Neotropical Clusiaceae

Mats H.G. Gustafsson

Department of Biological Sciences, Aarhus University, Denmark.  

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.

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.

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.

Distinguishing characters (always present)

Other important characters

  • Most Clusiaceae have a tendency to form adventitious roots.
  • White to yellow latex.
  • Coriaceous opposite leaves.
  • Parallel secondary and intersecondary veins.
  • Many stamens, often variously fused.

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.

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

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 carpelClusia
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 terminalTovomita
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 sessileGarcinia
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 fasciclePlatonia
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

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.

Status

  • Native

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 Garcinia sclerophyllous 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. Mammea americana L., Platonia insignis Mart. and Garcinia spp.
  • The latex has several traditional technical and medicinal uses, and contains compounds with antibiotic and antitumor activity.

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.

How to cite

Gustafsson, M.H.G. (2009). Neotropical Clusiaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Clusiaceae.htm.

Click images to enlarge


Flowers of Caraipa sp. © M. Gustafsson, University of Aarhus.



Fruits of Chrysochlamys membranacea showing ramiflory © M. Gustafsson, University of Aarhus.



Pistillate flower of an apparently beetle-pollinated species of Clusia (sect. Anandrogyne) © M. Gustafsson, University of Aarhus.



Staminate flower of an apparently beetle-pollinated species of Clusia (sect. Anandrogyne) © M. Gustafsson, University of Aarhus.



Fruits of Clusia ducu with seeds surrounded by orange aril © M. Gustafsson, University of Aarhus.



Staminate flower of Clusia fructiangusta with resiniferous synandrium © M. Gustafsson, University of Aarhus.



Pistillate flower of Clusia fructiangusta with resiniferous staminodial ring © M. Gustafsson, University of Aarhus.



Inflorescence of Clusia garciabarrigae © M. Gustafsson, University of Aarhus.



Stem of Clusia loretensis with adventitious roots surrounding "host" tree © M. Gustafsson, University of Aarhus.



Pistillate flower of Clusia pallida, with staminodial, resiniferous ring surrounding ovary © M. Gustafsson, University of Aarhus.



Staminate flower of Clusia pallida with resiniferous synandrium © M. Gustafsson, University of Aarhus.



Clusia species of sect. Anandrogyne with nectariferous flowers visited by wasps © M. Gustafsson, University of Aarhus.



Open fruit of Clusia sp. showing orange aril of seeds © M. Gustafsson, University of Aarhus.



Fruiting individual of Clusia sp. (sect. Anandrogyne) in montane scrub at 2,300 m © M. Gustafsson, University of Aarhus.



Garcinia macrophylla © William Milliken/RBG Kew.



Kielmeyera coriacea © William Milliken/RBG Kew.



Kielmeyera regalis © William Milliken/RBG Kew.



Kielmeyera petiolaris © William Milliken/RBG Kew.



Symphonia globulifera © Denise Sasaki/Programa Flora Cristalino.



Fruit of Tovomita weddeliana showing large, arillate seed and red inside of capsule valves © M. Gustafsson, University of Aarhus.