EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; 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 megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, axillary buds +[?], prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

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

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous... For other features such a a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), where on the tree a thicker nucellus and a stylar epidermal layer are acquired has not yet been indicated.

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels + [one position], elements with elongated scalariform perforation plates; axial parenchyma diffuse or diffuse-in-aggregate; tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

ASPARAGALES + COMMELINIDS: ?

The age of this clade has been estimated at 118-116 million years before present (Bremer 2000b; Leebens-Mack et al. 2005).

ASPARAGALES Bromhead  Main Tree, Synapomorphies.

Chelidonic acid +; (velamen +); (dimorphic root hypodermal cells +); anthers longer than wide, tapetal cells bi- to tetranuclear, microsporogenesis simultaneous; seeds exotestal, tegmen not persistent; endosperm helobial; mitochondrial sdh3 gene lost. - 14 families, 1122 genera, 26070 species.

Evolution. Stem group Asparagales are dated to ca 122 million years before present, crown group Asparagales to ca 119 million years before present (Janssen & Bremer 2004), although Wikström et al. (2001) had suggested dates of 107-98 and 101-94 million years before present respectively. The topology within Asparagales, especially near the base, in the latter study differs substantially from that used here, while Janssen and Bremer (2004), although not putting Orchidales sister to the rest of the order, placed it (in terms of time) near the beginning of divergence within it, and the topology of their tree also differs considerably in detail from that below. Comparable figures in Magallón and Castillo (2009) are ca 133.1 (stem) and 125 (crown) and 118.6 (stem) and 112.6 (crown) million years for relaxed and constrained penalized likelihood datings respectively.

Chemistry, Morphology, etc. Asparagales commonly have Arum-type arbuscular mycorrhizae, while in Liliales these mycorrhizae are commonly Paris-type (see F. A. Smith & Smith 1997). Three-trace tepals are found in Orchidaceae, Amaryllidaceae-Amaryllidoideae, Iridaceae, Asphodelaceae (but not Kniphofia, Ashpodelus), Asparagaceae-Agavoideae, Amaryllidaceae-Agapanthoideae, and Hemerocallidaceae; one-trace tepals in Ruscaceae (but not Maianthemum stellatum), Amaryllidaceae-Allioideae, Aphyllanthoideae, and Asparagoideae. Scilloideae have tepals with both kinds of vasculature, Urginea even having five traces in the outer whorls (see especially Chatin 1920). Where changes in microsporogenesis are to be placed on the tree is not clear. Furthermore, Rudall (2001a, see also 2002, 2003a) included an inferior ovary as a synapomorphy of the order, noting that in "higher" Asparagales there may well be a reversal to superior ovaries that is associated with the presence of infralocular septal nectaries (as in Xanthorrhoea and Johnsonia (Hemerocallidaceae)). However, since superior ovaries are also scattered through the "lower" Asparagales, where the evolution of different ovary morphologies are to be placed in the tree is unclear; ovary position seems a much more flexible character here (and elsewhere) than it has generally been given credit for.

For flavonoids, see Williams et al. 1988), for ovule and seed, see Shamrov (1999a) and Oganezova (2000a, b), for cytology, see Tamura (1995), for root morphology, see Kauff et al. (2000), for general morphology, see Rudall (2003a), for pollen of Japanese representatives, see Handa et al. (2001), for cytology and genome size, see Pires et al. (2006), and for the distribution of taxa with phytomelan and/or with baccate fruits, see Rasmussen et al. (2006).

Phylogeny. The tree is based largely on the analyses in Chase et al. (2000a) and Fay et al. (2000: successive weighting). These studies differ little in detail, although the analysis of Fay et al. (2000) hardly suprisingly had more nodes in the core Asparagales with strong support. For the Amaryllidaceae + Agapanthaceae node, see Meerow et al. (2000b); relationships between Aphyllanthaceae, Themidaceae and Hyacinthaceae might be better represented as trichotomy. A phylogeny presented by McPherson and Graham (2001) is largely congruent with this, although its sampling is much poorer. Understanding the relationships of Boryaceae and Orchidaceae is critical. Boryaceae have sometimes been placed as sister to Orchidaceae(e.g. Chase et al. 1995a; McPherson & Graham 2001), although with rather weak support, and there are other topologies, including the embedding of Orchidaceae in a paraphyletic Boryaceae-Hypoxidaceae clade (Li & Zhou: 2007: little support).

Recent work suggests that Orchidaceae are sister to other Asparagales (e.g. 76% bootstrap support in Graham et al. 2006; about the same in Givnish et al. 2006b; stronger [96-99%] in Pires et al. 2006, good sampling and seven genes from two compartments), and this is associated with more or less strong support for Boryaceae being sister to the Blandfordiaceae et al. clade. Rudall (2003a) had also suggested a close morphological relationship between Boryaceae and Blandfordiaceae. Note, however, while there is good support in Chase et al. (2006) for the position of Orchidaceae as sister to all other Asparagales, Boryaceae are placed immediately above the Blandfordiaceae et al. clade, albeit with very little support. Some phylogenetic reconstructions of Hilu et al. (2003: molecular data) had suggested Asparagales might be paraphyletic, with Orchidaceae separate from the rest. Rudall (2003a: morphological data) suggested that there was a close morphological relationship between Hypoxidaceae and Orchidaceae in particular. All in all the position of Orchidaceae as sister to the rest of Asparagales, with Boryaceae being part of the Blandfordiaceae et al. clade, seems the best hypothesis, and this hypothesis of relationships is adopted here. Note that this rather changes the characterisation of Asparagales, some characters previously considered to characterise the clade as a whole now being best moved to the subbasal node in the clade (cf. versions 7 and younger of this site). Diversification in Orchidaceae is indeed remarkable, but remember it has to be compared with that of all other Asparagales, somewhat less species-rich, perhaps, but a morphologically rather motley crew (see also below, under Orchidaceae).

Previous Relationships. Dahlgren et al. (1985) - basically about the time our understanding of relationships in monocots began to be seriously re-evaluated - recognised two major groups into which they placed most of the "lily-like" monocots. Although corresponding in part to the Asparagales and Liliales as recognised here, and divided by features including patterning of the tepals, absence of phytomelan (both features of their Liliales), etc., both Iridaceae and Orchidaceae were included along with families here included in Liliales. More recently, genera of Boryaceae have often been included in Anthericaceae, as by Takhtajan (1997).

Includes Amaryllidaceae, Asparagaceae, Asteliaceae, Blandfordiaceae, Boryaceae, Doryanthaceae, Hypoxidaceae, Iridaceae, Ixioliriaceae, Lanariaceae, Orchidaceae, Tecophilaeaceae, Xanthorrhoeaceae, Xeronemaceae.

Synonymy: Asparagineae J. Presl, Asphodelineae Thorne & Reveal, Hyacinthineae Link, Iridineae Engler - Agavales Hutchinson, Alliales Traub, Amaryllidales Bromhead, Apostasiales Blume, Asphodelales Doweld, Asteliales Dumortier, Gilliesiales Lindley, Hypoxidales Reveal & Doweld, Iridales Rafinesque, Ixiales Lindley, Narcissales Dumortier, Orchidales Rafinesque, Tecophilaeales Reveal, Xanthorrhoeales Reveal & Doweld - Iridanae Doweld, Orchidanae Doweld - Orchididae Heintze - Crinopsida Horaninov, Orchidopsida Bartling

ORCHIDACEAE Jussieu, nom. cons.   Back to Asparagales

Mycorrhizal herbs, protocorms mycoheterotrophic, root hairs often lacking [?distribution]; flavone C-glycosides, flavonols +, chelidonic acid?; SiO₂ bodies [stegmata] in leaf vascular bundle sheaths, these sheaths with fibres, (also fibre bundles in leaves); inflorescence racemose; flowers monosymmetric, resupinate, T free; A 3 [median of outer whorl and laterals of inner whorl], basally adnate to style, tapetal cells uninucleate, septal nectaries 0, ovary inferior, placentae branched, ca 1500+ tenuinucellate ovules/carpel, funicle not vascularised, style solid, stigma wet; seeds minute, phytomelan 0; fruit splitting down its sides, T deciduous; endosperm barely developing, embryo minute, undifferentiated, haustorial suspensor +.

880[list]/22,075 - five subfamilies below. World-wide. [Photo - Flower]

1. Apostasioideae Horaninov

Chemistry unknown; vessel elements simple; stomata tetracytic; leaves spiral, plicate; Apostasia not resupinate, T apiculate, carinate, (A 2, staminode +/0 - Apostasia); pollen reticulate, (operculate), (embryo sac bisporic - Allium type - Neuwiedia), micropyle bistomal; seeds entotestal, exotegmen sclerified [Neuwiedia], outer periclinal walls collapsing; n = 24.

2/16. Sri Lanka, N.E. India to N.E. Australia, Japan (map: from Pridgeon et al. 1999).

Synonymy: Apostasiaceae Lindley, Neuwiediaceae Reveal & Hoogland

Vanilloideae [Cypripedioideae [Orchidoideae + Epidendroideae]]: C-glycosyl flavones, (saponins), 6-hydroxy flavonols +; (velamen +; tilosomes +); leaves spiral or two-ranked; flowers strongly monosymmetric; (tepal nectaries +), labellum strongly differentiated, the style and A almost completely congenitally fused [gynostemium], anthers to 2x as broad as long, pollen sticky, placentation parietal, ovules not fully developed at pollination, stigma asymmetrical; fertilisation may take some months; seeds dispersed before maturity of embryo; radicle 0.

2. Vanilloideae Szlachetko

Plant sympodial or monopodial, (echlorophyllous and/or viny); SiO₂ bodies 0; stomata tetracytic; (venation reticulate); calyculus +; T often carinate, (margins of labellum fused with column - some Vanilleae); A 1 [median member of outer whorl], staminodes 2 [from inner whorl], anther incumbent [by massive expansion of the apical column/connective], "pollinia" soft, viscidium 0, (pollen in tetrads), (placentation axile), rostellum [ridge, part of median stigmatic lobe] +; (T persistent; fruit baccate)); seeds fusiform, crustose, exotestal [outer parietal wall well developed], tegmen persisting; endosperm to 16-nucleate; n = 9, 10, 12, 14-16, 18.

15/180: Vanilla (100), Epistephium (12). Pantropical, esp. Asia; Australia, some N. America (map: from Pridgeon et al. 2003).

Synonymy: Vanillaceae Lindley

Cypripedioideae [Orchidoideae + Epidendroideae]: ?

3. Cypripedioideae Kosteletzky

(Stomata paracytic); leaves conduplicate or plicate; 2 abaxial T of outer whorl connate, labellum saccate; A 2 [from inner whorl], staminode 1 [median member of outer whorl], pollen ± psilate (foveolate), operculate, (in tetrads), tapetal cells binucleate, microsporogenesis successive?, (pollen operculate), median stigma lobe largest, (embryo sac bisporic, eight celled [Allium-type]; micropyle bistomal); (T abscising); n = 9 or more.

5/130: Paphiopedilum (62), Cypripedium (50). Mostly (warm) temperate N. hemisphere, East Malesia and tropical South America (S. India) (map: from Hultén 1958; Pridgeon et al. 1999). [Photo - Flower]

Synonymy: Cypripediaceae Lindley

Orchidoideae + Epidendroideae: floral primordium tranversely elliptic-oval; labellum initiated first; A 1 [median member of outer whorl], (staminodes 2 [from inner whorl]), pollen forming pollinia that are variously attached to sticky viscidium, pollinarium stalk variously formed from part of the anther [caudicula], epidermis of the rostellum [tegula] or apex of the rostellum [hamulus], microsporogenesis simultaneous [tetrads tetrahedral], pollen usu. in tetrads, porate or ulcerate, median carpel developed before the others, rostellum [ridge, part of median stigmatic lobe, viscidium is also part of it] +; T not abscising; tegmen not persisting; n = 9 or more [19 common].

4. Orchidoideae Eaton

Plant sympodial; mycoheterotrophs rare, stem/root tuber common; SiO₂ bodies 0; sclerenchyma in leaf [as fibre bundles or associated with vascular bundles] and stem rare; stomata anomocytic; leaves usu. spiral, soft, herbaceous, deciduous; anther erect (incumbent), apex acute, pollinia soft/sectile, staminodes reduced, (hamulus [pollinium stalk from modified apical part of rostellum] +).

208/3755: Habenaria (600), Caladenia (376), Platanthera (200), Pterostylis (200), Disa (175), Cynorkis (125), Orchis (125), Corybas (120), Goodyera (80-100), Satyrium (90), Disperis (85), Prasophyllum (80), Cyclopogon (75), Pelexia (75), Peristylus (75), Zeuxine (70), Diuris (55), Goodyera (55), Holothrix (55), Cheirostylis (50), Dactylorhiza (50), Thelymitra (50). World-wide, esp. temperate (map: from Pridgeon et al. 2001, 2003; distribution in N. Asia and N. North America unclear).

Synonymy: Neottiaceae Horaninow, Limodoraceae Horaninow, Liparidaceae Vines, Ophrydaceae Vines

5. Epidendroideae Kosteletzky

Epiphytes common, (plant without shoots or leaves [Vandeae]; mycoheterotrophic); (SiO₂ bodies 0); velamen + (0), (roots with pneumathodes); stomata often para- or tetracytic; stems thick; leaves usu. distichous, conduplicate (plicate), articulated with sheathing base (not); anther incumbent (strongly convex) [by column elongation, or by early anther bending (vandoids)], with beak, operculate, pollinia hard (sectile), apical part of median stigma lobe forms stipe [= pollinium stalk]; (cotyledon visible).

650/18000: Bulbophyllum (2035), Epidendrum (1500), Dendrobium (1400, but see below), Pleurothallis (1000), Lepanthes (>800), Stelis (700), Oncidium (680, but generic limits?), Masdevallia (410), Liparis (320), Malaxis (300), Maxillaria (ca 300), Crepidium (280), Dendrochilum (265), Calanthe (260), Acianthera (200), Coelogyne (200), Specklinia (200), Angraecum (200), Eulophia (200), Phreatia (190), Oberonia (175), Taeniophyllum (170: major photosynthetic organs are green, flattened roots), Phreatia (160), Pinalia (160), Telipogon (160), Octomeria (150), Dracula (120), Encyclia (120), Dichaea (110), Trichosalpinx (110), Ceratostylis (100), Catasetum (100), Elleanthus (100), Glomera (100), Prostheacea (100), Sobralia (100), Camaridium (80), Platystela (75), Brachionidium (65), Appendicula (60), Callostylis (60), Neottia (60), Nervilia (60), Podochilus (60), Scaphyglottis (60), Sophronitis (60), Ornithidium (55), Lepanthopsis (50), Maxillariella (50), Restrepia (50). More or less world-wide, but most diverse in the tropics; rather poorly developed in Australia (map: from Pridgeon et al. 2005).

• Orchidaceae can be recognised by their strongly monosymmetric flowers in which the 1-3 stamens are at least basally adnate to the style. The flowers are usually held with the median member of the outer tepalline whorl in the adaxial position, while the median member of the inner whorl is often remarkably elaborated and forms a labellum. The ovary is inferior and the fruit, a capsule, opens down the sides to release the very numerous and minute seeds.

• Apostasioideae have 2-3 stamens that are connate only to the base of the style and apiculate, carinate tepals. The other subfamilies have a well developed labellum and the stamen(s) are more or less completely fused with the style, forming a gynostemium.

• Cypripedioideae have two stamens and a deeply saccate labellum. The other subfamilies have but a single stamen. In the other subfamilies the labellum is very variable in shape, but not often deeply saccate. Vanilloideae lack both pollinia and viscidia, while both are present in Epidendroideae and Orchidoideae. The former have rather tough leaves and stems and incumbent anthers, the latter often have softer, deciduous leaves and straight anthers.

Evolution. Stem group Orchidaceae are dated to ca 119 million years before present, crown group Orchidaceae to ca 111 million years before present (Janssen & Bremer 2004), and the minimum ages estimates of Ramírez et al. (2007: calibration by Miocene Goodyerinae pollinaria found in amber) are somewhat younger, 87-76 million years before present. Ramírez et al. (2007) suggest that the subfamilies had diverged by the end of the Cretaceous, ca 65 million years before present, or perhaps slightly later in the early Palaeocene, with diversification in the speciose, epiphytic Epidendroideae being Eocene in age, some 47-40 million years before present (see also Conran et al. 2009a). Wikström et al. (2001) had suggested divergence between the single member of Cypripediodeae and Epidendroideae included in their analysis as being much more recent, only ca 37-36 million years before present.

Note that although Orchidaceae are considered to be very diverse, much of the floral variation is at one level only intricate recombination on a rather limited theme. Orchidaceae are sister to the rest of Asparagales which are also morphologically very diverse and include some 6,850 species, and Asparagales as a whole are sister to commelinids, with some 22,750 species. Numbers and sister-group relationships alone are not yet telling much of a story, indeed, we still know little about the origin and biogeography of the family (see also Chase 2003 for a convenient summary). Normally neither orchids nor pollinating insects are diverse on oceanic islands, but angraecoid orchids are surprisingly diverse on the Mascarene islands, and Reunion in particular also has a diverse insect fauna (Micheneau et al. 2008).

Members of Orchidaceae are not often eaten by butterfly caterpillars (Janz & Nylin 1998) or by insect herbivores in general, although Riodininae-Riodininae larvae may be found on them (Hall 2003 and references).

Some of the distinctive features of the family seem to be biologically connected. Thus pollinia ensure the fertilization of numerous ovules; the minute seeds to which they give rise, usually devoid of endosperm or differentiated embryo, are well suited for wind dispersal; and the obligate myco-heterotrophy of the young plant may compensate for the absence of seed reserves (Johnson & Edwards 2000 in part). Gravendeel et al. (2004 and references; see also Peakall 2007) list the numerous hypotheses that have been advanced to explain the diversity of Orchidaceae; these include pollinator specialization, niche partitioning, habitat fragmentation, wide dispersal of orchid seeds, etc.. Tremblay et al. (2005) reviewed the evolutionary consequences of variation in sexual reproduction in orchids. Orchidaceae are of course noted for the diversity of pollination mechanisms they show, and orchid diversification is often explained in terms of the close association between pollinators and individual species of orchids. For a summary of pollination in Orchidaceae, see van der Cingel (1995, 2001) - and of course the classic study by Darwin (1862) is still worth reading.

Orchid flowers may be notably long-lived (months!), although some last only for a single day. Flowers of Orchidaceae are commonly resupinate, the ovary being twisted about 180°, the labellum ending up in the abaxial position. However, the amount of resupination often varies within a plant, especially when the inflorescence is arching; here all flowers of the inflorescence are often oriented so that their labellum is in the same position with respect to gravity, with the ovary sometimes being twisted 360° (as in Angraecum, etc.) or not at all. Fischer et al. (2007) discuss the variety of ways - of which twisting of the pedicel is but one mechanism involved - that flowers in the speciose Bulbophyllum present themselves. In other orchids such as Calopogon the flowers are never resupinate, and all flowers on the erect inflorescence show "normal" monocot orientation, the labellum here being adaxial. In the dioecious Catasetum resupination varies between staminate (resupinate) and carpellate (not resupinate) flowers; because of this and other striking differences, especially in labellum morphology, staminate and carpellate specimens were once put in separate genera, Myanthus and Monachanthus respectively.

The most conspicuous aspect of floral variation is in the labellum, a highly-differentiated member of the inner whorl of tepals, which shows a diversity of form and color that is truly remarkable; duplication of genes may be involved in this (Mondragón-Palomino & Theißen 2008). The spatial relationships of the labellum and column in particular force the pollinator to approach the flower in a particular way, and in general, the pollinaria are very precisely placed on the pollinator, closely related species differing in exactly where on the animal they are placed (e.g. Maad & Nilsson 2004). There is quite often movement of the pollinia after attachment to the pollinator to bring them into the proper position for pollination (for pollinia, see Selbyana 29: 1-86. 2008, and references). But as suggested above and below, other factors may have facilitated the diversification of Orchidaceae. Although artificial hybrids involving three or more genera have been reported, how these will look when generic boundaries are redrawn is unclear, nevertheless, many Orchidaceae can be crossed artificially, perhaps because of the presence of well-developed and effective premating barriers and hence the lack of any need for postmating barriers. However, it has been suggested (based on work on European orchids) that in those orchids with generalized food-deceptive mating mechanisms, barriers to crossing may in fact be postzygotic, whereas those that practice sexual deception have premating reproductive barriers (Cozzolino & Scopece 2008).

The plesiomorphic condition for the family may be to lack nectar (Jersáková et al. 2006), pollen alone being collected from flowers of Apostasioideae; all told, some 8,000 species of orchids lack nectar altogether. However, nectaries do occur in Orchidaceae, and they vary in position (Davies et al. 2005) although they are never septal; nectar is in fact a common reward (Bernadello et al. 2007 and references). In a number of species of Maxillaria hairs on the labellum contain protein and perhaps also starch and function as pseudopollen, so rewarding the pollinator (Davies et al. 2000; Davies 2009).

Cozzolini and Widmer (2005) suggested more particularly that orchid diversification is associated with the deceptive pollination that is so prevalent in the family; about one third of the species - some 6500 or so - are pollinated in this way (Schlüter & Schiestl 2008 for molecular mechanisms; Peakall 2009 for deceit and speciation). Interestingly, recent analyses suggest that when pollinators visit orchid flowers in the course of deceptive pollination or to pick up scent rewards, pollinator specifity is greater and species richness was greater than when pollinators visit for nectar (Schiestl & Schlüter 2009). Thus deceit pollination may under certain situations increase outcrossing and speciation, the latter perhaps because of the specificity of the pheromones produced by the plants (Jersáková et al. 2006; see also Ledford 2007). Perhaps 60% of orchids are pollinated by bees (Schoonhoven et al. 2005), deceived or otherwise. The European Ophrys (Orchidoideae) is well known for its distinctive and variable labellum and production of chemicals like pheromones which together allow the flower to mimick female insects: pollination is by pseudocopulation by bees and wasps in particular. Alkenes (hydrocarbons with at least one double bond) are involved in the chemical component in this mimickry (e.g. Stökl et al. 2009), and such chemicals are common in related genera as well (Schiestl & Cozzolino 2008). (Note that current estimates of species numbers in Ophrys range from 16 to 215, but the former number seems much closer to "reality" [Bateman et al. 2006a; Devey et al. 2008].) Flowers of Serapias (Bellusci et al. 2008 for a phylogeny) may even attract pollinators by mimicking a nest hole. Within Epidendroideae, pollination during pseudocopulation with fungus gnats has recently been reported in the large genus Lepanthes (Blanco & Barboza 2005).

Orchids that secrete oil in their flowers show convergence with the flowers of other oil-pollinated plants. Thus pollination of some South African Orchidoideae-Coryciinae is by oil-collecting bees; the flowers have paired, pouch-like structures like those of another local oil plant, Diascia (Scrophulariaceae: Pauw 2006). Moreover, the distinctive lip-like appendage that was a defining feature of the old Corycinae (Waterman et al. 2009) seems to have evolved in parallel. Orchids in Oncidiinae have elaiophores, sometimes on the labellum, and may be mimics of Malpighiaceae, both groups being visited by bees like Centris, etc. (Stpiczynaska & Davies 2008; Chase et al. 2009). Examples of butterfly, moth and even bird pollination are also well known. Thus the corolla tube of Angraecum sesquipedale, from Madagascar, is ca 30 cm long, and its pollinator for long remained unknown, although Darwin (1862) suggested that some moth with a proboscis that long would be found; Xanthopus morgani praedicta, with a proboscis length of ca 25 cm, was subsequently discovered (Nilsson et al. 1987; Nilsson 1988). For spurs, nectariferous and otherwise, in Orchidoideae-Orchidinae, see Bell et al. (2009).

Williams (1982) discussed the general importance of male euglossine bees in orchid (Epidendroideae) pollination in the neotropics; male bees searching for fragrances are usually involved, and the some 190 species of bees pollinate perhaps up to 25% of tropical American Orchidaceae, hence their common name, orchid bees. Pollination occurs especially in orchids growing at lower altitudes, and anywhere from >700-2,000 species may be so pollinated (Cameron 2004 and references [Photo: Pollinators.]; Zimmermann et al. 2009), not to mention hundreds of species of Zingiberales, Gesneriaceae, Lecythidaceae, etc. Although effective pollinators, the relationships between orchids and euglossine bees are non-specific on both sides (Cameron 2004). Closely related species of sympatric species of Euglossa showed greater disparity in the fragrances they had collected and stored in their hind tibial pockets than might be expected, the distinctive fragrance signals of males perhaps being involved in pre-mating isolation in the bees; the most dominant compounds in these fragrances were highly homoplasious (Zimmermann et al. 2009). The pollination of Catasetinae by male euglossine bees which visit the flowers for fragrance compounds is well known (Darwin 1862; Chase & Hills 1992 for a phylogeny).

A final distinctive feature of many orchid flowers is that the ovules are usually not fully developed at anthesis. There are many reports that fertilisation may be delayed relative to pollination, as in Cypripedium and a number of other genera; the time between pollination and fertilization ranges from four days to ten months (in Vanda), the normal time being one week to six months (Wirth & Withner 1959, also Sogo & Tobe 2005, 2006d for references: Fagales show the same correlation. ?Situation in Apostasioideae). Even after fertilization, it may be a month before embryo development begins, as in Sarcanthinae (Wirth & Withner 1959 for references).

There is a close association between basidiomycete - and some ascomycete - fungi and orchids. Rhizoctonia (= Ceratobasidium) is a common anamorph or form genus, and Russulaceae, Tuber, and Sebacinales B (autotrophic orchids) and B (mixotrophic and myco-heterotrophic orchids) are all involved; the most common families are Tuslasnellaceae and Ceratobasidiaceae and Sebacinaceae (see Currah et al. 1997 and Yukawa et al. 2009 for a list of the fungi; Otero et al. 2002; Roy & Selosse 2009; Weiß et al. 2009). This association starts on the germination of the orchid seed. At this stage sugars and nitrogen move from the fungus to the orchid (Zimmer et al. 2007), and the initial fungus-plant association results in a protocorm (Peterson et al. 1998). However, some orchids can be made to germinate in the absence of a fungus. The fungi form an intracellular mycorrhizal association and are basically modified ectomycorrhizae (Smith & Read 1997), and at least in some cases may also be ectomycorrhizal on forest trees (Bidartondo & Read 2008, see also below). It is perhaps a reflection of this association that Orchidaceae have particularly small seeds when compared with their immediate relatives (Moles et al. 2005a), the seeds being as little as 150 µm long or less. The specificity of the mycorrhizal association with fungi may be connected with the diversification of the family (Otero & Flanagan 2006), although the relationship is by no means one-on-one. Obligate mycoheterotrophy is not uncommon, a number of orchids having independently become achlorophyllous mycoheterotrophs; this may have happened some 20 times in the family (Molvray et al. 2000), and the fungi involved are different. The Australian Rhizanthella (Orchidoideae) is a subterranean mycoheterotroph, the flowers even opening underground. In Corallorhiza (Epidendroideae) the fungi involved are several species of Russula which form an ectomycorrhizal association with adjacent trees and an endomycorrhizal association with the orchid (Taylor & Bruns 1999). However, bi- or unidirectional (from fungus to orchid) movement of carbon has been detected even in chlorophyllous orchids (Cameron et al. 2008; Tsujita et al. 2009; Hynson et al. 2009a), although of course in mycoheterotrophic orchids carbon flow is unidirectional. It is likely that the fungi involved in orchid symbioses are in fact saprophytes living on decaying plant material that can also form close relationships with orchids (Ogura-Tsujita et al. 2009; Yukawa et al. 2009).

Details of the fungus-orchid association were until recently unclear in Apostasioideae, although the basiomycete Tulasniella was involved, and this genus is also found in Cypripedium, etc. (Kristiansen et al. 2004); Yukawa et al. (2009) have studied the associates of Apostasia in some detail. Stomatiferous root tubercules form in Apostasia as a result of the association with the fungus, and it has been suggested that these may make the plant better able to deal with wet conditions (Stern & Warcup 1994); seeds of Apostasioideae are rather larger than those of other orchids. Interestingly, the Australian Boryaceae also have a plant-fungus association - details?

It is often forgotten that Orchidaceae show considerable diversity in habit and other vegetative features despite their generally modest size; Tatarenko (2007) outlines the extensive vegetative variation of temperate orchids, i.e. especially Orchidoideae. The epiphytic habitat is particularly favoured by Epidendroideae, and about 70% of all Orchidaceae are epiphytes, and there are more species of epiphytic orchids than of all other vascular epiphytes combined (Benzing 1983). Indeed, speciation in Orchidaceae may increase in epiphytic clades, e.g. in Epidendroideae-Bulbophyllinae (Gravendeel et al. 2004). In the epiphytic habitat orchids have to deal with periodic drought and lack of nutrients (Gravendeel et al. 2004, see also Motomura et al. 2008); twig epiphytes in particular grow in extreme conditions, and some minute twig epiphytes of the New World Epidendroideae-Oncidiinae mature within a year (Chase 1987; Chase and Palmer 1997). In Epidendroideae-Vandeae, taxa that lack photosynthetic leaves or have very deciduous leaves but have photosynthetic roots (rarely the stem is photosynthetic) have evolved several times. The aptly named Taeniophyllum, with its distinctive flattened photosynthetic roots, is one example (e.g. Carlsward et al. 2006b); all told, over 200 species, all epiphytes, are involved. (These plants, and ?many other Epidendroideae, lack roots hairs, and the roots themselves are commonly rather fat. Root hairs ["rhizoids"] are sometimes described as being branched - Rasmussen 1999.) In addition to the pneumathodes common in Epidendroideae, these Vandeae have aeration units, made up of distinctive exodermal cells, a space beneath, and a pair of thin-walled cortical cells, and such aeration units are also found in related leafy Vandeae (Benzing et al. 1983; Carlsward et al. 2006a, b). A variant of crassulacean acid metabolism (CAM) photosynthesis is common among epiphytes (see Cameron et al. 2008 for Oncidiinae), and the adoption of CAM by epiphytic Epidendroideae growing at low altitudes has been associated with Tertiary radiation of that subfamily (Slivera et al. 2009); CAM has evolved perhaps ten times in the family, and has also reversed to C₃ photosynthesis. How carbon dioxide and water flux are controlled in epiphytes is unclear, especially because there are no stomata, although the aeration units may be stomata analogues (Benzing et al. 1983; Cockburn et al. 1985). Indeed, many of the epiphytic Epidendroideae, perhaps some 7,000 species, are likely to have CAM (Winter & Smith 1996b). (Note that taxa like Malaxis and Liparis - Epidendroideae - may be secondarily terrestrial.) New World epiphytic Epidendroideae in particular have distinctive tilosomes, cells of the innermost layer of the velamen that are adjacent to the passage cells of the exodermis and that have complex often lignified excrescences developing from the wall (Pridgeon et al. 1983), but such cells are also found in ground-dwelling Orchidoideae-Spiranthinae (Figueroa et al. 2008). Extrafloral nectaries are scattered in the family, and they can be found on the stems opposite the leaves in Vanilla, at the bases of the pedicels in Cymbidium, etc.

Chemistry, morphology, etc. For the anatomy of Apostasioideae, see Stern et al. (1993); the subfamily is poorly known.

Zygomorphy of the flower in many, but not all orchids - and in Hypoxidaceae and Doryanthaceae - is evident even in the earliest primordia (Kurzweil & Kocyan 2002 and references). A few Orchidaceae have more or less polysymmetric flowers, and in Telipogon (Epidendroideae - Oncidiineae) a polysymmetric perianth becomes evident only late in development (Pabón-Mora & González 2008). The sequence of organ initiation varies considerably within the family (Pabón-Mora & González 2008). Apostasioideae and Cypripedioideae have simultaneous initiation of members of the inner tepal whorl, the plesiomorphic condition for Asparagales (Kocyan & Endress 2001a); have Vanilloideae been studied? At least some Orchidaceae have placentoids (Weberling 1989). Prutch and Schill (2000) discuss variation in the morphology and ultrastructure of the stigma; variation seems to be at about the subfamilial level. Although the seeds are geneally minute and the testa cells have thin walls, Selenipedium (Cypripedioideae) has a hard, dark testa, although apparently it lacks phytomelan. There is much variation in chromosome number and size. Thus Apostasioideae and Orchidoideae have small chromosomes, while larger chromosomes occur in Cypripedioideae and Vanillioideae. matK in Apostasioideae may be in transition from a possibly functional gene to a pseudogene; in the other members of the family examined (but the sampling is poor) it is a pseudogene (Kocyan et al. 2004).

Additional information is taken from Swamy (1948: floral vasculature), Wirth and Withner (1959: embryology and development), van der Pijl and Dodson (1966: pollination), Rasmussen (1982: pollinarium morphology), Newton and Williams (1978: Cypripedioideae, Apostasioideae pollen), Schill and Pfeiffer (1977: pollen, general), Johnson and Edwards (2000: pollinia morphology), Pacini and Hesse (2002: pollen units), Freudenstein and Rasmussen (1997: sectile pollinia, pollinia that separate into groups of pollen grains), Pridgeon et al. (1983: tilosomes), Porembski and Barthlott (1988: velamen), Schlechter (1992, 1996, 2003: general), Stern et al. (1993b: vegetative anatomy), Dressler (1993: general), Endress (1994b: floral morphology), Freudenstein and Rasmussen (1999: morphological phylogeny), Pridgeon et al. (1999), Rasmussen (1999: terrestrial orchids), Kurzweil (2000), Molvray et al. (2000), Cameron and Chase (2000), Szlachetko and Rutkowski (2000: gynostemium), Kocyan and Endress (2001a: floral development), Kristiansen et al. (2001), Johansen and Frederiksen (2002: flowers), Cameron (2002), Szlachetko and Margonska (2002: gynostemium), Kurzweil (esp. 1987, 1993: floral development), Kurzweil and Kocyan (2002: a general survey of floral development), Freudenstein et al. (2002: pollinium development, 2004: phylogeny), Prychid et al. (2004: SiO₂ bodies), and Yeung (2005: embryogeny). For reticulate venation in Vanilloideae, see Cameron and Dickison (1998), for general information, see Pridgeon et al. (2003). For information on Orchidoideae, see Stern (1997a, b: anatomy), Stern et al. (1993a: anatomy), Pridgeon et al. (2001a, 2003: general), and Bell et al. (2009: nectar spurs), while for information on Epidendroideae, see Stern and Carlsward (2009: anatomy of Laeliinae).

Phylogeny. Cameron (2007) provides a summary of phylogenetic studies on the family. There is still some uncertainty over the position of Cypripedioideae. For instance, they may group (albeit weakly) with Vanilloideae (Freudenstein & Chase 2001) or be sister to Orchidaceae minus Apostasioideae, which would appear to make sense from the point of view of androecial evolution (Cameron et al. 1999, one gene, successive weighting). They may also be sister to Orchidaceae minus Apostasioideae and Vanilloideae (e.g. Kocyan et al. 2004; Cameron & Chase 2000; Cameron 2002, 2005b, 2006 [two genes; this study places them in a basal trichotomy in the family with atp alone]). This latter hypothesis is followed here, and it suggests that the monandrous condition may have evolved twice (see also Freudenstein et al. 2002). (A conservative topology might be that of a polychotomy [as found by Cameron 2004].) Furthermore, there are suggestions that Codonorchis is either sister to [Epidendroideae + Orchidoideae] (e.g. Clements et al. 2002) or basal in Orchidoideae (Cameron 2006), and perhaps should be recognised as a subfamily, Codonorchidoideae (it has whorled leaves - see Cameron 2006).

For a phylogeny of Apostasioideae, see Kocyan et al. (2004). Relationships within Vanilloideae are becoming fairly well resolved (Cameron 2004, 2009; Cameron & Molina 2006; Pansarin et al. 2008; ). Included are Pogoniinae (Erythrorchis - mycoheterotroph), Vanillinae, and Galeolinae and Lecanorchidinae (both mycoheterotrophs). For relationships in Cypripedioideae, including also a morphological survey, see Albert (1994).

Relationships within Orchidoideae are becoming fairly well resolved (e.g. Cameron 2004); see Górniak et al. (2006) for Spiranthinae. Clemens et al. (2002) clarify relationships of the Diuridae, a few of which are to be placed in Epidendroideae; for Codonorchis, see above. Orchidoideae include the erstwhile Spiranthoideae; there the anther is incumbent (as in Epidendroideae) and with apical rostellar tisssue. Disa is especially diverse in the Cape Region (see Bytebeier et al. 2007, 2008); for a phylogeny of Satyrium, see van der Niet and Linder (2008); it has diversified in the Fynbos region (Verboom et al. 2009). Prescottiinae s.l. have diversified at very high altitudes - to 4,900 m - in the Andes (Álvarez-Molina & Cameron 2009). For information about the speciose Caladenia, see Australian J. Bot. 57(4). 2009.

For general phylogenetic relationships in Epidendroideae, see van den Berg (2005), and for a general account of the subfamily, see Pridgeon et al. (2005). Support for branching along the spine of Epidendroideae is not strong (e.g. Cameron et al. 1997; Pridgeon et al. 2001b; Cameron 2004). Palmorchis may be sister to Neottieae, the combined clade being in turn sister to all other Epidendroideae (Rothacker & Freudenstein 2006). It has been suggested that "basal" clades tend to lack articulated leaves, they have no velamen, and the pollinia are sectile (Pridgeon et al. 2005), and this will clearly affect identification of apomorphies for the subfamily. Malaxis and Liparis may not be monophyletic, but are closely intertwined; it is likely that they are secondarily terrestrial. Dendrobium is turning out to be polyphyletic (Yukawa & Uehara 1996 [orchids in this area are vegtatively variable, florally less so], Yukawa et al. 1993, 1996, 2000; Clements 2003 and earlier work, a mix of revisionary studies and phylogeny). For a major study of Pleurothallidinae, see Pridgeon et al. (2001b); Pleurothallis itself is also not monophyletic; Abele et al. (2005) and Matuszkiewicz and Tukallo (2006) discuss the phylogeny of Masdevallia (Pleurothallidinae). For phylogenetic relationships in Laeliinae, see van den Berg et al. (2000), for diversification in Coelogyninae, see Gravendeel et al. (2005), for studies in Maxillarieae, see Whitten et al. (2000), Williams and Whitten (2003), Sitko et al. (2006), and especially Whitten et al. (2007: Maxillaria to be restricted, generic realignments needed; Blanco et al. 2007: many new combinations) and Stern et al. (2004: anatomy), Oncidiinae, Williams et al. (2001a, b, 2005), Stern and Carlsward (2006: anatomy), Cymbidieae, Whitten et al. (2005), Cieślicka (2006: Eulophia), Neubig et al. (2008: Dichaea, Zygopetalinae), and Chase (1987), Chase and Palmer (1997), and Williams et al. (2001) all dismantling the broadly-delimited and polyphyletic Oncidium, Epidendreae (Kulak et al. 2006), angraecoid orchids in general, (Stewart et al. 2006), and Aeridinae, see Hidayat et al. (2005).

Classification. Chase et al. (2003) provide a higher-level phylogenetic classification for the family, while Govaerts et al. (2003) is a provisional checklist of the family (see also World Checklist of Monocots). For an illustrated account of the genera, see Alrich and Higgins (2008).

Generic limits in the family are in the middle of a major overhaul to make them consistent with molecular findings, since it is clear that there is widespread homoplasy in floral features (e.g. Waterman et al. 2009; Chase et al. 2009 and references). Floral differences have been over-emphasized in the past, the features characterising the erstwhile broadly-delimited and polyphyletic Oncidium - basically, mimicry of Malpighiaceae oil flowers - being just a single example (Williams et al. 2001; Neubig et al. 2008). Thus generic limits suggested by molecular studies and those suggested by variation in floral morphology do not always agree (Kocyan et al. 2008 and references), indeed, in some cases vegetative variation may correlate better with clades evident in molecular phylogenies (e.g. Cameron 2005a). Generic limits in Epidendroideae as elsewhere were based on floral variation, but vegetative variation correlates better with clades evident in molecular phylogenies (Cameron 2005a), even if anatomical variation may suggest little in the way of major phylogenetic structure (Stern et al. 2004; Stern & Carlsward 2006). In Aeridinae, too, there is probably widespread parallelism in characters used to delimit genera (Hidayat et al. 2005). For a reclassification of Pleurothallidinae, see Pridgeon and Chase (2001); Pleurothallis was not monophyletic. There is some controversy about generic limits in the Masdevallia area, cf. Luer (2006) and Pridgeon (2007). Szlachetko et al. (2005 and references) give a statement of the "floral" position, maintaining that variation in column form, etc., yields taxonomically important characters. Such disagreements reflect fundamental differences in philosophies, differing beliefs in the ability of morphology when used alone alone to disclose relationships, and/or a preference for evolutionary classifications.

However, having a phylogeny does not mean that there will be agreement about generic limits. Clements (2006 and references) suggests a wholesale reorganization of the Dendrobium and relatives (note that the species numbers above do not reflect this), but Burke et al. (2008) present an alternative view of how things should be reclassified. Jones and Clements (2002a, esp. 2002b) divide Pterostylis; since the monophyly of Pterostylis s.l. was not in question, the division is perhaps questionable (if one likes broadly-drawn generic limits), similarly Jones et al. (2001) dismember the monophyletic Caladenia, Jones et al. (2002) attacking Diuridae as a whole. Such differences do not necessarily involve fundamentally different classificatory philosophies, simply conflicting preferences for narrow or broad genus limits, and they are something of a pain.

For an account ofAnacamptis, Orchis, etc., see Kretzschmar et al. (2007).

Boryaceae et al. + [Ixoliriaceae + Tecophilaeaceae] + Doryanthaceae + Iridaceae + Xeronemaceae + Xanthorrhoeaceae, [Amaryllidaceae + Asparagaceae]]]]]: fructans +; cuticle wax crystals as parallel platelets; (T ± connate; A inserted on T tube); seeds exotestal, (phytomelan +).

Chemistry, Morphology, etc. For the distribution of fructose oligosaccharides, see Pollard (1982). Although recorded there only for some Hypoxidaceae in the [Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]] clade, and not for some of the other smaller families in Asparagales, fructans seem to be widespread; they were not recorded from Orchidaceae.

Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]: ovules with hypostase, embryo sac with chalazal constriction.

BORYACEAE M. W. Chase, Rudall & Conran   Back to Asparagales

Plant xeromorphic; rhizome short, roots mycorrhizal; endodermis much thickened; leaves spiral, bundles with lateral phloem, base sheathing; inflorescence scapose, involucrate, raceme or spike; T tube short; A adnate to tube [not Alania], anthers (centrifixed), little longer than wide; septal nectaries external; many crassinucellate [parietal cell only] ovules/carpel, micropyle bistomal; T persistent in fruit; (seed papillate - Borya); endosperm helobial, without starch, embryo small, ovoid; n = 11, 14; seedling?

2[list]/12. Australia, scattered (map: see Brittan et al. 1987). [Photo - Borya Habit © M. Fagg]

• Boryaceae are xeromorphic, some being resurrection plants, with aerial stems and stilt roots; the inflorescence is scapose and involucrate, but the flowers are undistinguished.

Evolution. Stem group Boryaceae are dated to ca 109 million years before present, crown group Boryaceae to ca 54 million years before present (Janssen & Bremer 2004: note their position).

Chemistry, Morphology, etc. Borya has tuberculate roots, perhaps with coil-forming Rhizoctonia (cf. Orchidaceae); the plant is arborescent and dessication-tolerant (Barthlott 2006). The pedicels of Alania have several bracteoles. The nucellus is a single cell layer across. Additional information is taken from Dahlgren et al. (1985), Conran (1998: general), Conran and Temby (2000: floral morphology).

Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]: ovules with hypostase, and nucellar cap.

Phylogeny. An at most moderately well-supported - if persistently appearing - group (Rudall et al. 1998a; Chase et al. 2000a; Fay et al. 2000; Davis et al. 2004 - Lanariaceae not included; Graham et al. 2006; Chase et al. 2006; etc).

Chemistry, Morphology, etc. See Conran and Temby (2000) for general information esp. about ovules.

BLANDFORDIACEAE R. Dahlgren & Clifford   Back to Asparagales

Rhizome short; chemistry?; hairs 0; velamen +; raphides 0; leaves two-ranked, flat-curved, midrib prominent, sheath?; inflorescence a raceme; pedicels articulated, T large, tubular; A adnate to below middle of tube, latrorse, centrifixed, pollen trichotomosulcate; G stipitate, septal nectaries external, many ovules/carpel, outer integument ca 4 cells across, style short, stigma ± punctate, dry; capsule septicidal; exotesta papillate; embryo short; n = 17, 27; cotyledon photosynthetic.

1[list]/4. E. Australia (map: see Brittan et al. 1987). [Photo - Blandfordia Flower © B. Walters]

Evolution. Blandfordiaceae date to ca 100 million years before present (Janssen & Bremer 2004: note topology).

Previopus Relationships. Rudall (2003a) suggested that there was a close morphological relationship between Boryaceae and Blandfordiaceae.

Chemistry, Morphology, etc. Information is taken from Clifford and Conran (1998: general), Prakash and Ramsey (2000: embryology) and Kocyan and Endress (2001b: some floral morphology.

Lanariaceae [Asteliaceae + Hypoxidaceae]: hairs multicellular, often branched; stomata paracytic; lamina with distinct midrib; G ± inferior[?], micropyle bistomal.

LANARIACEAE R. Dahlgren & A. E. van Wyk   Back to Asparagales

Plant with vertical rhizome; biflavones +; raphides 0 (styloids +); indumentum dendritic; leaves two-ranked to spiral, sheath ?closed; inflorescence branched; T connate half-way; A adnate in mouth; G ± inferior, 2 apotropous fully crassinucellate ovules/carpel, micropyle zig-zag, integument ca 5 cells across, obturator +, style long, stigma punctate; capsule type?; seed 1; exotesta palisade, thin-walled, other cells rounded, tegmen persists, develops at micropyle; endosperm initially with starch; n = 18; seedling?

1[list]/1: Lanaria plumosa. Cape Province, South Africa. [Photo - Habit] [Photo - Habit]

• Lanariaceae are monocots that can be recognised by their shortly branched inflorescences covered by dendritic hairs and radially symmetric flowers with tepals connate half-way.

Evolution. The divergence of Lanariaceae dates from ca 113 million years before present (Janssen & Bremer 2004: note topology).

Chemistry, Morphology, etc. Information is taken from De Vos (1963 - embryology), Dora and Edwards (1991 - chemistry) and Rudall (1998 - general).

Asteliaceae + Hypoxidaceae: rosette-forming or caespitose; flavonols +; mucilage canals +; endosperm thin-walled; cotyledon not photosynthetic, ligule long.

ASTELIACEAE Dumortier   Back to Asparagales

Plant ± rhizomatous; saponins +; indumentum leipdote-stellate; leaves spiral, base sheathing or not; plant dioecious (flowers perfect), inflorescence branched raceme or spike, inflorescence bracts large; flowers rather small, T connate basally (free), A adnate to base of T/free, (G subinferior; placentation parietal), intra-ovarian trichomes +, few to many ovules/carpel, micropyle also zig-zag, hypostase +, nucellar cap 0, style divided or not (short), stigmas dry; fruit a berry; (seed with mucilaginous hairs), endosperm oily, no hemicellulose; n = 8, 30, ?35, chromosomes 4-6 µm long; seedling primary root well developed.

1[list]/36. New Zealand to New Guinea, Pacific Islands E. to Hawaii, Chile, the Mascarenes (map: see van Steenis & van Balgooy 1966; Brittan et al. 1987). [Photos - Milligania & Astelia Flowers © C. Howells - Australian Plants Society, Tasmania.]

Evolution. Stem group Asteliaceae are dated to ca 104 million years before present, crown group Asteliaceae to ca 92 million years before present (Janssen & Bremer 2004: note topology). For the biogeography of the family, see Birch et al. (2008).

Given the developing ideas of relationships in the family (see below), character evolution in it will repay investigation.

Chemistry, Morphology, etc. The nectaries may be on the outside of the ovary. For additional information, see Di Fulvio and Cave (1965) and Prakash and Ramsey (2000: both embryology), and Bayer et al. (1998a: general) for information.

Phylogeny. the phylogeny of the family has been clarified by Birch et al. (2009); Milligania, with loculical capsular fruits, a semi-inferior ovary and no intra-ovarian trichomes, perfect flowers, etc., and often considered rather different from other Asteliaceae, seems to be embedded in Astelia, as do the other small genera previously recognized in the family (Birch et al. 2008, esp. 2009).

Previous Relationships. Relationships between Milligania and Lanaria and Blandfordia have been suggested (Bayer et al. 1998a).

HYPOXIDACEAE R. Brown, nom. cons.   Back to Asparagales

Stem ± cormose, leaf bases persisting, contractile roots common; saponins 0; velamen +, dimorphic root hypodermis 0; stomata (tetracytic), with oblique or parallel cell divisions; leaves 3-ranked, (conduplicate-)plicate (conduplicate-flat), (unifacial), sheaths also closed; inflorescence various, scapose, axis compressed; (flowers 2-merous), T free to tubular, (A inserted lower down; many; 3, plus 3 staminodes adnate to style = gynostemium - Pauridia; extrorse), basifixed, sagittate, tapetum plasmodial, microsporogenesis successive [tetrads tetragonal], ovary inferior, apical beak common, (placentation parietal - Empodium), septal nectary 0, few to many ovules/carpel, stigma ± radiate, dry or wet; fruit capsular (circumscissile) or baccate; seeds globose, exotesta palisade or not, (endotegmen persistent), raphe prominent; (endosperm nuclear - Pauridia); n = 6-9, 11, chromosomes 2-5 µm long; embryo short, undifferentiated.

7-9[list]/100-220: Hypoxis (50-100, but apomictic...: ?inc. Rhodohypoxis, etc.). Seasonal tropics, esp. southern Africa (temperate) (map: see Fl. N. Am. 26: 2002; FloraBase 2005). [Photo - Inflorescence] [Photo - Flower]

• Hypoxidaceae can be recognised by their rosettes of plicate or at least folded leaves and persistent leaf bases; non-glandular indumentum is quite prominent. In the flowers, the outer tepal whorl tends to be green outside, and the ovary is inferior; there is often a narrowly tubular part at the apex of the ovary formed by the connate tepals or by an apical beak of the ovary itself.

Evolution. Stem group Hypoxidaceae are dated to ca 78 million years before present, crown group Hypoxidaceae to ca 100 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. Kocyan (2007) found that some flowers of Curculigo racemosa were polyandrous, however, they were not fasciculate. There is controversy over variation in tapetum type and the numbers of cells in the nuclei, and whether or not there is a velamen. The ovules have a parietal cell, so are not tenuinucellate.

Some information is taken from Rudall et al. (1998a: anatomy), Nordal (1998: general), Judd (2000 general), and Kocyan and Endress (2001b: floral morphology).

Previous Relationships.Rudall (2003a) suggested that there might be a close morphological relationship between Hypoxidaceae and Orchidaceae.

[Ixoliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae [Xanthorrhoeaceae [Amaryllidaceae + Asparagaceae]]]]]: ?

Evolution. The divergence of this clade (i.e. [Asparagaceae + Iridaceae + Cyanastraceae (= Tecophilaeaceae)]) has been dated to ca 84 million years before present (Eguiarte 1995).

Phylogeny. This clade is strongly supported in analyses using data from four plastid genes (Fay et al. 2000; see also Chase et al. 2000a; Soltis et al. 2007a), but no morphological characters have yet been found for it.

Ixioliriaceae + Tecophilaeaceae: cormose; leaves spiral, base sheathing; flowers quite large, outer T mucronate to aristate, T tube short; A inserted at mouth; embryo long; x = 12.

Chemistry, Morphology, etc. The outer tepals in at least some Iridaceae (and Orchidaceae!) are also mucronate to aristate.

Phylogeny. There is weak to moderate support for this sister-taxon pair in Chase et al. (2000a), Pires et al. (2006) and Givnish et al. (2006) and stronger support in Graham et al. (2006, but poor sampling); they have a very long branch in the three-gene analysis of Fay et al. (2000). Davis et al. (2004) found some support for a sister-group relationship between Ixoliriaceae and Iridaceae, although sampling was poor; Chase et al. (2006) find strong support for this relationship. Janssen and Bremer (2004) showed Ixoliriaceae diverging considerably before (although adjacent to) Tecophilaeaceae.... All told, their position is somewhat unclear, and they sometimes link with Doryanthaceae, [Ixoliriaceae + Iridaceae] being sister to Doryanthaceae in Chase et al. (2006: very little support), which are adjacent on the tree here.

Previous Relationships. Ixioliriaceae and Tecophilaeaceae are placed together (along with Eriospermaceae and Lanariaceae - for the former, see Ruscaceae s. str, Asparagaceae s.l.) in Takhtajan (1997).

IXIOLIRIACEAE Nakai   Back to Asparagales

Plant a tunicated corm; saponins 0?; dimorphic exodermis 0; peduncle with a sclerenchymatous ring; mucilage cells +; leaf shortly cylindrical at apex, base ?type; inflorescence terminal, subumbellate, leafy; T tube short; A centrifixed, ovary inferior, many ovules/carpel, stigma 3-lobed, dry; seeds angled, phytomelan +; endosperm walls pitted, starch in cells surrounding embryo; cotyledon remains white even when exposed to light!

1[list]/3. Egypt to Central Asia (map: from Traub 1942, rather approximate). [Photo - Flower © A. Shoob]

• Ixoliriaceae are cormose herbs with racemosely-arranged and quite large blue, shortly tubular flowers with an inferior ovary.

Evolution. The divergence of the Ixoliriaceae clade is dated to ca 112 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. Vascular bundles in the leaf are unequal in size, some in the inflorescence axis are arranged in a circle, enclosing additional scattered bundles. The inflorescence axis is leafy, the flowers are blue and there are no alkaloids, all unusual features for Amaryllidaceae, where Ixiolirion was often included.

Information is taken from Arroyo (1982) and Arroyo and Cutler (1984: both anatomy), and also from Kubitzki (1998b: general) and DÖnmez and Isik (2008); see Tillich (2003) for seedling morphology.

Previous Relationships. Both Dahlgren et al. (1985) and Takhtajan (1997) recognised relationships between Ixoliriaceae Tecophilaeaceae, as well as with a selection of other asparagalean families.

TECOPHILAEACEAE Leybold, nom. cons.   Back to Asparagales

Corm tunicate or not; ?saponins +; stomata paracytic, divisions parallel; (leaves petiolate, with midrib), sheaths open? (none); inflorescence a raceme, branched or not, or flowers axillary; flowers often monosymmetric, (T tube moderate), anthers ± poricidal, pollen operculate (not Kabayea and Cyanastrum), (stamens of very different sizes, or staminodes 2-3); G (semi-inferior; free - Cyanastrum), 2-many ana-campylotropous ovules/carpel, obturator +, stigma punctate; seed variable, phytomelan +/0, testa multilayered, (exotesta palisade), thick-walled; endosperm (nuclear - Cyanella), thick-walled, pitted or not, ?starch (0, chalazosperm + - Cyanastrum), embryo also short; n = 8, 10-12, 14, chromosomes 2-4 µm long; cotyledon not photosynthetic, (coleoptile +), primary root long (hypocotyl and primary root 0 - Cyanastrum).

9[list]/23. Africa, Chile, and U.S.A. (California - Odontostomum) (map: from Carter 1962; Scott 1991; Brummitt et al. 1998; Fl. N. Am. 26: 2002).[Photo - Flower, Flower.]

• Tecophilaeaceae are cormose herbs sometimes with broad-bladed, petiolate leaves; their flowers have spreading tepals and are monosymmetric because of the androecium, the stamens being strongly dimorphic. The anthers open by pores.

Evolution. The divergence of the Tecophilaeaceae clade is dated to ca 108 million years before present, and of the crown group to ca 87 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. Cells adjacent to stomata in Cyanastrum were described as having parallel cell divisions by Tomlinson (1974). The monosymmetry of the flower is largely caused by the androecium; enantiostyly also occurs in a few species of Cyanella. Odontostomum has been reported as having six staminodia alternating with the six stamens; these are more properly to be described as a corona or enations.

Some information is taken from Rudall (1997), Simpson and Rudall (1998) and Brummitt et al. (1998), that on seedlings, which are variable in their morphology, from Tillich (1995, 2003).

Synonymy: Androsynaceae Salisbury, Conantheraceae J. D. Hooker, Cyanastraceae Engler, Cyanellaceae Salisbury, Walleriaceae Takhtajan

Doryanthaceae [Iridaceae [Xeronemataceae [Xanthorrhoeaceae [Amaryllidaceae + Asparagaceae]]]]: ?

Phylogeny. Doryanthaceae may go around here; although there is only moderate support in Fay et al. (2000), there is 92% bootstrap support for a sister group relationship [Doryanthaceae + other Asparagales] in Graham et al. (2006: note sampling).

DORYANTHACEAE R. Dahlgren & Clifford   Back to Asparagales

Huge sub-bulbous tufted perennial; steroidal saponins +; styloids +, raphides 0; cuticular wax rodlets parallel, stomata paracytic; leaves spiral, when older with dry threads at apex; inflorescence a thyrse; T large, tube long, (A also adnate to the base of the tepal lobes), latrorse, centrifixed, endothecium thick, tapetal cells with several nuclei, pollen trichotomosulcate, ovary inferior, micropyle?, stigma punctate, dry; seeds flattened, testa many-layered, with phlobaphene; endosperm thin-walled, embryo flattened; n = 17, 18, 22, 24, bimodal; seedling with laterally compressed haustorium, coleoptile +.

1[list]/2. E. Australia (map: from O. Seberg, pers. comm). [Photo - Habit.]

• Doryanthaceae are massive, tufted, perennial monocots, the leaves having entire margins but disintegrating into fibres at the apex. Their subumbellate inflorescences borne at the end of long leafy stems have numerous large, radially symmetrical, bright red flowers with inferior ovaries.

Evolution. The divergence of Doryanthaceae from other Asparagales can be dated to ca 107 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. Much information is taken from Clifford (1998); Tillich (2003) described seedling morphology. Kocyan and Endress (2001b) note that the connective is massive, each stamen being supplied by 2-4 "vascular complexes".

Previous Relationships. Rudall (2003a) suggested that there was a close morphological relationship between Iridaceae and Doryanthaceae.

Iridaceae [Xeronemataceae [Xanthorrhoeaceae [Amaryllidaceae + Asparagaceae]]]: (secondary thickening +); Arabidopsis-type telomeres lost, (TTAGGG)_n [human-type telomeres] common.

Chemistry, Morphology, etc. Glucomannans are reported from some members of this clade(Buckeridge et al. 2000), but it is unclear what significance to attach to this. Note that the loss of Arabidopsis-type telomeres is not simple; human-type telomeres ((TTAGGG)_n)) may predominate, but there are other types, too. Hyacinthaceae agree with other members of this clade, although the Arabidopsis-type telomere is somewhat more common than in the other members sampled (Adams et al. 2001; especially Sýkorová et al. 2003b, 2006a, b). Acanthocarpus also lacks the telomere, alone among the taxa discussed as being out-groups; in fact it is a member of Laxmanniaceae, an ingroup, not Dasypogonaceae (see commelinids).

Phylogeny. A group with quite strong support in Fay et al. (2000) and Soltis et al. (2007a), etc.

IRIDACEAE Jussieu, nom. cons.   Back to Asparagales

Plant rhizomatous; roots mycorrhizal, root hairs 0; flavone C-glycosides, flavonols +, chelidonic acid 0?; dimorphic root hypodermis +; (stem endodermis +); raphides 0, styloids +; cuticular wax rodlets parallel; leaves two-ranked, often equitant and isobifacial [oriented edge on to the stem], plicate; flowers usu. large, T ± free, often aristate; A 3 (2 - Diplarrhena), extrorse, 1-many crassi- or tenuinucellate ovules/carpel, outer integument 4-6 cells across, micropyle endo- or exostomal, style branched (branches bifid), stigma on the edges of the complex/expanded style, dry; seed testal and tegmic, phytomelan 0, phlobaphene +, endotesta with lipids; endosperm thick-walled, hemicellulosic, embryo quite large; cotyledon not photosynthetic, (with ligule or coleoptile - e.g. Tigridia; photosynthetic - e.g. Sisyrinchium; hypocotyl short).

66[list]/2025 - eight subfamilies below. World-wide (map: see Heywood 1978 [S. America], Hultén & Fries 1986; Fl. N. Am. 26: 2002; Bahali et al. 2004; FloraBase 2005; Davies et al. 2005; Rodrigues & Sytsma 2006; Alexeyeva 2008. Note that Fig. 2b in Davies et al. 2005 suggests that Iridaceae grow throughout Africa, much of the Arabian Peninsula, Central Asia, etc. - this should be confirmed). [Photos - Collection]

1. Isophysidoideae Thorne & Reveal

Vessel elements in roots with scalariform perforation plates; biflavonoids [amentoflavone] +; flower solitary, with spathes; microsporogenesis?, nectary 0; G [3], style shortly branched, branches alternating with anthers; seedling unknown.

1/1: Isophysis tasmanica. Tasmania.

Synonymy: Isophysidaceae F. A. Barkley

Iridoideae [Patersonioideae [Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]]]: xanthone [mangiferin] +; vessel elements in roots with simple perforation plates; inflorescence with cymose units in which the flowers arise successively from axils of the prophylls, i.e. alternating [a rhipidium]; flowers short lived [open ca 1 day], (pollen operculate [often with two exine bands in a sulcus]), G inferior; endosperm nuclear.

2. Iridoideae Eaton

(Plant bulbous); gamma-glutamyl peptides, metacarboxy amino acids +; vessel elements in root with simple perforation plates; rhiphidia simple; (flowers long-lived; monosymmetric - Diplarrhena), T nectaries +, oil glands or oil hairs +, (septal nectaries + - Diplarrhena), endothecium with spiral thickenings, style branches long, tubular, (branches alternating with anthers, Sisyrynchium et al.); n = .

30/820: Iris (280, inc. Belamcanda), Moraea (200), Sisyrinchium (60). Worldwide, but esp. the spine of Central and South America.

Patersonioideae [Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]]: rhipidia 2, fused [binate], each unit with 2-many flowers; T connate, endothecium with base-plate or U-shaped thickenings; extra codon in rps4 gene.

3. Patersonioideae Goldblatt

Plant ± woody and rhizomatous; biflavonoids [amentoflavone] +; secondary growth +; flowers blue, (inner tepals reduced to scales or 0), filaments ± connate, pollen spherical, inaperturate, intectate, septal nectary 0; embryo small; n = 11, 21; two extra codons in rps4 gene.

1/20. More or less open conditions, Sumatra, Borneo, New Guinea, the periphery of Australia (map: from George 1986).

Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]: ?

4. Geosiridoideae ("Geosiridaceae") Goldblatt & Manning

Echlorophyllous, saprophytic; leaves heterobifacial; flowers sessile; T connate basally only, microsporogenesis successive, nectary 0; seeds minute, dust-like; endosperm helobial, starchy; n = ?

1/2. Madagascar, the Comores.

Aristeoideae [Nivenioideae + Crocoideae]: ?

5. Aristeoideae Vines

Plumbagin +; flowers ± blue, T connate basally only, septal nectay 0; embryo small; n = 16.

1/55. More or less open conditions, sub-Saharan Africa and Madagascar.

Nivenioideae + Crocoideae: flowers long-lived.

6. Nivenioideae

Plant with woody caudex; secondary thickening +; unit of rhipidium with 1-2 flowers; P long-tubular; 1(-4) shield-shaped [tangentially flattened] seeds per loculus; n = 16.

3/14. Restricted to the S.W. Cape region, South Africa

6. Crocoideae G. T. Burnett

Plant with corms; vessel elements in root with simple perforation plates; leaves when plane with pseudomidrib (not Pillansia), sheath closed; inflorescence spicate; rhipidium binate, with a single flower, pedicel 0; (flowers short-lived), variously monosymmetric (polysymmetric), endothecium with spiral thickenings, pollen exine tectate, perforate-scabrate, aperture with one or a pair of longitudinal bands forming operculum, septal nectaries +/0, ovules campylotropous, chalazal hypostase prominent; n = 9-17.

28/995: Gladiolus (260), Romulea (90: cytologically very variable, n = 9-17), Geissorhiza (85), Crocus (90), Hesperantha (80), Babiana (55), Watsonia (50), Ixia (50). Overwhelmingly southern African, to Europe, Madagascar and Central Asia.

Synonymy: Crocaceae Vest, Galaxiaceae Rafinesque, Gladiolaceae Rafinesque, Nivenioideae Goldblatt, Geosiridaceae Jonker, Ixiaceae Horaninow

• Iridaceae may be recognised by their usually ensiform, equitant, isobifacial leaves and their large flowers with showy tepals, three extrorse stamens, complex stigma, and inferior ovary.

Evolution. Stem group Iridaceae are dated to ca 103 million years before present, divergence of crown group Iridaceae to ca 96 million years before present (Janssen & Bremer 2004: Isophysis was included). Davies et al. (2005) discuss diversification rates within the family, finding them to be notably diverse (i.e. having clades with a disproportionate number of species) in e.g. southern Africa, but relatively less so in the north temperate zone. Iridaceae are one of the major geophytic groups of the Cape (Procheŝ et al. 2006) with more than 650 species there; Davies et al. (2004c) see this diversification as the result of the interaction of local features such as traits affecting reproductive isolation and the ecological and climatic heterogeneity of the area. For the radiation of the Cape genus Moraea, both cytologically and florally diverse, see Goldblatt et al. (2002); radiation in this and other iridaceous Cape genera may have begun in the fynbos in the Miocene some 25 million years before present, divergence in the succulent Karoo being more recent (Verboom et al. 2009).

In general Iridaceae show considerable floral diversification, whether based on the flower type of Iris et al. or on the tubular flowers such as occur in Gladiolus et al. (e.g. Bernhardt & Goldblatt 2006 and references; Rodrigues & Sytsma 2006; Wilson 2006). In Iris and its relatives, the tepaloid style overarches the stamen opposite it, the landing platform being a member of the outer perianth whorl. Thus the flower is a kind of revolver flower, with three points of entry for the pollinator - another way of thinking about the flower is that it appears to the pollinator as if were really three monosymmetric flowers. Interestingly, in the oil-flowers of Cypella the three landing platforms for the pollinating bee are members of the inner perianth whorl. Here the pollen deposited on the backs of the bees comes from half anthers of adjacent stamens and is deposited on the receptive surfaces of two adjacent half-stigmas (Vogel 1974): there are other oil flowers in Iridaceae. The flowers of Gladiolus are obliquely monosymmetrical, although this is hardly apparent in the open flowers due to changes in orientation as the flower and inflorescence grow. Tepal patterning, where it occurs, is usually on an adaxial lateral member of the outer whorl and adjacent members of the inner tepal whorl and is then clearly on the adaxial side of the flower, but it sometimes occurs on the adaxial lateral and abaxial members of the outer whorl and the member of the inner whorl between them (Eichler 1875; Choob 2001). Although it is likely that other Crocoideae show the same oblique monosymmetry, monosymmetry in Diplarrhena (Iridoideae) appears to be vertical; that genus has only two stamens and one staminode. Interesting infraspecific variation occurs, thus in some flowers of Crocosmia X crocosmiiflora the odd member of the outer whorl is adaxial, and in others it is abaxial, and patterning of tepals, etc., varies accordingly. All told, well over half the family has monosymmetric flowers of one sort or another; certainly, the evolution of monosymmetry in the family will repay further study (see also Davies et al. 2004b).

Babiana (Crocoideae) is pollinated by birds, scarab beetles, bees, moths, etc. (Bernhardt & Goldblatt 2006, esp. Goldblatt & Manning 2007). Much work has been carried out on pollination in Iridaceae, particularly those from the sub-Saharan (esp. South African) region (Goldblatt and Manning 2006, see also 2008 for a general account).

There is also considerable variation in vegetative morphology in the family, some taxa having terete, unifacial leaves, others apparently ordinary heterobifacial leaves (even some Iris), etc., and many have ensiform isobifacial leaves as in Gladiolus, most Iris, etc. (e.g. Arber 1925); Geissorhiza alone has ligulate leaves. Crocus has revolute leaves with a unifacial midrib, and Romulea seems to be a modification of this.

Chemistry, Morphology, etc. For the occurence of plumbagin in Aristea, see Harborne and Williams (2001). Homeria and Moraea (Iridoideae) have bufadienolides (Harborne & Williams 2001). Iris contains a greater diversity of isoflavonoids than any other group outside Fabaceae (Reynaud et al. 2005).

Goldblatt (1990) interpreted the paired "bracts" below the single flowers of Isophysis as representing a reduced rhipidium - a rhipidium may then be another synapomorphy for the family. Some species of Nivenia are heterostylous, a very uncommon condition in the monocots. Aristea is palynologically very variable, some members even having disulcate pollen (see Goldblatt & Le Thomas 1997; le Thomas et al. 2001). In Sisyrynchium and its relatives the style branches alternate with the stamens; elsewhere in the family, the two are on the same radius. For a discussion of the caruncles/arils of Iris, see Wilson (2006).

Additional information is taken from from Rudall et al. (1986) and Rudall (1995a), both anatomy, Goldblatt et al. (1998: general), Goldblatt (2001: general), Cocucci and Vogel (2001: nectaries), Tillich (2003a: seedlings - very variable), Rudall et al. (2003a: nectary evolution), DÖnmez and Isik (2008: pollen), and Goldblatt and Manning (2008: general account).

Phylogeny. Iridaceae are monophyletic in nearly all studies (but cf. Chase et al. 1995a). Initial resuls suggested that the monotypic Isophysidoideae were sister to the rest of the family, Crocoideae and Iridoideae appeared to be monophyletic, but the status of Aristeoideae was unclear. The four-gene analysis of Reeves et al. (2001a, b) suggested that Patersonia, Geosiris, and Aristea were successively sister to a large clade making up [Aristeoideae + Crocoideae]; support was mostly moderate (see also Teixeira de Souza-Chies et al. 1997). If these relationships are confirmed, either the circumscription of Crocoideae will have to be considerably extended, or three more subfamilies will be needed - basically, the classification that developed. Goldblatt et al. (2008: five plastid genes) opted for this latter classification; they found strong support for the pectinations basal to Crocoideae s. str., albeit using successive weighting, which tends to leave one a little uneasy. Within Crocoideae the five tribes are mostly only moderately supported and their relationships are unresolved, on the other hand, the five tribes recognised in Iridioideae are well supported and their relationships are better resolved: [Diplarreneae [Irideae [Sisyrincheae [Trimezieae + Tigridieae]]]] (Goldblatt & Manning 2008; seee also Golblatt et al. 2004, 2006).

Note that the Australian Diplarrhena, whose flowers have only two stamens and are monosymmetric, is perhaps sister to all other Iridoideae (Reeves et al. 2001a, b; Rudall et al. 2003a); Diplarrhena also has spherical, inaperturate, intectate pollen grains. For diversification of the American Tigridieae, see Rodrigues and Sytsma (2006); there is very extensive floral homoplasy (as in Iris - Wilson 2006). For a phylogeny of Iris, see Tillie et al. (2001) and Wilson (2004). Within Crocoideae Tritoniopsis, with a tubular cotyledonary sheath and tubular cataphyll, may be sister to the rest of the subfamily, but with at best moderate support (Goldblatt et al. 2006; Golblatt & Manning 2008). For a phylogeny of Crocus see Petersen et al. (2008, cf. in part Frello et al. 2004), and this is explained by Mathew et al. (2009); see Goldblatt and Manning (1998) for a treatment of much of Gladiolus.

Classification. I follow the classification suggested by Goldblatt et al. (2008); the subfamilies are for the most part well characterised. See Goldblatt and Manning (2008) for an account of the genera.

Xeronemataceae [Xanthorrhoeaceae [Amaryllidaceae + Asparagaceae]]: mitochondrial rpl2 gene lost.

A strongly supported group in Fay et al. (2000) and Soltis et al. (2007a). The loss of the mitochondrial rpl2 gene occurs either at this node or the next up the tree, according to its distribution in Adams et al. (2002b).

XERONEMATACEAE M. W. Chase, Rudall & Fay   Back to Asparagales

Plant rhizomatous; leaves two-ranked, equitant and isobifacial [oriented edge on to the stem]; inflorescence a dense spike; flower large, pollen boat-shaped, style solid; n = 17, 18.

1/2. New Zealand (Poor Knights Island) and New Caledonia.

• Xeronemataceae are large monocot herbs that may be recognised by their equitant isobifacial leaves and congested inflorescences with rather large, radially symmetrical, upwardly-facing flowers. The stamens are strongly exserted.

Evolution. The divergence of Xeronemataceae from other Asparagales has been dated to ca 100 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. The family is little known, although there is some information in Chase et al. (2000c); the style is scored as if it is hollow in Rudall (2003a).

Previous Relationships. Xeronemataceae were provisionally placed in Asphodelaceae by Takhtajan (1997) and in Hemerocallidaceae by Clifford et al. (1998).

Xanthorrhoeaceae [Amaryllidaceae + Asparagaceae]: (pedicels articulated); septal nectaries infralocular [see beginning of this page], ovary superior.

Chemistry, Morphology. For chromosome sizes of a number of taxa in the group, see Vijayavalli and Mathew (1990 - as Liliaceae).

Phylogeny. This group has strong support in Fay et al. (2000) and Chase et al. (2000b). The optimisation of successive microsporogenesis on the tree is uncertain, i.a. microsporogenesis varies within Xanthorrhoeaceae.

XANTHORRHOEACEAE Dumortier, nom. cons.   Back to Asparagales

Anthraquinones +; styloids +; inflorescence branches cymose; cotyledon not photosynthetic.

35/900. Esp. Old World, not Arctic, western South America.

1. Hemerocallidoideae Lindley

Habit various; flavonols, naphthoquinones, saponins +; roots often swollen; mucilage cells 0; raphides 0; cuticular wax rodlets parallel; leaves (spirally) two-ranked, conduplicate to flat-conduplicate, sheath closed, (immediately above the sheath semi-ensiform, unifacial - most of phormioid clade); inflorescence various, (bracteoles lateral); pedicel usu. articulated; (flowers monosymmetric, median tepal of outer whorl adaxial - Hemerocallis), T tube short (1/2 way - Hemerocallis; 0), filaments often ornamented/swollen, (anthers centrifixed; porate [Dianella and relatives]), pollen usu. trichotomosulcate, infra-locular septal nectaries +, 1-many tenuinucellate ovules/carpel, nucellar cap +, chalazal nucellus well developed; endosperm usu. helobial, stigma dry (wet); fruit also a berry (nut, schizocarp); seeds ovoid, (with strophiole/aril - Johnsonia et al.); endosperm hemicellulosic, embryo also short; n = 4 [Agrostocrinum], 8, 9, 11, 12, chromosomes 0.8-17.33 µm long; (cotyledon not photosynthetic - Dianella), epicotyl long or not (hypocotyl 0; collar +), primary root well developed, branched or not.

19[list]/85. Papuasia to New Zealand and the Pacific, esp. Australia (e.g. all 8 genera of Johnsoniaceae s. str., inc. Geitonoplesium, etc.), also Europe to Asia, Malesia, India, Madagascar, Africa, and two genera in South America (map: from Wurdack & Dorr 2009). [Photo - Habit, Flower, Flower].

Synonymy: Dianellaceae Salisbury, Geitonoplesiaceae Conran, Hemerocallidaceae R. Brown, Johnsoniaceae J. T. Lotsy (= Anthericaceae - Johnsonieae), Phormiaceae J. Agardh

Xanthorrhoeoideae + Asphodeloideae: secondary thickening +; A not adnate to T, hypostase +; seeds angled.

2. Xanthorrhoeoideae M. W. Chase, Reveal & M. F. Fay   Back to Asparagales

Stem thick, woody; plant resiniferous; raphides 0; layer of sclerenchyma below epidermis in leaves; stomata paracytic; leaves spiral, unifacial, not sheathing; inflorescence spike-like, of congested cymes; pedicels not articulated; T = 3 dry + 3 subpetaloid, free, microsporogenesis successive [tetrads tetragonal], pollen extended sulcate, infra-locular septal nectaries +, several ovules/carpel, apex of nucellus pointed, stigma ± punctate, ?wet; inner cuticle of tegmen +; seeds flattened; endosperm quite thick-walled, development?, little hemicellulose, embryo transverse to long axis of seed; n = 11 (bimodal); hypocotyl short.

1[list]/30. Australia (map: see Bedford et al. 1986). [Photo - Habitat, - Habit, Inflorescence.]

3. Asphodeloideae Burnett

Geophytic herbs to pachycaul trees, rosette-forming; tetrahydroanthracenones in roots, steroidal saponins 0; (velamen +); foliar vascular bundles often inverted, parenchymatous perhaps secretory cells [aloin cells] in the inner bundle sheath adjacent to the phloem; leaves spiral or two-ranked, often fleshy, margins often toothed, sheath closed (leaves not sheathing); inflorescence (branched) racemose or spicate; pedicels articulated ["basal" clades] or not, ± weak monosymmetry common, T ± free (connate - Kniphofia, Alooideae, (anthers centrifixed), microsporogenesis simultaneous [tetrads tetrahedral], 1-many straight [Asphodelus clade] to hemitropous [the rest] ovules/carpel, hypostase +, stigma dry (wet); (capsule fleshy); seed ± ovoid, with impressed funicular aril [arising as annular invagination at the apex of the funicle]; endosperm thick-walled, hemicellulosic[?], (perisperm +, slight), embryo long; n = (6 - Kniphofia) 7, chromosomes 1.5-20 µm long, bimodal; 3'-rps12 intron lost [Bulbine not sampled]; (coleoptile +).

Ca 15[list]/785: Aloe (400), Haworthia (54-70), Kniphofia (70, sp. limits difficult), Bulbine (60), Trachyandra (50). Africa, esp. South Africa, also South Africa + New Zealand (Bulbinella) and the Mediterranean to Central Asia (map: see Reynolds 1966; Seberg 2007). [Photo - Collection, Inflorescence, Flowers.]

• Xanthorrhoeoideae can be recognised by their habit, having a dense tuft of long, narrow leaves terminating a stout, woody stem (and persisting as a skirt of dried leaves if there are no fires) and an erect, densely spike-like inflorescence with small flowers and capsular fruits.

Evolution. Stem group Xanthorrhoeaceae s.l. are dated to ca 93 million years before present, divergence within crown group Xanthorrhoeaceae s.l. to ca 90 million years before present (Janssen & Bremer 2004). Excremis and Pasithea represent independent migrations of the phormioid clade to South America (Wurdack & Door 2009).

For an ecological account of Xanthorrhoea, see Lamont et al. (2004); some diversification in the genus may be associated with the aridification of the Nullarbor Plain some 14-13 million years ago separating eastern and western clades (Crisp & Cook 2007).

Members of Asphodeloideae have more or less succulent leaves, and species of Aloe and Haworthia in particular are commonly rosette plants with fleshy leaves; these can be spiral or distinctively two-ranked. As with Aizoaceae from southern Africa, there is great variation in the micromorphology of the epidermis in Aloe and Haworthia (Cutler 1982); the two grow in similar extreme habitats.

Within Asphodeloideae, many species of Aloe are pollinated by birds, but insect pollination is also known here, as in other groups of Asphodelaceae; the directionality of evolution of pollinator relationships is unclear (Hargreaves et al. 2008). The floral monosymmetry that occurs in Haworthia and relatives is rather weak. Hemerocallidoideae often have rather elaborate stamens.

Chemistry, Morphology, etc. There is variation in microsporogenesis in the clade, as in Hemerocallidoideae. Both Hemerocallidoideae and Xanthorrhoeoideae have ovaries that may be secondarily superior and that have infra-locular septal nectaries (Rudall 2002, 2003a).

In Asphodeloideae Bulbine, Trachyandra, and Kniphofia all have knipholone, an anthraquinone derivative (van Wyck et al. 2005), but it appears not to have been reported from the Asphodelus clade. Aloin cells are reported from Dianella (Hemerocallidoideae: see Rudall 2003a). In at least some species of Aloe the larger stamens are opposite the inner whorl of tepals. Note that ovule orientation at the basal node in the family is unclear (cf. Steyn & Smith 1998). Kniphofia has an bistomal micropyle and a nucellar endothelium (Takhtajan 1985); it lacks aloin cells, having a well developed sclerenchymatous cap in their place (as have some other Asphodelaceae, even some Alooideae).

Some information is taken from Riley and Majumdar (1979: biosystematics), Beaumont et al. (1985: leaf anatomy and chemistry), Van Wyk et al. (1995, 2005: chemotaxonomy), G. Smith and Van Wyk (1998: general), Steyn and Smith (1998: ovule morphology, 2001), McPherson et al. (2004 - loss of the 3'-rps12 intron), and Reynolds (2004: esp. Aloe).

Johnsonia (Hemerocallidoideae) has chelidonic acid (Ramstad 1953). Microsporogenesis in Hemerocallis was described as being successive (?alone in Hemerocallidoideae) and the endosperm as being nuclear by Di Fulvio and Cave (1965, but cf. Cave 1955). Hemerocallis also has isoflavones, monosulcate pollen and a wet stigma, but it lacks a nucellar cap and septal nectaries. In pollen morphology Hemerocallis was considered to be derived by Chase et al. (1996), indeed, it is unlikely to be sister to the rest of Hemerocallidoideae (McPherson et al. 2004). Hemerocallis also seems to have lateral bracteoles, as does Dianella; both may have "inverted" flowers (e.g. Eichler 1875; Ehrhardt 1992), although in Hemerocallis, at least, this seems to be variable, and although flowers with the median outer tepal adaxial are common, the seal of the Daylily Society shows a flower with the normal monocot orientation. The number of vascular bundles supplying the tepals in members of this subfamily varies from (1-)3-9(-25) (Clifford et al. 1998a).

Information is taken from Kosenko (1994: pollen), Chase et al. (1996: general), Clifford and Conran (1998 - Johnsoniaceae: general), and Clifford et al. (1998a: general).

Floral tube and chelidonic acid in Xanthorrhoea? Some information is taken from Chanda and Ghosh (1976: pollen), Rudall and Chase (1996: phylogeny) and Clifford (1998: general).

Phylogeny. There is strong support for Xanthorrhoeaceae s.l. in Fay et al. (2000), Wurdack and Dorr (2009), etc. See Kite et al. (2000) for the distribution of anthroquinones, McPherson et al. (2004) for taxa lacking the 3'-rps12 intron. However, relationships within the clade remain unclear. There is moderate support for [Xanthorrhoeoideae + Asphodeloideae] in the three-gene tree of Chase et al. (2000a), less in analyses including taxa with some sequences missing; see also Fay et al. (2000). However, Devey et al. (2006) find some support for a [Xanthorrhoeoideae + Hemerocallidoideae] clade (see also Pires et al. 2006; Wurdack & Dorr 2009 - slightly better than moderate support), while Chase et al. (2006, but see sampling) suggest a [Asphodeloideae + Hemerocallidoideae] clade. Rudall (2003a) suggested a close morphological relationship between Hemerocallidaceae (Hemerocallidoideae) and Asphodelaceae (Asphodeloideae) - and between Xanthorrhoeaceae s. str. and Iridaceae...

Within Asphodelaceae, Alooideae are very distinctive. They are sometimes shrubby plants, and secondary growth is common. They have 1-methyl-8-hydroxyanthraquinones [e.g. chrysophanol; in roots] and anthrone-C-glycosides [in leaves]. Their sieve tube plastids also have peripheral fibers in addition to the central protein crystal. They have tetracytic stomata (e.g. Cutler 1972), although this is questioned by G. Smith and van Wyk (1992); perhaps there is variation. The leaves are notably succulent, with white or concolorous spots and tubercles. The vascular bundles in the leaf form a circle and there are globules in the outer bundle sheath (also in Kniphofia); the central cells of the leaf are gelatinous. The karyotype is bimodal. However, Bulbine is sister to Alooideae, and then come other Asphodeloideae, including Kniphofia et al. and Eremurus et al.; the Asphodelus clade is sister to all the rest of the family (good support). Thus the recognition of Alooideae makes Asphodeloideae paraphyletic (see Devey et al. 2006 for a phylogeny, inc. details of that of Bulbine, also references below). Some species of Bulbine have a bimodal karyotype on n = 7 (4 long, 3 short: Spies & Hardy 1983), rather like the karyotype of Aloeae (4L + 3S: probably evolved independently, see Devey et al. 2006; Pires et al. 2006), and also the same medicinal properties... For relationships around Aloe, see Treutlein et al. (2003a, b).

There are two well supported clades within Hemerocallidoideae, the phormioid and [hemerocallid + johnsonioid] clades (Wurdack & Dorr 2009). Pasithea is siter to all other phormioids, and it has completely bifacial leaves, i.e. the plesiomorphic condition (Wurdack & Dorr 2009). The loss of the 3'-rps12 intron characterises a major clade [Johnsonieae + Hemerocallis + Simethis], i.e. the latter clade above, see McPherson et al. (2004) and Chase et al. (2000b).

Excremis coarctata is odd; it was initially placed here only with hesitation (see Clifford et al. 1998a; Devey et al. 2006): pedicel articulated; A connate basally, 3 [which whorl?] adnate to T; also a septicidal endocarp: 1 sp., Andean (for anatomy, see Ely & Luque Arias 2006, the base of the leaf is isobifacial). It has the habit of Phormium, and it was sometimes included in Phormiaceae (= Hemerocallidaceae s. str./Hemerocallidoideae). The only other South American member of Asphodeloideae is Pasithea. Wurdack and Dorr (2009) found that both were members of the phormioid clade.

Classification. A.P.G. II (2003) suggested as an option including Asphodelaceae, Xanthorrhoeaceae and Hemerocallidaceae in Xanthorrhoeaceae s.l., and this circumscription was adopted by A.P.G. III (2009). The subfamial classification follows that in Chase et al. (2009b).

Species estimates in Dianella (Hemerocallidoideae) range from 25-350+ (Carr 2007). Generic limits around Aloe itself are decidedly unsatisfactory (e.g. Treutlein et al. 2003a, b), while species limits are also problematic, as in Kniphofia and also Haworthia (Bayer 2009 for some comments and refernces). G. Smith and Steyn (2004) discuss the taxonomy of Alooideae (= Aloeae). I thank Syd Ramdhani and Matt Ogburn for useful discussion.

Previous Relationships. Three genera that used to be placed in Asphodelaceae s. str. are now in Hemerocallidoideae (Simethis), Asparagaceae-Asparagoideae (Hemiphylacus) and Asparagaceae-Agavoideae (Paradisea, Anthericaceae s. str.) respectively - the evidence is largely molecular (Chase et al. 2000b).

Synonymy: Aloaceae Batsch, Asphodelaceae Jussieu, nom. cons., Eccremidaceae Doweld

Amaryllidaceae + Asparagaceae: microsporogenesis successive [possible place]; endosperm development?

Evolution. This clade separates from Xanthorrhoeaceae s.l. ca 93 million years before present, divergence within it begins at ca 91 million years before present (Janssen & Bremer 2004), corresponding figures given by Wikström et al. (2001) are 61-54 and 58-51 million years before present respectively.

Chemistry, Morphology, etc. Microsporogenesis is uniform in this group. In other Asparagales that also have successive microsporogenesis, details of wall formation (centrifugal cell plates) is similar to those members of this clade that have been studied, however, where microsporogensis is simultaneous, plate formation may also be centripetal (Nadot et al. 2006). For chromosome size in Liliaceae s.l. and supposed relatives, see Vijayavalli and Mathew (1990).

Phylogeny. This is a strongly supported clade (e.g. Chase et al. 1995a; Fay et al. 2000; Chase et al. 2000b; Graham et al. 2005). However, inclusion of Aphyllanthaceae/Asparagaceae-Aphyllanthoideae in analyses has tended to decrease support for clades within it (Graham et al. 2006).

AMARYLLIDACEAE J. Saint-Hilaire, nom. cons.   Back to Asparagales

Lectins binding mannose; leaves two-ranked; inflorescence scapose, umbellate [cymose construction], with scarious spathe, inflorescence bracts 2 (or more - external), pedicels not articulated; (T free; A connate basally), (tapetal cells uninucleate), style long, stigma dry; endosperm nuclear or helobial; hypocotyl 0.

73/1605. Worldwide - three subfamilies below.

Evolution. Stem group Amaryllidaceae are dated to ca 91 million years before present, divergence within crown group Amaryllidaceae begins ca 87 million years before present (Janssen & Bremer 2004).

Fungi on Allium and other Allioideae are rather different from those on Amaryllidoideae (e.g. Savile 1962).

Chemistry, Morphology, etc. Distinctive, mannose-binding lectins (the specificity is absolute) are found in Allioideae and Amaryllidoideae (van Dammme et al. 1991); I do not know if they have been recorded from Agapanthus. Very large genomes with a C value of some 350 picograms or more are found in some Allioideae and Amaryllidoideae - also also Asparagaceae-Scilloideae and Orchidaceae (Leitch et al. 2005). For tapetal cells, see Wunderlich (1954), for inflorescence structure, see Weberling (1989).

Phylogeny. This is a very strongly supported clade (e.g. Fay et al. 2000, but cf. McPherson et al. 2004; Thomas et al. 2005), and it has some characters! Meerow et al. (1999), Fay et al. (2000: strong support), Givnish et al. (2006) and Pires et al. (2006) suggest a set of relationships [Agapanthaceae [Alliaceae + Amaryllidaceae]], although Meerow et al. (2000a) found Agapanthaceae to be sister to Amaryllidaceae, albeit with weak support.

Classification. Combining the three families Agapanthaceae, Alliaceae and Amaryllidaceae into Alliaceae s.l. was an option in A.P.G. II (2003), an option that was exercised in A.P.G. III (2009), although the name of the clade is there Amaryllidaceae. The infrafamial classification follows that in Chase et al. (2009b).

1. Agapanthoideae Endlicher

Plant rhizomatous; leaf ptyxis flat; inflorescence bracts connate along one side; flowers large, monosymmetric, T ± connate basally, ovules apotropous; seeds flat; endosperm with hemicellulose; n = (14) 15 (16), chromosomes 4-9 µm long; seedling as in Allioideae?

1/9. South Africa (map: from Leighton 1965). [Photo - Habit] [Photo - Flower]

• Agapanthoideae are rather robust, rhizomatous herbs that can be recognised by their rather fleshy and two-ranked leaves and their scapose umbellate inflorescence of generally large flowers with superior ovaries.

Chemistry, Morphology, etc. Information is taken from Kubitzki (1998b: general).

Synonymy: Agapanthaceae F. Voigt

Allioideae + Amaryllidoideae: geophytes, bulbs sympodial, tunicate, with contractile roots.

2. Allioideae Herbert

Flavonoids, cysteine-derived sulphur compounds +; also styloids +; laticifers +; leaves (spiral), sheath closed, long, shortly ligulate [Allium, at least]; floral bracts 0; C ± connate; A connate or adnate to free, 2-many tenuinucellate campylotropous ovules/carpel, (micropyle bistomal; embryo sac bisporic), obturator +, nucellar cap + (?0), (stigma wet); seeds angular, exotestal, other layers of testa collapsed or not; (endosperm pitted); chromosomes 2-20 µm long; (cotyledon not photosynthetic).

13[list]/795: Allium (690). Mainly South America, but Allium esp. N. Temperate Eurasia - three groups below. [Photo - Collection] [Photo - Inflorescence, Flower, Flower.]

2A. Allieae Dumortier

Bulbs lacking starch; (leaves ± unifacial); C basally connate, corona 0; A both basally connate and adnate to C, often with winged filaments, tapetal cells uninuclear, (G semi-inferior), 2 epitropous-14 ovules/carpel, style solid, gynobasic; (caruncle +); endosperm cellular, embryo curved; n = (7) 8 (9), chromosomes 9.0-19.3 µm long, telomeres distinctive [?how common in family].

1/260-780. North temperate, often seasonally dry, especially Central Asia, scattered in Africa (map: from Hultén 1962; de Wilde-Duyfjes 1976; Hanelt 1990; Hanelt et al. 1992; Fl. N. Am. 26: 2002, n.b. not native in Iceland).

Synonymy: Alliaceae Borkhausen, nom. cons., Cepaceae Salisbury, Milulaceae Traub

Tulbaghieae + Gilliesieae: bulbs with starch; endosperm helobial.

2B. Tulbaghieae Meisner

Plant often rhizomatous; leaf sheath short; flowers bracteate; C rather strongly connate, corona massive, lobes connate or not; A sessile, adnate to corolla tube and/or corona, 2-several ovules/carpel; seeds ± flattened; embryo?; n = 6, chromosomes 11.5-14.7 µm long.

1/22. Southern Africa (map: from Vosa 1975).

Synonymy: Tulbaghiaceae Salisbury

2C. Gilliesieae Baker

Corona +/0, (A 2-3; variously connate and adnate; extrorse; staminodes +), 2-many ovules/carpel, (inner integument 5-7 cells acros - Dichelostemma); embryo short; n = ³4.

10/80: Nothoscordum (22). South U.S.A., Mexico to South America (map: from Fl. N. Am. 26. 2002).

Synonymy: Gilliesiaceae Lindley

• Allioideae can be recognised by their smell, their often rather fleshy and soft leaves, and their scapose umbellate inflorescence with medium-sized flowers that have superior ovaries.

Evolution. In Gilliesieae, Gilliesia has very strongly monosymmetric flowers with only two stamens; the flowers may mimic insects (Rudall et al. 2002).

Chemistry, Morphology, etc. The apparently bifacial leaves of at least some species of Allium have inverted vascular bundles along the adaxial surface and vascular bundles with normal orientation along the abaxial surface (Mathew 1996); I do not know how widely this feature is spread in the family. The flowers of Allium are shown with the median member of the outer whorl in the adaxial position (Spichiger et al. 2004). Do Alliodeae have apotropous ovules? There has been major movement of ribosomal protein and succinate dehydrogenase genes from the mitochondrion in Allium (Adams & Palmer 2003), and that genus has also lost its minisatellite telomeres (S&ygrave;korová et al. 2006a).

Some information is taken from Berg (1996) and Berg and Maze (1966), both embryology, and Rahn (1998: general). For Allium, see Rabinowitch and Currah (2002: more horti-/agricultural), Fritsch and Friesen (2002 [and many other papers in same book]: general), Fritsch and Keusgen (2006: cysteine sulphoxide distribution).

In Gilliesieae, Schickendantziella has only three tepals; they are caudate. Nothoscordum has solid styles.

Phylogeny. Fay and Chase (1996) discuss relationships within the subfamily; the topology is [Allieae [Tulbaghieae + Gilliesieae]], although the support for the clades is rather weak. Nguyen et al. (2008) provide a phylogeny for Allium (see also Friesen et al. 2006; Hirschegger et al. 2010 - section Allium); Old and New Word species are mostly in two separate clades, although basal to the clade containing all North American members (they belong to subgenus Amerallium) are Eurasian taxa, while the relationships of members of the small subgenera Nectaroscordum and Microscordum are unclear (Nguyen et al. 2008). Diversity within North America is centered in the west, especially in California, and a number of species there are serpentine endemics (Nguyen et al. 2008).

Classification. Friesen et al. (2006) provide a subgeneric and sectional classification ofAllium; Gregory et al. (1998) list names included in it. See Vosa (1975) for a revision of Tulbaghia and details of its cytology.

3. Amaryllidoideae Burnett   Back to Asparagales

Norbelladine alkaloids, non-protein amino acids, chelidonic acid +, saponins 0; exodermis with long and short cells, 2-4-layered velamen; sclerechymatous ring in scape, bundles in rings; petiole bundles in arc; (lacunae formed by breakdown of parenchyma); leaves (spiral), flat or revolute to involute, (vascular bundles inverted; base sheathing); bracts equitant; flowers large, (monosymmetric, with median member of outer tepalline whorl adaxial; T ± free; corona +; A ± connate and with outgrowths or tepaloid webbing), ovary inferior, (outer integument 3< cells across), stigma capitate to deeply trifid, (wet); endosperm starchy or with hemicellulose (thin-walled), embryo poorly differentiated, small; n = (5-)11(12<), chromosomes (1.5-)3-28 µm long; cotyledon bifacial, (not photosynthetic), primary root well developed, contractile.

59[list]/800+ - fourteen groups below. Tropical (temperate), esp. South America and Africa, also Mediterranean (map: from Allan Meerow and O. Seberg, pers. comm.; Snijman 1984; Fl. N. Am. 26: 2002). [Photo - Flower, Fruit.]

3A. Amaryllideae Dumortier

Extensible [helically-thickened] fibers in leaf; leaves follow the flowers (perennial - many Crininae); A connate basally (not - Crininae; small appendages developed from filaments), pollen bisulcate, exine gemmate, with scattered spinules, intectate-columellate, ovules unitegmic, embryo sac Allium type, (style laterally displaced); seeds water-rich, non-dormant, phytomelan 0, testa to 25 cells thick, chlorophyllous, with stomata, or ± collapsed, or 0, endosperm usu. with a corky layer, chlorophyllous, starchy; embryo chlorophyllous; (n = 10, 12, 15), chromosomes 5.3-20.5 µm long.

11/146: Crinum (65), Strumaria (23). SubSaharan, especially South Africa, Crinum Pantropical.

Synonymy: Crinaceae Vest, Strumariaceae Salisbury

Cyrtantheae, etc.: bundle sheath cells parenchymatous.

3B. Cyrtantheae Traub

Scape lacking sclerenchymatous ring, subepidermal collecnchyma +; 1-layered rhizodermis +, velamen 0; seeds flat, horizontally stacked; n = (7) 8 (11).

1/50 (Cyrtanthus). Africa, especially the south.

Synonymy: Cyrtanthaceae Salisbury

Calostemmateae + Haemantheae: fruit indehiscent.

3C. Calostemmateae D. & U. Müller-Doblies

2-3 ovules/carpel; embryo germinates precociously producing a bulbil; fruit dry; phytomelan 0; n = 10, chromosomes 3.3-8 µm long.

2/4. Australia, Malesia.

3D. Haemantheae Hutchinson

(Plant rhizomatous); 1-layered rhizodermis +, velamen 0; scape lacking sclerenchymatous ring, subepidermal collenchyma +; inflorescence bracts connate, (flowers single - Gethyllis, etc.); (A 12); fruit baccate; seeds angled, etc.; phytomelan 0 (+); n = 6, 8, 9, 11, 12; chromosomes 5.2-24 µm long.

6/80: Gethyllis (32), Haemanthus (22). Tropical Africa, mostly in the South.

Synonymy: Gethyllidaceae Rafinesque, Haemanthaceae Salisbury

Lycoridae + Galantheae + Pancratieae + Narcisseae (Eurasian Clade): seeds subglobose, turgid.

3E. Lycoridae D. & U. Müller-Doblies

(Seeds irregularly discoid - Ungernia); n = 11 (etc.).

2/26: Lycoris (20). Temperate to subtropical East Asia to Iran.

Galantheae + Pancratieae + Narcisseae: ?

3F. Galantheae Parlatore

(Inflorescence bractes connate along one side); elaiosome + (0); n = 7-9, 11, 12.

8/31: Galanthus (17). Europe to N. Africa, the Crimea and the Caucasus.

Synonymy: Galanthaceae G. Meyer, Leucojaceae Batsch

3G. Pancratieae Dumortier

Staminal tube toothed; n = 11, chromosomes 8.7-22 µm long.

1/20. Mediterranean, southern Asia, to sub-Saharan Africa.

Synonymy: Pancratiaceae Horaninow

3H. Narcisseae Lamarck & de Candolle

Inflorescence bracts basally connate; (corona + - Narcissus); elaiosome + (0); n = (7) 11 (13), etc.

2/58: Narcissus (50). Europe to W. Asia and N. Africa.

Synonymy: Narcissaceae Jussieu, nom. cons.

Andean + extra-Andean/American clade: 1-layered rhizodermis +, velamen 0; scape lacking sclerenchymatous ring, subepidermal collenchyma +; bracts obvolute; (seeds flat, horizontally stacked).

3I. Hippeastreae Sweet

Inflorescence bracts often connate basally (along one side); flowers monosymmetric; A declinate, of varying lengths; seeds flattened, winged or D-shaped; n = 8-13, 17, etc., chromosomes 3-16.7 µm long.

11/218: Hippeastrum (55, the "Amaryllis" of many a windowsill), Zephyranthes (50), Habranthus (50). S.E./S.W. U.S.A., the Caribbean, and Central and South America.

Synonymy: Brunsvigiaceae Horaninow, Oporanthaceae Salisbury, Zephyranthaceae Salisbury

Eustephieae + Hymenocallideae + Stenomesseae + Eucharideae (Andean tetraploid clade): no palisade leaf mesophyll; x = 23 (tetraploid).

3J. Eustephieae Hutchinson

A of two lengths; seeds flattened, winged; (n = 21, etc.).

3/15: C. Andes (Peru, Bolivia, Argentina).

Hymenocallideae + Stenomesseae + Eucharideae: ?

3K. Hymenocallideae Small

Pollen grains auriculate [the two ends narrowed, with different sculpture]; testa thick, spongy, vascularised, phytomelan 0 (+ - Leptochiton); (n = 19, 20, 22), chromosomes 4-11.8 µm long.

3/65: Hymenocallis (50). S.E. U.S.A., the Antilles, Central America to Bolivia.

3L. Stenomesseae Traub

(Velamen + - Pamianthe); leaves petiolate, lorate; staminal cup + (0); seeds flattened, obliquely winged.

8/62: Stenomesson (35). Andean South America S. to Bolivia, Costa Rica (1 sp.).

3M. Eucharidae Hutchinson

Leaves petiolate; seeds globose, turgid, coat lustrous; chromosomes 2.3-10.7 µm long.

4/28: Eucharis (17). Central America, the Andes S. to Bolivia.

• Amaryllidoideae are usually bulbous herbs that can be recognised by their rather fleshy and two-ranked leaves and their scapose, umbellate inflorescence of generally large flowers with six stamens and an inferior ovary.

Evolution. Amaryllidoideae form an important component of the distinctive Cape geophyte flora (Procheŝ et al. 2006) having about 100 species endemic there. Petiolate leaves have evolved at least six times in the family. Monosymmetry in the family seems to be very flexible, perhaps being under simple genetic control (Meerow et al. 1999), and reversals and parallelisms seem to be common. The flowers are protandrous.

Wind dispersal of the inflorescences is common in Amaryllideae, the rigid, radiating pedicels allowing the infructescences to bowl along. The testa is commonly massive, green, and with anomocytic stomata in Amaryllidinae; it is photosynthetic, while in Crinum it is the endosperm that is green and photosynthetic. Seeds of some species of Crinum lack a testa and may have a corky outer endosperm; such seeds can float and remain viable in sea water for up to two years, while seeds of other species lack the corky layer, sink fast and can germinate without very much in the way of water at all (Snijman & Linder 1996; Bjorå et al. 2006). In Calostemmateae the bulbil, a precociously-germinated embryo, is the dispersal unit. Gethyllis (Haemantheae: to include Apodolirion) have single flowers with a subterranean ovary; the many-seeded fruit is indehiscent, and may be sweetly scented when ripe.

Chemistry, Morphology, etc. Norbelladine alkaloids, unique to Amaryllidoideae, are tyrosine derivatives; they are responsible for the poisonous properties of a number of the species. Over 200 different structures are known, or which 79 or more are known from Narcissus alone (Bastida & Viladomat 2002 and other references in the same volume, also Martin 1987). There are often crystals of calcium oxalate in the epidermis.

The flowers of Galanthus are shown with the median member of the outer whorl in the adaxial position (Spichiger et al. 2004), see also the similar position in Hippeastrum and several other monosymmetric Amaryllidoideae. Some species of Phaedranassa have slit-monosymmetric flowers, with all the stamens, etc., leaving the flower via an abaxial slit in the perianth tube; I do not know details of the symmetry of such flowers. Flowers of some species of Crinum are monosymmetric. In Galanthus in particular the inner whorl of tepals is very different from the outer whorl, although both are petaloid. Note that the "coronal" structures of e.g. Hymenocallis (evascularised outgrowths of the filaments) and those of Narcissus (vascularised, not associated with the stamens) are quite different (e.g. Arber 1937). Haemanthus has tepals with a single trace.

Flowers of Gethyllis have up to 18 stamens, and chromosomes in a single nucleus may be 3-16.1 µm long. Crinum has cellular endosperm; Zephyranthes has tenuinucellate ovules. Raymúndez et al. (2008) described megasporogenesis and megagmetogenesis in Hymenocallis caribaea; the ovules is crassinucellate ("pseudocrassincellate"), the micropyle ia zig-zag, and the vascularized outer integument is massive. x = 11 may be the basal chromosome number for the family (Meerow et al. 2006).

For anatomy, see Arroyo and Cutler (1984), for pollen, see DÖnmez and Isik (2008), and for general information, see Meerow and Snijman (1998).

Because of the leaf fibers in Amaryllideae, the coverings of the bulbs produce highly-extensible cotton-like fibers when torn. In Strumaria and Carpolyza the bases of the filaments are adnate to the style, while in Strumaria and Tedingia the base of the style may be much inflated, even bulbous. It is unclear if some ovules are ategmic. A very long-tubular dropper cotyledon sheath may be produced during germination; in Boophone and Cybistes germination may occur while the seeds are still enclosed by the fruit.

Phylogeny. For the phylogeny of Allioideae, see Fay et al. (2006b); part of Ipheion is embedded in Nothoscordum. Phylogenetic relationships within Amaryllidoideae are [Amaryllideae [Cyrtantheae [Calostemmateae, Haemantheae, Gethyllideae [Eurasian Clade [Andean Clade, Extra-Andean Clade]]]]] see Meerow et al. (1999, 2000a, 2000b). Note, however, that relationships between major clades of American and some southern African members are not well understood. Furthermore, Meerow et al. (2006) found that the inclusion of Hannonia, Lapiedra and Vagaria destabilised relationships in the European clade; Lledó et al. (2004) included the last two in Galantheae. Meerow and Clayton (2004) discuss relationships among African taxa.

Meerow and Snijman (2001, see also 2006) discuss relationships within Amaryllideae; Amaryllis and Boophone are successively sister to the rest of the tribe. Note that the former genus differs from other Amaryllidineae in not having a green testa, etc. Meerow et al. (2003) outline the phylogeny of Crinum, the only pantropical member of Amaryllidaceae. For a phylogeny of Crinum, see Kwembeya et al. (2007).

For relationships in Haemantheae, see Conrad et al. (2006). Meerow et al. (2006) provide a phylogeny for the Eurasian Clade, which includes daffodills, snowdrops, etc. The main dichotomy separates the Central and East Asian Lycorideae from the rest, which centre on the Mediterranean region. ITS and ndhF phylogenies are not congruent (Meerow & Snijman 2006). For a phylogeny of Galantheae, see Lledó et al. (2004).

Within Hippeastreae, Worsleya and Griffinia (Griffinieae: n = 10, 21; velamen + [Worsleya]; flowers blue; seeds whitish, globose, turgid [Griffinia]) are morphologically isolated and have an isolated position in Meerow et al. (2000a: as the "Hippeastroid" clade).

Classification. For the infrafamilial classification of Amaryllidaceae, I follow Chase et al. (2009: c.f. Fay and Chase 1996 for Allioideae). For a classification of Amaryllideae, see Meerow and Snijman (2001, 2006), and for generic limits in Galantheae, see Lledó et al. (2004).

ASPARAGACEAE Jussieu, nom. cons.   Back to Asparagales

153/2480. World-wide, but not Arctic - seven groups below.

Evolution. Stem group Asparagaceae s.l. are dated to ca 91 million years before present, divergence within crown group Asparagaceae s.l. begins ca 89 million years before present (Janssen & Bremer 2004). Eguiarte (1995), however, suggested that Agavaceae-Nolinaceae - the two are members of the two main clades here - diverged only some ca 47 million years ago,

Phylogeny. These seven subfamilies form a rather well supported clade in Fay et al. (2000: Hesperocallis not included), but there are no obvious characters for it. However, it is possible that "endosperm helobial, thick-walled, pitted, hemicellulosic" should be placed at this level. For details of relationships, see also Jang and Pfosser (2002) and Bogler et al. (2006). Fay et al. (2000), Pires et al. (2001), and Pires and Sytsma (2002) discuss uncertainties as to the immediate sister taxon to Themidaceae. Aphyllanthes has a very long branch in the three-gene analysis of Fay et al. (2000), and its phylogenetic position is unclear, indeed, its removal from some analyses rather dramatically changes support values (Chase et al. 2006). A position close to Hyacinthaceae was also found by McPherson and Graham (2001), but Pires et al. (2006) place it sister to Laxmanniaceae, but with only weak support. Themidaceae + Hyacinthaceae appear a moderately well supported pair in Fay and Chase (1996) and Meerow et al. (2000), but only weak support in the two-gene analysis of Jang and Pfosser (2002: Aphyllanthus not included) and in Chase et al. (2006) and Pires et al. (2006).

Classification. In addition to the absence of much in the way of synapomorphies for the clade, most of the subfamilies are difficult to recognise, for the most part having rather undistinguished "lily"-type flowers. Within some of them, e.g. Asparagoideae and especially Nolinoideae and Agavoideae, there is considereable variation, several segregate families having been recognised in the past. Hence the use of Asparagaceae s.l. to refer to the entire clade is justified (cf. A.P.G. II 2003, A.P.G. III 2009). The subfamilial classification follows that in Chase et al. (2009b).

Aphyllanthoideae [Brodiaeoideae + Scilloideae] Agavoideae: ?

1. Aphyllanthoideae Lindley   Back to Asparagales

Flavonols +; 2ndary thickening +; stems alone photosynthetic, with parallel wax scales; leaves two-ranked, supervolute-subinvolute, as non-photosynthetic scales, ligulate, base?; inflorescence scapose, flowers multibracteolate, sessile; T marcescent, basically free; A adnate to base, pollen spiraperturate, infra-locular septal nectaries +, 1 position? ovule/carpel, micropyle?, stigma trifid, dry; seeds slightly flattened, exotestal cells large, isodiametric; endosperm ?, 0; n = 16; cotyledon photosynetic, terete, first leaf terete.

1[list]/1: Aphyllanthes monspeliensis. W. Mediterranean. [Photo - Flower © E. Bourneuf]

• Aphyllanthaceae may be readily recognised; the above-ground plant consists of tufts of scapose infloresences, the scapes being the main photosynthetic organs since the scarious leaves at their bases are non-photosynthetic; the inflorescence is few-flowered, the spreading, usually blue tepals being moderate in size (the whole plant looks rather like Sisyrinchium!).

The stomata are in bands down the scape. The tepals have but a single bundle. Is there chelidonic acid?

General information is taken from Conran (1998); he mentions endosperm development as being helobial, but there is no mention of this in Schnarf and Wunderlich (1939), which appears to be the only source of information for ovule features.

Synonymy: Aphyllanthaceae Burnett

Brodiaeoideae + Scilloideae: steroidal saponins +; leaves spiral; pedicels bracteate; ovules anatropous; endosperm helobial or nuclear; cotyledon not photosynthetic.

For some characters of this pair, see Fay and Chase (1996); laticifer-like structures may occur in both families.

2. Brodiaeoideae Traub   Back to Asparagales

Fibrous monopodial corm storing starch; laticifers +; mucilage cells?; leaves (unifacial - Brodiaea), sheath closed; inflorescence scapose, umbellate, with several scarious inflorescence and internal bracts, pedicels often articulated; (T free; corona +); A (3), connate and/or adnate to T, (filaments flattened), (ovary stipitate, adnate to T by flanges opposite the outer tepals; nucellar epidermal cells enlarged; nucellar cap +; inner integument 3+ layers across), stigma capitate to trifid, dry (wet - Bloomeria); seeds angular; (endosperm helobial - Muilla, Triteleia), "embryo short"; n = 5-12+; hypocotyl?, primary root persistent.

12[list]/62. S.W. North America, to British Columbia and Guatemala (map: see Moore 1953; Fl. N. Am. 26: 2002). Flower, Flower, Fruit.]

• Asparagaceae-Brodiaeoideae are like Amaryllidaceae-Allioideae in their scapose, umbellate inflorescence, rather small flowers, more or less connate tepals sometimes with a corona, and a superior ovary, but the plants have a fibrous corm, not a bulb, and lack the distinctive smell common in the latter family, and there are usually 4 or more inflorescence bracts that are not as all-enveloping as the ca 3 bracts of Allioideae.

Chemistry, Morphology, etc. Little is known of the chemistry of the subfamily. The inner integument is massive or not, ditto base of the nucellus.

Information is taken from Moore 1953 (morphology), Berg (1978, 2003: embryology), Rahn (1998: general), Pires et al. (2001, 2002: phylogeny and morphological evolution) and Pires and Sytsma (2002: ditto).

Chemistry, Morphology, etc. There are two major clades in the Brodiaeoideae, albeit they have only only moderate support. One has a long tepalline tube and the other has appendages on the bases of the filaments that form a nectar cup; both characters arise in parallel in the opposing clade (Pires & Sytsma 2002). When the tepalline tube is adnate to the stipitate gynoecium, three narrow, ?nectar-containing tubes are formed.

Previous Relationships. Themidaceae/Brodiaeoideae have often been included in Amaryllidaceae-Allioideae because of their superficially similar umbellate inflorescence and rather "typical"-appearing and undistinguished monocot flowers (e.g. Takhtajan 1997).

Synonymy: Themidaceae Salisbury

3. Scilloideae Burnett  Back to Asparagales

Plant bulbous geophytes, roots often contractile, bulb leaves sheath closed or not; polyhydroxyalkaloids, flavones +, flavone C-glycosides +; little sclerenchyma in the leaf (well-developed); mucilage cells +; (leaf waxes with parallel platelets); inflorescence usu. scapose, (branched) raceme or spike, pedicels not articulated, bracteole 0; (corona +), 1-many ovules/carpel, stigma capitate to punctate and papillate; testa multi-layered; chromosomes 1.2-18 µm long; (hypocotyl 0; collar rhizoids +).

41-70[list]/770-1000 - six groupings below. Predominantly Old World, Mediterranean climates, esp. S. Africa and the Mediterranean, to Central Asia and Burma; some in South America.

3A. Oziroëeae M. W. Chase, Reveal & M. F. Fay

A basally connate and adnate to C; seeds rounded; embryo as long as seed; n = 15, 17; cotyledon?.

1/5. Western South America (map: see opposite, green colour).

See Guaglianone & Arroyo-Leuenberger (2002) for a revision (and distribution map) of Oziroëe.

Ornithogaleae + Urgineeae + Hyacintheae: rhexigenetic lacunae +; also styloids +.

3B. Ornithogaleae Rouy

Cardenolides +; (A 3; filaments flat, with appendages); seeds flattened/angled; protein crystals in nucleus; n = 2-10+; cotyledon photosynthetic or not.

4/312: Ornithogalum (160), Albuca (110-140). Europe, W. Asia, Africa.

Synonymy: Ornithogalaceae Salisbury

3C. Urgineeae Rouy

Bufadienolides +; bracts spurred [as small leaves in Bowiea]; seeds flattened/winged, testa brittle, not tightly adherent to endosperm; n = 6, 7, 10+.

2(-3?)/105: Drimia (100, inc. Urginea). Mainly Africa, Madagascar, the Mediterranean to India (map: from Pfosser & Speta 2001). [Photos - Boweia Collection.]

Asparagus (Asparagoideae) also has stem/branch leaves that have a backwardly-directed process like those on the leaves and/or bracts of Urgineeae.

Synonymy: Drimyidaceae Baillon, Hyacinthaceae Borkhausen

3D. Hyacintheae Dumortier

Homoisoflavanones +; (leaves with pustules or coloured spots); embryo sac type variable; seeds often rounded.

a. Pseudoprospero

Prophylls +; 2 ovules/carpel; 1 seed/loculus; n = 9; cotyledon not photosynthetic.

1/1: Pseudoprospero firmifolium. E. South Africa.

b. Massoniinae Bentham & Hooker f.

(Bracteoles +); (flowers monosymmetric); ovary and style sulcate, 2-many ovules/carpel; (elaiosomes +); n 5-10+; cotyledon not photosynthetic (photosynthetic).

Ca 10/235: Lachenalia (110), Ledebouria (70). Africa S. of the Sahara, Ledebouria to India (map: from Venter 2008).

Some species of Daubenya (Massoniinae) have a filament tube.

Synonymy: Eucomidaceae Salisbury, Lachenaliaceae Salisbury

c. Hyacinthinae Parlatore

(Bracts 0), prophylls quite common; 2-8(-many) ovules/carpel; (elaiosomes +); n = 4-8+; cotyledon photosynthetic or not.

21/265: Muscari (50), Bellevalia (50), Scilla (30), Prospero (25). Europe to the Mid (Far) East, North Africa (map: above, area in red, from Meusel et al. 1965, incomplete). [Photo: Scilla Collection.]

Synonymy: Scillaceae Vest

• Scilloideae are bulbous monocots with rather fleshy, mucilaginous leaves that are all basal. The scapose, racemose inflorescence bears for the most part rather undistinguished monocot flowers with more or less connate tepals; the fruit is a capsule.

Evolution. There are about 300 species of Scilloideae in the Cape flora (Procheŝ et al. 2006).

Lachenalia has monosymmetric flowers in which the median member of the outer whorl is in the adaxial position. The same is true of the remarkable monosymmetric flowers of Massonia (Daubneya) aurea that are on the outside of the inflorescence. These flowers have the three abaxial tepals greatly enlarged, while the inner flowers are polysymmetric and the tepals form a simple, lobed tube.

Chemistry, Morphology, etc. Some species of Scilloideae have terete, unifacial leaves, and even the bulb scales of some species of Rhadamanthus (= Drimia) are terete... Vegetative variation - in both leaf and bulb - is also considerable in Ledebouria (Venter 2007). However, there is little anatomical variation in the subfamily. Although mucilage cells are particlarly common in Scilloideae, they also seem to occur elsewhere (Lynch et al. 2006). Taxa that have tepals with single vascular traces are common here. Chromosome length can be bi- or even trimodal in the one karyotype. The leaves of seedlings are two-ranked.

Information is taken from Speta (1998a: subfamilial classification of Hyacinthaceae, 1998b: general), 2001 (subfamilial characters) and Pfosser and Speta (1999); for chemistry, see Kite et al. (2000) and Pfosser and Speta (2001); and for floral morphology in Ledebouriinae, see Lebatha and Buys(2006).

Phylogeny. There is little well-supported structure along the backbone of what are here called Hyacintheae and again within Hyacinthineae in the trnL-F spacer analysis of Wetschnig et al. (2002); the positions of Ornithogaleae and Urgineeae were also unclear. The topology [Oziroëeae [Ornithogaleae [Urgineeae + Hyacintheae]]] has moderate support in Manning et al. (2004). For phylogeny, see also Pfosser et al. (2003).

Classification Note that there is considerable disagreement over generic limits in Scilloideae; are there 15 or 45 genera in sub-Saharan Africa? (e.g. Stedje 2001a, b; Pfosser & Speta 2001; Lebatha et al. 2006). See Speta (1998a) for the dismemberment of Scilla. Manning et al. (2004) provide a generic synopsis of the family in sub-Saharan Africa that integrates some morphology with relationships and, like them, I have taken a generally broad view of genera. However, there are unresolved issues that include sampling, whether or not floral syndromes distort ideas of relationships (and so what effect characters taken from these syndromes have in combined analyses), the consequences of maintaining well-known generic names like Albuca and Galtonia as knowlege of phylogeny becomes clearer, and the role cytological data should play. As to Albuca, the genus is recognized in the recent reclassification of Ornithogaloideae by Manning et al. (2009).

See Guaglianone & Arroyo-Leuenberger (2002) for a revision (and distribution map) of Oziroëe.

Previous Relationships. Chlorogaloideae, until recently included in Hyacinthaceae/Scilloideae (e.g. Pfosser & Speta 1999), are here included in Agavoideae. The homoisoflavanones found in Scilloideae are rather uncommon in flowering plants, but they are also to be found in Camassia (Agavoideae, ex Chlorogaloideae) and Ophiopogon (Nolinoideae).

4. Agavoideae Herbert   Back to Asparagales

Plant rhizomatous; endosperm helobial, thick-walled, pitted, hemicellulosic.

23/637 - five groups below. More or less world-wide, esp. S.W. North America, few in Malesia, N. Australia, not cold temperate, New Zealand, etc. (map: see Ying et al. 1993; García-Mendoza & Galván V. 1995; Fl. N. Am. 26: 2002; Seberg 2007).

4A. Anemarrhena

Leaves ?spiral, base?; inflorescence a subspicate panicle; T ± free; A 3, opposite and adnate to middle of inner T, 2 apotropous ovules/carpel; seeds angled; embryo curved; n = 11; hypocotyl 0.

1/1: Anemarrhena asphodeloides. N. China, Korea.

Ubisch bodies are present in Anemarrhena, so there is probably a glandular tapetum. Information is taken from Conran and Rudall (1998 - confusion over stamen position) and Rudall et al. (1998b); Anemarrhenaceae were included in Anthericaceae (Agavoideae-Anthericum below) by Takhtajan (1997).

Synonymy: Anemarrhenaceae Conran, M. W. Chase & Rudall

Agave, etc. + Behnia + Herreria, etc. + Anthericum, etc.: nucellar cap, hypostase +.

This group has 100% support in three- and four-gene trees (Chase et al. 2000a; Fay et al. 2000; Bogler et al. 2006).

Agave, etc. + Hesperocallis: seeds flattened [?all]; n = 24, 30, chromosomes 0.4-10 µm long [bimodal: 4-6 long, rest short]

4B. Agave, etc.

Also caulescent, (bulbs, tunicated or not); non-protein amino acids, saponins, (homoisoflavanones - Chlorogaloideae), flavonols +; secondary growth +; also styloids +; (stomata para- or tetracytic), cuticular wax rodlets parallel; leaves spiral, (petiolate; vascular bundles inverted; margins serrate), apex (pungent-)pointed, base ?; inflorescence usually branched, and/or flowers in pairs or fascicles, (pedicel articulated - Chlorogaloideae); flowers large, (monosymmetric), tapetal cells several-nucleate, pollen semitectate, (operculate), ovary superior to inferior, many crassi- or tenuinucellate ovules, obturator + [Beschornea], (styles +), stigma wet to dry; (capsule septicidal - some Yucca; berry), T marcescent; (endosperm thin-walled; perisperm +, oily - Yucca, Agave); (cotyledon non-photosynthetic - Funkia), hypocotyl to 4 mm long, collar rhizoids +, primary root often branched.

10/340: Agave (210), Yucca (50). C. U.S.A. to N. South America, mostly S.W. North America, also East Asia (Hosta). [Photo - Flower]

Synonymy: Agavaceae Dumortier, nom. cons., Chlorogalaceae Doweld & Reveal, Funkiaceae Horaninov, nom. illeg., Hesperocallidaceae, Hostaceae B. Mathew (endosperm cells thin-walled), Yuccaceae J. Agardh (perisperm +)

Behnia + Herreria, etc. + Anthericum, etc.: ?

4C. Behnia

Secondary thickening +; tannin cells 0; velamen 1-layered; vessel elements also in the stem; leaves two-ranked, "supervolute", petiolate, with midrib and transverse secondaries, base not sheathing; dioecious; T marcescent, not twisting; A adnate to base of T, 2-3 tenuinucellate ovules/carpel, micropyle?, stigma 3-lobed, wet; fruit a berry; seeds angular, phytomelan 0, testa and tegmen thin-walled; endosperm walls not pitted and hemicellulosic; n = ?

1/1: Behnia reticulata. S.E. Africa.

Behnia was included in Luzuriagaceae (Liliales here) by Taktajan (1997), butit had also been placed in other families of that order and of Asparagales (Bogler et al. 2006); its broad leaves and campanulate flowers make it phenetically rather odd..

General information is taken from Conran (1998); details of ovules are unknown.

Synonymy: Behniaceae Conran, M. W. Chase & Rudall

Herreria, etc. + Anthericum, etc.: ?

4E. Herreria, Herreriopsis

Stems usu. climbing, prickly; saponins +, chelidonic acid?; (vessel elements in stem); mucilage cells 0; cuticular wax rodlets parallel; leaves ?spiral, ?cladodes, in fascicles, sheath?; pedicels not articulated; T and A free, 1-many [?type] ovules/carpel; fruit a septicidal capsule; seeds flattened; "embryo short"; n = 27, dimorphic [one large], chromosomes 0.7-3.7 µm long [Herreria]; "germination epigeal".

2[list]/9. South America (Brazil southwards), Madagascar. [Photo - Fruit]

The outer tepals of Herreriopsis have sac-like bases - possibly tepalline nectaries. The leaves are described as being cladode-like (Conran 1998) or cladodes (Stevenson in Takhtajan 1997).

Much information is taken from Conran (1998); the embryology is largely unknown.

5. Anthericum, etc.   Back to Asparagales

Rhizome short; chelidonic acid +; (velamen +); (vessel elements in the stems); mucilage cells +, tannin cells 0; cuticular wax rodlets parallel; leaves spiral to two-ranked, base sheathing; inflorescence thyrsoid (raceme); (pedicels not articulated); (flower monosymmetric), (T tube 0), 2-many ovules/carpel, embryo sac haustoria common, stigma dry; T persistent in fruit; seeds angular or flattened; tegmen?; n = 7, 8, 10, 11, 13-15, etc., chromosomes 2-10(-13.8) µm long; cotyledon not photosynthetic, coleoptile + [Chlorophytum].

8/285: Chlorophytum (150 - styloids +), Anthericum (65), Echeandia (60). More or less worldwide, but not cold temperate, few in Malesia, N. Australia, not New Zealand, etc. [Photo - Inflorescence] [Photo - Flower]

Synonymy: Anthericaceae J. Agardh

Evolution. For information on the Yucca-yucca moth (Tegeticula, Prodoxidae) association, a textbook example of mutualism, which may be some 40 million years old, see Pellmyr et al. (1996, 2007), Pellmyr and Leebens-Mack (1999), Pellmyr (2003), Gaunt and Miles (2002) and Althoff et al. (2006); there may have been another and recent radiation of yucca moths 3-2 million years ago. However Good-Avila et al. (2006) suggest that Agave et al. are only some 26-20 million years old, and Yucca 18-13 million years old; Wikström et al. (2001) give an age for the whole Agavoideae clade of some 35 million years before present that is also fairly recent compared to the first set of dates; Rocha et al. (2006) suggest ca 12.75 million years for the age of Agave etc. and ca 10.2 million years for Agave s.l. (Hesperaloe and everything above in the tree - Bogler et al. 2006; cf. also Smith et al. 2008); there are other possibilities for dates.

Good-Avila et al. (2006) discuss diversification in both Agave, which they suggest may be connected with the adoption of bat-pollination, and Yucca (see also Rocha et al. 2006). However, Smith et al. (2008) suggest that diversification was not significantly different in Yucca, with its 34 species, and Agave, with some 240 species. Note that close relatives of yucca moths are also found on Dasylirion and Nolina (Nolinoideae, this page) and other Prodoxidae on Saxifragaceae (Saxifragales).

Nobel (1988) discussed the eco-physiology of agaves and their relatives.

Chemistry, Morphology, etc. The raphides of Agave are hexagonal in transverse section. The flowers of Agave are shown with the median member of the outer whorl in the adaxial position (Spichiger et al. 2004). Camassia at least has single-trace tepals. In Polianthes the tapetal cells are multinucleate. In Hosta the stamens are sometimes inserted on the ovary. Germination of the pollen grain via the proximal pole has been reported in Beschorneria (Hesse et al. 2009). Furcraea has nuclear endosperm.

See Judd et al. (2007) for general information. For additional information about this group, especially the part that has been considered Agavaceae s. str. in the past, i.e. Agave, Yucca and their immediate relatives, see Alvarez & Köhler (1987: pollen), (Cave 1948: embryology), and Verhoek (1998: general), Kubitzki (1998b - Hostaceae), Speta (1998 - Hyacinthaceae - Chlorogaloideae).

Traub (1982) noted that Hesperocallis undulata smells of onions, and he even associated it with his Alliales. The genus was geographically odd in Hostaceae s. str., which is where other workers had placed it (cf. Kubitzki 1998b), but not in Agavoideae as here circumscribed; now Hosta is geographically a little odd!

The ovary and fruit of Leucocrinum (Anthericum group) are below the surface of the ground (Bogler et al. 2006). At least some mitochondrial genes show an accelerated rate of change (G. Petersen et al. 2006). Some information is taken from Conran (1998); ovule morphology in the group is taken from Leucocrinum alone.

Phylogeny. For relationships within this clade, see Pires et al. (2004) and especially Bogler et al. (2006: 2- and 3-gene analyses, the latter with missing data, but overall the same topology). I have followed the latter - which see for more details - in the topology above. Support for the subfamily as a whole is only 75%, that for the [Behnia + Herreria, etc. + Anthericum, etc.] clade 87%, and that for [Herreria, etc. + Anthericum, etc.] only 51% (and less in the two-gene tree); however, other nodes have close to 100% support, and there is a fair amount of detail resolved in relationships around Agave and Yucca (see below). Largely similar relationships were found by G. Petersen et al. (2006c) in their analysis of variation of four mitochondrial genes that are evolving particularly quickly in this clade.

The circumscription of group 4b above, Agave etc., corresponds to that of Agavaceae s.l. in Bogler et al. (2006). Genera like Camassia, etc. (ex Hyacinthaceae - Chlorogaloideae) are included; note that these latter taxa have rhexigenetic lacumae (Lynch et al. 2006) like many Hyacinthaceae (Scilloideae) themselves. Hesperocallis undulata is sister to the rest of the Agave, etc., clade (Bogler et al. 2006). (In Smith et al. 2008 Agavaceae s.l. includes Hosta etc. and excludes Anthericaceae, although support for Agavaceae so delimited was weak, and that for the still broader circumscription adopted here was stronger.) For other phylogenetic work on this group, see Bogler and Simpson (1996: molecular) and Sandoval (1995: morphological).

Classification. The broad concept of Agavoideae adopted here may not seem very satisfactory, but I fear that none of the alternative solutions is much better when it comes to communication. Agave should probably include Polianthes, Manfreda, etc., see e.g. Bogler and Simpson (1995), Bogler et al. (2006) and Rocha et al. (2006). Paradisea (ex Asphodelaceae/Asphodeloideae) is to be included here in the Anthericum group (e.g. Chase et al. 2000b).

Lomandroideae + Asparagoideae + Nolinoideae: steroidal saponins; pedicels articulated; fruit a capsule; endosperm helobial, thick-walled, pitted, hemicellulosic.

Phylogeny. There is moderate support for this clade in the four-gene chloroplast tree of Fay et al. (2000).

5. Lomandroideae Thorne & Reveal   Back to Asparagales

(Naphthoquinones +); (secondary thickening +; vessel elements in leaves); (inner T fimbriate; T connate basally; A adnate to tube), infra-locular septal nectaries +, especially basal dermal nucellar cells enlarged, nuclei of antipodals large; T persistent in fruit; seeds rounded to angular; chromosomes 0.6-2.4 µm long [records incomplete]; cotyledon photosynthetic or not, (coleoptile +; first leaves reduced).

14-15[list]/178. Predominantly Australian, also Madagascar, India, South East Asia to the Pacific, and South America.

5A. Lomandra group

Lamina with sclerenchymatous ribs extending from the inner sheath of the vascular bundle to the surface, outer bundle sheath with enlarged cells; leaves two-ranked, flat or curved, (margins prickly), (base auriculate); pedicel articulated (not - Xerolirion), 1-2 ovules/carpel, nucellus with axially oriented central conducting passage, stigma wet; testa thin, tegmen brown, collapsed, cellular; phytomelan 0; endosperm hemicellulosic; n = 7-9, chromosomes 2-7 µm long.

5/65: Lomandra (50). Australia, New Guinea, New Caledonia. [Photo - Inflorescence © K. Stüber.]

Synonymy: Lomandraceae Lotsy

5B. Laxmannia group

Plant with (ecto) vesicular-arbuscular mycorrhizae; storage roots +; mucilage +; leaves spiral, supervolute or conduplicate, (petiolate), (ligulate); flowers single or in groups, pedicel articulated or not; (anthers porose), nucellus with axially oriented central conducting passage, stigma wet; (seed arillate), testa with phytomelan, exotesta often papillate, rest of testa cellular, tegmen thin; endosperm thin-walled; n = 4, 11, chromosomes 0.5-2 µm long.

8/92: Thysanotus (50), Arthropodium (20). South East Asia to Australia, New Zealand and the Pacific.

Synonymy: Eustrephaceae Chupov, Laxmanniaceae Bubani

5C. Cordyline group

Rosette herbs to trees; storage roots +; mucilage +; stomata paracytic; leaves spiral, supervolute or conduplicate, (pseudopetiolate); flowers single, pedicel not articulated; testa with phytomelan, anatomy?; endosperm ?; n = 3, 6, 19 (stamens dimorphic), chromosomes 0.5-2.4 µm long.

2/17. India to the Pacific and New Zealand, tropical America. [Photo - Habit, Flower.]

Chemistry, Morphology, etc. There is considerable variation in seedling morphology, even within individual groups (Conran 1998).

Some information is taken from Chanda and Ghosh (1976: pollen, as Xanthorrhoeaceae), Rudall and Chase (1996), Chase et al. (1996), Conran (1998, as Lomandraceae), and Rudall (2000: ovule).

The leaf of Lomandra and its relatives has sclerenchymatous ribs extending from the inner sheath of the vascular bundles (cf. also Cordyline?); in Dasypogonaceae this sheath is absent, in Xanthorrhoeaceae s. str. it comes from the mesophyll, although the leaves of all three are xeromorphic and superficially similar. The pollen of Lomandra is very variable, sometimes being spiraperturate (cf. Aphyllanthes).

Classification. I have tentatively recognised three groups above, partly based on morphology, and partly based on molecular data (the Cordyline group - see Chase et al. 1996).

Xerolirion, from southwest Australia, has solitary, terminal carpellate flowers, staminate flowers in cymes, there is a single ovule per carpel, and it lacks silica bodies although there are cell wall ferulates; it is not entirely clear where it should be placed.

Asparagoideae + Nolinoideae: (velamen +); flowers rather small[!]; x = 10.

Baccate fruits containing seeds that lack phytomelan are common here, but I do not know at what level they are apomorphic. Indeed, since the capsular Hemiphylacus and [Comosperma + Eriospermum] are respectively sister to other Asparagaceae and Ruscaceae, baccate fruits are probably derived (cf. Judd et al.).

Chemistry, Morphology, etc. For phylloclade development in Asparagus and Ruscus and its relatives, see Cooney-Sovetts and Sattler (1987).

6. Asparagoideae Burmeister   Back to Asparagales

Horizontal or vertical rhizome; flavonols, saponins + +; vessels also in stem; cuticular wax rodlets parallel; leaves spiral (scarious and subtending phylloclades - Asparagus), base not sheathing; (plant mon- or dioecious) inflorescence ± fasciculate or paniculate; T tube at most short; A basally adnate to T (3, opposite inner T, outer A staminodial), 2-several ovules/carpel, embryo sac curved, nucellar epidermal cells enlarged, stigma wet or dry; fruit usu. a berry; seed rounded to ± angled (?endotestal); n = also 56, chromosomes 1-3 µm long.

2[list]/165-295: Asparagus (160-290!). Old World, but hardly in Australasia, also Mexico (map: from Hernandez S. 1995; Hultén & Fries 1986; FloraBase 2007; Seberg 2007). [Photo - Flower, Fruit.]

Evolution. Fukuda et al. (2005) discuss diversification in Asparagus; this seems to have been rapid and to have started in S. Africa.

Chemistry, Morphology, etc. Methyl mercaptans are known from Asparagus. The prophylls ("bracts") at the bases of the pedicels in Hemiphylacus are described as being lateral (Hernandez S. 1995). Hemiphylacus (G opposite inner T, fruit a capsule; n = 56) used to be placed in Asphodelaceae.

Some information is taken from Kubitzki and Rudall (1998) and Rudall et al. (1998b).

Synonymy: Hemiphylacaceae Doweld

7. Nolinoideae Burnett   Back to Asparagales

Flavonols, (azetidine-2-carboxylic acid [non-protein amino acid]), saponins +; (velamen +; vessels in stem - many ruscoids); (vascular bundles amphivasal); also styloids +; cuticular wax rodlets parallel; leaves (scarious), spiral or two-ranked (opposite - Polygonatum oppositifolium), (petiolate; broad, venation reticulate), margins spiny or not, (base not sheathing); inflorescence also racemose; (flowers 2-merous - some Maianthemum; T not connate; T-13, corona + [Aspidistra]; with a single trace); (A adnate to base of tube; connate); pollen often inaperturate/diffuse sulcate; stigma (much expanded), wet; ovules 1-6(-many; tenuinucellate)/carpel, (funicle or ovary wall obturator); fruit also a berry (± a drupe); seeds rounded (angled), (sarcotesta - Ophiopogoneae; testa 0 - Dracaena), phytomelan 0, (phlobaphene +); (endosperm nuclear); n = 5-7, 9, 18-21, chromosomes 0.5-19 µm long (bimodal); cotyledon not photosynthetic, (coleoptile +), primary root well developed, branched or not.

26/475: Dracaena (inc. Pleomele, Sanseviera, 100), Eriospermum (100 - see Perry 1994), Aspidistra (90), Polygonatum (60), Ophiopogon (55). N. hemisphere, esp. Southeast Asia-Malesia (Convallariaceae s. str.), Europe and the Near East (Ruscaceae s. str.), S.W. North America (Nolinaceae s. str.), Africa, esp. the Cape and S.W. (Eriospermaceae s. str.) (map: from Meusel et al. 1965; Hultén & Fries 1986; Perry 1994, incomplete). [Photo - Ruscus Flower, Eriospermum Flower © M. Elvin.]

Evolution. The flowers of Aspidistra, sometimes borne beneath litter and with a short corona at the apex of the perianth tube, also often have a large, fungiform stigma and perianth tube with included anthers, or anthers converging towards the centre of the flower, in both cases with easy access to the stigma apparently blocked; these flowers may be pollinated by amphipods (Conran & Bradbury 2007 and references) and/or fungus gnats; they look rather like flowers of Asarum and some Burmanniaceae. There are also more conventional sub-rotate flowers in the genus which have the stamens and stigma/style grouped in the centre, and some species have a dozen or more tepal lobes.

Vegetative variation is particularly impressive. Nolina (ex Nolinaceae) has secondary growth and is tree-like, while the initiation of the vascular system in the rhizome of Ophiopogon is similar to that in palm stems (Pizzolato 2009). The leaf blades of some species of Eriospermum have the most remarkable enations on the upper surface. These include fungiform protrusions on the small, crisped, ovate and fleshy blade (E. titanopsoides), a much-branched structure to 12 x 7.5 cm on a much smaller blade (E. ramosum), a bundle of enations with stellate hairs (E dregei), and paired enations that look as if they should grace the helmets of the Valkyries (E. alcicorne: see Perry 1994 for more details). Many other taxa, including Maianthemum, have more or less broadly elliptic leaf blades. Finally, Ruscus and its immediate relatives have cladodes, the flowers being born in the middle of a tough, more or less elliptical leaf-like structure. The prophylls are lateral or in some interpretations completely adnate to the axillary shoot, together they form an expanded cladode (Arber 1930). In any event, the leaves proper are small and scarious and subtend the cladode-like structures (cf. Asparagus above).

Chemistry, Morphology, etc. Convallarieae are monopodial. Dracaena and relatives have resin canals. The outer integument is only two cells thick, as in Bowiea. The absence of septal nectaries in some taxa of this group may be connected with the presence of prominent ovary wall obturators; the latter are possibly derived from the former. In Liriope, etc. (Ophiopogoneae), the seeds, with their fleshy testa (see above), are exposed early in development, so they are semi-gymnospermous. Finally, in Peliosanthes teta, the only species in Peliosanthes, the ovary varies from superior to inferior (Jessop 1976), although some recognise more species in the genus.

Ruscus and its immediate relatives also have chrysophanol in the roots; filaments connate, anthers extrose, in carpellate flowers the filaments of the staminodes almost completely enclose the gynoecium; embryo short; n = 20.

Additional information can be found in Björnstad (1970), Lu (1985), Tillich (1995: seed, etc.), Bogler and Simpson (1996: relationships of Nolinaceae, Dracaenaceae, etc.), Bos (1998: Dracaenaceae), Conran and Tamura (1998: Convallariaceae), Bogler (1998: Nolinaceae), Terry and Rudall (1998: Eriospermaceae), Yeo (1998: Ruscaceae s. str.), Rudall & Campbell (1999: floral morphology), Judd et al. (2002: general), Judd (2003: Ruscaceae S.E. U.S.A.) and Yamashita and Tamura (2004: chromosome evolution in Convallarieae).

Phylogeny Comosperma, ex Anthericaceae (= Agavoideae), comes here. It and and the very distinct Eriospermum are likely to be sister to the rest of the family; both have capsules and hairy seeds. Note, however, that the hairs on the seeds develop in different ways, and Comosperma has two tenuinucellate apotropous ovules/carpel, n = 20 vs. n = 7, etc., the two genera seemingly being unrelated (Rudall 1999.) The poorly understood Peliosanthes may also be in turn sister to the rest of the family (molecular data alone, e.g. Jang & Pfosser 2002). Relationships within the other Ruscaceae are poorly resolved, although major clades seem to correspond largely with tribes (see Conran & Tamura 1998) and the families that are also included here. However, Convallarieae may be paraphyletic with Aspidistreae and Ruscus and relatives embedded (Yamashita & Tamura 2000 - Eriospermum was not included; Rudall et al. 2000b); in Ruscus and immediate relatives a mitochondral cox2 intron is missing (Kudla et al. 2002).

Classification. There has been debate over the generic limits of Maianthemum, however, a broad circumscription seems appropriate; there is little support for infrageneric groupings within the clade that also includes Smilacina and the combined clade itself is well supported as being monophyletic (Kim & Lee 2007; Meng et al. 2008). For information on Aspidistra, see Hou et al. (2009) and references; there is a monograph in Li (2004).

Synonymy: Aspidistraceae Endlicher, Convallariaceae Horaninow, Dracaenaceae Salisbury, Eriospermaceae Endlicher (hypocotylar tuber; leaf sometimes with enations; pedicels not articulated; fruit a capsule; testa hairy; endosperm 0, perisperm +, embryo massive; n = (5-)7 (9, 10); cotyledon unifacial, photosynthetic), Nolinaceae Nakai, Ophiopogonaceae Endlicher (apotropous ovules), Peliosanthaceae Salisbury, Polygonataceae Salisbury, Ruscaceae Sprengel, nom. cons., Sansevieraceae Nakai, Tupistraceae Schnizlein