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UBC Theses and Dissertations

A monograph of the genus H̲y̲g̲r̲o̲h̲y̲p̲n̲u̲m̲ Lindb.(musci) Jamieson, David William 1976

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A MONOGRAPH OF THE GENUS HYGROHYPNUM LINDB. (MUSCI)  by DAVID WILLIAM JAMIESON B... A.  Humboldt S t a t e  U n i v e r s i t y , 1966  M.. A.  Humboldt S t a t e  U n i v e r s i t y , 1969  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE  REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY  i n t h e Department of BOTANY  We a c c e p t t h i s t h e s i s as conforming to the required standard  THE  UNIVERSITY OF BRITISH COLUMBIA August, 1976  (c) David William Jamieson  In  presenting  this  an a d v a n c e d d e g r e e the I  Library  further  for  agree  in p a r t i a l  fulfilment  of  at  University  of  Columbia,  the  make  it  freely  that permission  s c h o l a r l y p u r p o s e s may  by h i s of  shall  thesis  this  representatives. thesis  for  available for  financial  written permission.  Department  of  The U n i v e r s i t y  ]?oT^2/rtt^ of  British  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  12  XL  \ J 1L C  X  by  the  is understood gain  Columbia  for  extensive  be g r a n t e d  It  British  shall  requirements  reference copying of  Head o f  that  not  the  I  agree  and this  be a l l o w e d  that  study. thesis  my D e p a r t m e n t  copying or  for  or  publication  w i t h o u t my  Chairman:  P r o f e s s o r W.  B.  Schofield  ABSTRACT The  genus Hygrohypnum L i n d b .  f o r the f i r s t  time.  (Musci)  i s monographed f o r the  world  T h i r t y - n i n e s p e c i e s and a l a r g e number o f s u b s p e c i f i c  t a x a r e c o g n i s e d a t the i n c e p t i o n o f t h i s study were i n v e s t i g a t e d u s i n g t r a d i t i o n a l t o o l s o f herbarium  taxonomy, e x t e n s i v e f i e l d work  N o r t h America and the e x p e r i m e n t a l species  in  a uniform  c u l t i v a t i o n o f ten North  i n which the i m p o r t a n t  taxonomic c h a r a c t e r s a r e :  leaf cells,  p e r i c h a e t i a l l e a v e s and the annulus. l o w i n g 16 s p e c i e s are r e c o g n i s e d : culum (De Not.)  H.  norvegicum  comb.,  (Lindb.)  eugyrium  stem anatomy, shape o f  H.  Based on t h e s e c h a r a c t e r s the  H_. alpinum  smithii  (Sw.  H_. mo l i e  (Lindb.) Loesk. , in Lilj.)  _ . ochraceum  (Schmip.) B r o t h . ,  and H_. s t y r i a c u m are new  bestii  Broth., styriacum  a l p e s t r e (Hedw.) Loesk.,  closteri  (Ren.  e t Card.)  (Aust.)  Grout.  known p r e v i o u s l y as a Western N o r t h American endemic, i s now  The  _.  Grout, Hygrohypnum  i n Western North America and H_.  on the Upper P e n i n s u l a o f M i c h i g a n  fol-  ( W i l s . ex Turn.) Loesk.,  H_. subeugyrium and H.  H.  H.  the  H_. d u r i u s -  Broth.,  (Hedw.) Loesk.,  luriduirt (Hedw.) Jenn. ,  Loesk.,  differ-  sexuality, structure of  H_. c o c h l e a r i f o l i u m (Vent, i n De Not.)  H. montanum (Lindb.) B r o t h . , alpinum  H.  (Schimp.) Amann,  Broth.,  H. p o l a r e H.  nov.  e t Card.) H o l z . ,  (Limpr.)  Hemisphere genus,  i t s apex, l e a f c o n c a v i t y , c o s t a s t r u c t u r e , a l a r c e l l  e n t i a t i o n , l e n g t h o f the m a r g i n a l  (Ren.  American  environment.  These s t u d i e s show t h a t Hygrohypnum i s a N o r t h e r n  the l e a f and  throughout  bestii,  shown to be  and around the G u l f o f S t . Lawrence.  taxonomic t r e a t m e n t p r e s e n t s a g e n e r i c d e s c r i p t i o n and keys t o  the s p e c i e s and p r o v i d e s each s p e c i e s w i t h a d e s c r i p t i o n , a d i s c u s s i o n o f  iii variability  and taxonomic m a t t e r s ,  The  s t a t u s o f each ..excluded  The  g e n e r i c concept and  n i s e d s p e c i e s are examined by  i l l u s t r a t i o n s and a d i s t r i b u t i o n  map.  taxon i s d i s c u s s e d .  the p o s s i b l e r e l a t i o n s h i p s among the comparing the author's  mation d e r i v e d from an o r d i n a t i o n and two  cluster  opinion with  analyses.  recoginfor-  TABLE OF CONTENTS ABSTRACT LIST OF FIGURES LIST OF TABLES ACKNOWLE DGMENTS INTRODUCTION THE TAXONOMIC HISTORY OF THE GENUS METHODS OF THE MORPHOLOGICAL ANALYSIS METHODS OF CULTIVATION METHODS OF PREPARATION EXAMINATION  OF MATERIAL FOR MICROSCOPE  RESULTS OF THE MORPHOLOGICAL ANALYSIS Stem AnatomyL e a f Shape and Symmetry L e a f Apex Costa Leaf Concavity Median L e a f Alar  Cells  Cells  Response  to Drying  Sexuality P e r i c h a e t i a l Leaves Sporophyte TAXONOMIC TREATMENT Generic  Synonomy  Generic D e s c r i p t i o n  o f Hygrohypnum  Key t o t h e S p e c i e s o f Hygrohypnum  V  Hygrohypnum alpinum  98  Hygrohypnum d u r i u s c u l u m  113  Hygrohypnum s m i t h i i  136  Hygrohypnum b e s t i i  150  Hygrohypnum c o c h l e a r i f o l i u m  ...169  Hygrohypnum norvegicum  184  Hygrohypnum m o l l e  193  Hygrohypnum s t y r i a c u m  211  Hygrohypnum l u r i d u m  224  Hygrohypnum a l p e s t r e  270  Hygrohypnum p o l a r e  282  Hygrohypnum ochraceum  299  Hygrohypnum eugyrium  329  Hygrohypnum subeugyrium  347  Hygrohypnum montanum  36 3  Hygrohypnum c l o s t e r i  375  EXCLUDED SPECIES AND SPECIES OF QUESTIONABLE AFFINITIES  387  DISCUSSION  398  LITERATURE CITED  418  vi  LIST OF FIGURES FIG.  1  V a r i a t i o n i n stem anatomy  23  FIG.  2  Terminology o f l e a f shape  28  FIG.  3  Comparison o f l e a f shape v a r i a t i o n w i t h i n and between s p e c i e s  30  FIG.  4  Comparison o f l e a f shape v a r i a t i o n w i t h i n and between s p e c i e s  32  FIG.  5  Comparison o f l e a f shape v a r i a t i o n w i t h i n and ~ between s p e c i e s  34  FIG.  6  Comparison o f l e a f shape v a r i a i t o n w i t h i n and between s p e c i e s  36  FIG.  7  Comparison o f l e a f shape n a t u r a l l y growing p l a n t s and t h e i r response t o a r t i f i c i a l c u l t u r e  38  FIG.  8  Comparison o f l e a f shape i n n a t u r a l l y growing p l a n t s and t h e i r response t o a r t i f i c i a l c u l t u r e  40  FIG.  9  Comparison o f l e a f shape i n n a t u r a l l y growing p l a n t s and t h e i r response t o a r t i f i c i a l c u l t u r e  42  FIG.  10  D e s c r i p t i v e t e r m i n o l o g y f o r the shape o f the l e a f apex  46  FIG.  11  V a r i a t i o n i n the shape o f the l e a f  apex  48  FIG.  12  V a r i a t i o n i n the shape o f the l e a f  apex  50  FIG.  13  The n a t u r e o f l e a f  FIG.  14  V a r i a t i o n i n marginal l e a f  cells  57  FIG.  15  V a r i a t i o n i n marginal leaf  cells  59  FIG.  16  Variation i n alar c e l l  differentiation  64  FIG.  17  Variation i n alar c e l l  differentiation  66  FIG.  18  Variation i n alar c e l l  differentiation  68  FIG.  19  Variation i n alar c e l l  differentiation  70  FIG.  20  V a r i a t i o n i n the a l t e r a t i o n i n h a b i t from wet t o dry c o n d i t i o n s i n m o r p h o l o g i c a l l y d i f f e r e n t shoots  concavity  54  74  VI1  FIG.  FIG.  21  22  Variation i n a l t e r a t i o n i n habit from wet to dry conditions i n morphologically d i f f e r e n t shoots  76  The pseudo-paroicous inflorescence of Hygrohypnum styriacum  79  FIG.  23  Variation i n the p e r i c h a e t i a l leaf apex  83  FIG.  24  Variation i n the p e r i c h a e t i a l leaves  85  FIG.  25  Variation i n the p e r i c h a e t i a l leaves  87  FIG.  26  Variation i n the leaf shape and the habit of the shoots of Hygrohypnum alpinum C e l l u l a r d e t a i l of the p e r i c h a e t i a l and foliage leaves of Hygrohypnum alpinum  109  FIG.  27  107  FIG.  28  Distribution.map of Hygrohypnum alpinum  110  FIG.  29  Variation i'n_the habit of leaf shoots of Hygrohypnum duriusculum and comparisons . between the wet and dry conditions i n the shoots of Hygrohypnum duriusculum  124  Variation i n the leaf shape, length of marginal leaf c e l l s and the habit of leafy shoots of Hygrohypnum duriusculum  126  FIG.  FIG.  30  31  Variation i n the apices and the c e l l u l a r d e t a i l of the leaves of Hygrohypnum duriusculum  128  FIG.  32  D i s t r i b u t i o n map of Hygrohypnum duriusculum  129  FIG.  33  143  FIG.  34  Variation i n the leaf shape and the habit of leafy shoots of Hygrohypnum smithii C e l l u l a r d e t a i l s of the p e r i c h a e t i a l and foliage leaves of Hygrohypnum smithii  145  FIG.  35  D i s t r i b u t i o n map of Hygrohypnum smithii  146  FIG.  36  Variation i n the habit of the .shoots of. ,J Hygrohypnum b e s t i i and a comparison of i n d i v i d u a l shoots i n the wet and dry conditions  159  FIG.  37  Variation i n the l e a f shape of Hygrohypnum b e s t i i  161  FIG.  38  C e l l u l a r d e t a i l of the foliage leaves, of Hygrohypnum b e s t i i  16 3  D i s t r i b u t i o n map of Hygrohypnum b e s t i i  164  FIG.  39  viii  FIG. 40  V a r i a t i o n i n the l e a f shape and the h a b i t o f the moist shoots o f Hygrohypnum c o c h l e a r i f o l i u m  178  C e l l u l a r d e t a i l o f the l e a v e s o f Hygrohypnum cochlearifolium  180  FIG. 42  D i s t r i b u t i o n map o f Hygrohypnum c o c h l e a r i f o l i u m  181  FIG. 43  V a r i a t i o n i n the l e a f shape, t h e c e l l u l a r d e t a i l o f the f o l i a g e l e a v e s and the h a b i t o f the shoots o f Hygrohypnum norvegicum  190  FIG. 41  FIG.  44  D i s t r i b u t i o n map o f Hygrohypnum norvegicum  191  FIG.  45  V a r i a t i o n i n the l e a f shape and the h a b i t o f the shoots o f Hygrohypnum m o l i e  204  FIG.  46  C e l l u l a r d e t a i l o f the f o l i a g e and p e r i c h a e t i a l l e a v e s o f Hygrohypnum m o l i e  FIG.  47  206  D i s t r i b u t i o n map o f Hygrohypnum m o l i e  207  FIG. 48  V a r i a t i o n i n l e a f shape and shoot h a b i t o f Hygrohypnum s t y r i a c u m  218  FIG. 49  C e l l u l a r d e t a i l of the f o l i a g e leaves of Hygrohypnum s t y r i a c u m  220  FIG.  50  D i s t r i b u t i o n map o f Hygrohypnum s t y r i a c u m  FIG.  51  FIG.  52  V a r i a t i o n i n the h a b i t o f the m o i s t shoots o f Hygrohypnum l u r i d u m V a r i a t i o n i n the l e a f shape o f Hygrohypnum luridum  FIG. 53  FIG.  54  FIG. 55  221  249 251  V a r i a t i o n i n the l e a f shape o f Hygrohypnum luridum  253  V a r i a t i o n i n the c e l l u l a r d e t a i l o f the f o l i a g e l e a v e s o f Hygrohypnum l u r i d u m  255  Variation  i n alar c e l l differentiation of  Hygrohypnum l u r i d u m  257  FIG. 56  D i s t r i b u t i o n map o f Hygrohypnum luridum,  258  FIG.  57  V a r i a t i o n i n the l e a f shape and the h a b i t o f the shoot o f Hygrohypnum a l p e s t r e  2 76  FIG.  58  C e l l u l a r d e t a i l of the f o l i a g e Hygrohypnum a l p e s t r e  278  FIG. 59  leaves o f  D i s t r i b u t i o n map o f Hygrohypnum a l p e s t r e  279  XX  FIG.  FIG.  60  61  V a r i a t i o n i n t h e l e a f shape and the h a b i t o f the shoots o f Hygrohypnum p o l a r e  292  C e l l u l a r d e t a i l s o f the f o l i a g e l e a v e s and the stem o f Hygrohypnum p o l a r e  294  FIG.  62  D i s t r i b u t i o n map o f Hygrohypnum p o l a r e  295  FIG.  63  Variation:'-in the h a b i t o f l e a f y shots o f Hygrohypnum ochraceum and comparisons between the wet and d r y c o n d i t i o n s i n t h e s h o o t s  313  V a r i a t i o n i n the l e a f shape o f Hygrohypnum ochraceum  315  C e l l u l a r d e t a i l o f the f o l i a g e l e a v e s o f Hygrohypnum ochraceum  317  FIG.  FIG.  FIG.  64  65  66  The stem anatomy and the v a r i a t i o n i n the a l a r c e l l s o f Hygrohypnum ochraceum  319  FIG.  67  D i s t r i b u t i o n map o f Hygrohypnum ochraceum  320  FIG.  68  V a r i a t i o n i n t h e h a b i t o f the shoot and t h e l e a f shape o f Hygrohypnum eugyrium  339  C e l l u l a r d e t a i l o f t h e l e a v e s and the stem o f Hygrohypnum eugyrium  341  FIG.  69  FIG.  70  D i s t r i b u t i o n map o f Hygrohypnum eugyrium  342  FIG.  71  V a r i a t i o n i n the l e a f shape and the h a b i t o f the shoot o f Hygrohypnum subeugyrium i n Europe and N o r t h America  357  V a r i a t i o n i n the l e a f shape and the h a b i t o f the s h o o t o f Hygrohypnum subeugyrium i n Japan  359  FIG.  FIG.  72  73  C e l l u l a r d e t a i l o f the f o l i a g e l e a v e s o f Hygrohypnum subeugyrium  361  FIG.  74  D i s t r i b u t i o n map o f Hygrohypnum subeugyrium  362  FIG.  75  The h a b i t o f , t h e shoot and v a r i a t i o n i n l e a f shape o f Hygrohypnum montanum  370  FIG.  76  C e l l u l a r d e t a i l o f the f o l i a g e leaves o f Hygrohypnum montanum  372  D i s t r i b u t i o n map o f Hygrohypnum montanum  373  FIG.  77  X  FIG.  FIG.  78  79  The h a b i t o f the shoot and the v a r i a t i o n i n , l e a f shape o f Hygrohypnum c l o s t e r i  382  C e l l u l a r d e t a i l o f the f o l i a g e leaves of Hygrohypnum c l o s t e r i  384 385  FIG.  80  D i s t r i b u t i o n map o f Hygrohypnum c l o s t e r i  FIG.  81  Possible  relationships  among t h e s p e c i e s o f  Hygrohypnum  400  FIG.  82  Weighted p a i r group c l u s t e r a n a l y s i s  411  FIG.  83  Unweighted p a i r group c l u s t e r a n a l y s i s  412  FIG.  84  Ordination analysis  413  o f Hygrohypnum  xi  LIST OF TABLES  TABLE 1  Morphological features subject to analysis  18  TABLE 2  A comparison o f t h e s p e c i e s o f Hygrohypnum and the t a x o n o m i c a l l y i m p o r t a n t f e a t u r e s w i t h i n the genus  406-418  TABLE 3  Similarity  matrix  410  xii ACKNOWLEDGMENTS I would l i k e t o express s i n c e r e thanks to Dr. W i l f r e d S c h o f i e l d f o r h i s f r i e n d s h i p and h i s p a t i e n t s u p p o r t and ment throughout t h i s study.  Thanks...are a l s o due  encourage-  the N a t i o n a l  C o u n c i l o f Canada f o r f i n a n c i a l s u p p o r t through g r a n t A1217 Schofield The  and  o f the c u r a t o r s o f the  acknowledged ( a b b r e v i a t i o n s  GRO,  H,  BP,  H-BR,  according  BRNM, C, CAL,  WTU.  f o r advice  to  H-SOL, HSC,  t o Holmgren and Kueken, 1974) :  J E , KRAK, LD, P,  LE, MAK,  S-PA,  FLAS,  MICH, MIN,  TENN, TRTC, UAC,  My .thanks are a l s o extended to Dr. W. on n o m e n c l a t u r a l m a t t e r s .  FH,  C o l o r a d o , as the extremely a c c u r a t e b u t e d specimens a l l o w e d me  and  many o f my  work.  to r e t h i n k o t h e r s .  the p r o o f i n g o f the  source o f encouragement d u r i n g  Columbia.  my  distri-  I,..would l i k e  the  Stein.  Dr.  thesis  Many reinforce  Stein graciously  f i n a l manuscript and the  long tenure a t The  was  f i n a l months o f  L a s t l y , I acknowledge the encouragement I have r e c e i v e d  many f r i e n d s I have made d u r i n g my  Utrecht  Collins,  d i s c u s s i o n s w i t h Dr. Maze a l l o w e d me  t y p e d the c o r r e c t i o n s d u r i n g  US,  I would a l s o l i k e to o f f e r a  Jack R. Maze and J a n e t R.  i d e a s w h i l e f o r c i n g me  a much a p p r e c i a t e d  UC,  t o r e l o c a t e many c r u c i a l s p e c i e s .  t o Drs.  unorthodox and p r o v o c a t i v e  UBC,  c o l l e c t i n g d a t a on h i s w i d e l y  their constructive c r i t i c i s m of i t .  most s i n c e r e thank you  MNA,  F o r much o f the success i n  committee f o r t h e i r p a t i e n t e f f o r t s i n r e a d i n g  G,  MO,  D. Margadant o f  f i e l d work, I am most g r a t e f u l t o Dr. F. J . Hermann o f F o r t  t o thank my  Dr.  following herbaria i s g r a t e f u l l y  CANM, COLO, DUKE, F, FB,  NFLD.', ... NICH, NSPM, NY, 2NYS, OTT, UWO,  Research  the U n i v e r s i t y o f B r i t i s h Columbia Research Committee.  cooperation  ALTA, B, BM,  B.  from  University of  the British  1  INTRODUCTION Hygrohypnum i s a genus o f s e m i - a q u a t i c p l e u r o c a r p o u s mosses t h a t i s c u r r e n t l y t r e a t e d i n the f a m i l y Amblystegiaceae, i t s e l f a group o f t a x a o f predominantly a q u a t i c o r o t h e r w i s e damp environments. taxa p r e s e n t l y t r e a t e d w i t h i n the genus can be Species  Muscorum, but  t r a c e d to Hedwig's  (1801)  the o r i g i n a l concept o f the genus d a t e s from Schimper  (1853) i n B r y o l o g i a Europaea. and  Certain  Schimper named the genus Limnobium Schimp.  d i s t i n g u i s h e d i t from Hypnum on  the b a s i s o f the bdtoadly o v a t e -  l a n c e o l a t e t o almost o r b i c u l a r l e a v e s and  the nature o f the a l a r c e l l s  which were s a i d t o be h a r d l y d i f f e r e n t i a t e d from o t h e r b a s a l l e a f c e l l s and  l i t t l e o r not a t a l l i n f l a t e d o r excavated.  the genus grew, Schimper noted o n l y the h a b i t a t o f the f i v e o r i g i n a l The  manifold  systematic  Of the h a b i t a t i n which  t h a t the name, Limnobium, r e f l e c t e d  species.  problems i n the genus Hygrohypnum have t h e i r  o r i g i n i n the h a b i t a t i n which £he genus o c c u r s and modern g e n e r i c  the absence o f a sound,  concept.  Hygrohypnum n o r m a l l y o c c u r s on r o c k s o r stones between the h i g h low water l e v e l s ; o r , i n o r b e s i d e streams.  and  s m a l l , c o l d , s w i f t l y r u n n i n g mountain  L i k e o t h e r p l a n t s o f s i m i l a r environments v a r i o u s r e a l and  alleged  s p e c i e s o f Hygrohypnum have responded t o the f l u c t u a t i n g water l e v e l s by assuming a myriad o f forms which have b l u r r e d s p e c i e s l i m i t s .  I t i s to  be  expected then, t h a t the s p e c i e s c o n c e p t s h e l d by many workers have been badly  confused and  taxa have been Renauld generic  t h a t among the most v a r i a b l e s p e c i e s numerous: s u b s p e c i f i c  recognized.  (1883) r e v i s e d the European s p e c i e s and  c i r c u m s c r i p t i o n and  Brotherus  (1909  and  1925)  p r e s e n t e d a more p r e c i s e provided  s p e c i e s o r c e r t a i n groups o f s p e c i e s known t o those times.  The  keys to  the  genus has  been  2 t r e a t e d i n numerous r e g i o n a l f l o r a s Roth, 1905; Grout,  Dixon,  1931;  1896  and 1924;  Nyholm, 1965  a w o r l d wide b a s i s .  ( B r a i t h w a i t e , 1898;  Limpricht  B r o t h e r u s , 1923? Monkemeyer,  1904;  1927;  and o t h e r s ) , b u t i t has never been r e v i s e d  on  As a r e s u l t o f t h i s n e g l e c t many taxa have been des-  c r i b e d under d i f f e r e n t names, w h i l e o t h e r s o f d i v e r s e and  questionable  a f f i n i t i e s have been i n c l u d e d w i t h i n the genus, a p p a r e n t l y f o r the r e a s o n t h a t the p l a n t s o c c u r e d on r o c k s i n streams. g e n e i t y prompted Loeske  sole  This generic hetero-  (1910) t o r e f e r t o Hygrohypnum as a " b i o l o g i s c h e  Mischgattung." The o b j e c t i v e s o f t h i s monograph have been t o : 1.  a s s e s s a l l o f the  d e s c r i b e d s p e c i e s t o c i r c u m s c r i b e a more n a t u r a l genus, 2.  discern stable  m o r p h o l o g i c a l c h a r a c t e r s w i t h which t o d i s t i n g u i s h s p e c i e s , 3.  provide  keys t o and d e s c r i p t i o n s , i l l u s t r a t i o n s and maps o f a l l the r e c o g n i z e d s p e c i e s and 4.  put i n t o usuable d e s c r i p t i v e form an assessment o f the  v a r i a t i o n t h a t may To a c c o m p l i s h ways.  l  :  be encountered  i n each s p e c i e s .  these g o a l s Hygrohypnum has been i n v e s t i g a t e d i n t h r e e  F i e l d work was  conducted  a s s e s s the e c o l o g i c a l amplitude  throughout  North America i n o r d e r t o  o f the genus and t o g a i n f i e l d  to a s s e s s n a t u r a l v a r i a b i l i t y i n the r e c o g n i z e d t a x a . d i v e r s e l o c a l i t i e s from throughout environment t o determine 3.  Ten s p e c i e s from  North America were grown under u n i f o r m  the s t a b i l i t y o f c e r t a i n m o r p h o l o g i c a l c h a r a c t e r s .  More than 2,200 herbarium  microscope  2.  experience  speciments  were d i s s e c t e d and made i n t o  s l i d e preparations f o r c r i t i c a l  13,000 specimens were examined and  study.  annotated.  Subsequently  more than  2a . '  Many D a r w i n i a n s y s t e m a t i s t s h a v e e x p r e s s e d t h e v i e w t h a t a c l a s s i -  fication  system should r e f l e c t  g o a l , but  unfortunately  existence of best.  In  absence of  to  truly  H o w e v e r , when t h e o n l y adult  desirable  depends on  the  fragmentary  relationships  at  among e x -  or o c c a s i o n a l l y e s t a b l i s h e d from data  morphology  it  is  understand the b i o l o g i c a l  species.  too o f t e n i s  breeding systems, development,  from e x t e r n a l ,  a very  phylogeny  a f o s s i l record the  t a n t s p e c i e s may b e i n f e r r e d  phytogeography.  This i s  an u n d e r s t a n d i n g o f  a f o s s i l record, which a l l  the  karyology,  phylogeny.  chemistry,  a v a i l a b l e data are  relationsships  As a consequence I h a v e n o t  ecology  very d i f f i c u l t ,  '  if  on  and  derived  not  impossible  among a g r o u p  of  s p e c u l a t e d on the phylogeny  of  Hygrohypnum. The s e q u e n c e i n w h i c h t h e s p e c i e s o f H y g r o h y p n u m a r e p r e s e n t e d t h i s work r e p r e s e n t s variation. i n adult in  fig.  a s e r i e s of  The d e s i g n o f  morphology.  this  overlapping patterns  sequence serves only  The s e q u e n c e o f  81 o n p a g e 4 0 0 .  species i s  of  to  in  morphological  elucidate  graphically  The m o s t i m p o r t a n t m o r p h o l o g i c a l  trends  represented  trends  a r e as  follows; 1.  Leaves very broad to  2.  Leaves e x c l u s i v e l y s t r a i g h t  3.  A l a r c e l l s mostly  4.  Stem anatomy e x h i b i t i n g medulla of  larger,  narrow.  undifferentiated  thinner-walled  Plants mostly  autoicous  and/or  to v a r i o u s l y  a cortex of  a more v a r i a b l e p a t t e r n ..of 5.  to s t r a i g h t  small,  cells  falcate. differentiated.  thiek-walled~cells,  and a c e n t r a l  strand  a to  differentiation.  t o p l a n t s w i t h a more v a r i a b l e  sexuality.  3 THE  TAXONOMIC HISTORY OF HYGROHYPNUM Progress  i n t h e study o f mosses has been dependent upon t e c h n i c a l  advances i n m i c r o s c o p y .  The i n a b i l i t y t o observe m i c r o s c o p i c  caused e a r l y workers t o employ v e r y broad sequence, p l e u r o c a r p o u s  g e n e r i c concepts.  features As a con-  mosses, e x c l u s i v e o f F o n t i n a l i s , were o r i g i n a l l y  t r e a t e d i n t h e a l l i n c l u s i v e genus Hypnum.  Hedwig  (1801) was t h e f i r s t  to use m i c r o s c o p i c  f e a t u r e s i n c l a s s i f y i n g mosses and w i t h h i s work com-  menced t h e g r a d u a l  s u b d i v i s i o n o f t h e genus Hypnum sensu l a t u i n t o many  s m a l l e r , more n a t u r a l genera. In t h e f o l l o w i n g d i s c u s s i o n t h e taxonomic h i s t o r y o f Hygrohypnum i s t r a c e d t o t h r e e Hedwigian s p e c i e s and i s then f o l l o w e d through t h e desc r i p t i o n o f a d d i t i o n a l s p e c i e s and t h e i r subsequent treatment s e c t i o n a l and s u b g e n e r i c  groupings  i n various  o f Hypnum o r i n such segregate  genera  as Limnobium, Stereodon, Amblystegium, C a l l i e r g o n and u l t i m a t e l y Hygrohypnum .  In t h e f o l l o w i n g d i s c u s s i o n i t s h o u l d be understood  t h a t the  c l a s s i f i c a t i o n s a s c r i b e d t o each c i t e d author have been e d i t e d ao as t o d e a l o n l y w i t h those taxa t h a t a r e c o n s i d e r e d t o be modern s p e c i e s o f Hygrohypnum o r a synonym * t h e r e o f . The o r i g i n s o f t h e genus c a n be t r a c e d t o t h r e e s p e c i e s t r e a t e d by Hedwig w i t h i n the b r o a d l y c o n c e i v e d Muscorum Hedwig  genus Hypnum Hedw.  In h i s  Species  (1801) s u b d i v i d e d Hypnum i n t o e i g h t numbered groups t o  which he gave n e i t h e r f o r m a l nor h i e r a r c h i c a l r a n k i n g .  As c r i t e r i a f o r  d e l i m i t i n g h i s e i g h t groups Hedwig emphasized l e a f symmetry and t h e v a r i o u s a t t i t u d e s t h e l e a v e s e x h i b i t e d a l o n g t h e stems. a l s o used, b u t were s u b o r d i n a t e  Branching  to leaf characters.  the taxa i n c l u d e d i n these e a r l y groupings  p a t t e r n s were  By modern  standards,  are of diverse r e l a t i o n s h i p s .  However, i n terms o f t h e c h a r a c t e r s used, Hedwig's groups were  reasonably  4 sound,  Hedwig: (recognized and c l a s s i f i e d Hypnum a l p e s t r e , H. m o l l e and 1  H. l u r i d u m as f o l l o w s :  Group I I I , F o l i i s r e c t i s , a e q u a l i t e r e t a r e t e i m b r i c a t i s , teretibus).  (ramis  Hypnum a l p e s t r e Sw. ex Hedw.  Group IV, F o l i i s r e c t i s , a e q u a l i t e r e t l a x e incumbentibus, ramis pinnatis.  Hypnum m o l l e D i c k s . Hedw.  Group V I I I , F o l i i s s e c u n d i s ,  Bridel  incurvis.  Hypnum l u r i d u m Hedw.  (1801) r e c o g n i z e d Hypnum p a l u s t r e Huds, ex B r i d . , and H. m o l l e  D i c k s . , b u t d i d n o t t r e a t H, a l p e s t r e Sw.  L i k e Hedwig, B r i d e l  subdivided  Hypnum i n t o e i i j h t groups, b u t he a p p l i e d t o them f o r m a l names a p p a r e n t l y o f s e c t i o n a l rank.  In d i r e c t c o n t r a s t t o Hedwig, B r i d e l p l a c e d g r e a t e r weight  on b r a n c h i n g p a t t e r n i n d e l i m i t i n g h i s groups and r e l e g a t e d l e a f form and o r i e n t a t i o n upon t h e stems t o s u b o r d i n a t e of  roles.  Bridel's  classification  C u r r e n t l y r e c o g n i z e d s p e c i e s o f Hygrohypnum i s as f o l l o w s :  Group V, s e c t i o n S c u i r o i d e a , S u r c u l o c o n f e r t i m ramoso. subgroup 2, f o l i i s  secundis r e f l e x i s c o n t o r s i v e . Hypnum p a l u s t r e Huds. ex B r i d ,  Group V I I I , s e c t i o n Polymorpha, Surculo vage ramoso„ f o l i i s d i r e c t i o n ^ ; aequali. subgroup 4, Amphibia  Bridel  Hypnum m o l l e ,  Dicks.  (1812) m o d i f i e d h i s e a r l i e r c l a s s i f i c a t i o n system a l o n g t h e  l i n e s o f Hedwig's and s u b d i v i d e d Hypnum i n t o n i n e groups.  I h e s e groups and  a number o f t h e i r s u b d i v i s i o n s were g i v e n formal names a t s e c t i o n a l rank. Through a t y p o g r a p h i c a l e r r o r t h e number 2 was used t w i c e i n d e n o t i n g t h e groups and consequently  t h e n i n e groups nufabered  only eight.  In t h e new  system B r i d e l p l a c e d g r e a t e r v a l u e on l e a f c h a r a c t e r s and r e l e g a t e d b r a n c h -  5 ing  features to a lesser role.  B r i d e l ' s c l a s s i f i c a t i o n as a p p l i e d t o  Hygrohypnum i s g i v e n below: 1 Group 2 Section Illecebra, f o l i i s  i m b r i c a t i s appressa v e l appresso  p a t e n t i u s c u l u s , ramis p i n n a t i s v e l s u b p i n n a t i s , r a m u l i s teretibus obtusis.  Hypnum a l p e s t r e Sw. and Hypnum m o l l e D i c k s .  Section Cuspidata, f o l i i s  i m b r i c a t i s appressa v e l appresso  p a t e n t i u s c u l u s , ramis p i n n a t i s v e l subpinnato, compressisculus a c u t i s . Group 7, f o l i i s  ramulis  Hypnum negleetum B r i d .  secundis c i r c i n n a t o - f a l c a t i s  Section Lurida, caulibus  c o n € e r t i m ramosus. Hypnum l u r i d u m Hedw. and Hypnum s u b s p h a e r i c a r p o n  It  Brid.  seems apparent here t h a t B r i d e l r e c o g n i z e d t h e s i m i l a r i t i e s o f  Hedwig*s treatment o f Hypnum a l p e s t r e Sw. and Hedwig's and h i s own t r e a t ment o f Hypnum m o l l e f o r he i n c l u d e d them both i n s e c t i o n The  c l a s s i f i c a t i o n p r e s e n t e d by B r i d e l  from t h e system o f 1812.  (1827) was e s s e n t i a l l y unchanged  He d i d c o r r e c t t h e e r r o r i n the group numberings  and he e x c l u d e d Hypnum negleetum B r i d . in  Illecebra.  The most s i g n i f i c a n t change o c c u r c e d  t h e d i v i s i o n o f Hypnum i n t o two subgenera  based on endostomial c h a r a c t e r s .  Subgenus Hypnum, peristome i n t e r i o r i s c i l i i s  perforatis.  Subgenus Stereodon, p e r i s t o m e i n t e r i o r i s c i l i i s  imperforatis.  Hypnum m o l l e , H. a l p e s t r e , H. p a l u s t r i s and H. s u b s p h a e r i c a r p o n were a l l i n c l u d e d i n t h e subgenus  Stereodon.  1  I n d i c a t e s t h e second  and e r r o n e o u s l y numbered group 2  6 Sprengel  (1827) s u b d i v i d e d  was s u b d i v i d e d  Hypnum i n t o t h r e e groups, each o f which  i n t o a number o f s m a l l e r groups.  L i k e Hedwig and B r i d e l ,  S p r e n g e l used l e a f symmetry and l e a f a t t i t u d e upon t h e stem as c h a r a c t e r s o f p r i m a r y importance.  However, he appears t o have been t h e f i r s t t o  employ c o s t a s t r u c t u r e , smoothness o r p a p i l l o s i t y o f t h e l e a f c e l l s and t o o t h i n g o f t h e l e a f margin as c l a s s i f a c t o r y c r i t e r i a .  U s i n g these f e a t u r e s ,  S p r e n g e l d i v i d e d Hypnum i n t o t h r e e numbered groups o f u n s p e c i f i e d  Group 2, F o l i i s  secundis,  * nervoffis. ** F o l i i s  rank.  Hypnum spfaaericarpon  secundis e n e r v i s .  Spreng.  Hypnum p a l u s t r e  (no a u t h o r i t y  designated) Group 3, F o l i i s d i r e c t i o n e v a r i u s * F o l i i s directione variis Foliis  integerrimus,  e r e c t o - p a l u l i s , Nervo evanido, Foliis  fetisque  laevibus.  Hypnum m o l l e D i c k s , and Hypnum neglectum B r i d . Hypnum s p h a e r i c a r p o n carpon B r i d .  was an o b v i o u s m i s s p e l l i n g o f Hypnum  I t i s worth n o t i n g  t h a t where B r i d e l r e c o g n i z e d  subsphaerisufficient  s i m i l a r i t y between Hypnum m o l l e D i c k s , and H. a l p e s t r e Sw. t o i n c l u d e them w i t h i n t h e same g r o u p i n g , S p r e n g e l u n i t e d t h e two under H. m o l l e  Dicks.  Hiibner ^1833) d i v i d e d Hypnum i n t o s e v e r a l s e c t i o n s , f o r which he provided  elaborate  descriptions.  A l p e s t r i a he i n c l u d e d o n l y taxa hypnum .  H i s treatment i s unique f o r w i t h i n t h a t have s i n c e been i n c l u d e d w i t h i n  section Hygro-  H i s treatment i s as f o l l o w s :  Section A l p e s t r i a C a u l e f a s c i c u l a t i m ramoso, ramis c o n f e r t i s e r e c t i s s i m p l i c i b u s  t u r g i d o - t e r e t i b u s , f o l i i s dense i m b r i c a t i s b a s i n e r v i b u s ,  setis  laevibus. Hypnum m o l l e  Dicks.  Hypnum a l p e s t r e Sw., Hypnum p a l u s t r e L i n n . Hypnum s p h a e r i c a r p o n  Spreng.  Hampe (1837) d i v i d e d Hypnum i n t o two apparent subgenera which he called  Isothecium B r i d , and Stereodon e t Hypnum B r i d .  were s u b d i v i d e d  These two groups  f u r t h e r , b u t Hampe d i d n o t i n d i c a t e t h e c r i t e r i a he used.  H i s treatment i s unique f o r he t r e a t e d Hypnum l u r i d u m  Hedw. and H. s p h a e r i -  carpon B r i d . as v a r i e t i e s o f Hypnum p a l u s t r e L i n n , and p l a c e d Hypnum a l p e s t r e Sw, as a v a r i e t y o f De N o t a r i s his  molle  Dicks.  (1838) p r o v i d e d  a t r e a t m e n t s i m i l a r t o t h a t o f Hiibner,  In  Hypnum s e c t . P a l u s t r i a De N o t a r i s c l e a r l y i n c l u d e d t a x a t h a t have a l l  become r e c o g n i z e d De,Notaris"  s p e c i e s o f Hygrohypnum o r synonyms o f r e c o g n i z e d  species,  system i s g i v e n below:  Hypnum s e c t . P a l u s t r i a De Not. C a u l e r e p e n t e s c o n f e r t i m r a m o s i , f o l i a i m b r i c a t a plerumque secunda, s u b f a l c a t i v e concaviscula, tenuieve  seta  laevi.  Hypnum p a l u s t r e B r i d . Hypnum s u b s p h a e r i c a r p o n B r i d . Hypnum m o l l e Schimper  (1853, i n Bruch, Schimper and Gumbel) p r e s e n t e d  c e p t u a l s y n t h e s i s o f t h e modern genus Hygrohypnum. luridum  Dicks. the f i r s t  con-  Schimper e x t r a c t e d Hypnum  Hedw,, H. m o l l e Hedw, and H_. a l p e s t r e Hedw. from Hypnum and p l a c e d  them i n t h e new genus Limnobium Schimp.  Hypnum s u b s p h a e r i c a r p o n B r i d . and  H. negleetum B r i d . were shown t o be v a r i a n t s o f H. l u r i d u m  Hedw. and u n i t e d  8 with  i t . Simultaneously,  L. a r c t i c u m Schimp. L. ochraceum  he d e s c r i b e d Limnobium norvegicum Schimp.  Schimper  (Turn, ex W i l s . )  and  (1855) added Limnobium eugyrium Schimp, Schimp.  In so d o i n g ,  and  Schimper brought t o -  g e t h e r a group o f s p e c i e s e x h i b i t i n g such a d i v e r s i t y o f form as to have widely  spaced t o crowded branches, s t r a i g h t , l o o s l y appressed to  o r f a l c a t e - s e c u n d l e a v e s , p o i n t e d o r o b t u s e branch t i p s and double o r s i n g l e costae. shared  short  However, Schimper determined t h a t these  spreading and taxa  a s i m i l a r a r e o l a t i o n , a Seldom excavated r e g i o n o f p o o r l y d e f i n e d  a l a r c e l l s , a b r o a d l y o v a t e - l a n c e o l a t e o r o r b i c u l a r l e a f shape and h a b i t a t i n and Wilson  a common  around streams.  (1855) r e t a i n e d a more c o n s e r v a t i v e view and  s p e c i e s o f Hygrohypnum as Hypnum. upon l e a f symmetry and  treated present  day  He d i v i d e d Hypnum i n t o t h r e e groups based  l e a f a t t i t u d e upon the stem.  He  treated various  s p e c i e s o f Hypnum as f o l l o w s : Section I, leaves + spreading  i n everyway.  Section A l p e s t r i a A.  Stems c r e e p i n g ,  i r r e g u l a r l y branched; o r o c c a s i o n a l l y  arched  *** Leaves r o u n d i s h ,  pinnate,  e n t i r e , two  r a t h e r obtuse,  nerved, n e r v e l e s s . Hypnum m o l l e  Dicks.  Hypnum p a l u s t r e D i l l . , Hypnum a r c t i c u m SSction I I , leaves Section B.  Linn.  Sommerf.  secund.  Cuppressiformia Stem procumbent, + e n t i r e , f r u c t i f i c a t i o n s near  the  base; l e a v e f a l c a t e - s e c u n d , e n t i r e o r s e r r u l a t e , n e r v e l e s s o r two  nerved a t the base, * Leaves acpuminate.  9 Hypnum ochraceum T u r n . Commencing w i t h S u l l i v a n t (1856) t h e genus Limnobium Schimp. was subjected (1860  to a variety o f interpretations.  S u l l i v a n t (1856) and Schimper  and 1876) i n t e r p r e t e d t h e taxon as a subgenus o f Hypnum.  (1883), B o u l a y  (1884), Husnot  (1894) and Dixon  Limnobium as a s e c t i o n o f Hypnum.  Mitten  Renauld  (1896 and 1924) t r e a t e d  (1864) d i f f e r e d s l i g h t l y t r e a t -  i n g S e c t i o n Limnobium w i t h i n Stereodon B r i d . Renauld  (1883) c r i t i c a l l y r e v i s e d t h e European s p e c i e s as Hypnum  s e c t i o n Limnobium. the s e c t i o n .  I n so d o i n g he o f f e r e d some s i g n i f i c a n t i n s i g h t i n t o  He o b s e r v e d t h a t t h e s e c t i o n c o u l d be s u b d i v i d e d  groups based on l e a f morphology.  i n t o two  He c h a r a c t e r i z e d t h e groups as f o l l o w s :  Group 1, L e s f e u i l l e s s o n t e l a r g i e s o v a l e s ou s u b o r B i c u l a i r e , contractees  souvent  k l a base, a r r o n d e s ou sommet ou b r i e v i m e n t e t  obtusement a p i c u l e e s . Hypnum d i l a t a t u m Wis. Hypnum m o l l e  Dicks.  Hypnum alpinum Sch. 5*  a r c t i c u m Sommerf,  H. g o u l a r d i i Sch. H. o b t u s i f o l i u m Hook. Group 2, Les f e u i l l e s sont oblongues ou o b l o n g u e s - l a n c e o l e e s , ou moms r e t r e c i e s - a c u m m e e s . Hypnum p a l u s t r e L. H. p o l a r e  Lindb,  H, a l p e s t r e Sw. H. eugyrium B.S.G. H. l u s i t a n i c u m S c h .  plus  10 H. micans Wils. H. ochraceum Turn. De Bat (1885) presented a r e c l a s s i f i c a t i o n o f the genus Hypnum w i t h i n which modern species of Hygrohypnum were treated i n two  sections.  His systems p a r a l l e l e d that of Renauld (1883) emphasizing groups of species e x h i b i t i n g s t r a i g h t , o r b i c u l a r leaves which he c a l l e d section Eu-Limnobium and those species bearing s t r a i g h t o r f a l c a t e , more or less leaves, which he placed i n section Hetero-Limnobium.  lanceolate  De Bat's system i s  as follows: Eu-Hypnum a.  Rectifolia  re  1  Section, a r a m i f i c a t i o n i r r e g u l i e r e (correspond aux s i m p l i c i a )  SECTION Eu-LiMhobia A.  F e u i l l e s c i r c u l a i r e s ou o v a l e s - c i r c u l a i r e s ou avec obtus pondent aux o b t u s i f o l i a ) H. alpinum H. g o u l a r d i i H.  alpestre  H.  dilatatum  H. molle H. norvegicum H. d e f l e x i f o l i u m (= Scorpiurium) B.  F e u i l l e s ovales-acuminees peu ou appeine homotropes H.  lusitanicum  H. polare H. micans  (corres-  11 B. C u r v i f o l i a - F e u i l l e s ovales-oblongues  ou o v a l e s l a n c e o l e e s ,  nettement homotropes ou i n c u r v e e s . SECTION Hetero-Limnobia H. ochraceum H. subenerve H. eugyrium H. p a l u s t r e L i n d b e r g h 187 2) observed  t h a t Limnobium Schimp, was a l a t e r homonym  o f Limnobium R i c h . , a genus o f the H y d r o c h a r i t a c e a e .  As a s u b s t i t u t e f o r  Limnobium Schimp, L i n d b e r g c o i n e d t h e name Hygrohypnum L i n d b . Lindberg  (1879) reduced  Amblystegium Schimp,  Hygrohypnum L i n d b . t o s u b g e n e r i c  Braithwaite  Later,  s t a t u s under  (1898) t r e a t e d Hygroftfrpnum L i n d b . a s a  s e c t i o n o f Amblystegium and L i m p r i c h t  (1904) p l a c e d Hygrohypnum L i n d b . as  subgenus i n Hypnum. Kindberg  (1894) c o n t i n u e d  t o use Limnobium f o r c e r t a i n c u r r e n t l y  r e c o g n i z e d s p e c i e s o f Hygrohypnum and a t the same time e l e v a t e d Hypnum subg. C a l l i e r g o n S u l l . t o genus.  I n C a l l i e r g o n Kindberg  now r e f e r r e d t o C a l l i e r g o n and Hygrohypnum. h i s concept  Kindberg  treated species  (1897) e l a b o r a t e d  o f C a l l i e r g o n and d e s c r i b e d a number o f subgenera and s e c t i o n s  which i n c l u d e d s p e c i e s o f modern C a l l i e r g o n , Drepanocladus, Sematophyllum and Hygrohypnum.  Scorpidium,  Modern s p e c i e s o f Hygrohypnum were d i s -  t r i b u t e d i n Kindberg's t r e a t m e n t ^ o f  C a l l i e r g o n as f o l l o w s :  C a l l i e r g o n subg. Limnobium A.  Capsule not annulate,  Leaves e n t i r e , I I .  Leaves o f stem n o t  f a l c a t e , those o f t h e branches f a l c a t e , a l a r ^ c e l l s n o t l a r g e , c o s t a a b r e v i a t e sometimes double o r i n d i s t i n c t . capsule s h o r t i s h .  Stems c r e e p i n g o r p i n n a t e .  Pedicel of  12 Calliergon palustre  (L.) Kindb.  C a l l i e r g o n subg. Pseudo-Limnobium Kindb. B. ..Capsule a n n u l a t e , Leaves sometimes d e n t i c u l a t e , b. stems not p i n n a t e , sometimes r a d i c u l o s e , a l a r c e l l s  generally  distinct. S e c t i o n B a d i i f o r m i a Kindb. Leaves n o t d e c u r r e n t , those o f branches o f t e n  falcate.  D i o i c o u s and v e r y r . f r u i t i n g . r  C a l l i e r g o n " p o l a r e -(Lindb.)' Kindb.  S e c t i o n O c h r a c e i f o r m r a Kindb. bb.  C o s t a g e n e r a l l y s h o r t and i n d i s t i n c t , 2 .  Leaves  i  .....entire o r n e a r l y so, d e c u r r e n t and d i s t a n t , those o f branches o f t e n f a l c a t e , stems e r a d i c u l o s e , dioicous rarely  fruiting.  C a l l i e r g o n ochraceum  (Turn, ex W i l s . ) Kindb.  C . - t o r r e n t i s Kindb. C. g o u l a r d i i  (Schimp.)  Kindb.  S e c t i o n M o l l i f o r m i a Kindb. bb.  C o s t a g e n e r a l l y s h o r t and i n d i s t i n c t ,  3...Leaves n o t o r  i n d i s t i n c t l y d e c u r r e n t , sometimes d e n t i c u l a t e , crowded Often  generally  and n o t f a l c a t e , Stem not c r e e p i n g v s . r a d i c u l o s e .  monoicous. C a l l i e r g o n eugyrium C_. a r c t i c u m  (B.S.G.)"Kindb.  (Sommerf.) Kindb.  C_. m o l l e (Dicks.) Kindb. C_. a l p e s t r e  (Sw.)  Kindb.  C. c o l u m b i c o - p a l u s t r e C M .  §_t Kindb.  13 C_. c i r c u l i f o l i u m C M .  ert, Kindb.  C. submolle Kindb. S e c t i o n Montaniforme  Kindb.  Leaves s m a l l , sometimes d e c u r r e n t , u s u a l l y d e n t i c u l a t e , those o f the branches o f t e n f a l c a t e .  Stems c r e e p i n g v s .  monoecious. C a l l i e r g o n p s e u d o - a r c t i c u m Kindb. C_. montanum (Lindb.) Kindb. C_. pseudo-montanum Kindb. C. v i r i d u l u m Paris cations.-  (1905) and Roth  Loeske  (Hartm.) Kindb.  (1905) used Limnobium Schimp. i n t h e i r p u b l i -  (1903) recognized^: the p r i o r i t y o f Hygrohypnum L i n d b . ,  b u t i t was n o t u n t i l B r o t h e r u s (1909) p u b l i s h e d volume 1 o f Die N a t u r l i c h e n P f l a n z e n f a m i l i e n t h a t Hygrohypnum came i n t o g e n e r a l usage.  Brotherus also  determined t h a t the a f f i n i t i e s o f Hygrohypnum were n o t w i t h Hypnum, b u t w i t h h i s new  hypnaceous  s u b f a m i l y , the A m b l y s t e g i a e .  B r o t h e r u s (1923)  e l e v a t e d the s u b f a m i l y t o the rank o f f a m i l y . Grout  (1931) a l s o used the s u b f a m i l y A m b l y s t e g i a e .  More i m p o r t a n t l y ,  he determined t h a t Hygrohypnum c o u l d be s u b d i v i d e d i n t o f o u r groups. groups were g i v e n f o r m a l names, b u t no taxonomic rank. d i s t i n g u i s h e d as Group I ,  These  H i s groups were  follows:  Palustreformes Leaves more than t w i c e as l o n g as broad, v e r y concave w i t h the upper margins i n f o l d e d , e n t i r e o r s l i g h t l y  serrulate  above, a l a r c e l l s numerous. Group I I , A r c t i c i Leaves l e s s than twice as l o n g as wide, o f t e n n e a r l y  circular  i n o u t l i n e , more o r l e s s concave, e n t i r e o r s l i g h t l y  serrulate  a t apex, margins u s u a l l y p l a n e .  14 Group I I I , Montani Stems w i t h o u t a c e n t r a l s t r a n d , l e a v e s acuminate t o apiculate, d i s t i n c t l y lower margins  s e r r u l a t e above, s l i g h t l y d e c u r r e n t ,  reflexed.  Group IV, O c h r a c e i An o u t e r l a y e r o f stem c e l l s e n l a r g e d and h y a l i n e , ing  a hyaline  sheath.  Wijk, Margadant and F l o r s c h u t z  (1962) p r e s e n t e d a l i s t  i s h e d s p e c i e s and the s u b s p e c i f i c t a x a o f Hygrohypnum. emended t h a t l i s t  as f o l l o w s :  Hygrohypnum a l p e s t r e H. alpinum  (Lindb.)  (Hedw.)_Loesk. Loesk.  var.  v i r i s c e n s Amann  var.  t s u r u g i z a n i c u m (Card.) Nog.  H. aureum Herz. H. b r a s i l i a n s e H B E Z .  H_. c a u s s e q u i (Ren. et-Gard.)  Card.  H. c i r c i n a t u m Herz. H. c l o s t e r i  (Aust.)  Grout  H. c o r d i f o l i u m Okam. H_. coreanum Card. H. d i l a t a t u m (Wils.) Loesk. var.  c a l l i n e u r u m Amann  var.  d u r i u s c u l u m (Limpr.) Amann  H. d o i i Sak. var.  simplex Sak.  H. e l l i p t i c u m  Ther.  H. e n t o d o n t o i d e s  form-  ( B r o t h , e t Par.) B r o t h .  of a l l publ-  Wijk e t a l . (1969)  15 H_. eugyrium (Schimp.) Broth. ssp.  subeugyrium (Ren. et Card.) Grout  var. mackayi (Schimp.). Broth. var.  miquelonense (Ren. et Card.) Grout  var.  nervosum  var.  occidentale (Card. et. Ther.) Grout  (Roell) Podp.  H. fontinaloides Chen H. g l a c i a l e Warnst. H. hedbergii P. Vard. H. liukiuense Sak. H. luridium (Hedw.) Jenn. ssp. pseudomontanum (Kindb.) Wijk et Marg. var. crassinervium(Baur.).Podp. var.  e h l e i (Am.)  Wijk et Marg.  var.  malacocaulon (Herz.) Podp.  var.  obtusatum Podp.  var. pseudochraceum  * nom.  nud.  (Roth) Podp.  var.  subsphaericarpon (Brid.) C. Jans, i n Podp.  var.  tenellum (Schimp.) Podp.  H. lusitanicum (Schimp.) Corb. H. mizushimae Sak. H. molendinarium (Lam. etCand.) Wijk et Marg. H. molle (Hedw.) Losek. ssp.  b e s t i i (Ren. et Card.) Grout  var.  b e s t i i (Ren. et. Card.) Hab.  var.  japonicurn Sak.  var. pyrenaicum (Ren.) Podp. var.  schimperianum (Lor.) Loesk.  H. montanum (Lindb.) Broth. H. n i c h o l s i i Grout H. norvegicum (Schimp.) Amann H. novae-caesarae (Aust.) Grout H. ochraceum (Wils.) Loesk.  16 H. p e l i c h u e n s e H. p e r u v i e n s e  Williams Williams  H. p o e c i l o p h y l l u m D i x . H. p o l a r e  (Lindb.)  Loesk.  v a r . pseudostramineum (Lindb.) H. purpurascens  Broth.  H.  in Lilj.)  smithii  (Sw.  Podp.  Broth.  v a r . goulardii-;:(;S chimp.) - Wijk e t Marg. H.  styriacum  H.  szaferi  H.  tenquendamense Herz.  H.  validum  Bartram  (Limpr.)  Broth.  Szaf.  Herz.  (1965) added the most r e c e n t l y d e s c r i b e d s p e c i e s , a South  American p l a n t which he c a l l e d Hygrohypnum f a l c i f o l i u m B a r t .  17 METHODS OF THE MORPHOLOGICAL ANALYSIS The apparent inadequacy o f those c h a r a c t e r s t r a d i t i o n a l l y employed i s d i s t i n g u i s h i n g s p e c i e s o f Hygrohypnum i n d i c a t e d t h e need f o r a complete reassessment o f a l l p o t e n t i a l m o r p h o l o g i c a l c h a r a c t e r s .  Table 1 presents  a l i s t o f e i g h t y - s i x m o r p h o l o g i c a l c h a r a c t e r s which were examined d u r i n g t h i s study.  The assessment  conducted by 1.  o f t h e taxonomic  v a l u e o f these f e a t u r e s was  t h e t r a d i t i o n a l examination o f herbarium  specimens and  by 2.  c u l t i v a t i n g l i v i n g specimens under c o n d i t i o n s o f u n i f o r m e n v i r o n -  ment.  I n those cases where m a t e r i a l c o l l e c t e d i n t h e f i e l d bore mature  sporophytes o r where c u l t i v a t e d m a t e r i a l produced  sporophytes, the spores  so d e r i v e d were germinated and a l l o w e d t o produce  l e a f y gametophores.  Thus,  i n some c a s e s i t was p o s s i b l e t o compare gametophores grown from spores, gametophores d e r i v e d from new v e g e t a t i v e growth from a d u l t p l a n t s and the c o n t r o l herbarium  specimens.  18 Table  1  MORFHOLOGICAfc FEATORE8 HUBJ8CT TO RE* LYSIS Habit-.  P e r i c b a e t l E l leaves con't.  Color  plication  Stem  Areolation Length  Papillosity  Branching  pattern  P e i i g o n i a l leaves  Cross-section  Shape  Cortical c e l l color  Size  Fseudcpavaphyllia  Concavity  Orientation to the substrate phizoids  Costa Margins  Position  Areolation  Color  Sporophyte  C a l l wail d e t a i l  Seta  Frequency  Length  Abundance  Color  Leaves  Cross-section  Shape  Twisting  Siie  Surface texture  Symmetry  Capsule  Concavity  Shape  Apex  Size Color Curvature  Exothecial c e l l s  Toothing  Shape  Decurrency  Size  A t t i t u d e upon the stem  Wall thic-iriess  Changes i n appearance between the  Apophysal d i f f e r e n t i a t i o n  wet and dry c o n d i t i o n  C o n s t r i c t i o n below the nouth  Arcolaticr.  Stomates  C e l l shape  Numbers  C e l l size  Position  C e l l wall thickening'  Struct'ire  C e l l well d i s c o l o r a t i o n  Operculum  Cell vail pitting Papiilosity Regional  variation  Annulus • Peristome Exostome  Apical  Color  Median  Segment length and width  Marginal  Striations  Basal  Papillosity  Alar  Endostcme  Sexuality  Color  Autoicous, d i o i c o u s , pseudo-parolccus  Basal membrane height  P e r i c h a e t i a l leaves  Segment length anc width  Outer, middle and inner leaves General  Segment s p l i t t i n g  features  Papillosity  Si":ape  Cilia  Si-^e  Development  Attitude  Numbers  Apex Margins Curvature Toothing Coeta  Diameter Color VWll morphology  19 METHODS OF CULTIVATION A l l c u l t u r e d p l a n t s , e x c e p t those grown from spores, were m a i n t a i n e d i n a r e f r i g e r a t e d growth room h e l d between 6 ° and 9 ° C. grown on 4 l a y e r s o f paper towel d i s k s and moistened of  Hoagland's c u l t u r e medium (Hoagland  A l l p l a n t s were  w i t h a 50% s o l u t i o n  and A r n o l d , 1938).  Most p l a n t s  were grown i n 90 mm X 19 mm p l a s t i c p e t r i d i s h e s , w h i l e c e r t a i n  larger  specimens were grown i n 155 mm X 65 mm covered p l a s t i c d i s h e s .  The c u l -  t u r e s were i l l u m i n a t e d by 6 banks o f 60 i n c h l o n g c o o l w h i t e bulbs. tional  fluorescent  Each bank bore 4 b u l b s h e l d a t 38 i n c h e s above t h e p l a n t s . i l l u m i n a t i o n was p r o v i d e d by twelve 60 watt  e q u i d i s t a n t among t h e banks o f f l u o r e s c e n t l i g h t s .  Addi-  incandescent bulbs placed The t o t a l i l l u m i n a t i o n was  530 f o o t - c a n d l e s i n a 16/8 p h o t o p e r i o d . C u l t u r e s grown on paper unicellular  toweling  were f r e q u e n t l y contaminated by  green a l g a e and/or O s c i l l a t o r i a .  t r o l l e d by p e r i o d i c a l l y  Such c o n t a m i n a t i o n was con-  washing t h e p l a n t s w i t h t a p water and g e n t l y s c r u b -  b i n g them w i t h a s t i f f a r t i s t ' s b r u s h .  When n e c e s s a r y s e v e r a l washed stems  were t r a n s f e r r e d t o new c u l t u r e d i s h e s . E x p e r i m e n t a l p l a n t s grown from spores were m a i n t a i n e d i n 90 mm X 20 mm g l a s s p e t r i d i s h e s i n 50% Hoagland's medium i n 2% D i f c o Bacto-Agar. were m a i n t a i n e d  They  i n a growth room a t 10° C on a 12/12 p h o t o p e r i o d .  METHODS OF PREPARATION OF MATERIAL FOR MICROSCOPE EXAMINATION A l l m a t e r i a l s were examined u t i l i z i n g employing  Hoyer's mounting medium.  The  (1954) so as t o use 50 c c o f gum a r a b i c .  microscope  slide preparations  Hoyer's was m o d i f i e d from Anderson  20  RESULTS OF THE  MORPHOLOGICAL ANALYSIS  Stem Anatomy The  s t r u c t u r e o f the stem, as seen i n c r o s s - ^ s e c t i o n , i s extremely  important i n a s s e s s i n g b o t h s i m i l a r i t i e s and  d i f f e r e n c e s among the  various  species. The  stem i s u s u a l l y d i f f e r e n t i a t e d i n t o t h r e e h i s t o l o g i c a l  but t h e r e may and  only, be  two  several concentric  walled  and  the c o r t e x  yellow  o r as many as f o u r .  U s u a l l y the e p i d e r m a l  to red-brown i n c o l o r .  T h i s zone o f c e l l s i s c a l l e d  I n s i d e the c o r t e x t h e r e i s a broad r e g i o n o f  l a r g e , t h i n - w a l l e d , u s u a l l y h y a l i n e c e l l s c a l l e d the m e d u l l a T h i s m e d u l l a r y r e g i o n may  d i s c o l o r w i t h age.  stem c r o s s - s e c t i o n a group o f s m a l l c e l l s may  and  cells  rows o f c e l l s i n s i d e the e p i d e r m i s a r e s m a l l , t h i c k -  (Fig. 1 a - b).  from the m e d u l l a .  regions,  A t the v e r y c e n t r e o f be a b r u p t l y  i n most s p e c i e s .  the  differentiated  These c e l l s are c a l l e d the c e n t r a l s t r a n d  i t i s r e g u l a r l y present  (Fig. 1 a - f ) .  (Fig. 1 a - f)  Hygrohypnum b e s t i i and  H.  subeugyrium are unusual f o r i n them the c e n t r a l s t r a n d i s o n l y weakly  de-  or  veloped  are. a b s e n t .  centEal strand  Hygrohypnum montanum i s remarkable because i t l a c k s a  entirely.  In Hygrohypnum ochraceum, H. p o l a r e and the stem i s d i f f e r e n t i a t e d from the a d j a c e n t dermis. walled  H.  eugyrium the e p i d e r m i s o f  c o r t e x and  i s called a  In H. ochraceum the h y a l o d e r m i s i s a b r u p t l y e n l a r g e d (Fig. 1 c ) .  and  hyalo-  thin-  In o l d e r p l a n t s the t h i n o u t e r w a l l i s u s u a l l y worn  away l e a v i n g o n l y the c a v i t y c r e a t e d by the i n n e r t a n g e n t i a l w a l l the r a d i a l w a l l s o f each c e l l . to t h a t o f H. ochraceum, but  The  h y a l o d e r m i s o f H,  the e n l a r g e d  and  polare i s s i m i l a r  c e l l s u s u a l l y do not  completely  encompass the stem  ( F i g . 1 d & e ) , i n which case the hyalodermis i s r e f e r r e d  to as i n c o m p l e t e .  A h y a l o d e r m i s i s most p o o r l y d i f f e r e n t i a t e d i n H.  eugyrium  21  and i s evident only because the outer tangential wall of the  epidermis  i s s l i g h t l y thinner and less pigmented than the r a d i a l and inner tangent i a l c e l l s walls (Fig. l n e ) .  22 Fig. 1 a - f.  V a r i a t i o n i n stem anatomy.  a.  K.  duriusculum  b.  H.  luridum  c.  H.  ochraceum  d. and e. f.  H. p o l a r e  H. eugyrium  Scale:  I  lOOum  i  24  L e a f Shape and  Symmetry  L e a f shape i s a v e r y o f Hygrohypnum.  important c h a r a c t e r  In the p a s t ,  e r i o n has been b e s e t w i t h two shape and iability  i n d i s t i n g u i s h i n g species  the use o f l e a f shape as a taxonomic s e r i o u s problems.  l e a f symmetry are v a r i a b l e .  In some s p e c i e s  leaf  A n a t u r a l consequence o f t h a t  i s t h a t the v a r i a b i l i t y i n l e a f shape and  can o v e r l a p w i t h t h a t o f a n o t h e r .  symmetry o f one  the terms used t o ' d e s c r i b e  this a  from the f r e q u e n t l y ambiguous a p p l i c a t i o n o f l e a f shape.  v a r i o u s attempts to s t a n d a r d i z e  and q u a n t i f y the d e s c r i p t i o n o f  l e a f shape seem t o have y i e l d e d l i t t l e agreement among b o t a n i s t s . o f the problem seems t o i n v o l v e the e x t e n t apex a f f e c t s the l e a f shape. problem.  species  A second problem i s the employment o f l e a f shape as  taxonomic c r i t e r i o n d e r i v e s  The  var-  I t i s u n d e r s t a n d a b l e , then, t h a t many  p e o p l e have confused many s p e c i e s by p l a c i n g too much r e l i a n c e on single character.  crit-  No  Much  to which the n a t u r e o f the l e a f  attempt has been made here t o r e s o l v e  this  F o r the purposes o f t h i s study, Figig 2 i l l u s t r a t e s the usage o f  the v a r i o u s d e s c r i p t i v e terms as a p p l i e d t o l e a f shape i n Hygrohypnum. The  examination o f numerous herbarium specimens has  shape w i t h i n a s p e c i e s v a r i e s w i t h i n b r o a d l y s i g n i f i c a n t that those species  shown t h a t l e a f  definable l i m i t s .  I t i s very  t h a t have been a v a i l a b l e f o r e x p e r i m e n t a l  study e x h i b i t the same b a s i c v a r i a t i o n i n l e a f shape when grown i n c u l t u r e as they do under n a t u r a l c o n d i t i o n s . Figures  3-6  d i r e c t l y compare the v a r i a b i l i t y i n l e a f shape o f  f r e q u e n t l y confused s p e c i e s . the most o r one species.  The  Based on d a t a from herbarium m a t e r i a l , several  As a s t e r i s k a t the base o f a l e a f i n d i c a t e s  o f the most f r e q u e n t l y encountered l e a f shapes f o r a  l e a v e s o f s p e c i e s grown under n a t u r a l and  t i o n s a r e compared i n F i g u r e s  7-9.  experimental  given condi-  Lodge (1959, 1960) was  showed t h a t  e n v i r o n m e n t a l l y induced.  Zales  i n Drepanocladus l e a f f a l c a t i o n (1973) a l s o e s t a b l i s h e d  i a b i l i t y of l e a f curvature i n P h i l o n o t i s . on  Similarly, culture  Hygrohypnum have shown l e a f f a l c a t i o n to be  v a l i d as a taxonomic c r i t e r i o n .  the  and  responses to e x p e r i m e n t a l  same stem o r on  in-  i n very s p e c i f i c locations,  culture. or  c o n t i g u o u s stems.  f a l c a t e l e a v e s on  In;culture,  pro-  produce s t r a i g h t  d i f f e r e n t , but  Hygrohypnum montanum produces s t r a i g h t and stem, but  studies  highly variable  c o n d i t i o n s Hygrohypnum eugyrium may  f a l c a t e l e a v e s on  var-  However, those s p e c i e s f r e q u e n t l y  ducing f a l c a t e leaves e x h i b i t v a r i a b l e Under n a t u r a l  the  b o t h H.  the  same  eugyrium  and  H.  and  e x p e r i m e n t a l c o n d i t i o n s Hygrohypnum s t y r i a c u m produces b o t h s t r a i g h t  and  falcate leaves.  montanum produce e x c l u s i v e l y  produce e x c l u s i v e l y  Certain  straight leaves.  Under both  specimens o f Hygrohypnum ochraceum which  circinate-caniculate  l e a v e s i n n a t u r e , respond  e x p e r i m e n t a l c o n d i t i o n s by p r o d u c i n g both c i r c i n a t e - c a n i c u l a t e and  s t r a i g h t and  natural  p l a n e ones.  Herbarium specimens r e v e a l  leaves  t h a t Hygrohypnum  l u r i d u m can produce specimens w i t h s t r a i g h t and/or f a l c a t e l e a v e s different plants,  on  sequences a l o n g the  d i f f e r e n t but same stem.  c o n t i g u o u s stems o r  E x p e r i m e n t a l d a t a on  l e s s c l e a r because t h e r e i s l i m i t e d m a t e r i a l has  been shown t h a t  straight or in  i n one  H.  available  to  in  on  alternating  luridum  are  f o r study.  It  c o l l e c t i o n non-contiguous stems, which bore  f a l c a t e l e a v e s i n n a t u r e u n i f o r m l y produced s t r a i g h t  leaves  culture. The  v a r i a b i l i t y i n l e a f shape and  the o b s c u r i n g e f f e c t o f  upon l e a f shape d i c t a t e c e r t a i n procedures f o r e v a l u a t i n g In a l l c a s e s the  falcation  l e a f shape.  l e a v e s s h o u l d be o b s e r v e d under a c o v e r s l i p w i t h t h e i r  adaxial  s i d e down.  The  l e a f shape e x h i b i t e d  must be  a r r i v e d a t through the  subjective  by  a p a r t i c u l a r specimen  assessment o f a t l e a s t twenty-  26 f i v e l e a v e s d i s s e c t e d from h e a l t h y stems. b e a r i n g f a l c a t e l e a v e s may  The  l e a f shape i n p l a n t s  be a s s e s s e d i n s e v e r a l ways.  Frequently,  the l e a v e s borne d o r s a l l y o r v e n t r a l l y on p r o s t r a t e stems a r e o r l e s s f a l c a t e than those l a t e r a l l y p l a c e d .  straight  When o n l y f a l c a t e  leaves  are a v a i l a b l e one must attempt t o p l a c e the f a l c a t e l e a f a d a x i a l s i d e down under a c o v e r s l i p and e x t r a p o l a t e the t h e o r e t i c a l shape from the l e s s asymmetrical s i d e o f the l e a f .  27  F i g v 2 a - g. a.  Terminology  Orbicular  b.  Dblong-Elliptic  Ci.  Broadly  d.  Ovate  e.  Oblong  f.  Oblong-lanceolate  g.  Lanceolate  ovate  o f L e a f Shape  28  9 V  29  F i g . 3 a - v.  Comparison o f l e a f - s h a p e v a r i a t i o n w i t h i n and between  species. a - e.  K.  alpinum  f - i . H.  duriusculum  j - n.  H.  smithii  o - r.  H.  bestii  s - v.  H.  molle  Scale: a  -  ° "  j ,  R  ''  S  -  V J  I  I  1mm  2mm  I  I,  30  31 Pig.  4 a - x.  Comparison o f l e a f - s h a p e  species. a - c.  H.  norvegicum  d -  H.  cochlearifolium  f.  g - j .  H.  styriacum  k - s.  H.  luridum  t - x.  H.  alpestre  Scale: a  c;  d  j,  k  s;  I 0.5mm t - x; 1mm  v a r i a t i o n w i t h i n and between  33 Fig.  5 a - u.  Comparison o f l e a f - s h a p e  v a r i a t i o n w i t h i n and  species. a - d.  H.  polare  e - 1.  H.  ochraceum  in - u .  H. eugyrium f  Scale:  I  1 m m  I  between  34  35  F i g . 6 a - n.  Comparison o f l e a f - s h a p e  v a r i a t i o n w i t h i n and between  species. a - c.  H, subeugyrium from N o r t h America and Europe,  d - f.  H. subeugyrium from Japan,  g - i .  H.  j - n.  H. montanum  Scale:  closteri  37  Fig. 7 a - t.  Comparison o f l e a f - s h a p e  i n n a t u r a l l y growing p l a n t s o f  Hygrohypnum and t h e i r response t o a r t i f i c i a l c u l t u r e . H. alpinum a - b.  N a t u r a l p l a n t s ; c - d.  Cultivated plants  H. d u r i u s c u l u m e - f and i - j . N a t u r a l g - h and k - 1.  Cultivated plants  H. m o l l e m - ri. N a t u r a l o - p.  plants  Cultivated plants  H. b e s t i i  Scale:  q - r.  Natural  s - t.  Cultivated plants  I  1mm  plants  I  plants  38  39  Fig. 8 a - t.  Comparison o f l e a f - s h a p e  i n n a t u r a l l y growing p l a n t s o f  Hygrohypnum and t h e i r response t o a r t i f i c i a l c u l t u r e . H.  H.  H.  H.  smithii a - b.  Natural  plants  o - e.  Cultivated plants  styriacum f - g.  Natural  plants  h - j .  Cultivated plants  luridum k - m.  Natural  plants  n - p.  Cultivated plants  luridum q.  Natural  r.  L e a f d e r i v e d from growth o f v e g e t a t i v e  s - t.  Scale:  1  plants  Leaves d e r i v e d from gametophores  1mm  plants.  grown from  spores.  41 Pig. 9 a - t .  Comparison o f l e a f - s h a p e i n n a t u r a l l y growing p l a n t s  Hygrohypnum and t h e i r r e s p o n s e t o a r t i f i c i a l c u l t u r e . H.  ochraceum a - f.  Natural  plants  g - j . Cultivated plants H. eugyrium k - m.  Natural  plants  n - p.  Cultivated plants  H. montanum q - r.  Natural  plants  s - t.  Cultivated plants  Scale:  a - p,-  I  ,—imm_  q -  1  0.5mm  t;  43  Leaf Apex The shape of the leaf apex, l i k e that of the complete leaf, i s quite variable i n some species and less so i n others.  In some species the shape  of the l e a f apex often overlaps with the shape exhibited by other species. However, toothing and curvature of the margin and concavity modify the character of the l e a f apex so that i f often can be used as an e f f e c t i v e character. The description of the shape of the leaf apex has been subjected to the same terminological ambiguity as has l e a f shape.  For the purposes of  this study F i g . 1 0 i l l u s t r a t e s the a p p l i c a t i o n to Hygrohypnum of the terms describing the shape of the leaf apex. Figure 1 1 b i l l u s t r a t e s the abruptly acuminate leaf apex of Hygrohypnum styriacum.  Both Hygrohypnum luridum.and H. eugyrium have acute  leaf apices  whose margins are often i n r o l l e d imparting a weakly apicula.te  appearance.  The f i n e teeth at the leaf t i p s of H. eugyrium (Fig. 1 1 c & d)  d i f f e r from the h a b i t u a l l y entire leaf t i p s of H. luridum  (Fig. 1 1 a).  Hygrohypnum norvegicum and H. cochlearifolium have long been confused with each other.  The sharply acute l e a f apex of H. norvegicum d i f f e r s from  the rounded, obtuse apex of H. cochlearifolium (Fig. 1 2 a & b). The deep concavity of the leaf apex and i t s tiny squarrose apiculus make Hygrohypnum alpestre unique i n the genus (Big. 1 2 f ) . The but blunt and denticulate leaf apices  acute,  which are regularly observed i n  H* subeugyrium are also unique to t h i s species  (Fig. 1 1 e s f ) ,  Hygrohypnum molle has been confused regularly with H. duriusculum. The leaf apex of the former i s usually gradually tapered and acute with a s l i g h t l y blunted point (Fig. 1 2 c ) . r a r e l y acute, but normally  The apex of-H. duriusculum i s  i t i s rounded obtuse or rounded obtuse and  44 sometimes s l i g h t l y , b u t b l u n t l y a p i c u l a t e  ( F i g . 12  d>.  Hygrohypnum s t y r i a c u m and H. l u r i d u m a r e o f t e n confused w i t h other.  The a b r u p t l y acuminate apex o f the former  from the acute apex o f the l a t t e r  ( F i g . 11 a ) .  ( F i g . 11 b)  each  differs  45  F i g . 10 a - h. a - b.  D e s c r i p t i v e t e r m i n o l o g y f o r the shape o f the l e a f apex,  Rounded  c.  Obtuse  d.  Broadly  e.  Acute  f.  Acuminate  g - h.  acute  Apiculate  46  47  F i g . 11 a - f . a.  H.  luridum  b.  H.  styriacum  c - d.  H. eugyrium  e - f.  H.  Scale:  V  V a r i a t i o n i n t h e shape o f the l e a f  * I  subeugyrium  100um  I  apex.  48  49 F i g . 12 a - f .  V a r i a t i o n i n the shape o f the l e a f apex.  a.  H. norvegicum  b.  H.  c.  H. m o l l e  d.  H. d u r i u s c u l u m  e.  H. montanum  f.  H.  cochlearifolium  alpestre  Scale:  a - e; f  ;  ' I I  100um 0.5mm  I I  50  51Costa The value.  c o s t a o f Hygrohypnum i s v a r i a b l e and thus o f l i m i t e d  taxonomic  The c o s t a i s u s u a l l y s h o r t and double, b u t may a l s o be l o n g and  double o r s h o r t and/or l o n g and s i n g l e o r s i n g l e and f o r k e d . a t some time, e x h i b i t s a s i n g l e c o s t a .  Each s p e c i e s ,  However, the frequency o f o c c u r r e n c e  o f a s i n g l e c o s t a i s so low i n such s p e c i e s as Hygrohypnum eugyrium, H. montanum and H. m o l l e t h a t the c o s t a can be c o n s i d e r e d double.  as o n l y s h o r t and  Hygrohypnum p o l a r e i s unique i n the genus s i n c e i t s c o s t a i s  e x c l u s i v e l y s i n g l e , s t o u t and p e r c u r r e n t o r ending j u s t a few c e l l s below the apex.  A s h o r t , d o u b l e c o s t a predominates i n Hygrohypnum d u r i u s c u l u m ,  but a slender, s i n g l e costa occurs  r e g u l a r l y i n a few l e a v e s .  The c o s t a  o f Hygrohypnum s m i t h i i i s always s t o u t and a l t h o u g h i t i s b o t h s h o r t and double o r s i n g l e , a s i n g l e c o s t a predominates.  Hygrohypnum l u r i d u m i s  e q u a l l y unique f o r w i t h i n the s p e c i e s the c o s t a v a r i e s c o n t i n u o u s l y out  through-  i t s complete range o f v a r i a t i o n as e x h i b i t e d i n the e n t i r e genus.  Leaf  Concavity Leaf c o n c a v i t y i s r e a d i l y apparent i n almost a l l s p e c i e s o f Hygro-  hypnum and i s one o f t h e s e v e r a l f e a t u r e s which u n i f y the genus. it  However,  i s u s e f u l i n d i s t i n g u i s h i n g s p e c i e s o n l y i n cases where i t can be  d e s c r i b e d as d e e p l y  concave.  Deep c o n c a v i t y i s d e f i n e d here n o t i n quan-  t i t a t i v e terms, b u t i n terms o f what can be seen i n a microscope  slide  p r e p a r a t i o n when t h e l e a f has been p l a c e d a d a x i a l s i d e f a c i n g downward. I f t h e c o n c a v i t y i s such t h a t t h e weight o f t h e c o v e r s l i p w i l l impart curved  f o l d that i s nearly p a r a l l e l  a  t o the; l e a f margin, as i n F i g . 13  b s d, o r such t h a t a s e r i e s o f i r r e g u l a r o r l i n e a r f o l d s form throughout the l e a f , as i n F i g . 13 d, then t h e l e a f i s i n t e r p r e t e d as deeply  concave.  F i g u r e s 13 a & c i l l u s t r a t e t h e normal c o n f i g u r a t i o n o f these same l e a v e s . The  asymmetrical c u r v a t u r e o f f a l c a t e l e a v e s imparts  c a n i c u l a t e concavity to the leaves.  a channelled or  The d i f f e r e n c e s shown i n such concav-  i t y among f a l c a t e - l e a v e d s p e c i e s i s n o t s i g n i f i c a n t .  53  F i g . 13 a - d. H.  H.  The n a t u r e o f l e a f  concavity.  alpestre a.  A l e a f as seen w i t h a d i s s e c t i n g  microscope.  b.  The same l e a f as seen i n a microscope s l i d e p r e p a r a t i o n .  cochlearifolium c.  A l e a f as seen w i t h a d i s s e c t i n g  microscope.  d.  The same l e a f as seen i n a microscope s l i d e p r e p a r a t i o n .  55 Median L e a f C e l l s The broad v a r i a t i o n within individual cells of l i t t l e  i n the dimensions o f the median l e a f  cells  s p e c i e s and between d i f f e r e n t s p e c i e s r e n d e r s these  taxonomic  usefulness.  In most c a s e s the extremes o f  median l e a f c e l l l e n g t h v a r y independent o f l e a f s i z e i n a g i v e n s p e c i e s . Only r a r e l y w i l l a comparison be h e l p f u l  i n distinguishing  o f the average c e l l them from each o t h e r .  s i o n s were measured from m i d d l e l a m e l l a t o middle  l e n g t h s o f two s p e c i e s A l l l e a f c e l l dimenlamella.  Marginal Leaf C e l l s The length of the median marginal leaf c e l l s i s extremely useful i n one case.  Hygrohypnum b e s t i i has often been mistaken f o r H. molle or  H. duriusculum.  The median marginal leaf c e l l s of H_. b e s t i i vary from  60 to 250 um long, while those of H. molle or H. duriusculum are rarely more than 55 um.  Equally rarely one or two marginal c e l l s i n H.  and H. subeugyrium are from 65 to 70 um. are very p l a s t i c ,  The marginal c e l l s of H. ochraceum  ranging from 45 to 250 um.  a l l other species are under 50  eugyrium  The marginal l e a f c e l l s of  um.  Figures 14 and 15 i l l u s t r a t e some variation observed i n the marginal leaf c e l l s .  56  Fig.  14 a - g.  V a r i a t i o n i n marginal l e a f  a - b,  H.  alpinum  c - d.  H.  duriusculum  e - f.  Jl. b e s t i i  g - h.  H.  Scale:  L_  molle  50 um  J  cells,  58 F i g . 15 a - f .  V a r i a t i o n i n marginal l e a f  a - b.  H.  ochraceum  c - d.  H.  eugyrium  e - f.  H.  subeugyrium  Scale:  1"'  50^tm  |  cells,  60 Alar  Cells The  a l a r c e l l s a r e one  i n the genus.  o f the most important  taxonomic  criteria  Yet, many o f the problems w i t h i n the genus can be  traced  t o the f a c t t h a t the a l a r c e l l s have not been c a r e f u l l y examined. may  be due,  i n p a r t , to the comments o f Schimper  (1853).  the a l a r c e l l s a r e "wenig oder gar n i c h t a u s g e h o l t deselben may  kaum von der u b r i g e n  He ®aid t h a t  und' das Z e l l h e t z  B l a t t b a s i s verschieden."  His  observation  have r e s u l t e d from the f r e q u e n t o c c l u s i o n o f the a l a r c e l l  by dark c y t o p l a s m i c  This  c o n t e n t s which o b s c u r e any o b s e r v a b l e  lumens  differentiation.  I t i s t r u e t h a t the a l a r c e l l s o f Hygrohypnum a r e not as d r a m a t i c a l l y d i s t i n c t as those o f some s p e c i e s o f C a l l i e r g o n o r Hypnum.  However, i t  i s a l s o t r u e t h a t t h e r e i s s u f f i c i e n t a l a r d i f f e r e n t i a t i o n i n some s p e c i e s o f Hygrohypnum, such t h a t when they a r e c l e a r e d , these c e l l s r e v e a l s e v e r a l r e a d i l y observable Two  patterns of  differentiation.  p r e r e q u i s i t e s a r e a b s o l u t e l y e s s e n t i a l f o r the e f f e c t i v e  ment o f the a l a r c e l l s .  First,  t e n to t w e n t y - f i v e  assess-  c l e a n l e a v e s must be  c a r e f u l l y removed from a mature stem o r p r i m a r y branch segment.  Secondly,  the l e a v e s should be p l a c e d a d a x i a l s i d e downward i n a drop o f c l e a r i n g agent such as Hoyer's, l a c t o - p h e n o l o r Several d i f f e r e n t expressions the genus. shape, 2. 5.  Each e x p r e s s i o n cell  excavation  size, and  6.  3.  KOH.  o f a l a r d i f f e r e n t i a t i o n are evident i n  i s the r e s u l t o f a combination o f 1.  r e l a t i v e numbers o f c e l l s ,  4.  cell  wall thickness,  discoloration.  Quadrate, s h o r t r e c t a n g u l a r o r i r r e g u l a r shapes are those most f r e q u e n t l y encountered among the a l a r c e l l s o f Hygrohypnum.  C e l l s o f such  shapes d i f f e r markedly from the rhomboid, f u s i f o r m o r l i n e a r  flexuose  shapes o f median l e a f c e l l s .  However, the r e c o g n i t i o n o f a d i s c e r n i b l e  group o f a l a r c e l l s depends upon the r e l a t i v e numbers o f quadrate o r s h o r t r e c t a n g u l a r c e l l s and a s u b j e c t i v e assessment o f how a b r u p t l y they d i f f e r from s u r r o u n d i n g c e l l s .  A few i s o l a t e d quadrate  u l a r c e l l s may o c c u r i n Hygrohypnum m o l l e  to s h o r t r e c t a n g -  ( F i g . 16 a ) , b u t the b u l k o f  the c e l l s a r e s i m i l a r t o the l i n e a r o r l o n g r e c t a n g u l a r b a s a l o r lower median l e a f c e l l s .  Such a c o n d i t i o n i s i n t e r p r e t e d here as e x h i b i t i n g  no a l a r d i f f e r e n t i a t i o n .  F i g u r e s 16 b, c & d i l l u s t r a t e  the a l a r r e g i o n s o f H. c o c h l e a r i f o l i u m , H. norvegicum In these s p e c i e s t h e r e a r e s e v e r a l quadrate i n the a l a r r e g i o n . or fusiform c e l l s apparent.  respectively  and H. c l o s t e r i .  to short rectangular c e l l s  However, the t r a n s i t i o n from the s u r r o u n d i n g l i n e a r  i s so g r a d u a l t h a t a d i s t i n c t group o f c e l l s  i s not  T h i s c o n d i t i o n d i f f e r s markedly from t h a t o f H. m o l l e , b u t a l l  o f these t a x a a r e i n t e r p r e t e d as l a c k i n g any a l a r d i f f e r e n t i a t i o n .  In  a d d i t i o n , Hygrohypnum b e s t i i , H. s m i t h i i , H_. s t y r i a c u m and H. montanum a r e s i m i l a r l y i n t e r p r e t e d as e x h i b i t i n g no a l a r  differentiation,  A c l e a r l y d i s c e r n i b l e group o f numerous quadrate tea s h o r t r e c t a n g u l a r a l a r c e l l s o c c u r s i n Hygrohypnum p o l a r e ( F i g . 17 a) and H, l u r i d u m ( F i g , 17 c ) . F i g u r e 18 a i l l u s t r a t e s a s i m i l a r s i t u a t i o n i n H, d u r i u s c u l u m , which d i f f e r s i n the more f r e q u e n t o c c u r r e n c e o f i r r e g u l a r c e l l s and thicker walls.  The a l a r c e l l s o f H. p o l a r e a r e v e r y c o n s t a n t , b u t those o f  H, l u r i d u m and H. d u r i u s c u l u m a r e v a r i a b l e . i n the r e l a t i v e numbers o f a l a r c e l l s ,  i n c l u d i n g the l e a f margins  T h i s c o n t r a s t s w i t h the s i t u a t i o n seen i n F i g . 18 a.  Hygrohypnum d u r i u s c u l u m t h i s v a r i a t i o n o c c u r s i n b o t h r o b u s t and p l a n t s , p a r t i c u l a r l y those from Western North America. bility  little  H. d u r i u s c u l u m may e x h i b i t as few  as 3 v e r t i c a l , p a r a l l e l rows o f a l a r c e l l s , ( F i g . 18 b ) .  While H. l u r i d u m v a r i e s  In  depauperate  I t i s this varia-  t h a t has no doubt c o n t r i b u t e d t o the c o n f u s i o n between H, d u r i u s c u l u m  62 and H. m o l l e and emphasizes t h e need t o observe numerous l e a v e s making a judgement.  The a l a r c e l l s o f H. l u r i d u m and H.  vary f u r t h e r i n c e l l wall thickness, excavation  before  duriusculum  ( P i g . 17 d s 18 c) and  c e l l wall discoloration. Abrupt c e l l enlargement o r i n f l a t i o n i s another f r e q u e n t l y encountered mode o f a l a r d i f f e r e n t i a t i o n .  An abrupt  increase i n c e l l  size i s  i n t e r p r e t e d here t o mean a t l e a s t a two f o l d i n c r e a s e i n l e n g t h and/or width r e l a t i v e t o the adjacent Hygrohypnum eugyrium  cells.  T h i s c o n d i t i o n i s apparent i n  ( F i g . 19 a) and H. alpinum  ( F i g . 19 b ) .  In r a r e i n s t a n c e s a r e c o g n i z a b l e group o f a l a r c e l l s may o c c u r as a gradual and  transition in cell  basal c e l l s .  shape and s i z e from t h e s u r r o u n d i n g  F i g u r e 19 d i l l u s t r a t e s a s m a l l group o f such  w h i l e F i g u r e 19 c i l l u s t r a t e s a l a r g e r group.  median cells,  These two s t a t e s p e p r e s e n t  o p p o s i t e ends o f the spectrum o f a l a r d i f f e r e n t i a t i o n i n Hygrohypnum ochraceum. Another e x p r e s s i o n o f a l a r d i f f e r e n t i a t i o n may n o t be c o n v e n i e n t l y r e f e r r e d t o a geneasal  pattern,  which 1 o r 2 rows o f e n l a r g e d ,  fiigure  18 d i l l u s t r a t e s a s i t u a t i o n i n  i n c r a s s a t e , d i s c o l o r e d c e l l s extend  the l e a f base a t t h e l i n e o f i n s e r t i o n .  across  J u s t above t h i s b a s a l row and  a l o n g t h e l o a f margin i s a v a r i a b l e number o f quadrate o r i r c e g u l a r c e l l s . T h i s a l a r c e l l c o n f i g u r a t i o n i s t y p i c a l o f Hygrohypnum subeugyrium.  63 Fig.  16  a » d.  a.  H.  molle  b.  H.  cochlearifolium  c.  H.  norvegicum  d.  H.  closteri  Scale:  v  A  Variation in alar c e l l differentiation.  1  100um  i  64  65 F i g . 17 a - d.  Variation  a - b.  H.  polare  c - d.  H.  luridum  in alar c e l l  differentiation,  66  67  F i g . 18 a - d. a - c. d.  H.  Scale:  Variation  H.  i n alar c e l l  duriusculum  subeugyrium  I  100  um  differentiation,  68  69  F i g , 19 a - d.  Variation  a.  H. eugyrium  b.  H.  Co-  d.  Scale:  i n alar c e l l J  alpinum H.  ochraceum  1  um  |  differentiation.  70  71 Response t o  Drying  Upon d r y i n g the l e a v e s o f most s p e c i e s e x h i b i t l i t t l e o v e r a l l appearance o r i n t h e i r a t t i t u d e upon the stem. undergo v e r y dramatic  change i n  A few,  however,  changes, which when used w i t h o t h e r c h a r a c t e r s  be u s e f u l i n d i s t i n g u i s h i n g c e r t a i n t a x a .  can  F i g u r e s 20 & 21 compares t h e  m o i s t and d r y c o n d i t i o n s i n some o f these p l a n t s . S h r i n k a g e i n l e a f w i d t h i s the most common response t o d r y i n g r e s u l t s i n several recognizable patterns of l e a f convolution. 20 b i l l u s t r a t e s two i n r o l l can be  such p a t t e r n s .  One  seen i n a l e a f near the middle o f the f i g u r e *  have responded by t w i s t i n g about t h e i r m i d l i n e . i n F i g . 20 d.  o f F i g . 20 d. come s q u a r r o s e .  Figure  p a t t e r n where the l e a f margins  p a t t e r n can be seen i n the upper h a l f o f the same f i g u r e .  i s observable  and  The  second  Here the  leaves  A similar configuration  A t h i r d p a t t e r n can be seen i n the upper h a l f  In t h i s case the margins i n the lower h a l f o f the l e a f These d r y i n g responses c h a r a c t e r i z e Hygrohypnum  be-  duriusculum.  F i g u r e s 21 b & 21 e i l l u s t r a t e the d r y i n g response o f Hygrohypnum bestii.  Here severe  l e a f c o n t o r t i o n does not accompany s h r i n k a g e .  the l e a v e s remain n e a r l y p l a n e o r a r e s l i g h t l y t w i s t e d .  Instead,  In such cases  the  r e d u c t i o n i n apparent l e a f s u r f a c e a r e a i s more e v i d e n t . Hygrohypnum m o l l e has bestii. drying  l o n g been confused  w i t h H. d u r i u s c u l u m  However, H. m o l l e undergoes v e r y l i t t l e ( F i g , 21 c ) ,  A s l i g h t amount o f s h r i n k a g e  and  H.  change i n appearance upon and  t w i s t i n g i s evident,  but the l e a v e s tend to r e t a i n t h e i r i m b r i c a t e t o s l i g h t l y s p r e a d i n g a t t i t u d e . F o r a l l i t s remarkable polymorphism, Hygrohypnum l u r i d u m  shows  little  change i n appearance upon d r y i n g . An e n t i r e l y d i f f e r e n t response to d r y i n g i s seen i n F i g . 62 e. the l e a v e s a r e f a l c a t e and  Here  folded along t h e i r midline i n t o a c a n i c u l a t e  channel.  Shrinkage i n l e a f w i d t h i s n e g l i g i b l e , b u t t h e l e a f apex  undergoes  a c h a r a c t e r i s t i c t w i s t i n g t o resemble a f i l i f o r m l e a f t i p .  T h i s i s a common response i n Hygrohypnum ochraceum. The l e a v e s o f Hygrohypnum c l o s t e r i a r e w i d e l y spaced a l o n g t h e i r stems.  In t h e m o i s t c o n d i t i o n t h e l e a v e s a r e s p r e a d i n g ( F i g . 76 d ) .  Upon d r y i n g the l e a v e s s h r i n k and t w i s t and a c c e n t u a t e the d i s t a n t s p a c i n g by becoming almost s q u a r r o s e ( F i g . 76 c ) .  73 F i g . 20 a - h.  V a r i a t i o n i n a l t e r a t i o n i n h a b i t from wet to dry  conditions i n morphologically d i f f e r e n t shoots, H.  duriusculum a.  Moist;  b.  Dry  c.  Moist?  d.  Dry  e.  Moist;  f.  Dry  g.  Moist;  h.  Dry  74  75 Fig.  21 a - e.  V a r i a t i o n i n a l t e r a t i o n i n h a b i t from wet  donditions i n morphologically d i f f e r e n t H,  H.  shoots.  bestii a.  Moist;  b.  Dry  d.  Moist;  e.  Dry  molle c.  M o i s t ; unchanged i n the d r y c o n d i t i o n .  to dry  77  Sexuality The problems imposed on bryophyte taxonomy by t h e i n f r e q u e n c y o f s e x u a l p l a n t s i s w e l l known.  However, i n Hygrohypnum, t h e s e x u a l con-  d i t i o n p r o v i d e s c l e a r d i s c o n t i n u i t i e s between c e r t a i n s p e c i e s o r c o r r e s l a t e s w i t h d i s c o n t i n u i t i e s e v i d e n t from o t h e r c h a r a c t e r s . A l l b u t f o u r s p e c i e s o f Hygrohypnum a r e a u t o i c o u s . bestii,  H. ochraceum and H. p o l a r e a r e d i o i c o u s .  Hygrohypnum  The male and female  p l a n t s o f H. b e s t i i and H, p o l a r e a r e s i m i l a r i n appearance.  The male  p l a n t s o f H, ochraceum a r e more s l e n d e r and t h e i r leaves.more w i d e l y spaced than i n t h e females. Hygrohypnum s t y r i a c u m e x h i b i t s t h e most unusual i n t h e genus.  T h i s s p e c i e s has a s i n g l e perigonium  sexual c o n d i t i o n subtended by 2 t o  3 p e r i c h a e t i a which i n t u r n a r e subtended by 1 t o 3 s m a l l , s c a l e l i k e bracts  ( F i g . 13 a & b ) .  T h i s complete c l u s t e r o f s e x u a l i n f l o r e s c e n c e s  i s jfeorne i n t h e a x i l o f a f o l i a g e l e a f . as f o u r l e a v e s o r as few as one.  The perigonium  When o n l y one p e r i g o n i a l l e a f i s p r e s e n t  the a n t h e r i d i a d i r e c t l y abut t h e o u t e r p e r i c h a e t i a l l e a f adjacent.  may have as many  immediately  I n those cases where t h e a n t h e r i d i a a r e e n c l o s e d by 2 t o 4  p e r i g o n i a l leaves the o v e r a l l d i s p o s i t i o n o f the sexual i n f l o r e s c e n c e s d i f f e r s from t y p i c a l a u t o i e c y o n l y i n immediate p r o x i m i t y t o one another. When o n l y 1 p e r i g o n i a l l e a f i s p r e s e n t , however, t h e s i t u a t i o n i s reminescent o f paroicy. situation.  The term p s e u d o - p a r o i c y  i soffered to describe  this  78 Fig.  22 a - b.  The pseudo-paroicous i n f l o r e s c e n c e o f Hygrohjypnum  styriacum. a.  The i n f l o r e s c e n c e as i t appears n a t u r a l l y .  b.  A diagrammatic  representation  o f the i n f l o r e s c e n c e .  79  80 Perichaetial The  Leaves  p e r i c h a e t i a l l e a v e s e x h i b i t a number o f u s e f u l  s t r e n g t h e n the d i s c o n t i n u i t i e s between c e r t a i n Hygrohypnum alpinum and  by  the  inner p e r i c h a e t i a l leaves the o v e r r i d i n g  abaxial  o f the d i s t a l end  cochleaeifolium.  inner p e r i c h a e t i a l  leaf  genus.  weakly p l i c a t e . strong p l i c a e .  (See  s u r f a c e o f some a p i c a l c e l l s  walls.  These p a p i l l a e are  Occasionally  same p o s i t i o n o f the  one inner  or  formed  two  perichaetial  H.  montanum s i m i l a r l y stand a p a r t from  i n n e r p e r i c h a e t i a l l e a v e s are  the  e i t h e r plane or  F i g . 25  f, g &  h)  Hygrohypnum c o c h l e a r i f o l i u m  t h i s i n the The °f  exhibition of strongly  costa i s variable  Hygrohypnum.  very  p e r i c h a e t i a l l e a v e s o f a l l o t h e r s p e c i e s have 2 to 4  A p l a n e i n n e r p e r i c h a e t i a l l e a f margin i s the g e n e r a l c o n d i t i o n genus.  of  cells.  Their  The  by  A l l o t h e r s p e c i e s o f Hygrohypnum have smooth  Hygrohypnum c l o s t e r i and b u l k o f the  taxa.  ( F i g , 23 a & b ) .  p a p i l l a e have been observed i n the l e a v e s o f H.  that  H. m o l l e stand a p a r t from o t h e r s p e c i e s  the p a p i l l a e they e x h i b i t on the  characters  i n the  However, the  and  H.  in  smithii d i f f e r dramatically  r e c u r v e d margins  ( F i g . 24 d,  the from  e & f).  i n n e r p e r i c h a e t a i l l e a v e s o f most s p e c i e s  i n n e r p e r i c h a e t i a l l e a v e s o f H.  polare,  like  the v e g e t a t i v e l e a v e s , e x h i b i t a s t r o n g , s i n g l e p e r c u r r e n t c o s t a  ( F i g . 25  The  but  v e g e t a t i v e l e a v e s o f H.  s h o r t and f o r the  double c o s t a e do o c c u r .  costa i s exclusively  The  shape o f the  However, i t may o f the  closfeeri are  s i n g l y costate,  I t s inner p e r i c h a e t i a l leaves are  single  apex o f the  usually  ( F i g . 25  be o f some v a l u e i n c e r t a i n c a s e s .  a b r u p t i n Hygrohypnum alpinum and  H.  few  unusual  h).  inner peeichaetial  i n n e r p e r i c h a e t i a l l e a f apex i s a c u t e .  a  c),  The  l e a f apex i s v a r i a b l e . The  most common shape  acuteness v a r i e s  cochlearifolium  ( F i g . 24  a & f)  from to  81 g r a d u a l l y so i n most s p e c i e s . rounded o r o b t u s e  Hygrohypnum  ( F i g . 23 c & 24 d ) .  s m i t h i i d i f f e r s by b e i n g  R e f l e c t i n g the shape o f the apex  o f i t s v e g e t a t i v e l e a v e s , H. s t y r i a c u m has a s l i g h t l y aeuminate -innerp e r i c h a e t i a l l e a f apex ( F i g . 25 d ) . H  *  The i n n e r p e r i c h a e t i a l l e a v e s o f  c l o s t e r i a r e a l s o s l i g h t l y accuminate ( F i g . 25 h ) , a s t r i k i n g de-  p a r t u r e from t h e a c u t e , b u t b l u n t apex i n the v e g e t a t i v e  leaves.  The t o o t h i n g a l o n g the margin and i n the apex o f the i n n e r c h a e t i a l l e a v e s tends t o r e f l e c t the s i t u a t i o n i n the v e g e t a t i v e but i s g e n e r a l l y t o o v a r i a b l e to be u s e f u l .  perileaves,  82 F i g . 23 a - c.  Variation  a.  H.  molle  b.  H.  alpinum  c.  H.  smithii  Scale:  L  100um  i n the p e r i c h a e t i a l  \  l e a f apex.  84  F i g . 24 a - f .  V a r i a t i o n i n the p e r i c h a e t i a l  a.  H.  alpinum  b.  H.  duriusculum  c.  H.  molle  d.  H. s m i t h i i  e - f.  H.  cochlearifolium  leaves.  86 F i g . 25 a - g. a - b. c.  H.  H.  Variation luridum  polare  d - e.  H.  f - g.  H. montanum  h.  H.  styriacum  closteri  i n the p e r i c h a e t i a l  leaves,  Sporophyte The sporophyte o f f e r s few c h a r a c t e r s t h a t a r e o f v a l u e i n d i s t i n g u i s h i n g s p e c i e s o f Hygrohypnum.  The sporophytes a r e not u n i f o r m , but  the d i f f e r e n c e s they e x h i b i t a r e i n such v a r i a b l e f e a t u r e s as c a p s u l e and s e t a l e n g t h o r c o l o r . are  However, the annulus and the endostomial  cilia  useful i n c e r t a i n cases. All  Hygrohypnum s p e c i e s e x c e p t H. l u r i d u m have a w e l l developed  annulus.  The absence o f the annulus i n H. l u r i d u m p r o v i d e s a sharp c o n t r a s t w i t h H  -  s t y r i a c u m , a s p e c i e s w i t h which H. l u r i d u m i s o f t e n c o n f u s e d . Hygrohypnum d u r i u s c u l u m has 1 t o 3 w e l l developed c i l i a  between  a d j a c e n t e n d o s t o m i a l segments, w h i l e i n H. alpinum and H. m o l l e the are  rudimentary o r wanting.  cilia  Hygrohypnum c o c h l e a r i f o l i u m a l s o d i f f e r s  from  H. m o l l e by i t s 2 t o 3 w e l l d e v e l o p e d c i l i a , which u n f o r t u n a t e l y f a l l away very  early.  89 TAXONOMIC TREATMENT Hygrohypnum L i n d b . , A c t . Soc. Sc. Fenn. 10:277.  1872  Hypnum Hedw. s e c t . A l p e s t r i a Hub., Muse. Ger, 629. 1833. Hypnum Hedw. S e c t . P a l u s t r i a De Limnobium  Schimp.,  B r y o l . E u r . 6:65.  Hypnum Hedw. subg. Limnobium Bot.  Not., S y l , Muse. 45. 1838. 1853, hom. i l l e g .  (B.S. & G.) S u l l , i n Gray, Man.  N. U. S. ed. 2:677.  1856.  Stereodon B r i d . s e c t . Limnobium  (B.S. & G.) M i t t . , J . L i n n .  Soc. B o t . 8:42, 1864. Amblystegium B.S, & G, subg. Hygrohypnum  (Lindb.) L i n d b . ,  Musci Scand. 33. 1879. Hypnum Hedw. s e c t . Limnobium  (B.S. & G,) Ren., Rev. B r y o l .  1 0 ( 3 ) : 41. 1883. Hypnum Hedw. subg. Hypnum B o u l . s e c t . Limnobium  (B.S, & G.)  B o u l . , Muse. F r a n c e . 1884. Hypnum Hedw. s e c t . Eu-Limnobia Du Bat, B u l l . T r i m . Soc. B o t . Lyon 3:54. 1885. Hypnum Hedw. s e c t . Hetero-Limnobia Du Bat, Bot. Calliergon  B u l l . T r i m . Soc.  Lyon 3:55. 1885. ( S u l l . ) Kindb. i n p a r t , Canad. Rec. S c i . 6(2):72.  1894. Amblystegium B.S. & G. s e c t . Hygrohypnum B r i t . Moss Calliergon  (Lindb.) Braithw.,  F I . 3:17. 1898.  ( S u l l . ) Kindb. sugb. Limnobion Kindb., E u r . N. Am.  B r y i n . 1:79. 1897. Calliergon  ( S u l l . ) Kindb. subg, Pseudo-Limnobion Kindb., E u r .  N. Am. B r y i n . 1:80, 1897.  90  Calliergon  ( S u l l . ) Kindb. subg. Pseudo-Limnobion  Kindb. s e c t .  B a d i a f o r m i a Kindb. i n p a r t . , E u r . N. Am. B r y i n . 1:80. 1897. Calliergon  ( S u l l . ) Kindb. subg. Pseudo-Limnobion  Kindb. s e c t .  O c h r a c e i f o r m i a Kindb., E u r . N. Am. B r y i n . 1:80. 1897. Calliergon  ( S u l l . ) Kindb. subg. Pseudo-Limnobion  Kindb. s e c t .  M o l l i f o r m i a Kindb., E u r , N. Am, B r y i n . 1:80, 1897. Calliergon  ( S u l l . ) Kindb-. subg. Pseudo-Limnobion  Kindb. s e c t .  M o n t a n i f o r m i a Kindb., E u r . N. Am. B r y i n . 1:80, 1897. Names t r e a t e d as synonyms elsewhere, b u t f o r which t h e l i t e r a t u r e was u n a v a i l a b l e  f o r assessment d u r i n g t h i s study.  Hypnum Hedw. s u b s e c t . P a l u s t r & a (fie NotJ Rabenh,, Deutsch, K r y p t . P I . 2(3);270.  1848.  Hypnum Hedw. subg. Hygrohypnum  (Lindb.) S c h i f f n . , Lotos Prag,  55:211. 1907. Hypnum Hedw. s e c t . Hygrohypnum  (Lindb.) P a r . , I n d . B r y o l . 589.  1896,  A G e n e r i c D e s c r i p t i o n o f Hygrohypnum P l a n t s u s u a l l y growing on i r r i g a t e d r o c k and stones i n and b e s i d e s m a l l , c o l d , s w i f t l y r u n n i n g mountain l a r g e r i v e r s , r a r e l y a t low e l e v a t i o n s  streams o r r i v u l e t s , sometimes i n i n temperate zones, r a r e l y on wood,  o r submerged i n s t i l l water o r on damp, shaded c l i f f ledges? p l a n t s  var-  i o u s l y forming l o o s e l y t o v e r y t i g h t l y woven, appressed mats o r a s c e n d i n g patches, mats o r r a r e l y t u r f s ; c o l o r v a r i a b l e , d i r t y - y e l l o w , y e l l o w - g r e e n , b r i g h t o r dark, d u l l - g r e e n ,  yellow-brown,  olivaceous-green, blackish-green,  most s p e c i e s e x h i b i t i n g a r u s t y m o t t l i n g o r v a r i e g a t i o n , r a r e l y one ox two s p e c i e s e x h i b i t a b r i g h t m e t a l l i c r e d o r bronze p i g m e n t a t i o n , a l l s p e c i e s becoming yellow-brown, brown, reddish-brown o r b l a c k i s h - b r o w n w i t h age; stems  u s u a l l y 1 to 15 cm tips,  long,  r a r e l y t o 20 cm,  procumbent o r a s c e n d i n g a t  l e a f y throughout o r becoming denuded i n the o l d e s t  extremities;  b r a n c h i n g i r r e g u l a r , a r i s i n g from the a x i l s o f l a t e r a l o r v e n t r a l procumbent, a s c e n d i n g , r a r e l y e r e c t o r f a s t i g a t e ; usually revealing  2 to 4,  Stem  sometimes 5 rows o f s m a l l ,  the  leaves,  cross-sections  thick-walled,  yellow-  ish-brown, brown o r reddish-brown c o r t i c a l c e l l s , r a r e l y e x h i b i t i n g an i n c o m p l e t e l y d i f f e r e n t i a t e d o r w e l l ^.developed row'' o f s l i g h t l y l e s s t h i c k i n f l a t e d , thin-walled hyaline  and  outer c o r t i c a l c e l l s , medullary c e l l s l a r g e r ,  t h i n n e r w a l l e d , o f t e n becoming d i s c o l o r e d and  w i t h age? c e n t r a l s t r a n d p o o r l y  or  usually  thicker walled  t o w e l l developed, r a r e l y e n t i r e l y absent;  r h i z o i d s reddish-brown, a r i s i n g from the base o f v e n t r a l stem l e a v e s o r v e n t r a l s i d e o f the  the  perichaetia.  Leaves v a r i a b l e , u s u a l l y o r b i c u l a r , b r o a d l y e l l i p t i c , or ovate, ovate-lanceolate, usually straight, often  b r o a d l y ovate  oblong or oblong-lanceolate, r a r e l y  lanceolate;  f a l c a t e and/or secund, sometimes s q u a r r o s e r e c u r v e d ;  apex b r o a d l y rounded o r obtuse to a c u t e , r a r e l y s l i g h t l y acuminate o r b r o a d l y rounded and/or obtuse w i t h an a p i c u l u s , or f i n e l y denticulate, apex o r r a r e l y c o a r s e l y  a p i c a l margins u s u a l l y  sometimes s e r r u l a t e , r a r e l y a few  f i n e teeth at  entire the  d e n t i c u l a t e , a p i c a l margins u s u a l l y p l a n e , sometimes  i n r o l l e d ao as to appear a p i c u l a t e , r a r e l y r e c u r v e d o r r e f l e x e d as an margins o f the lamina u s u a l l y e n t i r e o r f i n e l y and  apiculus;  i r r e g u l a r l y denticulate,  :<  r a r e l y s e r r u l a t e throughout, u s u a l l y p l a n e , sometimes i n f o l d e d as wing a l o n g one  o r b o t h s i d e s o f the l e a f , r a r e l y n a r r o w l y r e c u r v e d ; c o n c a v i t y  u s u a l l y s h a l l o w t o d e e p l y so, r a r e l y almost p l a n e ; c o s t a v a r i a b l e , s h o r t and and  double w i t h one  double w i t h one  or stout,  variable, usually  o r b o t h arms ending below m i d l e a f , sometimes  o r b o t h arms e x t e n d i n g beyond m i d l e a f , o r s i n g l e ,  u s u a l l y ending between 1/2  t o 3/4  o f the l e a f l e n g t h ,  rarely  long slender pre-  92  c u r r e n t , sometimes f o r k e d one o r t w i c e ,  n e a r l y a l l the c o s t a l v a r i a t i o n  may be o b s e r v e d w i t h i n a s i n g l e p l a n t ; l e a v e s  a b r u p t l y narrowed a t t h e  i n s e r t i o n , sometimes c l a s p i n g t h e stem when t h e a l a r c e l l s a r e excavated; leaves  crowded t o d i s t a n t ; a t t i t u d e upon t h e stem when wet v a r i a b l e ,  u s u a l l y a p p r e s s e d i m b r i c a t e o r tumid j u l a c e o u s , wide s p r e a d i n g ,  upon d r y i n g the l e a v e s may change l i t t l e  undergo v a r y i n g degrees o f l a t e r a l Aceolation variable, a l l c e l l s bic  or spreading  o r f u s i f o r m t o long,  shrinkage  or rarely  i n attitude or  and c o n t o r t i o n .  smooth, median l e a f c e l l s  s h o r t rhom-  linear-flexuose, usually l i n e a r flexuose,  u s u a l l y o r u n i f o r m w i d t h ; sometimes u n u s u a l l y  cells  wide c e l l s a r e i r r e g u l a r l y  interspersed, walls u s u a l l y t h i n , r a r e l y thickened;  toward t h e l e a f apex  the c e l l s g e n e r a l l y become s h o r t e r , f u s i f o r m rhombic o r o v a l , sometimes changing l i t t l e  from t h e median c e l l s ; m a r g i n a l l e a f c e l l s u s u a l l y l e s s than  60 urn l o n g , r a r e l y e x c e e d i n g 60 urn, b u t when d o i n g ceeding  so o f t e n r e g u l a r l y ex-  100 urn; b a s a l c e l l s v a r i a b l e , becoming g r a d u a l l y wider, l o n g e r o r  s h o r t e r than the median c e l l s o r changing l i t t l e , o r changing l i t t l e ,  becoming t h i c k e r w a l l e d  p i t t e d o r u n p i t t e d , h y a l i n e o r v a r i o u s l y pigmented p a r -  t i c u l a r l y when t h i c k - w a l l e d , yellow-brown, r e d d i s h - y e l l o w ,  brown o r r e d d i s h -  brown; a l a r c e l l s v a r i a b l e , u n d i f f e r e n t i a t e d o r v a r i o u s l y f o r m i n g s m a l l and i r r e g u l a r t o l a r g e groups o f q u a d r a t e , s h o r t r e c t a n g u l a r o r i r r e g u l a r o r v a r i a b l e groups o r e n l a r g e d  and t h i n - w a l l e d  to i n f l a t e d c e l l s ,  cells  plane  o r excavated, h y a l i n e o r yellow^brown, brown, reddish-brown, r a r e l y b r i g h t red. Plants autoicous, ovate, laminal c e l l s  d i o i c o u s o r pseudo-paroicous; p e r i g o n i a l leaves  smooth, e c o s t a t e , margins e n t i r e o r weakly s e r r u l a t e ,  c o n c a v e - i m b r i c a t e e n c l o s i n g 4 t o 10 o b l o n g - ^ e l l i p t i c antthfe'tii&dia and s e v e r a l hyaline, u n i s e r i a t e periphyses;  outer  and m i d d l e p e r i c h a e t i a l l e a v e s v a r -  93 i a b l e , ovate to o v a t e - l a n c e o l a t e , s i n g l e to midleaf,  c o s t a absent, s h o r t and  squarrose-reflexed  i n the upper h a l f , l a m i n a l  smooth; i n n e r p e r i c h a e t i a l l e a v e s e r e c t , b r o a d l y , angular  double o r  l o n g l i n e a r or  cells tri-  l a n c e o l a t e , u s u a l l y 2 to 4 deep p l i c a e , r a r e l y p l a n e o r n e a r l y  so, c o s t a v a r i a b l e , absent, s h o r t and and/or f o r k e d ,  double, l o n g and  f a i n t o r s t o u t , margins u s u a l l y p l a n e ,  double or s i n g l e rarely  recurved,  e n t i r e o r f i n e l y to c o a r s e l y s e r r u l a t e , e s p e c i a l l y i n the apex, apex obtusely few  acute to l o n g t a p e r i n g acute,  c e l l s i n the  l a m i n a l c e l l s smooth, r a r e l y a  l e a f apex are p a p i l l o s e on  means o f d i s t a l l y o v e r l a p p i n g S e t a 6 t o 31 mm  the a b a x i a l l e a f s u r f a c e  by  endwalls.  long,, u s u a l l y 10 to 20 mm,  e r e c t or s l i g h t l y  when m o i s t , smooth, c o l o r v a r i a b l e , y e l l o w i s h - o r a n g e ,  inclined  yellowish-red,  reddish-brown, dark b r i g h t r e d ; c r o s s s e c t i o n s r e v e a l i n g 2 to s e v e r a l rows of small, thick-walled c e l l s , medullary c e l l s s l i g h t l y l a r g e r , thin-walled, becoming t h i c k e r w a l l e d w i t h  age;  c e n t r a l s t r a n d broad, w e l l developed;  upon d r y i n g the s e t a t w i s t s i n v a r i o u s d i r e c t i o n s . Capsules 1 to 3 mm c a l o r i n c l i n e d and  l o n g , o v o i d t o o b l o n g - c y l i n d r i c a l , e r e c t and  s l i g h t l y to s t r o n g l y a r c u a t e ;  symmetri-  color v a r i a b l e , yellow-  brown, brown, o r reddish-brown; e x o t h e c i a l c e l l s v a r i a b l e , rounded, quadrate, short rectangular, i r r e g u l a r , present  o r absent,,  r a r e l y absent;  c u c u l l a t e ; capsule  d i f f e r e n t i a t e d neck  stomates s u p e r f i c i a l on the neck and  v a r i a b l e i n number, 7 to 30; times 4,  t h i c k or t h i n walled;  lower  capsule,  annulus u s u a l l y o f 2 to 3 rows o f c e l l s , some-  operculum c o n i c o r c o n i c a p i c u l a t e ;  calyptra  -  response t o d r y i n g v a r i a b l e , g e n e r a l l y s h r i n k i n g , remain-  i n g e r e c t o r becoming s t r o n g l y a r c u a t e  c y l i n d r i c , the neck when d i f f e r e n t i a t e d  becoming w r i n k l e d  s t r o n g l y c o n t r a c t e d beneath the mouth  upon d r y i n g , c a p s u l e  or sometimes h a r d l y a t a l l . Peristome double, exostome o f 16 t e e t h , p a l e y e l l o w ,  yellow-brown, o r  94 l i g h t reddish-brown  below, h y a l i n e i n the upper h a l f , each segment i s  h o r i z o n t a l l y s t r i a t e i n the lower 1/2 upper  1/3  t o 1/2;  t o 1/3,  f i n e l y p a p i l l o s e i n the  endostome o f 16 h y a l i n e t o p a l e y e l l o w t e e t h ,  p a p i l l o s e t o almost smooth, f r e q u e n t l y c r a c k e d a l o n g the k e e l ,  finely especially  i n the lower h a l f ; 1 t o 3 r a d i m e n t a r y t o w e l l developed c i l i a between a d j a c e n t segments, b a s a l membrane 2 t o 4 t i e r s i n h e i g h t . Spores u s u a l l y dusky y e l l o w o r y e l l o w - g r e e n , f i n e l y  papillose.  95 Key  t o t h e s p e c i e s o f Hygrohypnum  1.  Stem c r o s s - s e c t i o n r e v e a l i n g an e p i d e r m i s o f s m a l l t h i c k - w a l l e d c e l l s which a r e s i m i l a r t o t h e s u b a d j a c e n t c o r t i c a l c e l l s . .  4  Stem c r o s s - s e c t i o n r e v e a l i n g an e p i d e r m i s i n which some o r a l l o f the c e l l s a r e e n l a r g e d and t h i n - w a l l e d r e l a t i v e t o t h e suba d j a c e n t c o r t i c a l c e l l s o r some c e l l s have a s l i g h t l y t h i n n e r and l e s s pigmented o u t e r t a n g e n t i a l c e l l w a l l compared w i t h t h e o t h e r epidermal c e l l s w a l l s and t h e s u b a d j a c e n t c o r t i c a l c e l l s  2  1.  2.  Costa uniformly  s t r o n g and s i n g l e , p e r c u r r e n t  H. p o l a r e  2.  C o s t a s h o r t and double, double t o m i d l e a f o r i f s i n g l e and/or f o r k i n g , then ending w e l l b e f o r e t h e apex  3  3.  A l a r c e l l s i n f l a t e d , forming a d i s t i n c t group o f c e l l s , o f t e n becoming r e d o r reddish-brown w i t h age; l e a f apex acute...H. eugyrium  3.  A l a r c e l l s w i d e r and s h o r t e r than a d j a c e n t c e l l s , b u t n e i t h e r i n f l a t e d n o r abruptly d i f f e r e n t from them, m a r g i n a l a l a r c e l l s i n c r e a s i n g i n l e n g t h d i s t a l l y from t h e p o i n t o f i n s e r t i o n , h y a l i n e o r y e l l o w i s h ? never r e d ; l e a f apex b l u n t H. ochraceum  4.  Leaves a l l s t r a i g h t ; sometimes secund.  4.  Leaves f a l c a t e o r some l e a v e s on the same o r d i f f e r e n t stems w i t h i n t h e same specimen f a l c a t e and/or s t r a i g h t .....18  5.  Leaves b r o a d l y o v a t e t o o r b i c u l a r  5.  Leaves o v a t e t o o b l o n g - o v a t e o r o v a t e - l a n c e o l a t e  6.  Median m a r g i n a l l e a f c e l l s 60 um o r l o n g e r  6.  Median m a r g i n a l l e a f c e l l s r a r e l y l o n g e r than 55 um  7.  Alar c e l l s c l e a r l y d i f f e r e n t i a t e d , either thin-walled or incrassate  8  A l a r c e l l s u n d i f f e r e n t i a t e d o r a few q u a d r a t e o r s h o r t c e l l s which a r e i n c r a s s a t e o r t h i n - w a l l e d  9  7.  5  6 ...11 H. b e s t i i .....7  rectangular  8.  A l a r c e l l s t h i n - w a l l e d , u s u a l l y h y a l i n e , e n l a r g e d , rounded r e c t a n g u l a r , forming a r e c t a n g u l a r group whose l o n g a x i s p a r a l l e l s the l e a f margins; i n e e r p e r i c h a e t i a l l e a v e s p a p i l l o s e on t h e .\baxial s u r f a c e near t h e apex. „H. alpinum  8.  A l a r c e l l s i n c r a s s a t e , c l e a r l y pigmented i n o l d e r l e a v e s , quadrate, s h o r t r e c t a n g u l a r o r i r r e g u l a r , f o r m i n g an i r r e g u l a r group; i n n e r p e r i c h a e t i a l l e a v e s smooth , H. d u r i u s c u l u m  96  9.  Costa usually single, stout, to midleaf or s l i g h t l y beyond, sometimes forked and/or stout, short and double; plants very coarse and r i g i d H. smithii  9.  Costa almost exclusively short and double, i f single, then the costa i s slender and the plants are soft  10  10. Leaves deeply concave to cochleariform, usually 0.8 to 1.2 mm long, apex obtuse or broadly rounded, inner p e r i c h a e t i a l leaf margins e n t i r e and recurved; endostomal c i l i a 2 to 3. H. cochlearifolium 10. Leaves concave, but never cochleariform, usually 1.0 to 1.7 mm long, apex tapering to an acute, but blunt point; inner p e r i chaetial leaf margins coarsely denticulate and plane; endostomal c i l i a rudimentary or wanting H. molle 11. Alar c e l l s c l e a r l y d i f f e r e n t i a t e d , either i n f l a t e d and mostly t h i n walled or smaller, incrassate and quadrate to short rectangular.... 12 11. Alar c e l l s undifferentiated or with but a few quadrate to short r e c tangular c e l l s which do not form a recognizable group 14 12.  Leaf apex narrowly recurved along the margin and reflexed as a small apiculus, e s p e c i a l l y i n leaves at or near stem or branch t i p s ; leaves deeply concave, e s p e c i a l l y near the apex H. alpestre  12. Margins of leaf apices plane or variously i n r o l l e d , never recurved./ apex never forming a reflexed apiculus.... 13 13. Leaf apices  always acute and entire  13. Someileaf apices denticulate  H.  luridum  with a few f i n e teeth or obtuse and d i s t i n c t l y , .H. subeugyrium  14. Leaf, apex abruptly acuminate, tapering to a slender t i p ; plants pseudo-paroicous. H. styriacum 14.  Leaf apex obtuse or acute, with or without a blunt t i p  15  15. Costa predominantly single to midleaf or beyond, sometimes short and double....  16  15. Costa usually short and double, r a r e l y single to midleaf  17  16. Leaf apex obtuse; plants coarse; inner p e r i c h a e t i a l leaves p l i c a t e , margins recurved H. s m i t h i i 16. Leaf apex acute with a blunt t i p ; plants coarse; inner p e r i c h a e t i a l leaves never p l i c a t e , margins plane H. c l o s t e r i 17. Leaf apex entire; leaves usually 0.5  to 0.8 mm  long...H, norvegicum  97 17. Margin o f t h e l e a f apex uneven t o d e n t i c u l a t e ; l e a v e s u s u a l l y 1.0 t o 1.7 iron l o n g H_. m o l l e 18.  Leaves e x c l u s i v e l y f a l c a t e  19  18. F a l c a t e and s t r a i g h t l e a v e s o c c u r r i n g s i m u l t a n e o u s l y on t h e same stem o r on d i f f e r e n t stems w i t h i n t h e same specimen*,  23  19.  20  A l a r c e l l s u n d i f f e r e n t i a t e d from a d j a c e n t  cells  19. A l a r c e l l s w e l l developed; s m a l l , numerous, quadrate t o s h o r t - r e c t a n g u l a r , i n c r a s s a t e o r i n f l a t e d ; p l a n e o r excavated 21 20.  Leaf margin f i n e l y t o c o a r s e l y s e r r u l a t e , e s p e c i a l l y i n t h e apex, and n a r r o w l y r e c u r v e d , p a r t i c u l a r l y i n the lower h a l f..of the l e a f  20.  H. montanum  L e a f margin always e n t i r e and p l a n e  21. A l a r c e l l s i n f l a t e d  H. eugyrium  21. A l a r c e l l s s m a l l , quadrate t o s h o r t r e c t a n g u l a r , 22. Leaf apex always acute, 22.  H_. s t y r i a c u m  t h e margin e n t i r e  incrassate  22  H. l u r i d u m  Some l e a f apices.; obtuse and d i s t i n c t l y d e n t i c u l a t e o r acute w i t h a few f i n e t e e t h  H_. subeugyrium  23. L e a f apex always e n t i r e 23. L e a f a p i c e s - b e a r i n g a few f i n e t e e t h  24 ...25  24.  L e a f apex acute; a l a r c e l l s numerous, quadrate t o s h o r t r e c t a n g u l a r ; annulus absent; p l a n t s a u t o i c o u s , H. l u r i d u m 24. L e a f apex a b r u p t l y acuminate; a l a r c e l l s u n d i f f e r e n t i a t e d o r j u s t a few quadrate c e l l s ; annulus p r e s e n t ; p l a n t s pseudo-paroicous H. s t y r i a c u m 25. A l a r c e l l s i n f l a t e d , t h i n - w a l l e d , l e a f apex always acute; stem e p i d e r m i s d i f f e r e n t i a t e d as an i l l - d e f i n e d hyalodermis..H. eugyrium 25.  A l a r c e l l s quadrate t o s l i g h t l y e n l a r g e d , i n c r a s s a t e ; some l e a f a p i c i e s o b t u s e and d e n t i c u l a t e ; stem e p i d e r m i s s i m i l a r t o c o r t i c a l stem c e l l s , , H, subeugyrium  98 Hygrohypnum alpinum Neotype:  (Lindb.) Loesk., Hedwigia 43:194.  Norway, Gudbrandsdalenj  B l y t t as Hypnum m o l l e .  Hypnum alpinum Schimp., Syn. ed. 2:777. Amblystegium  1904.  1876.  (S-PA).  hom.  m o l l e v a r . alpinum L i n d b . , Musci Scand,  Hypnum d i l a t a t u m v a r . alpinum Hypnum m o l l e v a r . alpinum  (Lindb.) Ren.,  Rev.  illeg. 33.  1879.  B r y o l . 10:51,  (Lindb.) B o u l . , Muscin. Prance 24.  Limnobium m o l l e v a r . a l p i n u m (Lindb.) Vent, at B o t t . , A t t i . c  Critt.  I t a l . s e r . 2. f a s c . 3:166.  Hypnum d i l a t a t u m v a r . a-tpinum Limnobium alpinum  Soc.  1884.  (Lindb.) Husn., Muse. G a l l . 413,  (Lindb.) Loeske. Hedwigia  Names o f taxa f o r which study m a t e r i a l was l i t e r a t u r e was  1884.  (Lindb.) Roth. E u r . Laubm. 2:642. 57f, 8.  Pla£yphyllum alpinum  1883.  50:243.  u n a v a i l a b l e and  1894. 1905.  1911.  available  inadequate f o r e v a l u a t i n g the taxon.  Hygrohypnum alpinum v a r . v i r e s c e n s Amann. F l . Mouss. S u i s s e 2:359.  1912.  Names o f t a x a f o r which the l i t e r a t u r e was Amblystegium 1883:7.  d i l a t a t u m v a r . alpinum  u n a v a i l a b l e f o r t h i s study, (Lindb.) A d l e r z . Bot.  Not,  1883.  P l a n t s u s u a l l y s o f t and p l i a b l e when m o i s t o r d r y , l e s s o f t e n somewhat s t i f f and b r i t t l e when d r y , e s p e c i a l l y i n North American m a t e r i a l ,  generally  f o r m i n g s m a l l l o o s e l y woven patches o r l e s s o f t e n l a r g e r more t i g h t l y woven mats, b o t h o f which e a s i l y fragment. yellow-brown,  Color v a r i a b l e , pale yellow, s i l t y  brownish-yellow-green o r b r i g h t y e l l o w - g r e e n , sometimes ex-  h i b i t i n g a t r a n s l u c e n t sheen. stems and stem l e a v e s g e n e r a l l y v a r i a b l e * , branches 1 - 2  Stems t o about 5 cm l o n g , l e a f y  throughout?  obscured w i t h i n the mat o r p a t c h .  Branching  (3.5) cm l o n g , c h a r a c t e r i s t i c a l l y a s c e n d i n g t o e r e c t  99 in  one  plane.  Stem c r o s s - s e c t i o n e x p o s i n g 2 to 3 (4) rows o f  small,  t h i c k - w a l l e d , y e l l o w o r organe-^rown c o r t i c a l c e l l s ; m e d u l l a r y larger, thin-walled, hyaline with l i t t l e and  d i s c o l o r a t i o n w i t h age;  ish. at  evidence of w a l l  c e n t r a l strand present,  cells  thickening  h y a l i n e o r brown-  R h i z o i d s abundant, dark reddish-brown, smooJbh w a l l e d ,  arising  the base o f v e n t r a l stem l e a v e s . Leaves v a r i a b l e ; c l o s e l y spaced o r d i s t a n t , d i f f e r i n g l i t t l e  the wet  to d r y c o n d i t i o n , though l e a f s h r i n k a g e  from  i s r e a d i l y e v i d e n t upon  d r y i n g ; c l o s e l y spaced l e a v e s a r e l o o s e l y a p p r e s s e d - i m b r i c a t e  and  either  e s s e n t i a l l y p l a n e o r r u f f l e d - c o n t o r t e d ; d i s t a n t l e a v e s are s p r e a d i n g erect-spreading Leaves  (1) 1.3  and v a r i o u s l y c o n t o r t e d by m a r g i n a l i n r o l l i n g o r - 1.7  (2) mm  long X  v a r i a b l e , u s u a l l y oblong e l l i p t i c  (0.9)  1.2  *•,].. 6  (1.7) mm  to  ruffling.  wide; shape  t o o r b i c u l a r , but v a r y i n g from o v a t e to  t r a n s v e r s e ; margins u s u a l l y e n t i r e , l e s s o f t e n f i n e l y s e r r a t e to s e r r u l a t e , e s p e c i a l l y i n the apex, v a r y i n g among l e a v e s on the same stem, u s u a l l y p l a n e throughout, o c c a s i o n a l l y s l i g h t l y r e c u r v e d  a t the base.  v a r i o u s l y p l a n e o r r u f f l e d o r l e s s o f t e n weakly concave. narrowly decurrent often very  or transverse.  f a i n t , one  base  C o s t a u s u a l l y double w i t h s l e n d e r arms,  o r both arms may  s i n g l e and/or f o r k e d w i t h  Leaf  Leaves  reach midleaf,  the main a x i s r e a c h i n g  r a r e l y slender  s l i g h t l y above m i d l e a f .  A r e o l a t i o n v a r i a b l e ; median l e a f c e l l s s h o r t f u s i f o r m t o l i n e a r uose, moderately t h i n - w a l l e d ,  (25)  35 - 55  (75) um  long X  (4) 6 - 7  f u s i f o r m ; m a r g i n a l l e a f c e l l s i n the upper h a l f o f the l e a f l i t t l e long, r a r e l y longer;  l e a f base c e l l s v a r i o u s l y become l o n g e r o r s h o r t e r i n l e n g t h o r wider o r changing v e r y  little  flex(9)  wide; a p i c a l c e l l s s h o r t e n i n g g r a d u a l l y , v a r i o u s l y o v a l , quadrate o r  from the median c e l l s , u s u a l l y 25 t o 50 um  and  um  short  different  toward  the  slightly  from the median c e l l s ; b a s a l c e l l s t h i c k e r  100 walled, few  h y a l i n e o r sometimes becoming y e l l o w i s h o r brown w i t h age, p i t s  t o none; a l a r c e l l s u s u a l l y forming  a w e l l d e f i n e d r e c t a n g u l a r group  o f t h i n - w a l l e d , sometimes s l i g h t l y i n f l a t e d and/or excavated, to r e c t a n g u l a r c e l l s , n o r m a l l y  irregular  h y a l i n e , sometimes becoming brownish w i t h  age. Plants autoicous;  p e r i g o n i a l leaves ovate,  0.5 t o 0.8 mm l o n g , eco-  s t a t e , e n t i r e , except f o r 2 t o 4 t e e t h a t t h e s h a r p l y acute apex; o u t e r p e r i c h a e t i a l leaves ovate t o ovate l a n c e o l a t e , ecostate, squarrose i n the upper h a l f ;  i n n e r p e r i c h a e t i a l l e a v e s l i n e a r t o l i n e a r - l a n c e o l a t e , up  to 4 mm l o n g , e c o s t a t e o r f a i n t l y s i n g l e o r double e c o s t a t e , 0, 2. o r 4 deep p l i c a e ; margins o f t h e m i d d l e and i n n e r p e r i c h a e t i a l l e a v e s  charac-  t e r i s t i c a l l y c o a r s e l y s e r r a t e a t t h e apex w i t h c e r t a i n c e l l s i n the apex c o a r s e l y p a p i l l o s e on t h e a b a x i a l l e a f s u r f a c e by o v e r r i d i n g d i s t a l Seta 8 t o 20 mm l o n g , u s u a l l y 9 t o 16 mm, orangish-red  endwalls.  color variously yellowish-red,  o r r e d , smooth, s t r a i g h t when wet, v a r i o u s l y t w i s t e d when d r y ;  c a p s u l e as p e r t h e genus. Outer p e r i s t o m e t y p i c a l f o r t h e genus; i n n e r p e r i s t o m e t e e t h  finely  p a p i l l o s e , c i l i a a b s e n t o r v e r y rudimentary; spores dusky y e l l o w o r smoky grey, 18  f i n e l y t o c o a r s e l y p a p i l l o s e , 13 t o 22 um i n diameter, u s u a l l y 14 t o  um.  Hygrohypnum alpinum i s a w e l l d e f i n e d s p e c i e s and can be r e c o g n i z e d r e a d i l y by i t s b r o a d l y o v a t e t o o r b i c u l a r l e a v e s which e x h i b i t a group o f t h i n - w a l l e d , u s u a l l y h y a l i n e and i r e e g u l a r t o r e c t a n g u l a r a l a r c e l l s .  The  l o n g a x i s o f t h e r e c t a n g u l a r group o f a l a r c e l l s i s p a r a l l e l t o t h e l e a f margin  ( P i g . 27 g & h ) .  C e r t a i n c e l l s i n t h e apex o f t h e i n n e r  perichaetial  l e a v e s a r e p a p i l l o s e on t h e a b a x i a l s u r f a c e by means o f o v e r r i d i n g d i s t a l  101 endwalls  ( P i g . 27 a ) .  and Meylan  T h i s u s e f u l f e a t u r e was f i r s t observed by Amann  (1912), b u t seems t o have gone u n n o t i c e d s i n c e then.  The s p e c i e s concepts o f Hygrohypnum alpinum and H. d i l a t a t u m ( W i l s . 2 ex Schimp;) Loesk.  have been i n t i m a t e l y i n v o l v e d w i t h a number o f mis-  c o n c e p t i o n s o f H. m o l l e Hedwig  (Hedw.) Loesk.  (1801) d e s c r i b e d Hypnum m o l l e and H. a l p e s t r e based on t h e e a r l i e r  concepts o f D i c k s o n Bridel  (Hedw.) Loesk. and H. a l p e s t r e  (1812, 1827),  (1790) and Swartz (1799) r e s p e c t i v e l y . s e v e r a l subsequent  Except f o r  workers t r e a t e d Hypnum a l p e s t r e Sw.  as synonymous w i t h Hypnum m o l l e D i c k s . ( S p r e n g e l , 1827; De N o t a r i s , 1838) or  r e l e g a t e d Hypnum a l p e s t r e Sw, t o v a r i e t a l s t a t u s w i t h Hypnum m o l l e D i c k s ,  (Hampe, 1837).  Schimper  (1853) p a r t i a l l y c l a r i f i e d t h e s i t u a t i o n , b u t a  broad and somewhat mistaken concept o f c e r t a i n s p e c i e s s e r v e d o n l y t o c l o u d the i s s u e .  L i k e Hedwig, Schimper based h i s concept o f Limnobium m o l l e on  that o f Dickson.  Schimper a l s o observed t h a t a l t h o u g h Swartz (1799)  a c c u r a t e l y d e s c r i b e d Hypnum a l p e s t r e , he r e p r e s e n t e d t h e s p e c i e s w i t h an i l l u s t r a t i o n t h a t Schimper f e l t t o be Hypnum m o l l e sensu D i c k s o n . emphasized  i n Schimper's  This i s  treatment o f Limnobium a l p e s t r e where he l i s t e d  Hypnum a l p e s t r e as t h e basionym, b u t s p e c i f i c a l l y e x c l u d e d Swartz* t r a t i o n , Tab. V I , r e f e r r i n g i t t o Limnobium m o l l e .  illus-  O b v i o u s l y , Schimper  noted t h a t Hypnum Stereddon a l p e s t r i s B r i d , from B r y o l b g i a U n i v e r s a I I i s synonymous w i t h Hypnum m o l l e D i c k s , and s i m i l a r l y Hypnum Stereodon B r i d . was synonymous w i t h Hypnum a l p e s t r e Sw.  However, B r i d e l  mollis  (1827) c l e a r l y  2 E v i d e n c e t o be p r e s e n t e d l a t e r has e s t a b l i s h e d t h a t the name Hypnum d u r i u sculum De Not, has p r i o r i t y o v e r Hypnum d i l a t a t u m W i l s . i n Schimp, for  However,  t h e sake o f c l a r i t y Hypnum d i l a t a t u m w i l l be employed i n t h e p r e s e n t d i s -  cussion.  102 i n d i c a t e d t h a t H. S_. a l p e s t r i s was based on Hypnum a l p e s t r e Sw, and H. S_. m o l l i s was based on Hypnum m o l l e The one,  Dicks.  concept o f Limnobium m o l l e Schimp* seems t o have been a v e r y broad  Schimper  (1853) i l l u s t r a t e d t h e s p e c i e s w i t h  two p l a t e s : Tab. I l l ,  F i g . 1 - 14, pg. 576 and Tab. IV, F i g . 2 - 6 , pg. 577.  Schimper  (1860)  e x t r a c t e d and d e s c r i b e d Hypnum alpinum Schimp. and H, d i l a t a t u m W i l s . i n Schimp. from t h e 1853 concept o f Limnobium m o l l e . new t a x a Schimper c i t e d Tab. I l l ,  To i l l u s t r a t e these two  F i g . 1 - 14. pg. 576 t o r e p r e s e n t  alpinum and Tab, IV, F i g . 2 - 6, pg. 577 t o r e p r e s e n t Fufclfcher, Sfchimper  Hypnum  Hypnum d i l a t a t u m .  (1860) c i t e d no i l l u s t r a t i o n f o r h i s more r e s t r i c t e d  t r e a t m e n t o f Hypnum m o l l e ,  b u t c l e a r l y based i t on D i c k s o n ' s concept.  An  e x a m i n a t i o n o f t h e few a v a i l a b l e Schimper specimens i n d i c a t e s t h a t he h e l d a sound concept o f Hypnum m o l l e ,  H. d i l a t a t u m ,  H, alpinum and H. a l p e s t r e .  I t i s , t h e r e f o r e , remarkable t h a t : 1. t h e i l l u s t r a t i o n c i t e d t o r e p r e s e n t Limnobium a l p e s t r e i n B r y o l o g i a Europaea and r e d e s i g n a t e d c l e a r l y Hygrohypnum m o l l e  i n Syn, ed I i s  (Hedw.) Loesk. and, 2. t h a t i n Syn. ed. I , he  c l e a r l y a s c r i b e s t h e Swactz i l l u s t r a t i o n o f Hypnum a l p e s t r e t o Hypnum alpinum Schimp. As p r e s e n t l y understood Hygrohypnum m o l l e (Wils. i n Schimp.) Loesk., H. alpinum  (Lindb.)  Loesk. a r e v e r y c l e a r l y d e f i n e d s p e c i e s .  (Hedw.) Loesk., H. d i l a t a t u m Loesk, and H. a l p e s t r e (Hedw,)  Of t h e i l l u s t r a t i o n s i n B r y o l o g i a  Europaea, Tab. I I I . F i g . 1 - 14, pg. 576 ace r e f e r a b l e t o Hygrohypnum alpinum, Tab.  IV. F i g . 2 - 6 , pg, 577 are r e f e r a b l e t o H, d i l a t a t u m and Tab. IV, F i g , 1,  l b and 7 - 21, pg. 577 are r e f e r a b l e t o H, m o l l e . I t i s worth n o t i n g t h a t Hedwig's alpestre i s misleading. The  I t erroneously  (1801) Tab. LXIV. F i g . 2 f o r Hypnum shows a s i n g l e c o s t a i n the l e a v e s .  c o s t a o f Hygrohypnum a l p e s t r e i s u s u a l l y s h o r t and double o r l o n g and  103 double.  Similarly,  shows a f a l c a t e  Tab. IV, P i g . 7, pg. 577 from B r y o l o g i a  Europaea  l e a f w i t h a s i n g l e c o s t a , a c h a r a c t e r combination t h a t  i s unknown i n Hygrohypnum m o l l e . As l a t e i s 1927 Monkemeyer e r r o n e o u s l y r e p r e s e n t e d Hygrohypnum d i l a tatum w i t h an i l l u s t r a t i o n o b v i o u s l y o f H. m o l l e ( F i g . 168 b ) .  The  sim-  i l a r i t y o f H. alpinum and H. d i l a t a t u m l e d Lawton (1971) t o employ a l e a f o f H. alpinum t o r e p r e s e n t H.  dilatatum  ^ P l a t e 156, F i g . 1 ) .  As t h e r e has been c o n s i d e r a b l e c o n f u s i o n o v e r s e v e r a l t a x a i n the ^Hygrohypnum m o l l e complex", so too has t h e r e been c o n f u s i o n c o n c e r n i n g the n o m e n c l a t o r i a l s t a t u s o f Hygrohypnum alpinum.  Schimper  (1876) d e s c r i b e d  Hypnum alpinum Schimp. as a s e g r e g a t e from Limnobium m o l l e Schimp. U n f o r t u n a t e l y , Hypnum alpinum Schimp. i s a l a t e r homojiym o f Hypnum alpinum Web.  e t Mohr.  The f i r s t name t o be v a l i d l y and l e g a l l y a p p l i e d t o Schimper's  taxon was Amblystegium  m o l l e v a r . alpinum i n L i n d b e r g (1879).  i n d i c a t e d t h a t the v a r i e t y was er  Of the f i v e specimens  Schimper  cited a Blytt  from the Norwegian D o v r e f j e l d and a specimen c o l l e c t e d Nedalen i n the Harjedalem r e g i o n o f Sweden. import.  Schimp-  specimens, but d e s i g n a t e d no h o l o t y p e .  two a r e i m p o r t a n t .  i s of l i t t l e  Lindberg c l e a r l y  based on Schimper's o r i g i n a l c o n c e p t .  (1876) based the s p e c i e s on 5 c i t e d  NSrway  (With.)  specimen  by Thedenius  T h a t Nedalen i s now  from  a part of  These two specimens have h o t been seen.  However, a B l y t t specimen c a l l e d Hypnum m o l l e from Gudbrandsdalen,  Norway,  a s i t e not f a r from the D o v r e f j e l d , and a Thedenius specimen c a l l e d Hypnum d i l a t a t u m from Tannas i n the H a r j e d a l e n d i s t r i c t o f Sweden and a g a i n a s i t e not  f a r from Hedalen, are p r e s e n t a t S-PA.  examples o f Hygrohypnum alpinum. o f the two.  Both o f t h e s e specimens a r e v a l i d  The B l y t t specimen  i s i n the b e t t e r c o n d i t i o n  In the absence o f any proven Schimper m a t e r i a l o f Hygrohypnum  alpinum, the B l y t t specimen i s s e l e c t e d as the neotype.  104 Hygrohypnum alpinum e x h i b i t s some v a r i a t i o n i n l e a f shape, c o s t a s t r u c t u r e , the n a t u r e o f the l e a f margin and the a t t i t u d e o f the l e a v e s upon the stem. 26 a - e ) .  L e a f shape v a r i e s from b r o a d l y o v a t e to o r b i c u l a r ( F i g .  The l e a f apex may  be b r o a d l y a c u t e o r rounded w i t h a s m a l l ,  a b r u p t a p i c u l u s ( F i g . 26 a s ~ e ) , b u t r a r e l y one may mid-leaf  The c o s t a i s u s u a l l y s l e n d e r and  encounter a s l e n d e r , s i n g l e and/or  ( F i g . 26 a - e ) .  Schimper's  s e r r u l a t e margin a t the l e a f apex.  double,  forked costa reaching  (1876) d e s c r i p t i o n noted the  finely  The p r e s e n t study has shown the margin  t o the l e a f apex t o v a r y from e n t i r e t o f i n e l y s e r r u l a t e between l e a v e s on the same o r d i f f e r e n t p l a n t s .  One  o f the most remarkable  features of  H. alpinum i s the d i s p o s i t i o n o f i t s l e a v e s a l o n g the stems. all  the l e a v e s o f North American  European  specimens and a s i g n i f i c a n t number o f  specimens e x h i b i t l e a v e s t h a t a r e c o n v o l u t e o r r u f f l e d  to t h e i r l o n g i t u d i n a l a x i s  ( F i g . 26 f , g & h ) .  Other European  e x h i b i t l e a v e s t h a t a r e e s s e n t i a l l y p l a n e o r s l i g h t l y concave i  Virtually  parallel specimens  ( F i g , 26  &j). Hygrohypnum alpinum i s most e a s i l y confused w i t h H. d u r i u s c u l u m .  The  f o l l o w i n g t a b l e o f c h a r a c t e r s w i l l h e l p t o s e p a r a t e them.  H.  alpinum  H. d u r i u s c u l u m  A l a r c e l l s t h i n - w a l l e d , more o r l e s s r e c t a n g u l a r , and h y a l i n e .  A l a r c e l l s t h i c k - w a l l e d , quadrate to i r r e g u l a r , u s u a l l y d i s c o l o r e d .  Apical perichaetial leaf cells p a p i l l o s e by a n t e r i o r l y o v e r r i d i n g end w a l l s .  Perichaetial leaf cells  Endostomial c i l i a absent o r v e r y rudimentary.  One o r two w e l l t o p o o r l y developed c i l i a between the e n d o s t o m i a l t e e t h .  Stem branches u s u a l l y l e s s than 2 cm l o n g , more o r l e s s ascends ing to e r e c t .  Stem branches o f v a r i a b l e l e n g t h , u s u a l l y p r o s t r a t e or ascending.  smooth.  105 O l d e r stems g e n e r a l l y r e t a i n ing t h e i r leaves.  In  O l d e r stems g e n e r a l l y denuded of leaves.  Europe Hygrohypnum alpinum and H. d u r i u s c u l u m can be  e a s i l y s e p a r a t e d by f i e l d o b s e r v a b l e c h a r a c t e r s .  Hygrohypnum  fairly alpinum  n o r m a l l y forms s o f t , e a s i l y fragmenting patches o f t i g h t l y woven,  stol-  o n i f e r o u s stems, from which a r i s e numerous s h o r t , a s c e n d i n g t o e r e c t branches.  The s t o l o n i f e r o u s stems a r e g e n e r a l l y f o l i o s e throughout  densely clothed i n r h i z o i d s v e n t r a l l y . to  Hygrohypnum d u r i u s c u l u m i s r i g i d  the touch and the p r o s t r a t e , i r r e g u l a r l y branched  stems a r e r e g u l a r l y  denuded i n the o l d e r e x t r e m i t i e s . R h i z o i d s a r e s p a r s e o r absent. of  and  Branches  i r r e g u l a r p o s i t i o n and v a r i a b l e l e n g t h u s u a l l y l i e p a r a l l e l t o the stem  and a r e p r o s t r a t e t o s l i g h t l y a s c e n d i n g a t the t i p s . do not fragment In  The patches o r  tufts  easily.  western North America  one must i n i t i a l l y r e l y ap m i c r o s c o p i c  c h a r a c t e r s t o s e p a r a t e H. alpinum and H. d u r i u s c u l u m .  In t h i s r e g i o n  H. d u r i u s c u l u m o f t e n assumes a h a b i t s i m i l a r t o t h a t o f H. alpinum, i . e . d e n s e l y f o l i o s e , and d e n s e l y r h i z o i d a l s t o l o n i f e r o u s stems which bear numerous s h o r t , e r e c t o r a s c e n d i n g b r a n c h e s . p e r i e n c e one (Fig.  With i n c r e a s e d f i e l d  can employ the r u f f l e d appearance o f the l e a v e s H.  ex-  alpinum  26 g & h) t o s e p a r a t e i t from H, d u r i u s c u l u m whose l e a v e s a r e g e n e r a l l y  l e s s crowded and more symmetrical, p l a n o - c o n c a v e - ( F i g . 29 c ) , My own America  f i e l d e x p e r i e n c e w i t h Hygrohypnumalpinum i n Western North  has been c o n f i n e d t o southern B r i t i s h Columbia.  In t h a t r e g i o n the  s p e c i e s o c c u r s e x c l u s i v e l y on l a r g e b o u l d e r s on the banks o f o r i n the middle o f s m a l l , s w i f t l y r u n n i n g mountain streams.  The o n l y time t h a t  this  s p e c i e s o c c u r s i n a submerged s i t u a t i o n f o r any l e n g t h o f time i s d u r i n g spring runoff.  Otherwise,  spray from t u r b u l e n t water.  i t o c c u r s on emergent r o c k s t h a t a r e s u b j e c t e d t o  106 Fig.  26 a - j . of  Hygrohypnum alpinum.  a - e. f.  V a r i a t i o n i n the l e a f shape and the h a b i t o f the s h o o t s  Variation i n foliage leaves.  J  A m o i s t s h o o t o f a specimen w i t h r u f f l e d  l e a v e s from  Western  N o r t h America. g - h.  Comparison ruffled  i  - j .  between the m o i s t  l e a f e d specimen  Comparison  (g) and d r y ; (h) shoot o f a  from western North America.  between the m o i s t  ( i ) and d r y ( j ) shoot o f a  p l a n e t o concave l e a f e d specimen from Europe and N o r t h America.  Scale: a - e?  I  f - j;  Each stem i s a p p r o x i m a t e l y 1 cm  1mm  I long.  107  108  F i g . 27 a - h .  C e l l u l a r d e t a i l o f t h e p e r i c h a e t i a l and f o l i a g e  of Hygrohypnum a l p i n u m . a.  Perichaetial  b - c. d.  Foliage  l e a f apex, leaf  apicies.  Median c e l l s o f f o l i a g e  leaves,  e - f.  Marginal c e l l s o f f o l i a g e  g - h.  Alar cells of foliage  leaves,  leaves.  Scale: a j  1  100 um  . I , 100"m  b - c, g - h ; d - f.  I  I  -•  100um  I I  —-  leaves  110  F i g . 28.  Hygrohypnum alpinum  Ill Exsiccati  Examined  Macoun, Canadian Musci # 357 as H, d i l a t a t u m . (TRTC) # 396 as !H. a l p e s t r e , b u t i n p a r t H. l u r i d u m . (USA, NY, MO) # 900 as H, d i l a t a t u m . (USA) C a n i d i a n Mosses, # 490 as H. d i l a t a t u m . (USA, NY) Grout, North American Musci P l e u r o c a r p i , # 427 as H. d i l a t a t u m . (MIN, MO, NY, TENN, UC, USA) Bauer, Musci europ. e t amer. a x s i c c a t i , # 2089. (BRNM, NY S-PA) Rabenhorst, B r y o t h e c a europaea, # 1348. (NY, S-PA) Husnot, Musci G a l l i a e , # 593 as H. p a l u s t r e v a r . julaceum. (S-PA) Verdoorn, Musci S e l e c t i e t C r i t i c i , s e r i e s V I , # 270. (MICH, NY, LE, TENN) S e l e c t e d Specimens  Examined  Canada  B r i t i s h Columbia G a r i b a l d i P r o v i n c i a l Park, S e n t i n e l G l a c i e r Area? S c h o f i e l d 32953 as H. a l p e s t r e . (CAMN, DUKE, TENN)-• North Vancouver, Lynn Canyon; S c h o f i e l d 35292 as H. d i l a t a t u m . (HBP) <i?C* C a r i b o u D i s t r i c t , end o f Long Lake; Boas 688 as H. d i l a t a t u m . (CAN, UBC) G l a c i e r ; Macoun as H. d i l a t a t u m IN Canadian Musci # 900 (USA) Kamloops Lake D i s t r i c t , Skuian Creek; Brinkman 259 as H. d i i a - . , tatum. (CAN, MICH) Lake Lindeman; W i l l i a m s 6 May 1896 as H. d i l a t a t u m . (NY) Salmo, Sheep Creek; Jamieson 5551. (UBC) United States Alaska St. ?acrous Bay; Howell 18 Aug 1895 as H. d i l a t a t u m i n • Howell's A l a s k a P a c i f i c C o a s t P l a n t s # 1818. (NY) Short Bay; Howell 8 Aug 1895 as H. m o l l e . (MIN) Washington Cascades Mtns, Copper Creek; A l l e n , June 1905 as H. d i l a t a t u m i n North American Musci P l e u r o c a r p i # 427. (MICH, NY, TENN, WTU, MO, UC, USA) Snohomish Co., Monte C r i s t o ; S c h o f i e l d 20085 as H. d i l a t a t u m . (CAN, S-PA) C l a l l a m Co, Olympic N a t i o n a l Park, Olympic H o t s p r i n g s ; S v i l h a 758 as H. d i l a t a t u m , (WTU; F) Mount R a i n i e r Region, P a r a d i s e V a l l e y ; F r y e 11 Aug 1904 as H. d i l a t a t u m . (WTU) Chelan Co., T r a n s e n Creek; Sharp 2112. (TENN) Idaho Bonner Co., P r e i s t Lake; I r e l a n d 8565 as H. d i l a t a t u m . (ALTA, UBC) Lemhi I n d i a n R e s e r v a t i o n , M i l l Creek; Henderson 3979 as H. dilatatum. (USA) V Montana G l a c i e r N a t i o n a l Park, Lake MacDonald; S t a n d l e y 18388 as H. alpinum. (USA)  112  M i s s o u l a Co. R a t t l e s n a k e Campground; Sharp & C l e b s c h M-16. (TENN) B e l t Mountains; W i l l i a m s 19 Sept 1891 as H. a r c t i c u m , admixture o f H. b e s t i i and H. d i l a t a t u m . (MIN) California S i s k i y o u Co., Duck Lake Creek; N o r r i s 23300 as H. d i l a t a t u m . (HSC) France Dep. de I ' I s e r e , V a l l o n des Etages; Culmann 2 Sept 1927 i n Musci europ. e t amer. e x s i c c a t i # 2089. (S-PA, TENN, NY, BRNM) Haute S a v o i e , Mt. B l a n c ; Payot, i n Musci G a l l i a e # 593. (S-PA) Switzerland V a l a i s , Mattmark; N i c h o l s o n & Dixon 7 J u l y 1913. (BM) Berner, Oberlander, Handegg; Culmann Sept 1911 i n Musci S e l e c t ! e t C r i t i c i , s e r i e s V I , # 270. (MICH, NY, TENN) T e s s i n , T i c i n o , Compolungopass; Traumann 8 June 1908, (S-PA) U r i , A l p e t t i ; Culmann 7 Aug 1882. (S-PA) Austria T i r o l , O t z a l e r A l p e n ; Berner J u l y 1931. (S-PA) Vorarlberg, Arlberg; Eggler. (S-PA) S a l z b u r g , Kresmuler; Loeske 2 Aug 1903. (S-PA) Karnten, M a l t a ; B r e i d l e r 1885. (S-PA) Italy Macuguaga, V a l Anyasca; N i c h o l s o n & Dixon 10 J u l y 1913. (BM) Iceland N. V, D y n j a u v i ; Hesselbo 26 June 1912. (NY) Norway Opdal, D r i v a ; K a u r i n Okt 1881. (S-PA) Gausta; JMderholm J u l y 1895. (S-PA) Hordaland, Hardanger; K o t i l a i n e n 2 Sept 1931. (S-PA) Gudbrandsdalen; B l y t t as H. m o l l e . NEOTYPE, (S-PA) tEroms, Lyngen, Kjosen; T. U l s i n e n 20 Sept 1968. (LE) Sweden H a r j e d a l e n , S t o r s j o ; Thedenius J u l y 1842. (S-PA) Jamtland, F r o s t v i k e n s ; Nyman 6 Aug 1947. (S-PA) A s e l e Lappmark, M a r s f j a l l e t , J i l k e l m i n i a r Hulphers 14 June 1943. (S-PA) L y c k s e l e Lappmark, S t o r f j a l l e t back v i d V a l l i n t j a k k o ; Normann 15 Aug 1945. (S-PA) P i t e Lappmark, A r j e p l o g s ; M o l l e r 30 J u l y 1918. (S-PA) L u l e Lappmark, Kamajokk; NymanaAug 1893. (S-PA) Torne Lappmark, Karevaggejokk; N i c h o l s o n & Dixon 26 Aug 1907. (BM) ,;  113 Hygrohypnum d u r i u s c u l u m (De Not.) Jamieson n o v . comb. Lectotype: Erbar. C r i t t o g . I t a l ,  s e r . I I , no. 204. (<§)  Limnobium d u r i u s c u l u m De Not., E r b a r . C r i t t .  Ital.  ser. I I ,  no. 204. 1869. Hypnum d i l a t a t u m W i l s . i n Schimp. Syn. Ed. 2. 776. 1876. (  Amblystegium  dilatatum  Limnobium d i l a t a t u m Soc. C r i t t .  (Wils.) L i n d b . Musci Scand. 33. 1879.  ( W i l s . i n Schimp.) Vent, e t B o t t . ,  Ital,  s e r . 2, 3:166.  Atti.  1884.  Hypnum m o l l e v a r . d i l a t a t u m ( W i l s . i n Schimp.) B o u l . Muscin. France.  24.  1884.  Hypnum p s e u d o - a r c t i c u m Kindb., B u l l . T o r r e y Bot. C l u b 17(11):280. 1890 Hypnum c i r c u l i f o l i u m Cs»: M. & Kindb. i n Macoun, C a t . Canad. P I . 6:242.  1892.  Calliergon dilatatum 6:72.  ( W i l s . i n Schimp.) Kindb,, Canad. Rec. S c i .  1894.  Calliergon circulifolium  (C. M. & Kindb, i n Macoun) Kindb.,  Canad, Rec. S c i . 6:72. Limnobium pseudo-arcticum 6:74.  (Kindb.) Kindb., Canad, Rec. S c i .  1894.  Hypnum m o l l e s s p . d i l a t a t u m Handb. B r i t . Mosses. C a l l i e r g o n pseudo-arcticum 1:85.  1894.  (Wils. i n Schimp.) D i x . , Stud. 483. 1896. (Kindb.) Kindb., E u r . N, Am,  Bryin,  1897.  Hypnum eugyrium v a r . d i l a t a t u m Handls. Mies. 345,  ( W i l s . i n Schimp.) Grout, Mosses  1903.  Hypnum d i l a t a t u m v a r . d u r i u s c u l u m (De Not.) Limpr., Laubm. Deutsch. 3:353.  1904.  11.4 Limnobium d i l a t a t u m v a r . d u r i u s c u l u m Laub. 2:645.  (De Not.)  Roth, E u r .  1905,  Hygrohypnum pseudo-arcticum  (Kindb.) B r o t h , ,  Nat. P f l . 1(3):1040,  1909. Hygrohypnum c i r c u l i f o l i u m Nat.  P f l . 1(3):1039.  (C, M. & Kindb. i n Macoun) B r o t h , , 1909.  Hygrohypnum d i l a t a t u m v a r . d u r i u s c u l u m Mouss. S u i s s e 2:361.  Amann, F l .  1912.  Hygrohypnum c o r d i f o l i u m Okam., J , C o l l . 36:30.  (De Not.)  Imp. U n i v . Tokyo  14. 1915.  Names o f taxa f o r which n e i t h e r t h e a p p r o p r i a t e specimens were a v a i l a b l e d u r i n g t h i s  l i t e r a t u r e nor any  study.  Hygrohypnum d i l a t a t u m v a r . callineurum-Amann, B u i . Soc. Vaudoise Sc. Nat. 53:123.  Plants u s u a l l y coarse  1920.  and s t i f f  t o t h e touch, r a r e l y s o f t and f l e x i b l e ,  forming l o o s e l y t o t i g h t l y woven mats o r p a t c h e s , r a r e l y deep c u s h i o n s o r tufts.  Color v a r i a b l e , b r i g h t yellow-green with r u s t y mottling, d u l l  green w i t h r u s t y m o t t l i n g , d i r t y y e l l o w mottling, a l l c o l o r s grading  olive-  t o yellow-brown w i t h o r w i t h o u t r u s t y  i n t o one another a t v a r i o u s times.  Stems (1)  2-7  (9) cm l o n g , f o l i o s e throughout o r denuded from t h e base t o more than  half  the l e n g t h o f the stem.  B r a n c h i n g i r r e g u l a r , u s u a l l y a s c e n d i n g from  procumbent stems; o f t e n t h e o n l y l e a f y p a r t o f t h e p l a n t , b u t sometimes denuded a t t h e base.  Stem c r o s s - s e c t i o n s r e v e a l i n g 3 t o 4 rows o f s m a l l ,  t h i c k - w a l l e d y e l l o w i s h t o reddish-brown c o r t i c a l c e l l s ; m e d u l l a r y  cells  l a r g e r , t h i n - w a l l e d , sometimes becoming t h i c k e r w a l l e d  and d i s c o l o r e d w i t h  age;  Rhizoids v a r i a b l e ,  c e n t r a l s t r a n d w e l l developed, o f t e n d i s c o l o r e d .  infrequent o r densely  c l o t h i n g t h e v e n t r a l s u r f a c e o f p r o s t r a t e stems, a r i s i n g  115 from t h e bases o f v e n t r a l stem o r b r a n c h l e a v e s . Leaves v a r i a b l e , c l o s e l y spaced t o w i d e l y d i s t i n c t , d i f f e r i n g i n appearance w i t h i n and between t h e wet and d r y c o n d i t i o n s !  Wet c o n d i t i o n ,  l e a v e s appressed i m b r i c a t e o r s p r e a d i n g and somewhat c o n t o r t e d ,  some c l o s e l y  spaced appressed i m b r i c a t e l e a v e s a r e o f t e n secund, o t h e r c l o s e l y leaves are spreading  to e r e c t spreading  s t i l l o t h e r c l o s e l y spaced l e a v e s s i m p l y  and v a r i o u s l y c o n t o r t e d , appressed i m b r i c a t e ;  spaced while  widely  spaced l e a v e s when wet, v a r i o u s l y l o o s e l y appressed i m b r i c a t e o r s p r e a d i n g : Upon d r y i n g , a l l l e a v e s whether l o o s e l y spaced o r d i s t a n t , a p p r e s s e d i m b r i c a t e , secund o r s p r e a d i n g , width, spreading  undergo v a r y i n g degrees o f s h r i n k a g e i n  and c o n t o r t i o n ; t h e c o n t o r t i o n may i n v o l v e t w i s t i n g ,  u s u a l l y toward the stem and t o t h e l e f t o r i n r o l l i n g on t h e margins w i t h concomitant t w i s t i n g . , one dry,  Leaves may v a r y  r e g i o n o f the stem t o another; sometimes a p p e a r i n g  moist.  Leaves  from c l o s e l y t o w i d e l y  spaced from  Leaves s t r a i g h t when wet, c o n t o r t e d when  f a l c a t e from the s h r i n k a g e ,  never t r u l y f a l c a t e when  (0.7) 1.25 - 1.8 (2.3) mm l o n g X (0.6) 1.0 - 1 . 4 (1.8) mm wide;  shape u s u a l l y o b l o n g - e l l i p t i c t o b r o a d l y o v a t e ,  l e s s o f t e n o r b i c u l a r o r ovate;  margins u s u a l l y e n t i r e , o f t e n uneven o r u n d u l a t i n g ,  rarely clearly serrulate,  o f t e n narrowly r e f l e x e d i n t h e a l a r r e g i o n ; p l a n e o r s h a l l o w l y concave; apex v a r i o u s l y acute, o b t u s e , o b t u s e w i t h ^a s m a l l a p i c u l u s o r rounded; l e a f base o f t e n narrdwly d e c u r r e n t  a t t h e p o i n t o f i n s e r t i o n and sometimes c l a s p i n g  the stem w i t h a narrow c o l l a r ; c o s t a v a r i a b l e , u s u a l l y s h o r t and double w i t h s l e n d e r f o r k s r e a c h i n g t o m i d l e a f , o f t e n s h o r t and double and v e r y  faint,  s h o r t and s i n g l e , o r s i n g l e and/or b i - o r t r i f u r c a t e r e a c h i n g w e l l above the m i d d l e . Areolatiqn  exteemely v a r i a b l e ; Median l e a f c e l l s s h o r t f u s i f o r m t o l o n g  l i n e a r f l e x u o s e , r a r e l y l o n g rhombic, v a r i o u s l y t h i n t o moderately t h i c k -  116 walled?  (26) 45 - 70 (104) um l o n g X (3) 5 - 6 (8) um wide, c e l l  dimen-  s i o n s v a r i a b l e between l e a v e s on t h e same s t e m and o f t e n d i f f e r i n g one  from  s i d e t o t h e o t h e r o f the same l e a f ? c e l l s s h o r t e n i n g g r a d u a l l y toward  the apex, a p i c a l c e l l s shape v a r i a b l e , f u s i f o r m , s h o r t rhombic, rounded rectangular  t o rounded quadrate? m a r g i n a l c e l l s u s u a l l y s h o r t e r than  cent laminal c e l l s ,  adja-  25 t o 50 um l o n g , a few i s o l a t e d c e l l s may r a r e l y ex-  ceed 60 um, e s p e c i a l l y i n l a r g e European specimens; c e l l s u s u a l l y i n c r e a s i n g in  l e n g t h , widfeh and w a l l t h i c k n e s s toward t h e l e a f base, shape d i f f e r i n g  little  from median c e l l s ?  b a s a l c e l l s t h i c k - w a l l e d , p i t s few t o none, be-  coming d i s c o l o r e d w i t h age, y e l l o w i s h t o reddish-brown d i s c o l o r a t i o n sometimes c r e a t i n g a weak s u n b u r s t  e f f e c t a c r o s s t h e base? a l a r c e l l s forming a w e l l  d e f i n e d group o f t h i c k - w a l l e d , q u a d r a t e t o s h o r t r e c t a n g u l a r c e l l s , o r i r r e g u l a r , plane to deeply and  excasrated,  reaching  3 t o 5 rows from t h e margin  4 t o 5 c e l l s up t h e margin from t h e l e a f base. P l a n t s autoicous?  long, ecostate,  p e r i g o n i a l l e a v e s o v a t e l a n c e o l a t e , 0.6 t o 0,75 mm  e n t i r e o r s l i g h t l y s e r r a t e a t t h e apex, d e e p l y  p e r i c h a e t i a l l e a v e s o v a t e l a n c e o l a t e , weakly e o s t a t e from above t h e middle?  concave;  to ecostate,  squarrose  i n n e r p e r i c h a e t i a l l e a v e s l i n e a r l a n c e o l a t e , up t o  3 mm l o n g , e c o s t a t e o r w i t h a v e r y f a i n t , l o n g s i n g l e o r f o r k e d c o s t a ing  outer  t o 3/4 o f t h e l e a f l e n g t h ,  reach-  2 t o 4 l o n g p l i c a e , margins e n t i r e o r sometimes  s l i g h t l y s e r r a t e i n t h e apex, l e a f c e l l s a l l smooth. Seta 9 t o 27 mm l o n g , u s u a l l y 12 t o 20, c o l o r v a r i o u s l y y e l l o w i s h - r e d , r e d o r maroon, smooth, s t r a i g h t when wet, v a r i o u s l y t w i s t e d when d r y . t y p i c a l f o r t h e genus, annulus o f 2 t o 3 rows o f deciduous  Capsule  cells.  Peristome c h a r a c t e r i s t i c o f t h e genus; endostome u s u a l l y w i t h J t o 3 w e l l defined c i l i a , (11)  sometimes rudimentary; spores smoky brown, f i n e l y p a p i l l o s e ,  15 - 19 (23) um i n d i a m e t e r . ^  117 Hygrohypnum d u r i u s c u l u m  i s a h i g h l y v a r i a b l e s p e c i e s and i s o n l y  s l i g h t l y l e s s complex than H. luraldum.  Under t h e microscope,  the species  can be r e c o g n i z e d b e s t by i t s u s u a l l y o b l o n g - e l l i p t i c t o b r o a d l y leaves  ovate  ( F i g . 30 a - c) and t h e w e l l d e f i n e d group o f t h i c k - w a l l e d , u s u a l l y  d i s c o l o r e d , quadrate,  s h o r t r e c t a n g u l a r c*r s l i g h t l y i r r e g u l a r a l a r  cells  ( F i g . 31 c , e - h ) . Coarseness and r i g i d i t y o f h a b i t and d e f o l i a t i o n o f t h e o l d e r p a r t s o f t h e stems have l o n g been used as f i e l d c r i t e r i a c h a r a c t e r i z i n g H. duriusculum.  Though u s e f u l c h a r a c t e r s , they s h o u l d be used w i t h  discretion.  C e r t a i n specimens from t h e A u s t r i a n and Swiss A l p s and t h e Norwegian mountains a r e v e r y s o f t t o t h e touch and c o u l d be confused In North America H. duriusculum, and have d e f o l i a t e d o l d e r stems. foliose The  w i t h H. m o l l e .  H. b e s t i i and H, s m i t h i i a r e c o a r s e , Furthermore, H_. d u r i u s c u l u m  stiff  i s sometimes  throughout. a t t i t u d e o f t h e l e a v e s on t h e stems a l l o w s one t o r e c o g n i z e s e v e r a l  growth forms, which o f t e n i n t e r g r a d e . geographically. mats o r patches  These forms can be r o u g h l y  Hygrohypnum d u r i u s c u l u m on r o c k s i n streams.  correlated  most commonly forms low appressed  I n Europe and E a s t e r n North America i t  i s common f o r t h e l e a v e s t o be r a t h e r d i s t a n t a l o n g t h e stems ( F i g . 29 a - d ) . When d r y , t h e l e a v e s o f European specimens a r e c o n s p i c u o u s l y shrunken In t h e shrunken c o n d i t i o n t h e l e a v e s become v a r i o u s l y c o n t o r t e d ,  ( F i g . 29 f ) ,  twisting  i n various d i r e c t i o n s o r spreading-squarrose.  There appears t o be no r e g u l a r i t y  i n the c o n t o r t i o n o r spreading o f the leaves.  L e a f s h r i n k a g e upon d r y i n g  sometimes g i v e t h e s u g g e s t i o n o f f a l c a t i o n , b u t t h i s i s l o s t upon r e w e t t i n g . Leaf shrinkage distant leaves.  i s l e s s pronounced i n E a s t e r n North American m a t e r i a l w i t h The p a t t e r n o f c o n t o r t i o n i s more r e g u l a r .  o f specimens from E a s t e r n North America a r e p r e d o m i n a n t l y  The d r y l e a v e s  widely to e r e c t  -118 s p r e a d i n g , sometimes almost s q u a r r o s e ( F i g . 29 d ) .  The l e a v e s a r e  f u r t h e r c o n t o r t e d ; e i t h e r t w i s t e d upon themselves o r w i t h t h e i r margins i n r o l l e d t o form a c h a n n e l and then t w i s t e d .  P l a n t s o f t h i s form a r e  most common i n t h e New England S t a t e s , New York, t h e n o r t h e r n A p p a l a c h i a n mountains, Quebec and t h e M a r i t i m e p r o v i n c e s .  Some specimens from Europe  and E a s t e r n North America b e a r i n g d i s t a n t l e a v e s have t h e appressed i m b r i c a t e d .  T h e i r l e a v e s a r e n o t so o b v i o u s l y  leaves  loosely  shrunken.  Another form common t o Europe and E a s t e r n North America, which may a l s o r e a c h i n t o Western North America b e a r s i t s .leaves r a t h e r c l o s e t o gether  ( F i g . 30 h ) .  The l e a v e s a r e n o t as c o n s p i c u o u s l y Shrunken and a r e  l o o s e l y apressed imbricate.  Though t h i i form  and t h e p r e v i o u s ones a r e  sometimes remarkable d i s t i n c t , they do i n t e r g r a d e .  T h i s would  the forms a r e responses t o some e n v i r o n m e n t a l phenomenon.  imply t h a t  Certain  specimens  can be found i n both Europe and North America which bear d i s t i n c t , more o r l e s s s p r e a d i n g l e a v e s i n t h e lower p a r t o f t h e stem and more o r l e s s  closely  spaced appressed i m b r i c a t e l e a v e s a t o r near t h e stem apex o r v i c e v e r s a . A v a r i a t i o n o f t h e s e p l a n t s w i t h c l o s e l y spaced, l o o s e l y appressed i m b r i c a t e l e a v e s , o c c u r r i n g e r r a t i c a l l y throughout t h e e n t i r e range o f t h e s p e c i e s , i s one i n which t h e appressed i m b r i c a t e l e a v e s a r e o f t e n swept t o one s i d e o f the stem c r e a t i n g a secund appearance  ( F i g . 29 g S h ) .  Certain  European  m a t e r i a l o f t h i s type has l e a v e s t h a t a r e s l i g h t l y more shrunken than those from S c a n d i n a v i a o r North America.  T h i s form, t o o , grades  imperceptibly  i n t o o t h e r forms. In Western North America h a b i t v a r i a t i o n e x h i b i t s another p a t t e r n . The p l a n t s tend t o form low, p r o s t r a t e , d e n s e l y woven mats o f denuded o r f o l i o s e stems b e a r i n g numerous a s c e n d i n g t o e r e c t , d e n s e l y f o l i a t e d b r a c h e s , which seldom exceed 1.5 cm i n l e n g t h .  Leaves i n t h i s form a r e u s u a l l y  distant,  119 though  they may  v a r y from s p r e a d i n g c o n t o r t e d o r  Growth forms i n Europe and North America l e a v e s u s u a l l y form l o o s e l y woven mats. difficult  t o d i s t i n g u i s h branches  t h a t have w i d e l y spaced  In these p l a n t s i t i s o f t e n  from stems as both may  impofctant i s the f a c t t h a t the stems o r branches than the 1.5  secund.  be l e a f y .  More  are u s u a l l y much l o n g e r  cm l o n g branches o f Western North American  material.  I f herbarium m a t e r i a l can be taken as a measure o f the h a b i t d i v e r s i t y and the q u a l i t y o f growth reached i n d i f f e r e n t l o c a l i t i e s and under d i f f e r e n t environmental c o n d i t i o n s , then i t can be s a i d t h a t Hygrohypnum d u r i u s c u l u m reaches i t s most f a v o r a b l e growth p o t e n t i a l i n the h i g h A l p s o f A u s t r i a , S w i t z e r l a n d and a d j a c e n t I t a l y ,  Numerous specimens c o l l e c t e d by  Glowacki, Ganders e t a l , i n the l a s t c e n t u r y show t h a t , i n the  Breidler,  aforementioned  a r e a s H. d u r i u s c u l u m forms l a r g e , t h i c k c u s h i o n s o r almost t u r f - l i k e mats. Only r a r e l y does m a t e r i a l from o t h e r a r e a s approach  the s i z e a t t a i n e d i n the  high A l p s . In  N o r t h America  and e x t r a - a l p i n e a r e a s o f Europe,  Hygrohypnum d u r i u s c u l u m  o c c u r s more f r e q u e n t l y as the w e l l known, e a s i l y fragmenting mat o r p a t c h o f stiff,  naked stems b e a r i n g many fewer l e a f y  branches.  C e r t a i n depauperate o r o t h e r w i s e anomalous specimens from E a s t e r n North America have been confused w i t h o t h e r taxa o r a r e p u z z l i n g .  Three  specimens  c o l l e c t e d by A, J . Sharp from L a u r e l F a l l s and R o a r i n g Fork i n the G r e a t Smokey Mountains  N a t i o n a l Park o f Tennessee have been c i t e d i n the p a s t as  e v i d e n c e f o r the presence o f H. c o c h l e a r i f o l i u m i n e a s t e r n North One  o f t h e s e specimens was  i n # 59 o f North American  issued  as Hygrohypnum c o c h l e a r i f o l i u m by A. J . Grout  Musci P l e u r c a r p i Supplement,  Comparison o f t h e s e  specimens w i t h H, d u r i u s c u l u m have shown them t o be depauperate duriusculum  .  Ovate e l l i p t i c  America.  t o o r b i c u l a r l e a v e s a r e borne  forms o f appressed-  120 i m b r i c a t e on e r e c t branches a r i s i n g from fragmented, denuded  stems.  T h i s compares f a v o r a b l y w i t h s i m i l a r , though more r o b u s t specimens from f a r t h e r n o r t h .  The a l a r : c e l l s , though v e r y p o o r l y d i f f e r e n t i a t e d ,  are those o f Ii. d u r i u s c u l u m . Another A. J . Sharp specimen from M i c a Bay i n the Algoma d i s t r i c t , N o r t h e r n O n t a r i o i s a l s o remarkable.  I t s crowded, o r b i c u l a r ,  almost  p l a n e l e a v e s borne on s h o r t stems and branches bear a s t r i k i n g resemb l a n c e t o Hygrohypnum alpinum.  However, the presence o f w e l l  differen-  t i a t e d quadrate, short rectangular, t h i c k - w a l l e d a l a r c e l l s r e v e a l s i t to be H. d u r i u s c u l u m . Critical  study o f a p p r o x i m a t e l y 300 specimens r e v e a l s t h a t an o b l o n g -  e l l i p t i c t o o r b i c u l a r l e a f i s the u s u a l c a s e . between l e a v e s on the same p l a n t .  Still,  some v a r i a t i o n o c c u r s  I n b r o a d l y o v a t e l e a v e s the g r e a t e s t  width i s reached a t a p o i n t somewhere between the lower q u a r t e r and the lower t h i r d o f the l e a f .  As the l e a f shape approaches o r b i c u l a r the w i d e s t  p o i n t r i s e s t o the m i d d l e o f the l e a f .  The n o t a b l e v a r i a t i o n o c c u r s when  the w i d e s t p a r t r i s e s above the m i d d l e o f the l e a f t o produce a obovate c o n d i t i o n  ( F i g . 30 e ) .  slightly  Leaves o f such shape are not common.  Ovate  o r o v a t e - e l l i p t i c l e a f shapes do o c c u r r a r e l y i n p l a n t s t h a t a r e o t h e r w i s e depauperate. The shape o f the apex r e a d i l y v a r i e s from b r o a d l y a c u t e , o b t u s e , rounded o r o b t u s e o r rounded w i t h a s m a l l a p i c u l u s . V i r t u a l l y a l l p u b l i s h e d d e s c r i p t i o n s o f Hygrohypnum d u r i u s c u l u m make r e f e r e n c e t o the weak s e r r u l a t i o n o f the a p i c a l l e a f margin. 576 from B r y o l . e u r . was (1876).  c l e a r l y r e f e r r e d t o Hypnum alpinum by  Tab, I I I , Schimper  However, t h i s p l a t e has been f r e q u e n t l y and e r r o n e o u s l y r e f e r r e d  12-1 t o H. d u r i u s c u l u m .  F i g u r e 4 a o f Tab.  Ill  no doubt has been r e s p o n s i b l e  f o r the n o t i o n t h a t H. d u r i u s c u l u m has a s e r r u l a t e apex. than 200 European s e r r u l a t e margin  Study o f more  specimens o f v a l i d H. d u r i u s c u l u m has shown t h a t a i s of infrequent occurrence.  Q u i t e o f t e n , however, the  o u t e r w a l l o f the m a r g i n a l c e l l s seems p a r t i a l l y r e s o r b e d , thus g i v i n g the i l l u s i o n o f t e e t h . The a r e o l a t i o n o f Hygrohypnum d u r i u s c u l u m i s h i g h l y v a r i a b l e . l e n g t h s o f median l e a f c e l l s v a r y from 25 t o 100 um.  The  A few European  spec-  imens have l a r g e l e a v e s w i t h median l e a f c e l l s u n i f o r m l y i n the 60 t o 100 range.  um  There i s no c o r r e l a t i o n between l e a f s i z e and median l e a f c e l l l e n g t h .  Median l e a f c e l l s may  v a r y i n l e n g t h from one  s i d e o f the l e a f t o the o t h e r .  Zdenk P i l o u s e r e c t e d the name Hygrohypnum d i l a t a t u m f a b u l b o s a and t r i b u t e d two  dis-  specimens o f t h i s form as numbers 429 and 485 o f h i s musci  slovenici exsiccati.  cecho-  He based the form on the presence o f s m a l l b u l b s found  a t the a p i c i e s o f v a r i o u s stems and branches. e n t a g l e d , denuded stems.  The p l a n t s are l i t t l e more than  Examination o f the b u l b s has shown them t o be  clusters  o f c l o s e l y spaced, t i g h t l y i m b r i c a t e d l e a v e s t h a t have enveloped the stem o r branch:  apicies.  The b u l b s , l i k e the stems, a r e dark brown i n c o l o r .  The  b u l b s a r e p r o b a b l y the r e s u l t o f a p i c a l growth under s e v e r e environmental c o n d i t i o n s and, as such, a r e o f no taxonomic  value.  Kindberg d e s c r i b e d Hypnum pseudo-arcticum  i n Macoun (1890) and  dis-  t i n g u i s h e d i t from H. a r c t i c u m by i t s c r e n u l a t e l e a f margin and the s h o r t double c o s t a . acuminate  Grout  (1930), however, s t a t e d t h a t the p l a n t had a b r u p t l y  apex and t r e a t e d the p l a n t as synonymous w i t h Hygrohypnum l u r i d u m .  An examination o f the h o l o t y p e XS-PA) r e v e a l s t h a t Grout e v a l u a t e d an admixed fragment o f Hygrohypnum l u r i d u m w h i l e Kindberg, h i m s e l f , d e s c r i b e d a s m a l l fragment o f what i s r a t h e r t y p i c a l Hygrohypnum d u r i u s c u l u m from Western North  122 America. An examination o f the h o l o t y p e  (NICH) o f Hygrohypnum c o r d i f o l i u m  Okara. r e v e a l s t h a t i t i s i d e n t i c a l w i t h Hygrohypnum d u r i u s c u l u m Jamieson.  (De Not,)  The Okamura specimen agrees w i t h H, d u r i u s c u l u m i n terms o f  l e a f shape, a l a r d i f f e r e n t i a t i o n and o v e r a l l  appearance.  In 1869 De N o t a r i s d i s t r i b u t e d Limnobium d u r i u s c u l u m as the exsiccatum Erbar. Crittogam, I t a l . b e a r s a complete  s e r . I I , no. 204.  The l a b e l o f t h i s  exsiccatum  L a t i n d e s c r i p t i o n and a r e f e r e n c e t o Lago Maggiore.,  as t h e type l o c a l i t y .  Two specimens  (at G and UC) o f t h i s important  c c a t i have been a v a i l a b l e f o r study.  Italy exsi-  Index Muscorum, volume 5 (1969)  i n d i c a t e d t h a t t h e De N o t a r i s name i s a nomen nudum.  A r t i c l e 31 o f t h e  S e a t t l e e d i t i o n o f t h e Rules o f Nomenclature i m p l i e s t h a t names w i t h accomp a n y i n g d e s c r i p t i o n s borne on e x s i c c a t a l a b e l s b e f o r e 1 January 1953 a r e va published. The  Consequently,  Limnobium duriiusculum De Not, i s a v a l i d name.  s i g n i f i c a n c e o f t h i s name i s c r u c i a l f o r Limnobium d u r i u s c u l u m De Not.  i s u n q u e s t i o n a b l y t h e same organism  as Hypnum d i l a t a t u m Schimp.  Hypnum  d i l a t a t u m was d e s c r i b e d by Schimper  (1876) based on a m a n u s c r i p t  submitted  t o him by W i l l i a m W i l s o n . o  v  e  r  T h e r e f o r e , Limnobium d u r i u s c u l u m has p r i o r i t y  Hyp"" " d i l a t a t u m and beetaues t h e l e g a l basionym o f Hygrohypnum d u r i u -  sculum  1  (De Not.) Jamieson.  The specimen o f t h e De N o t a r i s e x s i c c a t u m a t  G i s d e s i g n a t e d as t h e l e c t o t y p e .  123 Fig.  29 a - h.  V a r i a t i o n i n the h a b i t o f l e a f y shoots o f Hygrohypnum  d u r i u s c u l u m and comparisons  between the wet and d r y c o n d i t i o n s i n  the s h o o t s . Each p a i r o f f i g u r e s , a - b , the comparison shoots. each  c - d , e - f , and g - h r e p r e s e n t s  between the wet and d r y c o n d i t i o n i n f o u r d i f f e r e n t  F i g u r e s a, c, e, and g i l l u s t r a t e the m o i s t c o n d i t i o n i n  ehoot.  Scale: Each shoot i s a p p r o x i m a t e l y 1 cm l o n g .  124  125 Pig.  30 a - h . V a r i a t i o n i n f o l i a g e l e a f shape, l e n g t h o f m a r g i n a l l e a f c e l l s and t h e h a b i t o f l e a f y shoots o f Hygrohypnum d u r i u s c u l u m . a - e. f h.  Foliage leaves  - g. Marginal leaf  cells  Leafy shoot  Scale: a - e.  I  f - g.  I  h.  1mm •  100um  t h i s shoot i s  | |  approximately 1 cm long  126  127 P i g . 31 a - h.  Variation  i n t h e apices  l e a v e s o f Hygrohypnum a - b.  -Leaf a p i c i e s  c, e - h. d.  Alar  cells  Median l e a f  cells  Scale:  ,  a - c , e - h. d.  \  duriusculum.  I  '  I 100um  lMum_J I  and c e l l u l a r d e t a i l  o f the  128  129  Fig.  32.  Hygrohypnum  duriusculum  130 E x s i c c a t i Examined A u s t i n , Musci A p p a l a c h i a n ! # 436 as JL m o l l e . (MICH, NY, USA) Grout, Hand-lens Mosses # 72. (MIN, NY) North American Musci P e r f e c t i # 62. (MICH, NY, TENN, UC, USA) # 62a (UBC) # 290 as Hy m o l l e . (MICH, MIN, NY, TENN, UC, USA) North American Musci P i e u r o c a r p i # 260 as H. m o l l e . (MIN, MO, NY, TENN, UC) # 442 as H. m o l l e . (COLO, MIN, MO, NY, TENN, USA) # 446 as H. m o l l e . (MIN, MO, NY, TENN, USA) North American Musci P l e u r o c a r p d Supplement # 59 as H. c o c h l e a r i f o l i u m . (MIN, TENN, UC) Macoun, Canadian Cryptogams # 794 as H. pseudo-arcticum. (TRTC) Canadian Mosses # 394 as H. a r c t i c u m . (USA) # 490, i n p a r t . (CANM, MICH, USA) Canadian Musci # 357. (CANM, NY, UC.) # 358 as H. a r c t i c u m . (CANM, MIN, NY i n p a r t ) S u l l i v a n t & Lesquereux, Musci B o r e a l i - A m e r i c a n i # 304. (MICH, NY, USA.) # 451 as H, p s e u d o - a r c t i c u m . (MICH, NY, UC., USA) Bauer Musci e u r o p a e i e t amer. e x s i c c a t i # 1789. (BRNM, G, MO, NY) Musci europaea e x s i c c a t i # 1278 a. iG, NY) # 1278b. (BRNM, G, NY) # 1279 as H. m o l l e . (BRNM, G, NY) # 1663 as H. m o l l e v a r . schimperianum. (BRNM, NY) # 1664 (BRNM, CANM~,G, NY) Brotherus Musci t u r k e s t a n i c i # 120. (G, H, NY) # 121. (G, S-PA) De N o t a r i s , E r b a r . C r i t t . I t a l . S e r . I I # 204 as L. d u r i u s c u l u m . (G, UC) L e c t o t y p e a t G • # 404. (G, UC) ~ . D i s m i e r , B r y o t h e c a G a l l i a e # 19. (NY) F a m i l e r , Ffcora e x s i c c a t a B a v a r i c a ; Bryophyta # 381B. (S-PA) Husnot, Musci G a l l i a e # 293 as H. m o l l e . (G, NY) # 495 as H, a l p e s t r e . (G, NY) # 692. (G) Kerner, F l o r a E x s i c c a t a A u s t r o - H u i j g a r i c a # 1922. (BP, BRNM, G, L E , MIN, r r _  MO, NY) L e v i e r , Bryotheca L e v i e r # 711. (S-PA) L i m p r i c h t , Bryotheca S i l e s i a n a # 90. (B, BP, L E , NY)  131 Lisowski, Bryotheca P o l o n i c a F a s c . IX. # 265 as H. alpinum. (LE, S-PA) F a s c . X, # 297. (BP, CAN, LE) F a s c . XI. # 319. (BP, CAN) F a s c . XXII. # 593 as H. p a l u s t r e . (CAN) F a s c . XXV. # 668. (BP, CAN) F a s c . XLIX. # 1266. (BP. LE) Mougeot, N e s t l e r & Schimper, S t i r p e s Criptogamae Kogeso-Rhenanae # 730 as H. m o l l e . (G, UBC) Museo H i s t . Natur. V i n d o b o n e n s i , Cryptogamae e x s i c c a t a e # 4260 as H. m o l l e . (NY, t h i s p a r t i c u l a r specimen i s an admixture o f H. m o l l e and H. d i l a t a t u m ) Noguchi & H a t t o r i , Musci J a p o n i c a S e r . 8. # 372 as H_. t s u r u g i z a n i c u m , (MICH, TENN, UC) P i l o u s , Musci c e c h o s l o v e n i c i e x s i c c a t i # 412. (G) #466. (G) # 487. (G) i # 798. (G) # 1177 as H. m o l l e . (C, G) Rabenhorst, B r y o t h e c a europaea # 899. (G, NY) W i l s o n , Musci B r i t a n n i c i # 384. (G, NY) Verdoorn, Musci S e l e c t i e t C r i t i c i S e r . XI, # 272. (MICH, MO, NY, TENN, UC) Z e t t e r s t e d t , Z e t t . Muse. p y r . # 251 as H. m o l l e . (CAN, NY) ? Krytogamae e x s i c c a t i # 400. (NY) S e l e c t e d Specimens Examined Canada B r i t i s h Columbia New.Denver, S l o c a n Lake; McFadden. ^UBC) Osoyoos Lake; Macoun 29 May 1895 as C. a r c t i c u m . (CAN) Macleod's Lake, 55°N; Macoun 2396 as H. s m i t h i i . (NY) Beaton, S e l k i r k Mtns., S a b l e Creek; Tusco O c t . 1962. (UBC) C o f f e e Creek, 4 m i l e s S. o f A i n s w o r t h ; Jamieson 20 Sept 1975 (UBC) Alberta L i t t l e S l a v e Lake; Mrs. Roy as H. s m i t h i i . (USA) Kananaska Co.; Macoun 3 Aug 1880. (TENN) Ontario Algoma d i s t r i e t , near M i c a Bay; Sharp CM-636. (TENN, UBC) Thunder Bay d i s t r i c t , Kakabeka F a l l s ; Macoun 14 J u l y 1869. (CANM, NY) A l g o n q u i n Park, Petawana R i v e r ; Macoun 24 J u l y 1900. (CAN) Ottawa d i s t r i c t ; Macoun 8 Aug 1905. (CAN) Lake Nipogon; Moser 1 J u l y 1884. Quebec Gaspe Co., Mt, A l b e r t ; A l l e n 30 J u l y 1881. (NY) S t . P a u l , Montmagmy; Gagnon & Masson 11206, (NY) Mont S t . H i l a i r e ; Dupret 7 Sept 1907. (CAN) Pare du Mont Tremblant, R i v i e r e La D i a b l e j Herman 16664. (CAN) Mont S h e f f o r d , c t e de Sheffond; Le B l a n c 2530 (NY)  132  Table Top Mtn.j A l l e n 10 Aug 1881. (NY) New Brunswick Grand F a l l s , F i s h hatchery, Feirane F a l l s ? Habeeb 355, (NY) Queens Co., Cannaan Forks; Macoun 1899. (CAN) A l b e r t Co., Fundy National Park, t r a i l to Third V a u l t F a l l s 5 Ireland 11470. (UBC) Nova S c o t i a Cape Breton I s . , Cape Dauphin; Nichols July-Aug 1914.(NY) Cape Breton I s . , Indian Brook; Nichols 9 Aug 1909. (NY) Labrador Caribou River; Wickes 25 J u l y 1938. (USA) C h u r c h i l l F a l l s , Bridge Camp area; Brassard 5330. (NFLD) Newfoundland Squires National Park, N. of Deer Lake, Humber R,; N o r r i s 4085, (HSC) Birchy Cove; Waghorne 21 Aug 1895. (MIN, NY) United States Alaska Kodiak Group, Raspberry I . Raspberry S t r a i g h t , Port V i t a ; Eyerdam 880. (TENNJ S - P A ) Washington C l a l l a m Co., Olympic National Park, OlymptceHotsprings; S v i l h a 793. (WTU) California Inyo Co., Inyo National Forest, along t r a i l on S. side of Lake Sabrina; Jamieson 5389. (UBC) Idaho" Idaho Co., Nez Perce National Forest, Red River D i s t r i c t ; Lems M60 as H. molle, (MICH) Nevada White Pine Co., Snake Mtns,, Nevada National Forest; Lawton 2810 as H. molle. (USA) Arizona Santa Cruz Co., Santa R i t a Mtns.; Bartram 156. (MY, USA) Apache Co., White Mtns., W, fork o f L i t t l e Colorado R.s Dybdall R 264. (Museum of N. Arizona) Montana G l a c i e r Co., G l a c i e r National Park, Reynolds Creek; Hermann 20433. (Personal herbarium o f F. J . Hermann) Missoula Co., Rattlesnake Campground; Sharp M-41. (TENN) B e l t Mountains; Williams 19 Sept 1891 as H. arcticum. (MIN) Wyoming Carbon Co., M i l l Creek; Porter 27 June 1934 as H. molle. i n North American Musci P e r f e c t i # 290. (TENN, USA, MIN, MO, NY, UC) Johnson Co., B u f f a l o Tensleep Rd., Porter 1638. (TENN) Park Co., E. of Beartooth Lake; Conard 21 Aug 1953. (UBC) Medicine Bow Mtns., E. o f Saratoga; Sharp 550. (TENN) Colorado Larimer Co., Rocky Mountain National Park, Larkspur Creek; Hermann 25989a. (Personal herbarium of F. J . Hermann) Mineral Co., F a l l s Creek, NE o f Pagosa Springs; Hermann 23343. (COLO)  133  q  La P l a t a Co., Above V a l l e c i t o ; Knowlton 85 as H. alpinum. (USA) Boulder Co., NW of Eldora, Middle Branch Creek; Hermann 24311. (WTU) Outlet of Corona Lake; Grout 20 J u l y 1914 as H. dilatatum i n North American Musci P l e u r o c a r p i # 446. (TENN, MIN, MO, NY* USA) New Mexico Rio A r r i b a Co., Brazos Canyon; Stanley & Bollman 10794. (NY, USA) Santa Fe Canyon; Arsene 20335, (USA) Utah Duchesne Co,, Uinta Mtns,, Ottoson Basin; Flowers 9141, (WTU) Minnesota Cook Co., Grand Marais; Holzinger 19 J u l y 1902 i n North American Musci P l e u r o c a r p i # 260. (NY, TENN, MIN, MO, UC) New York Greene Co., C a t s k i l l Mtns,, Haines F a l l s ; Hermannl4338 1/2, (MICH) Essex Co., S t , Huberts; Ketchledge 723. (UBC) Vermont Windam Co., S t r a t t o n ; Grout 28 June 1926 i n North American Musci P e r f e c t i 62, (TENN) Bennington Co., Manchester; Grout 28 June 1931 i n North American Musci P e r f e c t i 62. (UBC) New Hampshire Belknap Co., Mt. Belknap, G i l f o r d ; Carter 5 Sept 1904, (USA) Warren, Baker's River; Faxon 10 J u l y 1886. (NY) Mt, Washington, Great Gulf; Faxon 29 J u l y 1886. (NY) Maine Mt, Desert I s l a n d , Witches Hole; Patterson 208. (NY) Massachusetts Berkshire Co., P i t t s f i e l d , Lulu Brook; Rice 4 June 1942. (NY, as H. dilatatum; TENN, as H, molle) Connecticutt Ansonia; A l l e n 29 Oct 1880. (NY) Virginia Madison Co., Shenandoah National Park; Patterson 1195, (NY) Tennessee Sevier Co., Great Smokey Mountain National Park, Laurel F a l l s ; Sharp 15 Sept 1935 as•- H. c o c h l e a r i f o 1 ium i n North American Musci P l e u r o c a r p i Supplement 59. (TENN, MIN, UC) Sevier Co., Roaring Fork; Sharp 36211. (TENN) North C a r o l i n a Yancey Co., Blue Ridge Parkway, Crabtree F a l l s ; Jamieson 4875. (UBC) Jackson Co., Soco F a l l s near Junaluska Lake; Welch 2860.(TENN) Greenland Nugssuag Pen., Nugssuag; Holmen 15582- (LE, S-PA, UBC) , Disco F j o r d , Kuanit; P o r s i l d 969. (S-PA) Faroes Fugelefjord Bygd; Jensen 19 May 1896. (CAN, NY)  134  Great B r i t a i n Scotland Clova; Fergusson J u l y 1876. Schottland; Schimper 1868. (S-PA) Glen Coe, Rievers Glen, Binstead & Dixon 16 J u l y 1898. (NY) Wales North Wales, Aber; Hunt 16 May 1868. (S-PA) Norway Gausta; Jaederholm J u l y 1895. (S-PA) Opdal; Kaurin Aug. 1881, (S-PA) Lomsaggerej Bryhn J u l y 1879. (S-PA) Nordland, Salten, Nyman 28 J u l y 1893. (S-PA) Finmarken, Kaafjord; Z e t t e r s t e d t 13 J u l y 1868. (NY) Sweden Smaland, Hults Hesslas dam; Larsson 11 Sept. 1941. (S-PA) Bahusia, K r i s t e n d a l ; Arven 3 J u l y 1913. (S-PA) Upland, A t l y l n Sundstra; Lindberg June 1855. (S-PA) Vestmanland, Kingsor Runnakvarn; A h r l i n g 23 Aug 1880. (S-PA) Varmland, Ostmarks sn pa Ranneberget; Larsson 24 Aug, 1936. (S-PA) Dalarna, Norrbarki; Hakelier 4 July 1965. (S-PA) Halsingland, Bergsjo, Elfasen- C o l l i n d e r 3 J u l y 1877. (S-PA) Harjedalen, Lyingdaleri S t o r s j o ; Thedenius fiug 1842. (S-PA) Jamtland, B o r g a f j a l l t , Rankanlet, Frostvikens; Nyman 1 Aug. 1947. (S-PA) Angermanland, Sabra Klochersbacen; A r n e l l 26 Aug. 1874, (S-BA) Asele Lappmark, V i l h e i n i m i a ; M o l l e r 22 J u l y 1914. (S-PA) Lycksele Lappmark, Tama; Stenholm 12 J u l y 1924 . (S-PA) Lapponia L u l e n s i s , Sarjekensis, Kotikjokk; Jens;eh & A r n e l l 24 J u l y 1902, (S-PA) Tornea Lappmark, J u k k a s j a r v i , Abisko National Park; Persson & G j a e r e v o l l 15 Aug. 1944. (S-PA) Finland Lapponia pomojensis, Bakalda; Brotherus 18 J u l y 1872. (NY) {  S o v i e t Union.  S i b e r i a , A l t a i Mtns., Belokurikka; Bardunov 5 June 1966. , ( N I C H ) S i b e r i a , Akmolinsk. D i s t r . Atbalarky Ulu-tau; Ganeschin 1717, (H-BR) Bryotheca Caucasia, Ossetia; Brotherus 26 May 1887. (H-BR) S i b e r i a , Jenesei, A n t i s ferova; A r n e l l 27 June 1876. (S-PA) Turkestan, Thian Schan, in^ywalle f l . Narinkol.; Brotherus 27 J u l y 1896. (S-PA) Lapponia lmandra; A. Kohlman 24 J u l y 1887 (C, as Amblystegium molle v a r . alpinum) France Chamonix; Bernet J u l l i e t 1868, (G) Switzerland V a l a i s , Grand S t , Bernard; Jaquet 18 Aug. 1926. (S-PA) V a l a i s , Mattmark; Micholson & Dixon 13. (BM) Tessin, Lago Campoluigo; Conti Aout 1894. (G) Austria Steiermark, Schladming; B r e i d l e r Aug 1871, (G) T i r o l , I n n e r v i l l g a r t e n ; Ganders 15 Sept 1881. (BP) Salzburg, Lungau, Lessachthal; B r e i d l e r 16 June 1885. (BP) Vorarlberg, St. G a l l e n k i r s c h ; Hooch 26 Oct. 1928. ( s - P A ) Karnten, Tandelapha b e i Malta; B r e i d l e r 4 Aug. 1880. ( s - P A )  135. Italy  Prov. Comensis, V a l l e de Darengo; A r t a r i a 15 Aug. 1898. (S-PA, LE) Campello-Monte, Porv. Noraro Pedemonti, Bei Dannai; Levier 5 Aug. 1904 i n Bryotheca I t a l i c a 102. (S-PA) Macuguaga, V a l Anyasca; Nicholson & Dixon 10 J u l y 1913. (BM) Czechoslovakia Slovenia, Vysoke Tatry; Smarda 13 J u l y 1937. (Moravian) Moravia, Hruly Jesenik; Smarda J u l y 1947. (Moravian) Riesengebirges (= Krkonse), K l e i n e r Teich; S c h i f f n e r 13 June 1886, (S-PA) Poland Montes T a t r i O c c ; Lisowski 17 Aug 1956, i n Bryotheca Polonica Fasc. X. # 297. (BP) Roumania Carpat, merid., Montes Fogarasi; Vajada 13 Aug. 1966. (BP) Carpat. merid., Montes R e t y l z l a t ; Vajada 3 Aug. 1969. (BP) Bulgaria R i l a P l a n i n a , c i r c a Gam Koruje; Podpera 7 Aug. 1908. (S-PA) Borovec; Kovacs 29 Oct. 1956. (BP) Balkan, Trojanskapt, Katoferskap; Kuc 30 Aug 1960. (BP) V i t o s a planima, Stara R e l l a ; Podpera 17 J u l y 1908. (G) Asia Kungei A l t a u , Brotherus Aug. 1896 i n Musci Turkestani # 120. (H-BR, NY) Kashmir, Gulmarg? Duthie 3 June 1892. (H-^BR) China' Chensi (=Shensi) Prov., Sinn t s a i ; L i c e n t 209. (BM) Japan Mutsu PVW. Towada? Uemotsu 14 Aug, 1910. (H-BR) Shinano Prov., Mt., Shirouma; T i s h i b a 17 Aug. 1909. (H-BR) Kukuoka Prov., Kyushu, Seburi Mtns.; Amakawa 610. (NICH) - ! Nigano Co,, Okura, Minami-saku; Shimiju Aug. 1952 as H, tsurugzanicum i n Musdi Japonica # 372. (TENN) "~ Mt. Hokkoda; Uemtsu J u l y 1910. (NY) r  136 Hygrohypnum s m i t h i i (Sw. i n L i l j . ) Broth, Nat. Pf, 1(3):1039. 1909. Ledtotype: Sweden, Harjedalen, Funnasdalen? Thedenius.  (S-PA)  Lesikea s m i t h i i Sw. i n L i l j . , Svensk. F l . ed. 3:549. 1816. Hypnum arcticum Somm. i n Wahlenb., F l . Lapp. Suppl.  65.2.  1826.  Hypnum molle var. arcticum (Somm.) Aongstr. i n F r i e s . , Summ, Beg. Skand.  1:83.  1846.  Limnobium arcticum (Somm.) Schimp., B r y o l . Eur. 6:70, 578.  1853.  Stereodon a r c t i c u s (Somm.) M i t t , , Journ. Limn. Soc. 8:42. 1864. Hypnum s m i t h i i (Sw.) Lindb,, Act. Soc. Sc. Fennica 10:277. 1872, Amblystegium s m i t h i i (Sw.) Lindb., Musci Scand. 33.  1879.  Hypnum t o r r e n t i s C. M u e l l . e t Kindb. i n Macoun, Cat. Canad, P I . 6:243. 1892. C a l l i e r g o n arcticum (Somm.) Kindb., Canad. Rec. Sc. 6(2):72. 1894. Hygrohypnum arcticum (Somm.) Loesk., Verh, Bot. Ver. Brandenburg 46:198. 1905. Hygrohypnum crassinervium Warnst. ex Bauer. Hedwigia 54:152 - 153. f. 22.  1913.  Names o f taxa f o r which the appropriate l i t e r a t u r e was not a v a i l a b l e during t h i s study. Amblystegium s m i t h i i (Sw.) Lindb. i n Hartm., Handb. Skand. F l . ed, 10.  2:6.  1871.  Plants forming very s t i f f , coarse, l o o s e l y to t i g h t l y woven t u f t s o r mats.  Color v a r i a b l e , d u l l dark green, oMvaceous-green, blackish-green o r  black, o f t e n modified by golden-green o r golden-brown m o t t l i n g , e s p e c i a l l y i n the stem o r branch t i p s . at the t i p s .  Stems 1 to 8 cm long, p r o s t r a t e o r ascending  Branching v a r i a b l e , nearly unbranched o r frequently and i r r e g -  u l a r l y branched, branches  remaining p r o s t r a t e and a t t a i n i n g dimensions  s i m i l a r to the stems, or branches short and ascending to erect? o l d e r extremities of both stems and branches becoming denuded or clothed i n p e r s i s t e n t , but shredded l e a f bases.  remaining  Stems normally eradicu-  lose but those plants which bear many short, erect or ascending branches may be densely r a d i c u l o s e .  Stem cross-sectionarevealing 3 to 4 rows o f  small, t h i c k - w a l l e d , brownish c o r t i c a l c e l l s ? medullary c e l l s becoming l a r g e r and thinner walled toward the middle, becoming brown with age; cent r a l strand w e l l developed. Leaves variable? (0.6) 0.8 - 1,2  (1.5) mm long X (0.4) 0.6 - 1.2  (1,4)  mm wide? shape ovate, broadly ovate to o r b i c u l a r , sometimes transverse ovate; apex obtuse or rounded, sometimes almost acute or abruptly tapering to an obtuse point; margins e n t i r e or weakly d e n t i c u l a t e , plane throughout or s l i g h t l y recurved near the base; leaves plane or broadly and shallowly concave; costa v a r i a b l e ; u s u a l l y coarse, s i n g l e to 3/4, often v a r i o u s l y forked, o c c a s i o n a l l y short and double; leaves sometimes rather s t i f f l y decurrent the base.  at  When wet the leaves are s t i f f l y , but l o o s e l y imbricated to erect  spreading, upon drying the leaves shrink l a t e r a l l y s l i g h t l y and t w i s t . A r e o l a t i o n v a r i a b l e : Median l e a f c e l l s rhomboid, fusiform, b a c i l i f o r m or fusiform to l i n e a r flexuose;  22 - 48 (64) mm long X (4) 5 - 7 (10) mm wide;  a p i c a l c e l l s o v a l rhombic or rounded; Margins o f t e n , but not always, bordered by an i l l - d e f i n e d row of short c e l l s ; c e l l s generally becoming longer, wider and more incrassate toward the base, i n o l d e r leaves the basal c e l l s are often shiny yellow i n c o l o r , p i t s few to none; a l a r c e l l s s i m i l a r l y incrassate and yellow, v a r i o u s l y quadrate or rectangular, but not forming a c l e a r l y recognizable group. Plants monoicous.  Perigonia ovate, p e r i g o n i a l leaves ovate, to 0.6  mm  138 long,  concave  imbricate, ecostate, e n t i r e .  Outer and middle p e r i -  c h a e t i a l leaves ovate, ovate-lanceolate or sometimes s l i g h t l y elongatet t i a n g u i a r ? costa s i n g l e to midleaf, short and double or almost absent? margins e n t i r e , plane or recurved at the base; apex obtuse, v a r i o u s l y e n t i r e or d e n t i c u l a t e , l e a f lamina plane o r p l i c a t e .  Inner p e r i c h a e t i a l  leaves up to 5 mm long, u s u a l l y 2.5 to 4 mm long, costa s i n g l e and strong, 2-4  p l i c a e , margins e n t i r e and recurved, apex obtuse. Seta 8 - 17 mm long, s t r a i g h t or s l i g h t l y curved, red or reddish-  brown, smooth; Capsule t y p i c a l f o r the genus. Peristome as i n the genus? annulus of two rows of c e l l s ? endostome w i t h 1 to 3 poorly to w e l l developed c i l i a . to 21 um i n diameter, u s u a l l y 16 to 19  Spores yellow to brown, 13  um.  Hygrohypnum s m i t h i i may be recognized by i t s coarse, r i g i d habit and the u s u a l l y broadly ovate to o r b i c u l a r , l o o s e l y imbricated to spreading leaves and the stout, g e n e r a l l y s i n g l e costa (Fig. 33 a - e ) .  Of these  features, the r i g i d h a b i t i s constant, while the others may vary. of t h i s v a r i a t i o n , these characters i n combination  In s p i t e  are d i a g n o s t i c f o r the  species. A number of features held by e a r l i e r authors as u s e f u l i n recognizing the species have been shown to be u n r e l i a b l e by the present  study.  Husnot (1894) and Lawton (1971) r e s p e c t i v e l y described the stem as ranging from 3 - 5 cm and 1 - 4 cm long.  The present studies have revealed  continuous v a r i a t i o n i n stem length from 1 - 8  cm.  North American p l a n t s  are o f t e n shorter than European ones, but the d i f f e r e n c e s do not appear significant. Schimper (1853, 1860, 1876), Braithwaite 11898), Dixon (1896, 1924), Grout (1931) and Lawton (1971) commented upon the slender nature of the  139 species.  I f the thickness of a stem i s c o r r e c t l y i n t e r p r e t e d here to  mean the distance between the t i p of one l e a f to that of another l e a f on the opposite side of a moist stem, then the feature i s too v a r i a b l e and too subjective a feature to be u s e f u l . The leaves of Hygrohypnum s m i t h i i are g e n e r a l l y described as broadly ovate to o r b i c u l a r .  I t i s not unusual to observe leaves that may be ovate,  oblong ovate or almost t r i a n g u l a r (Pig. 33 c, d, & e ) . are o f t e n described as being 1 mm or l e s s i n length. leaves a t t a i n i n g lengths of 1.5 mm are not unusual. have born leaves up to 2.0 mm  Further, the leaves In natural populations,  Plants grown i n c u l t u r e  long.  The margin i n the upper h a l f of the l e a f i s frequently described having an i l l - d e f i n e d border of short c e l l s ( F i g . 34 c ) .  as  Such i s not always  the case as can be seen i n F i g . 34 e. The median l e a f c e l l s are generally q u i t e uniform.  Occasionally the  median a r e o l a t i o n e x h i b i t s very wide c e l l s interspersed among c e l l s of more t y p i c a l dimensions ( F i g . 34 f ) . The a l a r c e l l s are u n d i f f e r e n t i a t e d (Fig. 34 g) o r they form a small, i l l - d e f i n e d girottp of small, quadrate, incrassate c e l l s ( F i g . 34 h). Occasionally Hygrohypnum s m i t h i i has been confused with H, b e s t i i and M* dilatatum.  I t i s s i g n i f i c a n t that a l l three are t y p i c a l l y very coarse  to the touch.  Hygrohypnum s m i t h i i i s most r e a d i l y d i s t i n g u i s h e d from H.  b e s t i i by the marginal l e a f c e l l s .  Compare F i g . 34 c & e with F i g . 37 e & f.  In Western North America very small specimens of H. s m i t h i i and H. duriusculum may be confused.  Both species may  assume a h a b i t i n which p r o s t r a t e stems  bear numerous short, ascending to erect branches. best on the basis of t h e i r costae.  They may be d i f f e r e n t i a t e d  Further, H.- duriusculum loses considerable  of i t s r i d l d i t y upon moistening, whereas H_. s m i t h i i does not.  The median l e a f  c e l l s of H. s m i t h i i are u s u a l l y 22 to 48 um long, whereas those of H, d u r i u -  140  sculum are u s u a l l y 45 to 70 um long.  The a l a r c e l l s o f H. duriusculum  t y p i c a l l y e x h i b i t more conspicuous d i f f e r e n t i a t i o n than those o f H. smithii. Personal f i e l d experience with Hygrohypnum s m i t h i i has been confined to Southern B r i t i s h Columbia.  I n t h i s area the species has been observed  under a v a r i e t y o f conditions.  In Kokanee Creek and a t Whistler Mtn.  H. s m i t h i i was observed growing on i r r i g a t e d rocks. at elevations i n excess o f 4000 feet.  At Stagleap Pass H. s m i t h i i was  c o l l e c t e d from a very sluggish and seepy stream. encased by dark, humic s o i l .  Both l o c a l i t i e s were  The plants were h e a v i l y  This feature i s most unusual, f o r species of  Hygrohypnum normally occur i n clean water.  A t Kokanee Lake, H, s m i t h i i was  observed i n another equally unusual s i t u a t i o n .  Only very r a r e l y have I seen  any species of Hygrohypnum growing completely submerged.  At the o u t l e t o f  Kokanee Lake H. s m i t h i i was seen growing i n extensive mats on large boulders whose tops were j u s t a few inches below the lake surface.  The mats of H.  s m i t h i i covered a surface area o f approximately 30 square f e e t . Hygrohypnum s m i t h i i was o r i g i n a l l y described as Leskea s m i t h i i by Swartz i n L i l j e b l a d ' s (1816) Svensk F l o r a .  Schimper (1853) described Limnobium  arcticum and based the taxon on Hypnum arcticum Sommerfeld, which was i t s e l f described i n Wahlenberg  (1826) .  In l i t e r a t u r e not seen LMdberg (Lindberg i n  Hartman, 1871) evidently recognized the c o n s p e c i f i c i t y o f the Swartz and Sommerfeldtitaxa and the p r i o r i t y o f the Swartz name f o r he recognized Amblystegium s m i t h i i (Sw.) Lindb. Although Lindberg's (1872, 1879) opinion of the generic p o s i t i o n o f the p l a n t v a r i e d from that o f 1871 i t i s c l e a r that he s t i l l held the Swartz and Sommerfeldt taxa as c o n s p e c i f i c .  Neither  Swartz nor Sommerfeldt designated any specimens, although they s p e c i f i e d geographical areas.  Swartz i n d i c a t e d Norrige and Sommerfeldt designated S a l t d a l e n ,  141 Nordlandiae.  Schimper (1853) c i t e d several specimens, o f which the  f o l l o w i n g are p e r t i n e n t :  S a l t d a l e n , Nordlandiae, Sommerfeldt; C h r i s t i a n a ,  Norvige, B l y t t , Schimper; Harjedalen prope Funnasdalen, Thedenius; K l e i n e r Teich, Sendtner. Apparent duplicates of the B l y t t , Thedenius and Sendtner specimens are a t S-PA.  In the Schimper herbarium at BM i s a specimen from  C h r i s t i a n a , Norvige f o r which a c o l l e c t o r i s not designated.  I t may w e l l  be Schimper's own c o l l e c t i o n ; t h i s cannot be c o n c l u s i v e l y proven.  These  specimens are p e r t i n e n t f o r they seem to e s t a b l i s h the c o n s p e c i f i c i t y of the Sommerfeldt and Schimper concepts of Hypnum arcticum and Limnobium a r c ticum.  Then, i f one can assume that Lindberg (1871, i n Hartmann) accurately  recognized the c o n s p e c i f i c i t y o f Leskea s m i t h i i Sw. and Limnobium arcticum Schimp,, then a neotype can be selected from the specimens c i t e d by Schimper, With t h i s r a t i o n a l e a Thedenius specimen at S-PA i s designated as the l e c t o type f o r Hygrohypnum s m i t h i i (Sw. i n L i l j . ) Broth.  142 P i g . 33 a - h. of  V a r i a t i o n i n l e a f shape and t h e h a b i t o f l e a f y shoots  Hygrohypnum  smithii.  a - e.  Foliage  leaves.  f - h.  L e a f y shoots i n the m o i s t c o n d i t i o n .  Scales a - e. f - h.  ;  I  1mm  I  shoots are approximately 0.75  cm long  143  144 P i g . 34 a - h.  C e l l u l a r d e t a i l s o f the p e r i c h a e t i a l and f o l i a g e leaves  of Hygrohypnum s m i t h i i . a.  F o l i a g e l e a f apex.  b.  P e r i c h a e t i a l l e a f apex.  c and e.  Marginal c e l l s o f f o l i a g e leaves  d and f.  Median l e a f c e l l s o f f o l i a g e leaves  g - h. A l a r c e l l s o f f o l i a g e leaves  Scale: a,  c - f.  b, g - h.  I  I  100um  100um I  I  145  146  4-  F i g . 3 5 . Hygrohypnum s m i t h i i  147 E x s i c c a t i Examined Bauer, Musci europaei e t americani e x s i c c a t i # 2289. (BRMN) De N o t a r i s , Erbar, C r i t t . I t a l , # 306 as H. arcticum. (NY, UBC) L e v i e r , Bryotheca I t a l i c a # 512 as L. molle. (S-PA) Limpricht, Bryotheca S i l e s i a c a # 292 as Hypnum arcticum. (BP, LE) L i s o w s k i , Bryotheca Pblonica Fasc. X # 295. (BP, LE) Fasc. XXIV # 639. (BP, LE) Macoun, Canadian Musci # 358 as arcticum. (NY, TENN, UC) # 503 as H. g o u l a r d i i . (MICH, MIN, TRTC, USA) Rabenhorst, Bfeyotheca europaea # 1142 as L. arcticum. (LE) S u l l i v a h t and Lesquereux, Musci B o r e a l i Americani # 194 as H. molle. (MICH, NY, TENN) Selected Specimens Examined Canada B r i t i s h Columbia Vancouver I s l a n d , Strathcona P r o v i n c i a l Park, Golden Hind? Boas 356. (UBC) Bulkley Range; Boas 714. (UBC) Lake of the Hanging G l a c i e r s ; MacFadden 1096. (UBC) Skeena R., south o f Harrison L., S c h o f i e l d and Boas 21269. (UBC) Haines Highway, 54 mi. NW of Haines? Hermann 21794. (CAN, MICH, NY, USA) Kokanee G l a c i e r P r o v i n c i a l Park, Kokanee L.? Jamieson 5614. (UBC) G a r i b a l d i P r o v i n c i a l Park, w h i s t l e r Mtn,? S c h o f i e l d and Jamieson. (UBC) Alberta Jasper National Park, Tonquin Valley? MacFadden 27 J u l y 1926. (UBC) Quebec Gaspe, S t , Anne R,; Macoun 29 Aug. 1890. (CAN) A n t i c o s t i I . , B e c e c i i R., Macoun Aug. 1890. (CAN) Nova S c o t i a Cape Breton I . , V a l l e y of Barrasois; N i c h o l s 1948. (NY) Newfoundland Humber R. D i s t r i c t , E. of Blow-Me-Down- Mtn., Williams 1601. (CAMN$ HSC, MICH) Sops Arm, Approx. 5 mi. S., N o r r i s 4198. (HSC) Trout R i v e r , approx. 4 mi. E. N o r r i s 4655. (HSC) United States Alaska Prince W i l l i a m Sound, Krii/fcrfat I . , Thrum Bay; Eyerdam 589 (MICH) Washington Olympic National Park, Solduc Hotsprings, Dear Lake T r a i l ? S c h o f i e l d , Ireland s Boas 19473. (UBC) Snohomish Co., Monte C s i s t o , T s a i l to G l a c i e r Basin? S c h o f i e l d 20123. (UBC)  148  California Mono Co. Harvey Monroe H a l l Natural Area, Cabin Creek; Catchenda 4788. (CAN, MICH, TENN) T r i n i t y Co., i n r i v u l e t below E. Weaver L,; N o r r i s 9415 (HSC) Montana Flathead Co., G l a c i e r National Park, Hidden Lake T r a i l S. of Logan Pass; Hermann 18315, (CAN, USA) L i n c o l n Co., Cabinet Mtns., Leigh Lake T r a i l ; S c h o f i e l d 11978. (CAN, HSC, NY) Colorado Boulder Co., NW of Eldora, Diamond Lake T r a i l ; Hermann 26660. (Herbarium o f F. J . Hermann, Dupl. - UBC) Greenland Nigerdleg; Holmen 63-146. (NY, UBC) Ilimaussag Peninsula, Dyrnaes; Holmen & Steere 62-584. (NY, UBC) T u n a g d l i a r f i k F j o r d , Narassarssuag; Holmen & Steere 62-607. (NY, UBC) Iceland N. W. Iceland, Strandasyala, Reyhyavik; Johannson 3 Sept 1967. (UBC) Faroes Bordo, Graverdal, Jansen 17 May 1896. (NY, UAC, UBC) Sweden Harjedalen, Tannas, Funnasdalen; Thedenius 1842, (S-PA, Lectotype) Jamtland, Frostviken; Hulphers 1 Sept 1934. (S-PA) Asele Lappmark, V i l h e l m i n a ; M o l l e r 20 J u l y 1914. (S-PA) Lycksele Lappmark, Tama; Angstrom. (S-PA) Lapponia Tornensis, Mellan, J i k k a s j a r v i ; Jaderhoto 29 J u l y 1914, (S-PA) Norway C h r i s t i a n i a , Bogstadaas; Z e t t e r s t e d t & Wickbom 16 June 1870, (S-PA) Kongsvold, Dovre; Kindberg 1885. (NY, S-PA) Nordland, Helgeland, Lerskaradalen S t o p e f j e l d ; B l y t t & A r n e l l 20 Aug. 1870. (S-PA) Finnmarken, Hammersfest; Jorgensen 15 Oct. 1888. (S-PA) Finland Lapponia imandrae, Urupjok, Haarakoski; Kihlman 3 J u l y 1892, (S-PA) Lps. S a l m i j a r v i , K u o t s j a r v i ; Roivainen J u l y 1927. (NY) S o v i e t Union  Peninsula Kolaensis, Montes Chiciny; Schljakov 20 Aug. 1947 as Hep a t i c a e e t Musci URSS E x s i c c a t i 9 49. (CAN) Austria T i r o l F e r w e l l ; Lorentz Sept 1866. (BP) Steiermark, Sturzbach am Ostablang des K n a l l s t e i n i n der Solk; B r e i d l e r 4 Aug. 1877. (BP) Vorarlberg, Gampadelthal i n der Rhatikonkette b e i Schruns; B r e i d l e r 19 J u l y 1882. (BP) Karnteri, " K l e i n Eland" im M a l t a t h a l ; B r e i d l e r 25 Aug. 1880. (BE) Switzerland Bern Canton, Oberlager am Faulhorn; Culmann 10 Aug. 1908. as Musci europaei e x s i c c a t i # 650. (S-PA) Scotland Aberdeen, Head of Glen C a l l a t e r ; C r o a l l 11 Aug 1853. (CAN)  149 Ben Lawers; Gardner 1838. (CAN) Ben Mhur; Schimper 1869. (BP, S-PA) Canlochan Glen, F o r f a r ; Hunt 13 J u l y 1868. (BP) Clova; ex dons E, G. B r i t t o n 1868 ?? (NY) France Pyrenees c e n t r a l e s , Crabioules e t Tousse de Maupas; Z e t t e r stedt 18 Sept 1856 as Zett. Musci pyr, # 252. (CAN, S-PA) Italy Monte Adamello, Schneebach an der Mondronhutte; Kern 27 Aug 1901. (BP) Poland Montes T a t r i A l t i ; Lisowski 29 Aug. 1956 as Bryoth, P o l . Fasc. XXIV # 639. (CAMN) C zecko slovak i a K l e i n e r Teich; Herausgeber 6 Aug 1868 as Bryo. S i l e s . # 292, (BP, NY)  150 Hygrohypnum b e s t i i (Ren. et Bryhn) Holz., B r y o l o g i s t 4:12,  22.  1901.  Lectotype; Montana, v i c i n i t y of Lake McDonald, Avalanche Basin. Leg: J . M. Holzinger and J . B. Blake 29 J u l y 1898.  (NY)  Hypnum molle ssp. b e s t i i Ren. et Bryhn, B u l l . d. L'Acad. I n t . d. Geog. Bot. 10:7.  1901.  Hypnum b e s t i i .(Ren, et Bryhn) Ren. 8.  et Bryhn, Rev. B r y o l . 28:  1901.  Limnobium b e s t i i (Ren. et Bryhn) Holz., B r y o l . 4:22.  1901.  Hygrohypnum molle ssp. b e s t i i (Ren. et Bryhn) Broth., Nat. P f l . Ed. 1, 2:1040.  1909.  Hygrohypnum molle var. b e s t i i (Ren. et Bryhn) Hab., 54:156.  Rhodora  1952.  P l a n t s very s t i f f and coarse, r a r e l y s o f t , i n l o o s e l y woven mats or tufts.  Color u s u a l l y d i r t y olive-green, often d i r t y - y e l l o w , often d u l l  yellowish-green  with golden m o t t l i n g , e s p e c i a l l y near stem and branch  apices, r a r e l y bright-green.  Stems(1) 3 - 7  (12) cm long, denuded i n o l d e r  extremities i n s t i f f forms, f o l i o s e throughout i n some s o f t forms; r i g i d l e a f bases and decurrent wings c h a r a c t e r i s t i c a l l y p e r s i s t e n t on the otherwise denuded p o s t e r i o r extremities of stems i n s t i f f , coarse forms.  Stem  cross-sections r e v e a l i n g 2 to 4 rows of small, t h i c k - w a l l e d c o r t i c a l c e l l s , yellow-brown to reddish-brown i n c o l o r ; medullary c e l l s l a r g e r , u s u a l l y thinner-walled, o c c a s i o n a l l y becoming incrassate with age i n s t i f f , r i g i d stems; c e n t r a l strand absent or very weak.  Branching i r r e g u l a r ; i n very  s t i f f forms branches ascending or erect from p r o s t r a t e , interwoven, denuded stems; i n s o f t e r forms l e a f y stems and branches intertwine i n l o o s e l y woven tufts.  Eradiculose.  151*  Leaves v a r i a b l e .  Somewhat d i s t a n t l y spaced.  A t t i t u d e on the stem  v a r i a b l e from the wet t o dry c o n d i t i o n ; when wet leaves weakly appressed, spreading o r e r e c t , sometimes s l i g h t l y secund; v a r i o u s l y contorted when dry, o f t e n t w i s t i n g t o the l e f t i n the upper h a l f .  Leaves (1) 1.5 - 2.5  (3) mm long X 1 - 1.5 (2.0) mm wide; shape ovate t o broadly ovate, sometimes s l i g h t l y asymmetrical; margins e n t i r e , becoming somewhat uneven i n the apex, r a r e l y weakly d e n t i c u l a t e ; apex obtuse o r somewhat acute; leaves plane t o broadly and shallowly concave; s t r o n g l y decurrent t o scarecely so, leaves o f t e n removed with d i f f i c u l t y from the stem because o f the strong bond between the i n c r a s s a t e basal l e a f c e l l s and the i n c r a s s a t e c o r t i c a l stem c e l l s ; costa v a r i a b l e , u s u a l l y strong and double from a massive base, one o r both branches reaching midleaf o r beyond, o r very strong and s i n g l e reaching w e l l beyond midleaf, o f t e n bearing $., 2 o r r a r e l y 3 l a t e r a l f o r k s , l e s s commonly weak and double, e s p e c i a l l y i n very young leaves and small branches, costa yellow-green o r yellow, becoming brownish w i t h age. A r e o l a t i o n v a r i a b l e ; median l e a f c e l l s l i n e a r flexuose to broadly rhombic flexuose, (40) 60 - 120 (170) am long X (2) 6 - 10 (12) um wide, median l e a f c e l l s may e x h i b i t a fa±£iy uniform width (10 - 12 um) may be i r r e g u l a r l y  o r very wide c e l l s  interspersed among much narrower c e l l s , such  v a r i a t i o n o c c u r r i n g between leaves on the same stem and from one side t o the other i n i n d i v i d u a l leaves; a p i c a l c e l l s u s u a l l y rounded quadrate o r rhombic, although l i n e a r flexuose c e l l s up t o 100 um long are not uncommon; marginal l e a f c e l l s between the lower 1/3 and the upper 1/4 of the l e a f vary from 60 t o 250 um long; t r a n s i t i o n from median t o basal c e l l s v a r i a b l e , l i t t l e changed o r not at a l l , becoming t h i c k e r o r thinner-walled, mostly becoming s l i g h t l y wider, b a s a l c e l l s u s u a l l y s t r o n g l y i n c r a s s a t e and y e l l o w i s h , t u r n ing yellow-brown with age, the pigmentation o f t e n c r e a t i n g a r a d i a t i n g sun-  152 burst e f f e c t , or sometimes l i t t l e o r not a t a l l incrassate? P i t s few t o none; A l a r c e l l s v a r i a b l e , hardly d i f f e r e n t i a t e d t o an imprecisely d i f f e r e n t i a t e d group o f quadrate t o short rectangular, moderately i n c r a s s a t e cells. Plants apparently d i o i c o u s , male and female plants similar.- P e r i g o n i a borne i n the a x i l s o f stem leaves and l a r g e branch leaves, inner p e r i g o n i a l leaves ovate, 0.75 to 0.9 mm long, deeply concave, margins a t Ifehe apex s l i g h t l y i r r e g u l a r to weakly d e n t i c u l a t e , ecostatej Outer and middle p e r i c h a e t i a l leaves ovate to ovate-lanceolate, ecostate, r e f l e x e d squarrose i n the upper h a l f ,  inner p e r i c h a e t i a l leaves long l i n e a r l a n c e o l a t e , up t o 2 mm,,  long, costa v a r i a b l e , short and double, s i n g l e t o midleaf o r absent, leaves m u l t i s t r a t o s e through the middle w i t h b i s t r a t o s e blades, a p i c a l margin uneven w  with 1 t o 4 coarse teeth.  Seta 15 t o 25 mm long u s u a l l y 18 t o 22 mm; red to dark maroon-red i n c o l o r ; smooth, s t r a i g h t when wet, v a r i o u s l y twisted when dry.  Capsule as  i n the genus, Annulus of 2 t o 3 o r 4 rows o f deciduous c e l l s . Peristome as i n the genus, Endostome with 1 t o 3 f i n e l y p a p i l l o s e , weakly appendiculate c i l i a between each tooth; spores pale yellow, f i n e l y p a p i l l o s e , 13 t o 21 um i n diameter, u s u a l l y 17 to 19 um. Of the broad-leafed species Hygrohypnum b e s t i i i s best recognized by 1.  the very long marginal l e a f c e l l s , which range from 60 to 250 um, 2.  the l a r g e leaves which reach 3 mm long X 2 mm wide, and 3. sexuality.  the dioicous  In many cases d i s c o l o r a t i o n i n the basal l e a f c e l l s imparts the  appearance o f a r a d i a t i n g '^sunburst" i n the l e a f base which i s t y p i c a l o f the species. Hygrohypnum b e s t i i i s a very d i s t i n c t i v e species.  However, c e r t a i n o f  153 i t s c h a r a c t e r i s t i c s are s u f f i c i e n t l y v a r i a b l e to overlap with other species o r are simply shared with other species. The broadly ovate l e a f shape which i s t y p i c a l o f H. b e s t i i ( F i g . 38 a, b ) , i s a l s o shared by H_. molle ( F i g . 44 a - e) . The leaves o f H. b e s t i i are u s u a l l y 1.5 t o 2,5 mm long and as such are much l a r g e r than the t y p i c a l lengths o f 0.9 to 1.3 mm a t t a i n e d by the leaves of H. molle i n Western North America.  However, small p l a n t s o f H. b e s t i i w i t h t i n y leaves are not i n f r e -  quent and may e a s i l y be confused with H_. molle.  The i d e n t i t y of such plants  can be e s t a b l i s h e d only by noting o f the very long marginal l e a f c e l l s i n H_. bestii. Both Hygrohypnum b e s t i i and H_. duriusculum are described as coarse to the touch o r r i g i d and s t i f f when dry.  Further, the leaves o f H, b e s t i i may  approach an o r b i c u l a r shape that i s more t y p i c a l o f H. duriusculum  ( F i g . 37 c ) .  Again, these two species may be r e a d i l y d i s t i n g u i s h e d by t h e i r marginal l e a f c e l l s , where those o f H_. b e s t i i are by f a r the longer (Compare F i g . 38 e & f w i t h F i g . 30 f & g ) . As ijtust discussed, H. b e s t i i i s g e n e r a l l y very coarse o r s t i f f to the touch when dry.  C e r t a i n specimens may be r i g i d even when wet. However, i t  i s not unusual to f i n d specimens that are s o f t and f l e x i b l e i n both the wet and dry s t a t e .  Therefore, t h i s feature should be employed i n the f i e l d with  some d i s c r e t i o n . One unique feature o f H. b e s t i i , which seems to be o f uncertain taxonomic value has t o do w i t h the d i f f i c u l t y with which leaves may be removed from the stem during d i s s e c t i o n .  The bond a t the p o i n t of i n s e r t i o n between the l e a f  and the stem i s exceedingly tenacious and i s r e f l e c t e d i n the shredded and p e r s i s t e n t l e a f bases on the o l d e r e x t r e m i t i e s o f both stems and branches. The leaves w i l l i n v a r i a b l y shred during d i s s e c t i o n .  A good deal o f c o r t i c a l  154 stem t i s s u e remains attached t o the leaves.when they are removed from the stem. The a t t i t u d e o f the leaves upon the stems may vary s l i g h t l y between and w i t h i n the wet and dry c o n d i t i o n s . When moist the leaves are g e n e r a l l y rather d i s t a n t and l o o s e l y imbricated t o s l i g h t l y spreading.  In some  p l a n t s , however, the leaves may be gently folded around the stem so as t o be s l i g h t l y secund ( F i g . 36 c ) . Upon d r y i n g the leaves normally e x h i b i t some l a t e r a l shrinkage and concomitant t w i s t i n g .  Sometimes the leaves  may be very s l i g h t l y f a l c a t e , but normally the leaves are q u i t e s t r a i g h t . The a l a r c e l l s i n H. b e s t i i vary from v i r t u a l l y u n d i f f e r e n t i a t e d (Fig. 3 8 h) t o an i l l - d e f i n e d group o f short rectangular c e l l s ( F i g . 3 8 g ) . 1  Such v a r i a t i o n can lead to confusion w i t h e i t h e r H. molle o r H_. duriusculum. The very long marginal l e a f c e l l s i n H_, b e s t i i w i l l c l e a r l y d i s t i n g u i s h i t from these two species. The median l e a f c e l l s a l s o e x h i b i t some rather p e c u l i a r v a r i a t i o n . most cases the median a r e o l a t i o n i s uniform. wide c e l l s may be i r r e g u l a r l y  In  However, i n some p l a n t s very  interspersed among c e l l s o f more t y p i c a l widths;  Curiously a s i m i l a r s i t u a t i o n occurs i n H. s m i t h i i , which i s i t s e l f a rather coarse p l a n t .  In leaves of H_. s m i t h i i there i s l a t e r a l shrinkage and l o n g i -  t u d i n a l t w i s t i n g that i s reminescent o f H_. b e s t i i .  This i s more a function of  the r i g i d i t y o f the p l a n t s than a fundamental r e l a t i o n s h i p . Hygrohypnum b e s t i i was described o r i g i n a l l y from s t e r i l e m a t e r i a l . Due to the confusion o f H. b e s t i i with H. duriusculum and H. molle the dioicous nature and the existence of sporophytes i n H. b e s t i i remained Unclear u n t i l Lawton's (1966) r e p o r t .  Of 128 specimens examined i n d e t a i l 11 were female,  4 were male and 8 were female and bore sporophytes,  Lawton (1966) pointed  out that the m u l t i s t r a t o s e inner p e r i c h a e t i a l leaves contrasted the u n i s t r a t -  155 ose c o n d i t i o n i n H. molle. The p a p i l l o s e a p i c a l c e l l s i n the p e r i c h a e t i a l leaves of H. molle d i f f e r from the smooth c e l l s i n those o f JI. b e s t i i , Hygrohypnum b e s t i i has one unusual form that occurs at f i v e widely spaced l o c a l i t i e s along the P a c i f i c Coast of North America.  The p l a n t  s u p e r f i c i a l l y resembles H. ochraceum, but d i f f e r s i n the absence o f the i n f l a t e d , hyaline c o r t i c a l c e l l s .  L i k e t y p i c a l H, b e s t i i the leaves u s u a l l y  exceed 2 mm long, but d i f f e r through a more oblong-lanceolate l e a f shape (Fig. 37 f - i ) . and forked. bestii.  The costa i s stout and v a r i o u s l y double and/or s i n g l e  The marginal l e a f c e l l s are very long as i s t y p i c a l f o r H,  The form a l s o d i f f e r s i n i t s complete lack o f a l a r d i f f e r e n t i a t i o n  (Fig. 38 i ) and a tendency f o r more regular pinnate branching. This form seems to be a lowland p l a n t occurring from sea l e v e l to about 2000 f t , whereas H. b e s t i i i s from much higher e l e v a t i o n s . may i n d i c a t e one of two s i t u a t i o n s .  The s t e r i l i t y o f t h i s form  One, that the p l a n t i s dioicous, or two  that the f i v e known populations represent s t e r i l e o u t l i e r s o f an otherwise f e r t i l e mountain species that cannot reproduce a t low e l e v a t i o n s .  That H,  b e s t i i i s dioicous i s no doubt s i g n i f i c a n t . Hygrohypnum b e s t i i has long been viewed as a Western North American endemic (Lawton, 1971; S c h o f i e l d , 1969). ;  That Habeeb (1952) reported the  species from A l b e r t and V i c t o r i a counties New Brunswick seems to have gone e n t i r e l y unnoticed. The present study has discovered other l o c a l i t i e s f o r H. b e s t i i i n Eastern North America, extending from the upper peninsula of Michigan to the Gaspe Peninsula, Quebec and Cape Breton Highlands, Nova Scotia. In Index Muscorum V o l . 5 Wijk e t a l . (1969) reported H. b e s t i i from Europe.  I t i s p o s s i b l e that the species might be discovered i n Scandanavia,  but as yet I have seen no m a t e r i a l to support t h i s a s s e r t i o n .  156 Although H. b e s t i i i s common throughout Western North America, my own f i e l d experience has been l i m i t e d to f i v e l o c a l i t i e s i n southern B r i t i s h Columbia and three i n northern C a l i f o r n i a . 3* b e s t i i has been observed i n a v a r i e t y of h a b i t a t s .  In these areas The e l e v a t i o n a l  range, as I know i t , extends from sea l e v e l to 6500 f e e t .  In .'some cases  the species grown on submerged rocks i n c l e a r , cold mountain streams.  In  other cases, i t grows on rocks i n small, s i l t - : a i l d sand-ladened, apparently seasonal r i v u l e t s .  In such r i v u l e t s the water tumbles over the rocks i n  small cascades or slowly seeps over them.  Frequently considerable humic  s i l t i s present. Organic or inorganic s i l t and sand accumulate i n the p l a n t s under such c o n d i t i o n s .  At sea l e v e l the form resembling H_, ochraceum  has been observed growing on sandstone i n a seepy roadside d i t c h .  I t has  a l s o been seen i n a c l e a r , s i l t - f r e e stream a t an e l e v a t i o n o f about 300 feet.  I have been unable to draw any c o r r e l a t i o n between the coarseness  or softness of H_. b e s t i i and v a r i a t i o n i n h a b i t a t . Renauld (1901) described Hypnum (Limnobium) molle Dicks, s s p . b e s t i i Ren .et Bryhn based on a J . M. Holzinger c o l l e c t i o n of 29 J u l y 1898 from Avalanche Basin, Montana at an e l e v a t i o n of 1500 m.  Renauld (1901a) pub-  l i s h e d a very s i m i l a r and somewhat redundant paper r e d e s c r i b i n g the p l a n t as Hypnum (Limnobium) b e s t i i Ren et Bryhn. to elevate the p l a n t to the rank of species.  I t would seem that he wished However, w i t h i n the t e x t  he remarks, " j e c r o i s que l e H. b e s t i i R. et B. sera mieux a sa place comme sous-espece subordonnee au H. molle Dicks, (sensu s t r i c t o ) . "  Here again,  he places the plant as subordinate to H_. molle. Wijk et a l .  (1964) c r e d i t e d Brotherus (1909) w i t h the authorship o f  Hygrohypnum b e s t i i . important p o i n t s .  An examination of Brotherus* treatment reveals two One the presentation o f H. b e s t i i was preceded by that of  157 Hygrohypnum molle (Dicks.)  Secondly, the name H. b e s t i i , i t s e l f , was  immediately preceded by an a s t e r i s k , which denoted s u b s p e c i f i c s t a t u s . For  some reason Wijk e t a l . overlooked the a s t e r i s k .  I t seems c l e a r that  Brotherus wished to t r e a t H. b e s t i i as a subspecies and not as a species. Brotherus (1925) presented a s i m i l a r treatment. John Holzinger (1901) published a note commenting on Hypnum (Limnobium b e s t i i Ren e t Rryhn.  In that paper he noted that Limnobium Schimp. was an  i l l e g a l homonym o f Limnobium Rich.  In that paper he attempted to t r a n s f e r  Hypnum b e s t i i Ren. e t Bryhn to Hygrohypnum, but an e d i t o r i a l e r r o r prevented the t r a n s f e r .  In a footnote, Holzinger (1901a) stated "By typographical e r r o r  i n the January i s s u e , page 12 l a s t l i n e , Hygrohypnum b e s t i i (Ren. e t Bryhn) Holz. was p r i n t e d Hygrohypnum b e s t i i Ren. e t Bryhn. ponsible f o r t h i s . "  The e d i t o r alone i s r e s -  Therefore, as noted by Lawton (1966) Holzinger should be  given r i g h t f u l c r e d i t f o r the t r a n s f e r . Hygrohypnum b e s t i i (Ren. et^ Bryhn) Holz. was described by Renauld £.1901) based on a Holzinger specimen from Avalanche Basin, Montana.  The Renauld  herbarium i s a t PC. A search o f the PC m a t e r i a l has f a i l e d t o produce any Holzinger specimens.  At NY there i s a specimen bearing a l l the data i n d i c a t e d  by Holzinger and c i t e d by Renauld.  This specimen has been appropriately  i d e n t i f i e d and i s designated as the l e c t o t y p e .  158 F i g . 36 a - e.  V a r i a t i o n i n the h a b i t o f the shoots o f Hygrohypnum  b e s t i i and a comparison of i n d i v i d u a l shoots i n the moist and • iV'V-  dry conditions. a.  Moist; b. Dry  c.  A moist shoot w i t h secund leaves  d.  Moist; e. Dry  Scale: Each shoot i s approximately 2 cm long.  159  160 Fig.  a  37  -  i .  V a r i a t i o n  i n  l e a f  a  -  e.  T h e  t y p i c a l  f  -  i .  T h e  l e a f  N o r t h  of  S c a l e :  I  2mm  s h a p e  t h e  o f  A m e r i c a .  I  l e a f  s h a p e  o f  H y g r o h y p n u m  b e s t i i .  s h a p e .  t h e  l o w l a n d  f o r m  f r o m  t h e  P a c i f i c  C o a s t  161  162 F i g . 38 a - i . C e l l u l a r  d e t a i l o f the f o l i a g e leaves o f Hygrohypnum  bestii. a - c. d.  Leaf apices  Median l e a f c e l l s  e - f . Marginal l e a f c e l l s g - h. i.  Alar cells  A l a r c e l l s from the lowland form from P a c i f i c Coastal North America.  Scale: a - c, g - i :  d - f.  I  |  100um *" l  '  100um  l  164  E i g . 3 9 . Hygrohypnum b e s t i i  165 E x s i c c a t i Examined Allen,. Mosses o f the Cascade Mountains # 141 as H. dilatatum. (MICH, MIN, NY, USA) # 139a as H. ochraceum. (TENN, MIN) Grout, North American Musci Pleurocarpi # 251. (MIN, MO, NY, TENN, UC, USA) # 382 as H. molle. (MIN, MO, NY, TENN, UC, USA) Macoun, Canadian Mosses # 398 as H. arcticum. (MICH, MO, USA) Canadian Musci # 357 as H. dilatatum. (NY, USA) # 358 as H. arcticum. (MICH, MO, USA) # 360 as H. ochraceum. (MO) # 483 as Hypnum turgescens. (MICH, NY) Holzinger, Mosses o f Nferth Western Montana # 56. (USA) # 62. (NY) Selected Specimens Examined Canada B r i t i s h Columbia Queen C h a r l o t t e Islands, Graham I . , Mamin R. , S c h o f i e l d 30054. (C, CAN, UBC) Queen Charlotte Islands, Moresby I . , Takakia L.; S c h o f i e l d 24991. (UBC) Manning P r o v i n c i a l Park, Windy Joe Mountain; Williams 329. (UBC) A t l i n , McKee Creek; Szczawinski 70/62. (USA) G a r i b a l d i P r o v i n c i a l Park, Whistler Mountain s k i bowl: ' Jamieson 5480. (UBC) Bulkley Range: Boas 629. (UBC) New Denver, Denver G l a c i e r ; MacPadden 13 Sept 1925. (UBC) Roger's Pass, S e l k i r k Mountains; Macoun 2 Aug 1890. (UBC) Vancouver I s l a n d , Burmah L.; Boas 254. (UBC) Pasulko Lake, N. of Lyttoh; .Schofield 17936. (UBC) Prince Rupert, Slopes o f Hays Mountain; S c h o f i e l d & Sharp 11 Sept 1964. (TENN) Vancouver, Poiftt Grey; S c h o f i e l d 13198. (CANM, UBC as Hygrohypnum ochraceum. Lowland form.) West Vancouver, Lynn Canyon; S c h o f i e l d 13347. (CAN, UBC) Vancouver I . , Bear Creek, San Juan R,; Boas 25. (CAN, UBC) S a l t s p r i n g I ; Weston L.; Boas 424. (CAN, UBC) Alberta K i c k i n g Horse Pass, K i c k i n g Horse L.; Macoun Aug. 1890 as Canadian Musci # 483. (NY) Waterton Lakes National Park, d i v i d e on E. side of Cameron L.; B i r d 6353. (NY) Oldman River Watershed, Race Horse Creek; B i r d and Lakusta 18227. (CAN, UACA) Quebec Gaspe Sud, Gaspe Bay near P e n o ^ i l l e ; Crum and Williams 10798. (UBC) Gaspe Nord, Ruisseau-a-Rebours; Crum & Williams 10747 as H. dilatatum. (UBC)  166  i.  Gaspe Co., St. Joachim de l a T o u r e l l e ; V i c t o r i n e t a l . (MICH) New Brunswick A l b e r t Co., Alma; Habeeb 1640 as H. molle v a r . b e s t i i . (NY) V i c t o r i a Co., Grand F a l l s ; Habeeb 918. (In the l i t e r a t u r e Habeeb 1952) Nova S c o t i a Cape Breton Highlands National Park, Beulach Ban F a l l s j Ireland 11845 as H. dilatatum. (UBC) Cape Breton Highlands National Park, Aspy R.; S c h o f i e l d 4890 as H. dilatatum. (CANM) United States Alaska. Denali Highway, NW of Paxson, Hermann 21225. (CANM,MICH) kodiak I s . Group, Raspberry I , , Port V i t a ; Eyerdam 880 as H. molle. (TENN) A l e u t i a n I s . , A t t u I s . , Massacre Bay; Howard 51 as H, dilatatum. (TENN) Skagway; Williams 27 Aug 1899 as H. b e s t i i (NY) Anchorage, F i r e Lake; Lepage 22312 „ (S-PA) A l e u t i a n I s . , Amchitka I s . , W. o f Banjo Pt.; Shacklette 7621 t (MICH) Unalaska ; S e t c h e l l 28, (UC) Washington Olympic Mountains; Piper Oct. 1890 as L. dilatatum. (NY) Mt. Ranier, Goat Mountain; A l l e n 24 Sept 1898 as H. dilatatum (UBC) Scenic; Frye 3222. (NY) King Co., 2 mi. N, of Cedar F a l l s ; Ireland 5817 (WTU as Hygrohypnum dilatatum. Lowland form.) Oregon Hood R, Co., Mt, Hood, Timberline Lodge; Hale 21576 as H, molle. (UBC) Klamath Co., Crater Lake National Park, Kerr Notch; Hermann 22873 as H. b e s t i i . (CSNM) Multnomah Co., Cloud Cap Inn; S c h o f i e l d 19668 as H. dilatatum. (CANM) Region of the Alsea (a r i v e r ) ; Van Wert May 1923 as H. dilatatum. (MIN) Benton Co., Oak Creek Lab Oregon St. Univ., Hughes. (UBC, lowland form) California Tehama Co., Bridge 8 - 69 on Deer Creek, Highway 32; Kowalski 4 May 1970 as H. molle. (CANM) Plumas Co., Indian F a l l s on fhdtan Creek, Route 89; Lawton 3136 as H. molle. (UBC) Fresno Co., Sequoia National Park, Stoney Creek; MacFadden 8 J u l y 1959 as H. molle. (C§NM) Del Norte Co. G r i f f i n g Creek at Smith River, Highway 199; N o r r i s 8849. (HSC) Siskiyou Co., Salmon R. near B i g Bar; N o r r i s 9280. (HSC) Modoc Co., Soup Creek Campground E, of L i k e l y ; N o r r i s 9451. (HSC)  167  Idaho  Calavaras Co., Bolander (USA) Shasta Co. Lassen N a t i o n a l Park, Dersch Meadows? Mueller 6739. (UC) San Bernadino Co., P a r r i s h 3884. as IH. pseudo-arcticum. (NY, UBC) Marin Co., Muir Woods National Mon,; Hermann 17477 (USA, lowland form) Humboldt Co., P r a i r e Creek, Redwoods State Park; Jamieson 0099. (HSC, lowland form)  Elmore Co. Boise National Forest, Hwy between A t l a n t a and Queen*s R.; MacFadden 18706 as H. molle. (UBC) Idaho Co., 80 mi. ENE o f Kooskis along route 12; Hermann 20210. (UBC) Lake Pend d ' O r i e l l e , Creek near B l u e s l i d e ; Leiberg 1891. (NY) Montana Flathead Co., V i c i n i t y o f Lake McDonald, Avalanche Basinj Holzinger & Blake 28 - 29 J u l y 1898 as Hypnum turgescens i n Mosses of North Western Montana # 62. (NY) Carbon Co., Red Lodge, Bench Creek P i c n i c Area; Conard 27 Aug. 1953 as H. b e s t i i . (NY) Lake Co. Flathead Lake, S t a t i o n Creek; S c h o f i e l d 12189 as H. dilatatum. (UBC) WVoming Yellowstone National Park, Black Sand Basin; Lawton 1871 as H. molle. (CANM) Teton Co., Cascade Canyon; Conard 14 Aug. 1953 as H. b e s t i i . (CANM) Fremont Co., To-Gwo-Tee Pass; Porter 1619 as H. molle. (TENN) Albany Co., Centennial H i l l s ; Nelson 2804, as H. s m i t h i i (TENN) Colorado P i t k i n Co., S. o f Redstone, route 133, C r y s t a l R.-Hays C; Hermann 25635 as H. molle. (UBC) Boulder Co., L e f t Hand Creek; Prettyman 25 Feb 1960. (COLO) Mineral Co. San Juan Mtns., NE of Pagosa Springs; Hermann 23340. (COLO) Jackson Co., Mt. Z i r k e l Wilderness, Lone Pine Creek T r a i l ; Hermann 26740. as H. b e s t i i . (UBC) Grand Co., 1/2 mi S. o f R o l l i n g Pass; Hermann 25660. (WTU) Larimer Co., Rocky Mountain National Park, M i l l Creek; Hermann 25996. (WTU) G a r f i e l d Co., 31 mi. S. o f G&enwood Springs, SW of Trappers Lake; Hermann 24258. (WTU) Clear Creek Co., 6 mi. SW o f S i l v e r Plume, Quale Creek; Hermann 24918. (WTU) New Mexico Rio A r r i b a Co., V i c i n i t y o f Brazos Canyon; Stanley S Bollman 10712 as H. dilatatum. (USA) South Dakota Lawrence Co. Eleven miles SW o f Lead; Hermann 25360. (WTU) Michigan Keweenaw Co., Hebard Park; Steere Sept 1936 as H, molle (MICH) Ontonagon Co., Shores of Lake Superior? N i c h o l s & Steeee 20 - 27 Aug. 1935 as H. molle. (MICH) V  168  Utah Brookbark, C i t y Creek; Flowers 7165  as H. ochraceum.  (DUKE)  169 Hygrohypnum cochlearifolium 1039.  (Vent, i n De Not.)  Broth., Nat. P f l . ,1(3):  1909.  Lectotype:  Erbar. C r i t t .  I t a l . , ser. 2, fasc. 10, n. 453.  (NY) •  Limnobium cochlearifolium Vent, i n De.Not., Erbar. C r i t t . ser. 2, fasc. 10, n 453. 1871.  Ital.,  (Lectotype NY)  Limnobium cochlearifolium Vent, i n De Not. & Bagl., Erbar. C r i t t . I t a l . , ser. 2, fasc. 9 - 11., no. 401 - 550. Genoa. 1871. Hedwigia (5):70 - 73. 1872. Hypnum g o u l a r d i i Schimp., Syn 2:778.  1876.  (Holotype BM) also  i n Rev. Bryol. 3(1):25 - 26. 1876. Hypnum cochlearifolium  (Vent.) Vent., Rev. Bryol. 6(4):62. 1879:  Hypnum arcticum ssp. g o u l a r d i i (Schimp.)  Ren., Rev. Bryol. 10:51.  1883. Hypnum arcticum var. g o u l a r d i i (Schimp.) Husn., Muse. G a l l . 414. 1894.. Calliergon g o u l a r d i i (Schimp.) Kindb., Canad. Rec. S c i . 1894. 72.  1894.  Limnobium g o u l a r d i i (Schimp.) Roth, Die.Eur. Laubm. 2:640. 1905. Hygrohypnum g o u l a r d i i (Schimp.) Loesk., Verh. Bot. Ver. Prov. 46: 198.  1905.  Hygrohypnum s m i t h i i var. cochlearifolium  (Vent.) Monk., Laubm.  Eur. 741. 168b. 1927. Hygrohypnum s m i t h i i var. g o u l a r d i i (Schimp.) Wijk et Marg. Ind. Muse. 5:577. 1969. Names of dubious or otherwise unclear status Hypnum cochlearifolium Vent., Known only from Index Muse. 3:39. 1964. "Limnobium cochlearifolium Vent.", Known only from the above source.  170 Names t r e a t e d as synonyms i n o t h e r p u b l i c a t i o n s , but whose o r i g i n a l l i t e r a t u r e was  unavailable during  this  study.  Amblystegium c o c h l e a r i f o l i u m (Vent. e t _ B o t t . ) et  Broth.,  Herb. Mus.  Genn. ed.  2, 2:66.  L i n b , i n Bomanss 1894.  Amblystegium g o u l a r d i i (Schimp.) C. Jenns., Medd. Groenland. 15:433.  P l a n t s very  1898.  s o f t , forming l o o s e l y woven, e a s i l y fragmenting, o f t e n  mud-clogged patches or s m a l l , t i g h t l y woven p a t c h e s . d i r t y green to y e l l o w ,  u s u a l l y yellow  rusty mottling.  (1) 2 - 3  quently  and  Stems  to y e l l o w - g r e e n w i t h  (5) cm  long.  Mostly  a r i s i n g i r r e g u l a r l y along  branches e i t h e r p r o s t r a t e o r a s c e n d i n g .  brownish c o r t i c a l c e l l s ; m e d u l l a r y c e l l s h y a l i n e or becoming brownish w i t h Plants  age;  age,  mm  and  from the wet  walled,  c e n t r a l strand small, poorly  Leaf  differ-  a t t i t u d e upon the  stem  be l o o s e l y  tumid; a t t i t u d e more v a r i a b l e upon d r y i n g , e i t h e r c o n d i t i o n or v i s i b l y  s h r i n k i n g , l e a v e s near  large plants frequently e x h i b i t considerable  lateral  the  shrink-  o t h e r c r i s p i n g , i n s m a l l e r p l a n t s the l e a v e s r e t a i n t h e i r  c o n c a v i t y upon d r y i n g . (1.3)  thick-walled,  to dry c o n d i t i o n ; when m o i s t the l e a v e s may  to s p r e a d i n g  though l i t t l e  of small,  1 to 2  eradiculose.  from the wet  stem t i p s o f v e r y  and  the stems, a l l  l a r g e r and mostly t h i n n e r  Leaves v a r i a b l e , Crowded to v e r y d i s t a n t .  changing l i t t l e  of s i m i l a r size  Stem c r o s s s e c t i o n s w i t h  rows (sometimes a t h i r d more or l e s s incomplete row)  imbricated  infre-  a r i s i n g from near the stem bases, o r branches s l e n d e r  s h o r t , seldom exceeding 1 cm,  varying  conspicuous  unbranched o r  i r r e g u l a r l y branched, branches e i t h e r l a r g e and  to the stems and  entiated.  Color v a r i a b l e , d u l l ,  Leaves  (0.5)  0.8  - 1.2  (1.5)  mm  long X  (0.3)  0.6  -  1.0  wide; shape u s u a l l y oblong or o b l o n g - e l l i p t i c , sometimes o r b i c u l a r ;  apex u s u a l l y obtuse or rounded, o c c a s i o n a l l y acute,  r e g u l a r l y c u c u l l a t e ; margins  e n t i r e , v a r i o u s l y recurved,  particularly  e s p e c i a l l y i n l a r g e r leaves; concavity s p e c i e s , very  i n the s m a l l e r l e a v e s , o r p l a n e ,  c l e a r l y evident  deep t o c o c h l e a r i f o r m , p l i c a e p r e s e n t  throughout the  throughout,  irregular  in.inumber and arrangement; c o s t a v a r i a b l e , u s u a l l y s h o r t and double one  arm sometimes r e a c h i n g m i d l e a f ,  to midleaf  l e s s o f t e n s h o r t and s i n g l e , s i n g l e  o r absent, i n a l l cases t h e c o s t a i s s l e n d e r ;  narrowly d e c u r r e n t  with  leaves  sometimes  a t the p o i n t o f i n s e r t i o n .  A r e o l a t i o n v a r i a b l e ; median l e a f c e l l s  short fusiform, short  f l e x o s e o r rhombic, 26 - 48 (55) um l o n g X (4) 5 - 6  linear  (8) um wide;  cells  becoming s l i g h t l y s h o r t e r and more rhombic toward the apex; changing  little  toward the margin; m a r g i n a l l e a f c e l l s s h o r t , l e s s than 50 um; b a s a l  cells  v a r i a b l e , g r a d u a l l y becoming s l i g h t l y  l o n g e r and wider or changing  p i t s few t o none; a l a r c e l l s t h i n w a l l e d ,  little;  u n d i f f e r e n t i a t e d or o f a few  quadrate o r s h o r t r e c t a n g u l a r c e l l s , never forming a c l e a r l y d e f i n e d group. S e x u a l i t y p o o r l y understood, p l a n t s u s u a l l y s t e r i l e , unknown.  Outer and middle p e r i c h a e t i a l l e a v e s t r i a n g u l a r l a n c e o l a t e , c o s t a  s h o r t and s i n g l e , margins r e c u r v e d , half.  s l i g h t l y s q u a r r o s e r e f l e x e d i n the upper  Inner p e r i c h a e t i a l l e a v e s oblong l a n c e o l a t e , up t o 2.5 mm  apex mostly acute,  long,  erect  e i t h e r e n t i r e o r f a i n t l y d e n t i c u l a t e , a few p a p i l l a e on  the a b a x i a l l e a f s u r f a c e , margins r e c u r v e d ;  C o s t a s i n g l e , f a d i n g above m i d l e  Seta y e l l o w i s h t o reddish-brown; 12 t o 16 mm in  Perigonial detail  l o n g , smooth; Capsule as  the genus; Spores immature, unmeasurable. Peristome as i n the genus, annulus p r e s e n t ,  with  o f two c e l l rows, endostome  2 to 3 c i l i a .  T h i s p l a n t e x h i b i t s s e v e r a l f e a t u r e s which, i n combination, may to d i s t i n g u i s h the taxon. orbicular.  No o t h e r  The l e a v e s  are broadly  serve  o v a t e , oblong-ovate o r  taxon i n the genus has such deeply  concave l e a v e s .  In  172 some specimens the c o n c a v i t y may leaf.  The  species i s exceedingly s o f t .  to the m o i s t conspicuous green  be as deep as 3/4  stems.  o f the l e n g t h o f the  There i s no r i g i d i t y  I t i s r a r e t h a t patches  whatever  o f these p l a n t s do not  exhibit  r u s t y m o t t l i n g o r v a r i e g a t i o n over a p r i m a r i l y y e l l o w o r y e l l o w -  coloration. C e r t a i n c h a r a c t e r s w i t h i n t h i s taxon e x h i b i t v a r i a t i o n t h a t  discussion.  The  warrants  v a r i e t y v a r i e s i n o v e r a l l s i z e , l e a f shape, shape o f  the  apex, r e c u r v a t u r e o f the l e a f margin, a t t i t u d e o f the l e a v e s upon the stem o r branches,  and  the s p a c i n g o f the l e a v e s along the stem.  In s m a l l e r i p l a n t s  i . e . those w i t h s h o r t stems, the s m a l l l e a v e s are crowded and imbricated. stems. i.e.,  The  The  c o n c a v i t y o f the l e a v e s imparts  t a n t l y spaced,  appressed-  a j u l a c e o u s a s p e c t to the  l e a v e s are r e g u l a r l y r e c u r v e d a l o n g the margin.  those whose stems vary from 3 to 4 cm  closely  In l a r g e r p l a n t s ,  l o n g , the l e a v e s are more d i s -  o f t e n e x c e e d i n g l y concave and more o r l e s s e r e c t - s p r e a d i n g to  sometimes almost  e r e c t when m o i s t .  o l d e r p o r t i o n s o f the stems w i l l be  In these l a r g e p l a n t s the l e a v e s i n the crowded and r a t h e r j u l a c e o u s , whereas  younger l e a v e s , though o f equal s i z e , w i l l be d i s t a n t and more s p r e a d i n g . Both V e n t u r i i n De N o t a r i s (1871, 1872) :  narrowly  r e c u r v e d l e a f margins.  Nyholm  and Schimper  (1876) noted  (1965) a l s o remarked upon t h i s  a c t e r i n d i s t i n g u i s h i n g the v a r i e t y from Hygrohypnum norvegicum. is variable. recurved  Among s m a l l p l a n t s the margin i s r e g u l a r l y and  ( F i g . 40 h - k ) .  ( F i g . 40 f ) .  looks v i r t u a l l y  l i k e a spoon.  char-  This feature  conspicuously  In l a r g e r p l a n t s the margin i s r e c u r v e d  a - e) o r p l a n e  the  ( F i g . 40  In those l e a v e s w i t h p l a n e margins the  leaf  These p l a n t s vary c o n s i d e r a b l y i n the placement o f the l e a v e s a l o n g stems and branches.  In some specimens they may  they are so w i d e l y spaced  as to b a r e l y o v e r l a p .  the  be crowded, w h i l e i n o t h e r s  173 The leaves of Hygrohypnum cochlearifolium are often i r r e g u l a r l y plicate.  However these p l i c a e are not a c l e a r l y integral part of the  leaves as are the p l i c a e of the p e r i c h a e t i a l leaves.  Instead, they appear  to be a function of the extremely f l a c c i d nature of the leaves and t h e i r corresponding  i n a b i l i t y to r e s i s t the various physical pressures of water  or the s o i l and sand p a r t i c l e s that accumulate i n the leaf concavity.  The  p l i c a e are generally more frequent among the larger leaves, perhaps as a r e s u l t of more exposed surface area. thoroughly  Generally leaves that have been  cleaned during examination have few i f any p l i c a e .  The sexual condition of Hygrohypnum cochlearifolium i s unclear. authors have described i t as monoicous. dioicous.  Only Kindberg (1897) has said i t i s  The species i s most frequently s t e r i l e .  perigonia has been examined. quently sexual or i t may  Most  The plants may  Only one plant bearing  be monoicous and very i n f r e -  be c l e a r l y dioicous.  This matter has yet to be  resolved. The nomenclature of Hygrohypnum cochlearifolium (Vent.; Broth, i s highly confused.  i n De  Not.)  This species was o r i g i n a l l y described very  c l e a r l y by Venturi as Limnobium cochlearifolium i n De Notaris' Erbar. Cirittog. I t a l . Ser. II n. 453 i n 1871.  The label of this exsiccatum  a complete description and a reference to a c o l l e c t i n g s i t e . specimen of this important  bears  The only  e x s i c c a t i available to me i s at NY, where i t i s  f i l e d with Hygrohypnum s m i t h i i .  The rules of nomenclature indicate that on  or a f t e r 1 January 1953 only descriptive material d i s t r i b u t e d independent of an e x s i c c a t i may  effect valid publication.  well within the l i m i t .  The 1871  date of issue i s  Further, De Notaris and Baglietto (1872) published  the label of the e x s i c c a t i . Index Muscorum, vol 3 c i t e s the name Hygrohypnum cochlearifolium De  Not.,  174 Erbar.  Critt.  Ital.  s e r . 2, 10:n. 453.  1871 as nom. nud. i n synon. Index  Muscorum f u r t h e r i n d i c a t e s t h i s name t o be synonomous w i t h c o c h l e a r i f o l i u m Vent.", an obvious  "Liimobium  m i s s p e l l i n g , which i s i n t u r n h e l d as  synonympjUS w i t h Hygrohypnum s m i t h i i (Sw.) B r o t h , v a r . g o u l a r d i i Wijk e t Marg. and  (Schimp.)  I t i s obvious t h a t Limnobium c o c h l e a r i f o l i u m Vent, i n De Not.  Hypnum c o c h l e a r i f o l i u m De Not. a r e based on the same e x s i c c a t i .  A  c a r e f u l examination o f the l a b e l borne on t h i s e x s i c c a t i c l e a r l y shows o n l y the name Limnobium c o c h l e a r i f o l i u m . Nowhere on the l a b e l i s t h e r e  an i n d i -  c a t i o n o f the use o r t h e p r e f e r e n c e f o r the use o f Hypnum i n s t e a d o f Limnobium. Further  confusion  cochlearifolia  i s added by the f a c t t h a t the basionym f o r Weymouthia  (Schweagr.) D i x . i s Hypnum c o c h l e a r f o l i u m  Muse. Suppl. 1(2):221. Not.  a reality,  88.  1816.  Schweagr., Spec.  Were the name Hypnum c o c h l e a r i f o l i u m De  i t would s t i l l be a l a t t e r homonym o f the e a r l i e r Schweag-  r i c h e n name. Schimper  (1876) d e s c r i b e d  G o u l a r d i n the Pyrenees.  Hypnum g o u l a r d i i from m a t e r i a l  c o l l e c t e d by  An examination o f the Schimper type a t BM r e v e a l s  t h a t i t u n q u e s t i o n a b l y the same organism as Limnobium c o c h l e a r i f o l i u m Vent Venturi  "(1879) c l o u d e d the i s s u e , h i m s e l f ,  by c i t i n g Hypnum c o c h l e a r i -  forme as a synonym o f "Hypnum g o u n o d i i Schimp." an obvious m i s s p e l l i n g o f Hypnum g o u l a r d i i Schimp.  T h i s would seem t o imply t h a t V e n t u r i  h i s own p l a n t as i d e n t i c a l t o Schimper's l a t t e r named p l a n t . Venturi  l o o k e d upon  I f i n this act  t r a n s f e r r e d h i s Limnobium c o c h l e a r i f o l i u m t o Hypnum c o c h l e a r i f o l i u m  as a new combination and a synonym o f Hypnum g o u l a r d i i Schimp., then the new combination becomes an i l l e g a l homonym o f Hypnum c o c h l e a r i f o l i u m Schwaegr. In 1884 V e n t u r i An  and B o t t i n i r e p o r t e d  Limnobium c o c h l e a r i f o l i u m f o r I t a l y .  u n f o r t u n a t e m i s i n t e r p r e t a t i o n o f the V e n t u r i  r e p o r t by Van d e r Wijk and  175  Margadant (1969) treated Venturi and B o t t i n i as the parenthetical authors of Hygrohypnum cochlearifolium as c i t e d by Brotherus '(1909) and H_. s m i t h i i var. cochlearifolium of Monkemeyer  (1927).  Brotherus (1909) c l e a r l y referred  his name to Limnobium cochlearifolium i n Venturi's e x s i c c a t i . (1927) also referred h i s name to Venturi. Venturi' s use of Hypnum.  Monkemeyer  However, Monkemeyer curiously c i t e d  I- have not been able to trace Venturi's  apparent use  of the name Hypnum of h i s species. Confusing though t h i s s i t u a t i o n i s , Limnobium cochlearifolium Vent, i n De Not. i s the basionym f o r Hygrohypnum cochlearifolium (Vent, i n De Not.) Broth. Various authors have treated Hygrohypnum cochlearifolium as a variety. (Husnot, 1894- Monkemeyer, 1927) or a subspecies (Renauld, 1883) of H_. s m i t h i i . However, neither these authors nor Boulay (1884) adequately c l a r i f i e d the s i m i l a r i t i e s between the two taxa.  Several features indicate that Hygrohypnum  cochlearifolium i s a species c l e a r l y d i s t i n c t from H. s m i t h i i . cochlearifolium i s exceedingly s o f t .  Hygrohypnum  No other taxon i n the genus i s as s o f t .  Hygrohypnum s m i t h i i i s exceedingly s t i f f and r i g i d .  Neither taxon exhibits  any r e c i p r o c a l tendency towards softness or r i g i d i t y . and Hygrohypnum cochlearifolium have concave leaves.  Both Hygrohypnum s m i t h i i However, the leaf concav-  i t y of Hygrohypnum smithii i s simply not comparable to the cochleariform.-concavity of H_. cochlearifolium  The o v e r a l l areolation of the two taxa i s d i f f e r e n t .  Associated with the r i g i d nature of Hygrohypnum s m i t h i i i s the incrassate nature of the leaf c e l l s .  On the contrary, the leaves of Hygrohypnum cochlearifolium  are very f l a c c i d and correspondingly  thin-walled.  are also incrassate and yellowish i n color. l e a r i f o l i u m are thinner-walled, brown i n color.  The basal c e l l s of H. s m i t h i i  The basal leaf c e l l s of H. coch-  sometimes lax and either hyaline or s l i g h t l y  The costa of H_. s m i t h i i i s usually thick, yellowish, single  176 and/or f o r k e d , although s h o r t and  double costae  s t o u t nature o f the c o s t a imparts c o n s i d e r a b l e On  are not i n f r e q u e n t . rigidity  t o the  The  leaves.  the o t h e r hand, the c o s t a of Hygrohypnum c o c h l e a r i f o l i u m i s u s u a l l y  slender,  faint,  s h o r t and  double.  The  s l e n d e r and  f a i n t q u a l i t y of  c o s t a no doubt i n f l u e n c e s the f l a c c i d nature o f the l e a v e s ,  The  the  leaf  margins o f Hygrohypnum s m i t h i i are always p l a n e whereas those o f H_. cochl e a r i f o l i u m may  be p l a n e o r r e c u r v e d .  Although the i n n e r p e r i c h a e t i a l  leaves  o f both s p e c i e s have r e c u r v e d margins, the' apex o f the i n n e r p e r i c h a e t i a l leaves  i s rounded, whereas t h a t of H.  cochlearifolium is  acute.  For a comparison between Hygrohypnum c o c h l e a r i f o l i u m and H. see  the d i s c u s s i o n under the l a t t e r  species.  norvegicum  177 F i g . 40 a - m.  V a r i a t i o n i n the l e a f shape and  shoots o f Hygrohypnum a - c , e - f , h - k .  the h a b i t o f the m o i s t  cochlearifolium. V a r i a t i o n i n l e a f shape.  d.  L e a f " c " as seen i n c r o s s - s e c t i o n .  Note the r e f l e x e d  g.  L e a f " f " as seen i n c r o s s - s e c t i o n .  Note the p l a n e  1 - m.  V a r i a t i o n i n the h a b i t o f the s h o o t s .  Scale: a - k;  1 - m;  I  1  m  m  1  Both shoots are a p p r o x i m a t e l y 0.6  cm  long.  margins,  margin.  178  179 F i g . 41 a - e.  C e l l u l a r d e t a i l of the f o l i a g e leaves of Hygrohypnum  cochlearifolium. a.  Leaf apex,  b - c.  Median l e a f c e l l s ,  d - e.  Alar c e l l s .  Scale:  \  a - c;  I  d - e;  I  '  IQOum  100um  )  |  181  4-  •  -  F i g . 42. Hygrohypnum c o c h l e a r i f o l i u m  182 E x s i c c a t i Examined De N o t a r i s , E r b a r . C r i t t . I t a l . s e r . 2, f a s c . 10, n. 453 as Limnobium cochlearifolium. (NY, L e c t o t y p e ) I n s t i t u t i u m Botanicum nomine V. L. Komarovii Academiae S c i e n t i a r u m URSS, H e p a t i c a e e t Musci URSS E x s i c c a t i # 48 as Hygrohypnum c o c h l e a r i f o l i u m . (CAN, LE, UBC) Lisowski, Bryotheca Polonica. F a s c . X, # 296. (BP, LE) F a s c . XXV, # 667. (BP. CAN) F a s c . XXIV, # 637 as H. v i r i d u l u m (BP, CAN, LE) F a s c . XXIV # 638 (BP, CAN) F a s c . XV, # 418. (LE) Museo H i s t . H a t u r . V i n d o b o n e a s i , Cryptggame e x s i c c a t i # 2895. (BP, UC, F) Selected  Specimens Examined  Canada Yukon, Haines Highway, M i l e 100; Crum and S c h o f i e l d 9637 as H. norvegicum. (CAN, MICH) Northwest T e r r i t o r i e s , B a f f i n I s . , Cape S e a r l e ; Hale 17 - 18 Aug 1950 as H_. s m i t h i i v a r . g o u l a r d i i . (UBC) United States Alaska Brooks Range, E n d i c o t t Mtns; S t e e r e 18115a as H_. c o c h l e a r i f o l i u m . (NY) Mt. McKinley N a t i o n a l Park, Mt. E i e l s o n ; Weber & V i e r c k 10301 as H_. c o c h l e a r i f o l i u m . (CAN, NY,), DUKE) Montana E l e c t r i c Peak; Rydberg & Bessey 18 Aug. 1897 as L . . g o u l a r d i i . (NY) . Colorado C l e a r Creek Co., Mt. Evans; Vaarama & Evans 11 O c t . 1963. .((COLO) Greenland • Godhavn; Berggren 1870 as H. norvegicum. (NY, S-PA, P) D i s c o I s . D i s k o f j o r d , K u a n i t ; P o r s i l d 954 as H. c o c h l e a r i f o l i u m . (CANM) Svartenhuk Penn. S i m i u t a p kua; Holmen 12350 as H_. c o c h l e a r i f o l i u m . (CAN) S c o r e s b y l a n d , M e s t e v i g ; Holmen 18043 as H. c o c h l e a r i f o l i u m . (CANM) Nugssuag Penn., ,Nugssuag; Holmen 15577. (S-PA, LE) M a r t l e k ; Beggren 1870.- (S-PA) Hurry I n l e t ; Dusen 7 Aug. 1899. (H) Spitzbergen Advent Bay; Ber^gr.eri 1868 as H_. m o l l e . (S-PA) Robbe Bay; Berggren 1868 as H. m o l l e . (S^-PA) Norway Tromso amt, N o r d r e i s e n G a l s l s o v a r e ; A r n e l l 23 Aug. 1893. as H_. goulardii. (S-PA) F i l e j e l d ; Bryhn Aug. 1887 as H. g o u l a r d i i . (S-PA) Dovre r e n s i s Foksjriko,* Bryhn Aug. 1906 as H. c o c h l e a r i f o l i u m . (S-PA)  183 Finmarken, T a l i r k , Jorgensen 27 J u l y 1890 as H_. g o u l a r d i i . (S-PA, H, NY) Sweden H a r j e d a l e n , Storsrjo, Smith 10 Aug. 1914 as H. c o c h l e a r i f o l i u m . (S-PA) Lule  Lappmark, T a r r e k a i s s e ,  Nilsson  12 Aug. 1897 as H_. norvegicum.  (S-PA) L u l e Lappmark, S a r j e k e n s i s , Rapsdalen, Jensen & A r n e l l 31 J u l y 1902 as A. c o c h l e a r i f o l i u m (S-PA, H) Torne Lappmark, F s a s i r u j a k a J u k a j a r v i , Hulphers 18 J u l y 1944 as H_. c o c h l e a r i f o l i u m (S-PA) Finland Karesuando, Smith & A r n e l l 14 Aug. 1939. (S-PA) F i n s k a Lapm. E u o n t e l i s , K o t i l a i n e n 10 J u l y 1920 as A. v i r i d u l u m . (S-PA) • Prov. L e . , Par O-Kahpeusvarri, R o i v a i n e n 352 as H. v i r i d u l u m . (S-PA, UAC) Soviet Union K o l a Pen., montes C h i b i n y , S c h l j a k o v 24 Aug. 1947. (UBC, CANM) J e n i s e i , N i s c h z n j e Tauguska, A r n e l l 13 J u l y 1876 as A. v i r i d u l u m . (S-PA) Switzerland V a l a i s , S t . Bernard, Amann 19 J u l y 1932 as H. c o c h l e a r i f o l i u m . (S-PA) V a l a i s , Mt. Mora, Culmann Aug. 1886 as H. g o u l a r d i i . (S-PA) Bern, F a u l h o r n , Culmann 16 Aug. 1885 as H. g o u l a r d i i . '(H) Austria T i r o l , Glungezer N o r d s e i t e , S c h i f f n e r 4 Sept. 1903. (S-PA) K a r n t e n , F o l l a c h , B r e i d l e r 6 Aug. 1861 as H. g o u l a r d i i (S-PA, H) S t e i e r m a r k , Schladming, B r e i d l e r 20 Aug. 1882 as H. c o c h l e a r i f o l i u m . (H) S a l z b u r g , S u l z b a c h t h a l , B r e i d l e r 13 Aug. 1890 as K. g o u l a r d i i . (H) Italy T i r o l I t a l i a n o , Saent, V e n t u r i , E r b a r , C r i t t , I t a l . s e r . 2 n. 453 as L. c o c h l e a r i f o l i u m . (NY) Spain G a l l i a m e r i d . M a l a d e t t a , G o u l a r d 187 3 as H_. g o u l a r d i i . (BM) Poland Montes T a t r i , L i s o w s k i 1 Sept 1957 as H_. c o c h l e a r i f o l i u m B r y o t h e c a P o l o n i c a # 638. (CAN) Czechoslovakia Vysoke T a t r y , M l y n i c a , P i l o u s 263 as H. c o c h l e a r i f o l i u m .  and  (CAN)  184 Hygrohypnum norvegicum 358.  (Schimp.) Amann, F I . Mouss. S u i s s e 1:188  e t -2:  1912. Hypnum v i r i d u l u m Hartm., Handb. F l . ed. 5:324. (Holotype,  1849.  horn, i l l e g .  S-PA)  Limnobium norvegicum Schimp., B r y o l . Eur. 6:70.  576.  Hypnum norvegicum  1860.  (Schimp.) Schimp., Syn.  Amblystegium v i r i d u l u m L i n d b . , Limnobium v i r i d u l u m Calliergon viridulum  (Lindb.)  637.  Musci Scand.  33.  1879.  Kindb., Canad. Rec.  (Lindb.)  1853.  Sc. 6(2):74.  Kindb., Eur. N. Am.  B r y i n . 1:86.  Hygrohypnum v i r i d u l u m (Hartm.) B r o t h . , Nat. P f l . 1(3):1039.  P l a n t s forming  s m a l l , s o f t , l o o s e l y to t i g h t l y woven t u f t s o r  Color u s u a l l y shiny, pale yellow-green, w i t h age.  Stems 1 to 3 cm  l o n g , ascending.  Branching  1897. 1909.  patches.  o c c a s i o n a l l y becoming a l i g h t brown  l o n g , u s u a l l y l e s s than 1.5  the t i p s s l i g h t l y ascending.-  1894.  cm,  p r o s t r a t e or w i t h  i r r e g u l a r , branches 1 . 0 - 1 . 2  (1.5)  cm  Stem c r o s s - s e c t i o n s r e v e a l i n g 2 to 3 rows o f s m a l l , t h i c k -  w a l l e d , brownish c o r t i c a l c e l l s ; medullary  c e l l s l a r g e r and  thinner-walled;  c e n t r a l strand present, often d i s c o l o r e d .  R h i z o i d s v a r i a b l e , red-brown,  sparse,  a r i s i n g from the base o f v e n t r a l stem l e a v e s . Leaves ovate, 0.4  - 0.5  (0.6)  b r o a d l y ovate  mm  r a r e l y b r o a d l y ovate;  (0.4)  0.5  l e a v e s ; sometimes the apex may  be  o r r a r e l y v e r y narrowly  slightly recurved,  but never e n c l u d i n g the l e a f apex, r e c u r v a t u r e may of the l e a f o r may  leaves plane arm  (0.9') mm  long X  (0.2)  wide; apex u s u a l l y acute, o c c a s i o n a l l y obtuse i n the more  e n t i r e , v a r i o u s l y plane  1/4  - 0.8  squarrose;  margins  recurvature v a r i a b l e ,  be c o n f i n e d t o the  lower  o c c u r throughout the margin e x c l u s i v e o f the apex;  t o s h a l l o w l y , but c l e a r l y ,  concave; c o s t a short'and  o f t e n r e a c h i n g m i d l e a f , very r a r e l y s l e n d e r and  double,  s i n g l e t o j u s t above  one mid-  185 leaf.  Leaf  a t t i t u d e upon the stem v a r y i n g from the wet t o d r y conditions,-  when wet the l e a v e s a r e s t r a i g h t and l o o s e l y i m b r i c a t e d t o l o o s e l y v e r y r a r e l y they may be s l i g h t l y  spreading,  f a l c a t e ; when d r y the l e a v e s tend t o become  crisped, usually shrinking l a t e r a l l y  and c o n c o m i t a n t l y  i n r o l l i n g or t w i s t i n g .  A r e o l a t i o n v a r i a b l e ; median l e a f c e l l s s h o r t rhombic, b a c i l l i f o r m , form o r l i n e a r f l e x u o s e , decreasing  slightly  (16) 20 - 30 (48) mm long X 5 - 6 mm wide;  i n l e n g t h toward the apex; m a r g i n a l  leaf cells  fusi-  cells always  l e s s than "50 um; b a s a l c e l l s v a r i a b l e , g r a d u a l l y e l o n g a t i n g and widening from the median c e l l s , o r widening s l i g h t l y , b u t changing l i t t l e changing l i t t l e w i t h age; p i t s  i n length, or  i n l e n g t h o r w i d t h , h y a l i n e or becoming s l i g h t l y golden-brown few t o none; a l a r c e l l s u n d i f f e r e n t i a t e d o r forming  a small  i r r e g u l a r group o f a few, quadrate t o s h o r t r e c t a n g u l a r , s l i g h t l y i n c r a s s a t e c e l l s , h y a l i n e o r becoming brown w i t h age. P l a n t s monoicous, mm l o n g , deeply  P e r i g o n i a ovate;  concave i m b r i c a t e d ,  p e r i g o n i a l l e a v e s ovate,  e c o s t a t e , margins e n t i r e .  0.2 t o 0.6  Outer and  middle p e r i c h a e t i a l l e a v e s ovate t o ovate l a n c e o l a t e , s q u a r r o s e - r e f l e x e d i n the apex, e c o s t a t e t o s h o r t s i n g l e o r s h o r t and double, margine e n t i r e . p e r i c h a e t i a l l e a v e s l a n c e o l a t e , e r e c t , up t o 1.8 mm t a p e r i n g t o a p o i n t o r acute, reaching midleaf,  Inner  l o n g , apex g r a d u a l l y  c o s t a v a r i a b l e s i n g l e and s l e n d e r o r double,  2 t o 4 p l i c a e , c e l l s from the base t o m i d l e a f  margins e n t i r e t o very f i n e l y and i r r e g u l a r l y  pitted,  toothed.  Seta 6 t o 10 mm l o n g , y e l l o w t o reddish-brown, smooth-walled, s t r a i g h t or s l i g h t l y  curved when wet, v a r i o u s l y t w i s t i n g and c o n t o r t i n g upon d r y i n g .  Capsule as i n the genus. Peristome as i n the genus. yellow,  9 t o 17 um i n diameter.  o f t e n f a l l i n g away.  Annulus o f two c e l l rows. Endostome w i t h  Spores dark, dusky-  2 t o 3 w e l l developed  cilia,  186 In s p i t e o f a monographic study, Hygrohypnum norvegicum remains a p o o r l y understood  species.  from which t o generate In combination, norvegicum.  There, are j u s t too few  specimens o f t h i s  taxon  a sound concept o f i t s v a r i a b i l i t y .  f o u r c h a r a c t e r s may  serve t o d i s t i n g u i s h Hygrohypnum  These c h a r a c t e r s are the v e r y t i n y l e a f s i z e , the ovate  leaf  shape, the acute l e a f apex and the l o o s e l y s p r e a d i n g nature of the m o i s t leaves. In h i s o r i g i n a l d e s c r i p t i o n Schimper  (1853) remarked upon the  i t i e s and d i f f e r e n c e s between L. norvegicum and what i s now hypnum l u r i d u m .  L a t e r , he  (Schimper,  1876)  similar-  known as Hygro-  noted the s i m i l a r i t y of H.  icum and H^ m o l l e , as d i d L i m p r i c h t (1904) .  norveg-  From the p r e s e n t s t u d i e s i t seems  e n t i r e l y u n l i k e l y t h a t H_. norvegicum c o u l d be confused w i t h e i t h e r H_. l u r i d u m o r H. m o l l e .  The  f o l l o w i n g c h a r t : may  serve t o c l a r i f y the  H. norvegicum L e a f Length L e a f Apex  0.4  t o 0.9  mm  Acute  L e a f Apex Margin  Entire  t o 2.0 mil  Alar cells  U n d i f f e r e n t i a t e d or s m a l l group o f quadrate to short r e c tangular c e l l s .  Plane  c l e a r l y s e p a r a t e them.  H_. l u r i d u m 0.9  t o 2.5  mm  Gradually tapering to a p o i n t . Entire  Plane t o i n r o l l i n g i n the upper 1/2.'  Undifferentiated  very s m a l l e s t l e a v e s of H^ l u r i d u m may  l a r g e s t l e a v e s o f H_. norvegicum.  and b l u n t  D e n t i c u l a t e or entire  Plane t o v a r i o u s l y recurved  will  0.9  Acute  L e a f Margins  The  H. m o l l e  situation.  Well-developed group o f quadrate to; s h o r t r e c t a n ular c e l l s .  o v e r l a p i n s i z e w i t h the very  However, the d i f f e r e n c e s mentioned above  ' 187 In Europe and North America  l e a f l e n g t h may  be used as a g e n e r a l  c r i t e r i o n f o r d i s t i n g u i s h i n g Hygrohypnum m o l l e from H. norvegicum. Europe the average l e a f l e n g t h f o r H_. m o l l e i s 1.2  t o 1.7  mm  l e a v e s have e v e r been observed e x h i b i t i n g s h o r t e r l e n g t h s . North America the average l e a f l e n g t h f o r H_. m o l l e i s 0.9 it  In  and very few In Western  t o 1.3 mm  and  i s , t h e r e f o r e , u n d e r s t a n d a b l e t h a t v a r i o u s r e p o r t s o f H_. norvegicum  from south o f 60°N. have been based upon m i s d e t e r m i n a t i o n s o f H_. m o l l e . The specimen o f H_. norvegicum, which e f f e c t s the remarkable of the s p e c i e s from A l a s k a i n t o s o u t h e r n B r i t i s h Columbia,  disjunction clearly  differs  from H. m o l l e by i t s e n t i r e , r e c u r v e d l e a f margins and i t s v e r y s h o r t l e a v e s (0.7' mm) . The s p e c i f i c l i m i t s between Hygrohypnum norvegicum and H_. c o c h l e a r i f o l i u m are weak.  Both t a x a tend t o merge as a consequence  of s i m i l a r patterns of  v a r i a t i o n i n l e a f dimensions, l e a f c o n c a v i t y , l e a f m a r g i n a l r e c u r v a t u r e and the nature o f the l e a f apex.  The f o l l o w i n g c h a r t w i l l p o i n t out the g e n e r a l d i f f e r -  ences .  H_. norvegicum L e a f Length  0.4  t o 0.9  L e a f Shape  Ovate  L e a f Apex  Acute  mm  H_. c o c h l e a r i f o l i u m 0.8  t o 1.5  mm  B r o a d l y ovate t o almost orbicular Obtuse  Leaf concavity  Plane t o Shallow  Deeply concave t o cochleariform  L e a f margins  Plane to r e c u r v e d  Recurved t o p l a n e  The l e a v e s o f H_. norvegicum are g e n e r a l l y l o o s e l y i m b r i c a t e d t o s l i g h t l y spreading.  The l e a v e s o f H. c o c h l e a r i f o l i u m may  a l s o be l o o s e l y  imbricated,  b u t i n v e r y s m a l l specimens, which are most r e a d i l y c o n f u s e d w i t h H.  norvegicum,  188 the deep l e a f c o n c a v i t y imparts a d i s t i n c t l y plants.  tumid appearance t o the  The l e a f a p i c e s o f H_. norvegicum are almost always a c u t e .  The  l e a f a p i c e s o f H_. c o c h l e a r i f o l i u m are g e n e r a l l y obtuse, b u t . i n s m a l l specimens c e r t a i n l e a f a p i c e s may Schimper  be a c u t e .  (1853) d e s c r i b e d Limnobium  from G u l b r a n d s d a l e n , Norway.  Nyholm  norvegicum based on a B l y t t  (1965) o f f e r e d two s e t s o f  t i o n s o f H. norvegicum, one o f which was  specimen  illustra-  a s c r i b e d t o the B l y t t t y p e .  In  p e r s o n a l communications Nyholm remembered t h a t the B l y t t specimen was i n the Schimper c o l l e c t i o n a t BM.  A thorough s e a r c h o f the BM m a t e r i a l has  not y i e l d e d the B l y t t specimen.  Hartmann  (1849) d e s c r i b e d Hypnum v i r i d u l u m .  Hartmann's s p e c i e s i s the same taxon as Limnobium a l a t e r homonym o f Hypnum v i r i d u l u m B r i d . Holmgren  norvegicum, however, i t i s  The Hartmann type, a t S-PA  specimen from S a r k e v a r e , L u l e Lappmark, Sweden.  illustrations  c i t e d by Nyholm  the Hartmann type.  is a  The second s e t o f  (1965) f o r Hygrohypnum norvegicum was based on  A c a r e f u l study o f a l l  the a v a i l a b l e d e s c r i p t i o n s and  specimens has shown t h a t I c o m p l e t e l y agree w i t h Nyholm's treatment o f the Scandanavian m a t e r i a l .  Based on my  agreement w i t h her work, I can o n l y assume  t h a t she a c c u r a t e l y a s s e s s e d the c o n s p e c i f i c i t y o f the Schimper and Hartmann types.  I have a c c o r d i n g l y based my  Hartmann type.  concept o f Hygrohypnum norvegicum on the  189 F i g . 43 a - j . V a r i a t i o n i n the l e a f shape, the c e l l u l a r d e t a i l o f the f o l i a g e l e a v e s and the h a b i t o f the shoots o f Hygrohypnum norvegicum. a - d.  L e a f shape  e - f.  The h a b i t o f the m o i s t  g - h.  Leaf  shoots.  apicies  i.  Alar  cells  j.  Median l e a f  cells  Scale:  a-e  e - f;  a  _  I 0.5mm  ;  1  Both shoots a r e a p p r o x i m a t e l y 0.5 cm l o n g . I  '  100um  I  190  191  -IF i g . 44-  Hygrohypnum norvegicum  192 S e l e c t e d Specimens Examined Canada B r i t i s h Columbia Salmo a r e a , Porcupine Greek; Jamieson # 5598. (UBC) United States Alaska Mt. McKinley N a t i o n a l Park, t r a i l above R i l e y Creek; C r o a s d a l e B-23. (USA) Greenland Scoresby Sund, R y p e f j o r d ; Holmen 18767 as Hygrohypnum v i r i d u l u m . (NY) Norway L i l l e E l v e d a l , T e a d f a l e r l a c a Gronne G r o t t o ; K a u r i n Aug. 1892 as Hypnum v i r i d u l u m . (S-PA)L i l l e E l v e d a l , prope F l a d f a t e r , "Gronne G r o t t o " ; K a u r i n Aug. 1888 as Hypnum v i r i d u l u m . (S-PA) Dovre, Knudshoe; Bryhn Aug. 1887. (S-PA) Dovre, Z e t t e r s t e d t 11 J u l y 1854. (C, S-PA)L i l l e E l e v e d a l , Gronne G r o t t o ; K a u r i n Aug. 1887. (S-PA) Sweden Norbotten, O v e r t o r n e a ; L o n n q v i s t 22 Aug. 1963 as Hygrohypnum norvegicum. (S-PA) Jamtland, Undersakers s;n S y l a r n a ; K r u s e n s t j e r n a 12 J u l y 1938 as Hypnum v i r i d u l u m . (S-PA) P i t e Lappmark, A r j e p l o g sn, L u l . I s t j a k k ; Wistrand 2 J u l y 1936 as Hypnum v i r i d u l u m . (S-PA) L u l e a Lappmark, Sarkavaare; Holmgren June 1848, c i t e d as Amblystegium v i r i d u l u m . - (Holotype f o r Hypnum v i r i d u l u m , S-PA) L u l e a Lappmark, S t . S j o f a l l e t ; Holmgren 9 J u l y 1867. (S-PA) Lapponia L u l e n s i s , Sareks; Jensen & A r n e l l 3 Aug. 1902 as Amblystegium v i r i d u l u m . (C, S-PA) Finland Lapp. ov., In a l p e S c h e l e s n a j a prope pagu K a n t a l a k s ; B r o t h e r u s 29 J u l y 1872. (H-SOL) Austria S t e i e r m a r k , Schladming; B r e i d l e r 27 Aug. 1869 as Limnobium sp. (BP)  193 Hygrohypnum m o l l e  (Hedw.) Loesk., M o o s f l . Harz. 320.  1903..  Neotype: S c o t l a n d , Ben Mac Dhui, by s t r e a m l e t s from p e r p e t u a l snow: G.-.-Ei Hunt, J u l y 1871.  (S-PA)  Hypnum m o l l e Hedw., .Spec. Muse. 273. • 70f. 7 - 1 0 .  1801.  Hypnum r u p e s t r e S c h l e i c h . ex Spreng. Mant. Prim. F I . Halens. 56. 1807. Hypnum m o l l e v a r . r i g i d u l u m Hartm., Handb. Skand. F I . ed. 5:324. 1849.  nom. i l l e g . i n c l .  type spec.  Limnobium m o l l e Schimp., B r y o l . e u r . 6:69. 576. 577. 1853. Hypnum m o l l e v a r . giganteum 1331.  1863.-  L o r . , V e r h . Zoo. B o t . Ges. Wien 13:  nom. nud.  Hypnum Schimpefianum L o r . , Moostud.  123. 5c. 1864. (Holotype, G)  Hypnum taurense M o l . i n L o r . , Moostud.  123. 1864. nom. nud. i n synon.  Hypnum a l p e s t r e v a r . t u r g e s c e n s L o r . , Moostud. 123. in  1864. nom. nud.  synon.  Hypnum m o l l e v a r . schimperianum  (Lor.) Schimp., Syn. ed. 2:775. 1876.  Hypnum m o l l e v a r . s c h i m p e r i L i n d b . , B o t . Not., 1877:30. Amblystegium m o l l e  (Hedw.) L i n d b . , Musci Skand.  Amblystegium m o l l e v a r . schimperianum 33.  1877.  33. 1879.  (Lor.) L i n d b . , Musci Skand.-  1879.  Hypnum d i l a t a t u m s s p . m o l l e  (Hedw.) Ren., Rev. B r y o l . 10:51.  1883.  Hypnum m o l l e v a r . maximum B o u l . , Muscin. France 24. 1884. Limnobium submolle Kindb., C a l l i e r g o n submolle C a l l i e r g o n molle  Rev. B r y o l . 22:87.  (Kindb.)Kindb.,' Bot. Not. 1896:132.  (Hedw.) Kindb., E u r . N. Am. B r y i n .  Hypnum m o l l e v a r . pyreniacum 10:8.  1901.  1895. 1896.  1:84. 1897.  Ren., B u l l . A c t . . I n t . Ge©gr. B o t .  194  Hypnum b e s t i i v a r . pyreniacum  (Ren.) Ren.,  Rev.  Bryol.  28:8.  1901. Hygrohypnum m o l l e v a r . 49:52.  schimperianum  (Lor.)  Loesk., Hedwigia  1909.-  Hygrohypnum m o l l e v a r . pyreniacum  (Ren.) Podp., Consp.  576.  1954.  Plants  s o f t , i n l o o s e l y to t i g h t l y woven mats o r p a t c h e s .  Color  v a r i a b l e , b r i g h t y e l l o w - g r e e n , o l i v e - g r e e n , y e l l o w - g r e e n or golden v a r i e gated y e l l o w - g r e e n above, yellow-brown, golden-brown or brown below, u s u a l l y yellowish-green, (1) 3 - 7  (10)  olive-green, cm  occasionally bright-green  throughout; stems  l o n g , procumbent. Branches i r r e g u l a r , v e r y w i d e l y spaced,  o f t e n a t t a i n i n g the l e n g t h o f the main stem, e s p e c i a l l y w i t h i n symmetrical p a t c h e s , o r branches s h o r t and  radially  a s c e n d i n g from p r o s t r a t e  Stems u s u a l l y l e a f y throughout, i n f r e q u e n t l y the lower 1 or 2 cm may  be  denuded.  o f o l d stems  Stem c r o s s - s e c t i o n s w i t h 2 to 3 rows, o c c a s i o n a l l y and  complete f o u r t h rows, of y e l l o w  to reddish-brown, s m a l l , t h i c k - w a l l e d  c e l l s ; medullary c e l l s l a r g e r , thinner-walled ate and  stems.  d i s c o l o r e d w i t h age;  often discolored.  Rhizoids  in-  cortical  though sometimes becoming i n c r a s s -  c e n t r a l s t r a n d weakly to s t r o n g l y d i f f e r e n t i a t e d , r e d to reddish-brown, a r i s i n g from the base o f  v e n t r a l stem l e a v e s , but o f i r r e g u l a r o c c u r r e n c e . Leaves v a r i a b l e ; c l o s e l y spaced to d i s t a n t , e s p e c i a l l y on young branchtips.  A t t i t u d e on  loosely imbricate  the stem l i t t l e d i f f e r e n t from the wet  sometimes s l i g h t e d t w i s t e d a t the apex when dry.  ovate, r a r e l y ovate o r almost o r b i c u l a r , g e n e r a l l y  gradually  to an acute, but b l u n t apex,  (0.6)  - 1.2  -.75  condition;  or s l i g h t l y spreading., Leaves s t r a i g h t , r a r e l y s l i g h t l y  secund, young branch l e a v e s Leaves b r o a d l y  to the dry  (1.25) mm  (0.8)  1.0  - 1.75  (2.0)  mm  tapering long X  wide; margin v a r i a b l e , e n t i r e , undulate o r  finely  195 d e n t i c u l a t e i n the upper h a l f , e s p e c i a l l y i n the apex, margins sometimes slightly  recurved  a t the i n s e r t i o n p o i n t ; s h a l l o w l y to deeply  narrowly or h a r d l y a t a l l d e c u r r e n t ; double w i t h  costa v a r i a b l e , usually short  s l e n d e r arms, the l o n g e s t arm  reaching midleaf  beyond, or double w i t h both arms r e a c h i n g m i d l e a f , and/or  concave;  or  and  slightly  r a r e l y s i n g l e to  midleaf  forked.  A r e o l a t i o n r a t h e r uniform; Median l e a f c e l l s rhomboid to l i n e a r flexuose, thin-walled, it  (24)  32 - 52  i s not uncommon f o r u n u s u a l l y  (74) mm  long X  (3) 5 - 6  l o n g or s h o r t c e l l s to occur  an o t h e r w i s e l o n g or s h o r t a r e o l a t i o n ; u s u a l l y l i t t l e  (8) mm  fusiform wide,  i n leaves  with  change from the median  t o the a p i c a l c e l l s , b u t i n some l e a v e s the a p i c a l c e l l s become s h o r t rhombic, to s h o r t f u s i f o r m ; m a r g i n a l c e l l s i n the upper h a l f o f the l e a f 15 to 60 long, u s u a l l y 30 to 50 um. ing  longer  c e l l s reaching  60 um  are r a r e ; b a s a l c e l l s becom-  or s h o r t e r , s l i g h t l y wider, s l i g h t l y more i n c r a s s a t e , p i t s  none, d i s c o l o r e d or not a t a l l ; a l a r c e l l s u n d i f f e r e n t i a t e d o r a few i n c r a s s a t e , u n p i t t e d quadrate to s h o r t r e c t a n g u l a r surrounding  apex, up leaves  few  to  slightly  c e l l s , hardly d i f f e r e n t  from  c e l l s , rarely discolored.  Plants autoicous; concave and  um  ecostate;  t o 1.1  mm  p e r i g o n i a l l e a v e s ovate w i t h  and  apex,  o u t e r p e r i c h a e t i a l l e a v e s ovate, t a p e r i n g to an  deeply acute  l o n g , weakly s h o r t double c o s t a t e ; i n n e r p e r i c h a e t i a l  l i n e a r l a n c e o l a t e , 2.0  absent, f a i n t  an-apiculate  to 3.5  mm  l o n g , 2 t o 4 long p l i c a e ,  double, or s i n g l e t o m i d l e a f ;  apex acute o r obtuse, c e r t a i n  c e l l s on the a b a x i a l s u r f a c e of the l e a f apex b e a r i n g by o v e r r i d i n g d i s t a l Seta 6 to 15 mm  costa  a few p a p i l l a e formed  endwalls. long, u s u a l l y 7 to 11 mm,  deep maroon, smooth, s t r a i g h t when wet  orangish-red,  reddish-brown to  v a r i o u s l y , t w i s t e d when dry; Capsule  i n the genus; Annulus of 2 to 3 rows of deciduous c e l l s .  as  196 Peristome as i n the genus;  Endostomal c i l i a  Spores smoky brown, f i n e l y p a p i l l o s e , 14.6  t o 16.6  12.5  rudimentary o r wanting;  t o 18.7  um i n diameter, u s u a l l y  um.  Hygrohypnum m o l l e can be d i s t i n g u i s h e d from o t h e r s p e c i e s i n the genus by 1.  the b r o a d l y o v a t e , concave l e a v e s  ( F i g . 45 a & b ) , which  t a p e r i n t o an a c u t e , b u t b l u n t , and o f t e n d e n t i c u l a t e apex 2.  the u n d i f f e r e n t i a t e d a l a r c e l l s  ( F i g . 46 f & h) and 3.  generally  ( F i g . 46 a & b ) , the i n n e r p e r i -  c h a e t i a l l e a v e s i n which the c e l l s on the a b a x i a l s u r f a c e o f the l e a f apex are p a p i l l o s e by means o f o v e r r i d i n g d i s t a l endwalls  ( F i g . 46 d ) .  Hygrohypnum m o l l e does e x h i b i t some v a r i a t i o n w i t h r e s p e c t to growth habit,  l e a f shape and s i z e , t o o t h i n g o f the l e a f apex and l e a f  Throughout Europe and i n some specimens  concavity.  from North America Hygrohypnum  m o l l e forms l o o s e l y woven t u f t s o r p a t c h e s i n which the stems branch a t i r r e g u l a r i n t e r v a l s and the branches q u i c k l y grow t o l e n g t h s c l o s e t o the l e n g t h s o f the stems.  On the o t h e r hand, most North  American'specimens,  e s p e c i a l l y those from Southern B r i t i s h Columbia and a d j a c e n t Washington, t i g h t l y woven, b u t s o f t patches or mats i n which b r a n c h i n g i s i r r e g u l a r  form and  both branches and stems are s h o r t and more or l e s s ascending. Leaf shape i s g e n e r a l l y b r o a d l y ovate may  e x h i b i t ovate  (Fig. 45 a & b).  ( F i g . 45 c) or almost o r b i c u l a r l e a v e s  c o n c a v i t y i s u s u a l l y more e v i d e n t i n the more o r b i c u l a r  Certain  ( F i g . 45 d & e ) .  t o 2.0 mm.  Lawton  l e a f l e n g t h v a r i e s from 1.2  t o 1.7 mm,  i n B r i t i s h Columbia and Washington,  varies  (1971) noted t h a t  o f North American p l a n t s were s l i g h t l y s m a l l e r than those o f European The p r e s e n t study has borne out those o b s e r v a t i o n s .  Leaf  leaves.  Throughout the e n t i r e range o f Hygrohypnum m o l l e the l e a f l e n g t h more o r l e s s c o n t i n u o u s l y from 0.8  specimens  leaves  specimens.  In Europe the average  whereas i n North America, and  the average l e n g t h i s 0.9  especially  t o 1.3 mm.  The  197  very largest leaves from North American plants have not been observed to exceed 1 . 5 mm. The l e a f apex i s generally described as acute, but blunt and f i n e l y denticulate (Fig. 46 a)•  However, i t may vary from  acute to obtuse and  coarsely denticulate to entire. Concavity seems to vary somewhat, r e l a t i v e to l e a f shape.  There i s a  tendency f o r large ovate leaves to be shallowly or not at a l l concave, whereas broadly ovate leaves are deeply concave.  However, the c o r r e l a t i o n  i s not absolute. The costa i s generally short and double, however, i t may vary from double with one arm reaching midleaf or beyond, double with both arms reaching midleaf or beyond or single and/or forked to midleaf or beyond. The early works of B r i d e l ( 1 8 1 2 , 1 8 2 7 ) , Sprengel (1827) and Hampe (1837)  noted a s i m i l a r i t y between Hygrohypnum molle and H. alpestre.  Further, Schimper.. (1853) i l l u s t r a t e d H_. alpestre with a drawing of H. molle. Both of these species have s t r a i g h t , concave leaves that are loosely imbricated or with the shoots sometimes julaceous.  At the apex of some  stems and branches the young leaves are so crowded that the stem or branch apex appears obtuse.  Though these characters r e f l e c t a common bond within  the genus, they mask very fundamental differences at the species l e v e l . The following chart w i l l i l l u s t r a t e the important differences.  198 H. molle  H. alpestre  Leaf shape  Broadly ovate  Oblong or oblong lanceolate .  Leaf apex  Acute but blunt, plane  Apiculate and reflexed  Concavity  Shallowly to deeply so  Deeply so, often quite boat shaped i n the apex.  Alar c e l l s  E s s e n t i a l l y undifferentiated.  Large group of quadrate c e l l s  Median leaf c e l l s  Usually 32 to' 52 um long  Usually 65 to 90 um long  Inner perichaet i a l leaves  Apex coarsely denticulate and p a p i l l o s e  Apex long tapering, entire, smooth.  I l l founded concepts of Hygrohypnum molle and H_. duriusculum have long caused confusion between these two clear-cut taxa.  The following chart w i l l  perhaps c l a r i f y , the differences.  H_. molle  H-. duriusculum  Leaf Shape  Broadly ovate, gradually tapering to an acute but blunt t i p .  O b l o n g - e l l i p t i c to broadly ovate, more abruptly tapered to an obtuse, broadly acute or apiculate apex.  Alar c e l l s  Undifferentiated  Well developed groups of quadrate, incrassate, discolored cells.  Stem cross section  2 to 3 rows of small incrassate c o r t i c a l cells  3 to 4 rows of small, incrassate c o r t i c a l c e l l s .  Inner perichaet i a l leaves  Apical cells- papillose by d i s t a l l y overriding endwalls.  Apical c e l l s smooth  Inner peristome  Endostome smooth, c i l i a rudimentary.  Endostome f i n e l y p a p i l l o s e with well developed c i l i a .  199 Hygrohypnum molle and H. cochlearifolium have a number of common features.  Both plants are quite s o f t , although H. cochlearifolium i s the  softer of the two.  The  leaves of both are generally loosely  to s l i g h t l y spreading (Fig. 45 f & g with F i g . 40 1).  imbricate  The leaves of H.  molle are generally broadly ovate and shallowly concave, but certain specimens may  exhibit broader leaves with a deeper concavity  (Fig. 45 d & e)  that are more comparable with H_. cochlearifolium (Fig. 13 d) .  Both taxa  frequently exhibit .a s i m i l a r pattern of golden-brown mottled coloration. The  costa morphology i s s i m i l a r and has already been pointed out i n the  discussion of H.  cochlearifolium.  The two taxa may  be distinguished from one another by the  following  features: H. molle  H_. cochlearifolium  Leaf Shape  Broadly ovate  Mostly o b l o n g - e l l i p t i c to almost o r b i c u l a r  Leaf Length  0.8  0.5  Leaf Margins  Plane; denticulate to entire  Usually recurved, sometimes plane; entire or weakly crenulate  Leaf Apex  Gradually tapering to an acute but blunt point  Apex broadly rounded obtuse  Leaf Concavity  Shallow  Very deep to  Alar C e l l s  Undifferentiated  Undifferentiated or a few quadrate c e l l s  Endostome  C i l i a rudimentary or wanting  C i l i a 2 to 3  Perichaetial  Apex coarsely denticulate; margins plane; apical c e l l s p a p i l l o s e  Apex e n t i r e ; margins recurved; apical c e l l s smooth, rarely 1 or 2 p a p i l l a e  to 2.0  mm  to 1.5  mm  cochleariform  200 As an autoicous species the presence of antheridia and archegonia on the same i n d i v i d u a l would t h e o r e t i c a l l y enable H. molle to freely reproduce sexually.  I f the number of herbarium specimens bearing sporophytes  i s a reasonable measure of a species fecundity, then i t appears that Hygrohypnum molle reproduces sexually very infrequently.  Barely ten percent of  the several hundred specimens studied bear sporophytes and the majority of f e r t i l e specimens are from Western North America. At the inception of this study four subspecific taxa were recognized for Hygrohypnum molle (Wijk et a l . , 1962).  Hygrohypnum molle ssp. b e s t i i  (Reri. et Bryhn) Broth, i s treated here as a species. cribed H_. molle var. japonicum.  Sakurai (1932) des-  Examination of the holotype from the Makino  Herbarium has revealed that the plant i s only a form of Hygrohypnum ochraceum. Hygrohypnum molle var. pyreniacum  (Ren.) Podp. was described by Renauld  i n two s l i g h t l y varying descriptions i n the Academe de Geographie B u l l e t i n Vol. 10 and Revue Bryolbgique V o l . 28 i n 1901. upon.the s i m i l a r i t y of the plant to Hypnum b e s t i i .  Botanique  Renauld remarked  A specimen whose label  bears the data given by Renauld for the type specimen i s at NY.  I t seems  l i k e l y that this specimen i s a duplicate of the type that Renauld may have sent to J . M. Holzinger. Examination of the plant reveals that i t i s a moderately large specimen of H_. molle.  I t does not have the very long marginal  leaf c e l l s that would otherwise a l i g n i t with H. b e s t i i . Lorentz (1864) described Hypnum molle var. schimperianum.  At G (Sheet  3079/247) there i s a specimen agreeing with the c o l l e c t i n g information i n d i cated by Lorentz.  A careful examination of this specimen reveals that i t i s  not s u f f i c i e n t l y d i f f e r e n t from other specimens of Hygrohypnum molle to warrant taxonomic recognition. Although the p r i n c i p l e of p r i o r i t y credits Hedwig.with the authorship of  201 Hygrohypnum molle (Hedw.) Loesk., i t i s clear that he based his concept on Hypnum molle Dicks.  Similarly, Schimper (1853) based his concept of  Limnobium molle, i n part, on Dickson's concept.  The type l o c a l i t y as given  by Hedwig (1801) i s "ad rivulorum ripas i n alpibus Scotius." Schimper (1876) narrowed the e a r l i e r heterogeneous Hypnum dilatatum and H_. alpinum.  concept of L. molle Schimp. by extracting The more r e s t r i c t i v e concept was again  equated with that of Hypnum molle Dicks, and several specimens were c i t e d as the basis for the concept.  Among the specimens was one collected by G. E.  Hunt from Ben Mac Dhui i n Scotland.  At S-PA there are two specimens bearing  these data and both specimens are equally representative of the concept of Hygrohypnum molle as perceived i n the present study. One problem i n the t y p i f i c a t i o n of Hygrohypnum molle l i e s i n the fact that the only specimen bearing the name Hypnum; mo M e Dicks, i n the Hedwig^ 1  Schwaegrichen herbarium at G. i s a mixture of Hygrohypnum luridum (Hedw.) Jenn. and H. duriusculum (De Not.) Jamieson.  That a specimen i s i n the 'Hedr-  wig herbarium does not necessarily mean that Hedwig ever saw i t .  A comparison  of the broadly ovate leaves of the troublesome specimen with Hedwig's type description of the leaves of Hypnum molle as " f o l i a ovato-lanceolata" and an examination of Tab. LXX c l e a r l y reveals a lack of agreement. anomalous specimen.from  Therefore, this  the Hedwig herbarium i s rejected as representative of  Hygrohypnum molle. In the absence of any Dickson material or appropriate Hedwig material and based on a'careful study of several hundred specimens bearing the names Hypnum molle and H_. duriusculum I select the following specimen, to serve as the neotype of Hygrohypnum molle (Hedw.) Loesk.: Scotland, Ben Mac Dhui, by streamlets from perpetual snow. Leg. G. E. Hunt, July 1871  202 ex h e r b . G. S t a b l e r & ex h e r b a r i o S. O. At  S-PA.  Lindberg.  203 F i g . 45 a - g.  V a r i a t i o n i n the l e a f shape and the h a b i t o f the shoots  o f Hygrohypnum molle. a - e.  Foliage leaves,  f - g.  The h a b i t of moist shoots.  Scale: a - e;  \  I  1mm  I  f;  The shoot i s approximately  0.6 cm long  g;  The shoot i s approximately  2 cm long  204  46 a - h. C e l l u l a r d e t a i l o f the f o l i a g e and p e r i c h a e t i a l leaves o f Hygrohypnum molle. a - b. Foliage l e a f apices c & e. Marginal l e a f c e l l s d.  P e r i c h a e t i a l l e a f apex  f S h. g.  Alar c e l l s  Median l e a f c e l l s  Scale: a - b;  I  100um  I  c, e and g,-  1  • 100um  d, f and h;  I  100um  I  \  206  207  4-  F i g . 47 . Hygrohypnum molle  208 E x s i c c a t i Examined A l l e n , Mosses o f the Cascade Mtns., Washington. # 142 as H. a r c t i c u m . (MIN, NY, TENN, USA, DUKE) Macoun, Canadian Mosses # 393 as H. pseudo-arcticum. (CANM) # 394 as H_. a r c t i c u m . (USA) # 398 as Hypnum a r c t i c u m . (LE) Bauer, 'Musci e u r o p a e i e x s i c c a t i # 649 as Hygrohypnum schimperianum. (S-PA, BRNM) # 1663 as H_. m o l l e v a r . schimperianum. (CANM, S-PA) Kerner, F l o r a E x s i c c a t a A u s t r o Hungarica # 1923 as Hypnum schimperianum. (BRNM, MIN, S-PA) # 1924. (BRNM, G, L E , MIN, MO, NY, UC, BP', C) L i s o w s k i , Bryotheca P b l o n i c a F a s c . XV # 418 as H. v i r i d u l u m . (CANM) F a s c . XV # 419. (CANM, LE) F a s c . XXV # 669. (CANM) F a s c . XXV # 668. (LE) F a s c . XXXI # 824. (CANM, NICH) Museo H i s t Natur Vindobonensi, Cryptogamae e x s i c c a t i # 4260 as'H. m o l l e . (G, L E , NY) # 4362 as H. m o l l e . (G, LE) Museum Botanicum U n i v e r s i t a t i s C l u j , F l o r a Romaniae E x s i c c a t i # 1411 as H_. m o l l e v a r . schimperianum. (MO, S-PA) Zmuda, Bryotheca P o l o n i c a # 150. (S-PA) :  S e l e c t e d Specimens Examined Canada British  Columbia  st Seymour P r o v i n c i a l Park, 1 Punp Peak; S c h o f i e l d 15968 as H. a l p e s t r e . (CANM) G a r i b a l d i P r o v i n c i a l Park, W h i s t l e r S k i Bowl; Jamieson 5486. (UBC) Vancouver I s l a n d , S t r a t h c o n a P r o v i n c i a l Park, Burman Lake; Boas 254. (UBC) G o l d Ranges, Macoun 10 Aug. 1889 as H. pseudo-arcticum i n . Canadian Mosses # 393. (CANM) Rocky Mountains, Hector; Macoun 8 Aug. 1904 as H. a r c t i c u m i n Canadian Mosses # 398. (CANM) E a g l e Pass Mtns., near R e v e l s t o k e ; T a y l o r 21 Aug. 1921 as H. dilatatum. (MICH, USA) United States Washington Mt. R a n i e r N a t i o n a l Park, Mt. R a n i e r ; A l l e n 119^as H_. a r c t i c u m i n Mosses o f the Cascade; Mtns., Washington # 142. (NY, TENN, USA, MIN) Oregon' Hood R i v e r Co., Cloud Capp Inn; F r y e 29 Aug. 1907. (WTU) Mt. Hood; R o e l l 1140 & 1411. (NY) Mt.  209 California Mono Co., H. M. H a l l N a t u r a l Area, S p u l l e r Lake; Catchende 47138. (MICH) E l d o r a d o Co., Echo Lake; Conard 5 Sept. 1947. (MICH) Montana F l a t h e a d Co., S p e r r y G l a c i e r T r a i l , A k a i y a n F a l l s ; Hermann 20757. (NY, WTU) Colorado L a r i m e r Co., Rocky Mountain N a t i o n a l Park, Longs Peak: K i e n e r 4113. (CANM, MICH) Alaska Juneau I c e f i e l d , Taku Anunatak; Ward 8-27-49-28 as H. subeugyrium. (MIN) Great B r i t a i n Scotland Ben Mac Dhui; Fergusson J u l y 1873. (S-PA) Ben N e v i s ; Boner 1870. (NY) Ben N e v i s ; i n the Red Burn; Dixon 11 J u l y 1898. (NY) Norway J o t u u k j e l d e n e supra hosptium Gjendeboden; K a u r i n Aug. 1891. (S'-PA) Finnmarken, aunt Tanen Burkelium; K a u r i n J u l y 1895. (s-PA) N o r e f j e l d ; Bryhn J u l y 1900. (S-PA) Knudshoe; Bryhn Aug. 1885. (S-PA) E f f e s vagen V i d a s o e t e r G r o s l l i ; Jensen 1 J u l y 1937. (S-PA) Opdal, i n monte Snehetta;. O l s s o n 31 J u l y 1885. (LE, NY) Sweden Harkedalen, p a r Tannas; E i n a n d e r 3 - 4 Aug. 1891. (S-PA) Jamtland, K a i l s sn L i l l a n j e s k u t a n ; H a k e l i e r 13 J u l y 1966. (S-PA) Brahnkyika; C l i v e 1857. (S-PA) Finland Lapp, r o s s i t . , K i p i n a ; Angstrom. (S-PA) Lapponia imandrau, Umptek; Kihlman 30 J u l y 1892. (S-PA) Kemi Lapp. K o l a r i ; K o t i l a i n e n 11 Aug. 1927. (S-PA) Lapponia murr., L i t s a ; B r o t h e r u s 3 June 1887. (NY) S o v i e t Union Lapponia murmanica, S i t s a , K o l a ; Brotherus Aug. 1887. (C, S-PA) Germany M a t r e i , A l g a u , S i e g e r l a n d h u s s e ; Leg. ? 8 Aug. 1929. (BRNM, a c c e s i o n number 08719) France Chamoix; Payot.1854. (G, 3079/210) Mt. B l a n c ; Leg. ?. (G, 3079/208). V a l l e de B a r b a r i n e , s u r l e s Roches S e l i c e u s e s a Fontanabran; Bernet 13 Aout 1903. (G, 3079/259) Switzerland La F i b b i a , S t . G o t t h a r d ; Weber 7 Aug. 1881. (G, 3079/228) V a l a i s , Grand S t . Bernard, Combe 'des Morts; Rhodes 1705 as H. m o l l e . (NY) ' Austria Tirol,  I n n e r v i l l g a r t e n ; Ganders 15 Sept.  1881.  1720/89 (B, —1972""^  210  S a l i s b u r g i a , Montes L a n s c h i t z b a r prope L e s s a c h ; B r e i d l e r as F l o r a E x s i c c a t a A u s t r o - H u n g a r i c a # 1924. (G, 3079/205) S t e i e r m a r k j ^ S , j n g s d o r f e r ; B r e i d l e r 25 J u l y 1887.  B '  . 1972  Italy Genoa, rupes F o n a l i s ; L o r e n t z 12 Aug. 1864. (S-PA) Czechoslovakia Vysoke T a t r y , F u r k o t a ; Smarda 3 J u l y 1957. (BRNM) ' Poland Montes A l t i , v a l l e D o l i n a ; L i s o w s k i 27 Aug. 1956 as B r y o t h e c a P o l o n i c a # 419. (CANM) Romania T r a n s y l v a n i a , d i s t r . Hunedosra, I n montes R e t e z a l ; P e t e r f i 24 J u l y 1914 i n F l o r a Romaniae E x s i c c a t i # 1411. (S-PA).  211 Hygrohypnum s t y r i a c u m  (Limpr.) Loesk., Nat.  Hypnum (Limnobium) s t y r i a c u m Eurhynchium Cad.  P f l . 1(3):1039.  Limp., F l o r a  (Jena) 65:201.  (Pseudo-Rhyncostegium) s t y r i a c u m  Rec.  S c i . 6:22.  Limnobium s t y r i a c u m  1909. 1882.  (Limpr.) Kindb.  1894.  (Limpr.) Roth, Eur.  Laubm. 2:638. 56 f .  2.  1905. Names from the  l i t e r a t u r e but not a v a i l a b l e f o r assessment.  Hypnum s t i r i a c u m Limpr. ex. 1900.  nom,  Par.,  i l l e g . orthogr.  Ind. B r y o l . S u p p l . pro H_.  P l a n t s v a r i o u s l y s o f t to s l i g h t l y r i g i d , p a t c h e s or mats.  213.  styriacum.  i n l o o s e l y to t i g h t l y woven  Color u s u a l l y d u l l yellow-green with a rusty  mottling,  l e s s o f t e n a d i r t y brownish y e l l o w - g r e e n or a u n i f o r m d u l l green. 1-4 in  (6) cm  the  l o n g ; v a r i o u s l y f o l i o s e , e i t h e r f o l i o s e throughout or denuded  lower p o r t i o n s , e s p e c i a l l y i n the v e r y  t r a t e , sometimes somewhat a s c e n d i n g . lose.  Stem c r o s s - s e c t i o n s  tical  Stems  long stems; stems mostly  Branching i r r e g u l a r .  Sparsely  pros-  radicu-  r e v e a l i n g 2 to 3 rows o f s m a l l , t h i c k - w a l l e d  c e l l s ; m e d u l l a r y c e l l s g r a d u a l l y becoming l a r g e r toward the  o f t e n becoming t h i c k e r - w a l l e d and d i s c o l o r e d w i t h age;  cor-  middle,  c e n t r a l strand  well  developed. Leaves v a r i o u s l y d i s p o s e d  along  crowded, crowding o f t e n more e v i d e n t A t t i t u d e v a r i a b l e from the wet spreading leaves various  the stem, d e c i d e l y d i s t a n t to  near the stem and branches a p i c i e s .  to dry c o n d i t i o n ; when wet  a l l around the stem, l e s s o f t e n s l i g h t l y  the l e a v e s  spreading,  weakly i m b r i c a t e d ,  are  f a l c a t e ; lipon d r y i n g  s h r i n k l a t e r a l l y to a marked degree, o f t e n concomitantly a t t i t u d e s , e i t h e r spreading,  slightly  e s p e c i a l l y so i n s m a l l e r  the  twisting i n branches,  p a r t i c u l a r l y near stem and branch a p i c i e s or  212 f a l c a t e ; Leaf  shape q u i t e uniform, ovate w i t h  acuminate apex, u s u a l l y s t r a i g h t , o r s l i g h t l y  an a b r u p t l y o r g r a d u a l l y f a l c a t e , i n which case the  apex u s u a l l y t a p e r s g r a d u a l l y ; apex i s sometimes s h a r p l y r e f l e x e d ; Leaves (0.7)  1.0 - 1.75 (2.0) mm  e n t i r e ; conspicuously  l o n g X (0.3) 0.5 - 1.0 )1.1) mm wide;,.margins  broadly  concave when wet; very weakly d e c u r r e n t  or •  not a t a l l ; c o r t i c a l  stem c e l l s o f t e n adhere t o the l e a f base; c o s t a  and  double, s i n g l e and/or f o r k e d , o r i n f r e q u e n t l y s i n g l e  t h i c k , short'and  strong  to above the m i d d l e . A r e o l a t i o n q u i t e uniform, median c e l l s g e n e r a l l y s h o r t , v a r i o u s l y rhombic, s h o r t f u s i f o r m or b a c i l l i f o r m ; t h i c k - w a l l e d l o n g X (4) 5 - 7  (9) um wide; l i t t l e  toward the base, those c e l l s n e a r e r  (18) 28 - 40 (50) um  changed toward the apex; c e l l s  varying  the c o s t a becoming l o n g e r and narrower  toward the base, those c e l l s near the margin becoming s h o r t e r and broader, these t e n d e n c i e s  a r e g e n e r a l , however a l l c e l l s become t h i c k e r - w a l l e d toward  and w i t h i n the l e a f base; a l a r c e l l s v a r i a b l e , b u t never s h a r p l y undifferentiated or recognizable gular c e l l s grading  delimited,  as a s m a l l group o f quadrate o r s h o r t  rectan-  i m p e r c e p t a b l e i n t o o t h e r b a s a l o r lower median c e l l s .  P l a n t s pseudo-paroicous; p e r i g o n i a and p e r i c h a e t i a c l u s t e r e d i n t o aggrega t i o n s and subtended by common b r a c t s ; a s i n g l e p e r i g o n i u m i s s i t u a t e d immedi a t e l y beside  a s i n g l e perichaetium  o r between 2 t o 4 p e r i c h a e t i a , i n a l l cases  the p e r i g o n i a l - p e r i c h a e t i a l complex i s e n c l o s e d  within 2 to 3 small bracts;  the e n t i r e complex i s borne i n the a x i l o f a v e g e t a t i v e are b r o a d l y - o v a t e  with  a rounded or 1 t o 3 lobed apex; p e r i g o n i a l l e a v e s a r e  ovate t o o v a t e - l a n c e o l a t e , leaves, ecostate, e n t i r e or very tial  leaves  l e a f ; common b r a c t s  the apex i s o f t e n a b r u p t l y acuminate l i k e the f o l i a g e  a r e o l a t i o n becoming r a t h e r i n c r a s s a t e and d i s t o r t e d , margins  finely  differing  s e r r u l a t e , 0.4 t o 0.6 mm l o n g ; o u t e r and i n n e r in little  perichae-  but s i z e , l i n e a r - l a n c e o l a t e , variously  ecostate  213 or f a i n t l y  s i n g l e c o s t a e , deeply p l i c a t e - s u l c a t e , margins e n t i r e o r  irregularly  toothed,  l e a v e s up to 2 mm  S e t a 9 - t o 21 mm or s l i g h t l y  long.  long, v a r i o u s l y red to reddish-yellow,  i n c l i n e d when wet,  c o n t o r t e d upon d r y i n g .  very  smooth, s t r a i g h t  t w i s t i n g to the l e f t and becoming v a r i o u s l y  Annulus deciduous, o f 1 or 2 rows o f c e l l s .  Capsule  as i n the genus. Peristome double; as per  the genus; Endostome w i t h  are sometimes p o o r l y developed. 23 um  Spores smoky y e l l o w ,  styriacum  f i n e l y p a p i l l o s e , 12  (1882) the most unique f e a t u r e e x h i b i t e d by  i s the pseudo-paroicous s e x u a l i t y .  productive  to  s t r u c t u r e s the s p e c i e s may  be  d i f f e r e n t i a t i o n and The  r e c o g n i z e d by a combination of  the s t r u c t u r e o f the  l e a v e s are b a s i c a l l y ovate,  to an acuminate p o i n t or e x i s t as a few s h o r t and  o f the l e a f apex, the  The  short rectangular c e l l s  Although Hygrohypnum s t y r i a c u m i s r e g u l a r l y confused  w i t h H.  ( F i g . 49  f.&  i s regarded  g).  The  (Limpr.) B r o t h ,  luridum  tapered  and  costa i s  In l e a v e s of compar-  i s s t o u t e r than those of H_.  i t o f t e n i s mistaken f o r H_. norvegicum. (Limpr.) B r o t h ,  abruptly  a l a r c e l l s are u n d i f f e r e n t i a t e d  double or s i n g l e to s l i g h t l y beyond m i d l e a f .  a b l e s i z e the c o s t a o f H_, s t y r i a c u m  alar  costa.  but the apex i s normally  ( F i g . 48 a - h ) .  H.  In the absence o f s e x u a l r e -  c h a r a c t e r s which i n c l u d e l e a f shape, the nature  and  which  i n diameter.  As noted by L i m p r i c h t  it  2 to 3 c i l i a ,  luridum.  is- a d i s t i n c t i v e  species,  i n P a c i f i c C o a s t a l North America  Grout  (1931) remarked t h a t "H_.  by L i m p r i c h t h i m s e l f as a s u b s p e c i e s  styriacum  of H . palustre  I have been unable to d i f f e r e n t i a t e American p l a n t s r e f e r r e d to i t from  forms o f p a l u s t r e . " taxa.  I t i s obvious t h a t Grout c o u l d not d i s t i n g u i s h the  two  However, I have seen ho l i t e r a t u r e c o r r o b o r a t i n g the a s s e r t i o n a s c r i b e d  to L i m p r i c h t .  214 When sex organs a r e a v a i l a b l e f o r examination  the unique p e r i g o n i a l -  p e r i c h a e t i a l complex o f Hygrohypnum s t y r i a c u m i s c l e a r l y d i f f e r e n t from the a u t o i c o u s c o n d i t i o n p r e s e n t i n H. l u r i d u m .  Further, c a r e f u l d i s s e c t i o n of  o p e r c u l a t e c a p s u l e s o f H_. s t y r i a c u m w i l l r e v e a l an annulus a t the base o f the peristome.  Hygrohypnum luridum does n o t possess  an  In the absence o f sex organs and sporophytes H. l u r i d u m may be d i s t i n g u i s h e d through acters presented  annulus.  Hygrohypnum s t y r i a c u m and  the c o n t r a s t i n g gametophytic  char-  i n the f o l l o w i n g c h a r t .  H_. s t y r i a c u m  H_. l u r i d u m  Leaf Shape  Ovate w i t h an a b r u p t l y t a p e r e d acumen.  U s u a l l y l a n c e o l a t e or oblong lanceolate with a s l i g h t a p i c u l u s .  Alar  L i t t l e d i f f e r e n t from other b a s a l c e l l s or a few quadrate o r s h o r t rectangular c e l l s .  W e l l d e f i n e d group o f quadrate to s h o r t r e c t a n g u l a r c e l l s .  Median L e a f Cells  U s u a l l y 28 - 40 um  U s u a l l y 30 - 55 um.  Costa  Stout  Slender  Cells  The  a b r u p t l y narrowed acumen i n H_. s t y r i a c u m v a r i e s i n a number o f ways.  In some cases i t ' i s (Fig.  squarrose  ( F i g . 48 j ) , i n o t h e r s i t may.be s t r a i g h t  48 i ) , o r i t m a y b e d e c i d e d l y tur.ned t o one s i d e i m p a r t i n g a s l i g h t l y  f a l c a t e appearance t o the l e a v e s  ( F i g . 48 j & k ) .  In some cases'the apex i s  more g r a d u a l l y t a p e r e d and such l e a v e s a r e u s u a l l y d e c i d e d l y f a l c a t e . g e n e r a l , however, t h e l e a v e s a r e u s u a l l y s t r a i g h t . tends  In  Upon d r y i n g , t h e apex  t o r o l l up and t w i s t , thus a c q u i r i n g a f i l i f o r m appearance  ( F i g . 48 i ) .  Specimens o f Hygrohypnum s t y r i a c u m e x h i b i t i n g s l i g h t l y o r d e c i d e d l y c a t e l e a v e s may be confused w i t h the f a l c a t e forms o f H. l u r i d u m . forms o f H. l u r i d u m the a l a r c e l l s a r e f r e q u e n t l y excavated, observed  i n H. s t y r i a c u m .  fal-  In f a l c a t e  a c o n d i t i o n never  215 Hygrohypnum  norvegicum i s b e s t d i s t i n g u i s h e d from H. s t y r i a c u m  the b a s i s o f i t s ovate l e a v e s , which have acute  on  apicies.  The a t t i t u d e o f the l e a v e s upon the stem and the l e n g t h o f the i n t e r nodes may v a r y .  In g e n e r a l , m o i s t l e a v e s are more o r l e s s s t r a i g h t and  loosely imbricate close together. leaves.  ( F i g . 48 i ) . Often  In the i m b r i c a t e d c o n d i t i o n the l e a v e s are  the i n t e r n o d e s  Those specimens w i t h  are l o n g i n specimens'with  f a l c a t e l e a v e s are a l s o g e n e r a l l y  spreading imbricate  ( F i g . 48 j & k ) . A number o f e a r l i e r authors o f Hygrohypnum  styriacum  reported observations  on c e r t a i n f e a t u r e s  f o r which t h i s study has r e v e a l e d g r e a t e r  i t y than p r e v i o u s l y known o r t h a t o r a r e d i f f e r e n t from p r e s e n t Limpricht  (1904) noted t h a t the common b r a c t s subtending  c h a e t i a l complex were 2 t o 3 l o b e d . 3 lobed. broadly Nyholm  Limpricht costate.  observations.  the p e r i g o n i a l - p e r i -  I have found them t o vary  from e n t i r e to  (1904) a l s o noted t h a t the i n n e r p e r i c h a e t i a l l e a v e s  Roth (1905), on the o t h e r hand, s a i d they were  (1965) showed i n an i l l u s t r a t i o n  and b r o a d l y  costate.  variabil-  were  ecostate.  ( f i g . 302) t h a t they were s t r o n g l y  T h i s study has r e v e a l e d t h a t p e r i g o n i a l l e a v e s and  o u t e r p e r i c h a e t i a l l e a v e s are e c o s t a t e , whereas middle and i n n e r p e r i c h a e t i a l l e a v e s may be e c o s t a t e or weakly t o s t r o n g l y s i n g l e c o s t a t e . d e s c r i b e d the stems o f H. s t y r i a c u m  as f i l i f o r m .  be no more s l e n d e r than o t h e r Hygrohypna.  (1904) and Roth (1905)  d e s c r i b e d the i n n e r p e r i s t o m e t e e t h as not b e i n g c r a c k e d  spores  by B r e i d l e r as the b a s i s f o r the s p e c i e s . type'from among these  specimens.  a number o f t h e i r d u p l i c a t e s .  along the m i d l i n e .  I  Monkemeyer' (1927) d e s c r i b e d the  as smooth w a l l e d , when, i n f a c t , they  In the o r i g i n a l d e s c r i p t i o n L i m p r i c h t  (1904)  I have found the s p e c i e s t o  Limpricht  have found them to be i r r e g u l a r l y c r a c k e d .  Limpricht  are very d e n s e l y p a p i l l o s e .  (1882) c i t e d f o u r specimens c o l l e c t e d However,  he d i d not d e s i g n a t e  a holo-  I have seen a l l o f the c i t e d specimens and  Of these  specimens I have s e l e c t e d the f o l l o w i n g  216 specimen from the L i m p r i c h t herbarium a t BP as b e i n g most r e p r e s e n t a t i v e o f the s p e c i e s and d e s i g n a t e i t as the l e c t o t y p e J L e g . J . B r e i d l e r  1 Sept  1880, Abhang am Schiedek d i e Patzenalm b e i Schladming c a 2800 m.  There i s  a s l i g h t d i s c r e p a n c y between the c i t a t i o n o f t h i s description  and the data borne on the l a b e l .  specimen i n the  The c i t a t i o n  i n the  original description  says "Nordabhand des.'......" and makes no mention o f the p r o x i m i t y t o Schladming.  217 F i g . 48 a - k.  V a r i a t i o n i n l e a f shape and shoot habit of Hygrohypnum  styriacum. a• r h.  V a r i a t i o n i n l e a f shape.  i - k.  V a r i a t i o n i n the h a b i t of the shoot i n the moist c o n d i t i o n .  Scale: a  T  h;  i - k;  I  1mm  |  Each shoot i s approximately 1 cm long  218  219  c i  '  F i g . 49 a - g.  C e l l u l a r d e t a i l o f the f o l i a g e leaves o f Hygrohypnum  styriacum. a.  Leaf apex.  b and d. Median l e a f c e l l s , c and e.  Marginal l e a f c e l l s ,  f and g.  Alar c e l l s .  Scale: a,  f and  g;  |  100um  I  220  221  222 E x s i c c a t i Examined Bauer, Musci europ. e t amer. e x s i c c a t i # 1790. (BRNM, S-PA) Lisowski, Bryotheca P o l o n i c a F a s c . XXIV, # 636. (BP, CANM, S-PA, LE, NIGH) F a s c . XXV, # 666. (BP, CANM, S-PA, LE) F a s c . XXXI, # 823. (BP, CANM, S-PA, LE, NI<C-H) S e l e c t e d Specimens Examined Canada B r i t i s h Columbia G a r i b a l d i P r o v i n c i a l Park, S e n t i n a l G l a c i e r a t t h e E . end of G a r i b a l d i L; W. B. S c h o f i e l d 32918. as Hygrohypnum. l u r i d u m . (UBC) G a r i b a l d i P r o v i n c i a l Park, W h i s t l e r Mtn. S k i Area; D. W. Jamieson # 5472. (UBC) Vancouver I s . , S t r a t h c o n a P r o v i n c i a l Park, Golden Hinde; Boas 297 as Hygrohypnum l u r i d u m . (UBC) S e l k i r k Mtns., Z i n c t o n , Rambler Mine; F. A. MacFadden 3880 as Hygrohypnum p a l u s t r e (UBC) K i t c h e n e r Krag; Boas 1960 as Hygrohypnum p a l u s t r e . (UBC) Z i n c t o n , Lucky J i m Mine; F. A. MacFadden 3864 as Hygrohypnum p a l u s t r e . (UBC) Wilmer, near Windemere, P a r a d i s e Mine; F. A. MacFadden 3878 as Hygrohypnum p a l u s t r e . (UBC) Sandon D i s t r i c t , Rambler Mine; F. A MacFadden 11 Sept... 1927 as Hygrohypnum p a l u s t r e . (MO, TENN) E a s t e r n P a c i f i c Coast D i s t r i c t , Skagway (Alaska Quadrangle) Along the Haines Hwy, 83 m i l e s NNW o f Haines; F. J . Hermann 21843 As H_. norvegicum. (USA) P a r a d i s e Mine; F. A MacFadden 1 Aug. 1928 as Hygrohypnum p a l u s t r e (MO, UC) Alberta J a s p e r N a t i o n a l Park, Tonquin V a l l e y ; F. A. MacFadden 3875 as Hygrohypnum p a l u s t r e (UBC) Bow R i v e r Watershed, S. t r i b u t a r y o f Three I s l e Creek. W. o f Upper Kamanaskis L.; B i r d & Glenn 13826 as Hygrohypnum luridum. (UAC & UBC) J a s p e r N a t i o n a l Park, W i l c o x F a l l s ; E . Whitehouse 25745 as Hygrohypnum p a l u s t r e . (USA) J a s p e r N a t i o n a l Park, N. o f Sunwapta Pass Campsite; O. D. B i r d 5232. as Hygrohypnum l u r i d u m (UAC) United States California ?; K e l l o g . (USA) Water courses near Summit; H. B o l a n d e r i 1864 - 1880. as Hypnum s t y r i a c u m (NY) T r i n i t y Co., B u l l a r d ' s B a s i n ; D. H. N o r r i s 9287. as Hygrohypnum l u r i d u m (HSC) Idaho C u s t e r Co., A d a i r Creek; F. A. MacFadden 7 Aug. 1941. as Hypnum p a l u s t r e (CANM, MICH, NY, TENN, TRTC) Lemhi Co., 8 m i l e s N. o f ghost town o f G i b b o n s v i l l e ; T. C. F r y e 1 Sept. 1929 as Hygrohypnum ochraceum (WTU)  223 Montana G l a c i e r N a t i o n a l Park, Hidden L. overlook-Logan Pass; F. J . Hermann 18094 as Hygrohypnum l u r i d u m (TENN) G l a c i e r N a t i o n a l Park, Hanging V a l l e y - L o g a n Pass; F. J . Hermann 18088 as Hygrohypnum l u r i d u m (NY) Between f o r k s o f C u t Bank Creek; R. S. W i l l i a m s 26 J u l y 1897 as Hypnum s t y r i a c u m (C, MO, NY) Utah On stream above S a l t Lake; P. A. Evans 15 Aug. 1924 as Hygrohypnum p a l u s t r e (NY) Wyoming B a t t l e Lake Mth.; A. Nelson 4202. (NY) Norway S o r f o l d e n ; M. K o t i l a i n e n 25 J u l y 1922 as Hygrohypnum s t y r i a c u m (C, S-PA) Sweden L u l e Lappmark, Roveguaare, K v i k k j o k k ; Hulphers 12 J u l y 1937 as Hygrohypnum v i r i d u l u m (S-PA) Austria T i r o l , Zunig b e i W i n d i s c h - M a t r e i o b e r h a l b ?; J . Baumgartner 16 Sept 1905 as Musci europ. e t a m e r . e x s i c c a t i # 1790. (BRNM, S-PA) S t e i e r m a r k , Abhang am Schiedek d i ^ Patzenalm b e i Schladming, ca. 2000 m; J . B r e i d l e r 1 Sept 1880. LECTOTYPE. (BP) Poland Montes T a t r i A l t i , v a l l i s D o l i n a ; L i s o w s k i 27 Aug. 1956 as B r y o t h e c a P o l o n i c a F a s c ' XXIV, # 636. (BP, CANM, S-PA)  224 Hygrohypnum l u r i d u m Neotype:  (Hedw.) Jenn., Man. Moss. West. P e n n s y l . 287.  " I n S u e c i a ad t r a b e s m o l e n d i n i c a " ; Swartz  1913.  (G, sheet  3079/156) Hypnum l u r i d u m Hedw., Spec. Muse. 291.  1801. (Lectotype a t  G, sheet 3079/156) Hypnum p a l u s t r e Huds. ex B r i d . , Musci Rec. 2(2):117. Hypnum molendinarium 1805.(Lectotype  1801.  Lam e t Cand., F l . F r a n c . Ed 2, 2:358. a t G)  Hypnum l u r i d u m v a r . ferrugineum B r i d . , Muse, Rec. Supp. 2:231. 1812. Hypnum neglectum  B r i d . , Muse. Rec. Supp. 2:120.  1812.  Hypnum s u b s p h a e r i c a r p o n S c h l e i c h . ex B r i d . , Muse. Rec. Supp. 2: 232.  1812.  Hypnum a q u a t i l e Mart.,  F l . Crypt. E r l . ,  Hypnum s p h a e r i c a r p o n Spreng.,  S y s t . Veg. 4(1):201.  Hypnum p a l u s t r e v a r . a q u a t i l i s 640.  19. I f . 2 A - K. 1817. 1827.  (Mart.) B r i d . , B r y o l . U n i v . 2:  1827.  Hypnum p a l u s t r e v a r . f e r r u g i n e u s (Brid.) B r i d . , B r y o l . U n i v . ••• 2:641.  1827.  Hypnum p a l u s t r e v a r . l u r i d u m  (Hedw.) Hamp., F l o r a  (Jena)  20:274.  1837, Hypnum p a l u s t r e v a r . s u b s p h a e r i c a r p o n Schwaegr. i n Hamp., F l o r a (Jena) 20:274. 1837. Limnobium p a l u s t r e . Schimp., B r y o l . E u r . 6:66. Amblystegium subenerve  574. 1853.  Schimp., B r y o l . E u r . 6:52. 563. 1853.  (Type a t S-PA) Limnobium p a l u s t r e v a r . s u b s p h a e r i c a r p o n 6:67.  575§- 1853.  (Brid.) Schimp.  Bryol. Eur.  225 Limnobium p a l u s t r e v a r . hamulosum Schimp., B r y o l . E u r . 6:67. 575/3  1853.  Limnobium p a l u s t r e v a r . julaceum Schimp., B r y o l . E u r . 6:67. 5756  1853.  Limnobium p a l u s t r e v a r . laxum Schimp., B r y o l i 575Y  E u r . 6:67.  1853.  Hypnum r o e s e i Schimp., C o r o l l . 131. 1856. Hypnum p a l u s t r e v a r . t e n e l l u m Schimp., Syn. 634. Hypnum subeverve (Schimp.) Schimp., Syn. 634. Hypnum p a l u s t r e v a r . hamulosum  1860.  1860.  (Schimp.) Schimp., Syn. 634.  1860. Hypnum p a l u s t r e v a r . julaceum  (Schimp.) Schimp., Syn. 635. 1860.  Hypnum p a l u s t r e v a r . laxum (Schimp.) Schimp., Syn. 634. Hypnum p a l u s t r e s s p . s u b s p h a e r i c a r p o n Medd. Soc. F. F I . Fenn. 3:192. Amblystegium p a l u s t r e  ( S c h l e i c h . ex-Brid.) L i n d b . , 1878.  (Brid.) L i n d b . , Musci Scand. 33.  Hypnum k r a u s e i i C. M u l l . , F l o r a  1860.  (Jena) 70:219.  1879.  1887. (Holotype  a t S-PA) Hypnum c o l u m b i c o - p a l u s t r e C. M u e l l . P I . 6:241.  e t Kindb. i n Macoun, C a t . Canad.  1892.  Scleropodium k r a u s e i  (C. Muell.) Macoun e t Kindb. i n Macoun, C a t .  Canad. P I . 6:203.  1892.  Hypnum pseudo-montanum Kindb. i n Macoun, Cat'. Canad. P I . 6:243. 1892. Limnobium pseudomontanum 74.  (Kindb.) Kindb., Canad. Rec. S c i . 6 ( 1 ) :  1894.  Calliergon palustre  (Brid.) Kindb., B o t . Not. 1896:132.  1896.  226  Brachythecium k r a u s e i  (C. M u e l l . ) P a r . , I n d . B r y o l . 136.  1894.  Limnobium pachycarpulum. C. M u e l l . , Nuov. G i o r n . B o t . I t a l . n. ser.  3:116.  1896.  C a l l i e r g o n pseudomontanum 1:86.  (Kindb.) Kindb., E u r . N. Am.  Bryin.  1897.  Amblystegium  p a l u s t r e v a r . hamulosum(B.S.G.)  Moss. 3:60.  Braithw., B r i t .  1898.  Hypnum p a l u s t r e v a r . r o e s e i Hypnum pachycarpulum  (Schimp.)  Roth, Hedwigia 38:265. 1899.  (C. Muell.) P a r . , Ind. B r y o l . Supp. 206. 1900.  Hypnum subeugyrium v a r . o c c i d e n t a l e Card, e t Ther. Proc., Wash. Ac. Sc.  4:342.  1902.  Hygrohypnum p a l u s t r e  (Hedw.) Loesk., M o s s f l . Harz. 319.  1903.  Hygrohypnum p a l u s t r e v a r . s u b s p h a e r i c a r p o n ( S c h l e i c h . ex B r i d . ) Loesk., M o o s f l . Harz. 320.  1903.  Hygrohypnum p a l u s t r e v a r . julaceum (B.S.G.) Loesk., M o o s f l . Harz. 320.  1903.  Hygrohypnum p a l u s t r e v a r . laxum 320.  (B.S.G.) Loesk., M o o s f l . Harz.  1903.  Limnobium s p h a e r i c a r p o n De Not. ex P a r . , Ind. B r y o l . ed. 2, 3:194. 1905.  Nom.  i n v a l . i n synon. e r r , p r e . L. s u b s p h a e r i c a r p o n  (Brid.) De Not. Limnobium p a l u s t r e v a r . t e n e l l u m (Schimp.)  Roth, E u r . Laubm. 2:637.  1905. ' Hygrohypnum s u b s p h a e r i c a r p o n (Brid.) Loesk., Verh. Bot. V e r Brandenburg. 49:198. Hygrohypnum subenerve 49:198.  1905.  1905. (B.S.G.) Loesk., Verh. Bot. V e r . Brandenburg.  227 Limnobium  subenerve  (B.S.G.) Roth, E u r . Lauta."  Hygrohypnum p a l u s t r e v a r . tenBllum FI.  Brandenburg  2:1060.  (Schimp.) Warnst.,  56f4. 1905. Krypt.  1906.  Hygrohypnum p a l u s t r e v a r . r o e s e i FI.  2:637.  Brandenburg 2:1060.  (Schimp.)  1906.  p r o . Hypnum p a l u s t r e v a r . r o e s e i  Roth ex Warnst.,  Nom. i n v a l . (Schimp.)  Krypt.  i n synon. e r r . Roth.  Hygrohypnum p a l u s t r e v a r . tenuissimum Warnst., K r y p t . F l . Brandenburg  2:1060.  1906.  Hygrohypnum s u b s p h a e r i c a r p o n v a r . cataractum Loesk., A l l g . B o t . Z e i t s c h r . 13:120.  1907.'  Hygrohypnum k r a u s e i (C. Muell.) P a r . , C o l l . 16.  1909.  Hygrohypnum subenerve v a r . hamulosum Glow., C a r n o l i a n. s e r . 4: 143.  1913.  Hygrohypnum p a l u s t r e f o . v u l g a r i s Moenk., S u s s w a s s f l . 151. f 5 2 a . 1914. Hygrohypnum p a l u s t r e f o . hamulosa f52f.  (B.S.G.)' Moenk., S u s s w a s s f l . 151.  1914.  Hygrohypnum p a l u s t r e v a r . s u b s p h a e r i c a r p a Moenk., S u s s w a s s f l . 151.  f52c.  ( S c h l e i c h . ) B.S.G. i n  1914.  Hygrohypnum p a l u s t r e v a r . s u b s p h a e r i c a r p a f o . j u l a c e a Moenk., S u s s w a s s f l . 151.  f52d.  Hygrohypnum p a l u s t r e v a r , t s n e l l a . 151.  S u s s w a s s f l . 151.  (B.S.G.) Mosnk., S u s s w a s s f l .  f o . subenervis  (Schimp.) Moenk.,  f52e. o n l y . 1914.  Hygrohypnum p a l u s t r e v a r . s u b e n e r v i s  Limnobium  1914.  1914.  Hygrohypnum p a l u s t r e v a r . t e n e l l a  151.  (B.S.G.)  (Schimp.) Moenk., S u s s w a s s f l .  1914. pseudochraceum  Amann ex Roth, Hedwigia 57:139.  4 f . 6.  228 1915. Hygrohypnum'palustre 56:267.  v a r . neglectum  (Brid.) R o e l l , Hedwigia  1915.  Hygrohypnum p a l u s t r e s s p . subenerve S u i s s e 2:358.  (B.S.G.) Amann, F l . Mouss.  1919.  Hygrohypnum p a l u s t r e v a r . alpinum Amann, F l . Mouss. S u i s s e 2: 358.  1919.  Hygrohypnum p a l u s t r e v a r . e u - p a l u s t r e f o . hamulosa Moenk., Laubm. E u r . 736.  1927.  Hygrohypnum p a l u s t r e v a r . e u - p a l u s t r e f o . l l a x a V -Laubm...Eur. 736.  (B.S.G.) Moenk.,  1927.  Hygrohypnum p a l u s t r e v a r . t e n e l l u m f o . j u l a c e a Laubm. E u r . 736.  (B.S.G.) Moenk.,  1927.  Hygrohypnum subenerve v a r . plumulosum Amann, Mem. H i s t . Nat. 3:63.  17.  (Kindb.) Grout, C h e c k l i s t  Pleuroc.  1929.  Hygrohypnum eugyrium v a r . o c c i d e n t a l e l i s t Pleuro.  Soc. Vaudoise  1928.  Hygrohypnum pseudomontanum Moss. N. Am.  (B.S.G.)  Moss. N. Am.  16.  (Card, e t Ther.) Grout, Check1929.  Hygrohypnum eugyrium s s p . subeugyrium v a r . o c c i d e n t a l e Ther.) Grout, C h e c k l i s t P l e u r o c Moss. N. Am. Hygrohypnum p a l u s t r e ' s s p . pseudomontanum F l . N. Am.  3:89.  Hygrohypnum 465.  1929.  (Kindb.) Grout, Moss  1931.  Hygrohypnum l u r i d u m f o . hamulosum 465.  16.  (Card, e t  (B.S.G.) C. Jens., Skad.  Bladmf1.  l u r i d u m f o . julaceum (B.S.G.) C. J e n s . , Skad.  Bladmf1.  1939.  1939.  229 Hygrohypnum l u r i d u m f o . laxum (B.S.G.) C. J e n s . , Skand. B l a d m f l . 465.  1939.  Hygrohypnum l u r i d u m f o . subenerve B l a d m f l . 465.  (Schimp.) C. Jens., Skand.  1939.  Hygrohypnum l u r i d u m v a r . s u b s p h a e r i c a r p o n Skand. B l a d m f l . 465.  ( S c h l e i c h . ) C. J e n s . ,  1939.  Hygrohyphum p a l u s t r e v a r . malacocaulon Herz., Hedwigia 82:87. 14.  1944.  Hygrohypnum nervosissimum P a r r . , Rev. B r y o l . L i c h e n . 21:17. 15f.  1-8.  1952.  Hygrohypnum l u r i d u m v a r . e u - l u r i d u m f o . v u l g a r e Podp. Consp. 571.  (Moenk.)  1954.  Hygrohypnum l u r i d u m v a r . eu-luridum f o . hamulosum Podp., Consp. 571.  (B.S.G.)  1954.  Hygrohypnum l u r i d u m v a r . eu-luridum f o . laxum (B.S.G.) Podp., Consp. 571.  1954.  Hygrohypnum l u r i d u m v a r . t e n e l l u m  (Schimp.) Podp., Consp. 571.  1954. Hygrohypnum l u r i d u m v a r . t e n e l l u m f o . subenerve Consp. 571.  (Schimp.) Podp.,  1954.  Hygrohypnum l u r i d u m v a r . t e n e l l u m f o . plumulosum (Amann) Podp., Consp. 571.  1954.  Hygrohypnum l u r i d u m v a r . t e n e l l u m f o . hamulosum Consp. 571.  (Glow.) Podp.,  1954.  Hygrohypnum l u r i d u m v a r . t e n e l l u m f o . tenuissimum Consp. 571.  (Warnst.) Podp  1954.  Hygrohypnum l u r i d u m v a r . malacocaulon  (Herz.) Podp., Consp. 572.  2 30 Hygrohypnum l u r i d u m v a r . julaceum Hygrohypnum l u r i d u m v a r . Podp., Gonsp. 572.  (B.S.G.) Podp., Consp. 572. 1954.  s u b s p h a e r i c a r p o n f o . cataractum  (Loesk.)  1954.  Hygrohypnum l u r i d u m v a r . pseudochraceum (Amann) Podp., Consp. 572.  1954.  Hygrohypnum l u r i d u m v a r . pseudochraceum f o . alpinum Consp. 572.  (Amann) Podp.,  1954.  Nomina Nuda Hygrohypnum l u r i d u m v a r . eu-luridum f o . complanata Consp. 571.  (Roell) Podp.,  1954.  Hygrohypnum l u r i d u m v a r . obtusatum Podp., Consp. 572.  (Loesk. e t S t o l l e i n R o e l l )  1954.  Hygrohypnum p a l u s t r e v a r . complanatum R o e l l , Hedwigia 38:265. 1899. Hygrohypnum p a l u s t r e v a r . obtusatum Loesk. e_t S t o l l e i n R o e l l , Hedwigia 56:267.  1915.  Hypnum p a l u s t r e v a r . a u r o n i t e n s P r o g e l i n Mol., J a h r e s b e r . Naturh. V e r . Passau 10:2'68.  1875.  Hypnum p a l u s t r e v a r . pachyneuron Glow., Verh. Z o o l . B o t . Ges. Wien 57:22.  1907.  Hypnum - p a l u s t r e v a r . o r t h o c a r p o n Saut., M i t t h e i l . Ges. S a l z b . Landesk. 10:69.  1871.  Hypnum p a l u s t r e v a r . p f e f f e r i L o r , i n Limpr.,. "Krypt. F l . S c h l e s . 1:63.  1876.  Hypnum p a l u s t r e v a r . stabianum B o t t . , A t t i Soc. Toscana S c . Nat. 92:190.  1913.  231 Nomena Nuda n o t seen Hypnum d o l i c h o n e u r o n I n g l . e t Rhodes Ann. Rep. Moss Each. C l u b . 21:145 - 167.  1916.  Hypnum p a l u s t r e v a r . g r a c i l u s Ren. i n H o l z i n g e r , B r y o l o g i s t . 13:53.  1913.  Names t r e a t e d as synonyms elsewhere, u n a v a i l a b l e f o r assessment d u r i n g t h i s  b u t for. which the l i t e r a t u r e  was  study.  Hygrohypnum l u r i d u m f o . intextum C. M u e l l . , Syn. 2:142. 1851. Hypnum intextum V o i t , i n S t u r n . F l . Deutsch.  2:11 i c . 1811.  Hypnum pseudo-plumosum v a r . s u b s p h a e r i c a r p o n  (Brid.) Spruce,  Mag. Nat. H i s t . s e r . 2. Limnobium ambiguum  3:281.  De Not., Cronac.  1849, Briol. Ital.  Limnobium r e p t i l e f o r m e De Not., Cronac. Limnobium s u b s p h a e r i c a r p o n  Ann,  2:27.  Briol. Ital.  1867.  2:27.  1867.  (Brid.) L i n d b . , Medd. Soc. F. F l . Fenn.  Names o f c e r t a i n taxa and t h e i r a l l e g e d synonyms f o r which the l i t e r a t u r e was examined, b u t found inadequate  f o r e f f e c t i v e assessment o f the taxon i n  the absence o f the type specimens. Hypnum r u d e r a l e v a r . i n t r i c a s t u m B r i d . , B r y o l . Univ. 2:584. 1827. Hygrohypnum l u r i d u m f o . i n t r i c a s t u m (Brid.) C. Jens. Skand. B l a d m f l . 465.  1939.  Hypnum p a l u s t r e v a r . p r o l i x u m Mat., Hedwigia 44:44. Hygrohypnum p a l u s t r e v a r . p r o l i x u m Brandenburg 2:1060.  1905.  (Mat.) Warnst., K r y p t . F l .  1906.  Hygrohypnum l u r i d u m f o . p r o l i x u m Podp., Consp. 571.  1954.  Names o f taxa n o t p r e s e n t a t the d e s i g n a t e d s i t e s i n the l i t e r a t u r e . Hygrohypnum l u r i d u m f o . ambiguum 1939.  f i d e Index Muscorum.  c f . Jens., Skand. B l a d m f l . 465.  232 Hygrohypnum l u r i d u m f o . i n t r i c a s t u m C. J e n s . , Skand. 465.  1939.  f i d e Index Muscorum.  Hygrohypnum l u r i d u m f o . r e p t i l e f o r m e Bladmf1. 465.  Bladmf1.  1939.  (De Not.) c f . C. J e n s . , Skand.  f i d e Index Muscorum.  Hygrohypnum luridum+var. t e n e l l u m f o . subenerve C. Jens., Skand. Bladmf1. 465. 1939. f i d e Pddper'a i n r C o n s p .  233 Plants slender to robust,  forming l o o s e l y t o t i g h t l y woven p a t c h e s  o r t u f t s , o f t e n forming e x t e n s i v e or t u f t s e a s i l y degrees.  fragment; p l a n t s  mats; when l o o s e l y woven the p a t c h e s frequently clogged with s i l t  times a l l c o l o r v a r i a t i o n s may  same p l a n t . but  Stems (1)  a s c e n d i n g at the  3-5  (6)  grade from one cm  be b a d l y  to another w i t h i n  the  creeping,  apex; stems sometimes hooked a t the apex; u s u a l l y  abraded, b u t  leaves  near the stem base  seldom are the stems c l e a r l y denuded.  i r r e g u l a r , branches 1 - 2 stem c r o s s - s e c t i o n s  mottling,  or brownish-black,  l o n g ; mainly p r o s t r a t e and  f o l i o s e throughout, i n some o l d specimens the may  varying  Color v a r i a b l e , yellow-green to yellow-green with rusty  l e s s o f t e n yellow-brown, b r i g h t green, b l a c k i s h - g r e e n at  to  (3)  cm  Branching  l o n g , g e n e r a l l y a s c e n d i n g , sometimes e r e c t ,  r e v e a l i n g 3 to 4 rows o f s m a l l , y e l l o w i s h o r brownish,  t h i c k - w a l l e d c o r t i c a l c e l l s ; medullary c e l l s thinner-walled,  increasing in  s i z e toward the middle; c e n t r a l s t r a n d w e l l developed, o f numerous s m a l l c e l l s ; m e d u l l a r y c e l l s and  the  c e n t r a l s t r a n d g e n e r a l l y becoming more  d a r k l y pigmented w i t h age;  r h i z o i d s o f v a r i a b l e frequency, reddish-brown,  a r i s i n g a t the base o f v e n t r a l stem Leaves v a r i a b l e , but (0.5)  1.0  - 1.5  (2.5)  mm  changing l i t t l e long X  (0.25) 0.4  able, lanceolate, oblong-lanceolate, leaves  leaves. from the wet - 0.75  ovate and  (1.1)  to dry mm  condition;  wide; shape  o c c a s i o n a l l y broadly  vari-  ovate;  s t r a i g h t f o r f a l c a t e , r a r e l y almost c i r c i n a t e ; apex i n l a n c e o l a t e  severely  f a l c a t e leaves  or  u s u a l l y t a p e r i n g g r a d u a l l y t o an acute p o i n t ; i n o b l o n g -  l a n c e o l a t e , ovate or l e s s severely  f a l c a t e leaves  the apex u s u a l l y t a p e r i n g i n t o  a s m a l l a b r u p t l y d i f f e r e n t i a t e d a p i c u l u s ; margins e n t i r e , i n r o l l i n g v a r i o u s l y , f r e q u e n t l y i n r o l l i n g on one oblong-lanceolate  s i d e o f the  l e a f c r e a t i n g i n f o l d e d wings; i n  o r o v a t e l e a v e s margins f r e q u e n t l y i n r o l l on b o t h s i d e s o f  the l e a f i n upper q u a r t e r  accentuating  the a b r u p t l y  tapered  apiculus.  Leaves  234 v a r i o u s l y d i s t a n t to crowded; a t t i t u d e upon stem v a r i a b l e , s p r e a d i n g , l o o s e l y to t i g h t l y i m b r i c a t e d , f a l c a t e from s l i g h t l y curved v a r i a t i o n s may  occur  almost j.ulaceous o r any  to almost c i r c i n a t e ;  as a . g r a d i e n t  or more o f  these  a l o n g a g i v e n l e n g t h o f stem o r branch,  the most f r e q u e n t v a r i a t i o n combination b e i n g versa;  two  v a r i a t i o n of  i m b r i c a t e - f a l c a t e or vice  f r e q u e n t l y a degree o f c o r r e l a t i o n e x i s t s between l e a f shape or  degree of f a l c a t i o n and  l e a f p o s i t i o n upon the s t e m — l a n c e o l a t e  t o e x h i b i t a g r e a t e r frequency  i n the s p r e a d i n g  l e a v e s seem  a t t i t u d e ; among f a l c a t e  l e a v e s t h e r e i s an o c c a s i o n a l tendency f o r l e a v e s p o s i t i o n e d d o r s a l l y on prostrate while  'stems to be  s t r o n g l y f a l c a t e w i t h d e c i d e d l y i n r o l l e d margins  l e a v e s p o s i t i o n e d on the f l a n k s o f such p r o s t r a t e stems are o f t e n  r a t h e r ovate, a wing, and  g e n t l y f a l c a t e w i t h the d o r s a l l y p r o x i m a l margin i n f o l d e d as  the v e n t r a l l y p l a c e d  l e a v e s may  o r v a r i o u s l y f a l c a t e i n the upper h a l f .  e i t h e r be  squarrose-caniculate  Leaves almost p l a n e ,  cave or c a n i c u l a t e , e s p e c i a l l y when f a l c a t e .  Costa v a r i e s  shallowly  con-  continuously  throughout the s p e c i e s and w i t h i n i n d i v i d u a l l e a v e s o f the same p l a n t ; s i n g l e and  s h o r t to p e r c u r r e n t ,  s h o r t and  double or absent.  A r e o l a t i o n extremely v a r i a b l e ; median l e a f c e l l s v a r i o u s l y s h o r t rhombic to long, l i n e a r flexuose,  (28)  30 - 55  (95)  um  the average dimensions o f median l e a f c e l l s may  long X vary  (4) 5 - 7 from one  (12) um  wide;  l e a f to another  on the same stem; a p i c a l l e a f c e l l s g e n e r a l l y s h o r t e n i n g g r a d u a l l y or changing little  from the median c e l l s ; b a s a l l e a f c e l l s become s h o r t e r and wider than  median l e a f c e l l s or only s l i g h t l y wider: i n e i t h e r case the b a s a l c e l l s be weakly to s t r o n g l y p i t t e d i f a t a l l ; defined  a l a r c e l l s v a r i a b l e , present  group o f q u a d r a t e , s h o r t r e c t a n g u l a r or i r r e g u l a r l y shaped  v a r y i n g from t h i n and h y a l i n e to i n c r a s s a t e and reddish-brown; p l a n e  may  as a w e l l cells;  d i s c o l o r e d yellow-brown or  or excavated, excavated c e l l s sometimes be  inflated  to  235 v a r y i n g degrees. P l a n t s a u t o i c o u s ; p e r i g o n i a ovate; l a n c e o l a t e 0.3  - 0.7  mm  l o n g , deeply  e n c l o s i n g s e v e r a l a n t h e r i d i a and  p e r i g o n i a l l e a v e s ovate to  concave i m b r i c a t e , e n t i r e ,  ovate-  ecostate,  numerous paraphyses'; O u t e r and middle p e r i -  c h a e t i a l l e a v e s ovate l a n c e o l a t e o r l a n c e o l a t e , squarrose  recurved  i n the  upper h a l f , c o s t a s i n g l e or absent; i n n e r p e r i c h a e t i a l l e a v e s l o n g t a p e r i n g l a n c e o l a t e , to 3.5  mm  l o n g , e r e c t , e n t i r e , c o s t a s t r o n g and  and double or almost absent, s t o u t or Seta 8 - 2 4  mm  arcuate,  long  e r e c t or a r c h i n g when moist,  var-  slender.  l o n g , smooth w a l l e d ,  i o u s l y t w i s t i n g upon d r y i n g .  s i n g l e or  Capsule 1 - 3 mm  l o n g ; when wet  usually inclined  l e s s o f t e n e r e c t c y l i n d r i c a l ; upon d r y i n g the capsule may  or  may  not c o n s t r i c t below the mouth, when c o n s t r i c t e d below the mouth the lower h a l f o f the c a p s u l e  i s rather bulbous.  Capsule w a l l c e l l s i n c r a s s a t e , y e l l o w  reddish-brown, o v a l , quadrate, r e c t a n g u l a r or i r r e g u l a r . e r i n g neck b e a r i n g  8-30  Peristome t y p i c a l  superficial  Capsules w i t h  tap-  stomata.  f o r the genus; annulus absent, endostome w i t h  p o o r l y to w e l l d e f i n e d c i l i a .  or  1-3  Spores dusky y e l l o w , p a p i l l o s e , 10 - 26 um  in  diameter.  Hygrohypnum luridum  (Hedw.) Jenn. i s an extremely v a r i a b l e s p e c i e s .  measure o f t h i s v a r i a b i l i t y and  subspecific levels.  i s seen i n the 115  However, t h i s study  names a s c r i b e d a t the  shows t h a t H.  p o l y m o r p h i c s p e c i e s t h a t cannot be  subdivided  c o n s i s t e n t taxa.  continuously  The  specific  l u r i d u m must be  viewed as a s i n g l e , e x c e e d i n g l y into morphologically  A  effectively  species varies v i r t u a l l y  i n nine or t e n f e a t u r e s , which have been used s i n g l y o r i n v a r i o u s  combinations as c r i t e r i a  f o r the r e c o g n i t i o n o f numerous s u b s p e c i f i c t a x a .  These are o v e r a l l p l a n t s i z e , l e a f shape, a t t i t u d e o f the l e a v e s on the degree o f f a l c a t i o n , degree of i n r o l l i n g o f the l e a f margin, i n t e r n o d e  stem, length,  236 c o s t a l s t r u c t u r e , degree o f l e a f c o n c a v i t y , a l a r d i f f e r e n t i a t i o n and the degree  t o which the a l a r c e l l s are excavated.  The  s i z e o r o v e r a l l robustness o f t h i s s p e c i e s v a r i e s w i d e l y .  variation i s d i f f i c u l t  t o measure and i s d e a l t w i t h i n q u a l i t a t i v e  which u n f o r t u n a t e l y are vague and i m p r e c i s e .  mats s e v e r a l decimeters a c r o s s .  mats may may  o r t i g h t l y woven w i t h l i t t l e  be t h i n due  b e r y e r y o l o o s e l y woven and  fragmentation.  easily  These patches o r  to the p r o s t r a t i o n o f t h e i r component branches  Stems can be s l e n d e r , t h a t i s l e s s than 1 mm  thicker,' ca. 2.5 mm.  form  to extensive  be t h i c k and almost t u r f l i k e as a consequence o f the ascending  and stem t i p s .  tip  These may  terms,  Hygrohypnum l u r i d u m can  s m a l l patches o r t u f t s a few c e n t i m e t e r s square a l l the way  fragmented  This  o r they branches  i n diameter, or  (Thickness i s the d i s t a n c e i n m o i s t p l a n t s from  to l e a f t i p from one s i d e of the stem t o the other.)  leaf  Variation i n stature  i s a f r e q u e n t environmental response among p l a n t s , thus i t seems r e a s o n a b l e to  assume t h a t v a r i a t i o n o f t h i s s o r t i s such a response. The  l e a v e s are e x c e e d i n g l y v a r i a b l e .  C e r t a i n f e a t u r e s v a r y and  v a r i a t i o n s i n t u r n i n f l u e n c e the v a r i a t i o n o f o t h e r l e a f f e a t u r e s . q u e n t l y , the v a r i a b l e l e a f c h a r a c t e r s are d i f f i c u l t  to t r e a t  B a s i c l e a f shape v a r i e s from narrowly l a n c e o l a t e (Fig.  51 b ) .  (Fig.  51 f , g, i , o, p, q e t c ) .  these Conse-  individually.  ( F i g . 51 m)  t o ovate  The most common shape i s ovate l a n c e o l a t e o r o b l o n g - l a n c e o l a t e Leaves  e x h i b i t i n g such shapes  on the same p l a n t s and i n r a r e i n s t a n c e s they may  o f t e n occur  o c c u r immediately a d j a c e n t '  one another w i t h out l e a v e s t r a n s i t i o n a l shapes between them. The  l e a f apex may  g r a d u a l l y t a p e r t o a sharp p o i n t  i n most i n s t a n c e s appears f, p, e t c ) .  ( F i g . 51 j & m),  t o t a p e r a b r u p t l y t o an a p i c u l a t e p o i n t  but  ( F i g . 51  T h i s i s most e v i d e n t i n oblong ovate o r oblong l a n c e o l a t e l e a v e s .  237 In r e a l i t y the a p i c e s g r a d u a l l y t a p e r t o an acute p o i n t  ( F i g . 53 a & b) b u t  appear a p i c u l a t e r e s u l t i n g from the i n r o l l i n g o f the l e a f margin j u s t below the apex.  T h i s accentuates the t a p e r and a r t i f i c i a l l y  d i f f e r e n t i a t e s the  apiculus. As  continuous as the v a r i a b i l i t y o f l e a f shape i s w i t h  modifying  its  f e a t u r e s , c e r t a i n f e a t u r e s show a c o r r e l a t i o n w i t h  Narrowly l a n c e o l a t e t o ovate l a n c e o l a t e l e a v e s from s l i g h t l y  leaf  attendent shapes.  are frequently v a r i o u s l y  falcate.  Such l e a v e s v a r y  circinate  ( F i g . 52 v ) A n e n t i r e l e a f may be f a l c a t e ( F i g . 52 u) o r o n l y  the upper h a l f ( F i g . 52 p ) .  f a l c a t e ( F i g . 51 n) t o almost  The asymmetry o f a l e a f as a consequence o f the  f a l c a t i o n may a l s o be more e v i d e n t on one s i d e o f the l e a f  ( F i g . 51 e ) .  Very commonly the degree o f f a l c a t i o n may depend on the d o r s a l , or v e n t r a l i n s e r t i o n o f the l e a v e s upon the p r o s t r a t e stem. l e a v e s may be s e v e r e l y f a l c a t e t o almost c i r c i n a t e and deeply  Dorsally placed tubulose.  l e a v e s a r c h h i g h above t h e i r p o i n t o f i n s e r t i o n , c u r v i n g a b r u p t l y s u b s t r a t e and o c c a s i o n a l l y t h e t i p s curve completely 50 d ) .  In o t h e r  substrate.  instances  tubulose  simply  Such  toward the  beneath t h e stem ( F i g . curve g r a d u a l l y toward the  Sometimes i n the more ovate l a n c e o l a t e l e a v e s the e x p r e s s i o n o f  f a l c a t i o n i s confined spicuous  the d o r s a l l e a v e s  lateral  there.  t o the upper h a l f o f the l e a f , o r i s a t l e a s t more con-  In these the f a l c a t e upper h a l f i s s e v e r e l y curved  ( F i g . 51  and deeply  1 ) . Leaves i n s e r t e d l a t e r a l l y on t h e p r o s t r a t e stem may  e i t h e r be s e v e r e l y f a l c a t e as those d o r s a l l y i n s e r t e d o r may be f a l c a t e by means o f an i n r o l l e d l e a f margin.  In t h i s l a t t e r s i t u a t i o n the l e a f  the d o r s a l s i d e o f the stem i s i n r o l l e d toward the l e a f m i d l i n e . may be g r a d u a l l y o r • a b r u p t l y  falcate.  The more ovate l a n c e o l a t e  Such a l e a f leaves  i n s e r t e d l a t e r a l l y g e n e r a l l y a r e more g r a d u a l l y f a l c a t e than narrower L e s s f a l c a t e ovate l a n c e o l a t e l e a v e s  nearer  leaves.  a r e p l a c e d a d a x i a l s i d e down on a m i c r o -  238  scope s l i d e and covered w i t h a cover s l i p , f a l c a t e appearance. a d a x i a l s i d e down.  Thus the i n r o l l e d wing opens when t h e l e a f i s p l a c e d The p r e s s u r e o f the cover s l i p  making i t appear more t r u l y ovate phenomenon. falcate.  they are seen t o l o s e t h e i r  lanceolate.  f l a t t e n s the l e a f ,  T h i s , however, i s a v a r i a b l e  Leaves s i t u a t e d v e n t r a l l y on a p r o s t r a t e stem are a l s o v a r i a b l y  Sometimes•these are n o t so much f a l c a t e as they a r e squarrose t u b -  u l o s e i n the upper h a l f  ( F i g . 51 c) and o c c a s i o n a l l y , the squarrose  upper h a l f i s d e f l e c t e d t o one s i d e by the s u b s t r a t e . primarily  On-some p l a n t s w i t h  f a l c a t e the v e n t r a l l e a v e s a r e p e r f e c t l y ovate  h i n t of f a l c a t i o n .  tubulose  l a n c e o l a t e w i t h no  I n these, the v e n t r a l l e a v e s a r e i m b r i c a t e d among them-  selves . Narrowly l a n c e o l a t e o r ovate  l e a v e s may vary s t i l l  further.  Generally  the l e a v e s a r e q u i t e crowded along the stem o r they may be q u i t e d i s t a n t . There d i s t a n t l e a v e s may.occur i n two r e g i o n s ; on s l e n d e r branches, p o s t e r i o r p o r t i o n s o f o l d e r stems. d i r e c t l y with f a l c a t i o n .  or i n  I f o f t e n appears t h a t c o n c a v i t y v a r i e s  The more pronounced the f a l c a t i o n ,  the more p r o -  nounced the l e a f c o n c a v i t y o r c a n i c u l a t i o n . F r e q u e n t l y , the narrowly e x h i b i t i n g no f a l c a t i o n .  l a n c e o l a t e o r ovate  l a n c e o l a t e l e a v e s are s t r a i g h t ,  I n these i n s t a n c e s , there i s v a r i a t i o n i n the  a t t i t u d e o f the l e a v e s upon the stem, s p a c i n g o f the l e a v e s along the stem, the l e a f c o n c a v i t y and the i n r o l l i n g o f the margin. t i g h t l y i m b r i c a t e d t o very w i d e l y distant.  In some cases,  spreading.and  These l e a v e s vary  they v a r y from crowded t o  those l e a v e s t h a t a r e w i d e l y s p r e a d i n g and d i s t a n t  are a l s o c r i s p e d when d r y and e x h i b i t c o n s i d e r a b l e l a t e r a l s h r i n k a g e w i t h some t w i s t i n g . spaced, narrowly  from  coupled  Not i n f r e q u e n t l y , those axes b e a r i n g c r i s p e d , w i d e l y  s p r e a d i n g l e a v e s a r e r a t h e r s p i n d l y secondary l a n c e o l a t e t o ovate  branches.  In s t r a i g h t ,  l a n c e o l a t e l e a v e s c o n c a v i t y when p r e s e n t i s more  239  e v i d e n t than i n f a l c a t e l e a v e s which o f t e n p a r t i c u l a r l y e v i d e n t when the l e a v e s are i m b r i c a t e d . l a t e to ovate  Sometimes i n very l o n g , o l d stems the narrowly  l e a v e s are w i d e l y  spaced and  normal from the stem base to the apex. r e s u l t of maturation  grade from v e r y s m a l l s i z e  lanceoto  T h i s phenomenon appears to be  the  from j u v e n i l e to a d u l t l e a v e s .  In a s i g n i f i c a n t l y l a r g e number o f specimens l e a v e s o c c u r i n such shapes as; o v a t e , but narrowing to an acuminate t i p ; oblong  t o oblong  acute  These b a s i c shapes  t i p ; and b r o a d l y ovate o r b r o a d l y  grade i m p e r c e p t i b l y from one ures.  Most i m p o r t a n t l y ,  Beyond t h i s ,  these  to another,  lanceolate.  feat-  5  l e a v e s of•, these shapes are most t y p i c a l l y  l e a v e s may  with  v a r y i n g i n a number o f minor  straight.  v a r y i n t h e i r a t t i t u d e upon the stems, t h e i r  s p a c i n g a l o n g the stem, t h e i r c o n c a v i t y and l e a f margins.  ovate  the degree o f i n r o l l i n g o f  A f r e q u e n t m o d i f i c a t i o n among these l e a v e s has  i n r o l l i n g of the l e a f margin and  the  to do w i t h  the d e l i m i t a t i o n o f a s m a l l a p i c u l u s .  In  a l l o f these l e a f shapes i t i s common f o r the margin to i n r o l l s l i g h t l y near m i d l e a f  to s h o r t l y below the apex.  As  a r e s u l t of the i n r o l l i n g ,  t a p e r of the l e a f near the apex i s a b r u p t l y changed such t h a t an  the  from the  apical  apiculus i s evident. ~The widely cated.  s p a c i n g o f these v a r i o u s l y shaped l e a v e s on the stem v a r i e s from very  spaced to very crowded.  They may  f u r t h e r vary from s p r e a d i n g  Sometimes they are concave or weakly tumid.  v i r t u a l l y plane.-  At o t h e r times they  F r e q u e n t l y , much o f t h i s v a r i a t i o n may  g r a d i e n t from p o s t e r i o r o f the stem t o the stem apex. a t the p o s t e r i o r stem e x t r e m i t i e s are w i d e l y  to i m b r i -  be e v i d e n t along  In such cases the  spaced and s p r e a d i n g  are a leaves  to almost  erect spreading.  Toward the stem apex the l e a v e s become g r a d u a l l y more crowded  and  imbricated.  conspicuously  widely  and  i n isome cases  The  c o n c a v i t y of these l e a v e s v a r i e s  qiite  the l e a v e s are almost c u c u l a t e , whereas, i n o t h e r s ,  the c o n c a v i t y i s v e r y s h a l l o w .  A consequence of the pronounced c o n c a v i t y i n  240 some l e a v e s the stems are  julaceous.  T y p i c a l l y Hygrohypnum l u r i d u m has  a w e l l d e f i n e d group o f quadrate  to s h o r t r e c t a n g u l a r or o c c a s i o n a l l y i r r e g u l a r l y 54 a & b ) .  shaped a l a r c e l l s  T y p i c a l l y the c e l l s are i n c r a s s a t e , but they may  thin walled.  They may  brown and may  vary  o f these v a r i a b l e s may  to d e c i d e d l y excavated  o c c u r on the same p l a n t and  does seem to be a g r e a t e r  frequency  l e a v e s than i n s t r a i g h t ' l e a v e s .  l e a v e s w i t h excavated.'alar s t r a i g h t , almost b r o a d l y  to red-  ( F i g . 54 d - f ) .  i n the same l e a f .  Each There  f o r excavated a l a r c e l l s i n f a l c a t e  A l a r c e l l s o f Hygrohypnum l u r i d u m have  never been d e s c r i b e d as i n f l a t e d .  However, a few  zones may  alar cells i n falcate  be q u i t e i n f l a t e d  ( F i g . 54 f ) .  ovate l e a v e s t h e r e i s a l s o a g r e a t e r  of thin-walled, hyaline or only s l i g h t l y d i s c o l o r e d a l a r The  be somewhat  be h y a l i n e or s t r o n g l y d i s c o l o r e d y e l l o w  from p l a n e  (Fig.  c o s t a o f Hygrohypnum l u r i d u m  i s exceedingly  independent o f any o t h e r l e a f c h a r a c t e r . s i n g l e and p e r c u r r e n t to s h o r t and  The  Among  frequency  cells.  v a r i a b l e and  i s so  c o s t a v a r i e s from s t r o n g ,  double to v i r t u a l l y absent.  Among the  v e r y s m a l l e s t ' l e a v e s the c o s t a i s more f r e q u e n t l y absent, but t h i s does not mean t h a t l a r g e r l e a v e s have more w e l l developed c o s t a e . too may  completely  lack a costa.  In f a c t ,  As w i t h o t h e r l e a f f e a t u r e s i t i s not  common f o r the e n t i r e range o f v a r i a t i o n to occur w i t h i n one  In the o r i g i n a l d e s c r i p t i o n o f Hypnum l u r i d u m Hedwig designate  a type  specimen.  c o l l e c t e d from d e c a y i n g  Thorough study o f the s p e c i e s has  un-  plant.  (1801) d i d not  Hedwig d i d g i v e h a b i t a t and g e o g r a p h i c a l  i n d i c a t i n g t h a t the p l a n t was Germany.  they  date  m a t t e r i n Sweden  r e v e a l e d t h a t o n l y two  and  specimens  t h a t are a t t r i b u t a b l e to Hedwig and/or h i s herbarium are i n e x i s t a n c e . specimens are a t G. subsphaericarpon.  One  Both  specimen b e a r i n g no c o l l e c t i n g date i s named Hypnum  I t bears the G a c c e s s i o n number 3079/184.  The  second  241 specimen i s e n t i r e l y r e p r e s e n t a t i v e o f the f a l c a t e forms o f the s p e c i e s , it  comes from an a c c e p t a b l e g e o g r a p h i c a l area and a t l e a s t o f f e r s  p o s s i b i l i t y o f having been seen by Hedwig. sheet number 3079/156 as the  the  Therefore, I designate  neotype- f o r Hygrohypnum l u r i d u m  G  (Hedw.)  Jenn. From out o f the mass o f c h a r a c t e r v a r i a t i o n t h a t i s Hygrohypnum l u r i d u m numerous p a s t workers have p e r c e i v e d and d e s c r i b e d as assortment s p e c i f i c taxa.  The  non-existence  of  these  f o r the purposes of t h i s study and  o f many of t h e i r type d e s c r i p t i o n s has made d i f f i c u l t  their reality.  sub-  o f the type specimens f o r many o f  taxa or a t l e a s t t h e i r u n a v a i l a b i l i t y ambiguity  of  the  an assessment  C a r e f u l study o f the a v a i l a b l e type d e s c r i p t i o n s , o t h e r  c r i t i c a l l i t e r a t u r e and  a d e t a i l e d a n a l y s i s o f over 500  specimens o f the  s p e c i e s from throughout  i t s range has p e r m i t t e d an assessment o f the s p e c i e s  and i t : a l l e g e d s u b s p e c i f i c taxa as r e c o g n i z e d by Index Muscorum  (1959  - 1969).  I t i s the author's o p i n i o n t h a t t h e r e are no m o r p h o l o g i c a l l y c o n s i s t e n t ent i t i e s w i t h i n the s p e c i e s .  A l l <nf the s u b s p e c i f i c taxa have been based on v a r -  i a b l e f e a t u r e s or have been e r r o n e o u s l y p l a c e d w i t h i n the species.' Hygrohypnum l u r i d u m v a r . s u b s p h a e r i c a r p o n l u r i d u m f o . alpinum  (Brid.) C. Jens., Hygrohypnum  (Amann) Podp. and Hygrohypnum l u r i d u m f o . nervosissimum  (Parr.) Podp. have been based p r i m a r i l y on the s t r e n g t h o f t h e i r c o s t a e . Hypnum s u b s p h a e r i c a r p o n ,  Bridel  (1812) noted  "nervo f u b e x c u r r e n t e "  Of  and went  on to say "supreme tantum f a l c a t o - f e c u n d o nervo c r a f f o f e r r u g i n e o - f u s c o ad apicem p r o d u c t o of  H.  fpecific diftinctum."  l u r i d u m v a r . alpinum  apex and  as h a v i n g  a strong c o s t a prolonged  Amann (1912) c h a r a c t e r i z e d the  leaves  c o n s p i c u o u s l y i n r o l l e d margins near the  to j u s t beneath the apex.  Parriat  t h a t Hygrohypnum nervosissimum had a p i c u l a t e l e a v e s with the margin  (1952) noted inrolled  and a s i n g l e c o s t a d i s a p p e a r i n g j u s t below the apex, sometimes with a c o s t a l  242  branch  reaching midleaf.  indisputable. throughout  That these p l a n t s are c l e a r l y r e c o g n i z a b l e i s  However, as d i s c u s s e d e a r l i e r the c o s t a v a r i e s c o n t i n u o u s l y  the s p e c i e s .  I t i s easy to f i n d p l a n t s resembling  taxa i n a l l r e s p e c t s , but p o s s e s s i n g a s h o r t double t u r e o f s t r o n g s i n g l e and  s h o r t and double  each o f  these  c o s t a o r a v a r i a b l e mix-  costae.  Each o f these  taxa  simply r e p r e s e n t t h a t end o f the spectrum o f c o s t a l v a r i a t i o n i n which the costae are a l l , o r almost  a l l , s t r o n g and  single.  Consequently,  these  taxa  do not warrant taxonomic r e c o g n i t i o n . Hygrohypnum l u r i d u m f o . laxum tenuissimum  (B.S.G.) Podp., Hygrohypnum l u r i d u m f o .  (Warnst.) Podp.,.Hygrohypnum l u r i d u m v a r . t e n e l l u m  and Hygrohypnum l u r i d u m f o . subenerve similar features. plants. atue.  Schimper  (Schimp.) Podp.  (Schimp.) C. Jens, are a l l based on  In g e n e r a l they are c h a r a c t e r i z e d as s l e n d e r o r s m a l l (1853) d e s c r i b e d v a r . laxum as " g r a c i l o r e s , p a r t i m denud-  F o l i a r e m o t i u s c u l a , undique p a t e n t i a , o v a t o - l a n c e o l a t a . "  Schimper  (1876) f u r t h e r c h a r a c t e r i z e d Limnobium p a l u s t r e v a r . t e n e l l u m as "minus, formis majoribus  H_. i n c u r v a t i s i m i l i ,  acuminatis." spitosum, folis vel  Schimper  folis  illis  var.  (Laxum) s a t s i m i l i b u s , s u b i t o  (1853) d e s c r i b e d Amblystegium subenerve as "laxue  r i g i d u l u m ; c a u l e d i v i s i o p a r t i m denudato vage p i n n a t i m  acuminate-ovatis,  ramuloso;  valde concavis, semi-costatis, costa lutescente s i m p l i c i  bifurca, interrimis  "  F i n a l l y , Warnstorf  (1906) d e s c r i b e d H. p a l u s t r e  v a r . tenuissimum as stems very s h o r t , asending,  , l e a v e s moderately  spaced,  to 0.8  mostly  c l e a r l y secund, s m a l l l a n c e o l a t e , 0.5  e c o s t a t e o r w i t h a very s h o r t , very t h i n simple c o s t a . stegium  subenerve Schimp.  (at S-PA)  specimens v a r i o u s l y determined examined.  cae-  was  as one  mm  X 0.2  loosely  to 0.3  mm,  Only the type o f Ambly-*.  a v a i l a b l e f o r study.  However numerous  o r the o t h e r o f these t a x a have been  In most cases, such p l a n t s are i n v a r i a b l y very poor and  specimens, o f t e n c o n s i s t i n g of o n l y a few  stems.  grown under l e s s than o p t i m a l environmental  fragmented  These p l a n t s appear to have  conditions.  It i s likely,  there-  243 f o r e , t h a t each o f these  a l l e g e d taxa are m o r p h o l o g i c a l  abberations  and  not  worthy o f taxonomic r e c o g n i t i o n . Overall size,  costalJmorphology,  l e a f f a l c a t i o n , and  a l l variable  c h a r a c t e r s , have been used to c h a r a c t e r i z e Hygrohypnum l u r i d u m sphaericarpon H.  luridum  ococaulon workers 1927;  (Brid.) O.  ssp. pseudomontanum  ( B r i d e l , 1812;  Nyholm, 1965;  l a r g e , very 3/4  1876;  Milde,  1869;  Husnot, 1894;  and Moenkemeyer, 1927)  losum as a s m a l l , s l e n d e r p l a n t w i t h  Kindberg  and  more r o b u s t  simple  Various  1827)  1898;  have a l l  as a very r o b u s t p l a n t a strong, slender same  costa  authors  Braithwaite,  1898;  v a r i o u s l y d e s c r i b e d f o . hamu-  falcatefeseeund l e a v e s .  However, o n l y  Husnot remarked on the s h o r t and  double  nature  (1897) observed the s i m i l a r i t y o f s s p . pseudo-montanum  d i s t i n g u i s h e d the s u b s p e c i e s  recurved  on the b a s i s o f i t s l o n g e r  stems, the l a r g e r , t h i n n e r more l o o s e l y d i s p o s e d ,  less d i s t i n c t l y was  He  mal-  Moehkemeyer,  w i t h Hygrohypnum montanum on the b a s i s o f the v a r i o u s l y i n c u r v e d or f a l c a t e leaves.  Jens.,  Braithwaite,  Sprengel,  S i m i l a r l y , many o f these  L i m p r i c h t , 1904;  Schimper, Dixon, B r a i t h w a i t e  1904;  Dixon, 1924;  luridum var. subsphaericarpon  Nyholm, 1965;  (B.S.G.) C.  (Warnst.) Podp.  Limpricht,  f a l c a t e secund, concave l e a v e s with  (Schimper, 1853,  o f the c o s t a .  1876;  Lawton, 1971;  Jensen 1939;  o f the l e a f l e n g t h .  Dixon, 1924;  f o . tenuissimum  Schimper, 1853,  Roth, 1905;  f o . hamulosum  (Kindb.) Wijk e t Marg., H_. l u r i d u m v a r .  Grout, 1931;  v a r i o u s l y d e s c r i b e d H.  reaching  H_. l u r i d u m  (Herz) Podp. and H_. l u r i d u m  Husnot, 1894;  with  Jens.,  v a r . sub-  longer  decurrent,  s e r r u l a t e l e a v e s , whose a l a r c e l l s were l a r g e r and whose c o s t a  and p r o l o n g e d  to above the m i d d l e .  v a r . tenuissimum as h a v i n g  very s h o r t  (1 cm),  Warnstorf ascending  (1906) c h a r a c t e r i z e d stems, b e a r i n g r h i z o i d s  o n l y near the base and whose l e a v e s were r a t h e r l o o s e l y spaced, c l e a r l y  secund,  narrowly l a n c e o l a t e , u s u a l l y 0.5  to  s h o r t c o s t a t e and  t o 0.6  mm  long X 0.2  mm  wide, e c o s t a t e  b e a r i n g a few quadrate to s h o r t r e c t a n g u l a r alar* c e l l s .  244  Study o f v a r . s u b s p h a e r i c a r p o n ,  f o . hamulosum and  f o . tenuissimum have  shown them to based on c o n t i n u o u s l y v a r i a b l e o v e r a l l p l a n t s i z e , f a l c a t i o n and may  c o s t a l morphology.  have s h o r t and  ilarly, double  double  Very  l a r g e , robust f a l c a t e leaved plants  or s i n g l e costae or they may  s m a l l , s l e n d e r f a l c a t e l e a v e d p l a n t s may costae.  The  ambiguous concepts  leaf  be e c o s t a t e .  Sim-  have s t r o n g s i n g l e or s h o r t  o f r o b u s t and s m a l l and s l e n d e r have  caused p r e v i o u s workers t o c a l l p l a n t s o f e s s e n t i a l l y t h e same s i z e b o t h subsphaericarpon dimensions. occurrence  and  f o . hamulosum r e g a r d l e s s o f the p l a n t ' s r e a l  var.  linear  A more fundamental problem among these t a x a i s the very  frequent  o f s t r a i g h t l e a v e s on stems b e a r i n g a m a j o r i t y o f f a l c a t e l e a v e s .  I t i s not unusual  to come upon stems t h a t have segments of f a l c a t e  leaves  a l t e r n a t i n g a t i r r e g u l a r i n t e r v a l s - w i t h segments b e a r i n g s t r a i g h t l e a v e s .  As  a consequence, none o f the c h a r a c t e r s s e r v i n g as the b a s i s f o r these taxa are s u f f i c i e n t l y c o n s i s t e n t to serve as a sound taxonomic base. not r e c o g n i z e these  T h e r e f o r e , I do  taxa.  The p o s i t i o n o f Hygrohypnum l u r i d u m ssp. pseudomontanum i n t h i s group i s somewhat ambiguous. subspecies  As noted  e a r l i e r , Kindberg  (1931) t r e a t e d the supposed  as a s p e c i e s a l l i e d w i t h Hygrohypnum montanum.  the supposed s u b s p e c i e s i s not even remotely v a r i a n t o f H,  luridum.  Kindberg  The  r e l a t e d to H_. montanum, b u t i s a  a c c u r a t e l y noted  the f r e q u e n t s i m i l a r i t y  the v a r i a b l e i n c u r v a t u r e and r e c u r v a t u r e of the l e a f margins. noted  the l e s s d i s t i n c t l y s e r r u l a t e l e a f margins.  in fact clearly entire.  Kindberg  f a c t s are t h a t  The  c o r r e c t l y observed  He  of  mistakenly  a l l e g e d subspecies i s  t h a t the l e a v e s o f the  c a l l e d s u b s p e c i e s were l a r g e r than those o f H.^montanum.  However, the  so  leaf  s i z e o f ssp. pseudomontanum s h o u l d be compared w i t h the s l e n d e r , f a l c a t e l e a v e d forms o f H. s u b s p e c i e s was t h i s a r e a and  l u r i d u m t h a t have been a t t r i b u t e d to f o . hamulosum.  o r i g i n a l l y d e s c r i b e d from Owen Sound, O n t a r i o .  The  P l a n t s from  the a d j a c e n t Great Lakes area are o f t e n s l i g h t l y l a r g e r than  the  245  t r a d i t i o n a l fo. hamulosum. blunter leaf  apices  Grout (1931) accurately noted the s l i g h t l y  which often occur i n ssp. pseudomontanum.  However,  the character merges imperceptibly with other small falcate leaved variants of H. luridum i n the Great Lakes area.  Grout (1931) accurately  noted the simultaneous occurrence of straight and falcate leaves i n ssp. pseudomontanum, but did not attach any significance to the f a c t .  Hygro-  hypnum luridum ssp. pseudomontanum i s not worthy of formal recognition. Plants bearing straight, julaceous or loosely imbricated, oblong ovate to broadly lanceolate leaves with more or less abruptly acuminate apicies and having strong single costae have long been treated as the variety or form julaceum of Hygrohypnum luridum. so treated by Schimper (1853, 1876), Limpricht (1924), Husnot (1894) and Roth (1905) .  These plants have been  (1904), Grout (1931), Dixon  As noted previously the costa varies  more or less continuously throughout the species.  Much weight has been  placed on straight and falcate leaves i n the recognition of subspecific taxa i n Hygrohypnum luridum.  Leaves of both shapes are frequently observed  on the same stems i n the species and thus invalidate the characters as usef u l i n this species. Muller  (1887) described Hypnum krausei from Takhim Valley, Alaska.  Grout (1931) treated the plant as synonymous with Hygrohypnum alpestre. Examination of a duplicate of the type at N Y reveals that the plant i s the common western North American form ;of Hygrohypnum luridum. Hygrohypnum luridum var. crassinervium form of Hygrohypnum smithii (Sw. i n L i l j . )  (Baur.) Podp. i s a depauperate Broth.  Hygrohypnum luridum var. e h l e i (Arn.) Wijk et Marg. i s a falcate leaved form of U. polare (Lindb.) Loesk.  For a complete discussion of the A r n e l l  plant see the treatment of H. polare.  246 Roell  (1899) used the e p i t h e t complanatum f o r what he viewed as new  v a r i e t y o f H_. l u r i d u m .  The name was used i n the absence o f a f o r m a l L a t i n  d e s c r i p t i o n n o r was a specimen c i t e d .  The name i s a nomen nudum.  L a t e r R o e l l (1915) p u b l i s h e d the name Hygrohypnum p a l u s t r e v a r . obtusatum which he c r e d i t e d t o Loeske.  The name was accompanied n e i t h e r by  a L a t i n d e s c r i p t i o n nor a c i t e d specimen and i s t h e r e f o r e a nomen nudum. Wijk e t a l . (1964) c r e d i t De N o t a r i s f o r a u t h o r i n g r e p t i l i f o r m e and L_. ambiguum.  N e i t h e r d e s c r i p t i o n s nor specimens o f such  p l a n t s have been a v a i l a b l e f o r study. Jensen  the names Limnobium  F u r t h e r , Wijk e t a l , (1962) noted t h a t  (1939) reduced the two De N o t a r i s taxa t o forms o f Hygrohypnum  luridum.  The Wijk r e f e r e n c e t o Jensen's work i s mistaken f o r Jensen c l e a r l y makes no mention o f the De N o t a r i s t a x a i n t h a t work. these  Consequently, the s t a t u s o f  two t a x a i s u n c l e a r . Wijk e t a l . (1962) noted t h a t Jensen  (1939) r e c o g n i z e d Hypnum  intextum  V o i t . i n Sturm and Hypnum r u d e r a l e v a r . i n t r i c a s t u m B r i d . as forms o f Hygrohypnum luridum.  Again,  Jensen d i d d i s c u s s these  the r e f e r e n c e t o Jensen's work i s m i s t a k e n , f o r i f t a x a i t was c l e a r l y n o t i n h i s Skandinavien  Bladmoss-  flora. Bridel of the type,  (1827) d e s c r i b e d Hypnum r u d e r a l e v a r . i n t r i c a s t u m .  the d e s c r i p t i o n i s inadequate t o c l e a r l y d e a l w i t h the name.  Simi'l-arly, the d e s c r i p t i o n o f Hypnum intextum study  In the absence  V o i t has been u n a v a i l a b l e f o r  and the taxon cannot be e f f e c t i v e l y d e a l t with.  The v a l i d i t y o f the  taxon i s a t b e s t ' d o u b t f u l . Hygrohypnum l u r i d u m v a r . pseudoehraceum Limnobium pseudochraceum by Amann i n Roth  (Roth) Podp. was d e s c r i b e d as  (1915).  Amann c h a r a c t e r i z e d the  taxon as b e i n g very s i m i l a r t o Limnobium ochraceum and noted i n p a r t i c u l a r the 'exceedingly  s t r o n g and wide c o s t a . - The  p l a n t vaguely  resembles Hygro-  hypnum ochraceum and seems o n l y t o be a c o a r s e v a r i a n t o f H. l u r i d u m . The type was examined from S-PA; Amann plumulosum.  (1928) d e s c r i b e d Hygrohypnum  subenerve (B.S.G.) Loesk. v a r .  None o f the specimens c i t e d by Amann have been a v a i l a b l e f o r  study and the d e s c r i p t i o n does n o t a l l o w f o r c l e a r r e c o g n i t i o n .  One f e a t u r e  noted by Amann which i s most unusual f o r a s p e c i e s o f Hygrohypnum b i s e r i a t e nature o f the l e a v e s . Podpera  i s the  The s t a t u s o f the t a x o n , i s , as y e t , u n c l e a r .  (1954) however, d i d reduce i t t o a form o f Hygrohypnum  luridum.  248  Fig.551 a - j . V a r i a t i o n i n the h a b i t o f m o i s t shoots o f Hygrohypnum luridum.  Scale:  249  250 F i g . 52 a - q.  V a r i a t i o n i n the l e a f shape o f Hygrohypnum luridum.  251  252 F i g . 53 a - v.  Scale:  \  \  V a r i a t i o n i n l e a f shape o f Hygrohypnum luridum.  I  1mm  I  F i g . 54  - h.  V a r i a t i o n i n the c e l l u l a r d e t a i l o f the f o l i a g e leaves  of Hygrohypnum luridum. a - b.  Leaf apices  c, f - h. d, e.  Median l e a f c e l l s  Marginal l e a f c e l l s  Scale: a,  b;  \  100um  l  255  .256 F i g . 55 a - f. V a r i a t i o n i n the a l a r c e l l s o f Hygrohypnum luridum.  Scale:  I  100"™  I  257  258  -t-  Fig. 5 6 .  Hygrohypnum luridum  259 E x s i c c a t i Examined Allen, Mosses of the Cascade Mountains, Washington # 140 as Hypnum palustre. (MICH, MIN, .NY, USA, TENN) Austin, Musci Appalachiani # 435 as Hypnum palustre. (MICH, NY, UBC, USA) Bauer, Musci europ. e t a m e r i . e x s i c c a t i . # 2090 as Hygrohypnum palustre. (NY) # 2091 as Hygrohypnum subenerve. (BRNM, NY) Musci eur. e x s i c c a t i # 1285a as Hygrohypnum palustre. (BRNM, NY','NICH) # 1285bLas Hygrohypnum palustre. (NY) Brotherus, Musci Turkestanici # 129. (H-BR, LE, MIN as Amblystegium palustre var. subsphaericarpon) Musci Turkestanici # 256 as Amblystegium palustre var. subsphaericarpon (H-BR) Musci Turkestanici # 257 as Amblystegium palustre var. subsphaericarpon (H-BR) Bryotheca Fennica # 286 as Hygrohypnum palustre (B, LE) De Notaris, Erbar. Crittogam. I t a l . #906 as Limnobium palustre. (B, NY) Ser. I I . # 3 as Limnobium subsphaericarpon. I(UC,- BM) Delogne, Les Mousses de l;Ardenne # 95 as Hypnum palustre. (C) Desmazieres, Plantes Cryptogams de France # 443 as Hypnum palustre. (G) Drummond, Musci Americani # 195 as Hypnum palustre. (CANM, MICH, NY) • Fejfoldy, F l o r a Hungarica # 423 as Hygrohypnum palustre. (B) Fleischer & Warnstorf Bryotheca Europ. meridion cent III # 298 asHygrohypnum palustre var. laxum. (C) Gravet, Bryotheca Belgica # 349 as Hypnum palustre. (G, NY) Grout, North American Musci P e r f e c t i # 118 as Hygrohypnum palustre. (MICH, LE, NY, TENN, USA,-.F) North American Musci Pleurocarpi # 362 as Hygrohypnum palustre. (MICH, MIN, MO, NY, TENN, UC, USA) Hesselbo, Bryophyta Danica Exsiccata # 333 (BP, LE; NICH, CANM) Husnot, Musci Galliae # 494 as Hypnum palustre var. subsphaericarpon. (NY; BM) # 292 as Hypnum palustre. (NY, BM) Jack, Leiner & Stizenberger, Kryptogamen Badens # 96 as Limnobium palustre. (G, NY,BM, B P ) Kerner, Flora E x s i c c a t i Austro-Hungarica # 1515 as Limnobium palustre. (G, LE, MO, UC, BP, C) Kopsch, Bryotheca Saxonica # 284 as Hygrohypnum palustre. (MICH) Limpricht, Bryotheca S i l e s i a n a # 249 as Hypnum palustre. (B, LE, NY) Lisowski, Bryotheca Polonica Fasc. IX # 266 as Hygrohypnum palustre. (CANM) Fasc. XV # 420 as Hygrohypnum palustre var. hamulosum. (B) Fasc. XXII # 593 as Hygrohypnum palustre (BP, -LE,-»'NICH) Fasc. XXXI # 819 as Hygrohypnum palustre var. tatricum. (B, LE) Fasc. XXXI # 820 as Hygrohypnum palustre var. julaceum. (B, BP, LE., NICH).  -  Fasc. LXVIII '# 1746'as Hygrohypnum var. julaceum. (LE," NICK) Macoun, Canadian Mosses # 398 as Hypnum palustre. (MICH, MO, USA, C) Canadian Musci # 341 as Limnobium palustre. (CANM)  260 # 356 as Hypnum p a l u s t r e (MIN, MO, NY, TRTC, UC USA, C) # 357 as Hypnum eugyrium. (MIN) # 493 as Hypnum c o l u m b i c o - p a l u s t r e . (MICH, MIN, NY, TRTC)• # 527 as Hypnum pseudomontanum. (TRTC) • # 535 as Hypnum d i l a t a t u m . (MO) • F l o r a Canadensis # 760 as Hypnum p a l u s t r e . (TRTC)• Matouschek, Kryptogamen e x s i c c a t i # 2195 as Hypnum p a l u s t r e . (B, LE, C,. NY) # 1587 as Hypnum p a l u s t r e v a r . subsphaericarpon (LE, C) M i g u l a , Cryptogamae Germaniae, A u s t r i a e , H e l v e t i a e e x s i c c a t i # 272 as Hypnum p a l u s t r e . (B, MICH, NY, UBC, C) G. M u l l e r , W e s t f a l i e n Laubmoose # 123 as Hypnum p a l u s t r e (BP) Mougeot, N e s t l e r & Schimper, S t i r p e s Cryptogamae Bogeso-Rhennanae # 521 as Hypnum p a l u s t r e (G, NY) P e t r a k , F l o r a Behomica e t Moraviae e x s i c c a t a I I S e r i e s L fg.# 1. # 48 as Hypnum p a l u s t r e v a r . hamulosum (LE) P i l o u s , Musci c e c h o s l o v e n i c i e x s i c c a t i # 74 as Hygrohypnum p a l u s t r e . (B) # 117 as Hygrohypnum subenerve. (G) # 237 as Hygrohypnum p a l u s t r e f o . julaceum. (B) # 279 as Hygrohypnum p a l u s t r e v a r . julaceum. (B) # 7 33 as Hygrohypnum p a l u s t r e . ( B ) # 819 as Limnobium p a l u s t r e v a r . t e n e l l u m f o . subenerve. (B,C) Rabenhorst, B r y o t h e c a europaea # 294 as Limnobium p a l u s t r e . (NY, BM, C) # 393 as Hypnum p a l u s t r e . (NY) _ # 550 as Hypnum p a l u s t r e v a r . subenerve. (G,- NY, BM C) # 1347 as Hypnum p a l u s t r e . (NY) # 1347b as Hypnum p a l u s t r e . (NY) Regensberg, b o t a n i s c h e n G e s e l l a c h a f t , F l o r a e x s i c c a t i B a v a r i c a Bryophyta # 381 as Hypnum p a l u s t r e . (B) # 384 as Hypnum subenerve. (B) S c h u l t z , herbarium mormale Cent 12, # 1183 as Hypnum p a l u s t r e (C) # 1183 b i s as Hypnum p a l u s t r e (C) S o c i e t e dauphinose # 3137 as Hypnum p a l u s t r e . (G) Wartmen & Schenk, S c h w e i z e r i s c h e Kryptogamen # 395 as Hypnum p a l u s t r e . (G, NY) W i l l i a m s , North American Mosses - Yukon Region # 767 as Hypnum p a l u s t r e . (USA) W i l s o n , Musci B r i t a n i c i # 383 as Hypnum p a l u s t r e v a r . s u b s p h a e r i c a r p o n (G, NY) Zmuda, F l o r a G a l i c i a e # 96 as Hygrohypnum s u b s p h a e r i c a r p o n (G) S e l e c t e d Specimens Examined Canada B r i t i s h Columbia Koothey N a t i o n a l Park, V e r m i l l i o n R. near V e r m i l l i o n Crum ana S c h o f i e l d 5490. (UBC) Vancouver I s l a n d : H a l b e r t 6187  Crossing;  261 N. o f F o r t N e l s o n along the A l a s k a Hwy, 58 40 N, 124 42 W; C o r r e l l 12051 as Hygrohypnum p a l u s t r e . (UBC) L y t t o n , 8 m i l e s S.; S c h o f i e l d & Boas 17815. (UBC) Armstrong; W i l s o n , March 1904. (UBC) Bowron R., 25 N. o f Bowron Lake; Boas 547. (UBC) Kamloops D i s t r i c t , Kamloops, Jamison Creek; MacFadden 3870. (TENN) Roger's Pass; Macoun 21 J u l y 1890 i n Canadian Musci # 341 (CANM) Spence's B r i d g e ; Macoun 21 May 1889 as Hypnum p a l u s t r e . (CANM) Yoho V a l l e y ; Macoun 478 as H. a l p e s t r e . (CANM) Mt. Queest, Cinnemosun Narrows, Shuswap L.; Macoun 27 J u l y 1889 as Hypnum p a l u s t r e . (CANM) ' K'oe-.tenay N a t i o n a l Park, Kimpton Creek below S i n c l a i r Pass; Crum C-65. (CANM) MacLeod's Lake; Macoun 26 June 1875. as Hypnum p a l u s t r e v a r . subjulaceum. (NY) P a v i l l i o n Lake; Boas 1960. (UBC) Queen C h a r l o t t e I s l a n d s , Moresby I s . , Kaisoun; S c h o f i e l d 31112. (UBC) Queen C h a r l o t t e I s l a n d s , Graham I s . , Van I n l e t ; S c h o f i e l d 32088. (UBC) F r u i t v a l e , K e l l e y Creek; Jamieson 5580. (UBC) M i c h e l Creek, B.C. Hwy 3 near Sparwood; Jamieson 6065. (UBC) Ten M i l e Creek, S l o c a n Lake; MacFadden 10 June 1926. (UBC) Inver Creek, between Skeena R. and snow p a t c h e s E. o f P r i n c e Rupert; S c h o f i e l d & Sharp 9/10-118a. (TENN) Wilmer, Lake o f Hanging G l a c i e r ; MacFadden 4376 as Hygrohypnum p a l u s t r e . (TENN) G a r i b a l d i P r o v i n c i a l Park, W h i s t l e r Mtn S k i A r e a ; < S c h o f i e l d & Jamieson. (UBC) Alberta' Lundbreck F a l l s , Cbleman-Kananaskis Road; Jamieson 6118. (UBC) Bearspaw Dam, S, s i d e o f Bow R. west"of C a l g a r y ; B i r d • 9039. (NY, UBC) J a s p e r N a t i o n a l Park, M i e t t e H o t s p r i n g s ; Crum & S c h o f i e l d 4703 (MICH, UAC, UBC) Bow R i v e r Watershed, E l p o c a Creek a l o n g Coleman-Kananaskis Road; B i r d ' s Glenn 13494. (UBC) Waterton N a t i o n a l Park, above Anderson Lake; Crum & S c h o f i e l d 5939. (UBC) B a n f f N a t i o n a l Park, L o u i s e Creek; Jamieson 6190. (UBC) J a s p e r N a t i o n a l Park, Sunwapta Pass Campground; B i r d 5232. (ALTA) Manitoba The Rock, on the Hays R i v e r ; Scoggan 75. (UBC) C h u r c h i l l , Eskimo P o i n t ; Crum & S c h o f i e l d 7155. (MICH) Lower Hays R i v e r , York F a c t o r y Region, Hudson Bay Lowlands; R i t c h i e 4130. (CANM) Herb Lake V i l l a g e , Wekusko Lake, N. E. o f the Pas; Scoggan 102. (CANM) Hansen's Creek, E. o f Rennie on Hwy 44; B i r d 3286. (CANM) :  262  Saskatchewan S w i f t C u r r e n t ; Macoun 6 Sept 1880 as Hypnum p a l u s t r e . (CANM) Ontario Grey Co., I n d i an F a l l s near Balmy Beach; Crum 11352. (TRTC) Bruce Co., Sauble F a l l s ; Moxley 24 May 1936. (5ERTC) P e e l Co., C r e d i t F o r k s ; C a i n 1413. (TRTC) Kenora D i s t r i c t , Cedar Lake, N. o f V e r n i l l i o n Bay; C a i n 4914. (TRTC) Owen Sound; Macoun 28 J u l y 1871 i n F l o r a Canadensis as Hypnum palustre. (TRTC) Thunder Bay D i s t r i c t , l a k e shore N. End o f Simpson I s ; Garton 6280. (CANM) Algoma D i s t r i c t , N. end o f Bachawana Bay; Sharp CM-631. (UBC) E l o r a , Grand R i v e r ; Moxley 7 June 1933. (TENN) M a n i t o u l i n I s l a n d , shores o f Manitou Lake: Hermann 16044. (USA) Quebec A n t i c o s t i I s l a n d , J u p i t e r R i v e r ; Macoun 20 Aug. 1883. (UBC) James Bay, Rupert House; Lepage 18969. (USA) Bonaventure Co., N o u v e l l e R i v e r , S t . Jean 1 E v a n g e l i s t e C o l i n s & F e r n a u l d 3313-AB. (MICH) Lake M i s t a s s i h i , New Quebec; Lepage 4328. (TRTC) S t . Rimon Rimouski; Lepage 2828. (TRTC) Pare de l a G a s p e s i i , S. o f S t . Anne-des-Monts; I r e l a n d 11170. (CANM) Montmorency R i v e r , F a i r y Creek; Macoun 30 June 1905. (CANM) New Brunswick V i c t o r i a Co., Grand F a l l s ; Habeeb 1 J u l y 1943. (UBC, F) A l b e r t Co., Fundy N a t i o n a l Park; I r e l a n d 11565. (UBC) Queens Co., Moser 1889. (CANM) Newfoundland Bay o f I s l a n d s , B r i c h y Cove; Waghorne 23. (MO). Lomond R. a t i t s i n t e r s e c t i o n w i t h P r o v i n c i a l highway 44. Norris 4584. (HSC) C a t a r a c t s P r o v i n c i a l Park, 7 km NW o f C o l i n e t ; Weber 534 (NFLD) Labrador C h u r c h i l l F a l l s , B r i d g e Camp Area; B r a s s a r d 5178. (NFLD) L'Anse au C l a i r ; Waghorne 8 Sept 1894.. (NY) Yukon Lake LeBarge; W i l l i a m s 765. (MO) R a n c h e r i a R i v e r , 60°10'N, 130°08'W.; C o r e l l 12160. (MICH) Dawson C i t y D i s t r i c t , Hunker Creek; Macoun 22 J u l y 1902. (CANM) Haines Highway, 83 m i l e s NW o f Haines, A l a s k a ; Hermann 21848. (CANM) F r a n c i s Lake, on Money Creek; Jamieson 1976. (UBC) North West T e r r i t o r i e s . E l l e s m e r e I s l a n d , Head o f Tanquary F j o r d ; B r a s s a r d 1614. (CANM) V i c t o r i a I s l a n d , Kaskyak R i v e r , Minto I n l e t ; Wynne 18860. (NY) G r e a t Bear Lake, E. end o f McTavish Arm, v i c i n i t y o f E l d o r a d o Mine; S t e e r e 10070a (NY) Campbell Lake, S c o t t e r 9742 (C, as Hygrohypnum a l p e s t r e . ) United States Alaska Skagway, W i l l i a m s 27 Aug. 1899 i n North American Mosses-Mukon Region # 767. (USA) 1  ;  263 Mount McKinley Region, Garner, B r a g s t a d Creek; S h e r r a r d C-18. (USA) Anchorage-Seward Highway, McHugh Creek Campground; Sharp 14 June 1971. )TENN) Brooks Range, S i k - S i k - P u k R i v e r , Gorgeous Gorge, 30 m i l e s W. o f A n a k t u v i k Pass; Spatzman 15 Aug. 1951. (NY) T a k h i n V a l l e y ; Krause 20 J u l y 1882. (NY, as Hypnum k r a u s e i ; isotype ) Brooks Range, S. o f Cape Thompson, Ogotoruk Creek; S t e e r e 63-652. (NY) Brooke Range, E n d i c o t t Mountains, SW o f Chandler Lake; S t e e r e 18333. (NY) Brooks Range, F r a n k l i n Mountains, . Schrader L. P e t e r s L. a r e a ; S t e e r e 18926a. (NY) Washington King Co., Green R i v e r Gorge, E. o f B l a c k Diamond; I r e l a n d 9606. . (CANM) Upper V a l l e y o f the N e a q u a l l y ; A l l e n 28 June 1898 i n Mosses o f t h e Cascade Mountains, Washington # 140. (USA, CANM) Olympic P e n i n s u l a , Mouth o f Sekim R i v e r ; S v i h l a 703. (MICH) California Sequoia N a t i o n a l Park, H a l s t e a d T r a i l ; Degener & P e i l e r 16914. (MO) New Mexico Hermit's Peak; Arsene 18479. (USA) Cumber's Pass; V r e e l a n d 31 Aug. 1900. (NY) San M i g u e l Co., Pecos V a l l e y Holy Ghost Creek; S t u d h a l t e r & Man 3089 (COLO) Taos Co., Angosturo Creek, T r e s R i t o s ; S t u d h a l t e r & Man 3258 (COLO) Arizona Coconine Co., West Fork o f Oak Creek Canyon; Dunn 417. (?) Idaho C u s t e r Co., Sawtooth W i l d e r n e s s A r e a , Forks o f Baron Creek; Morton 8537. (USA) K o o t e n a i Co., Lake Pend d ' O r e i l l e ; L e i b e r g June 1889. (NY) Utah S a l t Lake Co., B r i g h t o n ; Darker 5897. (MICH.) Marysvale, F a l l s above B u l l i o n Creek; Jones 5899. (MO, UC) Montana L i n c o l n Co., K o o t e n a i R i v e r , K o o t e n a i F a l l s Overlook, 7 m i l e s E. o f Troy; Hermann 22481. (TENN) F l a t h e a d N a t i o n a l F o r e s t , Echo Lake; Kaufman J u l y 1928. (TENN) Park Co., S i l v e r Gate, Wyoming Creek; Conard 25 Aug. 1953. (CANM, TRTC) Columbia F a l l s ; W i l l i a m s 24 J u l y 1895. (C, MO) S t . I g n a t i u s , near Mary's Lake; F r y e 2137. (CANM) G l a c i e r N a t i o n a l Park, SW shore o f Bowman Lake; Hermann 18017. (CANM) Carbon Co., SW o f Red Lodge, T r a i l t o F a l l s o f B a s i n Creek; Lawton 2156. (CANM) Wyoming Carbon Co., E l k Mtn,; Gooding 21 Aug." 1901. (MIN, USA)  264 Big.Horn Mountains, G r a n i t e CrEek; Whitehouse 25916. (USA) Albany Co., L i b b y Creek; P o r t e r 1536. (TENN)' Park Bo., Absaroka Range; K i e n e r 4047. (MICH) Y e l l o w s t o n e R i v e r ; Knowlton 10 Aug. 1888. (USA)Albany Co., M e d i c i n e Bow Mountains, 9 m i l e s W. o f C e n t i n a l Hermann 17692. (CANM) B e a r t o o t h Mountains; shores o f B e a r t o o t h Lake; Welch 16600. (CANM) Colorado La P l a t a R i v e r ; Baker, E a r l e & Tacey 20. (USA) Manitou, Ruxton Creek; Jewett 18 J u l y 1910. (USA) • C o l l e c t e d mostly w i t h i n 100 m i l e s o f Canon C i t y ; Brandegee ' 1874 - 1878. (USA) Gunnison Co., v i c i n i t y o f G o t h i c , E a s t R i v e r V a l l e y ; Welch 9096. L a r i m e r Co., Rochy Mountain N a t i o n a l Park, 7 m i l e s SE Chambers Lake; Hermann 26783. (UBC) Minnesota Cook Co., v i c i n i t y o f Grand M a r a i s ; H o l z i n g e r 16 J u l y - 7 Aug. 1902. (MIN) Cook Co., Schroeder, C r o s s R i v e r Gorge; O l s o n 761. (CANM) Michigan Ontanogon Co., P o r c u p i n e Mountains S t a t e Park, T r a i l t o Bucks h o t C a b i n ; I r e l a n d 5214. (USA) Keweenaw Co., G r a t i o t R., 2 m i l e s W. o f C l i f f ; Hermann 16389 <USA) A l g e r Co., Miner's F a l l s ; Sharp 22 J u l y 1955. (USA) I s l e Royale N a t i o n a l Park, S c o v i l l e P o i n t ; Van Dyk Aug. 1955. (USA) Cheboygan Co., M i l l Creek; V i t t 304. (ALTA) Marquette Go.,.Huron Mountains, C l i f f R i v e r , N i c h o l s Aug. 1934. (NY) Wisconsin B a y f i e l d Co., Near H e r b s t e r on Lake S u p e r i o r ; Cheney 6913. (TRTC) Ohio Ottawa Co., South Bass I s l a n d ; Schnooberger 3586. (USA) New York Watkins; A u s t i n 17 J u l y 1874 (NY) Herkimer Co., N e w v i l l e ; A u s t i n 1868. (NY) Cayuga Lake B a s i n , F o o t o f I s l a n d F a l l , F a l l Creek; Dudley 25 Nov. 1882. (NY) New J e r s e y . Sussex Co., West Vernon; A u s t i n Sept 1867. (NY) Vermont C a l e d o n i a Co., Peacham; B l a n c h a r d 5 Nov. 1885. (USA) New Hampshire Bekknap Co., G i l f o r d , Mt. Belknap;. C a r t e r 5 Sept 1904. (USA) West V i r g i n i a Mingo; Gray M-1408. (TENN) U f f i n g t o n ; H e r r o d 20 June 1930. (USA) Guatamala Dept. o f Huehuetenango, T a q u i a ; V o g e l B-9009. (NY, TENN) N  :  265 Greenland Qroge P e n i n s u l a , I n u k a v s a i t F j o r d , S a g d l i a r u s e q ; Holmen 17054. (NY) Scoresby Land, M e s t e r s v i g , Blyminen; Holmen 18045. (NY) Great B r i t a i n Scotland S u t h e r l a n d Co., near Inchnadamph; Crundwell 12 June 1951.(G) Wales B l a c k Mountain, Brechnoch, N s l o p e s o f Darenlwyd; L i f l i n 18 May 1959. (FLAS) Northern I r e l a n d B e l f a s t ; Stewart 12 May 18-7. (NY) Faroes StjzimjzJ, Gjoven a t Vestmannahavn; Jensen 9 June 1896. (NY) Iceland S . - I c e l . , R a n g a r v a l l a s j s t a , D r a n g s h l i d ; Johannson 14 J u l y 1965. (FLAS) Denmark J u t l a n d , b e i Bjornsholm; Jensen i n B r y o t h e c a europaea # 393. (NY) Zealand, Lake Fureso; Hesselbo 8 J u l y 1903 i n Bryophyta Danica ' E x s i c c a t i # 333. (S-PA) Bornholm, Bobea; Jensen 2-4-1882. (H) Netherlands. W. E . p l a s ; Wasschen 29 June 1956. (GRO) Dordrecht; van d e r Sande L a c o s t e . (GRO) Belgium Dinant (Namur); Gravet i n B r y o t h e c a B e l g i c a # 349. (G) France Chambery Sabaudiae, Cascade de Coux; P a r i s 10 J u l y 1861 as Hypnum p a l u s t r e . i n B r y o t h e c a europaea # 550. (G) V i l l a r e de.Lans, I s e r e ; Ravaud IN S o c i e t e dauphinose # 3137. as Hypnum p a l u s t r e . (G) Haute S a v o i e , F o r e t de l a g l a c i e r e au Mont Brezon; B e r n e t 29 J u l y 1883. (G, S-PA) Blanche, au Mont B l a n c ; l e g . ?. (B, a c c e s s i o n # 7720/345) 7972 Haute Garonne, D e v e s o i r du Lac V e r t p r e s Luchon; Husnot as Musci G a l l i a e # 592. (S-PA) Montauban; Lange 7 Sept 1857. (S-PA) Germany A l l g a u , H i n d e l a n g , W i l d b a c h t a b e l b e i Oberdorf; Bornmuller 26 Aug. 1922. (B) Krahenberg; Winter A p r i l 1902. (B) F i c h t e l g e b e b i r g e , b e i d e r Ruine W a l d s t e i n ; F a m i l e r & Schwab Aug 1903 as F l o r a E x s i c c a t i B a v a r i c a ; Bryophyta # 384. (B) Mark Brandenburg, Potsdam; Benkert Nov. 1960. (B) R u i s s e , Walkenreid; Loeske 1/6/1903. (B) Ruhland; Moenkemeyer J u l i 1910. (NY) Waren b e i Mwritz; S t r u c k . (S-PA) Bayern, P a r t e n k i r c h e n ; Suse March 1916. (S-PA) Wurtemburg, A l p i r s b a c h ; Walde 1897. (S-PA) R h e i n l a n d , M e r z i g a. d. Saar; Winter. (S-PA) R h e i n l a n d , Bensberg (near K o l n ) ; S c h l u e g e r . (S-PA)  266  H e s s i n , E b e r s t a d t ; l e g . ?. (S-PA) Baden, H e i d l e b e r g ; Arschoug Mai 1861 (S-PA) T h u r i n g e r , Jena; R o e l l 27 J u l y 1871. (S-PA) Austria N e i d e r O e s t e r e i c h , S c h r a t t e n s t e i n , S t i x t e n s t e i n ; Redinger 28 June 1931. • . N e i d e r O e s t e r e i c h , Schneeberg, A l p e l l e i t e n zwischen BaumgartnerhausLackerboden; Redinger 19 J u l y 1931. (B) D a c h s t e i n g e b e i t e s ; Morton 1923. (B) T i r o l i a c e n t r a l i s , i n v a l l e G s c h n i t z e l ; Kerner i n F l o r a E x s i c c a t i Austro-Hungarica' # 1515. V o r a l b e r g , W a l s c a t a l A u f s b e r g zwei F l e u ; B o r n m u l l e r 30 J u l y 1936. (B). N e i d e r O e s t e r e i c h , Obersee b e i Lunz; Redinger 16 June 1930. (B) Switzerland S i l v e r e t t a Gruppe; B o r n m u l l e r 26 J u l y 193 2. (B) In den Arve b e i Genf; B e r n e t J u l i 1863 i n S c h w e i z e r i s c h e K r y p t o gamen # 395. (G) V i a . M a l a ; Reuter Aug. 1854. (G) N e u c h a t e l ; Bruch. (S-PA) Z u r i c h , Huntwangen; Frymann Feb 1893. (S-PA) Zermatt; Husnot i n Musci G a l l i a e # 292 (S-PA)Gemhi Pass; R e n i s c h . (S-EA) Lugano, Monte Are; K i n d b e r g 27 June 1892. (S-PA) Italy C a r r a v e l l a ( G a n z e g l i o ) ; F l e i s c h e r 23 J u l y 1895. (B) D i n t o r i S e r r a v i l l e a l i a S c r i v i a ; F e r r a r i 1862. i n E r b a r , C r i t t o g a m . I t a l . # 906. (B) L i g u r i e n , im Bogo b e i R a p a l l o auf k a l k ; F l e i s c h e r Aug. 1896. i n B r y o t h e c a Europ. m e r i d i o n Cent. I l l # 298. (B) T a r v i s i n a , ad flumen, P l a v i n o ; Saccardo. (S-PA) Lago d'Como; Leg. ?. (S-PA) Czechoslovakia Moravia, M a r i e n t a l ad therman T e p l i t z p r . Mahr-Weisskirchen; Matouschek i n Kryptogamen e x s i c c a t i # 2195. (B) S l o v a k i a , Mont Mala F a t r a , prope Krasnany; P i l o u s May 1935. i n -Musci c e c h o s l o v e n i c i e x s i c c a t i # 74. (B) Bohemia, Trutnov, prope Volanov; P i l o u s June 1949 i n Musci c e c h o s l o v e n i c i e x s i c c a t i # 819. (B) S l o v a k i a , Montes B e l s k e T a t r y , prope Zdearska V i d i a ; P i l o u s Aug. 1947. i n Musci c e c h o s l o v e n i c i e x s i c c a t i # 279. (B) Marienbad (=Marianske L a z n e ) ; Suse 11/9/1915. (S-PA) Poland T a t r y Zachonie, Montes T a t r a O c c , v a l l i s D o l i n a ; L i s o w s k i 8 May 1956 i n B r y o t h e c a P o l o n i c a F a s c . XXXI # 820. (B) Mons P i l s k o , ( B e s k i d i A l t i ) , t o r r e n t i s Cebula; L i s o w s k i 5 May 1957. i n B r y o t h e c a Polonica.'Fasc. XXII # 593. (S-PA) G l e i w i t z i n O b e r b f l n f i n u (? s p e l l i n g ) ; Kabbatk. (S-PA) E l b i n g , Vogelsanger Wald, Kalmuss 1895. (S-PA) Hungary Comit. Borsod., v a l l i s Garadna-volgy supra Omassa; Boros 16 O c t . 1959. (B) Comit. Z a l a , l a c u s B a l t o n ad Tihany; Boros 4 A p r i l 1957. (B)  267 Comit. Esztergom, v a l l i s Ramszaka dek prope Domos; Boros 7 Aug. 19^49. (BP) Comit Veszprem, v a l l i s Zurokvolgy prope pag Bakouykuti; V a j d a 30 May 1954. (BP) Comit, P s s t . , montes Nagyszensas prope pag Nagykovasci; V a j d a 16 O c t . 1955. (BP) Comit. Heves, montes Tasko prope pag Parad; V a j d a 27 May 1952. (BP) Rumania C a r p a t , merid., Comit, C s i h , v a l l i s Uzvolgy prope pag C s i k s z e n t mahtom; V a j d a 28 Aug. 1970. (BP) C a r p a t , merid., v a l l i s R a d e s u l u i prope Gura A p i , montes R e t y e z a t ; V a j d a 12 J u l y 1968. (BP) Banat., r i v i Csernapatak, ad H e t f o r r a s prope pag B a i l e Herculane; V a j d a 7 Aug. 1967. (BP) Turkey Prov. R i s e , D i s t r i c t Hemsin, Ortakoy-Cat; Davis 21311. (C) Norway Nordland, S o r f o l d e n ; Nyman 19 Aug. 1899. (S-PA) • Troms, f j l k e Tronso; A r n e l l 11 J u l u 1891. (S-PA) . S o r - T r o n d l a g , Trondherim; A r n e l l J u n i 1869. (S-PA). T r o n d f j e l d e t ; Kindberg 22 J u l y 1897. (S-PA) . Gudbrandsdalen; Z e t t e r s t e d t ' 2 5 June 1858. (S-PA) Dovre; Hartman 1872. (S-PA) Hordaland, Hardangervidda s t r a s o om Dyrskar v i d vagen m e l l a n O s l o och Haugesund; S t r i d v a l l 4 Aug. 1970. (S-PA) Sweden Skane, V i n s l o f sn i baken N v i d landvagen 2 km USV on Lommarp; Johansson 10/7/1916. (S-PA) B l e c k i n g e , S i s s e b a c k , f r a s t e u ; Medelius 4/9/1922. (S-PA) H a l l a n d , G K a r u j i s k y r k a ; F l o r i n 15 June 1923. (S-PA) Smaland, Jonkoping Torpa; Arven 15 Sept 1896. (S-PA) G o t l a n d , Lummalunda; Jaederholm 1 J u l y 1923. (S-PA) O s t e r g o t l a n d , Omberg; Mosen 12 Sept 1871. (S-PA) V a s t e r g o t l a n d , Skovde, Havstena, Tomtangen; Hulphers J u l i 1930. (S-PA) , Bohuslan, F i s h e b a c k s k i l ogavdun; Hulphers Aug. 1927. (S-PA) - S a l s l a n d ; Bergstrom 16 Sept 1913. (S-PA) Narke, Karlsluund;.Weldheim 26 May 1923. (S-PA) Sodermanland, Vardinge sn Sjunda; F l o r i n 4/9/1927. (S-PA) Uppland, U p s a l a , Krasnbo; Jaederholm' 1888. (S-PA) Vastmanland, G r y t h y t t e S k a t v i k e n ; Stenholm 6/8/1925. (S-PA) Varmland, P e r s b e r g ; L a r s s o n 9/6/1914. (S-PA) D a l a r n e , . p a r Boda, Osmundsberg, A r n e l l 17 Aug. 1896. (S-PA) G a s t r i k l a n d , V a l b o ; Hartman June 1858. (S-PA) Medepad, Torp sn J o h a n n i s b e r g ; Huss 1882. (S-PA) H a r j e d a l e n , paroec Tannas L j u s n e d a l ; Ostmann 19 June 1905. (S-PA) Jamtland, O f f e r d a l s sn H a l l b e r g e t ; F l o r i n 25 June 1944. (S-PA) L y c k s e l e Lappmark, Tarna; Hulphers 15 July-1940. (S-PA) Angermanland, p a r T a s j o T a s j o b e r g e t ; A r n e l l 14 J u l y 1894. (S-PA) L u l e Lappmark, Jikkmokk; Nyman Aug. 1893. (S-PA) L a p o n i a T o r n e s n i s , Roukoajokk ( J u k k a s j a r v i ) ; Jaederholm 27 J u l y 1912. (S-PA) Lapponia T o r n e n s i s , A b i s k o j o k k ; Jaederholm 12 J u l y 1917. (S-PA)  268 Soviet Union Caucasus, i n f l . Rion pr. E l t s e v i ; Brotherus 10 July 1877. (NY) A l t a i Mountains, Katung pr. Techeposch; Grano. (H-BR) S i b e r i a , Yamarovka; Mikkvo 24 June 1904. (H-BR) Tomsk D i s t r i c t , Tomsk Junge; Matoejeff 12 July 1912. (H-BR) S i b e r i a , Jenesei, Hantai; Sahlberg 1876. (H-BR) S i b e r i a , Jenesei, Tostoinis; A r n e l l . (H-BR) Siberia, Lena, Kamachsur.; Nilsson-Ehle 6 Aug 1898. (C,S-PA) Sib e r i a , Lena, Balagnach; Nilsson-Ehle 15 July 1898. (C, LE) Siberia, Jenesei, Plachino; A r n e l l 22 July 1876. (C, S-PA) Karelia ladogensis, Par Svistamo, Kaaherjoki Laynavaara; Kotilainen 22 Sept 1926. (H, S-PA) Prov. Uchta, Archangelsk; Liskendrath June 1885. (H) Prov. Perm; K e l l e r . (H) Rossia Bor. porp Vologda; Dohle 1/7 1907'. (H) Irkutsk D i s t r i c t , Verzholensk i n v a l l e f l . Tachikan; Kusenezou Jenesei D i s t r i c t , Turuchansch Chantaish; Kusnezow a t Reverdatto 7 July 1914. (H-BR) Pamir i n angustatis Chargusti; Paulson 4/9/ 1898. (H-BR) A l t a i Tschemal; Verestchagin. (H-BR) A l t a i , ad f l . Mena; Krylow. (H-BR) Amur, Tukuringra an Ufer des Saja; Kuseneva 9/7/ 1915. (H-BR) Jakutsk, Jenkovshi spitze; Kusenova. (H-BR) Semiryelschensk, Almatinka minor props Vjernoyi; Regal 20/3/ 1876. (H-BR) Terskei Alatau, f l . Dschuka; Brotherus 5 Aug 1896 i n Musci Turkenstanici #130. (H-BR) Kungei Alatau, as fontes f l . Koissu; Brotherus Aug 1896 i n Musci Turkenstanici # 256. (H-BR) Thian Schan (=Tien Shan) Mtns., ad. f l . Narinhol (=Naryn) R.; Brotherus 25 July 1896. (H-BR) Siberia, Ust ' Kut, i n v a l l e flum Lena; Nilsson-Ehle 4 June 1898. (S-PA) Jenesei, Nischnaja Tunguska; A r n e l l 14 July 1876. (S-PA) Kashmir Karakorum Range, upper end of the Hushe Valley, Atosar Valley; Webster & Nasir 6090. (TENN) Lind Valley near Sonamarg; Duthie 17 Aug 1893. (H-NR) B a l l i s t o n , by water course W. of Dras; Duthie 24 Aug 1893. (H-BR) Gulmarg; Duthie 8 Jan 1892. (H-BR) Indian Uttar Pradesh, Kimaun, K u l t i Valley; Duthie 7 Sept 1894. (H-BR) Tibet Kargall, Darass Valley; Khan 26 June 1901. (H-BR) Finland Kuusamo, r i v u l o Kulmakkapuro; Hallstrom 18 July 1911 i n Bryotheca Fennica # 286. (B, LE) China Interior p r o v i n c i a , Schen-si sept, i n medio monte Kuan-tou; G i r a l d i 1894. (G) Yunnan Prov., head waters of the Mekong R.; Handel-Mazetti #1565. (H-BR) Setchwan Prov,, In montium Daliang-schan ( t e r r i t o r i Lolo), ad orientum urbis Ningyuen regione calide temperata ad rivulum prope vicum Sikwai; Handel-Mazetti # 391. (H-BR)  Japan Tshihau Prov. Y u b a r i Mtn; Yasuda 518. (H-BR) Spitsbergen, King's By D i s t r i c t ; A r n e l l & Mortensson 26 J u l y  1956  270 Hygrophypnum a l p e s t r e (Hedw) Loesk., Verh. Bot. Ver. Brandenburg 46: 198.  1905.  Neotype:  " n o l l e " ; from Swartz herbarium:  (S-PA).  annotated by Nyholm  . '  Hypnum r i v u l a r e Sw., Disp. s y s t . muse. Suec.  1799 - not  seen - c i t e d from Hedw. 1801 & Broth, 1924 nom.; i l l e g . horn. Hypnum a l p e s t r e Hedw., Spec. Muse. 247. 64f. 1-4. 1801 Hypnum molle var. alpestre (Hedw.) Hamp., F l o r a 20:274. 1837. Limnobium a l p e s t r e (Hedw.) Schimp., B r y o l . Eur. 6:68. 1853. Figure 577 not included. Amblystegium r i v u l a r e (Sw.) Lindb, Musci Scand.  33. 1879.  nom. i l l e g . C a l l i e r g o n alpestre (Hedw.) Kindb., Canad. Rec. Sc. 6(2): 72. 1894. Hygrophypnum r i v u l a r e (Sw.) Broth., Laubm. Fennosk. 500. 1923.  nom. i l l e g .  Plants u s u a l l y s o f t , r a r e l y somewhat s t i f f , u s u a l l y forming l o o s e l y woven patches o r t u f t s , often e a s i l y fragmenting.  Color v a r i a b l e , yellow  or yellow-green with or without rusty mottling above, older extremities d i r t y brown, reddish-brown or reddish-black, r a r e l y green or yellowgreen throughout.  Stems (1) 2-4 (9) cm long, p r o s t r a t e or ascending near  the apex; stems f o l i o s e throughout o r denuded toward the base or clothed i n p e r s i s t e n t l e a f bases, frequently clogged with s i l t . i r r e g u l a r , branches u s u a l l y ascending.  Branching  Stem cross sections r e v e a l i n g  2-3 concentric rows of brownish, t h i c k - w a l l e d c o r t i c a l c e l l s ; medullary c e l l s l a r g e r , thinner walled, h y a l i n e , becoming d i s c o l o r e d and sometimes incrassate w i t h age; c e n t r a l strand w e l l developed.  Rhizoids  infrequent,  271  a r i s i n g on young stem or branch segments from the bases of v e n t r a l leaves, generally absent or obscured i n the older e x t r e m i t i e s . Leaves v a r i a b l e , c l o s e l y spaced and julaceous, sometimes spreading, e s p e c i a l l y i n some young stem or branch t i p s , spreading leaves sometimes shrinking l a t e r a l l y and t w i s t i n g s l i g h t l y upon d r y i n g , otherwise changing l i t t l e from the wet to the dry c o n d i t i o n ; leaves oblong to oblong lanceol a t e ; s t r a i g h t , never f a l c a t e ; (0.9) 1.4-1.6 (1.9) mm long x (0.4) 0.6-0.7 (0.9) mm wide; deeply concave, almost boat shaped i n the apex, g e n e r a l l y tumid julaceous; apex acute to s l i g h t l y obtuse, becoming r e f l e x e d as a squarrose a p i c u l u s , apiculus g e n e r a l l y worn away i n older leaves; margins e n t i r e or uneven i n the apex, narrowly recurved, e s p e c i a l l y i n the upper 1/3 and the apex; s h o r t l y and narrowly decurrent a t the p o i n t of i n s e r t i o n costa v a r i a b l e , u s u a l l y long and double with one arm reaching midleaf, l e s s frequently s i n g l e to beyond midleaf or short and double, i n a l l cases slender and f a i n t . A e r o l a t i o n uniform, median l e a f c e l l s long l i n e a r flexuose  (55)  65-90 (105) um long x 5-6 um wide, a p i c a l c e l l s vary w i t h the shape of the apex, u s u a l l y longer i n acute  apices  c e l l s toward the base becoming  shorter, somewhat wider and more i n c r a s s a t e ; p i t s v a r i a b l e , few t o none; a l a r c e l l s s l i g h t l y enlarged, rounded quadrate or short rectangular, t h i c k - w a l l e d c e l l s , becoming d i s c o l o r e d with age, sometimes excavated. Plants autoicous; p e r i g o n i a l leaves ovate with a small t i p , concave imbricate, ecostate; outer and middle p e r i c h a e t i c a l leaves ovate to ovate l a n c e o l a t e , ecostate, squarrose i n the upper h a l f ; inner p e r i c h a e t i a l leaves l i n e a r lanceolate to l a n c e o l a t e , to 2.  mm long, s t r a i g h t ,  p l i c a t e , margins e n t i r e or sometimes toothed i n the apex; costa f a i n t and slender, double to midleaf, s i n g l e to midleaf or absent.  272 Seta 7 t o 16 nun l o n g , smooth, r e d d i s h brown; c a p s u l e t y p i c a l f o r the genus. Peristome cells;  t y p i c a l f o r the genus; a n n u l u s composed o f two rows o f  inner peristome  w i t h 3, sometimes 2, f i n e l y p a p i l l o s e ,  slightly  a p p e n d i c u l a t e c i l i a between a d j a c e n t segments; spores 10 t o 18  um.,  u s u a l l y 12 t o 16, f i n e l y p a p i l l o s e , dusky: y e l l o w . Among the s p e c i e s o f Hygrohypnum, H. a l p e s t r e i s most e a s i l y d i a g nosed by i t s narrowly r e c u r v e d , squarrose obscured  l e a f apex  ( F i g . '57h).  i n s l i d e p r e p a r a t i o n s f o r the compound microscope,  Tough  the squarrose  apex i s e a s i l y seen w i t h a d i s s e c t i n g microscope o r a hand l e n s . examine young stem o r branch  l e a v e s f o r the squarrose  One  must  apex f o r the apex  i s f r e q u e n t l y worn away i n o l d e r l e a v e s . Other u s e f u l c h a r a c t e r s a r e the boat shaped c o n c a v i t y o f the l e a v e s , the oblong  to oblong  lanceolate leaves  s l e n d e r and l o n g double  ( F i g . 57  a - g,..i -;.k) the normally  c o s t a e , the u n i f o r m l y l o n g median l e a f  ( F i g . 58 c and d) and the rounded quadrate  alar cells  ( F i g . 58 f - h ) .  The l e a f c o n c a v i t y i s v e r y deep, b u t does n o t extend to the apex.  cells  a l l the way  I n s t e a d , t h e c o n c a v i t y ends s h o r t l y below the apex, thus  d e l i m i n t i n g a p l a n a r zone j u s t below the squarrose a p i c u l u s ( F i g . 57e). The upper 1/3  t o 1/2 o f the l e a f thus appears boat shaped.  Normally ( F i g . 57h) .  the stems o f H_. a l p e s t r e a r e s t r o n g l y  O c c a s i o n a l l y the l e a v e s a r e l o o s e l y  tumid-julaceous  spreading.  Of a l l the Hygrohypjia, H_. a l p e s t r e remarkable e x h i b i t s a v e r y u n i f o r m median l e a f a r e o l a t i o n .  The median l e a f c e l l s do n o t have  u n u s u a l l y s h o r t o r u n u s u a l l y l o n g c e l l s s c a t t e r e d through an o t h e r w i s e u n i f o r m median a r e l a t i o n as do some s p e c i e s . In the p a s t , Hygrohypnum  a l p e s t r e has been confused w i t h H_. m o l l e ,  273  H.  l u r i d u m and H. eugryium.  For a comparison o f H. a l p e s t r e and  m o l l e , see the d i s c u s s i o n under the l a t t e r  species.  H.  The f o l l o w i n g  c h a r t w i l l p o i n t out the p e r t i n e n t d i f f e r e n c e s between the  remaining  three species.  H.  alpestre  H_. l u r i d u m  H_. eugyrium  Leaf shape  Oblong t o oblong l a n c e o late.  Lanceolate to oblong l a n c e o late .  Lanceolate to oblong l a n c e o l a t e  Leaf apex  Acute, w i t h a squarrose, r e curved a p i c u l u s .  Acute, s t r a i g h t , secondarily a p i c u l a t e by inrolled margins.  Acute, s t r a i g h t , one o r b o t h margins i n r o l l e d .  Leaf  Straight.  S t r a i g h t or falcate.  Falcate or straight.  Concavity  Deeply concave, b o a t shaped i n the apex.  Plane to s h a l l o w l y concave.  Concave.  Leaf  Tumid-julaceous throughout.  Julaceous to spreading.  Imbricate t o spreading.  Usually slender, l o n g and double.  Strong and single or short and double, o f equal frequency.  Usually short and double f a i n t .  Quadrate to slightly enlarged.  Quadrate s h o r t , rectangular, r a r e l y enlarged or i n f l a t e d .  Quadrate t o s h o r t rectangular, inflated.  symmetry  attitude  Costa  Alar  cells  The o n l y forms o f Hygrohypnum l u r i d u m w i t h which H. a l p e s t r e be e a s i l y confused a r e the s t r i g h t l e a v e d , j u l a c e o u s ones. two  Here, the  t a x a can be r e a d i l y s e p a r a t e d by the squarrose l e a f apex o f  alpestre.  can  H.  274  On rare occasions a few workers have confused E. eugryrium w i t h H. a l p e s t r e .  These two taxa may be separated most e a s i l y by the squar-  rose l e a f apex of II. a l p e s t r e .  I t i s a l s o worth noting t h a t the known  d i s t r i b u t i o n s of the two taxa are not sympatric. In the type d e s c r i p t i o n Hedwig (1801) based Hypnum a l p e s t r e on a Swartz specimen from a l p i n e Norway. imens of  At the present time there are no spec-  Hypnum a l p e s t r e i n the Hewwig herbarium.  At S-PA there are  two specimens o f Hygrohypnum a l p e s t r e i n the Swartz herbarium, both of which are e q u a l l y r e p r e s e n t a t i v e of the species as understood here. One specimen bears no other data other than the p e n c i l e d name "molle" and a Nyholm annontation l a b e l c a l l i n g the p l a n t H. a l p e s t r e .  The second  specimen i s mounted on a very f i b r o u s paper and bears a few notes concerning other l i t e r a t u r e c i t a t i o n s .  This second specimen i s selected  as the neotype. Warnstorf  (1915) described Hygrohypnum a l p e s t r e var. s c o r p i o i d e s .  The type of t h i s v a r i e t y has not been a v a i l a b l e f o r examination.  The  d e s c r i p t i o n p o i n t s out three characters that are unusual f o r the species, i f i n f a c t the v a r i e t y i s H_. a l p e s t r e .  The leaves of the v a r i e t y are  described as being up t o 2.5 mm. long.  This c l e a r l y exceeds the l e a f  s i z e f o r the species as determined by the present study.  The leaves of  the v a r i e t y are further described as being "secundo-imbricata." leaves of the species have never been observed t o be secund.  The  In other  species, that leaves be s t r a i g h t , secund o r f a l c a t e has been shown to be varibel.  The stem and branch t i p s are always s t r a i g h t , though they may  be " c a t k i n - l i k e " o r hakenformig.  The status of the v a r i e t y i s unclear  and c e r t a i n r e s e r v a t i o n s are j u s t i f i e d .  275 F i g . 57 a - k.  V a r i a t i o n i n the l e a f shape and the h a b i t o f the  shoot o f Hygrohypnum a l p e s t r e . a-g, h.  i - k . V a r i a t i o n i n l e a f shape,  The h a b i t o f a moist shoot.  Scale: a - g, i - k; h;  I  1mm  1-  The shoot i s approximately 1 cm long  276  277 F i g . 58 a - h.  C e l l u l a r d e t a i l o f the f o l i a g e leaves o f Hygrohypnum  alpestre. a,  Leaf apex.  b, e.  Marginal l e a f c e l l s . '  c, d. Median l e a f c e l l s , f - h. A l a r c e l l s . Scale: a - e;  I  f - h;  I I  '  100um  100um  I |  279  4-  F i g . 59.  Hygrohypnum alpestre  280  Exsiccati  Examined  Bauer, Musci europ. e x s . # 643. (BRNM, NY) #  1277.  (BRNM)  Husnot, Musci G a l l i a e # 750 as Hypnum alpinum. Rabenhorst, B r y o t h e c a europaea #913. (NY) S e l e c t e d Specimens  (S-PA)  Examined  Canada B r i t i s h Columbia Lake Lindeman; W i l l i a m s 27 May 1898 as Hypnum a l p e s t r e . (NY) Manitoba S e a l R i v e r , W. o f Great I s l a n d ; R i t c h i e 1946. (CANM) Ontario Thunder Bay D i s t r i c t , Oskondaga R i v e r NNE o f Shabaqua S t a t i o n , G o l d i e Township; Garton 10685 as Drepanocladus revolvens. (CANM) Thunder Bay; Macoun J u l y 1869 as Hypnum m o l l e . (MO, NY) C u r r e n t R i v e r ; Macoun 19 J u l y 1869. (MO) Labrador C h u r c h i l l F a l l s , B r i d g e Camp A r e a ; B r a s s a r d 5523. (NFLD) N o r t h West T e r r i t o r i e s Great Bear Lake, P o r t Radium, v i c i n i t y o f the E l d o r a d o Mine; S t e e r e 10246. (CANM) Campbell Lake, 68° 08' N, 133° 27' W; S c o t t e r 9742. (C) Hudson's Bay, B e l c h e r I s l a n d s , Tukarak I s . , near Long Lake; D o n t t #355. (MICH, NY) Queen E l i s a b e t h I s l a n d s B a f f i n I s l a n d , Head o f C l y d e I n l e t , F a l c o n R i v e r , Wynne-Edwards 9285. (CANM, NY) United  States Alaska Brooks Range, E n d i c o t t e Mountains, SW end o f Chandler Lake; S t e e r e 18252a. (NY) Brooks Range, n o r t h e r n s l o p e o f t h e DeLong Mountains, v i c i n i t y o f Noluck Lake; S t e e r e 63-380. (NY) Greenland Godhavn; Berggren 1870. (NY, S-PA) M a r t l e k ; Berggren 1870. (S-PA) Jacobshaven; Berggren 1870. (S-PA) Ritenbenk; Berggren 1870. (S-PA) Svartenhuk P e n i n s u l a , U m i a r f i k F j o r d , Amitsoq; Holmen 12002. (CANM, LE, NY) C h r i s t i a n h a l b ; Holmen 12066. (NY) Scoresby Sund, Sydlap; Holmen 18461. (NY) G e o g r a p h i c a l S o c i e t y I s l a n d s ; Holmen 18634. (NY) Renland, R y p e f j o r d ; Holmen 19166. (CANM, NY) Iceland E. I c e l . , E g i l s s t a o i r , E y v i n d a r a ; Johannsson 12 Aug. 1967. (FLAS)  281 Bear Island Mt. Misery; Berggren 1868. (S-PA) Spitzbergen Kobbe Bay; Berggren 1868. (MICH, S-PA) Sweden Halsingland, Abra, Hofsatter; Collinder 17 July 1878. (S-PA) Harjedalen, Storsjo, Lyungdalsforsen; F l o r i n 10 July 1931. (S-PA) Jamtland, Frostvikens; F l o r i n 13 Aug 1925. (S-PA) Nordbotten, Kiruna Katterjaure; Hulphers 14 July 1944. (S-PA) Asele Lappmark, Vilhelmina; Moller 20 July 1914. (S-PA) Lycksele Lappmark, Tama, N. Storsf j a l l e t ; Stenholm 16 July 1924. (S-PA) 1^ Pite Lappmark, Arjepluog; Moller 12 July 1918. (S-PA) Lule Lappmark, regio Sarjekensis; Jensen & A r n e l l 3 Aug. 1902. (S-PA) Torne Lappmark, Abisko; Hulphers 21 July 1941. (S-PA) Norway Kongswold; Kindberg July 1885. (S-PA) Lomseggere; Bryhn July 1879. (S-PA) Finmarken, mellan Nergaan ob f j e l l e t Aglappen; Maalseludalen; Holmgren 28 July 1868. (S-PA) Norlands, am Salten Bejeeren; Hagen 3 Aug 1894. (S-PA) Tromso, amt Nordreisen; A r n e l l 29 Aug 1891. (S-PA) Gausta; Jaederholm, July 1895. (S-PA) Finland N. E. Le Poroeno, Vallinkorva; L. & H. Roivainen 1 Aug 1961. (BP, UAC) Soviet Union Siberia, Yakutia, Lowlands of the Lena River, Buchta T i k s i ; Kildjushebski 12 Aug 1955. (LE) Kola Peninsula, Khibiny Mnt.; Shelyakov 1768a. (LE) T i l a j a k , Konjakovsk Serebryank Mountaints, Headwaters of the Iova River; Gorchakovski 9 June 1953. (LE) Middle Urals, Denezhkin Kamen; Teklemushev 10 July 1928. (LE) Novaya Zemlya, Guba Krestova; Brotherus 9-10 Aug 1901. (LE) Turukhansk District,, Jenesei I n l e t , Dudinka River, Matvjev 253. (LE) Siberia, Jenesei, Duninka; A r n e l l 7 Aug 1876. (C,NY) Lapponia ponojensis; Ponoj; M. Brenner 17 July 1863. (LE)  282 Hygrchypnum p o l a r e 198.  (Lindb.) Loeske, Verh. Bot, V e r . Brandenburg 46:  1905.  Lectotype:  Insulae Spetsbergensis,  Malmberg.  1861.  Wahlenbergs Bay.  Leg: A . J .  (S-PA)  Hypnum p o l a r e L i n d b . ,  Oefv. K. V e t . Ak. Foerh.  23:540.  Hypum p o l a r e v a r . pseudostramineum L i n d b . , Osfv. Foerh.  23:540.  1867.  Kobbe Bay.  Holotype:  1867  K. V e t . Ak.  H-SOL, S p i t z b e r g e n , ad  Leg. A . J . Malmberg, 1861.  Amblystegium p o l a r e  (Lindb.)  L i n d b . , Musci Scand. 33. 1879.  Hypnum p o l a r e v a r . l a e t e - v i r e n s E k s t r . , B o t . Not. 1880:198. 1880.  nom. nud.  Hypnum p a l u s t r e v a r . p o l a r e  (Lindb.) Husn., Muse. G a l l . 411  1894. Calliergon polare  (Lindb.) Kindb., E u r . N. Am. B r y i n .  1:82.  1897. Hypnum p o l a r e v a r . f a l c a t u m Bryhn, Nyt. N a t u r v i d . Hygrohypnum  polare  1902.  (Lindb.) B r o t h . , Nat. P f l . 1(3):1041. 190 .  Amblystegium e h l e i Am., Lectotype:  40:24.  Ark. Bot. 13 ( 2 ) : 74-75 f . 3 . 1913.  S i b e r i a , Lena, Kumashsur, 70° 30' N;  N i l s s o n - E h l e 1898.  (S-PA)  Hygrohypnum ehleL (Am.) B r o t h . , Trav. Mus. Bot. Ac. Sc. Petrograd.  16:42