Polygalaceae Hoffmanns. & Link

Polygalaceae, Jorge Paiva. Flora of Tropical East Africa, 2007

Morphology General Habit

Annual or perennial herbs, shrublets or shrubs, rarely small trees or woody climbers

Morphology Leaves

Leaves usually alternate, rarely opposite, fascicled or verticillate, sometimes in basal leaf-rosettes, sessile or shortly petiolate, simple, entire, usually exstipulate (stipules when present glandular or conical-truncate)

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, zygomorphic, in 1–several-flowered, terminal, axillary or pseudolateral racemes, secondarily spiciform or corymbiform

Morphology Reproductive morphology Flowers Calyx

Sepals 5, free, sometimes the 2 anterior ones joined; 3 external, 2 anterior (inferior) alike and 1 superior larger; 2 internal (lateral) ones often petaloid (wing-sepals), sometimes all nearly alike

Morphology Reproductive morphology Flowers Corolla

Petals 3–5, ± joined, at least near the base, or free, subequal or unequal and the 2 lateral ones often absent or vestigial; the 2 upper ones free or joined to the lower one, usually keel-shaped (keel or carina), often fringed (crested) or appendaged at the tip

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens (4–)5–10, but usually 8; filaments usually united into a slit tube; anthers basifixed, introrse, monothecate

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, (1–)2–5-locular, usually with 1 pendulous ovule per loculus [4 or (6–)8–20, very rarely ± 40 in Xantophylleae]; style terminal, simple; stigma with 2 spreading arms, the posterior (inferior) one papillose

Morphology Reproductive morphology Fruits

Fruit a capsule, samara or drupe

Morphology Reproductive morphology Seeds

Seeds often hairy, usually carunculate; endosperm mostly present, oily and proteinaceous.

Note

Of the 4 genera of the area, Carpolobia G.Don belongs to the tribe Carpolobieae B.Eriksen and the other three ( Securidaca L., Muraltia DC. and Polygala L.) to the tribe Polygaleae.

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

Distribution

A family of 21 genera and 800–1000 species, distributed worldwide throughout temperate and tropical areas. There are seven genera in New Guinea, of which two are arborescent.

Morphology General Habit

Herbs, climbers, shrubs and trees, sometimes with swollen nodes

Morphology General Indumentum

Indumentum when present of simple hairs

Morphology Leaves Stipules

Stipules absent, extra-floral nectaries sometimes present at junction of petiole and stem

Morphology Leaves

Leaves simple, spirally arranged, sometimes alternate or whorled, sometimes reduced or absent, petiolate or sessile

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary, usually unbranched, sometimes thyrsoid or in fascicles

Morphology Reproductive morphology Flowers

Flowers bisexual, zygomorphic, each subtended by a bract and 2 bracteoles; sepals 5, quincuncial, free or variously fused, usually with the lowest 2 lobes fused or the 2 lateral lobes expanded and petaloid; petals 3, sometimes 5, free or variously fused, lower petal commonly keel-shaped, pouched, lobed or with a fringed crest; stamens usually 8, filaments often fused, often adnate to petal base, anthers basifixed, 1- or 2-locular, usually dehiscing by an apical pore; intra-staminal disk sometimes present; ovary superior, usually sessile, rarely stipitate, usually 2-locular and with 1 ovule per locule, placentation usually apical. Fruit a berry, capsule, samara or drupe

Morphology Reproductive morphology Seeds

Seeds usually arillate, sometimes winged.

Persson, C. & Eriksen, B. (2009). Neotropical Polygalaceae.

Morphology

Description

Annual or perennial herbs, subshrubs, shrubs, trees or lianas (lianas sometimes with anomalous growth); branches sometimes with short emergences on young shoots (Moutabea Aubl.), or ending in a thorn (some species of Acanthocladus Klotzsch ex Hassk.). Stems sometimes with stalked or sessile glands ( extrafloral nectaries) at leaf and inflorescence nodes (sometimes incorrectly called stipules) or on petioles. Stipules absent. Leaves usually alternate , rarely opposite or verticillate , simple , sometimes scaly , needle-like; margins entire . Inflorescences terminal or axillary , usually racemose or variously elaborated panicles, rarely spikes, heads, or of solitary flowers; bracts and bracteoles often present, persistent or caducous . Flowers usually zygomorphic , rarely actinomorphic (Diclidanthera Mart.), bisexual ; sepals usually 5, usually distinct, quincuncial, the two inner sepals usually larger and petaloid (wings) but sometimes almost similar to outer sepals; petals (4-)5 (Moutabeae), or 3 (Polygaleae), sometimes apically equipped with a terminal tuft of narrow lobes (crest); androecium with (6-)8(-10) stamens, the filaments generally connate , at least in their lower half, forming a cleft sheath or tube, usually adnate to petals, the anthers basifixed, bi- to tetrasporangiate, usually with a falcate slit forming two apical or subapical pores, or less often forming a marginal or ventral, longitudinal slit; pollen polycolporate; annular nectary often present at base of ovary (possessing or lacking a process ) or on staminal sheath (as in Bredemeyera Willd.); ovary superior , carpels 2-8, locules 2 or pseudomonomerous in Polygaleae, 2-8 in Moutabeae; ovules 1 per locule , ( sub )apical, pendulous, anatropous, bitegmic. Fruits usually capsules, often with a ± pronounced wing (e.g., Bredemeyera and some species of Polygala L.), sometimes berries (Moutabeae), drupes (e.g. Monnina Ruiz & Pav.), dry and indehiscent (sometimes referred to as wingless samaras), or samaras, the samaras single- winged (Securidaca L.), or double- winged (Pteromonnina B.Eriksen and Ancylotropis B.Eriksen). Seeds 1 per locule , glabrous or pilose , sometimes villous or with a comb of long hairs (e.g. Bredemeyera); an exostome aril (caruncle) sometimes present; endosperm typically abundant.

Distribution

Distribution in the Neotropics

• Acanthocladus Klotzsch ex Hassk.: distributed in Colombia, Ecuador, Peru, Brazil to Argentina.
• Ancylotropis B.Eriksen: E Brazil.
• Badiera DC.: distributed in West Indies, Central America to Ecuador.
• Barnhartia Gleason: restricted to the Guianas, S Venezuela and Amazonian Brazil.
• Bredemeyera Willd.: distributed from Mexico to temperate parts of Argentina
• Diclidanthera Mart.: distributed in Colombia, Venezuela, Guyana, Brazil, Peru and Bolivia.
• Monnina Ruiz & Pav.: distributed from Mexico to Bolivia.
• Moutabea Aubl.: distributed from Costa Rica to Bolivia.
• Polygala L.: widely distributed in the Neotropics.
• Pteromonnina B.Eriksen: disjunctly distributed between the Andes from Ecuador to Chile and eastern South America from north-eastern Brazil to Argentina. One species with a disjunct range is also distributed (by humans?) in Mexico.
• Securidaca L.: distributed in the West Indies and from Mexico to Paraguay.

Diagnostic

Distinguishing characters (always present)

• Simple leaves.
• Stipules absent.
• Flowers hypogynous.
• Pistil with 1 ovule/locule.

Key differences from similar families

Species with papilionaceous flowers differ from Leguminosae (=Fabaceae) subfam. Papilionoideae in having:

• Simple leaves (vs. usually compound).
• A 3-merouscorolla with a standard of two petaloid sepals (vs. usually a 5-merouscorolla including a standard of one petal).
• One ovule per locule in 2-8 carpellate ovaries (vs. usually several ovules per locule and 1-carpellate ovaries).

Other important characters

• Alternate leaves (opposite in Acanthocladus, whorled in some Polygala).
• Flowers usually zygomorphic; papilionaceous in the vast majority (tribe Polygaleae) with 3 petals and 5 sepals, two of which are larger and petaloid.
• Stamens generally 8, sometimes fewer or up to 10, adnate to the petals.

Notable genera and distinguishing features

• Monnina: papilionaceous flowers and drupes.
• Polygala: usually herbaceous and with papilionaceous flowers and capsule.
• Securidaca: papilionaceous flowers and unilaterally winged samaras.

Useful tips for generic identification

Key to genera of Neotropical Polygalaceae

1. Flowers papilionaceous with a trimerous corolla and two petaloid sepals — 2
1. Flowers not papilionaceous; corolla pentamerous ... 9

2. Fruit dehiscent (capsule) — 3
2. Fruit indehiscent (drupe, samara, etc.) — 6

3. Seeds usually provided with a coma (hairs often as long as the seed), then plant leafless or leaves reduced to minute scales ... Bredemeyera
3. Seeds glabrous or with short hairs — 4

4. Plants herbaceous, sometimes woody; calyx usually persistent; when caducous, the keel of the corolla with a crest or a beak ... Polygala
4. Plants woody; calyx caducous; crest or beak on keel of the corolla absent — 5

5. Leaves opposite, sub-opposite or alternate; branches often spine-tipped; capsule with subglobose locules, woody... Acanthocladus
5. Leaves alternate, branches without spines; capsule laterally compressed, sub-coriaceous... Badiera

6. Fruit juicy ... Monnina
6. Fruit dry, sometimes winged — 7

7. Lianas or shrubs; fruit a unilaterally winged samara ... Securidaca
7. Herbs; fruit unwinged or bilaterally winged — 8

8. Keel apically deeply incised; style curved, apically entire with a tuft of hairs; nectary absent ... Ancylotropis
8. Keel apically emarginate; style geniculate, apically bifid without hairs; nectary unilateral ... Pteromonnina

9. Sepals free; ovary 2-3-locular ... Barnhartia
9. Sepals connate; ovary (2-)4-5(-7)-locular ... 10

10. Flowers distinctly zygomorphic; ovary usually (2-)4(-5)-locular ... Moutabea
10. Flowers actinomorphic or only slightly zygomorphic; ovary 5(-7)-locular ...Diclidanthera

General Description

General notes

The position and circumscription of several genera need to be elucidated and modern revisions of some genera, e.g. Securidaca, are still wanting.

Status

Native.

Notes on delimitation

• DNA-sequences of the trnL-F region and rbcL gene support the monophyly of Polygalaceae as well as its position in the order Fabales.

Number of genera

11 genera:

• Ancyloptropis: (2 spp.)
• Acanthocladus: (6 spp.)
• Badiera: (10-15 spp.)
• Barnhartia: (1 sp.)
• Bredemeyera: (ca 15 spp.)
• Diclidanthera: (8 spp.)
• Monnina: (ca 150 sp.)
• Moutabea: (8-10 spp.)
• Polygala: (ca 150 spp.)
• Pteromonnina: (ca 30 ssp.)
• Securidaca: (ca 80 sp.)

Literature

Important literature

Aymard, G.A. Berry, P.E. & Eriksen, B. 2004. Polygalaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych and Holst, B.K. (eds.) Flora of the Venezuelan Guayana, vol. 8. Missouri Botanical Garden, St. Louis.

Bernardi, L. 2000. Consideraciones taxonomicas y fitogeográficas acerca de 10 Polygalae Americanas. Cavanillesia Altera I: 1-456.

Blake, S. 1916. A revision of the genus Polygala in Mexico, Central America and the West Indies. Cambridge, MA: Harvard University Press.

Eriksen, B. 1993. A revision of Monnina subg. Pterocarya (Polygalaceae) in Northwestern South America. University of Gothenburg, Gothenburg, PhD thesis.

Eriksen, B. 1993. Phylogeny of the Polygalaceae and its taxonomic implications. Pl. Syst. Evol. 186: 33-55.

Eriksen, B., Ståhl, B. and Persson, C. 2000. Fam. 216. Polygalaceae. In: G. Harling and L. Andersson (eds). Flora of Ecuador, University of Gothenburg, Gothenburg.

Eriksen, B. and Persson, C. 2006(2007). Polygalaceae, pp. 345-363. In: K. Kubitzki (ed.) The Families and Genera of Vascular Plants vol. IX. Springer-Verlag, Berlin.

Forest, F., Chase, M.W., Persson, C., Crane, P.R. and Hawkings, J.A. 2007. The role of biotic and abiotic factors in evolution of ant dispersal in the milkwort family (Polygalaceae). Evolution 61-7: 1675-1694.

Marques, M. do C. 1980. Revisão das espécies do gênero Bredemeyera Willd. (Polygalaceae) do Brasil. Rodriguésia 67:3-33.

Marques, M. do C. 1996. Securidaca L. (Polygalaceae) do Brasil. Arch. Jard. Bot. Rio de Janeiro 34(1): 7-144.

Persson, C. 2001. Phylogenetic relationships in Polygalaceae based on chloroplast DNA sequences from the trnL-F region. Taxon 50: 763-779.

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

Morphology General Habit

Herbs, shrubs, climbers or rarely small trees

Morphology Leaves

Leaves alternate, rarely opposite, simple; stipules none

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, zygomorphic; pedicels often articulated

Morphology Reproductive morphology Flowers Calyx

Sepals 5, free, imbricate, the two inner larger, often petaloid, wing-like

Morphology Reproductive morphology Flowers Corolla

Petals 3–5, hypogynous, declinate, outer 2 free or united with the lowermost, upper 2 free or minute and scale-like or 0

Morphology Reproductive morphology Flowers Androecium

Stamens 8, rarely 5 or 4, monadelphous beyond the middle or rarely free, the sheath split above, often adnate to the petals; anthers erect, 1–2-celled, opening by an apical pore, rarely 2-valved or opening nearly to the base

Morphology Reproductive morphology Flowers Nectaries

Torus small or rarely expanded into an annular disk within the stamens

Morphology Reproductive morphology Flowers Gynoecium

Ovary free, 2-celled or rarely 1-celled or 3–5-celled; style simple; ovules solitary in each cell, rarely more, pendulous

Morphology Reproductive morphology Fruits

Fruit a capsule, samara or drupe

Morphology Reproductive morphology Seeds

Seeds often pilose, with a conspicuous strophiole; endosperm mostly present; embryo straight

Distribution

Fairly abundantly represented in our area

Polygalaceae, A. W. Exell. Flora Zambesiaca 1:1. 1960

Morphology General Habit

Small trees, woody climbers, shrublets or perennial or annual herbs

Morphology Leaves

Leaves usually alternate, simple, entire, exstipulate

Morphology Reproductive morphology Flowers Calyx

Sepals 5; two lateral (interior) ones often petaloid (wings); two anterior ones sometimes joined

Morphology Reproductive morphology Flowers Corolla

Petals 3–5; two upper ones free or joined to the lower one (keel); two lateral ones free, often absent or vestigial

Morphology Reproductive morphology Flowers Androecium

Stamens usually 5–8 (rarely 4); filaments usually united in a slit tube

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, usually 2-locular (1–5), with 1 pendulous ovule in each loculus

Morphology Reproductive morphology Fruits

Fruit a capsule, samara or drupe

Morphology Reproductive morphology Seeds

Seeds often sericeous, usually carunculate; endosperm usually present