UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

The biology of the host-specific epiphytic red alga Ceramium codicola and some other epiphytes of Codium… Lewis, Raymond Jerald 1982

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1982_A6_7 L49.pdf [ 3.4MB ]
Metadata
JSON: 831-1.0098709.json
JSON-LD: 831-1.0098709-ld.json
RDF/XML (Pretty): 831-1.0098709-rdf.xml
RDF/JSON: 831-1.0098709-rdf.json
Turtle: 831-1.0098709-turtle.txt
N-Triples: 831-1.0098709-rdf-ntriples.txt
Original Record: 831-1.0098709-source.json
Full Text
831-1.0098709-fulltext.txt
Citation
831-1.0098709.ris

Full Text

THE BIOLOGY OF THE HOST-SPEC IF IC EP IPHYTIC RED ALGA CERAMIUM COD I COLA AND SOME OTHER EPIPHYTES OF CODIUM SPP. IN B R I T I S H COLUMBIA by RAYMOND JERALD LEWIS A . B . , The U n i v e r s i t y of C a l i f o r n i a , B e r k e l e y , 1979 A THES I S SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS  FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Botany Department) We accept t h i s t h e s i s as con fo rm ing to the r e q u i r e d s t a n d a r d THE UN IVERS ITY OF B R I T I S H COLUMBIA May, 1982 © RAYMOND JERALD LEWIS , 1982 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of Botany The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date 2 8 J u l? 1 9 8 2 DE-6 (3/81) i i R e s e a r c h S u p e r v i s o r : D r . Rona ld E . Foreman ABSTRACT F i e l d and c u l t u r e s t u d i e s have been c a r r i e d out w i t h the e p i p h y t i c red alga.,. Ce r am i um cod i co l a J . A g a r d h T w i t h the o b j e c t i v e of de t e rm in i n g what f a c t o r s in i t s b i o l o g y cause it to be l i m i t e d to growth on one host p lant , . Cod i um f r a g i l e ( S u r i n g a r ) H a r i o t . A s u r ve y of e p i p h y t e s of C^ f r a g i l e and C ^ se tc he I I i i c o n f i r m the absence of C^ cod i co I a on C^ se tc he I I i i and shows that C^ cod i co l a i s the most abundant ep i phy te on C^ fr ag i I e . More s p e c i e s of e p i p h y t e s were p r e s e n t in wave s h e l t e r e d a r e a s . C^ cod i co I a was not p re sen t in samples from the S t r a i t of G e o r g i a . In culture.,. C^ cod i co I a grew to r e p r o d u c t i v e m a t u r i t y apar t from C. fr ag i I e from s p o r e s T but the l i f e h i s t o r y was not comple ted . The p re sence of l i v i n g or e x t r a c t e d C ^ fr ag i I e or C se t che I I i i d i d not promote g e r m i n a t i o n , g rowth or r e p r o d u c t i o n of C. c o d i c o l a T and t y p i c a l l y i n h i b i t e d i t . C. cod i co I a d i d not grow wel l at temperatures above 15°C in c u l t u r e ; C . fr ag i I e, Cer ami um pac i f i cum and C ^ gardner i grew we I I at 1 0 - 18°C . R h i z o i d s of C ^ cod i co l a have bu lbous tips.,, wh ich appears to be g e n e t i c a l l y de te rm ined . S u b s t r a t u m can a f f e c t l a te r development of these r h i z o i d s . , wh i ch may ob s cu re the bu lbous na tu re of the r h i z o i d t i p s . C. cod i co I a i s unab le to produce d i g i t a t e h o l d f a s t s on i t s r h i z o i d tips-, as was ob se rved in other Ceramium s p p . T and may o n l y be able to a t t ach by e n t a n g l i n g i t s r h i z o i d s in l o o s e l y c o n s t r u c t e d s u b s t r a t a such as Cod i um spp. C. se tche I I i i i s not s u i t a b l e because it i s too compact., and p e n e t r a t i o n of r h i z o i d s i s i n h i b i t e d . i i i F i e l d s t u d i e s show that C^ cod i c oI a i s r e p r o d u c t i v e th roughout the year.,, a l t h o u g h reduced in the w i n t e r . T h a l l u s l eng th was a l s o reduced in the w i n t e r . In a d d i t i o n , tha i I i were longer in wave s h e l t e r e d s i t e s , and longer in the upper r e g i o n s of t h e i r i n t e r t i d a l d i s t r i b u t i o n . Ma le and female gametophytes and t e t r a s p o r o p h y t e s of C. c o d i c o l a e x h i b i t a he te rmorph i sm in b r a n c h i n g p a t t e r n . Gametophytes d i f f e r from t e t r a s p o r o p h y t e s by h a v i n g more p r ima r y b ranches and l a r ge r ang le s of b r a n c h i n g . Females d i f f e r from males by h a v i n g a h i ghe r degree of p r o l i f e r o u s b ranch ing , , r e s u l t i n g in a h i gher number of p r imary b ranches and a h i ghe r b r a n c h i n g r a t i o . B r a n c h i n g p a t t e r n in females may be the r e s u l t of c o n t i n u e d growth of i n v o l u c r a l r a m u l i T and i t may op t im i ze r e p r o d u c t i v e po ten t i a I . TABLE OF CONTENTS Pag ABSTRACT i TABLE OF CONTENTS i L I S T OF TABLES L I S T OF F IGURES v ACKNOWLEDGEMENTS v i i INTRODUCTION MATERIALS AND METHODS RESULTS 1 A . E p i p h y t e s of Cod i um spp 1 B. C u l t u r e e x p e r i m e n t s . . . . C. R h i z o i d morpho logy D. P h e n o l o g i c a l phenomena. E . B r a n c h i n g p a t t e r n D I S C U S S I O N A . E p i p h y t e s of Cod i um spp B. C u l t u r e e x p e r i m e n t s . . . . C. R h i z o i d morpho logy D. P h e n o l o g i c a l phenomena. E . B r a n c h i n g p a t t e r n SUMVIARY L ITERATURE C ITED L I S T OF TABLES Tab le Page I . S i t e s and months of c o l l e c t i o n s of Cod i um f r a g i I e and C ^ s e t c h e I I i i in the v i c i n i t y of B a m f i e l d , and wave exposure of s i t e s . 10 I I . E p i p h y t e s of Cod i um f r a g i I e c o l l e c t e d in the v i c i n i t y of Bamf i e I d , p r e v i o u s l y r epo r t ed s u b s t r a t a , months of c o l l e c t i o n and wave expo su re of c o l l e c t i o n s i t e s . 18 I I I . E p i p h y t e s of Codi um s e t che I I i i c o l l e c t e d in the v i c i n i t y of B a m f i e l d , p r e v i o u s l y r epo r t ed s u b s t r a t a , months of c o l l e c t i o n and wave exposure of c o l l e c t i o n s i t e s . 19 IV . E p i p h y t e s of Cod i um f r a g i I e c o l l e c t e d from Hornby I s l a n d , l i s t e d from most to l e a s t abundant . 21 V . E p i p h y t e s of e p i p h y t i c Cerami um cod i c o I a and C ^ pac i f i cum, months of c o l l e c t i o n and wave exposure of c o l l e c t i o n s i t e s . 21 V I . F e r t i l i t y of c u l t u r e s of Cerami um cod i c o I a i n i t i a t e d from c a r p o s p o r e s and t e t r a s p o r e s at v a r i o u s t e m p e r a t u r e s . 26 V I I . I n i t i a l number of s p o r e s , % g e r m i n a t i o n and maximum l eng th s of Cerami um c o d i c o l a s p o r e l i n g s a f t e r 14 days of growth in e x t r a c t s of Cod i um f r a g i I e and C ^ se tche I I i i . 32 V I I I . D i amete r s of r h i z o i d s of Ceramium spp. and M i c r o c l a d i a c o u l t e r i from f i e l d - c o l l e c t e d and c u l t u r e d ( f rom spo re s or r egene ra ted from a p i c a l segments ) s pec imen s . 37 I X . F - v a l u e s and a s s o c i a t e d p r o b a b i l i t i e s of s i g n i f i c a n c e ob ta i ned from two-way anovas w i t h t rea tment s of month, s i t e and m o n t h - s i t e i n t e r a c t i o n on f r e q u e n c i e s of r e p r o d u c t i v e c l a s s e s . 42 X. F - v a l u e s and a s s o c i a t e d p r o b a b i l i t i e s of s i g n i f i c a n c e ob ta i ned from two-way anova s , one w i t h month and s i t e , the other w i t h he i gh t and s i t e as t reatments and l eng th s of p l a n t s of v a r i o u s r e p r o d u c t i v e c l a s s e s as the dependent v a r i a b l e . 49 X I . Mean number of p r imary b ranches and b r a n c h i n g o rde r s fo r 5 mm a p i c a l segments of t e t r a s p o r o p h y t e s , female gametophytes ( w i t h or w i t h o u t i n v o i u c r a l r a m u l i ) and male gametophytes of Ceramium c o d i c o l a . 58 V I L I S T OF FIGURES F i g u r e Page 1. S u r f a c e v iew of a) Cod i um fr ag i I e and b) C ^ se tc he I I i i showing u t r i c l e s . 4 2. S i t e s of col l e c t i o n of Co d i um fr ag i I e and se tche I I i i in the v i c i n i t y of B a m f i e l d . 12 3 . G e r m i n a t i o n p a t t e r n of spo re s of Ceramium cod i c o l a in c u l t u r e . 24 a. Spore be fo re d i v i s i o n . b . U n e q u a l l y d i v i d e d s p o r e . c . S p o r e l i n g showing d e v e l o p i n g r h i z o i d . d . S p o r e l i n g showing ap i ca l development and bu lbous r h i z o i d . e. S p o r e l i n g showing narrow r h i z o i d s g rowing out of s w o l l e n r h i z o i d . 4 . Pe r cen tage of g e r m i n a t i o n and s u r v i v a l of t e t r a s p o r e s and c a r p o s p o r e s of Cerami um cod i c o l a at 1 0 T 15 and 20°C w i t h t ime. 28 5 . Mean l eng th s of t e t r a s p o r e I i n g s and c a r p o s p o r e I i n g s of Cer ami um cod i co I a grown in c u l t u r e at 10 and 15°C at v a r i o u s t imes o f g r owth . 31 6. C o r t i c a t i o n o f Ceramium p a c i f i c u m from the f i e l d and in c u l t u r e . 36 a. Specimen c o l l e c t e d from the f i e l d as an ep iphy te of Cod i um fr ag i I e showing complete c o r t i c a t i o n . b. Specimen grown from a c a r po spo re in c u l t u r e showing naked i n ter nodes . c . Same as b ) T showing c o r t i c a t i o n d e v e l o p i n g more f u l l y f a r t h e r from the apex. 7. R h i z o i d m o r p h o l o g y . 40 a. D i ag ram of d i g i t a t e h o l d f a s t s t r u c t u r e produced on the t i p s of r h i z o i d s of some Ceramium spp. in c u l t u r e . b. R h i z o i d s o f M i c r o c I ad i a cou I ter i from a s p o r e l ing grown in c u I t u r e . c . R h i z o i d s of a specimen of Ce r am i um pac i f i cum c o l l e c t e d as an ep i phy t e of Cod i um f r a g i l e w i t h r h i z o i d s a r i s i n g away from the base of the p l a n t . V I I F i g u r e Page 8. Mean f r e q u e n c i e s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s of Ceramium c o d i c o l a e xp r e s s ed as a pe rcen tage of samples c o l l e c t e d at F i r s t Beach and S e p p i n g s I s l a n d in the months of the year 1981 . 44 9 . Mean f r e q u e n c i e s of a) n o n - c y s t o c a r p i c and b) c y s t o c a r p i c female gametophytes of Ceramium c o d i c o l a at F i r s t Beach and S e p p i n g s I s l a n d , e xp r e s s ed as pe rcen tage of samples c o l l e c t e d in the months of the year 1981. 47 10. Mean l eng th s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s of Ceramium cod i c o l a f rom F i r s t Beach and S epp i n g s I s l a n d from spec imens c o l l e c t e d in the months of the year 1981 . 51 -52 11. Mean l eng th s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s of Ceramium c o d i c o l a f rom F i r s t Beach (FB) and S epp i n g s I s l a n d ( S I ) from m i d - (open c i r c l e s ) and I o w - i n t e r t i d a I ( c l o s e d c i r c l e s ) r e g i o n s . 54 12. R e p r e s e n t a t i v e 5-mm a p i c a l segments of a) a female gametophyte, b) a t e t r a s p o r o p h y t e , and c) a male gametophyte of Ce r am i um cod i co I a showing o v e r a l l b r a n c h i n g p a t t e r n s . 56 13. Mean r a t i o s of h i ghe r to lower order branches of a) female g ametophy te s - l ( i n c l u d i n g i n v o l u c r a l r a m u l i ) , b) female g a m e t o p h y t e s - l I ( e x c l u d i n g i n v o l u c r a l r a m u l i ) , c) t e t r a s p o r o p h y t e s and d) male gametophy te s . 60 ACKNOWLEDGEMENTS I would l i k e to thank D r . R . E . Foreman for p r o v i d i n g a d v i c e , f a c i l i t i e s for r e s e a r c h , a n a l y s i s of data and w r i t i n g of t h i s t h e s i s at the B a m f i e l d Ma r i ne S t a t i o n , and f i n a n c i a l suppor t ( t h r o u g h a B .C . S c i e n c e C o u n c i l g r a n t ) . A d d i t i o n a l f i n a n c i a l suppor t was p r o v i ded by a U . B . C . g r adua te f e l l o w s h i p , for wh ich I am t h a n k f u l . Thank you to Dr s . R . E . deWreede and D. Ga rba r y for s u g g e s t i n g improvements in the m a n u s c r i p t and for ready a v a i l a b i l i t y d u r i n g my b r i e f v i s i t s to campus. Anong the people I am indebted to are E r i c Cabot for s t a t i s t i c a l c o n s u l t a t i o n and p r o v i s i o n of computer programs for a n a l y s e s ; the s t a f f of the Bamf i e l d Ma r i ne S t a t i o n ; V a l e r i e Harpham for e x p e r t l y d r a u g h t i n g the map and g r a p h s ; Lynn Y i p and R i c k Cohen for p h o t o g r a p h i c a s s i s t a n c e ; and C h r i s t i n e M i l l i k e n for s u g g e s t i n g s t y l i s t i c improvements of t h i s t h e s i s . I would a l s o l i k e to thank Laura R i c h a r d s for her c o n s t r u c t i v e remarks and f r i e n d s h i p . I thank my p a r e n t s whose ca re (and care p a c k a g e s ) , so s t r o n g l y f e l t over a long d i s t a n c e , have encouraged me to keep g o i n g . INTRODUCTION Ceramium cod i c o I a was d e s c r i b e d by J . Agardh ( 1894 ) as an ep iphy te of Cod i um fr ag i I e ( S u r i n g a r ) Ha r i o t from San t a C r u z , C a l i f o r n i a . I t i s a smaI I ( 1 - 2 . 5 cm) comp le te l y c o r t i c a t e d s p e c i e s of Ceramium that i s known to grow o n l y on C ^ f r a g i l e ; a t tachment is by many p igmented r h i z o i d s that pene t r a te between the u t r i c l e s of Cod i um and have bu lbous t i p s . I t has been found to occur on the P a c i f i c Coa s t over the e n t i r e range of C^ f r a g i l e , from S i t k a , A l a s k a to I s I a San B e n i t o , Baja C a l i f o r n i a (Abbot t and H o l l e n b e r g 1 9 7 6 ) . Cod i um f r a g i l e i s a s i phonou s green a l g a that i s known to occur in many temperate s e a s . There are a l s o f i v e other Codium spp. that occur on the P a c i f i c Coa s t of N o r t h A n e r i c a : C ^ se t che I I i i Gardner from S i t k a , A l a s k a to Punta B a j a , Ba ja C a l i f o r n i a ; C^ cuneatum S e t c h e l l and G a r d n e r , hubbs i i Dawson and C ^ j ohn s tone i S i l v a a l l found e x c l u s i v e l y south of P o i n t C o n c e p t i o n ( S i l v a , 1 9 5 1 ) ; and C ^ r i t t e r i S e t c h e l l and Gardner from A t t u I s l a n d to Campania I s l a n d , n o r t h e r n B r i t i s h Co lumbia and p o s s i b l y on the West Coa s t of Vancouve r I s l a n d (Hawkes, al . , 1 9 7 8 ) . Of these Cod i um s p p . , C. fr ag i I e i s the most common, f o l l owed by C^ setche I I i i . The o the r s are found on l y o c c a s i o n a l l y . Thai I i of Cod i um have a spongy c o n s t r u c t i o n , be i ng composed of c o e n o c y t i c f i l a m e n t s woven toge the r in to a f a i r l y m a s s i v e t h a l l u s w i t h e n l a r g e d b r a n c h l e t s ( u t r i c l e s ) f o rm ing the s u r f a c e . S i l v a (1951 ) comments on the r i c h a l g a l f l o r a found on C^ fr ag i I e and a t t r i b u t e s t h i s to i t s loose c o n s t r u c t i o n and the a b i l i t y of a l gae to en tang le in i t . The u t r i c l e s of C. f r a g i I e are unbranched and a p p r o x i m a t e l y 1 50 - 350 jjm w ide , w h i l e u t r i c l e s o f C^ s e t c h e I I i i are h i g h l y branched and 65 - 90 jjm w ide , fo rming a more compact t h a l l u s ( S i l v a 1951; a l s o see F i g u r e 1 ) . Abbot t and H o l l e n b e r g ( 1976 ) l i s t seven s p e c i e s of a l gae c h a r a c t e r i s t i c a l l y found e p i p h y t i c on C. fr ag i I e , but none on C^ se t che I I i i . L i t t l e and Jones ( 1 980 ) d e f i n e an ep iphy te as "an o r g a n i s m that grows on another p l a n t but i s not p a r a s i t i c on i t . " E p i p h y t e s in t e r r e s t r i a l e c o s y s t e m s , p a r t i c u l a r l y in t r o p i c a l f o r e s t s , i n c l ude v a s c u l a r p l a n t s such as o r c h i d s , b r o m e l l i a d s and f e rn s that are adapted to o b t a i n i n o r g a n i c n u t r i e n t s and m o i s t u r e apar t from s o i l (Odum 1 9 7 1 ) . However, a l gae do not r e q u i r e s o i l to grow, o b t a i n i n g i n o r g a n i c n u t r i e n t s and m o i s t u r e d i r e c t l y from the water in wh ich they l i v e , r e g a r d l e s s of t h e i r attachment s i t e . T h e r e f o r e , aqua t i c a l g ae can grow on s eve ra l s u b s t r a t a and many are found as e p i p h y t e s . The e p i p h y t e - h o s t r e l a t i o n s h i p can be c o n s i d e r e d a case of s y m b i o s i s , wh i ch l i t e r a l l y means " l i v i n g t o g e t h e r " and i n c l u d e s such i n t e r a c t i o n s as p a r a s i t i s m , commensal i sm and m u t u a l i s m (Odum 1 9 7 1 ) . E p i p h y t i s m is g e n e r a l l y c o n s i d e r e d to be commensaI i sm, in wh ich the e p i p h y t e , or commensal, o b t a i n s a p l a ce to a t t a c h and the host i s u n a f f e c t e d . Many e p i p h y t i c a l gae are l i m i t e d to growth on one or a few host s p e c i e s . T h i s i n d i c a t e s that they have r equ i r ement s that on l y a few h o s t s can p r o v i d e . These may i n v o l v e chemical a s pec t s of the h o s t - e p i p h y t e i n t e r a c t i o n , the p h y s i c a l nature of the h o s t , and other f a c t o r s . Chemica l a s p e c t s of the h o s t - e p i p h y t e i n t e r a c t i o n may i n v o l v e t r a n s l o c a t i o n of n u t r i e n t s between p l a n t s , p r o d u c t i o n of n o n - n u t r i t i v e s t i m u l a t o r y s u b s t a n c e s by the h o s t , and p r o d u c t i o n of i n h i b i t o r y compounds by the h o s t . Each of these may a f f e c t which e p i p h y t e s grow on the h o s t , and t h e r e f o r e a f f e c t s p e c i f i c i t y . F i g u r e 1. S u r f a c e v iew of a) Cod i um f r a g i I e and b) C ^ s e t c h e I I i i showing u t r i c l e s . S c a l e = 1 (mi. D e m o n s t r a t i o n of t r a n s l o c a t i o n of n u t r i e n t s from host to ep i phy te would i n d i c a t e that the i n t e r a c t i o n between s p e c i e s i s p a r a s i t i c r a the r than commensal. S e t c h e l l ( 1918 ) c o n s i d e r e d th ree m o r p h o l o g i c a l c r i t e r i a to dete rmine the p a r a s i t i c na tu re of red a l g a e : 1) p e n e t r a t i o n , 2 ) r e d u c t i o n of t h a l l u s and 3) l o s s of c o l o r . T r a n s l o c a t i o n of e i t h e r i n o r g a n i c n u t r i e n t s or p h o t o s y n t h a t e has been demons t ra ted to occur from host p l a n t s to ep i phy te s t ha t do not meet these c r i t e r i a of p a r a s i t i s m ( L i n s k e n s 1 963a , C i t h a r e l 1972 , H a r l i n 1973b, Penot 1974 , H a r l i n and C r a i g i e 1975 , Turner and Evans 1 9 7 7 ) . In some c a s e s , t r a n s l o c a t i o n from ep iphy te to host was a l s o o b s e r v e d . H a r l i n ( 1973b) emphas i zes that t r a n s f e r to and from the medium is an important i n d i r e c t route of t r a n s l o c a t i o n . T h i s s hou l d not be c o n s i d e r e d a case of p a r a s i t i s m , because the host may not be n e g a t i v e l y a f f e c t e d and adjacent but s e p a r a t e l y a t t a ched p l a n t s may a l s o e x h i b i t s i m i l a r t r a n s l o c a t i o n . One-way t r a n s l o c a t i o n of n u t r i t i v e s ub s t ance s has been demonst ra ted to occur from hos t to a l g a l p a r a s i t e s that do meet S e t c h e l l ' s c r i t e r i a (Evans et a l . 1973, G o f f 1 9 7 9 ) , but not in e p i p h y t e s that do not , c o n f i r m i n g t h e i r n o n p a r a s i t i c n a t u r e . N o n - n u t r i t i v e s u b s t a n c e s produced by s p e c i f i c h o s t s have been h y p o t h e s i z e d to a f f e c t the growth and development of two red a l g a l e p i p h y t e s . E r y t h r o c y s t i s s a c c a t a has been grown to r e p r o d u c t i v e m a t u r i t y in c u l t u r e apa r t from i t s normal h o s t . However, normal morpho logy was o n l y ob ta i ned in the p re sence of i t s host (Gonza l e z and G o f f 1 9 8 0 ) . J anczews k i a mor imoto i , a semi par a s i t i c red a l g a , would not ge rminate w i thou t a s u i t a b l e host p r e s e n t . However, s po re s in the p re sence o f , but s epa ra te from, a s u i t a b l e host grew to r e p r o d u c t i v e m a t u r i t y . The s ub s t ance that i n i t i a t e d t h i s re sponse was found to pass t h rough a d i a l y s i s membrane that would a l l o w m o l e c u l e s of 6 000 - 8000 Dal tons to pass (Nonomura and West 1 9 8 1 b ) . The growth of s eve ra l other e p i p h y t e s have been s t i m u l a t e d or m o d i f i e d by the p re sence of a s p e c i f i c host or host e x t r a c t ( W i l l i a m s 1963 , Boney 1980 , Hal lam et al . 1 9 8 0 ) . Some p l a n t s are known to produce s ub s t ance s which are i n h i b i t o r y to o ther o r g a n i s m s , i n c l u d i n g e p i p h y t e s ( e . g . S a r ga s s um, Conover and S i e b u r t h 1964; A s p a r a g o p s i s a rmata , Codomier et al . 1 9 8 1 ) . An ep iphy te o c c u r r i n g on such a p l an t must be able to t o l e r a t e these s u b s t a n c e s . S e v e r a l p h y s i c a l p r o p e r t i e s of hos t p l a n t s have been b r i e f l y s t u d i e d w i t h r e s p e c t to e p i p h y t i s m , i n c l u d i n g s u r f a c e t e x t u r e ( L i n s k e n s 1 9 6 6 ) , s u r f a c e t e n s i o n ( L i n s k e n s , 1963b) and g e l a t i n o u s c h a r a c t e r ( B a l l a n t i n e 1 9 7 9 ) . H a r l i n (1973a) found that Smi t ho r a na i adum, wh ich n o r m a l l y grows on sea g r a s s e s , would a l s o grow on p l a s t i c sea g r a s s which p h y s i c a l l y approx imates rea l sea g r a s s . She s p e c u l a t e d that S ^ na i adum i s unable to a t t a ch to host p l a n t s w i t h g e l a t i n o u s s u r f a c e s . B i r d et a l . ( 1982 ) found Pt i I o thamn ionops i s I e j o I i sea g row ing on four s p e c i e s of a r t i c u l a t e d c o r a l l i n e red a lgae and on Cod i um s e t c h e I I i i . Each of the c o r a l l i n e h o s t s p o s s e s s e d l a r ge r g e n i c u l a and was le s s~ b r anched , f o rm ing l oo se r clumps than d i d other c o r a l l i n e s p e c i e s t ha t were not e p i p h y t i z e d . T h i s may a l l o w p e n e t r a t i o n of the endophy t i c ba sa l p o r t i o n of P ^ Ie j o I i s e a . T h i s s u g g e s t s that p h y s i c a l c h a r a c t e r i s t i c s o f the host are important in d e t e r m i n i n g host s p e c i f i c i t y in P ^ Ie j o I i s e a . S e v e r a l other f a c t o r s may a f f e c t e p i p h y t i s m . Some p l a n t s shed t h e i r oute rmost layer of c e l l s , wh ich would remove e p i p h y t e s (Ende und O o r s c h o o t 1962, Moss 1 9 8 1 ) . G r a z e r s may a l s o remove e p i p h y t e s ( S l o cum 1 9 8 0 ) . The p h y s i c a l env i ronment in which the p o t e n t i a l host grows may l i m i t which p l a n t s can occur as e p i p h y t e s . H e i g h t in the i n t e r t i d a l zone, w i t h accompanying d e s i c c a t i o n , temperature and s a l i n i t y i n f l u e n c e s , and wave exposure are two major f a c t o r s to c o n s i d e r on mar ine s h o r e s . Temporal d i s t r i b u t i o n of both host and ep iphy te are a l s o important because a s u c c e s s f u l ep i phy te must be able to c o l o n i z e i t s h o s t T grow to m a t u r i t y and reproduce be fo re the host d i e s (den Ha r tog 1 9 7 2 ) . Therefore. , s e a s o n a l i t y in co I o n i z a t i o n T g rowth and r e p r o d u c t i o n may be e s p e c i a l l y impor tant in the l i f e of an e p i p h y t e . Go f f and Co l e ( 1976 ) have demonst rated tha t the red a l g a l p a r a s i t e Ha rveye I la mi rab i I i s e x h i b i t s a seasona l a l t e r n a t i o n of g e n e r a t i o n s . Nonomura and West (1981a) found la rge seasona l changes in the b iomass and r e p r o d u c t i o n of the p a r a s i t e J anc zewsk i a mo r imoto i in Hokka i do and sma l l e r changes in J . g a r d n e r i in C a l i f o r n i a , but no seasona l a l t e r n a t i o n of g e n e r a t i o n s . A s ea sona l a l t e r n a t i o n of g e n e r a t i o n s was demons t ra ted to occur in the l i f e h i s t o r y of s e ve r a l red a lgae in the v i c i n i t y of Woods Ho le ( Lew i s 1914 ) and in Ce r am i um d i aphanum in the B a l t i c Sea ( S v e d e l i u s 1929, Du R i e t z 1 9 3 0 ) . The r e s e a r c h r epo r ted in t h i s t h e s i s was under taken to determine the f o I Iowi ng : 1) What e p i p h y t e s occur on Co d i um fr ag i I e and C ^ se tche I I i i in v a r i o u s s i t e s in B a r k l e y Sound? T h i s was determined to compare the e p i f l o r a of C . fr ag i I e and C ^ se t che I I i i in a reas exposed to v a r i o u s l e v e l s of wave ac t i on , , and to sea rch for C ^ cod i co l a on C^ se tc he I I i i . 2) Wi I I C ^ cod i c o I a grow in l a b o r a t o r y c u l t u r e apart from C ^ fr ag i I e and i s g rowth m o d i f i e d or s t i m u l a t e d by the p re sence of C ^ fr ag i I e or C . se tc he I I i i ? T h i s s hou l d e s t a b l i s h whether or not C. c o d i c o l a i s c h e m i c a l l y dependent on fr ag i I e and whether chemical i n t e r a c t i o n i s impor tant for growth,, or lack t h e r e o f , of C^ cod i co l a on C^ se tc he I I i i . 3) Does C^ cod i c oI a e x h i b i t any m o r p h o l o g i c a l s p e c i a l i z a t i o n in i t s a t tachment o rgan compared to other Ceramium spp. that may enable i t to a t t ach to C ^ fr ag i I e ? O b s e r v a t i o n s from specimens c o l l e c t e d in the f i e l d and grown in c u l t u r e s hou ld e s t a b l i s h c h a r a c t e r i s t i c s of s u i t a b l e s u b s t r a t a for secure at tachment of these v a r i o u s Ceramium spp. 4) Does r e p r o d u c t i o n and p l a n t s i z e of C^ cod i c o I a e x h i b i t any d i f f e r e n c e s due to time of y e a r , degree of wave e xpo su r e , or he i gh t in the i n t e r t i d a l zone? T h i s s h o u l d demonst rate the manner in which C^ cod i c o I a d e a l s w i t h p o t e n t i a l l y adve r se c o n d i t i o n s . 5) Does C^ cod i c o I a e x h i b i t he te romorph i sm in b r a n c h i n g p a t t e r n in d i f f e r e n t r e p r o d u c t i v e phases s i m i l a r to that ob se r ved by G a r b a r y , Grund and McLach l an (1980 ) fo r C^ rubrum? How is t h i s e xp re s s ed in female and male gametophytes and t e t r a s p o r o p h y t e s of C^ cod i c o I a ? MATERIALS AND METHODS Cod i um f r a g i I e and C ^ se tche I I i i were c o l l e c t e d in the m i d - to low-i n t e r t i d a l zone on shore at low t i de or by SCUBA in the v i c i n i t y of B amf i e l d M a r i n e S t a t i o n (see Tab l e I and F i g u r e 2 f o r c o l l e c t i o n s i t e s and d a t e s ) and C. fr ag i I e f rom Hornby I s l a n d in the S t r a i t of G e o r g i a by SCUBA on 17 J u l y 1981. H e i g h t in the i n t e r t i d a l zone was e s t ima ted from the he i gh t r e l a t i v e to water l eve l at low t i de and is e xp r e s s ed as he i gh t above or below the Canad i an h y d r o g r a p h i c a I datum as determined from t i de t a b l e s (Anonymous 1980 , 1 9 8 1 ) . Wave exposure of these s i t e s was ranked on a s u b j e c t i v e s c a l e w i t h 1 = mode ra te l y exposed , 2 = m o d e r a t e l y s h e l t e r e d and 3 = f u l l y s h e l t e r e d . No spec imens of e i t h e r s p e c i e s of Cod i um was obse rved on shores f u l l y exposed to wave a c t i o n . M a c r o a l g a l e p i p h y t e s of both Cod i um spp. were i d e n t i f i e d u s i n g keys by Scage l ( 1 9 6 6 ) , Widdowson (1973 , 1 9 7 4 ) , Abbott and H o l l e n b e r g ( 1976 ) and G a r b a r y , Hansen and S cage l ( 1 9 8 0 ) . Two s i t e s were chosen for month l y s amp l i ng of C^ cod i c o I a on C^ fr ag i I e f rom Janua r y to December 1981 , one on rocks l oca ted southwest of F i r s t Beach a l o n g T r e v o r C h a n n e l , and the other on S e p p i n g s I s l a n d , f a c i n g the calm wa te r s of Dodger C h a n n e l . The F i r s t Beach s i t e is exposed to mode ra te l y heavy waves by s w e l l s that come in from the open P a c i f i c Ocean and many p l a n t s were to rn o f f in w i n te r s t o rms . The S e p p i n g s I s l a n d s i t e is exposed to some wave a c t i o n by s w e l l s coming in from Impe r i a l E a g l e Channe l , but is more s h e l t e r e d than F i r s t Beach. At both s tudy s i t e s , C^ cod i co I a was the most abundant ep i phy te on C^ f r a g i l e and both C^ f r a g i l e and Cj_ setche I I i i were p r e s e n t . Two hundred i n d i v i d u a l s of cod i co I a were sampled each month by s e p a r a t i n g 50 i n d i v i d u a l s of C^ cod i c o I a g r e a t e r than 1 . 5 mm in l eng th from each of four tha i I i of C. f r a g i l e from both s i t e s and examin ing them Tab le I . S i t e s and months of c o l l e c t i o n s of Cod i um fr ag i I e and C ^ se t che I I i i in the v i c i n i t y of B a m f i e l d and wave exposure of s i t e s . S I T E CODIUM MONTHS COLLECTED WAVE EXPOSURE 3 F i r s t Beach C. O f r ag i 1 e se tche 1 1 i i 6 /80 - 12/81 1 Sepp i ngs I s I and - I (Dodger C h a n n e I ) C. C ^ f r a g i l e se tche 1 1i i 6 /80 -12/81 2 Sepp i ngs I s I and - I I C f r a g i l e 6 - 7 / 8 0 ; 1 , 3 , 4/81 3 D i a n a I s l a n d * 3 (Dodger C h a n n e I ) fr ag i 1 e 7 , 11/80 ; 2 ,4/81 3 D i a n a I s I and ( K i r b y P t . ) fr ag i 1 e 2/81 1 D i ana I s 1 and (K i rby P t . , t i d e p o o l ) C ^ f r a g i l e 2/81 3 Second Beach f r a g i l e 6 , 9 - 1 2 / 8 0 ; 1 -6/81 3 Pachena Bay*3 C ^ fr ag i 1 e 6 , 7 , 9 / 8 0 ; 3/81° 2 Fo1ger 1s1 and c. c^  fr ag i 1 e se tche 1 1 i i 7/80 1 S c o t t ' s Bay c^  f r a g i l e 10/80 2 Di xon 1 s 1 and s e t c h e 1 1 i i 2/81 2 aWave e x p o s u r e : 1 = m o d e r a t e l y exposed , 2 = m o d e r a t e l y s h e l t e r e d , 3 = f u l l y s h e l t e r e d . ^No C ^ cod i co I a ob se r ved at these s i t e s . C^ pac i f i c um was p r e s e n t . c N o C ^ f r a g i l e p r e sen t at t h i s d a t e . F i g u r e 2. S i t e s of c o l l e c t i o n of Cod i um f r a g i l e and C ^ se tche I I i i in the v i c i n i t y of B a m f i e l d . m i c r o s c o p i c a l l y at 100X to dete rmine the p re sence of t e t r a s p o r a n g i a , c y s t o c a r p s , c a r p o g o n i a , or s p e r m a t a n g i a . A specimen l a c k i n g these s t r u c t u r e s was c l a s s i f i e d as a v e g e t a t i v e p l a n t . From J u l y to O c t o b e r , two C^ f r a g i l e p l a n t s were sampled from the h i ghe r r e g i o n s in the i n t e r t i d a l zone and two f rom the lower. These spec imens were examined f r e s h from m a t e r i a l m a i n t a i n e d in r u n n i n g seawater t a b l e s or p r e s e r ved in 5% fo rma ldehyde in seawater for l a t e r c o u n t i n g . The l eng th s of p l a n t s of C^ cod i c o I a sampled from March to December 1981 were measured to the nea re s t 0 . 1 mm. The f r e q u e n c i e s and l eng th s of the r e p r o d u c t i v e phases were ana l y zed by two- and th ree-way a n a l y s i s of v a r i a n c e (anova) w i t h r e p l i c a t i o n ( Soka l and R o h l f 1981 ) to detect any s i g n i f i c a n t d i f f e r e n c e s among months of c o l l e c t i o n , between s i t e s and between t i d a l h e i g h t s . R e p r o d u c t i v e phase f requency data was c onve r t ed to p r o p o r t i o n s and t r an s fo rmed by a r c s i n ( s q r t ) x as recommended by Soka l and R o h l f ( 1 9 8 1 ) . D i f f e r e n c e s r e vea l ed by anovas were e l u c i d a t e d u s i n g m u l t i p l e unp lanned compar i son s among p a i r s of means ( Soka l and R o h l f 1 9 8 1 ) . C o r r e l a t i o n a n a l y s i s was used to detect r e l a t i o n s h i p s between f r e q u e n c i e s of d i f f e r e n t r e p r o d u c t i v e g r o u p s . A l l s i g n i f i c a n t d i f f e r e n c e s in the text are at p < .05 u n l e s s o t he rw i s e no ted . I per formed these a n a l y s e s on an App le I I P l u s mic rocomputer w i t h programs prepared by E. Cabo t . A l g a e were grown in l a b o r a t o r y c u l t u r e in P r o v a s o l i ' s ES medium (PES) (McLach l an 1973) w i t h 10 mg/1 G e 0 2 added to prevent d ia tom growth (Lewin 1 9 6 6 ) . The medium was f i l t e r e d th rough a Whatman GF/D g l a s s f i b r e f i l t e r , then th rough a 0 . 4 5 ^im membrane f i l t e r ( S c h l e i c h e r and S c h u e l l or M i l l i p o r e C o r p . ) that had been washed i n ho t d i s t i l l e d wa t e r , at a vacu um of 10 p s i w i t h a vacuum pump. A l l g l a s s w a r e used for the c u l t u r e of a l gae or as medium c o n t a i n e r s were t r e a t e d w i t h a 10% aqueous s o l u t i o n of c o n c e n t r a t e d h y d r o c h l o r i c a c i d for 24 hou r s or l o n g e r , f o l l owed by r i n s e s in tap water and d i s t i l l e d wa te r , and were a u t o c l a v e d at 15 p s i steam p r e s s u r e (121 °C) for 15 m inu te s or l o n g e r . Pyrex deep s t o r a g e d i s h e s (100 X 80 rrm, C o r n i n g No. 3250 ) and p o l y s t y r e n e d i s p o s a b l e c u l t u r e d i s h e s (100 X 15, 60 X 15, and 100 X 25 mm) were used as c u l t u r e c o n t a i n e r s . Pyrex c u l t u r e d i s h e s were washed in a 2% aqueous s o l u t i o n of L i q u i - N o x ( p h o s p h a t e - f r e e d e t e r g e n t , A l c o n o x , I n c . ) o v e r n i g h t f o l l owed by a soak of one hour or longer in tap w a t e r , r i n s e s in d i s t i l l e d water and a u t o c l a v i n g . A l l c u l t u r e s were grown in P e r c i v a l u p r i g h t c u l t u r e chambers on a 1 6 : 8 hour LD c y c l e u s i n g c o o l - w h i t e f l u o r e s c e n t l i g h t s (Genera l E l e c t r i c F20T12-CW or F24T12-CW-HO) at an i r r a d i a n c e of a p p r o x i m a t e l y 30 pE/m^/s as determined by a L i - C o r quantum meter ( L I - 1 8 5 A ) w i t h a L I—190S sen so r he ld l e v e l . D u r i n g the cou r se of t h i s s t u d y , a l gae were c u l t u r e d at 5 , 10, 12, 15, 18, and 20°C. C u l t u r e s of Ce r am i um spp. and Mi c roc I ad i a cou I ter i were i n i t i a t e d from s po r e s or v e g e t a t i v e a p i c a l segments . S p o r u l a t i n g m a t e r i a l was washed in s t e r i l e PES medium befo re be ing p l aced over a c o v e r s l i p in PES medium u n t i l s u f f i c i e n t spore r e l e a s e was o b t a i n e d . Segments of t e t r a s p o r i c p l a n t s were washed by the " d i p and d r a g " method d e s c r i b e d in Tatewaki and P r o v a s o l i ( 1964 ) in 0 . 5 or 0.75%  agar in s t e r i l e medium befo re wash ing in s t e r i l e medium. I r e co rded the number of spo re s r e l e a s e d per d i s h , the number of these spo re s that ge rmina ted and, p e r i o d i c a l l y , the l eng th s of s p o r e l i n g s . C u l t u r e medium was changed every 1 0 - 1 6 d a y s , depend ing on the amount of b iomass p r e s e n t , the s i z e of the c u l t u r e v e s s e l and whether or not c on tam inan t s were p r e s e n t . C. cod i c o I a was grown on Cod i um spp. by r e l e a s i n g spores of C^ cod i c o I a d i r e c t l y onto the s u r f a c e s of Cod i um tha i I i . R e l a t i v e l y e p i p h y t e - f r e e p i e ce s of C^ f r a g i I e a p i c e s and C ^ s e t c h e I I i i ( a p p r o x i m a t e l y 1 - 2 cm^ of s u r f a c e area ) were cut w i t h a r a zo r b l a d e , t r ea ted in 2% Javex b leach in seawater for I m inu te s at 1 0 - 12°C , and washed in three changes of s t e r i l e seawater over a p e r i o d of 10 to 15 m i n u t e s . The above time and c o n c e n t r a t i o n of b leach t reatment was determined by t e s t i n g a s e r i e s of t imes and c o n c e n t r a t i o n s , and u s i n g the one that r e s u l t e d in the fewest s u r v i v i n g e p i p h y t e s w i thou t harming the Cod i um t h a l l u s . F ragments of Cod i um were t i ed onto g l a s s s l i d e s w i t h mono f i l ament n y l on l i n e in order to anchor them, because they tended to f l o a t in s t a t i c c u l t u r e , and grown in Py rex deep s t o r a ge d i s h e s w i t h a p p r o x i m a t e l y 250 ml of PES medum wh ich was changed week l y . In a d d i t i o n , C^ f r a g i l e was grown in c u l t u r e f ree of major c o n t a m i n a n t s . A s i n g l e p l an t a ro se in a c u l t u r e of C. c o d i c o l a , and may have s t a r t e d from a z ygo te or f r agment . I t was m a i n t a i n e d at 18°C for 3 months and at 12°C for 2 month s ; at the end of t h i s time it had grown into a mat of f i l a m e n t s , as d e s c r i b e d by A r a s a k i et a l . ( 1 955 ) and Borden and S t e i n ( 1 9 6 9 ) . Ramus ( 1972 ) found that a g i t a t i o n is n e c e s s a r y to o b t a i n the normal mo rpho l ogy of C. fr ag i I e in l a b o r a t o r y c u l t u r e . A g i t a t i o n was s u p p l i e d by a e r a t i o n from a smal l aquar ium pump (M in i -Comp MC/S, P e n n - P l a x , I n c . ) . W i t h i n 8 weeks , u t r i c l e s formed and the normal C^ fr ag i I e " f i n g e r s " were f o r m i n g . These c u l t u r e s remained u n i a l g a l and the " f i n g e r s " reached a p p r o x i m a t e l y 1 cm in l eng th (8 weeks l a t e r ) , when they were cut o f f and t i e d down on g l a s s s l i d e s in s epa ra te v e s s e l s for exper iment s w i t h C^ cod i c o I a . C u l t u r e of C ^ cod i c o I a was a l s o attempted in aqueous e x t r a c t s of C. fr ag i I e and C ^ se tche I I i i . I p l a ced 4 . 0 g of f r e e z e - d r i e d Cod i um in a l i t e r of PES medium for one hou r , then homogenized i t in a S o r v a l I Omni -Homogen izer for one m i n u t e . The r e s u l t i n g m i x t u r e was vacuum f i l t e r e d t h r ough four l a y e r s of cheese c l o t h , Whatman N o s . 4 , 1, and 3 paper f i l t e r s in s u c c e s s i o n , Whatman GF/D g l a s s f i b r e f i l t e r , and a 0 . 4 5 j^m membrane f i l t e r at 1 0 - 15°C . The f i l t r a t e was d i l u t e d w i t h PES medium to make s o l u t i o n s c o n t a i n i n g 100 , 50, 25 , 1 2 . 5 , 2 . 5 and 1 . 2 5 % e x t r a c t . S po re s r e l e a s e d onto c o v e r s l i p s were p l aced in 100 X 80 mn c u l t u r e d i s h e s w i t h 250 m l . of e x t r a c t . The w i d t h s of r h i z o i d s were measured to the nea re s t p n from f i e l d - c o l l e c t e d specimens of Ceramium s p p . , from v e g e t a t i v e ap i c a l segments wh i ch r e gene r a t ed r h i z o i d s in c u l t u r e and from s p o r e l i n g s in c u l t u r e at 500X on a L e i t z O r t h o p l a n l a r g e f i e l d compound m i c r o s c o p e . M i c r o p h o t o g r a p h s were taken u s i n g Kodak Pana tom i c - X f i l m . B r a n c h i n g p a t t e r n s of C^ cod i c o I a t e t r a s p o r o p h y t e s and male and female gametophytes were c h a r a c t e r i z e d by the method of S t r a h l e r ( 1953 ) as d e s c r i b e d by Barker et a l . ( 1 9 7 3 ) . F i v e mn a p i c a l segments of p l a n t s c o l l e c t e d in March 1981 at S e p p i n g s I s l a n d and A p r i l 1981 at F i r s t Beach were drawn on paper and the b r a n c h i n g c h a r a c t e r i z e d from these d r a w i n g s . Female gametophytes were c h a r a c t e r i z e d i n c l u d i n g and e x c l u d i n g the i n v o l u c r a l ramul i t ha t s u r r o u n d the c y s t o c a r p s . Q u a n t i t a t i v e  parameter s y i e l d e d by t h i s method were ana l y zed by the S t u d e n t ' s t - d i s t r i b i u t i o n . RESULTS A . Ep iphy tes on Codium spp. In B a r k l e y Sound, C^ f r a g i I e occu r s from a p p r o x i m a t e l y 0 . 3 to 1 . 8 m above datum on m o d e r a t e l y exposed to f u l l y s h e l t e r e d s h o r e s . C ^ s e t che l I i i appeared to occupy a narrower band at 0 to 0 . 5 m above datum, and it has been ob se r ved s u b t i d a l l y by SCUBA in t h i s area to a depth of 14 m (R . E . deWreede and J . Wat son , p e r s . comm.). C^ s e t che l I i i was found at mode ra te l y exposed and m o d e r a t e l y s h e l t e r e d s i t e s . M a c r o a l g a l e p i p h y t e s found in t h i s s tudy on C. f r a g i l e and C ^ s e t c h e l I i i in B a r k l e y Sound are l i s t e d in Tab I e s I I and I I I r e s p e c t i v e l y in approx imate order of abundance. The e p i p h y t i c a l gae were c h a r a c t e r i z e d as common or ra re based on the number of t imes they were ob se r ved as e p i p h y t e s . Rare e p i p h y t e s were those found on l y once or twice dur i ng t h i s s t u d y . A t o t a l of 30 taxa of a l gae were i d e n t i f i e d on C^ fr ag i I e , 13 o f which were c h a r a c t e r i z e d as common. Of the common e p i p h y t e s , C. cod i c o I a , E r y t h r o t r i ch i a pu I v i na ta and Audou i ne I la rh i z o i d e a are r epo r ted as n o r m a l l y o c c u r r i n g on C^ f r a g i l e . Cer ami um pac i f i cum i s o f t en found on C^ f r a g i l e , and it was a l s o found g row ing on mussel s h e l l s , other a l g a e , and r o c k s . I t was p r e sen t on C^ f r ag i I e in most of the s i t e s , but i t was more abundant in s h e l t e r e d s i t e s . On the f u l l y s h e l t e r e d shore of D i a n a I s l a n d , I found on l y C. pac i f i cum and no C^ cod i co I a d u r i n g a f u l l year of o b s e r v a t i o n , w h i l e nea rby at S e p p i n g s I s l a n d , both C^ cod i c o I a and C ^ pac i f i c u m occu r r ed as e p i p h y t e s . Both C^ cod i c o l a and C ^ pac i f i cum are f u l l y c o r t i c a t e d and they look ve r y s i m i l a r , but they were r e l i a b l y d i s t i n g u i s h e d on the b a s i s of r h i z o i d mo rpho l o g y . C^ cod i co l a p o s s e s s e s r e l a t i v e l y t h i c k r h i z o i d s w i t h l a r g e expanded t i p s w h i l e r h i z o i d s of C^ pac i f i c u m are narrower and do not Table I I . Epiphytes of Godiim fragile collected in the vic inity of Bamfield, previously reported substrata, months of collection (1 = January, 2 = February, etc.) and wave exposure of collection s i tes . SPECIES (DIVISION*) REPORTED SUBSTRATA15 1 2 3 4 MDNTH 5 6 7 8 9 10 11 12 W\VE EXP.C Cannon epiphytes: Ceramiim codicola J. Ag. (R) C fr ag i 1 e. + + + + + + + + + + + + 1,2,3 C. pacificim (Col l .) Kyi . (R) Rocks, plants. + + + + + + + + + + + + 1,2,3 C. gardneri Ky i . (R) Rocks. + + + + + + + + + 1,2,3 Erythrotrichia pulvinata Gardn. (R) C. fr ag i 1 e. + + + + + + + + + + + 1,2,3 Ulva sp. (C) Rocks, wood, plants. + + + + + + + + + 1,2,3 Navicula grevi l le i (B) + + + + + + + + + 1,2,3 Ectocarpoids (P) Rocks, plants. + + + + + + + 1,2,3 Audouinella rhizoidea C. fragile, (Drew) Garb. (R) other plants. + + + + 2,3 Audouinella sp. + + + + 1,2,3 Plocamiun tenue Kyi. (R) Rocks. + + + 1,3 Rhodynenia sp. (juvenile) (R) Rocks. + + + 2 Cal I i tharmion pikeanun Harv. (R) Rocks. + + + 1,2,3 Ant i tharm ion defectun Ky 1. (R) Plants. + + 1,2 Rare epiphytes: Ulothr ix sp. (C) Chaetcmorpha sp. (C) Polysiphonia hendryi Gardn. (R) Cal I i tharmion acutumKyl. (R) P. pacifica Hoilenb. (R) Gel idiun sp. (juvenile) (R) Porphyra sp. (R) Microcladia boreal is Rupr. (R) M. couIteri Harv. (R) Laminarian (juvenile) (P) Entercmorpha sp. (C) Platytharmion pectinatun Ky I . (R) Ceramitm californictm J. Ag. (R) Halosaccion glandi forme (OneI.) Rupr. (R) Sphacelar ia furcigera Kutz. (P) Colpcmenia sp. (P) CIadophora sp. (C) Rocks, wood, plants. Rocks , tangled in plants. Rocks. Rocks, crabs, plants. Rocks, p i l ings. Rocks. Rocks. Rocks. Plants. Rocks. Rocks, plants. Roc.ks, p i l ings . Graci laria, other plants. Rocks. Plants. Rocks, plants. Rocks, shel ls. + + + + + + + + + + 3 2 2 2 2 2 1 1 3 1 2 a R = Rhodophyta, P = Phaeophyta, C = Chlorophyta, B = Bacillariophyta. bSubstrata reported in Abbott and Hollenberg (1976). °Wave exposure as in Table 1. Table I I I . Epiphytes of Godium setchel I i i collected in the vicinity of Bamfield, previously reported substrata, months of collection (1 = January, 2 = February, etc.) and wave exposure of collection s ites. SPECIES (DIVISION) REPORTED NOMTH WAVE SUBSTRATA 1 2 3 4 5 6 7 8 9 10 11 12 EX 5 . Common epiphytes: Ectocarpus parvus (Saund.) Hollenb. (P) Audouinella sp. (R) Platytharmion vi I losim Ky I . (R) Ceramiun gardneri Kyi . (R) Audouinella rhizoidea (Drew) Garb. (R) Antithamnion defectum Kyi. (R) Ulva sp. (C) Rare epiphytes: Ke I ps. Rocks,.pi I ings. Rocks. C fr ag i Ie and other plants. Plants. Rocks, wood, plants. Polysiphonia pacifica Hollenb. (R) Rocks, p i l ings Herposiphonia vert ic i l lata (Harv.) Ky i . (R) H. pi inula (J. Ag.) Hollenb. (R) Erythrotr ichia carnea (Dillw.) J. Ag. (R) Erythrotr ichia pulvinata Gardn. (R) Pterosiphonia dendroidea (Mont.) FaIk. (R) Gr i f f i ths ia pacifica Ky I . (R) Ectocarpus si I iculosus (Di I Iw.) Lyngb. (P) Plants. Costar ia costata (C. Ag.) Saund. (P) Rocks. Desmarestia I igulata (Lightf.) Lamour. (P) Rocks. Entercmorpha sp. (C) Rocks, mud, plants. Plants. Rocks, plants. Plants. C fr ag i I e. Rocks. Rocks. + + + + + + + + + + + + + + + + + + + + + + 1,2 1,2 2 2 2 1,2 1,2 2 2 2 1 2 2 2 2 2 1 have " expanded t i p s . In a d d i t i o n , C^ pac i f i cum grows to a l a r g e r s i z e (up to 18 cm) than does C. c o d i c o l a (up to 2 . 5 cm) (Abbott and H o l l e n b e r g 1 9 7 6 ) . Most of the r a re e p i p h y t e s were not f e r t i l e , and many p r o b a b l y would not reach r e p r o d u c t i v e m a t u r i t y be fo re e i t h e r be ing d i s l o d g e d from C ^ fr ag i I e or t e a r i n g i t away because of t h e i r l a r ge s i z e . For example, Rhodymen i a s p . , G e I i d i um s p . , P o rphy r a s p . , Ha I o s a c c i on g I and i fo rme, and perhaps P Iocami um tenue , CaI I i thamn ion p i keanum, Mi c roc I ad i a b o r e a l i s and M^ c o u I t e r i p r o b a b l y would not reach r e p r o d u c t i v e m a t u r i t y because of t h e i r l a r ge s i z e . Of the 17 r a r e e p i p h y t e s , 14 were found e x c l u s i v e l y in s h e l t e r e d (2 or 3 ) a r e a s . E i g h t e e n taxa of mac roa l gae were i d e n t i f i e d from the e p i f l o r a of C . s e t c h e l I i i , 7 o f which were commonly found. Of t he se , E c t o ca rpu s pa rvu s and Audou i ne I I a sp . were found in n e a r l y every c o l l e c t i o n . Audoui nel la r h i zo i dea i s r epo r ted as a f requent ep iphy te of C^ fr ag i I e and o c c a s i o n a l on o the r a l gae (Abbot t and H o l l e n b e r g , 1 9 7 6 ) . I t was found almost as f r e q u e n t l y on C ^ se tche I I i i as on C^ f r a g i l e in B a r k l e y Sound. In a d d i t i o n , a sma I I red a l g a i d e n t i f i a b l e as E r y t h r o t r i c h i a p u I v i n a t a , based on i t s c r u s t o s e base w i t h u p r i g h t s a r i s i n g from i t , was found g row ing on the u t r i c l e t i p s of C . se t che I I i i . I t was s i m i l a r in appearance to E ^ pu I v i na ta i d e n t i f i e d from u t r i c l e t i p s of C^ fr ag i I e . Of the 11 t axa r a r e l y found on C^ s e t c h e l I i i , 9 were found e x c l u s i v e l y in m o d e r a t e l y s h e l t e r e d a r e a s . Of t h e s e , Cos tar i a cos t a t a , Desmar es t i a I i g u I a t a , and perhaps P t e r o s i phon i a dendro idea and Gr i f f i ths i a pac i f i ca would p r o b a b l y not reach r e p r o d u c t i v e m a t u r i t y on s e t che l I i i be fo re be ing d i s I o d g e d by waves . E p i p h y t e s found on C^ f r a g i l e c o l l e c t e d f r o m H o r n b y I s l a n d are l i s t e d in Tab l e I V . P I a t y t hamn ion v i I Iosum, the most abundant ep iphy te he re , was never found in B a r k l e y Sound on C^ f r a g i l e , a l t h o u g h i t was p re sen t as an ep iphy te Tab le I V . E p i p h y t e s of Cod i um f r a g i I e c o l l e c t e d from Hornby I s l a n d , l i s t e d from most to l e a s t abundant . P I a t y t h a m n i o n v i l l o s u m K y l . (R) Audou i ne I la rh i zo i dea (Drew) G a r b . (R) E c t o c a r p o i d s (P) Nav i c u I a g rev i I Ie i (B) Cer ami um ga rdne r i K y I . (R) C . p a c i f i c u m ( C o l l . ) K y i . (R ) T a b l e V . E p i p h y t e s of e p i p h y t i c Ceramium cod i c o I a and of c o l l e c t i o n and wave exposure of c o l l e c t i o n s i t e s . p a c i f i c u m , months MONTH S P E C I E S ( D I V I S I O N ) 1 2 3 4 5 6 7 8 9 10 11 12 WAVE EXP. E r y t h r o c I ad i a i r r e g u l a r i s f . sub i n t e g r a ( R o s e n v . ) G a r b . , Hans . & S c a g . (R) E r y t h ro t r i ch i a ca rnea ( D i I I w . ) J . Ag . (R) U l va s p . (C) sp. Audou i ne I I a P locamium tenue (R) Ky i P o r p h y r a sp. Ha Io sacc i on (R) , (R) g I and i forme ( G m e l T P R u p r . (R) M i c r o c I ad ia cou I ter i H a r v . Ca I I i thamn ion pi keanum H a r v . (R) Ant i thamn ion de fec tum K y I . P o I y s i phon i a hend ry i Gardn, Pt e ros i phon i a b i p i nnata ( P o s t , et R u p r . ) F a l k . Ca I I i thamn i on sp . (or P I e o n o s p o r i um s p . ) Enteromorpha s p . (C) Nav i c u I a grev i I Ie i (B) + + + (R) + (R) + . (R ) (R) (R) + + + + + + + + + + + + + + + 1 , 2 , 3 1 , 2 , 3 1 , 2 , 3 1 , 2 , 3 1 2 , 3 3 2 1,2 1 2 2 3 2 on C^ se t che I I i i . The other e p i p h y t e s l i s t e d in Tab le IV were a l s o p re sent on C^ fr ag i I e in B a r k l e y Sound. C ^ cod i c o I a i s n o t a b l y absent from Hornby I s l a n d , a l t h o u g h C^ pac i f i cum and C . g a r d n e r i are p r e s e n t . E p i p h y t i c mac roa l gae g row ing on C^ cod i c o I a and C ^ pac i f i cum which themse l ve s were e p i p h y t i c on C^ f r a g i l e are l i s t e d in Tab le V . Of the 15 s p e c i e s l i s t e d he re , 11 were a l s o found d i r e c t l y on C ^ fr ag i I e . E r y t h r o c I ad ia i r r e g u l a r i s f . s u b i n t e g r a and E r y t h r o t r i c h i a ca rnea were a lmost a lways found on both of these Ceramium s p p . , but they were not found d i r e c t l y on C ^ fr ag i I e . The other 2 e p i p h y t e s found on Cerami um spp. but not d i r e c t l y on fr ag i I e were o n l y ob se r ved once or tw i ce . Ul va sp . was common on Ceramium spp . and most smal I tha i I i of U I v a were on Ceramium spp . r a the r than on C^ f r a g i I e . As U I v a grows l a r g e r , i t may ove rg row the Ceramium on which i t is g rowing and a t t a c h d i r e c t l y to C ^ fr ag i I e . No Ce r am i um sp . was found as an ep iphy te of Ce r am i um on C^ fr ag i I e even though they are the most abundant e p i p h y t e s . B. C u l t u r e experiments Ceramium cod i c o I a w i l l grow to r e p r o d u c t i v e m a t u r i t y in l a b o r a t o r y c u l t u r e apar t from C . f r a g i l e . The p a t t e r n of g e r m i n a t i o n and development in c a r p o s p o r e s and t e t r a s p o r e s i s s i m i l a r , f o l l o w i n g a b i p o l a r "Ce ramium- type " p a t t e r n wh ich is t y p i c a l of red a l gae of the order Ce r am ia l e s ( D i x on 1 9 7 3 ) . The sequence for spo re s of C. c o d i c o l a shown in F i g u r e s 3 a - d t y p i c a l l y o c cu r s in 4 to 8 d a y s . Spo re s d i v i d e u n e q u a l l y ( F i g u r e 3b) and the sma l l e r daughter c e l l p roduces a r h i z o i d which e l o n g a t e s ( F i g u r e 3 c ) . When a p i c a l d i v i s i o n of the l a r ge r daughter c e l l o c cu r s the r h i z o i d t i p was ob se rved to be expanded i n to a bu lb ( F i g u r e 3 d ) . The r h i z o i d t i p was never obse rved to be expanded be fo re a p i c a l d i v i s i o n o c c u r r e d . As growth c o n t i n u e s in c u l t u r e , narrow F i g u r e 3 . G e r m i n a t i o n p a t t e r n of spo re s of Ceramium cod i c o I a in c u l t u r e . a. Spore be fo re d i v i s i o n . S c a l e = 25 pm. b . U n e q u a l l y d i v i d e d s p o r e . S c a l e = 25 jim. c . S p o r e l i n g showing d e v e l o p i n g r h i z o i d . S c a l e = 25 jjm. d. S p o r e l i n g showing a p i c a l development and bu lbous r h i z o i d . S c a l e = 100 jjm. e . S p o r e l i n g showing narrow r h i z o i d s g rowing out of swo l l en r h i z o i d . S c a l e = 100 jjm. r h i z o i d s l a c k i n g bu lbous t i p s a r i s e from the bu lbous s w e l l i n g ( F i g u r e 3 e ) . A t o t a l of 42 c u l t u r e s i n i t i a t e d from c a r p o s p o r e s and 25 from t e t r a s p o r e s , w i t h 2 to 10 s p o r e l i n g s per c u l t u r e , c o n t a i n e d p l a n t s which grew and s u r v i v e d at l ea s t 30 days a f t e r spore r e l e a s e . S p o r e l i n g s in c u l t u r e that d i d not s u r v i v e t h i s long were e i t h e r overgrown w i t h con taminan t s or d i e d of unknown c a u s e s . C o n t a m i n a t i o n by E r y t h r o c I ad i a i r r e gu I a r i s f . sub i n t e g r a , E r y t h r o t r i c h i a c a r n e a , e c t o c a r p o i d s and u n i c e l l u l a r a lgae was e s p e c i a l l y common in c u l t u r e s i n i t i a t e d from t e t r a s p o r e s . The number of s u c c e s s f u l c u l t u r e s that c o n t a i n e d p l a n t s that produced r e p r o d u c t i v e s t r u c t u r e s at v a r i o u s tempera tu re s are l i s t e d in Tab le V I . F e r t i l e c a r p o s p o r e I i n g s produced t e t r a s p o r a n g i a , and f e r t i l e t e t r a s p o r e l i n g s produced e i t h e r c a r p o g o n i a or s pe rmat i a on s epa ra te p l a n t s . No c y s t o c a r p s were o b t a i n e d in c u l t u r e . T e t r a s p o r e s produced in c u l t u r e were i s o l a t e d , but these d i ed p r i o r to or a f t e r g e r m i n a t i o n . In one t r i a l at 12°C, 51 t e t r a s p o r e s were i s o l a t e d , 37 o f which d ied w i thou t g e r m i n a t i n g and 14 o f wh i ch ge rm ina ted , p r o d u c i n g a r h i z o i d be fo re d y i n g . At 10 and 12°C, the 39 c a r p o s p o r e I i ngs that produced t e t r a s p o r a n g ia were at l e a s t 1 . 7 rrm in l e n g t h ; two c a r p o s p o r e I i n g s which reached t h i s l eng th d i d not produce t e t r a s p o r a n g i a . The two c a r p o s p o r e I i n g s that produced t e t r a s p o r a n g i a at 15°C were l e s s than 1 .7 mm in l e n g t h . At 5°C, no c y s t o c a r p s of C^ cod i c o l a r e l e a s e d c a r p o s p o r e s . T e t r a s p o r e s were r e l e a s e d , but these o f t e n remained grouped as t e t r a d s . They were m a i n t a i n e d at 5°C for 17 days u n t i l they a l l l o s t t h e i r p i g m e n t a t i o n ; none of them germinated in t h i s t ime. At 10, 15 and 20°C, both c a r p o s p o r e s and t e t r a s p o r e s ge rmina ted and grew. The pe rcen tage of c a r p o s p o r e s and t e t r a s p o r e s that ge rmina ted and s u b s e q u e n t l y s u r v i v e d at these temperatures a re p l o t t e d a g a i n s t time in F i g u r e 4 . C a r p o s p o r e s grew best at 10°C, Tab l e V I . F e r t i l i t y of c u l t u r e s of Ceramium c o d i c o l a i n i t i a t e d from c a r p o s p o r e s or t e t r a s p o r e s at v a r i o u s t empe ra t u r e s . S u c c e s s f u l c u l t u r e s were those that c o n t a i n e d s p o r e l i n g s that s u r v i v e d at l e a s t 30 d a y s . Length of f e r t i l e p l a n t s was measured when they f i r s t became f e r t i l e . The time u n t i l f e r t i l i t y i s the s h o r t e s t for c u l t u r e s at that t empera tu re . Temp. (°C) # s u c c e s s f u l # c u l t u r e s # p l a n t s Length (cm) of Days u n t i l c u l t u r e s fer t i I e f e r t i l e f e r t i l e p l a n t s f e r t i l e C a r p o s p o r e I i ngs : 10 20 12 12 2 1 15 17 2 18 2 0 20 1 0 36 1 . 7 - 3 . 8 37 3 1 . 7 - 2 . 3 38 2 0 . 4 - 1 . 6 37 T e t r a s p o r e I i n g s : 10 13 6 41 12 2 1 5 8 a 15 9 6 30 18 1 0 20 0 0 a T h i s c u l t u r e was p r o b a b l y f e r t i l e be fo re t h i s t ime. F i g u r e 4 . Pe r cen tage of g e r m i n a t i o n and s u r v i v a l of t e t r a s p o r e s and c a r p o s p o r e s of Cerami um cod i c o I a at 10, 15 and 20°C w i t h t ime. E r r o r ba r s a re 95% c o n f i d e n c e i n t e r v a l s for pe r cen tage s from R o h l f and Soka l ( 1 9 8 1 ) . DAYS f o l l o w e d by 15 and 20°C. At 20°C, no c a r p o s p o r e I i n g s s u r v i v e d past 31 d a y s , and the l a r g e s t s i z e ob t a i ned was 0 . 2 0 mm. A lower pe rcen tage of t e t r a s p o r e s ge rm ina ted than c a r p o s p o r e s . G e r m i n a t i o n and s u r v i v a l of t e t r a s p o r e s was not s i g n i f i c a n t l y d i f f e r e n t at 10 and 15°C, but was s i g n i f i c a n t l y l e s s at 20°C. Growth in l eng th of s p o r e l i n g s at 10 and 15°C i s p l o t t e d a g a i n s t time in F i g u r e 5 . I n i t i a l l e ng th s were measured when spore l i n g s we re > 0 . 10 mm. Growth of t e t r a s p o r e I i n g s and c a r p o s p o r e I i n g s ( second t r i a l ) was f a s t e r at 10 than at 15°C. Growth of car pos por e I i ngs ( f i r s t t r i a l ) was equal at 10 and 15°C, and l e s s than growth of car pos por e I i ng s in the second t r i a l at both t e m p e r a t u r e s . Many t e t r a s p o r e I i n g s and c a r p o s p o r e ! i n g s ( second t r i a l ) grown at 10°C were p r o d u c i n g r e p r o d u c t i v e s t r u c t u r e s at 40 to 50 d a y s , which may account for the reduced growth at t h i s t ime. C a r p o s p o r e s and t e t r a s p o r e s were grown for on l y 14 days in e x t r a c t s of Cod i um because of b a c t e r i a l c o n t a m i n a t i o n . G e r m i n a t i o n and growth of spo re s o b t a i n e d in t h i s time are l i s t e d in Tab l e V I I . Lower c o n c e n t r a t i o n s ( 1 . 2 5 to 12 . 5% ) of e x t r a c t had no e f f e c t on spore g e r m i n a t i o n pe rcen tage compared to g rowth in PES w i t hou t e x t r a c t . H i ghe r c o n c e n t r a t i o n s ( 2 5 . 0 to 100 .0% ) of C. fr ag i I e e x t r a c t i n h i b i t e d g e r m i n a t i o n ; h i g h e r c o n c e n t r a t i o n s of C. s e t c h e I I i i e x t r a c t had l e s s e f e c t . Growth of s p o r e l i n g s was u n a f f e c t e d in lower c o n c e n t r a t i o n s and i n h i b i t e d in h i ghe r c o n c e n t r a t i o n s of e x t r a c t s of bo th s p e c i e s of Cod i um. Fewer spo re s of C^ cod i co I a ge rminated and grew in c u l t u r e on l i v i n g C. fr ag i I e and C ^ se tche I I i i than in c u l t u r e w i t hou t Cod i um. Reduced s u r v i v a l may be due to the c o n s t a n t p re sence of con tam inan t s in these c u l t u r e s , or to i n h i b i t i o n by Cod i um as a r e s u l t of c o m p e t i t i o n for n u t r i e n t s or a l l e l o p a t h y . I t was d i f f i c u l t to o b t a i n accu ra te count s of spo re s on the s u r f a c e s of Cod i um tha i I i . On C. f r a g i l e , 7 . 7 % of c a r p o s p o r e s and 2 . 8 % of F i g u r e 5. Mean l eng th s of t e t r a s p o r e I i ng s and c a r p o s p o r e I i n g s of Ce r am i um cod i c o I a grown in c u l t u r e at 10 and 15°C at v a r i o u s t imes of g rowth. E r r o r ba r s are + 1 s t a n d a r d e r r o r . Tetraspores I.OH 0.5- • 10° c O I5°C 0 -Carpospores - Trial E E x h-e> z LU 0.5-0 -Carpospores — Trial 2 2.OH 0.5H 20 3 0 4 0 DAYS 50 60 70 Tab le V I I . I n i t i a l number of s p o r e s , per cent g e r m i n a t i o n and maximum l eng th s of Ceramium c o d i c o l a t e t r a s p o r e I ings and c a r p o s p o r e I ings a f t e r 14 days of growth in e x t r a c t s of Cod i um fr ag i I e and setche I I i i . M i s s i n g l eng th s i n d i c a t e that no s p o r e l i n g s grew to measu rab le l eng th ( 0 . 1 0 mm). E x t r a c t c o n c e n t r a t i o n (%) 0 1 . 25 2 . 5 0 1 2 . 5 2 5 . 0 50 .0 75 .0 1 00 . 0 C. f r a g i l e T e t r a s p o r e I i ngs : # s p o r e s 40 % g e r m i n a t i o n 35 .0 Max . l e n g t h (mm) 0 . 4 0 Ca rpo spo r eI i n g s : # s p o r e s 25 % germ. 88 .0 Max. l e n g t h (mm) 0 . 8 0 C. setche 11 i i T e t r a s p o r e I i n g s : # s p o r e s 40 % ge rm ina t ion 35 .0 Max. l e n g t h (mm) 0 . 4 0 Car pospor e I i n g s : # s po r e s 25 % germi nat i on 88 .0 Max. l e ng th (mm) 0 . 8 0 27 29 24 44 . 4 20 . 7 4 2 . 0 0 . 4 5 0 . 6 5 65 51 94 33 .8 43 .1 4 4 . 7 0 . 6 0 0 . 2 5 90 87 83 22 .2 4 2 . 5 13 .3 0 . 4 0 0 . 65 0 . 3 0 120 67 98 4 9 . 2 55 .2 6 7 . 3 0 . 90 0 . 3 0 0 . 90 25 22 27 24 20 .0 4 . 5 0 . 0 4 . 2 0 . 4 0 41 47 24 20 .0 8 .5 4 . 2 0.20 90 62 48 2 3 . 3 2 5 . 8 35 .4 0 . 55 0 . 2 0 0 . 2 0 90 90 52 14 .4 2 2 . 2 28 . 8 0 . 2 0 0 . 3 0 t e t r a s p o r e s counted grew and s u r v i v e d to 30 d a y s , w h i l e on C^ se t che I I i i , 0 . 6 and 0 . 2 % s u r v i v e d r e s p e c t i v e l y . S u r v i v a l of s po re s was s i m i l a r on C^ fr ag i I e f rom b l e a c h - t r e a t e d f i e l d - c o l l e c t e d specimens and c u l t u r e d spec imens . Growth of s p o r e l i n g s on thai I i of Cod i um was s lower than in c u l t u r e s w i t h o u t Cod i um p r e s e n t . On C^ f r a g i l e at 10°C, c a r p o s p o r e l i n g s grew to a maximum length of 0 . 6 4 mm in 60 days in one t r i a l , 0 . 6 4 mm in 38 days in a second and 0 . 4 9 mm in 38 days in a t h i r d . T e t r a s p o r e I i n g s grew to 0 . 5 5 mm in 61 d a y s , 0 . 6 0 mm in 59 days and 0 . 6 2 mm in 59 d a y s . On C^ s e t che l I i i , c a rpo s por e I i ng s grew to a maximum of 0 . 3 5 mm in 61 days and no t e t r a s p o r e I i n g s grew l a r g e r than 0 . 1 5 mm. No r e p r o d u c t i v e s t r u c t u r e s were produced on these p l a n t s . R h i z o i d s of C^ cod i co I a grown on C^ fr ag i I e in c u l t u r e t y p i c a l l y p e n e t r a t e d the tha i I us and produced bu lbous t i p s . The t i p s remained bu lbous in 40 o f 46 s p o r e l i n g s , u n l i k e those grown w i t hou t C^ f r a g i l e . The bu lbous t i p s of r h i z o i d s from these 40 s p o r e l i n g s had a mean d iameter of 6 0 . 4 pm ( S . D . = 1 4 . 1 1 ) , wh ich is not s i g n i f i c a n t l y d i f f e r e n t from bu lbous t i p s from r h i z o i d s of f i e l d - c o l l e c t e d specimens g rowing on C^ fr ag i I e ( see be l ow ) . R h i z o i d s of C. c o d i c o l a s p o r e l i n g s grown on C^ s e t c h e I I i i o f t e n d id not p e n e t r a t e the t h a l l u s and and grew over the s u r f a c e . When p e n e t r a t i o n was a c h i e v e d , the r h i z o i d t i p s were s l i g h t l y s w o l l e n , w i t h a mean diameter of 33 .1 pm ( S . D . = 1 4 . 2 8 , n = 8 ) , which is s i g n i f i c a n t l y (p < . 0 0 1 ) sma l l e r than r h i z o i d bu lb s of C^ cod i c o I a g rowing on C^ fr ag i I e . S p o r e l i n g s of Ce r am i um pac i f i c um and C ^ gardner i grew f a s t e r and l a rge r than C ^ cod i c o I a in c u l t u r e . C a r p o s p o r e l i n g s of C^ pac i f i c um ranged from 2 . 8 to 8 . 4 mm in l eng th 37 days a f t e r spo re s were r e l e a s e d from f i e l d - c o l l e c t e d spec imens at 12°C, and w i t h i n 105 days they had grown into a t ang l ed clump of p l a n t s wh ich produced t e t r a s p o r e s . T e t r a s p o r e s were i s o l a t e d and they ge rm ina ted and grew. Growth was s l o w e r , w i t h s p o r e l i n g s r e a c h i n g 10 mm in 71 d a y s . C a r p o g o n i a were produced w i t h i n 124 d a y s , but no c y s t o c a r p s were o b t a i n e d and the c u l t u r e s were te rm ina ted because of c o n t a m i n a t i o n . T e t r a s p o r e s r e l e a s e d from f i e l d - c o l l e c t e d specimens grew and produced c a r p o g o n i a and spe rmatang i a w i t h i n 45 days and c y s t o c a r p s w i t h i n another 16 days at 12°C. C a r p o s p o r e s were not s u c c e s s f u l l y i s o l a t e d because of c o n t a m i n a n t s . Male gametophytes were sma l l e r than female gametophytes in c u l t u r e . Of 24 t e t r a s p o r e I i n g s , 16 p roduced c a r p o g o n i a and 8 s p e r m a t a n g i a . S p o r e l i n g s of C. pac i f i cum a l s o grew we l l and produced r e p r o d u c t i v e s t r u c t u r e s at 18°C. F i e l d - c o l l e c t e d specimens of C^ pac i f i c um we re f u l l y c o r t i c a t e d , w i t h no gaps in the i n t e rnode s ( F i g u r e 6 a ) . S p o r e l i n g s of C^ pac i f i cum in l a b o r a t o r y c u l t u r e grew w i t h l a r ge i n t e r n o d a l gaps in the c o r t i c a t i o n ( F i g u r e 6 b ) . C o r t i c a t i o n was more deve loped d i s t a l from the ap i ce s ( F i g u r e 6 c ) . C a r p o s p o r e I i n g s of C^ gardner i grew into a clump of t ang l ed p l a n t s that were p r o d u c i n g t e t r a s p o r a n g ia w i t h i n 49 days of growth at 12 and 18°C. No t e t r a s p o r e s were s u b s e q u e n t l y i s o l a t e d . T e t r a s p o r e I i n g s grew to 2 . 5 - 2 . 7 mn in l e ng th in 17 days and were p r o d u c i n g c a r p o g o n i a or spe rmatang i a on s e p a r a t e p l a n t s by 53 days at 12 and 18°C. No c y s t o c a r p s were ob ta i ned in c u l t u r e . C . R h i z o i d morphology The d i amete r s of r h i z o i d s of f i v e s p e c i e s of Ce r am i um and M i c r o c lad ia c o u I t e r i under v a r i o u s c o n d i t i o n s are l i s t e d in Tab Ie V I I I . On I y C^ cod i c o I a p o s s e s s e s r h i z o i d s w i t h bu lbous t i p s ; spec imens c o l l e c t e d from C. f r a g i l e in the f i e l d had r h i z o i d s wh ich ave raged 6 8 . 3 jjm in d iameter ( S . D . = 1 2 . 8 0 , n = 2 0 ) . R h i z o i d s of C^ cod i co I a in C^ fr ag i I e were about 1 . 1 mm long , F i g u r e 6 . C o r t i c a t i o n of Ce r am i um pac i f i cum from the f i e l d and in c u l t u r e . a. Specimen c o l l e c t e d from the f i e l d showing complete c o r t i c a t i o n . b . Specimen grown from a c a r p o s p o r e in c u l t u r e showing naked i n t e r n o d e s . c . Same as b ) , showing d e v e l o p i n g c o r t i c a t i o n f a r t h e r from the apex. a b c Tab l e V I I I . D i amete r s of r h i z o i d s (jjm) of Ce r am i um spp. and Mi c roc I ad i a cou I ter i from f i e l d - c o l l e c t e d and c u l t u r e d ( f rom spo re s or r e g e n e r a t e d from a p i c a l segments) s p e c i m e n s . Mean + s t a n d a r d d e v i a t i o n (sample s i z e ) . S p e c i e s Na tu r a l C u l t u r e d c , cod i co1 a 2 4 . 6 + 5 .10 (19) 2 4 . 3 20 .0 I + I + 3 . 5 5 6 .53 (20) (16) ( S p o r e s ) (Regen . ) c ^ pac i f i cum: On Cod i um f r a g i l e On P r i o n i t i s l a n c e o l a t a 1 2 . 2 12.7 + + 1 .14 1.17 (10 ) (20 ) 9 . 9 : + 1 .20 (10 ) ( S p o r e s ) ga rdner i : On Cod i um f r a g i1e On C . s e t c h e 1 1 i i 2 7 . 1 27 .0 + + 6 .77 7.57 (10) (10 ) 16 .6 + 3 .38 (20 ) (Regen. ) ea ton ianum 1 7 . 4 + 4 .01 ( 10 ) 19 . 0 + 2 .58 (10 ) (Regen. ) c a1 i f o r n icum 14.3 + 1.57 (10) 14.5 + 3.37 (10) (Regen . ) ivu cou1te r i 1 2 . 5 + 1.82 ( 28 ) ( S p o r e s ) a p p r o x i m a t e l y the same l eng th as the u t r i c l e s . R h i z o i d s of C. c o d i c o l a and C. ga rdner i were p igmented, w h i l e the o t he r s had c o l o r l e s s r h i z o i d s . C. pac i f i cum, C. gardner i and C ^ cal i f o r n icum were ob se rved to produce d i g i t a t e h o l d f a s t s on the t i p s of r h i z o i d s as they con tac ted the s u b s t r a t u m in c u l t u r e . T h i s was ob se r ved once in C^ pac i f i c um and gardner i and twice in C^ cal i f o r n icum. T h i s s t r u c t u r e is drawn in F i g u r e 7 a . No d i g i t a t e h o l d f a s t s t r u c t u r e s were ob se rved in C^ cod i c o I a or C^ eaton i anum. R h i z o i d s o f spo re l i n g s of M^ cou I ter i were a l s o a r r anged in a p a t t e r n that cou ld serve as a h o l d f a s t s t r u c t u r e ( F i g u r e 7 b ) . The r h i z o i d s of a l l Ce r am i um spp. except C^ cod i co I a a ro se from the ba sa l and e rect p o r t i o n s of the p l a n t s in nature and in c u l t u r e . R h i z o i d s a r i s e from the nodes between a x i a l c e l l s . F i g u r e 7c shows such r h i z o i d s on C. pac i f i cum from a specimen e p i p h y t i c on C ^ fr ag i I e . R h i z o i d s of C. cod i co I a and cou I ter i a ro se o n l y from the basa l p o r t i o n of the p l an t both in na tu re and in l a b o r a t o r y c u l t u r e . D. P h e n o l o g i c a l phenomena Ma le and female gametophytes and t e t r a s p o r o p h y t e s of C. cod i c o I a and C. pac i f i c um were p re sen t in a l l months on C^ f r a g i l e . In a d d i t i o n , a l l r e p r o d u c t i v e s t age s of C^ gardner i were found in J a n u a r y , March , June, J u l y and November, as we l l as t e t r a s p o r o p h y t e s and female gametophytes in A p r i l and t e t r a s p o r o p h y t e s in F e b r u a r y , May and September , g rowing on e i t h e r C. fr ag i I e or C ^ se tche I I i i . Over a l l months and both s i t e s (n = 4400 C ^ cod i c o I a p l a n t s ) , t e t r a s p o r o p h y t e s made up 3 8 . 2 % of the sample, female gametophytes 3 2 . 8% , male gametophytes 1 7 . 5 % and v e g e t a t i v e p l a n t s 1 1 . 5% . These f r e q u e n c i e s are a l l s i g n i f i c a n t l y d i f f e r e n t from one a n o t h e r , and these d i f f e r e n c e s o b t a i n F i g u r e 7 . R h i z o i d mo rpho l o g y . a . D i a g r a m of d i g i t a t e h o l d f a s t s t r u c t u r e produced on the t i p s of r h i z o i d s of some Cerami um spp. in c u l t u r e . b . R h i z o i d s of Mi c roc I ad i a cou 11e r i from a s p o r e l i n g grown in c u l t u r e . c . R h i z o i d s of a specimen of Ceramium p a c i f i c u m c o l l e c t e d as an ep i phy te of Cod i um f r a g i l e w i t h r h i z o i d s a r i s i n g away from the base of the p l a n t . over most months of the y e a r . O v e r a l l , 5 0 . 3% of the sampled p l a n t s were f e r t i l e gametophytes , wh ich i s s i g n i f i c a n t l y more than f e r t i l e t e t r a s p o r o p h y t e s ( 3 8 . 2 % ) . V a l u e s of the F - s t a t i s t i c ob ta i ned from two-way anovas on f requency of r e p r o d u c t i v e p l a n t s w i t h s i t e and month as t rea tment s are l i s t e d in Tab le I X . They revea l that s i g n i f i c a n t v a r i a t i o n in f r e q u e n c i e s of t e t r a s p o r o p h y t e and v e g e t a t i v e p l a n t s o c cu r s among months but no s i g n i f i c a n t d i f f e r e n c e s in f r e q u e n c i e s of female and male gametophyte s . There is no s t a t i s t i c a l t d i f f e r e n c e in f r e q u e n c i e s between s i t e s . No s i g n i f i c a n t d i f f e r e n c e s in f r e q u e n c i e s were r e v e a l e d in three-way anovas w i t h month, s i t e and he i gh t as t r ea tment s over the months of J u l y t h rough O c t o b e r . F i g u r e 8 show the mean f r e q u e n c i e s of t e t r a s p o r o p h y t e s , female gametophytes , male gametophytes and v e g e t a t i v e p l a n t s over both s i t e s w i t h an e r r o r bar of minimum s i g n i f i c a n t d i f f e r e n c e for each r e p r o d u c t i v e group as c a l c u l a t e d by the T-method d e s c r i b e d in Soka l and R o h l f ( 1 9 8 1 ) . Two means are s i g n i f i c a n t l y d i f f e r e n t i f the e r r o r bars do not o v e r l a p w i t h an exper imentwi se p r o b a b i l i t y of e r r o r of 0 . 0 5 . These data i n d i c a t e that the h i g h l y s i g n i f i c a n t e f f e c t ob ta i ned for month of the year t reatment on f r e q u e n c i e s of t e t r a s p o r o p h y t e s and v e g e t a t i v e p l a n t s i s the r e s u l t of a dec rea sed f requency of t e t r a s p o r o p h y t e s and an i n c r e a s e d f requency of v e g e t a t i v e p l a n t s in J a n u a r y . There is a s l i g h t tendency for female gametophytes to be l e s s f requent in w i n te r months , a l t h o u g h t h i s was not s i g n i f i c a n t . C o r r e l a t i o n a n a l y s e s were a l s o performed between the v a r i o u s r e p r o d u c t i v e p l an t f r e q u e n c i e s and v e g e t a t i v e p l an t f r e q u e n c i e s . T e t r a s p o r i c and v e g e t a t i v e p l an t f r e q u e n c i e s are s t r o n g l y n e g a t i v e l y c o r r e l a t e d (r = - 0 . 7 2 3 7 , n = 88) w i t h a s lope f i t by l e a s t squa re s of - 0 . 9 6 . Female gametophytes and v e g e t a t i v e p l an t f r e q u e n c i e s a l s o showed a s i g n i f i c a n t n e g a t i v e c o r r e l a t i o n (r = - 0 . 4 4 5 1 , n = 88) w i t h a s lope of - 0 . 7 0 , Tab le I X . F - v a l u e s and a s s o c i a t e d p r o b a b i l i t i e s of s i g n i f i c a n c e ob ta i ned from two-way anovas w i t h t reatments of month (M), s i t e (S) and m o n t h - s i t e i n t e r a c t i o n (M*S ) on f r e q u e n c i e s of r e p r o d u c t i v e c l a s s e s , df = degrees of f r eedom. df T e t r a - Female Male V e g e t a t i v e s po r ophy t e s game tophy te s gametophytes p I ants M 9 6.97 < .001 9 .69 < .001 S 1 0 .005 > .75 0 . 7 5 9 < .50 M*S 9 2 .14 < . 05 4 . 82 < .001 Er ro r 60 M 10 3 .45 < .005 1 .45 < .25 0 . 874 < .75 6 .64 < .001 S 1 0 .054 > .75 1 .89 < . 25 2 .29 < .25 0 . 011 > . 75 M * S 10 1 .67 < .25 1 .21 <.50 0 . 892 < .75 1 .37 < .25 E r r o r 66 Female gametophy te s : C y s t o c a r p i c N o n - c y s t o c a r p i c females females F i g u r e 8 . Mean f r e q u e n c i e s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s o f Ceramium c o d i c o l a e xp r e s s ed as a pe rcentage of samples c o l l e c t e d at F i r s t Beach and S e p p i n g s I s l a n d in the months of the year 1981 (J = J anua r y , F = F e b r u a r y , e t c . ) . Each p o i n t r e p r e s e n t s the mean f requency from e i gh t samples of 50 t ha i I i of C. c o d i c o l a . E r r o r ba r s are minimum s i g n i f i c a n t d i f f e r e n c e (MSD) as c a l c u l a t e d by the T-method ( Soka l and R o h l f 1 9 8 1 ) . 4 C H 20-\ 4 C H 2CH LU o 30-LU CL •OH 4CH 2CH 0 Tetrasporophyte + 4 Female Gametophytes Male Gametophytes f Vegetative Plants iiim J F "m" -i 1 1 r-A M J J -1— A S 0 - r -N -T-D MONTHS which i s s i g n i f i c a n t l y l e s s than the s lope for t e t r a s p o r i c p l a n t s . Male gametophyte and v e g e t a t i v e p l a n t f r e q u e n c i e s were not s i g n i f i c a n t l y c o r r e l a t e d . T h i s i n d i c a t e s that v e g e t a t i v e p l a n t s are m a i n l y s t e r i l e t e t r a spo rophy te s - , some are s t e r i l e female gametophy te s T and few or none are s t e r i l e male gametophytes in the sample t aken . Female gametophytes were d i f f e r e n t i a t e d into those w i t h c y s t o c a r p s and those w i thou t c y s t o c a r p s (but w i t h c a r p o g o n i a ) . Two-way anova on month and s i t e r e v e a l s that a s i t e - m o n t h i n t e r a c t i o n s i g n i f i c a n t l y a f f e c t s both c y s t o c a r p i c and non-cys toca r p i c p l an t f r e q u e n c i e s . However.,. C. c o d i c o l a was sampled at d i f f e r e n t t imes of the month from each s i t e . Mean f r e q u e n c i e s are shown in F i g u r e 9 T w i t h s i t e s p l o t t e d s e p a r a t e l y . The two cu rve s for the s i t e s are l a r g e l y p a r a l l e l ; no s i t e - m o n t h i n t e r a c t i o n is apparent from these d a t a . At both s i t e s T the genera l t rend is that more female gametophytes have c y s t o c a r p s from late A p r i l or e a r l y May u n t i l Oc tober or November at both s i t e s . The minimum c y s t o c a r p i c p l a n t f r e q u e n c i e s were ob t a i ned at m i d - J a n u a r y at S e p p i n g s I s l a n d and e a r l y March at F i r s t Beach. The s i g n i f i c a n t s i t e - m o n t h i n t e r a c t i o n ob ta i ned in the anova p r obab l y r e f l e c t s these d i f f e r e n c e s p lu s a r t i f a c t u a l d i f f e r e n c e s between s i t e s in sampl ing t imes as noted above. C o r r e l a t i o n of c y s t o c a r p i c and n o n - c y s t o c a r p i c female gametophyte f r e q u e n c i e s r e vea l ed a s i g n i f i c a n t n e g a t i v e c o r r e l a t i o n ( r = - 0 .69 , . n = 86) w i t h a l e a s t squa re s l i n e of Y = - 1 . 5X + 35 . 8 (Y = c y s t o c a r p i c , . X = non - c y s toca r p i c) wh i ch i s s i g n i f i c a n t l y d i f f e r e n t from a s lope of - 1 . 0 . T h i s i n d i c a t e s that i f a l l female gametophytes bear c y s t o c a r p s T 3 5 . 8% of the p o p u l a t i o n would be female gametophytes,. w h i l e i f no female gametophytes bear c y s t o ca rp s , . 2 8 . 5% ( X - i n te r cep t ) wou ld be found. The r ema in i n g female gametophytes would p r o b a b l y be s t e r i l e and counted as vege ta t i ve p l a n t s . F i g u r e 9 . Mean f r e q u e n c i e s of a) n o n - c y s t o c a r p i c and b) c y s t o c a r p i c female gametophytes o f Ceramium c o d i c o l a at F i r s t Beach and Sepp i ng s I s l a n d , e x p r e s s e d as percentage of samples c o l l e c t e d in the months of the year 1981 . Each po i n t r e p r e s e n t s the mean f r e q u e n c i e s f rom 4 samples of 50 t ha i I i of C . cod i co I a . E r r o r bar equa l s MSD as c a l c u l a t e d by the T-method (Soka l and R o h l f 1 9 8 1 ) . A n a l y s i s of the l eng th s of 4000 t h a l l i of C^ cod i c o I a of d i f f e r e n t r e p r o d u c t i v e groups from the two s i t e s over 10 months and between two t i d a l h e i g h t s at both s i t e s over four months r e vea l ed important d i f f e r e n c e s due to a l l of the f a c t o r s . F - v a l u e s and a s s o c i a t e d p r o b a b i l i t i e s ob ta i ned from two-way anovas on month and s i t e from March to December and two-way anovas on s i t e and t i d a l he i gh t from J u l y to October are l i s t e d in Tab le X . Means and e r r o r bar s (minimum s i g n i f i c a n t d i f f e r e n c e ) of these l eng th s are p l o t t e d for each c e l l of the anovas in F i g u r e s 1 0 - 1 1 . A l l four c l a s s e s show a s i m i l a r sea sona l t r end , w i t h maximum leng th s o c c u r r i n g in the months of A p r i l t h rough June , and sma l l e r l eng th s from J u l y to December. W i t h i n each c l a s s of r e p r o d u c t i v e p l a n t , p l a n t s are s i g n i f i c a n t l y longer at S e p p i n g s I s l a n d than they are at F i r s t Beach. T h i s can be seen in the g raphs ( F i g u r e 1 0 ) , a l t h o u g h at most i n d i v i d u a l m o n t h - s i t e g r o u p s , the d i f f e r e n c e s are n o n s i g n i f i c a n t . V e g e t a t i v e p l a n t s are longer at S e p p i n g s I s l a n d from March to J u l y , and are a p p r o x i m a t e l y the same l eng th at both s i t e s from Augu s t to December. W i t h i n r e p r o d u c t i v e c l a s s e s d u r i n g J u l y to O c t o b e r , p l a n t s of C. cod i c o I a tended to be longer in the upper r e g i o n s of the in te r t i da I zone in which they were found than in the lower r e g i o n s ( F i g u r e 1 1 ) . In a d d i t i o n , p l a n t s were g e n e r a l l y longer at S epp i n g s I s l a n d than at F i r s t Beach w i t h i n a t i d a l h e i g h t . V e g e t a t i v e p l a n t s were of the same l eng th at both i n t e r t i d a l h e i g h t s , but p l a n t s at S epp i n g s I s l a n d were s i g n i f i c a n t l y sma l l e r than p l a n t s at F i r s t Beach d u r i n g t h i s s amp l i ng p e r i o d . E . B ranch ing pa t te rn s F i g u r e 12 shows t y p i c a l 5-rmi a p i c a l segments of a t e t r a s p o r o p h y t e , a female gametophyte and a male gametophyte of C. c o d i c o l a , in wh ich Tab le X . F - v a l u e s and a s s o c i a t e d p r o b a b i l i t i e s of s i g n i f i c a n c e ob ta i ned from two-way anova s , one w i t h month (M) and s i t e ( S ) , the other w i t h h e i g h t (H) and s i t e ( S ) as t rea tment s and l eng th of p l a n t s of v a r i o u s r e p r o d u c t i v e c l a s s e s as the dependent v a r i a b l e . T e t r a - Female Ma le V e g e t a t i v e s p o r o p h y t e s gametophytes gametophytes pI ants df F g F £> F g F p Two-way anova on month and s i t e : E r r o r df 1558 1302 677 350 M 9 35.17 < .001 4 5 . 63 < .001 40 . 20 001 30 .98 <. 001 S 1 62 .34 <.001 86. 59 < .001 60. 37 001 14.97 <. 001 M*S 9 6.47 < .001 5. 32 < .001 1. 60 <• 25 3 .23 <• 001 Two-way anova on he i gh t and s i t e : E r r o r df 669 552 248 96 H 1 24 .78 < .001 26. 75 < .001 7. 12 01 0 . 385 <. 75 S 1 53.80 < .001 52. 51 < .001 20. 29 001 5 .83 <• 025 H *S 1 5.41 < . 005 1. 90 < .25 7. 65 01 1.37 <. 25 F i g u r e 10. Mean l eng th s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s of Ceramium c o d i c o l a from F i r s t Beach and S e p p i n g s I s l a n d from specimens c o l l e c t e d in the months of the year 1981. E r r o r ba r s equa l s the MSD as c a l c u l a t e d from the GT2-method as d e s c r i b e d by Soka l and R o h l f ( 1 9 8 1 ) . I I I 1 I I 1 1 I I-M A M J J A S O N D Vegetative Plants 6H 5-4-3-2-E £ X o 6 H 2 LU 51 4 -3-2 -• First Beach O Seppings I. I V<5 Male Gametophyte —r-A —r~ 0 N "d" -1— A M F i g u r e 11. Mean l eng th s of a) t e t r a s p o r o p h y t e s , b) female gametophytes , c) male gametophytes and d) v e g e t a t i v e p l a n t s of Cerami um cod i co I a from F i r s t Beach (FB) and S e p p i n g s I s l a n d ( S I ) from m i d - (open c i r c l e s ) and l o w - i n t e r t i d a l ( c l o s e d c i r c l e s ) r e g i o n s . E r r o r bars equa l s the MSD as c a l c u l a t e d from the GT2-method as d e s c r i b e d in Soka l and R o h l f ( 1 9 8 1 ) . Tetrasporophytes \ Female Gametophytes r Male Gametophytes i i — — i Vegetative Plants 4) o T T o —i 1 1 r FB SI FB SI F i g u r e 12. R e p r e s e n t a t i v e 5-rmi a p i c a l segments of a) a female gametophyte, b) a t e t r a s p o r o p h y t e , and c) a male gametophyte of Ceramium cod i c o l a showing o v e r a l l b r anch i n g p a t t e r n s . d i f f e r e n c e s in o v e r a l l h ab i t can be seen. The S t r a h l e r method of o r d e r i n g b ranches p r o v i ded a u s e f u l q u a n t i t a t i v e measure of these d i f f e r e n c e s . The mean number of p r imary b ranches and o rde r s of b r a n c h i n g in segments of C. c o d i c o l a are g i v en in Tab le X I . In g e n e r a l , female gametophytes have s i g n i f i c a n t l y more p r imary b ranches than do male gametophytes which have more than t e t r a s p o r o p h y t e s . Ma le s at F i r s t Beach had a s i m i l a r number of p r imary b ranche s as females i n c l u d i n g i n v o l u c r a l r a m u l i . Except for ma l e s , there were no s i g n i f i c a n t d i f f e r e n c e s between s i t e s w i t h i n each r e p r o d u c t i v e c l a s s . The number of o r d e r s of b r a n c h i n g d i d not s i g n i f i c a n t l y d i f f e r among r e p r o d u c t i v e c l a s s e s or between s i t e s , except for males at F i r s t Beach which had s i g n i f i c a n t l y more. Few male gametophytes at F i r s t Beach exceeded 5 mm in l eng th and t h i s sample of s i x may not be c h a r a c t e r i s t i c of males as a who I e . The r a t i o s of f i r s t to second, second to t h i r d , and t h i r d to f o u r t h o rder b ranches w i t h 95% c o n f i d e n c e i n t e r v a l s are shown in F i g u r e 13 . F i r s t Beach male segments of 4 and 5 mm had s i m i l a r r a t i o s and were lumped. The r a t i o of f i r s t to second order b ranches at both s i t e s shows that females have a h i gher r a t i o than do males or t e t r a s p o r o p h y t e s , which are e q u a l . F i r s t Beach females have a h i ghe r r a t i o than those at S epp i n g s I s l a n d , but males and t e t r a s p o r o p h y t e s have equal r a t i o s at both s i t e s . B r a n c h i n g r a t i o s in females decrease s i g n i f i c a n t l y as branch order i n c r e a s e s w h i l e they remain the same in males and t e t r a s p o r o p h y t e s . The r a t i o s of t h i r d to f o u r t h order b ranches are equal or n e a r l y equal for a l l phases at both s i t e s . Tab le X I . Mean number of p r imary b ranches and b r anch i n g o rde r s +_ s t anda rd d e v i a t i o n fo r 5 rrm a p i c a l segments of t e t r a spo rophy te s - , female gametophytes ( w i t h or w i thout i n v o l u c r a l r amu l i ) and male gametophytes of C. c o d i c o l a . T e t r a -s p o r o p h y t e s Female gametophytes I n v o l u c r a l r a m u l i : Male gametophyt e s Wi th Wi thout # Pr imary b r anche s : F i r s t Beach S e p p i n g s I s l a n d # Br anch i ng order s : F i r s t Beach S epp i n g s I s l a n d Sample s i ze: F i r s t Beach Sepp i ng s I s l a n d 24 . 9 + 10 .03 89.1 + 39 .65 45 .1 + 21 . 79 79 . 3 + 27 .11 22 .3 + 11 .56 72.5 + 37.61 45 .0 + 29 .32 31.8 + 15 .95 4 . 55 + 0 .686 4 .45 + 0 . 759 20 20 4 . 1 3 + 0 . 834 4 .53 + 0 . 990 1 5 1 5 5 .50 + 0 . 837 4.80 + 1 .014 6 15 F i g u r e 13. Mean r a t i o s of h i ghe r to lower order b ranches of a) female gametophy te s - l ( i n c l u d i n g i n v o l u c r a l r a m u l i ) , b) female g a m e t o p h y t e s - l I ( e x c l u d i n g i n v o l u c r a l r a m u l i ) , c) t e t r a s p o r o p h y t e s and d) male gametophy te s . E r r o r bars equal +_ 1 s t anda rd e r r o r . Female Gametophytes- I 7H 6H 5 i I -< 4. CC H 0 2 1 O 3-< cr 00 2-3H Tetrasporophytes Female Gametophytes - I I • First Beach O Seppings I. Male Gametophytes 7 2 o 2 % q \y 2°/ 3 % DISCUSS ION A . Ep iphytes on Codium spp. Ceramium cod i c o I a was the most abundant ep iphy te among 30 s p e c i e s of e p i p h y t e s on C^ f r a g i l e , and was absent from C ^ s e t c h e l I i i . Two other a lgae r e p o r t e d as e p i p h y t e s of C^ fr ag i I e by Abbot t and H o l l e n b e r g ( 1 9 7 6 ) , Audou i ne I la rh i zo i dea and E r y t hr ot r i ch i a p u I v i n a t a , were found on both C. fr ag i I e and s e t che l I i i . A. rh i zo i dea i s r epo r ted to grow on other a l g a e , and it was commonly found on C^ se tche I I i i . Abbot t and H o l l e n b e r g ( 1976 ) c on s i de r E ^ pu I v i nata to be host s p e c i f i c , l i m i t e d to C^ fr ag i I e . I t was found twice in t h i s s t udy on C^ s e t c h e I I i i but i s much more abundant on C. f r a g i l e . The spongy c o n s t r u c t i o n of Cod i um spp. may not be as s u i t a b l e a s u b s t r a t u m for many a l gae as has been p o s t u l a t e d . B a l l a n t i n e (1979) found C. i s t hmocI ad i um in Pue r t o R i c o to harbor an i n t e rmed i a te number of e p i p h y t i c a l g a l s p e c i e s compared to other ho s t s he s u r v e y e d . He found 30 s p e c i e s of a l g a e on C^ i s t hmocI ad i um, wh ich compares we l l to the r e s u l t s of t h i s s tudy for C^ f r a g i l e . E p i p h y t e s on Cod i um spp. e x h i b i t at l e a s t three modes of a t t achment . Some e p i p h y t e s are p a r t i a l l y or e n t i r e l y en t ang l ed between the u t r i c l e s for a t tachment . T h i s was ob se r ved in Ce r am ium s p p . , Audoui n e l I a rh i zo idea and e c t o c a r p o i d s in t h i s s t u d y , and in Pt i I o thamn ionops i s Ie j o I i sea on C^ se t che I I i i by B i r d et a l . ( 1 9 8 2 ) . Other e p i p h y t e s cement themse lves to the " u t r i c l e s , such as Ant i thamn ion defectum and E r y t hr ot r i ch i a pu I v i n a t a . One other mode of attachment which was not ob se rved in t h i s s tudy i s that of growth w i t h i n the w a l l s of the u t r i c l e s , which is e x h i b i t e d by En t o c I ad i a cod i co I a (Abbott and H o l l e n b e r g 1 9 7 6 ) . O ' K e l l y and Y a r i s h (1981 ) found E. cod i co I a on u t r i c l e s of C^ fr ag i I e in B a r k l e y Sound. Many s p e c i e s of e p i p h y t e s were found on l y to occur in a reas s h e l t e r e d from wave e xpo su re . W i l s o n (1977 ) a l s o ob se rved g r ea te r numbers of e p i p h y t e s on O f r a g i l e in a s h e l t e r e d than in a wave-exposed s i t e in Rhode I s l a n d . Wave a c t i o n ac t s as an important s hap i n g f a c t o r of e p i p h y t i c a l g a l communit ies by removing those a l gae that are not we l l a t t a c h e d , are too l a r g e , or a comb ina t i on of bo th . The o b s e r v a t i o n of the i n t e r t i d a l d i s t r i b u t i o n of C^ fr ag i I e in B a r k l e y Sound agrees w i t h other s t u d i e s on C^ f r a g i l e on the west coas t of Vancouver I s l a n d (Widdowson 1965, Borden and S t e i n 1 9 6 9 ) . Widdowson (1965 ) a l s o r e p o r t e d C ^ setche I I i i to occur lower in the i n t e r t i d a l zone than C^ fr ag i I e and found it in a reas f u l l y exposed to wave a c t i o n . Hurd (1916) found C. f r a g i l e o c cu r r ed i n t e r t i d a l l y in Puget Sound . In the S t r a i t of G e o r g i a , C. f r a g i l e o ccu r s s u b t i d a l l y , 2 to 4 m below mean water leve l (MWL) and C. se tche I I i o c cu r s from 4 to 15 m below MiVL ( R . E . Foreman, unpub. o b s . ) S ou th (1968 ) r e p o r t s f i n d i n g s e ve r a l specimens of C^ fr ag i I e in the i n t e r t i d a l zone at G a b r i o l a I s l a n d . C^ f r a g i l e appears to occupy a lower p o s i t i o n in r e l a t i o n to water leve l in the S t r a i t of G e o r g i a than in B a r k l e y Sound. C. cod i co I a was not found as an ep iphy te in the one c o l l e c t i o n of C. fr ag i I e from the S t r a i t of G e o r g i a in t h i s s t u d y . Sou th ( 1968 ) t e n t a t i v e l y i n d e n t i f i e d some specimens of Ce r am i um e p i p h y t i c on C^ fr ag i I e as C. cod i co I a from G a b r i o l a I s l a n d . Other specimens of C^ f r a g i I e from the i S t r a i t of G e o r g i a in the U n i v e r s i t y of B r i t i s h Co lumbia He rba r i um d id not have C^ cod i c o I a as an e p i p h y t e . T h e r e f o r e , C^ cod i c o I a appears to be absent or rare in the S t r a i t of G e o r g i a . Many mar ine a lgae that occur in B a r k l e y Sound are not p re sen t in the S t r a i t of G e o r g i a (compare Scage l 1973 and Foreman and Root 1 9 7 5 ) . T h i s may be due to d i f f e r e n c e s in the temperature and s a l i n i t y of the sea water between these two a r e a s . Drueh l (1978) r e p o r t s that the s a l i n i t y of seawater at A n p h i t r i t e P t . ( a t the n o r t h e r n l i m i t of B a r k l e y Sound) ranged from 2 8 . 5 to 3 0 . 5 0/00 and the temperature ranged from 9 to 15°C, w h i l e at Chrome I s l a n d , near Hornby I s l a n d in the S t r a i t of G e o r g i a , the s a l i n i t y ranged from 2 5 . 0 to 2 8 . 5 0/00 and temperature from 7 to 18°C. In l a b o r a t o r y c u l t u r e , C^ cod i co I a from B a r k l e y Sound grew p o o r l y at 18°C, but C^ fr ag i I e from B a r k l e y Sound grew we l l at that t empera tu re . T h e r e f o r e , the h i gher temperature of seawater in the S t r a i t of G e o r g i a in the summer may l i m i t growth of C^ cod i co I a . Lower s a l i n i t y may a l s o render C. cod i co I a more s u s c e p t i b l e to h i gh t e m p e r a t u r e s . C^ cod i c o I a does occur south of P o i n t C o n c e p t i o n , C a l i f o r n i a , and into Baja C a l i f o r n i a , where the water temperature may exceed 18°C ( e . g . G u n n i l l 1 9 8 0 ) . T h i s may i n d i c a t e some e c o t y p i c d i f f e r e n t i a t i o n between p o p u l a t i o n s of C^ cod i c o I a between B r i t i s h Co lumbia and s ou the rn C a l i f o r n i a and Ba ja C a l i f o r n i a . B. Cu l t u re experiments The fact that C. cod i co I a grew from spo re s and produced r e p r o d u c t i v e s t r u c t u r e s in l a b o r a t o r y c u l t u r e i n d i c a t e s that i t i s n u t r i t i o n a l l y and c h e m i c a l l y independent of C. f r a g i l e . The p resence of e i t h e r l i v i n g Cod i um or e x t r a c t of Cod i um d id not enhance growth or r e p r o d u c t i o n and i n h i b i t e d i t in some c a s e s . I n h i b i t i o n by the host in c u l t u r e may be the r e s u l t of n u t r i e n t c o m p e t i t i o n or a l l e l o p a t h y in the l i m i t e d volume of the c u l t u r e v e s s e l and may not be as important in the f i e l d . The l i f e h i s t o r y of C. c o d i c o l a was not completed in c u l t u r e , but t h i s may be a r e s u l t of c u l t u r e c o n d i t i o n s used in t h i s s t u d y , and not n e c e s s a r i l y a d e m o n s t r a t i o n that C. cod i co I a r e q u i r e s C^ fr ag i I e in order to do so . C^ pac i f i cum reproduced in s i m i l a r c o n d i t i o n s , but the second g e n e r a t i o n s p o r e l i n g s grew much s lower than the f i r s t g e n e r a t i o n , i n d i c a t i n g a reduced v i a b i l i t y in the second g e n e r a t i o n . C. cod i co I a i s much sma l l e r than C. pac i f i cum, and a r e d u c t i o n in v i a b i l i t y may p revent growth from o c c u r r i n g at a l l . C . c o d i c o l a i s smal l both in the f i e l d and in c u l t u r e . Both C. pac i f i cum and C ^ gardner i grow l a r ge r than C^ cod i co I a in the sea and in c u l t u r e . Thai I i of C. c o d i c o l a sampled for r e p r o d u c t i v e data in t h i s s tudy were 1 . 5 rim in l eng th or l o n g e r , and 8 8 . 5 % were r e p r o d u c t i v e . Most c a r p o s p o r e l i n g s grown in c u l t u r e at 10 or 12°C that grew to a s i z e of 1 . 7 mm or g r ea te r were r e p r o d u c t i v e . I t i s p o s s i b l e that r e p r o d u c t i o n in C. c o d i c o l a i s r e l a t e d to s i z e of the t h a l l u s . Smal l s i z e may confer a c o m p e t i t i v e advantage to C. c o d i c o l a over other Ceramium spp . g row ing on the s u r f a c e of C. f r a g i l e in a reas exposed to wave a c t i o n . Sma l l e r s i z e would r e s u l t in l e s s drag from water m o t i o n , wh ich would decrease the p o s s i b i l i t y of be i ng washed away. In s h e l t e r e d a r e a s , C. c o d i c o l a i s longer and C. p a c i f i c u m and C. gardner i are more common. I n d i v i d u a l s of C. pac i f i c um e p i p h y t i c on C. fr ag i I e in s h e l t e r e d t i de poo l s have been obse rved to be up to 5 cm in l e n g t h . Cer ami um pac i f i cum, wh ich was c omp le te l y c o r t i c a t e d when c o l l e c t e d in the f i e l d , , produced la rge naked i n te rnode s in c u l t u r e . T h i s i n d i c a t e s that c o r t i c a t i o n may be a f f e c t e d by env i ronmenta l parameters and may not be a r e l i a b l e taxonomic c r i t e r i o n . C. rubrum from A t l a n t i c N o r t h A n e r i c a was f u l l y c o r t i c a t e d when grown under sho r t d a y l e n g t h s in c u l t u r e and deve loped naked i n t e rnode s when grown under long d a y l e n g t h s (Ga rba r y et a l . 1 9 7 8 ) . A l l c u l t u r e s in t h i s s t udy were grown in long d a y l e n g t h ( 1 6 : 8 LD) c o n d i t i o n s , These r e s u l t s i n d i c a t e that development of c o r t i c a t i o n may be r e l a t e d to d a y l e n g t h in C^ pac i f i cum, as in C;_ rubrum. Ky I i n ( 1925 ) d e s c r i b e d C. w a s h i n g t o n i e n s e as an a l g a s i m i l a r to C^ pac i f i c um, except that i t i s s m a l l e r , of a l i g h t e r c o l o r and has l i g h t e r c o r t i c a t i o n . S i z e and co l o r may a l s o be a f f e c t e d by env i ronmenta l pa ramete r s , and C. w a s h i n g t o n i e n s e may be c o n s p e c i f i c w i t h C^ pac i f i cum. C . R h i z o i d morphology P h y s i c a l c h a r a c t e r i s t i c s of the s u b s t r a t u m appear to be important for g rowth of C. cod i co I a in nature by a f f e c t i n g the a b i l i t y to a t t a ch s e c u r e l y . R h i z o i d s of some P o I y s i phon i a spp . produce f i l amen tou s or d i s c o i d r h i z o i d a l a t t a chmen t s , the l a t t e r in re sponse to f i r m s u b s t r a t u m contact (Chen and T a y l o r 1968, F l e t c h e r 1 9 7 9 ) . R h i z o i d s of Ce r am i um rub rum may be s h o r t , p r o d u c i n g a much d i v i d e d h o l d f a s t , or e l o n g a t e , r e l y i n g on t e n d r i l l a r a c t i v i t y to ach ieve attachment (Jones and Duerden 1 9 7 2 ) . In t h i s s t udy , r h i z o i d s of C^ pac i f i cum, C. gardner i and C^ c a l i f o r n icum were obse rved to produce both much d i v i d e d h o l d f a s t s at r h i z o i d t i p s and e l onga te r h i z o i d s , w h i l e C^ cod i co I a and C ^ ea ton i anum produced o n l y e l onga te r h i z o i d s . The a b i l i t y to produce a cement ing s t r u c t u r e such as a d i s c o i d or much d i v i d e d h o l d f a s t would f a c i l i t a t e attachment on s o l i d s u b s t r a t a . A l gae p roduc i ng o n l y e l onga te r h i z o i d s would be l i m i t e d to s u b s t r a t a into which the r h i z o i d s c o u l d e n t a n g l e . C^ f r a g i l e p r o v i d e s such a s u b s t r a t u m for C^ cod i c oI a and c u l t u r e exper iment s i n d i c a t e d that C. se t che I I i i i s too compact to be a s u i t a b l e s u b s t r a t u m for p e n e t r a t i o n by r h i z o i d s of C^ cod i co I a . The bu lbous t i p s of r h i z o i d s of C^ cod i co I a may he lp to ensure ancho r i n g between u t r i c l e s of C. fr ag i I e . Young s p o r e l i n g s of C. c o d i c o l a produced r h i z o i d s w i t h bu lbous t i p s , w h i l e s p o r e l i n g s of C. p a c i f i c u m and Mic roc I ad i a c o u l t e r i d i d not , i n d i c a t i n g that t h i s c ha r a c t e r is g e n e t i c a l l y determined in C. c o d i c o l a . R h i z o i d s of C. c o d i c o l a s p o r e l i n g s grown on C. fr a g i l e in c u l t u r e r e t a i n e d bu lbous t i p s w h i l e r h i z o i d s of those grown w i t hou t C. f r a g i l e in c u l t u r e had narrow non -bu lbou s r h i z o i d s g rowing out from expanded p o r t i o n s . The p h y s i c a l p r o x i m i t y of u t r i c l e s to these r h i z o i d s may p reven t the f o rmat i on of these narrow r h i z o i d s . Po I y s i phon i a ruborh i za and P. s a c c o r h i za , wh ich occur as e p i p h y t e s of Cod i um spp. in the t r o p i c a l P a c i f i c O c e a n T a l s o have bu lbous t i p s on r h i z o i d s that penet ra te between the u t r i c l e s ( H o l l e n b e r g 1 9 6 8 ) . D. Pheno log i ca l phenomena The f r e q u e n c i e s of v a r i o u s r e p r o d u c t i v e c l a s s e s of C^ cod i c oI a ob t a i ned du r i n g d i f f e r e n t months of the year i n d i c a t e that a l a r ge p r o p o r t i o n o f the p o p u l a t i o n is f e r t i l e th roughout the y e a r . A seasona l a l t e r n a t i o n of g e n e r a t i o n s i s not i n d i c a t e d by these d a t a T and C ^ cod i c oI a can be c o n s i d e r e d to be of the l i f e - f o r m Ephemerophyceae a c c o r d i n g to the t e r m i n o l o g y of G a r b a r y (1 976) or an a seasona l annual a c co rd i n g to Sea r s and Wi I ce ( 1 9 7 5 ) . C. pac i f i cum and C ^ gardner i were p re sen t and r e p r o d u c t i v e th roughout most of the year and showed no ev idence of a sea sona l a l t e r n a t i o n of g e n e r a t i o n s . In C. c o d i c o l a T seasona l t rends were apparent in the f r e q u e n c i e s of t e t r a s p o r i c - , c y s t o c a r p i c and v e g e t a t i v e p l a n t s and in l eng th s of a l l r e p r o d u c t i v e c l a s s e s , w i t h fewer i n d i v i d u a l s p r oduc i n g spores in w in te r and p l a n t s be ing s h o r t e s t in w i n t e r . Go f f and Co le ( 1976 ) showed that t e t r a s p o r o g e n e s i s T g a m e t o g e n e s i s T and c a r p o s p o r o g e n e s i s in Ha r veye I I a m i r ab i I i s in B r i t i s h Co lumbia is c o r r e l a t e d w i t h h i gher temperature and/or l i g h t a b v a i I a b i I i t y . Coo l e r temperatures and lower l i g h t a v a i l a b i l i t y in the w in te r may n e g a t i v e l y a f f e c t r e p r o d u c t i o n of C. c o d i c o l a at t h i s time of y e a r . Compar i s on s between two s i t e s of d i f f e r e n t wave exposure and between two i n t e r t i d a l h e i g h t s at each of the s i t e s d id not revea l any s i g n i f i c a n t e f f e c t s of these parameters on f r e q u e n c i e s of r e p r o d u c t i v e p l a n t s . The f requency of f e r t i l e gametophytes s i g n i f i c a n t l y exceeds that of f e r t i l e te tr a s p o r o p h y t e s . However.,, i f a l l or most of the v e g e t a t i v e p l a n t s sampled were immature t e t r a s p o r o p h y t e s T then gametophyte and spo rophyte f r e q u e n c i e s are not s i g n i f i c a n t l y d i f f e r e n t , , i n d i c a t i n g that a ba lance occu r s between these two a l t e r n a t e p h a s e s . Among gametophy te s T females were more numerous and of g r ea te r l e ng th than males th roughout the y e a r . T h e o r e t i c a l l y , , ma les and females a r i s e w i t h equal f requency from t e t r a s p o r e s . These d i f f e r e n c e s in f requency and l eng th i n d i c a t e e i t h e r that females are longer l i v e d than males,- or that more females s u r v i v e than males and females grow f a s t e r . The former e x p l a n a t i o n seems more l i k e l y . D i f f e r e n c e s in l eng th between s i t e s i n d i c a t e s that wave exposure,, the most obv i ou s f a c to r d i f f e r e n t between s i t e s T n e g a t i v e l y a f f e c t s the l eng th of C. cod i c o I a on C^ fr ag i I e . T h i s may be a t t r i b u t e d to the wash ing away of l a r g e r p l a n t s of C^ cod i c o I a or to the b r e a k i n g of b r a n c h e s . In f u l l y s h e l t e r e d a r e a s , C^ pac i f i cum was o f t e n abundant and p l a n t s up to 5 cm in l e n g t h were o b s e r v e d . C. cod i co I a was longer on fr ag i I e p l a n t s that were h igher in the i n t e r t i d a l zone,, e s p e c i a l l y at S e p p i n g s I s l a n d . T h i s may be the r e s u l t of l i g h t l i m i t a t i o n or i n c r ea sed g r a z i n g in the lower i n t e r t i d a l r e g i o n s . H i ghe r in the i n t e r t i d a l x i t i s u n l i k e l y that C^ cod i co I a undergoes much d e s i c c a t i o n s i n ce the spongy t h a l l u s of C^ fr ag i I e may p r o v i d e m o i s t u r e dur ing emergenee. E. B ranch ing pa t te rn T e t r a s p o r o p h y t e s and female gametophytes of C^ cod i c oI a ( t h i s t h e s i s ) and C. rubrum (Garbary,. Grund and McLach lan 1980 ) e x h i b i t s i m i l a r b r anch i ng p a t t e r n s . The p r ima r y to s econda ry b ranch r a t i o was the same for t e t r a s p o r o p h y t e s of both s p e c i e s at 2 . 5 and female gametophytes e x h i b i t e d a h i ghe r r a t i o than t e t r a s p o r o p h y t e s in both species., a l t h o u g h t h i s r a t i o was h i g h e r in C^ cod i co I a . The number of p r imary b ranches i s not comparable because d i f f e r e n t l eng th s of a p i c a l segments was used in these two s t u d i e s . The b r a n c h i n g p a t t e r n of male gametophytes of C^ cod i col a was c h a r a c t e r i z e d as h a v i n g a sma l l e r or equal number of p r imary b ranches per u n i t l eng th as females and a h igher number than t e t r a s p o r o p h y t e s . The r a t i o of p r imary to secondary b ranches in males was l e s s than that in females., but equal to that of t e t r a s p o r o p h y t e s . Male gametophytes may a l s o have more o r d e r s of b r a n c h i n g per u n i t l eng th than do females or t e t r a s p o r o p h y t e s . T h i s would account for the h i gher number of p r imary b ranches p re sent in males than in t e t r a s p o r o p h y t e s even though they have equal b r anch i n g r a t i o s . P a c k i n g of more o r de r s of b r a n c h i n g into the same l eng th may r e s u l t from h a v i n g sma l l e r segments between b r a n c h i n g po in t s , , l a r g e r b r a n c h i n g a n g l e s , or b o t h . G a r b a r y T Grund and McLach l an (1980 ) showed that female gametophytes of C. rubrum had l a rge r ang l e s at b r a n c h i n g p o i n t s than d id t e t r a s p o r o p h y t e s . B r a n c h i n g ang le s were not measured in t h i s s t udy , but in F i g u r e 12, i t i s apparent that both male and female gametophytes have l a r g e r ang l e s at b r a n c h i n g p o i n t s than do t e t r a s p o r o p h y t e s . Gametophytes p r i m a r i l y d i f f e r from spo rophy te s by h a v i n g l a r ge r b r a n c h i n g a n g l e s . Female gametophytes d i f f e r from males by h a v i n g a more p r o l i f e r o u s b r a n c h i n g p a t t e r n . The lower b r a n c h i n g r a t i o s ob ta i ned in female gametophytes at h i gher order b ranches ( F i g u r e 13) i n d i c a t e s that they e x h i b i t a more r e g u l a r l y d ichotomous b r a n c h i n g p a t t e r n e a r l y in t he i r development and deve lop more p r o l i f e r o u s b ranches l a t e r . G a r b a r y , Grund and McLach l an (1980 ) p o i n t out * that some i n v o l u c r a l ramul i s u r r o u n d i n g c y s t o c a r p s con t i nue to grow, which would i nc rea se the b r a n c h i n g r a t i o . Young females would p r o b a b l y not produce c y s t o c a r p s be fo re at l ea s t 1 or 2 b r a n c h i n g s have a l r e a d y o c c u r r e d , a f t e r wh ich the e f f e c t s of the a d d i t i o n of i n v o l u c r a l ramul i would be ev iden t in the b r a n c h i n g p a t t e r n . T h i s seems to be the case he re . I t i s p o s s i b l e that the g rea te r p r o l i f e r a t i o n of p r imary branches in female gametophytes is of a d a p t i v e s i g n i f i c a n c e . Females of Ceramium spp. produce c a r p o g o n i a near the ap i ce s of p r imary b r a n c h e s , w h i l e spe rmat i a are produced a long the branches from c o r t i c a l c e l l s , and t e t r a s p o r a n g i a a long the b ranches from p e r i c e n t r a l c e l l s ( F r i t s c h , 1945 ) . A female gametophyte that p roduces more ap i ce s would be able to produce more c a r p o g o n i a , w h i l e a s i m i l a r i n c rea se in the number of p r imary b ranches would not r e s u l t in an i n c r ea s e in the p r o d u c t i o n of r e p r o d u c t i v e c e l l s in male gametophytes and t e t r a s p o r o p h y t e s . T h e r e f o r e , he te romorph i sm in the d i f f e r e n t phases of C. c o d i c o l a , and other Ceramium s p p . , may occur to maximize r e p r o d u c t i v e po tent i a I . SUMMARY R e s u l t s of t h i s s t udy on the b i o l o g y of Cerami um cod i c o I a are summarized below: 1. Ce r am i um cod i co I a i s the most abundant ep i phy te of Cod i um fr ag i I e in a rea s exposed to wave a c t i o n . Other e p i p h y t e s may be more abundant in s h e l t e r e d a r e a s . C^ cod i co I a i s absent from C ^ se t che I I i i . 2 . R e s u l t s of c u l t u r e exper iment s a r e : a . Ceramium c o d i c o l a i s ab le to grow to r e p r o d u c t i v e m a t u r i t y in c u l t u r e apar t from Cod i um fr ag i I e , i t s o n l y known h o s t . b . The presence of l i v i n g or e x t r a c t e d C ^ fr ag i I e or C^ s e t c h e I I i i does not enhance growth or r e p r o d u c t i o n , and o f ten i n h i b i t s i t in c u l t u r e . c . R h i z o i d s of C^ cod i co I a u s u a l l y pene t r a te the t h a l l u s of C^ fr ag i I e in c u l t u r e w h i l e they u s u a l l y do not pene t r a te C^ se tc he I I i i . d . G e r m i n a t i o n and growth of spo re s of C^ cod i co I a i s be t te r at 10 and 12°C than at 15°C or h i g h e r . e . Ce r am ium pac i f i cum d i s p l a y s incomplete c o r t i c a t i o n in c u l t u r e , which may be the r e s u l t of d a y l e n g t h . C ^ pac i f i cum and C ^ w a s h i n g t o n i e n s e may t h e r e f o r e be c o n s p e c i f i c . 3. R h i z o i d s of some Ceramium spp . produce a d i g i t a t e h o l d f a s t s t r u c t u r e . T h i s s t r u c t u r e was not found on C. c o d i c o l a , i n d i c a t i n g that attachment by r h i z o i d s of C^ cod i co I a i s dependent on entang lement w i th a l o o s e l y c o n s t r u c t e d s u b s t r a t u m . 4 . Ce r am i um cod i c o l a i s r e p r o d u c t i v e th roughout the y e a r . R e p r o d u c t i o n and p l an t s i z e are reduced in the w i n t e r , wh ich i s p r o b a b l y the r e s u l t of c o o l e r temperatures and lower l i g h t a v a i l a b i l i t y d u r i n g t h i s t ime. P l an t s i z e is l a r ge r in a reas s h e l t e r e d from wave a c t i o n , and in the upper r e g i o n s of the i n t e r t i d a l d i s t r i b u t i o n of C. c o d i c o l a . 5. R e p r o d u c t i v e phases of C^ cod i col a e x h i b i t he te romorph i sm in b r anch i n g p a t t e r n which may be r e l a t e d to o p t i m i z a t i o n of r e p r o d u c t i v e p o t e n t i a l . L ITERATURE C I TED A b b o t t , I .A. and G . J . H o l l e n b e r g . 1976 . The Ma r i n e A l g a e of C a l i f o r n i a . S t a n f o r d U n i v e r s i t y P r e s s . S t a n f o r d , C a l i f o r n i a . 827 pp . A g a r d h , J .G. 1894. A n a l e c t a a l g o l o g i c a . C o n t . I . Lunds U n i v . A r s s k r . 29: 1 - 1 4 4 . Anonymous. 1980, 1981 . Canad ian T i de and C u r r e n t T a b l e s . V o l . 6 . B a r k l e y Sound and D i s c o v e r y Pa s s age to D i x o n E n t r a n c e . Government of Camada, F i s h e r i e s and O c e a n s . S c i e n t i f i c I n f o r m a t i o n and P u b l i c a t i o n s B ranch , O t tawa. A r a s a k i , S . , H. Tokuda, and K . F u j i a m a . 1955. The r e p r o d u c t i o n and morphogeny in Cod i um f r a g i l e . B o t . Mag. (Tokyo ) 69 : 3 9 - 4 5 . ( I n J apanese) B a l l a n t i n e , D . L . 1979 . The d i s t r i b u t i o n of a l g a l e p i p h y t e s on macrophyte h o s t s o f f s h o r e from La P a r g u e r a , Puer to R i c o . B o t . M a r . 2 2 : 1 07 - 110 . B a r k e r , S . B . , G. Cuming and K . H o r s f i e l d . 1973 . Q u a n t i t a t i v e  morphometry of b r a n c h i n g s t r u c t u r e of t r e e s . J . Theo r . B i o l . 4 0 : 3 3 - 4 3 . B i r d , C . J . , D. G a r b a r y and G. I . Han sen . 1982. O b s e r v a t i o n s on P t i I o t h a m n i o n o p s i s l e j o l i s e a ( F a r l . ) D i x . ( Ce ram iaceae , Rhodophyta) in N o r t h w e s t e r n N o r t h A n e r i c a . S y e s i s 14 : 1 0 9 - 1 1 3 . Boney, A .D . 1980 . P o s t - a t t a c h m e n t r e spon se s of monospores of some endophy t i c Audou i neI la spp . ( N e m a l i a l e s : F I o r i d e o p h y c e a e ) . Nova Hedw. 33: 4 9 9 - 5 0 7 . Bo rden , C .A . and J . R . S t e i n . 1969. R e p r o d u c t i o n and e a r l y development in Cod i um fr ag i I e ( S u r i n g a r ) H a r i o t : C h I o r o p h y c e a e . P h y c o l o g i a 8 : 9 1 - 9 9 . Chen, L . C . - M . and A . R . A . T a y l o r . I 9 6 8 . R h i z o i d development in P o I y s i phon i a . B r . p h y c o l . B u l l . 3 : 6 00 . ( A b s t r a c t ) C i t h a r e l , J . 1972 . P o I y s i phon i a I anosa ( L . ) Tandy est i l un s imp le e p i p h y t e ? Acadamie des S c i e n c e s . C .R . D274: 1 904 - 6 . Codomie r . L . , M. Segot et G . Combaut. 1981 . I n f l u e n c e de composes o r g an i que s ha l o gene s sur la c r o i s s a n c e d ' A s p a r a g o p s i s armata (Rhodophycee, Bonnema i son i a I e ) . B o t . Ma r . 24 : 509-51 3 . Conove r , J . T . and J .McN. S i e b u r t h . 1964 . E f f e c t of S a r g a s s um d i s t r i b u t i o n on i t s e p i b i o t a and a n t i b a c t e r i a l a c t i v i t y . B o t . Ma r . 6 : 1 47 - 157 . D i x o n , P . S . 1973 . B i o l o g y of the Rhodophyt a . O l i v e r <& Boyd . E d i n b u r g h . 285 pp. D r u e h l , L . D . 1978 . The d i s t r i b u t i o n of Macrocys t i s i n teg r i f o I i a in B r i t i s h Co lumbia as r e l a t e d to env i ronmenta l p a r a m e t e r s . Can . J . B o t . 56: 6 9 - 7 9 . Du R i e t z , G . E . 1930 . S t u d i e s in the taxonomy and e c o l o g y of Cer ami um d i aphanum in the B a l t i c . B o t a n i s k a N o t i s e r 4 3 3 - 4 5 8 . Ende, G. van den, und R. van O o r s c h o o t . 1962. W e i t e r e Beobachtungen ueber den Ep iphy tenbewuchs von H i m a n t h a l i a e I o n g a t a ( L . ) S . F . G r a y . B o t . M a r . 5: 1 1 1 - 1 2 0 . E v a n s , L . V . , J .A . C a l l o w , and M .E . C a l l o w . 1973 . S t r u c t u r a l and p h y s i o l o g i c a l s t u d i e s on the p a r a s i t i c red a l g a H o I m s e l l a . New P h y t o l . 72: 3 9 3 - 4 0 2 . F l e t c h e r . R . L . 1979 . S t u d i e s on attachment in the red a l g a l genus P o I y s i phon i a . B r . p h y c o l . J . 14 : 123. ( A b s t r a c t ) Foreman, R . E . and J . Roo t . 1975 . Macrophyte s t u d i e s in the S t r a i t of G e o r g i a : A r e v i e w of the c u r r e n t s t a t u s of macrophyte knowledge and r e s e a r c h . BERP Report 7 5 - 2 . 52 pp . F r i t s c h , F . E . 1945 . The S t r u c t u r e and Reproduct ion of the Al gae . V o l . I I . U n i v e r s i t y P r e s s . Cambr idge . 939 pp. G a r b a r y , D. 1976 . L i f e - f o r m s of a l g ae and t h e i r d i s t r i b u t i o n . Bo t . Ma r . 19: 9 7 - 1 0 6 . G a r b a r y , D . J . , D. Grund and J . M c L a c h l a n . 1978. The taxonomic s t a t u s of Ceramium rub rum (Huds . ) C . Ag . ( C e r a m i a l e s , Rhodophyceae) based on c u l t u r e e x p e r i m e n t s . P h y c o l o g i a 17: 8 5 - 9 4 . G a r b a r y , D . , D.W. Grund and J . M c L a c h l a n . 1980 . B r a n c h i n g p a t t e r n s and l i f e h i s t o r y s t a g e s in Ceramium rub rum (Hud s . ) C. A g . Nova Hedw. 33: 2 4 9 - 2 6 0 . G a r b a r y , D . J . , G . I . Hansen and R . F . S c a g e l . 1980. The mar ine a l gae of B r i t i s h Co lumbia and N o r t h e r n W a s h i n g t o n : D i v i s i o n Rhodophyta (Red a l g a e ) . C l a s s B a n g i o p h y c e a e . S y e s i s 1 3: 1 37 - 195 . G o f f , L . J . 1979. The b i o l o g y of Har veye I I a mi r ab i I i s ( C r y p t o n e m i a ! e s , Rhodophyceae ) . V I . T r a n s l o c a t i o n of p h o t o a s s i m i I a t e d C. J . P h y c o l . 15: 8 2 - 8 7 . G o f f , L . J . and K . C o l e . 1976. The b i o l o g y of Ha r veye I I a mi r ab i I i s (C r yp tonemia I e s , Rhodophyceae ) . IV. L i f e h i s t o r y and pheno l o g y . Can. J . B o t . 54 : 2 8 1 - 2 9 2 . G o n z a l e z , E . A . and L . J . G o f f . 1980 . E r y t h r o c y s t i s s a c c a t a : A b i o c h e m i c a l l y o b l i g a t e s ymb ion t ? J . P h y c o l . 16 ( s u p p l . ) : 14. ( A b s t r a c t ) G u n n i l l , F .C . 1980 . Rec ru i tment and s t a n d i n g s t o c k s in p o p u l a t i o n s of one g reen a l g a and f i v e brown a l gae in the i n t e r t i d a l zone near La J o I l a , C a l i f o r n i a d u r i n g 1 9 7 3 - 1 9 7 7 . Ma r . E c o l . P r o g . S e r . 3 : 2 3 1 - 2 4 3 . Ha I I am, H . D . , M.N. C l a y t o n and D . P a r i s h . 1980. S t u d i e s on the a s s o c i a t i o n between Nothe i a anomaI a and Hormos i r a banks i i ( P h a e o p h y t a ) . A u s t . J . B o t . 28: 2 3 9 - 2 4 8 . H a r l i n , M.M. 1973a . " O b l i g a t e " a l g a l e p i p h y t e : Smi t ho r a na i adum grows on a s y n t h e t i c s u b s t r a t e . J . Phycol 9 : 2 3 0 - 2 3 2 . H a r l i n , M.M. 1973b . T r a n s f e r of p r oduc t s between e p i p h y t i c mar ine a lgae and hos t p l a n t s . J . P h y c o l . 9 : 2 4 3 - 2 4 8 . H a r l i n , M.M. and J . S . C r a i g i e . 1975 . The d i s t r i b u t i o n of p h o t o s y n t h a t e in A s c o p h y I Ium nodos um as i t r e l a t e s to e p i p h y t i c P o I y s i phon i a I ano sa . J . P h y c o l . 11 : 1 0 9 - 1 1 3 . den H a r t o g , C. 1972 . S u b s t r a t u m . P l a n t s . M u l t i c e l l u l a r p l a n t s . In K i n n e , O. ( e d . ) Mar i ne E c o I o g y . V o l . 1 . Env i r onmenta l f a c t o r s . P t . 3 . W i I e y - l n t e r s c i e n c e . London. Pp. 1 2 7 7 - 1 2 8 9 . Hawkes, M.W., C . E . Tanne r , and P . A . L e b e d n i k . 1978 . The ben th i c mar ine a lgae o f n o r t h e r n B r i t i s h Co l umb i a . S y e s i s 11 : 8 1 - 1 1 5 . H o l l e n b e r g , G . J . 1968 . An account of the s p e c i e s of P o I y s i phon i a of the c e n t r a l and we s te rn t r o p i c a l P a c i f i c Ocean . I . 0 1 i g o s i p h o n i a . Pac . S c i . 2 2 : 5 6 - 9 8 . Hu rd , A .M. 1916 . Cod i um mucronatum. P u b l . Puget Sound mar. b i o l . S t n . 1: 1 0 9 - 1 2 3 . J o n e s , W.E. and R . C . Due rden . 1972 . V a r i a t i o n s in the morpho logy of the p r i m a r y r h i z o i d in Cerami um rubrum (Hud s . ) C . Ag . B r . phyco l J . 7: 281 . ( A b s t r a c t ) K y l i n , H. 1925 . The mar ine a l gae in the v i c i n i t y of the b i o l o g i c a l s t a t i o n at F r i d a y H a r b o r , W a s h i n g t o n . Lunds U n i v . A r s s k r . 2 1 : 1 - 8 7 . Lew in , J .C . 1966 . S i l i c o n m e t a b o l i s m in d i a t o m s . V . Germanium d i o x i d e , a s p e c i f i c i n h i b i t o r of d ia tom g rowth . P h y c o l o g i a 5: 1 - 1 2 . L e w i s , I . F . 1 914 . The sea sona l l i f e - c y c l e of some red a lgae at Woods H o l e . P l a n t Wor ld 17 : 3 1 - 3 5 . L i n s k e n s , H .F . 1963a . B e i t r a g zur F rage der Bez iehungen zw i schen Ep i phy t und B a s i p h y t bei mar i nen A l g e n . P u b l . S t a z . Zoo I . , N a p o l i . 33 : 2 7 4 - 2 9 3 . L i n s k e n s , H . F . 1963b. O b e r f I a c h e n s p a n n u n g an mar i nen a l g e n . P r o c . N e d e r l a n d s e Akademie V . Wetenschapfen C66: 2 0 5 - 2 1 7 . L i n s k e n s , H . F . 1966. A d h a e s i o n von F o r t p f I a n z u n g s z e I I en b e n t h o n t i s c h e r A l g e n . P l a n t a ( B e r l . ) 68: 9 9 - 1 1 0 . L i t t l e , R . J . and C . E . J one s . 1980 . A D i c t i o n a r y of B o t a n y . Van N o s t r a n d R e i n h o l d Co . New Y o r k . 400 pp . Markham, J .W. 1969. V e r t i c a l d i s t r i b u t i o n of e p i p h y t e s on the s t i p e of Ner eocys t i s I ue t keana ( M e r t e n s ) P o s t e l s and R u p r e c h t . S y e s i s 2: 2 2 7 - 2 4 0 . M c L a c h l a n , J . 1973 . Growth m e d i a - m a r i n e . In S t e i n , J .R . ( e d . ) Handbook of P h y c o I o g i ca I Me thod s : C u I t u r e Methods and Growth Mea su rement s . Cambr idge U n i v e r s i t y P r e s s , London. Pp. 2 5 - 5 1 . M o s s , B . S . 1981. The c o n t r o l of e p i p h y t e s by H a l i d r y s s i I i quosa ( L . ) L y n g b . I n t . B o t . C o n g r e s s . 13 : 170. ( A b s t r a c t ) Nonomura, A .M. and J .A . W e s t . 1981a . Seasona l growth of the p a r a s i t e J anczewsk i a on Lau renc i a ( Rhodophy ta , C e r a m i a l e s ) in C a l i f o r n i a (USA) and H o k k a i d o ( j a p a n ) . B o t . Ma r . 2 4 : 3 49 - 359 . Nonomura, A .M. and J . A . W e s t . 1981b. Host s p e c i f i c i t y of J anczewsk i a ( C e r a m i a l e s , R h o d o p h y t a ) . P h y c o l o g i a 20 : 2 5 1 - 2 5 8 . Odum, E . P . 1971 . Fundamenta l s of E c o I o g y , 3 rd ed. S a u n d e r s . P h i l a d e l p h i a . 574 pp . O ' K e l l y , C . J . and C. Y a r i s h . 1981 . O b s e r v a t i o n s on mar ine Chae topho raceae . I I . On the c i r c u m s c r i p t i o n of the genus E n t o c I ad i a R e i n k e . P h y c o l o g i a 20: 3 2 - 4 5 . Peno t , M. 1974 . I o n i c exchange between t i s s u e s of A s c o p h y I I um nodos um ( L . ) L e j o l i s and P o I y s i phon i a I a no sa ( I . ) Tandy . Z. P f I a n z e n p h y s i o I o g i e 73: 1 2 5 - 1 3 1 . Ramus, J . 1972 . D i f f e r e n t i a t i o n of the green a l g a Cod i um fr ag i I e . An . J . B o t . 59 : 4 7 8 - 4 8 2 . R o h l f , F . J . and R . R . S o k a l . 1981 . S t a t i s t i c a l Tab I e s , 2nd ed. W.H. Freeman and C o . San F r a n c i s c o . 211 pp . S c a g e l , R . F . 1966 . Ma r i ne a l gae of B r i t i s h Co lumbia and n o r t h e r n W a s h i n g t o n . Pa r t I . Ch I o r ophyceae ( g reen a l g a e ) . N a t . Mus . Can . B u l l . 207 . 257 pp. S c a g e l , R . F . 1973 . M a r i n e b e n t h i c p l a n t s in the v i c i n i t y of B a m f i e l d , B a r k l e y Sound, B r i t i s h Co l umb i a . S y e s i s 6 : 1 2 7 - 1 4 5 . S e a r s , J . R . and R .T . W i l c e . 1975 . S u b l i t t o r a l , ben th i c mar ine a l gae of s o u t h e r n Cape Cod and ad jacen t i s l a n d s : Seasona l p e r i o d i c i t y , a s s o c i a t i o n , d i v e r s i t y , and f l o r i s t i c c o m p o s i t i o n . E c o l . Monog r . 45 : 3 3 7 - 3 6 5 . S e t c h e l l , W.A. 1918 . P a r a s i t i s m among the red a l g a e . P r o c . An. P h i l . S o c . 57: 1 5 5 - 1 7 2 . S i l v a , P .C . 1951. The genus Cod i um in C a l i f o r n i a . U.C. Pub. B o t . 25 : 7 9 - 1 1 4 . S locum, C . J . 1980 . D i f f e r e n t i a l s u s c e p t i b i l i t y to g r a z e r s in two phases of an i n t e r t i d a l a l g a : advan tage s of he te romorph i c g e n e r a t i o n s . J . exp. mar . B i o l . E c o l . 4 6 : 9 9 - 1 1 0 . S o k a l , R .R . and F . J . R o h l f . 1981. B i o m e t r y . 2nd ed. W.H. Freeman and Co . San F r a n c i s c o . 859 pp. S o u t h , G .R . 1968 . I n t e r t i d a l mar ine a l gae from G a b r i o l a I s l a n d , B r i t i s h C o l u m b i a . S y e s i s 1 : 1 7 7 - 1 8 6 . S t r a h l e r , A .N . 1953 . R e v i s i o n of H o r t o n ' s q u a n t i t a t i v e f a c t o r s in e r o s i o n a l t e r r a i n . T r a n s . An. Geophy s . Un. 34 : 345. S v e d e l i u s , N. 1929 . The sea sona l a l t e r n a t i o n of g e n e r a t i o n s of Ce r am i um c o r t i c a t u l u m . Some a n t i c r i t i c a l r emark s . S v e n s k . B o t . T i d s s k r . 23 : 3 6 6 - 3 8 7 . T a t e w a k i , M. and L . P r o v a s o l i . 1964 . V i t a m i n r equ i r ement s of three s p e c i e s of An t i thamnion . B o t . M a r . 6 : 1 9 3 - 2 0 3 . T u r n e r , C . H . C . and L . V . E v a n s . 1977. P h y s i o l o g i c a l s t u d i e s on the r e l a t i o n s h i p between A s c o p h y I I um nodos um and P o I y s i phon i a I a n o s a . New Phy to l . 79 : 363 -371 . Widdowson, T .B . 1965 . A s u r ve y of the d i s t r i b u t i o n of i n t e r t i d a l a l gae a long a coas t t r a n s i t i o n a l in r e spec t to s a l i n i t y and t i d a l f a c t o r s . J . F i s h . Re s . Bd. Canada 22: 1 4 2 5 - 1 4 5 4 . Widdowson, T .B . 1973 . The mar ine a l gae of B r i t i s h Co lumbia and n o r t h e r n W a s h i n g t o n : r e v i s e d l i s t and k e y s . Par t I . Phaeophyceae (brown a l g a e ) . S y e s i s 6: 8 1 - 9 6 . Widdowson, T . B . 1974. The mar ine a l gae of B r i t i s h Co lumbia and n o r t h e r n W a s h i n g t o n : r e v i s e d l i s t and k e y s . Par t I I . Rhodophyceae ( red a l g a e ) . S y e s i s 7: 1 4 3 - 1 8 6 . W i l l i a m s , P .R . 1963. O b s e r v a t i o n s on two e p i p h y t i c members of the Phaeophyceae. B r . p h y c o l . B u l l . 2 : 3 9 0 - 3 9 1 . ( A b s t r a c t ) W i l s o n , L . 1977. Sea sona l d i s t r i b u t i o n of red a l g a l e p i p h y t e s on Cod i um f r a g i l e . J . P h y c o l . 13 ( s u p p l . ) : 74 . ( A b s t r a c t ) 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0098709/manifest

Comment

Related Items