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BEN # 168

Adolf Ceska aceska at CUE.BC.CA
Sat Jul 5 03:10:22 EST 1997

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No. 168                              July 5, 1997

aceska at freenet.victoria.bc.ca        Victoria, B.C.
 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2

          This issue of BEN is dedicated to bryologist

                     DR. WILF B. SCHOFIELD,

Professor  Emeritus of the University of British Columbia on the
occasion of his 70th birthday. Through his work in the fields of
bryology and phytogeography, Dr. Wilf Schofield  has  made  sig-
nificant contributions to our botanical knowledge of the Pacific
Northwest, Canada, and other parts of our planet.

The  Journal  of the Hattori Botanical Laboratory is planning to
publish a special Festschrift issue dedicated to  Dr.  Schofield
with  over  thirty  papers. Although BEN cannot compete with the
Hattori Journal, I hope that even those BEN readers  not  really
interested  in  mosses will appreciate the special focus of this

Dr. Wilf Schofield will celebrate his 70th birthday (July 19) on
a field trip to Alaska. All the best success  on  this  and  all
other endeavours!

From:  Pekka  Pakarinen,  Department of Ecology and Systematics,
      University of Helsinki, Finland
      <pakarine at cc.helsinki.fi>

A number of bryological and vegetational studies in  the  boreal
and   north-temperate   regions  have  shown  a  generally  high
resemblance  in  the  bryophyte  flora  and  bryophyte   ecology
throughout  the  northern  wetlands  (e.g.  in  Japan Suzuki and
Fujiwara acc. to Tuxen et al. 1972, in Quebec Gauthier 1980, and
in Minnesota Janssens 1992).

Sjors (1963) compared boreal wetlands in Ontario and  Fennoscan-
dia, and was able to identify corresponding ecological gradients
in  mire (peatland) vegetation of these two regions primarily on
the basis of the  bryophyte  composition  and  water  chemistry,
while the similarity in vascular strata was lower.

During  the  past  10-20 years, great advances have been made in
vegetational research of wetlands both in North America  and  in
Europe.  Wetland  classifications  of extensive regions, such as
Canada (National Wetlands Working Group 1988) or Alaska (Viereck
et al. 1992) tend to emphasize large, mappable  landscape  units
and  contain  vegetational classes at lower levels utilizing one
or two dominant species of each layer.

The question is then: if bryophytes are a significant  component
of  these  systems,  particularly  in peatlands, and if they are
widely distributed, common indicators of wetland ecology  across
the  circumarctic  and  circumboreal  bioclimatic zones, what is
their role in vegetational classifications?

The traditional Braun-Blanquet system  (BB)  basically  utilizes
the  information  of  all  plant  species  of  community. Within
Europe, a number of  national  vegetation  classifications  have
been  published  recently,  based on the European phytosociology
(Braun-Blanquet  school),  e.g.  those  of  Rodwell  (1991)  for
Britain, and Schaminee et al. (1995) for the Netherlands.

There  have  been attempts to expand the BB system also to other
regions than continental Europe. Tuxen  et  al  (1972)  compared
Sphagnum  bogs  of  Europe  with  selected  North  American  and
Japanese   materials   and   developed   a   tentative    global
phytosociological  classification  of  bog  hummock communities.
Recently,  Wells  (1996)  summarized  his  detailed   vegetation
studies  in  Newfoundland  developing a phytosociological class-
ification for peatlands of Atlantic Canada.  Wells  (1996)  dis-
cusses  the relationships with the European vegetation types and
develops in most cases a new nomenclature for his study region.

In his  new  book  (Vegetation  of  North  Europe,  in  German),
Dierssen  (1996)  who  has  earlier  studied  especially  the NW
European mires, expands the BB system to the north. The  problem
is   that  the  traditional  nomenclature  of  vegetation  units
developed originally for central Europe, is retained,  and  this
nomenclature  clearly  runs into complications in N Europe where
boreal and  subarctic  vegetation  prevails.  Bryophytes  appear
mostly  at  lower  levels as community types named by one or two
species, and for example the large class of rich fens is  called
Caricion  davallianae - the name refers to a sedge, Carex daval-
liana which is practically absent from the boreal and  subarctic
regions of Europe.

Instead of linking rich fen vegetation  to higher plants of gen-
erally  limited  distribution,  it would be preferable to define
circumboreal  rich fens  (Braunmoore, see Pakarinen 1995) by one
or  several bryophyte  species  characteristic  of these wetland 
habitats:  Campylium stellatum,  Cinclidium stygium, Tomentohyp-
num nitens.

Literature cited

Dierssen, K. 1996. Vegetation Nordeuropas. Eugen  Ulmer  Verlag,
      Stuttgart. 838 p.
Gauthier, R. 1980. La vegetation des tourbieres et les sphaignes
      du  parc  des  Laurentides,  Quebec. Etudes ecologiques 3.
      Laboratoire  d'ecologie  forestiere,   Universite   Laval,
Janssens,  J.A.  1992. Bryophytes. In: Wright, H.E. Jr., Coffin,
      B.A. & Aaseng,  N.E.  (ed.)  The  patterned  peatlands  of
      Minnesota,  p.  43-57. University of Minnesota Press, Min-
National Wetlands Working Group. 1988. Wetlands of Canada.  Eco-
     logical Land Classification Series 24.  Environment Canada,
     Ottawa. 452 p.
Pakarinen, P.  1995.  Classification  of boreal mires in Finland
      and Scandinavia: A review. Vegetatio 118: 29-38.
Rodwell,  J.S.  (ed.)  1991.  British plant communities. Vol. 2.
      Mires and heaths. Cambridge University  Press,  Cambridge.
      628 p.
Schaminee, J.H.J., Weeda, E.J. & Westhoff, V. 1995. De vegetatie
      van  Nederland. Deel 2. Opulus Press, Uppsala, Leiden. 357
Sjors, H. 1963. Bogs and fens on  Attawapiskat  River,  northern
      Ontario. National Museum of Canada Bulletin 186: 45-133.
Tuxen,  R.,  Miyawaki,  A.  & Fujiwara, K. 1972. Eine erweiterte
      Gliederung der Oxycocco-Sphagnetea. In: van der Maarel, E.
      & Tuxen, R. (ed.) Grundfragen und Methoden in  der  Pflan-
      zensoziologie, p. 500-520. Junk, The Hague.
Viereck,  L.A.,  Dyrness,  C.T.,  Batten,  A.R. & Wenzlick, K.J.
      1992: The Alaska  vegetation  classification.  Gen.  Tech.
      Rep.   PNW-GTR-286.   Portland,  OR:  U.S.  Department  of
      Agriculture, Forest Service,  Pacific  Northwest  Research
      Station, Portland. 278 p.
Wells,  E.D.  1996.  Classification  of  peatland  vegetation in
      Atlantic Canada. Journal of Vegetation Science 7: 847-878.

From: Dr. Jan-Peter Frahm [abbreviated from Frahm,  J.-P.  1994.
      Moose  -  lebende  Fossilien. Biologie in unseren Zeit 24:
      <frahm at uni-bonn.de>

We have been surprised to find that that mosses are one  of  the
most  conservative  plant  groups.  The  oldest  known bryophyte
fossils from the Palaeozoic era compare  well  with  the  recent
plants and can be assigned to living families or genera.

Mosses  have  changed very little in the last 300 million years.
Their origin goes even further back than this.  Initial  studies
of  mosses  fossilized in Dominican amber showed that even 25 to
40 million years ago the same species that  we  know  today  oc-
curred  on the Island of Hispaniola. Most of the fossil finds of
mosses in the last century were from the Tertiary and Quaternary
period. People believed that mosses did  not  preserve  well  as
fossils  because they have no woody parts. Only at the beginning
of this century the first moss from the Carboniferous period was
discovered in England. But by the  year  1957  33  species  were
known from the Palaeozoic and Mesozoic era (Savicz-Lyubitskaya &
Abramov 1959), and by 1967 the number grew to 68 species (Jovet-
Ast  1967,  Oostendorp 1987). Since then more fossil mosses have
been found in the Upper Permian rocks in Russia  (Ignatov  1990)
and  in  the  Permian  rocks  in  the Antarctica (Smoot & Taylor

Originally, fossil bryophytes were placed in several  artificial
"form" genera (e.g., Muscites, Hepaticites, Thallites etc.). New
studies  show that even the oldest known fossil mosses are often
so similar to the living mosses that they can be placed into the
recent families or genera. Because of this, we cannot  make  any
conclusions  about  the  evolution  of  mosses  based  on fossil

The origin of mosses goes further back than can be documented by
known fossils and we have to guess from what plant groups mosses
evolved. The oldest known bryophyte is Pallavicinites  devonicus
from  the  U.S.A., about 350 million years old. It can be placed
in the recent liverwort  order  Metzgeriales.  In  the  Devonian
period  we  can  find  thallose  liverworts  that are similar to
Anthoceros  or  Riccia.  The  oldest  "true"  moss  is  Muscites
plumatus from Carboniferous period, found in England that can be
placed  into  recent order Bryales. In the Permian period we can
find the first fossils related to peat moss (cells are differen-
tiated  into  green   cholorocysts   and   empty   water-storing
hyalocysts),  but  having leaves with midrib. These fossils were
placed into a separate fossil order Protosphagnales, but  it  is
questionable if they are indeed the ancestors of our recent peat
moss  (Sphagnum).  The first pleurocarpous mosses are also known
from the Permian period.

Mesozoic strata are relatively  poor  on  moss  fossils  due  to
larger  extent  of  dry  regions  that were not so favourable to
fossilization as the Palaeozoic or the Tertiary  swampy  forests
were.  From  the  recent  areas of distributions we can conclude
that in this time happened the  differentiation  of  the  mosses
adapted  to  dry  environments.  Fossils from this time resemble
Anthoceratales,  Marchantiales,  Metzgeriales,   Jungermaniales,
Sphagnales and Bryales.

In  Europe,  forty  species of mosses are known from Miocene and
seventy-nine from Pliocene (Dickson 1973). All except  four  are
mosses  that  still  exist  today.  The  species  composition of
Miocene and Pliocene mosses of Europe is similar to  the  recent
bryoflora  of  southern parts of North America or of Macronesian
Islands (Canary Islands, Madeira, Azores). Genera such  as  Gol-
lania  or  Echinodium spread to these volcanic islands from con-
tinents, but have disappeared from Europe during the glaciation.
The bryoflora of some parts of Macronesian Islands gives a  good
impression of the late tertiary bryoflora of Europe.

Amber  is  an  important  source  of fossils, including those of
mosses.  Mosses  are  known  from  Baltic,  Saxon,  Mexican  and
Dominican  amber,  all from the Tertiary. While the Baltic amber
originated from the pitch of conifers, Dominican amber came from
leguminous tree (Hymenaea spp. - Caesalpinaceae [recent trees of
this genus are source of copal - AC]) and is extremely clear and

There are eighteen species of liverworts known from Baltic amber
and twelve from the Dominican amber (Grolle 1988).  If they  are
identifiable,  these   are  all  extinct  species  and they were
described and classified as  artificial  form-genera.  A  recent
review of these samples resulted only in one new identification,
i.e.,  the type of  Muscites hauchecornei has been identified as
the recent Trachycystis flagellaris that now occurs from eastern
Asia  (China,  Korea,  Siberia) to Alaska.  In  the Miocene this 
species occurred in Poland.

Two  other  now  extinct  species  of  Trachycystis  have   been
described  from  Miocene in Silesia and from the Pliocene of the
border area between Germany and  Holland  (Miller  1984).  These
species  represent  examples of Asiatic elements in the Tertiary
flora of Europe  that  were  (as  many  flowering  plants,  e.g.
Ginkgo,  Sequoia,  Metasequoia)  wiped  out in Europe during the

In summary, bryophytes are very old in  an  evolutionary  sense.
Liverworts  existed  in  the  Devonian era 350 million yers ago.
Today's bryophyte orders  were  already  differentiated  in  the
Paleozoic  period.  Today's  genera are known from Tertiary fos-
sils. More than 90 % of the species of  mosses  present  towards
the  end  of  the  Tertiary  are  still extant today. Apart from
fluctuations in abundance  of  certain  species,  the  bryophyte
flora  of  the  Quaternary was already identical with the recent

Literature cited

Dickson, J.H. 1973. Bryophytes of the Pleistocene. Cambridge.
Grolle, R. 1988. Bryophyte fossils in Amber. Bryol. Times 47: 4-
Ignotov, M.S. 1990. Upper Permian mosses from the Russian  plat-
      form. Palaeontographica Abt. B. Bd. 217: 147-189.
Jovet-Ast,  S. 1967. Bryophyta. Pp. 17-186 in: Boureau, E. [ed.]
      Traite de Paleobotanique 2.
Miller, N.D. 1984. Tertiary and Quaternary  fossils.  Pp.  1194-
      1232 in: Schuster, R.M. [ed.] New manual of bryology, Vol.
Oostendorp,  C.  1987.  The  bryophytes  of  the  Palaeozoic and
      Mesozoic. Bryophytorum Bibl. 34.
Savicz-Lyubitskaya, L.I. & I.I. Abramov.  1959.  The  geological
      annals of Bryophyta. Revue Bryol. Lichenol. 28: 330-342.
Smoot,  E.L.  & T.N. Taylor. 1986. Structurally preserved plants
      from Antarctica: II. A  Permian  moss  from  the  Transan-
      tarctic Mountains. Amer. J. Bot. 73: 1683-1691.

From: Dr Weber <weberw at spot.Colorado.edu>

A  few  weeks  ago  Herbarium COLO received a gift from the Her-
barium at Gothenburg, Sweden. In moving into a new building  for
the  herbarium  there  the  remains  were  discovered of a long-
standing bryophyte  exchange  club  that  operated  out  of  the
botanical  garden.  The  club  dissolved  many years ago, and my
friend Uno Eliasson felt that since I know the Swedish  language
well  enough  to  decipher  thousands  of hand-written labels, I
might be the best place to send them  inasmuch  as  no  American
institutions  received  material  from  the club. Specimen dates
range from 1824, then mostly between 1860 and 1900,  and  a  few
from  the  1930s.  The  collection came in ten big cartons, each
containing at least 2,000 specimens. I have made a list of  over
250 contributors, each with a distinctive handwriting, All but a
few  of  the labels are hand-written on the packets. The collec-
tion is of very fine quality, and many of the well-known profes-
sionals, such as Zetterstedt, Du Rietz, Bryhn, Hagen,  Blomberg,
Kindberg, S. O. Lindberg, Wilhelm and Sigfrid Arnell, Schiffner,
Schimper  and  Warnstorf  are  represented,  along  with laymen,
priests, teachers, and landed gentry. This collection represents
a great addition to our Scandinavian collections.

Botanical exchange clubs were very important in  the  nineteenth
century;  there  were  several  of them on the continent, and in
America cryptogamic exchange clubs are still operating  although
at  a  declining  rate.  They brought together all levels of ac-
tivity and knowledge among devotees  of  mosses,  hepatics,  and
lichens.  A  very  vigorous exchange club in Lund dealt with all
groups of plants, especially vasculars,  and  died  only  a  few
decades  ago. The specimens fill the herbaria of Europe, and the
geographical coverage is extraordinary, despite the difficulties
of travel at the time. One  large  segment  of  this  collection
consists  of brophytes collected in Spitzbergen and North Green-
land from 1861 to 1870 by  Sven  Berggren  and  A.  L.  Malmgren
labelled  "Plantae  in  itineribus Succorum polaribus collectae"
(plants collected in the course of  polar  rescue  expeditions)!
The  data  are sparse, merely the country, province, parish, and
village; rarely is there any mention of the habitat.

When an interesting plant was found, it appears that many people
visited the spot over a period of years  and  collected  at  the
same  site. An important offshoot of this extraordinary collect-
ing activity was not only the creation of museum collections and
the instruction of hundreds of  serious  amateurs  in  the  sub-
science  of  bryology (certainly a lot more than we have today).
Some years ago I encountered scientific papers in which  it  was
shown  that  the  accumulation of heavy metals, especially lead,
"frozen" and dated in moss specimens  in  the  herbaria  enabled
scholars  to  trace  the origin and spread of pollution over the
course of the Industrial Revolution in Scandinavia!

The curator at Gothenburg should be congratulated on not letting
this collection go into the dumpster, as it well might have, and
certainly would have in many an American institution where space
is a problem, and where "classical"  taxonomy  is  considered  a
waste  of  good  academic  space.  I  receive many letters in my
present "exile" from the COLO  herbarium  that  tell  me  horror
stories  of herbaria everywhere being sent down the drain, their
value and importance being denied in favor of  the  more  glitzy
new  aspects of biology. Every innovational aspect of biology is
surprisingly important,  but  the  museum  collections  must  be
preserved  at  all costs, because they are our ultimate basis of
understanding biodiversity. We are also losing history. I  shud-
der at the loss of the life works of professors, whose notes and
unpublished  work  is  tossed  along with their human usefulness
upon retirement. Our library is now discarding journals  because
they  take  up  too  much  space,  and  will  replace  them with
microfilm unless they have colored plates and maps.  Is  western
civilization about to lose its Alexandrine library? Who is going
to rally the troops?

From: Nils Cronberg <Nils.Cronberg at sysbot.lu.se>

Cronberg,  N.,  Molau, U. & Sonesson, M. 1997. Genetic variation
      in the clonal bryophyte Hylocomium splendens at hierarchi-
      cal geographic scales in Scandinavia.  Heredity  78:  293-
      301. [Abstract]

Molecular  techniques can contribute to the solving of taxonomic
problems e.g., involving polyploidisation, the  distinctness  of
closely  related  taxa  and phylogenetic relationships. The dis-
tribution of molecular markers can also  give  new  insights  to
biogeographic  problems.  Thirdly,  they can be used to identify
genotypes which is useful in the  study  of  clonality  and  the
balance  between genotypic and phenotypic control of morphologi-
cal traits.

The clonal  bryophyte  Hylocomium  splendens  is  widespread  in
arctic  and  boreal  regions  of the Northern Hemisphere. Arctic
populations,  morphologically  differentiated  from  the  boreal
populations,  are  sometimes  referred  to as H. alaskanum. Most
bryologist, however, prefer to use the name  H.  splendens  also
for  this  material since the variation between the two extremes
is continuous.

In a first study H. splendens s. l. was sampled in a  hierarchi-
cal  fashion  from  populations  representing  four Scandinavian
vegetation zones (S  boreal,  N  boreal,  subalpine,  subarctic-
alpine).  Allozyme  electrophoresis revealed variation at 11 out
of  13  screened  loci,  allowing  accurate  identification   of
genotypes. Clonal diversity was high; from a total sample of 298
shoots  79  genotypes  could  be  detected. Surprisingly, clonal
diversity was highest at the subarctic-alpine site, even at  the
level of 10x10 cm patches. The mean number of genotypes detected
from  samples of 5 shoots taken from each of 15 such patches was
2.9 at the subarctic-alpine site.

The total allelic diversity (HT) based on polymorphic  loci  was
0.274, which is comparable to the more variable vascular plants.
The relative differentiation among populations was low, only 7%,
indicating  a  high  level  of gene flow. As matter of fact, the
genetic structure of the Hylocomium populations  appears  to  be
similar  to  the genetic structure of the conifer species, under
which canopy it is often growing.

The 7% variation distributed among  populations  indicated  that
the  subarctic-alpine  population  (belonging to "H. alaskanum")
was slightly differentiated versus the three lowland population.
Preliminary results from  ongoing  studies  (including  material
also  from  Siberia) confirm this observation. Most probably, H.
splendens s. l. contains several geographic races with different
postglacial history, some of which  share  morphological  traits
attributed  to  H.  alaskanum.  These races are only weakly dif-
ferentiated genetically and therefore expected to  have  evolved

From: Adolf Ceska <aceska at freenet.victoria.bc.ca>

Dr.  Wilf  Schofield loves poetry and classical music. As a part
of this Festschrift I wanted to include an essay on the  use  of
bryophytes  in  these  two fields. The task proved too difficult
considering a short time available and the amount of poetry  one
would  have  to  read  or  the amount of music one would have to
listen to. I can only present few suggestions that the  Internet
gurus brought to my attention.

In  the anthology "A Book of Luminous Things" (edited by Czeslaw
Milozs) a poem "Moss-Gathering" by  Theodore  Roethke  describes
techniques  of  moss  collecting.  This technique is exactly the
same as that applied by Dr. Wilf Schofield:

   To loosen with all ten fingers held wide and limber
   And lift up a patch, dark-green, the kind for lining
                                cemetery baskets,
   Thick and cushiony, like an old-fashioned doormat,
   The crumbling small hollow sticks on the underside mixed
                                with roots,
   And wintergreen berries and leaves still stuck to the top,-
   That was moss-gathering.

[Roethke feels remorse for gathering so much moss:]

   And afterwards I always felt mean, jogging back over the
                                logging road,
   As if I had broken the natural order of things
                                in that swampland;
   Disturbed some rhythm, old and of vast importance.
   By pulling off flesh from the living planet;
   As if I had committed, against the whole scheme of life,
                                a desecration.

Dr. Schofield does not feel a similar guilt, on the contrary, he
can be proud of his collecting record. As of June 15,  1997,  he
has  collected  107,990  specimens (not counting duplicates) and
made them immortal in the herbarium collections.

In classical  music,  mosses  are  only  seldom  mentioned.

In Richard  Wagner's Parsifal,  Kundry describes to Parsifal how
his mother took care of him as a child:

   Gebettet sanft auf weichen Moosen
   den hold geschlafert sie mit Kosen.
   Gently bedded in soft moss
   she lulls carressingly her darling boy.

Czech composer Bedrich Smetana was a follower of Wagner  and  as
such had a similar moss theme. In his opera "Tajemstvi [Secret]"
the  smuglers  walk  through  a forest and sing (quite loudly) a
nice chorus:

   Quiet, quiet, [step] on moss ...

If you are really interested in moss and classical music, I  was
told,  you  should  try  the music of American composer Lawrence
Moss (1927 - ), although most of it sounds rather  academic.  My
anonymous  advisor  said that Flight for brass quintet is pretty
good, as is Incidental Music for two pianos & percussion.

I have not found any  references  to  liverworts,  hornworts  or
Takakiopsida in poetry or classical music.

Submissions, subscriptions, etc.:  aceska at freenet.victoria.bc.ca
BEN is archived on gopher freenet.victoria.bc.ca. The URL is:

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