Passifloraceae Juss. ex Roussel

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Vines, trailing or climbing by means of axillary unbranched tendrils, herbaceous or woody, or rarely erect shrubs or trees

Morphology Leaves

Leaves alternate, simple, entire or lobed, or very rarely compound, the petiole usually bearing glands; stipules 2, thread-like or large and ovate.

Morphology Reproductive morphology Flowers

Flowers axillary, perfect or rarely unisexual, solitary or variously cymose- racemose or paniculate; peduncles usually jointed; bract and bracteoles 3, small and distant from the flower, or large, foliaceous and close to the flower, forming an involucre

Morphology Reproductive morphology Flowers Calyx

Calyx-tube (receptacle) almost flat to saucer-like or bell-shaped, giving rise in the middle to a gynophore (ovary-stalk), and nearly always bearing toward the margin or within its throat a ring of one or several series of erect or radiate filamentous or membranous outgrowths (called the corona); sepals usually 5, inserted in the throat of the receptacle, overlapping, often with a small horn-like process on the back near the apex, and usually coloured inside

Morphology Reproductive morphology Flowers Corolla

Petals either lacking or usually 5, alternate with the sepals, free, overlapping, and withering-persistent

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens usually 5, attached below the ovary on the gynophore

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior and free, located at the apex of the gynophore, 1-celled; style simple or 3–4-branched, or else 3 separate styles present; ovules numerous, attached to 3 or 5 parietal placentas

Morphology Reproductive morphology Fruits

Fruit berry-like or sometimes a capsule opening by 3 apical valves; seeds numerous, usually compressed-ovoid, covered with a fleshy aril or surrounded by pulp; endosperm fleshy; cotyledons often foliaceous.

Distribution

A widespread tropical and subtropical family of about 12 genera and 600 species, especially concentrated in S. America.

Passifloraceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:1. 1954

Morphology General Habit

Herbaceous climbers bearing tendrils, or erect trees or shrubs

Morphology Leaves

Leaves alternate, entire or lobed, often with a glandular petiole; stipules usually small and deciduous

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual

Morphology Reproductive morphology Flowers Calyx

Sepals 5, imbricate, persistent, free or partially united

Morphology Reproductive morphology Flowers Corolla

Petals 5, rarely absent, free or shortly united, imbricate

Morphology Reproductive morphology Flowers Corona

Corona of one or more rows of thread-like processes or scales, or annular

Morphology Reproductive morphology Flowers Androecium

Stamens 5 or more, hypogynous or perigynous, shortly united or in bundles, sometimes springing from the gynophore; anthers 2-celled, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-celled, with 3 or rarely 4–5 parietal placentas; ovules usually numerous; styles free or united; stigmas often capitate

Morphology Reproductive morphology Fruits

Fruit a capsule or berry, indehiscent or loculicidally 3-valved

Morphology Reproductive morphology Seeds

Seeds with pitted testa, surrounded by a pulpy aril; endosperm fleshy; embryo large, straight

Timothy Utteridge & Gemma Bramley (2020). The Kew Tropical Plant Families Identification Handbook, Second Edition. Kew Publishing Royal Botanic Gardens, Kew

Note

Climbers with tendrils, herbs or shrubs. Leaves alternate. Flowers actinomorphic; sepals and petals alternating; extrastaminal corona; stamens. on androgynophore or hypanthium; ovary superior. Fruit a berry or loculicidal capsule. Seeds arillate.

Recognition

Characters of similar families: Convolvulaceae: tendrils absent. Cucurbitaceae: tendrils 90˚ to petiole base, unisexual flowers, inferior ovary. Passifloraceae (Turneroideae): petals contorted; corona absent or much reduced; stigmas brush-like.

Morphology General Habit

Climbers with axillary tendrils, or erect herbs, shrubs or small trees

Morphology Leaves Stipules

Stipules usually present, often small, sometimes foliaceous, caducous

Morphology Leaves

Leaves alternate; simple, seldom compound; lamina variable, lobed or unlobed; venation pinnate, palmate or pedate; petioles or leaf base often with nectaries

Morphology Reproductive morphology Inflorescences

Inflorescences axillary, usually cymose, or racemose

Morphology Reproductive morphology Flowers

Flowers usually bisexual and actinomorphic (rarely zygomorphic), hypanthium saucer-shaped to tubiform; sepals (3–)5(–8); petals as many as and alternating with sepals; extrastaminal corona of 1 or more whorls of scales or processes of varying width, or tubular or cup-shaped, sometimes absent; nectary extrastaminal, annular or of 5-parts; stamens (4–)5 or 8(–numerous) borne on androgynophore or hypanthium; ovary superior, (2–)3(–5) carpels, unilocular, sessile or on elongate androgynophore; placentation parietal; styles (1–)3–5, free or partially united; stigmas capitate to subglobose, sometimes divided

Morphology Reproductive morphology Fruits

Fruit a berry or loculicidal capsule

Morphology Reproductive morphology Seeds

Seeds few to many, surrounded by membranous or mostly pulpy aril.

Distribution

Following APGIII we include Turneraceae and Malesherbiaceae in Passifloraceae here, but we present only the most species-rich Passifloroideae subfamily (16 genera, c. 800 species). The largest and most well-known genus is Passiflora (c. 625 species, mostly New World); Adenia (100 species) is also sizeable in the Old World, particularly Africa. Several Passiflora species are used as attractive garden plants; others are grown for their fruits or juice.

Description Author

Gemma Bramley and Sara Edwards

Nunes, T. (2009). Neotropical Passifloraceae.

Morphology

Description

Climbing herbs or lianas with axillary tendrils, shrubs or trees, rarely forked or with adhesive discs at apices of tendrils. Leaves simple or rarely compound , alternate , stipulate (rarely absent), petiole glandless or with glands (2-10); laminas unlobed or 2-3-5-9- lobed , edge entire or serrate , with or without glands , ocellus usually present. Inflorescence axillary , cymose, sessile or pedunculate, primary axis often a tendril , secondary axis often reduced; bracts linear , lanceolate or foliaceous , sometimes glandular ; pedicels articulate . Flowers actinomorphic (rarely zygomorphic ), hermaphrodite , functionally unisexual; perianth persistent , segments free or partially fused at base, usually with a cup-shaped to tubular hypanthium, red, pink, mauve; purple, blue, green or white (rarely yellow); sepals (3-)5(-6), petals equal to sepals, imbricate , rarely absent; corona with one to many series of filaments; stamens 4(-5), inserted on androgynophore ; anthers dorsifixed; ovary (1-)3(-5)-carpellate, superior , unilocular, sessile or stipitate on elongate androgynophore ; placentation parietal ; ovules many; styles (1-)3(-5), free to partially fused at base; stigmas globose , capitate , papillate or divided . Fruit a berry or loculicidally 3-5-valved capsule . Seeds single to numerous, compressed , sub - globose , reticulate , foveolate, pitted or laterally grooved (3)6(11); endosperm copious; cotyledons foliaceous . Germination epigeal or hypogeal. Several species night flowering and pollinated by bats.

Distribution

Distribution in the Neotropics

• Mainly in tropical and subtropical areas, particularly diverse in the New World tropics.

Diagnostic

Distinguishing characters (always present)

• Vegetatively easily recognizable by the alternate leaves; tendrils present.
• Flower with corona with one to many series of filaments.
• Ovary unilocular, many ovules; placentation parietal.
• Seeds globose, compressed, reticulate, pitted or transversely grooved.

Key differences from similar families

• Passifloraceae can't be confused with any other family of the Malpighiales because of the presence of the filamentous corona and the androgynophore, combined with the presence of tendrils and extrafloral nectaries. 
• It is very close to the Turneraceae and Malesherbiaceae, but differs by the flowers possessing a corona, consisting of 1 or more rows of filaments, usually borne on a hypanthium; ovary usually on a short to long gynophore; often vines with tendrils, arilfleshy.
• Differs from Cucurbitaceae by usually simple leaves, opposite to the tendril; stipules lacking, petioles always without glands. Inflorescences determinate, sometimes reduced to a single flower, axillary.

Other important characters

• Leaves alternate; simple or lobate (3-9-lobate).
• 3 styles; 5-10 stamens; ovary tricarpellate, unilocular.

Notable genera and distinguishing features

• Mitostemma: Restricted to Brazil and Guiana. Scandents shrubs; flowers usually in short, terminal or axillary racemes; calyx much reduced; sepals 4; petals 4; corona in 3 series of filaments; stamens 8 or 10, inserted on the floor of the hypanthium near the base of the ovary, free, or united close to the base; gynophoreerect; ovary 1-celled, with 4 parietal placentae; styles 4, distinct to the base; stigmas reniform-capitate; fruitovoid.
• Dilkea: Confined to the Amazon (Peru and Brazil). Woody vines or small trees, without tendrils, or rarely with a few poorly developed ones; flowers in axillary or terminal glomerules or short-spicate, rarely solitary, hermaphrodite, red or white; sepals 4-5; petals 4-5; corona with 2 series of filaments; stamens, hypogynous, free except at the very base; ovary subsessile or short-stipitate, with 4 parietal placentae; styles 4, united below the middle; stigmas reniform-capitate; fruitglobose or ovoid.
• Passiflora: Distributed throughout the tropics and subtropics, although the vast majority are endemic to the New World. Can be distinguished by the straight gynophore (rarely curved in P. mucronata and P. galbana); stamens that are free from their tips to the gynophore, and 3 styles; corona in various series; hypanthium well-developed; stigmas capitate; flowers mostly pentamerous.

General Description

Number of genera

14 genera with 4 occuring in the Neotropics:

• Ancistrotryrsus Harms (2 spp.).
• Dilkea Mast. (6 spp.).
• Mitostemma Mast. (3 spp.).
• Passiflora L. (including Tetrastylis Barb. Rodr., c. 450 spp.).

General notes

• Some species have medicinal benefits and have been used in folk medicine (Passiflora incarnata) - in infusion of leaves to combat various diseases, P. edulis - diuretic and inflammations; P. incarnata - sedative; P. quadrangularis - analgesic).
• Other species are known for their economic value such as fruit juice (P. edulis, P. alata, P. cincinnata, P. platyloba, P. maliformis, P. foetida, P. tarminiana, P. ligularis, P. quadrangularis), and ornamental because of the nature of the exotic nature of their flowers (P. racemosa, P. caerulea, P. quadrangularis, P. amethystina).
• The bark of P. suberosa and P. holosericea are used as a vermifuge, the flower buds and flowers of P. alata are made into a tea-like drink and taken as a vermifuge or sedative.

Status

• All genera are Native of Neotropics.
• Several species are cultivated throughout the Neotropics because of their edible fruits and the beauty of their flowers (P. caerulea, P. cincinnata, P. racemosa, P. amethystina, P. kermesina, P. quadrangularis, P. alata, P. maliformis, P. laurifolia, P. ligularis).

Notes on delimitation

• The Passifloraceae is placed in the Malpighiales, by DNA-sequences of the atpB and rbcL gene, together with 28(-30) families which also occur in the Neotropics, including the Euphorbiaceae, Flacourtiaceae, Malpighiaceae, Rhizophoraceae, Salicaceae and Violaceae.

Literature

Important literature

ANGIOSPERM PHYLOGENY GROUP [APG.] 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II.Botanical Journal of the Linnean Society 141: 399-436.

Brizicky, G.L., 1961. The genera of Turneraceae and Passifloraceae in the southeastern United States. Journal of the Arnold Arboretum, Cambridge, 42:204-218.

Escobar, L.K. 1988. Passifloraceae. Flora da Colômbia. Univ. Nac. de Colombia  10: 1-143. Falta da 71-143.

Feuillet, C. & MacDougal, J.M. 1997. New infrageneric names in Passiflora (Passifloraceae). Biollania, Edición Especial 6: 335-340.

Killip, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19(1-2):1-613.

Ulmer, T. & MacDougal, J.M. 2004. Passiflora: Passionflowers of the world. Timer Press, Portland.

Passifloraceae, W. J. J. O. de Wildem (Rijksherbarium, Leiden). Flora of Tropical East Africa. 1975

Morphology General Habit

Climbers or sometimes erect herbs, mostly provided with axillary tendrils (tribe Passifloreae), or erect shrubs or trees (tribe >i>Paropsieae), glabrous or hairy, rarely thorny

Morphology Leaves

Leaves mostly alternate, simple or compound, entire or lobed, often with glands on petiole and blade; stipules small, sometimes caducous

Morphology Reproductive morphology Inflorescences

Inflorescences axillary, either cymose (>i>Passifloreae), ending in 1–several tendrils or not, or racemose (>i>Paropsieae); bracts and bracteoles usually small

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or functionally unisexual (then plants often dioecious or polygamous); stipe articulate to pedicel; hypanthium saucer-shaped to tubiform

Morphology Reproductive morphology Flowers Calyx

Sepals 4–5(–6), imbricate, free or partially connate >i>(Adenia in part), often persistent

Morphology Reproductive morphology Flowers Corolla

Petals (3–)4–5(–6), imbricate, rarely absent

Morphology Reproductive morphology Flowers Corona

Corona extrastaminal, inserted on the hypanthium, various, composed of hairs, or of 1 or more whorls of thread-like processes or scales, or tubiform or cup-shaped, or absent (>i>Adenia in part)

Morphology Reproductive morphology Flowers Nectaries

Disk mostly extrastaminal, annular or composed of 5 mostly strap-shaped parts (>i>Adenia), or absent

Morphology Reproductive morphology Flowers Androecium

Stamens 4-many, inserted on the hypanthium or on an androgynophore, if few alternipetalous, free or partially connate; anthers 2-thecous, basifixed to dorsifixed, versatile or not, sometimes apiculate, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, sessile or on a gynophore or androgynophore, 1-locular, 3–5(–6)-carpellate, with 3–5(–6) parietal placentas; ovules mostly numerous, anatropous; styles 1 or 3–5, very short to distinct, free or partially united; stigma capitate to subglobose, sometimes much divided (>i>Adenia)

Morphology Reproductive morphology Fruits

Fruit a loculicidally 3–5-valved capsule, or berry-like

Morphology Reproductive morphology Seeds

Seeds mostly compressed, enveloped by a membranous or mostly pulpy aril; funicles often distinct; testa crustaceous, mostly pitted; endosperm horny; embryo large, straight, with foliaceous cotyledons

Passifloraceae, R. and A. Fernandes. Flora Zambesiaca 4. 1978

Morphology General Habit

Trees, shrubs or herbs, erect or more usually tendrillous climbers, sometimes with a napiform root

Morphology Leaves

Leaves alternate, rarely opposite, entire or variously lobed or 3–7-foliolate, sessile or petiolate, often with glands at the margin and on the abaxial surface of the lamina and at the apex of the petiole False stipules present in some species of Basananthe

Morphology Leaves Stipules

Stipules 0 or 2, caducous or persistent

Morphology Reproductive morphology Inflorescences

Tendrils, when present, axillary (sterile peduncles) or at the apex of the floriferous peduncle

Morphology Reproductive morphology Flowers

Flowers solitary, in racemes or in paniculate cymes, pedicellate, the pedicel often articulate and 3-bracteolate Flowers hermaphrodite or unisexual or polygamous, actinomorphic male flowers often with a vestigial ovary female flowers usually with staminodes

Morphology Reproductive morphology Flowers Hypanthium

Hypanthium ± developed, broad or narrow

Morphology Reproductive morphology Flowers Calyx

Calyx-tube short or ± elongate, coriaceous or herbaceous; lobes 3–? (3–6 in FZ species), valvate or imbricate, persisting with the tube

Morphology Reproductive morphology Flowers Corolla

Petals 0 or as many as the calyx-lobes, inserted at the base or mouth of the calyx-tube or in an intermediate position, the margin entire or fimbriate

Morphology Reproductive morphology Flowers Corona

Corona annular, single or double, rarely 0, tubular and often fimbriate at the margin with the laciniae erect or radiant, or reduced to a rim of hairs

Morphology Reproductive morphology Flowers Androecium

Stamens 5, rarely 6–10 or ?; filaments free or partially connate into a tube, inserted at the base of the flower or on an androgynophore; anthers oblong or linear, basifixed or dorsifixed, introrse, 2-locular, dehiscing by longitudinal slits, sometimes apiculate

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 3–5(6)-carpellar, 1-locular, often stipitate (borne on an androgynophore) or rarely subsessile; ovules many or few, on 3–5(6) parietal placentas, pendulous, anatropous, with the funicle elongate; styles 1 or 3-5, with capitate or clavate, sometimes fimbriate stigmas

sex Male

male flowers often with a vestigial ovary

sex Female

female flowers usually with staminodes

Morphology Reproductive morphology Flowers Nectaries

Disk-glands sometimes present

Morphology Reproductive morphology Fruits

Fruit baccaceous or a 3–5-valved capsule, usually many-seeded

Morphology Reproductive morphology Seeds

Seeds 1–?, on long funicles, ovate, compressed, rarely oblong or tumid, mostly arillate, with a furrowed and ridged seed-coat; endosperm fleshy, more or less abundant Embryo large, with leafy cotyledons and cylindrical radicle

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Herbs or shrubs, rarely trees, glabrous or pubescent, the hairs simple or branched; leaves alternate, simple, usually serrate, often 2-glandular at the base; stipules small or lacking

Morphology Reproductive morphology Flowers

Flowers perfect, regular, axillary, solitary or few, sessile or stalked, rarely in racemes; peduncles free or united with the petiole, often jointed and 2-bracteolate

Morphology Reproductive morphology Flowers Calyx

Calyx tubular, 5-lobed, soon deciduous

Morphology Reproductive morphology Flowers Corolla

Petals 5, often yellow, clawed and inserted in the throat of the calyx, sometimes bearing a fimbriate scale at the apex of the claw

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 5, inserted at the base, middle, or throat of the calyx tube, rarely hypogynous; filaments free; anthers oblong, opening inwardly

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, free, 1-celled; styles 3, terminal, thread-like, simple or divided; stigmas brush-like or rarely merely dilated; ovules numerous, 2-seriate on 3 parietal placentas

Morphology Reproductive morphology Fruits

Fruit a capsule splitting partially or completely by 3 valves; seeds oblong-cylindrical, slightly curved, arillate, with hard pitted coat; endosperm abundant; cotyledons plano-convex.

Distribution

A pantropical family of 8 genera and about 150 species, the majority American.

Turneraceae, R. Fernandes. Flora Zambesiaca 4. 1978

Morphology General Habit

Annual or perennial herbs, shrubs or rarely trees with simple or stellate hairs and sometimes setae, frequently glandular

Morphology Leaves

Leaves alternate, usually simple, subentire, serrate to more or less deeply divided, sometimes with sessile glands; petiole short or absent; stipules small or absent

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary, of racemes, panicles or cymes, or flowers solitary and axillary

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic, sometimes heterostylous

Morphology Reproductive morphology Flowers Calyx

Calyx usually with a short to long tube and 5 lobes, imbricate in aestivation

Morphology Reproductive morphology Flowers Corolla

Petals 5, usually brightly coloured, contorted in aestivation, adnate to the calyx-tube forming a short to fairly long hypanthium, sometimes with a ligule at the base

Morphology Reproductive morphology Flowers Androecium

Stamens with the filaments more or less adnate to the calyx-tube; anthers 2-celled, dorsifixed, introrse, dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-locular and usually several-ovulate, rarely 1-ovulate; ovules anatropous Styles 3, with fimbriate or laciniate stigmas

Morphology Reproductive morphology Fruits

Fruit a 3-valved capsule

Morphology Reproductive morphology Seeds

Seeds 1 to several with a unilateral aril; testa hard, reticulate-alveolate or longitudinally striate-sulcate

Turneraceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:1. 1954

Morphology General Habit

Herbs or undershrubs with entire or lobed alternate leaves; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic, yellowish

Morphology Reproductive morphology Flowers Calyx

Calyx tubular, 5-toothed, teeth imbricate

Morphology Reproductive morphology Flowers Corolla

Petals 5, inserted on the calyx-tube, free, clawed, contorted

Morphology Reproductive morphology Flowers Androecium

Stamens 5, from the base of the calyx-tube; filaments free; anthers 2-celled, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-celled with 3 parietal placentas; styles 3, terminal, slender; stigmas fringed; ovules numerous

Morphology Reproductive morphology Fruits

Fruit a capsule, opening loculicidally by 3 valves with the placenta in the middle of each

Morphology Reproductive morphology Seeds

Seeds arillate, pitted; endosperm horny or fleshy; embryo straight, large

Turneraceae, J. Lewis. Flora of Tropical East Africa. 1954

Morphology General Habit

Herbs, shrubs or trees, usually pubescent, sometimes stellately

Morphology Leaves

Leaves alternate, simple, often glandular; stipules small or absent

Morphology Reproductive morphology Flowers

Flowers solitary to numerous, axillary and sometimes terminal, in racemes, panicles or cymes, regular, hermaphrodite, sometimes heterostylous

Morphology Reproductive morphology Flowers Calyx

Calyx 5-merous, connate at least near the base, lobes imbricate in aestivation (2 wholly inside, 2 wholly outside)

Morphology Reproductive morphology Flowers Corolla

Petals (contorted in aestivation) and stamens 5, both adnate to the calyx-tube forming an hypanthium (see Fig. 1, p. 3) which may be very short (0.25 mm.)

Morphology Reproductive morphology Flowers Androecium

Filaments often flattened or narrowly winged; anthers introrse, dorsifixed, 2-celled, dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, unilocular; placentae usually parietal and pluriovulate, rarely basal and uniovulate (Stapfiella only); ovules anatropous Styles 3; stigmas apically divided (at least in our species), usually fimbriate or laciniate

Morphology Reproductive morphology Fruits

Fruit a 3-valved capsule

Morphology Reproductive morphology Seeds

Seeds arillate; aril dry (in our species a unilateral scarious membrane); testa hard, either with a raised rectangular network, the spaces of which form longitudinal lines of contiguous pits, or longitudinally striate-submuricate (>i>Stapfiella only)

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Herbs, subshrubs, shrubs, treelets or rarely trees, stems erect or decumbent, frequently with serial axillary buds; hairs generally present, simple, stellate or porrect- stellate; glandular hairs often present

Morphology Leaves Stipules

Stipules generally small or wanting, sometimes developed

Morphology Leaves

Leaves alternate simple, often bearing extrafloral nectaries at apex of petiole or base of blade margin, rarely abaxial; blades entire , margins entire, crenate, or toothed, not often pinnately cleft

Morphology Reproductive morphology Inflorescences

Inflorescences axillary or terminal, cymose or racemose, usually of solitary flowers, sometimes epiphyllous by adnation of peduncle and petiole; 2 bracteoles generally subtending flowers

Morphology Reproductive morphology Flowers

Flowers actinomorphic, bisexual, ephemeral, frequently showy, sometimes fragrant, distylous or homostylous; sepals 5, nearly distinct or united into a 10-veined tube, the lobes quincuncial; corolla commonly yellow, sometimes red, orange, pink, or white, frequently with purple spot or dark convergent lines at base of each petal blade , 5 petals distinct, alternate with sepals, with contorted aestivation, sometimes ligulate, clawed,  claw adnate to calyx thus forming floral tube; very rarely with corona, narrow, fringed, inserted at the throat, on bases of petal blades and sepal lobes; androecium with 5 stamens, these alternate with petals, filaments inserted at base of floral tube (then flowers hypogynous), or each filament with basal nectary and margins adnate to floral tube up to throat, forming 5 nectariferous pockets (then flowers perigynous), anthers dehiscing by longitudinal slits, commonly dorsifixed, sometimes nearly basifixed; pollen tricolporate, reticulate, binucleate; gynoecium syncarpous,  ovary generally superior, or slightly half-inferior, carpels 3,  locule 1, styles 3, distinct, erect or reflexed, glabrous or pilose , stigmas generally brush-like; placentation parietal , ovules 1-numerous per placenta, anatropous, bitegmic, crassinucellate

Morphology Reproductive morphology Fruits

Fruits loculicidal capsules, the surface smooth, verrucous or tuberculate, 3 valves with a median placenta

Morphology Reproductive morphology Seeds

Seeds 1-60, curved or straight, sometimes ridged, commonly reticulate, the areoles concave or with 1-2 punctiform pits; with a distinct exostome, chalaza rounded, sometimes prominent, navel-like; aril always present, fleshy, membranous when dry, glabrous to exceptionally hairy, generally inserted around hilum .

Note

Notes on delimitation: In Cronquist's system (1981) Turneraceae are located in the order Violales, with Achariaceae, Violaceae, Malesherbiaceae, Passifloraceae, and Lacistemataceae. According to the Angiosperm Phylogeny Group (APG II 2003), Turneraceae is placed in the order Malpighiales. Passiflora L., Turnera L. and Malesherbia Ruiz & Pav. form a clade that corresponds to Passifloraceaesensu lato, where an inclusion of Turneraceae and Malesherbiaceae into Passifloraceae was suggested. This clade shows 100% support in Chase et al. (2002); Davis and Chase (2004), as well as in the three-gene study of Soltis et al. (2000), but it is not clear whether there is paraphyly or not among these families. In a recent study of the Malpighiales, by means of plastid DNA sequences, a clade called Violids comprising Achariaceae, Violaceae, Malesherbiaceae, Turneraceae, Passifloraceae, and a Lacistemataceae-Salicaceae (including Flacourtiaceae) lineage was recovered as monophyletic with significant statistical support (Korotkova et al. 2009). In most of the previous studies, Turneraceae and Malesherbiaceae were sister to Passifloraceae, whereas data from Davis and Chase (2004) and Korotkova et al. (2009) provide evidence that Turneraceae and Passifloraceae are sister groups. Among the 40 or so families in the Malpighiales (APG, 2003), only four (Erythroxylaceae, Hypericaceae, Linaceae and Turneraceae) possess distylous species (Shore et al. 2006), and within the clade Violids, Turneraceae is the only one. Interestingly, the families of the Violid clade are the ones assigned by Cronquist (1981) to the order Violales, except for Salicaceaes. str., which was separated in the order Salicales. However, Cronquist drew attention to the fact that Salicales and Flacourtiaceae share many features (many stamens, parietal placentation, separate styles and the presence of salicin in Salix L., Populus L. and Idesia Maxim.) and pointed out that they could be placed nearby. It is noteworthy that all the molecular phylogenies have been made upon one species of each family, so most probably there will be changes when more species are taken into account. Number of genera: Adenoa Erblichia Piriqueta Turnera The chromosome number is known in some species of Piriqueta and Turnera.  In Piriqueta the basic number is x=7, while in Turnera there are 3 basic numbers: x=7, x=13 and x=5.

Distribution

Adenoa Arbo, monotypic, endemic to Cuba. Erblichia Seem., 1 species in Mesoamerica and 4 species in Madagascar. Piriqueta Aubl., 44 species in America and 1 in South Africa. Turnera, 140 species in America and 2 in Africa. All genera are native, some species are cultivated (Erblichiaodorata Seem. (flor de fuego, butterfly tree), Turnera subulata Sm. (buttercup, chanana, guarujá), Turnera ulmifolia L. (cat bush, ram-goat dash-along, sundrops, yellow alder). Some species are weedy, e.g. Turnera subulata in the North of Brazil and Turnera ulmifolia in the Caribbean Islands.  There is a very high rate of endemism in the family; Adenoa cubensis (Britton & Wilson) Arbo is endemic to Cuba, and there are many endemic and rare species within Turnera and Piriqueta.

Diagnostic

Distinguishing characters (always present): Alternatesimple leaves. Flowers actinomorphic, bisexual, ephemeral. Sepals 5, quincuncial aestivation. Petals 5, distinct, clawed. Stamens 5, anthers dehiscing by longitudinal slits. Carpels 3, ovary uniloculate, parietal placentation, anatropous ovules, styles 3. Fruit a capsule, loculicidal dehiscence. Seeds small (1-6mm long) and provided with an aril. Adenoa - shrub with stellate hairs, coriaceoustomentose leaves, without extrafloral nectaries. Flowers homostylous, solitary, peduncle developed, floral tube with nectariferous inner face, petals with colleters along the margin; corona absent; free staminal filaments, anthers dorsifixed; exerted stigmas. Capsuletomentose, dehiscent down to the middle, dressed at maturity with the remnant parts of the flower.  Seed obovoid, striate, chalaza rounded not prominent, aril narrow. Erblichia - tree with simple hairs, large leaves with small marginal nectaries. Flowers large, showy, fragrant, homostylous, solitary; peduncle and pedicel developed; sepals almost distinct; petals ligulate; stamens free, anthers dorsifixed; exerted stigmas. Capsule ellipsoidal, granulose. Seed obovoid, striate, chalaza rounded, not prominent; aril wide, oily, yellow or reddish, membranous when dry. Piriqueta - herbs, subshrubs or shrubs with porrect-stellate hairs, long glandular hairs with swollen base in most species. Leaves rarely with nectaries. Flowers usually distylous, sometimes homostylous, solitary, sometimes gathered in cymose inflorescences, hardly ever in an apical raceme; peduncle and pedicel developed, bracteoles generally wanting; floral tube with a membranouscrown at the throat; stamens adnate only at the base to the floral tube; anthers generally dorsifixed; styles sometimes bifurcate, stigmas generally brush-like. Capsule granulose, in a few species smooth. Seeds obovoid, reticulate, areoles slightly concave, in many species with a punctiform cavity, in a few species with 2 punctiform cavities; chalaza generally rounded, not prominent; aril narrow or wide. Turnera - herbs, subshrubs, shrubs or treelets with simple hairs, stellate in a few species, glandular hairs sessile-capitate, stipitate-capitate or microcapitate. Leaves often with nectaries. Flowers usually distylous, sometimes homostylous, solitary or gathered in cymose or racemose inflorescences; peduncle absent or developed, often adnate to the petiole, then flowers epiphyllous; pedicel generally not developed; bracteoles 2, persistent; floral tube formed by calyx and corolla, or by calyx, corolla and androecium; petals ligulate in a few species; anthers dorsifixed, sometimes almost basifixed; stigmas generally brush-like. Capsule granulose or smooth. Seeds obovoid, in a few species almost globose, chalaza rounded or prominent, sometimes concave; seed coat sometimes striate, usually reticulate, areoles sometimes with a punctiform cavity; aril narrow or wide, hardly ever pilose. Key to Neotropical genera of Turneraceae 1.  Trees 7-30m high, trunk up to 1m wide; showy homostylous flowers 6-8 cm long, with ligulate petals; sepals nearly distinct. Mesoamerica — Erblichia1.  Herbs, subshrubs, shrubs or treelets up to 4 m high; flowers less than 6 cm long; sepals united into 10-nerved tube — 2 2.  Petals with colleters all along the margin; homostylous flowers; styles divergent at the base — Adenoa (Cuba)2.  Petals without colleters along the margin; distylous flowers, homostylous in some species; styles converging at the base — 3 3.  Flowers with a membranouslaciniatecrown, inserted at the floral tube gorge on sepals and petals; peduncle always free; porrect-stellate hairs (central ray longer). Southern USA to Northern Argentina and Uruguay — Piriqueta 3.  Flowers without a crown; peduncle frequently adnate to the petiole (epiphyllous flowers), sometimes free; simple hairs, exceptionally stellate (rays of similar length). Southern USA to 39ºS in Argentina — Turnera Key to series of genus Turnera 1.  Staminal filaments adnate at the base to the floral tube, rarely free or coherent between them — 21.  Staminal filaments adnate along their margins to the petal claws up to the throat, forming nectariferous pockets between each filament and the corresponding sepal — 12 2.  Floral peduncle developed; solitary flowers, sometimes gathered in inflorescences — 32.  Floral peduncle not developed or very brief; flowers gathered in terminal or axillary inflorescences, exceptionally axillary and solitary — 10 3.  Floral peduncle totally free or inserted at the petiole base being the remainder free; fruit tuberculate or granulose — 43.  Floral peduncle partially or totally adnate to the petiole, at least the apical flowers epiphyllous — 8 4.  Pedicell developed, rarely very brief; solitary flowers, sometimes gathered in axillary cymose inflorescences — Series Salicifoliae4.  Pedicell absent; flowers solitary or gathered in terminal or axillary racemose inflorecences — 5 5.  Leaves with nectaries at the petiole or at the blade base, on the margin — 65.  Leaves without nectaries — 7 6.  Plants without glandular hairs; stipules inserted at both sides of the foliar base; anthers almost basifixed; seed obovoid, rapheal side often straight — Series Stenodictyae6.  Plants with capitate-sessile or microcapitate glandular hairs; stipules adnate to the foliar base and slightly to the petiole; anthers dorsifixed; seed subglobose, rapheal side convex — Series Annulares 7.  Plants with capitate-sessile or setiform glandular hairs; leaves small (0.4-4.5 cm long.) with stipules well developed — Series Microphyllae7.  Plants with capitate-stipitate glandular hairs, visible at least at the calyx; leaves small or medium (up to 10cm long.) with stipules reduced or replaced by colleters — Series Papilliferae 8.  Sepals coherent 1/3-1/2 of their length; staminal filaments free; fruit smooth or slightly depressed-reticulate — 98.  Sepals united 2/3 of their length; staminal filaments irregularly connate, to different heights; fruit granulose — Series Conciliatae 9.  Flowers generally small, gathered in terminal or lateralcapitate racemes; eaves generally medium (2-12 cm long); stipules developed, 0.5-4.5 mm long; fruit slightly depressed-reticulate; seedstriate-reticulate; chalaza prominent and concave — Series Capitatae9.  Flowers medium or small, solitary or gathered in terminal or axillary racemes; leaves small to medium (0.5-12 cm long), frequently with nectaries at apex of petiole or at blade base margin; stipules generally absent or rudimentary, developed in some species (up to 3 mm long); fruit smooth (warty in T. sidoides and T. melochioides var. rugosa); seedreticulate, sometimes the nodes elevated (tuberculate in T. sidoides); chalaza generally rounded, slightly prominent, rarely conical — Series Leiocarpae 10.  Leaves medium to large (4-34 cm long), petiolate, not revolute; stipules well developed, foliar nectaries present or absent — 1110.  Leaves very small (up to 16 mm long), sessile, entire, revolute; stipules absent, foliar nectaries absent — Series Sessilifoliae 11.  Fruit tuberculate or warty; seed puberulous — Series Salicifoliae11.  Fruit slightly depressed-reticulate or smooth; seed generally glabrous... Series Capitatae 12.  Floral peduncle free or absent; leaves with or without nectaries — Series Anomalae12.  Floral peduncle adnate to the petiole, sometimes the apical portion free; leaves with nectaries at the petiole-blade linkage, with a secretory device that looks like a pore — Series Turnera   Hairs simple and unicellular in Erblichia and most Turnera, stellate in Adenoa, porrect-stellate in Piriqueta. Glandular hairs long and micro-capitate with swollen bases in Piriqueta, stipitate-capitate in Turnera series Papilliferae, sessile-capitate in most Turnera series Annulares and Microphyllae, micro-capitate in many Turnera (series Anomalae, Capitatae, Leiocarpae and Turnera). Serial axillary buds frequent in Turnera and Piriqueta. Serial branches often developed in Turnera. Stipules developed in Erblichia, and some species of Turnera (series Annulares, Anomalae, Capitatae, Leiocarpae, Microphyllae and Salicifoliae). Foliar nectaries present in Erblichia, a few species of Piriqueta and many species of Turnera, those of series Turnera with a secretory device that looks like a pore. Cymose or racemose inflorescences in some species of Piriqueta and Turnera (series Salicifoliae, Capitatae, Leiocarpae and Anomalae). Prophylls (bracteoles) rudimentary or absent in species of Piriqueta with solitary flowers. Epiphyllous flowers in Turnera (series Conciliatae, Leiocarpae, Turnera and some species of series Anomalae and Capitatae). Distyly in a number of species of Turnera and Piriqueta. Sepals nearly distinct in Erblichia, joint into 10-veined tube with petal claws adhering in Adenoa, Piriqueta and Turnera. Stamens adnate to the floral tube only at the base in Adenoa, Piriqueta and Turnera (series Annulares, Capitatae, Conciliatae, Leiocarpae, Microphyllae, Papilliferae, Salicifoliae, Sessilifoliae and Stenodictyae). Stamen filaments united to petal claws along the margins up to the throat, forming nectariferous pockets, thus floral tube formed by the adnation of calyx, corolla and androecium, in Turnera (series Anomalae and Turnera). Anthers nearly basifixed in Erblichia and Turnera (series Stenodictyae and Turnera). Ovary slightly half-inferior in Turnera (series Turnera and Anomalae), Fruit usually naked, dressed with the remnants of the floral pieces in Adenoa and a few species of Turnera. Seed somewhat to considerably curved, straight in Turnera series Annulares and a few species of series Leiocarpae; chalaza usually rounded, prominent and navel-like in Turnera series Anomalae, Capitatae, Conciliatae, Salicifoliae, Stenodictyae and Turnera subseries Umbilicatae. Seed coat slightly striate in Adenoa and Erblichia; reticulate in Piriqueta, areoles concave, often with 1 or 2 punctiform pits; usually reticulate, areoles concave sometimes with 1 punctiform pit, occasionally ridged or tuberculate in Turnera . Leaves petiolate. Blade entire. Margin entire or crenate-serrate. Extrafloral nectaries at the petioleapex or at the blade base margin. Flowers distylous, epiphyllous, morning flowering. Prophylls (bracteoles) 2, persistent in the fruit. Sepals joint in a 10-nerved tube with adhering petal claws thus forming a floral tube. Stamens with nectaries at the base, on the outer side. Key differences from similar families: The predominant reproductive system in Turnera and Piriqueta is distyly. About 30% of the species are homostylous, and some species have both distylous and homostylous populations. Distylous species show reciprocal herkogamy, and are usually strongly self-incompatible, while homostylous species are self-compatible. Neither Malesherbiaceae nor Passifloraceae have distylous species. Many species of Turnera and Piriqueta look like Malvaceae or Sterculiaceae: the shape and colour of the flowers are similar, but in the latter families the stamens are not free and the filaments are connate into a tube. Benitez-Vieyra et al. (2007) show that there is floral mimicry between Turnera sidoides Vell. and species of Malvaceae in Argentina. In Malesherbiaceae and Passifloraceae: The ovary is raised. The corollaaestivation is cochlear. The corona is often well developed and distinctly coloured. In the flowers of the Malesherbiaceae: There is an androgynophore. The styles are inserted below the ovaryapex. The seeds are straight and lack an aril. The calyx is always persistent in the fruit. In Passifloraceae: The ovary is raised by a gynophore or androgynophore. The number of stamens is variable (4-5-many). Pollen grains may be 3-12-colporate or foraminate. Plants are often climbers with tendrils.

Arbo, M.M. (2009). Neotropical Turneraceae.

Morphology

Description

Herbs, subshrubs, shrubs, treelets or rarely trees, stems erect or decumbent, frequently with serial axillary buds; hairs generally present, simple , stellate or porrect- stellate ; glandular hairs often present. Stipules generally small or wanting, sometimes developed. Leaves alternate , simple , often bearing extrafloral nectaries at apex of petiole or base of blade margin, rarely abaxial ; blades entire , margins entire , crenate , or toothed, not often pinnately cleft . Inflorescences axillary or terminal , cymose or racemose, usually of solitary flowers, sometimes epiphyllous by adnation of peduncle and petiole ; 2 bracteoles generally subtending flowers. Flowers actinomorphic , bisexual , ephemeral, frequently showy, sometimes fragrant , distylous or homostylous; sepals 5, nearly distinct or united into a 10-veined tube, the lobes quincuncial; corolla commonly yellow, sometimes red, orange, pink, or white, frequently with purple spot or dark convergent lines at base of each petal blade , 5 petals distinct, alternate with sepals, with contorted aestivation , sometimes ligulate , clawed,  claw adnate to calyx thus forming floral tube; very rarely with corona , narrow, fringed, inserted at the throat, on bases of petal blades and sepal lobes; androecium with 5 stamens, these alternate with petals, filaments inserted at base of floral tube (then flowers hypogynous), or each filament with basal nectary and margins adnate to floral tube up to throat, forming 5 nectariferous pockets (then flowers perigynous ), anthers dehiscing by longitudinal slits, commonly dorsifixed, sometimes nearly basifixed; pollen tricolporate, reticulate , binucleate; gynoecium syncarpous,  ovary generally superior , or slightly half-inferior, carpels 3,  locule 1, styles 3, distinct, erect or reflexed , glabrous or pilose , stigmas generally brush-like; placentation parietal , ovules 1-numerous per placenta , anatropous, bitegmic, crassinucellate. Fruits loculicidal capsules, the surface smooth, verrucous or tuberculate , 3 valves with a median placenta . Seeds 1-60, curved or straight, sometimes ridged, commonly reticulate , the areoles concave or with 1-2 punctiform pits; with a distinct exostome, chalaza rounded , sometimes prominent , navel-like; aril always present, fleshy , membranous when dry, glabrous to exceptionally hairy, generally inserted around hilum .

General Description

Notes on delimitation

• In Cronquist's system (1981) Turneraceae are located in the order Violales, with Achariaceae, Violaceae, Malesherbiaceae, Passifloraceae, and Lacistemataceae.
• According to the Angiosperm Phylogeny Group (APG II 2003), Turneraceae is placed in the order Malpighiales. Passiflora L., Turnera L. and Malesherbia Ruiz & Pav. form a clade that corresponds to Passifloraceaesensu lato, where an inclusion of Turneraceae and Malesherbiaceae into Passifloraceae was suggested. This clade shows 100% support in Chase et al. (2002); Davis and Chase (2004), as well as in the three-gene study of Soltis et al. (2000), but it is not clear whether there is paraphyly or not among these families.
• In a recent study of the Malpighiales, by means of plastid DNA sequences, a clade called Violids comprising Achariaceae, Violaceae, Malesherbiaceae, Turneraceae, Passifloraceae, and a Lacistemataceae-Salicaceae (including Flacourtiaceae) lineage was recovered as monophyletic with significant statistical support (Korotkova et al. 2009).
• In most of the previous studies, Turneraceae and Malesherbiaceae were sister to Passifloraceae, whereas data from Davis and Chase (2004) and Korotkova et al. (2009) provide evidence that Turneraceae and Passifloraceae are sister groups.
• Among the 40 or so families in the Malpighiales (APG, 2003), only four (Erythroxylaceae, Hypericaceae, Linaceae and Turneraceae) possess distylous species (Shore et al. 2006), and within the clade Violids, Turneraceae is the only one.
• Interestingly, the families of the Violid clade are the ones assigned by Cronquist (1981) to the order Violales, except for Salicaceaes. str., which was separated in the order Salicales. However, Cronquist drew attention to the fact that Salicales and Flacourtiaceae share many features (many stamens, parietal placentation, separate styles and the presence of salicin in Salix L., Populus L. and Idesia Maxim.) and pointed out that they could be placed nearby.
• It is noteworthy that all the molecular phylogenies have been made upon one species of each family, so most probably there will be changes when more species are taken into account. 

Number of genera

• Adenoa
• Erblichia
• Piriqueta
• Turnera

General notes

• The chromosome number is known in some species of Piriqueta and Turnera.  In Piriqueta the basic number is x=7, while in Turnera there are 3 basic numbers: x=7, x=13 and x=5.

Status

• All genera are native, some species are cultivated (Erblichiaodorata Seem. (flor de fuego, butterfly tree), Turnera subulata Sm. (buttercup, chanana, guarujá), Turnera ulmifolia L. (cat bush, ram-goat dash-along, sundrops, yellow alder).
• Some species are weedy, e.g. Turnera subulata in the North of Brazil and Turnera ulmifolia in the Caribbean Islands. 
• There is a very high rate of endemism in the family; Adenoa cubensis (Britton & Wilson) Arbo is endemic to Cuba, and there are many endemic and rare species within Turnera and Piriqueta.

Distribution

Distribution in the Neotropics

• Adenoa Arbo, monotypic, endemic to Cuba.
• Erblichia Seem., 1 species in Mesoamerica and 4 species in Madagascar.
• Piriqueta Aubl., 44 species in America and 1 in South Africa.
• Turnera, 140 species in America and 2 in Africa.

Diagnostic

Useful tips for generic identification

Key to Neotropical genera of Turneraceae

1.  Trees 7-30m high, trunk up to 1m wide; showy homostylous flowers 6-8 cm long, with ligulate petals; sepals nearly distinct. Mesoamerica — Erblichia
1.  Herbs, subshrubs, shrubs or treelets up to 4 m high; flowers less than 6 cm long; sepals united into 10-nerved tube — 2

2.  Petals with colleters all along the margin; homostylous flowers; styles divergent at the base — Adenoa (Cuba)
2.  Petals without colleters along the margin; distylous flowers, homostylous in some species; styles converging at the base — 3

3.  Flowers with a membranouslaciniatecrown, inserted at the floral tube gorge on sepals and petals; peduncle always free; porrect-stellate hairs (central ray longer). Southern USA to Northern Argentina and Uruguay — Piriqueta
3.  Flowers without a crown; peduncle frequently adnate to the petiole (epiphyllous flowers), sometimes free; simple hairs, exceptionally stellate (rays of similar length). Southern USA to 39ºS in Argentina — Turnera

Key to series of genus Turnera

1.  Staminal filaments adnate at the base to the floral tube, rarely free or coherent between them — 2
1.  Staminal filaments adnate along their margins to the petal claws up to the throat, forming nectariferous pockets between each filament and the corresponding sepal — 12

2.  Floral peduncle developed; solitary flowers, sometimes gathered in inflorescences — 3
2.  Floral peduncle not developed or very brief; flowers gathered in terminal or axillary inflorescences, exceptionally axillary and solitary — 10

3.  Floral peduncle totally free or inserted at the petiole base being the remainder free; fruit tuberculate or granulose — 4
3.  Floral peduncle partially or totally adnate to the petiole, at least the apical flowers epiphyllous — 8

4.  Pedicell developed, rarely very brief; solitary flowers, sometimes gathered in axillary cymose inflorescences — Series Salicifoliae4.  Pedicell absent; flowers solitary or gathered in terminal or axillary racemose inflorecences — 5

5.  Leaves with nectaries at the petiole or at the blade base, on the margin — 6
5.  Leaves without nectaries — 7

6.  Plants without glandular hairs; stipules inserted at both sides of the foliar base; anthers almost basifixed; seed obovoid, rapheal side often straight — Series Stenodictyae
6.  Plants with capitate-sessile or microcapitate glandular hairs; stipules adnate to the foliar base and slightly to the petiole; anthers dorsifixed; seed subglobose, rapheal side convex — Series Annulares

7.  Plants with capitate-sessile or setiform glandular hairs; leaves small (0.4-4.5 cm long.) with stipules well developed — Series Microphyllae
7.  Plants with capitate-stipitate glandular hairs, visible at least at the calyx; leaves small or medium (up to 10cm long.) with stipules reduced or replaced by colleters — Series Papilliferae

8.  Sepals coherent 1/3-1/2 of their length; staminal filaments free; fruit smooth or slightly depressed-reticulate — 9
8.  Sepals united 2/3 of their length; staminal filaments irregularly connate, to different heights; fruit granulose — Series Conciliatae

9.  Flowers generally small, gathered in terminal or lateralcapitate racemes; eaves generally medium (2-12 cm long); stipules developed, 0.5-4.5 mm long; fruit slightly depressed-reticulate; seedstriate-reticulate; chalaza prominent and concave — Series Capitatae
9.  Flowers medium or small, solitary or gathered in terminal or axillary racemes; leaves small to medium (0.5-12 cm long), frequently with nectaries at apex of petiole or at blade base margin; stipules generally absent or rudimentary, developed in some species (up to 3 mm long); fruit smooth (warty in T. sidoides and T. melochioides var. rugosa); seedreticulate, sometimes the nodes elevated (tuberculate in T. sidoides); chalaza generally rounded, slightly prominent, rarely conical — Series Leiocarpae

10.  Leaves medium to large (4-34 cm long), petiolate, not revolute; stipules well developed, foliar nectaries present or absent — 11
10.  Leaves very small (up to 16 mm long), sessile, entire, revolute; stipules absent, foliar nectaries absent — Series Sessilifoliae

11.  Fruit tuberculate or warty; seed puberulous — Series Salicifoliae
11.  Fruit slightly depressed-reticulate or smooth; seed generally glabrous... Series Capitatae

12.  Floral peduncle free or absent; leaves with or without nectaries — Series Anomalae
12.  Floral peduncle adnate to the petiole, sometimes the apical portion free; leaves with nectaries at the petiole-blade linkage, with a secretory device that looks like a pore — Series Turnera

 

• Hairs simple and unicellular in Erblichia and most Turnera , stellate in Adenoa , porrect-stellate in Piriqueta .
• Glandular hairs long and micro-capitate with swollen bases in Piriqueta , stipitate-capitate in Turnera series Papilliferae, sessile-capitate in most Turnera series Annulares and Microphyllae, micro-capitate in many Turnera (series Anomalae, Capitatae, Leiocarpae and Turnera).
• Serial axillary buds frequent in Turnera and Piriqueta .
• Serial branches often developed in Turnera .
• Stipules developed in Erblichia , and some species of Turnera (series Annulares, Anomalae, Capitatae, Leiocarpae, Microphyllae and Salicifoliae).
• Foliar nectaries present in Erblichia , a few species of Piriqueta and many species of Turnera , those of series Turnera with a secretory device that looks like a pore.
• Cymose or racemose inflorescences in some species of Piriqueta and Turnera (series Salicifoliae, Capitatae, Leiocarpae and Anomalae).
• Prophylls (bracteoles) rudimentary or absent in species of Piriqueta with solitary flowers.
• Epiphyllous flowers in Turnera (series Conciliatae, Leiocarpae, Turnera and some species of series Anomalae and Capitatae).
• Distyly in a number of species of Turnera and Piriqueta .
• Sepals nearly distinct in Erblichia , joint into 10-veined tube with petal claws adhering in Adenoa , Piriqueta and Turnera .
• Stamens adnate to the floral tube only at the base in Adenoa , Piriqueta and Turnera (series Annulares, Capitatae, Conciliatae, Leiocarpae, Microphyllae, Papilliferae, Salicifoliae, Sessilifoliae and Stenodictyae).
• Stamen filaments united to petal claws along the margins up to the throat, forming nectariferous pockets, thus floral tube formed by the adnation of calyx, corolla and androecium, in Turnera (series Anomalae and Turnera).
• Anthers nearly basifixed in Erblichia and Turnera (series Stenodictyae and Turnera).
• Ovary slightly half-inferior in Turnera (series Turnera and Anomalae),
• Fruit usually naked, dressed with the remnants of the floral pieces in Adenoa and a few species of Turnera .
• Seed somewhat to considerably curved, straight in Turnera series Annulares and a few species of series Leiocarpae; chalaza usually rounded, prominent and navel-like in Turnera series Anomalae, Capitatae, Conciliatae, Salicifoliae, Stenodictyae and Turnera subseries Umbilicatae.
• Seed coat slightly striate in Adenoa and Erblichia ; reticulate in Piriqueta , areoles concave, often with 1 or 2 punctiform pits; usually reticulate, areoles concave sometimes with 1 punctiform pit, occasionally ridged or tuberculate in Turnera .

Other important characters

• Leaves petiolate.
• Bladeentire.
• Margin entire or crenate-serrate.
• Extrafloral nectaries at the petioleapex or at the blade base margin.
• Flowers distylous, epiphyllous, morning flowering.
• Prophylls (bracteoles) 2, persistent in the fruit.
• Sepals joint in a 10-nerved tube with adhering petal claws thus forming a floral tube.
• Stamens with nectaries at the base, on the outer side.

Distinguishing characters (always present)

• Alternatesimple leaves.
• Flowers actinomorphic, bisexual, ephemeral.
• Sepals 5, quincuncial aestivation.
• Petals 5, distinct, clawed.
• Stamens 5, anthers dehiscing by longitudinal slits.
• Carpels 3, ovary uniloculate, parietal placentation, anatropous ovules, styles 3.
• Fruit a capsule, loculicidal dehiscence.
• Seeds small (1-6mm long) and provided with an aril.

Key differences from similar families

• The predominant reproductive system in Turnera and Piriqueta is distyly. About 30% of the species are homostylous, and some species have both distylous and homostylous populations. Distylous species show reciprocal herkogamy, and are usually strongly self-incompatible, while homostylous species are self-compatible. Neither Malesherbiaceae nor Passifloraceae have distylous species.
• Many species of Turnera and Piriqueta look like Malvaceae or Sterculiaceae: the shape and colour of the flowers are similar, but in the latter families the stamens are not free and the filaments are connate into a tube. Benitez-Vieyra et al. (2007) show that there is floral mimicry between Turnera sidoides Vell. and species of Malvaceae in Argentina.

In Malesherbiaceae and Passifloraceae:

• The ovary is raised.
• The corollaaestivation is cochlear.
• The corona is often well developed and distinctly coloured.

In the flowers of the Malesherbiaceae:

• There is an androgynophore.
• The styles are inserted below the ovaryapex.
• The seeds are straight and lack an aril.
• The calyx is always persistent in the fruit.

In Passifloraceae:

• The ovary is raised by a gynophore or androgynophore.
• The number of stamens is variable (4-5-many).
• Pollen grains may be 3-12-colporate or foraminate.
• Plants are often climbers with tendrils.

Notable genera and distinguishing features

• Adenoa - shrub with stellate hairs, coriaceoustomentose leaves, without extrafloral nectaries. Flowers homostylous, solitary, peduncle developed, floral tube with nectariferous inner face, petals with colleters along the margin; corona absent; free staminal filaments, anthers dorsifixed; exerted stigmas. Capsuletomentose, dehiscent down to the middle, dressed at maturity with the remnant parts of the flower.  Seed obovoid, striate, chalaza rounded not prominent, aril narrow.
• Erblichia - tree with simple hairs, large leaves with small marginal nectaries. Flowers large, showy, fragrant, homostylous, solitary; peduncle and pedicel developed; sepals almost distinct; petals ligulate; stamens free, anthers dorsifixed; exerted stigmas. Capsule ellipsoidal, granulose. Seed obovoid, striate, chalaza rounded, not prominent; aril wide, oily, yellow or reddish, membranous when dry.
• Piriqueta - herbs, subshrubs or shrubs with porrect-stellate hairs, long glandular hairs with swollen base in most species. Leaves rarely with nectaries. Flowers usually distylous, sometimes homostylous, solitary, sometimes gathered in cymose inflorescences, hardly ever in an apical raceme; peduncle and pedicel developed, bracteoles generally wanting; floral tube with a membranouscrown at the throat; stamens adnate only at the base to the floral tube; anthers generally dorsifixed; styles sometimes bifurcate, stigmas generally brush-like. Capsule granulose, in a few species smooth. Seeds obovoid, reticulate, areoles slightly concave, in many species with a punctiform cavity, in a few species with 2 punctiform cavities; chalaza generally rounded, not prominent; aril narrow or wide.
• Turnera - herbs, subshrubs, shrubs or treelets with simple hairs, stellate in a few species, glandular hairs sessile-capitate, stipitate-capitate or microcapitate. Leaves often with nectaries. Flowers usually distylous, sometimes homostylous, solitary or gathered in cymose or racemose inflorescences; peduncle absent or developed, often adnate to the petiole, then flowers epiphyllous; pedicel generally not developed; bracteoles 2, persistent; floral tube formed by calyx and corolla, or by calyx, corolla and androecium; petals ligulate in a few species; anthers dorsifixed, sometimes almost basifixed; stigmas generally brush-like. Capsule granulose or smooth. Seeds obovoid, in a few species almost globose, chalaza rounded or prominent, sometimes concave; seed coat sometimes striate, usually reticulate, areoles sometimes with a punctiform cavity; aril narrow or wide, hardly ever pilose.

Literature

Important literature

APG. 2003. An update of the angiosperm phylogeny group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 339-436

Arbo M.M. 1977. Adenoa, nuevo genero americano de Turneraceae. Hickenia 1(16): 87-91

Arbo M.M. 1979. Revisión del género Erblichia (Turneraceae). Adansonia, ser. 2, 18(4): 459-482.

Arbo M.M. 1995. Turneraceae Parte I. Piriqueta. Flora Neotropica (Monograph 67, 156 pages, 59 figs.

Arbo, M.M. 1997. Estudios sistemáticos en Turnera (Turneraceae). I. Series Salicifoliae y Stenodictyae. Bonplandia (Argentina) 9(3-4): 151-208Arbo M.M. 2000. Estudios sistemáticos en Turnera (Turneraceae). II. Series Annulares, Capitatae, Microphyllae y Papilliferae. Bonplandia (Argentina) 10: 1-82.Arbo M.M. 2004. Turneraceae in Smith N. et al. (eds.) "Flowering Plants of the Neotropics". Princeton University Press.

Arbo M.M. 2005. Estudios sistemáticos en Turnera (Turneraceae). III. Series Anomalae y Turnera. Bonplandia (Argentina) 14(3-4): 115-318.

Arbo M.M. 2007. Turneraceae in Kubitzki K., The families and genera of vascular plants, Springer. Alemania. Vol. IX: 458-466.

Arbo M.M. 2008. Estudios sistemáticos en Turnera (Turneraceae). IV. Series Leiocarpae, Sessilifoliae y Conciliatae. Bonplandia (Argentina) 17(3-4): 107-334.

Benitez-Vieyra S., Hempel de Ibarra N., Wertlen A.M. & Cocucci A.A.  2007. How to look like a mallow: evidence of floral mimicry between and Malvaceae. Proc. R. Soc. B. 274: 2239-2248.

Chase M.W., Zmartzy S., Lledó M.D., Wurdack K.J., Swensen S.M. & Fay M.F. 2002. When in doubt, put it in Flacourtiaceae: a molecular analysis based on plastid rbcL sequences. Kew Bull.57: 141-181.

Cronquist A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York.

Davis C.C. & Chase M.W. 2004. Elatinaceae are sister to Malpighiaceae; Peridiscaceae belong to Saxifragales. Amer. J. Bot. 91: 262-273.

Korotkova N., Schneider J.V., Quandt D., Worberg A., Zizka G. & Borsch T. 2009. Phylogeny of the eudicot order Malpighiales: analysis of a recalcitrant clade with sequences of the petD group II intron. Pl. Syst. Evol. 271, on line.

Shore J.S., Arbo M.M. & Fernández A. Breeding system variation, genetics and evolution in the Turneraceae. New Phytologist 171: 539-551.

Soltis D.E., Soltis P.S., Chase M.W., Mort M.E., Albach D.C., Zanis M., Savolainen V., Hahn W.H., Hoot S.B., Fay M.F., Axtell M., Swensen S.M., Prince L.M., Kress W.J., Nixon K.C., Farris J.S. 2000. Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Bot. J. Linn. Soc. 133: 381-461.

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 [and more or less continuously updated since]. http://www.mobot.org/MOBOT/research/APweb/.