<?xml version="1.0" encoding="ISO-8859-1"?><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance">
<front>
<journal-meta>
<journal-id>0122-9761</journal-id>
<journal-title><![CDATA[Boletín de Investigaciones Marinas y Costeras - INVEMAR]]></journal-title>
<abbrev-journal-title><![CDATA[Bol. Invest. Mar. Cost.]]></abbrev-journal-title>
<issn>0122-9761</issn>
<publisher>
<publisher-name><![CDATA[INSTITUTO DE INVESTIGACIONES MARINAS Y COSTERAS "JOSE BENITO VIVES DE ANDRÉIS" (INVEMAR)    INSTITUTO DE INVESTIGACIONES MARINAS Y COSTERAS -JOSE BENITO VIVES DE ANDRÉIS- (INVEMAR)]]></publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id>S0122-97612004000100011</article-id>
<title-group>
<article-title xml:lang="en"><![CDATA[ANNUAL SKELETAL EXTENSION OF TWO REEF-BUILDING CORALS FROM THE COLOMBIAN CARIBBEAN SEA]]></article-title>
<article-title xml:lang="es"><![CDATA[Extensión esquelética anual de dos corales formadores de arrecifes del Mar Caribe colombiano]]></article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Charry]]></surname>
<given-names><![CDATA[Henry]]></given-names>
</name>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Alvarado]]></surname>
<given-names><![CDATA[Elvira M.]]></given-names>
</name>
<xref ref-type="aff" rid="A01"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Sánchez]]></surname>
<given-names><![CDATA[Juan A.]]></given-names>
</name>
<xref ref-type="aff" rid="A02"/>
</contrib>
</contrib-group>
<aff id="A01">
<institution><![CDATA[,Universidad de Bogotá Jorge Tadeo Lozano Centro de Investigaciones Científicas (Museo del Mar) ]]></institution>
<addr-line><![CDATA[Bogotá ]]></addr-line>
<country>Colombia</country>
</aff>
<aff id="A02">
<institution><![CDATA[,Universidad de los Andes Departamento de Ciencias Biológicas ]]></institution>
<addr-line><![CDATA[Santafé de Bogotá ]]></addr-line>
<country>Colombia</country>
</aff>
<pub-date pub-type="pub">
<day>00</day>
<month>12</month>
<year>2004</year>
</pub-date>
<pub-date pub-type="epub">
<day>00</day>
<month>12</month>
<year>2004</year>
</pub-date>
<volume>33</volume>
<numero>1</numero>
<fpage>209</fpage>
<lpage>222</lpage>
<copyright-statement/>
<copyright-year/>
<self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_arttext&amp;pid=S0122-97612004000100011&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_abstract&amp;pid=S0122-97612004000100011&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_pdf&amp;pid=S0122-97612004000100011&amp;lng=en&amp;nrm=iso"></self-uri><abstract abstract-type="short" xml:lang="en"><p><![CDATA[The skeletal growth of the scleractinian corals Diploria labyrinthiformis (Linnaeus 1758) and Montastraea annularis (Ellis and Solander 1786) from Isla Grande (north of Rosario islands), Colombian Caribbean, was determined from annual extension increments revealed by X-radiography of 4-6 mm thick slabs obtained along the axis of maximum growth. The skeletal extension average rate for the last 22 years for D. labyrinthiformis was 3.8 mm year-1 (SE 0.10; N = 87). The correlation between growth of D. labyrinthiformis and hours of sunlight was significantly negative. The annual variances of sunlight enhanced annual statistically significant differences of growth in this species. M. annularis showed an average extension growth of 10.6 mm year -1 (SE 0.32; N = 55) during the last 14 years. The increase of M. annularis growth was coincident with the end of nearby dredging activities as well as the decrease of nutrient levels. Nevertheless, the growth rate of this species showed no statistically significant differences through the years and no correlation with variation in sunlight. A low density annual band, wider than high density band in D. labyrinthiformis and narrower in M. annularis, seems to form approximately during April and May in both species, which is coincident with the end of dry season (increase of sea surface temperature, decrease of hours of sunlight and high influence of Dique Channel waters over Rosario islands). The annual bands forming on both species at the study site might be highly related with time of reproduction of each species. Further investigation will permit a better understanding of how some local conditions or coral physiology are related with coral growth at Rosario islands.]]></p></abstract>
<abstract abstract-type="short" xml:lang="es"><p><![CDATA[El crecimiento del esqueleto de los corales escleractíneos Diploria labyrinthiformis (Linnaeus 1758) y Montastraea annularis (Ellis & Solander 1786) en Isla Grande (zona norte de las islas del Rosario), Caribe colombiano, se determinó a partir de los incrementos anuales en la extensión revelados por rayos-X de láminas de 4-6mm de grosor obtenidas a lo largo de los ejes de máximo crecimiento. La tasa promedio de extensión esquelética en los últimos 22 años para Diploria labyrinthiformis fue de 3.8 mm año-1 (ES 0.10; N= 87). La correlación entre crecimiento de D. labyrinthiformis y las horas solares fue significativamente negativa. Las variaciones anuales de brillo solar aumentaron diferencias anuales estadísticamente significativas en el crecimiento de esta especie. M. annularis mostró un promedio de extensión anual de 10.6 mm año-1 (ES 0.32; N= 55) durante los últimos 14 años. El incremento del crecimiento de M. annularis fue coincidente con la finalización de actividades de dragado en una localidad cercana y con la disminución de los niveles de nutrientes. No obstante, esta especie no mostró diferencias significativas a través de los años ni correlación con la variación de brillo solar. Una banda anual de baja densidad, más amplia que la de alta densidad en D. labyrinthiformis y más angosta en M. annularis, parece formarse aproximadamante durante abril y mayo en ambas especies, lo cual es coincidente con el final de la época seca (incremento en la temperatura del agua, disminución de horas de brillo solar y mayor influencia del Canal del Dique sobre las islas del Rosario). Las bandas anuales que se forman en las dos especies en el sitio de estudio podrían estar muy relacionadas con la época de reproducción de cada especie. Investigación adicional permitirá un mejor entendimiento de cómo algunas condiciones locales o la fisiología del coral están relacionadas con el crecimiento coralino en las islas del Rosario.]]></p></abstract>
<kwd-group>
<kwd lng="en"><![CDATA[Growth rates]]></kwd>
<kwd lng="en"><![CDATA[Diploria labyrinthiformis]]></kwd>
<kwd lng="en"><![CDATA[Montastraea annularis]]></kwd>
<kwd lng="en"><![CDATA[Caribbean Sea]]></kwd>
<kwd lng="en"><![CDATA[Colombia]]></kwd>
<kwd lng="en"><![CDATA[Coral reefs]]></kwd>
<kwd lng="es"><![CDATA[Tasas de crecimiento]]></kwd>
<kwd lng="es"><![CDATA[Diploria labyrinthiformis]]></kwd>
<kwd lng="es"><![CDATA[Montastraea annularis]]></kwd>
<kwd lng="es"><![CDATA[Mar Caribe]]></kwd>
<kwd lng="es"><![CDATA[Colombia]]></kwd>
<kwd lng="es"><![CDATA[Arrecifes coralinos]]></kwd>
</kwd-group>
</article-meta>
</front><body><![CDATA[ <p>&nbsp;</p>  <font face="verdana" size="2">      <p align="center"><font size="3"><b>ANNUAL SKELETAL EXTENSION OF TWO REEF-BUILDING CORALS FROM THE COLOMBIAN CARIBBEAN SEA</b></font> </p>      <p>&nbsp;</p>      <p align="center"><font size="3"><b>Extensi&oacute;n esquel&eacute;tica anual de dos corales formadores de arrecifes del Mar Caribe colombiano</b></font>.</p>      <p>&nbsp;</p>      <p><b>Henry Charry, Elvira M. Alvarado and Juan A. S&aacute;nchez </b></p>      <p>&nbsp;</p>  <hr size="1">      <p><b>ABSTRACT</b></p>      <p>The skeletal growth of the scleractinian corals <i><i>Diploria labyrinthiformis</i></i>    (Linnaeus 1758) and <i>Montastraea annularis</i> (Ellis and Solander 1786) from Isla    Grande (north of Rosario islands), Colombian Caribbean, was determined from    annual extension increments revealed by X-radiography of 4-6 mm thick slabs    obtained along the axis of maximum growth. The skeletal extension average rate    for the last 22 years for <i>D. labyrinthiformis</i> was 3.8 mm year&#8211;1 (SE 0.10;    N = 87). The correlation between growth of <i>D. labyrinthiformis</i> and hours of    sunlight was significantly negative. The annual variances of sunlight enhanced    annual statistically significant differences of growth in this species. <i>M. annularis</i>    showed an average extension growth of 10.6 mm year &#8211;1 (SE 0.32; N = 55)    during the last 14 years. The increase of <i>M. annularis</i> growth was coincident    with the end of nearby dredging activities as well as the decrease of nutrient    levels. Nevertheless, the growth rate of this species showed no statistically    significant differences through the years and no correlation with variation    in sunlight. A low density annual band, wider than high density band in <i>D. labyrinthiformis</i>    and narrower in <i>M. annularis</i>, seems to form approximately during April and May    in both species, which is coincident with the end of dry season (increase of    sea surface temperature, decrease of hours of sunlight and high influence of    Dique Channel waters over Rosario islands). The annual bands forming on both    species at the study site might be highly related with time of reproduction    of each species. Further investigation will permit a better understanding of    how some local conditions or coral physiology are related with coral growth    at Rosario islands.</p>      <p>KEY WORDS: Growth rates, <i>Diploria labyrinthiformis</i>, <i>Montastraea annularis</i>, Caribbean Sea, Colombia, Coral reefs</p>  <hr size="1">      ]]></body>
<body><![CDATA[<p><b>RESUMEN</b></p>      <p> El crecimiento del esqueleto de los corales escleract&iacute;neos <i>Diploria labyrinthiformis</i> (Linnaeus 1758) y <i>Montastraea annularis</i> (Ellis &amp; Solander    1786) en Isla Grande (zona norte de las islas del Rosario), Caribe colombiano,    se determin&oacute; a partir de los incrementos anuales en la extensi&oacute;n    revelados por rayos-X de l&aacute;minas de 4-6mm de grosor obtenidas a lo largo    de los ejes de m&aacute;ximo crecimiento. La tasa promedio de extensi&oacute;n    esquel&eacute;tica en los &uacute;ltimos 22 a&ntilde;os para <i>Diploria labyrinthiformis</i>    fue de 3.8 mm a&ntilde;o-1 (ES 0.10; N= 87). La correlaci&oacute;n entre crecimiento    de <i>D. labyrinthiformis</i> y las horas solares fue significativamente negativa.    Las variaciones anuales de brillo solar aumentaron diferencias anuales estad&iacute;sticamente    significativas en el crecimiento de esta especie. <i>M. annularis</i> mostr&oacute;    un promedio de extensi&oacute;n anual de 10.6 mm a&ntilde;o&#8211;1 (ES 0.32;    N= 55) durante los &uacute;ltimos 14 a&ntilde;os. El incremento del crecimiento    de <i>M. annularis</i> fue coincidente con la finalizaci&oacute;n de actividades de    dragado en una localidad cercana y con la disminuci&oacute;n de los niveles    de nutrientes. No obstante, esta especie no mostr&oacute; diferencias significativas    a trav&eacute;s de los a&ntilde;os ni correlaci&oacute;n con la variaci&oacute;n    de brillo solar. Una banda anual de baja densidad, m&aacute;s amplia que la    de alta densidad en <i>D. labyrinthiformis</i> y m&aacute;s angosta en <i>M. annularis</i>,    parece formarse aproximadamante durante abril y mayo en ambas especies, lo cual    es coincidente con el final de la &eacute;poca seca (incremento en la temperatura    del agua, disminuci&oacute;n de horas de brillo solar y mayor influencia del    Canal del Dique sobre las islas del Rosario). Las bandas anuales que se forman    en las dos especies en el sitio de estudio podr&iacute;an estar muy relacionadas    con la &eacute;poca de reproducci&oacute;n de cada especie. Investigaci&oacute;n    adicional permitir&aacute; un mejor entendimiento de c&oacute;mo algunas condiciones    locales o la fisiolog&iacute;a del coral est&aacute;n relacionadas con el crecimiento    coralino en las islas del Rosario.</p>     <p>PALABRAS CLAVE: Tasas de crecimiento, <i>Diploria labyrinthiformis</i>, <i>Montastraea annularis</i>, Mar Caribe, Colombia, Arrecifes coralinos</p>  <hr size="1">      <p>&nbsp;</p>     <p><font size="3"><b>INTRODUCTION</b></font></p>     <p>Rosario islands reefs are one of the most important coral ecosystems in Colombia.    This area was declared as a National Natural Park in 1978 to preserve the coral    community. However, the use of dynamite as a fishing-method, uncontrolled tourism,    launches and boats, increase in sea surface temperature and sediment discharge    due to dredging of Dique Channel resulted in the loss of coral coverage (Alvarado    <i>et al</i>., 1986). Bar&oacute;n <i>et al</i>., (1984) reported a sediment discharge amount    of 12,000 tons per day from Dique Channel since 1982, when dredging began. By    1986 a 90% mortality of <i>Acropora palmata</i> and <i><i>A. cervicornis</i></i>    was evident (Ram&iacute;rez 1986) and coral coverage decreased (Alvarado <i>et al</i>., 1986). This phenomenon was coincident with El Ni&ntilde;o event of 1982    - 1983, when mortality of corals at all Caribbean sites increased and death    of the sea urchin <i>Diadema antillarum</i> was reported. Recent studies suggest    a low retrieval of Rosario islands coral reef which could be related with dredging    detention in 1990 (Alvarado pers. observ., Sch&ouml;nwald unpublished). Sclerochronology    provides annual growth rates of specimens which can be used to understand relationships    between coral growth and environmental conditions (Dodge and Vaisnys 1980, Hudson    <i>et al</i>., 1989, Guzm&aacute;n <i>et al</i>., 1991). Knutson <i>et al</i>., (1972) demonstrated    the presence of alternating high (dark images on the X-radiography positive    print) and low (light images on the X-radiography positive print) density bands    which are formed annually. Dodge and Vaisnys (1977), investigated the ecological    effects of dredging on corals in Bermuda using growth-band analysis. Other environmental    factors have been suggested to control coral growth rates: available sunlight    (Buddemeier <i>et al</i>., 1974), water temperature (Dodge 1981, Dodge and Lang 1983,    Hubbard and Scaturo 1985), oil effects (Dodge <i>et al</i>., 1984, Guzm&aacute;n <i>et al</i>., 1991, Guzm&aacute;n and Jarvis 1996), suspended particulate matter (Tomascik    and Sander 1985) and sedimentation (Rogers 1990). However, the effect of sediments    and eutrophication over coral growth remain uncertain. Previous studies have    shown the negative effects of both dredging and sediments over coral skeletal    extension (Dodge and Vaisnys 1977, Bak 1978) and positive effects (Hudson 1981,    Carricart-Ganivet and Merino 2001), as well as contradictory effects of environmental    variables over corals from the same study sites (Dodge and Thomson 1974, Logan    and Tomascik 1991). Dodge and Brass (1984) found that density, calcification    and extension growth rates are complementary variables and when they are combined,    a better understanding between growth rates and the environment is obtained.    Consequently, recent studies suggest that massive corals growth rates may be    poor indicators of coral reef health where they are influenced by sedimentation    (Edinger <i>et al</i>., 2000) and indeed they can modulate their skeletal growth independently    of the amount of calcium carbonate available (Carricart-Ganivet and Merino 2001).    In this article we determine the extension growth rates of <i>Diploria labyrinthiformis</i>    (Linnaeus 1758) and <i>Montastraea annularis</i> (Ellis and Solander 1786) at    Isla Grande (north of Rosario islands), including the possible role of time    of reproduction over growth rate results. <i>D. labyrinthiformis</i> is a common    species at Isla Grande, but has been poorly studied in Colombia and elsewhere    in the Caribbean. <i>M. annularis</i> is a dominant species at the study area.    According to the description of Knowlton <i>et al</i>., (1992), this study was done    sensu stricto with morphotype I of the sibling species <i>Montastraea annularis</i>.</p>     <p>&nbsp;</p>     <p><b><font size="3">MATERIALS AND METHODS</font></b></p>     <p><b>Collecting site</b></p>     <p> Coral specimens of <i>D. labyrinthiformis</i> and <i>M. annularis</i> were    collected on May 1998, from the Isla Grande north reef at Rosario islands (<a href="#fig1">Figure    1</a>), Colombia (10&ordm;10&#8217;21&#8221; - 10&ordm;11&#8217;28&#8221;N and    75&ordm;42&#8217;36&#8221; - 73&ordm;43&#8217;52&#8221; W). Trade winds are    the most important macroclimatic factor in the zone. Two principal climatic    periods are known: dry season, between December and April, with high influence    of Trade winds and, wet (rainy) season, between May and November. Dique Channel    waters (an artificial arm of Magdalena River which crosses the country) pass    through Barbacoas Bay and discharge into Rosario islands during the wet season.    During the dry season the Dique Channel influences Rosario islands waters less    (Leble and Cuignon 1987, Alvarado and Corchuelo 1992).</p>          ]]></body>
<body><![CDATA[<p>    <center><a name="fig1"><img src="img/revistas/mar/v33n1/v33n1a11fig1.gif"></a></center></p>        <p><b>Collection and laboratory description</b></p>     <p> Fifteen colonies of each species between 20 &#8211; 50 cm in diameter were    obtained at depths of 6.0 &#8211; 11.0 m by SCUBA diving, using hammer and chisel.    Corals were sun exposed for one week, soaked in fresh water with 2% sodium hypochlorite    for 24 &#8211; 72 h, rinsed and dried with sun exposure. Five specimens of <i>M.    annularis</i> and four specimens of <i>D. labyrinthiformis</i> were cut using    a 24&#8221; diamond rock saw into 4 &#8211; 6 mm thick slabs through the axis    of maximum growth of each colony. The slabs were X-rayed with an Ergophos IV-SiemensTM    machine with an exposure ranging from 42 &#8211; 60 kv and 12 &#8211; 30 mas    over 0.12&#8211;0.30 s, and a film-target distance of ca. 30 cm. The negative    prints of the X-rays were developed on photographic paper to obtain positive    contact prints. One transect along the axis of maximum growth was analyzed by    measuring with vernier calipers (resolution 0.1 mm) the distance between each    couplet of light and dark bands which were defined as annual growth increments    (Knutson <i>et al</i>., 1972, Buddemeier <i>et al</i>., 1974, Dodge and Vaisnys    1980, Logan and Tomascik 1991). In cases where the banding was not clearly defined,    another transect near the axis of maximum growth was measured. </p>     <p><b>Environmental data</b></p>     <p> Hours of sunlight at Rosario islands were obtained from Instituto de Hidrolog&iacute;a,    Meteorolog&iacute;a y Estudios Ambientales (IDEAM) archives. Sea-surface temperature    was taken with the help of nighttime satellite observations by the National    Oceanic and Atmospheric Administration (NOAA). Nutrient and turbidity Information    was obtained from previous studies (Bar&oacute;n <i>et al</i>.,1984, Ram&iacute;rez    1986, Alvarado and Corchuelo 1992, among others).</p>     <p><b>Statistical analysis</b></p>     <p> To transform the mean and variance into more homogeneous data with respect    to time, standardization procedure of ring-width analysis was followed (Fritts    1976). The annual linear growth values of each coral were divided by its particular    mean growth rate, providing index chronologies with an approximate value of    1.0. The index values of all corals were averaged by year to form an index master    chronology (<a href="#fig2">Figure 2</a>). To smooth the year-to-year variability    (high frequency variation) a three-year moving average was applied to each chronology.    The index master chronology was correlated with sea-surface temperature and    hours of sunlight using the Spearman test. To quantify yearly variation in growth    a Repeated Measures ANOVA was used, with dredging (presence/non presence) and    time (years) as factors. This kind of analysis is required because successive    growth bands are measured in the same corals. The null hypothesis tested was    that there are no statistically discernible differences in the mean growth rates    between the years analyzed on each species, the index master chronologies of    both species, and between the years 1975 &#8211; 1981 (before dredging period),    1983 &#8211; 1989 (dredging period) and 1991 - 1997 (after dredging period).    With Levene and Bartlett tests (i.e. homogeneity of variance tests) and with    normal probability plots (i.e. normality test) the ANOVA assumptions were tested.</p>          <p>    <center><a name="fig2"><img src="img/revistas/mar/v33n1/v33n1a11fig2.gif"></a></center></p>        ]]></body>
<body><![CDATA[<p>&nbsp;</p>     <p><b><font size="3">RESULTS</font></b></p>     <p>The skeletal extension growth rate of <i>D. labyrinthiformis</i> was 3.8 mm    year <sup>-1</sup> (SE &plusmn; 0.10; N = 87) and 10.6 mm year <sup>-1</sup>    (SE &plusmn; 0.32; N = 56) for <i>M. annularis</i> at Isla Grande. The number    of annual growth bands ranged from 21 to 22 years for <i>D. labyrinthiformis</i>    and 6 to 14 years in <i>M. annularis</i>. There was not a clear pattern related    to growth variation of <i>D. labyrinthiformis</i> (<a href="#fig2">Figure 2A</a>),    in fact the annual growth shifted into an alternated variation along the years    studied (up and down 1.0). In contrast, <i>M. annularis</i> (<a href="#fig2">Figure    2B</a>) showed a clear coincidence to increase growth when last dredging activities    finished (1990-1991). In accordance with the time of collection and what is    shown in the apex of corals on the contact prints, low density (LD band) of    both species seems to be formed (or to have just formed) in April-May. The LD    band formation coincides with the increase of sea-surface temperature, the decrease    of hours of sunlight, and a higher influence of Dique Channel waters over Rosario    Islands. High density (HD) bands were generally narrower than LD band portions    in <i>D. labyrinthiformis</i>, while in <i>M. annularis</i> HD band were wider    than LD band portions. Samples X-radiograph is shown in <a href="#fig3">figure    3</a>.</p>          <p>    <center><a name="fig3"><img src="img/revistas/mar/v33n1/v33n1a11fig3.gif"></a></center></p>        <p> The Repeated Measures ANOVA of <a href="#tab1">table 1</a> indicates significant    differences in annual growth along 1975 - 1997 period in <i>D. labyrinthiformis</i>    (F = 2.39; P &lt; 0.005) and no significant differences in the annual growth    of the 1984 - 1997 period in <i>M. annularis</i>. Tukey HSD test shows that    recent years (1995, 1996) have significant differences in relation to the years    1977, 1983 and 1985 in <i>D. labyrinthiformis</i>. We found significant statistical    differences between growth variation of both species (Two-way Repeated Measures    ANOVA: F = 4.97; P &lt; 0.000) during 1984 - 1997 period (<a href="#fig4">Figure    4</a>). Repeated-measures ANOVA of <i>D. labyrinthiformis</i> growth rates were    not significant for before (1975 - 1981), during (1983 - 1989) and after (1991    - 1997) the last dredging (F = 3.1798; P &gt; 0.05: <a href="#tab1">Table 1</a>).    Using years as a factor, significant differences for the annual growth rate    during 1991 - 1997 period were found in this species, therefore another repeated-measures    ANOVA was done and it shows no significant differences before or during the    last dredging (excluding 1991 - 1997 period). Growth rates in <i>M. annularis</i>    during and after dredging were not statistically significant (F = 2.705; P &gt;    0.05: <a href="#tab1">Table 1</a>) in spite of the increase of growth after    last dredging activities. </p>          <p>    <center><a name="fig4"><img src="img/revistas/mar/v33n1/v33n1a11fig4.gif"></a></center></p>          <p>    <center><a name="tab1"><img src="img/revistas/mar/v33n1/v33n1a11tab1.gif"></a></center></p>        ]]></body>
<body><![CDATA[<p> Results of Spearman correlation test for annual growth rates of <i>D. labyrinthiformis</i>    (&plusmn; standard error) and <i>M. annularis</i> (&plusmn; standard error)    with sea-surface temperature (SSTs in &deg;C) and annual sunlight (hours &plusmn;    standard error) are included in <a href="#tab2">Table 2</a>. Master index of    both species are different in each test because the growth period analyzed with    sea surface temperature was 1984-1996 while with annual sunlight was 1984-1997.    Sea surface temperature at Isla Grande varied very little during 1984-1996 (average    range = 27.9 - 29.0 &deg;C). Indeed, sea surface temperature was not significantly    negatively correlated with growth of <i>D. labyrinthiformis</i> (<a href="#tab2">Table    2</a>). Contrastingly, growth of <i>D. labyrinthiformis</i> was significantly    negatively correlated with hours of sunlight (r = - 0.60; P &lt; 0.039). Annual    growth of <i>M. annularis</i> was positively but not significantly correlated    with both sea surface temperature and hours of sunlight (<a href="#tab2">Table    2</a>). </p>            <p>    <center><a name="tab2"><img src="img/revistas/mar/v33n1/v33n1a11tab2.gif"></a></center></p> 	     <p>&nbsp;</p>     <p><b><font size="3">DISCUSSION</font></b></p>     <p>In the present study, growth rates of <i>D. labyrinthiformis</i> and <i>M.    annularis</i> are slightly higher than those reported in previous studies at    high and low latitudes of the Caribbean (<a href="#tab3">Table 3</a>). Latitudinal    growth rate differences are controlled by reduction in winter water temperatures    and light levels with increasing latitude. Consequently, coral growth rates    are inversely related to latitude in the Atlantic (Logan and Tomascik 1991,    Logan <i>et al</i>., 1994). </p>            <p>    <center><a name="tab3"><img src="img/revistas/mar/v33n1/v33n1a11tab3.gif"></a></center></p> 	     <p> However, growth rates are also controlled by local environmental conditions.    <i>D. labyrinthiformis</i> growth rates at Isla Grande seems to be influenced    by changes in hours of sunlight, while a clear increment in skeletal extension    of <i>M. annularis</i> was seen when dredging activities finished (1990 - 1991).    Nevertheless, it was not statistically significant. It is possible that tolerance    to light and sediment level are the principal causes of the inverse results    showed between correlation of growth in both species with hours of sunlight.    In addition, this could be producing the significant differences in skeletal    extension of <i>D. labyrinthiformis</i> and <i>M. annularis</i> in the last    14 years. Symbiont species of hosts have diverse mechanisms of photo-acclimation    (Chang <i>et al</i>., 1983). Iglesias-Prieto and Trench (1994) showed that different    species of symbiotic dinoflagellates have several photosynthetic responses under    identical farming conditions. Leletkin and Zvalinsky (1981) provided evidence    that at a low light intensity the photosynthesis rate of zooxanthellae is greater    compared with those adapted to a normal light intensity. Increase of nutrients    may also have an effect on zooxanthellae. For example, Marubini and Davies (1996)    report an increase in areal density of dinoflagellates and a decrease in extension    growth in Porites porites and <i>M. annularis</i> due to nitrate enrichment.  </p>     <p> Sediment levels per se affect coral species in a different way. It is well    known that species differ in their capability to remove sediments which are    in contact with their colonies (e.g. Rogers 1990). <i>D. labyrinthiformis</i>    and D. strigosa have shown better capability to remove sediments than <i>M.    annularis</i> and M. cavernosa (Hubbard and Pocock 1972, Rogers 1990 and references    therein). Dredging activities at the Dique Channel started again in 1982 and    its influence at the Rosario islands was evident (Bar&oacute;n <i>et al</i>.,    1984). Alvarado and Corchuelo (1992), observed the influence of the Dique Channel    waters in the southern Rosario islands during wet season (May - November), while    during dry season they found a slight influence in the north. It is also important    to note that amounts of nutrients and turbidity increased between 1982 - 1990    at Isla Grande (Alvarado <i>et al</i>., 1986, Alvarado and Corchuelo 1992).    In previous studies (e.g. Dodge <i>et al</i>., 1974, Dodge and Vaisnys 1977,    Bak 1978, Dodge 1981, Cort&eacute;s and Risk 1985, Tomascik 1990), dredging    activities near coral reefs were found to produce a reduction in coral growth    and increase of coral mortality. Dredging activities near coral reefs increase    turbidity, reducing the available light for photosynthesis and increasing the    sediment load on corals, which have sub-lethal and lethal effects (Rogers 1990).    Notwithstanding, Tomascik and Sander (1985) explained that coral extension may    increase at sites with moderate increments of suspended particulate matter in    the water column, because they are a significant source of additional food for    the polyps. Dodge and Brass (1984) and Heiss (1996) determined that higher levels    of phosphate reduce calcification and growth, while other nutrients promote    coral extension. Hence, recent studies have discussed that growth rates are    poor indicators of reef health and suggest that high extension and low density    skeletal growth could be a result of eutrophic conditions (Edinger <i>et al</i>.,    2000). </p>     ]]></body>
<body><![CDATA[<p> We found that low density (LD) band formation of both species studied is formed    when Dique Channel waters have high incidence over Rosario Islands, as has been    suggested by some authors (Edinger <i>et al</i>., 2000, Carricart-Ganivet and    Merino 2001). Generally, the annual band formation has been related with environmental    changes, such as available light and water temperature (Highsmith 1979), salinity    (related with precipitation changes), and time of reproduction (Guzm&aacute;n    and Cort&eacute;s 1989). In this study, LD band deposition of both <i>D. labyrinthiformis</i>    and <i>M. annularis</i> seems to happen during April-May, during the transition    from dry to rainy season (highest water temperature and lowest time of sunlight).    Dodge and Thomson (1974) found that formation of LD band of <i>D. labyrinthiformis</i>    in Bermuda takes place in summer (at high water temperature), while Logan and    Tomascik (1991) found that LD band formation in the same species at the same    locality occurs during winter period (lowest sea water temperature). Previous    studies of <i>M. annularis</i> in the Caribbean have related HD band formation    with summer and autumn when sea water temperature increases and hours of sunlight    decrease (Macintyre and Smith 1974, Highsmith 1979, Hudson 1981). At Rosario    islands, sea surface temperature variation is insignificant between dry and    rainy period (27.6 &plusmn; 0.54 &deg;C and 28.6 &plusmn; 0.58 &deg;C, respectively),    this indicates that sea surface temperature does not have a significant effect    on seasonal growth rates of both studied species at Isla Grande. Guzm&aacute;n    and Cort&eacute;s (1989) also suggested that sea water temperature does not    influence coral growth rate in the eastern Pacific at Costa Rica (low latitude)    due to the small variation between dry and rainy period. </p>     <p> Wellington and Glynn (1983) suggested that the lower growth rate in Panama    corals could be related to the energy spent in reproduction. Guzm&aacute;n and    Cort&eacute;s (1989) found that in Porites lobata LD band was formed during    the dry season (higher hours of sunlight) and when corals were not reproducing.    At Isla Grande, gametogenic cycle of <i>D. labyrinthiformis</i> begins between    August-October and finishes during May-June, coinciding in general with dry    period and higher hours of sunlight. According to this, main expenditure of    energy (maturation of gonads) occurs during the dry season, while during May-June    this expenditure declines due to gamete liberation (Alvarado <i>et al</i>.,    in press.). In contrast, gametogenic cycle of <i>M. annularis</i> at Isla Grande    begins on April and finishes with gamete liberation during September - October    (S&aacute;nchez <i>et al</i>., 1999), coinciding with rainy season and less    hours of sunlight. In summary, the difference observed between annual bands    deposition of both species at Isla Grande might be highly related with time    of reproduction of each species inasmuch as LD band formation in <i>M. annularis</i>    coincides with the rainy season and the beginning of the gametogenic cycle using    a high expenditure of energy, while in <i>D. labyrinthiformis</i> coincides    with the dry season and the end of the cycle. Coral reproductive cycle is influenced    by the life strategy of each organism. In corals, and generally in any organism,    reduction of growth means a reduction in the future reproduction and the survival    (Hall and Hughes 1996). Corals do not stop growing in order to reproduce, they    decrease their growth rate during reproduction periods. It appears to be clear    in both studied species in which skeletal extension is wider before their reproductive    cycle begins. </p>     <p> This is the first sclerochronological study in the Colombian Caribbean, which    allowed retrospective skeletal extension growth analysis. It is difficult for    us not to refer to the diminishing of coral population at Isla Grande during    at least the two previous decades. At the first sight, some authors related    this significant occurrence with dredging activities, but indeed there were    other confounding environmental changes (i.e. El Ni&ntilde;o event, death of    the sea urchin Diadema antillarum, among others). We attempted to find any skeletal    coral growth rate change during the last two decades over the study area, even    though the direct incidence of the dredging activities was not the main purpose    of this research. However, we can not confirm any assumption due to the lack    of information such as environmental data, sub-annual bands formation or complementary    growth rates variables. Nowadays, it is well known that density, calcification    and extension growth rates are complementary variables and when they are studied    in concert a better understanding between growth rates and the environment is    obtained (Dodge and Brass 1984). Nevertheless, we report here LD band formation    during high incidence of Dique Channel waters over Rosario islands and an apparent    relation of density bands formation with time of reproduction of each species.    Additional investigation of these species is necessary to understand how coral    physiology or other local conditions are related with coral growth at Rosario    Islands. </p>     <p>&nbsp;</p>     <p>ACKNOWLEDGMENTS</p>     <p>This research was part of the project &#8220;Reproduction, growth and transplantation survivorship of the coral species <i>Montastraea annularis</i>, <i>Diploria labyrinthiformis</i> and Porites astreoides in the National Natural Park Corales del Rosario and San Bernardo, Colombian Caribbean&#8221;, grant to EM Alvarado, JA S&aacute;nchez and R Garc&iacute;a (Jorge Tadeo Lozano University and COLCIENCIAS 1202-09-221-96). We thank  R Garc&iacute;a for the contribution with the project. R Garc&iacute;a, OL Arenas, MF Gil, LH Chasqui, A Alfonso and M Morelos for the help in the field work. HM Guzm&aacute;n and RD Llin&aacute;s for useful advice and material support. RE Dodge (Nova Southeastern Oceanographic Center) for providing important articles. I Barraquer for X-raying. M Vargas and INGEOMINAS for cutting the corals. LM Moncada, OL Rodriguez and all laboratory personnel at Universidad de Bogot&aacute; Jorge Tadeo Lozano for their support in preliminary chemical tests. S Dell and A Vaughn for radiograph scanning. D Trujillo and A Gal&aacute;n for providing statistical programs. J Dillard and R Slusar for grammar review. Comments from JP Carricart-Ganivet and three anonymous reviewers notably improved the manuscript. Contribution No. 61 Museo del Mar, Centro de Investigaciones Cient&iacute;ficas, Universidad de Bogot&aacute; Jorge Tadeo Lozano.</p>     <p>&nbsp;</p>     <p><b><font size="3">REFERENCES</font></b></p>     <!-- ref --><p>Alvarado, E.M., F. Duque, L. Fl&oacute;rez-Gonzalez and R. Ram&iacute;rez.    1986. Evaluaci&oacute;n cualitativa de los arrecifes coralinos de las islas    del Rosario (Cartagena-Colombia). Bol. 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Bull. 21 (8): 376-381.&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;[&#160;<a href="javascript:void(0);" onclick="javascript: window.open('/scielo.php?script=sci_nlinks&ref=000105&pid=S0122-9761200400010001100045&lng=','','width=640,height=500,resizable=yes,scrollbars=1,menubar=yes,');">Links</a>&#160;]<!-- end-ref --><!-- ref --><p> Wellington, G.M. and P.W. Glynn. 1983. Environmental influences on skeletal    banding in eastern Pacific (Panama) corals. Coral Reefs. 1: 215-222.&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;[&#160;<a href="javascript:void(0);" onclick="javascript: window.open('/scielo.php?script=sci_nlinks&ref=000106&pid=S0122-9761200400010001100046&lng=','','width=640,height=500,resizable=yes,scrollbars=1,menubar=yes,');">Links</a>&#160;]<!-- end-ref --><p>&nbsp;</p>     <p align="center">DATE RECEIVED: 13/05/03 DATE ACCEPTED: 24/09/04</p>     <p>&nbsp;</p>     <p>ADDRESSES OF THE AUTHORS:    <br>   503 West End Avenue, Elizabeth, NJ 07202 USA E-mail: <a href="mailto:hcharry@yahoo.com">hcharry@yahoo.com</a>    (HC).    <br>   Centro de Investigaciones Cient&iacute;ficas (Museo del Mar), Universidad de    Bogot&aacute; Jorge Tadeo Lozano, calle 22 No. 3-30 piso 7, Bogot&aacute; D.C.,    Colombia. E-mail: <a href="mailto:elvira.alvarado@utadeo.edu.co">elvira.alvarado@utadeo.edu.co</a>    (EMA).    ]]></body>
<body><![CDATA[<br>   Departamento de Ciencias Biol&oacute;gicas, Universidad de los Andes, carrera    1E No 18 A &#8211;10 (J304A) PO Box 4976, Santaf&eacute; de Bogot&aacute; D.C.,    Colombia. E-mail: <a href="mailto:juansanc@uniandes.edu.co">juansanc@uniandes.edu.co</a>    (JAS).    <br> </p>  </font>      ]]></body><back>
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