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Sid

Percent of colonies with recent Injury

1606 colonies

4 K Rude and W. Starter, BioScim 20, 222 (1970).

5. C. Bertand. A A Reimer, J. R. Young. Survey of Marine Communities in Perem 30

a Esperances with Od (Publ. EPA-600/3-76-028, Environmental Protection

Agency Ecological Research Series, Narragansen, RI, 1976).
6. J. D. Cubu a d., in 1907 Oil Spill Conference (Publ. 452, Amencan Petroleum

Institute. Washington, DC, 1987), pp. 401-406.
677 colonica
Dip

7. J. Cubit and S. Williams, Atoll Ra. Bull. 269, 1 (1983)
20

& J. D. Cubit, RC. Thompson, H. M. Caffey, D. M. Windsor, Smithsonian Contrib.

Mer. Sci. 32. 1 (1988); LG Macintyre and P.W. Glynn, Am. Asser. Petrol Grol

Buell. 60. 1054 (1976).
(10%
Mortality

9. G L Hendlar, in Proceedings of the Third Intermahonial Coral Reef Symposium. D.L

Tavlor, Ed. (University of Miami, Miami, FL 1977), vol. I, pp. 217-223. ). D

Cubil. D. M. Windsor, R. C. Thompson, J. M. Burger, Externe Conial Shelf Se 10 1126 colonies

22, 719 (1986) Por

10. J. C Ogden and E. H. Gladichter, UNESCO Tech. Pap. Mar. Sci. 23, 1 (1983) Sid

> 10%

Important species in common throughout the tropical western Adanac are red Dip Por

Mortality

mangrove R. negle, sagrass T testudinum, most reef corals and algas, and

commercially important lobsters (Paulinus spp) and oysters (C. ruzophorae).

11. Ongoing drilling and refining of Yucatan and possible production along the west Nono Moderato

coast of Florida Amount of alling

12. The oil cope was 70% Venezuelan crade, 30% Mexican Isthmus crude, specific

pravny 27 (American Petroleum Institute). Percent compoution determined at the Fig. & Frequency of injury for the three most common species of massive Bermuda Biological Station by column chromatography with the use of alumina coraks in relation to the amount of oiling a 12 rects (4 unoiled, 3 lightly to and silica gads: 47.4% surates (fraction I, herane); 6.4% light aromatics moderately oned, and 5 heavily oiled). Open bars represent coral colonies (fraction 2. 10% ccher/90% horanc); 4.8% heavy aromatics (fraction 3, 20% with injuries that did not exceed 10% of the surface area of the coral; filled

methylene chloride/80% hecanc); 18.9% polar fraction (fraction 4. mcthylene

chlorid); and 22.5% werecovered. bars represent injuries greater than 10% (maximum observed, 100%).

13. CD Getter, G. I Scont, J. Michel, in 1981 Oil Spill Conference (Publ. 75-4161, Sample sizes are shown above cach category of oiling Abbreviations; Dip

American Petroleum Institute, Washington, DC. 1981). pp. 535-540. Duplonia diversaPor, Parites aspreides; Sid, Siderestre sideres, s, shallow (0 to 1

14. Refinery oficials animated that <21.000 liters of Corcut 9527 (Enxons Chemicals m), and d, deep (1 to 2 m).

Amenica) were spraved from aircraft (6). STRI personnel observed thus spraving

barween Acana Chaquta and Punta Galeta 15. J. D. Woodley a dl. Sommer 214, 749 (1981), ) H Connell, ithul 199, 1302

(1973) natural populations from laboratory-based physiological data a

16. Surveys beyond Punta Galera began the first week of June 1986 and lasted 1

monch Videotapes were made of the coast berween Rio Chagres and Islas small-scal, short-term press perturbation aperiments in the field Naranjos, and sell photographs were made all along the coast. Visual assessments (2, 3, 48, 52, 53).

were made of degree of oiling (heavy, moderate, light, or absent) and the habitats Third, sublethal effects are extensive and may be more important

and nypos of organisers obviously affected

17. J. D. Cubat, G. Batista de Yec, A. Roman, V. Banista. in Agonia de la Naturaleza, S. in the long term chan initial mortality (3, 48). Changes in stomato Hectandon Moreno and ). Espinou Gonzalez. Eds. (Impretes, Panama, 1985), pp. pod behavior and population structure were more pronounced than

183-199, km Medica Perane 9. 56 (1984).

18. Od was observed commonly uncil the ame of manuscnpt revision (July 1988) might be aapected from a simple reduction in numbers. Likewise,

around Isla Pavardi. where it emerged directly from the coral landhl on which the injury of corals has allowed colonization of their bare skeleton by chinery is buik, and Punua Muerto, Bahua Cativa, castern Isla Largo Remo, and algae and other sessile organisms that may overgrow parts of

Punta Galcea

19. Withun this area ad commonly escaped from sediments when we walked among colonies that survived the initial effects of oil (3, 54). Corals stressed

mangroves or cored into shallow subudal sagrass beds. Insertidal surfaces were by oil are probably also more susceptible to epidemic disease and heavily coated with oil

20. Most of the ou char escaped from Bahua Cativa was driven west. Coasts facing north likely to grow and reproduce more slowly than unaffected colonics

northeast were hcanhv oled, whereas adjacent areas facing west or south (3, 5, 52). Any combination of these effects may further reduce received back oil. The best studied cumple is at the northwestern side of Isla Largo overall abundance of corals as much as the initial spill (54). Such a

Ramo (Table 2).

21. These low rides occur annually at this are of vear (8,9). Sea levels during most of chain reaction could continue long afrer any petroleum hydrocar.

this came were as much as 10 cm below the 9-year average for thus period of bons are present in the environment or coral cissues, just as staghorn normally low sea level. coral continued to decline precipitously following a severe hurricane

22. Corals were collected by divers on days when on surface slicks were visible, placed

in solvent-rinsed aluminum foil, and frozen. Tissues were removed from skeletons in Jamaica (55)

with an air pick and homogenized with a tissue grinder. To minimize variations The response of organisms to an oil spill, or any other major due to water a carbonate inclusions, standard Lowrey protein determinations

vere used as one measure of issue mass. After solvent atraction, tocal lipid disturbance, will depend on the conditions in which they normally

weights were determined microgravimetrically Extracts were fractonated by live (15, 48, 56). Morcover, the suite of organisms able to survive dorpion chromatography. Hydrocarbons were analyzed with a capillary gas under conditions of chronic pollution, and their resistance to further chromatograph cqwpped with a flame ionization detector and 25-m SÉ 52 fused

silia columns stress, is typically different from that in similar unpolluted habicats 23. R H Groen Sampling Design and Statistical Methods for Environmental Biologists (48, 57). Much of Bahia las Minas has been subjected to human (Wikcy, New York, 1979). disturbance, beginning with decades of excavation, dredging, and

24. J. B. Ć. Jacuon, Bredl. Mer Sa. 23, (1973), K. L. Heck. Mar. Biol. 41.335 (1977)

R Vasquez-Montoya, An. Inst. Cien. Mar Lommel. Univ. Nar. Autor. Mex. 10,1 landfilling for the construction of the Panama Canal and the City of

(1983). Colon, drainage and spraying of mangroves for mosquito control 25. This cancimate was atrapolated from aerial photographs of 36 km of the 82-km

shoreline, of which 76 km contains mangroves or the photographed shoreline, construction of the refinery and a large cement plant on landfill, a 36% contained a band of dead mangroves major od spill in 1968, and unknown amounts of chronic oil 2. Sprousing success was measured for seedlings collected from an unoiled site, and pollution from the refinery and ships passing to and from the Canal

planted in one unoiled and two oted sites. Or 25 seedlings transplanted into each

site, 16 (64%) sprouted as the unoiled site and I (2%) sprouted at the oiled sites (4, 5, 17, 58). It is a measure of the severity of the 1986 oil spill that

27. J. P. Sucherland. Mer Bial 58.75 (1980); W. E. Odum, C. C. McIvor, T. J. Smith the biological consequences were so detectable despite this history HI, The Ecology of the Mangroves of South Florida: A Community Profile (Publ. 81/24,

U.S. Fish Wildlife Service, Washington, DC, 1982). of environmental abuse.

2 Roots were sampled three times before the spill in nven and channels (Seprember

o October 1981, January 1982, and June 1982) and twice in open habitats REFERENCES AND NOTES

(September to October 1981 and January 1982). Several areas of shore within each

habiat were chosen haphazardly. Fifteen to 25 interadal roots 220 cm long that 1. National Recarch Council, alate Selts, & Efras (Nacional Academy had not become firmly attached to the mud were lifted from the water, and cover of Pram. Wathington DC, 1985)

attached organisms was measured for each root by 100 random point counts after 2 RB Clark. Alles T. R. Soc. Lando Se B 297. 18S (1982); R S. Carncy, the oil spill was artimated visually before the spill. Afte the spill previously Am Sa 74, 298 (1906).

sampled moverine and channel habitats included both ouled and unosled sites but all 3. Y lon and B Rinkevich, Me Ecol. Prop. Sc. 3, 167 (1980).

open sitos vere ailed. Thus it was necessary to choose sites near Portobelo (Fig. 2),

6 JANUARY 1989

ARTICLES 43

Pp. 111-118

the nearest equivalent unoiled shoreline, for unoiled open habitat similar to oiled open habicar. One hundred roots were sampled in each oiled and unoiled habitat

after the spil. 29. A K OGower and J. W. Wacancy, Buall. Mar. Sci. 17, 175 (1967); I. B C Jackson, Mar. Bial: 14, 304 (1972);

D. K. Young and M. W. Young, in Ecology of Manne Benches, B. C Coull. Ed. (University of South Carolina Preams, Columbia 1977), pp. 359-382; I C Ogden, in Handbook of Seagrass Biology An Ecerystem Perspective, R. C. Phillips and C. P. McRoy, Eda (Garland, New York, 1980), PP

173-198, A. W. Sconcr, Bull. Mer Sai 30, 537 (1980) 30. All seagrass beds sampled be adjacent to mangrove shoreline and are 21 m deep:

unoiled beds meeting the criteria occur near Portobelo and Isla Grande (Fig. 2). Infauna were collected with 10.2 cm inside diameter polyvinyl chloride (PVC) tubes pushed into the sediment to a depth of at least 10 cm. Each sample consiss of three pooled cores (240 cm) At cach census cight samples were collected haphazardly at each site Samples were sieved over 500 un mesh and preserved in 10% formalin with rose bengal

. Animals were sorted to major con and counted or weighed wet. 31. M B. Bertness, Ecology 62, 751 (1981). 32. J. D. Cubit, in Proceedings of the Fifth international Coral Reef Congress, C. Gabrc, .L

Todart B. Salvat, Eds Antenne Muscun-Ephe, Moorca, French Polynesia

1985), vol. 3, 33. Percent cover of sessile species attached to the substratum was estimated by point

sampling along ten permanent transects 9 to 22 m long, perpendicular to the reef edge and spaced at random intervals within a 20-m strata. The number of points sampled per survey was between 1510 and 1560. The servad 6 m of thek transects were more directly exposed to ou than the landward portions. This 6-m band forms the "heavily oued zone" of Fig. 6. Se urchins were counted in rwo

1 x 20 m transects, one at the reef edge, the other 26 m farther landward. 3. For the reef edge transect ontv, residuals from regressions of changes in abundance

from Mav to June were hughh negaove in 1986 compared with all other transects

in all other vears (-41.0.88, P < 0.001) 35. RL Caldwell and H. Dingle, Sa. Am 234, 80 (January 1976); R. Steger, Ecology

60. 1520 (1987) 36 ML Reaka, in Proceedings of the Fifth Iniemational Coral Reef Congren, M. Harmchan

Vivien and B. Salvat, Eds. Antenne Museum-Ephe, Moorra, French Polynesia, 1985), vol. 5, pp. 139-14 RL Caldwell G. Rodenck, S. Shuster, lul Zool

Union Sfono, in press; R L Caldwell, Bull Mer. Sa 41, 135 (1987) 37. The four collection sites ranged from 100 to 125m x 25 to 40 m (all larger than

3000 m) and were selected before the spull to have interodal Thalassia bods with form, sanch substratum. and moderate amounts of rubble and coralline algal nodules. Densioes of stomatopods were estimated from 30 quadrats (0.5 m) chosen a random within 400 m plos a cach site. AB hard substrata were removed from each quadrat and broken, and all stomatopods were retained. A minimum of 300 individuals was collected, including ones from pieces of coral rubble collected haphazardly within cach site, but outside the plor used for

quadrats. These animals were included in the analysis of injuna and growth. 38 Cuocular imunes were recorded for all anunais as descnbed by LK Berzins and R

L Caldwell (Mer Beher Physiol. 10. 83 (1983) 39 Wc estimated growth bv companng the length of the carapace of animals newly

molted in the laborator with its length before molting, using only animals that

moired within 3 days of caprure and were longer than 35 mm 40. Almost every piece of rubble in the heart oued ploos was covered by hundreds of

hermit crabs. Such densides were nor observed in unoued plou or in any years

previoush 41. See P W Glumn (Bull Biol Sor Wash 2.13 (1972)) and J. W. Porter (bud., p. 89)

for general descnpoons of Panamanian reets. We selected fringing reefs that entended at least 100 m along the shoreline and to a depth of at least 12 m. Ar cach reet four or five une transects were extended from haphazardly chosen points at the shoreward edge of the reet to the deepest point of the reef Along cach line, 1 m quadrats were placed contiguously or up to 3 m apart, depending on transcat length 18 to 81 quadrats per transect Quadrats were divided into 100 cm celk,

and total cover was ourated by me within cach cell 12. We determined differences in coral cover before and after the spell (ful to October

1985 and July to August 1946) by combining a
(0 m2, 3 to sad 6 to 12 m; prepi.
respectively, poutapil - 29, 200, and 1871 et
(mean cover before the pill cover sterowd). The
by one-way ay of variance of the value for one hem
anled, and four unoiled rocks, - 2 For Own depe F-

Dom F-444; P - 0.13, 6 - 12 F-990, P-0.49. 12. Dead areas were bare a colonised by thin powd of Game

colonia of S. more are realed in plasac bag undene, brown laboratory, and placed in clan tantos and art. A fendal speared and wurface of each and when the flamentou alge powing over the end

areas vere quecued by hand 4. A. Anton. In Proceedings of the Fourth American Comel Rey Sy

Gomez ad., Ede (Marine Sciences Center, Universary of the Phulppee Queen

Ciry, Philippina, 1981), vol. 2 pp. 3-14, d., vol 27-14 45. Recent injurias vert counted on 12 rede for all on 2x 50

haphazardly located on I mand 1 l m depech Ljuries were when the appeared the word bere are only lightly apo te me

algae. Sie al injury was really imated percent of the tonal 46. Significance was tested with hierarchical log hones modes (SPSSIC

the following variables: coral pesies (chrex leves), water depeda amount of injury (three level), and amount of osling (see bed por disebuted equally a fraction moun odling for any of the there pece (P < 0.001); injuries increased in frequency and every culing increased amongst the feet of alling on coral munta sa independent of depoh (P < 0.001), injuries or more frequar = shoe depehs. The three coral species difered in frequency of muna (P < 0.001) ve See > Parisererede > Diplarie diron. La April 1977, sbour lates all spillcoral injuria vere saill nor dameributed equally a faction of

oiling among sites (P < 0.001). 17. A J. Southward and E C. Southward, J. Fisk Ra. Board Com 36.60 (17. 48. A) Southward, Philes. Tres R. Sa London Se B 291, 241 (192) 49. RJ. Nadeau and E T. Bergquin in 1971 o Spl Co Ni

Amenican Petroleum Lacine, Washington, DC, 1977). pp AS-544 L I Can hd.pp. 539-542, 1. E. G. Cincron and Y. Schacha Novela UNESCO TOP

Mu Sa. 23, 87 (1982). 50. J. H. Vandermeulen, Philos. Trans. R. Sa Lada Se B 397, 135 (1980) 51. A H. Knapad., O Petreden. Pol. I, 157 (1983), R E Dodged.Com

3, 191 (1984); A H. Knap, Muer Pollen. Bl. 18. 119 (19) 52. B. E. Brown and L S. Howard, Adv. Mar Bel. 22, 1 (1985) 53. P. Yodzi, Ecology 67, SOB (1988). 54. S. R. Palumbi and J. B. C Jackson, J. Exp. Me Heal Evol. 66, 103 (1982). T?

Hughes and J. C Jackson Exol. Monogr SS, 141 (1985). 55. N. Knowtoon. J.C. Lang MC Rooney, P. Cliford, Nont 29. 231 (1981) N

Knowlton, J. C Lang, & D. Keller, Similiamuan Cant. Me sa 56. J.H. Connell, in Population Dyna, R M. Anderson B D Turner. LR Tete

Eds. (Blackwell Scientific, Oxford 1979), pp. 141-163, AT Pxed SA Levin, Ecal Manag. 51, 145 (1981); STA Pident and PS Waste The

of Natural Distro Price Demo (Academic Pron Orlando, FL 1943 57. R P. M. Bak, MaPolies 16,534 (1987) 58. H. H. Rousseau, in The Pen Canada anomal by Care (Nea San

Francisco, 1915), pp. 374432; W. G Comber, a The Code Enginering Treatise, & W. Goedhals, Ed (McGraw-HI New York 1916 459-493, D. P. Curry, Am J. Trop. Med. S. 1 (1925), 1 G Clayton the Pena Canal (A. Kroch, Chicago, 1931), E C Webwer, La Dar

(Editorial Universitaria Panama Panama, 1973) 59. J. Brawn and R. Green helped with anda, and G KOK D. ME

Mon provided technical assistance. B. Carey D Dedge, R Green I kn Knowlton, Y. Loys and A. Smith reviewed the man and helped mam other ways. This work was supported by the US Mens Meme Son under contraa 14-12-0001-30393 and by the Smuchek Trs kan Institute and the Southsonian Intubon Éavion Soccer

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