SOLANALES Dumortier Main Tree, Synapomorphies

Habit Usually bisexual (sometimes monoecious, andromonoecious or dioecious), evergreen trees, shrubs or lianas, perennial, biennial or annual herbs (sometimes climbing).

Vegetative anatomy Roots diarch. Phellogen ab initio superficially or deeply seated. Secondary lateral growth sometimes anomalous (via cylindrical cambium or concentric cambia) or absent. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits usually alternate (sometimes scalariform), simple or bordered pits. Imperforate tracheary xylem elements fibre tracheids, libriform fibres or tracheids with simple or bordered pits, usually non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, aliform, lozenge-aliform, confluent, reticulate, unilateral or banded (sometimes absent). Wood often fluorescent. Intraxylary phloem usually present. Sieve tube plastids S type. Nodes 1:1–3, unilacunar with one to three leaf traces (rarely multilacunar with several traces). Latex cells and articulated laticifers sometimes frequent. Resins present or absent. Cystoliths often present. Prismatic or acicular calciumoxalate crystals, druses, styloids and crystal sand often abundant.

Trichomes Hairs unicellular or multicellular, uniseriate, simple, furcate, dendritic or stellate (sometimes peltate-lepidote); multicellular glandular hairs often abundant.

Leaves Usually alternate (spiral; sometimes opposite), usually simple (sometimes pinnately or palmately compound or), entire or pinnately lobed, sometimes coriaceous, often with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate or palmate. Stomata usually anomocytic or paracytic (sometimes anisocytic, diacytic, tetracytic or parallelocytic). Cuticular wax crystalloids? Domatia as hair tufts or absent. Mesophyll without mucilage cells, with or without sclerenchymatous idioblasts. Leaf margin and leaflet margins serrate or entire.

Inflorescence Terminal or axillary, usually simple or compound dichasia or cincinni (rarely corymbose or paniculate cymose, capitate or spicate), or flowers solitary terminal or axillary. Floral prophylls (bracteoles) rarely absent.

Flowers Actinomorphic to obliquely zygomorphic. Usually hypogyny (rarely epigyny or half epigyny). Sepals (three to) five (to seven), usually with imbricate or plicate (rarely valvate or open) aestivation, often persistent, free or connate. Petals (three to) five (to seven), with contorted, plicate, imbricate, induplicate or valvate aestivation, usually connate into infundibuliform, campanulate or tubular corolla. Nectariferous disc intrastaminal, annular or cupular (sometimes tuberculate, discoid or pulvinate, rarely absent).

Androecium Stamens (three to) five (to seven), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, from tepals or adnate to corolla (epipetalous), sometimes of two different lengths. Anthers often connivent around style, usually dorsifixed to basifixed (rarely ventrifixed), usually non-versatile, tetrasporangiate, usually introrse (rarely extrorse), longicidal (dehiscing by longitudinal slits or sometimes by two apical slits) or poricidal (dehiscing by one apical pore, or two separate apical pores). Tapetum secretory. Female flowers sometimes with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3–5(–8)-colpate or (2–)3–5(–6)-colpor(oid)ate (sometimes polycolpate, pantoporate or inaperturate), usually shed as monads (rarely tetrads), usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with usually columellate (occasionally granular) infratectum, perforate, reticulate, punctate or striate, psilate, spinulate or echinate. Pollen tube with callose plugs.

Gynoecium Pistil composed of usually two (sometimes five, rarely one or three) connate antepetalous carpels, often displaced relative to floral symmetry. Ovary usually superior (rarely inferior or semi-inferior), usually unilocular or bilocular (rarely trilocular, quinquelocular or multilocular [due to secondary septa]). Style single, simple, usually terminal (rarely gynobasic), or stylodia two (to five) usually more or less connate. Stigma capitate, truncate, punctate, filiform, peltate, infundibuliform or lobate, papillate or non-papillate, Dry or Wet type. Male flowers sometimes with pistillodium.

Ovules Placentation usually axile to basal (often with large placentae; rarely parietal). Ovules one to several hundred per carpel, usually anatropous or hemianatropous (sometimes amphitropous, campylotropous or pleurotropous), ascending to pendulous, apotropous, unitegmic or bitegmic, usually tenuinucellar (sometimes weakly crassinucellar). Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, or tetrasporous, Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent. Endosperm development cellular or nuclear (rarely [seemingly?] helobial). Endosperm haustoria micropylar and/or chalazal (sometimes absent). Embryogenesis usually solanad or caryophyllad (rarely onagrad).

Fruit A loculicidal or septicidal (rarely irregularly dehiscent) capsule, a pyxidium or berry (rarely a drupe, nut or schizocarp with five to numerous nutlike mericarps), often with persistent and accrescent calyx.

Seeds Aril absent. Exotesta and endotesta persistent. Theoidal exotestal thickenings often present. Perisperm not developed. Endosperm sometimes ruminate, usually copious, oily (rarely starchy). Embryo straight to curved (rarely annular, U-shaped or spirally twisted), usually well differentiated, with or without chlorophyll. Cotyledons usually two (rarely four). Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), O-methyl flavones, O-methyl flavonols and flavone glycosides, cyanidin, acylated anthocyanins, coumarins, ursolic acid, esters of caffeic acid, glycine betaines, nicotinic acid compounds, hygroline alkaloids (hygrines), nortropane-3α-ols, tropane-3α-ols (e.g. hyoscyamine, apoatropine, littorine), tropane-3β-ols (e.g. tigloidine), hydroxyl tropines (e.g. scopolamine, daturamine), tropane alkaloids (e.g. teloidines, tropinone, cuscohygrine, convolvine, convolamine), pyrrolizidine and pyrrolidine alkaloids (e.g. nicotinic pyrrolidine), saponins, cyanogenic compounds, withanolide steroidal lactones, and polyacetate-derived arthroquinones present. Tannins rare. Montinioside (iridoid gentiobioside) present in Montiniaceae. Iridoids and ellagic acid not found. Carbohydrates stored as oligosaccharides. Cell walls with arabinoxyloglucanes. Pyrrolidine, pyrrolizidine and tropane alkaloids synthesized from ornithine precursor.

Systematics Solanales may be sister-group to Plantaginales, although the support is fairly low.

The clade [Montiniaceae+[Sphenoclea+Hydrolea]] has the potential synapomorphies: petiole bundle transection arcuate; and presence of alkaloids (Stevens 2001 onwards). Sphenoclea and Hydrolea share the characters: swollen placentae; numerous ovules per carpel; cellular endosperm development; multicellular micropylar and chalazal haustoria; and endosperm at most scanty.

Convolvulaceae and Solanaceae form a highly supported clade, which is characterized by the following potential synapomorphies, according to Stevens (2001 onwards): presence of intraxylary phloem; leaves with conduplicate ptyxis; floral symmetry oblique; corolla aestivation usually contorted-plicate or induplicate-valvate; corolla tube formation late; carpels antepetalous; ovules numerous per carpel; integument (5–)9–20(–40) cell layers thick; presence of integumentary tapetum; persistent calyx conspicuous in fruit; testa often multiplicative; young seeds with starch; cotyledons incumbent; presence of flavonol and flavone glycosides, acylated anthocyanins, coumarins, caffeic acid esters, tropane (polyhydroxy nortropanes), pyrrolizidine and pyrrolidine alkaloids (synthesized from ornithine precursor); and absence of tannins.

Convolvulales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 247. Jan-Apr 1820 [‘Convolvulaceae’]; Cuscutales Bercht. et J. Presl, Přir. Rostlin: 247. Jan-Apr 1820 [‘Cuscutae’]; Cressaceae Raf. in Ann. Gén. Sci. Phys. Bruxelles 8: 270. 1821 [’Cressaria’]; Evolvulaceae Bercht. et J. Presl, Přir. Rostlin 2: 130, 132. 1825; Cuscutaceae (Dumort.) Dumort., Anal. Fam. Pl.: 20, 25. 1829, nom. cons.; Dichondraceae Dumort., Anal. Fam. Plant.: 20, 24. 1829, nom. cons.; Cuscutineae Link, Handbuch 1: 594. 4-11 Jul 1829 [‘Cuscutinae’]; Erycibaceae Endl. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 272, Comm. 185. 5-11 Apr 1840 [’Erycibeae’]; Convolvulopsida Brongn., Enum. Plant. Mus. Paris: xviii, 54. 12 Aug 1843 [’Convolvulineae’]; Evolvulineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1088, 1097. 1846 [‘Evolvulaceae‘]; Poranaceae J. Agardh, Theoria Syst. Plant.: 364. Apr-Sep 1858; Convolvulineae Engl., Syllabus, ed. 2: 175. Mai 1898; Humbertiaceae Pichon in Notul. Syst. (Paris) 13: 23. Jul-Sep 1947, nom. cons.

Distribution Cosmopolitan except polar areas, with their largest diversity in subtropical regions in Asia and America.

Fossils Pollen fossils (Perfotricolpites digitatus) resemble Convolvulus and other extant Convolvulaceae. Other pollen fossils (Calystegiapollis microechinatus, Perfotricolpites digitatus, Tricolpites trioblatus, Xenostegia tridentata) are known from the Eocene of Africa, southern Australia, Brazil and Europe.

Habit Usually bisexual (in Hildebrandtia dioecious), usually climbing and winding perennial or annual herbs (rarely evergreen trees, shrubs or lianas). Some species are xerophytic. Cuscuta are achlorophyllous, winding root-less stem holoparasites with haustoria and often filiform stems and branches.

Vegetative anatomy Main root in Cuscuta ephemeral, without mycorrhiza. Phellogen ab initio usually superficial (sometimes pericyclic). Medulla often septated through diaphragms. Secondary lateral growth usually anomalous (usually from concentric/successive cambia) or absent. Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or somewhat heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric, unilateral or banded. Wood sometimes fluorescent. Tyloses sometimes abundant. Intraxylary (concentric) phloem usually present (absent in Cuscuta). Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Latex cells and articulated laticifers abundant. Fibres and sclereids sometimes present. Heartwood sometimes with gum-like substances. Calciumoxalate present as acicular or prismatic crystals, styloids, druses and/or elongate crystals, crystal sand etc. in some representatives.

Trichomes Hairs unicellular or multicellular, uniseriate, furcate or multi-armed (sometimes stellate or dendritic); glandular hairs multicellular (also lepidote).

Leaves Alternate (spiral), usually simple (sometimes pinnately or palmately compound), entire or lobed, sometimes coriaceous (in Cuscuta very small, scale-like, or absent), often with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection usually arcuate (in Humbertia annular). Venation usually palmate (sometimes pinnate). Stomata usually paracytic (sometimes anomocytic, anisocytic or parallelocytic). Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts. Leaf margin serrate or entire (occasionally lobed). Extrafloral nectaries present on petiole and lamina in numerous species.

Inflorescence Terminal or axillary, usually simple or compound dichasia or cincinni (rarely dense head or spike), or flowers solitary axillary. Floral prophylls (bracteoles) sometimes large and forming involucre.

Flowers Usually actinomorphic (rarely zygomorphic; in Humbertia with oblique irregular corolla), often large. Hypogyny. Sepals (three to) five, with imbricate quincuncial aestivation, often large (sometimes unequal in size), persistent, usually free. Petals (three to) five, with induplicate-valvate or contorted-plicate (in Cuscuta imbricate [valvate?]) aestivation, connate into usually infundibuliform (sometimes campanulate or tubular) corolla, in Cuscuta with fimbriate corona especially below and behind stamens; corolla tube formation late. Nectariferous disc intrastaminal, annular or cupular (absent in Humbertia).

Androecium Stamens (three to) five, haplostemonous, antesepalous, alternipetalous, often unequal in size. Filaments free from each other, adnate to corolla tube (epipetalous). Anthers basifixed (in Humbertia) or dorsifixed, non-versatile, tetrasporangiate, usually introrse, longicidal (dehiscing by longitudinal slits; anthers rarely septate). Anther placentoid absent. Tapetum usually secretory (in Cuscuta sometimes amoeboid-periplasmodial), with multinucleate cells. Staminodia usually absent (staminodia in Cuscuta five, intrastaminal, scale-like, alternating with fertile stamens).

Pollen grains Microsporogenesis simultaneous. Pollen grains tri- to zonopolycolpate or pantoporate, shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, imperforate, perforate, reticulate, punctate, psilate, echinate or spinulate.

Gynoecium Pistil composed of two (to five) connate median antepetalous carpels (gynophore present in Humbertia). Ovary superior, usually unilocular or bilocular (rarely trilocular or quinquelocular; in Dichondra and Falkia deeply bilobate or quadrilobate, with lobes fused at base of bifid gynobasic style; in Mina with secondary septa). Style single or stylodia two (to five), usually more or less connate (in Cuscuta simple or two free; style sometimes, e.g. in Dichondreae, gynobasic). Stigma capitate, punctate, filiform or lobate (rarely peltate, truncate, infundibuliform etc.), papillate (with multicellular papillae), Dry type. Pistillodium absent.

Ovules Placentation usually subbasal to basal (in Humbertia axile). Ovules (one or) two (to four; in Humbertia numerous) per carpel, usually anatropous, ascending, apotropous, unitegmic, tenuinucellar (reduced tenuinucellar, with meiocyte semi-inferior) to weakly crassinucellar (in Cuscuta incompletely tenuinucellar). Integument usually 9–20 (in Cuscuta 15–17) cell layers thick. Placental obturator often present. Parietal tissue one or two cell layers thick. Megagametophyte usually monosporous, Polygonum type (in Cuscuta usually disporous, octacellular, Allium type). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually caryophyllad (in Cuscuta sometimes solanad).

Fruit Usually a loculicidal capsule or a pyxidium (sometimes irregularly dehiscing; rarely a nut or, e.g. in Erycibe, a berry); sepals usually persistent and often lignified in fruit.

Seeds Aril absent. Testa usually multiplicative (not in Cuscuta), with complex anatomy. Outer three cell integument layers specially differentiated. Exotesta often with papillae or hairs, somewhat thickened. Outer hypodermis consisting of small cells, little thickened. Inner hypodermis consisting one or more palisade layers. Inner hypodermal cells often elongate, thickened, or somewhat elongate. Inner layers two to eight, consisting of sclereidal cells. Endotesta? Perisperm not developed. Endosperm copious, often chartaceous, oily. Embryo usually large, straight or curved, with chlorophyll (in Cuscuta filiform, spirally coiled, poorly differentiated). Suspensor haustorium present. Cotyledons two, often plicate or coiled, bifid (almost absent in Cuscuta). Germination phanerocotylar.

DNA Deletion of 6–15 bp present in plastid gene atpB. Deletion of 150 bp present in plastid gene trnF. Plastid gene rpl2 intron lost. Plastid gene ndhF pseudogene in Cuscuta and probably in many other Convolvulaceae. Plastid inverted repeat reduced by 7 kb in Cuscuta reflexa: hence genes rpl2, rpl23, trnI, region homologous to Nicotiana ORF2280, and one rps12 intron lost. Pseudogene ψndhB in Cuscuta europaea reduced and cis-spliced intron of rps12 absent. Plastid gene ycf15 lost. Plastid ORF244 and intron in rpl2 lost in numerous Convolvulaceae, including Cuscuta. Plastid gene infA lost (Convolvulus). Mitochondrial coxI intron present (Ipomoea).

Phytochemistry Flavonols (kaempferol, quercetin), flavonol glycosides, flavones, O-methyl flavonols, O-methyl flavones, cyanidin (in Cuscuta), acylated anthocyanins, coumarins, caffeic acid esters, ornithin-derived tropane alkaloids (polyhydroxy nortropanes, e.g. convolvine, convolamine, cuscohygrine; not found in Cuscuta and Humbertia), ergoline alkaloids, pyrrolidine alkaloids (not found in Cuscuta), glycine betaines, saponins, cyanogenic compounds (derived from phenylalanine?) latex, and glucoretinous resin present. Ellagic acid and tannins not found. Some secondary metabolites are synthesized by endophytic fungi or bacteria in, e.g., Ipomoea and Turbina.

Use Ornamental plants, starch source (Ipomoea batatas), vegetables (Ipomoea aquatica), forage plants, medicinal plants.

Systematics Convolvulaceae are sister-group to Solanaceae.

Humbertia is sister to the remaining Convolvulaceae.

Humbertioideae Roberty in Candollea 14: 22. Oct 1952

1/1. Humbertia (1; H. madagascariensis; Madagascar). – Large tree. Wood with scent similar to sandalwood. Vascular bundles collateral. Secondary lateral growth normal. Wood rays uniseriate, homocellular, with exclusively procumbent cells. Usually with latex cells only in flowers or with articulated laticifers. Petiole vascular bundle transection annular. Flowers solitary, axillary, strongly obliquely zygomorphic. Sepals with five traces. Nectariferous disc absent. Filaments curved in bud, adnate to corolla base (epipetalous). Anthers basifixed. Gynophore present. Style clavate. Placentation axile. Ovules numerous per carpel. Fruit a few-seeded drupe. Endosperm copious. n = ? Ornithin-derived tropane alkaloids absent. – The phytochemistry of Humbertia is very insufficiently known.

Convolvuloideae Burnett, Outl. Bot.: 1002, 1095, 1105. Feb 1835 [‘Convolvulidae’]

c 60/1.590–1.625. Cardiochlamyeae Stafanović et D. F. Austin in S. Stefanović, D. F. Austin et R. G. Olmstead in Syst. Bot. 28: 796. 13 Nov 2003. Cordisepalum (2; C. phalanthopetalum, C. thorelii; Southeast Asia), Cardiochlamys (2; C. madagascariensis, C. velutina; Madagascar), Dinetus (6; D. decorus, D. dinetoides, D. duclouxii, D. grandiflorus, D. racemosus, D. truncatus; the Himalayas to Burma, southern China and Southeast Asia, Malesia), Porana (2; P. nutans, P. volubilis; Southeast to the Philippines, Mexico), Duperreya (3; D. commixta, D. halfordii, D. sericea; western and southern Australia), Poranopsis (3; P. discifera, P. paniculata, P. sinensis; India, Tibet and Burma to southern China and Southeast Asia, Malesia), Tridynamia (4; T. bialata, T. megalantha, T. sinensis, T. spectabilis; India and Burma to Hainan, Southeast Asia and West Malesia). – Erycibeae Hogg, Veg. Kingd.: 535. 1858. Erycibe (67; tropical Asia to Japan and tropical Australia). – Dichondreae Choisy ex G. Don, Gen. Hist. 4: 253, 302. 1837-8 Apr 1838. Dichondra (14; tropical and subtropical regions on both hemispheres), Falkia (3; F. canescens, F. oblonga, F. repens; Africa), Nephrophyllum (1; N. abyssinicum; Ethiopia), Petrogenia (1; P. repens; the Chihuahuan Desert in Mexico), Metaporana (5; M. conica, M. densiflora, M. parvifolia, M. sericosepala, M. verdcourtii; East Africa, Madagascar, Socotra), Calycobolus (c 30; tropical Africa, Madagascar, tropical America), Dipteropeltis (3; D. macrantha, D. mayumbensis, D. poranoides; tropical West and Central Africa), Rapona (1; R. tiliifolia; Madagascar). – Cresseae C. B. Clarke in J. D. Hooker, Fl. Brit. India 4: 180. Jun 1883. Hildebrandtia (13; Africa, Madagascar, the Arabian Peninsula), Seddera (c 20; tropical and subtropical regions in Africa, Madagascar, the Arabian Peninsula), Evolvulus (c 100; southern United States, Mexico, Central America, the West Indies, tropical South America, two species, E. alsinoides and E. nummularius, also in the Old World tropics), Cressa (4–5; C. aphylla, C. australis, C. cretica, C. nudicaulis, C. truxillensis; tropical to mediterranean regions on both hemispheres), Bonamia (c 45; tropical and subtropical regions on both hemispheres), Stylisma (8; southern and eastern United States), Wilsonia (3; W. backhousei, W. humilis, W. rotundifolia; southern Australia, Tasmania), Itzaea (1; I. sericea; Central America), Neuropeltis (18; tropical Africa, tropical Asia), Neuropeltopsis (1; N. alba; Borneo), Keraunea (1; K. brasiliensis; Bahia in Brazil). – Maripeae Webb et Berthel., Hist. Nat. Iles Canaries 3(2,3): 27. Apr 1844. Dicranostyles (15; tropical America), Maripa (20; tropical and subtropical America, with their highest diversity in northern South America), Lysiostyles (1; L. scandens; tropical South America). – Jacquemontieae Stefanović et D. F. Austin in S. Stefanović, D. F. Austin et R. G. Olmstead in Syst. Bot. 28: 802. 13 Nov 2003. Jacquemontia (80–100; tropical and subtropical regions on both hemispheres, with their largest diversity in the Neotropics). – Cuscuteae Dumort., Fl. Belg.: 50. 1827. Cuscuta (c 170; almost cosmopolitan). – Aniseieae Stefanović et D. F. Austin in S. Stefanović, D. F. Austin et R. G. Olmstead in Syst. Bot. 28: 796. 13 Nov 2003. Aniseia (6; A. argentina, A. cernua, A. harmandii, A. heterophylla, A. martinicensis, A. minor; southern Mexico, Central America, the West Indies, tropical South America), Odonellia (2; O. eriocephala, O. hirtiflora; tropical America), Tetralocularia (1; T. pennellii; Colombia). – Convolvuleae Dumort., Fl. Belg.: 50. 1827.Convolvulus (c 125; cosmopolitan, with their highest diversity in temperate regions). – Ipomoeeae Hallier f. in Engl. Bot. Jahrb. Syst. 16: 562, 583. 27 Jun 1893. Polymeria (10; East Malesia to northern and eastern Australia and New Caledonia); ‘Merremia’ sibirica, ‘Meremia’ poranoides, Remirema (1; R. bracteata; Thailand), Distimake (35; tropical Africa, Madagascar, the Seychelles, India and Sri Lanka to southern China, tropical Australia and islands in the Pacific), Camonea (5; C. bambusetorum, C. kingii, C. pilosa, C. umbellata, C. vitifolia; tropical Asia, one species also in tropical West and East Africa, Madagascar and tropical America from southern Mexico and the West Indies to northern Argentina), ‘Merremia’ caloxantha, Operculina (c 15; tropical regions on both hemispheres), ‘Merremia’ pro parte, Hyalocystis (2; H. popovii, H. viscosa; tropical Africa), Hewittia (1; H. malabarica; tropical Africa, Madagascar, tropical Asia to New Guinea), Xenostegia (5; X. filiformis, X. media, X. pinnata, X. sapinii, X. tridentata; tropical Africa, Madagascar, tropical Asia to tropical Australia and Solomon Islands), Decalobanthus (c 13; Southeast Asia, Malesia to islands in the Pacific; one species, D. peltatus, in tropical East Africa and Madagascar to islands in the Pacific), Merremia (10–20?; South, East and Southeast Asia, Malesia to New Guinea and northern Australia), Daustinia (1; D. montana; Brazil), ’Ipomoea’ (c 650; warm-temperate to tropical regions on both hemispheres; non-monophyletic), Stictocardia (12; tropical regions in the Old World; paraphyletic), Turbina (15; tropical and southern Africa, New Caledonia, tropical America; in Stictocardia?), Lepistemon (10; tropical regions in the Old World), ‘Ipomoea’ pro parte; unplaced Ipomoeeae: Argyreia (90–95; tropical Asia to Australia), Astripomoea (12; tropical and southern Africa), Blinkworthia (2; B. convolvuloides, B. lycioides; Burma, southern China), Lepistemonopsis (1; L. volkensii; tropical East Africa), Paralepistemon (1; P. shirensis; southern tropical Africa). – Unplaced Convolvuloideae Saccia (1; S. elegans; Colombia, Bolivia). – Distribution as for Convolvulaceae. Herbaceous (to woody), dextrorsely twining vines and lianas (rarely trees). Secondary lateral growth anomalous. Fibres and sclereids sometimes present. Hairs often unicellular, T-shaped (sometimes stellate). Leaf margin usually entire (sometimes lobed or serrate). Bract sometimes adnate to pedicel and accrescent in fruit (in Cardiochlamyeae often foliaceous); floral prophylls (bracteoles) in Convolvulus sometimes strongly enlarged. Flowers sometimes somewhat obliquely disymmetrical. Sepals with less than five traces. Pollen grains pantoporate or tri- to polycolpate. Exine in Cuscuta sometimes reticulate. Gynophore absent. Ovary in Mina with secondary septum. Placentation subbasal to basal. Ovules (one or) two (to four) per carpel, tenuinucellar to weakly crassinucellar. Integument vascularized, with unbranched bundle. Archespore in Cuscuta multicellular. Megagametophyte in at least Ipomoea strongly elongated. Fruit in Cardiochlamyeae one-seeded, utriculate, with papery pericarp. Testal cells in Erycibeae and Maripeae little thickened and elongated. Fruit usually a capsule with varying types of dehiscence. Endosperm usually with galactomannans. Ergoline alkaloids (produced by Clavicipitales fungi) sometimes present.

Phylogeny (simplified) of Convolvulaceae based on DNA sequence data (Stefanović, Krueger & Olmstead 2002; Stefanović, Austin & Olmstead 2003).

Hydroleales Bercht. et J. Presl, Přir. Rostlin: 247. Jan-Apr 1820 [‘Hydroleae’]

Habit Bisexual, annual or perennial herbs or shrubs, sometimes with axillary-sublateral spines. Helophytic.

Vegetative anatomy Mycorrhiza absent. Phellogen? Vessel elements with usually simple perforation plates (rarely scalariform); lateral pits? Vestured pits occasionally present. Imperforate tracheary xylem elements fibre tracheids, usually non-septate. Wood rays? Axial parenchyma? (diffuse parenchyma absent). Sieve tube plastids S type? Nodes? Crystals clustered in cells surrounding air-canals in primary cortex. Calcium oxalate druses in leaf veins.

Trichomes Hairs unicellular or multicellular, uniseriate; stalked glandular hairs abundant.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transaction arcuate. Venation pinnate, brochidodromous. Stomata anomocytic. Cuticular wax crystalloids? Leaf margin serrate or entire.

Flowers Actinomorphic (with diagonally inserted carpels). Hypogyny. Sepals four or five, with valvate aestivation, connate at base; first sepal adaxial, second sepal abaxial (median sepal abaxial in Hydrolea palustris). Petals four or five, with imbricate aestivation, postgenitally? connate into campanulate or tubular perigone; corolla tube formation late. Nectariferous disc intrastaminal, consisting of four or five interstaminal tubercles, or absent.

Androecium Stamens four or five, haplostemonous, antesepalous, alternipetalous. Filaments free from each other, swollen at base, adnate to corolla tube in lower part (epipetalous). Anthers basifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads?, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, without supratectal sculpturing.

Gynoecium Pistil composed of two (to four) connate, diagonally arranged/initiated carpels; carpel synascidiate (in upper part symplicate). Ovary superior, usually bilocular (sometimes trilocular or quadrilocular). Stylodia two (to four), incurved, largely free, spreading. Stigmas somewhat infundibuliform or capitate, Wet type? Pistillodium absent.

Ovules Placentation axile, with large bilobate placentae. Ovules c. 200 to c. 300 per carpel, usually anatropous, unitegmic, tenuinucellar. Integument six to eight cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells early degenerating. Endosperm development cellular. Endosperm haustoria micropylar and chalazal, multicellular. Embryogenesis solanad.

Fruit A usually septicidal (sometimes also loculicidal) capsule (rarely irregularly dehiscing).

Seeds Aril absent. Seeds ruminate, with longitudinal ridges. Exotestal cells thin-walled. Endotestal cells tanniniferous, with cuticle. Perisperm not developed. Endosperm sparse, with oil and aleuron. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Virtually unknown. Alkaloids? O-methyl flavonols? O-methyl flavones? Inulin not found.

Habit Dioecious (Montinia, Grevea) or monoecious (Kaliphora), evergreen trees or shrubs (Grevea bosseri a liana), often with peppery scent. Nodes with axillary hair tufts (in Kaliphora weakly developed).

Vegetative anatomy Phellogen? Medulla in Grevea with vascular bundles (medullary bundles). Pericyclic fibres usually absent (sometimes weakly developed). Young stem with vascular tissue usually as cylinder (sometimes as separate bundles). Cambium sometimes storied. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present (Grevea). Imperforate tracheary xylem elements fibre tracheids (Grevea) or libriform with simple or bordered pits, usually non-septate (in Grevea septate). Wood rays uniseriate or multiseriate, usually heterocellular (in Grevea homocellular). Axial parenchyma paratracheal scanty (Montinia, Grevea), or absent (Kaliphora). Sieve tube plastids S type. Nodes unilacunar or multilacunar with one (Kaliphora), three (Montinia) or four to eleven (Grevea) leaf traces (1–11:1–11). Calciumoxalate as crystal sand, acicular crystals and styloids usually present.

Trichomes Hairs unicellular or multicellular, usually uniseriate (sometimes biseriate or multiseriate); glandular hairs absent.

Leaves Alternate (spiral) or (in Grevea) more or less opposite, simple, entire, usually coriaceous (in Grevea thin), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate (sometimes rounded); petiole sometimes with additional vascular strands. Venation pinnate. Stomata usually anomocytic (in some species of Grevea anisocytic). Cuticular wax crystalloids? Leaf margin entire.

Inflorescence Terminal or axillary, cyme, or corymbose to paniculate cymose (female flowers in Montinia and Grevea solitary terminal). Floral prophylls (bracteoles) absent.

Flowers Actinomorphic. Epigyny (Montinia, Grevea) or half epigyny (Kaliphora). Sepals in male flowers three or four (or five), in female flowers four or five, with open? aestivation, connate. Petals in male flowers three or four (or five), in female flowers four or five, with imbricate (Montinia, Grevea) or valvate (Kaliphora) aestivation, caducous, free. Nectariferous disc present at least in female flowers, pulvinate to discoid, vascularized.

Androecium Stamens three or four (or five), haplostemonous, antesepalous, alternipetalous. Filaments short, thick, free from each other and from tepals. Anthers dorsifixed (Montinia, Grevea) or basifixed (Kaliphora), non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers often with staminodia (absent in Kaliphora).

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolporate (sometimes tetracolporate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate or retipilate (Kaliphora).

Gynoecium Pistil composed of usually two connate carpels (sometimes one carpel). Ovary inferior or semi-inferior, bilocular. Stylodia two, recurved (Kaliphora) or style single, short, stout, bifid, hollow, persistent in fruit (Montinia, Grevea). Stigmas decurrent (Kaliphora) or stigma large and bilobate (Montinia, Grevea), type? Male flowers sometimes with rudimentary pistillodium.

Ovules Placentation intrusively parietal to subaxile or axile. Ovules one (Kaliphora), four to six (Grevea) or up to twelve (Montinia) per carpel, anatropous (Grevea, Montinia) or campylotropous (Kaliphora), ascending, apotropous (Kaliphora) to pendulous, unitegmic, tenuinucellar? Integument ? cell layers thick. Parietal tissue approx. one cell layer thick. Megasporangial base in Montinia thin. Endothelium? Megagametophyte monosporous, Polygonum type. Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule with persistent style and stigma (Montinia, Grevea) or a drupe with two pyrenes (Kaliphora).

Seeds Aril absent. Testa winged or unwinged, thin-walled, in Grevea juicy when moistened. Exotesta lignified, with thickened periclinal cell walls (exotesta in Grevea and Kaliphora thin), usually persistent (in Grevea ephemeral). Adjacent mesotestal cell walls thickened in Montinia. Endotesta? Perisperm not developed. Endosperm copious (Grevea) or thin (Kaliphora) with hemicellulose and stratified often thick cell walls, or absent (Montinia). Embryo straight, well differentiated, chlorophyll? Cotyledons two, accumbent, foliaceous (cotyledons in Montinia with connate petioles, laterally penetrated by plumule). Radicula often oblique. Germination phanerocotylar.

Phytochemistry Route I iridoids, Route II secoiridoids, montinioside (iridoid glucoside [iridoid gentiobioside] related to valepotriates), tannins, and proanthocyanidins present. Flavonoids?

Hyoscyamaceae Vest, Anleit. Stud. Bot.: 272, 294. 1818 [’Hyoscyamoideae’]; Atropaceae Martinov, Tekhno-Bot. Slovar: 57. 3 Aug 1820; Browalliaceae Bercht. et J. Presl, Přir. Rostlin: 243. Jan-Apr 1820 [’Brovalliae’]; Daturaceae Bercht. et J. Presl, Přir. Rostlin: 243. Jan-Apr 1820 [’Datureae’]; Nicotianaceae Martinov, Tekhno-Bot. Slovar: 416. 3 Aug 1820 [’Nicotianeae’]; Nolanaceae Bercht. et J. Presl, Přir. Rostlin: 244. Jan-Apr 1820 [’Nolaneae’], nom. cons.; Cestraceae Schlechtend. in Linnaea 8: 250. Jan-Jul 1833 [’Cestrineae’]; Nolanales Lindl., Nix. Pl.: 18. 17 Sep 1833; Cestrales Schltdl. in C. F. P. von Martius, Consp. Regn. Veg.: 22. Sep-Oct 1835 [‘Cestrineae’]; Lyciaceae Raf., Autik. Bot. 1: 15. 1840 [’Lycioides’]; Solanopsida Brongn., Enum. Plant. Mus. Paris: xix, 57. 12 Aug 1843 [’Solanineae’]; Daturineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1117, 1130. 1846 [‘Datureae’]; Hyoscyamineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1117, 1134. 1846; Nolanineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1117, 1118. 1846; Solanineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1117, 1118. 1846 [‘Solaneae’]; Sclerophylacaceae Miers in London J. Bot. 7: 57, 58. Feb 1848 [‘Sclerophylaceae’]; Goetzeaceae Miers in Trans. Linn. Soc. London 27: 191. 1870 [’Goetziaceae’]; Duckeodendraceae Kuhlm. in Arq. Serv. Florest. 3: 7. 1950; Salpiglossidaceae (Benth.) Hutch., Evol. Phylog. Fl. Pl.: 631. 28 Aug 1969

Distribution Tropical, subtropical and temperate regions on both hemispheres, with their highest diversity in tropical and subtropical America.

Fossils There are only few unambiguous Solanaceae fossils known. Some macrofossils from Eocene and later layers (North America etc.) have been assigned to Solanites and they may belong in Solanaceae. However, there are several fossils that fairly well fit into other lineages of Solanales. A fossil fruit similar to Physalis were found in Eocene layers of Argentina.

Habit Usually bisexual (sometimes monoecious, andromonoecious or dioecious), evergreen trees, shrubs or lianas, perennial, biennial or annual herbs. Some species are xerophytes. Some species are succulent. Often evil-smelling (foetid).

Vegetative anatomy Roots diarch (lateral roots tetrastichous). Phellogen ab initio superficially or deeply seated. Endodermis sometimes prominent. Secondary lateral growth sometimes anomalous (from cylindrical cambium). Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes scalariform), simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids or (sometimes very long) libriform fibres (rarely tracheids) with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, lozenge-aliform, confluent, vasicentric, reticulate, or banded. Wood often fluorescent. Stem and petioles usually with intraxylary phloem (absent in, e.g., Tsoala). Sieve tube plastids S type. Nodes 1:1–3, unilacunar with one to three leaf traces. Secretory cavities present in Duckeodendron. Resins present or absent. Cystoliths present. Prismatic calciumoxalate crystals, druses and crystal sand abundant. Raphides present in Sclerophylax.

Trichomes Hairs unicellular or multicellular, usually uniseriate, often dendritic (sometimes peltate-lepidote, echinoid or stellate); glandular hairs frequent.

Leaves Usually alternate (spiral; sometimes opposite especially in inflorescence), usually simple (sometimes pinnately compound or trifoliolate), entire or pinnately lobed (in Goetzeoideae usually coriaceous), often with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles arcuate or annular. Venation pinnate. Stomata usually anomocytic or anisocytic (sometimes paracytic or diacytic; in Duckeodendron diacytic). Cuticular wax crystalloids? Domatia as hair tufts or absent. Mesophyll usually without mucilaginous idioblasts. Leaf margin and leaflet margins serrate or entire. Extrafloral nectaries present on lamina in, e.g., some species of Solanum.

Inflorescence Terminal or axillary, thyrsoid with various shape and with usually monochasial cymes, or flowers solitary axillary.

Flowers Actinomorphic to more or less (obliquely) zygomorphic (in Schizanthus resupinate and strongly zygomorphic, 3:2 type). Usually hypogyny (in Sclerophylax sometimes epigyny). Sepals (four or) five (to seven), usually with imbricate (rarely valvate) aestivation, often persistent, more or less connate (sometimes more or less inflated). Petals (four or) five (to seven), with valvate, valvate-induplicate, valvate-plicate, valvate-conduplicate, valvate-supervolute, cochleate, cochleate-conduplicate, cochleate-plicate, contorted-induplicate, contorted-conduplicate, contorted-plicate, quincuncial or reciprocative aestivation, connate into infundibuliform, campanulate or tubular perigone (odd or median petal adaxial); corolla tube formation late. Nectariferous disc intrastaminal (rarely absent). Extrafloral nectaries often present on abaxial side of corolla or on abaxial side of leaves (rarely on calyx).

Androecium Stamens (three to) five (to seven; in Schizanthus two fertile), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate to corolla tube (epipetalous), sometimes of two different lengths. Anthers often densely connivent around style, usually dorsifixed to basifixed (rarely ventrifixed), sometimes versatile, tetrasporangiate, usually introrse (sometimes extrorse or latrorse), longicidal (dehiscing by longitudinal slits or by two apical slits) or poricidal (with separate apical pores or with common apical pore due to anthers connivent or connate). Tapetum secretory, with often quadrinucleate cells. Staminodia usually absent (in Salpiglossis one, corresponding to adaxial median stamen; in Schizanthus three, corresponding to adaxial median stamen and adaxial lateral staminal pair), sometimes with four fertile stamens or with two fertile stamens and two staminodia, or sometimes with five staminodia or with two lateral or dorsal mobile fertile stamens and three staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3–5(–9)-colpate or (2–)3–5(–6)-colpor(oid)ate (sometimes inaperturate, rarely pantoporate), usually shed as monads (in Bouchetia, Nierembergia, Reyesia and Salpiglossis as tetrads; rarely as massulae), usually bicellular (sometimes tricellular) at dispersal. Exine usually tectate to semitectate (sometimes intectate), with usually columellate (occasionally granular) infratectum, perforate, reticulate, scabrate, striate, regulate, punctate-foveolate, striate-reticulate or striate-rugulate (sometimes echinate, psilate, microcechinate or granulate, rarely regulate-reticulate, regulate-striate, microgranulate, scabrate-gemmate, fossulate or smooth).

Gynoecium Pistil composed of usually two (sometimes five, rarely one or three; in Nicandra three to five; in Nolana primarily five, often secondarily transversely and/or longitudinally septate) connate antepetalous carpels, usually displaced relative to floral symmetry with oblique orientation (odd carpel often displaced to adaxial position; not so in Nicandra). Ovary usually superior (rarely inferior), usually bilocular (sometimes quinquelocular, rarely unilocular or trilocular, sometimes multilocular due to secondary septa; in Nolana usually quinquelocular, often with secondary septa; in Goetzeoideae occasionally unilocular and pseudomonomerous; in, e.g., Capsicum apically unilocular). Style single, simple, usually terminal (in Nolana sometimes gynobasic). Stigma usually bifid (sometimes capitate; in Nolana peltate), papillate or non-papillate, usually Wet type (in Solandra Dry type). Pistillodium?

Ovules Placentation axile (with large placentae). Ovules usually several to numerous (in, e.g., Duckeodendron one, in Goetzeoideae two) per carpel, usually anatropous or hemianatropous (sometimes amphitropous, anacampylotropous or hemicampylotropous), unitegmic, tenuinucellar. Integument usually 9–20 cell layers thick. Megagametophyte monosporous, Polygonum type, disporous, Allium type, or tetrasporous, Adoxa type. Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent. Endosperm development usually cellular (rarely nuclear or helobial). Endosperm haustorium chalazal (sometimes absent). Embryogenesis usually solanad (rarely onagrad).

Fruit Usually a septicidal (sometimes a loculicidal or septicidal-loculicidal) capsule, a pyxidium or many-seeded berry (rarely diclesium; in Goetzeoideae except Metternichia drupe; in Duckeodendron single-seeded drupe; in Nolana schizocarp with five to numerous nutlike mericarps), often with persistent and accrescent calyx.

Seeds Aril absent. Exotestal cells usually with thickened inner periclinal and anticlinal walls. Endotesta persistent, with lignified cell walls. Perisperm not developed. Endosperm usually copious (in Duckeodendron and Goetzeoideae sparse or absent), usually oily (rarely starchy). Embryo straight or curved (sometimes annular, rarely spirally coiled, in Duckeodendron U-shaped), usually well differentiated, without chlorophyll. Cotyledons usually two (in Goetzeoideae sometimes four). Germination phanerocotylar.

Cytology x = 10 (Schizanthoideae); x = 12, 13 (Goetzeoideae); x = 11 (Benthamielleae); x = 8, 10–12 (Cestroideae); x = 7 (Calibrachoa), 8 (Bouchetia, Hunzikeria, Nierembergia), 9 (Petunia, Fabiana, Nierembergia), 10 (Leptoglossis), 11 (Brunfelsia); x = 12 (Schwenckioideae); x = 9–23 (Solanoideae); x = 8–12 (Nicotianoideae) – Protein bodies present in nuclei. Cestreae without Arabidopsis type telomeres, and with much larger chromosomes than in remaining Solanaceae.

DNA Genome duplication supposed to have taken place c. 50 Mya (Schlueter & al. 2004). Duckeodendron and Goetzeoideae share unique deletion in trnL/F spacer. Plastid gene infA lost (present in nuclear genome).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, acylated anthocyanins, coumarins, ursolic acid, caffeic acid esters, hygroline alkaloids (hygrines), tropane-3α-ols (e.g. hyoscyamine, apoatropine, littorine), tropane-3β-ols (e.g. tigloidine), nortropane-3α-ols, hydroxyl tropines (e.g. scopolamine, daturamine), teloidines, tropinone, cuscohygrine and other tropane alkaloids, nicotinic pyrrolidine and other pyrrolidine alkaloids, steroidal alkaloids (e.g. tomatine, solanine, solanocapsine), saponins, cyanogenic compounds, polyacetate-derived arthroquinones (in Fabiana), withanolides (steroidal lactones), and nicotinic acid compounds present. Ellagic acid, tannins and proanthocyanidins not found. Aluminium accumulated in some species. Carbohydrates stored as oligosaccharides. Cell walls containing arabinoxyloglucanes and sometimes galactoxyloglucan hemicelluloses.

Use Ornamental plants, spices (Capsicum), fruits and vegetables (Solanum, Physalis, Capsicum annuum, Espadaea), starch sources (tubers of Solanum tuberosum), medicinal plants, stimulants (tobacco from Nicotiana spp., ethanol from tubers of Solanum tuberosum, narcotics from Scopolia carniolica, Atropa belladonna, Hyoscyamus, Datura, Brugmansia), insecticides (Nicotiana), timber.

Systematics Solanaceae are sister to Convolvulaceae.

Analyses of the nuclear gene SAMT (coding for salicylic acid methyltransferase; Martins & Barkman 2005) and the plastid genes ndhF and trnL/F (e.g. Olmstead & al. 2008) indicate that Schizanthus (x = 10) is sister to the remaining Solanaceae.

According to Olmstead & al. (2008), Sclerophylax is recovered in the same subclade within Atropeae as Lycium and Nolana. The morphology of Sclerophylax differs considerably from other Solanaceae and the genus has often formed a monogeneric family, Sclerophylacaceae. Crystal sand present in some species. Leaves opposite, entire, succulent, usually asymmetrical. Petiole sometimes absent. Stomata diacytic. Flowers solitary, axillary. Sometimes epigyny. Sepals five, unequal in size, connate at base into asymmetrical calyx. Petals five, with conduplicate-contorted aestivation, connate into somewhat zygomorphic infundibuliform corolla. Stamens five. Filaments epipetalous. Anthers dehiscing by longitudinal slits. Pollen grains tricolporate. Exine perforate to reticulate. Pistil composed of two oblique carpels. Ovary bilocular, with septum oblique relative to median floral plane. Style single, terminal. Stigma simple, papillate. Placentation subapical-axile. Ovules two or three per carpel, pendulous. Fruit one- to three-seeded, dry, indehiscent, with membranous pericarp, and surrounded by persistent and accrescent spiny calyx. Endosperm present. Embryo straight or curved. n = 12.

A possible topology of Solanaceae is the following: [Schizanthoideae+[[Duckeodendron+Goetzeoideae]+[[Nicotianoideae+Solanoideae]+Schwenckieae+Petunioideae+[Benthamielleae+Cestroideae]]]] (Olmstead & al. 2008).

Schizanthoideae Hunz. in Kurtziana 28: 56. Jul 2000

1/12. Schizanthus (12; Chile, Argentina). – Annual herbs. Phellogen pericyclic. Pericyclic fibres absent. Flowers strongly zygomorphic, resupinate. Abaxial pair of petals connate into keel. Stamens two abaxial-lateral fertile and three staminodial. Endosperm development ab initio nuclear. Fruit a septicidal capsule. Embryo curved. n = 10. Typical tropane alkaloids present.

[[Duckeodendron+Goetzeoideae]+[[Petunioideae+[Schwenckieae+[Nicotianoideae+Solanoideae]]]+[Benthamielleae+Cestroideae]]]

[Duckeodendron+Goetzeoideae]

Endosperm sparse or absent. Unique deletion in trnL/F spacer region.

Duckeodendron

1/1. Duckeodendron (1; D. cestroides; Amazonian Brazil). – Tree. Wood with large open radial canals. Secretory cavities present. Stomata diacytic. Carpels oblique. Ovule one per carpel. Fruit a single-seeded drupe with thick fibrous mesocarp. Endosperm sparse. Embryo U-shaped. Cotyledons small. n = ? – Duckeodendron is sister to Goetzeoideae, according to Santiago-Valentin & Olmstead (2003), or sister to the remaining Solanaceae above Goetzeoideae, according to some analysis by Olmstead (2013).

Goetzeoideae (Miers ex Airy Shaw) Thorne et Reveal in Bot. Rev. (Lancaster) 73: 132. 29 Jun 2007

6/7. Metternichia (1; M. principis; eastern Brazil), Coeloneurum (1; C. ferrugineum; Hispaniola), Henoonia (1; H. myrtifolia; Cuba), Espadaea (1; E. amoena; Cuba), Goetzea (2; G. elegans: Puerto Rico; G. ekmanii: Hispaniola), Tsoala (1; T. tubiflora; Madagascar, probably extinct). – Madagascar, Cuba, Hispaniola, Puerto Rico, eastern Brazil. Trees or shrubs. Pollen grains tricolpate. Ovules two per carpel. Exine tectate-perforate, echinate. Fruit usually a drupe (in Metternichia capsule). Endosperm sparse to almost absent. Cotyledons often large, fleshy, sometimes four. x = 12, 13. – A probable topology is, according to Santiago-Valentin & Olmstead (2003): [Metternichia+[Coeloneurum+[Henoonia+[Espadaea+Goetzea]]]]. The position of Tsoala tubiflora is unknown (the species has not been observed since 1959 and may be extinct).

[[Petunioideae+[Schwenckieae+[Nicotianoideae+Solanoideae]]]+[Benthamielleae+Cestroideae]]

[Petunioideae+[Schwenckieae+[Nicotianoideae+Solanoideae]]]

Petunioideae (Horan.) Thorne et Reveal in Bot. Rev. (Lancaster) 73: 132. 29 Jun 2007

8/135–140. Petunia (40; tropical and subtropical South America), Fabiana (c 15; warm-temperate South America, especially in the Andes); Brunfelsia (45–50; tropical America); Leptoglossis (6; L. acutiloba, L. albiflora, L. darcyana, L. ferreyraei, L. lomana, L. schwenckioides; coastal Peru and Chile, Argentina), Nierembergia (c 20; Mexico, South America south to Chile and Argentina), Bouchetia (4; B. anomala, B. arniatera, B. erecta, B. procumbens; southern United States to Brazil), Hunzikeria (3; H. coulteri, H. steyermarkiana, H. texana; southwestern United States, Mexico, Venezuela), Plowmania (1; P. nyctaginoides; Mexico, Guatemala). – Southern United States to southern South America. Phellogen usually superficial (rarely deeply seated). Bordered pits present. Pericyclic fibres usually present. Druses usually absent. Flowers sometimes zygomorphic. Stamens usually four (rarely five), usually didynamous. Embryo straight or slightly curved. x = 7–9, 11. Tropane alkaloids (calystegines) sometimes present. – The position of Petunioideae is unresolved and sometimes they are placed as sister to [Nicotianoideae+ Solanoideae] (e.g. Olmstead & al. 1999).

[Schwenckieae+[Nicotianoideae+Solanoideae]]

Schwenckieae Hunz. in Kurtziana 10: 42. 25 Apr 1977

3/25–30. Heteranthia (1; H. decipiens; Brazil), Melananthus (7; M. cubensis, M. dipyrenoides, M. fasciculatus, M. guatemalensis, M. luetzelburgii, M. multiflorus, M. ulei; Central America, the West Indies, tropical South America), Schwenckia (c 20; tropical America, one species, S. americana, also in tropical West Africa). – Tropical America, tropical West Africa. Annual herbs. Pericyclic fibres present. Flowers zygomorphic. Each corolla lobe in Melananthus and Schwenckia trilobate. Stamens two longer and two shorter, or two fertile and two or three staminodia. Embryo straight, short. x = 12 (Schwenckia).

[Nicotianoideae+Solanoideae]

Stigma Wet type. Cotyledons sometimes with margins against radicula. x = 12. Tropane alkaloids and nicotine (a pyridine alkaloid) sometimes present.

Nicotianoideae Miers in London J. Bot. 7: 58. 1848 [‘Nicotianeae’]

8/c 110. Nicotiana (c 75; southwestern United States, northern Mexico, southern South America east of the Andes, Australia, islands in the South Pacific, one species, N. africana, in Namibia), Symonanthus (2; S. aromaticus, S. bancroftii; southwestern Western Australia), Anthocercis (c 15; southwestern Western Australia, southern South Australia), Grammosolen (2; G. dixonii, G. truncatus; southeastern Western Australia, southern South Australia),’Cyphanthera’ (9; southern Australia; polyphyletic), Anthotroche (3; A. myoporoides, A. pannosa, A. walcottii; southwestern Western Australia), Crenidium (1; C. spinescens; continental southwestern Western Australia), 'Duboisia' (4; D. arenitensis, D. hopwoodii, D. leichhardtii, D. myoporoides; eastern Australia, New Caledonia; probably non-monophyletic). – Namibia, Australia, New Caledonia, America. Phellogen superficial. Pericyclic fibres often present. Stamens four (staminodium sometimes present) or five, sometimes didynamous. Embryo usually straight (sometimes curved). Radicula short. x = 8–12. Tropane alkaloids (calystegines) sometimes present. Nicotine often present.

Solanoideae Burnett, Outlines Bot.: 985, 1095, 1106. Feb 1835 [‘Solanidae’]

57/2.170–2.270. Atropeae Kitt. in A. Richard, Nouv. Elém. Bot., ed. 3, Germ. transl.: 796. 1840. Latua (1; L. pubiflora; southern Chile); Jaborosa (23; Chile, southern Argentina); Sclerophylax (14; Uruguay, Paraguay, Argentina), Nolana (89; Peru to Chile and western Argentina, the Galápagos Islands), Lycium (90–95; warm-temperate and subtropical regions on both hemispheres), Atrichodendron (1; A. tonkinense; Vietnam; in Lycium?); Atropa (5; A. acuminata, A. baetica, A. belladonna, A. komarovii, A. pallidiflora; northern Morocco, Europe to the Caucasus, northern Iran, Central Asia and the Himalayas, Mongolia), Jaborosa (22; southern Peru, Chile, southern Argentina), Anisodus (5; A. acutangulus, A. carniolicoides, A. luridus, A. mariae, A. tanguticus; the Himalayas in Nepal, India and Bhutan to western China), Atropanthe (1; A. sinensis; China), Hyoscyamus (c 17; Europe, the Mediterranean, Madeira, the Canary Islands, North Africa to Somalia, southwestern and Central Asia to China), Przewalskia (1; P. tangutica; western China), Scopolia (2; S. carniolica: the Alps, the Carpathians, the Caucasus; S. japonica: the Korean Peninsula, Japan), Physochlaina (6; P. capitata, P. infundibularis, P. macrocalyx, P. macrophylla, P. physaloides, P. praealta; the Caucasus to India and China). – Exodeconus clade Exodeconus (6; E. flavus, E. integrifolius, E. maritimus, E. miersii, E. prostratus, E. pusillus; South America, the Galápagos Islands). – Nicandreae Lowe, Man. Fl. Madeira 2: 98. Jan-Apr 1872. Nicandra (1; N. physalodes; Peru to northern Argentina). – Solandreae Miers in Ann. Mag. Nat. Hist., ser. 2, 3: 166. Mar 1849. Solandra (10; Mexico, Central America, the West Indies, tropical South America south to Bolivia and southeastern Brazil), Schultesianthus (7; S. coriaceus, S. crosbyanus, S. dudleyi, S. leucanthus, S. megalandrus, S. uniflorus, S. venosus; southern Mexico, Central America, Venezuela and Colombia to Peru), ‘Markea’ (20; Central America, northern South America; polyphyletic), Trianaea (5; T. brevipes, T. naeka, T. nobilis, T. speciosa, T. spectabilis; Venezuela, Colombia, Ecuador, Peru), ‘Merinthopodium’ (3; M. neuranthum, M. pendulum, M. vogelii; southern Mexico, Central America to Venezuela and Colombia; non-monophyletic), Poortmannia (1; P. speciosa; Colombia, Ecuador), Ectozoma (2; E. pavonii, E. ulei; southwestern Colombia, Ecuador, northern Peru), Dyssochroma (4; D. albidoflava, D. eximia, D. longipes, D. viridiflora; southeastern Brazil), Juanulloa (8–11; Mexico, Central America, the West Indies, tropical South America to Bolivia), Hawkesiophyton (4; H. klugii, H. ochraceum, H. panamense, H. ulei; Panamá to Colombia and Amazonia in Brazil). – Mandragoreae Rchb., Handb. Nat. Pfl.-Syst.: 201. 1-7 Oct 1837. Mandragora (4; M. autumnalis, M. caulescens, M. officinarum, M. turcomanica; the Mediterranean, Central Asia, the Himalayas). – Datureae Dumort., Anal. Fam. Plant.: 24. 1829. Trompettia (1; T. cardenasiana; Bolivia), Datura (12; southern United States, Mexico), Brugmansia (6; B. arborea, B. longifolia, B. pittieri, B. sanguinea, B. suaveolens, B. versicolor; the Andes in Venezuela to Bolivia). – Salpichroa clade Salpichroa (22; southern United States and Mexico to southern Andes). – Physalideae Miers in Ann. Mag. Nat. Hist., ser. 2, 3: 179. Mar 1849 [‘Physaleae’]. Cuatresia clade: Cuatresia (13; tropical America); Withaniinae Bohs et Olmstead in R. G. Olmstead et al. in Taxon 57: 1171. 26 Nov 2008: Deprea (c 50; Central America, tropical South America, with their highest diversity in northern and central Andes), Aureliana (15; eastern Brazil, central South America), Withania (c 20; warm-temperate to tropical regions in the Old World, one species, W. begoniifolia, on St. Helena), Nothocestrum (4; N. breviflorum, N. latifolium, N. longifolium, N. peltatum; the Hawaiian Islands), Tubocapsicum (1; T. anomalum; southern and eastern Asia), Discopodium (1; D. penninervium; tropical African mountains); Iochromeae Miers in Ann. Mag. Nat. Hist., ser. 2, 3: 178. Mar 1849: Dunalia (8; the Andes), ’Iochroma’ (c 25; western South America south to north-western Argentina; non-monophyletic), Saracha (3; S. andina, S. punctata, S. quitensis; the Andes from Venezuela to Bolivia), Acnistus (1; A. arborescens; tropical America; in Iochroma?), Eriolarynx (3; E. fasciculata, E. iochromoides, E. lorentzii; Bolivia, Argentina), Vassobia (3; V. breviflora, V. dichotoma, V. fasciculata; central South America); Physalis clade: Witheringia (c 14; southeastern Mexico, Central America to Bolivia, the West Indies), Leucophysalis (5; L. grandiflora, L. kimurai, L. nana, L. savatieri, L. viscosa; North America, Mexico, Central America, Colombia, Venezuela), ’Physalis’ (c 90; nearly cosmopolitan, with their highest diversity in Mexico to northern South America; paraphyletic), Schraderanthus (1; S. viscosus; Mexico, Guatemala, Venezuela; in Physalis?), Capsicophysalis (1; C. potosina, southern Mexico to Guatemala and Honduras; in Physalis?), Calliphysalis (1; C. carpenteri; southeastern United States), Darcyanthus (1; D. spruceanus; Peru, Bolivia), Oryctes (1; O. nevadensis; southwestern United States), Quincula (1; Q. lobata; southwestern United States to northern Mexico), Chamaesaracha (13; North America, northern South America). – Capsiceae Dumort., Fl. Belg.: 38. 1827. Lycianthes (c 140; East Asia, tropical America; paraphyletic?), Capsicum (c 32; southern United States to central Argentina; incl. Lycianthes?). – Solaneae Dumort., Anal. Fam. Plant.: 24. 1829. Jaltomata (c 60; tropical and subtropical America), Solanum (1.300–1.400; nearly cosmopolitan, with their largest diversity in Australia and South America). – Subcosmopolitan. Herbs or shrubs (sometimes small trees). Vestured pits absent. Crystal sand sometimes present. Stamens (four or) five, sometimes didynamous. Filament base often enlarged, lobed etc. Pistil sometimes composed of three carpels or carpels sometimes divided into single-seeded units (e.g. Nolana with primarily five antepetalous carpels). Style sometimes gynobasic. Endosperm development cellular. Fruit usually a berry (rarely drupe or schizocarp, in Hyoscyameae etc. pyxidium). Calyx sometimes strongly accrescent in fruit. Seeds flattened. Exotestal cells sometimes anticlinally elongate. Embryo curved, often coiled. x = 9–23. Tropane alkaloids (calystegines), nicotine and steroidal alkaloids sometimes present. Glycine betaines present in Lycium.

[Benthamielleae+Cestroideae]

Benthamielleae Hunz. in Kurtziana 28: 61. Jul 2000

3/15. Benthamiella (13; southern Patagonia in Chile and Argentina), Combera (2; C. minima, C. paradoxa; the Andes in temperate Chile and Argentina), Pantacantha (1; P. ameghinoi; Patagonia in central Argentina). – Southern South America. Exine with special irregular ornamentation. x = 11. – Benthamielleae are sister to Cestroideae, according to Olmstead & al. (2008).

Cestroideae Burnett, Outlines Bot.: 985, 1095, 1106. Feb 1835 [‘Cestridae’]

6/160–210. Salpiglossideae Benth. in Edwards’s Bot. Reg. 21: ad t. 1770. 1 Feb 1835. Salpiglossis (1; S. sinuata; southern Andes), Reyesia (4; R. cactorum, R. chilensis, R. juniperoides, R. parviflora; northern Chile)? x = 11. – Browallieae Hunz. in Lorentzia 8: 6. 31 Mai 1995. Browallia (5; B americana, B. demissa, B. eludens, B. speciosa, B. viscosa; southeastern Arizona to central South America, the West Indies). x = 10–12. – Cestreae Dumort., Anal. Fam. Plant.: 24. 1829 [‘Cestrineae’]. Cestrum (150–200; tropical and subtropical America), Vestia (1; V. foetida; Chile), Protoschwenckia (1; P. mandonii; Brazil, the Andes in Bolivia). x = 8. – Southern United States to southern Andes. Phellogen superficial or deeply seated. Bordered pits present. Pericyclic fibres present. Stamens four or five, often didynamous. Staminodia often present. Pollen grains in Salpiglossis shed as tetrads. Fruit sometimes fleshy. Exotesta in Cestrum with all cell walls somewhat thickened. x = 8, 10–12. Chromosomes larger than other Solanaceae. Arabidopsis type telomeres absent at least in some Cestroideae. Steroid alkaloids present in Cestrum. – A probable topology of Cestroideae is [Salpiglossideae+[Browallieae+Cestreae]]. A plausible topology of Cestreae is [Protoschwenckia+[Vestia+[Cestrum+Sessea]]] (Olmstead & al. 2008).

Cladogram (simplified) of Solanaceae based on DNA sequence data (Olmstead 2013).

Cladogram (simplified) of Solanaceae based on DNA sequence data (Olmstead & al. 2008).

Cladogram of Petunioideae based on DNA sequence data (Olmstead & al. 2008).

Cladogram of Nicotianoideae based on DNA sequence data (Olmstead & al. 2008).

Cladogram of Solanoideae based on DNA sequence data (Olmstead & al. 2008).

Pongatiaceae Endl. ex Meisn., Plant. Vas. Gen.: Tab. Diagn.: 242, Comm.: 151. 18-24 Aug 1839 [’Pongatieae’], nom. illeg.; Sphenocleales Doweld, Tent. Syst. Plant. Vasc.: xlvii. 23 Dec 2001

Habit Bisexual, annual herbs. Relatively succulent. Helophytic. Stem hollow.

Vegetative anatomy Phellogen mid-cortical? Large vertical cortical air canals (aerenchyma) present in stem. Vessel elements with simple perforation plates; lateral pits scalariform or alternate, simple pits? Imperforate tracheary xylem elements tracheids? with simple pits. Wood rays absent? Axial parenchyma paratracheal scanty. Pericyclic sclerenchyma present. Sieve tube plastids S type? Nodes? Laticifers and latex absent. Calciumoxalate druses present.

Leaves: Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata tetracytic. Cuticular wax crystalloids? Mesophyll with calciumoxalate druses. Leaf margin entire.

Flowers Actinomorphic, small. Epigyny or partial epigyny (upper parts of ovary free). Sepals five, with imbricate aestivation, persistent, connate. Petals five, with imbricate quincuncial aestivation, connate in lower part, inflexed, caducous; corolla tube formation early. Nectary absent. Disc absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments short, free from each other, adnate to base of corolla tube (epipetalous). Anthers dorsifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate to tricolporoidate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, imperforate to finely perforate, smooth or almost so.

Gynoecium Pistil composed of two connate carpels. Ovary inferior or semi-inferior, bilocular. Style single, simple, short, or absent. Stigma capitate, Wet type. Pistillodium absent.

Ovules Placentation axile, with large placentae. Ovules numerous per carpel, anatropous, pendulous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Hypostase absent. Megagametophyte monosporous, Polygonum type. Synergids elongated. Antipodal cells degenerating, ephemeral. Endosperm development cellular. Endosperm haustoria chalazal and micropylar, multicellular. Embryogenesis onagrad.

Seeds Aril absent. Exotesta with thickened inner cell walls; Exotestal cells polygonal, with thickened inner walls and spine-shaped radial processes. Endotesta developing into endothelium? Perisperm not developed. Endosperm very sparse, with thick cell walls, or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Very insufficiently known. Cyclic thiosulphinates (zeylanoxides) present. O-methylflavonols? O-methylflavones? Alkaloids? Iridoids not found. Secoiridoids sometimes present. Fructose with isokestose linkages present (inulin has been reported).

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SOLANALES Juss. ex Bercht. et J. Presl

CONVOLVULACEAE Juss.

HYDROLEACEAE R. Br. ex Edwards

MONTINIACEAE Nakai

SOLANACEAE Juss.

SPHENOCLEACEAE (Lindl.) Baskerville