Ericales

▪ plant order

Introduction

  rhododendron order of flowering plants, containing 25 families, 346 genera, and more than 11,000 species.

      The relationships of the order are unclear. It belongs to neither of the two major asterid groups (Asterids I or Asterids II), but with Cornales it is basal to the core asterid clade of flowering plants in the Angiosperm Phylogeny Group II (APG II) botanical classification system. The families included in it have previously been placed in several other orders, including Theales. The circumscription of the order and its placement in the asterids are something of a surprise, although both are well supported by morphological and molecular data.

      Morphologically, the order is rather generalized and unspecialized for the asterids. Most members have at least weakly fused petals and radially symmetric flowers (flower), a superior 3- or 5-locular ovary, and 5 or 10 (sometimes more) stamens that are often borne free of the petals. The fruit is most often a capsule, and the seed coat is usually thin. Iridoids, distinctive chemicals perhaps involved in protection of the plant against herbivores, are scattered through the order.

      Molecular studies suggest that there are eight family groupings in Ericales, plus one isolated family ( Theaceae). The Ericaceae group contains Ericaceae, Clethraceae, and Cyrillaceae. The Balsaminaceae group contains Balsaminaceae, Marcgraviaceae, and Tetrameristaceae (including Pelliciera). The Polemoniaceae group contains Polemoniaceae and Fouquieriaceae. The Pentaphylacaceae group contains Pentaphylacaceae (including Ternstroemiaceae) and Sladeniaceae. The Styracaceae group contains Symplocaceae and Diapensiaceae. The Lecythidiaceae group includes only Lecythidaceae. The Primulaceae group contains Primulaceae, Theophrastaceae, Myrsinaceae, Maesaceae, Sapotaceae, and Ebenaceae. The Sarraceniaceae group contains Sarraceniaceae, Actinidiaceae, and Roridulaceae. These groupings and their member families are described in this article.

Ericaceae group

      The Ericaceae group contains Ericaceae, Clethraceae, and Cyrillaceae, which are characterized by having a hollow style as well as by features of the endosperm, or seed reserve, and stem anatomy.

Ericaceae

 Ericaceae, or the heath family, is the largest family in Ericales, containing more than a third of its genera (128) and species (about 4,000), with many new species still being discovered. Members of the family occur mainly in tropical montane, Mediterranean, and temperate to arctic climates. Major genera include Rhododendron (about 850 species, including azalea, Ledum, Menziesia, and Tsusiophyllum), Erica (almost 800 species, including Philippia and other small genera), Vaccinium (about 400 species), Gaultheria (235 species, including Pernettya), Leucopogon (about 230 species), and Cavendishia (155 species).

 In Ericaceae the petals are usually fused, and the corolla is often urn-shaped or tubular; the 10 stamens often release their pollen by pores or short slits, and they sometimes have appendages on the anthers or the tops of the filaments. The pollen is often in groups of four grains, or tetrads. The ovary is usually divided into five parts.

      Ericaceae prefer acidic habitats, well-lit conditions, and a temperate climate. Most of the family live in close association with fungi (fungus), which inhabit the roots; however, the fungal hyphae do not penetrate the cells. In fact, the fungi facilitate the plants' uptake of nutrients, especially nitrogen, an element often in short supply in the acidic habitats that the family favours. The overall result is that Ericaceae, although a large family, has a rather distinct ecology. Some species, such as Monotropa ( Indian pipe), lack chlorophyll and completely depend on their fungi for nutrients. These fungi are also associated with trees, and nutrients move from the trees to the ericaceous plant via the fungi. Different species of fungi are associated with different achlorophyllous Ericaceae, a much closer association than elsewhere in the family.

      Many Ericaceae have xeromorphic leaves, which apparently evolved for life under conditions of limited water (or nutrient) availability. Erica, in particular, have leaves that are small and narrow, with a channel on the lower surface, while other genera have leaves that are thick and hard, with close parallel venation. Both groups are abundant in Mediterranean climates, and many species have starch-rich tubers or burls and resprout after fire. Many of the genera related to Vaccinium are lianas (liana) or epiphytes (epiphyte) (grow on other plants) in the montane tropics. Their leaves are often thick, and the base of the stem can be very swollen.

 Ericaceae often have large and conspicuous flowers, and a variety of relationships with pollinators have been demonstrated. Rhododendron and some of its relatives have viscin threads, very fine structures that hold the pollen tetrads together in clumps. Large insects, and in the tropics birds and perhaps bats, are the main pollinators by dislocating strands of threads in which the pollen tetrads are entangled. Erica and its relatives have smaller flowers, but, even so, birds and a variety of insects (including flies in South Africa) are pollinators (pollination); some species of Erica are pollinated by wind, as are Empetrum (crowberry) and its relatives. Many of the large-flowered Vaccinium relatives are pollinated by birds. Less-common forms of pollination include buzz pollination (bees dislodge pollen by bugging), as in some species of Vaccinium, including Vaccinium oxycoccus ( cranberry). Most species of kalmia have the anthers held under tension, being released only when the pollinator lands, so pollination here is explosive.

      Ericaceae with capsular fruits have wind-dispersed seeds, and wings or tails of various sorts on the seed may aid in this process. Fleshy-fruited Ericaceae are dispersed mostly by birds, sometimes by mammals, with the hard seeds being excreted by the animal. Most species of Gaultheria combine the two methods of dispersal, having a dry, capsular fruit surrounded by succulent and brightly coloured sepals. Some Vaccinium relatives have seeds with a mucilaginous covering, and they may be dispersed more like mistletoe. These species also have green embryos and are either epiphytic or epilithic (grow on rocks).

      In summary, there are perhaps four lines of ecological specialization in Ericaceae, which are to a remarkable extent mutually exclusive. About 1,300 species have xeromorphic leaves; about 1,500 species have fleshy fruits; about 900 species have pollen with viscin threads; and about 400 species are either epiphytic or epilithic.

      Ericaceae are much valued in horticulture, especially in temperate regions, for their showy flowers. However, a number of species of Vaccinium, in particular, are cultivated for their fruits. A few (e.g., Rhododendron ponticum) are locally seriously invasive.

Clethraceae

      Clethraceae contains two genera. Clethra, with 75 species, grows from East Asia to Malesia (biogeographic region), in the southeastern United States, and from Mexico southward along the Andes; a single species grows on the Atlantic island of Madeira. All are woody deciduous to evergreen plants with spiral, often toothed leaves that tend to be clustered at the ends of branches. They have racemose inflorescences (inflorescence) at the ends of the branchlets and rather small flowers. These flowers have only basally fused (or free) sepals and petals and 10 late-inverting stamens with distinctive arrow-shaped anthers that open by pores. The fruit has three compartments. Purdiea includes about 12 species native to the Caribbean, Central America, and northern South America. Its flowers are considerably larger and more showy than those of Clethra.

Cyrillaceae

      Cyrillaceae is a small family of two genera of trees or shrubs that grow in the Caribbean region, from the southeastern United States to northern South America and the West Indies. Cyrillaceae have spirally arranged, toothless leaves, with short petioles, long-racemose inflorescences, and rather small flowers. The flowers appear to be have separate petals; the anthers are arrow-shaped; the stigma is often three-lobed; and the fruit has three compartments. The number of species in Cyrilla is disputed, with some botanists recognizing a single widespread and variable C. racemiflora and others recognizing up to 10 or 12 distinct species.

Balsaminaceae group

      The Balsaminaceae group members—Balsaminaceae, Marcgraviaceae, and Tetrameristaceae (including Pelliciera)—are distinct from the rest of the order, although they are highly variable in growth and appearance. All have cells with bundles of needle-shaped crystals of calcium oxalate and lack a nectary disc. They have a short style, and their petals do not form a tube. Marcgraviaceae and Tetrameristaceae have leaves that elongate while still folded in bud, and there are faint lines on the lower surface of the expanded leaf where the edges of the leaves are pressed against the surface.

Balsaminaceae

 Balsaminaceae, or the touch-me-not family, includes 2 genera and about 1,000 species of fleshy herbs. Hydrocera, with one species, is Indo-Malesian, while Impatiens (touch-me-not genus), with all the other species, grows throughout the family range, which is mostly Old World—mainly Africa (especially Madagascar) to the mountains of Southeast Asia. Balsaminaceae are rather fleshy herbs that have toothed leaves and strongly zygomorphic, spurred flowers, with the five stamens closely arching over the ovary. The spur is a modified sepal, and the flower is held upside down. The stamens make a cap over the stigma and may get knocked off by the pollinators, usually bees. Nectar is secreted in the spur, and the showy flowers usually have three petal-like sepals and five petals; two pairs of petals are joined at their bases. The rather large seeds are thrown some distance when the capsule dehisces explosively, as it does when it is touched.

Marcgraviaceae

      Marcgraviaceae are often lianas or epiphytes and are found only in the Neotropics. There are 7 genera and about 130 species in the family, of which Marcgravia includes 60. The family has often rather thick leaves with indistinct venation and inflorescences with flower bracts that are modified as flask-shaped nectaries. The stamens are often quite numerous. The fruits, with many small seeds exposed on a fleshy, brightly coloured placenta, are distinctive.

      Heterophylly (different leaf types on the same plant) is common in Marcgravia. The climbing form of the plant has small leaves without stalks. The leaves are arranged in two rows on the branches and are pressed against the trunk of the tree on which the plant is growing. Short adventitious (aerial) roots develop along these shoots and enable the plant to climb. The upper shoots, which bear pendulous flower clusters at their ends, have much larger stalked, spirally arranged leaves and lack the adventitious roots of the climbing stems. (Similar growth patterns occur in other climbers, including some Aracaeae.) The transition between shoot types is often abrupt, although the cause is unknown; it is thought that increasing light received by the shoots as they climb may be involved. Pollination in Marcgraviaceae is mostly by birds and bats, which take nectar from the modified bracts. In the simplest case a bract subtends each flower. In Marcgravia, however, the flower cluster is pendulous and umbellate—the flowers are on stalks that radiate from a common point, like an umbrella. The central flowers of the umbel are sterile, and their bracts are enlarged to form erect, pitcherlike structures that are superficially similar to the insectivorous pitchers of Nepenthes, which hang below the outer ring of fertile flowers. Nevertheless, birds sometimes avoid brushing the stigma when taking nectar from these bracts, and self-pollination may occur. Birds and sometimes mammals eat the fleshy fruits of Marcgraviaceae.

Tetrameristaceae

      Tetrameristaceae have glands on the inner surfaces of the sepals and only a single ovule in each part of the ovary. The flowers have only five stamens. Pelliciera rhizophorae are evergreen trees in mangrove vegetation on the Pacific or, rarely, Atlantic coast of Central America and northern South America. They have long pointed terminal buds, spirally arranged toothed leaves with asymmetrical bases and short stalks, and large flowers in the axils of leaves. The large sharply pointed single-seeded fruits are also distinctive. Until its affinity with Tetrameristaceae was recognized, Pelliciera had been its own family.

      The other two genera of Tetrameristaceae are rather small woody plants. The three species of Tetramerista grow in west Malesia. The one species of Pentamerista grows in the Guiana Highlands of Venezuela. These genera have spiral, short-stalked leaves with indistinct venation and marginal glands. Tetramerista has glistening dots on the inner surface of both calyx and corolla. Both genera have fleshy fruits, which are presumably dispersed by animals.

Polemoniaceae group

      The Polemoniaceae group includes Polemoniaceae and Fouquieriaceae. There are few distinctive morphological characters that unite them. Both have strongly fused petals, a three-lobed stigma, and capsular fruits. Both families are diverse in southwestern North America. Polemoniaceae also shows a close connection at the generic level between western North America and southern South America, which is known from other plants of the drier, warmer habitats of those areas. Fouquieriaceae is restricted to the deserts of the southwestern United States and northwestern Mexico.

Polemoniaceae

      Polemoniaceae, or the Jacob's ladder or phlox family, includes about 20 genera and nearly 400 species of mostly herbaceous plants, which are mainly north temperate in distribution, although they also occur in South America. Phlox (70 species) is largely North American, but there is one species in northeastern Asia. Polemonium (27 species) is predominantly north temperate but grows at higher elevations in Mexico and in Chile. Some genera, such as Gilia (25 species), are mainly western North American, but several, such as Linanthus (35 species), Navarretia (30 species), and Collomia (15 species), also have a few species in southern South America. Polemoniaceae are herbs, or sometimes shrubs or lianas, with opposite leaves, and the plant may smell unpleasant. The sepals are fused, and each often has a green central portion, the rest being whitish. The corolla has a long tube, and the five stamens are frequently inserted at different levels in the tube, or they may be of different lengths. The seeds are mucilaginous when wetted.

      The radiation of Polemoniaceae in southwestern North America is associated with much floral diversification and pollination by a variety of birds, bats, and insects, including beetles and butterflies, although some of the annuals self-pollinate. There is a group of vines and shrubs that is more tropical in distribution, including Cobaea. The seeds are dispersed by wind, although they may stick to animals when they are wetted.

Fouquieriaceae

 Fouquieriaceae, or the ocotillo family, are shrubs that grow in drier parts of western North America. There is a single genus, Fouquieria (including Idria), with 11 species. They are often little-branched shrubs with spirally arranged leaves. Leaves on some shoots are borne close together (short shoots), while those on others are well separated (long shoots) and often have the petioles modified as spines after the rest of the leaf has fallen. The flowers have free sepals but a strongly fused corolla; the 10 or more stamens are free from the corolla. Species of Fouquieria are pollinated by hummingbirds or bats; the seeds are wind-dispersed.

Pentaphylacaceae group

      Members of the Pentaphylacaceae group—Pentaphylacaceae (including Ternstroemieae) and Sladeniaceae—all have only unicellular hairs; the sepals and petals are basically free; there is usually no nectary; and the capsular fruit has a central column. The group is mainly tropical and subtropical.

Pentaphylacaceae

      Pentaphylacaceae have rather short filaments, and the embryos are curved. The smallish flowers are usually borne singly in the leaf axils or in some modification of this. The three groups in this family were previously placed in different families.

      The first group includes a single genus (Pentaphlyax), with a single species (P. euryoides) that is scattered from Sumatra to China. It has spirally arranged evergreen leaves with entire margins (smooth, without teeth). In the flowers the pollen sacs appear to be borne transversely on the stout filaments, and they open by flaps. There are only two ovules in each ovary locule, and the midribs of the valves separate from the rest as the capsule opens.

      The second group consists of subfamily Ternstroemieae, with two genera of evergreen shrubs to trees that is especially abundant in Malesia, Central America, and South America. Ternstroemia (more than 100 species) is pantropical, although it has only a few species in Africa.

      The third group, subfamily Frezierieae, includes 9 genera and more than 230 species. Eurya (75 species) occurs from Asia and Malesia to the western Pacific. Adinandra (75 species) is Indo-Malesian. Freziera (57 species) is entirely American. The leaves in the family are often two-ranked and toothed and may remain rolled up as they elongate, so the lower surface of the blade has longitudinal markings. The flowers also occur in clusters in the leaf axils. The fruit is usually a berry.

      Ternstroemieae have evolved fleshy, animal-dispersed fruits. Eurya and Freziera, in particular, tend to grow in montane and disturbed conditions; both genera have male and female flowers growing on different plants. Ternstroemia has leaves inserted all around the stem; they lack teeth and occur only at the end of each growth increment. The genus looks quite different from other members of the group.

Sladeniaceae

      Sladeniaceae are trees that grow in semitropical conditions. The family includes two genera: Sladenia (two species) grows from southern China to Burma, and Ficalhoa (one species) grows on the mountains of East Africa. The family has toothed leaves and distinctive cymose inflorescences in the axils of the leaves. The flowers are small, lack stalks, and have more or less free sepals and petals. The two genera differ considerably in details of their flowers and fruit. Their current geographical distributions may be deceptive, since fossil wood apparently of Sladenia has been found from northeast Africa.

Styracaceae group

      The link between Styracaceae and Diapensiaceae is most obvious when DNA sequences are compared. However, in both families there are close relationships between members growing in widely separated areas in the Northern Hemisphere. Both have rather conspicuous insect-pollinated flowers and capsular fruits with small, wind-dispersed seeds. Diapensiaceae have often been thought to be close to Ericaceae, but this is not supported by molecular studies.

Styracaceae

 Styracaceae, or the silver bells family, are evergreen or deciduous trees or shrubs of warm north temperate to tropical regions, including Malesia, North America, and South America. There are some 11 genera and 160 species in the family. Styrax (about 120 species) is by far the largest genus, occurring throughout much of the family range. Rehderodendron (nine species), Halesia (five species), and Alniphyllum (three species) are concentrated in Southeast Asia and Malesia. Styracaceae characteristically have stellate or scaly hairs, spirally arranged toothed leaves, and often rather bell-shaped flowers with a more or less completely fused and/or small calyx, and a more or less inferior ovary.

      Styracaceae also have winged fruits that are dispersed as wholes or, rarely, as drupes (Parastyrax). Styrax, in particular, can develop the most elaborate galls that superficially look like fruits of Annonaceae. A number of Styracaceae are cultivated for their flowers, which are usually white. Styrax also yields a medicinally important resin.

Diapensiaceae

 Diapensiaceae is a small family with 6 genera and 18 species. All are perennial herbs or subshrubs that grow in the Arctic and north temperate region, especially in East Asia and the eastern United States. Diapensia (four species) is circumboreal, with the other genera being much more localized. For example, Galax (two species) grows only in the eastern United States; Berneuxia (one species) grows only in the Himalayas; Shortia (including Schizocodon, six species) grows in the eastern United States, where it is rare, and in East Asia. The family has moderate-sized flowers, whose petals are only weakly fused into a tube that is sometimes held together by the flattened filaments of the five stamens, which alternate with the petals. The anthers are incurved. There are often five sterile stamens, or staminodes. The stigma is shortly three-lobed, and the fruit is a capsule with small seeds. The family is of rather minor importance in the horticulture trade, although the leaves of Galax are sometimes used in salads.

Symplocaceae

      Symplocaceae is a group of tropical to subtropical evergreen trees. There is a single genus, Symplocos, with about 320 species that grow in North America, South America, Southeast Asia, Indo-Malesia, and especially New Caledonia. The toothed leaves of Symplocos often dry yellowish because the plants tend to accumulate aluminum. The racemose inflorescences have rather small flowers that look rather like those of Rosaceae (rose family). The petals are fused only at the base; there are often many stamens; and the ovary is inferior. The fruit is fleshy, with a stone.

Lecythidaceae

 Lecythidaceae, or the Brazil nut family, is a pantropical group of evergreen trees of about 25 genera and 310 species. There are several groups in the family with distinctive geographical distributions. The Brazil nut group includes about 10 genera and 215 species, all Neotropical; in particular, the group includes the larger genera Eschweilera (about 100 species) and Gustavia (40 species). An Old World group of 6 genera and 58 species includes Barringtonia (40 species), which grows from eastern Africa to the Pacific. Three smaller groups include 9 genera and 49 species; they occur in South America and Africa, and one includes genera that were until recently placed in Scytopetalaceae.

      Members of the Brazil nut family usually have spiral leaves borne in tufts at the ends of the branches, or the leaves may be two-ranked. The margins are often serrate or minutely stipulate or both. The flowers are often large, with free petals or what appear to be petals but are really modified stamens, and numerous functional stamens (up to 1,200) that are free to fused. The ovary is more or less inferior.

      The great diversity of flowers is accompanied by a great diversity in pollinators and pollination mechanisms. Many Neotropical Lecythidaceae have complex flowers in which the stamens are fused and form a hood covering the ovary. Large bees force their way into the flower to reach the nectar in the centre or to collect a special pollen. Bats and small bees also pollinate Neotropical Lecythidaceae. Malesian members have flowers with the many stamens radiating like pins in a pincushion; at least some are pollinated by bats. Monkeys eat the fleshy parts of the seeds or fruits of many Neotropical Lecythidaceae, although others have their fruits dispersed by wind, water, fish, birds, or scatter-hoarding rodents. Bats and other mammals probably disperse the fruits of Barringtonia and its relatives. There is great variation in the morphology of both the embryo and the seedling.

      Lecythidaceae includes a number of ornamental trees. Bertholletia excelsa (Brazil nut tree) has nutritious oily seeds (not nuts) with very thick coats; the woody fruits have to be smashed open by collectors to free the seeds. The wood of Scytopetalum tieghemii is used in Sierra Leone and Ghana for house poles because of its resistance to decay.

Primulaceae group

      The Primulaceae group contains Theophrastaceae, Myrsinaceae, Primulaceae, Maesaceae, Sapotaceae, and Ebenaceae. The first three families have long been considered to be closely related, but details of the relationships between them became clearer after DNA studies. These families have much in common, including chemistry and small glandular hairs. Secretory canals containing yellow, red, or brown tannins, for example, are frequent. The petals are fused into a tube, and the five stamens are opposite the petals, rather than alternating with them, as is usual. The style is often short, and the ovary is not divided by partitions; the placenta, on which the ovules are borne, is very much swollen.

      Theophrastaceae, Myrsinaceae, and Primulaceae all have one or more genera that are herbs with rather small, rotate corollas; that is, the flowers have a fairly short, narrow corolla tube and a spreading limb. Indeed, it is possible that the woody members of the first two families have evolved from a plant of this kind. The three woody families all have some members with tufts of large leaves at the ends of the branches, the plant itself even being unbranched and with a stout stem. Species like this are found in Maesa (Maesaceae), Clavija (Theophrastaceae), and Discocalyx (Myrsinaceae). Insect pollination is common in the order.

Theophrastaceae

      Theophrastaceae includes 6 to 9 genera and 105 species of mostly shrubs and small trees that are largely restricted to the New World. Samolus (15 species) is the only herbaceous genus, and it also grows in Europe and the Antipodes. Jacquinia (35 species) is Central American and Caribbean, while Clavija (50 species) grows in both Central and South America. Theophrastaceae have petal-like staminodes (nonfunctional stamens) borne on the corolla tube opposite the sepals. In several woody members the anthers form a cone in the centre of the flower when it opens, but they later spread. The result is that there are very distinct male and female phases. Most have tough, more or less whorled leaves with toothed or spiny margins or a sharp apex.

Myrsinaceae

 Myrsinaceae, or the Myrsine family, is pantropical and temperate, especially north temperate, with species from trees to herbs. There are about 41 genera and 1,435 species in the family. Ardisia (about 450 species) is found in much of the family's range but not in Africa. Myrsine (155 species, including Rapanea and Suttonia) is pantropical to warm temperate. Lysimachia (150 species) is mostly herbaceous and temperate. Discocalyx (115 species, including Tapeinosperma) grows from Malesia to the Pacific. Embelia (100 species) grows in the Old World. Parathesis (85 species) and Stylogyne (60 species) are restricted to the Americas. Anagallis (28 species) occurs in Europe, Africa, and South America, and there is one widespread species. Cyclamen (about 20 species) is found from Europe to Iran and Somalia. Many Myrsinaceae species have distinctive yellowish to blackish dots or streaks on the often spiral leaves (and often obvious on the persistent calyx and on the fruit). The ovary is superior, and the seeds are often rounded. In many woody Myrsinaceae, the point of insertion of the branches on the main stem is vertically elongated.

      Cyclamen, which used to be included in Primulaceae, has a swollen underground storage structure called a corm that can live for more than a century. Aegiceras is a mangrove plant, and its seeds have embryos much larger than those in other members of the family. In Myrsinaceae such as Myrsine, male and female flowers are on different plants. Several species of Lysimachia are pollinated by bees that visit the flowers to collect oils secreted by small glandular hairs. Herbaceous Myrsinaceae provide a number of horticultural plants, of which Cyclamen is most notable.

Primulaceae

 Primulaceae, or the primrose family, are herbs with perhaps 9 genera (their limits are currently unclear) and some 900 species. They are common in the Northern Hemisphere and are scattered elsewhere. The major genera are Primula (some 500 to 600 species, including Dodecatheon, Dionysia, and Cortusa) and Androsace (about 160 species, including Douglasia and Vitaliana). Primulaceae are usually rosette herbs with a scapose inflorescence—that is, there are no leaves along the inflorescence stalk—and medium-sized flowers with fused sepals and spreading petals. The fruit is a capsule with many angular seeds.

 Primulaceae include a number of cushion plants, as well as a floating aquatic, Hottonia, with much divided submerged leaves. Many Primulaceae species have heterostylous flowers; that is, some plants have a long style and short stamens (pin flowers), while in others the relationship is reversed (thrum flowers). This promotes cross-fertilization. The distinctive flowers of Dodecatheon ( shooting star), with its recurved petals and anthers forming a central cone, are visited by bees, which dislodge pollen from the anthers by buzzing (buzz pollination); this “genus” is really only a Primula modified for and adapted to buzz pollination. Primula, in particular, is commonly cultivated.

Maesaceae

      Maesaceae are evergreen lianas to shrubs or trees found in the Old World tropics to Japan, the Pacific, and Australia; there is one genus, Maesa, and about 150 species. The veins of the leaves are often not very obvious, even when the leaf is dry, but there are well-developed and conspicuous secretory canals. The small flowers are urn-shaped and have an inferior ovary. The fruit is fleshy, but there is also a stony layer; there are many angular seeds.

Sapotaceae

 Sapotaceae is a largely tropical family of evergreen trees and shrubs. There are 53 genera and about 1,100 species in the family, but generic limits in the family are notoriously difficult and changeable. Pouteria (200–305 species, including Planchonella), Chrysophyllum (80 species), Manilkara (80 species), and Mimusops (40 species) are found throughout the humid tropics. Palaquium (120 species) grows from Southeast Asia to the Pacific, while Madhuca (110 species) is Indo-Malesian. Sideroxylum (75 species, including Dipholis) grows in the Americas and Africa to the Mascarenes, while Micropholis (38 species) is confined to the New World.

      Sapotaceae have rather small, naked terminal buds with adpressed, brownish, often T-shaped unicellular hairs. The axillary branches often have a prominent pair of buds at the very base, then long internodes, and finally a tuft of leaves. The leaf blades tend to have rather closely parallel secondary veins, and their margins have no teeth. The twigs usually exude copious gutta or latex. The flowers, with their persistent sepals, fused petals, stamens as numerous as the petals and opposite them, and often protruding style, are distinctive, as are their seeds, which are large and have a thick, shiny, brown seed coat with a very large pale-coloured scar.

      Sapotaceae species have smaller and apparently much simpler flowers than those of Lecythidaceae. However, there is much variation in the number and lobing of petals and the presence and nature of staminodes, although simply urn-shaped flowers are common. Little is known of the family's pollination, although bats and insects of various sizes are suspected. Some taxa lack nectaries, and the sweet and fleshy corolla may be eaten by the pollinator, so providing a reward for it. The fleshy fruits of Sapotaceae are dispersed by bats and various mammals, including monkeys, and by birds and even fish (some species of Pouteria in the Amazon).

      Latex of Sapotaceae is a source of gutta-percha, balata, and chicle, either pure trans-polyisoprene polymers or a mixture of cis and trans constituents. The berries of a number of species are edible.

Ebenaceae

 Ebenaceae, or the persimmon or ebony family, includes trees and shrubs placed in four genera, with about 490 species found throughout the tropics and some also in temperate regions. Diospyros (about 500 species) occurs throughout the family's range. Ebenaceae often have two-ranked leaves that lack teeth but have flat, dark-coloured glands on the lower surface. The flower buds often have adpressed, brown, T-shaped hairs and are often pointed; the petals are fused at the base, and their lobes overlap regularly. The sepals commonly increase considerably in size in fruit, which is a rather large-seeded berry. The bark, even of twigs, is black with a yellow undersurface; the heartwood is also black and the leaves too may dry blackish.

      Species limits in Diospyros are difficult to delineate, but in parts of the Asian tropics many clearly very different species grow together. Flowers are unisexual, usually with male flowers on different plants from the females. Pollination is mainly by insects, with dispersal by birds and mammals that eat the berries, but few details are known. Lissocarpus, which used to be placed in its own family (Lissocarpaceae), has bisexual flowers with an eight-lobed corolla tube and an inferior ovary.

      Species of Diospyros are of economic importance for the wood that several produce and for their fruits. The wood, which is either uniformly dark (ebony) or variously streaked and marbled, has been much used in furniture making. The fruit (date plums, persimmons) can be very astringent if eaten before they are fully ripe.

Sarraceniaceae group

      The Sarraceniaceae group is made up of Roridulaceae, Sarraceniaceae, and Actinidiaceae. Members of the group have racemose inflorescences with at least medium-sized, pendulous flowers. The stamens initially face the outside of the flower, but they invert during development, and the anthers end up facing inward; the anthers often open by pores or short slits. The style is sunken into the apex of the ovary, and the fruit is a capsule with many small seeds.

Sarraceniaceae

 Sarraceniaceae, or the pitcher plant family, are insectivorous herbs in 3 genera with about 15 species. Darlingtonia, with a single species, grows in California; Sarracenia, with eight species, is found in southern and eastern North America, while Heliamphora, with 5–10 species, occurs only in the Guiana Highlands in northern South America. All are pitcher plants. Their pitchers are long with short petioles and are borne in a rosette. The flowers are quite large, with free rather petal-like sepals, free petals, and usually numerous stamens; the style is either unbranched or like an umbrella.

      The hollow tubular leaves of Sarraceniaceae are shaped like urns, trumpets, or small pitchers. They are exquisitely constructed pitfalls that entrap insects lured to the mouth of the pitcher by nectar-secreting glands and glistening surfaces. Downward-pointing hairs in the throat of the pitcher prevent the insect's escape, and the exhausted prey slide down the slippery throat and fall into the liquid in the pitcher, where they are either digested by enzymes secreted by glands in the pitcher or eaten by the animals living in the pitcher, their remains being excreted into the liquid. Like most other insectivorous plants, Sarraceniaceae live in acidic, boggy habitats.

Roridulaceae

      Roridulaceae contains a single genus, Roridula, with two species of small southern African shrubs. They have linear leaves that are covered with capitate, resin-secreting hairs. The flowers are medium-sized with free sepals and petals and only five stamens that invert early in their development. Although Roridula also appears to be insectivorous, its long leaves being covered by sticky hairs, the leaves are not rolled up in buds, nor are the hairs sensitive (as in Droseraceae, Drosophyllaceae, and Byblidaceae). There is as yet no evidence of insectivory.

Actinidiaceae

      Actinidiaceae are usually shrubs, small trees, or lianas; they are largely tropical and especially abundant from Southeast Asia to Malesia. There are 3 genera and 355 species in the family. Saurauia (300 species) grows throughout the range of the family, while Actinidia (some 30 species) is Indo-Malesian and East Asian. The leaves of Actinidaceae are spiral and often strongly toothed. The flowers appear to be perfect, but they are usually either male or female. They have apparently free sepals and petals and many stamens that invert just before the flower opens; the anthers dehisce by a slit that is much broader toward the apex. The fruit is usually a berry. kiwi fruit comes from species of Actinidia.

Theaceae

 Theaceae, or the tea family, includes 7 genera and about 200 species of shrubs and trees that grow mostly in the Southeast Asia–Malesia region, although a few species are found in the southeastern United States. Camellia (some 120 species) grows from Southeast Asia to Indo-Malesia. Pyrenaria (42 species) grows in Southeast Asia and western Malesia. Stewartia (9 species) grows largely in East Asia, mostly China, with a few in the southeastern United States. Franklinia, with 1 species from Georgia in the United States, is now extinct in the wild.

      Theaceae have quite thick and toothed leaves that often turn red just before falling. The flowers are usually quite large, with free sepals and petals. There are numerous stamens with long filaments. The fruit is usually a capsule with a persistent central column, and the seeds are often flattened.

      Bees are probably major pollinators in Theaceae, with pollen being the main reward. Pseudopollen is also known from the family. This consists of cells from the inside of the anthers that are about the same size as the pollen but with very different cell-wall thickenings. Its exact role in pollination is unclear. Franklinia matures its fruit during the summer of the year after it flowers. Seed dispersal is largely by wind.

      The single most economically important plant in Ericales is certainly Camellia sinensis, the leaves of which are the source of tea. C. sinensis is a native of Assam (a state in northeastern India), but it was first cultivated by the Chinese, from whom India and Ceylon (now Sri Lanka) imported seeds. C. japonica and other species and hybrids are the source of the many cultivars of Camellia that are grown for their flowers, while the genus also yields a valuable oil from the seed variously used as a hair oil, in soap, or for cooking or lubricating. Several other genera of Theaceae are cultivated for their flowers or bark, including Stewartia and Franklinia.

James L. Luteyn Peter Stevens

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