ATOLL RESEARCH BULLETIN NO. 172 Comparative Investigations of Tropical Reef Ecosystems: Background fo r an integrated coral reef progray , Edited by Marie-Helene Sachet and Arthur L . Dahl Issued by THE SMITHSONIAN INSTITUTION Washington, D.C., U.S.A. CONTENTS Introduction . . . . . . . . . . . . . . . . . . . . . . S. V . Smith. 1 A preliminary coral reef ecosystem model A . L . Dahl, 5. C . Patten, S . V . Smith, . . . . . . . . . . . . . . . . . and J . C . Zieman, J r . 7 A comparative survey of coral reef research s i t e s A . L . Dahl, I . G . Macintyre, and A . Antonius. . . . . . . . . . . . . . . . . . . . . . 37 State of knowledge of coral reefs as ecosystems . . . . . . . . . . . . . . . . . . . . . M . - H . Sachet 121 INTRODUCTION CITRE and IMSWE studies i n B r i t i s h Honduras S. V. Smith A se r ies o f events a t the Smithsonian I n s t i t u t i o n beginning about 1970 and cont inu ing t o the present have l e d t o the f o l l o w i n g group of r e l a t e d papers dea l ing w i t h var ious aspects o f co ra l reef ecology, p r i m a r i l y i n r e l a t i o n t o the r e e f s o f B r i t i s h Honduras (Be l i ze ) . A b r i e f h i s t o r i c a l sketch o f those events provides usefu l background in fo rmat ion t o r e l a t e the papers presented here. During t h e summer o f 1970 researchers from several i n s t i t u t i o n s began consider ing the p o s s i b i l i t y o f seeking funding f o r co ra l reef research from the I n t e r n a t i o n a l Decade o f Ocean Exp lora t ion (IDOE) o f f i c e o f the Nat ional Science Foundation. This considerat ion, together w i t h d r a f t proposals and in formal d iscussion, l e d i n the spr ing o f 1971 t o a grant from IDOE t o t h e Smithsonian I n s t i t u t i o n f o r the fo rmula t ion and coord ina t ion of a long-term, mu1 t i - d i s c i p l inary , mu1 t i - i n s t i t u t i o n a l i n v e s t i g a t i o n o f cora l r e e f ecosystems. That p r o j e c t was named "Comparative Inves t i ga t i ons o f Trop ica l Reef Ecosystems" (CITRE). A1 so i n the sp r ing o f 1971, the Smithsonian s c i e n t i s t s received i n t e r n a l f i n a n c i a l support f o r a more l i m i t e d e f f o r t designed t o describe b i o t i c and a b i o t i c c h a r a c t e r i s t i c s o f co ra l ree fs . That p r o j e c t was named " Inves t i ga t i ons o f Marine Shallow-Water Ecosystems" (IMSWE). Smithsonian s c i e n t i s t s invo lved i n the one p r o j e c t were f o r the most p a r t invo lved i n the o the r a s we l l , so an e f f o r t was made t o coordinate the two a c t i v i t i e s as much as poss ib le . As soon as CITRE planning funds became ava i l ab le , Smithsonian s c i e n t i s t s began gather ing a v a i l a b l e i n fo rma t ion from which t o s e l e c t s i t e s f o r e s t a b l i s h i n g f a c i l i t i e s on a cora l r e e f . Po ten t i a l s i t e s were then v i s i t e d . Dahl, Macintyre, and Antonius repor ted on these s i t e se lec t i on e f f o r t s , and t h e i r r e p o r t i s presented here i n rev i sed form. G lover 's Reef, B r i t i s h Honduras, was selected as the i n i t i a l s i t e f o r CITRE e f f o r t s ; a dec i s ion on Caribbean secondary s i t e s and on a P a c i f i c pr imary s i t e was deferred f o r a l a t e r stage i n the CITRE p r o j e c t . Dur ing November 1971, a two-week workshop was h e l d a t G l o v e r ' s Reef. The purpose o f t h a t workshop was t o i n t r o d u c e workshop p a r t i c i p a n t s t o techn iques o f q u a n t i t a t i v e to ta l -ecosys tem model ing and t o g a t h e r m a t e r i a l f o r p r e p a r i n g t h e formal CITRE p r o p o s a l . Workshop p a r t i c i p a n t s had been s e l e c t e d t o be i n one o r more o f n i n e work ing groups which had been e r e c t e d as b a s i c u n i t s w i t h i n t h e CITRE p r o j e c t . P a r t i c i p a n t s and t h e i r p r i m a r y work ing groups a r e g i v e n i n Table 1 . That t a b l e does n o t c a p t u r e t h e i n t e r a c t i o n among t h e p a r t i c i p a n t s ; most i n d i v i d u a l s were i n v o l v e d t o some e x t e n t i n a t l e a s t one work ing group bes ides t h e p r i m a r y one i n which t h a t t a b l e l i s t s them. The CITRE proposal was submi t ted t o IDOE i n January 1972. That proposa l was n o t funded, b u t t h e e f f o r t s produced v a l u a b l e r e s u l t s . We a r e u s i n g t h e A t o l l Research B u l l e t i n t o make these r e s u l t s g e n e r a l l y a v a i l a b l e r a t h e r than l o s i n g them because o f t h e f a t e o f t h e proposa l i t s e 1 . f . Dahl , Pat ten, Smith, and Zieman summarize t h e conceptua l model which was developed a t G l o v e r ' s Reef. That model i s v e r y p r e l i m i n a r y , and p a r t s o f i t a r e be ing o r a l r e a d y have been r e v i s e d f o r p u b l i c a t i o n e l sewhere. Sachet p resen ts t h e b i b l i o g r a p h i c s e c t i o n o f t h e CITRE p roposa l . It should be emphasized t h a t t h e r o l e o f t h e i n d i v i d u a l s p r e s e n t i n g s e c t i o n s o f t h e p roposa l has been e d i t o r i a l ; a l l o f t h e workshop p a r t i c i p a n t s (Tab le 1 ) and many o t h e r i n d i v i d u a l s c o n t r i b u t e d m a t e r i a l 1 y t o these papers. I n a d d i t i o n t o t h e papers adapted f rom t h e CITRE s i t e s e l e c t i o n r e p o r t and p roposa l , severa l r e l a t e d papers have been produced. L a r g e l y as a r e s u l t o f e f f o r t s d u r i n g t h e s i t e survey and workshop, Tsuda and Dawes p r e s e n t a check1 i s t o f mar ine a lgae a t G l o v e r ' s Reef. Severa l papers on t h e geographic and t e r r e s t r i a l aspects o f t h e B r i t i s h Honduran r e e f a rea w i l l appear i n a subsequent i s s u e o f t h e A t o l l Research B u l l e t i n . We hope t h a t t h e papers p resen ted i n t h i s i s s u e o f ARB w i l l c o n t i n u e t h e c o n s i d e r a b l e i n t e r e s t which has been engendered i n t h e i n t e g r a t e d approach t o c o r a l - r e e f eco logy as v i s u a l i z e d by t h e CITRE p a r t i c i p a n t s . Moreover, we a r e c o n f i d e n t t h a t c o n t i n u e d e f f o r t s i n B r i t i s h Honduras w i l l serve t o e l u c i d a t e t h e n a t u r e o f a complex, impor tan t , b u t poor ly-known a rea o f Caribbean c o r a l r e e f s . Table 1 . Part icipants a t the Glover's Reef Workshop Stephen V . Smith Principal Investigator Smithsonian Ins t i tu t ion (presently a t Hawaii I n s t i t u t e of Marine Biology) S i r Maurice Yonge External Advisory Committee Member Edinburgh, Scotland George D. Grice NSF Observer National Science Foundation Ecosystem Analysis Working Group Bernard C . Patten Group Leader University of Georgia Thelma Richardson University of Georgia Joseph C. Zieman University of Virginia Detritus and Nutrients Working Group Robert E. Johannes Group Leader University of Georgia Donald W . Kinsey Mauri Bros. & Thompson, Sydney, Australia Nelson Marshall University of Rhode Island Michael E . Q. Pilson University of Rhode Island Kenneth L . Webb Virginia In s t i t u t e of Marine Science Benthic Plants Working Group Arthur L . Dahl Group Leader Smithsonian Ins t i tu t ion Michael S. F o s t e r U n i v e r s i t y o f C a l i f o r n i a , Santa Barbara ( p r e s e n t l y a t C a l i f o r n i a S t a t e U n i v e r s i t y , Hayward) Roy T. Tsuda U n i v e r s i t y o f Guam I n v e r t e b r a t e Working Group P e t e r W . Glynn Group Leader Smi thson ian T r o p i c a l Research I n s t i t u t e A r n f r i e d An ton ius Smi thson ian I n s t i t u t i o n ( p r e s e n t l y Harbor Branch Foundat ion, F l o r i d a ) Judy Lang Smi thson ian T r o p i c a l Research I n s t i t u t e ( p r e s e n t l y U n i v e r s i t y o f Texas, A u s t i n ) James P o r t e r Smi thson ian T r o p i c a l Research I n s t i t u t e ( p r e s e n t l y U n i v e r s i t y o f M ich igan) Amada Reimer Smi t h s o n i a n T r o p i c a l Research I n s t i t u t e (p resen t1 y Pennsy lvan ia S t a t e U n i v e r s i t y ) K laus R u e t z l e r Smi thson ian I n s t i t u t i o n P l a n k t o n Working Group Tom S. E n g l i s h U n i v e r s i t y o f Washington Group Leader A r t h u r B a r n e t t S c r i p p s I n s t i t u t i o n o f Oceanography Frank F e r r a r i Texas A & N U n i v e r s i t y Robin Ross U n i v e r s i t y o f Washington V e r t e b r a t e s Working Group Ray S . B i rdsong O ld Dominion U n i v e r s i t y Group Leader James E. Boh lke P h i l a d e l p h i a Academy o f N a t u r a l Sc iences E d i t h t i . Chave U n i v e r s i t y o f Hawai i Dav id W . G r e e n f i e l d U n i v e r s i t y o f I l l i n o i s C . L a v e t t Smi th American Museum o f N a t u r a l H i s t o r y , New York Frank H. T a l b o t The A u s t r a l i a n Museum Geology Working Group I a n G. M a c i n t y r e Smi thson ian I n s t i t u t i o n K e i t h E. Chave U n i v e r s i t y o f Hawai i Clyde Moore L o u i s i a n a S t a t e U n i v e r s i t y Jon N. Weber Pennsy lvan ia S t a t e U n i v e r s i t y Group Leader T e r r e s t r i a l Phenomena Working Group F. Raymond Fosberg Group Leader Smi thson ian I n s t i t u t i o n Marie-He1 ene Sachet Smi t h s o n i a n I n s t i t u t i o n Ralph W. S c h r e i b e r U n i v e r s i t y o f South F l o r i d a Dav id R. S t o d d a r t Cambridge U n i v e r s i t y Oceanography and Meteoro logy Working Group Andrew C . Vastano Group Leader Texas A & M U n i v e r s i t y Bruce W. M c A l i s t e r N a t i o n a l Science Foundat ion A PRELIMINARY CORAL REEF ECOSYSTEM PODEL E d i t e d by A. L. Dahl , 8. C. Pa t ten , S. V. Smith, and J. C . Zieman, Jr. A ma jo r goa l o f t h e CITRE p l a n n i n g e f f o r t was t o deve lop a concep- t u a l framework f o r a systems model o f a c o r a l r e e f ecosystem s u i t a b l e f o r c o m p u t e r i z a t i o n . Bo th t h e process o f development and t h e p r e l i m i n - a r y r e s u l t s may be o f some v a l u e t o f u t u r e comprehensive s t u d i e s o f complex r e e f systems. The p r e l i m i n a r y model was conce ived n o t o n l y t o o u t 1 i n e a p o s s i b l e mathemat ica l model, h u t a l s o t o i n t e g r a t e t h e proposed r e s e a r c h o f many s p e c i a l i s t s i n t o a coheren t framework w i t h d e f i n e d i n t e r a c t i o n s . Wh i le t h e s p e c i a l i s t s generated t h e model s t r u c t u r e , t h e y a l s o had t o m o d i f y t h e i r r e s e a r c h p l a n s t o i n c o r p o r a t e a l l m a j o r aspec ts o f t h e system as i d e n t i f i e d i n t h e model . No one model can dea l w i t h a l l aspects o f an ecosystem. I t s p r o p e r - t i e s , however, shou ld u l t i m a t e l y resemble those o f t h e system b e i n g modeled and i t s l e v e l o f a b s t r a c t i o n shou ld s u i t i t s i n t e n d e d purpose. Thus no model i s s t a t i c ; i t must e v o l v e as a v a i l a b l e knowledge o f t h e ecosystem i n c r e a s e s . The p r e l i m i n a r y model d e s c r i b e d h e r e i s one f i x e d p o i n t i n such an e v o l v i n g process. I t r e p r e s e n t s an i n t e r m e d i a t e l e v e l o f e c o l o g i c a l a b s t r a c t i o n , which, because o f t h e d i v e r s i t y o f t h e r e e f system, i s n e v e r t h e l e s s h i g h l y complex. I t i s based on t h e g r e a t d i v e r - s i t y o f v i e w p o i n t s and p r o f e s s i o n a l exper iences o f t h e i n v e s t i g a t o r s i n v o l v e d . Yet , w h i l e i t s d e t a i l s a r e open t o q u e s t i o n , i t r e p r e s e n t s a usefu l benchmark f r o m which t o proceed. The mode l was developed d u r i n g a two-week workshop a t G l o v e r ' s Reef, B r i t i s h Honduras, i n November 1971. Fo r t y -one p a r t i c i p a n t s ( s e e I n t r o - d u c t i o n ) r e p r e s e n t i n g a wide v a r i e t y o f r e e f - r e l a t e d d i s c i p l i n e s and w i t h exper ience i n many c o r a l r e e f areas a t t e n d e d t h i s workshop. The p r i n c i p a l a c t i v i t y was t h e development o f a conceptua l model s u f f i c i e n t l y comprehensive t h a t i t c o u l d be a p p l i e d t o most r e e f areas. However, t h e s h a l l o w r e e f f l a t a reas o f G l o v e r ' s Reef served as t h e p r ime f o c u s f o r t h e i n i t i a l e f f o r t and t h i s may be r e f l e c t e d i n some o f t h e d e t a i l s p resen ted here . O v e r a l l , t h e r e s u l t r e p r e s e n t s a un ique s y n t h e s i s o f c u r r e n t i n f o r m a t i o n about ' t h e t r o p i c a l r e e f ecosystem. Process o f Model Development CITRE p a r t i c i p a n t s were i n i t i a l l y chosen t o be members o f one o r more o f t h e n i n e work ing groups ( I n t r o d u c t i o n ) w i t h r e s p o n s i b i l i t y f o r a ma jo r aspec t o f c o r a l r e e f eco logy . Most work ing groups met b e f o r e t h e workshop f o r i n i t i a l d i s c u s s i o n o f t h e i r s u b j e c t a rea and an i n t r o - d u c t i o n t o mode l ing concepts . The workshop i t s e l f i n c l u d e d l e c t u r e s i n t r o d u c i n g t h e s c i e n t i s t s t o some aspects o f mode l ing t h e o r y , and p r e s e n t a t i o n s by work ing groups as t h e y f i t t e d t h e i r s u b j e c t m a t t e r i n to t h e genera l model ing f ramework. The work ing groups proceeded by i n t e r n a l d i s c u s s i o n s , c o n s u l t a t i o n s w i t h t h e modelers and w i t h o t h e r work ing groups, and f i e l d o b s e r v a t i o n s t o check on t h e genera l a p p l i c a b i l i t y o f t h e model e lements deve loped. The m inor inconven iences o f work ing a t a remote s i t e were f a r outweighed by t h e freedom f rom d i s t r a c t i o n s and t h e a b i l i t y t o " c o n s u l t t h e env i ronment . " The model was des igned t o d e p i c t t h e f l o w o f carbon t h r o u g h t h e c o r a l r e e f ecosystem. Each w o r k i n g group f i r s t d e f i n e d t h e "compartments," o r f u n c t i o n a l u n i t s between wh ich t h e f l o w s were t o be measured. An approx imate l i m i t o f twen ty compartments p e r w o r k i n g g roup was s e t t o f i x t h e o v e r a l l model c o m p l e x i t y w i t h i n p r e s e n t computer c a p a b i l i t i e s . I n t e r - g r o u p d i s c u s s i o n s r e s o l v e d problems o f o v e r l a p p i n g o r o v e r l o o k e d f u n c t i o n a l u n i t s . Diagrams were then developed f o r each compartment r e l a t i n g i t t o o t h e r compartments and f l o w s i n t h e system. The compart- ments were l i s t e d i n a m a t r i x , and t h e i n t e r a c t i o n s between compartments determined j o i n t l y by t h e w o r k i n g groups i n v o l v e d . Some q u a n t i f i c a t i o n o f these i n t e r a c t i o n s was a t tempted ( a t l e a s t i n terms o f o r d e r s o f magnitude), b u t t h i s was n o t completed and i s o m i t t e d here . R e v i s i o n s and ad jus tmen ts c o n t i n u e d t h r o u g h o u t t h e workshop. The t e r r e s t r i a l s e c t i o n was k e p t s i m p l i f i e d as t h e p r i m a r y concern was i t s i n t e r a c t i o n s w i t h t h e r e e f system. The model developed t h r o u g h a c y c l i c a l p r o g r e s s i o n . A s ta tement o f t h e purpose o f t h e model and t h e d e f i n i t i o n o f compartments was f o l l o w e d by t h e c o n c e p t u a l i z a t i o n o f t h e i n i t i a l model and f i e l d o b s e r v a t i o n s t o check t h e model v a l i d i t y , l e a d i n g t o changes i n t h e c o n c e p t u a l i z a t i o n and (more r a r e l y ) i n compartment d e f i n i t i o n s . W i t h each p e r m u t a t i o n t h e model became more r e f i n e d and ( h o p e f u l l y ) a c c u r a t e . The Model Compartments Three b a s i c e lements c h a r a c t e r i z e t h e model : compartments, f l o w s ( f l u x e s ) and e x t e r n a l d r i v i n g f o r c e s ( f o r c i n g f u n c t i o n s ) . The compart- ments a r e t h e carbon o r c a r b o n - e q u i v a l e n t s t o r a g e u n i t s o f t h e system and may be d e f i n e d as p l a n t o r an imal t ypes t h a t a c t as p r o c e s s i n g u n i t s f o r t h e food which t h e y i n g e s t o r produce, o r as m a t e r i a l p o o l s such as d e t r i t u s o r carbon d i o x i d e t h r o u g h which substances pass as t h e y c y c l e i n t h e system. Compartments i n t h e CITRE model may be c o n s i d e r e d " f u n c t i o n a l groups" i n t h a t t h e y a r e a b s t r a c t condensa t ions o f groups o f organisms o r substances t h a t a p p a r e n t l y have t h e same o r s i m i l a r f u n c - t i o n s i r t h e ecosystem. I n t h i s sense t h e y a r e " e c o l o g i c a l spec ies " as opposed t o c o n v e n t i o n a l taxonomic spec ies . Fo r example, Hal imeda, P e n i c i l l u s , Rh i oce ha lus , Udotea, Jan ia , and C o r a l l i n a a r e d i s t i n c t genera o f a l g a e i?r_P___ even be long ing t o two p h y l a ) ; y e t i n t h e model t h e y a r e a l l cons ide red p a r t o f t h e same compartment-- the noncrustose, c a l c i u m ca rbona te -p roduc ing macroalgae d e s i g n a t e d XCBALG* [6] ( s e e T a b l e I ) . L i k e w i s e two compartments a r e d i s t i n g u i s h e d on whether m o t i l e organisms remove s o l i d subs t ra tum as t h e y f w d ( " s c r a p e r s " : des igna ted XSCRAP [35] ) , o r e a t o n l y t h e o v e r l y i n g organisms ( "b rowsers " : XBROWS [ 3 6 ] ) . A l l d i s s o l v e d o r g a n i c compounds c o n t a i n i n g n i t r o g e n , and n o t a s s o c i a t e d w i t h some o t h e r compartment, a r e s i m i l a r l y grouped (XDON [ 6 7 ] ) . Compartment boundar ies n o t o n l y combine d i s s i m i l a r spec ies on f u n c t i o n a l grounds, i n some cases t h e y even separa te p a r t s o f organisms w i t h d i s t i n c t r o l e s o r l o c a t i o n s . Mar ine g rass b lades (XBLADE [9] ) were separated f r o m mar ine g rass r o o t s (XROOT [ l o ] ) because o f t h e f u n c t i o n a l d i f f e r e n c e s i n h e r e n t i n t h e l o c a t i o n o f these p a r t s above versus below t h e sed iment s u r f a c e . T r a n s p o r t f rom one p a r t o f t h e p l a n t t o ano the r t h u s becomes a f l o w between compartments. When i t appeared t h a t t h e i n o r g a n i c ca rbon c y c l e c o u l d be mos t e a s i l y t r e a t e d as a d i s - t i n c t system, separa te compartments were e s t a b l i s h e d f o r t h e c a l c i u m carbonate i n t h e w a l l s o f l i v i n g organisms (XORG [84] ) and t h e r e m a i n i n g ( o r g a n i c ) carbon i n t h e organisms. The organisms were t h e r e f o r e d e f i n e d i n t h e model as c o n t r o l l i n g t h e f l o w o f carbon t o t h e i r s k e l e t o n s r a t h e r t h a n f l o w i n g t h a t carbon t h r o u g h t h e organ ism compartments. Tab le I enumerates t h e p r e l i m i n a r y w o r k i n g compartments i d e n t i f i e d f o r t h e CITRE model, w i t h t h e i r acronyms and numbers. I n o r d e r t h a t compartments may f u n c t i o n w i t h i n t h e model , t h e y mus t be l i n k e d o r coupled; f l o w s ( o r f l u x e s ) between them a r e measured i n whatever i s t h e c u r r e n c y t h e model --i .e., t h e m a t e r i a l w i t h wh ich exchanges a r e made. Flows, 1 i ke compartments, a r e based on t h e genera l o b j e c t i v e s o f t h e mode l ing program. I n t h e case o f t h e CITRE mode l the f o l l o w i n g m a t e r i a l s were cons ide red s i g n i f i c a n t t o t h e model o f a c o r a l r e e f ecosystem: carbon, i n o r g a n i c carbon, o r g a n i c carbon, n i t r o g e n , phosphorus, ca l c ium, biomass, and energy. It was f i n a l l y agreed t o f l o w carbon t h r o u g h t h i s model because o f i t s mutua l impor tance t o t h e b iochemica l and geochemical c y c l e s , even though o t h e r f l o w s a r e a l s o i m p o r t a n t . Fo r example, t h e n u t r i e n t s (PO , NO3, e t c . ) wh ich a r e i m p o r t a n t i n feed-back l o o p s c o n t r o l l e d by pho tosyn th?s is , a r e f l o w e d i n t h e model, b u t f o r mathemat ica l c o n s i s t e n c y , a r e conver ted t o ca rbon e q u i v a l e n t s . * A l though t h e r e i s no s tandard c o n v e n t i o n f o r naming compartments, f l o w s , o r o t h e r components, c e r t a i n c o n s i s t e n c i e s make bookkeeping e a s i e r and more accura te , such as t h e use o f acronyms. Here X---- s tands f o r com- par tment , t hus XCBALG f o r CarBonate p r o d u c i n g U a e and XDON f o r g i s s o l v e d Organ ic N i t r o g e n . For convenience, a1 1 compartments were numbered - consecu tTve ly as t h e y appeared i n t h e m a t r i x ( F i g . 9 ) . Forcing func t ions Forc ing func t ions a r e d r i v i n g forces o r va r i ab les which o r i g i n a t e outs ide the system o f re fe rence b u t which i n f l u e n c e the behavior o f t h e system. These var iab les i r ~ c l ude 1 i g h t , temperature, inputs o f mater ia ls , and o the r in f luences no t under the cont ro l o f t h e system. The diagrams used i n t h i s model t o i l l u s t r a t e compartments and f lows are feedback dynamics o r Fo r res te r diagrams (Forrester , 1961). The u n i t s o f compartments and f lows i n the CITRE model a r e grams o f carbon per square meter (gC m-2) and grams o f carbon per square meter per day (gC m-2 day- l ) respect ive ly . F igure 1 shows the symbols used i n the d ia - grams and b r i e f l y describes meanings attached t o these symbols. Although these diagrams may become very complex, they are usefu l f o r a graphic representa t ion o f the model and as a means t o f a c i l i t a t e discussion. The general c h a r a c t e r i s t i c s o f the t o t a l CITRE ecosystem model as descr ibed by these diagrams i s out1 ined below. The f o r c i n g func t ions (ex te rna l d r i v i n g fo rces) , compartments and f lows between compartments, together del i neate a p re l im ina ry t o t a l ecosystem model f o r a co ra l r e e f . Both the l a r g e number o f compartments (104) and the number o f f lows between them form a very complex model. Figure 2 g r a p h i c a l l y i l l u s t r a t e s the coup1 ings and eco log ica l r e l a - t ionsh ips o f one compartment, t h e f leshy algae (XFLALG [51 . C02 i n p u t from XDISOL (80), a compartment o f common i n t e r e s t t o the Geology and Nu t r i en ts -De t r i t us groups, i s c o n t r o l l e d by f o u r i n t e g r a t i n g funct ions, three of which (#1 through #3) a re a b i o t i c func t ions t h a t regu la te poten- t i a l photosynthesis. I n t e g r a t i n g func t i on #4, coupled w i t h the o t h e r three, gives r e a l i z e d photosynthesis. Dashed arrows f rom t h e i n t e g r a t i n g func t i ons t o the valves on the f lows show t h a t the i n t e g r a t i n g func t i ons in f luence f low w i thou t c o n t r i b u t i n g m a t e r i a l t o it, t h a t i s they are in fo rmat iona l o r cont ro l coupl ings r a t h e r than mater ia l f lows o r f luxes . S i x feedback loops o r c y c l i c a l f lows, o f t h r e e bas i c types, a f f e c t t h i s compartment. Two s i m i l a r loops (diagrammed as one) i n d i c a t e the c y c l i n g o f carbon, o r carbon equ iva len ts o f oxygen, respec t i ve l y , between the d issolved organic carbon (XDOC [62] ) o r oxygen (X02 [70] ) compart- ments and XFLALG (5). These loops a r e c o n t r o l l e d by the amount o f the "upstream" o r donor compartment present as w e l l as by temperature, s a l i n i t y , and exposure ( i n t e g r a t i n g func t i on # I ) . Another type o f loop i n the diagram shows the c y c l i n g o f carbon t o XTURF (4) , XPHPL2 (12), and XPHPL3 (13). Flows t o these th ree compartments from XFLALG ( 5 ) are reproduct ion, and f lows back f rom them are rec ru i tmen t from s e t t l i n g and growth. The t h i r d loop type i s t h a t diagramming C02 uptake from XDISOL (80) dur ing photosynthesis, and i t s r e t u r n du r ing r e s p i r a t i o n . Outputs t o XDON (67) and XDOP (69) are excre t ion , as i s i n p a r t t h e flow t o XDOC (62). F loa t i ng macroalgae, XDTPLT (8) break o f f from XFLALG ( 5 ) and cont inue l i v i n g ; d e t r i t u s (XDETR3 [651 and XOETR4 1661 ) i s s i m i l a r l y derived. XFLALG (5) a l so furn ishes carbon to XBROWS ( 3 6 ) , XGRAZV (48) and XBROWV (49 ) , because these animals e a t a lgae. The dot ted arrows on t h i s l a s t s e t of flows show t h a t these flows a r e con t ro l l ed i n p a r t by t h e amount of a lga l mater ial a v a i l a b l e , and i n p a r t by t h e biomass o f t h e grazer . S ix more " s i n g l e compartment" models a r e a l s o shown i n a s i m i l a r diagram format (Figures 3-8). Tliese diagrams a l l resemble Figure 2 in t h e i r complexity and mode of working. Space does not permit t h e inc lus ion of diagrams f o r a l l 104 compartments, although these were developed during t h e two-week workshop.* Connectivity matr ix While feedback dynamics diagrams a r e a convenient method f o r showing individual compartments i n d e t a i l , they cannot be combined f o r a model of this s c a l e without becoming unmanageable mazes. An a l t e r n a t i v e graphical r ep resen ta t ion t h a t has d e s i r a b l e proper t ies i s the connect iv i ty matr ix (Figure 9) . This matr ix i s a square binary matrix showing t h e presence o r absence of flows from each compartment to each o the r one. The s i z e o f t h e matrix i s determined by t h e number of compartments (104 i n t h e case of t h e CITRE model ) . Each compartment, from X1 through X104, i s 1 i s t e d both across t h e v e r t i c a l ax i s and down the horizontal ax i s . The d i r e c t i o n of flow i s from compartments on the horizontal t o those on t h e v e r t i c a l ax i s . Flows a r e indica ted by a dot " " loca ted a t the i n t e r s e c t i o n square o f two components o f the matrix. Thus, i n Figure 9 f o r example, t h e s i n g l e d o t on t h e bottom 1 ine ( X O R G C 104 ) i n d i c a t e s a flow from XORGC (104) t o XDOC (62) but not the reverse . The connect iv i ty matrix shows o t h e r information of i n t e r e s t . As i t s name implies , i t i l l u s t r a t e s t h e t o t a l "connect iv i ty" o r percentage of poss ib le i n t e r a c t i o n . The CITRE matrix with 104 compartments has (104)2 o r 10,816 potent ia l i n t e r a c t i o n s . In t h i s coral r ee f ecosystem model t h e r e a r e about 2,000 i n t e r a c t i o n s o r 20% connect iv i ty . Since connect iv i ty i s highly dependent on compartment d e f i n i t i o n and a t t h i s level o f r e so lu t ion concerns only material flows, i t i s not poss ib l e t o d i s t ingu i sh a t this time between p rope r t i e s r e s u l t i n g from t h e modeling approach, and those inhe ren t i n t h e ecosystem. Complex ecosystems may be charac ter ized by a much .higher percentage of information "flows" o r non-material i n t e r a c t i o n s . However, these i n t e r a c t i o n s a r e even more dependent than material flows on t h e level o f model reso lu t ion . * Copies of t h e working d r a f t s may be obtained from A. L. Dahl a t Department of Botany, Smi thsonian I n s t i t u t i o n , Washington, D. C. 20560. Some working groups a r e continuing t o develop t h e i r p a r t s of the model. There are two disadvantages to the connectivity matrix: i t does not show external forcing functions (temperatdre, currents, e t c . ) or information processing (which are shown by dashed arrows on the diagrams), and i t does not indicate the magnitude of the flows. A more complex form of matrix, a coeff ic ient matrix, subst i tu tes numerical values fo r the binary indicators of the connectivity matrix. Turnover rates then appear in the principal diagonal ( a l l , a 2 , a33,. . .) 5 o f the matrix, and f lux or t ransfe r rates appear in the of f - ~agonal elements. The CITRE matrix of turnovers and t ransfers i s incomplete and is therefore omitted here. While there i s no immediate prospect o f continuing t o develop the CITRE model in i t s present form, i t should now be possible t o begin to piece together the major elements, and to quantify the essential relationships of the coral reef ecosystem. Only in t h i s way will an overall picture of t h i s most fascinating biological community u1 timately be assembled. Li tera ture Cited Forrester, J . W . 1961. Industrial dynamics. Cambridge, Mass.: MIT Press, 464p. Table 1. List of Model Compartments and Their Characteristics Mnemonic Compartment Number Compartment Name Characteristics -- Benthic Plants (10) Nitrogen-fixing algae XNFIX Probably important in reef nutrient cycling Coral 1 ine crusts Benthic microalgae Turf Macro-a1 gae, fleshy XCRUST Important in forming and cementing reef framework through their calcium carbonate production XBMALG Primary unicellular forms in sediments and surface films XTURF Composed of species such as Pterocladia, Polysiphonia, and Cladophora less than 2 cm. high XFLALG Laroe fleshv alaae over 2 cm. high, suc6 as ~urbinaha, Sargassum, and Dictyota that may produce signifi- cant macro-habitats Carbonate-producing macroalgae XCBALG Carbonate producing algae other than crusts, such as Halimeda, Rhipocephal us, Udotea , and Penicillus that make a significant -- contribution to carbonate sediments. Plankton (23) XI1 XI2 Boring algae Detached plants Marine grasses - blades Marine grasses - roots XBORE Includes filamentous and siphonaceous greens such a s Ostreobium, t ha t occur within the carbonate matrix of the reef and l iv ing animals. XDTPLT Plants such as Turbinaria,Sargassum, Thalassia, and Syrinqodium t h a t often remain a l i ve fo r some time a s f loat ing aglomerations, as well as Acanthophora and Laurencia, t h a t sometimes form d r i f t populations on the bottom. XROOT XBLADE The emergent portions of Thalassia, Syringodium, Diplanthera, and Hal ophyea Roots and rhizomes of the seagrasses, as well as the portions of such algae as Halimeda, Udotea, Rhipocephalus, Penicil lus , and Avrainvillea tha t penetrate in to the sediment Heterotrophi c phytoplankton XPHPLl Less than 10 u Autotrophic phytoplankton XPHPL2 Less than 10 u Table 1. List of Model Compartments and Their Characteristics Mnemonic Compartment Number Compartment Name Characteristics Plankton (23) Xl 3 X14 X15 Xl 6 XI7 XI8 XI9 X20 X2 1 X22 X23 X24 X25 Autotrophic phytoplankton Autotrophic phytoplankton Microholoplanktonic omnivores Mesoholoplanktonic omnivores Macroholoplanktonic omnivores Neuston omnivores Microepi ben thic omnivores Mesoephibenthic omnivores Macroepibenthic omnivores Mesoholoplanktonic ca>nivores Macroholoplanktonic carnivores Neuston carnivores Mesoepibenthic carnivores XPHPL3 XPHPL4 XZOOHl XZOOH2 XZOOH3 XZOON XZOOEl XZOOE2 XZOOE3 XZOCH2 XZOCH3 XZOCN XZOCE2 10-100 u Greater than 100 u Less than 200 u 200-500 u Greater than 500 u All sizes Less than 200 u 200-500 u Greater than 500 u 200-500 u Greater than 500 u All sizes 200-500 u Invertebrates (14) X34 Macroepi benthic carnivores XZOCE3 Greater than 500 u Microholoplanktonic de t r i t u s XZODHl Less than 200 u feeders Mesoholoplanktonic de t r i t u s XZODH2 200-500 u feeders Kacrohol opl anktonic de t r i tus XZODH3 Greater than 500 u feeders Neuston de t r i tus feeders XZODN All s izes Microepibenthic de t r i tus XZODEl Less than 200 u feeders Mesoepibenthic de t r i tus XZODE2 200-500 u feeders Macroepibenthic de t r i tus XZODE3 Greater than 500 u feeders Animal-plant symbionts Invertebrate scrapers Invertebrate browsers XHERM Sedentary or s e s s i l e ; derive a portion of t he i r nutr i t ion from symbiotic algae XSCRAP Motile animals tha t remove sol id substratum along w i t h food XBROWS Motile animals tha t do not remove sol id substratum along w i t h food ,Table 1. Lis t of Model Compartments and Their Characterist ics Compartment Number Compartment Mnemon i c Name Characterist ics Passive suspension feeders XAHERM Sedentary or s e s s i l e ; feed on materials suspended i n the water column, passively col lect ing food brought by ambient water current Active suspension feeders XSPONG Sedentary o r s e s s i l e ; feed on materials suspended i n the water column, act ively create a water current t o bring food through the food gathering apparatus Microbrowsers (meiofauna) XFlEIOl Live on or in the sediment or reef framework and feed by passing sediment through t h e i r gut ( a l l a re l e s s than ? mm. i n smallest diameter) Macro-deposit feeders XDEPOS Feeds by passing sediment through i t s gut (more than 2 mm in smallest diameter) Sedentary micropredators XPREDl Sessi le or sedentary; capture individual prey organisms passing i n the water. Predators on small prey X P R E D 2 Motile animals t ha t capture small invertebrates and/or vertebrates Vertebrates (1 1 ) X48 Predators on medium prey XPRED3 Micropredators (meiofauna) XMEI02 Parasites/pathogens XI PARA Parasi te pickers XPICKI Invertebrate eggs attached XBEGGI t o reef Grazers Browsers Plankton feeders, bottom Motile animals t h a t capture medium- sized invertebrates and/or vertebrates Live within the sediment or i n the reef i n t e r s t i ce s and capture small micro-browsers and/or other micro- predators. Relatively sedentary predator deriving i t s nourishment from one (or very few) prey individuals Motile animals t h a t remove paras i tes from other animals Attached invertebrate eggs XGRAZV Feed by scraping the substra te and in so doing remove a portion of the substra te XBROWV Feed by nipping the substra te b u t do not remove portions of the substra te XPLNKB Feed on epibenthic plankton, regard- l e s s of t h e i r position i n the water column Table 1. L i s t o f Model Compartments and The i r C h a r a c t e r i s t i c s Mnemonic Compartment Number Compartment Name C h a r a c t e r i s t i c s Plankton feeders, midwater XPLNKM Feed on holoplankton neuston regard less o f t h e i r p o s i t i o n i n t he water column Predators, smal l XPREDS Feed on ver tebra tes o r i nve r teb ra tes by methods o the r than browsing o r graz ing. Size l e s s than 50 mm standard l e n g t h Predators, medium XPREDM Feed on ver tebra tes o r i nve r teb ra tes by methods o the r than browsing o r grazing. S ize 51 mm t o 250 mm standard l e n g t h Predators, l a r g e Predators, t o p Pa ras i t e p i cke rs D e t r i t u s feeders XPREDL Feed on ver tebra tes o r i nve r teb ra tes by methods o the r than browsing o r grazing. S ize 251 mm t o 500 mm standard l e n g t h XPREDT Feed on ver tebra tes o r i nve r teb ra tes by methods o t h e r than browsing o r grazing. Size 501 mm and g rea te r XPICKV Feed on ve r teb ra te ectoparas i tes XDETV Feed on d e t r i t u s Attached f i s h eggs NO3 NO2 N"3 Dissolved organic carbon Suspended de t r i t u s Suspended de t r i t u s Suspended de t r i t u s Trapped de t r i t u s Dissolved organic nitrogen Dissolved inorganic PO4 Dissolved organic P Dissolved 02 I n t e r s t i t i a l dissolved NO3 XBEGGV XN03 XN02 XNH3 XDOC XDETRl XDETR2 XDETR3 XDETR4 XDON XP04 XDOP X02 XIN03 Vertebrate eggs which are attached t o the substratum o r supers t ra te , brooded or guarded (i .e . , non- pelagic) Dissolved n i t r a t e Dissolved ni t r i te Dissolved ammonia Smaller than 10 u Greater t h a n 100 u Detri tus t h a t f a l l s t o the bottom or moves by s a l t a t i on along the bottom N N Table 1. Lis t of Model Compartments and Their Characterist ics Comoartment Number Comoartment Mnemonic Name Characterist ics X78 X79 Geology (9) X80 I n t e r s t i t i a l dissolved NO2 I n t e r s t i t i a l dissolved NH4 I n t e r s t i t i a l dissolved organic C I n t e r s t i t i a l par t i cu la te organic C (dead) I n t e r s t i t i a l dissolved organic N I n t e r s t i t i a l dissolved organic P I n t e r s t i t i a l dissolved PO4 I n t e r s t i t i a l dissolved 02 Dissolved inorganic C XI DET X 1 DON XIDOP XDISOL Grams carbon, dissolved in the seawater, in the form of Cop, HC03- and CO3. In motion. Suspended inorganic C. Bedload inorganic C Frame inorganic C Non-framehon-sediment inorganic C Inorganic C in rubble Inorganic C in sand Inorganic C in mud Interstitial dissolved XSUSP XBED XFRAME XORG XRUBBL XSAND XMUD Grams carbon, suspended in the seawater, in the form of fine CaC03. In motion. Grams carbon in the form of coarse CaC03. In motion, on or near the sea floor. Grams carbon in the form of CaC03, representing the rigid framework of the reef Grams carbon in the form of CaC03, tied up in living non-frame organisms Sediment. Grams carbon in the form of CaC03 in the loose sediment. Greater than 4 mm. Sediment. Grams carbon in the form of CaC03 in the loose sediment, 62 u- 4 mm Sediment. Grams carbon in the form of CaC03 in the loose sediment less than 62 u. Grams carbon dissolved in the interstitial waters of the sediment in the form of C02, HC03 and C03. "l U . - e "l .w L w C , U 5 L 5 s U L . r w L : t- u s m V) .v s w E C , L 5 n E 0 U 7 w -0 0 I: LC 0 I-' "l . r 2 , - - w 7 n e "l U .,- I-' "l . - L w I-' U 5 s. 5 s U U . r s E E 5 w z s E C , s '2 w L 5 n E 0 U L w .Q 5 Z 4 2 s w E w L 5 a E 0 U 7 3 s 5 w L a le - = 0 n o "l a 0 E w w s 5 w . - W L r w I-' 3 s E 0 aJ L s % - e N O z w w e > ." 0 E U w u CL 5 a 25 LL z >< " l C , s 5 7 a m 8 . r X .? U- I s w m 0 L C , . r Z c cn X w s e w L Z E $ 0 L L L C , - - O m U -w " l.7 + - E E 5 U " l W " l > L s s w ' -5 w 5 5 " l s w L h 5 0 5 % - .o m m w 3 w L s W " l h O 'r-7 3 L 3 5 s T U V ) L V ) 5 W I - ' w w - s a J o 5 4 J V C L U . r N T I - '= O L O E rn 0 W U 0 O W - -V L n w - 5 - s w r n ~ m " a m w e s w w w w u c " l U 0 - C L - w . r o w + w s . - % - .r- B 5 L s L C s m w C ,W .V W O L 3 m a s 0 U O C W O S O 7 2 .,-. V ) LI: w X r n n C w w m 5 - u r - 5 .- a s o a E 5 3 - 0 7 3 U U C , 0 G . p c "l a s w 5 " la r - 0 s 0 e a e L s V)w m s .r U 0 S U L I :e .,- . - w 3 3 u r v 0 5 3 3 5 a v at!- L 0 V ) U L C " l 7 I;! .r- s 5 " l ? - I;! . r s 5 r - 5 .?- I-' s W V I VI C , VI E aJ s 0 VI E e m s w w E U w 5 7 L W I - ' ==, ' : . - - 0 w L e .?- m I- '= E I-' w VI W I - ' L s 5 w E VI . r 't 22 : V V I L " l 3 E L w 0 0 ) L : 0 a L VI I-' .,-- 0 s Z a . - - 5 L L w w L I-' 5 s w s VI .?- I-' .? E I-' 5, VI m 3 s .F VI o w L s u m w . - - s E .F . - - I-' L . r 5 0 I-' 0 L T I v a V ) a (0 0 0 w L T I aJ E I-' 5 F i g u r e 1 Symbols f o r feedback dynamics diagrams Des ignates a com ar tment , o r a f u n c t i o n a l -7--- group gC m 2). M a t e r i a l f l o w . T h i s would be f l u x e d grams carbon o r n u t r i e n t s ( g C m-2 Flows a r e c o n t r o l l e d by " v a l v e s " . A dashed l i n e i s i n f o r m a t i o n . T h i s symbol i s a d e c i s i o n f u n c t i o n , wh ich serves t o i n t e g r a t e i n f o r m a t i o n abou t i n f l u e n c e s on a f l o w . I n a d d i t i o n t o f l o w s between compart- ments, m a t e r i a l may come f r o m a source, e x t e r n a l t o t h e system o f d e f i n i t i o n , o r be sequestered i n an e x t e r n a l s i n k . An open c i r c l e w i t h a name i n i t i s a v a r i a b l e which comes f r o m o u t s i d e t h e system o f r e f e r e n c e (e.g., s u n l i g h t ) . T h i s symbol i s used i n t h e i l l u s t r a t i o n o f smal l compartment g r o u p i n g s o r submodel s t o show f l o w f r o m a compart- ment w i t h i n t h e system o f d e f i n i t i o n , b u t n o t o f i n t e r e s t i n t h e p r e s e n t d iagram. T h i s symbol shows a f l o w f r o m a compart- ment t o a n o t h e r compartment w i t h i n t h e system o f d e f i n i t i o n b u t n o t o f i n - t e r e s t i n t h e p r e s e n t d iagram. 4' XDTPLT (8) TEMPERATURE . XDOC (62) X02 (70) XDON (67) XDOP (69) Figure 2. Diagram representing the fleshy macro-algal compartment and i t s re la t ion to other compartments. Figure 5. Diagram representing the detritus feeder compartment and i t s relation t o other compartments. I) TEMP XZOCH2 (22) XZOCH3 (23) 2) DlEL CYCLE XZODH2 (28) 3)TURBIDITY XZODH3 (29) 4 ) SHELTER XBROWS (36 ) 5) PARASITES 8 DISEASE XPRED2 (42 ) XPRED3 (43) \ / X I PARA (45) XNFIX( I ) \ y -: XPREDS x M L 1 ~ ~ 1 ( 5 4 3 X BMALG (3) - / XPREDT (55 XZOOH3(17) / 1 XSBlRD (93) XZOCH3(23) XALAND (95) X ZODH3(29) / X N H 3 (61) X DETR2 (64) X DETR3 (65) X DETR4 (66) X DON (67) X P O 4 (68) X DOP (69) X DlSOL (80) X SUSP (81 d V kc / XMElOl (39) I R -- X DETR 4 (66 ) I \-+' I C - XME102(44) ' ? X IDET (75) I XDETV ( 57 ) 2 - - - PLANTS 8 SYMBIONTS - - - X NO3 (59) v 1 - Figure 6 . Diagram representing the dissolved n i t r a t e compartment and i t s re la t ion t o other compartments. TERRESTRIAL NUTRIENTS XNlT (96 ) XPHOS (97) XCAL (98) XPOTAS (99) XMINER (100) X PDECM (92) XADECM (94) Figure 8. Diagram representing the seabird compartment and i t s re la t ion t o other compartments. F i g u r e 9. CITRE p r e l i m i n a r y ecosystem m a t r i x ; r e e f - f l a t submodel. The d o t s i n d i c a t e carbon o r ca rbon-equ iva len t f lows f rom compartments on t h e l e f t t o compartments a l o n g t h e t o p . B lanks i n d i c a t e no c o n n e c t i v i t y , and t h e q u e s t i o n marks i n d i c a t e p o s s i b l e carbon f l o w between v a r i o u s o r g a n i c compounds and b e n t h i c p l a n t s . A COMPARATIVE SURVEY OF CORAL REEF RESEARCH SITES* Arthur L . Dahl, Ian G. Macintyre, and Arnfried Antonius The complexity of coral r e e f environments has long a t t r a c t e d s c i e n t i f i c i n t e r e s t , b u t as y e t few i f any research programs have compared r e e f s from d i f f e r e n t a reas o r oceans with a view t o understand- ing the overal l funct ioning o f a reef ecosystem. Plans f o r such programs recent ly were i n i t i a t e d a t the Smithsonian I n s t i t u t i o n , and i t was found t h a t i n the undertaking of an i n t e r d i s c i p l i n a r y inves t iga t ion of r ee f ecosystems, the f i r s t c r i t i c a l s t e p i s t o choose research s i t e s t h a t can meet s t r i n g e n t s c i e n t i f i c c r i t e r i a . The Smi th son ian ' s comparati ve i nformatiori on potent ia l r ee f research s i t e s in both t h e Caribbean and P a c i f i c i s being presented here in the hope t h a t o t h e r i n v e s t i g a t o r s wi l l f i n d i t useful i n planning fu tu re r ee f s t u d i e s . The data were o r i g i n a l l y compiled as a r epor t on the research s i t e s e l e c t i o n process f o r t h e Smithsonian programs, and t h i s has l a r g e l y determined the p resen t format. The observa t ions a r e pr imari ly qua1 i t a t i v e i n na tu re , present ing a broad view of comparative reef s t r u c t u r e and composition. Thei r value l i e s i n t h e i r common perspec t ive , having been compiled by a c lose ly in t eg ra t ed s i t e s e l e c t i o n team whose members, within a s h o r t t ime, v i s i t e d many coral reef a reas throughout the Caribbean and much of the P a c i f i c , applying common c r i t e r i a to achieve a common goal . The s i t e search was conducted f o r two programs of the Smithsonian I n s t i t u t i o n . The f i r s t , Inves t iga t ions of Marine Shallow Water Ecosystems (IMSWE), was organized within t h e National Museum of Natural History with t h e support of the Smi thsonian Environmental Sciences program t o conduct an in t eg ra t ed b io logica l and geological ana lys i s of coral reef communities. The second was t o be a much l a r g e r , mu1 t i - i n s t i t u t i o n a l - . program, the Comparative Inves t iga t ions o f Tropical P.eef Ecosystems (CITRE). d e v e l o ~ e d under the ausoices o f t h e Smithsonian Off ice of Environmental sciences with a p lenning g r a n t from the In t e rna t iona l Decade of Ocean Exploration Off ice a t the National Science Foundation. The plans f o r the CITRE program included t h e development of a complete systems ana lys i s model of t h e r e e f ecosystem based on energy and material flows through various reef components. For both programs, t h e s c i e n t i f i c advantages and g r e a t e r research po ten t i a l o f a s c i e n t i f i c a l l y " i d e a l " reef s i t e were given f i r s t p r i o r i t y , with purely l o g i s t i c a l cons idera t ions * Xnvestigatjons of Marine Shallow Water Ecosystems (IMSWEI con t r ibu t ion No. 3 ; supported i n p a r t by t h e Comparative rnves t iga t ions of Tropical Reef Ecosystems p ro jec t under NSF Grant No. G X - 2 8 6 7 6 a s p a r t of the In terna t ional Decade of Ocean Exploration program. cons ide red s e c o n d a r i l y , and thus a f r e s h l o o k was t a k e n a t many p o t e n t i a l r e e f r e s e a r c h areas w i t h t h e c o n t e x t o f t h e programs i n mind. An " i d e a l " r e e f was cons ide red t o b e one t h a t i s s u b j e c t t o l i t t l e e x t e r n a l s t r e s s , and t h a t i s c h a r a c t e r i z e d by as many o f t h e s c i e n t i f i c c r i t e r i a as p o s s i b l e . S ince t h e c r i t e r i a by which s i t e s were judged have a c r u c i a l b e a r i n g on t h e c o n c l u s i o n s o f t h e s i t e a n a l y s i s , t h e y a r e o u t l i n e d below i n some d e t a i l : Ample development o f a l l c h a r a c t e r i s t i c r e e f zones, f r o m t h e " reef f l a t " t o deep w a t e r communit ies i n depths o f 50 t o 100m. A s teep f o r e - s l ope, t o f a c i 1 i t a t e f i e l d o b s e r v a t i o n s by t e l e s c o p i n g t h e zonat ion. Vigorous r e e f growth where i n t e r a c t i o n s w i t h t h e su r round ing w a t e r mass can be measured and w i t h a good g e o l o g i c a l r e c o r d o f p a s t development. U n i d i r e c t i o n a l c u r r e n t f l o w f o r p e r i o d s l o n g enough t o p e r m i t c ross - r e e f m e t a b o l i c s t u d i e s o f t h e t y p e s u c c e s s f u l l y used a t Eniwetok (Johannes -- e t a l . 1972). No o v e r r i d i n g un ique c h a r a c t e r i s t i c s w i t h r e s p e c t t o t h e r e e f s i n t h e same area. Freedom f r o m m a j o r t e r r e s t r i a l , human, o r n a t u r a l c a t a s t r o p h i c i n f l u e n c e s i n t h e p r e s e n t o r r e c e n t pas t . Some p r a c t i c a l c r i t e r i a c o u l d n o t be ignored : 1. S u f f i c i e n t a c c e s s i b i l i t y t o meet t h e needs o f a l a r g e program. 2. A h a r b o r and some accommodations and f a c i l i t i e s , o r t h e p o s s i b i l i t y o f deve lop ing these a t reasonab le c o s t . 3. A v a i l a b i l i t y o f r e s e a r c h vesse l suppor t . 4. P r o b a b i l i t y o f a s t a b l e government w i t h a f a v o r a b l e a t t i t u d e towards t h e program. 5. S u i t a b i l i t y f o r r e s e a r c h needs such as mu1 t i p l e sampl i ng , m o n i t o r i n g w i t h shore-based i n s t r u m e n t a t i o n , d r i l l i n g f o r g e o l o g i c a l samples, e tc . S ince t h e p u b l i s h e d l i t e r a t u r e has o n l y s c a t t e r e d i n f o r m a t i o n on inany o f these f e a t u r e s , a q u e s t i o n n a i r e ( F i g . 1 ) was developed and s e n t t o numerous s c i e n t i s t s w i t h f i e l d exper ience i n t h e Car ibbean and t h e t r o p i c a l P a c i f i c . The response was g e n e r a l l y good, p a r t i c u l a r l y f o r t h e Caribbean, and p r o v i d e d many suggest ions f o r p o t e n t i a l c o r a l r e e f research s i t e s . F o l l o w i n g p r e l i m i n a r y e v a l u a t i o n o f t h e q u e s t i o n n a i r e s , survey teams were s e n t t o t h e most p r o m i s i n g r e e f a reas t o i n v e s t i g a t e the i r appropriateness in terms of the established c r i t e r i a . The standardized reports of the survey teams, based primarily on the i r f i r s t hand observations, make u p the body of the present paper. For the sake of brevity, no attempt has been made to c i t e the many ea r l i e r published reports on the areas described. Reef descriptions were compiled from f i e l d notes and photographs. A t l e a s t one, and generally two or more of the authors participated on a l l survey teams i n order to provide a basis for valid comparative judgements. In a l l instances the surveys focused on the s e t c r i t e r i a although additional de ta i l s included in the reports may r e f l ec t the special i n t e r e s t s of individual team members rather than the unusual prominence of a particular group. The conclusions in terms of s i t e preference, were conditioned by the c r i t e r i a for the proposed ecosystem programs, and would d i f fe r i f other factors were to be included. Charges for f a c i l i t i e s or transportation, where given, are those a t the time of the surveys (generally 1971) and are included only where such information i s d i f f i c u l t to obtain to give a rough measure of p rac t ica l i ty . For convenience and c l a r i t y , and Caribbean and Pacific s i t e s are treated separately. The Pacific survey was inevitably l e s s thorough, and many other areas warrant detailed examination. The Indian Ocean and more d i s tan t Pacific areas such as Australia were excluded a s being impractical for a U.S.-based program. CARIBBEAN AREA Until recently, Caribbean coral reefs were generally considered infer ior to t he i r Indo-Pacific counterparts, mainly because previous studies concentrated on the southern Florida and northern Bahama area (e.g. A . G . Mayor; T . W . Vaughan), which i s noted for i t s marginal coral-reef development. Recent investigations, however, indicate tha t t h i s ea r l i e r assessment of Caribbean reefs i s not al together accurate. A1 though reef growth in the Caribbean i s accomplished by considerably fewer coral species than in the Indo-Pacific, the reef construction processes are comparable. In other words, the same variety of reef types occur i n both oceans, as well as an overall s imilar i ty in zonation w i t h depth. Data on potential research s i t e s were compiled for the Caribbean Sea and areas to the north. Regions of coral growth i n the southwestern Atlantic, south of the barren area of heavy sedimentation off the Orinoco and Amazon r ivers , were excluded from consideration owing t o log is t ica l problems and lack of s c i en t i f i c jus t i f ica t ion (see Laborel, 1967). Three sources of data were used: (a) the l i t e r a t u r e ; (b) standard questionnaires sent to experienced workers in the Caribbean region; and (c) report from the site-survey teams. Information from the questionnaires i s condensed i n Table 1 . From accumulated information and the personal r epor t s of program members, a number of s i t e s were se lec ted f o r d e t a i l e d f i e l d examination, including Acklins Is land (Bahamas), S t . Croix (U.S. Virgin I s l a n d s ) , Discovery Bay (Jamaica), Glover ' s Reef ( B r i t i s h Honduras) and the San Blas I s lands (Panama). The data in Table 1 incorpora te t h e conclusions of the survey team r e p o r t s , which follow. Acklins I s l and , Bahamas* The r e e f s o f f the north and e a s t coas t of Acklins Is land ( ~ 2 ~ 3 0 ' ~ 7 4 ? 0 0 ' ~ ) i n the eas t e rn Caribbean were surveyed during two v i s i t s ; t h e f i r s t on Oct. 20-24, 1970 by a two-man team, and the second from April 13-23, 1971 by an eight-man team. Acklins Is land i s p a r t of the Crooked Is land D i s t r i c t of the Bahamas, b u t i n f a c t i s separated from Crooked Is land by "The Going Through", a t i d a l channel (Fig. 2 ) . The nea res t populated cen te r i s Nassau, and from t h e r e access t o Ackl i n s i s by small plane o r boat. An old a i r p o r t a t P inef ie ld Point wil l soon be replaced by t h e new one under construc- t i o n a t Pompey Bay. A t p r e sen t , c h a r t e r a i r l i n e s land on a road near t h i s bay. Apart from t h i s s e r v i c e , two weekly f l i g h t s a r e ava i l ab le from Nassau t o Crooked Is land . The t r i p t o Acklins i s completed by t a x i and f e r r y (small outboard) . A 27 m vessel based i n Chester on northern Acklins i s used pr imar i ly f o r shipping and f e r ry ing from Nassau t o Chester , b u t would be a v a i l a b l e f o r c h a r t e r a t $100/day, and i t could f e a s i b l y be considered f o r a f l o a t i n g hotel o r f e r r y f o r r ee f diving. Small boats a r e r e a d i l y a v a i l a b l e , but outboards a r e not . The population of Acklins i s sparse and s c a t t e r e d along the western and northern coas t s . Only a few se t t lements can be ca l l ed communities. The c o s t of l i v i n g i s very high because near ly a l l commodities a r e imported. Local f a c i l i t i e s f o r v i s i t o r s a r e few and a l s o expensive (e .g . a small co t tage with ba res t e s s e n t i a l s c o s t s $ll(US)/day/person, and a half- ton t ruck $24/day with d r i v e r ) . A research team would probably have t o bring "bed and board" with i t . The i s l and i s composed of P le is tocene c a l c a r e n i t e , apparent ly predominantly o o l i t i c ma te r i a l . I t s topography i s r e l a t i v e l y f l a t b u t a s e r i e s of s t r i k i n g " f o s s i l dune r idges" (up t o 30 m r e l i e f ) p a r a l l e l t h e present c o a s t l i n e . The water t a b l e i s c lose t o the su r face (2-3 m) so t h a t f r e s h water i s r e a d i l y ava i l ab le . The s o i l i s very th in except in some depressions. Most of t h e vegetat ion i s a heavy growth of low brush, genera l ly about s i x t o t en f e e t high, b u t t h e r e a r e a few sca t t e red t r e e s (coconut, mahogany, casuarina and tamar ind) . Mangrove swamps a r e well developed on the northern end of the Bight of Ackl i n s . xField survey: October 20-24, 1970, by W . H . Adey, and I . G . Macintyre. Second f i e l d survey: April 13-23, 1971, by W . H . Adey, A . Antonius, A. L. Dahl, P. M. Kier, I . G . Macintyre, M . E . Rice, and T . R . Waller. F i r s t V i s i t : October 20-24, 1970 The e a s t e r n and n o r t h e r n areas o f A c k l i n s were i n v e s t i g a t e d as w e l l as t h e c e n t r a l l agoon area ( B i g h t o f Ackl i n s ) . Heavy swe l l s c r e a t e d rough weather c o n d i t i o n s and t u r b i d i t y f o r two days, a c o n d i t i o n common i n t h e area f rom October t o A p r i l . The f o l l o w i n g s i t e s were examined: 1 . West s i d e o f Atwood Harbour, n o r t h c o a s t o f A c k l i n s I s l a n d . The p a t c h r e e f here i s no more t h a n 15 m f r o m shore, and i t abounds w i t h f i s h ; a hawk's b i l l t u r t l e a l s o was observed i n t h e area. Much o f t h e r e e f s u r f a c e i s covered w i t h c r u s t o s e c o r a l l i n e a laae , e s ~ e c i a l l v - - o f t h e genera P o r o l i t h o n and N e o g o n i o l i t h o n b u t l a r g e growths 0 f ' ~ o n t a G r e a a n n u l a r i s , Acropora pa lmata, M i l l e p o r a , P o r i t e s a s t r e o i d e s , and P o r i t e s p o r i t e s were a l s o p r e s e n t . The sandy bot tom of t h e harbor , where examined, had we l l -deve loped T h a l a s s i a beds w i t h l a r g e heads o f N e o g o n i o l i t h o n s t r i c t u m . I n t h e southwest where t h e c u r r e n t was s t r o n g o n l y a t h i n sediment c o v e r was p r e s e n t o v e r t h e r o c k ledge and smal l heads o f S i d e r a s t r e a r a d i a n s and Fav ia f ragum occur red . 2. Pa tch r e e f d i r e c t l y shoreward o f bank b a r r i e r r e e f o f f e a s t c o a s t o f A c k l i n s I s l a n d , o p p o s i t e Golden Grove. The r e e f he re i s about one and a h a l f k i l o m e t e r s o f f s h o r e . The abundant p a t c h r e e f s i n t h i s area a r e surrounded by a sandy bot tom and have abou t 6 m of r e l i e f . A v e r y h i g h percentage o f t h e r e e f s u r f a c e i s . . covered by Poro l i thon, ~ e o g o n i o l i t h o n and o t h e r - c o r a l 1 i nes . No l a r g e caverns were observed as d e s c r i b e d e lsewhere on t h e Bahama Banks ( S t o r r , 1964), b u t t h e r e were "overhangsN here and t h e r e . Large c o l o n i e s o f Acropora pa lmata and M i l l e p o r a were observed a l o n g w i t h abundant b u t smal l c o l o n i e s o f A g a r i c i a a g a r i c i t e s . A d d i t i o n a l common c o r a l s i n c l u d e t h r e e spec ies o f D i p l o r i a , D ichocoenia s t o k e s i i , P o r i t e s a s t r e o i d e s , and Montast rea a n n u l a r i s . A g r e a t abundance o f a l c y o n a r i a n s was a l s o no ted . 3. I n n e r edge o f bank b a r r i e r r e e f , n o r t h e a s t c o a s t o f A c k l i n s I s l a n d o f f P i n e f i e l d P o i n t . Going seaward f rom t h e p a t c h r e e f t h e r e i s a zone o f abundant Montast rea a n n u l a r i s and D i p l o r i a c o r a l heads l e a d i n g t o t h e t u r b u l e n t i n n e r edqe o f t h e r e e f f l a t , where t h e r e a r e l a r s e o v e r t u r n e d q rowths o f Acropora-palmata (some w e r e - l i v e g rowths ) and laFge growths o f " ~ i 1 l e p o r a . Rock s u r f a c e s and c o r a l d e b r i s a r e h e a v i l y coa ted w i t h c o r a l l i n e a l g a e . F i s h were abundant, i n c l u d i n g a number o f d i f f e r e n t t ypes o f p a r r o t f i s h , some q u i t e l a r g e , wh ich seem t o i n h a b i t an open network o f sma l l t u n n e l s under t h e c o r a l - r o c k f l o o r . 4. O f f S p r i n g P o i n t on t h e west c o a s t o f A c k l i n s I s l a n d i n A c k l i n s B i g h t . Fo r a d i s t a n c e o f abou t 150 m f rom shore a v e r y t h i n l a y e r o f sediment c o v e r s t h e smooth r o c k pavement. R i c h a l g a l growths o f Halimeda, P e n i c i l l u s , Batophora and A c e t a b u l a r i a were observed, a l o n g w i t h two spec ies o f c o r a l s : S i d e r a s t r e a r a d i a n s and F l a v i a fragum. A few c o l o n i e s o f Man ic ina a r e o l a t a were a l s o noted. S i t e Colombia I s l a s del Rosario TABLE I : Quest ionnaire Response - Caribbean Area Reef S t ruc tures* Undisturbed Accessible Santa Marta Curacao Venezuela Los Roques Cubagua, Coche, Margari t a , Los Frayles Is1 a s Lesser Anti1 l e s Dominica Puerto Rico Mona Is land Virgin I s l ands S t . Croix Hispaniola S t . Domingo area Jamaica Discovery Bay Bar r i e r Reef and Cays, Pedro Cays, Pedro Bank Banner ~ e e f Grand Cavman - Bahamas Ackl ins Is land Andros Is land Bimini Bermuda Mexico Xslas de Lobos Local F a c i l i t i e s - t + 0 - - - t 0 + - 0 0 0 t t - P o l i t i c a l S t a tus+ S i t e B r i t i s h Honduras B a r r i e r Reef Turneffe I s l a n d Lighthouse Reef G lover 's Reef Colombian I s l ands (o f f Nicaragua) Serrana Bank Roncador Bank Old Providence I s l a n d S t . Andrews I s l a n d Courtown Cays A1 buquerque Cays Great Corn I s l a n d Panama Holandes Cays, San B las I s l ands Reef S t ruc tu res Undisturbed Accessib le + = f avo rab le o r present 0 = unknown o r poss ib le + - = v a r i a b l e - = unfavorable o r imprac t i ca l * see s e l e c t i o n c r i t e r i a above. D. + b a s i c a l l y t he a n t i c i p a t e d go;ebnment a t t i t u d e towards a l a r g e U.S. funded program P o l i t i c a l S ta tus + t + + - - - - - - - + Quest ionna i res and equ i va len t in fo rmat ion on Caribbean s i t e s were supp l ied by the f o l l o w i n g : W. H. Pdey, A. AnWnius, R. W. Bauer, A. L. Dahl, R . F. D i l l , J. Ge is te r , P. W. Glynn, L. S. Land, J. C. Lang, I. G. Macintyre, J. D. M i l l iman, J. L. Munro, R. P f a f f , J. K. Rigby, and K. Ruetz le r . 5. Atwood Harbor, North Coast o f Ack l i ns I s l and (see F ig . 3 ) Back Reef This area i s character ized by smal l patch ree fs sca t te red on a rock p l a t f o r m which genera l l y has a t h i n sand cover. These protected patch r e e f s have a r e l a t i v e l y dense co ra l growth cons i s t i ng p r i m a r i l y o f Acropora palmata, Montastrea annu la r i s , Po r i t es ast reoides, Agar ic ia aga r i c i t es , Dichocoenia s t o k e s i i , Dichocoenia s t e l l a r i s and Por i t es p o r i tes . Reef Crest co ra l composit ion on the l e e s ide o f the r e e f c r e s t i s almost i d e n t i c a l t o t h a t o f the patch ree fs . However, the remainder o f the r e e f c r e s t i s dominantly composed o f Acropora palmata and Montastrea annu lar is which form an open framework marked by the common occurrence o f dead overturned co ra l communities. Upper Fore-Reef Slope The dominant charac teP is t i c o f t h i s area o f the r e e f i s a g e n t l y d ipp ing rock pavement. The two d i s t i n c t zones observed are a) barren zone 1.5-6 m and b) grooved zone 6-18 m. Barren Zone: Predominantly a knobby rock bottom w i t h a t h i n cover- i n g ( 2 cm) o f benth ic algae and f i n e t o coarse carbonate sand. Crustose cora l 1 ines genera l l y are t h r i v i n g under t h i s a lga l -sediment cover. Small co ra l co lon ies sca t te red over t h e rock surface inc lude D i p l o r i a s t r igosa, Siderastrea siderea, Siderastrea radians, Dichocoenia stokesi i , Montastrea cavernosa, Montastrea annu la r i s, Agar ic ia a q a r i c i tes, Po r i t es ast reoides, Po r i t es p o r i tes, Co lphy l l i a natans, I s o p h y l l i a sinuosa, Meandrina meandrites. Sargassum sp., gorgonians, and M i l l epo ra sp. increase no t i ceab ly i n depths l ess than 4 m and l a r g e patches o f bor ing sponges are common. Scattered shal low (1 m) depressions i n the rock pavement have a w e l l - r i p p l e d coarse sand o r rubb le bottom. A few Diadema sp. occur under ledges i n t h e w a l l s o f these depressions. Grooved Zone: The most s t r i k i n g feature o f t h i s depth zone i s the grooves which are c l e a r l y v i s i b l e i n a e r i a l photographs. O f f Atwood Harbour they are genera l l y l ess than 30 m apar t and are commonly about 60 m long. The gen t l y d ipp ing rock pavement t ransected by these grooves i s encrusted w i t h a v a r i e t y o f t r o p i c a l - r e e f cora ls , bu t the colonies are genera l l y smal l , well-spaced and the re fo re do no t form an i n t e r - l ock ing framework. Dominant c o r a l s i nc lude the f o l l o w i n g species: D i p l o r i a s t r igosa, Po r i t es ast reoides, Siderastrea siderea, Dichocoenia s t o k e s i i , Dichocoenia s t e l l a r i s , Montastrea cavernosa, Montastrea annular is , Favia fraqum, Meandrina meandri tes, D i p l o r i a l a b y r i n t h i f o r m i s, Co lpophy l l ia natans, Ste hanocoenia michel i n i i . Gorgonians and sponges + are a l s o abundant i n t h i s dept zone. The groove system begins i n a general depth range o f 5 t o 10 m as an i n d i s t i n c t l a b y r i n t h o f channels between the sca t te red co ra l growth. I n our l i m i t e d observat ions we d i d n o t f i n d grooves o r i g i n a t i n g as "spoon-shaped i n c i s i o n s " o r "eros ional p i t s , " as repor ted f o r s i m i l a r features o f f Andros I s land (Newel1 e t a l . , 1951, p. 25) and o f f Saipan I s land (Cloud, 1959, p. 406). These i n d i s t i n c t channels coalesce t o form a we l l -def ined u-shaped groove which i s r e l a t i v e l y s t r a i g h t and has a smooth sediment-free sur face covered w i t h a t h i n a l g a l growth and cora ls here and there. These u-shaped grooves, which a r e about 10 t o 15 m long, widen from 15 cm wide and 15 cm deep t o 1 m wide and 1 m deep where they open ou t i n - t o a s t i l l deeper and wider sed imen t - f i l l ed groove. I n some .places, up t o th ree separate u-shaped grooves were noted t o be feeding i n t o the same broad sediment-f i 11 ed one. The sed imen t - f i l l ed sec t ions are genera l l y about 50 t o 60 m long w i t h a maximum w id th o f about 10 m. The lower sed imen t - f i l l ed sect ions tend t o taper i n t o a ser ies o f lobes a t the ou te r l i m i t o f the grooves. The thickness o f accumulated sediment i n the troughs was n o t es tab l i sh - ed; however, a t the head o f t h e sed imen t - f i l l ed sect ions the wa l l s were 2 t o 2.5 m h igh and gradua l ly diminished i n r e l i e f t o about 1 m a t the terminus. It i s i n t e r e s t i n g t o note t h a t the grooves i nves t i ga ted were completely enclosed w i t h no ready o u t l e t i n t o deeper water f o r t h e sediments. At the head o f the sed imen t - f i l l ed sect ions the re i s a band 5 m wide o f cora l rubb le which grades down-slope i n t o a w e l l - r i p p l e d sand (wave length, 60 t o 150 cm, ampl i tude, 12 t o 15 cm). This i s an ex- t remely coarse t o medium-grained ske le ta l ca l ca ren i te w i t h gravel s i z e ma te r ia l c o l l e c t i n g i n the r i p p l e troughs. Texture gradat ion o f the sediments cont inues down the groove so t h a t about half-way down, t h e gravel i s absent and there i s s imply a poor ly sorted, extremely coarse t o medium-grained ske le ta l ca l ca ren i te , which i n t u r n grades i n t o a we1 1-sorted medium t o coarse-grained ske le ta l ca lcaren i t e a t the terminus o f t h e groove. Without subsurface data i t i s d i f f i c u l t t o e s t a b l i s h whether these l i n e a r features have been formed by eros ion o r d i f f e r e n t i a l accumula- t i o n o f organic l imestone. Newel1 e t . a l . (1951) found t h a t grooves o f f Long Cay i n t h e Bahamas c u t i n t o o o l i t e bedrock, which c l e a r l y i n - d ica ted an eros ional o r i g i n . Lower Fore-Reef Slope This area s t a r t s a t depths ranging from 12 t o 18 m, where t h e s lope o f the sea f l o o r increases markedly, and cont inues t o t h e depth l i m i t s o f our observat ions, approximately 40 m. Large areas o f well-developed and d iverse deep-water co ra l commu- n i t i e s are predominant i n the upper sec t ion o f t h i s area. Sediment chutes t h a t t raverse the co ra l communities here and there coalesce a t depths of 20 t o 25 m t o form an extensive sand f l a t . Below t h i s sand f l a t the coralsdo not show any appreciable change in species composition b u t tend t o construct large coral mounds covering areas of u p t o 220 sq. meters with approximately 3 m of r e l i e f above the surrounding sand slope. The dominant corals of the lower fore-reef slope consist of the following species, in order of t h e i r abundance: Montastrea annularis , Agaricia aga r i c i t e s , Montastrea cavernosa, Mussa angulosa, Diploria labyrinthiformis, Meandrina meandrites, Eusmilia f a s t i g i a t a , Mycetophyl l i a lamarckiana, Col pophyll i a natans, Porites as t reoides , Stephanocoenia michelinii , Dichocoenia s tokes i i , Dichocoenia s t e l l a r i s , Porites por i tes , Isophyll i a sinuosa, and Diploria s t r igosa. East Coast of Acklins Island 6. Off Pinefield Point The area observed begins a t a depth of 20 m where there i s a d i s t i nc t sand f l a t . Coral heads a re sparse and widely scat tered, b u t gorgonians and benthic algae are very common. Below a depth of 20 m the slope of the sea f loor increases and coral growths tend t o be res t r ic ted t o l i nea r ridges which a re separated by sand and gravel-fi l led grooves generally 3 m wide and 1 m deep. The coral assemblage i s similar t o t h a t of the deep-water coral communities noted on the lower fore-reef slope off Atwood Harbour. Corals give way t o a sand and gravel slope a t depths of 35 t o 40 m. 7. Misconception Rock Misconception Rock consists of a small (100 sq. m), re la t ive ly f l a t rock pavement (well-sorted f i ne t o medium calcareni te) i n the reef t r a c t tha t i s exposed a t low t ide . I t i s approximately 1 kilometer north of Creek Point off the eas t coast of Acklins Island. The rock surface i s heavily pi t ted and etched, the charac te r i s t i c erosional form of limestones i n the splash zone (Newel1 e t . a l . , 1951), and the outer edges are covered w i t h a heavy algal growth. Many large gastropods (Livona pica), Diadema sp., and chitons were observed in numerous shallow depressions. As found i n other areas of the Bahamas (Newel1 e t . a l . , 1951) the bank barr ier reefs off eastern Acklins Island appear t o be a Holocene reef veneer over pre-existing sand ridges t h a t are exposed above sea level in places. 8. South of Golden Grove The broad (12 m) platform which i s a charac te r i s t i c feature of the insular shelf off the eas t coast of Acklins Island separates the reef c res t and fore-slope communities by distances of over 1.5'km. The bottom i s a smooth rock pavement which has a t h i n (1 cm) sedi- ment-algal cover, consisting of a dense benthic algal layer with trapped sand- t o s i l t - s ized sediment. Small coral mounds, having r e l i e f t o l e s s than 1 m are widely spaced on the rock pavement. Coral heads, f o r the most part dead, are scattered over t h i s rock surface. Dominant l i ve corals a re Oiploria s t r igosa , Eusmilia f a s t i g i a t a , Agaricia agarici t e s and Oichocoenia s tokes i i . Rich epifaunal and infaunal assemblages occur under the l i ve and dead coral material , including the bivalve Barbatia domingensis, the gastropod, Tegula fasc ia ta , and the echinoids, Eucidaris tribuloides, Arbacia punctulata and Echinoneus cyclostomus. Gorgonians are common and some are encrusted by Millepora sp. Summary Atwood Harbour would be an excellent location f o r a base s ta t ion as i t i s a good harbour fo r boats, and the Bahamian government i s will ing t o cooperate with s c i e n t i s t s working in t h e i r t e r r i t o ry . The northeast corner of Acklins Island, eas t of Atwood Harbour and North of Pinefield Point, i s Crown Land and arrangements could probably be made t o s e t t h i s area aside as a research s i t e . The reefs have been rela- t i ve ly unaffected by human ac t i v i t y . Fishing appears t o be a minor occupation carried out i n a primitive fashion on an occasional basis. When the new a i r s t r i p i s completed, access ib i l i ty t o Acklins Island will be improved with regular weekly f l i g h t s scheduled fo r the near future. Bi-weekly f l i g h t s t o Crooked Island are currently avai lable from Nassau. A number of vacated small houses on the island might be used as accommodation f a c i l i t i e s . They might be rented a t reasonable r a t e s , b u t i t should be recognized tha t amenities are lacking. Apart from some of the small patch reefs , the reefs on the insular she1 f a r e not we1 1 developed. The exposure of calcareni t e in the reef t r a c t off the eas t coast (Misconception Rock) indicates t ha t many of the reefs may be veneers over Pleistocene sediment ridges. The shallow reef c res t s a re commonly separated from the fore-reef slope coral communities by a very broad rock platform. Rough seas common during the months of October through April would prevent work on the reef c r e s t and fore-reef slope during these months. St . Croix, U.S. Virgin Islands* St . Croix (17"45'N, 64"401W) i s the la rges t of the U.S. Virgin Islands, having a surface area of 22 sq. km (45 km long and u p t o 11 km wide; see Fig. 4). I t s population i s about 35,000, and apar t from *Fie d su vey: January 26-28, 1971, by I . G . Macintyre, ass i s ted by H. &. ~ u i t e r . tour ism, t h e i s l a n d supports a few o the r i n d u s t r i e s such as baux i te r e f i n i n g , o i l r e f i n i n g , and the assembly o f w r i st-watches. This i s l a n d i s r e a d i l y accessib le as i t i s served by several major a i r 1 ines. Transportat ion on the i s l a n d i s ava i l ab le through car r e n t a l a t about $12/day o r scooters a t $7/day. Boats may be chartered, o r rented l ess expensively through t h k F a i r l e i g h Dickinson Laboratory. Hotel accommodations are cos t l y , p a r t i c u l a r l y dur ing the w i n t e r season, bu t the marine l abo ra to ry has some rooms a t reasonable ra tes f o r v i s i t - i n g s c i e n t i s t s . T e r r e s t r i a l Condit ions The topography o f S t . Croix i s " h i l l y " (maximum r e l i e f 355 m, M t . Eagle). The i s l a n d i s composed o f rocks having d i ve rse o r i g i n s and compositions, p r i m a r i l y Upper Cretaceous volcanic sandstones and mud- stones, tu f faceous sandstones, Upper Oligocene m o n t m o r i l l o n i t i c mud- stones, and Lower Miocene a rg i l l aceous and sandy cora l l imestones. Despite the abundance o f vo lcan ic debr is , no volcanoes are present on S t . Croix; however, two small igneous bodies (a d i o r i t e and a gabbro) i n t r u d e the Cretaceous sedimentary rocks. Numerous small d ikes are a l so present i n the h i l l y ranges occur r ing on t h e eastern and northwestern sect ions of the i s l a n d (Whetten, 1966). S t . Cro ix Reefs Although S t . Cro ix has a v a r i e t y o f r e e f types t h a t would be exce l l en t f o r s c i e n t i f i c study, t h e r e appears t o be no evidence o f a r e e f marg i i~a l t o deep oceanic waters, a ser ious drawback t o the se lec t - i o n o f S t . Cro ix as a pr imary s i t e . 1. Bank B a r r i e r Reefs. As expected, the best developed ree fs occur off the eastern o r windward coast, and the most s t r i k i n g developments a r e the bank b a r r i e r ree fs on the landward edge o f the i n s u l a r she l f ; these reefs p a r a l l e l the nor theastern and southeastern coasts f o r distances o f over 5 km. The bank b a r r i e r r e e f o f f Tague Bay (F ig. 2, S i t e 1 ) has about 5 m r e l i e f o f f the shoreward edge and i s approximately 90 m wide. Th is r e e f i s dominated by Acropora palmata, bu t an i n d i s t i n c t zonat ion was noted around the i nne r edge o f the ree f , where t h e base i s charac ter ized main ly by Montastrea annular is ; the edge by Acropora almata and l a r g e mounds o f Po r i t es p o r i m t o 2.5 m h i g h and 4.5 2- and the t o p by Acropora palmata, D i p l o r i a sp., and M i l l epo ra sp. The seaward edge o f t h i s r e e f was n o t i nves t i ga ted dur ing t h i s b r i e f v i s i t ; however the t ransec t (F ig. 6 ) based on in fo rmat ion received from D r . Mu l te r i nd i ca tes t h a t t h i s seaward slope drops o f f t o a depth o f 14 m and gives a graphic representa t ion o f the dominant co ra l d i s t r i b u t i o n across t h e ree f . 2. A lga l Cup Reefs There was n o t enough t ime t o i n v e s t i g a t e the f r i n g e ree fs common around most o f the promontories along the nor theastern coast o f St. Croix. However, the nearshore "a lga l cup r e e f s " occu r r i ng around Cottongarden Po in t were i nves t i ga ted (F ig. 5, S i t e 2). These ree fs , which are charac ter ized by pronounced r ims and over hangs, have a r e l i e f o f about 2 m. The i r upper surfaces are covered by f l e s h y algae i nc lud ing Sarqassum sp. and a few f l a t co lonies o f Po r i t es ast reoides. Chips from these a l g a l ree fs , broken o f f w i t h g reat d i f f i c u l t y , revealed a dense agglomeration cons i s t i ng main ly of crustose co ra l 1 i n e algae and the d i s t i n c t i v e red encrust ing fo ramin i fe ra , Homotrema sp. An unusual occurrence o f Acropora p r o l i f e r a i n an area o f h igh a g i t a t i o n was noted next t o these a l g a l reefs. Abundant algae are present on the sea f l o o r surrounding the cup ree fs , no tab ly P e n i c i l l u s sp. and Halimeda sp. (sediments are p a r t i c u l a r l y r i c h i n Halimeda- der ived debris). 3. Buck I s land T r a i l A r a p i d survey o f the Buck I s l a n d Nat ional T r a i l (Fig. 5, S i t e 3) i nd i ca ted t h a t t h i s i s a wel l-developed "palmata r e e f . " A1 though separated from shore by a narrow sandy b e l t , i t can be considered a form o f f r i n g e ree f . 4. Deep Patch Reef A deep patch r e e f d i r e c t l y eas t o f Buck I s land (Fig. 5, S i t e 4) cons is ts o f an area o f l a rge open frameworks, p r i m a r i l y Acro ora palmata +- This patch r e e f has a r e l i e f o f about 8-9 m above i t s base, w l c h IS covered w i t h l a r g e heads o f Montastrea annular is , D i p l o r i a sp., Montastrea cavernosa, and surrounded by a great abundance o f alcyonarians. M i l l epo ra was found coat ing some o f t h e dead a lcyonar ian branches. 5. Shel f Edge North o f Buck I s l a n d This area was i nves t i ga ted i n o rder t o determine whether a s h e l f - edge r e e f e x i s t s a t t h i s l o c a t i o n (F ig. 5, S i t e 5). The reconnaissance was c a r r i e d out by a towed swimmer, and a l l observat ions were made from the sur face o f t h e water. Although no evidence o f a shelf-edge r e e f was found, an i n t e r e s t i n g t r a n s i t i o n i n bottom c h a r a c t e r i s t i c s was noted: a t depths o f about 30 m a se r ies o f sediment chutes t rave rse a r e l a t i v e l y s teep ly d ipp ing (about 10-20') rock surface t h a t i s covered by alcyonarians, sponges, algae and scat te red co ra l heads. From t h e surface, there appeared t o be l i t t l e r e l i e f between t h e sediment chutes and i n te rven ing rock surfaces. The s p a r s i t y o f co ra l s suggests t h a t r e e f framework cons t ruc t i on was n o t occur r ing a t t h i s depth. Although a spur and groove system appeared to be present a t t h i s location from the a i r survey, firsthand obser- vations showed tha t the "buttress systems" described off Jamaica by Goreau (1959) are not present in t h i s area off St . Croix. Shoreward, the sea f loor gradually grades in to a rocky bottom having sand patches and scattered alcyonarians, sponges, algae and coral heads. Around depths of 15 m , large scattered colonies of Acropora palmata appear, and d i rec t ly shoreward there i s a zone of Millepora. Abundant colonies of Acropora palmata are present in depths of 3-6 m along w i t h other corals such as Montastrea annularis and Diploria sp. 6. Other Reefs off St . Croix During the R/V Eastward cruise, two other reef types were investi- gated in t h i s area: inactive and submerged reefs. The most prominent reef type off St . Cro-ix i s the inactive reef , of which Lang Bank i s the best example. Inactive reefs are reefs t ha t occur within depths of potential vigorous coral growth, but which a re characterized by an absence of reef framework construction. Lang Bank, si tuated a t the shelf edge east (windward) of St. Croix (Fig. 5 ) a t depths of around 9-12 m, i s ideally located for act ive barr ier reef development. From 1 imi ted bottom photographs, however, i t i s evident tha t an interlocking reef framework does not occur on tliis ridge which i s covered w i t h abundant alcyonarians and sponges and large sand patches, but only scattered coral heads. Dredge hauls from t h i s area contained abundant alcyonarians, sponges, and small colonies of Porites por i tes , Porites astreoides, Diploria sp., Agaricia agar ic i tes , Stephanocoenia michel i n i i , Acropora cervicornis, and Millepora sp. Coral debris heavily encrusted by coral 1 ine algae was a1 so abundant. Meyerhoff (1926) suggested tha t the topography of the eastern St. Croix insular shelf i s related t o subaerial erosion of a Tertiary reef complex. Submerged ridges were noted on bathymetric prof i les off the southern coast of St. Croix, established a t depths of about 40 m. These ridges are interpreted t o be submerged reefs t ha t were establish- ed i n relation to a pre-existing lower sea leve l , and tha t occur i n depths greater than are commonly associated w i t h vigorous growth of tropical reef corals. They appear t o be common features off most eastern Caribbean islands (Macintyre, 1972). Pollution Extensive pol 1 ution of the marine environment i s occurring off the southwestern coast of St. Croix, where a garbage dump, bauxite plant and o i l refinery are located (see Fig. 4 ) . I t i s expected that pollution levels will r i s e in the near future because sewage will be piped from Christiansted ( the island cap i t a l ) over t o this s ide of the island. Despite the unfortunate aspects of t h i s circumstance,, i t would offer an opportunity for detai led compari son between pol 1 uted reefs off t h i s coast and the non-polluted reefs t o the east . Such studies might yield valuable information on the degrees of s t r e s s imposed on reefs by pollution, and how well they survive i t . West Indies Laboratory of Fairleigh Dickinson University The West Indies Laboratory i s located on Tague Bay, on the north- eastern coast of S t . Croix, and i t overlooks Buck Island National Park (Fig. 5 ) . This laboratory o f f e r s s c i e n t i f i c f a c i l i t i e s fo r researchers as well as some accommodations, when available. There a re 4 small laboratories equipped with aquaria, and an outdoor aquarium area supplied by a dual saltwater system. The Laboratory a lso owns two 4.9 m ( 16 ' ) Boston whalers and three 5.8 m (19 ' ) Rabollo boats which may be rented by v i s i t ing s c i e n t i s t s a t a minimal cost . Owing to an extensive summer program, v i s i t ing s c i en t i s t s are advised t o plan v i s i t s f o r l e s s crowded periods of the year. An a f f i l i a t e program i s gradually being developed which will permit outside univers i t ies t o have facul ty and research space as well as teaching space privileges on the basis of a yearly charge. Summary Apart from the polluted reefs on the south coast of St . Croix, the coral reefs around t h i s island have been re la t ive ly untouched by man because there i s l i t t l e need t o exploit them f o r food. However, the absence of a reef marginal t o deep water i n the region i s a drawback, as i s the general lack of well-developed reef - f la t zones on the shelf reefs (a d i s t i n c t disadvantage i n d r i l l i ng operations). On the other hand, the numerous advantages of t h i s location make St. Croix worthy of consideration as a research s i t e , par t icular ly fo r a smaller sca le program. The cooperation and available f a c i l i t i e s of W.I.L. would make operational costs on S t . Croix considerably lower than elsewhere in the Caribbean; a t the same time, W.I.L. would probably f a c i l i t a t e detailed and sophist icated experimentation which might be d i f f i c u l t t o undertake and complete a t l e s s well -equipped s i t e s . Following i s a brief out l ine of the merits of St . Croix as a study s i t e : 1. Fac i l i t i e s of the West Indies Laboratory, notably; a. inexpensive accommodations; b. f i s h pens; c. outdoor and indoor aquaria supplied by a l l P.V.C. non- toxic dual saltwater systems and backed u p by independent emergency power uni t ; d . laboratory and o f f i ce space; motor boats; e . reference l i b r a ry with microfilmed periodicals and pr int - out service; f . radiocommunication with f i e l d uni ts ; 9. monitoring service which W.I.L. intends t o es tabl ish off northwestern coast . 2. The labora tory s t a f f would be ab le t o a s s i s t i n serv ic ing f i e l d equipment. Technicians working on Navy p r o j e c t s on western S t . Croix might a l s o be e n l i s t e d t o a id with e l e c t r o n i c s problems. 3. J e t a i r p o r t with d a i l y f l i g h t s t o Washington, Miami, and New York cos t ing l e s s than $200 r e t u r n . San Juan, P . R . , i s 45 minutes away by hourly scheduled f l i g h t s . 4 . A v a r i e t y of reef types a re present , including f r i n g i n g , bank b a r r i e r , i nac t ive , submerged, a lga l cup, pol lu ted and non-polluted r e e f s . 5. Because S t . Croix i s an American t e r r i t o r y t h e r e should be a minimum of red tape involved in having a sec t ion of r ee f t r a c t s e t a s ide f o r s c i e n t i f i c i nves t iga t ion ; ground has a1 ready been broken with the Buck Is land National Underwater Park. 6. There would be l e s s d i f f i c u l t y in shipping equipment t o a U.S. i s l a n d , and f inanc ia l advantages of a duty-free i s l and in the purchase of equipment. Discovery Bay, Norrh Cohrt of Jamaica* Jamaica, an independent member nat ion of the B r i t i s h Commonwealth, i s a l a rge t rop ica l i s l and loca ted i n the northwestern Caribbean Sea a t the extreme northwest t i p of the Nicaraguan Rise. I t i s 140 miles long and 60 miles wide and has mountain peaks i n excess of 7,000 f e e t above sea l e v e l . Fringe r e e f s extend discontinuously along the north coas t and a r e noted f o r t h e i r cora l spec ies d i v e r s i t y (Goreau and Wells, 1967). Discovery Bay ( 1 8 ~ 3 0 ' ~ , 77O25'W), the s i t e of the Discovery Bay Marine Laboratory, i s loca ted about midway along the northern Jamaica c o a s t , and i t l i e s ad jacen t t o a well developed sec t ion of the f r i n g e r ee f (Fig. 7 ) . The r e e f s i n Discovery Bay (Fig . 8 ) gene ra l ly may be divided i n t o the following major zones ( a f t e r Kinsey, 1970; J . Lang, personal communication, 1971 ) . Lagoon Zone: This zone i s about 400 to 500 m wide, and about 1 t o 3 m deep. I t has a submerged sinkhole 25 m deep, but the wal l s and bottom a r e mud covered and there a r e no rock exposures. Loose sand i s the main c o n s t i t u e n t of t h e f l o o r in the lagoon zone, and t h i s precludes development o f s e s s i l e benthic epifauna. Here and the re , bare rock patches and coral heads occur w i t h s ca t t e red gorgonians. Corals and rocks become more abundant seaweed, toward t h e r e a r zone. Rear Zone: The inshore f ace of the reef c r e s t makes u p t h i s zone, which i s about 0.3 t o 2 m deep. Apart from the reef f l a t , t h i s i s the only reef a rea s t rong ly a f f ec t ed by t i d a l f l u c t u a t i o n s and charac ter ized by s trong water movement. Bottom types range from ca lcareous sand, and sand and coral fragments, t o bare coral-rock exposures. Living coral i s *y 1. G . Macintyre, a s s i s t e d by P . Dustran, E . A. Graham, L . S. Land, and J . C . Lang. common b u t never covers the bottom en t i re ly . Richest coral growth occurs in the area abutting the shoreward face of the reef f l a t . Reef Flat: The reef f l a t comprises the area j u s t submerged a t low t ide . I t i s subjected to strong breaking wave action as well as occasional exposure to a i r and rainwater. The main component i s a r e t i cu l a t e ridge-1 i ke s t ructure 5 t o 20 m wide, composed of dead colonies of Acropora palmata, which separates the lagoon from the sea. The unconsolidated skeletons of A . palmata form a l a t t i c e - l i ke s t ructure . Protected areas within the reef a r e locations for a res t r ic ted gorgonian fauna. Channels and pools in the reef f l a t may reach a depth of 2 m . Palmata Zone: This narrow zone, 5 t o 10 m wide, abuts the seaward face of the reef f l a t and extends seaward to a depth of 4 to 6 m . The fu l l force of wave action occurs in t h i s zone where Acropora palmata i s predominant. The bottom consists of coarse sand and sand-scoured coral rock l i t t e r e d with dead and fa l len - A . palmata colonies. Barren Zone: A band of reduced coral d ivers i ty occurs seaward, and i t can be up to 20 m wide, ranging from 5 t o 8 m in depth. The most common organism, Oiadema antillarum, which grazes on the bot tom, may be responsible fo r the scarci ty of s e s s i l e organisms. Isolated colonies of A. palmata occur along with small heads of Diploria s t r igosa, Montastrea annulari s , and Mil 1 epora complanata. Buttress/Mixed Zone: I n the Ocho Rios area studied by Goreau (1959) as well as other areas along the northern coast , the buttress zone merges abruptly with the barren zone. Although the component species remain the same, Discovery Bay d i f f e r s s l i gh t ly in architecture from the other areas in t h a t the normal buttress zone becomes f l a t t e r , with a gradual increase in coral species and coral s ize . This "mixed" zone i s 15 t o 40 m wide, and slopes from 6 to 15 m in depth. Montastrea annularis , one of the dominant corals in the buttress formations, occurs in Discovery Bay in the form of huge rounded masses scattered in f i e l d s of A . cervicornis. Diploria spp. i s also common, along with Oendrogyra cylTndrus, Agaricia agar ic i tes , Porites spp., and Colpophyllia natans. This i s an area of r ich gorgonian divers i ty . Cervicornis Zone: A . cervicornis becomes dominant seaward, and M . annularis reduced t o small patches. This zone, which i s 30 t o 100 m - wide and 20 to 25 m deep, i s characterized by huge mounds of A. cervicornis 15 t o 40 m wide with t h e i r long axis normal to the axis of the reef c r e s t . These reefs (1 to 5 m r e l i e f ) are separated by sand channels 3 t o 50 m wide. Upper surfaces of these reefs arc generally l eve l , and they gradually slope to about 10 m in depth a t t he i r seaward extremity, where they drop abruptly to sand level a t about 20-25 m on t he i r seaward face. Consolidated coral rubble covered with l iv ing A. cervicornis i s typical (Land and Goreau, 1970). Other coral spec ies appear o n the steep seaward faces and la te ra l reef flanks adjacent to the sand areas , including Pori tes as t reoides , Mycetophyll ia sp . , and Montastrea cavernosa as well as M. annularis. In Discovery Bay the A . cervicornis reefs drop steeply a t the seaward edge to a zone of sand- about 40 to 60 m wide. Upper S i l l Reefs: A se r ies o f elongate bu t d iscont inuous ree fs a re es tab l ished a t 40 t o 50 m depths, and r i s e t o about 25 m depths. P l a t y colonies o f Montastrea annu la r i s are dominant along w i t h a mixed co ra l community composed p r i m m ~ q a r i c i a lamarck i , Mycetophyll i a sp., o the r species of Agar ic ia, Scolymia sp., f l e s h y algae, sponges and ant ipa thar ians . Fore Reef Slope: The p l a t y growths o f Montastrea annu la r i s along w i t h Agar ic ia sp. and abundant sponges, gorgonians and ant ipa thar ians cont inue down the r e e f f o r e slooe t o a depth o f 50 m, below which e i t h e r Agar i c ia sp. dominate, o r mixed 'cora l growths occur cons i s t i ng of Agar ic ia sp., Madracis sp., Montastrea cavernosa, and Mycetophy l l ia sp. Lower S i l l Reefs: This zone, a t depths o f 60 t o 70 m, i s charact- e r i z e d by e i t h e r elongate r i dges having l ess than 10 m r e l i e f o r i s o l a t e d rocky outcrops. Corals occu r r i ng on these fea tures i nc lude Agar i c ia sp., Montastrea cavernosa, Madracis sp., and Mycetophy l l ia sp. Sponges and ant ipa thar ians are a l s o abundant. Deep Fore Reef Slope: Scl e r a c t i n i an co ra l s become inc reas ing l y sparse w i t h depth, and t h e dominant benth ic organisms are sc le ro- sponges and demosponges. The Discovery Bay Laboratory The Discovery Bay Laboratory i s j o i n t l y operated by t h e Marine Sciences Research Center o f the Sta te U n i v e r s i t y o f New York, and the U n i v e r s i t y o f the West Indies. The Laboratory s i t e cons is ts o f .06 sq. k i lometers (15 acres) o f coasta l proper ty . F a c i l i t i e s have been designed p r i m a r i l y f o r co ra l - ree f research w i t h emphasis on on-s i te i n v e s t i g a t i o n s (SCUBA f a c i l i t i e s ) r a t h e r than f o r t r a i n i n g programs. Therefore, on l y l i m i t e d space i s a v a i l a b l e t o v i s i t i n g s c i e n t i s t s and the re are no dormi to ry accommodations. The new c e n t r a l b u i l d i n g cons i s t s o f an a i r - cond i t i oned research u n i t housing fou r small l abo ra to r i es , a l a r g e "wet" labora tory , a dark room, instrument s to re , museum and two o f f i c e s . The "wet" l a b i s sub- d i v i d e d i n t o s i x semi-enclosed research bays provided w i t h a l l serv ices i n c l u d i n g seawater and cen t ra l t a b l e s f o r aquaria and s o r t i n g and hold- i n g tanks. A separate reading and conference room i s nearby. Support f a c i l i t i e s inc lude a machine shop, a boat and wood workshop, and a d i v i n g locker . Three boats are a v a i l a b l e a 6.7 meter (22- foot) t w i n outboard-motor vessel, a 4.6 meter (15- foot) work boat, and a 3.7 meter (12- foot) i n f l a t a b l e boat t h a t can be c a r r i e d by c a r t o remote s i t e s . A landrover and Volkswagen bus serve t o t ranspor t equipment as we l l as personnel. The l abo ra to ry i s equipped w i t h a g l a s s - d i s t i l l e d water supply, d i s s e c t i n g and compound microscopes, h i s t o l o g i c a l apparatus, d ry ing oven, pH meter, t o p load ing and a n a l y t i c a l balances, photographic equip- ment and darkroom f a c i l i t i e s , r e f r i g e r a t o r , f reezer , cent r i fuge, collecting gear, e t c . The diving f a c i l i t y contains two high-capacity a i r compressors, a recompression chamber with a i r bank, SCUBA tanks, regulators and auxi l iary diving equipment. Summary The reefs off Discovery Bay Laboratory probably are some of the most extensively studied reefs in the world, and the data already available would be a d i s t i n c t asse t t o a research program. The Discovery Bay reefs a re r ich and well developed both in shallow and deep water. Since they occur very close t o the coast , t h e i r location would f a c i l i t a t e close monitoring of various parameters. More- over, the re la t ive ly narrow horizontal extent of the reef biotope would be advantageous in instrument monitoring, as well as in on-site investigations. This s i t e i s readily accessible from the U.S. mainland, and accommodations are available local ly a t inexpensively priced guest houses (ca. $7/day s ingle) f o r as many a s 40 s c i en t i s t s . On the other hand, there i s a s ignif icant b u t a s yet undetermined t e r r e s t r i a l influence on the reef ecosystem in t h i s area. The internal sediment found i n association with the present reefs i s character is t ic- a l l y brown owing t o the presence of bauxite or iron oxide tha t has been incorporated from t e r r e s t r i a l sources. From 1964 t o 1967, a ship channel was dredged and blasted in to Discovery Bay t o permit docking of large ocean-going vessels in t h i s bay. The channel i s about 300 to 400 m from some of the key areas of potential research. The construction of t h i s channel as well as frequent v i s i t s of large ships in to the bay provide negative influences on the reef ecosystem i n t h i s area. The paucity of larger reef f i sh i s notable in Discovery Bay. Although f i sh populations are sparse, f i s h t raps a re s t i l l used exten- sively, and therefore t h i s location has limited potential fo r any proposed f i s h studies in association with the reef ecosystem. The poor development of a but t ress zone in the area i s an additional drawback. The Discovery Bay Laboratory i s not eqippped t o handle large coral - reef study programs because of i t s re la t ive ly small s ize . Research projects might a lso be hampered by customs delays and the need fo r import l icenses owing t o the problem of bringing onto the island sc i en t i f i c equipment and material from another country. Glover's Reef, Bri t ish Honduras* A f i r s t reconnaissance v i s i t t o Bri t ish Honduras, 17-22 January 1971, included several diving s ta t ions along the bar r ie r reef and "Preliminary survey: January 17-22, 1971 by A. Antonius and J . N. Weber. Field survey: June 20-27, 1971 by A. Antonius, A. L. Dahl, and K. Ruetzler. Turnef fe Is land, and an a i r survey o f the b a r r i e r ree f , Turnef fe Island, Lighthouse Reef and Glover 's Reef. The in fo rmat ion obtained s t r o n g l y suggested t h a t an extensive d i v i n g survey be focused on Glover 's Reef, and t h i s was conducted 20-27 June. Desc r ip t i on o f Reefs G love r ' s Reef (160501N, 87"501W) i s the southernmost o f the th ree B r i t i s h Honduras a t o l l s , about 75 km SE o f Be l i ze City, 45 km E o f the mainland and 25 km E o f t h e b a r r i e r r e e f . It i s elongated i n NNE-SSW d i r e c t i o n , about 28 km long and 10 km wide. The a t o l l (F ig. 9 ) i s surrounded by deep water (350-7000 m) w i t h i n 1-2 km from the i n t e r t i d a l per iphera l ree f . The we l l developed surface-breaking a t o l l reef f l a t (mainly co ra l and c o r a l l i n e algae) i s i n t e r r u p t e d by two major openings: NE-entrance and SW-entrance (12 m deep). On i t s SE pa r t , i t supports a chain o f s i x cays which are d i s t r i b u t e d over a d is tance o f 10 km. The cays vary between 150 m and over 1 km i n length and are more o r l e s s covered by coconut palms. The lagoon, i n cont ras t t o the o ther two B r i t i s h Honduras a t o l l s , i s r a t h e r deep, 8-15 m i n most par ts , w i t h hundreds o f patch ree fs r i s i n g t o the surface. O f t h e s i x cays on Glover 's Reef, on l y Long Cay and adjacent L i t t l e Cay are permanently inhabi ted. Both i s l ands are used as a r e s o r t f o r d i v ing -o r i en ted t o u r i s t s . From the seaward margin o f t h e per iphera l r e e f a g radua l ly s lop ing r e e f - f r o n t extends t o t h e d rop -o f f . The r e e f f r o n t va r i es i n w id th from about 400 m (SW cays) t o 1.5 km (Long Cay) and i s about 500 m wide on the leeward s ide. The edge o f t h e drop-o f f occurs a t a depth o f 15-25 m. A system o f deep p a r a l l e l grooves runs perpendicular t o i t . These grooves are p a r t i c u l a r l y well-developed on the windward ( E ) s i de o f the a t o l l . Sediments produced i n the breaker zone are t ransported down the grooves p e r m i t t i n g undisturbed co ra l growth i n between. The leeward r e e f s a re considerably more in f luenced by sediments which are d r i v e n ou t o f t h e lagoon by wind generated cur ren ts . The windward s ide shows a dominating, 1 ush, growing ree f - co ra l community i n c l u d i n g sponges and gorgonians (Fig. 10) . No attempt was made t o gain a complete l i s t o f species dur ing the survey, bu t t h e general appearance o f the r e e f and the unpara l le led s i z e o f most o f the co ra l and sponge specimens observed s t r o n g l y suggest t h a t a l l o f t h e known Caribbean species can be found there. On the sketches of the r e e f t ransec ts on ly dominant species are l i s t e d . Compared w i t h the windward r e e f , t h e leeward s ide shows a s l i g h t decrease i n number o f s c l e r a c t i n i a n species and i s s l i g h t l y l e s s populated, probably due t o sedimentat ion (F ig. 11). The co ra l cover ends around 45 m and gives way t o a sand slope. The fo re - ree f slope i s very steep a t NE Cay, almost v e r t i c a l a t Long Cay and v e r t i c a l t o overhanging a t SW Cay. It was explored a t f i v e d i f f e r e n t l oca t i ons t o a depth of 60 m. There, and as f a r down as one could see i n the c lear water (horizontal v i s i b i l i t y a t l eas t 50 m ) , lush coral , sponge and algal growth continues. The patch reefs r i s e t o the surface from the lagoon f loo r , which i s generally covered with sand and Thalassia. These reefs are mainly of Acropora palmata and A. cervicornis s t ructure on top and Montastrea annularis below, with coral growth occurring t o a depth of 3-4 m and occasionally 10 m. Every patch reef i s a small individual reef-enti ty and ideal for a variety of experiments. The winds in the v ic in i ty of the Brit ish Honduras a t o l l s blow steadily from the East for most of the year, being replaced between November and February by occasional Northern winds, which occur only a few times every winter and l a s t four or f ive days a t a time. Thus, the wind-generated western currents in the a to l l area are by f a r predominant over the main Caribbean current, which impinges on the British Honduras a t o l l s i n a northerly or southerly direct ion, and regularly changes i t s direction several times a year. Water temperatures a t the surface vary between 25OC in winter and 31?C in the summer (open water); no s ignif icant temperature decrease was noticed during the dives ( for detailed information on climatic, biologic and geologic features of Glover's Reef, see Stoddart, 1962). Summary British Honduras i s a po l i t i ca l ly s table country, and i t s o f f ic ia l language i s English. Re1 ations between the government and resident North Americans or Europeans a r e cordial. The Brit ish Honduras govern- ment seems t o look favorably upon educational and research projects, and duty exemption for a l l s c i e n t i f i c and educational equipment and materi a1 s has a1 ready been granted. The remoteness of Glover's Reef makes i t l ike ly tha t i t will remain undisturbed; s t i l l , there i s good access ib i l i ty t o Belize by a i r , and the a to l l can be reached in any weather by boat because two-thirds of the t r i p takes place i n a sheltered lagoon between the mainland and the barrier reef. There are no research f a c i l i t i e s on the a to l l a t present, and i t i s not always possible t o make sat isfactory arrangements for an extended sc i en t i f i c program using tourist-oriented f a c i l i t i e s . Glover's Reef i s well suited for d i rec t comparison with Indo- Pacific a to l l s as there are no land influences, no pollution, or other human disturbances; yet the large barr ier r ee f , and two comparable b u t physiographical l y d i f fe ren t a t o l l s are readily accessible. While Glover's Reef i s the only one of the three a t o l l s which was not h i t by the devastating hurricane Hattie i n 1961, there i s a hurricane potential; a small one passed over the a to l l in November 1971 without causing major damage. In contrast t o the other reefs surveyed i n the Caribbean area, Glover's Reaf a to l l appears t o offer the greatest variety of reef types and the optimum r e e f development i n terms o f populat ion dens i ty and species d i v e r s i t y o f r e e f co ra l s and associated organisms. San Blas Is lands , Panama* The San Blas Is lands (S039'N, 78'45'W) l i e o f f t h e Caribbean coast o f Panama between San Bl as Po in t and Cape T i buron (F ig. 12) . During the Spanish conquest o f Panama, the Cuna Indians were d r i ven from the mainland t o f i n d re fuge on these is lands . Today, they have t i t l e t o over 365 is lands and a s t r i p o f land along the adjacent coast . They have j u r i s d i c t i o n over agreements t o buy, s e t t l e o r e s t a b l i s h any forms o f business on t h e i r i s lands . Standard access t o the San Blas Is lands i s by means o f l i g h t plane o r small boat. The i s l ands are about 110 km east o f Colon, and i t takes about 8 hours from the re by boat. There are scheduled f l i g h t s t o several San Blas l o c a l i t i e s from Panama City, and a f l o a t plane serv ice from Colon. One ser ious problem t o be considered concerning t r a v e l by sea i s t h a t rough waters are common du r ing the months o f December t o A p r i l , so the i s l ands genera l l y are n o t accessib le by boat. A i r s t r i p s such as a t C a r t i on the mainland (see Fig. 12) a re accessib le year round t o l i g h t a i r c r a f t t h a t can t ranspor t passengers t o and from Panama C i t y a t a cos t o f $9.25 per one-way t r i p . Although t h i s provides a r a p i d and i n - expensive form o f t ranspor ta t i on t o t h e area, and could be used by s c i e n t i s t s when the is lands are inaccess ib le by water, t h i s a i r c a r r i e r cannot always operate on a s p e c i f i c schedule owing t o t h e frequency o f poor weather condi t ions. I n a d d i t i o n , the sho r t and uneven d i r t a i r - s t r i p s preclude the use o f l a r g e r a i r c r a f t so t h a t on ly f o u r o r f i v e passengers and l i g h t baggage loads can be accommodated on any one f l i g h t . The on l y housing f a c i l i t i e s p resen t l y ava i l ab le i n t h e area are located on Picofeo Is land, which i s t h e on l y p r i v a t e l y owned San Blas is land. These f a c i l i t i e s c o n s i s t o f several sheds and some machinery ( i nc lud ing a 6 - v o l t generator) t h a t a re p a r t o f a disused copra- processing p lan t . One b u i l d i n g serves as t h e Picofeo Hotel which i s operated by Jose Garcia, one o f two Panamanian brothers who own the i s l a n d and i t s bu i l d ings . The h o t e l , which i s an o l d wooden s t ruc tu re , o f f e r s minimal accommodations t o v i s i t o r s , who are expected t o b r i n g t h e i r own food suppl ies a t $2.50/night. Extended stays can be arranged a t cheaper ra tes . Outsiders are genera l l y discouraged from s e t t i n g up reisdence i n t h e o the r i s lands . However, a few t o u r i s t reso r t s a re now being developed i n the San Blas Is lands, which should prov ide f o r improved access and accommodations. A sho r t i n i t i a l v i s i t conf irmed t h e p o t e n t i a l o f the area, and *Prel iminary survey: May 1-2, 1971 by A. L. Dahl and.A. Childs. F i e l d survey: August 3-7, 1971 by A. Antonius, J. C. Land, and I . G. Macintyre (ass is ted by C. Eiirkeland, D. Meyer, M. McCosker, W. K. Sacco). located some s i t e s fo r fur ther study. Ten locations were then vis i ted during the four-day survey. These areas a re marked on maps redrawn from sections of HO chart No. 2771, San Blas Point t o Concepcion Bay (Figs. 13, 17, and 22). Following are brief descriptions of the areas i n the order t ha t they were v i s i t ed : 1. Off Sail Rock, Profi le A-A' Sail Rock i s an exposed rock knob marking an isolated reef not f a r from Porvenir Island, where there i s a very shor t a i r s t r i p (Fig. 13). The water was very turbid during the survey of the reefs i n t h i s area ( l e s s than 2 m v i s i b i l i t y a t a depth of 20 m ) and there was a marked thermocline a t a depth of about 3 m. The zonation across the prof i le i s given i n Fig. 14. Points worth noting in t h i s area are the dominance of Asaricia sp. corals a t the edge of the shallow platform and the lack of coral growth below about 20 m where the accumulation of f i n e sediments (r ich in organic mater ia l ) apparently prevents corals--other than the bladed Agaricia tenuifol ia--from growing. 2. Picofeo Island The Picofeo Hotel i s s e t on p i les over the water a t the north end of Picofeo Island (Fig. 13). This area i s characterized by typical back-reef coral communities (Fig. 15) dominated by Porites pori tes . As was found a t Sail Rock, the outer edge of the platform i s marked by a r i ch Agaricia sp. coral community. The reef terminates in a shallow sand f l a t a t a depth of about 6 m. 3. Off Sardingan Point, Prof i le B-B' Sardingan Point i s on the mainland near San Blas Point and west of Picofeo Island (Fig. 13). The reef in t h i s area drops off steeply t o a depth of about 30 m (Fig. 16) . An in teres t ing aspect of t h i s reef i s t ha t Agaricia sp. corals tend t o dominate throughout the depth zones except in very shallow water, where Pori t e s pori t e s i s the most abundant coral . 4. Off Salar, Profi les C - C 1 , D - D ' , and E - E ' The group of islands a t Salar are among the inner San Blas Islands north of Macolla Point (Fig. 12 ) . As names for the individual islands were not available, they have been numbered from southwest t o northeast (Fig. 17). The survey concentrated primarily on the area between islands 2 and 3. Several dives were completed in t h i s area of vigorous coral growth, spectacular drop-offs, and overhangs which a re described in prof i les C-C' (Fig. 18) , D - D o (Fig. 19) , and E-E' (Fig. 20). The general zonation of the reef f l a t of island no. 3 i s a lso shown in a diagramatic sketch (Fig. 21 ). 5. Off Ogopuquip, Prof i le F-F' Three d i f fe ren t s i t e s were vis i ted in the Holandes Cays (Fig. 22). The f i r s t was a t Ogopuquip on the southern s ide o f the keys, where a p r o f i l e (F-F ' ) was prepared o f f t h e southvest p o i n t o f Ogopuquip Is land (Fig. 23). This area had the g rea tes t d i v e r s i t y o f co ra l species o f any s i t e v i s i t e d i n the San Blas Is lands . 6. Holandes Cays Algal Ridge, P r o f i l e G - G ' The d i s t i n c t i v e s t ruc tu re o f the nor thern r e e f around the Holandes Cays i s i l l u s t r a t e d by p r o f i l e G-G' (F ig. 24) . This r e e f has a prominent a l g a l r i d g e t h a t has been described i n more d e t a i l elsewhere (Glynn, i n press). 7 . O f f Holandes Cays, P r o f i l e H-H' Beyond the a l g a l r i d g e o f f t h e eas t end o f Holandes Cays, there i s a broad rock p l a t f o r m s i m i l a r t o t h a t observed o f f many o f the Bahamian Is lands ( p r o f i l e H-H' , Fig. 25). The survey d i d n o t extend t o the drop- o f f which, according t o the char t , probably occurs 200 m beyond the outer end o f the p r o f i l e . Summary Some r e e f areas, i n p a r t i c u l a r t h e r e e f s o f f Salar Is land, meet the s c i e n t i f i c c r i t e r i a es tab l ished above f o r a research s i t e i nc lud ing : well-developed ree fs , steep fo re - ree f slope, l a c k o f d is turbance by human a c t i v i t y , and a c lose p r o x i m i t y t o shore. The well-developed a l g a l r i d g e o f f the Holandes Cays o f f e r s an oppor tun i ty t o make compar- a t i v e s tud ies w i t h s i m i l a r r e e f s t ruc tu res i n the P a c i f i c . Bu i ld ings are ava i l ab le f o r conversion i n t o research f a c i l i t i e s . However, they are n o t located adjacent t o well-developed ree fs . Smithsonian Trop ica l Research I n s t i t u t e mainta ins marine l a b o r a t o r i e s and a research vessel elsewhere i n Panama, and conducts research i n t h e San Blas I s land area. Two a t y p i c a l c h a r a c t e r i s t i c s o f San Blas ree fs , i n comparison w i t h other Caribbean ree fs , are the predominance o f Agar ic ia sp. cora ls i n a l l depth zones, and the presence o f a well-developed a l g a l r i d g e o f f Holandes Cays. The i n a c c e s s i b i l i t y o f t h i s area by boat f o r f i v e months o f the year and the l i m i t e d a i r t r anspor t serv ice o f fe red present a ser ious l o g i s t i c a l problem i n t ranspor t i ng personnel and mater ia l i n and o u t o f the San Blas area. As f a c i l i t i e s on Picofeo I s l a n d are too d i s t a n t from good r e e f development t o be usefu l , i t would become necessary t o negot ia te lease arrangements w i t h the Cuna Indians f o r one o f the un- inhab i ted is lands . PACIFIC AREA In the Pacific Ocean, coral reefs are scattered over a vast geo- graphic area. The only safe generalization about Pacific reefs i s t ha t no two reefs are ident ical . However, in sp i t e of the great d ivers i ty in form and composition tha t characterizes these communities, a number of features demonstrate the underlying relationships within t h i s assemblage. The coral fauna i s remarkably uniform throughout the Indo- Pacific (Wells, 1969), with the highest d ivers i ty and the center of evolutionary radiation in the western tropical Pacific (Stehl i and Wells, 1971). This f a c t frequently leads t o the statement t ha t reefs in the western Pacific are "rich" while those t o the ea s t are "impoverished." Whether t h i s applies t o a l l elements of the f lo ra and fauna, however, has ye t t o be demonstrated. The a to l l reef form i s another common Pacific feature , a1 though involving many s t ructural variat ions on the basic theme. The presence of a Porol i thon algal ridge i s often believed to be charac te r i s t i ca l ly Pacific, and i s cer ta inly common there , while i t rarely occurs on Atlantic ree fs . The great number and d ivers i ty of tropical Pacific reefs has made the search for data on research s i t e s very d i f f i c u l t . Information was compiled from many sources on reef s t ructure and composition, and on log i s t i c and practical arrangements. Most published descriptions of Pacific Islands include only t e r r e s t r i a l geography; specif ic local descriptions a re rare , and only the shallow reef i s mentioned. Charts from the U. S. Army Map Service and the Navy Hydrographic Office in- dicate with f a i r accuracy the presence or absence of shallow reef s t ruc ture , and sometimes the nature of the offshore slope and the depth t o which coral development might be expected. A few s c i e n t i f i c papers include reef descriptions of some s o r t , b u t not fo r the deeper areas important t o a complete reef program,and generally i n l o c a l i t i e s t h a t are otherwise unsuitable. Most of the past research on Pacific reefs was conducted e i t he r by ship-borne expeditions t o otherwise inaccessible islands, or in areas t ha t have since been disturbed by development. Also, published information tends t o be too dated t o be r e l i ab l e fo r current reef conditions; much of i t predates World War 11. While many questionnaires were returned by Pacific reef special- i s t s , t h e i r information did not always meet program requirements, perhaps because of the great s i z e and complexity of the area. Personal experience tended t o be limited t o a few s i t e s and to what could be seen from above the water. Areas of f i e ld experience and interpret - a t ions of reef quali ty depended on the respondant's special i n t e r e s t (an interes t ing reef t o a coral spec i a l i s t would not necessarily be selected by a bird or sea-snake expert , f o r example). Also, the information was often too generalized, referring t o island groups rather than t o specif ic reefs. The areas recommended and a summary of the questionnaire responses fo r the Pacific area are l i s t ed i n Table 11. Reef Site Structures* - Cook Islands Hervev Islands Line Islands Chrisbnas Island Fanning Island 0 Society 1s1ands Tahiti Other New Caledonia Port Moresby, Papua i New Guinea Ashore Reef, 0 Timor Sea Heron Island, G.B.R. i Low Isles, 0 N. @eensland Aldabra, Indian Ocean - Undisturbed + Accessible - 0 0 + + i - + i - Political Facilities Status+ Number of Recommendations Sites suggested by name only: Saipan, Marianas Islands; Nukuoro, Caroline Islands; Wotho, Marshall Islands; Onotoa Atoll, Gilbert Islands; Butaritari Atoll (Makin), Gilbert Islands; Phoenix Islands; Buka, Bougainville, Solomon Islands; Rabaul; New Britain; Bismarck Archipelago. Key: + = favorable or present 0 = unknown or possible - = unfavorable or impractical + = variable * see selection criteria above, p. 38 + basically the anticipated government attitude to a large international but U.S. funded program Questionnaires and equivalent information on Pacific sites were returned by the following: A. Antonius, A. L. Bloom, A. L. Dahl, M. S. Doty, F. R. Fosberg, R. Hagemeyer, H. Heatwole, R. Johannes, H. S. Ladd, E. G. Menez, H. A. ReMer, R. W. Schreiber, J. Sieburth, W. A. Starck, D. R. Stoddart, J. N. Weber, C. M. Yonge, and University of Hawaii. 64 It was obvious from the quest ionnaires and the o the r in fo rmat ion a v a i l a b l e t h a t data on t h e r e e f s t a t e and s t ruc tu re , and on cu r ren t l o g i s t i c arrangements, could on l y be obtained by ac tua l s i t e v i s i t s by s c i e n t i s t s f a m i l i a r w i t h t h e basic c r i t e r i a f o r a research s i t e , as was done i n the Caribbean. As i t would have been p h y s i c a l l y impossible t o v i s i t every p o t e n t i a l s i t e , i t was necessary t o s e l e c t from those on which some data were a v a i l a b l e the areas w i t h the bes t prospects f o r a s a t i s f a c t o r y s i t e . One important r o l e o f the quest ionnaires was t o help p i n p o i n t such areas f o r more d e t a i l e d examination. I n e v i t a b l y o the r areas which might be i dea l have been overlooked because they were n o t s u f f i c i e n t l y we l l known, o f t e n because they are more remote. Trans- p o r t a t i o n i n the P a c i f i c however i s improving r a p i d l y , and w i l l make more such areas l o g i s t i c a l l y p r a c t i c a l . A number o f i s l ands i n the P a c i f i c have r e c e n t l y been recommended f o r preservat ion as " Is lands f o r Science" ( E l l i o t t , 1971). These were selected f o r t h e i r l ack o f human disturbance, and i n many cases, have exce l l en t cora l reef development. A number o f them were i n f ac t considered a t an e a r l y stage i n t h e screening process. However, they are undisturbed p r e c i s e l y because they are inaccessib le, and thus l o g i s t i c a l l y impossible f o r a major research program. S i t e Screening An i n i t i a l screening o f poss ib le s i t e s i n the P a c i f i c reduced t o a more manageable number the l i s t o f areas being considered al though many excluded areas have c h a r a c t e r i s t i c s t h a t might be o f g rea t i n t e r e s t f o r more spec ia l i zed programs. The f o l l o w i n g c r i t e r i a were appl ied i n the screening. Because o f t h e need t o f i nd a r e e f w i t h as few compl icat ing external in f luences as possib le, preference was g iven t o a t o l l s o r b a r r i e r r e e f areas f a r from h igh i s l a n d in f luences. For a l a r g e program, the need f o r regu la r a i r serv ice o r an immediately adjacent a i r s t r i p f o r poss ib le cha r te r se rv i ce e l im ina ted many l ess accessib le s i t e s . The p o s s i b i l i t y o f acqu i r i ng o r cons t ruc t ing and main ta in ing a f a c i l i t y f o r a t l e a s t 20 people a t reasonable cos t was a l so considered important. A l l French t e r r i t o r i e s were omit ted from cons idera t ion because o f un- c e r t a i n t i e s concerning long-term cooperat ion w i t h government a u t h o r i t i e s . Great d is tance o r t r a v e l t ime from the Uni ted States was a l so considered t o be l ess des i rab le . The r e s u l t s o f the screening are given below by i s l a n d group, w i t h t h e reasons f o r i n c l u s i o n o r exc lus ion from f u r t h e r considerat ion. Excluded from f u r t h e r considerat ion: a. Hawaiian Islands--Reef s t r u c t u r e and d i v e r s i t y inadequate. Disturbed. b. Mariana Islands--High is lands . Reefs marginal i n q u a l i t y . Serious d is turbance by war and Acanthaster. c. Guam--Reefs se r ious l y disturbed. d. L ine Islands--Poor a c c e s s i b i l i t y . Far from center o f co ra l d i v e r s i t y . e. Phoenix I s 1 ands--1naccessi b le . Canton d is turbed. f . Tokelau Islands--Inaccessible. g . Tonga--Poor accessi bil i ty . h . Cook Islands--Poor access ib i l i ty . i . Bismarck Archipel ago, Solomon Is1 ands, New Hebrides--High islands or inaccessible. j. New Caledonia, French Polynesia--French administration. K . Great Barrier Reef and other Australian reef areas--Great distance. A fur ther breakdown was made of those areas not excluded. Those dis- cussed in more detail l a t e r in t h i s paper are marked with a n as te r i sk (*) . Carol ine Islands U . S . administration, good a i r service to d i s t r i c t centers , many a t o l l s , in center of coral d ivers i ty . Palau*. Many recommendations. Greatest biological d ivers i ty in Pacific Islands. Regular a i r service. Some f a c i l i t i e s . Considerable barrier reef area and one small a to l l (Kayangl). Helen Reef. Rich and undisturbed, b u t inaccessible. Yap*. High island. Regular a i r service. Some r ich reef areas . Ulithi Atoll*. Ai rs t r ip and weekly Coast Guard f l i g h t , large a t o l l . Some disturbance during World War 11. Woleai Atoll*. Small, undisturbed a to l l with abandoned a i r - s t r i p . Excellent reefs . Ifaluk Atoll . Very small a t o l l , inaccessible. Truk. Accessible b u t reefs seriously disturbed. Ponaoe*. Reqular a i r service. Hiqh island with barr ier ree f . some' reef disturbance. - A n t Atoll*. Near Ponape, undisturbed except fo r recent Acanthaster damage. Pakin Atoll*. Near Ponape, undisturbed. Nukuoro. Inaccessible. Kapingamarangi Atoll , Heavily populated, inaccessible. Marshal 1 Is1 ands U. S. administration. Many large a t o l l s , mostly e i t he r disturbed or inaccessible. 1 . Majuro*. Regular a i r service, disturbed. 2. Arno*. Near Majuro, undisturbed. 3. Wotho. Inaccessible. 4 . Ailuk. Undisturbed b u t inaccessible. 5. Eniwetok. Accessible. Excellent f a c i l i t i e s , disturbed. Clearance for foreign par t ic ipants d i f f i c u l t to obtain. Gilbert and El l ice Islands Soon to have improved regular a i r service. 1 . Funafuti. Serious World War I 1 damage and h e a v i l y p o p u l a t e d . Sta te o f o the r r e e f s unknown. F i j i I s l ands Large h igh i s l ands access ib le . Rich r e e f s i n o u t l y i n g areas, no i n fo rma t i on on poss ib le s p e c i f i c s i t e s . Samoa Is lands Accessible. Good f r i n g i n g r e e f development. 1. Tu tu i l a * . Reefs d i s t u r b e d i n more populated areas. 2. Manua Islands*. Undisturbed. 3. Rose A t o l l . Small and t o t a l l y undis turbed. A c c e s s i b i l i t y d i f f i c u l t . On t h e bas is o f t h e i n i t i a l screening, a number o f areas most l i k e l y t o o f f e r s u i t a b l e s i t e s were selected, i n c l u d i n g Palau, Yap, U l i t h i , Woleai, Ponape ( i n c l u d i n g adjacent Ant and Pakin) , Arno, and Samoa ( i n c l u d i n g Rose At01 1 ) . A survey team was the re fo re sent t o examine these areas. F i j i and t h e G i l b e r t and E l l i c e I s l ands a l s o have g r e a t p o t e n t i a l , bu t too l i t t l e i n fo rma t i on was a v a i l a b l e on which t o p lan a s i t e v i s i t , and t ime d i d n o t permi t t he more leng thy survey t h a t would t he re fo re be requ i red . U.S. T r u s t T e r r i t o r l * O f t he enormous number o f r e e f areas sca t te red throughout t he Marianas, Caro l ine and Marshal l Is lands, most were too i naccess ib le t o be considered, bu t even the access ib le areas i nc lude v a s t r e e f t r a c t s w i t h innumerable p o t e n t i a l s i t e s . The experience and r e p o r t s o f t he Acanthaster surveys conducted i n Micronesia by the U n i v e r s i t y o f Guam (Tsuda, 1971) and the Acanthaster c o n t r o l teams operated by the U.S. T rus t T e r r i t o r y Admin i s t ra t i on were o f p a r t i c u l a r va lue i n s e l e c t i n g promising areas f o r t h e survey team v i s i t s . T ranspor ta t ion i n Micrqnesia i s improving. Cont inenta l A i r 1 ines-- A i r Micronesia now serves a l l t he d i s t r i c t centers (Koror, Yap, Saipan, Truk, Ponape, and Majuro) 2 t o 3 t imes per week. I n add i t i on , A i r P a c i f i c has a c h a r t e r se rv i ce a v a i l a b l e f o r $180/hour (7-passenger a i r - planes), and the re i s a new c h a r t e r serv ice by I s l a n d Av ia t i on , Inc., ( r a t e s around $120/hour). However, f l y i n g by o t h e r than Navy seaplanes i s s t i l l r e s t r i c t e d t o those few i s l ands w i t h a i r s t r i p s . Vessels se rv i ce o u t l y i n g i s l ands a t i n t e r v a l s o f two weeks t o several months. * F i e l d survey: August 14-September 5, 1971 by A. Antonius and A. L. Dahl [ass is ted by R. Randall (Palau and Yap), R. T. Tsuda (Ponape) and B. Sablan (Majuro and Arno)] Local transportation i s d i f f i c u l t , although small motor boats can usually be hired. The Acanthaster control teams have diving compressors and good motor boats a t Palau, T r u k , Ponape, and Majuro, and provided much of the l og i s t i c support f o r the survey team. There i s an excellent marine laboratory and s ta f f a t the University of Guam, and the U.S. Trust Terri tory Government i s highly cooperative. There i s some resistance from the local populations t o outside interference b u t i n general they appeared favorably disposed towards s c i e n t i f i c as opposed t o commercial or t ou r i s t ac t iv i t i e s . There are small hotels in the d i s t r i c t centers, b u t in outlying areas i t i s necessary t o depend on the hospitali ty of the local people. Palau (7'3O1N, 1340301E) i s considered t o have the r ichest reefs of any Pacific island area. The archipelago of volcanic and high and low limestone islands contains a complex of fringing and patch reefs par t ly surrounded by a bar r ie r reef (Fig. 26). A small a to l l occurs immediately no r th of the barr ier reef. After a detailed examination of charts and consultations with local spec i a l i s t s , a number of s i t e s ranging throughqut the archipelago were chosen for detailed examination. This was a wholly inadequate sample of the divers i ty of s i t e s available, So no generalizations should be made from the few descriptions presented here. 1. Ngeregong Island The f i r s t dive was a t Ngeregong Island, on the windward (eas t ) s ide of the barr ier reef about 32 km south of Koror (Fig. 26) . Ngeregong has an abandoned Japanese a i r s t r i p t ha t , i f restored, could simplify l og i s t i c arrangements. The outer reef slope, on the SE s ide of the island consists of a gentle sandy slope, w i t h scattered coral growth, becoming even sandier with depth. The reef was poorly developed. 2. Angaur Angaur Island l i e s 40 km beyond Ngeregong, a t the southern end of Palau, beyond the barr ier reef (Fig. 26). I t has an excellent Coast Guard a i r f i e l d serviced weekly from Guam. The island, formerly an important source of phosphates, lacks major reef development except on the south and west sides. The team dived on the west (leeward) s ide of Angaur (Fig. 26), in the center of reef development. There is a smooth rocky f l a t w i t h small scattered corals extending 700 m offshore t o a depth of 5 m , followed by a gentle slope 200 m wide and going down t o 8 m , w i t h larger , more abundant corals. A steeper slope, 100 m wide w i t h lush l iving coral coverage, extends down t o 38 m , where i t i s interrupted by a sand f l a t 30 m wide. A slope with patches of coral interspersed w i t h sand continues down from 40 m a t 1 km offshore. The water was warm and c lear . The lack of diverse coral habi ta ts , the amount of sediment, and the extended zonation interrupted by terraces were considered undesirable for a research s i t e . The Ngemelis I s lands a r e 40 km SW of Koror on t h e western b a r r i e r r ee f (Fig. 26). The i s l ands a r e pa r t s of a s l i g h t l y e leva ted coral platform, and f r o n t on t h e ou te r r ee f margin on t h e west and a she l t e red channel on t h e sou theas t , as well as t h e lagoon t o t h e north. The i s l ands can be reached from Koror by boat without going outs ide the b a r r i e r r e e f . Two d ives were made, f i r s t on the SE s i d e , o f f a v e r t i c a l drop-off going down t o 240 m , and then on the s loping west s i d e . 3. Ngemelis I s lands , SE On t h e she1 te red SE s i d e of Ngemelis (Fig. 28), t h e shallow reef f l a t i s 300 m wide, with few c o r a l s inshore , increasing t o dense coverage of the r ee f edge a t 0 .5 m depth, with s o f t c o r a l s predominant. Beyond t h e edge i s a v e r t i c a l s lope , mostly overhnging, dropping t o 260 m . Alcyonarians dominate down t o 30 m , with s c i e r a c t i n i a n s and gorgonians equal ly represented. The s lope begins t o p ro jec t outward a t 40 m , c o l l e c t i n g calcareous sediment; co ra l s a re scarce and appear not t o grow beyond 30-60 m. The water temperature decreases with depth. The drop-off i s spec tacu la r , but s o f t co ra l s a r e dominant, and t h e r e i s apparent ly a cons tant flow of sediment. 4. Ngemelis Islands,W The west (leeward) s i d e of t h e Ngemelis Is lands (Fig. 29) has a broader reef f l a t 500 m wide with s tony co ra l s dominant a t the seaward edge ( 0 . 5 m depth) . There i s a s t e e p drop-off t o 12 m with good coral coverage, with big bu t t r e s ses dominated by Por i t e s heads and rubble- f i l l e d chutes extending down t o 20 m. A gen t l e s lope extends down t o 40 m w i t h Pachyseris dominant, a f t e r which the slope becomes increas ingly sandy and co ra l s dwindle. Again water temperature decreased with depth. The s i t e has high spec ie s d i v e r s i t y and a good drop-off , but sediment- a t i o n l i m i t s coral growth a t around 50 m. A t both s i t e s t h e drop-offs a r e very near t h e su r face , and sponges a re almost completely lacking. Current pa t te rns in t h e area appeared q u i t e complex, and t h i s , toge ther with t h e inadequacies i n the reef s t r u c t u r e , would r u l e t h i s area out f o r c e r t a i n types of r ee f research. Kayangl Atoll i s several hours away by small boa t , t r a v e l i n g u p i n s i d e t h e western b a r r i e r r e e f , and then beyond t h e northern t i p of Palau (Fig. 26). Kayangl would genera l ly be access ib l e from Koror except in rough weather. There was some d i f f i c u l t y a t f i r s t because of a r ecen t ly i n s t i t u t e d v i l l a g e pol icy of charging a l l v i s i t i n g Americans $50, but t h e requirement was waived a f t e r the D i s t r i c t Administrator explained t h e value of t h e research program t o t h e i s l ands . Three dives were made on Kayangl ; in t h e shallow lagoon, and on the o u t e r leeward and windward s lopes. 5. Kayangl lagoon The f i r s t dive was on the west (leeward) s i d e of t h e lagoon near the en t rance channel (Fig. 30) . The sandy bottom, 4-6 m deep, becoming sha l lower toward t h e r e e f , separates t h e abundant pa tch ree fs and c o r a l heads. These a r e co ra l - covered f rom t o p t o bottom, and show v e r y g r e a t spec ies d i v e r s i t y . The area i s t y p i c a l o f s h a l l o w P a c i f i c lagoons. 6. Kayangl leeward o u t e r r e e f The leeward o u t e r r e e f o f Kayangl ( F i g . 30) i s topped by a 50 m wide f l a t o f ca lcareous a lgae, l e a d i n g t o a r e e f edge w i t h l u s h c o r a l growth. There i s a s teep d r o p - o f f t o 10 m depth, s t i l l w i t h v e r y good c o r a l coverage and h i g h spec ies d i v e r s i t y , f o l l o w e d by a g e n t l e s l o p e t o 40 m w i t h Pachyser is dominant. The s l o p e f l a t t e n s between 40 and 50 m, and becomes v e r y sandy beyond 50 m. The temperature decreases markedly w i t h depth, and t h e wa te r becomes i n c r e a s i n g l y murky. The l i v i n g r e e f ends between 50 and 60 m. 7. Kayangl windward o u t e r r e e f The e a s t s i d e o f Kayangl i s ve ry exposed, making i t i m p o s s i b l e i n most weather t o anchor a smal l boat . The windward o u t e r ree f (no diagram) commences o f f t h e i s l a n d s w i t h a b road a1 ga l - tu r f - covered f l a t , and then drops t o a p l a t f o r m about 1000 m wide and 7-10 m deep w i t h l i t t l e c o r a l growth. Th is was n o t f o l l o w e d t o t h e d r o p - o f f . The l a c k o f a developed windward c o r a l community and t h e s h a l l o w lagoon p r e v e n t t h i s f r o m be ing as d e s i r a b l e a s i t e as l o g i s t i c s and d i v e r s i t y m i g h t i n d i c a t e . Yap (g030'N, 138O05'E), as a d i s t r i c t c e n t e r , i s a c c e s s i b l e ( 3 f l i g h t s p e r week f rom Guam), and has e x c e l l e n t f r i n g i n g r e e f s , espec- i a l l y on t h e n o r t h w e s t s ide , w i t h some lagoon development i n s i d e t h e r e e f (F ig . 31). It i s a h i g h i s l a n d , so t h a t t h e r e a r e c o n s i d e r a b l e t e r r e s t r i a l i n f l u e n c e s , and t h e r e e f s have exper ienced some Acan thas te r i n f e s t a t i o n . I n f o u r d i v e s on t h e west s ide , a s u p e r f i c i a l coun t o f c o r a l s by R. Randal l y i e l d e d o v e r 100 species. P r o f i l e s f rom t h e d i v e s on d i f f e r e n t p a r t s o f t h e r e e f have been combined t o y i e l d a composi te s e c t i o n ( F i g . 32 ) . 8. O f f G o r r o r Two d i v e s were made i n t h e lagoon, on t h e SW s ide , between t h e r e e f and Yap I s l a n d . The f i r s t , o f f Gorror , was i n a s h a l l o w sand f l a t area w i t h t u r t l e grass and occas iona l c o r a l patches o f h i g h c o r a l d i v e r s i t y (approx imate ly 50 spec ies ) . The wa te r was r a t h e r murky. 9. Off N i f The second d i v e , o f f N i f , was i n one o f t h e deep lagoon areas. The su r round ing r e e f w a l l s a r e covered w i t h l a r g e c o r a l s down t o t h e sandy bot tom a t T O m. The r e e f t o p i s exposed a t low t i d e , w i t h an a l g a l t u r f cover and o n l y sma l l specimens o f F a v i t e s . 10. O f f Okau The leeward ou ter r e e f o f f Okau begins w i t h a rocky f o r e r e e f f l a t , 1500 m wide, c u t by long surge channels 2 m deep and 1-2 m wide. Coral coverage increases t o t h e r e e f edge a t a depth o f 7 m, w i t h on l y a few species l ess than i n Palau. The slope then drops s teep ly t o 25 m w i t h good co ra l growth, beyond which a more gent le slope i s dominated by Pachyseris. The water changes a t around 20-25 m; above i t i s warm and c lea r , below cool and tu rb id . Coral growth seems t o end a t around 40 m. 11. M i l Entrance A f i n a l d i ve was made i n the M i l Entrance (no diagram) on t h e west s ide between Yap and Rumung Is lands . The r e e f f l a t on the n o r t h s ide of the channel extends down t o 2 m w i t h good cora l coverage. There i s a d rop -o f f w i t h good co ra l growth down t o 10 m, and a gent le slope w i t h t u r t l e grass deeper down. A s t rong cu r ren t o f t u r b i d water was moving ou t o f the channel, which conta ined many l a r g e sharks. Although the r e e f q u a l i t y observed was high, t h e t e r r e s t r i a l in f luences, d is tance o f the r e e f f r o n t from shore, and shal low l i m i t t o c o r a l development are major disadvantages. There i s a l so more chance o f an tagon is t i c f e e l i n g s aga ins t Americans on the p a r t o f t h e l o c a l popu la t ion i n d i s t r i c t centers. U l i t h i and Woleai It was n o t poss ib le dur ing t h e sho r t survey t o arrange t ranspor ta t i on t o U l i t h i o r Woleai A t o l l s i n t h e Yap d i s t r i c t , bu t from o the r evidence and from conversat ions w i t h c h i e f s from both areas, some usefu l in form- a t i o n was co l l ec ted . U l i t h i (lOOOO'N, 13g045'E) i s a l a r g e a t o l l nor theas t o f Yap. It has an a i r s t r i p w i t h weekly Coast Guard f l i g h t s from Guam o r the p o s s i b i l i t y o f cha r te r f l i g h t s ($500-$650 f o r 5-7 passengers, one day round- t r i p ) . There i s a l so sh ip serv ice from Yap every two weeks. There are some sunken ships i n the lagoon, and o the r remnants o f war damage, as w e l l as an increas ing c iguatera problem, bu t w i t h the l a r g e s i ze o f t h e a t o l l undis turbed areas must remain. Th is i s one o f very few r e l a t i v e l y undisturbed at01 1s d i r e c t l y accessib le by plane. Woleai (7"201N, 143'45'E) i s a small, t o t a l l y undisturbed a t o l l p resen t l y inaccess ib le but w i t h an abandoned Japanese a i r s t r i p . If the a i r s t r i p were ever repa i red t h i s cou ld be an important s i t e t o consider, even though l o g i s t i c s would be somewhat more d i f f i c u l t . E a r l i e r suggestions t h a t the a i r s t r i p might be res tored soon have n o t been confirmed. A. Antonius d ived on t h e ree fs dur ing t h e 1969 Acanthaster- survey, and they appeared t o be w e l l developed. Ponape Ponape (70001N, 1580001E) i s another d i s t r i c t center, c o n s i s t i n g o f a large volcanic island and several smaller islands surrounded by a barr ier reef (Fig. 33). The reefs are generally subjected t o consider- able t e r r e s t r i a l influences. There are three f l i g h t s weekly from Guam t o Ponape and two from Hawaii, making t h i s the most accessible of the Caroline Islands. Not f a r from Ponape, however, are two small a t o l l s , A n t and Pakin. A n t i s an a to l l of moderate s i ze only 10 miles from Ponape, while Pakin, somewhat smaller, i s 30 miles away. Both can be reached by small boat from Ponape within 3 hours. One dive was made on the outer slope of each a t o l l , and both had excellent reef development. 12. A n t The dive was on the leeward side j u s t beyond the northern t i p a t the northernmost i s l e t (Fig. 33). The reef c r e s t a t t h i s point i s 100 m wide, with a fore reef f l a t 200 m wide, extending down to the reef edge a t 10 m (Fig. 34). Coral growth begins a t 2 m depth, 30 m offshore, increasing t o 90 percent coverage a t 50 m offshore, and 100 percent a t 100 m out. From the reef edge there i s a steep slope down t o 30 m depth with a coral cover of over 100 percent because of the overlapping table Acroporas which a re dominant. The slope continues down to 50 m w i t h interspersed sand cover increasing t o 50 percent, although coral development continues much deeper. The water was warm and c l ea r , w i t h no change in temperature. A n t i s an excellent ree f , with a deep entrance, a good drop-off, and deep coral development. 13. Pakin The dive s i t e a t Pakin was near the center of the northern reef (Fig. 33) , i n an exposed though not windward area. The 80 m wide reef c r e s t merges w i t h a sloping fore reef 50 m wide and 7 m deep a t the edge (Fig. 35). The f i r s t 20 m i s bare of coral cover, b u t coral density increases t o the rugged edge, w i t h headshaped Porites dominant. From there an almost ver t ical slope descends as f a r a s could be observed, w i t h a good coral cover of large specimens and a considerable amount of algae. Sand patches begin a t 50 m , b u t a t 60 m the coral cover i s s t i l l 30 percent and continues t o the l imi t of v i s i b i l i t y . Again there was c lear warm water with no temperature change w i t h depth. The reef i s excellent fo r research, w i t h a steep drop-off and coral development t o beyond normal SCUBA range. Six gray sharks were observed. A n t has a single owner, an ardent conservationist , which might make arrangements fo r a long-term project d i f f i c u l t . Pakin has no channel in to the lagoon a t present, b u t there are government plans t o open one and to build a pier in the lagoon. There would not appear t o be any d i f f i cu l ty in locating a f a c i l i t y there , and log is t i ca l and construction support on Ponape seems excellent . One dive was a l so made on the Ponape barr ier reef (see below), b u t i t was disappointing. 14. Mant Passage The brief dive in the Mant Passage, a northeast entrance through the windward wide of the Ponape bar r ie r reef revealed a smooth rocky f l a t a t 4 m depth with 20-30 percent coral coverage and low species divers i ty . The water was rather turbid , and one gray shark 2.5 m long was seen. The frequent heavy r a in fa l l s i n Ponape might cause technical problems for a research program, as well as affecting the reef . Arno and Majuro Majuro (7'05'N, 171?10'E) and adjacent Arno (7'05'N, 171?45'E), two large a t o l l s in the southern Marshall Islands, are the only accessible a t o l l s i n the area without government entry res t r ic t ions . Majuro has two weekly f l i g h t s from Hawaii, and Arno can be reached eas i ly from Majuro. Majuro has been reported as being disturbed, while Arno i s re la t ive ly untouched b u t dives on both a t o l l s revealed rich reefs with an excel l en t configuration. 15. Kinajon (Arno) The outer reef off Kinajon on Arno i s on the sheltered south side (Fig. 36). The rocky fore reef f l a t 100 m wide develops good coral coverage and surge channels towards i t s edge a t 10 m depth (Fig. 37). A steep slope with Porites heads dominant drops t o 30 m where the angle of slope lessens and sand patches appear. Pachyseris then becomes dominant. A t 60 m the coral cover i s s t i l l 20 percent and continues as f a r as can be seen. The water was warm and exceptionally c lear ( the water surface was v i s ib le from 60 m ) , with no change i n temperature. The reef in general seemed excellent; on the other side where the deep entrance i s , i t i s reportedly even be t te r developed. 16. Laura (Majuro) On Majuro the reef was surveyed off Laura, a sheltered location on the southwest side (Fig. 36). The 60 m wide fore reef consisted of 20 m of bare surface, 20 m w i t h a dense algal cover, and 20 m with corals , mainly Acro ora, leading t o the rugged reef edge a t 5-8 m depth (Fig. 38 r"- Deep surge channels cut in to the reef . There i s no real drop-off, b u t a gently rounded slope with valleys and ridges perpendic- ular t o the shore, and a l so many shore-parallel steps. The coral cover decreases from 90 percent a t the edge (5-8 m) t o 60 percent a t 12 m , t o 50 percent a t 20 m w i t h increasing algal cover. The slope gets steeper with depth, becoming ver t ical a t 40 m , where the coral cover i s 20 percent. This s i t e i s accessible by road from Majuro, and has good reef features and c lear water. Fishes were very abundant, b u t no sharks were observed. AMERICAN SAMOA* The Samoa Islands (14O15'S, 170?00'W) a re a chain of volcanic *Field survey: September 7-12, 1971 by A. L. Dahl and A. Antonius (ass is ted by S. Ritterbush). i s l a n d s w i t h f r i n g i n g c o r a l r e e f s i n t h e South P a c i f i c . Whi le r a t h e r f a r f rom t h e c e n t e r o f c o r a l d i v e r s i t y , t h e i r ready a c c e s s i b i l i t y by d i r e c t f l i g h t f r o m Hawai i and t h e s t r o n g i n t e r e s t o f t h e l o c a l govern- ment warranted d e t a i l e d examina t ion o f t h e r e e f s . The Manua I s l a n d s a r e remote f r o m t h e main i s l a n d o f T u t u i l a and can be reached by government o r commercial b o a t i n about 8 hours . There has a l s o been an i n t e r m i t t e n t f l o a t p l a n e s e r v i c e . Because t r a n s p o r t a t i o n i s d i f f i c u l t i t was o n l y p o s s i b l e t o d i v e on t h e r e e f s i n t h e v i c i n i t y o f t h e main anchorage o f each i s l a n d . S ince t h e r e e f s i n t h e Manua I s l a n d s were n o t t r a n s e c t e d no diagrams a r e g i v e n . 17. Tau (Manua) A t Tau t h e d i v e was made o f f t h e n o r t h w e s t s i d e ( F i g . 39) , a l e e - ward b u t exposed area, about 300 m o f f s h o r e . There i s a r o c k y f l a t w i t h huge bou lde rs a t 20-25 m depth . Few c o r a l spec ies a r e p resen t , m a i n l y P o r i t e s and P o c i l l o p o r a , and specimens a r e sma l l and scarce. 18. Olosega (Manua) The d i v e o f f Olosega was on t h e s h e l t e r e d leeward ( w e s t ) s i d e about 200 m o f f s h o r e , a t 20-25 m d e p t h ( F i g . 39 ) . The b o t t o m i s rocky , w i t h huge b l o c k s , f o r m i n g v a l l e y s and r i d g e s . Cora l cover i s more e x t e n s i v e and w i t h more spec ies t h a n Tau, w i t h huge coheren t c o l o n i e s o f t a b l e - 1 i ke Acropora and P o r i t e s ( l o b a t a ? ) . Many a1 cyonar ians a r e a l s o p resen t . 19. Ofu (Manua) The anchorage o f Ofu i s on t h e west s i d e i n a v e r y s h e l t e r e d l o c a t i o n (F ig . 39), 200 m o f f s h o r e f r o m A lau fau , where t h e wa te r i s 10-20 m deep. The bo t tom has marked t o p o g r a p h i c r e l i e f , w i t h sma l l c o r a l s and r e d a l g a e on t o p o f t h e e l e v a t i o n s , dense c o r a l cover and good spec ies d i v e r s i t y on t h e w a l l s , and w h i t e sand i n t h e bot toms of t h e t roughs . The w a t e r was c l e a r and warn. F ishes were abundant, b u t no sharks were observed. Two d i v e s were made on t h e f r i n g i n g r e e f s on t h e n o r t h and sou th s i d e s o f T u t u i l a , t h e main i s l a n d o f American Samoa. 20. Leone Bay ( T u t u i l a ) A t Leone Bay on t h e windward sou th west s i d e ( F i g . 39), t h e r e e f was surveyed o u t t o 350 m f r o m t h e shore nearLogologo P o i n t (F ig . 40 ) . The r e e f s t r u c t u r e i s i r r e g u l a r , somewhat resemb l ing a spur and groove system, w i t h a s h a l l o w r e e f f l a t , and then l a r g e r e e f patches i n deeper water , e x t e n d i n g down t o 25 rn. The c o r a l cover i s v e r y v a r i a b l e , some- t imes a lmos t 100 pe rcen t , as a t Ofu, b u t many h e l i o p o r a s a r e a l s o p resen t . A l a r g e f l a t o f w h i t e comp le te l y d e t r i t u s - f r e e coarse sand occurs a t 15 m depth . The w a t e r was warm and c l e a r , w i t h many f i s h e s and no sharks . 21. Ogegasa Point (Tutuila) On the north side of Tutuila, a t Ogegasa Point (Fig. 39) , there i s a ver t ical basal t ic rock slope from the surface t o 3 m , followed by a rocky f l a t with small scattered corals extending 50 m offshore to a depth of 7 m (Fig. 41). From here a slope w i t h spur and groove configurations drops t o 15 m and then a very steep slope down to 30 m , ending i n an extensive sand f l a t . This slope has the best coral cover- age and r iches t species divers i ty of a l l the Samoan s i t e s observed. The water was warm and s l igh t ly turbid , perhaps from a recent r a in fa l l . Again the area was r ich in f ishes b u t lacked sharks. Apparently there are no good near-shore drop-offs around the islands of American Samoa. I t was not possible t o reach Rose Atol l , as the seaplane was damaged shor t ly before our a r r i va l , and time and weather precluded boat transportation. Discussions with the Office of Marine Resources which recently surveyed the at01 1 , indicated t ha t the land area was inadequate fo r any f a c i l i t y , so t h a t i t could only be used fo r shor t v i s i t s . CONCLUSIONS The overall evaluation of the s i t e information leads t o the following conclusions fo r the Smithsonian-planned programs. In the Caribbean, the l og i s t i c problems and unique character of the San Blas Islands, the disturbance and pol i t ical problems a t Discovery Bay, and the poor reef s t ructure of St . Croix and Acklins Island l e f t Glover's Reef, Brit ish Honduras as the preferred s i t e . In the Pacific no f inal decision was possible without fur ther f i e l d surveys, b u t several areas showed good potent ia l , including Pakin, Ul i th i , and Arno. Ulithi was not vis i ted by a survey team, and so requires fur ther examination. A more deta i led study of areas in F i j i and the Gilbert and Ell ice Islands could a l so be productive. However, once the s c i en t i f i c s u i t a b i l i t y of a s i t e was determined, i t would s t i l l be necessary t o negotiate with the local inhabitants fo r space and permission t o work on t h e i r reefs . I t i s important t o remember t ha t the areas reported on here were selected and described i n accordance with the specif ic program c r i t e r i a l i s t e d i n the introduction, not a l l of which would necessarily apply t o other projects. We hope t h a t others searching fo r a su i tab le location fo r a coral reef research program will be able t o use t h i s information,with due caution fo r i t s l imita t ions , in se lect ing a s i t e most appropriate t o t h e i r needs. ACKNOWLEDGEMENTS We should p a r t i c u l a r l y l i k e t o acknowledge t h e support of the Environmental Sciences Program a t t h e Smithsonian I n s t i t u t i o n and of t h e In terna t ional Decade of Ocean Exploration o f f i c e of the National Science Foundation (Grant GX-28676). In add i t ion , the cooperation and support of t h e governments of t h e Bahamas, Br i t i sh Honduras, t h e U.S. Trust T e r r i t o r y and American Samoa were g r e a t l y apprec ia ted , as was t h e a s s i s t ance of t h e West Indies Laboratory of Fa i r le igh Dickenson Universi ty on S t . Croix, t h e Discovery Bay Marine Laboratory on Jamaica, t h e Smithsonian Tropical Research I n s t i t u t e in Panama, and the Universi ty of Guam. The ind iv idua l s who cooperated in t h i s survey, e i t h e r as p a r t i c i p a n t s on survey teams, con t r ibu to r s of ques t ionnai res o r s i t e suggest ions, members of t h e CITRE and IMSWE programs, o r supp l i e r s of local advice and support a r e too numerous t o mention although many a r e acknowledged a t appropr ia te places i n the r epor t . REFERENCES CITED Cloud, P. E. 1959. Geology of Saipan, Mariana Is lands . 4. Submarine topography and shoal-water ecology. U.S. Geol. Survey Prof. Paper 280-K: 361 -445. El 1 i o t , H . 1971. Pac i f i c oceanic i s l ands recommended f o r des ignat ion a s i s l ands f o r science. South Pac i f i c Commission. Regional Symposium on Conservation o f Nature--Reef and lagoons. SPC/RSCN/ WP. 17. 26 Ju ly 1971. 16 p. Glynn, P . W. In press . Aspects of t h e ecology of coral r e e f s i n the western At l an t i c region. In R . Endean and 0. A. Jones (eds. ), Biology and Geology of C o r a Reefs. 2 vols . New York, Academic Press. Goreau, T. F . 1959. The ecology of Jamaican coral r e e f s . 1 . Species composition and zonation. Ecology .40: 67-90. Goreau, T. F . , and J . W . Wells. 1967. The shallow water S c l e r a c t i n i a of Jamaica: revised l i s t o f spec ies and t h e i r v e r t i c a l d i s - t r i b u t i o n range. Bull. Marine Sc i . 17: 442-453. Johannes, R . E. -- e t a l . 1972. The metabolism of some coral r ee f communities: a team study of n u t r i e n t and energy f l u x a t Eniwetok. BioScience 22: 541-543. Kinzie, R . A. 1970. The ecology of t h e gorgonians (Cnidaria , Oc tocora l l i a ) of Discovery Bay, Jamaica. Unpublished Ph.D. t h e s i s , Yale Universi tv. " Laborel, J . 1967. Madreporaires des c8 te s du Br6si l . Deuxisme th6se: 1 'Univers i te d' Aix-Marseil l e . Land, L . S . , and T. F. Goreau. 1970. Submarine l i t h i f i c a t i o n of Jamaican r ee f s . J . Sed. Petrology 40: 456-462. Macintyre, I . G . 1972. Submerged r e e f s of Eastern Caribbean. Amer. Assoc. Petroleum Geologists Bull. 56: 720-738. Meyerhoff, H. A. 1926. Geology of t h e Virgin I s l ands , Culebra and Vieques: Physiography. N . Y. Acad. Sci . S c i e n t i f i c Survey of Puerto Rico and the Virgin Is lands 4: 72-219. Newell, N. D . , J . K. Rigby, A. J . Whiteman, and J . S. Bradley. 1951. Shoal -water geology and environments, ea s t e rn Andros Is1 and, Bahamas. Bull. h e r . Museum Nat. History 97: 1-30. S t e h l i , F. G . , and J . W . Wells. 1971. Diversi ty and age pa t t e rns i n hermatypic cora ls . Syst. Zoo1 . 20: 115-126. S toddar t , D. R . 1962. Three Caribbean a t o l l s , Turneffe I s l and , ~ i g h t h o u s e Reef and Glover ' s Reef, ~ r i t i s h - ~ o n d u r a s . Atol l - ~ e s . Bull. No. 87. 151 D. ~~~ . ~ . - . ~ t o r r , ~ . F. 1964. ~ c b i o ~ ~ and oceanography of t h e cora l - reef t r a c t , Abaco Is land , Bahamas. Spec. Pap. Geol. Soc. America 79. 98 p. Tsuda, R. T. (ed.). 1971. S ta tus of Acanthaster planci and cora l r e e f s in t h e Mariana and Carol ine I s l ands , June 1970 t o May 1971. Universi ty of Guam, The Marine Laboratory, Technical Report No. 2, October 1971. 127 p. Wells, J . W . 1969. Aspects of P a c i f i c coral r e e f s . Micronesica 5(2) : 317-322. Whetten, J . T. 1966. Geology of S t . Croix, U. S. Virgin Is lands . Geol. Soc. Amer. Memoir 98: 177-239. Site name Ocean area General location Longitude Latitude Charts General references OIsland: acontinental: OAtoll; a~ringing reef on Jvolcanic or Dother base; UBarrier reef. Description of site Features: Adequate comparable reef area for sampling (perhaps 1 km frontage) Well developed reef flat for drilling Reef development to at least 50 m depth Width across reef less than 300 m Reef near enough to shore to permit shore-based instrumentation Reef undergoing active construction Considerable species diversity Reef not obviously unique Major terrestrial influences absent Reasonably continuous reef accumulation Current and tidal flow patterns permitting cross-reef metabolic studies Varietv of subsidiarv site tvoes in area No Notes Undrs~urbcd by dcvolopncni, :~iascrcpi~~.c siorxs, vsr, or pc!.lu~l~n Prcbsblli iy oi re,, ;ln:.ny .:,dlsLurbed Weather permitting year-round work Accessible, within 1 day's travel of a major airport and harbor facility Anchorage and landing for work boats Accommodations available Research soace available Buildings &ailable for conversion Electricity Fresh water Local suppliers (food, fuel, building materials) Research vessels available in general area Governmental authority Possible attitude toward project Previous scientific work (include references) Aeria l Photoqraphs (Source) Other notes Information supplied by Date A sketch map on the reverse would be helpful., Indicate p a l e or approximate distances if possible. (Also i d e a l i z e d c r o s s seCt1OnS ~ 7 t h r e e zona t ions ) F igu re 1 - QUESTIONNAIRE FOR SITE SURVEY The Bight of Acklins Figure 2 - ACKLINS ISLAND 22030'N 74000'W Palrnata Zone Figure 6 - ST. CROlX - SITE 1 - BANK BARRIER REEF OFF TAGUE BAY (data supplied by L. Firth, D. Jennus and N. B. Ogden) ,.-? / D e e ~ Fore Reef Slope .:.. I , .. I I Lower Sill Reefs .... . . . . . . . . . . .... . . ... '.. . / / ( -.. - -. . . Fore Reef Slope .-. .... .-... - - . . . . _ . . . / ,--; I.--.. . . 0 500 Upper Sill Reefsa, ' ; .. . j i . . ,: ... :.. . . Meters Figure 7 -D ISCOVERY BAY (after Land, unpublished) 18030'N 77025'W Figure 9 - GLOVER'S REEF (modified after Stoddan, 1962) 16050'N 87050'W Meters 500 M e t e r s 4 -dominant on the fore-reef \ 0 -dominant on the reef-edge A- tdominan t on the slope 1 10 x Vertical Exaggeration GLOVER'S REEF Figure 10 - TYPICAL WINDWARD REEF 5 500 Meters 4 Acropora palmata l o x Vertic esiderastrea s- *Diploria strigosa -- *Diploria clivosa -- lsophyllia sinuosa Madracis decactis -- Favia fragum -- AAgaricia agaricites AAgaricia tenuifolia AMontastrea cavernosa Porites astreoides Meandrina meandrites Solenastrea bournoni Dichocoenia stokesi + - dominant on the fore-reef *- dominant on the reef-edge A- dominant on the slope al Exaggeration GLOVER'S REEF Figure 11 -TYPICAL LEEWARD REEF - - - - - - sand Colpophyllia natans Acropora cewicornis Siderastrea siderea Meandrina meandrites A& sp. Porites porites -- Millepora sp. Porites astreoides Algal covered rock pavement. Small colonies of Porites astreoides and Agaricia sp. Diadema are abundant Thalassia six beds. Small coral colonies. sideraStrei radians ( in balls), Manicina areolata and Porites branneri -- Sediment Floor I Large head-coral I or branchinpcoral I zone I I I Finaer or I Rock I bladkd-coral I pavement I zone I I I I Thalassia I beds I SAN BLAS ISLANDS, PANAMA - OFF SAIL ROCK Figure 14 - PROFILE A - A ' eters This depth interval is characterized by an abundance of Agaricia (up to 6 species). Other significant corals include - Diploria strigosa -- Meandrina meandrites Mycetophyllia lamarckiana Montastrea annularis Madracis decactis -- Colpophyllia natans Stephanocoenia michelinii ------- - The dominant coral below 20 meters is Agaricia lamarcki Other significant corals include Montastrea cavernosa Madracis decactis -- Siderastrea + Stephanocoenia michelinii Mussa angulosa -- Scolymia cubensis Scolymia lacera -- Mycetophyllia @ Manicina areolata -- SAN BLAS ISLANDS, PANAMA - OFF SARDINGAN POINT Figure 16 - PROFILE 6-B' Scattered large Acropora -a colonies. Other corals include Porites porites -- Acropora ce~icornis Agaricia sp. - Montastrea a- - - - - - - - a . ~ - f l o 0 ~ : ! Poorly developed buttresses, composed mainly of Montastrea annularis and Colpophyllia m. Some not well defined sand channels. Agaricia sp. also dominant. I -, . . -.p..: ..I. A Halimeda-rich sand occurs between platy growths of Montastrea annularis, Agaricia lamarcki and Porites sp. Other corals include Montastrea cavernosa Mycetophyllia sp. Sideracrrea s i d e r ~ Platy growths of Agaricia sp. dominate. - Scattered heads of Montastrea cavernosa Mycetophyllia sp. Scolymia sp. SAN BLAS ISLANDS - OFF SALAR Figure 18 - PROFILE C-C' ster Acropora palmata and Montastrea a-s dominant - Gradually being replaced seaward by a Montastrea -s, Colpophyllianatans and Acropora cervicornis community I - between and under the coral ledges Antipatharians are very common. 3 - 3 0 Isolated coral-rock blocks Agaricia undata -- Agaricia agaricites Agaricia larnarcki Montastrea cavernosa . -- , Mycetophyllia aliciae Large sponges . , . . . . . . . SAN BLAS ISLANDS - OFF SALAR Figure 19 - PROFILE D- D' Verv slight seaward dip in this zone which lacks sand channels. . - Dominant corals are Agaricia sp., Porites porites, Montastrea -s. Acropora palmata and Acropora cervicornis. Millepora sp. is also abundant. Buttress and sand channel zone. Buttresses are composed primarily of Montastrea a- and Colpophyllia -. Porites astreoides and Mycetophyllia larnarckiana are also abundant. G z 5 T Agaricia sp. corals are dominant on this steep slope. Other corals - include Montastrea annularis, Montastrea cavernosa, Madracis decactis, Porites sp. and Scolymia sp. - Crustose coralline algae and sponges are abundant both between and under the platy coral colonies Halimeda is also abundant. - - - - - - - - - - - Sand-mud bottorn,scattered algal growtns and coral heads. Agaricla sp. Madracis decactis - -- Montastrea cavernosa Porites sp. Siderastrea siderea Scolymia sp. , SAN BLAS ISLANDS - OFF SALAR Figure 20 - PROFILE E - E' . . :..;:: 3 . .. . ,( '.' . . . . . . . . .: ,.., " . . .; .:.>>,.. ,. I " ' .a x! .* . . .< j:., p : q . . . . . .:re ; . . . .! @ . . , i.: . . . . .?& x>"... . . . . . *?$- .' .: _ . . . . , , *$ ; ,:<,,a: . ete 3 - 1 - 1 - . , 3 - Crustose coralline algal coated coral rock. Scattered coral heads of Acropora palmata, Montastrea -s, Siderastrea siderea, Porites astreoides and Manicina areolata. -- - - - - - - -- - - 3fi + /;;a- p "<> ,l,p-w - There are small sand patches in this zone which is dominated by :: porites. Other corals include Diploria - - labyrinthiformis, Acropora cervicornis, lsophyllia sinuosa. Miltepora sp. i s abundant ------ - - - - - - - - Below a depth of about 10 meters there is a great diversity of corals. Down to a depth of 20 meters the dominant corals are Colpophyllia natans, Montastrea annularis, Diploria strigosa and Mycetophyllia sp. - There is a marked decrease in the slope of the sea floor a t 20 meters, and corals occur in isolated masses surrounded by a sand-mud bottom. The dominant corals here are Montastrea annularis and Siderastrea siderea. Other common corals in the 10-30 meter depth range include Stephanocoenia michelinii, -s sp., Dichocoenia w, Dichocoenia s-s, Montastrea cavernosa, Agaricia tenuifolia, Madracis d-s, Mycetophyllia lamarckiana, Eusmilia fastigiata, Mussa angulosa, Meandrina meandrites, Scolymia lacera, Scolymia cubensis and Helioseris c-. --- , SAN BLAS ISLANDS, PANAMA - OFF OGOPUOUIP Figure 23 - PROFILE F - F' Algal Ridge. Nodular on sides and smooth on the surface (nodular --growths coalescetoform the smooth surface). Distinct surge channels and blow holes. Exposed about 0.6 meters above the low tide sea level. Outer Trough. i n deep areas there are abundant corals (=s astreoides, Agaricia sp. and Diploria -sp.) and Diadema -sp. In shallow areas the bottom is covered with - Padina sp. Middle Ridge. The surface, which is about 0.3 meters above sea level -- a t low tide, is extensively pitted and etched. This is in marked contrast to the smooth rock floors of the troughs or tidal pools. Sampling indicated that this ridge is dominantly composed of crustose coralline algae. Inner Trough. Shallow rock bottom covered with algae, mainly Caulerpa -- - sp. and Halimeda sp. over crustose corallines. Porites porites is common. -- Inner Ridge. As in the case of the Middle Ridge the surface is -- extensively etched and pitted. However, this ridge is composed, for the most pan, of corals in a coarse calcarenite matrix. Back-Ridge Area. Corals (Porites porites, Porites astreoides, Agaricia sp.) - - --- Millepora sp. and - Diadema sp. thrive in these shallow waters shoreward of the ridges. Most coral debris has a heavy crustose coralline coating. (For a detailed description of this algal ridge see Glynn, in press.) SAN BLAS ISLANDS, PANAMA - HOLANDES CAYS ALGAL RIDGE Figure 24 - PROFILE G - G' 2 0 0 M e t e r s -4 C Rock ledge with about a 6' seaward dip Madracis sp. encrusting underside of this ledge. 0 algae (below which there is generally a layer of crustose coralline Acropora =a %. Rippled and well algae). Sponges are abundant (especially boring sponges which Montastrea a-s a rounded coarse commonly cover several sq. meters of the bottom). The rock Siderastrea -a % carbonate sand. pavement has been broken here and there to form % to 1 meter $ ledges. Large cavities have been formed under some of the ledges. Gorgonians are abundant but small. Coral colonies are small and widely scattered. They include Diploria - clivosa, Diploria strigosa, Porites astreoides, Favia fragum, Agaricia sp. - Siderastrea siderea. Manicina areolata and Acropora palmata. cock consists of coral debris in a coarse calcarenite. Fresh nature of skeletal components suggests that this is Post Pleistocene in age. SAN BLAS ISLANDS - OFF HOLANDES CAYS Figure 25 - PROFILE H - H' ATOLL ANGUAR I. Figure 26 - PALAU 7030'N 134030'E 0 0 0 - 0 0 0 , 0 0 w 0 0 r-. Meters rock reef f l a t dense coral growth I hard corals dominant shoots and buttresses, Porites heads dominant - sand increasing . "4 I I I f I I I I I 100 200 300 400 500 600 7 00 8 00 900 Mete rs Figure 29 -SITE 4 - NGEMELIS, WEST SIDE 5 - KAYANGLE - LAGOON Meters 6- KAYANGLE - LEEWARD OUTER REEF . -- 'r- dense coral growth hiah s~ecies diversitv algal flat Meters Figure 30 -SITES 5 & 6 - KAYANGLE Figure 31 - Y A P 9030'N 138005'E - 4 krn Figure 36 - ARNO P1 MAJURO 7005'N 171030'E coral cover I I I 50 100 150 Meters Figure 38 -SITE 16- MAJURO STATE OF KNOWLEDGE OF CORAL REEFS AS ECOSYSTEMS ~ a r i e - ~ 6 1 6 n e Sachet INTRODUCTION The f o l l o w i n g b r i e f rev iew o f c u r r e n t knowledge about co ra l r e e f s as ecosystems was w r i t t e n e a r l y i n 1972, most ly from mate r ia l fu rn ished by the CITRE working groups. I t attempts t o descr ibe the s ta tus of r e e f ecology papers which are p a r t i c u l a r l y re1 evant t o the model i ng approach. No in fo rmat ion on the systematics o f r e e f b i o t a i s inc luded i n t h i s summary, a1 though i n v e n t o r i e s are obvious1 y fundamental t o any b io -eco log ica l work. This rev iew has become out-of -date i n c e r t a i n respects and should, i d e a l l y , be thoroughly rev ised. Time f o r t h i s i s n o t ava i l ab le , and on l y a c e r t a i n amount o f updat ing has been done. The b ib l i og raphy has been gone over very c a r e f u l l y , cor rec ted and very much enlarged. I acknowledge w i t h thanks the work o f D r . Bryce Decker i n t h i s matter, and the help o f former CITRE p a r t i c i p a n t s , e s p e c i a l l y those working i n the Smithsonian. I n i t s present form, and w i t h apologies f o r i t s shortcomings, I be1 ieve t h i s review may s t i l l be o f value, espec ia l l y t o readers n o t thoroughly f a m i l i a r w i t h t h e co ra l r e e f f i e l d , o r those f a m i l i a r w i t h o the r aspects o f cora l r e e f research, e.g. systematics. Since the November 1971 workshop, and the January 1972 complet ion o f the CITRE proposal, the r e s u l t s o f a number o f Symposia and Seminars have been pub1 ished, as we l l as several specia l issues o f j ou rna l s , devoted l a r g e l y t o cora l r e e f research. They demonstrate how a c t i v e and va r ied such research i s , and should be consul ted f o r t o p i c s ou ts ide the scope o f t h i s review, f o r d e t a i l s o f s tud ies o n l y b r i e f l y mentioned here, and f o r more recent developments n o t covered here. Among such volumes can be c i t e d the Proceedings o f the 1969 "Mandapam Symposium" issued i n 1972 (Mukundan and P i l l a i , eds.), those o f the 1972 I n t e r n a t i o n a l Helgoland Symposium, "Man i n the sea," assembled i n a whole volume (24:1973) o f ~ e l g o l g n d e r w issenschaf t l i che Meeresuntersuchungen, the March and June 1973 issues o f the B u l l e t i n o f Marine Science, conta in ing 17 papers " I n Memory o f D r . Thomas F. Goreau," and a recent issue o f A t o l l Research B u l l e t i n (1 66-1 70) devoted t o f i v e papers on Acanthaster i n f e s t a t i o n s . (Manuscript completed March 1974) The r e s u l t s o f an Acanthaster symposium a t t h e Second In te r -Congress meet ing i n Guam, May 1973, have j u s t been p u b l i s h e d i n M ic rones ica (Dec. 1973). A lso j u s t r e c e i v e d i s t h e f i r s t volume o f t h e l o n g a w a i t e d t r e a t i s e on B i o l o g y and geo logy o f c o r a l r e e f s (Jones and Endean, 1973) . Several o t h e r symposia and r e v i e w volumes a r e s t i l l i n press, i n c l u d i n g t h e r e s u l t s o f t h e " F l o a t i n g Symposium" (2nd I n t e r n a t i o n a l Symposium on Coral Reefs, Grea t B a r r i e r Ree f ) , June 1973. H o t o f f t h e p r e s s a t t h e t i m e o f t h e G l o v e r ' s Reef workshop and t h e p r e p a r a t i o n o f t h e CITRE proposal were Regional V a r i a t i o n i n I n d i a n Ocean Coral Reefs ( S t o d d a r t and Yonge, 1971) and t h e Supplement t o I s l a n d B i b l i o g r a p h i e s (Sachet and Fosberg, 1971) . BACKGROUND F o l l o w i n g Char les D a r w i n ' s d e s c r i p t i o n and t h e o r y o f a t o l l o r i g i n (1842, l 9 6 2 ) , a growing c o n t r o v e r s y o v e r t h e " c o r a l r e e f problem" ( i . e . o r i g i n o f a t o l l s and r e e f s ) l e d t o v i g o r o u s debates based m a i n l y on i n f o r m a t i o n c o l l e c t e d f r o m p u b l i s h e d b a t h y m e t r i c c h a r t s and i n s u f f i c i e n t f i e l d d a t a (e.g. Daly, 1910, 1915; Vaughan, 1916a, 1916b; Dav is , 1928) . An e x c e p t i o n was t h e r e s u l t s o f t h e F u n a f u t i e x p e d i t i o n s o f t h e Royal S o c i e t y (1904) and o f t h e A u s t r a l i a n Museum (Hedley, 1896-1 900). Over t h e p a s t 20 years , t h e " c o r a l r e e f problem" c o n t r o v e r s y has abated, p a r t i c u l a r l y s i n c e d r i l l i n g techn iques have made i t p o s s i b l e t o e s t a b l i s h t h a t a t o l l f o r m a t i o n i s indeed a s s o c i a t e d w i t h subs idence- -no tab ly i n s t u d i e s by Ladd and c o - i n v e s t i g a t o r s (Ladd e t a1 ., 1953; Ladd and Sch langer , 1960; Ladd u., 1967) . The enormous volume o f work c a r r i e d o u t o n r e e f s and a t o l l s i n t h e e a r l i e r p e r i o d s and i n r e c e n t y e a r s i s summarized i n l i t e r a t u r e r e v i e w s such as those o f S t o d d a r t (1969a), Ladd ( i n p ress ) and Glynn ( i n p r e s s ) . Ma jo r t o p i c s d i scussed i n p r e v i o u s r e p o r t s i n c l u d e r e e f f o r m a t i o n (Ladd and Tracey, 1949) , geo logy o f r e e f s (Ladd, 1961; F a i r b r i d g e , 1950; Newe l l , l 9 5 g ) , b i o l o g i c a l z o n a t i o n o f r e e f s ( W e l l s , 1954, 1957a, Goreau 1959) , and c o r a l p h y s i o l o g y (Yonge, 1940, 1963, 1968) t o l i s t b u t a few. I n a d d i t i o n , s e v e r a l b i b l i o g r a p h i e s o f c o r a l - r e e f s t u d i e s have been compi led r e c e n t l y (Pugh, 1950; Sachet and Fosberg, 1955, 1971; We l l s , 1957b; Ranson, 1958; M i l 1 iman, 1965). As i n t h e e a r l y e x p e d i t i o n s , most o f t h e work on s p e c i f i c a t o l l s i n t h e l a s t 20 y e a r s has c o n s i s t e d o f d e t a i l e d d e s c r i p t i o n s and i n v e n t o r i e s (Ladd, ed. 1954-date; Sachet 1962b; D i r e c t i o n des Cent res d ' ~ x p e ' r i m e n t a t i o n s n u c l e b i r e s , 1969). A qua l i t a t i v e d e s c r i p t i o n o f t h e c o r a l a t o l l ecosystem was prepared i n 1957 by Fosberg (1961, 1963b). These d e s c r i p t i o n s f u r n i s h an e x c e l l e n t base1 i n e f o r compara t i ve s t u d i e s and a s t a r t i n g p o i n t f o r t h e q u a n t i t a t i v e o b s e r v a t i o n s needed f o r model c o n s t r u c t i o n . I t can be s a f e l y s t a t e d t h a t n e i t h e r t h e qua l i t a t i v e c o n s t i t u t i o n n o r t h e r o l e o f a s i n g l e f u n c t i o n a l component has been c o m p l e t e l y e l u c i d a t e d i n any one r e e f env i ronment . E c o l o g i c a l processes on t r o p i c a l r e e f s have been es t ima ted on occas ion by p r o j e c t i n g da ta f r o m long- te rm o r d e t a i l e d s t u d i e s o f temperate mar ine communit ies (Pa ine and Vadas, 1970; Neman, 1970). An a s p e c t o f r e e f eco logy which has h a r d l y been developed as y e t i s any work p e r m i t t i n g an e s t i m a t i o n o f c a r r y i n g c a p a c i t y f o r e x p l o i t a t i o n by human p o p u l a t i o n s . GENERAL REEF SURVEYS S i n c e D a r w i n ' s r e e f s t u d i e s , t h e r e have been many i m p o r t a n t genera l i n v e s t i g a t i o n s o f t h e n a t u r e o f these s t r u c t u r e s f rom v a r i o u s v i e w p o i n t s . Some o f these were one-man e n t e r p r i s e s , such as those o f A lexander Agass iz a t t h e end o f t h e 1 9 t h c e n t u r y aboard t h e ALBATROSS, i n a l l t h e m a j o r r e e f areas o f t h e wor ld , those o f J . S t a n l e y Gard ine r i n t h e I n d i a n and P a c i f i c Oceans i n t h e f i r s t p a r t o f t h e t w e n t i e t h c e n t u r y , t h a t o f Wood-Jones on Cocos-Keel ing, r e s u l t s pub1 i s h e d i n 1909 and 1910, and t h a t o f G i b s o n - H i l l , a l s o o n Cocos-Keel ing j u s t b e f o r e World War 11. Dav id S t o d d a r t made t h r e e d e t a i l e d geomorphological s t u d i e s of t h e B r i t i s h Honduras r e e f s and cays between 1959 and 1965. No tab le were a s u b s t a n t i a l number o f c o o p e r a t i v e i n v e s t i g a t i o n s i n v o l v i n g peop le w i t h d i f f e r e n t i n t e r e s t s and backgrounds. Among these have been t h e Royal S o c i e t y ' s and t h e A u s t r a l i a n Museum's e x p e d i t i o n s t o F u n a f u t i , E l l i c e I s l a n d s , r e p o r t e d on by Dav id and Sweet i n a volume p u b l i s h e d by t h e Royal S o c i e t y o f London i n 1904 and i n papers e d i t e d by Hedley (1896-1900); t h e G r e a t B a r r i e r Reef E x p e d i t i o n i n 1928-1929, under t h e l e a d e r s h i p o f C.M.Yonge (1 %Oa, 19306, 1931 ) ; t h e Carneg ie I n s t i t u t i o n o f Washington 's Cora l Reef Program, w i t h work e s p e c i a l l y a t t h e Dry Tor tugas L a b o r a t o r y and i n American Samoa and T a h i t i i n t h e f i r s t t h i r d o f t h i s c e n t u r y . I n v o l v e d i n t h i s were e s p e c i a l l y A l f r e d G. Mayor (Mayer) , W.A. S e t c h e l l , and T. Wayland Vaughan; some o f t h e i r p u b l i c a t i o n s a r e l i s t e d i n t h e r e f e r e n c e s . A f t e r World War I 1 came a comprehensive program o f work on B i k i n i and En iwetok A t o l l s i n c o n n e c t i o n w i t h t h e a tomic weapons t e s t s t h e r e (Ladd, 1973) . T h i s work i s p u b l i s h e d as an enormous s e r i e s o f papers c o l l e c t i v e l y known as USGS P r o f e s s i o n a l Paper 260, p a r t s o f wh ich a r e s t i l l appear ing . The USGS-Army Map S e r v i c e Far E a s t su rvey o f t h e N o r t h e r n Marsha l l I s l a n d s i n 1951 -1952 l e d t o many papers, e s p e c i a l l y t h e M i l i t a r y Geography o f t h e N o r t h e r n Marsha l l I s l a n d s and A t o l l Research B u l l e t i n 113 (Fosberg s., 1956, Fosberg and C a r r o l l , 1965), and a r e v i e w by MacNeil (1972) . D u r i n g t h e 1 9 5 0 ' s t h e P a c i f i c Science B o a r d ' s Cora l A t o l l Program s e n t mu1 t i - d i s c i p l i n a r y e x p e d i t i o n s t o Arno A t o l l , Onotoa A t o l l , Raro ia A t o l l , Kapingamarangi A t o l l , and I f a l u k . I n 1958 and 1960, under t h i s same program were two e x p e d i t i o n s t o J a l u i t A t o l l t o s t u d y t h e e f f e c t s o f Typhoon Ophel ia . Repor ts were p u b l i s h e d i n t h e A t o l l Research B u l l e t i n . ORSTOM, t h e French overseas r e s e a r c h o r g a n i z a t i o n , sponsors a m u l t i d i s c i - p l i n a r y s t u d y o f c o r a l r e e f s around Nosy-~e' , Malagasy. From 1967 t o t h e p r e s e n t t i m e t h e Royal S o c i e t y has sponsored c o n t i n u i n g work on A ldabra A t o l l and o t h e r southwest I n d i a n Ocean c o r a l i s l a n d s and r e e f s . A smal l r e s e a r c h s t a t i o n i s now m a i n t a i n e d on A ldabra t o c o n t i n u e t h i s work. P r e l i m i n a r y r e s u l t s a r e p resen ted i n P h i l o s o p h i c a l T r a n s a c t i o n s o f t h e Royal S o c i e t y o f London v o l . B, 260, 1971. A comprehensive d i s c u s s i o n o f t h e geo logy o f A ldabra has been p u b l i s h e d r e c e n t l y ( B r a i t h w a i t e A. 1973) . The Fonda t ion S inger -Po l ignac d u r i n g t h e 1 9 6 0 ' s sponsored a number o f e x p e d i t i o n s t o New Caledon ia and French Po lynes ia , and m a i n t a i n e d a r e s e a r c h vesse l i n t h e area f o r t h i s purpose. R e s u l t s a r e p u b l i s h e d i n Cahiers du P a c i f i q u e and i n a s e r i e s o f b e a u t i f u l l y i l l u s t r a t e d ~ 6 m o i r e s . I n c o n n e c t i o n w i t h t h e n u c l e a r t e s t s on Mururoa, Tuamotus, t h e French government sponsored an e x t e n s i v e s e r i e s o f i n v e s - t i g a t i o n s i n t h e Tuamotus, e s p e c i a l l y Mururoa, and i n C l i p p e r t o n I s l a n d . Repor ts a r e p u b l i s h e d i n Cah ie rs du P a c i f i q u e and i n a s e r i e s o f d u p l i c a t e d papers on C l i p p e r t o n I s l a n d . I n 1971, a smal l r e s e a r c h s t a t i o n ( "Antenne de T a h i t i " ) o f t h e ~use 'um N a t i o n a l d ' H i s t o i r e N a t u r e l l e , P a r i s , was c r e a t e d i n Moorea, near T a h i t i , t o c o n t i n u e such research , and an i m p r e s s i v e l i s t o f papers i s a l r e a d y a v a i l a b l e , severa l o f which a r e c i t e d be low ( C h e v a l i e r , Den izo t , S a l v a t , e t c . ) The mar ine s t a t i o n o f Endoume-Marsei l le has f o r some y e a r s sponsored c o r a l r e e f i n v e s t i g a t i o n s a t t h e mar ine s t a t i o n a t Tu lea r , Malagasy, comprehensive r e p o r t s have been p u b l i s h e d i n s p e c i a l i s s u e s o f t h e Recuei l des Travaux s e r i e s and l a t e r o f ~ e ' t h y s . A genera l d e s c r i p t i o n o f t h e r e e f s o f Madagascar was p resen ted by P ichon (1972) . The U n i v e r s i t y o f Hawai i r e c e n t l y a c q u i r e d a s t a t i o n on Fanning I s l a n d i n t h e L i n e I s l a n d s and d u r i n g 1969 s taged a b road-sca le NSF- sponsored i n v e s t i g a t i o n o f t h e Fanning I s l a n d s r e e f s . The r e s u l t s have been published i n P a c i f i c Sc ience ( A p r i l 1971 ) and i n a s p e c i a l r e p o r t i s s u e d by t h e Hawai i I n s t i t u t e o f Geophysics (Chave, ed., 1970); t h i s i s a c o n t i n u i n g program, and f u r t h e r r e s u l t s can be expected. S ince 1968, a U n i v e r s i t y o f Hawai i Sea Gran t p r o j e c t t o s t u d y v a r i o u s q u a n t i t a t i v e aspec ts o f r e e f s and r e e f b i o t a i n Kaneohe Bay, Oahu, has been underway. The f i r s t r e s u l t s o f t h a t program have been p u b l i s h e d ( S . V . S m i t h - e t . - a l . , 1973) . The work o f severa l o t h e r surveys i s ment ioned t h r o u g h o u t t h e f o l l o w i n g s e c t i o n s though t h e p r e s e n t d i s c u s s i o n i s by no means e x h a u s t i v e . PHYSICAL ENVIRONMENT Much o f t h e oceanograph ic work c a r r i e d o u t i n t h e t r o p i c a l seas i n t h e p a s t , and e s p e c i a l l y i n t h e l a s t 25 years , i s a source o f env i ronmenta l d a t a on c o r a l r e e f s . One example i s work on s u r f a c e c u r r e n t s , o f s i g n i f i c a n c e i n t h e d i s t r i b u t i o n o f r e e f and i s l a n d b i o t a . S p e c i f i c r e s e a r c h on a t o l l s and r e e f s , however, has been pursued o n l y by a sma l l number o f i n v e s t i g a t o r s and w i t h a l i m i t e d scope. Work a t B i k i n i A t o l l (von Arx, 1954; Munk and Sargen t 1954) has d e l i n e a t e d t h e c i r c u l a t i o n p a t t e r n w i t h i n t h e lagoon t h a t i s e s t a b l i s h e d i n response t o p r e v a i l i n g winds and i n f l u e n c e d by t i d e s , waves, and l o c a l ocean c u r r e n t s . The B i k i n i s tudy, as w e l l as t h e r e s e a r c h a t Fanning I s l a n d ( G a l l a g h e r G., 1971), p r o v i d e s v o l u m e t r i c e s t i m a t e s o f exchanges o f wa te r between lagoon and ocean. Fanning I s l a n d i n v e s t i g a t i o n s a l s o produced measurements o f s a l t and h e a t f l u x e s . The wa te rs o f t h e G r e a t B a r r i e r Reef have been s t u d i e d by Brandon (1973). These s t u d i e s and o t h e r s devoted t o s p e c i f i c p h y s i c a l parameters on r e e f s can c o n t r i b u t e g u i d e l i n e s f o r a n ecosystem model, b u t t h e m a j o r i t y o f oceanograph ic p r o j e c t s on r e e f s have n o t i n c l u d e d c o o r d i n a t e d s t u d i e s o f i n t e r a c t i o n s o f t h e p h y s i c a l env i ronment w i t h the b i o l o g i c a l r e e f communities; see f o r ins tance the work o f Van Dorn, and Vastano and Reid, on Wake I s land . A t o l l meteorology was most ly l i m i t e d , p r i o r t o World War 11, t o sporadic weather records (see f o r instance those used by Sachet, 19571, observat ions on storms and hurr icanes, and the l i k e . Wi th World War I 1 ac t i on , and the atomic t e s t s i n the P a c i f i c , as we l l as the increase i n geophysical research genera l l y , much more i n fo rma t ion i n meteorology and c l imato logy has become ava i l ab le . Weather data are more complete and c o l l e c t e d from more s ta t i ons . Summaries such as those o f R . C. Taylor (1973) and Z ipser and Taylor ( l968) , and volumes o f data such as those prepared f o r U.S. J o i n t Task Force Seven, a r e a few examples among many, which supply important data i n the study o f at011 ecosystems. See a l so s tud ies by Montgomery (1973) and Quinn and B u r t (1970), and the very d e t a i l e d work o f Blumenstock and Rex (1960) on Eniwetok. Even more c l o s e l y r e l a t e d t o the ecosystem approach are papers descr ib ing hurr icanes, typhoons and o the r storms, and t h e i r e f f e c t s on r e e f s and is lands . The work o f Blumenstock and o thers (1958, 1961) on J a l u i t , t h a t o f S toddar t on the B r i t i s h Honduras ree fs (1363, 1969d), and the Solomon I s . (1973), o f Sachet r e l a t i v e t o C l ipper ton (1962b) a re a few examples. More general work on hurr icanes and o ther meteorological phenomena i s inc luded i n the work o f Ramage and others, i n the U.S. Navy At lases (1956-68) and many o the r use fu l sources. A t o l l s have a l so been used as "observat ion p la t fo rms" (Lavoie, 1963) i n atmospheric and o the r s tudies, no tab ly by the Hawaii I n s t i t u t e o f Geophysics and the U.S. J o i n t Task Force Seven (e.g. McCreary 1959). GEOLOGY The geological record o f f o s s i l r ee fs has been the sub jec t o f exhaust ive studies, e s p e c i a l l y by petroleum geologists,and some o f the reviews and b ib1 iographies c i t e d above a r e concerned w i t h f o s s i l reef s tudies, i n c l u d i n g paleoecology. Those references w i l l n o t be d e t a i l e d here (but cf. Ladd, 1957; Laporte 1974). S u f f i c e i t t o say t h a t a vas t amount o f knowledge o f f o s s i l r e e f s has accumulated, from which s tud ies o f modern r e e f s can de r i ve data as we l l as ideas (Hedgepeth, 1957). To a c e r t a i n extent , o f course, the f o s s i l and modern reefs cannot be separated. The modern sect ions o f the r e e f are considered here t o be ma te r ia l deposited i n r e l a t i o n t o present sea l e v e l s , approximately dur ing the l a s t 5,000 years. Since the l a t e 19 th century, modern co ra l r e e f s have been s tud ied by geo log is ts p r i m a r i l y t o p rov ide i n fo rma t ion on p re -ex i s t i ng envi ron- mental cond i t ions documented i n the l i t h o l o g i c a l record o f the geological column. With few except ions these s tud ies have concentrated on desc r ip t i ons o f r e e f components r a t h e r than on the processes respons ib le f o r component format ion. However, much a t t e n t i o n has been pa id t o the i n f l uence o f sea l e v e l changes on the nature o f present-day r e e f s (Fa i rb r i dge 1958; c f . Stoddart 1971 ) . S t u d i e s o f r e e f morpho logy ( rev iewed by S t o d d a r t , 1969a) g e n e r a l l y have i n c l u d e d mapping bo t tom topography a t i n t e r m e d i a t e s c a l e s (1 :50,000) and d e s c r i b i n g t o p o g r a p h i c a l z o n a t i o n and s i z e o f f e a t u r e s i n such terms as mean r e e f w id th , k n o l l d imens ions, e t c . (e.g. Emery -- e t a1 . 1954) . There have been few q u a n t i t a t i v e s t u d i e s o f r e e f f e a t u r e s , a p a r t f r o m t h e work i n Kaneohe Bay, Oahu, by Roy (1970a), a t Fann ing I s l a n d by Roy and Smi th (1971), and i n t h e Ma ld i ves and e lsewhere by S t o d d a r t ( u n p u b l i s h e d ) . Many s t u d i e s concern t h e m ine ra logy , compos i t i on , and t e x t u r e o f r e e f and n e a r - r e e f sediments w i t h a v iew t o u n d e r s t a n d i n g sequences i n t h e g e o l o g i c a l r e c o r d (e.9. Ginsburg, 1956; Neumann, 1965, M a c i n t y r e , 1970) . Recent t r e n d s have been t o examine CaC03 budgets i n a reas o f ca rbona te sed imen ta t ion (Stockman e t a1 . , 1967; Neumann and Land, 1969; Land, 1970; S . V . Smi th u., 1970,1971; S . V . S m i t h 1971a, 1973; Chave e t a1 ., 1971) . Such s t u d i e s have o n l y cons ide red some ( o r even o n l y o n e ) f t h e components i n t h e budget; o r t h e y have i n t e g r a t e d t h e components o f t h e budget a c r o s s a v e r y r e s t r i c t e d env i ronment ; some have n o t even d e a l t w i t h t r o p i c a l r e e f systems. What i s needed now i s t h e g e n e r a t i o n o f a t o t a l c a r b o n a t e budget th rough t ime , i n o r d e r t o d e s c r i b e t h e r e l a t i o n s h i p s among r e e f development, sediment g e n e r a t i o n , sediment t r a n s p o r t , and subsequent d e p o s i t i o n . L i t t l e i n f o r m a t i o n i s a v a i l a b l e on f o r m e r z o n a t i o n s i n v a r i o u s s tages o f r e e f development, y e t work on s p e c i f i c problems a s s o c i a t e d w i t h c o r a l r e e f s has i n d i c a t e d t h a t s i g n i f i c a n t d a t a may r e s u l t f rom expos ing i n t e r n a l r e e f s t r u c t u r e s : Shinn (1 963) i n v e s t i g a t e d t h e o r i g i n o f spu rs i n r e e f s o f f F l o r i d a and subsequen t l y (G insburg a., 1967) s t u d i e d mar ine cementa t ion and i n t e r n a l s e d i m e n t a t i o n w i t h i n r e e f s o f f Bermuda. To da te , r a t h e r l i t t l e c o r i n g has been a t tempted (Mayor, 1924; Cary, 1931 ; Ladd and Sch langer , 1960; S t o d d a r t 1971 ) on modern c o r a l r e e f s , w i t h t h e r e s u l t t h a t t h e i n t e r n a l s t r u c t u r e s and l i m e s t o n e f a c i e s o f these r e e f s a r e l a r g e l y unknown. Most o f t h i s work has i n v o l v e d o n l y one o r a few c o r e h o l e s p e r r e e f . More d e t a i l e d d r i l l i n g w i t h a submers ib le d r i l l t h a t can p e n e t r a t e 2 mete rs i n t o t h e r e e f i s underway on r e e f s o f Jamaica ( L . Land, pe rsona l communication, 1970) . I t has been suggested t h a t l i v i n g c o r a l r e e f s c o n s i s t o f a t h i n veneer o f modern r e e f g r o w t h o v e r o l d e r f o u n d a t i o n s wh ich g e n e r a l l y d i c t a t e t h e i r p resen t -day morpho logy (Newel1 , 1962; S t o d d a r t , 1969a, 1973). However, modern r e e f c o n s t r u c t i o n has been shown t o f o r m mass ive w a v e - r e s i s t a n t frameworks, n o t a b l y i n Jamaica (Goreau, 1961 b, Goreau and Land, 1974) and i n B r i t i s h Honduras (Purdy, 1974) . Numerous workers have i n v e s t i g a t e d g rowth r a t e s (e.g. Vaughan, 1915; Mayor, 1924; Shinn, 1966) o r c a l c i u m carbonate d e p o s i t i o n r a t e s (e.g. , Kawaguti and Sakumoto, 1948; Goreau 1959; Goreau and Goreau, 1959; Goreau, 1961a) o r genera l r e e f CaC03 p r o d u c t i o n r a t e s (e .g . Chave e t a l . 1971; S.V. Smi th g., 1971, S.V. Smith, 1973, Glynn e t a1. 1971), as we1 1 as ske l e t a 1 framework d e s t r u c t i o n by m e c h a n i c a l w d d a r t , 1963; Glynn s., 1965; B a l l e t a1 ., 1967; P e r k i n s and Enos, 1968) and b i o l o g i c a l means ( D u e r d e n n 2 ; O t t e r , 1937; Yonge, 1963; Goreau and Hartman, 1963; Bakus 1964, 1967; Neumann, 1966) . However, c o n c u r r e n t i n v e s t i g a t i o n s o f most of these processes have n o t been c a r r i e d o u t a t a s i n g l e r e e f s i t e i n o r d e r t o determine t h e r e l a t i v e impor tance o f t h e v a r i o u s parameters c o n t r o l l i n g t h e n e t accumu la t ion o f modern r e e f framework. The success o f framework c o n s t r u c t i o n depends l a r g e l y on t h e g rowth r a t e o f r e e f organisms. O ther f a c t o r s a f f e c t i n g c o n s t r u c t i o n i n c l u d e p r e d a t i o n , c o m p e t i t i o n between c o r a l s and o t h e r organisms, v a r i a t i o n i n d i s t r i b u t i o n o f c o r a l and o t h e r spec ies , b i o l o g i c a l e r o s i o n o f s k e l e t a l framework and o f f rame-cement ing agents , mechanical e r o s i o n , and t h e r a t e o f sea l e v e l change. Cement i n f i l l i n g and l i t h i f i c a t i o n of i n t e r n a l sediment i n v a r i o u s c a v i t i e s tend t o nega te t h e d e s t r u c t i v e a c t i o n o f c o r a l b o r e r s . For example, some c o r a l heads c o l l e c t e d from submerged r e e f s i n t h e e a s t e r n Car ibbean have been a lmos t c o m p l e t e l y a1 t e r e d - - t h r o u g h repea ted b o r i n g , i n f i l l i n g , and li t h i f i c a t i o n - - t o a dense m i c r i t i c 1 in iestone ( M a c i n t y r e , 1972) . B i o t a have l o n g been t h o u g h t t o be t h e m a j o r f rame-cement ing agents , b u t i n r e c e n t y e a r s e x t e n s i v e M g - c a l c i t e cementa t ion o f r e e f framework and i n n e r framework has been recogn ized (summarized i n B r i c k e r , 1971) . The r e l a t i o n , i f any, o f b i o l o g i c a l agents t o t h i s cementa t ion i s n o t a l t o g e t h e r c l e a r . The o r i g i n o f magnesian c a l c i t e cement remains unexp la ined, b u t chemica l a n a l y s i s o f r e e f i n t e r s t i t i a l wa te rs may o f f e r some c l u e s t o t h e processes r e s p o n s i b l e f o r i t s p r e c i p i t a t i o n . W i t h i n t h e r e e f ecosystem, i n t e r a c t i o n s between wa te rs and s o l i d CaCO appear t o be l a r g e l y the r e s u l t o f b i o l o g i c a l c a l c i f i c a t i o n processes, o r i h e r a c t i o n s between b i o g e n i c r o c k , d e t r i t u s , and wa te rs separa ted f rom open ocean waters--e.g. , t h e waters w i t h i n t h e i n t e r s t i c e s o f s k e l e t a l f ragments o r t h e r e e f f rame i t s e l f . D i r e c t chemical p r e c i p i t a t i o n and s o l u t i o n r e a c t i o n s o f CaCO i n seawater appear t o be i n h i b i t e d o r p reven ted by i n t e r a c t i o n s of dqsso lved o r g a n i c compounds i n sea w a t e r w i t h CaCO m i n e r a l s u r f a c e s (Chave, 1965; Chave and Suess, 1967, 1970; Suess, 19703. Chemical r e a c t i o n s between seawater and ca rbona te m i n e r a l s may occur , f o r example, i n t h e f o r m a t i o n o f o o l i t h s , grapestone, w h i t i n g s , and beachrock; b u t these r e a c t i o n s a r e , a t best , m i n o r i n a c o r a l - r e e f ecosystem, i f t h e y a r e i n o r g a n i c a t a l l . Two i n t e r a c t i o n s between r e e f wa te rs and b i o g e n i c ca rbona tes appear t o be i m p o r t a n t when c o n s i d e r i n g t h e f l o w o f carbon i n a r e e f ecosystem. F i r s t , r e c e n t r e p o r t s i n d i c a t e t h a t chemical o r b iochemica l p r e c i p i t a t i o n o f CaCO w i t h i n t h e r e e f i s as i m p o r t a n t , o r even more i m p o r t a n t , t h a n b i o g e n i z c a l c i f i c a t i o n w i t h r e s p e c t t o b i n d i n g t h e components t o f o r m a r i g i d , w a v e - r e s i s t a n t framework (see B r i c k e r , 1971) . The second i n t e r a c t i o n between r e e f waters and b i o g e n i c carbonates i s t h e removal o f d i s s o l v e d carbonate f r o m i n t e r s t i t i a l wa te rs o f sediments and d e p o s i t i o n o f CaC03 w i t h i n t h e sediment components (see B r i c k e r , 1971 ) . The impor tance o f t h i s i n t e r a c t i o n i s n o t o n l y t h a t i t r e p r e s e n t s a f l o w o f carbon f r o m seawater i n t o t h e r e e f ecosystem b u t t h a t t h e r e s u l t o f t h i s f l o w d i r e c t l y a f f e c t s t h e c h a r a c t e r and h y d r a u l i c p r o p e r t i e s o f t h e sed iment . In order t o understand f u l l y the processes responsible fo r the precipitat ion of CaCO , both in the framework and within sediment grains, i t i s necessa?y to invest igate the whole reef CaCO system-- namely both water and rock chemistry. By studying the whofe chemical system, i t should be possible to determine whether the cementation process involves only an internal cycling of CaCO , o r whether s ign i f ican t amounts of CaC03 a re derived from external source?. The CO system i n seawater provides a unique l ink between geochemical and biologigal processes. The 1 ink i s par t icular ly important i n coral ree fs , where both ca lc i f ica t ion and organic carbon production-consumption greatly a f f ec t the system. The organic carbon t ransfe r should be mentioned; i t i s suf f ic ien t f o r t h i s paper to point out t ha t (as discussed i n some detai l by Park, 1969) measurement of two parameters in the marine CO system i s suf f ic ien t t o par t i t ion the system changes into inorganic ca803 precipitat ion-solution and organic C production-respiration. Separating changes in deep ocean CO in to solution and oxidation has been undertaken by several authors (g .g . Park, 1968; S . V . Smith, 1971a). Both S.V. Smith (1973) and Kinsey (unpub. data and personal comm.) have studied die1 va r i ab i l i t y in the C02 system in coral ree fs . S.V. Smith (1973) has related t h i s var iab i l i ty to organic carbon pro- duction and u t i l i z a t i on and t o ca lc i f ica t ion . The present major l imita t ion of these short-term studies i s the small amount of change i n the marine COZ Tystem caused by inorganic chemical processes and organic metabolic a t i v i t i e s of the reef communities. Kinsey's studies were performed on water which "aged" over several hours and consequently experienced eas i ly measurable changes in the C03 system. Smith's studies dea l t w i t h water "aged" l e s s than one hour therefore showing barely measurable perturbations i n the C02 system. BIOTIC ENVIRONMENT The biot ic components of the surrounding sea should not be ignored. Perhaps most important i s the plankton, which i s a s ign i f ican t food supply of the corals and many other reef animals. This has been studied w i t h re la t ion to reef animals by A . R . Emory (1968). Small pelagic f i s h and f r y a re also food f o r reef-dwelling carnivorous f i s h , a s well a s fo r sea-birds. Predaceous pelagic f i s h v i s i t the reefs and prey upon the reef f i shes . The same is true of porpoises, other small cetaceans and pinnipeds. Detritus of organic or igin brought to the reefs by water movement i s also an important i n p u t in to the reef system ( N . Marshall 1965). All these biot ic components a re discussed in more deta i l below. TERRESTRIAL ECOSYSTEM To the best of ou r knowledge, the only attempt a t a generalization of the funct ioning of the t e r r e s t r i a l cora l a t o l l ecosystem i s the desc r ip t ion wr i t t en i n 1957 by Fosberg (1961, 1963b). That desc r ip t ion at tempts t o conceptual ize t h e system in a b s t r a c t , non-quant i ta t ive , non-mathematical terms, and t o i n d i c a t e funct ional groups of e n t i t i e s i n t h e system which can be s tudied i n terms of t h e i r r e l a t i o n s h i p s t o the whole. This work was not followed u p except f o r extending i t t o high i s l ands . I t r e su l t ed l o g i c a l l y from the general program f o r a t o l l , s t u d i e s proposed a t two coral a t o l l research symposia sponsored i n 1951 by the Pac i f i c Science Board ( c f . papers i n Atoll Research Bu l l e t in s 1 , 2, 1951, Fosberg ed . ) . Five expedi t ions planned t o provide comparable d e s c r i p t i v e and inventory da ta on a t o l l s of d i f f e r e n t types were c a r r i e d out ( t o Arno, Onotoa, Raroia, Kapingamarangi and I f a luk ) . T h e i r r e s u l t s , a s well a s those of many o the r s tud ie s of land (and marine) a spec t s of a t o l l s were published over the next 20 yea r s i n t h e Atoll Research Bul le t in and elsewhere. An a t tempt a t bringing together t h i s g r e a t mass of data was made by Wiens (1962), b u t i t was not e n t i r e l y successful i n represent ing t h e world at01 1 s a s an ecosystem. Other qua1 i t a t i v e desc r ip t ions of the i s l and ecosystem a r e included i n Numata (1967) e s p e c i a l l y those papers by Jackson, and Sachet . See a l s o S todda r t , 1 9 6 9 ~ . An idea of the enormous amount of basic information a v a i l a b l e on t h e t e r r e s t r i a l a spec t s of coral a t o l l s may be gained by perusing the 261 pages devoted t o t h i s sub jec t i n Is land Bibl iographies and 236 i n i t s supplement (Sachet and Fosberg 1955, 1971 ) . From t h i s wealth of d a t a , and c i t i n g only a few s p e c i f i c r e fe rences , we can de r ive a f a i r idea of components and processes involved i n t h e t e r r e s t r i a l o r cay ecosystem and inf luences on the adjacent submerged reef ecosystems. Some components and processes, such a s n u t r i e n t content of ground water , s a l t spray, r a i n water , animal and p lan t bodies, and sediments, have never been quan t i f i ed b u t can be measured by standard ana ly t i ca l procedures. Standing crops of the macroscopic organisms present no problem, b u t again, t h e r e a r e no q u a n t i t a t i v e da ta f o r coral i s l a n d s . Estimates can be reasonably made by simple sampling, counting, and weighing procedures. Imports of organic mat te r ( f i s h , squid, e t c . ) by seab i rds and shore and wading b i rds i n t o t h e t e r r e s t r i a l at01 1 ecosystem have never been es t imated . Some observat ions have been made on t h e ground water of a t o l l s , beginning with Charles Darwin ( l839) , but mostly within t h e l a s t few years (Fosberg, 1959; Cox, 1951 ; Arnow, 1954, 1955; Tracey u.? 1961) . The general nature of t h e a t o l l ground water l e n s has been e s t ab l i shed , b u t d e t a i l e d s t u d i e s of i t s behavior under varying condi t ions of geological s t r u c t u r e and r a i n f a l l amounts and regimes, a s well a s of t i d a l ranges and regimes, a r e s t i l l requi red . The t e r r e s t r i a l aeoloav of c e r t a i n a t o l l s has been s tudied by a number of expedi t ions4and 6 d i v i d u a l s (e .g . Royal Socie ty , 1904; ~ e n t w o r t h , 1931 ; Fosberg e t a1 ., 1956, Newell, 1956; Fosberg, 1957a; McKee, 1958, 1959; Fosberq a n d a r r o l l , 1965; Sachet 1962b; S todda r t , 1962, l96gb). Very few q u a i t i t a t i v e measurements of sediments a r e avai1,abl e . F a i r l y d e t a i l e d s o i l s t u d i e s a r e a v a i l a b l e f o r a few P a c i f i c a t o l l s (Stone, 1951 ; Fosberg, 1954; Fosberg e t a1 . , 1956; Fosberg and C a r r o l l , 1965; Te rc in i e r 1956, 1969) a s well a s a general d iscuss ion of a t o l l s o i l s (Stone, 1953). Most of the a t o l l so i l types a r e very widespread and t h e pa t t e rns r a t h e r s i ~ n p l e . There has been no systematic inves t iga t ion to determine i f Caribbean a t o l l s o i l types correspond t o those i n t h e Pac i f i c . Pac i have 1963 The occurrence and o r i g i n of a t o l l phosphate rock i n t h e cen t r a l f i c a t o l l s were e luc ida ted by Fosberg(l957b); and f u r t h e r observat ions been reported by Stone (I%?,), Fosberg a. (1956), Niering (1961, ) , and Roy (l97Ob). S imi lar rock was described f o r t h e Indian Ocean a t o l l s by Piggot t (1968) and r ecen t ly was found on Glover 's Reef, B r i t i s h Honduras by Fosberg. Hutchinson (1950) d e a l s i n g r e a t d e t a i l with background information on phosphate accumulation. Subs tant ia l information i s a v a i l a b l e on a t o l l f l o r a s and vegetat ion (Fosberg 1949; Hatheway, 1953, 1955; Fosberg, 1953; Fosberg u.1956; Fosberg, 1957; Sachet 1962a; S todda r t , 1962; e t c . ) Pure s tands of c e r t a i n t r e e s and shrubs a r e f r equen t ly found, an unusual occurrence i n the t r o p i c s . The land animals of coral i s l a n d s a re reasonably well known, b u t information, except f o r t h a t on b i r d s , i s mostly s c a t t e r e d i n monographs and papers on the groups concerned. A few papers s p e c i f i c a l l y on a t o l l faunas a r e found i n , t h e Atoll Research Bul le t in s e r i e s and i n the r e p o r t s of the B r i t ~ s h Indian Ocean expedit ions (Percy Sladen, e t c . ) of t h e e a r l y twentieth century. There i s an extensive l i t e r a t u r e on a t o l l b i rds ( l i s t e d and annotated by Sachet and Fosberg 1955, 1971). The a t o l l b i rd da ta have never been reviewed a s a whole, but t h e Smithsonian f i e l d guides (Watson u.1963; King, 1967) and a number of regional papers (Amerson, 1959; Baker, 1951 ; Pelzl , ms.) a r e s t e p s i n t h i s d i r e c t i o n . Atoll i n s e c t s , a s well a s t h e i n s e c t s of high i s l a n d s , a r e discussed i n t h e Pac i f i c Insec t s and I n s e c t s of Micronesia s e r i e s ( G r e s s i t t , e d . ) . Papers t h a t deal with the i n t e r a c t i o n s of a t o l l faunas a r e a v a i l a b l e f o r Arno a t o l l (Marshall , 1951) and t h e Tokelaus (Hinckley, 1969) . Data on t h e consequences t o t h e r e s t of t h e a t o l l ecosystem of the presence of l a r g e numbers of seabi rds a r e brought together by Hutchinson (1950). No work has been done on e i t h e r t e r r e s t r i a l primary p roduc t iv i ty o r ni t rogen f i x a t i o n on a t o l l s , nor i s t he re much information on t e r r e s t r i a l food chains , predat ion o r decomposition of organic matter under a t o l l condi t ions . Extrapolat ions mav be made from o the r ecosvstems, but i t would be important' to know jus't how well these extrapol~tions~correspond t o ac tua l a t o l l pa t t e rns . HUMAN INFLUENCES The complex of human inf luences on and i n t e r a c t i o n s with t h e cay ecosystem and the t o t a l r ee f ecosystem i s q u i t e apparent but d i f f i c u l t t o c h a r a c t e r i z e and summarize s a t i s f a c t o r i l y (Johannes 1970-71). The p e s t i c i d e o r , b e t t e r , b ioc ide , component i s of general importance and i s very probably pervasive in a l l p a r t s of the system. This p a r t i c u l a r problem i s d e a l t with s p e c i f i c a l l y below. The na ture of changes brought about by human a c t i v . i t y v a r i e s f rom obv ious ( S t o d d a r t 1968a) and e a s i l y measured t o v e r y obscure and h a r d t o e s t i m a t e , b u t t h e impor tance o f these changes t o t h e system may n o t n e c e s s a r i l y be p r o p o r t i o n a l t o t h e i r obv iousness. P o l l u t i o n i n terms o f a d d i t i o n o f human wastes (sewage, e t c . ) can be e s t i m a t e d f rom s tandard f i g u r e s and t a b l e s . The amounts and e f f e c t s o f s o l i d - w a s t e p o l l u t i o n a r e p r o b a b l y i n p r o p o r t i o n t o amount o f human a c t i v i t y on t h e cays and w i t h i n t h e lagoon, w i t h a s u b s t a n t i a l a d d i t i o n f rom d r i f t m a t e r i a l s washed i n f rom t h e open ocean. An FA0 con fe rence on Mar ine P o l l u t i o n (Ruivo, ed., 1972) i n c l u d e d severa l accounts o f p o l l u t i o n o f r e e f s , a t o l l s and lagoon (see e s p e c i a l l y Johannes, Chan, B a g n i s ) . I nc reased s e d i m e n t a t i o n i s ano the r f r e q u e n t r e s u l t o f man's a c t i v i t i e s , as e x e m p l i f i e d by Kaneohe Bay, Oahu (Roy, l 97Oa) . T h i s l a t t e r problem i s l i k e l y t o be more severe on h i g h i s l a n d s than on a t o l l s . One o f t h e most s i g n i f i c a n t e f f e c t s o f human a c t i v i t y i s t h e d e s t r u c t i o n and/or a l t e r a t i o n o f h a b i t a t s o f o t h e r organisms (Sachet , 1963; Jackson, 1967). The rep lacement o f n a t u r a l f o r e s t by coconut p l a n t a t i o n i s a good example ( c f . S t o d d a r t , 1968). Except f o r s i m p l e measurement o f a rea , we know o f no way t o measure such changes o r t h e i r e f f e c t s , o r , i n many cases, even t o a s s i g n s p e c i f i c e f f e c t s t o p a r t i c u l a r causes. T h i s does n o t i n any way m i n i m i z e t h e i r impor tance. A volume e d i t e d by Fosberg (1963a), d e a l s w i t h i s l a n d s i n c l u d i n g a t o l l s , w i t h s p e c i a l r e f e r e n c e t o man's r o l e i n t h e i r eco logy . The presence o f a mar ine l a b o r a t o r y i n Kaneohe Bay, Oahu, Hawa i i , has l e d t o a l a r g e number o f s t u d i e s concern ing v a r i o u s aspects o f human e f f e c t s on t h e b a r r i e r and f r i n g i n g r e e f complex t h e r e . Among t h e papers d e a l i n g w i t h t h a t bay a r e Bathen (1968) ; Roy (1970a); Smith e t a l . (1 970, 1973) ; Smi th ( 1 971 b) ; Caperon s.(1 971 ) ; C l u t t e r ( 1 972) ;and Johannes (1970,1972). These papers have d e a l t p r i m a r i l y w i t h t h e h i g h n u t r i e n t l e v e l and r a p i d d e p o s i t i o n r a t e i n t h e bay. P o l l u t i o n o f t h e b iosphere by man-made chemica l compounds has reached a l e v e l such t h a t a l l f auna l e lements o f t h e e a r t h a r e contaminated (R isebrough s., 1970) . These compounds a r e o f two t ypes : 1 ) b i o c i d e s ( p r i m a r i l y c h l o r i n a t e d hydrocarbons) manufactured and d i s t r i b u t e d s p e c i f i c a l l y t o e r a d i c a t e o r c o n t r o l " p e s t " organisms; and 2) chemica ls manufactured f o r and used i n i n d u s t r i a l p rocesses wh ich "escape" from t h e i r i n t e n d e d area o f use ( p r i m a r i l y p o l y c h l o r i n a t e d b i p h e n y l s and heavy m e t a l s ) . The accumulated 1 i t e r a t u r e on t h e d i s t r i b u t i o n o f these com- pounds i s e x t e n s i v e b u t o n l y r e c e n t l y have t h e i r p h y s i o l o g i c a l e f f e c t s on m e t a b o l i c processes begun t o be e l u c i d a t e d . New i n f o r m a t i o n on i n d u c t i o n o r i n h i b i t i o n o f v a r i o u s enzyme systems and chemical r e a c t i o n s by man-made chemica ls becomes a v a i l a b l e w i t h each e d i t i o n of t h e p e r t i n e n t j o u r n a l s . These p h y s i o l o g i c a l e f f e c t s a r e e s p e c i a l l y r e l e v a n t t o mar ine organisms, s i n c e accumu la t ion and b i o l o g i c a l con- c e n t r a t i o n o c c u r most r e a d i l y i n a q u a t i c systems. Wh i le b i o c i d e s have been d e t e c t e d by a l l s t u d i e s t h u s f a r des igned t o i n v e s t i g a t e t h e i r presence o r absence, few such s t u d i e s have been c a r r i e d o u t on c o r a l a t o l l s ; and l i t t l e i n v e s t i g a t i o n has been made o f biocide accumulation in any level of the food web of a coral a t o l l . To our knowledge, the only non-marine data available fo r a tropical island are in unpublished information on Sooty Tern eggs from the Dry Tortugas, Florida, in which DDE and PCBs were present i n a l l samples (W.B . Robertson, J r . pers. comm.). By an accident of geography, a t o l l s have been the most numerous s i t e s of atomic and nuclear atmospheric bomb-tests and, inevi tably , the s i t e s of pollution by radionuclides. Between 1946 and 1958, more than 59 t e s t s took place a t Eniwetok and Bikini a t o l l s (Welander, 1969). Later Christmas and Johnston Islands were involved, and more recently, Mururoa and Fangataufa Atolls in the Tuamotus. In the Marshall Islands, a survey of the ecosystem had been carried out prior to any t e s t s (Ladd, 1973), and periodic resurveys followed. They gave r i s e to an enormous l i t e r a t u r e much of i t in c lass i f ied or hard-to-get AEC repor ts . Only a f t e r many years did papers appear in s c i en t i f i c journals of more general d i s t r ibu t ion , as well as i n proceedings of conferences and symposia (see fo r instance papers by Beasley, Beasley and others , Held, Held and others , Nelson and Evans, Templeton Ual, and many others) . Such information has not ye t been integrated as a whole picture of radionuclide pollution on coral reefs and a t o l l s , b u t much data continues t o be accumulated. There i s much l e s s eas i ly available information on Johnston Island, Christmas Island (Palumbo e t a1 ., 1966) o r the Tuamotus. In the Marshalls, as the r e s u l t of a catastrophic incident (Operation Castle, 1954) the e f f ec t s of exposure to radiation on a to l l human population became avai lable fo r study. Detailed medical surveys of the exposed islanders have been carried out and repeated a t regular in tervals . See f o r example papers by Conard e t a1 ., Robbins e t a l . and Lisco and Conard. The sociological e f f ec t s of the Castle d i sas te r , as well as those of the displacement of the Eniwetok and Bikini populations a re other aspects of human interference with the coral island ecosystem (Stoddart 1968a). Many others , the impact of war on a t o l l s , of organized migration, of changes in economic patterns, could be mentioned, b u t cannot be detailed here. This type of information i s beyond the scope of t h i s review, b u t i t i s obvious tha t anthropology and history can furnish evidence relevant to the tota l picture of coral island ecosystems. MARINE BIOTA: BENTHIC PLANTS Benthic marine algae make major contributions t o primary pro- du'ctivi t y , nitrogen f ixa t ion , community s t ructure , organism d i s t r i - bution, carbonate production, and reef consolidation and destruction. Setchell (1926, 1928) was probably the f i r s t biologist t o recognize f u l l y the variety and importance of these roles of algae in coral reefs , and much work has followed his pioneer e f fo r t . Setchell (1928) described zonation of algae across the reef , a s have Kanda (1944), Doty and Morrison (1954), Gilmartin (1960), Doty (1967, 1970) and Tsuda (1 970). Seasonal occurrence has been described by Bernatowicz (1952) and Denizot (1969), and measured on Guam by Tsuda (1972), and Dahl (1972) has analyzed community s t r u c t u r e o f Samoan a lgae . Dahl (1971) has a l s o demonstrated t h a t c e r t a i n benthic a lgae a r e useful as ecological i nd ica to r s and can even provide a continuous record o f environmental condi t ions . S tudies on temperate and subtropical a lgae have demonstrated t h e i r importance in ecological i nves t iga t ions and have developed techniques f o r surveying f i e l d populations. Line t r a n s e c t s can provide a useful bas is f o r descr ib ing community s t r u c t u r e and measuring seasonal v a r i a t i o n s , a s i n the work by Neushul (1967) on subt ida l vegeta t ion i n western Washington, and t h e r ecen t survey of subt ida l ecology o f f southern Cal i forn ia (Neushul a., 1967; Clarke and Neushul, 1967). Experimental techniques can add considerably t o the f i e l d da ta on community composition and ecology (Neushul and Dahl , 1967). Crustose c o r a l l i n e a lgae (Melobesioidae) may be a t l e a s t a s important a s c o r a l s i n t h e development of r ee f s t r u c t u r e s (Setchel l , 1926; J.H. Johnson, 1961 ; Gross u., 1969). These a lgae have been e s p e c i a l l y noted in the P a c i f i c , where they have given t h e i r name t o thelLithothamnion r i d g e , " a s t r i k i n g b u t misnamed topographic f e a t u r e of P a c i f i c a t o l l s ; however, the importance of c o r a l l i n e a lgae t o r ee f cons t ruc t ion i n general i s probably g r e a t e r than t h e ex tan t reef l i t e r a t u r e would ind ica t e (Denizot , 1972). In many a reas of t h e P a c i f i c ( e .g . Ladd u.,1970; J.H. Johnson 1961) a s well a s i n the Tethys Seaway (e .g . Lemoine, 1939; J.H. Johnson, 1965) and even i n the A t l a n t i c (Iams, 1969) encrus t ing c o r a l l i n e a lgae have been important o r even the primary con t r ibu to r s t o sedimentary formations. Considering the widespread occurrence of encrust ing coral 1 ine a lgae , t h e i r importance i n determining t h e na ture of the substratum-water i n t e r f a c e , and t h e i r presence i n the Fossil record, they have been r a t h e r neglected. They a r e considered t o be a d i f f i c u l t group ( s e e f o r example W . R . Taylor 1960; Adey 1970) and have o f t en been t r e a t e d a s p a r t of the "dead" s u b s t r a t e . Adey has underway a d e t a i l e d s tudy of the r o l e of c o r a l l i n e a lgae i n t h e r ee f ecosystem. Calcareous green a lgae (most1 y Hal imeda spp.) con t r ibu te g r e a t l y t o loose sediments i n c e r t a i n a reas of the r e e f , e s p e c i a l l y c e r t a i n zones of lagoons (Chapman, 1901; Emery e t a1 ., 1954; Hoskin, 1963). Stockman e t a l . (1967) and Neumann and L-1969) have a l s o shown t h e importance of the genus P e n i c i l l u s i n the formation of lime muds i n Florida and the Bahamas. Land (1971) has s tudied the importance of sediment production by Melobesia, a red a lga which commonly enc rus t s sea g ras ses . The r o l e s of boring a l g a e and microbenthic a lgae have y e t t o be de l i7ea ted in r e spec t t o calcium carbonate breakdown though Nesteroff (1956) and o t h e r s a s c r i b e t o them a major r e s p o n s i b i l i t y f o r i n t e r t i d a l e ros ion . Some at tempts a t including c rus tose c o r a l l i n e a lgae i n regional s tud ie s of calcium carbonate budgets (S.V. Smith c., 1970, l972a) and in inves t iga t ions of primary p roduc t iv i ty (Marsh, 1970; L i t t l e r , 1971 have been undertaken. The r e s u l t s a r e prel iminary, however, a s there i s l i k e l y a va r i a t ion by several orders of magnitude i n growth and metabolic r a t e s , a s a funct ion of l i g h t , temperature, s p e c i e s , and even p a r t of an individual p l an t considered (Adey and McKibbin, 1970). Some prel iminary product iv i ty measurements have a l s o been made on reef a lgae by Doty (1971) and Soegiar to (1972). Recent s t u d i e s have indeed shown t h a t the alga-covered reef f l a t , a v i s u a l l y unimpressive and o f t e n ignored component of t h e r ee f community (Dahl , 1972), was in f a c t , twice a s productive a s a reas of r i c h coral cover and supported l a rge roving populations of herbivorous f i s h e s (Johannes e., 1972). Symbiosis, o r the occurrence of zooxanthel lae within l i v i n g coral t i s s u e s has long been known, b u t only r ecen t ly has t h e i r r o l e as n u t r i e n t acceptors been demonstrated (Goreau and Goreau, 1960; Muscatine and Hand, 1958). The zooxanthel lae a l s o play a s i g n i f i c a n t r o l e i n t h e c a l c i f i c a t i o n of some c o r a l s (Goreau, 196la , Goreau and Goreau, 1960). There i s s t i l l cons iderable debate over the i n t e r - ac t ion between p lan t and animal components. Recent work has only begun to explore the v a r i e t y of marine symbiotic r e l a t i o n s h i p s (see f o r example, D . L . Taylor 1969a, 1969b, 1971). Information on grazing of a lgae by reef herbivores includes general observat ions on predat ion upon a lgae via s t u d i e s of g u t contents and r e s u l t s from caging; herbivore d e n s i t i e s have not been reported beyond presence o r absence, nor have the p lant communities in which t h e experiments were c a r r i e d o u t been r igorous ly defined (Randall , 1961, 1965; Mathiesen u., 1972; Earle 1972). The s l a t e -penc i l urchin, Heterocentrotus mammil l a t u s , has been observed t o feed heavily on Porolithon sp . on the a lga l r idge of Bikar Atoll i n t h e Marshall I s l ands , and t o have a s i g n i f i c a n t inf luence in the e ros ion of the reef edge (Fosberg, pers . comm.). &a grasses" a r e l o c a l l y important i n t rop ica l nearshore environments. In h i s monograph, den Hartog (1970) summarizes under each spec ie s what i s known o f i t s biology, i t s r o l e a s food f o r herbivores ( e .g . Randall , 1965), and i t s funct ion a s a substratum s t a b i l i z e r . Recently, the echinoid Diadema has been shown t o graze heavi ly on Thalassia in c e r t a i n s i t u a t i o n s i n the Caribbean (Ogden g., 197-e have been r a t h e r few q u a n t i t a t i v e s t u d i e s on the product iv i ty of t u r t l e g ra s s (Thalassia testudinum), one of t h e more abundant and the bes t known spec ie s . Early s t u d i e s were made by Pomeroy (1960), Odum (1957), and Jones (1968) on product iv i ty measured by t h e 0 method. However, c l o s e r inspec t ion by Zieman (1968) has shown ghat t h i s method i s suspec t f o r Thalassia produc- t i v i t y , a s t h e leaves have the capac i ty t o expand and s t o r e gasses in i n t e r s t i t i a l lacunae. This phenomenon has a l s o been demonstrated by Hartman and Brown (1967) f o r t h e fresh-water spec ies Elodea canadensis and Ceratophyllum dimersum. INVERTEBRATES It i s i m p o s s i b l e he re t o d i s c u s s t h e i n d i v i d u a l r o l e s o f t h e numerous groups o f i n v e r t e b r a t e s i n t h e r e e f ecosystem, o r t h e enormous amount o f a v a i l a b l e work b a s i c t o an i n t e g r a t e d s t u d y of t h e system. Many p e r t i n e n t papers on i n v e r t e b r a t e s have appeared i n Cahiers du P a c i f i q u e , wh ich a l s o i n c l u d e b i b l i o g r a p h i e s of c e r t a i n groups, and i n r e c e n t Symposium volumes c i t e d pp. 1-2. T h i s r e v i e w w i l l o n l y t o u c h on a few h i g h l i g h t s o f r e c e n t r e s e a r c h on i n v e r t e b r a t e s i n t h e i r f u n c t i o n s i n t h e system. S ince c o r a l r e e f s a r e a lmos t e n t i r e l y t h e r e s u l t o f b i o l o g i c a l a c t i v i t y , an a n a l y s i s o f r e e f development may be approached by examin ing s t r u c t u r a l a c t i v i t i e s ( c o n s t r u c t i o n , maintenance, and d e s t r u c t i o n ) and t h e c o n t r i b u t i o n o f t h e v a r i o u s b i o t a i n v o l v e d ( c a l c i f i c a t i o n by hermatypes, b i n d i n g and b a f f l i n g p r o p e r t i e s o f some s o f t - b o d i e d spec ies , and b i o d e g r a d a t i o n by a v a r i e t y o f i n v e r t e b r a t e s ) . Work on growth r a t e s o r c a l c i u m ca rbona te d e p o s i t i o n r a t e s has been ment ioned i n p r e v i o u s s e c t i o n s . The min imal energy requ i rements o f c o r a l s were i n v e s t i g a t e d by Coles (1969) , who conc luded t h a t t h r e e common r e e f spec ies a r e capab le o f c a p t u r i n g and i n g e s t i n g s u f f i c i e n t zoop lank ton t o account f o r d a i l y maintenance under l a b o r a t o r y c o n d i t i o n s . Y e t i t was de te rm ined f rom a f i e l d s t u d y i n Bermuda t h a t t h e energy needs o f c o r a l s were o f an o r d e r of magni tude g r e a t e r t h a n c o u l d be met by t h e s u p p l y o f d r i f t i n g n e t zoop lank ton (Johannes e t a1 . , 1970) . Obv ious ly these m e t a b o l i c s t u d i e s a r e n o t c o n c l u s i v e F u r t h e r s t u d i e s o f c o r a l me tabo l i sm shou ld i n c l u d e examina t ion o f c o r a l f e e d i n g and d i g e s t i v e mechanisms and t h e r e l a t i o n o f these mechanisms t o c o r a l morphology and food t ype . Hartman and Goreau (1970) have c a l l e d a t t e n t i o n t o t h e s c l e r o - sponges as an i m p o r t a n t c o n s t r u c t i o n a l e lement bes ides s c l e r a c t i n i a n f r a m e b u i l d e r s i n t h e sub- ree f and deeper r e e f f ramework. I t i s a l s o c l e a r t h a t many b e n t h i c r e e f i n v e r t e b r a t e s p l a y an i m p o r t a n t r o l e i n maintenance processes. An example i s t h a t o f t h e sponge Mycale l a e v i s , wh ich p r o t e c t s t h e l o w e r su r faces o f some massive c o r a l s f r o m t h e d e s t r u c t i v e e f f e c t s o f b o r i n g sponges (Goreau and Hartman, 1966) . A l though c o r a l s a r e commonly t h e p r i n c i p a l animal c o n t r i b u t o r s o f s k e l e t a l m a t e r i a l s t o t h e f ramework and l o o s e sediments t h a t make up t h e r e e f , severa l o t h e r i n v e r t e b r a t e groups can make s u b s t a n t i a l a d d i t i o n s ( e s p e c i a l l y mo l lusks , echinoderms, and F o r a m i n i f e r a ) . The l a t t e r , i n a d d i t i o n t o t h e i r commonly recogn ized r o l e as s t r a t i g r a p h i c markers, a r e f r e q u e n t l y a prominent component of l o o s e sediments. Some e n t i r e a t o l l beaches i n t h e c e n t r a l P a c i f i c a r e made up o f t h e worn t e s t s o f C a l c a r i n a and Bacu logyps ina, two genera t h a t i n h a b i t windward r e e f f l a t s . Hometrema forms consp icuous d a r k r e d c r u s t s on t h e c o r a l f ragments seen on G l o v e r ' s Reef and o t h e r a t o l l s . S k e l e t a l framework d e s t r u c t i o n by b i o l o g i c a l means has been d iscussed above. Some processes a r e d i r e c t and obv ious, such as t h e f e e d i n g o f f i s h e s , c rabs, worms, s n a i l s , e t c . on l i v i n g c o r a l s . No tab le a1 so i s t h e r e c e n t p o p u l a t i o n e x p l o s i o n o f Acanthaster p l a n c i which feeds upon and l o c a l l y d e v a s t a t e s r e e f s i n P a c i f i c a reas (An ton ius 1 W l a , Chesher, 1969; Newman 1970; Randal l 1972; A t o l l Res. B u l l . 166-170, 1973; M ic rones ica , 9(2) , Dec. 1973; and numerous o t h e r r e c e n t papers ) . f e e d i n g o f f i s h e s on i n v e r t e b r a t e s , e.g. c o r a l s and sponges, was s t u d i e d by Randal l (1967), Randal l and Hartman (1968), and Bakus (1967) . Many e n d o l i t h i c b o r i n g and bur row ing forms a r e a l s o known (e .g . sponges: Goreau and Hartman 1963, and R u e t z l e r , 1971 ; s i p u n c u l i d s : Rice, 1969; mo l lusks : R . Robertson, 1970, and r a t e s o f d e s t r u c t i o n have been c a l c u l a t e d f o r some of them (Neumann, 1966; Glynn u. 1971) . I n t e r a c t i o n s i n v o l v i n g i n v e r t e b r a t e s have been summarized f o r Western A t l a n t i c r e e f s by Glynn ( i n p r e s s ) . Lang (1970, 1971) o b t a i n e d i n f o r m a t i o n on t h e i n t e r s p e c i f i c aggress ive behav io r o f hermatyp ic c o r a l s , r e p r e s e n t i n g a feedback l o o p w i t h i n a s i n g l e f u n c t i o n a l component. Research on t h e r o l e o f smal l i n t e r s t i t i a l an imals i n t h e c o r a l r e e f ecosystem has lagged behind o t h e r areas o f s tudy . P r e l i m i n a r y sampl ing has i n d i c a t e d , f o r example, t h a t groups such as T u r b e l l a r i a , Nematoda, Po lychaeta, and Crustacea a r e abundant i n r e e f sediments, r a i s i n g t h e p o s s i b i l i t y t h a t t h e y may p r o v i d e a s i g n i f i c a n t food resource f o r d e p o s i t feeders znd p o s s i b l y g r a z e r s and browsers (Renaud-Mornant, e t a1 ., 1971 ; Thomassin 1972; papers presented a t t h e " F l o a t i n g ~ y m p o s i u m " s e e p.2) by Thornassin, J.F. Grass le and o t h e r p a r t i c i p a n t s ) . FISH (AND OTHER VERTEBRATES) The p l a c e o f f i s h e s i n t h e budget o f t h e r e e f ecosystem seems t o have been e l u c i d a t e d perhaps i n more d e t a i l t han t h a t o f most o t h e r v e r t e b r a t e groups. Except f o r t h e green t u r t l e , t h e b i o l o g y of which i s be ing s t u d i e d i n v a r i o u s p a r t s o f t h e t r o p i c s (e.g. F r a z i e r ms.; E h r e n f e l d 1974), l i t t l e c o n s i d e r a t i o n has been g i v e n t o such o t h e r v e r t e b r a t e s as s e a l s o r sea-snakes, which a r e o n l y l o c a l l y s i g n i f i c a n t . Sea b i r d s have a l r e a d y been ment ioned w i t h r e s p e c t t o t h e t e r r e s t r i a l p a r t o f t h e ecosystem. Est imates o f ' f i s h biomass have been made by Odum and Odum, 1955 (425 k g l h e c t . ) ; Bardach, 1959 (450 k g l h e c t . ) ; Randal l , 1963b (1590 k g l h e c t . ) Brock, 1954 (1850 k g l h e c t . ) ; and Goldman and T a l b o t , i n p ress (200- 2100 k g l h e c t . f r o m d i f f e r e n t r e e f a r e a s ) . These w i d e l y d i f f e r e n t e s t i m a t e s may be due i n p a r t t o d i f f e r e n t sampl ing techn iques and i n p a r t t o t h e v a r i a t i o n t h a t i s now known t o occur between d i f f e r e n t p a r t s o f a s i n g l e r e e f . As y e t , t h e r e i s no good i n f o r m a t i o n on d i f f e r e n c e s between w i d e l y separated r e e f s , p a r t i c u l a r 1 y f r o m ocean t o ocean. Some o f t h e f a c t o r s i n f i s h b i o l o g y which a f f e c t biomass may be d e t a i l e d as f o l l o w s : H a b i t a t s e l e c t i v i t y i s marked i n r e e f f i s h e s . D i s t r i b u t i o n i s o b v i o u s l y a f f e c t e d by depth, s h e l t e r , food a v a i l a b i l i t y and c o m p e t i t i v e i n t e r a c t i o n s . Var ious areas o f t h e r e e f have d i f f e r e n t s p e c i f i c i t y i n many zones. Up t o 25 p e r c e n t o f spec ies may be r e s t r i c t e d t o s i n g l e areas o f t h e r e e f (Ta l b o t , unpub l i shed i n f o r m a t i o n ) . From t h e work done on t h e movements o f f i s h e s on c o r a l r e e f s ( S p r i n g e r and McErlean, 1962; Randal 1 , 1961 , 1962, 1963a; Bardach, 1958; C.L. Smi th and T y l e r , 1972; Moe, 1969; Winn and Bardach, 1960; Reese, 1964) i t seems t h a t t h e m a j o r i t y o f r e e f f i s h e s a r e r e s t r i c t e d t o a r e e f o r even i n d i v i d u a l p a t c h r e e f s , a t l e a s t f o r l o n g p e r i o d s . Many spec ies a r e t e r r i t o r i a l . Some few p e l a g i c spec ies such as some scombrids, carangids, and sharks, a p p a r e n t l y move between r e e f s ( T e s t e r and Wass, personal communicat ion). These spec ies feed m o s t l y on r e e f f i s h e s . W i t h i n t h e i r t e r r i t o r i e s , many f i s h e s have d i f f e r e n t f e e d i n g and r e s t i n g areas and may move up t o a m i l e o r even more on d a i l y f e e d i n g m i g r a t i o n s (Winn and Bardach, 1960; Randal l and Randa l l , 1963; Randa l l , 1963; Bardach and Menzel , 1957) . Some f i s h e s may have dai ly b reed ing m i g r a t i o n s . Others may r e s t i n caves down t h e deep r e e f f r o n t and f e e d on t h e r e e f f l a t o r r e s t on p a t c h r e e f s by day and f e e d o v e r Tha lass ia beds a t n i g h t . Some d i e 1 changes i n d i s t r i b u t i o n have been documented by Hobson (1965, 1968, 1972), S ta rck and Dav is ( 1 966), and C o l l e t t e and Tal b o t (1 972) . The food h a b i t s o f West I n d i a n and P a c i f i c c o r a l f i s h e s have been s t u d i e d by a number of workers, and t h i s i n f o r m a t i o n forms a s o l i d q u a l i t a t i v e b a s i s f o r f u r t h e r s t u d y ( H i a t t and S t rasburg , 1960, Suyehi ro , 1962, Randal 1, 1967; P l e s s i s 1972; Star&, Emery, pe rsona l communicat ions). Few da ta on f e e d i n g e f f i c i e n c y and convers ion a r e a v a i l a b l e f o r c o r a l r e e f f i s h e s , b u t a f a i r amount i s known abou t m e t a b o l i c r a t e s i n temperate f r e s h w a t e r and mar ine f i s h e s . It appears t h a t convers ion f a c t o r s c o u l d be used t o c a l c u l a t e t h e metabol ism o f c o r a l r e e f f i s h e s . Some f u r t h e r work i s needed, however, on s e l e c t e d spec ies o f d i f f e r e n t s i z e s , a c t i v i t y p a t t e r n s , and f e e d i n g h a b i t s f o r accura te ecosystem model ing. Most i n v e s t i g a t o r s who have worked on t h e problem o f age and g rowth r a t e s have found t h a t c o n v e n t i o n a l techn iques a r e n o t a lways a p p l i c a b l e t o t r o p i c a l f i s h e s . Recent work (Moe, 1969) has shown, however, t h a t o t o l i t h s can be used f o r some spec ies, and s i ze - f requency techn iques as w e l l as ag ing p o p u l a t i o n dynamics may y i e l d d a t a f o r r e p r e s e n t a t i v e species (Gul l and , 1970). R e l a t i v e l y 1 i t t l e i s known o f t h e r e p r o d u c t i v e b i o l o g y o f c o r a l r e e f f i s h e s (Breder and Rosen, 1966). F ishes e x h i b i t a wide v a r i e t y o f r e p r o d u c t i v e mechanisms, f r o m extreme p a r e n t a l c a r e (e.g. mouth b reed ing) t o random s c a t t e r i n g o f eggs. A few spec ies have.been shown t o undergo d a i l y o r annual b reed ing m i g r a t i o n s (Randal l and Randa l l , 1963; Reinboth, 1973; C.L. Smi th and T y l e r 1972). Sex r a t i o s a r e g r e a t l y v a r i a b l e , and severa l t y p e s o f hermaphrod i t ism a r e common Fecundi ty , r e c r u i t m e n t , and egg and l a r v a e l o s s e s a r e perhaps most poor1 y known. Reef f i s h e s can a l s o p l a y a r o l e i n r e e f d e s t r u c t i o n . Bardach (1961) has documented t h e r o l e o f d e s t r u c t i o n by f i s h e s on Bermudian ree fs , w h i l e Glynn e t a l . (1971) have done l i k e w i s e f o r f i s h e s on t h e P a c i f i c r e e f s o f Pan= Graz ing by p a r r o t - f i s h e s and some o t h e r r e e f f i s h e s i s commonly observed on r e e f s i n a l l a reas. PLANKTON A t p resen t , t h e q u a n t i t a t i v e c o n t r i b u t i o n o f phyto- and zooplankton t o t h e r e e f ecosystem i s unknown. E s t i m a t i o n s o f t h e s t a n d i n g c r o p o f zooplankton around c o r a l a t o l l s and i n t h e lagoon areas can be found i n M.W. Johnson (1949, 1954), G i l m a r t i n (1958) , Mahnken (1966) , B a r n e t t (ms.), O.A. Math isen (1964), Odum and Odum (1955), and A.R. Emery (1968) . Most i n v e s t i g a t o r s have found t h a t t h e biomass o f zooplankton i n t h e v i c i n i t i e s o f r e e f s and a t o l l s and i n s i d e c o r a l r e e f lagoons i s h i g h e r than t h e biomass i n t h e open ocean a t t h e same l a t i t u d e s , y e t t h e r e e f zooplankton biomass has p r o b a b l y been underes t ima ted due t o t h e inade- quate sampl ing o f e p i b e n t h i c ( h o v e r i n g above t h e water -bot tom i n t e r f a c e o r w i t h i n t h e c o r a l heads) and n e u s t o n i c ( n e a r s u r f a c e ) forms. P resen t e s t i m a t e s o f zooplankton biomass, however, a r e o r d e r s o f magni tude (Odum and Odum, 1955) s m a l l e r than e s t i m a t e s o f b e n t h i c organism and f i s h biomass. The s t a n d i n g s t o c k o f p h y t o p l a n k t o n has been expressed by c h l o r o p h y l l - A va lues (N. M a r s h a l l ) , b u t convers ion o f these va lues i n t o carbon i s u n r e a l i s t i c w i t h o u t f u r t h e r a n a l y s i s . The l i t e r a t u r e does c o n t a i n i n f o r m a t i o n on such necessary i n p u t s t o t r o p i c a l p h y t o p l a n k t o n as l i g h t , n u t r i e n t s , and t r a c e meta ls ( S t r i c k l a n d , 1960; J e f f r e y , 1968) . One common component o f t h e c o r a l r e e f wa te r column, Trichodesmium, has been s t u d i e d e x t e n s i v e l y (Prabhu e t a1 . , 1966; C a l e f and Gr ice, 1966; Goer ing e t a1 ., 1966; Ramamurthy a n z h a d r i , 1966a, 1 g66b; Ramamurthy and Kr ishnamurthy, 1967) . I t i s obv ious t h a t many r e e f organisms f e e d on zooplankton. Q u a n t i t y o f p l a n k t o n u t i l i z e d by one r e e f assemblage i n Puer to R ico was determined by Glynn (ms). Glynn ( i n press and ms.) has a l s o g i v e n q u a n t i t a t i v e d a t a on f e e d i n g a c t i v i t y o f f i s h e s and i n v e r t e b r a t e s on zooplankton as w e l l as data on zooplankton r e c r u i t m e n t (Glynn, i n p r e s s ) . NUTRIENTS AND DETRITUS S t u d i e s o f t o t a l community metabo l i sm o n a t o l l r e e f f l a t s s i t u a t e d i n l a r g e l y u n i d i r e c t i o n a l c u r r e n t s were i n i t i a l l y developed by Sargeant and A u s t i n (1949, 1954) . T h e i r method has been a p p l i e d i n a number o f l a t e r s t u d i e s (e.g. Odum and Odum, 1955; Kohn and H e l f r i c h , 1957; Gordon and K e l l y , 1962; M i l l iman and Mahnken, 1961 ; Qasim and Sankaranarayanan, 1970; Odum m., 1959) . The method was f i r s t used i n a comprehensive manner a t Eniwetok i n 1971 d u r i n g P r o j e c t Symbios (Johannes a., 1972). Tha t p r o j e c t proved w i t h o u t doub t t h e e f f i c a c y o f t h i s approach; f o r i n s t a n c e , t h e symbios s t u d i e s e x p l a i n e d why r e e f communit ies a r e so p r o d u c t i v e b i o l o g i c a l l y though bathed i n waters v e r y l o w i n p l a n t n u t r i e n t s N i t r o g e n f i x a t i o n and u n u s u a l l y e f f i c i e n t r e c y c l i n g o f phosphorus w i t h i n t h e community a r e a p p a r e n t l y r e s p o n s i b l e . These two phenomena, t h e r e f o r e , shou ld now be focused upon a t t h e subcommunity and spec ies l e v e l s . Random search ing, spec ies by spec ies , would p r o b a b l y n o t have r e v e a l e d t h e g r e a t q u a n t i t a t i v e s i g n i f i c a n c e o f t h e s e processes. Net r e e f pho tosyn thes is and t o t a l n i g h t t i m e r e s p i r a t i o n can be determined by measur ing t h e i n c r e a s e i n t h e oxygen c o n c e n t r a t i o n i n t h e wa te r as i t crosses t h e r e e f d u r i n g t h e day, and i t s decrease a t n i g h t . Another method which can he1 p m o n i t o r community p r o d u c t i v i t y and r e s p i - r a t i o n uses changes i n t h e CO system (S.V. Smith, 1972) . The system was ana lyzed accord ing t o s o d o f t h e c a l c u l a t i o n s o u t l i n e d by Park (1969) . T h i s approach has a number o f advantages ( c f . S.V. Smi th , 1973; Smi th and Marsh, 1973; Kanwisher, 1963) . S t i l l ano the r approach t o community metabo l ism c o n s i s t s o f measure- ments made i n fenced enc losu res w i t h t h e wa te r s u r f a c e open t o t h e atmos- phere; i t has been s u c c e s s f u l l y used t o a dep th o f 1 m on t h e G r e a t B a r r i e r Reef by Kinsey (1972, and unpub l i shed o b s e r v a t i o n s ) . The r e e f i m p o r t s o r g a n i c m a t t e r f rom upstream i n t h e f o r m o f zoo- p l a n k t o n and e x p o r t s i t downstream i n t h e f o r m o f mucus aggrega tes and a l g a l d e t r i t u s (N. M a r s h a l l , 1965; Johannes, 1967, Johannes ., 1972) . T h i s t r a n s f e r i s l i k e l y t o be o f g r e a t s i g n i f i c a n c e i n l agoon e c o l o g y and p o t e n t i a l aquacu l tu re . A c u r i o u s r e s u l t o f r e c e n t s t u d i e s a t Eniwetok ( P i l s o n and B e t z e r , 1973) has been t o demonst ra te t h a t l e v e l s o f d i s s o l v e d phosphorus change remarkab ly l i t t l e ac ross t h e r e e f . The mechanism by wh ich t h i s constancy i s m a i n t a i n e d i s by no means c l e a r as y e t . I t may be t h a t organ isms p a v i n g t h e r e e f c o n s t i t u t e a s i g n i f i c a n t phosphorus s i n k and t h a t exchange between t h i s s i n k and t h e o v e r l y i n g wa te r c o n s t i t u t e s a mechanism f o r b u f f e r i n g phosphorus l e v e l s i n t h e wa te r . Such a mechanism has been demonstrated between sediments and t h e o v e r l y i n g wa te r i n s a l t marshes by Pomeroy .(1 965, 1967, 1971 ) . ECOSYSTEM ANALYSIS I n t e r e s t i n a mathemat ica l r e p r e s e n t a t i o n o f e c o l o g i c a l phenomena developed e a r l y i n c o n n e c t i o n w i t h t h e dynamics o f p o p u l a t i o n s and ep idemics . Names such as Ma1 thus, P e a r l , M e r h u l s t , Lo tka , and V o l t e r r a w i l l a lways be c i t e d i n any h i s t o r y o f mathemat ica l eco logy . A t t h e p r e s e n t t ime, much a c t i v e r e s e a r c h c e n t e r s on e x t e n s i o n s and e l a b o r a t i o n s o f t h e o r i g i n a l L o t k a - V o l t e r r a c o m p e t i t i o n e q u a t i o n s . T h i s approach i s p a r t i c u l a r l y p o p u l a r i n p o p u l a t i o n eco logy . I n community and ecosystem eco logy, t h e dominant models c u r r e n t l y used a r e compartment models. They deal w i t h s t o r a g e s and f l o w s o f energy and m a t e r i a l s i n systems and have been u s e f u l l y complementary t o chemical and r a d i o b i o l o g i c a l t echn iques developed and u t i l i z e d i n eco logy d u r i n g t h e l a s t twen ty y e a r s . They w i l l c o n t i n u e t o be i m p o r t a n t , d e s p i t e sugges t ions t h a t e c o l o g i s t s expand t h e i r mathemat ica l h o r i z o n s (e .g . Clymer, 1972) p r e c i s e l y because o f t h i s c o m p a t i b i l i t y w i t h c u r r e n t and p r o s p e c t i v e exper imen ta l methodology. The c u r r e n t s t a t u s o f systems model ing i n eco logy i s summarized by P a t t e n (1971 , 1972). A model f r o m a g e o l o g i c a l v i e w p o i n t i s p resen ted by M a c i n t y r e u ,1974. The I n t e r n a t i o n a l B i o l o g i c a l Program ( IBP) has been i n s t r u m e n t a l i n a c c e l e r a t i n g t h e pace o f development o f t o t a l ecosystem mode l ing ( i n p a r t i c u l a r w i t h t h e A n a l y s i s o f Ecosystems P r o j e c t o f t h e U.S. IBP) . A1 1 t h e Biome Programs (Grass1 and, Deciduous Fores t , Deser t , Tundra, Coni ferous Fores t , e t c . ) a r e commit ted t o some fo rm o f mathemat ica l ecosystem a n a l y s i s . P rog ress t o d a t e has been v a r i a b l e , and t h e degree o f emphasis a l s o d i f f e r s f rom program t o program. The Grassland Biome Program i s e x p l i c i t l y engaged i n t o t a l ecosystem mode l ing ( c f . Bledsoe u., 1971) . Others (e.g. Deciduous F o r e s t , D e s e r t ) a r e emphasiz ing process models as p o t e n t i a l modules f o r even tua l t o t a l ecosystem models. 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