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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.Sc,  U n i v e r s i t y of N o r t h D a k o t a ,  1970  M o S c , U n i v e r s i t y of N o r t h D a k o t a ,  1971  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR O F PHILOSOPHY  i n the  Department of BOTANY  We a c c e p t t h i s t h e s i s as conforming t o required standard  THE  UNIVERSITY MAY,  OF  BRITISH  1974  COLUMBIA  the  In p r e s e n t i n g an  this  thesis  i n partial  advanced degree a t t h e U n i v e r s i t y  the  Library  s h a l l make i t f r e e l y  f u l f i l m e n t of the requirements f o r of B r i t i s h  that  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 that permission f o r extensive for  Columbia, I agree  copying of this  thesis  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 t h e Head o f my D e p a r t m e n t o r  by  h i s representatives.  of  this  written  thesis  forfinancial  gain  permission.  Department o f  firTTfiKil  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a  »ate  I t i s understood that  Kacj  3^J93!±  Columbia  shall  copying or publication  n o t be a l l o w e d w i t h o u t my  "The and  tapers  The  sun  The  shooting  stars  And  chaos  at  is  moon,  run  the  down end  gods,  like in  waxen  purple  globes; jellies,  hand."  Dryden's sudden after  all  of  Oedipus,  appearance a  rain  fall.  referring of  Nostoc  to  the  balls  iii Chairman:  Professor J . Stein ABSTRACT  The genus Nodularia /Mertens i n JuergensJ Bornet et Flahault  1888  i s considered with regard to i t s ecology, d i s t r i b u t i o n , physiology and taxonomy. Ecology was  studied by periodic sampling of organisms and moni-  toring environmental  parameters i n four saline ponds i n the B r i t i s h  Columbia i n t e r i o r , as well as by sampling a variety of brackish, marine and other inland saline water bodies. i n habitats of a l k a l i n e pH 8.2 s a l t s (4 - 60 °/  oo  - 10.0)  Nodularia was  collected  and medium to high dissolved  ) and i s associated with a s p e c i f i c group of eury-  haline bluegreen algae.  However, environments i n which i t grows are  usually dominated by species of green algae, with Nodularia only occasi o n a l l y becoming dominant. The d i s t r i b u t i o n of Nodularia was c o l l e c t i o n s and l i t e r a t u r e reports.  examined by reference to herbarium  Although many bluegreen algae are  considered cosmopolitan, Nodularia has a temperate and subtropical d i s t r i b u t i o n with few reports from polar or t r o p i c a l l a t i t u d e s . The growth and physiology was  investigated using 16 i s o l a t e s  from Canada, United States, Great B r i t a i n and A u s t r a l i a .  Isolates  were grown i n a wide range of conditions of l i g h t , temperature, pH and dissolved s ^ l t s i n defined media.  Generally, the highest growth  rates were obtained with r e l a t i v e l y high l i g h t i n t e n s i t y (550 - 600 f t - c ) , temperature of 25 - 30 C, pH near 10 and dissolved s a l t s of 5-30  °/ 00  Highest rates of growth were obtained with NO3  nitrogen source as opposed to NH^ nitrogen i n pure culture.  -  +  or urea.  Nodularia was  as the  found to f i x  iv  N o d u l a r i a was e s t a b l i s h e d i n 1822 by M e r t e n s , but not published u n t i l  1888 by Bornet and F l a h a u l t .  been 28 taxa d e s c r i b e d . The v a l i d i t y  At p r e s e n t t h e r e  have  of these and the v a r i a b i l i t y  nature and of i s o l a t e s i n c u l t u r e i n a v a r i e t y c o n d i t i o n s was c o n s i d e r e d .  validily  in  of c h e m i c a l and p h y s i c a l  An e v a l u a t i o n 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 t h a t sheath and a k i n e t e  character-  i s t i c s are v a r i a b l e while vegetative  location  and a s p e c t s of a k i n e t e  c e l l shape, heterocyst  f o r m a t i o n are more s t a b l e c h a r a c t e r i s t i c s .  On the b a s i s of the o b s e r v a t i o n s r e p o r t e d , a l l be i n c l u d e d w i t h i n two s p e c i e s , N. Bornet et F l a h a u l t i n other  genera.  the d e s c r i b e d t a x a c a n  spumigena ^Mertens i n Juergens7  1888 or N. harveyana (Thw.) T h u r e t  1875,  or p l a c e d  V  TABLE  OF C O N T E N T S PAGE  ABSTRACT  i i i  LIST  OF T A B L E S  LIST  OF F I G U R E S  LIST  OF A P P E N D I X  vi vii TABLES  ix  ACKNOWLEDGEMENTS  x  INTRODUCTION  1  ECOLOGY  3 Methods  4  Results  9  DISTRIBUTION  27  GROWTH AND P H Y S I O L O G Y  36  Materials  and  methods  Results  45  TAXONOMY  65  DISCUSSION  105 Ecology  and  Physiology  Taxonomy LITERATURE APPENDIX  36  113  CITED  TABLES  105  118 I  -  X  138  vi  LIST  OF  TABLES  TABLE I.  PAGE Locations of  Nodularia  collections,  7  1971-1973 II.  III.  IV.  V.  O r i g i n of i s o l a t e s used i n physiology experiments Light intensity experiments  used in  M i c r o b i o l o g i c a l media p u r i t y of c u l t u r e s  growth  and  culture  for  checking  Comparative morphology of Nodularia in axenic culture, contaminated c u l t u r e and n a t u r e  37  42  44  47  Vii  LIST  OF F I G U R E S  FIGURE  PAGE  1.  Study  areas  2.  Seasonal  changes  in  saline  3.  Seasonal  changes  in  temperature,  organisms  in  for  the  interior  of  Columbia  ponds  1972-1973  in  of  ponds  6 12  salinity  Ctenocladus  and  14  Pond  4.  Some f e a t u r e s  5.  Seasonal changes i n temperature, s a l i n i t y and 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 Lake  20  6.  Seasonal  and  21  and  24  changes  organisms  for  saline  British  in  17  temperature,  1972-1973  in  Inks  salinity Lake  7.  Seasonal changes i n temperature, o r g a n i s m s f o r 1972-1973 i n W h i t e  8.  North  American  distribution  of  N.  spumigena  30  9.  North  American  distribution  of  N.  harveyana  30  10.  European  distribution  of  spumigena  32  11.  European  distribution  o f _N« h a r v e y a n a  32  12.  World ( e x c l u s i v e of North America d i s t r i b u t i o n of N. spumigena  and  Europe)  34  13.  World ( e x c l u s i v e of North America d i s t r i b u t i o n o f N. h a r v e y a n a  and  Europe)  34  14.  Standard curves c e l l numbers  15.  Quality  16.  Comparison of cultures with  17.  Effect  of  temperature  18.  Effect  of  light  19.  Effect  of  pH o n t h e  of  light  of  N.  salinity Lake  optical  used in  growth of bacterial  density  culture  versus  experiments  axenic cultures contamination  on  isolates  intensity growth  of  on growth of  40  42  and  47  Nodularia  49  of  Nodularia  Nodularia  52 54  v i i i  FIGURE  PAGE  20.  Effect  of  salinity  on growth  21.  E f f e c t of s a l i n i t i e s u s i n g s a l t s N a C l on g r o w t h o f Nodularia  other  22.  E f f e c t of Nodularia  growth  23*  E f f e c t of s a l i n i t y , y l e n e r e d u c t i o n by  24.  The the  25.  Possible explanation for i s t i c s of N. armorica  26.  Comparison of the v a r i a b i l i t y of s i z e a t i v e c e l l s , h e t e r o c y s t s and a k i n e t e s  nitrogen  source  of  Nodularia  on t h e  than  59  of  61  acet-  64  s p e c i e s 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 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)  69  isolates  under  pH a n d t e m p e r a t u r e Nodularia  57  a variety  character-  of  culture  of  N.  of of  vegettwo  86  88  conditions  27.  Comparison of c e l l N. sphaerocarpa  28.  Akinete v a r i a b i l i t y i s o l a t e s as r e l a t e d  29.  Comparison of N. harveyana  30.  Cultural  31.  The  morphological  variation  of  N.  spumigena  101  32.  The  morphological  variation  of  N.  harveyana  103  cell  variation  sizes  akinete  on  of to  three B r i t i s h salinity  sizes  of  harveyana  of  N_.  and  Columbia  spumigena  and  Nodularia  91  94  96  99  ix  LIST OF APPENDIX TABLES  TABLE  PAGE Procedures used i n water analysis  138  Water chemistry f o r Ctenocladus Pond  139  Water chemistry f o r Wallender Lake  140  TV  Water chemistry f o r Inks Lake  141  V  Water chemistry f o r White Lake  142  I II III  VI . Water chemistry f o r the lakes i r r e g u l a r l y ' sampled or sampled only once ;  VII VIII IX X  World d i s t r i b u t i o n of Nodularia  143 144  Medium BG-11 (Stanier et a l . 1 9 7 1 )  157  Herbarium material of Nodularia examined  158  Synonomy of Nodularia  i**  X  ACKNOWLEDGEMENTS  I  would  guidance my  to NRC  persons  them  I  to  the  for  thank  course of  their  lent  Janet this  comments  freely  J.R.  Dr.  of The  provided help  in  the  field,  errands  and  aided  well  as  most  as  for  and t o  criticisms  was  time  of  help  and  members the  thesis.  and a d v i c e  s u p p o r t e d by  of  and  funds  from  Stein. Carolyn  study  Stein  study,  and  study  to  ran  R.  equipment,  gratitude  worried,  the  to  am g r a t e f u l .  A1035 My  She  during  committee  Many  like  is  insufficiently typed, in  many  glued, other  supplying understanding  difficult  of  times.  and  expressed xeroxed,  aspects smiles  of at  here.  retyped, the even  1 INTRODUCTION  Drotiet and D a i l y ' s green algae pretation  caused  considerable  inherent  to a v a r i e t y  have b e e n d e s c r i b e d (Golubic to  1969) and e v e r y  year  more  Physiological  and b i o c h e m i c a l  i f an i n c o r r e c t  plied.  Anacystis  point.  The commonly  been 1968;  algal  species with  confusion.  value  regarding  taxa  of i n t e r p r e t a t i o n s . ' Many as new  nidulans,  taxa  1970).  by r e s e a r c h e r s  who  has been  justification  a r e p r o p o s e d to add  c h a r a c t e r i z a t i o n a r e of  a coccoid isolate  binomial  of K r a t z  However A n a c y s t i s  are apparently  i s ap-  form, i s a case i n and A l l e n has  shown t o be a S y n e c h o c o c c u s s p . (Padmaja Komarek  inter-  Cyanophyceae  little  or unacceptable  studied  blue-  of the Cyanophyceae.  i n bluegreen  the taxonomic  little  of the c o c c o i d  controversy  o f taxonomy and m o r p h o l o g y  The v a r i a b i l i t y subject  (1956) r e v i s i o n  and  nidulans  Desikachary i s employed  u n w i l l i n g to use t h e c o r r e c t  binomial. Taxonomic culture that  work i s o f l i t t l e  and i n n a t u r e  both  field  complementary (Pringsheim  i s taken  value  unless  into  account.  It i s  and l a b o r a t o r y s t u d i e s be c o o r d i n a t e d  to g i v e  1967).  insight  Despite  into  the b i o l o g y  the acknowledged  type  o f an a p p r o a c h f o r t h e b l u e g r e e n  only  one s t u d y  observations,  variation  (Kann and Komarek cultural  study  algae  important and  o f an a l g a  need  for this  (Desikachary  1970), u s i n g  and a c r i t i c a l  both i n  ecological  review  of the  1970),  literature, for  has  been  such studies The  genus  present  1972,  delimit  the  large  studying its ered  in  as  /Mertens  in  of  the  JuergensV  The  genus  such as  Nos t o e sense.  occurrence and  three  its  this,  filamentous,  species.  generic  of  the  need  apparent.  undertaken  the  growth,  light  was  a  p.  In  240)  genera,  defined  are  investigation  Nodularia  Stafleu  reported.  and  or  is  to  more  study  The  problem  distribution,  chapters;  et  Flahault  the  and was  and  than  the  appears  variation.  well-  approached  factors  (see  genus  manageable  Anabaena,  morphological  separate  Bornet  heterocystous  by  controlling  These  are  Ecology; Physiology;  consid  Taxonomy  3  ECOLOGY  Of  primary  ining  its  iated  with,  at  a  role  near  ponderosa and the is  in  pine  atures  -  winter. with  29  a n d why  it  in  by  the  area  Artemisia low  as age 12  27.4  snow i n daily  Kamloops has snow i n  average  eratures  was made  of  in  in  a  organism i s is  particular  the  and as  average 0 and  -6  daily  of an  are  rain  average or  Nutt.  -2  of  in  maximum  24.3  January  defassoc-  place  of  May  or  -9  C.  with  lower  areas  most  occuring  cm/yr  Nodularia  preciprain  temperand  Penticton most  falling  The are  29  and minimum J a n u a r y  1972;  vall-  Both  June.  of  the  I965)  (Krajina  maximum  temperatures  Department  in  and most  daily  C and  British  are  a n d minimum J u l y  average  July  which  zone  precipitation,  May-September  ponds i n  areas  ponds and t h e  precipitation,  C (Canada  sampling,  four  interior  These  the  maximum  maximum a n d m i n i m u m  Periodic  an  organisms i t  tridentata  C and the  average  December  are  were  December  daily  C a n d 13  cm/yr  C and the  what  occurs  of  amounts  minimum J a n u a r y t e m p e r a t u r e s receives  of  bunch grass b i o g e o c l i m a t i c  most  The  are  study  environment:  sampling areas  relatively  June.  the  Kamloops and P e n t i c t o n .  dominated  itation,  in  time.  vegetation  receive the  the  a n d when  primary  Columbia  in  in  particular The  eys  importance  averC  and  temp-  Transport).  April-July regularly  1973,  occured.  4  Three o f t h e s e ponds ( C t e n o c l a d u s Pond, W a l l e n d e r L a k e ,  Inks  Lake) are l o c a t e d southwest o f Kamloops and one (White Lake) l o c a t e d southwest o f P e n t i c t o n ( F i g . l ) .  B l i n n (I969) has d o c -  umented some a s p e c t s o f C t e n o c l a d u s Pond and W a l l e n d e r L a k e . In a d d i t i o n to these four h a b i t a t s , ically,  which were sampled p e r i o d -  a number o f 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 o r o n l y on a s i n g l e o c c a s i o n ( T a b l e 1 ) .  Methods: A t each s a m p l i n g t h e t e m p e r a t u r e  and d i s s o l v e d oxygen were  measured (YSI model 54 oxygen m e t e r - t h e r m i s t o r and a g l a s s thermometer);  pH r e c o r d e d (Metrohm model E280A pH m e t e r ) ;  water ( d u p l i c a t e 500 ml s a m p l e s ) , p l a n k t o n and grab samples c o l l e c t e d and g e n e r a l o b s e r v a t i o n s made (water l e v e l s , present,  species  and e s t i m a t i o n of abundance o f organisms on a s u b j e c t -  i v e b a s i s s i n c e a t t e m p t s a t q u a n t i t a t i v e measurements extremely d i f f i c u l t ) .  The w a t e r samples were  proved  filtered  t h r o u g h Whatman (#1) and M i l l i p o r e (.45 Um) f i l t e r s ,  within  s i x h o u r s o f c o l l e c t i o n and m a i n t a i n e d a t 5 C f o r r e t u r n t o the laboratory.  I n t h e l a b o r a t o r y t h e organisms p r e s e n t  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 t h e 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 , ethanol.  were  Analyses of ortho-phosphate,  L u g o l ' s i o d i n e o r 70% 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 o u t , w i t h i n 48 h o u r s , a c c o r d i n g  5  F i g u r e 1.  Study areas i n the i n t e r i o r  of B r i t i s h Columbia,  s c a l e 1:62,500 a.  B r i t i s h Columbia  1 i n c h = 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. ' S a l t Pond 4» 9. S a l s o l a Pond 10. ' S a l t Pond 3' 1 1 . S a l t w o r t Pond  c.  South west o f P e n t i c t o n B r i t i s h Columbia 12. White Lake 13. Mahoney Lake 14. Green Lake  7  T a b l e 1. A.  B.  Locations of Nodularia  Locations regularly  collections,  1971-1973.  sampled  1.  C t e n o c l a d u s Pond 50°39'N, 120°32'W N o r t h o f T r a n s - C a n a d a Highway (BC Hwy 97), 9 mi west o f Kamloops B.C. C i t e d b y B l i n n ( 1 9 6 9 ) a s ' C h e r r y C r e e k Pond'  2.  W a l l e n d e r Lake 50°38'N , 120°26.5'W E a s t s i d e o f L a c l a J e u n e Road, 3 mi f r o m i t s j u n c t i o n w i t h t h e T r a n s - C a n a d a Highway (BC Hwy 97)  3.  Inks Lake 5 0 ° 3 7 ' N , 120°27'W West s i d e o f L a c l a J e u n e Road, 4 mi f r o m i t s j u n c t i o n w i t h t h e T r a n s - C a n a d a Highway (BC Hwy 97)  4.  White Lake 49°19'N , 119°38'W One m i l e s o u t h w e s t o f t h e D o m i n i o n R a d i o A s t r o p h y s i c a l O b s e r v a t o r y a n d 5 mi s o u t h o f P e n t i c t o n B.C.  Locations 5.  irregularly  or singly  sampled  ' S a l t pond 4' 50°40'N , 120°35'W A s m a l l unnamed pond s o u t h w e s t o f S a l s o l a Pond ( S a l s o l a Pond c i t e d b y B l i n n a s S a l t M i n e Pond  1)  6.  ' S a l t pond 3' 50°40'N , 1 2 0 ° 3 4 ' W A s m a l l unnamed t e m p o r a r y pond between S a l s o l a Pond a n d S a l t w o r t Pond ( S a l t w o r t Pond c i t e d b y B l i n n I969 a s S a l t M i n e Pond 2)  7.  B i g Q u i l l Lake, Saskatchewan 5 1 ° 3 0 ' N , 104°40'W N o r t h o f K a n d a h a r on Hwy 14 ( S a s k . ) a n d 100 mi east of Saskatoon  8.  Devils  9.  S a n de F u c a , W a s h i n g t o n 4 8 ° 1 0 ' N , 122°48'W A d j a c e n t t o W a s h i n g t o n Highway 525  10.  L a k e , N o r t h D a k o t a a t Camp G r a f t o n  R i e f e l W i l d l i f e Refuge Near L a d n e r B.C.  49°08'N  48°N  , 99°W  , 123°10'W  Continued...  8 T a b l e 1 (Contd.)  .  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 , 1971-1973  C u l t u r e s were i s o l a t e d f r o m s o i l samples c o l l e c t e d a t the f o l l o w i n g l o c a t i o n s 11.  S p o t t e d Lake 49°05'N , 1 1 9 ° 3 4 » W I n R i c h t e r P a s s , 6 mi n o r t h w e s t o f Osoyoos B.C. on B.C. Hwy 3  12.  S m a l l unnamed pond 1 1 9 ° 3 8 ' N , 49°09'W I n low a r e a 1 mi s o u t h o f R i c h t e r Lake, 4 mi southwest o f S p o t t e d Lake a d j a c e n t t o Hwy 3  13.  Bowers Lake 50°40'N , 120°26'W A t t h e j u n c t i o n o f Trans-Canada Highway (Canada Hwy 1 , BC Hwy 9 7 ) and Lac l a Jeune Road  14.  Water h o l e Simpson's Gap, N o r t h e r n T e r r i t o r i e s , Australia 20°45'S , 134'E C o l l e c t e d by D r . J.R. S t e i n , 18 i x I969  9  t o Standard Methods (A.P.H.A. I965).  The one e x c e p t i o n t o  t h i s treatment of water samples i s t o be found i n t h e data from August  1972 when t h e water was not a n a l y z e d u n t i l two  weeks a f t e r c o l l e c t i o n . c a t i o n s (Na , Mg , Ca (model  Analyses of t h e water f o r t h e major , K ) were made with a Perkin-Elmer  303) atomic a b s o r p t i o n spectrophotometer  anions (C0^  }  HCO^, SO^, C l ) by Standard Methods from water  samples s t o r e d at 5 C. as Appendix,  and f o r major  D e t a i l s of water a n a l y s i s a r e g i v e n  T a b l e 1.  When r e f e r e n c e i s made t o s a l i n i t y i t i s i n t h e broad sense of t h e amount o f d i s s o l v e d s a l t  c o n t a i n e d i n t h e water  without any i m p l i e d i o n i c content r a t h e r than i n t h e narrower oceanographic sense of c h l o r i n i t y .  The term b r a c k i s h i s  used i n r e f e r e n c e t o d i l u t e sea water,  and t h e term  saline  r e f e r s t o i n l a n d water bodies of s a l t content g r e a t e r than  0.5 °/oo.  Results: In t h e s p r i n g ( l a t e March and A p r i l ) t h e l a k e s a r e at t h e i r l a r g e s t volume and c o n t a i n r e l a t i v e l y low amounts of d i s s o l v e d s a l t s due t o t h e h i g h volume o f water  resulting  from snow-melt and drainage i n t o these d e p r e s s i o n s .  As  s p r i n g and summer p r o g r e s s , t h e s a l i n i t y i n c r e a s e s as water volume decreases due t o e v a p o r a t i o n i n excess of r a i n f a l l and  10  runoff.  In  late  the  ponds r e a c h  dry  up  summer  their  completely  The described  by  minimum  (Fig  dynamics of Blinn  and f a l l  (late  volume  the  ions  in  saline  In  (1971)*  any  can be  a great  year  water  levels  of  For  example,  in  precipitation. levels tion  in  were  1973  rophic  ponds are  growth,  (April-July)  lower  it  than  ponds of  deal  these was  this  of  it  type  is  must  be  variation  ponds depending observed that  at  1972  diversity as  of  the  variations  characterized  principally  followed  invertebrates  A.  instances  from on  total  water  corresponding  collec-  dates. The  ses  some  September)  interpretation  there  year  and i n  and  2).  emphasised that to  August  algae,  organisms  salinity single  from  occurs  ponds must  Ctenocladus  in  large spring  July  in  To  be  amount and  growth  to  spring  increases.  regularly  a  a heterotrophic  and b a c t e r i a ,  Locations 1.  by  by  of  autot-  early  summer  phase,  September. and  examine  Maximum  diversity these  grazing  decrea-  trends  and  considered.  sampled.  Pond has  Na  and SO.  +  as  the  dominant  4 ions. summer to  The  pond n o r m a l l y  and t h e  more  dominant  than alga  salinity  400 is  °/oo the  in  dries  up  may v a r y the  fall  chlorophyte  during  from  the  course of  30-50 ° / o o i n t h e  (Appendix, Ctenocladus  Table  II).  circinnatus  the spring The Borzi.  11  F i g u r e 2.  Seasonal changes i n s a l i n e  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  ponds  13  Nodularia in  small  algae  harveyana numbers  present  tenerrima Teodor. algae  were  sp.,  Ulothrix  Ag.  of  in  and  were  July,  the  latter  Cladophora  and U l o t h r i x .  the  but  layer  in  less  d e p o s i t e d by Around the  rubra  Nels.  Ruppia quito  larvae  Artemia  the  L.  (Aedes  pond i s  salina  In sp.)  and  sp.;  diatoms  sp.  All  April were  A mat  but  largest  reduced in Lyngbya  former an  in  the  the  the  April most  disappear-  maximum  on dead  remained  pockets  or  dying  throughout  under  the  salt  water.  pond i s  a  is  zone  large  large  Salicornia growth  numbers  numbers  purple  of  a dense  observed, J n June  of  or  isolated  and h i d d e n  a n d May  by  in  summer,  reached  epiphyte  Lyngbya  Dunaliella  numbers  the  Ag. ,  Amphora  numbers  during  pond i t s e l f  inhabited  Leach.  Gomont  evaporating  the  bluegreen  (Menegh.) the  Other  (Dunal)  C.A.  quantity  of  the  May  Ulothrix  salina  Kutz;  Ctenocladus  shoreline  and w i t h i n  maritima  August  the  as  sp.,  brevis  and  the  and  0.  salinity  Dunaliella  June or  Spirogyra  Ag. ,  their  greatly  April  June-September.  Dunaliella  C.A.  in  during  (Dillw.)  Navicula  increasing  algae  open w a t e r ,  summer  algae  Euglena  present  completely.  numbers of  With  an  only  from  sp.,  amphibia  Wille;  and Lyngbya were  species  green  Phormidium tenue  coffeaeformis  and May.  absent  Cladophora fracta  nordgaardii  ed  the  Oscillatoria  Anabaena  collected  a n d was  Kutz.,  and  was  of  of  mos-  through of  sulphur  brine  shrimp,  bacteria  14  30 .25 •  u a »s  20 15 10  5 0 400 r o o  300  CO  200  100  0  fc  Ctenocladus  w e w •rl c bo c o •p c ca  Nodularia  Aedes  c  E o o  Artemia  Dunaliella  1972  APR  Figure 3 .  MAY  JUN  JUL  AUG  SEP  1973 APR  MAY  JUN  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 t h e organisms i n d i c a t e d by t h e v e r t i c a l bars  are  JUL  AUG  15  covers  a thick,  cod and the pond are  and  black,  rotifer,  egg c a s e s ,  around the  dynamics  of  the  dissolved  salts  larvae  more  and  edge.  An u n i d e n t i f i e d  is  adult The  important  Lake  and  mud.  Brachionus p l i c a t i l i s  water's  Wallender  2.  anerobic  Muller,  salt  flies  periodicity  organisms i s  characterized  less variable  salinity  inhabit  a  and  population  lower  range  Fig.3« amount  Ctenocladus Pond.  a n d SO^ these  (Appendix,  lower  organisms icornia the  Table of  is  peripheral  mat  is  in  present  aria  the  is  community  geminata  all  cells  the  in  by  May  of  algae  Menegh. ,  0_. t e n u i s  C.A.  a  of Sal-  margin  of  dominant (April  rapid  and  growth  characteristic N.  and  harveyana) typically  D u r i n g May, of  the  cells  filaments  Associated with c o n s i s t i n g of  Ag. ,  ,  variety  spring  and June  few  Mg of  The  (primarily  J u n e many  .  result  around the  early  very  Na  -H-  c o n s i s t of  D u r i n g May  akinetes.  bluegreen  a  itself.  for  July  of  as  larger  C l a d o p h o r a mat.  and by as  L.  Nodularia  vegetative,  akinetes  nearly  lake  are  plants  formation  maximum n u m b e r s  with a  the  (Fig.4c).  primarily  converted to  the  Cladophora f r a c t a . in  a  vascular  season except  associated with  is  The  ions  Probably  and S c i r p u s m a r i t i m u s  Cladophora results  occurs  III).  and Ruppia w i t h i n  May),  principal  dissolved salt,  present.  throughout  early of  are  rubra  lake  alga  levels  The  of  (l4-88°/oo)  + than  the  (Ephedridae)  shown i n  by  ostra-  are  Nodulhave seen,  Nodularia  Oscillatoria  0_. n i g r o v i r i d u s  Thwaites,  16  Figure a.  4»  Some f e a t u r e s Wallender the  of  Lake w i t h  shoreline  (iv  the  large  A c l o s e r view of diatom o x i m a t e l y 1 0 cm l o n g )  c.  Peripheral ender Lake  d.  Shoreline  e.  of  numbers  bloom i n  C l a d o p h o r a mat (vi 1973) Wallender  bacteria  ponds of  studied. diatoms  along  1973)  b.  sulphur  saline  a.  around the  Lake w i t h  and S c i r p u s  (vi  (pencil  edge  of  Cladophora,  is  appr-  Wall-  purple  1973)  Inks Lake showing d e n s i t y of b r i n e shrimp ( v i i 1973). A l s o present are Cladophora and Ruppia (thermometer i s a p p r o x i m a t e l y 3 5 cm l o n g )  18  Spirulina  major  nordgaardii most  other  Kutz.  appears  early  Cladophora  mat,  remarkable  bloom  coffeaeformis are  in July  bluegreen  During  present  and Phormidium  (April)  before  a diatom  maximum  occurs.  of  diatoms  other  occurred  times  of  Cladophora  and t h e bluegreen  and two U l o t h r i x  Duj.  found  i n much l e s s  (July  1972)  Ceratium in  localized  spring  also small  animal  of  Ceratium  Lake  the  Amphora  These  by r a p i d  diatoms  reduced growth  Kutz. (O.F.M.)  appearance  since  it  is  in  A species of  on t h e s h o r e l i n e  time  a large Euglena  such  normally  A t t h e same  (see below)  of  Nodularia  hirundinella  Its  a  1973  and U.? aequalis  1973.  present. areas  April  associated with  interesting  Inks  of  b u t i n much  conditions.  community  (Cladocera),  (Rotatoria) spring. numbers  of  number was  of  found  mud t h r o u g h o u t  a n d summer.  The sp.  i n nearby  were  is  saline  In  followed  tenerrima  number  ° / o o ) waters  (48  is  algae  was o b s e r v e d b u t n o t i n  saline  the growth  (principally  the year  maximum  s p p . , U.  a large  1972  Lyngbya  i n numbers  spp.) (Fig.4a,b).  The diatom  July  decline  spring  numbers.  In  the  Gamont.  algae.  and N a v i c u l a  at  after  fragile  Diaptomus  late  salina  spring  the  sp.  and Chironomidae,  Artemia in  of  lake  is  composed of  (Copepoda) , all  present  (Branchiopoda)  a n d summer  a  Brachionus primarily  is  ( F i g . 4e).  present  in in  Moina plicat; .lis !  the large  Ephidridae  19  are  present  iment  is  around  c o v e r e d by  The  periodicity  The  maximum v a r i e t y  summer  By  number  of  late  Inks  IV)  ities  (Fig.6).  and the  validus are in  the  at  Vahl.  is  noted  Inks  Lake  salinities  evidently Evidence to  is is  further 4.  and  harveyana  Lake  sed-  (Fig.4d).  as  spring  Fig.5.  or  early  increasing reduces  to  higher plant  and N.  60  the  Lake pH  sal-  the  animal  spumigena  rather  Nodularia  Kutz.)  are  pattern and  commun-  Scirpus  variabilis  the  slightly  Cladophora  of  present occur-  disappearance  increasingly  °/oo,  section  has  (plus  spring  most  differs  (Appendix,  and  same  in  becoming  but  peripheral  Anabaena  growth  salinity  in  Inks  The  Lake  Lake  components  numbers.  approximately  support  shown  the  Wallender  animal  water  presented  is  salinity  distinctive  greatest  related  White  to  cyanophyte  the  in  bottom  bacteria  response to  similar  and W a l l e n d e r of  in  extreme  has  same  large  with  organisms  and a s l i g h t l y  plant  with  sulphur  some d e t a i l s .  B o t h N. in  of  similar  and t h e  spring  purple  and the  minimum.  in  The  same  Nodularia  both  Lake  the  a  is  lake  species occurs  otherwise  present.  ence of  but  of  the  decreases  dissolved salts  Table  mat  of  summer  Lake  from Wallender  of  and dynamics  species to  3.  edges  a mat  and d i v e r s i t y  inity.  less  the  saline.  In  disappears  disappearance  than  pH o r  on growth  and  ionic  changes.  physiology  this. is  100  miles  southeast  of  the  other  three  20  30  — 1972  Figure  5«  •  MAY  •  '  JUN  JUL  '  AUG  .  SEP  f  I  L-  APR  ,  MAY  i_  JUN  Seasonal changes i n temperature, salinityorganisms for 1972-1973 i n Wallender Lake  21  Cladophora  Diaptomus Moina  •  '  1 2 9 7  Figure  6.  m  Y  ZZ]  L  • JUN  *  JUL  |  •  .  AUG  SEP  /  / 1  9  7  3  , APR  MAY  , JUN  Seasonal changes i n temperature, s a l i n i t y o r g a n i s m s f o r 1972-1973 i n I n k s L a k e .  , JUL  and  ,_ AUG  22  lakes.  The  dominant  ions  are  Na ,  SO  +  a n d CO ; 'r }  inity  varies  Table  V).  from  The  the  algae  at  the  at  the  being  Tydb.  harveyana  of  the  from  occurs  in  Lake  absent  has  concentration,  The  factor  the as  the  destruction  well  water  as  in  with  tenuis,  0.  the  lake  is  does  of  other the  Nodularia  side  of  of  range not  early  the  of  habitat  to  pattern as  the  summer  the  depends on t h e  lowest  vascular  the  bluegreen  Phormidium r e t z i  Lyngbya a e s t u a r i i  (Mertens)  of  salin-  factor.  Nodularia  recedes  level The  algae  and  relatively  limiting  plants.  cattle  the  presence of  algae,  a  (May-June)  and  line  bases  Nodularia  growth  water  (both  in  However  of  The  times  around the  lake.  be  stricta  at  prints  mat  three  Nodularia  year.  plants.  are  both  fluctuation  appear the  the  other  Distichlis  hoof  mat  bacteria  the  o c c u r s on and in  different  peripheral  sulphur  itself.  and  s a l -  (Appendix,  notably  no  the  rest  chalbea Mertens,  sp.,  purple  spring to  vascular  above  present  Anabaena  the  most  being  is  of  plants  responsible for  bases of  are  characterizes  south  a narrow  low  is  9.4-10.0  as  spumigena)  on t h e  White  generally  no  waterbody  vascular  area  which  the  and N.  emergent  corralling  margin  Lake  There  and  interface  pH i s  and S c i r p u s n e v a d e n s i s W a t s . ,  emergent  N.  White  lakes.  shoreline  Around the  (Torr.)  and the  organisms of  mud-water  lakes.  ity,  15-40 °/oo  previous three  of  is  from  the  si  beyond  Nodularia,  of  the  other  algae  Oscillatoria  (C.A.  Liebmann  Ag.)  Gomont,  and  Spirulina  23  major;  and t h e  cinnatus.  The  iodicity is  green  and  present  algae  preceding  limitations  only  in  Spirogyra bluegreen  as  April  sp.  and  algae  Nodularia.  a n d May  Ctenocladus  show t h e The  and the  same  Spirogyra  Ctenocladus  cirper-  sp.  June  in  or J u l y . In A p r i l anders  the  1973  (Ambystoma  diatoms oceran  Amphora Moina  sp.).  sp.  and  a  1972  a  large  washed  shore.  The  White  B.  Lake  is  shown  Locations  1972  at  Na ,  SO^  +  a  Oscillatoria 'Salt  alga  in  salinity  1972  May  (Appendix, 7.  °/oo,  the  of  at  Table  Big  a  was  the  cladIn  floating  and  dynamics  of  0_.  with Table  The  circinnatus  tenuis  contained salinity  of  spumigena  primary  VI).  with  and Anabaena N.  4  spumigena °/oo  once.  (Na , +  in  ions  May  being  dominant lesser  alga  amounts  sp. as  the  SC^,  dominant  C0~)  VI).  Quill 26  47  brevis, 3'  of  Diaptomus.  sampled only  Nodularia  Ctenocladus  Pond  the  population  contained  (Appendix,  of  and  sp.  salam-  consist  spp.;  genus  Nostoc  sampled or  and  p o n d was  the  of  Fig.7«  of  this  of  numbers  organisms  Navicula  periodicity  salinity CO^  and  amount  as  large  plankton  copepod of  P o n d 4'  of  6.  The  irregularly  'Salt  5.  contained  coffeaeformis  September on  lake  °/oo  Lake, of  Saskatchewan,  predominantly  Na  in +  August a n d SO^  1972,  had  (Appendix,  a  24  30  25 20  a 15  B  9  EH  10 5 0 50  o o  •P  25  •rl  e  •H  H  CO CO  N. spumigena  e  00 •H C «J .  N. h a r v e y a n a  bs  O  Lyngbya  •P  c ca  e a o  •rt  o  Diaptomus Moina  Nostoc  1972 Figure  7.  MAY  JUN  Seasonal organisms  J L.  JUL  AUG  changes for  in  SEP  temperature,  1972-1973  in  White  1973 salinityLake.  25  Table  VI).  ter.  Nodularia  Associated with  Boyer  and  epods  (Diaptomus  sp.  large  which  grew  detached 8. (Na ,  numbers sp.).  August  Kutz.,  Cladophora  sp.  harveyana  18  Nodularia  C.  and  Enteromorpha Fuca,  °/oo  (Appendix,  Percursaria  was  with  an  large  elmorei and  cop-  Enteromorpha numbers  of  shore. had a  salinity  Table  VI).  open water  12°/oo  of  Nodularia  where  Morren  shoreline  it  was  and M i c r o c y s t i s  associated  with  sp.  Washington,  c o l l e c t e d from  was  a brackish Table  VI).  in  May  (Na , +  1973  Nodularia  Cl  pond  )  Associated with  percursa  (C.  Refuge  a  Ag.)  Rosenvinge  of the and  aestuarii.  10.  Riefel  had  and  also  a  Wildlife  salinity  contained  Hugeotia  sp.  and along the  San de  was  alga  (Appendix, from  (Corixidae)  Aphanizomenon f l o s - a q u a e  aeruginosa  Near  lee  phytoplank-  Chaetoceras  boatmen  Dakota,  collected both  principal  diatom  bottom  on t h e  1972  the  dominant the  North  associated with  a  to  Lake,  was  office  The  attached  the  water  Devils  spumigena  Lyngbya  of  up  in  salinity  was  blown  4  9.  it  plants  S0 )  +  VI)  s p u m i g e n a was  sp.,  being  Isolates Blinn  of  8.4  Nodularia  with  °/oo  -  pond near  (Na , +  harveyana.  Lyngbya a e s t u a r i i  Cl  )  The and  an  the  refuge  (Appendix, dominant  Table  alga  Oscillatoria  present. from  (1969)  soil  (See  reports  N.  also  Table  I)  spumigena from Wallender  Lake,  was  26  Ctenocladus Salsola  Pond,  Bowers  Pond and S a l t w o r t  Wallender  Lake  and  from  1971-1973^  salt  from  the  techniques  however  dry  lake  (Allen  Spotted  Lake  ment  of  an A u s t r a l i a n  also  yielded  an  Nodularia Salsola  collections 100 ible  °/oo) in  failed  to  was  produce  from any  were  Bowers  in  an  Lake  sample  of  N.  Saltwort  these  be  from  Pond,  salinity  to  of  in  Richter  collected  of  dry  soil  and  N_.  spumigena.  isolates Lake.  near  from  Enrich-  Alice Springs  spumigena.  collected  the  of  was  from  Enrichment  isolate  resulted  Lake,  obtained  akinetes.  pond south  soil  Nodularia  soil  revealed  resulted  never  made  Pond.  Ironmask  microscopic examination  small  Pond or  Isolates  and t e c h n i q u e s  isolate  were  for  the  a  Pond.  bed  1973)  observations and  Polygon Pond,  Ctenocladus  Similar  Lake,  Lake,  was  isolates.  and  Pond,  p o s s i b l y because  growing.  sites  Polygon  too No  high  when  (greater  akinetes  enrichment  Ironmask  were  than vis-  procedures  DISTRIBUTION  Some as  being  ination crete  individual  (1932)  distribution  Umezaki  (I96I)  without  any  restricted. and never many  tropical. in  in  a  of  general  4  pH  restricted  a  marine  plankton  ments,  tolerances  ution  patterns  inent  data  as " s u b c o s m o p o l i t a n "  rarely  algae  found  in  Dawson  differentiated  species).  is  or  hot  a n d more  have  preferences.  acid  into  more  The  states  the  that  temperate  and  algae  found  of  are  tropical  subtle  nature  very  situations  (I966)  misidentification,  particularly  are  spring species or  Others  o b s c u r e d by  and  prefix.  1973)•  (some  describes  cosmopolitan"  s p e c i f i c s e n s e many b l u e g r e e n  habitats  dis-  distribution  (1955)  sense bluegreen  (Brock  s p e c i e s may b e  as  very  exam-  point.  describe the  harveyana  Cyanophytes are  In  this  Chapman  algae  closer  illustrates  (1943)  the  However, reveal  harveyana  d e s c r i b e s N.  below  marine  N.  bluegreen  s p e c i e s may  cosmopolitan.  explanation  However,  or  Nodularia  of  most  distribution.  and L i n d s t e d t  spumigena as  regard  1945)  genera  distribution.  N.  the  (Fritsch  cosmopolitan in of  Gietler of  authors  require-  these lack  of  confused taxonomic  distribpertsit-  uation. The review  of  distribution the  of  literature  Nodularia and  was  s t u d i e d by  examination  of  a  herbaria  critical material  28  (Appendix, the  great  tropical  areas  the  areas;  had  Some o f  it  temperate  regarding  the  from  obvious  from  ularia  is  the  revealed  1937, 1938;  Others  1893;  Arctic  (Apstein,  and West  1911;  uncommon i n  not  from  Field  have  in  been  in  Field  of  reported  Wheldon  Wille  Nod(Des-  1949). higher  more  Herbarium; information  1957).  but  1897)  it  is  (Croasdale 1924)  (60°-90°  of  Herbarium;  surveys  climates  temperate  Nordstedt  and S u b a r c t i c 1947;  not  surveys  Skuja  contain  cited  collection  polar  have  latitudes  are in  equator,  Woodhead and Tweed  (Foslie  1959)  Arctic  do  sub-  presence  1929;  Standey  and  more  details  tropical  Fremy  that  the  as  the  Nodularia  1968;  (Mobius  Fukushima  of  is  more  habitat.  New E n g l a n d c o a s t lakes  have  in  show  areas  collections  would  (Karim  of  tropical  sufficient  1914).  other  south  been  of  survey  temperate  have  reports  the  in  and  that  extensive  this  were  north  argued  of  that  Nod-  north  or  south  distribution  is  the  equator).  What type  Drouet  Antarctic  West  as  which  altitude  1912;  of  of  West  Fritsch  1973;  may b e  altitudes  Collections  collections  25°-60°  tropical  1908;  Tilden  results  occurred frequently  1959; the  of  there  areas  ikachary  or  It  subject  tropical  The  (i.e.  however,  ularia  IX).  majority  8-13).  Figs. been  Table  (i.e.,  important  perhaps  B r a c k i s h marine  in  habitats  of  North  America)  and a r i d  western  North  America)  are  (i.e.,  areas  or in  Baltic  semi-arid which  Sea, inland  repeated  29  Fig.  8.  North  American d i s t r i b u t i o n  of  N.  spumigena  Fig.  9^  North  American d i s t r i b u t i o n  of  N.  harveyana  •  herbarium  specimen  o  literature  report  30  31 Fig.  10.  European d i s t r i b u t i o n  of  N.  spumigena  Fig.  11.  European d i s t r i b u t i o n  of  N.  harveyana  •  herbarium  o  literature  specimen report  33  Fig.  12.  World  (exclusive  distribution  Fig.  13.  World  of  of N.  (exclusive  of N.  North  America  and  Europe)  spumigena  North  distribution  of  •  herbarium  specimen  o  literature  America  harveyana  report  and  Europe)  34  35  c o l l e c t i o n s have been made.  The f a c t o r s  enabling Nodularia  t o occur i n these types of waters i s the a b i l i t y of these a l g a e t o grow i n a wide range o f s a l i n i t i e s , a b i l i t y t o i s t rapid fluctuations cation.  Bristol  i n s a l i n i t y and r e s i s t a n c e  to  res-  desgic-  (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 d r y s o i l sample s t o r e d f o r 5 9 y e a r s .  Laboratory  obser-  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 a r e a l l o w e d t o d r y i n t h e c u l t u r e chamber  (15-30 C) and t h e n s t o r e d a t room  t e m p e r a t u r e , t h e a l g a l m a t e r i a l i s v i a b l e and growth  will  resume when a p o r t i o n o f t h e d r i e d m a t e r i a l o f any o f t h e 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 Because o f t h i s r e s i s t a n c e  two y e a r s s t o r a g e .  to d e s i c c a t i o n , the akinetes  p r o b a b l y d i s t r i b u t e d by w i n d as a r e many o t h e r organisms  (Maguire 1 9 6 3 ) .  iso-  small  are  aquatic  36  GROWTH AND P H Y S I O L O G Y  The lates of  growth  were  the  and p h y s i o l o g y o f  studied  isolates the  under given  study  was  of  logical  observations  conditions  of  of  obtaining  erns  of  the  of  variety  alga  in  of  The  of  into  in  by  to  salinity  aid  growing  in  the  the patt-  investig-  isolates  under  a  variables.  is  a modification  were  (1958)  tried  spp.  were  (Appendix,  but  BG-11  wide made  in  the  field  manipulation  and  repeatedly  of  with  ecoof  and growth  which  of  this  a variety  (1971),  Nodularia  agar.  some  occurrence  Second,  chemical  clarify  Stanier,  collected  the  and  sources of  of  considering the  response  isolates  pH,  the  to  iso-  purpose  BG-11  media  Removal  The  The  medium u s e d was  Gorham and Z e h n d e r  factory  the  plasticity  and  II. First  light,  Nodularia  methods:  Cohen-Bazire  other  growing  nature.  physical  and  Table  of  conditions.  twofold.  insight  morphological  Materials  and  by  in  temperature,  hope  ation  laboratory  are  portion  a number  bacterial the  on  (w/v)  cultures  agar  restreaking  across were  was the  be  the  the  aided agar  maintained  most  algal  material manual  filaments.  by  the  surfaces on  satis-  Isolations  and by  the  Hughes,  Several  used.  plates of  Mandel  from  VIII).  (6-11)  streaking  contamination  hormogonia  Stock  proved to  pH r a n g e by  1%  Table  Kunisawa,  agar  phototactic or  through  plates  or  37  Table  II.  O r i g i n of i s o l a t e s experiments. Details  isolate  of  used in  locations  are  No.  growth  given  in  isolated  Table  2  A l k a l i n e s o i l from R i c h t e r Lake B . C .  3  White  4  'Salt  5  Ctenocladus  6  'Salt  7  Inks  8  Wallender  9  B i g Q u i l l Lake Sask. (N. spumigena, form a k i n e t e s , has gas vacuoles)  B.C.  p o n d 4'  pond Lake  Spotted  (N. (N.  Lake  B.C.  (N.spumigena")  (N.  harveyana)  spumigena)  spumigena) (N.  harveyana) does  not  S o i l c o l l e c t e d at S i m p s o n ' s Gap, Northern T e r r i t o r i e s A u s t r a l i a by D r . J . R . S t e i n (N. spumigena) LB-1452/l and  from  Protozoa  Butcher  (N.  Alkaline  Culture  Collection  of  England,  isolated  harveyana,  does not  form  soil  Lake  the  Cambridge  from  B.C.  Bowers  (N.  Inks  14  White  15  Unnamed p o n d n e a r (N. harveyana)  S a n de F u c a  Washington  16  Devils  Dakota  spumigena)  Lake  Lake  (N.  Lake B . C .  B.C.  North  reasons for using these taxonomy s e c t i o n .  (N.  Algae by  akinetes)  13  # The the  of  spumigena) (N.  B.C. (N.  B.C.  harveyana")  Pond B . C .  B.C.  Lake  near a s m a l l pond s o u t h (N. spumigena)  B.C.  3'  I.  from  Alkaline  Lake  from  physiology  1  10  soil  and  spumigena)  harveyana) spumigena)  specific  (N.  names  are  given  in  38  in  liquid  culture.  Nodularia grown  in  growth  were  medium  tube For  medium  in  seven  day  of  optical  optical  are  Bellco  3 to  was  curves  trolled  Environments  umbia,  for model  20 C  aliquot  each  to  in was  and  closures. light-  1 0 0 ml  liquid  measured  data  ml  25  homogeneous  after  based  on  times. of  a  at  culture  the  20  number  isolate.  was  nm u s i n g  660  Lomb S p e c t r o n i c  related  nitrogen  containing  grown  noted  of  and  steel  Growth  three  phase  salinity tubes,  were  otherwise  varied type  Marshall, Ltd.,  Boone,  (Bell  2 0 0 2 ) .  log  were  measured either  a  spectrophotometer. of  These  cells  by  prep-  comparison  Fig.14.  reach-in  Ltd.,  the  insure  flasks.  and  for  were  Ltd.,  to  (absorbance)  a Bausch  experiments  stainless  cultures  5 ml  density  Co.  Co.  daily  replicated  (Sherer  chamber  turned  unless  shown a s  chambers:  Percival  with  Erlenmeyer  Temperatures ment  light,  each,  density  standard  curves  a  in  2 0 mm t e s t  cultures  B e c k m a n DBG o r  aring  and f i l a m e n t s  done u s i n g  were  ml  initiating  Temperature,  period  Growth  The  used for  pH e x p e r i m e n t s 250  duplicate  the  in  racks  ing.  as  flasks  were u s e d .  liquid  a  ml  500  experiments  Test  cultures  by for  growth 15  C,  Michigan, Winnipeg,  Iowa,  Craft  model  in 25  controlled C,  model  C and  30  35  RT-18B-5E;  Manitoba,  model  I-36-L) a n d a  Industries,  environ-  Surrey,  C  Con-  T18L; walk-in  British  Col-  39  Figure  14.  Standard numbers. a.  The from  curves  group 8-12  of  of m  relationship  optical  isolates (i.e. between  density  with  N.  versus  filament  spumigena).  optical  density  cell  widths The at  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 (•) and 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 between t h a t of s t r a i n s 12 a n d 9* b.  Those i s o l a t e s with filament widths from 5-7 m ( i . e N. h a r v e y a n a ) . The relationship 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 ) . All other 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  Light  was  cool-white  s u p p l i e d by  florescent  General  tubes.  The  Electric  20  spectrum of  a n d 40 light  watt  from  400-720 nm was m e a s u r e d b y u s i n g a n a p p a r a t u s s i m i l a r t o one  d e s c r i b e d by B u r r  ity  of  light  the was  varying light a  light set  the  is  at  tubes  shown as F i g .  four  number from  (1972).  and Duncan  levels  of  the  light  of  15.  and the  vessel.  1.67-873 f o o t - c a n d l e m e t e r  Gossen  The  spgectral  quantity  illumination  tubes  culture  The  (Table  distance  Light  was  the qual-  of  the  III)  by  of  the  measured  and a radiometer  with  (YSI  2 model  calibrated  60)  otherwise  9-H  range The poor  use  of  two  or  adding  an  isolates  such  that  buffers  (in  was  unit  1.0  the  unless  increments)  investigated. tried  overlapping  in  was range  method of  adjusting  used.  photoperiod  (light:dark).  no b u f f e r  pH e x p e r i m e n t s  1 M H C l was  The  hours  pH 6-11  and S k o o g (1952) o f  1 M NaOH o r  The  of  increments)  more  For  hours:8  16  range  unit  0.2  results.  a Metrohm  four  the  pH u s e d w a s  Fitzgerald with  in  (in  of  ergs/cm / s e c .  s p e c i f i e d was  Growth and  in  the  The  The  wide  adequate. gave  very  Gerloff,  pH e v e r y  hours  12  pH was m e a s u r e d  with  E280A pH m e t e r . salinity  was  varied  in  a range  of  appropriate  amount  of  NaCl to  the  were  isolates  grown (Nos  on t h e 7,  9,  above  10,  11)  range were  of  1-100  °/oo  medium.  NaCl  and i n  grown w i t h  by All addition,  Na£S0^,  M^SO^  42  Figure  Table  15.  III  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 B a r r and Duncan .19 72)  Light intensities  used i n c u l t u r e /  Light  Level 1  Light  Level  Light  Level 4  (LL1)  L i g h t L e v e l 2 (LL2)  3 (LL3) (LL4)  experiments.  2 ,  foot-candles  ergs/cm /sec  550-600 200  1.8xl0  4  60-75  0.9xl0  4  25  4xl0  l.lxlO  4  3  lux  6000-6400 2200 650-8( 240  43  and Na^CO^ anions  as t h e  or  dominant  cations  had  salt  effects  to  on  discover  rates  of  if  different  growth  or  salinity  tolerance. The  effect  of  nitrogen  using  NO,,  NH~^ a n d u r e a .  salts  (i.e.  osmolality)  riate  amounts  3  BG-11 w i t h o u t A  level 100  algal  cells  ence  growth  nitrogen  amount The  of  purity  of  from  below)  patterns  level  and as is  isolates  a number  of  phase  contrast  microscopy.  material was  and under  measured by  1970)  at  Nodularia  White was  laboratory  (May  present  as  approp-  grown  growth  rates  with  contrast was  cells/  microscopy).  little  with  differ-  data the  No.  6 was  media  were  and Inks  a major  algal  above.  checked  (Table  IV)  studied with  conditions. method  (except same  mentioned  growth  Nitrogen (Stewart  Lake  of  low  5 bacterial  1 and No.  on  inoculation.  consequence a l l  reduction 1972)  was  cultures  there  investigations  acetylene Lake  phase  dissolved  adding  whether  than  investigated  contamination  bacteriological  fixation  by  b a s e d on c u l t u r e s  bacterial  axenic  a  of  before  identical (less  was  material  days  showed t h a t  using  Nitrogen  20-30  measured with  fixation)  low  constant  c o n s i d e r a t i o n was  as  (details  for  kept  contamination  Results in  for  differed  bacterial  concentration  Inoculation  nitrogen  cultures  was  NaCl.  preliminary  axenic  The  4  of  s o u r c e s on growth  (June  component.  and  field  fixation et  1972)  al. when  Axenic  44  Table  1.  2.  IV.  M i c r o b i o l o g i c a l media of cultures.  Nutrient Broth beef extract SST  Peptone  -  glucose 4-  - 1.0 water  g; tryptone - 100 m l  -  5.0  g;  -  1.0  g;  yeast  1.0  g;  Potato-dextrose  peptone  -  1.0  dextrose  -  20  g;  water  water  -  1.0  1  extract  -  -  1.0  1  agar g;  agar  -  20  g;  Thioglycolate 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 5 g; N a C l - 2.5 g ; 1 - c y s t i n e - 0.75 g; l a t e - 0.3 g ; a g a r - 0.75 gi methylene .002 g ; w a t e r 1.0 1  6.  purity  Glucose -  p o t a t o - 200 g; w a t e r - 1.0 1 5.  peptone  checking  Media  glucose 0.5 g j 3.  g;  3.0  for  Sodium c a s e i n a t e  dextrose thioglycoblue -  agar  s o d i u m c a s e i n a t e - 3.0 g ; a g a r - 12 g; w a t e r - 1.0  peptonized 1  milk  -  7*0  g;  45  Cultures the  were  gas  used f o r  p h a s e was  600C, W i l k i n s six  with  a  foot  apak  R and w i t h gas.  for  components.  of:  acetic  acid,  algae  The  apparatus  controls  sisted  algae  fixed  stainless  hydrogen flame  carrier  and t h r e e  (O.D.)  Three for  each run  algae  TCA t h e n  at  replicates were  incubated then  2% w / v ) ; with  set  of  each  used.  Por-  helium  the  1 ml  as of  45  C  sample  controls  TCA  but  Ltd.)  column of  The  with  incubated  acetylene  Co.  a temperature  killed  not  with  of  (model  now V a r i a n  steel  ionization  was  Analysis  chromatograph  and Research Inc.  in  0.25  experiments.  an A e r o g r a p h gas  Instrument  with  all  laboratory  con-  (trichloro-  fixed  with TCA;  added and sample  incubated.  Results: Comparison difference bacterial cultures The  rates  between  axenic  cultures  contamination. grew  than  at  a  lates  the  but  most  cultures  grew  35  arently  less  than  occurred  (although  5 C, quite  axenic was  tolerance The the  slowly)  at  in of  to  was  with  the  growth  the  for  at  all  limit  25  growth  the  in  V).  among  isolates  temperature 5 C for  level  (Fig.16).  (Table  either  of  low  morphology  cultures  O.D.)  l i t t l e  contaminated  some v a r i a b i l i t y  lower  lov/est  there  cultures  rate  closer  (highest  C (Fig.17).  and  Nodularia  were  There  best  showed t h a t  higher  morphology of  maximum t e m p e r a t u r e C and  The  slightly  Temperature.  1.  30  growth  contaminated  nature  The  of  iso-  C or  was was  30  C.  between app-  studied. 5 isolates  Growth which  46  F i g u r e 16.  Comparison of growth of a x e n i c c u l t u r e s c u l t u r e s w i t h b a c t e r i a l contamination.  and  data a t 25 C, 1.6 °/oo and pH 9 f o r i s o l a t e 1 ( p o i n t s are mean of s i x v a l u e s . V e r t i c a l bar i n d i c a t e s range) o A  contaminated axenic  47  .10  s c  .08  o o O  e rt X> l< o 10  .06  rt o .04  .02  (ft  4 Days  Table  V.  Comparative morphology of N o d u l a r i a i n axenic 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 and n a t u r e .  axenic  contaminated  nature  rare  very  straight  straight  regular  regular  1.  autolysis  not  2.  filament shape  twisted  3.  spacing of heterocysts.  uncommon  rare  irregular  48  F i g u r e 17.  Effect  of temperature on i s o l a t e s of N o d u l a r i a .  R e s u l t s shown are f o r pH 9, L L 1 w i t h b a s a l medium ( s a l i n i t y about 1.6 /oo) a f t e 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)  d.  i s o l a t e 13 (o)  14 (X)  11 (•) 15 (•)  12 ( A ) 16 ( A )  O.D.  (absorbance 660 nm)  O.D.  (absorbance 660  nm)  50  were t e s t e d . 2. level,  Light.  Most  isolates  LL1 (Fig.18).  Two i s o l a t e s  LL2  and i s o l a t e  ing  on t h e temperature.  light to  extent  isolates  any other 3.  N o . 12 g r e w  at  than  pH.  at the higher  (Nos. 2,  at  either  it  lower  slightly  best  at  LL1 o r LL2 depend-  temperatures  d i d at very  9) g r e w  light  (35 C) t h e  to inhibit  light at  growth  levels.  Thus  LL4 but not at  a l l  intensities.  The i s o l a t e s a-d).  showed o p t i m a l  growth  at  or  near  F o u r t een  isolates  h a d pH g r o w t h  between  1 0 . 0 a n d 10.4, w h e r e a s  isolates  N o . 2 a n d N o . 8 grew  best  at  p H 9*4 a n d  4.  Salinity.  pH  10 (Fig.19  best  the temperature  35 C g r e w  light  best  At high  combined with  a greater  most at  levels  grew  9.6 r e s p e c t i v e l y A wide  among t h e i s o l a t e s .  range  optima  (Fig.19 e , f ) .  of v a r i a b i l i t y  Some i s o l a t e s  ( N o s . 6,  was s e e n  8, 1 0 , 1 2 )  grew  o , in  a maximum  lates  salinity  showed an u p p e r  salinity  of i n i t i a l  Cultures  initiated  had if  of  then  were  isolation from  However,  tolerance affected  of  i n t h e range with  60  ° / o o .  ° / o o .  The  (1.6 ° / o o )  o f 40-50 ° / o o .  60  iso-  limit.  BG-11  i n o c u l u m grown  was i n c r e a s e d t o  most  t h e upper  inoculum i n normal  initiated  the tolerance  / o o NaCl.  salinity  a maximum t o l e r a n c e cultures  30  at  However,  40 ° / o o  Growth  of  o / inoculum er  at  60 / o o a n d a t t e m p t s  salinities  were  unsuccessful.  at  growing  cultures  The s a l i n i t y  at  at  high-  which  51  Figure  18.  Effect  of  light  intensity  Growth c u r v e s f o r c , d , b a s a l m e d i u m ( a b o u t 1.6  a.  growth  of  isolate  30 C (X), 35 C (o). b.  growth  1 at  25 C (A),  of  isolate  30 C (X),  e, f at °/oo)  25  different  20 C ( • ) ,  2 at  25 C (A),  on growth  different  20 C ( • ) ,  of C,  Nodularia pH 9  temperatures  temperatures  15 C ( o ) ,  c.  isolates  1  (o),  2  (X),  3  (• h  4  (A)  d.  isolates  5  (o),  6  (X),  7  (•),  8  (A)  e.  isolates  9  (o),  10  f.  isolates  13  (o),  14  (X),  11 15  -  15 C ( o ) ,  35 C (•)  (X),  and  (•), (•),  12 16  (A) (A)  -  52  X  O  A D O  D O A  D O A  • O  A  x  O  •  6 8 A  O A  O A  • x  LL3  LL2  LL1  60  200  9A  • O Ax.  LL4  LL3  LL2  L L 1  25  60  200  600  light level ft.-c.  L L 4  25  •  600  53  Figure  19. a.  Effect growth  3 (•), b.  growth  7 (•), c.  growth  of at  growth  at  f.  6-11  pH  6-11  at  pH 6-11  at  for  isolates  1  (o),  2  (X),  for  isolates  5  (o),  6  (X),  for  isolates  9  (o),  10  for  isolates  13  (X),  pH 6-11  (o),  14  (X),  16 (A) 9.2-10.0  at  pH  growth  at  pH o f  (O)  Nodularia  12 (A)  growth  13  of  8 (A)  15 ( • ) ,  e.  pH  growth  4 (A)  11 . ( • ) , d.  pH o n t h e  for  10.2-10.8  isolates for  2 (X),  isolates  8 (A)  6 (X),  54  55  optimum growth  for  a l l isolates Use  of  results. inities for  salts  the  other  Isolate  No.  all  than  isolates  for  major  5 °/oo  NaCl.  NaCl gave  and  °/oo  20  nor  growth  showed s i m i l a r growth  or  S0^  were  were  NaCl  the  same  higher  of  at  the  major  sal-  tolerance  evident  as the  C0^  at  levels  rates  No d i f f e r e n c e s  cation  essentially  showed b e t t e r  9  used and h i g h e s t  as  between  (Fig.20).  salts  but  inities as  o c c u r e d was  same with  salMg  anion  (Fig.21). Nitrogen.  5. than  with  with  22  NH^  as  +  c,d).  nitrogen  the  at  of  any  isolates  source of  Growth  than  centrations  All  was  level  much b e t t e r of  NH^ .  N H ^ C l s h o w e d some g r o w t h , NH^"*~ u s e d i n i t i a l l y  using  as  the  on N H ^  field  to  very  be  used cover  to  than  two  axenic  effect  of  fixation.  the  was  without at  added  lower  perhaps the  similar  NO  Fig.22a,b  con-  high  inhibitory.  source gave  as  a measure  (approx.  (less  investigate  the  nitrogen  reduction,  the  media  with  con-  Growth  results  to  e,f).  22  conditions low  laboratory,  nitrogen  (Fig.  +  Acetylene under  in  (compare  Cultures  +  of  growth  much b e t t e r  nitrogen  centration urea  grew  50,000  1 nmole/l isolates,  rates  nitrogen, Nitrogen  of  of  nitrogen  cells/liter)  pond w a t e r ) . No.  nitrogen  1 and No. fixation  pH a n d s a l i n i t y in  the  growth  fixation, was In  the  6,  were  and to  on r a t e s  media  found  dis-  of  inhibited  56  F i g u r e 20.  E f f e c t of s a l i n i t y on growth of N o d u l a r i a . Growth a t s a l i n i t i e s o f 1.6 - 60 °/oo a t LL2, pH 9> 25 C, inoculum grown at 1.6 BG-11)  o  /oo  (normal  a.  growth f o r i s o l a t e s 1 ( o ) ,  2 (x),  3 (•),  4  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 ) ,  d.  growth f o r i s o l a t e s 13 ( o ) ,  14 ( X ) ,  11 ( • ) , 15 ( • ) ,  (A)  12 16  (A) (A)  57  58  Figure  21.  Effect NaCl  of  salinities  on growth  Growth  at  of  using  salts  salinities  1.6  -  60  °/oo  1.6 ° / o o  a.  isolates  10  MgSO  b.  isolates  9  c.  isolates  7  d.  isolates  11 Na  N  a  SO S 2  2  z  (A)  °4 ^  Na S0 SO  than  Nodularia.  pH 9, i n o c u l u m g r o w n a t Na  other  4  4  M  g  s  (•) (°)  0 4  LL2,  at  25  BG-11. Na N a  C 2  CO  (o)  °3  (A)  MgSO^ (•)  Na^O^  (o)  (A)  MgSO  Na  (o)  4  (•)  CO  -"3  C,  59  60  Figure  22.  Effect of nitrogen Nodularia.  s o u r c e on t h e  A l l d a t a a t 25 C, LL2, pH 9 a n d (salinity a b o u t 1.6 °/oo).  growth  basal  medium  a.  growth 3 (•),  o n NO  by  isolates  1  (o),  2  (X),  b.  growth  o n N0^  by  isolates  5  (o),  6  (X),  c.  growth  o n NH  by  isolates  1  (o),  2  (X),  by  isolates  5  (o),  6  (X),  7 (•), 3  d.  (•),  growth  7 (•),  4 (A) 8 (A)  4  (A7  o n NH.  8 (A)  e.  growth  on urea  3 (•),  4 (A)  f.  growth  on u r e a  7 (•),  8 (A)  of  by  isolates  1  (o),  2  (X),  by  isolates  5  (°),  6  (X),  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 w h i c h rates increased.  Highest r a t e s of acetylene  reduction  o c c u r e d a t c o n d i t i o n s s i m i l a r t o t h o s e f o r maximum growth (pH 1 0 , s a l i n i t y 5-10 ° / o o , t e m p e r a t u r e 30 C) ( F i g . 2 3 ) .  This  perhaps i s a r e f l e c t i o n o f t h e b i o c h e m i c a l l i n k between  photo-  synthesis  and n i t r o g e n f i x a t i o n ( p h o t o s y n t h e t i c  phosphoryl-  a t i o n s u p p l y i n g t h e energy r e q u i r e m e n t s o f t h e p r o c e s s , art  1970).  Stew-  63  Figure  23.  E f f e c t o f s a l i n i t y , pH and t e m p e r a t u r e 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  on  o f s a l i n i t y on i s o l a t e 1 , L L 2 , 2 5 C, b a s a l m e d i a ( a b o u t 1 . 6 °/oo)  a.  Effect pH 1 0 ,  b.  E f f e c t o f pH o n i s o l a t e 1 , L L 2 , 2 5 C, b a s a l m e d i a ( a b o u t 1 . 6 °/oo)  c.  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 1 , L L 2 , pH 1 0 , b a s a l m e d i a ( a b o u t 1 . 6 °/oo)  65  TAXONOMY  The  genus N o d u l a r i a  was  e s t a b l i s h e d by Mertens i n 1822  d i s t r i b u t i n g specimens of the a l g a as Decas XV #4  of the  s i c c a t a e Algae A q u a t i c a e of G.H.B. Juergens (1822). c o l l e c t i o n was  made i n 1821  Germany (53.4°N, 7.0°E). m a t e r i a l of N. 3925-64, NBV The  The  type  S e v e r a l specimens of the (UC  436361, NBV  3925-67, NBV  isotype  3925-51,  NBV  3925-68, F 951302)""".  l a s t specimen (F 951302) i s i n very good c o n d i t i o n .  g e n e r i c name was  "conserved" by a d e c i s i o n of the  a l Committee on B o t a n i c a l Nomenclature (I969), used the name t o d e s c r i b e No  ex-  on the i s l a n d of Norderney,  spumigena e x i s t  3925-66, NBV  published  by  The  Internation-  as Link i n  1809  a r e d a l g a , Lemanea ( S t a f l e u 1972).  d e s c r i p t i o n of N o d u l a r i a  e x i s t e d u n t i l 1888  Bornet and F l a h a u l t r e v i s e d the h e t e r o c y s t o u s bluegreen T h i s work by Bornet and F l a h a u l t  (1886-1888) has  when algae.  been d e s i g -  nated by the I n t e r n a t i o n a l Committee on B o t a n i c a l Nomenclature as the taxonomic s t a r t i n g p o i n t of t h i s group of b l u e g r e e n  * L e t t e r i n i t i a l ( s ) are standard a b b r e v i a t i o n s of h e r b a r i a (UC - U n i v e r s i t y of C a l i f o r n i a B e r k e l e y ; FH - Farlow Herbarium, Harvard U n i v e r s i t y ; F - F i e l d Museum of N a t u r a l H i s t o r y , C h i c ago; NBV - R i j k s h e r b a r i u m , Leiden Netherlands; NY - New York B o t a n i c a l Garden; PC - Cryptogamic Herbarium of the P a r i s Museum; BM - B r i t i s h Museum, London; UBC - U n i v e r s i t y of B r i t i s h Columbia, Vancouver). The number f o l l o w i n g the herbarium desi g n a t i o n i s the a c c e s s i o n number of the i n s t i t u t i o n (or l o a n number i n the case of NBV). I f no a c c e s s i o n number has been designated, a number from 1000-1400 f o l l o w i n g the name "Nordin" r e f e r s t o an i d e n t i f i c a t i o n number of the annotation l a b e l a t t a c h e d t o the herbarium sheet a f t e r examination.  66  algae job  (Briquet  in  1935).  reducing the taxonomic  time.  Between  Nodularia closely  were  1822  described  and F l a h a u l t confusion  a n d 1888  several  were formed s i n g l y formed i n  to  Spermosira  its  They  also  1834,  in  whereas  groups.  characteristic and  because in  Bornet be of  Kutzing  Bornet  (translated from  and  separated  the  that  Lyngbya,  were  consider  the genus  describe  this  genus Nodularia.  JL. a n n u l a t a  and F l a h a u l t  the  akinetes  included the in  of  this  akinetes  d i d not  species  of  existed at  material  Spermosira  signifance  admirable  (1843) e s t a b l i s h e d  Juergen's  accompanying  the genus.  that  and F l a h a u l t  included a species  follows  in  d i d an  additional species  and K i i t z i n g  r e l a t e d genus Spermosira.  genus from N o d u l a r i a  as  Bornet  Suhr  the  genus  Latin):  "Filaments free within a sheath. S t e r i l e trichomes exceptionally uniform. Sheath h y a l i n e , c l o s e , commonly t h i n , m u c i l a g i n o u s , sometimes l a s t i n g o n l y a short time. Cells short, flattened, disc-shaped, h e t e r o c y s t s compressed. Akinetes spherical, subs p h e r i c a l or disc-shaped, formed i n the i n t e r v a l between h e t e r o c y s t s w i t h a smooth e p i s p o r e . " After  examining  m a t e r i a l from European  u c e d 16 t a x a t o  3 species  ieties).  also  They  which Bornet Their  1.  described  and F l a h a u l t  validity is  N.  names  Flahault,  one  later in  (Mertens i n  Ann.  Sci.  species  they  having  new s p e c i e s .  recognized  discussed  spumigena  (one  herbaria,  Nat.  are  The  discussed  this  3  redvarspecies  below.  section.  Juergens])  Bornet  Bot.  7.  ser.  and  7 : 243.  1888.  67  The  s p e c i e s has the c h a r a c t e r i s t i c s of the genus with  aments 8-18  ym wide.  Bornet  v a r i e t i e s of t h i s s p e c i e s . 8-18  and F l a h a u l t d e s c r i b e d t h r e e The  v a r . genuina had  ym wide and t h e a k i n e t e s subglobose.  had f i l a m e n t s 12-16  ym wide and a k i n e t e s  had f i l a m e n t s 12-18  elliptical.  /urn wide and  filaments  The v a r .  litorea  spherical-compressed.  The v a r . major (not maior as i n c o r r e c t l y c i t e d by 1932)  fil-  akinetes  Geitler,  flattened-  F i g . 24d.  2. N. armorica T h u r e t ,  i n Bornet  and Thuret,  A l g o l . 2, p.122, p l a t e 29, F i g . 12,13, T h i s s p e c i e s was  first  near L e C r o i s i c , France.  Notes  1880.  d e s c r i b e d from m a t e r i a l c o l l e c t e d  I t was  s e p a r a t e d from N.  spumigena  by the unique s t r u c t u r e of i t s a k i n e t e s which i n l i v i n g p l a n t s had biconcave  end c e l l w a l l s , and the ends of v e g e t a t i v e  f i t t i n g i n t o the adjacent ("pileatae").  The  f i l a m e n t s were 10-12  compressed and d i s c - s h a p e d wide as l o n g .  a k i n e t e l i k e a head i n t o a  and  3-4  cells  times  as  F i g . 24c.  3. N. harveyana (Thwaites) Nost., p.37S, The  cap  ym wide, the  before d i v i s i o n ;  cells  a l g a was  first  Thuret,  Essai class.  1875.  d e s c r i b e d by Thwaites i n W.H.  P h y c o l o g i a B r i t a n n i a (1848) as Spermosira harveyana.  Harvey's  68  F i g u r e 24.  - a.  The 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 and s i n c e t h e r e v i s i o n o f B o r n e t 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 ) . N. h a r v e y a n 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 Harvey ( 1 8 4 8 ) , see a l s o F i g . 32.x  b.  N. s p h a e r o c a r p a  c.  N. a r m o r i c a  d.  250  f r o m Fremy ( 1 9 2 9 ) . x 500  d r a w i n g of t y p e m a t e r i a l from B o r n e t and T h u r e t ( 1 8 8 0 ) . x 65O N. spumigena drawings o f Merten's t y p e m a t e r i a l f r o m B o r n e t and T h u r e t ( 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. p a l u d o s a d r a w i n g of t y p e m a t e r i a l f r o m W o l l e ( 1 8 8 7 ) . x approx. 250  g.  N. t u r i c e n s i s  h.  N. h a w a i i e n s i s  i.  N. t e n u i s d r a w i n g f r o m 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 ( 1 9 2 9 ) on r i g h t , x 1400  d r a w i n g of t y p e m a t e r i a l from H a n s g i r g ( 1 8 9 2 ) . x 400 d r a w i n g from T i l d e n ( 1 9 1 0 ) . x approx. 400  c o n t i n u e d p.  75  69  70  Thuret ( 1 8 7 5 ) t r a n s f e r r e d t h e s p e c i e s t o N o d u l a r i a .  Bornet  and F l a h a u l t d e s c r i b e d t h e f i l a m e n t s as 4 - 6 ym wide; a k i n e t e s subglobose,  6 - 8 ym wide; v e g e t a t i v e c e l l s b e f o r e d i v i s i o n not Fig. 24a.  much l o n g e r than wide.  4« N. sphaerocarpa Nat. B o t . s e r . 7.  Bornet and F l a h a u l t , Ann. S c i . 7 : 245. 1888.  T h i s s p e c i e s was d e s c r i b e d as new by Bornet and F l a h a u l t from s e v e r a l c o l l e c t i o n s : Meneghini  i n I t a l y on s o i l ; R o u s s e l on t r e e sap i n F r a n c e .  The R o u s s e l specimen  was examined as a l o a n from t h e P a r i s  Museum (PC Nordin 1 3 1 5 ) . the authors.  No type specimen  was d e s i g n a t e d by  N. sphaerocarpa was d e s c r i b e d as b e i n g d i f f e r -  ent from N. harveyana (6-7  Bory i n Belgium; Bornet i n France;  by f i l a m e n t s s l i g h t l y wider i n width  ym), s l i g h t l y wider width o f t h e a k i n e t e s ( 7 - 1 0 ym) and  t h e shape o f t h e a k i n e t e s ( s p h e r i c a l - c o m p r e s s e d ) .  F i g . 24b•  The f o l l o w i n g s p e c i e s were d e s c r i b e d a f t e r 1888 or were not t r e a t e d by Bornet and F l a h a u l t .  T h e i r present s t a t u s i s  d i s c u s s e d and t h e v a l i d i t y of each taxon i s c o n s i d e r e d .  5.  Spermosira a t l a n t i c a 11  D i c k i e , J . L i n n . Soc. B o t .  : 458, Fig. 4, 1871.  The m a t e r i a l on which t h e name was based was c o l l e c t e d from h u r r i c a n e d e b r i s i n t h e A t l a n t i c , 200 m i l e s o f f t h e  71  A f r i c a n coast. described.  (1907) c i t e d t h i s as b e i n g incompletely-  Forti  The type c o l l e c t i o n was t h e o n l y c o l l e c t i o n made,  no type m a t e r i a l c o u l d be l o c a t e d and t h e inadequate d e s c r i p t i o n and u n c o n v i n c i n g f i g u r e suggest t h a t t h e name s h o u l d be rejected.  F i g . 24«s.  6.  N. paludosa  Wolle, Fresh-water Algae o f t h e U.S.  p. 2 9 1 , p l a t e 198, F i g . 3 , 4 . 1 8 8 7 . The b a s i s f o r d e s c r i b i n g t h i s as a new s p e c i e s was t h e mistaken i m p r e s s i o n t h a t european  c o l l e c t i o n s o f t h e genus were  a l l made i n b r a c k i s h waters and always o c c u r r e d i n c o n s i d e r a b l e masses.  W o l l e ' s m a t e r i a l was c o l l e c t e d from Colorado and  Pennsylvania. eyana  The c e l l dimensions a r e i d e n t i c a l t o N. h a r v -  and from t h e d e s c r i p t i o n and i l l u s t r a t i o n i t i s p o s s i b l y  a N. harveyana although no a k i n e t e s were seen. r e p o r t e d t h a t no specimens herbarium.  Drouet  (l93o)  of t h i s s p e c i e s e x i s t e d i n W o l l e ' s  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 (I896), A c k l e y (1931), Coyle (1930) and  made by Clements M c l n t e e r (1939)•  Forti  d e s c r i b e d and G e i t l e r  (1907) c o n s i d e r e d i t i n c o m p l e t e l y  (1932) d i d not l i s t  s h o u l d be r e j e c t e d because l a c k o f type m a t e r i a l . .  7.  it.  The s p e c i e s  o f t h e inadequate d e s c r i p t i o n and  Fig.24f.  N. mainensis F.L. Harvey, B u l l . T o r r e y Bot. Club 16  :  188, 1889.  72  This Maine.  alga It  and c e l l s ure  was c o l l e c t e d f r o m  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  2-6 ym l o n g .  was i n c l u d e d w i t h  type ies  i s unknown  No a k i n e t e s  the description.  reported.  on t h e b a s i s o f b e i n g  Bohm.  incompletely  by  Cramer  to  t h e genus N o d u l a r i a  the  taxon  erial  Forti  with  show i t  despite  t o be i d e n t i c a l  Algae  that  of the  the  spec-  rejected  Prod.  Alg.F l .  in  synonomize  of herbarium  mat-  1236; NBV 3925-1)  N. harveyana.  Tilden,  and F l a -  A l g e n S a c h s e n s #994:  1262; PC N o r d i n  with  the species  N. harveyana  (1932)  Examination  turicensis  Bornet  synonomy w i t h  (Rabenhorst's  9. N . h a w a i i e n s i s  ican  the fact  (1907) a n d G e i t l e r  1084; NY N o r d i n  Described  to  N. harveyana.  Ann.  Hansgirg,  d e s c r i b e d as Spermosira  c o l l e c t e d by Cramer  FH N o r d i n  collection,  fig-  F i g 27, 1892.  r e d u c e d S_. t u r i c e n s i s (p.244).  1888  a n d no  The l o c a t i o n  H a n s g i r g i n 1892 t r a n s f e r r e d  (i860).  \m w i d e  33-38  reported  Orono,  described.  (Cramer)  2 : 74  was o r i g i n a l l y  near  The s p e c i e s s h o u l d be  8. N . t u r i c e n s i s  hault  were  and s i n c e the o r i g i n a l  has not been  This  Pushaw S t r e a m  F i g . 24g.  Hawaiian  Almanac and  (I902.) p . 1 1 2 , 1 9 0 1 .  from  a s #484.  Hawaii  and d i s t r i b u t e d  Examination  indicates  in Tilden's that  Amer-  isotypes  73  (UC  740394;  only  1250;  PC N o r d i n  1339;  other  two c o l l e c t i o n s ,  by Tilden  in Tahiti  ific  Algae  #11; U C 233460; NBV 3925-47; NY N o r d i n 1257) a n d  that  by Crossland from T a h i t i  Rather  they  NY N o r d i n  a r e most  and Grunow. it  Kahn  i s apparently  ipines,  be  easily  having  erocysts, ution.  and t h e  (South  Pac-  are not Nodularia. (? s o l u t u m )  (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 marine  habitats  Bornet Hawaii, (Phill-  1962; F l o r i d a , T a y l o r 1928; D a w s o n I966).  for i t  variable  lack  Hormothamnion  i s superficially  mistaken  very  696203),  common i n t r o p i c a l  Velasquez  Hormothamnion  likely  (UC  UBC 2971)  similar  (May 1 9 4 6 ,  trichome  of akinetes  size,  to Nodularia  1951)  and can  but differs  irregular  anda d i s t i n c t l y  in  spacing of  tropical  het-  distrib-  F i g . 24h.  10.  N. tenuis  G . S . West,  J . Linn.  S o c . B o t . 38  :  171, 1907. The 1347)  type  specimen from  andt h e m a t e r i a l  It  i s probably  of  Geitler  was e x a m i n e d  of the attributes  (BM N o r d i n  of t h e genus.  confirms the suggestion  F i g . 24i.  N. quadrata Exped.  The  h a s none  an Anabaena s p . which  (1932).  11.  Tanganyika  Fritsch,  p.45, p l a t e  Freshw.  A l g . Nat. Antarct.  2, F i g . 109-115, 1912.  s p e c i e s was d e s c r i b e d f r o m  material  collected  in the  74  Antarctic.  Fritsch  described it  is  smaller  (3-4 ym)  and because  some o f  the  trichome  Nordin  (BM  Fritsch  was  acteristic The  1934)  was  not  seen.  of  the  heterocyst  shape  square be  is  variable show a  (West  freshwater  and West  algae,  its  variability  of  and  lack  akinetes,  sp.  rather  of  the  than  12.  the  a Nodularia.  N.  spumigena  Nat.  one  important  is  unclear  c o n s i d e r e d N.  work  is  been  Wille  1959)  since  with  quadrata it  to  1924)  and more  Antarctic  s h o u l d be  c o n s i d e r e d an  an  des-  of  Because of  heterocyst  to  limi-  surveys  irregular  the  spacing Anabaena  24j.  minor  Exped.,  be  recollected  questionable.  Fig.  char-  Iyengar  spumigena  considers  never  by  spaced heterocysts.  material. N.  material  observed  (1932)  var.  Antarct.  akinete  of  1911;  alga  type  The  figure  1942  heterocyst,  one  its  (4-6ym)  harveyana  square.  lacks  the  Fukishima status  are  species because N.  regularly  s p e c i e s has  I964;  (Holm-Hanson  his  of  heterocysts  in  Geitler  Since the  detailed  of  new  that  material  square  in  a  and the  that  the  s p e c i e s but  Anabaena.  ted  The  of  heterocysts.  a valid  pite  examined  (1944)  and D e s i k a c h a r y  than  heterocysts  genus,  significance  as  Fritsch,  p.44,  plate  Freshw. 2,  Fig.  Alg. 104  108, 1912. This in  the  variety  Antarctic.  was  also  The  described from  type  material  material  (BM 1815)  was  collected examined  -  75  Figure  24 j.  cont'd. N.  quadrata  drawing  by  material, k.  N.  spumigena v a r . of  1.  N.  harveyana  minor  the  var.  N.  epiphytica  (1927) . n.  N.  willei  o.  N.  fusca  p.  N.  skujae  drawing of (1927).  of  Guerro q.  r.  N.  N.  spumigena  t.  N.  N.  aerophila  of x  type  (1912)  approx  500  from  approx  material  Skuja  type material x approx 500  type  by  type  zujaris  drawing  Gonzalez-Guerro of  (1940).  x  Taylor  by  Gonzalez-  material  of  type  (1930).  material 1750  drawing  (1940).  x  drawn  (1929). x  type  aerophila  (1928).  500  approx  Brabez  Gardner  Gardner  approx  and R i c h  drawing  by  material  (1928). x  by  300  Fritsch  from  type  820  from  spumigena v a r .  x  drawing  vacuolata  spumigena v a r .  Fritsch  material,  var.  by s.  by  drawing of type material x approx 400 drawing  the  500  drawing  sphaerocarpa  drawing  (1912) o f  approx  type  (1926) . x m.  Fritsch x  of  x  material approx  from  type  approx  1000  400  Brabez  material 500  77  and  is  have  identical  to  N.  harveyana.  been  reported  but N.  Antarctic  by A p f e l  (F  (1959).  F i g . 24k.  13.  N.  harveyana  1299674;  harveyana  hault)  No o t h e r  var.  Elenkin,  of  collected  from  has been  1299702)  F  collections  Jard.  the  and by Fukishima  sphaerocarpa  Bull.  this  (Bornet  Imp.  and F l a -  Bot. Pierre  le  G r . 16:331, 1916. Elenkin  considered that  ecological  variant  al  is  studies  14.  of  presented  N.  1254),  in  collection  (UC  is  verrucosum).  ytica  it  (see G e i t l e r Geitler  an Anabaena  have  evidently  from  been  1255) of  often  only  as w e l l were  (NY  F i g . 241.  7:65,  as a  second  examined.  Nostoc  844 N .  Gardner's  sp.  Fig.  24m.  The  Young  muscorum; diagrams,  collections  from  Nordin  do n o t r e s e m b l e  No s u b s e q u e n t made.  this.  collected  material  401800)  1932, p .  seeing  sp.  material  stages  of Nostoc  support  experiment-  1927.  NY N o r d i n  juvenile  filaments  filaments  ered  401800a;  (UC  from  an  Mem. N . Y . B o t . G a r d .  The type  material  actually  hormogonial ure  Gardner,  Rico.  was m e r e l y  Evidence  (p.89) t o  12, F i g . 1 6 ,  Puerto  the isotype  material  below  s p e c i e s was d e s c r i b e d  freshwater  sphaerocarpa  harveyana.  epiphytica  plate This  N.  N.  the  mat-  p.855  N.  consid-  of N.  epiph-  taxon  78  15.  N.  willei  plate This  freshwater  in  only  collection  (NY  is  12, F i g .  s p e c i e s was  from  identical  to  16.  N.  also  Puerto  ugas  s p e c i e s was  Islands,  provisionally  (personal  ies  be  is  to  de T o n i willei Lyngbya  It  Gardner  N.  taxon  (Ulmus  and  it  skujae  Hist. This  both  in  was  Nat.  be  is  1940 not  placed  longer  in  Casa  a  de  the  and  at  Tortgenus  akinetes.  the  spec-  present  Drouet  1-3,  material  and Fig.  Bol.  J .  Cyanothrix  growth  a Nodularia.  from  the  Johannesbaptistia  may b e  cited  1928.  in in  Pap.  considers this  position  genus  28:435, F i g .  at  pools  synonomous w i t h it  the  material  Wash.,  1, F i g . 2 3 ,  Gonzalez-Guerro,  described  Populus)  the  collected  and the  heterocysts  taxonomic  1927) or  a n d De T o n i  7:65,  evidently  Inst.  marine  s p e c i e s was  1 9 3 8 a ) , i t may  Nevertheless  17.  sap  belong  Gard.  24n.  from  no  and  examined  Carnegie  lacked  Its  (Gardner  (Fremy  Fig.  communication)  may  (Drouet  1956).  it  Bot.  material  type  25:48, p l a t e  The  a Nodularia.  unclear.  1 2 5 6 ) was  collected  because  from  The  Taylor,  Florida.  Taylor  Rico.  Lab.  N„Y.  15, 1927.  spumigena.  N_. f u s c a  Mem.  described  Nordin  Tortugas The  Gardner,  form  of  Daily 24o.  Real.  Soc.  Espan.  1928. collected  Compo a n d M a d r i d ,  in  tree  Spain.  79  (1932)  Geitler type  material  and f i g u r e  lists  c o u l d be  the  gas  This  the  of  the  very  abnormal ler  is  collected  akinete unlike  from  the  Most  only  it  and  from  zujaris  Zujar match  preserved  The  those  figure  to  abnormal  collection  N.  and  spumigena. does  not  24q.  or  Spain.  of  N.  s h o w n may was  the  dim-  7-12  ym  have  exhibiting  Anabaena in  The  harveyana,  ym l o n g ;  an Anabaena  reported  Bol.  7-8, 1930.  (4-12  it  an  of  regularly  (1922)  probably  River,  material is  presence  collections  for  Fig.  probably be  Roy.  Gonzales-Guerro,  and v a r i a t i o n genus.  the  Fig.  30:224,  cells  size  poorly  considers the  var.  Espan.  vegetative  growth.  (1932)  material  was  Fig.24p.  Trans.  collected  vacuoles  a variety.  wide)  drawn  gas  of  Sjostedt  of  alga  No  description  Fritsch  Baltic  (I898),  establishment  Soc.  the  harveyana.  are  1285)  variable  the  been  1284,  from  is  however, is  material  spumigena  N.  because  attribute  N.  harveyana.  from  vacuolata  variety  reported  have  to  N.  1 8 : 8 8 , 1929.  (1943)  Real  ensions  var.  Lemmerman  19.  The  this  NY N o r d i n  particular  justify  identical  vacuoles.  Lindstedt  however  Afr.  However,  (i. e. O s t e n f e l d , with  is  South  created  vacuoles.  synonomy w i t h  spumigena  Soc.  gas  in  located,  species  N.  18.  Fritsch  it  sp. sp.  GeitThe  literature  type and  80  this  variety  Fig.  24>.  s h o u l d be  excluded from  the  N_. f e r t i i i s s i m a R a n d a w a ,  20.  genus  Proc.  Nodularia.  Ind.  Acad.  Sci.  B. 4:41, 1936. This ern a  s p e c i e s was  India.  nomen  No  This new  description  N.  aerophila  6lA:214,  s p e c i e s was  reported  ments  and  ative  characteristic  lack  justify  its  ical  cell  ection ure  was  are  times  given,  of  basis  size  not  and the  N.  soil.  22.  and  from  collection  from  thus  rejected  it  is  Northas  N.  but  harveyana Fig.  from  variety  Austria  and  soil  habitat,  The  lack  of  other N.  the  two  Zent-  type  described short  akinetes  to  which  is  material  it  which  has  a  been  negdo  is  reported  not  ident-  and o n l y  published description  as  f i l a -  characteristics  harveyana The  and  collfig-  many  24s.  spumigena v a r .  was  zum B o t .  its  aerophila  zum B o t . Z e n t r a b l a t t This  Beihefte  F i g . 8 a , 1941.  shape.  examined  of  of  akinetes.  separation  that  from  Brabez,  rablatt  s p e c i e s on t h e  in  was  a  nudum.  21.  a  published from  collected  in  Brabez,  Beihefte  6lA:215, 1941. the  same  sample  as  N.  aer-  81  ophila  a n d was  habitat; cribed ical  short  from  to  separated  N.  a  filaments; single  Fig.  23.  N.  advocated genus  implexa  i n c l u s i o n of  (as  compared to  as  being the or  genera  A u l o s i r a has  flattened).  wide  The  distinctly  was  longer  than  wide  is  ident-  completely  prior been  resembles  a Hormothamnion  Dixit).  Another  A.  N.  long  implexa  Nodularia sheath  and t h e  long  division). as  implexa  round or  heterocyst.  wider The  are  and In  not the  as only  Aulosira  long  var.  than  thinr  2-4x  as  s c h a u i n s l a n d i i Lemm. and A.  the  because  spumigena as  and  were  was  cells  distinct  as wide  cell  Les  difference  morphological feature  either the  of  considered either  sp.  distinct  from  to  to  and t h e  sheath  1.5-2x  Bourrelly  only  cells  width)  than  (except  are  the  the  (N.  long  separate  is  des-  genus A u l o s i r a w i t h i n  wider  as  always  implexa  is  as wide have  always  was  p.418, F i g . 5 - 6 , 1970.  genus N o d u l a r i a  harveyana  Nodularia  (aerial  description  separation  that  a thicker  N.  of  its  It  and F l a h a u l t )  the  that  their  long;  cells  etes  of  He m a i n t a i n e d  thinner  are  its  Aulosira  part  two  cells  and by  d'Eau Douce.III.  the  as  akinetes).  (Bornet  transferred  Nodularia.  (i.e.,  of  therefore  between  her  lack  characteristics  24t.  Algues Bourrelly  similar  filament  spumigena,  unjustified.  on  wide  which  crassa is  that  long  akinetes  of  akin-  and Aulosira  82  are or  invariably adjacent  ocysts but  of  to  A u l o s i r a has  define  itat  was the  soil.  marine  the  and  genus  arrangement  other  and  only  commonly genus  sira  any  part  24. The only  Nodularia  N.  is  reported  from  The  is  filament,  distinct found  in  occurs primarily  in  should not  in  a  the  or  soil or  two  temperate  and  areas.  include  marine  the  and o n l y  tropical  hab-  brackish of  temperate  char-  freshwater  patterns  primarily  This  primarily  subtropics  heter-  characteristic  a  found  from  rarely  subtropical,  For  these  genus  Aulo-  it.  f.  crassa  citation  of  this  to  and f i g u r e  from  also  typically  rarely  spumigena  reference  is  hand,  the  Aulosira  of  is  a major  distribution  Nodularia  original  description specimen  The  1959).  occasionally in  on t h e is  removed  heterocysts.  as  There  only  occasionally in  the  or  and  of  (1875)  Thuret  either  spaced along the  Nodularia  different.  tropics.  less  (Desikachary  genus N o d u l a r i a .  reasons  The  random  Aulosira,  a n d may b e  regularly  situations  environments, in  are  environments.  very  wide  heterocyst  used by  and freshwater  are  than  differentiation.  and b r a c k i s h on  the  Nodularia  acteristic to  longer  Siberia  it, (p. with  in  (Woronichin)  taxon  Starmach  518-519)  c o u l d not  (I966),  to  particularly  be  of  large  Elenkin. be  appears an N. cell  located. from  the  spumigena dimensions.  83  Validity preceding since be  1888  the  can be  are  taxa  were the  shown t o  a  distinct single  the  N.  merely  of  N.  are  N.  described  by  establish  a  or  The  validity  questions  distinct  harveyana  entity;  is  and N.  halves  of  Bornet  and F l a h a u l t  can  Bornet the  justified;  a  The  genus  defined  armorica  spumigena  two  the  and F l a h a u l t .  N.  taxa:  species  from  Nodularia  in  truncate  France  1061;  note  that  1924).  hidden this  Evidently  by  name  by  N.  sphae-  spumigena  continuum  of  a  The  of  North  UC  1087,  and t h e three  this  the  first  Three America  North  end of  in  is  was  (1.  only  type  and  (Freray  in  been coll-  were  Gardner  9808I6,  F  adja-  have  the  is  the  collections  France  American  an  seen  Osterhout  3925-30,  fifth  species  collections  100568; 3. NBV  (1888)  concave end of  armorica.  Coast  the  the  five  1873.  1302)  which  only  602,  of  characteristic  in  FH N o r d i n  Of  in  Thuret  Gardner  PC N o r d i n  akinetes  is  Pacific  100567; 2.  characteristic  cell  specific  on t h e  PBA  armorica:  distinguishing  material.  ected  Meslin  of  species or  They  the  1213,  excluded  is  species;  vegetative  given  ner  names  the  necessary to  distinct  akinete.  made  is  all  Nodularia  species?  the  living  it  considered are:  biconcave,  cent  be  d e s c r i b e d by B o r n e t  Validity state  that  s p e c i e s of  (1888),  varietal  rocarpa  and F l a h a u l t ' s  indicates  included within  that  UC  Bornet  summary  and F l a h a u l t of  of  436, Gard-  NY  Nordin  and  collections  one  84  (UC  100568)  The  other  from  two  did  would  be  to  spumigena.  N.  were  ure N.  placed  from shown  in  The  the  akinete  two  and M e s l i n ' s  structure.  a microscopic artifact specimens  identified  between  the  This  odd s i n c e the  is  living ing  akinetes  cells  growth  N.  (Bornet  show t h a t  acteristics  far  tative species With  cells on  this  variant  are  of  N.  a  large  N.  spumigena of  number  are  identical  25a,b).  variable  and  (Fig.  varietal of  is  names  herbarium  in  several  is  did  for  det-  some  fig-  the may  be  herbarium  show b i c o n c a v e  areas  25c,d,e).  (Fig.  supposedly occurs only  1888). the  than  Studies  stability  shape,  size  size  alone  is  of  considered in  of  vary-  charof  separation doubtful an  in  taxonomic  shape  vegof  a  value.  ecological  synonomy w i t h  spumigena:  specimens,  under  and w a l l  and  Thus  26).  N.  very  characteristic However  placed of  is  except  752514)  UC  akinete  armorica  (Setchell  poor.  characteristics  evidence,  differed  602)  (1924)  spumigena  heterocysts  akinete  Validity of  or  more  (436,  report  determine  characteristics  Gardner  identical  The  and F l a h a u l t to  and were  as  description.  characteristic  conditions  akinete  (1888)  761550;  (UC  condition.  "doubtfully"  Indeed t h i s  (Fig.  as  by  specimens  spumigena  poor  material  armorica  the  in  characteristic  and F l a h a u l t ' s  N.  was  collections  species  Fremy and  Wash,  preserved  since  Bornet in  Is.  show t h e  with  1903)  armorica  unique  not  expected  and Gardner ails  Whidbey  it.  Examination  observation  of  the  85  Figure  25.  a.  Possible explanation o f N. a r m o r i c a .  for  A simplified  representation  etes  in  view,  with  respect  side  to  with  slide  akinete  showing  filament  not  characteristics  a  chain being  of  akin-  horizontal  surface.  b.  If t h e f i l a m e n t were not h o r i z o n t a l , t h e n a t o p view 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 overl a p p i n g to give the appearance of " d i s c s " between the akinetes.  c. d.  a n d e show i s t i c o f N.  "disc" areas armorica.  between  akinetes  character-  87 Figure  26.  Comparison of ive  cells,  lates  under  Count of t i o n s of salinity  the  variability  heterocysts a variety  and of  of  size  akinetes  culture  of  of two  vegetatiso-  conditions.  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 pH ( 8 ^ 9 , 1 0 ) , t e m p e r a t u r e (20 C , 25 C ) a n d (1.6 / o o , 10 / o o , 20 / o o ) n = 360.  a.  i s o l a t e 7 (i) vegetative (iii) akinetes.  cells  (ii)  b.  i s o l a t e 3 (i) vegetative (iii) akinetes.  cells  (ii  heterocysts  heterocysts  88  I'l  200  i l l  C.V.=8.15#  s =„52 C.V.=9.60^ 2  o CM  O  100  U  4) E  .O  3 S3  4.5  5.0  5.5  6.0  5.5  s =.12 C.V.=6.80fo  11  2  6.0  6.5  7.0  s =.12 C.V.=6.18# 2  6  .  0  6.5  7.0  7-5  111  8.0  8  -5  9.0  9-5  s =,63 C.V.=8.69£ 2  200 L  0]  H H  0)  u CM 0  100  0)  .o  B 3 sz;  4.5  5.0  5.5  6.0  5.0  5.5  6.0  6.5  5-5  6.0  C e l l w i d t h ( ym)  6.5  7.0  7.5  8.0  8.5  9.0  89  variability  of  conditions  and  that  seems  there  isolates  a  number  large l i t t l e  ietal  names  ially  established  12-18  um)  population akinete  of  several  and  N.  akinete  or  on t h e  formation  maturation  and  etes,  heterocysts  of  harveyana  N.  and  collection  ences a  were  aration  and  sites  into  of  There  appears  to  carpa  collected  be more  harveyana.  Of  frequently  Of  of  were  24 N.  artific-  1 2 - 1 6 ym among  from of  the  and  if  collector)  27)  there  Habitat differ-  "freshwater",  is  with  N.  no  sep-  is  gradation.  with  32.3%,  akin-  specimens  specimens,  a  of  show t h e r e  N.  than  harveyana  sphaerocarpa  its  shape  habitat  soil  N.  of  valueless.  compared.  habitat  clonal  during  are  However,  in  a  and  stages  herbarium  merely  var-  subspherical,  size  herbarium  collections  brackish-marine,  respectively.  cells  rather  13.2% ats  The  smaller.  according to  from  ym,  as  (Fig.  of  of  were  shapes  akinetes  results  (identification were  of  The  difference  68  8-12  see  groups,  a  retention  compared to  cells  being  distinct  indicates  variable  variety  the  nature,  Developmental  sphaerocarpa  were  size  generally  very  filament. a  culture  dimensions  sphaerocarpa:  N.  for  among  are  and v e g e t a t i v e  and  in  is  in  varying  flattened-elliptical  significant.  difference  harveyana  shapes  encompass  N.  isolates  The  choice  categorizing  of  widely  justification  same  spherical-compressed Validity  of  spumigena. (the  under  sphaeroN. examined  54.4%, and  and  soil  collections  habit-  (species  90  Figure  27.  Comparison of N.  cell  sizes  of  N.  and  sphaerocarpa. o  N.sphaerocarpa  (identification arium  A  N.harveyana  vegetative  b.  heterocysts  c.  akinetes  cells  on  herb-  on  herb-  sheet)  (identification arium  a.  harveyana  sheet)  Cell  length  (y  m)  C e l l length  > o  > 0  On  m)  Cell  length  (y  m  )  o >  «  L a  (y  9  rj  > t>  o> o  O  o ogo cp  o o o  o o o  O  o  O O  92  identification figures  two  of  interesting the  three  grown  over  itive  correlation  The  larger  the  their  and N.  ify.  harveyana  treating  with  This  field  the of  was  Scattergrams  and v e g e t a t i v e  (Fig.29).  (1968)  that  is  However  the  to  a  N.  make  the in  classes.  the  respective  a  fact  the  same  akinete are  akinete  as  pos-  (Fig.  28).  formed,  is  N.  hard  such  harveyto  just-  (1916)  Elenkin's  that  the  shows no  between  size  support  when  show a  size  isolate  that  portion  of  N.  dimensions.  by  and  N.  spumigena  herbarium  of  in  the  material of  with  for two  and  are  heterocyst  into  two  view  of  Nodularia  the are  the  readily  distinct Bursa  s h o u l d be  maintenance often  material,  laboratory.  akinete,  separation  s p e c i e s of  place  distinct  of  contrasts  the  as  observation  material  support  that  and  is  harveyana,  distinction  show a  single  N.  studies  (1-40 °/oo),  third  seem t o  herbarium  This  as  sphaerocarpa  size  culture  akinetes  b a s e d on  harveyana  An o b v i o u s  same t i m e size  only  which  and growth  cell  groups  species  salinity  in  examined  from  from  salinities  largest  N.  collections  sidered.  of  observations  Validity taxa:  salinity  sphaerocarpa  of  collector)  identified  between  Thus,  These  proposal  range  size.  correlation. ana  correlation  isolates  a wide  higher  original  22.6%, 52.1% a n d 25.3%.  were  An  according to  of  collected separable  the at  contwo the  into  two  93  Figure  28.  Akinete isolates  variability as  related  of to  three  British  salinity.  a.  isolate  7  1.61o  10%= A  20%n  b.  isolate  5  I . 6 & 0  10%>A  20%D  c.  isolate  3  1.6^o  10&A  20%D  Columbia  Number of akinetes  Number of akinetes H O  Cn O  Cn  O  Cn  o  oo O  oo Cn  o  Cn  to O  O  o  Number of akinetes O  95 Figure  2 9 .  Comparison of N. harveyana.  cell  sizes  of  N.  spumigena  and  o  N. spumigena  (identification arium sheet)  on  herb-  A  N» harveyana  (identification arium sheet)  on  herb-  a.  vegetative c e l l  b.  heterocyst  c.  akinete  size  size  size  C e l l length Oj  J*.  Lr,  (v  O  Cell  )  length  (  m  0) Co  00  > >  O  o» o cb°o° o o O  O  0(9  oo o  QJJ  Um)  a  o  o §(> o@ o o  03  Q  97  One of  approach used to  growth  and B o l d  patterns  scopically  identical  characteristics, genera  in  The is  this  manner  Thus  genus  The  genus  is  morphologically genus  cysts)  distinct  being  very  itions.  Other  presence;  filament  shape, the  texture  the  generic  Nodularia  under  length;  the  genus  investigation  in  growth  species or  two  this  even  study  range  quite  vegetative  cell  shown i n  presented  Juergens}  harv-  Fig.  as  the  of  color;  here  is  other  an  of  spacing of  the  hetero-  growth  variable  An i l l u s t r a t i o n is  N.  characteristics  between  wide  were  from  regular  midway  the  species,  separable  long;  species description (Mertens  micro-  (Fig.30).  Certain  akinete  and c o l o r ) .  and  different  Baker  Mertens.  easily  than  characteristics  morphology of From the  and  wider  little  only  spumigena  genera.  of  I969,  that  IX.  and N.  formation  varied  strikingly  taxa,  Isolates  considered during  Table  similar  (cells  heterocysts;  unworkable.  should contain  Thuret  algal  and B o l d  proved impossible  material  Appendix,  (Thw.)  completely  (Kanfcz  and r e c o g n i z i n g d i f f e r e n t  herbarium  the  bluegreen  plates  exhibited  shown i n  eyana  the  on agar  proved  1970)  separate  cond-  (sheath akinete  of  variability  31  and  in  32.  emendation  of  follows:  Bornet  and F l a h a u l t  emend.  Filaments with c e l l s d i s t i n c t l y wider than l o n g , h e t e r o c y s t s spaced at r e g u l a r i n t e r v a l s a l o n g the 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 the heterocysts.  98  Fig.  30.  Cultural  variation  Comparison after  two  at  C pH  20  refer sp.  to  which  of  growth  weeks 8 and  isolate has  of  Nodularia. patterns  growth LLl.  several 1 %  "IUCC  mis-identified  agar  numbers  583" as  isolates  (W-V)  Identification  numbers.  been  of  o n BG-11  is  a  Calothrix  Nodularia.  99  100  The  morphological variation  of  N . spumigena.  from F r i t s c h 1951 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 , sheath and l o c a t i o n of f o r m a t i o n of akinetes. from Bornet ament.  and T h u r e t  1880  x650,  vegetative  from Bornet akinetes.  and Thuret  1880  x650,  filament  f i l -  with  from B o r n e t and T h u r e t 1880 x650, t h e i r drawing of M e r t e n ' s t y p e m a t e r i a l , shows a v e g e t a t i v e filament and 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 . from Bharadwaja 1935 xl370, filament with akinetes.  vegetative  from P l a y f a i r filament with  1914 x800, vegetative immature a k i n e t e s .  from Prescott filament with  1962 x900, akinetes.  vegetative  filament  filament  filament  and  and  and  t o  H  (fflwiiiiiiiiiiiim in  D  102  F i g u r e 32.  The m o r p h o l o g i c a l v a r i a t i o n of N. harveyana.  a.  from Bornet and Thuret 1880 ament .  x650, v e g e t a t i v e  fil-  b.  from Harvey I848 x 2 5 0 , f i l a m e n t w i t h a k i n e t e s (drawing of type m a t e r i a l ) .  c.  from Smith 1950  d.  from H o r t o b a g y i 19 59 xlOOO, v e g e t a t i v e f i l a m e n t and filament with akinetes.  e.  from Hardingl971  f.  from G e i t l e r 1932  g.  from M a b i l l e 1954 xlOOO, v e g e t a t i v e f i l a m e n t and filament with akinetes.  h.  from C a r t e r 1933  i.  from Skuja 1926 x 8 2 0 , f i l a m e n t s w i t h immature and mature a k i n e t e s .  x800, v e g e t a t i v e f i l a m e n t .  xlOOO, v e g e t a t i v e f i l a m e n t . x900, v e g e t a t i v e f i l a m e n t .  x 6 l 3 , vegetative filament.  103  104.  Vegetative length  2:1  vacuole.  gas  um  -  wider  long.  than  disc-shaped;  8-18  series  in  a  two's).  colorless ish,  inland  Temperate  or  usually  or  and s u b t r o p i c a l N.  spumigena  Bornet  in  a  or  thin  Habitat marine,  'hard'  to  single  with  sheath.  euplankton;  saline,  to  wide,  U«n l o n g ,  6-15  (occasionally  Filaments  tychoplankton  ym  8-16  subspherical  urn w i d e ,  transparent  width/ with  subspherical  long  Akinetes  usually  in  pm w i d e ,  sometimes  10:1;  Heterocysts  disc-shaped; 2-10  7.5-16.0  cells  ratio  brack-  freshwater. distribution.  [Mertens  and F l a h a u l t  in  Juergens]  emend.  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 ; wider 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 b e e q u a l ) ; width/ l e n g t h r a t i o 2.5:1 - 1:1. Heterocysts subspherical (wider than long) to s p h e r i c a l ; 4•0-7«5 um w i d e . Akinete subspherical ( 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 wide, 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 present. Habitat tychoplankton or benthic; marine, brackish or inland s a l i n e situations^ o c c a s i o n a l l y in freshwater; also found on s o i l o r i n t r e e s a p . Temperate and s u b tropical in distribution. m  N.  The in  synonomy Appendix,  of  other  Table  X.  taxa  harveyana  attributed  (Thw.)  to  Thuret  Nodularia  are  emend.  shown  105  DISCUSSION  Ecology  and P h y s i o l o g y :  Few  c o l l e c t o r s have  environment ually ence  no in  in  which  observation,  work  will  quantitative  Nodularia  laboratory nature  made  be  has  spp.  been  were  done  reports  collected  on t h e m .  considered with  literature  measurements  and growth  and Its  reference  of  to  studies  the  virtoccurr-  ecological in  the  lab-  oratory. Temperature: at  temperatures  a wide  range  of  of  1911;  Brannon  Nodularia  to  reports 12.9  C.  limit  N.  only  (20-30  springs  springs  (Welch  instance  measured f o r  1049304,  in  more C).  I964)  but  any  which  the  maximum t e m p e r a t u r e  in  Baltic  of  large  harveyana  bloom  temperature collection  30-35  of  were a  at  or  given.  "hot"  The  is  26  C (Groesbeck  and t h i s  is  well  C from  laboratory  studies  from  warm  spring  below  at high  reported  1888)  not  (1922)  occur  been  1909;  al.1966;  quantities  numbers  and F l a h a u l t  give  does  Sjostedt  has  study  West  Woodson et  the  California),  1878;  Temperature  no t e m p e r a t u r e s  Nodularia  Mono C o . ,  1955;  this  literature  C (Francis  since  Bornet  during  the  some c a s e s  N.  in  in  1973).  typically  (Danjoy  in  Dellow  Toetz  spumigena from  temperatures "hot"  growth  However  12.9-33  1922;  1968;  collected  Reports  from  Sjostedt  Proshkina-Lavrenko appear  C.  15-30  values  was  was F  the  (p.45)«  106  The to  temperature be  similar  bilis  to  tially  of  under  field In  the  salt  white  light  the  measured Dellow in  in in  same r a n g e  area  with of  maximum g r o w t h the  field  the  effect  less  (1955) who  the is  rates  light  in  laboratory are  Cells  only in  Some c u l t u r e  by  the work  a  30  C may  of  Nodularia  shallow  is  to  the  of  is  experiments  that  of  in  was  ft-c.  a  restric-  algal  cultures.  given  distance  As  in  assessing Because  from  cells  at  the  surface  center  of  the  vessel  bluegreen  is  gave  (p.50). in  sea  This  which  encountered  of  intensity  zonation  ft-c)  shading  turbidity  light  300-750  and  high  r e d u c e d by  N_. h a r v e y a n a  involving  par-  to  water  and by  (200-600  intensity  at  of  The  difficult  Nodularia  that  intensity  on b l u e g r e e n  received  vessel.  note  intensity  the  are  describe algal  difficulties  light  that  light.  light  light  studies, of  They  growth  report  to  varia-  areas.  Ctenocladus)  regard  appears  1955a).  contribute  only  spp.  Anabaena  than  conditions  intensity  The  with  New Z e a l a n d .  the  culture  field  and C a s s i e  an  or  for  ponds,  light  i.e.,  less  tropical  shoreline  (Cladophora  the  to  source  the  of  values  saline  some l a k e s .  ted  shading  but  Nodularia  and Meyers  waters  intensity  around the  algae  water  to  absence from  inland  for  Nostocaceae,  and l a b o r a t o r y  intensity,  other  cave  its Light  assess.  the  some o t h e r  Nodularia  explain  Light:  of  maximum g r o w t h  and N o s t o c muscorum ( K r a t z  restriction  both  for  the of  of  light the  receive algae  107  (i.e.  M c L a c h l a n a n d G o r h a m 1962)  ities  (7500-9000 f t - c )  used.  In  dilute only  the  results range  growth  i n o c u l u m was  slightly  growth  from  show t h a t  of  light  were  to  compensate f o r  experiments  used, one  side  by  grown  grew  than  of  the  very  pH:  lowest  where  Hortobagyi  Woodson et  genus  is  than  present,  was from  pH i s  also  lates  of  9,  al.  8,  pH i s not pH  of  10.0  reflected  in  the  differed  other.  high  as the  as  The wide  cultures  of  direct  energy  source  except  Nodularia  Plinski  from  When  above a  less  laboratory  high  as  1973).  The  of  than  are  10.4*.  study although  8.2,  This  studies. rates  pH  containing  amounts  surveyed.  1955; Labbe  During this  8.  pH o f  pH 8  been  and  habitats  large  has  (Dellow  I966; S e r p e t t e  1972;  were  as  (p.36)  maximum r a t e s  as  that  s h o w e d maximum g r o w t h  some i s o l a t e s  densities  relatively  7.0-9.35  from  Nodularia.  to  intens-  (LL4).  Ortega  at  the a  grown  indicate  invariably  cell  but  c o l l e c t i o n s below  5«2  with  ft-c)  and W i l l i a m s  the  light  intensity  under  daylight  range  collected  Nodularia  grow  incubator  level  1966;  light  vessel to  commonly r e p o r t e d  with  numbers  the  Nodularia  than  Bayly  much more  small  habitats  light  high  intensities  with  the  pH v a l u e s  1959;  1966;  only  spp.  Published reports  collected  greater  any  the  light  Cultures  the  (2 5-600  intensities  sunlight.  at  of  high  described previously  so t h a t  Nodularia  inhibited  slower  used very  response All  at No  a  to  iso-  pH  growth  greater  108  occurred  at  pH 6 a n d v e r y  of  bluegreen  is  by  algae  Gerloff  et  little  at  pH 7.  The  s h o w i n g maximum g r o w t h  (1952)  al.  at  only  reports  a high  pH  (10)  (1962)  and M c L a c h l a n and Gorham  with Microcystis aeruginosa. Salinity: the  most  reports  thoroughly describe  "especially study 3-65  This  a wide  °/oo  Hutchinson  et  Verch  and B l i n n  reports  ++ Ca of  from  HCO^ , of  1900;  waters  C0^  the  Literature  c o l l e c t i o n of  a n d S0^  1917;  1937;  Rawson and Moore  Sjostedt  show from  1922; 1944;  and T i e s s  Nodularia  Na  a n d Mg  as  as  dominant  anions.  1964;  Ortega has  dominant  the  from  Nodularia,  Kalbe  i.e.,  reports  Moore  c o l l e c t i o n s where  many  present  salinities  1973).  Plinski with  Nodularia  In  is  terms,  P r o s h k i n a - L a v r e n k o 1968;  1966;  1972;  however,  general  1962).  II-VI). for  Nodularia  1972;  been  cations  There  dominant  are  ions  and no  are  + or  K  .  dominant  atory  much more  c o l l e c t e d at  Trahms  1932;  and W i l l i a m s  ,  Tables  (Levander  Bayly  CI  ecology of  R a t h s a c k - K u t z e n b a c h 1961;  1955;  reported  in  were  salinities  a_l.  the  (Prescott  isolates  of  0.12-35.7  Dellow  collections  (Appendix,  range  of  documented q u a n t i t a t i v e l y ;  hard water"  Nodularia °/oo  aspect  This Ca  or  may o n l y K  are  observations with  ( p . 5 5 ) show t h a t affected  by  the  growth dominant  reflect  the  uncommon o n  different rate  fact  a world  types  of  and s a l i n i t y  anion  or  that  cation  saline  basis.  dominant tolerance  present.  lakes Labor-  ions are  not  109  T h e maximum occurs  appears  showed t h a t  level  to  above  approximately  limit  of  tolerant.  Thus,  60 ° / o o .  f o r most  of  salinity  t h e most  technically  the isolates  growth  freshwater  of  salinity basal  itional  medium  Nitrogen: or  with  Sjostedt  ort  this  as  grew  i n medium  ability  of  is  (1922)  e n h a n c e d b y sewage view.  of  The l a b o r a t o r y as w e l l  lacking  Nodularia  with  5-20  i n which  it  occurs  not  different  On  in  refnature.  organisms.  patterns  isolates  when  of grown  isolate  grew  best  in  declining  with  any  add-  grew  better  correlation  at  10-20°/oo  of  nitrogen  or  absence of  Nodularia  that  t h e growth  of  but  presented  results  with  of  medium.  various  grow  less  °/oo,  "freshwater"  levels  NH.  +  of  no d a t a  N0^  source.  or urea  as  in  Nodularia to  ( p . 5 5 ) show t h a t  any combined n i t r o g e n to  upper  two i s o l a t e s  at  growth  stated  was t h e  some w e r e  that  bluegreen  little  60°/oo  rose  best  isolates  outflows  the s a l i n i t y  growth  47 ° / o o a n d 5 8 ° / o o .  grows  the presence  was  isolates  report  in the basal  There  levels  nature.  (1 ° / o o ) w i t h  than  observations  The freshwater  The marine  (NaCl)  Field if  noted  and marine  measurable  although  were  (1971)  gradients.  salts.  salinity  source  isolates,  (1933)  which  culture  common s a l i n i t i e s  and van Baalen  the  In  Nodularia  Batterton  on  60 ° / o o .  s h o w e d maximum t o l e r a n c e  gradient  lecting  at  was n o t p r e s e n t  Hof and Fremy  Nodularia a  salinity  be about  Nodularia  growth  of  suppa l l  (0.25-1.0 The i n nitrogen  g/1)  110  source have  is  interesting,  been too  high  for  however  the  growth,  since the  (1.0-6.1 m g / l ) .  much l o w e r  (0.5-1.0 g / l )  levels  Kratz  levels  in  that  Anabaena v a r i a b i l i s  and N o s t o c muscorum grew  ilar  concentrations  g/l  with  Nodularia  was  but  fix  The  cultures  present  nitrogen  in  in  medium w i t h o u t  ability  of  the  is  effect  that  other of  at  fixation  took  pH f i g u r e  800  place  these with  (and ant  made  the  at  data  those  the  the  sim-  used  fix  the  with  °/oo)  nearly (pH  a  salinity  the  The  data  only  fixation  marine  of  of  nitrogen  5 °/oo  Except  with  evidence  seen f o r  a  rates  proof  isolates  on n i t r o g e n  (1969) f o r  identical  the  (but  for  the  results  10).  observations  parameter  was  a n d pH 8.5.  environmental  the  two  nitrogen.  highest  fixed  definite  and c i r c u m s t a n t i a l  parameters  C,  30  are  regarding  of  to  reported  1-50  Nodularia  related  aspect  at  shows t h a t  and Stewart  They  and l a b o r a t o r y  can be  Jones  ft-c,  Considering ount  to  impure,  combined nitrogen)  isolates  isolate.  fixation  obtained  by  were  culture  environmental  reported  Calothrix  study  pure  (growth  on t h e  NH^ a n d u r e a )  on  study.  because the  lacking.  tested  this  of  well  are  report  (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  Stewart nitrogen  in  (0.5  nature  (1955a)  and Meyers  may  biology of  ecology of  pH) the  parameters  made,  some  taken  acc-  generalizations  of  Nodularia.  is  probably  genus.  into  The  the  Salinity most  algae,  import-  both  Ill  species,  typically  °/oo)  (5-30 or  are  higher  reduced sudden  and  than  although these  amounts. or  observations  erior  salinity  lag  salts  of  aria  rose  period  spp.  such as  nately  affect  Maine;  Fjerdingstad  ularia  has  umed t o waters  be  reported Moore tedt  (1922)  of  in  Baltic  from  the  (1917)  (1932) the  in  Lake  blooms  salt  in  Devils  South Sea  from in  Baltic  Africa at  at  2.3-2.5  Sea  21  Australia  do n o t  salinity  state  int-  weeks)  and  medium  with  dissolved  reports  marked  indicate  salinity  rainfall  water  and  (Collins  (or or  near  Dakota  14•5  °/oo;  Kalbe  29.1  values,  may  9.5  °/oo;  from  in  are Sjoset  (I964)  Early the  al. from  (I966)  and W i l l i a m s  but  ass-  situations  Hutchinson  °/oo.  Nod-  be  are  recorded  and T i e s s  and B a y l y at  at  from where  Bloom  was  alter-  1908  favorable),  salinity  a  Nodul-  surf  which  saline.  only  changes  Locations  quantities,  °/oo;  Corangamite,  Columbia  into  North  °/oo;  from  with  Denmark).  at  a few  evident  grew  bloom  Lake,  in  much  tolerate  is  British  in  1 °/oo  of  where  to  lower  at  where  brackish  present  This  the  waters  salinities  seem  °/00  where  conditions  either  the  in  65  content  1969  in  saline  usually  also  grown  pools  literature  in  to  15  recorded  are  ponds  introduced  seacoast  favorable  which  are  Some l i t e r a t u r e  occur,  been  occur  or  salinity.  conditions  the  may  spp.  (inoculum  when  in  the  brackish  they  in  from  °/oo).  live  values  (in  studies  40  they  changes  field  short  in  Nodularia  gradual  laboratory  found  records  locations  112  it  can be  inferred  (1878) t h e  tidal  and Thuret  at  Bornet Suhr  that  Lake  the  Alexandrina,  Deauville,  and F l a h a u l t  waters  France  and Hofman-Bang a l o n g the  (Kalbe  and T i e s s  1964  I878) were t o x i c  on t h e  brackish:  in  Sjostedt  bloom r e p o r t s Baltic  in  by  coast.  German B a l t i c  and r e s u l t e d  Francis  South A u s t r a l i a ;  (cited  cite  (1888)  were  death  of  Bornet 1922);  Schmitz,  Two  coast;  blooms Francis  livestock  and  poul-  try. Light salinity role  in  and temperature  and  areas  temperature  The  c o u l d be  bluegreen  algae  factors  saline  difficult  (Armitage  in  factor  to  water,  s h a d i n g by  other  affect  the  of  aria  spp.  amount at  temperature  higher is  explained  In  tropics.  or  Spirulina Jenkin  and t r o p i c a l in  nature  1932;  p o s s i b l y a more  in  the  than  spp. of  from light  and  inland  saline  tropical  inland  the Rich  between  areas.  dominant These  1932). brackish  or  Light  is  a  since turbidity  of  the  daylength  received.  altitudes  are  role  important  terms  the  plants,  light  in  from  1971;  measure  a more  Nodularia  obvious differences  temperate  lesser  s p p . a r e absent  Arthospira  would be  areas  in  a  but  absence of  since Nodularia  situations  growth  better  and b r a c k i s h a r e a s  saline  two  controlling  distribution.  tropical  lakes  pH i n  possibly play  and l a t i t u d e  The  presence of  tropics  important  all Nodul-  indicates  factor  than  that  light.  113  Taxonomy: There term  exists  "species"  organisms  is  arbitrary  in  of  the  variability  population uals  of  is  one  algae  has  are  algae  amount  among  of  in  the  The  and w i t h o u t  taxonomic  scheme i s natural  al  is  necessary for  The of  A great  distribution) ions sense  of  many  that  is  deal  can be  the  is  taxonomic  not  data  obtained  workers.  The  The  area  (for  is  also  sufficient algae  neglected (on  of  or  in  literature  confusion is  A  individ-  volume  bluegreen  example  a  -  workable obser-  and t h a t  (Komarek  cultur-  1971).  and misused  aspect  ecology, observat-  misused in  i n c r e a s e d by  Crow  I966,  apparent  accumulated is  of  than  algae.  variability,  from the  and  variability  described  variability,  It  often of  defined  this  an  Knowledge  collection  years  l i t t l e  surrounds  bluegreen  a  higher  characteristics  easily  for  the  in  1962, 1963; F j e r d i n g s t a d  bluegreen  an  it  variability  many  into  material  literature  study.  of  that  term has  taxonomy.  a  impossible.  of  than  more  within  concern for  vations work  is  in  variability.  the  extent  insight  The  sense of  1924; C a n a b a e u s 1929; D r o u e t 1969)  of  algae  same s e n s e a s  certain  "populations"  living  196 7 ) .  the  other  to  "population"  defined  been  in  prokaryotes.  essential  species  (Hutchinson  bluegreen  organisms with  is  certain  with  applicable  an u n d e f i n e d  variability a  not  bluegreen  group  possessing  within  problem  since they  significance  the  a  new  the species  114  or  variety  Nodularia being out  descriptions s p e c i e s have  retained,  akinetes),  var.  or  aerophila,  lacking  chemical  and  of  and the  from  terminal of  the  strain used  Berkeley  original  in  was  also  and  same  investigations  and Gorham  (1964).  B-1466)  in  the  Allen's  isolate  is  a  The  Nordin  Calothrix et  (N.  1181)  as  of  The  Granhall  Allen  This  (1967),  a  well  as  (#583)  shows  characteristics  (1972)  also  Mat-  material  sphaerocarpa, is  strain  derived  7103)  sp.  a_l.  nidulans  Bacteriology  sphaerocarpa)  et  collection  the  Collection  al.  bio-  a Nodularia.  (strain  Nodularia  I964).  not  examined.  Edelman  improper  by M.B.  Department  by Kenyon  by  is  differentiation  isolate  Gottingen (Koch  were  and b a s a l  reached  The  7103-  the  spumigena  filament  example,  Culture  California  culture,  heterocysts  (F  University  N.  (with-  Anacystis  isolated  evidently  at  with  For  culture  material  material  single  problem  problem  sphaerocarpa" is  type  (e.g.  a  Also,  p h y s i o l o g i c a l and  (p.l).  collection  Rivulariaceae  clusion  used in  mentioned  Indiana  and Immunology  material  similar  The  original  culture  the  Another  by Drouet  this  incomplete  d e s c r i b e d from  isolates  been  justification.  described without  inadequate  was  "Nodularia  from  cultures  from  p.80  of  identified  erial  with  investigations.  already  11.1.1  been  little  described with  akinetes).  identification  has  or  with  same  con-  studying has  been  Gorham  (i960)  (Culture  subculture  and H e n d r i c k s o n  of  (1969)  115  isolated  an  subculture istics late  of  alga of  this  isolate  genus N o d u l a r i a  the  generic  of  two  conditions  Certain  characteristics of  salinity).  The  IJ.  prior  regularly  20-30  Thuret  (1875)  midway  between  trichome  istic  is  sheath  spumigena and  in  to  cells),  are cell  a  other  is than  culture  the  character-  5 ym w i d e . also  the  different  the  a fairly  The  cited  iso-  by  bluegreen  algae  and m a t e r i a l genera  vary  always  (light,  wider  than  division) . filament  presence of  1966).  The  field  the pH,  long  in  (except  close  akinetes from  the  are  together both  are  Again t h i s  formed  center  of  character-  (1951).  Fritsch  in  under  emphasized by  heterocyst.  harveyana);  different  The h e t e r o c y s t s  and mature  by  have  temperature,  and f a i r l y  (1951).  present  may  under  little  taxon indicate  Pearson and K i n g s b u r y  Nodularia  detail  stable  morphological characteristics  usually N.  other  may v a r y  heterocysts  towards  be  characteristic  and F r i t s c h the  to  conditions used  cells  described in  However The  of  spaced along the  (every  the  with  1963;  (Stein  growth  harveyana  spumigena. A  and has  filaments  appears  characteristics  culture  range  examined  N.  s p e c i f i c name was  Studies  characteristics  wide  was  as  (1972).  morphologically. that  identified  with  incorrect  and Berg  The  they  a N_. h a r v e y a n a  using this  Granhall  which  material  but  it  is  the  sheath  not is  are  variable.  (more  always  so  N.  present  e v e n more  variable,  116  being Ca  absent  much more  of  the  hardness  as  a factor  a n d Mg  Oscillatoria  limosa  great  deal.  cells  to  several  alent  in  culture.  both  in  bright  nature  Because  akinetes  all  was t h e  of  The  but as  may b e  brown,  are  formed in  aspect  from  which  c o u l d be  be  extreme  distinctions  more  a  of  s p e c i e s of  result  plasticity of  many  species Anabaena  Whitton  several  has  is  or  more  prevdeal a  blue.  variable.  cells,  there  color  akinete.  correlated  a  dozen  a great  very  shape,  the  few  typically  green is  varies  are  and The  with  any  texsize  physical  a very  the  poor  a  been  Nodularia  in  the  judgement  in  the  extent  of  organism.  naming  variability, The  (especially questioned  literature  aptness  the  algae  of  distinct  grouping of  amorphous group  I968; B o u r crit-  Cylindrospernmm  forms;  and t h a t  generic  Oscillator-  (Drouet  commented t h a t  of  than  (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  bluegreen  represented was  of  of  Cyanophyta  sensu G e i t l e r )  for  is  in  parameter.  and m i s u n d e r s t a n d i n g the  eria  light  of  a  varies  color  size,  maturity  length  from  vegetative  formation;  species  1970).  color  cites  formation  condition  p r e v i o u s l y mentioned  multiplicity  elly  may v a r y  The  to  iaceae  sheath Filament  cell  appears  any  in  latter  culture.  depend upon t h e  chemical  The  (1964)  Foerster  Ag.  filaments  Vegetative  stages  only  C.A.  hundred.  and i n  akinete  a variety  or  nature  blue-green  The  ture  In  (Roth)  time.  whereas,  genera  in  the  117  Oscillatoriaceae  were  ears  to  be  Thus  it  must  be  ness  of  the  taxa  proper  similar  taxonomic  to  very  arbitrary.  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A manual o f f r e s h w a t e r a l g a e o f N o r t h C a r o l i n a . Tech. B u l l . # 1 8 8 , N o r t h C a r o l i n a Agr. Exp. S t a t . , R a l e i g h . 313 pp. W h i t t o n , B.A. 1962. M o r p h o l o g y - h a b i t a t a s s o c i a t i o n s i n b l u e green a l g a e . B r . P h y c o l . B u l l . 2: 167-171. W i l l e , N. 1897. 0m F a e r o e r n e s F e r s k v a n d s a l g e r og om F e r s k v a n d s a l g e r n e s Spredningsmaader. B o t . Not. 1897: 1-32. W i l l e , N. I924. S i i s s w a s s e r a l g e n von der Deutschen S i i d p o l a r E x p e d i t i o n auf dem S c h i f f "Gauss". T_n F. von D r y g a l s k i . , ed. , Deutsche S i i d p o l a r - E x p e d i t i o n 1901-1903. 8: 373-445. W i l l i a m s , W.D. 1970. Energy t r a n s f o r m a t i o n s i n s a l t B u l l . A u s t . Soc. L i m n o l . , 1: 6-8.  lakes. I  137  W i t t r o c k , V., Nordstedt, 0. and Lagerheim, G. 1897Algae aquae d u l c i s e x s i c c a t a e . Bot. Not. 1897: 75-94. W o l l e , F. 1887. Fresh-water Algae of the U n i t e d S t a t e s . Bethlehem, Penn. 364 pp. Woodhead, N. and Tweed, R.D. 1957. A c h e c k l i s t of t r o p i c a l West A f r i c a n a l g a e . H y d r o b i o l o g i a 11: 299-396. Woodson, B.R., Holoman, V.A. and Quick, A. I966. A d d i t i o n s t o f r e s h - w a t e r algae i n V i r g i n i a . I I . D i n w i d d i e County. J . E l i s h a M i t c h e l l Soc. 82: 154-159. Yondeda, Y. Geobot.  1937. Cyanophyceae of Japan, I . 6: 179-209.  A c t a Phytotax.  Young, R.T. 1924. The l i f e of D e v i l s Lake North P u b l . North Dakota B i o l . S t a t . 116 pp.  Dakota.  Zhadin, V . I . and Gerd, S.V. 1961. Fauna and f l o r a of the r i v e r s , l a k e s and r e s e r v o i r s of the U.S.S.R. I s r a e l Prog. S c i . T r a n s . , Jerusalem, No. 1903, 626 pp.  APPENDIX TABLES I  138  Table I.  Na , +  K, +  Procedures Used i n Water  Ca  , Mg''  :  Analysis  Perkin-Elmer Atomic A b s o r p t i o n Spectrophotometer Model 303.  S0^  :  T u r b i d i m e t r i c method (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  Cl  :  A r g e n t o m e t r i c method (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  :  P o t e n t i o m e t r i c t i t r a t i o n (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  Ortho-phosphate :  Stannous C h l o r i d e method (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  NH.  :  N e s s l e r i z a t i o n method (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  •  P h e n o l d i s u l f o n i c A c i d method (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , 1965.  HCO^  and  CO3  +  *4  N0^  Total Dissolved Solids  E v a p o r a t i o n at 105 C (Standard Methods) American P u b l i c H e a l t h A s s o c i a t i o n , I965.  Table I I .  pH temp C  Water c h e m i s t r y f o r Ctenocladus  30v72  21vii72  9.1  9.0  23viii72 8.8  Pond  24ix72 8.4  28iv73 8. 8  3vi73  21vii73  8.9  8.6  27  24  24  8  14  27  29  Na mg/1  8100  42500  56000  72000  13700  36000  95000  Mg mg/1  1800  5200  9700  12600  1200  4200  9200  Ca^mg/l  56  144  172  209  65  87  87  280  865  1235  1024  193  522  835  23500  108000  130000  170000  42000  100000  250000  C0~mg/1  0  100  480  1240  40  640  620  HCO~ mg/1  0  40  580  260  330  240  380  650  1750  5000  6400  600  1500  550  16  12  12  14  10  12  14  +  ++  K mg/l +  S0 "mg/1 4  Cl~mg/1 NH mg/l +  4  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 .  PH temp C  Water chemistry f o r Wallender  30v72  21vii72  8.7  8.65  23viii72 8.5  Lake  24ix72  28iv73  3vi73  21vii7  8.85  8.5  8.9  8.8  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  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  550  650  700  800  300  550  800  +  K mg/l +  CI"mg/1 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  T a b l e IV.  Water chemistry f o r Inks Lake  30v72 8.9  pH  21vii72 8.8  23viii72 8.8  24ix72  28iv73  3vi73  21vii73  9.0  8.6  8.8  9.1  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  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  550  650  700  800  300  550  800  temp C +  +  S0~  Cl"  mg/1  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  0.3  0.2  0.2  0.1  0.2  0.3  0.1  o-PO,  4  mg/1  TDS g/1  35  52  58  66  18  44  65  T a b l e V.  Water chemistry f o r White Lake  29v72  20vii72  PH  10.0  9.5  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  HC0 " mg/1  400  750  280  1050  490  600  1200  1000  900  1700  +  ++  +  3  Cl~  mg/1  23viii72 no c o l l e c t i o n  23ix73  27±v73  2vi73  9.4  9.7  9.7  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  0.2  0.3  0.1  0.3  0.3  4  TDS g / l  15  27  40  14  27  Table VI.  PH temp C  Water chemistry f o r t h e l a k e s i r r e g u l a r l y sampled o r sampled o n l y once.  S a l t Pond  S a l t Pond  4  3  39v72  21vii72  9.9  9.3  Big Q u i l l Lake  23viii72 8.6  Devils Lake  24viii72 9.1  Spotted Lake  Riefel Refuge  San d . Fuca  29v72  241x73  20v73  7.8  8.2  8.4  30  26  25  21  33  19  18  11400  2200  10300  1475  34100  2675  5370  41  30  3400  384  17000  320  651  404  0  126  56  92  200  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  Na mg/l +  Mg^mg/l Ca  1 |  mg/l  K mg/l +  6.5  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-P0  3.5  1.9  0.2  1.2  4.5  0.2  0.1  4  mg/l  TDS g/1  47  4  26  12  166  8.4  18  144  Table VII.  World d i s t r i b u t i o n of Nodularia  *  N. h a r v e y a n a N. s p u m i g e n a both  -  identification  only  to  genus  CANADA B r i t i s h Columbia * * B l i n n (1969) Kamloops *-::-Sismey (1922) n o l o c a t i o n *-Sismey (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.  Okanagan  L. V a s e u x  L.  Manitoba  **Bajkov  (1935)  several  Saskatchewan *-"-Kuehune (1941) -"-"-Rawson & M o o r e  locations  7 l o c a t i o n s i n south c e n t r a l (1944) 8 l o c a t i o n s i n s o u t h  Sask. central  Sask.  U.S.A. Alaska -"•Gardner  3963  Sitka  Arizona -"-Cameron (I963) -"--"-Cameron (I963) **Taylor & Colton  UC  661526  F  1047757  Yuma C o . & G i l a C o . Cochise Co. (1928) C o c o n i n o C o .  Arkansas -"-Demarree & T h o m a s o n  24582  Drew Co.  F  1133627  California -xx-Drouet (1943) M o d o c C o . -»-Drouet & M c B r i d e 4569 M o n o C o . & S a n B e r n a d i n o C o . UC 664554 F H N o r d i n 1076 N B V 3925-43 NY N o r d i n 1268 F 1101885 ( D r o u e t 1943) ^-Gardner 989 ( P B A IO63) M a r i n Co UC 341420 F H N o r d i n  1088  -"Gardner -"-Gardner ^-"Gardner -"Gardner  NBV 3925-48 1498  F 980816  Oakland  NY N o r d i n 1282  UC 202866  F Nordin  PC N o r d i n 1314  1182  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 F H N o r d i n 1078 F 1032981 NY N o r d i n 1266 1604  NBV 3925-42  145  ^Gardner 6552 B e r k e l e y UC 440248 •^Gardner 6568 B e r k e l e y UC 661705 FH N o r d i n 1075 NBV 3925-46 F 1033552 NY Nordin 1264 ---^Gardner 7231 San Mateo Co. FH Nordin 1081 ^-Gardner 7670 B e r k e l e y UC 641592 FH Nordin 1080 NBV 3 9 2 5 - 4 0 F 1 0 3 6 3 9 1 NY Nordin 1270 *-*Gardner 7946 Contra Costa Co. UC 661724 NBV 3 9 2 5 - 5 6 FH Nordin 1056 F 1043544 NY Nordin 1293 ^-Gardner 7963 San F r a n c i s c o UC 661654 FH Nordin 1077 NBV 3 9 2 5 - 4 1 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 3 9 3 9 3 1 UC 202712 NBV 3925-54 FH Nordin 1066 PC Nordin 1328 -"-"-Osterhout & Gardner (PBA 1 0 6 1 ) Oakland FH Nordin IO87 NBV 3 9 2 5 - 3 0 F 980816 NY Nordin 1263 PC Nordin 1302 **Setchell 1628 San B e n i t o Co. UC 752513 F 1221362 #Setchell 1679 San F r a n c i s c o UC I O O 5 6 3 *-x-Setchell & Jepson S t a n i s l a u s - M a r i p o s a Co. UC 202 713 NBV 3 9 2 5 - 5 3 Colorado *Louderback  17  Denver  F 1235673  Connecticut **Holden 1 4 5 8 - 1 4 6 1 B r i d g e p o r t FH Nordin 1062 NY N o r d i n 1290 ( C o l l i n s 1900, 1905) Florida *Brannon 141A Gainesville F 1124079 PC Nordin 1329 (Brannon 1952) -"-Drouet, Madsen & Crowson 1 1 5 H Wakulla Co. F 1 3 2 4 3 8 5 ^Drouet, Madsen & Crowson 11513 Wakulla Co. UC 912044 F 1324476 *Drouet & N i e l s e n 11232 T a y l o r Co. UC 912045 F 1 3 0 8 8 7 1 ---Drouet & N i e l s e n 11681 F r a n k l i n Co. UC 912043 F 1 3 1 7 6 1 6 -"-Standley 73311 Lee Co. F 1030027 Illinois -"Drouet, Glassman & Chapp 12696 Cook Co. F Nordin 1202 *Drouet, Glassman & Chapp 1 2 7 1 1 Cook Co. F Nordin 1203 -x--::--x-Tiffany & B r i t t o n (1952) no l o c a t i o n -x-x-x-T ran seau 63 Coles Co. NY Nordin 1283 ^Velasquez, R i c h a r d s & Drouet 2507 Cook Co. F 980549  146  Indiana **Palmer (1928) Knox Co. *Palmer et a l . 2518 I n d i a n a p o l i s **Transeau (1913) Decker Iowa -x-x-Prescott  (1931)  Dickenson  Kentucky -"Mclnteer  (1939)  no l o c a t i o n  F  980816  Co.  Louisiana ^-Drouet 8766 C a l c a s i e u P a r i s h F 1332708 PC Nordin 1311  UC  910404  NBV  3925-45  Maine , ^Collins (PBA 1062) Casco Bay F 545439 NY Nordin 1280 PC Nordin 1307 ---Collins 22 Cape R o s i e r UC 687705 FH Nordin 1072 NBV 3925-39 F 1209180 NY Nordin 1278 PC Nordin 1310 -"-Collins 2387 Cape R o s i e r FH Nordin 1070 FH N o r d i n 1073 NY Nordin 1275 * * C o l l i n s 2439 Cape R o s i e r FH Nordin 1060 NY Nordin 1287 -"-•'"-Collins 3459 Cape R o s i e r NY N o r d i n 1289 ^-Collins 5094 Casco Bay UC 752510 F 1219372 NBV 3925-27 **Collins 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 B r o o k s i s t e r 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 P t . FH Nordin 1071 Maryland *Wolle & Drouet 2302 Somerset Co. FH Nordin 1074 NY Nordin 1271 F 939834 UBC 49983 (Drouet 1939)  1  a p p a r e n t l y when C o l l i n s made a c o l l e c t i o n with t h e two s p e c i e s t o g e t h e r he made two sheets of t h e m a t e r i a l , c a l l i n g one f o r i n s t a n c e N o d u l a r i a harveyana and t h e o t h e r N o d u l a r i a spumigena. Both sheets show both s p e c i e s c o n t a i n e d i n them. He u s u a l l y gave both sheets t h e same c o l l e c t i o n number.  147  Massachusetts ^Chapman ( 1 9 4 0 ) Essex Co. *Collins Woods Hole UC 752511 •KFarlow ( 1 8 8 1 ) Cambridge **Farlow  Woods Hole  UC  100562  FH  4555  *Setchell Cambridge UC 100565 **T'aylor 3084 Nantucket I s l a n d UBC ^Webber ( 1 9 6 7 ) Essex Co. Michigan #Ackley (1931) Montcalm Co., **Ackley (1931) ^Johnson  **Phinney  Ann  (Drouet  L i v i n g s t o n e Co., Cheboygan C i t y & Washtenaw Co. & Shiawassee Co. Branch Co.  Arbor  23M40  49982  UC  752512  Emmet Co.  PC  N o r d i n 1300  F  1138192  F  1310578  F  1935)  Co.,  1157488  Mississippi **Drouet  *Scott Missouri *Casebolt  9844  Bay  Hancock Co.  St. Louis  167  F  C l a y Co.  IO8587O  F Nordin I I 9 8  ( G i e r & Johnson  1954)  Nebraska -"Anderson & Walker (1920) Cherry Co. -/-Clements ( I 8 9 6 & 1 9 0 1 ) L a n c a s t e r Co. & South Bend •$:-*Kiener 10401a F i l m o r e Co. F 1099262 x-Kiener 10466 Red W i l l o w Co. F 1099440 ^Kiener 13617 L a n c a s t e r Co. F 1127250 •K--«-Kiener 1 3 7 7 7 Cherry Co. UC 6 7 9 6 9 1 F 1127792 -SB'-Kiener  13885  Dodge Co.  -"--"Kiener  14139 14140  L a n c a s t e r Co. L a n c a s t e r Co.  -"r-"-Kiener  XKiener -"-Kiener *Kiener ^Kiener -"-Kiener -"-Kiener •5'-"Kiener  *~"-Kiener  F  1131624  F 1132181 F 1132185  15687 K e i t h Co. F 1139911 16499 Kearney Co. UC 6 8 9 1 7 4 F 1144655 20613a S h e r i d a n Co. F 1219490 21834-21835 Dundy Co. F 1220286 F 1220345  22773a Scott's B l u f f 22778 Scott's B l u f f 23130 Garden Co. F 23602-23604 Lincoln  Co. F 1249366 Co. F 1249569 Nordin I I 8 4 Co. F Nordin 1185-1187  Nevada -"--"Christiansen 1731 Washoe Co. F Nordin *-"-LaRivers 616 Washoe Co. F Nordin 1168  1169  148  New Hampshire X C o l l i n s Hampton * * C o l l i n s Hampton * * C o l l i n s Hampton NY Nordin 1 2 9 6 (Collins 1884)  NY Nordin 1 2 7 7 NY N o r d i n 1 2 7 9 ( C o l l i n s 1 8 8 4 ) NY N o r d i n 1 2 8 6 NY N o r d i n 1 2 9 2 F Nordin 1 1 7 8 FH N o r d i n 1 0 5 9  New J e r s e y x-X-peters A t l a n t i c C i t y  F  1081208  New York X-Martindale (1889) Staten Island *Pike (1886) Long I s l a n d x-X-Smith ( 1 9 2 4 ) Bergen North C a r o l i n a *Cocke ( 1 9 4 9 ) D u p l i n Co. **Cocke ( 1 9 4 9 ) Macon Co. x-Cocke (1967) Piedmont & C o a s t a l P l a i n **Cocke ( 1 9 6 7 ) Coastal Plain x-Whitford (1943) New Hanover Co. x-x-whitford ( 1 9 4 3 ) Craven Co. x-Whitford and Schumacher (I969) Piedmont and C o a s t a l P l a i n x-Whitford and Schumacher (I969) C o a s t a l P l a i n North Dakota x-x-Brannon (1911) D e v i l s Lake K-Moore ( 1 9 1 7 ) D e v i l s Lake -"-"-Moore & C a r t e r (1923) Ramsay, Nelson, Eddy & McLean Cos. x-x-yerch & B l i n n ( 1 9 7 2 ) D e v i l s Lake x-x-Young ( 1 9 2 4 ) D e v i l s Lake Ohio x - L i l l i c k & Lee -oi;-Lillick & Lee """"Riddle ( 1 9 0 5 ) Oklahoma x-x-Toetz  Columbus (1934) Cincinnati Champaigne Co. (1934)  Payne Co.  (1973)  Pennsylvania x-x-Habeeb 3 7 3 8  Pike Co.  F  Nordin  1176  Tennessee x-x-Bold G o o d e l e t s v i l l e F Nordin 1 1 7 4 x-Bold B 1 4 3 Nashville F 1211634 Utah x-Harding  (1971)  Utah Co.  149  Virginia *Louver & S t r i c k l a n d 1 1 3 0 York Co. UC 6 8 0 4 1 1 *0tt (1973) Yorktown **-Woodson e t a l . (I966) Dinwiddle Co.  F 1109792  Washington •x-x-x-Fairchild & Wilson (I967) Grant Co. ^-Gardner 3 3 5 (PBA 1 0 1 3 ) LaConner S k a g i t Co. UC NBV 3 9 2 5 - 3 8 F 546279 NY Nordin 1 2 6 9 PC Nordin ( S e t c h e l l & Gardner 1 9 0 3 ) -*-::-Gardner 4 1 1 (PBA 1 0 1 2 ) Whidby I s l a n d UC I O O 5 6 6 3925-72  NY N o r d i n 1 2 9 7  F 980816  F 1047781  IOO569 1306  NBV  PC N o r d i n  1334 ( S e t c h e l l & Gardner 1 9 0 3 ) ^Gardner 4 3 6 Port Townsend UC IOO567 ( S e t c h e l l & Gardner 1 9 0 3 ) ^Schumacher & Muenscher (1952) Whatcom Co. Wisconsin -x-Prescott (1962) x*-Prescott (I962)  no l o c a t i o n no l o c a t i o n  EUROPE Austria *x-Brabez ( 1 9 4 1 ) Franzenbader x-Brabez ( 1 9 4 1 ) Franzenbader •5'Forti (1907) c i t e s c o l l e c t i o n by Beck -x--x-Forti (1907) c i t e s c o l l e c t i o n s made by Hansgirg Belgium -x-Beeftink NBV  4 5 0 & 451  Fort St. P h i l l i p e  Loitlesberger,  NBV 3 9 2 5 - 3 2  3925-33  -x-Forti (1907) one by Bory  c i t e s two c o l l e c t i o n s by De Wildemann  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 a t C a r i n t h i a e -x-Hansgirg Hermanmestec F Nordin 1 2 3 3 -x-Hansgirg L i b o c h o v i c e F Nordin 1 2 3 4 -x--x-Kol ( 1 9 2 6 ) Lomniczi -x-x-Rosa ( 1 9 5 1 ) Bohmen •*Tarnavschi (1931) Bucovina  150  ' Denmark  **Cleve  (1897)  *Wille  (1897)  Helleback *"-*Fjerlingstad (I969) Knudshoved, S j a e l l a n d I s l a n d *-*0stenfeld Falster Island NY Nordin 1285 **0stenfeld 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 a t 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 & HofmanBang a t Hofmansgave; O s t e n f e l d at Taarback, Anhatt, Fyr, Storebaeltj Rosenvinge at Svendborg, V e j l e f j o r d , Faeno-Sund, L i m f j o r d e n ; Rosenvinge, I f f i l g e & Ostenf e l d at Oresund, Kjerteminde, F r e d r i d e s h a v e n , Groves F l a k , A a l b o r g Bugt, Nordre Runner, Hirsholmene; Rosenvinge & Th. Mortensen at Nykohing; Schmidt at Bornholm I s l a n d *-Sporring (1942) S k a l l i n g e n s Kirkebo,  Faeroe I s l a n d s  1901)  (Borgesen  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 S h i r e 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 a t Barmouth, R a l f s at D o l g e l l e y , Thwaites at Shirehampton "--x-Joshua C i r e n c e s t e r F Nordin 1249 -"-Petersen (1935) no l o c a t i o n #*--"-Stewart & Pugh (1963) L i n c o l n s h i r e *-*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  Helsinki  PC N o r d i n  1317-1319  France ---Bornet & F l a h a u l t (W&N 895) Cosne 1309 FH Nordin 1082 NBV 3925-34 F Nordin 1235 NY N o r d i n 1281 -x-x-Fremy (1934) B e l l e - I l e & B r e s t -"-"-Fremy & M e s l i n (1924) La Meaffe  UC NBV  (Nylander  759399 PC 3925-35  1861)  Nordin  151  -IH'-Gomont T a b l e s de l a L o i r e 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 ' H u b e r v i l l e NBV 3925-16 PC N o r d i n 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) B e r t h e n i c o u r t 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 N o r d i n 1063 PC Nordin 1327b (Bornet and Thuret 1880) Thuret ( i n Bornet & Thuret 1880) Croisic •Thuret? Cosne PC Nordin 1313 Thuret D e a u v i l l e FH Nordin 1065 NBV 3925-22 NBV 392524 PC Nordin 1327 PC N o r d i n 1331 x-x-villeret (1953) Bretagne Germany x-Anagnostidis & Schwabe (I966) Fehmarn I s l a n d x-x-Arndt et a l . (1966) Wismar-Bucht x-x-Bandel (1940) Rostoc x-X-Bornet & F l a h a u l t (1888) c i t e c o l l e c t i o n s by Braun ( v a r . gen. " i n s t a g n i s aquae d u l i s " ) ( v a r . l i t . " i n herb. Thuret") x-Bornet & F l a h a u l t (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 F r e i b u r g NBV 3925-15 PC Nordin 1338 x-x-Braun 1847 F r e i b u r g 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, R e i n b o l d , R i c h t e r not seen x-Forti (1907) c i t e s c o l l e c t i o n s by Brand, V o l k , M. Schmidt not seen x-x-Frohlich? S c h l e s w i g NBV 3925-57 NBV 3925-63 NBV 3925-70 PC Nordin 1326 PC Nordin 1335 x-x-Kalbe & T i e s s (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  (1910) Ostsee *Lindstedt (1943) & Forti Reinke, R e i n b o l d  **Kolwitz  **Mertens  Norderney  NBV 3 9 2 5 - 6 4 F 951302  NBV  ( 1 9 64)  *-*Pankow -"-Pankow -"-Petersen  UC 4 3 6 3 6 1 3925-66  Rostoc Ostsee,  (1971)  (1935)  (1907)  NBV  Kieler  NBV  c i t e c o l l e c t i o n s by 3925-15  3925-67  NBV  NBV  3925-59  3925-68  Forde  no l o c a t i o n  -*-"-Rathsack-Kutzenbach (I96I) Rugen I s l a n d -"-^•Rathsack-Kutzenbach (1961a) M i t t l e r e n Ostsee **Richter (1894) P l o n e r Sea •H-Rose B e r r a (Rabh. Algen 2 3 7 ) NBV 3 9 2 5 - 1 4 F 999057 NY Nordin 12 59 ^-"-Schmitz (H&R 1 4 2 ) G r e i f s w a l d e r Bodden UC 7 6 0 9 3 6 UC 9 5 3 5 4 2 NBV 3 9 2 5 - 5 8 NBV 3 9 2 5 - 6 5 PC Nordin 1 3 2 5 PC Nordin 1 3 3 2 *-*-Trahms ( 1 9 3 7 ) Rugen #-"Waldemann ( 1 9 5 9 ) middle Ostsee a l s o c i t e s c o l l e c t i o n s by Merkle, D r i v e r , H e s s l e and V a l l i n , O s t e n f e l d , Brandes, Rothe, Bandel Hungary -::->"-Fritsch (I964) c i t e s c o l l e c t i o n by P a l i k -"Hortobagyi (1959) Lake S z e l i d *-"-Palik  (1961)  *Tamas  (1958)  B a l a t o n Sea  Iceland ^--"Petersen  -"-Petersen  (1932)  (1932)  Einarsnes  Hvalnes & Knararnes & E i n a r s n e s  Ireland -"-Bornet & F l a h a u l t  **Rees  (1888)  c i t e s c o l l e c t i o n by Harvey  Cork Co.  (1935)  Italy -"Bornet & F l a h a u l t (1888) c i t e c o l l e c t i o n by Menghini -"-"Forti (1907) c i t e s c o l l e c t i o n by F o r t i Netherlands -"-Beeftink NBV  -"-Bierbr'auer  -"-Bilio ---Bilio -"-Bilio  0-199 & 0-201  Zeeuwsch-Vlaanderen  3925-11  2 3A  Ostvoorne  3925-12  Goeree NBV 392 5 - 7 Goeree NBV 3 9 2 5 - 9 Goeree NBV 3 9 2 5 - 2  61025-2  22B  NBV  NBV  3925-10  153  *Bilio 6 Noord-Beveland NBV 3925-4 x-Bilio 9 Noord-Beveland NBV 392 5-3 *Bilio W Noord-Beveland NBV 3925-5 x-Bilio M Noord-Beveland NBV 3925-6 x-Bilio M107 Ostvoorne NBV 3925-8 *Forti (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, S u r i n i g a r a t Leeuwarden *Simons & Vroman (1973) De P u t t e n xx-Van den Bosch Goes PC Nordin 1320 Norway **Foslie (WN&LI1344) Bugonaes UC 761551 NBV 3925-21 F 975766 NY Nordin 1295 Poland -**Collector unknown Danzig Bay -"-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  UC 100562  NBV 3925-23  Rumania •x-Hof & Fremy  (1933)  Szovata  Siebenburgen  Scotland «Bornet & F l a h a u l t (1888) c i t e c o l l e c t i o n by B a t t e r s •x-x-Bornet & F l a h a u l t (1888) c i t e c o l l e c t i o n by B a t t e r s a t t h e mouth o f t h e Clyde -x-Stewart (i960) Hebrides Spain •x~x-Gonzalez-Guerro  (1928)  Madrid  Sweden -:i--"-Borge (1907) Vaddo, Edeby •x-Cedergren (1926) K o l v i k e n •x-x-Cleve (1897) Moseskar Vaderdarna •x-Granhall & H e n r i k s s o n (I969) x-x-Lindstedt (1943) Torekovj Bohuslan: Fiskebiickskil Stockevik -x-Lindstedt (1943) s e v e r a l l o c a t i o n s -x~x-Nordstedt (W&N I98) Lomma FH Nordin 1090 NBV 3925-50 NBV 3925-61 NY N o r d i n 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 e x s i c e I I #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 W i n k l e r from west B a l t i c Coast Switzerland ---Cramer (Rabh. Alg.Sach. 994) Z u r i c h FH N o r d i n IO84 NBV 3925-1 F 1002450 NY N o r d i n 1262  EXCLUSIVE OF NORTH AMERICA AND Algeria *Debray  EUROPE  (1893)  Antarctica *-Apfel 51 & 52 Burger Lakes F 1299674 F 1299701 ^Fritsch Winter Harbour BM 1815 ( F r i t s c h 1912) •x-Fukushima (1959) Ongul I s l a n d A n t i l l e s (Dutch) W a n den Hoek et a l . -"-*Wagenear-Humelinck  (1972) Curacao 641 Aruba NBV 3925-52 (Koster  i960)  Argentina •5--"Borge (1901) Patagonia •"-"-Borge (1907a) Puna de Atacama -"-DeHalperin (1970) G u l f o Nuevo Australia *~"Bayly & W i l l i a m s (I966) Lake Corangamite -"-"-Francis? Lake A l e x a n d r i n a FH Nordin 1064 *~"-Playf a i r (1914) Lismore **Schmidle (I896) -x-ftWilliams (1970) Barbados -"-West  (1904)  Chancery  Lane E s t a t e  China -"Skvartzow (1927) North Manchuria *~"-Skvartzow (192 7) H a r b i n , North Manchuria Columbia -"-West (1914) Cundinamarca  ( F r a n c i s I878)  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 B i r k e t Quarun BM 302 (West 1909) Falkland Islands x-x-Guarrera & Kuhnemann at M a l v i n a s Formosa **0kada Guatemala **Standey x-Tilden  (1932)  (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  Kotosho  65787 R i o P u c a l Dept. (1908) Lake A m a t i t l a n  F 1023960  India -x-x-Bharadwaja (1935) Benares -Chacko (1972) Madras -Franklin (1972) Madras *--x-Iyengar & D e s i k a c h e r y (1944) Hare I s l a n d , South •x-x-Kamat (1968) A l i b a g , Maharashtra x-x-Kumar (1970) Sardhana x-Raju (1972) Kamat **Raju (1972) Kanpur -x-x-Randawa (1936) N. I n d i a x--x-Subrahmanyam (1972) B a s t a r , J a g d a l p u r x--x-Vasishta (1961 & i960) Hoshiarpur Israel -Jabotinsky  (I96I)  Lake  India  Zohar  Japan -x-Umezaki (I96I) s e v e r a l l o c a t i o n s **Yoneda (1937) Shinano Java -x-Mobius (1893) S o l o **Van Oye (1922) B a t a v i a W a n Oye (1923) T a s i k m a l a j a -x-Jutono (1973) Jogjakarta Mexico -x-Drouet & R i c h a r d s 3416 Empalme, Sonora -Ortega (1972) Lake Texcoco -x-Patrick Lake Texcoco F Nordin 1241  F 1031425  156  New Zealand -"-Chapman (1955) Stanmore Bay ^-Chapman l b S t a n l e y Bay F 1246844 *Dellow (1955) Hauraki G u l f *Dellow & C a s s i e (1955) Whangaparaoa Bay Peru **Ibanez Trujillo F Nordin 1238 **---Maldonaldo Lago V i l l a F 1102709 Puerto R i c o **Wille 1817a Laguna Guanica (Gardner 1927) S i e r r a Leone ***Woodhead & Tweed  (1957)  no  F 1102707  UC 4 6 3 7 3 5  NY N o r d i n  1256  location  South A f r i c a * * F r i t s c h & Rich (1929) Kimberley -"-^•Hutchinson et a l . ( 1 9 3 2 ) T r a n s v a a l ( R i c h 1 9 3 0 ) ( R i c h 1934) -^-Pearson 26 Orange R i v e r BM Nordin 1348 ( F r i t s c h 1918) South West A f r i c a "Welch (I964) Swakop R i v e r , R i e t f o n t i e n S p r i n g -xx-Welch (I964) Gross Barmen, Etosha Reserve *~--Welch ( 1 9 6 5 ) Okandu *West (1912) L i t t l e Namaqualand Sudan **Karim  (I968)  J e b e l Marra  Tunisia -"-Serpette & Labbe (I966) between Rades and M i l i a n e * * S e r p e t t e & Labbe (1966) O a s i s de Tozeur U.S.S.R. *Elenkin (1916) K i s l o v o d s k -"Kosinskaja T a r t a r 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 ( 1 9 7 2 ) 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 ( 1 9 6 1 ) Lake Balkash  157  Table VIII.  Medium BG-11 ( S t a n i e r et a l . 1971)  Compound  Amount (g/1)  NaN0  1.5  3  K HP0 2  0.04  4  MgS0 -7H 0  0.075  CaCl .2H 0  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  4  2  2  2  A5  H B0 3  2.86  3  MnCl .4H 0  1.81  ZnS0 .7H 0  0.222  2  2  4  2  Na Mo0 .2H 0  0.39  CuS0 .5H 0  0.079  2  4  2  4  2  Co(N0 ) .6H 0 3  notes:  :  2  0.0494  2  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 after cooling. t h e Na C0 should be a u t o c l a v e d sepa r a t e l y i f t h e d e s i r e d 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 c o o l i n g . 2  158  T a b l e IX.  Herbarium  m a t e r i a l of N o d u l a r i a examined.  N o d u l a r i a spumigena B e r k e l e y no date B r i s t o l , England NBV 3 9 2 5 - 2 7 Bold 28 i i i 1953 G o o d e l e t s v i l l e , Tenn. F Nordin 1174 Braun 1847 F r e i b u r g , Germany NBV 3 9 2 5 - 1 5 PC Nordin 1338 (type of Kiitz. S. V r i e s i a n a ) Braun 1847 F r e i b u r g , Germany NBV 3 9 2 5 - 2 5 PC Nordin 1336 Brebisson? no date no l o c a t i o n NBV 3 9 2 5 - 7 1 Christensen 1731 14 v 1953 Washoe Co., Nev. F Nordin 1169 c o l l e c t o r unknown ( K l e i b a d e n ? ) v i i 1839 Wangerooge, Germany NBV 3 9 2 5 - 1 3 c o l l e c t o r unknown 5 v i i i 1887 Danzig Bay, Poland NBV 3 9 2 5 -  23  Collins no date West B r o o k s i s t e r , Maine NY Nordin 1276 Collins v i i i I 8 8 4 Rockingham Co., N.H. FH Nordin 1059 F Nordin 1178 NY Nordin 1286 NY Nordin 1292 NY Nordin 1296 Collins v i i i I884 Hampton, N.H. NY Nordin 1279 Collins 2439 v i 1892 Cape R o s i e r , Maine FH Nordin 1060 NY Nordin 1287 Collins 3459 14 v i i 1897 Cape R o s i e r , Maine NY Nordin 1289 C o l l i n s 5094 v i i 1904 Casco Bay, Maine UC 752514 F 1219336 Collins 5526 (Phycotheca B o r e a l i s Americana 1307) 14 v i i 1906 H a r p s w e l l , 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., M i s s . F 1310578 Farlow v i i I 8 8 9 Woods Hole, Mass. UC 100562 FH 4555 Foslie ( W i t t r o c k , Nordstedt & Lagerheim, Algae E x s i c c a t a e 1344) 10 v i i i 1889 Bugonaes, Norway UC 761551 UC 100562 NBV 3 9 2 5 - 2 1 F 975766 NY Nordin 1295 Francis?* I878? Lake A l e x a n d r i n e near A d e l a i d e , A u s t r a l i a FH Nordin IO64 Gardner 4 1 1 (PBA 1 0 1 2 ) v i 1 9 0 1 Whidbey I s l a n d , Wash. UC 100566 NBV 3 9 2 5 - 7 2 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 I 9 3 6 Contra Costa Co., C a l i f . UC 661724 FH Nordin 1056 NBV 3 9 2 5 - 5 6 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 F r a n c i s a p p a r e n t l y d i s t r i b u t e d h i s samples of N o d u l a r i a from t h i s s i t e . Bornet & F l a h a u l t ( 1 8 8 8 ) r e p o r t examining m a t e r i a l from him. 1  159  Gomont? 30 v i i i 1885 T a b l e s de l a L o i r e , France PC Nordin 1323 Gomont? 15 v i i i 1894 Auvergne, France PC Nordin 1324 Gomont ( W i t t r o c k , Nordstedt & Lagerheim, Algae E x s i c c a t a e 1343) v i i i 1894 Auvergne, France UC 761550 NBV 3925-49 F 975767 NY Nordin 1294 Habeeb 3738 15 v 1951 P i k e Co., Pa. F Nordin 1176 Holden 1 4 5 8 - 1 4 6 1 28 v 1899 B r i d g e p o r t , Conn. FH Nordin 1062 NY Nordin 1290 Holden 30 v i 1899 no l o c a t i o n NY Nordin 1291 Ibanez 27 x 1952 T r u j i l l o , Peru F Nordin 1238 Joshua no date C i r e n c e s t e r , England F Nordin 1249 Kiener 13777 22 v i i 1936 Cherry Co., Nebr. UC 67969I F 1127792 Kiener 21 v i i 1941 F i l l m o r e Co., Nebr. F 1099262 10401a Kiener 23 i v 1943 Dodge Co., Nebr. F 1131624 13885 Kiener L a n c a s t e r Co. Nebr. 14139 & 14140 6 v i 1943 F 1132185 F 1132181 Kiener 16499 6 i v 1944 Kearney Co., Nebr. UC 689174 F 1144655 Kiener 21834 F 1220345 27 i i i 1947 Dundy Co., Nebr, Kiener 21835 F 1220286 27 i i i 1947 Dundy Co., Nebr, Kiener 23130 2 i v 1948 Garden Co., Nebr. F Nordin II84 Kiener 23602-23604 17 v 1948 L i n c o l n Co., Nebr. F Nordin 1185 F Nordin 1186 F Nordin 1187 Koch 3 v i i 1845 Borkum, Germany NBV 3 9 2 5 - 1 7 NBV 3 9 2 5 - 2 9 a Koch 3 v i i I846 no l o c a t i o n NBV 3 9 2 5 - 2 9 b LaRivers 6lb 22 i x 1951 Washoe Co., Nev. F Nordin 1168 Lebel 415 14 v i 8 6 0 Mont d ' H u b e r v i l l e , France NBV 3 9 2 5 - 1 6 PC Nordin 1337a L e b e l 58O 12 i v 1862 N e g r e v i l l e , France PC Nordin 1337b Maldonado 41 i 1942 Lago V i l l a , Peru F 1102707 F 1102709 Mertens v i 1 8 2 1 Norderney, Germany UC 4 3 6 3 6 1 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 Nordstedt ( W i t t r o c k & Nordstedt, Algae E x s i c c a t a e I98) 11 v i 1877 -Lomma, Sweden FH Nordin 1090 NBV 3 9 2 5 - 5 0 NBV 3 9 2 5 - 6 1 F Nordin 1250 NY Nordin 1298b Nylander i 8 6 0 Helsinki, Finland PC Nordin 1317 PC Nordin 1318 PC Nordin 1319 Ostenf e l d 5 v i i i 1901 F a l s t e r , Denmark NY Nordin 1285 Ostenf e l d 1 v i i i 1904 S t o r e B e e l t , Denmark NY Nordin 1284 Osterhout 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 3 9 2 5 - 5 4 FH Nordin 1066 PC Nordin 1328 Osterhout & Gardner (PBA 1061) 30 v 1902 Oakland, C a l i f . FH Nordin IO87 NBV 3 9 2 5 - 3 0 F 9808I6 NY Nordin 1263 PC Nordin 1302  160  P e t e r s 14 v 1891 A t l a n t i c C i t y , N.J. F 1081208 Phinney 23M40 8 i i 1 9 4 0 Emmet Co., Mich. F 1138192 Rose v i i 1852 B e r r a , Germany NBV 3925-14 F 999057 NY Nordin 1259 Schmidt 1899-1900 Siam? PC Nordin 1322 Schmitz, Hauck & R i c h t e r (Phykotheca U n i v e r s a l i s 1 4 2 ) v i i i 1886 G r e i f s w a l d e r Bodden, Germany UC 760936 UC 953542 NBV 3925-58 NBV 3 9 2 5 - 6 5 PC Nordin 1325 PC Nordin 1332 Setchell 1628 14 i v 1897 San B e n i t o Co., C a l i f . UC 752513 F 1221362 S e t c h e l l & Jepson v i i I 8 9 6 between La Grange, S t a n i s l a u s Co. & C o u l t e r v i l l e , Mariposa Co., C a l i f . UC 202713 NBV 3 9 2 5 - 5 3 Standey 65787 20 i i 1939 R i o P u c a l Dept., Guatemala F 1023960 Suhr ( F r o l i c h ? ) v i i 1834 S c h l e s w i g NBV 3925-57 NBV 3925-63 NBV 3925-70 PC Nordin 1326 PC Nordin 1335 (type of K u t z i n g s N. s u h r i a n a = N. spumigena Mertens & Suhr's Lyngbya annulata) Taylor 3084 29 v i i i 1 9 2 0 Nantucket I s l a n d , Mass. UBC 49982 Transeau 63 14 v 1911 C h a r l e s t o n , 1 1 1 . NY Nordin 1283 Thuret 30 v i i i 1874 Cherbourg, France FH Nordin IO63 PC Nordin 1327b Thuret? v i i i 1874 D e a u v i l l e , France FH Nordin 1065 NBV 3 9 2 5 - 2 2 NBV 3925-24 PC Nordin 1327 PC Nordin 1 3 3 1 Thwaites? v 1867 Shirehampton, England PC Nordin 1330 Van den Bosch no date Goes, Netherlands PC Nordin 1 3 2 0 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 B i r k e t Quarun, Egypt BM 302 Wille 1 8 1 7 a Laguna Guanica, Puerto R i c o UC 4 6 3 7 3 5 NY Nordin 1256 Nodularia Apfel  harveyana  51 & 52 19 i 1948 Burger Lakes, A n t a r c t i c a F 1299674 F 1 2 9 9 7 0 1 Atkinson 1895 no l o c a t i o n PC Nordin 1301 A t k i n s o n 1895 no l o c a t i o n PC Nordin 1321 Beeftink 0 - 1 9 9 29 v i i i 1951 Zeeuwsch-Vlaanderen, Netherlands NBV 392 5-10 Beeftink 0 - 2 0 1 29 v i i i 1951 Zeeuwsch-Vlaanderen, Netherlands NBV 3 9 2 5 - 1 1 B e e f t i n k 4 5 0 28 v i i 1955 F o r t S t . P h i l l i p e , Belgium NBV 3925-33  161  Beeftink  451  NBV  28 v i i 1955  Fort St. P h i l l i p e ,  Belgium  3925-32  Bierbrauer 6 v i 1953 Ostvoorne, N e t h e r l a n d s NBV 3925-12 Bilio 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 Bilio 6 6 i x I966 Noord-Beveland, Netherlands NBV 3925-4 Bilio 9 6 i x 1966 Noord-Beveland, Netherlands NBV 3925-3 Bilio W 6 i x 1966 Noord-Beveland, Netherlands NBV 3925-5 Bilio M 6 i x I966 Noord-Beveland, Netherlands NBV 3925-6 Bilio 22B 17 x I966 Goeree, Netherlands NBV 3925-2 Bilio 23A 17 x I966 Goeree, Netherlands NBV 3925-7 Bold B143 12 i 1947 N a s h v i l l e , Tenn. F 1211634 Bornet & F l a h a u l t ( W i t t r o c k & Nordstedt, Algae E x s i c c a t a e 895) 25 i x 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 C l a y Co., Mo. F Nordin 1198 Chapman l b i x 1947 S t a n l e y Bay, New Zealand F 1246844 Collins 4844 no date no l o c a t i o n NY Nordin 1273 NY Nordin 1274 Collins v i i I884 Hampton, N.H. NY Nordin 1277 Collins 2387 14 v i i 1892 Cape R o s i e r , Maine FH Nordin 1070 FH Nordin 1073 NY Nordin 1275 Collins 22 v i i 1894 Cape R o s i e r , Maine UC 687705 FH Nordin 1072 NBV 3925-39 F 1209180 NY N o r d i n 1278 PC Nordin 1310 Collins (PBA 1062) 14 v i i 1903 Casco Bay, Maine F 545439 NY Nordin 1280 PC Nordin 1307 Collins 5094 v i i 1904 Casco Bay, Maine UC 752510 NBV 3925-37 F 1219372 Collins 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 i 8 6 0 Z u r i c h , S w i t z e r l a n d FH Nordin IO84 NBV 3925-1 NY N o r d i n 1262 F  1002450  Demaree & Thomason Ark.  Drouet NBV  F  24582  1 viii  1943  Collins,  Drew  Co.,  1133627  8766  28 x 1948  3925-45  C a l c a s i e u P a r i s h , La.  F 1332708  PC N o r d i n  UC  910404  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 N o r d i n 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 & N i e l s e n 11232 23 i 1949 T a y l o r Co., F l a . UC 912045 F 1308871 Drouet & N i e l s e n 11681 31 i 1949 F r a n k l i n Co., F l a . UC 912043 F 1317616 Drouet & R i c h a r d s 3416 23 x i i 1939 Empalme, Mexico F 1031425 Fritsch no date Winter Harbour, A n t a r c t i c a 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 B e r k e l e y , C a l i f . UC 202865 u c 274104 Gardner 3963 v i i 1917 S i t k a , A l a s k a 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 B e r k e l e y , C a l i f . UC 440248 Gardner 6568 12 i 1931 B e r k e l e y , C a l i f . UC 661705 FH Nordin 1075 NBV 3925-46 F 1033552 NY Nordin 1264 Gardner 7670 4 i i 1934 B e r k e l e y , 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 P a r i s , France PC Nordin 1303 Gomont 11 v i i 1892 Cosne, France PC N o r d i n 1304 Groesbeck 90 12 v i 1940 Mono Co., C a l i f . F 1049304 H a n s g i r g v i i 1888 L i b o c h o v i c e , C z e c h o s l o v a k i a F Nordin 1234 (Hansgirg 1892) H a n s g i r g 1891 Hermanmestec, C z e c h o s l o v a k i a F Nordin 1233 Hollenberg 1553 23 i i i 1934 Santa Ana R i v e r , 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 L a n c a s t e r Co., Nebr. F 1127250 Kiener I5687 24 i x 1943 Keystone, Nebr. F 1139911 Kiener 20613a 23 v 1946 S h e r i d a n Co., Nebr. F 1219490 Kiener 22773a 23 v i i i 1947 S c o t t ' s B l u f f Co., Nebr. F 1249366  163  Kiener  22778  24 v i i i  1947  S c o t t ' 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, C o l o . F 1235673 Louver & S t 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 P t . , Maine FH Nordin 1071 Palmer, Webster, Prettyman, Webster & Drouet 2518 17 v i i i 1939 I n d i a n a p o l i s , Ind. F 9808I6 Patrick 197 24 v i i 1947 X o c h i m i l c o , Mexico F Nordin 1241 Pearson 26 no date South A f r i c a BM Nordin 1348 R o u s s e l 16 v i i i I869 T a l a t u s ? , France PC Nordin 1315 Scott 30 i i i 1941 Bay S t . L o u i s , M i s s . F IO8587O Setchell 1679 no date San F r a n c i s c o , C a l i f . UC IOO563 Setchell 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, R i c h a r d s & 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 f o l l o w i n g specimens were i n t o o poor a c o n d i t i o n o r i n i n s u f f i c i e n t numbers t o make a judgement on t h e i r i d e n t i f ication. c o l l e c t o r unknown i x 1885 P a v i a , Germany NBV 3925-26 Drouet & Louderback 5739 21 v i i i 1946 S a l t Lake Co., Utah F 1198553 Drouet & R i c h a r d s 2709 26 x 1939 S i 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 I s l a n d , 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 E x s i c c a t u s 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 ( T a y l o r 1928a) uncatalogued m a t e r i a l a t UBC  164  West West West West West  44 1 v i i i 1904 Niamkolo, Tanganyika BM Nordin 1344 22 23 v i 1904 Nkata Bay, Tanganyika BM Nordin 1346 134 10 x i 1904 Komba Bay, Tanganyika BM Nordin 1347 208 10 i 1905 Toa, Tanganyika BM Nordin 1345 432 10 i 1905 Toa, Tanganyika BM 303  The f o l l o w i n g m a t e r i a l i s more p r o p e r l y p l a c e d i n a genus other than N o d u l a r i a 7 v i i 1853 L e i p z i g , Germany FH Nordin 1068 = Scytonema sp. Allen 5079a 2 i v 1953 from herbarium specimen F Nordin 1181 = C a l o t h r i x sp. Crossland 7292 23 x 1929 T a h i t i UC 696203 - Hormothamn i o n (solutum?) Born. & Grun. Fritsch 363 no date Winter Harbour, A n t a r c t i c a BM 1934 = Anabaena sp. Gardner 3305 9 v 1916 B e r k e l e y , 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 U l t d o b e r n , Germany FH Nordin 1067 NBV 3925-28 NY Nordin 1260 = Scytonema sp. Rabenhorst (Algen Sachsens 470) v i 1855 Z u r i c h , S w i t z e r l a n d UC 432853 FH Nordin IO89 NBV 3925-60 NBV 3925-69 F 1015345 F 1015346 F 1015349 NY Nordin 1261 = Anabaena sp. R o u s s e l 4 i x 1851 "Meloduuo", France PC Nordin 1312 = Anabaena sp. Rrealea x 1857 A t t e l o b e r n , Germany NBV 3925-19 NBV 392520 = Scytonema sp. Setchell 23 v i i i 1889 W a l i t e H i l l Pond, R.I. UC IOO564 = Nostoc (carneum?) Ag. Tilden (American Algae 484) 22 v 1900 Oahu, Hawaii UC 740394 NY Nordin 1258 PC Nordin 1339 UBC 2971 - Hormothamn i o n sp. Tilden (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. Wille 282d 6 i 1915 Coamo S p r i n g s , Puerto R i c o UC 401708 FH Nordin 1085 NY Nordin 1255 = Scytonema sp. Nostoc sp. Wille 1415a no date Puerto R i c o UC 401800 NY Nordin 1254 = Scytonema sp.Nostoc sp. Aavd?  165  Table  X.  Synonomy et  Nodularia  of  Nodularia  Flahault  spumigena  /Mertens  in  Juergens7  1888  /Mertens  in  Juergens.7  Bornet  IT. s p u m i g e n a  var.  genuina  Bornet  et  Flahault  19.  spumigena  var.  litorea  Bornet  et  Flahault  19.  spumigena  var.  ma j o r  II. s p u m i g e n a  var.  vacuolata  Fritsch  19.  var.  aerophila  Brabez  spumigena  II. s p u m i g e n a N.  armorica  N.  willei  Nodularia 19.  f.  e r a s sa  harveyana  Elenkin  Thuret  sphaerocarpa  1875  (Bornet  et  Flahault)  Gonzalez-Guerro  spumigena  N.  turicensis  Bornet  var.  minor  (Cramer)  et  Flahault  Fritsch Hansgirg  Excludendae  epiphytica  Gardner  II. f e r t i l i s s i m a II. f u s c a  = Nostoc  Randawa  Taylor  =  nom.  sp.  (juvenile)  nud.  conglomerate  N.  hawaiiensis  N.  implexa (Bornet et F l a h a u l t ) Bourrelly = A u l o s i r a implexa Bornet et Flahault  19.  quadrata  II. s p u m i g e n a N.  Elenkin  Brabez  N.  N_.  1888  Flahault  (Woronichin)  (Thwaites)  var.  II. s p h a e r o c a r p a  Species  et  Flahault  Gardner  harveyana  s k u j ae  Bornet  et  Thuret  II. a e r o p h i l a N_.  Bornet  tenuis  Tilden  Fritsch var.  G.S.  = Hormothamnion  = Anabaena  zu j a r i s  West.  ?solutum  et  sp.  Gonzalez-Guerro  = Anabaena  Bornet  = Anabaena  sp.  continued.  sp .  Flahault  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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