EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline; primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway [ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, smaller than inner members, A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther, tapetum glandular, binucleate, microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions, pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA + C/PHYB + E gene pairs.

Possible apomorphies are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged.

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

VITALES + ROSIDS: anthers articulated [± dorsifixed, transition to filament narrow, connective thin].

ROSIDS: embryo long; genome duplication; chloroplast infA gene defunct, mitochondrial coxII.i3 intron 0.

MALVIDAE = [[GERANIALES + MYRTALES] [PICRAMNIALES [SAPINDALES [HUERTEALES [MALVALES + BRASSICALES]]]]]: ?

Evolution. Wang et al. (2009: penalized likelihood dates) suggested that rapid radiation with Malvidae occured (113-)107-83(-76) million years ago.

[GERANIALES + MYRTALES]: ellagic acid +; K persistent in fruit[!]

Phylogeny. Geranium, the only representative of the order studied, was sister to all other rosids except Vitaceae (Zhu et al. 2007, support weak). Their position is unstable in a rbcL analysis of all angiosperms (Hilu et al. 2003). In some earlier trees, Crossosoma (Crossosomatales) was also included or was nearby, see e.g. Morgan and Soltis (1993), Chase et al. (1993), while in Price and Palmer (1993: rbcL analysis) Biebersteinia (see Sapindales here) was still tentatively included in Geraniales. Savolainen et al. (2000a) found the order to be monophyletic, but with only 52% support (see also Savolainen et al. 2000b); Crossosomatales were still its sister group, but with still less support. For further discussion of the relationships of Geraniales, see the Crossosomatales page.

GERANIALES Dumortier  Main Tree, Synapomorphies.

Non-hydrolysable tannins 0 [but see Geraniaceae]; nodes also 3<:3<; leaf margins gland-toothed; inflorescence cymose; nectary outside A, A obdiplostemonous, G opposite petals, (outer integument largely dermal in origin), ?style, stigma dry; fruit dehiscence?; seed testal. - 5 families, 17 genera, 836 species.

Evolution. Stem group Geraniales are dated to 89-83 million years before present, crown group members 86 to 80 million years before present (Anderson et al. 2005), however, Wang et al. (2009: penalized likelihood dates) suggested potentially older dates, with the stem group originating 114-76 million years ago, while the crown group divergence began 109-62 million years ago.

Chemistry, Morphology, etc. Geraniaceae and Vivianiaceae have a layer of small hypodermal druse-containing cells in the calyx (Kenda 1956). Indeed Weigend (2006: 217) suggests that there might be "a close and possibly exclusive relationship between Geraniaceae and Ledocarpaceae (= Ledocarpaceae and Vivianaceae here)." He lists numerous characters suggesting such a relationship, and these will need to be critically evaluated. Thus he notes that in both the basal ovules in the ovary tend to develop into seeds, while Boesewinkel (1997) mentions that it is the upper ovules of Caesarea (Vivianaceae) develop into seeds, the nature of the campylotropy of the ovules differs, the fruits of Rhyncotheca (Ledocarpaceae) differ rather strongly from the superficially similar fruits of Geraniaceae, etc. (although both do have a column). Furthermore, Weigend (2006) suggests that the fruits of his Ledocarpaceae are septicidal or septifragal, while Boesewinkel (1997) describes them (apart from Rhyncotheca) as being loculicidal; in fact, the fruits of Balbisia, at least, show variation. Nevertheless, as Weigend's lists and the descriptions here suggest, there are a number of similarities between the two. Bortenschlager (1967) provides a comprehensive pollen survey of Geraniaceae in the old sense, i.e. including things like Dirachmaceae (Rosales), also Vivianaceae and Ledocarpaceae which, although no longer in Geraniaceae, at least are still in Geraniales. Details of the distribution of the rpl16 intron in the order are unclear (see Geranioideae: cf. Downie & Palmer 1992b).

Phylogeny. Geraniales are poorly understood, and although a smallish group, they are morphologically quite heterogeneous, which, coupled with uncertainty over their exact phylogenetic position, makes thinking of apomorphies difficult (Kubitzki 2006a). Savolainen et al. (2000a) found the order to be monophyletic, but with only 52% support (see also Savolainen et al. 2000b). The tree here is based on these and other (see above) earlier studies, although their sampling is inconsistent, etc.

Includes Francoaceae, Geraniaceae, Ledocarpaceae, Melianthaceae, Vivianiaceae.

Synonymy: Francoales Takhtajan, Ledocarpales Doweld, Melianthales Doweld - Geranianae Reveal - Geraniopsida Meisner

GERANIACEAE Jussieu, nom. cons.   Back to Geraniales

Herbs; hydrolysable tannins +; leaves spiral (opposite), pinnate, leaflets not articulated, 2ndary veins often palmate; inflorescence cymose, often pseudoumbellate; C contorted, nectaries opposite sepals, vascularised, pollen ?often starchy, ovules often campylotropous, micropyle zig-zag; K persistent, endotesta palisade, with light line, crystalliferous, much thickened, unlignified, exotegmen palisade, anticlinal walls sinuous.

7[list]/805 - 2 groups below. Temperate and warm temperate. [Photos - Collection]

1. Hypseocharis

Perennial acaulescent herbs; anatomy?; stipules 0; A 15, ca 12 ovules/carpel, style filform, stigma capitate, ?surface; fruit a loculicidal capsule (septicidal, almost with mericarps - H. tridentata), filaments also persistent; endosperm scanty, cotyledons spiral; n = ?

1/1-3. S.W. Andean South America.

Synonymy: Hypseocharitaceae Weddell

2. The Rest

(Shrubs); hairs glandular; cork epidermal; (vessel elements with scalariform perforation plates); rays often 0; nodes 3:3; stem often jointed; petiole bundles annular (medullary bundles +); cuticle waxes 0 (rodlets); leaves (opposite), also simple and lobed, conduplicate-plicate, (petiolar spines + - Sarcocaulon), stipules 2, often well developed, interpetiolar or cauline, colleters +; (flowers monosymmetric), K with nectariferous spurs, (adaxial nectary only, "adnate" to pedicel; nectary ± enclosed, on abaxial base of A), aristate, (C 4), (A 15 [=5 x [3]]; 2-5; 7; antepetalous whorl staminodial), filaments ± connate basally, (tapetum amoeboid), pollen trinucleate, (colpate), 40-140 µm long, G [(2-)5], 1-2 apical ovules/carpel, (micropyle exostomal; nucellar cap +), true style short, stout, hollow [?always], stigma lobed; fruit with upper part of ovary elongating [the "stylar" beak], septicidal, dividing into mericarps which curl upwards and separate from columella, whether or not seeds disperse separately; exotestal cells undistinguished, stellate, (endotegmen slightly lignified); endosperm 0, embryo green (Geranium, at least), curved, cotyledons accumbent or longitudinally folded; n = 4, 7-14, etc; inversions in cp DNA, group II intron between nad1 b/c exons, rpl2 and rpl16 introns all 0.

5/865: Geranium (430), Pelargonium (280: the "geranium" of the window sill, fl. monosymmetric, A 2-7, nectary single, in pedicel, either x = 11, chromosomes 1.5³ µm long or x = 9, chromosomes 1.5-3.0 µm long), Erodium (80: see Fiz et al. 2006 for a phylogeny; there has been substantial dispersal), Monsonia (40: inc. Sarcocaulon). Temperate and warm temperate, esp. southern Africa (Map: from van Steenis & van Balgooy 1966; Hultén 1971; Meusel et al. 1978; Aedo et al. 2005).

Synonymy: Erodiaceae Horaninow

• Geraniaceae are often herbs with jointed stems; the stipulate leaves have palmate venation or are palmately or pinnately compound. The inflorescence is cymose, and all parts of the flower apart from the carpels are free. The sepals are aristate, the androecium is obdiplostemonous, the carpels are usually beaked, there are few (1-2(-12)) ovules per carpel, and the embryo is curved.

Evolution. Geraniaceae (excl. Hypseocharis) are some 40 million years old (Wikström et al. 2001). Diversification of Pelargonium (and Monsonia) in South Africa seems to have occured ca 30-10 million years before present as aridification set in, while diversification of the Geranium-Erodium clade occured at roughly the same time in Eurasia and the Mediterranean, perhaps in response to climate change and mountain uplift there (Fiz et al. 2008). Recent estimates suggest divergence of the Cape fynbos clades of Pelargonium at some (13.6-)10.6(-3.7) million years ago, diversification in the succulent karoo starting somewhat before (Verboom et al. 2009).

Struck (1997) and Bakker et al. (1999, 2000, 2004, 2005) discuss the phylogeny and diversification of Pelargonium in the Cape region; there is striking vegetative and floral variation, and almost 100 species are geophytes (Procheŝ et al. 2006). Leaf shape and scent is notably variable; Jones et al. (2009) discuss foliar evolution. The two adaxial petals of Pelargonium alone may bear markings, rather unusual for flowering plants. Darwin (1859) noted that the central flower of a Pelargonium inflorescence might lose its adaxial markings and also its nectary. This would be expected of a peloric flower which has become "ventralised", i.e. the morphology of the abaxial sector of the flower extends to the adaxial sector, so the markings on the adaxial petals and the adaxial nectary disappear.

Palmer et al. (1987) noted extensive expansion of the inverted repeat in Pelargonium; the chloroplast genome is the largest of all flowering plants, largely because of the expansion, and it has undergone very extensive rearrangement (Chumley et al. 2006). Parkinson et al. (2005) and Bakker et al. (2006a) found very great increases in the rate of evolution of the mitochondrial gene nad1 throughout this part of the family, and especially in Pelargonium, but not in Hypseocharis (see also Palmer et al. 2000); this has been observed in other genes, as well (Guisinger et al. 2008).

Chemistry, Morphology, etc. Boesewinkel (1988) described the corolla of Hypseocharis as being often imbricate, while Weddell (1861: vol. 2: 289) illustrated it as being contorted (see also Takhtajan 1997). Monsonia may have nectaries axillary to the sepals or on the abaxial bases of the stamens (Aldasoro et al. 2001). When there are fifteen stamens, there are antepetalous stamen pairs, as is common in obdiplostemony (Ronse Decraene et al. 1996; see also Rama Devi 1991), or there may be five groups of three connate stamens (Aldasoro et al. 2001). Campylotropy is by inpushing of the inner integument (Albers & Van der Walt 2006 and references). There has been major movement of ribosomal protein and succinate dehydrogenase genes from the mitochondrion in Erodium (Adams & Palmer 2003).

Information on embryology is taken from L. Narayana (1970), on nectaries from Link (1994), on floral morphology from Erbar (1998), on the seed coat from Meisert et al. (2001), on Monsonia, perhaps paraphyletic, from Aldasoro et al. (2001), and general information from Albers and Van der Walt (2006). There is information on chemistry in references included in Lis-Balchin (2002).

Phylogeny. Hypseocharis is sister to the other members of the family (e.g. Price & Palmer 1993; bakker et al. 1998). The genus is in need of further study; if H. tridentata, with septicidal (and ventricidal) capsule dehiscence, is sister to the rest of the genus, simple optimisiation of fruit characters on the tree becomes interesting. Other relationships are [Pelargonium [Monsonia [Geranium + Erodium]]]; the position of the monotypic California is unclear(Fiz et al. 2008).

Classification. Whether or not Hypseocharis is recognised as a family is of little importance; this was an option in A.P.G. II (2003) and there are certainly phenetic differences between it and the Geraniaceae s. str., but a broad circumscription of the family is adopted here

Previous Relationships. Hypseocharis used to be included in Oxalidaceae (Hutchinson 1973; Cronquist 1981), but nectaries, testa anatomy, etc., place it unambiguously here (e.g. Boesewinkel 1988; Rama Devi 1991). Also, its leaflets are not strongly articulated with the petiole, as they are in Oxalidaceae.

Previous Relationships. Geraniaceae used to include taxa like Biebersteiniaceae (Sapindales) and Dirachmaceae (Rosales). Hypseocharis itself has been placed in a monotypic order, Hypseocharitales (Takhtajan 1997), although placed near Geraniales in the sequence.

[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]: ?

Melianthaceae + Francoaceae: at least traces of inulin +; leaves conduplicate, spiral, insertion rather broad; inflorescence terminal, racemose, sterile bract(s) at apex, bracteoles 0; G lobed, placentae intrusive, ovules in two rows, micropyle bistomal, style long; endosperm copious, embryo short.

Erstwhile Greyiaceae + Francoaceae make a strongly supported pair (Morgan & Soltis 1993; Price & Palmer 1993; Soltis & Soltis 1997). Obdiplostemony develops after the initiation of the androecium, which is initially diplostemonous. The nectaries are not vascularised. Neither family has myricetin, common in Saxifragaceae with which Francoa and Tetilla have often been associated or even included in the past.

Melianthaceae and Francoaceae are placed in a single family by Savolainen et al. (2000b); many of the characters given there may be plesiomorphic. A.P.G. II suggest as an option separating Melianthaceae and Francoaceae, an option which is followed here.

Information on seed anatomy is taken from Danilova (1996), for Melianthaceae, see also Corner (1976); Linder (2006) provides a general account for the two families under Melianthaceae.

MELIANTHACEAE Berchtold & J. Presl, nom. cons.   Back to Geraniales

Shrubs to trees; glucuronide triterpenoid saponins [Bersama], bufadienolides +; cork subepidermal to outside phloem; (largely phloic medullary bundles +); cambium storied; nodes 5:5, 9:9, etc.; petiole bundles annular (cortical or medullary bundles +); styloids or raphides +; cuticle waxes usu. 0; leaves odd-pinnate and leaflets articulated, or simple, hydathodes common, (margins entire), stipules single, intrapetiolar, or 2, ± cauline; (inflorescence axillary); flowers monosymmetric, resupinate or not [Greyia], K 5, or 2 connate + 3, adaxial spurred or not, C 4-5(-6), clawed, unequal, recurved, nectary large, adaxial or annular, A 4, opposite sepals (-8; connate basally; 10, uniseriate), anther endothecium not fibrous [Melianthus], pollen biaperturate, G [(3-)4-5(-7)], 1-5 apical or basal apotropous ovules/carpel, stigma punctate or slighly expanded and ridged, wet; fruit a loculicidal capsule (valves opening internally - Greyia); seed arillate or not, testa multiplicative [20+ layers], exotesta palisade, crystalliferous, outer wall thick, endotesta fibrous, radial walls thickened and lignified [Greyia], tegmen crushed; endosperm starchy, thick-walled [Bersama, Greyia], or with amyloid [Melianthus]; n = 16-18 (19).

3[list]/11. Africa (Map: modified from Culham et al. 2007). [Photo - Inflorescence, Inflorescence, Inflorescence.]

• Melianthaceae are at most small trees that may be recognised by their serrate leaves or leaflets, and large, toothed stipules. They have terminal racemes of large, more or less monosymmetric flowers with the parts (apart from the ovary and sometimes calyx) free; the nectary is outside the stamens.

Melianthus - spurred K, 2-5 ovules, smells nasty; Bersama - 1 erect ovule, may smell of mustard. Melianthus zenkeri has a shortly branched inflorescence and may lack stipules. Leaf insertion in at least some species of Bersama is not notably broad. Hutchinson (1973) described a disc lining the inside of the K. The number of nuclei in the pollen grain is unclear (Dahlgren & van Wyk 1988). For the ovules of Bersama, which may face away from each other, see Danilova (1996); what is the level of this character? In Bersama the fruits are partly septicidal as well. In Melianthaceae s. str. the tegmen is crushed and unlignified, but in Greyia the exotegmen and next layer is fibrous, but although the mesotegmic cells elongate, they are not persistent, there is also a massive hypostase.

In Greyia the leaf sheaths are adnate to the stem; the sheath, petiole and lamina are detached together, helped by the activity of a cambium. Stipules may occur rarely (Dahlgren & van Wyk 1988). Greyia also has a distinctive chemistry; the B-ring deoxyflavonoids found there are scattered in broad-leaved angiosperms (Bohm & Chan 1992).

For node and petiole anatomy, see Hilger (1978a, b), for general anatomy, see Gornall and Al-Shammary (1998), for floral development, see Ronse Decraene et al. (2001b - stipule morphology varies more than in the species studied there), for anthers, see Endress & Stumpf (1991), and for flower and fruit, see Doweld (2001a: cf. micropyle type). For Greyia in particular, further information is taken from Hideux and Ferguson (1976: pollen), Ramamonjiarisoa (1980: anatomy), Steyn et al. (1986), Steyn and van Wyk (1987), Nemirovich-Danchenko (1995: seed coat anatomy), Gregory (1998) and Ronse Decraene and Smets (1999: floral development).

Synonymy: Bersamaceae Doweld, Greyiaceae Hutchinson, nom. cons.

FRANCOACEAE A. Jussieu, nom. cons.   Back to Geraniales

Perennial herbs; flavonols, myricetin, ellagic acid 0, anthocyanin in roots, inulin?; cork ?; young stem with pseudosiphonostele; nodes 3:5; hairs uniseriate, secretory; leaves pinnate or simple, conduplicate-plicate (2ndary veins pinnate); inflorescence scapose; flowers poly- or monosymmetric; K 4-5, induplicate-valvate, C (3) 4-5, clawed or not, stamens = and opposite sepals, or 8, tapetum multinucleate [Francoa], pollen with complex endapertures, nectary lobed, G [4(-5)], deeply sulcate, ovules numerous, funicular obturator +, style short, stigmas sessile, commissural; fruit dehiscence variable; exotestal cells elongated, thickened, endotesta elongated, with thickened anticlinal walls, tegmen of pigmented cells; n = 26. chal. haust?

2/2. Chile. [Photo - Francoa Habit © Gardenweek.org]

Francoa: A 8, capsule with loculicidal dehiscence, Tetilla: A = K, septicidal dehiscence (both described as diplostemonous and septicidal in Linder 2006). The stem has an endodermoid layer.

For seed anatomy, see Nemirovich-Danchenko (1995), for vegetative anatomy of Francoaceae, see Gregory (1998), for embryology, Gaümann (1919).

Krach (1976) compared the testa anatomy with that of Cunoniaceae.

Vivianiaceae + Ledocarpaceae: (thorns +); ?nodes; rays 0; leaves opposite, simple, (margins entire), stipules 0, but a line across the stem; inflorescence terminal; K aristate, C contorted, pollen grains spherical, 23-40 µm long, polypantoporate, micropyle endostomal, apices of integuments 3-4 cells across, funicle with hairs, style short, styles/stigmas separate, long, stigma dry, margins ± revolute; fruit a loculicidal capsule; exotesta and endotegmen more or less tanniniferous; endosperm ± copious, walls thick, pitted, embryo curved.

The two are placed in a single family by Savolainen et al. (2000b), but the support is not strong. Boesewinkel (1997) describes ovules and seeds in the group, while Weigend (2005) details some aspects of floral morphology and pollination, and later (Weigend 2006) provides an account of the two families (under Ledocarpaceae, also with data on wood and leaf anatomy, inflorescence morphology, etc.). There is some conflict in the various accounts (see also introduction to Geraniales). Endosperm type/development?

VIVIANIACEAE Klotzsch   Back to Geraniales

Woody herb (annual); glandular hairs +; chemistry?; cork?, cambium storied; rays 0; nodes 1:1; cuticle waxes ± band-like or 0; leaves white-hairy below, 2ndary veins subpalmate; (flowers 4-merous); K valvate, basally connate, strongly 8- or 10-ribbed, (2-lobed) nectary glands alternating with C, (A 4, 5, 15), G [(2) 3], odd member abaxial, 2 superposed ovules/carpel; seed (hairy), raphe tanniniferous; (exotegmen thick-walled, elongated, not lignified - Caeserea); endosperm copious, initially with some starch, embryo green, root long; n = 7.

1 or 4 (Lefor 1975)[list]/6. Chile, S. Brasil (Map: from Lefor 1975).

The inflorescence may be represented all or in part by a branched thorn. The vessel elements may have a single bar across the perforation.

The exotestal seeds are unlike those in Geraniaceae s. str., but like that family the wood lacks rays and the floral anatomy of the two is similar. The stamens are much longer than the stigma/styles.

See Carlquist (1985b) for wood anatomy, Narayana and Rama Devi (1995) for general information, and M. S. Dunthorn (pers. comm.) for nodal anatomy.

LEDOCARPACEAE Meyen   Back to Geraniales

Shrubs; anatomy?; cuticle waxes as platelets; epicalyx + or 0; (K acute; C 0), (pollen inaperturate - Balbisia, Wendtia), nectary 0, G [3, 5], opposite sepals, 2 collateral or superposed-many ovules/carpel, nucellar cap +, stigmas drying dark; (fruit septicidal or more or less septifragal); (exotesta of slime cells - Balbisia); endosperm scanty (thin-walled), cotyledons spiral (straight); embryo colour?; n = 9.

3/12. W. South America, especially the Andes (Map: modified from Culham 2007).

The leaves can be deeply lobed. The fruits of Balbisia may be more or less septifragal (with a loculicidal slit, too) or clearly loculicidal.

Rhynchotheca is a morphologically very distinctive genus: C 0, G opposite sepals, stigma wet; fruit a septicidal, pointed, long-ovoid capsule with columella; endosperm cells thin-walled, embryo straight. It is possibly wind pollinated. Note that the fruits lack a true beak, unlike the apparently rather similar fruits of Geraniaceae.

Wendtia is sister to Balbisia (G opposite sepals); the pair are sister to Greyiaceae + Francoaceae (Price & Palmer 1993).

For carpel orientation, see Baillon (1874) and Knuth (1931).

Synonymy: Rhyncothecaceae Endlicher .