches, faintly similar to those of A. palmata. There appears to be a trade-off

between branching and colony growth (measured by either increases in weight or length).

Unlike A. cervicornis, where growth and branching are directly correlated, branching in A.

prolifera occurs predominantly during periods of low growth (Fig 2.17a-f).

16

ACKNOWLEDGEMENTS

The work was funded by the National Undersea Research Program (UNCW in Key

Largo, Florida and the Caribbean Marine Research Center on Lee Stocking Island in the

Bahamas) and the National Science Foundation (9203327,9702032,9906976). We would like to

thank Dr. Steve Miller, Otto Rutten, Mike Birns and others, for continued logistical support from

the NURC/UNCW Day-boat Program in Key Largo. Drs. John Marr, Tom Bailey, Steve Jury, as

well as Brian Kakuk, Jeremy, Ester, both Kerleens, and Craig and Tara Dahlgren of CMRC Lee

Stocking Island Bahamas. Thanks to Dan Thornhill, Mark Warner, Todd LaJuenesse, Tom

Shannon, Nathan Jess, and Peter Anziano for field assistance. Permitting in the United States

thanks to Harold Hudson, John Hallace and others from NOAA Florida Keys.

17

REFERENCES

Bell, A. (1991). Plant Form. New York, Oxford University Press. Blackstone, N. W. (2003). “Redox signaling in the growth and development of colonial

hydroids.” Journal of Experimental Biology 206(4): 651-658. Bottjer, D. J. (1980). “Branching Morphology of the Reef Coral Acropora-Cervicornis in

Different Hydraulic Regimes.” Journal of Paleontology 54(5): 1102-1107.

Rinkevich, B. (2002). “The branching coral Stylophora pistillata: Contribution of genetics in shaping colony landscape.” Israel Journal of Zoology 48(1): 71-82.

Vollmer, S. V. and S. R. Palumbi (2002). “Hybridization and the evolution of reef coral

diversity.” Science 296(5575): 2023-2025.

18

Figure 2.1aAll Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reef), A cervicornisAverage New Branches Added Since Previous Sampling Period

0

0.5

1

1.5

2

2.5

3

3.5

Feb-99

Apr-99

Jun-99

Aug-99

Oct-99

Dec-99

Feb-00

Apr-00

Jun-00

Aug-00

Oct-00

Dec-00

Feb-01

Apr-01

Jun-01

Aug-01

Oct-01

Dec-01

New

Bra

nche

s

ADM AC 8/99

ADM AC 11/99

ADM AC 3/00

LG AC 6/99

LG AC 11/99

LG AC 5/00

LSI AC 1/99

LSI 5/99 AC

LSI 8/99 AC

Figure 2.1bAll Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reef), A. cervicornisTotal Branches Since Intial Transplantation

0

2

4

6

8

10

12

Jan-99

Mar-99

May-99

Jul-99

Sep-99

Nov-99

Jan-00

Mar-00

May-00

Jul-00

Sep-00

Nov-00

Jan-01

Mar-01

May-01

Jul-01

Sep-01

Nov-01

Tota

l Bra

nche

s `

LSI AC 1/99

LSI AC 5/99

LSI AC 8/99

LSI AC 11/99

ADM AC 8/99

ADM AC 11/99

ADM AC 3/00

ADM AC 5/00

ADM AC 8/00

LG AC 6/99

LG AC 11/99

LG AC 5/00

LG AC 8/00

LG AC 11/00

19

Figure 2.2All Sites Bahamas (LSI) and Florida (Admiral and Little Grecian Reefs)A. cervicornis, A. proliferaAverage New Branches per Day Since Previous Sampling Period

0

0.02

0.04

0.06

0.08

0.1

Feb-99

May-99

Jul-99

Oct-99

Dec-99

Mar-00

Jun-00

Aug-00

Nov-00

Jan-01

Apr-01

Jun-01

Sep-01

Nov-01

Feb-02

New

Bra

nche

s/da

y

ADM 8/99

ADM 11/99

ADM 3/00

LG 6/99

LG 11/99

LG 5/00

LSI 1/99

LSI 5/99

LSI 8/99

LSI prolifera

Figure 2.3Total Branches Since Initial TransplantationBahamas (LSI) (A. cervicornis, A. prolifera )Little Grecian Reef (LG) and Admiral Reef (ADM), Florida (A. cervicornis )

0

5

10

15

20

25

30

35

Dec-98

Mar-99

Jun-99

Sep-99

Dec-99

Mar-00

Jun-00

Sep-00

Dec-00

Mar-01

Jun-01

Sep-01

Dec-01

Tot

al B

ranc

hes

LSI A. prolifera1/99

ADM AC 8/99

LG AC 6/99

LSI AC 8/99

20

Figure 2.4(A-I)ADM AUG 1999 A. cervicornis Original transplant size (length) v. average new branches since previous sampling period

0.0

0.5

1.0

0 10 20 30 40 50 60

Original Size ofTransplant

New

Bra

nche

s .

11/99 Branches

A

y = 0.0295x - 1.1233R2 = 0.2149

0.0

0.5

1.0

1.5

2.0

2.5

0 10 20 30 40 50 60


New

Bra

nche

s .

3/00 Branches

B

y = 0.0037x + 0.2267R2 = 0.0035

0

0.5

1

1.5

2

2.5

0 10 20 30 40 50 60Original Size ofTransplant

New

Bra

nche

s . 5/00 Branches

C

y = -0.0234x + 1.8313R2 = 0.0439

0

1

2

3

4

5

6

0 20 40 60


New

Bra

nche

s .

9/00 Branches

E

y = 0.0481x - 1.2352R2 = 0.0642

0

1

2

3

4

5

6

0 10 20 30 40 50 60


New

Bra

nche

s

.

11/00 Branches

F

y = 0.0316x + 0.0486R2 = 0.0196

0

1

2

3

4

5

6

7

0 10 20 30 40 50 60


New

Bra

nche

s .

3/01 Branches

G

y = -0.0887x + 6.1414R2 = 0.2058

0

1

2

3

4

5

6

7

0 10 20 30 40 50 60


New

Bra

nche

s .

5/01 Branches

H

y = 0.0064x + 0.535R2 = 0.0028

0

1

2

3

4

5

6

7

0 10 20 30 40 50 60


New

Bra

nche

s .

8/01 Branches

I

y = 0.0211x + 1.2411R2 = 0.0139

0.0

1.0

2.0

3.0

4.0

5.0

6.0

0 10 20 30 40 50 60


New

Bra

nche

s .

8/00 Branches

D

21

Fig 2.5(A-I) ADM Aug 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

0

0.5

1

1.5

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s 11/99 Branches

B

y = 0.0295x - 0.1233R2 = 0.2149

0

0.5

1

1.5

2

2.5

3

3.5

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s

3/00 Branches

C

y = 0.0739x - 1.5583R2 = 0.4616

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5


Num

ber o

f Bra

nche

s

5/00 Branches

E

y = 0.0833x + 1.1333R2 = 0.112

00.5

11.5

22.5

33.5

44.5

55.5

66.5

77.5

88.5


Num

ber o

f Bra

nche

s

9/00 Branches

F

y = 0.0644x + 2.0416R2 = 0.0739

00.5

11.5

22.5

33.5

44.5

55.5

66.5

77.5

88.5

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s

11/00 Branches

G

y = 0.0773x + 2.7377R2 = 0.0353

0123456789

101112131415


Num

ber o

f Bra

nche

s

3/01 Branches

H

y = -0.1605x + 14.648R2 = 0.1247

0123456789

101112131415

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s

5/01 Branches

I

y = 0.0252x + 7.5136R2 = 0.0056

0123456789

1011121314

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s

8/01 Branches

D

y = 0.0956x - 0.3492R2 = 0.1881

00.5

11.5

22.5

33.5

44.5

55.5

66.5

77.5

0 10 20 30 40 50 60


Num

ber o

f Bra

nche

s . 8/00 Branches

22

Fig. 2.6(A-H)ADM Nov 1999 A. cervicornisOriginal transplant size (length) v. new branches produced since previous sampling period

A

y = 0.0467x - 0.0838R2 = 0.4672

0

1

2

3

4

5

6

0 20 40 60 80 100 120


New

Bra

nche

s

3/00 Branches

B

y = -0.0106x + 1.2836R2 = 0.1111

0

1

2

3

4

0 50 100 150


New

Bra

nche

s

5/00 Branches

C

y = 0.0004x + 0.5197R2 = 0.0002

0

1

2

3

4


New

Bra

nche

s

7/00 Branches

D

y = -0.0003x + 0.2445R2 = 0.0002

0

1

2

3

0 50 100 150


New

Bra

nche

s

9/00 Branches

E

y = -0.021x + 2.1915

R2 = 0.2349

0

1

2

3

4

0 20 40 60 80 100 120


New

Bra

nche

s

11/00 Branches

F

y = 0.0039x - 0.1238R2 = 0.1901

0

1

2

0 20 40 60 80 100 120


New

Bra

nche

s

3/01 Branches

G

y = 0.0476x + 0.2516R2 = 0.2052

0

1

2

3

4

5

6

7

8

9

10

11

0 20 40 60 80 100 120


New

Bra

nche

s

5/01 Branches

H

y = 0.03x + 0.8322R2 = 0.1974

0

1

2

3

4

5

6

7

8

0 20 40 60 80 100 120


New

Bra

nche

s

8/01 Branches

23

Fig. 2.7(A-H)ADM Nov 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

y = 0.0633x + 0.4582R2 = 0.6615

012345678

0 50 100 150


Num

ber o

f Bra

nche

s

3/00 Branches

B

y = 0.052x + 1.5777R2 = 0.7011

0123456789

0 50 100 150


Num

ber o

f Bra

nche

s

5/00 Branches

C

y = 0.0531x + 2.1424R2 = 0.5571

0123456789

10

0 50 100 150


Num

ber o

f Bra

nche

s

7/00 Branches

D

y = 0.0489x + 2.0363R2 = 0.6751

012345678

0 50 100 150

Original Size ofTransplantN

umbe

r of B

ranc

hes

9/00 Branches

E

y = 0.0329x + 3.7475R2 = 0.2465

0

1

2

3

4

5

6

7

8

9

0 20 40 60 80 100 120


Num

ber o

f Bra

nche

s

11/00 Branches

F

y = 0.033x + 3.2314R2 = 0.2546

012

34

56

789

10

0 20 40 60 80 100 120


Num

ber o

f Bra

nche

s

3/01 Branches

G

y = 0.0677x + 3.7492R2 = 0.5118

0123456789

10111213141516

0 20 40 60 80 100 120


Num

ber o

f Bra

nche

s

5/01 Branches

H

y = 0.103x + 4.1563R2 = 0.4802

02468

1012141618202224

0 20 40 60 80 100 120


Num

ber o

f Bra

nche

s

8/01 Branches

24

Fig 2.8(A-G) ADM March 2000 A. cervicornisOriginal transplant size (length) v. new branches produced since previous sampling period

A

y = 0.0187x + 0.6724R2 = 0.2605

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

New

Bra

nche

s

5/00 Branches

B

y = 0.0066x + 0.0711R2 = 0.1941

0

0.5

1

1.5

2

2.5

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

New

Bra

nche

s

7/00 Branches

C

y = 0.0012x + 0.505R2 = 0.003

00.5

11.5

22.5

33.5

0 50 100 150 200

Initial Length (mm)

New

Bra

nche

s

9/00 Branches

D

y = -0.0168x + 1.4499R2 = 0.2897

0

0.5

1

1.5

2

2.5

0 50 100 150 200

Initial Length (mm)

New

Bra

nche

s

11/00 Branches

E

y = -0.016x + 2.0146R2 = 0.0968

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 50 100 150 200

Initial Length (mm)

New

Bra

nche

s

3/01 Branches

F

y = 0.0481x - 0.0493R2 = 0.1263

0

2

4

6

8

10

12

14

0 50 100 150 200

Initial Length (mm)

New

Bra

nche

s

5/01 Branches

G

y = -0.0087x + 1.2942R2 = 0.046

0

0.5

1

1.5

2

2.5

3

3.5

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

New

Bra

nche

s

8/01 Branches

25

Fig 2.9(A-G) ADM March 2000 A. cervicornisOriginal transplant size (length) v. total branches

B

y = 0.0476x + 0.5012

R2 = 0.7719

0

1

2

3

4

5

6

7

8

9

10

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

Tota

l Bra

nche

s

7/00 Branches

A

y = 0.0434x + 0.5394R2 = 0.6837

0

1

2

3

4

5

6

7

8

9

10

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

Tota

l Bra

nche

s

5/00 Branches

C

y = 0.0455x + 0.8174R2 = 0.6356

0

2

4

6

8

10

12

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

Tota

l Bra

nche

s 9/00 Branches

D

y = 0.0055x + 2.9607R2 = 0.0065

0

1

2

3

4

5

6

7

8

0 50 100 150 200

Initial Length (mm)To

tal B

ranc

hes

11/00 Branches

E

y = -0.0259x + 5.597R2 = 0.0685

0

1

2

3

4

5

6

7

8

9

10

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

Tota

l Bra

nche

s

3/01 Branches

F

y = 0.0222x + 5.5477R2 = 0.0181

0

2

4

6

8

10

12

14

16

18

20

0 20 40 60 80 100 120 140 160 180 200

Initial Length (mm)

Tota

l Bra

nche

s

5/01 Branches

G

y = 0.0018x + 7.8061R2 = 0.0001

0

2

4

6

8

10

12

14

16

18

20

0 20 40 60 80 100 120 140 160 180 200Initial Length (mm)

Tota

l Bra

nche

s

8/01 Branches

26

Fig 2.10(A-I) LG June 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

y = 0.0117x + 1.1201R2 = 0.1813

0

1

2

3

4

5

0 20 40 60 80 100 120 140 160 180

Length of Transplant

Tota

l Bra

nche

s

8/99 Branches

B

y = 0.0094x + 1.044R2 = 0.1283

0

1

2

3

4

5

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

11/99 Branches

C

y = 0.0107x + 1.367R2 = 0.1243

0

1

2

3

4

5

6

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

5/00 Branches

D

y = 0.0649x - 0.2204R2 = 0.5389

0

2

4

6

8

10

12

14

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

8/00 Branches

E

y = 0.0643x - 0.2893R2 = 0.5545

0

2

4

6

8

10

12

14

0 20 40 60 80 100 120 140 160 180Length of Transplant

Tota

l Bra

nche

s

9/00 Branches

F

y = 0.0632x - 0.1645R2 = 0.5316

0

2

4

6

8

10

12

14

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

11/00 Branches

G

y = 0.0686x + 0.1703

R2 = 0.7733

0

2

4

6

8

10

12

14

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

3/01 Branches

H

y = 0.0579x + 2.0222R2 = 0.5759

0

2

4

6

8

10

12

14

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

5/01 Branches

I

y = 0.0648x + 3.5626R2 = 0.4579

0

2

4

6

8

10

12

14

16

0 20 40 60 80 100 120 140 160 180


Tota

l Bra

nche

s

8/01 Branches

27

Figure 2.11(A-I)LG June 1999 A. cervicornis Original transplant size (length) v. average new branches since previous sampling period

A

y = 0.0008x + 0.5319R2 = 0.0009

0

1

2

3

4

0 20 40 60 80 100 120 140 160 180

Original Transplant length

New

Bra

nche

s

8/99 Branches

B

0

1

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

11/99 Branches

C

y = -0.0009x + 0.6527R2 = 0.001

0

1

2

3

4

5

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

5/00 Branches

D

y = 0.0541x - 1.5874R2 = 0.5835

0

1

2

3

4

5

6

7

8

9

10

11

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

8/00 Branches

E

0

1

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

9/00 Branches

F

0

1

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

11/00 Branches

G

y = 0.0053x + 0.3348R2 = 0.0458

0

1

2

3

4

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

3/01 Branches

H

y = -0.0041x + 1.5222R2 = 0.0128

0

1

2

3

4

5

6

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

5/01 Branches

I

y = 0.0069x + 1.5404R2 = 0.0445

0

1

2

3

4

5

0 20 40 60 80 100 120 140 160 180


New

Bra

nche

s

8/01 Branches

28

Fig 2.12(A-G) LG Nov 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

y = 0.0441x + 0.1033R2 = 0.2413

0

1

2

3

4

5

6

7

8

0 20 40 60 80 100 120


Tota

l Bra

nche

s

5/00 Branches

B

y = 0.0765x - 0.2098R2 = 0.4236

0

1

2

3

4

5

6

7

8

9

10

0 20 40 60 80 100 120


Tota

l Bra

nche

s

8/00 Branches

C

y = 0.0683x + 0.0088R2 = 0.4387

0

1

2

3

4

5

6

7

8

0 20 40 60 80 100 120


Tota

l Bra

nche

s 9/00 Branches

D

y = 0.0917x - 0.6562R2 = 0.4513

0

1

2

3

4

5

6

7

8

9

10

11

0 20 40 60 80 100 120


Tota

l Bra

nche

s

11/00 Branches

E

y = 0.0655x + 0.9796R2 = 0.3286

0

1

2

3

4

5

6

7

8

9

10

0 20 40 60 80 100 120


Tota

l Bra

nche

s

3/01 Branches

F

y = 0.1031x + 0.7499R2 = 0.6084

0

1

2

3

4

5

6

7

8

9

10

11

12

13

0 20 40 60 80 100 120


Tota

l Bra

nche

s

5/01 Branches

G

y = 0.0906x + 1.1855R2 = 0.4287

0

1

2

3

4

5

6

7

8

9

10

11

12

0 20 40 60 80 100 120


Tota

l Bra

nche

s

8/01 Branches

29

Figure 2.13(A-G)LG Nov 1999 A. cervicornis Original transplant size (length) v. average new branches since previous sampling period

A

y = 0.0316x - 0.5104R2 = 0.1419

0

1

2

3

4

5

6

7

0 20 40 60 80 100 120


New

Bra

nche

s

5/00 Branches

B

y = 0.0327x - 0.3088R2 = 0.2526

0

1

2

3

4

5

0 20 40 60 80 100 120


New

Bra

nche

s

8/00 Branches

C

y = 0R2 = #N/A

0

1

0 20 40 60 80 100 120


New

Bra

nche

s

9/00 Branches

D

y = 0.0158x - 0.1624R2 = 0.1134

0

1

2

3

4

0 20 40 60 80 100 120


New

Bra

nche

s

11/00 Branches

E

y = -0.0202x + 1.5178R2 = 0.2297

0

1

2

3

4

0 20 40 60 80 100 120Length of Transplant

New

Bra

nche

s 3/01 Branches

F

y = 0.0613x - 1.3069R2 = 0.5341

0

1

2

3

4

5

6

7

0 20 40 60 80 100 120


New

Bra

nche

s

5/01 Branches

G

y = -0.0056x + 0.599R2 = 0.0374

0

1

2

3

0 20 40 60 80 100 120


New

Bra

nche

s

8/01 Branches

30

Fig 2.14 (A-H) LSI Jan 1999 A. cervicornisOriginal transplant size (length) v. average new branches since previous sampling period

A

y = 0.028x + 0.9851R2 = 0.0662

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s 5/99 branches

B

y = 0.0444x + 0.9616R2 = 0.1907

0

0.5

1

1.5

2

2.5

3

3.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s 8/99 branches

C

y = 0R2 = #N/A

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

0 5 10 15 20 25 30 35

Initial Weight

Tota

l Bra

nche

s 11/99 branches

D

y = 0.0888x - 0.6349R2 = 0.5471

0

0.5

1

1.5

2

2.5

3

3.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s 3/00 branches

E

y = -0.0771x + 2.5517R2 = 0.1461

0

0.5

1

1.5

2

2.5

3

3.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s

5/00branches

F

0

0.5

1

1.5

2

2.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s

8/00 branches

G

y = -0.1586x + 3.6641R2 = 0.0788

0

1

2

3

4

5

6

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s

11/00 branches

H

y = -0.0442x + 1.3526R2 = 0.0154

0

0.5

1

1.5

2

2.5

3

3.5

0 5 10 15 20 25 30 35

Initial Weight

New

Bra

nche

s

3/01 branches

31

Fig 2.15 (A-H) LSI Jan 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

y = 0.0045x + 0.9055R2 = 0.0853

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s 5/99 branches

B

y = 0.008x + 0.755R2 = 0.2967

0

0.5

1

1.5

2

2.5

3

3.5

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s 8/99 branches

S

y = 0.0071x + 0.7774R2 = 0.242

0

0.5

1

1.5

2

2.5

3

3.5

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s

11/99 branches

D

R2 = 0.6316

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s 3/00 branches

E

y = 0.0118x + 2.202R2 = 0.0907

0

1

2

3

4

5

6

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s

5/00 branches

F

y = 0.0127x + 2.1341R2 = 0.0783

0

1

2

3

4

5

6

7

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s

8/00 branches

G

y = 0.0081x + 3.8067R2 = 0.0627

0

1

2

3

4

5

6

7

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s

11/00 branches

H

y = -0.0048x + 6.0549R2 = 0.0035

0

1

2

3

4

5

6

7

8

9

10

0 50 100 150 200 250

Initial Length

Tota

l Bra

nche

s

3/01 branches

32

33

Fig 2.16 (A-H) LSI May 1999 A. cervicornisOriginal transplant size (length) v. average new branches since previous sampling period

A

y = 0.0099x - 0.5429R2 = 0.3377

0

0

0

1

1

1

1

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

8/99 branches

B

y = 0.0055x - 0.3124R2 = 0.185

0

0

0

1

1

1

1

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

11/99 branches

C

y = 0.0184x - 0.8792R2 = 0.4138

0

1

1

2

2

3

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

3/00 branches

D

y = -0.003x + 1.1212R2 = 0.0023

0

1

2

3

4

5

6

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

5/00 branches

E

y = -0.0155x + 1.7577R2 = 0.2688

0

1

1

2

2

3

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

8/00 branches

G

y = -0.0016x + 1.4655R2 = 0.0002

0

1

1

2

2

3

3

4

4

5

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

3/01 branches

H

y = 0.3333x - 27.667R2 = 1

0

1

2

3

4

5

6

7

8

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

5/01 branches

F

y = -0.004x + 0.5485R2 = 0.021

0

0

0

1

1

1

1

0 20 40 60 80 100 120

Initial Length

New

Bra

nche

s

11/00 branches

Fig 2.17 (A-H) LSI May 1999 A. cervicornisOriginal transplant size (length) v. total branches

A

y = 0.0099x + 0.4571R2 = 0.3377

0

0.5

1

1.5

2

2.5

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

8/99 branches

B

y = 0.0118x + 0.2308R2 = 0.6699

0

0.5

1

1.5

2

2.5

0 20 40 60 80 100 120 140 160

Initial Length

Tota

l Bra

nche

s

11/99 branches

C

R2 = 0.824

0

0.5

1

1.5

2

2.5

3

3.5

0 20 40 60 80 100 120 140 160 180 200

Initial Length

Tota

l Bra

nche

s

3/00 branches

D

y = 0.0309x + 0.3867R2 = 0.2606

0

1

2

3

4

5

6

7

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

5/00 branches

E

y = 0.0153x + 1.8618R2 = 0.0539

0

1

2

3

4

5

6

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

8/00 branches

F

y = 0.0325x + 0.9619R2 = 0.1637

0

1

2

3

4

5

6

7

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

11/00 branches

G

y = -0.002x + 4.8319R2 = 0.0002

0

1

2

3

4

5

6

7

8

9

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

3/01 branches

H

y = 0.1905x - 9.8095R2 = 1

0

2

4

6

8

10

12

0 20 40 60 80 100 120

Initial Length

Tota

l Bra

nche

s

5/01 branches

34

Fig 2.18(a-i)LSI Feb 99 A. proliferaOriginal transplant size (length) v. total branches

A

y = 0.2199x - 8.1237R2 = 0.6354

0

2

4

6

8

10

0 20 40 60 80

Original Transplant Size

Tota

l Bra

nche

s5/99Branches

B

y = -0.0211x + 9.1657R2 = 0.0176

0

2

4

6

8

10

12

0 20 40 60 80


Tota

l Bra

nche

s

8/99Branches

C

y = 0.0264x + 8.0428R2 = 0.0066

02468

10121416

0 20 40 60 80Original Transplant Size

Tota

l Bra

nche

s

11/99Branches

D

y = 0.1811x + 2.3933R2 = 0.6494

0

5

10

15

20

0 20 40 60 80


Tota

l Bra

nche

s 3/00Branches

E

y = 0.2284x + 1.41R2 = 0.4743

0

5

10

15

20

0 20 40 60 80


Tota

l Bra

nche

s

5/00Branches

F

y = 0.4422x - 6.5036R2 = 0.3573

05

1015202530

0 20 40 60 80


Tota

l Bra

nche

s

8/00Branches

G

y = 0.3653x + 2.1118R2 = 0.2052

05

10152025303540

0 20 40 60 80


Tota

l Bra

nche

s

11/00Branches

H

y = 0.5017x - 1.9072R2 = 0.1623

05

1015202530354045

0 20 40 60 80


Tota

l Bra

nche

s

3/01 Branches

I

y = 1.2545x - 40.205R2 = 0.7537

0

10

20

30

40

50

0 20 40 60 80


Tota

l Bra

nche

s

5/01Branches

35

Fig. 2.19a-iLSI Feb 99 A. proliferaOriginal Tranplants size (length) v. average new branches since previous sampling time

A

y = 0.2199x - 9.1237R2 = 0.6354

012345678

0 20 40 60 80


New

Bra

nche

s 5/99 Branches

B

y = -0.0085x + 4.4663R2 = 0.0019

0

1

2

3

4

5

6

0 20 40 60 80


New

Bra

nche

s

8/99Branches

C

y = 0.0045x + 1.6273R2 = 0.0002

01234567

0 20 40 60 80


New

Bra

nche

s

11/99Branches

D

y = 0.0296x + 1.368R2 = 0.0104

012345678

0 20 40 60 80

Original Transplant WeightN

ew B

ranc

hes

3/00Branches

E

y = 0.0727x - 2.3822R2 = 0.0391

0

2

4

6

8

0 20 40 60 80

Original Transplant Weight

New

Bra

nche

s

5/00Branches

F

y = 0.1657x - 4.5115R2 = 0.0756

0

5

10

15

0 20 40 60 80


New

Bra

nche

s

8/00Branches

G

y = -0.0904x + 9.7335R2 = 0.065

0

2

4

6

8

10

0 20 40 60 80


New

Bra

nche

s

11/00Branches

H

y = 0.1094x - 1.7827R2 = 0.0418

0

2

4

6

8

10

12

0 20 40 60 80


New

Bra

nche

s

3/01Branches

I

y = 0.0091x + 2.5182R2 = 0.0004

01234567

0 20 40 60 80


New

Bra

nche

s

5/01Branches

36

Fig. 2.20aADM Aug99 A. cervicornisNew branches/day v. percent growth length/day, percent growth weight/day (95%CI)

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Nov-99

Dec-99

Jan-00Feb-00M

ar-00A

pr-00M

ay-00

Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01

Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

bra

nche

s/da

y `

0

0.002

0.004

0.006

0.008

0.01

0.012

0.014

0.016

0.018

0.02

perc

ent g

row

th/d

ay

ADM 8/99branches/day

8/99 percentlength/day

8/99 percentweight/day

Fig. 2.20bADM Nov99 A. cervicornisNew branches/day v. percent growth length/day, percent growth weight/day (95%CI)

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Nov-99

Dec-99

Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay

ADM 11/99newbranches/day



37

Fig. 2.20c ADM March00 A. cervicornisNew branches/day v. percent growth length/day, percent growth weight/day (95%CI)

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Nov-99

Dec-99

Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

Bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay

ADM 3/00 NewBranches/day



Fig. 2.21aLG Jun99 A. cervicornisNew branches/day v.percent growth length/day and percent growth weight /day

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Aug-99

Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

Bra

nche

s/da

y

0.000

0.004

0.008

0.012

0.016

0.020

perc

ent g

row

th/d

ay

LG 6/99newbranches/day

6/99percentlength/day

6/99percentweight/day

38

Fig. 2.21bLG Nov99 A. cervicornisNew branches/day v.percent growth length/day and percent growth weight /day

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Aug-99

Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

Bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay

LG 11/99 newbranches/day



Fig. 2.21cLG may00 A. cervicornisNew branches/day v.percent growth length/day and percent growth weight /day

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Aug-99

Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01

New

Bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay




39

Fig. 2.22aLSI, Bahamas Jan99 A. cervicornisNew Branches/day v percent growth length, percent growth weight

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Jan-99Feb-99M

ar-99A

pr-99M

ay-99Jun-99Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01

new

bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay

LSI 1/99 newbranches/day



Fig. 2.22bLSI, Bahamas May99 A. cervicornisNew Branches/day v percent growth length, percent growth weight

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Jan-99Feb-99M

ar-99A

pr-99M

ay-99Jun-99Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01

new

bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay LSI 5/99 new

branches/day

5/99 %length/day

5/99 % wt/day

40

Fig. 2.22cLSI, Bahamas Aug99 A. cervicornisNew Branches/day v percent growth length, percent growth weight

0

0.01

0.02

0.03

0.04

0.05

0.06

0.07

0.08

Jan-99Feb-99M

ar-99A

pr-99M

ay-99Jun-99Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01

new

bra

nche

s/da

y

0

0.004

0.008

0.012

0.016

0.02

perc

ent g

row

th/d

ay LSI 8/99 new

branches/day

8/99 %length/day

8/99 %wt/day

Fig. 23All Sites (A. cervicornis, A. prolifera )Total grams/total mm

0

0.05

0.1

0.15

0.2

0.25

0.3

Jan-99

Mar-99

May-99

Jul-99

Sep-99

Nov-99

Jan-00

Mar-00

May-00

Jul-00

Sep-00

Nov-00

Jan-01

Mar-01

May-01

Jul-01

Sep-01

Nov-01

grow

th w

eigt

h/ g

row

th le

ngth

(gra

ms/

mm

)

ADM 8/99

ADM 11/99

ADM 3/00

ADM 5/00

ADM 8/00

LG 6/99

LG 11/99

LG 5/00

LG 8/00

LG 11/00

LSI 1/99

LSI 5/99

LSI 8/99

LSI 11/99

LSI 3/00

Prolifera

41

Fig. 2.24Little Grecian ReefJune 1999 Tranplants, New Branches/day, Total grams/mm

0

0.05

0.1

0.15

0.2

0.25

May-99

Jul-99

Sep-99

Nov-99

Jan-00

Mar-00

May-00

Jul-00

Sep-00

Nov-00

Jan-01

Mar-01

May-01

Jul-01

Sep-01

Nov-01

new

bra

nche

s/da

y, to

tal g

ram

s/m

m


LG 6/99 Totalgrams/mm

Fig. 2.25Little Grecian Reef, A. cervicornis Total Grams/Total Length (g/mm)

0.00

0.05

0.10

0.15

0.20

0.25

0.30

Aug-99

Oct-99

Dec-99

Feb-00

Apr-00

Jun-00

Aug-00

Oct-00

Dec-00

Feb-01

Apr-01

Jun-01

Aug-01

g/m

m

`

LG 6/99 Back Reef(2m)

LG 11/99 Reef Crest(2m)

LG 5/00 Fore ReefChannel (3-4m)



42

CHAPTER 3

EFFECT OF SEASON AND CORAL SIZE ON THE GROWTH RATE OF TRANSPLANTED

ACROPORA CERVICORNIS1


43

ABSTRACT

Growth of the Staghorn coral Acropora cervicornis was monitored in situ at three sites, two in

the Florida Keys and one in the Bahamas for two years quarterly. Highest growth rates, in terms

of linear extension and buoyant grams added since last sampling period, were observed in late

spring and early summer (March – August), lowest in late summer through winter (September-

March). Ratios of accretion (g):linear growth (mm) were relatively constant within a particular

sight, with corals from higher wave energy sites in the Bahamas and fore-reef sites in the Florida

Keys having higher ratios than corals from low energy inshore patch reefs. Growth rates were

generally positively correlated (p=.05) with colony size, with extension positively correlated

(p=.05)about 40% of the time and accretion about 90% of the time. Time of transplantation

appears to be important for subsequent growth. Corals transplanted during the late-summer

early-fall time period, when seawater temperatures were highest, or winter (November- March

time period), when seawater temperatures were lowest, showed low initial linear extension and

accretion which often continued throughout the study. In contrast, those corals transplanted

during early spring exhibited higher growth rates than those transplanted during warmer seasons.

Highest growth rates were observed on the low wave-energy inshore patch reef (Admiral reef) in

the Florida Keys compared to the higher energy fore-reef site (Little Grecian Reef) in the Florida

Keys and a high-current patch reef in the Bahamas.

Key Words: Acropora, branching, staghorn coral, restoration, buoyant weight

44

INTRODUCTION

Growth of branching corals can be characterized by increases in (a) weight and (b) length

along two growth axes (axial and radial). The major environmental factors that influence growth

of corals are temperature and the amount o f light or sun hours that are available (Yap, 1984).

Other factors that can influence growth include predation (Baums 2003), eutrification and

sedimentation (Roy and Smith 1971, Dodge et al 1974, Rogers 1979, Hoegh-Goldberg 1997)

algal over-growth, breakage (Tunnicliffe 1981), and food abundance (Lewis 1974, Wellington

1982); although these factors seem to influence the coral to a lesser extent than light and

temperature.

Many studies have looked for a relationship between extension and accretion. Density

bands that form in boulder corals are evidence that there is some functionality between the two at

least seasonally (Highsmith 1979, Wellington and Glynn 1983). Highsmith (1979), Schneider

and Smith (1982), and Wellington and Glynn (1983) all showed that calcification and extension

are out of phase at least in some corals. Fitt et al (2000) showed seasonal changes in tissue

biomass and Zooxanthellae concentrations that could result in different amounts of calcification

and thus different banding in A. cervicornis and A. palmata. All corals in genus Acropora do not

band. This is due to secondary infilling of the skeleton which does not happen in many of the

boulder species (Gladfelter 1984). Because of this infilling any sclerochronology is all but

impossible with the acroporids. Although they do not band, extension and calcification can be

measured easily with the buoyant weight technique and linear extension measurements. It can

also be deduced by x-radiography which has been used extensively in the past twenty years.

45

Some experiments have been designed to study the roles of light and temperature on the

growth of corals. Shinn (1964) found highest growth rates when the temperature is at its

warmest between 28°C and 30°C and death at 13.3°C Lough and Barnes (2000) found that

extension and calcification were directly related to the average sea surface temperatures (SST).

For a 1° C rise in SST, calcification and extension both increased in both high and low latitude

corals. They also found that extension and calcification rates were greater on top than the sides

of the colonies indicating light may be enhancing calcification. Although higher temperatures

mean higher growth, corals do have thermal limits. A worldwide rise in the SSTs has had

detrimental effects on coral reefs. A few degrees above the average annual high mixed with

doldrums conditions can “bleach” corals. This bleaching is the loss of symbiotic algae of

upwards 90% and leaves the coral without photosynthetic products and photo protection.

Globally, temperature induced bleaching is blamed for a decrease of the worlds coral reefs by

1/10 in the past fifteen years (Wilkerson, 1998).

Light has a large role in the growth of corals. Kawagutti (1948) was one of the first to

show that light enhances deposition of calcium carbonate in hermatypic corals. Since then it

seems to have been studied more than any other of the environmental factors yet is still one of

the most poorly understood. Different species have significantly different responses to light, and

most usually have an excess of light leading to photo-inhibition in the endosymbiotic algae.

Chalker (1977) found that A.cervicornis shows a daily rhythm in calcification with the maxima

at sunrise and sunset and no change in the dark rate. This suggests a daily rhythm that is directly

proportional to the daily rhythm in calcification capacity of the endosymbionts. In fact, he saw

these rhythms persist for at least on day in the total darkness. Recent DNA analysis of the

endosymbionts has led to the discovery of many different types of zooxanthellae, referred to as

46

clades, and each of these seem to respond in a different way to different light and temperature

regimes (i.e. photoinhibition or algal death at increased light and temp).

Linear extension rate has been found to be independent of colony size in Pocillopora

damicornis (Kinizie and Sarmiento 1986) from Western Austrailia, Samoa, the Great Barrier

Reef and Hawaii. This data suggest determinate growth in colonial coral forms.

Elevated nutrients have been found to increase calcification in P. damicornis and

Stylophora pistillata in a long term nutrient enrichment experiments on the Great Barrier Reef

(Hoeugh-Guldberg, 1997). In this experiment both nitrogen and phosphorus were elevated in

mircroatolls for two years. Even though most was taken up by organisms other than corals, the

small amount that reached the coral colonies had a statistical impact thus suggesting that even

very small amount of nutrients can elevate coral calcification.

47

METHODS

Transplantation experiments were conducted on a patch reef due north of Lee Stocking

Island, Bahamas (N 25° 46.990’ W 076° 07.852’) and at two sites off Key Largo, Florida,

Admiral Reef (N 25° 02.732’ W 080° 23.675’) a 3 m deep inshore patch reef and Little Grecian

Reef (N 25°07.164’ W 080° 18.107’) a 3m deep offshore reef crest.

Fifteen A. cervicornis and A. prolifera fragments, ranging in size from 2-19 cm in length,

were attached to cement bricks adjacent to the reef they were collected. Transplants were glued

into a 2″ section of ½ ″ PVC with Oately® epoxy putty then attached to submerged concrete

blocks. The weight of the epoxy and PVC pipe were measured so that the initial weight of the

transplant could be determined.

Fragments were weighed using the Bouyant Weight method to the nearest milligram.

Temperature (25oC) and salinity (36 ppt) were held constant during measurements at every

season. Length was measured using vernier calipers to the nearest millimeter. Branching was

analyzed by either calculating the total number of branches (total) arising since the original

transplant date, or the number of branches added (new) since the previous sampling date.

Initially almost all fragments (ca. 75 per site) had a single axial corallite (no radiating branches

from a single stick) and new branches produced from this single stick were tallied over time.

The only exceptions were one transplant from LG 6/99 which started with three branches (one

main stick with two radiating branches), three transplants from LG 11/99 which started with two

branches, two transplants from ADM 3/00 which started with two branches and two that started

with three branches. The dead parts where not counted in the linear measurements although

since they cannot be removed they were measured with the buoyant weight technique.

48

Data was analyzed using one-way ANOVA’s with 95% confidence intervals. All linear

regressions correlated at the 5% level of significance using all live replicates for that season

(D.F. = ~15). Data was plotted either as percent of the original size per day or as the total

growth per day since the previous period. Total weight per day is stated as “total weight” on

graphs and percent of the original size is plotted as “percent of original size.”

49

RESULTS

Highest growth rates at each site occurred in the late-spring, early-summer period (May-

August) in most cases, with the lowest in late winter (Fig 3.1-3.4). Transplants monitored in the

Bahamas (Fig 3.4a-d). had lower seasonal differences in growth rate compared to the inshore and

offshore Florida reefs (Fig 3.1a-d, Fig. 3.3a-d), indicating a strong correlation between

temperature and growth rates.

Ratios of weight to length, indicating density of the branches, were generally consistent

within a particular habitat at a site, but varied with site (Fig. 3.5). Transplants at the least wave-

exposed site, the patch reef at Admiral Reef in Florida, had the lowest ratios while transplants at

the high-current fringing reef in the Bahamas had higher ratios. Transplants from the outer-reef

Little Grecian Reef in Florida exhibited three distinct patterns, depending on source of Acropora

cervicornis. The fragments for the first transplant in June 1999 came from the high-wave energy

reef-flat on the SW-portion of the reef and were consistently thicker (highest weight: length

ratio) than any other corals used during the entire two years of subsequent monitoring. The

second sets of corals were transplanted from the back-reef just adjacent to where all transplants

were kept and monitored. Their weight: length ratios were intermediate for all sets of transplants

from this reef, and very similar to corals from the Bahamas. All subsequent transplants from

Little Grecian were obtained from the relatively low-energy channel bordering the south side of

the reef; ratios from all of those transplants are as low as the ratios from transplants from

Admiral Reef.

Growth of transplants in relation to initial size over time generally showed a positive

slope, both in terms of increased total length (about 44% of the correlations were significant at

50

p=0.05) and increased buoyant weight (about 78% were significant at p=0.05 level)(Table 3.1,

3.2). The linear regressions of length verses colony size over time, about a third of these

correlations were significant (p<0.05) during any time interval at the Florida sites, while about

half were significant at any time interval at the Bahamas site. In fact, only one out of 100+

correlations of linear growth and size was significantly negative; meaning 99% of the growth

determinations over time were positively correlated with colony size (about 44% significant at

the p<0.05 level). For linear regressions of added weight vs. colony size over time, 100% of the

correlations were positive (about 78% data were significant at p<0.05).

51

DISCUSSION

Skeletal growth of corals can be characterized by increases in (a) weight and (b) length

along one or more growth axes. Environmental factors that influence growth of corals, in terms

of weight and length, include temperature (Shinn 1966, Highsmith 1979, Barnes and Crossland

1980, Crossland 1984, add more refs) and light (Buddemeier et al. 1974, Highsmith 1979,

Wellington 1982, Wellington and Glynn, 1983, Huston 1985, add more refs).

Much of the early work (e.g. Buddemeier et al. 1974, Highsmith 1979) on increases in

coral calcification has utilized X-radiographs of coral skeletons, calcium isotope depletion rates

from seawater, and/or alizarin-red staining of skeletons of live corals. X-radiographs have

revealed annual banding patterns in several massive coral species, such that the high-density

band is deposited during times of seasonally-cooler seawater in contrast to the low-density band

that is deposited when the coral is growing the fastest in the warmer seasons. Massive corals

typically grow at rates of about 10 mm per year, as seen in their seasonal density banding,

though decreases in the average rate of growth over time varies with coral species and

environmental stress while branching acroporids often grow at ten times that rate (e.g.

Shinn1966, Hudson 1981, Sneider and Smith 1982).

The growth of corals in the genus Acropora has been investigated on diurnal, seasonal

and decadal scales. Branching corals do not show annual bands of growth. However, both linear

extension and accretion of calcium carbonate vary on a diel cycle in the development of the axial

corallite of branching acroporid corals (Gladfelter 1984). In the Pacific staghorn coral Acropora

acuminata, linear extension is higher at night than during the day, though overall accretion is

greater during the day (Barnes and Crossland 1978. 1980). Gladfelter (1983) found the same

52

pattern in the Caribbean staghorn coral, A. cervicornis, and further documented two processes in

the construction of the skeleton of Acroporids: (1) linear extension in the form of deposition of a

framework of micritic fusiform crystals, possible of calcite, and occurring during the night, and

(2) subsequent nucleation and growth of aragonite needles on this framework, primarily during

the day, and essentially filling in the lattice work and contributing far more to accretion than the

initial linear extension. Le Tissier (1988) found similar patterns in P. damicornis where the

deposition of fusiform crystals on skeletal spines at the branch tip apex at night is proposed to

promote deposition of fasciculi during the day which results in the apical growth of the branch

tips.

Light is a major factor in controlling growth rates of tropical reef corals. The growth rate

of virtually all coral species decreases with depth over their normal depth range (e.g. Huston

1985), though there are a few species like Montipora vericosa in Hawaii that achieves maximum

photosynthesis and the highest population density at intermediate depths, it’s symbiotic

dinoflagellates apparently not able to efficiently handle high light intensity (Chang refs).

Generally, the number of hours of full sunlight is significantly correlated with coral growth rates

(Bak 1974). The increased growth seen in light is thought to involve intra- and intercellular

changes in pH and biochemistry associated with photosynthesis of the symbiotic dinoflagellates

(zooxanthellae) resulting in increased calcification (Barnes and Crossland 1974,Goroeau and

Hayes 197,Highsmith 1979). Chalker (1977) found daily rhythms in calcification rates of

Acropora cervicornis, with maxima occurring at sunrise and sunset, dips in the middle of the day

when photo inhibition was highest, and lowest rates as measures with Ca45 occurring at night;

all correlated with photosynthetic rates of the symbiotic dinoflagellates.

53

Optimal skeletal growth of acroporid corals appears to lie between 26° and 30°C; the

actual peak temperature range appears to be quite narrow with a 2-4°C range depending on

species (Shinn 1966, Gladfeleter 1984). Growth rates, in terms of linear extension and accretion,

decrease on both the cooler and warmer ends of the optimal temperature range. For instance,

Gladfelter (1984) shows two “peaks” of growth of Acropora cervicornis from St. Croix, Virgin

Islands; the first in mid-summer (July-August) and a second broad period of high growth in the

late fall (October-December) following a lower-growth period of warmest seawater in late

summer/early fall (August – October) and another decrease during the coldest seawater (March-

April) in this part of the Caribbean. Similarly, Yap and Gomez (1984) found decreased linear

extension of acroporids in the Philippines at temperatures greater than the seasonal average

maximum of 30oC, but no decreased growth at the normal minimum temperatures of about

26oC. Shinn (1966) documented decreased growth rates and significant mortality associated

with low winter-time temperatures (<20oC) of Acropora cervicornis living on the high-latitude

reefs of the Florida Keys, although these corals were transplanted to an area where they did not

normally exist.

Seasonal growth of corals follows the light and temperature minimums and maximums as

discussed above. Suresh and Mathew (1995) found that calcification rates of acroporids in the

Indian Ocean varied significantly with marked reductions during the monsoon season from June

through September; extension rates showed the same pattern but were more variable and thus not

significantly different over a seasonal scale. They attribute the seasonal reduction with reduced

light from increased clouds and sediments in the water. Barnes and Crossland (1980) study of

Acropora acuminata from the GBR is one of the few published studies that has documented

significant changes in linear extension of branching corals seasonally, in this case peaks were in

54

spring and early winter, lows were in the coldest part of winter (late June – October) as well as in

the hottest part of the summer. Wellington and Glynn (1983) concluded that light level is a

better correlate to skeletal density than temperature. Gladfelter (1984) surmises that in acroporid

corals linear extension appears to be influenced by temperatures above and below an optimal

range, but is not influenced by flucuations within that range.

It has long been known that branching corals living in high-energy habitats exhibit thick

break-resistant morphology, compared with long, thinner branches of the same species growing

in more protected environments. Harriott (1998) used differences in growth rates to explain

these different morphologies; while linear extension changed little between sites, those corals in

high-energy sites exhibited less linear extension (13 to 93mm) compared to cohorts living in

protected sites (31-115mm). Huston (1985) found that growth rates decrease with depth in some

species with highest a short distance from the surface and decreasing with depth.

The size of a coral colony appears to influence growth in some instances, depending on

the species of coral studied. For instance, Kinzie and Sarmiento (1986) found that linear

extension rate is independent of colony size for Pocillopra damicornis. Other studies have found

evidence of decreased growth rates with age of the coral (Alcala and Gomez 1982). Smith and

Hughes (1999) found low fecundity on newly fragmented corals compared to the control corals.

This would suggest a minimal size to reproduce and this could have an impact on the growth of

the coral colony. Only the data with A. prolifera show decreased growth during the second year

following transplantation (Fig. 3.need graph). Whether this slower growth reflects a “normal”

physiological change related to colony age, or are due to disease or other stresses impacting

acroporids in the Caribbean, is unknown.

55

REFERENCES

Alcala, A. C., E. D. Gomez, et al. (1982). “Survival and Growth of Coral Transplants in Central Philippines.” Kalikasan 11(1): 179-184.

Bak, R. P. M. (1974). “Available Light and Other Factors Influencing Growth of Stony

Corals Through the Year in Curacao.” Proceedings 2nd Int Coral Reef Sym 2: 229-233. Barnes, D. J. and C. J. Crossland (1980). “Diurnal and Seasonal Variations in the Growth

of a Staghorns Corals Measured by Time-Lapse Photography.” Limnology & Oceanography 25(6): 1113-1117.

Baums, I. B., M. W. Miller, et al. (2003). “Ecology of a Corallivorous Gastropod,

Coralliophila abbreviata, on Two Scleractinian Hosts. 1: Population Structure of Snails and Corals.” Marine Biology 142(6): 1083-1091.

Buddemeier, R. W., J. E. Maragos, et al. (1974). “Radiographic Studies of Coral Reef

Exoskeltons: Rates and Patterns of Coral Growth.” Journal of Experimental Marine Biology & Ecology 14: 179-200.

Caribbean Marine Research Center: 1996-2002 Normas Pond Temperature Data Lee

Stocking Island Bahamas. www://httpcmrc.org/lsi_seawater_temp.htm Chalker, B. E. (1977). “Daily Variation in the Calcification Capacity of Acropora

Cervicornis.” Proceedings 3rd Int Coral Reef Sym 2: 417-423. Crossland, C. J. (1984). “Seasonal Variations in the Rates of Calcification and

Productivity in the Coral Acropora Formosa on a High-Latitude Reef.” Marine Ecology-Progress Series 15(1-2): 135-140.

Fitt, W. K., F. K. McFarland, et al. (2000). “Seasonal Patterns of Tissue Biomass and

Densities of Symbiotic Dinoflagellates in Reef Corals and Relation to Coral Bleaching.” Limnology and Oceanography 45(3): 677-685.

Harriott, V. J. (1998). “Growth of the Staghorn Coral Acropora formosa at Houtman

Abrolhos, Western Australia.” Marine Biology (Berlin) 132(2): 319-325. Highsmith, R. C. (1976). “Coral Growth Rates and Environmental Control of Density

Banding.” Journal of Experimental Marine Biology & Ecology(37): 105-125. Hoeugh-Guldberg, O., M. Takabayashi, et al. (1997). “The Impact of Long-Term

Nutrient Enrichment On Coral Calcification And Growth.” Proceedings 8th Int Coral Reef Sym(1): 861-866.

Hudson, J. H. (1981). “Growth Rates in Montasterea annularis: a Record of Environmental Change in Key Largo Reef Marine Sanctuary, Florida.” Bull. Mar. 31: 444-459.

56

Huston, M. (1985). “Variation in Coral Growth Rates With Depth at Discovery Bay,

Jamaica.” Coral Reefs(4): 19-25. Gladfelter, E. H. (1984). “Skeletal Development in Acropora Cervicornis 3. A

Comparison of Monthly Rates of Linear Extension and Calcium Carbonate Accretion Measured Over a Year.” Coral Reefs 3(1): 51-57.

Kawagutti 1948 The Effect of Light On the Calcium Deposition of Corals. Bull

Oceanogr. Inst. Taiwan No.4:65-70. 1948 Kinzie, R. A. and T. Sarmiento (1986). “Linear Extension Rate is Independant of Colony

Size in the Coral Pocillopora damicornis.” Coral Reefs(4): 177-181. Le Tissier, M’D. A. A. (1988). “Diurnal Patterns of Skeleton Formation in Pocillopora

damicornis (Linnaeus).” Coral Reefs 7: 81-88. Loya, Y. (1976). “The Red Sea Coral Stylophora pistillata is an R-Stragetist.” Nature

259: 478-480. Lough, J. M. and D. J. Barnes (2000). “Environmental Controls on Growth of the

Massive Coral Porites.” Journal of Experimental Marine Biology & Ecology 245(2): 225-243.

NOAA (1/98-9/02) Historical Temperature Data Set - Molasses Reef Station http://www.coral.noaa.gov/seakeys/hist_data.shtml Shinn, E. A. (1966). “Coral Growth-Rate, An Environmental Indicator.” Journal of Paleontology (40): 233-240.

Smith, L. D. and T. P. Hughes (1999). “An Experimental Assessment of Survival, Re-

Attachment and Fecundity of Coral Fragments.” Journal of Experimental Marine Biology & Ecology 235(1): 147-164.

Suresh, V. R. and K. J. Mathew (1995). “Growth of Staghorn Coral Acropora aspera

(Dana) (Scleractinia: Acroporidae) in Relation to Environmental Factors at Kavaratti Atoll (Lakshadweep Islands), India.” Indian Journal of Marine Sciences 24(3): 175-176.

Tunnicliffe, V. (1981). “Breakage and Propagation of the Stony Coral Acropora

Cervicornis.” Proceedings of the National Academy of Sciences of the United States of America-Biological Sciences 78(4): 2427-2431.

Wellington, G. M. and P. W. Glynn (1983). “Environmental Influences on Skeltal

Banding in (Panama) Corals.” Coral Reefs 1: 215-222. Wellington, G. M. (1982). “An Experimental Analysis of Light and Zooplankton on

Coral Zonation.” Oecologia 52: 311-320.

57

Wilkerson, C. R. (1998). Status of Coral Reefs of the World. Townsville, Austrailian Institute of Marine Science.

Yap, H. T. and E. D. Gomez (1984). “Growth of Acropora Pulchra .2. Responses of

Natural and Transplanted Colonies to Temperature and Day Length.” Marine Biology 81(2): 209-215.

Kawagutti 1948 The Effect of Light On the Calcium Deposition of Corals. Bull

Oceanogr. Inst. Taiwan No.4:65-70. 1948 (with Sakumoto)

58

Fig. 3.1aAdmiral Reef, A. cervicornis Percent growth (length, mm/day) of orginal size for each sampling period

0

0.002

0.004

0.006

0.008

0.01

0.012

0.014

0.016

0.018

S-99

O-99

N-99

D-99

J-00

F-00

M-00

A-00

M-00

J-00

J-00

A-00

S-00

O-00

N-00

D-00

J-01

F-01

M-01

A-01

M-01

J-01

J-01

A-01

S-01

perc

ent g

row

th p

er d

ay (m

m/d

8/99 length/ day

11/99 length/ day

3/00 length/ day

5/00 length/ day

8/00 length/ day

Figure 3.1bAdmiral Reef , A. cervicornis Growth length (mm/day) for each sampling period

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

S-99

O-99

N-99

D-99

J-00

F-00

M-00

A-00

M-00

J-00

J-00

A-00

S-00

O-00

N-00

D-00

J-01

F-01

M-01

A-01

M-01

J-01

J-01

A-01

S-01

mm

/day

8/99 length/ day

11/99 length/ day

3/00 length/ day

5/00 length/ day

8/00 length/ day

59

Fig. 3.1cAdmiral Reef, A. cervicornis Percent growth (weight, g/d) of original size for each sampling period

0

0.005

0.01

0.015

0.02

0.025

0.03

S-99

O-99

N-99

D-99

J-00

F-00

M-00

A-00

M-00

J-00

J-00

A-00

S-00

O-00

N-00

D-00

J-01

F-01

M-01

A-01

M-01

J-01

J-01

A-01

S-01

perc

ent g

row

th p

er d

ay (g

ram

s/day

)

8/99weight/day

11/99weight/day

3/00weight/day

5/00weight/day

7/00weight/day

Fig. 3.1dAdmiral Reef, A. cervicornis Growth weight (g/d) for each sampling period

0

0.05

0.1

0.15

0.2

0.25

0.3

Jul-99A

ug-99

Sep-99O

ct-99

Nov-99

Dec-99

Jan-00

Feb-00M

ar-00

Apr-00

May-00

Jun-00Jul-00

Aug-00

Sep-00O

ct-00

Nov-00

Dec-00

Jan-01Feb-01

Mar-01

Apr-01

May-01

Jun-01Jul-01

Aug-01

Sep-01

gram

s/da

y

8/99 weight/day

11/99 weight/day

3/00 weight/day

5/00 weight/day

7/00 weight/day

60

Fig. 3.2aAdmiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length (mm/d) for Aug 1999 transplants

0

0.001

0.002

0.003

0.004

0.005

0.006

0.007

0.008

0.009

Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01Sep-01

perc

ent g

row

th (m

m/d

, g/d

)

8/99 length

8/99 weight

Fig. 3.2bAdmiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length (mm/d) for Nov 1999 transplants

0

0.001

0.002

0.003

0.004

0.005

0.006

0.007

0.008

0.009

Jul-99

Aug-99

Sep-99

Oct-99

Nov-99

Dec-99

Jan-00Feb-00

Mar-00

Apr-00

May-00

Jun-00

Jul-00

Aug-00

Sep-00

Oct-00

Nov-00

Dec-00

Jan-01

Feb-01M

ar-01

Apr-01

May-01

Jun-01

Jul-01

Aug-01

Sep-01

perc

ent g

row

th (m

m/d

, g/d

)

11/99 length

11/99 weight

61

Fig. 3.2cAdmiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length (mm/d) for March 2000 transplants

0

0.005

0.01

0.015

0.02

0.025

0.03

Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01Sep-01

perc

ent g

row

th (m

m/d

,g/d

) 3/00 length

3/00 weight

Fig. 3.2dAdmiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length (mm/d) for May 2000 transplants

0

0.002

0.004

0.006

0.008

0.01

0.012

0.014

0.016

0.018

Jul-99A

ug-99Sep-99O

ct-99N

ov-99D

ec-99Jan-00Feb-00M

ar-00A

pr-00M

ay-00Jun-00Jul-00A

ug-00Sep-00O

ct-00N

ov-00D

ec-00Jan-01Feb-01M

ar-01A

pr-01M

ay-01Jun-01Jul-01A

ug-01Sep-01

perc

ent g

row

th (m

m/d

, g/d

)

5/00 length

5/00 weight

62

Fig. 3.2eAdmiral Reef, A. cervicornis Percent growth weight (g/d) and percent growth length (mm/d) for Aug 2000 transplants

0

0.002

0.004

0.006

0.008

0.01

0.012

0.014

0.016

0.018

Jul-99

Aug-99

Sep-99O

ct-99

Nov-99

Dec-99

Jan-00

Feb-00

Mar-00

Apr-00

May-00

Jun-00

Jul-00

Aug-00

Sep-00

Oct-00

Nov-00

Dec-00

Jan-01

Feb-01

Mar-01

Apr-01

May-01

Jun-01Jul-01

Aug-01

Sep-01

perc

ent g

row

th (m

m/d

, g/d

)

8/00 length

8/00 weight

Fig. 3.3aLittle Grecian ReefPercent growth length (mm/d) of original size for each sampling period

0.000

0.002

0.004

0.006

0.008

0.010

0.012

0.014

0.016

0.018

J-99F-99M

-99A

-99M

-99J-99J-99A

-99S-99O

-99N

-99D

-99J-00F-00M

-00A

-00M

-00J-00J-00A

-00S-00O

-00N

-00D

-00J-01F-01M

-01A

-01M

-01J-01J-01A

-01S-01

perc

ent g

row

th/d

ay (m

m/d

ay)

6/99 %length/day

11/99 % length/day

5/00 %length/day

8/00 %length/day

11/00 %length/day

63

Fig. 3.3bLittle Grecian ReefTotal Growth (length, mm/day) for each sampling period

0.0

0.5

J-99F-99M

-99A

-99

M-99

J-99J-99A

-99

S-99O

-99N

-99D

-99

J-00F-00M

-00A

-00

M-00

J-00J-00A

-00S-00

O-00

N-00

D-00

J-01

F-01M

-01A

-01M

-01

J-01J-01A

-01S-01

1.0

1.5

2.0

2.5

3.0m

m/d

ay

64

6/99 length/day

11/99 length/day

5/00 length/day

8/00 length/day

11/00 length/day

Fig. 3.3cLittle Grecian ReefPercent growth weight (grams/day) of original size for each sampling period

0.000

0.002

0.004

0.006

0.008

0.010

0.012

0.014

J-99F-99M

-99A

-99M

-99J-99J-99A

-99S-99O

-99N

-99D

-99J-00F-00M

-00A

-00M

-00J-00J-00A

-00S-00O

-00N

-00D

-00J-01F-01M

-01A

-01M

-01J-01J-01A

-01S-01

perc

ent g

row

th/d

ay

6/99 %wt/day

11/99 %wt/day

5/00 %wt/day

8/00 %wt/day

11/00 %wt/day

Fig. 3.3dLittle Grecian Reef, A. cervicornisTotal growth weight (grams,day) for each samling period

0.00

0.05

J-99F-99M

-99A

-99M

-99J-99J-99A

-99S-99O

-99N

-99D

-99J-00F-00M

-00A

-00M

-00J-00J-00A

-00S-00O

-00N

-00D

-00J-01F-01M

-01A

-01M

-01J-01J-01A

-01S-01

0.10

0.15

0.20

0.25

0.30to

tal g

row

th/d

ay (g

ram

s/day

)

65

6/99weight/day

11/99weight/day

5/00weight/day

8/00weight/day

11/00weight/day

Fig. 3.4aLee Stocking Island, A. cervicornisPercent growth (length, mm/day) of original size for each sampling period

0.000

0.003

0.006

0.009

0.012

0.015

0.018

J-99

M-99

M-99

J-99

S-99

N-99

J-00

M-00

M-00

J-00

S-00

N-00

J-01

M-01

M-01

J-01

S-01

perc

ent g

row

th/d

ay

1/99 %length/day

5/99 %length/day

8/99 %length/day

11/99 %length/day

3/00 %length/day

Fig. 3.4bLee Stocking Island, A. cervicornisGrowth length (mm/day) for each sampling period

0

0.5

J-99

M-99

M-99

J-99

S-99

N-99

J-00

M-00

M-00

J-00

S-00

N-00

J-01

M-01

M-01

J-01

S-01

1

1.5

2

2.5

3

3.5

4

4.5m

m/d

ay

66

1/99length/ day5/99length/ day8/99length/ day11/99length/ day3/00length/ day

Fig. 3.4cLee Stocking Island, A. cervicornisPercent growth (weight, g/day) of original size for each sampling period

0.000

0.005

0.010

0.015

0.020

0.025

0.030

J-99F-99M

-99A

-99M

-99J-99J-99A

-99S-99O

-99N

-99D

-99J-00F-00M

-00A

-00M

-00J-00J-00A

-00S-00O

-00N

-00D

-00J-01F-01M

-01A

-01M

-01J-01J-01A

-01S-01

perc

ent g

row

th/d

ay

1/99 %wt/day

5/99 %wt/day

8/99 %wt/day

11/99 %wt/day

3/00 %wt/day

Fig. 3.4dLee Stocking Island, A. cervicornisGrowth weight (g/day) for each sampling period

0

0.04

J-99F-99M

-99A

-99M

-99J-99J-99A

-99S-99O

-99N

-99D

-99J-00F-00M

-00A

-00M

-00J-00J-00A

-00S-00O

-00N

-00D

-00J-01F-01M

-01A

-01M

-01J-01J-01A

-01S-01

0.08

0.12

0.16

0.2gr

ams/d

ay

67

1/99weight/day

5/99weight/day

8/99weight/day

11/99weight/day

3/00weight/day

Fig. 3.5All Sites (A. cervicornis, A. prolifera )Total grams/total mm

0

0.05

0.1

0.15

0.2

0.25

0.3

Jan-99

Mar-99

May-99

Jul-99

Sep-99

Nov-99

Jan-00

Mar-00

May-00

Jul-00

Sep-00

Nov-00

Jan-01

Mar-01

May-01

Jul-01

Sep-01

Nov-01

grow

th w

eigt

h/ g

row

th le

ngth

(gra

ms/

mm

)

ADM 8/99

ADM 11/99

ADM 3/00

ADM 5/00

ADM 8/00

LG 6/99

LG 11/99

LG 5/00

LG 8/00

LG 11/00

LSI 1/99

LSI 5/99

LSI 8/99

LSI 11/99

LSI 3/00

Prolifera

Fig. 3.6Lee Stocking Island, Normans Pond (daily averages)4/20/96-7/11/02 *no data for Jan 99-Oct 99data from <http://www.cmrc.org/lsi_seawater_temp.htm>

15N-95

F-96

M-96

A-96

N-96

J-97

A-97

J-97

O-97

J-98

A-98

J-98

O-98

J-99

A-99

J-99

O-99

J-00

A-00

J-00

68

20

25

30

35

* no data

Cel

cius

O-00

J-01

A-01

J-01

O-01

J-02

A-02

J-02

O-02

D-02

`

Fig. 3.7Molasses Reef Florida 1/98-9/02 (Daily Average Sea Temp) C-Man Station (CRAMP)data from <http://www.coral.noaa.gov/seakeys/hist_data.shtml>

15

20

25

30

35

Jan-98

Apr-98

Jul-98

Oct-98

Jan-99

Apr-99

Jul-99

Oct-99

Jan-00

Apr-00

Jul-00

Oct-00

Jan-01

Apr-01

Jul-01

Oct-01

Jan-02

Apr-02

Jul-02

Deg

rees

Cel

cius

Table 3.1Orginal Transplant Size v. growth Length (mm)p=.05 significanceSite Jan-99 May-99 Aug-99 Nov-99 Mar-00 May-00 Aug-00 Sep-00 Nov-00 Mar-01 May-01 Aug-01LG Jun99 X X U (0.46) U (0.17) U (0.26) U (0.21) U (0.38) U (0.26) U (0.38) U (0.45)

X X U (0.26) U (0.391) U (0.380) U (0.220)U (0.205) U (0.377) U (0.471)

X U (0.422) U (0.302) U (0.272) U (0.303)X U (0.301) U (0.471)

U (0.33) U (0.2) U (0.06) U (0.35) U (0.45) U (0.26) U (0.06)X U (0.060) U (0.080) U (0.212) U (0.212)

U (0.443) U (0.429) U (0.01)U (0.435) U (0.173) U (0.117) U (0.073) U (0.312)

X U (0.175) U (0.069) U (0.382)U (0.003) U (0.001) U (0.433) U (0.043)

X U (0.438) U (0.104) U (0.467) U (0.166)X U (0.040) X X

X U (0.380) U (0.396) U (0.006)X U (0.130) U (0.218) U (0.374)

U (0.21) U (0.02) U (0.04) U (0.28) U (0.02) U (0.4) U (0.001) U (0.005) U (0.22)Red=not significa p=.05)

X S (0.55)LG Nov99 X X X S (0.676) S (0.530) S (0.649)LG May00 X X X X X X S (0.804) S (0.558) S (0.557)LG Aug00 X X X X X X U (0.050)LG Nov00 X X X X X X X X S (0.527)ADM Aug99 X X X S (0.56) S (0.55)ADM Nov99 X X X S (0.648) S (0.787) S (0.427) S (0.534)ADM Mar00 X X X X X S (0.530) S (0.592) S (0.672) S (0.738)ADM May00 X X X X X X S (0.675)ADM July00 X X X X X X S (0.521) S (0.618)LSI Jan99 X S (0.66) S (0.56) S(0.541) X X X XLSI May99 X S (0.880) S (0.652) S (0.711) X X XLSI Aug99 X X S (0.927) S (0.979) S (0.974) X X XLSI Nov99 X X X S (0.683) S (0.73) S (0.842) X XLSI Mar00 X X X X S (0.675) S (0.592) X XLSI prolifera Jan99 X X XLG=Little Grecian nt (ADM=Admiral Reef Blue=significant (p=.05)LSI=Lee Stocking Island

69

Table 3.2Orginal Transplant Size v. Weight Growth (grams)p=.05 significanceSite Jan-99 May-99 Aug-99 Nov-99 Mar-00 May-00 Aug-00 Sep-00 Nov-00 Mar-01 May-01 Aug-01LG Jun99 X X U (0.484) U (0.004)

X U (0.119)X U (0.471)

U (0.234) U (0.020)U (0.482) U (0.075) U (0.081) U (0.358) U (0.359)

X U (0.303)U (0.401) U (0.000)

XU (0.396) U (0.370) U (0.172)

U (0.539)X X

U (0.271)U (0.426)

U (0.288) U (0.167) U (0.015)red=no ignificant (p=.05)

S (0.778) X S (0.728) S (0.837) S (0.812) S (0.933) S (0.818) S (0.892)LG Nov99 X X X X S (0.764) S (0.60) S (0.802) S (0.651) S (0.607) S (0.897)LG May00 X X X X X S (0.862) S (0.911) S (0.837) S (0.977) S (0.937)LG Aug00 X X X X X X X S (0.649) S (0.655) S (0.844) S (0.889) S (0.872)LG Nov00 X X X X X X X X X S (0.884)ADM Aug99 X X X S (0.554) S (0.629) S (0.403) S (0.774)ADM Nov99 X X X S (0.648) S (0.530) S (0.950) S (0.627) S (0.827) S (0.526) S (0.866)ADM Mar00 X X X X X S (0.973) S (0.887) S (0.966) S (0.915) S (0.827)ADM May00 X X X X X S (0.819) S (0.736) S (0.636) S (0.894) S (0.968) S (0.779)ADM July00 X X X X X X X S (0.917) S (0.602)LSI Jan99 X S (0.94) S (0.904) S (0.899) S (0.591) S (0.965) S (0.909) S (0.54) X X X XLSI May99 X X S (0.685) S (0.989) S (0.804) S (0.834) S (0.897) S (0.670) X X XLSI Aug99 X X X S (0.767) S (0.912) S (0.998) S (0.992) X X XLSI Nov99 X X X X S (0.912) S (0.901) S (0.922) S (0.885) S (0.861) X XLSI Mar00 X X X X X S (0.905) S (0.869) S (0.601) S (0.637) X XLSI prolifera Jan99 X S (0.851) S (0.704) S (0.812) S (0.866) S (0.707) S (0.869) X XLG=Little Grecian t sADM=Admiral Reef Blue=significant (p=.05)LSI=Lee Stocking Island

70

CHAPTER 4

EFFECT OF SEASON AND CORAL SIZE ON THE SURVIVAL OF TRANSPLANTED

ACROPORA CERVICORNIS 1


Abstract

Survival of fragments of the staghorn coral, Acropora cervicornis, transplanted seasonally in

PVC-holders to cement blocks adjacent to their collection site, was highest in the Bahamas and

in the Florida Keys in the spring and early summer when water temperatures were moderate (23-

28oC). Survival rates decreased at both sites during the summertime, when temperature was

highest. Transplants made during late summer (August-September) exhibited lower survival

over the first 3-month recovery period than at any other time during the year. Survival of

transplants in 1999-2000 was higher in the Florida Keys compared to the Bahamas. Smaller

fragments (< 6 cm) exhibited higher mortality and lower growth rates compared to larger

fragments (> 6cm). Reef restoration projects might optimize survival by transplanting larger

pieces of Acropora cervicornis during seasons with moderate seawater temperatures, avoiding

the high-temperatures of late summer and early fall.

Key words: Acropora cervicornis, restoration, transplantation, mortality

72

Introduction

Survival of transplanted Acroporid corals has been studied since the early 1900’s (Mayer

1914, Vaughn 1915) and has come more into the mainstream in the 1960’s when Ginn Shinn,

with Shell Oil Company, transplanted A. cervicornis to several different areas in the Florida

Keys (Shinn 1966). Since then hundreds of transplantation experiments have taken place

worldwide. In this study the staghorn coral A. cervicornis and the hybrid A. prolifera were

transplanted in the Florida Keys and Bahamas to study the effects of season and size on the

survival of transplanted colonies.

73

Methods

Transplantation experiments were conducted on a patch reef due north of Lee Stocking

Island (LSI), Bahamas and at two sites off Key Largo, Florida; Admiral Reef (ADM) (N 25°

02.732’ W 080° 23.675’) a 3m deep inshore patch reef and Little Grecian Reef (LG) (N

25°07.164’ W 080° 18.107’) a 3m deep offshore reef crest. Artificially broken fragments of

Acropora cervicornis were transplanted four times corresponding to the seasons and monitored

quarterly between 1999 and 2001.

In the Bahamas, coral colonies were transplanted January, May, and August of 1999 and

March of 2000. Florida coral colonies were transplanted first at Little Grecian Reef on June 17,

1999, then both Little Grecian and Admiral Reef in August and November 1999, just Admiral

reef on March 12, 2000 (Little Grecian had 6-8ft seas), then both sites in May and August 2000

and finally just Little Grecian Reef on November 29, 2000. Due to its scarcity, Acropora

prolifera was transplanted only once in the Bahamas on February 21, 1999 and monitored

quarterly until May 2001.

Fifteen A. cervicornis and A. prolifera fragments, ranging in size from 25 mm to 200

mm, were glued to PVC pipe and attached to cement bricks adjacent to the reef they were

collected. Transplants were glued into a 2″ section of ½ ″ PVC with Oately® epoxy putty. The

weight of the epoxy and PVC pipe were measured so that the initial weight of the transplant

could be determined.

Fragments were weighed using the Buoyant Weight method in reef water to the nearest

.001 gram. Temperature (25oC) and salinity (36 ppt) were held constant in the weighing

apparatus for every season. Length was measured to the nearest millimeter using vernier

74

calipers. Growth was measured from the top of the PVC or where the live tissue starts at the

base. Mortality was measured when a transplanted colony had lost all tissue. Graphs were

generated in Excel, with 95% confidence intervals.

75

Results

The rates of survival of transplants of A. cervicornis as indicated by the slopes of the

lines in Figure 1, as well as survival over the course of the study (1999-2001), was higher in the

Florida Keys compared to the Bahamas site (Figure 4.1). However, survival of smaller colonies

was usually much lower than that for larger transplants (> 4cm) at all sites except Admiral Reef

in the Florida Keys (Figures 4.2, 4.3 a-e). All transplants (100%) of A. prolifera survived during

the experiment.

Florida transplants appeared to be more prone to mortality during winter months (cold

temperatures), compared to summer months (warmer months) (Figures 4.1). Bahamas

transplants experienced more mortality during summertime high-temperature periods compared

to low-temperature periods (Figure 4.4). Relatively lower survival was observed at both sites

during summertime transplants when initial survival (first three months) was compared to

seasonal seawater temperature (Figure 4.4 a,b).

76

Discussion

Survival of acroporid corals transplanted to similar reef environments is widely reported

as ‘variable’, often with no clear environmental or physiological correlations to help with

interpretations of the data (e.g. Alcala et al. 1982, Auberson 1982,). A number of factors appear

to negatively affect survival of transplanted corals, including natural events such as storms,

predation, disease, and various physiological parameters such as size of the transplant and

physiological stress associated with seasonal temperature.

While storms and predacious corallivorous snails and fire worms can seriously reduce

cover and numbers of individual coral colonies (NMFS web site, Sept. 2002), only a few studies

have good data documenting effects of storms (Tunnicliffe 1983) and coral predators (Miller

2001). For instance, Bak and Criens (1982) found that survival of transplants depends on their

stability, increasing when A. cervicornis fuses to other nearby branches thereby reducing

scouring by sediments and rolling of small pieces on the benthos (e.g. Woodley 1992). We made

similar observations with the passage of Tropical Storm Harvey and Hurricane Irene by Key

Largo during the fall of 1999; virtually all PVC-cemented transplants placed on anchored cement

bricks survived 20 foot waves on top of Little Grecian Reef, while most natural colonies in the

surrounding rubble perished.

It seems logical that larger transplants might survive better than small pieces of coral.

Smith and Hughes (1999) documented about 30% survival of larger pieces of Acropora

intermdia 17 months post-transplant compared to only 8% of smaller pieces. Colonies

transplanted from the reef flat survived better than those placed on the reef crest and on the outer

slope (Smith and Hughes 1999). However, Yap (1998) found no significant differences in

77

survivorship between two sizes of Porites rus. Rinkevich (2000) found high survivorship of

Styllophora pistillata, irregardless of whether transplanted pieces were small (<4 cm) or larger.

If size is important in survival, then relative growth rates leading to larger individuals is

also important to increased survival. Porites cylindrical in shallow water grew more rapidly than

colonies in deeper water (Yap 1998), a pattern prevalent amongst the photo-synthetically active

symbiotic corals.

High survivorship clearly depends on an optimal temperature range for growth. In the

current study lower survival was documented at high (>300C) temperatures common during the

late summer at both sites (Figure 4) and elsewhere in the Caribbean,

(http://www.cmrc.org/lsi_seawater_temp.htm). Colonies of Acropora cervicornis living at the

higher latitudes of the Florida Keys also exhibit higher mortality during the coldest winter

months (Figure 4.1) usually with lower survival amongst smaller (<4cm) transplants (Figure

4.2,4.3) (Shinn 1966, Porter et al. 1982). Yap and Gomez (1984) found that Acropora pulchra

transplants respond to temperature and day length such that longer light and higher temperatures

result in more mortality.

78

References

Alcala, A. C., E. D. Gomez, et al. (1982). “Survival and growth of coral transplants in

central Philippines.” Kalikasan 11(1): 179-184.

Auberson, B. “Preliminary note on morphological diversity in colonial transplants of six coral species”

Bak, R. P. M. and S. R. Criens (1982). “Experimental Fusion in Atlantic Acropora (Scleractinia).” Marine Biology Letters 3(2): 67-72.

Mayer, A.G. (1914). “The effects of temperature on tropical marine animals”. Carnegie

Inst. Washington D.C. 183: 1-24.

Miller, M. W. (2001). “Corallivorous snail removal: evaluation of impact on Acropora palmata.” Coral Reefs 19(3): 293-295.

NMFS Website :http://www.nmfs.noaa.gov/ Porter JP, JF Battey and GJ Smith (1982) Perturbation and change in coral reef

communities. Proc natl Acad Sci 79:1678-1681 Rinkevich, B. (2000). “Steps towards the evaluation of coral reef restoration by using small branch fragments.” Marine Biology 136(5): 807-812. Shinn, E.A. (1966). “Coral growth-rate, an environmental indicator” J. Paleontology 40:233-240.

Smith, L. D. and T. P. Hughes (1999). “An experimental assessment of survival, re-attachment and fecundity of coral fragments.” Journal of Experimental Marine Biology & Ecology 235(1): 147-164.

Tunnecliffe, V. (1983) “Caribbean staghorn coral populations: pre-Hurricane Allen

conditions in Discovery Bay, Jamaica” Bulletin of Marine Sciences (33)1: 132-151. Woodley, J. D. (1992). “The incidence of hurricanes on the north coast of Jamaica since

1870: Are the classic reef descriptions atypical?” Hydrobiologia 247(1-3): 133-138. Vaughan, T.W. (1915) The geological significance of the growth-rates of Floridian shoal-

water corals” Washington Acad. Sci. 5:591-600

Yap, H. T. and E. D. Gomez (1984). “Growth of Acropora-Pulchra .2. Responses of Natural and Transplanted Colonies to Temperature and Day Length.” Marine Biology 81(2): 209-215.

79

Yap, H. T., R. M. Alvarez, et al. (1998). “Physiological and ecological aspects of coral transplantation.” Journal of Experimental Marine Biology and Ecology 229(1): 69-84.

Yap, H. T. and E. D. Gomez (1985). “Growth of Acropora-Pulchra 3. Preliminary Observations on the Effects of Transplantation and Sediment on the Growth and Survival of Transplants.” Marine Biology (Berlin) 87(2): 203-210.

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Fig. 4.1Tranplant Survival (Acropora cervicornis ) Little Grecian Reef, FLA (LG), Lee Stocking Island, Bahamas (LSI), Admiral Reef , FLA (Adm)

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LG Nov1999

LG May00

LG Aug00

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LSI Aug99

LSI Nov99

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Adm Aug99

Adm Nov99

Adm Mar00

Adm May00

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Fig. 4.2LSI percent survival vs size for all transplants (Jan 1999-May 2001)

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.5-2 (n=23) 2-7 (n=32) 7-25 (n=21)Size class (bouyant g)

% su

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Fig 4.3aSurvival of corals in relation to initial size transplantedLittle Grecian Reef June 1999 Tranplants

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Fig 4.3bSurvival of corals in relation to initial size transplantedLittle Grecian Reef Nov 1999 Tranplants

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(n=12)

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Fig 4.3cSurvival of corals in relation to intial size transplantedLittle Grecian Reef May 2000 Tranplants

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Fig 4.3dSurvival of corals in relation to initial size transplantedLittle Grecian Reef AUG 2000 Tranplants

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Fig 4.3eSurvival of corals in relation to initial size transplantedLittle Grecian Reef Nov 2000 Tranplants

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Fig. 4.4a Survival at first sampling period LSI, Bahamas

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Fig. 4.4bFlorida (Admiral)Survival at first sampling period (Number of days in period in parenthesis)

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CHAPTER 5

SCIENTIFICALLY PRODUCED LESIONS IN REEF CORALS: SCARRED FOR LIFE? 1


86

ABSTRACT

The recovery rates of scientifically produced scars or lesions were investigated in the Caribbean

reef coral Montastrea faveolata. Fastest rates of re-growth occurred in the fall and spring; and

were the slowest during the winter. Colonies appeared to stop re-growth during the late fall

when temperatures were highest on the reef. Artificial lesions on Montastrea faveolata filled

with epoxy took approximately twice the recovery time as those allowed to recovery without the

use of filler compounds; however differences in growth rates were only seen in the first three

months.

87

INTRODUCTION

Destructive sampling of live corals provides information on coral physiology and the

health of reef corals that cannot be obtained by non-destructive methods. Pieces of coral colonies

can be collected with a steel punch, chipping with hammer and chisel or hydraulic corer.

Sampling the coral colony, regardless of technique, will leave a lesion or scar that may or may

not fully recover.

Studies designed to investigate recovery of artificial damage on whole colonies have

found that, recovery rates are species- specific (Meesters et al 1997) as well as morphology-

specific. For instance, Hall (1997) documented fastest to slowest recovery of arborescent, bushy,

tabular, massive, and submassive colony morphologies (Hall, 1997). In addition, massive corals

recover from injuries quicker if the entire injury is surrounded by live tissue, in contrast to

injuries occurring on the edge of the colony. More recently, Loya et al. (2002) has noted that

photo synthetically-fixed carbon appears to be translocated from nearby healthy tissue towards

the site of injury, implying whole colony recognition and response to such injuries.

In recent years underwater epoxy has been used to fill , injuries and lesions, but no

published studies have been conducted to determine if this practice is beneficial to the recovery

of the coral colony or not. In this study, sampling lesions were either filled with epoxy or

allowed to heal without the addition of filler compounds in order to see which recovered the

fastest.

88

METHODS

Lesions were generated in the Florida Keys and the Bahamas as part of an ongoing multi-

year study. Several colonies of M. faveolata were labeled in May 2000 and photographed again

on August 2000, while five others were labeled March 2001 and photographed May 2002 and

two of those five were found and photographed in Aug 2002. In Florida other species, S. siderea

and Porites asteroides were photographed on may 2002 and again on Aug 2002. All lesions in

Florida were allowed to recover naturally without filler compounds.

Colonies of the boulder coral Montastrea faveolata were sampled at Lee Stocking Island,

Bahamas and Admiral Reef, an inshore reef near Key Largo, Fla. In the Bahamas one M.

faveolata colony, at the depth of 3m, contained ten lesions produced for a different study on

August 13, 2000. Six of these lesions were filled completely with Z-Spar® epoxy while the

other four lesions were not filled, except for a very small amount of epoxy used to attach small

numbered tags to each lesion. Care was taken to place the tag in the center of each lesion to

minimize the impact of the tag and epoxy on the control lesions. It was essential to label the

lesions so that pictures could be taken over the course of the experiment.

A Nikonos III, 35mm lens, and close up kit were used to photograph each lesion.

Photographs were scanned and digitized and then analyzed using NIH shareware ImageJ.

Perimeter and surface area of the lesions were calculated. The resulting data was analyzed using

Statview statistical software. A student T-test was used to see if initial lesion size was

statistically different between the control and experimental regimes.

89

RESULTS

Non-filled lesions recover faster than epoxy-filled lesions (Fig 5.1, Table 5.1). Within

the three months, the control lesions had recovered 61% (+/- 4%) while the epoxy-filled lesions

had only recovered 12% (+/- 4%). This lag in recovery is evident throughout the study and by

seven months (March 2001) the epoxy-filled lesions (19% +/- 8% recovery) are still well behind

the control lesions (67% +/- 7% recovery). A linear regression drawn for both data sets minus

the initial starting point suggests that full recovery of epoxy-filled lesions takes approximately

twice the amount of time as control lesions (1.0 – 2.0 yrs). While only 6 months of data was

analyzed from the Florida Keys (Table 5.2) these corals show similar recovery rates to those

observed in the Bahamas for non-epoxy filled lesions (1.0-2yrs).

90

DISCUSSION

Filling sample lesions with epoxy significantly retards the recovery in the coral

M. faveolata. Rates of tissue growth of epoxy-filled lesions were half of those allowed to

recover without any filler materials. Interestingly, only the initial recovery rates were

significantly different; after 3 months recovery rates for epoxy and non-epoxy filled lesions were

not significantly different (Fig. 5.1). There are several explanations of the initial slow growth of

the epoxy-filled lesions. First, coverage of the damaged area with epoxy may hinder the release

of chemical cues emanating from the damaged tissue and thus slow the recovery of the coral

(ref). Second, the epoxy, although non-toxic when cured, may release water soluble chemicals

that could retard tissue growth around the lesions. Although the epoxy-fixed tags used in this

particular experiment were placed on both filled and non-filled lesions, reduction of initial

recovery rates was only evident for the former colonies. Tissue re-growth into the non-filled

lesions made contact with the epoxy-fixed labels only after nine months into the experiment and

rates of re-growth were not negatively impacted at that time, suggesting leeching of toxic

chemicals during the curing process as the most likely source of the phenomenon.

Tissue re-growth into lesions resulting from scientific collecting appears to mirror

recovery from other forms of injury and damage. Recent literature has suggested several modes

of recovery from damage. For instance type, size and location of lesion has been shown to

significantly influence all influence the recovery in some corals and not in others. Meesters et al

(1994) suggest in the species Montastrea annularis, recovery is fuelled directly from the polyps

that border the damaged area and is not dependant upon colony size (localized regeneration

hypothesis). Also, Lirman (2000) also found, that in the branching coral Acropora palmata,

91

colony size did not affect recovery rate; however, the initial size of lesions seemed to correlate

with recovery, with; the smallest lesions recovering the fastest. C-14 labeling has shown that

translocation of materials from other parts of the colony move into the damaged area. Oren

(1997) found that lesion size is correlated with translocation of materials from other areas than

those that border the lesion in Favia favus; he suggests that except for small lesions (>1cm2) the

larger the lesion size the further the C-14 is translocated for repair. These differences may be

explained related to morphology of the different species by Hall’s (1997) paper that found

different recovery rates of different morphological types of corals:

(i.e.arborescent>bushy>tabular>massive>submassive).

Seasonality may also influence speed of recovery. Kramarsky-Winter and Loya (2000)

found that in the Fungid coral granulose small corals (>5cm) and not sexually mature did not

repair regardless of season or water temp while large corals (<5cm) repaired at different rates

during different seasons and virtually stopped in the spring when the corals become fecund and

all energy is diverted into reproduction. This is a small single polyp coral that is a brooder and

not a broadcast spawner as in Montastrea faveolata. These different types of reproduction could

influence the mode recovery and where energy is diverted: i.e. reproduction or recovery.

Hall (1997) found that the type of injury (scraping, tissue injury, or regrowth of a whole branch)

affects the recovery rate in different morphotypes of corals. Repair of scraping injuries was

greater than tissue injuries, while regrowth of a new branch was slowest of all. With the

exception of Porites mayeri which repaired more of its central injury than its edge injury,

recovery rate of central and edge injuries was not significantly different for Acropora robusta, A.

hyacinthus, A. palifera, Pocillopora damcornis, and Porites lichen. These results imply that the

92

amount of damage caused by an injury differs, both for the species it is inflicted upon, and the

type of injury generated.

Since little scientific experimentation has been published to support sampling recovery,

resource managers need to take into account type of coral, size of lesion, location of lesion and

whether filling with compounds will be beneficial to the recovery of sample lesions. As shown

by our data, filling lesions in M. faveolata seems to retard the recovery in this species yet

resource managers are requiring scientists to fill sample lesions.

93

REFERENCES

Hall, V. R. (2001). “The response of Acropora hyacinthus and Montipora tuberculosa to three different types of colony damage: scraping injury, tissue mortality and breakage.” Journal of Experimental Marine Biology and Ecology 264(2): 209-223.

Hall, V. R. (1997). “Interspecific differences in the regeneration of artificial injuries on

scleractinian corals.” Journal of Experimental Marine Biology and Ecology 212(1): 9-23. Kramarsky-Winter, E. and Y. Loya (2000). “Tissue regeneration in the coral Fungia

granulosa: the effect of extrinsic and intrinsic factors.” Marine Biology 137(5-6): 867-873. Lirman, D., P. W. Glynn, et al. (2001). “Combined effects of three sequential storms on

the Huatulco coral reef tract, Mexico.” Bulletin of Marine Science 69(1): 267-278. Lirman, D. (2000). “Fragmentation in the branching coral Acropora palmata (Lamarck):

growth, survivorship, and reproduction of colonies and fragments.” Journal of Experimental Marine Biology and Ecology 251(1): 41-57.

Meesters, E. H., W. Pauchli, et al. (1997). “Predicting regeneration of physical damage on a reef-building coral by regeneration capacity and lesion shape.” Marine Ecology-Progress Series 146(1-3): 91-99.

Meesters, E. H., M. Noordeloos, et al. (1994). “Damage and Regeneration - Links to Growth in the Reef-Building Coral Montastrea Annularis.” Marine Ecology-Progress Series 112(1-2): 119-128.

Oren, U., B. Rinkevich, et al. (1997). “Oriented intra-colonial transport of C-14 labeled

materials during coral regeneration.” Marine Ecology-Progress Series 161: 117-122.

94

Table 5.1. Percent recovery of the epoxy-filled lesion from the initial damage (Bahamas). Percent Growth I.D.

Number Epoxy Filled

Lesions

Aug 00 - Nov 00

(3 months)

Aug 00 -March 01 (7 months)

Aug 00 -May 01

(9 months)

Aug 00 – Sept 01

(13 months)

Aug 00 - Dec 01

(16 months)

Aug 00 -June 02

(22 months)

599 16 28 27 38 34 58 592 7 14 27 41 54 81 584 8 21 39 53 69 88 568 14 24 28 52 70 90 563* n.d. 0 1 0 13 14 564 15 25 36 47 61 82

Average 12 19 26 38 50 69 Stddev 4 11 13 20 23 29 95%ci 4 8 11 16 18 23

95

Table 5.2. Percent recovery of the control lesions from the initial damage (Bahamas). Percent Growth I.D.

Number Non-filled

Lesions

Aug 00 - Nov 00

(3 months)

Aug 00 -March 01 (7 months)

Aug 00 -May 01

(9 months)

Aug 00 – Sept 01

(13 months)

Aug 00 - Dec 01

(16 months)

Aug 00 -June 02

(22 months) 586 58 61 75 85 90 Recovered 562 65 78 85 92 85 Recovered 578 56 66 69 76 88 Recovered 598 60 64 67 71 76 81

Average 61 67 74 81 85 95 stddev 4 8 8 9 6 10 95%ci 4 7 8 9 6 9

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Fig. 5.1aRecovery of sampling scars Montastrea faveolata LSI, Bahamas Aug 2000(initial lesion) - June 2000

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Fig. 5.1bRecovery of sampling scars Montastrea faveolata with linear regression LSI, Bahamas Aug 2000(initial lesion) - June 2000

R2 = 0.9587

R2 = 0.9965

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Fig. 5.2Admiral Reef, Florida, Montastrea faveolataMF1,MF2,MF4 March 2001 (initial damage) - Aug 2001

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Fig. 5.3Monthly growth rates for each seasonMontastrea faveolata , LSI, Bahamas

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Documents

Growth and Survival of Acropora Cervicornis · Acropora cervicornis fragments were transplanted seasonally (every 3 months) near Key Largo, Florida (patch reef-Admiral Reef and fore