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Stemflow and throughfall mycobiota of a trembling aspen-red alder forest MacKinnon, J. Andrew 1982

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STEMFLOW  AND THROUGHFALL OF  TREMBLING  MYCOBIOTA  A  ASPEN-RED  ALDER  FOREST  by J .  B . S c ,  University A  THESIS THE  accept  THE  MACKINNON  B r i t i s h  Columbia,  Vancouver,  SUBMITTED IN PARTIAL FULFILLMENT REQUIREMENTS FOR T H E DEGREE OF MASTER OF SCIENCE in  THE  We  Of  ANDREW  OF  F A C U L T Y OF GRADUATE STUDIES (Department of Botany)  this thesis required  UNIVERSITY  OF B R I T I S H  May  J .  as conforming standard  Andrew  to  COLUMBIA  1982  MacKinnon,  1982  the  1979  In  presenting  requirements of  B r i t i s h  i t  freely  agree for  this for  an  available  that  I  understood  that  f i n a n c i a l  by  his  that  or  be  her or  s h a l l  of  V6T 1Y3  Date  DE-6  (3/81)  the  University  and  study.  I  copying  granted  by  the  of  publication be  allowed  Columbia  of  make  further this  head  representatives.  not  the  s h a l l  /^^)T/9/UV  The U n i v e r s i t y o f B r i t i s h 1956 M a i n M a l l Vancouver, Canada  at  of  Library  permission.  Department  fulfilment  the  extensive  may  copying  gain  degree  reference  for  purposes  or  p a r t i a l  agree  for  permission  scholarly  in  advanced  Columbia,  department  for  thesis  It  this  without  thesis  of  my  is thesis my  written  ii  ABSTRACT The  s t e m f l o w and  forest  were sampled o v e r  preserved,  concentrated,  spores.  R e c o r d e d on  Seimatosporium, spore Ovate  types  species  to  five  Gyoerffyella yakuensis,  of and  dates  one  date  Dicranidion  Cruciform found  to  possible  and  Conidium. be role  stemflow/throughfall  i  and  Crescent  were  Conidium,  Phragmospore,  and  H-Shaped were  Gyoerffyella  forest  ecosystem  Triposporina  the  Dematiaceous  Cornutispora  with  on  of C e r a t o s p o r e l l a ,  Conidium  correlated  recorded  C o n i d i u m ; and  sporulation  i s postulated.  Pestalotiopsis,  uni-inflata,  Branched  and  Ceratosporium  aquatica,  species  Triradiate  Seasonal  Spore,  biappendiculata,  T r i c l a d i u m , and  Lateriramulosa  Didymospore,  Helicospore;  elodeae,  the  fungal  Fusarium, Tripospermum, T i t a e a ,  Clavariopsis  Tiny  Samples were  of  were  positively i n the  dates  aspen  species  were T e t r a p l o a a r i s t a t a ,  fragile,  Dendrospora,  ten  the T a p e r e d  Varicosporium  only  year.  s i x to ten dates  gemellipara,  C o n i d i u m and  one  F l a b e l l o s p o r a , Helicoma,  Stauronema, T e t r a c l a d i u m , T-Shaped  than  Large on  of  examined m i c r o s c o p i c a l l y f o r  Double  recorded  Scolecobasidium,  two  and  Alternaria,  Phragmospore,  cornutum,  the c o u r s e  more  designated  Scolecospores;  and  t h r o u g h f a l l i n a red a l d e r - t r e m b l i n g  a  and  of  recorded  on  1ichenicola, species  of  Dematiaceous fungi  was  precipitation.  A  for fungal  most  propagules  in  iii  TABLE OF CONTENTS  ABSTRACT  i i  L I S T OF TABLES  iv  L I S T OF FIGURES  v  ACKNOWLEDGEMENTS  vi  Introduction  1  Materials  7  a n d methods  Results  15  Spore  t y p e s r e c o r d e d on more t h a n  ten dates  Spore  t y p e s r e c o r d e d on s i x t o t e n d a t e s  45  Spore  types  49  Spore  t y p e s r e c o r d e d on one d a t e  r e c o r d e d on two t o f i v e  17  dates  only  53  Discussion  57  Appendix  1 calculation  Appendix  2. M e d i a e m p l o y e d  Malt-yeast-peptone  of stemflow  Oat  agar  Appendix  3.  rainfall References  volumes  (Bandoni  (SORB-T) medium  1972)  (Bandoni  75  1981) .... 75  (OA) medium Variations  72 75  (MYP) medium  Sorbose-tetracycline  and t h r o u g h f a l l  76 in  spore  concentration during a 77 78  i v  LIST  Table  I.  Chemical  modification  OF  TABLES  of  r a i n f a l l  by  the  forest 2  canopy Table  II.  Table  III.  a  Collection Variation  r a i n f a l l  numbers in  and  dates  stemflow  spore  12 concentration  during 77  V  LIST  OF  FIGURES  Figure  1.  Collecting  Figure  2.  Spore  types  recorded  from  the  sites  18  Figure  3.  Spore  types  recorded  from  the  sites  20  Figure  4.  Spore  types  recorded  from  the  sites  22  Figure  5.  Spore  types  recorded  from  the  sites  24  Figure  6.  Spore  types  recorded  from  the  sites  26  Figure  7.  Spore  types  recorded  from  the  sites  28  Figures  8,9.  Annual  Alternaria (Fig.  sp.  variation  (Fig.  8)  and  in  numbers  Double  of  spores  Crescent  of  Conidium 30  10,11.  Didymospore 12,13.  Large  Annual (Fig.  variation  10)  Annual  and  Ovate  variation  Phragmospore  (Fig. Figures  8  9)  Figures  Figures  Apparatus  in  (Fig.  in  numbers  of  Phragmospore numbers  12)  (Fig.  of  and  spores 11)  spores  H-Shaped  of . of  Spore  13)  35  14,15.  Lunate  33  Spore  Annual  variation  (Fig.  14)  and  in  number  of  Seimatosporium  sp.  spores  of  (Fig.  15) 37  Figures  16,17.  Fusarium  sp.  Annual (Fig.  variation 16)  and  in  number  Tripospermum  of  spp.  spores (Fig.  of 17)  40 Figures  18,19.  Titaea Figure  sp.  20.  Annual (Fig.  Annual  variation 18)  and  variation  in  number  Scolecospores in  number  of  of  spores  (Fig. spores  of  ?  19) of  43 Titaea  vi  sp  46  Figures  21  and  p r e c i p i t a t i o n , s i t e s . Figure  23.  Figure  22.  Figure  stemflow 22.  P r i n c i p a l  Monthly  21.  and  throughfall  mean  Components  Monthly  temperature  Analysis  incident  volumes at  Scatter  the Plot  at  the  sites  60 63  vii  ACKNOWLEDGEMENTS I both  would for  fungi.  his  I  Thesis  his  Bradfield,  for  of  of  this the  and his  data  thesis  used  and  in  u t i l i z e d  Research  Grant  my  s p i r i t u a l ) this  work;  for  also  l i k e  Dr.  G.  for  his  help  to  Data  in  A-801  over  the  design  Seifert  this  and  and  Dr.  Bandoni.  F i n a l l y ,  is  and  of  studies  my  dedicated  to  support  them.  E .  technical  rough  draft  Dave  Z i t t i n  computer  Materials  supported would  by like  (financial  allowed  me  of  advice G.  the  I  of  been  members  Dr.  a  to  were  have  with  read  me  study  helpful  thesis.  help  the  comments.  introduced  Bandoni,  two  his  research  years  to  other  for  helpful  J .  not  expertise,  Keith  of  me  would  Hughes,  many  R.  this  whose  thesis  thank  sampling  Centre  to  many  work  editing  with  Dr.  introducing  this  C.  provided  parents,  this  and  production  equipment  thank  supervisor,  manipulation.  Bio-Sciences  systems  my  assistance  Committee: fungi  of  thank  would  concerning  aspects  to  supervision  Without  possible. my  like  to  and NRC to and  complete  1  INTRODUCTION That both  the  leaves  p a r a s i t i c  time.  Plant  one  hundred  years,  studied  fungi  in  has  since  the the  that  that  populations  1 980)  as  in of  as  or  the  Some  a  by long  for  over  have  been  of  these  canopies  not  only  and  p r e c i p i t a t i o n , parameters  in  Table  terms  work  of  is  & Carroll  that 1977;  (Pseudotsuga  of  of  of  menziesii  r a i n f a l l  effects  • the  tree  and  canopy fluxes  suggesting  and  1980;  his  Carroll  (Mirb.)  of  organisms  nutrient  C a r r o l l  Carroll  plants  decomposers  the  stemflow/throughfall,  two,  of  documenting  measuring  the  I.  considered in  the  including  populations  in  on  stemflow,  communities  and  of  effect  modifications  canopies  The  the  of  profound  summarized  forest  Douglas-fir  a  rhizosphere  micro-organisms,  between  role  ecology  various  have  well  are  them.  the  between  modify  generally  (Bernstein  with  interface  horizons.  in  for  phylloplane  Yet, the  known  pathogens  the  and  inhabited  studied.  and  p r e c i p i t a t i o n  connections workers  have  of  are  been  leaf  throughfall  reported  exchange  live  between  s o i l  been  Workers  floor,  has  1950's.  modifications  cauloplane,  various  nutrient  the  receiving  forest  have  early  well  tree  studied  saprophytes  stemflow,  These  phylloplane,  in  been  a  fungi  populations,  and  communities  have  the  canopies,  throughfall r a i n f a l l .  not  of  saprophytic  and  canopy  Forest  on  and  branches  pathologists  widely  general,  and  coet  al  Franco)  in  Oregon. In  forests  throughfall  such  water,  as and  the  one  stemflow  studied water,  in  here, that  l i t t e r f a l l ,  order,  are  the  2  Table I. Chemical m o d i f i c a t i o n canopy.  of r a i n f a l l by the  forest  3  Values are i n percent c o n c e n t r a t i o n increase or decrease from i n c i d e n t r a i n f a l l . Element  Throuqhfall or Stemflow  Change  Reference  Na  throughfall throughfall throughfall  +81% + 111% +33%  C a r l i s l e e t a l 1966 Eaton e t a l 1963 S o l l i n s e_t a l 1980  K  throughfall stemflow throughfall throughfall throughfall stemflow  + 138% +1154% +993% +1625% + 189% +918%  B r i n s o n e t a l 1980 B r i n s o n e t a l 1980 C a r l i s l e e t a l 1966 S o l l i n s e t a l 1980 V o i g t 1960 V o i g t 1960  Mg  throughfall stemflow throughfall throughfall throughfall  +510% + 1362 + 133% +1422% + 180%  B r i n s o n e t a l 1980 B r i n s o n e_t a l 1980 C a r l i s l e e t a l 1966 S o l l i n s e t a l 1980 S o l l i n s e t a l 1980  ci  throughfall throughfall  + 179% + 18%  Eaton e t a l 1963 S o l l i n s e t a l 1980  H  throughfall throughfall  -85% -17%  Eaton e t a l 1963 S o l l i n s e t a l 1980  s  throughfall stemflow throughfall throughfall  +81% + 545% + 425% + 88%  B r i n s o n e t a l 1980 B r i n s o n e t a l 1980 Eaton e t a l 1963 S o l l i n s e_t a l 1980  p  throughfall • stemflow throughfall throughfall throughfall stemflow  + 233% +1483% + 251% + 883% + 267% + 567%  B r i n s o n e t a l 1980 B r i n s o n e t a l 1980 C a r l i s l e e t a l 1966 S o l l i n s e t a l 1980 V o i g t 1960 V o i g t 1960  N (inorg.)  throughfall throughfall throughfall  -30% + 173% -92%  C a r l i s l e e t a l 1966 Eaton e t a l 1963 S o l l i n s e t a l 1980  N (org.)  throughfall  +77%  C a r l i s l e e t a l 1966  C  throughfall stemflow  + 371% +2719%  B r i n s o n e t a l 1980 B r i n s o n e t a_l 1980  organic matter  throughfall  +400%  C a r l i s l e e t a l 1966  4  important is  means  because,  kg/ha,  of  when  the  incident  of  the  of  output  most  p r e c i p i t a t i o n allow  channelled  into  the  channelled  The  this  stations  required  the  approach  amount  of  leaving; kg/ha  the  of  kg/ha  values  multiplying flow  are  the  is  provide in the and  the  0.5-9.0%  the than  portion  Regression  p r e c i p i t a t i o n  in  and  of  small  stemflow. of  different on  throughfall  of  that forest  (Based  in  in  Though  stemflow  a  estimated  70-96%  in  1965).  stemflow  period  measuring  such  for  Figure  my  21).  In  p r e c i p i t a t i o n  p r e c i p i t a t i o n  concentrations  throughfall the  s i t e s ,  the-'nutrients  b i o l o g i c a l l y  water  or  for  reason  as  This  expressed  small.  only  stemflow  Patric  is  higher  amount  and  the  for  reduces  c o l l e c t i n g  The  the  f l o o r .  ends  is  up  as  1).  units  variable  1973).  of  and  stemfall  measure  much  ground  stemflow,  usual  r a i n f a l l ,  the  forests,  (Appendix  quite  throughfall,  deciduous  into  throughfall  in  sampling  forest  elements  is  are  and  1],  the  measured  throughfall  [Appendix  temperate  or  Helvey  to  stemflow  estimation  calculations  north  of  reaches  (eg.  over  of  elements  communities  sites  movement  p r e c i p i t a t i o n  equations is  the  contribution  concentrations in  element  kg/ha.  coefficient thus  a  of  reducing  measure  researchers entering  a  useful  a  poor  system.  concentration  who  the  unit  idea  is  of  for  These of  this  an  number  use  to  is  system,  and  t h i s . is  On  by  the  by  that  measure the  the  are  Kimmins is  hand,  happening  calculated  the  number  the  amount  other  actually  values  highly  sampling  (see used  it  in  of  sample  element  value  S t a t i s t i c a l l y ,  commonly  what  nutrients  variation  the  representative  kg/ha  d i v i d i n g  are  of  volume of  by of  hectares  5  covered  by t h e s y s t e m  fairly  good  throughfall, it  idea which  i s entirely  deposited  trees,  up  to  Table  I ) .  areas  of  more  20  (Helvey  elements  throughfall,  is  very  away  i s  that  the  Also,  values. but i s  bases  of  i n some  the  rough-  the concentrations  much  higher  those  of r a i n f a l l (see  than  stemflow  than  or  area,  area,  low  than  creates  kg/ha  in  localized  n u t r i e n t and i o n c o n c e n t r a t i o n ,  biologically  a  rainfall  study  the trunk  1965).  higher  to  the stemflow  around from  provides  on t h e s t u d y  in a  are in turn  a r e much  high  significant  randomly  & Patric  The n e t r e s u l t  evenly  band  cm  this  happening  f o r measuring  i n stemflow  which  While  approximately  i n a narrow  about  species  most  fall  what  i s not deposited  instead  of  of  studied.  inappropriate  Stemflow  barked  being  which a r e  values  might  suggest. Another  possible role  throughfall, terrestrial  i s  (1981)  maple  (Acer He  (Arnold)  stemflow.  The  (Thuja  macrophyllum found  Ceratosporella,  of  Bandoni piicata  Gyoerffyella  also  study  stemflow  Pursh)  flow,  and, t o a  microbial  present  innoculum  f o rthe  was  initiated  to  by  The  Bandoni  i n v e s t i g a t e d the stemflow  and  Douglas-fir,  red  alder  Gyoerffyella  and G y o e r f f y e l l a  Ceratosporium,  extent,  studied  gemellipara  reported  lesser  and t h r o u g h f a l l .  been  Donn.),  the hyphomycetes  in alder He  of  has p r e v i o u s l y  (1976).  cedar  Ingold,  sp.  source  mycobiota  of stemflow  red  Bong).  Titaea  the  and Gonczol  western  a  ecosystem.  investigate mycobiota  as  f o r stemflow  species  big-leaf  (Alnus  rubra  biappendiculata  Marvanova, gemellipara of  of  and  ?  i n maple  Tripospermum,  Titaea, Cornutispora,  and s e v e r a l  6  unidentified conidia (1976)  reported  conidium,  from stem flow of v a r i o u s  the occurrence  Titaea  Cornutispora  sp. ,  lichenicola  of  a  trees.  Gonczol  dematiaceous  T-shaped  fragile  Harkness,  Dicranidion Hawksw. , ?  T i t a e a sp. , and c o n i d i a  t e n t a t i v e l y i d e n t i f i e d as those of a N e c t r i a stemflow  gathered a t the base of a beech  For  the present  each of red a l d e r Michx.) and species.  the  and  I sampled stemflow  trembling  throughfall  aspen  beneath  two  B r i t i s h Columbia,  species  accounted  foam  of  tree. from two t r e e s  (Populus  tremuloides  the canopies  Hardwoods are common c o n s t i t u e n t s of  i n southwestern these  study  sp. from  of  both  disturbed  areas  and i n the area that I s t u d i e d  f o r almost a l l of the arborescent  spec i e s . The first  study was i n i t i a t e d with s e v e r a l goals  The  was t o r e c o r d the types of f u n g a l propagules that occur i n  stem flow and, where p o s s i b l e , i d e n t i f y them. describe  the  distribution  of these c o n i d i a something the  i n mind.  data  i n an  (temporal and s p a t i a l ) and abundance  attempt  to  suggest  their,  of t h e i r e c o l o g i c a l c h a r a c t e r i s t i c s . gathered  t h r o u g h f a l l mycobiota ecosystem.  The second was to  might  suggest  might  play  in  roles  source  Finally,  that  I hoped  stemflow  the p o p l a r - a l d e r  and  and  forest  7  MATERIALS AND Five British  s a m p l e s t a t i o n s were e s t a b l i s h e d on Columbia  Endowment  Lands  i n t e r s e c t i o n o f an  unnamed c r e e k  site  directly  is  located  University H i l l The to  Elementary  following  old.  It  Teversham 1977). species  provide  contains  salmonberry  (Vaccinium  (Pteridium  aquilinum  species  one  were  and  100%  Smith)  the  Boulevard.  The from  t r e m b l i n g aspen  early  successional  until early  1936  other  Presl.)  ( L . ) K u h n ) , and  red  two  understory  sword  bracken  ferns ferns  numerous s p e c i e s o f  ages  herbs.  t h e two  dominant  of  t r e e s were  the  &  huckleberry  shrubs, and  l a r g e r t r e e of e a c h of Actual  these  The  up  stage  (Klassen  autumn,  canopy c o v e r .  and  (Kaulf)  selected.  of  Boulevard  (Rubus s p e c t a b i l i s P u r s h ) ,  muni turn  s m a l l and  an  continued  than  parvifolium  (Polystichum  Chancellor  r e d a l d e r and  summer  greater  south  School.  which  During  immediately  across  represents  logging  t h e U n i v e r s i t y of  with Chancellor  f o r e s t i s d o m i n a t e d by  45 y e a r s  One  METHODS  tree not  determined. C o l l e c t i n g devices were  mounted  plastic  tubing  around  the  approximately supported the of 1).  on  the  split tree  similar  test  to those  trees.  u s e d by  A l e n g t h of  (1981)  inch diameter  was  tacked  in  a  angled  at  30 d e g r e e s .  The  tubing emptied i n t o a p l a s t i c  bag  i n an  aluminum can  n a i l e d t o the  t u b i n g was  fastened  T h r o u g h f a l l was  to the  spiral  once  trunk  flow of water from the t r e e i n t o the the  1/2  down t h e m i d d l e l e n g t h w i s e descending  Bandoni  tree.  increase  t u b i n g , t h e u p p e r edge  tree with ducting  sampled at a f i f t h  To  station  tape  (Figure  l o c a t e d under  a  8  Figure  1.  Collecting  Apparatus.  9  10  m i x e d c a n o p y of (10  cm. wide  trees.  the  at  A short  plastic  bag,  two d o m i n a n t  the  mouth)  length  of  was  tree  suspended  tubing  supported  in  species.  led  an  A plastic  with wires  from t h e  aluminum  funnel  between  funnel  neck  can hanging  two  to  a  from  the  wires. It the to  was  not  plastic  thought  bags due t o  an e l e m e n t a r y  as  soon  as  collecting  possible  to  the  Forty-one  of  the  without with  v/v  95% e t h y l  extensive  The d e n s i t y  in top  four  160 m l .  of  of of  in  Company M o d e l  a  preservatives  the  stations  were  collected  samples  place,  in  sampling  and  the  into  bottles  distortion higher conidia  material  preserved  20 ml i n e a c h t u b e .  of  sample  m a t e r i a l to  soon  added to  of  after  each  alcohol in  retrieval.  bottle  proved  germination, occurred  concentrations.  the  samples  was  was  required.  stemflow  so  low  From e a c h  o r t h r o u g h f a l l was  Thirty  centrifuge  minutes  95% of  the  concentrate  that sample placed  centrifugation  (International  Two c e n t r i f u g a t i o n 40 ml of  1980  collection,  spore morphology t h a t  CL) removed more t h a n  top  each  and i n h i b i t i n g s p o r e  samples  clinical  i n examining  alcohol  organisms  the  made between November 6,  the  t u b e s and c e n t r i f u g e d .  speed  any  The bags were e m p t i e d  in  involved  to p r e s e r v e  some s p e c i e s a t  bottle  put  (Table time  in k i l l i n g  concentration  rain.  II).  Because  effective  after  were  1981  Five percent  Consequently,  collections  necessary  put  lab.  3,  was  to  p r o x i m i t y of  new bags  and November  it  the  school.  bottles,  transported  desirable  conidia  steps  at  Equipment from  reduced  the  160 ml  with a r e s u l t i n g  4X  11  Table  II.  C o l l e c t i o n numbers a n d d a t e s .  Number 1 2 3 4 5  Date 6 November 1980 7 November 1980 8 November 1980 9 November 1980 10 November 1980 18 November 1980 19 November 1980 20 November 1980 27 November 1980 29 November 1980 1 December 1980 2 December 1980 10 December 1980 11 December 1980 14 December 1980 15 December 1980 22 December 1980 27 December 1980 9 J a n u a r y 1981 20 J a n u a r y 1981 24 J a n u a r y 1981  6  7 8 9 10 1 1 12 13 14 15 16 17 18 19 20 21  increase  in c o n i d i a l  glass  t h e c o n c e n t r a t e were p i p e t t e d o n t o  s l i d e , a d r o p of  glycerine  p l a c e d pn t o p a n d f a s t e n e d i n slide Site  was (No.  place  with 1000X  (No.  a  slip  nail  polish.  Each  1-41)  and C o l l e c t i o n  1-5).  i l l u m i n a t i o n on a  an O r t h o m a t were  of  when  Leitz  Wetzler  necessary,  conidia.  that,  at the m a g n i f i c a t i o n  three  "sample sweeps" of Four  100X m a g n i f i c a t i o n u s i n g p h a s e -  camera a t t a c h m e n t .  used,  photographing  sampling.  a d d e d , and a c o v e r  with  coded w i t h C o l l e c t i o n Date  The s l i d e s were e x a m i n e d a t contrast  Date 27 J a n u a r y 1981 30 J a n u a r y 1981 12 F e b r u a r y 1981 14 F e b r u a r y 1981 16 F e b r u a r y 1981 18 F e b r u a r y 1981 26 F e b r u a r y 1981 3 M a r c h 1981 8 M a r c h 1981 25 M a r c h 1981 29 M a r c h 1981 1 A p r i l 1981 8 A p r i l 1981 8 May 1981 12 J u n e 1981 20 J u l y 1981 5 S e p t e m b e r 1981 28 S e p t e m b e r 1981 5 O c t o b e r 1981 3 November 1981  density..  Two 0 . 1 m l s a m p l e s of clean  Number 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41  or  five  A  compound  microscope  M a g n i f i c a t i o n s of for  450X and  identification  minimum a r e a c a l c u l a t i o n  (100X)  the cover  necessary  to spot  s l i p allowed  sweeps d i d n o t  and showed  the c o n i d i a ,  most  efficient  s i g n i f i c a n t l y change  the  12  number  of  three  sweeps  coverage  species  of  The  as  descriptive genus  Measures into  conidia  for  18%  give  ml,  conidia  of  preserved November Three from  ml  in  slides, and  of  the  was  by  1981 to  interface at  see  of to  was  and  re-  modified  were  drawings  of  by  identified  were  made  for  conidia  in  the to  only  was  collected  January  was  and  fungi. a  0.1  ml  of  the  of of  conidia  number  be  confirmed  not  possible  from  the  February  Conidia fine  of  from  were  from in  1982. removed  under  with  the  by  sites of  needle  magnification  water  of  reciprocal  malt-yeast-peptone 2);  number  number give  of  sample.  this  80X  The the  give  were  concentration  by  0.25  using  Appendix  conidia  actual  As  isolate  to  (MYPT;  and  could  and  transferred  tetracycline  total  morphological  and  spores  the  by  fungi  or  and  a  sweep  Century  (unconcentrated)  32X  illumination,  19th  throughfall.  10  material  used  microscope  s t r i c t l y  multiplied  cultures.  of  of  number  actual  air-water  dissecting  s l i p  the  were  for  each  (1979), the  and  m u l t i p l i e d  December  methods the  stemflow  fresh  on  abundance  estimate  m u l t i p l i e d  pure  the  Raj  possible,  total  then  using  in  relative  cover  the  identity  isolation  in  mm.  The  case.  rough  type  and  The  a  then  per  each  square  type  Photographs  the  one  concentrate, per  of  c o n i d i a l  to  in  Nag  abundance.  s l i p .  recorded  &  Where  19.8  cover  Saccardo  species.  reference  converted  by  Kendrick  or  relative  c o n i d i a l  were  terms.  their  each  each  defined  by  or  approximately of  of  Conidia  defined  each  18%  abundance  categories  future  covered  about  recorded.  to  recorded  a  incident  agar samples  with was  13  spread  on  water  then  blocks  using  a  of  cutting  Mechanical were  onto 2)  were  medium  induces  transferred at  favour  growth  lower  arm  designed  by  and  the  discourage  to  to  MYPT;  The  and  microscope  sample  streaks  (SORB-T;  see  tetracycline-enhanced  b a c t e r i a l thus  and  Botany/Zoology  tetracycline  and  dry,  dissecting  U.B.C.  germinate.  formation  failed  oat  agar  15  and  of  to  a  the  with  allowed  growth;  prevents  the  sorbose  'weed  fungi'  medium.  that  5,  agar  2),  under  transferred  to  colony  to  grown  at  out  employed  Isolates  Appendix  cut  allowed  overgrowing  see  were  sorbose  and  media  (WA;  agar  Shop,  made  Appendix  from  agar  (OA; 20  aquatic  to  sporulate  see  Appendix  C.  ;  the  low  Hyphomycetes,  on 2)  these  or  media  WA.  Fungi  temperature  which  were  was  generally  were  used  grow  to  well  temperatures.  Fungi  isolated  and  c o l l e c t i o n  numbers  Individual  collection  identified  and  placed  were in  numbers  are  given  the  J . A .  MacKinnon  UBC C u l t u r e  noted  C o l l e c t i o n .  in  the  abundance  were  culture  descriptions. Fungal  spore  characterize  each  total  205  of  d i v e r s i t y  Collection  and  Site  Date-Sites  on  (5  each  Collection  Sites  on  41  used  to  Date.  A  Dates)  were  characterized. These Components abundance analysis amounts  205  Analysis  axes  variation  were  (PCA),  characterizing with  of  Date-Sites  the  with  fungal  samples.  drawn in  included  the  A  spore PCA i s  representing data  in  a  Principal  d i s t r i b u t i o n a  multivariate  successively  matrix.  The  and  f i r s t  lesser axis  14'  accounts for a  the  for  the  second  largest  largest  multi-dimensional  reflects  their  Computer (MTS),  (an  Research  amount,  the  470v/6 MIDAS  Laboratory  of  in  v a r i a t i o n ,  PCA  II),  program  was  run  d i s t r i b u t e d of  second  and  proximity  Michigan  University  the  stepwise  which  The  Model  the  of  continuing  ordination  s i m i l a r i t y .  Amdahl  using  amount  on  producing of  points  the  U.B.C.  Terminal by  Michigan  the  axis  System  S t a t i s t i c a l  (Fox  &  Guire  1976). The, with  C o l l e c t i o n  different types were  matrix  and  spore a l l  unsealed.  for  this  program  Date-Sites types  Date-Sites  is  cases-variable's  representing  representing were  a  used  in  the the  the  cases  matrix, and  variables.  A l l  ordination:  the  the spore data  15  RESULTS Conidia in  sample  sweeps  different in  6  recorded  b-g  in  for  as  absence  from  and  were  plates  were  Leucosporidium that  branched  i n t h e samples  formed  short  such  Alternaria  commonly  such  as  and  Large  from  presence  numbers  of  i n the genera anamorph  showed  up  my  yeasts,  and  several commonly  (Figure 6h).  p r e s e n t were h y p h o m y c e t e s . saprobes  Others  many  plates;  ("mirror")  that  an  a l l of  Finally,  1981),  and Mucor  of  48);  on  of  slides.  PeniciIlium  from a m e r o s p o r e s .  their  h e r e due t o  preserved  ( J . A. M a c K i n n o n  and  fungi"  not imply  c h a i n s (pseudomycelium)  seen  or Fusarium.  Imperfecti, "lower  does  e t a l (Summerbell  of the o t h e r f u n g i  hyphomycetes,  them  fashion  not  to  were n o t i n c l u d e d  sporobolomycetaceous Fell  were  designed  of  Ptychogaster  derived  in a multiradiate  Most were  the  scott i i  they  were  samples  a s Zygomycetes  Basidiomycete  included  used  throughfall.  the  t y p e s shown  t h e y o c c u r r e d o u t s i d e of t h e  from my  which  as w e l l  presumably  y e a s t s were seen  yeasts  and  and  several  though  methods  identifying  species,  representing  Chytridiomycetes  revealed  Aphyllophoraleous  these  because  observed  the c o n i d i a l  observed,  present, of  slips,  t a x a were  Basidiomycetes, Fungi  and  stemflow  Rhizopus,  these  also  fungal  As w e l l ,  The i s o l a t i o n  difficulty  Aspergillus  groups.  were  Oomycetes  Streaked  cover  and t h e a b s e n c e  amerospores the  the  Ascomycetes,  Zygomycetes, such  37 d i f f e r e n t  the d a t a base  sweeps.  select  of  taxonomic  Figure  sample  of a t l e a s t  were  or p a r a s i t e s  a s s p e c i e s of  representative  Flabellospora,  Titaea,  Several  "aquatic"  Clavariopsis,  16  Gyoerffyella,  Tetracladium,  Dendrospora.  Of  Tripospermum  have  hyphomycetes  (Ingold  growing these  or  situat  these,  1975c),  have  been  It  not  unusual  throughfall. as  they  spring  to  Both  reach  peaks  when  the  Bacteria  were  abundant  Numerous  protozoans  identify  the  d i s t i n c t i v e only were more  on  than  date  only.  7)  the  Taxa  reported  in  or  a l l  of  semi-aquatic  in  on  rather  sporulation  during  without  also  observed.  No  attempt  the  bacteria.  Desmids  are  types  dates,  those  five  are  2-6)  Records  more of  and  on  or  to  recorded ten  dates, on  one  (Figure  graphs  dates  spore  and  commonly  drawings  Sporulation  these  how  to  highly  observed,  those  ten  with  made  the  recorded  line  than  was  of  six  those  plates  tetracycline.  of  into  recorded and  winter  overgrown  algae  indication  provided. on  only  divided  dates,  (Figures  recorded  8-20).  an  or  leaves,  Streaked  with  the  stemflow  the  supplemented  were  recorded.  than  soon  spore  types  in  species  leaves.  samples.  material  As  Photographs  (Figures  wood  on  a  were  were  or  to  1ichenicola genera  occasions.  two  types  of  a l l  Micrasterias  two  Seimatosporium,  these  are  were  were  ten  spore  spore  provided  media  recorded,  recorded  of  unpreserved  genus  of  grow  trees  protozoans  those  those  species  months  unless  of  aquatic  been  t e r r e s t r i a l  Cornutispora  find  their  bacteria  on  species  apparently  from  they  a  and  prepared  these  never  as  and  ions.  Pestalotiopsis, is  considered  streams.  in  and  Lateriramulosa  have  in  collected  Lateriramulosa,  of  been  but  submerged  Three, coelomycetes, of  species  t r a d i t i o n a l l y  sporulating  genera  Tricladium,  are  types  for also from  17  other  authors are noted.  noted  f o r those  J . A. MacKinnon c o l l e c t i o n  species in culture  Spore  types  numbers a r e  (U.B.C. C u l t u r e C o l l e c t i o n ) .  r e c o r d e d on more t h a n  ten dates  A l t e r n a r i a sp. This  spore  type  peak  i n the f a l l ,  was  recorded  common p l a n t quite  aid  a n d a l a r g e peak  from  parasites,  dispersal  sp.  from  Double Crescent This  2 f , g ) from  This  spore  type occurs  The  F i g u r e 2g.  (1974)  on w a t e r  foam.  8).  It  are f a i r l y  dictyospores are  mentioned  (see D i s c u s s i o n ) .  spore  the stemflow  February.  o c c u r s on p o p l a r s . genus.  (Figure  dematiaceous  and f l o a t  stream  i s t h e same  and  i n January  that  films,  Dudka  spores of which  may  (1965) r e c o r d e d  F i g u r e s 2h,8.  Conidium  Fig.  January  Bandoni  the year, with a small  S p e c i e s of A l t e r n a r i a  and t h e i r  a r e hydrophobic  in their  Alternar ia  in  a l l sites.  distinctive.  A l t e r n a r ia  occurs throughout  type  recorded  of b i g l e a f  from November t o It  by  Bandoni  maple and D o u g l a s - f i r . March,  with  the conidia  o f any  end o f one o f t h e c r e s c e n t s may s p l i t , Figures  peaks  i s most common on a l d e r s ,  I t i s hot l i k e  2d,e,f,g,9.  (1981  in  but a l s o described  a s i s shown  18  Figure  2. Spore types recorded from the s i t e s , a) Large P h r a g m o s p o r e b) O v a t e P h r a g m o s p o r e c ) H - S h a p e d S p o r e dg) D o u b l e C r e s c e n t C o n i d i u m h) A l t e r n a r i a sp. . A l l b a r s = 20 n.  19  20  Figure  3. S p o r e t y p e s r e c o r d e d from t h e s i t e s , a) L u n a t e S p o r e b) F u s a r i u m s p . c) Seimatosporium sp. d-f) Tripospermum spp. g) ? T i t a e a sp. HT S c o l e c o s p o r e s . A l l b a r s = 20 ?.  22  Figure  4. Spore types recorded from the s i t e s , a) Clavariopsis aquatica b) Stauronema sp. c) C e r a t o s p o r i u m s p . d) C e r a t o s p o r e l l a s p . e) Tapered Helicospore f) Flabellospora sp. g) H e l i c o m a sp. h) Titaea sp. . A l l bars = 20 v, e x c e p t a) T ~ b a r = 200  a.  24  Figure  5. Spore types recorded from the s i t e s , a) D e m a t i a c e o u s C r u c i f o r m C o n i d i u m b) D e n d r o s p o r a sp. c,d) Tricladium sp. e,f) Gyoerffyella gemellipara g) D e m a t i a c e o u s T - S h a p e d C o n i d i u m h) T i n y T r i r a d i a t e Conidium. A l l b a r s = 2 0 », except g,h) , bars = 10 » .  25  26  Figure  6. Spore t y p e s r e c o r d e d Lateriramulosa uni-inflata ? Speiropsis irregularis a c u m i n a t a I n g o l d e) S e c o n d Unknown h) B r a n c h e d Y e a s t .  from t h e s i t e s , a) b) ? C e r a t o s p o r e l l a sp. c) P e t e r s e n d) A l a t o s p o r a H-Shaped S p o r e f ) Unknown g) A l l b a r s = 20JI.-  28  Figure  7. S p o r e t y p e s r e c o r d e d from t h e s i t e s . A) D i d y m o s p o r e XI00 b) P e s t a l o t i o p s i s s p . X320 c ) S c o l e c o b a s i d i u m s p . X300 d) G y o e r f ' f y e l l a b i a p p e n d i c u l a t a X220 e) T e t r a c T a d i u m s p . X70 f ) T r i p o s p o r i n a y a k u e n s i s X30 g) V a r i c o s p o r i u m e l o d e a e X120 h) B r a n c h e d C o n i d i u m X10 i ) T e t r a p l o a a r i s t a t a X240 j ) C o r n u t i s p o r a 1 i c h e n i c o l a X100 k) P i c r a n i d i o n f r a g i l e X75.  30  F i g u r e s 8,9. Annual v a r i a t i o n i n numbers of s p o r e s of A l t e r n a r i a s p . ( F i g . 8) and Double C r e s c e n t Conidium ( F i g . 9) T (month 0 = November, 1980). Collecting S i t e s : "o"=small a l d e r "•"=large a l d e r " A = s m a l l p o p l a r " A " = l a r g e p o p l a r "«"=throughfall. n  31  FIGURE 8 30-  25-  20—  15—  o  03  "D •  c o  10-  MM  i 5-  :  0-  I  1  N  S  I  S  I  month ^FIGURE 9 o 150H  IOOH •  03 •  MM  •  MM  o  A  5(Ho  Al  o  O  u  P N  A  o c»  « A  CD  •  vi  month  32  Didymospore There  were s e v e r a l  category.  Ovate  They o c c u r r e d y e a r  didymospores  round  included  at a l l s i t e s .  in  this  F i g u r e s 7,J_0. «  Phragmospore This  commonly  Large  spore  type  occurred  year  i n t h e w i n t e r and s p r i n g .  round  at a l l sites,  more  F i g u r e s 2b,11.  Phragmospore This  was  different  spore  t y p e was v e r y common i n t h e s p r i n g  not recorded  species  of  i n t h e summer.  Fusarium.  It  may  perhaps  and f a l l , but represent  I t was r e c o r d e d from a l l s i t e s .  a  Figures  2a,12 .  H-Shaped  Spore  This to  spore type  February.  stemflow  Lunate  Bandoni  has  found  from  this  November spore  in  F i g u r e s 2c,13.  Spore spore  t y p e was r e c o r d e d y e a r  i n the winter.  ( p e r s . comm.) Figures  common a t a l l s i t e s  ( p e r s . comm.)  of Douglas-fir.  This common  i sfairly  has  round,  a n d was  I t was . r e c o r d e d from a l l s i t e s .  found  this  spore  i n stemflow  specially Bandoni  of D o u g l a s - f i r .  3a, 14.  Seimatosporium sp. Spores  of t h i s  genus were r e c o r d e d y e a r  round  except  for  33  Figures 10,11. Annual v a r i a t i o n in numbers of spores of Didymospore (Fig. 10) and Ovate Phragmospore (Fig. 11) (month 0 = November, 1980). Collecting Sites: O"=small a l d e r "•'^large alder "A"=small poplar "A"=large poplar "•"=throughfall. n  .  ^FIGURE 10 30 25H  2015— 105—  A J" J* A.J month  I A  FIGURE 11 302520—  O A  15O  A A  10pO^Oi  5H  A J  O A«& O)  A  A month i!i  A A\  J  A  TT  35  Figures 12,13. Annual variation i n numbers of spores of Large Phragmospore (Fig. 12) and H-Shaped Spore (Fig. 13) . (month 0 = November, 1980). Collecting Sites: "o"=small a l d e r "•"=large alder "A"=small poplar "A"=large poplar "•'^throughfall.  100—r—  FIGURE 12  80H  6(H 4(H  A A A  20H  O4i «» •  GQSD  rt  •  o  A»**»<&  I"  Ji  month  •  ]  A  s rt"  _ FIGURE 13 sols— 20 15-  ion 5H  rt  ]  Ui Hi ~ ~T month  s  iJ"  37  Figures 14,15. Annual v a r i a t i o n i n number of spores of Lunate Spore (Fig. 14) and Seimatosporium sp. (Fig. (month 0 = November, 1980). C o l l e c t i n g Sites: "0"=small alder "•'^large alder "A"=small poplar "A"=large poplar B"=throughfall. n  15)  38  FIGURE 14  lOOOH  •TH  500H  a o  o  o  J  M  j  M  m o n t h  _ FIGURE 15 o  150—I  100-H  A  •H  5<H  AA •  o  o QAO»  • mo  A  # a&mt& o N  -i  •  S  J i  o  month  ]~  39  March  and  April,  January.  and  were e s p e c i a l l y  They were r e c o r d e d  from  a l l sites.  genus i n c l u d e s p a r a s i t e s of Rosa  , Vitis  1964),  among  not  Populus  .  Fusarium  others,  Figure  were  (1973)  spores  most  were v e r y  common  recorded  Beck, E n g l a n d , spring,  and  One  but  and  (Shoemaker  from  Alnus  or  autumn t o w i n t e r .  plant  Jones  beech  parasites.  wood  &  a  in  the  Smooth  winter  (1964) r e c o r d e d stream.  this  Archer  as common i n foam of  Oliver in  sites,  Members of  Willoughby  particularly and  a l l  and  Fusarium  Figures  3a,16.  spp. these  spore  undescribed.  types  ( F i g . 3f)  They were not  were v e r y common Bandoni  Tripospermum  from  & Archer  camelopardus as spring.  decaying  from  90,91.  a l l sites.  Willoughby T.  from  round,  Gareth  of  t o be  months, from  coelomycete  Cornus  common a l l y e a r ,  c a m e l o p a r d u s I n g o l d , Dann & M c D o u g a l l ;  appear  of  year  Mackinnon  Tripospermum  T.  , and  been r e c o r d e d  s p e c i e s of F u s a r i u m  as c o l o n i z i n g  J . A.  This  to  3c,15.  genus a r e v e r y common  spp.  has  November  sp.  Fusarium and  but  common from  leaves  (Figs  to  3d,e) summer  i n t h e autumn, w i n t e r , and  spring,  (1981, F i g . 2d)  recorded  also  s t e m f l o w o f maple and  r a r e i n stream &  others  close  i n the  (1973) r e c o r d e d  Hudson  the  is  Sutton  in a t e r r e s t r i a l  T.  foam  myrti  from  street  (Lind)  i n England,  (1964) site.  recorded  fact,  spora  Hughes i n the  r e p o r t e d T. In  species .  and  winter  myrti  from  members of  this  40  F i g u r e s 16,17. A n n u a l v a r i a t i o n i n number o f s p o r e s o f Fusarium sp. ( F i g . 16) a n d T r i p o s p e r m u m s p p . ( F i g . ~ (month 0 = November, 1 980) . Collecting Sites: " 0 " = s m a l l a l d e r "#"=large a l d e r A " = s m a l l p o p l a r "A"=large poplar "a"=throughfall. N  41  FIGURE 16 600H  o  40(H  o  •iH  /  200H  o  O  a o a  N  month FIGURE 17  30H 25H  20H 15H •rH  o  AO  A  io-  ••  5H  N  month  42  genus have "aquatic leaves stream  not  submerged foam,  i n a stream. of  origin.  Vancouver,  in  as  habitat,  spores  Tripospermum  ?  reported  hyphomycete"  terrestrial  (T.  been  Canada.  that  this  flow  in  genus  Figures  almost  (1981)  from  a  typical  i s , g r o w i n g on d e c a y i n g  Though o c c a s i o n a l l y  Bandoni  water  sporulating  also  collected  certainly found  in  have a  conidia  of  the s u r f a c e of a paved lane i n  3d,e, f_,J_7.  J . A. MacKinnon  74  camelopardus).  T i t a e a sp. Bandoni  (1981  F i g . 2c)  recorded  spores  of t h i s  type  from  stemflow of r e d a l d e r as V o l u c r i s p o r a sp.  I t i s a l s o one o f t h e  spore  stream  types  represent Fig.  the  14).  I  throughout bears  figured  described  spore  figured  recorded  the  some  by I n g o l d  this  winter  by G o n c z o l spore  & Toth  type,  foam, and  chickweed  to  Titaea  leaves.  always  on  o r n i thomorpha  Figures  may  (1974 T a b l e I I , alder,  and s p r i n g a l w a y s a t low f r e q u e n c y .  resemblance  from  (1975b) from  It  Trotter,  3g,18.  Scolecospores This category They  are,  represent  for  spring  or  occurring  the  propagules  Scolecospores frequency  r e p r e s e n t s a l l of the s c o l e c o s p o r e s  occur  i n the f a l l summer. spore  type  most  part,  of  several  in  low  indistinguishable different  frequency  and w i n t e r ,  and p r o b a b l y  fungal  in early  f o r the winter  were,  the  species.  autumn,  and were n o t r e c o r d e d  Scolecospores  recorded.  higher i n the  most f r e q u e n t l y  and s p r i n g months.  Figures  43  Figures 18,19. Annual v a r i a t i o n i n number of s p o r e s of ? Titaea sp. ( F i g . 18) a n d S c o l e c o s p o r e s ( F i g . 19) . (month 0 = November, 1980). Collecting Sites: O" = s m a l l a l d e r " • "=l a r g e alder "A*' =small poplar "A =large poplar "B"=throughfall. n  n  44  FIGURE 18 30-H 252015•H  10-  •H  o o  ••CO  •  •  o  5-1  rt  T  ii. J .!  m o n t h  £LGURE 19 3000H  2000-H  c3 1000-  O  o  •• AA  4& rt  M  ji r  m o n t h  F~T  45  3h,J_9.  Titaea  sp.  This  genus has a l s o been  maple  (Bandoni  3,4).  I t was  mostly  on  small  This  spore  and  (Gonczol from  the  1976 fall  but a l s o of  T.  on  of  Plate to  bifgleaf I I I , Figs  the  spring,  the a l d e r s .  triradiata  This  Hansford.  both  submerged  ,  was  Ingold  This  but  It  was  resemble  appears  always  any  wood  (1942,  1965,  Ingold  1) n o t e d i n Great  in  identical  that  a  (1975a, he  &  the  Archer and  Ingold &  also Ellis  F i g . 2-12,  commonly  Britain.  present  . to  stream,  1967),  t o M a r c h on  described  d e s c r i b e d by W i l l o u g h b y  (1960).  foam s a m p l e s  Helicoma  not  Fraxinus  13 s p .  dates  r e c o r d e d from December  poplars.  does  Flabellospora development  s i x to ten  sp.  It  Nilsson  t y p e s r e c o r d e d on  s p o r e t y p e was  numbers.  Fig.  the  stemflow  4h,20.  Flabellospora  (1968),  beech  t r e m b l i n g aspen  resembles  Spore  alders  and  r e c o r d e d many t i m e s  spore c l o s e l y Figures  1981)  r e p o r t e d from  found  both  in small  species  of  s p e c i e s whose (1969) by  from  Willoughby  (1952),  F i g . 11;  and  1975b,  this  conidium  in  t o be  identical  to  Figure 4f.  sp. dematiaceous  helicospore  appears  46  Figure  20. Annual variation i n number of s p o r e s of Titaea sp. . (month 0 = November, 1980). Collecting Sites: O"=small a l d e r "•"=large alder A =small poplar "A"=large poplar " "=throughfall. n  n  B  n  JIGURE 20  100  80A  A  60H  A  4(H  A  A  2(H  A •  A  £9*Q A ^ A  m o n t h  48  the  one  recorded  litter and  of  by  Ceratosporium This  both  1)  five  sites,  from t h e  recorded  Figure  terrestrial  only  i n November  4q.  Matsushima  type  has  been  collected  from D o u g l a s - f i r t h r o u g h f a l l . in winter-spring.  by  I t was  Figure  Carroll  recorded  (1981, at  all  4c.  Helicospore  This stemflow  is  undoubtedly  the  of D o u g l a s - f i r and  2h),  and  in  on  stream  sp.  He1icornyces.  This helicospore  the  I t may  s p r i n g , on  same  species  as  stems of J u n c u s by  foam  Helicosporium  through  F i g . 1g)  I t was  alders.  cornutum  spore  Fig.  Fig.  (1981  Scirpus microcarpus.  December, on  Tapered  Bandoni  by  represent  poplars.  Bandoni  (1965  in  (1981  P l a t e 2j)  as  a s p e c i e s of H e l i c o s p o r i u m  or  was  Tubaki  collected  recorded  Figure  from  the  autumn  4e.  r Pestalotiopsis Spores fall  and  from  recorded small  illustrated Britain  of  this  winter,  conidia/ml. was  sp.  in  usually one  from  which  of  low  may  the  December.  unidentified also  a very  Pestalotiopsis  sample  the  with  32  i n December.  It  exception  Ingold  spore  from  belong similar are  during  concentrations  l a r g e a l d e r stemflow  in  ( p e r s . comm.) f o u n d Species  in  genus were r e c o r d e d  large a l d e r , with  alder an  coelomycete  in'" t h i s  very  of  (1975a,  stream  spore  but  in  common  one  spore  F i g . 39-14)  foam genus. street  in  Great Bandoni  spora  plant parasites.  49  Figure  7b.  J . A. M a c K i n n o n 55.  Scolecobasidium sp. This  s p o r e t y p e h a s been  f r o m washed S c i r p u s up  in  eight  recorded  microcarpus  litter.  o f my c o l l e c t i o n s .  was most common on l a r g e a l d e r .  Spore types  Gyoerffyella  November  to  throughfall. stemflow a  January, Bandoni  in  s a p r o b e on c o n i f e r wood  showed  on  each  This  (Webster  dates  Ingold  stemflow  trees.  type  a t a l l s i t e s , but  on two t o f i v e  of of  (1981) r e p o r t e d  of r e d a l d e r  spore  I t occurred  (Arnold)  One s p e c i m e n was r e c o r d e d  This  (1981 F i g . 1b)  Figure 7f.  recorded  biappendiculata  by B a n d o n i  three large  this  dates, alder  species,  and  also  species  h a s been  & Descals  1982).  from in  in  the  reported Figure  as  7d.  C e r a t o s p o r e l l a sp. Three September  s i n g l e spores of t h i s and  poplar.  once  Bandoni  stemflow  type  were  recorded,  i n J a n u a r y , on b o t h a l d e r s  (1981)  o f an u n s p e c i f i e d  reported tree  this  species.  twice  in  a n d on t h e l a r g e  same Figure  spore  type  in  4c.  Stauronema s p . These occurs  spores  represent  i n stemflow,and  has  another also  been  coelomycete recorded  genus from  that  streams  50  (Ingold an  1974?  I n g o l d 1975b F i g . 39,10; I n g o l d  unknown c o n i d i u m .  from  Five  the large alder  spores of t h i s  and b o t h p o p l a r s ,  1977 F i g . 1c) a s  s p e c i e s were  in  winter  recorded,  and  spring.  F i g u r e 4b.  Clavariopsis This streams,  species  by Dyko  1964). both  times  common  in  fungus  collecting  a  terrestrial  (Webster  on  & Descals  (Gareth  a n d on  and  Oliver  Jones  winter  and  & Archer spring  angiosperm  1982),  i n November a n d once  Willoughby  litter  i n January,  (1973)  recorded  i n stream  foam i n  Conidium  was r e p r e s e n t e d by one s p o r e  dates.  It  poplars,  and t h r o u g h f a l l .  Gonczol  (1976  Pi  Gyoerffyella  was  recorded This  I I I , Figs  (1975 F i g 27) from  (Bandoni  as  sporulating in  F i g u r e 4a.  This  This  reported  I t occurs s a p r o b i c a l l y  i n streams  alder.  D e m a t i a c e o u s T-Shaped  Miura  reported  r e c o r d e d once  on l a r g e  most  England.  was  commonly  localities  wood submerged  It  as  been  (1976).  in terrestrial  angiosperm  it  i s most  but has a l s o  decomposer wood  a q u a t i c a de W i l d  from  spore  streams.  each  large  was  1,2) from  on  also  beech  of  four  alder,  both  reported  by  tree  h o l l o w s and  litter  decomposer  F i g u r e 5g.  g e m e l l i p a r a Marvanova  species i s 1972;  Park  a  common 1974;  terrestrial  Gams 1975).  were r e c o r d e d on f o u r d a t e s , a l l on  small  Spores alder  of t h i s in  fungus  December.  51  Bandoni  (1981)  stemflow  of b i g - l e a f  record  recorded  conidia  maple,  G. c r a g i n i f o r m i s  and  (Petersen)  common i n t h e w i n t e r  Gyoerffyella  gemellipara  habitats.  conidia  of G y o e r f f y e l l a  the J.  denticles  &  from  Archer  the  (1973)  Marvanova a n d G y o e r f f y e l l a foam  in  h a s n o t been  r e c o r d e d here  are  England.  recorded the  from  secondary  g e m e l l i p a r a M a r v a n o v a , a s c a n be seen by  the  base o f t h e p r i m a r y  axis.  Figure 5e,f.  A. M a c K i n n o n 83.  Tiny  Triradiate This  once It  at  species  i n stream  Marvanova  The s p o r e s  this  Willoughby  sp. as f a i r l y  aquatic  of  in  tiny  Conidium a m e r o s p o r e was r e c o r d e d once  the spring,  was a l s o  from  foam  this  study,  r e c o r d e d by I n g o l d in  a stream  sp.  Species  of  decomposers  1957,  Hudson 1972,  F i g . 39-3) a s an  in Scotland.  &  genus  ( e . g. Sutton  Park  I t was r a r e l y  spore  was  and  March  from  in  occurred  alder. unknown  recorded i n  was n o t e d .  in  (Willoughby  the  a r e commonly  Scourfield 1964,  1974,  Tetracladium  England  of the l a r g e  and  alder,  a  F i g u r e 5h.  » this  litter  Wild,  (1975c,  of s i x c o n i d i a / m l . stemflow  Tetracladium  two  i n stemflow  winter,  b u t on one o c c a s i o n i n December, on l a r g e  concentration  Makela  both times  i n the  Dyko  1940,  Gams  winter  & Archer  Hirst  e t a l 1969, B a n d o n i  1972,  1973).  Gregory  Bandoni large  poplar. and  terrestrial &  1976,  r e c o r d e d from small  recorded  alder  One  i n February,  T. m a r c h a l i a n u m  spring The  1981).  in  spore  stream  de  foam i n  recorded  here  52  probably  belongs  species;  i t bears  early  in their  Tricladium  some r e s e m b l a n c e  development  Dyko  beech  from  ( 1 9 6 6 ) , and  from  Webster  Descals  &  collection  the  wood  like  any  described  T. m a r c h a l i a n u m 1961  F i g . 7).  conidia  Figure  the  rest  from  gutter  T.  7e.  for  this  by  by  Park  o t h e r s , from  the  water  (1976  herbaceous Gregory  Plate  plant  &  by  Hirst  II, Yadav  (1957).  s p l e n d e n s as a saprobe  on  (1964) r e p o r t e d t h e same s p e c i e s in  streams.  There  were  three  s p o r e t y p e , a l l i n t h e autumn,  s p l e n d e n s and  throughfall. Tr i c l a d i u m  d u r i n g w i n t e r and  the y e a r .  flow  a  S p e c i e s of  litter  Gonczol  (1982) r e p o r t e d T.  i n England of  of  & Oliver  Ingold.  terrestrial  by  spora  l a r g e p o p l a r , and  (1973) r e c o r d foam  stem  air  splendens  ( 1 9 8 1 ) , and  submerged  dates  alder,  stream  T.  trunks  the  Gareth Jones  on a n g i o s p e r m  small  (Greathead  (1976), Bandoni  2 ) , ' and  wood, and  i s not  to  have been r e c o r d e d f r o m  the h o l l o w s of  Fig.  g e n u s , but  spore type resembles  Tricladium  in  this  sp.  This  (1974),  in  Bandoni  i n the s t r e e t s  Willoughby spp. as  spring,  & Archer  abundant  and  less  Canada.  in  common  r e c o r d e d a s p o r e of t h i s  of Vancouver,  from  genus  Figures  5c ,d.  Triposporina This  y a k u e n s i s Matsushima  species  (1965  F i g . 2a;  (1960  Plate  stream  foam  has  been c o l l e c t e d ,  1975a F i g . 2-7;  11-6), in  and  unknown, by  1975b F i g . 3 9 - 5 ) ,  Willoughby  England.  as an  This  & Archer  Hudson & I n g o l d  (1973  s p o r e t y p e was  Ingold  F i g 4k)  from  r e c o r d e d , i n the  53  s t e m f l o w of Matsushima  the  l a r g e and  (1971) d e s c r i b e d  terrestrial).  Figure  Varicosporium  elodeae  This  species  terrestrial,  i s a common decomposer  semi-aquatic,  as  aero-aquatic they  totally  small  sporulated  There are poplar  and  he  three one  December.  Willoughby  commonest  in  streets  that  at  leaves,  also  since  water  (presumably  & Archer  Varicosporium  of  wide  foam  elodeae Kegel  of V a n c o u v e r , C a n a d a .  Figure  1982).  incubated  i t mainly these  the  than  from  when  stagnant  spores,  and  of  considered  rather  two  on  a l l i n November-  (1973) r e p o r t e d stream  (Park  when he  alder,  variety  i s better  surface  isolated  small  a  litter  species  collections on  of  aquatic  this  the  winter-spring  (1981) r e c o r d s the  and  hyphomycete,  submerged;  pools.  i t from d e c a y i n g  January.  Kegel  (1964) s u g g e s t e d  leaves,  a l d e r s , i n November and  7f.  Nilsson an  small  this  species  in England. from g u t t e r 7g.  J . A.  as  Bandoni flow  in  MacKinnon  80.  Spore types  Branched One poplar,  recorded  on  one  date  only  Conidium s p o r e was in  recorded,  December.  qiganteum Crane.  Figure  It 7h.  in  the  appears  stemflow similar  to  of  the  small  Varicosporium  54  Dendrospora One  sp.  s p o r e of t h i s  stemflow  pof  have been  reported  1977,  Bandoni  stemflow belong has  to  D.  by  1981).  Descals &  angiosperm  wood  spring  found  spores  Figure  5b.  Dematiaceous One poplar,  of  aristata  This  species  and  s p o r e was February.  stream  D.  of  of  Dendrospora  (1974,  tenella  is  to  from  F i g . 3j)(both a  saprobe  on  Willoughby & Archer  Bandoni  this  appears  same s p o r e t y p e  Iqbal  in England.  members  genus  in  the  winter  ( p e r s . comm.) has  on  stems of  Juncus.  Conidium  recorded, Figure  Berk. has  foam  7i.  This  (Webster  r e c o r d e d , from  spore  & D e s c a l s 1982).  in  the  stemflow  of  the  large  5a.  & Br. been  recorded  in  Uganda  r e c o r d e d , i n the stemflow Figure  genus  This  1d) and  ( 1 9 6 1 ) , from t e r r e s t r i a l oak  from  S p e c i e s of t h i s  I n g o l d as most common  foam  Cruciform  Tetraploa  Gregory  erecta  i n December.  from  t y p e was  species  1980).  (Webster  s p o r e was  study,  D e s c a l s & Webster. unknown  this  terrestrial litter  spore of t h i s  (i960,Fig.  i n stream  in  i n December.  i n December.  Webster  (1973) r e c o r d e d D. and  One  tenella  Ingold  found  from d e c o m p o s i n g  r e c o r d e d as an  streams  was  alder  of l a r g e a l d e r ,  been  fide  large  type  from  litter (Ingold of  J . A. M a c K i n n o n  the 81.  the  by  a i r s p o r a by  Webster  1966  (1977),  F i g . 11q).  small  poplar,  One in  55  Cornutispora This  1ichenicola  coelomycete  from A u s t r a l i a ,  Hawksw. i s a lichen  the B r i t i s h  parasite  Isles,  Italy,  s p o r e s were r e c o r d e d , f r o m t h e s t e m f l o w one  sampling  collected, of  date  in  November.  as C o r n u t i s p o r a  red cedar, alder,  and  sp.  ; a s an unknown, by G o n c z o l h o l l o w s of beech  trees;  Dicranidion This  fragile  Plate  stemflow  from stems o f Fig  7)  F i g 53)  on  genus were  (1981) from  III,  (1974  this  alder,  Juncus  from  from  the  streams.  is  Harkness  the s p e c i e s  r e c o r d e d by G o n c z o l  a s an unknown from s t e m f l o w Hungary.  It  foam.  was The  (Butterfield the  of  Four  Ii-  Figure  stream  Spores  by M i u r a  Switzerland.  of the l a r g e  maple, and  (1976  and  and  , by B a n d o n i  bigleaf  previously recorded  Lateriramulosa  in  (1971)  from  One  suggests  that  conidia.  it but  should simply  S p o r e s of t h i s  very  from  variable of  7k.  the  s p e c i e s was  leaves  (1973) c o l l e c t e d  F i g . 1-30)  in  Matsushima  This  Marvanova  (1966  tree  r e c o r d e d , i n the stemflow  Figure  r e c o r d e d , from  decaying  Ingold  F i g . III-6)  of a beech  s p e c i e s c a n be  s p o r e was  uni-inflata  November.  hyphomycete,  of t h i s  i n November.  s p o r e was  alder,  r e c o r d e d by  conidia  1973).  large alder,  One  also  foam a t t h e base  (1976  in  a  as  be a  species  of  described  the  by  debris  considered  small  Matsushima  terrestrial  i t from p l a n t not  stemflow  situation.  i n a stream; a s an  hyphomycete  with  i n streams  increase  she  "aquatic"  water-borne greatly  in  56  numbers  after  of  this  fungus  in  flowing  Ingold  r a i n f a l l , are  water  & E l l i s  foam.  a  stimulated  (Marvanova  1952,  Willoughby sp.  October,  stream  a  F i g . &  "Unidentified in  and  1",  in  by  and  sporulation  immersing  1973).  Several  have  reported  1f)  Archer F i g .  growth  4i)  strips  this  record  as  rare,  England.  Figure  6a.  cultures  the  culture  researchers  (1973) very  of  of  spore  this  (e. in  stream  species  occurring  g.  only  (as in  57  DISCUSSION The  results  interesting especially present,  points  concerning with  fungi relation  of  Of  fungi  sporulation.  about  fungi  H-Shaped  recorded  Spore,  Flabellospora,  between  here,  and  their  allows fungi  t h e Double Crescent  Ceratosporium,  spp. ,  a in  Clavariopsis,  the  the Dematiaceous C r u c i f o r m  Lateriramulosa common p o i n t  a l l have t h r e e  Seimatosporium,  all  p o s s e s s appendaged  not  branched  spores.  o r appendaged that  such  a  or appendaged.  appendaged  Ingold are  but  Zygomycetes, and A l g a e two  tetraradiate  also of  explanations shape: e i t h e r  Triradiate,  the Branched  Conidium,  C o n i d i u m , T e t r a p l o a , and  Of t h e o t h e r  fungi  Cornutispora, Several  of  the  only  of  Ascomycetes,  aquatic  habitats.  for this  convergent  spores  It  spores  that  of  this  is  are  branched  water-transported. not  the  spores are  Fusarium).  (1966,1975b) n o t e d often  recorded,  and D i c r a n i d i o n  o f t h e common  percentage  spores are c h a r a c t e r i s t i c  Imperfecti,  Gyoerf f y e l l a  Tiny  (theScolecospores, high  spores  spp.,  o r more b r a n c h i n g arms j o i n e d a t a  o r t o a common a x i s .  Pestalotiopsis,  significant  Varicosporium,  Conidium,  Titaea  Scolecobasidium,  Dendrospora,  proposed  sites,  t h e r o l e of canopy  Triposporina,  Fungi  canopies  the information  Tripospermum  Ceratosporella,  branched  forest  t o the d i v e r s i t y of f u n g i  Tricladium,  or  several  ecosystem. the  branched  indicate  of  As w e l l ,  amount of s p e c u l a t i o n  forest  spp.,  investigation  d i s t r i b u t i o n of these  certain  the  this  information  seasonality  a  of  many  These aquatic  Basidiomycetes, Ingold  (1975b)  evolution  shape  settle  of the more  58  slowly;  or  surfaces.  rates  that  underwater  to  in  these  &  at  base  presence of  a  the  spora passed  Bandoni explanations dispersed  suggested for  (1974, r e l i e d  under  applicable  to  with  torn a  several  the  off  s e t t l i n g .  on  water  (a  films  this  being  spore  films  l i t t e r .  The  conidiophore  by  And branched  water A  might  be  spores  t e r r e s t r i a l  tension  l i t t e r .  with  be  this  present and  in are  mechanism. various and  was  not He  advantageous layers might  than arms  refloated  might  also  fungi.  also  spore  the  in  this  between  spore  in  liberated,  these  was  eject  aerosol  these  that  of  branched  area.  'caltrop')  on  noted  trapped  the  are  that  shape  them  1957),  produced,  and  carry  (1960)  explosion  out  that  can  in  ),  Hirst  occurrences  branched  surface  &  noted  reported  sp.  bubble  water,  Tubaki  fungi  trapped  collected  spores  Tricladium  pointed  conidia  water  its  smaller  after the  in  leaf  planes  1975)  t e r r e s t r i a l  dispersal  easily  on  the  Gregory by  be  bubbles.  of  and  surface  can  hyphomycete  and  a i r  flowing  that  t e r r e s t r i a l  the  they  also  shapes.  (1975)  the  in  He  were  propagules  at  underwater  spores.  other  Bandoni  and  Two  1961;  of  on  difference  spores  spores  bursting  a r i s t a t a  into  unbranched  bursting  aquatic  (Gregory  and  tetraradiate  waterfall.  (Tetraploa  perhaps  of  appreciable  and  a i r ,  above  no  than  (1973)  the  trapped  branched  numbers  into  study, a i r  found  Bubbles  immediately the  effectively  branched  concentrate  surfaces.  noted the  between  Webster  spores  aerosol  (1959)  higher  could  water  more  tetraradiately  Iqbal bubbles  are  Webster  settling found  they  moved  be a  of more spore  occupying  more more  easily easily  59  F i n a l l y , Many  some  aquatic  sigmoid  (and  plane,  most  varied with  seasonally.  When  a  p r e c i p i t a t i o n correlation  and a  when  and is  production  concurrent  not  production  a  one,  but  sporulation  of  in  spore  (Figures  with  the  As  the  well,  trees  many  of  between  is  not  between rainy  temperature  the  fungi  reported  season  in  d i s p e r s a l ,  that and  Vancouver,  can  is the  there spore in  not  an  is  amount  of  However,  the  the  spore  the  spring,  autumn  without  be  shown  (Figure  22)  and  correlation  is  an  during  peaks  in  produce  months inverse The  probably  of  the the  spores  correlation  unexpected.  production.  October,  that  also  here  coldest is  compared  greatest  the  sites  seen  in  the  numbers  the  be  when  production  leaves.  during  can the  the  without  greatest  surprising  It  are  p r e c i p i t a t i o n  is  temperature  spore  with  8-20) it  in  production  sporulation  it  again  spore  aquatic  at  in  obvious  large  fungi  decreases  here,  the  in  The  and  no  liberated.  8-20);  for  is  21),  p a r t i c u l a r l y  specialized  p r e c i p i t a t i o n  a l l  between  p r e c i p i t a t i o n  straight  possessed found  produce  transport.  mainly  (Figures  spores  Scolecospores.  water  of  (Figure  perfect,  before  the  hyphomycetes  were  sporulation  correlation  months.  Since  They  production.  is  when  were  rise  spore  months  study  increases  there  coldest  this  number  of  for  p r e c i p i t a t i o n  that  inverse  adapted  correlation  the  drops  the  are  precipitation  positive  made  t e r r e s t r i a l )  dispersal, The  is  be  non-sigmoid.  water  total  more  in  samples.  the  there  is,  for  should  that  recorded  that  adaptations in  even  scolecospores  scolecospores one  mention  of  And  since  the  year,  correlation onset the  of  time  the when  60  Figures 21 a n d 2 2 . F i g u r e 21. Monthly incident p r e c i p i t a t i o n , stemflow and t h r o u g h f a l l volumes at the sites. F i g u r e 22. M o n t h l y mean t e m p e r a t u r e at the sites. (month 0 = November 1980) Figure 20: " c o i n c i d e n t precipitation "#"=throughfall A"=stemflow. n  300-9- F I G U R E J 2 1  o  a a :  1  200-  O  o o  o o  o o  lOO-i  I  •8 A  o  rt ]  20-r-  *  »  '  ^-  A ,  .A -A •  J  i  iJi Ui J i J m o n t h t  FIGURE 22  I5H  U o, ,  <D  KH  +-» 05 i_ CD  a £  5H  O  N  month  A  |  62  most of  the  remained fungi  "tree  fungi"  begin  dormant  during  the  begin  to  sporulate  season a f t e r having the  summer.  the  late  autumn  into  the  spring. the  possibility the  p r e s e n c e of  investigated. determine spore was  rest a,  Point 17)  was  b, a  of  (the  1  (0.99) w i t h the  the this  23).  the  of  s a m p l e s on  of  in  only  and  are  through  increases  through  drop off  again  sporulation,  the  reflecting  in others, performed  distribution with  that  the  respect  do  was  not  points  to  of  the  to  site  ordinate  in Figure  23  d. stemflow  on  the  PCA  axis;  x-axis  on  x-axis  spore this  sample  the  sample on from  type)  indicates  most  has  a high  that  separation  the  abundance large  (3500 c o n i d i a / m l . ) .  Points  b,  axis  whose s e p a r a t i o n  Variable  2  (Fusarium  second a x i s )  and  Variable  3  (Lunate Spore;  (the  weight  a very  an  a)  samples.  had  y-axis,  (point  on  b  C o l l e c t i o n Date  i s b a s e d p r i m a r i l y on  Scolecospores  Point  rainy  wood  2 was  primarily  axis).  the  not  1 and  points  having  the  variation,  spatial  samples  This  d were s e p a r a t e d  on  the  does not  sample p o i n t s  Scolecospore  the  of  sporulating  of axes  obvious  large alder  first  i n or  i n some t r e e s  The  the  Scolecospores.  based  an  clumping  and  onset  and  after  A l t e r n a t i v e l y , these  is also a spatial  separated  (the  concentration and  c,  the  winter,  ordination  of  clearly  Variable  of  No  with  variation  fungi  was  (Figure  the  labelled  these  i f there  evident  with  there  A PCA  types.  the  again  season.  species  temporal  that  grow  grown v e g e t a t i v e l y  number of  Besides  dry  again  The  until  to  large alder  sp.  ; 0.69 0.71  weight weight  c, is  with with  s t e m f l o w on C o l l e c t i n g  63  F i g u r e 23. P r i n c i p a l Components A n a l y s i s S c a t t e r P l o t . axis=PCA a x i s 1; Y-axis=PCA a x i s 2.  X-  64  65  Date  19)  is  conidia/ml.) on  unusual of  stemflow (540  Date  of  on  indicates  that  spore  type  seemed  sp.  The  the  sites  similar,  same  (though  s l i g h t l y  The  more  number  of  surface  area  to  the G.  for  types  then  may  be  of  recently was With  lower  the  due  l i t t l e the  to  to  me  recorded  aquatic  information information  the  aspen  and  found  in  species  bark  alder  some  due  the  Bandoni  to  in  support by  the  increased perhaps  larger  trees.  the  unknown  lichen  parasite  red  trees  1981,  alder.  has  been  Gonczol  1976)  d i s t r i b u t i o n  presented  my are  those  to  larger in  their  at  or  of  The  from  on  as  cracking,  that  types.  judged  be  the  the  spore red  on  what  data;  seemed  in  based  throughfall  as  may  the  clustering  confirms  of  hyphomycetes  1982,  points,  s i m i l a r ,  This  parasites.  (Carroll  four  different  This  lichens  was  alder  concentration  concentration)  two  recorded.  lichen  (560  large  This  populations,  suggested  lichenicola  there  abundance.  has  (the  these  obviously  types  the  fungal  of  of  trembling  of  stemflow  .  markedly  in  alder  high  observation  spore  trees  d  a  abundance.  colonization  presence  only  the  types  C a r r o l l  Cornutispora  noted  diverse  point has  are  possess  (1100  concentration  significant.  casual  of  small  high  also  sp.  and  generally  presence  C.  The  and  larger  spore  18)  concentration  (the  and  more  mycobiota  are  '  not  c  a  points  from do  point  separation  is  these  obvious  stemflow.  spore  rest  high  ;  Fusarium  clear  the  stemflow  sites  of  Date  d i s t r i b u t i o n  different  and  Fusarium  the  of  Spore; has  conidia/ml.)  clustering  its  18)  Collection  Despite  Dr.  Lunate  C o l l e c t i o n  conidia/ml.)  in  here,  and  limited  66  speculation is  on  the  role  these  fungi  in  the  forest  ecosystem  possible. The  the  greatest  above-ground  t e r r e s t r i a l  temperate Most  of  and  hardwood  stream  fungi  that  throughfall  (with  the  P e n i c i l l i u m  ,  important been  in  and  the  from  acuminata,  Tricladium  not  observed to  these  species  actively  form  main  species  regions on  are  study  99%  of  of  ecosystem, mosses  in  of  The  systems  the  f i r s t energy  with the  systems  of  believed  conidia  high  fungi  in  suggest  to  , be  have  t e r r e s t r i a l Alatospora  elodeae Populus  the  concentrations  would  and  they  of  Varicosporium  these  1978).  Aspergillus  moist  and  north  al  although  in  Alnus  et  to  stemflow  not  land,  sites,  and  of  are  leaves my  from  input  in  (Gosz  species  on  matter  were leaves.  l i t t e r of  were  spores  that  they  of were  there. the  t r u l y  to  streams)  stream  matter  a  ,  leaves  allochthonous,  t e r r e s t r i a l  At  the  restricted  organic  the  leaves  growth  some  role  decomposition.  of  t e r r e s t r i a l  however,  growing  The  on  on  of  sp.  organic  leaves  Zygomycetes)  1981).  demonstrate  employed;  tree  decomposing  (Bandoni  of  i d e n t i f i e d  exception  situations  Methods  is  have  decomposition  isolated  commonly  I  mass  decomposing  forests  the  a  of  the  in  systems that  produced  (the order  they  the  in  to  other  1% b e i n g  Of  in  t h i s ,  New  be  was  (those  in  temperate almost in  the  & Likens  Hampshire,  largely was  to  depend  Fisher  stream  47%  north  plants  watershed). stream  hyphomycetes  appears  by  input  stream.  "aquatic"  from  leaf  forested entirely adjoining (1973),  showed  the  that  t e r r e s t r i a l  photosynthesized  dissolved  in  organic  by  matter  67  (particles  less  particulate  organic matter  in  diameter).  their in  organic  diameter).  passed  study  The  site.  tell  by  organisms,  The d a t a  component  (Suberkropp  form  which  in  organic  matter  I973a,b). matter  The  input  utilized biomass and  can in  hyphomycetes, stream  animals  directly  i s well  an  fungi,  important  a  stream  than  1 mm.  to  diameter) and  The  from  (Barlocher  important  to  the  be  the  particulate &  Kendrick  but  organic can  and b a c t e r i a by  be  leaves  invertebrates.  including  link  on  i n the food  the chain  by Cummins  ( 1 9 7 4 ) a n d Cummins  The  aquatic of the  F u r t h e r p r o c e s s i n g of the o r g a n i c matter  documented  the  microbial  t o be t h e most appear  fine  "processing"; the microbial  be  are  from  invertebrates  of fungi  the  dioxide in  of the allochthonous  after  utilized  1  matter  degradation,  i s passed  by t h e g r o w t h  particular  to carbon  1974,1976,1980),  invertebrates  ecosystems.  streams  appear  components  than  organic  invertebrates.  to  these  produced  microbes,  microbial  matter  major  coarse  i s converted  less  organic the  9 6 % was  story.  are unavailable to the  by  twigs  & Klug  microns  e t a_l ( 1 9 7 3 )  (particles  the fungi  0.5  greater  total  organic matter  "shredder"  which  of the  of Sedell  of a b r a s i o n ,  the  among  (particles  t h e same  a n d 5 3 % was  greater than  3 4 % was c o n v e r t e d  organic matter  of  diameter)  organic matter,  percent  particulate  a combination  actions  and  in  (particles  essentially  coarse  particulate  microns  matter  Sixty-six  downstream,  Oregon  0.5  Of t h e p a r t i c u l a t e  particulate mm.  than  &  in Klug  (1979). The  fungi  reported  in this  study  are probably  perthophytes  68  on is  wood the  has  or  case  been  support  shown  sampled As  noted  chemical floor  leaves  trapped  considered,  be  spores  stemflow  spores reach  both  are the  hyphomycetes from  fallen  and leaves  considerable  well,  in  systems,  hollows  in  v i s i b l e  as  a  It  tree  may  (Gonczol  species  1976;  found  in  this  any  of  the  in  the  source  organisms  important  of  of  only  inoculum Many  of  fungi  the  floor;  and  t e r r e s t r i a l  and  it  few  the  trees.  the is  fungi  possible  hyphomycetes  long  after  c e l l u l o l y t i c analagous probably  to  w i l l  well  convert  that  is,  they  are  by from  they  are  of  the  Many  i n i t i a l l y  leaves,  at  important  in  isolate  aquatic  Trichoderma) these  fungi  c a p a b i l i t i e s .  studied  the  bacteria,  isolated  and  l i g n o l y t i c  that  mycobiota  (Penicillium,  abscission,  and  the  The  to  stemflow  represented  that  that  forest  to  l o c a l i t i e s . l i k e l y  of It  be  seems  from  for  inoculum  the  on  These  the  contains  the  forest  throughfall  in  role  microbial  can  other  fungi  to  and  r a i n f a l l .  throughfall  very  not  stemflow  incident  equally  well.  floor  situation the  or,  parasites.  fungi  I,  the  the  degradation.  have  a  as  trees  contain  ultimate  a  and  forest  in  were  contributions  water-borne,  abscission, their  algae  material  in  as  though  and  saprobes  An  the  protozoans,  lichen  the  Table  important  system.  Here  in from  are  may  of  Fusarium)  study.  different  floor.  decomposing  different  no  changes  in  many  some  decomposer  forest  1ichenicola, trapped  of  this  A l t e r n a r i a ,  leaves  previously  throughfall  (  including  in  chemically  be  that  However,  trees  parasites  Cornutispora  1981),  study.  and  for  tree  populations  Bandoni  are  bark,  in  c e l l u l o s i c  As  aquatic plant  69  m a t e r i a l to m i c r o b i a l 'shredders'. (Cummins  Most  biomass  available  to  invertebrate  of these 'shredders' lack c e l l u l a s e enzymes  1974).  There are other sources of inoculum f o r abscission.  Once  a  leaf  hundreds of thousands course  the  of  of  has  conidia  i t s degradation.  d i s p e r s e d by water f i l m s  been of  leaves  inoculated, a  after  i t may  bear  over  the  hyphomycete  These c o n i d i a , which c o u l d then be  (Bandoni  & Koske 1974),  in spreading the fungus to other l e a v e s . spores present i n stemflow  the  are  important  In f a c t , the number of  and t h r o u g h f a l l c o u l d not account f o r  the l a r g e number of c o l o n i e s of these fungi present on l e a v e s on the  forest  largely  floor.  gone  from  Since the l e a v e s from the p r e v i o u s year are the  forest  floor  by  the  time  of  next  a b s c i s s i o n , they do not represent a source of inoculum f o r newly f a l l e n leaves.  However, spores from the s o i l can move, on water  films,  the s o i l to i n o c u l a t e the f r e s h l y f a l l e n l e a v e s .  up  in  At l e a s t one of the fungi r e p o r t e d here has a l s o been r e p o r t e d from s o i l s It  seems  likely  the  (eg. Waid  elodeae)  1944).  that spores i n s t e m f l o w / t h r o u g h f a l l a l s o  serve as inoculum f o r the f r e s h l y fungi l i s t e d  (Varicosporium  fallen  leaves.  A few  of  the  i n the R e s u l t s s e c t i o n of t h i s t h e s i s were found on  leaves around  the base of the sample t r e e s .  Gonczol  (1976)  found that l e a v e s i n hollows i n the trunks of beech t r e e s , which had not been i n c o n t a c t with the ground,  possessed  rich  p o p u l a t i o n s i n c l u d i n g s e v e r a l of the fungi found i n t h i s One mycobiota  role  for  stemflow/throughfall,  then,  fungal study.  would  inoculum to the t e r r e s t r i a l decomposing systems.  be  as Both  70  tree species  studied drop t h e i r  t h e months  of  greatest  stemflow and t h r o u g h f a l l , months least  of  i n November.  fungal  and a r e a l s o  thestemflow/throughfall.  of  spores that a r espread  These l e a v e s  their  documented  They a r e  with  well  b r o k e n down m a i n l y  transported  decompose i n t h e t e r r e s t r i a l  fungi  inoculated  At  spores  other  leaves.  t o streams,  (Kaushik  by t h e a q u a t i c  leaves  the  p r o d u c e l a r g e numbers  in water f i l m s t o i n f e c t are  h a s been  coincident  volume ( F i g u r e 21).  Some o f t h e s e i n f e c t e d l e a v e s fate  coincident  become i n o c u l a t e d by t h e s a p r o b e  in  first  spore c o n c e n t r a t i o n s i n  maximum s t e m f l o w / t h r o u g h f a l l  some o f t h e l e a v e s  The  s p e n d on t h e g r o u n d a r e  f o u r months t h a t t h e s e l e a v e s with  leaves  & Hynes  where 1971).  hyphomycetes.  ecosystem; again  from s t e m f l o w / t h r o u g h f a l l  Other  some o f t h e  may be i m p o r t a n t i n  decomposition. All the  of this  trees.  fungi  r a i s e s the q u e s t i o n  A s was m e n t i o n e d  recorded  inthis  s t u d i e s by T u b a k i  (1960)  of t h e c r e e k  sufficient some  most  of  present  i na e r o s o l , there  Though  t o generate  any  was no d i s t u r b a n c e  amount  f u n g i may have been t r a n s m i t t e d  f o r the diversity in the trees.  imperfect  possess branched c o n i d i a .  birds t o the treetops, but t h i s explanation account  the  h a v e shown t h a t b r a n c h e d s p o r e s c a n be  t o some h e i g h t  of  earlier,  study  transmitted  Spores  o f how t h e s e f u n g i g e t i n t o  of  t h e most  aerosol.  on t h e f e e t o f  seems  and abundance  Perhaps  of  unlikely fungal  likely  to  species  explanation  i n v o l v e s t h e teleomorphs o f these hyphomycetes and Coelomycetes. Where  known,  wind-dispersed  the teleomorphs spores.  of the fungi  M o s t o f t h e known  i n t h i s study  teleomorphs  have  of the  71  aquatic  hyphomycetes  in t e r r e s t r i a l  a r e Ascomycetes  found as saprobes on wood  s i t u a t i o n s near streams (Webster & D e s c a l s 1979).  The w i n d - d i s p e r s e d a s c o s p o r e s of t h e s e teleomorphs c o u l d be  transported  to  the  easily  tops of t r e e s , where t h e i r g e r m i n a t i o n  c o u l d produce the anamorphic hyphomycete s t a t e .  72  APPENDIX  1 CALCULATION OF STEMFLOW AND THROUGHFALL  Regression equations precipitation  in  d a t a of H e l v e y  & Patric  for  temperate  predicting  hardwood f o r e s t s ,  ( 1965),  (1977),  and H u t c h i n s o n & R o b e r t s  (summer  values)  S=0.  stemflow  Lawson  VOLUMES volume  derived  ( 1967), Henderson  (1981),  from  from the e_t  al  are:  005P-0.014  to S=0.087P-0.001 and (winter  values)  S=0.017P-0.017 to  S=0.048P-0.001  where S=stemflow  volume,  P=precipitation or  from  Similar equations  values)  T=0.697P-0.064 to T=0.955P-0.002 and  volume  l e s s t h a n 0 . 5 % t o a l m o s t 9% of  as stemflow. (summer  and  for  the t o t a l  throughfall  precipitation  are:  73  (winter  values)  T=0.789P-0.038 to  T=0.959P-0.001  where T=throughfall  volume, and  P=precipitation or  from  96%.  about  Using  (summer  volume  70% o f t h e p r e c i p i t a t i o n  the g e n e r a l i z e d  equations  as t h r o u g h f a l l , t o almost of Helvey  & Patric  (1965):  values)  S=0.041P-0.005 and T=0.901P-0.031 and (winter  values)  S=0.062P-0.005 and  T=0.914P-0.015  where S=stemflow volume T=throughfall  volume, and  P=precipitation  volume  and  November  (the  considering months  that  t o March  most  trees  inclusive are  as  without  winter  months  leaves),  the  74  precipitation station the  were e n t e r e d  stemflow  sampling  data  period  and  from into  Environment the  equations  throughfall  (Figure  21).  volumes  Canada  meteorological  to g i v e rough v a l u e s f o r at  my  site  over  the  75  APPENDIX  2.  MaIt-yeast-peptone 7.0  g.  1.0  g.  0.5  g.  yeast  7.0  g.  ICN  1000  malt  MEDIA  (MYP) medium  ml.  When  extract  agar  d i s t i l l e d  medium  used  for  water  was  used  for  isolations  that  soluble  tetracycline  were  maintenance from  contaminated was  added  to  inhibit  b a c t e r i a l  Sorbose-tetracycline 4.0  g.  0.5  g.  yeast  7.0  g.  ICN  by  added  malt-yeast-peptone-tetracycline  100  1972)  extract  cultures  1000  (Bandoni  soytone  This  was  EMPLOYED  of  nature,  or  bacteria. after  (MYPT)  axenic for 100  transferring mg/1  autoclaving  agar.  cultures.  The  to  waterproduce  tetracycline  growth.  (SORB-T)  medium  tetracycline  (added  (Bandoni  1981)  sorbose  ml. mg/1  extract  agar  d i s t i l l e d  water  water-soluble  after  autoclaving)  76  This sorbose  medium  as  a  PeniciIlium other added  to  carbon or  fungi  was  used  source,  Aspergillus on  the  discourage  7.0  the  fungi  colonies  As  b a c t e r i a l  isolations  "weed"  form  plate.  Oat Oat  for  with  and  MYPT,  from such do the  nature. as  With  species  of  overgrow  the  tetracycline  was  not  growth.  agar  (OA)  medium  flakes g.  1000  ICN  ml.  agar  d i s t i l l e d  The  autoclaved  autoclaved induced other  water  oat  flakes  sporulation  media.  water  of  and  (6-10 some  agar  were  flakes/petri  fungi  that  would  poured plate). not  over This  the medium  sporulate  on  77  APPENDIX  3.  VARIATIONS  Spore remain from  concentrations  constant Table  beginning  of  termination  a of  after  certain  plateau  sporulation  Table  I I I . a  during  III,  sharply  during  IN SPORE  a  the  the a  this  the  or  r a i n f a l l .  from  rain.  As is  spores The  and then  represents  the  A RAINFALL  throughfall  concentration  period,  can  be  highest  produced  do  seen at  gradual  the  following  concentration increases  not  drops  slowly  to  increase  r a i n f a l l .  in  Large  stemflow alder  sampling period (hours after beginning 0 0-0.25 0.25-0.50 0.50-0.75 0.75-1.0 1.0-1.5 1.5-2.0 2.0-3.0 3.0-4.0 4.0-5.0 5.0-6.0  of  probably  short  Variation  r a i n f a l l .  stemflow  period  previous  level;  during  in  spore  r a i n f a l l ,  CONCENTRATION DURING  of  spore  stemflow,  r a i n f a l l )  concentration March  14,  1982.  spore concentration (conidia/ml) 00 44 21 07 1 1 16 31 35 40 36 42  a in  78  REFERENCES  A r c h e r , J . 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