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WOOD ANATOMY of SOLANACEAE: a SURVEY Sherwin Clqu1st’

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WOOD ANATOMY of SOLANACEAE: a SURVEY Sherwin Clqu1st’ Allertonia, 6(4), April 1992, pp. 279—326 WOOD ANATOMY OF SOLANACEAE: A SURVEY SHERwIN CLQu1sT’ ABSTRACT Quantitative and qualitative wood anatomy data are presented for 106 collections of Solanaceae, representing 21 genera and 82 species. Wood expressions are diverse in Solanaceae: wood is ring porous to diffuse porous; vessels have simple perforation plates (rarely vestiges of bars); lateral wall pitting is alternate; grooves andlor helical thickenings are present in vessels ofsome species; imperforate tracheary elements range from true tracheids with pits 5—7 m in diameter (Brunfelsia, Fabiana) to fiber-tracheids with pit cavities I m in diameter; vasicentric tracheids and vascular tracheids occur in a scattering of species; axial parenchyma is diffuse, diffuse-in-aggregates, narrow banded, ray- adjacent, and vasicentric scanty; rays are basically Heterogeneous Type IIB, but in particular species they approach or attain Heterogeneous III, Homogeneous I, Homogeneous III, Paedomorphic I, and Paedomorphic III; crystal sand and rhomboidal crystals are present in fIbriform thin-walled idioblasts, fiber-tracheids, axial parenchyma, and ray cells in a scattering of species. Solanaceae range into many habitats, and degree of wood xeromorphy or mesomorphy sensitively correlates these with respect to vessel diameter, vessel density, vessel element length, presence of vasicentric tracheids, and presence of helical sculpture on vessels. There is in Solanaceae as well as in other families a continuum, demonstrable with SEM, between the phenomena of grooves interconnecting pit apertures (confluent pit apertures), grooves accompanied by thickening bands, and presence of helical thickenings (without grooves) in vessels of Solanaceae; these conditions are included under the inclusive heading of helical sculpture. Degree of grouping of vessels is proportional to xeromorphy except in Brunfelsia and Fabiana, which have tracheids that deter vessel grouping as in other dicotyledonous woods. Wood of scandent Solanaceae is distinctive and similar to that of other climbing dicotyledons. A predominance of upright ray cells, tallness of multiseriate rays, and other features indicate paedomorphosis clearly in wood of some Solanaceae (e.g., Datura meteloides, Solanum xantii), but the family appears to have had a woody ancestry. Imperforate tracheary elements with living contents show minimal diameter of pit cavities. A feature such as crystal sand presence links Solanaceae closely to the satellite families Duckeodendraceae, Goetzeaceae, and Nolanaceae. Newly reported for the family are silica bodies in rays (Acnistus) and bordered pits in sclerosed tyloses (Solanum gayanum). INTRODUCTION The family Solanaceae contains about 85 genera and 3600 species (D’Arcy 1979, 1986). The present study describes wood features of 21 genera and 82 species, which may seem like a small proportion of the family. However, several reasons may be given for this representation. Many Solanaceae are herbaceous and offer too little wood for study of wood anatomy; wood of herbs and near- herbs is essentially primary xylem or like primary xylem, and is not really com parable with data obtained from woody cylinders of appreciable size (a few some what woody herbs have nevertheless been included in this study for purposes of ‘Rancho Santa Ana Botanic Garden and Department of Biology, Pomona College, Claremont, California 91711. 279 280 ALLERTONIA 6.4 comparison). Most xylaria contain material primarily of trees; Solanaceae are not typically trees, and thus xylaria offer only a very small fraction of the family. Furthermore, Solanaceae are most abundant in tropical areas and wood oftropical genera has not been sampled so extensively (based on species number) as has that of temperate genera. More significantly, identification of tropical Solanaceae— particularly in large genera such as Solanum—is often difficult, so that not only are wood collectors tempted to avoid some large genera, but specimens when taken are sometimes not reliably identified to species, if they are identified to species at all. If these various circumstances are considered, the proportion of species included in the present study does not seem so scanty. In fact, samples determined only to genus have not been included. Despite the number of woody Solanaceae, there has been no previous survey of wood anatomy in the family, other than the familial summaries of Solereder (1908) and Metcalfe & Chalk (1950). Details on wood anatomy have been offered for a few species (a single species in some papers listed) by Ahmad (1964), Baas & Schweingruber(1987), Bonnemain(1970), Cariquist & Hoekman (1985), Cozzo (1946), D’Arcy (1970), Descole & O’Donell (1937), Fahn et al. (1986), Gottwald & Parameswaran (1964), Greguss (1959), Melville (1949), Norverto (1989), Tor torelli (1940), and Williams (1936). Inamdar & Murthy (1977) and Murthy et al. (1980) surveyed vessel elements in selected Solanaceae, but these studies are mainly concerned with primary xylem, and their results are not really applicable here. The present study is thus the first attempt to survey wood anatomy for the family. The present effort must be termed a survey rather than a monograph, and hopefully the inherent interest ofsolanaceous wood as revealed here will encourage further collection of woods that can lead to monographs of wood of particular solanaceous genera. The inherent interest in wood anatomy of Solanaceae is considerable because the family shows a wide range of features. Although perforation plates are char acteristically simple in the family, a very few vestigial scalariform or aberrant plates may be observed. Imperforate tracheary elements range from cells that must be termed tracheids (according to the scheme of Bailey, 1936, or the IAWA Committee on Nomenclature, 1964) to fiber-tracheids with vestigial borders on pits; some species have vasicentric tracheids. Axial parenchyma is exclusively diffuse in some species, exclusively vasicentric in others. Rays are predominantly multiseriate in some species, predominantly uniseriate in others, and in histology, ray cells vary from exclusively procumbent to exclusively upright. Crystal types include rhomboidal crystals and crystal sand in axial parenchyma, ray cells, and in fibriform idioblasts. This wide range of wood features invites comparison to taxonomic systems. Systems for subfamilial and tribal groupings reviewed by D’Arcy (1979) and Hunziker (1979) offer frameworks for comparison to wood data. The wide range of wood features in Solanaceae represents more numerous instances of ecological patterning of wood than character state distributions that primarily relate to taxonomic groupings. Certainly Solanaceae occupy a wide range of habitats. The following classification of the species in the present study can be offered, together with subsequent modifying comments: 1992 CARLQUIST: SOLANACEAE 281 Tropical trees or large shrubs: Brugmansia sanguinea, B. suaveolens, Cypho mandra hartwegii, Solanum auriculatum, S. australe, S. erianthum, S. gran diflorum, S. hayesii, S. hirtum, S. hispidum, S. leucocarpon, S. nigricans, S. nudum, S. paludosum, S. rugosum, S. saponaceum, and S. triste. Subtropical trees or large shrubs: Acnistus arborescens, A. grandiflorus, Nico tiana otophora, N. raimondii, N. setchellii, N. tomentosa, Nothocestrum (all species), Solanum acropterum, S. albidum, S. bahamense, S. kauaiense, S. oblongifolium, S. sandwicense, and S. trichoneuron. Temperate tree or large shrubs: Duboisia myoporoides. Subtropical shrubs: Acnistus parviflorus, Brunfelsia calycina, B. nitida, Cestrum (all species), Dunalia (both species), Iochroma tubulosa, Lycium sandwicense, Nicotiana cordifolia, N. glauca, Solanum crispum, S. gayanum, and S. nel sonii. Temperate shrubs: Anthocercis littorea, Fabiana (all species), Grabowskya (all species), Lycianthes lycioides, Lycium (all species except L. sandwicense), Solanum nitidum, and S. simile. Temperate subshrubs: Solanum douglasii, S. xantii. Temperate herbs: Datura meteloides, Lycopersicon esculentum. Subtropical climbers: Solandra guttata, Solanum appendiculatum, S. jasmi noides, S. sodiroi, S. tetrapetalum, and Streptosolen jamesonii. Within the above categories, different climatic regimes are represented. For example, “subtropical” is applied to Andean shrubs that experience minimal frost, despite latitude near the equator, as well as to shrubs from higher latitude but lower elevation (e.g., Solanum bahamense). Some of the shrubs termed temperate here are from tropical latitudes but at high elevation where frost is prevalent (Solanum nitidum), whereas others are from extreme habitats such as the Atacama Desert of Chile (Fabiana bryoides) or areas of Patagonia that are both dry and cold (Fabiana viscosa, Grabowskya ameghinoi). Some of the species of Lycium are from areas that are moderately dry (L. europaeum, shores ofthe Mediterranean Sea), whereas others are from desert areas (L. brevipes, L. fremontii). Although at first glance these seem like quite different kinds of habitats in terms of total rainfall, the adaptations of wood in Mediterranean-type climates and desert areas are very similar (Cariquist & Hoekman, 1985), a fact that reflects the necessity of wood to maintain an intact water conducting system during a prolonged dry season. MATERIALS AND METHODS TLE 1 indicates the sources of materials. Numerous samples were provided by xylaria. These are supplemented by specimens I have obtained during field work in California, Chile, Peru, and the Hawaiian
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