BULLETIN OF MARINE SCIENCE, 69(1): 267–278, 2001
COMBINED EFFECTS OF THREE SEQUENTIAL STORMS ON THE HUATULCO CORAL REEF TRACT, MEXICO
Diego Lirman, Peter W. Glynn, Andrew C. Baker and Gerardo E. Leyte Morales
ABSTRACT Reefs of the Huatulco area, southern Mexico, were exposed to an unprecedented se- quence of three major storms (Olaf, Pauline, and Rick) over a 2-mo period (September– November 1997). The prior establishment of monitoring transects, as well as the timing of our surveys just 1 mo after the passage of Hurricane Rick, provided an unique oppor- tunity to document storm impacts on these recently described reef communities of the eastern Pacific. Considering the lack of prior hurricane damage to these reefs, the domi- nance of branching pocilloporid corals, and the intensity and high frequency of the 1997 storms that affected the area, it was hypothesized that storm-generated damage patterns would be significant and consistent among the reefs of Huatulco. However, the damage patterns documented were limited in severity and variable in spatial distribution. Of the six reefs surveyed prior to the first storm in July–August 1997, only three showed signifi- cant decreases in live coral cover, whereas the remaining reefs showed slight, non-sig- nificant increases in coral cover between surveys. The most common type of damage observed was the fragmentation of Pocillopora spp. colonies; at some locations, the den- sity of surviving Pocillopora spp. fragments exceeded 20 m–2. Fragmentation of the mas- sive coral Pavona gigantea (Verrill) was observed only at a single site. At several sites, large sections of reef framework (up to 245 cm in diameter) still exhibiting live Pocillopora spp. colonies on their upper surfaces were detached and transported away from their original locations. Even if the immediate damage observed was significantly less than predicted in light of the physical characteristics of the storms, the long-term effects of these storms will depend on the survivorship of detached colonies and fragments, the regeneration of damaged colonies, and the future impacts of bioerosion.
The negative, and often catastrophic effects of El Niño–Southern Oscillation (ENSO) disturbances on the coral reefs of the world have been well documented. As a result of the 1982–1983 ENSO, there was significant coral mortality throughout the equatorial east- ern Pacific (Glynn, 1984, 1990, 2000). Similarly, the 1997–1998 ENSO, which was ranked as the strongest event in recorded history (Enfield, this issue), resulted in bleaching and coral mortality at local and regional levels across the tropical oceans (Wilkinson, 1998; Goreau et al., 2000). Although the main mechanisms of coral stress and mortality during ENSO events are elevated sea surface temperatures (SST) and changes in irradiance lev- els (Glynn and D’Croz, 1990; Gleason and Wellington, 1993; Brown, 1997), other ENSO– associated stressors such as storm damage, elevated nutrients, sedimentation, aerial ex- posure, and algal blooms are also known to cause coral mortality (Glynn, 2000). In this study, we document the effects of storm-generated physical disturbance on the reefs of Huatulco, Mexico. Altered storm paths during the 1997–1998 ENSO exposed the reefs of this area to an unprecedented sequence of physical disturbances where three major storms (Olaf, Pauline, and Rick) made landfall on the southern Mexican coast over a 2-mo period (September–November 1997). The prior establishment of transects on sev- eral of the affected reefs, as well as the timing of our surveys just one month after the passage of Hurricane Rick, provided an unique opportunity to document storm impacts
267 268 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 1, 2001
Figure 1. Tracks of the three storms that impacted the Huatulco area in the latter part of 1997. Tropical Storm Olaf (September 26–October 12; maximum sustained winds = 110 kph; maximum wind gusts = 140 kph), Hurricane Pauline (5–10 October; 215 kph; 260 kph), and Hurricane Rick (7–10 November; 140 kph; 170 kph).
on these recently described reef communities of the eastern Pacific (Glynn and Leyte Morales, 1997). The effects of major storms on live coral cover, species diversity, and reef morphology are well documented (e.g., Stoddart, 1963; Glynn et al., 1964; Woodley et al., 1981; Done, 1992; Witman, 1992; Harmelin-Vivien, 1994). The physical characteristics of each storm as well as the timing and sequence of disturbances are key factors in describing damage patterns (Lirman and Fong, 1997a). The disturbance history of an area has also been shown to play a major role in determining damage and recovery patterns (Hughes, 1989). In general, reefs that have not been exposed to major physical disturbances for prolonged periods are believed to be more susceptible to storms as reef complexity and the abun- dance of less robust, fast-growing coral species with high susceptibility to physical dam- age proliferate (Porter et al., 1981; Rogers, 1993; Aronson and Precht, 1995). For ex- ample, the devastating effects of Hurricane Allen on Jamaican reefs in August 1980 have been attributed to the high abundance of fragile branching species such as Acropora cervicornis (Lamarck) that developed during a 36-yr period without major storm impacts in that area (Woodley et al., 1981; Woodley, 1992). Similarly, a sequence of strong storms over a short time period can have compounding effects on reef communities. In Florida, three major storms impacted reefs over a 2-yr period, and damage patterns documented after Hurricane Andrew (August 1992), were exacerbated during two subsequent storms (Lirman and Fong, 1997a). In this case, the additional damage was attributed in part to the remobilization of coral fragments during subsequent storms (Lirman and Fong, 1997a,b). LIRMAN ET AL.: HUATULCO REEF STORM DAMAGE 269
METHODS
The 1997 tropical storm season in the eastern Pacific was very active, with a total of 17 named tropical storms and hurricanes. More notable for the region, however, was the unprecedented se- quence of major storms that made landfall along the coast of Mexico in the latter part of the season (Fig. 1). Hurricane Nora (16–25 September; maximum sustained winds = 205 kph; maximum wind gusts = 260 kph), which affected the Baja California region, and Tropical Storm Olaf (26 Septem- ber–12 October; 110 kph; 140 kph), Hurricane Pauline (5–10 October; 215 kph; 260 kph), and Hurricane Rick (7–10 November; 140 kph; 170 kph), which affected the southern Mexican states of Oaxaca and Guerrero, resulted in severe flooding and coastal destruction [Fig. 1; Mexican Na- tional Weather Service (http://smn.can.gob.mx); U.S. National Hurricane Center (http:// nhc.noaa.gov)]. Land observations during the storms indicated variable wind and swell direction for each storm (Leyte Morales, pers. obs.). Swells from Tropical Storm Pauline and Hurricane Olaf impacted the Huatulco coastline from a SE direction. In contrast, swells from Hurricane Rick origi- nated from a NW direction. The reefs of the Huatulco area were described in detail previously by Glynn and Leyte Morales (1997). To determine the severity of hurricane damage, reefs in the Huatulco area were surveyed on 12–22 December 1997, only 1 mo after Hurricane Rick (Fig. 2). Changes in percent cover of live coral were evaluated on six reefs (Cacaluta, Chachacual, Entrega, Dos Hermanas, San Agustín, Montosa) that were initially surveyed in July–August 1997, 2 mo prior to the landfall of Tropical Storm Olaf. Survey transects were established perpendicular to the main axis of each reef along a depth contour (from the upper to the bottom edge of each reef), and 1 m2-quadrats were flipped on-
Figure 2. Map of the Huatulco area with the location of reefs surveyed on December 1997. 270 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 1, 2001 end along the transects to estimate live coral cover. Quadrats were divided into 100 subdivisions and the number of subdivisions occupied by live coral cover was visually estimated. The number of quadrats surveyed along each belt varied according to the width of each reef. Data from all quadrats were combined for each reef. In addition, the abundance of coral fragments and detached coral colonies were assessed on six reefs where such damage patterns were observed (Riscalillo, Mixteca, Órgano, Rincón Sabroso, Montosa, Maguey). The abundance of live Pocillopora fragments was quantified within transects placed along the depth contour of these reefs. Furthermore, to quantify the transport of detached colonies and fragments, transects were surveyed along the bottom edge on sandy substrata on three reefs (Órgano, Rincón Sabroso, Maguey). The abundance of live Pocillopora colonies and frag- ments were quantified within 1 m2 quadrats along these transects. Although storm damage was most evident on the branching pocilloporid corals that predominate on the reefs sampled, damage to the massive coral Pavona gigantea was noted on Riscalillo reef. Here, surveys were carried out to quantify the abundance and size of attached and detached P. gigantea colonies, as well as the percent live and dead tissue on both attached and dislodged colo- nies. The abundance of P. gigantea was quantified within three parallel transects located along the depth contour of Riscalillo Reef, as well as within a transect located along the bottom edge of the reef on sandy substratum. The maximum diameter of each P. gigantea colony was measured with a flexible tape and the percent live and dead tissue area were visually estimated. The data were examined for conformity to the assumptions of each statistical test prior to analy- sis. Normality was tested with the Shapiro-Wilk test and homoscedasticity was tested with the Bartlett’s test. When violations of the normality assumptions were encountered for percent cover estimates, and arcsine transformations (arcsine ÷x) were not successful, non-parametric statistical methods were employed for comparisons instead (Sokal and Rohlf, 1981).
RESULTS
The most abundant coral species on the reefs of Huatulco were Pocillopora verrucosa (Ellis & Solander), Pocillopora capitata (Verrill), and Pocillopora damicornis (Linnaeus). Live Pocillopora spp. fragments were abundant on storm-damaged Huatulco reefs (Table 1, Fig. 3). Our surveys showed that Pocillopora spp. fragments were both retained within the reef framework as well as deposited along the deep edges of reefs. Three reefs, Riscalillo, Mixteca, and Órgano, had the highest abundance of coral fragments. Mean abundance of fragments was not related to coral cover on these reefs (linear least-square regression, P > 0.1).
Table 1. Mean density (# m-2) of live Pocillopora fragments on Huatulco reefs that experienced storm damage in December 1997. Surveys were conducted on the reef surface and at the reef base on sandy substrata. n = number of 1-m2 quadrats surveyed. Rneef L)ocatio Mmean density (SEM Mnaximu Rfiscalillo R)ee 266.2 (4.9 132 4 Mfixteca R)ee 253.3 (5.3 922 Óorfgan R)ee 230.9 (3.3 531 Rf. Sabroso R)ee 96.9 (2.8 28 Mfontosa R)ee 73.5 (1.3 527 Mfaguey R)ee 42.8 (2.2 201 Óoregan B)as 210.3 (2.4 502 Re. Sabroso B)as 194.9 (1.6 202 Meaguey B)as 183.9 (1.6 252 LIRMAN ET AL.: HUATULCO REEF STORM DAMAGE 271
Figure 3. Photograph of a section of Montosa reef showing numerous live and dead fragments of Pocillopora spp. over live Pocillopora spp. colonies. Many of the live fragments were already fused to the underlying live tissue when this photograph was taken on 11 December 1997. Depth = 6 m.
In many cases, entire live Pocillopora colonies or large pieces of reef framework were broken, overturned, or transported away from the reef (Fig. 4). On Mixteca reef, a total of 35 detached live Pocillopora colonies were encountered within a 50 m2-belt surveyed along the reef base. Mean diameter of these detached colonies was 63.7 cm (SEM = 6.9) with a maximum diameter of 245 cm. Large, detached pieces of reef framework with live Pocillopora spp. colonies growing on their upper surfaces were also seen at Montosa, Chachacual, Cacaluta, La Entrega, Riscalillo, and Mixteca. At Mixteca reef, two large cavities where large sections (<3 m in diameter) of reef framework were removed were observed. Another mechanism for damage was the transport of rocks from the steep is- land shores down and across the reef face, resulting in coral fragmentation and abrasion. Such damage was most evident at Montosa reef where recently deposited boulders and coral debris were observed at the reef base. The massive coral P. gigantea experienced significant damage on Riscalillo reef. Out of a total of 158 colonies encountered, 22% of these colonies (n = 35) were detached and either remained in situ or were transported to the bottom of the reef. Detached P. gigantea colonies were significantly smaller (mean size = 24.7 cm, SEM = 3.5) than those colonies that remained attached to the reef (mean size = 48.2 cm, 2.9; t test, P < 0.01). Tissue mortality on these detached colonies was high at 62.4%, 5.1, but the proportion of tissue loss was not related to colony size (linear least-square regression, P > 0.05). 272 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 1, 2001
Figure 4. Photograph of La Entrega reef (12 December 1997) of a large section of reef framework dislodged during one of the major storms to impact the Huatulco area. Depth = 8 m.
Comparisons of live coral cover estimates measured before and after the storms on six reefs provided varied results (Fig. 5). Significant decreases in mean percent live coral cover were detected on three reefs, Cacaluta, Chachacual, and La Entrega (Mann-Whitney U tests, P < 0.05). In contrast, no significant increases in percent coral cover were docu- mented at Dos Hermanas, San Agustín, and Montosa (P > 0.05). To test the hypothesis that the orientation of each reef with respect to incoming waves and surge would influence the extent of hurricane damage, reef orientation and damage estimates were combined in a single plot (Fig. 6). This approach revealed no discernable patterns or distinct groupings of damage extent based on reef orientation. LIRMAN ET AL.: HUATULCO REEF STORM DAMAGE 273
Figure 5. Mean percent cover of live corals (± SEM) on Huatulco reefs before (white bars, July– August 1997) and after (stripped bars, December 1997) the last of three significant storms to affect the area. Numbers on top of the bars represent significance values for Mann-Whitney U tests. Numbers within the columns are sample sizes (number of 1 m2 quadrats) for each reef.
DISCUSSION
Considering the lack of recent hurricane damage to the reefs of Huatulco (Glynn and Leyte Morales, 1997), the dominance of branching corals on most reefs, and the intensity and high frequency of the three storms that affected the area (Glynn et al., 1998), it was hypothesized prior to this study that storm-generated damage patterns would be signifi- cant and consistent among the reefs of Huatulco. However, results from our surveys indi- cated that damage patterns were very patchy in severity and spatial distribution. Pocillopora, the most abundant coral genus on the reefs of Huatulco, was the most susceptible to storm damage as evidenced by the high abundance of broken branches and dislodged colonies on many of the reefs surveyed. The high susceptibility of branching morphotypes to storm damage is well documented (Woodley et al., 1981; Lirman and Fong, 1997a,b). The high susceptibility of Pocillopora spp. to fragmentation during storms was also documented at Punta Arenas in the Gulf of California, Mexico (González-Peláez et al., 2001). At this location, Hurricane Isis (2 September 1998; maximum sustained winds = 120 kph; maximum wind gusts = 150 kph) caused significant fragmentation and dislodgment of Pocillopora spp. colonies and tissue losses exceeding 59%. At Huatulco, live coral fragments were retained on the reef surface, where many fused to the underlying live coral colonies in a matter of weeks. Additionally, many surviving fragments accumulated on the sandy substrate at the base of reefs. The survivorship of coral fragments after detachment has been directly linked to substrate type (Yap and Gomez, 1984, 1985; Heyward and Collins, 1985). For example, fragments of the branching coral Acropora palmata exhibited high survivorship within the reef-flat and back reef areas in 274 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 1, 2001
Figure 6. Diagram of reef orientation and hurricane-related damage to reefs of the Huatulco area surveyed in December 1997. Orientation is defined as the perpendicular direction from the main longitudinal axis of the reef. SAB = Rincón Sabroso, MON = Montosa, MAG = Maguey, ENT = La Entrega, AGU = San Agustín, TEJ = Tejoncito, DAR = Dársena, MIX = Mixteca, RISC = Riscalillo, HER = Dos Hermanas, ORG = Órgano, CHA = Chachacual. Reefs within ovals were surveyed prior to the passage of the storms in July–August 1997.
Florida where they fused to live and dead A. palmata colonies. In direct contrast, those fragments that were transported away from the reef and onto sand, exhibited high mortal- ity patterns associated with sedimentation (Lirman, 2000). Accordingly, the retention of Pocillopora spp. fragments within the reef structure at Huatulco will be key to their long- term survivorship. Of the six reefs surveyed in July–August 1997, only three showed significant decreases in coral cover attributable to the passage of the storms (Fig. 5). The remaining reefs showed slight, non-significant increases in coral cover between surveys. Moreover, contrasting results were found even within the same reef. At Montosa, while a high density of –2 Pocillopora fragments were observed (mean = 7.5 fragments m , maximum = 53 frag- ments m–2) in a clear demonstration of storm damage, live coral cover increased from 60% to over 75% (Fig. 5). Several potential explanations exist for the observed variable patterns: (1) The physical damage caused by major storms on coral reefs is notoriously patchy at spatial scales ranging from regions to reef zones (e.g., Hubbard et al., 1991; Rogers, 1992; Bythell et al., 1993). Therefore, it is possible that our survey methods and sample sizes were insufficient to fully characterize damage patterns within reefs. (2) Be- cause of the slender, branching morphology and rapid regeneration and growth rates of Pocillopora spp., it is possible that significant fragmentation may have taken place with- out a major loss in coral cover. Thus, if mainly branch-tips were removed by the storms and the damaged colonies healed rapidly, the quadrat method used would still yield high LIRMAN ET AL.: HUATULCO REEF STORM DAMAGE 275
Figure 7. Photograph of a sea urchin (Diadema mexicanum) population grazing on the exposed reef framework at Dársena reef (17 December 1997). Depth = 6 m. estimates of coral cover even if significant coral biomass may have been removed. (3) Although the reefs of Huatulco are not normally exposed to hurricane or tropical storms during non-ENSO years, they are commonly exposed to high wave impact seasonally. Furthermore, the location of most reefs within small bays with rocky headlands may expose them to periodic strong wave action as the incoming swells are reflected by the steep shoreline. These chronic, seasonal disturbances may interact to remove the most susceptible coral colonies and branches, resulting in reef communities resistant to storm- generated physical disturbance. Previous studies have shown that reefs exposed to prior disturbances can be more resistant to subsequent storm impacts (Harriot and Fisk, 1986; Witman, 1992; Lirman and Fong, 1997a). Furthermore, this same physical setting may be responsible for the lack of correlation found here between reef orientation and the degree of storm-related damage (Fig. 6). The rocky shoreline and the development of reefs within small bays may have protected some reefs from direct impacts while exposing others to increased wave activity. Storm damage was not limited to the fragmentation of live coral colonies. At the most severely impacted sites such as Montosa, Chachacual, Cacaluta, Riscalillo, and Mixteca, large sections of reef framework still exhibiting live Pocillopora spp. colonies on their upper surfaces were dislodged and transported away from their original locations. Similar storm-generated damage to large pocilloporid blocks has been documented for the Gulfs of Chiriquí and Panama, Panama (Glynn et al., 1972; Glynn and Macintyre, 1977). In addition to the immediate reduction in reef structure and live coral cover, the exposure of the reef framework to bioeroders may compound the initial losses. Previous studies have shown that increases in abundance and/or redistribution of bioeroders following ENSO- associated coral mortality can intensify natural levels of bioerosion and result in the loss 276 BULLETIN OF MARINE SCIENCE, VOL. 69, NO. 1, 2001
of reef framework (Glynn, 1988; Glynn and Colgan, 1992; Eakin 1996). The intense activities of bioeroders are already evident in the Huatulco area at Dársena reef where a large population of Diadema mexicanum (Agassiz) was observed (Fig. 7). Live coral cover on this reef was significantly reduced due to the sedimentation produced by dredg- ing and port-building activities, and the grazing activities of the sea urchins have effec- tively flattened the exposed reef framework since then. A limited number of studies have reported storm disturbance on Pacific reefs during the 1982–1983 ENSO event. Laboute (1985) and Harmelin-Vivien and Laboute (1986) reported significant damage caused by cyclones in 1982–1983 at French Polynesia. Simi- larly, Robinson (1985) described damage patterns to branching and massive corals in the Galápagos Islands in 1982–1983 associated with increased swell size and reversals of swell direction. The present study is the first to document storm effects associated with the 1997–1998 ENSO event on the recently described coral reefs of the Huatulco region. Damage on these reefs was manifested as significant losses of live coral cover at three sites, widespread fragmentation and dislodgment of colonies of the branching genus Pocillopora and the massive coral P. gigantea, and the breakage and transport of large sections of reef framework. However, considering the magnitude and the short interval among the three storms, the immediate damage observed was significantly less than pre- dicted in light of the physical characteristics of the storms. Nevertheless, the long-term effects of these storms will depend on the survivorship of detached colonies, the regen- eration of damaged colonies, and the future impacts of bioerosion.
ACKNOWLEDGMENTS
Support for this study was received from the Universidad del Mar and the National Geographic Society (grant # 5969-97). Partial support for publication of this research was provided by NOAA’s COP grant NA67RJO149.
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