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Caryophyllales

(Order)

Overview

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Caryophyllales is an of flowering plants that includes the cacti, carnations, amaranths, ice plants, and most carnivorous plants. Many members are succulent, having fleshy stems or leaves.

Description

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The Caryophyllales includes about 6% of dicot species.[1] This order is considered a core eudicot, which is also referred to as the core tricolpates. [2] Currently, the Caryophyllales contains 33 families, 692 genera and 11,155 species. [3] The monophyly of the Caryophyllales has been supported by DNA sequences, cytochrome c sequence data and heritable characters such as anther wall development and vessel-elements with simple perforations. [2]

Circumscription

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As with all taxa, the circumscription of the Caryophyllales has changed within various classification systems. All systems recognize a core of families with centrospermous ovules and seeds. More recent treatments have expanded the Caryophyllales to include most carnivorous plants.

Although the monophyly of the order has been strongly supported, their placement is still uncertain. Systematists are undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid clade. [2] Caryophyllales is separated into 2 sub-orders: Caryophyllineae and Polygonineae. [2] These two sub-orders were formerly (and sometimes still are) recognized as two orders, the Polygonales and Caryophyllales. [2]

Apg Ii

As circumscribed by the APG II system (2003), this order includes well-known plants like cacti, carnations, spinach, beet, rhubarb, sundews, venus fly traps, and bougainvillea. Recent molecular and biochemical evidence has resolved additional well-supported clades within the Caryophyllales.

Apg

Carnegiea gigantea
Sweet William Dwarf from the family Caryophyllaceae

This represents a slight change from the APG system, of 1998

Cronquist

Chenopodium album

The Cronquist system (1981) also recognised the order, with this circumscription:

The difference with the order as recognized by APG lies in the first place in the concept of "order". The APG favours much larger orders and families, and the order Caryophyllales sensu APG should rather be compared to subclass Caryophyllidae sensu Cronquist.

A part of the difference lies with what families are recognized. The plants in the Stegnospermataceae and Barbeuiaceae were included in Cronquist's Phytolaccaceae. The Chenopodiaceae (still recognized by Cronquist) are included in Amaranthaceae by APG.

New to the order (sensu APG) are the Asteropeiaceae and Physenaceae, each containing a single genus, and two genera from Cronquist's order Nepenthales.

Earlier Circumscriptions

Earlier systems, such as the Wettstein system, last edition in 1935, and the Engler system, updated in 1964, had a similar order under the name Centrospermae.

Photos

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Taxonomy

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The Order Caryophyllales is further organized into finer groupings including:

Families

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Achatocarpaceae

Shrubs or small trees. Branchlets spinescent, sometimes with scattered thorns. Leaves alternate, sometimes fasciculate, exstipulate; blade spatulate to oblanceolate to elliptic, margins entire, venation pinnate. Inflorescences axillary racemes or panicles, or flowers solitary; bracts 0.5-1 mm; bracteoles present or absent. Flowers unisexual, staminate and pistillate on same plant; perianth simple; tepals 4-5, imbricate, distinct, herbaceous. Staminate flowers: stamens [10-]12-14[-20], exserted; filaments filiform, 1-1.5[-2] mm; anthers basifixed, oblong, ex-trorsely dehiscent. Pistillate flowers: pistils 2-carpellate; ovary 1-locular; style absent; stigmas 2, filiform. Fruits berries. Seed 1, lenticular, without true endosperm; embryo annular, surrounding perisperm.[1] [more]

Agdestidaceae

[more]

Aizoaceae

[more]

Amaranthaceae

Herbs, clambering subshrubs, shrubs, or lianas. Leaves alternate or opposite, entire, exstipulate. Flowers small, bisexual or unisexual, or sterile and reduced, subtended by 1 membranous bract and 2 bracteoles, solitary or aggregated in cymes. Inflorescences elongated or condensed spikes (heads), racemes, or thyrsoid structures of varying complexity. Bracteoles membranous or scarious. Tepals 3-5, membranous, scarious or subleathery, 1-, 3-, 5-, or 7(-23) -veined. Stamens as many as tepals and opposite these, rarely fewer than tepals; filaments free, united into a cup at base or ± entirely into a tube, filament lobes present or absent, pseudostaminodes present or absent; anthers (1- or) 2-loculed, dorsifixed, introrsely dehiscent. Ovary superior, 1-loculed; ovules 1 to many; style persistent, short and indistinct or long and slender; stigma capitate, penicillate, 2-lobed or forming 2 filiform branches. Fruit a dry utricle or a fleshy capsule, indehiscent, irregularly bursting, or circumscissile. Seeds lenticular, reniform, subglobose, or shortly cylindric, smooth or verruculose.[2] [more]

Barbeuiaceae

[more]

Basellaceae

Vines herbaceous or herbs twining, usually fleshy, glabrous. Leaves simple, alternate, usually petiolate, margin entire. Inflorescences of spikes, racemes, or panicles; bracts 3, caducous; bracteoles 2, persistent. Flowers bisexual, actinomorphic, small. Perianth segments 5, white or reddish, distinct or connate at base, imbricate in bud, persistent in fruit. Stamens 5, opposite to petals; filaments inserted on perianth. Ovary superior, 1-loculed; ovule 1, basal, campylotropous. Pistil united from 3 carpels. Style simple with 3 stigmas or 3 free styles. Fruit a utricle, dry or fleshy, often surrounded by persistent bracteoles and perianth. Seed globular; testa membranous; endosperm copious; embryo spirally twisted or semicircular to horseshoe-shaped.[3] [more]

Cactaceae

Fleshy perennials, shrubs, trees or vines, terrestrial or epiphytic. Stems jointed, terete, globose, flattened, or fluted, mostly leafless and variously spiny. Leaves alternate, flat or subulate to terete, vestigial, or entirely absent; spines, glochids (easily detached, small, bristlelike spines), and flowers always arising from cushionlike, axillary areoles (modified short shoots) . Flowers solitary, sessile, rarely clustered and stalked (in Pereskia), bisexual, rarely unisexual, actinomorphic or occasionally zygomorphic. Receptacle tube (hypanthium or perianth tube) absent or short to elongate, naked or invested with leaflike bracts, scales, areoles, and hairs, bristles, or spines; perianth segments usually numerous, in a sepaloid to petaloid series. Stamens numerous, variously inserted in throat and tube; anthers 2-loculed, dehiscing longitudinally. Ovary (pericarpel) inferior, rarely superior, 1-loculed, with 3 to many parietal (rarely basal) placentas; ovules usually numerous; style 1; stigmas 2 to numerous, papillate, rarely 2-fid. Fruit juicy or dry, naked, scaly, hairy, bristly, or spiny, indehiscent or dehiscent, when juicy then pulp derived from often deliquescent funicles (except in Pereskia) . Seeds usually numerous, often arillate or strophiolate; embryo curved or rarely straight; endosperm present or absent; cotyledons reduced or vestigial, rarely leaflike.[4] [more]

Caryophyllaceae

Herbs annual or perennial, rarely subshrubs or shrubs. Stems and branches usually swollen at nodes. Leaves opposite, decussate, rarely alternate or verticillate, simple, entire, usually connate at base; stipules scarious, bristly, or often absent. Inflorescence of cymes or cymose panicles, rarely flowers solitary or few in racemes, capitula, pseudoverticillasters, or umbels. Flowers actinomorphic, bisexual, rarely unisexual, occasionally cleistogamous. Sepals (4 or) 5, free, imbricate, or connate into a tube, leaflike or scarious, persistent, sometimes bracteate below calyx. Petals (4 or) 5, rarely absent, free, often comprising claw and limb; limb entire or split, usually with coronal scales at juncture of claw and limb. Stamens (2--) 5--10, in 1 or 2 series. Pistil 1; carpels 2--5, united into a compound ovary. Ovary superior, 1-loculed or basally imperfectly 2--5-loculed. Gynophore present or absent. Placentation free, central, rarely basal; ovules (1 or) few or numerous, campylotropous. Styles (1 or) 2--5, sometimes united at base. Fruit usually a capsule, with pericarp crustaceous, scarious, or papery, dehiscing by teeth or valves 1 or 2 × as many as styles, rarely berrylike with irregular dehiscence or an achene. Seeds 1 to numerous, reniform, ovoid, or rarely dorsiventrally compressed, abaxially grooved, blunt, or sharply pointed, rarely fimbriate-pectinate; testa granular, striate or tuberculate, rarely smooth or spongy; embryo strongly curved and surrounding perisperm or straight but eccentric; perisperm mealy.[5] [more]

Chenopodiaceae

Herbs annual, subshrubs, or shrubs, rarely perennial herbs or small trees. Stems and branches sometimes jointed (articulate) ; indumentum of vesicular hairs (furfuraceous or farinose), ramified (dendroid), stellate, rarely of glandular hairs, or plants glabrous. Leaves alternate or opposite, exstipulate, petiolate or sessile; leaf blade flattened, terete, semiterete, or in some species reduced to scales. Flowers monochlamydeous, bisexual or unisexual (plants monoecious or dioecious, rarely polygamous) ; bracteate or ebracteate. Bractlets (if present) 1 or 2, lanceolate, navicular, or scale-like. Perianth membranous, herbaceous, or succulent, (1-) 3-5-parted; segments imbricate, rarely in 2 series, often enlarged and hardened in fruit, or with winged, acicular, or tuberculate appendages abaxially, seldom unmodified (in tribe Atripliceae female flowers without or with poorly developed perianth borne between 2 specialized bracts or at base of a bract) . Stamens shorter than or equaling perianth segments and arranged opposite them; filaments subulate or linear, united at base and usually forming a hypogynous disk, sometimes with interstaminal lobes; anthers dorsifixed, incumbent in bud, 2-locular, extrorse, or dehiscent by lateral, longitudinal slits, obtuse or appendaged at apex. Ovary superior, ovoid or globose, of 2-5 carpels, unilocular; ovule 1, campylotropous; style terminal, usually short, with 2(-5) filiform or subulate stigmas, rarely capitate, papillose, or hairy on one side or throughout. Fruit a utricle, rarely a pyxidium (dehiscent capsule) ; pericarp membranous, leathery, or fleshy, adnate or appressed to seed. Seed horizontal, vertical, or oblique, compressed globose, lenticular, reniform, or obliquely ovoid; testa crustaceous, leathery, membranous, or succulent; embryo annular, semi-annular, or spiral, with narrow cotyledons; endosperm much reduced or absent; perisperm abundant or absent.[6] [more]

Didiereaceae

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Gisekiaceae

[more]

Halophytaceae

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Hectorellaceae

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Illecebraceae

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Molluginaceae

Herbs annual or perennial, subshrubs, or shrubs, rarely dioecious, glabrous or rarely hairy. Stems erect or prostrate. Leaves simple, alternate, rarely opposite, often in a basal rosette or in pseudowhorls on stems, margin entire; stipules absent or membranous. Inflorescences terminal or in seemingly axillary cymes, rarely as a solitary flower. Flowers bisexual, rarely unisexual, actinomorphic, hypogynous, rarely perigynous. Tepals 5, rarely 4, free or connate below into a tube, lobes white or pink to purple, sometimes yellow inside (in Glinus) . Petals absent or few to many, white, pink, or purple. Stamens 3-5 or many, arranged in several rings, free or connate at base in bundles; anthers dehiscing by longitudinal slits. Ovary superior, syncarpous (in Gisekia developmentally syncarpous; fruit a deeply lobed schizocarp) ; carpels 2-5 or many, placentation axile, rarely seemingly basal. Stigmas as many as locules. Ovules 1 to many per locule. Fruit usually a loculicidal capsule or deeply (3-) 5-15-lobed mericarps (in Gisekia), rarely breaking into 2 nutlets [in Limeum Linnaeus, not in Flora area]. Seeds with embryo curved around a hard, starchy perisperm.[7] [more]

Nyctaginaceae

Herbs, shrubs, trees, or sometimes spiny vines. Leaves opposite, alternate, or whorled; stipules absent; petiole usually present, well defined; leaf blade simple, herbaceous or slightly fleshy, margin entire. Inflorescences mostly terminal, less often axillary, of cymes, umbels, or verticils, sometimes 1-flowered or fasciculate, often grouped into panicles; bracts often inconspicuous, sometimes forming calyxlike involucre, or large and brightly colored. Flowers bisexual, rarely unisexual or polygamous, actinomorphic. Perianth constricted beyond the ovary, base persistent, closely enclosing ovary which appears inferior, limb petaloid beyond constriction, tubular, funnelform, or campanulate, apex 5-10-lobed, lobes plicate or valvate in bud, persistent or caducous. Disk absent. Stamens (1-) 3-5(-many), hypogynous, free or connate at base, involute in bud; anthers 2-loculed, dehiscence longitudinal. Ovary superior, 1-loculed; ovule 1. Style 1; stigma globose. Fruit an achenelike anthocarp enclosed by persistent perianth, ribbed or winged, often glandular. Seed 1; endosperm present; embryo straight or curved.[8] [more]

Petiveriaceae

[more]

Phytolaccaceae

Herbs or shrubs, rarely trees, erect, rarely scandent, mostly glabrous, rarely dioecious. Leaves simple, alternate, entire; stipules absent or tiny. Inflorescences terminal, axillary, or leaf-opposed, racemose, cymose, panicled, or spicate. Flowers small, bisexual or rarely unisexual, actinomorphic, rarely zygomorphic. Tepals 4 or 5, persistent, free or connate at base, equal or unequal, imbricate in bud, green or sometimes other colored. Stamens 4 to many, inserted on a fleshy disk; filaments usually persistent, free or slightly connate at base; anthers 2-loculed, dorsifixed, longitudinally dehiscent. Ovary superior, globose; carpels 1 to many, free or connate; ovule solitary in each carpel, basal, campylotropous. Styles persistent, short or absent, erect or curved, as many as carpels. Fruit fleshy, a berry or drupe, rarely a capsule. Seeds reniform or oblate, small; testa membranous or hard and fragile, smooth or wrinkly; embryo large, curved, surrounding copious mealy endosperm.[9] [more]

Portulacaceae

Herbs annual or perennial, rarely ± shrubby, usually succulent, usually glabrous except for nodal hairs and/or scales. Leaves alternate or opposite; true stipules absent, nodes sometimes with axillary scales and/or hairs; petiole usually poorly defined or absent; leaf blade simple, usually fleshy, margin entire. Inflorescences usually terminal, less often axillary, in cymes or racemelike panicles, forming heads of sessile flowers surrounded by an involucre of leaves, or reduced to solitary flowers. Bracts inconspicuous. Flowers