Caprifoliaceae Juss.

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

Shrubs, vines, lianas, rarely trees

Morphology Leaves

Leaves simple, opposite, sometimes ternate, with vestigial interpetiolary stipules, entire, margins often crenate . Inflorescence generally axillary, cymose, bracteate

Morphology Reproductive morphology Flowers

Flowers showy, hermaphrodite, slightly to strongly zygomorphic; sepals (4-)5, united in short tube; corolla (4-)5- lobed, joined in a zygomorphic tube, white, pink or red; stamens 3-5, fused to the corolla -tube, anthers 2-locular; ovary inferior, (2-)3 carpellate, style long, exserted, stigma capitate, placentation axile, ovules 1-many per locule .  Fruit fleshy, generally berries (nutlet in Vesalea), 1-many seeded.

Note

Number of genera: Vesalea R. Br. (formerly Abelia) - Temperate Asia, reaching Mexico, 2 - 5 species. Lonicera L.  - honeysuckle, from Asia, N. Africa and North America south to Costa Rica, 5 species. Symphoricarpos  Duhamel - snow berry, C. China; North America, reaching Guatemala, two species. In former classifications, the Caprifoliaceae used to include species with actinomorphic flowers, such as Viburnum and Sambucus, but those are nowadays placed within the Adoxaceae after molecular studies (APG II, 2003). The old circumscription is still followed by Delprete in Smith (2004). In 'APGII' (Angiosperm Phylogeny Group, 2003) an expanded Dipsacales was promoted in the 'Euasterids II', including a broader concept of the Caprifoliaceae, which encompassed the Dipsacaceae and Valerianaceae known from the Neotropics, along with the Adoxaceae. If sunk into the Caprifoliaceae the Dipsacaceae and Valerianaceae would form very discrete subfamilies. However, in Neotropikey the three families (Caprifoliaceae, Dipsacaceae and Valerianaceae) are treated separately, q.v.

Diagnostic

Notable genera and distinguishing features: Lonicera japonica is often cultivated in the Neotropics. Key to genera of Neotropical Caprifoliaceae 1. Climbing plants with twinning stems ... Lonicera 1. Shrubs ...  2. Berries white, inflated ... Symphoricarpus 2. Nutlets dry, with persistent calyx... Vesalea Key differences from similar families: The opposite leaves, inferior ovary and tubular corolla may generate confusion with the Rubiaceae, but the Caprifoliaceae flowers are mostly zygomorphic and their stipules are vestigial. Distinguishing characters (always present): Climbers or shrubs. Leaves opposite, margins rarely entire. Flowers with bilateral symmetry. Ovary inferior with 3 carpels. Stipules vestigial-Fruitsucculent berries (sometimes nutlets).

Distribution

Many species of Caprifoliaceae are cultivated in the Neotropics, especially Lonicera and Vesalea. However, this is not a very prominent family in the region, with only a few species ocurring only marginally in the area. In the Neotropics, the Caprifoliaceae is a marginal family with a few genera ocurring at high altitude and reaching Mexico from the West of the USA, and none are native in South America.

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition

Characters of similar families: Viburnaceae: actinomorphic flowers, lobed stigma on sessile or short style, lacking a nectary in the corolla. Rubiaceae: interpetiolar stipules, leaf margins always entire. Hydrangeaceae: petals free, stamens twice as many as petals. Cornaceae: flowers typically 4-merous. Compositae: (confused with Dipsacoideae) anthers united into cylinder; fruit an achene. Apiaceae: (confused with Dipsacoideae) leaves alternate, fruit a schizocarp of 2 mericarps.

Morphology General Habit

Herbs, shrubs, and woody climbers

Morphology Leaves Stipules

Stipules absent, sometimes interpetiolar stipule-like structures apparent

Morphology Leaves

Leaves simple or pinnately compound, opposite, decussate, sometimes whorled, connate or in a basal rosette; margins entire, toothed or pinnatifid

Morphology Reproductive morphology Inflorescences

Inflorescences usually axillary, thyrsoid cymes 1-, 2- or 3-flowered, bracteate, or a compact terminal capitulum (Dipsacoideae and Scabiosoideae)

Morphology Reproductive morphology Flowers

Flowers bisexual, usually zygomorphic, sometimes appearing nearer actinomorphic: campanulate to funneliform, often restricted into a narrow tube; calyx persistent, of 2–5 fused sepals, sometimes accrescent, sometimes reduced to short or long teeth only, or setae, involucel or epicalyx of connate bracteoles sometimes present; corolla sympetalous, lobes 4–5, imbricate, bilabiate unless regular, sometimes gibbous with basal nectaries, dense glandular hairs in throat, or with short abaxial spur; stamens 4–5, alternating with petals, equal in length or didynamous, filaments attached to corolla tube, anthers free, 2-locular, often prominently exserted; ovary inferior, 2–8 fused carpels, some aborting, with 1–5 fertile locules and 1–several ovules per locule, style elongate, exserted, stigma capitate

Morphology Reproductive morphology Fruits

Fruit dry dehiscent or indehiscent capsule, berry, pitted drupe, achene or cypsela Capsules and berries many-seeded, drupes with 2–4 pyrenes, achenes and cypselae single-seeded.

Distribution

About 28 genera and ca. 825 species, predominantly temperate, mostly in the Northern hemisphere. An expanded family concept including Dipsacaceae and Valerianaceae is largely accepted. Shrubby genera Sambucus and Viburnum, which were often included in past classifications of Caprifoliaceae, are now strongly supported as Viburnaceae.

Note

Leaves simple or pinnately compound, opposite. Flowers 5-merous, synsepalous and sympetalous, usually zygomorphic, tubular or 2-lipped, ovary inferior. Fruit a capsule, berry, drupe or achene.

Description Author

Lesley Walsingham

Valerianaceae, J. O. Kokwaro (East African Herbarium). Flora of Tropical East Africa. 1968

Morphology General Habit

Annual (occasionally biennial) or perennial herbs, rarely subshrubs; often with strongly scented rhizomes

Morphology Leaves

Leaves opposite or sometimes forming basal rosettes, exstipulate, often pinnately much divided but sometimes entire; cauline leaves sometimes few, small or none; basal leaves pinnatifid or once- or twice-pinnatisect; base often sheathing

Morphology Reproductive morphology Inflorescences

Inflorescence a many-flowered compound dichasial cyme, thyrse or monochasium, sometimes condensed and capitate, bracteate and usually bracteolate (the subtended bracteoles often mistaken for the calyx)

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual by abortion (plants then dioecious, as in some Valeriana spp.), irregular or almost regular, usually 5-merous

Morphology Reproductive morphology Flowers Calyx

Calyx often small or obsolete at the time of flowering, sometimes enlarging as the fruit matures, and then variously lobed; lobes often forming a pappus

Morphology Reproductive morphology Flowers Corolla

Corolla funnel-shaped or tubular, often attenuated at the base, generally basally spurred or saccate, wider at the throat; limb 3–5-fid, oblique or 2-lipped; lobes imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens 1–4, epipetalous, alternating with the corolla-lobes; anthers versatile, 2- or 4-lobed, 2- or 4-thecous, introrse, dehiscing longitudinally; pollen grains tricolpate, echinate

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior, tricarpellate, 3-locular but only 1 locule fertile; ovule solitary and pendulous, anatropous; style single and slender; stigma subtruncate, entire or shortly 2–3-lobed

Morphology Reproductive morphology Fruits

Fruit a 1-seeded achene

Morphology Reproductive morphology Seeds

Seed pendulous; embryo straight, large; cotyledons oblong; radicle superior; endosperm very thin or absent

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

Terrestrial annual, biennial, or perennial herbs sometimes subshrubs or cushion-forming shrubs or shrublets, or climbers or scandent plants, usually with characteristic odour, especially when dry (associated with valeric acid); monoecious, dioecious or gynodioecious

Morphology Roots

Rootstock fibrous, sometimes rhizomatous, or possessing conical, napiform or fusiform taproots

Morphology Stem

Stems often hollow, often glabrous, or variously pubescent, sometimes glaucous

Morphology Leaves

Leaves opposite or alternate, whorled or basal, sometimes decussate, exstipulate simple or pinnatisect, variously shaped, base usually clasping, sessile or petiolate, margins entire or serrate

Morphology Reproductive morphology Inflorescences

Inflorescences cymose, usually compound, often densely so, with many small flowers, bracteate

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual, sessile or short-pedicellate, usually zygomorphic (especially with funnel-shaped corollas, but symmetrical with rotate corollas) and bilaterally symmetrical-Calyx persistent and often becoming woody in fruit, 0-5- lobed, each lobe of few to many segments, very small in flower and markedly inrolled and imbricate, often enlarging to a pappus on apex of fruit, pappus frequently plumose, sometimes absent

Morphology Reproductive morphology Flowers Corolla

Corollas white, pink red or lilac, or very rarely yellow, funnel-shaped, subsalverform or rotate, corolla tube usually longer than corolla lobes, often bulging on one side, saccate or spurred below; corolla lobes 5; stamens 1-4, usually inserted near base of corolla tube and alternating with lobes, usually exserted from corolla, sometimes included; anthers sessile, on short filaments, or filaments distinct, versatile, 2- or 4- lobed, dehiscing introrsely; stigma 2-fid, 3-fid, or subentire, exserted

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary inferior, 3-celled, 1 cell with pendent ovule and other 2 sterile and often much reduced in fruit

Morphology Reproductive morphology Fruits

Fruit dry and indehiscent, glabrous or variously setuliferous or hairy; seed endospermic with straight embryo.

Diagnostic

Distinguishing characters (always present): The family is a very natural one characterized by: Thyrsoid inflorescence. Sympetalous asymmetric flowers. Inferior 3-carpellate ovaries, one fertile carpel. A single style. Fruit an achene. In addition, dried members of the tribe Valerianeae have a characteristic odour of valeric acid, although this is absent in the tribe Patrinieae. Modifications of the calyx which, if present, form an often distinctive plumose pappus. Stamen number. Corolla lobe number. Key differences from similar families: Most recent analyses of the Dipsacales place the Valerianaceae as sister to the Dipsacaceae itself. The easiest character to perhaps separate these two families is the presence of: A characteristic capitulum in the Dipsacaceae surrounded by an involucre of more than 2 bracts, and a fruit surrounded by an epicalyx. In the Valerianaceae a capitulum is never present, and the epicalyx is absent.

Note

Number of genera: Eriksen (1989a) reduced all of the South American representatives of the Valerianaceae to one amorphous Valeriana, s.l. She maintained this exceptional lumping in her treatment for the Flora of Ecuador (Eriksen 1989b). Weberling (2001) commented on this and, with the supporting evidence of Backlund & Donoghue (1996), maintained the status of the smaller South American genera together with a more restricted sense of Valeriana. As now accepted, there are between 10-13 genera, and some 350-450 species, worldwide. Mainly distributed in the northern hemisphere, South America, and southeastern South Africa. One or two weedy species found in the Neotropics. A total of eight genera are recorded for Central and South America: Aretiastrum (DC.) Spach Astrephia Dufr. Belonanthus Graebn. Phyllactis Pers. Plectritis DC. Stangea Graebn Valeriana L. (the largest genus in the family), widespread in both the Old and New World. Notes on delimitation: The only apparent delimitation problem appears to be with the inclusion of Triplostegia Wall. ex DC. in the family; it is often placed in its own family within the Dipsacales. It does not pose any problems with delimitation of the Valerianaceae in the New World. In 'APGII' (Angiosperm Phylogeny Group, 2003) an expanded Dipsacales was promoted in the 'Euasterids II', including a broader concept of the Caprifoliaceae, which encompassed the Dipsacaceae and Valerianaceae known from the Neotropics, along with the Adoxaceae. If sunk into the Caprifoliaceae the Dipsacaceae and Valerianaceae would form very discrete subfamilies. However, in Neotropikey the three families (Caprifoliaceae, Dipsacaceae and Valerianaceae) are treated separately, q.v.

Distribution

Centranthus ruber (L.) DC., an Old World species, is reportedly a weed in some areas of South America, as is Valeriana officinalis L. Valeriana L. is widespread throughout the Neotropics from Mexico south to Argentina. Most of the other genera in the family are restricted to the high altitude areas of the Andes throughout the region.

Valerianaceae, J. F. M. Cannon. Flora Zambesiaca 7:1. 1983

Morphology General Habit

Perennial or, less commonly, annual herbs, often with aromatic rhizomes; rarely slightly shrubby

Morphology Leaves

Leaves opposite or occasionally basal; exstipulate, frequently pinnately divided, sometimes entire; bases often sheathing

Morphology Reproductive morphology Inflorescences

Inflorescence a many–flowered compound dichasial cyme, thyrse or monochasium, sometimes sub–capitate; bracts and bracteoles usually present

Morphology Reproductive morphology Flowers

Flowers hermaphrodite (occasionally unisexual and the plants then dioecious); normally 5–merous and irregular to nearly regular

Morphology Reproductive morphology Flowers Calyx

Calyx obsolete or minute, sometimes expanded in fruit and lobed to form a pappus

Morphology Reproductive morphology Flowers Corolla

Corolla tubular or funnel–shaped, often with a basal spur or sacoate; limb 3–5 fid, oblique or divided into 2 distinct lips

Morphology Reproductive morphology Flowers Androecium

Stamens 1–4 alternating with the lobes of the corolla, epipetalous; anthers versatile, 2–4 lobed

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior, with 3 locules only 1 of which is fertile; ovule solitary, anatropous; style, stigma subtruncate, entire or shortly 2–3 lobed

Morphology Reproductive morphology Fruits

Fruit an achene

Morphology Reproductive morphology Seeds

Seed pendulous; embryo straight with oblong cotyledons; endosperm sparse or lacking

Ghazanfar, S. A., Edmondson, J. R. (Eds). (2013). Flora of Iraq, Volume 5, Part 2: Lythraceae to Campanulaceae. Kew Publishing

Morphology General Habit

Annual or perennial herbs, rarely suffruticose or shrubby, sometimes with scented rhizomes or tubers, dichasially branched

Morphology Leaves

Leaves exstipulate, opposite or radical, pinnatifid or entire

Morphology Reproductive morphology Flowers

Flowers in dense cymose inflorescences, hermaphrodite (in ours) or unisexual, zygomorphic or sometimes nearly regular

Morphology Reproductive morphology Flowers Calyx

Calyx annular or toothed (Valerianella), often forming in fruit a feathery pappus (Valeriana, Kentranthus)

Morphology Reproductive morphology Flowers Corolla

Corolla funnel-shaped, mostly with 5 imbricate lobes, sometimes with a long spur

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 1–3, rarely 4, inserted in the corolla tube

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary 3-celled, two sterile cells of a different size and shape (smaller or bigger) than the fertile one

Morphology Reproductive morphology Fruits

Fruit an achene or nut

Morphology Reproductive morphology Seeds

Seeds without endosperm.

Distribution

About 16 genera and 400 species, generally distributed except for S Africa, Australia and Polynesia. Three genera in Iraq. Valeriana and Kentranthus are very rare and found in northern Iraq in the mountains; Valerianella is more common.

Note

Under the APG III (2009) classification, Caprifoliaceae is expanded to include Valerianaceae, together with families Diervillaceae, Dipsacaceae, Linnaeaceae and Morinaceae.

Ghazanfar, S. A., Edmondson, J. R. (Eds). (2013). Flora of Iraq, Volume 5, Part 2: Lythraceae to Campanulaceae. Kew Publishing

Morphology General Habit

Annual or perennial herbs, rarely shrubs

Morphology Leaves

Leaves simple or compound, opposite or rarely whorled, exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence usually a head, or rarely a spike of false whorls (Morina) with a calyx-like involucre of bracts

Morphology Reproductive morphology Flowers

Flowers bisexual, zygomorphic, each with a tubular calyx-like involucel and borne in the axils of imbricate bracteoles

Morphology Reproductive morphology Flowers Calyx

Calyx small, cup-like or tubular, or divided into pappus-like segments

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, 4- or 5-lobed, imbricate

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens usually 4, free, alternate with the corolla lobes and inserted towards the top of the tube

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary inferior, unilocular, with a solitary ovule pendulous from the apex of the locule

Morphology Reproductive morphology Fruits

Fruit indehiscent, dry, enclosed within the calyx-like involucel and often crowned with a persistent calyx.

Distribution

The family consists of nine genera and about 160 species. It occurs primarily in the Mediterranean area and the Near East.

Note

Under the APG III (2009) classification, Caprifoliaceae is expanded to include Dipsacaceae, together with families Diervillaceae, Linnaeaceae, Morinaceae and Valerianaceae.

Dipsacaceae, D. M. Napper. Flora of Tropical East Africa. 1968

Morphology General Habit

Annual or perennial herbs or subshrubs

Morphology Leaves

Leaves opposite, entire, lobed or variously dissected, often markedly heterophyllous

Morphology Reproductive morphology Inflorescences

Inflorescences of one or several long-pedunculate dense heads each surrounded by an involucre

Morphology Reproductive morphology Flowers Receptacle

Receptacle convex, globose or cylindric, naked, shortly pilose or with paleaceous bracts each subtending a single flower

Morphology Reproductive morphology Flowers

Flowers sessile, bisexual (excepting sometimes the marginal ones), with the ovary surrounded by and enclosed in a highly characteristic tubular involucel formed by the fusion of 2 or 4 bracts often with a membranous cupular or spreading lobed limb

Morphology Reproductive morphology Flowers Calyx

Calyx cupular or saucer-shaped with an entire, irregularly 4–5-lobed or toothed margin, sometimes with long naked or plumose bristles

Morphology Reproductive morphology Flowers Corolla

Corolla gamo-petalous with an irregularly 4- or 5-lobed limb and spreading lobes

Morphology Reproductive morphology Flowers Androecium

Stamens 2–4, alternating with the corolla-lobes and inserted near the top of the tube; anthers dithecous, oblong, exserted

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior and unilocular with a solitary ovule; style filiform with a terminal or lateral fusiform oblique or bifid stigma

Morphology Reproductive morphology Fruits

Fruit an achene closely surrounded by the persistent involucel and crowned by the persistent calyx

Dipsacaceae, Margaret J. and J. F. M. Cannon. Flora Zambesiaca 7:1. 1983

Morphology General Habit

Usually perennial herbs or subshrubs, less commonly annual or biennials

Morphology Leaves

Leaves opposite, estipulate, entire to pinnatifid, to finely divided, sometimes noticeably heterophyllous, rarely connate at the base

Morphology Reproductive morphology Inflorescences

Inflorescence usually capitate on a long peduncle, rarely verticillate or in lax panicles Heads surrounded by well–developed involucres

Morphology Reproductive morphology Flowers Receptacle

Receptacle sometimes with short hairs, otherwise glabrous or with a paleaceous bract subtending each flower

Morphology Reproductive morphology Flowers

Flowers sessile, hermaphrodite (rarely the outer female only); the ovary surrounded by an involucel of fused bracts, which are frequently extended to form a cupule

Morphology Reproductive morphology Flowers Calyx

Calyx cupular, often with a toothed margin and long–spreading bristles

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, somewhat zygomorphic or otherwise irregular, and with a 4–5 lobed limb

Morphology Reproductive morphology Flowers Androecium

Anthers frequently on rather long filaments and exserted at anthesis Stamens 2–4, inserted towards the top of the tube and alternating with the corolla lobes

Morphology Reproductive morphology Flowers Gynoecium

Style filiform, simple, clavate or shortly 2–lobed Ovary inferior with a solitary ovule

Morphology Reproductive morphology Fruits

Fruit an achene within the persistent involucel and frequently with a conspicuous persistent calyx

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

Annuals, biennial or perennial herbs, rarely shrubs

Morphology Roots

Roots fibrous or sometimes tuberous

Morphology Stem

Stems glabrous or variously pubescent, sometimes with prickles, often with basal rosettiform leaves, especially when young, or leaves cauline

Morphology Leaves

Leaves opposite or verticillate, exstipulate, sessile, pseudopetiolate or petiolate, bases sometimes connate, sometimes heterophyllous with lower leaves usually simply but upper leaves variously lobed, lamina entire or deeply lobed, laciniate, lyrate - pinnatifid, or 1-2- pinnatisect, margins entire or variously toothed, glabrous or variously pubescent, sometimes the prickles on lower surface (especially on venation)

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary, of dense cymose capitula, subtended by phyllaries 1-3-seriate, phyllaries free or rarely connate in basal half, flowering occurring at several points in capitulum, or rarely from apparent middle and working outwards (i.e. Dipsacus L.)

Morphology Reproductive morphology Inflorescences Capitulum

Capitula appearing radiate/radiant, ovoid or subglobose, hemispherical or cylindrica

Morphology Reproductive morphology Flowers Florets

Florets hermaphrodite or female, each usually with a basal epicalyx (involucel) of connate bracteoles sometimes expanded into a variously shaped, often scarious, corona and often subtended by a receptacular scale, involucel variously angled or ridged, receptacular scales linear - lanceolate, sometimes spine -tipped, glabrous or variously pubescent; calyx small, cupuliform or divided into 4-5 teeth or of numerous teeth or setae; corollas usually zygomorphic, lobes 4-5, subequal, or corolla 2-lipped, blue, lilac, white, yellow, pink, red, purple; stamens free, 2 or 4, epipetalous, alternating with corolla lobes, and usually conspicuously exserted from corolla, anthers dorsifixed, dehiscing introrsely via longitudinal slits; stigma solitary or 2- lobed; ovary inferior, unilocular, ovule pendent

Morphology Reproductive morphology Fruits

Fruit dry, indehiscent, enclosed by epicalyx and usually surrounded by persistent calyx, fruit body variously angled (4-), very rarely with an elaiosome at base (Knautia L.); seed 1, endospermic, embryo straight.

Note

Notes on delimitation: The family appears to be reasonably naturally delimited, although some floras expand the family concept with the inclusion of the Morinaceae (Morina L.) with its spikes of false verticellasters, and Triplostegiaceae (Triplostegia Wall. ex DC.), with its characteristic double epicalyx. In the Neotropics the three taxa commonly encountered: Dipsacus fullonum L., D. sativus (L.) Honckeny and Scabiosa atropurpurea L., are all easily placed in the family sensu stricto. In 'APGII' (Angiosperm Phylogeny Group, 2003) an expanded Dipsacales was promoted in the 'Euasterids II', including a broader concept of the Caprifoliaceae, which encompassed the Dipsacaceae and Valerianaceae known from the Neotropics, along with the Adoxaceae. If sunk into the Caprifoliaceae the Dipsacaceae and Valerianaceae would form very discrete subfamilies. However, in Neotropikey the three families (Caprifoliaceae, Dipsacaceae and Valerianaceae) are treated separately, q.v. Number of genera: The greater part of the family (7 genera and c. 250 spp.) are found in the Old World in Asia, Eurasia and northern Africa. Only tow are found in the Neotropics. Only two are found in the Neotropics.

Distribution

Only two genera, and three species, have been commonly recorded within the Neotropics: Dipsacusfullonum and D. sativus (both probably cultivated but occasionally naturalized). Scabiosa atropurpurea (often cultivated, and escaped). Both representatives of the family found in the Neotropics have been/are cultivated in a number of countries, and have naturalized.

Diagnostic

The presence of radiate/radiant capitula in most genera is a useful character. In Dipsacus the odd flowering pattern (acropetal/basipetal) is characteristic, along with the distinctly prickly stem and underside of the leaves, together with the connate leaf bases. Distinguishing characters (always present): Capitulum, surrounded by a few-seriate involucre. Florets with zygomorphic corollas. Basal epicalyx or involucel. Persistentcalyx. Freeexserted stamens. Cypsela. Key differences from similar families: Similar families include the Calyceraceae and Compositae in the presence of numerous florets arranged in capitula sometimes with receptacular bracts present. The Compositae is easily recognized because of the presence of: Included or exserted stamens with connate anthers (vs. stamens exserted with free anthers). Achenes with or without a pappus of scales or hairs (vs. the presence of the involucel and persistent coroniform calyx in the Dipsacaceae). The Dipsacaceae also have either 2 or 4 stamens; the number of stamens in Compositae is rarely reduced to 4. The Calyceraceae differ from the Dipsacaceae as follows: Stamens in the Calyceraceae are alternating with the corolla lobes whereas they are epipetalous in the Dipsacaceae. Although both the Dipsacaceae and Calyceraceae have persistent calyces those in the Calyceraceae typically become lignified and spiny; those in the Dipsacaceae become cupuliform or divided into a number of setae. An involucel is only found in the Dipsacaceae. Anthers in the Calyceraceae dehisce antrorsely whereas those in the Dipsacaceae dehisce introrsely. Dipsacus L. is easily recognized by: The prickly stem. Underside of the leaves with conspicuous prickles. Characteristic capitulum. Scabiosa L. often possesses: Basal rosette of leaves. The leaves are heteromorphous with entire or toothed lower leaves and pinnatifid cauline leaves. Typically radiate/radiant capitula.

Dipsacaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Perennial or annual herbs

Morphology Leaves

Leaves opposite or verticillate, entire or pinnately divided; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, zygomorphic, often crowded into heads with an involucre of leafy bracts

Morphology Reproductive morphology Flowers Calyx

Calyx epigynous, cupular or divided into pappus-like segments

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, lobes imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens usually 4, rarely 2-3, alternate with the corolla-lobes and inserted usually towards the bottom of the tube; anthers 2-celled, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior, 1-celled; style slender; ovule solitary, pendulous from the top

Morphology Reproductive morphology Seeds

Seeds with large straight embryo in scanty endosperm