Golden guinea vine Hibbertia scandens, copyright Casliber.

Belongs within: Gunneridae.

The Dilleniaceae are a pantropical and warm-temperate group of flowering plants, mostly trees and shrubs. Members of the family have toothed leaves with often strong and parallel secondary veins extending into the teeth (Angiosperm Phylogeny Website). Hibbertia, guinea flowers, is a genus of shrubs and climbers found in Australasia and Madagascar whose vernacular name refers to their production of radial yellow flowers.

The Dilleniaceae: tropical enigmas
Published 22 October 2012

In recent years, molecular analyses of often very large data sets have given us a reasonably good picture of the evolution of flowering plants, with most higher taxa settling down to reasonably comfortable positions. The subject of today’s post, however, is still something of a phylogenetic enigma.

Flower and opened fruit of the ‘red beech’, Dillenia alata, from here.

The Dilleniaceae are a family of about 500 species found mostly in the tropics, though one genus, Hibbertia, is also diverse in temperate Australia. Members of the family are very diverse in appearance: though the majority are trees or shrubs, some are lianes or even herbs. Dilleniaceae also show a remarkable diversity in features that are relatively stable in other families, such as floral symmetry and merosity (the number of flower organs such as stamens or carpels) (Horn 2009). Despite this diversity, Dilleniaceae are constant enough in other features that they have been recognised as a unified group since at least the 1800s. More questionable is their relation to other flowering plants: they are certainly members of the Pentapetalae, but the presence in some species of seemingly ‘basal’ characters (such as ladder-like perforation plates in the xylem and leaves with disorganised venation) lead some authors to regard them as an evolutionary link between the more basal magnoliids and a group of pentapetalous plants with centrifugal (starting from the centre and moving outwards) stamen development, called the Dilleniidae. ‘Dilleniids’ are now recognised as polyphyletic, including members of both the major clades Rosidae and Asteridae, but the Dilleniaceae themselves are not well resolved beyond basal Pentapetalae. Soltis et al. (2011) recently placed the Dilleniaceae as related to the clade of Asteridae + Caryophyllales + Santalales, but other analyses have placed them closer to Rosidae + Saxifragales, or even sister to all other Pentapetalae. Just to confuse matters, phylogenetic analysis within the Dilleniaceae suggests that at least some of their ‘primitive’ characters are in fact derived reversals of specific subtaxa (Horn 2009).

Erect guinea-bush Hibbertia riparia, photographed by Williewonker.

Relationships within the Dilleniaceae are perhaps better understood. The pantropical genus Tetracera was placed by horn (2009) as the sister to all other Dilleniaceae, which are divided between a strictly Neotropical clade (the Doliocarpoideae) and a strictly Old World clade. The Old World clade is in turn divided between two biogeographically distinct subclades, one centred in southern Asia (the Dillenioideae) and the mostly Australasian genus Hibbertia. Hibbertia and the Dillenioideae overlap only in northernmost Australia, southern New Guinea, Fiji and Madagascar (where both Dillenia and Hibbertia have representatives on the eastern side of the island). The Neotropical Doliocarpoideae are mostly lianes or scandent shrubs, with the only tree being the savannah species Curatella americana. The liane form is much rarer among Old World Dilleniaceae, most of which are trees or shrubs, though the small southern Asian genus Acrotrema contains rhizomatous herbs (and may be phylogenetically within the genus Dillenia). A group of succulent Australian species with photosynthetic stems, previously recognised as the genus Pachynema, have been reclassified by Horn (2009) as a derived subgroup of Hibbertia.

Hibbertia juncea, previously Pachynema junceum, photographed by Russell Cumming.
Systematics of Dilleniaceae

Characters (from Angiosperm Phylogeny Website): Trees and shrubs (occasionally lianes or perennial herbs); distinctive flavonols, myricetin, ellagic acid present; hairs more or less stellate and sclerified; primary stem with continuous vascular cylinder; cork cambium deep-seated; true tracheids present; raphides present, also common in wood; rays often broad; epidermis silicified; branching from previous flush; hairs unicellular; leaves spiral, lamina vernation conduplicate, surface often scabrid, margins toothed, secondary veins parallel, proceeding straight to the teeth, tertiary venation more or less scalariform, fine veins areolate, teeth with clear glandular expanded apex, base rather broad, stipules absent, long petiolar flanges usually present; pedicels articulated; flowers often yellow; sepals (3-)5(-20), large, petals (2-)5, usually crumpled in bud; androecium often asymmetric, anthers (2-)many, from a ring primordium or fasciculate, fascicles opposite sepals, supplied by trunk bundles, connective often well-developed, anthers basifixed, exodermis well developed; nectary absent; gynoecium with (1-3)4-8[-20] carpels, styluli long, stigmas capitate to punctate; ovules 1-many/carpel, apotropous, often campylotropous, micropyle zigzag or exostomal, outer integument 2(-3) cells across, inner integument 2-6 cells across, parietal tissue 6-14 cells across, nucellar cap ca. 2 cells across, chalazal area massive; calyx persistent in fruit; aril present, funicular, often laciniate, exotesta often fleshy, exotegmen with spiral or annular thickenings, endotegmen tanniniferous; zygote with distinctive wall and protrusions into the endosperm [“mantle”]; n = 4, 5, 8-10, 12, 13; germination phanerocotylar.

<==Dilleniaceae [Dilleniales, Dilleniinae]
    |--Burtonia Salisbury 1807FT93
    |--Delima [Delimoideae, Tetraceroideae]T-W89
    |    `--D. sarmentosa [=Tetracera sarmentosa]T-W89
    |--Wormia alataC16
    |--Dillenites microdentatus (Hollick) Berry 1916CBH93
    |--Curatella americanaCBH93, V-MR-M03
    |--Adrastaea salicifolia [=Hibbertia salicifolia]H90
    |--Dillenia [Dillenioideae]CD07
    |    |--D. alataR96
    |    |--D. excelsaP88
    |    |--D. grandifoliaK03
    |    |--D. indica Linnaeus 1753CD07
    |    |--D. pentagynaS02
    |    `--D. suffruticosaP88
         |--H. acerosaGK00
         |--H. acicularisH08
         |--H. acuminataH90
         |--H. amplexicaulisGK00
         |--H. asperaH90
         |--H. auriculifloraLK14
         |    |--H. a. ssp. auriculifloraLK14
         |    `--H. a. ssp. minorLK14
         |--H. axillibarbaG04b
         |--H. billardieriB96
         |    |--H. b. var. billardieriB96
         |    `--H. b. var. obovataB96
         |--H. bracteataH90
         |--H. calycinaH90
         |--H. carinataG04b
         |--H. ciliolataLK14
         |--H. circumdansH90
         |--H. cistifloraH90
         |--H. cistoideaH90
         |--H. commutataOS04
         |--H. covenyanaH90
         |--H. cuneiformisJK80
         |--H. cunninghamiiRL05
         |--H. dentataH87
         |--H. diffusaC08
         |    |--H. d. var. diffusaB96
         |    `--H. d. var. dilatataB96
         |--H. echiifoliaLK14
         |--H. elataH90
         |--H. empetrifoliaH90
         |--H. enerviaLK00
         |--H. exasperataG04b
         |--H. fasciculataMB08
         |    |--H. f. var. fasciculataMB08
         |    `--H. f. var. clavata Maiden & Betche 1908MB08
         |--H. fractiflexaLK14
         |    |--H. f. ssp. fractiflexaLK14
         |    `--H. f. ssp. filicaulisLK14
         |--H. glaberrimaEF04
         |--H. glomerataGK00
         |--H. gracilipesG04b
         |--H. hemignostaOS04
         |--H. hermanniifoliaH90
         |--H. hexandraH90
         |--H. hooglandiiLK14
         |--H. huegeliiRL05
         |--H. kaputarensisH90
         |--H. ledifoliaLK14
         |--H. lepidocalyxG04b
         |--H. lepidotaLK14
         |--H. linearisM65
         |--H. marginataH90
         |--H. microphyllaOS04
         |--H. monogynaH90
         |--H. nitidaH90
         |--H. oblongataLK14
         |    |--H. o. ssp. oblongataLK14
         |    `--H. o. ssp. brevifoliaLK14
         |--H. obtusifoliaC70 [=H. linearis var. obtusifoliaB96]
         |--H. orientalisLK14
         |--H. pedunculataH90
         |--H. perfoliataP05
         |--H. polystachyaGK00
         |--H. procumbensH90
         |--H. pulchraGK00
         |--H. pungensG04b
         |--H. racemosaGK00
         |--H. recurvifoliaOS04
         |--H. ripariaH90
         |--H. rostellataG04b
         |--H. rufaH90
         |--H. rupicolaOS04
         |--H. salignaH90
         |--H. scandensMM09
         |--H. sericeaH90
         |--H. serpyllifoliaC08
         |--H. silvestrisGK00
         |--H. spicataG04a
         |--H. stellarisJK80
         |--H. strictaMM09
         |--H. subvaginataGK00
         |--H. vaginataRL05
         |--H. vestitaC08
         |--H. villosaH90
         |--H. virgataH90
         `--H. volubilisH87

*Type species of generic name indicated


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[MB08] Maiden, J. H., & E. Betche. 1908. Notes from the Botanic Gardens, Sydney. No. 13. Proceedings of the Linnean Society of New South Wales 33: 304–319.

[M65] Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bulletin of the American Museum of Natural History 130: 1–362.

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[P05] Pinder, A. M. 2005. A review of biodiversity in wetlands with organic sediments on the Swan Coastal Plain, Western Australia, with an emphasis on aquatic invertebrates. Journal of the Royal Society of Western Australia 88 (3): 129–132.

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