pdf wkg Dav.14-14 - Davidsonia

Volume 14, Number 4 

October 2003 

Davidsonia 

A Journal of Botanical Garden Science

Davidsonia 

Editor 

Iain E.P. Taylor 

UBC Botanical Garden and Centre for Plant Research 

University of British Columbia 

6804 Southwest Marine Drive 

Vancouver, British Columbia, Canada, V6T 1Z4 

Editorial Advisory Board 

Quentin Cronk 

Fred R. Ganders 

Daniel J. Hinkley 

Carolyn Jones 

Lyn Noble 

Dorina Palmer 

Moura Quayle 

David Tarrant 

Roy L. Taylor 

Nancy J. Turner 

Associate Editors 

Mary Berbee (Mycology/Bryology) 

Moya Drummond (Copy) 

Aleteia Greenwood (Art) 

Michael Hawkes (Systematics) 

Richard Hebda (Systematics) 

Douglas Justice (Systematics and Horticulture) 

Daniel Mosquin (Publication) 

Hailey Pappin (Production) 

Andrew Riseman (Horticulture) 

Charles Sale (Finance) 

Janet R. Stein Taylor (Phycology) 

Sylvia Taylor (Copy) 

Roy Turkington (Ecology) 

Jeannette Whitton (Systematics) 

Davidsonia is published quarterly by the Botanical Garden of the University of British 

Columbia, Vancouver, British Columbia, Canada V6T 1Z4. Annual subscription, 

CDN$48.00. Single numbers, $15.00. All information concerning subscriptions should 

be addressed to the editor. Potential contributors are invited to submit articles and/ 

or illustrative material for review by the Editorial Board. 

ISSN 0045-09739 

Cover: Disanthus cercidifolius. Photo: Daniel Mosquin. 

Back cover: Polystichum kruckebergii frond. Photo: Gary Lewis.

Davidsonia 14:4 

109 

Editorial 

Small, Fragmented and Local Ecosystems 

Krajina’s biogeoclimatic zone system (for review see Beil et al, 1976) for 

vegetation classification is one of the foundation frameworks for management 

of British Columbia’s forest. As with all systems, it has undergone 

considerable modification as more concise information and modern methods 

of vegetation analysis have been developed, but there is no doubt that it has 

provided a useful tool for those charged with planning forest harvest policy 

and practices. However, it is perhaps not as useful as a tool for small-scale 

management, because it does not work particularly well when dealing with 

ecosystems that are not on the successional path to the defined forest climax 

states. 

The local, often small scale, occurrence of serpentine soils is a case in 

point. The paper by Lewis and Bradfield in this issue reports substantial 

new floristic and ecological analyses about an ecosystem that can be defined 

as rare, if not necessarily endangered. It is almost 25 years since Kruckeberg’s 

paper appeared in Davidsonia (Kruckeberg, 1979) and the Lewis and 

Bradfield paper is the first of two, which we have invited, to report new 

developments in the study of serpentine flora and ecology. Lewis has agreed 

to prepare a paper, scheduled for 2004, based on his doctoral thesis research 

that will allow us to re-publish Kruckeberg’s article and present an update. 

We plan a series of papers on these ‘smaller’ systems, including the muchfragmented 

Garry Oak system on Vancouver Island, the interior temperate 

rainforest on the eastern side of the Cariboo region, and the many coastal 

marshes on the Pacific rim. 

Conservation legislation in Canada, unlike that in the USA, has a strong 

emphasis on habitat conservation. We have an obligation to describe and 

document these more fragmented ecosystems so that their existence will be 

known, their importance understood and the enormous biological diversity, 

Iain E.P. Taylor, Professor of Botany and Research Director. 

UBC Botanical Garden and Centre for Plant Research. 

6804 SW Marine Drive, Vancouver, BC, Canada, V6T 1Z4. 

iain.taylor@ubc.ca

110 

which has been given to the care of British Columbians, can be passed to 

our children’s children. I hope that researchers and experts will help 

Davidsonia to assist industry, government and the public in meeting these 

obligations. 

References 

Beil, C.E., Taylor, R.L., and Guppy, G.A., 1976. The Biogeoclimatic Zones of 

British Columbia. Davidsonia 7: 45-55 

Kruckeberg, A.R., 1979. Plant that grow on serpentine - A hard life. Davidsonia 

10: 21-29


111 

Autumn Colours – Nature’s Canvas is a Silk Parasol 

The Adaptive Value of Autumn Foliage 

Abstract 

The variety and widespread nature of leaf colour change in autumn has led to 

investigation of the biochemical pathways and compounds responsible. The synthesis of 

bright red colouration initiated by longer nights prior to leaf abscission in deciduous 

species points to some adaptive value for this expensive ephemeral trait. It is 

hypothesized that during the breakdown of the unstable chlorophyll and the dismantling 

of the nutrient-rich photosynthetic apparatus, red anthocyanins provide a more 

biochemically parsimonious alternative to the elaborate xanthophyll system. This 

alternative enables leaves to screen out excess light energy and circumvent 

photooxidative damage to leaf cells, while allowing photosynthesis to persist at low rates 

in support of metabolic processes and phloem loading required for nutrient resorption 

from leaves. 

Introduction 

People continue to marvel at the spectacle of familiar green woods and 

local trees changing into a coat of blazing colours in autumn. Scientific 

investigations beginning in the 19 th century (see Wheldale 1916) have led to 

a very good understanding of how this occurs. Research continues today in 

the fields of physiology, biochemistry, and molecular genetics to further 

elucidate the complex signalling pathways and mechanisms involved. 

During the growing season, healthy leaves are green, and appear so due to 

the high concentrations of chlorophyll within the chloroplasts, organelles 

which act as microscopic factories converting water and carbon dioxide into 

carbohydrates and oxygen, using the energy in specific wavelengths of 

sunlight. Chlorophyll efficiently absorbs red and blue light during this process, 

but reflects or transmits the green light we observe. In autumn the chlorophyll 

breaks down and leaves show the yellow and orange colours of carotenoids 

already present in chloroplasts, but previously invisible because of the 

overwhelming green of the chlorophyll. In many species, new flavonoid 

pigments called anthocyanins are synthesized during this period imparting a 

Robert D. Guy and Jodie Krakowski. 

Department of Forest Sciences, Faculty of Forestry, University of British Columbia, 

2424 Main Mall, Vancouver, BC, Canada, V6T 1Z4. 

Corresponding author: guy@interchg.ubc.ca

112 

red colouration, becoming purple as the pH increases (the compounds react 

with the cellular solution producing a visible colour change). These 

compounds are stored in vacuoles in the mesophyll and/or epidermal cells 

of leaves and fruits (Hrazdina et al., 1982). 

Explaining why leaves develop brilliant hues in the autumn is not so straightforward. 

One North American aboriginal legend claims that when mighty 

ancestral hunters slew the celestial Spirit Bear, commemorated in the constellation, 

his red blood caused the tree leaves to redden in sympathy (Philp 

2001). Jack Frost, charged with comprehensively painting every leaf with 

the onset of frost, would probably be relieved at recent developments in 

plant physiology which point to the adaptive value of leaf colour change in 

the autumn. His innocence is proven by the observation that leaves often 

change colour before temperatures reach the freezing point. Several 

environmental cues interact with the physiology of leaves to induce colour 

change well before the damaging temperatures of autumn arrive. Plant 

biologists and ecologists have proposed several direct fitness benefits of 

this transient adaptation. 

Chlorophyll is a complex molecule. It is responsible for the vast majority 

of light capture to power carbon fixation in plants. Chlorophyll a and b 

molecules are associated with light harvesting complex proteins, which act 

as antennae funnelling light energy into photosystems I and II (Figure 1). 

Electrons, removed from water by the oxygen-evolving complex, are then 

propelled through these photosystems in a series of dozens of oxidation 

and reduction reactions that constitute the “photosynthetic electron transport 

chain”. Proton (H + ) transport driven by the flow of electrons, in combination 

with the release of protons from water, creates a pH gradient across 

chloroplast membranes. The pH gradient is harnessed to make ATP (an 

energy currency) while the electrons are ultimately used to reduce NADP to 

NADPH. These products of the “light reactions” are consumed by the 

Calvin cycle, which, through numerous chemical steps, “fixes” CO 2 

into the 

products of photosynthesis. 

Photosynthesis ultimately produces carbon and energy stores available to 

the plant in the form of sugar, or stored in plastids such as chloroplasts as 

starch grains. These reserves provide the means by which all higher functions 

of the plant can occur: cellular division and repair, growth, nutrient uptake 

reproduction, translocation, etc..


113 

Although ideally suited to their primary role, chlorophyll molecules easily 

break down due to their inherent instability, especially under the high-intensity 

photon bombardment of sunlight. The photons stimulate chlorophyll 

molecules to release excited electrons, disrupting the molecular bonds which 

hold the component atoms together. This is an exothermic reaction, after 

which the molecules achieve a more stable state in another form. This 

photooxidation of chlorophyll takes place throughout the leaf ’s green life. 

Normally, the chlorophyll thus destroyed is replaced by newly synthesized 

chlorophyll. In autumn, however, the synthesis of replacement chlorophyll 

stops. 

Environmental triggers contribute to autumn colour change 

In addition to providing the driving energy for photosynthesis, light influences 

the timing, magnitude and degree of leaf colour change and not only 

by destroying chlorophyll. Brighter sunlight tends to produce the most vivid 

colours, primarily when temperatures are low. Leaf age is also a factor, 

since the physiology and functions inherent in younger leaves differ from 

older leaves and they are less apt (or “competent”) in responding to the 

longer nights and lower temperatures of autumn. Younger leaves, therefore, 

turn colour after mature leaves on the same tree (Figure 2). The foliage of 

Larix lyallii (sub-alpine larch) turns colour later into the autumn in years 

where leaf emergence has been delayed by cold spring weather (Worrall, 1993). 

On the other hand, drought and nutrient stress (Schaberg et al., 2003) 

contribute to earlier fading of green and the appearance of other colours. 

Often it is the tissues immediately adjacent to the veins that are the last to 

turn colour (Figure 3). 

A reddening of foliage and other green tissues is not restricted to the 

autumn or to deciduous trees (Steyn et al., 2002). Even Pinus banksiana (jack 

pine) seedlings become stained with purple in the autumn (Nozzolillo et al., 

2002). Other examples include the frequent bright reddish or purple tinge 

in many high-elevation plant species emerging through brilliant white snow, 

the red “snow algae” in montane and alpine sites, and the reddish cast in the 

winter foliage of several conifers (especially the Taxaceae, including the yews 

and redwoods, and Cupressaceae, including the cypresses) (Weger et al., 1993; 

Han et al., 2003). Many sun-exposed plants will also redden in summer in 

response to drought, nutrient or salt stress (Figure 4).

114 

The interaction between the two stressors of high light and near-freezing 

temperatures has a more pronounced effect than either factor individually. 

This is easily observed: leaves at the top and outer edges of a canopy (unless 

they’re young!), subject to the strongest light intensity, turn far brighter colours 

than shaded leaves (Feild et al., 2001). This suite of phenomena points 

towards a role for autumn pigmentation in photoprotection. 

Natural history enthusiasts have long noted that leaves vary in colour, intensity 

and rate of change. This has been found among and within stands 

of trees of the same age, among parts of the same tree, even on different 

sides of a single leaf. Ramets of Populus tremuloides (trembling aspen) clones, 

where each individual tree is actually an aboveground shoot of a single root 

system, and thus all represent clones of a single genotype, have also been 

noted to vary widely in the same respect (Figure 5; Chang et al., 1989). Year 

to year variation was found in the peak colours and chemical signatures of 

different compounds within the same tree: yellow pigments were always 

detected, even in trees with green, orange or red foliage, while red pigments 

were only evident in red and orange leaves, and only expressed in some years 

(Chang et al., 1989). This variable and facultative expression of autumn 

colours may be correlated with environmental factors such as temperature 

and moisture availability. 

Preparations for winter 

A quick glance at the profusion of nature’s ebullience may cause one to 

wonder, ‘why bother’. These leaves are on the brink of imminent death – 

they will soon fall off, and assume new functions as fodder or fertilizer as 

they decompose. However, the leaf still has one critically important function 

before this occurs. Large amounts of nutritive reserves must be recovered 

for winter storage so they can boost the array of activity which begins with 

new growth each spring. 

Phloem-loading 

When fully functional, deciduous leaves are rich in nitrogen, sulphur, 

potassium, phosphorus and numerous other essential plant nutrients. The 

main role of these elements, in free form or as components of proteins and 

other cellular constituents, is for photosynthesis. Nutrients are in short supply 

in most soils and it is therefore advantageous for plants to resorb and recycle


115 

them. Indeed, over one-third of the yearly nitrogen and phosphorous requirements 

of forest trees are typically met this way. Studies show that 

following longer nights, nitrogen-containing amino acids and other nutrientrich 

compounds move into leaf veins and then through the petiole, to be 

stored in the living bark (which includes the phloem) over winter (Greenwood 

et al., 1986). This translocation process is driven by osmotic gradients in the 

phloem created by the amount of dissolved solutes. The only way materials 

can exit leaves back to the stem is via this process, called phloem-loading. If 

available sugar is completely consumed and not replaced by photosynthesis, 

then transport will not be possible. 

Phloem is a vascular tissue whose function is to move the products of 

photosynthesis and other phytochemicals throughout the plant, from their 

production site (source) to where they will be consumed or stored (sink). 

This movement depends on the active (energy-requiring) loading of sugars 

or sugar alcohols into phloem sieve and companion cells at the source end, 

and passive unloading at the sink end. Water follows by osmosis, generating 

a pressure gradient that pushes the phloem sap along. The sugar 

concentration of phloem sap is on the order of 15-25%. Hence large amounts 

of photosynthate are required. 

A plant’s ability to transport material through the phloem requires a 

functioning photosynthetic system to provide the sugar and energy needed 

for phloem-loading. Researchers have shown the existing energy reserves 

of leaves, consisting of sugars and stored starches primarily in the plastids, 

are too meagre to account for the recovery of leaf reserves without ongoing 

photosynthesis. The plant therefore faces a conundrum: at the same time 

enzymatic machinery is to be dismantled for seasonal recycling, some pieces 

of equipment must remain functioning to provide the energy and raw 

materials for this process. 

Photoprotection 

Normally functioning photosynthetic mechanisms dissipate light energy 

in excess of photosynthetic requirements. Light energy not captured for 

photosynthesis or otherwise diverted presents a danger to cells by reacting 

with unstable molecules and releasing free oxygen radicals, which then begin 

a series of oxidation reactions, damaging the cell (Yamasaki 1997). This type 

of damage due to light is called photooxidation. In particular, permanent

116 

damage may be incurred by photosystem II as a result of excessive photon 

barrage. In leaves, the photosynthetic apparatus itself is the main source of 

these radicals. 

If the speed of the carbon reactions is restricted by environmental stress 

(e.g., drought, poor nutrition, low temperatures), or if more light energy is 

absorbed than the photosystems are capable of processing, then the 

photosynthetic electron transport chain may become “over-reduced”. Under 

these conditions of high excitation pressure, excess electrons may flow to 

oxygen (O 2 

) to produce superoxide (O 2* 

) and other highly reactive radicals. 

These radicals destroy membranes and other cellular components on contact. 

Plants have efficient enzymatic systems for detoxifying these radicals, but 

these systems may be overwhelmed under high light, low temperature 

conditions. The risk will be especially high if light penetrates more deeply 

into canopies and leaf tissues as chlorophyll begins to degrade and leaves 

abscise. As summer ends, photosystems I and II, the chlorophyll-containing 

components of the photosynthetic apparatus, and the proteins of Calvin 

cycle are beginning to be dismantled and can no longer fully utilize the 

abundant light energy. Low temperatures, especially at dawn, exacerbate the 

situation by inhibiting remaining Calvin cycle activity (Figure 1). The leaf 

must rely on the properties of alternative pigments, some of which are already 

present in the photosynthetic complexes but were not previously visible due 

to masking by the strong green of chlorophyll. 

First and most common are the carotenoids which absorb blue to green 

light and reflect yellow to orange wavelengths. The carotenoids include 

carotenes such as lutein (a brilliant red pigment) and xanthopylls such as 

zeaxanthin (which gives corn its yellow colour). Carotenoids, and especially 

zeaxanthin, are able to accept energy from chlorophyll and dissipate it safely 

as heat. Zeaxanthin is synthesized from another xanthophyll, violaxanthin, 

via the xanthophyll cycle. The amount of zeaxanthin present in a leaf at any 

one time is dynamically regulated by this pathway and may change within 

minutes (Demmig-Adams and Adams, 1992). The enzymes in the xanthophyll 

cycle are less efficient at low temperatures, but, depending on the species 

and whether the leaf is adapted to shade or sun, the presence of these 

pigments may be sufficient to prevent photooxidation and protect the 

senescent leaf for the remaining tasks at hand (Demmig-Adams and Adams, 

1992).


117 

The complex and energetically expensive xanthophyll cycle is a finely-tuned 

and highly sophisticated system. The simpler, although less precisely 

regulated, alternative for plants is to utilize anthocyanins or other screening 

compounds that block the light before it reaches chlorophyll (Figure 1). 

These molecules absorb higher-energy blue light and reflect or transmit 

lower-energy red light, so tissues with anthocyanins appear red to purple. 

Unlike the xanthophyll cycle, anthocyanins require no enzymes to function 

and are energetically much less costly to the plant to produce and maintain. 

Like the carotenoids, energy absorbed by anthocyanins is simply lost as heat. 

They are also effective antioxidants (Neill and Gould, 2003). Thus, this 

group of phytopigments acts as a molecular sunscreen or parasol to greatly 

reduce the amount of light impinging on remaining chlorophyll, preventing 

the initiation of unstable redox reaction chains and the release of damaging 

free oxygen radicals within the cell. 

For plants which can potentially synthesize both xanthophylls and 

anthocyanins, there is clearly an advantage in the latter, cheaper compound, 

especially as leaves become more stressed and biochemical pathways less 

stable. Thus the vermillion and scarlet hues deepen as autumn progresses. 

Adaptive benefits of colour change 

Many different explanations have been proposed for the occurrence of 

autumn colour change in leaves. The association with light availability was 

noted very early on (Wheldale, 1916). Previously scientists thought that the 

synthesis of these chemicals did not have any evolutionary benefit, but it is 

highly improbable that such a persistent and widespread phenomenon would 

be selectively neutral. The distribution and frequency of autumn colour 

change suggests that this response evolved in many plant taxa independently, 

which infers it has some fitness benefit (e.g., Jaenike 2001). Similarly, a theory 

that compounds producing autumn colour were serving waste functions, 

emptying vacuoles of toxic products prior to leaf abscission (Ford 1986) 

seems unlikely on the same grounds. Red, orange or purple colours in fruit 

and possibly adjacent leaves are thought to provide a signal to herbivores 

that seeds are developmentally ready for dispersal, but many leaves that change 

colour have wind-dispersed fruit, are too immature to reproduce, or change 

colour long after seeds are dispersed. 

An interesting recent hypothesis proposed that red leaves afford some

118 

protection against aphid herbivory, suggesting that coevolution led to host 

specificity based on visual leaf colour cues as a signal mechanism for tree 

health during autumn oviposition (Archetti 2000; Hamilton and Brown, 2001). 

While many genera and species were reviewed, this does not take into account 

oviposition at other times and herbivory by other organisms; in autumn leaf 

herbivory in general is quite low. 

As reviewed here, recent physiological explanations for autumn anthocyanin 

production focus on its role in capturing light and preventing 

photooxidative damage to the photosynthetic apparatus, especially during 

stress induced by low temperatures during autumn (Smillie and Hetherington, 

1999; Hoch et al., 2001). Weger et al. (1993) proposed a similar role for 

winter rhodoxanthin in red cedar. The more efficient operation of 

anthocyanins at low temperatures relative to chlorophyll also suggests an 

adaptive role. Localized anthocyanin and rhodoxanthin expression at leaf 

surfaces which receive the most sunlight substantiates a protective role for 

these compounds (Gould et al., 1995; Feild et al., 2001; Hoch et al., 2001). 

DO try this at home! 

Simply appreciating nature’s grandeur is eminently rewarding for those 

lucky enough to live in temperate continental climates featuring deciduous 

forests. The most spectacular examples are found in North America: the 

forests of New England, the Canadian maritime region and Quebec are major 

tourist attractions in autumn. These changes, although more subtle, can also 

be observed in some deciduous conifers. In the Rockies and Cascades, golden 

yellow stands of sub-alpine larch make for particularly enchanting and popular 

hiking destinations (Figure 6). Angiosperms such as Betula (birches), 

Liriodendron tulipifera (tulip trees) and most Populus (poplars, cottonwoods 

and aspens) turn yellow before becoming brown; even in these trees it is 

simple to observe which parts of the tree change colour first. 

Research has demonstrated that ecological successional roles also influence 

the timing and expression of autumn colour: pioneer species, which colonize 

disturbed habitats and tend to grow rapidly, are adapted to maximize carbon 

capture in high light environments by inherent photoprotection mechanisms 

(Hoch et al., 2001). These species generally have less spectacular colours 

than the more shade tolerant species which typically grow in older forests.


119 

There is a simple experiment anyone can do to test and observe these 

physiological changes and the effects of the environment on their expression. 

This works especially well on species with dramatic autumn changes, such as 

Liquidambar styraciflua (sweetgum), Rhus (sumac), Cornus (dogwood) or Acer 

saccharum (sugar maple). Before the leaves begin to change colour, cover up 

a portion of a leaf, still attached to the tree, with a piece of opaque material, 

such as thick paper. Do this on both sides of the leaf and secure it with a 

paper clip. An alternative approach is to cover the leaf with an image drawn 

in black on transparent film. Wait until the rest of the leaves on the branch 

show their autumn pigmentation and remove the masking: the covered 

portion should retain the yellow carotenoid colours, and little or no green 

since sustained chlorophyll synthesis requires direct light. You can even add 

an element of creative design or stencil mysterious messages (Figure 7) along 

your favourite wooded trail! 

Bibliography 

Archetti, M. 2000. The origin of autumn colours by coevolution. Journal of 

Theoretical Biology 205:625-630. 

Chang, K.-G., Fechner, G.H., and Schroeder, H.A. 1989. Anthocyanins in 

autumn leaves of quaking aspen. Forest Science 35:229-236. 

Demmig-Adams, B., and Adams, W.W. III. 1992. Photoprotection and other 

responses of plants to high light stress. Annual Review of Plant Physiology 

and Plant Molecular Biology 43:599-626. 

Feild, T.S., Lee, D.W., and Holbrook, N.M. 2001. Why leaves turn red in autumn. 

The role of anthocyanins in senescing leaves of red-osier dogwood. Plant 

Physiology 127:566-574. 

Ford, B.J. 1986. A theory of excretion in higher plants. Journal of Biological 

Education 20:251-254. 

Gould, K.S., Kuhn, D.N., Lee, D.W., and Oberlauer, S.F. 1995. Why leaves are 

sometimes red. Nature 378:241-242. 

Greenwood, J.S., Stinissen, H.M., Peumans, W.J., and Chrispeels, M.J. 1986. 

Sambucus nigra agglutinin is located in protein bodies in the phloem 

parenchyma of the bark. Planta 167:275-278. 

Hamilton, W.D., and Brown, S.P. 2001. Autumn tree colours as a handicap signal. 

Proceedings of the Royal Society of London, B 1489-1493. 

Han, Q., Shinohara, K., Kakubari, Y., and Mukai, Y. 2003. Photoprotective role 

of rhodoxanthin during cold acclimation in Cryptomeria japonica. Plant, Cell

120 

and Environment 26:715-723. 

Hoch, W.A., Zeldin, E.L., and McCown, B.H. 2001. Physiological significance of 

anthocyanins during autumnal leaf senescence. Tree Phyisology 21:1-8. 

Hrazdina, G., Marx, G.A., and Hoch, H.C. 1982. Distribution of secondary 

plant metabolites and their biosynthetic enzymes in pea (Pisum sativum L.) 

leaves. Plant Physiology 70:745-748. 

Jaenike, J. 2001. Sex chromosome meiotic drive. Annual Review of Ecology and 

Systematics 32:25-49. 

Neill, S.O., and Gould, K.S. 2003. Anthocyanins in leaves: light attenuators or 

antioxidants Functional Plant Biology 30:865-873. 

Nozzolillo, C., Isabelle, P., Andersen, O.M., and Abou-Zaid, M. 2002. 

Anthocyanins of jack pine (Pinus banksiana) seedlings. Canadian Journal of 

Botany 80:796-801. 

Philp, J. 2001. Why leaves change color. University of Maine Cooperative 

Extension Bulletin #7078. 

Schaberg, P.G., van den Burg, A.K., Murakami, P.F., Shane, J.B., and Donnely, J.R. 

2003. Factors influencing red expression in autumn foliage of sugar maple 

trees. Tree Physiology 23:325-333. 

Smillie, R.M., and Hetherington, S.E. 1999. Photoabatement by anthocyanin 

shields photosynthetic systems from light stress. Photosynthetica 36:451- 

463. 

Steyn, W.J., Wand, S.J.E., Holcroft, D.M., and Jacobs, G. 2002. Anthocyanins in 

vegetative tissues: a proposed unified function in protoprotection. New 

Phytologist 155:349-361 

Weger, H.G., Silim, S.N., and Guy, R.D. 1993. Photosynthetic acclimation to low 

temperature by western red cedar seedlings. Plant, Cell and Environment 

16:711-717. 

Wheldale, M. 1916. The anthocyanin pigments of plants. Cambridge University 

Press, Cambridge, UK. 318 pp. 

Worrall, J. 1993. Temperature effect on bud-burst and leaf fall in subalpine 

larch. Journal of Sustainable Forestry 1:1-18. 

Yamasaki, H. 1997. A function of colour. Trends in Plant Science 2:7-8.


121 

A floristic and ecological analysis at the Tulameen 

ultramafic (serpentine) complex, southern British 

Columbia, Canada 

Abstract 

While distinct floristic and ecological patterns have been reported for ultramafic 

(serpentine) sites in California and Oregon, those of British Columbia are muted which is 

thought to be related to the moderating influence of increased precipitation, a short time 

since glaciation, and the presence of non-ultramafic glacial till over ultramafic sites. 

Despite these factors, we found clear floristic and ecological differences with respect to 

soil type at our study site on Grasshopper Mountain, part of the Tulameen ultramafic 

complex in southern British Columbia. Ultramafic soils support 28% of the local species 

richness and host more rare taxa than non-ultramafic soils. Many species show patterns 

of local restriction to or exclusion from ultramafic soil habitats. Patterns of plant family 

diversity also show differences between substrates. 

Introduction 

Ultramafic (serpentine) soils and the plants that they support have long 

been of interest to botanists (Whittaker 1954; Proctor and Woodell 1975; 

Brooks 1987). They frequently support vegetation that is distinct from 

surrounding areas in species composition and structure as well as high levels 

of plant endemism and diversity. For these reasons, and because of 

phenomena related to speciation and plant physiological response, Brooks 

(1987) and Proctor (1999) asserted that the biological importance of 

ultramafics far outweighs the less than one percent of the earth’s surface 

they occupy. 

The chemical and physical properties of ultramafic soils often have adverse 

effects on plant growth (termed the “serpentine effect”). These soils generally 

contain elevated concentrations of the heavy metals nickel, chromium, and 

cobalt, and high levels of magnesium, all potentially toxic to plants. They 

are generally deficient in nitrogen, phosphorus, potassium, and calcium, 

thereby further restricting plant growth. The reduced vegetation cover 

combined with rugged terrain frequently associated with ultramafic sites 

Gary J. Lewis and Gary E. Bradfield. 

Department of Botany, University of British Columbia, 

3529-6270 University Blvd., Vancouver, BC, Canada, V6T 1Z4. 

Corresponding author: garylewis@shaw.ca

122 

results in poorly-developed, unstable, and often dry soils. These soils also 

exhibit high heterogeneity, both between- and within-sites, a result of the 

inherent variability of ultramafic rocks and the pedological processes that 

weather them. Consequently, no single chemical or physical factor, nor single 

group of these factors can be said to be responsible for the vegetation of 

serpentine soils (Brooks 1987; Proctor and Nagy 1992; Roberts and Proctor 

1992). 

The vegetation of ultramafic soils can range in physiognomy from 

serpentine barrens to well-developed forests, but is usually floristically and 

structurally distinct from adjacent non-ultramafic soils. Some genera and 

families of vascular plants have shown a particular affinity or aversion to 

serpentine soils within certain regions (Dearden 1979; Kruckeberg 1969, 1992; 

Rune and Westerbergh 1992). Plant functional groups have also shown strong 

patterns relative to soil types. Species of dry habitats are often wellrepresented 

on ultramafic soils, whereas species of mesic to moist habitats 

are largely excluded (Kruckeberg 1979). Deciduous elements are 

conspicuously diminished in importance on ultramafics in Oregon (Whittaker 

1954). 

Kruckeberg (1979, 1992) described four categories of floristic response 

to serpentine: endemic, indicator, bodenvag (widespread), and excluded species. 

Serpentine endemics are those species restricted to ultramafic soils. Indicator 

species are those within a local or regional context that are restricted or 

nearly restricted to ultramafic substrates and whose presence, therefore, 

indicates serpentine soils. The term bodenvag (“soil wanderer”) refers to species 

that are either indifferent to soil type or that have developed serpentine 

ecotypes or races and, thus, occur commonly on and off ultramafic sites in a 

given region. Excluded species are those found commonly in surrounding 

areas but are unable to successfully colonize ultramafic substrates. 

Ultramafic soils have also been shown to host species outside of their 

main ranges. For example, Polystichum kruckebergii, a serpentine indicator fern 

known to also occur on non-serpentine soils (Lellinger 1985), was once 

thought to reach the northern limits of its range in southern British Columbia 

(Hitchcock and Cronquist 1973). However, it has more recently been found 

to track ultramafic outcrops through the interior of the province as far north 

as the Cassiar Mountains of northwestern British Columbia, a range extension 

of at least 580 km (Kruckeberg 1982; Douglas et al. 1998). Species’ altitudinal


123 

ranges are affected at ultramafic sites in the state of Washington. Several 

tree and shrub species occur at higher and lower elevations on serpentine 

than their normal elevation ranges on non-serpentine soils (Kruckeberg 1969). 

Little information is available for the plant communities and vegetational 

responses to serpentine soils in British Columbia. There is some support 

for the hypothesis that the vegetational response to ultramafics is less 

pronounced with increasing latitude in western North America, a result of 

the increasing precipitation, the presence of non-ultramafic glacial till 

deposited ca 12,000 years ago, and the relatively short time available for 

speciation since glacial retreat (Whittaker 1954; Kruckeberg 1979, 1992; D. 

Lloyd, pers. comm.; R. Scagel, pers. comm.). 

This report is part of a larger study directed to characterize the extent of 

the serpentine effect and to expand the knowledge of floristics and ecology 

of ultramafic sites in British Columbia. Through a detailed comparison of 

adjacent ultramafic and non-ultramafic soils, we sought to understand the 

uniqueness of ultramafic sites within a British Columbia context in order to 

help inform the decisions of conservationists and land managers. 

Study Site 

We compared plant communities and associated soils at Grasshopper 

Mountain, part of the Tulameen ultramafic complex (49 o 20’ N, 120 o 50’ W) 

of southern British Columbia (Figure 8). This is a relatively flat-topped 

mountain (elevation 1487 m) with a vertical rise of approximately 565 m. It 

is 6 km long and 2.5 km wide. Grasshopper Mountain was chosen because 

there were adjacent sections of ultramafic and non-ultramafic soils that 

minimized the confounding influences of aspect, topography, history, biota, 

and climate on the developing plant communities and permitted differences 

in vegetation to be directly attributed to edaphic factors. 

The Tulameen ultramafic complex lies within a climatic transition zone 

between humid coastal British Columbia and the dry interior. The complex 

is overlaid by coniferous forests dominated by Pseudostuga menziesii (Douglasfir), 

Pinus contorta var. latifolia (lodgepole pine), and P. ponderosa (ponderosa 

pine) at lower elevations, and by Pseudotsuga menziesii, Abies lasiocarpa (subalpine 

fir), and Picea engelmannii (Engelmann spruce) at higher elevations. Previous 

studies provide information on the geology (Cook and Fletcher 1993; Fletcher

124 

et al. 1995) and pedology (Bulmer 1992; Hope 1997) of Grasshopper Mountain 

and limited information on the vegetation (Kruckeberg 1979; Hope 

1997). 

Methods 

Vegetation and soils were sampled in a total of seventy-one 10-metre 

radius circular plots on adjacent ultramafic and non-ultramafic sections of 

the mountain’s southern face during July and August 2002. In each section, 

plots were selected randomly within a stratified design based on the degree 

of overstorey canopy cover (open, moderate, and closed forest; Figures 9, 

10, 11), slope position (top, upper, mid, lower, and toe), and elevation. The 

percent cover of understorey vegetation as well as forest structural data were 

recorded in plots. Soil chemical analyses were carried out in the laboratory 

of Dr Les Lavkulich, Faculty of Agricultural Sciences, University of British 

Columbia, for percent total C and N (Leco CN2000 Analysis), available P 

(Bray 1 Extraction), CEC and exchangeable K, Ca, Mg and Na (using the 

ammonium acetate method at pH 7.0), available Ni, Cr, Co, Mn, Al, Fe, Cu 

and Zn (DTPA Extraction), and pH (in 0.01 M CaCl 2 

). 

Means of soil variables for three soil types (ultramafic, glacial till-influenced, 

and non-ultramafic), and means for plot-level species richness and diversity 

(Shannon and Simpson diversity indices) were compared using analysis of 

variance (ANOVA) with a post-hoc Bonferroni adjustment provided by 

SYSTAT 10.2 (SYSTAT 2002). Floristic observations were evaluated using 

summary tables, and species distributions examined in relation to the different 

soil types. In this fashion ultramafic indicators, excluded, and bodenvag species 

were identified. PC-Ord (McCune and Mefford 1999) was used for plot 

summary statistics. Taxonomic nomenclature follows Douglas et al. (1998- 

2002). 

Results and Discussion 

Soils 

Soil chemical analysis indicated the occurrence of three general soil types 

at Grasshopper Mountain: ultramafic, non-ultramafic, and glacial tillinfluenced 

soils (Table 1). Ultramafic plots showed elevated levels of Mg 

and Ni and decreased levels of Ca and the Ca:Mg ratio, a general index of 

soil nutrient favourability (Proctor and Nagy 1992), relative to non-ultramafic


125 

plots. The values for till-influenced plots are intermediate, though only statistically 

so in the case of Mg. These till plots occurred in ravines and at 

lower elevations where non-ultramafic, glacial till accumulated through 

colluvial (emplaced by gravitational forces) processes over ultramafic bedrock. 

Floristic Patterns 

One hundred and seventy-seven vascular plant species from 35 families 

were recorded on Grasshopper Mountain: 111 species in 26 plots on 

ultramafic soils, 70 species in 10 plots on till, and 119 species in 35 plots on 

non-ultramafic soils. While these differences in total species richness may 

be partly explained by the different sample sizes, ANOVA results indicated 

no significant differences (p > 0.05) among plot mean values of species 

richness, Shannon diversity, and Simpson diversity for the three soil types. 

Ultramafic studies conducted elsewhere have reported conflicting results 

(Wilson et al. 1990). Whereas Huston (1979) predicted decreased species 

diversity on sites with extreme nutrient deficiency and toxicity, and Kruckeberg 

(1969) and Brooks (1987) characterized species composition of ultramafic 

sites as depauperate, Proctor and Woodell (1975) suggested that ultramafic 

sites may actually have higher diversity. 

Of the total species recorded, 49 (28 percent) were found solely or primarily 

in ultramafic plots. This finding suggests that Grasshopper Mountain, and 

potentially other ultramafic occurrences in BC, contribute greatly to the local, 

and regional, species pools. As noted by Kruckeberg (1979), the majority of 

this richness is derived from the presence of species common to other regions 

(especially the dry interior in our case) which attain a local foothold on the 

habitats available at ultramafic sites. The results are a flora distinct from 

that on adjacent non-ultramafic soils, and increased local and regional diversity. 

Taking into account the differences in sample size, our study also indicated 

trends in the representation of families on the three soil types (Table 2). 

Some families are more common on the ultramafic side (e.g. Apiaceae, 

Asteraceae, Caryophyllaceae, Poaceae, and Pteridophytes 1 ), whereas others 

are more common on till and non-ultramafic soils (e.g. Liliaceae, Rosaceae, 

Ranunculaceae, Betulaceae, Caprifoliaceae, Grossulariaceae and Salicaceae). 

The latter four families were not observed on ultramafic soils, while 

representatives of two families (Juncaceae and Polygonaceae) were observed 

only on ultramafic soils. Gymnosperms 2 and the remaining families were

126 

similarly represented across soil types. 

These patterns of restriction and exclusion indicate that there is an effect 

of soil type on floristics at Grasshopper Mountain. In Oregon and California 

the Ranunculaceae, Rosaceae, Fabaceae, Primulaceae and Scrophulariaceae 

are generally absent from ultramafic soils (Kruckeberg 1992), while the 

Caryophyllaceae have a particular affinity for ultramafics in Newfoundland 

(Dearden 1979) and Sweden (Rune and Westerbergh 1992). These patterns 

may be due to a combination of direct effects of soil chemical and physical 

properties on plant species and indirect effects through species interactions. 

Rare Taxa 

Ten rare vascular plant taxa, eight of which are provincially red- or bluelisted, 

were found at Grasshopper Mountain, eight from the ultramafic side 

and two from open, rocky cliffs on the non-ultramafic side (Table 3). The 

serpentine subspecies Adiantum pedatum subsp. calderi (maidenhair fern, Figure 

12), is included in this list though it has not received red- or blue-listed 

status. Aspidotis densa is included because it is reported by Douglas et al. 

(1998-2002) to be restricted to ultramafic outcrops east of the Coast-Cascade 

ranges. The Tulameen ultramafic complex is the only known site in 

British Columbia for Polystichum scopulinum (Douglas et al. 1998); however, 

no species are currently recognized as being endemic to serpentine sites in 

British Columbia. 

The distribution patterns of rare taxa on Grasshopper Mountain suggest 

that, in addition to their contribution to local and regional diversity, ultramafics 

may also be important to the maintenance of rare taxa in the province. 

Similarly, California ultramafics provide habitat for some of the last remnant 

patches of native California grasslands and their highly endangered flora 

which, on non-ultramafic soils, have been almost entirely replaced by 

Mediterranean grass species (Harrison 1999). At Grasshopper Mountain 

the rare ferns Polystichum scopulinum, P. kruckebergii (Figure 13, back cover), 

Aspidotis densa and Adiantum aleuticum (A. pedatum subsp. calderi) were observed 

only on ultramafic substrates, while Cheilanthes gracillima (Figure 14) was observed 

only on non-ultramafic rock outcrops. Similar substrate relationships 

for these species were observed by Kruckeberg (1964) at many sites in 

Washington state. The mechanisms maintaining rare taxa at Grasshopper 

Mountain require further study but may be related to the presence of open


127 

habitats within a forested matrix since most of the rare species were found 

in open areas. Harrison (1999) has investigated two hypotheses on California 

serpentines that may also apply at Grasshopper Mountain: 1. that there is 

edaphic control of competitive dominance and 2. that there is edaphic resistance 

to invasion of non-native species. 

Species Ranges 

The location of Grasshopper Mountain within the coast-interior climatic 

transition zone results in a mix of floristic elements with phytogeographical 

affinities to the coast, the interior, and the south; consequently, many species 

occur at the edge of their ranges (Tables 4 through 6). Several of these are 

species of interior BC occurring at the western edge of their ranges on the 

dry, ultramafic sites of Grasshopper Mountain. The presence of a few taxa, 

including Pseudoroegneria spicata subsp. inermis and Eriogonum ovalifolium var. 

nivale, may represent westward range extensions (Douglas et al. 1998-2002). 

Two conifers, Pinus albicaulis and Pinus ponderosa, occur at the lower and upper 

limits of their ranges, respectively. 

Plant Species as Soil Indicators 

Following the indicator classification scheme proposed by Kruckeberg 

(1979, 1992), the plant species of Grasshopper Mountain were grouped into 

local ultramafic indicator species, local ultramafic excluded species, and 

widespread (bodenvag) species (Tables 4 through 6). Within each indicator 

group, taxa were further subdivided into functional groups based on site 

moisture affinity (see Douglas et al. 1998-2002), and plant life forms that 

have previously been shown to respond to ultramafic soil conditions 

(Whittaker 1954; Kruckeberg 1979, 1992). Species occurring in less than 

10% of plots on a given substrate, and species found primarily on tillinfluenced 

soils, have been omitted from this classification. 

Thirty-five species are good indicators of local ultramafic conditions at 

Grasshopper Mountain (Table 4). However, the majority of these species 

are known to occur elsewhere on non-ultramafic soils. Their association 

with ultramafic soils at the study site may be a function of exclusion from 

the mostly well-developed mesic forests of the non-ultramafic side. For 

instance, 24 of the 35 indicator species are dry habitat associated herbs, and 

a number of species from other functional groups in this category are also

128 

associated with dry habitats. Similarly, Kruckeberg (1979, 1992) reported 

higher richness of dry habitat associated species for ultramafic sites in British 

Columbia and Washington as compared with the surrounding floras. 

Thirty-seven species are entirely or nearly excluded from ultramafic soils 

at the study site (Table 5). The main functional groups are deciduous broadleaved 

trees and shrubs (18 species) and mesic to moist habitat associated 

herbs (11 species). The exclusion of broad-leaved trees and shrubs from 

ultramafic soils has been previously documented (Whittaker 1954). These 

functional groups may be restricted to non-ultramafic substrates partly 

because of soil chemistry, and partly because the greater canopy cover, shade 

and soil moisture conditions better meet the habitat requirements of the 

particular species. The exclusion of dry habitat-associated herbs (six species) 

and the fern Cheilanthes gracillima, from ultramafic substrates is an interesting 

pattern since ample habitat appeared to be available on the ultramafic side 

of the mountain. These species may be directly excluded by ultramafic soil 

factors. 

Thirty-four species representing all functional groups except ferns were 

widespread on all three soil types (Table 6). These bodenvag species may be 

exhibiting one of two responses to ultramafic soils: 1. they may be indifferent 

to the adverse chemical and physical soil environment or 2. those individuals 

occurring on ultramafic soils may represent edaphic races or ecotypes tolerant 

of soil conditions. Evidence for ecotypic differentiation has been shown 

for bodenvag species from California (Kruckeberg 1951; Rajakaruna and Bohm 

1999) and the Pacific Northwest (Kruckeberg 1967). For example, 

Kruckeberg (1967) found strong ecotypic response in Achillea millefolium and 

Potentilla glandulosa, partial ecotypic response in Antennaria racemosa, Juniperus 

communis, Pinus contorta, Pseudoroegneria spicata and Taxus brevifolia. These species, 

therefore, may exist as ecotypic races on the different soil types. He found 

no ecotypic response in Rubus parviflorus which may simply be indifferent to 

ultramafic soil conditions. 

We conclude that the ultramafic soils of Grasshopper Mountain exert an 

influence on plant growth which is similar to the vegetational response in 

Oregon and California. While richness and diversity levels are similar across 

substrates at Grasshopper Mountain, the floristics and ecological relationships 

are distinct. The presence of ultramafic soils within a matrix of nonultramafic 

soils is important for increasing local and regional diversity and


129 

Figure 1. Simplified diagram of photosynthetic apparatus. PS I and II are photosystem I 

and II, respectively; OEC is the oxygen-evolving complex; P i 

is inorganic phosphate; e - 

represents electrons; H + represents protons; CH 2 

O represents carbohydrate products of 

photosynthesis. 

Photos: Rob Guy 

Figure 2. Colour changes later in younger leaves. 

Figure 3. Isolated tissues turn before areas adjacent to major veins in this Acer rubrum 

(red maple) leaf.

130 


Figure 4. a) anthocyanin in autumn foliage of Acer palmatum (Japanese maple), b) 

astaxanthin in Chlamydomonas nivalis (red snow algae), c) rhodoxanthin in western 

Thuja plicata (western red cedar), d) betacyanin in salt-adapted Salicornia europaea 

subsp. rubra (samphire). 


Figure 5. Carotenoids in autumn Populus tremuloides (aspen) foliage. 

Figure 6. Autumn foliage of Larix lyallii (sub-alpine larch).


131 

Photo: Rob Guy 

Figure 7. The name of the first author’s eldest son, Lachlan, “stencilled” onto Euonymus 

alatus (winged Euonymus). 

Figure 8. Ultramafic occurrences in British Columbia, Canada showing the location of the 

Tulameen ultramafic complex. Adapted from Hulbert (2000-2001).

132 

Photo: Gary Lewis 

Figure 9. Canopy cover types at Grasshopper Mountain: a) closed forest over tillinfluenced 

soil 


Figure 10. b) moderately-closed forest over non-ultramafic soil.


133 


Figure 11. c) open forest over ultramafic soil and talus 


Figure 12. Adiantum pedatum subsp. calderi, recognized by Cody and Britton (1989) as 

the serpentine subspecies of maidenhair fern, is not recognized by Douglas et al. (1998- 

2002) who include it in the taxon Adiantum aleuticum.

134 


Figure 13. Polystichum kruckebergii, Kruckeberg’s holly fern, growing in gravel on Olivine 

Mountain. 


Figure 14. Cheilanthes gracillima growing on the non-ultramafic side of Grasshopper 

Mountain.


135 

Photo: Hugh Daubeny 

Figure 15. Rubus strigosus resistant to cane spur blight compared to a red raspberry 

cultivar susceptible to the disease 


Figure 16. Rubus strigosus plant with resistance to Phytophthora-incited root rot 

compared to plant of susceptible red raspberry cultivar.

136 


Figure 17. Fruit of Rubus strigosus compared to fruit of 4th generation derivative. 

Photo: Hugh Daubney 

Figure 18. Fruit of ‘Tulameen’, a leading fresh market cultivar.


137 

for the maintenance of rare taxa. Decisions related to conservation and 

management of ultramafic sites in British Columbia, as in other regions, 

should take into account their potential biological significance. 

Acknowledgements 

We would like to thank Jocie Ingram for her excellent field assistance, Les 

Lavkulich for providing facilities for the soil chemical analyses at UBC, and 

Fred Ganders for help with plant identification. Thank you to Rose 

Klinkenberg, Jack Maze, Terry McIntosh and Randall Rae for reviewing earlier 

versions of this manuscript. Financial assistance was provided by a Natural 

Sciences and Engineering Research Council of Canada (NSERC) scholarship 

and a University Graduate Fellowship (both to GJL). Additional funding 

was provided through the Studies on BC Ecosystems Fund at UBC. 

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140 

Mean and Standard Errors for Selected Soil Variables for Ultramafic, 

Till-influenced and Non-Ultramafic Plots 

Soil Variables 

Soil Type 

Ultramafic 

n = 26 

Till 

n = 10 

Non- 

Ultramafic 

n = 35 

Ca meq/100g 

4.045 

+/- 0.415 

a 

4.633 

+/- 0.962 

a 

10.126 

+/- 0.926 

b 

Mg meq/100g 

8.204 

+/- 0.656 

a 

3.502 

+/- 1.012 

b 

1.102 

+/- 0.099 

c 

Ca:Mg 

0.502 

+/- 0.031 

a 

1.887 

+/- 0.432 

a 

9.583 

+/- 0.395 

b 

Ni ppm in soil 

20.819 

+/- 2.622 

a 

2.585 

+/- 0.716 

b 

0.569 

+/- 0.097 

b 

Table 1: Means and standard errors for selected soil variables for ultramafic, tillinfluenced 

and non-ultramafic plots. Shared letters denote a non-significant difference 

(p > 0.05) based on ANOVA results and the post-hoc Bonferroni adjustment. 

Rare Taxa Found on Grasshopper Mountain Including Their 

Provincial Ranking and the Soil Type on Which They Occurred 

Species 

Adiantum aleuticum 

(A. pedatum subsp. calderi) 

Aspidotis densa 

Arabis holboellii var. pinetorum 

Cheilanthes gracillima 

Crepis atrabarba subsp. atrabarba 

Lupinus arbustus subsp. pseudoparviflorus 

Melica bulbosa var. bulbosa 

Polemonium elegans 

Polystichum kruckebergii 

Polystichum scopulinum 

Ranking 

— 

— 

blue 

blue 

red 

red 

blue 

blue 

blue 

red 

Soil Type 

Ultramafic 

Ultramafic 

Ultramafic 

Non-Ultramafic 

Primarily Ultramafic 

Ultramafic 

Ultramafic 

Non-Ultramafic 

Ultramafic 

Ultramafic 

Table 2: Rare taxa found on Grasshopper Mountain including their provincial ranking and 

the soil type on which they occurred. Rare vascular plant taxa have been defined 

through the work of the BC Conservation Data Centre and are summarized in Douglas et 

al. (1998). Red-listed species are taxa considered “candidates for legal designation as 

endangered or threatened species.” Blue-listed species are “vulnerable rare taxa that 

could become candidates for the Red List in the foreseeable future.”



Total Number of Species per Family for Ultramafic, Till and Non- 

Ultramafic Plots 

Family 

Ultramafic 

Till 

Non- 

Ultramafic 

Total 

Apiaceae 

5 

1 

2 

5 

Asteraceae 

11 

5 

6 

14 

Betulaceae 

0 

2 

3 

3 

Caprifoliaceae 

0 

1 

2 

2 

Caryophyllaceae 

3 

1 

1 

3 

Ericaceae 

2 

2 

1 

3 

Fabaceae 

2 

1 

2 

2 

Grossulariaceae 

0 

1 

2 

2 

Gymnosperms 

8 

5 

7 

10 

Hydrophyllaceae 

1 

0 

1 

1 

Juncaceae 

1 

0 

0 

1 

Liliaceae 

4 

5 

7 

7 

Onagraceae 

2 

0 

1 

2 

Orchidaceae 

2 

2 

2 

2 

Poaceae 

7 

2 

4 

8 

Polygonaceae 

4 

0 

0 

4 

Pteridophytes 

4 

1 

1 

4 

Pyrolaceae 

1 

1 

1 

2 

Ranunculaceae 

1 

0 

3 

3 

Rosaceae 

5 

5 

11 

12 

Salicaceae 

0 

0 

3 

3 

Saxifragaceae 

1 

0 

1 

2 

Scrophulariaceae 

3 

1 

4 

5 

Other 

5 

5 

9 

9 

Table 3: Total number of species per family for ultramafic, till, and non-ultramafic plots, 

and total across all three soil types. The gymnosperms, pteridophytes, and “other” are 

represented by three, two and 13 families, respectively.


Local Ultramafic Indicator Species: Species Restricted to or Found 

Primarily on Ultramafic Soils at Grasshopper Mountain 

Functional Group 

Deciduous broad-leaved shrubs and trees 

Evergreen coniferous shrubs and trees 

Evergreen broad-leaved shrubs 

Mesic to moist habitat-associated herbs 

Dry habitat-associated herbs 

Ferns 

Total Number of Indicator Species 

Species 

Rosa nutkana 

Juniperus communis var. montana 

Pinus albicaulis * 

Taxus brevifolia † 

Arctostaphylos uva-ursi 

Cirsium edule ‡ 

Lupinus arcticus subsp. subalpinus 

Achillea millefolium var. lanulosa 

Antennaria racemosa ‡ 

Arenaria capillaris subsp. americana ‡ 

Astragalus miser var. serotinus ‡ 

Bromus carinatus 

Castilleja hispida var. hispida (yellow form) 

Cirsium hookerianum ‡ 

Crepis atrabarba subsp. atrabarba ‡ 

Epilobium minutum 

Eriogonum heracleoides var. angustifolium ‡ 

Eriogonum ovalifolium var. nivale ‡ 

Eriogonum umbellatum ‡ 

Koeleria macrantha ‡ 

Lomatium ambiguum ‡ 

Lomatium macrocarpum‡ 

Luzula multiflora subsp. multiflora 

Melica bulbosa var. bulbosa ‡ 

Melica subulata 

Phacelia hastata var. hastata ‡ 

Pseudoroegneria spicata subsp. inermis ‡ 

Senecio canus ‡ 

Senecio integerrimus var. exaltatus ‡ 

Senecio streptanthifolius ‡ 

Silene parryi 

Adiantum aleuticum (A. pedatum subsp. calderi) 

Aspidotis densa 

Polystichum kruckebergii 

Polystichum scopulinum 

35 

Table 4: Local ultramafic indicator species: species restricted to or found primarily on 

ultramafic soils at Grasshopper Mountain. ‡ denotes an interior species occurring at the 

western edge of its range. † denotes a coastal species occurring at the eastern edge of 

its range. * denotes a species occurring at the lower limits of its altitudinal range.



Local Ultramafic Excluded Species: Species Restricted to or Found 

Primarily on Non-Ultramafic Soils at Grasshopper Mountain 






Ferns 

Total Number of Excluded Species 

Species 

Acer glabrum var. douglasii 

Alnus viridis subsp. sinuata 

Betula papyrifera var. papyrifera 

Holodiscus discolor 

Lonicera involucrata 

Lonicera utahensis 

Philadelphus lewisii 

Populus tremuloides 

Prunus virginiana 

Ribes lacustre 

Ribes viscosissimum ‡ 

Rosa gymnocarpa 

Salix spp. - approximately 3 species 

Shepherdia canadensis ‡ 

Spiraea betulifolia subsp. lucida ‡ 

Symphoricarpos albus 

Ceanothus velutinus var. velutinus ‡ 

Penstemon fruticosus ‡ 

Actaea rubra 

Arnica cordifolia ‡ 

Aster conspicuus ‡ 

Clintonia uniflora 

Fragaria vesca var. americana 

Fragaria virginiana var. platypetala 

Orthilia secunda var. secunda 

Pedicularis bracteosa var. latifolia 

Prosartes hookeri var. oregana 

Thalictrum occidentale 

Valeriana sitchensis 

Allium cernuum var. cernuum 

Antennaria rosea 

Arabis exilis ‡ 

Arabis holboellii 

Artemisia michauxiana ‡ 

Heuchera cylindrica ‡ 

Cheilanthes gracillima + 

37 

Table 5: Local ultramafic excluded species: species restricted to or found primarily on 

non-ultramafic soils at Grasshopper Mountain. ‡ denotes an interior species occurring at 

the western edge of its range. + denotes a southerly species occurring at the northern 

edge of its range.


Widespread (Bodenvag) Species: Species Found Commonly on All 

Soil Types at Grasshopper Mountain 



Evergreen coniferous shrubs and trees 




Dry to moist habitat associated herbs 

Total Number of Bodenvag Species 

Species 

Amelanchier alnifolia 

Prunus emarginata 

Rubus parviflorus 

Vaccinium membranaceum 

Abies lasiocarpa var. lasiocarpa ‡ 

Pinus contorta var. latifolia 

Pinus monticola 

Pinus ponderosa ‡# 

Picea engelmannii ‡ 

Pseudostuga menziesii var. (study site on 

border of varietal ranges) ‡† 

Mahonia aquifolium 

Pachistima myrsinites 

Angelica arguta 

Aquilegia formosa subsp. formosa 

Aster engelmannii ‡ 

Bromus vulgaris 

Epilobium angustifolium subsp. angustifolium 

Erythronium grandiflorum subsp. grandiflorum 

Goodyera oblongifolia 

Lilium columbianum 

Maianthemum racemosum subsp. amplexicaule 

Osmorhiza sp. 

Viola glabella 

Agoseris aurantiaca subsp. aurantiaca 

Calamagrostis rubescens ‡ 

Carex rossii 

Fritillaria affinis var. affinis 

Hieracium scouleri var. griseum ‡ 

Lomatium dissectum var. multifidum ‡ 

Pedicularis racemosa 

Piperia unalascensis 

Sedum lanceolatum var. lanceolatum 

Castilleja miniata (orange form) 

Moehringia macrophylla + 

34 

Table 6: Widespread (bodenvag) species: species found commonly on all soil types at 

Grasshopper Mountain. ‡ denotes an interior species occurring at the western edge of its 

range. † denotes a coastal species occurring at the eastern edge of its range. + denotes 

a southerly species occurring at the northern edge of its range. # denotes a species 

occurring at the upper limits of its altitudinal range.



The North American Red Raspberry - A Genetic 

Resource Awaiting Exploitation 

Abstract 

Rubus idaeus L., the red raspberry consists of two subspecies, R. idaeus subsp. strigosus 

Michx, which is native to North America and far eastern Asia, and R. idaeus subsp. 

vulgatus Arrhen, which is native to Europe and Asia Minor. The subspecies are usually 

referred to as R. strigosus and R. idaeus, respectively. Neither is adequately represented 

in the red raspberry breeding gene pool. This paper describes efforts to find potentially 

useful parents with disease or insect resistances from previously unexploited selections of 

R. strigosus. Selections with resistances to aphid, root rots, and cane diseases were 

identified and have been incorporated into the Pacific Agriculture Research Centre’s 

breeding programme. Several derivatives of the selections have the potential to become 

cultivars with entirely new sources of resistances. 

Red raspberries (Rubus idaeus) are native to temperate regions of Europe, 

Asia, and North America (Jennings, 1988). Those occurring throughout 

North America and into Eastern Asia are usually designated Rubus idaeus 

subsp. strigosus and those occurring in Europe, from near the polar circle to 

the mountains of the Caucasus in Asia Minor, designated R. idaeus subsp. 

vulgatus (Jennings, 1988). Although the two subspecies are completely crossfertile, 

they are often given specific rank as R. strigosus and R. idaeus, 

respectively. I will follow the usual custom of breeders and other researchers 

and use the species designations (Jennings, 1988; Daubeny, 1996). R. idaeus 

is distinguished by glandless inflorescences and thimble shaped-fruit and R. 

strigosus by glandular inflorescences and round fruit. Both R. idaeus and R. 

strigosus grow in a wide range of environments. 

Several years ago I came to appreciate the extent of the range for R. strigosus 

in North America. In August, I collected fruit from plants growing in swampy 

woods bordering White Lake, approximately 110 km west of Ottawa, and 

then in October I collected fruit from plants growing on a dry, desert-like 

bank bordering a highway over a mountain pass in southern Utah. During 

the same period, I received seed from fruit collected in geographically diverse 

regions throughout Canada and the United States. The casual traveler can 

Hugh M. Daubeny, Emeritus Research Scientist. 

Pacific Agriculture Research Centre, Agassiz, BC, Canada. 

Correspondence address: 3558 W 15th Avenue, Vancouver, BC, V6R 2Z4.


also appreciate the range of environments. Traveling by train from Montreal 

to Ottawa, the gravely banks on either side of the tracks appear to be 

landscaped by it. Plants grow in the chinks of the old log structures in 

British Columbia’s historic Barkerville in the centre of the Province and 

along the road side going up to the glorious alpine meadows of Manning 

Park in the southern part of BC. At least one popular Canadian film used, 

probably inadvertently, R. strigosus as part of the scenery. In the National 

Film Board’s “Margaret’s Museum”, a vigorous plant figured prominently 

beside the house, later to become the Museum that Margaret and her husband 

built on a wind swept, desolate-appearing Cape Breton peninsula above the 

pounding Atlantic surf. The only problem with this - the plant remained the 

same size as the years passed. 

Until recently, most raspberry cultivars were derived from the crosses 

between selected plants of R. strigosus and R. idaeus that were made in the 

19th and early part of the 20th centuries (Daubeny, 1996). Considering the 

very wide distribution of R. strigosus, it is amazing that only nine individual 

selections of the species appear to be involved in the derivations of these 

cultivars (Dale et al., 1993; Daubeny, 2002; Finn and Knight, 2002). As far 

as can be determined, these selections were obtained from seedling 

populations growing in or near the following: New York City; Oswego County, 

New York; New Haven, Connecticut; Ottawa, Ontario; Ohio; Illinois; New 

Jersey; North Carolina, and Wyoming. The records on the origins of some 

of these are incomplete or vague, but it is obvious that the nine selections 

represent only an infinitesimal part of the genetic variability within the species. 

There are two compelling reasons why this variability must be exploited. 

First, pest resistance, in the broadest sense, is urgently required since key 

chemicals for control have been or soon will be eliminated. Resistance has 

both environmental and economic connotations, and many, if not all of the 

necessary resistances, are available in the wild species. 

Second, red raspberries are increasingly being grown in environments 

considered to be less than ideal for the currently available cultivars. For 

example, production is increasing in harsh continental-type climates, such as 

the Canadian Prairies, and, also in mild Mediterranean-type climates, such as 

the Santiago area of Chile, Southeastern Australia, and Central Coastal and 

Southern California. Continental-type climates need cultivars with winter 

hardiness and Mediterranean-type climates need cultivars with the minimal



chilling requirements necessary to break dormancy. Even in climates that 

are optimal for present production, such as the Pacific Northwest, more 

plantings are to be found on heavier, poorly drained soils where Phytophthorainfected 

root rots can be devastating. The wide range of habitats, in which 

plants of the wild species flourish, indicates the presence of variability that 

can eventually meet the requirements of many of the diverse environments 

in which red raspberry culture now takes place or will take place in the future. 

Concomitant with the urgent need for more pest and disease resistances 

and wider environmental adaptations, unique market outlets for red raspberry 

fruit are being developed (Finn and Knight, 2002). There are few fruits that 

give rise to as many popular processed products. For example, raspberry 

juice by itself or in blends with other fruit such as cranberry, has become 

popular. Raspberry jam has always been popular and fruits processed into 

ice cream and yogurt and as whole “individual quick freeze” are equally 

popular. Consumers demand that fresh red raspberries be available for 12 

months of the year, just as strawberries and apples are. This has stimulated 

production in the Southern Hemisphere, where fresh fruit is air-freighted to 

the Northern Hemisphere from November through to April. Greenhouse 

and tunnel production for “out-of-season” fruit has also increased (Pritts et 

al., 1999). Recently there has been increased recognition of the health value 

of red raspberries, which have high levels of antioxidants or “nutraceutical” 

compounds such as anthocyanins, phenolics and ellagic acid, each of which 

is reported to be effective against the effects of the degenerative diseases of 

aging (Finn and Knight, 2002). Red raspberries figure prominently in 

advertisements for cereals and other products that are equated with healthy 

diets. 

In recent years, the Agriculture and Agri-Food Canada red raspberry 

breeding programme at the Pacific Agriculture Research Centre (PARC) has 

been using selections from previously unexploited populations of R. strigosus 

(Daubeny, 2002). Seedling populations of the species were obtained from 

sites in BC, the Yukon, Ontario, Quebec, Nova Scotia, Washington, 

Minnesota, Wisconsin, New York, and North Carolina. Most seedlings were 

initially screened for reaction to the aphid, Amphorophora agathonica, a vector 

of the raspberry mosaic virus complex. This complex can have a devastating 

effect on growth and fruiting of susceptible cultivars and the easiest way to 

control it is through vector resistance. Genes giving resistance to the aphid


were identified in selected seedlings obtained from one site in Quebec and 

two in Ontario (Daubeny and Stary, 1982). They have been incorporated 

into breeding programme selections and appear to give resistance to a 

resistance-breaking biotype of the aphid. Until recently, resistance was based 

entirely on a single gene obtained many years ago from R. idaeus via the old 

cultivar ‘Lloyd George’. It is not surprising that resistance from this source 

would ultimately break down after many years of selection pressure. 

Some R. strigosus seedlings show resistance or reduced susceptibility to 

Phytophthora-incited root rots and cane diseases, including spur blight (Didymella 

applanata) (Figure 15), cane botrytis (Botrytis cinerea), and cane spot (Elsinoe 

veneta), as well as fruit rot (mostly caused by Botrytis cinerea), black vine weevil 

(Otiorhynchus sulcatus), and root lesion nematode (Pratylenchus penetrans) (Dale 

et al., 1989; Daubeny, 1996; Vrain and Daubeny, 1986). Other valuable traits 

observed include winter hardiness, self-supporting habit, early ripening fruit 

on floricanes (summer) and/or primocanes (autumn), and bright nondarkening 

red fruit that is easy to remove from the receptacle. The last is 

important to improve harvesting efficiency whether it be by hand or machine 

and also to reduce risk of fruit rot. 

Root rot resistance is particularly important. The PARC breeding 

programme has identified resistance in seedling populations of R. strigosus 

from six sites, one each in BC, New York, Minnesota, North Carolina, Ontario, 

and Quebec (Lévesque and Daubeny, 1999; Daubeny, unpublished data) (Figure 

16). There are some selections, 2 to 5 generations removed from these, 

that show at least some resistance and have acceptable fruit size and coherence 

(invariably R. strigosus selections produce small, crumbly fruit [Figure 

17]). Several of the selections appear to have cultivar potential and are 

being extensively evaluated under commercial conditions. 

It is obvious that some progress has been made in broadening the base of 

R. strigosus germplasm available to modern day red raspberry breeding. 

However, efforts have been curtailed in recent years because public sector 

resources for collecting and exploiting wild population germplasm have been 

reduced or even eliminated. Concurrently the private sector has not shown 

much interest in acquiring representative selections from these populations 

because it usually takes a long time to incorporate useful traits from the wild 

into commercially acceptable cultivars. At the same time the wild populations 

are endangered by humanity’s myriad activities with many potentially useful



traits being lost. It is imperative that this situation be remedied and that 

there be a thorough and systematic examination of representative selections 

from the remaining populations. The use of DNA genetic markers to 

determine variability and relationship to important economic traits will 

certainly be of value in this task (Daubeny, 2002; Finn and Knight, 2002). 

While conservation of R. strigosus populations is important, there is recognition 

that the species, along with two other Rubus species, R. spectablis Pursh. 

and R. parviflorus Nutt., can be aggressive invaders of areas disturbed by 

logging, burning, and site preparation activities and can impede reforestation 

efforts by competing for nutrients, moisture, and light (Oleskevich et 

al., 1996). It is obvious that there must be an appropriate balance between 

conservation and eradication. It seems that it would be a relatively simple 

matter to take representative plants from sites on which there is proposed 

eradication. 

The genetic base for modern day red raspberry breeding is being expanded 

by the introduction of genes from the native North American red 

raspberry. Efforts are also being made to introduce new genes from the 

European red raspberry, the original genetic base, which is probably as narrow 

as that of its North American counterpart (Jennings et al, 1991). 

Already there has been some success with the introduction of genes from 

related Rubus species into red raspberry breeding programmes (Daubeny, 

2001). For example, the successful cultivar ‘Tulameen’ (Figure 18), released 

from the PARC breeding programme in 1989, has genes from the black 

raspberry, R. occidentalis L. (Daubeny and Kempler, 2003). These genes were 

introduced into the red raspberry gene pool by the breeding programme at 

the East Malling Research Station (now Horticulture Research International) 

in the United Kingdom. It has taken four to six generations to recover 

acceptable red raspberry qualities combined with black raspberry traits, such 

as fruit firmness, extended shelf life and late ripening. In raspberry breeding, 

a generation may be as long as seven to eight years, which means that the 

original interspecific crosses were made in the 1950s. Other related species, 

now appearing in the derivations of new cultivars, include R. spectabilis (Pacific 

Coast salmonberry), R. arcticus L. (Arctic raspberry), R. odoratus L. (Eastern 

North American purple flowering raspberry), and the Asiatic species, R. 

coraneus Mig., R. cockburianus Hemsl., R. crataegifolius Bge. and R. phoenicolasius 

Maxim. Each species has genes for useful plant traits, some of which are

150 

similar to the traits being introduced from the native R. strigosus and R. idaeus. 

Rubus spectabilis is of special interest as a source of early ripening on both 

floricanes and on primocanes. These traits are important for regional selfsufficiency 

in red raspberry production both for the fresh market and for 

processing. 

Clearly, various Rubus species offer enormous potential for improvement 

of the red raspberry. Wild species biodiversity must be maintained both in 

natural habitats and in germplasm repositories. In present day breeding 

programmes most of the species, except for R. strigosus and R. idaeus, are 

represented by only one genotype. Continued development of a range of 

adapted cultivars thus requires use of many more genotypes from all sources 

(Dale et al., 1993). Subsequently there must be sustained and effective efforts 

to evaluate representative genotypes for potentially useful traits. The 

genotypes identified can then be made available on an international basis for 

incorporation into all red raspberry breeding programmes. 

References 

Dale A., Moore, P.P., McNicol, R.J., Sjulin, T.M., and Burmistrov, L.A. 1993. 

Genetic diversity of red raspberries throughout the world. Journal of the 

American Society for Horticultural Science 118:119-129. 

Daubeny, H.A. 1996. Brambles. In Fruit Breeding Volume II. Vines and Small 

Fruits. Edited by J. Janick and J.N. Moore. John Wiley. pp109-190 

Daubeny, H.A. 2001. Raspberries revisited. Biodiversity 2:23 

Daubeny, H.A., 2002. Raspberry breeding in the 21st century. Acta 

Horticulturae 585: 69-72. 

Daubeny, H.A. and Stary, D. 1982. Identification of resistance to Amphorophora 

agathonica in the native North American red raspberry. Journal of the 

American Society for Horticultural Science 107:593-597. 

Daubeny, H.A. and Kempler, C. 2003. ‘Tulameen’ red raspberry. Journal 

American Pomological Society 57: 42-44. 

Finn C. and Knight, V.H. 2002. What’s going on in the world of Rubus 

Breeding Acta Horticulturae 585: 31-35. 

Jennings, D.L. 1988. Raspberries and Blackberries: Their Breeding, Diseases and 

Growth. Academic Press. London, UK. 

Jennings, D.L., Daubeny, H.A., and Moore, J.N. 1991. Blackberries and 

Raspberries (Rubus). In: Genetic Resources of Temperate Fruit and Nut


151 

Crops. Edited by J.N. Moore and J.R. Ballington. International Society for 

Horticultural Science, Wageningen, The Netherlands. pp331-389. 

Lévesque, C.A. and Daubeny, H.A. 1999. Variations in reaction to Phytophthora 

fragariae var. rubi in raspberry genotypes. Acta Horticulturae 505: 231-235. 

Olesevich C., Shamoun, S.F., and Punja, Z.K. 1996. The Biology of Canadian 

Weeds. 105. Rubus strigosus Michx., Rubus parviflorus Nutt., and Rubus 

spectablis Pursh. Canadian Journal of Plant Science 76:187-201. 

Pritts, M.P., Langhams, R.W., Whitlow, T.H., Kelly, M.J., and Roberts, A. 1999. 

Winter raspberry production in greenhouses. HortTechnology 9:13-15. 

Vrain, T.C. and Daubeny, H.A. 1986. Relative resistance of red raspberry and 

related genotypes to the root lesion nematode. HortScience 21:1435-1437.

152 

November in the Garden 

A November walk in the garden may be cool or wet, but there are many 

wonderful autumn days. There are trees, especially the maples, still showing 

their brilliant colours, and there are the flowers that grow well when the days 

are cooler. 

But when I walk through the November garden I have my eye open for 

plants that might look good in a wreath. There is Thuja plicata (western red 

cedar) and there are many evergreen shrubs, but what I really need are good 

sources of berries. Even if the berries are not usable in a wreath, they are all 

wonderful to look at. 

In the David C. Lam Asian Garden some of the Sorbus spp. (mountain 

ash) have already dropped some or all of their fruit, but others will keep 

them for another month or more. You really must walk along the lower path 

in the Asian Garden where you will surely find three different plants with 

blue fruit. At the first bridge is Decaisnea fargesii (dead man’s toes) with 

wonderful blue bean-like fruit. A little further on the right, below the path is 

a grouping of several Dichroa febrifuga with rich dark blue berries. Almost at 

the end of the path on your left you will find Alangium platanifolia var. 

macrophyllum with its wonderful blue drupaceous fruits scattered on the ground 

or still clinging to the branches. The contrast between the blue fruit and the 

golden leaves is almost magical. 

If purple is your colour, Callicarpa bodinieri is the shrub for you. The birds 

are attracted to the clusters of violet drupaceous fruit in early winter. Just 

before you enter the tunnel you will find Berberis morrisonensis with berries a 

lovely shade of red. At the top of the path from the tunnel, against the 

Garden Pavilion, is Pyracantha coccinea (fire thorn). The brilliant orange fruit 

of this member of the Rosaceae is a pome, like an apple. 

The only berry left in the Food Garden is the Actinidia deliciosa (kiwi fruit). 

These will be harvested soon and will ripen indoors. Beside it is Actinidia 

Judy Newton, Education Coordinator. 

UBC Botanical Garden and Centre for Plant Research, 

6804 SW Marine Drive, Vancouver, BC, Canada V6T 1Z4. 

judy.newton@ubc.ca


153 

arguta with much smaller, smooth-skinned fruit that usually ripens in October. 

You need to look right up into the canopy to spot the fruit. Before you 

leave, take a few minutes to look at all the winter vegetables that you can 

grow and harvest. 

In both the Winter Garden and the E.H. Lohbrunner Alpine Garden, you 

will find my favourite berries, Gaultheria mucronata (syn. Pernettya mucronata), 

an evergreen shrub from Chile. The plump berries are white, red, pink or 

mauve and will remain on the plants until late spring. 

The Sorbus hupehensis ‘Pink Pagoda’ in the bottom corner of the Winter 

Garden as you walk along the path below the Alpine Garden is the easy to 

see. The fruit, a small pome, starts pink and fades to white as the winter 

progresses. Then one day in spring the migrating robins find the tree and 

very soon the fruit is gone. Malus ‘Adirondack’ grows near the ‘Pink Pagoda’ 

and its wonderful red-orange fruit remain for most of the winter. Cotoneaster 

and many Ilex (holly) species, including the deciduous hollies Ilex verticillata 

and Ilex ‘J.C. van Toll’ provide further interest and it is hard to miss Arbutus 

unedo (strawberry tree) with its vivid red-orange fruit. You can get a closer 

look at this plant in the planting outside the Shop In The Garden. If you 

walk along the grass towards the stream you will see a delightful small tree 

with bright red fruit, Crataegus grignonensis. If you are lucky enough to see it 

capped with snow, you will have the perfect picture. 

As you approach the Garden exit, on the left side of the main path, you 

will see the red berries of the female Skimmia japonica. On the ground to the 

left, opposite the purple flowering, Aconite are bright red, cone-shaped seed 

heads of Arisaema consanguineum. 

There is an enormous variety of shrubs and trees in fruit, including 

Mahonia, Chaenomeles, Sarcococca and Stranvaesia, to name a few. But don’t leave 

without one last stop to see the many beautiful small pomes on Sorbus ‘Eastern 

Promise’ growing on the boulevard outside the main gate. It is hard to 

describe the exact colour… maybe orange with pink overtones. What do 

you think

154 

Gleanings 

Journal Articles 

Big old cottonwoods 

Stewart B. Rood and Mary Louise Polzin 

Canadian Journal of Botany 81: 764-767 

When we think of old trees, we look to the giants of Clayquot Sound, 

Vancouver Island or the oaks of Europe. This report of Populus trichocarpa 

that are up to 400 years old points to potential for substantial longevity of 

section Tacamahaca cottonwoods as opposed to the shorter-lived section 

Aigeiros species. The authors also point out that such long-lived trees 

contribute to habitat structure and reveal stable floodplain locations. 

Impacts of golf course construction and operation on headwater streams: 

bioassessment using benthic algae 

Jennifer G. Winter, Peter J. Dillon, Carolyn Peterson, Ron A. Reid and Keith 

M. Somers 

Canadian Journal of Botany 81: 848-858 

The authors show that golf course land management on the Canadian 

Precambrian Shield is associated with significant differences in the abundance 

of certain benthic algae in headwater streams. There was a lower proportion 

of diatoms and a high proportion of cyanobacteria and filamentous green 

algae in the operational golf course streams. 

Review Article 

Pharmaceutical discoveries based on ethnomedicinal plants: 1985-2000 and 

beyond 

Walter H. Lewis 

Economic Botany 57: 126-134 

A short review by a UBC alumnus coinciding with publication of the 2 nd 

edition of “Medical Botany”


155 

Volume 14 Index 

Authors, Titles, Illustrations & Key Words 

Abies chensiensis 86, 14:(3) front 

cover 

Abies delavayi 87 

Abies densa 88 

Abies fargesii 87 

Abies grandis 84, 89 

Abies koreana 86 

Abies lasiocarpa 84 

Abies pinsapo 85, 14:(3) back cover 

Abies sp., silver firs, UBC Botanical 

Garden 71 

Acer palmatum, leaf pigmentation 130 

Acer rubrum, colour change 129 

Achenes, Bidens cernua, Bidens 

amplissima 83 

Adiantum pedatum subsp. calderi 133 

Arctostaphylos manzanita 14:(1) front 

cover 

August in the Garden 105 

Autumn colours, leaf biochemistry 111 

Bidens amplissima 81 

Bidens amplissima, endemic status, 

range extension 63 

Bidens cernua 82 

Bidens cernua, taxonomy 63 

Bradfield, G.E. 121 

British Columbia 

serpentine, floristics, ecology 121 

strawberry 5, 12 

ultramafic regions, map 131 

Burns Bog, editorial 1 

Canopy cover, forest 132, 133 

Catling, P.M. 12 

Cheilanthes gracillima 134 

Chlamydomonas nivalis, pigmentation 

130 

Climatological data 27 

Conservation, Bidens spp., endemic 

status 63 

Cronk, Q. 3 

Daubeny, H. 5, 145 

Directors’ Note 3 

Disanthus cercidifolius 14:(4) front 

cover 

Ecology, serpentine, British Columbia 121 

Ecosystems, editorial 109 

Editorial 1, 29, 61, 109 

Endemic, Bidens amplissima, range 

extension 63 

Ethics, pesticides, plant introductions, 

organic gardening 79 

Euonymus alatus, leaf stencilling 131 

Exploration, Hydrangea 31 

Floristics, serpentine, British Columbia 

121 

Forest, canopy cover 132, 133 

Fragaria chiloensis 

accessions from BC 15 

British Columbia, taxonomy, variation, 

germplasm 12 

diversity 5 

illustrations 13, 14, 15 

Fragaria sp., distribution maps 16, 17 

Fragaria x ananassa nothosubsp. 

cuneifolia, taxonomy, variation, 

germplasm 12 

Fragaria x ananassa, origins, Fragaria 

chiloensis 5 

Ganders, F.R. 63 

Germplasm, Fragaria, variation 12 

Germplasm, Rubus sp., pest resistance 


Gleanings 107, 154 

Guy, R.D. 111 

Hinkley, D.J. 31 

Hydrangea 

anomola subsp. petiolaris 14:(2) back 

cover 

arborescens subsp. radiata 41 

aspera subsp. strigosa ‘Elegant Sound 

Pavilion’ 43

156 

Hydrangea (cont.) 

integrifolia 46 

involucrata 43 

scandens subsp. chinensis 44 

seemannii 47 

serrata ‘Kiosumi’ 45 

serrata ‘Iza No Hana’ 14:(2) front 

cover 

sikokiana 42 

systematics, exploration 31 

Hydrangea sp. (with an affinity to H. 

peruviana) 46 

Instructions to Authors 28 

January in the Garden 26 

Justice, D. 71 

Klinkenberg, B. 63 

Klinkenberg, R. 63 

Larix lyallii 130 

Leaf 

biochemistry, autumn colours 111 

colour change 129, 130, 131 

pigmentation, Acer, Thuja, Salicornia 

130 

stencilling, Euonymus alatus 131 

Lewis, G.J. 121 

Map, ultramafic regions, British Columbia 

131 

May in the Garden 59 

Newton, J. 26, 59, 105, 152 

November in the Garden 152 

Organic gardening, ethical issues 79 

Peer review and the Editor, editorial 61 

Pest, resistance, red raspberry, genetic 

resource 145 

Pesticides, ethical issues 79 

Photosynthesis, schematic diagram 110 

Pigmentation, Acer, Chlamydomonas, 

Thuja, Salicornia 130 

Plant introductions, ethical issues 79 

Polystichum kruckebergii 134, 14:(4) 

back cover 

Populus tremuloides 130 

Range extension, Bidens amplissima 63 

Raspberry, red, pest resistance, genetic 


Resistance, red raspberry, genetic 


Rubus 

pest resistance, genetic resource 145 

strigosus 135, 136 

‘Tulameen’ 136 

Salicornia europaea subsp. rubra, leaf 

pigmentation 130 

Serpentine 

British Columbia, map 131 

floristics, ecology, British Columbia 121 

Silver Firs, see Abies sp. 71 

Stewartia koreana 14:(1) back cover 

Strawberry 

cultivated origins 5 

hybrid, distribution, taxonomy, variation, 

germplasm 12 

Systematics, Hydrangea, exploration 31 

Taxonomy, 

Bidens amplissima, Bidens cernua, 

endemic status 63 

Fragaria spp. 5, 12 

Rubus spp. 145 

Taylor, I.E.P. 1, 29, 61, 79, 109 

Thuja plicata, leaf pigmentation 130 

Ultramafic regions, British Columbia, map 

131 

Vancouver Island Beggarticks, see Bidens 

amplissima

Table of Contents 

Editorial 

Taylor 

Autumn Colours – Nature’s Canvas is a Silk Parasol 

Guy and Krakowski 

A Floristic and Ecological Analysis at the Tulameen 

Ultramafic (serpentine) Complex, southern British Columbia, 

Canada 

Lewis and Bradfield 

The North American Red Raspberry - A Genetic Resource 

Awaiting Exploitation 

Daubeny 

November in the Garden 

Newton 

Gleanings 

Index to Volume 14 

meter 

109 

111 

121 


152 

154 

156

Printed in Canada 

A UBC Botanical Garden and Centre for Plant Research Publication

pdf wkg Dav.14-14 - Davidsonia

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