Euphorbia

Herbs, subshrubs, or shrubs [trees, cactoid succulents, geophytes, vines], annual, biennial, or perennial, monoecious [dioecious]; hairs unbranched or absent; latex white. Leaves persistent, deciduous, or small and caducous proximally, alternate, opposite, or whorled, sometimes bractlike and subtending floral structures, simple; stipules absent or present, persistent or deciduous; petiole absent or present, glands absent; blade unlobed, margins entire, crenulate, crenate-dentate, or serrulate, laminar glands absent; venation palmate, palmate at base and pinnate distally, or pinnate, often only midvein prominent. Inflorescences bisexual [unisexual], terminal or axillary, pseudanthia (each consisting of cuplike involucre bearing glands on rim, these sometimes with petaloid appendages, enclosing solitary pistillate flower surrounded by (0–)1–80 staminate flowers, entire structure termed the cyathium), in monochasia, dichasia, pleiochasia, cymose clusters, capitate glomerules, or solitary; glands subtending each bract 0. Pedicels present. Staminate flowers: sepals 0; petals 0; nectary absent; stamen 1; pistillode absent. Pistillate flowers: sepals 0 (ovary subtended by a calyxlike structure in E. floridana, E. inundata, E. mesembrianthemifolia, E. porteriana, E. rosescens, and E. telephioides); petals 0; nectary absent; pistil 3-carpellate; styles 3, distinct or connate basally to most of length, unbranched or 2-fid. Fruits capsules, tardily dehiscent and with spongy mesocarp in E. lathyris [drupes]. Seeds globose to ovoid, oblong, cylindric, deltoid, pyramidal, or bottle-shaped; caruncle present or absent. x = 6, 7, 8, 9, 10.

Distribution

North America, Mexico, West Indies, Bermuda, Central America, South America, Eurasia, Africa, Atlantic Islands, Indian Ocean Islands, Pacific Islands, Australia.

Discussion

Species ca. 2000 (139 in the flora).

Euphorbia is one of the two or three most species-rich angiosperm genera worldwide. Members of the genus occur in almost all habitat types, and many species prefer disturbed areas. Species in the genus are vegetatively highly diverse; growth forms include diminutive ephemerals, tuberous geophytes, taprooted perennial herbs, vines, various types of shrubs, trees to 25 m tall, and many xerophytic stem-succulents. Although succulents are primarily restricted to the Old World, a handful of independently derived succulents are native to the New World (B. L. Dorsey et al. 2013). Within the flora area these include E. antisyphilitica and E. tithymaloides. The striking vegetative similarity between the Old World succulent Euphorbia and New World cacti is one of the most commonly cited examples of convergent evolution.

The most distinctive feature of Euphorbia is its unique pseudanthial inflorescence, the cyathium (G. Prenner and P. J. Rudall 2007). This structure is so similar in appearance to a bisexual flower that many early botanists, including Linnaeus, actually mistook it for one. As with all Euphorbiaceae, the flowers of Euphorbia are unisexual, but in contrast to most other members of the family, they are extremely reduced. The pistillate flower comprises a single, perianth-less ovary, and the staminate flower comprises a single perianth-less stamen. Most species are likely insect-pollinated, but a few species are hummingbird-pollinated (R. L. Dressler 1957).

The latex of all of the species is abundant and in some instances highly caustic, and care should be taken to avoid exposure to it. Nevertheless, some species in the flora area have been used for medicinal purposes (for example, Euphorbia corollata and E. ipecacuanhae, C. F. Millspaugh 1892).

Euphorbia is best known for its ornamental taxa, in particular E. pulcherrima Willdenow ex Klotzsch (the Christmas poinsettia), a native of Mexico. This species is widely grown throughout the flora area but has not become naturalized. Other commonly cultivated species in the flora area include E. milii Des Moulins (crown-of-thorns), E. rigida M. Bieberstein, E. characias (Mediterranean spurge), E. marginata (snow-on-the-mountain), and E. antisyphilitica (candelilla); the last two are native species. Euphorbia “Diamond Frost” has in recent years become a popular cultivar to grow in pots or flowerbeds. Its progenitor, E. graminea, is introduced in the flora area and appears mainly associated with plantings in several southern states. In addition to several herbaceous European species that have become naturalized in North America, several others that have been recorded in the flora area in the past appear not to be persistent. These include species such as E. amygdaloides Linnaeus, E. epithymoides Linnaeus (sometimes treated as E. polychroma Kerner), E. lucida Waldstein & Kitaibel, E. paralias Linnaeus, and E. segetalis Linnaeus. In addition to these leafy taxa, numerous succulent species are commonly cultivated in botanical gardens and by private growers. Among the most popular are E. lactea Haworth, E. neriifolia Linnaeus, E. obesa Hooker, E. tirucalli Linnaeus, and E. trigona Miller. Although some of these species may persist around areas where they were previously cultivated, there is no evidence that they are actually naturalized in the region.

One of the most troublesome noxious weeds in the northern part of the flora area is leafy spurge, which was introduced from Eurasia. This species has been widely treated in North America as Euphorbia esula Linnaeus, but it turns out to be a misapplication of that name. The true leafy spurge in North America is more appropriately treated as E. virgata, a weedy species that is broadly distributed throughout temperate Europe and Asia (D. V. Geltman 1998). The actual E. esula is a related species of more restricted distribution in Europe that lacks the weedy tendencies of E. virgata (see discussion under 124. E. virgata for characters that distinguish the two). As with some of the herbaceous European waif species mentioned above, the real E. esula has been recorded historically in different parts of the flora area, but it does not appear to have persisted. It is therefore excluded here, and this should help to dispel the incorrect application of that name to leafy spurge in North America.

Historically, distinctive clades within Euphorbia were segregated into a number of satellite genera. In the flora area, these include Chamaesyce, Pedilanthus, and Poinsettia. Although these segregate genera are morphologically well-defined and monophyletic assemblages, recent molecular phylogenetic research has demonstrated that they are all nested within a broadly defined Euphorbia (V. W. Steinmann and J. M. Porter 2002; J. W. Horn et al. 2012). These molecular analyses both show Euphorbia as comprising four distinct clades, each of which is treated as a subgenus. Three of these subgenera, subg. Chamaesyce (Gray) Caesalpinius ex Reichenbach, subg. Esula Persoon, and subg. Euphorbia, are represented in the flora area. Subgenus Esula is treated here as a morphologically cohesive unit, whereas subg. Chamaesyce and subg. Euphorbia are divided into well-defined sections, which are keyed out and treated below.

The pleiochasia in Euphorbia are determinate. Each bears a whorl of pleiochasial bracts, which subtends multiple dichasial cymes, termed pleiochasial branches, that arise from a common point. The pleiochasium usually is terminated by a cyathium, but that sometimes aborts.

Selected References

Dorsey, B. L. et al. 2013. Phylogenetics, morphological evolution, and classification of Euphorbia subgenus Euphorbia. Taxon 62: 291–315.

Horn, J. W. et al. 2012. Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae). Molec. Phylogen. Evol. 63: 305–326.

Prenner, G. and P. J. Rudall. 2007. Comparative ontogeny of the cyathium in Euphorbia (Euphorbiaceae) and its allies: Exploring the organ-flower-inflorescence boundary. Amer. J. Bot. 94: 1612–1629.

Lower Taxa

Euphorbia sect. Alectoroctonum, Euphorbia sect. Anisophyllum, Euphorbia sect. Crepidaria, Euphorbia sect. Nummulariopsis, Euphorbia sect. Poinsettia, Euphorbia subg. Esula

Key

1	Stems usually prostrate, sometimes erect, ascending, reclining, or decumbent; leaves opposite (rarely whorled in E. fendleri), blade bases usually asymmetric; stipules interpetiolar (except in E. acuta where at base of petiole, deciduous, sometimes appearing absent).	Euphorbia sect. Anisophyllum
1	Stems usually erect or ascending, rarely decumbent or prostrate; leaves alternate, opposite, or whorled, blade bases symmetric; stipules at base of petiole or absent.	> 2
2	Stems semisucculent to succulent, zig zag; involucres strongly zygomorphic, spurred and forming tube that encloses glands.	Euphorbia sect. Crepidaria
2	Stems not both succulent and zig zag; involucres ± actinomorphic, not spurred.	> 3
3	Involucral gland appendages usually petaloid, occasionally rudimentary; leaf margins entire.	Euphorbia sect. Alectoroctonum
3	Involucral gland appendages not petaloid (except in E. bifurcata, E. eriantha, and E. exstipulata in sect. Poinsettia, but then leaf margins usually toothed); leaf margins entire or toothed (teeth sometimes inconspicuous in E. eriantha with linear leaves).	> 4
4	Cyathia in terminal monochasia, dichasia, or condensed pleiochasia; involucral glands shallowly cupped to deeply concave, 1–3 per cyathium (if 4–5 in E. exstipulata then involucral gland appendages present; if 4–5 in E. radians then terminal clusters of cyathia subtended by white to pinkish, leafy bracts; if 4–5 in E. eriantha, then involucral gland appendages fringed, canescent, and folded over glands); involucral gland appendages petaloid, fringed, or absent.	Euphorbia sect. Poinsettia
4	Cyathia in pleiochasia; involucral glands slightly concave, flat, or slightly convex, 4–5 per cyathium (2–3 in E. oblongata), terminal clusters of cyathia never subtended by white to pinkish bracts; involucral gland appendages hornlike or absent.	> 5
5	Ovary and capsule not subtended by calyxlike structure; seeds with caruncle; involucral gland appendages hornlike or absent.	Euphorbia subg. Esula
5	Ovary and capsule subtended by calyxlike structure; seeds without caruncle; involucral gland appendages absent.	Euphorbia sect. Nummulariopsis