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extraordinary
orchids
Sandra Knapp
With a Foreword by Mark Chase
30 29 28 27 26 25 24 23 22 21 1 2 3 4 5
Preface 6
Roots of love 38
Monstrous forms 66
Interdependence 96
Postscript 154
Bibliography 156
Index 159
Foreword
I am one of those people who was passionate about orchids before I even became a botanist
and, in fact, was drawn to botany by my fascination with them. Initially I studied the history
of East Asia and grew African violets (Streptocarpus) as a hobby, but became bored with
them and started experimenting with more exotic members of that family Gesneriaceae.
Then I bought my first orchid, a Cattleya hybrid, and when it flowered just a few weeks later,
my Koehleria, Achimines and Columnea plants, beautiful as they were, had to go. My orchid
collection grew swiftly and I was compelled to switch to study biology so I could research
orchids and understand why they are so unusual. So began my lifelong entrancement.
My focus was on orchid taxonomy and my PhD on a small, mostly Mexican genus,
Leochilus. At that time in 1985, it had just become possible to carry out DNA studies, in
an emerging field of specialization called molecular systematics. This was a perfect
vehicle to answer so many questions: what are the major groups of orchids, how are
these groups related to each other, when did orchids evolve, and what other families
of plants are they related to. When I started my work none of these questions could be
satisfactorily answered. A lot has changed. Part of that major change is in the number
of orchid scientists. In the 1980s, a worldwide meeting of orchid scientists would have
numbered less than 20, but now we have at least two orders of magnitude more. Many
serious scientists had been discouraged from orchid studies by the sheer size of the
family and the lack of a scientific and phylogenetic framework on which to base their
research. Those who did study orchids were often characterized as fanatics and
dilettantes. Now those attitudes have changed, as evidenced by this book.
What we have learned about these extraordinary plants is remarkable. We now
know there are five major groups, recognized taxonomically as subfamilies:
Apostasioideae, Vanilloideae, Cypripedioideae, Orchidoideae and Epidendroideae, the
last two much larger than the first three. Orchids are members of the large order
Asparagales, which includes asparagus, daffodil, amaryllis, onion, agave, aloe, yucca,
grass tree and daylily (but not the true lilies, tulips etc. which are members of the
distantly related order Liliales). DNA data can be used as a ‘molecular clock’ to determine
the time of origin for a group of plants without a fossil record. Previously, orchids were
thought, due to the lack of a fossil record, to be recently evolved. However, by using this
DNA-based clock, we have learned that orchids are one of the oldest families of plants
FOREWORD 5
down wind gusts and providing shade, in effect creating the tropical rainforest
environment we usually think of. No more than a fraction of the light available at the top
makes it to the forest floor to be available for photosynthesis by the herbs and shrubs
of the forest understorey. But up in the canopy, when the sun shines it is unrelenting,
and when it rains there is no respite. Orchids deal with these challenges in a number of
special ways.
First, the leaves. Epiphytic orchid leaves are usually thick, stiff and somewhat fleshy;
some are even round in cross section. Many species also have swollen stems, some so
inflated that they are called pseudobulbs, or ‘false’ bulbs, because they are not buried
underground like those of tulips and daffodils, which are made up of overlapping leaf
bases rather than being stems. These thick leaves and swollen stems help orchid
plants retain water in times of drought – think of a cactus, the stem of which can store
water for long periods, and then imagine these leaves and stems performing the same
function, but high in the canopy.
Plants need water not only to grow, but also to perform photosynthesis, the reaction
in which water and carbon dioxide are converted to sugars and oxygen, much to our
benefit! The basic reaction of photosynthesis requires three main things: water, carbon
dioxide and sunlight. The energy in sunlight drives the whole process. Water is usually
taken up by plant roots, and all plants take in atmospheric gases such as carbon
dioxide through tiny pores in the leaves called stomates. The problem with these pores
is they also cause the plant to lose water via evaporation, especially if it is hot, as in the
middle of the day. Plants can lose around 97 per cent of the water they absorb through
their roots to evaporation. How can this be efficient in an environment where water
can be limiting? As a result most, if not all, epiphytic orchids use a special form of
photosynthesis called crassulacean acid metabolism (CAM), which compartmentalizes
the reactions so that the stomata can be closed during the day when water loss is a
risk, and open at night to take up carbon dioxide. CAM is a photosynthetic pathway
that was first discovered in succulents of the family Crassulaceae, which includes the
jade plant and sedums, and is also found in many other plant families living in stressful
environments. In CAM photosynthesis, carbon dioxide taken up through the stomata
at night is converted to an acid and stored in vacuoles – hollow spaces in cells – during
the hours of darkness to be then transported
to the chloroplast and reconverted to
carbon dioxide for use in photosynthesis
once the sun comes out. This specialized
form of photosynthesis allows the plants to
save water in environments where it is at a
premium, including high in the forest
canopy of the tropics.
Orchid roots are also specialized for
living in the air. If you have ever looked at
the roots of an epiphytic orchid, you might
notice they are covered by a spongy white
layer that looks almost like very thin foam.
This white layer is called the velamen, and
its structure greatly increases uptake of
water and atmospheric gases, allowing the
plant better access to both water and nutrients. The velamen is formed of layers of
cells that die as the root matures. If you look at the very tip of an epiphytic orchid root,
it is green; sometimes the velamen cells can harbour cyanobacteria – these nitrogen-
fixing bacteria could be helping the plant by providing extra nutrition in the low
nutrient environment of the forest canopy. Some of the roots of epiphytic orchids
anchor the plant to tree trunks or branches, where they form part of an epiphytic
community. And community it is – orchids are rarely the only ones up high in the
canopy. Along with orchids, the branches of tropical trees bear communities of mosses,
lichens and sometimes other epiphytic plants. These aggregations can create soil, and
even supply the tree itself with minerals that diffuse from the soil through the bark
into the host tree.
Epiphytes also create mini-ecosystems for insects and other arthropods, and
certainly contribute to the extremely high species diversity of tropical forests. The
most specialized of these ecosystems is known as an ant garden. Here, various species
of ant build their nests among the tangled roots of epiphytes and sometimes even use
orchid stems themselves as nesting places; the hollow pseudobulbs of some orchids
like Schomburgkia and Myrmecophila harbour colonies of ants, and the orchid itself
takes up nutrients from the detritus the ants accumulate. Most tropical orchids are
not obligate members of these ant garden ecosystems, but when they are, their tiny
seeds are removed and moved about by the ants, potentially to new branches and
growing sites. These complex mutualisms in tropical forests, where both partners in
the relationship derive benefit, are common and becoming ever better understood as
scientists are able to access the canopy in new ways to study how it works.
The sheer variety of form found in tropical orchids spurred a mania for orchid
collection and cultivation in the nineteenth century. The glamour and rareness of these
plants was the ultimate prize for gardeners wanting to show off their collections. Termed
‘Orchido-mania’ by James Bateman in his 1843 mega-book The Orchidaceae of Mexico
and Guatemala, this new condition ‘pervades all (and especially the upper) classes to
such a marvellous extent.’ That ‘Orchido-mania’ was especially prevalent in the ‘upper
classes’ was due to the fact that cultivating tropical orchids required a considerable
infrastructure. First, the glasshouses, then the heating, then the light. All this cost
money, and a lot of it. But in the beginning, it did not go all the way of the aristocratic
growers. Shipment upon shipment of tropical epiphytes came to Europe, and most
either died on route or were potted up carefully in heavy, rich soils, only to die later on.
It was assumed that being from the tropics, orchids needed hot and steamy
conditions, so in they went to over-heated humid glasshouses. It took a lot of
experimentation to tease out the best conditions for cultivating these seemingly
delicate plants. In the early 1800s, Joseph Banks – of the voyage of HMS Endeavour
fame and then President of the Royal Society – invented a hanging basket filled with
moss and twigs in which to grow his specimens; he was more successful than most.
The code of orchid growing was finally cracked when growers began to think
carefully about where each of these wonderful plants was found. Some, yes, were from
the hot, humid lowland tropics, but many of the new and glamorous plants coming into
cultivation were from the higher mountains of the tropics – quite a different
environment. High elevation epiphytic orchids suffer from heat more than from cold,
something which seems counterintuitive but which growers began to master. Bateman
LIFE IN THE AIR 15
ABOVE: The culture of orchids away from their native tropical forest
habitats was revolutionized by the realization that they did not need
constant water and heat. Bateman’s sketch of the ideal orchid house
was one stimulus for orchid cultivation for the masses, although he
felt orchids and the aristocracy belonged together!
LIFE IN THE AIR 17
pointed out that the orchids he was treating in his masterpiece were ‘more abundant
in the higher latitudes and purer air, than in the hot and pestiferous jungles of the
coast’ – so these tropical plants were different. He described his growing techniques in
wonderfully flowery language, and a plan for an orchid conservatory completes the
instruction. The rules were simple: ‘1st The plants can scarcely have too much light or
too much sun’, ‘2nd Take care of the roots’, ‘3rd Beware of noxious insects’, ‘4th Give
the plants a season of rest’, ‘5th Attend to the condition of the air’ and ‘6th Do not over-
water’. Pretty simple, really.
The publication of John Charles Lyons’s manual, Remarks on the Cultivation of
Orchidaceous Plants, published in 1843, the same year as Bateman’s tome, gave clear
instructions that didn’t cost the earth – orchid cultivation no longer needed to be the
preserve of the aristocracy with money to spare for glasshouses or skilled gardeners.
In the middle part of the nineteenth century, orchid cultivation became something for
(almost) everyone, sparking a revolution in cultivation. A series of articles entitled
‘Orchids for the million’ was published in the 1850s by a gardener called Benjamin
Williams at the instigation of the great Victorian orchid taxonomist John Lindley. This
manual provided even simpler instructions and eventually became The Orchid
Grower’s Manual, a set of instructions by which anyone could grow epiphytic orchids,
even in their living rooms – something we still see today.
Victorian orchid fanciers preferred the large-flowered show-offs of the orchid
world – as still seems the fashion today, given what is available for sale in greenhouses
and supermarkets – but orchids in nature are more diverse than just those we see on
the supermarket shelves. Orchids come in all shapes and sizes, from the cattleyas
with their blousy flowers to Lepanthes or Oberonia with flowers one can barely see.
Fashions in orchid cultivation in the nineteenth century favoured the showy and
magnificent, and Victorian collectors satisfied this desire with more and more new
species and genera from the tropics. But they were also collecting the less obvious
species, sending these to specialists such as John Lindley in Cambridge, whose interest
in orchids spanned the lot. So how did these collectors reach the canopy to collect
these orchidaceous rarities? How does one access an ecosystem 30 metres (98 feet) or
more in the air?
Sometimes branches fall or trees are felled; when I collected plants in the American
tropics I would follow the road builders and harvest the epiphytes from the fallen
trees, making specimens for scientific study. That was highly opportunistic though,
and not a specialized collection of orchids for cultivation. The collectors who supplied
the Victorian enthusiasts often lived in tropical areas and would employ local people
to climb, or cut down, trees to get at the plants high in the branches. Sometimes they
even used trained monkeys to access the exotic ‘air plants’ that grew out of reach. I’m
sure they, like me, also took advantage of fallen trees and branches. They probably
observed plants in situ to see if the flowers were exciting enough to warrant sending
to Europe, and maybe even cultivated them. These collectors and growers were
focusing on individual plants as commodities, not attempting to understand how they
fit into a functioning ecosystem or what they did in their everyday lives.
Real access to the canopy in order to study it as a functioning ecosystem came with
the adaptation of mountain-climbing techniques for scaling trees safely, first in the old
growth forests of the Pacific Northwest of the Americas, and later on in the tropical
rainforests. Today, the canopy can be accessed by a variety of methods, from tree-
climbing to canopy rafts and walkways. Scientists are able to study in situ how plants
and animals adapt to their environment and just how many organisms live in the
canopy itself. Using tree-fogging, entomologists estimate that each tropical tree
contains thousands of species of insects – multiply that by the number of trees, and it
becomes mind-boggling. By being at the tops of the trees themselves for more than a
fleeting moment, scientists discovered the stresses associated with living in the
canopy, and untangled the adaptations that epiphytes, including orchids, have evolved
to deal with these challenges. It is extraordinary how many challenges have been
overcome to bring us that supermarket orchid – by orchid growers over the generations
and by the orchids themselves in their life above the ground.
ABOVE: The leaves of the Australian tongue orchid, ABOVE: The Australian tongue orchid’s spidery white flowers
Dendrobium linguiforme, are egg-shaped, thick and fleshy; are produced in groups, or inflorescences, from the join of
their grooved surface and overall shape give this orchid both the leaf with the stem. The sepals are long and delicate and
its common and scientific name. The thick leaves help the give the flowers their overall appearance; the pale purple
plant conserve water during dry periods in its sclerophyll and yellow petals are smaller and less striking.
forest habitat.
OPPOSITE: The flowers of most orchids are upside down;
the pedicel or ovary twists so that the uppermost petal is
lowermost in flower. These flowers are known as resupinate.
The clamshell or cockleshell orchid, Prosthechea cochleata,
is unusual in being non-resupinate, with the dark, hooded
lower petal, called the lip or labellum, remaining as the
uppermost part of the flower, arched over the spidery
green sepals.
ABOVE: In his The Orchidaceae of Mexico and Guatemala, James
Bateman called this showy orchid, Laelia superbiens, from Mexico
and northern Central America ‘the Gorgeous Laelia’. And gorgeous
it is, with inflorescences described by its collector Mr Skinner as
‘flower stems four yards in length, and bearing upwards of twenty
flowers’. Behind the flowers here you can see the large pseudobulbs
with their yearly rings of growth, clearly showing their stem origins.
ABOVE: If you don’t look closely, the flowers of this tiny orchid, Stelis
ophioglossioides, look like mere triangles. But focus in, as the artist
Franz Bauer did in June 1825, and you can see the tiny petals held
inside the triangular outline of the sepals – an orchid in super-
miniature. This species was first discovered in the Caribbean by the
Austrian explorer Nikolaus von Jacquin and is the type species of
Stelis whose genus name refers to a mistletoe and the early, but
mistaken, idea that orchids were parasites.
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