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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
BRYOPHYTES AND CLASSIFICATION SYSTEMS OF NORTHERN WETLANDS
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-
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.
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.
MOSSES - LIVING FOSSILS
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
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.
A WINDFALL OF BRYOPHYTES
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?
MOLECULAR TECHNIQUES IN THE STUDY OF HYLOCOMIUM SPLENDENS
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-
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-
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
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
MOSSES IN POETRY AND CLASSICAL MUSIC
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
Thick and cushiony, like an old-fashioned doormat,
The crumbling small hollow sticks on the underside mixed
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
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,
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
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: