Table of Contents
Volume II: Perennials and Annuals
Due to the number of plants covered, listings given here cannot be comprehensive but instead
cover the major groups of plants
Sphagnum moss and Polytrichum
Ferns
Thalictrum, Buttercup, and Hellebore
Poppy and Corydalis
Silene and Carnation
Polygonum and Fallopia
Saxifrage, Astilbe, and Heuchera
Geranium and Pelargonium
Bean, Lupin, and Lathyrus
Stock and Aubrieta
Primrose and Cyclamen
Tobacco and Petunia
Pulmonaria
Mullein and Figwort
Mint, Origanum, and Sage
Campanula
Doronicum, Rudbeckia, Sunflower, and Aster
Nymphaea
Butomus
Lilium, Fritillaria, and Tulip
Orchis
Iris, Gladiolus, and Crocus
Daylily, Daffodil, and Snowdrop
Hyacinth, Muscari, and Hosta
Sedges and Grasses
Glossary
Bibliography
Index
Forewords & Introductions
The Botanical Garden Volume II: Perennials and Annuals
IntroductionOur aim in this book is to provide new information and a new way of looking at plants and gardening from a more botanical viewpoint. The plant families are covered systematically, and the relationships between them are discussed; readers will be able to put the knowledge they have acquired piecemeal into a framework, and understand the botanical groups and the similarities and differences between them.
DNA studies in plantsThe discovery of the structure of DNA by Watson and Crick in 1953 opened up a whole new method for studying the relationships between living things; much more recently, the use of computers to compare large amounts of simple data has revealed new evidence for the ancestry of plants. These new studies have not proved to be a Rosetta Stone that will reveal all, but they have provided some important new information to help solve old problems. Hitherto unsuspected relationships have been suggested, and interesting variations within a single species have also been shown up. The details of the method are complex, but depend on studying the behaviour of three different bodies within the plant cell. The DNA in two of these, the mitochondrion (involved in respiration) and the chloroplast (involved in photosynthesis), is inherited maternally, while the DNA of the third, the nucleus, is derived from both parents. The genes of mitochondria are too unstable to be of use in these studies, but fortunately the genes of chloroplasts are very stable; rearrangements of their DNA sequences are rare enough to be used to indicate major evolutionary groups, butfrequent enough to be interesting and worth looking for. Research since the 1980s is now beginning to be used to describe new relationships between genera and families, and new arrangements have been published. These sometimes confirm the classical view based on the morphology of plants, and sometimes bring surprises.
Major groupingsThe main division of the flowering plants into monocotyledons and dicotyledons is upheld by DNA studies, with the exception of a few primitive plants that fall outside both categories; this indicates that the monocotyledons arose as a group within the primitive dicotyledons, rather than separately. Some of the important groupings of monocotyledons are described below. Within the main body of the dicotyledons, around six groups are shown to be rather isolated. Two of these are the Saxifragales and the Caryophyllales, while two others, the Ranunculales and the Proteales, respectively include the largely herbaceous Ranunculaceae (see pp.38-59), Berberidaceae (see pp.60-63), and Papaveraceae (see pp.64-71), and a diverse group related to
Protea, which includes the familiar genera
Platanus and
Nelumbo: this superficially crazy association of the waterlily-like
Nelumbo, the sacred lotus, with totally different-looking trees and shrubs, must rank as one of the great surprises of DNA research. The rest of the main body of dicotyledons fall into two large clades, the Rosids and Asterids, a clade being a group of families or genera with a common ancestor, an evolutionary lineage. The artificiality of several previously recognised families of plants has been shown up by DNA studies. In particular, traditional Scrophulariaceae has been found to be a diverse assemblage, and although the traditional name has been retained here, the plants have been arranged as they naturally fall (see pp.250-63, pp.268-69, and p.277).
Unanswered questionsThe key question of what the first flowering plant looked like is still unanswered, but we are left with a number of interesting speculations. The modest water plant
Ceratophyllum (see p.385), for example, appears to be a very early offshoot from the flowering plant ancestry, but its exact position still remains uncertain; could this be what the first flowering plant was like?
Hellebore Epimedium, poppy, and CorydalisThis group of families, which includes the buttercup family, Ranunculaceae (see pp.38-59), was traditionally considered primitive, as many had very simple flowers, with indefinite numbers of separate stamens and carpels. Recent studies have confirmed this, as well as the close relationship of the Berberidaceae (see pp.60-63), which includes several distinct herbaceous genera. Poppies, the Papaveraceae (see pp.64-71), and
Corydalis, both of which often have a milky sap, are related to this group. The beautiful Japanese
Glaucidium, which was of doubtful affinity, has been shown to fall within Ranunculaeae, but the rather similar
Paeonia is now thought to be closer to Saxifragaceae (see pp.98-108).
Dianthus, cactus, polygonum, and mesembryanthemumPinks and carnations,
Dianthus, and
Silene are the main garden genera of their family, the Caryophyllaceae (see pp.74-81); several garden weeds from this family, such as chickweed and mouse-ear chickweed, are almost universal. Related to this group are the mainly succulent families Cactaceae (see p.87), which is so characteristic of the American deserts, and the Aizoaceae or Mesembryanthemaceae (see p.86), which takes its place in southern Africa; both of these families have an unusual metabolism that enables them to withstand extreme drought conditions. Other families of this group, the Caryophyllaies, are edible and furnish such vegetables as beet, spinach, and rhubarb, and such diverse ornamentals as
Limonium,
Mirabilis,
Drosera, and
Tamarix.
Saxifrage, Sedum, and peonyThe group called Saxifragales, as identified by DNA studies, is unusually diverse. The Saxifragaceae (see pp.98-108), many of which are mountain plants, are close to the usually succulent Crassulaceae (see p.109), which includes
Sedum, the stonecrop.
Paeonia, a remarkably isolated genus, whose affinities have long been in dispute, probably belongs here, although here Paeonlaceae (see pp.58-59) is listed in its traditional position next to Ranunculaceae; it appears to be closest to
Daphniphyllum. The woody plants (see Volume I) traditionally placed in Hamamelidaceae, such as
Liquidambar,
Cercidiphyllum, and
Itea, are now thought to belong here, as does
Ribes, the gooseberry and currant.
Geranium and FrancoaThe important horticultural family Geraniaceae (see pp. 112-15) includes the hardy
Geranium, found all over the world, and the tender
Pelargonium, which is mainly South African. Both of these genera have a large number of species with attractive characteristics, such as scented leaves and brightly coloured flowers, as well as large seeds that are dispersed far from the parent plant, by a spring in the case of
Geranium (or rarely as a burr), or by wind, with a silky tall, in the case of
Pelargonium. DNA indicates that the southern hemisphere plants
Melianthus and
Francoa are related to
Geranium.
Sarracenia, Primula, Phlox, and ImpatiensThe family Primulaceae (see pp.198-205), though small in number of genera, is important in temperate gardens, mainly because
Primula itself has undergone such an explosion of beautiful species in the Himalayas and the Alps. The family also includes
Anagallis and
Lysimachia. Closely related to
Primula is the mainly American
Phlox,
Polemonium, and the climbing
Cobaea, in the Polemoniaceae (see pp.196-97). These are shown by DNA studies to fall within the Asterids, and within that to belong to the Ericales, a large group that includes
Impatiens and the pitcher plants
Sarracenia, as well as shrubs such as
Styrax,
Camellia, and
Rhododendron (see Volume I).
Peas, beans, Lathyrus, and PolygalaPeas, beans, and vetches, in the family Leguminosae (see pp.124-37), are shown to be a distinct group within the Rosids. This huge family is mainly tropical and mostly woody, but the temperate Leguminosae are important in gardens, providing sweet peas and many other ornamental species of
Lathyrus, as well as peas and beans, which are vital to us for their protein-rich seeds. The capacity to fix atmospheric nitrogen is well developed in Leguminosae, and also appears to have arisen independently in a few other families. Milkwort,
Polygala, in the Polygalaceae (see p.123), a dwarf herb in the northern hemisphere but often tall and shrubby in the south, is related to Leguminosae.
Foxglove, Mimulus, and SalviaThe Lamiales, grouped around
Lamium, the dead nettle, mostly have characteristic 2-lipped, tubular flowers. The largest families are Lablatae (see pp.278-99), mostly aromatic plants including mints and sage, and Scrophularlaceae (see pp.250-263, pp.268-269, and p.277). DNA studies show that most of the genera of traditional Scrophulariaceae, such as
Antirrhinum,
Digitalis,
Mimulus, and
Nemesia, are distinct from a second group, containing
Verbascum,
Scrophularia,
Sutera, and some other small South African genera, which are thought to be closer to Gesneriaeae and Lentibulariaceae (see pp.264-65). Partially parasitic members of the former Scrophulariaceae, such as Rhinanthus, are now put in with the fully parasitic Orobanchaceae (seep.276). Even more surprisingly,
Veronica is shown to be closest to
Campsis and
Catalpa (see Volume I).
Salvia and the rest of the
Labiatae are associated with the shrubs
Callicarpa and
Clerodendrum (see Volume I), formerly in the Verbenaceae (see pp.266-67).
Daisies and CampanulaThe daisy family, Compositae (see pp.336-81) have long been considered the most advanced of the flowering plants in evolutionary terms. The flowerhead, with its numerous small, I-seeded flowers, the outer ones resembling petals, has proved very successful, especially in dry, semi-desert climates. Many species also have a parachute to disperse the seeds on the wind. In Campanulaceae (see pp.324-31),1 which is associated with Compositae in the Asterales, evolution has gone in different directions: the seeds are very small and simple, and the flowers may be large and few, as in
Canarina, which has just one flower at the end of each branch, or very small and crowded in a daisy-like head, as in
Jasione. In
Lobelia, very close to
Campanula, the flowers have become 2-lipped for more specialised pollinators, such as hummingbirds in the red-flowered American species.
Nymphaea and the most primitive flowering plantsDNA studies suggest that the most primitive of the flowering plants are a mixed group of herbs and woody plants, which include magnolias,
Laurus, and
Drimys (see Volume I). Primitive herbaceous plants include
Chloranthus,
Aristolochia,
Asarum, and
Peperomia, with Nymphaeaceae (see pp.282-83) and
Ceratophyllum the most primitive of all. Many of these plants must have survived unchanged for millions of years, while the rest of the flowering plants continued to evolve. Fossil flowers similar to
Nymphaea have been found in early Cretaceous deposits in Portugal; they are around 120 million years old. The reduced flowers of
Peperomia and the simple 3-petalled
Saruma lead into the monocotyledons, with similarities to
Acorus and
Sagittaria.
Lilium, Trillium, Alstroemeria, and ColchicumIn traditional classifications Liliaceae (see pp.416-29), the lily family, included, hundreds of genera -- almost any plant with six perianth segments, six stamens, and a superior ovary. More detailed studies, aided by DNA, have enabled smaller groupings to be recognised and the new, more narrowly defined Liliaceae is restricted to
Lilium,
Tulipa,
Calochortus,
Tricyrtis,
Prosartes, and
Scoliopus and their close relatives.
Smilax,
Trillium,
Alstroemeria, and
Colchicum are related and come within the Liliales, but most of the other former Liliaceae come within a widely designated Asparagales.
Orchids, Iris, Narcissus, and AlliumThe major group Asparagales now encompasses the old families Orchidaceae (see pp.430-33), Iridaceae (see pp.438-55) and Amaryllidaceae (see pp.464-73), as well as Alliaceae (see pp.474-81) and Convallariaceae (see pp.498-503). Relationships within the group show Orchidaceae, with its complex and advanced pollination and fungus-dependent seeds, to be closest to the simple-flowered
Astelia and
Rhodohypoxis. The closeness of Amaryllidaceae and Alliaceae is confirmed, while Iridaceae is shown to be rather isolated.
Grasses, sedges, and rushesGrasses, sedges, and rushes were traditionally put together by lovers of wild flowers, and DNA studies have confirmed their closeness, along with the burr reed
Sparganium, and
Xyris, a rush-like plant with coloured petals. A more surprising member of this group is the Bromeliaceae (see p.512), familiar to Europeans in the pineapple or as "air plants" but very frequent in the warmer parts of both North and South America, where many species are common, forming moss-like grey tufts on trees and even telephone wires. Spanish moss,
Usnea tillandsioides, is a conspicuous feature of the scenery in the southern United States and Mexico.