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Shrubs or perennial to annual herbs, rarely trees, rosette-forming or ephemerals, sometimes with tuberous or gemmiferous roots, or with tubers or stolons; stems occasionally with heteroblastic growth or with cauline spines; plants glabrous, frequently viscose or slightly or densely wooly-tomentose, with a diverse array of non-glandular and glandular trichomes. Leaves alternate, often in pairs, sometimes becoming opposite in the inflorescence, usually simple, entire, infrequently pinnatifid to deeply dissected or compound, exstipulate. Flowers perfect, rarely functionally unisexual in dioecious or andromonoecious plants, sessile to mostly pedicellate, fragrant or not, solitary or more commonly in axillary, extra-axillary, or terminal multi-flowered lax panicles, cymes or fascicles. Perianth (4)5(6)-merous; calyx actinomorphic, rarely zygomorphic, undivided or slightly to much divided, usually persistent and variously accrescent; corolla actinomorphic or zygomorphic, rotate, stellate, tubular, infundibuliform or hypocrateriform, exceptionally papilionaceous, tube glabrous or hairy inside, lobes longer or shorter than the tube, similar or dissimilar in size, aestivation valvate, valvate-induplicate, valvate-plicate, valvate-conduplicate, valvate-supervolute, cochlear, cochlear-conduplicate, cochlear-plicate, contorted-induplicate, contorted-conduplicate, contorted-plicate, quincuncial, or reciprocative; androecium included or exserted, 5-merous, 6-merous (Goetzea), 4-merous (then stamens equal in Nothocestrum, otherwise didynamous: 4 fertile or 2 fertile stamens and 2 staminodes; sometimes also with a fifth staminode) or with 2 fertile mobile stamens in lateral or dorsal position and 3 staminodes with vestigial or sometimes without anthers, filaments straight or declinate, inserted at different heights on the corolla tube, distinct or connate in a basal ring fused to the corolla, filament base appendages adnate to petal tube (“stapet”) absent, inconspicuous or conspicuous (with or generally without basal auricles), anthers dorsifixed, basifixed, dorsi-basifixed, or ventrifixed, extrorse, introrse or latrorse, frequently connivent (in Solanum sect. Lycopersicon with sterile apices and joined in a column), thecae generally non confluent apically, equal or unequal, dehiscence longitudinal or by terminal pores or slits, connective inconspicuous, wide, forming a dorsal layer of uniform and slight thickness, or thick with an emerging hump; gynoecium with oblique orientation (except Nicandra), usually bicarpellate, 3–5-carpellate in Jaborosa, Nicandra, Trianaea, and up to 30-carpellate in Nolana, ovary generally bilocular, sometimes 4-locular due to false septa, superior or sometimes partly inferior, glabrous or with trichomes or prickles; style heteromorphic or homomorphic, straight or declinate, terminal or subterminal, stigma variously shaped, usually wet and papillate (papillae rarely lacking); nectary absent, cryptic, or evident, then annular, invaginated or with 2 prominent lobes. Fruit generally a many-seeded berry or a septifragal, septicidal or septicidal-loculicidal capsule, rarely a pyxidium, drupe, diclesium or schizocarp. Seeds 1 to ca. 5000, compressed, then discoid, lenticular, reniform, irregularly ovoid or not compressed, then generally angular, cuboidal, ovoid, prismatic, polyhedric, subspherical, reniform; embryo straight to coiled, U-shaped only in Duckeodendron, cotyledons incumbent or oblique, less frequently accumbent, usually equal, slightly longer or shorter (up to 12 times shorter in the Australian endemic genera) and as wide or rarely broader than rest of embryo; endosperm rarely absent, generally copious, storing mainly oil and starch, with cellular type of endosperm formation (nuclear only in Schizanthus).

Cosmopolitan family of 96 genera and ca. 2400 species, much more diversified in South America than elsewhere.

Vegetative Morphology. Annual, biennial or more commonly perennial plants, succulents in Sclerophylax and Nolana; shrubs (0.5–6 m tall but occasionally stunted, rarely exceeding 20 cm high), sometimes lianas (up to 30 m high) or small trees (5–10 m tall, rarely up to 30 m), rarely greatly reduced (annuals, hemicryptophytes or chamaephytes); occasionally with fleshy, thick taproots, or with gemmiferous roots, or root tuberosities, or tubers, stolons or rhizomes. Branching sympodial, sometimes little branched, zigzag, or strongly dichasial; stems usually erect, or reclinate to prostrate, sometimes climbing, occasionally with heteroblastic growth, or sometimes with spines or prickles, or leafless, internodes long or short, hollow or usually solid. Leaves entire, infrequently toothed or cleft (pinnatifid, pinnatipartite or pinnatisect), generally simple, or occasionally compound (usually imparipinnate or trifoliolate), always exstipulate, occasionally thick, coriaceous, occasionally highly succulent (especially Nolana and Sclerophylax), alternate or frequently anisogeminate (the members of a pair both on the same side of the stem), often markedly unequal, sometimes in whorls of three to six, fasciculate or not, sessile or petiolate; basal leaves sometimes in a rosette. In some taxa (especially in Solanum), hair tuft type domatia on the leaves (Brouwer and Clifford 1990).

Various systems of vegetative reproduction are known. For example, in Nierembergia aristata, aerial stolons arise at the basal tuber-like nodes of the erect plants; the species of Solanum sect. Petota (S. tuberosum, S. chacoense, etc.) have subterraneous stolons arising from the underground stem and form a terminal tuber each. In other cases, root buds (Fig. 56C) are responsible for the origin of subterranean organs; in this way the horizontal rhizomes of Nierembergia rigida (Cosa 1989) or the vertical rhizomes of some Jaborosa spp. are formed (Barboza and Hunziker 1987). In Bouchetia anomala, Leptoglossis linifolia (Fig. 56B) and Solanum elaeagnifolium, there are lateral root buds which show two growth phases: a horizontal followed by a vertical phase, the buds arising at the point of curvature of the roots (Cosa 1989). Finally, the vertical root ends in a prominent tuberosity in S. juvenale and S. hieronymi (Cosa et al. 1998, 2000).

Fig. 56.
figure 1

Solanaceae. Vegetative anatomy. A–C Leptoglossis linifolia. A Diarch primary root. B Vegetative propagation. C Secondary root with root buds. D–F Brunfelsia spp.; G–I Browallia spp.; J–L Nicotiana spp.; M–O Schizanthus spp.; P–R Reyesia spp. D, G, J, M, P Primary stem diagrams (observe the different position and kind of tissues). E, H, K, N, Q Secondary stem diagrams (note the different position of the periderm). F, I, L, O, R Foliar structures. F, I Dorsiventral, L, O Isolateral, R Homogeneous. (Orig.)

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Vegetative Anatomy. All members of the family are distinctive in having internal phloem and 1-traced, unilacunar nodes. The foliar epidermis (trichomes and stomata) and the internal structure of leaves and stems have relevant features for the delimitation of the different taxa. The primary root is usually diarch (Fig. 56A), such as in Nicotiana tabacum, Solanum juvenale, S. elaeagnifolium, S. tuberosum, S. sisymbriifolium and Bouchetia anomala (Dottori et al. 2000; Hadid et al. 2007), but sometimes triarch types are present such as in Solanum palinacanthum (Pericola et al. 2004), or even tetrarch, pentarch and hexarch in other Solanum species (Nurit-Silva et al. 2007, 2011). The first phellogen originates from pericyclic layers (Combera, Benthamiella, Nicotiana), or immediately below the epidermis (Solanum sisymbriifolium). In some species (e.g. Solanum hieronymi) the roots develop into small fleshy storage organs.

In the primary stem, the epidermis consists of a single layer of cells, while the cortex shows distinct features according to the position and kinds of tissues (Metcalfe and Chalk 1950; Cosa 1991, 1993, 1994; Liscovsky et al. 2002; Liscovsky and Cosa 2005; Liscovsky 2007, Figueroa et al. 2008); in some genera, e.g. Petunia, Benthamiella and Combera, the cortex consists exclusively of chlorenchyma; the chlorenchymatic subepidermal layers are arranged palisade-like in Leptoglossis, Salpiglossis and Reyesia (Fig. 56P). In other genera, the cortex consists of chlorenchyma and collenchyma: sometimes, as in Lycium, Jaborosa, Sclerophylax and Brunfelsia (Fig. 56D), the collenchyma is superficial, while in Browallia (Fig. 56G), Nicotiana (Fig. 56J), Schizanthus (Fig. 56M), Cestrum, Sessea, some Solanum and the majority of Fabiana spp. it is located on the innermost layers of the cortex. Finally, in genera such as Datura and Brugmansia the cortex consists of chlorenchyma, collenchyma and parenchyma, with the collenchyma located beneath the superficial chlorenchyma. In some species of Melananthus, Protoschwenckia and Schizanthus, the innermost cortical layer usually contains many large starch grains (starch sheath); in other cases (e.g. Nierembergia, Combera, Schwenckia, Streptosolen), these cells have Casparian strips. Two patterns of arrangement of the vascular tissues are characteristic: the most common is an amphiphloic siphonostele, but in some Solanum spp. and in Sclerophylax a eustele with bicollateral bundles is present. In the secondary stem, the origin of the periderm and the presence and distribution of fibres and crystals (druses, solitary crystals and crystal sands) are taxonomically important. The periderm may have a superficial or a deep origin; in the former case, it arises from the epidermal layer (e.g. Brunfelsia, Fig. 56E, Schwenckia, Nierembergia and Leptoglossis), or from subepidermal layers (Browallia, Fig. 56H, Cestrum, Latua); in the latter case, the periderm originates at the outer limit of the external phloem, such as in Salpiglossis and Reyesia (Fig. 56Q) and Schizanthus (Fig. 56N). The pericycle consists commonly of isolated groups of fibres (Fig. 56E, H, Q) or, in some cases, a continuous ring of fibres is present, generally with cellulose or with slightly or well-lignified cell walls. Sclerenchyma also occurs in the cortex or in the external and/or internal phloem and/or in the pith, such as in Brunfelsia (Fig. 56E), Cestrum, Sessea, Browallia (Fig. 56H), Streptosolen, Nicotiana (Fig. 56K); sclereids also appear in the epidermis (Cestrum and Sessea). In Schizanthus (Fig. 56N) and Combera, there are no fibres at all. Datura shows atypical growth in the vascular system by forming an extra-parenchyma as a water storage tissue (Liscovsky et al. 2001).

Wood structure is diverse in Solanaceae: ring porous (e.g. Fabiana and Lycium) to diffuse porous (Capsicum rhomboideum, Nothocestrum latifolium), and the vessels have simple perforation plates, or rarely vestiges of bars (Carlquist 1988, 1992); wood of Duckeodendron is clearly mesomorphic, that of Espadaea is intermediate and that of Henoonia is relatively xeromorphic, based on vessel element dimensions, vessel density, and presence of vasicentric tracheids (Carlquist 1988). Carlquist (1987) reviewed the wood anatomy in Nolana and suggested it was indicative of paedomorphosis. Both in primary and secondary stems druses (Fig. 56H, N), solitary crystals or crystal sand (Fig. 56K) are present in the cortex and pith and sometimes also in the mesophyll. Stem and leaves in Salpiglossis and Reyesia contain druses and solitary crystals but Browallia, Streptosolen, Brunfelsia and Schizanthus have only druses; the stems of Benthamiella, Combera, Pantacantha, Nierembergia and Bouchetia completely lack any kind of crystals.

The anatomy of the petiole is uniform. The vascular system consists of a central arc-shaped bicollateral bundle accompanied by 1–4 smaller bundles on each side (e.g. Browallia, Salpiglossis, Reyesia, Solanum). Dorsiventral (Fig. 56F, I), isolateral (Fig. 56L, O) and homogenous (Fig. 56R) leaves are present in the family, but the dorsiventral type is the most common. Some genera show only one type of leaf structure, e.g. Jaborosa, Schultesianthus, Brunfelsia and Schwenckia, while others (such as Lycium, Calibrachoa, Reyesia, Solanum and Nierembergia) have two and even all three structural types (Cosa 1991, 1993; Reis et al. 2002; Araújo et al. 2010). The vascular system consists of a main bicollateral bundle from which smaller veins branch. In some species, the phloem is surrounded by collenchymatous cells (Nicotiana, Browallia, Streptosolen) or fibre groups (Pantacantha, Benthamiella, Brunfelsia, some Solanum); in Duckeodendron, the main vein is surrounded by a sclerenchymatous sheath. Stomata occur either on both sides of the leaf—e.g. some species of Calibrachoa, Datura, Schwenckia, Protoschwenckia and Solanum—or only on the lower side, as in some Solanum spp. (Benítez de Rojas 2007; Nurit-Silva and Agra 2011), Cestrum, Atropa, Brunfelsia, Latua, Streptosolen and Duckeodendron.

The presence of different types of stomata on one leaf is frequent in Solanaceae. The most common types are anomocytic and anisocytic stomata; however, Brunfelsia has mostly paracytic stomata and, in Dyssochroma, Juanulloa, Withania, Physalis, Schwenckia and Melananthus, the diacytic type is predominant (Patel and Inamdar 1971; Bessis and Guyot 1979). Apparently, Fabiana is the only genus where the stomata are placed on projections of the epidermis. Raphides are present in the mesophyll of some Sclerophylax (Di Fulvio 1961).

Trichomes are present on nearly all parts of the plants and are of taxonomic importance (Seithe 1962). The array, density, and variety of trichomes (i.e. morphology, size, and ornamentation of the cell walls) produce a great diversity of indumentum types (Roe 1971). Non-glandular, simple to variously dendritic, uniseriate, multicellular trichomes as well as glandular trichomes (head and stalk uni- or multicellular) are the most widespread in the family. Simple short trichomes result in a puberulose or strigose appearance, as in some Jaltomata, Larnax, etc., while longer trichomes which are the most common result in a villose (Hyoscyamus, some Aureliana, Nolana and Solanum, etc.), tomentose (some Australian genera, Solanum, etc.) or viscid indumentum (e.g. Salpiglossis, Reyesia, Nicotiana, Nolana, Petunia, Fabiana, Calibrachoa, Exodeconus, some Athenaea, and Solanum spp.). Dendritic trichomes vary from few-branched as in Leptoglossis, Withania, Brunfelsia, etc. to many-branched as in Solanum (Mentz and de Oliveira 2004), Sessea (Benítez de Rojas and Jáuregui 2005), Cestrum, Nolana, Juanulloa, Anthotroche and Symonanthus. In general, the surface of the trichomes is smooth or slightly to strongly verrucate, especially within the non-glandular types; rarely striate non-glandular trichomes (Vestia, Leptoglossis, some Solanum spp.) or trichomes with verrucate stalk (Schwenckia, Protoschwenckia, Heteranthia, and some Jaborosa, Capsicum, and Lycianthes spp.) appear within the glandular type. The stellate, echinoid and lepidote trichomes of Solanum subgen. Brevantherum and subgen. Leptostemonum are very variable in structure (Mentz et al. 2000). The stellate trichomes are sessile or stalked (stalk uniseriate or multiseriate, multicellular, emergent or intrusive), the lateral rays (more than 4) are in one plane or multiangulate (arms radiating outwards in all directions) while the short or long central ray is glandular or not. The most conspicuous trichomes are: a) unicellular non-glandular trichomes, b) glandular trichomes with long multiseriate stalk and multicellular head (shaggy hairs) present only in Schizanthus, c) glandular trichomes with a multiseriate, multicelled head (more than 20 cells) and a long, multicellular, uniseriate, smooth stalk characteristic of Salpiglossis and Reyesia, d) whorled branched trichomes mixed with other dendritic trichomes in the majority of the Australian genera, e) glandular scale-like peltate trichomes, sunken in the mesophyll of Schultesianthus, f) vesicular, shiny, strongly verrucate trichomes only in Quincula, g) viscid glandular trichomes (head uni- or multicelled) in some Exodeconus, Nicotiana, Nolana, Petunia, Calibrachoa, and oil-secreting trichomes on the inner surface of the corolla in Nierembergia and Atropa, h) bayonet trichomes (two-celled trichomes, their basal cell very large and thick-walled, the apical cell small and thin-walled) of Solanum sect. Basarthrum, i) non-glandular and glandular trichomes, present in Oryctes, Lycium and Lycianthes, j) branched-glandular trichomes in Sclerophylax, Physochlaina orientalis, Eriolarynx lorentzii and some Physalis, or stellate-glandular ones with many short glandular branches (e.g. Solanum velleum and S. robustum), and k) dendritic-echinoid (S. axillifolium) or paleaceous (S. castaneum) trichomes. The presence of prickles is peculiar to Solanum subgen. Leptostemonum; they may be straight, curved or uncinate, acicular, mammiform or conic, compressed or not, patent to retrorse, glabrous or with non-glandular or glandular trichomes, rarely with stellate trichomes at their base and sometimes with a tuft of unicellular rays at their apex. In Saracha, the young stems are covered with brownish, dendriform, multicellular emergences.

Inflorescence Structure. Danert (1958, 1967) and Troll (1969) intensively studied the inflorescence structure in the family; before them, an account had been provided by Eichler (1875, 1878). The flowering shoots are basically of the monotelic (closed) thyrse type. They are often strongly modified by serial (basis to top) differentiation in the degree of branching, by recaulescence (shift of the subtending leaf along their axillary shoots), and concaulescence (fusion of axillary shoot to the primary shoot). These basic types of branching are consistent in the family. However, the diverse patterns found are still difficult to connect with each other. Each flowering shoot ending in a flower commonly continues growth by sympodial branching resulting in cymose systems. These represent the repetition of modules (sympodial units), each one consisting of a flower and its underlying node and internodes (under-structure or Unterbau). The number of nodes and internodes of the under-structure varies considerably. A tendency to the reduction of the under-structure to its minimum is evident, that is, three internodes delimited by the nodes of the first (α) and second (β) prophylls. The buds of the β-prophyll most commonly continue branching. End flowers may recede to their secondary flowers, whereby they are shifted to a lateral position. Thus, sympodial systems, superficially resembling racemes, may arise in which flowers, though lateral in position, lack a subtending leaf and have instead one opposite to it. These are the so-called spur shoots (Child 1979). This pattern is often further modified by recaulescence in such a way that each flower is accompanied by two leaves (the so-called geminate leaves), of which one, usually the smaller, corresponds to the β-prophyll of the same flower and the other to the β-prophyll of the preceding flower. In many Solanaceae, and more often in Solanoideae, these sympodial systems are particularly rich in number of sympodial units in the upper parts of the main shoot. This is the so-called acrotonic development promotion (Troll 1969). If these branching systems continue forming normal leaves, they represent the main photosynthetic part of the whole plant. Such foliose sympodial chains are the so-called anthocladia (Troll 1969). Anthocladia may further become independent from the main shoot’s under-structure if they produce adventitious roots. Reduction of the main shoot’s under-structure leads to a creeping habit of the anthocladia. Complete loss of the under-structure and independence of the anthocladia is clearly evident in Lycianthes asarifolia and Nierembergia repens (Troll 1969). Anthocladia may further become zonalized by abortion or caducity of the basal sympodial members (Przewalskia tangutica). Genetic control of branching system development is just beginning to be understood (Lippman et al. 2008).

Flower Structure. Perianth usually 5-lobed, rarely 5-parted or unevenly 4-toothed or 4-lobed (Nothocestrum, some Lycium, etc.), or 6-lobed (Goetzea). Calyx synsepalous, actinomorphic, sometimes zygomorphic (e.g. Brugmansia, Nicotiana undulata), exceptionally asymmetric (Sclerophylax, Nolana), or circumscissile after anthesis (Datura), entire (Tubocapsicum, Capsicum, etc.) or with 5–10 linear teeth or appendages (Lycianthes, Capsicum). In some genera tube almost absent (Athenaea, Nectouxia, Markea, Schizanthus, etc.). Calyx usually persistent, very frequently accrescent around the berry or capsule, sometimes inflated and the upper sector almost closed (Physalis, Quincula), or tightly appressed to the pericarp or loosely enclosing it and widely open in the upper sector (Chamaesaracha, Leucophysalis), or reflexed (Jaltomata spp., Schraderanthus, Capsicophysalis) and exposing the mature berry. Nicandra is unique for having auriculate calyx segments.

Corolla gamopetalous, very small (2.5 mm long.) to very large (100–370 mm long in Solandra), actinomorphic, slightly (e.g. Sclerophylax) or distinctly zygomorphic (e.g. Browallia, Streptosolen, Schultesianthus), in general rotate, stellate, tubular, infundibuliform or hypocrateriform (papilionate in Schizanthus); aestivation valvate (Aureliana, Dyssochroma), valvate-induplicate (Cestrum, Jaltomata, Protoschwenckia), valvate-plicate (Capsicum), valvate-conduplicate (Jaborosa, Oryctes, Schwenckia), valvate-supervolute (each lobe has one of its edges rolled inwards and overlapped by its opposite edge, two lobes being dextroinvolute and the other three sinistroinvolute, tribe Anthocercideae), cochlear (Petunia, Nierembergia, Lycium, Markea, Schultesianthus), cochlear-conduplicate (Latua), cochlear-plicate (Nicandra, some Lycium), contorted-induplicate (Exodeconus), contorted-conduplicate (Sclerophylax, Metternichia, Datura), quincuncial (Duckeodendron), reciprocative (the anterior induplicative lobe covers the four others which are conduplicate, Calibrachoa); exceptionally tube very short (Athenaea), lobes usually entire, or deeply laciniate in Schizanthus, and generally equal (except Browallia with unequal lobes). In Lycium and Cestrum, the corollas leave, after falling off, an annular persistent basal portion around the ovary base (called cyathium by some authors).

Several filament and anther features are of interest in recognizing genera and suprageneric taxa. These can be summarized as follows. The basic number of stamens in the family is five. This condition is maintained in Solanoideae. There is an exceptional increase or decrease in stamen number due to alteration of the symmetry of the whole flower (Robyns 1931; Knapp 2002a). In Cestroideae and Nicotianoideae, reduction in the androecium by abortion of 1 or 3 stamens is frequent. When 4 stamens are fertile, reduction or loss of the anterior stamen has occurred (Fig. 57A, N, O, Q). Four fertile stamens are found in some genera of tribes Anthocercideae, Salpiglossideae, and Schwenckieae. The lateral pair is normally longer than the dorsal pair (Fig. 57A). If two fertile stamens are present, this condition may be achieved by further reduction of either the lateral, as in Leptoglossis (Fig. 57P) and Hunzikeria, or the ventral stamens as in Schwenckia and also in Schizanthus (Fig. 57R). Six stamens are exceptional, such as in Goetzea.

Fig. 57.
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Solanaceae. Androecium. A–M Examples of major stamen modifications. A Bouchetia anomala, ventrifixed anthers. B Nectouxia formosa, apical geniculation of the filament. C Solanum herculeum, filament fusion, basifixed anthers and anther dehiscence by apical slits. D Nicandra physalodes, basal geniculation of the filaments. E Symonanthus aromaticus, anther adnation. F Browallia americana, anther asymmetry. G Solanum sciadostylis, dorsal and ventral connective outgrowths. H Anthotroche pannosa. I Nierembergia browallioides, filament adnation. J Juanulloa wardiana, confluent thecae. K Capsicum chacoense, wing-like expansions of the filament. L Jaborosa integrifolia. M Solanum lycopersicum. N–R Diagrams showing reduction in the androecium. N Bouchetia, Leptoglossis (in part), Melananthus, Plowmania, Protoschwenckia, Reyesia (in part), Salpiglossis. O Browallia and Streptosolen. P Hunzikeria, Leptoglossis (in part), Reyesia (in part). Q Brunfelsia, Crenidium, Cyphanthera, Duboisia. R Schizanthus and Schwenckia. (A–M from Hunziker 2001, N–R orig.)

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Fusion at the filament level takes place when an organ complex is formed in which the boundary of the individual stamens cannot be recognized anatomically. This condition is found in Solanum, though it may be masked by additional fusion of the filaments to the corolla tube. In some species, however, it is clearly evident because the filaments are fused also above their fusion with the corolla tube. Adnation, i.e. the bonding of stamens without losing anatomical boundaries, may take place at the anther level—e.g. in Solanum sect. Lycopersicon (Fig.  57M) and Symonanthus (Fig.  57E)—and at the filament level, as in Nierembergia.

Filaments may have lateral auriculate appendages at their base. In addition, these expanded segments may be fused to the corolla tube to form the so-called stapet (Ritterbusch 1976), which provides important features in recognizing Capsicum (Fig.  57K) and allied genera. The filament expansions are functionally related to nectar conduction from the disc to the place of presentation. Geniculation or curvature of the filament may occur near the anther (e.g. Atropa, Mandragora, Solandra, Sclerophylax and Jaborosa, Fig.  57L). In Nicandra (Fig.  57D) and Cestrum, filaments are geniculate near the base.

Filament attachment is also variable. The most common conditions are dorsifixed and basifixed. Ventrifixed anthers are present in many genera such as Cuatresia, Jaltomata, Withania, Trianaea, Schwenckia, Melananthus, Heteranthia, Nierembergia, Leptoglossis, Bouchetia (Fig.  57A), and in the Australian genera.

Connective outgrowths are of systematic and ecological importance in Solanum sect. Pachyphylla (ex Cyphomandra, Fig. 57G); lateral connective expansions are also present in Nectouxia (Fig. 57B). Thecal fusion occurs when the two thecae of an anther share the sporogenous tissue. Thus, anthers have two loculi instead of four and one semicircular dehiscence line instead of two straight lines (Fig. 57H, J, Q). Such synthecous anthers are found in Brunfelsia, five genera of Anthocercideae, in Markea spp. (ex Hawkesiophyton), Juanulloa spp. (ex Rahowardiana ) and some Fabiana species. The dehiscence line is longitudinal in most genera but may be restricted to a longitudinal apical slit (Fig. 57C) or pore (Solanum in part, Lycianthes). In some Solanum, in addition to the larger end pore, a longitudinal row of short slits is found which may enlarge and be fused after opening (Carrizo García et al. 2008). Anther asymmetry by the unequal development of the thecae in one anther may result in size differences (Nierembergia, Fig. 57I) or in abortion of one theca (Browallia and Streptosolen, Fig. 57F, O).

The ovary in the Solanaceae is typically superior, bicarpellate, and bilocular (Fig. 58R). Solandra stands out with a partly inferior ovary (Fig. 58V), as well as Nothocestrum and Datura. In Nicandra (Fig. 58Y) and Trianaea, there are 5- or 4-carpellate ovaries while in some Jaborosa species ovaries are 3- to 5-carpellate. In Nolana, the development of the carpels is unique in the subdivision of the loculi into locelli (Bondeson 1986; Di Fulvio 1971; Huber 1980).

Fig. 58.
figure 3

Solanaceae. Fruit and gynoecium. A–J Fruits. A Berry with accrescent and inflated calyx (partially removed) of Nicandra physalodes. B Berry with two macroscopic stone cells of Lycium bridgesii. C Pyxidium of Scopolia japonica with accrescent calyx removed. D Berry of Lycium gilliesianum. E Berry with scattered sclerosomes of Lycianthes rantonnei. F Capsule of Vestia foetida. G Berry of Atropa belladonna. H–J Fruit of Lycium boerhaviifolium. H Lateral view, I Lateral view of pyrene, J Cross section showing the two pyrenes. K–Z′ Gynoecia. K, L Melananthus multiflorus. K Longitudinal section of basal part of flower showing ovary and nectary. L Pistil. M, N Cross sections of ovary in the upper and basal part of Datura ferox. O, P Vassobia iochromoides. O Pistil. P Longitudinal section of ovary. Q Pistil of Browallia americana. R, S Solanum nitidibaccatum. R Cross section of ovary. S Pistil. T, U Lycium cestroides. T Longitudinal section of pistil showing cryptic nectary. U Pistil. V Longitudinal section of ovary Solandra grandiflora. W Ovary base with nectary of Mandragora officinarum. X Stigma and upper part of style of Leptoglossis lomana. Y Cross section of ovary of Nicandra physalodes. Z, Z′ Cross section of upper part of style, and style and stigma of Bouchetia anomala respectively. (From Hunziker 2001)

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Regarding the number of ovary locules, some variation is found: Trianaea has 8–10 locules, Solandra has 4 locules while Coeloneurum and Henoonia have only one. In some genera, the number of locules varies along the ovary due to the development of false septa; thus, some Lycium ovaries are 4-locular in the upper half and bilocular in the lower half, whereas in Datura (Fig. 58M, N) and some Vassobia species they are 4-locular only in the lower half. The ovules are mostly borne on more or less fleshy axile placentas, or on parietal (Tsoala) or basal placentas (Melananthus); in Sclerophylax, ovules are pendant. Four different types of ovules have been reported so far (Barboza 1991): anatropous, anacampylotropous, hemitropous and hemicampylotropous. There are usually numerous ovules per locule (Fig. 58M, P, R, V, W), but sometimes they can be reduced to a few or one (Fig. 58K).

Styles are mostly cylindrical, solid, and terminal (Fig. 58K, O, U); occasionally, they can be hollow with a stylar canal (Fig. 58Z), and less frequently subterminal. Browallia (Fig. 58Q) and Streptosolen stand out by having a corrugated style. Stylar heteromorphism was recorded in some Solanum spp., Athenaea and Aureliana. Leptoglossis (Fig. 58X) and allied genera are typical for the presence of a distally broadened style with two thin lateral expansions. Stigmata are chiefly capitate and bilobed with an evident slit (Fig. 58O), except for Schizanthus with a very small inconspicuous stigma. The receptive surface has small to medium-sized 1- to 4-cellular papillae which are absent in some taxa, e.g. Nierembergia linariaefolia and Solanum nigrum. The majority of the analyzed stigmata are wet with the exception of Solandra where they are dry. Although the stigma has a comparatively large surface in Browallia (Fig. 58Q), only two small lateral areas are receptive. Most genera have a floral nectary while a few of them completely lack it. When present, the nectary (Fig. 58K, O, P, T, U, W) is located at the ovary base. Nectar is secreted through modified stomata (Bernardello 2007). There are some variations in structure, shape and colour which are of taxonomic interest: 1. macroscopically inconspicuous and of the same colour as the ovary, but microscopically detectable (e.g. Browallia, many Lycium spp., Fig. 58T, U, Vassobia, etc.); 2. macroscopically conspicuous and orange to red carotinoid-coloured (e.g. some Lycium, Salpichroa, Jaltomata, Fig. 58O); 3. invaginated of two types: invaginated-pelviform (Schwenckia) or invaginated-bilobed (e.g. Fabiana, Petunia). Showy special cases of the second type are Mandragora (Fig. 58W) with two opposite commissural lobes, and Melananthus with a cup-shaped nectary (Fig. 58K, L).

Extrafloral nectaries occur on the outer surface of the corollas (Anderson and Symon 1985) or on the abaxial surface of the leaves. Extrafloral nectaries on the calyces of Physalis viscosa and Lycium (sub nom. Phrodus and Grabowskia) have been reported (Bernardello 1987; Rodríguez 2000).

Embryology. Anther wall formation follows the basic as well as the dicotyledonous type; posterior divisions are frequent in the middle layers. In the mature anther wall, the epidermis may become papillate, the endothecial cells develop lignified thickenings following different patterns (reticulate, helical, annular), and sometimes also middle layer cells remain (Carrizo García 2002a; Barboza and Carrizo García 2005); in species with poricidal dehiscence, the lignified layers usually are restricted to the zone surrounding the pores (Siddiqui and Khan 1988; Carrizo García 2002b). The tapetum is glandular and belongs to the pachymerous type (inner tapetum cells larger and radially more elongated than the outer tapetum cells); a very common feature is the presence of an early placentoid arisen from the connective tissue (Hartl 1963). In many species, the so-called resorption tissue differentiates in the septum, usually in hypodermal position; this tissue is placed in subhypodermal position in poricidal anthers, except in the pore zone (e.g. Solanum betaceum, S. glaucophyllum, Carrizo García et al. 2008). The resorption tissue is formed by a variable number of cells of which the cytoplasm is gradually replaced by calcium oxalate crystals in the first stage of development (D’Arcy et al. 1996). Regarding tissue thickness, some are well developed, up to 12 cells wide in cross section (e.g. Nicotiana glauca), and others are very narrow, only two to four cells wide (e.g. Solanum).

Normally, two rows of archaesporial cells occur; cytokinesis of pollen mother cells is simultaneous; the microspore tetrads are mainly tetrahedral, but isobilateral (e.g. Hyoscyamus and Atropa, Sharma et al. 1987) and decussate tetrads (e.g. Nicandra physalodes, Prasad and Singh 1978) can also be found. Pollen is usually 2-celled at shedding but 3-celled pollen has been found in species of Capsicum (Davis 1966), Nicotiana (Jagannadham 1988), Jaborosa, and Salpichroa (Barboza 1989).

The ovules are always tenuinucellate and unitegmic, with a well-developed endothelium. The nucellus forms a hypodermal archaesporial cell, which enlarges and functions directly as megaspore mother cell; rarely two archaesporial cells are found (e.g. Solanum spp.). The megaspore mother cell usually gives rise to a linear tetrad, T tetrads have been reported in Nicotiana tabacum (Goodspeed 1947) and some Solanum spp. (Mohan and Singh 1969; Siddiqui and Khan 1986); in both tetrad types, the proximal megaspore is functional. The female gametophyte is usually monosporic and develops according to the Polygonum type; bisporic embryo sacs of either the Allium or the Endymion type (Modilewski 1935; Pavari 1957; Dharamadhaj and Prakash 1978), or of the tetrasporic Adoxa type (Nanetti 1912) have also been observed; in all cases, the antipodals are ephemeral. Embryogenesis conforms to the Solanad type; a few species follow the Onagrad type (e.g. Przewalskia tangutica, Lu et al. 1999). Polyembryony has been reported in a few members of the family (e.g. Solanum torvum, Mohan and Singh 1969). The development of the endosperm is cellular, exceptionally nuclear (e.g. Schizanthus pinnatus, Samuelsson 1913) or helobial (Hyoscyamus niger, Svensson 1926).

Pollen Morphology. Pollen grains show a broad range of variation in size, shape, apertures, aggregation and exine ornamentation. Palynological characters have been important in contributing to the delimitation at different taxonomic levels (Barboza 1986; Moncada and Fuentes 1993; Persson et al. 1994; Bernardello and Luján 1997; Knapp et al. 1997) and in providing support for hypotheses of relationships between genera (Stafford and Knapp 2006).

Almost 50% of the genera have medium-sized pollen grains (range: 25–50 μm), the smallest ones have been reported in some Lycianthes taxa, i.e. L. laevis (mean 10.7 μm, sub nom. Solanum bigeminatum) and L. lysimachioides (mean 11.6 μm, sub nom. Solanum lysimachioides; Sharma 1974), and in Schwenckia, Melananthus and some Solanum where pollen grains measure 12–20 μm. Some Nierembergia spp., Hyoscyamus, Salpichroa, Tsoala tubiflora and Plowmania nyctaginoides have the largest grains (60–96 μm). The shape is commonly subspheroidal (ranging from suboblate to subprolate), only few genera have also oblate (Browallia, Leucophysalis, Tubocapsicum and Espadaea) or prolate (Exodeconus, Atropa, Chamaesaracha, etc.) or exclusively oblate grains (Markea, sub nom. Hawkesiophyton and Schultesianthus; Persson et al. 1994); Schizanthus is the only genus with prolate spheroidal to perprolate grains (Fig. 59G). The typical condition is the presence of trizonocolporate apertures. However, tri-, tetracolporate (Latua, Fabiana, Witheringia, Nothocestrum, Schwenckia, Plowmania, Protoschwenckia and Solanum) or tetra-, pentacolporate (Physochlaina) grains may co-occur, and in Browallia and Streptosolen grains are up to 9-colpate or colporoidate (Fig. 59C). Exclusively tetra- or pentaporate (some Jaborosa) or exclusively tricolpate (Metternichia, Coeloneurum, Goetzea, Henoonia) or tri- or tetracolpate or colporoidate grains (Benthamiella, the Australian genera, some Brunfelsia, Tsoala and Espadaea) are more unusual in Solanaceae. Anisodus and Mandragora have pantoporate pollen grains. In Solanum, inaperturate grains appear in functionally female flowers in all cases of dioecy (S. appendiculatum, S. confertiseriatum, and many spp. of subgen. Leptostemonum). There exist some remarkable peculiarities concerning the morphology of the apertures. One is the case of the cryptaperturate pollen of Mandragora where there are no ectoapertures but a symmetrical pattern of endoapertural thinnings occurs on a thick and homogeneous endexine layer (Diez and Ferguson 1984); in Datura and Brugmansia the endoaperture is a continuous endocingulum with indistinct margins (Fig. 59A; Persson et al. 1999); finally, in some genera the exine is greatly thickened around the ectoapertures forming an annulus as in Jaborosa sect. Jaborosa (Barboza 1986) or a distinct margo as in Merinthopodium (Persson et al. 1994), tribe Benthamielleae, Petunia, Calibrachoa, Reyesia chilensis, and Salpiglossis spinescens (Stafford and Knapp 2006) or some Solanum spp.

Fig. 59.
figure 4

Solanaceae. Pollen SEM micrographs. A Brugmansia suaveolens, ectocolpus, endocingulum, striate exine, x 2000. B Leptoglossis ferreyrae, reticulate exine, x 4000. C Streptosolen jamesonii, hepta-aperturate pollen, rugulate exine, x 2000. D Metternichia principis, echinate exine, x 5800. E Salpiglossis sinuata, tetrad, gemmate exine, x 1200. F, G Schizanthus pinnatus. F Detail of apocolpium, exine microgranulate, x 5400. G Equatorial view, x 2000. H Brunfelsia sp., alveolate exine, x 2000. (Orig.)

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Pollen grains are normally shed as monads; tetrads also occur in Nierembergia, Bouchetia, Salpiglossis (Fig. 59E) and Reyesia, and only rarely pollen in massulae is found in some Nierembergia spp. (e.g. N. rigida, N. hatschbachii, etc.). Intra- and intergeneric differences are observed in the ornamentation of the exine (Gentry 1979, 1986; Barboza 1986; Persson et al. 1994, 1999; Bernardello and Lujan 1997; Knapp et al. 2000; Stafford and Knapp 2006). In general, the exine is reticulate (Fig. 59B), scabrate, striate (Fig. 59A), rugulate (Fig. 59H), punctate-foveolate, striate-reticulate or striate-rugulate. The presence of a psilate (some spp. of Cestrum, Nicotiana, Physalis and Salpichroa), echinate (Metternichia, Fig. 59D, Exodeconus, Markea lopezii, Tsoala pubiflora, Goetzea, Coeloneurum, Espadaea and Henoonia), microechinate (Latua, Lycianthes, some Physalis spp.), or faintly granular exine (Nothocestrum, Solanum spp.) is less frequent, and exceptionally the exine is rugulate-reticulate or rugulate-striate (Browallia), microgranulate (Schizanthus, Fig. 59F, Heteranthia), scabrate-gemmate (Anisodus), fossulate (some Salpichroa spp.), or smooth (some Jaborosa). In Mandragora, many blunt spines (1.5–2 μm high) are distributed on the dense and fine gemmate-clavate exine (Diez and Ferguson 1984).

The structure of the exine has been little studied in Solanaceae. In general, it is completely tectate (Brunfelsia, Cestrum, Solanum, etc.), or tectate in the equatorial area and with bacula not joined to a tectum in the polar region (some spp. in Juanulloeae), or semitectate (e.g. Lycium) or intectate (Mandragora). The sexine may be up to 6 times thicker or more rarely of the same thickness or thinner than the nexine; however, the nexine is thicker than the sexine in Vestia and Juanulloa (sub nom. Rahowardiana). The imperforate or perforate tectum is usually supported by short and thick (Brunfelsia, Reyesia, Salpiglossis, Merinthopodium, etc.) or by short and granular (Hunzikeria, Streptosolen) or by not well-defined columellae (Cestrum, Solanum); in Sclerophylax, the perforate tectum is on prominent columellae, some of which are bifurcate (Gentry 1986). The foot layer is extremely thin (e.g. Brunfelsia), or barely perceptible (Streptosolen). Melananthus, however, is characterized by an undulating continuous foot layer. Some Australian genera have crowded columellae more or less uniform in length and an irregular and moderately thick foot layer (Knapp et al. 2000). Schizanthus stands apart from the other genera and is readily distinguished by the radially and transversely channelled tectum and foot layer. This character is shared only with Schwenckia (Gentry 1979).

Karyology. Solanaceae exhibit a dysploid series from x = 7 to x = 14 (15?), although other basic numbers such as x = 17, x = 19 and x = 23 have been recorded (Moscone 1992; Hunziker 2001). The chromosome number distribution in the subfamilies (fide Olmstead et al. 2008) is as follows (the figures within taxa are arranged according to their decreasing frequency): Goetzeoideae: x = 12, 13 (Espadaea: x = 12; Metternichia: x = 13); Cestroideae: x = 12, 8, 11, 10 (Cestreae: x = 8; Browallieae: x = 11, 10, 12; Salpiglossideae: x = 11); Nicotianoideae: x = 9, 10, 12, 8 (Nicotiana: x = 9, 10, 12, 8; Anthocercideae: x = 9, 10); Solanoideae: x = 12, 17, 23, 14, 13, 11, 10, 15, 9 (exceptions to x = 12 are Nicandra physalodes: x = 10; Hyoscyameae p.p.: x = 17, 14, 11; Physalideae: Physalis lanceolata: x = 10, Quincula lobata: x = 11; Capsiceae: Capsicum p.p.: x = 13; Solaneae p.p.: i.e. Solanum subgen. Archeosolanum: x = 23, one cultivar of Solanum lycopersicum: x = 13, Solanum mammosum: x = 11, Solanum bullatum: x = 15). In tribes and genera not assigned to a subfamily, the chromosome number distribution is: Schizanthus: x = 10; Benthamielleae: x = 11; Petunieae: x = 8, 9, 7, 10, 11 (Fabiana: x = 9; Calibrachoa: x = 9; Petunia: x = 7; Brunfelsia: x = 11; Leptoglossis: x = 10; Nierembergia: x = 8, 9; Bouchetia: x = 8; Hunzikeria: x = 8); Schwenckieae: x = 12. By far the most common base number is x = 12 which was found in more than 50% of the species studied, including the large subfam. Solanoideae where it is almost universal.

Twenty-five of the 96 genera of Solanaceae are karyologically unknown, some of them comprising several species [e.g. Cuatresia (15), Juanulloa (11), Larnax (32) and Markea (9)], and another 13 being monotypic (Darcyanthus, Duckeodendron, Heteranthia, Oryctes, Plowmania, Protoschwenckia, Tsoala, Calliphysalis) or unispecific (Coeloneurum, Discopodium, Henoonia, Nectouxia). The main gaps are in tribe Schwenckieae where only one count was done and in tribe Juanulloeae where just two reports for one of 4 genera are available. Furthermore, 8 genera belonging to tribe Physalideae remain unexplored regarding their chromosome numbers. Finally, in Solanum, less than half of its ca. 1400 spp. have been examined.

A comprehensive hypothesis on possible paths of chromosome number changes during the evolution of the family is still lacking, and even its original base number is a matter to be clarified. Raven (1975) postulated x = 12 as being plesiomorphic in Solanaceae; based on more recent chloroplast DNA phylogenetic studies, however, Olmstead et al. (2008) have suggested that this number is apomorphic. Evidence for dysploid changes from x = 12 to x = 13 via Robertsonian translocations, i.e. centric fissions, has been reported in Capsicum (Moscone et al. 1993, 2007) and Solanum (sub nom. Lycopersicon, Banks 1984).

Polyploidy has occurred at least in 22 genera belonging to subfamilies Cestroideae (tribe Browallieae), Nicotianoideae (Nicotiana and tribe Anthocercideae), Solanoideae (Mandragora, Lycium, tribes Solaneae and Hyoscyameae), and in tribe Physalideae. The highest chromosome number was found in Scopolia japonica (2n = 14x = 168). Polyploids are remarkably frequent in Nicotiana where ca. 40% of the species are tetraploids often arisen by amphidiploidy, e.g. N. tabacum (2n = 2x + 2x = 48), as demonstrated by genomic in situ hybridization (Kenton et al. 1993). Furthermore, polyploid series are conspicuous in some genera with x = 12, i.e. Mandragora (2x, 7x, 8x), Lycium (2x, 3x, 4x, 8x, 10x), Physalis (2x, 4x, 6x), Chamaesaracha (2x, 3x, 4x, 6x), Withania (2x, 3x, 4x, 8x), Solanum (sect. Solanum: 2x, 4x, 6x, 8x; sect. Petota: 2x, 3x, 4x, 5x; sect. Lathyrocarpum: 2x, 4x, 6x) and Scopolia (4x, 7x, 14x).

The appearance of B chromosomes has only been recorded in few species of the following genera (the number of species and Bs is indicated in brackets): Cestrum (1: 1–6 Bs), Combera (1: 1–5 Bs), Datura (1: 1–6 Bs), Nicandra (1: 1 B), Nicotiana (1: 1 B), Pantacantha (1: 1–3, 6 Bs), Petunia (2: 1–2 Bs) and Solanum (11: 1–3 Bs; Jones and Rees 1982; Moscone 1989).

Karyotype analyses have revealed that the family shows high variation in chromosome size with a range of mean lengths from 1.0–1.5 μm in Metternichia, Solanum sect. Basarthrum and Atropa, to 6.5–11.51 μm in Cestrum (Las Peñas et al. 2006), and Solanum (sect. Pachyphylla and sect. Cyphomandropsis). Nevertheless, most species have comparatively small or medium-sized chromosomes with average lengths of 1.5 to 5.0 μm (Moscone 1989; Bernardello and Anderson 1990; Berg and Greilhuber 1993; Pringle and Murray 1993; Badr et al. 1997; Acosta et al. 2005; Las Peñas et al. 2006; Chiarini and Barboza 2008; Chiarini et al. 2010). Measurements of the nuclear DNA content, available only for few genera, show that genome size (1C-value) of species with diploid complements varies from 0.6 pg (588 Mbp) in Solanum chacoense and S. tripartitum to 24.8 pg (24304 Mbp) in Solanum huilense (sub nom. Cyphomandra hartwegii var. ramose, Bennett et al. 2000; Stiefkens and Bernardello 2000).

In general, karyotypes are symmetrical with a majority of metacentric chromosomes of rather similar size; exceptions to this are found in Nicotiana and Solanum sect. Acanthophora where complements mostly composed of subtelocentric and submetacentric chromosomes are frequent. In the latter cases, the chromosomes often have different lengths although bimodal karyotypes are rare. Many genera show a high constancy in chromosome number, shape and size between species. However, in some genera (Nicotiana, Capsicum, Solanum) evidence accumulates that the direction of evolutionary changes due to chromosome rearrangements implies karyotype re-patterning with asymmetry increase in the advanced taxa.

Chromosome banding approaches and gene mapping by fluorescent in situ hybridization have revealed useful markers for chromosome identification to prompt phylogenetic conclusions in some genera such as Cestrum (Berg and Greilhuber 1993, etc.), Capsicum (Moscone et al. 1993, 1995, 2007, 2011), Solanum (Pringle and Murray 1993) and Nicotiana (Parokonny and Kenton 1995; Lim et al. 2000). These studies have demonstrated that the heterochromatin is highly variable between species with respect to amount and distribution. Furthermore, in Cestrum, it has special properties as it appears in clustered, rather than blocked, bands which in part behave as cold-sensitive regions.

Pollination. Solanaceae often have flowers adapted to specialized types of pollination (Cocucci 1999; Knapp 2010a). Almost every known pollination syndrome is present in the family: bee-, bird-, moth-, butterfly-, fly- and bat-pollination. However, unspecialized pollination systems are also found in, for example, Lycium. Reward systems include almost all known kinds of primary attractants, except food-bodies and resins: perfume, oil, nectar and pollen. Pollination by deceit, i.e. without any reward, is present in some blowfly-pollinated species of Jaborosa.

Mechanical flower devices for pollen presentation and pollen placement adapted to particular pollinators are noteworthy among butterfly- and bee-pollinated species. These include mobile anthers (rotatory in Leptoglossis or balancing in Browallia and Brunfelsia), explosive pollen release, buzz pollination, and bellow mechanisms.

At the tribal level two or several syndromes commonly occur. At the generic level contrasting patterns are found: Solanum is presumably bee-pollinated throughout, whereas Nicotiana shows infrageneric syndrome radiation with bee-, moth-, bird- and bat-pollination. In the Physalideae (Acnistus, Cuatresia, Physalis, Vassobia, Aureliana, Athenaea), where bee-pollination prevails, flowers attract pollen-collecting females, fragrance-collecting Euglossine males or nectar-feeding bees of both sexes. In Euglossine-pollinated species of Solanum (ex Cyphomandra), fragrance is produced by the prominent anther connective. Bat-pollination is widespread in the epiphytic genera (Dyssochroma, Juanulloa, Markea, Trianaea), while butterfly-pollination is prevalent in Salpiglossis and Reyesia. Nierembergia is unique for its association with oil-collecting bees. One Coleoptera genus, Epicante, feeds exclusively on Nolana corollas and may complete pollination, and on at least one occasion Phodopis vesper (a northern Chilean hummingbird) has been observed pollinating Nolana villosa; Nolanomelissa toroi has been found collecting pollen and nectar from Nolana rostrata in Chile (Rozen 2003).

Reproductive Systems. In general, Solanaceae have hermaphrodite flowers, wet stigmata and binucleate pollen when shed. The stigma surface consists of a thick lipidic upper layer and a thin aqueous under-layer. Pollen captured on the lipidic layer sinks through the aqueous layer and hydrates (Konar and Linskens 1966a, 1966b). In agreement with the binucleate pollen and the stigma features, the gametophytic type of self-incompatibility system prevails, i.e. genetic recognition and rejection of incompatible pollen takes place during the passage of the pollen tube through the style (Heslop-Harrison and Shivanna 1977).

Breeding systems derived from this general pattern have been described. A number of species has unisexual flowers distributed monoeciously or dioeciously. Andromonoecious species are common in Solanum subgen. Leptostemonum (Whalen and Costich 1986; Miller and Diggle 2003). The few reported dioecious species are found in Solanum (Levine and Anderson 1986; Anderson and Symon 1989; Knapp et al. 1998; Knapp 2010b), Deprea (Sawyer and Anderson 2000), Dunalia (Hunziker 2001), Lycium (Minne et al. 1994; Miller and Venable 2002) and Withania (Hepper 1991; Anderson et al. 2006). Gender dimorphism has evolved independently in six lineages, primarily in the Physalideae (sub Solaneae, Sawyer and Anderson 2000). In some Australian and Central American Solanum spp., flowers are morphologically hermaphrodite but functionally unisexual: male flowers form abortive ovules and female flowers sterile pollen (Anderson 1979; Anderson and Symon 1989; Knapp et al. 1998; Martine et al. 2006). The two spp. of Symonanthus are truly dioecious, and flowers have either rudimentary pistils or stamens (Haegi 1981). Dunalia solanacea has long-styled and short-styled flowers with reciprocity in stamen lengths, as expected from distyly; however, the long-styled flowers are male sterile, but it is not known if the short-styled flowers are female sterile (Rodríguez 1998). Finally, four South African Lycium spp., Withania coagulans from Asia, W. aristata from the Canary Islands (Anderson et al. 2006) and Deprea paneroi from South America are functionally dioecious as well (Sawyer and Anderson 2000). In Nolana stigmas were found to be receptive during a wide range of floral developmental stages, and pollen remained viable throughout the open flower period and into senescence (Douglas and Freyre 2010).

Solanaceae are an experimental model to study gametophytic self-incompatibility systems based on S-RNase (Newbigin et al. 1999; Wang et al. 2001). Self-incompatibility factors of the style, controlled by a single multialeletic locus (S), are glycoproteins with ribonuclease activity. S-RNase from the pistil extracellular matrix is taken up by both incompatible and compatible pollen tubes, but it causes degradation of RNA only in incompatible pollen tubes (McClure and Franklin-Tong 2006). In addition, other factors are also required for pollen rejection (e.g. HT-B). A model of S-RNase compartmentalization in the vacuoles has been suggested recently, in which S-RNase is selectively released later in incompatible pollen tubes (Goldraij et al. 2006); the HT-B factor is involved in the process. The cytotoxic activity of S-RNases causes growth arrest of incompatible pollen tubes by inhibiting translation (McClure and Franklin-Tong 2006).

Fruit and Seed. The family is very diverse in fruit types: berry, capsule, drupe, schizocarp and diclesium; this diversity has been analyzed in a phylogenetic framework (Knapp 2002b). The two most common types are berries or capsules. Regardless of fruit type, the calyx is generally persistent (Fig. 58A, D, G), and rarely circumscissile after anthesis (Datura); sometimes, there are different degrees of calyx accrescence which just surrounds the fruit (Fig. 58G) or envelopes it completely; in some cases, it is clearly inflated, as in Nicandra (Fig. 58A), Physalis, Przewalskia, Withania, etc. (Hunziker 2001).

Capsules, plesiomorphic in the family (Knapp 2002b), are variable in number of seeds: from one in Melananthus, four or five in Metternichia, 30–50 in Fabiana to ca. 5000 in Nicotiana. Capsule dehiscence is almost always longitudinal (Fig. 58F), septicidal, septifragal or septicidal-loculicidal; some genera (Hyoscyamus, Atropanthe, Physochlaina, Przewalskia and Scopolia) have capsules with circumscissile dehiscence (pyxidium).

Berries are clearly derived in the family and seem to have three separate origins (Knapp 2002b). Some berries show different degrees of sclerification with scattered stone cells variable in size: microscopic, small but macroscopic (Fig. 58E), or conspicuous (Fig. 58B). The highest degree of sclerification is reached in the drupe (i.e. two to eight seeds enclosed in a wooden endocarp surrounded by a fleshy mesocarp) observed in some Lycium spp. (Fig. 58H–J), Lycianthes lycioides and Saracha, which are considered derived (Bernardello 1983; Knapp 2002b). Duckeodendron has a very peculiar drupe in which a pericarp with a multi-layered epicarp (epidermis and the loose parenchyma with groups of brachysclereids), a yellow and fibrous mesocarp (long adpressed rough fibres) and a bony dark brown endocarp enclose one seed (Machado et al. 2011). The origin of this fruit type most probably is entirely independent of the drupes found in Lycium and Lycianthes (Knapp 2002b). The unusual sclerified schizocarp of Nolana (Fig. 67J, K) is probably also derived from a berry (Tago-Nakazawa and Dillon 1999). Some Nolana species have 3–6 many-seeded mericarps broadly affixed to the receptacle, others have (2–)5–30 free mericarps.

Some fruits dry after maturity, resulting in the presence of a thin leathery pericarp, as in genera of tribe Hyoscyameae (except Atropa), Oryctes, and some spp. of Solanum subgen. Leptostemonum. Sclerophylax is very peculiar in having a dry indehiscent fruit, generally with one seed per locule, with a membranous pericarp included in the accrescent and sclerified calyx (= diclesium). The origin of this fruit type inferred from the phylogeny of the family has not yet been explored.

Seeds (Fig. 60) are not compressed or more or less compressed, thick, with various forms: subspherical, reniform, discoid, cuboidal, prismatic, polyhedric, ovoid, or more rarely elongated and thin, lenticular or bacilliform. Some Datura spp. have elaiosomes. Seed size varies from 1.2 to 2.5 mm long, but sometimes seeds are smaller (0.3–1 mm long) or larger (4–35 mm long). The testa is variable. It can form a thin (Fig. 60D′) or broad wing, or be smooth, finely reticulate (Fig. 60F″′), tuberculate, honeycombed, etc. The testa cells may be thick-walled, with an undulate or wavy outline (Fig. 60C, H), foveolate or reticulate-foveolate, etc. (Axelius 1992; Freire de Carvalho et al. 1999; Lester et al. 1999).

Fig. 60.
figure 5

Solanaceae. Seed. A Nierembergia linariaefolia. A lateral view, A′ cross section, A″ longitudinal section. B Reyesia parviflora. B cross section, B′ lateral view, B″ longitudinal section. C Schultesianthus leucanthus. C lateral view, C′ embryo, C″ cross section. D Sessea vestioides. D cross section, D′ lateral view, D″ longitudinal section. E Latua pubiflora. E longitudinal section, E′ cross section, E″ hilum view. F Juanulloa verrucosa. F lateral view of embryo, F′ seed, cross section, F″ frontal view of embryo, F″′ seed, lateral view. G Duboisia hopwoodii. G lateral view, G′ cross section, G″ longitudinal section. H Schizanthus grahamii. H lateral view, H′ longitudinal section, H″ cross section. I Lycium gilliesianum. I longitudinal section, I′ cross section, I″ lateral view. J Nicotiana longiflora. J lateral view, J′ cross section, J″ embryo. K Solanum nitidibaccatum. K cross section, K′ embryo, K″ lateral view. (From Hunziker 2001)

Full size image

The embryo has important features for the delimitation of the taxa: form, ratio of its total size and the size of the cotyledons, and how the cotyledons are arranged in relation to the rest of the embryo (Hunziker 2001) are relevant. Regarding their form, embryos are thick (Fig. 60C′, F) or slender (Fig. 60K′), long or short, straight (Fig. 60A″, J″), slightly curved (Fig. 60G″, E), vermiform, annular, coiled (Fig. 60B″, K′) or subcoiled (Fig. 60H′, I), and exceptionally U-shaped. Furthermore, Cestrum, Sessea, Vestia, Merinthopodium, Markea and Juanulloa (Fig. 60F) have embryos with broad cotyledons, unlike all the other taxa where their width is about the same as that of the rest of the embryo. As regards the ratio between total embryo size and cotyledon size, two groups are found: most commonly the cotyledons are equal, slightly longer or up to 2.5–3 times shorter than the rest of the curved or straight embryo; in the endemic Australian genera the cotyledons are between 6–8 times shorter than the rest of the straight embryo (Fig. 60G″). In the majority of the genera (Fig. 60A, B, H, I, K), the embryo has incumbent or oblique cotyledons; some epiphytic genera (Juanulloa, Merinthopodium, Schultesianthus) have accumbent cotyledons (Fig. 60C, D, E, F, J).

There are two main possibilities of seed dispersal: wind dispersal in case of very small seeds (e.g. Nicotiana, Schwenckia) and winged seeds (e.g. Sessea), and animal dispersal mainly by birds eating the berries (as in many Solanum) and probably by ants in Datura spp. with elaiosomes.

In some genera (Solanum, Tubocapsicum, Nicotiana, Capsicum, Nolana), germination is phanerocotylar (Dottori et al. 2000; D’Arcy et al. 2001; Perícola et al. 2004; Machado et al. 2008; Cabrera et al. 2010).

Phytochemistry. Hunziker (2001) and Eich (2008) provided a thorough review of the vast phytochemical literature on the secondary compounds found in Solanaceae. Two recent reviews deal with the withanolides and related steroids in subfam. Solanoideae (Chen et al. 2011; Misico et al. 2011). Eich (2008) and Misico et al. (2011) discussed and summarized the chemical information in a phylogenetic framework. Nearly 75% of the genera have been investigated phytochemically at least in part. Tropane alkaloids, steroidal alkaloids/glycoalkaloids, and withanolides (see below) are important distinctive groups of metabolites in Solanaceae.

According to Eich (2008), eight types of alkaloids have been detected in the family: a) simple pyrrolidines (hygrine and its derivates) found mainly in subfam. Solanoideae, and in members of Schizanthus, Brunfelsia, Anthocercis, Duboisia and Nierembergia liniariaefolia; b) pyrrolidine amides (conioidines), very uncommon and detected only in Chamaesaracha coniodes; c) pyrrolydine-type nicotinoids (nicotine and/or its congeners) in large amounts in genera of subfam. Nicotianoideae (Nicotiana, Crenidium, Cyphanthera, Duboisia) and in extremely low concentrations in some species of Petunia and members of subfam. Cestroideae (Salpiglossis sinuata, Streptosolen jamesonii, Cestrum spp.), and Solanoideae (Solanum lycopersicum, S. melongena, Atropa belladonna, Capsicum annuum, Withania somnifera, Brugmansia candida, Datura spp.); d) tropanes (some very poisonous), the most common alkaloids produced by genera of subfam. Nicotianoideae (except Nicotiana), Solanoideae (Datureae, Hyoscyameae, Mandragora, Solandra, Nicandra, Physalis, Withania, Latua), and Schizanthus and Nierembergia; e) calystegines (polyhydroxylated nortropanes), widespread in subfam. Solanoideae (some Hyoscyameae, Datura spp., Solanum spp., Capsicum spp., Physalis spp., Withania spp., Lycium chinense, Nicandra physalodes) and in Duboisia leichhardtii and Brunfelsia nitida; f) pyrrolizidines, rare in the family and present in Datura stramonium and Nicotiana tabacum; g) β-carbolines, in Vestia foetida, Datura stramonium and two Solanum spp.; h) steroidal alkaloids, common and predominant metabolites of Solanum but also of some other genera (Capsicum, Lycianthes, Saracha, Cestrum, Nicotiana); these alkaloids are pharmacologically and toxicologically very well investigated (see Eich 2008 and bibliography therein). Some alkaloids are unique to one or few species, such as: fabianine (tetrahydroquinoline alkaloid) in Fabiana imbricata; 2-methoxy-3-isobutylpyrazine, mainly responsible for the aroma of the fruit of Capsicum annuum; nicotianamine in Nicotiana spp., Solanum spp., Lycium chinense, and Datura metel; solamines from the roots of two Solanum spp. and Duboisia leichhardtii; pyrrole alkaloids like the poisonous brunfelsamidine in Brunfelsia grandiflora and Nierembergia lineariaefolia, and solsodomines in Solanum sp.; benzodiazepines and catecholamines in Solanum tuberosum; betaines in Solanum wendlandii and Nicotiana tabacum; homo-cholestane glycoside (aethiosides A–C) in Solanum aethiopicum; and cardenolides in Nierembergia rigida.

Pharmacologically more interesting steroidal lactones are the withanolides (Chen et al. 2011; Misico et al. 2011), i.e. polyoxygenated steroids with a C28 ergostane skeleton. Almost 650 withanolides have been isolated so far from 23 genera (ca. 70 spp.) belonging to subfam. Solanoideae, and from only one species of subfam. Cestroideae (Browallia viscosa). Some withanolides have been demonstrated to be good chemotaxonomical markers at different levels (Misico et al. 2011); thus, physalins and related withanolides appear only in subtribe Physalidinae (Physalideae); acnistins are exclusive to subtribes Iochrominae and Withaniinae (Physalideae); the withametelin skeleton is exclusive to Datura; trechonolides, sativolides, and spiranoid withanolides at C-22 are present only in Jaborosa sect. Lonchestigma while spiranoid-γ-lactones have been isolated mainly from species of Jaborosa sect. Jaborosa. The wide biological activity of many withanolides is usually associated with their structural variety. These compounds show insecticidal, phytotoxic, antiparasitic, antimicrobial, anti-inflammatory, antistress, immunosuppressive and immunomodulatory, cancer-related, and CNS-related activities (Chen et al. 2011; Misico et al. 2011).

Flavonoids, especially flavones and flavonols, are common in many Solanum species (Sarmento da Silva et al. 2003), and flavonoid aglycones in Nicotiana, Petunia, Solanum, Datura, Physalis, Chamaesaracha, Iochroma, Atropa, Hyoscyamus and Salpiglossis (Wollenweber et al. 2005). Anthocyanins are the flower, tuber and fruit pigments best studied in the family, such as in Petunia/Calibrachoa (Ando et al. 1999, 2004), Brunfelsia (Vaknin et al. 2005), and Solanum (Brown 2005; Lachman and Hamouz 2005; Ichiyanagi et al. 2005) species. Monoterpenoids and sesquiterpenoids were studied as volatile compounds in the fragrance of flowers and other plant parts of Cestrum, Nicotiana, Brugmansia, Datura, Solanum, Capsicum, Fabiana and Lycium, while different diterpenoids were reported in Nicotiana spp., Petunia x hybrida, Cestrum parqui, Physalis coztomatl, and Nolana and Capsicum spp. Finally, triterpenoids and phytosterols were detected in many genera of the family (for details, see Eich 2008).

Other compounds are very rare in the family and appear in individual species or genera, such as β-phenylethylamine, tyramine and its N-methyl derivatives in Nierembergia linariaefolia; phenylpropanoid acids, phenylethanoids, and phenylmethanoids and others with good phytotoxic activity in Cestrum parqui; two unusual glycosides (cesternosides A and B) in Cestrum nocturnum; capsaicinoids (ca. 25 compounds, capsaicin the dominating alkaloid) in Capsicum spp. fruits; hydroxycoumarins and/or long chain alkyl esters of hydroxycinnamic acids in Atropa belladonna, Fabiana imbricata, Nicotiana tabacum, Schultesianthus leucanthus, Datura stramonium, and Brunfelsia and Solanum spp.; some lignans in Nierembergia rigida and Solanum spp., and lignanamides in Capsicum annuum and Hyoscyamus niger (Eich 2008).

Distribution and Habitats. Solanaceae are widely distributed throughout the world, with the greatest concentration in the New World. The family ranges from sea level (Aureliana, Athenaea, Nolana, some Solanum and Calibrachoa) to 4900–5000 m (e.g. Fabiana bryoides in South America, and Przewalskia tangutica in China). Although the family has many temperate members, the majority of genera and species is tropical; some genera are found in semiarid and arid settings, even deserts, while others are well adapted to montane rainforests. The highest diversity occurs from Central America southwards into South America along the Pacific coast and in the Andean mountains (D’Arcy 1991). The main centre of diversification is South America, where 37 endemic genera occur together with some endemic species of the cosmopolitan genera Solanum, Lycium, Lycianthes and Nicotiana. In the uplands of western South America, endemic species of Saracha, Dunalia, Salpichroa, Iochroma, some Capsicum, Solanum sect. Petota, Nicotiana sect. Tomentosae and Undulatae and some spp. of sect. Paniculatae, etc. are found; several genera have representatives, including most of the lianas and epiphytes, in tropical forests at different elevations, like Acnistus, Streptosolen, Browallia, Deprea, Larnax, Duckeodendron, Juanulloa, Markea, Trianaea, Schultesianthus, Merinthopodium, most Solanum, etc.; the western coast of South America (5–30°S) is dominated by desert conditions except for localities where recurring fogs along the shore allow for the development of plant communities termed lomas formations; in the lomas formations occur 16 genera and ca. 130 species (no fewer than 95 endemic species) belonging mainly to Exodeconus, Jaltomata, Leptoglossis, Lycium, Nicotiana, Nolana, Reyesia, Schizanthus, Salpiglossis and Solanum (Dillon 2005). Nolana, with 75 endemic lomas species, stands out as the largest and most wide-ranging genus to be encountered in all lomas formations. Towards eastern and southern South America, the diversity is less pronounced; to the north in the subtropics, Vassobia, Eriolarynx, Schwenckia, Metternichia, Protoschwenckia, etc. are found; from temperate areas in Chile to the south, with some genera reaching Argentina, Latua, Vestia, Schizanthus, Reyesia, Salpiglossis, Nierembergia, Fabiana, Jaborosa, Combera, Benthamiella, Pantacantha, etc. grow; towards north-eastern South America, mostly in tropical and subtropical lowlands, Aureliana, Athenaea, Petunia, Calibrachoa, some Capsicum, etc. are centred. Sclerophylax is endemic to Argentina and adjacent parts of Paraguay and Uruguay.

Other centres of Solanaceae diversity are 1) Mexico and North America, including Physalis and its close relatives (Chamaesaracha, Tzeltalia, and Quincula), Datura, and the two endemic genera Nectouxia (Mexico) and Oryctes (western North America), plus some endemic species of Cestrum, Solanum, Lycianthes, Lycium and Nicotiana; Plowmania and some Bouchetia and Hunzikeria species are also centred in this area but also reach South America; finally, some species have probably been introduced by man (Petunia, Nierembergia) or arrived by local migrations from the south (e.g. Jaltomata); 2) Central America, with centres of upland and lowland diversity, including species of Cestrum, Brunfelsia, Witheringia, Lycianthes and Solanum sect. Geminata; Goetzea, Espadaea, Henoonia and Coeloneurum are mostly endemic to the semiarid lowlands of the Greater Antilles; 3) Australia and neighbouring islands, with some endemic taxa (7 genera), Nicotiana sect. Suaveolantes (exception: N. africana of Namibia), and Solanum subgen. Archaeosolanum and some groups of subgen. Leptostemonum; 4) a zone in Eurasia from Europe to Japan, particularly in the Himalayan region where Hyoscyameae are the best represented tribe together with Mandragora and Withania (Tu et al. 2010).

Solanaceae are little diversified in the rest of the world. For example, in Africa only a few genera are present, namely the two widely ranging genera Solanum and Lycium, three in common with Europe and Asia (Mandragora, Hyoscyamus and Withania), the extraordinary Nicotiana africana, and the endemic Discopodium. In Asia the strictly Asian Tubocapsicum, species of Solanum and Lycianthes, and species of the European genera Atropa, Mandragora, Hyoscyamus and Withania are found. Finally, Nothocestrum is a geographically isolated genus on Hawaii.

Economic Importance. The economic importance of Solanaceae is well documented (Hawkes 1999 and references therein), as well as the role they have played in the development of traditional cultures and civilizations. The family contains not only major food plants such as potato, chili and tomato, but also others locally important and little known outside their ranges.

Among the food plants, the potato (Solanum tuberosum) has a remarkable position, being the world’s fourth most important crop after wheat, rice and maize (Hawkes 1990); its wild close relatives have been used in programs aimed at breeding for disease resistance, environmental tolerances, and other agronomic traits of interest (Hijmans et al. 2002). Solanum lycopersicum (= Lycopersicon esculentum), the only domesticated tomato, is a very popular vegetable crop in the modern human diet worldwide, with significant phenotypic variation in fruit size and shape; wild tomato species all produce fruits which are almost invariably round and small; mutations at six genetic loci seem to have been essential in transforming the small, inconspicuous berries of wild tomatoes to the extremely large fruits now found in modern cultivars (Tanksley 2004). Some other Solanum spp. are cultivated at a commercial scale for their fruits, especially S. betaceum (= Cyphomandra betacea), the “tree tomato” or “tamarillo”. Brazil is the native region of this and New Zealand the main country of fruit production; the interest in the “tamarillo” as a potential new crop is increasing in many frost-free climatic areas of the world (Prohens and Nuez 2000). Solanum melongena (“eggplant”, “berengena”, “aubergine”, “brinjal”), indigenous to Indochina and domesticated in the Indo-Burma region, has medicinal, culinary and even ornamental uses. In India, eggplants are still used for medicinal purposes, but they are cultivated worldwide mostly for their edible berries (Daunay et al. 2007); S. macrocarpon, the “gboma eggplant”, is cultivated throughout much of Africa, especially in Ghana, for its edible fruits and leaves as is the “scarlet eggplant”, S. aethiopicum, widely planted in many parts of tropical Africa, less in Brazil and occasionally in S Italy and the Black Sea Russian republics (Daunay et al. 2001). In South America, S. quitoense (“naranjilla”, “lulo”), S. muricatum (“pepino”, “pepino dulce”) and S. sessiliflorum are reputed for their sweet or fragrant fruits; some cultivars have been developed with a higher sugar content in their fruits (Prohens and Nuez 2001) or plants resistant to root-knot nematodes, such as the S. quitoense ‘Lulo la Selva’ cultivar (Heiser 2001).

Considering their uses as vegetables and condiments, peppers are superb culinary plants. Cultivars commercially grown throughout the world belong to Capsicum annuum and allied species (C. frutescens, C. chinense, C. pubescens, C. baccatum var. pendulum); some—known as “sweet pepper”, “bell pepper”, “pimiento”—produce non-pungent fruits while the plants with pungent fruits are called “chili”, “yellow chilli”, “ají”, “red pepper”, “paprika”, etc.; pungency is due to the presence of a group of metabolites unique to Capsicum spp. known as capsaicinoids, a mixture of phenolic amides in which capsaicin is prevalent. These substances have also shown a wide range of pharmacological properties, such as antigenotoxic, antimutagenic and anticarcinogenic effects (Manirakiza et al. 2003).

Of the great number of Physalis species, very few are used for their fruit. The Old World species P. alkekengi is grown as an ornamental and is popular for its large, bright orange to red husks. Among New World Physalis, some have been domesticated mainly in Mexico from where they were taken to Europe and other parts of the world (Montes Hernández and Aguirre Rivera 1992); the edible fruits of the “tomatillo” or “husk-tomato” (P. philadelphica), the “cape gooseberry” (P. peruviana), and the “ground cherries” (P. pubescens, P. subglabrata, etc.) are consumed raw or cooked in salsas, gazpacho and guacamole (Morton 1987; Montes Hernández and Aguirre Rivera 1992) or are preserved and canned to be exported. In many countries (South Africa, Australia, Greece, Turkey), these species are commercially cultivated at different scale.

Nicotiana is very well known as “tobacco”; economically important spp. are N. tabacum and N. rustica. Nicotiana tabacum is a natural amphidiploid hybrid; its maternal S-genome donor is an ancestor of N. sylvestris and the paternal T-genome donor is derived from a particular lineage of N. tomentosiformis (Murad et al. 2002; Chase et al. 2003). Only a few species are ornamental, although some are cultivated as garden plants for their flowers or their foliage, such as N. alata, while many varieties of diverse flower colour have been produced experimentally (Japan Tobacco Inc. 1994).

The roots of the Mediterranean Mandragora spp., which contain tropane alkaloids, were highly praised as amulets or for their medicinal properties (narcotic, anti-inflammatory, disinfectant, aphrodisiac), and used for all kinds of magic purposes due to their human shape in the Middle Ages. Furthermore, they furnished the most ancient curative drug for humanity. Highly esteemed by the Egyptians, it was mentioned in the Bible, and was also well known to Dioscorides, Theophrastus, Plinius and many other classical authorities. On account of the extraordinary role which “mandrake” played in the popular superstitions and folklore of Europe, it is undoubtedly the most relevant magic plant (Evans 1979).

Atropa belladonna has been known since ancient times for its potent toxic properties, due to the presence of tropane alkaloids in roots, stems, leaves and fruits. It was involved in witchcraft practices in the Middle Ages. Its generic name refers to the goddess Atropos, one of the three Fates in Greek and Roman mythology; this goddess, together with Lachesis and Cloto, was in charge of spinning the thread of human destiny, which Atropos could cut off with her shears whenever she pleased. Nowadays, A. belladonna (“beautiful lady”) and A. baetica are cultivated as a source of atropine, a drug used for therapeutical purposes on account of its depressant effect on the parasympathetic nervous system (Evans 1979).

Hyoscyamus niger (“henbane”, “beleño negro”, “judquiame noire”, “schwarzes Bilsenkraut”) is well known in Europe for its extreme toxicity and has been valued medicinally since very early times. Dioscorides, as well as Plinius, wrote about it and it is thought to have been used in witches’ brews and other toxic preparations during the Middle Ages, together with other species, above all H. muticus. The “white henbane” (H. albus) was used medicinally for its narcotic, anti-inflammatory and disinfectant effects (Daunay et al. 2007). Hyoscyamus niger has been included in the pharmacopoeia of most European countries, being used in general as an external analgesic.

The alkaloid content (especially scopolamine) of the “daturas” (Datura spp.) explains their use as hallucinogens in healing, initiation or divining rites in traditional cultures of Africa, India, the south-western USA, Mexico and South America. The species of greatest importance for these ceremonies and uses are D. inoxia, D. discolor, D. wrightii, D. quercifolia, D. leichhardtii and D. ferox. The “chamico” (D. ferox) in South America and, worldwide, the “jimson weed” or “thorn apple” (D. stramonium) are toxic weedy plants; their toxic seeds contaminate commercial seeds and grains and also invade all sorts of crops (40 crops in almost 100 countries according to Holm et al. 1997).

Solanum glaucophyllum grows in southern Brazil, Uruguay, Paraguay and northern Argentina, and is one of the most important poisonous plants of Argentina (“duraznillo blanco”), being responsible for a serious disease characterized by wasting and calcification of soft tissues in cattle; the toxicity is caused by high levels of 1,25-dihydroxyvitamin D3 (Costa et al. 1998). Another South American toxic species is Nierembergia linariaefolia, known as “chuscho” or “chucho”, causing severe disease in cattle, sheep, goats, horses and rabbits (Gallo 1987; Botha et al. 1999); pyrrole-3-carbamidine is responsible for its lethality (Buschi and Pomilio 1987).

The rare Latua pubiflora (“latúe”, “palo mato”, “palo de bruja”), endemic of S Chile, is a species with narcotic and toxic properties due to its content of atropine and hyosciamine and, to a lesser extent, of scopolamine in stems, seeds and leaves (Schultes 1979). Also some Cestrum spp. (C. sendtnerianum, C. laevigatum, C. dumetorum, C. nocturnum, C. diurnum, C. parqui, C. mathewsii) have poisonous properties or are used as medicinal plants in many countries of the American continent. Cestrum parqui is considered a major problem because of its toxicity to livestock, causing death after ingestion of small quantities of leaves, twigs or fruit, and even dry leaves from plants which have been cut down or sprayed (Gallo 1987).

As regards ornamentals, “petunia” (breeding lines and commercial hybrid varieties of Petunia x hybrida) is a popular ornamental species, a model for flower colour (anthocyanin biosynthesis), and first among bedding plants (Sink 1984). Of some Argentinean Calibrachoa spp. (ex Petunia spp.), genotypes suitable for breeding pot plants were selected for their ornamental value (Facciuto et al. 2006). Species of Brugmansia (“trumpet flower”, “floripondio”), Brunfelsia (“jazmín del Paraguay”, “galán de moda”), Capsicum (“peppers”), Cestrum (“dama de noche”), Physalis (“cape gooseberry”, “winter cherry”, “chinese lantern-plant”), Schizanthus (“butterfly flower”, “fringe flower”, “flor de pajarito”), Solandra (“chalice vine”, “tetona”, “copa de oro”), some Nierembergia (“cup flower”) and Nicotiana, and a few Datura spp., such as D. wrightii, are ornamentals. Nolana paradoxa (“bluebird” and “snowbird”) and N. humifusa (referred to as “N. prostrata”) have found their way into the horticultural trade and wider usage of artificial hybrids amongst Nolana species has been investigated (Freyre et al. 2005).

Finally, the timber of Duckeodendron cestroides is of good quality and similar in texture and properties to ‘marupá’ (Simaruba amara; Record and Hess 1942). In Brazil, where this species is endemic, it is used to make fine wooden furniture as well as smaller products (e.g. matches, Freitas da Silva et al. 1977).

Conservation. Habitat destruction due to deforestation and clearing for agriculture and urban development is the main threat to various species of Solanaceae, especially given the small population size and the restrictive distribution of some. This problem is largest in Central and South American forests, both regions with endemic genera at risk, such as Duckeodendron (Cramer et al. 2007), Latua, Plowmania, Darcyanthus, as well as many threatened species of other genera (Barboza et al. 2010, 2011; Stehmann and Greppi 2011). Some species are listed in the IUCN Red List (2007) in different categories: Solanum diploconos and Aureliana fasciculata, both from Brazil (Lower Risk); some Nothocestrum spp. from the Hawaiian Islands (Critically Endangered), and Nicotiana africana (Least Concern). Australia has provided a list of ca. 35 rare and threatened Solanaceae species (Electronic Flora of South Australia, http://www.flora.sa.gov.au/lucid_keys/Solanaceae/Solanaceae_rare_and_threatened.shtml). Within Solanum, many species are also listed in different risk categories (Queensland Nature Conservation Act 1992; Threatened Species Protection Act 1995; Bañares et al. 2004), while others have been considered in the literature as Critically Endangered (Knapp 2010b), Endangered (Knapp 2007, 2010c; Granados-Tochoy et al. 2007; Knapp and Nee 2009; Giacomin and Stehmann 2011), or Vulnerable species (Farruggia and Bohs 2010; Stern and Bohs 2012). Finally, Solanum fernandezianum is an endemic species in danger of extinction on the Robinson Crusoe Island, Chile (Stuessy et al. 1998; Ricci 2006) where some conservation measures have been taken (Solano et al. 2011).

Classification and Phylogeny. D’Arcy (1979, 1991) compiled in detail the development of thought on the classification of the Solanaceae from classical times to the Third International Solanaceae Congress held in Colombia in July 1988. Since then, Solanaceae have been divided into two subfamilies: Solanoideae and Cestroideae (D’Arcy 1979, 1991; Hunziker 1979; Olmstead and Palmer 1992). The last formal classification was presented by Hunziker (2001). This author recognised 6 subfamilies (Cestroideae, Juanulloideae, Solanoideae, Salpiglossideae, Schizanthoideae and Anthocercidoideae), 27 tribes and 92 genera, excluding 15 genera considered members of other families. The main achievement of this classification was a comprehensive morphological analysis of all genera, revealing unknown characters for many taxa. He also provided palynological, karyological and phytochemical evidence to support his system. Seed shape, cotyledon features, and corolla aestivation were the main characters used to circumscribe the subfamilies. From a phylogenetic point of view, Hunziker’s system recognizes many non-monophyletic groups (i.e. subfam. Cestroideae, tribe Salpiglossidae, tribe Solaneae, etc.; cf. Stafford and Knapp 2006; Olmstead et al. 2008). In 1999, Olmstead et al. proposed a phylogenetic and provisional classification based on chloroplast DNA sequences; their analysis included 52 genera (79 spp.), some of them previously excluded from the Solanaceae, such as Nolana and the Antillean genera of the Goetzeaceae. Additional recent studies strongly support Duckeodendron (Fay et al. 1998; Santiago-Valentín and Olmstead 2003), the four Antillean genera (Santiago-Valentín and Olmstead 2003), and Sclerophylax (Olmstead and Bohs 2007) as nested within Solanaceae. Olmstead et al. (2008) published a new classification of the family based on the chloroplast regions ndhF and trnLF of 89 genera (190 spp.); this classification recognised four subfamilies (Goetzeoideae, Cestroideae, Nicotianoideae and Solanoideae), two unranked informal clades within Solanoideae (Atropina and Salpichroina), and many genera and some tribes unassigned to subfamily or within a more inclusive clade. Olmstead et al.’s (2008) work is the most comprehensive and complete for the family so far, with nearly 94% of the genera examined. The data produced an estimate of the phylogeny of Solanaceae with a number of well-supported clades (i.e. Goetzeoideae, Petunieae, Physalideae, Hyoscyameae) which accommodated many genera, and others with little resolution (i.e. basal clades in Solanoideae).

Olmstead et al.’s (2008) main modifications of the Hunziker (2001) classification are: 1) Nolana and Sclerophylax (in subfam. Solanoideae), Goetzea, Espadaea, Henoonia, Coeloneurum and Tsoala (in subfam. Goetzeoideae), and Duckeodendron (unassigned to a subfamily) are nested in Solanaceae; 2) reorganization of subfam. Solanoideae with 7 tribes also including re-accommodated genera (and other genera not assigned to any tribe or subtribe); 3) a much-reduced subfam. Cestroideae (3 tribes); 4) acceptance of subfam. Nicotianoideae including the Australian endemic genera (tribe Anthocercideae) + Nicotiana; 5) recognition of a reorganized tribe Physalideae with three subtribes (Withaniinae, Iochrominae and Physalidinae); 6) incorporation of Metternichia in subfam. Goetzeoideae; 7) acceptance of Petunia, Calibrachoa, Tzeltalia and Tsoala as valid genera; 8) inclusion of Cyphomandra, Lycopersicon, Normania and Triguera in Solanum. Olmstead et al. (2008) conclude: “Although significant advancements have been made in the understanding of relationships within Solanaceae, a complete phylogenetic classification has not been yet provided for the whole family. The classification we summarize below is the current framework for Solanaceae based on molecular evidence”.

Solanales, Solanaceae and Related Families. Within Lamiidae, Solanales are one of the four major clades together with Boraginales, Lamiales and Gentianales, among which there is no well-supported resolution (Soltis et al. 2011). Solanaceae are assigned to Solanales along with Convolvulaceae, Hydroleaceae, Montiniaceae and Sphenocleaceae (APG III 2009). These families comprise altogether ca. 165 genera and 4080 species. Solanaceae and Convolvulaceae are the larger families with an almost worldwide distribution. Authors differ in the dating of the stem group of Solanales. According to Wikström et al. (2001, Sphenoclea not included), it may date from the Campanian-Santonian 86–82 m.y.a., diversifying 78–76 m.y.a., while Janssens et al. (2009) date stem group Solanales to 101 m.y.a. and Bremer et al. (2004) to ca. 106 m.y.a.

The current topology of the tree for Solanales, obtained with a large number of genes (17) and more samples of Solanaceae and Convolvulaceae (Soltis et al. 2011), is congruent with prior molecular studies (Soltis et al. 2000; Bremer et al. 2002) but has stronger internal support. The order is characterized by the presence of O-methyl flavonols (flavones), inflorescence terminal, pollen tube usually with callose and persistent calyx in fruit (Stevens 2001 onwards).

Bremer (1996) and Soltis and Soltis (1997) found Montiniaceae, a family from Africa and Madagascar, sister to [Solanaceae + Convolvulaceae]. Soltis et al. (2000) reported strong support for the association of Montinia and Hydrolea (Sphenoclea was not included). Later, in a broader sampling in Montiniaceae, Bremer et al. (2002) provided stronger support, though just above 50%, for the association of Montiniaceae with Sphenoclea and Hydrolea. Finally, Soltis et al. (2011) found Montiniaceae are sister to Spenocleaceae + Hydroleaceae, and this well-supported clade (BS = 98%) in turn is sister to Convolvulaceae + Solanaceae (BS = 100%).

Classification of Solanaceae

  1. I.

    Subfam. Goetzeoideae Thorne & Reveal (2007).

    (Gen. 1–6).

  2. II.

    Subfam. Cestroideae Burnett (1835).

    1. 1.

      Tribe Cestreae Dumort. (1829).

      (Gen. 7–9).

    2. 2.

      Tribe Browallieae Hunz. (1995).

      (Gen. 10–11).

    3. 3.

      Tribe Salpiglossideae Benth. (1835).

      (Gen. 12–13).

    4. 4.

      Cestroideae: Incertae sedis:

      (Gen. 14–15: Protoschwenckia Soler., Heteranthia Nees & Mart.).

  3. III.

    Subfam. Nicotianoideae Miers (1848).

    1. 1.

      Tribe Anthocercideae G. Don (1838).

      (Gen. 16–22).

    2. 2.

      Tribe Nicotianeae G. Don

      (Gen. 23: Nicotiana L.).

  4. IV.

    Subfam. Solanoideae Burnett (1835).

    1. 1.

      Tribe Lycieae Lowe (1872).

      (Gen. 24).

    2. 2.

      Tribe Hyoscyameae Endl. (1839).

      (Gen. 25–31).

    3. 3.

      Tribe Juanulloeae Hunz. (1977).

      (Gen. 32–35).

    4. 4.

      Tribe Datureae Dumort. (1829).

      (Gen. 36–37).

    5. 5.

      Tribe Physalideae Miers (1849).

      1. a.

        Subtribe Withaniinae Bohs & Olmstead (2008).

        (Gen. 38–43).

      2. b.

        Subtribe Iochrominae Reveal (2012).

        (Gen. 44–49).

      3. c.

        Subtribe Physalidinae Reveal (2012).

        (Gen. 50–57).

      4. d.

        Physalideae: Incertae sedis:

        (Gen. 58–63: Capsicophysalis Averett & M. Martínez, Cuatresia Hunz., Larnax Miers, Deprea Raf., Schraderanthus Averett, Tzeltalia E. Estrada & M. Martínez).

    6. 6.

      Tribe Capsiceae Dumort. (1827).

      (Gen. 64–65).

    7. 7.

      Tribe Solaneae Dumort. (1829).

      (Gen. 66–67).

    8. 8.

      Solanoideae: Incertae sedis:

      (Gen. 68–80: Latua Phil., Jaborosa Juss., Sclerophylax Miers, Nolana L.f., Exodeconus Raf., Nicandra Adans., Schultesianthus Hunz., Trianaea Planch. & Linden, Solandra Sw., Mandragora L., Nectouxia Kunth, Salpichroa Miers, Darcyanthus Hunz.).

  5. V.

    Solanaceae: Incertae sedis:

    (Gen. 81–96: Genera not assigned to a subfamily: Schizanthus Ruiz & Pav., Duckeodendron Kuhlmann, Tribes not assigned to a subfamily: Tribe Benthamielleae Hunz. (2000): Pantacantha Speg., Combera Sandw., Benthamiella Speg., Tribe Petunieae Horan. (1847): Fabiana Ruiz & Pav., Calibrachoa La Llave & Lex., Petunia Juss., Brunfelsia L., Leptoglossis Benth., Nierembergia Ruiz & Pav., Bouchetia Dunal, Hunzikeria D’Arcy, Plowmania Hunz. & Subils, Tribe Schwenckieae Hunz. (1977): Schwenckia L., Melananthus Walp.).

Key to the Genera of Solanaceae

  1. 1.

    Fruit a drupe, 1-seeded; embryo U-shaped; endosperm oily; corolla aestivation quincuncial; trees up to 30 m high. Brazil, Central Amazonia 82. Duckeodendron

  • Fruit a berry, capsule, diclesium, or schizocarp, few- to many-seeded, rarely 1-seeded; embryo straight or slightly curved, vermiform, annular or coiled; endosperm mostly starchy or missing; corolla aestivation various; herbs, shrubs or small trees 2

  1. 2.

    Leaves always entire and coriaceous; corolla orange; fruit fleshy, yellow or orange, mesocarp gelatinous at maturity; embryo straight; endosperm missing 3

  • Leaves entire or variously lobed or divided, membranaceous, less commonly coriaceous; corolla variously coloured; fruit fleshy or dry, mesocarp rarely gelatinous at maturity; embryo of various shape; endosperm scanty or abundant 6

  1. 3.

    Corolla funnel-shaped, cup-shaped or tubular-curved, > 10 mm long; ovary bilocular 4

  • Corolla campanulate or shortly tubular, < 10 mm long; ovary unilocular 5

  1. 4.

    Corolla funnel-shaped or cup-shaped; calyx and corolla hexamerous; stamens 6, equal. Greater Antilles 6. Goetzea

  • Corolla tubular-curved; calyx and corolla pentamerous (rarely 4- or 6-merous); stamens 5, unequal. Cuba 5. Espadaea

  1. 5.

    Corolla campanulate, lobed nearly to base; style pubescent; stamens inserted at base of corolla. Greater Antilles 3. Coeloneurum

  • Corolla shortly tubular; style glabrous; stamens inserted at middle of corolla. Cuba 4. Henoonia

  1. 6.

    Aestivation valvate, valvate-induplicate, valvate-plicate, valvate-conduplicate, cochlear, cochlear-conduplicate, cochlear-plicate, contorted-induplicate, contorted-conduplicate, or reciprocative; corolla actinomorphic or zygomorphic; anthers mostly dithecal; endosperm starchy; embryo variously curved or almost straight, cotyledons usually up to 2.5–3 times shorter, equal, slightly longer or frequently much longer than rest of embryo 7

  • Aestivation valvate-supervolute; corolla actinomorphic; anthers mostly monothecal (dithecal in Anthocercis and Symonanthus); endosperm oily and starchy (starchy only in Duboisia); embryo straight, cotyledons very short (ca. 5–12 times shorter than the rest of the embryo) 92

  1. 7.

    Placentation parietal; style strongly exserted and pubescent at the apex; stigma capitate, very thick. Madagascar 2. Tsoala

  • Placentation mostly axile (basal in Melananthus; apical in Sclerophylax); style mostly included, glabrous or pubescent in its basal half; stigma, when capitate, thin or thick 8

  1. 8.

    Aestivation valvate-induplicate (valvate also in some Jaltomata spp.); pedicels articulated 9

  • Aestivation valvate, valvate-plicate, valvate-conduplicate, cochlear, cochlear-conduplicate, cochlear-plicate, contorted-induplicate, contorted-conduplicate, or reciprocative; pedicels not articulated (articulated in some Solanum spp.) 12

  1. 9.

    Ovary with an annular persistent corolla remaining at its base (cyathium); anthers dorsifixed, embryo straight or almost curved 10

  • Ovary without cyathium; anthers ventrifixed, embryo strongly curved. Americas 66. Jaltomata

  1. 10.

    Seeds long and thin (up to 10 times longer than broad), compressed, winged, imbricate in the fruit; capsules few-seeded (1–8 seeds). South America and Haiti 8. Sessea

  • Seeds slightly longer than broad, thick, subpolyhedrical, wingless, adjacent in the fruit; few-seeded berry or many-seeded capsule 11

  1. 11.

    Anthers included; ovary stipitate, ovules few (4–20, exceptionally more); berry few-seeded. Mexico, Central America, the Antilles, South America 7. Cestrum

  • Anthers exserted; ovary sessile, ovules many (more than 50); capsules many-seeded. Chile 9. Vestia

  1. 12.

    Seeds not compressed and angular, mostly prismatic, polyhedrical, and cuboidal, more rarely subglobose, subreniform, or long, thin and very narrow (in Benthamiella and Combera seeds slightly compressed) 13

  • Seeds compressed and not angular, mostly discoid or reniform 35

  1. 13.

    Stamen fertile 4 or 2 14

  • Stamens fertile 5 (rarely also 3, 2 or 1 in Benthamiella) 25

  1. 14.

    Anthers basifixed 15

  • Anthers ventrifixed or dorsifixed 17

  1. 15.

    Aestivation cochlear; anthers monothecic; style smooth; pollen grains triporate or 3–4-colpate, rarely 3-colporate. Central and South America 89. Brunfelsia

  • Aestivation reciprocative or cochlear-conduplicate; anthers dithecic, with partial or complete abortion of one theca in the upper stamen pair; style strongly corrugated; pollen grains 4–5/9-colporate 16

  1. 16.

    Herbs 0.08–1(1.5) m high; corolla hypocrateriform, ventricose, not twisted, aestivation reciprocative; stamens inserted in upper half of corolla, lower posterior pair with filaments usually two times longer than anthers, upper lateral pair with compressed filaments shorter than anther. Americas 10. Browallia

  • Shrubs 1.5–2 m high; corolla infundibuliform, not ventricose, its lower part spirally twisted, aestivation cochlear-conduplicate; stamens inserted in both halves of the corolla, lower posterior pair with filaments 4–5 times longer than anthers, upper lateral pair with terete filaments as long as anthers or somewhat longer. Colombia, Ecuador, Peru 11. Streptosolen

  1. 17.

    Anthers dorsifixed; nectary cushion-shaped. Bolivia, Brazil 14. Protoschwenckia

  • Anthers ventrifixed; nectary absent or when present annular, sheath-like or invaginate-bilobed 18

  1. 18.

    Nectary absent 19

  • Nectary always present 20

  1. 19.

    Corolla infundibuliform, slightly bilabiate; stamens inserted at lower half of corolla tube and not connivent on style; thecae unequal; oil-secreting glandular trichomes inside the corolla missing. Brazil 15. Heteranthia

  • Corolla hypocrateriform; stamens inserted at apex of corolla tube and generally connivent on style; thecae equal; oil-secreting glandular trichomes inside the corolla present. Mexico, South America 91. Nierembergia

  1. 20.

    Corolla zygomorphic, limb with five undivided lobes 21

  • Corolla actinomorphic or zygomorphic, limb with five trilobulate or trifid lobes 24

  1. 21.

    Style cylindrical, not broadened distally, lacking lateral expansions; stigma laminar; nectary sheath-like and bilobed; pollen grains in monads or tetrads. Americas 92. Bouchetia

  • Style broadened distally, with 2 lateral expansions or wings; stigma inconspicuous, linear; nectary annular and prominent or invaginate-bilobed; pollen grains in monads 22

  1. 22.

    Calyx campanulate; corolla 42–50 mm long, funnel-shaped, tube comparatively wide; fertile stamens 4 plus one staminode, thecae distinct; nectary annular; shrubs. Mexico, Guatemala 94. Plowmania

  • Calyx not campanulate; corolla 9–30 mm long, cylindrical or salver-form, tube narrow; fertile stamens 4 or 2 plus 2–3 staminodes, thecae confluent; nectary invaginate-bilobed; small herbaceous xerophytic perennials or ephemerals 23

  1. 23.

    Corolla ventricose, limb narrow (3–6 mm); calyx cylindrical; fruits fusiform; seeds 0.3–0.5 mm long. Peru, Chile, Argentina 90. Leptoglossis

  • Corolla salver-form, limb broad (10–23 mm); calyx funnel-shaped, campanulate or obconical; fruits almost subglobose; seeds 1.2–1.5 mm long. USA, Mexico, Venezuela 93. Hunzikeria

  1. 24.

    Placentation axile; nectary invaginate-pelviform; aestivation valvate-conduplicate; capsules bilocular, with 6–70 seeds; seeds cuboidal, 0.6–0.8 mm long, testa reticulate. Central and South America 95. Schwenckia

  • Placentation basal; nectary invaginate-pelviform or slightly bilobulate; aestivation valvate-induplicate; capsules unilocular, 1-seeded; seeds elongated, 1–1.5 mm long, testa smooth. Americas 96. Melananthus

  1. 25.

    Fruit a berry or long capsule (35–45 mm long); pollen grains echinate; shrubs or small trees 26

  • Fruit a short capsule (3–28 mm long); pollen grains not echinate; usually herbs or small shrubs 27

  1. 26.

    Fruit a thin capsule; seeds very long (30–39 x 2–3.5 mm); embryo straight, endosperm scanty; corolla aestivation contorted-conduplicate. Brazil 1. Metternichia

  • Fruit a globose berry; seeds slightly longer than broad, ca. 2 mm diam.; embryo slightly curved, endosperm abundant; corolla aestivation cochlear-conduplicate. Chile 68. Latua

  1. 27.

    Flowers usually subtended by two opposite bracteoles similar to the leaves; capsules few-seeded (1–5 or up to 7 seeds) 28

  • Flowers without a pair of opposite bracteoles; capsules many-seeded 30

  1. 28.

    Herbs; leaves petiolate, sheathless, spathulate, oblong-spathulate, deltoid-ovate or rhomboide-ovate. Argentina, Chile 84. Combera

  • Plants woody; leaves sessile, sheathed, linear, narrowly elliptic to obovate or linear-subulate 29

  1. 29.

    Low shrubs; leaves not densely imbricate; leaves and calyx teeth rigid and spiny; corolla aestivation valvate-induplicate; seed surface ridged-winged. Argentina 83. Pantacantha

  • Chamaephytes or lax cushions; leaves densely imbricate; leaves and calyx teeth neither rigid nor spiny; corolla aestivation contorted-conduplicate; seed surface foveolate or reticulate. Argentina, Chile 85. Benthamiella

  1. 30.

    Nectary absent; stamens attached to apex of corolla tube and generally connivent on style; oil-secreting glandular trichomes inside the corolla present. Mexico and South America 91. Nierembergia

  • Nectary always present; stamens attached to lower half of corolla tube (in some Nicotiana spp. near apex of corolla tube) and not connivent on the style; oil-secreting glandular trichomes missing 31

  1. 31.

    Anthers dorsifixed; aestivation contorted-conduplicate 32

  • Anthers ventrifixed; aestivation cochlear or reciprocative 33

  1. 32.

    Flowers in many-flowered terminal paniculoid inflorescences; glandular trichomes generally with multicellular head (rarely head unicellular), branched trichomes frequent; stamens equal or in two (4 + 1) or three groups (2 + 2 + 1); seeds ca. 100–5000. Americas, Australia, Africa 23. Nicotiana

  • Flowers solitary or arising from the axil of one leaf; glandular trichomes with unicellular head, branched trichomes absent; stamens equal (only in 2 species) or in two groups (2 + 3); seeds ca. 30–52. Peru, Bolivia, Chile, Argentina 86. Fabiana

  1. 33.

    Fertile stamens 5 (2 + 2+ 1); nectary sheathed-bilobed; stigma laminar; pollen grains in monads or tetrads. Americas 92. Bouchetia

  • Fertile stamens 5 (2 + 2+ 1, rarely 2 + 3); nectary bilobed; stigma capitate, discoid or bilobate; pollen grains always in monads 34

  1. 34.

    Corolla aestivation reciprocative: the anterior induplicative lobe covers the four others which are conduplicate; calyx usually lobed halfway, lobes narrowed towards apex; seeds with straight anticlinal walls. Americas 87. Calibrachoa

  • Corolla aestivation cochlear; calyx usually deeply lobed, lobes linear or enlarged towards apex; seeds with undulating anticlinal walls. Southern South America 88. Petunia

  1. 35.

    Woody epiphytes or occasionally shrubs or trees; embryo almost straight or slightly curved (annular in Schultesianthus), cotyledons accumbent or oblique; endosperm comparatively scanty 36

  • Herbs, shrubs or trees (rarely vines in Solanum); embryo annular, coiled or subcoiled, cotyledons incumbent; endosperm usually abundant 41

  1. 36.

    Aestivation valvate 37

  • Aestivation cochlear 38

  1. 37.

    Anthers basifixed; flowers solitary on brachyblasts; cotyledons oblique, shorter and not broader than rest of embryo. Brazil 32. Dyssochroma

  • Anthers dorsifixed; inflorescences on long peduncles; cotyledons accumbent, as long as (or somewhat shorter or longer) but wider than rest of embryo. Mexico to northern South America 35. Merinthopodium

  1. 38.

    Anthers ventrifixed; ovary (4) 5–10-carpellate. Colombia to Peru 75. Trianaea

  • Anthers dorsi- or basifixed; ovary 2-carpellate 39

  1. 39.

    Flowers zygomorphic, fragrant; calyx thick and coriaceous; gynoecium and androecium usually bent; seeds 5–5.5 mm long; embryo strongly annular. Mexico to Bolivia 74. Schultesianthus

  • Flowers actinomorphic, not fragrant; calyx membranous; gynoecium and androecium straight; seeds 2.2–3.8 mm long; embryo usually slightly curved 40

  1. 40.

    Anthers dorsifixed; corolla thick and fleshy, tubular. Mexico to Bolivia 33. Juanulloa

  • Anthers basifixed; corolla membranous, funnel-form or infundibuliform. Panama to Peru 34. Markea

  1. 41.

    Fertile stamens five; corolla aestivation cochlear, valvate, valvate-conduplicate or contorted-conduplicate; berry frequently with stone cells, rarely with pyrenes, or fruit a capsule, diclesium or schizocarp 42

  • Fertile stamens four or two; corolla aestivation cochlear; fruit a capsule 90

  1. 42.

    Fruit mostly a circumscissile capsule (pyxidium) 43

  • Fruit a berry or a non-circumscissile capsule, or diclesium or schizocarp; gynoecium 2–5-carpellate (in Nolana up to 30-carpellate) 48

  1. 43.

    Filaments 1.5–4 times longer than anthers, inserted at base of corolla or further up (in some Hyoscyamus in upper half of corolla but not near its upper margin); stamens equal or unequal 44

  • Filaments shorter than anthers, inserted near upper margin of corolla; stamens equal. China 26. Przewalskia

  1. 44.

    Flowers solitary (in Hyoscyamus solitary at base and in condensed cymes above); fruiting calyces inflated or not 45

  • Inflorescences paniculate; fruiting calyces inflated. Asia 28. Physochlaina

  1. 45.

    Flowers pendant on long pedicels; stamens included, filaments inserted at base of corolla; nectary annular, prominent; lobes of fruiting calyx never spine-tipped 46

  • Flowers sessile or very shortly pedicellate; stamens usually exserted, filaments inserted at middle of corolla, at a lower level or further up; nectary absent or cryptic; fruiting calyx rigid, lobes spine-tipped. Europe, Asia, Africa 25. Hyoscyamus

  1. 46.

    Corolla actinomorphic; filaments straight; calyx lobes usually unequal in size 47

  • Corolla zygomorphic (lobes unequal); filaments circinate in bud, curved at anthesis; calyx lobes equal. China 29. Atropanthe

  1. 47.

    Corolla and calyx usually almost of the same length; stamens equal, anthers 5–7 mm. Asia 30. Anisodus

  • Corolla ca. two times longer than calyx; stamens slightly unequal, anthers 2.5–4 mm. Europe and Asia 27. Scopolia

  1. 48.

    Gynoecium (3–4)5-carpellate or up to 30-carpellate 49

  • Gynoecium bicarpellate (two species of Jaborosa with 2–5 carpels) 50

  1. 49.

    Berry enclosed by accrescent, stiff and inflated calyx; flowers actinomorphic; calyx segments auriculate; filaments equal, geniculate at their expanded base; carpels 3–5; annual herbs or subshrubs. Peru to Argentina 73. Nicandra

  • Mericarps fused or free from each other; flowers commonly zygomorphic; calyx segments not auriculate; filaments unequal (3 + 2), not geniculate nor expanded basally; carpels 2–30; annuals or perennial herbs or weak shrubs. Chile, Peru, Galapagos Islands 71. Nolana

  1. 50.

    Nectary with two opposite lobes; pollen grains cryptoaperturate, exine echinate. Europe, Asia 77. Mandragora

  • Nectary, when present, without lobes; pollen grains usually aperturate (a few species of dioecious Solanum with inaperturate monads), exine not echinate 51

  1. 51.

    Filaments straight 52

  • Filaments declinate, or incurved towards centre of flower 86

  1. 52.

    Ovary with cyathium; corolla aestivation cochlear or cochlear-plicate. Cosmopolitan 24. Lycium

  • Ovary without cyathium; corolla aestivation valvate, valvate-plicate, valvate-conduplicate, contorted-conduplicate, contorted-induplicate or contorted-plicate 53

  1. 53.

    Corolla aestivation contorted-conduplicate; anthers basifixed; endosperm with protein and/or oil, lacking starch 54

  • Corolla aestivation valvate, valvate-plicate, valvate-conduplicate, contorted-induplicate or contorted-plicate; anthers mostly dorsifixed or dorsi-basifixed, more rarely ventrifixed or basifixed; endosperm starchy 56

  1. 54.

    Fruit a many-seeded capsule or berry; calyx neither accrescent nor thorny; stamens equal; placenta axile; many ovules per locule; embryo coiled; herbs, shrubs or small trees 55

  • Fruit a diclesium, generally 2-seeded; calyx accrescent, lobes becoming thorny; stamens unequal; placenta apical; ovules 1–2 per locule; embryo straight or slightly curved; annual or perennial herbs. Argentina, Paraguay, Uruguay 70. Sclerophylax

  1. 55.

    Annual herbs or short-lived perennials; branching mostly dichasial; flowers erect; calyx circumscissile forming a reflexed annulus subtending the fruit; ovary partially inferior, tetralocular in lower half, nectary invaginate; capsule dehiscent, mostly with spines or tubercles; seeds with or without a funicular elaiosome. Americas 37. Datura

  • Shrubs or small trees; branching mostly monochasial; flowers pendant or inclined; calyx not circumscissile, falling off completely or persisting around the fruit; ovary superior, bilocular throughout, nectary annular; berry pendant, unarmed; seeds lacking an elaiosome. South America 36. Brugmansia

  1. 56.

    Ovary with basal nectary; filaments usually inserted individually on corolla; anthers opening by longitudinal slits 57

  • Ovary without nectary; filaments basally united into a uniform and continuous ring fused at base of corolla; anthers strictly poricidal or opening by terminal pores which sometimes expand into longitudinal slits 85

  1. 57.

    Anthers extrorse, ventrifixed 58

  • Anthers introrse, dorsifixed, basifixed or dorsi-basifixed 59

  1. 58.

    Corolla usually glabrous within; calyx usually entire at anthesis; pericarp lacking stone cells. Guatemala to Bolivia 58. Cuatresia

  • Corolla usually with tufts of dendroid trichomes within; calyx sometimes 5-apiculate, shortly dentate or more frequently divided; pericarp with few stone cells. Europe, Africa, Asia 43. Withania

  1. 59.

    Filaments minute, anthers sessile or subsessile; perianth tetramerous; ovary half-inferior. Hawaii 39. Nothocestrum

  • Filaments at least half as long as anthers; perianth usually pentamerous; ovary superior 60

  1. 60.

    Calyx entire or the five main nerves overtopping the smooth margin as cylindrical or dentate appendages 61

  • Calyx 5-divided 63

  1. 61.

    Corolla generally with an inside ring of non-glandular trichomes above the filament insertion; pericarp usually with stone cells. Mexico to Bolivia 50. Witheringia

  • Corolla glabrous inside or with some tiny glandular trichomes; pericarp lacking stone cells (few in one Capsicum species) 62

  1. 62.

    Stapet auriculate; calyx frequently with five or ten appendages or teeth prolonging the nerves; corolla aestivation valvate-plicate; placenta and endocarp never fuse. Southern USA to South America 65. Capsicum

  • Stapet not auriculate; calyx without appendages, lacking nerve prolongations; corolla aestivation valvate; placenta centrifugally enlarged and fused with endocarp. Asia 40. Tubocapsicum

  1. 63.

    Corolla aestivation valvate 64

  • Corolla aestivation valvate-induplicate, valvate-plicate or valvate-conduplicate 70

  1. 64.

    Corolla lobes short, tube always longer 65

  • Corolla lobes as long as tube or slightly longer 67

  1. 65.

    Corolla urceolate-campanulate or campanulate-infundibuliform; anthers dorsifixed; calyx not accrescent; pericarp lacking stone cells 66

  • Corolla generally infundibuliform; anthers basifixed; calyx accrescent, slightly or much-inflated, completely surrounding the berry, stone cells frequent in pericarp. Venezuela to Bolivia 60. Deprea

  1. 66.

    Corolla urceolate-campanulate; stamens included; style and anthers pubescent. Africa 38. Discopodium

  • Corolla campanulate-infundibuliform; stamens exserted; style and anthers glabrous. Mexico to South America 49. Acnistus

  1. 67.

    Corolla with a dense ring of trichomes inside; stapet not auriculate, filaments hairy, inserted near middle of corolla. Mexico and Central America 51. Brachistus

  • Corolla glabrous inside (glabrescent in some Larnax species); stapet auriculate, filaments glabrous, inserted at base of corolla 68

  1. 68.

    Herbs or shrubs; calyx membranous, strongly accrescent, tightly enveloping the berry; styles long, always of the same length; filaments and anthers frequently unequal. Central America to Peru 59. Larnax

  • Shrubs or trees, calyx rather thick, not accrescent or, if accrescent, surrounding the berry completely or partly, but always open upwards; styles heteromorphic; filaments and anthers equal 69

  1. 69.

    Shrubs; calyx almost dialisepalous (tube very short or sometimes almost absent), strongly accrescent. Brazil 41. Athenaea

  • Shrubs or small trees (up to 8 m high); calyx with evident tube, not accrescent. Brazil, Paraguay, Argentina, Peru, Bolivia 42. Aureliana

  1. 70.

    Corolla infundibuliform or tubular-infundibuliform; fruiting calyx not inflated 71

  • Corolla rotate or campanulate-rotate; fruiting calyx inflated or not 75

  1. 71.

    Herbs, generally small to very small (3–20 cm high); stamens unequal 72

  • Shrubs or trees (up to 10 m, or more); stamens usually equal 73

  1. 72.

    Calyx with equal lobes; corolla 20–40 mm long, 5-angulate; berry lignified at apex or sometimes with stone cells; seeds wingless. Ecuador, Peru, Chile, Argentina 72. Exodeconus

  • Calyx with unequal lobes; corolla 5–8 mm long, with well-developed lobules; fruit a capsule; seeds with a hyaline peripheral wing. North America 54. Oryctes

  1. 73.

    Stapet large, with conspicuous appendages and lateral wings; style subterminal. Colombia to Argentina 44. Dunalia

  • Stapet not prominent, lacking apical appendages or lateral wings; style terminal 74

  1. 74.

    Young stems densely hairy with deciduous brown specialized trichomes (thickened, multiseriate, bifurcate or not); filaments inserted at base of corolla; berry with pyrenes, pericarp purple to shining black; calyx not accrescent. Venezuela to Bolivia 46. Saracha

  • Young stems with simple trichomes; filaments inserted between 1/4 and 3/4 of corolla length; berry usually with many stone cells, pericarp yellowish to orange or red, sometimes purple at apex; calyx accrescent. South America 45. Iochroma

  1. 75.

    Corolla inside usually with a ring of trichomes above; filaments longer than anthers; stapet not auriculate and not with incipient auricles 76

  • Corolla glabrous inside; filaments more or less as long as or slightly longer than anthers; stapet auriculate 84

  1. 76.

    Herbs or weak shrubs; stapet not auriculate; calyx strongly accrescent; style subterminal 77

  • Small trees; stapet with incipient auricles; calyx slightly accrescent; style terminal. Argentina, Bolivia 48. Eriolarynx

  1. 77.

    Fruiting calyx inflated, urceolate, angled and strongly veined (not angled in Calliphysalis), completely closed or with a small apical opening, hiding the berry 78

  • Fruiting calyx not inflated, neither angled nor strongly veined, reflexed and not enclosing the berry or enclosing it only partly 81

  1. 78.

    Fruiting calyx coriaceous, partly open at mouth; soft-stemmed reclining shrubs or subshrubs. Mexico 62. Tzeltalia

  • Fruiting calyx papery, completely closed; herbs, rarely shrubs 79

  1. 79.

    Aerial parts lacking vesiculose water-containing trichomes; flowers solitary or 2–6; corolla usually campanulate-rotate or campanulate, pericarp of uniform thickness, never disintegrating at maturity 80

  • Aerial parts with a granular aspect, covered almost completely by vesiculose, whitish, water-containing trichomes; flowers 2–4(5); corolla rotate; pericarp unevenly thickened, disintegrating almost completely in thinnest and weakest parts. USA, Mexico 55. Quincula

  1. 80.

    Flowers solitary; fruiting calyx angled, invaginated at base; rhizomatous perennial herbs or annual herbs. Mostly Americas 57. Physalis

  • Fascicles of 2–6 flowers; fruiting calyx rounded, not invaginated at base; taprooted perennial herbs. USA 53. Calliphysalis

  1. 81.

    Inflorescence fasciculate (6–8 flowers) from axils; corolla lobed to divided, 4–5 cm wide; fruiting calyx reflexed under berry at maturity. Mexico, Guatemala, Venezuela 61. Schraderanthus

  • Flowers 1 or 2 per axil; corolla little or slightly lobed, not divided, 2–4 cm wide; fruiting calyx reflexed or closely appressed and partially or completely enclosing berry at maturity 82

  1. 82.

    Corolla rotate-campanulate, irregular; berry red; fruiting calyx red, reflexed under the berry at maturity; testa tuberculate; erect annual herbs 63. Capsicophysalis

  • Corolla rotate, actinomorphic; berry green; fruiting calyx green, closely appressed and partially or completely enclosing the berry at maturity; testa rugose-reticulate; erect annual or ascending or spreading perennial herbs 83

  1. 83.

    Corolla 8–9 mm long, aestivation contorted-plicate; pericarp generally dry, lacking stone cells. USA, Mexico 56. Chamaesaracha

  • Corolla 20–24 mm long, aestivation valvate-plicate; pericarp generally juicy, with stone cells. North America 52. Leucophysalis

  1. 84.

    Shrubs or small trees; calyx not accrescent (or very slightly accrescent), never surpassing half of the berry; corolla 5-lobed. Southern South America 47. Vassobia

  • Herbs; calyx much accrescent, tightly enveloping berry; corolla 5-angulate, lobes inconspicuous. Peru, Bolivia 80. Darcyanthus

  1. 85.

    Calyx entire, truncate, with five to ten linear nerve prolongations; anthers strictly poricidal at tips; berry juicy, exceptionally with pyrenes. Americas, Asia 64. Lycianthes

  • Calyx 5-parted into lobes or segments; anthers poricidal or with a terminal pore expanding into a longitudinal slit; berry juicy or nearly dry, never with pyrenes. Cosmopolitan 67. Solanum

  1. 86.

    Corolla zygomorphic or slightly zygomorphic; aestivation cochlear; filaments declinate 87

  • Corolla actinomorphic; aestivation valvate, valvate-induplicate or valvate-conduplicate; filaments incurved towards the centre of the flower or curved at least at the apex 88

  1. 87.

    Corolla 10–37 cm long; stamens equal, anthers basifixed; ovary partly inferior tetralocular; scandent shrubs or lianas. Mexico to Brazil 76. Solandra

  • Corolla 1.9–3.5 cm long; stamens unequal, anthers dorsifixed; ovary superior bilocular; perennial herbs. Europe, Asia, Africa 31. Atropa

  1. 88.

    Leaves dentate to pinnatisect, not darkening when dry; flowers 1–7 per axil, or in dense glomerules or fascicles; calyx tube well-developed; ovary 2–5-carpellate; berry globose-depressed; embryo annular, endosperm copious. Peru to Argentina 69. Jaborosa

  • Leaves entire, darkening when dry; flowers solitary; calyx tube absent or very short; ovary bicarpellate; berry elongated; embryo coiled, endosperm scanty 89

  1. 89.

    Corolla lacking a corona; distal end of filaments broadened or not, lacking a laminar enlargement on the dorsal side of anthers; scandent shrubs, occasionally herbs. Venezuela to Argentina 79. Salpichroa

  • Corolla with a corona; distal end of filaments broadened into a laminar enlargement on the dorsal side of anthers; herbs. Mexico 78. Nectouxia

  1. 90.

    Lateral stamens aborted or with smaller anthers, posterior pair always fertile or with larger anthers; flowers not resupinate 91

  • Lateral stamens fertile, posterior pair aborted; flowers resupinate. Chile, Argentina 81. Schizanthus

  1. 91.

    Corolla 2.3–5 cm long; anthers dorsifixed, 2–4 mm long. Chile, Argentina 12. Salpiglossis

  • Corolla 0.6–1.3(–1.5) cm long; anthers basifixed, 0.8–1 mm long. Chile, Argentina 13. Reyesia

  1. 92.

    Anthers dithecal; plants dioecious or hermaphrodite 93

  • Anthers monothecal; plants hermaphrodite 94

  1. 93.

    Plants hermaphrodite; stamens 4, didynamous or subequal; corolla lobes long and narrow, usually longer or sometimes equal or slightly shorter than tube. Australia 17. Anthocercis

  • Plants dioecious; stamens (3) 4 or 5, equal; corolla lobes as long as broad, four to six times shorter than tube. Australia 16. Symonanthus

  1. 94.

    Fruit a capsule; endosperm oily and starchy; plants usually with dense indumentum of dendritic trichomes 95

  • Fruit aberry; endosperm starchy; plants glabrous or glabrate. Australia, New Caledonia 22. Duboisia

  1. 95.

    Leafy shrubs; corolla 5–22 mm long 96

  • Shrubs appearing leafless, the small leaves early-deciduous; corolla 2.5–4 mm long; stamens 4 (2 + 2) and a sterile staminode usually present. Australia 21. Crenidium

  1. 96.

    Stamens 5 97

  • Stamens 4, didynamous (sometimes with a staminode present). Australia 19. Cyphanthera

  1. 97.

    Stamens unequal (2 long, 3 short); corolla lobes long and narrow, glabrous inside. Australia 18. Grammosolen

  • Stamens equal; corolla lobes broad and short, densely pubescent inside. Australia 20. Anthotroche

  1. I.

    Subfam. Goetzeoideae Thorne & Reveal (2007).

Trees to shrubs; pollen tricolpate, exine echinate; fruit a capsule pauciseminate or a berry pauciseminate; embryo straight, cotyledons fleshy mostly large; endosperm at most slight.

  1. 1.

    Metternichia J.C. Mikan Fig. 61

    Fig. 61.
    figure 6

    Solanaceae. Metternichia principis. A Flowering branch. B Flower. C Opened corolla. D Half an anther showing the curved apex of the filament and its basal insertion. E Anther, ventral view. F Ovary. G Stigma, commissural view. H Fruit. I Seed. J Embryo, front view. K Embryo, lateral view. (Hunziker 2001; drawn by L. Sánchez)

    Full size image

    Metternichia J.C. Mikan, Del. fl. faun. bras. 3, tab. 13 (1823).

Trees. Leaves entire. Flowers 5-merous, solitary or in small fascicles, bracts deciduous; calyx campanulate, zygomorphic; corolla strongly scented, white, long, actinomorphic, infundibuliform, lobes short and wide, aestivation contorted-conduplicate; stamens unequal, 2 + 2 + 1 or 2 + 3, filaments declinate; anthers basifixed; ovary bilocular, ovules 10–16, anatropous; nectary annular. Capsule subcylindric-fusiform, septicidal-loculicidal, 4-valved. Seeds 4–5 per locule, erect, long, very narrow and thin, triquetrous, each angle with a narrow wing, hilum near the base of the ventral face; embryo straight, cotyledons compressed, not wider than the slightly longer radicle; endosperm scanty. 2n = 26.

A single species from Brazil (Rio de Janeiro state, Bahia, Minas Gerais): Metternichia principis J.C. Mikan.

  1. 2.

    Tsoala Bosser & D’Arcy

    Tsoala Bosser & D’Arcy, Bull. Mus. Natl. Hist. Nat., B, Adansonia 14: 8 (1992).

Shrubs or small trees, much branched; young stems and leaves with dense reddish pubescence. Leaves deciduous, entire, petiole canaliculate. Flowers terminal, solitary, 5-merous; calyx lobes subequal, triangular; corolla greenish, hypocrateriform, entirely glabrous, tube thin and very long (up to 15 cm), limb infundibuliform, lobes apiculate; stamens strongly exserted, subequal, filaments glabrous, inserted near apex of tube; anthers basifixed; ovary unilocular or partly bilocular, placentation parietal; nectary annular; style longer than stamens, pubescent at apex, stigma thick, capitate. Capsule apiculate, pericarp green, thin and fragile. Seeds 4–9; embryo thick, straight, cotyledons very short; endosperm abundant.

Monotypic. Tsoala tubiflora Bosser & D’Arcy, Madagascar.

  1. 3.

    Coeloneurum Radlk.

    Coeloneurum Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 19: 280 (1890); Liogier, La Flora de la Española VI, vol. 70, ser. Científica 27: 361 (1994).

Shrubs or small trees; stems with dense rusty pubescence. Leaves petiolate, entire, glabrous, leathery. Flowers extra-axillary, solitary, sessile or subsessile; calyx broadly campanulate, shortly 5-lobed, rusty pubescent; corolla campanulate, yellowish or cream, lobed nearly to base, lobes oblong; stamens 5(4), filaments glabrous, inserted basally; ovary unilocular; style sometimes absent, pubescent at base, stigma bilobed. Berry globose, yellow or orange, 1-seeded. Seed large; embryo straight, cotyledons fleshy, thick, radicle tiny; endosperm very scanty.

A single species, C. ferrugineum Urb., Hispaniola (Greater Antilles).

  1. 4.

    Henoonia Griseb.

    Henoonia Griseb., Cat. pl. Cub.: 166 (1866); Fuentes Fiallo, Fl. República de Cuba 10(4): 2–14 (2005), rev.

Shrubs; branches greyish, sometimes lateral branches spinescent. Leaves subsessile, leathery, entire, base cordate or wedge-shaped, margins revolute, glabrous and brilliant adaxially, pubescent abaxially. Flowers axillary, solitary or in fascicles, subsessile; calyx cupola-shaped, shortly 5-lobed, pubescent; corolla shortly tubular, deeply lobed, lobes reflexed; stamens 5, filaments glabrous, inserted at middle of corolla tube; anthers dorsifixed; ovary unilocular, uniovulate; style glabrous. Berry globose-apiculate, 1-seeded. Seeds up to 8 mm, broadly ovoid; embryo straight, cotyledons fleshy, thick; endosperm very scanty.

A single species, H. myrtifolia Griseb., Cuba.

  1. 5.

    Espadaea A. Rich.

    Espadaea A. Rich., Hist. phys. Cuba, Pl. vasc. 11: 147 (1850); Fuentes, Revista Jard. Bot. Nac. Cuba 3: 51–67 (1982); Fuentes, Fl. República Cuba 10(4): 2–14 (2005), rev.

Trees; stems greyish, rusty pubescent when young. Leaves petiolate, entire, leathery, glabrous and brilliant adaxially, pubescent abaxially. Flowers axillary, solitary, subsessile, (4)5(6)-merous; calyx cup-shaped, shortly dentate; corolla yellow, funnel-shaped, slightly curved, lobes reflexed; stamens 5, unequal, glabrous, strongly exserted, inserted basally; anthers basifixed; ovary tomentose, bilocular, each locule uniovulate; style glabrous. Berry globose-apiculate, orange or yellow, 1–2-seeded. Seeds up to 7.5 mm, broadly ovoid or dorsally flattened; embryo straight, cotyledons fleshy, thick, radicle tiny; endosperm very scanty. 2n = 48.

A single species, Espadaea amoena A. Rich., Cuba. Planted as an ornamental.

  1. 6.

    Goetzea Wydl.

    Goetzea Wydl., Linnaea 5: 423 (1830); Liogier, Descriptive Flora of Puerto Rico and Adjacent Islands IV: 444 (1995).

Trees or shrubs. Leaves shortly petiolate, entire, leathery, broad. Flowers axillary, solitary, long-pedicellate, pendant; calyx campanulate or cup-shaped, 6-lobed; corolla yellow, conspicuously funnel-shaped, 6-lobed, lobes reflexed; stamens 6, filaments glabrous, exserted, inserted basally; anthers pubescent; ovary bilocular, pubescent, ovules few, pendant; style glabrous, stigma bi- or trilobed. Berry ovoid, orange, 1–7-seeded. Seeds large; embryo straight, cotyledons fleshy, thick, radicle tiny; endosperm very scanty.

Two endemic and endangered species in the Greater Antilles: G. elegans Wydl. (Puerto Rico) and G. ekmanii O.E. Schulz ex O.C. Schmidt (Hispaniola).

  1. II.

    Subfam. Cestroideae Burnett (1835).

Androecium 4 or 5, often didynamous or heterodynamous, staminode usually present. Cotyledons incumbent; endosperm copious.

  1. II.1.

    Tribe Cestreae Dumort. (1829).

Shrubs. Flowers sessile or with articulate pedicels; corolla aestivation valvate-induplicate, slightly contorted; stamens equal or unequal, filaments straight, generally geniculate at their insertion; ovary with a small annular remainder of the deciduous corolla (cyathium). Embryo straight or slightly curved cotyledons broader than the rest of the embryo.

  1. 7.

    Cestrum L.

    Cestrum L., Sp. pl. 1: 191 (1753); Francey, Candollea 6: 46–398 (1935), 7: 1–132, pl. I–III (1936), rev.; Benítez de Rojas & D’Arcy, Ann. Missouri Bot. Gard. 85(2): 273–351 (1998); reg. rev.; Romanutti & Hunziker, Fl. Fanerog. Argent. 55: 4–12 (1998), reg. rev.; Nee, Solanaceae V. Advances in Taxonomy and Utilization: 109–136 (2001), rev.

Shrubs or trees, rarely vines, some fragrant. Leaves entire, unequal. Inflorescence terminal, monochasial cymes, up to 40-flowered. Bracts deciduous or persistent; calyx campanulate, urceolate, tubular, or cup-shaped; corolla variously coloured, tubular, infundibuliform, hypocrateriform, or slightly urceolate; stamens equal or unequal, geniculate and inserted at different heights in corolla tube, filaments with or without appendages; anthers dorsifixed; ovary stipitate pedicellate, annular nectary prominent; stigma wet, papillate. Berry ovoid or ellipsoid. Seeds subpolyhedrical; cotyledons usually shorter than rest of embryo. n = 8, 16; 2n = 16.

Nearly 150–200 spp. in tropical and subtropical America. Some species cultivated as garden plants (C. nocturnum L., etc.), others are poisonous (C. parqui L’Hérit., C. laevigatum Schltdl.).

Cestrum, Sessea and Vestia appear to form a monophyletic group (tribe Cestreae) well supported by molecular (Santiago-Valentín and Olmstead 2003; Martins and Barkman 2005; Montero-Castro et al. 2006; Olmstead et al. 2008), morphological (Hunziker 2001) and cytological (Sykorova et al. 2003) characters.

  1. 8.

    Sessea Ruiz & Pav

    Sessea Ruiz & Pav., Fl. peruv. prodr.: 21, tab. 33 (1794); Bitter, Repert. Spec. Nov. Regni Veg. 18 (513/523): 199–225 (1922), rev.; Benítez de Rojas & Nee, Solanaceae V. Advances in Taxonomy and Utilization: 153–159 (2001), reg. rev.

    Sesseopsis Hassler (1917).

Shrubs to trees, frequently with dendroid or candelabra hairs. Leaves entire, membranous or subcoriaceous, some looking like stipules. Inflorescence axillary or terminal, mostly several- to many-flowered. Flowers 5-merous, commonly with bracteoles; calyx tubular, lobes equal or unequal; corolla tubular or slightly infundibuliform; stamens geniculate and inserted at different heights, filaments pilose and thickened at insertion; anthers included; ovary pubescent, 4-lobed; stigma lobed, deflexed or capitate. Capsule bivalvate, valves bifid. Seeds 6–19, elongate and imbricate in fruit, with a peripheral narrow hyaline wing. 2n =16.

15 spp., Andes of South America.

Montero-Castro et al. (2006) found evidence for the paraphyly of Sessea in relation to Vestia which needs to be validated with additional sampling and inclusion of related genera.

  1. 9.

    Vestia Willd.

    Vestia Willd., Enum. pl. 1: 208 (1809).

Much-branched shrubs, ill-scented. Leaves entire. Flowers 5-merous, pendant, shortly pedicellate; calyx tubular, minutely dentate, glabrous; corolla greenish yellow, hypocrateriform; stamens strongly exserted, attached slightly above basal fourth of corolla, filaments unequal, thickened at insertion and pubescent from there downwards; anthers basifixed; gynoecium longer than stamens; ovary bilocular, nectary annular; stigma capitate. Capsule 2-valved, valves bifid. Seeds ca. 50, subpolyhedrical, angulate, wingless; testa tuberculate. n = 8; 2n = 16.

A single species in Chile. Vestia foetida (Ruiz & Pav.) Hoffmanns. is a well-known medicinal plant.

  1. II.2.

    Tribe Browallieae Hunz. (1995).

Herbs or small shrubs. Corolla zygomorphic; stamens 4 (anterior aborted), didynamous; pollen grains 4–9-colporate; style geniculate and tortuose at apex, strongly corrugated above; stigma broad, two-horned. Seeds cuboidal or cuboidal-elongated; embryo slightly curved, cotyledons two times shorter than the rest of the embryo.

  1. 10.

    Browallia L.

    Browallia L., Sp. pl. 2: 631 (1753).

Annual herbs. Leaves cordate-ovate to linear-oblong. Inflorescence a raceme or flowers solitary, showy. Calyx actinomorphic; corolla superiorly ventricose, occasionally slightly bilabiate, aestivation reciprocative; upper lateral pair of stamens with filaments flattened, glabrous or with branched trichomes, and anthers with one thecae aborted, lower posterior pair terete and narrow, glabrous or pilose at apex, and anthers with both thecae fertile; ovary glabrous or with rigid trichomes, with cryptic or annular and thick nectary. Capsule globose, elongated or obconic, calyx accrescent and papery. Seeds angulate 10–45, testa reticulate-foveolate. n = 10, 11, 22, 44; 2n = 20, 22, 44.

About eight spp. from southern Arizona (USA) to Andean South America, reaching Bolivia. At least two species, B. americana L. and B. speciosa Hook., have ornamental value.

A critical treatment of the genus is strongly needed to clarify its obscure taxonomy. Browallia and Streptosolen form a sister pair well-supported by morphological (Hunziker 2001), palynological (Stafford and Knapp 2006) and molecular (Martins and Barkman 2005; Olmstead et al. 2008) data.

  1. 11.

    Streptosolen Miers

    Streptosolen Miers, Ann. Mag. Nat. Hist. II, 5 (27): 207 (1850).

Evergreen scabrous-pubescent shrubs, densely branched. Leaves ovate, petiolate. Inflorescence subcorymbose cymes. Flowers pedicellate; calyx zygomorphic, tubular, 4–5-lobed; corolla infundibuliform, tube spirally twisted, not ventricose, limb rather narrow, aestivation cochlear-conduplicate; upper lateral pair of stamens with short filaments and small anthers (usually one theca aborted), lower posterior pair with long filaments and larger anthers; ovary pubescent above, nectary annular, thick. Capsule globose to ovate. Seeds 60–80, cuboidal-elongated; testa reticulate. n = 11, 12.

A single species, S. jamesonii (Benth.) Miers, Colombia, Ecuador and Peru. Ornamental.

  1. II.3.

    Tribe Salpiglossideae Benth. (1835).

Herbs or shrubs. Pubescence of glandular trichomes with multicellular, multiseriate heads and long uniseriate stalk. Flowers mostly solitary on long pedicels; corolla aestivation cochlear; androecium 4–5-merous, stamens fertile didynamous, the posterior pair with larger anthers than the lateral pair; pollen in monads or in tetrads; style expanded or spoon- shaped at apex, stigma linear. Seeds compressed; embryo coiled, cotyledons up to three times shorter than the rest of the embryo.

  1. 12.

    Salpiglossis Ruiz & Pav. Fig. 62

    Fig. 62.
    figure 7

    Solanaceae. Salpiglossis sinuata. A Basal portion of branch. B Flowering branch. C Glandular trichome. D Flower. E Inside view of two calyx segments, showing venation. F Opened corolla. G Anther, lateral view. H Anther, ventral view. I Anther, dorsal view. J Pollen tetrad. K Gynoecium. L Transverse section of ovary. M Dehiscent fruit. (Hunziker 2001; drawn by N. Flury)

    Full size image

    Salpiglossis Ruiz & Pav., Fl. peruv. prodr.: 94, tab. 19 (1794); Hunziker & Subils, Bot. Mus. Leafl. 27(1–2): 1–43 (1980), rev.

Herbs or shrubs, densely viscid-pubescent; stems leafy or almost without leaves, sometimes xeromorphic (S. spinescens) with spine-like processes. Flowers solitary or in panicles, chasmogamous or cleistogamous; perianth 5-merous; calyx actinomorphic, campanulate; corolla zygomorphic, infundibuliform; androecium with one or without staminode (S. sinuata); anthers ovoid, dorsifixed; ovary bicarpellate, nectary annular; style hollow, expanded distally. Capsule ovate 2–4-valved. Seeds up to 300 or more, testa reticulate. n = 22.

Two spp. endemic to southern Andean South America (Chile and Argentina).

  1. 13.

    Reyesia Gay

    Reyesia Gay, Fl. Chile 4: 418 (1840); Hunziker & Subils, Bot. Mus. Leafl. 27(1–2): 1–43 (1980), rev.

Annual or perennial herbs, dichasially branched, glutinose. Lower leaves entire to pinnatifid or pinnatisect, the upper smaller or linear scales. Flowers solitary, terminal, small; calyx actinomorphic, strongly glanduliferous; corolla zygomorphic, tubular or infundibuliform; the posterior pair of stamens always fertile, the lateral pair with fertile or sterile anthers; anthers basifixed, thecae usually unequal, divaricate; ovary bicarpellate, nectary pelviform, bilobed, style hollow or solid, spoon-shaped distally. Capsule 2–4-valved hidden at the base of the persistent calyx. Seeds (3)20–25, testa reticulate or granulate.

Four spp. in Andean northern Chile, one reaching Argentina.

  1. II.4.

    Cestroideae: Incertae sedis

  1. 14.

    Protoschwenckia Soler.

    Protoschwenckia Soler., Ber. Deutsch. Bot. Ges. 16: 243 (1898); Freire de Carvalho, Sellowia 18(18): 67–72 (1966), reg. rev.

    Schwenkiopsis Dammer (1916).

Small shrubs, profusely branched, with non-glandular and glandular trichomes, and occasionally branched trichomes. Leaves entire, ovate, base cordate. Inflorescence paniculate. Flowers pedicellate; calyx actinomorphic, with 5 equal uninerved lobes; corolla actinomorphic, tubular, lobe margins inflexed; aestivation valvate-induplicate; androecium and gynoecium included; stamens 4 (2 + 2), filaments filiform, pubescent from insertion to tube downwards; anthers dorsifixed; ovary with a cushion-shaped nectary; style short, stigma discoidal-capitate. Capsule septicidal-loculicidal, with 2 bifid valves. Seeds ca. 22, subreniform; embryo slightly curved, cotyledons 3 times shorter than rest of embryo.

A single species, P. mandonii Soler., Andes of Bolivia, Brazil.

  1. 15.

    Heteranthia Nees & Mart.

    Heteranthia Nees & Mart., Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: 41 (1823).

Perennial herbs. Leaves petiolate, ovate to obovate. Inflorescence racemose. Calyx actinomorphic, 5-lobed; corolla zygomorphic, funnel-shaped, slightly bilabiate, basal tube narrow and short, lobules very short and broad, aestivation unknown; stamens 2 + 2; anthers ventrifixed, thecae unequal; ovary subconical or pyriform, nectary absent; style exserted, curved or almost uncinate. Capsule septicidal, 2-valved; calyx accrescent. Seeds 20–28, cuboidal; embryo straight, very small, cotyledons unequal.

Monotypic, H. decipiens Nees & Mart., eastern Brazil.

  1. III.

    Subfam. Nicotianoideae Miers (1848).

Mostly herbs or shrubs. Calyx non-accrescent or scarcely accrescent; androecium tetramerous (staminode present), or pentamerous (stamens of two lengths). Embryo mostly straight or slightly curved, cotyledons short to very short relative to the rest of the embryo.

  1. III.1.

    Tribe Anthocercideae G. Don (1838).

Shrubs. Calyx non-accrescent; corolla broadly actinomorphic, aestivation valvate-supervolute; anthers ventrifixed, extrorse, attached low in the corolla tube. Mostly capsule. Seeds reniform to elongate, testa reticulate; embryo terete, cotyledons 5–12 times shorter than the rest of the embryo; endosperm with a small oil sector.

  1. 16.

    Symonanthus Haegi Fig. 63

    Fig. 63.
    figure 8

    Solanaceae. Symonanthus aromaticus. A Flowering branch. B Leaf, abaxial view. C Transverse section of leaf. D Dendritic trichome. E Glandular trichome. F Staminate flower. G Staminate flower, opened corolla. H Carpellate flower, longitudinal section. I Apex of corolla lobe. J Anther, lateral view. K Anther, ventral view. L Rudimentary ovary of staminate flower. M Fruit. N Seed. O Seed, transverse section (the stippled lower area is oily). P, Q Two views of embryo. (Hunziker 2001; drawn by J. de Ugarte)

    Full size image

    Symonanthus Haegi, Telopea 2: 175 (1981); Purdie, Symon & Haegi, Fl. Australia 29: 13–17 (1982).

Dioecious or hermaphrodite shrubs, indumentum of glandular and non-glandular branched hairs. Leaves long and narrow, margin revolute. Flowers solitary or in 2–3-flowered cymes; calyx 5-lobed; corolla narrowly tubular or cylindrical-campanulate, upper half funnel-shaped, almost truncate or with 5 equal, very short and broad lobules; staminate flower: stamens equal, 5 or 4 (then staminode sometimes present); anthers dithecal, gynoecium much reduced; pistillate flower: stamens sterile, ovary large, nectary annular, ovules 4–10; style well developed, stigma bilobed. Capsule subglobose, septicidal-loculicidal, 4-valved. Seeds few, reniform; embryo slightly arcuate; endosperm copious. n = 36; 2n = 32.

Two spp., south-western Australia.

  1. 17.

    Anthocercis Labill.

    Anthocercis Labill., Nov. Holl. pl. 2: 19, tab. 158 (1806); Purdie, Symon & Haegi, Fl. Australia 29: 6–13 (1982).

Shrubs, glabrous or pubescent, occasionally viscid, usually with glandular trichomes. Leaves variously shaped, sometimes thick and fleshy or leathery, usually entire, or minutely toothed or serrulate-crenulate. Inflorescences cyme-, raceme- or panicle-like. Flowers perfect; calyx actinomorphic, 5-partite; corolla narrowly 5-lobed; stamens 4, didynamous or subequal (sometimes with a staminode); anthers dithecal; nectary annular; stigma bilobed. Capsule pear-shaped or ovoid, septicidal-loculicidal, 2–4-valved. Seeds up to 100, reniform; endosperm copious. n = 36.

10 spp., mainly south-western temperate Australia. Some species have occasionally caused poisoning of children or cattle.

  1. 18.

    Grammosolen Haegi

    Grammosolen Haegi, Telopea 2: 178 (1981); Purdie, Symon & Haegi, Fl. Australia 29: 28–29 (1982), rev.

Shrubs; branches and leaves densely tomentose, with dendritic trichomes. Leaves crowded or well-spaced, subsessile. Inflorescences cymose, 1–3-flowered, sometimes forming leafy spikes. Flowers perfect; calyx 5-lobed, densely tomentose; corolla tubular, lobes long and narrow; stamens 5, unequal (2 + 3); anthers monothecal; nectary annular; stigma bilobed. Capsules septicidal-loculicidal, 4-valved. Seeds reniform; endosperm copious. n = 56.

Two spp., South Australia.

  1. 19.

    Cyphanthera Miers

    Cyphanthera Miers, Ann. Mag. Nat. Hist. II, 11 (65): 376 (1853); Purdie, Symon & Haegi, Fl. Australia 29: 21–28 (1982).

    Anthocercis sect. Cyphanthera (Miers) F. Muell. (1861).

Shrubs, usually pubescent, viscid or slightly to densely woolly-tomentose, trichomes glandular or dendritic. Leaves usually sessile, variously shaped. Inflorescence a cyme, or raceme-, panicle- or thyrse-like, or flowers solitary. Flowers perfect; calyx small; corolla funnel-shaped to campanulate; stamens 4 (2 + 2), sometimes also a staminode present; anthers monothecal; nectary annular; stigma slightly bilobed. Capsule septicidal-loculicidal, 4-valved. Seeds ca. 6, reniform. n = 30; 2n = 60.

Nine spp., southern temperate Australia.

A phylogenetic analysis (six spp. sampled) revealed that Cyphantera is not monophyletic (García and Olmstead 2003; Clarkson et al. 2004; see comment under Duboisia).

  1. 20.

    Anthotroche Endl.

    Anthotroche Endl. in Endl. & Fenzl, Nov. stirp. dec. 1: 6 (1839); Purdie, Symon & Haegi, Fl. Australia 29: 30–34 (1982), rev.

Shrubs with a very dense indumentum of dendritic and glandular trichomes. Leaves shortly petiolate, variously shaped. Flowers solitary or in 2–6-flowered clusters, sometimes forming leafy pseudospikes; calyx actinomorphic; corolla campanulate to rotate, densely pubescent, lobes broad and short; stamens 5, equal, included or exserted, filaments sometimes incurved and pubescent; anthers monothecal, horseshoe-shaped. Capsules ovoid to ellipsoid or globose, septicidal-loculicidal, 4-valved. Seeds reniform; endosperm copious. n = ca. 31.

Four spp., arid inland of south-western Western Australia.

  1. 21.

    Crenidium Haegi

    Crenidium Haegi, Telopea 2: 179. 1981; Purdie, Symon & Haegi, Fl. Australia 29: 34 (1982), rev.

Intricately branched shrubs, leafless except on young twigs; branches spinescent, tomentose with dendritic trichomes. Leaves sessile, margin slightly revolute. Flowers in cymose clusters. Flowers perfect; calyx minutely 5-lobed; corolla narrowly tubular with ovate to broadly ovate and short lobes; stamens 4 (2 + 2 and a staminode usually present), exserted; anthers monothecal; nectary annular; stigma shortly bilobed. Capsules ovoid, septicidal-loculicidal, valves 2, bifid. Seeds 20–100, reniform. n = 35.

Monotypic, C. spinescens Haegi, inland south-western Western Australia.

  1. 22.

    Duboisia R. Br.

    Duboisia R. Brown, Prodr.: 448 (1810); Purdie, Symon & Haegi, Fl. Australia 29: 17–21 (1982), rev.

Shrubs or trees, glabrescent to glabrous; bark corky and thick; wood extremely light. Leaves narrowly obovate to obovate-elliptic, or linear. Inflorescence paniculate. Flowers perfect; calyx campanulate; corolla campanulate-funnel-shaped; stamens 4 (2 + 2 and sometimes a vestigial staminode); anthers monothecal; nectary annular; stigma bilobed or bifid. Berry globose or subglobose, rarely ellipsoid to ovoid. Seeds 2–12, reniform; endosperm copious, starchy, oily sector absent. n = 30; 2n = 60.

Four spp. in New Caledonia and eastern Australia. Species of Duboisia known as “corkwood” are harvested commercially to obtain scopolamine for medicinal purposes, and are toxic to cattle, horses and humans.

Duboisia does not form a monophyletic group (García and Olmstead 2003; Clarkson et al. 2004), and a re-circumscription of this genus and also of Cyphanthera (C. albicans is resolved in the Duboisia clade) is needed.

  1. III.2.

    Tribe Nicotianeae G. Don

Mostly herbs or shrubs. Inflorescence terminal, paniculoid, many-flowered; corolla aestivation contorted-conduplicate; anthers dorsifixed; stamens 5, equal or 4 + 1 or 2 + 2 + 1. Capsule with up to ca. 5000 seeds.

  1. 23.

    Nicotiana L.

    Nicotiana L., Sp. pl. 1: 180 (1753); Goodspeed, The genus Nicotiana: 1–536 (1954), rev.; Cocucci & Hunziker, Flora Fanerog. Argent. 89: 3–14 (2005); Knapp, Chase, & Clarkson, Taxon 53: 73–82 (2004), rev.

Herbs to small trees. Basal leaves spathulate, sometimes rosulate, cauline leaves increasingly smaller and similar to inflorescence bracts, margin entire or undulate. Flowers nocturnal, diurnal or cleistogamous; corolla actino- or zygomorphic,<90 mm, tubular, salverform, or campanulate, variously coloured, tube straight or curved, sometimes very narrow or broadly inflated, lobes short, often emarginate; stamens included or exserted, sometimes strongly geniculate; anthers circular to elongated. Capsule bivalvate. 2n = 18, 20, 24, 32, 36, 40, 48. In some species, 2n = 38, 42, 44, 46.

Near 76 spp., southern South America east of the Andes (40 spp.), Africa (1 sp.), Australia (25 spp.), SW North America (8 spp.).

  1. IV.

    Subfam. Solanoideae Burnett (1835).

Herbs to small trees; corolla aestivation valvate, cochlear or contorted. Androecium 5-merous (rarely 4-merous). Fruit mostly a berry but also schizocarp and less frequently drupe, diclesium or pixidium. Seeds flattened.

  1. IV.1.

    Tribe Lycieae Lowe (1872).

Shrubs or small trees. Perianth pentamerous, sometimes tetramerous; corolla aestivation cochlear or cochlear-plicate; stamens 5, anthers dorsifixed; ovary 2–4-locular. Fruit a berry or a drupe. Embryo annular or subcoiled, cotyledons incumbent; endosperm copious.

  1. 24.

    Lycium L.

    Lycium L., Sp. pl. 1: 191 (1753); Bernardello, Bol. Acad. Nac. Ci. Republ. Argent. 57: 173–356 (1986), part. rev.; Chiang-Cabrera, Ph.D. diss., The University of Texas at Austin (1981), part. rev.

    Grabowskia Schltdl. (1832).

    Phrodus Miers (1849).

Mostly shrubs, usually with cauline spines. Flowers axillary, solitary or in clusters; calyx actinomorphic or zygomorphic; corolla tubular or infundibuliform; stamens exserted or included, filaments cylindrical or enlarged at proximal end; thecae free in their basal third or half; ovary bicarpellate, completely bilocular or tetralocular in its upper part, 1–∞ ovules per carpel, nectary annular or invaginate-bilobed, inconspicuous and greenish or conspicuous and red. Berry generally many-seeded, rarely drupaceous with two 1-seeded pyrenes or intermediate with stone cells or sclerenchyma. Seeds discoidal-compressed or ovoid and plano-convex. n = 12, 18, 24, 36, 48, 60; 2n = 24, 36, 48.

Cosmopolitan genus of ca. 92 spp. Recent phylogenetic studies (Levin and Miller 2005; Levin et al. 2007, 2011) have shown that the Chilean monotypic Phrodus (L. bridgesii (Miers) Levin, Miller & Bernardello) and the few species of the exclusively American genus Grabowskia, reduced to one (L. boerhaviifolium L.f.), must be included in Lycium.

  1. IV.2.

    Tribe Hyoscyameae Endl. (1839).

Perennial herbs. Flowers usually solitary; corolla aestivation cochlear. Fruit mostly a pyxidium many-seeded and hidden in the accrescent calyx. Embryo strongly curved to subcoiled; endosperm rather copious.

  1. 25.

    Hyoscyamus L.

    Hyoscyamus L., Sp. pl. 1: 179 (1753); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 26 (1997), rev.

    Archihyoscyamus Lu (1997).

Herbs with rhizomes or napiform, woody or fleshy roots; glabrous, villose or glandular pubescent. Leaves in a rosette. Lower flowers axillary, solitary, upper ones in condensed inflorescences; calyx variously shaped, densely veined, 5-toothed or 5-lobed; corolla zygomorphic, tube short, expanding upwards, or subrotate, lobes unequal; androecium and gynoecium mainly exserted; filaments unequal; anthers dorsifixed; nectary indistinct; style filiform, stigma capitate. Pyxidium elongate, hidden by the accrescent calyx, calyx teeth spine-tipped and rigid. Seeds 200–500, discoid; embryo strongly curved, cotyledons unequal, as long as or shorter than rest of embryo. n = 14, 17, 34; 2n = 28, 34, 68.

About 17 spp., from Madeira, Canary Islands, and North Africa to India and China.

  1. 26.

    Przewalskia Maxim.

    Przewalskia Maxim., Bull. Acad. Imp. Sci. Saint-Pétersbourg. 27: 507 (1881); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 27 (1997), rev.

Perennial herbs with fleshy roots; indumentum glandular. Leaves densely aggregated. Inflorescence with clusters of 1–6 axillary flowers; calyx actinomorphic, cylindrical-campanulate; corolla actinomorphic, tubular; stamens equal, exserted, filaments glabrous, attached above; anthers dorsifixed; nectary annular; stigma discoidal-capitate, exserted. Pyxidium globose, included in the much accrescent, urceolate-ovoid and strongly veined calyx, calyptra convex and apiculate. Seeds reniform. 2n = 44.

A single species, P. tangutica Maxim., western China, 3200–5000 m.

  1. 27.

    Scopolia Jacq.

    Scopolia Jacq., Observ. bot.: 32, tab. 20 (1764), nom. cons.; Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 27 (1997), rev.

Perennial herbs with rhizomes. Leaves obovate or oblong. Flowers axillary, solitary; calyx campanulate; corolla campanulate or tubular-campanulate, shallowly 5-lobed, lobes very short and broad; stamens slightly unequal, included, pubescent; anthers probably basifixed; nectary annular; stigma depressed. Pyxidium globose, enclosed in the somewhat enlarged calyx. Seeds discoid; embryo subcoiled, cotyledons slightly shorter than rest of embryo. n = 24; 2n = 44, 46, 48, 84, 168.

Two spp. with disjunct distribution: S. carniolica Jacq. in Europe and S. japonica Maxim. in East Asia.

Within Hyoscyameae, Scopolia is inferred to be paraphyletic in relation to Przewalskia (Olmstead et al. 2008).

  1. 28.

    Physochlaina G. Don

    Physochlaina G. Don, Gen. hist. 4: 470 (1838); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 27 (1997), rev.

Perennial herbs with fleshy, tuberous roots; indumentum non-glandular and glandular. Leaves entire or sinuate-dentate. Inflorescence paniculate, terminal or axillary. Calyx actinomorphic, campanulate, tubular-campanulate or tubular-urceolate, lobes equal; corolla actinomorphic with a basal tube expanding upwards; stamens exserted or included, inserted below middle of corolla; anthers dorsifixed; nectary annular, prominent; stigma discoid. Pyxidium subglobose or elongated included in the much accrescent, inflated, elongated calyx; embryo subcoiled, cotyledons shorter than rest of embryo. 2n = 28, 41, 42, 82.

About eight spp., Asia.

  1. 29.

    Atropanthe Pascher

    Atropanthe Pascher, Oesterr. Bot. Z. 59(9): 529 (1909); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 26 (1997), rev.

Perennial herbs with thick rhizomes. Leaves entire, acuminate. Flowers solitary, axillary, pendant; calyx campanulate-urceolate, lobes subequal; corolla slightly zygomorphic, cylindrical-campanulate, lobes unequal, revolute; stamens included, filaments unequal, circinate in bud, curved at anthesis, inserted basally; anthers ventrifixed; nectary annular prominent; stigma bilobed. Pyxidium pendant, small, globose and apiculate, included in the enlarged, urceolate and inflated calyx. Seeds ca. 50, discoid; embryo subcoiled, cotyledons slightly shorter than rest of embryo. 2n = 48.

Monospecific, A. sinensis (Hemsley) Pascher, southern and central China, 1400–3000 m.

  1. 30.

    Anisodus Link

    Anisodus Link ex Sprengel, Syst. veg. 1: 699 (1825); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27: 25 (1997), rev.

Cryptophytic herbs, glabrous or pubescent, roots woody, napiform. Leaves entire, sinuose or dentate. Flowers axillary, solitary, on long pedicels; calyx campanulate or urceolate, zygomorphic, irregularly 5-lobed; corolla campanulate or campanulate-urceolate, lobes equal, auriculate, revolute; stamens equal, included, filaments straight, adnate basally to corolla tube; anthers dorsifixed; ovary conical, nectary annular, prominent, thick; stigma discoid. Pyxidium usually globose, 2–4 times shorter than the ribbed calyx. Seeds discoid. 2n = 24, 42, 48, 50.

Four spp. in the mountains of Nepal, India, Bhutan, and China, 2000–4500 m.

  1. 31.

    Atropa L.

    Atropa L., Sp. pl.: 181 (1753); Hoare & Knapp, Bull. Nat. Hist. Mus. London, Bot. 27(1): 26 (1997), rev.

    Pauia Deb & R. M. Dutta (1965).

Perennial herbs, glabrous or glandular-pubescent. Basal leaves alternate, upper geminate and anisophyllous. Flowers solitary, paired or in triplets, pendant or erect; calyx campanulate, tube shorter than lobes; corolla slightly zygomorphic, tubular-campanulate, bell-shaped or infundibuliform, lobes rounded, slightly unequal and curved outwards; androecium and gynoecium included or exserted; filaments unequal, incurved, attached basally; anthers dorsifixed; nectary annular; style somewhat arched, stigma capitate. Berry black and juicy, calyx somewhat enlarged. Seeds ca. 25–160, reniform; embryo annular, cotyledons almost as long as rest of embryo. n = 36; 2n = 50, 72, 74.

Three spp. in Europe, in Asia to Mongolia, and in northern Morocco.

  1. IV.3.

    Tribe Juanulloeae Hunz. (1977).

Mostly epiphytes or lianas, some genera myrmecophilous, others bat-pollinated. Corolla aestivation cochlear or valvate. Fruit a large berry. Embryo straight to annular; cotyledons accumbent or oblique, shorter than the rest of the embryo; endosperm very scarce to copious.

  1. 32.

    Dyssochroma Miers

    Dyssochroma Miers, Ann. Mag. Nat. Hist. II, 4(22): 250 (1849).

Shrubs or climbers, epiphytic or terrestrial; brachyblasts cauliflorous. Leaves coriaceous. Inflorescence terminal, a congested cyme or a single flower. Flower pendant, 5-merous; calyx subcampanulate, coriaceous, deeply divided; corolla thick, shortly cylindrical, expanded upwards into an infundibuliform-campanulate or ventricose part, limb 5-partite, aestivation valvate; stamens straight, attached basally to corolla; anthers basifixed; nectary annular, well developed; style articulate, stigma saddle-shaped or very long and decurrent. Berry ovoid-globose. Seeds boomerang- or rod-shaped; embryo curved, cotyledons oblique, as wide as rest of embryo; endosperm abundant. 2n = 24.

Two spp. in Brazil.

Knapp et al. (1997) suggested Dyssochroma to be congeneric with Trianaea.

  1. 33.

    Juanulloa Ruiz & Pav. Fig. 64

    Fig. 64.
    figure 9

    Solanaceae. Juanulloa speciosa. A Flowering branch. B Glandular trichome. C Flower bud, top view showing aestivation. D Young flower, longitudinal section. E Stamen, ventral view. F Stamen, dorsal view. G Fruit, accrescent calyx cut longitudinally. H Seed. I Seed, cross section. J Embryo. (Hunziker 2001; drawn by N. Flury)

    Full size image

    Juanulloa Ruiz & Pav., Prodr. Fl. peruv. prodr.: 27, tab. 4 (1794); Hunziker & Subils, Kurtziana 21: 209–235 (1991), rev.

    Laureria Schltdl. (1833).

    Sarcophysa Miers (1849).

    Ectozoma Miers (1849).

    Rahowardiana D’Arcy (1973).

Climbers, shrubs or small trees, epiphytic or terrestrial. Leaves coriaceous or membranous. Inflorescences terminal or lateral, variously modified cymes or raceme-like, usually many-flowered or reduced to 1–2 flowers, inflorescence rhachis short or very long, pendant. Calyx fleshy, 5-lobed, or deeply divided or unlobed and inflated; corolla fleshy, tubular or funnel-form, slightly zygomorphic, aestivation cochlear; stamens generally included, attached at different levels of corolla, pubescent at point of insertion; anthers dorsifixed; nectary annular, well developed; stigma saddle-shaped. Berry ovoid or subglobose, calyx accrescent. Seeds shortly subreniform or boomerang-shaped; embryo curved, cotyledons accumbent, wider than rest of embryo.

About 11 spp. from southern Mexico to Bolivia.

  1. 34.

    Markea Rich.

    Markea Rich., Actes Soc. Hist. Nat. Paris 1: 107 (1792); Hunziker, Kurtziana 25: 67–113 (1997), rev.

    Hawkesiophyton Hunz. (1977).

Epiphytic, shrubs or small trees, frequently with root tuberosities. Leaves in pseudowhorls, coriaceous or membranous. Flowers solitary or in a cyme of 2–17, usually pedicellate, on long and flexible or short and rigid peduncles; calyx tube very short or well developed; corolla thick, funnel-form or infundibuliform, lobes not reflexed at anthesis, aestivation cochlear; stamens adnate basally and pubescent at point of insertion; anthers basifixed, frequently connivent; nectary well developed; stigma capitate, saddle-shaped or with 2 short branches. Berry ellipsoid, calyx accrescent. Seeds 40–100; embryo slightly curved, cotyledons oblique, wider than rest of embryo.

About nine spp. from Panama to Peru.

  1. 35.

    Merinthopodium Donn. Sm.

    Merinthopodium Donn. Sm., Bot. Gaz. 23: 11, pl. I (1897).

Epiphytic shrubs or small trees, rarely terrestrial. Leaves subcoriaceous or membranous, glabrous or minutely pubescent on the main vein beneath. Inflorescence a pendant raceme, peduncles long (12–90 cm), flexible, slender, bearing 1–14 pedicellate flowers. Calyx campanulate, actinomorphic, tube very short; corolla broadly campanulate, lobes short, reflexed at anthesis, aestivation valvate; stamens attached basally; anthers dorsifixed and somewhat versatile, thecae not confluent; nectary annular; style straight, stigma saddle-shaped. Berry ovoid, surrounded by the slightly accrescent calyx. Seeds boomerang-shaped; embryo thick and short, curved, cotyledons accumbent; endosperm rather scarce.

About three spp. from southern Mexico to Venezuela and Colombia. The taxonomy of this genus is obscure.

  1. IV.4.

    Tribe Datureae Dumort. (1829).

Herbs, shrubs or small trees. Flowers solitary, large; corolla aestivation contorted-conduplicate; anthers basifixed, pubescent, dehiscence latrorse or extrorse. Embryo annular; cotyledons shorter than rest of embryo; endosperm scarcely copious, rich in protein.

  1. 36.

    Brugmansia Pers.

    Brugmansia Pers., Syn. pl. 1: 216. 1805; Nee, Solanaceae Newslett. 3 (2): 27–35 (1991); Preissel & Preissel, Engelstrompeten Brugmansia und Datura: 8–101 (1997), rev.

    Methysticodendron R.E. Schult. (1955).

Shrubs or trees. Leaves large, dentate or sinuate-repand. Flowers pendant or inclined, 5-merous; calyx zygomorphic, unequally toothed, sometimes splitting along one side (spathe-like), falling off after anthesis, or persisting around the fruit; corolla very large, broadly infundibuliform, rarely narrow-infundibuliform to almost cylindrical; stamens inserted near middle of corolla; ovary superior, completely bilocular, nectary annular; stigma long, saddle-shaped. Berry large, pendant. Seeds up to 300, large, testa usually thick, sometimes suberose. n = 12; 2n = 24.

Six spp. in Andean South America (Venezuela to Bolivia), some used as ornamentals or for medicinal and psychotropic preparations, for example: B. arborea (L.) Lagerh., B. sanguinea (Ruiz & Pav.) D. Don.

  1. 37.

    Datura L.

    Datura L., Sp. pl. 1: 179 (1753); Avery et al.: The genus Datura: 1–289 (1959), rev.; Symon & Haegi, in Hawkes et al. (eds.) Solanaceae III. Taxonomy, Chemistry, Evolution: 197–210 (1991); Preissel & Preissel. Engelstrompeten. Brugmansia und Datura: 102–137 (1997), rev.

Annual herbs or short-lived perennials dichasially branched. Leaves sinuate-dentate or lobed. Flowers erect or oblique; calyx cylindrical, shortly (3)5(6)-lobed, circumscissile after anthesis; corolla broadly funnel-form; stamens included, attached basally; ovary partially inferior, tetralocular in lower half, with small fleshy spines; nectary invaginate-pelviform; stigma bilobed. Capsule erect or deflexed, septifragal with 2–4 valves (D. ceratocaula with smooth berry), pericarp with spines, bristles, or blunt tubercles; calyx remnants accrescent and reflexed. Seeds up to ca. 500, sometimes with elaiosome; endosperm with lipid vesicles and rich in protein bodies. n = 12, 24; 2n = 24, 48.

About 9 to 11 spp. from Mexico to south-western USA. Datura ferox L. and D. stramonium L. are toxic weeds.

  1. IV.5.

    Tribe Physalideae Miers (1849).

Mostly shrubs, small trees or perennial herbs. Flowers in axillary fascicles; corolla aestivation usually valvate, rarely contorted; stapet frequently present, inconspicuous or conspicuous, with or without basal auricles. Embryo subcoiled to strongly bent.

  1. IV.5.a.

    Subtribe Withaniinae Bohs & Olmstead (2008).

Corolla campanulate, campanulate-urceolate, rotate or salver-form, aestivation valvate; filaments often with conspicuous lateral appendages; calyx often not accrescent.

  1. 38.

    Discopodium Hochst.

    Discopodium Hochst., Flora 2: 22 (1844).

Shrubs or small trees. Leaves frequently anisophyllous. Flowers in fascicles of 5–10; calyx cup-shaped, subtruncate, denticulate; corolla campanulate-urceolate, tube and lobes with dense indumentum of simple trichomes; stamens included, inserted basally; anthers dorsifixed and pubescent on ventral surface, connective thick; nectary annular, protruding, almost as high as the rest of the ovary; style with antrorse trichomes, stigma capitate-discoidal. Berry red, globose; calyx not accrescent. Seeds ca. 20, subreniform, compressed; embryo annular, cotyledons shorter than rest of embryo; endosperm copious.

One sp., D. penninervium Hochst., tropical Africa.

  1. 39.

    Nothocestrum A. Gray

    Nothocestrum A. Gray, Proc. Amer. Acad. Arts. 6: 48 (1866).

Shrubs or trees. Leaves often geminate. Inflorescences lateral, with (1)2–3(10) pedicellate and fragrant flowers, on prominent brachyblasts. Flowers 4-merous; calyx unevenly toothed or lobed, or almost bilabiate; corolla salver-form, with 4 well-developed folds; stamens included or exserted; anthers sessile or subsessile, dorsi-basifixed; ovary partly inferior, nectary annular, slightly protruding; style short, stigma saddle-shaped. Berry globose or fusiform, few-seeded, calyx accrescent or not. Seeds subreniform, compressed; embryo annular, cotyledons as long or slightly shorter than rest of embryo; endosperm scanty. n = ca. 24.

A genus of five or six species endemic to the Hawaii Archipelago, some of them federally listed as endangered.

  1. 40.

    Tubocapsicum (Wettst.) Makino

    Tubocapsicum (Wettst.) Makino, Bot. Mag. (Tokyo) 22: 18 (1908); D’Arcy et al., Bot. Bull. Acad. Sin. 42: 67–84 (2001), rev.

Perennial herbs; stems stout, dichasially branched, glabrate. Leaves alternate or anisogeminate. Flowers in fascicles of 2–6. Pedicels obconically thickened at apex; calyx cup-shaped, truncate; corolla broadly campanulate, divided almost to middle or to 2/3 into reflexed segments; stamens basally broad, inserted usually in upper half of corolla; anthers basifixed and versatile; nectary hardly visible; stigma capitate. Berry juicy, red, shining, pericarp lacking stone cells, calyx persistent, hardly accrescent. Seeds ca. 100, suborbiculate, laterally compressed; embryo annular; endosperm rather abundant. n = 24.

One species from southern and eastern Asia: Tubocapsicum anomalum (Franch. & Sav.) Makino.

  1. 41.

    Athenaea Sendtn.

    Athenaea Sendtn. in Mart., Fl. bras. 10 (fasc. 6): 133 (1846), nom. cons.; Barboza & Hunziker, Bol. Soc. Argent. Bot. 26(1–2): 91–105 (1989), rev.

Shrubs with conspicuous non-glandular or viscid-glandular indumentum. Fascicles of up to 6 flowers, rarely solitary; calyx deeply divided, segments equal or unequal, rarely cordiform; corolla rotate, segments 2–4 times longer than the very short tube, white and with green or red spots inside; stapet prominent with two short lateral auricles; anthers dorsifixed; nectary annular; styles heteromorphic, stigma discoidal-depressed. Berry many-seeded, surrounded completely or in part by the accrescent calyx, pericarp glabrous to pubescent, lacking stone cells. Seeds subreniform, discoid; embryo slightly curved, cotyledons much shorter than rest of embryo; endosperm abundant.

Seven spp., south-eastern Brazil.

  1. 42.

    Aureliana Sendtn.

    Aureliana Sendtn. in Mart., Fl. bras. 10 (fasc. 6): 138 (1846); Hunziker & Barboza, Darwiniana 30 (1–4): 95–113 (1990), rev.

Shrubs or small trees. Leaves geminate or alternate. Flowers axillary, usually in up to 26-flowered fascicles; calyx urceolate in bud, later splitting into 5 unequal and short lobes, or not urceolate and lobules equal; corolla rotate, segments mostly with greenish spots inside; conspicuous stapet bidenticulate, anthers dorsifixed; nectary annular; style heteromorphic, stigma discoidal-depressed. Berry many-seeded, calyx not accrescent, pericarp and placenta with parenchymatic proliferations, pericarp with few small stone cells. Seeds reniform, discoid; embryo subcoiled, cotyledons much shorter than rest of embryo; endosperm abundant. 2n = 24.

Eight spp. in eastern Brazil, Paraguay, Peru, Bolivia, and north-eastern Argentina.

  1. 43.

    Withania Pauquy

    Withania Pauquy, Belladone: 14, f. 4 a–d (1825), nom. cons.; Hepper, in Hawkes et al. (eds.) Solanaceae III. Taxonomy, Chemistry, Evolution: 211–227 (1991), part. rev.; Zhang, Lu & D’Arcy, Fl. China 17: 309–310 (1994), reg. rev.

    Mellisia Hook. f. (1867–1871).

    Physaliastrum Makino (1914).

    Archiphysalis Kuang (1966).

Shrubs or geophytes. Leaves sometimes opposite and frequently anisogeminate. Flowers usually in groups of 2–6, perfect or functionally unisexual, pendant; calyx segments equal or very unequal; corolla broadly campanulate or somewhat infundibuliform, with dendritic trichomes inside, shortly 5-lobulate and with or without folds; perfect flowers: stamens 5, connivent, equal, usually included, filaments generally expanded into a stapet with 2 inconspicuous lateral extensions fused to the corolla; anthers ventrifixed; nectary annular, somewhat protruding; style articulate; pistillate flowers: androecium rudimentary; staminate flowers: small ovary lacking style and stigma. Berry tightly or loosely enclosed by the accrescent urceolate and sometimes rough and muricate calyx. Seeds up to 150, reniform or discoid; embryo strongly bent. n = 12, 24, 36; 2n = 24, 48, 96.

Almost 20 Old World spp. The taxonomy and phylogeny of the genus need clarification.

  1. IV.5.b.

    Subtribe Iochrominae Reveal (2012).

Corolla tubular, funnel-shaped, salver-form or campanulate, aestivation valvate; stapet usually auriculate, anthers mostly dorsifixed. Berry with stone cells; calyx usually not accrescent.

  1. 44.

    Dunalia Kunth

    Dunalia Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 3: 43, tab. 194 (1818), nom. cons.; Hunziker, Bol. Acad. Nac. Ci. Republ. Argent. 41(2): 211–244 (1960), rev.

Shrubs or small trees. Leaves large to small. Inflorescences axillary, sessile, usually 1–5-flowered (20–30-flowered in D. solanacea). Flowers hermaphrodite or functionally unisexual; corolla hypocrateriform or infundibuliform, 5-lobulate, limb narrow, lobes broader than long; stamens glabrous, usually equal, inserted basally, stapet with two lateral, narrow and free wings; anthers basi-dorsifixed, fertile ones larger and exserted, sterile ones small and included; fertile ovary with long style and exserted stigma, nectary included. Berry globose, calyx not accrescent, stone cells rarely absent. Seeds ca. 30 or more; embryo annular. n = 12.

Five spp., Colombia to central Argentina.

  1. 45.

    Iochroma Benth.

    Iochroma Benth., Bot. Reg. 31, tab. 20 (1845), nom. cons.

Shrubs, exceptionally spiny; young stems with simple trichomes. Leaves petiolate, rarely microphyllous. Flowers axillary, pedicellate, solitary or usually in fascicles of 2–13(–18). Perianth 5-lobulate; calyx urceolate in bud, at anthesis campanulate; corolla funnel-shaped, tubular or salver-form, variously coloured, lobes wide, tube with folds; stamens equal or not, inserted at different level of corolla; ovary bicarpellate (rarely 4-carpellate); stigma shortly saddle-shaped. Berry globose to elongated, with many stone cells, calyx usually accrescent. Seeds up to ca. 250; embryo subcoiled or almost hippocrepiform; endosperm abundant. n = 12; 2n = 24.

Around 26 spp. in western South America, reaching north-western Argentina; many are ornamentals due to their coloured long corollas.

A phylogenetic analysis revealed that Iochroma species do not form a monophyletic group (22 spp. sampled); two alternatives have been proposed: to transfer some species to Acnistus, or to consider Acnistus as a monophyletic group within a monophyletic Iochroma s.l. (Smith and Baum 2006).

  1. 46.

    Saracha Ruiz & Pav.

    Saracha Ruiz & Pav., Fl. peruv. prodr.: 31, t. 34 (1794); Álvarez, Systematics of Saracha (Solanaceae), Master thesis, University of Missouri, St. Louis, USA: 1–172 (1996), rev.

Shrubs or small trees; young stems densely hairy with deciduous brown trichomes (thickened, multiseriate, bifurcate or not, ending in a uniseriate row of cells). Leaves coriaceous or subcoriaceous. Inflorescences terminal or axillary, 1–3- or up to 15-flowered. Flowers pendant; calyx campanulate; corolla tubular-infundibuliform or broadly campanulate; stamens somewhat expanded or forming a stapet when inserted basally to corolla; anthers basifixed; nectary annular; stigma bilobed. Berry globose, usually with thick stone cells or with one or two many-seeded pyrenes, pericarp thick, coriaceous, calyx not accrescent. Seeds up to 35, embryo strongly bent; endosperm rather abundant. 2n = 24.

Two spp. from Venezuela to Bolivia, at 2500–3800 m.

Although this genus is morphologically well-defined, its monophyly is still uncertain (Smith and Baum 2006).

  1. 47.

    Vassobia Rusby

    Vassobia Rusby, Bull. New York Bot. Gard. 4: 422 (1927); Hunziker, Kurtziana 17: 91–118 (1984), rev.

Shrubs or small trees, with cauline spines. Leaves large. Inflorescence terminal, a cymose corymb, ca. 50-flowered, or axillary, fasciculate, ca. 8-flowered. Flowers inodorous, 5-merous; calyx urceolate in bud, irregularly lobed at anthesis; corolla rotate or campanulate-rotate, tube with 5 plaits and glabrous inside; stamens equal, stapet auriculate; anthers basi- or dorsi-basifixed; nectary included; stigma bilobate. Berry juicy, pericarp coalescent with placenta, stone cells many, calyx scarcely accrescent. Seeds discoid; embryo annular. n = 12.

Two spp., Bolivia, eastern Paraguay, southern Brazil, northern Argentina and north-western Uruguay.

  1. 48.

    Eriolarynx (Hunz.) Hunz.

    Eriolarynx (Hunz.) Hunz., Kurtziana 28: 66, f. 1 (2000).

    Vassobia Rusby sect. Eriolarynx Hunziker, Kurtziana 10: 24 (1977), ibid. 17: 91–118 (1984), rev.

Small shrubs or trees. Leaves glabrous to pubescent. Inflorescences axillary, sessile, fasciculate, up to 4-flowered or flowers solitary, pendant. Calyx urceolate in bud, later campanulate, 5-apiculate or shortly 5-lobed; corolla campanulate-rotate or subinfundibuliform-rotate, with a ring of hairs inside, lobes 5, short, aestivation valvate-plicate; stamens unequal, glabrous, stapet with incipient auricles; anthers basi-dorsifixed; nectary slightly protruding; stigma shortly saddle-shaped. Berry juicy, pericarp coalescent with the placenta, stone cells abundant, calyx accrescent. Embryo slightly curved or annular, cotyledons as long as rest of embryo. n = 12.

Three spp., one in the Bolivian Andes, the others in north-western Argentina.

Eriolarynx might be monophyletic after inclusion of Iochroma australe Griseb. (Smith and Baum 2006).

  1. 49.

    Acnistus Schott Fig. 65

    Fig. 65.
    figure 10

    Solanaceae. Acnistus arborescens. A Part of branch with flowering brachyblast. B Flower bud showing entire calyx. C Flower. D Inside view of a corolla sector. E Anther, dorsal view. F Anther, ventral view. G Dorsal view of anther base, showing insertion of filament. H Gynoecium from a bud. I Gynoecium from a flower at anthesis. J Fruit. K Transverse section of fruit. L Stone cell. M Seed. N Seed, longitudinal section. O Embryo. (Hunziker 2001; drawn by N. Flury)

    Full size image

    Acnistus Schott, Wiener Z. Kunst. 1829(4): 1180 (1829); Hunziker, Kurtziana 15: 81–102 (1982), rev.

Shrubs or small evergreen trees; indumentum of simple and branched trichomes. Leaves entire, decurrent. Inflorescences axillary, many-flowered fascicles. Flowers small, fragrant, pedicellate, 5-merous; calyx urceolate in bud, torn into somewhat unequal lobules at anthesis; corolla campanulate-infundibuliform; stamens inserted in lower half of corolla; anthers dorsifixed, exserted; nectary annular, prominent; stigma broad, discoidal-capitate. Berry many-seeded, juicy, stone cells rather scanty; calyx not accrescent. Seeds subreniform; embryo strongly bent. n = 12; 2n = 24.

A single species, A. arborescens (L.) Schltdl., southern Mexico to eastern Brazil and Paraguay. A redefinition of Acnistus will be necessary if some Iochroma spp. are transferred to this genus (Smith and Baum 2006).

  1. IV.5.c.

    Subtribe Physalidinae Reveal (2012).

Corolla with an internal ring of trichomes, aestivation valvate or contorted. Calyx normally accrescent enveloping partly or entirely the berry.

  1. 50.

    Witheringia L’Hérit.

    Witheringia L’Hérit., Sert. angl.: 1, 33, tab. I (1788); Hunziker, Kurtziana 5: 101–153 (1969), rev.; Sousa-Peña, M., Systematics and reproductive biology of the genus Witheringia L’Her. (Solanaceae), PhD thesis, University of Connecticut: 1–197 (2001), rev.

Herbs or shrubs; stem glabrous or with simple or branched trichomes. Leaves geminate, unequal in size. Inflorescences axillary, many-flowered, peduncle short or long. Perianth 4–5-merous; calyx cyathiform, truncate; corolla in general 5-parted, tube shorter than elongated lobes, usually with an internal ring of trichomes near stamen insertion, aestivation valvate; stamens equal or not, inserted in upper half of corolla; anthers dorsifixed or dorsi-basifixed; nectary annular; stigma discoidal-depressed or subglobose. Berry red or yellow, stone cells 2–10, calyx sometimes slightly accrescent. Seeds reniform or subreniform; embryo strongly bent. n = 12; 2n = 24.

About 11 spp., southern Mexico to Bolivia.

  1. 51.

    Brachistus Miers

    Brachistus Miers, Ann. Mag. Nat. Hist. II, 3 (16): 262 (1849); Hunziker, Kurtziana 5: 153–163 (1969), rev.

Shrubs or small trees, frequently anisophyllous. Inflorescences axillary, many-flowered fascicles, or flowers solitary, small, odourless. Calyx margin 5-angulate or with 5 triangular teeth; corolla with a campanulate tube and an inner ring of trichomes, segments longer than broad, aestivation valvate; stamens inserted at middle of corolla, stapet not auriculate; anthers dorsi-basifixed; nectary annular. Berry red or orange, many-seeded, calyx strongly or slightly accrescent, pericarp fleshy, stone cells few; embryo annular.

Three spp. in Mexico and Central America. Brachistus is a poorly known genus.

  1. 52.

    Leucophysalis Rydb.

    Leucophysalis Rydb., Mem. Torrey Bot. Club 4: 365 (1896).

Annual or perennial herbs, pubescent, anisophyllous. Inflorescences axillary, 1–2-flowered. Flowers showy, pedicels reflexed to pendant; calyx 5-parted, segments unequal; corolla rotate or shallowly bell-shaped, 5-angled or faintly 5-lobuled, inside with hairy ring, aestivation valvate-plicate; stamens equal or unequal, inserted basally; anthers basifixed or dorsi-basifixed; nectary annular; stigma very shortly saddle-shaped. Berry green, ovoid or globose, pericarp with abundant stone cells, calyx accrescent, tightly or loosely appressed to berry, partially or completely enclosing it. Seeds many, discoid or reniform; embryo subcoiled or annular. n = 12; 2n = 24.

Two spp., Canada and USA.

  1. 53.

    Calliphysalis Whitson

    Calliphysalis Whitson, Rhodora 114: 137 (2012).

Erect perennial herbs with taproot, viscid pubescent. Leaves alternate and geminate, long petiolate. Inflorescence axillary in fascicle of 2–6 flowers. Flowers pendant, actinomorphic, 5-merous. Calyx campanulate, shallowly 5-lobed; corolla broadly campanulate, with five patches of olive-green to mustard-yellow spots at the interior base; stamens inserted at the corolla base, its maturation staggered; anthers basifixed; ovary with many ovules, nectary annular, stigma capitate. Berry mustard-yellow, entirely hidden by the non-angled and scarcely papery calyx. Seeds discoid to reniform; stone cells numerous. n = 12.

Monotypic, C. carpenteri (Riddell) Whitson, south-eastern USA.

  1. 54.

    Oryctes S. Watson

    Oryctes S. Watson, Botany (fortieth parallel): 274, pl. 28, f. 5–10 (1871).

Very small viscid-pruinose annuals. Leaves entire, slightly undulate. Flowers in fascicles of 2–5 in axils, on short peduncles. Calyx zygomorphic, campanulate, with unequal segments; corolla tubular, purple, lobes short, deltoid, slightly unequal, as long as broad, aestivation valvate-conduplicate; androecium heteromorphic: 2 longest stamens exserted or equalling the corolla, filaments inserted basally; anthers basifixed, thecae slightly unequal; nectary included; stigma discoidal-depressed. Capsule almost as long as accrescent, faintly veined calyx. Seeds strongly compressed, with a hyaline peripheral wing; embryo coiled, cotyledons incumbent; endosperm scanty.

Monotypic, O. nevadensis S. Watson, western North America.

  1. 55.

    Quincula Raf.

    Quincula Raf., Atl. J. 1: 145 (1832); Barboza, Kurtziana 28 (1): 69–79 (2000), rev.

Herbs with dense indumentum of water-containing, vesicular, whitish trichomes. Leaves pinnatifid or pinnatipartite. Flowers axillary, up to five. Calyx campanulate; corolla rotate, lobes inconspicuous, inside with five pads of branched trichomes; filaments inserted basally, anthers basi-ventrifixed; nectary inconspicuous; style curved. Berry with few seeds, completely enveloped by the inflated calyx with invaginated base, pericarp thin, thickness uneven, easily breaking into irregular pieces. Seed margin clearly irregular or somewhat crenate; embryo coiled, endosperm abundant. n = 11, 12; 2n = 20, 22, 44.

One species, Q. lobata (Torrey) Raf., south-western USA to northern Mexico.

  1. 56.

    Chamaesaracha (A. Gray) Benth. & Hook.

    Chamaesaracha (A. Gray) Benth. & Hook., Gen. pl. 2: 891 (1876); Averett, Rhodora 75: 325–365 (1973), rev.

Perennial ascending or spreading herbs, glabrous to variously pubescent. Leaves variously shaped. Flowers axillary 1–2, 5-merous. Calyx lobed; corolla rotate, slightly lobed, with white tomentose pads in the throat, aestivation contorted-plicate; stamens inserted basally; anthers dorsifixed; nectary strongly developed; stigma bilobed. Fruit a dry, green berry, tightly and partially enclosed by the accrescent calyx, pericarp thin. Seeds up to 40, testa rugose-reticulate; embryo subcoiled. n = 12, 18, 24, 36.

Small, mainly Mesoamerican genus with ca. 10 spp.

  1. 57.

    Physalis L.

    Physalis L., Sp. pl. 1: 182 (1753); Benítez de Rojas & Magallanes, Acta Bot. Venez. 21(2): 11–42 (1998), reg. rev.; Martínez, Anales Inst. Biol. Univ. Nac. Autón. México, Bot. 69(2): 71–117 (1998), part. rev.

    Margaranthus Schltdl. (1838).

Herbs, rarely shrubs; taproot thick or roots gemmiferous; stems occasionally viscid; indumentum varied, of simple or branched, non-glandular or glandular trichomes. Leaves entire or pinnatifid. Flowers solitary, pendant. Calyx invaginate; corolla rotate or campanulate-rotate, rarely urceolate, aestivation contorted-plicate, in general with a dense hairy ring inside; anthers basifixed, connective broad; nectary included; stigma shortly saddle-shaped. Berry many-seeded, completely enclosed by inflated calyx, reticulate-veined, strongly urceolate. Seeds enveloped by parenchymatic cells of placental and pericarpic origin; embryo subcoiled or coiled. n = 12, 24, 36; 2n = 24, 48, 72.

About 90 spp., mostly American (Mexico, USA and Central America, reaching South America), except the Eurasian P. alkekengi L.

A recent molecular study (Whitson and Manos 2005) revealed paraphyly of Physalis in its current circumscription which calls for taxonomic changes to correct the paraphyly.

  1. IV.5.d.

    Physalideae: Incertae sedis

  1. 58.

    Cuatresia Hunz.

    Cuatresia Hunz., Kurtziana 10: 15, f. 4 (1977); Hunziker, Opera Botanica 92: 73–82 (1987), rev.

Shrubs or small trees. Leaves geminate, strongly anisophyllous. Flowers 2–45, in sessile or subsessile axillary inflorescences or on long peduncle; calyx cup-shaped or urceolate, margin almost entire at anthesis, or obscurely divided at fruiting stage; corolla usually glabrous within, campanulate or infundibuliform, 5-lobed or 5-parted, aestivation valvate-induplicate; stamens equal; anthers ventrifixed; nectary included or slightly prominent, stigma shortly saddle-shaped. Berry juicy, pericarp lacking stone cells, calyx not accrescent or investing the entire berry. Seeds reniform, discoid; embryo strongly curved; endosperm very abundant.

About 15 spp., Guatemala to eastern Bolivia.

  1. 59.

    Larnax Miers

    Larnax Miers, Ann. Mag. Nat. Hist., ser. II, 4: 37 (1849); Sawyer, Monogr. Syst. Bot. 104: 259–285 (2005); Leiva & Barboza, Arnaldoa 16: 29–36 (2009), key.

Herbs, shrubs or small trees. Inflorescence axillary, fascicles up to 30-flowered; calyx shortly apiculate or toothed, exceptionally with long segments; corolla mostly stellate, variously coloured, lobes 2–4 times longer than tube, aestivation valvate; androecium heterodynamous or homodynamous, filaments broadened at their insertion into a conspicuous or inconspicuous stapet with or without auricles; anthers basifixed; nectary conspicuous; stigma discoid, saddle-shaped or bilobed. Berry completely enveloped by the accrescent calyx, calyx tightly appressed to berry or inflated and loosely surrounding it. Seeds up to ca. 150, reniform; embryo strongly bent, cotyledons sometimes very short.

About 32 spp., Central America to Peru. A taxonomic revision is needed.

  1. 60.

    Deprea Raf.

    Deprea Raf., Sylva tellur.: 57 (1838); Barboza & Hunziker, Kurtziana 23: 45–65 (1994), rev., Sawyer, Monogr. Syst. Bot. 104: 259–285 (2005).

Herbs or shrubs. Leaves rarely geminate. Flowers generally in fascicles of 1–7, perfect or exceptionally functionally unisexual. Calyx tube 2.5–3 times longer than unequal lobes; corolla narrowly campanulate, funnel-shaped or urceolate, lobes always shorter than tube, aestivation valvate; filament base almost always with an inconspicuous stapet; anthers basifixed; nectary conspicuous; stigma discoid or saddle-shaped. Berry subglobose or ellipsoidal, loosely or tightly enveloped by the strongly accrescent calyx. Seeds subreniform; embryo strongly bent, cotyledons shorter than rest of embryo.

Ten spp., Venezuela to Bolivia.

  1. 61.

    Schraderanthus Averett

    Schraderanthus Averett, Phytologia 91: 54 (2009).

Erect herbs or soft-wooded shrubs, viscid, glandular-pubescent throughout. Leaves geminate, anisophyllous. Inflorescences axillary, fasciculate, 6–8(–10)-flowered. Calyx deeply 5-parted, exceeding length of corolla, segments acute; corolla rotate and reflexed, 5-lobed to parted, inside with a hairy ring and with greenish spots in the throat, aestivation valvate-plicate; stamens unequal, exserted, attached basally to corolla, anthers bluish, dorsi-basifixed. Berry globose, fleshy, red to orange-red, calyx accrescent, broadly campanulate, exceeding berry but becoming reflexed at maturity. Seeds ca. 50–75, reniform; embryo subcoiled, endosperm abundant. 2n = 24 (sub nom. Leucophysalis viscosa).

Monotypic, S. viscosus (Schrad.) Averett, Mexico, Guatemala, Venezuela.

  1. 62.

    Tzeltalia E. Estrada & M. Martínez

    Tzeltalia E. Estrada & M. Martínez, Brittonia 50: 289 (1998).

Weak shrubs; stem fistulose, glabrous to densely villose. Leaves alternate or geminate, petiole winged. Inflorescence axillary (1)2–7-flowered fascicles. Calyx thin-chartaceous, glabrous at anthesis, truncate or lobed; corolla rotate, whitish or cream-white usually with 5 purplish-green or olive green spots near the base of the lobes, inside throat and base of lobes with 5 more or less confluent pads of branched trichomes surrounding and above the filaments insertion, lobes 2–3 times longer than tube; filaments blue or purplish; anthers subdorsifixed. Berry greenish, calyx accrescent, open at mouth and not invaginate at base, strongly 5- to 10-angled and strongly reticulate-veined. Seeds reniform, alveolate.

Three spp., southern Mexico and Guatemala.

  1. 63.

    Capsicophysalis (Bitter) Averett & M. Martínez

    Capsicophysalis (Bitter) Averett & M. Martínez, J. Bot. Res. Inst. Texas 38(1): 72 (2009).

Annual or weak perennial herbs. Leaves petiolate, thin to membranous. Flowers axillary 1–2. Calyx campanulate, rounded at the base; corolla campanulate-subrotate, unequally 5-lobed, irregular, white, yellowish or yellow-green with villous pads in the throat; filaments equal, anthers basifixed. Berry fleshy, shiny red or orange-red, calyx accrescent, at first loosely investing the berry, then splitting and becoming reflexed below the berry. Seeds discoidal, testa tuberculate with rod-like projections.

One species, C. potosina (B.L. Rob. & Greenm.) Averett & M. Martínez, southern Mexico to Guatemala and Honduras.

  1. IV.6.

    Tribe Capsiceae Dumort. (1827).

Shrubs, trees, or perennial herbs. Flowers solitary or in fascicle pauciflowered. Calyx entire, truncate, with 0–10 linear nerve prolongations (teeth); corolla rotate or stellate, rarely campanulate, aestivation valvate. Calyx not accrescent. Endosperm copious.

  1. 64.

    Lycianthes (Dunal) Hassl.

    Lycianthes (Dunal) Hassl., Ann. Conserv. Jard. Bot. Genève 20: 180 (1917), nom. cons.; Bitter, Abh. Naturwiss. Verein Bremen 24(2): 292–520 (1919), rev.; Barboza & Hunziker, Darwiniana 31(1–4): 17–34 (1992), reg. rev.; Benítez de Rojas & D’Arcy, Ann. Missouri Bot. Gard. 84: 167–200 (1997), reg. rev.; Dean, Bot. J. Linnean Soc. 145: 385–424 (2004), part. rev.

Shrubs, trees, vines or perennial herbs. Leaves various. Flowers axillary or extra-axillary; calyx with (0)5–10 linear teeth; corolla rotate, apically 5-angled or deeply divided; androecium usually heterodynamous, filaments fused in a ring adnate to the basal edge of corolla; anthers basifixed, tightly coherent or connate in a column, dehiscing with a terminal pore; nectary absent; stigma discoid or saddle-shaped. Berry sometimes with stone cells or occasionally with 1–2-seeded pyrenes. Seeds discoid or subreniform; embryo coiled, cotyledons as long as rest of embryo. n = 12; 2n = 24.

American-Asian genus of ca. 150 spp. or more, mostly neotropical.

  1. 65.

    Capsicum L.

    Capsicum L., Sp. pl.1: 188 (1753); Hunziker, Darwiniana 36: 201–203 (1998), reg. rev.; Barboza et al., Syst. Bot. 36 (3): 768–781 (2011), key.

Shrubs, herbs or trees. Leaves entire, alternate or geminate. Flowers axillary, solitary or in fascicles of 2–9 (18), pedicels usually geniculate distally. Calyx in general with (0)5–10 linear-compressed teeth; corolla rotate, stellate, campanulate, or campanulate-urceolate, lobes usually shorter than rest of corolla, aestivation valvate, tube with plaits; stamens usually equal, filament basis with two lateral, short and thick auricles fused to the corolla (except C. chacoense); anthers dorsifixed; nectary included; style sometimes heteromorphic, stigma bilobed. Berry usually with capsaicinoids in the epidermis of the septum and adjacent areas and consequently hot. Seeds many; embryo annular. n = 12, 13; 2n = 24, 26.

About 32 spp., southern USA to central Argentina. The cultivated peppers belong to five spp., the most important is C. annuum L. with many cultivars.

  1. IV.7.

    Tribe Solaneae Dumort. (1829).

Herbs, shrubs or trees. Corolla rotate, campanulate, stellate, infundibuliform, tubular, urceolate; aestivation valvate; stamen usually equal in length, markedly enlarged at base where adnate to base of corolla; anthers dehiscing longitudinally or by pores then expanding into a longitudinal slit; stigma capitate or bilobed; calyx slightly or strongly accrescent.

  1. 66.

    Jaltomata Schltdl.

    Jaltomata Schltdl., IndexSem. (Halle): 8 (1838); Mione, Anderson & Nee, Brittonia 45: 138–145 (1993), part. rev.; Mione, Leiva G. & Yacher, Taxon 17: 49–58 (2007).

    Hebecladus Miers (1845).

Perennial herbs or shrubs. Leaves entire, repand or toothed. Inflorescence axillary or terminal dichasia or umbellate, pedunculate; flowers solitary, 2–3 or up to 12–41, pendant, usually protogynous; pedicels basally articulated. Calyx 5-lobed; corolla rotate, campanulate, urceolate, crateriform or tubular, 5-lobed, or lobes alternating with lobules totalling 10, aestivation valvate or valvate-induplicate; androecium usually homomorphic, filaments enlarged at insertion, stapet usually swollen and hairy; anthers ventrifixed; nectary well developed, nectar orange, yellowish, or red. Berry frequently edible, many-seeded, calyx strongly accrescent, rotate to reflexed, never enclosing the berry. Seeds small; embryo strongly bent; endosperm abundant. n = 12; 2n = 24, 48.

Ca. 60 spp. in two centres of diversity: south-western USA and Mexico (J. antillana (Krug & Urban) D’Arcy in Greater Antilles), and western South America from Venezuela and Colombia to Bolivia (J. werfii D’Arcy in Galápagos).

  1. 67.

    Solanum L.

    Solanum L., Sp. pl. 1: 184 (1753); Hawkes & Hjerting, The potatoes of Argentina, Brazil, Paraguay and Uruguay: 1–525 (1969), reg. rev.; Morton, A revision of the Argentine species of Solanum: 1–260 (1976), reg. rev.; Whalen, Gentes Herb. 12: 179–282 (1984); Francisco-Ortega et al., Pl. Syst. Evol. 185: 189–205 (1993), rev.; Bohs, Fl. Neotrop. Monogr. 63: 1–175 (1994), part. rev.; Knapp, Fl. Neotrop. Monogr. 84: 1–404 (2002), part. rev.; Spooner et al., Syst. Bot. Monogr. 68: 1–209 (2004), reg. rev.; Peralta, Spooner & Knapp, Syst. Bot. Monogr. 84: 1–186 (2008), part. rev.

    Lycopersicon Mill. (1754).

    Melongena Mill. (1754).

    Bassovia Aubl. (1775).

    Triguera Cav. (1786), nom. cons.

    Androcera Nutt. (1818).

    Cyphomandra Sendtn. (1845).

    Normania Lowe (1872).

    Pheliandra Werderm. (1940).

Herbs to trees, rarely vines, sometimes tuber-bearing, glabrous or with a profuse variety of trichomes, sometimes prickly. Leaves alternate or paired, frequently unequal, simple or compound, sometimes with pseudostipules. Inflorescences terminal, axillary or extra-axillary, cymes branched or unbranched, in racemose, paniculate or scorpioid structures. Flowers sessile or on pedicels, articulated or not, actinomorphic or zygomorphic, usually perfect, rarely unisexual, rarely dioecious. Perianth 4–5-merous, occasionally 4–6(10)-merous; calyx cyathiform or variously divided; corolla rotate, campanulate-urceolate, stellate, infundibuliform or urceolate, aestivation valvate or valvate-induplicate; stamens equal or unequal, filaments basally fused in a ring adnate to corolla tube, sometimes also into an upper ring free from the corolla; anthers equal or unequal, connivent or not, basifixed to basi-ventrifixed, exclusively poricidal or with a terminal pore expanding into a longitudinal slit, or anthers opening introrsely with longitudinal slits, sometimes with a thick glandular excrescence at the base or with a prolonged and sterile apical sector; nectary absent. Fruit a berry, sometimes dry and dehiscent, stone cells usually present, calyx slightly or strongly accrescent. Seeds often flattened; embryo coiled, endosperm abundant. n = 11, 15; 2n = 24, 48, 72, 96.

Cosmopolitan; ca. 1400 spp., mainly American. Genus of worldwide economic importance (see section “Economic Importance”).

Phylogenetic studies in this genus have included increasingly more species in recent years. Currently, 13 major clades are recognized (Bohs 2005; Weese and Bohs 2007; Stern et al. 2011).

  1. IV.8.

    Solanoideae: Incertae sedis

  1. 68.

    Latua Phil.

    Latua Phil., Bot. Zeitung (Berlin) 16 (33): 241 (1858).

Shrubs or trees with heteroblastic growth and cauline spines. Leaves fasciculate on short shoots or scattered on long shoots. Flowers solitary, pedicels tomentose, arising in the axil of a spine from an involucre of scales; calyx campanulate, lobes equal; corolla actinomorphic, densely pilose throughout, aestivation cochlear-conduplicate; stamens 5, filaments unequal, adnate basally to corolla tube; anthers basifixed; nectary annular. Berry globose, yellow or orange-yellow, calyx accrescent, splitting irregularly. Seeds numerous, thick; embryo bent, cotyledons shorter than rest of embryo; endosperm abundant. 2n = 24.

A single species, L. pubiflora (Griseb.) Baill., coastal mountains of southern Chile.

  1. 69.

    Jaborosa Juss.

    Jaborosa Juss., Gen. pl.: 125 (1789); Barboza & Hunziker, Kurtziana 19: 77–153 (1987), rev.

    Trechonaetes Miers (1845).

Perennial herbs, frequently with gemmiferous roots and rhizomes. Leaves entire to pinnatisect. Flowers in 1–7-flowered fascicules or in dense glomerules (ca. 100 or more flowers). Calyx campanulate, 5-lobed; corolla tube cylindrical, rotate, campanulate-rotate, urceolate or hypocrateriform, pubescent inside, aestivation valvate-induplicate or valvate-conduplicate; stamens 5, usually included, inserted at different levels on corolla, bent towards the centre of the flower, tip inflexed; ovary bicarpellate, rarely 3–5-carpellate, nectary annular; stigma lobulate or branched. Berry globose-depressed, calyx accrescent. Seeds up to ca. 90; embryo annular; endosperm copious. n = 12; 2n = 24.

22 spp., southern Peru to Argentina.

  1. 70.

    Sclerophylax Miers Fig. 66

    Fig. 66.
    figure 11

    Solanaceae. A–D Sclerophylax arnottii. A Flowering branch. B Flower. C Calyx. D Fruiting calyx. E–H Sclerophylax ruiz-lealii. E Flowering branch. F Flower. G Calyx. H Fruiting calyx. (Di Fulvio 1961; drawn by H. García)

    Full size image

    Sclerophylax Miers, London J. Bot. 7: 18 (1848); Di Fulvio, Kurtziana 1: 9–103 (1961), rev.

Branched annual or perennial herbs, indumentum eglandular or glandular or plants subglabrous. Two entire leaves per node, generally not opposite, often anisophyllous. Flowers solitary (rarely paired) in leaf axils, 5-merous; sepals united at base, unequal, accrescent with lobes becoming more or less thorny in fruit; corolla white or violet, infundibuliform, slightly zygomorphic, deciduous, lobes inconspicuous, aestivation contorted-conduplicate; stamens included; anthers dorsifixed; ovary bicarpellate, bilocular; placentation apical, ovules apotropous, pendant, 1–2 per locule; style terminal, stigma simple. Fruit dry, indehiscent, generally one seed per locule, pericarp membranous included in the accrescent and sclerified calyx (diclesium). Seeds with aleuron, exarillate; embryo straight or curved; endosperm oily. n = 12; 2n = 24.

14 spp. mainly in Argentina, also in Uruguay and Paraguay.

  1. 71.

    Nolana L.f. Fig. 67

    Fig. 67.
    figure 12

    Solanaceae. Nolana lezamae. A Flowering branch. B Flower. C Floral trichome. D Calyx with connate lobes. E Dissected corolla. F Anther, ventral view. G Anther, lateral view. H Anther, dorsal view. I Gynoecium. J Calyx expanding with mericarps. K Mericarps. (Drawn by S. Leiva García)

    Full size image

    Nolana L.f., Dec. pl. horti upsal.1: 3 (1762).

Annuals to shrubs, sometimes with basal rosettes from taproots. Leaves succulent, terete to laminar. Flowers 5-merous; calyx regular, actinomorphic, to bilabiate, zygomorphic, enclosing the ovary; corolla regular, or more commonly irregular to weakly zygomorphic, infundibuliform, campanulate or tubular, more rarely suburceolate or salver-form; stamens adnate basally to the corolla tube, often pubescent, unequal (3 long, 2 short); ovary 5-carpellate. Fruit a schizocarp, mericarps 2–30, highly variable in shape, from polygons to spheres, and external sculpturing smooth to rugulose. Seeds with copious oily endosperm. 2n = 24.

89 species in Chile, Peru and one in the Galapagos Islands (Ecuador). The genus has been traditionally divided into subgenera Alona and Nolana; however, recent studies have shown that additional monophyletic groups are in need of formal classification (Dillon, pers. obs.).

  1. 72.

    Exodeconus Raf.

    Exodeconus Raf., Sylva tellur.: 57 (1838); Axelius, Pl. Syst. Evol. 193: 153–172 (1994), rev.

    Cacabus Bernh. (1839).

Small annual viscid herbs. Leaves entire to sinuate-dentate. Flowers solitary, 5-merous. Calyx tubular, infundibuliform, or cyathiform to urceolate, deeply divided; corolla tubular-campanulate or infundibuliform with a very narrow base, aestivation contorted-induplicate; stamens included, usually heterodynamous, filaments unequal, inserted basally; anthers basifixed; nectary annular, inconspicuous; style terminal, stigma much depressed. Berry sometimes sclerified apically to in upper half, base with sclereids and stone cells, calyx accrescent, innervation very conspicuous. Seeds up to ca. 300; embryo bent, cotyledons shorter than rest of embryo. 2n = 24.

Six spp. mostly in southern Ecuador, Peru, northern Chile and northern Argentina.

  1. 73.

    Nicandra Adans.

    Nicandra Adans., Fam. pl. 2: 219 (1763), nom. cons.; Carrizo García, Flora Fanerog. Argent. 88: 3–4 (2005), reg. rev.

    Physalodes Boehm. (1760), nom. rej.

Annual herbs; stems hollow. Leaves generally sinuate. Flowers extra-axillary, solitary, 5-merous.Calyx aestivation reduplicate-valvate, segments auriculate or cordate at base and fused almost to middle; corolla broadly campanulate, aestivation cochlear-plicate; stamens geniculate, incurved; anthers basifixed, connivent; ovary 3–5-carpellate, nectary included, crenulate; stigma capitate-lobed. Berry pendant, pericarp almost dry, semitranslucent, easily ruptured, stone cells present, calyx strongly accrescent, chartaceous, inflated, with conspicous venation. Seeds discoidal-reniform; embryo coiled, cotyledons shorter than rest of embryo; endosperm abundant. n = 10; 2n = 19, 20, 21, 40.

Three spp., two endemic to Peru and N. physalodes (L.) Gaertn. from Peru to northern Argentina; this species is an ornamental and ruderal in tropical and subtropical areas throughout the world.

  1. 74.

    Schultesianthus Hunz.

    Schultesianthus Hunz., Kurtziana 10: 35, f. 9 (1977); Bernardello & Hunziker, in Hawkes et al. (eds.) Solanaceae III. Taxonomy, Chemistry, Evolution: 1–17 (1991), rev.

Climbing epiphytes or scandent shrubs. Leaves coriaceous, variously shaped, abundantly punctate below. Inflorescence terminal cymes, 2–10-flowered, occasionally reduced to a single flower. Flowers showy, fragrant, 5-merous; calyx usually actinomorphic, lobes thick, coriaceous; corolla zygomorphic, funnel-form to strongly infundibuliform, lobes auriculate, aestivation cochlear; stamens declinate or straight, inserted basally and pubescent at point of insertion; anthers basifixed; nectary annular, slightly exserted; style declinate or straight. Berry globose or ovoid; calyx accrescent, woody. Seeds ca. 30, reniform, large; embryo strongly annular, cotyledons accumbent, as broad as long and not wider than rest of embryo; endosperm scarce.

Eight spp., southern Mexico to Bolivia.

Knapp et al. (1997) defined the genus more broadly and included three species recognized under Markea by Hunziker (1997, 2001).

  1. 75.

    Trianaea Planch. & Linden

    Trianaea Planch. & Linden in Linden, Cat. pl. exot.: 717 (1853).

Shrubs or lianas, usually epiphytic, sometimes with a woody tuberosity, exceptionally small trees. Leaves thick, shiny. Flowers terminal or axillary, solitary or in groups of 2–3, showy, 5- or 4-merous, pendant. Calyx campanulate; corolla campanulate, large and fleshy, tube short, lobules rounded, reflexed, aestivation cochlear; stamens strongly geniculate at base and attached basally; anthers ventrifixed; ovary 4- or 5-carpellate, nectary prominent; stigma capitate to clavate. Berry large (up to 8 cm diam.), surrounded by the accrescent calyx. Seeds reniform, elongated, thin; embryo curved, cotyledons oblique, longer than broad; endosperm rather abundant.

About six spp., north-western South America from Colombia to northern Peru.

  1. 76.

    Solandra Sw.

    Solandra Sw., Kongl. Vetensk. Acad. Nya Handl. 8: 302, tab. XI (1787), nom. cons.; Bernardello & Hunziker, Nordic J. Bot. 7: 639–652 (1987), rev.

Climbing epiphytic shrubs or lianas or small trees, glabrous or pubescent. Leaves coriaceous or subcoriaceous, lustrous. Flowers solitary, terminal, usually zygomorphic; calyx tubular-campanulate; corolla large, limb short, tube cylindrical basally and expanded apically, aestivation cochlear; stamens equal, declinate, attached near middle of corolla or more basally; anthers basifixed; ovary bicarpellate, tetralocular, ∞-ovuled, partly inferior, nectary annular; style declinate, stigma discoidal-capitate. Berry ∞-seeded. Seeds discoid or reniform, compressed; embryo subcoiled, cotyledons incumbent or slightly oblique, shorter than rest of embryo; endosperm abundant. n = 12; 2n = 24.

10 spp., Mexico and West Indies to Peru, Bolivia, and Brazil; some species are widely cultivated as ornamentals, such as S. grandiflora Sw., S. longiflora Tussac, and S. maxima (Sessé & Mociño) P.S. Green.

  1. 77.

    Mandragora L.

    Mandragora L., Sp. pl. 1: 181 (1753); Ungricht, Knapp & Press, Bull. Nat. Hist. Mus. London, Bot. 28: 17–40 (1998), rev.

Perennial rosette-forming herbs, with fleshy, thick taproots. Leaves rugose-bullate abaxially. Flowers 5(6)-merous; calyx deeply lobed; corolla campanulate, divided to almost half its length or more, aestivation cochlear; stamens homomorphic, incurved towards the centre of the flower and attached basally, filaments with a conspicuous basal tuft of trichomes; anthers dorsifixed, connective thick and fleshy; nectary prominent, two-lobed; stigma capitate. Berry globose, with a pleasant odour at maturity, later odour heavy and disagreeable, calyx accrescent. Seeds up to 50, reniform, large; embryo coiled, cotyledons unequal, longer than rest of embryo; endosperm scanty. 2n = 24, 48, 84, 96.

Three spp., disjunct in Mediterranean Europe, central Asia, and the Sino-Himalaya region.

  1. 78.

    Nectouxia Kunth

    Nectouxia Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 3: 10, tab. 193 (1818).

Hemicryptophytic herbs, somewhat fetid; rhizome fleshy with minute cataphylls; stems glandular-pubescent. Leaves cordate. Flowers solitary, pendant. Calyx divided almost to base into equal segments, densely hirsute; corolla yellow or yellow-green, hypocrateriform, aestivation valvate-induplicate, with a ring-like corona at junction of tube and limb; stamens included or subincluded, inserted distally, thecae dorsally with a laminar enlargement of the filament apex; nectary rather prominent; stigma discoid. Berry usually apiculate. Seeds up to 10; embryo coiled; endosperm rather scanty.

One species, N. formosa Kunth, Mexico.

  1. 79.

    Salpichroa Miers

    Salpichroa Miers, London J. Bot. 4: 324 (1845); Keel, PhD thesis, University of New York: 1–196. University Microfilms International, Michigan, USA (1984), rev.; Barboza & Hunziker, Fl. Fanerog. Argent. 54: 19–24 (1998), reg. rev.

Scandent, pendant, prostrate or straggling shrubs. Leaves solitary or geminate. Flowers solitary, rarely in pairs, pendant. Calyx 5-parted almost to base or with a tube as long as segments; corolla cylindrical, urceolate, or urceolate in basal half and cylindrical upwards, aestivation valvate or slightly valvate-induplicate, inside normally glabrous (not S. origanifolia); stamens inserted at various levels in upper half of corolla, filament apex usually spathulate; anthers dorsifixed, frequently connivent; nectary prominent; stigma discoid. Berry elongated. Seeds up to 28; embryo subcoiled; endosperm rather scanty. n = 12; 2n = 24.

16 spp., Venezuela to north-western Argentina and Chile; S. origanifolia (Lam.) Baillon naturalized in the USA, Europe, North Africa, and Australia.

  1. 80.

    Darcyanthus Hunz. Fig. 68

    Fig. 68.
    figure 13

    Solanaceae. Darcyanthus spruceanus. A Flowering branch. B, C Glandular trichomes. D Flower. E Inside view of a corolla sector. F Anther, dorsal view. G Anther, ventral view. H Gynoecium. I Fruit. J Stone cell. K Seed. L Seed, longitudinal section. (Hunziker 2001; drawn by N. Flury)

    Full size image

    Darcyanthus Hunz., Bol. Soc. Argent. Bot. 35: 345 (2000).

Annual sticky herbs, indumentum of long glandular multicellular trichomes. Inflorescences fasciculate, sessile, flowers 7–10, pedicellate, 5-merous. Calyx teeth triangular, unequal, shorter than tube; corolla delicate, rotate or rotate-campanulate, angulate, cream with 5 purple spots within, tube very short; stapet conspicuous, with lateral outgrowths fused to corolla, filaments glabrous, anthers dorsi-basifixed; nectary included, stigma discoidal-capitate. Berry globose, red, pericarp thin, not fleshy, with stone cells, tightly surrounded by accrescent and thin calyx. Seeds small; embryo annular.

Monotypic, D. spruceanus (Hunz.) Hunz., oriental rainforests of Peru and Bolivia.

  1. V.

    Solanaceae: Incertae sedis

Genera not assigned to subfamily

  1. 81.

    Schizanthus Ruiz & Pav.

    Schizanthus Ruiz & Pav., Fl. peruv. prodr.: 6 (1794); Grau and Gronbach, Mitt. Bot. Staatssamml. München 20: 111–203 (1984), rev.; Hunziker and Cosa, Fl. Fanerog. Argent. 74: 3–6 (2001), reg. rev.

Annual or biennial herbs, sometimes woody at base, usually sticky, with non-glandular unicellular trichomes and glandular shaggy hairs. Leaves entire, slightly serrate, lobed to pinnatisect. Inflorescences terminal. Flowers 5-merous, resupinate (twist of ca. 72° at anthesis); calyx segments slightly unequal, linear or spathulate, tube almost absent; corolla zygomorphic, papilionate (segments unequal, deeply laciniate: one anterior, two lateral and the two posterior ones fused forming a keel), aestivation cochlear; 2 fertile stamens in lateral position, and 3 staminodes; anthers dorsifixed, dehiscing explosively; ovary with annular nectary; style filiform, stigma inconspicuous, lacking papillae. Capsule septicidal, 2-valved. Seeds compressed, up to 40, ellipsoidal or reniform; embryo arcuate, cotyledons incumbent, clearly shorter than rest of embryo; endosperm nuclear, copious. n = 10.

A genus of 12 spp., Chile and Argentina (2). Three spp. (S. pinnatus Ruiz & Pav., S. retusus Hook. and S. grahamii Gillies) and a few hybrid strains (S. x wisetonensis Hort., S. hybridus Hort., etc.) are cultivated as ornamentals.

  1. 82.

    Duckeodendron Kuhlmann

    Duckeodendron Kuhlmann, Arch. Jard. Bot. Rio de Janeiro 4: 361, tab. 31 h–l (1925); Martins et al. in Ribeiro et al., Flora da Reserva Ducke 583–587 (1999); Nee in Smith et al., Flowering Plants of the Neotropics: 133–134 (2004).

Trees ca. 30 m high. Leaves entire, coriaceous, pubescent abaxially; petiole canaliculate, with dense ochre indumentum. Inflorescence terminal, cymose, ca. 15-flowered, brownish-pubescent. Flowers subtended by one tiny pilose bract, pedicels articulated; calyx aestivation quincuncial, lobes broad, as long as tube; corolla greenish, tubular, slightly infundibuliform, pubescent inside, lobes rounded, aestivation quincuncial; stamens equal, inserted near middle of corolla; anthers exserted, dorsi-basifixed; ovary bicarpellate, each locule uniovulate; style strongly exserted, stigma minutely bilobed. Drupe 1-seeded. Seed ovoid; embryo U-shaped, cotyledons extremely short; endosperm oily.

Monotypic, central Amazonia (Manaus, Brazil): D. cestroides Kuhlmann.

Tribes not assigned to subfamily

Tribe Benthamielleae Hunz. (2000).

Chamaephytes, small shrubs or perennial microphyllous herbs. Leaves sometimes rigid and spinescent. Flowers subtended by two opposite bracteoles. Stamens usually 5 (rarely 1–3), anthers dorsifixed. Capsule pauciseminate. Embryo curved; endosperm copious.

  1. 83.

    Pantacantha Speg.

    Pantacantha Speg., Anales Soc. Ci. Argent. 53: 171 (1902); Rossow, Fl. Patag. 6: 278–279 (1999), rev.; Hunziker, Fl. Fanerog. Argent. 64: 9–10 (2000), rev.

Low shrubs; stems branched, densely leafy. Leaves sessile, sheathing, linear-subulate to narrow-elliptic, rigid, spinescent. Flowers 5-merous, axillary, actinomorphic, solitary, subtended by two leaf-like bracteoles; calyx campanulate or tubular, teeth equal or subequal, rigid, spiny; corolla greenish-yellow or yellow, cylindrical or slightly infundibuliform, aestivation valvate-induplicate; stamens inserted near base of corolla, filaments equal, pubescent; ovary with prominent annular nectary, ovules ca. 20 per locule; stigma discoid, somewhat bilobate. Capsule septicidal or partly loculicidal. Seeds (1)2–3, angulate-subreniform, strongly ridged or winged; embryo curved, cotyledons shorter than rest of embryo. 2n = 22.

One species, P. ameghinoi Speg., western Argentina.

  1. 84.

    Combera Sandw.

    Combera Sandw. in Hook., Icon. pl. 34, t. 3325 (1939); Rossow, Fl. Patag. 6: 235–237 (1999) reg. rev.; Hunziker, Fl. Fanerog. Argent. 64 (9): 10–11 (2000), reg. rev.

Small, erect perennial herbs; stems densely leafy, partly subterranean. Leaves petiolate, sheathless, spathulate to deltate-ovate or rhombic-ovate, fleshy, closely imbricate. Flowers clustered among the uppermost leaves. Calyx actinomorphic to slightly zygomorphic, 5-dentate, with long unequal teeth; corolla actinomorphic, infundibuliform, aestivation contorted-conduplicate; stamens 5, filaments equal, attached at lower half of corolla; ovary with an included annular nectary, ovules 4–10 per locule. Capsule septicidal or partly loculicidal. Seeds up to 7, embryo curved. 2n = 22.

Two spp., Patagonia (Argentina and Chile).

  1. 85.

    Benthamiella Speg.

    Benthamiella Speg., Anales Soc. Ci. Argent. 15: 109 (1883); Arroyo, Bot. Not. 133: 67–76 (1980), rev.; Arroyo-Leuenberger, Fl. Fanerog. Argent. 64: 4–9 (2000), reg. rev.

    Saccardophytum Speg. (1902).

Pulvinate chamaephytes or lax cushions; stems branched, densely leafy. Leaves sessile, sheathed, membranous or fleshy, closely imbricate. Flowers normally at the end of branches. Calyx actinomorphic, campanulate or tubular, with equal or subequal, acute or truncate lobes; corolla whitish or yellow, actinomorphic, infundibuliform or subcampanulate; aestivation contorted-conduplicate; fertile stamens 1, 2 or 5, staminodes sometimes present, filaments equal or not, attached at 1 or 2 levels on the upper or the lower half of corolla tube; ovules 4–10 per locule. Capsule septicidal or partly loculicidal. Seeds 1–4¸ embryo curved cotyledons shorter than rest of embryo. 2n = 22.

12 spp., southern Patagonia (Argentina and Chile).

Tribe Petunieae Horan. (1847).

Herbs or shrubs. Corolla commonly zygomorphic, funnel-shaped or salver-form, aestivation cochlear, reciprocative or less frequently contorted; androecium usually heterodynamous, stamens fertile 5 or 4 or 2, staminodes present.

  1. 86.

    Fabiana Ruiz & Pav.

    Fabiana Ruiz & Pav., Fl. peruv. prodr.: 18, pl. 34 (1794); Barboza & Hunziker, Kurtziana 22: 109–153 (1993), rev.

Chamaephytes or microphyllous shrubs, of homoblastic or heteroblastic growth; stems and leaves with dense resiniferous indumentum. Leaves sessile, imbricate, fasciculate or rosulate. Flowers solitary, 5-merous, actinomorphic; calyx lobes shorter than tube; corolla whitish, rarely lilac or bluish, funnel-shaped or salverform, aestivation contorted-conduplicate; stamens homodynamous or heterodynamous (2 + 3), filaments usually geniculate; anthers dorsifixed, thecae confluent or free; ovary with a prominent invaginate-bilobed nectary; stigma capitate or saddle-shaped. Capsule septicidal, 2-valved. Seeds ca. 50, polyhedric, angular; embryo straight or slightly curved. n = 9; 2n = 18.

15 spp., western South America, usually Andes. Fabiana imbricata Ruiz & Pav. is an ornamental species and a medicinal plant in Argentina and Chile.

  1. 87.

    Calibrachoa La Llave & Lex.

    Calibrachoa La Llave & Lex., Nov. veg. descr. 2: 3 (1825).

    Leptophragma Benth., in Dunal, De Candolle, Prodr. 13(1): 578 (1852).

Small shrubs, more rarely annual herbs, generally viscid-pubescent; stem usually with brachyblasts. Leaves variously shaped. Flowers in monochasia, apparently solitary and arising from a pair of subopposite equal bracts; pedicels mostly deflected at fruiting time; calyx usually lobed halfway, lobes narrowing towards apex; corolla slightly zygomorphic, funnel-shaped, aestivation reciprocative (the anterior induplicative lobe covers the four others, which are conduplicate); stamens 2 + 2 + 1, filaments sometimes strongly geniculate; anthers ventrifixed; nectary bilobed. Capsules septicidal, 2-valved. Seeds up to 60, reticulate-foveate with straight thick-walled ridges. n = 9; 2n = 18.

26 spp., south-eastern and southern Brazil, Paraguay, Uruguay, and Argentina, C. parviflora (Juss.) D’Arcy disjunct between South and North America.

  1. 88.

    Petunia Juss.

    Petunia Juss., Ann. Mus. Hist. Nat. 2: 215, t. 47 (1803).

    Stimoryne Raf., Fl. Tellur. 3: 76 (1857).

Annual herbs, rarely chamaephytes (P. patagonica), viscid-pubescent; brachyblasts absent. Leaves variously shaped. Inflorescence sympodial; flowers in monochasia arising from a pair of subopposite equal leaves. Calyx usually deeply lobed, lobes linear or enlarged towards the apex; corolla actinomorphic or slightly zygomorphic, tube funnel-shaped, salverform or cylindric, aestivation cochlear; stamens usually 2 + 2 + 1; anthers ventrifixed; nectary bilobed. Capsules septicidal, many-seeded, 2-valved, fruiting pedicels usually thin and deflexed. Seeds < 1 mm, reticulate-foveate with wavy thick-walled ridges. n = 7; 2n = 14.

14 spp., Brazil, Bolivia, Uruguay, Paraguay, Argentina. Petunia x hybrida (Hook.) Vilm., a hybrid between P. integrifolia (Hook.) Schinz & Thell. and P. axillaris (Lam.) Britton, Stern & Poggenb., is among the most popular annual bedding plants.

  1. 89.

    Brunfelsia L.

    Brunfelsia L., Sp. pl. 1: 191 (1753); Plowman, Fieldiana, Bot. 39: 1–135 (1998), rev.

    Franciscea Pohl (1826).

Evergreen shrubs or trees. Leaves entire, petioles glabrous to pilose. Flowers in terminal cymes. Calyx actinomorphic or slightly zygomorphic, lobes subequal; corolla zygomorphic, hypocrateriform, aestivation cochlear, lobes unequal; stamens 4, didynamous, upper lateral pair with long filaments and anthers occasionally scarcely exserted, lower posterior pair shorter, with anthers always included; anthers basifixed and monothecous; ovary bicarpellate, nectary annular; style curved at apex. Capsule bivalved or fruit subbaccate. Seeds 2–40, of different form; embryo straight or slightly curved, cotyledons incumbent, shorter than or as long as rest of embryo. n = 11; 2n = 22.

Around 50 spp., Central America and the Antilles to South America. Some spp. are widely cultivated as garden plants; a few others have medicinal properties, or are hallucinogenic or toxic species.

  1. 90.

    Leptoglossis Benth.

    Leptoglossis Benth., Bot. Voy. Sulphur: 143 (1845); Hunziker & Subils, Bot. Mus. Leafl. 27: 1–43 (1980), rev.

Xerophytic annual or perennial herbs. Leaves narrow or linear, rarely dimorphic. Calyx actinomorphic, cylindrical, 5-cleft (in L. albiflora zygomorphic, 7–10-cleft); corolla zygomorphic, ventricose, aestivation cochlear; fertile stamens 4 (2 + 2) or 2 (the remaining aborted), filaments inserted basally or halfway to the enlarged distal part of corolla tube; anthers circular, thecae always confluent; nectary invaginate-bilobed, sometimes pelviform; style with two lateral expansions above. Capsules septicidal, fusiform. Seeds ca. 100, polyhedric; embryo slightly curved, cotyledons shorter than rest of embryo. n = 10; 2n = 20.

Seven spp., one in Argentina and six in coastal Peru and Chile.

  1. 91.

    Nierembergia Ruiz & Pav.

    Nierembergia Ruiz & Pav., Fl. peruv. prodr.: 23 (1794); Cocucci & Hunziker, Fl. Fanerog. Argent. 15: 1–15 (1995), reg. rev.

Herbs, sometimes rhizomatous or tuberous, or shrubs. Leaves linear to circular. Flowers solitary or in lax cymes, pedicels short. Calyx funnel-shaped or bell-shaped with 10 longitudinal veins; corolla zygomorphic, tube filiform, abruptly expanded into funnel-, bell-, or cup-shaped limb, inside with elaiophores, aestivation cochlear; stamens 5 or 3 + 2 (occasionally 4), attached to the mouth of tube; anthers ventrifixed; nectary absent; stigma, when crescent-shaped, embracing the stamens. Capsule septicidal-loculicidal, 4-valved. Seeds 20–100, polyhedric or trigonal; embryo slightly curved, cotyledons slightly longer than rest of embryo. n = 8, 9, 24; 2n = 16, 18.

20 spp., disjunct between South America and Mexico.

Two main clades within the genus were identified (Tate et al. 2009): one composed primarily of herbaceous species with broad stigmas and the other containing woody species with crescent-shaped stigmas.

  1. 92.

    Bouchetia Dunal

    Bouchetia Dunal in DC., Prodr. 13(1): 589 (1852); Hunziker & Subils, Bol. Soc. Argent. Bot. 22 (1–4): 275–295 (1983), rev.

Rhizomatous or root sprouting, decumbent herbs. Basal leaves oblong-spathulate or rhombic-ovate, upper leaves narrowly elliptic. Flowers solitary, axillary. Calyx tubular-campanulate, corolla zygomorphic, cylindrical below, funnel-shaped above, aestivation cochlear; stamens fertile 5 (2 + 2 + 1) or 4 (2 + 2, one staminode present or absent); anthers ventrifixed; ovary and style compressed, nectary sheath-like and bilobed. Capsules septicidal-loculicidal, 4-valved. Seeds 50–60, polyhedric; embryo straight or slightly curved, cotyledons shorter than rest of embryo. n = 8.

One species in South America and two in North America.

  1. 93.

    Hunzikeria D’Arcy

    Hunzikeria D’Arcy, Phytologia 34: 283 (1976); D’Arcy, Ann. Missouri Bot. Gard. 65: 705–708 (1978), rev.

Small herbs up to 25 cm, usually much branched from a perennial root. Leaves ovate-elliptic. Flowers solitary or paired. Calyx funnel-shaped, campanulate or obconic; corolla salver-form, ventricose, aestivation cochlear; fertile stamens 4 (staminode sometimes present), didynamous, anthers ventrifixed, equal or heteromorphic; style with two lateral expansions above, nectary invaginate-bilobed. Capsule subglobose, septicidal-loculicidal, 4-valved. Seeds 7–40, reniform; embryo straight or slightly curved, cotyledons shorter than rest of embryo. n = 16.

Three spp., south-western USA, Mexico, Venezuela.

  1. 94.

    Plowmania Hunz. & Subils

    Plowmania Hunz. & Subils, Kurtziana 18: 127, f. 1–3 (1986).

Plants up to 1.5 m high, usually scandent. Leaves ovate-elliptic. Flowers 5-merous; calyx campanulate; corolla brightly coloured, zygomorphic, funnel-shaped, slightly ventricose, aestivation cochlear; stamens 4 (sometimes with a vestigial fifth stamen), didynamous, the lateral longer, posterior shorter; anthers ventrifixed; ovary with annular prominent nectary; style with two lateral expansions above. Capsule apparently septicidal, 2-valved, pericarp coriaceous. Mature seeds unknown.

Monotypic, P. nyctaginoides (Standl.) Hunz. & Subils, Mexico (Chiapas) and Guatemala.

Tribe Schwenckieae Hunz. (1977).

Herbs or small shrubs. Corolla with 5 trilobulate or trifid lobes; anthers ventrifixed; stamens fertile 2–4. Embryo tiny, cotyledons 2–3 times shorter than the rest of the embryo.

  1. 95.

    Schwenckia L.

    Schwenckia L., Gen. pl. 6: 577 (1764); Freire de Carvalho, Rodriguesia 44: 307–524 (1978), reg. rev.

Annual or perennial herbs, sometimes shrubs. Leaves sessile to petiolate. Inflorescences racemose or paniculate, axillary or terminal. Calyx tubular to campanulate, actinomorphic or slightly zygomorphic; corolla actino- or zygomorphic, 5-lobate, each lobe trilobulate or trifid, aestivation valvate-conduplicate; 2 stamens + 3 staminodes, or 2 + 2 fertile stamens; ovary bilocular, nectary invaginate-pelviform; style smooth, stigma discoid. Capsule commonly globose. Seeds up to 70, cuboidal; embryo straight; endosperm copious. n = 12.

Around 25 spp., Central America and Antilles to north-eastern Argentina.

  1. 96.

    Melananthus Walp.

    Melananthus Walp., Bot. Zeitung (Berlin) 8: 788 (1850); Freire de Carvalho, Sellowia 18 (18): 51–66 (1966), reg. rev.

    Microschwenckia Benth. (1882).

Tiny herbs, or small shrubs; indumentum glandular. Leaves sessile or subsessile. Inflorescences spiciform, terminal. Calyx zygomorphic, segments 5, unequal, triangular-elongate or linear; corolla zygomorphic or actinomorphic, tubular, straight or slightly curved, tube very narrow, cylindrical, aestivation valvate-induplicate; stamens 2 + 2; ovary unilocular (unicarpellate?), uniovulate , nectary invaginate-pelviform or slightly bilobed. Capsule zygomorphic, septicidal, valves 2, unequal. Seed 1, smooth; embryo straight or slightly curved; endosperm scanty.

Five or six spp., Cuba, Brazil, Mexico, Guatemala, Belize, Venezuela, Argentina.

While this account was in press, two genera have been sunk into synonymy: Larnax was included in Deprea (Deanna et al. 2015) and Aureliana in Athenaea (Zamberlan et al. 2015; Knapp et al. 2015).