1. Marcgraviaceae Bercht. & J.Presl

    1. This family is accepted.

[NTK]

Dressler, S. (2009). Neotropical Marcgraviaceae.

Morphology
Description

Terrestrial, hemiepiphytic or epiphytic lianas or shrubs, rarely small trees. Roots sometimes adventitious . Hypophyllous glands on the lower surface of the leaf blade , raphide cells and variously shaped sclereids frequent. Leaves simple , alternate , exstipulate , glabrous , margins entire or minutely crenate . Inflorescences terminal , racemose, sometimes pseudo-umbels or pseudo-spikes, erect or pendulous. Bracts transformed into variously shaped nectaries. Flowers bisexual , actinomorphic , hypogynous, 2-bracteolate, the bracteoles sepaloid or deltoid, sometimes lacking. Sepals 4-5, unequal, free or nearly so, imbricate , persistent . Petals 3-5, imbricate , free or connate . Stamens 3 to many; filaments free or basally connate , uniseriate; anthers basifixed or nearly so, dithecal, tetrasporangiate, introrse, longitudinally dehiscent . Ovary superior , completely or incompletely 2-20-locular; ovules numerous. Fruits subglobose, apiculate with a persistent style and stigma , capsular, loculicidally and septifragously dehiscent from the base (sometimes appearing berry -like), pulpy inside. Seeds hemispherical to reniform , few to numerous with a shiny reticulate testa.

General Description
Notes on delimitation

Two subfamilies:

  • Marcgravioideae: leaves distichous; conspicuous heterophylly between juvenile, plagiotropous and adult orthotropous branches; perianth tetramerous, petals fully fused to form a caducous cap; only central (sterile) flowers of inflorescence fused with nectarybract, other flowers lacking nectary (Marcgravia L.).
  • Noranteoideae: leaves spiral; perianth usually pentamerous, petals distinct to somewhat fused; each flower usually subtended by nectarybract  (all other genera).
  • Vegetatively similar to the Ternstroemiaceae (non-climbing!) this family was formerly considered to be close to Theaceae/Ternstroemiaceae. Recent molecular data associate it rather with Balsaminaceae, and the former Theaceae-satellites Pellicieraceae and Tetrameristaceae.
Number of genera

Seven genera (see "Distribution in the Neotropics").

Status
  • Native and endemic to the Neotropics, Norantea guianensis sometimes cultivated in countries outside its range (e.g. Jamaica, Costa Rica, Trinidad).
General notes
  • The elaborate inflorescences with the variously shaped bracteal “nectary -containers” are frequented by a wide range of visitors (insects, lizards, birds, bats, non-flying mammals). Different pollination syndromes are exemplified in the various taxa. Whereas Ruyschia and Souroubea seem predominantly insect-pollinated (flies?, butterflies, hawk-moths?), Norantea, Sarcopera and Schwartzia brasiliensis (Choisy) Bedell ex Gir.-Cañas are probably best adapted to bird-pollination, especially by perching birds. Sarcopera sessiliflora (Triana & Planch.) Bedell was the first plant where pollen transmission by birds' feet was described. Most Schwartzia, Marcgraviastrum and Marcgravia spp. seem to be primarily bat-pollinated, but the ornithophilous syndrome seems to occur as well. This is a coarse generalisation and surprisingly many Marcgraviaceae are probably autogamous as well (and even cleistogamous, as experimentally proven in Marcgravia coriacea Vahl).
  • The brightly coloured pulp with the small seeds exposed when the capsules split open suggest endozoochorous dispersal.
  • Apart from occasional horticultural use in the tropics the family has no significant economic value.
Distribution
Distribution in the Neotropics
  • Marcgravia (ca. 65 spp.): S Mexico, Mesoamerica, South America, Antilles.
  • Marcgraviastrum (15 spp.): S Nicaragua to Peru, Bolivia plus 2 spp. in E Brazil.
  • Norantea (2 spp.): Caribbean and Amazonian basin of NE South America.
  • Ruyschia Jacq. (9 spp.): Mesoamerica, N Andes, Lesser Antilles.
  • Sarcopera (ca. 10 spp.): Honduras to N Bolivia, Guyayana Highlands.
  • Schwartzia Vell. (ca. 15 spp.): Costa Rica through the Andes south to Bolivia, in the Caribbean basin and 1 sp. in E Brazil.
  • Souroubea (19 spp.): Mexico to Bolivia (absent from the Antilles).
  • The Marcgraviaceae are endemic to the Neotropics.  Species range from Southern Mexico to Northern Bolivia and Eastern Brazil including the Antillean arc. Marcgraviastrum (Wittm. ex Szyszyl.) de Roon & S.Dressler, Souroubea Aubl., Norantea Aubl. and Sarcopera Bedell are missing in the Antilles (although Norantea guianensis Aubl. is rarely cultivated as an ornamental).
  • Taxa are mostly found in primary humid tropical lowland forests or montane rain or cloud forests.
Diagnostic
Distinguishing characters (always present)
  • Woody plants.
  • Bracts with extrafloral nectaries adapted into variously shaped nectar -containers.
  • Terminal inflorescences: racemes, umbels or spikes.
  • Leaves alternate and simple with abaxialglands.
  • Fruit capsular with seeds embedded in fleshy pulp.
Other important characters
  • Usually climbing.
  • Often hemiepiphytic.
  • Youngest leaf enclosing the terminalbud.
  • Young growth often reddish coloured by anthocyanins.
  • Flowers with variously connate petals.
  • Stamens 3 or 5 or many.
Useful tips for generic identification

Key to genera of the Neotropical Marcgraviaceae 

1. Inflorescence umbellate or subumbellate ... 2
1. Inflorescence spicate or racemose ... 3

2. Inflorescence completely fertile; sepals and petals 5; petals free or variously connate; leaves spirally arranged ... Marcgraviastrum
2. Inflorescence partly sterile (central flowers aborted with only the bracteal nectaries developed); sepals and petals 4; petals calyptrately connate; leaves distichously arranged ... Marcgravia

3. Inflorescence spicate; nectaries attached to the rachis next to the flowers ... Sarcopera
3. Inflorescence racemose; nectaries variously attached to the pedicel... 4

4. Stamens 3-5, rarely 4 or 6-8; nectaries attached to the pedicel at the base of the calyx, rarely at some distance ... 5
4. Stamens (5-)12-35(-80); nectaries attached to the pedicel at various distances from the calyx, but never at its base ... 6

5. Ovary 2-locular; nectaries gibbose or somewhat leaf-like, solid or nearly so ... Ruyschia
5. Ovary 3-5-locular; nectaries tubular, hollow, often auriculate... Souroubea

6. Inflorescence an elongated raceme, 25-60(-85) cm; pedicels 2-5(-7) mm; nectaries adnate above the middle of the pedicel, never attached at the base ... Norantea
6. Inflorescence a short raceme, 4-25(-35) cm; pedicels (20-)30-70 mm; nectaries adnate below the middle of the pedicel, rarely at the base ... Schwartzia

Notable genera and distinguishing features
  • See first image: inflorescence structure.
Literature
Important literature

Bedell, H.G. 1985. A generic revision of Marcgraviaceae I. The Norantea complex. Ph.D. Dissertation (ined.), Univ. Maryland, College Park, MD.

Bedell, H.G. 1989. Marcgraviaceae. In: Howard, R.A. (ed.). Flora of the Lesser Antilles 5: 300-310.

Dressler, S. 2000. Marcgraviaceae. In: Flora de República de Cuba, Ser. A, Fasc. 5: 1-14.

Dressler, S. 2001. Marcgraviaceae. In: Steyermark, J.A., P.E. Berry, K. Yatskievych & B.K. Holst (eds.), Flora of the Venezuelan Guayana vol 6, pp. 248-260.  Missouri Botanical Garden, St. Louis.

Dressler, S. 2004. Marcgraviaceae. In: Kubitzki, K. (ed.). The Families and Genera of Vascular Plants. vol. 6, pp. 258-265. Springer-Verlag, Berlin.

Dressler, S. 2004. Marcgraviaceae, pp. 236-238. In: Smith, N.P., Heald, S.V., Henderson, A., Mori, S.A. & Stevenson, D.W. (eds.). Flowering Plants of the Neotropics. Princeton University Press/New York Botanical Garden Press.

Giraldo-Cañas, D. 2002. Los géneros Marcgraviastrum y Norantea (Marcgraviaceae) en Brasil. Revista Acad. Colomb. Ci. Exact. 26: 469-476.

Giraldo-Cañas, D. 2003. Revisíon de las especies colombianas del género Schwartzia (Marcgraviaceae). Caldasia 25: 1-21.

Giraldo-Cañas, D. 2005. Validation of a New Species of Schwartzia (Marcgraviaceae) and Synopsis of the genus for Ecuador. Novon 15: 123-127.

Giraldo-Cañas, D. & Fiaschi, P. 2005. Las Marcgraviaceae (Ericales) de Brasil: Las especies del complejo Norantea. Caldasia 27(2): 173-194.

Giraldo-Cañas, D. 2006. Lectotipificación para Schwartzia magnifica (complejo Norantea, Marcgraviaceae) y Revisión del género para Bolivia y Perú. Caldasia 28(2): 275-283.

Hammel, B.E. 2007. Marcgraviaceae. In: Hammel, B.E. et al. (eds.). Manual de Plantas de Costa Rica vol 6, pp. 374-391. Missouri Botanical Garden, St Louis.

Lens, F., Dressler, S., Jansen, S., van Evelghem, L. & Smets, E. 2005. Relationships within balsaminoid Ericales: a wood anatomical approach. American Journal of Botany 92(6): 941-953.

Lens, F., Dressler, S., Vinckier, S., Janssens, S., Dessein, S., Van Evelghem, L. & Smets, E. 2005. Palynological variation in balsaminoid Ericales. I. Marcgraviaceae. Ann. Bot. 96: 1047-1060.

Roon, A.C. de 1967. Foliar sclereids in the Marcgraviaceae. Acta Bot. Neerl. 15: 585-623.

Roon, A.C. de 1970. Flora of Panama, Fam. 121. Marcgraviaceae. Ann. Missouri Bot. Gart. 57: 29-50.

Roon, A.C. de 1975. Contributions towards a monograph of the Marcgraviaceae. Thesis. Utrecht.

Utley, J.F. 1976. A synopsis of the Mexican Marcgraviaceae. Brenesia 9: 51-59.

Utley, J.F. 2001. Marcgraviaceae. In: Stevens, D.W., C. Ulloa U., A. Pool & O.M. Montiel (eds.), Flora de Nicaragua vol 2, pp. 1335-1338. Missouri Botanical Garden, St Louis.

Images from La Selva Digital Flora Project http://sura.ots.ac.cr/local/florula3/index.htm

Images

Marcgraviaceae Bercht. & J.Presl appears in other Kew resources:

First published in Prir. Rostlin Aneb. Rostl. 218. 1820 [Jan-Apr 1820] (1820)

Accepted by

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

Sources

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