Primulaceae, the primrose family, are herbs. The family includes 58 genera and some 2,600 species. Phylogenetic studies led to a major revision of the family, including the consolidation of the former families Maesaceae, Myrsinaceae, and Theophrastaceae into Primulaceae.
Primuloideae
There are some 900 species across nine genera in the tribe Primuloideae. They are common in the Northern Hemisphere and are scattered elsewhere. The major genera are Primula (some 500 to 600 species, including the former genera Dodecatheon, Dionysia, and Cortusa) and Androsace (about 160 species, including the former 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.
Primuloideae include a number of cushion plants as well as a floating aquatic, Hottonia, with much divided submerged leaves. Many 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.
Myrsinoideae
Formerly the family Myrsinaceae, the Myrsine tribe of Primulaceae is pantropical and temperate, especially north temperate, with species ranging 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 geographic 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 the former 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 of these 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 species, the point of insertion of the branches on the main stem is vertically elongated.
Cyclamen 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. Some genera, such as Myrsine, have male and female flowers on different plants. Several species of Lysimachia are pollinated by bees that visit the flowers to collect oils secreted by small glandular hairs. Herbaceous members provide a number of horticultural plants, of which Cyclamen is most notable.
Theophrasteae
Formerly placed as its own family (Theophrastaceae), the tribe Theophrasteae includes four genera and 90 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. Theophrasteae 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.
Maesoideae
The former 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 there are about 180 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 the former genus 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 the former genus 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, the persimmon or ebony family, includes trees and shrubs placed in four genera, with about 855 species found throughout the tropics and some also in temperate regions. Diospyros (about 800 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.