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non-core Caryophyllales


Pam Soltis, Doug Soltis, and Monica Arakaki
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taxon links [down<--]Caryophyllales Interpreting the tree
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Containing group: Caryophyllales


The non-core Caryophyllales clade (sensu Cuénoud et al. 2002), which corresponds to the Polygonales of Judd et al. (2002), has only recently been identified through molecular phylogenetic analysis. It includes several families that were previously classified in Cronquist’s (1981) Rosidae or Dilleniidae.


Possible synapomorphies for the non-core Caryophyllales (= Polygonales sensu Judd et al. 2002) are scattered secretory cells containing plumbagin, naphthaquinone (which has been lost in several clades), an indumentum of stalked, gland-headed hairs, basal placentation (with shifts to parietal in some Droseraceae and axile in Nepenthaceae), and starchy endosperm (Judd et al. 2002). Polygonaceae and Plumbaginaceae are united by ovaries with a single ovule and generally indehiscent fruits such as achenes or nuts. The clade of Droseraceae, Nepenthaceae, and relatives is probably supported by carnivory, circinate leaves, and pollen grains in tetrads (Judd et al. 2002).

Discussion of Phylogenetic Relationships

There are three major clades within non-core Caryophyllales:

  1. Polygonaceae (43 genera, 1,100 species; cosmopolitan) and Plumbaginaceae (22 genera, 800 species; cosmopolitan, but centered in the Mediterranean region) form a strongly supported clade, as they have in many previous analyses (e.g., Giannasi et al. 1992; Rettig et al. 1992).
  2. Tamaricaceae (5 genera, 90 species; Eurasia and Africa) and Frankeniaceae (monogeneric with 90 species; cosmopolitan)
  3. Carnivorous plants classified in Droseraceae (3 genera; 115 species; cosmopolitan), Drosophyllaceae (monospecific; Spain and Morocco), Dioncophyllaceae (3 monospecific genera; western Africa), and Nepenthaceae (monogeneric with 90 species; Madagascar to New Caledonia), together with Ancistrocladaceae (monogeneric with 12 species; tropical Africa to Borneo and Taiwan).

The relationship between Droseraceae and Nepenthaceae, and their relationship to Caryophyllidae sensu Cronquist (1981), was first recognized on the basis of rbcL sequences (Albert et al. 1992; Chase et al. 1993; Williams et al. 1994). Analyses of matK sequences alone for several species of Nepenthes, Drosera, and Ancistrocladus, plus Drosophyllum and all three genera of Dioncophyllaceae, resulted in a well-supported tree with Droseraceae sister to a clade of Nepenthaceae , (Drosophyllum , (Dioncophyllaceae , Ancistrocladaceae)) (Meimberg et al. 2000).

Other Names for non-core Caryophyllales


Albert, V. A., S. E. Williams, and M. W. Chase. 1992. Carnivorous plants: phylogeny and structural evolution. Science 257:1491-1495.

Chase, M., Soltis, D. and R. E. A. Olmstead. 1993. Phylogenetics of seed plants: an anlysis of nucleotide sequences from the plastid gene rbcL. Ann. Miss. Bot. Gard. 80: 526-580.

Cronquist. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York.

Cučnoud, P., Savolainen, V. Chatrou, L. W., Grayer, P. M., R. J., and M. Chase. 2002. Molecular

Giannasi, D. E., Zurawski, G., Learn, G. and M. T. Clegg. 1992. Evolutionary relationships of the Caryophyllidae based on comparative rbcL sequences. Systematic Botany 17: 1-15.

Judd, W. S., Campbell, C. S., Kellogg, E. A. and P. F. Stevens. 2002. Plant systematics: a phylogenetic approach. Sinauer Associates, Inc., Sunderland, Massachusetts.

Meimberg, H., P. Dittrich, G. Bringmann, J. Schlauer, and G. Heubl. 2000. Molecular phylogeny of Caryophyllidae s. l. based on matK sequences with special emphasis on carnivorous taxa. Plant Biology 2:218-228.

Rettig, J. H., Wilson, H. D. and J. M. Manhart. 1992. Phylogeny of the Caryophyllales--Gene sequence data. Taxon 41: 201-209.

Williams, S., Albert, V., and M. Chase. 1994. Relationships of Droseraceae: a cladistic analysis of rbcL sequence and morphological data. American Journal of Botany 81: 1027-1037.

Information on the Internet

Title Illustrations
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Scientific Name Polygonum bistorta
Copyright © 2004 Kurt Stüber
Scientific Name Drosera intermedia
Location coastal Mississippi, USA
Acknowledgements Photograph courtesy InsectImages.org (#1241028)
Specimen Condition Live Specimen
Source Collection Bugwood Network/Forestry Images
Copyright © James Henderson, Gulf South Research Corporation
Scientific Name Nepenthes rajah
Location Sabah, Malaysia
Specimen Condition Live Specimen
Identified By Ch'ien Lee
Life Cycle Stage Mature plant
Body Part Lower pitcher
Size c. 20 cm long
Copyright © 2006 Chien Lee
About This Page

Florida Museum of Natural History and the Genetics Institute, Gainesville, Florida, USA

Doug Soltis
Department of Botany and the Genetics Institute, Gainesville, Florida, USA

Monica Arakaki
University of Florida, Gainesville, Florida, USA

Correspondence regarding this page should be directed to Monica Arakaki at

Page: Tree of Life non-core Caryophyllales. Polygonales. Authored by Pam Soltis, Doug Soltis, and Monica Arakaki. The TEXT of this page is licensed under the Creative Commons Attribution License - Version 3.0. Note that images and other media featured on this page are each governed by their own license, and they may or may not be available for reuse. Click on an image or a media link to access the media data window, which provides the relevant licensing information. For the general terms and conditions of ToL material reuse and redistribution, please see the Tree of Life Copyright Policies.

Citing this page:

Soltis, Pam, Doug Soltis, and Monica Arakaki. 2006. non-core Caryophyllales. Polygonales. Version 07 June 2006 (under construction). http://tolweb.org/non-core_Caryophyllales/20968/2006.06.07 in The Tree of Life Web Project, http://tolweb.org/

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