1. Clusiaceae Lindl.

    1. This family is accepted.

[FZ]

Guttiferae, N. K. B. Robson. Flora Zambesiaca 1:2. 1961

Habit
Trees, shrubs, woody climbers or perennial or annual herbs; juice resinous
Leaves
Leaves opposite or rarely subopposite to alternate or whorled, simple, usually entire, exstipulate, containing various glandular secretions
Flowers
Flowers actinomorphic, bisexual, dioecious or polygamous
Calyx
Sepals (2) 4–5 (6 or more), quincuncial or decussate
Corolla
Petals 4–5 (6 or more), free, usually convolute (sometimes imbricate or decussate, rarely absent), alternating with the sepals
Androecium
Androecium basically of two whorls of stamen fascicles, the outer (antisepalous) one often sterile (fasciclodes) or absent, filaments variously united or free (when the androecium may appear polyandrous); antipetalous fascicles variously united or free, very rarely each one reduced to a single stamen
Gynoecium
Ovary superior, 1–5-locular (rarely to 12-locular), placentation usually axile, sometimes ± parietal, loculi 1–?-ovulate; styles free, ± united or absent; stigmas equal in number to the loculi
Fruits
Fruit a septicidal (rarely loculicidal) capsule or a berry or drupe
Seeds
Seeds sometimes winged, carinate or arillate, without endosperm
[FWTA]

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

Habit
Trees or shrubs, with resinous juice
Leaves
Leaves opposite, simple; stipules absent
Flowers
Flowers actinomorphic, unisexual, polygamous or dioecious, rarely hermaphrodite
Calyx
Sepals 2–6; rarely more, imbricate
Corolla
Petals the same number, hypogynous, contorted or imbricate, very rarely subvalvate
Androecium
Stamens mostly numerous, hypogynous, free or variously united in the lower part or into bundles opposite the petals; anthers 2-celled, opening lengthwise
Sterile Parts
Rudimentary ovary sometimes present in the male flower; staminodes often present in the female flower
Gynoecium
Ovary sessile, superior, 1- to many-celled; ovules 1 to many, on the inner angle of the cells or erect from their base, rarely parietal (Allanblackia); stigmas various, sometimes radiating
Fruits
Fruit dehiscent or not, sometimes large and globose
Seeds
Seeds often with an aril, without endosperm; embryo large; cotyledons often minute
[NTK]

Gustafsson, M.H.G. (2009). Neotropical Clusiaceae.

Morphology
Description

Terrestrial or hemiepiphytic trees, shrubs or lianas, frequently with adventitious roots ; usually glabrous and evidently laticiferous ; exudate mostly white to yellow, sometimes orange, rarely clear or not evident. Leaves simple , exstipulate , entire and usually opposite, with laticifers or glands . Inflorescences terminal or axillary , cymose or less commonly racemose. Flowers hermaphroditic or unisexual (plants then mostly dioecious ), with pedicel often with one or more pairs of bracteoles, these sometimes similar to or grading into sepals. Sepals (2-)4-many, usually free (though may be joined at base). Petals 4-9, often white or cream, less often green, pink, red, orange, purple or blackish. Androecium extremely variable, consisting of 4 to several hundred stamens that may be fused into fascicles or synandria of varying shape, not uncommonly accompanied by staminodes and frequently secreting resin or oils. Ovary superior and syncarpous, consisting of 2-8(-20) carpels, often surrounded by staminodes. Styles 1-8 or stigma sessile . Fruit a dry or fleshy capsule or a berry , rarely a drupe . Seeds often arillate.

Distribution
Distribution in the Neotropics

The family occurs throughout the Neotropics:

  • Calophyllum L., Chrysochlamys Poepp. & Endl., Clusia L., Clusiella Planch. & Triana, Garcinia L., Marila Sw., Symphonia L.f. and Tovomita Aubl. are found in rainforests throughout most of mainland South and Central America.
  • Caraipa Aubl., Haploclathra Benth., Kielmeyera Mart. & Zucc., Mahurea Aubl., Moronobea Aubl., Neotatea Maguire, Platonia Raf., and Thysanostemon Maguire occur in northern South America east of the Andes.
  • Dystovomita (Engl.) D'Arcy and Mammea L. are predominantly Central American.
Diagnostic
Distinguishing characters (always present)
  • Woody plants with entireexstipulate leaves with glands and/or canals.
  • Viscous exudate.
  • Radially symmetrical, choripetalous flowers.
  • Superior, syncarpous ovary.
Key differences from similar families
  • Apocynaceae may be vegetatively similar to some Clusiaceae, but have abundant, fast-flowing latex.
  • Caraipa, Mahurea and Kielmeyera have alternate leaves and sparse latex and may be confused with a number of other families, e.g. the Bonnetiaceae, which completely lack latex, and the Sapotaceae which have latex, and sometimes similar venation and even occasionally opposite leaves, but can be distinguished vegetatively based on the presence of T-shaped hairs.
Other important characters
  • Most Clusiaceae have a tendency to form adventitiousroots.
  • White to yellow latex.
  • Coriaceous opposite leaves.
  • Parallel secondary and intersecondary veins.
  • Many stamens, often variously fused.
Notable genera and distinguishing features
  • Clusia (over 400 spp. all Neotropical). Widespread hemiepiphytes, lianas, small trees and shrubs with fleshy capsules and seed less than 6 mm long with an orange aril. Flowers often with resin.
  • Chrysochlamys (over 60 spp. all Neotropical).  Common understorey treelets in rainforests. Like Clusia but usually ramiflorous, and seeds ca 10 mm long.
  • Tovomita (over 30 spp., all Neotropical).  Like Chrysochlamys but inflorescence usually terminal, sepals only two, and inner fruit wall bright red.
  • Symphonia (one or two spp., ca. 20 in total).  Common trees in swamp forests with bright red globular flowers and stamens fused into a tube with 5 lobes, each bearing 3 anthers.
  • Kielmeyera (ca. 50 spp. most Brazilian).  Small trees of dry open habitats with alternate leaves with distant secondary veins, often large flowers with white or pink contorted petals.
  • Caraipa (ca. 30 spp. all Neotropical).  Common forest trees on acidic soils with sparse latex, alternate leaves, stellate trichomes and white contorted petals.
  • Calophyllum (ca. 10 spp. in Neotropics, ca 180 in total).  Tall forest and beach trees with leaves with dense, parallel secondary veins.
Useful tips for generic identification

Key to genera of Neotropical Clusiaceae

1. Leaves opposite—2
1. Leaves alternate —3

2. Fruit a fleshy capsule, seeds arillate—6
2. Fruit a dry capsule or indehiscent —9

3. Secondary veins closely parallel, tertiary veins not evident—Neotatea
3. Tertiary veins well developed—4

4. Plant with stellate hairs, seeds 1-3—Caraipa
4. Without stellate hairs, seeds numerous—5

5. Seeds (including wing) >2 mm—Kielmeyera
5. Seeds <2 mm—Mahurea

6. Seed less than 6 mm long, usually several per carpel —Clusia
6. Seed longer than 10 mm, one per carpel —7

7. Leaves almost as wide as long, apex rounded, leaf bases deeply excavated—Dystovomita
7. Leaves at least twice as long as wide, leaf bases not strikingly excavated—8

8. Inside of capsule valves dark red, inflorescence usually terminal —Tovomita
8. Inside of capsule pale, inflorescence often pendulous, below leaves—Chrysochlamys

9. Fruit a dry capsule.... 10
9. Fruit indehiscent —. 11

10. Inflorescence racemose, seeds numerous—Marila
10. Inflorescence not racemose, seeds 8 or fewer—Haploclathra

11. Liana with resiniferous flowers, fruit with numerous minute seeds—Clusiella
11. Trees or shrubs with flowers without resin, seeds 1-40—12

12. Style and stigma undivided—13
12. Style distally branched or stigma sessile —14

13. Leaves with secondary veins closely parallel, fruit one-seeded—Calophyllum
13. Venation reticulate, fruit 4-seeded—Mammea

14. Petals not contorted, stamens not obviously fasciculate, stigma sessile —Garcinia
14. Petals contorted, stamens fasciculate, style distally 5-branched—15

15. Flowers umbel-like petals dark red, stamen fascicles fused into 5- lobed tube—Symphonia
15. Flowers borne singly, petals white, pink or orange, fascicles free —16

16. Stamens >15 per fascicle —Platonia
16. Stamens <14 per fascicle —17

17. Petals broad, fascicles spirally twisted—Moronobea
17. Petals narrow, fascicles not twisted—18

18. Anthers 8-9 mm long—Thysanostemon
18. Anthers 10-40 mm long—Lorostemon Ducke

General Description
Number of genera
  • Calophyllum
  • Caraipa
  • Chrysochlamys (perhaps including Tovomitopsis Planch. & Triana)
  • Clusia (including Oedematopus Planch. & Triana, Havetia Kunth, Havetiopsis Planch. & Triana, Decaphalangium Melch., Pilosperma Planch. & Triana, Quapoya Aubl., Renggeria Meisn.)
  • Clusiella
  • Dystovomita
  • Garcinia (including Rheedia L.)
  • Haploclathra
  • Kielmeyera
  • Mahurea
  • Mammea
  • Marila
  • Moronobea
  • Neotatea
  • Platonia
  • Symphonia
  • Thysanostemon
  • Tovomita
Notes on delimitation
  • Clusiaceae are members of the large and heterogeneous order Malpighiales.
  • The two subfamilies Clusioideae and Kielmeyeroideae are well supported groups, but with few uncontradicted characters uniting them.
  • The Hypericaceae have often been included in Clusiaceae, but have recently been shown to be more closely related to Podostemaceae.
General notes
  • The family occurs in most Neotropical habitats, from sea level to 3,500m altitude (spp. of Clusia). Diversity of genera is highest on the Guayana shield and surrounding areas whereas species diversity is highest in and near the Northern Andes.
  • The West Indies are poor in species.
  • Most species are found in rainforests but species of Clusia, Garcinia and Kielmeyera also occur in dry, open habitats (Clusia then often with almost succulent leaves and Garciniasclerophyllous with spine -tipped leaves).
  • Pollination mechanisms are highly diverse and pollinators include bees, wasps, perching birds, bats, flies, beetles, moths, cockroaches and perhaps small arboreal mammals.
  • Several lineages in the family offer resin as a reward for some groups of bees that use it in nest construction, a phenomenon otherwise known only from Dalechampia Plum. ex L. (Euphorbiaceae).
  • The family comprises some important timber trees, particularly Calophyllum spp.
  • Several species are cultivated as fruit trees, e.g. Mammeaamericana L., Platoniainsignis Mart. and Garcinia spp.
  • The latex has several traditional technical and medicinal uses, and contains compounds with antibiotic and antitumor activity.
Status
  • Native
Literature
Important literature

Cuatrecasas, J. 1950.  Notas a la flora de Colombia. X.  Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 8: 33-64.

D'Arcy WG. 1980.  Flora of Panama. Family 123. Guttiferae.  Ann Missouri Bot Gard 67:969-1043.

Gentry, A. H. 1993. A Field Guide to the Families and Genera of Woody Plants of Northwest South America (Colombia, Ecuador, Peru) with supplementary notes on herbaceous taxa. The University of Chicago Press, Chicago.

Gustafsson, M. H. G., V. Bittrich and P. F. Stevens. 2002.  Phylogeny of the Clusiaceae based on rbcL sequences. International Journal of Plant Sciences 163: 1045-1054.

Gustafsson, M. H. G. and V. Bittrich 2003.  Evolution of morphological diversity and resin secretion in flowers of Clusia L. (Clusiaceae): insights from ITS sequence variation. Nordic Journal of Botany 22: 183-203.

Gustafsson, M. H. G., V. Bittrich and K. Winter 2007.  Diversity, phylogeny and classification of Clusia. In U. Lüttge (ed.) Ecological studies vol. 194. Clusia: a woody Neotropical genus of remarkable plasticity and diversity, pp. 95-116. Springer, Heidelberg.

Hammel, B. E. 1986.  New species of Clusiaceae from Central America with notes on Clusia and synonymy in the tribe Clusieae. Selbyana 9: 112-120.

Hammel, B. E. 1999.  Synopsis of Chrysochlamys (Clusiaceae: Clusioideae: Clusieae) in Mesoamerica. Novon 9: 360-374.

Pipoly, J. J., Kearns, D. E. and Berry, P. E. 1998. Clusia. In: Steyermark, J. A., Berry, P. E. & Holst, B. K. (eds), Flora of the Venezuelan Guayana, vol. 4. Timber Press, Portland, pp. 260-294.

Pipoly, J. J. and Graff, A. 1995a.  A synopsis of the genus Clusia sections Criuvopsis and Brachystemon (Clusiaceae) in northern South America. Sida 16: 505-528.

Pipoly, J. J. and Graff, A. 1995b.  The genus Clusia section Criuva (Clusiaceae) in Guayana. Sida 16: 649-678.

Kearns, D. E. & Berry, P. E. 1998.  Clusia. In: Steyermark, J. A., Berry, P. E. and Holst, B. K. (eds), Flora of the Venezuelan Guayana, vol. 4. Timber Press, Portland, pp. 260-294.

Pipoly, J.J. and M.H.G. Gustafsson 2002.  Clusiaceae. In: S.A. Mori, G. Cremers, C.A. Gracie, J.-J. de Granville, S.V. Heald, M. Hoff, and J.D. Mitchell (eds.)., Guide to the vascular plants of central French Guiana. Part 2. Dicotyledons. Memoirs of the New York Botanical Garden 76(2): 211-223.

Ribeiro, J.E.L.S., Hopkins, M.J.G., Vicentini, A., Sothers, C.A., Costa, M.A.S., Brito, J.M., Souza, M.A.D., Martins, L.H.P., Lohmann, L.G., Assunção, P.A.C.L., Pereira, E.C., Silva, C.F., Mesquita, M.R. & Procópio, L.C. 1999. Flora da Reserva Ducke: Guia de identificação das plantas vasculares de uma floresta de terra firme na Amazônia Central. INPA, Manaus. 816 pp.

Stevens, P.F. 2006.  Clusiaceae-Guttiferae, pp. 48-66, in Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. Volume IX. Flowering Plants. Eudicots. Berberidopsidales, Buxales, Crossosomatales. Springer, Berlin.

[FTEA]

Guttiferae, P. Bamps, N. Robson & B. Verdcourt. Flora of Tropical East Africa. 1978

Habit
Trees, shrubs, subshrubs or herbs, rarely climbers, mostly containing a yellow, orange or clear resinous latex, usually glabrous but occasionally with an indumentum of simple or stellate hairs
Leaves
Leaves usually opposite, sometimes verticillate or alternate, simple, entire or very obscurely crenate, exstipulate, usually with glands and resin channels which are often translucent
Inflorescences
Inflorescences terminal or axillary, 2-fid cymes, false racemes, panicles, fascicles or umbels or sometimes the flowers solitary
Flowers
Flowers regular, often fleshy, hermaphrodite, dioecious or polygamous
Calyx
Sepals (2–)4–5(–6 or more), imbricate, often decussate
Corolla
Petals free, 4–5(–6 or more), imbricate or contorted, sometimes decussate
Androecium
Stamens usually numerous, free or very often in fascicles or fused into groups, anthers usually with 2 thecae dehiscing by longitudinal slits; the outer fascicles are often sterile “fasciclodes”, frequently forming what is often called the disc, or are absent
Gynoecium
Ovary superior, 1–several-locular, with parietal, axile, apical or basal placentation; ovules anatropous, l–? per locule; styles free or fused, sometimes very short or absent; stigmas or stigma-lobes mostly equal in number to the locules
Fruits
Fruit a berry, drupe or septicidal (rarely loculicidal) capsule
Seeds
Seeds arillate or not, without albumen, sometimes winged; embryo with cotyledons often very reduced

Images

Clusiaceae Lindl. appears in other Kew resources:

First published in Intr. Nat. Syst. Bot., ed. 2. 74. 1836 [13 Jun 1836] , nom. alt.: Guttiferae (1836)

Accepted by

  • APG IV (2016) http://dx.doi.org/10.1111/boj.12385

Sources

Flora Zambesiaca
Flora Zambesiaca
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of Tropical East Africa
Flora of Tropical East Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of West Tropical Africa
Flora of West Tropical Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2019. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

Neotropikey
Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
http://creativecommons.org/licenses/by/3.0