Neotropical Dilleniaceae

Gerardo A. Aymard

UNELLEZ-Guanare, Estado Portuguesa, Venezuela. 

Description

Lianas, shrubs, small trees with tortuous branches (Curatella Loefl.), or evergreen trees up to 30 m (Dillenia L.); lianas generally with stems > 5 cm diameter, the vascular bundles disposed in bands or concentric rings separated by abundant parenchyma (Doliocarpus Rol., Neodillenia Aymard, Pinzona Mart. & Zucc., several spp. of Davilla Vand.). Leaves alternatesimple, spirally arranged, blades entire or dentate, usually with numerous parallel lateral veins, petiolate, frequently coriaceous and scabrous, glabrous or pubescent with sclerified or silicified simple or fasciculate trichomes, occasionally stellate (Tetracera L.); stipules absent or, if present, soon deciduous, infrequently winged and adnate to the petiole. Inflorescence terminal, axillary, or ramiflorous, paniculate, racemose, cymose, glomerate, or fasciculate (Doliocarpus), or flowers solitary. Flowers actinomorphic or rarely zygomorphic, hypogynous, bisexual or androdioecious (Tetracera). Sepals 2–7(to 14 in Tetracera), frequently 5, imbricate at bases except the two inner sepals of Davilla, cucullate and imbricate for most of length, two larger than others in Davilla; petals 2–7, free or connate at the base, imbricate, often crumpled in bud, deciduous, usually thin and delicate, yellow, white, or rarely pinkish. Stamens usually numerous (50-500), centrifugal, free and persistent; connective broadened and sometimes thickened or connective linear; anthers basifixed, opening by longitudinal slits or apical pores. Carpels 1–20, free or connate along the ventral side (Curatella and Pinzona); placentation parietal to basal, ovules 1 or 2 or numerous in each carpel, apotropous or anatropous; styles free, usually slender and elongate; stigmas terminal, capitate or peltate. Fruits dry and dehiscent follicles or capsule, or berry-like, surrounded by two accrescent sepals in Davilla. Seeds often arillate; endosperm abundant, carnose; embryo small, straight.

Notes on delimitation

  • Some evidence, including seed coat anatomy, suggests a relationship between Dilleniales and Vitales, but relationships between Dilleniales and Caryophyllales have also been suggested (e.g. D. Soltis et al. 2003a; Soltis et al. 2007a). Horn (2006) lists a number of features suggesting a relationship between Dilleniaceae and Rhabdodendraceae, probably sister to the rest of Caryophyllales (Stevens 2001 onwards).

Distribution in the Neotropics

  • Tetracera is the only genus that occurs in both hemispheres, with 45 species of which ca. 20 occur in the neotropical area.
  • Doliocarpus and Davilla are the largest genera that occur in the neotropical region with ca. 40 and 30 species respectively.
  • The remaining genera in the neotropical region are: Neodillenia (three species), Curatella and Pinzona, each with one species.

Distinguishing characters (always present)

Neotropical Dilleniaceae are recognisable by:

Number of genera

  • Curatella Loefl.
  • Davilla Vand.
  • Dillenia L.
  • Doliocarpus Rolander
  • Neodillenia Aymard
  • Pinzona Mart. & Zucc.
  • Tetracera L.

Useful tips for generic identification

Key to genera of Neotropical Dilleniaceae

1. Erect shrubs to trees up to 15 m tall ... 2
1.Scandent to creeping climbers or lianas, or else small erect shrubby, generally < 3 m tall … 3

2. Plants not cultivated with twisted stems and branches; flowers 0.5-3 cm broad, stamens connective broadened and sometimes thickened; receptacle among carpels absent … Curatella.
2. Plants cultived without twisted stems and branches; flowers 8-20 cm broad, stamens connective linear; receptacle among carpels present … Dillenia

3. Carpel 1 per flower … 4
3. Carpels 2-5 per flower … 6

4. Sepals unequal in size, the 2 inner ones larger, covering the fruit entirely; inflorescence in terminal or lateral panicles … Davilla
4. Sepals ± equal in size, never covering the fruit; inflorescence ramiflorous, fasciculate, glomerate, or flowers solitary … 5

5. Stamens free, not forming a ring around the carpel, aril white … Doliocarpus
5. Stamens connate at the base, forming a ring around the carpel, aril red … Neodillenia

6. Carpels 3-5 per flower; fruits follicles or berries … 7
6. Carpels 2 per flower; fruits capsules … 8

7. Inflorescence paniculate; connective broadened, fruits follicles, the aril fimbriate or deeply laciniateTetracera
7. Inflorescence racemose or flowers solitary; connective linear, fruits berries, the aril entireNeodillenia

8. Carpels free; aril white … Davilla
8. Carpels connate ventrally from the base to the apex of the ovary; aril orange … Pinzona

Notable genera and distinguishing features

  • Curatella and Pinzona: Carpels connate ventrally.
  • Davilla: Sepals unequal, the 2 inner ones larger, covering the fruit.
  • Doliocarpus: Vascular bundles disposed in bands or concentric rings separated by abundant parenchyma; inflorescence ramiflorous; the carpel 1, seed with a white aril.
  • Neodillenia: Sepals 1.5--4 cm long; stamens 100--300 or more, fused forming a ring around the carpels; anthers 3--6 mm long, connective linear; aril red.
  • Tetracera: Inflorescences paniculate; fruits follicles, the aril fimbriate or deeply laciniate.

Status

  • Curatella (Endemic).
  • Davilla (Endemic).
  • Dillenia (Cultivated).
  • Doliocarpus (Endemic).
  • Neodillenia (Endemic).
  • Pinzona (Endemic).
  • Tetracera (Pantropical).

General notes

Kubitzki (2004) and Horn (2006) placed the genus Neodillenia (Aymard, 1997) inside of the genus Doliocarpus. By its vascular bundles disposed in bands or concentric rings separated by abundant parenchyma, the connective linear (only in Doliocarpus grandiflorus Eichl. in Mart. and D. magnificus Sleumer), ramiflorous inflorescences and single carpel (only in Neodillenia venezuelana Aymard), the genus Doliocarpus is similar to the genus Neodillenia. However, Neodillenia has ramiflorous inflorescences, racemes or solitary flowers, flower buds 1--4 long cm, sepals 1.5--4 cm long; 100--300 or more stamens, fused forming a ring around the carpels; anthers 3--6 mm long, connective linear, red aril, and 1—5 carpels. In contrast, Doliocarpus always has ramiflorous inflorescences , flower buds ca. 5 mm long, sepals 0.3--1.8 cm long; 25--80 stamens, free not forming a ring around the carpels; anthers 2--3 mm long, broadened and sometimes thickened connective; white aril, and always one carpel.

Important literature

Aymard G. 1991. Dilleniaceae Novae Neotropicae. I. Dos nuevas especies y una nueva subespecie de Doliocarpus de la Guayana Venezolana. Ann. Jar. Bot. de Madrid. 49(2):195-200.

Aymard G. 1993. Dilleniaceae Novae Neotropicae II. Tres nuevas especies y dos nuevas subespecies del género Doliocarpus. Novon 3(4):317-320.

Aymard G. 1995. Dilleniaceae Novae Neotropicae IV. Dos nuevas especies y una nueva subespecie del género Doliocarpus. Ernstia 5(1):27-35.

Aymard G. 1995. Dilleniaceae Novae Neotropicae IV. Dos nuevas especies y una nueva subespecie del género Doliocarpus. Ernstia 5(1):27-35.

Aymard G. 1996. Dilleniaceae Novae Neotropicae VI. A new species of Tetracera from Panama. Novon 6:325-327.

Aymard G. 1997. Dilleniaceae Novae Neotropicae V. El género Doliocarpus en Colombia. Anales Jardín Bot. de Madrid. 55:17-30.

Aymard G. 1997. Dilleniaceae Novae Neotropicae IX. Neodillenia a new genus from Amazon basin. Harvard Papers in Botany 10:121-131.

Aymard G. 1998. Dilleniaceae Novae Neotropicae VIII. Two new species of Davilla from Brazil. Brittonia 50(1):51-55.

Aymard G. 1998. Dilleniaceae. P. Berry et al. (Eds.) Flora of Venezuelan Guayana. Vol. 4:676-685. Missouri Botanical Garden.

Aymard, G. 1998. Four new species of Doliocarpus from Brazil-Amazonian region. Kew Bulletin 53(3):133-140.

Aymard, G. 1999. Dilleniaceae. In: Catalogue of  the vascular plants of Ecuador. P. Jorgensen & S. León-Yánez. (eds.) Monogr. Syst. Bot. Missouri Bot. Gard. 75: 437-439.

Aymard, G. 2000. DILLENIACEAE NOVAE NEOTROPICAE: XI. A new subandean species of Doliocarpus. Brittonia 52(2): 196-199.

Aymard, G. 2001. Dilleniaceae Novae Neotropicae: XII. Doliocarpus humboldtianus, una nueva especie de la Amazonia Brasilera. BioLlania Edic. Esp. No. 7: 8-12.

Aymard, G. 2002. Davilla papyracea (Dilleniaceae) a new species from Brazil. Kew Bull. 57: 487-490.

Aymard, G. 2003. A new species of Davilla (Dilleniaceae) amongst the Flora Săo Paulo, Brazil. Acta Botánica Venezuelica 25(2): 153-159.

Aymard, G. 2003. A new Species of Doliocarpus and a New Species of Tetracera (Dilleniaceae) from Brazil. Novon 13: 1-4.

Aymard, G. 2003. Dilleniaceae. In: The checklist das Plantas do Nordeste do Brasil. http://www.cnip.org.br/bdpn/checklistNE.pdf.

Aymard, G. & B. Boom. 2002. A new species of Tetracera (Dilleniaceae) from Guyana. Brittonia 54 (4): 275-278.

Aymard G. & C . Kelloff. 1993. DILLENIACEAE. In: J. Boggan et al. (eds.) Checklist of the Plants of Guyana. Biological Diversity of the Guianas Program 8:. 87.Smithsonian Institution, Washington, DC.

Aymard G. & C . Kelloff. 1997. DILLENIACEAE. In: J. Boggan et al. (eds.) Checklist of the Plants of Guyana (Guyana, Surinam, French Guiana). 2nd Edition. Biological Diversity of the Guianas Program Publication Series 30:95-96. Smithsonian Institution, Washington, DC.

Aymard, G. & C. Kelloff 2001. DILLENIACEAE. In: T. Hollowell et al. (eds.). Preliminary checklist of the Plants of Guiana Shield (Venezuela; Guyana; Surinam; French Guiana). Biological Diversity of the Guianas Program Publication Series 57: 75-76. Smithsonian Institution, Washington, DC.

Aymard G. & J. Miller. 1994. Dilleniaceae Novae Neotropicarum III. Sinopsis y Adiciones a las Dilleniaceae del Perú. Candollea 49(1):169-182.

Aymard, G. & S. A. Mori. 2002. Dilleniaceae (Dillenia Family). Guide to the Vascular Plants of Central French Guiana: part 2 (Dicotyledons). Mem. New York Bot. Gard. 76(2): 247-251.

Aymard G. & B. Stergios. 1985. Flora de la Mesa de Cavacas. IV Dicotyledoneae: DILLENIACEAE. Biollania 2:18-22.

Dickison, W. C. 1967a. Comparative morphological studies in Dilleniaceae I. Wood anatomy. J. Arnold Arbor. 48: 1--29.

Dickison, W. C. 1967b. Comparative morphological studies in Dilleniaceae II. The pollen. J. Arnold Arbor. 48: 231--240.

Dickison, W. C. 1968. Comparative morphological studies in Dilleniaceae. III. The carpels. J. Arnold Arbor. 49: 317--329.

Dickison, W. C. 1969. Comparative morphological studies in Dilleniaceae IV. Anatomy of the node and vascularization of the leaf. J. Arnold Arbor. 50: 384--398.

Dickison, W. C. 1970a. Comparative morphological studies in Dilleniaceae V. Leaf anatomy. J. Arnold Arbor. 51:89-98.

Dickison, W. C. 1970b. Comparative morphological studies in Dilleniaceae. VI. Stamens and young stem. J. Arnold Arbor. 51: 403-416.

Dickison, W. C. 1971. Comparative morphological studies in Dilleniaceae VII. Additional notes on Acrotema. J. Arnold Arbor. 52: 319--330.

Gilg, E. 1893. Dilleniaceae. pp. 100--128. in: Die Natürlichen Pflanzenfamilien III(6). Engler, A. and K. Prantl (eds.). Germany.

Gilg, E. and E. Werdermann. 1925. Dilleniaceae. Nat. Pflanzenfam. band 21: 7--22.

Horn, J. W. 2006. Dilleniaceae. Pp. 132-154, in Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. Volume IX. Flowering Plants. Eudicots. Berberidopsidales, Buxales, Crossosomatales.... Springer, Berlin.

Kubitzki, K.1970. Die gattung Tetracera (Dilleniaceae) Mitt. Bot. Staatssamml. München 8: 1--98.

Kubitzki, K.1971. Doliocarpus, Davilla, und verwandte Gattungen (Dilleniaceae). Mitt. Bot. Staatssamml. München 9: 1--105.

Kubitzki, K.1973. Neue und bemerkenswerte neotropische Dilleniaceen. Mitt. Bot. Staatssmml. München 9: 707--720.

Kubitzki, K.1980. Eine neue Davilla art aus Venezuela. Mitt. Bot. Staatssmml. München 16: 501--502.

Kubitzki, K.1981. Zwei neuer Doliocarpus arten aus Venezuela. Mitt. Bot. Staatssmml. München 17: 237--238.

Kubitzki, K. 2004. DILLENIACEAE. In: Flowering Plants of the Neotropics. 128-130 Pp. N. P. Smith et al. (eds.). Princenton University Press, Princeton. NJ.

Soltis, D. E., A. E. Senters, M. J. Zanis, S. Kim, S., J. D. Thompson, P. S. Soltis, L. P. Ronse Decraene, P. K, Endress, & J. S. Farris, J. S. 2003a. Gunnerales are sister to other core eudicots: Implications for the evolution of pentamery. American J. Bot. 90: 461-470.

Soltis, D. E., M. A. Gitzendanner & P. S. Soltis. 2007a. A 567-taxon data set for angiosperms: The challenges posed by Bayesian analyses of large data sets. Int. J. Plant Sci. 168: 137-157.

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 7, May 2006 http://www.mobot.org/MOBOT/research/APweb/

How to cite

Aymard, G.A. (2009). Neotropical Dilleniaceae. In: Milliken, W., Klitgĺrd, 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/Dilleniaceae.htm.