Saxifragales

▪ plant order

Introduction

 the saxifrage order of dicotyledonous flowering plants, consisting of 16 families, 112 genera, and nearly 2,500 species. It belongs to the core eudicots, and, although its phylogenetic position is not well resolved, it is probably sister to the Rosid group in the Angiosperm Phylogeny Group II botanical classification system (see angiosperm).

Order characteristics

      Saxifragales encompasses a wide variety of plant types distributed throughout the world, including shrubs and trees, such as witch hazel and witch alder (Hamamelidaceae), rock-garden plants such as saxifrage (Saxifragaceae), familiar garden ornamentals such as peonies (peony) (Paeoniaceae), and bushes that yield currants (currant) and gooseberries (gooseberry).

      Although the order shows wide structural variations, some common traits unite Saxifragales. Flowers in the order are nearly always radially symmetric, with petals that are free rather than fused together, and the seeds typically have abundant endosperm (nutrient tissue). Most members of the order have separate (unfused) carpels, at least toward the apex. Many of the families now included in the order were once considered unrelated and were placed in three different subclasses in the earlier botanical classification systems of the American Arthur Cronquist and the Soviet-Armenian Armen Takhtajan. However, they are now supported as a monophyletic group on the basis of molecular studies. Although the internal relationships among the families are largely unresolved, this has been suggested to be the result of a rapid ancient radiation. The fact that many of the families in the order are quite small and geographically restricted offers support to the ancient radiation hypothesis, and several of the woody families have an extensive fossil history.

Major families

       Crassulaceae, or the stonecrop ( sedum) family, is the largest family in Saxifragales, with 34 genera and about 1,400 species. The family is widespread from tropical to boreal regions but is concentrated in arid regions and is characterized by very thick, fleshy leaves and by water-storage tissues in the leaves and stem. A specialized system of photosynthesis (light capture) adaptive to these kinds of conditions has been named after the family—Crassulacean Acid Metabolism (CAM). Special hairs, aerial roots, and surface cells may enable some species to absorb water directly from the air. Members of the genus Kalanchoe produce plantlets on the indentations of the leaf margins. Flowers in the family are bisexual, with the perianth parts (sepals and petals) in sets of four or five. A nectar-producing gland sits at the base of each of the 3–10 separate carpels. The family also includes widely grown ornamental perennials such as Echeveria and Crassula. The commonly cultivated Crassula ovata (jade plant) is native to South Africa. Sedum (420 species) contains many favourite rock-garden plants, and other species are grown as pot plants. Surprisingly, some species of Crassula have evolved into aquatic plants.

       Saxifragaceae, with about 33 genera and 540 species, is almost entirely herbaceous. Most species of the family grow in moist woodlands in cold northern regions. Members of the genus Saxifraga (saxifrage) (literally “rock breaker”), with their tenacious ability to thrive on rocky crags in cold, harsh climates, have a tough root system and penetrating taproots. Among the herbaceous families in the order, Saxifragaceae are perhaps the most important horticulturally. Saxifraga, Astilbe, and Heuchera contain many species cultivated in rock and woodland gardens. Flowers are usually bisexual and often somewhat showy, and the leaves vary from simple to compound and are sometimes grouped in a basal rosette. Shoots and leaves of some plants are eaten or brewed like tea. People of the Appalachian Mountains make salads with Saxifraga erosa (mountain lettuce, or deer-tongue).

 Grossulariaceae consists of a single genus, Ribes (about 150 species), many members of which bear edible fruits of commercial importance. Chief among them are R. uva-crispa, formerly called R. grossularia (European gooseberry); R. nigrum (black currant), used in jams, as a drink flavouring, and to make the liqueur cassis; R. rubrum (red currant); and R. americanum (American black currant). Other species of gooseberry and currant produce splendid shrubs with fragrant flowers and colourful fruits, and they are cultivated as garden ornamentals. Certain Ribes species act as alternate hosts to Cronartium ribicola, a parasitic fungus that causes white-pine blister rust, an extremely destructive forest disease found especially in the United States and Canada. Fungal spores that germinate on Ribes plants are blown to the pines, where the disease eventually kills the trees. In North America, eradication and strict controls have severely limited the growth of currant and gooseberry bushes in areas containing white pines (pine).

 Haloragaceae, or the water milfoil family, comprises 8 genera and 145 species of land, marsh, and water herbs with small leaves and small flower clusters. The flowers are unisexual, generally wind-pollinated, with a three- to four-chambered ovary and a similar number of styles (pollen-receptive parts at the upper end of the ovary). Representative genera are Myriophyllum (60 species), Haloragis (27 species), Laurembergia (22 species), Proserpinaca (2–3 species), Gonocarpus (1 species, Gonocarpus chinensis, or Chinese raspwort), and Meziella (1 species, Meziella trifida). Of these, Myriophyllum (water milfoil) is the principal genus of horticultural interest, mainly used in aquaria. The only shrubby genus, Haloragodendron (5 species), is native to southern Australia.

  Hamamelidaceae, or the witch hazel family, comprises 27 genera and 82 species. Its major distribution centres are East Asia and the Malaysian region, but it also extends into Australia, North America, Central America, tropical and southern Africa, and Madagascar. Members of this family thrive in moist woodlands and even on steep slopes in forested areas. Some are understory plants in forests, where they tend to be scattered, and several genera grow in the mountains of tropical and subtropical regions. Members of the family may be recognized by the presence of stellate (star-shaped) hairs and palmately veined leaves that are two-ranked—that is, lying on a horizontal plane rather than spirally arranged along the stem. The flowers are usually small, with evident perianth parts, and borne in dense clusters; the fruits are capsular with an inferior or semi-inferior ovary. Both wind and insect pollination occur in the family. Several genera are important ornamental shrubs or trees, including Hamamelis (witch hazel), Corylopsis (winter hazel), and Fothergilla (witch alder). The bark of witch hazel is often used as an astringent.

  Paeoniaceae, or the peony family, consists of the genus Paeonia, with 33 species. These are mainly native to temperate Eurasia, but there are two species native to western North America. The genus includes perennial herbs or small shrubs with large compound leaves and large showy flowers. These have a nearly continuous spiral of bracts, sepals, and petals, then numerous stamens, a nectary disc, and separate carpels with expanded stigmas. The fruits are follicles with seeds having an aril that is attractive to ants and other seed dispersers. These plants have an unusual embryology and large chromosomes (chromosome). Peonies have been cultivated since at least 900 BCE, with most herbaceous varieties derived from the Asian Paeonia lactiflora. The shrubby varieties are mostly derived from the similarly Asian P. suffruticosa.

      Iteaceae consists of the genus Itea, with 18 species. One species of Itea is native to eastern North America, while the rest are native to the area from the Himalayas to Japan and then western Malesia. The North American Itea virginica and the Chinese Itea ilicifolia are cultivated ornamental shrubs. Iteaceae have spirally arranged serrate leaves, rather small flowers with valvate petals in racemose or paniculate clusters, and capsular fruits whose two carpels remain attached by the stigma.

 Altingiaceae, or the sweet gum family, consists of one or two genera, Altingia and Liquidambar (sometimes included in Altingia). If Liquidambar is recognized as separate, it includes four species, one native to eastern North America and eastern Mexico and Central America, one from the eastern Mediterranean, and two from East Asia. Altingia then consists of about 10 species of trees, native to Southeast Asia; some of the species are timber trees that grow to heights of 60 metres (nearly 200 feet). The family is characterized by palmately lobed leaves with petiolar stipules and unisexual, spherical inflorescences.

      Cercidiphyllaceae comprises a single genus and two deciduous tree species from China and Japan. They are wind-pollinated; the flowers are unisexual and lack perianth parts, and there are separate male and female plants. Both the male and female structures that appear to be flowers are actually clusters of flowers, each flower being reduced to a few stamens or a single carpel. Cercidiphyllum japonicum ( katsura tree) is the largest deciduous tree in Japan, and it is capable of producing strong timber for house and furniture construction. During the Tertiary Period (65.5 to 2.6 million years ago), this family was widely distributed throughout the Northern Hemisphere.

Paul E. Berry

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