Plant Life in the Scottish Highlands
Flowers of the Mixed Wood and Copses Part 2

Wood Sorrel (Oxalis Acetosella)

The present subject is a very typical woodland plant, and a very common one in woods throughout the Highlands.  It is very fond of damp, mossy banks and dells, wherever there is shade.

One can say, without prejudice, that it is one of our most beautiful and dainty plants with its delicate, hanging flowers and shapely, trifoliate leaves.  There are few lovelier sights than a secluded dell in the depths of the woods, where the soil is carpeted by deep green mosses which form a perfect setting to the pale chaste flowers of the Wood Sorrel, displayed in company with the pale yellow stars of the Yellow Pimpernel and the delicate blue blooms of the Speedwell.

The Wood Sorrel is specially adapted to its woodland habitat, and that is very successful in the life struggle is show by its abundance and the extensive colonies that it forms.

It perennates by means of a long, slender, pinkish, creeping rhizome which is partially buried in the humus and mosses.  In the winter it is covered by a blanket of fallen leaves and sleeps snugly in its comfortable bed, whilst the winter gales howl through the naked branches of the long-suffering trees.

Fine roots are give off from the nodes, and anchor the plant securely in the soil.  Each year the rhizome lengthens by means of a terminal bud, and as it is gregarious plant it soon possesses a large territory which it is continually increasing.

The leaves, which are given off from each node, have fine, long, pinkish stalks surmounted by three heart-shaped, thin, pale green leaflets which are usually pinkish on the under side, an are covered with short, soft hairs.  The thin texture of the leaves is an adaptation to the shady conditions prevailing in woodlands, permitting the light to penetrate more easily into the tissues.

The leaves are very interesting subject from the botanical point of views, because they illustrate the rare occurrence of movement in the plant world.

In daytime, under ordinary conditions, the three leaflets spread outwards until they are in the same plane where they obtain the maximum illumination.

In the evening, or on the approach of cold weather, or in strong sunlight, the leaflets commence to move.  The two lateral leaflets move backwards toward each other, until their under surfaces meet, whilst the upper one bends backwards and downwards until it covers the two lateral ones.  In such a position the stomata on the under surfaces are closely protected and transpiration is reduced to a minimum.

The movement results in the delicate tissues being less exposed to the sun’s rays during the day and diminishes loss of heat from the leaves by radiation at night.

The movement is caused by changes in the turgidity of the thickened stalks or pulvini of each leaflet.

The flowers are produced singly at the summit of a slender stalk which has two tiny bracts about half-way up and is covered by fine, weak hairs.

Each flower is delicately beautiful structure.  The calyx is a tiny green cup formed by five obtuse thin sepals which are united together towards the base.  The corolla is also cup-shaped and if formed of five fairly large egg-shaped white petals which have a very delicate texture and are beautifully marked by fine lilac veins, and are often pinkish on the outer surface.

Each flower contains ten stamens arranged in two whorls, the stamens in one whorl being longer than those in the other.

The ovary is a conical green body surmounted by five spreading styles with receptive stigmatic surfaces on the upper side.

The flowers produce little or no nectar and are visited by small bees and flies for the sake of their pollen.  The stamens and the stigmas mature at the same time but as the latter are above the stamens, cross-pollination is assured.

Besides the showy flowers, however, the Wood Serrel produces small blind flowers, known as cleistogamous flowers.

If we pulled up a plant and examined it, we should notice some tiny, green, bud-like object on long, slender stalks, produced at the base of the plant.  If one of these buds was carefully opened, it would be found to be hollow and to contain a pistil and stamens.  These flowers, however, have no entrance and are never visited by insects, their only function being to produce self-fertilized seed.

Many plants produce such cleistogamous flowers and by their means the plant is assured of setting seed.  Experiments have shown that these seeds are quite fertile and give rise to strong plants.  They are formed with the greatest economy, only four hundred pollen grains being produced in the cleistogamous flowers, whilst over a million may be produced in the showy flowers.

No plant can afford to depend on self-fertilized seed alone and lose the advantage of cross-fertilization, hence the showy flowers that can only be cross-pollinated.

Hedge Woundwort (Stachys sylvatica)

Some plants impress themselves upon us by the striking beauty of their flowers, other by their sweet perfume, and others, like our present subject, by their abundance.

The Hedge Woundwort is one of our most common plants and is to be found on the edges of woods, in thickets and bushy places and in hedgerows, but it does not attain very high altitudes.

It is a tough, robust plant, the companion of nettles, docks and Figworts, rough members of the plant world, jostling each other for space in the limited place available in the bushy thickets and hedges.

In spite of its abundance, however, it s a very interesting plant and a good illustration of the fact that we can learn more from a detailed study of common things than from many of the rarer things in life.

It has set out to conquer in the struggle for life and has done so by means of marvelous adaptation to ensure perennation, to colonize, to defeat competitors, and to ensure cross-fertilized seed in abundance as well as its efficient distribution.

It has a tough, creeping rootstock which is often woody in texture, and is buried in the soil.  From the nodes, long, fibrous roots are given off which anchor the rhizome firmly in the soil.  In the autumn, the rootstock produces buds which lengthen into short branches call scions.  Tiny, fringed, scale-leaves are given off at the nodes of both the scions and the rhizome.  The scions end in a large bud which in the spring gives rise to the tall flowering shoots.  The rhizome often decays, with the result that the scions form new, independent plants.

The creeping rhizomes advance outwards from the centre of the colony and thus the area of the colony is continually increasing, so that in time the Hedge Woundwort occupies large areas to the complete exclusion of other species, whose seeds are unable to thrive in the shade of the tall stems and leaves of the colony, and whose creeping stems are unable to penetrate the dense mass of roots.

The flowering stems, which attain two to four feet in height and are often much branched above, are covered with coarse hairs and are square in section with strong, woody fibres at each corner which make the stems very tough.

The leaves, which have long, hairy stalks, are produced in opposite pairs, the blades being ovate, cordate and crenate and covered with coarse hairs, which often impart a dull grayish-green colour to them.  The leaves, like the rest of the plant, are strong smelling and this sves them from destruction by herbivorous animals.

The upper part of the stem is taken up by the inflorescence consisting of a tall, leafy spike in which the flowers are arranged in whorls in the axils of opposite leaves, the lower whorls being fairly widely separated.  Toward the summit of the inflorescence the leaves become sessile, narrow and bract-like in order not to hide the flowers.

Each whorl is actually two whorls, one being produced in the axil of each leaf.   The flowers are arranged in a cymose manner, i.e. as in the Red Campion, but in this case the flower stalks are so short that the cyme appears to be a compact whorl.

The result of this arrangement s that the flowers in any particular whorl do not all bloom at the same time.  There is thus a succession of blooms, but owing to the fact that the whorls are arranged in a spike, the lower ones have finished flowering before the upper ones have commenced to show any blooms.  This results in a long flowering period which gives a good chance of much fertile seed being set.  The flowers are specially constructed and adapted to humble-bees.

Each flower is surrounded by a campanulate, hairy calyx which is fringed by five, fine teeth.  Sticky glands are scattered among the hairs, their function being that of preventing creeping insects reaching the flowers.

The corolla is in the form of a tube, about one-third of an inch in length, and is suddenly narrowed towards the base, a ring of hairs completing blocking the entrance to the narrow portion, where the nectar is secreted.  This is an additional protection against creeping or small, flying insects that might try to steal the nectar.

The upper part of the tube widens out to form two lips.  The upper one is concave and projects beyond the tube like a hood, into which the anthers fit and are perfectly protected against rain and damp. The lower lip, which is grooved in the upper part, is hanging and three-lobed, the middle lobe being broad and forming a landing stage for the bees.  It also forms the conspicuous part of the flower, being purple-red in colour, beautifully mottled with white and violet.

The four stamens are arranged in two pairs, which differ in length and are placed one behind the other in the roof of the hood-like upper lip.

The ovary, consisting of four green compartments, sends up a long style, the bifid stigma being situated between the two pairs of anthers.

In the young flower only the first par of anthers are open, and when they have shed their pollen they turn outwards, the second pair then opening to turn outwards also when their pollen is shed.

The lobes of the stigma which occupy the same place as the anthers now open out and, all danger of self-fertilization having passed, they become receptive.

When a bumble-bee alights upon the beautiful lower lip, it pushes its face into the entrance of the tube and in so doing its head brushes the under side of the upper lip. Thus, in a young flower it will be dusted with pollen, but in an older flower it will touch the stigma.  The bee then pushes its proboscis down the corolla tube, through the ring of hairs and so to the nectarines at the base of the ovary, the humble-bee being the only bee which has a tongue long enough to reach so far.

These bees always start with the lower flowers (the older stigmatic flowers) first, visiting the younger upper ones afterwards, and it is thus obvious that they will always bring pollen from one plant to the stigmas of the next plant visited.  Cross-pollination is thus assured, self-fertilization being impossible, and this explains the great success and abundance of this plant.

After fertilization the corolla withers, but the hairy calyx enlarges and completely protects the four developing seeds.  When ripe the seeds are black and shiny and are scattered as the wind shakes the tall flowering stems.

Ground Ivy (Nepeta hederacea)

This is favourite spring flower with many people, and coming into bloom as it does, so soon after the gloomy, cold days of winter, it is doubly welcome.

Grassy woodland banks and hedgerows are its favourite habitat, and here it forms large colonies, often covering the surface with its kidney-shaped leaves and purple-blue flowers.

It is well adapted to colonization as its weak stems creep over the surface of the soil and grasses and root at the nodes.  It is a perennial and produces runners from which new plants are eventually formed, so that in time large colonies appear which are penetrated with difficulty by other plants, and those who find themselves caught up in their advance have a hard struggle to survive.

The stems bend upwards and become erect in spring, and are covered in pairs of long stalked, dark green ,kidney-shaped leaves coated with roughish hairs, and with large rounded teeth around the their margins,

The flowers are a deep blue in colour and produced three or four together n the axils of the upper leaves of the erect portion of the stems, so that they overtop the surrounding vegetation which at this early season is still short.  Later on the colonies disappear in the grasses and rank vegetation, and hence the reason for its early flowering season.

The flower are similar in form to those of the Hedge Woundwort, but these are two kinds of blooms.  Some are large and showy and are hermaphrodite, the others are smaller and less conspicuous and contain a pistil only.

The great botanist Muller believed that this was an aid to cross-pollination, the smaller less conspicuous flowers being visited after the showy ones, they were therefore sure to be pollinated, the bees being covered by pollen from the showy flowers.

Early flying bumble-bees are the only insects capable of reaching the nectarines and of pollinating the flowers, and for this reason they are blue in colour.

The whole plant contains an aromatic, bitter substance which prevent it being eaten by herbivorous animals.

Bugle (Ajuga reptans)

This is another very common woodland plant, but it comes into flower in summer and hence is more common in glades and clearings and thin parts of the wood where there is less shade from the trees.

It possesses a short, thick, erect rootstock producing buds from which long, leafy runners are given off, and these root at the extremities to give rise to new plants.  The Bugle thus forms large colonies and is well adapted to spread over surrounding grasses and other lowly vegetation.

The stock gives rise to a fairly stout, flowering stem covered by several pairs of large, stalked, smooth leaves.

The inflorescence consists of a crowded spike of several whorls of blue flowers which are very conspicuous upon their tall stem, smaller whorls being produced in the lower pairs of leaves.

The flowers are similar in structure to those of the Hedge Woundwort, but as the upper lip is reduced to a small tooth, the anthers are without protection from the corolla.  They are, however, sheltered by the bracts surrounding the next whorl of flowers above them  This is an example of the economy effected by Nature, for as the anthers are effectually protected by the bracts, there is no need for a hood, as in the case of the Hedge Woundwort, and it has thus been dispensed with.

When the pollen is shed, the style lengthens and the stigma is produced in the place originally held by the anthers, which in the meantime have turned outwards in order not to be in the way.

The flowers are pollinated by bumble-bees, and only one kind of flower is produced.

WOODLAND SPEEDWELLS

Two beautiful Speedwells are found in the Highland woods, the Common Speedwell (Veronica officinalis) and the Germander Speedwell (V. Chamaedrys). The former is a typical woodland plant and adapted to shady conditions, but the latter is less tolerant of shad and is much more abundant on the edges of woods and copses, on sunny banks, in clearings and in hedgerows.

The Germander Speedwell is a beautiful little plant, forming dense colonies which, when the flowers are fully open, are covered by a mass of exquisite, bright blue blooms which rival the summer skies in glory and are hardly surpassed for purity of colour by any other British flower.

As a boy I was always attracted to these lovely flowers which grew in profusion on a garden bank of my old home is southern England, but the Scottish plants seem to have deeper, more intensely blue flowers.  This I have remarked in several species of flowers that grow both in the south and in the Highlands, the Dog Rose and the Harebell being good examples.

The present plant is a perennial with long trailing weak stems which clamber over the surrounding vegetation and turn up at the extremities, their lower parts producing roots which anchor them securely.

The stems are covered by pairs of opposite ovate leaves, which are almost stalkless, have a wrinkled surface and are covered with weak hairs.

The flowers are produced on long stalks from the axils of the upper leaves, one stalk arising in each opposite leaf.  They are terminated by a raceme of very beautiful, bright blue flowers with a white conspicuous eye.

The anthers and stigmas mature together, but as the former open outwards there is no danger from self-fertilization.

The flower are pollinated by hover-flies in exactly the same way as those of the Alpine Speedwell (see p. 31).

The Common Speedwell is a rather similar plant, but has thinner leaves as an adaptation to is more shady habitat.

Its many creeping stems, which root at the base, form colonies, although they are not as dense as those of the Germander Speedwell.

The leaves are stalked, but are similar in shape to those of the preceding and are hairy.  The stems do not become so erect and the plant has a much more trailing appearance than  in the preceding.

Its flowers are similarly produced, but are much paler lilac-blue in colour and are rather smaller.  They are pollinated in the same way as those of the Alpine Speedwell, but are less visited by insects than the conspicuous flowers of their woodland cousin and are often self-fertilized.

Yellow Pimpernel (Lysimachia nemorum) 

This is a common plant in shady places along the edges of woodland rills and streams, by springs and in damp places.  It is a member of the Primrose Family.

It is a pretty little plant with its trailing reddish stems rooting at the nodes, its pairs of smooth, dark green, egg-shaped leaves and its solitary yellow flowers on long, fine stalks.

It is a typical shade plant with think leaves into which the weak light can penetrate without difficulty.  The absence of hairs also means that nothing obstructs the passage of the light to the leaf.  As it grows in moist places, it does not fear drought and hence has made no provision against it, thus it is not found where the soil is dry or where it would be exposed to winds.

The flower has a smooth cup-shaped calyx with five pointed teeth, whilst the corolla is a shallow, bright yellow cup about half an inch across with a tiny collar as in the Veronica.  The five stamens have long filaments which spread out from the centre of the flower, so that although the anthers open inwards, the danger of pollen falling on the stigma is averted.

The flowers do not produce nectar and are visited for their pollen by small flies and bees.

Dog’s Mercury (Mercurialis perennis)

This plant is very typical of the woodlands and is one of the first to be found in flower, its bright green stems shooting up almost as soon as the snow has disappeared from the woods.

It comes into flower in March in sheltered spots in the Highlands before the trees have even begun to think of producing their leaves, and thus it can profit from all the available light.  When the leaves throw their dense shade in summer, the plant dies down till the following spring.

This is hardly exact for the Dog’s Mercury is a perennial with a creeping underground rhizome in which food is stored, and which lies dormant in the soil during late summer, autumn and winter.  The rhizomes spread through the soft woodland humus in all directions, with the result that the flowering stems appear in countless numbers beneath the naked trees.

The annual stems, which may attain a foot in height, are at first leafless as the shoot up rapidly into the sunshine, but later they are covered by many dark green, oblong leaves which are crowded together toward the summit of the stem.  They are roughly hairy and toothed, and contain crystals of calcium oxalate in their tissues, which render them unpalatable to herbivorous animals, and even snails and slugs do not enjoy such a bitter meal.

The flowers are unisexual, the females being produced on one plant and the males upon another so that cross-pollination must be effected.  The pollen is carried to the female flowers by the wind and this is another reason for their early flowering, as the wind can blow through the naked woods quite easily whereas in summer the woods are remarkable for their peace and stillness.

The male flowers are produced on long stalks in the axils of the upper leaves and they overtop the plant by several inches.  They are very inconspicuous, being green in colour and produced in clusters toward the summit of the stalks.  The actual flower is reduced to three green segments which correspond to the calyx of other flowers, and about twelve stamens are present in each flower.  A vast amount of pollen is produced, witness to the great waste involved when pollination depends on the wind.

The female flowers are even less conspicuous and as they are borne in a single cluster at the summits of slender stalks which are no long enough to overtop the upper leaves, one must look carefully for them.  The flower has three green sepals as in the males and the rounded ovary produces two longish styles.

After fertilization the ovary becomes warty and often produces prickles so that the seeds are often transported by animals, the prickles catching in their hairy coats as the pass through the undergrowth.  This is another marvelous adaptation to the woodland habitat, as many hairy coated animals make their homes there.

Enchanter’s Nightshade (Circaea)

Under this name we have two British species one, the Alpine Enchanter’s Nightshade (C. alpina), is very abundant in Highland woods and is confined to the northern parts of Britain, the other, the Common Enchanter’s Nightshade (C. lutetium), is abundant in all British woodlands.

These strangely named plants grow in fairly dense shade and flower in full summer when the light intensity is at its lowest.  They are thus typical shade plants and like others of the same class have thin, dark green leaves and white flowers which are more conspicuous than colored ones would be in the semi-light.

The common species is a perennial plant with a creeping rhizome which branches here and there until compact colonies are produced.  This the reason for its persistence as a garden weed.

The annual stems are about one foot in height and are covered with pairs of long-stalked ovate leaves.  These are very thin in texture and are covered like the stems with fine white hairs.

Naked flowering stalks are given off from the axils of the upper leaves, each producing a raceme of small white or pinkish flowers.

Each flower is produced upon a slender pedicel and consists of a calyx of two reflexed  green sepals which are found upon the summit of the ovary.  The corolla has only two petals which spread horizontally.  They are thin textured and often crinkled.  Only two stamens are formed and these are placed in the same way as those of the Speedwell.  The style is situated between the two stamens.

The flowers are visited by small bees and flies.  These alight upon the style and the stamens, the anthers of which dust the under part of the insect’s body with pollen.  As the insect descends upon the style any pollen will be left upon the stigma before the anthers touch its body.  Thus cross-pollination is assured.

The alpine species is smaller in all its parts and has no hairs; its leaves are even thinner and it produces a single raceme of very small flowers.

This plant can be found in the greatest profusion in the beautiful birch woods of the Trossachs between Loch Achray and Loch Katrine.  There, in company with the Yellow Cow-wheat, the Yellow Pimpernel, the Chickweed Wintergreen and tall Mosses, it forms a soft starry carpet under the graceful birches.  The beautiful woods can compete with any Britain for their loveliness and verdure, with their mossy, flower-adorned dells, where one would never be surprised to find a fairy frolic in progress.

This little plant is quite common elsewhere in the western Highlands and in Pertshire wherever the woods are damp and shady.

After fertilization the ovary, in both cases, becomes covered with curved prickles and fors a burr.  These catch in the fur of animals or upon the clothing of passers-by and are transported far from the parent plant before falling to the ground.  In woods where there is a little wind, parachute seeds would be of little use, so the plant has forced animals and man to distribute its seeds.  Thus, even the most humble plants are amazingly adapted to their chosen habitats.

Wild Hyacinth (Scilla non-scripta)

Everyone has their favourite flower which brings back to them sweet memories of the past.  Some prefer the beautiful Rose, others the sweet-scented, shy Violet, others the Primrose, but for me there is no flower to compare with the lovely Wild Hyacinth.

There is probably nothing so glorious in this country as the sight of a blue sea of Wild Hyacinths sweeping away beneath the noble trees of an oakwood.  Probably no other plant can show such glory when in the mass, and in no other country does one plant form such a magnificent display of colour and beauty.  For truly, there is nothing more beautiful than their thousands of spikes, each with up to twenty dainty bells swaying gently in the breeze, perfuming the air with their delicate bells swaying gently in the breeze, perfuming the air with their delicate sweet odour, a nostalgic odour associated with spring, the cuckoo, the sweet tenor of the blackbird on its favourite branch and the twittering of the red-breasted swallow as it sweeps by overhead.

In the Highlands is to be found in every county, but rarely does it make such a show of colour as in the English woodlands.  This is mainly due to the fact that sheep are widespread in the Highland and they prevent the development of great masses of plants.  Wherever woods have been fenced in, however, we can find them in abundance.

The Wild Hyacinth is a typical plant of the oakwoods, and is to be found at its best in the deep humus beneath their noble branches.  It is also fairly common in some birchwoods and is quite frequent in the rough, scrubby woods which straggle along the deep valleys of streams and rills where sheep have difficulty in penetrating.

If we visit the woods in the depths of winter, when the ground is not covered by snow, we should see no sign of the beauty to come.  Deep down in the warm, decaying leaves and twigs, the hyacinth bulb sleeps snugly out of reach of the cold winter winds and frosts.  If we dug up one of these bulbs, we should find that already life was stirring in the centre of the bulb.  If we cut it open, we should find that the centre was occupied by the embryo flower stalk, flowers and leaves.  The bulb itself is remarkable for the fact that the scale leaves are filled with a sticky, unwholesome mucilage which prevents them from being eaten by predatory insects and animals.

In early spring the leaves begin to lengthen and burst above the ground to form a rosette of shiny, bright green, linear foliage.  From their midst arises the tall, cylindrical, flower stalk.  At first it is perfectly erect and the flower buds, protected by the bracts, are pressed closely to the stem so that they will not be damaged in their passage upwards through the soil from the bulb.  At the end of April the flower stalk curves over gracefully at the top and from it the beautiful flowers hang like fairy bells.  Each flower sways delicately upon a tiny bluish stalk, which arises from the axil of a pair of narrow, bluish colored bracts.

The flowers are not really blue in colour, in spite of the popular name of Bluebells, and it is remarkable how few truly blue flowers exist.  They are more accurately described as purple-blue in which the blue predominates.  A white-flowered variety sometimes occurs ad plants with pinkish flowers may be found.

The tubular flowers consist of six segments which spread outwards toward their tips, the edges being slightly recurved.  If we carefully dissected a flower, we should find that a stamen was seated below the middle of each segment so that the anthers formed a ring round the entrance to the flower.  At the base of the flower we should find the round ovary surmounted by a longish style which projected just beyond the ring of anthers.

It is a sad and sobering thought that all this beauty is after all not for our own enjoyment.  The flowers are but signboards displayed for the benefit of the hosts of hive-bees and humble-bees for whom the nectar is reserved at the bottom of the corolla tube, whilst the perfume serves a similar purpose.

If we watched a bee visiting a flower, we should see that it grasped the recurved tips of the corolla on alighting and then pushed its head into the flower, touching the projecting stigma and then the stamens.  Obviously, any pollen brought from another flower will be left upon the stigma and cross-pollination assured.

On visiting the wood in late summer and autumn, we should find that the beautiful flowers had gone, the leaves would be withered and yellow, and all that remained of so much loveliness was an upright stalk with several, brownish capsules, opening at the top by three valves.  As the stalks swayed in the wind, the seeds would be thrown out to fall fairly far from the parent plant.  Several years elapse before the seedlings are mature, flower-bearing plants.

Wild Garlic (Allium Ursinum)

This plant, like the Wild Hyacinth, belongs to the Lily Family and is a close relative of such common garden vegetables as the onion, garlic and leek.  Seen from a distance in early summer in the depths of a damp, shady wood, it is quite an attractive plant with its large umbels of pure white flowers and its rosette of large, shiny-green, spreading leaves.

Unfortunately, all its beauty disappears on nearer approach, as from the whole plant emanates a strong, overpowering, garlic-like odour.  I have a pressed specimen in my herbarium which was collected in 1936, but the odour is still there and can be smelt at some distance as soon as the sheet is exposed.  The strong smell and acrid juices with which the plant is permeated make it objectionable to animals and insects.

It is to be found throughout the lower parts of the Highlands, in damp oakwoods, and in those rough thickets of rowan, aspen and oak which fill the deep, rocky valleys of so many Scottish streams.  It is usually in company with such woodland plants as the wild hyacinth, primrose, and woodland grasses.

It perennates by means of a white, evil-smelling bulb sunk deep in the woodland humus.  It spring the bulb sends up several large, broad, deep green leaves which are attached to a short stalk.

>From the midst of the leaves arises a three-sided, leafless flower stalk terminated by an umbel of about twelve white flowers, each seated upon a slender pedicel.  The umbel is concealed by two white membranous scales, known as the spathe.  When the umbel expands, the spathe is pushed apart and usually falls back and hangs below the flowers.

The perianth consists of six ovate, white segments of delicate texture.  The ovary is formed of three rounded chambers surmounted by a short, awl-shaped style, and is surrounded by a ring of six stamens.

Between the bases of the stamens and ovary are nectarines which are visited by bees and flies which do not seem to be perturbed by the strong odour.  The visiting insect must align upon the centre of the flower, that is to say upon the stigman.  As it partakes of the nectar it becomes liberally dusted with pollen which it will deposit on the stigma of the next flower visited, thus effecting cross-pollination.  As the stigma lies below the level of the anthers, the flowers must often by self-fertilized.

Giant Bell-flower (Campanula latifolia)

This is one of the most handsome plants to be found in the Highlands, but unfortunately it is not a common plant and is confined to the area south of the Caledonia Canal.  It prefers rough woody glens where it is sheltered from strong winds.

It is a perennial with a thick, short stock giving rise to stout, furrowed stems which attain as much as five feet in height in luxuriant specimens.  The stem is leafy, the lower leaves being stalked, often as much as six inches long and over two inches wide, ovate-lanceolate in shape, with a serrate margin and a covering of soft hairs.  The upper leaves are sessile and decrease in size as the summit of the stem is approached.

The flowers are produced in a beautiful, long, terminal raceme, each flower being seated on a short pedicel, subtended by a leafy bract.

Each flower is a lovely structure, being deep purplish-blue in colour or pure white, and having the form of a large bell over one and a half inches in length and over one and a quarter inches in diameter at its mouth. The bell has five pointed lobes, which curve backwards towards the tips.

The interior of the bell is covered with soft hairs, which prevent small insects creeping into the flower to steal the nectar secreted by the nectarines at the base of the ovary.

The flowers are adapted to pollination by large bees, which alone are able to push into the bell-shaped corolla in spite of the hairs, and have a tongue long enough to reach the nectar.  For details of the pollination the reader is referred to the Harebell (see Chapter XIX).

Common Twayblade (Listera ovate)

The Orchid Family is well represented in the Highlands and species may be found in every type of situation.  Our present subject can be found here and there in woodlands throughout the Highland area.

It is another member of the oakwood flora and loves deep leafy humus.  On digging a plant up we should find that its roots are composed of a mass of thickish fibres.  They are remarkable from being inhabited by a mycorrhiza as in the case of many plants which inhabit humus.

This is necessary because, although humus contains a large amount of nutriment, it is not readily available to higher plants.  This is the reason why an orchid dug up from its natural haunts and planted in a garden will not flourish, the necessary fungus not being present in garden soil.

When in March or April the Twayblade breaks through the soil, it has the appearance of an Arum.  It only produces a pair of large, broadly ovate leaves during the season, but these are pressed close together, to protect the young flower stem as the plant bursts through the ground. The stem keeps growing until the leaves are about one foot above the ground, when they spread outwards.

>From between the leaves arises the tall flowering stem, which possesses no leaves, but is terminated by a very long spike of greenish flowers, each of which is fairly widely separated from the next.  Each flower has the appearance of a tiny green man, the labellum being in the form of a narrow two-lobed, hanging segment.  The upper part of the labellum is deeply furrowed and in this furrow nectar is secreted.

The mechanism of pollination in Orchids is dealt with in Chapter XX.  In this case the flowers are pollinated by small bees and flies and are no so constructed that they can only be fertilized by insect aid.  Cross-pollination only is possible, thus in the absence of insect visitors no seed to set, but this very rarely happens in nature.

Any reader who would like to know more about the amazing way orchids are adapted to insect visitors for pollination, should read The Fertilization of Orchids by Charles Darwin, in which all British orchids are fully treated.

Knotted Figwort (Scrophularia nodosa)

Our present subject, which is a common plant in the lower woodlands and shady, damp, waste places, cannot be called a lovely plant.  It has nothing to charm the eye, being tall, coarse and angular with a long, terminal spike of brown, dingy flowers.  It is even less pleasing to the nose as its leaves, stems and flowers exude a most disagreeable odour.

Below ground it possess a short, tough, perennial, knotted rhizome from which arise the stout flower stems, which may attain three feet in height.  The stem is very distinctly four-sided with prominent corners, and it is clothed with many pairs of large, stalked, ovate leaves with serrate margins.  They are usually quite glabrous and a bright, shiny green above. 

The upper part of the stem is occupied by the long panicle, consisting of a number of horizontal branches, each producing a few dull flowers. 

The plant is remarkable for the fact that the flowers are constructed for pollination exclusively by wasps.

The deep brown corolla, tinged with green towards the base, has the form of a coal-scuttle placed horizontally with the spout, which consists of two forward pointing loves, uppermost.  In the roof of the corolla is a tiny flap which projects towards the mouth.

If we examine a newly-opened flower, we shall find a conical ovary at its base, surrounded by a fleshy gland which secretes nectar, and surmounted by a long style which protrudes just beyond the corolla mouth.  The four stamens at this stage are curled up inside the corolla.

This disagreeable odour and brown colour seems to be attractive to wasps and these yellow and black banded visitors are to be seen in the large numbers around the plants.  On visiting the newly-opened flowers, it hangs on to the corolla by its curved basal rime and pushed it head inside to reach the nectar, but it is pushed downward by the flap in the roof and its thorax and abdomen rub against the stigma projecting from the base of the corolla.

In an old flower, we shall find that the stigma has bent down under the corolla where it is out of the way of visitors.  At the same time the four stamens lengthen and occupy the place originally held by the stigma.  When a wasp visit’s the flower its under body touches the stamens and becomes powdered with pollen, which it transfers to the stigma of the next newly-opened flower that it visits.

The plants are beautifully adapted to the wasps as the lower ones open first and are thus staminate when the newly-opened upper flowers are still pistillate.  The insects visit the upper flowers first and work down to the lower ones, where they become covered with pollen, for wasps work in the opposite direction to bees.  On going to the next plant, they visit the upper, pistillate flowers first and thus cross-pollination is assured.

THE WOODLAND ST. JOHN’S-WORTS

Several members of the St. John’s-wort Family (Hypericaceae) are found in the lower parts of the Highlands and three of them are woodland plants.  The Tutsan (Hypericum androsaemum) is practically confined to the western Highlands and the Hebrides and is far from common, but it does occur here and there in thickets, woods and hedges.  The Square-stalked St. John’s-wort (H. quadrangulum) is even less widely distributed and is hardly found beyond the limits of Argyll and the islands as far north as Mull. It prefers the borders of woods and damp, bushy places.  The Common St. John’s-wort (H. perforatum) has a rather irregular distribution, and is mainly restricted to the Highlands area south of the Caledonia Canal, although it does penetrate into Ross and Cromarty.  It prefers more open woods and scrubby places on dry soil, especially if limestone be present.

The Tutsan is one of those plants which once seen will never be confused with any other British plant.  It is rather fine looking, shrubby plant, with a short, woody stock giving rise to several, erect, flowering stems which may reach two feet in height.  They are covered with very large, ovate leaves often three inches in length.  The base of the leaf is heart-shaped, whilst the margin is without any trace of teeth or lobes.  The leaves which are quite smooth and shiny are arranged in pairs, one above the other, along the stem at intervals of two or three inches and they distinguish the plant from any other.  If held up to the light, they will be seen to dotted with tiny, pellucid spots.  These are sac containing resin and they are of quite common occurrence in the Hypericaceae.

The stems are terminated by small corymbs of rather small dull yellow flowers which arise in axils of two leafy bracts.  The calyx is peculiar, consisting of five green sepals of which two are large and ovate, and the other three are small and narrow.  The saucer-shaped corolla consists of five yellow petals, which are rather shorter than the sepals.

The stamens are very numerous, but if we carefully dissect the flower we shall find that they are actually united into five bundles.  At the centre of the flower is the spherical white, shiny ovary, which is surmounted by three styles.

The flowers possess no nectar and depend upon the abundant pollen produced by the large number of stamens to attract pollen-seeking bees, flies and beetles.  They are compelled to alight upon the ovary by the fence of stamens, and hence transported pollen will be left upon the styles.  Self-fertilization can occur as the stamens stand well above the styles.

The fruit is a glossy, black, berry-like capsule containing a large number of seeds, so that here again we have a woodland plant dependent upon birds for the distribution of its seeds.

The Square-stalked St. John’s-wort is a herbaceous perennial with, as its name suggests, four-sided stems.  They are clothed with small, oppoistie, ovate leaves which have numerous pellucid dots and veins and few black, glandular dots round the margins.  The flowers are produced in terminal corymbs and are rather small and of a pale yellow, and the stamens are united into three bundles.  The petals and anthers sometimes have a few black glands upon them.   This plant may be distinguished from the next by its narrow, pointed sepals and four-angled stem.

The Common St. John’s-wort is a rather similar plant, but coarser and larger and much branched.  As each branch terminates in a corymbs of bright, golden yellow flowers, the plant is quite handsome when in full bloom and is visited by numerous bees and flies for pollen.  The petals and anthers are much more frequently spotted with blank glands.  The fruit, as in the case of the preceding, is a many-seeded capsule.

THE WOOD-RUSHES          

We have already made the acquaintance of two Wood-rushes in the section on the alpine plants, and in the woods we can find two more of them, the Great Wood-rush (Luzula maxima) and the Hairy Wood-rush (L. pilosa).

The former s a fine plant and the largest of the genus.  It is to be found n all parts of the Highlands and I have found really luxuriant specimens in the wooded area of Glen Nevis.  It is a perennial and consists of a large tussock of bright green, grass-like leaves over one foot in length.  The flower stems attain over two feet in height and are terminated by a large, branching, compound panicle of small brownish flowers.

The flowers are clustered, two or three together, and consist of six scarious, brown, finely-pointed segments.  They are adapted for pollination by the wind, and to this end the style is very long so that the three spreading stigmas are produced well above the flowers where they are exposed to the wind.  Self-fertilization is avoided as the stigmas mature well before the stamens.

The Hairy Wood-rush is another common woodland plant to be found throughout the Highlands.  It is more aggressive than the preceding, as the stock is branched and tufted and gives rise to creeping runners which root and form new plants.  Thus it gradually increases its range and forms colonies often of large size.

The grass-like leaves are mainly radical, although two or three occur on the flowering stems, and are fringed with long, white, cottony hairs.  The flowering stems attain about one foot and are terminated by a panicle of pale brown flowers, the centre one being sessile, the other solitary on short, erect stalks.  The perianth segments is this plant are very narrow and finely pointed.  The flowers are wind pollinated.

THE WOODLAND GRASSES

Quite a number of grasses are to be found in the Highland woods and they contain several characteristic species.  They do not form the same dense carpets of turf as the grasses of the mountain pastures or the meadows, this being due to the much less intense light in the shade of the trees, and also to the more restricted water supply.  They are more usually found in glades, woodland borders and in open scrubby places.

Two very lovely and delicate grasses belong to the genus Melica. The Wood Melic (M. uniform) is an elegant, slender plant which supports shade very well and forms patches here and there beneath the trees.  The flowers are produced in a fine branching, panicle of a few slender branches, each producing two or three one-flowered spikes with brown or purple glumes.  The plants make a lovely sight when the breezes sweep through the woods, the delicate panicles dancing in the wind.  It is of very local distribution in the Highlands, being confined to Perthshire, Angus, Argyll and Mull.

The Mountain Melic (M. nutans) is more widely distributed than the preceding and is found in most of the central and northern counties, in both woods and on rocky hillsides.  It also is quite a slender plant and produces a one-sided panicle of from ten to fifteen spike lets, each consisting of two flowers, with brown or purple glumes.

A woodland grass, common in some places, but wanting in others, is the Creeping Soft-grass (Holcus mollis). It has an underground, creeping rhizome giving rise to tufts of leaves at the nodes and forming new plants by the decay of the intervening portion of the rhizome.  It is thus a colonizing grass and an aggressive species.  It is easily known by its leaves and stems covered with very soft, white down and hairs, and its dense, soft spike of very pale green flowers, theouter glumes of which bear short points or awns.  Each spikelet, if carefully dissected, will be seen to consist of two flowers, the upper one of which bears stamens only, whilst the lower one has both stamens and a pistil.  These details can only be made out under a lens.

Another frequent inhabitant of the woods is the Wood Poa (Poa nemoralis). It is found throughout the Highlands, but is more common south of the Caledonia Canal and is rare in the Hebrides.  It is a tufted plants which sends out short, creeping stems.  Its leaves are narrow, rather weak and slender, whilst the panicle terminates a slender flower stalk.  It is composed of very small spike lets produced along the very fine branches of the panicle.  It is thus a very graceful and airy grass and not likely to be confused with any other.

A very different grass is the Giant Fescue (Festuca gigantea), which is a stout perennial to be found in damp woods and thickets in all but the northern Highlands. It is destitute of hairs and its tall stems often reach four feet in height and are clothed with long, smooth leaves.  The drooping panicle is composed of long, pale green spike lets, each of which contains several flowers and each glume is terminated by a long, thin, hairy awn.

Another large coarse grass is the Hairy Brome (Bromus ramous). This plant may attain as much as six feet height in good soil and in sheltered positions.  The leaves are very hairy with long, hairy sheaths. 

The panicle is very large and branching, and the spikelets resemble to a certain extent those of the oat, being about one inch in length, slender and green, whilst the glumes are terminated by a stiff awn.

The Slender False Brome (Brachypodium sylvaticum) is found throughout the Highlands in woods and shrubby places.  It produces a tuft of long leaves which have a few weak cottony hairs upon their under surface.  The flowers are produced in a narrow, loose spike, each spikelet being arranged alternatively upon the central axis.  They are about one inch in length, are pale green in colour, and the flowering glumes are terminated by a long weak awn about as long as themselves.