RUBIACEAE

Piero G. Delprete

The fourth largest family of flowering plants, with 600–650 genera and more than 13,000

species. A predominantly tropical family with a few genera in temperate regions; 217–220

genera and more than 5,000 species in the Neotropics. These are most commonly terrestrial,

rarely epiphytic, rheophytic, semi-aquatic (amphibious) or aquatic, presenting a wide range of

habits, such as herbs, shrubs, small to large trees, and herbaceous to woody climbers. Climbers

are represented by 31 genera and a total of 259 species occurring in a wide range of habitats,

including moist, wet or dry forests, scrublands, savannas, and sometimes open disturbed biomes;

0–2,400 m elevation.

Diagnostics: In the absence of fertile material, climbing Rubiaceae are distinguished from

climbers in other families that have simple, commonly opposite (rarely whorled) leaves, with

entire margins by the interpetiolar stipules, and the watery, clear exudates of stems. Climbing

mechanisms are diverse but never include tendrils. Twining woody Rubiaceae may be confused

with species of Trigonia (Trigoniaceae) but in the absence of fertile material, Trigonia can be

distinguished by the stem cross section with a square medulla and the presence of four, shallow

phloem wedges or arcs, a combination of characters never present in climbing Rubiaceae.

General Characters

1. STEMS. Herbaceous or woody with moderate secondary growth, some species attaining 25–

30 m in length and up to 20 cm in diam. in some genera (e.g., Schradera, Uncaria,

Chiococca, and Manettia). Most climbing neotropical Rubiaceae are herbaceous, although

sometimes they are woody at the base, and only a few genera are lianas with thick, woody
Figure 217. Cross sections of stems in Rubiaceae. A. Simple stem in Uncaria guianensis, young stems
quadrangular (see square medulla) becoming cylindrical with age. B. Asymmetrical, lobed stem in Randia sp.
C. Sub-quadrangular stem with deep phloem wedges in Hexasepalum sarmentosum. D. Cylindrical stems in
Schradera exotica, axial elements divided in radial segments. E. Neoformations in Chiococca alba during late
stages of secondary growth. F. 4-lobed, fistulose in Emmeorhiza umbellata. Photos by P. Acevedo.
Figure 218. Climbing mechanisms in Rubiaceae. A. Root-climber, Malanea macrophylla. B. Scrambler,
Hexasepalum sarmentosum. C. Scrambler aided by recurved, hook-like thorns, Uncaria guianensis. D
Scrambler aided by straight thorns in Randia retroflexa. Photos by P. Acevedo.
 stems throughout (e.g., Chiococca, Guettarda, Manettia, Schradera, and Uncaria). Stems are

cylindrical, quadrangular, lobed, or asymmetrical in cross section (Figure 217). Most genera

have regular anatomy with xylem forming a continuous cylinder, however, the following

anatomical variations have been documented in the family: neoformations in Chiococca alba

(L.) Hitchc. (Figure 217E); phloem wedges in Hexasepalum sarmentosum (Sw.) J.H. Kirkbr.

& Delprete (Figure 217C); deeply lobed stems in Randia and Emmeorhiza (Figure 217B, F);

and axial elements divided in radial segments in Schradera (Figure 217D). Metcalfe & Chalk

(1957) report that stems in Randia asperifolia (Sandw.) Sandw. (Guyana and Suriname) has

successive, concentric cambia, however, we only have seen young stems with regular

anatomy in this species. Stems of Uncaria exhibit overlapping layers of sclerenchyma in the

cortex (Figure 217A).

2. EXUDATES. Generally watery, odorless, and colorless. The genus Salzmannia has stipular

colleters that produce resin droplets, which later covers the stems.

3. CLIMBING MECHANISMS. Most climbing Rubiaceae are scramblers that lean on and

grow over the phorophytes, these are sometimes aided by the presence of thorns (modified

axillary branches) that are straight, stout or acicular (2–4 per node in Randia, Figure 218D;

or 2 per node in Chomelia, Guettarda and Machaonia), or recurved and hook-like in Uncaria

(Figure 218C). Some Rubiaceae (e.g., Chiococca, Malanea, and Morinda) are shrubby in

early stages, later becoming scramblers. Twiners are present in Emmeorhiza, Manettia,

Paederia, Sabicea, and in some species of Chiococca, Cosmibuena, Malanea, Morinda, and

Randia. Root climbers are known in Malanea, Hillia and Schradera (the latter two genera

have dimorphic juvenile and adult stages).

4. STIPULES. An important diagnostic character at the generic level. They are interpetiolar in

all climbing genera, minute to large (sometimes foliaceous, e.g., Galium), entire, divided or

bifid, laciniate, or fimbriate, early or late caducous, persistent and withering on the stem, or

falling off by irregular fragmentation.

5. LEAVES. Simple, commonly opposite and decussate, less frequently ternate or 4 (6– 8) per

node (e.g., Galium). Leaf margins entire or exceptionally undulate. Leaf domatia, when

present, always on the abaxial side and located on secondary (less often tertiary) vein axils,

these with sparse or dense tufts of hairs, hairy pockets, ciliated pits, and crypts, ciliated or

not.

6. INFLORESCENCES. Most genera either have terminal or axillary inflorescences, but some

genera may have both, terminal and axillary inflorescences; inflorescence position represents

a good diagnostic character at the generic level. Sometimes, the inflorescences are frondose,

i.e., bearing leaf-like bracts (pherophylls) that subtend lateral branches. Flowers are produced

in cymes, panicles, umbels, racemes, spikes, heads, or sometimes solitary. In several

dioecious genera of the tribe Gardenieae, female flowers are solitary, while male flowers are

produced in inflorescences, e.g., Randia where male inflorescence are fasciculate (or rarely

uniflorous).

7. FLOWERS. Commonly heterostylous, actinomorphic or rarely zygomorphic (e.g.,

Coutarea), bisexual, rarely unisexual or functionally unisexual where plants are dioecious or

polygamous-dioecious, mostly 4–6(–10)-merous in climbing genera. Calyx commonly

tubular and distally truncate, undulate, toothed or lobed, or sometimes the tube reduced or

absent with free lobes; sepals of similar shape and size, slightly unequal or conspicuously

unequal; sometimes one or several calyx lobes are enlarged into colored calycophylls that
 may serve to attract pollinators and/or to aid seed dispersal; corolla sympetalous, tubular,

hypocrateriform, infundibuliform, campanulate or rotate, with valvate, contorted or imbricate

aestivation (a generic diagnostic character); stamens as many as the corolla lobes and

alternate to them, with dorsifixed or basifixed anthers with longitudinal dehiscence; ovary

inferior, 2(–7)-locular, surrounded by a hypanthium, placentation axile, rarely incompletely

axile or parietal (e.g., Randia), with ovules solitary or numerous.

8. FRUITS. Simple or rarely multiple (e.g., Morinda), including berries, either fleshy (e.g.,

Sabicea), leathery or thinly woody with a gelatinous pulp (e.g., Randia); drupes with woody

pyrenes (e.g., Chiococca, Palicourea); or dry capsules, either loculicidal (e.g., Rondeletia) or

septicidal (e.g., Emmeorhiza); schizocarps, either separating into free mericarps (e.g.,

Machaonia) or with mericarps suspended by thin carpophores (e.g., Paederia).

9. SEEDS. One to many per locule; one (rarely two) per pyrene in drupes, or mericarp in

schizocarps, and sometimes in capsules; numerous in berries and capsules. They are smooth

or sculptured (e.g., reticulate, foveolate, etc.), wingless, rarely winged (e.g., Coutarea and

Hintonia) or comose (e.g., Hillia).

USES

Rubiaceae is a family with great economic importance, being the source of coffee, one of

the most important crops in the world. Many genera are the source of commercial and traditional

medicines. In addition, many species are cultivated as ornamentals for their beautiful flowers,

foliage and forms, and several genera are a significant source of timber in the tropics. Climbing

Rubiaceae are not the exception, with several species cultivated as ornamentals in private and

public gardens (e.g., Chiococca, Guettarda, Manettia and Nertera). Other species are used in
traditional medicine, e.g., the bark of Coutarea hexandra (Jacq.) K. Schum. as a secondary

source of quinine for the treatment of malaria; the roots, stems and fruits of species of Morinda

as a tonic, appetite stimulant, purgative or aphrodisiac; the bark, leaves and fruits of the

neotropical species of Uncaria, the most celebrated medicinal climbing Rubiaceae throughout

tropical America (commonly known as uña de gato, cat claw, or gambier) are used as a

sustainable source of tannins, iridoid glycosides and alkaloids for the treatment of diarrhea,

inflammation, diabetes, rheumatic pains, acne, and respiratory problems (Azevedo et al. 2018;

El-Saber et al. 2020; Honório et al. 2018; Zanetti et al. 2020; Zhang et al. 2015).

Key to genera of climbing Rubiaceae

1. Plant with a strong fetid odor; fruit a laterally compressed, dry, schizocarp, with mericarps

suspended by thin carpophores upon splitting ............................................................. Paederia

1. Plant without fetid odor; fruit various, if schizocarpic, then not suspended by a carpophore

..................................................................................................................................................2

2. Stipules fimbriate, with (1)3–9 setae ...........................................................................................3

2. Stipules entire or bifid (not fimbriate) .........................................................................................6

3. Fruit schizocarpic, breaking up into 2 indehiscent mericarps................................. Hexasepalum

3. Fruit capsular, septicidal ..............................................................................................................4

4. Inflorescences terminal, paniculate, with secondary branches terminating in many-flowered

umbels ................................................................................................................... Emmeorhiza

4. Inflorescences terminal and/or axillary, thyrsoid with flowers in fascicles or glomerules, or

capitate ......................................................................................................................................5
5. Inflorescence thyrsoid, with flowers in fascicles, or flowering branches with terminal and

axillary glomerules subtended by minute or small bracts; fruit capsular, dehiscing

septicidally from the apex down to the seed insertion, with both cocci dehiscent; seed

margin with a very narrow wing ............................................................................. Denscantia

5. Flowering branches with few- to many-flowered glomerules, these usually subtended by leaf-

like bracts; fruit schizocarpic, separating into two indehiscent mericarps, or a septicidal

capsule with two dehiscent cocci; seed margin without a narrow wing .................Spermacoce

6. Stipules truncate or bifid (connate or sheathing at base, with two triangular to linear lobes on

each side of the stem) ...............................................................................................................7

6. Stipules entire ..............................................................................................................................8

7. Stipules connate at base to sheathing, often connate to the petioles, truncate or bifid, with

interpetiolar appendages, these conical, linear, shortly bifid, bilobed, fimbriate at the apex,

with glandular apical projections or laminar and erose or laciniate ....................... Notopleura

7. Stipules sheathing at base, bifid, with two short to long triangular lobes on each side of the

stem, without interpetiolar appendages ...................................................................Palicourea

8. Stems with hook-like thorns, slightly or strongly recurved, or coiled ............................. Uncaria

8. Stems unarmed or with straight thorns ........................................................................................9

9. At least some portions of the stems armed with straight thorns ...............................................10

9. Stems unarmed ...........................................................................................................................13

10. Thorns 2–4 per node, simple, without vestigial nodes or reduced leaves; plant dioecious;

flowers functionally unisexual; male flowers in few- to many-flowered fasciculate or

corymbose inflorescence (rarely uniflorous); female flowers solitary; ovules many per locule

........................................................................................................................................Randia
10. Thorns 2 per node, simple or compound (sometimes dendroid in Machaonia), often with

vestigial nodes, sometimes with reduced leaves; plant monoecious or polygamous-

dioecious; inflorescences with flowers of both sexes or with hermaphroditic flowers; ovules

1 per locule .............................................................................................................................11

11. Fruit schizocarpic, chartaceous or thinly woody, splitting into 2 indehiscent mericarps with

corky ornamentation ...................................................................... Machaonia martineziorum

11. Fruit drupaceous, fleshy or succulent, with 2–7 woody pyrenes .............................................12

12. Flowers 4-merous, bisexual; corolla lobes valvate; style 2-lobed; ovary 2-locular; ovules

pendulous; drupes with 2 pyrenes ..................................................................Chomelia obtusa

12. Flowers (4–)5–7(–9)-merous, bisexual, or unisexual and bisexual on the same individual;

corolla lobes imbricate-quincuncial; style commonly subcapitate or shortly (2)4–7(–9)-

lobed; ovary (2–)4–7(–9)-locular; ovules inserted at the middle of the placenta, vertical;

drupes with (2–)4–7(–9) pyrenes .............................................................................. Guettarda

13. Leaves whorled, commonly 4 (6–8) per node, including 2 leaf-like stipules; stipules

commonly of same shape and size of leaves ................................................................ Galium

13. Leaves opposite or rarely ternate; stipules much smaller than leaves .....................................14

14. Inflorescence a globose head of many flowers with connate ovaries; infructescence forming a

fleshy, oblong-globose, ellipsoid to ovoid, multiple fruit ........................................... Morinda

14. Inflorescence not capitate, flowers free from each other; infructescence of free fruits ...........15

15. Fruits dry, capsular or schizocarpic .........................................................................................16

15. Fruits fleshy or spongy, baccate or drupaceous, or a leathery or thinly woody berry with a

gelatinous pulp in Randia .......................................................................................................24

16. Fruit schizocarpic, chartaceous, splitting into 2 indehiscent mericarps, and leaving a

persistent septum ............................................................................. Machaonia martinicensis

16. Fruits capsular, septicidal or loculicidal ..................................................................................17

17. Plant epiphytic or hemi-epiphytic, rarely terrestrial in juvenile stages; young branchlets

fleshy; leaf secondary veins inconspicuous; corolla large, tube 24–150 mm long; capsules

(2–)3–15 cm long ....................................................................................................................18

17. Plant terrestrial (rarely hemiepiphytic lianas in a few species of Manettia); young branchlets

woody or herbaceous; leaf secondary veins abaxially conspicuous or prominent; corolla

small to medium-sized, tube less than 24 mm long or 1.8–45 mm long in Manettia; capsules

less than 2.5 cm long ..............................................................................................................19

18. Stipules shallowly fused at base; calyx tubular, truncate or shallowly to deeply lobed; corolla

lobes imbricate (quincuncial) or rarely convoluted; capsules cylindrical, sometimes

lenticellate; seeds elliptic, dorsoventrally compressed, with a narrow rhombic,

membranaceous, entire or irregularly erose wing.................................................. Cosmibuena

18. Stipules free at base; calyx with free lobes, not tubular; corolla lobes convoluted (i.e., right-

contorted); capsules oblong to narrowly oblong, slightly compressed, septicidal, mericarps

secondarily splitting at apex; seeds fusiform, comose at distal end ................................ Hillia

19. Terrestrial, herbaceous or woody twiners, rarely hemiepiphytic lianas with adventitious roots

at nodes; inflorescence axillary, thyrsoid, dichasial or fasciculate, pauci- or multi-florous,

rarely uniflorous; capsule septicidal ........................................................................... Manettia

19. Terrestrial shrubs or small to medium-sized trees, rarely climbing shrubs or woody vines;

inflorescence terminal or axillary, cymose, paniculate or corymbose, pauci- or multi-florous;

capsules loculicidal (secondarily septicidal) or septicidal ......................................................20

20. Corollas campanulate or broadly infundibuliform, tube 2–7.5 cm long; filaments basally

connate, inserted at base of the corolla tube; capsules more than 1.8 cm long ......................21

20. Corollas hypocrateriform, tube up to 2 cm long; filaments free, inserted at median or distal

portion of the corolla tube; capsules less than 1.8 cm long ....................................................22

21. Flowers actinomorphic, 8-merous; corolla straight, with equal stamens; corolla tube 2.5–6 cm

long; capsule septicidal, ellipsoid or globose, 2–3.5 cm long, terete in cross section

.................................................................................................................... Hintonia octomera

21. Flowers zygomorphic, 6-merous; corollas curved, with unequal stamens; corolla tube 1.9–7.5

cm long; capsule loculicidal, obovate to oblong-ovate in outline, 1.8–7 cm long, laterally

compressed ................................................................................................ Coutarea hexandra

22. Corolla mouth glabrous, lacking a thick fleshy ring; capsules septicidal or loculicidal

............................................................................................................................ Arachnothryx

22. Corolla mouth with a thick fleshy ring, or a ring of yellow or white hairs; capsules loculicidal

................................................................................................................................................23

23. Corolla with a ring of yellow or white hairs within the mouth; seeds horizontal

(perpendicular to the septum), small, angular, globose, fusiform or falciform, narrowly or

partially winged; wing bipolar or unipolar, rarely absent ............................................ Rogiera

23. Corolla with a thick fleshy glabrous ring at mouth; seeds vertical, with a central hilum,

dorsoventrally compressed, fusiform or irregularly elliptic, winged; wing bipolar Rondeletia

24. Fruit drupaceous, fleshy or spongy, with 2–several woody pyrenes .......................................25

24. Fruit a fleshy, many-seeded berry ............................................................................................35

25. Herbaceous vines; stipules withering on the stem ..................................................... Didymaea

25. Woody vines (lianas), or scrambling shrubs; stipules not withering on stems ........................26
26. Stipules persistent, becoming indurate, distal portion falling off by fragmentation, entire,

dentate, or with 1–2 short acumens or with 1–2 aristae, these sometimes with 1–2 caducous

glandular appendages; pyrenes with 2 marginal germination slits extending to the middle

and sometimes also with several shorter dorsal slits ............................................... Eumachia

26. Stipules not becoming indurate; pyrenes with germination apertures not marginal ................27

27. Distal portion of branches covered with resinous exudates; apical stipules producing a small

globe of resinous exudate; scrambling shrubs ....................................................... Salzmannia

27. Distal portion of branches not covered with resinous exudates; apical stipules not producing a

globe of resinous exudate; terrestrial or epiphytic shrubs, woody vines, lianas, twining or

scrambling, sometimes shrubs, treelets or trees with scandent or clambering branches ........28

28. Stipules with interpetiolar appendages, these conical, linear, shortly bifid, bilobed or

fimbriate at apex, with glandular apical projections or laminar and erose or laciniate; fruits

with 2 pyrenes ......................................................................................................... Notopleura

28. Stipules without interpetiolar appendages; fruits with 2–7 pyrenes ........................................29

29. Stipules bifid, with two short to long triangular lobes on each side of the stem .........................

................................................................................................................................ Palicourea

29. Stipules entire, broadly to narrowly triangular, acuminate (rarely bifid at tip), narrowly ovate,

obovate, elliptic, oblong or oblanceolate ................................................................................30

30. Ovary 2-locular, with 2 ovules per locule, sometimes appearing as 4-locular due to thick

placenta within each locule; pyrenes (2–3) 4 per fruit ........................... Appunia megalantha

30. Ovary 2–7-locular, with 1 ovule per locule; pyrenes 2–7 per fruit ..........................................31

31. Corolla tubular-subcylindrical; filaments inserted at base of corolla tube (forming a minute

tube at base of corolla); fruits with 2 pyrenes, or rarely with 1 pyrene (by abortion) ............32
31. Corolla hypocrateriform or infundibuliform; filaments inserted just below or at the corolla

mouth; fruit with (1)2–7 pyrenes ............................................................................................33

32. Stems not resinous; inflorescence paniculate or racemose; calyx extremely reduced, lobes

broadly to narrowly triangular; corolla campanulate or broadly infundibuliform, white,

cream-white to pale yellow throughout, the tube narrowly reduplicate; fruits globose to

oblong-cylindrical, sometimes laterally compressed ................................................ Chiococca

32. Stems often slightly resinous; inflorescence cymose; calyx tube narrowly obovoid, shallowly

lobed; corolla purple-brown outside and deep yellow inside, the tube not reduplicate; fruit

subglobose to obovoid (endemic to eastern Cuba) .............................................. Ramonadoxa

33. Flowers (4–)5–7(–9)-merous, bisexual or unisexual and bisexual on same individual

(polygamous-dioecious); corolla lobes imbricate (quincuncial), with entire, undulate or

fringed margin; ovary (2–)4–7(–9)-locular; fruit globose, ovoid or oblong-ovoid, with (2–

)4–7(–9) pyrenes ....................................................................................................... Guettarda

33. Flowers 4(5)-merous, bisexual; corolla lobes valvate or valvate-induplicate, with entire

margin; ovary (1)2–3-locular; fruit fusiform, ellipsoid, oblong-ellipsoid or cylindrical, with

(1) 2–3 pyrenes .......................................................................................................................34

34. Flowers 4(5)-merous, heterostylous or possibly homostylous; corolla lobes valvate or

valvate-induplicate, ovate, lanceolate, oblong, oblong-ovate, linear-lanceolate or linear,

round, ovate or acute at apex, sometimes short- or long-acuminate at apex, sometimes with a

rhomboidal or triangular internal appendix at apex ...................................................Chomelia

34. Flowers 4-merous, heterostylous; corolla lobes valvate, narrowly ovate, lanceolate, oblong or

ligulate, round or acute at apex, without an internal apical appendix ........................ Malanea
35. Twining or scrambling vines, rarely a shrub with scrambling branches; plants monomorphic,

without adventitious roots; stipules persistent, free at base; inflorescences axillary, 1 or 2 per

node, compact-verticillate, fascicled, glomerulate, capitate or laxly paniculate; flowers free

from each other; ovary 4–5(–7)-locular; fruits free from each other ............................ Sabicea

35. Terrestrial or epiphytic vines; plants with distinct juvenile and adult stages (juvenile with

adventitious roots); stipules caducous, sheathing at base; inflorescences axillary or terminal,

2 or 3 per node, a pedunculate capitulum subtended by a discoid involucre; flowers

congested, free or basally connate; ovary 2(4)-locular, basally connate with those of the

same head; fruits usually basally connate forming a head ....................................... Schradera

Identification of genera based on vegetative characters

Stipules (interpetiolar).

A) Entire. Triangular, ovate, elliptic, obovate, oblong, ligulate, linear: Appunia,

Arachnothryx, Chiococca, Cosmibuena, Coutarea, Didymaea, Eumachia, Guettarda,

Hillia, Hintonia, Machaonia, Malanea, Manettia, Morinda, Paederia, Randia,

Rogiera, Rondeletia, Sabicea, Salzmannia, Schradera, and Uncaria;

B) Fimbriate. With (1)3–7(–9) setae: Denscantia, Emmeorhiza, and Hexasepalum.

C) Withering. Withering on the stem (no abscission layer). Stipules persistent, deltoid to

narrowly triangular, and remaining on stem as a series of triangular units on either

side of the nodes: Didymaea.

D) Indurate. Stipules become hardened when old, with the distal portion falling off by

fragmentation; they are dentate or with 1–2 short acumens or aristae, these

sometimes with 1–2 caducous glandular appendages: Eumachia.

 E) Resinous. Stipules with abundant resinous exudate, forming a globose structure at the

distal nodes, and covering stems: Salzmannia.

F) Foliaceous. Stipules are leaf-like, giving the impression of whorled leaves, 4–6(–8) per

node: Galium.

Thorns. Present in a few genera. Some species may develop thorns or not, which may be variable

on different portions of the stem. In Randia, thorns are straight stout or acicular and 2–4

per node. In Chomelia and Guettarda, they are straight or slightly curved, stout or acicular,

simple (without vestigial nodes) and two per node. In Uncaria, thorns are hook-shaped,

slightly or strongly recurved and one or two per node.

Involucral bract. The pedunculate flowering heads subtended by an involucral, discoid or cup-

shaped bract that sometimes splits irregularly with age: Schradera.

Heterophylly. Juvenile shoots with small leaves, thin flattened stems, climbing by means of

adventitious roots and adult shoots with large leaves, with thick, woody stems, that hang

down from the phorophyte: Hillia and Schradera.

APPUNIA J. D. Hooker in Bentham & J. D. Hooker, Gen. Pl. 2: 120. 1873.

 Weakly branching shrubs or small trees, sometimes scrambling lianas, unarmed. Stipules

persistent or

caducous, free or

briefly connate at

base, broadly to

narrowly triangular,

acuminate or bifid at

tip. Leaves opposite

(ternate in non-

climbing species),
Appunia guatemalensis, photo by Jan Meerman.
petiolate; blades

elliptic, ovate, lanceolate to oblanceolate, membranaceous, chartaceous to subcoriaceous;

domatia absent. Inflorescences leaf-opposed, with 1 or 3 leaves opposed to the peduncles (due to

extreme reduction of internode), or occasionally terminal, pedunculate, sometimes with

secondary peduncles, capitate, with 2–10(–15) flowers per head; ovaries of individual flowers

congested but not fused. Flowers 4–5(6)-merous, heterostylous. Calyx persistent, extremely

reduced, tubular or funnelform, truncate, undulate or with small, broadly triangular lobes; corolla

hypocrateriform or narrowly infundibuliform, pale green to white, tube adaxially glabrous or

pilose near the base; lobes 4–5(6), valvate, reflexed, linear-lanceolate, margins entire or ciliate,

dorsally corniculate; stamens included, inserted near middle of tube or near mouth of corolla,

glabrous, anthers oblong; ovary 2-locular, with 2 ovules per locule, sometimes appearing as 4-

locular due to thick placenta within each locule; style included or exserted just beyond corolla

mouth, obtuse to capitate, or shortly bifid. Infructescence a head of a few to many fruits
congested but not fused. Fruit drupaceous, globose to oblong-ovoid, passing from red to maroon

or purple-black at maturity; pyrenes 4 or fewer per fruit (due to the abortion of 1–2 ovules),

vertical, woody, ovoid to reniform, angled at the ventral edge.

Distinctive features: Recognized by the leaf-opposed inflorescences. Inflorescences often with

2 or 3 peduncles per node, capitate, with individual flowers congested but not fused; ovary 2-

locular, commonly with 2 ovules per locule. Fruits fleshy drupes, congested but not fused, with

(2–3) 4 pyrenes.

Distribution: About 12 species distributed from Central America to northern South America,

with a few species in the Amazon basin; highest diversity in Venezuela and the Guianas. Only A.

megalantha C. M. Taylor & Lorence, from Colombia, Ecuador and northern Peru, is reported as

a liana; wet forests; 320–850 m.

ARACHNOTHRYX Planchon, Fl. Serres Jard. Eur. 5: 442. 1849.

Shrubs or small trees, rarely scrambling shrubs or woody vines, unarmed. Stipules

caducous, free or sheathing at base, oblong-ovate, acute to acuminate. Leaves opposite,

isophyllous or anisophyllous, short- to long-petiolate; blades elliptic to oblanceolate,

chartaceous, domatia absent; tertiary venation clathrate. Inflorescence terminal, frondose or not,

thyrsoid, pauci- or multi-florous. Flowers 4(5)-merous, heterostylous, fragrant. Calyx lobes

lanceolate, acute; corolla actinomorphic, hypocrateriform, pink, glabrous and lacking a fleshy

ring at mouth, tube arachnoid-tomentose or glabrous outside, lobes imbricate, round, margin

eroded or crisped; stamens included (in long-styled flowers) or partially exserted (in short-styled

flowers), filaments inserted at median or distal portion of the tube, anthers short, narrowly

oblong, basifixed; style exserted just beyond the corolla mouth (in long-styled flowers) or
included (in short-styled flowers), terete, glabrous, with 2 oblong lobes; ovary 2-locular;

placentation axile, ovules many per locule. Fruit a globose, crustaceous capsule dehiscing

basipetally. Seeds horizontal, rhomboid or multi-angular, margins undulate, fringed, or narrowly

winged; testa foveolate, rugulose.

Distinctive features: Similar to Rondeletia and Rogiera in being woody and by having capsular

fruits but distinguished by the glabrous corolla mouth lacking a fleshy ring or a ring of hairs (vs.

thick fleshy ring in Rondeletia or with a ring of yellow or white hairs in Rogiera).

Distribution: About 80 species, of which only A. darienensis (Standl.) Borhidi, from Panama

(Darien) and Colombia (Chocó), has a variable habit that includes scrambling shrubs or lianas;

moist forests; 20–274 m.

CHIOCOCCA P. Browne, Civ. Nat. Hist. Jamaica 164. 1756.

Scrambling or twining lianas, shrubs or treelets with scrambling branches; unarmed. Lianas

with twisted, deeply furrowed

stems, reaching 10–15 m in

length and up to 6 cm in diam.;

commonly with short,

opposite, plagiotropic

branches; cross sections

(known only for C. alba (L.)

Hitchc.) with neoformed

Chiococca alba, photo by P. Acevedo.
vascular cylinders in the cortex

(Figures 4F; 217E). Stipules persistent, connate at base, deltoid or narrowly triangular. Leaves
 Chiococca alba. A. Flowering branch. B. Cross section of mature stem showing neoformations. C.
Inflorescence. D. Flower, longitudinal section & lateral view. E. Stamens. F. Distal portion of style
showing stigmatic area. G. Infructescence. H. Fruit, lateral view and cross section. Drawing courtesy
of Bobbi Angell.

opposite, subsessile or short-petiolate; domatia absent. Inflorescences axillary, paniculate or

racemose, sparsely branched. Flowers 4–5(–6)-merous, homostylous. Calyx persistent, extremely

reduced, lobes broadly to narrowly triangular; corolla < 1 cm long, campanulate or broadly

infundibuliform, white, cream-white, greenish white or yellow; tube narrowly reduplicate,

glabrous or puberulous outside, glabrous or pubescent inside, lobes valvate or narrowly

imbricate, broadly to narrowly triangular; stamens included or partially exserted, filaments

inserted at base of corolla, basally connate forming a short tube, anthers elliptic to linear; ovary

2-locular, with 1 pendulous ovule per locule, style exserted just beyond corolla mouth, glabrous,
capitate. Fruit drupaceous, fleshy or spongy, white, globose to oblong-cylindrical, terete or

laterally compressed, with (1)2 woody, 1-seeded pyrenes. Seeds pendulous, laterally

compressed.

Distinctive features: Corolla small, campanulate or broadly infundibuliform, white, cream-

white, greenish white or yellow, with tube narrowly reduplicate, and lobes valvate or narrowly

imbricate. Fruits are fleshy or spongy drupes, white, circular to ovate more or less laterally

compressed. The widespread C. alba is easily recognized by the twining branches, deeply

furrowed stems with numerous neoformations of vascular cylinders, and white laterally

compressed drupes.

Distribution: A neotropical genus with ~20 species, seven of which are reported as climbers;

distributed in United States (Florida), Mexico through Central America, the West Indies, and

South America in Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, Brazil, Paraguay

and northern Argentina; dry to moist forests; 0–2,400 m.

CHOMELIA Jacquin, Enum. Syst. Pl. 1, 12. 1760 (nom. cons.).

 Shrubs or trees, sometimes with leaning branches, a few species growing as scrambling

vines or lianas up to 6 m long; branches short and opposite; unarmed or with axillary pairs of

straight, acicular thorns,

simple, or compound

(brachyblasts, sometimes

with vestigial nodes and

vestigial leaves). Stipules

persistent or caducous, free

or connate at base, broadly to

narrowly triangular. Leaves

Chomelia psilocarpa, photo by A. Hernández.
opposite, short- to long-

petiolate; blades chartaceous to subcoriaceous, sometimes with sunken secondary veins; domatia

tufts of dense hairs or absent. Inflorescence axillary or terminal on axillary short shoots of 3–5

nodes, cymose, pauci- or multi-florous, sometimes with scorpioid branches, or dichasial cymes,

fasciculate or uniflorous. Flowers 4(5)-merous, heterostylous or possibly homostylous;

hypanthium ovoid, narrowly obovoid, obovoid, oblate or turbinate. Calyx persistent, short- to

long-tubular, lobes ovate, oblong, narrowly triangular or linear; corolla narrowly

infundibuliform, white to cream-white or yellowish white, glabrous, the tube sometimes strigose

or sericeous outside, lobes valvate or valvate-induplicate, acute (acuminate) at apex, with a

rhomboidal or triangular internal appendix at apex; stamens included or partially exserted,

inserted near the corolla mouth, filaments short, equal, glabrous, anthers subsessile, elliptic

narrowly elliptic, oblong to narrowly oblong, sometimes with pointed extension or tailed at base,

dorsifixed near the base; ovary 2(3)-locular; placentation axile, ovules 1 per locule, apically
inserted, pendulous, style 2(3)-lobed. Fruit drupaceous, fleshy, fusiform or ellipsoid, turning red

to purple to bluish black when mature; pyrenes 2(3), pendulous, woody, dorsoventrally convex.

Seeds cylindrical.

Distinctive features: Similar to Guettarda but differs by the 4(5)-merous flowers,

valvate corolla lobes in bud, style 2(3)-lobed, ovary 2(3)-locular and fruits with (1)2(3) pyrenes

[vs. flowers (4–)5–7(–9)-merous, corolla lobes imbricate-quincuncial, style subcapitate or (2–)4–

7(–9)-lobed, ovary (2–)4–7(–9)-locular and fruits with (2–)4–7(–9) pyrenes in Guettarda].

Distribution: A neotropical genus of ~70 species, five of which sometimes grow as

scrambling vines; from Panama to Bolivia, including the Guianas and the Amazon basin;

lowland, moist forests; 0–1,200 m.

COSMIBUENA Ruiz & Pavón, Fl. Peruv. 3: 2. 1802 (nom. cons.).

Shrub or trees, epiphytic, rarely terrestrial, rarely climbing with twining or scrambling,

unarmed branches; terminal branchlets

commonly succulent. Stipules caducous

(those subtending flowers somewhat

persistent), connate to sheathing at base,

free portion obovate to oblanceolate.

Leaves opposite, petiolate; blades ovate,

elliptic, obovate, or oblong,

subcoriaceous, coriaceous to succulent;

Cosmibuena grandiflora, photo by C. Galdames. domatia absent. Inflorescences terminal,

cymose, multiflorous or rarely uniflorous. Flowers (4–)5–6(–7)-merous, homostylous, nocturnal.

Calyx persistent or deciduous, membranaceous to coriaceous, truncate or shallowly to deeply

divided; corolla hypocrateriform, white, cream-white, greenish white, turning cream to yellow

after anthesis, tube long and narrow, glabrous (rarely papillose) inside and outside, lobes

imbricate or rarely convolute, elliptic, elliptic-oblong to oblong, or rarely ovate; stamens

included or partially exserted, filaments inserted at distal portion of tube, anthers oblong; ovary

2-locular; ovules many per locule, vertical, imbricate, style exserted just beyond corolla mouth,

glabrous, branches 2, elliptic to ovate. Fruit capsular, septicidal, cylindrical, woody, dark brown

when dry, smooth or with several lenticels. Seeds elliptic, dorsoventrally compressed, with a

narrow rhombic, membranaceous wing, margin entire or irregularly erose.

Distinctive features: Cosmibuena and Hillia are similar in being epiphytic or hemi-epiphytic,

shrubs or trees, with succulent terminal branches, coriaceous or succulent leaves, large flowers,

and large capsules. Cosmibuena differs from Hillia by the stipules shallowly fused at base, calyx

truncate or divided, and seeds with a membranaceous wing; while in Hillia stipules are free,

calyx lobes free at base (tube absent), and seeds with a tuft of long hairs at distal end.

Distribution: A genus of four species, commonly of erect shrubs or trees, sometimes with

scrambling lateral branches; Mexico to Peru, Bolivia and southern Brazil; Cosmibuena

grandiflora (Ruiz & Pav.) Rusby, although commonly a tree, is known to grow in Venezuela

(Tillett 45481) as a liana 30 m long; moist forest, savanna; 0–1,300 m.

COUTAREA Aublet, Hist. Pl. Guiane 1: 314. 1775.

 Shrubs or small to medium-sized (rarely tall) trees, or rarely scrambling (?) lianas;

unarmed; bark fissured; young

branches often lenticellate.

Stipules free at base, broadly

triangular to deltoid, persistent.

Leaves opposite, short-

petiolate; blades ovate to

elliptic, chartaceous to thinly

coriaceous; domatia tufts of

Coutarea hexandra, photo by A. Popovkin. sparse or dense hairs, or absent.

Inflorescences terminal, paniculate, frondose or not, sparsely branched. Flowers (5–)6–7-merous;

flower buds curved. Calyx cup-shaped, persistent, lobes linear; corolla campanulate,

zygomorphic, white, cream-white, pale green, pink, red, violet to purple, tube reduplicate at

edges, glabrous throughout, lobes imbricate, ovate to deltoid, margin entire, rounded at apex;

stamens partially exserted, filaments long, unequal, glabrous throughout or puberulent at basal

portion inserted at base of corolla, anthers linear, dorsifixed near base; ovary 2-locular,

placentation axile, peltate to entire length of septum, ovules many per locule, style exserted just

beyond corolla, glabrous, style 2-branched. Fruit capsular, dehiscing loculicidally from apex,

obovate to oblong-ovate in outline, laterally compressed, woody, often lenticellate. Seeds many,

ascendingly imbricate, oblong-ovate in outline; wing concentric, membranous, with entire

margin.

Distinctive features: Characterized by the large, campanulate, zygomorphic corollas, with a

curved tube, reduplicate at edges, lobes imbricate in bud; hypanthium laterally compressed; and
filaments connate at base, inserted at base of corolla tube, and of different lengths. Capsules

laterally compressed, obovate to oblong-ovate in outline and often lenticellate.

Distribution: A genus of two species; Mexico throughout Central America to Peru, Bolivia,

Brazil to northern Argentina. Coutarea hexandra (Jacq.) K. Schum., although commonly a shrub

or a small to large tree, sometimes grows as a shrub with scrambling branches or as a liana;

scrublands to dense forests; 0–1,730 m.

DENSCANTIA E.L. Cabral & Bacigalupo, Darwiniana 39: 353. 2002.

Scrambling herbaceous or subwoody vines; unarmed; stems basally woody. Stipules

persistent, often withering on the stem, sheathing

and connate to the petioles, fimbriate, with 1 or 3–

8 setae, each seta with an apical colleter. Leaves

opposite, sometimes seeming whorled by the

presence of axillary short shoots (brachyblasts),

subsessile or short-petiolate; blades elliptic,

obovate, narrowly elliptic to elliptic-lanceolate,

membranaceous, chartaceous or papyraceous.

Inflorescence terminal, thyrsoid, frondose or not,

lateral branches with flower glomerules, or

flowering branches with terminal and axillary

Denscantia cymosa, photo by W. Oliveira flower fascicles or glomerules. Flowers 4-merous,

Fonseca.
homostylous. Calyx persistent, lobes small, narrowly triangular; corolla infundibuliform to

tubular, white, cream-white to pink, tube externally glabrous or puberulent, internally glabrous,
puberulent or pubescent with moniliform hairs, lobes valvate, narrowly ovate, narrowly

triangular or oblong; stamens exserted well beyond the corolla, filaments inserted near the mouth

of the corolla tube or at lobe sinuses, anthers elliptic or oblong; ovary 2-locular, ovules 1 per

locule, centrally inserted, vertical, style exserted well beyond the corolla, capitate-bilobed. Fruit

capsular, ovoid, obovoid, oblong-ovoid, dehiscing septicidally from the apex down to the seed

insertion, both cocci dehiscent, thinly woody or crustaceous. Seeds vertical, plano-convex,

oblong-ovate in outline, ventrally sulcate, margin with a very narrow wing.

Distinctive features: Climbing herbs or subshrubs, with sheathing fimbriate stipules bearing

(1)3–8 setae, often withering on the stem, and a thyrsoid inflorescence with flowers in fascicles

or glomerules. Capsules with both cocci dehiscing septicidally from the apex down to the seed

insertion. Similar to Spermacoce (see discussion under that genus).

Distribution: A genus of five species endemic to the Atlantic coastal forests of Brazil, from

Alagoas to Rio de Janeiro; 50–800 m.

DIDYMAEA J. D. Hooker in Bentham & J. D. Hooker, Gen. Pl. 2: 150. 1873.

 Procumbent herbs or scrambling vines to 4.5 m long; stems terete, unarmed. Stipules

persistent, free at base,

deltoid to narrowly

triangular, withering on

the stem as a series of

triangular units on either

side of the nodes. Leaves

opposite; subsessile, or

short- to long-petiolate;

Didymaea mexicana, photo by M. Jiménez Chimil. blades ovate, broadly

ovate, narrowly ovate,

elliptic, broadly elliptic, narrowly lanceolate, lanceolate, cordate, or orbicular; membranaceous

or chartaceous; domatia absent. Inflorescence axillary, cymose, pauciflorous, or uniflorous;

bracts small. Flowers 4-merous, homostylous. Calyx tube extremely reduced, persistent, truncate

or undulate; corolla campanulate or rotate, actinomorphic, greenish brown, yellowish white,

purplish, violet to purple, tube glabrous, lobes valvate, narrowly triangular; stamens partially

exserted or exserted just beyond the corolla, filaments inserted on upper portion of tube or near

the mouth of the corolla, short or long, shorter than the corolla tube, equal, glabrous, anthers

elliptic, round at base and at apex; ovary 2-locular, ovules 1 per locule, basally inserted, style

exserted or partially exserted, glabrous, stigmatic lobes 2, ovate. Fruit drupaceous, fleshy,

didymous or globose, bluish black to purplish black. Pyrenes (1)2, ascending, hemispherical,

woody, 1-seeded. Seeds minute, reniform.

Distinctive features: Scrambling herbs or herbaceous vines to 4.5 m long, with deltoid to

narrowly triangular stipules withering on the stem as a series of triangular units on both sides of

the nodes. Drupes fleshy, bluish black to purplish black.

Distribution: A neotropical genus of eight species of which only D. mexicana Hook. f. grows as

a vine that reaches more than 2 m in length; northeastern Mexico to Panama; moist tickets, cloud

forests and rocky slopes; 1,500–2,300 m.

EMMEORHIZA Pohl ex Endlicher, Gen. Pl. 565. 1838.

Twining vines, often with short plagiotropic branches; stems woody at the base, unarmed

5–10 m long and ~1 cm

in diam.; cross section

deeply 4-lobed and often

fistulose (Figure 217F).

Stipules persistent, often

withering on stem,

sheathing, connate to

petioles, fimbriate, with

3–9 setae, each seta with

Emmeorhiza umbellata, photo by P. Acevedo.
an apical colleter.

Leaves opposite or ternate, often seemingly whorled by the presence of axillary short shoots

(brachyblasts) with fasciculate leaves, sessile to short-petiolate; blades ovate, narrowly ovate,

broadly to narrowly elliptic, rhombic to oblong, chartaceous to papyraceous; domatia absent.

Inflorescences terminal, paniculate, with secondary branches terminating on many-flowered

umbels. Flowers 4-merous, homostylous. Calyx persistent, tube extremely reduced, lobes minute

or small, narrowly triangular, lanceolate, narrowly lanceolate to linear; corolla broadly

infundibuliform, white or cream-white, tube externally glabrous, internally pubescent or villous

at medial-basal portion, lobes valvate, narrowly ovate; stamens exserted well beyond the corolla,

filaments inserted near the mouth of corolla tube or at lobes sinuses, anthers elliptic to narrowly

elliptic; ovary 2-locular, ovules 1 per locule, centrally inserted, style exserted well beyond the

corolla mouth, branches 2, oblong. Fruit capsular, narrowly obconical to turbinate, dehiscing

septicidally from the apex down to the seed insertion, valves indehiscent. Seeds 2 per capsule,

vertical, laterally compressed, narrowly oblong in outline; wings bipolar, somewhat acute at both

ends, margin entire.

Distinctive features: Twining, subwoody vine; stems deeply lobed, fistulose; inflorescences

paniculate with flowers clustered in umbels. Fruit capsular, dehiscing septicidally from the apex

down to the seed insertion, with indehiscent valves.

Distribution: A genus of a single species, E. umbellata (Spreng.) K. Schum.; Colombia,

Venezuela, Trinidad, the Guianas south to Brazil, Bolivia and northern Argentina; moist forests;

30–2,500 m.

EUMACHIA A.P. de Candolle, Prodr. 4: 478. 1830.

 Subshrubs, shrubs or trees, rarely creeping or climbing herbs, or scrambling vines;

unarmed; stem in E. microdon

(DC.) Delprete & J.H. Kirkbr.

quadrangular to slightly flattened,

2–2.5 m long. Stipules persistent,

becoming indurate, distal portion

falling off by fragmentation, free at

base, entire. Leaves opposite, long-

petiolate; blades ovate, elliptic,

Eumachia microdon, photo by M. Caraballo-Ortiz. obovate, oblong or narrowly

elliptic, chartaceous or papyraceous; domatia absent. Inflorescences terminal, pedunculate,

subcapitate or paniculate with 2–several pairs of secondary branches, few- to many-flowered, with

2–8 bracts, either subtending inflorescence or inserted on inflorescence branches. Flowers 5-

merous, heterostylous. Calyx persistent, minute, cyathiform or cup-shaped, lobed or undulate,

lobes (when present) small, broadly to narrowly triangular; corolla actinomorphic,

hypocrateriform, infundibuliform or campanulate, white or cream-white, tube glabrous outside,

lobes valvate, ovate-triangular, oblong-ovate, ovate to oblong-lanceolate to elliptic-ovate,

appendiculate on the adaxial side; stamens included in long-styled flowers, as long as the corolla

tube or exserted in short-styled flowers, anthers ovoid to ellipsoid; ovary 2-locular, ovules 1 per

locule, basally inserted, erect, style glabrous, 2-branched. Fruit drupaceous, subglobose to

ellipsoid, fleshy, with 2 pyrenes, turning from orange-red to red to purplish red to bluish or

purplish black at maturity; pyrenes woody, plano-convex, ovate to elliptic in outline, with 2
marginal germination slits extending to the middle and sometimes also with several shorter

dorsal slits, dorsally multicostate, ventrally flat.

Distinctive features: Scrambling shrubs with persistent, indurate stipules with distal portion

falling off by fragmentation; drupes subglobose to ellipsoid, turning from orange-red to red to

purplish red to bluish or purplish black at maturity; pyrenes dorsally multicostate, ventrally flat,

with 2 marginal germination slits extending to the middle, and sometimes also with several

shorter dorsal slits.

Distribution: A pantropical genus of ~90 species, with 27–30 in the Neotropics, of which E.

microdon is the only species that grows as a scrambling shrub in addition to being an erect shrub;

Mexico to Panama, Greater and Lesser Antilles, Guyana, Venezuela, Colombia, Ecuador, and

Peru; understory of evergreen or seasonal forests; 50–1,050 m.

GALIUM Linnaeus, Sp. Pl. 105. 1753.

Erect, sprawling or scrambling herbs (to 4 m long), annual or perennial herbs, or

subshrubs; unarmed.

Stipules persistent,

commonly of same

shape and size of leaves.

Leaves whorled,

commonly 4 (rarely 6–8)

per node (including the 2

leaf-like stipules),
Galium hypocarpium, photo by B. Hammel. subsessile or sessile;
blades ovate, narrowly ovate, elliptic, narrowly elliptic, oblong, lanceolate or linear,

membranaceous or chartaceous; domatia absent. Inflorescences axillary or terminal, paniculate

or cymose, frondose or not, 1-, few- or many-flowered. Flowers (3–)4–5-merous, bisexual,

unisexual or functionally unisexual (male and female flowers on separate individuals), or

unisexual and bisexual on same individual (polygamous-monoecious). Calyx lobes free,

caducous, ovate, lanceolate to narrowly ovate; corolla campanulate, urceolate or rotate, white,

cream-white, greenish white, yellow, yellowish white, pink, red or maroon, tube (when present)

glabrous, internally glabrous, pubescent or hispid, lobes valvate, ovate, margin entire; stamens

inserted at sinuses of corolla lobes, filaments short, glabrous, anthers elliptic or oblong; ovary 2-

locular, placenta axile, reduced, ovules 1 per locule, centrally inserted, vertical; styles 2,

sometimes united at base, capitate, glabrous. Fruits didymous, baccate, fleshy, white, orange, red

or blackish red, or dry, schizocarpic, splitting into 2 dehiscent cocci. Seeds 2, vertical, dorsally

convex.

Distinctive features: Scrambling herbs to 4 m long; leaves 4 (6–8) per node, including the 2

leaf-like stipules. Fruits baccate, fleshy, didymous, orange.

Distribution: A cosmopolitan genus of ~400 species occurring in tropical and temperate

environments; in the New World there are ~160 species ranging from Alaska to southern

Argentina and Chile, 92 of which are found in the Neotropics, with only five species consistently

reported as vines that attain 2 or more m in length; forest understory, forest margins, high

altitude mountain vegetation, secondary vegetation; 50–3,400 m.

GUETTARDA Linnaeus, Sp. Pl. 991. 1753.

 Shrubs, small to tall canopy trees, or scrambling lianas. Axillary thorns when present,

straight or slightly

curved, stout, with

vestigial nodes and

sometimes with reduced

leaves. Stipules

persistent or readily

caducous, free at base,

deltoid or narrowly
Guettarda comata, from Schunke-Vigo 6636 (US).
triangular. Leaves

opposite or whorled, 3–4 per node, short- to long-petiolate; blades ovate, elliptic or oblong,

chartaceous, papyraceous or subcoriaceous; domatia of hairy-pockets, or absent. Inflorescences

axillary, sessile or pedunculate, cymose or dichasial, with 2–7 scorpioid branches, or rarely

subcapitate, frondose or not, few- to many-flowered. Flowers (4–)5–7(–9)-merous, bisexual,

sometimes unisexual flowers also present (polygamous-dioecious), sessile or pedicellate. Calyx

persistent, cup-shaped, lobes small, broadly triangular, deltoid or ovate; corolla hypocrateriform

or broadly infundibuliform, white, cream-white, yellow, bluish white, pale blue, violet to purple,

tube externally pubescent, internally glabrous or pubescent, with or without a pubescent ring at

base, lobes (4–)5–7(–9), imbricate (quincuncial), ovate, round or oblong, margin entire, undulate

or fringed; stamens included or partially exserted, filaments inserted near mouth of corolla tube,

anthers subsessile, oblong or linear; ovary (2–)4–7(–9)-locular, ovules 1 per locule, pendulous,

inserted near the roof of the locule, style included, terete, glabrous or antrorsely puberulent,

capitate or with (2–)4–7(–9) short lobes, these ovate, oblong to linear. Fruit drupaceous, fleshy,
globose, ovoid or oblong-ovoid, white, red, purple to black. Pyrenes (2–)4–7(–9), woody, plano-

convex. Seeds pendulous, compressed, oblong.

Distinctive features: Scrambling lianas, sometimes armed with axillary thorns; flowers (4–) 5–

7(–9)-merous, bisexual or unisexual; calyx persistent, cup-shaped with broadly triangular,

deltoid or ovate lobes; drupes fleshy, white, red, purple to black, and with (2–)4–7(–9) woody

pyrenes inserted near the apex of the locule.

Distribution: A pantropical genus of ~120 species, 80–90 of which occur in the Neotropics, of

these, only G. comata Standl. (Colombia), and G. tikalana Lundell (Mexico and Belize) are

reported as lianas; primary and secondary forests; 100–800 m.

HEXASEPALUM Bartling ex A.P. de Candolle, Prodr. 4: 561. 1830.

Annual or perennial, erect or decumbent herbs, sometimes scrambling subwoody vines;

unarmed. Stems sub-quadrangular, reaching 8–10 m

in length and ~2 cm diam. in H. sarmentosum (Sw.)

J.H. Kirkbr. & Delprete; cross section with deep

phloem wedges in Hexasepalum sarmentosum

(Figure 217C). Stipules persistent, often withering

on stem, sheathing and connate to petioles, fimbriate,

with 3–13 setae, each seta commonly with an apical

colleter. Leaves opposite, sessile, subsessile to short-

petiolate; blades ovate, elliptic, oblong, narrowly

oblong to lanceolate, chartaceous, papyraceous or

Hexasepalum sarmentosum, photo by P.
Acevedo. subcoriaceous; domatia absent. Inflorescences
axillary, glomerules, 1- to few-flowered. Flowers 4-merous, homostylous. Calyx persistent, tube

extremely reduced, lobes ovate, oblong, lanceolate, narrowly lanceolate to linear; corolla

hypocrateriform or narrowly infundibuliform, white, cream-white, pinkish white, pink, bluish

white, pale blue to lilac, tube externally glabrous, internally pubescent, usually with moniliform

hairs, lobes valvate, ovate, oblong-ovate to narrowly ovate; stamens partially exserted or shortly

exserted, filaments inserted near mouth of corolla tube or at lobes sinuses, glabrous, anthers

elliptic, narrowly elliptic or short-oblong; ovary 2-locular, ovules basally or medially inserted, 1

per locule, style exserted just beyond or well beyond corolla, glabrous, capitate or with 2 short,

elliptic, oblong or narrowly oblong branches. Fruit schizocarpic, obovoid, thinly woody to

coriaceous, breaking up into 2 mericarps; mericarps indehiscent, plano-convex, 1-seeded. Seeds

vertical, obovate or subellipsoid, plano-convex, dorsal side smooth or reticulate-foveolate,

ventral side with a narrowly ellipsoid depression and two apical extensions, Y-shaped.

Distinctive features: Scrambling vines with fimbriate stipules bearing 3–13 setae, and

schizocarpic fruits that split into 2, indehiscent, 1-seeded mericarps.

Distribution: A genus of ~12 species naturally found in the Americas and Africa, of which H.

gardneri (K. Schum.) J.H. Kirkbr. & Delprete (from Brazil), H. sarmentosum (from Tropical

America and Tropical Africa), and H. scandens (Sw.) J.H. Kirkbr. & Delprete (from Hispaniola)

are scrambling vines; thickets, pastures, forest edges, and secondary growths, in moist areas; 0–

1,600 m.

HILLIA Jacquin, Enum. Syst. Pl. 3, 18. 1760.

Erect or scrambling epiphytic shrubs or small trees, sometimes with pendulous branches,

glabrous throughout; unarmed. Stipules readily caducous, free at base, ovate, oblong,
 oblanceolate or ligulate. Leaves opposite, isophyllous

or anisophyllous, subsessile to short-petiolate; blades

ovate, elliptic, oblanceolate to lanceolate,

subcoriaceous to coriaceous, sub-succulent to

succulent. Inflorescences terminal, cymose, 3-

flowered, or uniflorous. Flowers 4–10-merous. Calyx

lobes free, round, narrowly triangular or lanceolate.

Corolla tubular, hypocrateriform or broadly

infundibuliform, often inflated at the middle, white,

cream-white, yellow, or pale to bright green, orange,

pink, red or violet to purple; tube glabrous throughout,

Hillia parasitica, photo by P. Acevedo. lobes right-contorted, round, elliptic, broadly to

narrowly triangular or linear, reflexed at anthesis; stamens included, partially exserted or

exserted just beyond the corolla mouth, filaments inserted at middle or near the corolla mouth,

anthers subsessile, narrowly elliptic or oblong; ovary 2-locular, ovules many per locule, style

included or as long as corolla tube, branches 2, ovate or long-linear. Fruit capsular, septicidal,

oblong to narrowly oblong, crowned by the permanent calyx and the disk, woody, valves

secondarily splitting at apex. Seeds many, imbricate, fusiform, with a tuft of long hairs (exotestal

extensions) at distal end.

Distinctive features: Hillia and Cosmibuena are similar in being epiphytic or hemi-epiphytic

(rarely terrestrial), vines with succulent distal branches, coriaceous or succulent leaves, large

flowers, and large, septicidal capsules. See discussion under Cosmibuena.

Distribution: A neotropical genus of ~24 species, seven of which are often described as

climbing shrubs or lianas; Mexico throughout Central America to Colombia, Bolivia, and Brazil,

including the Greater and Lesser Antilles; wet or cloud forests; 600–2,750 m.

HINTONIA Bullock, Hooker’s Icon. Pl. 33(4): tab. 3295. 1935.

Shrubs, or small to medium-sized trees, rarely scrambling woody vines > 5 m long, with

short, opposite branches; unarmed.

Stipules persistent, or sub-caducous,

free or connate at base, broadly

triangular. Leaves opposite, short- to

long-petiolate; blades ovate, elliptic,

oblong, or lanceolate, chartaceous or

papyraceous; domatia tufts of sparse

or dense hairs, or absent.

Hintonia octomera, photo by W.J. Hayden.
Inflorescence axillary, cymose,

pauciflorous or uniflorous. Flowers 6- or 8-merous. Calyx lobes free or shallowly connate at

base, caducous, narrowly lanceolate or linear; corolla actinomorphic, campanulate, narrowly

campanulate, or broadly infundibuliform, white, cream-white, yellowish white, pink, purplish,

violet to purple; tube reduplicate, glabrous or basally pubescent outside, lobes imbricate or left-

contorted, broadly triangular, margin entire, round or obtuse at apex; stamens partially exserted,

filaments inserted at base of the corolla tube, basally connate into a minute tube, slender, long,

equal, glabrous, anthers linear, basifixed; ovary 2-locular, placentation axile, with many ovules

per locule, perpendicular to the central septum, style exserted just beyond the corolla, terete,
glabrous, unlobed, with stigmatic lines along style. Fruit capsular, septicidal, dehiscing

basipetally, ellipsoid or subglobose, woody, sometimes lenticellate, the valves sometimes

secondarily splitting at apex. Seeds imbricate, horizontal or basipetal, large, dorsoventrally

compressed, elliptic to narrowly elliptic in outline, with concentric wing along the entire margin.

Distinctive features: Scrambling woody vines > 5 m long, flowers 5–7 cm long, 6- or 8-merous,

with campanulate or broadly infundibuliform corolla, filaments basally connate, inserted at base

of the corolla, anthers linear, basifixed. Similar to Coutarea but distinguished by the

actinomorphic flowers with straight corolla, the stamens of equal length, and by the terete,

septicidal capsules (vs. zygomorphic, curved corollas with unequal stamens and laterally

compressed, loculicidal capsules).

Distribution: A genus of four species; Mexico, Guatemala, El Salvador and Costa Rica of which

Hintonia octomera (Hemsl.) Bullock (from SE Mexico and Guatemala) is often reported as a

liana or a climbing shrub; primary and secondary forests, seasonal forests, and rarely on coastal

dunes; 5–100 m.

MACHAONIA Bonpland in Humboldt & Bonpland, Pl. Aequin. 1: 101, pl. 29. 1806.
 Shrubs or small trees, sometimes with scandent lateral branches, and rarely subwoody

scrambling shrubs or vines, reaching 5 m in length; young stems often lenticellate. Thorns when

present, 2 per node, stout,

straight, simple or dendroid,

often with vestigial nodes, and

sometimes with reduced

leaves. Stipules persistent, free

at base, broadly triangular.

Leaves opposite, but often

seemingly fasciculate due to

the extreme reduction of nodes

Machaonia martinicensis, photo by Reinaldo Aguilar.
in lateral shoots (brachyblasts),

subsessile or short-petiolate; blades ovate, elliptic to very narrowly elliptic or obovate,

chartaceous or papyraceous; domatia tufts of sparse hairs, or absent. Inflorescence terminal,

frondose or not, paniculate or cymose, densely or sparsely branched, few- to many-flowered.

Flowers 4–5-merous, heterostylous. Calyx persistent, tube extremely reduced, rotate or cup-

shaped, lobes small, elliptic, ligulate, oblong, ovate; corolla < 1 cm long, broadly

infundibuliform, white, cream-white or yellowish white, tube pubescent at mouth but not

forming a ring, lobes imbricate, ovate or round; stamens well exserted in short-styled flowers or

exserted among the corolla lobes in the long-styled flowers; ovary 2–3-locular, ovules 1 per

locule, inserted at top of the locule, pendulous, style branches 2, ovate or elliptic. Fruit

schizocarpic, narrowly turbinate, laterally compressed, chartaceous or thinly woody, splitting

into 2 pendulous, indehiscent, 1-seeded cocci that remain attached to a persistent septum. Seeds

narrowly oblong-cylindrical.

Distinctive features: Scrambling shrubs, sometimes with axillary, straight, simple thorns; fruits

small, schizocarpic, splitting into 2, indehiscent, pendulous, 1-seeded cocci that remain attached

to a persistent septum.

Distribution: A neotropical genus of ~20 species ranging from Mexico throughout Central

America to South America and the West Indies. Machaonia martineziorum Lorence (armed

species ranging from southern Mexico and Guatemala) and M. martinicensis (DC.) Standl.

(unarmed species ranging from Nicaragua to Colombia, Jamaica and Martinique) are sometimes

reported as scrambling shrubs or lianas; flooded forests and thickets and coastal vegetation; 0–

160 m.

MALANEA Aublet, Hist. Pl. Guiane 1: 106, t. 49. 1775.

Twining, root-climbing or scrambling, subwoody vines or lianas, shrubs, or small trees;

unarmed. Stipules

partly or readily

caducous, free or

connate at base,

narrowly triangular,

narrowly ovate,

obovate, elliptic,

oblong or oblanceolate.
Malanea glabra, photo by Alex Popovkin.

Leaves opposite,
subsessile or short-petiolate; blades broadly ovate to ovate, broadly elliptic to elliptic,

papyraceous or subcoriaceous; domatia absent. Inflorescence axillary, paniculate or spiciform,

with short-fasciculate lateral branches, or cymose with the flowers borne in sessile glomerules or

cymules along the primary, secondary and tertiary axes. Flowers 4-merous, heterostylous. Calyx

persistent, campanulate, lobes ovate or broadly to narrowly triangular; corolla broadly

infundibuliform, white, cream-white, yellowish white to orange; tube internally pubescent, not

forming a ring, lobes 4, valvate, narrowly ovate, lanceolate, oblong or ligulate, round or acute at

apex; stamens well exserted above the corolla lobes in short-styled flowers or included or

partially exserted among the corolla lobes in long-styled flowers; ovary (1) 2–3-locular, ovules 1

per locule, inserted at top of septum, pendulous, style 2–3-branched. Fruit drupaceous, fleshy,

oblong-ellipsoid, fusiform or cylindrical, red, purple or black at full maturity; pyrenes (1) 2–3,

plano-convex, dorsal side 3–4-costate, ventral side flat. Seed 1 per pyrene, pendulous,

cylindrical.

Distinctive features: Twining, root-climbing or scrambling vines or lianas with 1–3-locular

ovaries with a pendulous ovule per locule; pyrenes dorsally 3–4 costate and ventrally flat.

Similar to Guettarda in having drupaceous fruits with pendulous pyrenes but differs by the 4-

lobed corollas, valvate in bud, with entire margin, and the (1) 2–3-locular ovaries and fruits [vs.

(4–)5–7(–9)-lobed corollas, which are valvate, imbricate or left-contorted in bud, with entire,

undulate or fringed margins, and the 2–7-locular ovaries and fruits]. In addition, climbing

Guettarda are scramblers, never twiners or root-climbers.

Distribution: A neotropical genus of ~33 species, 29 of which are reported as climbers,

however, several species are erect shrubs during early stages and only become climbers as they

grow old; known from Guatemala, Belize, and from Costa Rica to SE Brazil and the Lesser
Antilles; seasonally dry forests, gallery forests, wet forests, and secondary vegetation; 0–1,400

m.

MANETTIA Mutis ex Linnaeus, Mant. 2: 553, 558. 1771 (nom. cons.).

Twining herbaceous or woody vines to 10 m long, rarely hemiepiphytic, root-climbing

lianas or very rarely erect

subshrubs. Stipules

persistent, free at base,

broadly triangular to deltoid.

Leaves opposite, short- to

long-petiolate; blades ovate,

elliptic, oblong, lanceolate or

almost linear, chartaceous or

Manettia cordifolia, photo by P. Acevedo.
papyraceous; domatia tufts

of sparse hairs, or absent. Inflorescence axillary, frondose or not, thyrsoid, dichasial or

fasciculate, rarely uniflorous. Flowers commonly heterostylous. Calyx persistent, tube extremely

reduced or cup-shaped, truncate or with small to foliose, lobes 4(5, or 8) narrowly triangular,

lanceolate, ovate or linear; corolla tubular, urceolate, hypocrateriform or narrowly

infundibuliform, white, cream-white, yellow, pink, red, bluish white to pale blue, or bicolorous

(tube red and lobes yellowish or greenish), tube glabrous, puberulent or pubescent, lobes 4(5),

valvate, deltoid, narrowly triangular or elliptic; stamens included or partially exserted in long-

styled flowers or exserted just beyond the corolla mouth in short-styled flowers, filaments

inserted on distal half of the corolla tube, anthers narrowly elliptic or oblong; ovary 2-locular,
 Manettia reclinata. A. Flowering branch. B. Node showing interpetiolar stipules. Flower bud with
nectary, frontal, lateral, & dorsal views & longitudinal section of nectary. C. Flower bud. D. Flower,
longitudinal section & top view. E. Fruiting branch. F. Partly dehisced fruit. G. Seed. Drawing
courtesy of Bobbi Angell.

placenta peltate to the entire length of the septum, ovules many per locule, style included in

short-styled flowers or exserted just beyond the corolla in long-styled flowers, glabrous,
branches 2, oblong or ligulate. Fruit capsular, dehiscing septicidally from apex, oblong, ovoid to

subglobose, often laterally compressed, crustaceous or thinly woody, the valves sometimes

secondarily splitting at apex. Seeds many, membranaceous, round, broadly to narrowly elliptic in

outline, with a concentric, narrow, membranaceous wing, with entire or irregular margin.

Distinctive features: Twining or less often root-climbing vines or lianas, with slender stems and

small to large corollas that are tubular, urceolate, hypocrateriform or narrowly infundibuliform,

white, cream-white, yellow, pink, red, bluish white to pale blue, or bicolorous; capsules

septicidal, oblong, ovoid to subglobose, often laterally compressed.

Distribution: A genus of 90–100 species most of which are reported as climbers; Mexico,

throughout Central America, Greater and Lesser Antilles to Colombia, Bolivia, Brazil, and

northern Argentina; primary and secondary forests, seasonal forests, Andean cloud forests; 200–

3,000 m.

MORINDA Linnaeus, Sp. Pl. 176. 1753.

Shrubs, trees or less often twining or scrambling, subwoody vines, 3–4 m in length;

unarmed. Stipules caducous or persistent, free or connate at base or sheathing, oblong, triangular,

ligulate, spatulate or bifid. Leaves opposite or ternate, isophyllous or anisophyllous, short- to

long-petiolate; blades ovate, broadly elliptic, oblong, oblanceolate, chartaceous or stiffly

chartaceous; domatia tufts of sparse or dense hairs, or absent. Inflorescence axillary or distal,

solitary or fasciculate, pedunculate, capitulum of many fused flowers. Flowers homostylous or

distylous. Calyx caducous, tube extremely reduced or short tubular, truncate, undulate or lobed,
 lobes when present,

4–7, broadly

triangular; corolla

hypocrateriform or

narrowly

infundibuliform,

white, cream-white,

or rarely pale blue;

tube externally
Morinda royoc, photo by P. Acevedo.
glabrous or

puberulous , glabrous throughout or puberulent inside, lobes 4–7, valvate, lanceolate or oblong,

with narrow dorsal extension at apex (usually caducous); stamens 4–7, included, partially

exserted, or exserted just beyond the corolla mouth, filaments attached at the middle of the

corolla tube, long, anthers narrowly oblong or elongate, with acute extensions at apex; ovary 2-

locular, with 2 ovules per locule, or partly 4-locular, with 1 ovule per locule, ovules basally

inserted, ascending, style included or exserted just beyond the corolla, capitate, bilobed or 4-

lobed, lobes ovate, oblong or linear. Fruits multiple, fleshy, oblong-globose, ellipsoid to ovoid,

grayish white to yellowish white at maturity; single fruits drupaceous, fleshy; pyrenes commonly

4 per fruit, or 2–3 per fruit by the abortion of 1–2 ovules, oblong or narrowly ovoid. Seeds

vertical, ovoid to obovoid or reniform.

Distinctive features: Twining vines with multiple, fleshy fruits, formed by the fusion of many

ovaries.
Distribution: A genus of 60–80 species worldwide, 20 of which are present in the Neotropics,

with M. royoc L. as the only species of vine; distributed in SE United States (Florida), southern

Mexico to NW South America (Venezuela, Colombia, Ecuador), Cuba, Jamaica, Hispaniola and

the Bahamas; wet forests, coastal scrubs, dry and semi deciduous forests and disturbed

vegetation; 0–400 m.

NOTOPLEURA (Bentham) Bremekamp, Recueil Trav. Bot. Néerl. 31: 289. 1934.

Mostly terrestrial, erect

shrubs or trees, rarely

subshrubs, epiphytic herbs,

scrambling vines or root-

climbing vines, unarmed;

stems commonly succulent.

Stipules persistent to caducous,

connate at base or sheathing,

Notopleura pithecobia, photo by Monica Álvarez.

often connate to the petioles,

truncate or bifid, with interpetiolar appendages, these conical, linear, shortly bifid, bilobed,

fimbriate at the apex, with glandular apical projections or laminar and erose or laciniate. Leaves

opposite, short- to long-petiolate; blades ovate, elliptic, obovate, oblong, oblanceolate or

lanceolate, chartaceous, papyraceous, subcoriaceous to succulent in epiphytic species; domatia

absent. Inflorescence pseudo axillary or terminal, paniculate, sparsely or densely branched, or

capitate, sometimes subtended by bracts. Flowers 4–5(6)-merous, heterostylous, rarely

homostylous. Calyx persistent, tube extremely reduced or cup-shaped, lobes small, broadly to
narrowly triangular; corolla tubular or narrowly infundibuliform, white, cream-white, pinkish

white, yellow, yellowish orange or orange, tube externally and internally glabrous or pubescent;

lobes 4–5(6), valvate, deltoid to narrowly triangular, short-acuminate at apex, sometimes with

pronounced dorsal appendages; stamens 4–5, filaments short or long, included, inserted at basal

or median portion of corolla tube in long-styled flowers, or included or partially exserted and

inserted at medial or distal portion of corolla tube in short-styled flowers, anthers narrowly

elliptic, oblong or narrowly oblong; ovary 2–4(6)-locular, with 1 ovule per locule, basally

inserted, style branches 2–4(6), oblong to linear. Fruit drupaceous, fleshy or spongy, subglobose,

ellipsoid, narrowly ovoid, white, orange, red or black, with 2–4(6) pyrenes. Pyrenes ascending,

dorsoventrally convex, dorsal side smooth or with 1–5 ridges, ventral side sulcate.

Distinctive features: Stems commonly succulent; stipules connate at base or sheathing, truncate

or bifid, and with interpetiolar appendages; inflorescence pseudoaxillary or terminal, paniculate

or capitate, sometimes subtended by bracts; ovary 2–4(6)-locular, with 1 ovule per locule,

basally inserted; fruit drupaceous, fleshy or spongy, white, orange, red or black, with 2–4(6)

pyrenes.

Distribution: A genus of ~73 species, distributed from Mexico to South America, and the West

Indies (Taylor, 2001). Only two species, i.e., N. episcandens C.M. Taylor & Lorence from

Panama and N. pithecobia (Standl.) C.M. Taylor from Costa Rica to Ecuador, in addition to

growing as epiphytic shrubs are reported as scrambling or root-climbing vines that can reach 2–3

m in length; wet forests; 400–1,800 m.

PAEDERIA Linnaeus, Syst. Nat., ed. 12: 135, 189. 1767.

 Twining herbaceous or woody vines, with strong fetid odor; unarmed. Stipules caducous,

free at base,

triangular, rarely

bilobed at apex.

Leaves opposite or

whorled, 3–4 per

node, long-

petiolate; blades

ovate, elliptic or

cordate, chartaceous

or papyraceous;
Paederia brasiliensis, from Venturi 9052 (US).

domatia tufts of

hairs or hairy-pockets, or absent. Inflorescence terminal or terminal on lateral branches,

paniculate or thyrsoid, sparsely branched. Flowers 4–6-merous, bisexual or unisexual and

bisexual on the same individual (polygamous-dioecious), homostylous. Calyx persistent, tube

extremely reduced, with small, broadly to narrowly triangular or linear lobes; corolla tubular,

reddish to purple inside, greenish white outside, tube narrowly infundibuliform, internally

densely pubescent at distal portion, lobes 4–6, valvate, narrowly triangular or narrowly ovate,

margin undulate or fringed; stamens included, inserted at about the middle of the corolla tube,

anthers subsessile, narrowly oblong; ovary 2-locular, with 1 ovule per locule, basally inserted,

style bifid, divided almost to the base. Fruit schizocarpic, dry, ovate or elliptic in outline,

laterally compressed, crustaceous to chartaceous, with 2 flat mericarps that remain suspended by

thin carpophores; mericarps indehiscent, with seed enclosed in endocarp tissue, dorsoventrally
flat, narrowly to broadly elliptic in outline, with narrow or large, concentric, membranaceous

wings, and entire margin.

Distinctive features: Twining vines with strong fetid odor, due to Sulphur containing

compounds (Takeda et al., 1991); fruits are unique in being laterally compressed, crustaceous to

chartaceous, brittle, splitting into two flat mericarps that remain suspended by thin carpophores

and later dispersed by the wind.

Distribution: A pantropical genus of ~30 species distributed in Asia, Africa and Madagascar,

with only two species native to the Neotropics, i.e., P. ciliata (Bartl. ex DC.) Standl., endemic to

Mexico, and P. brasiliensis (Britton) Standl., distributed in Brazil, Peru, Bolivia, Paraguay and

Argentina; common in calcareous substrates; 0–1,200 m.

PALICOUREA Aublet, Hist. Pl. Guiane 1: 172. 1775, sensu lato (incl. Psychotria subgen.

Heteropsychotria, Cephaelis)

Erect shrubs, small trees, or scrambling or clambering shrubs; unarmed. Stipules

persistent, sheathing at base, bifid, with two short–long, triangular lobes on each side of the stem

(without interpetiolar appendages). Leaves opposite, petiolate; blades chartaceous, papyraceous,

or subcoriaceous. Inflorescence (in climbing species) terminal or subaxillary, paniculate or

corymbiform, terminating in flowering heads or cymules. Flowers 4–5-merous, heterostylous.

Calyx persistent, tube reduced, lobed; lobes narrowly triangular or lanceolate; corolla narrowly

funnelform, white or cream-white, tube glabrous outside, glabrous or puberulent, without a

pubescent ring at base inside (in climbing species); lobes valvate, ovate to narrowly ovate,

margin entire; stamens included (in long-styled flowers), partially exserted or exserted just

beyond or well beyond the corolla (in short-styled flowers), filaments inserted at the lower,
middle or upper part of the corolla tube or near the mouth of corolla tube, short to long, anthers

elliptic; ovary 2-locular (in climbing species); ovules 1 per locule, basally inserted, style

included (in short-styled flowers) or exserted just beyond or well beyond the corolla (in long-

styled flowers), branches as many as the locules, ovate, elliptic, oblong or linear. Fruit

drupaceous, fleshy, subglobose, with 2 pyrenes (in climbing species). Pyrenes woody,

dorsoventrally convex, dorsally costate, ventrally sulcate.

Distinctive features: Palicourea, as currently delimited, has a very variable habit, ranging from

erect, trailing, decumbent, scrambling or rarely epiphytic, herbs, shrubs or trees. It is

characterized by the stipules basally sheathing with two triangular lobes on each side of the stem,

and without interpetiolar appendages (present in Notopleura).

Distribution: Palicourea (sensu lato) is a neotropical genus of ~650 species occurring in

Mexico, Central America, Greater and Lesser Antilles, and South America south to Bolivia and

southern Brazil. At least P. scandens C.M. Taylor from the Pacific slopes of the Colombian

Andes is sometimes reported as a climber (Taylor, 2019); primary and secondary forests, Andean

cloud forests; 100–1,900 m.

RAMONADOXA Paudyal & Delprete, Bot. J. Linn. Soc. 187: 389. 2018.

Erect shrubs, treelets or scrambling woody vines; unarmed. Branches glabrous, terete;

young branches sometimes slightly resinous; basal internodes often laterally compressed below

nodes. Stipules persistent, connate at base, deltoid to narrowly triangular. Leaves opposite,

petiolate; blades chartaceous to coriaceous, ovate to oblong, domatia absent. Inflorescence

axillary, paniculate, pedunculate, many-flowered; bracts small, lanceolate or triangular. Flowers

4-merous, homostylous. Calyx persistent, tube narrowly obovoid, shallowly lobed; corolla
tubular-subcylindrical, slightly inflated at the middle, purple-brown outside, deep yellow inside;

tube not reduplicate, glabrous throughout, lobes 4, narrowly imbricate in bud, short, ovate,

obtuse at tip; stamens included, filaments connate into a short tube at base, inserted at base of

corolla tube, lower half puberulent, anthers linear; ovary 2-locular, with 1 ovule per locule,

apically inserted, pendulous, style exserted, filiform, basally puberulent, capitate, with 2 minute,

connivent style branches. Fruit drupaceous, spongy, white, subglobose to obovoid; pyrenes 2,

woody, hemi-ellipsoidal, 1-seeded. Seeds cylindrical, terete.

Distinctive features: A recent segregate of Chiococca, distinguished by the narrowly obovoid,

shallowly lobed calyx tube, the tubular-subcylindrical corollas, with tube not reduplicate and

slightly inflated at the middle, purple-brown outside and deep yellow inside; while in Chiococca

the calyx tube is extremely reduced, with lobes broadly to narrowly triangular, the corollas are

campanulate to funnelform, with tube narrowly reduplicate, white, cream-white to pale yellow

throughout.

Distribution: A monotypic Cuban genus; thickets and pinelands of eastern Cuba; 300–450 m.
RANDIA Linnaeus, Sp. Pl. 1192. 1753.

Basanacantha J. D. Hooker (1873).

Dioecious shrubs or small to medium-sized trees, sometimes scrambling shrubs or rarely

twining lianas; unarmed or armed with 2–4

axillary, straight, stout, or acicular thorns per

node (Figure 218D). Stipules persistent,

connate at base, broadly to narrowly

triangular, broadly ovate or lanceolate. Leaves

opposite, sessile or short- to long-petiolate;

blades broadly ovate, ovate, elliptic, obovate,

oblong to lanceolate, chartaceous,

papyraceous or subcoriaceous; domatia tufts

of sparse or dense hairs, or absent.

Inflorescence axillary or terminal on axillary,

short shoots, rarely cauliflorous; male

inflorescence fasciculate or corymbose, few-

to many-flowered, or rarely uniflorous; female

Randia retroflexa, photo by P. Acevedo.

inflorescences 1-flowered. Staminate and

pistillate flowers dimorphic with respect to the perianth. Calyx persistent, tube cupular or

tubular, truncate, with small teeth or lobed, or tube absent, with free small to foliose lobes;

corolla hypocrateriform or narrowly infundibuliform, white to cream-white, or yellowish white,

turning pale yellow to yellow at later stages of anthesis, tube internally glabrous, pubescent or

villous without a pubescent ring inside, lobes left-contorted, ovate, suborbicular, oblong, oblong-
ovate to lanceolate. Staminate flowers: 4–6-merous; hypanthium and ovary absent; stamens

included, partially exserted or exserted just beyond the corolla, filaments inserted at distal

portion of corolla tube or near the corolla mouth, short or long, anthers elliptic, narrowly elliptic

to oblong; style nonfunctional, included. Pistillate flowers: 5–7-merous; stamens non-functional,

included or partially exserted; ovary 1-locular, with incomplete placenta, or 2-locular, with

parietal placentation, ovules many per locule, style partially exserted or exserted just beyond the

corolla, with 2 ovate branches. Fruit baccate, leathery or somewhat woody, ovoid to oblate, with

seeds surrounded by a gelatinous pulp that turns black when exposed to air. Seeds horizontal,

perlaceous, dorsoventrally compressed, irregularly elliptic in outline.

Distinctive features: Dioecious scrambling shrubs or rarely twining lianas with straight, stout or

acicular thorns that are opposite or whorled, 3–4 per node, and lack vestigial nodes (present in

Chomelia, Guettarda and Machaonia). Inflorescence in Randia is axillary or terminal on lateral

short shoots, or rarely cauliflorous. In staminate individuals, the inflorescence is fasciculate or

corymbose, few- to many-flowered, or very rarely uniflorous. In pistillate individuals, the

inflorescences are always single-flowered. Fruits are characteristic in being externally leathery or

thinly woody with seeds surrounded by a gelatinous pulp that usually turns black when exposed

to air.

Distribution: A neotropical genus of ~90 species, nine of which are reported as lianas which are

mostly found in Mexico with a few species in Central America, and lowlands of South America;

moist forests; 0–900 m.

ROGIERA Planchon, Fl. Serres Jard. Eur. 5: 442. 1849.

Shrubs, or small to medium-sized trees, rarely scrambling shrubs or woody vines;

unarmed. Stipules persistent or

caducous, free or connate at

base, triangular to linear, erect

or reflexed. Leaves opposite

(rarely ternate), isophyllous or

anisophyllous, short- to long-

petiolate; blades ovate, elliptic,

or ovate-oblong, chartaceous,

subcoriaceous or coriaceous;
Rogiera edwardsii, photo by N. Ramírez Marcial.
domatia tufts of sparse or dense

hairs, or hairy pockets, or absent. Inflorescence terminal or axillary, cymose, paniculate or

corymbose, pauci- or multi-florous. Flowers heterostylous, fragrant. Calyx lobes free or

shallowly connate at base, 4–6(–7), usually unequal, oblong to ovate-triangular to narrowly

triangular, persistent; corolla actinomorphic, hypocrateriform, white, yellowish white, yellow,

red, or purple, tube glabrous or pubescent and with a ring of yellow or white hairs at the mouth,

lobes 4–6(–7), imbricate, oblong, triangular-ovate to round, margin entire, undulate or crisped,

rounded at apex; stamens included in long-styled flowers or partially exserted in short-styled

flowers, filaments inserted at median or distal portion of tube, shorter than the tube or absent,

equal, glabrous, anthers narrowly oblong to linear, basifixed; ovary 2-locular with axile

placentation, ovules many per locule, style exserted just beyond the corolla mouth in long-styled

flowers or included in short-styled flowers, terete, glabrous, 2 lobed. Fruit capsular, loculicidal,
woody, globose, ellipsoid or obovoid. Seeds horizontal, small, angular, globose, fusiform or

falciform, narrowly or partially winged; wing bipolar or unipolar, rarely absent.

Distinctive features: Rogiera, Arachnothryx and Rondeletia are similar in being woody plants

with capsular fruits. Rogiera is similar to Rondeletia by the loculicidal capsules, from which it

differs by the corolla with a ring of yellow or white hairs at mouth (vs. corolla with a thick fleshy

glabrous ring at mouth in Rondeletia), and the horizontal, angular, globose, fusiform or falciform

seeds (vs. vertical, dorsoventrally compressed, fusiform, irregularly elliptic or bipolar seeds in

Rondeletia). Arachnothryx differs from Rogiera by the corolla mouth glabrous, without a thick

fleshy ring or a ring of yellow or white hairs.

Distribution: A neotropical genus of 10 species, of which R. breedlovei (Lorence) Borhidi

always grows as a climbing shrub, while R. edwardsii (Standl.) Borhidi, R. gratissima Planch. &

Lind, and R. ligustroides (Hemsl.) Borhidi are trees or shrubs that sometimes grow as scrambling

shrubs, these are distributed from southern Mexico to Guatemala and Honduras; evergreen

primary and secondary forests, cloud forests, and disturbed vegetation; 600–2,800 m.

RONDELETIA Linnaeus, Sp. Pl. 72. 1753.

Shrubs, or small to medium-sized trees or rarely scrambling shrubs or woody vines;

unarmed. Stipules persistent or caducous, free at base, ovate, triangular, narrowly triangular to

subulate, erect. Leaves opposite, short- petiolate; blades ovate or elliptic, chartaceous; domatia

tufts of hairs. Inflorescence terminal, cymose, paniculate-thyrsoid, multi-florous. Flowers 5-

merous, heterostylous, fragrant. Calyx lobes equal, ovate, narrowly triangular, persistent. Corolla

actinomorphic, hypocrateriform, white, with a thick fleshy glabrous ring at mouth; tube glabrous

outside; glabrous inside; lobes imbricate, equal to subequal, obovate, margin entire or slightly
undulate, round at apex. Stamens included or partially exserted (only tips exserted); filaments

inserted at distal portion of the corolla tube, short, equal, glabrous; anthers narrowly oblong;

round at base and at apex, dorsifixed. Style exserted just beyond the corolla mouth (long-styled

flowers) or included (short-styled flowers), terete, glabrous; lobes 2, oblong to linear. Ovary 2-

locular; placentation axile; placenta peltate to the entire length of the septum; ovules many per

locule. Fruit capsular, loculicidal, dehiscing basipetally, globose, woody. Seeds vertical, with a

central hilum, dorsoventrally compressed, fusiform or irregularly elliptic, winged, wing bipolar.

Distinctive features: Rondeletia is similar to Rogiera and Arachnothryx by being woody plants

with small, globose, subglobose, ellipsoid or obovoid capsular fruits. The diagnostic characters

to differentiate these three genera are compared under Rogiera (see above).

Distribution: A genus of ~140 species, with R. purdiei Hook. f., native to Colombia and

northern Venezuela, as the only species that grows as a climber in addition to being a tree or a

shrub (Bernal et al. 2015, onwards); primary and secondary dry forests, Caribbean coastal

forests, 5–900 m.
SABICEA Aublet, Hist. Pl. Guiane 1: 192, t. 75. 1775.

Herbaceous to woody, twining or scrambling vines, rarely subshrubs; unarmed; stems

slender (up to 1.7 cm in

diam.) and in some

species reaching > 10

m in length; cross

section with regular

anatomy, the xylem

with shallow phloem

arcs and inconspicuous

rays. Stipules
Sabicea panamensis, photo by P. Acevedo.
persistent, free at base,

ovate to broadly triangular, sometimes ligulate or lanceolate. Leaves opposite, petiolate; blades

narrowly elliptic to obovate, ovate to lanceolate, papyraceous to subcoriaceous. Inflorescence

axillary, one or two per node, compact-verticillate, fascicled, glomerulate, capitate or laxly

paniculate cymes, sometimes subtended by broadly lanceolate or ovate involucral bracts.

Flowers 4–5-merous, homostylous, or rarely heterostylous, sessile to shortly pedicellate. Calyx

persistent, campanulate to infundibuliform, lobes antrorse to reflexed, lanceolate to elliptic,

ligulate, triangular or linear; corolla salverform or hypocrateriform, white, cream-white or pink

(rarely purple), lobes valvate, ovate to lanceolate; stamens 4–5(–10), included, filaments short,

anthers oblong; ovary 4–5(–7)-locular, with many ovules per locule, style included in short-

styled flowers or exserted beyond corolla mouth in long-styled flowers, style branches 4–5(–10),

oblanceolate, elliptic, oblong or linear. Fruit baccate, fleshy, globose, subglobose, ellipsoid to
turbinate, red to red-purple, dark violet or maroon, crowned by the persistent calyx. Seeds many,

minute, variously angular, brownish orange.

Distinctive features: Twining herbaceous or subwoody vines with ovate or widely triangular

stipules, or less often scrambling vines; corolla salverform or hypocrateriform, white, cream-

white or pink with valvate lobes; fruit a fleshy, multi-seeded berry, crowned by the persistent

calyx.

Distribution: A pantropical genus of ~170 species, represented in the Neotropics by ~54

species of which 42 are climbers that reach two or more m in length; Mexico, Central America,

Greater Antilles, most of South America, to southern Brazil and Bolivia (Khan 2007); seasonal

forests, savanna formations, low altitude rainforests, primary and secondary cloud forests; 0–

1,800 m.

SALZMANNIA A.P. de Candolle, Prodr. 4: 617. 1830.

Shrubs or trees with erect or scandent branches, sometimes leaning shrubs; unarmed;

branches covered with copious resinous exudates. Stipules persistent, sheathing at base,

triangular, shallowly orbicular or truncate, glabrous, with internal colleters exuding abundant

resinous exudate. Leaves opposite, short- to long-petiolate; blades lanceolate, elliptic, broadly

elliptic, oblong to obovate, chartaceous, subcoriaceous to coriaceous; domatia absent.

Inflorescence axillary or subterminal, sessile, subsessile or pedunculate, uniflorous or 3–18-

flowered heads; bracts when present often foliaceous, lucid, glabrous. Flowers 4–5-merous,

homostylous, sessile or pedicellate. Calyx persistent, cupular, shortly lobed, lobes broadly

triangular, ovate, orbicular-ovate, triangular or deltoid, imbricate, persistent, glabrous or

pubescent; corolla actinomorphic, tubular, pale green to greenish yellow, rarely with reddish or
purplish tinge on lobes, tube cylindrical or tetragonal, not reduplicate, glabrous, lobes narrowly

imbricate in bud, much shorter than tube, ovate, deltoid or narrowly triangular, glabrous; stamens

included, filaments inserted at base of the corolla tube, connate at base into a minute, slender,

glabrous or pubescent, anthers narrowly oblong to linear, basifixed; ovary 2-locular with axile

placentation, ovules 1 per locule, inserted near the roof of the locule, style as long as the corolla

tube, terete, glabrous, with 2 linear lobes. Fruit drupaceous, fleshy, globose, ovoid to ellipsoid,

pinkish red, lavender, red to purple-blue; pyrenes 2, woody, plano-convex. Seeds pendulous,

narrowly ovoid to narrowly obovoid.

Distinctive features: Scrambling shrubs, similar to Chiococca but distinguished by the stems

covered by abundant resinous exudate, the tubular corollas with a non-reduplicate tube, and the

fruits that are pink to red or purplish blue; whereas in Chiococca the stems aren’t resinous, the

corollas are campanulate or broadly infundibuliform with a narrowly reduplicate tube, and the

fruits are white or yellowish white.

Distribution A neotropical genus of 4–5 species with disjunct distribution, one species in

Venezuela, and 3–4 species in the coastal dunes “restingas” and forested vegetation of north-

eastern Brazil; only S. nitida DC. from NE Brazil is reported as a scrambling shrub; 0–140 m.

SCHRADERA Vahl, Eclog. Amer. 1: 35. 1797 [“1796”].

 Herbaceous or woody vines, rarely terrestrial, hemiepiphytic or epiphytic shrubs, or

woody lianas; unarmed.

Plants with a juvenile

phase morphologically

different from the adult

phase. Juvenile plants < 1

m long, slender, sparsely

branched, with

adventitious roots at the

Schradera exotica, photo by P. Acevedo. nodes, and small,

membranaceous, crenate leaves. Adult plants developing into woody vines with scandent or

hanging branches and larger, coriaceous, entire leaves; stem up to 20 cm diam. and 25–30 m

long; cross section with axial elements divided in radial segments by wide rays in Schradera

exotica (J. F. Gmel.) Standl. (Figure 217D). Stipules large, caducous, sheathing at base, forming

a basal tube, lobes round, ovate to triangular, appressed to each other or imbricate. Leaves

opposite, petiolate; blades elliptic, elliptic-oblong to oblanceolate, chartaceous to coriaceous;

domatia absent. Inflorescences axillary or terminal, 2 or 3 per node, pedunculate, hemispherical

or globose capitula, commonly pauci- or multi-florous (rarely uniflorous), subtended by a

discoid, small to large involucre, that is truncate, undulate or irregularly divided at later stages.

Flowers free or basally connate, 5–6(–10)-merous, bisexual, sessile, heterostylous, fragrant, and

nocturnal. Calyx persistent, tubular, margin truncate or undulate; corolla hypocrateriform, white,

tube pilose at distal portion inside, lobes valvate, reflexed, narrowly lanceolate to linear, fleshy,

and entire; stamens 5–6(–10), included, inserted near middle of the corolla tube in long-styled
flowers or partially exserted, inserted near the mouth of corolla tube in short-styled flowers,

filaments glabrous, anthers oblong; ovary 2(4)-locular, with many horizontal ovules per locule,

style included in short-styled flowers or exserted in long-styled flowers, stigmatic branches 2(4),

linear. Fruits commonly basally connate, individual fruits few to many (per head), baccate,

globose to subglobose, fleshy, smooth, greenish white, white or yellowish white. Seeds many,

horizontal, dorsoventrally compressed, quadrangular.

Distinctive features: Root-climbing or epiphytic lianas with juvenile and adult phases, bearing

pedunculate heads subtended by a discoid involucre, with congested, free or basally connate

flowers.

Distribution: A pantropical genus of ~65 species distributed in tropical America, Southeast Asia

and New Guinea, of which 40 are distributed in the Neotropics, 22 of which are reported as

climbers; Costa Rica south to the Amazon basin and the Atlantic Forest in Brazil, including the

West Indies; premontane forests, low and high altitude rainforests, cloud forests; 150–1,700 m.

SPERMACOCE Linnaeus, Sp. Pl. 102. 1753.

Borreria G. Mey. (1818).

Annual or perennial herbs, subshrubs, sometimes scrambling or leaning, up to 4 m long;

unarmed. Stipules persistent, often withering on the stem, sheathing and connate to the petioles,

fimbriate, with 5–9 setae, each seta with an apical colleter. Leaves opposite, sometimes

seemingly whorled, subsessile or short-petiolate; blades narrowly ovate, elliptic to narrowly

elliptic, membranaceous, chartaceous or papyraceous; domatia tufts of sparse hairs, or absent.

Flowering branches with 3–13 terminal and axillary flowering glomerules; glomerules 13–32-

flowered, usually subtended by leaf-like bracts. Flowers 4-merous, heterostylous. Calyx

persistent, lobes narrowly triangular or linear; corolla infundibuliform, white to bluish white,

tube glabrous or puberulent outside, glabrous, puberulent or pubescent at medial or distal portion

inside, with moniliform hairs, lobes valvate, ovate-triangular; stamens partially exserted or

exserted just beyond the corolla, inserted at distal portion, near the mouth of the corolla tube or

at lobes sinuses in short-styled flowers or included, inserted at median portion of corolla tube in

long styled flowers, filaments short or long, anthers oblong; ovary 2-locular, ovules 1 per locule,

centrally inserted, vertical, style glabrous, branches 2, oblong. Fruit (in climbing species) dry,

schizocarpic, separating into two indehiscent mericarps, or a septicidal capsule with two

dehiscent cocci. Seeds vertical, centrally inserted, plano-convex, obovate or elliptic in outline,

dorsoventrally compressed, finely foveolate or transversally sulcate on the dorsal side, with a

well-developed strophiole on the ventral side.

Distinctive features: Scrambling herbs (sometimes woody at base); stipules basally sheathing

and fimbriate, with 5–9 setae, each of which has an apical colleter; flowers heterostylous; fruits

schizocarpic, separating into two indehiscent mericarps (e.g., S. schumannii (Standl. ex

Bacigalupo) Delprete) or a septicidal capsule (e.g., S. valens (Standl.) Govaerts). Spermacoce

resembles Denscantia in the fimbriate stipules and the capsular fruits. However, Denscantia is

distinguished by the thyrsoid inflorescences with branchlets terminating in fascicles or

glomerules, homostylous flowers, capsules apically dehiscing down to the seed insertion and

seeds with very narrow wings (vs. inflorescence capitate, usually subtended by leaf-like bracts,

and sometimes on flowering branches as a series of few- to many-flowered glomerules, flowers

heterostylous, capsules usually dehiscing down to the base, and seed wingless in Spermacoce).

Distribution: A pantropical genus of ~300 species, with ~80 species in the Neotropics, of which

S. valens from Minas Gerais, Brazil, and S. schumannii from Colombia, Brazil, Paraguay and
Argentina, sometimes grow as scrambling herbs up to 4 m in length; humid and seasonal forests,

savanna formations, rocky outcrops, secondary vegetation and ruderal areas; 170–1,200 m.

UNCARIA Schreber, Gen. 125. 1789 (nom. cons.).

Monoecious or rarely dioecious, scrambling lianas or shrubs with scrambling lateral

branches. Stems initially quadrangular, turning

cylindrical with age, up to 15 cm in diam. and >

20 m in length; cross section regular, often with a

quadrangular medulla, inconspicuous rays, and

the cortex exhibiting overlapping layers of

sclerenchyma (Figure 217A). Axillary, hook-

shaped thorns (Figure 218C) helping the plant to

hold on to the phorophyte. Stipules persistent or

caducous, free at base, deltoid, obovate or

cordate. Leaves opposite, short- to long-petiolate;

blades ovate, elliptic, obovate or oblong,

Uncaria guianensis, photo by P. Acevedo. subcoriaceous to coriaceous; domatia tufts of

sparse hairs, hairy-pockets or tuft-pits, or absent. Inflorescence terminal, paniculate, with

branches terminating in globose few- to many-flowered heads, sometimes at the top of lateral

thorns. Flowers 4–5(-6), bisexual or functionally unisexual (staminate and pistillate flowers on

separate individuals). Calyx persistent, funnel-shaped or short tubular, lobes broadly to narrowly

ovate; corolla hypocrateriform, white to cream-white, or yellow when young and turning red at

later stages, tube externally retrorsely puberulous, villous or sericeous, internally glabrous or
sericeous, lobes narrowly imbricate, round, ovate, narrowly ovate or oblong; stamens partially

exserted, filaments inserted near the mouth of the tube, anthers elongate, with a pointed

extension at base and an acute extension at apex; ovary 2-locular with many ovules per locule,

peltate or along the entire length of the septum, style exserted well beyond the corolla, glabrous,

capitate. Fruit capsular, dehiscing septicidally from the apex, thinly woody, ovoid, narrowly

ovoid to turbinate, sessile or long-pedicellate. Seeds many, ascendingly imbricate, laterally

compressed, membranaceous, irregularly narrowly oblong to fusiform in outline, wings bipolar,

margin entire.

Distinctive features: Scrambling lianas or shrubs with hook-shaped axillary thorns;

inflorescences terminal, paniculate, with branches terminating in globose heads.

Distribution: A genus occurring in tropical Africa and tropical America, with ~34 species, two

of which are found in the Neotropics. Uncaria guianensis (Aubl.) J.F. Gmel. from South

America is recognized by the strongly recurved thorns, glabrous branches and leaves and the long-

pedicellate capsules, while U. tomentosa (Schult.) DC., from southern Mexico south to the

Amazon Basin, is recognized by the slightly curved thorns, sparsely pubescent branches and

leaves, and the sessile capsules; along rivers, moist forests, secondary forests; 0–300 m.