Neotropical Brunelliaceae

Clara Inés Orozco

Universidad Nacional de Colombia, Bogotá, Colombia. 

Description

Tree or treelets, sometimes with clear lenticels in cortex. Plants dioecious or gynodioecious. Stem and branches terete, unbranched in lower part; branch nodes swollen.  Leaves petiolate, stipulate, unifoliolate or multifoliolate with 3-26(30) leaflets, opposite or whorled; interpetiolar stipules present varying in shape and number; leaf lamina olive green and often shiny adaxially, frequently leathery, broadly oblong, ovate or broadly elliptic, indumentum of simple hairs, always present on abaxial side of lamina, with few exceptions, trichomes glandular or not, patulous, appressed, curled or woolly, sometimes arachnoid, base often asymmetric, cuneate or obtuse, apex acute or obtuse, margin serrate or crenate; secondary venation semi-craspedodromous, with (7)15-23(32) pairs of secondary veins, petiole sulcate adaxially, stipels often present on petioles of species with 1 to 3 leaflets, in species with more than 3 leaflets stipels present on leaf rachisInflorescences proliferating monotelic thyrso-paniculate, generally pedunculate, varying from simple triads to pleio-thyrsoid paracladia.  Flowers apetalous, hermaphroditic or unisexual (then always female), 4-16 mm in diameter, usually largest in unifoliolate species; sepals (3-)4-5(-6), lobes valvate, persistent in fruit, even or uneven in size due to weak fusion of sepals; stamens inserted on a disk, in two whorls, outer alternisepalous and inner oppositisepalous, sometimes apparently polyandrous, increase or reduction of obdiplostemonous condition is frequent; ovary usually 2-3-, sometimes 4-5- and rarely 6-8-carpellate, carpels free, sometimes weakly fused at base; style subulate, apex inclined, stigma sutural, decurrent, papillose; ovules 2-3 per carpel, anatropous, epitropous.  Fruit of various follicles, these covered by dense red or yellow indumentum of either bristly hairs or smooth trichomes, dehiscent ventrally; endocarp thick and hard, either U-shaped, urceolar or navicular.  Seeds red, 2 to 3 per follicle, surface shiny, expelled at dehiscence.

Notes on delimitation

  • Although several morphological studies were conducted in Brunellia SEM studies of floral ontogeny are lacking, and molecular data are desirable to build a comprehensive phylogeny of the family.
  • In the APG system the Brunelliaceae belongs to the order Oxalidales which also includes the Huaceae, Cephalotaceae, Connaraceae, Cunoniaceae, Elaeocarpaceae and Oxalidaceae.  Within the Oxalidales, the Brunelliaceae is part of a clade that includes Cunoniaceae, Cephalothaceae and Eleaocarpaceae, and is considered sister to the Cephalotaceae (Bradford & Barnes, 2001).
  • Savolainen et al. (2000) consider Brunelliaceae as sister to (Eleocarpaceae +(Cephalotaceae+ Cunoniaceae)).  Orozco (1997) used morphological characters and did not include the same families, but provided a topology with Connaraceae as as sister to (Cunoniaceae, exception Spiraeanthemum and Ascmithia + (Brunelliaceae +Spiraeanthemun and Ascmithia)).

Distribution in the Neotropics

  • The family is monogeneric: Brunellia has c. 54 species, distributed widely in the Andes, the Neblinatepui in Venezuela, Central America and the Antilles.

Key differences from similar families

  • The presence of interpetiolar stipules and stipels on the rachis of the compound leaves or on the petiole of the unifoliolate leaves are diagnostic and are particularly useful to distinguish them from high altitude species of Turpinia (Staphylaceae). The latter present the same variation of the leaves but without stipules and stipels.

Number of genera

  • One: Brunellia Ruiz & Pavón

Useful tips for generic identification

The genus can be recognized by the following vegetative characters:

  • Trees or treelets, with opposite or whorled stipulate leaves.
  • Other vegetative characters are the dense simple indumentum on the abaxial side of the leaves, the adaxial side often shiny and the leaves often leathery. The craspedodromous venation is another useful leaf character.
  • The absence of petals, the apocarpic ovary, the ventral dehiscence of the follicles, the frequent presence of prickly hairs especially on the fruit of the species with more than one leaflet are useful characters to recognize Brunellia.  The hard endocarp and the clearly exposed bright red seeds, visible following ventral dehiscence of the fruit, which is composed of independent follicles, are also very important characters in Brunellia.

Status

  • Endemic.

Important literature

Bradford, J. C. & Barnes RW. 2001. Phylogenetics and classification of Cunoniaceae (Oxalidales) using chloroplast DNA sequences and morphology. Systematic Botany 26: 354 - 385.

Cuatrecasas, J. 1970.  Brunelliaceae. Flora Neotropica Monograph 2: 1 - 189.

Cuatreacasas, J.  1985.  Brunelliaceae. Flora Neotropica Monograph 2 (suppl.): 29 - 103.

Merran, L. & P. K. Endress 2002.  Comparative floral structure and systematics in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae Botanical Journal of the Linnean Society .140 321-381.

Orozco, C. I. 1997.  Sobre la posición sistemática de Brunellia Ruiz & Pavon. In: Evolutionary biology of Brunellia Ruiz & Pavón  (Brunelliaceae, Oxalidales) (eds. Academia de Ciencias Exactas, Físicas y Naturales, Universiteit van Amsterdam), pp. 35 - 57. Guadalupe, Bogotá

Orozco, C. I. & F. Weberling  1999.  A comparative study of inflorescence in Brunellia Ruiz & Pavón and related taxa.  Beitr. Biol. Pflanzen 71: 261-279.

Orozco, C. I. 2001- 2002.  Evolutionary biology of Brunellia Ruiz & Pavón (Brunelliaceae, Oxalidaldes) (eds. Academia de Ciencias Exactas, Físicas y Naturales and Universiteit  van Amsterdam), Guadalupe, Bogotá

Orozco, C. I. 2001. Pollen morphology of Brunellia Ruiz & Pavón and related taxa.  Grana 40: 241 - 255.

Orozco, C. I. 2002. Leaf anatomy in Brunellia Ruiz & Pavón. In : Evolutionary biology of Brunellia   Ruiz & Pavón  (Brunelliaceae, Oxalidales) (eds. Academia de Ciencias Exactas, Físicas y Naturales and  Universiteit van Amsterdam), pp.59- 79. Guadalupe, Bogotá

Orozco, C. I. 2002. A study of floral morphology in Brunellia Ruiz & Pavón.  In: Evolutionary biology of Brunellia Ruiz & Pavón  (Brunelliaceae, Oxalidales) (eds. Academia de Ciencias Exactas, Físicas y Naturales and  universiteit van Amsterdam), pp. 102 - 118 Guadalupe, Bogotá

Savolainen V, Fay MF, Albach DC, Backlund A, van der Bank M, Cameron KM, Johnson SA, Lledó MD,Pintaud J-C, Powell M, Sheanhan MC, Soltis DE, Soltis PS, Weston P, Whitten WM, Wurdack KJ & Chase MW.2000. Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bulletin 55: 257-309.

How to cite

Orozco, C.I. (2009). Neotropical Brunelliaceae. 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/Brunelliaceae.htm.

Click images to enlarge


Brunellia acostae: A. habit, A1. stipules, A2 inflorescence, B follicle closed, C. flower in fruit, D. indumentums of appresed simple hairs, E. nervation, F. seed, G. part of the inflorescences © Clara Inés Orozco, instituto de Ciencias naturales, Universidad Nacional de Colombia.



Brunellia cayambensis : A. habit, B. and C. nervation, D. follicles with ventral dehiscence, bristle-like hairs on the fruit, E. seed, F. interpetiolar stipules © Clara Inés Orozco, Insituto de Ciencias Naturales, Universidad Nacional de Colombia.



Detail of the flowers, calyx as the unique verticil of the perianthium, carpels and stamens of Brunellia comocladifolia © Paul Maas, Utrecht University.



Habitat of Brunellia goudotii © Alvaro Neira, Universidad Nacional de Colombia.



Whorled leaves (a leaflet per leaf ). Follicles developed of Brunellia goudotti © Alvaro Neira, Universidad Nacional de Colombia.



Habit of Brunellia macrophyla (species of very humid Andean forests) © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



The stipels on the rachis of the leaves and the axillary position of the thyrsoid inflorescences of Brunellia propingua © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



Unifoliolate species with opposite leaves. Interpetiolar stipules. Ventral dehiscence of the follicles of Brunellia rufa © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



Unifoliolate species with opposite leaves. Ventral dehiscence of the fruits of Brunellia rufa © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



Plurifoliate leaves, infrutescence of Brunellia sibundoya © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



Margin of the foliar lamina and open flowers of Brunellia sibundoya © Clara Inés Orozco, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.



Young leaves and stipules of Brunellia sp. © Maximilian Weigend, Freie Universität Berlin.



Stipules of Brunellia sp. © Maximilian Weigend, Freie Universität Berlin.



Young fruit of Brunellia sp. (follicles closed) © Maximilian Weigend, Freie Universität Berlin.



Mature fruit of Brunellia sp. (follicles open) © Maximilian Weigend, Freie Universität Berlin.



Brunellia tomentosa : A. habit, B. opened follicles C. inflorescence, D. indumentum on the adaxial side of the leaf lamina, E. stipules and the whorled disposition of the leaves F. seed, G. inflorescence © Clara Inés Orozco, Insituto de Ciencias naturales, Universidad Nacional de Colombia.



Unifoliolate leaves, opposite insertion of Brunellia trianae © Juan Lázaro Toro Corantioquia, Colombia.



Inflorescence of Brunellia trianae © Juan Lázaro Toro Corantioquia, Colombia.



Follicles and endocarp of Brunellia trianae © Juan Lázaro Toro Corantioquia, Colombia.