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BEN # 196
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No. 196 June 23, 1998
aceska@victoria.tc.ca Victoria, B.C.
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Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2
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ANNUAL GENERAL MEETING - NPSBC NATIVE PLANT SOCIETY
OF BRITISH COLUMBIA - SATURDAY, MAY 23RD, 1998
From: NPSBC Native Plant Society of British Columbia
<npsbc@hotmail.com>
The NPSBC Native Plant Society of British Columbia held its
second Annual General Meeting on Saturday, May 23rd at the
Penticton Lakeside Resort and Conference Centre in Penticton,
BC. The meeting introduced two new projects of the Society: an
Atlas of British Columbia Flora that will involve members of the
Society and the public in surveying native plants and habitats
in regions throughout the province; and Ethical Use Guidelines
and Principles.
The Atlas project was introduced by the Biodiversity/Research
Committee. Later, Al Oliver of the Agriculture and Agri-Food
Canada Food Inspection Agency spoke on the issue of ethics using
the impact on native plant nurseries and habitats from gypsy
moth infestation as an example. There were also a number of
exhibits by businesses and organizations with a related interest
in native plants and habitats.
During the business portion of the Annual General Meeting, four
new directors were elected to the Board of fifteen members
including Brian Compton of Vancouver, Malcolm Martin of Vernon,
Brenda Ramsay of Terrace and Claudia Schaefer of Vancouver; four
other directors were re-elected to the Board and eight continued
for the second year of their two-year terms.
The current Executive will continue for another year, with
Douglas Justice, New Westminster as President.; Tom Wells, Delta
as Vice-President; Ross Waddell, Vancouver as Secretary; and
Sylvia Mosterman, Chilliwack as Treasurer. Other continuing
directors include Adolf Ceska, Victoria; Theresa Duynstee,
Delta; Verna Miller, Spences Bridge; John Olafson, Victoria;
Giles Stevenson, Duncan; Paulus Vrijmoed, Langley; and David
Williams, Kamloops.
Prior to the meeting, the Society held a popular workshop on
"The Identification of Grasses" in cooperation with the Ministry
of Forests in Penticton. The workshop was led by Board member,
Dr. Adolf Ceska of the BC Conservation Data Centre in Victoria,
assisted by Oluna Ceska. This two-day event was one of the most
requested in the Society's recent membership survey.
The attendees spent Saturday afternoon visiting the Ornamental
Gardens at the Pacific Research Centre of Agriculture and Agri-
Food Canada in Summerland. That evening, the Annual Dinner of
the Society featured Dr. Roy L. Taylor, Executive Director of
the Rancho Santa Ana Botanic Garden speaking on "California
Native Plant Initiatives - A Model for British Columbia". Dr.
Taylor, who is the former Director of the University of British
Columbia Botanical Garden, is an internationally recognized
expert on the flora of the Queen Charlotte Islands.
The weekend ended with two field trips in the South Okanagan
Region: Adolf and Oluna Ceska led a trip to the grassland bluffs
at Vaseux Lake; and Harold Baumbrough hosted members at the
Nature Trust in Naramata. The Society will now focus on the
implementation of the Atlas of British Columbia Flora Project
and solicits the interest of persons from throughout the
province who wish to become involved with this project. Dr. Bob
Maher and the Centre for Environment and Information Management
(CEIM) of the Royal Roads University in Victoria, expressed the
interest to co-ordinate this project.
Individual membership fees are $20 per year (applications should
be sent to NSPBC, 2012 William Street, Vancouver, B.C., V5L 2X6)
and include subscriptions to MENZIESIA, a newsletter published
3-4 times a year.
ARE HORNWORTS THE BASAL LINEAGE OF LAND PLANTS?
From: David J. Garbary, Department of Biology, St. Francis
Xavier University, Antigonish, Nova Scotia, Canada, B2G 2W5
<dgarbary@juliet.stfx.ca>
and
Karen S. Renzaglia, Department of Plant Biology, Southern
Illinois University, Carbondale, IL 62901-6509, U.S.A.
<renzaglia@plant.siu.edu>
[In the cover letter to this article, Dr. Garbary wrote: "I
tried to make this ms as simple as possible in terms of jargon
and taxonomic detail. The only assumption that I have made is
that readers have some familiarity with the language of
phylogenetic analysis." I found a nice glossary of terms used in
cladistics on the following web page:
http://www.ucmp.berkeley.edu/glossary/gloss1phylo.html
and a good introduction to cladistics is at
http://www.ucmp.berkeley.edu/clad/clad1.html
A. Ceska]
The identification of the primary lineages of land plants and
their phylogenetic relationships is a fundamental question in
plant evolutionary biology (Graham 1993, Kenrick and Crane
1997). A primary aspect of this problem involves the three
groups of bryophytes (i.e., hornworts, liverworts and mosses)
and their relationships to each other and to remaining lineages
of terrestrial plants. These bryophyte groups are central to the
problem because of conflicting interpretations regarding whether
or not liverworts or hornworts are the sister group to remaining
land plants, and whether or not mosses plus liverworts form a
monophyletic lineage. Recent phylogenetic interpretations in-
volving hornworts (Anthocerotopsida) have led to two widely
accepted points:
1. that they are part of a grade of organisms called bryophytes
which comprise one of the oldest lineages of land plants,
and
2. that there are many specializations of these organisms that
set them apart from other land plants and other bryophytes
(see Schofield 1985 for summary).
Here we review the current state of our knowledge on hornwort
phylogeny, and their relationships with other land plants.
Monophyly of land plants is an assumption of this essay.
The phylogenetic trees of Mishler & Churchill (1984, 1985)
provided a seminal contribution to interpretations of land plant
evolution. The central feature of their cladograms was the
paraphyletic nature of the bryophytes with three lineages occur-
ring in sequence (liverworts, hornworts and mosses) at the base
of the land plant clade (Figure 1). Two consequences of this
arrangement were that the liverworts were the sister group of
all other land plants and that mosses were the sister group to
vascular plants (and as was suggested by Kenrick & Crane 1991,
1997 - the protrachiophyte grade).
Although the Mishler & Churchill scheme has been widely
reproduced and accepted, several conflicting interpretations of
hornwort relationships have been proposed subsequent to their
groundbreaking study. Garbary et al. (1993) used male
gametogenesis and sperm architecture to base phylogenetic
analyses of land plants. The premise behind utilization of these
characters is that male reproductive development provides an
unparalleled suite of unequivocal characters that are truly
homologous across all land plants with flagellated sperm. This
study suggested that all bryophyte groups formed a monophyletic
assemblage that was sister to remaining vascular plants (Figure
2). The only counter intuitive result was the placement of
Selaginella basal on the bryophyte clade. Subsequent analyses of
Selaginella based on new data that emphasized development rather
than mature sperm morphology has placed Selaginella in a more
intuitively correct position within a lycophyte assemblage
(Maden et al. 1997, Renzaglia et al. 1998). The position of
hornworts external to a moss plus liverwort assemblage is a
primary feature of the phylogenetic hypothesis based on male
gametogenesis.
More recent approaches to this problem have included a number of
molecular sequence studies and a more comprehensive analysis of
morphological, developmental and ultrastructural features across
a wide range of bryophyte, lycophyte and pteridophyte as-
semblages (Hedderson et al. 1996, 1998; Malek et al. 1996;
Garbary & Renzaglia 1998). The pattern of relationships resolved
in these studies suggests that hornworts are the basal lineage
of land plants. In addition, these studies suggest that mosses
and liverworts form a monophyletic group that is the sister-
clade to vascular plants (Figure 3). The molecular studies use
the 18s gene for rRNA and the cox3 mRNA. Conflicting conclusions
are derived from sequences of the chloroplast gene for rbcL by
Lewis et al. (1997) where hornworts are resolved as the sister
group to vascular plants.
In summary, several studies are congruent in indicating the
basal position of hornworts in land plants, as well as the
monophyly of a moss plus liverwort clade. Although these conclu-
sions must be regarded as tentative pending more comprehensive
taxon sampling for both molecular and non-molecular characters,
the overall evidence seems to be away from the Mishler and
Churchill hypothesis, and to a more fundamental role for
hornworts in interpretations of land plant phylogeny. Clearly,
the hypothesis that hornworts are the most ancient extant
lineage of land plants remains a viable option that requires
further testing and exploration.
References:
Garbary D.J., K. Renzaglia, & J.G. Duckett. 1993. The phylogeny
of land plants: a cladistic analysis based on male
gametogenesis. Plant Systematics and Evolution 188: 237-269.
Garbary D.J. & K.S. Renzaglia. 1998. Bryophyte phylogeny and the
evolution of land plants: Evidence from development and
ultrastructure. In: Bryology for the Twenty-First Century.
Proceedings of the Centenary Symposium of the British
Bryological Society (Bates J.W., N.W. Ashton, & J.G. Duckett
- Editors). Maney and The British Bryological Society. Leeds
(in press).
Graham L.E. 1993. Origin of land plants. John Wiley: New York.
Hedderson T.A. & R.L. Chapman. 1998. The origins and diver-
sification of land plants; new evidence from molecular data.
In: Bryology for the Twenty-First Century. Proceedings of the
Centenary Symposium of the British Bryological Society (Bates
J.W., N.W. Ashton, & J.G. Duckett - Editors). Maney and The
British Bryological Society. Leeds (in press).
Hedderson T.A., R.L. Chapman, & W.L. Rootes. 1996. Phylogenetic
relationships of bryophytes inferred from nuclear encoded
rRNA gene sequences. Plant Systematics and Evolution 200:
213-224.
Kenrick P. & P.R. Crane. 1991. Water-conducting cells in early
fossil land plants: implications for the early evolution of
tracheophytes. Botanical Gazzette 152: 335-356.
Kenrick P. & P.R. Crane. 1997. The origin and early evolution of
land plants, a cladistic study. Smithsonian Institution
Press: Washington.
Maden A.R., D.D. Whittier, D.J. Garbary, & K.S. Renzaglia. 1997.
Ultrastructure of the spermatozoid of Lycopodiella lateralis
(Lycopodiaceae). Canadian Journal of Botany 75: 1728-1738.
Malek O., K. Luettig, R. Hiesel, A. Brennicke, & V. Knoop. 1996.
RNA editing in bryophytes and a molecular phylogeny of land
plants. EMBO Journal 15: 1403-1411.
Mishler B.D. & S.P. Churchill. 1984. A cladistic approach to the
phylogeny of "bryophytes." Brittonia 36: 406-424.
Mishler B.D. & S.P. Churchill. 1985. Transition to a land flora:
phylogenetic relationships of the green algae and bryophytes.
Cladistics 1: 305-328.
Renzaglia K.S., D.L. Bernhard, & D.J. Garbary. 1998. Development
of the male gamete of Selaginella australiensis. Interna-
tional Journal of Plant Sciences (in press).
Schofield W.B. 1985. Introduction to bryology. Macmillan New
York.
FIGURES - Three phylogenetic hypotheses for the relationships of
the three groups of bryophytes to each other and to vascular
plants. [In some e-mail systems, the Microsoft is trying to have
the last word, and the cladograms will come out slightly
misaligned. - AC]
Figure 1. Mishler and Churchill model with liverworts
as the sister group to all other land plants.
.....................................................
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....... L ......... H ......... M ........ V ........
....... I ......... O ......... O ........ A ........
....... V ......... R ......... S ........ S ........
....... E ......... N ......... S ........ C ........
....... R ......... W ......... E ........ U ........
....... W ......... O ......... S ........ L ........
....... ......... ......... ........ ........
....... \\ ........ \\ ........ \\ ..... //.........
......... \\ ........ \\ ........ \\ .. //...........
........... \\ ........ \\ ........ \\//.............
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Figure 2. Revised hypothesis based on male gametogenesis
with bryophytes as monophyletic assemblage.
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....... V ......... H ......... L ........ M ........
....... A ......... O ......... I ........ O ........
....... S ......... R ......... V ........ S ........
....... C ......... N ......... E ........ S ........
....... U ......... W ......... R ........ E ........
....... L ......... O ......... W ........ S ........
....... ......... ......... ........ ........
....... \\ ........ \\ ........ \\ ..... //.........
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Figure 3. New model with hornworts as sister group to all
other land plants and with mosses and liverworts as
monophyletic.
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....... H ......... V ......... M ........ L ........
....... O ......... A ......... O ........ I ........
....... R ......... S ......... S ........ V ........
....... N ......... C ......... S ........ E ........
....... W ......... U ......... E ........ R ........
....... O ......... L ......... S ........ W ........
....... ......... ......... ........ ........
....... \\ ........ \\ ........ \\ ..... //.........
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RED ALDER (ALNUS RUBRA BONG. - BETULACEAE) IN MONTANA
From: Toby Spribille <Spribille_Toby/r1_kootenai@fs.fed.us>
Over the past couple years there have been several reports of
red alder (Alnus rubra) in Montana, mostly originating from the
vicinity of Troy, in the extreme western portion of the state.
Several suspected sightings turned out to be "false alarms", but
a report last year from a Forest Service timber stand exam
inspector, Alan Lane, of a 22-inch dbh specimen found near the
Idaho border clearly ruled out our other common species, A.
sinuata ssp. crispa and A. incana ssp. tenuifolia. On 16 June I
visited one of the reported locations southwest of Troy with
Mike Arvidson, a Kootenai National Forest botanist based in
Troy, and we collected samples:
Montana, Lincoln Co., southwest of Troy and east of Idaho bor-
der, South Fork of Callahan Creek, frequent in creek bottom
with Thuja plicata, Acer glabrum, Asarum caudatum etc. 16 Jun
1998 Spribille & Arvidson (Fortine Herbarium, duplicates will
go to WTU, MONTU, COLO).
The material we found clearly belongs to Alnus rubra, adding a
new native tree species to the flora of Montana!
Alnus rubra is a coastal species familiar to botanists west of
the Coast Ranges and Cascades but relatively rare inland. The
Montana stations represent the easternmost limits of the
species' range. Red alder is occasional in northern Idaho, where
it is suspected of hybridizing with Alnus incana ssp. tenuifolia
(Lorain 1988), a phenomenon which may also be occurring in the
Montana populations.
Red alder joins a suite of other coastal vascular plants, mosses
and lichens which are disjunct in the "inland wet belt" stretch-
ing from west-central Idaho and western Montana north into
central B.C. (and it has been a very wet belt this year). These
species are particularly conspicuous in the valleys along the
Clark Fork and Kootenai Rivers and in the Cabinet and Purcell
Mountains of northwest Montana. If the Pacific Northwest is ever
blessed with a distribution atlas for our flora it would be
interesting to gauge the participation by percentage of various
phytogeographic elements -- e.g., coastal, interior northwest,
boreal and cordilleran -- in these local floras and compare them
with coastal areas. But that will have to await better budgets!
Reference:
Lorain, C.C. 1988. Floristic history and distribution of coastal
disjunct plants of the northern Rocky Mountains. M.Sc.
Thesis, Univ. Idaho, Moscow. xii + 212 p.
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