A search for phylogenetically informative wood characters within Lecythidaceae s.l

The wood structure of 71 species representing 24 genera of the pantropical Lecythidaceae s.l., including the edible Brazil nuts (Bertholletia excelsa) and the spectacular cannon-ball tree (Couroupita guianensis), was investigated using light and scanning electron microscopy. This study focused on finding phylogenetically informative characters to help elucidate any obscure evolutionary patterns within the family. The earliest diverging subfamily Napoleonaeoideae has mixed simple/scalariform vessel perforations, scalariform vessel-ray pitting, and high multiseriate rays, all features that are also present in Scytopetaloideae. The wood structure of Napoleonaea is distinct, but its supposed close relative Crateranthus strongly resembles Scytopetaloideae. The isolated position of Foetidia (Foetidioideae) can be supported by a unique type of vessel-ray pitting that is similar in shape and size to intervessel pitting (distinctly bordered, <5 μm). The more derived Planchonioideae and Lecythidoideae share exclusively simple perforations and two types of vessel-ray pitting, but they can easily be distinguished from each other by the size of intervessel pitting, shape of body ray cells in multiseriate rays, and the type of crystalliferous axial parenchyma cells. The anatomical diversity observed is clearly correlated with differences in plant size (shrubs vs. tall trees): the percentage of scalariform perforations, as well as vessel density, and the length of vessel elements, fibers, and multiseriate rays are negatively correlated with increasing plant size, while the reverse is true for vessel diameter.     

Comparative wood anatomy of epacrids (Styphelioideae,Ericaceae s.L.)

The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel-ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze-thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.     

A review of the platanaceous woods from the Eocene paratropical rainforest of south-east England

Small diameter pyritized axes, commonly referred to as 'twigs', of fossil platanaceous wood are described from the Lower Eocene London Clay Formation of south-east England. These twigs are characterized by solitary vessels with scalariform perforation plates, opposite intervessel pits, and tall, multiseriate rays that dilate in the phloem region. The wood anatomy supports close relationship to members of extant Platanaceae and the material is placed in the organ genus Plataninium Unger erected for fossil woods with close anatomical similarity to Platanus L. This material supplements the fossil record of platanaceous type wood from the Eocene London Clay and documents the first record of Plataninium decipiens Brett in the twig flora.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 181–191.     

Relationships within balsaminoid Ericales: a wood anatomical approach

Wood samples of 49 specimens representing 31 species and 11 genera of woody balsaminoids, i.e., Balsaminaceae, Marcgraviaceae, Pellicieraceae, and Tetrameristaceae, were investigated using light microscopy and scanning electron microscopy. The wood structure of Marcgraviaceae, Pellicieraceae, and Tetrameristaceae is characterized by radial vessel multiples with simple perforation plates, alternate vessel pitting, apotracheal and paratracheal parenchyma, septate libriform fibers, and the presence of raphides in ray cells. Tetrameristaceae and Pellicieraceae are found to be closely related based on the occurrence of unilaterally compound vessel-ray pitting and multiseriate rays with long uniseriate ends. The narrow rays in Pelliciera are characteristic of this genus, but a broader concept of Tetrameristaceae including Pelliciera is favored. Within Marcgraviaceae, wide rays (more than five-seriate) are typical of the genus Marcgravia. Furthermore, there is evidence that the impact of altitude and habit plays an important role in the wood structure of this family. The wood structure of Balsaminaceae cannot be compared systematically with other balsaminoids because of their secondary woodiness. Balsaminaceae wood strongly differs due to the presence of exclusively upright ray cells in Impatiens niamniamensis, the absence of rays in Impatiens arguta, and the occurrence of several additional paedomorphic features in both species.     

Systematic implications of wood and bark anatomy in the Pacific Island genus Meryta (Araliaceae)

Wood anatomy was examined in 16 species of Meryta (a genus of c . 35 species) and bark anatomy was studied in 12 species. All but two of these taxa form an assemblage corresponding to the Northern Arc clade, one of two major groups identified by a recent molecular phylogenetic study. M. sinclairii and M. tenuifolia (corresponding to the New Zealand/Fiji clade) differ distinctly by having more numerous simple perforation plates, multiseriate rays with few marginal rows, and the absence of sclerified cells in collapsed secondary phloem, a bark feature that has not yet been found elsewhere in Araliaceae. The increase in abundance of simple perforation plates in the wood of these two species is not accompanied by a decrease in the number of bars on scalariform perforation plates. The wood structure of Meryta bears a strong resemblance to members of the Pacific Schefflera clade, sharing similar ranges of variation of several features. Bark characters, such as the diameter of the cortical secretory canals, the types of crystal in cortical cells, the types of axial parenchyma cell in collapsed secondary phloem, and the presence of sheath cells by phloem rays, appear to be of diagnostic value for some species of Meryta .  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 363–379.     

Anatomical features, monomer lignin composition and accumulation of phenolics in 1-year-old branches of the Mediterranean Cistus ladanifer L.

Mediterranean shrub species are described as having phenology, habitus , reproductive biology and anatomical alterations in certain tissues, allowing their survival during the dry season and protecting them from herbivory. Anatomical and chemical analyses were conducted in 1-year-old branches of Cistus ladanifer L. in order to investigate the role played by shoot structure in the adaptive strategies of this species in the Mediterranean environment. Results showed that both xylem and pith underwent lignification. Pith parenchyma cells had thickened walls, higher lignin content than xylem and different monomer composition. Xylem presented features aiding safe water transport. A large accumulation of phenolic substances was found in xylem, pith and cortical parenchyma. Observations reported in this paper suggest the occurrence of adaptive strategies in 1-year-old branches of C. ladanifer whose structural features: (1) allow mechanical reinforcement of tissues to withstand drought without suffering permanent damage; (2) favour safety rather than efficiency in water transport; (3) defend the plants from animal predation and pathogens by accumulating phenolics in various tissues, and (4) protect inner tissues against UV-B radiation through deposition of phenolic compounds in cortical layers.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 361–371.     

Re-interpretation of Wexfordia hookense from the Upper Devonian of Ireland as an arborescent lycophyte

New collections of pyritized axes of the lycophyte Wexfordia hookense have been made from the Upper Devonian (uppermost Famennian) type locality at Sandeel Bay, County Wexford, in south-eastern Ireland. The specimens reveal additional histological features that permit reinterpretation of the morphology of this taxon and reevaluation of its taxonomic affinities. Wexfordia is shown to possess both secondary xylem, with narrow, uni- to biseriate rays, and periderm. The range of variation in relative amounts of primary and secondary xylem can be correlated with position in the mature plant. This evidence indicates that Wexfordia was a small tree rather than an herbaceous form. Fine structure of tracheids and additional anatomical features strongly support affinities with Carboniferous arborescent Isoetales, rather than Devonian Protolepidodendrales, and further support the hypothesis that radiation in this lineage was well underway prior to the Carboniferous.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 275–287.     

Wood structure of Aegiceras corniculatum and its ecological adaptations to salinities

We describe the wood structure of Aegiceras corniculatum and its differences under various soil salinities. This species had diffuse-porous wood with poorly defined growth rings. Vessels which had single perforations occurred abundantly and in multiples and were storeyed. Intervascular pits between contiguous vessels were alternate bordered ones while half-bordered pit-pairs existed between both vessel-ray and vessel-parenchyma. Homogenous xylem rays were multiseriate and uniseriate. Fiber-tracheids with bordered pits often had thinner walls. Xylem parenchyma cells were scant and distributed diffusely and paratracheally. Differences in the structural and quantitative characters of vessels, xylem rays and fiber-tracheids under diverse soil salinities are described. This revised version was published online in August 2006 with corrections to the Cover Date.     

广义锦葵科火绳树属4个种的枝条木材解剖及其分类学意义

研究了广义锦葵科火绳树属4个种枝条的木材解剖。火绳树属枝条为散孔至半环孔材,管孔主要为单管孔和2～3个管孔组成的径列复管孔;导管间纹孔式和射线导管间纹孔式互列、小;侵填体和螺纹加厚缺如。射线主要为单列射线,2～3列射线较多;射线细胞多为方形,射线组织主要为异型,边缘直立细胞常1行;射线组织稀为同型;鞘细胞和瓦形细胞缺如。轴向薄壁组织傍管和离管型,主要为带状。晶体丰富,主要在射线、纤维和薄壁组织中。研究的4个种可以通过枝条木材解剖特征加以区分。     

Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics

CARLQUIST, S., 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Qualitative and quantitative data, based mostly upon liquid-preserved specimens, are presented for Akebia, Roquila, Decaisnea, Holbodia, Lardizabala, Sinofranchetia and Stauntonia . Because Decaisnea is a shrub whereas the other genera are vines, anatomical differences attributable to the scandent habit can be considered. These include exceptionally wide vessels, a high proportion of vessels to tracheids (or other imperforate trdcheary elements) as seen in transection, simple perforation plates, multiseriate rays which are wide and tall, and pith which is partly or wholly sclerenchymatous. With respect to ecology, two features are discussed: spirals in narrower vessels may relate to adaptation to freezing in the species of colder areas, and crystalliferous sclereids seem adapted in morphology and position to deterrence of phytophagous insects or herbivores. The wood may provide mechanisms for maintaining conduction even if wider vessels are deactivated temporarily by formation of air embolisms. Wood and stem anatomy of Lardizabalaceae compare closely to those of Berberidaceae and of Clematis (Ranunculaceae), as well as to other families of Berberidales. Decaisnea is more primitive than these in having consistently sralariform perforation plates and in having scalariform pitting on lateral walls of vessels. A tentative listing of anatomical features which may correspond to generic limits is given.     

Systematic implications of seed coat diversity in Gaultherieae (Ericaceae)

The seed morphology of 90 samples from 83 species of tribe Gaultherieae (Chamaedaphne, Diplycosia, Eubotryoides, Eubotrys, Gaultheria and Leucothoe) and relatives in tribes Andromedeae (Andromeda and Zenobia) and Vaccinieae (Satyria) was investigated with stereoscopic and scanning electron microscopy. Seeds exhibit variation in shape, colour, size, wing, hilum region, primary ornamentation and epidermal cells. Non‐metric multidimensional scaling (NMDS) analysis based on selected seed characters supports the affinities of some groups within Gaultherieae at various taxonomic levels. Seed characters corroborate the delimitation of Andromeda, Chamaedaphne, Leucothoe, Satyria and Zenobia and Gaultheria series Trichophyllae, series Hispidulae, section Amblyandra and section Brossaeopsis. Parsimony optimization of seed characters onto a previously published phylogenetic estimate of Gaultherieae reveals that small seeds have evolved from larger seeds and an areolate seed coat has evolved from a reticulate seed coat. Optimization also suggests that several seed character states are synapomorphies or potential synapomorphies for some major clades of Gaultherieae. Seeds of Gaultherieae from East Asia, temperate North America and the Pacific are more diverse than those from tropical America. Samples from the eastern Himalaya possess the highest variation in seed morphology. The wing and bulging edge cells observed in seeds of Leucothoe suggest dispersal by wind. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 477–495.     

Wood and stem anatomy of woody Amaranthaceae s.s.: ecology, systematics and the problems of defining rays in dicotyledons

Wood and stem anatomy is studied for seven species of six genera (root anatomy also reported for one species) of Amaranthaceae s.s. Quantitative data on vessels correlate closely with relative xeromorphy of respective species, agreeing with values reported for dicotyledons without successive cambia in comparable habitats. Libriform fibre abundance increases and vessel diameter decreases as stems and roots of the annual Amaranthus caudatus mature. Long, thick-walled fibres in Bosea yervamora may be related to the upright nature of elongate semi-climbing stems. Non-bordered or minutely bordered perforation plates characterize Amaranthaceae, as they do most other Caryophyllales. Amaranthaceae have idioblastic cells containing druses, rhomboidal crystals or crystal sand: these forms intergrade and seem closely related. Rays are present in secondary xylem of the Amaranthaceae studied. Cells intermediate between ray cells and libriform fibres occur in Charpentiera elliptica . Degrees of diversity in rays and reports of raylessness in Amaranthaceae induce discussion of definition and identification of rays in dicotyledons; some sources recognize both rays and radial plates of conjunctive tissue in Amaranthaceae. The action of successive cambia is described: lateral meristem periclinal divisions produce secondary cortex externally, conjunctive tissue internally and yield vascular cambia as well. Vascular cambia produce secondary phloem and secondary xylem, in both ray and fascicular zones, as in a dicotyledon with a single cambium. Identification of meristem activity and appreciation of varied ray manifestations are essential in understanding the ontogeny of stems in Amaranthaceae (which have recently been united with Chenopodiaceae).  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 1–19.     

Caryophyllales: a key group for understanding wood anatomy character states and their evolution

Definitions of character states in woods are softer than generally assumed, and more complex for workers to interpret. Only by a constant effort to transcend the limitations of glossaries can a more than partial understanding of wood anatomy and its evolution be achieved. The need for such an effort is most evident in a major group with sufficient wood diversity to demonstrate numerous problems in wood anatomical features. Caryophyllales s.l., with approximately 12 000 species, are such a group. Paradoxically, Caryophyllales offer many more interpretive problems than other ‘typically woody’ eudicot clades of comparable size: a wider range of wood structural patterns is represented in the order. An account of character expression diversity is presented for major wood characters of Caryophyllales. These characters include successive cambia (more extensively represented in Caryophyllales than elsewhere in angiosperms); vessel element perforation plates (non‐bordered and bordered, with and without constrictions); lateral wall pitting of vessels (notably pseudoscalariform patterns); vesturing and sculpturing on vessel walls; grouping of vessels; nature of tracheids and fibre‐tracheids, storying in libriform fibres, types of axial parenchyma, ray anatomy and shifts in ray ontogeny; juvenilism in rays; raylessness; occurrence of idioblasts; occurrence of a new cell type (ancistrocladan cells); correlations of raylessness with scattered bundle occurrence and other anatomical discoveries newly described and/or understood through the use of scanning electron microscopy and light microscopy. This study goes beyond summarizing or reportage and attempts interpretations in terms of shifts in degrees of juvenilism, diversification in habit, ecological occupancy strategies (with special attention to succulence) and phylogenetic change. Phylogenetic change in wood anatomy is held to be best interpreted when accompanied by an understanding of wood ontogeny, species ecology, species habit and taxonomic context. Wood anatomy of Caryophyllales demonstrates problems inherent in binary character definitions, mapping of morphological characters onto DNA‐based trees and attempts to analyse wood structure without taking into account ecological and habital features. The difficulties of bridging wood anatomy with physiology and ecology are briefly reviewed. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 342–393.     

Illicioxylon, an Element of Gondwanan Polar Forests? Late Cretaceous and Early Tertiary Woods of Antarctica

The Late Cretaceous and early Tertiary sediments from the northernPeninsula region of Antarctica yield a rich assemblage of fossilwood with well preserved anatomy. Wood specimens of a previouslyrecognized morphotype are described. The woods are characterizedby diffuse porous wood, mainly solitary vessels with long scalariformperforation plates, scalariform and opposite vessel-ray pitting,generally uniseriate and biseriate heterogeneous rays, and tracheidswith obvious uniseriate, circulate, bordered pits. These fossilspecimens show greatest anatomical similarity to the organ genusIllicioxylon Gottwald and extant members of the Illiciaceae.The occurrence of illiciaceous-like wood in Gondwana suggeststhat the distribution of this family may have been more widespreadin the geological past and that a relatively warm temperateclimate prevailed over the northern Peninsula region of Antarcticaduring the Late Cretaceous and early Cenozoic. Copyright 2000Annals of Botany Company Fossil, wood, Illiciaceae, Illicioxylon, Illicium, Cretaceous, Tertiary, Gondwana, Antarctica     

Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales)

Chrysobalanaceae s.l. , one of the few suprafamilial subclades of Malpighiales that is supported by molecular phylogenetic analyses, and containing Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, and Trigoniaceae, was comparatively studied with regard to floral structure. The subclade is well supported by floral structure. Potential synapomorphies for Chrysobalanaceae s.l. are the following shared features: floral cup; flowers obliquely monosymmetric; sepals congenitally united at base; sepals of unequal size (outer two shorter); fertile stamens concentrated on the anterior side of the flower and sometimes united into a strap; staminodes absent in the posteriormost antepetalous position; anthers extremely introrse, with thecae almost in one plane; endothecium continuous over the dorsal side of the connective; dorsal anther pit; gynoecium completely syncarpous up to the stigma; carpel flanks slightly bulged out transversely and thus carpels demarcated from each other by a longitudinal furrow; flowers with dense unicellular, non-lignified hairs, especially on the gynoecium; light-coloured, dense indumentum on young shoots and inflorescences. Potential synapomorphies for Chrysobalanaceae + Euphroniaceae include: spur in floral cup; clawed petals; lignified hairs on petals; nectary without lobes or scales and mostly annular. Potential synapomorphies for Dichapetalaceae + Trigoniaceae include: special mucilage cells in sepals in mesophyll (in addition to epidermis); anthers almost basifixed; gynoecium synascidiate up to lower style; nectary with lobes or scales and semi-annular.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 249–309.     

WOOD ANATOMY OF ILLICIUM (ILLICIACEAE): PHYLOGENETIC,ECOLOGICAL, AND FUNCTIONAL INTERPRETATIONS

Quantitative and qualitative features are reported for 23 collections of ten species of Illicium, sole genus of Illiciaceae. Vessel elements are long, thin-walled, and angular; perforation plates are scalariform and range from long to moderately long; vessel-ray pitting is scalariform to opposite. Tracheids bear fully bordered pits. Axial parenchyma is sparse, abaxial to vessels with some diffuse cells also present. Rays are both multiseriate and uniseriate; the former are lacking near the pith in some species. No erect sheathing cells are present on multiseriate rays, and procumbent cells become more abundant as stems increase in size. Four species from montane subequatorial highlands lack growth rings and helical sculpture in vessels; the remainder of Illicium species have these features. Notably narrow vessels and large numbers of vessels per sq. mm characterize the temperate species from northern limits for the genus, I. anisatum, I. floridanum, and I. parviflorum. Greater vessel density offers redundancy and greater safety and is correlated with greater frost and possibly greater fluctuation in moisture availability. These two ecological features are probably also related to the narrow vessel diameter, which may retard entry of air embolisms (lowered air entry values) in accord with the physiological considerations of Slatyer. Reduction of number of bars per perforation plate within Illicium is also correlated with the more northerly climates. An additional hypothesis for evolution of shorter vessel elements in dicotyledons is introduced: if, as claimed by Slatyer, air embolisms in vessels tend to stop at ends of vessel elements and thus do not disable entire vessels, shorter vessel elements would maximally localize air embolisms. Presence of helical sculpture and other forms of relief within vessels has been difficult to explain in conifers and dicotyledons; such features might help resist cavitations by increasing adhesion of water molecules to cell walls (hydration).     

Systematic and comparative anatomy of Maxillarieae (Orchidaceae), sans Oncidiinae

On the basis of floral and vegetative morphology, 63 tropical American genera have been recognized within Maxillarieae. We were able to examine anatomical material of all subtribes, excluding Oncidiinae. Stegmata with conical silica bodies occur in leaves and stems of all subtribes excluding Ornithocephalinae, and pericyclic stegmata found in roots are characteristic of Lycastinae. Lycastinae and Maxillariinae are characterized by foliar glands, foliar fibre bundles and tilosomes. Endodermal cells are U-thickened in most Zygopetalinae; O-thickened in most Lycastinae, Ornithocephalinae and Telipogoninae; variously thickened in Maxillariinae; and thin-walled in Cryptarrhena lunata . Water-storage cells varied from thin-walled to variously banded throughout Maxillarieae. Cladistic analyses using anatomical characters yielded no resolution among subtribes, illustrating that anatomical characters are of limited value in assessing relationships within this tribe.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 251–274.     

Wood anatomy of Penaeaceae (Myrtales): comparative, phylogenetic, and ecological implications

Wood samples of stems, lignotubers, and roots of the majority of species of Penaeaceae were analyzed with respect to qualitative and quantitative features. Virtually no data have hitherto been presented on xylem features of this family, restricted to Cape Province, South Africa. Presence of vestured pits in vessels, septate crystalliferous parenchyma in wood, intraxylary phloem, predominantly erect ray cells in the typically narrow, multiseriate rays and in the uniseriate rays, and amorphous deposits in ray cells place Penaeaceae securely in Myrtales and help to define that order. By comparison of ecological preferences of the species, as observed during field work, with quantitative analysis of conductive tissue, close correspondence of the wood structure to habit and habitat is demonstrated.     

A comparison of the wood anatomy of 11 species from two cerrado habitats (cerrado s.s. and adjacent gallery forest)

A comparative study of the secondary xylem (wood) anatomy of 11 species (38 specimens) occurring in cerrado s.s. and the adjacent gallery forest (both cerrado s.l. habitat) was made with the aim of identifying the anatomical characteristics of ecological value and correlating them with the environmental conditions. The anatomical features that vary, in general, between the two habitats are: growth ring distinctness (well or poorly defined); tyloses and deposits (more abundant in cerrado specimens); gelatinous fibres (more evident in cerrado specimens and in different patterns between habitats); variation in paratracheal and banded parenchyma (more abundant in cerrado); and more cells per parenchyma strand in cerrado. In general, gallery forest specimens have wider vessels, fewer vessels per square millimetre and larger intervessel pits, indicating more efficient water conduction, whereas cerrado s.s. specimens are the opposite, with low vulnerability and mesomorphy indices, demonstrating greater safety under conditions of water stress. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ?? , ??–??.     

Wood anatomy of Polygonaceae: analysis of a family with exceptional wood diversity

Quantitative and qualitative data are presented for woods of 30 species of woody Polygonaceae. Wood features that ally Polygonaceae with Plumbaginaceae include nonbordered perforation plates, storeying in narrow vessels and axial parenchyma, septate or nucleate fibres, vasicentric parenchyma, pith bundles that undergo secondary growth, silica bodies, and ability to form successive cambia. These features are consistent with pairing of Plumbaginaceae and Polygonaceae as sister families. Wood features that ally Polygonaceae with Rhabdodendraceae include nonbordered perforation plates, presence of vestured pits in vessels, presence of silica bodies and dark-staining compounds in ray cells, and ability to form successive cambia. Of the features listed above, nonbordered perforation plates and ability to form successive cambia may be symplesiomorphies basic to Caryophyllales sensu lato . The other features are more likely to be synapomorphies. Wood data thus support molecular cladograms that show the three families near the base of Caryophyllales s.l. Chambered crystals are common to three genera of the family and may indicate relationship. Ray histology suggests secondary woodiness in Antigonon, Atraphaxis, Bilderdykia, Dedeckera, Eriogonum, Harfordia, Muehlenbeckia, Polygonum , and Rumex . Other genera of the family show little or no evidence of secondary woodiness. Molecular data are needed to confirm this interpretation and to clarify the controversial systematic groupings within the family proposed by various authors. Vessel features of Polygonaceae (lumen diameter, element length, density, degree of grouping) show an extraordinary range from xeromorphy to mesomorphy, indicating that wood has played a key role in ecological and habital shifts within the family; the diversity in ecology and habit are correlated with quantitative wood data.  © 2003 The Linnean Society of London. Botanical Journal of the Linnean Society , 2003, 141 , 25−51.