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, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem complex; 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(-8) mm/mm²; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores] +, 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, 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 sieve plate, companion cells from same mother cell that gave rise to the tube, the sieve tube with P-proteins; nodes unilacunar; stomata with ends of guard cells level with aperture, paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein 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, exine 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 ?type, stylulus short, hollow, stigma ± decurrent, wet [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; 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/PHYCgene 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. Furthermore, 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; nucleus of egg cell sister to one of the polar nuclei; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

See the Dilleniales page for discussion on the relationships of these groups, which have no firm position as yet, although it is increasingly likely that Carophyllales are close to the asterids; see also the Dilleniales Dilleniales page.

CARYOPHYLLALES Perleb  Main Tree, Synapomorphies.

(Odd ecology and/or physiology); not mycorrhizal; root hair cells in vertical files [sampling!]; (tracheids +); perforation plates not bordered; only alternate vascular pitting; scanty vasicentric parenchyma; both uni- and multiseriate rays present; leaves entire; anther with outer parietal cells developing directly into the endothecium, pollen colpate, tectum spinulose, G [3], when G = K or P, opposite them, micropyle endostomal, style branch long; fruit a loculicidal capsule; seed testal; embryo long. - 33 families, 692 genera, 11155 species.

Caryophyllales contain ca 6.3% of eudicot diversity (Magallön et al. 1999) and may date to the Albian, 111-104 million years before present, although Rhabdodendraceae do not split off until 90-83 million years before present (Wikström et al. 2001); Anderson et al. (2005: Rhabdodendraceae also included) suggests figures of 116-114 million years before present for stem group Caryophyllales, 102-99 million years before present for the crown group. The Droseraceae et al. and Simmondsiaceae et al. clade may have diverged 90-83 million years before present, the two main groups in it in turn diverging 82-76 million years before present (Wikström et al. 2001). However, if Rhabdodendraceae are sister to the Simmondsiaceae et al. clade (see below)... Some chrysomelid beetles - Alticinae, Cassidinae-Hispinae - seem notably more common on this clade than others (Jolivet & Hawkeswood 1995).

The shikimic acid pathway, particularly phenyalanine, is a starting point for the synthesis of nitrogen-containing benzylisoquinoline alkaloids and the betalains of core Caryophyllales; Kubitzki (1994) suggested a relationship between core Caryophyllales, Magnoliidae and monocots because all contained such compounds.

Hilu et al. (2003: matK analysis alone) suggest that Caryophyllales are sister to Asterids, a relationship that has been found in some other studies (e.g. Soltis et al. 1997, cf. also Nandi et al. 1998). A relationship between Caryophyllales and Dilleniales has also been suggested (D. Soltis et al. 2003a), see below for more details. However, Caryophyllales alone seem to be sisterto asterids, although the support is only moderate; see the Dilleniales page for further discussion.

Cuénoud (2002a, b) summarises variation in the order. Given the variation in carpel number in the clade, it is with some hesitation that three carpels is suggested as the plesiomorphic condition. There are many other unusual characters here, but their phylogenetic significance is unclear, partly because of sampling problems; e.g. knowledge of anther wall development is poor (Dahlgren & Clifford 1982). Furthermore, Rhabdodendraceae and members of the basal pectinations in both major clades are poorly known. Many families are tolerant of saline/arid conditions and/or have a distinctive habit (e.g. climbers with distinctive grapnel organs) or physiology (carnivory, C₄ pathway, CAM) or both. It is interesting that Landis et al. (2002) found that both Polygonales and Caryophyllales (here just the one order) commonly lack mycorrhizae, although there are some exceptions (e.g. some Nyctaginaceae and Amaranthaceae). Isoflavonoids are scattered in the group (Mackova et al. 2006, perhaps especially in the core Caryophyllales). Flavonol sulphates occur in Plumbaginaceae, Polygonaceae, and Amaranthaceae (Chenopodiaceae s. str.), and sulphated betalains in Phytolaccaceae. Placement of several features of wood anatomy on the tree need checking (cf. Carlquist 2002b, 2003a). Non-bordered perforation plates may be a synapomorphy for Caryophyllales or Caryophyllales and Santalales (Carlquist 2000a). Anomalous secondary thickening by successive cambia is widespread, as are maximally biseriate rays (the latter inc. Asteropeiaceae, but not core Caryophyllales - Nandi et al. 1998). Similarities in sieve tube plastids between Triplarieae (Polygonaceae) and core Caryophyllales are here treated as parallelisms (cf. Judd & Olmstead 2004). The outer stamens are often initiated in pairs, especially in core Caryophyllales, but also elsewhere in the order (Ronse Decraene & Smets 1993); C and adjacent (antepetalous) A are developmental units in Plumbaginaceae and Caryophyllaceae (Friedrich 1956; Ronse Decraene et al. 1998). Trinucleate pollen is common. Carpels open in development are known both from Polygonaceae and core Caryophyllales (Tucker & Kantz 2001). The rpl23 gene is a pseudogene in the few Caryophyllales examined (Logacheva et al. 2008). Purple-spored smuts and Uromyces rusts parasitize several families, including Plumbaginaceae, Polygonaceae and core Caryophyllales (Savile 1979b: he considered this to be a strong sign that the groups were close).

Rhabdodendraceae are sister to all other Caryophyllales in the rbcL analysis of Fay et al. (1997b) and in the Bayesian analysis of Soltis et al. (2007a). Cuénoud et al. (2002) found that Simmondsia was grouped with Rhabdodendron in a matK analysis, but with only weak support, but in two- and four-gene analyses (with poorer sampling) it was associated with core Caryophyllales; in trees shown by Drysdale et al. (2007) and Brockington et al. (2007) a position of Rhabdodendron as sister to the rest of core Caryophyllales was again found in most analyses, and is adopted here. Hilu et al. (2003: matK analysis alone) also suggested relationships between Rhabdodendraceae and this part of the tree, however, there were basically two main groups within Caryophyllales, the core Caryophyllales (with associated families) and Polygonaceae, etc. This latter clade, including Plumbaginaceae, Polygonaceae, Nepenthaceae, Droseraceae and Drosophyllaceae, is well-supported (Morton et al. 1997b), and has been expanded to include four carnivorous families (see also Albert et al. 1992; Meimberg et al. 2000; Cuénoud et al. 2002; Cameron et al. 2002) and other families with distinctive vegetative morphologies (see also Heubl et al. 2006). Asteropeiaceae and Physenaceae form a well supported pair, in turn showing a well-supported sister group relationship to core Caryophyllales (Källersjö et al. 1998), or Asteropeiaceae and Simmondsiaceae (the only two included) successively sister groups to the core (D. Soltis et al. 2000). The tree below is based largely on those presented by Meimberg et al. (2000), Cameron et al. (2002: 4 genes) and Cuénoud et al. (2002: matK alone), for the position of Achatocarpaceae, see Müller and Borsch (2005).

Takhtajan's (1997) Plumbaginanae are monotypic; Nepenthanae included Droseraceae and some other Caryophyllales, but also families now in Ericales, etc.

Includes Achatocarpaceae, Aizoaceae, Amaranthaceae, Anacampseros, etc., Ancistrocladaceae, Asteropeiaceae, Barbeuiaceae, Basellaceae, Cactaceae, Caryophyllaceae, Didiereaceae, Dioncophyllaceae, Droseraceae, Drosophyllaceae, Frankeniaceae, Giseckiaceae, Halophytaceae, Limeaceae, Lophiocarpaceae, Molluginaceae, Montiaceae, Nepenthaceae, Nyctaginaceae, Physenaceae, Phytolaccaceae, Plumbaginaceae, Polygonaceae, Portulacaceae, Rhabdodendraceae, Sarcobataceae, Simmondsiaceae, Stegnospermataceae, Talinaceae, Tamaricaceae.

Synonymy: Aizoineae Doweld, Basellineae Doweld, Cactineae Bessey, Caryophyllineae Bessey, Chenopodiineae Engler, Nyctaginineae Doweld, Phytolaccineae Engler, Portulacineae Doweld, Stegnospermatineae Doweld - Aizoales C. Y. Wu et al., Amaranthales Dumortier, Ancistrocladales Reveal, Atriplicales Horaninow, Cactales Dumortier, Chenopodiales Dumortier, Dioncophyllales Reveal, Droserales Grisebach, Nepenthales Dumortier, Nyctaginales Dumortier, Opuntiales Willkom, Petiveriales Lindley, Phytolaccales Doweld, Plumbaginales Lindley, Polygonales Dumortier, Portulacales Dumortier, Rhabdodendrales Doweld, Scleranthales Dumortier, Silenales Lindley, Simmondsiales Reveal, Stellariales Dumortier, Tamaricales Hutchinson - Caryophyllanae Takhtajan, Nepenthanae Reveal, Plumbaginanae Reveal, Polygonanae Reveal, Rhabdodendranae Doweld, Simmondsianae Doweld - Caryophyllidae Takhtajan, Plumbaginidae C. Y. Wu, Polygonidae C. Y. Wu - Amaranthopsida Horaninov, Cactopsida Brogniart, Caryophyllopsida Bartling, Opuntiopsida Endlicher, Polygonopsida Brongniart, Plumbaginopsida Endlicher

Droseraceae group + Tamaricaceae + Polygonaceae groups: acetogenic napthoquinones [plumbagin - in Plumbaginaceae, Droseraceae, Nepenthaceae, Dioncophyllaceae, related compounds in Polygonaceae] +; endosperm starchy.

It is unclear where the character of starchy endosperm is to be put on the tree. The condition is not known for the clades just below core Caryophyllales, and while Netolitzky (1926) noted that the core Caryophyllales lack starchy endosperm, and starch was not recorded from the thin endosperm found in the seeds of some Amaranthaceae (Shepherd et al. 2005b), Narayana and Lodha (1963) found starch in the young endosperm of Orygia (and Corbichonia: Lophiocarpaceae). In the Flora of China, several families of core Caryophyllales are reported to have starchy endosperm (e.g. Dequan & Gilbert 2003), but this must be a mistake.

For napthoquinones, particularly plumbagin, see Culham and Gornall (1994) and Ková&ath;&ik et al. (2006).

Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]]]: plants carnivorous; vascularised multicellular stalked or sessile glands; inflorescence ± cymose; C contorted, anthers extrorse, ovary unilocular.

Williams et al. (1994) drew atttention to possible relationships between Dioncophyllaceae and Drosophyllum, and Drosophyllum and Nepenthaceae were also found to be weakly associated (Morton et al. 1997b). For detailed relationships of this group, see Meimberg et al. (2000) and Cameron et al. (2002); the acquisition of carnivory may have happened once here, or it occured once and then was lost, perhaps more likely given the topologies found: Schlauer (1997) also discusses the evolution of carnivory. For a synapomorphy scheme for the whole group, see in part Albert and Stevenson (1996) and Meimberg et al. (2000: the floral characters listed are mostly plesiomorphies), but especially Heubl et al. (2006). The latter suggest that fly-paper traps are the plesiomorphic condition for the group, but note that where features like this or the possession of circinate leaves and pollen tetrads are placed on the tree will depend on the mode of character optimisation (e.g. ACCTRAN, DELTRAN) used. Heubl and Wistuba (1997) suggested that both Droseraceae and Nepenthaceae had ploidy levels of 8 or 16, based on x = 5 or thereabouts.

For acetogenic quinones and alkaloids, see Hegnauer (1986), Bringmann and Pokorny (1995) and Bentley (1998). For carnivory, see Lloyd (1942) and Juniper et al. (1989).

DROSERACEAE Salisbury, nom. cons.   Back to Caryophyllales

Insectivorous rosette herbs; flavonols, ellagic acid +; cork?; young stem with separate bundles, medullary rays broad; cambium 0; (medullary bundles +); nodes 1:1; petiole bundles various; stomata also tetracytic or actinocytic; mucilage hairs with xylem only; leaves adaxially circinate, glandular, stipules intrapetiolar, ?cauline, or 0; inflorescence terminal; K often connate at base, C ± marcescent, stamens = and opposite sepals (-15 - Dionaea), (introrse), (connective expanded), pollen in tetrads, bi- or trinucleate, intectate, 3-multiporate, operculate or not, G [3(-5)], median member abaxial, placentation parietal (basal - Dionaea), 3-many ovules/carpel, parietal tissue often absent, nucellar endothelium +, styles +, often bifid, (style + - Dionaea), stigmas expanded, papillate; (fruit indehiscent); exotesta palisade or not (endotesta with U thickenings), endotegmic cells small, ± sclerotic, or mucilaginous; endosperm nuclear, (embryo short); (germination cryptocotylar); n = 5<, chromosomes 1.5³ µm (<6µm [Dionaea]); chloroplast rpl2 intron 0 [one species!].

3[list]/115: Drosera (110). World-wide (Map: from Hultén 1971; George 1982; Correa A. & Silva 2005). [Photos - Collection, Collection.]

• Droseraceae are insectivorous herbs usually with adaxially coiled or folded leaves and moderate-sized flowers; there are three carpels usually terminated by branched styles.

The beginning of diversification within Drosera may date to ca 42 million years before present, although a pre-continental drift origin has also been suggested (Yesson & Culham 2006 and references). Drosera is exceptionally diverse in SW Australia, which has about one third of the species in the whole genus; diversification may be linked to the onset of the Mediterranean climate there some 15-10 million years before present.

Aldrovandra and Dionaea (both n = 6) may be sister taxa. Both have snap-traps, multicellular trigger hairs, etc. (Cameron et al. 2002; Rivadavia et al. 2003: little support for the relationship). The traps of Dionaea close in about 100 ms, the movement being aided by the leaf blades changing from concave to convex (Forterre et al. 2005); Volkov et al. (2008 and references) provide physiological details of the mechanisms involved.

Some Droseraceae have tenuinucellate ovules; the anthers are extrorse (Le Maout & Decaisne 1868; Baillon 1887).

See Hegnauer (1966, 1989) for chemistry, Boesewinkel (1989) for ovule and seed anatomy, Hoshi and Kondo (1998) for chromosomes, Cutler and Gregory (1998) for general anatomy, Conran et al. (2007) for gland morphology, Williams et al. (1994) and especially Cameron et al. (2002) and Rivadavia et al. (2003: focus on Drosera) for phylogeny, and Kubitzki (2002d) and the Carnivorous Plants Database for general information. Metcalfe and Chalk (1950) describe remarkable vascular patterns in the inflorescence axis and petiole.

Synonymy: Aldrovandaceae Nakai, Dionaeaceae Rafinesque

Nepenthaceae [Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]]: fibriform vessel elements +; rays 1-2 cells wide; cortical bundles in stem; anthers basifixed.

Heubl et al. (2006) place a character, "petiole with cortical vascular bundles" at this node.

NEPENTHACEAE Berchtold & J. Presl, nom. cons.   Back to Caryophyllales

Insectivorous, usu. viny; flavonols +, ellagic acid 0; cork pericyclic; (also medullary bundles +); (vessel elements with scalariform perforation plates); true tracheids +; large spirally-thickened cells in pith, pericycle, etc.; nodes 5-9:5-9; peltate glands +; leaves abaxially circinate, with involute blade and twining unbranched tendril terminated by pitchers, base broad; plant dioecious, inflorescence a raceme, bracts and bracteoles 0; P (3-)4, decussate, with large flat nectariferous glands adaxially, staminate flowers: A (4-)8-25, connate into a central column, pollen in tetrads, trinucleate, apertures indistinct, pistillode 0; carpellate flowers: staminodes?, G [(3-)4(-6)], placentation axile, many ovules/carpel, style short, stigma single, broad, papillate; seeds numerous, minute, exotesta with much thickened inner walls; endosperm +, nuclear; n = 40.

1[list]/90. Madagascar to New Caledonia (Map: see Meimberg & Heubl 2006). [Photo - Leaf; Collection.]

• Nepenthaceae are insectivorous plants that are easy to recognise because of the lidded pitchers borne on the end of a twining prolongation of the leaf. The leaf base itself is broad, further widening to form a laminar portion that then narrows to form the twining portion. The plants are dioecious, the inflorescences are racemes, the flowers are rather inconspicuous, and the seeds are very small.

The expanded part of leaf is developed from the leaf base, as in many monocots, the pitcher from the rest. How insects are trapped in the pitchers has long been unclear. Recent work suggests that the rim (peristome) of the pitcher is extremely wettable, and insects may aquaplane when they step on it, falling in to the pitcher below where they die and get digested; only when the rim is dry can insects walk on it easily, and then they may get trapped when they walk on to the wax-covered inner pitcher walls (Bohn & Federle 2004). Interestingly, the ant Camponotus schmitzi lives in close association with Nepenthes bicalcarata, and it can run across even the wetted rim. For the fauna of the pitchers, see Kitching (2000), while Pavlovic et al. (2007) discuss the physiology of lamina and trap.

The outer integument develops greatly after fertilisation and forms an exostome (Goebel 1933); there is a hair-pin bundle in the testa (Takhtajan 1988). The parietal cell in the ovule does not divide further.

For the biogeography and relationships of the group, see Meimberg and Heubl (2006). Some analyses suggest that the Malesian Nepenthes (including species from New Caledonia and Australia) are derived from a stock represented by the extant extra-Malesian taxa, but different relationships are suggested by different genes.

For general information, see Cheek and Jebb (2001: almost a monograph), Kubitzki (2002d) and the Carnivorous Plants Database, for chemistry, see Hegnauer (1966, 1990), for anatomy, Pant and Bhatnagar (1977), for the fauna of the pitchers, see Kitching (2000), and for the trapping of insects, see Bohn and Federle (2004: aquaplaning common).

Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]: ?

DROSOPHYLLACEAE Chrtek, Slavíkovà & Studnicka   Back to Caryophyllales

Small insectivorous woody plants; chemistry?; cork superficial; cortical bundles inverted; ?nodes; ?stomata; petiole bundles three, arcuate, inverted; mucilage hairs with xylem and phloem; leaves linear, abaxially circinate, with stalked glands in lines; flowers large, C contorted, ± marcescent, A 10, attachment?, pollen trinucleate, tectate, pantoporate, G [5], opposite the K, placentation basal, styles +, stigmas capitate; fruit septicidal; seeds operculate, exotesta not palisade, endotesta crystalliferous, with U thickenings, exotegmen thick-walled; endosperm ?, embryo short; germination cryptocotylar; n = 6, chromosomes ³15 µm long.

1/1: Drosophyllum lusitanicum. Southern Iberian Peninsula, Morocco (Map: from Ortega et al. 1995). [Photos - Collection]

• Drosophyllaceae are subwoody carnivorous plants that may be recognised by their long, abaxially coiled leaves that have sticky, glandular hairs in long lines on their abaxial surfaces; these hairs are not sensitive. The flowers are relatively large and short-lived (they remain open only a single day), but the petals dry and persist around the capsule.

Stem/leaf anatomy would repay investigation; both the cortical and petiole bundles appear to be inverted (Metcalfe & Chalk 1950, as Droseraceae). The flowers are relatively large; the stamens opposite the calyx are longest. Dehiscence of the fruit is down the ribs of the capsule and the valves are opposite the calyx.

Drosophyllaceae are sister to Dioncophyllaceae + Ancistrocladaceae, with good support, in an analysis of matK sequences, the position of Nepenthaceae being uncertain (Cuénoud et al. 2002).

For ovule and seed, see Boesewinkel (1989), and for general information, see Kubitzki (2002d) and the Carnivorous Plants Database.

Ancistrocladaceae + Dioncophyllaceae: lianes; (acetogenic napthyl isoquinoline alkaloids +); cortical bundles 0; petiole with sclerenchyma bundles; stomata actinocyclic; A introrse.

For the distinctive napthyl isoquinoline alkaloids, see Bringmann (1986), Bringmann and Pokorny (1995), and Bringmann et al. (2008, and references); they are synthesised from polyketide precursors, not from aromatic amino acids.

ANCISTROCLADACEAE Walpers, nom. cons.   Back to Caryophyllales

Plant twining, or with grapnels along one side of the branch opposite the leaves; myricetin +, ellagic acid?; cork mid-cortical; nodes 3:3; petiole bundle annular and with a peripheral ring of small bundles; (stomata actinocyclic); leaves supervolute, with surface glands; flowers small, pedicels articulated; C basally connate, A (5) 10, whorl opposite petals larger, filaments ± connate basally and adnate to C, G [3(-4)], half or more inferior, with 1 basal hemitropous ovule, micropyle?, styles articulated with apex of ovary, stigmas hippocrepiform or pinnatifid, ?type; fruit a nut, K much enlarged; seed ruminate, exotestal?; endosperm cellular, cotyledons much folded; n = ?

1[list]/12. Tropical Africa to W. Borneo and Formosa (Map: from van Steenis 1949a; Freson 1967; C. Taylor, pers. comm.). [Photo - Fruits] [Photo - Grapnels]

• Ancistrocladaceae are lianes that can be recognised by their shortly petiolate, extsipulate, entire leaves with glands in pits on the abaxial surface, and the complex, branched stem grapnels that are often opposite the leaves.

The pollen is like that of Dioncophyllaceae. Minute stipules and four carpels are reported by Takhtajan (1997) and Porembski (2002). 1/3 species tested had fluorescing wood.

In the past Ancistrocladaceae have often been included in Theales or Theanae (Cronquist 1981; Takhtajan 1997).

For chemistry, see Hegnauer (1989) and for general information, see Porembski (2002).

DIONCOPHYLLACEAE Airy Shaw, nom. cons.   Back to Caryophyllales

Insectivorous lianes or shrubs with anomalous secondary growth; cyclopentenoid cyanogenic glycosides +, ellagic acid?; cork?; xylem with included phloem; nodes ?; cortical bundles inverted; petiole bundle arcuate; glandular hairs +; leaves abaxially circinate, first leaves with stalked glands, later leaves with paired, recurved hooks; K valvate or open, A 10-30, G [2, 5], placentation parietal, many ovules/carpel, (short style +), stigmas punctate, capitate or feathery; capsule opening before maturity; seeds flattened, on elongated funicles, winged or not, ?ruminate; coat ?; endosperm ?nuclear; n = 12, 18 [both Triphyophyllum peltatum]; germination epigeal, cryptocotylar.

3[list]/3. Tropical West Africa (Map: only approximate).

• Dioncophyllaceae are distinctive in their rather long rosette leaves with closely parallel venation, these are followed by leaves with a short blade and glandular hairs on the abaxial surface of the prolonged midrib, and then, in the climbing phase, by leaves with paired grapnel hooks at the apex. The fruits open early and expose the still-developing flattened seeds.

Dioncophyllales were included in Theanae by Takhtajan (1997).

For morphology, see Gottwald and Parameswaran (1968) and Schmid (1964), for chemistry, Hegnauer (1966, as Flacourtiaceae, 1989) and Spencer and Siegler (1985), for carnivory, Bringmann et al. (2001), and for general information, see Porembski and Barthlott (2002) and the Carnivorous Plants Database.

[Frankeniaceae + Tamaricaceae] [Plumbaginaceae + Polygonaceae]: vessel elements with minute lateral wall pits +; sulphated flavonols, ellagic acid +; seed exotestal.

Frankeniaceae, Tamaricaceae and Plumbaginaceae all have flat, multicellular glands of subepidermal origin (Conran et al. 2007). This is perhaps an apomorphy here (or still higher), with a loss in Polygonaceae.

Frankeniaceae + Tamaricaceae: halophytic; bisulphated flavonols +, myricetin 0; wood storied; nodes ?; ?SiO₂ bodies +; (stomatal orientation transverse); leaves small, with salt-excreting glands; flowers small, 4-6-merous, C with basal adaxial appendages, pollen not spinulose, G with median member abaxial, placentation (basal; intrusive-)parietal; fruit a capsule; exotestal cells bulging or as hairs; endosperm +.

The monophyly of the two families and their sister-group relationship have recently been confirmed by Gaskin et al. (2004). It is equally parsimonious to assume that petal appendages are apomorphies for the family pair as it is to assume that they have evolved independently; in Tamaricaceae the Reamuria clade, members of which have these appendages, is sister to the rest of the family. Seeds with copious endosperm have the same distribution.

Tamaricaceae were placed in Violanae by Takhtajan (1997), probably because of their parietal placentation.

FRANKENIACEAE Gray, nom. cons.   Back to Caryophyllales

Herbs to shrubs; cork pericyclic or subepidermal; fibriform vessel elements +; rays 0; cuticle wax crystalloids 0; leaves opposite, often ericoid; flowers also 7-merous, K connate, lobes induplicate-valvate, C clawed, A (3-)6(-24; inner whorl staminodial), slightly connate at the base or not, versatile, tapetal cells binucleate, pollen trinucleate, G [(2-)3(-4)], (1-)2-6(-many) tenuinucellate ovules/carpel, nucellar cap +, funicles long, style +, stigma surface branched; exotestal cells large, papillate, papillae with terminal nail-like thickenings, endotestal cells thin-walled [?fibers], endotegmen with thick cuticle, tanniniferous; n = 10, 15.

1/90: Frankenia. ± World-wide in warm, dry areas, but scattered (Map: from Surgis 1921; George 1982; Whalen 1987; FloraBase 2004). [Photos - Collection]

There are no medullary rays. The endosperm has a coenocytic micropylar haustorium.

Some information is taken from Walia and Kapil (1965), Whalen (1987: taxonomy Old World Frankenia) and Olson et al. (2003: anatomy); for a general account, see Kubitzki (2002d).

TAMARICACEAE Link, nom. cons.   Back to Caryophyllales

Woody, also xeromorphic; (gypsum crystals +); cuticle waxes as tubes or curled rodlets; leaf bases often broad; inflorescence racemose (flowers solitary), bracteoles 0; K connate below or not, stamens = or 2x C or more, most connate at base into 5 bundles, borne with C on nectary (or nectary inside or outside A, or 0), anthers variously attached, G [(2-)3-4(-5)], opposite petals, ovules 2-many/carpel, embryo sac tetrasporic [a variety of types, even in one species, often 16-nucleate bipolar], styles +, (style +, short), stigmas usu. expanded, wet; seeds with hairs, exotestal cells periclinally elongated and thick-walled, endotestal cells thin-walled, crystalliferous; endosperm usu. scanty, oil and protein as reserves, perisperm common, thin; n = (11) 12.

5[list]/90: Tamarix (55). Eurasia and Africa, esp. Mediterranean to Central Asia, commonly naturalised in North America (map: from Hultén & Fries 1986; Meusel et al. 1978). [Photos - Collection]

• Tamaricaceae are woody plants with small leaves and flowers; the styles are long and are terminated by distinctive expanded stigmas. The capsular fruits release hairy seeds.

Relationships within the family are [[Holachna + Reamuria] [Myricaria + Tamarix]] (Gaskin et al. 2004).

The nucellus is very thin, the parietal cell not dividing.

See Hegnauer (1973, 1990) for chemistry, Czaja (1978) for seed storage, Zhang et al. (2001) for pollen and Gaskin (2002) for a general account.

Synonymy: Reamuriaceae Lindley (single terminal flowers, C contorted, basal adaxial scale on the petals, c.f. Frankeniaceae, many centrifugal stamens from 10 primordia, nectary 0, ovules ascending, stigma not expanded; endosperm + - Ronse Decraene 1990).

Plumbaginaceae + Polygonaceae: plants herbaceous; O-methylflavonols, myricetin, quinones +; (wood storeyed); successive cambia 0; cortical and/or medullary vascular bundles +; nodes 3:3; leaf bases broad; pollen usu. starchy, G with median member adaxial, 1 loculus and 1 basal ovule; fruit surrounded by accrescent calyx which forms part of the dispersal unit; exotesta ± persistent, otherwise seed coat undistinguished; mitochondrial coxII.i3 intron 0.

Lycaeninae caterpillars are quite commonly found on this group, probably because of the polyphenolics in their leaves (Fielder 1995).

For sterol composition in comparison to that of core Caryophyllales, see Wolfe et al. (1989), for anatomy, see Carlquist and Boggs (1996).

PLUMBAGINACEAE Jussieu, nom. cons.   Back to Caryophyllales

Often salt tolerant; glycine betaine, choline-O-sulphate +, little oxalate accumulation; vascularized mucilage glands and epidermal glands +; cork subepidermal or cortical; secondary thickening odd; rays multiseriate; petiole bundles arcuate; cuticle wax crystalloids 0 (irregular platelets); stomata also paracytic; K connate, ribbed, C contorted, stamens = and opposite petals, pollen with irregular columellae, tectum continuous, itself with columellae, with rather coarse blunt spines, nectary often on adaxial side of filament bases, G [5], ovule anatropous, funicle long and curled, obturator from wall at apex of ovary, embryo sac tetrasporic, 4- or 8-nucleate, stigmas capitate or not, (multicellular papillae +); endotegmen persistent; endosperm 4n or 5n, little persisting, embryo green.

27[list]/836 - three groups below. Predominantly Mediterranean to Central Asia, scattered elsewhere. [Photos - Collection]

1. Plumbaginoideae Burnett

Herbs or shrubs; 5-O-methylated flavonols +; stems angled and striate; leaves (deeply lobed), petioles often short, (cauline stipules - Plumbago); inflorescence racemose, vegetative and reproductive shoots similar; (heterostyly +); K herbaceous, glandular, C (connate), lobes truncate-emarginate and then apiculate, style +, receptive areas in bouquet-like aggregations along branch; fruit a basally circumscissile capsule, calyx herbaceous; n = 6, 7.

4/36. East Asia and Africa, Plumbago pantropical (Map: from Baker 1948, probably over-optimistic, Plumbago in particular commonly cultivated).

Many Plumbaginoideae seem to lack a protruding obturator (Dahlgren 1916). The sticky calyx glands of Plumbago persist in fruit.

2. Staticoideae Kosteletzky

Plumbagin 0, glycine betaines rare, betaalanine betaines +; leaves cartilaginous, with 5-10 marginal rows of whitish cells; C connate, stamens adnate to corolla, styles + (and style ± developed), stigma capitate (filiform).

2A. Aegialitideae (Link) Peng

Shrublet; ellagic acid +; successive cambia +; cortical vascular bundles +; branched sclereids; leaves involute, basal sheath surrounding stem; fruit?; n = ?

1/2. Indo-Malesian, Australia, in mangroves (Map: from van Steenis 1949c, in blue).

Little known.

Synonymy: Aegialitidaceae Linczewski

2B. Staticeae Bartling

Herbs or shrubs; (glycine betaines), betaalanine betaines +; leaves more or less clustered at base, (deeply lobed), inflorescence leaves reduced or absent, petioles often short; inflorescence capitate or cymose, axis channelled, vegetative and reproductive shoots different; (heterostyly +), K scarious (also petaloid), pollen often dimorphic, columellae regular, tectum incomplete, reticulate; fruit an achene or circumscissile capsule; calyx scarious; n = 8, 9; deletion of rpl16 intron.

22/800: Limonium (350), Acantholimon (165), Armeria (90: half on the Iberian Peninsula). Mostly Irano-Turanian (Mediterranean), but also S. Africa, S. South America, and W. Australia (Map: from Hultén; Baker 1948; FloraBase 2004).

Linczevski segregated 13 genera from Limonium - only 31 species involved. For a phylogeny focusing on Limonium, see Lledó et al. (2005).

The "style" of Armeria is papillate all around for its entire length. According to Dahlgren (1916), the embryo sac is tetrasporic but eight-nucleate. The calyx sometimes forms a scarious wing in fruit.

Synonymy: Armeriaceae Horaninow, Limoniaceae Seringe, nom. cons., Staticaceae Cassel

• Plumbaginaceae are perennial herbs or shrubs, rarely climbers. They can be recognised by their leaves which have broad bases and are frequently only shortly petiolate and/or with entire margins, and their flowers which have a relatively large, persistent, and sometimes colored calyx (which can be scarious in fruit) and antepetalous stamens. The single ovules borne at the ends of long, curved funicles are easily enough seen on dissection.

Betaines are quaternary ammonium compounds that are involved in salt excretion. For wood anatomy, which may be paedomorphic, the family perhaps having a more or less herbaceous ancestry, see Carlquist and Boggs (1996). There is extensive gross anatomical variation that probably can be integrated with the tribes/subfamilies: e.g. there is a continuous ring of sclerenchyma outside the phloem in Plumbaginoideae, separate fascicles in Staticoideae, etc. (see Maury 1886). Williams et al. (1994) suggested that it was not known if the mucilage glands were vascularized, although in their data matrix the family was scored as having such glands. Leaf vernation is flat, convolute or involute.

Lledó et al. (1998, 2001) suggest phylogenetic relationships within the group; these are consistent with the classification used here. It has also been suggested that Aegialitis may be sister to the rest of the family (Savolainen et al. 2000 - rbcL only); some of the characters attributed to Plumbaginaceae as a whole may need confirmation. Glycine betaine is known from only some species of Limonium (and from Plumbago), but not from Aegalitis and Armeria (Rhodes & Hanson 1993).

Primulaceae used to be associated with Plumbaginaceae. Both have stamens opposite the petals, common petal-stamen primordia, and a ± connate corolla (the latter especially in Staticoideae), but the two are not close - for Primulaceae, see Ericales.

Baker (1948, 1953) discussed variation in floral morphology (pollen, stigmas, etc.), de Laet et al. (1995) discuss floral development, Hegnauer (1969, 1990) chemistry, and there is much general information in Kubitzki (1993b).

POLYGONACEAE Jussieu, nom. cons.   Back to Caryophyllales

Shoots monopodial, branching from previous flush; soluble oxalate accumulation; cork subepidermal (pericyclic); dark-staining deposits, esp. in rays; pits vestured; nodes also 5 or more:5 or more; petiole with a (D-shaped) ring of bundles, (bundles scattered - some Coccoloba); mucilage cells common; soluble calcium oxalate accumulation; cuticle waxes as platelets or rodlets; (stomata dia- aniso- or paracytic); leaves revolute (convolute - Muhlenbeckia; margins lobed), 2ndary veins also palmate, stipule sheathing stem ["ochrea"], colleters +; (plant dioecious), inflorescences with flowers in fascicles, flowers small, pedicels articulated; flowers basically 3-merous, hypanthium ± developed, P spiral (4, 6), basally ± connate, one with two veins, stamens = to and alternate with P to 3 x P, pollen tricolporate to pantoporate, nectary a disc, or between A, G [(2) 3 (4)], ovule atropous, (unitegmic), nucellar beak +, (archesporium multicellular), (style short), stigma ± penicillate or capitate; fruit a trigonous (lenticular) achene; endosperm ruminate or not, embryo straight to curved, lateral or not; expansion of the chloroplast inverted repeat.

43[list]/1110 - two groups below. World-wide (Map: from Hultén 1971; FloraBase 2008; Tanya Schuster, pers. comm.). [Photos - Collection]

1. Polygonoideae Eaton

Shrubby, vines, herbs; (A 9), funicle long or short; n = 7 and up.

15/590: Polygonum (200+, sometimes split - as in Brandbyge 1993 - and then Persicaria 150), Rumex (200), Calligonum (80). Especially (warm) temperate.

Some species of Rumex have an X-autosome balance system determining the 'sex' of the plant. Stamens in Fagopyrum are both introrse and extrorse (Le Maout & Decaisne 1868).

Synonymy: Calligonaceae Chalkuziew, Persicariaceae T. Post & Kuntze, Rumicaceae Martynov

2. Eriogonoideae Arnott

Commonly woody, habit various; (plant dioecious); (inflorescence ± cymose, with involucre - Eriogonum et al.).

28/520: Eriogonum (250, but paraphyletic), Coccoloba (120). Largely tropical, especially America, Eriogonum and relatives esp. W. North America.

Interestingly in view of the paucity of mycorrhizae in the order, endomycorrhizae are reported from Eriogonum and ectomycorrhizae from Coccoloba (Malloch et al. 1980).

Synonymy: Eriogonaceae G. Don

• The flowers of Polygonaceae are rather small, all parts (bar the carpels) are more or less free, and the 5 or 6 tepals are persistent or enlarged in fruit; the achene itself is often angled. A sheathing stipule is common and the nodes of the herbaceous taxa in particular are more or less swollen. Eriogonum and relatives lack the distinctive stipule and have a more or less cymose and involucrate inflorescence.

Lycaeana and Heliophorus (Lycaenini) are found on this family throughout its extratropical range (Ehrlich & Raven 1964). Eriogonum and relatives are very diverse in the drier regions of southwest North America (with links with southern South America), and may represent a relatively recent radiation (Sanchez & Kron 2008).

Williams et al. (1994) noted that although no plumbagin had been reported from the family, other quinones were known there. Antigonon has leaf tendrils and successive cambia. There are often subepidermal strands of collenchyma or sclerenchyma in the stem in Polygonaceae. It has been thought that the flowers of Polygonaceae are basically 3-merous; the carpels are opposite the outer perianth whorl when the perianth is 3 + 3. Flowers with five tepals were derived from those with six, perhaps by fusion of two of the members. Recent work, however, suggests that the basic condition for the family is to have five tepals (Lamb Frye & Kron 2003). The exact nature of the funicle is unclear; it might be a reduced basal placenta. Sieve tube plastids with protein fibers are reported from Triplarieae (Behnke 1999).

The largely herbaceous Eriogonoideae s. str., i.e. Eriogonum and its immediate relatives, used to be kept separate from Polygonoideae, of variable habit, because the former lacked a sheathing stipule, their inflorescence was cymose and with an involucre, while the latter had a sheathing stipule and a racemose inflorescence that lacked an involucre. However, studies show a division of the family into two moderately supported clades, largely woody and largely herbaceous respectively. The former includes Eriogonoideae s. str., and this is derived from within the woody clade, while Coccolobeae appear to be paraphyletic (e.g. Cuénoud et al. 2002; Lamb Frye & Kron 2003); [Antigonon + Brunnichia], both lianes, are sister to the rest of the woody clade (Sanchez & Kron 2008). Eriogonum itself is paraphyletic and includes taxa like Chorizanthe and Dedeckera (Sanchez & Kron 2006, 2008). Polygonoideae s. str. are more or less herbaceous. Generic limits around Polygonum are difficult, and the mostly viney Muehlenbeckia (nestled in Fallopia) is also included in Polygonoideae (Schuster & Kron 2008).

For more information, see Hegnauer (1969, 1990: chemistry), Haraldson (1978: general), Brandbyge (1993: general), Carlquist (2003a: wood anatomy) and Logacheva et al. (2008: especially expansion of the inverted repeat). I thank Adriana Sanchez for comments.

Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]]: styles stigmatic their entire length; endosperm slight.

RHABDODENDRACEAE Prance   Back to Caryophyllales

Woody; ellagic acid +; cork?; (successive cambia 0); dark-staining deposits esp. in rays; nodes multilacunar; sieve tube plastids with protein crystalloids and starch; secretory cavities with resin; foliar fibre-like sclereids +; hairs peltate, cells with SiO₂ bodies; leaves revolute, obscurely punctate; inflorescence axillary, branched, ?with terminal flower; hypanthium +, short, K short, ± connate, C rather thick, A many, filaments very short, anthers very long, exodermis tanniniferous, nectary?, G 1, with 1 (2) basal campylotropous ovules, bitegmic zone short, nucellar cap 0, archesporium multicellular, stylodium basal, stigma much elongated, ?type; fruit a drupelet, surrounded by K, pedicel swollen; exotestal cells tangentially elongated, underlying cells short-tracheidal; endosperm type?, 0, embryo green, curved, with large cotyledons; n = 10.

1[list]/3. Tropical South America (Map: see Prance 1972c). [Photo - Flower]

• Rhabdodendraceae are evergreen trees that may be recognised by their fringed-peltate hairs, their rather large, entire and exstipulate leaves, flowers with stamens that have short filaments and long anthers, and a single carpel with a basal style. The fruit is distinctive, the drupes being surrounded by the persistent calyx and swollen pedicel apex. The punctations in the lamina may not be at all obvious.

I have not seen stipules (cf. Prance 1972c), but the rather broad petiole base can be confused with them. The ovule is often described as being unitegmic (e.g. Nandi et al. 1998, following Puff & Weber 1976; but see Tobe & Raven 1989). The styles may be stigmatic for only part of their lengths. The embryo is surrounded largely by testa that develops from the unitegmic part of the ovule.

The position of Rhabdodendraceae has long been uncertain. Thus they were placed in Rutales by Takhtajan (1997), although Prance (1968) had suggesyted a position in this general area.

For general information, see Prance (2002).

Simmondsiaceae [[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]]: nodes 1:1; C 0.

This clade may have diverged from the [Droseraceae group + Tamaricaceae + Polygonaceae] clade 82-76 million years before present, but diversification of the core Caryophyllales did not occur until substantially later at 47-39 million years before present (Wikström et al. 2001).

The morphology, etc., of the basal members of this clade seem rather different from those of the core members.

SIMMONDSIACEAE van Tieghem   Back to Caryophyllales

Evergreen shrubs; ellagic acid 0; cork pericyclic; true tracheids +; stomata cyclocytic and laterocytic; hairs uniseriate; leaves opposite, articulated near stem, flat, 2ndary veins ascending from near base; plant dioecious; flowers small (4, 6-merous), C 0, nectary 0, staminate plant: inflorescence usu. terminal, cymose; A 2x P, extrorse, anthers basifixed, longer than connective, pollen ± porate, central part operculoid, spinules minute; carpellate plant: flowers axillary, G [3], 1 apical pendulous apotropous ovule/carpel, outer integument ca 10 cells and inner integument 4-5 cells thick, styles papillate all around; fruit a capsule, K accrescent, columella persistent; seed 1, testa multiplicative, vascularised, exotestal cells palisade, walls thickened, mesotesta aerenchymatous, rest collapsed; endosperm reserve?; n = 13.

1[list]/1: Simmondsia chinensis (!: note the epithet). S.W. North America, the Sonoran Desert (Map: see Sherbrooke & Haase 1974). [Photos - Collection]

• Simmondsiaceae are evergreen shrubs that may be recognised by their opposite leaves that are clearly articulated near the stem and have flat, rather thick blades with entire margins. The staminate flowers are small and borne in terminal inflorescences while the carpellate flowers are single and axillary; the calyx is much enlarged in fruit.

Simmondsiaceae have usually been included in Buxaceae or placed in a separate family, but close to Buxaceae; Simmondsiales are included in Hamamelididae (Takhtajan 1997).

The large embryo contains liquid wax, esters of high molecular weight, mono-ethylenic acids. The stamens are described as being latrorse (Takhtajan 1997).

For general information, see Mathou (1939), for chemistry, see Hegnauer (1989, as Buxaceae), for testa anatomy, etc., see Tobe et al. (1992), for general information, see Köhler (2002), and for wood anatomy, see Carlquist (2002b).

[[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]]: ?

[Asteropeiaceae + Physenaceae]: successive cambia 0; young stem with vascular cylinder; abaxial-confluent parenchyma; vasicentric tracheids +, fiber tracheids +; rays 1-2 cells wide; A latrorse; fruit single-seeded.

Some information on the general anatomy of these two families is taken from Harms (1893); Carlquist (2006) compares their wood anatomy.

ASTEROPEIACEAE Reveal & Hoogland   Back to Caryophyllales

Evergreen trees or scrambling shrubs; ellagic acid?; pericyclic fibers +; petiole bundle annular; epidermal mesophyll sclereids +; ?stomata; inflorescences terminal, branched, pedicels with many bracteoles; C +, deciduous, A 9-15, basally connate, (pollen 6-rugate), G [(2) 3], 2-many ± apical ovules/carpel, ?micropyle, (style short, stigma lobed), stigma continuous across G; fruit nutlike, (several-seeded), K accrescent and forming wing, A persistent; seed coat?; endosperm reserve?, embryo curved, cotyledons spirally coiled; n = ?

1[list]/8. Madagascar. [Photos - Collection]

• Asteropeiaceae are evergreen trees that may be recognised by their shortly petiolate leaves whose blades are entire, exstipulate and have rather indistinct venation. The flowers are small are in terminal panicles, the fruit has a single seed, and the calyx is much accrescent and spreading.

Asteropeia is ectomycorrhizal (Ducousso et al. 2008).

Asteropeiaceae were previously often included in Theaceae or Theales (Cronquist 1981; Takhtajan 1997), but are very different in wood anatomy (Baretta-Kuipers 1976); the rays are uniseriate.

Some information is taken from Beauvisage (1920: anatomy), Morton et al. (1997b: general), Schatz et al. (1999: revision), and Kubitzki (2002d: general).

PHYSENACEAE Takhtajan   Back to Caryophyllales

Shrub or tree; ellagic acid?; pericyclic sclereids +; cuticle wax crystalloids?; leaves two-ranked; plant dioecious; inflorescence racemose; P 5-9, staminate flowers: A (8-)10-14(-25), basifixed, anthers long; carpellate flowers: G [2], septae incomplete, 2 ± subbasal campylotropous ovules/carpel, funicle long; fruit subdrupaceous?; seed large, coat vascularised, 16-20 layers thick, walls not notably thickened; endosperm 0, reserve?, cotyledons unequal; n = ?

1[list]/2. Madagascar. [Photos - Collection]

• Physenaceae are rather undistinguished with their more or less two-ranked, entire, exstipulate and coriaceous leaves. The flowers lack a corolla, the stamens have a very thin filaments and long anthers, and the styles are long. The fruit is inflated and contains a single seed which apparently has a vascularised seed coat.

The petiole is often described as being articulated; it commonly breaks transversely above the base. The leaf bundles are completely surrounded by mechanical tissue. There aare brachysclereids in the secondary phloem and the placental bundles are inverted (Dickison & Miller 1993).

Physenaceae were included in Urticales by Cronquist (1981) and placed in a monotypic Physenales in Dilleniidae by Takhtajan (1997).

General information is taken from Morton et al. (1997b) and Dickison (2002).

FROM NOW ONWARDS, THINGS ARE SOMEWHAT IN A STATE OF FLUX (but there have been recent improvements).

[Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]: Plant herbaceous; CAM [especially pervasive in succulents] and C₄ photosynthesis common; ferulic acid ester-linked to primary unlignified cell walls; (O-methylated) flavonols, quinones, betalains [chromoalkaloids], triterpenoid saponins +, tannins, myricetin 0; (phytoferritin +); sieve tube plastids with a ring of proteinaceous filaments and a central angular crystalloid (also with starch); pericyclic fibers 0 [phloem-derived fibers quite widespread]; (mucilage cells); (stomatal orientation transverse); inflorescence cymose; (stamens equal and opposite perianth members), pollen trinucleate, (polyaperturate), foot layer thin, nectaries on adaxial bases of stamens, placentation free central or basal, G with median member adaxial, ovules campylotropous, (funicles long), stigmas papillate, little expanded; seeds with exotestal and endotegmic cells thickened, (space between testa and tegmen), bar-like thickenings on endotegmic cells; endosperm 0, perisperm +, starch grains clustered, embryo curved; mitochondrial rps 10 gene and chloroplast rpl2 [latter present in some Portulaca?] gene intron lost.

Core Caryophyllales contain ca 5.3% of eudicot diversity (Magallön et al. 1999). Fossils identified as Amaranthaceae are dated to the Santonian/Campanian, ca 83 million years before present (Magallön et al. 1999), but molecular estimates of the age of the clade are only some 28-40 million years before present (Wikström et al. 2001) - something is clearly wrong. Taxa growing in saline and/or dry conditions are noticeably well represented here, and taxa that can grow on gypsum (hydrous calcium sulphate) are scattered throughout the clade (Douglas & Manos 2007). Succulents are common, and many taxa have C₄ photosynthesis of CAM or their variants. Core Caryophyllales are little liked by butterfly caterpillars (Ehrlich & Raven 1964).

Sterol composition may be of systematic interest (Wolfe et al. 1989; Patterson & Xu 1990), with distinctive sterols common or dominant in Caryophyllaceae, Phytolaccaceae, Amaranthaceae, and "Portulacaceae"; isoflavonoids (Reynaud et al. 2005) and phytoecdysones are scattered, but perhaps not in the Cactaceae area, and the former are sometimes quite diverse. Stomatal morphology is variable, but anomocytic stomata are common in all families. However, in Cactaceae and relatives, parallelocytic and other kinds of stomata are found; some families in this area have predominantly paracytic stomata. Stomatal orientation on stem and/or leaf is commonly transverse apparently throughout the order (Butterfass 1987, ?Amaranthaceae s. str.?), however, it is unlear what taxa have vertically or unoriented stomata. Any "corolla" present, as in Caryophyllaceae, is usually described as being of staminal origin (e.g. Ronse Decraene & Smets 1993; Leins et al. 2001); it arises at the same time or after the androecium, not before it, and the "petals" and stamen(s) opposite them may form a developmental unit, and the corona - in Lychnis viscaria, at least - arises from two bulges on the adaxial side of the "corolla", perhaps representing anther thecae. When the stamens are equal in number to the perianth members they are opposite to them, when there are many stamens the initial primordia either alternate with them (Aizoaceae), or continue the spiral of the tepals (Pereskia - Cactaceae: see Leins & Erbar 1994); development is centrifugal. Carpels are quite commonly open in development - also in Polygonaceae (Tucker & Kantz 2001). Placentation is quite variable, although one commonly thinks of this group as being free central of its variants; the basal condition for the clade may indeed be free-central. There may be a subepidermal layer of cells in the ovary with conspicuous calcium oxalate deposits, as in some Amaranthaceae and Polygonaceae(!), although in Nyctaginaceae, for example, cells immediately below the ovary have conspicuous raphide deposits (Guéguen 1901); there is little information on this feature. The apical cells of the nucellus are commonly elongated radially (e.g. in Cactaceae, "Portulacaceae", Aizoaceae, Phytolaccaceae, and Amaranthaceae: see Johri et al. 1992), i.e., they form a nucellar pad, but it is unclear if this is a feature of systematic significance. There are often short hairs on the funicle that are directed towards the micropyle (Neumann 1935). Seeds of a number of taxa have an operculum, although not necessarily identical in morphology (Bittrich & Ihlenfeldt 1984). Finally, there are commonly bar-like thickenings on the walls of the endotegmic cells (e.g. Netolitsky 1926; Bittrich 1993), although these are absent from most Caryophyllaceae, at least - a complete survey would be useful. For the loss of the intron of the rpl2 gene, see Logacheva et al. (2008).

The course of evolution of betalains and anomalous secondary thickening in this group has long been uncertain, but it now seems unlikely that the presence of anthocyanins is plesiomorphic (Cuénoud et al. 2002; Cuénoud 2002a; see also Clement & Mabry 1996), and normal secondary thickening may also be apomorphic.

Most of this group was included in the old Centrospermae (so named because of the basal or free-central placentation that is common in the clade) or Caryophyllales in the strict sense. Achatocarpaceae + Amaranthaceae + Caryophyllaceae form a moderately well supported clade, the rest of the core Caryophyllales another (Källersjö et al. 1998), however, although 13 families were included, sampling within them was poor. Similar relationships were suggested by Savolainen et al. (2000a). D. Soltis et al. (2000) found that Phytolaccaceae, Nyctaginaceae and Delosperma (Aizoaceae) formed a group, also Amaranthaceae plus Caryophyllaceae, but again the sampling was very sparse. For other suggestions of relationships, see Rodman (1994) and Downie and Palmer (1994: structural variation in chloroplast DNA). For relationships here I have largely followed those suggested by Cuénoud et al. (2002: note, they excluded the Delosperma sequence), largely similar to those found by Källersjö et al. (1998) and other workers, albeit the sampling is much more detailed. Cuénoud et al. (2002) found two quite well supported clades within core Caryophyllales (see tree), but sampling still needs to be improved. Cuénoud et al. (2002, along with others) found Aizoaceae to be monophyletic, albeit with only slightly better than marginal (52%) support in an analysis of matK sequences, the only gene for which they had moderately good sampling. Giseckia moved position in analyses of rbcL and matK sequences (cf. Cuénoud et al. 2002), and Sarcobatus is sister to Nyctaginaceae, albeit with only weak support, in matK analyses, while in a rbcL analysis it groups with Agdestis (cf. Cuénoud et al. 2002). In general, morphological distinctions between clades are slight; combination of families may well be order.

Additional information is taken from Zandonella (1977: nectaries), Rutishauser (1981: "stipules" and similar structures), Hegnauer (1989: general chemistry), Wolfe et al. (1989: sterols), Patterson & Zu (1990: sterols), Steglich and Strack (1990: betalains), Barthlott (1994: waxes), Behnke (1994a: sieve tube plastids, phytoferritin), Gibson (1994: stem anatomy), Hofmann (1994: floral morphology), Nowicke (1994: pollen), Werker (1997: seed coat), Jansen et al. (2000c: successive cambia), Cuénoud (2006: summary) and Muhaidat et al. (2007 and references: C₄ pathway).

Caryophyllaceae [Achatocarpaceae + Amaranthaceae]: (phytoecdysteroids +); esp. outer wall of exotesta thickened and with stalactite-like projections; mitochondrial rps1 and 19 genes lost.

The condition of these characters in Achatocarpaceae is unknown. For phytoecdysteroids, see Báthori et al. (1987), Dinan et al. (1998), and Zibareva et al. (2003). Sukhorukov (2007) described the exotgmic cells of Chenopodiaceae s. str. as often having tannin deposits in the outer walls of the exotegmic cells that projected into the cell lumen.

CARYOPHYLLACEAE Jussieu, nom. cons.   Back to Caryophyllales

Herbs (shrubs, lianes); cyclopeptides, anthocyanins, glycoflavones, anthraquinones +; cork usu. pericyclic; true tracheids, fibers +; pericyclic fibers +; nodes often swollen; stomata often diacytic; (cuticle waxes as rodlets); leaves opposite, conduplicate or ± flat, often connate at the base; (plant dioecious); flowers 4-5-merous, "C" often bilobed, A (1-) 5, 10 (obdiplostemonous; 15), outer secondary parietal cell dividing, (pollen 6(+) porate; nectary on receptacle), G [2-5], also opposite "C", (short gynophore), (1 basal-)many ovules/carpel, initially distinct air space between the cotyledons [?all taxa], nucellar cap +, (style +), placentation often axile basally when young, funicle?; fruit a septicidal and loculicidal capsule (circumscissile); (funicular strophiole - Moehringia; endotesta thickened), endotegmen (± thickened), lacking bars; n = 7--15, 17; protein bodies in nuclei; mitochondrial coxII.i3 intron 0.

86[list]/2200: Silene (720), Dianthus (300), Stellaria (175), Gypsophila (150), Arenaria (150: polyphyletic), Minuartia (120: polyphyletic), Paronychia (110), Cerastium (100), Acanthophyllum (75), Drynaria (50). Mostly temperate (Map: from Vester 1940; Hultén 1971). [Photos - Collection, Minuartia Habit,Microphyes Flower]

• Caryophyllaceae can be recognised by its herbaceous habit, opposite, entire leaves that often have little in the way of a petiole and are joined by a line at the base; geotropic adjustments occur at the swollen nodes. The inflorescence is cymose and the flowers appear to have a clearly distinguishable calyx and corolla, the latter being bilobed and/or clawed; there are usually 5 or 10 stamens. The fruit is a capsule.

Ectomycorrhizae have been reported from the family (Wang & Qiu 2006). Silene and its relatives may be pollinated by moths which at the same time lay eggs on the ovary, rather like the yucca - moth association (Kephart et al. 2006 for literature, also other papers in that issue of the New Phytologist [169(4)]).

The old tripartite division of the family into Silenoideae, Alsinoidae and Paronychioideae is not confirmed by any recent work (e.g. Nepokroeff et al. 2002; Smissen at al. 2002; Fior et al. 2006). Paronychioideae are a basal grade, with Corrigioleae (Telephium, Corrigola) sister to the rest of the family. Dicheranthus, Polycarpon, etc. may be the next clade, Paronychia, etc., the next. Drymaria and Pycnophyllum, both morphologically distinctive taxa, may be sister (Smissen et al. 2002 - they noted that Pycnophyllum [and Pentastemonodiscus] were not to be included in Caryophyllaceae-Alsinoideae, but they did not suggest where they should go; Fior et al. 2006). In the erstwhile Alsinoideae the calyx is free and the corolla has ± open venation. Alsinoideae for the most part break down into two groups: one, including Cerastium, Stellaria, etc., has capsules with split valves, and the other including Minuartia, etc., is very diverse, but has capsules with entire valves; the corolla is often bilobed. For Moehringia, the evolution of its strophiole, and its allies, see Fior and Karis (2007 and references). Caryophylloideae have a connate calyx and clawed corolla with more or less closed venation and adaxial appendages (ligules). The tribes Sileneae and Caryophylleae are perhaps monophyletic, and together are sister to or form a polychotomy with part of Arenaria (Nepokroeff et al. 2002; Fior et al. 2006); for Silene and its relatives, perhaps not monophyletic, see Erixon and Oxelman (2008).

The calyx can be very scarious, as in Brachystelma, while herbaceous Amaranthaceae also often have swollen nodes (and opposite leaves)!

Weberling (1989 and references) discusses the placentation of the family, which varies from axile to free central to the single, basal ovule of Uebelinia - which is getting close to a circinotropous basal ovule. Whether styles are free or there is obviously a stylar region as well, and also carpel number, have provided generic characters... Members of the old Paronychioideae have solanad rather than caryophyllad embryo development. Some species of Silene have an X-Y 'sex' determination system.

Some information is taken from Bittrich (1993), for stem anatomy, see Schweingruber (2007), general chemistry, see Hegnauer (1964, 1989), for the distribution of phytoecdyosteroids, see Zibareva et al. (2003), and of cyclopeptides, see Jia et al. (2004).

Synonymy: Alsinaceae (Lamarck & Candolle) Bartling, nom. cons., Dianthaceae Vest, Cerastiaceae Vest, Corrigiolaceae (Dumortier) Dumortier, Herniariaceae Martynov, Illecebraceae R. Brown, nom. cons. (cuticle waxes with long low crystals, or none), Lychnidaceae Lilja, Ortegaceae Martynov, Paronychiaceae Jussieu, Polycarpaeaceae Schur, Scleranthaceae Berchtold & J. S. Presl, Silenaceae Bartling, Spergulaceae Adanson, Stellariaceae Dumortier, Telephiaceae Martynov

Achatocarpaceae + Amaranthaceae: pollen porate.

ACHATOCARPACEAE Heimerl, nom. cons.   Back to Caryophyllales

Woody; C-glycosylflavonoids +, betalains?; cork?; secondary growth normal; nodes ?; cuticle waxes as ± lobed platelets in clusters; P 4-5, A 10-20, basally connate or not, pollen 6-porate, G [2], lateral or superposed, 1 (2) basal ovules, funicle?; fruit a 1-seeded berry; seeds with small aril; n = ?

3[list]/7. S.W. USA to South America (Map: from Fl. N. Am. 4: 2003; GBIF 2008). [Photo © C.E. Hughes - Fruits, Fruiting branch].

Some information is taken from Bittrich (1993).

AMARANTHACEAE Jussieu, nom. cons.   Back to Caryophyllales

Succulent, herbaceous or shrubby (lianes), often in saline conditions; betaines, anthraquinones, (isoquiniline alkaloids), 6-7-methylene-dioxyflavonols, isoflavonoids +, soluble oxalate accumulation; often a few pericyclic fibers +; cork pericyclic (and elsewhere) [esp. chenopods]; wood storied, rayless; crystal sand + [less common in chenopods], soluble calcium oxalate accumulation; cortical and/or medullary bundles + [less common in amaranths s. str.]; sieve tube plastids lacking protein crystalloid; nodes often swollen, (1:3, 1:5); petiole bundles ± annular; stomata also paracytic (dia- and anisocytic); hairs variable, often uniseriate; (leaves opposite), margins often toothed; (bracts and bracteoles scarious); P (1-)5(-8), ± herbaceous [chenopods] to scarious, stamens = P, joining disc (± connate; with appendages [pseudostaminodes]), (coloured vesicular anther appendages - Caroxyloneae), anther wall development monocotyledonous, (tapetum plasmodial), pollen multiporate, often starchy, foot layer well developed, G [1-3(-6)], (median member abaxial), with 1(-many) basal ovules, (chalazal region ± digested by the embryo sac), style ± developed, stigmas capitate; fruit surrrounded by a persistent (subfleshy) perianth [anthocarp], bracts and bracteoles persistent and also often part of disseminule, or dry, circumscissile capsule or indehiscent (berry, drupe); seed horizontal or not, endotegmen ± thickened and lignified, tanniniferous; (perisperm 0), embryo green or white (spiral - Salicornia etc.; straight); n = (6-)8, 9(-17).

174[list]/2050-2500: Chenopodium (100: C. quinoa), Atriplex (300), Gomphrena (120), Salsola (100), Alternanthera (100), Iresine (80), Amaranthus (60), Celosia (45). ± World-wide, esp. warm and dry temperate and subtropics, esp. saline habitats (Map: from Hultén & Fries 1986; Jalas et al. 1999). [Photo - flowers, fruits, Collection.]

• Amaranthaceae are more or less succulent herbs often with swollen nodes; they prefer dry and/or saline habitats. The small flowers often have 1-3 carpels and 1-2 basal ovules. The fruits either have a circumscissile capsule and a chartaceous to scarious perianth and bracts or are indehiscent and are closely invested by the more or less fleshy perianth and bracts.

Many taxa are halophytes (Jacobs 2001; Sage 2002). Although the family is apparently largely without mycorrhizae, vesicular-arbuscular mycorrhizae have been reported from chenopods in the Red Desert of Wyoming - but only on native taxa and under undisturbed conditions (Miller 1979).

There are several types of C₄ photosynthesis with ca 17 different kinds of leaf anatomy and probably 10+ independent acquistions of this photosynthetic pathway in Amaranthaceae (Kadereit et al. 2003; Sage et al. 2007; for details of the evolution of enzymes involved in Alternanthera, see Gowick et al. 2006); with some 800 species with C₄ photosynthesis, Amaranthaceae are by far the largest BLA group with this photosynthetic pathway. In three Suaedeae s.l. all the different aspects of C₄ photosynthesis occurs within a single cell (i.e. there is no Kranz anatomy), and this has evolved independently at least twice (Kapralov et al. 2006); partitioning of the organelles within the cell is maintained by the distinctive organization of the cytoskeleton (Chuong et al. 2006), although there is plasticity induced by the light-environment (Lara et al. 2008).

Amaranthaceae (inc. Chenopodiaceae) are palynologically fairly homogeneous (Nowicke 1975; Skvarla & Nowicke 1976), both having a similarly thickened tectum, apertures with reduced pointed flecks of exine underlain by lamellar plates, and a thickened endexine. Müller and Borsch (2006c) discuss the evolution of the distinctive stellate pore ornamentation of some Amaranthaceae s. str. - independent gains and losses.

Amaranthaceae are morphologically heterogeneous. Some problem taxa: Pleuropetalum (leaves spiral; inflorescence racemose, P 5, A 8, connate basally, G 5-6, several basal ovules, fruit initially fleshy; n = 8, 9 - A paired in development [Ronse Decraene et al. 1999]), in Amaranthoideae (Townsend 1993); Microtea (leaves spiral; inflorescence racemose, flowers in groups of up to 3; A (2-)5-9, pollen pantoporate, G [2-4], orientation variable, unilocular, funicle quite long [Baillon 1886], styles diverging; fruit an achene. Phytolaccaceae-Microteoideae Nowicke. 2/13. Central and South America, Antilles; S. Africa [Urban 1885], and with Lophiocarpus (Rohwer 1993) in Phytolaccaceae - Lophiocarpaceae Doweld & Reveal.

For discussion on the cortical vascular system, see Beck et al. (1982). Stem collenchyma is well developed; there are nucleated xylem fibers (Rajput 2002). 2-carpellate members of the family usually have transverse carpels, but occasionally they are median. The chalazal region of the ovule is more or less digested by the embryo sac in at least some Amaranthaceae - and this is also once recorded from Nyctaginaceae (Maheshwari 1950).

Amaranthus (Amaranthoideae: anthers 4-locellate at maturity, dehisce by 2 slits; 1-many ovules) sister to Beta, etc., in ORF 2280 phylogenies, and this whole group sister to Celosia [Celosieae - monophyletic, several ovules, derived) and Froelichia, etc. + Gomphreneae - monophyletic, anthers monothecal, pollen metareticulate, the mesocolpium raised (see Downie et al. 1997)] which together have a monadelphous androecium (and very scarious perianth and bracts). Amarantheae are polyphyletic. Cuénoud et al. (2002) found Amaranthaceae s. str. to be monophyletic, with very strong (97%) support, and Chenopodiaceae s. str. were perhaps monophyletic, but the branch collapsed in a strict consensus tree; the sampling was moderately good, but only one gene - matK - was sequenced and analysed. In an extensive rbcL analysis, much of the old Chenopodiaceae were monophyletic, but with little bootstrap support, ditto the old Amaranthaceae (incl. Polycnemoideae), Betoideae were paraphyletic (G. Kadereit et al. 2003). Other studies suggest paraphyly of Chenopodiaceae and sometimes even potential polyphyly of Amaranthaceae (Pratt et al. 2001). In a recent analysis of matK/trnK sequences, Müller and Borsch (2005b; see also Müller & Borsch 2005c), Polycnemum and Nitrophila (100% support) were sister to the rest; they have ordinary secondary thickening, imperfect flowers, basally connate filaments, and unithecate anthers. The rest of the Amaranthaceae + Chenopodiaceae had <70% support (and still lower posterior probabilities), while the Amaranthaceae s. str. had 100% support and the Chenopodiaceae s. str. again <70% support, but this time 100% posterior probabilities. Within Amaranthaceae - at least some flowers imperfect - Bosea and Charpentiera were successively sister to the rest, but Amaranthoideae and Amarantheae were paraphyletic. Some of the extreme halophytic genera are morphologically much modified, and generic limits are difficult; in the reduced perianth of the Australian Tecticornia (Salicornioideae) the odd member is abaxial (Shepherd et al. 2004, 2005a, esp. Shepherd & Wilson 2007, also nomenclatural changes). See also Schütze et al. (2003), G. Kadereit et al. (2005: Australian chenopods, 2006: Salicornioideae) and Akhani et al. (2007: Old World Salsoleae) for studies on groups of Chenopodiaceae sensu stricto.

Amaranthaceae sensu stricto have cuticle waxes lacking platelets; scarious bracts and perianth, and the filaments are often connate; n = (6-)8-9(-13, etc). The embryogeny is chenopodiad[!]. Chenopodiaceae sensu stricto quite commonly have isoflavonoids; cuticle waxes as platelets; bracts and P ± fleshy, pink to red; fruit rarely circumscissile; n = 9; 300 bp deletion in chloroplast DNA inverted repeat.

Additional information is taken from Hegnauer (1964, 1989: chemistry), Blunden et al. (1999: betaine distribution, very common and widespread), Robertson (1981: general), Kühn (1993), Townsend (1993: general), Judd and Ferguson (1999: general), Rajput (2002: anatomy), Carlquist (2003c: anatomy), Shepherd et al. (2005b; seed anatomy), and Sukhorukov (2007: fruit wall anatomy); see also Pratt (2003: phylogeny).

Synonymy (stamens = Amaranthaceae s. str., C = Chenopodiaceae s. str.): Achyranthaceae Rafinesque (A), Atriplicaceae Durande (C), Betaceae Burnett (C), Blitaceae T. Post & Kuntze (C), Celosiaceae Martynov (A), Chenopodiaceae Ventenat, nom. cons., Corispermaceae Link (C), Deeringiaceae J. Agardh (A), Dysphaniaceae (Pax) Pax, nom. cons. (C: cuticle waxes absent), Gomphrenaceae Rafinesque, Polycnemaceae Menge (C), Salicorniaceae Martynov (C), Salsolaceae Menge (C), Spinaciaceae Menge (C)

Stegnospermataceae [Limeaceae [Lophiocarpaceae [Aizoaceae, Nyctaginaceae, etc.]] [Molluginaceae, Cactaceae, etc.]]: ?

STEGNOSPERMATACEAE Nakai   Back to Caryophyllales

Woody, ± scandent; plant glabrous; true tracheids +; leaves fleshy; inflorescence racemose; "C" (2-)5, A (5) 8-10, connate basally, nectaries in depressions at base of G, G [2-5], opposite "C", placentation becoming free-central, 1 basal epitropous amphitropous ovule/carpel, placental obturator +, stigmas recurved; fruit a capsule; seeds arillate, exotesta ± palisade, unlignified, endotegmen enlarged, persistent; n = ?

1[list]/3. Central America, the Antilles (Map: from Bedell 1980). [Photo - Fruit]

• Stegnospermataceae are more or less scandent or sprawling plants with rather fleshy leaves and racemose inflorescences that have apparently petaloid flowers with a more or less rotate corolla.

There is no nucellar cap. Are the seeds endospermic?

Stegnospermaceae have often been included in Phytolaccaceae, which they looks like and which have a somewhat similar gynoecium, but they are most obviously distinguishable by their flowers with "petals". They also have pollen with a prominent foot layer and massive endexine - this is thin in Phytolaccaceae. The ovule is epitropous, in pluricarpellate Phytolaccaceae it is apotropous (Rogers 1985). Like Caryophyllaceae, there are idioblasts in the wood with sphaerites; there is only diffuse axial xylem parenchyma.

For more information, see Friedrich (1956: cf. carpel position), Hoffmann (1977: general), Bedell (1980: general), Horak (1981: secondary thickening), Narayana and Narayana (1986: embryology) and Rohwer (1993a: general).

Wide-band tracheids are scattered through this group, occuring especially in the very succulent members. Limeaceae, Cactaceae and "Portulacaceae" have cells in rows along the dorsal junction of the seed.

LIMEACEAE Reveal   Back to Caryophyllales

Herbs or subshrubs; anthocyanins +; cork?; (secondary thickening normal); leaves spiral, stipules 0; "C" + (0), adnate to base of staminal tube, A connate basally, G [2-7], opposite sepals, (pseudomonomerous and secondarily divided [Limeum]), placentation axile, 1-3 ovules/carpel, antipodal cells persist, placental obturator +; fruit a membranaceous capsule or schizocarp; seeds arillate or not; testa with cells in rows along the dorsal junction; n = 9.

2/23: Limeum (21). Southern Africa, to Ethiopia, S. Asia, also Australia (Macarthuria) (Map: from Culham 2007; FloraBase 2007).

For further information, see Hoffmann (1973: flower, growth), Behnke et al. (1983b: Macarthuria), Endress and Bittrich (1993: general, as Molluginaceae), and Hassan et al. (2005a: seed coat).

[[Lophiocarpaceae [Aizoaceae, Nyctaginaceae, etc.]] [Molluginaceae, Cactaceae, etc.]]: sieve tube plastids with globular crystalloids.

[Lophiocarpaceae [Nyctaginaceae, Aizoaceae, etc.]]: ?

LOPHIOCARPACEAE Doweld & Reveal   Back to Caryophyllales

Anthocyanins?; inflorescence a raceme or leaf-opposed cyme [Corbichonia, inflorescence evicted]; C 0 [Lophiocarpus], or C many, A many, centrifugal [Corbichonia], pollen tricolpate, G [2], 1-locular, ovule single, or [5], many ovules/carpel, placentation axile, placental obturator +; fruit an achene or capsule.

2/6. Africa, esp. S.W. Africa.

For Corbichonia flowers, see Ronse de Craene (2007), for embryology, see Narayana and Lodha (1963: the ovules are more or less anatropous and thinly crassinucellate).

Barbeuiaceae + Aizoaceae + Phytolaccaceae + Sarcobataceae + Nyctaginaceae: ?

BARBEUIACEAE Nakai  Back to Caryophyllales

Lianes; betalains?; libriform fibers, diffuse axial parenchyma, true tracheids +; sieve tube plastids with polygonal crystalloids[!]; cortical fibers +; druses +; leaves spiral; P 5, A many, pollen tricolporoidate, G [2], one ovule/carpel; fruit a loculicidal capsule; seeds 1 or 2, arillate, testa cells elongated, with sinuous anticlinal walls; n = ?.

1[list]/1: Barbeuia madagascariensis. Madagascar (Map: from Culham 2007).

The plant dries black.

See Hoffmann (1977), Rohwer (1993a: general, under Phytolaccaceae) and Carlquist and Schneider (2000: anatomy).

Aizoaceae + Phytolaccaceae + Sarcobataceae + Nyctaginaceae: soluble oxalate accumulation; raphides +.

For soluble oxalate accumulation, see Zindler-Frank (1976).

AIZOACEAE Martynov, nom. cons.   Back to Caryophyllales

Leaf succulents; growth sympodial; CAM +; cork from inner cortex or endodermis; wood storied; fibers ± in bands; wood rayless; wide-band tracheid pith cells; cuticular waxes as ribbons or rodlets; stomata also para- and anisocytic; leaf trace bundles forming reticulum in cortex; leaves opposite, with bladder-like cells on epidermis, bases broad, membranous; growth sympodual; hypanthium +, P colored internally, often with subapical abaxial appendage ["horn"], A often many, centrifugal, primordia 5, pollen tricolp(oroid)ate, nectary as ring, G septate, placental obturator +; seeds brown, exotesta ± palisade, or tangentially elongated; x = 8.

123[list]/ca 2020 - four subfamilies below. Esp. southern Africa, also Australia, etc., tropical and subtropical, arid. [Photos - Collection]

1. Sesuvioideae Lindley

Nodes also 3:3; stipules +; prophylls often prominent; (hypanthium +), A primordia opposite P, or development rather chaotic, G (1-)[2(-5]), 2-many ovules/carpel; capsule circumscissile (compound, fused with spiny bracts - Tribulocarpus); seeds shiny black, arillate (not).

4/36: Trianthema (17), Sesuvium (12). Tropics and Subtropics; Sesuvium portulacastrum is pantropical on beaches (Map: see George 1984; Hartmann 2001a, b; Fl. N. Am. 4: 2003). [Photos - Habit; Flower]

Tribulocarpus (ex Tetragonioideae, for which, see Aizooideae) is sister to the other Sesuvioideae, in which it is included here (Klak et al. 2003); it has an indehiscent fruit and hardly surprisingly lacks arillate seeds.

For a phylogeny of Sesuvioideae, see Hassan et al. (2005b).

Synonymy: Sesuviaceae Horaninow

Aizooideae [Mesembryanthemoideae + Ruschioideae]: inflorescence not distinct from vegetative plant, bract/eoles foliaceous; A primordia alternating with T; fruit a hygrochastic capsule.

2. Aizooideae

Bladder hairs with large terminal cell and multicellular stalk[?]; accessory lateral branches + [?]; inflorescence leafy; G [2-10], to inferior, 1 apical apotropous/basal-many ovules/carpel; (fruit septicidal - Gunniopsis; indehiscent - Tetragonia); seeds upright [?], (cell walls of seed coat little thickened).

7/135: Tetragonia (85). Drier parts of S. Africa, also Australia (Gunniopsis), few N. Africa and Asia Minor, N. America, etc. (Aizoon) (Map: see George 1984; Hartmann 2001a, b). [Photo - Flower]

Tetragonia is embedded in Aizooideae (Klak et al. 2003), however, it has rays, it lacks the bands of fibers of other Aizoaceae, and has vascicentric parenchyma adjacent to the fibers (Carlquist 2007).

Synonymy: Galeniaceae Rafinesque, Tetragoniaceae Link

Mesembryanthemoideae + Ruschioideae: leaves very succulent; P green, sepalline, "C" [staminodial] many, G more or less inferior, nectary interrupted; x = 9.

There is a combined [list] of genera recognised in these two subfamilies, but genus (and species) boundaries are uncertain (see below).

3. Mesembryanthemoideae Ihlenfeldt, Schwantes & Straka

Distinctive alkaloids [in Phyllobolus, etc.] +; cortical bundles +; (succulent persistent green cortex in stem); flowers 4-5-merous, nectary hollow or shell-shaped [koilomorphic], G [(3-)4-5(-6)], semi-inferior, placentation axile; expanding keels of fruit purely septal.

1/100. S. Africa, a few species also W. South America, Australia, N. Africa, the Mediterranean and the Near East, naturalised in W. North America (Map: see George 1984; Pascale Chesselet, pers. comm. 2004).

Synonymy: Mesembryanthaceae Burnett, nom. cons.

4. Ruschioideae Schwantes

Wide-band tracheids + [not in basal taxa]; (leaves spiral); bladder cells uncommon; inflorescence distinct; bracts/inflorescence often distinct; hypanthium 0[?], filaments papillate or hairy at base, nectaries often crest-like [lophomorphic], radial or annular, G [(3-)5-15(-25)], inferior, placentation basal or parietal; expanding keels of fruit largely valvar.

Ca 110/ca 1585: Ruschia (290-350), Conophytum (87-290: see Opel 2005a for leaf anatomy, 2005b for a morphological analysis), Lampranthus (180-220), Delosperma (155-165), Phyllobolus (150), Drosanthemum (100-110), Psilocaulon (65), Antimima (6-60). Southern Africa, esp. the western coastal Succulent Karroo (Map: Pascale Chesselet, pers. comm. 2004). [Photos - Flower; Flower]

Apatesieae and Dorotheantheae are successively sister to the remainder of the subfamily, and they are not very speciose. The much more speciose core Ruschioideae - Drosanthemeae and Ruschieae - have a crest-like (lophomorphic) nectary, hygrochastic capsules with a distinctive anatomy (for which, see Kurzweil 2006) that release only a few seeds at a time, wide band tracheids are common (absent in sister taxa), and leaves that are cylindrical or trigonous and succulent, not more or less flattened (Klak et al. 2004; Chesselet et al. 2004) and without the bladder-like epidermal cells of the rest of the family. They also have lost the chloroplast rpoC1 intron (Thiede et al. 2007)- cf. Cactoideae!

For a general account of Lithops, see Cole and Cole (2005).

• Aizoaceae are more or less prostrate herbs that may be recognised by their fleshy, opposite leaves the bases of which are membranous and usually surround the stem and the bladder-like epidermal cells. The flowers either have five perianth members that are green outside and colored inside or numerous linear "petals". There are usually many stamens, a more or less inferior and septate ovary, and a fruit that opens on moistening as the septal or other tissue swells.

Aizoaceae, in particular Mesembryanthemoideae and Ruschioideae, dominate much of the Succulent Karoo of southwestern Africa, making up more than 50% of the species and up to an astounding 90% of the biomass. Variation in growth characters - leaf size and shape, internode elongation, etc. - is considerable (Ihlenfeldt 1994). Although a distinction is sometimes made between plants with foliaceous bracts or bracteoles in which the inflorescence is not distinct from the rest of the plant, and plants with smaller bracts and distinct inflorescences (e.g. Hartmann 1993), it is unclear to me what the real growth characters are and where they go on the tree. Many taxa consist of paired leaves more or less flush with the surface of the ground, and these can be almost invisible in the stony habitats in which they grow, being greyish or brownish and looking like pebbles themselves - hence "flowering stones". The exposed surfaces of the leaves sometimes has distinctive "windows"; these leaves are bracteoles, the flower is terminal, and shoots develop in the axils of the bacteoles so giving rise to the next flowering unit (Hartmann 2004, 2006 for a summary). Indeed, in some species of Conophytum the leaves are almost completely connate except for a slit across the top out of which the flower, etc., appear. The bladder-like cells on the leaf surface ("idioblasts") may be involved with water uptake from dew or mist; other taxa may have massively-thickened outer walls that contain layers of calcium oxalate crystals (e.g. Ihlenfeldt & Hartmann 1982). In addition, individual cells may be variously papillate or the surface otherwise sculpted and/or with epicuticular waxes, the stomatal openings may be deeply sunken, etc. (e.g. Ihlenfeldt & Hartmann 1982; Hartmann 2002; Opel 2005a). Hartmann (1988) described the intricate morphology of the capsules, which are often hydrochastic, indeed, seed dispersal is by "jet action" using the kinetic energy of falling raindrops (ombro[hydro]chory: Parolin 2006). There is considerable variation in dispersal and establishment "strategies". Interestingly, Klak et al. (2004) suggest that the radiation in Ruschioideae in S.W. Africa, at least, is both recent (3.8-8.7 million years before present) and very fast, indeed, the "meganiche" dominated by the whole family there - arid winter rainfall area with moderate temperatures - is only some 5 million years old (Ihlenfeldt 1994). Edaphic specialization - soils can vary considerably locally - may be involved in the diversification of Aizoaceae (Ellis & Weis 2006).

I follow Klak et al. (2003) for basic groupings in the family; Aizoaceae s. str. (e.g. Chesselet et al. 1995) would seem to be paraphyletic. Both species and generic limits are difficult. In the early twentieth century Mesembryanthemum included the whole of the Ruschioideae and Mesembryanthemoideae, and until recently the Mesembryanthemoideae, by far the smaller of the two subfamilies, was divided into numerous genera. However, Klak et al. (2007) in a comprehensive study of the subfamily, obtained quite detailed phylogenetic resolution within it - however, Mesembryanthemum itself, although quite a small genus, was polyphyletic, and any attempt to maintain current genera would have caused the recognition of numerous and often poorly characterised taxa; only one genus was recognised. Hammer in 1993 noted that there were then about 1,800 known populations of Conophytum (Ruschioideae) - for which there were 450 names.

Studies of the wood anatomy of Aizooideae and Sesuvioideae are needed to clarify wood evolution in the family (see Carlquist 2007a). Wide-band tracheid pith cells in succulents (Aizoaceae, Cactaceae, Portulacaceae) are also found in the leaf away from the midrib in Aizoaceae; bands are narrow but very tall (= "wide"), so the cell lumen is locally very narrow (Mauseth et al. 1995 - similar in Hectorella - Carlquist 1998b). Studies of the perianth in Sesuvioideae show that the petaloid basal part is equivalent to a sheathing leaf base, and the apical "horn" the rest of the leaf, rather as in monocot leaf development (cf. Vorlaüferspitze!); B floral genes were not expressed, although they were in the petaloid staminodes of Ruschioideae (Frohlich et al. 2007). The androecium may arise as a ring meristem or as five separate primordia. Smets (1986) records the presence of a receptacular nectary disc. Hartmann (1993) noted that a nucellar cap occurs in the family, but under this term he described the radially elongated cells of the nucellar epidernmis that are found in several other core Caryophyllales. Aptenia has a wet stigma.

For more information, see Schwantes (1957: esp. fruit dehiscence), Hegnauer (1964, 1989: chemistry), Hofmann (1973: morphology), Haas (1976: esp. flower and fruit), Bittrich (1986: general, esp. Mesembryanthemoideae), Hartmann (1993), Landrum (2001: wide band tracheids), Chesselet et al. (1995, 2002: esp. information on Mesembryanthemoideae and Ruschioideae) and Interactive Mesembs. The books edited by Hartmann (2001a, b) include thousands of photographs.

Sarcobataceae + Phytolaccaceae + Nyctaginaceae: ORF 2280 sequence similarity, 210 bp deletion in chloroplast genome.

Note that Phytolaccaceae - Rivinioideae and Nyctaginaceae have gynoecia made up of a single carpel (Cuénoud et al. 2002), but if Sarcobataceae is placed somewhere around here, perhaps even within the already polymorphic Phytolaccaceae, its carpel number is a reversal... Family limits in this area may need adjustment; Douglas and Manos (2007) found only moderate support for the monophyly of Nyctaginaceae and vanishing little support for the monophyly of Phytolaccaceae (including Sarcobataceae).

SARCOBATACEAE Behnke   Back to Caryophyllales

Thorny shrub; cork etc.?; wood rayless; ?stomata; plant monoecious, bracteoles 0; staminate inflorescence catkinate, flowers with peltate scales ["bracts"], A 1-4, anthers long, pollen pantoporate, pore margins raised; carpellate flowers single, P tubular, bilobed [?bracteoles], G [2], [?position], ovule 1, funicle?; embryo green; n = 9.

1/2. S.W. North America (Map: from Fl. N. Am. 4: 2003). [Photos - Collection]

Ex Chenopodiaceae, but sieve tube plastids with globular inclusions, etc., suggest that it goes somewhere here. Is it really worth a separate family (cf. Behnke 1997)?

Some information is taken from Carlquist (2000a).

Phytolaccaceae + Nyctaginaceae: cork subepidermal; stomata also paracytic; 1 basal ovule/carpel; protein bodies in nuclei.

PHYTOLACCACEAE R. Brown, nom. cons.   Back to Caryophyllales

Herbs, vines (small trees); styloids and raphides +; cuticular waxes as platelets; leaves (opposite), conduplicate, (?stipules +); inflorescences (leaf-opposed), ± racemose; P 4-5(-10), A 2x P (= P, many, centrifugal), G (1)[-16], often pseudapocarpous, nucellar cap +, funicle?, (obturator +), styles ± gynobasic (style +); P and A persistent in fruit [?level]; embryo white; n = 9.

18[list]/65 - three groups below. Tropical and warm temperate, esp. America (Map: see George 1984; Fl. N. Am. 4: 2003). [Photos - Collection]

1. Phytolaccoideae

Fibers vasicentric; P usu. 5, ovule apotropous; G [3-16]; fruit a berry.

4/31: Phytolacca (25). Chile, Mexico, or cosmopolitan (Phytolacca).

The carpels are initiated in a ring around the apex of the axis (Zheng et al. 2004).

Synonymy: Sarcocaceae Adanson

2. Rivinioideae Nowicke

Habit various, inc. thorny trees; styloids, elongate crystals +; P usu. 4, (pollen pantoporate - Petiveria), G 1, stigma capitate [?always]; fruit various, inc. samaras, indehiscent.

9/13. Central and South America, Antilles, Florida, tropical Africa (some Hilleria), Australia, New Hebrides and New Caledonia (Monococcus).

Gallesia smells of onions.

Synonymy: Hilleriaceae Nakai, Petiveriaceae C. Agardh (nucellar beak developed; Monococcus and Petiveria have P4, diagonal), Riviniaceae C. Agardh, Seguieriaceae Nakai

3. Agdestidoideae Nowicke

Liane; diffuse axial parencyma, true tracheids +; wood rayless; Ca oxalate?; cuticle waxes with ± rounded platelets; inflorescence branches cymose; P 4 (5), A 12-16(25), G [(3-)4], seminferior; fruit a 1-seeded achene with sepalline wings

1/1: Agdestis clematidea. S. U.S.A. to Nicaragua (Map: from Fl. N. Am. 4: 2003; Culham 2007).

Synonymy: Agdestidaceae Nakai

• Phytolaccaceae can be recognised by their racemose inflorescences of apetalous flowers which have many, connate carpels - or a single carpel - each with a single basal ovule and a style; the fruit is often a berry. The plants are herbs, trees, or lianes, and the leaves tend to be rather thick and fleshy, lacking prominent fine venation.

Fossil fruits from the Upper Cretaceous (late Campanian