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The biology of Nodularia Cyanophyceae Nordin, R. N. 1974

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THE BIOLOGY OF NODULARIA (CYANOPHYCEAE) by RICHARD NELS NORDIN B . S c , U n i v e r s i t y of Nor th Dakota , 1970 M o S c , U n i v e r s i t y of Nor th Dakota , 1971 A THESIS SUBMITTED IN THE REQUIREMENTS DOCTOR O F PARTIAL FULFILMENT OF FOR THE DEGREE OF PHILOSOPHY i n the Department of BOTANY We accept t h i s t h e s i s as conforming to the r e q u i r e d s tandard T H E U N I V E R S I T Y OF B R I T I S H C O L U M B I A MA Y , 1974 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an advanced degree a t the U n i v e r s i t y o f B r i t i s h C o l u m b i a , I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n ot be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f firTTfiKil The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada »ate Kacj 3^J93!± " T h e t a p e r s o f t h e g o d s , T h e s u n a n d m o o n , r u n down l i k e waxen g l o b e s ; T h e s h o o t i n g s t a r s a l l e n d i n p u r p l e j e l l i e s , A n d c h a o s i s a t h a n d . " D r y d e n ' s O e d i p u s , r e f e r r i n g t o t h e s u d d e n a p p e a r a n c e o f N o s t o c b a l l s a f t e r a r a i n f a l l . i i i Chairman: Professor J. Stein ABSTRACT The genus Nodularia /Mertens in JuergensJ Bornet et Flahault 1888 i s considered with regard to i t s ecology, distribution, physiology and taxonomy. Ecology was studied by periodic sampling of organisms and moni-toring environmental parameters in four saline ponds in the British Columbia interior, as well as by sampling a variety of brackish, marine and other inland saline water bodies. Nodularia was collected in habitats of alkaline pH 8.2 - 10.0) and medium to high dissolved salts (4 - 60 °/ ) and is associated with a specific group of eury-oo haline bluegreen algae. However, environments in which i t grows are usually dominated by species of green algae, with Nodularia only occas-ionally becoming dominant. The distribution of Nodularia was examined by reference to herbarium collections and literature reports. Although many bluegreen algae are considered cosmopolitan, Nodularia has a temperate and subtropical distribution with few reports from polar or tropical latitudes. The growth and physiology was investigated using 16 isolates from Canada, United States, Great Britain and Australia. Isolates were grown in a wide range of conditions of light, temperature, pH and dissolved s^lts in defined media. Generally, the highest growth rates were obtained with relatively high light intensity (550 - 600 f t - c ) , temperature of 25 - 30 C, pH near 10 and dissolved salts of 5 - 3 0 °/ 0 0- Highest rates of growth were obtained with NO3- as the nitrogen source as opposed to NH^+ or urea. Nodularia was found to fix nitrogen in pure culture. i v Nodular ia was es tab l i shed i n 1822 by Mertens, but not v a l i d i l y publ ished u n t i l 1888 by Bornet and F l a h a u l t . At present there have been 28 taxa d e s c r i b e d . The v a l i d i t y of these and the v a r i a b i l i t y i n nature and of i s o l a t e s i n c u l t u r e in a v a r i e t y of chemical and p h y s i c a l cond i t ions was cons idered . An eva luat ion of the r e l i a b i l i t y of the taxonomic c h a r a c t e r i s t i c s i n d i c a t e s that sheath and akinete c h a r a c t e r -i s t i c s are v a r i a b l e whi le vegetat ive c e l l shape, heterocyst l o c a t i o n and aspects of ak inete formation are more s tab le c h a r a c t e r i s t i c s . On the bas is of the observat ions repor ted , a l l the descr ibed taxa can be inc luded w i t h i n two s p e c i e s , N. spumigena ^Mertens in Juergens7 Bornet et F lahau l t 1888 or N. harveyana (Thw.) Thuret 1875, or p laced i n other genera. V T A B L E OF CONTENTS PAGE ABSTRACT i i i L I S T OF T A B L E S v i L I S T OF F I G U R E S v i i L I S T OF APPENDIX T A B L E S i x ACKNOWLEDGEMENTS x INTRODUCTION 1 ECOLOGY 3 M e t h o d s 4 R e s u l t s 9 D I S T R I B U T I O N 27 GROWTH AND PHYSIOLOGY 36 M a t e r i a l s a n d m e t h o d s 36 R e s u l t s 45 TAXONOMY 65 D I S C U S S I O N 105 E c o l o g y a n d P h y s i o l o g y 105 T a x o n o m y 113 L I T E R A T U R E C I T E D 118 APPENDIX T A B L E S I - X 138 v i L I S T OF T A B L E S T A B L E PAGE I. L o c a t i o n s o f N o d u l a r i a c o l l e c t i o n s , 7 1971-1973 I I . O r i g i n o f i s o l a t e s u s e d i n g r o w t h a n d 37 p h y s i o l o g y e x p e r i m e n t s I I I . L i g h t i n t e n s i t y u s e d i n c u l t u r e 42 e x p e r i m e n t s I V . M i c r o b i o l o g i c a l m e d i a f o r c h e c k i n g 44 p u r i t y o f c u l t u r e s V . C o m p a r a t i v e m o r p h o l o g y o f N o d u l a r i a 47 i n a x e n i c c u l t u r e , c o n t a m i n a t e d c u l t u r e a n d n a t u r e V i i L I S T OF F I G U R E S F I G U R E PAGE 1. S t u d y a r e a s i n t h e i n t e r i o r o f B r i t i s h C o l u m b i a 6 2 . S e a s o n a l c h a n g e s i n s a l i n e p o n d s 12 3. S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y a n d 14 o r g a n i s m s f o r 1972-1973 i n C t e n o c l a d u s P o n d 4. Some f e a t u r e s o f s a l i n e p o n d s 17 5. S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y a n d 20 o r g a n i s m s f o r 1972-1973 i n W a l l e n d e r L a k e 6. S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y a n d 21 o r g a n i s m s f o r 1972-1973 i n I n k s L a k e 7. S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y a n d 24 o r g a n i s m s f o r 1972-1973 i n W h i t e L a k e 8 . N o r t h A m e r i c a n d i s t r i b u t i o n o f N . s p u m i g e n a 30 9. N o r t h A m e r i c a n d i s t r i b u t i o n o f N . h a r v e y a n a 30 10. E u r o p e a n d i s t r i b u t i o n o f N . s p u m i g e n a 32 11. E u r o p e a n d i s t r i b u t i o n o f _N« h a r v e y a n a 32 12. W o r l d ( e x c l u s i v e o f N o r t h A m e r i c a a n d E u r o p e ) 34 d i s t r i b u t i o n o f N . s p u m i g e n a 13. W o r l d ( e x c l u s i v e o f N o r t h A m e r i c a a n d E u r o p e ) 34 d i s t r i b u t i o n o f N . h a r v e y a n a 14. S t a n d a r d c u r v e s o f o p t i c a l d e n s i t y v e r s u s 40 c e l l n u m b e r s 15. Q u a l i t y o f l i g h t u s e d i n c u l t u r e e x p e r i m e n t s 42 16. C o m p a r i s o n o f g r o w t h o f a x e n i c c u l t u r e s a n d 47 c u l t u r e s w i t h b a c t e r i a l c o n t a m i n a t i o n 17. E f f e c t o f t e m p e r a t u r e on i s o l a t e s o f N o d u l a r i a 49 1 8 . E f f e c t o f l i g h t i n t e n s i t y o n g r o w t h o f N o d u l a r i a 52 19. E f f e c t o f pH o n t h e g r o w t h o f N o d u l a r i a 54 v i i i F I G U R E PAGE 20. E f f e c t o f s a l i n i t y o n g r o w t h o f N o d u l a r i a 57 21. E f f e c t o f s a l i n i t i e s u s i n g s a l t s o t h e r t h a n 59 N a C l on g r o w t h o f N o d u l a r i a 22. E f f e c t o f n i t r o g e n s o u r c e o n t h e g r o w t h o f 61 N o d u l a r i a 23* E f f e c t o f s a l i n i t y , pH a n d t e m p e r a t u r e o n a c e t - 64 y l e n e r e d u c t i o n b y N o d u l a r i a 24. T h e s p e c i e s o f N o d u l a r i a d e s c r i b e d i n a n d s i n c e 69 t h e r e v i s i o n o f B o r n e t a n d F l a h a u l t (1888) 25. P o s s i b l e e x p l a n a t i o n f o r a k i n e t e c h a r a c t e r - 86 i s t i c s o f N . a r m o r i c a 26. C o m p a r i s o n o f t h e v a r i a b i l i t y o f s i z e o f v e g e t - 88 a t i v e c e l l s , h e t e r o c y s t s a n d a k i n e t e s o f two i s o l a t e s u n d e r a v a r i e t y o f c u l t u r e c o n d i t i o n s 27. C o m p a r i s o n o f c e l l s i z e s o f N . h a r v e y a n a a n d 91 N . s p h a e r o c a r p a 28. A k i n e t e v a r i a b i l i t y o f t h r e e B r i t i s h C o l u m b i a 94 i s o l a t e s a s r e l a t e d t o s a l i n i t y 29. C o m p a r i s o n o f c e l l s i z e s o f N_. s p u m i g e n a a n d 96 N . h a r v e y a n a 30. C u l t u r a l v a r i a t i o n o f N o d u l a r i a 99 31. T h e m o r p h o l o g i c a l v a r i a t i o n o f N . s p u m i g e n a 101 32. T h e m o r p h o l o g i c a l v a r i a t i o n o f N . h a r v e y a n a 103 i x LIST OF APPENDIX TABLES TABLE PAGE I Procedures used in water analysis 138 II Water chemistry for Ctenocladus Pond 139 III Water chemistry for Wallender Lake 140 TV Water chemistry for Inks Lake 141 V Water chemistry for White Lake 142 ; VI . Water chemistry for the lakes irregularly 143 ' sampled or sampled only once VII World distribution of Nodularia 144 VIII Medium BG-11 (Stanier et a l . 1 9 7 1 ) 157 IX Herbarium material of Nodularia examined 158 X Synonomy of Nodularia i * * X ACKNOWLEDGEMENTS I w o u l d l i k e t o t h a n k D r . J a n e t R . S t e i n f o r h e l p a n d g u i d a n c e d u r i n g t h e c o u r s e o f t h i s s t u d y , a n d t o members o f my c o m m i t t e e f o r t h e i r comments a n d c r i t i c i s m s o f t h e t h e s i s . Many p e r s o n s l e n t f r e e l y o f e q u i p m e n t , t i m e a n d a d v i c e a n d t o t h e m I am g r a t e f u l . T h e s t u d y was s u p p o r t e d b y f u n d s f r o m NRC A1035 t o J . R . S t e i n . My g r a t i t u d e t o C a r o l y n i s i n s u f f i c i e n t l y e x p r e s s e d h e r e . S h e p r o v i d e d h e l p i n t h e f i e l d , t y p e d , g l u e d , x e r o x e d , r e t y p e d , w o r r i e d , r a n e r r a n d s a n d a i d e d i n many o t h e r a s p e c t s o f t h e s t u d y a s w e l l a s s u p p l y i n g u n d e r s t a n d i n g a n d s m i l e s a t e v e n t h e m o s t d i f f i c u l t o f t i m e s . 1 INTRODUCTION Drotiet and D a i l y ' s (1956) r e v i s i o n of the c o c c o i d b l u e -green algae caused c o n s i d e r a b l e c o n t r o v e r s y r e g a r d i n g i n t e r -p r e t a t i o n of taxonomy and morphology of the Cyanophyceae. The v a r i a b i l i t y i n h e r e n t i n bluegreen a l g a l taxa has been su b j e c t to a v a r i e t y of i n t e r p r e t a t i o n s . ' Many Cyanophyceae have been d e s c r i b e d as new s p e c i e s with l i t t l e j u s t i f i c a t i o n ( G o l u b ic 1969) and every year more taxa are proposed to add to the taxonomic c o n f u s i o n . P h y s i o l o g i c a l and b i o c h e m i c a l c h a r a c t e r i z a t i o n are of l i t t l e value i f an i n c o r r e c t or unacceptable b i n o m i a l i s ap-p l i e d . A n a c y s t i s n i d u l a n s , a c o c c o i d form, i s a case i n p o i n t . The commonly s t u d i e d i s o l a t e of Kratz and A l l e n has been shown to be a Synechococcus sp. (Padmaja and Desikachary 1968; Komarek 1970). However A n a c y s t i s n i d u l a n s i s employed by r e s e a r c h e r s who are a p p a r e n t l y u n w i l l i n g to use the c o r r e c t b i n o m i a l . Taxonomic work i s of l i t t l e v a lue unless v a r i a t i o n both i n c u l t u r e and i n nature i s taken i n t o account. I t i s important that both f i e l d and l a b o r a t o r y s t u d i e s be co o r d i n a t e d and complementary to give i n s i g h t i n t o the b i o l o g y of an a l g a (Pringsheim 1967). Despite the acknowledged need f o r t h i s type of an approach f o r the bluegreen algae (Desikachary 1970), only one study (Kann and Komarek 1970), using e c o l o g i c a l o b s e r v a t i o n s , c u l t u r a l study and a c r i t i c a l review of the l i t e r a t u r e , h a s b e e n r e p o r t e d . I n l i g h t o f t h i s , t h e n e e d f o r s u c h s t u d i e s a r e a p p a r e n t . The p r e s e n t i n v e s t i g a t i o n o f t h e f i l a m e n t o u s , h e t e r o c y s t o u s g e n u s N o d u l a r i a / M e r t e n s i n J u e r g e n s V B o r n e t e t F l a h a u l t ( s e e S t a f l e u 1972 , p . 240) was u n d e r t a k e n to s t u d y t h e g e n u s and d e l i m i t t h e s p e c i e s . T h e g e n u s i s more m a n a g e a b l e t h a n t h e l a r g e g e n e r a , s u c h a s Nos t o e o r A n a b a e n a , and a p p e a r s w e l l -d e f i n e d i n a g e n e r i c s e n s e . The p r o b l e m was a p p r o a c h e d by s t u d y i n g t h e o c c u r r e n c e and d i s t r i b u t i o n , f a c t o r s c o n t r o l l i n g i t s g r o w t h , and i t s m o r p h o l o g i c a l v a r i a t i o n . T h e s e a r e c o n s i d e r e d a s t h r e e s e p a r a t e c h a p t e r s ; E c o l o g y ; P h y s i o l o g y ; Taxonomy 3 ECOLOGY Of p r i m a r y i m p o r t a n c e i n t h e s t u d y o f an o r g a n i s m i s d e f -i n i n g i t s r o l e i n t h e e n v i r o n m e n t : what o r g a n i s m s i t i s a s s o c -i a t e d w i t h , a n d when a n d why i t o c c u r s i n a p a r t i c u l a r p l a c e a t a p a r t i c u l a r t i m e . T h e p r i m a r y s a m p l i n g a r e a s w e r e i n t h e i n t e r i o r o f B r i t i s h C o l u m b i a n e a r K a m l o o p s a n d P e n t i c t o n . T h e s e a r e a s a r e i n t h e p o n d e r o s a p i n e - b u n c h g r a s s b i o g e o c l i m a t i c z o n e ( K r a j i n a I965) a n d t h e v e g e t a t i o n i n t h e a r e a o f t h e p o n d s a n d t h e l o w e r v a l l -e y s i s d o m i n a t e d b y A r t e m i s i a t r i d e n t a t a N u t t . B o t h a r e a s r e c e i v e r e l a t i v e l y l o w a m o u n t s o f p r e c i p i t a t i o n , m o s t o c c u r i n g i n t h e w i n t e r . K a m l o o p s h a s an a v e r a g e o f 24.3 c m / y r p r e c i p -i t a t i o n , w i t h m o s t snow i n D e c e m b e r o r J a n u a r y a n d m o s t r a i n i n J u n e . T h e a v e r a g e d a i l y maximum a n d minimum J u l y t e m p e r -a t u r e s a r e 29 C a n d 13 C a n d t h e a v e r a g e d a i l y maximum a n d min imum J a n u a r y t e m p e r a t u r e s a r e -2 C a n d -9 C . P e n t i c t o n r e c e i v e s 27.4 c m / y r a v e r a g e p r e c i p i t a t i o n , w i t h m o s t f a l l i n g a s snow i n D e c e m b e r a n d a s r a i n i n May o r J u n e . T h e a v e r -a g e d a i l y maximum a n d minimum J u l y t e m p e r a t u r e s a r e 29 C a n d 12 C a n d t h e a v e r a g e d a i l y maximum a n d min imum J a n u a r y t e m p -e r a t u r e s a r e 0 a n d -6 C ( C a n a d a D e p a r t m e n t o f T r a n s p o r t ) . P e r i o d i c s a m p l i n g , M a y - S e p t e m b e r 1972; A p r i l - J u l y 1973, was made o f f o u r p o n d s i n w h i c h N o d u l a r i a r e g u l a r l y o c c u r e d . 4 Three of these ponds (Ctenocladus Pond, Wal lender Lake , Inks Lake) are l o c a t e d southwest of Kamloops and one (White Lake) l o c a t e d southwest of P e n t i c t o n ( F i g . l ) . B l i n n (I969) has doc-umented some aspects of Ctenocladus Pond and Wal lender Lake . I n a d d i t i o n to these four h a b i t a t s , which were sampled p e r i o d -i c a l l y , a number of l o c a t i o n s were sampled on an i r r e g u l a r b a s i s or o n l y on a s i n g l e o c c a s i o n (Table 1 ) . Methods: At each sampl ing the temperature and d i s s o l v e d oxygen were measured (YSI model 54 oxygen me te r - the rmis to r and a g l a s s thermometer); pH recorded (Metrohm model E280A pH meter ) ; water ( d u p l i c a t e 500 ml samples ) , p lank ton and grab samples c o l l e c t e d and gene ra l o b s e r v a t i o n s made (water l e v e l s , spec ie s p resen t , and e s t i m a t i o n of abundance of organisms on a s u b j e c t -i v e b a s i s s i n c e attempts at q u a n t i t a t i v e measurements proved ext remely d i f f i c u l t ) . The water samples were f i l t e r e d through Whatman (#1) and M i l l i p o r e (.45 Um) f i l t e r s , w i t h i n s i x hours of c o l l e c t i o n and main ta ined a t 5 C f o r r e t u r n t o the l a b o r a t o r y . In the l a b o r a t o r y the organisms present were i d e n t i f i e d and N o d u l a r i a i s o l a t e d . Then the samples were f i x e d w i t h e i t h e r n e u t r a l 4% f o r m a l i n , L u g o l ' s i o d i n e or 70% e t h a n o l . Ana lyses of or tho-phosphate , ammonium-nitrogen and n i t r a t e n i t r o g e n were c a r r i e d ou t , w i t h i n 48 hours , a c c o r d i n g 5 Figure 1. Study areas i n the i n t e r i o r of B r i t i s h Columbia, scale 1:62,500 a. B r i t i s h Columbia 1 inch = approx. 650 km b. West of Kamloops B r i t i s h Columbia 1. Inks Lake 2. Wallender Lake 3« Bowers Lake 4 . Ironmask Lake 5« unnamed pond 6. Polygon Pond 7. Ctenocladus Pond 8. 'Salt Pond 4» 9. Salsola Pond 10. 'Salt Pond 3' 11. Saltwort Pond c. South west of Penticton B r i t i s h Columbia 12. White Lake 13. Mahoney Lake 14. Green Lake 7 T a b l e 1. L o c a t i o n s of N o d u l a r i a c o l l e c t i o n s , 1 9 7 1 - 1 9 7 3 . A. L o c a t i o n s r e g u l a r l y sampled 1. Ctenocladus Pond 50°39'N, 120°32'W North o f Trans-Canada Highway (BC Hwy 97), 9 mi west of Kamloops B.C. C i t e d by B l i n n ( 1 9 6 9 ) as 'Cherry Creek Pond' 2. Wallender Lake 5 0 ° 3 8 ' N , 120°26 .5'W East s i d e of Lac l a Jeune Road, 3 mi from i t s j u n c -t i o n with t h e Trans-Canada Highway (BC Hwy 97) 3 . Inks Lake 50°37'N , 120°27'W West s i d e of Lac l a Jeune Road, 4 mi from i t s j u n c -t i o n w i t h t h e Trans-Canada Highway (BC Hwy 97) 4. White Lake 49°19'N , 119°38'W One m i l e south west of the Dominion Radio A s t r o -p h y s i c a l Observatory and 5 mi south of P e n t i c t o n B.C. B. L o c a t i o n s i r r e g u l a r l y or s i n g l y sampled 5. ' S a l t pond 4' 50°40'N , 120°35'W A s m a l l unnamed pond southwest of S a l s o l a Pond ( S a l s o l a Pond c i t e d by B l i n n as S a l t Mine Pond 1) 6. ' S a l t pond 3' 50°40'N , 120°34'W A s m a l l unnamed temporary pond between S a l s o l a Pond and S a l t w o r t Pond ( S a l t w o r t Pond c i t e d by B l i n n I969 as S a l t Mine Pond 2) 7 . B i g Q u i l l Lake, Saskatchewan 51°30'N , 104°40'W North of Kandahar on Hwy 14 (Sask.) and 100 mi east of Saskatoon 8. D e v i l s Lake, North Dakota at Camp G r a f t o n 48°N , 99°W 9. San de Fuca, Washington 48°10'N , 122°48'W Adjacent t o Washington Highway 525 10. R i e f e l W i l d l i f e Refuge 4 9 ° 0 8'N , 123°10'W Near Ladner B.C. Continued... 8 T a b l e 1 (Contd.) Locations of Nodularia c o l l e c t i o n s , 1971-1973 . C u l t u r e s were i s o l a t e d from s o i l samples c o l l e c t e d at the f o l l o w i n g l o c a t i o n s 1 1 . Spotted Lake 49°05'N , 1 19°34»W In R i c h t e r Pass, 6 mi northwest of Osoyoos B.C. on B.C. Hwy 3 1 2 . Small unnamed pond 1 1 9 ° 3 8 ' N , 49°09'W In low area 1 mi south of R i c h t e r Lake, 4 mi southwest of Spotted Lake adjacent t o Hwy 3 1 3 . Bowers Lake 50°40'N , 120°26'W At the j u n c t i o n of Trans-Canada Highway (Canada Hwy 1 , BC Hwy 97) and Lac l a Jeune Road 14. Water hole Simpson's Gap, Northern T e r r i t o r i e s , A u s t r a l i a 20°45'S , 134'E C o l l e c t e d by Dr. J.R. S t e i n , 18 i x I969 9 to Standard Methods (A.P.H.A. I965). The one exception to t h i s treatment of water samples i s to be found i n the data from August 1972 when the water was not analyzed u n t i l two weeks afte r c o l l e c t i o n . Analyses of the water f o r the major cations (Na , Mg , Ca , K ) were made with a Perkin-Elmer (model 303) atomic absorption spectrophotometer and fo r major anions (C0^} HCO^, SO^, Cl ) by Standard Methods from water samples stored at 5 C. D e t a i l s of water analysis are given as Appendix, Table 1. When reference i s made to s a l i n i t y i t i s i n the broad sense of the amount of dissolved s a l t contained i n the water without any implied i o n i c content rather than i n the narrower oceanographic sense of c h l o r i n i t y . The term brackish i s used i n reference to d i l u t e sea water, and the term s a l i n e refers to inland water bodies of s a l t content greater than 0.5 °/oo. Results: In the spring (late March and Ap r i l ) the lakes are at t h e i r largest volume and contain r e l a t i v e l y low amounts of dissolved s a l t s due to the high volume of water r e s u l t i n g from snow-melt and drainage into these depressions. As spring and summer progress, the s a l i n i t y increases as water volume decreases due to evaporation i n excess of r a i n f a l l and 10 r u n o f f . I n l a t e summer a n d f a l l ( l a t e A u g u s t a n d S e p t e m b e r ) t h e p o n d s r e a c h t h e i r min imum v o l u m e a n d i n some i n s t a n c e s d r y u p c o m p l e t e l y ( F i g 2) . T h e d y n a m i c s o f t h e i o n s i n s a l i n e p o n d s o f t h i s t y p e i s d e s c r i b e d b y B l i n n (1971)* I n a n y i n t e r p r e t a t i o n i t m u s t b e e m p h a s i s e d t h a t t h e r e c a n b e a g r e a t d e a l o f v a r i a t i o n f r o m y e a r t o y e a r i n w a t e r l e v e l s o f t h e s e p o n d s d e p e n d i n g o n t o t a l p r e c i p i t a t i o n . F o r e x a m p l e , i t was o b s e r v e d t h a t w a t e r l e v e l s i n 1973 w e r e l o w e r t h a n 1972 a t c o r r e s p o n d i n g c o l l e c -t i o n d a t e s . T h e p o n d s a r e c h a r a c t e r i z e d b y a l a r g e amount o f a u t o t -r o p h i c g r o w t h , p r i n c i p a l l y a l g a e , i n s p r i n g a n d e a r l y summer ( A p r i l - J u l y ) f o l l o w e d b y a h e t e r o t r o p h i c g r o w t h p h a s e , g r a z i n g i n v e r t e b r a t e s a n d b a c t e r i a , f r o m J u l y t o S e p t e m b e r . Maximum d i v e r s i t y o f o r g a n i s m s o c c u r s i n s p r i n g a n d d i v e r s i t y d e c r e a -s e s a s t h e s a l i n i t y i n c r e a s e s . T o e x a m i n e t h e s e t r e n d s a n d v a r i a t i o n s s i n g l e p o n d s must b e c o n s i d e r e d . A . L o c a t i o n s r e g u l a r l y s a m p l e d . 1. C t e n o c l a d u s P o n d h a s N a + a n d S O . a s t h e d o m i n a n t 4 i o n s . T h e p o n d n o r m a l l y d r i e s u p d u r i n g t h e c o u r s e o f t h e summer a n d t h e s a l i n i t y may v a r y f r o m 30-50 ° / o o i n t h e s p r i n g t o m o r e t h a n 400 ° / o o i n t h e f a l l ( A p p e n d i x , T a b l e I I ) . T h e d o m i n a n t a l g a i s t h e c h l o r o p h y t e C t e n o c l a d u s c i r c i n n a t u s B o r z i . 11 Figure 2. Seasonal changes i n sa l i n e ponds a. White Lake, 2 v i 1973 b. White Lake, 22 v i i 1973 c. Ctenocladus Pond, 3 v i 1973 d. Ctenocladus Pond, 21 v i i 1973 1 3 N o d u l a r i a h a r v e y a n a was c o l l e c t e d o n l y d u r i n g A p r i l a n d May i n s m a l l n u m b e r s a n d was a b s e n t f r o m J u n e - S e p t e m b e r . O t h e r a l g a e p r e s e n t w e r e t h e g r e e n a l g a e S p i r o g y r a s p . , U l o t h r i x  t e n e r r i m a K u t z . , U l o t h r i x s p . , D u n a l i e l l a s a l i n a ( D u n a l ) T e o d o r . a n d C l a d o p h o r a f r a c t a ( D i l l w . ) K u t z ; t h e b l u e g r e e n a l g a e O s c i l l a t o r i a a m p h i b i a C . A . A g . , 0 . b r e v i s C . A . A g . , A n a b a e n a s p . , P h o r m i d i u m t e n u e ( M e n e g h . ) Gomont a n d L y n g b y a  n o r d g a a r d i i W i l l e ; an E u g l e n a s p . ; t h e d i a t o m s A m p h o r a  c o f f e a e f o r m i s A g . a n d N a v i c u l a s p . A l l b u t t h e D u n a l i e l l a a n d L y n g b y a w e r e p r e s e n t i n t h e i r l a r g e s t n u m b e r s i n A p r i l a n d M a y . W i t h i n c r e a s i n g s a l i n i t y d u r i n g t h e summer , m o s t s p e c i e s o f a l g a e w e r e g r e a t l y r e d u c e d i n n u m b e r s o r d i s a p p e a r -e d c o m p l e t e l y . D u n a l i e l l a a n d L y n g b y a r e a c h e d maximum n u m b e r s i n J u n e o r J u l y , t h e f o r m e r i n t h e i s o l a t e d p o c k e t s o f o p e n w a t e r , t h e l a t t e r a s a n e p i p h y t e o n d e a d o r d y i n g C l a d o p h o r a a n d U l o t h r i x . C t e n o c l a d u s r e m a i n e d t h r o u g h o u t t h e summer b u t i n l e s s q u a n t i t y a n d h i d d e n u n d e r t h e s a l t l a y e r d e p o s i t e d b y t h e e v a p o r a t i n g w a t e r . A r o u n d t h e s h o r e l i n e o f t h e p o n d i s a z o n e o f S a l i c o r n i a  r u b r a N e l s . a n d w i t h i n t h e p o n d i t s e l f i s a d e n s e g r o w t h o f R u p p i a m a r i t i m a L . I n A p r i l a n d May l a r g e n u m b e r s o f m o s -q u i t o l a r v a e ( A e d e s s p . ) w e r e o b s e r v e d , J n J u n e t h r o u g h A u g u s t t h e p o n d i s i n h a b i t e d b y l a r g e n u m b e r s o f b r i n e s h r i m p , A r t e m i a s a l i n a L e a c h . A mat o f p u r p l e s u l p h u r b a c t e r i a 14 30 .25 • u 20 a s 15 » 10 5 0 400 r o o CO 300 200 100 0 f c w e w •rl c bo c o •p c ca c E o o Ctenocladus  Nodularia Aedes Artemia D u n a l i e l l a 1972 APR Figure 3 . MAY JUN JUL AUG SEP 1973 APR MAY JUN JUL AUG Seasonal changes i n temperature, s a l i n i t y and organisms f o r 1972-1973 i n Ctenocladus Pond. The r e l a t i v e abundance of the organisms are indicated by the v e r t i c a l bars 15 c o v e r s a t h i c k , b l a c k , a n e r o b i c mud . An u n i d e n t i f i e d o s t r a -c o d a n d t h e r o t i f e r , B r a c h i o n u s p l i c a t i l i s M u l l e r , i n h a b i t t h e p o n d a n d e g g c a s e s , l a r v a e a n d a d u l t s a l t f l i e s ( E p h e d r i d a e ) a r e a r o u n d t h e w a t e r ' s e d g e . T h e p e r i o d i c i t y a n d p o p u l a t i o n d y n a m i c s o f t h e m o r e i m p o r t a n t o r g a n i s m s i s shown i n F ig.3« 2. W a l l e n d e r L a k e i s c h a r a c t e r i z e d b y a l o w e r amount o f d i s s o l v e d s a l t s a n d l e s s v a r i a b l e s a l i n i t y r a n g e ( l 4 - 8 8 ° / o o ) + -H-t h a n C t e n o c l a d u s P o n d . T h e p r i n c i p a l i o n s a r e Na . Mg , a n d SO^ ( A p p e n d i x , T a b l e I I I ) . P r o b a b l y a s a r e s u l t o f t h e s e l o w e r l e v e l s o f d i s s o l v e d s a l t , a l a r g e r v a r i e t y o f o r g a n i s m s a r e p r e s e n t . T h e v a s c u l a r p l a n t s c o n s i s t o f S a l -i c o r n i a r u b r a a n d S c i r p u s m a r i t i m u s L . a r o u n d t h e m a r g i n o f t h e l a k e a n d R u p p i a w i t h i n t h e l a k e i t s e l f . T h e d o m i n a n t a l g a t h r o u g h o u t t h e s e a s o n e x c e p t f o r e a r l y s p r i n g ( A p r i l a n d e a r l y M a y ) , i s C l a d o p h o r a f r a c t a . D u r i n g May r a p i d g r o w t h o f C l a d o p h o r a r e s u l t s i n t h e f o r m a t i o n o f a c h a r a c t e r i s t i c p e r i p h e r a l mat ( F i g . 4 c ) . N o d u l a r i a ( p r i m a r i l y N . h a r v e y a n a ) i s p r e s e n t i n maximum n u m b e r s i n May a n d J u n e a n d t y p i c a l l y o c c u r s a s s o c i a t e d w i t h t h e C l a d o p h o r a m a t . D u r i n g M a y , N o d u l -a r i a i s p r i m a r i l y v e g e t a t i v e , b y J u n e many o f t h e c e l l s h a v e c o n v e r t e d t o a k i n e t e s a n d b y J u l y v e r y f e w f i l a m e n t s a r e s e e n , w i t h n e a r l y a l l c e l l s a s a k i n e t e s . A s s o c i a t e d w i t h N o d u l a r i a i s a c o m m u n i t y o f b l u e g r e e n a l g a e c o n s i s t i n g o f O s c i l l a t o r i a  g e m i n a t a M e n e g h . , 0_. t e n u i s C . A . A g . , 0_. n i g r o v i r i d u s T h w a i t e s , 16 F i g u r e 4» Some f e a t u r e s o f t h e s a l i n e p o n d s s t u d i e d . a . W a l l e n d e r L a k e w i t h l a r g e n u m b e r s o f d i a t o m s a l o n g t h e s h o r e l i n e ( i v 1 9 7 3 ) b . A c l o s e r v i e w o f d i a t o m b l o o m i n a . ( p e n c i l i s a p p r -o x i m a t e l y 1 0 cm l o n g ) c . P e r i p h e r a l C l a d o p h o r a mat a r o u n d t h e e d g e o f W a l l -e n d e r L a k e ( v i 1 9 7 3 ) d . S h o r e l i n e o f W a l l e n d e r L a k e w i t h C l a d o p h o r a , p u r p l e s u l p h u r b a c t e r i a a n d S c i r p u s ( v i 1 9 7 3 ) e . I n k s L a k e s h o w i n g d e n s i t y o f b r i n e s h r i m p ( v i i 1 9 7 3 ) . A l s o p r e s e n t a r e C l a d o p h o r a a n d R u p p i a ( t h e r m o m e t e r i s a p p r o x i m a t e l y 35 cm l o n g ) 18 S p i r u l i n a m a j o r K u t z . a n d P h o r m i d i u m f r a g i l e G a m o n t . L y n g b y a  n o r d g a a r d i i a p p e a r s i n J u l y a f t e r t h e d e c l i n e i n n u m b e r s o f m o s t o t h e r b l u e g r e e n a l g a e . D u r i n g e a r l y s p r i n g ( A p r i l ) b e f o r e t h e g r o w t h o f t h e C l a d o p h o r a m a t , a d i a t o m maximum o c c u r s . I n A p r i l 1973 a r e m a r k a b l e b l o o m o f d i a t o m s o c c u r r e d ( p r i n c i p a l l y A m p h o r a  c o f f e a e f o r m i s a n d N a v i c u l a s p p . ) ( F i g . 4 a , b ) . T h e s e d i a t o m s a r e p r e s e n t a t o t h e r t i m e s o f t h e y e a r b u t i n much r e d u c e d n u m b e r s . T h e d i a t o m maximum i s f o l l o w e d b y r a p i d g r o w t h o f C l a d o p h o r a a n d t h e b l u e g r e e n a l g a e a s s o c i a t e d w i t h N o d u l a r i a a n d two U l o t h r i x s p p . , U . t e n e r r i m a a n d U . ? a e q u a l i s K u t z . I n J u l y 1972 a l a r g e number o f C e r a t i u m h i r u n d i n e l l a ( O . F . M . ) D u j . was o b s e r v e d b u t n o t i n 1973. I t s a p p e a r a n c e i n s u c h s a l i n e (48 ° / o o ) w a t e r s i s i n t e r e s t i n g s i n c e i t i s n o r m a l l y f o u n d i n much l e s s s a l i n e c o n d i t i o n s . A t t h e same t i m e ( J u l y 1972) i n n e a r b y I n k s L a k e ( s e e b e l o w ) a l a r g e number o f C e r a t i u m were a l s o p r e s e n t . A s p e c i e s o f E u g l e n a was f o u n d i n l o c a l i z e d s m a l l a r e a s o n t h e s h o r e l i n e mud t h r o u g h o u t s p r i n g a n d summer . T h e a n i m a l c o m m u n i t y o f t h e l a k e i s c o m p o s e d o f a M o i n a s p . ( C l a d o c e r a ) , D i a p t o m u s s p . ( C o p e p o d a ) , B r a c h i o n u s p l i c a t ; ! . l i s ( R o t a t o r i a ) a n d C h i r o n o m i d a e , a l l p r e s e n t p r i m a r i l y i n t h e s p r i n g . A r t e m i a s a l i n a ( B r a n c h i o p o d a ) i s p r e s e n t i n l a r g e n u m b e r s i n l a t e s p r i n g a n d summer ( F i g . 4e) . E p h i d r i d a e 19 a r e p r e s e n t a r o u n d t h e e d g e s o f t h e l a k e a n d t h e b o t t o m s e d -i m e n t i s c o v e r e d b y a mat o f p u r p l e s u l p h u r b a c t e r i a ( F i g . 4 d ) . T h e p e r i o d i c i t y a n d d y n a m i c s o f o r g a n i s m s i s shown a s F i g . 5 . T h e maximum v a r i e t y o f s p e c i e s o c c u r s i n t h e s p r i n g o r e a r l y summer a n d d i v e r s i t y d e c r e a s e s i n r e s p o n s e t o i n c r e a s i n g s a l -i n i t y . B y l a t e summer t h e e x t r e m e s a l i n i t y r e d u c e s t h e number o f s p e c i e s t o a m i n i m u m . 3. I n k s L a k e i s s i m i l a r t o W a l l e n d e r L a k e b u t d i f f e r s f r o m W a l l e n d e r L a k e i n some d e t a i l s . I n k s L a k e h a s s l i g h t l y l e s s d i s s o l v e d s a l t s a n d a s l i g h t l y h i g h e r pH ( A p p e n d i x , T a b l e I V ) b u t o t h e r w i s e h a s s i m i l a r p l a n t a n d a n i m a l commun-i t i e s ( F i g . 6 ) . T h e same d i s t i n c t i v e p e r i p h e r a l C l a d o p h o r a mat a n d t h e same p l a n t a n d a n i m a l c o m p o n e n t s ( p l u s S c i r p u s  v a l i d u s V a h l . a n d t h e c y a n o p h y t e A n a b a e n a v a r i a b i l i s K u t z . ) a r e p r e s e n t . B o t h N . h a r v e y a n a a n d N . s p u m i g e n a a r e p r e s e n t i n t h e s p r i n g i n l a r g e n u m b e r s . T h e same p a t t e r n o f o c c u r -e n c e i s n o t e d w i t h g r e a t e s t g r o w t h i n s p r i n g a n d d i s a p p e a r a n c e o f N o d u l a r i a w i t h t h e w a t e r b e c o m i n g i n c r e a s i n g l y s a l i n e . I n b o t h I n k s L a k e a n d W a l l e n d e r L a k e m o s t N o d u l a r i a d i s a p p e a r s a t s a l i n i t i e s o f a p p r o x i m a t e l y 60 ° / o o , t h e d i s a p p e a r a n c e e v i d e n t l y i s r e l a t e d t o s a l i n i t y r a t h e r t h a n pH o r i o n i c c h a n g e s . E v i d e n c e i s p r e s e n t e d i n t h e s e c t i o n o n g r o w t h a n d p h y s i o l o g y t o f u r t h e r s u p p o r t t h i s . 4. W h i t e L a k e i s 100 m i l e s s o u t h e a s t o f t h e o t h e r t h r e e 20 30 — • • ' ' . f I L- , i_ 1972 MAY JUN JUL AUG SEP APR MAY JUN F i g u r e 5« S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y -o r g a n i s m s f o r 1 9 7 2 - 1 9 7 3 i n W a l l e n d e r L a k e 21 Cladophora Diaptomus Moina L ZZ] | • ' • * • . / / , , , ,_ 1 9 72 m Y JUN JUL AUG SEP 1 9 7 3 APR MAY JUN JUL AUG F i g u r e 6. S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y a n d o r g a n i s m s f o r 1972-1973 i n I n k s L a k e . 22 l a k e s . T h e d o m i n a n t i o n s a r e N a + , SO a n d CO ; t h e s a l -'r } si i n i t y v a r i e s f r o m 1 5 - 4 0 ° / o o a n d t h e pH i s 9 . 4 - 1 0 . 0 ( A p p e n d i x , T a b l e V ) . T h e o r g a n i s m s o f W h i t e L a k e a r e n o t a b l y d i f f e r e n t f r o m t h e p r e v i o u s t h r e e l a k e s . T h e r e i s no p e r i p h e r a l mat o f a l g a e a t t h e s h o r e l i n e a n d no p u r p l e s u l p h u r b a c t e r i a mat a t t h e m u d - w a t e r i n t e r f a c e a s c h a r a c t e r i z e s t h e o t h e r t h r e e l a k e s . A r o u n d t h e m a r g i n o f t h e l a k e i s D i s t i c h l i s s t r i c t a ( T o r r . ) T y d b . a n d S c i r p u s n e v a d e n s i s W a t s . , b o t h a t t i m e s b e i n g e m e r g e n t f r o m t h e w a t e r b o d y i t s e l f . N o d u l a r i a ( b o t h N. h a r v e y a n a a n d N . s p u m i g e n a ) o c c u r s o n a n d a r o u n d t h e b a s e s o f t h e e m e r g e n t v a s c u l a r p l a n t s a n d i n h o o f p r i n t s i n a c a t t l e c o r r a l l i n g a r e a o n t h e s o u t h s i d e o f t h e l a k e . N o d u l a r i a o c c u r s i n W h i t e L a k e i n s p r i n g t o e a r l y summer ( M a y - J u n e ) a n d i s g e n e r a l l y a b s e n t t h e r e s t o f t h e y e a r . H o w e v e r t h e s a l i n -i t y , w h i c h h a s a n a r r o w r a n g e o f f l u c t u a t i o n a n d r e l a t i v e l y l o w c o n c e n t r a t i o n , d o e s n o t a p p e a r t o b e t h e l i m i t i n g f a c t o r . T h e f a c t o r r e s p o n s i b l e f o r t h e p a t t e r n o f g r o w t h o f N o d u l a r i a i s t h e d e s t r u c t i o n o f h a b i t a t a s t h e w a t e r l i n e r e c e d e s b e y o n d t h e b a s e s o f t h e v a s c u l a r p l a n t s . T h e p r e s e n c e o f N o d u l a r i a , a s w e l l a s m o s t o t h e r a l g a e , d e p e n d s o n t h e l e v e l o f t h e w a t e r b e i n g a b o v e t h e l o w e s t v a s c u l a r p l a n t s . T h e o t h e r a l g a e p r e s e n t w i t h N o d u l a r i a a r e t h e b l u e g r e e n a l g a e O s c i l l a t o r i a  t e n u i s , 0 . c h a l b e a M e r t e n s , P h o r m i d i u m r e t z i ( C . A . A g . ) G o m o n t , A n a b a e n a s p . , L y n g b y a a e s t u a r i i ( M e r t e n s ) L i e b m a n n a n d S p i r u l i n a 23 major ; a n d t h e g r e e n a l g a e S p i r o g y r a s p . a n d C t e n o c l a d u s c i r -c i n n a t u s . T h e p r e c e d i n g b l u e g r e e n a l g a e show t h e same p e r -i o d i c i t y a n d l i m i t a t i o n s a s N o d u l a r i a . T h e S p i r o g y r a s p . i s p r e s e n t o n l y i n A p r i l a n d May a n d t h e C t e n o c l a d u s i n June or J u l y . In A p r i l 1973 t h e l a k e c o n t a i n e d l a r g e n u m b e r s o f s a l a m -a n d e r s (Ambystoma s p . ) . T h e p l a n k t o n o r g a n i s m s c o n s i s t o f t h e d i a t o m s A m p h o r a c o f f e a e f o r m i s a n d N a v i c u l a s p p . ; t h e c l a d -o c e r a n M o i n a s p . a n d a c o p e p o d o f t h e g e n u s D i a p t o m u s . I n S e p t e m b e r 1972 a l a r g e amount o f N o s t o c s p . was f l o a t i n g a n d w a s h e d o n s h o r e . T h e p e r i o d i c i t y a n d p o p u l a t i o n d y n a m i c s o f W h i t e L a k e i s shown a s F ig.7« B . L o c a t i o n s i r r e g u l a r l y s a m p l e d o r s a m p l e d o n l y o n c e . 5. ' S a l t P o n d 4' c o n t a i n e d N o d u l a r i a s p u m i g e n a i n May 1972 a t a s a l i n i t y o f 47 ° / o o , w i t h t h e p r i m a r y i o n s b e i n g Na +, S O ^ a n d CO^ ( A p p e n d i x , T a b l e V I ) . T h e d o m i n a n t a l g a o f t h i s p o n d was C t e n o c l a d u s c i r c i n n a t u s w i t h l e s s e r a m o u n t s o f O s c i l l a t o r i a b r e v i s , 0_. t e n u i s a n d A n a b a e n a s p . 6. ' S a l t P o n d 3' c o n t a i n e d N . s p u m i g e n a a s t h e d o m i n a n t alga i n May 1972 a t a s a l i n i t y o f 4 ° / o o ( N a + , S C ^ , C0~) ( A p p e n d i x , T a b l e V I ) . 7. B i g Q u i l l L a k e , S a s k a t c h e w a n , i n A u g u s t 1972, h a d a s a l i n i t y o f 26 ° / o o o f p r e d o m i n a n t l y N a + a n d SO^ ( A p p e n d i x , 24 a B 9 EH 30 25 20 15 10 5 0 o o 50 •P •rl e •H H CO CO 25 N. spumigena e 00 •H C «J . bs O •P c ca e •rt a o o N. harveyana Lyngbya Diaptomus Moina Nostoc J L. 1972 MAY JUN JUL AUG SEP 1973 F i g u r e 7 . S e a s o n a l c h a n g e s i n t e m p e r a t u r e , s a l i n i t y -o r g a n i s m s f o r 1 9 7 2 - 1 9 7 3 i n W h i t e L a k e . 25 T a b l e V I ) . N o d u l a r i a s p u m i g e n a was t h e p r i n c i p a l p h y t o p l a n k -t e r . A s s o c i a t e d w i t h i t was t h e d i a t o m C h a e t o c e r a s e l m o r e i B o y e r a n d l a r g e n u m b e r s o f w a t e r b o a t m e n ( C o r i x i d a e ) a n d c o p -s p . w h i c h g r e w a t t a c h e d t o t h e b o t t o m w i t h l a r g e n u m b e r s o f d e t a c h e d p l a n t s b l o w n u p o n t h e l e e s h o r e . 8 . D e v i l s L a k e , N o r t h D a k o t a , h a d a s a l i n i t y o f 1 2 ° / o o ( N a + , S 0 4 ) i n A u g u s t 1972 ( A p p e n d i x , T a b l e V I ) . N o d u l a r i a  s p u m i g e n a was c o l l e c t e d b o t h f r o m o p e n w a t e r w h e r e i t was a s s o c i a t e d w i t h A p h a n i z o m e n o n f l o s - a q u a e M o r r e n a n d M i c r o c y s t i s  a e r u g i n o s a K u t z . , a n d a l o n g t h e s h o r e l i n e a s s o c i a t e d w i t h C l a d o p h o r a s p . a n d E n t e r o m o r p h a s p . 9 . N e a r S a n d e F u c a , W a s h i n g t o n , i n May 1973 N o d u l a r i a  h a r v e y a n a was c o l l e c t e d f r o m a b r a c k i s h ( N a + , C l ) p o n d o f s a l i n i t y 1 8 ° / o o ( A p p e n d i x , T a b l e V I ) . A s s o c i a t e d w i t h t h e N o d u l a r i a was P e r c u r s a r i a p e r c u r s a ( C . A g . ) R o s e n v i n g e a n d L y n g b y a a e s t u a r i i . 1 0 . R i e f e l W i l d l i f e R e f u g e - a p o n d n e a r t h e r e f u g e o f f i c e h a d a s a l i n i t y o f 8 . 4 ° / o o ( N a + , C l ) ( A p p e n d i x , T a b l e V I ) a n d c o n t a i n e d N o d u l a r i a h a r v e y a n a . T h e d o m i n a n t a l g a was a H u g e o t i a s p . , w i t h L y n g b y a a e s t u a r i i a n d an O s c i l l a t o r i a s p . a l s o b e i n g p r e s e n t . e p o d s ( D i a p t o m u s s p . ) . T h e d o m i n a n t a l g a was a n E n t e r o m o r p h a C . I s o l a t e s f r o m s o i l ( S e e a l s o T a b l e I ) B l i n n ( 1 9 6 9 ) r e p o r t s N . s p u m i g e n a f r o m W a l l e n d e r L a k e , 26 C t e n o c l a d u s P o n d , B o w e r s L a k e , P o l y g o n P o n d , I r o n m a s k L a k e , S a l s o l a P o n d a n d S a l t w o r t P o n d . I s o l a t e s w e r e o b t a i n e d f r o m W a l l e n d e r L a k e a n d C t e n o c l a d u s P o n d . B o w e r s L a k e was d r y f r o m 1971-1973^ h o w e v e r m i c r o s c o p i c e x a m i n a t i o n o f s o i l a n d s a l t f r o m t h e d r y l a k e b e d r e v e a l e d a k i n e t e s . E n r i c h m e n t t e c h n i q u e s ( A l l e n 1973) r e s u l t e d i n an i s o l a t e o f N_. s p u m i g e n a . S i m i l a r o b s e r v a t i o n s a n d t e c h n i q u e s r e s u l t e d i n i s o l a t e s f r o m S p o t t e d L a k e a n d a s m a l l p o n d s o u t h o f R i c h t e r L a k e . E n r i c h -ment o f a n A u s t r a l i a n s o i l s a m p l e c o l l e c t e d n e a r A l i c e S p r i n g s a l s o y i e l d e d a n i s o l a t e o f N . s p u m i g e n a . N o d u l a r i a was n e v e r c o l l e c t e d f r o m P o l y g o n P o n d , I r o n m a s k L a k e , S a l s o l a P o n d o r S a l t w o r t P o n d , p o s s i b l y b e c a u s e when c o l l e c t i o n s w e r e made t h e s a l i n i t y was t o o h i g h ( g r e a t e r t h a n 100 ° / o o ) f o r N o d u l a r i a t o be g r o w i n g . No a k i n e t e s w e r e v i s -i b l e i n t h e s o i l f r o m t h e s e s i t e s a n d e n r i c h m e n t p r o c e d u r e s f a i l e d t o p r o d u c e a n y i s o l a t e s . D I S T R I B U T I O N Some a u t h o r s ( F r i t s c h 1945) r e g a r d m o s t b l u e g r e e n a l g a e a s b e i n g c o s m o p o l i t a n i n d i s t r i b u t i o n . H o w e v e r , c l o s e r e x a m -i n a t i o n o f i n d i v i d u a l g e n e r a o r s p e c i e s may r e v e a l v e r y d i s -c r e t e d i s t r i b u t i o n . N o d u l a r i a i l l u s t r a t e s t h i s p o i n t . G i e t l e r (1932) a n d L i n d s t e d t (1943) d e s c r i b e t h e d i s t r i b u t i o n o f N . s p u m i g e n a a s c o s m o p o l i t a n . Chapman (1955) d e s c r i b e s t h e d i s t r i b u t i o n o f N . h a r v e y a n a a s c o s m o p o l i t a n " a n d U m e z a k i (I96I) d e s c r i b e s N . h a r v e y a n a a s " s u b c o s m o p o l i t a n " w i t h o u t a n y e x p l a n a t i o n o f t h e p r e f i x . H o w e v e r , i n a g e n e r a l s e n s e b l u e g r e e n a l g a e a r e v e r y r e s t r i c t e d . C y a n o p h y t e s a r e r a r e l y f o u n d i n a c i d s i t u a t i o n s a n d n e v e r b e l o w pH 4 ( B r o c k 1973)• Dawson (I966) s t a t e s t h a t many m a r i n e s p e c i e s may b e d i f f e r e n t i a t e d i n t o t e m p e r a t e a n d t r o p i c a l . I n a s p e c i f i c s e n s e many b l u e g r e e n a l g a e a r e f o u n d i n r e s t r i c t e d h a b i t a t s (some h o t s p r i n g s p e c i e s o r t r o p i c a l m a r i n e p l a n k t o n s p e c i e s ) . O t h e r s h a v e m o r e s u b t l e r e q u i r e -m e n t s , t o l e r a n c e s o r p r e f e r e n c e s . T h e n a t u r e o f t h e s e d i s t r i b -u t i o n p a t t e r n s i s o b s c u r e d b y m i s i d e n t i f i c a t i o n , l a c k o f p e r t -i n e n t d a t a a n d more p a r t i c u l a r l y t h e c o n f u s e d t a x o n o m i c s i t -u a t i o n . T h e d i s t r i b u t i o n o f N o d u l a r i a was s t u d i e d b y a c r i t i c a l r e v i e w o f t h e l i t e r a t u r e a n d e x a m i n a t i o n o f h e r b a r i a m a t e r i a l 28 ( A p p e n d i x , T a b l e I X ) . T h e r e s u l t s o f t h i s s u r v e y show t h a t t h e g r e a t m a j o r i t y o f c o l l e c t i o n s w e r e i n t e m p e r a t e a n d s u b -t r o p i c a l a r e a s ( i . e . 25°-60° n o r t h a n d s o u t h o f t h e e q u a t o r , F i g s . 8-13). I t may b e a r g u e d t h a t t r o p i c a l a r e a s h a v e n o t b e e n t h e s u b j e c t o f a s e x t e n s i v e c o l l e c t i o n s a s more t e m p e r a t e a r e a s ; h o w e v e r , t h e r e h a v e b e e n s u f f i c i e n t d e t a i l s s u r v e y s o f t r o p i c a l a r e a s w h i c h w o u l d h a v e r e v e a l e d t h e p r e s e n c e o f N o d -u l a r i a h a d i t o c c u r r e d f r e q u e n t l y i n t r o p i c a l l a t i t u d e s ( D e s -i k a c h a r y 1959; D r o u e t 1937, 1938; F r e m y 1929; S k u j a 1949). Some o f t h e t r o p i c a l r e p o r t s o f N o d u l a r i a a r e f r o m h i g h e r more t e m p e r a t e a l t i t u d e s ( K a r i m 1968; S t a n d e y i n F i e l d H e r b a r i u m ; T i l d e n 1908; W e s t 1914). O t h e r s do n o t c o n t a i n i n f o r m a t i o n r e g a r d i n g a l t i t u d e ( M o b i u s 1893; W o o d h e a d a n d T w e e d 1957). C o l l e c t i o n s f r o m t h e A r c t i c ( F o s l i e c i t e d i n N o r d s t e d t 1897) o r t h e A n t a r c t i c ( A p s t e i n , c o l l e c t i o n i n F i e l d H e r b a r i u m ; F r i t s c h 1912; F u k u s h i m a 1959) h a v e b e e n r e p o r t e d b u t i t i s o b v i o u s f r o m o t h e r A r c t i c a n d S u b a r c t i c s u r v e y s ( C r o a s d a l e 1973; W e s t a n d W e s t 1911; W h e l d o n 1947; W i l l e 1924) t h a t N o d -u l a r i a i s uncommon i n p o l a r c l i m a t e s (60°-90° n o r t h o r s o u t h o f t h e e q u a t o r ) . What i s more i m p o r t a n t p e r h a p s i n d i s t r i b u t i o n i s t h e t y p e o f h a b i t a t . B r a c k i s h m a r i n e h a b i t a t s ( i . e . , B a l t i c S e a , New E n g l a n d c o a s t o f N o r t h A m e r i c a ) a n d a r i d o r s e m i - a r i d i n l a n d l a k e s ( i . e . , w e s t e r n N o r t h A m e r i c a ) a r e a r e a s i n w h i c h r e p e a t e d 29 F i g . 8. N o r t h A m e r i c a n d i s t r i b u t i o n o f N . s p u m i g e n a F i g . 9^ N o r t h A m e r i c a n d i s t r i b u t i o n o f N . h a r v e y a n a • h e r b a r i u m s p e c i m e n o l i t e r a t u r e r e p o r t 30 31 F i g . 1 0 . E u r o p e a n d i s t r i b u t i o n o f N. s p u m i g e n a F i g . 1 1 . E u r o p e a n d i s t r i b u t i o n o f N. h a r v e y a n a • h e r b a r i u m s p e c i m e n o l i t e r a t u r e r e p o r t 33 F i g . 12. W o r l d ( e x c l u s i v e o f N o r t h A m e r i c a a n d E u r o p e ) d i s t r i b u t i o n o f N . s p u m i g e n a F i g . 13. W o r l d ( e x c l u s i v e o f N o r t h A m e r i c a a n d E u r o p e ) d i s t r i b u t i o n o f N . h a r v e y a n a • h e r b a r i u m s p e c i m e n o l i t e r a t u r e r e p o r t 34 3 5 c o l l e c t i o n s have been made. The f a c t o r s e n a b l i n g N o d u l a r i a t o occur i n these types of waters i s the a b i l i t y of these a lgae t o grow i n a wide range of s a l i n i t i e s , a b i l i t y t o r e s -i s t r a p i d f l u c t u a t i o n s i n s a l i n i t y and r e s i s t a n c e t o d e s g i c -c a t i o n . B r i s t o l (1919) r e p o r t e d c u l t u r i n g N o d u l a r i a harveyana from a dry s o i l sample s t o r e d f o r 59 y e a r s . Labora to ry obse r -v a t i o n s i n d i c a t e t h a t i f agar p l a t e c u l t u r e s are a l l owed t o d ry i n the c u l t u r e chamber (15-30 C) and then s t o r e d at room temperature , the a l g a l m a t e r i a l i s v i a b l e and growth w i l l resume when a p o r t i o n of the d r i e d m a t e r i a l of any of the i s o -l a t e s i s i n t r o d u c e d i n t o l i q u i d media a f t e r two years s t o r a g e . Because of t h i s r e s i s t a n c e t o d e s i c c a t i o n , the ak ine t e s are p robably d i s t r i b u t e d by wind as are many o ther s m a l l a q u a t i c organisms (Maguire 1 9 6 3 ) . 36 GROWTH AND PHYSIOLOGY T h e g r o w t h a n d p h y s i o l o g y o f a number o f N o d u l a r i a i s o -l a t e s w e r e s t u d i e d u n d e r l a b o r a t o r y c o n d i t i o n s . T h e s o u r c e s o f t h e i s o l a t e s a r e g i v e n i n T a b l e I I . T h e p u r p o s e o f t h i s p o r t i o n o f t h e s t u d y was t w o f o l d . F i r s t t o c l a r i f y some e c o -l o g i c a l o b s e r v a t i o n s b y g r o w i n g t h e i s o l a t e s i n a v a r i e t y o f c o n d i t i o n s o f t e m p e r a t u r e , l i g h t , p H , a n d s a l i n i t y w i t h t h e h o p e o f o b t a i n i n g i n s i g h t i n t o t h e o c c u r r e n c e a n d g r o w t h p a t t -e r n s o f t h e a l g a i n n a t u r e . S e c o n d , t o a i d i n t h e i n v e s t i g -a t i o n o f m o r p h o l o g i c a l p l a s t i c i t y b y g r o w i n g i s o l a t e s u n d e r a v a r i e t y o f p h y s i c a l a n d c h e m i c a l v a r i a b l e s . M a t e r i a l s a n d m e t h o d s : T h e medium u s e d was BG-11 o f S t a n i e r , K u n i s a w a , M a n d e l a n d C o h e n - B a z i r e (1971), w h i c h i s a m o d i f i c a t i o n f r o m H u g h e s , G o r h a m a n d Z e h n d e r (1958) ( A p p e n d i x , T a b l e V I I I ) . S e v e r a l o t h e r m e d i a w e r e t r i e d b u t BG-11 p r o v e d t o b e t h e m o s t s a t i s -f a c t o r y c o n s i d e r i n g t h e w i d e pH r a n g e (6-11) u s e d . I s o l a t i o n s o f N o d u l a r i a s p p . w e r e made b y s t r e a k i n g t h e a l g a l m a t e r i a l c o l l e c t e d i n t h e f i e l d on 1% ( w / v ) a g a r p l a t e s a n d b y m a n u a l m a n i p u l a t i o n a n d r e p e a t e d l y r e s t r e a k i n g o f t h e f i l a m e n t s . R e m o v a l o f b a c t e r i a l c o n t a m i n a t i o n was a i d e d b y t h e p h o t o t a c t i c r e s p o n s e o f t h e h o r m o g o n i a a c r o s s t h e a g a r s u r f a c e s o r t h r o u g h t h e a g a r . S t o c k c u l t u r e s w e r e m a i n t a i n e d o n a g a r p l a t e s o r 37 T a b l e I I . O r i g i n o f i s o l a t e s u s e d i n g r o w t h a n d p h y s i o l o g y e x p e r i m e n t s . D e t a i l s o f l o c a t i o n s a r e g i v e n i n T a b l e I. i s o l a t e N o . i s o l a t e d f r o m 1 A l k a l i n e s o i l f r o m S p o t t e d L a k e B . C . ( N . s p u m i g e n a " ) 2 A l k a l i n e s o i l f r o m n e a r a s m a l l p o n d s o u t h o f R i c h t e r L a k e B . C . ( N . s p u m i g e n a ) 3 W h i t e L a k e B . C . ( N . h a r v e y a n a " ) 4 ' S a l t p o n d 4' B . C . ( N . s p u m i g e n a ) 5 C t e n o c l a d u s P o n d B . C . ( N . h a r v e y a n a ) 6 ' S a l t p o n d 3' B . C . ( N . s p u m i g e n a ) 7 I n k s L a k e B . C . ( N . s p u m i g e n a ) 8 W a l l e n d e r L a k e B . C . ( N . h a r v e y a n a ) 9 B i g Q u i l l L a k e S a s k . ( N . s p u m i g e n a , d o e s n o t f o r m a k i n e t e s , h a s g a s v a c u o l e s ) 10 S o i l c o l l e c t e d a t S i m p s o n ' s G a p , N o r t h e r n T e r r i t o r i e s A u s t r a l i a b y D r . J . R . S t e i n ( N . s p u m i g e n a ) L B - 1 4 5 2 / l f r o m t h e C u l t u r e C o l l e c t i o n o f A l g a e a n d P r o t o z o a C a m b r i d g e E n g l a n d , i s o l a t e d b y B u t c h e r ( N . h a r v e y a n a , d o e s n o t f o r m a k i n e t e s ) A l k a l i n e s o i l f r o m B o w e r s L a k e B . C . ( N . s p u m i g e n a ) 13 I n k s L a k e B . C . ( N . h a r v e y a n a ) 14 W h i t e L a k e B . C . ( N . s p u m i g e n a ) 15 Unnamed p o n d n e a r S a n de F u c a W a s h i n g t o n ( N . h a r v e y a n a ) 16 D e v i l s L a k e N o r t h D a k o t a ( N . s p u m i g e n a ) # T h e r e a s o n s f o r u s i n g t h e s e s p e c i f i c names a r e g i v e n i n t h e t a x o n o m y s e c t i o n . 38 i n l i q u i d c u l t u r e . N o d u l a r i a c u l t u r e s u s e d f o r i n i t i a t i n g e x p e r i m e n t s w e r e g r o w n i n 500 m l f l a s k s a n d f i l a m e n t s i n t h e l o g p h a s e o f g r o w t h w e r e u s e d . T e m p e r a t u r e , l i g h t , s a l i n i t y a n d n i t r o g e n e x p e r i m e n t s w e r e d o n e u s i n g 2 0 mm t e s t t u b e s , c o n t a i n i n g 2 5 m l l i q u i d medium i n e a c h , w i t h B e l l c o s t a i n l e s s s t e e l c l o s u r e s . T e s t t u b e r a c k s w e r e t u r n e d d a i l y t o i n s u r e h o m o g e n e o u s l i g h t -i n g . F o r pH e x p e r i m e n t s c u l t u r e s w e r e g r o w n i n 1 0 0 m l l i q u i d medium i n 250 m l E r l e n m e y e r f l a s k s . G r o w t h was m e a s u r e d a f t e r a s e v e n d a y p e r i o d u n l e s s o t h e r w i s e n o t e d a n d d a t a b a s e d o n d u p l i c a t e c u l t u r e s r e p l i c a t e d t h r e e t i m e s . G r o w t h o f a 3 t o 5 m l a l i q u o t o f a c u l t u r e was m e a s u r e d a s t h e o p t i c a l d e n s i t y ( a b s o r b a n c e ) a t 660 nm u s i n g e i t h e r a B e c k m a n DBG o r a B a u s c h a n d Lomb S p e c t r o n i c 2 0 s p e c t r o p h o t o m e t e r . T h e o p t i c a l d e n s i t y was r e l a t e d t o t h e number o f c e l l s b y p r e p -a r i n g s t a n d a r d c u r v e s f o r e a c h i s o l a t e . T h e s e c o m p a r i s o n c u r v e s a r e shown a s F ig .14. T e m p e r a t u r e s w e r e v a r i e d b y g r o w t h i n c o n t r o l l e d e n v i r o n -ment c h a m b e r s : r e a c h - i n t y p e f o r 15 C , 25 C , 30 C a n d 35 C ( S h e r e r C o . L t d . , M a r s h a l l , M i c h i g a n , m o d e l RT -18B-5E; C o n -t r o l l e d E n v i r o n m e n t s L t d . , W i n n i p e g , M a n i t o b a , m o d e l T 1 8 L ; P e r c i v a l C o . L t d . , B o o n e , I o w a , m o d e l I-36-L) a n d a w a l k - i n c h a m b e r f o r 2 0 C ( B e l l C r a f t I n d u s t r i e s , S u r r e y , B r i t i s h C o l -u m b i a , m o d e l 2 0 0 2 ) . 39 F i g u r e 14 . S t a n d a r d c u r v e s o f o p t i c a l d e n s i t y v e r s u s c e l l n u m b e r s . a . T h e g r o u p o f i s o l a t e s w i t h f i l a m e n t w i d t h s f r o m 8-12 m ( i . e . N . s p u m i g e n a ) . T h e r e l a t i o n s h i p b e t w e e n o p t i c a l d e n s i t y a t 660 nm a n d c e l l n u m b e r s f o r i s o l a t e s 12 ( A ) , 1 ( X ) , 6 ( • ) a n d 9 ( 0 ) . A l l o t h e r N . s p u m -i g e n a i s o l a t e s (2, 4, 10, 12, 13, 16) r e l a t i o n s h i p l i n e s f e l l b e t w e e n t h a t o f s t r a i n s 12 a n d 9* b . T h o s e i s o l a t e s w i t h f i l a m e n t w i d t h s f r o m 5-7 m ( i . e N . h a r v e y a n a ) . T h e r e l a t i o n s h i p b e t w e e n O . D . a t 660 nm a n d c e l l n u m b e r s f o r i s o l a t e s 5 (A), 8 ( X ) , a n d 14 ( o ) . A l l o t h e r i s o l a t e s o f N . h a r v e y a n a (7, H ? 15) f e l l b e t w e e n t h e two e x t r e m e s (5, 14). 40 41 L i g h t was s u p p l i e d b y G e n e r a l E l e c t r i c 20 a n d 40 w a t t c o o l - w h i t e f l o r e s c e n t t u b e s . T h e s p e c t r u m o f l i g h t f r o m 400-720 nm was m e a s u r e d b y u s i n g an a p p a r a t u s s i m i l a r t o t h e o n e d e s c r i b e d b y B u r r a n d D u n c a n ( 1 9 7 2 ) . T h e s p g e c t r a l q u a l -i t y o f t h e l i g h t i s shown a s F i g . 1 5 . T h e q u a n t i t y o f t h e l i g h t was s e t a t f o u r l e v e l s o f i l l u m i n a t i o n ( T a b l e I I I ) b y v a r y i n g t h e number o f l i g h t t u b e s a n d t h e d i s t a n c e o f t h e l i g h t t u b e s f r o m t h e c u l t u r e v e s s e l . L i g h t was m e a s u r e d w i t h a G o s s e n 1.67-873 f o o t - c a n d l e m e t e r a n d a r a d i o m e t e r ( Y S I 2 m o d e l 60) c a l i b r a t e d i n e r g s / c m / s e c . T h e p h o t o p e r i o d u n l e s s o t h e r w i s e s p e c i f i e d was 16 h o u r s : 8 h o u r s ( l i g h t : d a r k ) . G r o w t h i n t h e r a n g e o f pH 6-11 ( i n 1.0 u n i t i n c r e m e n t s ) a n d 9 - H ( i n 0.2 u n i t i n c r e m e n t s ) was i n v e s t i g a t e d . T h e w i d e r a n g e o f pH u s e d was s u c h t h a t no b u f f e r t r i e d was a d e q u a t e . T h e u s e o f two o r more b u f f e r s o v e r l a p p i n g i n r a n g e g a v e v e r y p o o r r e s u l t s . F o r pH e x p e r i m e n t s t h e m e t h o d o f G e r l o f f , F i t z g e r a l d a n d S k o o g (1952) o f a d j u s t i n g t h e pH e v e r y 12 h o u r s w i t h 1 M NaOH o r 1 M H C l was u s e d . T h e pH was m e a s u r e d w i t h a M e t r o h m E280A pH m e t e r . T h e s a l i n i t y was v a r i e d i n a r a n g e o f 1-100 ° / o o b y a d d i n g an a p p r o p r i a t e amount o f N a C l t o t h e m e d i u m . A l l i s o l a t e s w e r e g r o w n o n t h e a b o v e r a n g e o f N a C l a n d i n a d d i t i o n , f o u r i s o l a t e s (Nos 7, 9, 10, 11) were g r o w n w i t h N a £ S 0 ^ , M^SO^ 42 F i g u r e 15. Q u a l i t y of l i g h t used i n c u l t u r e experiments. (see Barr and Duncan .19 72) T a b l e I I I L i g h t i n t e n s i t i e s used i n c u l t u r e experiments. / 2 , f o o t - c a n d l e s ergs/cm /sec l u x L i g h t L e v e l 1 (LL1) 550-600 4 x l 0 4 6000-6400 L i g h t L e v e l 2 (LL2) 200 1 . 8 x l 0 4 2200 L i g h t L e v e l 3 (LL3) 60-75 0 . 9 x l 0 4 650-8( L i g h t L e v e l 4 (LL4) 25 l . l x l O 3 240 43 a n d Na^CO^ a s t h e d o m i n a n t s a l t t o d i s c o v e r i f d i f f e r e n t a n i o n s o r c a t i o n s h a d e f f e c t s o n r a t e s o f g r o w t h o r s a l i n i t y t o l e r a n c e . T h e e f f e c t o f n i t r o g e n s o u r c e s o n g r o w t h was i n v e s t i g a t e d u s i n g N O , , NH~^ a n d u r e a . T h e c o n c e n t r a t i o n o f d i s s o l v e d 3 4 s a l t s ( i . e . o s m o l a l i t y ) was k e p t c o n s t a n t b y a d d i n g a p p r o p -r i a t e a m o u n t s o f N a C l . I n o c u l a t i o n m a t e r i a l was g r o w n o n BG-11 w i t h o u t n i t r o g e n f o r 20-30 d a y s b e f o r e i n o c u l a t i o n . A p r e l i m i n a r y c o n s i d e r a t i o n was w h e t h e r g r o w t h r a t e s o f a x e n i c c u l t u r e s d i f f e r e d f r o m i d e n t i c a l c u l t u r e s w i t h l o w l e v e l b a c t e r i a l c o n t a m i n a t i o n ( l e s s t h a n 5 b a c t e r i a l c e l l s / 100 a l g a l c e l l s a s m e a s u r e d w i t h p h a s e c o n t r a s t m i c r o s c o p y ) . R e s u l t s ( d e t a i l s b e l o w ) s h o w e d t h a t t h e r e was l i t t l e d i f f e r -e n c e i n g r o w t h p a t t e r n s a n d a s a c o n s e q u e n c e a l l d a t a ( e x c e p t f o r n i t r o g e n f i x a t i o n ) i s b a s e d o n c u l t u r e s w i t h t h e same amount o f l o w l e v e l b a c t e r i a l c o n t a m i n a t i o n m e n t i o n e d a b o v e . T h e p u r i t y o f a x e n i c i s o l a t e s N o . 1 a n d N o . 6 was c h e c k e d u s i n g a number o f b a c t e r i o l o g i c a l g r o w t h m e d i a ( T a b l e I V ) a n d p h a s e c o n t r a s t m i c r o s c o p y . N i t r o g e n f i x a t i o n i n v e s t i g a t i o n s w e r e s t u d i e d w i t h f i e l d m a t e r i a l a n d u n d e r l a b o r a t o r y c o n d i t i o n s . N i t r o g e n f i x a t i o n was m e a s u r e d b y a c e t y l e n e r e d u c t i o n m e t h o d ( S t e w a r t e t a l . 1970) a t W h i t e L a k e (May 1972) a n d I n k s L a k e ( J u n e 1972) when N o d u l a r i a was p r e s e n t a s a m a j o r a l g a l c o m p o n e n t . A x e n i c 44 T a b l e I V . M i c r o b i o l o g i c a l m e d i a f o r c h e c k i n g p u r i t y o f c u l t u r e s . 1. N u t r i e n t B r o t h b e e f e x t r a c t - 3.0 g; p e p t o n e - 5.0 g; w a t e r - 1.0 1 2 . SST M e d i a g l u c o s e - 1.0 g; t r y p t o n e - 1.0 g ; y e a s t e x t r a c t -0.5 g j w a t e r - 100 m l 3. P e p t o n e - G l u c o s e g l u c o s e - 1.0 g ; p e p t o n e - 1.0 g; w a t e r - 1.0 1 4- P o t a t o - d e x t r o s e a g a r p o t a t o - 200 g; d e x t r o s e - 20 g; a g a r - 20 g ; w a t e r - 1.0 1 5. T h i o g l y c o l a t e y e a s t e x t r a c t - 5.0 g ; c a s i t o n e - 15 g j d e x t r o s e -5 g ; N a C l - 2.5 g ; 1 - c y s t i n e - 0.75 g; t h i o g l y c o -l a t e - 0.3 g ; a g a r - 0.75 gi m e t h y l e n e b l u e -.002 g ; w a t e r 1.0 1 6. S o d i u m c a s e i n a t e a g a r s o d i u m c a s e i n a t e - 3.0 g ; p e p t o n i z e d m i l k - 7*0 g; a g a r - 12 g; w a t e r - 1.0 1 45 C u l t u r e s w e r e u s e d f o r l a b o r a t o r y e x p e r i m e n t s . A n a l y s i s o f t h e g a s p h a s e was w i t h a n A e r o g r a p h g a s c h r o m a t o g r a p h ( m o d e l 600C, W i l k i n s I n s t r u m e n t a n d R e s e a r c h I n c . now V a r i a n C o . L t d . ) w i t h a s i x f o o t 0.25 i n ( O . D . ) s t a i n l e s s s t e e l c o l u m n o f P o r -a p a k R a n d w i t h h y d r o g e n f l a m e i o n i z a t i o n w i t h h e l i u m a s t h e c a r r i e r g a s . T h e a p p a r a t u s was s e t a t a t e m p e r a t u r e o f 45 C f o r a l l c o m p o n e n t s . T h r e e r e p l i c a t e s o f e a c h 1 m l s a m p l e a n d t h r e e c o n t r o l s f o r e a c h r u n w e r e u s e d . T h e c o n t r o l s c o n -s i s t e d o f : a l g a e i n c u b a t e d t h e n k i l l e d w i t h T C A ( t r i c h l o r o -a c e t i c a c i d , 2% w / v ) ; a l g a e n o t i n c u b a t e d b u t f i x e d w i t h T C A ; a l g a e f i x e d w i t h T C A t h e n a c e t y l e n e a d d e d a n d s a m p l e i n c u b a t e d . R e s u l t s : C o m p a r i s o n o f g r o w t h r a t e s s h o w e d t h a t t h e r e was l i t t l e d i f f e r e n c e b e t w e e n a x e n i c c u l t u r e s a n d c u l t u r e s w i t h l o w l e v e l b a c t e r i a l c o n t a m i n a t i o n . T h e N o d u l a r i a i n t h e c o n t a m i n a t e d c u l t u r e s g r e w a t a s l i g h t l y h i g h e r r a t e o f g r o w t h ( F i g .16). T h e c o n t a m i n a t e d c u l t u r e s w e r e c l o s e r t o t h e m o r p h o l o g y i n n a t u r e t h a n t h e m o r p h o l o g y o f a x e n i c c u l t u r e s ( T a b l e V ) . 1. T e m p e r a t u r e . T h e r e was some v a r i a b i l i t y among i s o -l a t e s b u t m o s t g r e w b e s t ( h i g h e s t O . D . ) a t e i t h e r 25 C o r 30 C . T h e maximum t e m p e r a t u r e t o l e r a n c e f o r a l l i s o l a t e s was b e t w e e n 30 C a n d 35 C ( F i g .17). T h e l o w e r l i m i t o f g r o w t h was a p p -a r e n t l y l e s s t h a n 5 C , t h e l o v / e s t t e m p e r a t u r e s t u d i e d . G r o w t h o c c u r r e d ( a l t h o u g h q u i t e s l o w l y ) a t 5 C f o r t h e 5 i s o l a t e s w h i c h 46 Figure 16. Comparison of growth of axenic cultures and cultures with b a c t e r i a l contamination. data at 25 C, 1.6 °/oo and pH 9 f o r i s o l a t e 1 (points are mean of s i x values. V e r t i c a l bar indicates range) o contaminated A axenic 47 .10 s c o o O e rt X> l< o 10 rt .08 .06 o .04 .02 (ft 4 Days T a b l e V . C o m p a r a t i v e m o r p h o l o g y o f N o d u l a r i a i n a x e n i c c u l t u r e , c o n t a m i n a t e d c u l t u r e a n d n a t u r e . 1. a u t o l y s i s 2. f i l a m e n t s h a p e a x e n i c n o t uncommon t w i s t e d 3. s p a c i n g o f i r r e g u l a r h e t e r o -c y s t s . c o n t a m i n a t e d r a r e s t r a i g h t r e g u l a r n a t u r e v e r y r a r e s t r a i g h t r e g u l a r 48 Figure 17. E f f e c t of temperature on i s o l a t e s of Nodularia. Results shown are f o r pH 9, L L 1 with basal medium ( s a l i n i t y about 1.6 /oo) afte r 7 days growth. a. i s o l a t e 1 (o) 2 (X) 3 (•) 4 (A) b. i s o l a t e 5 (o) 6 (x) 7 (•) 8 (A) c. i s o l a t e 9 (o) 10 (X) 11 (•) 12 (A) d. i s o l a t e 13 (o) 14 (X) 15 (•) 16 (A) O.D. (absorbance 660 nm) O.D. (absorbance 660 nm) 50 were t e s t e d . 2 . L i g h t . M o s t i s o l a t e s g r e w b e s t a t t h e h i g h e r l i g h t l e v e l , L L 1 ( F i g . 1 8 ) . Two i s o l a t e s ( N o s . 2 , 9) g r e w b e s t a t L L 2 a n d i s o l a t e N o . 12 g r e w b e s t a t e i t h e r L L 1 o r L L 2 d e p e n d -i n g o n t h e t e m p e r a t u r e . A t h i g h t e m p e r a t u r e s (35 C) t h e l i g h t l e v e l s c o m b i n e d w i t h t h e t e m p e r a t u r e t o i n h i b i t g r o w t h t o a g r e a t e r e x t e n t t h a n i t d i d a t l o w e r l i g h t l e v e l s . T h u s m o s t i s o l a t e s a t 35 C g r e w v e r y s l i g h t l y a t L L 4 b u t n o t a t a l l a t a n y o t h e r l i g h t i n t e n s i t i e s . 3. p H . T h e i s o l a t e s s h o w e d o p t i m a l g r o w t h a t o r n e a r pH 10 (F ig .19 a - d ) . F o u r t e e n i s o l a t e s h a d pH g r o w t h o p t i m a b e t w e e n 1 0 . 0 a n d 10.4, w h e r e a s i s o l a t e s N o . 2 a n d N o . 8 g r e w b e s t a t pH 9*4 a n d 9.6 r e s p e c t i v e l y (Fig.19 e , f ) . 4. S a l i n i t y . A w i d e r a n g e o f v a r i a b i l i t y was s e e n among t h e i s o l a t e s . Some i s o l a t e s ( N o s . 6, 8, 1 0 , 12) g r e w o , i n a maximum s a l i n i t y o f 30 / o o N a C l . H o w e v e r , m o s t i s o -l a t e s s h o w e d an u p p e r s a l i n i t y t o l e r a n c e o f 60 ° / o o . T h e s a l i n i t y o f i n i t i a l i s o l a t i o n a f f e c t e d t h e u p p e r l i m i t . C u l t u r e s i n i t i a t e d f r o m i n o c u l u m i n n o r m a l B G - 1 1 (1.6 ° / o o ) h a d a maximum t o l e r a n c e i n t h e r a n g e o f 40-50 ° / o o . H o w e v e r , i f c u l t u r e s w e r e i n i t i a t e d w i t h i n o c u l u m g r o w n a t 40 ° / o o t h e n t h e t o l e r a n c e was i n c r e a s e d t o 60 ° / o o . G r o w t h o f o / i n o c u l u m a t 60 / o o a n d a t t e m p t s a t g r o w i n g c u l t u r e s a t h i g h -e r s a l i n i t i e s w e r e u n s u c c e s s f u l . T h e s a l i n i t y a t w h i c h 51 F i g u r e 18. E f f e c t o f l i g h t i n t e n s i t y o n g r o w t h o f N o d u l a r i a G r o w t h c u r v e s f o r c , d , e , f a t 25 C, pH 9 a n d b a s a l medium ( a b o u t 1.6 ° / o o ) a . g r o w t h o f i s o l a t e 1 a t d i f f e r e n t t e m p e r a t u r e s -30 C (X), 25 C (A), 20 C (•), 15 C ( o ) , 35 C (o). b . g r o w t h o f i s o l a t e 2 a t d i f f e r e n t t e m p e r a t u r e s -30 C (X), 25 C (A), 20 C (•), 15 C ( o ) , 35 C (•) c . i s o l a t e s 1 (o), 2 (X), 3 ( • h 4 (A) d. i s o l a t e s 5 (o), 6 (X), 7 (•), 8 (A) e . i s o l a t e s 9 (o), 10 (X), 11 (•), 12 (A) f . i s o l a t e s 13 ( o ) , 14 (X), 15 (•), 16 (A) 52 LL4 25 LL3 60 LL2 200 A x O 8 A • O Ax. L L 1 light level L L 4 600 f t . - c . 25 O • 6 • O D O A O A 9A • X A D O D O A O A • x LL3 60 LL2 200 LL1 600 53 F i g u r e 19. E f f e c t o f pH o n t h e g r o w t h o f N o d u l a r i a a. g r o w t h a t pH 6-11 f o r i s o l a t e s 1 ( o ) , 2 ( X ) , 3 (•), 4 (A) b. g r o w t h a t pH 6-11 f o r i s o l a t e s 5 (o), 6 ( X ) , 7 (•), 8 (A) c . g r o w t h a t pH 6-11 f o r i s o l a t e s 9 ( o ) , 10 ( X ) , 11 . ( • ) , 12 (A) d . g r o w t h a t pH 6-11 f o r i s o l a t e s 13 ( o ) , 14 ( X ) , 15 (•), 16 (A) e. g r o w t h a t pH 9.2-10.0 f o r i s o l a t e s 2 ( X ) , 8 (A) f . g r o w t h a t pH o f 10.2-10.8 f o r i s o l a t e s 6 ( X ) , 13 ( O ) 54 55 o p t i m u m g r o w t h o c c u r e d was b e t w e e n 5 ° / o o a n d 20 ° / o o N a C l f o r a l l i s o l a t e s ( F i g . 2 0 ) . U s e o f s a l t s o t h e r t h a n N a C l g a v e e s s e n t i a l l y t h e same r e s u l t s . I s o l a t e N o . 9 s h o w e d b e t t e r g r o w t h a t h i g h e r s a l -i n i t i e s b u t a l l i s o l a t e s s h o w e d s i m i l a r l e v e l s o f t o l e r a n c e f o r s a l t s u s e d a n d h i g h e s t g r o w t h r a t e s w e r e a t t h e same s a l -i n i t i e s a s f o r N a C l . No d i f f e r e n c e s w e r e e v i d e n t w i t h Mg a s t h e m a j o r c a t i o n n o r S0^ o r C0^ a s t h e m a j o r a n i o n ( F i g . 2 1 ) . 5. N i t r o g e n . A l l i s o l a t e s g r e w much b e t t e r w i t h NO t h a n w i t h N H ^ + a s t h e s o u r c e o f n i t r o g e n ( c o m p a r e F i g . 2 2 a , b w i t h 22 c , d ) . G r o w t h was much b e t t e r i n m e d i a w i t h o u t a d d e d n i t r o g e n t h a n a t a n y l e v e l o f N H ^ + . C u l t u r e s a t l o w e r c o n -c e n t r a t i o n s o f N H ^ C l s h o w e d some g r o w t h , p e r h a p s t h e h i g h c o n -c e n t r a t i o n o f NH^"*~ u s e d i n i t i a l l y was i n h i b i t o r y . G r o w t h u s i n g u r e a a s t h e n i t r o g e n s o u r c e g a v e s i m i l a r r e s u l t s t o g r o w t h o n N H ^ + ( F i g . 22 e , f ) . A c e t y l e n e r e d u c t i o n , a s a m e a s u r e o f n i t r o g e n f i x a t i o n , u n d e r f i e l d c o n d i t i o n s ( a p p r o x . 50,000 c e l l s / l i t e r ) was f o u n d t o b e v e r y l o w ( l e s s t h a n 1 n m o l e / l p o n d w a t e r ) . I n t h e l a b o r a t o r y , t h e two a x e n i c i s o l a t e s , N o . 1 a n d N o . 6, w e r e u s e d t o i n v e s t i g a t e t h e r a t e s o f n i t r o g e n f i x a t i o n a n d t o d i s -c o v e r t h e e f f e c t o f n i t r o g e n , pH a n d s a l i n i t y on r a t e s o f n i t r o g e n f i x a t i o n . N i t r o g e n i n t h e g r o w t h m e d i a i n h i b i t e d 56 Figure 20. E f f e c t of s a l i n i t y on growth of Nodularia. Growth at s a l i n i t i e s of 1.6 - 60 °/oo at LL2, o pH 9> 25 C, inoculum grown at 1.6 /oo (normal BG-11) a . growth f o r i s o l a t e s 1 ( o ) , 2 (x), 3 (•), 4 (A) b. growth f o r i s o l a t e s 5 ( o ) , 6 ( X ) , 7 ( a ) , 8 (A) c. growth f o r i s o l a t e s 9 ( o ) , 10 ( X ) , 11 (•), 12 (A) d. growth f o r i s o l a t e s 13 ( o ) , 14 ( X ) , 15 (•), 16 (A) 57 58 F i g u r e 21. E f f e c t o f s a l i n i t i e s u s i n g s a l t s o t h e r t h a n N a C l o n g r o w t h o f N o d u l a r i a . G r o w t h a t s a l i n i t i e s 1.6 - 60 ° / o o a t L L 2 , 25 C, pH 9, i n o c u l u m g r o w n a t 1.6 ° / o o BG-11. a . b . c . d . i s o l a t e s 10 Na SO (A) MgSO ( • ) Na CO (o) i s o l a t e s 9 N a 2 S ° 4 ^ M g s 0 4 ( ° ) N a 2 C ° 3 i s o l a t e s 7 N a 2 S 0 4 (A) MgSO^ ( • ) N a ^ O ^ (o) i s o l a t e s 11 Na SO (A) MgSO ( • ) Na CO (o) z 4 4 - " 3 59 60 F i g u r e 22. E f f e c t o f n i t r o g e n s o u r c e o n t h e g r o w t h o f N o d u l a r i a . A l l d a t a a t 25 C, LL2, pH 9 a n d b a s a l med ium ( s a l i n i t y a b o u t 1.6 ° / o o ) . a. g r o w t h o n NO b y i s o l a t e s 1 ( o ) , 2 ( X ) , 3 ( • ) , 4 (A) b. g r o w t h o n N0^ b y i s o l a t e s 5 ( o ) , 6 ( X ) , 7 ( • ) , 8 (A) c . g r o w t h o n NH b y i s o l a t e s 1 ( o ) , 2 ( X ) , 3 ( • ) , 4 ( A 7 d. g r o w t h o n NH. b y i s o l a t e s 5 ( o ) , 6 ( X ) , 7 (•), 8 (A) e. g r o w t h o n u r e a b y i s o l a t e s 1 ( o ) , 2 ( X ) , 3 ( • ) , 4 (A) f. g r o w t h o n u r e a b y i s o l a t e s 5 ( ° ) , 6 ( X ) , 7 ( • ) , 8 (A) O.D. (absorbance 660 nm) O .D . (absorbance 660 nm) °« D > (absorbance 660 nm) ON 62 f i x a t i o n t o a l e v e l o f 0.25 g m / l (about 40 ppm) below which r a t e s i n c r e a s e d . Highes t r a t e s of ace ty l ene r e d u c t i o n occured a t c o n d i t i o n s s i m i l a r t o those f o r maximum growth (pH 10, s a l i n i t y 5-10 ° / o o , temperature 30 C) ( F i g . 2 3 ) . T h i s perhaps i s a r e f l e c t i o n of the b i o c h e m i c a l l i n k between photo-s y n t h e s i s and n i t r o g e n f i x a t i o n (pho tosyn the t i c p h o s p h o r y l -a t i o n s u p p l y i n g the energy requirements of the p rocess , Stew-a r t 1970). 63 F i g u r e 2 3 . E f f e c t of s a l i n i t y , pH and temperature on a c e t y l e n e r e d u c t i o n by N o d u l a r i a a. E f f e c t of s a l i n i t y on i s o l a t e 1, LL2, pH 1 0 , 25 C, b a s a l media (about 1 . 6 °/oo) b. E f f e c t of pH on i s o l a t e 1 , LL2, 2 5 C, b a s a l media (about 1 . 6 °/oo) c. E f f e c t of temperature on i s o l a t e 1 , LL2, pH 1 0 , b a s a l media (about 1 . 6 °/oo) 65 TAXONOMY The genus Nodularia was established by Mertens i n 1822 by d i s t r i b u t i n g specimens of the alga as Decas XV #4 of the ex-siccatae Algae Aquaticae of G.H.B. Juergens (1822). The type c o l l e c t i o n was made i n 1821 on the i s l a n d of Norderney, Germany (53.4°N, 7.0°E). Several specimens of the isotype material of N. spumigena exist (UC 436361, NBV 3925-51, NBV 3925-64, NBV 3925-66, NBV 3925-67, NBV 3925-68, F 951302)""". The l a s t specimen (F 951302) i s i n very good condition. The generic name was "conserved" by a decision of the Internation-a l Committee on Botanical Nomenclature (I969), as Link i n 1809 used the name to describe a red alga, Lemanea (Stafleu 1972). No published description of Nodularia existed u n t i l 1888 when Bornet and Flahault revised the heterocystous bluegreen algae. This work by Bornet and Flahault (1886-1888) has been desig-nated by the International Committee on Botanical Nomenclature as the taxonomic s t a r t i n g point of t h i s group of bluegreen * Letter i n i t i a l ( s ) are standard abbreviations of herbaria (UC - University of C a l i f o r n i a Berkeley; FH - Farlow Herbarium, Harvard University; F - F i e l d Museum of Natural History, Chic-ago; NBV - Rijksherbarium, Leiden Netherlands; NY - New York Botanical Garden; PC - Cryptogamic Herbarium of the Paris Mus-eum; BM - B r i t i s h Museum, London; UBC - University of B r i t i s h Columbia, Vancouver). The number following the herbarium des-ignation i s the accession number of the i n s t i t u t i o n (or loan number i n the case of NBV). I f no accession number has been designated, a number from 1000-1400 following the name "Nordin" refers to an i d e n t i f i c a t i o n number of the annotation l a b e l attached to the herbarium sheet a f t e r examination. 66 a l g a e ( B r i q u e t 1935). B o r n e t a n d F l a h a u l t d i d a n a d m i r a b l e j o b i n r e d u c i n g t h e t a x o n o m i c c o n f u s i o n t h a t e x i s t e d a t t h a t t i m e . B e t w e e n 1822 a n d 1888 s e v e r a l a d d i t i o n a l s p e c i e s o f N o d u l a r i a w e r e d e s c r i b e d a n d K i i t z i n g (1843) e s t a b l i s h e d t h e c l o s e l y r e l a t e d g e n u s S p e r m o s i r a . K u t z i n g s e p a r a t e d t h i s g e n u s f r o m N o d u l a r i a b e c a u s e i n J u e r g e n ' s m a t e r i a l a k i n e t e s w e r e f o r m e d s i n g l y w h e r e a s i n S p e r m o s i r a t h e a k i n e t e s w e r e f o r m e d i n g r o u p s . B o r n e t a n d F l a h a u l t d i d n o t c o n s i d e r t h i s c h a r a c t e r i s t i c t o b e o f s i g n i f a n c e a n d i n c l u d e d t h e g e n u s S p e r m o s i r a a n d i t s a c c o m p a n y i n g s p e c i e s i n t h e g e n u s N o d u l a r i a . T h e y a l s o i n c l u d e d a s p e c i e s o f L y n g b y a , JL. a n n u l a t a S u h r 1 8 3 4 , i n t h e g e n u s . B o r n e t a n d F l a h a u l t d e s c r i b e t h e g e n u s a s f o l l o w s ( t r a n s l a t e d f r o m L a t i n ) : " F i l a m e n t s f r e e w i t h i n a s h e a t h . S t e r i l e t r i c h o m e s e x c e p t i o n a l l y u n i f o r m . S h e a t h h y a l i n e , c l o s e , c ommon l y t h i n , m u c i l a g i n o u s , s o m e t i m e s l a s t i n g o n l y a s h o r t t i m e . C e l l s s h o r t , f l a t t e n e d , d i s c - s h a p e d , h e t e r o c y s t s c o m p r e s s e d . A k i n e t e s s p h e r i c a l , s u b -s p h e r i c a l o r d i s c - s h a p e d , f o r m e d i n t h e i n t e r v a l b e t w e e n h e t e r o c y s t s w i t h a s m o o t h e p i s p o r e . " A f t e r e x a m i n i n g m a t e r i a l f r o m E u r o p e a n h e r b a r i a , t h e y r e d -u c e d 16 t a x a t o 3 s p e c i e s names ( o n e s p e c i e s h a v i n g 3 v a r -i e t i e s ) . T h e y a l s o d e s c r i b e d o n e new s p e c i e s . T h e s p e c i e s w h i c h B o r n e t a n d F l a h a u l t r e c o g n i z e d a r e d i s c u s s e d b e l o w . T h e i r v a l i d i t y i s d i s c u s s e d l a t e r i n t h i s s e c t i o n . 1 . N . s p u m i g e n a (Mertens i n J u e r g e n s ] ) B o r n e t a n d F l a h a u l t , A n n . S c i . N a t . B o t . s e r . 7. 7 : 243. 1888. 67 The species has the c h a r a c t e r i s t i c s of the genus with f i l -aments 8-18 ym wide. Bornet and Flahault described three v a r i e t i e s of t h i s species. The var. genuina had filaments 8-18 ym wide and the akinetes subglobose. The var. l i t o r e a had filaments 12-16 ym wide and akinetes spherical-compressed. The var. major (not maior as i n c o r r e c t l y c i t e d by G e i t l e r , 1932) had filaments 12-18 /urn wide and akinetes f l a t t e n e d -e l l i p t i c a l . F i g . 24d. 2. N. armorica Thuret, i n Bornet and Thuret, Notes Al g o l . 2, p.122, plate 29, F i g . 12,13, 1880. This species was f i r s t described from material c o l l e c t e d near LeCroisic, France. I t was separated from N. spumigena by the unique structure of i t s akinetes which i n l i v i n g plants had biconcave end c e l l walls, and the ends of vegetative c e l l s f i t t i n g into the adjacent akinete l i k e a head into a cap ("pileatae"). The filaments were 10-12 ym wide, the c e l l s compressed and disc-shaped before d i v i s i o n ; and 3-4 times as wide as long. F i g . 24c. 3. N. harveyana (Thwaites) Thuret, Essai c l a s s . Nost., p.37S, 1875. The alga was f i r s t described by Thwaites i n W.H. Harvey's Phycologia Britannia (1848) as Spermosira harveyana. 6 8 F i g u r e 24. The species of N o d u l a r i a d e s c r i b e d i n and s i n c e the r e v i s i o n of Bornet and F l a h a u l t ( 1 8 8 8 ) . (Copied or photocopied from o r i g i n a l s ) . - a. N. harveyana drawing of type m a t e r i a l from Harvey ( 1 8 4 8 ) , see a l s o F i g . 32.x 250 b . N. sphaerocarpa from Fremy ( 1 9 2 9 ) . x 500 c. N. armorica drawing of type m a t e r i a l from Bornet and Thuret ( 1 8 8 0 ) . x 65O d. N. spumigena drawings of Merten's type m a t e r i a l from Bornet and Thuret ( 1 8 8 0 ) , a l s o see F i g . 3 1 . x 650 e . Spermosira a t l a n t i c a drawing of type m a t e r i a l from D i c k i e ( 1 8 7 1 ) . x approx. 500 f . N. paludosa drawing of type m a t e r i a l from Wolle ( 1 8 8 7 ) . x approx. 250 g. N. t u r i c e n s i s drawing of type m a t e r i a l from Hansgirg ( 1 8 9 2 ) . x 400 h. N. h a w a i i e n s i s drawing from T i l d e n ( 1 9 1 0 ) . x approx. 400 i . N. t e n u i s drawing from West ( I 9 I 4 ) on l e f t , x 500 and F r i t s c h and R i c h (1929) on r i g h t , x 1400 continued p. 75 69 7 0 Thuret ( 1875) transferred the species to Nodularia. Bornet and Flahault described the filaments as 4 - 6 ym wide; akinetes subglobose, 6-8 ym wide; vegetative c e l l s before d i v i s i o n not much longer than wide. F i g . 2 4 a . 4« N. sphaerocarpa Bornet and Flahault, Ann. S c i . Nat. Bot. ser. 7. 7 : 2 4 5 . 1 8 8 8 . This species was described as new by Bornet and Flahault from several c o l l e c t i o n s : Bory i n Belgium; Bornet i n France; Meneghini i n I t a l y on s o i l ; Roussel on tree sap i n France. The Roussel specimen was examined as a loan from the Paris Museum (PC Nordin 1 3 1 5 ) . No type specimen was designated by the authors. N. sphaerocarpa was described as being d i f f e r -ent from N. harveyana by filaments s l i g h t l y wider i n width ( 6 - 7 ym), s l i g h t l y wider width of the akinetes ( 7 - 1 0 ym) and the shape of the akinetes (spherical-compressed). F i g . 24b• The following species were described a f t e r 1888 or were not treated by Bornet and Flahault. Their present status i s discussed and the v a l i d i t y of each taxon i s considered. 5 . Spermosira a t l a n t i c a Dickie, J. Linn. Soc. Bot. 11 : 4 5 8 , F i g . 4 , 1 8 7 1 . The material on which the name was based was collected from hurricane debris i n the A t l a n t i c , 200 miles off the 71 A f r i c a n coast. F o r t i (1907) c i t e d t h i s as being incompletely-described. The type c o l l e c t i o n was the only c o l l e c t i o n made, no type material could be located and the inadequate descrip-t i o n and unconvincing f i g u r e suggest that the name should be rejected. F i g . 24«s. 6 . N. paludosa Wolle, Fresh-water Algae of the U.S. p. 2 9 1 , plate 198, F i g . 3,4. 1 8 8 7 . The basis f o r describing t h i s as a new species was the mistaken impression that european c o l l e c t i o n s of the genus were a l l made i n brackish waters and always occurred i n considerable masses. Wolle's material was c o l l e c t e d from Colorado and Pennsylvania. The c e l l dimensions are i d e n t i c a l to N. harv-eyana and from the description and i l l u s t r a t i o n i t i s possibly a N. harveyana although no akinetes were seen. Drouet (l93o) reported that no specimens of t h i s species existed i n Wolle's herbarium. C o l l e c t i o n s i d e n t i f i e d as N. paludosa have been made by Clements (I896), Ackley (1931), Coyle (1930) and Mclnteer (1939)• F o r t i (1907) considered i t incompletely described and G e i t l e r (1932) did not l i s t i t . The species should be rejected because of the inadequate description and lack of type material.. Fig . 2 4 f . 7. N. mainensis F.L. Harvey, B u l l . Torrey Bot. Club 1 6 : 188, 1889. 72 T h i s a l g a was c o l l e c t e d f r o m Pushaw S t r e a m n e a r O r o n o , M a i n e . I t was d e s c r i b e d a s h a v i n g t r i c h o m e s 33-38 \m w i d e a n d c e l l s 2-6 ym l o n g . No a k i n e t e s w e r e r e p o r t e d a n d no f i g -u r e was i n c l u d e d w i t h t h e d e s c r i p t i o n . T h e l o c a t i o n o f t h e t y p e i s unknown a n d s i n c e t h e o r i g i n a l c o l l e c t i o n , t h e s p e c -i e s h a s n o t b e e n r e p o r t e d . T h e s p e c i e s s h o u l d b e r e j e c t e d o n t h e b a s i s o f b e i n g i n c o m p l e t e l y d e s c r i b e d . 8. N . t u r i c e n s i s ( C r a m e r ) H a n s g i r g , P r o d . A l g . F l . B o h m . 2 : 74 F i g 27, 1892. T h i s was o r i g i n a l l y d e s c r i b e d a s S p e r m o s i r a t u r i c e n s i s b y C r a m e r ( i 8 6 0 ) . H a n s g i r g i n 1892 t r a n s f e r r e d t h e s p e c i e s t o t h e g e n u s N o d u l a r i a d e s p i t e t h e f a c t t h a t B o r n e t a n d F l a -h a u l t r e d u c e d S_. t u r i c e n s i s t o s y n o n o m y w i t h N . h a r v e y a n a i n 1888 ( p . 2 4 4 ) . F o r t i (1907) a n d G e i t l e r (1932) s y n o n o m i z e t h e t a x o n w i t h N . h a r v e y a n a . E x a m i n a t i o n o f h e r b a r i u m m a t -e r i a l c o l l e c t e d b y C r a m e r ( R a b e n h o r s t ' s A l g e n S a c h s e n s #994: FH N o r d i n 1084; NY N o r d i n 1262; PC N o r d i n 1236; NBV 3925-1) show i t t o b e i d e n t i c a l w i t h N . h a r v e y a n a . F i g . 24g. 9. N . h a w a i i e n s i s T i l d e n , H a w a i i a n A l m a n a c a n d A n n . (I902.) p .112, 1901. D e s c r i b e d f r o m H a w a i i a n d d i s t r i b u t e d i n T i l d e n ' s A m e r -i c a n A l g a e a s #484. E x a m i n a t i o n i n d i c a t e s t h a t i s o t y p e s 73 (UC 740394; NY N o r d i n 1250; PC N o r d i n 1339; UBC 2971) a n d t h e o n l y o t h e r two c o l l e c t i o n s , b y T i l d e n i n T a h i t i ( S o u t h P a c -i f i c A l g a e #11; UC 233460; NBV 3925-47; NY N o r d i n 1257) a n d t h a t b y C r o s s l a n d f r o m T a h i t i (UC 696203), a r e n o t N o d u l a r i a . R a t h e r t h e y a r e m o s t l i k e l y H o r m o t h a m n i o n (? s o l u t u m ) B o r n e t a n d G r u n o w . K a h n (I969) r e p o r t s H o r m o t h a m n i o n f r o m H a w a i i , i t i s a p p a r e n t l y common i n t r o p i c a l m a r i n e h a b i t a t s ( P h i l l -i p i n e s , V e l a s q u e z 1962; F l o r i d a , T a y l o r 1928; Dawson I966). H o r m o t h a m n i o n i s s u p e r f i c i a l l y s i m i l a r t o N o d u l a r i a a n d c a n b e e a s i l y m i s t a k e n f o r i t (May 1 9 4 6 , 1951) b u t d i f f e r s i n h a v i n g v e r y v a r i a b l e t r i c h o m e s i z e , i r r e g u l a r s p a c i n g o f h e t -e r o c y s t s , l a c k o f a k i n e t e s a n d a d i s t i n c t l y t r o p i c a l d i s t r i b -u t i o n . F i g . 24h. 10. N . t e n u i s G . S . W e s t , J . L i n n . S o c . B o t . 38 : 171, 1907. T h e t y p e s p e c i m e n f r o m T a n g a n y i k a was e x a m i n e d (BM N o r d i n 1347) a n d t h e m a t e r i a l h a s n o n e o f t h e a t t r i b u t e s o f t h e g e n u s . I t i s p r o b a b l y a n A n a b a e n a s p . w h i c h c o n f i r m s t h e s u g g e s t i o n o f G e i t l e r (1932). F i g . 24 i . 11. N . q u a d r a t a F r i t s c h , F r e s h w . A l g . N a t . A n t a r c t . E x p e d . p .45, p l a t e 2, F i g . 109-115, 1912. T h e s p e c i e s was d e s c r i b e d f r o m m a t e r i a l c o l l e c t e d i n t h e 74 A n t a r c t i c . F r i t s c h d e s c r i b e d i t a s a new s p e c i e s b e c a u s e i t s t r i c h o m e i s s m a l l e r (3-4 ym) t h a n t h a t o f N . h a r v e y a n a (4-6ym) a n d b e c a u s e some o f t h e h e t e r o c y s t s a r e s q u a r e . T h e m a t e r i a l (BM N o r d i n 1934) was e x a m i n e d a n d t h e o n e a k i n e t e o b s e r v e d b y F r i t s c h was n o t s e e n . T h e m a t e r i a l l a c k s o n e i m p o r t a n t c h a r -a c t e r i s t i c o f t h e g e n u s , t h a t o f r e g u l a r l y s p a c e d h e t e r o c y s t s . T h e s i g n i f i c a n c e o f t h e s q u a r e h e t e r o c y s t s i s u n c l e a r s i n c e h e t e r o c y s t s h a p e i s v a r i a b l e i n t h e t y p e m a t e r i a l . I y e n g a r a n d D e s i k a c h a r y (1944) show a f i g u r e o f N . s p u m i g e n a w i t h s q u a r e h e t e r o c y s t s . G e i t l e r (1932) c o n s i d e r e d N . q u a d r a t a t o b e a v a l i d s p e c i e s b u t i n h i s 1942 w o r k c o n s i d e r s i t t o b e an A n a b a e n a . S i n c e t h e s p e c i e s h a s n e v e r b e e n r e c o l l e c t e d d e s -p i t e d e t a i l e d (West a n d W e s t 1911; W i l l e 1924) a n d more l i m i -t e d ( H o l m - H a n s o n I964; F u k i s h i m a 1959) A n t a r c t i c s u r v e y s o f f r e s h w a t e r a l g a e , i t s s t a t u s i s q u e s t i o n a b l e . B e c a u s e o f t h e v a r i a b i l i t y o f t h e h e t e r o c y s t , i r r e g u l a r h e t e r o c y s t s p a c i n g a n d l a c k o f a k i n e t e s , t h e a l g a s h o u l d be c o n s i d e r e d an A n a b a e n a s p . r a t h e r t h a n a N o d u l a r i a . F i g . 2 4 j . 12. N . s p u m i g e n a v a r . m i n o r F r i t s c h , F r e s h w . A l g . N a t . A n t a r c t . E x p e d . , p .44, p l a t e 2, F i g . 104 -108, 1912. T h i s v a r i e t y was a l s o d e s c r i b e d f r o m m a t e r i a l c o l l e c t e d i n t h e A n t a r c t i c . T h e t y p e m a t e r i a l (BM 1815) was e x a m i n e d 75 F i g u r e 24 c o n t ' d . j . N . q u a d r a t a d r a w i n g b y F r i t s c h (1912) o f t h e t y p e m a t e r i a l , x a p p r o x 500 k . N . s p u m i g e n a v a r . m i n o r d r a w i n g b y F r i t s c h (1912) o f t h e t y p e m a t e r i a l , x a p p r o x 500 1. N . h a r v e y a n a v a r . s p h a e r o c a r p a d r a w i n g f r o m S k u j a (1926) . x 820 m. N . e p i p h y t i c a d r a w i n g o f t y p e m a t e r i a l b y G a r d n e r (1927) . x a p p r o x 300 n . N . w i l l e i d r a w i n g o f t y p e m a t e r i a l b y G a r d n e r (1927). x a p p r o x 500 o . N . f u s c a d r a w i n g o f t y p e m a t e r i a l b y T a y l o r (1928). x a p p r o x 400 p . N . s k u j a e d r a w i n g o f t y p e m a t e r i a l b y G o n z a l e z -G u e r r o (1928). x a p p r o x 500 q . N . s p u m i g e n a v a r . v a c u o l a t a t y p e m a t e r i a l d r a w n f r o m F r i t s c h a n d R i c h (1929). x 1000 r . N . s p u m i g e n a v a r . z u j a r i s d r a w i n g o f t y p e m a t e r i a l b y G o n z a l e z - G u e r r o (1930). x a p p r o x 400 s . N . a e r o p h i l a d r a w i n g o f t y p e m a t e r i a l f r o m B r a b e z (1940). x a p p r o x 1750 t . N . s p u m i g e n a v a r . a e r o p h i l a d r a w i n g o f t y p e m a t e r i a l f r o m B r a b e z (1940). x a p p r o x 500 77 a n d i s i d e n t i c a l t o N . h a r v e y a n a . No o t h e r c o l l e c t i o n s o f t h i s t a x o n h a v e b e e n r e p o r t e d b u t N . h a r v e y a n a h a s b e e n c o l l e c t e d f r o m t h e A n t a r c t i c b y A p f e l (F 1299674; F 1299702) a n d b y F u k i s h i m a (1959). F i g . 24k. 13. N . h a r v e y a n a v a r . s p h a e r o c a r p a ( B o r n e t a n d F l a -h a u l t ) E l e n k i n , B u l l . J a r d . Imp. B o t . P i e r r e l e G r . 16:331, 1916. E l e n k i n c o n s i d e r e d t h a t N . s p h a e r o c a r p a was m e r e l y an e c o l o g i c a l v a r i a n t o f N . h a r v e y a n a . E v i d e n c e f r o m e x p e r i m e n t -a l s t u d i e s i s p r e s e n t e d b e l o w ( p .89) t o s u p p o r t t h i s . F i g . 241. 14. N . e p i p h y t i c a G a r d n e r , Mem. N . Y . B o t . G a r d . 7:65, p l a t e 12, F i g . 1 6 , 1927. T h i s s p e c i e s was d e s c r i b e d f r o m m a t e r i a l c o l l e c t e d f r o m f r e s h w a t e r i n P u e r t o R i c o . T h e t y p e m a t e r i a l (NY N o r d i n 1254), t h e i s o t y p e m a t e r i a l (UC 401800) a s w e l l a s a s e c o n d c o l l e c t i o n (UC 401800a; NY N o r d i n 1255) w e r e e x a m i n e d . T h e m a t e r i a l i s a c t u a l l y j u v e n i l e s t a g e s o f N o s t o c s p . Y o u n g h o r m o g o n i a l f i l a m e n t s o f N o s t o c o f t e n do n o t r e s e m b l e t h e m a t -u r e f i l a m e n t s ( s e e G e i t l e r 1932, p . 844 N . m u s c o r u m ; p.855 N . v e r r u c o s u m ) . G e i t l e r s e e i n g o n l y G a r d n e r ' s d i a g r a m s , c o n s i d -e r e d i t an A n a b a e n a s p . No s u b s e q u e n t c o l l e c t i o n s o f N . e p i p h -y t i c a h a v e e v i d e n t l y b e e n m a d e . F i g . 24m. 78 15. N. w i l l e i G a r d n e r , Mem. N „ Y . B o t . G a r d . 7:65, p l a t e 1 2 , F i g . 1 5 , 1 9 2 7 . T h i s s p e c i e s was a l s o d e s c r i b e d f r o m m a t e r i a l c o l l e c t e d f r o m f r e s h w a t e r i n P u e r t o R i c o . T h e t y p e a n d e v i d e n t l y t h e o n l y c o l l e c t i o n (NY N o r d i n 1256) was e x a m i n e d a n d t h e m a t e r i a l i s i d e n t i c a l t o N . s p u m i g e n a . F i g . 2 4 n . 16 . N_. f u s c a T a y l o r , C a r n e g i e I n s t . W a s h . , P a p . T o r t u g a s L a b . 2 5 : 4 8 , p l a t e 1, F i g . 2 3 , 1 9 2 8 . T h e s p e c i e s was c o l l e c t e d f r o m m a r i n e p o o l s i n t h e T o r t -u g a s I s l a n d s , F l o r i d a . T h e s p e c i e s was p l a c e d i n t h e g e n u s p r o v i s i o n a l l y b e c a u s e i t l a c k e d b o t h h e t e r o c y s t s a n d a k i n e t e s . T a y l o r ( p e r s o n a l c o m m u n i c a t i o n ) no l o n g e r c o n s i d e r s t h i s s p e c -i e s t o b e a N o d u l a r i a . I t s t a x o n o m i c p o s i t i o n a t t h e p r e s e n t i s u n c l e a r . I t may b e l o n g i n t h e g e n u s J o h a n n e s b a p t i s t i a J . d e T o n i ( D r o u e t 1 9 3 8 a ) , i t may b e s y n o n o m o u s w i t h C y a n o t h r i x  w i l l e i G a r d n e r ( G a r d n e r 1927) o r i t may b e a g r o w t h f o r m o f L y n g b y a ( F r e m y a n d De T o n i 1940 c i t e d i n D r o u e t a n d D a i l y 1 9 5 6 ) . N e v e r t h e l e s s i t i s n o t a N o d u l a r i a . F i g . 24o. 1 7 . N . s k u j a e G o n z a l e z - G u e r r o , B o l . R e a l . S o c . E s p a n . H i s t . N a t . 2 8 : 4 3 5 , F i g . 1-3, 1 9 2 8 . T h i s t a x o n was d e s c r i b e d f r o m m a t e r i a l c o l l e c t e d i n t r e e s a p ( U l m u s a n d P o p u l u s ) a t C a s a de Compo a n d M a d r i d , S p a i n . 79 G e i t l e r (1932) l i s t s i t i n s y n o n o m y w i t h N . h a r v e y a n a . No t y p e m a t e r i a l c o u l d be l o c a t e d , h o w e v e r f r o m t h e d e s c r i p t i o n a n d f i g u r e t h e s p e c i e s i s i d e n t i c a l t o N . h a r v e y a n a . F i g.24p. 18. N . s p u m i g e n a v a r . v a c u o l a t a F r i t s c h T r a n s . R o y . S o c . S o u t h A f r . 18:88, 1929. F r i t s c h c r e a t e d t h i s v a r i e t y b e c a u s e o f t h e p r e s e n c e o f g a s v a c u o l e s . H o w e v e r , m a t e r i a l f r o m B a l t i c c o l l e c t i o n s ( i . e . O s t e n f e l d , NY N o r d i n 1284, 1285) a r e c o l l e c t e d r e g u l a r l y w i t h g a s v a c u o l e s . Lemmerman ( I 8 9 8 ) , S j o s t e d t (1922) a n d L i n d s t e d t (1943) h a v e r e p o r t e d g a s v a c u o l e s f o r N . s p u m i g e n a . T h i s p a r t i c u l a r a t t r i b u t e i s v a r i a b l e a n d p r o b a b l y d o e s n o t j u s t i f y t h e e s t a b l i s h m e n t o f a v a r i e t y . F i g . 2 4 q . 19. N . s p u m i g e n a v a r . z u j a r i s G o n z a l e s - G u e r r o , B o l . R e a l S o c . E s p a n . 30:224, F i g . 7-8, 1930. T h e a l g a was c o l l e c t e d f r o m Z u j a r R i v e r , S p a i n . T h e d i m -e n s i o n s o f t h e v e g e t a t i v e c e l l s m a t c h t h o s e o f N . h a r v e y a n a , h o w e v e r , t h e a k i n e t e s i z e a n d v a r i a t i o n (4-12 ym l o n g ; 7-12 ym w i d e ) i s v e r y u n l i k e t h e g e n u s . T h e f i g u r e shown may h a v e b e e n d r a w n f r o m p o o r l y p r e s e r v e d m a t e r i a l o r was e x h i b i t i n g a b n o r m a l g r o w t h . M o s t p r o b a b l y i t i s an A n a b a e n a s p . G e i t -l e r (1932) c o n s i d e r s i t t o b e an a b n o r m a l A n a b a e n a s p . T h e t y p e m a t e r i a l i s t h e o n l y c o l l e c t i o n r e p o r t e d i n t h e l i t e r a t u r e a n d 80 t h i s v a r i e t y s h o u l d b e e x c l u d e d f r o m t h e g e n u s N o d u l a r i a . F i g . 24>. 20. N_. f e r t i i i s s i m a R a n d a w a , P r o c . I n d . A c a d . S c i . B. 4:41, 1936. T h i s s p e c i e s was p u b l i s h e d f r o m a c o l l e c t i o n f r o m N o r t h -e r n I n d i a . No d e s c r i p t i o n was g i v e n , t h u s i t i s r e j e c t e d a s a nomen n u d u m . 21. N . a e r o p h i l a B r a b e z , B e i h e f t e zum B o t . Z e n t -r a b l a t t 6lA:214, F i g . 8 a , 1941. T h i s s p e c i e s was r e p o r t e d f r o m A u s t r i a a n d d e s c r i b e d a s a new s p e c i e s o n t h e b a s i s o f i t s s o i l h a b i t a t , s h o r t f i l a -m e n t s a n d l a c k o f a k i n e t e s . T h e l a c k o f a k i n e t e s i s a n e g -a t i v e c h a r a c t e r i s t i c a n d t h e o t h e r two c h a r a c t e r i s t i c s do n o t j u s t i f y i t s s e p a r a t i o n f r o m N . h a r v e y a n a t o w h i c h i t i s i d e n t -i c a l i n c e l l s i z e a n d s h a p e . T h e t y p e m a t e r i a l a n d o n l y c o l l -e c t i o n was n o t e x a m i n e d b u t t h e p u b l i s h e d d e s c r i p t i o n a n d f i g -u r e a r e t h a t o f N . h a r v e y a n a w h i c h h a s b e e n r e p o r t e d many t i m e s f r o m s o i l . F i g . 24s. 22. N . s p u m i g e n a v a r . a e r o p h i l a B r a b e z , B e i h e f t e zum B o t . Z e n t r a b l a t t 6lA:215, 1941. T h i s v a r i e t y was c o l l e c t e d i n t h e same s a m p l e a s N . a e r -81 o p h i l a a n d was s e p a r a t e d o n s i m i l a r c h a r a c t e r i s t i c s ( a e r i a l h a b i t a t ; s h o r t f i l a m e n t s ; l a c k o f a k i n e t e s ) . I t was d e s -c r i b e d f r o m a s i n g l e f i l a m e n t a n d b y i t s d e s c r i p t i o n i s i d e n t -i c a l t o N . s p u m i g e n a , t h e r e f o r e i t s s e p a r a t i o n i s c o m p l e t e l y u n j u s t i f i e d . F i g . 24t. 23. N . i m p l e x a ( B o r n e t a n d F l a h a u l t ) B o u r r e l l y L e s A l g u e s d ' E a u D o u c e . I I I . p.418, F i g .5-6, 1970. B o u r r e l l y t r a n s f e r r e d A u l o s i r a i m p l e x a t o N . i m p l e x a a n d a d v o c a t e d i n c l u s i o n o f p a r t o f t h e g e n u s A u l o s i r a w i t h i n t h e g e n u s N o d u l a r i a . He m a i n t a i n e d t h a t t h e o n l y d i f f e r e n c e b e t w e e n t h e two g e n e r a was t h a t t h e c e l l s o f N o d u l a r i a w e r e t h i n n e r ( a s c o m p a r e d t o t h e i r w i d t h ) a n d t h e s h e a t h was t h i n r he r ( i . e . , A u l o s i r a h a s a t h i c k e r s h e a t h a n d t h e c e l l s a r e n o t a s f l a t t e n e d ) . T h e g e n u s N o d u l a r i a i s d i s t i n c t b e c a u s e t h e c e l l s a r e d i s t i n c t l y w i d e r t h a n l o n g ( N . s p u m i g e n a 2-4x a s w i d e a s l o n g ; N . h a r v e y a n a 1.5-2x a s w i d e a s l o n g a n d o n l y b e i n g a s w i d e a s l o n g p r i o r t o c e l l d i v i s i o n ) . I n A u l o s i r a t h e c e l l s h a v e a l w a y s b e e n c o n s i d e r e d e i t h e r a s l o n g a s w i d e o r l o n g e r t h a n w i d e ( e x c e p t A . s c h a u i n s l a n d i i Lemm. w h i c h r e s e m b l e s a H o r m o t h a m n i o n s p . a n d A . i m p l e x a v a r . c r a s s a D i x i t ) . A n o t h e r d i s t i n c t m o r p h o l o g i c a l f e a t u r e i s t h a t a k i n -e t e s o f N o d u l a r i a a r e e i t h e r r o u n d o r w i d e r t h a n l o n g a n d a l w a y s s e p a r a t e f r o m t h e h e t e r o c y s t . T h e a k i n e t e s o f A u l o s i r a 82 a r e i n v a r i a b l y l o n g e r t h a n w i d e a n d may b e e i t h e r r e m o v e d f r o m o r a d j a c e n t t o t h e h e t e r o c y s t ( D e s i k a c h a r y 1959). T h e h e t e r -o c y s t s o f N o d u l a r i a a r e r e g u l a r l y s p a c e d a l o n g t h e f i l a m e n t , b u t A u l o s i r a h a s r a n d o m a r r a n g e m e n t o f h e t e r o c y s t s . T h i s c h a r -a c t e r i s t i c was u s e d b y T h u r e t (1875) a s a m a j o r c h a r a c t e r i s t i c t o d e f i n e t h e g e n u s N o d u l a r i a . T h e r e i s a l s o a d i s t i n c t h a b -i t a t d i f f e r e n t i a t i o n . N o d u l a r i a i s t y p i c a l l y f o u n d i n m a r i n e a n d b r a c k i s h s i t u a t i o n s a n d o n l y o c c a s i o n a l l y i n f r e s h w a t e r o r o n s o i l . A u l o s i r a , o n t h e o t h e r h a n d , i s p r i m a r i l y a s o i l a n d f r e s h w a t e r g e n u s a n d i s o n l y r a r e l y f o u n d i n b r a c k i s h o r m a r i n e e n v i r o n m e n t s . T h e d i s t r i b u t i o n p a t t e r n s o f t h e two a r e v e r y d i f f e r e n t . N o d u l a r i a o c c u r s p r i m a r i l y i n t e m p e r a t e e n v i r o n m e n t s , o c c a s i o n a l l y i n t h e s u b t r o p i c s a n d o n l y r a r e l y i n t h e t r o p i c s . A u l o s i r a i s p r i m a r i l y t r o p i c a l a n d s u b t r o p i c a l , a n d l e s s commonly r e p o r t e d f r o m t e m p e r a t e a r e a s . F o r t h e s e r e a s o n s t h e g e n u s N o d u l a r i a s h o u l d n o t i n c l u d e t h e g e n u s A u l o -s i r a o r a n y p a r t o f i t . 24. N . s p u m i g e n a f . c r a s s a ( W o r o n i c h i n ) E l e n k i n . T h e o r i g i n a l c i t a t i o n o f t h i s t a x o n c o u l d n o t b e l o c a t e d . T h e o n l y r e f e r e n c e t o i t , i n S t a r m a c h (I966), a p p e a r s f r o m t h e d e s c r i p t i o n a n d f i g u r e ( p . 518-519) t o b e o f a n N . s p u m i g e n a s p e c i m e n f r o m S i b e r i a w i t h p a r t i c u l a r l y l a r g e c e l l d i m e n s i o n s . 83 V a l i d i t y o f B o r n e t a n d F l a h a u l t ' s N o d u l a r i a t a x a : T h e p r e c e d i n g summary i n d i c a t e s t h a t a l l t h e s p e c i e s d e s c r i b e d s i n c e 1 8 8 8 c a n b e shown t o be e x c l u d e d f r o m t h e g e n u s o r c a n b e i n c l u d e d w i t h i n s p e c i e s o f N o d u l a r i a d e f i n e d b y B o r n e t a n d F l a h a u l t ( 1 8 8 8 ) , i t i s n e c e s s a r y t o e s t a b l i s h t h e v a l i d i t y o f t h e t a x a d e s c r i b e d b y B o r n e t a n d F l a h a u l t . T h e q u e s t i o n s t h a t w e r e c o n s i d e r e d a r e : i s N . a r m o r i c a a d i s t i n c t e n t i t y ; a r e t h e v a r i e t a l names o f N . s p u m i g e n a j u s t i f i e d ; i s N . s p h a e -r o c a r p a a d i s t i n c t s p e c i e s ; a r e N . h a r v e y a n a a n d N . s p u m i g e n a d i s t i n c t s p e c i e s o r m e r e l y two h a l v e s o f a c o n t i n u u m o f a s i n g l e s p e c i e s ? V a l i d i t y o f N . a r m o r i c a : B o r n e t a n d F l a h a u l t (1888) s t a t e t h e d i s t i n g u i s h i n g c h a r a c t e r i s t i c o f t h i s s p e c i e s i s t h e b i c o n c a v e , t r u n c a t e a k i n e t e s i n w h i c h t h e e n d o f an a d j a -c e n t v e g e t a t i v e c e l l i s h i d d e n b y t h e c o n c a v e e n d o f t h e a k i n e t e . T h e y n o t e t h a t t h i s c h a r a c t e r i s t i c i s s e e n o n l y i n l i v i n g m a t e r i a l . E v i d e n t l y o n l y f i v e c o l l e c t i o n s h a v e b e e n g i v e n t h e s p e c i f i c name a r m o r i c a . T h e f i r s t was t h e t y p e c o l l -e c t e d i n F r a n c e b y T h u r e t i n 1873. T h r e e c o l l e c t i o n s w e r e made o n t h e P a c i f i c C o a s t o f N o r t h A m e r i c a (1. G a r d n e r 436, UC 100567; 2. G a r d n e r 602, UC 100568; 3. O s t e r h o u t a n d G a r d -n e r PBA 1061; FH N o r d i n 1087, NBV 3925-30, F 9808I6, NY N o r d i n 1213, PC N o r d i n 1302) a n d t h e f i f t h i n F r a n c e (Freray a n d M e s l i n 1924). Of t h e t h r e e N o r t h A m e r i c a n c o l l e c t i o n s o n e 84 (UC 100568) f r o m W h i d b e y I s . W a s h , was i n p o o r c o n d i t i o n . T h e o t h e r two d i d n o t show t h e c h a r a c t e r i s t i c a k i n e t e a s w o u l d b e e x p e c t e d w i t h p r e s e r v e d m a t e r i a l a n d w e r e i d e n t i c a l t o N . s p u m i g e n a . T h e two c o l l e c t i o n s b y G a r d n e r (436, 602) w e r e p l a c e d i n t h e s p e c i e s a r m o r i c a " d o u b t f u l l y " ( S e t c h e l l a n d G a r d n e r 1903) s i n c e t h e s p e c i m e n s d i f f e r e d i n s e v e r a l d e t -a i l s f r o m B o r n e t a n d F l a h a u l t ' s (1888) d e s c r i p t i o n . T h e f i g -u r e shown i n F r e m y a n d M e s l i n ' s r e p o r t (1924) i s v e r y p o o r . N . a r m o r i c a a n d N . s p u m i g e n a a r e i d e n t i c a l e x c e p t f o r t h e u n i q u e a k i n e t e s t r u c t u r e . I n d e e d t h i s c h a r a c t e r i s t i c may b e a m i c r o s c o p i c a r t i f a c t ( F i g . 25a,b). H o w e v e r some h e r b a r i u m s p e c i m e n s i d e n t i f i e d a s N . s p u m i g e n a d i d show b i c o n c a v e a r e a s b e t w e e n t h e a k i n e t e s (UC 761550; UC 752514) ( F i g . 25c,d , e ) . T h i s i s o d d s i n c e t h e c h a r a c t e r i s t i c s u p p o s e d l y o c c u r s o n l y i n l i v i n g c e l l s ( B o r n e t a n d F l a h a u l t 1888). S t u d i e s u n d e r v a r y -i n g g r o w t h c o n d i t i o n s t o d e t e r m i n e t h e s t a b i l i t y o f t a x o n o m i c c h a r a c t e r i s t i c s show t h a t a k i n e t e s h a p e , s i z e a n d w a l l c h a r -a c t e r i s t i c s a r e f a r m o r e v a r i a b l e t h a n s i z e a n d s h a p e o f v e g -t a t i v e c e l l s o r h e t e r o c y s t s ( F i g . 26). T h u s s e p a r a t i o n o f a s p e c i e s o n a k i n e t e c h a r a c t e r i s t i c s a l o n e i s o f d o u b t f u l v a l u e . W i t h t h i s e v i d e n c e , N . a r m o r i c a i s c o n s i d e r e d a n e c o l o g i c a l v a r i a n t o f N . s p u m i g e n a a n d i s p l a c e d i n s y n o n o m y w i t h i t . V a l i d i t y o f v a r i e t a l names o f N . s p u m i g e n a : E x a m i n a t i o n o f a l a r g e number o f h e r b a r i u m s p e c i m e n s , o b s e r v a t i o n o f t h e 85 F i g u r e 25 . P o s s i b l e e x p l a n a t i o n f o r a k i n e t e c h a r a c t e r i s t i c s o f N. a r m o r i c a . a . A s i m p l i f i e d r e p r e s e n t a t i o n s h o w i n g a c h a i n o f a k i n -e t e s i n s i d e v i e w , w i t h f i l a m e n t n o t b e i n g h o r i z o n t a l w i t h r e s p e c t t o s l i d e s u r f a c e . b . I f t h e f i l a m e n t w e r e n o t h o r i z o n t a l , t h e n a t o p v i e w m i g h t show t h e e n d s o f t h e a k i n e t e s v i s u a l l y o v e r -l a p p i n g t o g i v e t h e a p p e a r a n c e o f " d i s c s " b e t w e e n t h e a k i n e t e s . c . d . a n d e show " d i s c " a r e a s b e t w e e n a k i n e t e s c h a r a c t e r -i s t i c o f N. a r m o r i c a . 87 F i g u r e 26. C o m p a r i s o n o f t h e v a r i a b i l i t y o f s i z e o f v e g e t a t -i v e c e l l s , h e t e r o c y s t s a n d a k i n e t e s o f two i s o -l a t e s u n d e r a v a r i e t y o f c u l t u r e c o n d i t i o n s . C o u n t o f 20 r a n d o m c e l l s o f e a c h t y p e u n d e r v a r i a -t i o n s o f pH (8^9,10) , t e m p e r a t u r e (20 C , 25 C ) a n d s a l i n i t y (1.6 / o o , 10 / o o , 20 / o o ) n = 360. a . i s o l a t e 7 ( i ) v e g e t a t i v e c e l l s ( i i ) h e t e r o c y s t s ( i i i ) a k i n e t e s . b . i s o l a t e 3 ( i ) v e g e t a t i v e c e l l s ( i i h e t e r o c y s t s ( i i i ) a k i n e t e s . 88 2 0 0 o CM O U 4) .O E 3 S3 1 0 0 I'l C.V .=8.15# i l l s 2 = „ 5 2 C.V . = 9 . 6 0 ^ 4 . 5 5 . 0 5 . 5 6 . 0 5 . 5 6 . 0 6 . 5 7 . 0 6 . 0 6 . 5 7 . 0 7 - 5 8 . 0 8 - 5 9 . 0 9 - 5 200 L 0] H H 0) u CM 0 100 0) .o B 3 sz; s 2=.12 C.V .=6.80fo 11 s 2=.12 C.V.=6.18# 4 . 5 5 . 0 5 . 5 6 . 0 5 . 0 5 . 5 6 . 0 6 . 5 111 s 2 = , 6 3 C.V . = 8 . 6 9 £ 5 - 5 6 . 0 6 . 5 7 . 0 7 . 5 8 . 0 8 . 5 9 . 0 C e l l width ( ym) 89 v a r i a b i l i t y o f s e v e r a l i s o l a t e s u n d e r w i d e l y v a r y i n g c u l t u r e c o n d i t i o n s a n d a l a r g e number o f i s o l a t e s i n n a t u r e , i n d i c a t e s t h a t t h e r e seems l i t t l e j u s t i f i c a t i o n f o r r e t e n t i o n o f v a r -i e t a l names o f N . s p u m i g e n a . T h e d i m e n s i o n s w e r e a r t i f i c -i a l l y e s t a b l i s h e d ( t h e c h o i c e i s among 8 - 1 2 ym, 1 2 - 1 6 ym a n d 1 2 - 1 8 um) a n d a k i n e t e s h a p e s a r e v e r y v a r i a b l e among a c l o n a l p o p u l a t i o n o r o n t h e same f i l a m e n t . D e v e l o p m e n t a l s t a g e s o f a k i n e t e f o r m a t i o n e n c o m p a s s a v a r i e t y o f s h a p e s d u r i n g i t s m a t u r a t i o n a n d c a t e g o r i z i n g t h e a k i n e t e s a s s u b s p h e r i c a l , s p h e r i c a l - c o m p r e s s e d a n d f l a t t e n e d - e l l i p t i c a l a r e v a l u e l e s s . V a l i d i t y o f N . s p h a e r o c a r p a : T h e s i z e a n d s h a p e o f a k i n -e t e s , h e t e r o c y s t s a n d v e g e t a t i v e c e l l s o f h e r b a r i u m s p e c i m e n s o f N . h a r v e y a n a a n d N . s p h a e r o c a r p a w e r e c o m p a r e d . H a b i t a t a n d c o l l e c t i o n s i t e s w e r e c o m p a r e d t o s e e i f h a b i t a t d i f f e r -e n c e s w e r e s i g n i f i c a n t . T h e r e s u l t s ( F i g . 2 7 ) show t h e r e i s a d i f f e r e n c e i n s i z e o f c e l l s o f h e r b a r i u m s p e c i m e n s , w i t h N . h a r v e y a n a g e n e r a l l y b e i n g s m a l l e r . H o w e v e r , t h e r e i s no s e p -a r a t i o n i n t o d i s t i n c t g r o u p s , r a t h e r m e r e l y a g r a d a t i o n . T h e r e a p p e a r s t o b e a d i f f e r e n c e i n h a b i t a t w i t h N . s p h a e r o -c a r p a c o l l e c t e d more f r e q u e n t l y f r o m t h e s o i l t h a n N . h a r v e y a n a . Of 6 8 c o l l e c t i o n s o f N . h a r v e y a n a e x a m i n e d ( i d e n t i f i c a t i o n a c c o r d i n g t o c o l l e c t o r ) 3 2 . 3 % , 5 4 . 4 % , a n d 1 3 . 2 % w e r e f r o m b r a c k i s h - m a r i n e , " f r e s h w a t e r " , a n d s o i l h a b i t -a t s r e s p e c t i v e l y . Of 2 4 N . s p h a e r o c a r p a c o l l e c t i o n s ( s p e c i e s 90 F i g u r e 27. C o m p a r i s o n o f c e l l s i z e s o f N . h a r v e y a n a a n d N . s p h a e r o c a r p a . o N . s p h a e r o c a r p a ( i d e n t i f i c a t i o n o n h e r b -a r i u m s h e e t ) A N . h a r v e y a n a ( i d e n t i f i c a t i o n o n h e r b -a r i u m s h e e t ) a . v e g e t a t i v e c e l l s b . h e t e r o c y s t s c . a k i n e t e s C e l l length ( y m) C e l l length (y m) > > 0 o > t> o o > o « o O n L a O o o > 9 rj o ogo cp o o o o C e l l length (y m ) O O O 92 i d e n t i f i c a t i o n a c c o r d i n g t o o r i g i n a l c o l l e c t o r ) t h e r e s p e c t i v e f i g u r e s w e r e 22.6%, 52.1% a n d 25.3%. A n i n t e r e s t i n g c o r r e l a t i o n f r o m c u l t u r e s t u d i e s i s t h a t two o f t h e t h r e e i s o l a t e s i d e n t i f i e d a s N. h a r v e y a n a , when g r o w n o v e r a w i d e r a n g e o f s a l i n i t i e s (1-40 °/oo), show a p o s -i t i v e c o r r e l a t i o n b e t w e e n s a l i n i t y a n d a k i n e t e s i z e ( F i g . 28). T h e h i g h e r t h e s a l i n i t y i n w h i c h a k i n e t e s a r e f o r m e d , t h e l a r g e r t h e i r s i z e . H o w e v e r t h e t h i r d i s o l a t e shows no s u c h c o r r e l a t i o n . T h u s , t o make a d i s t i n c t i o n b e t w e e n N . h a r v e y -a n a a n d N . s p h a e r o c a r p a b a s e d o n a k i n e t e s i z e i s h a r d t o j u s t -i f y . T h e s e o b s e r v a t i o n s seem t o s u p p o r t E l e n k i n ' s (1916) p r o p o s a l o f t r e a t i n g N . s p h a e r o c a r p a a s t h a t p o r t i o n o f N . h a r v e y a n a w i t h t h e l a r g e s t d i m e n s i o n s . V a l i d i t y o f N . h a r v e y a n a a n d N. s p u m i g e n a a s d i s t i n c t t a x a : T h i s was e x a m i n e d b y o b s e r v a t i o n o f h e r b a r i u m m a t e r i a l , f i e l d c o l l e c t i o n s a n d g r o w t h o f m a t e r i a l i n t h e l a b o r a t o r y . S c a t t e r g r a m s f r o m h e r b a r i u m m a t e r i a l o f a k i n e t e , h e t e r o c y s t a n d v e g e t a t i v e c e l l s i z e show a s e p a r a t i o n i n t o t w o d i s t i n c t g r o u p s ( F i g . 2 9 ) . T h i s c o n t r a s t s w i t h t h e v i e w o f B u r s a (1968) t h a t o n l y a s i n g l e s p e c i e s o f N o d u l a r i a s h o u l d b e c o n -s i d e r e d . An o b v i o u s s u p p o r t f o r t h e m a i n t e n a n c e o f t h e two s p e c i e s i s t h e f a c t t h a t t h e two a r e o f t e n c o l l e c t e d a t t h e same t i m e i n t h e same p l a c e a n d a r e r e a d i l y s e p a r a b l e i n t o two s i z e c l a s s e s . 93 F i g u r e 28. A k i n e t e v a r i a b i l i t y o f t h r e e B r i t i s h C o l u m b i a i s o l a t e s a s r e l a t e d t o s a l i n i t y . a . i s o l a t e 7 1 . 6 1 o 10%= A 20%n b. i s o l a t e 5 I . 6 & 0 10%>A 20%D c . i s o l a t e 3 1 . 6 ^ o 1 0 & A 20%D Number of akinetes Number of akinetes Number of akinetes H O t o O O o O Cn O Cn O Cn o oo O oo Cn o Cn 95 F i g u r e 2 9 . C o m p a r i s o n o f c e l l s i z e s o f N. s p u m i g e n a a n d N. h a r v e y a n a . o N. spumigena A N» harveyana a. vegetative c e l l s i z e b. heterocyst s i z e c. akinete s i z e ( i d e n t i f i c a t i o n o n h e r b -a r i u m s h e e t ) ( i d e n t i f i c a t i o n o n h e r b -a r i u m s h e e t ) C e l l length (v m) Oj J*. Lr, O 0 0 o » o cb°o° O O O QJJ 0 (9 o o o o o C e l l length ( Um) Co > > o a o (§> o @ o o 0 3 Q 0) 97 One a p p r o a c h u s e d t o s e p a r a t e b l u e g r e e n a l g a l t a x a , t h a t o f g r o w t h p a t t e r n s o n a g a r p l a t e s (Kanfcz a n d B o l d I969, B a k e r a n d B o l d 1970) p r o v e d c o m p l e t e l y u n w o r k a b l e . I s o l a t e s m i c r o -s c o p i c a l l y i d e n t i c a l e x h i b i t e d s t r i k i n g l y d i f f e r e n t g r o w t h c h a r a c t e r i s t i c s , a n d r e c o g n i z i n g d i f f e r e n t s p e c i e s o r e v e n g e n e r a i n t h i s manner p r o v e d i m p o s s i b l e ( F i g . 3 0 ) . T h e h e r b a r i u m m a t e r i a l c o n s i d e r e d d u r i n g t h i s s t u d y i s shown i n A p p e n d i x , T a b l e I X . T h u s t h e g e n u s s h o u l d c o n t a i n o n l y two s p e c i e s , N . h a r v -e y a n a ( T h w . ) T h u r e t a n d N . s p u m i g e n a M e r t e n s . T h e g e n u s i s d i s t i n c t a n d e a s i l y s e p a r a b l e f r o m o t h e r m o r p h o l o g i c a l l y s i m i l a r g e n e r a . C e r t a i n c h a r a c t e r i s t i c s o f t h e g e n u s ( c e l l s b e i n g w i d e r t h a n l o n g ; r e g u l a r s p a c i n g o f t h e h e t e r o c y s t s ; f o r m a t i o n o f a k i n e t e midway b e t w e e n t h e h e t e r o -c y s t s ) v a r i e d v e r y l i t t l e u n d e r t h e w i d e r a n g e o f g r o w t h c o n d -i t i o n s . O t h e r c h a r a c t e r i s t i c s w e r e q u i t e v a r i a b l e ( s h e a t h p r e s e n c e ; f i l a m e n t l e n g t h ; v e g e t a t i v e c e l l c o l o r ; a k i n e t e s h a p e , t e x t u r e a n d c o l o r ) . A n i l l u s t r a t i o n o f v a r i a b i l i t y i n t h e m o r p h o l o g y o f t h e g e n u s i s shown i n F i g . 31 a n d 3 2 . F r o m t h e i n v e s t i g a t i o n p r e s e n t e d h e r e an e m e n d a t i o n o f t h e g e n e r i c a n d s p e c i e s d e s c r i p t i o n i s a s f o l l o w s : N o d u l a r i a ( M e r t e n s i n J u e r g e n s } B o r n e t a n d F l a h a u l t e m e n d . F i l a m e n t s w i t h c e l l s d i s t i n c t l y w i d e r t h a n l o n g , h e t e r o c y s t s s p a c e d a t r e g u l a r i n t e r v a l s a l o n g t h e t r i c h o m a . I n i t i a t i o n a n d f o r m a t i o n o f a k i n e t e s midway b e t w e e n t h e h e t e r o c y s t s . 98 F i g . 30. C u l t u r a l v a r i a t i o n o f N o d u l a r i a . C o m p a r i s o n o f g r o w t h p a t t e r n s o f s e v e r a l i s o l a t e s a f t e r two weeks g r o w t h o n BG-11 1 % (W-V) a g a r a t 20 C pH 8 a n d L L l . I d e n t i f i c a t i o n n u m b e r s r e f e r t o i s o l a t e n u m b e r s . " IUCC 583" i s a C a l o t h r i x s p . w h i c h h a s b e e n m i s - i d e n t i f i e d a s N o d u l a r i a . 99 100 T h e m o r p h o l o g i c a l v a r i a t i o n o f N . s p u m i g e n a . f r o m F r i t s c h 1 9 5 1 x 8 5 0 , shows t e r m i n a l a n d i n t e r -c a l a r y h e t e r o c y s t s , s h e a t h a n d l o c a t i o n o f f o r m a t i o n o f a k i n e t e s . f r o m B o r n e t a n d T h u r e t 1880 a m e n t . f r o m B o r n e t a n d T h u r e t 1880 a k i n e t e s . x 6 5 0 , v e g e t a t i v e f i l -x 6 5 0 , f i l a m e n t w i t h f r o m B o r n e t a n d T h u r e t 1880 x 6 5 0 , t h e i r d r a w i n g o f M e r t e n ' s t y p e m a t e r i a l , s h o w s a v e g e t a t i v e f i l a m e n t a n d a f i l a m e n t w i t h a s i n g l e i s o l a t e d a k i n e t e . f r o m B h a r a d w a j a 1935 x l 3 7 0 , v e g e t a t i v e f i l a m e n t a n d f i l a m e n t w i t h a k i n e t e s . f r o m P l a y f a i r 1914 x 8 0 0 , v e g e t a t i v e f i l a m e n t a n d f i l a m e n t w i t h i m m a t u r e a k i n e t e s . f r o m P r e s c o t t 1962 x 9 0 0 , v e g e t a t i v e f i l a m e n t a n d f i l a m e n t w i t h a k i n e t e s . t o H (fflwiiiiiiiiiiiim in D 1 0 2 Figure 32. The morphological v a r i a t i o n of N. harveyana. a. from Bornet and Thuret 1880 x650, vegetative f i l -ament . b. from Harvey I848 x250, filament with akinetes (drawing of type material). c. from Smith 1950 x800, vegetative filament. d. from Hortobagyi 19 59 xlOOO, vegetative filament and filament with akinetes. e. from Hardingl971 xlOOO, vegetative filament. f . from G e i t l e r 1932 x900, vegetative filament. g. from Mabille 1954 xlOOO, vegetative filament and filament with akinetes. h. from Carter 1933 x 6 l 3 , vegetative filament. i . from Skuja 1926 x820, filaments with immature and mature akinetes. 103 104. V e g e t a t i v e c e l l s 7.5-16.0 p m w i d e , w i d t h / l e n g t h r a t i o 2:1 - 10:1; s o m e t i m e s w i t h gas v a c u o l e . H e t e r o c y s t s s u b s p h e r i c a l t o d i s c - s h a p e d ; w i d e r t h a n l o n g 8-16 ym w i d e , 2-10 u m l o n g . A k i n e t e s s u b s p h e r i c a l t o d i s c - s h a p e d ; 8-18 urn w i d e , 6-15 U«n l o n g , u s u a l l y i n a s e r i e s ( o c c a s i o n a l l y s i n g l e o r in t w o ' s ) . F i l a m e n t s u s u a l l y w i t h a t h i n c o l o r l e s s t r a n s p a r e n t s h e a t h . H a b i t a t t y c h o p l a n k t o n o r e u p l a n k t o n ; m a r i n e , b r a c k -i s h , i n l a n d s a l i n e , o r ' h a r d ' f r e s h w a t e r . T e m p e r a t e a n d s u b t r o p i c a l i n d i s t r i b u t i o n . N . s p u m i g e n a [ M e r t e n s i n J u e r g e n s ] B o r n e t a n d F l a h a u l t e m e n d . V e g e t a t i v e c e l l s 3.5-7*2 y m w i d e ; w i d e r t h a n l o n g ( e x c e p t j u s t b e f o r e d i v i s i o n when w i d t h a n d l e n g t h may be e q u a l ) ; w i d t h / l e n g t h r a t i o 2.5:1 - 1:1. H e t e r o c y s t s s u b s p h e r i c a l ( w i d e r t h a n l o n g ) t o s p h e r i c a l ; 4•0-7«5 um w i d e . A k i n e t e s u b s p h e r i c a l ( w i d e r t h a n l o n g ) t o s p h e r i c a l 4-9 U m w i d e , 4-8 ym l o n g . T h i n s h e a t h may o r may n o t b e p r e s e n t . H a b i t a t t y c h o p l a n k t o n o r b e n t h i c ; m a r i n e , b r a c k i s h o r i n l a n d s a l i n e s i t u a t -i o n s ^ o c c a s i o n a l l y i n f r e s h w a t e r ; a l s o f o u n d on s o i l o r i n t r e e s a p . T e m p e r a t e a n d s u b -t r o p i c a l i n d i s t r i b u t i o n . N . h a r v e y a n a ( T h w . ) T h u r e t e m e n d . The synonomy o f o t h e r t a x a a t t r i b u t e d to N o d u l a r i a a r e shown i n A p p e n d i x , T a b l e X . 105 D I S C U S S I O N E c o l o g y a n d P h y s i o l o g y : Few c o l l e c t o r s h a v e made q u a n t i t a t i v e m e a s u r e m e n t s o f t h e e n v i r o n m e n t i n w h i c h N o d u l a r i a s p p . w e r e c o l l e c t e d a n d v i r t -u a l l y no l a b o r a t o r y work h a s b e e n d o n e o n t h e m . I t s o c c u r r -e n c e i n n a t u r e w i l l b e c o n s i d e r e d w i t h r e f e r e n c e t o e c o l o g i c a l o b s e r v a t i o n , l i t e r a t u r e r e p o r t s a n d g r o w t h s t u d i e s i n t h e l a b -o r a t o r y . T e m p e r a t u r e : N o d u l a r i a was c o l l e c t e d d u r i n g t h i s s t u d y a t t e m p e r a t u r e s o f 15-30 C . R e p o r t s i n t h e l i t e r a t u r e g i v e a w i d e r a n g e o f v a l u e s f r o m 12.9-33 C ( F r a n c i s 1878; W e s t 1909; B r a n n o n 1911; S j o s t e d t 1922; D e l l o w 1955; Woods on e t al.1966; P r o s h k i n a - L a v r e n k o 1968; T o e t z 1973). T e m p e r a t u r e d o e s n o t a p p e a r t o l i m i t g r o w t h i n some c a s e s s i n c e S j o s t e d t (1922) r e p o r t s N. s p u m i g e n a f r o m t h e B a l t i c i n b l o o m q u a n t i t i e s a t 12.9 C . H o w e v e r more t y p i c a l l y l a r g e n u m b e r s o c c u r a t h i g h t e m p e r a t u r e s (20-30 C ) . N . h a r v e y a n a h a s b e e n r e p o r t e d f r o m " h o t " s p r i n g s ( D a n j o y i n B o r n e t a n d F l a h a u l t 1888) o r warm s p r i n g s ( W e l c h I964) b u t no t e m p e r a t u r e s w e r e g i v e n . T h e o n l y i n s t a n c e i n w h i c h t h e t e m p e r a t u r e o f a " h o t " s p r i n g was m e a s u r e d f o r a n y N o d u l a r i a c o l l e c t i o n i s 26 C ( G r o e s b e c k F 1049304, Mono C o . , C a l i f o r n i a ) , a n d t h i s i s w e l l b e l o w t h e maximum t e m p e r a t u r e o f 30-35 C f r o m l a b o r a t o r y s t u d i e s (p.45)« 106 T h e t e m p e r a t u r e f o r maximum g r o w t h f o r N o d u l a r i a s p p . a p p e a r s t o b e s i m i l a r t o some o t h e r N o s t o c a c e a e , i . e . , A n a b a e n a v a r i a -b i l i s a n d N o s t o c m u s c o r u m ( K r a t z a n d M e y e r s 1955a). T h e r e s t r i c t i o n o f N o d u l a r i a t o w a t e r s o f l e s s t h a n 30 C may p a r -t i a l l y e x p l a i n i t s a b s e n c e f r o m t r o p i c a l a r e a s . L i g h t : L i g h t i n t e n s i t y v a l u e s f o r g r o w t h o f N o d u l a r i a b o t h u n d e r f i e l d a n d l a b o r a t o r y c o n d i t i o n s a r e d i f f i c u l t t o a s s e s s . I n t h e i n l a n d s a l i n e p o n d s , t h e s h a l l o w w a t e r a n d t h e w h i t e s a l t a r o u n d t h e s h o r e l i n e c o n t r i b u t e t o t h e h i g h l i g h t i n t e n s i t y , b u t t h e l i g h t i n t e n s i t y i s r e d u c e d b y s h a d i n g o f o t h e r a l g a e ( C l a d o p h o r a o r C t e n o c l a d u s ) a n d b y t u r b i d i t y o f t h e w a t e r i n some l a k e s . T h e o n l y r e p o r t o f l i g h t i n t e n s i t y m e a s u r e d i n t h e f i e l d w i t h r e g a r d t o N o d u l a r i a i s t h a t o f D e l l o w a n d C a s s i e (1955) who d e s c r i b e a l g a l z o n a t i o n i n a s e a c a v e i n New Z e a l a n d . T h e y n o t e t h a t N_. h a r v e y a n a was r e s t r i c -t e d t o an a r e a w i t h l i g h t i n t e n s i t y o f 300-750 f t - c . T h i s i s t h e same r a n g e o f l i g h t i n t e n s i t y (200-600 f t - c ) w h i c h g a v e maximum g r o w t h r a t e s i n l a b o r a t o r y e x p e r i m e n t s (p.50). A s i n t h e f i e l d s t u d i e s , d i f f i c u l t i e s a r e e n c o u n t e r e d i n a s s e s s i n g t h e e f f e c t o f l i g h t o n b l u e g r e e n a l g a l c u l t u r e s . B e c a u s e o f s h a d i n g t h e l i g h t i n t e n s i t y a t a g i v e n d i s t a n c e f r o m t h e l i g h t s o u r c e i s t h a t r e c e i v e d o n l y b y c e l l s a t t h e s u r f a c e o f t h e c u l t u r e v e s s e l . C e l l s i n t h e c e n t e r o f t h e v e s s e l r e c e i v e l e s s l i g h t . Some c u l t u r e w o r k i n v o l v i n g b l u e g r e e n a l g a e 107 ( i . e . M c L a c h l a n a n d G o r h a m 1962) u s e d v e r y h i g h l i g h t i n t e n s -i t i e s (7500-9000 f t - c ) t o c o m p e n s a t e f o r h i g h c e l l d e n s i t i e s u s e d . I n t h e g r o w t h e x p e r i m e n t s d e s c r i b e d p r e v i o u s l y (p.36) d i l u t e i n o c u l u m was u s e d , s o t h a t t h e l i g h t i n t e n s i t y d i f f e r e d o n l y s l i g h t l y f r o m o n e s i d e o f t h e v e s s e l t o t h e o t h e r . T h e r e s u l t s show t h a t N o d u l a r i a s p p . g r o w u n d e r a r e l a t i v e l y w i d e r a n g e o f l i g h t i n t e n s i t i e s (2 5-600 f t - c ) b u t maximum r a t e s o f g r o w t h w e r e i n h i b i t e d b y l i g h t i n t e n s i t i e s a s h i g h a s d i r e c t s u n l i g h t . C u l t u r e s g r o w n w i t h d a y l i g h t a s t h e e n e r g y s o u r c e g r e w s l o w e r t h a n a n y o f t h e i n c u b a t o r g r o w n c u l t u r e s e x c e p t a t t h e v e r y l o w e s t l i g h t l e v e l (LL4). p H : P u b l i s h e d r e p o r t s i n d i c a t e t h a t N o d u l a r i a h a s b e e n c o l l e c t e d w h e r e pH v a l u e s r a n g e f r o m 7.0-9.35 ( D e l l o w 1955; H o r t o b a g y i 1959; B a y l y a n d W i l l i a m s I966; S e r p e t t e a n d L a b b e 1966; W o o d s o n e t a l . 1966; O r t e g a 1972; P l i n s k i 1973). T h e g e n u s i s much more commonly r e p o r t e d f r o m h a b i t a t s o f pH g r e a t e r t h a n 8, w i t h t h e c o l l e c t i o n s b e l o w pH 8 c o n t a i n i n g o n l y s m a l l n u m b e r s o f N o d u l a r i a . When l a r g e a m o u n t s a r e p r e s e n t , t h e pH i s i n v a r i a b l y a b o v e 8. D u r i n g t h i s s t u d y N o d u l a r i a was n o t c o l l e c t e d a t a pH o f l e s s t h a n 8.2, a l t h o u g h h a b i t a t s f r o m pH 5«2 t o 10.0 w e r e s u r v e y e d . T h i s r e s p o n s e t o pH i s a l s o r e f l e c t e d i n t h e l a b o r a t o r y s t u d i e s . A l l i s o -l a t e s o f N o d u l a r i a s h o w e d maximum g r o w t h r a t e s a t a pH g r e a t e r t h a n 9, w i t h some i s o l a t e s a s h i g h a s 10.4*. No g r o w t h 108 o c c u r r e d a t pH 6 a n d v e r y l i t t l e a t pH 7. T h e o n l y r e p o r t s o f b l u e g r e e n a l g a e s h o w i n g maximum g r o w t h a t a h i g h pH (10) i s b y G e r l o f f e t a l . (1952) a n d M c L a c h l a n a n d G o r h a m (1962) w i t h M i c r o c y s t i s a e r u g i n o s a . S a l i n i t y : T h i s a s p e c t o f t h e e c o l o g y o f N o d u l a r i a i s t h e m o s t t h o r o u g h l y d o c u m e n t e d q u a n t i t a t i v e l y ; h o w e v e r , many r e p o r t s d e s c r i b e c o l l e c t i o n s i n much more g e n e r a l t e r m s , i . e . , " e s p e c i a l l y h a r d w a t e r " ( P r e s c o t t 1962). I n t h e p r e s e n t s t u d y N o d u l a r i a i s o l a t e s w e r e c o l l e c t e d a t s a l i n i t i e s f r o m 3-65 ° / o o ( A p p e n d i x , T a b l e s II-VI). L i t e r a t u r e r e p o r t s show a w i d e r a n g e o f s a l i n i t i e s f o r c o l l e c t i o n o f N o d u l a r i a , f r o m 0.12-35.7 ° / o o ( L e v a n d e r 1900; M o o r e 1917; S j o s t e d t 1922; H u t c h i n s o n e t a_l. 1932; T r a h m s 1937; Rawson a n d M o o r e 1944; D e l l o w 1955; R a t h s a c k - K u t z e n b a c h 1961; K a l b e a n d T i e s s 1964; B a y l y a n d W i l l i a m s 1966; P r o s h k i n a - L a v r e n k o 1968; O r t e g a 1972; V e r c h a n d B l i n n 1972; P l i n s k i 1973). N o d u l a r i a h a s b e e n r e p o r t e d f r o m w a t e r s w i t h Na a n d Mg a s d o m i n a n t c a t i o n s a n d C I , HCO^ , C0^ a n d S0^ a s d o m i n a n t a n i o n s . T h e r e a r e no r e p o r t s o f N o d u l a r i a c o l l e c t i o n s w h e r e t h e d o m i n a n t i o n s a r e ++ + Ca o r K . T h i s may o n l y r e f l e c t t h e f a c t t h a t s a l i n e l a k e s o f d o m i n a n t Ca o r K a r e uncommon o n a w o r l d b a s i s . L a b o r -a t o r y o b s e r v a t i o n s w i t h d i f f e r e n t t y p e s o f d o m i n a n t i o n s ( p .55) show t h a t g r o w t h r a t e a n d s a l i n i t y t o l e r a n c e a r e n o t a f f e c t e d b y t h e d o m i n a n t a n i o n o r c a t i o n p r e s e n t . 1 0 9 T h e maximum l e v e l o f s a l i n i t y a t w h i c h m e a s u r a b l e g r o w t h o c c u r s a p p e a r s t o be a b o u t 60 ° / o o . F i e l d o b s e r v a t i o n s s h o w e d t h a t N o d u l a r i a was n o t p r e s e n t i f t h e s a l i n i t y r o s e a b o v e a p p r o x i m a t e l y 60 ° / o o . I n c u l t u r e 6 0 ° / o o was t h e u p p e r l i m i t o f g r o w t h f o r m o s t i s o l a t e s , a l t h o u g h some w e r e l e s s t o l e r a n t . H o f a n d F r e m y (1933) r e p o r t t h a t two i s o l a t e s o f N o d u l a r i a s h o w e d maximum t o l e r a n c e o f 47 ° / o o a n d 5 8 ° / o o . On a g r a d i e n t o f s a l i n i t y N o d u l a r i a g r o w s b e s t a t 5-20 ° / o o , r e f -l e c t i n g t h e m o s t common s a l i n i t i e s i n w h i c h i t o c c u r s i n n a t u r e . T h u s , t e c h n i c a l l y t h e i s o l a t e s w e r e n o t " f r e s h w a t e r " o r g a n i s m s . B a t t e r t o n a n d v a n B a a l e n (1971) n o t e d d i f f e r e n t p a t t e r n s o f g r o w t h o f f r e s h w a t e r a n d m a r i n e b l u e g r e e n i s o l a t e s when g r o w n o n s a l i n i t y g r a d i e n t s . T h e f r e s h w a t e r i s o l a t e g r e w b e s t i n t h e b a s a l medium (1 ° / o o ) w i t h g r o w t h d e c l i n i n g w i t h a n y a d d -i t i o n a l s a l t s . T h e m a r i n e i s o l a t e s g r e w b e t t e r a t 10-20 ° / o o s a l i n i t y ( N a C l ) t h a n i n t h e b a s a l m e d i u m . N i t r o g e n : T h e r e i s l i t t l e c o r r e l a t i o n o f n i t r o g e n s o u r c e o r l e v e l s w i t h t h e p r e s e n c e o r a b s e n c e o f N o d u l a r i a i n n a t u r e . S j o s t e d t (1922) s t a t e d t h a t t h e g r o w t h o f N o d u l a r i a was e n h a n c e d b y s e w a g e o u t f l o w s b u t p r e s e n t e d no d a t a t o s u p p -o r t t h i s v i e w . T h e l a b o r a t o r y r e s u l t s ( p .55) show t h a t a l l i s o l a t e s g r e w a s w e l l w i t h v a r i o u s l e v e l s o f N0^ (0.25-1 . 0 g/1) a s i n medium l a c k i n g a n y c o m b i n e d n i t r o g e n s o u r c e . T h e i n -a b i l i t y o f N o d u l a r i a t o g r o w w i t h N H . + o r u r e a a s n i t r o g e n 110 s o u r c e i s i n t e r e s t i n g , h o w e v e r t h e l e v e l s (0.5-1.0 g / l ) may h a v e b e e n t o o h i g h f o r g r o w t h , s i n c e t h e l e v e l s i n n a t u r e a r e much l o w e r (1.0-6.1 m g / l ) . K r a t z a n d M e y e r s (1955a) r e p o r t t h a t A n a b a e n a v a r i a b i l i s a n d N o s t o c m u s c o r u m g r e w w e l l o n s i m -i l a r c o n c e n t r a t i o n s (0.5 g / l o f NH^ a n d u r e a ) t o t h o s e u s e d w i t h N o d u l a r i a i n t h i s s t u d y . S t e w a r t (I965) r e p o r t e d t h a t N o d u l a r i a p r e s u m a b l y f i x e d n i t r o g e n b u t b e c a u s e t h e c u l t u r e s w e r e i m p u r e , d e f i n i t e p r o o f was l a c k i n g . T h e p r e s e n t s t u d y shows t h a t t h e two i s o l a t e s t e s t e d f i x n i t r o g e n i n p u r e c u l t u r e a n d c i r c u m s t a n t i a l e v i d e n c e ( g r o w t h i n m e d i u m w i t h o u t c o m b i n e d n i t r o g e n ) was s e e n f o r t h e a b i l i t y o f t h e o t h e r i s o l a t e s t o f i x n i t r o g e n . T h e o n l y d a t a o n t h e e f f e c t o f e n v i r o n m e n t a l p a r a m e t e r s o n n i t r o g e n f i x a t i o n i s t h a t r e p o r t e d b y J o n e s a n d S t e w a r t (1969) f o r a m a r i n e C a l o t h r i x i s o l a t e . T h e y r e p o r t e d h i g h e s t r a t e s o f n i t r o g e n f i x a t i o n a t 800 f t - c , a t 30 C , w i t h a s a l i n i t y o f 5 ° / o o ( b u t f i x a t i o n t o o k p l a c e a t 1-50 ° / o o ) a n d pH 8.5. E x c e p t f o r t h e pH f i g u r e t h e s e d a t a a r e n e a r l y i d e n t i c a l w i t h t h e r e s u l t s o b t a i n e d w i t h N o d u l a r i a (pH 10). C o n s i d e r i n g t h e e n v i r o n m e n t a l p a r a m e t e r s t a k e n i n t o a c c -o u n t a n d l a b o r a t o r y o b s e r v a t i o n s m a d e , some g e n e r a l i z a t i o n s c a n b e made r e g a r d i n g t h e b i o l o g y o f N o d u l a r i a . S a l i n i t y ( a n d t h e r e l a t e d p a r a m e t e r o f pH) i s p r o b a b l y t h e m o s t i m p o r t -a n t a s p e c t o f t h e e c o l o g y o f t h e g e n u s . T h e a l g a e , b o t h I l l s p e c i e s , a r e t y p i c a l l y f o u n d i n b r a c k i s h o r s a l i n e w a t e r s (5-30 ° / o o ) a n d a l t h o u g h t h e y may o c c u r i n s a l i n i t i e s l o w e r o r h i g h e r t h a n t h e s e v a l u e s t h e y a r e u s u a l l y p r e s e n t i n much r e d u c e d a m o u n t s . N o d u l a r i a s p p . a l s o seem t o t o l e r a t e s u d d e n o r g r a d u a l c h a n g e s i n s a l i n i t y . T h i s i s e v i d e n t f r o m f i e l d o b s e r v a t i o n s ( i n t h e p o n d s i n t h e B r i t i s h C o l u m b i a i n t -e r i o r s a l i n i t y r o s e f r o m 15 t o 65 °/00 i n a f e w w e e k s ) a n d l a b o r a t o r y s t u d i e s ( i n o c u l u m g r o w n a t 1 ° / o o g r e w w i t h o n l y a s h o r t l a g p e r i o d when i n t r o d u c e d i n t o medium w i t h d i s s o l v e d s a l t s o f 40 ° / o o ) . Some l i t e r a t u r e r e p o r t s i n d i c a t e N o d u l -a r i a s p p . l i v e i n c o n d i t i o n s w h e r e m a r k e d s a l i n i t y c h a n g e s o c c u r , s u c h a s s e a c o a s t p o o l s w h e r e r a i n f a l l a n d s u r f a l t e r -n a t e l y a f f e c t t h e s a l t c o n t e n t o f w a t e r ( C o l l i n s 1908 f r o m M a i n e ; F j e r d i n g s t a d 1969 f r o m D e n m a r k ) . L o c a t i o n s w h e r e N o d -u l a r i a h a s b e e n r e c o r d e d i n b l o o m q u a n t i t i e s , w h i c h may b e a s s -umed t o b e f a v o r a b l e c o n d i t i o n s ( o r n e a r f a v o r a b l e ) , a r e i n w a t e r s w h i c h a r e e i t h e r b r a c k i s h o r s a l i n e . B l o o m s i t u a t i o n s r e p o r t e d i n t h e l i t e r a t u r e w h e r e s a l i n i t y was r e c o r d e d a r e M o o r e (1917) i n D e v i l s L a k e , N o r t h D a k o t a a t 9.5 ° / o o ; S j o s -t e d t (1922) i n t h e B a l t i c S e a a t 14•5 ° / o o ; H u t c h i n s o n e t a l . (1932) i n S o u t h A f r i c a a t 21 ° / o o ; K a l b e a n d T i e s s (I964) f r o m t h e B a l t i c S e a a t 2.3-2.5 ° / o o ; a n d B a y l y a n d W i l l i a m s (I966) f r o m L a k e C o r a n g a m i t e , A u s t r a l i a a t 29.1 ° / o o . E a r l y r e c o r d s o f b l o o m s do n o t s t a t e s a l i n i t y v a l u e s , b u t f r o m t h e l o c a t i o n s 112 i t c a n b e i n f e r r e d t h a t t h e w a t e r s w e r e b r a c k i s h : F r a n c i s (1878) t h e t i d a l L a k e A l e x a n d r i n a , S o u t h A u s t r a l i a ; B o r n e t a n d T h u r e t a t D e a u v i l l e , F r a n c e ( c i t e d i n S j o s t e d t 1922); B o r n e t a n d F l a h a u l t (1888) c i t e b l o o m r e p o r t s b y S c h m i t z , S u h r a n d H o f m a n - B a n g a l o n g t h e B a l t i c c o a s t . Two b l o o m s ( K a l b e a n d T i e s s 1964 o n t h e German B a l t i c c o a s t ; F r a n c i s I878) w e r e t o x i c a n d r e s u l t e d i n d e a t h o f l i v e s t o c k a n d p o u l -t r y . L i g h t a n d t e m p e r a t u r e p o s s i b l y p l a y a l e s s e r r o l e t h a n s a l i n i t y a n d pH i n c o n t r o l l i n g g r o w t h b u t a more i m p o r t a n t r o l e i n d i s t r i b u t i o n . T h e a b s e n c e o f N o d u l a r i a s p p . f r o m t r o p i c a l a r e a s c o u l d b e b e t t e r e x p l a i n e d i n t e r m s o f l i g h t a n d t e m p e r a t u r e s i n c e N o d u l a r i a s p p . a r e a b s e n t f r o m i n l a n d s a l i n e l a k e s a n d b r a c k i s h a r e a s i n t h e t r o p i c s . I n t r o p i c a l i n l a n d s a l i n e s i t u a t i o n s A r t h o s p i r a o r S p i r u l i n a a r e t h e d o m i n a n t b l u e g r e e n a l g a e ( A r m i t a g e 1971; J e n k i n 1932; R i c h 1932). T h e s e t w o f a c t o r s w o u l d b e o b v i o u s d i f f e r e n c e s b e t w e e n b r a c k i s h o r s a l i n e a r e a s i n t e m p e r a t e a n d t r o p i c a l a r e a s . L i g h t i s a d i f f i c u l t f a c t o r t o m e a s u r e i n n a t u r e s i n c e t u r b i d i t y o f t h e w a t e r , s h a d i n g b y o t h e r p l a n t s , d a y l e n g t h a n d l a t i t u d e a l l a f f e c t t h e amount o f l i g h t r e c e i v e d . T h e p r e s e n c e o f N o d u l -a r i a s p p . a t h i g h e r a l t i t u d e s i n t h e t r o p i c s i n d i c a t e s t h a t t e m p e r a t u r e i s p o s s i b l y a more i m p o r t a n t f a c t o r t h a n l i g h t . 113 T a x o n o m y : T h e r e e x i s t s a p r o b l e m w i t h b l u e g r e e n a l g a e i n t h a t t h e t e r m " s p e c i e s " i s n o t a p p l i c a b l e i n t h e same s e n s e a s i n h i g h e r o r g a n i s m s s i n c e t h e y a r e p r o k a r y o t e s . T h e t e r m h a s l i t t l e s i g n i f i c a n c e i n b l u e g r e e n a l g a e o t h e r t h a n i t s u r r o u n d s a n a r b i t r a r y g r o u p o f o r g a n i s m s w i t h c e r t a i n c h a r a c t e r i s t i c s a n d p o s s e s s i n g a n u n d e f i n e d amount o f v a r i a b i l i t y . K n o w l e d g e o f t h e v a r i a b i l i t y i s e s s e n t i a l t o t h e t a x o n o m y . T h e v a r i a b i l i t y w i t h i n a c e r t a i n " p o p u l a t i o n " i s more e a s i l y d e s c r i b e d t h a n t h e v a r i a b i l i t y among " p o p u l a t i o n s " f o r b l u e g r e e n a l g a e . A p o p u l a t i o n i s d e f i n e d i n t h e s e n s e o f a c o l l e c t i o n o f i n d i v i d -u a l s o f o n e s p e c i e s l i v i n g w i t h i n a d e f i n e d a r e a o r v o l u m e ( H u t c h i n s o n 196 7). T h e e x t e n t o f v a r i a b i l i t y i n b l u e g r e e n a l g a e h a s b e e n o f c o n c e r n f o r many y e a r s ( f o r e x a m p l e - Crow 1924; C a n a b a e u s 1929; D r o u e t 1962, 1963; F j e r d i n g s t a d I 9 6 6 , 1969) a n d w i t h o u t i n s i g h t i n t o t h i s v a r i a b i l i t y , a w o r k a b l e t a x o n o m i c s c h e m e i s i m p o s s i b l e . I t i s a l s o a p p a r e n t o b s e r -v a t i o n s o f n a t u r a l m a t e r i a l i s n o t s u f f i c i e n t a n d t h a t c u l t u r -a l w o r k i s n e c e s s a r y f o r b l u e g r e e n a l g a e ( K o m a r e k 1971). T h e l i t e r a t u r e i s a n o f t e n n e g l e c t e d a n d m i s u s e d a s p e c t o f s t u d y . A g r e a t d e a l o f d a t a ( o n v a r i a b i l i t y , e c o l o g y , d i s t r i b u t i o n ) c a n b e o b t a i n e d f r o m t h e a c c u m u l a t e d o b s e r v a t -i o n s o f many w o r k e r s . T h e l i t e r a t u r e i s m i s u s e d i n t h e s e n s e t h a t t h e t a x o n o m i c c o n f u s i o n i s i n c r e a s e d b y new s p e c i e s 114 o r v a r i e t y d e s c r i p t i o n s w i t h l i t t l e j u s t i f i c a t i o n . A l s o , N o d u l a r i a s p e c i e s h a v e b e e n d e s c r i b e d w i t h o u t t y p e m a t e r i a l b e i n g r e t a i n e d , o r d e s c r i b e d w i t h i n c o m p l e t e m a t e r i a l ( w i t h -o u t a k i n e t e s ) , o r w i t h i n a d e q u a t e m a t e r i a l ( e . g . N . s p u m i g e n a v a r . a e r o p h i l a , p.80 was d e s c r i b e d f r o m a s i n g l e f i l a m e n t l a c k i n g a k i n e t e s ) . A n o t h e r s i m i l a r p r o b l e m i s i m p r o p e r i d e n t i f i c a t i o n o f i s o l a t e s u s e d i n p h y s i o l o g i c a l a n d b i o -c h e m i c a l i n v e s t i g a t i o n s . T h e p r o b l e m w i t h A n a c y s t i s n i d u l a n s h a s a l r e a d y b e e n m e n t i o n e d ( p . l ) . F o r e x a m p l e , t h e s t r a i n 11.1.1 o f " N o d u l a r i a s p h a e r o c a r p a " i s o l a t e d b y M . B . A l l e n a n d i d e n t i f i e d b y D r o u e t i s e v i d e n t l y n o t a N o d u l a r i a . M a t -e r i a l f r o m t h i s o r i g i n a l c u l t u r e (F N o r d i n 1181) a s w e l l a s c u l t u r e s f r o m I n d i a n a U n i v e r s i t y C u l t u r e C o l l e c t i o n (#583) a n d t h e c u l t u r e c o l l e c t i o n a t t h e D e p a r t m e n t o f B a c t e r i o l o g y a n d I m m u n o l o g y B e r k e l e y C a l i f o r n i a ( s t r a i n 7103) d e r i v e d f r o m t h e o r i g i n a l c u l t u r e , w e r e e x a m i n e d . T h e m a t e r i a l shows t e r m i n a l h e t e r o c y s t s a n d b a s a l d i f f e r e n t i a t i o n c h a r a c t e r i s t i c s o f t h e R i v u l a r i a c e a e a n d i s a C a l o t h r i x s p . T h i s same c o n -c l u s i o n was a l s o r e a c h e d b y K e n y o n e t a l . (1972) s t u d y i n g s t r a i n 7103- T h e same i s o l a t e ( N . s p h a e r o c a r p a ) h a s b e e n u s e d i n i n v e s t i g a t i o n s b y E d e l m a n e t a_l. (1967), G o r h a m (i960) a n d G o r h a m (1964). T h e N o d u l a r i a s p h a e r o c a r p a , ( C u l t u r e B-1466) i n t h e G o t t i n g e n c o l l e c t i o n i s a l s o a s u b c u l t u r e o f A l l e n ' s i s o l a t e ( K o c h I964). G r a n h a l l a n d H e n d r i c k s o n (1969) 115 i s o l a t e d an a l g a w h i c h t h e y i d e n t i f i e d a s N . s p u m i g e n a . A s u b c u l t u r e o f t h i s i s o l a t e was e x a m i n e d a n d h a s t h e c h a r a c t e r -i s t i c s o f a N_. h a r v e y a n a w i t h f i l a m e n t s 5 ym w i d e . T h e i s o -l a t e u s i n g t h i s i n c o r r e c t s p e c i f i c name was a l s o c i t e d b y G r a n h a l l a n d B e r g (1972). T h e g e n u s N o d u l a r i a a p p e a r s t o be a f a i r l y s t a b l e t a x o n m o r p h o l o g i c a l l y . S t u d i e s w i t h o t h e r b l u e g r e e n a l g a e i n d i c a t e t h a t g e n e r i c c h a r a c t e r i s t i c s may v a r y a n d m a t e r i a l may h a v e t h e c h a r a c t e r i s t i c s o f two d i f f e r e n t g e n e r a u n d e r d i f f e r e n t c u l t u r e c o n d i t i o n s ( S t e i n 1963; P e a r s o n a n d K i n g s b u r y 1966). C e r t a i n c h a r a c t e r i s t i c s o f N o d u l a r i a v a r y l i t t l e u n d e r t h e w i d e r a n g e o f g r o w t h c o n d i t i o n s u s e d ( l i g h t , t e m p e r a t u r e , p H , s a l i n i t y ) . T h e c e l l s a r e a l w a y s w i d e r t h a n l o n g ( e x c e p t i n IJ. h a r v e y a n a p r i o r t o c e l l d i v i s i o n ) . T h e h e t e r o c y s t s a r e r e g u l a r l y s p a c e d a l o n g t h e f i l a m e n t a n d f a i r l y c l o s e t o g e t h e r ( e v e r y 20-30 c e l l s ) , a c h a r a c t e r i s t i c e m p h a s i z e d b y b o t h T h u r e t (1875) a n d F r i t s c h (1951). T h e a k i n e t e s a r e f o r m e d midway b e t w e e n t h e h e t e r o c y s t s a n d m a t u r e f r o m t h e c e n t e r o f t h e t r i c h o m e t o w a r d s t h e h e t e r o c y s t . A g a i n t h i s c h a r a c t e r -i s t i c i s d e s c r i b e d i n d e t a i l b y F r i t s c h (1951). H o w e v e r o t h e r m o r p h o l o g i c a l c h a r a c t e r i s t i c s a r e v a r i a b l e . T h e s h e a t h i s u s u a l l y p r e s e n t i n f i e l d m a t e r i a l (more s o N. s p u m i g e n a t h a n N . h a r v e y a n a ) ; b u t i t i s n o t a l w a y s p r e s e n t a n d i n c u l t u r e t h e p r e s e n c e o f t h e s h e a t h i s e v e n more v a r i a b l e , 116 b e i n g a b s e n t much m o r e o f t h e t i m e . F o e r s t e r (1964) c i t e s C a a n d Mg h a r d n e s s a s a f a c t o r i n s h e a t h f o r m a t i o n i n O s c i l l a t o r i a l i m o s a ( R o t h ) C . A . A g . F i l a m e n t l e n g t h v a r i e s a g r e a t d e a l . I n n a t u r e f i l a m e n t s may v a r y f r o m a f e w d o z e n c e l l s t o s e v e r a l h u n d r e d . T h e l a t t e r c o n d i t i o n i s m o r e p r e v -a l e n t i n c u l t u r e . V e g e t a t i v e c e l l c o l o r v a r i e s a g r e a t d e a l b o t h i n n a t u r e a n d i n c u l t u r e . T h e c o l o r i s t y p i c a l l y a b r i g h t b l u e - g r e e n b u t may b e b r o w n , l i g h t g r e e n o r b l u e . T h e a k i n e t e a s p r e v i o u s l y m e n t i o n e d i s v e r y v a r i a b l e . B e c a u s e a k i n e t e s a r e f o r m e d f r o m v e g e t a t i v e c e l l s , t h e r e a r e a v a r i e t y o f s t a g e s i n f o r m a t i o n ; s i z e , s h a p e , c o l o r a n d t e x -t u r e a l l d e p e n d u p o n t h e m a t u r i t y o f t h e a k i n e t e . T h e s i z e was t h e o n l y a s p e c t w h i c h c o u l d b e c o r r e l a t e d w i t h a n y p h y s i c a l o r c h e m i c a l p a r a m e t e r . T h e m u l t i p l i c i t y o f s p e c i e s o f N o d u l a r i a i n t h e l i t e r a t u r e a p p e a r s t o b e more a r e s u l t o f p o o r j u d g e m e n t i n t h e n a m i n g o f s p e c i e s a n d m i s u n d e r s t a n d i n g t h e e x t e n t o f v a r i a b i l i t y , t h a n a n y e x t r e m e p l a s t i c i t y o f t h e o r g a n i s m . T h e a p t n e s s o f g e n e r i c d i s t i n c t i o n s o f many C y a n o p h y t a ( e s p e c i a l l y t h e O s c i l l a t o r -i a c e a e s e n s u G e i t l e r ) h a s b e e n q u e s t i o n e d ( D r o u e t I968; B o u r -e l l y 1970). W h i t t o n (1962) i n e v a l u a t i n g t h e t a x o n o m i c c r i t -e r i a f o r s e v e r a l b l u e g r e e n a l g a e commented t h a t C y l i n d r o s p e r n m m s p e c i e s r e p r e s e n t e d a d i s t i n c t g r o u p i n g o f f o r m s ; w h e r e a s , A n a b a e n a was a v e r y a m o r p h o u s g r o u p a n d t h a t g e n e r a i n t h e 117 O s c i l l a t o r i a c e a e w e r e v e r y a r b i t r a r y . N o d u l a r i a , t h u s a p p -e a r s t o b e s i m i l a r t o C y l i n d r o s p e r m u m i n i t s d i s t i n c t i v e n e s s . T h u s i t m u s t b e e m p h a s i z e d t h a t t h e s t a b i l i t y a n d d i s t i n c t -n e s s o f t h e t a x a d e p e n d t o a g r e a t e x t e n t o n s e l e c t i n g t h e p r o p e r t a x o n o m i c c r i t e r i a . 118 L I T E R A T U R E C I T E D A c k l e y , A . B . 1931. P r e l i m i n a r y r e p o r t o f M i c h i g a n a l g a e . M i c h . A c a d . S c i . A r t . L e t . 15: l - 4 9 « A l i , S . a n d S a n d h u , G . R . 1972. B l u e - g r e e n a l g a e o f t h e s a l i n e s o i l s o f t h e P u n j a b . O i k o s 23: 268 -272 . A l l e n , M . M . 1973. M e t h o d s f o r C y a n o p h y c e a e . i n S t e i n , J . R . , e d . , H a n d b o o k o f P h y c o l o g i c a l M e t h o d s C a m b r i d g e U n i v e r s i t y P r e s s , C a m b r i d g e 448 p p . A m e r i c a n P u b l i c H e a l t h A s s o c i a t i o n . 1965« S t a n d a r d M e t h o d s f o r t h e E x a m i n a t i o n o f W a t e r a n d S e w a g e . 1 2 t h . e d . A . P . H . A . New Y o r k . 769 p p . A n a g n o s t i d i s , K . a n d S c h w a d e , G . H . 1966. U b e r a r t e n r e i c h e B e s t a n d e v o n C y a n o p h y t e n u n d B a c t e r i o p h y t e n i n e i n e m F a r -b s t r e i f e n s a n d w a t t s o w i e i i b e r d a s A u f t r e t e n G o m o n t i e l l a -A r t i g d e f o r m i e r t e r O s c i l l a t o r i a t r i c h o m e . Nova H e d w i g i a 11 : 417 -441 . A n d e r s o n , E . M . a n d W a l k e r , E . R . 1920 A n e c o l o g i c a l s t u d y o f t h e a l g a e o f some s a n d h i l l l a k e s . T r a n s . A m e r . M i s c r o s c . S o c . 39: 51 -85 . A r m i t a g e , K . B . 1971 . A h i g h l y a l k a l i n e l a k e i n N i c a r a g u a . H y d r o b i o l o g i a 38: 437 -439 -A r n d t , E 0 A . , P a n k o w , H . a n d K e l l , V . I 9 66 . l i b e r d a s P h y t o -p l a n k t o n d e r W i s m a r - B u c h t . I n t . R e v . g e s . H y d r o b i o l . H y d r o g r . 51: 127 -146 . B a j k o v , A . 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Stat. 116 pp. Zhadin, V.I. and Gerd, S.V. 1961. Fauna and f l o r a of the r i v e r s , lakes and reservoirs of the U.S.S.R. I s r a e l Prog. S c i . Trans., Jerusalem, No. 1903, 626 pp. APPENDIX TABLES I 138 Table I. Procedures Used i n Water Analysis Na +, K +, Ca , Mg'' : Perkin-Elmer Atomic Absorption Spectrophotometer Model 303. S 0 ^ : Turbidimetric method (Standard Methods) American Public Health Association, I965. Cl : Argentometric method (Standard Methods) American Public Health Association, I965. HCO^ and C O 3 : Potentiometric t i t r a t i o n (Standard Methods) American Public Health Association, I965. Ortho-phosphate : Stannous Chloride method (Standard Methods) American Public Health Association, I965. NH.+ : N e s s l e r i z a t i o n method (Standard Methods) *4 American Public Health Association, I965. N 0 ^ • Phenoldisulfonic Acid method (Standard Methods) American Public Health Association, 1965. T o t a l Dissolved Solids Evaporation at 105 C (Standard Methods) American Public Health Association, I965. Table I I . Water chemistry f o r Ctenocladus Pond 30v72 2 1 v i i 7 2 2 3 v i i i 7 2 2 4 i x 7 2 2 8 i v 7 3 3 v i 7 3 2 1 v i i 7 3 pH 9.1 9.0 8.8 8.4 8. 8 8.9 8.6 temp C 27 24 24 8 14 27 29 Na+mg/1 8100 42500 56000 72000 1 3 7 0 0 36000 95000 Mg++mg/1 1800 5200 9700 12600 1200 4 2 0 0 9200 Ca^mg/l 56 144 172 209 65 87 87 K +mg/l 280 865 1235 1024 193 522 835 S04"mg/1 23500 108000 130000 170000 4 2 0 0 0 1 0 0 0 0 0 250000 C0~mg/1 0 100 480 1240 40 640 620 HCO~ mg/1 0 40 580 260 330 240 380 Cl~mg/1 650 1750 5000 6400 600 1 5 0 0 550 NH 4 + mg/l 16 12 12 14 10 12 14 N0~ mg/1 4.1 1.2 3.4 3.0 3. 5 3.0 3.0 o-PO^mg/l 1.5 2.2 2.0 2.2 2. 0 3.1 3.5 TDS g/1 35 180 226 306 48 140 408 Table I I I . Water chemistry for Wallender Lake 30v72 21vii72 23viii72 24ix72 28iv73 3vi73 21vii7 PH 8.7 8.65 8.5 8.85 8.5 8.9 8.8 temp C 22 24 25 9 14 24 23 Na+mg/1 3200 6400 7700 9900 3700 7100 8000 ++ Mg mg/1 2900 3800 4300 2800 600 1600 5600 „ ++ /, Ca mg/1 74 119 250 237 96 151 285 K +mg/l 220 707 570 1031 404 374 921 S0~ mg/1 20000 31000 41000 46000 12000 22000 43000 C0~ mg/1 70 160 300 300 0 140 180 HC0~ mg/1 25 160 220 200 0 150 760 CI"mg/1 550 650 700 800 300 550 800 NH+ mg/1 10.2 8.8 8.6 9.5 9.5 8.0 6.3 N0~ mg/1 2.8 1.9 1.7 0.7 1.9 0.6 0.7 o-PO^ mg/1 0.4 0.2 0.4 0.1 0.2 0.3 0.2 TDS g / l 25 48 70 88 14 30 64 Table IV. Water chemistry f o r Inks Lake 30v72 21vii72 23viii72 24ix72 28iv73 3vi73 21vii73 pH 8.9 8.8 8.8 9.0 8.6 8.8 9.1 temp C 20 25 26 9 15 24 23 Na +mg/l 9800 11300 11800 8400 5500 8800 9800 Mg^mg/1 1600 1950 2400 4000 900 2200 5800 Ca^mg/l 121 148 133 201 60 144 173 K +mg/l 770 1030 1360 1220 390 1260 1390 S 0 ~ mg/1 27000 35000 37500 44000 14000 34000 46000 CO^ mg/1 25 70 140 180 0 100 380 HCO~ mg.l 80 310 550 580 0 240 720 C l " mg/1 550 650 700 800 300 550 800 NH+ mg/1 8.8 8.2 7.2 6.0 10.4 8.2 6.5 NO^ mg/1 3.1 1.9 2.1 1.0 2.1 0.8 0.8 o-PO, mg/1 4 0.3 0.2 0.2 0.1 0.2 0.3 0.1 TDS g/1 35 52 58 66 18 44 65 Table V. Water chemistry f o r White Lake 29v72 20vii72 23viii72 23ix73 27±v73 2vi73 PH 10.0 9.5 no c o l l e c t i o n 9.4 9.7 9.7 temp C 25 23 14 11 22 Na +mg/l 3000 7200 9100 3800 7500 Mg + +mg/l 1050 2000 3980 1400 2100 Ca^mg/l 455 350 320 0 0 K +mg/l 131 219 635 420 881 S0~ mg/1 11000 21000 32000 10500 23000 CO^ mg/1 760 500 340 1380 1500 HC03" mg/1 400 750 280 1050 490 C l ~ mg/1 600 1200 1000 900 1700 NH* mg/1 20.4 15.4 12.0 26.5 12.0 N0~ mg/1 2.1 1.0 1.1 2.8 2.2 o-PO. mg/1 4 0.2 0.3 0.1 0.3 0.3 TDS g / l 15 27 40 14 27 Table VI. Water chemistry f o r the lakes i r r e g u l a r l y sampled or sampled only once. Sa l t Pond 4 Salt Pond 3 Big Q u i l l Lake Devils Lake Spotted Lake R i e f e l Refuge San d . Fuca 39v72 21vii72 23viii72 24viii72 29v72 241x73 20v73 PH 9.9 9.3 8.6 9.1 7.8 8.2 8.4 temp C 30 26 25 21 33 19 18 Na +mg/l 11400 2200 10300 1475 34100 2675 5370 Mg^mg/l 41 30 3400 384 17000 320 651 1 | Ca mg/l 404 0 126 56 6.5 92 200 K +mg/l 128 55 560 175 1610 90 192 S0~ mg/l 14500 1330 35000 3600 140000 1120 . 2304 C0= mg/l 8300 963 130 120 120 0 10 HCO^ " mg/l 2000 312 320 250 110 29 61 C l " mg/l 5500 612 5500 650 1200 4620 9690 NH+ mg/l 2.5 0.9 4.5 0.4 5.0 0.1 0 NO" mg/l 3.0 2.2 2.3 6.1 4.0 1.4 0.5 0-P04 mg/l 3.5 1.9 0.2 1.2 4.5 0.2 0.1 TDS g/1 47 4 26 12 166 8.4 18 144 T a b l e V I I . W o r l d d i s t r i b u t i o n o f N o d u l a r i a * N . h a r v e y a n a N. s p u m i g e n a b o t h - i d e n t i f i c a t i o n o n l y t o g e n u s CANADA B r i t i s h C o l u m b i a * * B l i n n (1969) K a m l o o p s *-::-Sismey (1922) no l o c a t i o n * - S i smey (1922) P e n t i c t o n - S t o c k n e r e t a l . (1972) S k a h a L . O k a n a g a n L . V a s e u x L. M a n i t o b a **Bajkov (1935) s e v e r a l l o c a t i o n s S a s k a t c h e w a n *-"-Kuehune (1941) 7 l o c a t i o n s i n s o u t h c e n t r a l S a s k . -"-"-Rawson & M o o r e (1944) 8 l o c a t i o n s i n s o u t h c e n t r a l S a s k . U . S . A . A l a s k a -"•Gardner 3963 S i t k a UC 661526 F 1047757 A r i z o n a -"-Cameron (I963) -"--"-Cameron (I963) * * T a y l o r & C o l t o n Yuma C o . & G i l a C o . C o c h i s e C o . (1928) C o c o n i n o C o . A r k a n s a s -"-Demarree & T h o m a s o n 24582 D r e w C o . F 1133627 C a l i f o r n i a -xx-Drouet (1943) Modoc C o . -»-Drouet & M c B r i d e 4569 Mono C o . & S a n B e r n a d i n o C o . UC 664554 FH N o r d i n 1076 NBV 3925-43 NY N o r d i n 1268 F 1101885 ( D r o u e t 1943) ^ -Ga rdne r 989 ( PBA IO63) M a r i n Co 1088 - " G a r d n e r - " -Gardner ^ - "Ga rdne r - " G a r d n e r UC 341420 FH N o r d i n NBV 3925-48 F 980816 NY N o r d i n 1282 PC N o r d i n 1314 1498 O a k l a n d UC 202866 F N o r d i n 1182 1604 B e r k e l e y UC 202865 UC 274104 3295 O a k l a n d UC II4844 4151 O a k l a n d UC 66I636 FH N o r d i n 1078 NBV 3925-42 F 1032981 NY N o r d i n 1266 145 ^Gardner 6552 Berkeley UC 440248 •^Gardner 6568 Berkeley UC 661705 FH Nordin 1075 NBV 3925-46 F 1033552 NY Nordin 1264 ---^Gardner 7231 San Mateo Co. FH Nordin 1081 ^-Gardner 7670 Berkeley UC 641592 FH Nordin 1080 NBV 3925-40 F 1036391 NY Nordin 1270 *-*Gardner 7946 Contra Costa Co. UC 661724 NBV 3925-56 FH Nordin 1056 F 1043544 NY Nordin 1293 ^-Gardner 7963 San Francisco UC 661654 FH Nordin 1077 NBV 3925-41 F 1034223 NY Nordin 1267 *-Groesbeck 90 Mono Co. & San Bernadino Co. F 1049304 -"Hof & Fremy (1933) Marina *Hollenberg 1553 Orange Co. UC 634466 **0sterhout 559 Alameda Co. UC 393931 UC 202712 NBV 3925-54 FH Nordin 1066 PC Nordin 1328 -"-"-Osterhout & Gardner (PBA 1061) Oakland FH Nordin IO87 NBV 3925-30 F 980816 NY Nordin 1263 PC Nordin 1302 * * S e t c h e l l 1628 San Benito Co. UC 752513 F 1221362 #Setchell 1679 San Francisco UC I O O 5 6 3 *-x-Setchell & Jepson Stanislaus-Mariposa Co. UC 202 713 NBV 3925-53 Colorado *Louderback 17 Denver F 1235673 Connecticut **Holden 1458-1461 Bridgeport FH Nordin 1062 NY Nordin 1290 ( C o l l i n s 1900, 1905) F l o r i d a *Brannon 141A G a i n e s v i l l e F 1124079 PC Nordin 1329 (Brannon 1952) -"-Drouet, Madsen & Crowson 115H Wakulla Co. F 1324385 ^Drouet, Madsen & Crowson 11513 Wakulla Co. UC 912044 F 1324476 *Drouet & Nielsen 11232 Taylor Co. UC 912045 F 1308871 ---Drouet & Nielsen 11681 F r a n k l i n Co. UC 912043 F 1317616 -"-Standley 73311 Lee Co. F 1030027 I l l i n o i s -"Drouet, Glassman & Chapp 12696 Cook Co. F Nordin 1202 *Drouet, Glassman & Chapp 12711 Cook Co. F Nordin 1203 -x--::--x-Tiffany & B r i t t o n (1952) no location -x-x-x-T ran seau 63 Coles Co. NY Nordin 1283 ^Velasquez, Richards & Drouet 2507 Cook Co. F 980549 146 Indiana **Palmer (1928) Knox Co. *Palmer et a l . 2518 Indianapolis F 980816 **Transeau (1913) Decker Iowa -x-x-Prescott (1931) Dickenson Co. Kentucky -"Mclnteer (1939) no lo c a t i o n Louisiana ^-Drouet 8766 Calcasieu Parish UC 910404 NBV 3925-45 F 1332708 PC Nordin 1311 Maine , ^ C o l l i n s (PBA 1062) Casco Bay F 545439 NY Nordin 1280 PC Nordin 1307 ---Collins 22 Cape Rosier UC 687705 FH Nordin 1072 NBV 3925-39 F 1209180 NY Nordin 1278 PC Nordin 1310 -"-Collins 2387 Cape Rosier FH Nordin 1070 FH Nordin 1073 NY Nordin 1275 * * C o l l i n s 2439 Cape Rosier FH Nordin 1060 NY Nordin 1287 -"-•'"-Collins 3459 Cape Rosier NY Nordin 1289 ^-Collins 5094 Casco Bay UC 752510 F 1219372 NBV 3925-27 * * C o l l i n s 5094 Casco Bay UC 752514 F 1219336 *-::-Collins 5526 (PBA 1307) Harpswell UC 693286 F 545467 FH Nordin 1061 NY Nordin 1296a NY Nordin 1288 NBV 3925-55 PC Nordin 1333 ^-"-Collins West Brooksister NY Nordin 1276 ^-Collins 5845 Casco Bay FH Nordin IO69 NY Nordin 1272 ( C o l l i n s 1908) -x-x-x-Collins (1908) Casco Bay *Muxter? K i t t e r y Pt. FH Nordin 1071 Maryland *Wolle & Drouet 2302 Somerset Co. FH Nordin 1074 NY Nordin 1271 F 939834 UBC 49983 (Drouet 1939) 1 apparently when C o l l i n s made a c o l l e c t i o n with the two species together he made two sheets of the material, c a l l i n g one f o r instance Nodularia harveyana and the other Nodularia  spumigena. Both sheets show both species contained i n them. He usually gave both sheets the same c o l l e c t i o n number. 147 Massachusetts ^Chapman (1940) Essex Co. * C o l l i n s Woods Hole UC 752511 •KFarlow (1881) Cambridge **Farlow Woods Hole UC 100562 FH 4555 * S e t c h e l l Cambridge UC 100565 **T'aylor 3084 Nantucket Island UBC 49982 (Drouet 1935) ^Webber (1967) Essex Co. Michigan #Ackley (1931) Livingstone Co., Cheboygan C i t y & Co., Montcalm Co., Washtenaw Co. & Shiawassee Co. **Ackley (1931) Branch Co. ^Johnson Ann Arbor UC 752512 PC Nordin 1300 F 1157488 **Phinney 23M40 Emmet Co. F 1138192 M i s s i s s i p p i **Drouet 9844 Hancock Co. F 1310578 *Scott Bay St. Louis F I O 8 5 8 7 O Missouri *Casebolt 1954) 167 Clay Co. F Nordin II98 (Gier & Johnson Nebraska -"Anderson -/-Clements •$:-*Kiener x-Kiener ^Kiener •K--«-Kiener -SB'-Kiener -"--"Kiener -"r-"-Kiener XKiener -"-Kiener *Kiener ^ K i e n e r -"-Kiener -"-Kiener •5'-"Kiener *~"-Kiener & Walker ( 1 9 2 0 ) Cherry Co. ( I 8 9 6 & 1901) Lancaster Co. & South Bend 1 0 4 0 1 a Filmore Co. F 1099262 10466 Red Willow Co. F 1 0 9 9 4 4 0 13617 Lancaster Co. F 1127250 13777 Cherry Co. UC 679691 F 1127792 13885 Dodge Co. F 1131624 14139 Lancaster Co. F 1 1 3 2 1 8 1 14140 Lancaster Co. F 1 1 3 2 1 8 5 15687 Keith Co. F 1 1 3 9 9 1 1 16499 Kearney Co. UC 689174 F 1 1 4 4 6 5 5 20613a Sheridan Co. F 1219490 2 1 8 3 4 - 2 1 8 3 5 Dundy Co. F 1220286 F 1 2 2 0 3 4 5 22773a Scott's B l u f f Co. F 1249366 22778 Scott's B l u f f Co. F 1249569 23130 Garden Co. F Nordin I I 8 4 2 3602-23604 Lincoln Co. F Nordin 1 1 8 5 - 1 1 8 7 Nevada -"--"Christiansen 1731 Washoe Co. F Nordin 1169 *-"-LaRivers 616 Washoe Co. F Nordin 1168 1 4 8 New Hampshire XCo l l i n s Hampton NY Nordin 1 2 7 7 * * C o l l i n s Hampton NY Nordin 1 2 7 9 ( C o l l i n s 1 8 8 4 ) * * C o l l i n s Hampton NY Nordin 1 2 8 6 NY Nordin 1 2 9 2 NY Nordin 1 2 9 6 F Nordin 1 1 7 8 FH Nordin 1 0 5 9 ( C o l l i n s 1 8 8 4 ) New Jersey x-X-peters A t l a n t i c C i t y F 1 0 8 1 2 0 8 New York X-Martindale ( 1 8 8 9 ) Staten Island *Pike ( 1 8 8 6 ) Long Island x-X-Smith ( 1 9 2 4 ) Bergen North Carolina *Cocke ( 1 9 4 9 ) Duplin Co. **Cocke ( 1 9 4 9 ) Macon Co. x-Cocke ( 1 9 6 7 ) Piedmont & Coastal P l a i n **Cocke ( 1 9 6 7 ) Coastal P l a i n x-Whitford ( 1 9 4 3 ) New Hanover Co. x-x-whitford ( 1 9 4 3 ) Craven Co. x-Whitford and Schumacher (I969) Piedmont and Coastal P l a i n x-Whitford and Schumacher (I969) Coastal P l a i n North Dakota x-x-Brannon ( 1 9 1 1 ) Devils Lake K-Moore ( 1 9 1 7 ) Devils Lake -"-"-Moore & Carter ( 1 9 2 3 ) Ramsay, Nelson, Eddy & McLean Cos. x-x-yerch & B l i n n ( 1 9 7 2 ) Devils Lake x-x-Young ( 1 9 2 4 ) Devils Lake Ohio x-Lillick & Lee ( 1 9 3 4 ) Columbus -oi;-Lillick & Lee ( 1 9 3 4 ) C i n c i n n a t i """"Riddle ( 1 9 0 5 ) Champaigne Co. Oklahoma x-x-Toetz ( 1 9 7 3 ) Payne Co. Pennsylvania x-x-Habeeb 3 7 3 8 Pike Co. F Nordin 1 1 7 6 Tennessee x-x-Bold Goodeletsville F Nordin 1 1 7 4 x-Bold B 1 4 3 Nashville F 1 2 1 1 6 3 4 Utah x-Harding ( 1 9 7 1 ) Utah Co. 1 4 9 V i r g i n i a *Louver & S t r i c k l a n d 1 1 3 0 York Co. UC 6 8 0 4 1 1 F 1 1 0 9 7 9 2 *0tt ( 1 9 7 3 ) Yorktown **-Woodson et a l . (I966) Dinwiddle Co. Washington •x-x-x-Fairchild & Wilson (I967) Grant Co. ^-Gardner 3 3 5 (PBA 1 0 1 3 ) LaConner Skagit Co. UC I O O 5 6 9 NBV 3 9 2 5 - 3 8 F 5 4 6 2 7 9 NY Nordin 1 2 6 9 PC Nordin 1 3 0 6 ( S e t c h e l l & Gardner 1 9 0 3 ) -*-::-Gardner 4 1 1 (PBA 1 0 1 2 ) Whidby Island UC I O O 5 6 6 NBV 3 9 2 5 - 7 2 NY Nordin 1 2 9 7 F 9 8 0 8 1 6 F 1 0 4 7 7 8 1 PC Nordin 1 3 3 4 ( S e t c h e l l & Gardner 1 9 0 3 ) ^Gardner 4 3 6 Port Townsend UC IOO567 (Setc h e l l & Gardner 1 9 0 3 ) ^Schumacher & Muenscher ( 1 9 5 2 ) Whatcom Co. Wisconsin -x-Prescott ( 1 9 6 2 ) no lo c a t i o n x*-Prescott (I962) no lo c a t i o n EUROPE Austria *x-Brabez ( 1 9 4 1 ) Franzenbader x-Brabez ( 1 9 4 1 ) Franzenbader •5'Forti ( 1 9 0 7 ) c i t e s c o l l e c t i o n by Beck -x--x-Forti ( 1 9 0 7 ) c i t e s c o l l e c t i o n s made by Loitlesberger, Hansgirg Belgium -x-Beeftink 4 5 0 & 451 Fort St. P h i l l i p e NBV 3 9 2 5 - 3 2 NBV 3 9 2 5 - 3 3 -x-Forti ( 1 9 0 7 ) c i t e s two c o l l e c t i o n s by De Wildemann one by Bory Czechoslovakia •x-Forti ( 1 9 0 7 ) c i t e s a c o l l e c t i o n by Hansgirg at Carinthiae -x-Hansgirg Hermanmestec F Nordin 1 2 3 3 -x-Hansgirg Libochovice F Nordin 1 2 34 -x--x-Kol ( 1 9 2 6 ) Lomniczi -x-x-Rosa ( 1 9 5 1 ) Bohmen •*Tarnavschi ( 1 9 3 1 ) Bucovina 150 ' Denmark **Cleve (1897) Helleback *"-*Fjerlingstad (I969) Knudshoved, Sjaelland Island *-*0stenfeld F a l s t e r Island NY Nordin 1285 * * 0 s t e n f e l d Store Beelt NY Nordin 1284 •^Schmidt (1899) c i t e s c o l l e c t i o n s by Warming at S k a l l -ingen, Th. Mortensen at Nymindegab -x-"-Schmidt (1899) c i t e s c o l l e c t i o n s by Lyngbye & Hofman-Bang at Hofmansgave; Ostenfeld at Taarback, Anhatt, Fyr, Storebaeltj Rosenvinge at Svendborg, V e j l e f j o r d , Faeno-Sund, Limfjorden; Rosenvinge, Iffilge & Osten-f e l d at Oresund, Kjerteminde, Fredrideshaven, Groves Flak, Aalborg Bugt, Nordre Runner, Hirsholmene; Rosen-vinge & Th. Mortensen at Nykohing; Schmidt at Born-holm Island *-Sporring (1942) Skallingens *Wille (1897) Kirkebo, Faeroe Islands (Borgesen 1901) England -"-^Berkeley no date B r i s t o l NBV 3925-27 ---Bristol (1919 & 1920) Broadbalk -"-Carter (1933) Essex -«-Grove et a_l. (1920) Birmingham -"-*-Grove et a l . (1920) Birmingham -"-Harvey (I848) c i t e s c o l l e c t i o n by Thwaites at Shire-hampton near B r i s t o l -"--"-Harvey (I848) c i t e s c o l l e c t i o n s by Salway at Barmouth, Ralfs at Dolgelley, Thwaites at Shirehampton "--x-Joshua Cirencester F Nordin 1249 -"-Petersen (1935) no l o c a t i o n #*--"-Stewart & Pugh (1963) Lincolnshire *-*Thwaites? Shirehampton PC Nordin 1 3 3 0 *West (1899) Cambridgeshire Finland -"-"-Cedercreutz (1934) Alands-Sea -x-5:-Forti (1907) c i t e s 2 c o l l e c t i o n s by E l f v i n g *-"-Levander (1900) Alands-Sea ---x-Nylander Hel s i n k i PC Nordin 1317-1319 (Nylander 1861) France ---Bornet & Flahault (W&N 895) Cosne UC 759399 PC Nordin 1309 FH Nordin 1082 NBV 3925-34 NBV 3925-35 F Nordin 1235 NY Nordin 1281 -x-x-Fremy (1934) B e l l e - I l e & Brest -"-"-Fremy & Meslin (1924) La Meaffe 151 -IH'-Gomont Tables de l a Loire PC Nordin 1323 x-Gomont? Paris PC Nordin 1303 x-Gomont Cosne PC Nordin 1304 x-x-Gomont Auvergne PC Nordin 1324 *-*-Gomont (W.N.&L. 1343) Auvergne UC 761550 NBV 3925-49 F 975767 NY Nordin 1294 *Koster 6121 F i n i s t e r e NBV 3925-31 :-Lebel 415 Mont d'Huberville NBV 3925-16 PC Nordin 1337a -x-Lebel 580 Negreville PC Nordin 1337b x-Lebel 998 LeHavre PC Nordin 1316 x-Mabille Berthenicourt F Nordin 1232 PC Nordin 1299 x-Mabille (1954) Berthenicourt x-Roussel Talatus PC Nordin 1315 x-Thuret Cherbourg FH Nordin 1083 NBV 3925-36 PC Nordin 1308 x-Thuret Cherbourg PC Nordin 1298 PC Nordin 1305 Thuret Cherbourg FH Nordin 1063 PC Nordin 1327b (Bornet and Thuret 1880) Thuret ( i n Bornet & Thuret 1880) C r o i s i c •Thuret? Cosne PC Nordin 1313 Thuret Deauville FH Nordin 1065 NBV 3925-22 NBV 3925-24 PC Nordin 1327 PC Nordin 1331 x-x-villeret (1953) Bretagne Germany x-Anagnostidis & Schwabe (I966) Fehmarn Island x-x-Arndt et a l . (1966) Wismar-Bucht x-x-Bandel (1940) Rostoc x-X-Bornet & Flahault (1888) c i t e c o l l e c t i o n s by Braun (var. gen. " i n stagnis aquae d u l i s " ) ( v a r . l i t . " i n herb. Thuret") x-Bornet & Flahault (1888) c i t e a c o l l e c t i o n by "Hantzsch i n herb. Grunow" x-x-Braun Freiburg NBV 3925-15 PC Nordin 1338 x-x-Braun 1847 Freiburg NBV 3925-25 PC Nordin 1336 x-x-Bursa (1968) Gulf of Gdansk x-x-Forti (1907) c i t e s c o l l e c t i o n s by Lemmerman, Reinbold, Richter not seen x-Forti (1907) c i t e s c o l l e c t i o n s by Brand, Volk, M. Schmidt not seen x-x-Frohlich? Schleswig NBV 3925-57 NBV 3925-63 NBV 3925-70 PC Nordin 1326 PC Nordin 1335 x-x-Kalbe & Tiess (I964) Rostoc xx-Kleiboden? Wangerooge NBV 3925-13 x-x-Klock (1930) Unterwarnow x-x-Koch Borkum NBV 3925-17 NBV 3925-26a xx-Koch no l o c a t i o n NBV 3925-29b 152 * * K o l w i t z ( 1 9 1 0 ) Ostsee *Lindstedt ( 1943 ) & F o r t i ( 1907 ) c i t e c o l l e c t i o n s by Reinke, Reinbold **Mertens Norderney UC 436361 NBV 3 9 2 5 - 1 5 NBV 3 9 2 5 - 5 9 NBV 3 9 2 5 - 6 4 NBV 3 9 2 5 - 6 6 NBV 3 9 2 5 - 6 7 NBV 3 9 2 5 - 6 8 F 951302 *-*Pankow (19 64) Rostoc -"-Pankow ( 1 9 7 1 ) Ostsee, K i e l e r Forde -"-Petersen ( 1935 ) no l o c a t i o n -*-"-Rathsack-Kutzenbach (I96I) Rugen Island -"-^•Rathsack-Kutzenbach ( 1 9 6 1 a ) M i t t l e r e n Ostsee **Richter ( 1894 ) Ploner Sea •H-Rose Berra (Rabh. Algen 237) NBV 3 9 2 5 - 1 4 F 999057 NY Nordin 12 59 ^-"-Schmitz (H&R 142 ) Greifswalder Bodden UC 760936 UC 953542 NBV 3 9 2 5 - 5 8 NBV 3 9 2 5 - 6 5 PC Nordin 1325 PC Nordin 1332 *-*-Trahms ( 1937 ) Rugen #-"Waldemann ( 1959 ) middle Ostsee also c i t e s c o l l e c t i o n s by Merkle, Driver, Hessle and V a l l i n , Ostenfeld, Brandes, Rothe, Bandel Hungary -::->"-Fritsch (I964) c i t e s c o l l e c t i o n by Palik -"Hortobagyi ( 1959 ) Lake S z e l i d *-"-Palik ( 1961 ) *Tamas ( 1 9 5 8 ) Balaton Sea Iceland ^--"Petersen ( 1932 ) Einarsnes -"-Petersen ( 1 9 3 2 ) Hvalnes & Knararnes & Einarsnes Ireland -"-Bornet & Flahault ( 1888 ) **Rees ( 1935 ) Cork Co. c i t e s c o l l e c t i o n by Harvey I t a l y -"Bornet & Flahault ( 1888 ) c i t e c o l l e c t i o n by Menghini -"-"Forti ( 1 9 0 7 ) c i t e s c o l l e c t i o n by F o r t i Netherlands -"-Beeftink 0 - 1 9 9 & 0 - 2 0 1 Zeeuwsch-Vlaanderen NBV 3 9 2 5 - 1 0 NBV 3 9 2 5 - 1 1 -"-Bierbr'auer Ostvoorne NBV 3 9 2 5 - 1 2 -"-Bilio 2 3A Goeree NBV 392 5 - 7 ---Bilio 6 1 0 2 5 - 2 Goeree NBV 3 9 2 5 - 9 -"-Bilio 22B Goeree NBV 3 9 2 5 - 2 153 * B i l i o 6 Noord-Beveland NBV 3925-4 x-Bilio 9 Noord-Beveland NBV 392 5-3 * B i l i o W Noord-Beveland NBV 3925-5 x-Bilio M Noord-Beveland NBV 3925-6 x-Bilio M107 Ostvoorne NBV 3925-8 * F o r t i (1907) c i t e s l i s t s of c o l l e c t i o n s •x-x-Forti (1907) c i t e s c o l l e c t i o n s by Van den Bosch, Surinigar at Leeuwarden *Simons & Vroman (1973) De Putten xx-Van den Bosch Goes PC Nordin 1320 Norway * * F o s l i e (WN&LI1344) Bugonaes UC 761551 UC 100562 NBV 3925-21 F 975766 NY Nordin 1295 Poland -**Collector unknown Danzig Bay NBV 3925-23 -"-Plinski (1973) Leczyca •x-Starmach (1966) no l o c a t i o n -x-x-Starmach (1966) no l o c a t i o n Rumania •x-Hof & Fremy (1933) Szovata Siebenburgen Scotland «Bornet & Flahault (1888) c i t e c o l l e c t i o n by Batters •x-x-Bornet & Flahault (1888) c i t e c o l l e c t i o n by Batters at the mouth of the Clyde -x-Stewart (i960) Hebrides Spain •x~x-Gonzalez-Guerro (1928) Madrid Sweden -:i--"-Borge (1907) Vaddo, Edeby •x-Cedergren (1926) Kolviken •x-x-Cleve (1897) Moseskar Vaderdarna •x-Granhall & Henriksson (I969) x-x-Lindstedt (1943) Torekovj Bohuslan: Fiskebiickskil Stockevik -x-Lindstedt (1943) several locations -x~x-Nordstedt (W&N I98) Lomma FH Nordin 1090 NBV 3925-50 NBV 3925-61 NY Nordin 1298b F Nordin 1250 x-Nordstedt (1897) c i t e s c o l l e c t i o n by C. Ag. i n herb Thuret -x-x-Nordstedt (1897) c i t e s c o l l e c t i o n by C. Ag. at Bastad and Areschoug exsice II #193 from same area *-x-Sjostedt (1922) Oresund 154 *-*-Skuja (1924) from Lappland Coast *Skuja (1926) Staburags **-Skuja (1926) c i t e s c o l l e c t i o n by Winkler from west B a l t i c Coast Switzerland ---Cramer (Rabh. Alg.Sach. 994) Zurich FH Nordin IO84 NBV 3925-1 F 1002450 NY Nordin 1262 EXCLUSIVE OF NORTH AMERICA AND EUROPE Alg e r i a *Debray (1893) Antarctica *-Apfel 51 & 52 Burger Lakes F 1299674 F 1299701 ^ F r i t s c h Winter Harbour BM 1815 ( F r i t s c h 1912) •x-Fukushima (1959) Ongul Island A n t i l l e s (Dutch) Wan den Hoek et a l . (1972) Curacao -"-*Wagenear-Humelinck 641 Aruba NBV 3925-52 (Koster i960) Argentina •5--"Borge (1901) Patagonia •"-"-Borge (1907a) Puna de Atacama -"-DeHalperin (1970) Gulfo Nuevo Au s t r a l i a *~"Bayly & Williams (I966) Lake Corangamite -"-"-Francis? Lake Alexandrina FH Nordin 1064 (Francis I878) *~"-Playf a i r (1914) Lismore **Schmidle (I896) -x-ftWilliams (1970) Barbados -"-West (1904) Chancery Lane Estate China -"Skvartzow (1927) North Manchuria *~"-Skvartzow (192 7) Harbin, North Manchuria Columbia -"-West (1914) Cundinamarca 155 Egypt x~x-Fritsch (I964) c i t e s c o l l e c t i o n by Nayal **West 393 Birket Quarun BM 302 (West 1909) Falkland Islands x-x-Guarrera & Kuhnemann (1949) c i t e c o l l e c t i o n by V a l l e n t i n at Malvinas Formosa **0kada (1932) Kotosho Guatemala **Standey 65787 Rio Pucal Dept. F 1023960 x-Tilden (1908) Lake Amatitlan India -x-x-Bharadwaja (1935) Benares -Chacko (1972) Madras -Franklin (1972) Madras *--x-Iyengar & Desikachery (1944) Hare Island, South India •x-x-Kamat (1968) Alibag, Maharashtra x-x-Kumar (1970) Sardhana x-Raju (1972) Kamat **Raju (1972) Kanpur -x-x-Randawa (1936) N. India x--x-Subrahmanyam (1972) Bastar, Jagdalpur x--x-Vasishta (1961 & i960) Hoshiarpur I s r a e l -Jabotinsky (I96I) Lake Zohar Japan -x-Umezaki (I96I) several locations **Yoneda (1937) Shinano Java -x-Mobius (1893) Solo **Van Oye (1922) Batavia Wan Oye (1923) Tasikmalaja -x-Jutono (1973) Jogjakarta Mexico -x-Drouet & Richards 3416 Empalme, Sonora F 1031425 -Ortega (1972) Lake Texcoco -x-Patrick Lake Texcoco F Nordin 1241 1 5 6 New Zealand -"-Chapman (1955) Stanmore Bay ^-Chapman l b Stanley Bay F 1246844 *Dellow (1955) Hauraki Gulf *Dellow & Cassie (1955) Whangaparaoa Bay Peru **Ibanez T r u j i l l o F Nordin 1238 **---Maldonaldo Lago V i l l a F 1102709 F 1102707 Puerto Rico **Wille 1817a Laguna Guanica UC 463735 NY Nordin 1256 (Gardner 1927) S i e r r a Leone ***Woodhead & Tweed ( 1957) no l o c a t i o n South A f r i c a * * F r i t s c h & Rich (1929) Kimberley -"-^•Hutchinson et a l . ( 1932) Transvaal (Rich 1930) (Rich 1934) -^-Pearson 26 Orange River BM Nordin 1348 ( F r i t s c h 1918) South West A f r i c a "Welch (I964) Swakop River, Rietfontien Spring -xx-Welch (I964) Gross Barmen, Etosha Reserve *~--Welch ( 1965) Okandu *West ( 1912) L i t t l e Namaqualand Sudan **Karim ( I 9 6 8 ) Jebel Marra Tunisia -"-Serpette & Labbe ( I 9 6 6 ) between Rades and Miliane **Serpette & Labbe ( 1966) Oasis de Tozeur U.S.S.R. *Elenkin (1916) Kislovodsk -"Kosinskaja Tartar Autononous SSR ( c i t e d i n F r i t s c h I964) -"-Proshkina-Lavrenko (I968) Caspian Sea -"-"Proshkina-Lavrenko (I968) Caspian Sea -"-Shtina (1972) Khirov S.S.S.R -"--"-Starmach ( 1 9 6 6 ) c i t e s c o l l e c t i o n by Woronichin i n S i b e r i a *--"-Zhadin & Gerd (1961) Lake Balkash 157 Table VIII. Medium BG-11 (Stanier et a l . 1971) Compound Amount (g/1) NaN03 1.5 K 2HP0 4 0.04 MgS04-7H20 0.075 CaCl 2.2H 20 0.036 Na CO 0.02 F e r r i c ammonium c i t r a t e 0.006 EDTA (disodium s a l t ) 0.001 Trace metal mix A5 1 ml/1 A5 H 3B0 3 2.86 MnCl 2.4H 20 1.81 ZnS0 4.7H 20 0.222 Na 2Mo0 4.2H 20 0.39 CuS0 4.5H 20 0.079 Co(N0 3) 2.6H 20 0.0494 notes: The f e r r i c ammonium c i t r a t e should be autoclaved separately to eliminate p r e c i p i t a t i o n and added a s e p t i c a l l y a f t e r cooling. : the Na 2C0 should be autoclaved sep-arately i f the desired f i n a l pH i s above 8, and added a s e p t i c a l l y a f t e r cooling. 158 Table IX. Herbarium material of Nodularia examined. Nodularia spumigena Berkeley no date B r i s t o l , England NBV 3925-27 Bold 28 i i i 1953 Goodeletsville, Tenn. F Nordin 1174 Braun 1847 Freiburg, Germany NBV 3925-15 PC Nordin 1338 (type of Kiitz. S. Vriesiana) Braun 1847 Freiburg, Germany NBV 3925-25 PC Nordin 1336 Brebisson? no date no lo c a t i o n NBV 3925-71 Christensen 1731 14 v 1953 Washoe Co., Nev. F Nordin 1169 c o l l e c t o r unknown (Kleibaden ? ) v i i 1839 Wangerooge, Germany NBV 3925-13 c o l l e c t o r unknown 5 v i i i 1887 Danzig Bay, Poland NBV 3 925 -23 C o l l i n s no date West Brooksister, Maine NY Nordin 1276 C o l l i n s v i i i I884 Rockingham Co., N.H. FH Nordin 1059 F Nordin 1178 NY Nordin 1286 NY Nordin 1292 NY Nordin 1296 C o l l i n s v i i i I884 Hampton, N.H. NY Nordin 1279 C o l l i n s 2439 v i 1892 Cape Rosier, Maine FH Nordin 1060 NY Nordin 1287 C o l l i n s 3459 14 v i i 1897 Cape Rosier, Maine NY Nordin 1289 C o l l i n s 5094 v i i 1904 Casco Bay, Maine UC 752514 F 1219336 C o l l i n s 5526 (Phycotheca Borealis Americana 1307) 14 v i i 1906 Harpswell, Maine UC 693286 FH Nordin 1061 NBV 3925-55 F 545467 NY Nordin 1288 NY Nordin 1296a PC Nordin 1333 Drouet 9844 9 x i i 1948 Hancock Co., Miss. F 1310578 Farlow v i i I889 Woods Hole, Mass. UC 100562 FH 4555 F o s l i e (Wittrock, Nordstedt & Lagerheim, Algae Exsiccatae 1344) 10 v i i i 1889 Bugonaes, Norway UC 761551 UC 100562 NBV 3925-21 F 975766 NY Nordin 1295 Francis?* I878? Lake Alexandrine near Adelaide, A u s t r a l i a FH Nordin IO64 Gardner 411 (PBA 1012) v i 1901 Whidbey Island, Wash. UC 100566 NBV 3925-72 F 980816 F 1047781 NY Nordin 1297 PC Nordin 1334 Gardner 436 15 v i 1901 Port Townsend, Wash. UC IOO567 Gardner 3295 29 i v 1916 Oakland, C a l i f . UC 114844 Gardner 7231 14 i v 1933 San Mateo Co., C a l i f . FH Nordin 1081 Gardner 7946 11 v I936 Contra Costa Co., C a l i f . UC 661724 FH Nordin 1056 NBV 3925-56 F 1043544 NY Nordin 1293 ^Herbarium sheet FH Nordin IO64 has no c o l l e c t o r ' s name on i t but Francis apparently d i s t r i b u t e d 1 h i s samples of Nodularia from t h i s s i t e . Bornet & Flahault (1888) report examining material from him. 159 Gomont? 30 v i i i 1885 Tables de l a Loire, France PC Nordin 1323 Gomont? 15 v i i i 1894 Auvergne, France PC Nordin 1324 Gomont (Wittrock, Nordstedt & Lagerheim, Algae Exsiccatae 1343) v i i i 1894 Auvergne, France UC 761550 NBV 3925-49 F 975767 NY Nordin 1294 Habeeb 3738 15 v 1951 Pike Co., Pa. F Nordin 1176 Holden 1458-1461 28 v 1899 Bridgeport, Conn. FH Nordin 1062 NY Nordin 1290 Holden 30 v i 1899 no l o c a t i o n NY Nordin 1291 27 x 1952 T r u j i l l o , Peru F Nordin 1238 no date Cirencester, England F Nordin 1249 13777 22 v i i 1936 Cherry Co., Nebr. UC 67969I Ibanez Joshua Kiener F Kiener Kiener Kiener F Kiener F Kiener Kiener Kiener Kiener 1127792 10401a 13885 21 v i i 1941 Fillmore Co., Nebr. F 1099262 23 i v 1943 14139 & 14140 6 1132185 F 1132181 16499 6 i v 1944 1144655 21834 21835 23130 Dodge Co., Nebr. F 1131624 v i 1943 Lancaster Co. Nebr. Kearney Co., Nebr. UC 689174 F 1220345 F 1220286 F Nordin II84 F Nordin 27 i i i 1947 Dundy Co., Nebr, 27 i i i 1947 Dundy Co., Nebr, 2 i v 1948 Garden Co., Nebr. 23602-23604 17 v 1948 Lincoln Co., Nebr. 1185 F Nordin 1186 F Nordin 1187 Koch 3 v i i 1845 Borkum, Germany NBV 3925-17 NBV 3925-29a Koch 3 v i i I846 no location NBV 3925-29b LaRivers 6 l b 22 i x 1951 Washoe Co., Nev. F Nordin 1168 Lebel 415 14 v i 8 6 0 Mont d'Huberville, France NBV 3925-16 PC Nordin 1337a Lebel 58O 12 i v 1862 Negreville, France PC Nordin 1337b 1942 Lago V i l l a , Peru F 1102707 F 1102709 Norderney, Germany UC 436361 NBV 3925-15 NBV 3925-64 NBV 3925-66 NBV 3925-67 F 951302 Nordstedt (Wittrock & Nordstedt, Algae Exsiccatae I98) 11 v i 1877 -Lomma, Sweden FH Nordin 1090 NBV 3925-50 NBV 3925-61 F Nordin 1250 NY Nordin 1298b Nylander i 8 6 0 H e l s i n k i , Finland PC Nordin 1317 PC Nordin PC Nordin 1319 5 v i i i 1901 F a l s t e r , Denmark NY Nordin 1285 1 v i i i 1904 Store Beelt, Denmark NY Nordin 1284 559 (not PBA 1061 as marked on UC specimens) 28 v i 1902 Alameda Co., C a l i f . UC 393931 UC 202712 NBV 3925-54 FH Nordin 1066 PC Nordin 1328 Osterhout & Gardner (PBA 1061) 30 v 1902 Oakland, C a l i f . FH Nordin IO87 NBV 3925-30 F 9808I6 NY Nordin 1263 PC Nordin 1302 Maldonado 41 i Mertens v i 1821 NBV 3925-59 NBV 3925-68 1318 Ostenf eld Ostenf eld Osterhout 160 Peters 14 v 1891 A t l a n t i c City, N.J. F 1081208 Phinney 23M40 8 i i 1940 Emmet Co., Mich. F 1138192 Rose v i i 1852 Berra, Germany NBV 3925-14 F 999057 NY Nordin 1259 Schmidt 1899-1900 Siam? PC Nordin 1322 Schmitz, Hauck & Richter (Phykotheca U n i v e r s a l i s 142) v i i i 1886 Greifswalder Bodden, Germany UC 760936 UC 953542 NBV 3925-58 NBV 3925-65 PC Nordin 1325 PC Nordin 1332 S e t c h e l l 1628 14 i v 1897 San Benito Co., C a l i f . UC 752513 F 1221362 S e t c h e l l & Jepson v i i I896 between La Grange, Stanislaus Co. & C o u l t e r v i l l e , Mariposa Co., C a l i f . UC 202713 NBV 3925-53 Standey 65787 20 i i 1939 Rio Pucal Dept., Guatemala F 1023960 Suhr (Frolich?) v i i 1834 Schleswig NBV 3925-57 NBV 3925-63 NBV 3925-70 PC Nordin 1326 PC Nordin 1335 (type of Kutzings N. suhriana = N. spumigena Mertens & Suhr's Lyngbya annulata) Taylor 3084 29 v i i i 1920 Nantucket Island, Mass. UBC 49982 Transeau 63 14 v 1911 Charleston, 1 11 . NY Nordin 1283 Thuret 30 v i i i 1874 Cherbourg, France FH Nordin IO63 PC Nordin 1327b Thuret? v i i i 1874 Deauville, France FH Nordin 1065 NBV 3925-22 NBV 3925-24 PC Nordin 1327 PC Nordin 1331 Thwaites? v 1867 Shirehampton, England PC Nordin 1330 Van den Bosch no date Goes, Netherlands PC Nordin 1320 Wagenear-Hummelinck 641 11 v 1955 Aruba, Dutch A n t i l l e s NBV 3925-52 West 393 8 x i i 1911 Birket Quarun, Egypt BM 302 Wi l l e 1817a Laguna Guanica, Puerto Rico UC 463735 NY Nordin 1256 Nodularia harveyana Apfel 51 & 52 19 i 1948 Burger Lakes, Antarctica F 1299674 F 1299701 Atkinson 1895 no location PC Nordin 1301 Atkinson 1895 no location PC Nordin 1321 Beeftink 0-199 29 v i i i 1951 Zeeuwsch-Vlaanderen, Nether-lands NBV 392 5-10 Beeftink 0-201 29 v i i i 1951 Zeeuwsch-Vlaanderen, Nether-lands NBV 3925-11 Beeftink 450 28 v i i 1955 Fort St. P h i l l i p e , Belgium NBV 3925-33 161 Beeftink 451 28 v i i 1955 Fort St. P h i l l i p e , Belgium NBV 3925-32 Bierbrauer 6 v i 1953 Ostvoorne, Netherlands NBV 3925-12 B i l i o 61025-2 29 v i 1961 Goeree, Netherlands NBV 3925-9 B i l i o M107 24 i x 1965 Ostvoorne, Netherlands NBV 3925-8 B i l i o 6 6 i x I966 Noord-Beveland, Netherlands NBV 3925-4 B i l i o 9 6 ix 1966 Noord-Beveland, Netherlands NBV 3925-3 B i l i o W 6 ix 1966 Noord-Beveland, Netherlands NBV 3925-5 B i l i o M 6 ix I966 Noord-Beveland, Netherlands NBV 3925-6 B i l i o 22B 17 x I966 Goeree, Netherlands NBV 3925-2 B i l i o 23A 17 x I966 Goeree, Netherlands NBV 3925-7 Bold B143 12 i 1947 Nashville, Tenn. F 1211634 Bornet & Flahault (Wittrock & Nordstedt, Algae Exsiccatae 895) 25 ix 1886 Cosne, France UC 759399 FH Nordin 1082 NBV 3925-34 NBV 3925-35 F Nordin 1235 NY Nordin 1281 PC Nordin 1309 Brannon 141A 5 i i 1943 G a i n s v i l l e , F l a . F 1124079 PC Nordin 1329 Casebolt 167 5 v i 1950 Clay Co., Mo. F Nordin 1198 Chapman l b i x 1947 Stanley Bay, New Zealand F 1246844 C o l l i n s 4844 no date no l o c a t i o n NY Nordin 1273 NY Nordin 1274 C o l l i n s v i i I884 Hampton, N.H. NY Nordin 1277 C o l l i n s 2387 14 v i i 1892 Cape Rosier, Maine FH Nordin 1070 FH Nordin 1073 NY Nordin 1275 C o l l i n s 22 v i i 1894 Cape Rosier, Maine UC 687705 FH Nordin 1072 NBV 3925-39 F 1209180 NY Nordin 1278 PC Nordin 1310 C o l l i n s (PBA 1062) 14 v i i 1903 Casco Bay, Maine F 545439 NY Nordin 1280 PC Nordin 1307 C o l l i n s 5094 v i i 1904 Casco Bay, Maine UC 752510 NBV 3925-37 F 1219372 C o l l i n s 16 v i i i 1904 Woods Hole, Mass. UC 7525H C o l l i n s 5845 10 v i i 1908 Casco Bay, Maine FH Nordin 1069 NY Nordin 12 72 Cramer (Rabenhorst Algen Sachsens 994 ) v i & v i i i860 Zurich, Switzerland FH Nordin IO84 NBV 3925-1 NY Nordin 1262 F 1002450 Demaree & Thomason 24582 1 v i i i 1943 C o l l i n s , Drew Co., Ark. F 1133627 Drouet 8766 28 x 1948 Calcasieu Parish, La. UC 910404 NBV 3925-45 F 1332708 PC Nordin 1311 Drouet, Glassman & Chapp 1 2 7 1 1 2 v i i 1957 Chicago, 111. F Nordin 1203 Drouet, Glassman & Chapp 12696 2 v i i 1957 Chicago, 111. F Nordin 1202 Drouet, Madsen & Crowson 11511 27 i 1949 Wakulla Co., F l a . F 1324385 162 Drouet, Madsen & Crowson 11513 27 i 1949 Wakulla Co., F l a . UC 912044 F 1324476 Drouet & McBride 4569 H x 1941 San Bernardino Co., C a l i f . UC 664554 FH Nordin 1076 NBV 3925-43 NY Nordin 1268 F 1101885 Drouet & Nielsen 11232 23 i 1949 Taylor Co., F l a . UC 912045 F 1308871 Drouet & Nielsen 11681 31 i 1949 Franklin Co., F l a . UC 912043 F 1317616 Drouet & Richards 3416 23 x i i 1939 Empalme, Mexico F 1031425 F r i t s c h no date Winter Harbour, Antarctica BM 1815 Gardner 335 (PBA 1013) 19 v 1901 La Conner, Wash. UC 100569 NBV 3925-38 F 546279 NY Nordin 1269 PC Nordin 1306 Gardner 989 (PBA 1063) 29 v 1903 Marin Co., C a l i f . UC 341420 FH Nordin 1088 NBV 3925-48 F 9808I6 NY Nordin 1282 PC Nordin 1314 Gardner I498 v i i 1905 Oakland, C a l i f . UC 202866 F Nordin 1192 Gardner 1604 x i 1905 Berkeley, C a l i f . UC 202865 uc 274104 Gardner 3963 v i i 1917 Sit k a , Alaska UC 661526 F 1047757 Gardner 4151 i 1918 Oakland, C a l i f . UC 66I636 FH Nordin 1078 NBV 3925-42 F 1032981 NY Nordin 1266 Gardner 6552 x i i 1930 Berkeley, C a l i f . UC 440248 Gardner 6568 12 i 1931 Berkeley, C a l i f . UC 661705 FH Nordin 1075 NBV 3925-46 F 1033552 NY Nordin 1264 Gardner 7670 4 i i 1934 Berkeley, C a l i f . UC 641592 FH Nordin 1080 NBV 3925-40 F 1036391 NY Nordin 1270 Gardner 7963 7 v i 1936 Lake Merced, C a l i f . UC 661654 FH Nordin 1077 NBV 3925-41 F 1034223 NY Nordin 1267 Gomont 8 v i 1887 Paris, France PC Nordin 1303 Gomont 11 v i i 1892 Cosne, France PC Nordin 1304 Groesbeck 90 12 v i 1940 Mono Co., C a l i f . F 1049304 Hansgirg v i i 1888 Libochovice, Czechoslovakia F Nordin 1234 (Hansgirg 1892) Hansgirg 1891 Hermanmestec, Czechoslovakia F Nordin 1233 Hollenberg 1553 23 i i i 1934 Santa Ana River, C a l i f . UC 634466 Johnson 6 v 1893 Ann Arbor, Mich. UC 752512 F 1157488 PC Nordin 1300 Kiener IO466 23 v i i 1941 Redwillow Co., Nebr. F 1099440 Kiener 13617 17 x i 1942 Lancaster Co., Nebr. F 1127250 Kiener I5687 24 i x 1943 Keystone, Nebr. F 1139911 Kiener 20613a 23 v 1946 Sheridan Co., Nebr. F 1219490 Kiener 22773a 23 v i i i 1947 Scott's B l u f f Co., Nebr. F 1249366 163 Kiener 22778 24 v i i i 1947 Scott's B l u f f Co., Nebr. F 1249569 Koster 6121 16 i v 1957 Penze, France NBV 3925-31 Lebel 791 & 792 no date France NBV 3925-16 NBV 3925-62 Lebel 998 24 v i i 1869 LeHavre, France PC Nordin 1316 Louderback 17 22 v i i i 1947 Denver, Colo. F 1235673 Louver & St r i c k l a n d 1130 9 v 1942 York Co., Va. UC 680411 F 1109792 Mabille 8 1 x i 1952 France F Nordin 1232 PC Nordin 1299 Muxter? 10 v 1916 K i l l e r y Pt., Maine FH Nordin 1071 Palmer, Webster, Prettyman, Webster & Drouet 2518 17 v i i i 1939 Indianapolis, Ind. F 9808I6 Patrick 197 24 v i i 1947 Xochimilco, Mexico F Nordin 1241 Pearson 26 no date South A f r i c a BM Nordin 1348 Roussel 16 v i i i I869 Talatus?, France PC Nordin 1315 Scott 30 i i i 1941 Bay St. Louis, Miss. F IO8587O S e t c h e l l 1679 no date San Francisco, C a l i f . UC IOO563 S e t c h e l l 17 i 1889 Cambridge, Mass. UC IOO565 Standley 14 i i i 1940 Punta Rossa Lee Co., F l a . F 1030027 Thuret 31 v i i 1874 Cherbourg, France PC Nordin 1298 PC Nordin 1305 Thuret 19 v i i i 1874 Cherbourg, France FH Nordin 1083 NBV 3925-36 PC Nordin 1308 Thuret? v i i I889 Cosne, France PC Nordin 1313 Velasquez, Richards & Drouet 2507 4 v i i i 1939 Cook Co., 111. F 980549 Wolle & Drouet 2302 26 v i i i 1938 Somerset Co., Md. FH Nordin 1074 NY Nordin 1271 F 939834 UBC 49983 The following specimens were i n too poor a condition or i n i n s u f f i c i e n t numbers to make a judgement on t h e i r i d e n t i f -i c a t i o n . c o l l e c t o r unknown i x 1885 Pavia, Germany NBV 3925-26 Drouet & Louderback 5739 21 v i i i 1946 Salt Lake Co., Utah F 1198553 Drouet & Richards 2709 26 x 1939 Si e r r a Co., N. Mex. F 1038909 Fan 10130 19 v i 1954 Hubbard Co., Minn. UBC 49981 (Drouet 1956) Gardner 602 v i i 1899 Whidbey Island, Wash. UC 100568 Kiener 13940 23 i v 1943 Dodge Co., Nebr. F 1131688 Runyon & L i l l i c k 609 25 i x 1933 Columbus, Ohio FH Nordin 1058 Stockmayer (Vindobon Kryptogamas Exsiccatus 428) v i i i 1893 Frankenfels, Austria FH Nordin 1086 Taylor 1923 P u r c e l l Range, B r i t i s h Columbia (Taylor 1928a) uncatalogued material at UBC 164 West 44 1 v i i i 1904 Niamkolo, Tanganyika BM Nordin 1344 West 22 23 v i 1904 Nkata Bay, Tanganyika BM Nordin 1346 West 134 10 x i 1904 Komba Bay, Tanganyika BM Nordin 1347 West 208 10 i 1905 Toa, Tanganyika BM Nordin 1345 West 432 10 i 1905 Toa, Tanganyika BM 303 The following material i s more properly placed i n a genus other than Nodularia Aavd? 7 v i i 1853 Leipzig, Germany FH Nordin 1068 = Scytonema sp. Al l e n 5079a 2 i v 1953 from herbarium specimen F Nordin 1181 = Calothrix sp. Crossland 7292 23 x 1929 T a h i t i UC 696203 - Hormotham-nion (solutum?) Born. & Grun. F r i t s c h 363 no date Winter Harbour, Antarctica BM 1934 = Anabaena sp. Gardner 3305 9 v 1916 Berkeley, C a l i f . UC 661514 FH Nordin 1079 NBV 3925-44 F 1033572 NY Nordin 1265 = Anabaena v a r i a b i l i s Rabenhorst (Algen Sachsens 469) v i i 1855 Ultdobern, Ger-many FH Nordin 1067 NBV 3925-28 NY Nordin 1260 = Scytonema sp. Rabenhorst (Algen Sachsens 470) v i 1855 Zurich, Switzerland UC 432853 FH Nordin IO89 NBV 3925-60 NBV 3925-69 F 1015345 F 1015346 F 1015349 NY Nordin 1261 = Anabaena sp. Roussel 4 ix 1851 "Meloduuo", France PC Nordin 1312 = Anabaena sp. Rrealea x 1857 Attelobern, Germany NBV 3925-19 NBV 3925-20 = Scytonema sp. Se t c h e l l 23 v i i i 1889 Walite H i l l Pond, R.I. UC IOO564 = Nostoc (carneum?) Ag. Tild e n (American Algae 484) 22 v 1900 Oahu, Hawaii UC 740394 NY Nordin 1258 PC Nordin 1339 UBC 2971 - Hormotham-nion sp. Tild e n (South P a c i f i c Algae 11) x 1909 T a h i t i UC 233460 NBV 3925-47 NY Nordin 1257 = Hormothamnion sp. Wi l l e 282d 6 i 1915 Coamo Springs, Puerto Rico UC 401708 FH Nordin 1085 NY Nordin 1255 = Scytonema sp. Nostoc sp. Wi l l e 1415a no date Puerto Rico UC 401800 NY Nordin 1254 = Scytonema sp.Nostoc sp. 165 T a b l e X . Synonomy o f N o d u l a r i a / M e r t e n s i n Juergens7 B o r n e t e t F l a h a u l t 1888 N o d u l a r i a s p u m i g e n a / M e r t e n s i n J u e r g e n s . 7 B o r n e t e t F l a h a u l t 1888 IT. s p u m i g e n a v a r . g e n u i n a B o r n e t e t F l a h a u l t 19. s p u m i g e n a v a r . l i t o r e a B o r n e t e t F l a h a u l t 19. s p u m i g e n a v a r . ma j o r B o r n e t e t F l a h a u l t II. s p u m i g e n a v a r . v a c u o l a t a F r i t s c h 19. s p u m i g e n a v a r . a e r o p h i l a B r a b e z II. s p u m i g e n a f . e r a s s a ( W o r o n i c h i n ) E l e n k i n N . a r m o r i c a T h u r e t N. w i l l e i G a r d n e r N o d u l a r i a h a r v e y a n a ( T h w a i t e s ) T h u r e t 1875 19. h a r v e y a n a v a r . s p h a e r o c a r p a ( B o r n e t e t F l a h a u l t ) E l e n k i n II. a e r o p h i l a B r a b e z N_. s k u j ae G o n z a l e z - G u e r r o II. s p h a e r o c a r p a B o r n e t e t F l a h a u l t N . s p u m i g e n a v a r . m i n o r F r i t s c h N . t u r i c e n s i s ( C r a m e r ) H a n s g i r g S p e c i e s E x c l u d e n d a e N_. e p i p h y t i c a G a r d n e r = N o s t o c s p . ( j u v e n i l e ) II. f e r t i l i s s i m a Randawa nom. n u d . II. f u s c a T a y l o r = c o n g l o m e r a t e N . h a w a i i e n s i s T i l d e n = H o r m o t h a m n i o n ? s o l u t u m B o r n e t e t F l a h a u l t N . i m p l e x a ( B o r n e t e t F l a h a u l t ) B o u r r e l l y = A u l o s i r a i m p l e x a B o r n e t e t F l a h a u l t 19. q u a d r a t a F r i t s c h = A n a b a e n a s p . II. s p u m i g e n a v a r . zu j a r i s G o n z a l e z - G u e r r o = A n a b a e n a sp . N . t e n u i s G . S . W e s t . = A n a b a e n a s p . c o n t i n u e d . 166 Species Inguirendae N_. mainensis F.L. Harvey IJ. paludosa Wolle Spermosira a t l a n t i c a Dickie 

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