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BEN # 226
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BBBBB EEEEE NN N N BOTANICAL
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No. 226 July 1, 1999
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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BEN # 226 and BEN # 277 are dedicated to
DR. THOMAS CHRISTOPHER (CHRIS) BRAYSHAW,
Curator Emeritus of the Royal British Columbia Museum,
Victoria, British Columbia, Canada,
on the occasion of his 80th birthday, July 2, 1999.
BOTANICAL HIGHLIGHTS OF A MARCH 1982 TRIP TO SOUTHERN CHILE
WITH CHRIS BRAYSHAW AND DEKALB RUSSELL
From: Rudolf Schmid <schmid@socrates.berkeley.edu>
From 5-22 March 1982 Chris Brayshaw, Dekalb Russell, an apple
grower and fervent naturalist from West Virginia, and I
botanized in southern Chile, concentrating on Myrtaceae, con-
ifers, and monogeneric families (the #-marked below) or
monotypic, often endemic genera (the *-marked below). Some
sought-after plants such as Gomortega keule* (Gomortegaceae#) we
saw only in the splendid botanical garden at Valdivia. We camped
several times but otherwise spent nights in low-cost shelters
(refugios) and especially inexpensive hotels and motels.
We escaped Santiago the capitol as quickly as possible, heading
south on the Pan-American highway to Valdivia 835 km south.
Other than remnant espinal (Acacia caven woodland) there is
little to see along the highway until around Collipulli, some
575 km south of Santiago because the roadside vegetation has
been badly mangled.
The Lake Region south of Valdivia, especially Lago Llanquihue,
is very pretty, but the vegetation has been greatly disturbed by
farming, particularly dairying. However, the flanks of Volcan
Osorno (2661 m) are nicely intact. Plants of interest here, many
especially from an alien northern-hemisphere perspective, in-
cluded: Coriaria ruscifolia (Coriariaceae#), Embothrium coc-
cineum (Proteaceae), Empetrum rubrum (Empetraceae), Eucryphia
cordifolia (Eucryphiaceae#), Euphrasia trifida (Scrophularia-
ceae), Gevuina avellana* (Proteaceae), Griselinia ruscifolia
(Cornaceae), Misodendrum linearifolium (Misodendraceae#)
parasitic on Nothofagus dombeyi (Fagaceae), Ugni molinae (Myrta-
ceae), the cushion Nassauvia revoluta (Compositae), Gunnera
chilensis and G. magellanica (Gunneraceae#), Laurelia sp. (Moni-
miaceae), Pernettya myrtilloides (Ericaceae), Quinchamalium
ericoides (Santalaceae), Chloraea piquichen (Orchidaceae),
Cortaderia sp. (Gramineae), and Philesia magellanica (Philesia-
ceae).
I found the allerce forest of Fitzroya cupressoides* (Cupressa-
ceae) east of Lenca, about 35 km southeast of Pto. Montt, the
most impressive part of the trip. We got permission from the
manager of a lumber mill to go onto their private logging road
(I forget why there was no logging activity, even though it was
the middle of the week). We were offered a jeep but took our
cheap rental van, a Furgon Daihatsu, 6 km up the road until it
pooped out, camped nearby, and the next morning walked up the
steep road to its end at about 800 m el. The flora here was very
rich and spectacular, particularly the Fitzroya trees, both the
ones still standing and the ones slaughtered. One presumes the
trees on the steep slopes of the ravines still survive. At this
site there were many pteridophytes (especially Asplenium and
Blechnum spp.), many Myrtaceae such as Luma apiculata, Myr-
ceugenia spp., Myrteola nummularia, which is a delightfully
delicate prostrate shrub, Tepualia stipularis,* which is the
only capsular myrt (subfamily Leptospermoideae) native to the
New World, as well as other interesting taxa such as Saxegothaea
conspicua* (Podocarpaceae), Aextoxicon punctatum* (Aextoxica-
ceae#), Asteranthera ovata* (Gesneriaceae), Azara serrata (Fla-
courtiaceae), Campsidium valdivianum* (Bignoniaceae),
Crinodendron hookerianum (Elaeocarpaceae), Desfontainia spinosa
(Desfontainiaceae#), Griselinia sp. (Cornaceae), Hydrangea
serratifolia (Hydrangeaceae), Mitraria coccinea* (Gesneriaceae),
Tristerix (now Macrosolen) tetrandrus (Loranthaceae, parasitic
on the Campsidium), and Chusquea (Gramineae). One could easily
spend several days in a botanically rich place such as this.
Returning to Santiago and southeast of Collipulli we toured the
easily accessible and very spectacular Araucaria araucana forest
in Parque National Conguillio on the east flank of Volcan Llaima
(3060 m). During out visit, we did not see a single person in
the entire park. Here also was Austrocedrus chilensis*
(Cupressaceae) and Misodendrum linearifolium (Misodendraceae#)
parasitic on Nothofagus antarctica (Fagaceae).
The trip ended with a jaunt to the Farellones, a ski resort area
in the Andes just northeast of Santiago. This was also the
highest point (3000+ m) of the trip. The lower elevations are
covered with mattoral, which is especially impressive with its
numerous taxa of columnar cacti, such as Trichocereus sp.
parasitized by Tristerix aphyllus (Loranthaceae), and many
individuals of Kageneckia angustifolia and K. oblonga (Rosa-
ceae). At our highest elevation around 3000 m prostrate and
cushion plants of Calycera sp. (Calyceraceae) and especially the
umbels Azorella spp. and Laretia acaulis* were striking. Also
here were Ephedra chilensis (Ephedraceae#) and Oxalis geminata
(Oxalidaceae).
It was also fascinating to see taxa familiar to us from cultiva-
tion but growing wild in Chile, for instance: Calceolaria spp.
(Scrophulariaceae), Drimys winteri var. chilensis (Winteraceae),
Fuchsia magellanica (Onagraceae), Alstroemeria spp.
(Alstroemeriaceae), Lapageria rosea* (Philesiaceae), and so many
others. In contrast, we were less enthralled to see familiar
Californian taxa such as Eschscholzia californica (Papaveraceae)
and especially plantations of Pinus radiata (Pinaceae). The
latter have contributed so much to the impoverishment of the
native vegetation, such as at the classic collecting site at
Corral, down the river from Valdivia.
We never made it to the Juan Fernandez Islands or to Concepcion
area. Otherwise these would have figured in the above narrative.
To Chris, the best of birthday wishes for your 80th (on 2 July)
from me and all of your plant friends mentioned above.
SNOWPATCH BUTTERCUPS OF THE WESTERN MOUNTAINS OF NORTH AMERICA
From: Fulton Fisher <ffisher@sfu.ca>
Near treeline in the mountains of western North America,
wherever snowpatches persist well into the summer, one or more
of a closely inter-related group of species of Ranunculus is
likely to be found (Fisher et al. 1972, Rowley et al. 1975). All
seem to require an assured supply of meltwater from adjacent
snowpatches. Most occur mainly in full sunlight although one,
the well known mountain buttercup R. eschscholtzii, usually
barely emerges from partial shade. Although both natural and
artificial hybrids connecting the species are always sterile,
they do show a closely shared common ancestry. Their chromosome
numbers form a polyploid series from 2n=16 to 96, the high
number sets probably having evolved step by step by combinations
of lower number sets. The actual phylogeny should today be
easily determined by DNA analysis.
The best known and most widely ranging of the snowpatch butter-
cups is R. eschscholtzii Schlecht. This hexaploid (2n=48) was
first found near its northern limit on Unalaska Is. in the
Aleutians near sealevel but at the southernmost end of its range
in N.Mexico it reaches nearly 15000 ft, almost always in moist
shady places on the edge of the treeline or not far from it.
Measurements have shown that this species requires less light
than the others but with running water alongside, it may venture
well into the open.
Although Ranunculus eschscholtzii itself is often observed at
altitudes as low as 4000 ft in southern British Columbia (BC),
probably only one of its six other relatives, the circumpolar
R. nivalis L.(2n=96), has been found regularly in BC and only in
the far north. The other five diploid and tetraploid species
have almost mutually exclusive geographic ranges in the western
mountains of the US even though geographically they all overlap
R. eschscholtzii; only R. suksdorfii and R. eximius in the south
east corner of BC, appear to cross the Canadian border.
There are two western diploids, R. oxynotus Gray of the Sierra
Nevada, and R. suksdorfii Gray. This last is well known in the
Olympics and Cascades and as far south as the Siskiyou Mts of
northern California. But this species also crosses into the
Bitterroot Mts of Idaho (and has been reported even in the
Flathead and Waterton Lakes region of Canada). Its range forms a
big loop back southwards and again westwards into the drier
mountains of Idaho and eastern Oregon, and reaches even eastern
Nevada. This is a giant horseshoe with differences between its
ends.
The eastern populations of R. suksdorfii have formed at least
two new geographic races distinguishable as subspecies from the
original Cascades version first collected from Mt Hood by
Suksdorf. One of these subspecies was originally given full
species status as the glaucous R. trisectus Eastwood from
Oregon, but genetic studies suggest it to be much more closely
related to typical R. suksdorfii further west and it has been
accordingly designated as R. suksdorfii subsp. trisectus. A
second outlying race of R. suksdorfii occurs in the high moun-
tains of eastern Nevada near the famous Bristlecone Pine popula-
tion on Mt Wheeler. This has been named R. suksdorfii subsp.
caespitosus to reflect its compact bushy form and to distinguish
it from the very similar R. adoneus (see below). The populations
of R. suksdorfii in Idaho and Montana north of the famous Lewis
and Clarke Trail along the Clearwater River probably warrants
distinction as yet another subspecific race with its regularly
taller habit and less sharply pointed leaf indentations.
The leaf subdivisions of R. suksdorfii subsp. caespitosus are
very narrow, almost tubular, so that this glaucous diploid has
often been misidentified as R. adoneus Gray which is the mag-
nificent large flowered Rocky Mountain snow-buttercup (sometimes
called "The Colorado Buttercup") also possessing very narrow
leaf segments. Ranunculus adoneus is found only in the southern
Rockies of Wyoming and Colorado and the higher parts of Utah and
may even reach northern New Mexico. However R. adoneus is a
tetraploid and hence totally separate from its look-alike, the
diploid R. suksdorfii subsp. caespitosus from further west.
To almost complete the story there is another tetraploid in the
Rockies, R. eximius Greene, which extends from beyond the north-
ern range limit of R. adoneus at Togwotee Pass, from the Grand
Tetons, through Yellowstone Park and Montana to the Canadian
border (and possibly beyond like R. suksdorfii) and reaches
eastwards to the Bighorn Mts of Wyoming. But R. eximius is quite
unlike R. adoneus, its fellow tetraploid further south; for its
leaves are elliptical and barely divided at all with three to
five deep parallel lobes, often giving the appearance of little
raised hands. Finally there is a third diploid, R. macauleyi
Gray, south of R. adoneus in southern Colorado and New Mexico;
this species has long elliptical leaves even narrower than those
of R. eximius to the north. However the teeth across the leaf
tips of R. macauleyi are very shallow, in some cases almost
nonexistent.
In most places these diploids and tetraploids occur at higher
altitudes than does the hexaploid R. eschscholtzii so they
seldom meet that wide-ranging species because they occur at
different levels on the same mountains. However, good evidence
has been found to explain anomalous floral changes in
R. eschscholtzii (Fisher et al. 1979), whereby apetalous
variants have sometimes locally replaced the normally petalled
form (apetaly is a regular mutation in R. eschscholtzii but
usually at a low frequency). Favorably wet conditions sometimes
occur that allow a few colonies of R. eschscholtzii to tolerate
even full sunlight beside the normal habitat of the much more
numerous diploids (e.g., R. suksdorfii on the Olympics and on Mt
Adams) or tetraploids (e.g., R. adoneus at Teton Pass). These
sometimes form sterile hybrids with R. eschscholtzii. It has
been shown that the large alpine bee-flies (=hover-flies or
Syrphidae) seem unable to distinguish one large shining butter-
cup from another, so that over the years their promiscuity has
caused the selective replacement of the normally attractive
petalled flowers in R. eschscholtzii by apetalous forms. Normal
and viable seed set thus still occurs, even though the usual
long range bee-flies avoid the tiny flowers, because short-range
pollinators including ants successfully climb up and down ad-
jacent stems. Futile sterile cross pollination of
R. eschscholtzii to the diploids or tetraploids is thus avoided
even though the keen-eyed Syrphids are busy only a few yards
away.
Key to the leaves of Snow Buttercup species and subspecies
1. Leaves orbicular with 3-5 shallow lobes (California, Nevada)
........................................ R. oxynotus (2n=16)
1. Leaves elliptical or reniform, lobes deeper
2. Leaves elliptical in outline, untoothed lobes longer than
wide
3. Leaves with 3 to 5 deep, almost parallel, pointed lobes
(Wyom. Mont. BC?) ................. R. eximius (2n=32)
3. Leaves with 3 to 7 shallow end teeth, otherwise unlobed
(Utah Colo. N.Mex.) ............. R. macauleyi (2n=16)
2. Leaves reniform in outline deeply 3-5 lobed
4. Lobes deeply trisected
5. Plants short, compact, slightly glaucous (Nevada
Utah?) ... R. suksdorfii subsp. caespitosus (2n=16)
5. Plants loose, not glaucous (Wyom. Utah Colo. N.Mex)
................................ R. adoneus (2n=32)
4. Lobes toothed but not trisected
6. Teeth rounded, central lobe often untoothed, almost
lingulate (all western mountains, Alaska to northern
Mex. and southern Cal.)
.......................... R. eschscholtzii (2n=48)
6. Teeth sharply distinct, central lobe +/- 3-5 toothed
7. Leaf surfaces green, sharply toothed lobes wider
than long (Cal. Oreg. Wash. Mont. Wyom. BC?)
........ R. suksdorfii subsp. suksdorfii (2n=16)
7. Leaf surfaces glaucous, trisected lobes longer
than wide (Oreg, Idaho)
.......... R suksdorfii subsp. trisectus (2n=16)
(The predominantly far-northern 12-ploid R. nivalis L.
[incl.R. sulphureus Soland.ex Phipps] has very distinctive black
hairs on the sepals. An outlying population occurs at Beartooth
Pass, Wyoming.)
References:
Fisher, F.J.F., J.A. Rowley, & C.J. Marchant. 1973. The biogeog-
raphy of western snow-patch Ranunculi of North America.
Compte rendu sommaire des seances. Societe de Biogeographie
438:32-43.
Fisher, F.J.F., A. Warner, & E.M. Reimer. 1979. Anomalous
apetaly: localized character displacement in Ranunculus
eschscholtzii. Canadian Journal of Botany 57: 2097-2106.
Rowley, J.A., F.J.F. Fisher, & C.J. Marchant. 1975. Distribu-
tion, ecology and photosynthesis of alpine Ranunculi in
Western North America. Proc. New Zealand Ecological Society
22(1975): 113-115.
WHERE HAS ALL THE CLOVER GONE?
From: Nancy J. Turner <nturner@uvic.ca>
In September, 1991, T.C. Brayshaw collected a modest specimen of
Trifolium wormskioldii on Dallas Road, at the west point south
of Menzies Street ("along seashore; open, south-facing, eroding
bank above beach, mainly grassy" V 151489). As he has been doing
for many, many years, with this collection he was helping us,
with his careful, meticulous work, to understand our flora
better and to document the changes that have occurred to our
local ecosystems. We owe him so much for his critical contribu-
tions. In the case of this collection, it is particularly sig-
nificant because, as far as I can determine, it represented the
last population of T. wormskioldii along the Victoria
waterfront. Bob (Turner) and I went to this site a couple of
weeks ago and found and enormous cement storm drain enclosure
and no sign of this native clover, although there were plenty of
introduced clovers and grasses and even some Camassia, Sanicula
bipinnatifida and other characteristic native species in the
vicinity.
Trifolium wormskioldii Lehm. was once common - even abundant -
all along this stretch of Victoria's coastline. James Douglas,
when he arrived here in 1842, commented on the clover, which
Chris Brayshaw, Adam Szczawinski and others conclude must have
been, at least in part, T. wormskioldii:
"Both Kinds [of soil], however, produce Abundance of Grass, and
several varieties of Red Clover grow on the rich moist
Bottoms.... In Two Places particularly, we saw several Acres of
Clover growing with a Luxuriance and Compactness more resembling
the close Sward of a well-managed Lea than the Produce of an
uncultivated Waste." (Douglas's report, Fort Vancouver, HBC,
July 12, 1842)
In fact, the point where Douglas first disembarked from the
Beaver he himself named Clover Point, from the fact that "a
large area of ground here was found covered with a species of
red clover, growing most luxuriently" (Walbran 1971: 96). For
Douglas, the clover and meadowlands he found were an indicator
of the potential of the area for modification and management,
for re-creating England and the English style of land use.
"Being pretty well assured of the Capabilities of the Soil as
respects the Purposes of Agriculture, the Climate being also
mild and pleasant, we ought to be able to grow every Kind of
Grain Raised in England..."
The clover was seen as a promising indication of the pos-
sibilities for improvements: "The growth of indigenous vegeta-
tion is more luxuriant, than in any other place, I have seen in
America, indicating a rich productive soil. Though the survey I
made was rather laborious, not being so light and active of foot
as in my younger days, I was nevertheless delighted in ranging
over fields knee deep in clover, tall grasses and ferns reaching
above our heats, at these unequivocal proofs of fertility".
(James Douglas in Hargrave 1938: 420-421, as cited by Akrigg and
Akrigg 1975: 357.)
Unfortunately, the scale and scope of landscape change that has
emanated from this concept has not been easily measured or
acknowledged, and the cultural and biological impacts have not
been fully assessed. For First Peoples, the loss of valued
cultural resources caused profound grief and sadness, still felt
today by Elders who remember some of the lands as they used to
be. For most people, however, the loss of lands and resources,
if recognized at all, is seen as the inevitable result of
progress. Many people still do not grasp the importance of
maintaining "wild" ecosystems or species, and, if they value
greenspace, it makes little difference to them whether it is a
green lawn, and agricultural field, or a natural wood or meadow.
Many people still view any uncultivated area as "waste", just as
James Douglas did over 150 years ago. The value in this land and
its ecosystems, for them, is in their potential, not in them-
selves.
This is well reflected in the words of Berhold Seeman, a
naturalist with the expedition which sailed in H.M.S. Herald,
under Captain Henry Kellett, D.B., 1845-51, who wrote in June
27, 1846: "In walking from Ogden Point round to Fort Victoria, a
distance of little more than a mile, we thought we had never
seen a more beautiful country; it quite exceeded our expecta-
tion; and yet Vancouver's descriptions made us look for some-
thing beyond common scenery. It is a natural park; noble oaks
and ferns are seen in the greatest luxuriance; thickets of the
hazel [?] and the willow, shrubberies of the poplar and the
alder, are dotted about. One could hardly believe that this was
not the work of art; more particularly when finding signs of
cultivation in every direction--enclosed pasture-land, fields of
wheat, potatoes and turnips. Civilization had encroached upon
the beautiful domain, and the savage could no longer exist in
the filth and indolence of mere animal life.... The fort of
Victoria was founded in 1843, and stands on the east shore of
the harbour, or rather creek, about a mile from the entrance.
The approach is pretty by nature, though somewhat rude by art.
The first place we came to was the dairy, an establishment of
great importance to the fort.... We were astonished at all we
saw. About 160 acres are cultivated with oats, wheat, potatoes,
turnips, carrots, and other vegetables, and every day more land
is converted into fields. Barely three years had elapsed since
the settlement was made, yet all the necessaries and most of the
comforts of civilized life already existed in what was a
wilderness..." (Scholefield, 1914)
Yet, unrecognized by the colonists, the lands they were changing
were already under a degree of care and management by the local
First Peoples, and the existing resources were already being
used, just not in the ways that the newcomers could perceive. As
a result, resources that were desired and once plentiful for
First Peoples were intentionally and unintentionally diminished,
at least in part because their value was not acknowledged. Of
course, as these species dwindled in range and abundance, they
became less salient to the First Peoples themselves, and as this
occurred, the cultural memory about them diminished. Thus, in a
spiraling downward cycle, species deteriorated and disappeared
and only a few noticed or cared. Those who cared were not able
to communicate their concerns to those who were causing the
changes. In short, there was a terrible lack of communication
that has continued to the present day.
As noted, Trifolium wormskioldii, because it is a perennial, and
because when it grows luxuriantly it forms a very showy display,
is likely the major species referred to by James Douglas and
after which Clover Point off Dallas Road is named. Its rhizomes
were also a valued food of First Nations and were undoubtedly
used by the Lekwungen peoples who used the area around Ross Bay
and Beacon Hill Park along with the rest of the Victoria area.
Dr. Newcombe collected this species in June,1917 off Dallas
Road, Victoria, on "moist ground near the sea" (V: 42028, 42029;
8951), and specimens in the Royal B.C. Herbarium (V) show that
it has grown in many sites around the city . It is characterized
as "common" (J.R. Anderson, 1920) and growing in "open grass"
(J.R. Anderson, 1920), "rich meadows" (J.R. Anderson, 1894),
"rich open lands (J.R. Anderson, 1896), "damp ground" (M.C.
Melburn, 1970), "moist meadow" (T.C. Brayshaw, 1981); or "dry
open ground (V.E.I. Goddard, 1930). Yet, although other native
clover species can still be readily found on rocky sites and
refugia around the city, T. wormskioldii, cannot be called
"common" around Victoria today by any stretch of the imagina-
tion. It is, in fact, virtually absent from the entire Victoria
coastline. The work of Chris Brayshaw has helped us to piece
together its passing, and, maybe it will help us to bring it
back again.
References:
Akrigg, G.P.V. & H.B. Akrigg. 1975. British Columbia chronicle
1178-1846: Adventures by sea and land. Discovery Press,
Vancouver, B.C.
Douglas, James. Douglas' original field journal. Provincial
Archives, British Columbia. File: A-B-40-D75.4A
Hargrave, J. 1938. The Hargrave correspondence, 1821-1843;
edited with introduction and notes by G. P. de T. Glazebrook.
The publications of the Champlain society, xxiv. The
Champlain society, Toronto.
Scholefield, E.O.S. 1914. British Columbia from the earliest
times to the present. Volume I and II. S.J. Clarke Publishing
Company, Vancouver, B.C.
Walbran, J.T. 1971. (originally published 1909) British Columbia
coast names, 1592-1906, to which are added a few names in
adjacent United States territory; their origin and history.
Vancouver Public Library & J.J. Douglas, Vancouver, B.C.
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