Roseli B. Torres
Herbário IAC - Campinas, SP, Brazil
Trees or shrubs, sometimes climbing, sometimes armed, monoecious, dioecious or polygamous. Leaves alternate, usually distichous, rarely opposite, deciduous or persistent, simple, margin entire to serrate, venation pinnate, usually trinerved, cystoliths present, stipules lateral. Inflorescence axillary, racemose, fasciculate or rarely reduced to only one flower. Flowers unisexual (rarely bisexual), not showy, usually 4-5 tepals, free or connate; stamens 4-5, opposite the tepals, anthers longitudinally dehiscent; ovary superior, 2-carpels, 1-locular, locule 1-ovulate, placentation pendulous. Fruit drupaceous; seeds rounded, embryo curved.
Notes on delimitation
- Celtis, Trema and other taxa of woody Cannabaceae were formerly placed in Ulmaceae, subfamily Celtidoideae or Celtidaceae. Although some species of Aphananthe have secondary veins of leaves ending in the teeth, as in the Ulmaceae, it is considered that this feature evolved independently from that of Ulmaceae.
- The presence of unisexual flowers in Cannabaceae is an important feature for the separation from Ulmaceae. However, Trema micrantha has rare bisexual flowers, which are protogynous and sometimes functionally male.
Distribution in the Neotropics
- Aphananthe Planch. - Mexico, Guatemala, El Salvador, Honduras and Nicaragua.
- Celtis L. - Mexico, Central America and Caribbean islands to Argentina.
- Lozanella Greenm. - Mexico to Bolivia.
- Trema Lour. - Mexico, Central America and Caribbean islands to Argentina
Distinguishing characters (always present)
- Trees, or shrubs, sometimes scandent, leaves simple, alternate, generally distichous, rarely opposite (Lozanella), with stipules, cystoliths present; flowers unisexual, inconspicuous; fruit a drupe.
Other important characters
- Trees or scandent shrubs with spines (Celtis).
- Lozanella is very similar to Trema, but with opposite leaves and fused stipules.
Key differences from similar families
- Plants without latex, sometimes armed, never urticating (latex in all parenchymatic tissues in Moraceae or restricted to bark in Urticaceae; plants unarmed in Ulmaceae, Moraceae (except Maclura) and Urticaceae; sometimes urticating in Urticaceae).
- Leaves simple, alternate, distichous, rarely opposite (leaves simple or lobed, alternate, distichous, spiral or opposite in Moraceae and Urticaceae).
- Leaves usually 3-nervate from base (penninerved with secondary venation ending in the teeth of the margin in Ulmaceae; penninerved or palmate in Moraceae and Urticaceae).
- Flower unisexual with apical placentation (flower unisexual or bisexual in Ulmaceae; basal placentation in Urticaceae).
- Fruit drupaceous (samara or nut in Ulmaceae; drupe or achene in Moraceae and Urticaceae, sometimes forming a compound fruit).
Number of genera
About 9 genera, with 4 genera and 20 species in the Neotropics:
- Aphananthe - A. monoica (Hemsl.) J.-F. Leroy.
- Celtis - C. australis L.; C. berteroana Urb.; C. brasiliensis (Gardner) Planch.; C. caudata Planch.; C. chichape (Wedd.) Miq.; C. ehrenbergiana (Klotzsch) Liebm.; C. fluminensis Carauta; C. iguanaea (Jacq.) Sarg.; C. loxensis C.C. Berg; C. pubescens (Kunth) Spreng.; C. reticulata Torr.; C. schippii Standl.; C. spinosa Spreng.; C. tala Gillies ex Planch.; C. trinervia Lam.
- Lozanella - L. enantiophylla (Donn. Sm.) Killip & C.V. Morton; L. permollis Killip & C.V. Morton.
- Trema - T. micrantha (L.) Blume; T. integerrima (Beurl.) Standl.
Useful tips for generic identification
Key to genera of Neotropical Cannabaceae
1. Leaves opposite; stipules united, leaving a conspicuous interpetiolar scar .... Lozanella1. Leaves alternate; stipules free or united only at their base .... 22. Monoecious plants, leaves pinnately veined, the secondary veins ending at the margin .... Aphananthe2. Monoecious, dioecious or polygamous plants, leaves pinnately veined, usually 3-nerved at base, the secondary veins ending before the margin .... 33. Trees, unarmed; tepals induplicate-valvate in bud, stigmas 2, simple.... Trema3. Trees or shrubs, sometimes scandent, sometimes armed; tepals imbricate in bud, stigmas 2, simple or bifurcate .... Celtis
- Lozanella endemic. Alphananthe, Celtis and Trema occur in other regions and continents.
- The Celtis species are hard to identify because their flowers are very similar and the morphological characters usually cited by different authors have great variation. The taxonomy of the genus requires extensive revision, because the available studies do not adequately account for the status of the many published names.
- Trema micrantha is a species of wide geographical distribution and great morphological variability. Studies on its biology in two natural populations revealed that several plants do change their sex during the breeding season. These studies revealed also the occurrence of rare bisexual flowers. We accept two species for the Neotropical region but the genus, like Celtis, also needs extensive revision.
- Some species of Celtis provide hardwood, some are ornamental or have edible fruits. The fruits of Aphananthe monoica are also edible. Trema micrantha is an important species in the regeneration of deforested areas; its fruits are enjoyed by birds; the bark can be used to make handmade paper and its timber for manufacturing industrial paper.
Berg, C.C. & Dahlberg, S.V. 2001. A revision of Celtis subg. Mertensia (Ulmaceae). Brittonia 53(1): 66-81.
Killip, E.P. & Morton, C.V. 1931. The genus Lozanella. Journal of the Washington Academy of Sciences21: 336-339.
Nee, M. 1984. Ulmaceae. In: Flora de Vera Cruz, Fascículo 40, pp. 34-38. Instituto Nacional de Investigaciones sobre Recursos Bióticos, Vera Cruz.
Pennington, T.D., Reynel, C. & Daza, A. 2004. Illustrated guide to the trees of Peru. pp. 93-96. David Hunt, Sherborne.
Systma, K.J.; Morawetz, J., Pires, J.C., Nepokroef, M., Conti, E., Zjhra, M., Hall, J.C. & Chase, M.W. 2002. Urticalean Rosids: circumscription, Rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89(9): 1531-1546.
Romanczuk, M.C. & Martinez, MA.P. 1978. Las especies del género Celtis (Ulmaceae) en la flora Argentina. Darwiniana 21(2-4): 541-577.
Song, B-H., Wang, X-Q., Li, F.-Z. & Hong, D.-Y. 2001. Further evidence for paraphyly of the Celtidaceae from the chloroplast gene matK. Plant Systematics and Evolution 228: 107-115.
Todzia, C.A. 1993. Ulmaceae. In: K. Kubitzki, J.G., Rohwer & Bittrich, V. (eds). The families and genera of vascular plants, pp. 603-611. Springer-Verlag, Berlin.
Torres, R.B. 1996. Biologia da reprodução de Trema micrantha (L.) Blume (Ulmaceae). Tese de Doutorado. Campinas: Universidade Estadual de Campinas.
Torres, R.B. & Luca, A.Q. 2005. Ulmaceae. In: M.G.L. Wanderley, G.J. Shepherd, T.S. Melhem & A.M. Giulietti (coord.). Flora Fanerogâmica do Estado de São Paulo, pp. 361-369. Fapesp & Rima, São Paulo.
Ueda, K., Keiko, K. & Tobe, H. 1997. A molecular phylogeny of Celtidaceae and Ulmaceae (Urticales) based on rbcl nucleotide sequences. Journal of Plant Reseach 110: 171-178.
Wiegrefe, S.J., Systma, K.J. & Guries, R.P. 1998. The Ulmaceae, one family or two? Evidences from chloroplast DNA restriction site mapping. Plant Systematics and Evolution 210: 249-270.
How to cite
Torres, R.B. (2011). Neotropical Cannabaceae. 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/Cannabaceae.htm.
Click images to enlarge
Branch with fruits of Celtis sp. @ Daniela Zappi, RBG, Kew.
Branch with fruits of Celtis pubescens © R.B. Torres, Herbário IAC.
Leaves, flowers, and fruits of Trema sp. (Peru) Mac H. Alford, University of Southern Mississippi.
Branch with male flowers and young fruits of Trema micrantha © R.B. Torres, Herbário IAC
Branch with mature and imature fruits of Celtis micrantha © R.B. Torres, Herbário IAC.