Primulaceae Batsch ex Borkh.

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

Habit

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

Leaves

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

Flowers

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

Calyx

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

Corolla

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

Androecium

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

Gynoecium

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

Fruits

Fruit a capsule with valvate or circumscissile dehiscence, rarely indehiscent

Seeds

Seeds few to numerous, often angular

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

Habit

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

Leaves

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

Flowers

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

Calyx

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

Corolla

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

Androecium

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

Gynoecium

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

Fruits

Fruit a capsule, valvate or circumscissile or rarely indehiscent

Seeds

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

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).

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

Habit

Trees, shrubs and lianes

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

Inflorescences

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

Flowers

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

Calyx

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

Corolla

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

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

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

Fruits

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

Seeds

Seeds with copious, often ruminate, endosperm

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

Habit

Trees, shrubs or rarely subherbaceous

Leaves

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

Flowers

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

Calyx

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

Corolla

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

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

Gynoecium

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

Fruits

Fruit a berry or drupe, rarely irregularly dehiscent

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

Habit

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

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

Inflorescences

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

Flowers

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

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

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

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

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

Fruits

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

Seeds

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