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Davidsonia Sep 1, 1979

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Array DAVIDSONIA
VOLUME 10 NUMBER 3
Fall 1979 Cover:
Sarracenia purpurea subsp. purpurea, Northern Common Pitcher-plant, occurs in northeastern British Columbia, x 0.37.
Amanita porphyria, Grey-brown Amanita,
x 0.66. This Amanita is present in summer to
late fall in coniferous forests, especially along
trails and in clearings.
VOLUME 10
NUMBER 3
Fall 1979
Davidsonia is published quarterly by The Botanical Garden of The University of British
Columbia, Vancouver, British Columbia V6T 1W5. Annual subscription, six dollars. Single
numbers, one dollar and fifty cents, except for special issues. All information concerning
subscriptions should be addressed to the Director of The Botanical Garden. Potential
contributors are invited to submit articles and/or illustrative material for review by the
Editorial Board.
A cknowledgements
The pen and ink illustrations are by Mrs. Lesley Bohm. The photographs on page 53 and
facing page 64 were taken by Mrs. Sylvia Taylor. The article on Lonicera was researched by
Mrs. Sylvia Taylor. Editorial and layout assistance was provided by Mrs. Sylvia Taylor and
Mrs. Pam Morgan Robin.
ISSN 0045-9739
Second Class Mail Registration Number 3313 Insectivorous Plants in British Columbia
SYLVIA TAYLOR
For centuries people have been fascinated by the unusual adaptations and lifestyles of the insectivorous
(insect-eating) plants, also often called carnivorous plants. Several species have been cultivated as curiosities for
over 300 years. Sundews were important in medicine and in mythology in 15th and 16th century Europe. The
American Indians and early explorers and settlers in North America thought that a plant as strange as the
Pitcher-plant (Sarracenia) must be a possible source of medicine and drugs. Carnivorous plants have excited the
imagination of many writers — the man-eating plants of many stories are probably based on a superficial
knowledge of the small, harmless Venus'-flytrap (Dionaea muscipula) or other species with an active trapping
mechanism, coupled with a very vivid imagination. Needless to say, there are no scientific facts to substantiate
"man-eating plants". Certainly, there have been reports of one of the tropical Pitcher-plants (Nepenthes
chelsonii) digesting giant rats — but even this is difficult to substantiate. The majority of insectivorous plants
trap and digest small insects such as flies, ants and mosquitoes, and occasionally small frogs and lizards are
caught by the larger pitcher plants.
There are at least 450 species (in 13 genera belonging to six families) of insectivorous plants distributed
throughout the world. Their basic characteristic is their unique method of survival — they all lure and trap
insects, and have an enzyme system to digest the prey. The plants then absorb nutrients from the digested insects.
Like all green plants, insectivorous ones can manufacture their own food by photosynthesis utilizing the raw
materials of sunlight, water and carbon dioxide. But they also live in nutrient-poor habitats and supplement
mineral deficiencies by "feeding" on insects, absorbing necessary nitrogen-containing compounds. A 1
There is some disagreement among scientists as to the importance of the insectivorous habit. The plants
usually live in nitrogen-poor soils, and scientists had always assumed that the habit had been evolved to enable
the plants to survive in such conditions, where most other plant species cannot grow. However, it is now known
that some species can survive quite well without an insect diet, even in nutrient-deficient habitats. It appears that
insects provide supplemental nutrients, such as nitrogen and phosphorus, and that these nutrients allow the
plants to grow faster and produce more flowers and seed than when the supplementary diet is absent. Charles
Darwin and his son Francis first showed convincingly that Sundews in cultivation were more vigorous when fed
artificially by supplying insects to the leaves than ones that were not fed (Darwin, 1876). More recent
experiments have shown that it is likely that most of the plant's energy and essential organic compounds result
from photosynthesis, and that the majority of minerals used are obtained from the prey. Unfortunately, these
suggestions have not yet been definitely proved, and more research is needed. Certainly, the ability to absorb
certain essential nutrients from insects enables the plants to survive in nutrient-poor environments, where there is
little competition because of the absence of other plants.
All the insectivorous plants trap insects on the outside surface of a modified leaf, and most then secrete
enzymes onto or into the trap to digest the insect. Some of the simple end products of digestion are absorbed into
the plant through the outer cell walls. A few species do not appear to secrete enzymes, digestion being accomplished by decomposition of the insect by bacteria present within the trap.
Insectivorous plants may be divided into three groups based on the method of catching the prey:—
a) Active trap — typified by the Venus'-flytrap (Dionaea muscipula), which has a fast-acting reflex movement
making it the most animal-like trap. The most common active traps are found in the Bladderworts, which
have small, elastic-walled bladders with fast-moving trap doors. The prey is swept into the "bladder in a
current of water produced when the walls of the bladder spring apart following the opening of the entrance.
b) Semi-active trap or Flypaper trap — utilizes movement as a supplement to an adhesive that captures the prey.
This type is found in the Sundews and Butterworts. Glands on the leaf surface secrete adhesive droplets,
which trap insects attracted by odor, color, or refraction of the drops. Tentacles on the leaf then gradually
move over the prey, thus enclosing it. 42
c) Passive trap or Pitfall — relies entirely on highly specialized modifications of the leaves to trap the victim.
The prey is enticed to enter the pitcher of Pitcher-plants such as Sarracenia and Nepenthes by color and odor,
is then trapped, drowned, and digested in the fluid at the base of the pitcher.
Insectivorous plants usually grow in special and geographically restricted habitats, such as sphagnum bogs.
Even one new ditch or one load of fill can change the drainage pattern sufficiently to dry up a small bog, thus
making it unsuitable as a habitat for bog-loving plants. The addition of fertilizer and the control of fires also
have adverse effects, the latter by allowing forest vegetation to gradually grow in the bogs. These factors are
combining to make insectivorous plants scarce, and possibly even endangered in some areas. In addition, large
quantities or whole populations are sometimes dug up by plant collectors or commercial dealers. Because of
these concerns, several insectivorous plant habitats in the United States have been protected, and plants growing
in the wild should all be treated as rare and threatened.
In British Columbia there are eleven species of insectivorous plants, representing four genera in three families.
Droseraceae (Sundew Family)
The family Droseraceae contains four genera, three of which contain only one species each and are of limited
distribution — Dionaea muscipula, Venus'-flytrap; Drosophyllum lusitanicum, Portuguese Dewy Pine; and
Aldrovanda vesiculosa, an aquatic plant. The fourth genus, Drosera or the Sundews, contains a large number of
species (estimates range from about 85 to 144 species), of which six occur in North America, and two in British
Columbia.
Sundews are found in every continent of the world, typically inhabiting damp heaths, bogs, swamps, and
other wet habitats with acid soil (preferably with a pH of 3.5-4.0). It is a particularly hardy genus, and these
plants are among the first ones to grow on fire-ravaged forest soils, or other newly disturbed habitats, such as
roadside ditches and abandoned logging trails. In general, they are among the smallest of the insectivorous
plants, many being only 2.5-5 cm in diameter and often completely hidden in the surrounding sphagnum moss
except at flowering time. One exception is Drosera regiae of South Africa, which has leaves 61 cm long, and
which is reputed to be capable of catching and digesting small animals.
The genus consists of low, stemless, herbaceous perennials with a slender rhizome that produces a small
number of fine fibrous roots each year. The leaves are tiny, flat and solid or thick and clublike, and form a
rosette on the surface of the ground. The leaves may be round, spoon-shaped, oblong, or filiform (narrow and
threadlike). The shape of the leaf is the most important characteristic for distinguishing the North American
species. There is an obvious petiole (leaf stalk), except when the leaf is filiform. The margins and upper surface
of the leaf are covered with many stalked glands or "tentacles" (up to 400 have been counted in Drosera
rotundifolia, and 200 may be the average number) that secrete a highly viscid, clear mucilage that glistens in the
sunlight. The glands tend to be short and stubby near the centre of the upper surface, becoming longer towards
the margin. In some species, or in some habitats, the glands are bright red, contrasting with the transparent
globules. All the species flower in early summer, producing a slender, leafless, upright flower stalk that is
15-50(-76) cm tall and bears 1-20 erect flowers on the upper part. The flowers are bisexual, regular, 2.5 cm or less
across, and open one at a time only in sunshine, and often for only one morning. There are 5 petals, which are
white to pink or deep red, depending on the species. There are 5 to many stamens, arranged in whorls of 5. There
are 3-5 styles, which are often divided. The fruit is a capsule, and the seeds are very tiny (almost dust-like). The
seed is mature in 6-8 weeks. Many species are capable of self-fertilization if the flower has not been cross-
fertilized by the time it closes.
Insects are attracted to the leaves because of the odor, color, or, perhaps, refractions of the droplets. The
glands secrete both an adhesive substance and digestive enzymes. The viscid secretion droplets accumulate on
each gland head and are held there until the gland is touched by the prey, when more secretion is released. When
an insect lands on the leaf it becomes stuck in the mucilaginous droplets, and its struggles to escape stimulate the
secretion of more sticky fluid until the insect may be completely covered. At the same time, the stalked glands
begin to bend inward over the insect, soon followed by curling of the longer outer "tentacles". In a matter of
time (maybe several hours) the insect is completely caught within a sticky web of "bars". Most insects die within
about 15 minutes of capture of suffocation caused by the mucilage blocking the respiratory holes or trachea
along the abdomen. The glands now secrete digestive enzymes, which decompose the insect. It has been shown
that Sundews contain the enzymes peroxidase, estarase, acid-phosphatase, and protease in the digestive fluid.
After about a week the "tentacles" unfurl, and the leaf is able to catch more prey. It has been shown that each
"tentacle" can repeat the bending and unbending process about three times. There are minute stalkless glands scattered over the upper leaf surface and on the stalks of the larger glands.
Their function is not known, but there are two theories — one is that they function in the absorption of the
digestive products, and the other is that they are concerned with the movement of the larger glands.
There are two theories concerning the movement of the stalked glands. The first proports that bending is
caused by uneven growth of the stalk (Shetler, 1974b) — an acceleration of growth begins near the base on the
outside of the stalk and moves upward towards the gland at the top. This means that the outside of the stalk
grows faster than the inside, so that it bends inwards. Unbending is caused by accelerated growth on the inside of
the stalk. The second theory states that bending results from a loss of turgor (rigidity) in groups of cells along the
side of the stalk closest to the stimulus, that is, on the inside of the stalk. In this theory, the stalkless glands are
thought to be responsible for the withdrawal of fluid from the cells, which causes the loss of turgor. Unbending
is caused by the cells regaining turgor (Heslop-Harrison, 1978).
The Sundews are restricted to catching only small or very weak prey — flies and even ants are capable of
escaping from the sticky fluid or from between the "tentacles". Even so, they may catch large numbers of
insects — an English naturalist once estimated that plants covering 1 to 2 hectares of land killed about 6,000,000
insects in one season (Savage and Savage, 1979).
Sorcerers and alchemists both held Sundews in high esteem, because of the "dew". Until relatively recently,
many ordinary people in Europe believed that a single plant brought into the house would cause "pernicious
fever", and that a person who searched for a Sundew on Midsummer's Eve and rubbed the leaves over his skin
would become indefatigable. An accidental discovery, however, was of no use! Sundews were particularly
effective when collected at midnight on Midsummer's Eve — but one had to walk backwards all the time to
avoid being followed by the Devil! Extracts and teas are still prescribed in France and Germany as an antispasmodic for tuberculosis, asthma, catarrh and whooping cough. The active ingredient has been found to be
plumbagin, which is present in the British Columbian species.
The genus name is derived from the Greek droseros, 'dewy' or 'moist with dew', because the drops of sticky
mucilage glisten like dewdrops in the sun.
"To return to our native bog-plants; the Sundews, all very much
alike, and all, therefore, to be treated of under Drosera rotundifolia, are pretty and interesting, though not brilliant. Evil little
things they are, with their carnivorous habit. One wonders what
crime the past lives of Drosera can have held, that now the race
should be compelled to dree so ominous and unpleasant a weird
of murder and fraud. When will the Sundews be free of the
burden, through some self-sacrificing individual plant who shall
starve to death rather than take life, and so redeem his race into
the happier paths of peace and virtue?"
(Farrer, 1908, p. 245)
Drosera anglica Hudson Great Sundew
The Great Sundew is circumboreal, being found in swamps and bogs in Europe, Japan and North America. In
British Columbia it can be found in suitable bogs and swamps throughout the Province, sometimes occurring in
dense colonies that cover many acres. This is the most common species at moderate to high elevations in British
Columbia.
There is considerable variation in the shape of the leaf blade from narrowly ovate to spatulate-oblong or
cuneate-obovate. The leaves are 10-30 mm long, (2-)3-5(-7) mm broad, gradually narrowing to the petiole, and
often standing erect above the moss. The petiole is glabrous or sparingly glandular-hairy, and 1.5-8 cm long. The
scape (flower stalk) appears in June to August and is 6-18 cm tall, bearing 2-7 whitish flowers in a racemose
inflorescence at the top. The calyx is (4-)5-6 mm long, connate for about one-third of its length, and the lobes are
ovate-oblong. The petals are oblong and slightly to considerably longer than the calyx. There are (4)5 stamens,
and (3)4-5 styles, which are bifid for two-thirds of their length. The seeds are 1-1.4 mm long, and the testa is
loose, longitudinally striated, and prolonged but not flattened at each end of the seed.
Drosera anglica is believed by some authorities to be a fertile hybrid derived from a cross between D. rotundifolia and another species (possibly D. linearis Goldie). Some of the forms with the broader spatulate to obovate
leaves have been shown to be sterile hybrids between D. anglica and D. rotundifolia.
The specific name anglica is derived from Anglia, meaning England.
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FIGURE 1. Drosera rotundifolia, Round-leaved Sundew,
as seen from above, X 4.8.
Drosera rotundifolia Linnaeus (Figure 1) Round-leaved Sundew
The Round-leaved Sundew is also a circumboreal species, being found in sphagnum bogs and swamps
throughout the Northern Hemisphere. It is present in suitable habitats throughout British Columbia, and may
also be found growing on old floating logs in stagnant lakes, or among the mosses in marshy ground around
pools. This is the predominant North American species, being found in most areas except the extreme south and
most of the Rocky Mountain and Great Plains regions.
The leaves are spreading, often hidden in the moss, and may be bright red if the plant is growing in full
sunlight. The rosette of leaves is 6-8 cm in diameter. The leaf blades are round to broadly obovate-cuneate,
(5-)6-12 mm long, and usually at least as broad. The petiole is stout and 2-9 cm long. The scape appears in June
to early September, and is up to 25 cm tall, bearing (l-)3-10(-20) whitish flowers. In some parts of the range the
flowers may be pink or even yellow, mauve or purple. The flowers are very similar to those of the Great Sundew,
except that there are usually only 3 styles and they are bifid nearly to the base. The seeds are about 1.5 mm long,
and the testa is loose, longitudinally striated, and prolonged and distinctly flattened at each end of the seed.
In winter the rosette of leaves withers, and winter buds, which withstand freezing, are formed on the
rhizome. Axillary buds, which also form on the rhizome, create secondary rosettes below the main one. As the
rhizome decays, these separate and become new plants.
The specific name rotundifolia means 'with round leaves'. A potion made from the leaves of D. rotundifolia
has been used to cure coughs, whooping cough, and respiratory diseases such as asthma. The fresh juice from
the leaves has also been used in various parts of the world as a cure for corns, calluses and warts.
Both native species are easy to grow in live sphagnum moss, preferably the small-growing type that tends to
form tufts. The moss should be kept constantly moist, but not sodden, and there should be high humidity. These
plants should receive about 40-50% sunlight all day, and the temperature should be between 18-30°C during the
growing season. All Sundews need a dormant period, with reduced light and a temperature of -4 to 4°C. Forcing
the end of dormancy is not successful as the plants usually rot and die. Propagation is by seeds, or vegetatively
from young and vigorous leaves or offshoots. Lentibulariaceae (Butterwort Family)
The family is usually divided into five genera containing about 500 species between them. Four of the genera
are small and fairly homogenous; the fifth, Utricularia or the Bladderworts, is very complex and variable, and
includes an estimated 150 to over 300 species. Two of the genera in this family, Pinguicula (Butterwort) and
Utricularia (Bladderwort), occur in British Columbia.
Pinguicula Butterwort
There are between 30 and 46 species of Butterwort, widely distributed throughout the Northern Hemisphere
and in the South American Andes. They are usually found in wet places, often with mosses, such as on wet soil
or among rocks along shores and streambanks; on hummocks in swamps; or in low, wet, sandy pinelands.
Most of the species are very similar in appearance. They are fibrous-rooted perennial herbs with a compact
basal rosette of leaves that lies flat on the ground. The leaves are oval or tongue-shaped, pale yellowish-green,
and the margins curl inwards to form a shallow bowl shape. The flowers are solitary at the ends of erect bractless
scapes, and are somewhat reminiscent of the flowers of the genus Viola. The calyx is 5-lobed, and the lobes are
partly connate to form an upper lip of 3 lobes and a lower lip of 2 lobes. The corolla is usually white or light to
dark violet, or occasionally bright yellow, bilabiate with an upper lip of 2 lobes and a lower lip of 3 lobes, and is
prolonged into a basal spur. The fruit is a 2-valved capsule.
The insectivorous adaptations are not as obvious in this genus as in other genera. The only unusual feature is
the greasy appearance of the leaves. The upper surface of the leaf is covered with large stalked glands that secrete
a film of colorless sticky mucilage over the leaf. There are also stalkless glands, raised slightly above the leaf
surface and appearing rather wart-like, which secrete digestive enzymes. An insect landing on the leaf causes
pressure, which results in the slow inrolling of the margin, thus pushing the insect towards the centre of the leaf,
and also stimulating the secretion of mucilage. A small insect is soon fast in the sticky secretion, and struggling
only causes it to sink deeper into the fluid. The stalkless glands secrete the digestive enzymes only if nitrogenous
compounds are present, thus a grain of sand blown on to the leaf may cause the inrolling of the margin but there
will be no secretion of enzymes. The digestive fluid is even more viscous than the mucilage, and contains the
enzymes ribonuclease, esterase, acid-phosphatase, amylase, and protease. Digestion is usually complete in about
24 hours, and the leaf slowly uncurls to re-set the trap. The hard indigestible remains of the insect dry out and
eventually may be blown away by the wind. The Butterworts usually capture small insects, such as mosquitoes 45
and gnats, which are attracted by a fungus-like or decaying odor.
The generic name is the diminutive of the Latin pinguis, 'fat' or 'fatty', referring to the greasy appearance of
the leaves, and ula 'little one'. The common name Butterwort also refers to the greasy appearance — wort is the
old Middle English word for 'plant'. The leaves of Butterwort were used in Scandinavia as a medicine, and were
rubbed on open sores of animals to make them heal faster. In fact, the digestive fluid does contain an antibiotic
to stop the decaying of the insects. Butterwort was also commonly used as a milk coagulant in home dairy
recipes.
Pinguicula villosa Linnaeus Hairy Butterwort
The Hairy Butterwort is found in many northern sphagnum bogs and swamps in North America and in
Eurasia, but it is usually so inconspicuous that it is rarely collected. In British Columbia it has been collected on
the Queen Charlotte Islands, at one station on Vancouver Island, and also apparently near Prince Rupert.
The leaves are 0.4-1.5 cm long, 0.2-0.7 cm broad, glabrous below and villous above. The scape is produced in
June to August and is 2-6(-12) cm tall and villous with capitate hairs, with a single nodding lavender-blue flower.
The sepals are (0.8-)l-2 mm long, and the corolla is 6-9 mm long, including the spur. The capsule is 3-5 mm long
and erect.
The specific name means 'covered with soft hairs', referring to the hairs on the leaf surface and on the scape.
Pinguicula vulgaris Linnaeus Common Butterwort
The common Butterwort is the most widespread species in the genus, being found throughout the arctic and
subarctic regions and up to 2300 m in the mountains of Eurasia, northernmost Africa, Iceland, Greenland and
North America. However, it is seldom abundant in its preferred habitats of bogs, mossy seeps, and wet soil on
beaches, meadows, or even on bare soil. In British Columbia it is found at relatively high elevations on
Vancouver Island and in the Coastal Mountains, and in valleys in the northern part of the Province.
The leaves are succulent, elliptic or oblanceolate, entire, 2-5(-6) cm long, and 7-18 mm wide. The petioles are
short and winged. The scape appears in (May) July to August, is 5-15 cm tall, and is obscurely glandular. The
calyx of the flower is 3-5 mm long. The corolla is deep violet-purple to lavender-blue or, rarely, nearly white. It is 46
(12-) 15-25 mm long including the slender spur (which is
about 10 mm long), and has a flaring throat and broad,
rounded lobes. The capsule is 4-6 mm long and is erect.
The species is represented by two subspecies in British
Columbia. Pinguicula vulgaris subsp. macroceras (Link)
Calder & Taylor (Figure 2) has flowers 24 mm long with
large broad, rounded lobes that are suddenly narrowed to
form a long filiform blunt spur up to 32 mm long. The lower
corolla lobes are oblong-obovate. This subspecies is found in
very different habitats ranging from sunny moist slopes to
glacier moraines and dripping rocks at elevations between
50-2300 m in the north Pacific coastal area, including Japan,
the Kamchatka Peninsula, the Aleutian Islands and Alaska.
In British Columbia it is present across the southern part of
the Province to the Rocky Mountains, but becomes almost
purely coastal (except for a few stations along the Skeena
River) in the northern parts. Pinguicula vulgaris subsp.
vulgaris is distinguished by the lower corolla lobes, which are
oblong rather than oblong-obovate.
The specific name vulgaris means 'common', 'general' or
'ordinary', while the subspecific macroceras is derived from
two Greek words, macro and ceras, meaning 'large horn',
referring to the large spur.
Both species of Pinguicula can be cultivated, preferring
moderate or cool summer temperatures with a relative
humidity of 60-100%. The Hairy Butterwort is probably best
grown in small compact-tufted species of living sphagnum
moss, but the Common Butterwort may be grown in a
mixture of Vi black peat, Vi sand and lA perlite. In
cultivation, all Butterworts are subject to attack by snails,
aphids and the fungus Ustilago pinguicula. This fungus
causes a smut and produces dark brown or black masses of
spores in the anthers. The disease is present in wild
populations of the Common Butterwort in British Columbia. Propagation is by seed, or vegetatively from young and
vigorous leaves. Butterworts do not like to have their root
systems disturbed, so are difficult to transplant.
Utricularia
Bladderwort
Most species of Bladderwort are aquatic (either anchored in
open water or free-floating), some are terrestrial on wet or
moist soil, and a few are epiphytic in moss. One or two
species are known that are restricted to the pools of water
found at the base of the leaves of large bromeliads in South
America. Representatives of the genus are found on all
continents of the world, with about 12 species occurring in
North America and five in British Columbia.
All species lack true roots and show little distinction
between the stem and leaves. In the aquatic species, the
leaves are submerged and very finely divided. All our native
species have alternate leaves, but in other species the leaves
may be partly or wholly whorled. There are bearded, hollow
bladders among the leaves or growing from specialized
stalks. The scape is tall and slender, and bears a loose raceme
of yellow flowers, each of which is subtended by a bract. The
flowers are strongly bilabiate, the lower lip is either entire or
3-lobed, and the upper lip is either entire or shallowly
FIGURE 2. Pinguicula vulgaris subsp. macroceras", Common Butterwort, in fruit, X1. 2-lobed. In most species the lower lip is raised at the base to form a prominent palate, which partially or
completely closes the opening to the corolla tube. The base of the corolla tube is extended backward and
downward to form a spur or sac. The fruit is a capsule. Most species form winter buds at the ends of the stems in
the fall, but some do not. These buds fall off and lie dormant at the bottom of the pond during the winter, ready
to produce new plants in the spring.
The bladders are modified leaves and there are many variations in form, with the smallest known being
0.3 mm and the largest about 6 mm in diameter. The best known one is that of the Greater Bladderwort
(Utricularia vulgaris). This is slightly less than 6 mm across, flattened, pear-shaped, and is attached to the plant
by a short stalk at one side. There is an opening at the narrow (free) end protected by a "door", which is
attached at the top and hangs downward over the opening, swinging free at the bottom where it rests against a
semi-circular collar or threshold. At the edge of the opening are a pair of branched antennae plus a number of
long slender bristles along the side. These form a funnel to guide the prey into the opening of the bladder. There
are (1)4 stiff bristles near the lower free edge that constitute the tripping or trigger mechanism. Scattered over the
outside of the bladder walls and on the threshold of the door are numerous stalkless glands, which secrete a
mucilaginous substance. On the outer surface of the door are many stalked glands, which also secrete mucilage
plus a sugary substance that is believed to attract the prey. The inside walls of the bladder are covered with many
2- to 4-headed glandular hairs.
When the trap is set, the door is closed, the trigger hairs are in position, the bladder is almost empty of water,
and the walls are straight or slightly concave. If a passing insect touches the trigger hairs, their movement alone
distorts the bottom edge of the door causing it to move away from the threshold. The vacuum inside the bladder
causes a suction so that water rushes into the bladder, carrying with it any insect or animal small enough to pass
through the opening. The bladder is now full of water and the walls bulge outwards. Specialized cells in the walls
of the bladder are concerned in the active transport of ions from inside the bladder to the water outside. This sets
up an osmotic gradient that results in water being moved out of the bladder, thus re-setting the trap. The
re-setting takes only about 30 minutes, although the prey may remain alive inside the trap for several days before
eventually dying. The mechanism of digestion is uncertain, but the insect quickly disintegrates after death,
therefore, there seems to be some kind of digestive fluid secreted into the bladder. In addition, there are reports
that the cells of the bladder become a red-brown color, suggesting some sort of metabolic activity. It has also
been found that the small pool of water that remains at the bottom of the bladder does contain the enzymes 4 /
esterase, acid-phosphatase, and protease. The indigestible, hard remains of the insect stay in the bladder.
The movement and trigger mechanism of Utricularia was the subject of an early movie film produced by Dr.
F. E. Lloyd of McGill University in the 1930's. Dr. Lloyd, who was interested in the physiological aspects of the
plants, also wrote one of the first comprehensive books on carnivorous plants (Lloyd, 1942).
Bladderworts can capture a wide range of organisms, including insects and insect larvae, diatoms, desmids,
protozoans, and small crustaceans such as Daphnea and Cyclops. They can also catch the occasional tadpole or
newly hatched fish, provided these are very tiny. Occasionally, bladders are found that have half a tadpole
sticking out through the door — obviously the victim was too large!
The generic name is derived from the Latin utriculus, 'little bag' or 'little bottle', referring to the bladders.
Utricularia gibba Linnaeus Humped Bladderwort
The Humped Bladderwort is found mainly in Central and South America and Africa, but is also present in the
eastern and western parts of North America. In British Columbia it is found on Vancouver Island, being
common in several lakes near Victoria and in Ash, Patterson and Turtle Lakes near Port Alberni, and on the
mainland in Beaver Lake in Vancouver.
The stolons (stems) are 10-25 cm long and are often a characteristic zig-zag shape when floating. The leaves
are 3-10 mm long, filiform, and dichotomously branched from the base. One branch is usually straight and
without bladders, while the other is usually further branched and has 1-3 bladders. The bladders are 0.5-1.5 mm
long, 0.5-1.0 mm wide, and have long branched hairs at the membrane opening. There are no winter buds. The
scape is 3-7 cm tall and there is either 1 or 0 scale. The inflorescence contains 1-2 bright yellow flowers, each
subtended by a semi-amplexicaulous bract. The pedicel is 5-10 mm long and is straight after anthesis. The calyx
lobes are equal, 1.5-2.5 mm long, and orbiculate. The upper lip of the corolla is 2-6 mm long, flat, and is at right
angles to the lower lip, which is 3.5-6 mm long. The palate is conspicuous and 3-4.5 mm long, and the spur is
cylindrical, 3-5 mm long, straight and parallel to the lower lip. The capsule is about 5 mm in diameter.
The species is represented in British Columbia by the subspecies gibba, the typical form.
The specific name gibba means 'swollen on one side'. Utricularia intermedia Hayne in Schrader Flat-leaved Bladderwort
Flat-leaved Bladderwort is a circumpolar boreal species, but is absent from Iceland, and is usually free-
floating in shallow standing or slowly moving water. In North America it is present as far south as California,
Indiana and Delaware. It is of scattered distribution in British Columbia, where it is often associated with Carex
lasiocarpa. Sometimes it grows in the sub-littoral zone of pools, anchored in the mud and forming large mats.
The stolons are very slender and are of two kinds:— leafy stolons 10-50 cm long, which lack bladders; and
leafless ones with bladders. The leaves are 5-30 mm long, and are divided at the base into 3 branches, which then
dichotomously branch into 6-20 slender segments. The segments are flat, have a central nerve, a blunt obtuse tip
with an abruptly starting bristle, and are all about the same width. The bladders are 1.5-4.5 mm long and 2-4 mm
wide. The winter buds are (2-)5-7(-15) mm long, 3-10 mm wide, and ovoid or ellipsoid. The scape appears in July
to August, and is 5-20 cm tall with one or two bract-like scales. The inflorescence has 2-4(-5) bright yellow
flowers, each subtended by a bract that is 1.5-4 mm long. The pedicel is 3-15 mm long. The calyx lobes are
subequal, 2.5-3.5 mm long and 2-3 mm wide. The upper corolla lip is 4-9 mm long, and the lower lip is
8-12(-18) mm long and 7-20 mm wide. The palate is well developed and about 7 mm long, and the spur is
cylindrical, straight, 8-12 mm long, and is at an acute angle to the lower lip. The capsule is about 3 mm in
diameter.
The specific name means 'intermediate', usually referring to the color, form or habit of the plant. This
particular species is intermediate in size between U. minor and U. vulgaris.
Utricularia minor Linnaeus Lesser Bladderwort
Lesser Bladderwort is a circumpolar species with small discontinuities in the distribution in Asia and America,
and is usually found in shallow standing or slowly moving water. In North America it extends as far south as
California, Colorado, Indiana and New Jersey. It is considered to be less common than the Flat-leaved Bladderwort (U. intermedia), but its small size may mean that it is frequently overlooked and, therefore, it may be more
common than the records indicate. In British Columbia it can be found in and around peaty bogs and swamps,
commonly occurring in small-channelled drainage areas, along and under the margins of these channels.
The stolons are 15-75 cm long and are differentiated into leafy ones with few bladders, and ones with fewer
leaves and more bladders. The leaves are numerous, 3-10 mm long, and dichotomously branched into 2-17 ultimate segments. These segments are slender, flat, without a central nerve, and the ultimate ones are strongly
acuminate. The bladders are borne on the leaves, usually 2-6 per leaf. They are 1.5-2 mm long and 0.7-1.5 mm in
diameter. The scape appears in June to September and is 4-15 cm tall with 1-4 bract-like purple scales. The
inflorescence contains 2-10 pale yellow flowers, each subtended by a purple auriculate bract that is 1-2 mm long.
The pedicel is 2-10 mm long, and is recurved after anthesis and in fruit. The calyx lobes are equal, 0.5-2.5 mm
long, 0.5-2.5 mm wide, and the lower lobes are emarginate. The upper lip of the corolla is 2-4 mm long, and the
lower lip is 4-8 mm long and 4-4.5 mm wide. The palate is about 3.5 mm long, and poorly developed. The spur is
small, 1-2 mm long, saccate, and poorly developed. The capsule is globose and about 2-2.5 mm in diameter.
A more robust form, which has the stolons 50-120 cm long, has been collected from several places in British
Columbia. The leaves of this form are 9-20 mm long, orbiculate, and divided into 20-40 segments.
The specific name minor means 'smaller', 'inferior', or 'less', referring to the habit of the plant. It is rarely
seen in flower in British Columbia.
Utricularia ochroleuca R. W. Hartman
This species is found in northern, central and western Europe, but is considered rare there, and in North
America from the Northwest Territories and Alaska east to Nova Scotia, and south to Washington, Oregon,
Colorado and Illinois. Its distribution in British Columbia is poorly known.
The stolons are 5-30 cm long and are differentiated into leafy and leafless ones. The leaves are 3-15 mm long,
and are divided into 3 parts at the base, these then further divide to form 5-19 segments. The segments have a
central nerve, and are gradually narrowed into a bristle at the apex. The leaves occasionally bear one bladder.
The bladders usually form on the leafless Stolons, and are about 1.5-2.5 mm long, and 1-2 mm wide. The winter
buds are subglobose, 1.5-5 mm long, and 1.2-2.5 mm wide. The scape appears in July to August, is 5-25 cm tall,
and has 1-5 bract-like scales. The inflorescence contains 2-10 lemon yellow or pale yellow flowers, each of which
is subtended by an auriculate bract about 0.5-4 mm long. The pedicel is 2-15 mm long, and is straight in anthesis
but becomes slightly recurved afterwards. The calyx lobes are subequal, about 2.5-3.5 mm long, 1.5-2 mm wide,
and the lower lobe is emarginate. The upper lip of the corolla is 3-9 mm long, and the lower lip is 5-12 mm long
and a little narrower than the upper lip. The palate is about 5 mm long, and the spur is 3-5.5 mm long, pyramidal
with a broad base, and at right angles to the lower lip. Non-flowering specimens are difficult to distinguish from non-flowering plants of the Flat-leaved Bladderwort (U. intermedia), and it may be that many collections have been misidentified. Flowering specimens are
easily referrable to this species because of the pyramidal spur at right angles to the lower lip, a characteristic that
is found in no other species in the area.
The specific name is derived from two Greek words — ochros, 'pale' or 'pale yellow', and leucon, 'white' —
meaning 'yellowish-white', referring to the color of the flowers.
FIGURE 3. Utricularia vulgaris, Greater Bladderwort.
A. Stolon and bladders, X 1, B. bladder, x 5.
49 Utricularia vulgaris Linnaeus (Figure 3) Greater Bladderwort
Greater Bladderwort is a circumpolar boreal species, with a break in the distribution in Iceland and
Greenland. It is present in ponds, lakes, marshes, and slow-moving streams. In North America it is widely distributed in a variety of suitable habitats as far south as California, Arizona and Texas.
The stolons are 20-180 cm long, and rather coarser than in the other native species. The leaves are numerous,
l-5(-9) cm long, pinnatifid, and divided into 20-150 filiform segments. The segments are terete and progressively
more slender after each division so that the ultimate ones are filiform and strongly acuminate. The bladders are
1-4 mm long, 1-3 mm wide and are attached near the base of the points of branching in the leaves by stalks
0.1-1.0 mm long. The bladders on the primary branches are larger than on other branches. There may be as
many as 10-50 bladders per leaf. The winter buds are ovoid or ellipsoid, 7-20(-30) mm long, and 6-15 mm wide.
The scape appears in May to August, and is stout, 6-30(-40) cm tall, and has 1-5 bract-like scales. The
inflorescence contains (3-)6-20 bright yellow flowers subtended by bracts 2.5-8 mm long. The pedicel is 6-30 mm
long, and is recurved after anthesis and in fruit. The calyx lobes are subequal, 3-6 mm long, 2-4 mm wide, and
the lower lobe is emarginate. The upper lip of the corolla is 3-17 mm long, and the lower lip is 5-20 mm long and
slightly lobed. The palate is gibbous, as large as the upper lip, and sometimes has reddish-brown veins on it. The
spur is 10 mm long, well-developed, falcate and directed forward. The capsule is globose, and about 6 mm in
diameter.
The American plants tend to have the spur somewhat more slender and pointed than in the Eurasian forms,
and have been separated out as U. vulgaris subsp. macrorhiza (Le Conte) R. T. Clausen. This is the form that is
present in British Columbia.
The specific name vulgaris means 'common', 'general' or 'ordinary', and the subspecific macrorhiza is
derived from two Greek words — macro, 'large', and rhiza, 'root' — meaning 'large-rooted'.
The aquatic species of Utricularia can be propagated from seeds in water, with plants reaching flowering size
in one to two years. They can also be propagated vegetatively very easily by taking stem cuttings.
£3 O Sarraceniaceae (Pitcher-plant Family)
This is a small family of low perennial herbs inhabiting bogs and wet savannahs, and having pitcher-like leaves
for trapping insects and other tiny animals. It contains about seventeen species in three genera, all endemic to
three widely separated regions of the New World. Heliamphora contains 4-6 species, which are found only on
isolated mountains of the Guiana Highlands of northern South America. Darlingtonia californica, California
Pitcher-plant or Cobra-plant, is the sole member of its genus, and is highly localized in montane meadows,
between 90-1830 m elevation, of northern California and adjacent Oregon. The third genus, Sarracenia,
contains between 8 and 12 species, all but Sarracenia purpurea being found only in the southeastern states of
America.
Sarracenia species grow only in very wet habitats, in waterlogged soil or even in standing water. Bogs are an
especially typical habitat. They are commonly found growing in acid soil, but this seems to be mainly because of
the lack of competition — specimens of S. purpurea have been found growing in humus derived from sphagnum
moss with a pH of 4.0, but also in an alkaline marl bog with a pH of 8.0. They were one of the first insectivorous
plants to be discovered, although not recognized as such at the time. The first drawing of S. purpurea was done
in Europe in 1601, and was reproduced in the second edition (1636) of John Gerard's Herball. This was one of
the first botanical books in English (instead of Latin) when it was originally published in 1597 as "The Herball
or General Historie of Plantes".
The genus consists of herbaceous perennials with round, horizontal to vertical rhizomes. The rhizome is
8-30 cm long when mature, and may survive for 20-30 years. The leaves are in a basal rosette and appear in two
or sometimes three forms each year. The most obvious are the pitchers, modified entire leaves that are produced
in the spring, with a second crop sometimes appearing in the fall, although these are often different in shape.
Secondly, half or all of the leaves in the fall (depending on the species) may be sword-like and tubeless, forming
the overwintering leaves or phyllodia. Finally, all species produce small scale-like leaves. The pitchers show great
variation between species, but in general either lie flat on the ground or arch upward, are 30-60 (-122) cm long,
and are strikingly colored, varying from yellowish-green to dark green to purplish and mottled with red to dark
red (the intensity of the red depending partly on the amount of sunlight in the habitat or the season). They are
fleshy, and a prominent wing runs along the outer surface of the leaf from top to bottom on the side facing the
centre of the plant. At the top of the leaf this wing broadens to form a narrow flange or lip around the front of the mouth of the pitcher and a hood on the back side. The hood is sometimes quite long and may arch over the
opening, thus forming a protection against rainwater entering the pitcher. The flowers appear in the spring to
early summer, and are large and showy with a more or less agreeable scent. There is a solitary leafless scape,
30-60(-122) cm tall (usually about as long as the leaves), which bears a single, nodding flower at its apex. There
are usually 5 (occasionally 4) sepals that are green or sometimes colored and petal-like, broad and spreading, and
persistent. The five petals are free, large, yellow to greenish-yellow or dark red, oblong to obovate, or they may
be very short and almost absent. They hang straight down and are incurved. There are twelve to many free
stamens. The base of the style is slender, but it is expanded at the top to form a very broad umbrella-like body
with 5 angles and 5 rays. There is a small hooked stigma at the end of each ray. The ovary is subglobose and
there are numerous ovules. The fruit is a 5-valved capsule with a granular surface. The seeds are numerous,
small, plump, and tan to dusty lavender in color.
The petals are persistent for several weeks and then drop, but the remainder of the flower persists until the
seeds are mature in July to September. The umbrella-shaped style is believed to catch pollen released by the
anthers, holding it where insects can easily find it. Only a minority of plants in a population seem to flower in
any one year. The species hybridize freely where their natural ranges overlap or in cultivation, and there are a
large number of natural hybrids, the most common probably being Sarracenia x catesbaei Elliott (S. purpurea
subsp. venosa x S. flava).
The prey are attracted to the pitcher by the flower-like appearance and odors. There are nectar-secreting
glands extending from the lip downwards on the outside. Insects land on the surface and gradually move into the
mouth of the pitcher as they feed on this nectar. The lip of the pitcher has stiff hairs that point downward into
the trap, and the insect cannot return once it has passed through this area. Just below the lip, in the steepest
portion of the throat, is a highly polished, slippery area without hairs, thus forcing the insect down into another
region of downward pointing hairs and then into a pool of liquid at the bottom of the pitcher. This liquid is a
mixture of digestive enzymes, rainwater (in some species), and bacteria, which are believed to help in digestion.
It is also believed that the liquid may contain a wetting agent to reduce the surface tension so that the trapped
insect is unable to crawl across the surface. It has been shown that the enzymes invertase and protease are
present. The chemical coniine has recently been shown to be present in very low concentration, sufficient to kill
any insect before digestion (Dr. G. H. N. Towers, personal communication). This is the same chemical that is
present in Poison-hemlock and that was used to kill Socrates. The trapped insects accumulate in the fluid, and £) 1
the digestive products are continuously withdrawn as the plant needs them.
The usual insects caught are ants, beetles and flies, although small toads, frogs and centipedes may also be
trapped. The quantity of prey captured is sometimes quite large — the pitchers usually live for several months
and may be almost completely filled with decaying remains. The odors of decay are often evident. Some animals
have learned to use the pitchers for food, shelter, or in part of their life-cycle. Small tree frogs often shelter
duiing the day in the mouths of large pitchers, using the suction pads on their feet to cling to the inside walls.
The fly Sarcophaga sarraceniae lays its eggs among the insect remains at the bottom of the pitcher, the larvae
feed on these and then bore their way out through the wall to pupate in the soil. It has been discovered that the
larvae produce an anti-enzyme as a protection against digestion. Some small mosquitoes also breed within the
pitchers in the wild.
Pitcher-plants have been cultivated as curiosities for many years, and have been used medicinally to a
limited extent. The Indians of Newfoundland used the roots of the northern form of the Common Pitcher-plant
as a medicine for smallpox, and the treatment was also used in England for a short period about 1865. Steam
distillation of the roots of S. flava produces sarasin, which was used to relieve tic douloureux, a painful facial
nerve irritation.
The generic name honors Dr. Michel Sarrasin (or Sarrazin) de l'Etang (1659-1734), a botanist and physician at
the Court of Quebec, who sent a specimen of Sarracenia purpurea subsp. purpurea to the French botanist
Joseph Pitton de Tournefort (1656-1708) about 1700.
Sarracenia purpurea Linnaeus Common Pitcher-plant
This is the most widespread species, being found in sphagnum bogs from the subarctic to the sub-tropics of
North America. It is now known to occur in all ten provinces of Canada, being found about eleven years ago in
northeastern British Columbia (Krajina, 1968).
The rhizome is stocky and more or less vertical. The pitchers are more or less suffused with purple or red,
(5-)10-20 cm long, 1-5 cm wide, obovoid, and curved-ascending from the base. They tend to be smaller and more
reddish when growing in an exposed situation. The wing is broad, and the hood is erect, open, reniform, and covered with reflexed bristles. The scape appears in late May to August, is erect, and 30-50 cm tall. The flower is
subglobose, 5-7 cm wide, and commonly dark purple-red or, rarely, yellowish. The style is greenish-yellow to
red.
There are two subspecies of the Common Pitcher-plant, which integrade in New Jersey. Sarracenia purpurea
subsp. purpurea or Northern Common Pitcher-plant occupies the northern part of the range, and is the form
found in British Columbia. It is characterized by long narrow pitchers that are usually more than three times as
long as broad, and by having smaller hood wings, which extend only slightly beyond the margin of the pitcher.
The petals and summit of the style are red. Sarracenia purpurea subsp. venosa is the southern form, which has
shorter, broader pitchers that are usually less than three times as long as broad.
The specific and subspecific epithet purpurea is from the Latin purpureus, 'purple', referring to the color of
the flowers and/or leaves. Sarracenia purpurea subsp. purpurea was designated as the Provincial flower of
Newfoundland in 1954, but was first used as an emblem on the Newfoundland penny about 1876 by order of
Queen Victoria. A potion of the roots and leaves was used as a cure-all for stomach, diuretic and menstrual
complaints and as a laxative, as well as for smallpox.
The Common Pitcher-plant is somewhat difficult to cultivate, and is probably best grown on live sphagnum
moss with semi-shade and cool root temperatures during the hot summer months, after it has reached maximum
development. Full sun is needed before this for best flower and pitcher formation. High humidity is essential.
However, we are presently growing the northern form successfully in The E. H. Lohbrunner Alpine Garden at
UBC in a peat bed using a mix of one-third peat moss, one-third coarse sand and one-third loam. The plants are
in full sun, but are never allowed to dry out in the summer. It has been stated that water from a water softener or
deionizer should not be used as the added sodium is detrimental to the plant. Propagation is by seeds, which
reach flowering size in 3-5 years, or by division of the rhizome.
Insectivorous Fungi and Seeds
In addition to the insectivorous plants, it has been found that some fungi are carnivorous. Arthrobotrys
obligiospora and Dactylaria brochopaga both trap eelworms in loops of their hyphae, and then ingest needed
nutrients from the body.
There are also indications that the seeds of some non-insectivorous plants may be carnivorous, or capable of
carnivory. John T. Barber of New Orleans has found that seeds of the Common Shepherd's-purse (Capsella
bursa-pastoris) are apparently capable of trapping prey. The seeds are mucilaginous, that is, they release a
gummy covering or pellicle when they take up water. The mucilage appears to contain a substance that will
attract motile soil bacteria, nematodes and protozoans. Barber showed that nematodes accumulated in
statistically significant numbers around Shepherd's-purse seeds, and that a significant number of them would be
dead (75% versus 7% in the absence of seeds after 8 days). Therefore, he suggested, and later proved, that the
seeds released a toxin. He also showed that Shepherd's-purse seeds released proteases on imbibition (uptake of
water), and that the proteolytic activity was confined to the mucilage. Experiments using labelled amino-acids
proved that the germinating seeds were able to take up products of digestion, and that they were incorporated
into the seedling. The seeds are small and contain minimum food reserves, and the plants grow in semi-arid soils
of low fertility. Thus, although the plants do not appear to needprey to germinate, or for the seedlings or plants
to be healthy, the potential for carnivory seems to be present — the seeds can attract, trap, kill, digest, and
absorb the resultant nutrients.
REFERENCES
Barber, J. T. 1978. Capsella bursa-pastoris seeds. Are they "carnivorous"? Carnivorous Plant Newsletter 7:39-42.
Ceska, A. and M. A. M. Bell. 1973. Utricularia (Lentibulariaceae) in the Pacific Northwest. Madrono 22:74-84.
Carnivorous Plant Newsletter, volumes 1-7 (1972-1978).
Darwin, C. 1876. Insectivorous Plants. John Murray, London.
Emboden, W. A. 1974. Bizarre Plants. Magical, Monstrous, Mythical. Macmillan Publishing Co., Inc., New York.
Farrer, R. 1908. Alpines and Bog-Plants. Edward Arnold, London.
Heslop-Harrison, Y. 1978. Carnivorous Plants. Scientific American 238(2):104-115.
Krajina, V. J. 1968. Sarraceniaceae, a new family for British Columbia. Syesis 1:121-124. Lloyd, F. E. 1942. The Carnivorous Plants. Republished in 1976 by Dover Publications Inc., New York.
Savage, C. and A. Savage. 1979. Canada's carnivorous plants. Nature Canada 8(1):4-12.
Schwartz, R. 1974. Carnivorous Plants. Edited by Deborah Leavy. Praeger Publ., New York.
Shetler, S. G. 1974a. Sarraceniales. In: Encyclopaedia Brittanica, 15th edition, pp. 252-256.
 . 1974b. Nepenthales. In: Encyclopaedia Brittanica, 15th edition, pp. 958-962.
Taylor, R. L. and B. MacBryde. 1977. Vascular Plants of British Columbia: A descriptive resource inventory. Technical Bulletin No. 4. The Botanical Garden of The University of British Columbia. University of British Columbia Press, Vancouver,
B.C.
All-America Selection Display Garden
The UBC Botanical Garden had an All-America Selection Display Garden for the first time this summer. In
this garden were displayed All-America flower winners from the immediate past and present years. The garden
was located in the entrance area to the Main Garden, and was open to the public at all times. Displayed in the
Garden were plants grown from seed supplied by three seed houses as well as the All-America Selection
Committee. The garden will be maintained in future years, with a changing display of flowers each year.
53
FIGURE 4. A view of the All-America Selection Display Garden in late September. The Genus Lonicera in British Columbia
Member of the Family Caprifoliaceae
LONICERA CAERULEA Linnaeus
Bluefly Honeysuckle
LONICERA CILIOSA (Pursh) A. P. de Candolle
Western Trumpet Honeysuckle
LONICERA DIOICA Linnaeus
Glaucous-leaved Honeysuckle
LONICERA ETRUSCA Santi
Etruscan Honeysuckle
LONICERA HISPIDULA (Lindley) D. Douglas ex Torrey & Gray
Hairy Honeysuckle
LONICERA INVOLUCRATA (J. Richardson) Banks ex K. P. J. Sprengel
Twinberry Honeysuckle
LONICERA x NOTHA Zabel
Tartary Hybrid Honeysuckle
LONICERA UTAHENSIS S. Watson
Utah Honeysuckle
Description of the Genus Lonicera
Lonicera is a genus of vigorous erect shrubs or woody vines. The shrub forms may be bushy or climbing,
deciduous to evergreen, and 1.2-3 m tall. The woody vines are sometimes more or less evergreen. All forms have
thin stems and hollow branchlets.
The twigs are rounded and mostly slender. The pith is moderate, pale or brown, and is excavated at the nodes
in some species. The leaf scars are opposite, crescent -shaped, small, and on the narrowed extremities of raised
bases that are more or less connected by transverse lines. Bundle traces 3. Stipule scars lacking.
The winter buds are often superposed, with the lowermost being the largest or developing into an inflorescence
in the first season. They are sessile, variously shaped, and have 2 to rather numerous 4-ranked scales.
The leaves are opposite, simple, mostly entire, short-petiolate or sessile, and exstipulate or rarely stipulate,
when the stipules are usually small and adnate to the petiole. Few show fall color.
The flowers are either in axillary pairs on a slender petiole subtended by 2 bracts and 4 bractlets, or in sessile
whorls at the ends of the branches. They are perfect, white or yellow to pink, scarlet or purple, usually small but
showy, and often have a fragrance that is most marked towards the evening thus attracting night-flying hawk-
moths and hummingbirds. The flowers are subtended by bracts and bractlets, the bractlets being distinct,
connate or sometimes wanting. The calyx is small, with the 5 sepals united to form an ovoid to nearly globulose
tube, with a shallowly 5-lobed limb, the lobes sometimes being obsolete. The corolla is regularly or nearly
regularly 5-lobed or often evidently bilabiate with a 4-lobed upper lip and a single strap-shaped lower lip. It is
tubular to funnelform or campanulate, and is often gibbous (swollen) or spurred near the base. Nectar is
secreted at the base of the corolla tube. There are 5 stamens, which are inserted on the corolla tube and alternate
with the corolla lobes. The anthers are linear or oblong. The style is slender and elongate, and the stigma is
capitate. The ovary is inferior and usually 2-3(-5)-celled. The ovules are pendulous, and there are 3-8 per cell.
The flowers generally appear in the spring, but in late summer in some species. C X 0.65
B xl.3
55
_i.5m
FIGURE 5.   Lonicera invotucrata var. involucrata. A. Habit, B. flowers, C. fruiting branch, D. fruit and seeds. 56
The fruit is a small fleshy berry and may be bright red to bright yellow or orange, blue or black. They are ripe
in the summer or early fall, and those of paired flowers may be partially or completely united. The fruits are
attractive to birds. The seeds are few to many depending on the species. They are oval or oblong but appear
elliptic in longitudinal section, small, and the outer epidermis is often thick, pitted and lignified. There is no data
available to show how old plants must be to produce good seed crops. Seed dispersal is primarily by birds and
animals.
There are more than 150 species of Lonicera distributed over the temperate and subtropical regions of the
Northern Hemisphere. About 20 species are native to North America, of which 6 occur naturally in British
Columbia. Two introduced species may also be found growing wild in certain parts of the Province.
Key to the species in British Columbia
Lower leaves with well developed, connate, stipule-like appendages at the base of the short petiole; flowers
in terminal or terminal and axillary inflorescences with several compact whorls; corolla strongly
bilabiate. Vine or climbing shrub Lonicera hispidula
Lower leaves without stipule-like appendage; flowers in terminal whorls or axillary pairs; corolla weakly or
strongly bilabiate.
Corolla strongly bilabiate.
Evergreen or half-evergreen vine (occasionally deciduous), 2-4(-9) m long; flowers in terminal
dense spikes. Adventive species Lonicera etrusca
Deciduous shrubs; flowers in terminal clusters or axillary pairs.
Flowers in terminal almost stalkless clusters; bushy shrub, 1.5-6 m tall with twining or
trailing stems about 60 cm long Lonicera dioica
Flowers in axillary pairs; shrubs, 2-3 m tall. Adventive species Lonicerax notha
Corolla regular or only weakly bilabiate.
Twining vine with widely branched stems often 5-6(-9) m long; upper 1-3 pairs of leaves on
flowering branches connate (united around stem) at base; flowers in terminal
compact spikes of 1-3 whorls Lonicera ciliosa
Shrubs; upper leaves not connate; flowers in axillary pairs.
Leaves (3-)5-12(-15) cm long; flower pairs closely subtended by two pairs of large leaflike green or purple conspicuous bracts; erect bushy shrub, 0.5-3(-4) m
'   tall Lonicera involucrata
Leaves less than 9 cm long; flowers not subtended by obvious bracts; shrubs less than
2 m tall.
Fruit red and often of unequal size with one member of pair not developing;
leaves 2-5(-8) cm long; ovaries united at base when mature; erect or
somewhat straggly shrub, 0.6-2 m tall Lonicera utahensis
Fruit blue or blue-black (occasionally red), both usually of equal size; leaves
1.2-7.5(-9) cm long; ovaries apparently united but actually separate
within a narrow-mouthed cup formed by the united bractlets;
sturdy shrub, (0.2-)0.5-l .5(-2) m tall Lonicera caerulea
The Native Species
Natural Distribution and Habitat
Lonicera caerulea is a widespread species with several varieties, and is present throughout many areas of
northern and central Europe, Asia and North America. In North America it is present south of approximately
latitude 60°N, extending south to California, Nevada, Wyoming, Minnesota and Pennsylvania. It is usually present on streambanks and in other moist to sometimes rather dry places, at moderate elevations in the mountains. In British Columbia it is common at moderate to rather high elevations in the mountains of the Interior
Hemlock Zone of the southern part of the Province, extending east to Alberta. The northern limit in British
Columbia may be about latitude 52°N.
Lonicera ciliosa is present from British Columbia south to northern California, especially west of the Cascade
summits, and east to Montana, Utah and Arizona. It is widespread in open woods and thickets, hillsides and
ridges, from sea level to moderate elevations in the mountains. In British Columbia the species is common in the
southern half of the Province in open woods and brush below 460 m west of the Cascades and in the Wet Interior
Zone, and there is sporadic occurrence to the Kootenay River Valley. It is often found clambering through small
conifers in the woods.
Lonicera dioica occurs from Quebec south to North Carolina and Iowa, and west to the Mackenzie District,
eastern British Columbia, South Dakota and Oklahoma. It is usually found in woods, often in wet places. In
British Columbia it is present in the northeastern and southeastern regions of the Province, in semi-open dry
forests above 610 m in the Rocky Mountain region. Collections have been made at Fairmont, Windermere,
Field, Peace River, Hudson Hope and on the Liard River near Smith River.
Lonicera hispidula occurs from southwestern British Columbia south to southern Oregon and southern California, and on Santa Cruz and Santa Catalina Islands. It is found in open woods and thickets west of the
Cascade summits. In British Columbia it occurs west of the Cascades on rocky dry open hillsides, woods and
thickets, often sprawling over broken rock or low shrubs. It is present in the southern Coastal region, the Gulf
Islands, and on south Vancouver Island.
Lonicera involucrata occurs from southern Alaska south along the coast to Santa Barbara County in
California and Chihuahua in Mexico, and east to Quebec, Lake Superior, New Brunswick, Colorado and Arizona. It is present on fairly moist to wet soil in woodlands and thickets from sea level to rather high elevations in
the mountains. In British Columbia it is rather sporadic but often locally abundant on moist sites with rich soil
to at least 1525 m throughout the Province. It is especially abundant in the Coastal and Interior Wet Belt Zones.
It forms a conspicuous element of the lowland coastal vegetation throughout the Queen Charlotte Islands. On
the coast it is associated with Thimbleberry, Red-osier Dogwood and Pacific Crabapple. In the interior it is
usually found in more open situations with Salmonberry, Red-osier Dogwood, Water Birch and Cow Parsnip. tD /
Lonicera utahensis is present from southern British Columbia and the Olympic Mountains of Washington
south to Crater Lake and the Blue Mountains of Oregon, and east to Alberta, Montana, Wyoming and Utah.
There are varied reports of its occurrence in northern California, although Munz and Keck (1973) state that it
probably does not occur in that state. It is widespread on moist wooded or open slopes, by stream banks and at
the edges of bogs, at moderate to rather high elevations in the mountains. In British Columbia it is widely distributed across the southern parts of the Province in moist places from valley bottoms to the timberline east of
the Cascades to the Rockies, and also at subalpine elevations at the Coast. It also extends north in the mountains
to latitude 52°N at Anahim Lake and in the Rockies.
Description
Lonicera caerulea — a highly variable species. Branchlets stiff and glabrous, or hairy only when young,
although some forms have branches that are much more hairy or downy. The bark is often shredded and light
brown. Winter buds are often superposed, short, spreading, and with valvate lower scales. The leaves are bright-
green, elliptic to oblong-obovate or oblong, have a rounded or obtuse apex, and are 0.6-3(-4) cm broad. They are
glabrous or sometimes pubescent above, but are more or less hairy beneath, especially on the midrib and veins.
The margin is sometimes villous-ciliate. The petiole is 2-5 mm long and hairy. The flowers are in axillary pairs,
pale-yellow to yellowish-white, sometimes tinged with red, and appearing in (April-)June to July throughout the
range. The corolla is funnelform, (0.9)1.0-1.3(-l.9) cm long, and 5-lobed to scarcely bilabiate. There is a short
thick or gibbous spur at the base of the tube. The corolla tube is hairy within and without. The bracts are paired,
green, oblong-linear, and narrow, 3-6(-10) mm long, and the longer ones are somewhat foliaceous in texture but
still inconspicuous. The bractlets are wholly connate to form a narrow-mouthed cup completely enclosing the
ovaries. The peduncle is axillary on twigs of the season. The fruit is oval to ellipsoid, and 0.5-1.0 cm diameter.
2n = 18, 36.
Lonicera cilosa — a twining vine, often growing through and around the branches of other shrubs or small
conifers. The twigs are hollow, and glaucous when young, becoming glabrous with age. They are sometimes
more or less evergreen. The leaves are mostly elliptic to occasionally ovate or obovate, with a rounded to acute
apex and wedge-shaped base, 4-10 cm long and 2.5-5 cm broad. They are green and glabrous above, and glaucous blue-green and slightly downy beneath. The margins are ciliate. The petiole is 3-5(-12) mm long. The
flowers are yellowish to reddish-orange, sometimes tinged purple on the outside, and appear in May to July (to
August) throughout the range. The flowers are fragrant. The corolla is tubular-funnelform to narrowly funnel-
form, (2-)2.5-4 cm long, and shallowly or not very strongly bilabiate. The corolla tube is downy outside and
pubescent on the inside adjacent to the stamens, and there is a gibbosity on one side at the base on the outside.
The stamen filaments and style are slightly exserted and hairy. The flowers are sessile. The fruit is orange-red to
coral-red with orange pulp inside, 5-6(-10) mm diameter, and in groups of 3 to 4 that develop at the expense of
the remainder. The seeds are large and yellow. The fruit is ripe in September.
Lonicera dioica — a bushy shrub, with essentially glabrous stems. The bark is gray or straw-colored. The
winter buds are solitary, ovoid, and have ovate to narrowly triangular scales. The leaves are usually elliptic to
obovate but are variable, tapered at the apex and base, 3.5-10 cm long and 2.5-5 cm broad. They are green and
glabrous above, glaucous and more or less pubescent below. The margin is finely hairy. The upper one or two
pairs of leaves are united around the stem at the base forming a concave disc. The remainder of the leaves are
almost sessile. The flowers are yellowish, but often tinged purplish or reddish, and appear in June and July
throughout the range. Two or more pairs of leaves beneath the inflorescence are united. The corolla is strongly
bilabiate, with the lobes varying from about half the length of the tube to approximately equal to the tube in
length. The tube is slightly swollen at the base, and is glabrous outside and glabrous to densely hairy within. The
filaments of the stamens are attached nearly at the opening of the tube, and have downy bases. The style is
usually glabrous to somewhat pubescent, and is distinctly longer than the tube. The fruit is coral-red, nearly
10 mm in diameter, and fleshy. The seeds are light brown, elliptic, and 3.4 mm long by 2.6 mm wide. A slight
groove or ridge may be apparent. The fruit is mature in July to August or September, and is dispersed during
August to September or October.
Lonicera hispidula — vine or climbing shrub with slender, freely-branching stems l-3(-4) m long. The
branches are villous-hirsute with glandular hairs, varying to glabrous. The young stems are often hairy, varying
to glaucous or glabrous. The leaves are ovate to ovate-oblong or somewhat elliptic, often cordate or subcordate
at the base, 2-7 cm long and 1.5-5 cm broad. They are green and glabrous or hairy above, glaucous and more or
less pubescent to glabrate beneath. The upper 1-3 pairs of leaves are connate around the stem. The petioles are
short. The central axis of the inflorescence varies from glabrous to hirsute or glandular. The flowers are reddish
to purplish, often being yellowish inside in the latter state, or sometimes pink or yellow tinged with purple. They
appear in June to August over the range. The corolla is 1-2.5 cm long, and is strongly bilabiate with the lobes
about as long as, or longer than, the tube. The tube is slightly gibbous near the base, and is densely hairy inside
and glandular-pubescent outside. The stamens and style are long exserted, and the filaments are hairy below.
The fruit is red, almost 10 mm in diameter, and is rather juicy. Only a few fruits will mature in most years.
2n = 18.
Lonicera involucrata — an erect to spreading or rounded shrub. The twigs are stout, 4-angled, and glabrous or
sometimes long-hairy, especially when young. The bark is light-brown or purplish, becoming gray with age, and
freely exfoliating. The winter buds are often superposed, glabrate, and the bud-scales are parted. The leaves are
narrowly ovate to oblong, oblong-ovate or obovate, with an acuminate and acute to rounded apex, and a cordate to cuneate base. The leaves are 2-7.5(-8) cm broad, and are prominently veined. They are dull dark green
and usually glabrous above, occasionally being sparsely pubescent on the veins, and bright green and glabrous to
slightly downy beneath, especially along the main veins. The margin is often ciliate. The petiole is 5-6(-12) mm
long. The flowers are yellow, sometimes tinged with red, and appear in March or April to August, partly
depending on the elevation. The corolla is narrowly funnelform to subcylindrical, l-1.5(-2) cm long, and nearly
regular with the lobes almost equal. The tube has a gibbosity or short thick spur on one side at the base, and is
glandular-hairy outside. The stamens are shorter than the tube, and are glabrous, or nearly so, and glandular.
The anthers are often slightly exserted. The style is slender, longer than the stamens, and is glabrous. The bracts
are (0.8-)l-l .5(-2) cm long, ovate or oblong, glabrous or often glandular-pubescent, and dark-red. The bractlets
are axillary, slender, and 0.5-5 cm long. The fruit is shining purple-black or black, globose to ovoid, 8-10 mm in
diameter, paired and wholly distinct above the bracts. The seeds are slightly shiny, black, elliptic, 2.7 mm long
and 1.8 mm wide. The fruits are mature in July to August, and are said to be poisonous by the Indians, although
they are so bitter and nauseous to taste that there is little danger of a sufficient number being eaten to cause
concern. 2n = 18.
Lonicera utahensis — a shrub, with a number of irregular straggling branches from a long thin stem. The
branches are slender, spreading and glabrous. The bark is gray, and often rather dead looking. The leaves are
elliptic to somewhat ovate or oblong, with a broadly rounded to obtuse apex and an obtuse to blunt or
sometimes subcordate base. They are 1-4 cm broad, pale green and glabrous above, and glabrous to more or less pubescent to hirsute below. The margin is coarsely ciliate near the base of the leaf. The petioles are 2-5 mm long.
The flowers are in axillary pairs, although one member of the pair is often dwarfed, and are white to pale yellow,
often fading to salmon-yellow. They appear in May to July. The corolla is tubular, 1-2 cm long, and nearly regular with the lobes more or less equal in size and shape but arranged to seem obscurely bilabiate. The base of the
tube is gibbous at one side, or the gibbosity is sometimes enlarged to form a short thick spur, and it is occasionally pinkish in color in that area. The stamens and style are glabrous. The ovaries are divergent and weakly or
scarcely united at their bases when young, becoming firmly united when mature. There is 1 pair of bracts, which
are narrow and (l-)1.5(-3) mm long. The bractlets are about 1 mm long or less, or obsolete. The peduncle is common to two flowers, and is slender and l-1.5(-2) cm long. The fruit is globose, pulpy and soft, and 6-10 mm in
diameter. There are 2-4 seeds. The peduncles elongate in fruit to 35 mm long.
Varieties and Ornamental Cultivars
The members of the genus Lonicera hybridize readily, and there are therefore a number of ornamental
cultivars in some species.
Lonicera caerulea is a variable shrub with several varieties throughout its range. The form found in British
Columbia is L. caerulea var. cauriana (Fernald) Boivin, which is the cordilleran form of the species. This variety
differs from the type in that it is 0.2-1 m tall, the branches are pruinose or puberulent and sparsely hirsute, and
the leaf margins are always villous-ciliate. The fruit is often red instead of blue or blue-black, and the seeds are
whitish-brown, orbicular and 1-1.7 mm long.
Lonicera ciliosa has one named variety, var. occidentalis, in other parts of the range, which has more brightly
colored fruit than the type.
Several varieties of Lonicera dioica are recognized. It is represented in British Columbia by L. dioica var.
glaucescens (Rydberg) Butters in Clements, Rosendahl & Butters. This form occupies the more western part of
the range of the species, and differs from the type in that the leaves are obovate to narrowly elliptic and are
5-8 cm long. The flowers are more purplish, and become reddish with age. The corolla tube is often downy
outside as well as inside.
There are several forms of Lonicera involucrata. The form occurring in British Columbia is the typical var.
involucrata, which has the corolla rarely tinged with red, and the plant is less than 1 m tall.
Propagation
The genus is easily propagated by either seeds or vegetative means, but the species hybridize readily so that
forms may not come true from seed. However, the seed is an excellent means of raising new forms and is used
for plant breeding purposes. The seeds of many species show double dormancy, so that germination may take a
long time.
The fruit should be collected when it is ripe, and the seeds extracted by macerating the fruit in running water,
allowing the empty seeds and pulp to float away. The seeds should be dried, and then either stratified or stored in
a sealed container at 1-3°C for up to one year. In the absence of other information, the seed should be stratified
for 3 months at 5CC and then sown. If germination does not occur within 4 months, the seeds can be stratified
for a further 3 months at 5°C.
Most of the North American species, except L. dioica, will germinate best if they are warm stratified at room
temperature for 3 months, followed by 3 months cold stratification at 5°C before sowing. The germination
period should have alternating temperatures of 30°C day and 20°C night. This regime, should give satisfactory
results. Lonicera dioca may be sown in the spring without treatment with up to 95% germination after
80-100 days.
The seeds should be covered with 3-6 mm of soil when sown. Mulching with 5-7.5 cm of straw helps to prevent
excessive drying during the germination period.
Softwood and hardwood cuttings are both easy to root, although it is probably best to use moderately firm
young shoots taken in July or August. The cuttings should be planted either in straight sand or in peat moss, and
kept moist and shaded. The use of a rooting hormone has been found to be advantageous. It is also possible to
take cuttings and plant them in very sandy soil under a cloche outside. Each node can be split in half lengthwise
and will root to produce two plants.
In addition, Lonicera species may be propagated by layering and by the division of large individual plants
using a sharp spade. Transplanting
Shrubs can be transplanted successfully, particularly after growth has been completed for the season. Pruning
of weak wood and removal of some new growth will enhance success of the transplant.
Conditions for Cultivation
Lonicera species all like a well drained good loamy soil with a pH about 6.0. All forms prefer positions where
the aerial parts are in the sun but the roots are shaded and cool (much like Clematis). All are easily grown within
their hardiness limits (some are hardy to Zone 2 Canadian, or even possible to Zone 1), and will usually produce
a wealth of foliage, flowers and fruit with very little care. The vine forms require a trellis, arbor or other
support.
Lonicera ciliosa is hardy to USDA Zone 6. Lonicera dioica is hardy to USDA Zone 2, and will succeed very
well in good garden soil, forming a low spreading rather elegant bush if given the support of a stout central
stake. Lonicera doica var. glaucescens is hardy to Canadian Zone 2 or USDA Zone 4. Lonicera involucrata is
hardy to Canadian Zone 1 or USDA Zone 4, and is a robust adaptable species growing equally well in seaside
gardens and in industrial areas. Lonicera utahensis is hardy to USDA Zone 6.
Lonicera species require little regular pruning. The shrubby forms benefit from an occasional shortening of
long growths that are spoiling the shape of the plant, and old weak growths of the climbing forms may be pruned
out. They should all be pruned immediately after flowering.
Landscape Value
The genus contains a number of extremely beautiful species, and every climbing species that is hardy is worth
growing. Several species are among the best flowering shrubs for the colder areas of Canada. Some forms, however, may be too large for the home garden, although they are excellent for shrubbery or in a wild garden. Several species have outstanding fragrance, and many have a good display of flowers, colorful fruit, and an
attractive habit of growth. The fruits are very attractive to birds.
The climbing species are best scrambling over other bushes or tree stumps, trellises or pergolas, although some
may be trained to grow as small standards. Some of the less rambling forms may be trained up stout posts
1.25-1.80 m tall, and then allowed to form loose spreading shrubs with no further support.
Lonicera caerulea has little or no merit for gardens, although it is botanically interesting. The native form,
variety cauriana, is of value for its habit of growth, flowers, and fruit.
Lonicera ciliosa is rare in gardens, but is the showiest native honeysuckle, and should be more widely cultivated. The orange flowers are particularly attractive to hummingbirds.
Lonicera dioica will form a low spreading, rather elegant, bush if given the support of a stout central stake. It
has no great beauty of flower, but is striking because of the very glaucous undersurfaces of the leaves. Lonicera
dioca var. glaucescens has more attractive flowers than the species, and can be used as a perennial vine.
Lonicera hispidula should be grown in a native garden for its attractive habit of growth.
Lonicera involucrata is a loose straggly bush that has no real ornamental value, except that it is hardy to
Canadian Zone 1, and would therefore be useful in such very cold areas. It is also useful in a native garden, and
as ground cover bush plantings in revegetation programs.
Lonicera utahensis has an attractive habit of growth as well as attractive flowers, and is a particularly
charming small shrub for shady gardens.
Availability
The native species are apparently not available from local nurseries, although Lonicera caerulea var.
dependens may be found.
Other uses
Hummingbirds and long-tongued insects are attracted to the flowers because of the nectar, and birds eat the
berries. Bears also apparently consider the berries of Lonicera involucrata as a favorite food — in fact this
species was called Bearberry or Grizzly Berries by the native Indians.
Lonicera dioca var. glaucescens and L. involucrata are both used as habitat or food by wildlife, and L. involucrata may also be used as watershed plantings or in forest management.
The Indian tribes of British Columbia and western Washington had various uses for several of the native
Lonicera species. In British Columbia, the children of the Lower Lillooet tribe at Pemberton used to suck the nectar from the
flowers of L. ciliosa. The Thompson Indians used to peel the thicker stems of this species, boil them and then
drink the decoction as a tonic. They also used a fibre obtained from the stems as a thread or twine. The Sechelt
Indians used to crush the tender green shoots and steep them in water, which was then used as a hair tonic. The
Saanich tribe believed that the berries were poisonous. The Squamish Indians believed that the vine was used as a
swing by ghosts — their common name for the plant means "swing of the ghosts".
The western Washington tribes made more use of Lonicera ciliosa than did the British Columbian Indians.
The Swinomish boiled the bark as a tea for colds and sore throats, or swallowed the juice from chewed leaves for
a cold. Leaves were bruised and soaked in hot water, and a woman then held her breasts over the steaming water
to stimulate the flow of milk after childbirth. The Chehalis tribe crushed leaves in water, which was then used to
bathe little girls so that their hair would grow long and shiny. The Chehalis and Squaxin tribes dipped leaves in
water that the women then drank as a contraceptive, but the Squaxin also drank this water as a treatment for
"womb troubles". The Klallam chewed the leaves and then put them on bruises. The Snohomish believed that
crows used the plant as a swing.
Lonicera involucrata was used by the Indians much more than any other Honeysuckle. The Thompson and
Lillooet tribes of British Columbia believed the berries to be highly poisonous — but the Flathead Salish of
Montana ate them as a powerful laxative. Most tribes considered the berries to be at least inedible if not actually
poisonous, except for birds and bears. The Bella Coola Indians chewed the leaves and applied the cud to an
"itch", or applied it to an opened boil to draw out the poison. The leaves were crushed and applied to burns and
to the sores of gonorrhoea. Gonorrhoea was also treated by applying toasted, pulverized bark. The bark was
boiled and the decoction drunk for a cough. The Southern Carrier boiled the bark for five hours, and then used
the decoction daily as an eyewash. The Gitksan used the fresh juice of berries for sore eyes, or the inner bark
(when berries were unavailable) was soaked in water to make a milky solution used as an eyewash. The
Thompson Indians boiled the leaves and then applied them to any part of the body that was swollen. Sometimes
the leaves were bruised before boiling as this was believed to greatly increase their strength. They also used a
decoction of leaves and twigs as a liniment. The Squamish chewed the bark and blew it onto the cheek for a
toothache, or blew the fresh saliva onto a boil — this was said to be particularly effective if done in the morning
immediately after waking, but before speaking. The Haida rubbed the berries into the scalp to prevent gray hair.
Lonicera involucrata was an important medicinal plant for the Kwakiutl tribe on Vancouver Island. They boiled
the leaves (or roots in winter), strained them and applied them as a hot compress to swollen shoulders and feet.
The bark was boiled and placed on the breasts of a woman who had just given birth to make her milk flow, or
the leaves were mixed with the basal leaves of Yarrow (Achillea millefolium) for the same purpose. The bark,
berries or leaves were mixed with Sea Wrack (Fucus gardneri) or Sea Lettuce (Ulva lactuca), dried Tobacco
(Nicotiana sp.), and Red Alder bark (Alnus rubra) to make a poultice for swellings and sores. They never ate the
berries, believing that they would lose their voices and become stupid if they did so, but they did make a purple
dye from the berries mashed with Salal berries (Gaultheria shallon).
The Quileute Indians of western Washington used the juice from L. involucrata berries to paint dolls' faces.
They chewed the leaves as an emetic when poisoned. The women of the Quinault and Makah tribes chewed the
leaves during childbirth. All the western Washington tribes associated the plant with the crow.
Indian hunters in the Okanagan sometimes ate the very juicy berries of L. utahensis, believing them to be a
good emergency supply of water.
Diseases and Problems of Cultivation
The climbing members of the genus are very subject to aphid attacks in summer, especially during hot dry
spells. However, these attacks are often overcome naturally if the plants have been given good loamy soil and
cool moist root conditions. Some vines are very susceptible to infestations of spider mites.
In general, the genus is not susceptible to severe disease problems, although Leaf Curl makes the leaves
thicken, curl and become deformed, and Powdery Mildew may be a problem in some areas, including the Lower
Mainland of British Columbia.
The following diseases have been identified on the native species in the wild in British Columbia:—
Lonicera ciliosa — Cercospora antipus. Leaf Spot; C. periclymeni; Poria ferrea, Stem Rot; and Stereum sp.
Lonicera hispidula — Microsphaera penicillata, Powdery Mildew; and M. alni. Lonicera involucrata — Kabatia lonicerae var. involucratae, Leaf Spot; Microsphaera penicillata, Powdery
Mildew; M. alni; Leptosphaeria dumetorum; and Ophiobolus minor.
Lonicera involucrata — Kabatia lonicerae var. involucratae, Leaf Spot; and Leptothyrium periclymeni var.
periclymeni.
The Adventive Species
Two other species of Lonicera have been reported either as adventive, that is, they have escaped from cultivation and have persisted in the wild for short periods of time, or as naturalized, that is, a thoroughly established
alien reproducing without cultivation.
Lonicera etrusca Santi, Etruscan Honeysuckle, is a native of the Mediterranean region that has now become
naturalized and established in thickets along the coast of Oregon (in Lane and Curry Counties), in Del Norte and
Humboldt Counties of California, and in southwestern British Columbia. In British Columbia, it is established
and common on southern Vancouver Island, particularly near Saanich and Sidney, as well as on the Lower
Mainland and north to the Queen Charlotte Islands. It is a vigorous vine, with glabrous to sometimes pubescent
twigs. The young shoots are reddish-purple, especially in cultivated forms. The leaves are oval to obovate with
an obtuse apex and a rounded or broadly tapered base, 4-9 cm long and 2.5-5 cm broad. They are glaucous and
usually somewhat downy beneath. The upper pairs of leaves are sessile and united at the base around the stem,
although sometimes all the leaves are free. The inflorescence is a terminal group of dense spikes, each containing
many flowers, and there are also sometimes two or three axillary heads. The flowers are yellowish-white or
creamy-white tinged with purplish-red, becoming a deeper yellow with age, 3-5 cm long, and fragrant. The
corolla is conspicuously bilabiate with the tube and lobes nearly equal in size. The fruit is red, and the berries are
never united. There are several varieties and cultivars of L. etrusca, the variety pubescens being the form usually
found in cultivation. The cv. Superba is a very vigorous form, with panicles larger than the species and reddish-
purple shoots. The species is hardy to USDA Zone 7, and revels in full sun. It is not often seen in gardens in
North America, but probably should be grown more widely as a garden and trellis plant. It is perhaps the most
striking of all Honeysuckles when growing well, with long shoots that branch and form immense bouquets of
flowers. It is probably best grown in the drier areas of the region. 2n = 18.
Lonicera x notha Zabel, Tartary Hybrid Honeysuckle, is actually a group of hybrids raised from seed of a garden cross between L. ruprechtiana and L. tatarica, and closely resembling L. ruprechtiana. It has become
naturalized in some areas, and has been reported as an adventive from the Boreal Spruce Zone of British
Columbia. It is a deciduous shrub, with ovate to oblong leaves. The leaves are pointed at the apex and tapered at
the base, 3.5-10 cm long, and more or less downy on both sides. The paired flowers are in shades of pink, turning
yellowish with age, 1.9 cm long, and not fragrant. The fruit is bright red and rather transparent. There are a
number of cultivars of this species: cv. Grandiflora with large rose-tinted flowers; cv. Carneorosea (or
Carneo-Rosea), which has the deepest rose-colored flowers of all forms; cv. Alba; cv. Ochroleuca; and cv.
Carnea. The two cultivars cv. Grandiflora and cv. Carneorosea are among the best of the numerous hybrid bush
honeysuckles, with attractive fruits as well as flowers and habit.
Origin of the Name
The Family name Caprifoliaceae is derived from Caprea, a she-goat, which has twisted horns in certain
breeds. These horns are suggested by the twisting stems of the climbing Honeysuckles. The generic name
Lonicera commemorates Adam Lonitzer (1528-1586), a German physician and naturalist who wrote a herbal
(Kreuterbuch) that was reprinted many times between 1557 and 1783. The specific name caerulea means 'dark
blue' or 'blue', referring especially to the shade of deep blue seen at midday in the Mediterranean sky. The origin
of the subspecific cauriana seems obscure, although it may be derived from the Latin counts, 'northwest' or
'northwest wind' and the suffix -iana, 'belonging to', referring to the geographical distribution of the plant. The
specific name ciliosa means 'fringed' or 'ciliate', referring to the ciliate hairs of the leaf margin. The specific
epithet dioica, 'dioecious' or 'coming from two', means that the male and female flowers are on separate plants.
This meaning, however, is obscure in L. dioica, which is not dioecious. The subspecific glaucescens means
'having some bloom', referring to the glaucous underside of the leaves. The specific epithet hispidula refers to
the stiff and rigid hairs on the plant. The specific name utahensis commemorates the fact that the plant was first
described from a specimen collected in Utah.
The type locality of Lonicera caerulea var. cauriana is "Alpine meadows, Mt. Paddo [Adams]" in Washington, where it was collected by Wilhelm Nikolaus Suksdorf (1850-1932). The type locality ofL. ciliosa is "On the
banks of the Kooskoosky" where it was collected by Meriwether Lewis. It was first introduced to cultivation in
Great Britain in 1824, and was awarded an Award of Merit by The Royal Horticultural Society in 1919. The type locality of L. dioica var. glaucescens is obscure, as several specimens are cited. It was first cultivated in Great
Britain in 1890, while L. dioica itself was first cultivated in 1636. The type locality of L. hispidula is "woods of
North West America". It is believed to have been described from garden plants grown from seed collected in
northwestern America by David Douglas. The type locality of L. involucrata is "wooded country between 54
and 64 degrees latitude" [Canada], where it was collected by Sir John Richardson (1787-1865). It was first
cultivated in Great Britain about 1824. The type locality of L. utahensis is "Wahsatch Mountains, Utah, in
Cottonwood Canyon; 9,000 feet altitude" where it was collected by Sereno Watson (1826-1892).
The specific name etrusca means 'from Tuscany', an area in northern Italy known as Etruria in classical times.
The type locality is "region of Tuscany, Italy", and it was first cultivated about 1750. The specific name notha
derives from the Greek nothos, 'false', 'baseborn', or 'bastard', referring to its hybrid origin. The plants were
raised from seed in Munich Botanic Garden in about 1878, the seed having been obtained from St. Petersburg
(Leningrad) Botanic Garden in Russia where the cross had occurred.
REFERENCES
Abrams, L. andR. S. Ferris. 1960. Illustrated Flora of the Pacific States. Vol. IV. Bignoniaceae to Compositae. Stanford University Press, Stanford, California. OO
Bean, W. J. 1973. 8th ed. rev. Trees and Shrubs Hardy in the British Isles. Volume 2. D-M. John Murray (Publishers) Ltd.,
London, in collaboration with The Royal Horticultural Society.
Clark, L. J. 1973. Wild Flowers of British Columbia. Gray's Publishing Ltd., Sidney, B.C.
Garman, E. H. 1973. 5th ed. rev. The Trees and Shrubs of British Columbia. Handbook No. 31. British Columbia Provincial
Museum, Victoria, B.C.
Gunther, E. 1945. Ethnobotany of Western Washington. University of Washington Publications in Anthropology 10(l):l-62.
Hillier's Manual of Trees and Shrubs. 1972. Hillier and Sons, Winchester, England.
Hitchcock, C. L. et al. 1959. Vascular Plants of the Pacific Northwest. Part 4. Ericaceae through Campanulaceae. University
of Washington Press, Seattle.
Lyons, C. P. 1965. Rev. ed. Trees, Shrubs and Flowers to Know in British Columbia. J. M. Dent & Sons (Canada) Ltd.,
Vancouver, B.C.
Munz, P. A. and D. D. Keck. 1973. A California Flora and Supplement. University of California Press, Berkeley, California.
Taylor, R. L. and B. MacBryde. 1977. Vascular Plants of British Columbia: A descriptive resource inventory. Technical Bulletin No. 4, The Botanical Garden of The University of British Columbia. University of British Columbia Press, Vancouver,
B.C.
Toms. H. N. W. 1964. Plant Diseases of Southern British Columbia. A Host Index. Reprinted from: Canadian Plant Diseases
Survey 44:143-225. Canada Department of Agriculture, Ottawa.
Turner, N. J. 1975. Food Plants of British Columbia Indians. Part 1. Coastal Peoples. Handbook No. 34. British Columbia
Provincial Museum, Victoria, B.C.
 . 1978. Food Plants of British Columbia Indians. Part 2. Interior Peoples. Handbook No. 36. British Columbia
Provincial Museum, Victoria, B.C.
U.S.D.A., Forest Service.  1974. Seeds of Woody Plants in the United States. U.S.D.A., Forest Service, Agriculture
Handbook 450. Announcement — Change in Subscription Rates
The subscription price of the journal will increase to ten dollars per annum, effective Spring 1980 (Volume
11, Number 1). The cost of single numbers will increase to two dollars and fifty cents, except for special issues.
64
Climatological Summary'
Data                                                     1979
JULY
AUGUST
SEPTEMBER
Average maximum temperature
21.2°C
20.6°C
18.8°C
Average minimum temperature
13.2°C
13.6°C
12.4°C
Highest maximum temperature
28.5°C
23.5°C
27.3°C
Lowest minimum temperature
7.5°C
11.1°C
9.3°C
Lowest grass minimum temperature
5.0°C
3.2°C
5.1°C
Rainfall/no. days with rain
25.8 mm/7
20.0 mm/8
115.5 mm/14
Total rainfall since January 1,1979
375.4 mm
395.4 mm
510.9 mm
Snowfall/no. days with snowfall
0
0
0
Total snowfall since October 1,1978
9.6 cm
9.6 cm
9.6 cm
Hours bright sunshine/possible
290.6/482.5
252.9/439.0
167.7/372.2
Ave. daily sunshine/no. days total overcast
9.4hr/l
8.2hr/2
5.6hr/4
"Site: The University of British Columbia, Vancouver, B.C., Canada V6T1W5
Position: lat. 49° 15'29"N; long. 123° 14 '58" W. Elevation: 104.4 m A successful and happy buyer at the third Friends of the UBC Botanical Garden Plant Sale for Students, which
was held in September during the first week of classes. More than seven thousand house plants were sold during
the three-day event. Pleurotus ostreatus, Oyster Mushroom,
xO.66. The Oyster Mushroom is common
from spring to late fall on the trunks of
dead trees, especially Alder. It is edible
when young.
Volume 10
Number 3
DAVIDSONIA
Fall 1979
Contents
Insectivorous Plants in British Columbia   41
All-America Selection Display Garden   53
The Genus Lonicera in British Columbia   54
Announcement — Change in Subscription Rates   64
Climatology   64

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