Rev. BioI. Trop., 46 Sup!. 5: 145-156, 1998

Epizoism: a new threat to health in reefs

Amfried Antonius 1 and Enric Ballesteros2 1 Ins1. Paleontology, Univ. Vienna, Althanstrasse 14, A-1090 Vienna, AuStria. 2 CentreEstudis Avanyats-CSIC, C. Santa Bm-bara sin, E-17300 Blanes, Spain.

(Rec. 25-VIl-I997. Rev. 26-1II-1998. Acep. 5-V-1998)

Abstract: Coral reefs around Carrie Bow , Belize, Central America were investigated in 1972 and reefs of the John Pennekamp Coral ReefState Park offKey Largo, Florida in 1973. Both sites were resurveyed in 1997 and anew threat to coral health was found: a rather epizootic occurrence of other organisms overgrowing living coral. On the -flat of the Barrier Reef north of Carrie Bow Cay, there are fields of cervicornis and Poriles poriles, scattered over shallow sand-bottom. A large number of these show either "tufts" of blue-green algae, or fila­ mentous "flags". Both may cover substantial portions ofindividualcoralla; the living tissue disappeared at the ofcon­ tact area. Another type ofovergrowth on living corals is represented by the red alga Melapeyssonnelia cora/lepida, as well as the brown alga Lobophora variegala. Both overgrow mainly Mi/lepora complanala and M. alcicornis, but also other corals with smooth surface, such as Poriles poriles and P aSlreoides. This syndrome occurs about equally abundant around Carrie Bow Cay, Belize, and in reefs offKey Largo, Florida. A similar kind oftightly attached over­ growth is represented by the sponge Cliona caribbea. It was found regularly in the reefs around Carrie Bow Cay on a wide variety of coral species. An identical type ofepizoism is represented by Chondri/la cf nucula, which, so far, was observed only in Belize. All these new syndromes together amount to a considerable coral-degrading factor, which was not found in these reefs 25 years ago.

Key words: Reefcorals, space competitors, epizoism, coral death, reef degradation.

Coral reefs around Carrie Bow Cay, Belize, When both reef-sites were resurveyed III Central America, located at 16° 48' N, 88° 05' early 1997, the most striking differences in W (Fig. 1) were investigated in 1972, and reefs comparison to the old surveys were that : 1) of the John Pennekamp State Park much larger portions of reef-crest and back­ off Key Largo, Florida, located at 25° 07' N, reef areas are dead today, and 2) a qualitatively 80° 18' W (Fig.2) in 1973 (Antonius 1974). new threat to coral health has emerged, consist­ Sinc~ 1972 was the year of discovery of the ing of a rather epizootic occurrence of certain black-band disease (BBD) of reef corals Cyanophyta, Rhodophyta, Phaeophyta, and (Antoni~ 1973), special emphasis in both sur­ Porifera overgrowing living corals. Thus, sub­ veys was put on coral health. Thus, along cer­ ject ofthis paper is a qualitative description of tain transects, live, diseased, otherwise impact­ newly observed phenomena of epizoism ed, and dead corals were recorded. Impacts on degrading coral health. live, corals other than diseases was found to be This does not mean that the "old" maladies mainly feeding damage by the fireworm have disappeared. On the contrary, BBD and carunculata (Antonius 1973, 1975) white-band disease (WBD) are still common at in these old surveys. both locations (Antonius 1981), coral-tissue- 146 REVISTA DE BIOLOGIA TROPICAL

! t N ....,,-,) Victoria 16°50' ('" l.-.~_...... o I ~ . Sillee Point ../ (_...•..•...... ~ Carrie Bow Cay l ...... ! ! I 1; ,I / Channel { 5km \

88°10'

Fig. I. To the left (= west) the mainland of Belize just south ofthe town of Dangriga with the mouth ofthe Sittee River. To the right (= east) the patch-reefsystem ofthe lagoon and the barrier reefwith Tobacco Reef in the north, Curlew Bank in the south and Carrie Bow Cay in the center.

Hawk

p~ ROCkS:'?'·

J 25°07' Greci~·~ Rocks if :-'{:" 1 km

80°18' Fig. 2. To the left (= west) Key Largo with Largo Sound (L. S.) and South Sound Creek (S). To the right (= east) the White Banks area with the Key Largo and Grecian Rocks reefs. The shelfedge marks the 301n isobath. bleaching (TBL) is rampant especially in MATERIALS AND METHODS Belize (Antonius, in prep.), while Florida is being invaded by an array ofpreviously little or Underwater, on-site observations were unknown syndromes, such as white-pox, yel­ made along a total oftwelve transects: 3 north low-blotch, white-plague (Goreau et al. 1998), and 3 south ofCarrie Bow Cay, Belize, as well white-plague type 11 (Richardson et al. 1998), as 3 transects on Key Largo Dry Rocks and 3 or rapid-wasting disease (Cervino et al. 1997). on Grecian Rocks, Florida. However, in our opinion, the combined phe­ The Belt Method was used throughout in nomena of epizoism, described in this paper, order to register all syndromc:s on corals in the may pose an even more severe threat to coral transects as indicators ofthe health condition of health and wellbeing of the entire reef ecosys­ the respective reefs (Antonius 1995a). It is a tem. semi-quantitative time-count technique that uses 30-min scans to quantify frequency of occurrence of coral diseases and other coral- ANTONIUS & BALLESTEROS: A new threat to coral health 147 degrading syndromes on a scale from zero to ing data on the changes that have taken place in six. However, for the purpose ofthis paper, per­ these reefs over time, will be forthcoming at a centages of afflicted specimens of the respec­ future date (Antonius, in prep.). tive coral species were calculated. An overall The second point listed in the Introduction, account ofcoral (and thus reef) health, dealing the new phenomena of other organisms over­ with all the presently known syndromes on reef growing living coral, is the subject of this corals (Antonius 1995b), will be published paper. We would like to point out that the des­ elsewhere (Antonius, in prep.). ignation as "new" phenomena shall not suggest Samples taken in the field were preserved in that these syndromes had not existed in the 4% formalin-seawater, as well as in 70% dena­ past, but merely that they had not been a threat tured ethanol. All voucher specimens were to coral and to coral reef health a few decades deposited in the Herbario del Centre d'Estudis ago. Avancats, CSIC, in Blanes, Spain. The respec­ tive accession-numbers are listed in the Blue-green algae: The Atlantic Barrier "Results" section. Reef immediately north of Carrie Bow Cay, Belize, consists ofan approximately 50m-wide reefcrest, made up mainly ofAcropora palma­ RESULTS ta and Millepora complanata, followed by a

For somebody who has not been back to these reefs for 25 years, the first sight was a TABLE 1 real shock. Especially the reefs around Carrie Summary ofpercentages ofcoral populations Bow Cay, in Belize which were teeming with overgrown by either cyanophytes, or macroal­ grouper, snapper, triggerfish, other colorful reef gae, or sponges in reefs ofCarrie Bow Cay, Belize and Key Largo, Florida. (n.r. = not reg­ fish, and with lobster and conch in the old days, istered) are now ghostly empty. The only larger fish seen during one month of survey was one par­ rotfish and two barracudas. Fish populations Belize Cyanophytes Florida % growing on % inside the National Marine Sanctuary at Key Largo, Florida, appeared to be in better shape. 50 Acropora cervicornis n.r. However, as mentioned under point one in the 50 Poriteslorites nx. n.r. reporte on 80 Introduction, large portions of reef-crest and and back-reef, which were documented as alive and Gorgonia in good condition 25 years ago, were found dead today. And this is true for Carrie Bow Belize Macr~algae Florida % growmgon % Reef in Belize, as well as for Grecian Rocks and Key Largo Dry Rocks in Florida. In all of 5 Millepora complanata IQ 10 these reefs, large expanses ofstill standing, rel­ 5 5 Porites astreoides 2 atively undamaged coral skeletons (mainly 10 Porites porites 2 Acropora palmata and Millepora complanata) can be found, as well as other areas where these Belize Sponges Florida % growing on % skeletons are weathered down to stumps, and others again where even these are obliterated IQ Dendrogyra cylindrus n.r. and reduced to rubble. Carrie Bow Reef was 5 Acropora palmata 1 5 Millepora complanata 1 surveyed in 1972 (Antonius unpublished), and 2 Montastrea annularis n.r. the Key Largo reefs in 1973 (Antonius 1974). 2 Diploria strigosa n.r. An accounting ofthese old (1972/73) results in 2 Diploria labyrinthiformis n.r. comparison with the new (1997) ones, contain- 1 Agaricia tenuifolia n.r. 148 REVISTA DE BIOLOGIA TROPICAL

Fig. 3. The typical "tufts" ofOscillatoria submembranacea on branches ofPorites porites. One ofthe branches has been com­ pletely covered by the cyanophyte.

roughly 200m-wide reef flat that is about lm swept in from the outer reef, getting caught in deep behind the reef crest and 2m deep toward the branches, is not known at present. the lagoon (RUtzler and Macintyre 1982). Right Following Drouet's , we identi­ behind the reefcrest, the bottom is mostly rub­ fied the "tufts" as Oscillatoria submem­ ble inbetween numerous Montastrea annularis branacea and the "flags" as Schizothrix mexi­ and Porites astreoides heads. Towards the cana, while according to classic taxonomy the lagoon, the floor becomes a sand-flat, studded "tufts" also match the characteristics of with Acropora cervicornis and Porites porites Microcoleus lyngbyaceus, and the "flags" those colonies. of Oscillatoria margaritifera. Since the taxon­ As mentioned earlier, compared to 1972, omy ofblue-greens is notoriously difficult, def­ the 1997 survey showed a much larger percent­ inite identification will undoubtedly require age of still standing, but dead coral colonies. further investigation. This was the case on the reefcrest as well as on Absolutely unequivocal, however, are the the reef flat. A substantial part of this coral effects ofthis kind ofovergrowth on the living mortality could be traced to cyanophytic over­ coral. There seems to be a positive correlation growth. About 50% ofall A. cervicornis and P. between the size ofthe area ofcontact between porites colonies (Table 1) scattered over the alga and coral, and the degree ofdegradation of reefflat show either "tufts" ofblue-green algae coral tissue: with increasing size ofthe area, the (Fig. J) or filamentous "flags" (Fig. 4) waving results of this contact become increasingly in the current. Both may cover substantial por­ severe. The coral tissue is usually found tions of these branching corals. Whether these bleached when the area ofcontact is as small as cyanophytes had grown on the coral branches a few square-millimeters; when the alga covers from minute germs, or whether they were several square-centimeters, the coral tissue is ANTONIUS & BALLESTEROS: A new threat to coral health 149

Fig. 4. Waving "flags" of Schizothrix mexicana on branch-tips of Acropora cervicornis. Coral tissue is being killed by the cyanophytic cover. mostly encountered in the process of dying; coral/epida is under way, since it is a new once cyanophytic overgrowth occupies areas in species. Both algae overgrow mainly Millepora the order of magnitude of square-decimeters, complanata (Fig. 5) and Millepora alcicornis the coral tissue has, in most cases, died off and from the bottom upwards, but also other corals is completely gone. with smooth surface, such as Porites porites This particular problem of cyanophytes (Fig. 6) and Porites astreoides. This syndrome invading coral populations was, in February occurs about equally abundant around Carrie 1997, encountered only to a minimal degree in Bow Cay, Belize, and in the reefs off Key Florida. However, blue-green algae can be ­ Largo, Florida with percentages of afflicted and have been - a problem in Key Largo reefs individuals of these species ranging from 2 to (Table 1) and will be dealt with again in the 10 % (Table 1). "Discussion". Both algae, Metapeyssonnelia coral/epida Samples of Oscillatoria submembranacea as well as Lobophora variegata form a tightly are deposited as EB-CBC-12 and EB-CBC-13, attached "skin" on the coral surface, with no while Schizothrix mecicana is represented by trace of living coral tissue left below the algal EB-CBC-14. cover. In the field, the two species are hard to distinguish, since both are very dark with razor­ Macroalgae: Another type of overgrowth sharp edge. In comparison, however, M on living corals is represented by the red alga coral/epida appears even darker than L. varie­ Metapeyssonnelia coral/epida, and the brown gata, with a violet hue and showing rather alga Lobophora variegata. While species deter­ straight edges, while L. variegata is tinted dark mination ofL. variegata did not pose any prob­ olive with more rounded edges. Both algae start lem, systematic evaluation ofMetapeyssonnelia to grow at the base ofcoral colonies which are 150 REVISTA DE BIOLOGIA TROPICAL

Fig. 5. The rhodophyte Metapeyssonnelia corallepida growing over Millepora complanata. The edge ofthe alga is relatively straight and its calor is a dark purple.

Fig. 6. The phaeophyte Lobophora variegata overgrowing some fmgers ofPorites porites. The edges ofthe alga are usually rounded and its color is a dark, drab olive-green. ANTONIUS & BALLESTEROS: A new threat to coral health 151

Fig. 7. The hadromerid sponge Cliona caribbea growing on a large Diploria labyrinthifonnis. It probably started out at some spot of denuded coral skeleton, but proceeds to overgrow living coral surface.

"dead", void of coral tissue and thus defense­ This does not mean, however, that less. From there they proceed upwards, now Metapeyssonnelia cora/lepida could not be overgrowing and destroying polyps and lethal for M comp/anata. Many cases have coenosarc. been registered of large frrecorals totally over­ Lacking observation time, the speed of the grown by the rhodophyte. Thus, these colonies process is not yet known, but the impact is evi­ still perfectly exhibit the shape and growth­ dent. Although coral polyps appear undisturbed form of M complanata, but they are now pur­ immediately in front of the advancing alga, plish-black instead of the typical live yellow. there is no trace of coral tissue left below the The same is true for L-c variegata overgrowing tightly attached algal cover. In fact, even the P porites : colonies of this coral showing fm­ surface structure of the coral skeleton appears gers which are olive-black instead ofthe normal to be eroded. Quite in contrast to other syn­ live, bright greenish-grey, have been killed and dromes, M cora/lepida and 1. variegata over­ overgrown by the phaeophyte. growing and thus killing off living coral tissue, Samples are deposited under the numbers do not appear to open up the way for other coral EB-CBC-7 and 11, as well as EB-KLF-l bioerosion, such as boring organisms. through 6. However, in some instances these overgrow­ ing macroalgae seem to trigger WBD (Antonius Sponges: A similar kind oftightly attached 1995b). This was definitely observed in cases of overgrowth, also starting at the base of the Lobophora variegata overgrowing Porites coral colony is represented by the hadromerid porites and is suspected - but could not be veri­ sponge Cliona caribbea. It was found regularly fied beyond a reasonable doubt - in cases of in reefs around Carrie Bow Cay on Acropora Metapeyssonnelia cora/lepida overgrowing palmata (Fig. 8), Millepora complanata, Millepora complanata. 152 REV1STA DE BIOLOGIA TROPICAL

Fig. 8. Close-up ofthe advancing front of Cliona caribbea on an Acropora palmata. The sponge not only kills off the coral polyps, but also penetrates the uppermost layer ofthe skeleton.

Montastrea annularis, Diploria strigosa, Of the substantial A. palmata and M com­ Diploria labyrinthiformis (Fig. 7), Agaricia planata populations south of Carrie Bow Cay, tenuifolia, and relatively frequently on an estimated 5% of all colonies may be afflict­ Dendrogyra cylindrus, but only very rarely on ed, while not more than 1-2% ofthe other coral Acropora palmata and Millepora complanata species are overgrown by C. caribbea (Table in Florida (Table I). I). The striking exception is D. cylindrus which However, Cliona caribbea covering live is not a frequent species around Carrie Bow corals may be the second stage of the invasion. Cay, but under heavy attack by C. caribbea The sponge readily grows on dead coral sur­ (Table I). Interestingly, this syndrome seems to .faces, often enough with no living coral any­ be much less of a problem in Florida where where near. From these footholds it proceeds to only very few cases were observed on A. overgrow live portions of the coral colony (Fig. palmata and M complanata (Table I). 7). In doing so, C. caribbea not only eliminates Another sponge, Chondrilla cl nucula, the coral's polyps and coenosarc, but also erodes apparently is behaving the same way. It starts the skeletal surface structure and even penetrates out at some dead spot of a coral colony, but the uppermost 1-2 cm of the coral skeleton. proceeds from there to overgrow living coral However, just as was observed in the case of surface. At present it does n9t seem to be a overgrowing macroalgae, coral polyps appear to threat to reef health though, since only three be undisturbed immediately in front ofthe lead­ cases were found in Carrie Bow Reef, and ing edge ofthe sponge. The speed ofadvance of none, so far, in Florida. One ofthese, however, C. caribbea is not yet known, but the position of had almost completely enveloped a large several ofthese leading edges have been marked colony ofDiploria strigosa (Fig. 9). in the field and will yield results in the future. ANTONIUS & BALLESTEROS: A new threat to coral health 153

Fig. 9. Another sponge, Chondrilla cf nucu/a, in the process ofovergrowing a large colony ofDip/oria strigosa. Although the sponge will kill living coral surface, polyps in the immediate vicinity appear undisturbed.

DISCUSSION 1997. However, just as was noticed in Belize, dead reef areas in Florida have also increased As summarized before (Antonius 1995b), enormously when compared to the situation in the fact that different organisms can overgrow the early 1970s. Among several other syn­ and kill reef corals is nothing new. What may dromes, it is not out of the question that be different from previous observations is the cyanophytes too may have contributed to this fact that these syndromes have now become a decline. After all, just as evidenced in case of factor controlling growth and development, at Phormidium corallyticum, the pathogen of the least ofthe reefs investigated. BBD (Antonius 1985), other cyanophytes are Overgrowing blue-green algae were noticed also seasonal. These seasons are much more in the past. For instance, investigations of coral pronounced in the waters ofFlorida, than in the reef decline in Mauritius in the early 1990s, more southerly Caribbean, such as Belize. revealed a widespread occurrence ofcyanophytes BBD, for instance, disappears in Florida in in various reef areas. However, being overshad­ winter, but remains active in Belize (Antonius owed by so many other coral-killing syndromes, 1981). Thus, it is entirely possible that over­ they were not recognized as active coral-killers growing blue-greens play a major role in at that time (Antonius 1991a, b; Ballesteros Floridian reefs in the hot season ofthe year. 1994, in press). In the very recent past, for example, span­ Although overgrowing cyanophytes were ningseveral consecutive summer seasons, registered as rampant at Carrie Bow Reef, there has been a massive invasion of filamen­ Belize in the present investigation (Table 1), tous blue-green algae in a large patch reefto the there was hardly any evidence ofthis problem NW of Key Largo Dry Rocks. These in the reefs ofKey Largo, Florida, in February cyanophytes grew in dense bushes, up to 154 REVISTA DE BIOLOGIA TROPICAL almost half-a-meter in height and covering Millepora colonies inside a field of living, large patches, killing off not only stony corals, bright yellow ones. but also the very frequent sea-fans (1. Halas What both algae seem to have in common is pers. comm.). Thus, the reef is locally known as the fact that they can trigger WBD, a phenome­ the "algae reef' since then, and the cyanophyte non previously only observed to be linked with was tentatively identified as Lyngbya sp. cyanophytes (Antonius 1987, 1988). Through this Today, most ofthe stony corals are dead, and so mechanism L. variegata as well as M cora/lepi­ is the dense population of Gorgonia ventalina da may amount to a force much more destructive (Table I). The large skeletons are still there, than any ofthem would be on their own. waving in the surge, while a new generation of Among coral-overgrowing and coral-killing now hand-sized fans has begun to grow. sponges, the most notorious to date is probably At Carrie Bow Reef in Belize, which is far Terpios sp. (Bryan 1973, Plucer-Rosario 1987 away from sources of man-made pollution, we and. Antonius 1991b), or Terpios hoshinota suspect that the problem is related to a decline respectively (Antonius 1993, Riitzler & Muzik in Caribbean water quality in general. Where 1993). T hoshinota which can infest hundreds should cyanophyte-promoting water-conditions of meters of coral reef in Micronesia (Bryan come from, if not from the open Caribbean 1973), has also been reported from the northern Sea? After all, northeasterly trade-winds pre­ Pacific, from Samoa to Taiwan (Plucer-Rosario vail in Belize during 70% ofthe year (Riitzler 1987), from southern Japan (Rtitzler & Muzik & Ferraris 1982). 1993), as well as from Mauritius (Antonius Of the array of coral-overgrowing organ­ 1993) and Sinai. (Antonius 1996). But it was isms, the brown alga Lobophora variegata is never observed in Atlantic waters. the only one previously observed by the However, there seems to exist a true coun­ authors to damage corals. During investiga­ terpart in the Caribbean. Cliona caribbea tions of coral health in Mauritius, mentioned turned out to amount to a major coral-killing earlier, L. variegata was observed to overgrow agent in reefs around Carry Bow Cay, Belize, small specimens of Porites sp., Favia ste/lig­ and also occurring in Florida. Cliona species, era., and others (Antonius 1991 b, 1995b, such as C. caribbea, C. lampa, and C. varians Ballesteros, in press). However, even in the par­ have been observed as successful competitors tially very sick coral populations of Mauritius for space before (Vincente 1978). However, (Antonius 1993, Ballesteros, in press), the one species, C. caribbea, now occurring in Lobophora-syndrome (LOB) was not encoun­ numbers controlling coral reef growth, may be tered as frequently as in the reefs around Carrie a new development. In contrast to the relative­ Bow Cay. Especially among the Porites porites ly species-specific coral-overgrowing algae populations of the back-reef area, LOB really described here, C. caribbea does not seem to amounts to a controlling factor. have preferences, and is overgrowing a wide While LOB seems to be less of a threat in variety of different coral species (Table 1) in a Florida, another overgrowing macroalga, the rather indiscriminate manner. rhodophyte Metapeyssonnelia cora/lepida is As mentioned earlier, the sponge does not dominant there. In Florida, this red alga is prey on corals, and probably can not even settle mainly attacking Millepora complanata in reef~ on live, healthy coral tissue. But it flourishes on crest areas. To the best of our knowledge, M denuded coral limestone surfaces, and from this corallepida was an undescribed species secure foothold has no difficulty in subsequent­ (Ballesteros et al., in prep.) and, therefore, it ly overgrowing the living coral . Just as was never observed before or it was misidenti­ Terpios hoshinota is aided by cyanobacterial fied. Just as L. variegata can envelope entire symbionts in overpowering corals, Cliona coralla, so can M cora/lepida. It is a striking caribbea harbors large quantities ofzooxanthel­ sight to fmd totally black, alga-overgrown lae which impart the same advantage. ANTONIUS & BALLESTEROS: A new threat to coral health 155

Another true cyanobacteriosponge is por parte de diversos organismos bent6nicos. Las poblacio- . Chondrilla sp. in Carrie Bow Reef, which nes de Acropora cervicornis y Poriles poriles situadas sobre 105 fondos arenosos someros de la plataforma arrecifal del could be C. nucula, described as coral-over­ norte de Carrie Bow Cay estaban, en parte, recubiertas por growing from Puerto Rico (Vincente 1990), but penachos 0 laminas de las cianoficeas Osci/laloria submem­ might be a new species (K. ROtzler, pers. branacea y Schizolhrix mexicana. Aproximadamente el comm.). The Chondrilla species found in 50% de 105 corales estaban afectados, con destrucci6n del tejido vivo del coral en las areas de contacto con las ciano­ Belize, however, is a whole order ofmagnitude ficeas. Desconocemos si las cianoficeas se desarrollaron di­ less frequent than Cliona caribbea and does not rectamente sobre los corales 0 bien si se engancharon a ellos (yet?) pose a threat to the health of local reefs. provenientes del sistema pelagico. Sea como fuere, su asen­ Summing up these observations, it is evident tamiento degrada 105 corales de forma considerable. Otro ti­ po de epizoismo es el producido por la rodoficea Melapeys­ that 'a substantial percentage of the coral popu­ sonnelia cora/lepida y la feoficea Lobophora variegala. lations one of us (A. Antonius) investigated are Ambas especies crecen principalmente sobre Mi/lepora afflicted with syndromes discussed here (Table complanala y M alcicornis, aunque tambien recubren otros 1). Thus, there can be no doubt that these coral­ corales lisos como Poriles porites y Poriles aslreoides. Tan­ to Melapeyssonnelia como Lobophora estlln fijadas fuerte­ killing phenomena together amount to a consid­ mente aI coral y destruyen sus tejidos, y avanzan desde la erable factor degrading coral reefs. What should base hacia 105 apices de 105 corales. Ambas especies estan be cause for more concern, however, is the fact presentes en Carrie Bow Cay y Key Largo. Un tipo de recu­ that none ofthese syndromes amounted to a rec­ brimiento parecido 10 constituye la esponja Cliona carib­ bea, recolectada sobre Acropora palmala, Mi/lepora com­ ognizable factor in any of these reefs some 25 planala, Monlaslrea annularis, Diploria strigosa, D. laby­ years ago. They certainly did not show up in the rinlhiformis, Agaricia lenuifolia, y, especialmente, Den­ painstakingly recorded old transects. How this drogyra cylindrus, en Carrie Bow Cay. La esponja Chondri­ situation is going to develop in the future should /la cf nucula crece sobre Diploria strigosa de forma identi­ ca a Cliona caribbea. Todos estos nuevos sindromes no be monitored carefully. existian hace 25 ai'los y actualmente representan, en conjun­ to, un importante factor de degradaci6n de 105 arrecifes.

ACKNOWLEDGMENTS REFERENCES Support of these investigations by the Austrian Science Foundation (FWF) and the Antonius, A. 1973. New observations on coral destruction Smithsonian Institution (for fieldwork in in reefs. Abs. Assoc. 151. Mar. Lab. Carib. 10: 3. Belize) is gratefully acknowledged. Joint Antonius, A. 1974. Final report of the coral reef group of research by the co-authors was facilitated by a the Florida Keys Project for the project year 1973. Harbor Branch Foundation, Fort Pierce, Florida. 201 p. travel grant from the bureau of "Acciones Integradas Hispano-Austriacas". Cordial per­ Antonius, A. 1975. Predation by the Hermodice sonal thanks go to K. Riitzler and to J. Halas for carunculala on reef corals. In H.G. Multer (ed.). Field guide to some carbonate rock environments. Kendall invaluable assistance in the field. Hunt. pp. 133K-133M. Antonius, A. 1981. The "band" diseases in coral reefs. Proc. Fourth Internal. Coral Reef Symp., Univ. RESUMEN PHilippines, Manila 2: 7-14.

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