Primulaceae Batsch ex Borkh.

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: Campanulaceae: latex often present, flowers actino- or zygomorphic, ovary usually inferior, with annular disk. Saxifragaceae: stamens 5 or 10, anthers 2-locular; intrastaminal disk present, ovary inferior to superior.

Morphology General Habit

Perennial or annual herbs, suffrutescent (woody-based), rarely tuberous, sometimes tufted or cushion-forming, shrubs, rarely small trees, rarely aquatic- Glandular hairs often present

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves simple, often forming a basal rosette, alternate, opposite or whorled; margins entire to serrate or rarely finely pinnate (Hottonia); sometimes glandular punctate

Morphology Reproductive morphology Inflorescences

Inflorescences often scapose, terminal or axillary, solitary, racemose, spicate or umbellate, bracts usually present

Morphology Reproductive morphology Flowers

Flowers bisexual, actinomorphic, rarely zygomorphic, usually 5-merous; sepals basally connate, persistent in fruit; petals basally connate, corolla tube short to very long, campanulate to salverform, lobes usually overlapping, entire, notched, fringed or reflexed (Cyclamen and some Primula); stamens the same number as and opposite the petals, spreading or connivent forming a cone, rarely with staminodes, often heterostylous, anthers opening by slits; ovary superior, rarely semi-inferior (Samolus), 1-locular, placentation free central; several to many ovules; style 1, stigma not lobed, capitate

Morphology Reproductive morphology Fruits

Fruit a dehiscing capsule, sometimes a drupe, with persistent calyx

Morphology Reproductive morphology Seeds

Seeds few to numerous, angular.

Distribution

55 genera with ca. 3,000 species in four subfamilies. Most temperate species are in the Primuloideae subfamily. Temperate species of Myrsinoideae tend to be herbs or shrubs, rarely small trees. Mainly occurring in northern temperate regions.

Ecology

Preferring alpine habitats of meadows or rocky crevices, or wet, boggy habitat and woodlands. Some genera are widely cultivated and important horticulturally.

Note

Herbs, shrubs. Leaves often forming basal rosettes. Hairs often present. Inflorescences often scapose. Flowers bisexual, actinomorphic, 5-merous; stamens equal to and opposite the corolla lobes; ovary usually superior. Fruit a capsule.

Description Author

Clare Drinkell

Primulaceae, P. Taylor. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Annual or perennial herbs or rarely shrubs; stems erect or procumbent and rooting at the nodes

Morphology Leaves

Leaves mostly basal, rarely cauline, alternate, opposite or verticillate, simple or lobate, often dentate; stipules absent

Morphology Reproductive morphology Flowers

Flowers solitary to paniculate, actinomorphic, hermaphrodite

Morphology Reproductive morphology Flowers Calyx

Calyx persistent, often leafy

Morphology Reproductive morphology Flowers Corolla

Corolla hypocrateriform, campanulate or tubular, lobes 5, imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens inserted on the corolla, the same number as and opposite to the lobes

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, rarely semi-inferior, 1-celled with a free basal placenta and numerous, or very rarely few ovules

Morphology Reproductive morphology Fruits

Fruit a capsule, many-seeded or very rarely 1-seeded

Morphology Reproductive morphology Seeds

Seeds angular with a small straight embryo in copious endosperm

Primulaceae, F. K. Kupicha. Flora Zambesiaca 7:1. 1983

Morphology General Habit

Annual or perennial herbs or rarely shrubs; stems erect or prostrate and rooting at nodes

Morphology Leaves

Leaves exstipulate, basal or cauline, alternate, opposite or verticillate, simple or lobed, entire or dentate

Morphology Reproductive morphology Flowers

Flowers actinomorphic or very rarely zygomorphic, hermaphrodite, often heterostylous, solitary or in racemose, spicate, paniculate, umbellate or verticillate inflorescences

Morphology Reproductive morphology Flowers Calyx

Calyx gamosepalous, free or rarely partially adnate to the ovary, (4)5(9)–partite, usually persistent

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, rotate to campa–nulate with a very short to long tube and 4–9–lobed limb, rarely absent

Morphology Reproductive morphology Flowers Androecium

Stamens equalling corolla–lobes, usually adnate to the corolla, oppositipetalous, sometimes alternating with staminodes

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior or rarely semi–inferior, unilocular; placentation free–central; ovules 2–3 or more, usually many

Morphology Reproductive morphology Fruits

Fruit a capsule with valvate or circumscissile dehiscence, rarely indehiscent

Morphology Reproductive morphology Seeds

Seeds few to numerous, often angular

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

Note

Dark dots throughout especially in the leaves and flowers. Leaves alternate; calyx persistent (usually with dark dots). Stamens as many and opposite the corolla lobes. Fruit drupaceous with a fleshy/spongy mesocarp.

Recognition

Characters of similar families: Rubiaceae: stipules present, leaves opposite, inferior ovary. Theaceae: flowers usually solitary, stamens numerous, lacking dark dots in floral parts. Solanaceae: lacking dark dots, ovary with many locules, many seeds per locule. Actinidiaceae: young parts scaly or thick hairy, stamens numerous. Sapotaceae: white sap present, stipules often present, lacking dark dots throughout.

Morphology General Habit

Shrubs, trees, or climbers, sometimes subherbaceous; branches flattened-triangular at the base (‘gusseted’)

Morphology General Hair

Hairs present, simple to complex glandular hairs

Morphology Leaves Stipules

Stipules lacking

Morphology Leaves

Leaves alternate, sometimes subverticillate; simple, with dark dot-like or linear glands; lamina margins entire, or crenate, or rarely serrate

Morphology Reproductive morphology Inflorescences

Inflorescences various, racemes, panicles, cymose, corymbose, terminal or axillary, often at apices of lateral branches

Morphology Reproductive morphology Flowers

Flowers uni- or bisexual, actinomorphic, 4- or 5-merous; sepals basally connate, usually glandular and persistent in fruit; petals basally connate (rarely free), lobes usually overlapping (imbricate, quincuncial or contorted), white to pink; stamens the same number as and opposite the petals, anthers 2-celled opening by slits or sometimes by pores; ovary superior, 1-locular, placentation free central with few to many ovules; style 1, terminal-Fruit drupaceous with a crustaceous endocarp and a thin fleshy to spongy mesocarp, usually globose, often with conspicuous dots or linear glands in the exocarp (‘skin’) and persistent calyx

Morphology Reproductive morphology Seeds

Seed usually 1.

Distribution

A pantropical family with c. 40 (perhaps up to 55) genera and c. 2,000 species; this group has been traditionally treated in its own family Myrsinaceae. Ardisia (pantropical), Myrsine (including Rapanea, pantropical), Embelia (Old World tropics), and Cybianthus (New World tropics) are the largest genera.

Description Author

Timothy Utteridge

Timothy M. A. Utteridge and Laura V. S. Jennings (2022). Trees of New Guinea. Kew Publishing. Royal Botanic Gardens, Kew

Distribution

Members of the family are found throughout lowland habitats including mangroves, but with the greatest diversity in montane habitats in New Guinea. The family is represented in New Guinea by 11 genera (including climbing/herbaceous taxa) and c. 150 species, including the endemic genus Fittingia and many endemic species.

Morphology General Habit

Trees (often unbranched), shrubs, climbers (rarely herbs)

Morphology General Indumentum

Indumentum of simple or multi-celled hairs or scales. Stipules absent

Morphology Leaves

Leaves simple, alternate, distichous, rarely opposite or whorled, venation pinnate, often glandular with linear or circular glands. Inflorescences terminal, axillary, or at apices of lateral branches, racemose or compound racemose, spicate corymbose, cymose, or umbellate or fasciculate; pedicels subtended by a floral bract or with a pair of bracteoles below the calyx (Maesa)

Morphology Reproductive morphology Flowers

Flowers bisexual or unisexual and then plants polygamodioecious, androdioecious or dioecious, 4- or 5- (or 6-)merous, actinomorphic; sepals basally connate or free, persistent, usually glandular; petals basally connate or rarely free, usually glandular; stamens as many as and opposite to petals, usually adnate to the corolla; anthers dehiscing by longitudinal slits or apical pores or rarely transversely septate (Aegiceras); ovary superior or semi-inferior (Maesa), 1-celled, ovules 1 to many; style one, stigma simple or lobed

Morphology Reproductive morphology Fruits

Fruits drupaceous with a fleshy or spongy exocarp

Morphology Reproductive morphology Seeds

Seeds 1 to many.

Primulaceae, P. Taylor. Flora of Tropical East Africa. 1958

Morphology General Habit

Annual or perennial herbs, or rarely shrubs; stems erect or prostrate and rooting at the nodes

Morphology Leaves

Leaves basal or cauline, alternate, opposite or verticillate, simple or lobed, entire or dentate; stipules absent

Morphology Reproductive morphology Flowers

Flowers axillary and solitary, racemose, verticillate or paniculate, hermaphrodite, regular, sometimes heterostylous

Morphology Reproductive morphology Flowers Calyx

Calyx gamosepalous, free or rarely adnate to the ovary, 4–9-partite, usually persistent

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, rotate, hypocrateriform or campanulate, with a very short to long tube and 4–9-partite limb, or corolla rarely absent

Morphology Reproductive morphology Flowers Androecium

Stamens equal in number to and inserted opposite the corolla-lobes

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior or rarely half inferior, 1-locular; ovules 2–3 or usually many on a free central placenta

Morphology Reproductive morphology Fruits

Fruit a capsule, valvate or circumscissile or rarely indehiscent

Morphology Reproductive morphology Seeds

Seeds small, 1, few or usually numerous, often angular

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 or small trees, usually evergreen, hermaphrodite or in Clavija Ruiz & Pav Usually dioecious or subdioecious "Myco-heterotrophic (""saprophytic""), small herbs without chlorophyll; root system tuberous or rarely cylindrical, roots filiform, glabrous; stems unbranched"

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or lateral, basically racemose but through reductions sometimes appearing as few-flowered umbels, rarely single-flowered, each flower subtended by a small bract

Morphology Reproductive morphology Flowers

Flowers regular or slightly asymmetric because of unequal size of the corolla lobes, 5- or 4- merous, bisexual or in Clavija often unisexual, the aestivation imbricate Flowers terminal, solitary, 5-20 mm long, bisexual, actinomorphic to sometimes zygomorphic, the cylindric to urceolate floral tube with a circular throat, surrounded by an ornamented annulus; tepals 6, forming 2 whorls each of different shape and size, sometimes connate into a miter; stamens 6, pendent, often alternating with 6 interstaminal lobes, filaments short, anthers dithecal, extrorse, longitudinally dehiscent, connective dilated; perianth basallly circumscissile; style 1, cylindrical, much shorter than the floral tube, 3-branched, each branch bearing a stigma, or unbranched with 1 capitate stigma; ovary inferior, 1-locular with 3 parietal placentas or with 3 placental columns in the center, septal nectaries absent

Morphology Reproductive morphology Flowers Calyx

Calyx persistent, lobes free to base, glandular punctuate, margins membranaceous

Morphology Reproductive morphology Flowers Corolla

Corolla sympetalous, usually firm and waxy in texture, lobes usually somewhat unequal in size

Morphology Reproductive morphology Flowers Androecium Staminodes

Staminodal structures present, fused to the corolla tube, alternating with the lobes

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens homomerous, antepetalous; filaments flattened, ± fused at base and to the lower part of the corolla tube, free to base or in Clavija often united into a tube; anthers basifixed, extrorsely dehiscent with longitudinal slits, the upper and lower parts of thecae filled with calcium oxalate crystals

Morphology Reproductive morphology Flowers Gynoecium

Gynoecium superior; ovary ovoid to subglobose, undivided, unilocular; style short to somewhat longer than ovary; stigma entire or vaguely lobed, truncate or capitate; ovules few to numerous, spirally inserted on a basal column

Morphology Reproductive morphology Fruits

Fruit a berry with a dry and sometimes woody pericarp, indehiscent, subglobose, oblong, or ovoid, yellow or orange Fruit cup-shaped, fleshy, crowned by a rim formed by the persistent basal part of the floral tube, dehiscent by withering of the wall

Morphology Reproductive morphology Seeds

Seeds dark brown to brownish-yellow, partly or entirely embedded in placental tissue, endosperm abundant; embryo straight or slightly curved, cotyledons linear or foliaceous . "Seeds many, small and ""dustlike"", (narrowly) ellipsoid to ovoid ."

Diagnostic

Distinguishing characters (always present): Schizogenous secretory cavities lacking. Free central placenta. Stamens epipetalous; anthers extrorsely dehiscent partly filled with accumulations of calcium oxalate crystals. Corolla tube with a ring of staminodal structures, alternating with the petals. Fleshy fruits with a dry exocarp and few to many large seeds. Leaves with subepidermal layers or bundles of extraxylary sclerenchyma. Key differences from similar families: Lacking secretory cavities (present in Myrsinaceae, Primulaceae, Samolaceae). Ring of petaloid staminodes (alsoin Samolaceae). Fleshy fruits (differing from Primulaceae, Samolaceae) with more than one seed (present Myrsinaceae). Key to genera of the Neotropical Thismiaceae 1. Filaments with distinct, fimbriate appendages ... Tiputinia .1. Filaments without fimbriate appendages ... Thismia . Notable genera and distinguishing features: Thismiaceae are very much undercollected in the Neotropics and of most species (except for the quite common Thismia panamensis (Standl.) Jonker) only 1 collection has been known. Two species from the Alto Macahe (in the state of Rio de Janeiro, Brazil) were only collected once each in 1889. Clavija, Neomezia, Theophrasta: monocaulous shrubs and treelets. Bonellia and Jacquinia: often spine -tipped leaves. Deherainia: large green flowers with foetidscent. Flowers sometimes zygomophic; one of the whorls of tepals often provided with long and filiform appendages, sometimes with a swollen top (osmophores?). Key differences from similar families: Formerly placed in Burmanniaceae from which it differs by having: Circumscissile perianth. 6 pendent (instead of 3 erect) stamens. Short style far exceeded by the floral tube. (For differences with Triuridaceae see under Burmanniaceae).

Note

Number of genera: Bonellia Bertero ex Colla Clavija  Ruiz & Pav. Deherainia Decne. Jacquinia L. Neomezia Votsch Theophrasta L. Votschia Ståhl Notes on delimitation: Not including Samolaceae, a sister group occasionally merged with the Theophrastaceae. Notes on delimitation: Formerly placed in Burmanniaceae (see Merckx, 2008). Number of genera: 2 genera: Thismia (12 spp.) and Tiputinia (1 sp.).

Morphology Leaves

Leaves (2-)4, alternate, sessile simple, entire, scale-like, sometimes forming a rosette just below the flower Leaves alternate, often pseudo- verticillate, exstipulate, petiolate simple, glandular - punctate, mostly with bundles or layers of subepidermal, extraxylary sclerenchyma

Distribution

Native. NW Mexico and S Florida to S Brazil and Paraguay. Endemic. Thismiaceae: from Costa Rica in the North to Bolivia and SE Brazil in the South; absent in the West Indian Islands. Tiputinia P.E.Berry & C.L.Woodw. - Ecuador. Thismia Griff. - from Costa Rica in the North to Bolivia and SE Brazil in the South; absent in the West Indian Islands.

Ståhl, B. (2009). Neotropical Theophrastaceae.

Morphology

Description

Shrubs or small trees, usually evergreen , hermaphrodite or in Clavija Ruiz & Pav. usually dioecious or subdioecious. Leaves alternate , often pseudo- verticillate , exstipulate , petiolate , simple , glandular - punctate , mostly with bundles or layers of subepidermal, extraxylary sclerenchyma. Inflorescences terminal or lateral , basically racemose but through reductions sometimes appearing as few-flowered umbels, rarely single-flowered, each flower subtended by a small bract . Flowers regular or slightly asymmetric because of unequal size of the corolla lobes, 5- or 4- merous , bisexual or in Clavija often unisexual, the aestivation imbricate . Calyx persistent , lobes free to base, glandular punctuate, margins membranaceous. Corolla sympetalous, usually firm and waxy in texture, lobes usually somewhat unequal in size. Staminodal structures present, fused to the corolla tube, alternating with the lobes. Stamens homomerous, antepetalous; filaments flattened, ± fused at base and to the lower part of the corolla tube, free to base or in Clavija often united into a tube; anthers basifixed, extrorsely dehiscent with longitudinal slits, the upper and lower parts of thecae filled with calcium oxalate crystals. Gynoecium superior ; ovary ovoid to subglobose, undivided, unilocular; style short to somewhat longer than ovary ; stigma entire or vaguely lobed , truncate or capitate ; ovules few to numerous, spirally inserted on a basal column. Fruit a berry with a dry and sometimes woody pericarp , indehiscent , subglobose, oblong , or ovoid , yellow or orange. Seeds dark brown to brownish-yellow, partly or entirely embedded in placental tissue, endosperm abundant; embryo straight or slightly curved, cotyledons linear or foliaceous .

General Description

Notes on delimitation

• Not including Samolaceae, a sister group occasionally merged with the Theophrastaceae.

Number of genera

• Bonellia Bertero ex Colla
• Clavija  Ruiz & Pav.
• Deherainia Decne.
• Jacquinia L.
• Neomezia Votsch
• Theophrasta L.
• Votschia Ståhl

Status

• Endemic.

Distribution

Distribution in the Neotropics

• NW Mexico and S Florida to S Brazil and Paraguay.

Diagnostic

Other important characters

• Leaves with subepidermal layers or bundles of extraxylary sclerenchyma.

Distinguishing characters (always present)

• Schizogenous secretory cavities lacking.
• Free central placenta.
• Stamens epipetalous; anthers extrorsely dehiscent partly filled with accumulations of calcium oxalate crystals.
• Corolla tube with a ring of staminodal structures, alternating with the petals.
• Fleshy fruits with a dry exocarp and few to many large seeds.

Key differences from similar families

• Lacking secretory cavities (present in Myrsinaceae, Primulaceae, Samolaceae).
• Ring of petaloid staminodes (alsoin Samolaceae).
• Fleshy fruits (differing from Primulaceae, Samolaceae) with more than one seed (present Myrsinaceae).

Notable genera and distinguishing features

• Clavija, Neomezia, Theophrasta: monocaulous shrubs and treelets.
• Bonellia and Jacquinia: often spine -tipped leaves.
• Deherainia: large green flowers with foetidscent.

Literature

Important literature

Ståhl, B. Theophrastaceae. Flora Neotropica Monographs (in press).

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

Morphology General Habit

Trees or shrubs, glabrous or pubescent; leaves alternate, simple, entire or with crenulate or serrate margins, the surfaces often gland-dotted; stipules none A pantropical family of about 30 or 40 rather weakly differentiated genera and 900 species, few if any of economic value.

Morphology Reproductive morphology Flowers

Flowers perfect, or unisexual and dioecious, regular, with parts in 4s or 5s in terminal or axillary clusters, racemes, or panicles

Morphology Reproductive morphology Flowers Calyx

Calyx segments free or somewhat connate, often ciliate and gland- dotted; corolla with segments united at least toward the base.

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens as many as the corolla-lobes, shorter than these and opposite them, almost free or inserted on and somewhat connate with the corolla-tube; anthers opening by slits or apical pores; staminodes none

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, sessile, 1-celled, with a free central placenta; ovules numerous or few; style short or long; stigma simple and variously capitate, lobed, or fimbriate

Morphology Reproductive morphology Fruits

Fruit a 1-seeded drupe; seed with copious endosperm.

Myrsinaceae, F. K. Kupicha. Flora Zambesiaca 7:1. 1983

Morphology General Habit

Trees, shrubs and lianes

Morphology Leaves

Leaves exstipulate, alternate, simple, entire or toothed, often clustered at branch ends, always with schizogenous resiniferous dots or lines but these varying from very obvious to obscure

Morphology Reproductive morphology Inflorescences

Inflorescences lateral or terminal (not in FZ area), racemose, paniculate, umbellate or fasciculate

Morphology Reproductive morphology Flowers

Flowers usually small and individually inconspicuous, 5– or 4–merous, actinomorphic, hermaphrodite or plants very often dioecious

Morphology Reproductive morphology Flowers Calyx

Calyx of free or connate sepals, often ciliate, often with dark spots, aestivation valvate, imbricate or contorted

Morphology Reproductive morphology Flowers Corolla

Petals usually white or pink, less frequently purple or yellow, free or more usually connate, often with dark dots or stripes, often papillose

Morphology Reproductive morphology Flowers Androecium

Stamens as many as and opposite petals; anthers introrse, dehiscing by longitudinal slits or rarely by apical pores (not in FZ area); filaments long or short, adnate to corolla, sometimes almost free

Morphology Reproductive morphology Flowers Gynoecium

Ovary globose, ovoid or clavate, superior or (in Maesa) semi–inferior, unilocular; number of carpels obscure, probably 3–4; style long and slender or short and thick or rarely absent; stigma punctiform, capitate, discoid or lobed; placenta free–central, bearing few to many ovules in one or more rows

Morphology Reproductive morphology Fruits

Fruit an indehiscent berry or drupe, 1·seeded except for Maesa which is many–seeded

Morphology Reproductive morphology Seeds

Seeds with copious, often ruminate, endosperm

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

Trees, treelets to shrubs or subshrubs, herbs or epiphytes; leaves, flowers and fruit with secretory cavities appearing as dark dots or dashes

Morphology Leaves

Leaves simple, alternate, subopposite, opposite or pseudowhorled; blades often coriaceous, margins entire to variously serrate, stipules absent; petioles usually short, occasionally absent

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary; racemes, panicles to simple or compound corymbs, fascicles or verticils, or solitary flowers

Morphology Reproductive morphology Flowers

Flowers bisexual or unisexual, with staminodes (on pistillate flowers) and pistillodes (on staminate flowers); partially connate sepals and petals, rarely petals free, epipetalous, distinct or connate stamens, and connate carpels (3) 4-5 (6); anthers dehiscing by longitudinal slits to pores; ovary superior, with free -central placentation; ovules few to several

Morphology Reproductive morphology Fruits

Fruit a single-seeded drupe or a valved or opercular capsule with few to many seeds.

Diagnostic

Ardisia: shrubs and small trees.  The inflorescence is paniculate, the lateral branches with the usually 5-parted flowers arranged in glomerules or cymes, typically more or less umbellate to much reduced ramiflorous inflorescence. The stamens are always included, the anthers rather long, and the filaments usually short.  Commonest in middle-elevation cloud forests and more diverse in southern Central American than in South America. Cybianthus: from small subshrubs to subcanopy trees, with racemose inflorescence, sometimes slightly branched at the apex, corolla rotate or tabulate. Rapanea: Small trees typical of middle-elevation forests especially in rather exposed situations.  Distinctive in the sessile axillary and ramiflorus flowers, typically densely clustered along the twigs below the leaves on suppressed short-shoots. Parathesis: shrubs and small trees.  Frequently the branchlets and lower leaf surface near midrid have stellate pubescence.  The flowers, always 4-merous, have unusually large yellow anthers and are always in panicles with valvateperianth and densely pubescentcalyx and petals. The fruit is distinctive in being ribbed and is also characterized by the pubescentpersistentcalyx lobes. Habit (epiphytes, herbs, trees or shrubs). Disposition of leaves. Type of inflorescences. Petals: free or partially connate. Fusion and insertion of filaments. Anthers exserted or included. Type and texture of the corolla. Imbrication of the calyx (regular or irregular). Imbrication of the corolla (valved or imbricated). Key to genera of neotropical Myrsinaceae. 1. Epiphytes; leaves very narrow, sessile, leaf apices mucronate — Grammadenia 1. Not epiphytes; leaves not like above — 2 2. Herbs; flowers solitary ... 32. Trees to shrubs; flowers in inflorescences ... 4 3. Leaves opposite; seeds not winged — Anagallis 3. Leaves alternate; seeds winged — Lysimachia 4. Flower in fascicles or verticils — Rapanea 4. Flowers not in fascicles or verticils — 5 5. Petals free — 65. Petals partially connate... 7 6. Filaments united (Mexico) — Heberdenia 6. Filaments free (Cuba) — Solonia 7. Exserted anthers (filaments longer than the corolla) — 87. Included anthers (filaments shorter than the corolla) — 9 8. Tubular coriaceous corolla, remains of styles not persistent on fruit (Caribbean) — Wallenia 8. Not tubular and not coriaceous corolla, remains of long styles persistent on fruit (cloud and elfin forests) — Gentlea 9. Plants with long glandular trichomes, campanulate corolla — Synardisia 9. Plants without glandular trichomes, not campanulate corolla — 10 10. Umbellate clusters of flowers arranged in panicles — Ctenardisia 10. Others kinds of inflorescences — 11 11. Calyx closed in bud, irregularly rupturing into 2-7 lobes (also visible in fruit) — Geissanthus 11. Calyx regularly divided — 12 12. Valvate perianth — Parathesis 12. Imbricate perianth — 13 13. Filaments partially united at base, free portion of filaments originating from near the middle of the corolla (sometimes filaments completely fused to the corolla); inflorescence generally racemose — Cybianthus 13. Filaments free, inserted near the base of the corolla, inflorescence usually in corymb or panicle — Ardisia Key differences from similar families: In the Neotropics, the Myrsinaceae differs of the Theophrastaceae for absences of the appendages (probably staminodes) inserted on corolla, alternating with petal lobes or for semi-inferior ovary (Samolus traditionally includes in Primulaceae) and of the Primulaceae (Primula, exclusively cultivated in Neotropics) for scapose inflorescence and flower commonly with heterostyly. Trees to shrubs with alternate, entire, and coriaceous leaves, without stipules, petiole short.  Sepals and petals partially connate.  Fruit a single seeded drupe. Distinguishing characters (always present): Leaves, flowers and fruit with secretory cavities, appearing as dark dots or lines. Stamens (or staminodes) epipetalous. Ovary syncarpous, superior, locule 1, with free -central placentation. Style 1.

Distribution

Native, naturalised (weeds) and cultivated. Anagallis L. - southern South America (including naturalised species). Ardisia Sw.- more diverse in southern Central American than in South America. Ctenardisia  Ducke - few species in Central and South America. Cybianthus Mart. - mostly middle-elevation cloud forests but also in Chocó and lowland Amazonia. Geissanthus Hook.f. - Andean. Gentlea Lundell - restricted to cloud and elfin forests. Grammadenia Benth. - Caribbean Island and northwestern South America. Heberdenia Banks ex DC. - one species in Mexico. Lysimachia L. - one species of western Peruvian Andes. Parathesis (DC.) Hook.f. - Caribbean Island and northwestern South America. Rapanea Aubl. - typical of middle-elevation forests especially in rather exposed situations. Solonia Urb. - monotypic species of Cuba. Synardisia (Mez) Lundell - one species Mexico to Nicaragua. Wallenia Sw. - Caribbean Islands.

Note

Very little is known about the reproductive biology of the family.  Most species seem to be pollinated by insects, and the fuits are usually dispersed by birds.  Autogamy has been documented in Ardisia and may occur in other genera as well. The family is of little economic importance.  Locally, the wood of several species is used for rustic construction (fencing) and as fuel (wood/charcoal), and fruit with thick mesocarps are often eaten. Species of Ardisia, Cyclamen, Lysimachia and Rapanea are cultivated as ornamental plants, including trees. Some species of Anagallis are introduced weeds.  In other regions of world some species are used in folk medicine. Notes on delimitation: Historically the Myrsinaceae has usually been included in the order Primulales near the Primulaceae and Theophrastaceae. Recent phylogenetic studies suggest, however, that Myrsinaceae is resolved in the Ericales, closely related to groups within the Primulaceae.  At present, the Myrsinaceae includes the genera Anagallis, Cyclamen, Lysimachia and Pelletiera which were previously considered to belong to the Primulaceae. Number of genera: 14 native genera and one genus exclusively cultivated:  Anagallis L.(including naturalised species). Ardisia Sw. (including Stylogyne and others segregrate genera, many of which have not been accepted). Ctenardisia Ducke. Cybianthus Mart. Cyclamen L. (cultivated). Geissanthus Hook. Gentlea Lundell Grammadenia Benth. Heberdenia Banks ex A. DC. Parathesis (A. DC.) Hook. Pelletiera A. St-Hil. Rapanea Aubl. Solonia Urb. Synardisia (Mez.) Lundell Wallenia Sw.

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

Morphology General Habit

Trees, shrubs or rarely subherbaceous

Morphology Leaves

Leaves alternate, rarely subopposite, simple, punctate or with schizogenous lines

Morphology Reproductive morphology Flowers

Flowers small, usually hermaphrodite, in clusters, racemes or panicles

Morphology Reproductive morphology Flowers Calyx

Sepals free or connate, often gland-dotted, valvate, imbricate or contorted, persistent

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous or rarely petals free; lobes contorted, imbricate or valvate

Morphology Reproductive morphology Flowers Androecium

Stamens the same number as and opposite the petals, the filaments more or less adnate to the corolla; anthers introrse, opening by slits or apical pores

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior to half-inferior, 1-celled; style simple; ovules numerous, on a free central placenta

Morphology Reproductive morphology Fruits

Fruit a berry or drupe, rarely irregularly dehiscent

Morphology Reproductive morphology Seeds

Seeds with smooth or rarely ruminate endosperm, with the embryo sometimes placed transversely

Luís C. Bernacci (2009). Neotropical Myrsinaceae.

Morphology

Description

Trees, treelets to shrubs or subshrubs, herbs or epiphytes; leaves, flowers and fruit with secretory cavities appearing as dark dots or dashes.  Leaves simple , alternate , subopposite, opposite or pseudowhorled; blades often coriaceous , margins entire to variously serrate , stipules absent; petioles usually short, occasionally absent.  Inflorescences terminal or axillary ; racemes, panicles to simple or compound corymbs, fascicles or verticils, or solitary flowers.  Flowers bisexual or unisexual, with staminodes (on pistillate flowers) and pistillodes (on staminate flowers); partially connate sepals and petals, rarely petals free , epipetalous, distinct or connate stamens, and connate carpels (3) 4-5 (6); anthers dehiscing by longitudinal slits to pores; ovary superior , with free -central placentation; ovules few to several.  Fruit a single-seeded drupe or a valved or opercular capsule with few to many seeds.

General Description

Notes on delimitation

• Historically the Myrsinaceae has usually been included in the order Primulales near the Primulaceae and Theophrastaceae.
• Recent phylogenetic studies suggest, however, that Myrsinaceae is resolved in the Ericales, closely related to groups within the Primulaceae.  At present, the Myrsinaceae includes the genera Anagallis, Cyclamen, Lysimachia and Pelletiera which were previously considered to belong to the Primulaceae.

General notes

• Very little is known about the reproductive biology of the family.  Most species seem to be pollinated by insects, and the fuits are usually dispersed by birds.  Autogamy has been documented in Ardisia and may occur in other genera as well.
• The family is of little economic importance.  Locally, the wood of several species is used for rustic construction (fencing) and as fuel (wood/charcoal), and fruit with thick mesocarps are often eaten.
• Species of Ardisia, Cyclamen, Lysimachia and Rapanea are cultivated as ornamental plants, including trees.
• Some species of Anagallis are introduced weeds.  In other regions of world some species are used in folk medicine.

Number of genera

14 native genera and one genus exclusively cultivated: 

• Anagallis L.(including naturalised species).
• Ardisia Sw. (including Stylogyne and others segregrate genera, many of which have not been accepted).
• Ctenardisia Ducke.
• Cybianthus Mart.
• Cyclamen L. (cultivated).
• Geissanthus Hook.
• Gentlea Lundell
• Grammadenia Benth.
• Heberdenia Banks ex A. DC.
• Parathesis (A. DC. ) Hook.
• Pelletiera A. St-Hil.
• Rapanea Aubl.
• Solonia Urb.
• Synardisia (Mez.) Lundell
• Wallenia Sw. 

Status

• Native, naturalised (weeds) and cultivated.

Distribution

Distribution in the Neotropics

• Anagallis L. - southern South America (including naturalised species).
• Ardisia Sw.- more diverse in southern Central American than in South America.
• Ctenardisia  Ducke - few species in Central and South America.
• Cybianthus Mart. - mostly middle-elevation cloud forests but also in Chocó and lowland Amazonia.
• Geissanthus Hook.f. - Andean.
• Gentlea Lundell - restricted to cloud and elfin forests.
• Grammadenia Benth. - Caribbean Island and northwestern South America.
• Heberdenia Banks ex DC. - one species in Mexico.
• Lysimachia L. - one species of western Peruvian Andes.
• Parathesis (DC.) Hook.f. - Caribbean Island and northwestern South America.
• Rapanea Aubl. - typical of middle-elevation forests especially in rather exposed situations.
• Solonia Urb. - monotypic species of Cuba.
• Synardisia (Mez) Lundell - one species Mexico to Nicaragua.
• Wallenia Sw. - Caribbean Islands.

Diagnostic

Distinguishing characters (always present)

• Leaves, flowers and fruit with secretory cavities, appearing as dark dots or lines.
• Stamens (or staminodes) epipetalous.
• Ovary syncarpous, superior, locule 1, with free -central placentation.
• Style 1.

Other important characters

• Trees to shrubs with alternate, entire, and coriaceous leaves, without stipules, petiole short. 
• Sepals and petals partially connate. 
• Fruit a single seeded drupe.

Key differences from similar families

• In the Neotropics, the Myrsinaceae differs of the Theophrastaceae for absences of the appendages (probably staminodes) inserted on corolla, alternating with petal lobes or for semi-inferior ovary (Samolus traditionally includes in Primulaceae) and of the Primulaceae (Primula, exclusively cultivated in Neotropics) for scapose inflorescence and flower commonly with heterostyly.

Useful tips for generic identification

• Habit (epiphytes, herbs, trees or shrubs).
• Disposition of leaves.
• Type of inflorescences.
• Petals: free or partially connate.
• Fusion and insertion of filaments.
• Anthers exserted or included.
• Type and texture of the corolla.
• Imbrication of the calyx (regular or irregular).
• Imbrication of the corolla (valved or imbricated).

Key to genera of neotropical Myrsinaceae.

1. Epiphytes; leaves very narrow, sessile, leaf apices mucronate — Grammadenia
1. Not epiphytes; leaves not like above — 2

2. Herbs; flowers solitary ... 3
2. Trees to shrubs; flowers in inflorescences ... 4

3. Leaves opposite; seeds not winged — Anagallis
3. Leaves alternate; seeds winged — Lysimachia

4. Flower in fascicles or verticils — Rapanea
4. Flowers not in fascicles or verticils — 5

5. Petals free — 6
5. Petals partially connate... 7

6. Filaments united (Mexico) — Heberdenia
6. Filaments free (Cuba) — Solonia

7. Exserted anthers (filaments longer than the corolla) — 8
7. Included anthers (filaments shorter than the corolla) — 9

8. Tubular coriaceous corolla, remains of styles not persistent on fruit (Caribbean) — Wallenia
8. Not tubular and not coriaceous corolla, remains of long styles persistent on fruit (cloud and elfin forests) — Gentlea

9. Plants with long glandular trichomes, campanulate corolla — Synardisia
9. Plants without glandular trichomes, not campanulate corolla — 10

10. Umbellate clusters of flowers arranged in panicles — Ctenardisia
10. Others kinds of inflorescences — 11

11. Calyx closed in bud, irregularly rupturing into 2-7 lobes (also visible in fruit) — Geissanthus
11. Calyx regularly divided — 12

12. Valvate perianth — Parathesis
12. Imbricate perianth — 13

13. Filaments partially united at base, free portion of filaments originating from near the middle of the corolla (sometimes filaments completely fused to the corolla); inflorescence generally racemose — Cybianthus
13. Filaments free, inserted near the base of the corolla, inflorescence usually in corymb or panicle — Ardisia

Notable genera and distinguishing features

• Ardisia: shrubs and small trees.  The inflorescence is paniculate, the lateral branches with the usually 5-parted flowers arranged in glomerules or cymes, typically more or less umbellate to much reduced ramiflorous inflorescence. The stamens are always included, the anthers rather long, and the filaments usually short.  Commonest in middle-elevation cloud forests and more diverse in southern Central American than in South America.
• Cybianthus: from small subshrubs to subcanopy trees, with racemose inflorescence, sometimes slightly branched at the apex, corolla rotate or tabulate.
• Rapanea: Small trees typical of middle-elevation forests especially in rather exposed situations.  Distinctive in the sessileaxillary and ramiflorus flowers, typically densely clustered along the twigs below the leaves on suppressed short-shoots.
• Parathesis: shrubs and small trees.  Frequently the branchlets and lower leaf surface near midrid have stellate pubescence.  The flowers, always 4-merous, have unusually large yellow anthers and are always in panicles with valvateperianth and densely pubescentcalyx and petals. The fruit is distinctive in being ribbed and is also characterized by the pubescentpersistentcalyx lobes.

Literature

Important literature

Agostini, G. 1980.  Una nueva classificación del género Cybianthus (Myrsinaceae). Acta Bot. Venez. 10(2): 129-185.

Anderberg, A.A. & Stahl, B.1995. Phylogenetic interrelations in the order Primulales, with special emphasis on the family circumscriptions. Canad.J.Bot.73: 1699-730.

Bernacci, L.C. & Jung-Mendaçolli, S.L. 2001.  Considerações taxonômicas e novas combinações em Ardisia Swartz (Myrsinaceae) do Sudeste do Brasil. Acta Bot. Bras. 14(3): 243-250.

Freitas, M.F. 2003. Estudos taxonômicos das espécies de Myrsine L. (Myrsinaceae) nas regiões sudeste e sul do Brasil. Tese de Doutorado. UNICAMP.

Jung-Mendaçolli, S.L. 2005.  Myrsinaceae. In: Wanderley, M.G.L. (ed.). Flora Fanerogâmica do Estado de São Paulo. São Paulo, Rima, vol. 4, p. 279-300.

Kallersjo, M. et al. 2000. Generic realignment in primuloid families of the Ericales s.l. (Angiosperms): a phylogenetic analysis based on DNA sequences of rbcl and ndhF. Amer.J.Bot.87: 1325-1341.

Lourteig, A. 1942. Primulaceae argentinae. Lilloa 8: 231-67.

Lourteig, A. 1967. Primuláceas. In: P.R.Reitz (ed.), Flora ilustrada Catarinense, Pp. 1-17. Itajaí, Brasil: Herbario "Barbosa Rodrigues".

Lundell, C.L. 1966a. Myrsinaceae. In: P.C.Standley & L.O.Williams (eds.), Flora of Guatemala, Fieldiana Bot. 24(8): 135-200.

Lundell, C.L. 1966b. The genus Parathesis of the Myrsinaceae. Contr.Texas Res. Found., Bot.Stud. 5: 1-06.

Lundell, C.L. 1971. Myrsinaceae. In: Woodson, R.E. (ed.), Flora of Panama. Ann.Missouri Bot.Gard. 58: 285-353.

Otegui, M. 1998.  Sinopsis del género Myrsine (Myrsinaceae) en el cono sur de América del Sur. Candollea 53(1): 133-157.

Pipoly, J.J. 1987.  A systematic revision of genus Cybianthus subgen. Gramnadenia (Myrsinaceae). Mem. New York Bot. Gard. 43: 1-75.

Pipoly, J.J. 1991.  Notas sobre el género Ardisia Sw. en Colombia. Caldasia 16(78): 277-284.

Pipoly, J.J. 1992. The genus Cybianthus (Myrsinaceae) in Ecuador and Peru. Sida 18: 1-160.

Pipoly, J.J. 1993.  Notes on Cybianthus subgenus Cybianthus (Myrsinaceae) in southeasten Brazil. Novon 3(4): 459-462.

Pipoly, J.J. 1996.  New species of Ardisia (Myrsinaceae) from Ecuador and Peru. Sida 17(2): 445-458.

Pipoly, J.J. 1998.  The genus Cybianthus (Myrsinaceae) in Ecuador and Peru. Sida 18(1): 1-60.

Ribeiro et al. 1999.  Flora da Reserva Ducke. Guia de identificaçao deas plantas vasculares de uma floresta de terra-firme na Amazonia Central.  INPA, Manaus.

Ricketson, J.M. & Pipoly, J.J. 1997.  Nomenclatural notes and a synopsis of Mesoamerican Stylogyne (Myrsinaceae). Sida 17(3): 591-597.

Ståhl, B.1990. Primulaceae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador vol. 39: 23-35. Goteborg, Sweden: Department of Systematic Botany, University of Goteborg.

Stearn, W.T. 1969. A synopsis of Jamaican Myrsinaceae. Bull.Brit.Mus.(Nat.Hist.)Bot. 4: 145-78.

Myrsinaceae, P. Halliday. Flora of Tropical East Africa. 1984

Morphology General Habit

Trees, shrubs, climbing shrubs or rarely subherbaceous, usually glandular-hairy or glabrous

Morphology Leaves

Leaves alternate, occasionally opposite or verticillate, entire or variously toothed, very rarely pinnately-partite (Ardisia), with or without visible resinous dots or streaks; stipules absent

Morphology Reproductive morphology Inflorescences

Inflorescence cymose, umbellate, fasciculate, racemose or paniculate, axillary or on older wood

Morphology Reproductive morphology Flowers

Flowers ?,?orhermaphrodite  small, regular, often marked with darker resinous dots or streaks

Morphology Reproductive morphology Flowers Calyx

Calyx hypogynous (adnate to ovary in >i>Maesa), 4-6-lobed; lobes usually contorted or imbricate, sometimes valvate, usually persistent, often ciliate or punctate

Morphology Reproductive morphology Flowers Corolla

Corolla usually with petals united into a short tube, occasionally free (>i>Embelia), rarely campanulate; lobes or free petals (3-)4-6(-7), contorted, imbricate or occasionally valvate

Morphology Reproductive morphology Flowers Androecium

Stamens equal in number to the petals and opposite to them, attached to the corolla, free or±joined, erect or incurved (especially in bud); anthers sagittate, oblong or shortly ovoid, 2-thecous, introrse, splitting longitudinally or with apical pores; staminodes in ? flower often almost as large as stamens, rarely present in ? flower

Morphology Reproductive morphology Flowers Gynoecium

Ovary globose or ovoid, superior or semi-inferior and free, to inferior and adnate to calyx (>i>Maesa), 1-locular; ovules few to many, inserted on or embedded in the shortly stalked or free-central placenta, 1-several-seriate; style short or long; stigma acute to discoid or capitate, sometimes lobed

Morphology Reproductive morphology Fruits

Fruit a drupe or berry, globose to ovoid, l(-few)-seeded, indehiscent

Morphology Reproductive morphology Seeds

Seed globose or angular, mostly with smooth endosperm (occasionally ruminate); embryo straight or curved; cotyledons subequal