"CONTENTdm"@en . "http://resolve.library.ubc.ca/cgi-bin/catsearch?bid=1201090"@en . "University Publications"@en . "2015-07-16"@en . "1979"@en . "https://open.library.ubc.ca/collections/davidsonia/items/1.0115077/source.json"@en . "application/pdf"@en . " DAVIDSONIA\nVOLUME 10 NUMBER 3\nFall 1979 Cover:\nSarracenia purpurea subsp. purpurea, Northern Common Pitcher-plant, occurs in northeastern British Columbia, x 0.37.\nAmanita porphyria, Grey-brown Amanita,\nx 0.66. This Amanita is present in summer to\nlate fall in coniferous forests, especially along\ntrails and in clearings.\nVOLUME 10\nNUMBER 3\nFall 1979\nDavidsonia is published quarterly by The Botanical Garden of The University of British\nColumbia, Vancouver, British Columbia V6T 1W5. Annual subscription, six dollars. Single\nnumbers, one dollar and fifty cents, except for special issues. All information concerning\nsubscriptions should be addressed to the Director of The Botanical Garden. Potential\ncontributors are invited to submit articles and/or illustrative material for review by the\nEditorial Board.\nA cknowledgements\nThe pen and ink illustrations are by Mrs. Lesley Bohm. The photographs on page 53 and\nfacing page 64 were taken by Mrs. Sylvia Taylor. The article on Lonicera was researched by\nMrs. Sylvia Taylor. Editorial and layout assistance was provided by Mrs. Sylvia Taylor and\nMrs. Pam Morgan Robin.\nISSN 0045-9739\nSecond Class Mail Registration Number 3313 Insectivorous Plants in British Columbia\nSYLVIA TAYLOR\nFor centuries people have been fascinated by the unusual adaptations and lifestyles of the insectivorous\n(insect-eating) plants, also often called carnivorous plants. Several species have been cultivated as curiosities for\nover 300 years. Sundews were important in medicine and in mythology in 15th and 16th century Europe. The\nAmerican Indians and early explorers and settlers in North America thought that a plant as strange as the\nPitcher-plant (Sarracenia) must be a possible source of medicine and drugs. Carnivorous plants have excited the\nimagination of many writers \u00E2\u0080\u0094 the man-eating plants of many stories are probably based on a superficial\nknowledge of the small, harmless Venus'-flytrap (Dionaea muscipula) or other species with an active trapping\nmechanism, coupled with a very vivid imagination. Needless to say, there are no scientific facts to substantiate\n\"man-eating plants\". Certainly, there have been reports of one of the tropical Pitcher-plants (Nepenthes\nchelsonii) digesting giant rats \u00E2\u0080\u0094 but even this is difficult to substantiate. The majority of insectivorous plants\ntrap and digest small insects such as flies, ants and mosquitoes, and occasionally small frogs and lizards are\ncaught by the larger pitcher plants.\nThere are at least 450 species (in 13 genera belonging to six families) of insectivorous plants distributed\nthroughout the world. Their basic characteristic is their unique method of survival \u00E2\u0080\u0094 they all lure and trap\ninsects, and have an enzyme system to digest the prey. The plants then absorb nutrients from the digested insects.\nLike all green plants, insectivorous ones can manufacture their own food by photosynthesis utilizing the raw\nmaterials of sunlight, water and carbon dioxide. But they also live in nutrient-poor habitats and supplement\nmineral deficiencies by \"feeding\" on insects, absorbing necessary nitrogen-containing compounds. A 1\nThere is some disagreement among scientists as to the importance of the insectivorous habit. The plants\nusually live in nitrogen-poor soils, and scientists had always assumed that the habit had been evolved to enable\nthe plants to survive in such conditions, where most other plant species cannot grow. However, it is now known\nthat some species can survive quite well without an insect diet, even in nutrient-deficient habitats. It appears that\ninsects provide supplemental nutrients, such as nitrogen and phosphorus, and that these nutrients allow the\nplants to grow faster and produce more flowers and seed than when the supplementary diet is absent. Charles\nDarwin and his son Francis first showed convincingly that Sundews in cultivation were more vigorous when fed\nartificially by supplying insects to the leaves than ones that were not fed (Darwin, 1876). More recent\nexperiments have shown that it is likely that most of the plant's energy and essential organic compounds result\nfrom photosynthesis, and that the majority of minerals used are obtained from the prey. Unfortunately, these\nsuggestions have not yet been definitely proved, and more research is needed. Certainly, the ability to absorb\ncertain essential nutrients from insects enables the plants to survive in nutrient-poor environments, where there is\nlittle competition because of the absence of other plants.\nAll the insectivorous plants trap insects on the outside surface of a modified leaf, and most then secrete\nenzymes onto or into the trap to digest the insect. Some of the simple end products of digestion are absorbed into\nthe 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.\nInsectivorous plants may be divided into three groups based on the method of catching the prey:\u00E2\u0080\u0094\na) Active trap \u00E2\u0080\u0094 typified by the Venus'-flytrap (Dionaea muscipula), which has a fast-acting reflex movement\nmaking it the most animal-like trap. The most common active traps are found in the Bladderworts, which\nhave small, elastic-walled bladders with fast-moving trap doors. The prey is swept into the \"bladder in a\ncurrent of water produced when the walls of the bladder spring apart following the opening of the entrance.\nb) Semi-active trap or Flypaper trap \u00E2\u0080\u0094 utilizes movement as a supplement to an adhesive that captures the prey.\nThis type is found in the Sundews and Butterworts. Glands on the leaf surface secrete adhesive droplets,\nwhich trap insects attracted by odor, color, or refraction of the drops. Tentacles on the leaf then gradually\nmove over the prey, thus enclosing it. 42\nc) Passive trap or Pitfall \u00E2\u0080\u0094 relies entirely on highly specialized modifications of the leaves to trap the victim.\nThe prey is enticed to enter the pitcher of Pitcher-plants such as Sarracenia and Nepenthes by color and odor,\nis then trapped, drowned, and digested in the fluid at the base of the pitcher.\nInsectivorous plants usually grow in special and geographically restricted habitats, such as sphagnum bogs.\nEven one new ditch or one load of fill can change the drainage pattern sufficiently to dry up a small bog, thus\nmaking it unsuitable as a habitat for bog-loving plants. The addition of fertilizer and the control of fires also\nhave adverse effects, the latter by allowing forest vegetation to gradually grow in the bogs. These factors are\ncombining to make insectivorous plants scarce, and possibly even endangered in some areas. In addition, large\nquantities or whole populations are sometimes dug up by plant collectors or commercial dealers. Because of\nthese concerns, several insectivorous plant habitats in the United States have been protected, and plants growing\nin the wild should all be treated as rare and threatened.\nIn British Columbia there are eleven species of insectivorous plants, representing four genera in three families.\nDroseraceae (Sundew Family)\nThe family Droseraceae contains four genera, three of which contain only one species each and are of limited\ndistribution \u00E2\u0080\u0094 Dionaea muscipula, Venus'-flytrap; Drosophyllum lusitanicum, Portuguese Dewy Pine; and\nAldrovanda vesiculosa, an aquatic plant. The fourth genus, Drosera or the Sundews, contains a large number of\nspecies (estimates range from about 85 to 144 species), of which six occur in North America, and two in British\nColumbia.\nSundews are found in every continent of the world, typically inhabiting damp heaths, bogs, swamps, and\nother wet habitats with acid soil (preferably with a pH of 3.5-4.0). It is a particularly hardy genus, and these\nplants are among the first ones to grow on fire-ravaged forest soils, or other newly disturbed habitats, such as\nroadside ditches and abandoned logging trails. In general, they are among the smallest of the insectivorous\nplants, many being only 2.5-5 cm in diameter and often completely hidden in the surrounding sphagnum moss\nexcept at flowering time. One exception is Drosera regiae of South Africa, which has leaves 61 cm long, and\nwhich is reputed to be capable of catching and digesting small animals.\nThe genus consists of low, stemless, herbaceous perennials with a slender rhizome that produces a small\nnumber of fine fibrous roots each year. The leaves are tiny, flat and solid or thick and clublike, and form a\nrosette on the surface of the ground. The leaves may be round, spoon-shaped, oblong, or filiform (narrow and\nthreadlike). The shape of the leaf is the most important characteristic for distinguishing the North American\nspecies. There is an obvious petiole (leaf stalk), except when the leaf is filiform. The margins and upper surface\nof the leaf are covered with many stalked glands or \"tentacles\" (up to 400 have been counted in Drosera\nrotundifolia, and 200 may be the average number) that secrete a highly viscid, clear mucilage that glistens in the\nsunlight. The glands tend to be short and stubby near the centre of the upper surface, becoming longer towards\nthe margin. In some species, or in some habitats, the glands are bright red, contrasting with the transparent\nglobules. All the species flower in early summer, producing a slender, leafless, upright flower stalk that is\n15-50(-76) cm tall and bears 1-20 erect flowers on the upper part. The flowers are bisexual, regular, 2.5 cm or less\nacross, and open one at a time only in sunshine, and often for only one morning. There are 5 petals, which are\nwhite to pink or deep red, depending on the species. There are 5 to many stamens, arranged in whorls of 5. There\nare 3-5 styles, which are often divided. The fruit is a capsule, and the seeds are very tiny (almost dust-like). The\nseed is mature in 6-8 weeks. Many species are capable of self-fertilization if the flower has not been cross-\nfertilized by the time it closes.\nInsects are attracted to the leaves because of the odor, color, or, perhaps, refractions of the droplets. The\nglands secrete both an adhesive substance and digestive enzymes. The viscid secretion droplets accumulate on\neach gland head and are held there until the gland is touched by the prey, when more secretion is released. When\nan insect lands on the leaf it becomes stuck in the mucilaginous droplets, and its struggles to escape stimulate the\nsecretion of more sticky fluid until the insect may be completely covered. At the same time, the stalked glands\nbegin to bend inward over the insect, soon followed by curling of the longer outer \"tentacles\". In a matter of\ntime (maybe several hours) the insect is completely caught within a sticky web of \"bars\". Most insects die within\nabout 15 minutes of capture of suffocation caused by the mucilage blocking the respiratory holes or trachea\nalong the abdomen. The glands now secrete digestive enzymes, which decompose the insect. It has been shown\nthat Sundews contain the enzymes peroxidase, estarase, acid-phosphatase, and protease in the digestive fluid.\nAfter about a week the \"tentacles\" unfurl, and the leaf is able to catch more prey. It has been shown that each\n\"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.\nTheir function is not known, but there are two theories \u00E2\u0080\u0094 one is that they function in the absorption of the\ndigestive products, and the other is that they are concerned with the movement of the larger glands.\nThere are two theories concerning the movement of the stalked glands. The first proports that bending is\ncaused by uneven growth of the stalk (Shetler, 1974b) \u00E2\u0080\u0094 an acceleration of growth begins near the base on the\noutside of the stalk and moves upward towards the gland at the top. This means that the outside of the stalk\ngrows faster than the inside, so that it bends inwards. Unbending is caused by accelerated growth on the inside of\nthe stalk. The second theory states that bending results from a loss of turgor (rigidity) in groups of cells along the\nside of the stalk closest to the stimulus, that is, on the inside of the stalk. In this theory, the stalkless glands are\nthought to be responsible for the withdrawal of fluid from the cells, which causes the loss of turgor. Unbending\nis caused by the cells regaining turgor (Heslop-Harrison, 1978).\nThe Sundews are restricted to catching only small or very weak prey \u00E2\u0080\u0094 flies and even ants are capable of\nescaping from the sticky fluid or from between the \"tentacles\". Even so, they may catch large numbers of\ninsects \u00E2\u0080\u0094 an English naturalist once estimated that plants covering 1 to 2 hectares of land killed about 6,000,000\ninsects in one season (Savage and Savage, 1979).\nSorcerers and alchemists both held Sundews in high esteem, because of the \"dew\". Until relatively recently,\nmany ordinary people in Europe believed that a single plant brought into the house would cause \"pernicious\nfever\", and that a person who searched for a Sundew on Midsummer's Eve and rubbed the leaves over his skin\nwould become indefatigable. An accidental discovery, however, was of no use! Sundews were particularly\neffective when collected at midnight on Midsummer's Eve \u00E2\u0080\u0094 but one had to walk backwards all the time to\navoid 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\nplumbagin, which is present in the British Columbian species.\nThe genus name is derived from the Greek droseros, 'dewy' or 'moist with dew', because the drops of sticky\nmucilage glisten like dewdrops in the sun.\n\"To return to our native bog-plants; the Sundews, all very much\nalike, and all, therefore, to be treated of under Drosera rotundifolia, are pretty and interesting, though not brilliant. Evil little\nthings they are, with their carnivorous habit. One wonders what\ncrime the past lives of Drosera can have held, that now the race\nshould be compelled to dree so ominous and unpleasant a weird\nof murder and fraud. When will the Sundews be free of the\nburden, through some self-sacrificing individual plant who shall\nstarve to death rather than take life, and so redeem his race into\nthe happier paths of peace and virtue?\"\n(Farrer, 1908, p. 245)\nDrosera anglica Hudson Great Sundew\nThe Great Sundew is circumboreal, being found in swamps and bogs in Europe, Japan and North America. In\nBritish Columbia it can be found in suitable bogs and swamps throughout the Province, sometimes occurring in\ndense colonies that cover many acres. This is the most common species at moderate to high elevations in British\nColumbia.\nThere is considerable variation in the shape of the leaf blade from narrowly ovate to spatulate-oblong or\ncuneate-obovate. The leaves are 10-30 mm long, (2-)3-5(-7) mm broad, gradually narrowing to the petiole, and\noften standing erect above the moss. The petiole is glabrous or sparingly glandular-hairy, and 1.5-8 cm long. The\nscape (flower stalk) appears in June to August and is 6-18 cm tall, bearing 2-7 whitish flowers in a racemose\ninflorescence at the top. The calyx is (4-)5-6 mm long, connate for about one-third of its length, and the lobes are\novate-oblong. The petals are oblong and slightly to considerably longer than the calyx. There are (4)5 stamens,\nand (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\nloose, longitudinally striated, and prolonged but not flattened at each end of the seed.\nDrosera 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\nleaves have been shown to be sterile hybrids between D. anglica and D. rotundifolia.\nThe specific name anglica is derived from Anglia, meaning England.\n43 WJ M*% *^*sl:^&*m*\u.\nV * \\ny *^ ^*\"\\n^*^ll^*^!lll Jr '^\n^**^SaL 'j \"\u00E2\u0080\u00A2r^VSe\n44\nFIGURE 1. Drosera rotundifolia, Round-leaved Sundew,\nas seen from above, X 4.8.\nDrosera rotundifolia Linnaeus (Figure 1) Round-leaved Sundew\nThe Round-leaved Sundew is also a circumboreal species, being found in sphagnum bogs and swamps\nthroughout the Northern Hemisphere. It is present in suitable habitats throughout British Columbia, and may\nalso be found growing on old floating logs in stagnant lakes, or among the mosses in marshy ground around\npools. This is the predominant North American species, being found in most areas except the extreme south and\nmost of the Rocky Mountain and Great Plains regions.\nThe leaves are spreading, often hidden in the moss, and may be bright red if the plant is growing in full\nsunlight. The rosette of leaves is 6-8 cm in diameter. The leaf blades are round to broadly obovate-cuneate,\n(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\nto early September, and is up to 25 cm tall, bearing (l-)3-10(-20) whitish flowers. In some parts of the range the\nflowers may be pink or even yellow, mauve or purple. The flowers are very similar to those of the Great Sundew,\nexcept that there are usually only 3 styles and they are bifid nearly to the base. The seeds are about 1.5 mm long,\nand the testa is loose, longitudinally striated, and prolonged and distinctly flattened at each end of the seed.\nIn winter the rosette of leaves withers, and winter buds, which withstand freezing, are formed on the\nrhizome. Axillary buds, which also form on the rhizome, create secondary rosettes below the main one. As the\nrhizome decays, these separate and become new plants.\nThe specific name rotundifolia means 'with round leaves'. A potion made from the leaves of D. rotundifolia\nhas been used to cure coughs, whooping cough, and respiratory diseases such as asthma. The fresh juice from\nthe leaves has also been used in various parts of the world as a cure for corns, calluses and warts.\nBoth native species are easy to grow in live sphagnum moss, preferably the small-growing type that tends to\nform tufts. The moss should be kept constantly moist, but not sodden, and there should be high humidity. These\nplants should receive about 40-50% sunlight all day, and the temperature should be between 18-30\u00C2\u00B0C during the\ngrowing season. All Sundews need a dormant period, with reduced light and a temperature of -4 to 4\u00C2\u00B0C. Forcing\nthe end of dormancy is not successful as the plants usually rot and die. Propagation is by seeds, or vegetatively\nfrom young and vigorous leaves or offshoots. Lentibulariaceae (Butterwort Family)\nThe family is usually divided into five genera containing about 500 species between them. Four of the genera\nare small and fairly homogenous; the fifth, Utricularia or the Bladderworts, is very complex and variable, and\nincludes an estimated 150 to over 300 species. Two of the genera in this family, Pinguicula (Butterwort) and\nUtricularia (Bladderwort), occur in British Columbia.\nPinguicula Butterwort\nThere are between 30 and 46 species of Butterwort, widely distributed throughout the Northern Hemisphere\nand in the South American Andes. They are usually found in wet places, often with mosses, such as on wet soil\nor among rocks along shores and streambanks; on hummocks in swamps; or in low, wet, sandy pinelands.\nMost of the species are very similar in appearance. They are fibrous-rooted perennial herbs with a compact\nbasal rosette of leaves that lies flat on the ground. The leaves are oval or tongue-shaped, pale yellowish-green,\nand the margins curl inwards to form a shallow bowl shape. The flowers are solitary at the ends of erect bractless\nscapes, and are somewhat reminiscent of the flowers of the genus Viola. The calyx is 5-lobed, and the lobes are\npartly connate to form an upper lip of 3 lobes and a lower lip of 2 lobes. The corolla is usually white or light to\ndark violet, or occasionally bright yellow, bilabiate with an upper lip of 2 lobes and a lower lip of 3 lobes, and is\nprolonged into a basal spur. The fruit is a 2-valved capsule.\nThe insectivorous adaptations are not as obvious in this genus as in other genera. The only unusual feature is\nthe greasy appearance of the leaves. The upper surface of the leaf is covered with large stalked glands that secrete\na film of colorless sticky mucilage over the leaf. There are also stalkless glands, raised slightly above the leaf\nsurface and appearing rather wart-like, which secrete digestive enzymes. An insect landing on the leaf causes\npressure, which results in the slow inrolling of the margin, thus pushing the insect towards the centre of the leaf,\nand also stimulating the secretion of mucilage. A small insect is soon fast in the sticky secretion, and struggling\nonly causes it to sink deeper into the fluid. The stalkless glands secrete the digestive enzymes only if nitrogenous\ncompounds are present, thus a grain of sand blown on to the leaf may cause the inrolling of the margin but there\nwill be no secretion of enzymes. The digestive fluid is even more viscous than the mucilage, and contains the\nenzymes ribonuclease, esterase, acid-phosphatase, amylase, and protease. Digestion is usually complete in about\n24 hours, and the leaf slowly uncurls to re-set the trap. The hard indigestible remains of the insect dry out and\neventually may be blown away by the wind. The Butterworts usually capture small insects, such as mosquitoes 45\nand gnats, which are attracted by a fungus-like or decaying odor.\nThe generic name is the diminutive of the Latin pinguis, 'fat' or 'fatty', referring to the greasy appearance of\nthe leaves, and ula 'little one'. The common name Butterwort also refers to the greasy appearance \u00E2\u0080\u0094 wort is the\nold Middle English word for 'plant'. The leaves of Butterwort were used in Scandinavia as a medicine, and were\nrubbed on open sores of animals to make them heal faster. In fact, the digestive fluid does contain an antibiotic\nto stop the decaying of the insects. Butterwort was also commonly used as a milk coagulant in home dairy\nrecipes.\nPinguicula villosa Linnaeus Hairy Butterwort\nThe Hairy Butterwort is found in many northern sphagnum bogs and swamps in North America and in\nEurasia, but it is usually so inconspicuous that it is rarely collected. In British Columbia it has been collected on\nthe Queen Charlotte Islands, at one station on Vancouver Island, and also apparently near Prince Rupert.\nThe leaves are 0.4-1.5 cm long, 0.2-0.7 cm broad, glabrous below and villous above. The scape is produced in\nJune to August and is 2-6(-12) cm tall and villous with capitate hairs, with a single nodding lavender-blue flower.\nThe 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\nand erect.\nThe specific name means 'covered with soft hairs', referring to the hairs on the leaf surface and on the scape.\nPinguicula vulgaris Linnaeus Common Butterwort\nThe common Butterwort is the most widespread species in the genus, being found throughout the arctic and\nsubarctic regions and up to 2300 m in the mountains of Eurasia, northernmost Africa, Iceland, Greenland and\nNorth America. However, it is seldom abundant in its preferred habitats of bogs, mossy seeps, and wet soil on\nbeaches, meadows, or even on bare soil. In British Columbia it is found at relatively high elevations on\nVancouver Island and in the Coastal Mountains, and in valleys in the northern part of the Province.\nThe leaves are succulent, elliptic or oblanceolate, entire, 2-5(-6) cm long, and 7-18 mm wide. The petioles are\nshort and winged. The scape appears in (May) July to August, is 5-15 cm tall, and is obscurely glandular. The\ncalyx of the flower is 3-5 mm long. The corolla is deep violet-purple to lavender-blue or, rarely, nearly white. It is 46\n(12-) 15-25 mm long including the slender spur (which is\nabout 10 mm long), and has a flaring throat and broad,\nrounded lobes. The capsule is 4-6 mm long and is erect.\nThe species is represented by two subspecies in British\nColumbia. Pinguicula vulgaris subsp. macroceras (Link)\nCalder & Taylor (Figure 2) has flowers 24 mm long with\nlarge broad, rounded lobes that are suddenly narrowed to\nform a long filiform blunt spur up to 32 mm long. The lower\ncorolla lobes are oblong-obovate. This subspecies is found in\nvery different habitats ranging from sunny moist slopes to\nglacier moraines and dripping rocks at elevations between\n50-2300 m in the north Pacific coastal area, including Japan,\nthe Kamchatka Peninsula, the Aleutian Islands and Alaska.\nIn British Columbia it is present across the southern part of\nthe Province to the Rocky Mountains, but becomes almost\npurely coastal (except for a few stations along the Skeena\nRiver) in the northern parts. Pinguicula vulgaris subsp.\nvulgaris is distinguished by the lower corolla lobes, which are\noblong rather than oblong-obovate.\nThe specific name vulgaris means 'common', 'general' or\n'ordinary', while the subspecific macroceras is derived from\ntwo Greek words, macro and ceras, meaning 'large horn',\nreferring to the large spur.\nBoth species of Pinguicula can be cultivated, preferring\nmoderate or cool summer temperatures with a relative\nhumidity of 60-100%. The Hairy Butterwort is probably best\ngrown in small compact-tufted species of living sphagnum\nmoss, but the Common Butterwort may be grown in a\nmixture of Vi black peat, Vi sand and lA perlite. In\ncultivation, all Butterworts are subject to attack by snails,\naphids and the fungus Ustilago pinguicula. This fungus\ncauses a smut and produces dark brown or black masses of\nspores in the anthers. The disease is present in wild\npopulations of the Common Butterwort in British Columbia. Propagation is by seed, or vegetatively from young and\nvigorous leaves. Butterworts do not like to have their root\nsystems disturbed, so are difficult to transplant.\nUtricularia\nBladderwort\nMost species of Bladderwort are aquatic (either anchored in\nopen water or free-floating), some are terrestrial on wet or\nmoist soil, and a few are epiphytic in moss. One or two\nspecies are known that are restricted to the pools of water\nfound at the base of the leaves of large bromeliads in South\nAmerica. Representatives of the genus are found on all\ncontinents of the world, with about 12 species occurring in\nNorth America and five in British Columbia.\nAll species lack true roots and show little distinction\nbetween the stem and leaves. In the aquatic species, the\nleaves are submerged and very finely divided. All our native\nspecies have alternate leaves, but in other species the leaves\nmay be partly or wholly whorled. There are bearded, hollow\nbladders among the leaves or growing from specialized\nstalks. The scape is tall and slender, and bears a loose raceme\nof yellow flowers, each of which is subtended by a bract. The\nflowers are strongly bilabiate, the lower lip is either entire or\n3-lobed, and the upper lip is either entire or shallowly\nFIGURE 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\ncompletely closes the opening to the corolla tube. The base of the corolla tube is extended backward and\ndownward to form a spur or sac. The fruit is a capsule. Most species form winter buds at the ends of the stems in\nthe fall, but some do not. These buds fall off and lie dormant at the bottom of the pond during the winter, ready\nto produce new plants in the spring.\nThe bladders are modified leaves and there are many variations in form, with the smallest known being\n0.3 mm and the largest about 6 mm in diameter. The best known one is that of the Greater Bladderwort\n(Utricularia vulgaris). This is slightly less than 6 mm across, flattened, pear-shaped, and is attached to the plant\nby a short stalk at one side. There is an opening at the narrow (free) end protected by a \"door\", which is\nattached at the top and hangs downward over the opening, swinging free at the bottom where it rests against a\nsemi-circular collar or threshold. At the edge of the opening are a pair of branched antennae plus a number of\nlong slender bristles along the side. These form a funnel to guide the prey into the opening of the bladder. There\nare (1)4 stiff bristles near the lower free edge that constitute the tripping or trigger mechanism. Scattered over the\noutside of the bladder walls and on the threshold of the door are numerous stalkless glands, which secrete a\nmucilaginous substance. On the outer surface of the door are many stalked glands, which also secrete mucilage\nplus a sugary substance that is believed to attract the prey. The inside walls of the bladder are covered with many\n2- to 4-headed glandular hairs.\nWhen the trap is set, the door is closed, the trigger hairs are in position, the bladder is almost empty of water,\nand the walls are straight or slightly concave. If a passing insect touches the trigger hairs, their movement alone\ndistorts the bottom edge of the door causing it to move away from the threshold. The vacuum inside the bladder\ncauses a suction so that water rushes into the bladder, carrying with it any insect or animal small enough to pass\nthrough the opening. The bladder is now full of water and the walls bulge outwards. Specialized cells in the walls\nof the bladder are concerned in the active transport of ions from inside the bladder to the water outside. This sets\nup an osmotic gradient that results in water being moved out of the bladder, thus re-setting the trap. The\nre-setting takes only about 30 minutes, although the prey may remain alive inside the trap for several days before\neventually dying. The mechanism of digestion is uncertain, but the insect quickly disintegrates after death,\ntherefore, there seems to be some kind of digestive fluid secreted into the bladder. In addition, there are reports\nthat the cells of the bladder become a red-brown color, suggesting some sort of metabolic activity. It has also\nbeen found that the small pool of water that remains at the bottom of the bladder does contain the enzymes 4 /\nesterase, acid-phosphatase, and protease. The indigestible, hard remains of the insect stay in the bladder.\nThe movement and trigger mechanism of Utricularia was the subject of an early movie film produced by Dr.\nF. E. Lloyd of McGill University in the 1930's. Dr. Lloyd, who was interested in the physiological aspects of the\nplants, also wrote one of the first comprehensive books on carnivorous plants (Lloyd, 1942).\nBladderworts can capture a wide range of organisms, including insects and insect larvae, diatoms, desmids,\nprotozoans, and small crustaceans such as Daphnea and Cyclops. They can also catch the occasional tadpole or\nnewly hatched fish, provided these are very tiny. Occasionally, bladders are found that have half a tadpole\nsticking out through the door \u00E2\u0080\u0094 obviously the victim was too large!\nThe generic name is derived from the Latin utriculus, 'little bag' or 'little bottle', referring to the bladders.\nUtricularia gibba Linnaeus Humped Bladderwort\nThe Humped Bladderwort is found mainly in Central and South America and Africa, but is also present in the\neastern and western parts of North America. In British Columbia it is found on Vancouver Island, being\ncommon in several lakes near Victoria and in Ash, Patterson and Turtle Lakes near Port Alberni, and on the\nmainland in Beaver Lake in Vancouver.\nThe stolons (stems) are 10-25 cm long and are often a characteristic zig-zag shape when floating. The leaves\nare 3-10 mm long, filiform, and dichotomously branched from the base. One branch is usually straight and\nwithout bladders, while the other is usually further branched and has 1-3 bladders. The bladders are 0.5-1.5 mm\nlong, 0.5-1.0 mm wide, and have long branched hairs at the membrane opening. There are no winter buds. The\nscape is 3-7 cm tall and there is either 1 or 0 scale. The inflorescence contains 1-2 bright yellow flowers, each\nsubtended by a semi-amplexicaulous bract. The pedicel is 5-10 mm long and is straight after anthesis. The calyx\nlobes 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\nangles 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\ncylindrical, 3-5 mm long, straight and parallel to the lower lip. The capsule is about 5 mm in diameter.\nThe species is represented in British Columbia by the subspecies gibba, the typical form.\nThe specific name gibba means 'swollen on one side'. Utricularia intermedia Hayne in Schrader Flat-leaved Bladderwort\nFlat-leaved Bladderwort is a circumpolar boreal species, but is absent from Iceland, and is usually free-\nfloating in shallow standing or slowly moving water. In North America it is present as far south as California,\nIndiana and Delaware. It is of scattered distribution in British Columbia, where it is often associated with Carex\nlasiocarpa. Sometimes it grows in the sub-littoral zone of pools, anchored in the mud and forming large mats.\nThe stolons are very slender and are of two kinds:\u00E2\u0080\u0094 leafy stolons 10-50 cm long, which lack bladders; and\nleafless ones with bladders. The leaves are 5-30 mm long, and are divided at the base into 3 branches, which then\ndichotomously branch into 6-20 slender segments. The segments are flat, have a central nerve, a blunt obtuse tip\nwith an abruptly starting bristle, and are all about the same width. The bladders are 1.5-4.5 mm long and 2-4 mm\nwide. The winter buds are (2-)5-7(-15) mm long, 3-10 mm wide, and ovoid or ellipsoid. The scape appears in July\nto August, and is 5-20 cm tall with one or two bract-like scales. The inflorescence has 2-4(-5) bright yellow\nflowers, each subtended by a bract that is 1.5-4 mm long. The pedicel is 3-15 mm long. The calyx lobes are\nsubequal, 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\n8-12(-18) mm long and 7-20 mm wide. The palate is well developed and about 7 mm long, and the spur is\ncylindrical, straight, 8-12 mm long, and is at an acute angle to the lower lip. The capsule is about 3 mm in\ndiameter.\nThe specific name means 'intermediate', usually referring to the color, form or habit of the plant. This\nparticular species is intermediate in size between U. minor and U. vulgaris.\nUtricularia minor Linnaeus Lesser Bladderwort\nLesser Bladderwort is a circumpolar species with small discontinuities in the distribution in Asia and America,\nand is usually found in shallow standing or slowly moving water. In North America it extends as far south as\nCalifornia, 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\ncommon than the records indicate. In British Columbia it can be found in and around peaty bogs and swamps,\ncommonly occurring in small-channelled drainage areas, along and under the margins of these channels.\nThe stolons are 15-75 cm long and are differentiated into leafy ones with few bladders, and ones with fewer\nleaves 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\nacuminate. 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\ndiameter. The scape appears in June to September and is 4-15 cm tall with 1-4 bract-like purple scales. The\ninflorescence contains 2-10 pale yellow flowers, each subtended by a purple auriculate bract that is 1-2 mm long.\nThe pedicel is 2-10 mm long, and is recurved after anthesis and in fruit. The calyx lobes are equal, 0.5-2.5 mm\nlong, 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\nlower 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\nsmall, 1-2 mm long, saccate, and poorly developed. The capsule is globose and about 2-2.5 mm in diameter.\nA more robust form, which has the stolons 50-120 cm long, has been collected from several places in British\nColumbia. The leaves of this form are 9-20 mm long, orbiculate, and divided into 20-40 segments.\nThe specific name minor means 'smaller', 'inferior', or 'less', referring to the habit of the plant. It is rarely\nseen in flower in British Columbia.\nUtricularia ochroleuca R. W. Hartman\nThis species is found in northern, central and western Europe, but is considered rare there, and in North\nAmerica from the Northwest Territories and Alaska east to Nova Scotia, and south to Washington, Oregon,\nColorado and Illinois. Its distribution in British Columbia is poorly known.\nThe stolons are 5-30 cm long and are differentiated into leafy and leafless ones. The leaves are 3-15 mm long,\nand are divided into 3 parts at the base, these then further divide to form 5-19 segments. The segments have a\ncentral nerve, and are gradually narrowed into a bristle at the apex. The leaves occasionally bear one bladder.\nThe bladders usually form on the leafless Stolons, and are about 1.5-2.5 mm long, and 1-2 mm wide. The winter\nbuds 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,\nand has 1-5 bract-like scales. The inflorescence contains 2-10 lemon yellow or pale yellow flowers, each of which\nis subtended by an auriculate bract about 0.5-4 mm long. The pedicel is 2-15 mm long, and is straight in anthesis\nbut becomes slightly recurved afterwards. The calyx lobes are subequal, about 2.5-3.5 mm long, 1.5-2 mm wide,\nand 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\nand a little narrower than the upper lip. The palate is about 5 mm long, and the spur is 3-5.5 mm long, pyramidal\nwith 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\neasily referrable to this species because of the pyramidal spur at right angles to the lower lip, a characteristic that\nis found in no other species in the area.\nThe specific name is derived from two Greek words \u00E2\u0080\u0094 ochros, 'pale' or 'pale yellow', and leucon, 'white' \u00E2\u0080\u0094\nmeaning 'yellowish-white', referring to the color of the flowers.\nFIGURE 3. Utricularia vulgaris, Greater Bladderwort.\nA. Stolon and bladders, X 1, B. bladder, x 5.\n49 Utricularia vulgaris Linnaeus (Figure 3) Greater Bladderwort\nGreater Bladderwort is a circumpolar boreal species, with a break in the distribution in Iceland and\nGreenland. 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.\nThe stolons are 20-180 cm long, and rather coarser than in the other native species. The leaves are numerous,\nl-5(-9) cm long, pinnatifid, and divided into 20-150 filiform segments. The segments are terete and progressively\nmore slender after each division so that the ultimate ones are filiform and strongly acuminate. The bladders are\n1-4 mm long, 1-3 mm wide and are attached near the base of the points of branching in the leaves by stalks\n0.1-1.0 mm long. The bladders on the primary branches are larger than on other branches. There may be as\nmany as 10-50 bladders per leaf. The winter buds are ovoid or ellipsoid, 7-20(-30) mm long, and 6-15 mm wide.\nThe scape appears in May to August, and is stout, 6-30(-40) cm tall, and has 1-5 bract-like scales. The\ninflorescence contains (3-)6-20 bright yellow flowers subtended by bracts 2.5-8 mm long. The pedicel is 6-30 mm\nlong, and is recurved after anthesis and in fruit. The calyx lobes are subequal, 3-6 mm long, 2-4 mm wide, and\nthe 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\nslightly lobed. The palate is gibbous, as large as the upper lip, and sometimes has reddish-brown veins on it. The\nspur is 10 mm long, well-developed, falcate and directed forward. The capsule is globose, and about 6 mm in\ndiameter.\nThe American plants tend to have the spur somewhat more slender and pointed than in the Eurasian forms,\nand have been separated out as U. vulgaris subsp. macrorhiza (Le Conte) R. T. Clausen. This is the form that is\npresent in British Columbia.\nThe specific name vulgaris means 'common', 'general' or 'ordinary', and the subspecific macrorhiza is\nderived from two Greek words \u00E2\u0080\u0094 macro, 'large', and rhiza, 'root' \u00E2\u0080\u0094 meaning 'large-rooted'.\nThe aquatic species of Utricularia can be propagated from seeds in water, with plants reaching flowering size\nin one to two years. They can also be propagated vegetatively very easily by taking stem cuttings.\n\u00C2\u00A33 O Sarraceniaceae (Pitcher-plant Family)\nThis is a small family of low perennial herbs inhabiting bogs and wet savannahs, and having pitcher-like leaves\nfor trapping insects and other tiny animals. It contains about seventeen species in three genera, all endemic to\nthree widely separated regions of the New World. Heliamphora contains 4-6 species, which are found only on\nisolated mountains of the Guiana Highlands of northern South America. Darlingtonia californica, California\nPitcher-plant or Cobra-plant, is the sole member of its genus, and is highly localized in montane meadows,\nbetween 90-1830 m elevation, of northern California and adjacent Oregon. The third genus, Sarracenia,\ncontains between 8 and 12 species, all but Sarracenia purpurea being found only in the southeastern states of\nAmerica.\nSarracenia species grow only in very wet habitats, in waterlogged soil or even in standing water. Bogs are an\nespecially typical habitat. They are commonly found growing in acid soil, but this seems to be mainly because of\nthe lack of competition \u00E2\u0080\u0094 specimens of S. purpurea have been found growing in humus derived from sphagnum\nmoss 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\nplants to be discovered, although not recognized as such at the time. The first drawing of S. purpurea was done\nin Europe in 1601, and was reproduced in the second edition (1636) of John Gerard's Herball. This was one of\nthe first botanical books in English (instead of Latin) when it was originally published in 1597 as \"The Herball\nor General Historie of Plantes\".\nThe genus consists of herbaceous perennials with round, horizontal to vertical rhizomes. The rhizome is\n8-30 cm long when mature, and may survive for 20-30 years. The leaves are in a basal rosette and appear in two\nor sometimes three forms each year. The most obvious are the pitchers, modified entire leaves that are produced\nin the spring, with a second crop sometimes appearing in the fall, although these are often different in shape.\nSecondly, half or all of the leaves in the fall (depending on the species) may be sword-like and tubeless, forming\nthe overwintering leaves or phyllodia. Finally, all species produce small scale-like leaves. The pitchers show great\nvariation between species, but in general either lie flat on the ground or arch upward, are 30-60 (-122) cm long,\nand are strikingly colored, varying from yellowish-green to dark green to purplish and mottled with red to dark\nred (the intensity of the red depending partly on the amount of sunlight in the habitat or the season). They are\nfleshy, and a prominent wing runs along the outer surface of the leaf from top to bottom on the side facing the\ncentre 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\nopening, thus forming a protection against rainwater entering the pitcher. The flowers appear in the spring to\nearly summer, and are large and showy with a more or less agreeable scent. There is a solitary leafless scape,\n30-60(-122) cm tall (usually about as long as the leaves), which bears a single, nodding flower at its apex. There\nare usually 5 (occasionally 4) sepals that are green or sometimes colored and petal-like, broad and spreading, and\npersistent. The five petals are free, large, yellow to greenish-yellow or dark red, oblong to obovate, or they may\nbe very short and almost absent. They hang straight down and are incurved. There are twelve to many free\nstamens. The base of the style is slender, but it is expanded at the top to form a very broad umbrella-like body\nwith 5 angles and 5 rays. There is a small hooked stigma at the end of each ray. The ovary is subglobose and\nthere are numerous ovules. The fruit is a 5-valved capsule with a granular surface. The seeds are numerous,\nsmall, plump, and tan to dusty lavender in color.\nThe petals are persistent for several weeks and then drop, but the remainder of the flower persists until the\nseeds are mature in July to September. The umbrella-shaped style is believed to catch pollen released by the\nanthers, holding it where insects can easily find it. Only a minority of plants in a population seem to flower in\nany one year. The species hybridize freely where their natural ranges overlap or in cultivation, and there are a\nlarge number of natural hybrids, the most common probably being Sarracenia x catesbaei Elliott (S. purpurea\nsubsp. venosa x S. flava).\nThe prey are attracted to the pitcher by the flower-like appearance and odors. There are nectar-secreting\nglands extending from the lip downwards on the outside. Insects land on the surface and gradually move into the\nmouth of the pitcher as they feed on this nectar. The lip of the pitcher has stiff hairs that point downward into\nthe trap, and the insect cannot return once it has passed through this area. Just below the lip, in the steepest\nportion of the throat, is a highly polished, slippery area without hairs, thus forcing the insect down into another\nregion of downward pointing hairs and then into a pool of liquid at the bottom of the pitcher. This liquid is a\nmixture of digestive enzymes, rainwater (in some species), and bacteria, which are believed to help in digestion.\nIt is also believed that the liquid may contain a wetting agent to reduce the surface tension so that the trapped\ninsect is unable to crawl across the surface. It has been shown that the enzymes invertase and protease are\npresent. The chemical coniine has recently been shown to be present in very low concentration, sufficient to kill\nany insect before digestion (Dr. G. H. N. Towers, personal communication). This is the same chemical that is\npresent in Poison-hemlock and that was used to kill Socrates. The trapped insects accumulate in the fluid, and \u00C2\u00A3) 1\nthe digestive products are continuously withdrawn as the plant needs them.\nThe usual insects caught are ants, beetles and flies, although small toads, frogs and centipedes may also be\ntrapped. The quantity of prey captured is sometimes quite large \u00E2\u0080\u0094 the pitchers usually live for several months\nand may be almost completely filled with decaying remains. The odors of decay are often evident. Some animals\nhave learned to use the pitchers for food, shelter, or in part of their life-cycle. Small tree frogs often shelter\nduiing the day in the mouths of large pitchers, using the suction pads on their feet to cling to the inside walls.\nThe fly Sarcophaga sarraceniae lays its eggs among the insect remains at the bottom of the pitcher, the larvae\nfeed on these and then bore their way out through the wall to pupate in the soil. It has been discovered that the\nlarvae produce an anti-enzyme as a protection against digestion. Some small mosquitoes also breed within the\npitchers in the wild.\nPitcher-plants have been cultivated as curiosities for many years, and have been used medicinally to a\nlimited extent. The Indians of Newfoundland used the roots of the northern form of the Common Pitcher-plant\nas a medicine for smallpox, and the treatment was also used in England for a short period about 1865. Steam\ndistillation of the roots of S. flava produces sarasin, which was used to relieve tic douloureux, a painful facial\nnerve irritation.\nThe generic name honors Dr. Michel Sarrasin (or Sarrazin) de l'Etang (1659-1734), a botanist and physician at\nthe Court of Quebec, who sent a specimen of Sarracenia purpurea subsp. purpurea to the French botanist\nJoseph Pitton de Tournefort (1656-1708) about 1700.\nSarracenia purpurea Linnaeus Common Pitcher-plant\nThis is the most widespread species, being found in sphagnum bogs from the subarctic to the sub-tropics of\nNorth America. It is now known to occur in all ten provinces of Canada, being found about eleven years ago in\nnortheastern British Columbia (Krajina, 1968).\nThe rhizome is stocky and more or less vertical. The pitchers are more or less suffused with purple or red,\n(5-)10-20 cm long, 1-5 cm wide, obovoid, and curved-ascending from the base. They tend to be smaller and more\nreddish 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\nsubglobose, 5-7 cm wide, and commonly dark purple-red or, rarely, yellowish. The style is greenish-yellow to\nred.\nThere are two subspecies of the Common Pitcher-plant, which integrade in New Jersey. Sarracenia purpurea\nsubsp. purpurea or Northern Common Pitcher-plant occupies the northern part of the range, and is the form\nfound in British Columbia. It is characterized by long narrow pitchers that are usually more than three times as\nlong as broad, and by having smaller hood wings, which extend only slightly beyond the margin of the pitcher.\nThe petals and summit of the style are red. Sarracenia purpurea subsp. venosa is the southern form, which has\nshorter, broader pitchers that are usually less than three times as long as broad.\nThe specific and subspecific epithet purpurea is from the Latin purpureus, 'purple', referring to the color of\nthe flowers and/or leaves. Sarracenia purpurea subsp. purpurea was designated as the Provincial flower of\nNewfoundland in 1954, but was first used as an emblem on the Newfoundland penny about 1876 by order of\nQueen Victoria. A potion of the roots and leaves was used as a cure-all for stomach, diuretic and menstrual\ncomplaints and as a laxative, as well as for smallpox.\nThe Common Pitcher-plant is somewhat difficult to cultivate, and is probably best grown on live sphagnum\nmoss with semi-shade and cool root temperatures during the hot summer months, after it has reached maximum\ndevelopment. Full sun is needed before this for best flower and pitcher formation. High humidity is essential.\nHowever, we are presently growing the northern form successfully in The E. H. Lohbrunner Alpine Garden at\nUBC in a peat bed using a mix of one-third peat moss, one-third coarse sand and one-third loam. The plants are\nin full sun, but are never allowed to dry out in the summer. It has been stated that water from a water softener or\ndeionizer should not be used as the added sodium is detrimental to the plant. Propagation is by seeds, which\nreach flowering size in 3-5 years, or by division of the rhizome.\nInsectivorous Fungi and Seeds\nIn addition to the insectivorous plants, it has been found that some fungi are carnivorous. Arthrobotrys\nobligiospora and Dactylaria brochopaga both trap eelworms in loops of their hyphae, and then ingest needed\nnutrients from the body.\nThere are also indications that the seeds of some non-insectivorous plants may be carnivorous, or capable of\ncarnivory. John T. Barber of New Orleans has found that seeds of the Common Shepherd's-purse (Capsella\nbursa-pastoris) are apparently capable of trapping prey. The seeds are mucilaginous, that is, they release a\ngummy covering or pellicle when they take up water. The mucilage appears to contain a substance that will\nattract motile soil bacteria, nematodes and protozoans. Barber showed that nematodes accumulated in\nstatistically significant numbers around Shepherd's-purse seeds, and that a significant number of them would be\ndead (75% versus 7% in the absence of seeds after 8 days). Therefore, he suggested, and later proved, that the\nseeds released a toxin. He also showed that Shepherd's-purse seeds released proteases on imbibition (uptake of\nwater), and that the proteolytic activity was confined to the mucilage. Experiments using labelled amino-acids\nproved that the germinating seeds were able to take up products of digestion, and that they were incorporated\ninto the seedling. The seeds are small and contain minimum food reserves, and the plants grow in semi-arid soils\nof low fertility. Thus, although the plants do not appear to needprey to germinate, or for the seedlings or plants\nto be healthy, the potential for carnivory seems to be present \u00E2\u0080\u0094 the seeds can attract, trap, kill, digest, and\nabsorb the resultant nutrients.\nREFERENCES\nBarber, J. T. 1978. Capsella bursa-pastoris seeds. Are they \"carnivorous\"? Carnivorous Plant Newsletter 7:39-42.\nCeska, A. and M. A. M. Bell. 1973. Utricularia (Lentibulariaceae) in the Pacific Northwest. Madrono 22:74-84.\nCarnivorous Plant Newsletter, volumes 1-7 (1972-1978).\nDarwin, C. 1876. Insectivorous Plants. John Murray, London.\nEmboden, W. A. 1974. Bizarre Plants. Magical, Monstrous, Mythical. Macmillan Publishing Co., Inc., New York.\nFarrer, R. 1908. Alpines and Bog-Plants. Edward Arnold, London.\nHeslop-Harrison, Y. 1978. Carnivorous Plants. Scientific American 238(2):104-115.\nKrajina, 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.\nSavage, C. and A. Savage. 1979. Canada's carnivorous plants. Nature Canada 8(1):4-12.\nSchwartz, R. 1974. Carnivorous Plants. Edited by Deborah Leavy. Praeger Publ., New York.\nShetler, S. G. 1974a. Sarraceniales. In: Encyclopaedia Brittanica, 15th edition, pp. 252-256.\n . 1974b. Nepenthales. In: Encyclopaedia Brittanica, 15th edition, pp. 958-962.\nTaylor, 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,\nB.C.\nAll-America Selection Display Garden\nThe UBC Botanical Garden had an All-America Selection Display Garden for the first time this summer. In\nthis garden were displayed All-America flower winners from the immediate past and present years. The garden\nwas located in the entrance area to the Main Garden, and was open to the public at all times. Displayed in the\nGarden were plants grown from seed supplied by three seed houses as well as the All-America Selection\nCommittee. The garden will be maintained in future years, with a changing display of flowers each year.\n53\nFIGURE 4. A view of the All-America Selection Display Garden in late September. The Genus Lonicera in British Columbia\nMember of the Family Caprifoliaceae\nLONICERA CAERULEA Linnaeus\nBluefly Honeysuckle\nLONICERA CILIOSA (Pursh) A. P. de Candolle\nWestern Trumpet Honeysuckle\nLONICERA DIOICA Linnaeus\nGlaucous-leaved Honeysuckle\nLONICERA ETRUSCA Santi\nEtruscan Honeysuckle\nLONICERA HISPIDULA (Lindley) D. Douglas ex Torrey & Gray\nHairy Honeysuckle\nLONICERA INVOLUCRATA (J. Richardson) Banks ex K. P. J. Sprengel\nTwinberry Honeysuckle\nLONICERA x NOTHA Zabel\nTartary Hybrid Honeysuckle\nLONICERA UTAHENSIS S. Watson\nUtah Honeysuckle\nDescription of the Genus Lonicera\nLonicera is a genus of vigorous erect shrubs or woody vines. The shrub forms may be bushy or climbing,\ndeciduous to evergreen, and 1.2-3 m tall. The woody vines are sometimes more or less evergreen. All forms have\nthin stems and hollow branchlets.\nThe twigs are rounded and mostly slender. The pith is moderate, pale or brown, and is excavated at the nodes\nin some species. The leaf scars are opposite, crescent -shaped, small, and on the narrowed extremities of raised\nbases that are more or less connected by transverse lines. Bundle traces 3. Stipule scars lacking.\nThe winter buds are often superposed, with the lowermost being the largest or developing into an inflorescence\nin the first season. They are sessile, variously shaped, and have 2 to rather numerous 4-ranked scales.\nThe leaves are opposite, simple, mostly entire, short-petiolate or sessile, and exstipulate or rarely stipulate,\nwhen the stipules are usually small and adnate to the petiole. Few show fall color.\nThe flowers are either in axillary pairs on a slender petiole subtended by 2 bracts and 4 bractlets, or in sessile\nwhorls at the ends of the branches. They are perfect, white or yellow to pink, scarlet or purple, usually small but\nshowy, and often have a fragrance that is most marked towards the evening thus attracting night-flying hawk-\nmoths and hummingbirds. The flowers are subtended by bracts and bractlets, the bractlets being distinct,\nconnate or sometimes wanting. The calyx is small, with the 5 sepals united to form an ovoid to nearly globulose\ntube, with a shallowly 5-lobed limb, the lobes sometimes being obsolete. The corolla is regularly or nearly\nregularly 5-lobed or often evidently bilabiate with a 4-lobed upper lip and a single strap-shaped lower lip. It is\ntubular to funnelform or campanulate, and is often gibbous (swollen) or spurred near the base. Nectar is\nsecreted at the base of the corolla tube. There are 5 stamens, which are inserted on the corolla tube and alternate\nwith the corolla lobes. The anthers are linear or oblong. The style is slender and elongate, and the stigma is\ncapitate. The ovary is inferior and usually 2-3(-5)-celled. The ovules are pendulous, and there are 3-8 per cell.\nThe flowers generally appear in the spring, but in late summer in some species. C X 0.65\nB xl.3\n55\n_i.5m\nFIGURE 5. Lonicera invotucrata var. involucrata. A. Habit, B. flowers, C. fruiting branch, D. fruit and seeds. 56\nThe fruit is a small fleshy berry and may be bright red to bright yellow or orange, blue or black. They are ripe\nin the summer or early fall, and those of paired flowers may be partially or completely united. The fruits are\nattractive to birds. The seeds are few to many depending on the species. They are oval or oblong but appear\nelliptic in longitudinal section, small, and the outer epidermis is often thick, pitted and lignified. There is no data\navailable to show how old plants must be to produce good seed crops. Seed dispersal is primarily by birds and\nanimals.\nThere are more than 150 species of Lonicera distributed over the temperate and subtropical regions of the\nNorthern Hemisphere. About 20 species are native to North America, of which 6 occur naturally in British\nColumbia. Two introduced species may also be found growing wild in certain parts of the Province.\nKey to the species in British Columbia\nLower leaves with well developed, connate, stipule-like appendages at the base of the short petiole; flowers\nin terminal or terminal and axillary inflorescences with several compact whorls; corolla strongly\nbilabiate. Vine or climbing shrub Lonicera hispidula\nLower leaves without stipule-like appendage; flowers in terminal whorls or axillary pairs; corolla weakly or\nstrongly bilabiate.\nCorolla strongly bilabiate.\nEvergreen or half-evergreen vine (occasionally deciduous), 2-4(-9) m long; flowers in terminal\ndense spikes. Adventive species Lonicera etrusca\nDeciduous shrubs; flowers in terminal clusters or axillary pairs.\nFlowers in terminal almost stalkless clusters; bushy shrub, 1.5-6 m tall with twining or\ntrailing stems about 60 cm long Lonicera dioica\nFlowers in axillary pairs; shrubs, 2-3 m tall. Adventive species Lonicerax notha\nCorolla regular or only weakly bilabiate.\nTwining vine with widely branched stems often 5-6(-9) m long; upper 1-3 pairs of leaves on\nflowering branches connate (united around stem) at base; flowers in terminal\ncompact spikes of 1-3 whorls Lonicera ciliosa\nShrubs; upper leaves not connate; flowers in axillary pairs.\nLeaves (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\n' tall Lonicera involucrata\nLeaves less than 9 cm long; flowers not subtended by obvious bracts; shrubs less than\n2 m tall.\nFruit red and often of unequal size with one member of pair not developing;\nleaves 2-5(-8) cm long; ovaries united at base when mature; erect or\nsomewhat straggly shrub, 0.6-2 m tall Lonicera utahensis\nFruit blue or blue-black (occasionally red), both usually of equal size; leaves\n1.2-7.5(-9) cm long; ovaries apparently united but actually separate\nwithin a narrow-mouthed cup formed by the united bractlets;\nsturdy shrub, (0.2-)0.5-l .5(-2) m tall Lonicera caerulea\nThe Native Species\nNatural Distribution and Habitat\nLonicera caerulea is a widespread species with several varieties, and is present throughout many areas of\nnorthern and central Europe, Asia and North America. In North America it is present south of approximately\nlatitude 60\u00C2\u00B0N, 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\nHemlock Zone of the southern part of the Province, extending east to Alberta. The northern limit in British\nColumbia may be about latitude 52\u00C2\u00B0N.\nLonicera ciliosa is present from British Columbia south to northern California, especially west of the Cascade\nsummits, and east to Montana, Utah and Arizona. It is widespread in open woods and thickets, hillsides and\nridges, from sea level to moderate elevations in the mountains. In British Columbia the species is common in the\nsouthern half of the Province in open woods and brush below 460 m west of the Cascades and in the Wet Interior\nZone, and there is sporadic occurrence to the Kootenay River Valley. It is often found clambering through small\nconifers in the woods.\nLonicera dioica occurs from Quebec south to North Carolina and Iowa, and west to the Mackenzie District,\neastern British Columbia, South Dakota and Oklahoma. It is usually found in woods, often in wet places. In\nBritish Columbia it is present in the northeastern and southeastern regions of the Province, in semi-open dry\nforests above 610 m in the Rocky Mountain region. Collections have been made at Fairmont, Windermere,\nField, Peace River, Hudson Hope and on the Liard River near Smith River.\nLonicera 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\nCascade summits. In British Columbia it occurs west of the Cascades on rocky dry open hillsides, woods and\nthickets, often sprawling over broken rock or low shrubs. It is present in the southern Coastal region, the Gulf\nIslands, and on south Vancouver Island.\nLonicera involucrata occurs from southern Alaska south along the coast to Santa Barbara County in\nCalifornia 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\nthe mountains. In British Columbia it is rather sporadic but often locally abundant on moist sites with rich soil\nto at least 1525 m throughout the Province. It is especially abundant in the Coastal and Interior Wet Belt Zones.\nIt forms a conspicuous element of the lowland coastal vegetation throughout the Queen Charlotte Islands. On\nthe coast it is associated with Thimbleberry, Red-osier Dogwood and Pacific Crabapple. In the interior it is\nusually found in more open situations with Salmonberry, Red-osier Dogwood, Water Birch and Cow Parsnip. tD /\nLonicera utahensis is present from southern British Columbia and the Olympic Mountains of Washington\nsouth to Crater Lake and the Blue Mountains of Oregon, and east to Alberta, Montana, Wyoming and Utah.\nThere are varied reports of its occurrence in northern California, although Munz and Keck (1973) state that it\nprobably does not occur in that state. It is widespread on moist wooded or open slopes, by stream banks and at\nthe 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\nthe Cascades to the Rockies, and also at subalpine elevations at the Coast. It also extends north in the mountains\nto latitude 52\u00C2\u00B0N at Anahim Lake and in the Rockies.\nDescription\nLonicera caerulea \u00E2\u0080\u0094 a highly variable species. Branchlets stiff and glabrous, or hairy only when young,\nalthough some forms have branches that are much more hairy or downy. The bark is often shredded and light\nbrown. Winter buds are often superposed, short, spreading, and with valvate lower scales. The leaves are bright-\ngreen, elliptic to oblong-obovate or oblong, have a rounded or obtuse apex, and are 0.6-3(-4) cm broad. They are\nglabrous or sometimes pubescent above, but are more or less hairy beneath, especially on the midrib and veins.\nThe margin is sometimes villous-ciliate. The petiole is 2-5 mm long and hairy. The flowers are in axillary pairs,\npale-yellow to yellowish-white, sometimes tinged with red, and appearing in (April-)June to July throughout the\nrange. The corolla is funnelform, (0.9)1.0-1.3(-l.9) cm long, and 5-lobed to scarcely bilabiate. There is a short\nthick or gibbous spur at the base of the tube. The corolla tube is hairy within and without. The bracts are paired,\ngreen, oblong-linear, and narrow, 3-6(-10) mm long, and the longer ones are somewhat foliaceous in texture but\nstill inconspicuous. The bractlets are wholly connate to form a narrow-mouthed cup completely enclosing the\novaries. The peduncle is axillary on twigs of the season. The fruit is oval to ellipsoid, and 0.5-1.0 cm diameter.\n2n = 18, 36.\nLonicera cilosa \u00E2\u0080\u0094 a twining vine, often growing through and around the branches of other shrubs or small\nconifers. The twigs are hollow, and glaucous when young, becoming glabrous with age. They are sometimes\nmore or less evergreen. The leaves are mostly elliptic to occasionally ovate or obovate, with a rounded to acute\napex 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\nflowers are yellowish to reddish-orange, sometimes tinged purple on the outside, and appear in May to July (to\nAugust) throughout the range. The flowers are fragrant. The corolla is tubular-funnelform to narrowly funnel-\nform, (2-)2.5-4 cm long, and shallowly or not very strongly bilabiate. The corolla tube is downy outside and\npubescent on the inside adjacent to the stamens, and there is a gibbosity on one side at the base on the outside.\nThe stamen filaments and style are slightly exserted and hairy. The flowers are sessile. The fruit is orange-red to\ncoral-red with orange pulp inside, 5-6(-10) mm diameter, and in groups of 3 to 4 that develop at the expense of\nthe remainder. The seeds are large and yellow. The fruit is ripe in September.\nLonicera dioica \u00E2\u0080\u0094 a bushy shrub, with essentially glabrous stems. The bark is gray or straw-colored. The\nwinter buds are solitary, ovoid, and have ovate to narrowly triangular scales. The leaves are usually elliptic to\nobovate but are variable, tapered at the apex and base, 3.5-10 cm long and 2.5-5 cm broad. They are green and\nglabrous above, glaucous and more or less pubescent below. The margin is finely hairy. The upper one or two\npairs of leaves are united around the stem at the base forming a concave disc. The remainder of the leaves are\nalmost sessile. The flowers are yellowish, but often tinged purplish or reddish, and appear in June and July\nthroughout the range. Two or more pairs of leaves beneath the inflorescence are united. The corolla is strongly\nbilabiate, with the lobes varying from about half the length of the tube to approximately equal to the tube in\nlength. The tube is slightly swollen at the base, and is glabrous outside and glabrous to densely hairy within. The\nfilaments of the stamens are attached nearly at the opening of the tube, and have downy bases. The style is\nusually glabrous to somewhat pubescent, and is distinctly longer than the tube. The fruit is coral-red, nearly\n10 mm in diameter, and fleshy. The seeds are light brown, elliptic, and 3.4 mm long by 2.6 mm wide. A slight\ngroove or ridge may be apparent. The fruit is mature in July to August or September, and is dispersed during\nAugust to September or October.\nLonicera hispidula \u00E2\u0080\u0094 vine or climbing shrub with slender, freely-branching stems l-3(-4) m long. The\nbranches are villous-hirsute with glandular hairs, varying to glabrous. The young stems are often hairy, varying\nto glaucous or glabrous. The leaves are ovate to ovate-oblong or somewhat elliptic, often cordate or subcordate\nat the base, 2-7 cm long and 1.5-5 cm broad. They are green and glabrous or hairy above, glaucous and more or\nless pubescent to glabrate beneath. The upper 1-3 pairs of leaves are connate around the stem. The petioles are\nshort. The central axis of the inflorescence varies from glabrous to hirsute or glandular. The flowers are reddish\nto purplish, often being yellowish inside in the latter state, or sometimes pink or yellow tinged with purple. They\nappear in June to August over the range. The corolla is 1-2.5 cm long, and is strongly bilabiate with the lobes\nabout as long as, or longer than, the tube. The tube is slightly gibbous near the base, and is densely hairy inside\nand glandular-pubescent outside. The stamens and style are long exserted, and the filaments are hairy below.\nThe fruit is red, almost 10 mm in diameter, and is rather juicy. Only a few fruits will mature in most years.\n2n = 18.\nLonicera involucrata \u00E2\u0080\u0094 an erect to spreading or rounded shrub. The twigs are stout, 4-angled, and glabrous or\nsometimes long-hairy, especially when young. The bark is light-brown or purplish, becoming gray with age, and\nfreely exfoliating. The winter buds are often superposed, glabrate, and the bud-scales are parted. The leaves are\nnarrowly 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\nand usually glabrous above, occasionally being sparsely pubescent on the veins, and bright green and glabrous to\nslightly downy beneath, especially along the main veins. The margin is often ciliate. The petiole is 5-6(-12) mm\nlong. The flowers are yellow, sometimes tinged with red, and appear in March or April to August, partly\ndepending on the elevation. The corolla is narrowly funnelform to subcylindrical, l-1.5(-2) cm long, and nearly\nregular with the lobes almost equal. The tube has a gibbosity or short thick spur on one side at the base, and is\nglandular-hairy outside. The stamens are shorter than the tube, and are glabrous, or nearly so, and glandular.\nThe anthers are often slightly exserted. The style is slender, longer than the stamens, and is glabrous. The bracts\nare (0.8-)l-l .5(-2) cm long, ovate or oblong, glabrous or often glandular-pubescent, and dark-red. The bractlets\nare axillary, slender, and 0.5-5 cm long. The fruit is shining purple-black or black, globose to ovoid, 8-10 mm in\ndiameter, paired and wholly distinct above the bracts. The seeds are slightly shiny, black, elliptic, 2.7 mm long\nand 1.8 mm wide. The fruits are mature in July to August, and are said to be poisonous by the Indians, although\nthey are so bitter and nauseous to taste that there is little danger of a sufficient number being eaten to cause\nconcern. 2n = 18.\nLonicera utahensis \u00E2\u0080\u0094 a shrub, with a number of irregular straggling branches from a long thin stem. The\nbranches are slender, spreading and glabrous. The bark is gray, and often rather dead looking. The leaves are\nelliptic to somewhat ovate or oblong, with a broadly rounded to obtuse apex and an obtuse to blunt or\nsometimes 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.\nThe flowers are in axillary pairs, although one member of the pair is often dwarfed, and are white to pale yellow,\noften 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\ntube 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\nscarcely united at their bases when young, becoming firmly united when mature. There is 1 pair of bracts, which\nare 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\ndiameter. There are 2-4 seeds. The peduncles elongate in fruit to 35 mm long.\nVarieties and Ornamental Cultivars\nThe members of the genus Lonicera hybridize readily, and there are therefore a number of ornamental\ncultivars in some species.\nLonicera caerulea is a variable shrub with several varieties throughout its range. The form found in British\nColumbia is L. caerulea var. cauriana (Fernald) Boivin, which is the cordilleran form of the species. This variety\ndiffers from the type in that it is 0.2-1 m tall, the branches are pruinose or puberulent and sparsely hirsute, and\nthe leaf margins are always villous-ciliate. The fruit is often red instead of blue or blue-black, and the seeds are\nwhitish-brown, orbicular and 1-1.7 mm long.\nLonicera ciliosa has one named variety, var. occidentalis, in other parts of the range, which has more brightly\ncolored fruit than the type.\nSeveral varieties of Lonicera dioica are recognized. It is represented in British Columbia by L. dioica var.\nglaucescens (Rydberg) Butters in Clements, Rosendahl & Butters. This form occupies the more western part of\nthe range of the species, and differs from the type in that the leaves are obovate to narrowly elliptic and are\n5-8 cm long. The flowers are more purplish, and become reddish with age. The corolla tube is often downy\noutside as well as inside.\nThere are several forms of Lonicera involucrata. The form occurring in British Columbia is the typical var.\ninvolucrata, which has the corolla rarely tinged with red, and the plant is less than 1 m tall.\nPropagation\nThe genus is easily propagated by either seeds or vegetative means, but the species hybridize readily so that\nforms may not come true from seed. However, the seed is an excellent means of raising new forms and is used\nfor plant breeding purposes. The seeds of many species show double dormancy, so that germination may take a\nlong time.\nThe fruit should be collected when it is ripe, and the seeds extracted by macerating the fruit in running water,\nallowing the empty seeds and pulp to float away. The seeds should be dried, and then either stratified or stored in\na sealed container at 1-3\u00C2\u00B0C for up to one year. In the absence of other information, the seed should be stratified\nfor 3 months at 5CC and then sown. If germination does not occur within 4 months, the seeds can be stratified\nfor a further 3 months at 5\u00C2\u00B0C.\nMost of the North American species, except L. dioica, will germinate best if they are warm stratified at room\ntemperature for 3 months, followed by 3 months cold stratification at 5\u00C2\u00B0C before sowing. The germination\nperiod should have alternating temperatures of 30\u00C2\u00B0C day and 20\u00C2\u00B0C night. This regime, should give satisfactory\nresults. Lonicera dioca may be sown in the spring without treatment with up to 95% germination after\n80-100 days.\nThe seeds should be covered with 3-6 mm of soil when sown. Mulching with 5-7.5 cm of straw helps to prevent\nexcessive drying during the germination period.\nSoftwood and hardwood cuttings are both easy to root, although it is probably best to use moderately firm\nyoung shoots taken in July or August. The cuttings should be planted either in straight sand or in peat moss, and\nkept moist and shaded. The use of a rooting hormone has been found to be advantageous. It is also possible to\ntake cuttings and plant them in very sandy soil under a cloche outside. Each node can be split in half lengthwise\nand will root to produce two plants.\nIn addition, Lonicera species may be propagated by layering and by the division of large individual plants\nusing a sharp spade. Transplanting\nShrubs can be transplanted successfully, particularly after growth has been completed for the season. Pruning\nof weak wood and removal of some new growth will enhance success of the transplant.\nConditions for Cultivation\nLonicera species all like a well drained good loamy soil with a pH about 6.0. All forms prefer positions where\nthe aerial parts are in the sun but the roots are shaded and cool (much like Clematis). All are easily grown within\ntheir hardiness limits (some are hardy to Zone 2 Canadian, or even possible to Zone 1), and will usually produce\na wealth of foliage, flowers and fruit with very little care. The vine forms require a trellis, arbor or other\nsupport.\nLonicera ciliosa is hardy to USDA Zone 6. Lonicera dioica is hardy to USDA Zone 2, and will succeed very\nwell in good garden soil, forming a low spreading rather elegant bush if given the support of a stout central\nstake. Lonicera doica var. glaucescens is hardy to Canadian Zone 2 or USDA Zone 4. Lonicera involucrata is\nhardy to Canadian Zone 1 or USDA Zone 4, and is a robust adaptable species growing equally well in seaside\ngardens and in industrial areas. Lonicera utahensis is hardy to USDA Zone 6.\nLonicera species require little regular pruning. The shrubby forms benefit from an occasional shortening of\nlong growths that are spoiling the shape of the plant, and old weak growths of the climbing forms may be pruned\nout. They should all be pruned immediately after flowering.\nLandscape Value\nThe genus contains a number of extremely beautiful species, and every climbing species that is hardy is worth\ngrowing. 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\nattractive habit of growth. The fruits are very attractive to birds.\nThe climbing species are best scrambling over other bushes or tree stumps, trellises or pergolas, although some\nmay be trained to grow as small standards. Some of the less rambling forms may be trained up stout posts\n1.25-1.80 m tall, and then allowed to form loose spreading shrubs with no further support.\nLonicera caerulea has little or no merit for gardens, although it is botanically interesting. The native form,\nvariety cauriana, is of value for its habit of growth, flowers, and fruit.\nLonicera 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.\nLonicera dioica will form a low spreading, rather elegant, bush if given the support of a stout central stake. It\nhas no great beauty of flower, but is striking because of the very glaucous undersurfaces of the leaves. Lonicera\ndioca var. glaucescens has more attractive flowers than the species, and can be used as a perennial vine.\nLonicera hispidula should be grown in a native garden for its attractive habit of growth.\nLonicera involucrata is a loose straggly bush that has no real ornamental value, except that it is hardy to\nCanadian Zone 1, and would therefore be useful in such very cold areas. It is also useful in a native garden, and\nas ground cover bush plantings in revegetation programs.\nLonicera utahensis has an attractive habit of growth as well as attractive flowers, and is a particularly\ncharming small shrub for shady gardens.\nAvailability\nThe native species are apparently not available from local nurseries, although Lonicera caerulea var.\ndependens may be found.\nOther uses\nHummingbirds and long-tongued insects are attracted to the flowers because of the nectar, and birds eat the\nberries. Bears also apparently consider the berries of Lonicera involucrata as a favorite food \u00E2\u0080\u0094 in fact this\nspecies was called Bearberry or Grizzly Berries by the native Indians.\nLonicera 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.\nThe Indian tribes of British Columbia and western Washington had various uses for several of the native\nLonicera species. In British Columbia, the children of the Lower Lillooet tribe at Pemberton used to suck the nectar from the\nflowers of L. ciliosa. The Thompson Indians used to peel the thicker stems of this species, boil them and then\ndrink the decoction as a tonic. They also used a fibre obtained from the stems as a thread or twine. The Sechelt\nIndians used to crush the tender green shoots and steep them in water, which was then used as a hair tonic. The\nSaanich tribe believed that the berries were poisonous. The Squamish Indians believed that the vine was used as a\nswing by ghosts \u00E2\u0080\u0094 their common name for the plant means \"swing of the ghosts\".\nThe western Washington tribes made more use of Lonicera ciliosa than did the British Columbian Indians.\nThe Swinomish boiled the bark as a tea for colds and sore throats, or swallowed the juice from chewed leaves for\na cold. Leaves were bruised and soaked in hot water, and a woman then held her breasts over the steaming water\nto stimulate the flow of milk after childbirth. The Chehalis tribe crushed leaves in water, which was then used to\nbathe little girls so that their hair would grow long and shiny. The Chehalis and Squaxin tribes dipped leaves in\nwater that the women then drank as a contraceptive, but the Squaxin also drank this water as a treatment for\n\"womb troubles\". The Klallam chewed the leaves and then put them on bruises. The Snohomish believed that\ncrows used the plant as a swing.\nLonicera involucrata was used by the Indians much more than any other Honeysuckle. The Thompson and\nLillooet tribes of British Columbia believed the berries to be highly poisonous \u00E2\u0080\u0094 but the Flathead Salish of\nMontana ate them as a powerful laxative. Most tribes considered the berries to be at least inedible if not actually\npoisonous, except for birds and bears. The Bella Coola Indians chewed the leaves and applied the cud to an\n\"itch\", or applied it to an opened boil to draw out the poison. The leaves were crushed and applied to burns and\nto the sores of gonorrhoea. Gonorrhoea was also treated by applying toasted, pulverized bark. The bark was\nboiled and the decoction drunk for a cough. The Southern Carrier boiled the bark for five hours, and then used\nthe decoction daily as an eyewash. The Gitksan used the fresh juice of berries for sore eyes, or the inner bark\n(when berries were unavailable) was soaked in water to make a milky solution used as an eyewash. The\nThompson Indians boiled the leaves and then applied them to any part of the body that was swollen. Sometimes\nthe leaves were bruised before boiling as this was believed to greatly increase their strength. They also used a\ndecoction of leaves and twigs as a liniment. The Squamish chewed the bark and blew it onto the cheek for a\ntoothache, or blew the fresh saliva onto a boil \u00E2\u0080\u0094 this was said to be particularly effective if done in the morning\nimmediately after waking, but before speaking. The Haida rubbed the berries into the scalp to prevent gray hair.\nLonicera involucrata was an important medicinal plant for the Kwakiutl tribe on Vancouver Island. They boiled\nthe leaves (or roots in winter), strained them and applied them as a hot compress to swollen shoulders and feet.\nThe bark was boiled and placed on the breasts of a woman who had just given birth to make her milk flow, or\nthe leaves were mixed with the basal leaves of Yarrow (Achillea millefolium) for the same purpose. The bark,\nberries or leaves were mixed with Sea Wrack (Fucus gardneri) or Sea Lettuce (Ulva lactuca), dried Tobacco\n(Nicotiana sp.), and Red Alder bark (Alnus rubra) to make a poultice for swellings and sores. They never ate the\nberries, believing that they would lose their voices and become stupid if they did so, but they did make a purple\ndye from the berries mashed with Salal berries (Gaultheria shallon).\nThe Quileute Indians of western Washington used the juice from L. involucrata berries to paint dolls' faces.\nThey chewed the leaves as an emetic when poisoned. The women of the Quinault and Makah tribes chewed the\nleaves during childbirth. All the western Washington tribes associated the plant with the crow.\nIndian hunters in the Okanagan sometimes ate the very juicy berries of L. utahensis, believing them to be a\ngood emergency supply of water.\nDiseases and Problems of Cultivation\nThe climbing members of the genus are very subject to aphid attacks in summer, especially during hot dry\nspells. However, these attacks are often overcome naturally if the plants have been given good loamy soil and\ncool moist root conditions. Some vines are very susceptible to infestations of spider mites.\nIn general, the genus is not susceptible to severe disease problems, although Leaf Curl makes the leaves\nthicken, curl and become deformed, and Powdery Mildew may be a problem in some areas, including the Lower\nMainland of British Columbia.\nThe following diseases have been identified on the native species in the wild in British Columbia:\u00E2\u0080\u0094\nLonicera ciliosa \u00E2\u0080\u0094 Cercospora antipus. Leaf Spot; C. periclymeni; Poria ferrea, Stem Rot; and Stereum sp.\nLonicera hispidula \u00E2\u0080\u0094 Microsphaera penicillata, Powdery Mildew; and M. alni. Lonicera involucrata \u00E2\u0080\u0094 Kabatia lonicerae var. involucratae, Leaf Spot; Microsphaera penicillata, Powdery\nMildew; M. alni; Leptosphaeria dumetorum; and Ophiobolus minor.\nLonicera involucrata \u00E2\u0080\u0094 Kabatia lonicerae var. involucratae, Leaf Spot; and Leptothyrium periclymeni var.\npericlymeni.\nThe Adventive Species\nTwo 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\nalien reproducing without cultivation.\nLonicera etrusca Santi, Etruscan Honeysuckle, is a native of the Mediterranean region that has now become\nnaturalized and established in thickets along the coast of Oregon (in Lane and Curry Counties), in Del Norte and\nHumboldt Counties of California, and in southwestern British Columbia. In British Columbia, it is established\nand common on southern Vancouver Island, particularly near Saanich and Sidney, as well as on the Lower\nMainland and north to the Queen Charlotte Islands. It is a vigorous vine, with glabrous to sometimes pubescent\ntwigs. The young shoots are reddish-purple, especially in cultivated forms. The leaves are oval to obovate with\nan obtuse apex and a rounded or broadly tapered base, 4-9 cm long and 2.5-5 cm broad. They are glaucous and\nusually somewhat downy beneath. The upper pairs of leaves are sessile and united at the base around the stem,\nalthough sometimes all the leaves are free. The inflorescence is a terminal group of dense spikes, each containing\nmany flowers, and there are also sometimes two or three axillary heads. The flowers are yellowish-white or\ncreamy-white tinged with purplish-red, becoming a deeper yellow with age, 3-5 cm long, and fragrant. The\ncorolla is conspicuously bilabiate with the tube and lobes nearly equal in size. The fruit is red, and the berries are\nnever united. There are several varieties and cultivars of L. etrusca, the variety pubescens being the form usually\nfound in cultivation. The cv. Superba is a very vigorous form, with panicles larger than the species and reddish-\npurple shoots. The species is hardy to USDA Zone 7, and revels in full sun. It is not often seen in gardens in\nNorth America, but probably should be grown more widely as a garden and trellis plant. It is perhaps the most\nstriking of all Honeysuckles when growing well, with long shoots that branch and form immense bouquets of\nflowers. It is probably best grown in the drier areas of the region. 2n = 18.\nLonicera 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\nnaturalized in some areas, and has been reported as an adventive from the Boreal Spruce Zone of British\nColumbia. It is a deciduous shrub, with ovate to oblong leaves. The leaves are pointed at the apex and tapered at\nthe base, 3.5-10 cm long, and more or less downy on both sides. The paired flowers are in shades of pink, turning\nyellowish with age, 1.9 cm long, and not fragrant. The fruit is bright red and rather transparent. There are a\nnumber of cultivars of this species: cv. Grandiflora with large rose-tinted flowers; cv. Carneorosea (or\nCarneo-Rosea), which has the deepest rose-colored flowers of all forms; cv. Alba; cv. Ochroleuca; and cv.\nCarnea. The two cultivars cv. Grandiflora and cv. Carneorosea are among the best of the numerous hybrid bush\nhoneysuckles, with attractive fruits as well as flowers and habit.\nOrigin of the Name\nThe Family name Caprifoliaceae is derived from Caprea, a she-goat, which has twisted horns in certain\nbreeds. These horns are suggested by the twisting stems of the climbing Honeysuckles. The generic name\nLonicera commemorates Adam Lonitzer (1528-1586), a German physician and naturalist who wrote a herbal\n(Kreuterbuch) that was reprinted many times between 1557 and 1783. The specific name caerulea means 'dark\nblue' or 'blue', referring especially to the shade of deep blue seen at midday in the Mediterranean sky. The origin\nof the subspecific cauriana seems obscure, although it may be derived from the Latin counts, 'northwest' or\n'northwest wind' and the suffix -iana, 'belonging to', referring to the geographical distribution of the plant. The\nspecific name ciliosa means 'fringed' or 'ciliate', referring to the ciliate hairs of the leaf margin. The specific\nepithet dioica, 'dioecious' or 'coming from two', means that the male and female flowers are on separate plants.\nThis meaning, however, is obscure in L. dioica, which is not dioecious. The subspecific glaucescens means\n'having some bloom', referring to the glaucous underside of the leaves. The specific epithet hispidula refers to\nthe stiff and rigid hairs on the plant. The specific name utahensis commemorates the fact that the plant was first\ndescribed from a specimen collected in Utah.\nThe 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\nbanks of the Kooskoosky\" where it was collected by Meriwether Lewis. It was first introduced to cultivation in\nGreat 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\nBritain in 1890, while L. dioica itself was first cultivated in 1636. The type locality of L. hispidula is \"woods of\nNorth West America\". It is believed to have been described from garden plants grown from seed collected in\nnorthwestern America by David Douglas. The type locality of L. involucrata is \"wooded country between 54\nand 64 degrees latitude\" [Canada], where it was collected by Sir John Richardson (1787-1865). It was first\ncultivated in Great Britain about 1824. The type locality of L. utahensis is \"Wahsatch Mountains, Utah, in\nCottonwood Canyon; 9,000 feet altitude\" where it was collected by Sereno Watson (1826-1892).\nThe specific name etrusca means 'from Tuscany', an area in northern Italy known as Etruria in classical times.\nThe type locality is \"region of Tuscany, Italy\", and it was first cultivated about 1750. The specific name notha\nderives from the Greek nothos, 'false', 'baseborn', or 'bastard', referring to its hybrid origin. The plants were\nraised from seed in Munich Botanic Garden in about 1878, the seed having been obtained from St. Petersburg\n(Leningrad) Botanic Garden in Russia where the cross had occurred.\nREFERENCES\nAbrams, L. andR. S. Ferris. 1960. Illustrated Flora of the Pacific States. Vol. IV. Bignoniaceae to Compositae. Stanford University Press, Stanford, California. OO\nBean, W. J. 1973. 8th ed. rev. Trees and Shrubs Hardy in the British Isles. Volume 2. D-M. John Murray (Publishers) Ltd.,\nLondon, in collaboration with The Royal Horticultural Society.\nClark, L. J. 1973. Wild Flowers of British Columbia. Gray's Publishing Ltd., Sidney, B.C.\nGarman, E. H. 1973. 5th ed. rev. The Trees and Shrubs of British Columbia. Handbook No. 31. British Columbia Provincial\nMuseum, Victoria, B.C.\nGunther, E. 1945. Ethnobotany of Western Washington. University of Washington Publications in Anthropology 10(l):l-62.\nHillier's Manual of Trees and Shrubs. 1972. Hillier and Sons, Winchester, England.\nHitchcock, C. L. et al. 1959. Vascular Plants of the Pacific Northwest. Part 4. Ericaceae through Campanulaceae. University\nof Washington Press, Seattle.\nLyons, C. P. 1965. Rev. ed. Trees, Shrubs and Flowers to Know in British Columbia. J. M. Dent & Sons (Canada) Ltd.,\nVancouver, B.C.\nMunz, P. A. and D. D. Keck. 1973. A California Flora and Supplement. University of California Press, Berkeley, California.\nTaylor, 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,\nB.C.\nToms. H. N. W. 1964. Plant Diseases of Southern British Columbia. A Host Index. Reprinted from: Canadian Plant Diseases\nSurvey 44:143-225. Canada Department of Agriculture, Ottawa.\nTurner, N. J. 1975. Food Plants of British Columbia Indians. Part 1. Coastal Peoples. Handbook No. 34. British Columbia\nProvincial Museum, Victoria, B.C.\n . 1978. Food Plants of British Columbia Indians. Part 2. Interior Peoples. Handbook No. 36. British Columbia\nProvincial Museum, Victoria, B.C.\nU.S.D.A., Forest Service. 1974. Seeds of Woody Plants in the United States. U.S.D.A., Forest Service, Agriculture\nHandbook 450. Announcement \u00E2\u0080\u0094 Change in Subscription Rates\nThe subscription price of the journal will increase to ten dollars per annum, effective Spring 1980 (Volume\n11, Number 1). The cost of single numbers will increase to two dollars and fifty cents, except for special issues.\n64\nClimatological Summary'\nData 1979\nJULY\nAUGUST\nSEPTEMBER\nAverage maximum temperature\n21.2\u00C2\u00B0C\n20.6\u00C2\u00B0C\n18.8\u00C2\u00B0C\nAverage minimum temperature\n13.2\u00C2\u00B0C\n13.6\u00C2\u00B0C\n12.4\u00C2\u00B0C\nHighest maximum temperature\n28.5\u00C2\u00B0C\n23.5\u00C2\u00B0C\n27.3\u00C2\u00B0C\nLowest minimum temperature\n7.5\u00C2\u00B0C\n11.1\u00C2\u00B0C\n9.3\u00C2\u00B0C\nLowest grass minimum temperature\n5.0\u00C2\u00B0C\n3.2\u00C2\u00B0C\n5.1\u00C2\u00B0C\nRainfall/no. days with rain\n25.8 mm/7\n20.0 mm/8\n115.5 mm/14\nTotal rainfall since January 1,1979\n375.4 mm\n395.4 mm\n510.9 mm\nSnowfall/no. days with snowfall\n0\n0\n0\nTotal snowfall since October 1,1978\n9.6 cm\n9.6 cm\n9.6 cm\nHours bright sunshine/possible\n290.6/482.5\n252.9/439.0\n167.7/372.2\nAve. daily sunshine/no. days total overcast\n9.4hr/l\n8.2hr/2\n5.6hr/4\n\"Site: The University of British Columbia, Vancouver, B.C., Canada V6T1W5\nPosition: lat. 49\u00C2\u00B0 15'29\"N; long. 123\u00C2\u00B0 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\nwas held in September during the first week of classes. More than seven thousand house plants were sold during\nthe three-day event. Pleurotus ostreatus, Oyster Mushroom,\nxO.66. The Oyster Mushroom is common\nfrom spring to late fall on the trunks of\ndead trees, especially Alder. It is edible\nwhen young.\nVolume 10\nNumber 3\nDAVIDSONIA\nFall 1979\nContents\nInsectivorous Plants in British Columbia 41\nAll-America Selection Display Garden 53\nThe Genus Lonicera in British Columbia 54\nAnnouncement \u00E2\u0080\u0094 Change in Subscription Rates 64\nClimatology 64"@en . "Periodicals"@en . "Vancouver (B.C.)"@en . "QK71.D39 B75"@en . "QK71_D39_B75_10_03_1979"@en . "10.14288/1.0115077"@en . "English"@en . "Vancouver : University of British Columbia Library"@en . "Vancouver: The Botanical Garden of The University of British Columbia"@en . "Images provided for research and reference use only. Permission to publish, copy, or otherwise use these images must be obtained from the University of British Columbia Botanical Garden: http://www.botanicalgarden.ubc.ca"@en . "Original Format: University of British Columbia. Archives"@en . "University of British Columbia. Botanical Garden"@en . "Davidsonia"@en . "Text"@en . ""@en .