Asteraceae subfamily Barnadesioideae (ten genera, c. 90 species), confined to South America, are sister to the remainder of the family. The relative antiquity of the barnadesioids might lead one to expect that they contain more wood features plesiomorphic for the family, but only one character clearly falls in that category. Pits on imperforate tracheary elements are bordered (except for annuals), whereas simple pits occur in two related families, Calyceraceae (part) and Stylidiaceae (all that have been examined); in Goodeniaceae bordered pits only occur. By attaining fully bordered pits in Chuquiraga, the imperforate tracheary elements qualify as an apomorphy, 'neotracheids', valuable for resisting embolism formation in dry and cold South American habitats. Neotracheids are found also in Loricaria (Asteraceae: Inuleae), also from these habitats. Neotracheids, like plesiomorphic tracheids, are conductive, unlike fibre tracheids and libriform fibres. Other barnadesioid wood characters adapted to cold and drought include grouping of vessels, high vessel density, shorter vessel elements and helical sculpture (including helical thickenings on lumen-facing walls) of secondary xylem vessels. In Chuquiraga and Dasyphyllum, these helical thickenings are bordered in some species (new report for angiosperms). Some of the barnadesioid adaptations to cold and drought can be found in North American Artemisia spp. (Asteraceae: Anthemideae), especially in montane and desert areas. Wood features of barnadesioids match their respective habits and habitats: a few trees; shrubs of humid, dry or desert areas; a distinctive rhizomatous succulent in the pampas (Schlechtendalia); a scree/gravel perennial (Huarpea) and two genera of annuals, one with succulent leaves (Duseniella) and one with rayless (at least at first) stems in arid and open soils (Doniophyton). Diversity is unusual considering the small size of the subfamily. Examples of endodermal crystals (Arnaldoa only), pith sclereids and primary xylem fibres are cited.

Argophyllaceae (Argophyllum, 14 spp.; Corokia, 6 spp.; Lautea, 1 sp.), are shrubs that occur in the southwestern Pacific and eastern Australia. They occur in habitats where moisture is relatively common but dry days and mild frost may occur. The woods of these genera show enough distinctive features to justify their grouping in a single family: perforation plates with 10–20 bars, vessel elements narrow and numerous per mm2, imperforate tracheary elements about 50% longer than the vessel elements, axial parenchyma scarce, diffuse, multiseriate rays narrow and heterocellular (upright cells common in uniseriate rays), crystals absent, gum deposits common. These features group the genera of Argophyllaceae more closely with each other than with the nearest families in Asterales (Alseuosmiaceae, Phellinaceae). Probable apomorphies of the genera include helical thickenings in vessels and tracheids, together with abundant tracheids and rare septate fiber-tracheids (Corokia); almost total absence of axial parenchyma and tracheids combined with maximal abundance of septate fiber-tracheids and no helical thickenings (Argophyllum, Lautea). Lautea, formerly included within Corokia, has floral and foliar distinctions and is endemic to a single island, Rapa Iti. Woods of Argophyllaceae are alike in their ecological adaptations (perforation plates, vessel diameter and density) but the presence of tracheids and helical thickenings in Corokia suggest adaptations to frost and mild drought. As expected, vessels group more prominently in the tracheid-free species (Argophyllum, Lautea) but very little in the tracheid-rich genus Corokia.

The genus Bidens (Compositae) comprises c. 230 species distributed across five continents, with the 41 Polynesian species displaying the greatest ecomorphological variation in the group. However, the genus has had a long and complicated taxonomic history, and its phylogenetic and biogeographic history are poorly understood. To resolve the evolutionary history of the Polynesian Bidens, 152 individuals representing 91 species were included in this study, including 39 of the 41 described species from Polynesia. Four chloroplast and two nuclear DNA markers were utilized to estimate phylogenetic relationships, divergence times, and biogeographic history. Bidens was found to be polyphyletic within Coreopsis, consistent with previous assessments. The Polynesian radiation was resolved as monophyletic, with the initial dispersal into the Pacific possibly from South America to either the Hawaiian or Marquesas Islands. From the Marquesas, Bidens dispersed to the Society Islands, and ultimately to the Austral Islands. The initial diversification of the crown group in the Pacific is estimated to have occurred ~1.63 mya (0.74-2.72, 95% HPD), making Polynesian Bidens among the youngest and most rapid plant diversification events documented in the Pacific. Our findings suggest that relatively rare long-distance dispersal and founder-event speciation, coupled with subsequent loss of dispersal potential and within-island speciation, can explain the repeated and explosive adaptive radiation of Bidens throughout the archipelagoes of Polynesia.

The very different evolutionary pathways of conifers and angiosperms are very informative precisely because their wood anatomy is so different. New information from anatomy, comparative wood physiology, and comparative ultrastructure can be combined to provide evidence for the role of axial and ray parenchyma in the two groups. Gnetales, which are essentially conifers with vessels, have evolved parallel to angiosperms and show us the value of multiseriate rays and axial parenchyma in a vessel-bearing wood. Gnetales also force us to re-examine optimum anatomical solutions to conduction in vesselless gymnosperms. Axial parenchyma in vessel-bearing woods has diversified to take prominent roles in storage of water and carbohydrates as well as maintenance of conduction in vessels. Axial parenchyma, along with other modifications, has superseded scalariform perforation plates as a safety mechanism and permitted angiosperms to succeed in more seasonal habitats. This diversification has required connection to rays, which have concomitantly become larger and more diverse, acting as pathways for photosynthate passage and storage. Modes of growth such as rapid flushing, vernal leafing-out, drought deciduousness and support of large leaf surfaces become possible, advantaging angiosperms over conifers in various ways. Prominent tracheid-ray pitting (conifers) and axial parenchyma/ray pitting to vessels (angiosperms) are evidence of release of photosynthates into conductive cells; in angiosperms, this system has permitted vessels to survive hydrologic stresses and function in more seasonal habitats. Flow in ray and axial parenchyma cells, suggested by greater length/width ratios of component cells, is confirmed by pitting on end walls of elongate cells: pits are greater in area, more densely placed, and are often bordered. Bordered pit areas and densities on living cells, like those on tracheids and vessels, represent maximal contact areas between cells while minimizing loss of wall strength. Storage cells in rays can be distinguished from flow cells by size and shape, by fewer and smaller pits and by contents. By lacking secondary walls, the entire surfaces of phloem ray and axial phloem parenchyma become conducting areas across which sugars can be translocated. The intercontinuous network of axial parenchyma and ray parenchyma in woods is confirmed; there are no “isolated” living cells in wood when three-dimensional studies are made. Water storage in living cells is reported anatomically and also in the form of percentile quantitative data which reveal degrees and kinds of succulence in angiosperm woods, and norms for “typically woody” species. The diversity in angiosperm axial and ray parenchyma is presented as a series of probable optimal solutions to diverse types of ecology, growth form, and physiology. The numerous homoplasies in these anatomical modes are seen as the informative results of natural experiments and should be considered as evidence along with experimental evidence. Elliptical shape of rays seems governed by mechanical considerations; unusually long (vertically) rays represent a tradeoff in favor of flexibility versus strength. Protracted juvenilism (paedomorphosis) features redirection of flow from horizontal to vertical by means of rays composed predominantly or wholly of upright cells, and the reasons for this anatomical strategy are sought. Protracted juvenilism, still little appreciated, occurs in a sizeable proportion of the world’s plants and is a major source of angiosperm diversification.

All Onagraceae for which data are available have vestured pits on vessel-to-vessel pit pairs. Vestures may also be present in some species on the vessel side of vessel-to-ray pit pairs. Herbaceous Onagraceae do not have fewer vestures, although woods with lower density (Circaea L. and Oenothera L.) have fewer vestures. Some Onagraceae from drier areas tend to have smaller vessel pits, and on that account may have fewer vestures (Epilobium L. and Megacorax S. Gonzalez & W. L. Wagner). Pit apertures as seen on the lumen side of vessel walls are elliptical, occasionally oval, throughout the family. Vestures are predominantly attached to pit aperture margins. As seen from the outer surfaces of vessels, vestures may extend across the pit cavities. Vestures are usually absent or smaller on the distal portions of pit borders (except for Ludwigia L., which grows consistently in wet areas). Distinctive vesture patterns were observed in the several species of Lopezia Cav. and in Xylonagra Donn. Sm. & Rose. Vestures spread onto the lumen-facing vessel walls of Ludwigia octovolvis (Jacq). P. H. Raven. Although the genera are presented here in the sequence of a recent molecular phylogeny of Onagraceae, ecology and growth forms are more important than evolutionary relationships with respect to abundance, degree of grouping, and morphology of vestured pits. Designation of vesture types is not warranted based on the distribution of named types in Onagraceae and descriptive adjectives seem more useful, although more data on vesturing in the family are needed before patterns of diversity and their extent can be fully ascertained. Vestures are less common and may have a more complicated genetic basis than helical thickenings in vessels, but may be a more effective form of water column maintenance.

The four species of Lennoaceae have strands of primary plus secondary xylem in a background of starch-rich parenchyma. These strands constitute a cylinder with large primary rays. The wood within these strands is markedly different from that of other families in the crown group of Boraginales such as Cordiaceae and Ehretiaceae, most of which are woody. Lennoaceae differ because they lack fibrous cells (libriform fibers), lack rays within the vascular strands, and have markedly elliptical vessel-to-vessel pits without vestures. Lennoaceae have secondary xylem with short, wide vessel elements with thick walls, horizontally elongate elliptical pits, simple perforation plates much narrower than the vessel lumen; variously uneven vessel wall thickenings; and axial parenchyma. The wood of Lennoaceae shows resemblances to unrelated succulents such as Kalanchoe (Crassulaceae) and Lithops (Aizoaceae). The vessel features also suggest adaptation to high water tensions as root parasites in desert areas, whereas the lack of imperforate tracheary elements may relate to support of the underground stem portions by sand or rock detritus. Habit and ecology are more important in the architecture of lennoaceous xylem than systematic affinities. The four species of Lennoaceae differ from each other in minor xylary features.

-Wood of Bruniaceae is very primitive according to widelyaccepted criteria. Vessels are relatively long, with scalariform perforation plates having numerous bars, fully or vestigially bordered. Many aberrations in disposition of bars and perforations in perforation plates are present, and no species has exclusively normal perforation plates. Lateral wall pitting of vessels, both intervascular and between vessels and rays, is scalariform to opposite. Vessels are solitary or nearly so, angular to round in transection. Helical thickenings are present in vessels of only a single species. All imperforate elements are thick-walled tracheids, some with gelatinous walls. Tracheids are only slightly longer (in two instances shorter) than the vessel elements they accompany in any given species. Axial parenchyma is diffuse, with very slight tendencies toward aggregates or vasicentric scanty. Rays are heterocellular, with a predominance of upright cells, or upright cells exclusively in species with very narrow multiseriate rays. Multiseriate rays vary from biseriate to an average of more than five cells in width, according to species. Uniseriate wings are present on at least some multiseriate rays in all species. Borders are frequently present on the tangential walls of ray cells, which tend to be thick. U niseriate and multiseriate rays are present in all species. Growth rings vary from absent to conspicuous, the latter mostly in montane species. Inconspicuous growth rings feature narrow vessels or narrow tracheids in latewood. More pronounced growth rings have more numerous as well as wider vessels in earlywood; vessels may be absent or nearly so in latewood, which would thus constitute an all-tracheid band produced annually. Crystals are present in rays of most species, sometimes chambered in subdivided ray cells. A few species have crystals in chambered axial parenchyma cells. Droplets or massive deposits of amorphous dark-staining compounds are present in all species. Woods of Bruniaceae are exemplary, considering their primitiveness, for adaptation to xeromorphy, based on current concepts of wood xeromorphy. Vessels are notably narrow or numerous per mm2 of transection, or both. The nature of growth rings and of perforation plate modifications are suggestive of xeromorphy. Although species of moister habitats do have quantitatively more mesomorphic wood features, the span within Bruniaceae is not great, suggesting that wood of the family as a whole is adapted to extremes of water stress in accordance with the dry, hot summers, high incidence of wind, and porous sandstone soils of Cape Province, South Africa. Lignotubers have woods more parenchymatous and mesomorphic than wood of stems in the lignotuber-bearing species. Wood of stems does not differ appreciably from that

Molecular studies indicate that Penaeaceae, Oliniaceae, and the monospecific families Alzateaceae and Rhynchocalycaceae form a clade of Myrtales. Of these four families, Penaeaceae have tracheids with vestured pits, whereas the others have septate fibers lacking vestures; all have vestured pits in vessels. Tracheid presence in Penaeaceae may be related to the arid South African habitats of the family. Presence of vestures on tracheids in families with vestured vessel pits is one indication that imperforate elements are tracheids and are conductive cells, whereas fiber-tracheids and libriform fibers are non-conductive. Tracheids occur widely in angiosperms and may be plesiomorphies or apomorphies. Combretaceae, the first branch of the Myrtales clade, has a great diversity of vesture features in vessels compared to the Penaeaceae alliance families. Alzatea has vestures that spread over the inside of the vessels, whereas in most taxa of the alliance, vestures are confined to the pit cavities and pit apertures. Vestures in the alliance tend to be globular in shape, and are bridged together by strands of wall material. Lignotubers and roots in Penaeaceae have vestures much like those in stems. Only a few species and genera (notably Alzatea) of the alliance have vesture features the pattern of which correlates with the current taxonomic system. Vestured pits should be viewed from the inside surface of vessels as well as the outer surface, and although sectional views of vestured pits are infrequent, they are very informative. Studies that explore diversity from one order or family to another are needed and offer opportunities for understanding the evolutionary significance of this feature.

Vestured pits are present on pits of secondary xylem vessels of all studied Echium species, roots as well as stems. Variations in vesturing presence do occur in the genus (across wide circular pit cavities; along margins of elliptical pit apertures; aggregated to various degrees; variously abundant), but these are difficult to define precisely and are related primarily to organography and ecology rather than to the taxonomic system. The Macaronesian species have been reported to form a single clade. Wood anatomical features other than vesturing are also closely keyed to species ecology and, in particular subclades, growth forms. Woodiness, exemplified by most species of the insular clade of Echium, has been claimed to have evolved as a mechanism to promote outcrossing on islands. This alleged indirect selection is questioned here. The bases for secondary woodiness on island areas are multiple and are reviewed here (lessened temperature and moisture extremes; retention of branches instead of reproducing by seed as a form of economy; lack of mammalian herbivores; superior dispersal and colonization abilities of less woody clades; ability to occupy geologically new or recently altered areas). These features may, individually and collectively, serve to increase the ability of species to become woody on islands. Vesture presence can be cited for some groups that have radiated well on Macaronesian islands (Boraginaceae, Brassicaceae) as well as on certain major land areas (Acacia and Eucalyptus on Australia all have vestured pits on vessels). Plant species can be viewed as having an umbrella of features relevant to woodiness and growth form, with thresholds in tolerance of cold, drought, etc. In an island environment, there is alleviation, moderation, and transcendence of these thresholds and restrictions so that survival of vegetative structures into the next growing season is achieved. Advantageous radiation into new growth forms is made possible by this extension of the growing season.

Wood anatomical data for the 19 families of Brassicales are presented, based on light microscopy and scanning electron microscopy (SEM), arranged according to recent molecular phylogenetic evidence. Because of large species numbers and diversity in ecology and growth form, Brassicales are an ideal case study group for understanding wood evolution. Features newly reported include vestured pits in Cleomaceae, Koeberliniaceae, Pentadiplandraceae, Salvadoraceae, and Setchellanthaceae. Vesturing of primary xylem helices is shown for Raphanus (first report in angiosperms). Fiber dimorphism is newly reported in some genera of the crown group (Capparaceae + Cleomaceae + Brassicaceae). The fiber-tracheid is probably the ancestral imperforate tracheary element type for Brassicales, and from it, libriform fibers, living fibers (including septate fibers), and tracheids have likely been derived. The Baileyan concept of unidirectional evolution from tracheids to libriform fibers must have many exceptions in angiosperms, and tracheids are not uniform. Tracheids occur in Emblingiaceae, Koeberliniaceae, Pentadiplandraceae, Stixaceae, and Tropaeolaceae. Synapomorphies can be identified, as in the Akaniaceae-Tropaeolaceae clade (rays of two sizes, living fibers, scalariform perforation remnants) and the Moringaceae-Caricaceae clade (ground tissue of wood composed of thin-walled fibers or similar parenchymatous cells). Wood of Brassicales is mostly not paedomorphic, although paedomorphic characters suggesting secondary woodiness occur within the families Brassicaceae (abundance of upright ray cells, raylessness), Caricaceae, Cleomaceae, and Moringaceae. Brassicales are probably ancestrally woody, and wood of Sapindales and Malvales has a number of key character states (plesiomorphies) like those in Brassicales, as would be predicted by current molecular phylogenies. Surveys of large taxonomic groupings, such as Brassicales, tend to yield more examples of homoplasies and apomorphies that can be interpreted in terms of adaptation and functional interlinkage (e.g., ray evolution paralleling imperforate tracheary element evolution). In turn, these features can be interpreted in terms of ecology (e.g., xeric habitats) and growth forms (e.g., tree succulents). The assemblages of wood character information in a reasonably well known order of angiosperms permits hypotheses about wood evolution in angiosperms as a whole. Some of the more important hypotheses presented include: (1), that evolution of wood (and other) characters is always progressive, with gene overlays (silencing, modification, etc.) and simultaneous changes in multiple features, so that ancestral conditions are never truly re-attained. (2). Not all characters are of equal value in water economy of any given plant; some (presence of tracheids) may supersede others, and xeromorphic characters can be arranged relative to each other in tiers, although various taxonomic groups have different rosters of conductive safety features. (3). Heterochrony (protracted juvenilism, accelerated adulthood) is extensively represented in angiosperms, and acts as an overlay that is a source of diversity that angiosperms have drawn on since their inception (probably as minimally woody plants). (4). There may be no "purely taxonomic" characters, because genes of an organism relate primarily to changes, ancient and new, that are of adaptive significance, although we may not be able to detect selective value, past or present.Although many families of Brassicales are small and represent occupancy of specialized or extreme habitats (Batis, Koeberlinia, Moringa), active speciation in Brassicaceae and Capparaceae is related to tolerance of drought and cold with mechanisms such as vestured pits, narrow vessels, and abbreviation in life cycle length.

The diversity of expression in axial parenchyma (or lack of it) in woods is reviewed and synthesized with recent work in wood physiology, and questions and hypotheses relative to axial parenchyma anatomy are offered. Cell shape, location, abundance, size, wall characteristics and contents are all characteristics for the assessment of the physiological functions of axial parenchyma, a tissue that has been neglected in the consideration of how wood histology has evolved. Axial parenchyma occurrence should be considered with respect to mechanisms for the prevention and reversal of embolisms in tracheary elements. This mechanism complements cohesion-tension-based water movement and root pressure as a way of maintaining flow in xylem. Septate fibres can substitute for axial parenchyma ('axial parenchyma absent') and account for water movement in xylem and for the supply of carbohydrate abundance underlying massive and sudden events of foliation, flowering and fruiting, as can fibre dimorphism and the co-occurrence of septate fibres and axial parenchyma. Rayless woods may or may not contain axial parenchyma and are informative when analysing parenchyma function. Interconnections between ray and axial parenchyma are common, and so axial and radial parenchyma must be considered as complementary parts of a network, with distinctive but interactive functions. Upright ray cells and more numerous rays per millimetre enhance interconnection and are more often found in woods that contain tracheids. Vesselless woods in both gymnosperms and angiosperms have axial parenchyma, the distribution of which suggests a function in osmotic water shifting. Water and photosynthate storage in axial parenchyma may be associated with seasonal changes and with succulent or subsucculent modes of construction. Apotracheal axial parenchyma distribution often demonstrates storage functions that can be read independently of osmotic water shifting capabilities. Axial parenchyma may serve to both enhance mechanical strength or, when parenchyma is thin-walled, as a tissue that adapts to volume change with a change in water content. Other functions of axial parenchyma (contributing resistance to pathogens; a site for the recovery of physical damage) are considered. The diagnostic features of axial parenchyma and septate fibres are reviewed in order to clarify distinctions and to aid in cell type identification. Systematic listings are given for particular axial parenchyma conditions (e.g. axial parenchyma 'absent' with septate fibres substituting). A knowledge of the axial parenchyma information presented here is desirable for a full understanding of xylem function. (C) 2015 The Linnean Society of London.

Raylessness occurs in several hundred species belonging to about 40 families (fewer depending on taxonomic delineation). Fibre distribution (raylessness at first, followed by origin of rays), fibre wall thickness and sclerenchyma at pith margins support the idea that rapid acquisition of mechanical strength is basic to most instances of raylessness. Raylessness may be the most readily available process for achieving mechanical strength in ancestrally herbaceous groups lacking large amounts of phloem and cortical fibres. Raylessness is not a uniform phenomenon and a small number of instances suggest alternative causation, as in two lianas (Cobaea, Thunbergia). Raylessness occurs in only a small number of trees and annuals, but is found in woody herbs, subshrubs and some shrubs. It is indicative of secondary woodiness and wood paedomorphosis. Raylessness would seem to block the radial flow that rays typically provide, but a surprising number of rayless woods have moderately pitted fibres (indicative of flow) and septate or non-septate living fibres. Three-dimensional networks of conjunctive tissues in rayless species with successive cambia (Aizoaceae, Amaranthaceae, Nyctaginaceae) could also provide radial flow avenues. Ontogenetic changes from raylessness to ray presence within the stem of a given species are described and illustrated. Pseudo-raylessness, late-onset raylessness and early-onset raylessness are recognized. Systematic distribution and pertinent literature are given for known instances of raylessness and pseudo-raylessness. Raylessness shows that wood evolution involves not merely change in the abundance and position of cell types, but also redesign and diversification in cell types. (c) 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 529-555.

SEM studies of tracheary elements of subfamily Orontioideae (Lysichiton, Orontium, Symplocarpus) of Araceae show unexpected features. The plants are entirely vesselless. There are small pores in pit membranes of end walls of tracheids in roots and stems, but pit membranes remain intact. End wall pit membranes of stems have a coarse fibrillar texture, somewhat reminiscent of (but different from) those of Nymphaeaceae and Cabombaceae. Acoraceae, which are also vesselless, represent the first branch of the monocot tree, according to phylogenies, and the orontioids form the next branch. Vessellessness is therefore a potentially plesiomorphic feature in monocots, but it may also be related to the highly mesic habitats of Acoraceae and the orontioids. Various other non‐submersed monocots have vesselless or near‐vesselless xylem. Sectioned xylem of Orontioideae is also very suggestive of stages in the development of the pit membranes of both end walls and lateral walls of tracheids: open networks of cellulosic fibrils apparently precede the addition of denser fibrillar meshes, key information in assessing to what extent perforations in scalariform perforation plates of vascular plants may stop formation at the open network stage, and to what extent a thicker pit membrane experiences lysis and disintegration as the vessel element matures.

Dimorphic fibres in angiosperm woods are designated when zones of two different kinds of fibres can be distinguished in transverse sections. The usage of most authors contrasts wider, thinner-walled, shorter (sometimes storied) fibres with narrower, thicker-walled fibres that have narrower lumina. The wider fibres can be distinguished in longitudinal sections from axial parenchyma, which usually consists of strands of two or more cells each surrounded by secondary walls (and thus different from septate fibres). This phenomenon occurs in some Araliaceae, Asteraceae, Fabaceae, Myrtales (notably Lythraceae), Sapindales (especially Sapindaceae), Urticales and even some Gnetales. Additional instances can doubtless be found, especially if instances of wide latewood fibres together with narrow earlywood fibres are included. There is little physiological evidence on differential functions of dimorphic fibres, except in Acer, in which hydrolysis of starch in the wide fibres is known to result in transfer of sugar into vessels early in the growing season. Starch storage in axial parenchyma may, in a complementary way, serve for embolism reversal and prevention and thus for maintenance of the water columns. Crystalliferous fibres (Myrtales, Sapindales) can be considered a form of fibre dimorphism that deters predation. Gelatinous fibres, often equated with tension wood, can also be considered as a form of fibre dimorphism. The evolutionary significance of fibre dimorphism is that a few small changes in fibre structure can result in the accomplishment of diversified functions.(c) 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174, 44-67.

Early angiosperms were minimally woody; increase in woodiness and changes in wood histology yielded trees, lianas, and shrubs in various clades. Many eudicot herbs have been derived from variously woody ancestors. Some of those derivatives have, at various stages, evolved secondary woodiness to various degrees. Categories of information by which we can trace these progressions are presented: length-on-age curves for vessel elements, perforation plate morphology, ray histology, DNA-based phylogenies, geological and ecological factors, dispersal capabilities, and speciation ability. Trajectories that angiosperms have followed are analyzed in terms of growth forms: sympodial habits, cane shrubs, lianas, trees, various herb-related forms, stem succulents, and plants with successive cambia. Phylogenetic modalities that are related to degree of woodiness are discussed: retention of and departure from juvenile wood features in basal angiosperms, overlay effects (additive or modifying effects of factors on woodiness), character independence and interdependence, and degrees and types of transitions between more woodiness and less woodiness. Production of procumbent ray cells (which excel at radial conduction) is the result of not just subdivision of ray initials but also infrequent tangential divisions in ray initial derivatives. In juvenilistic woods, this process runs in parallel with shortening of fusiform cambial initials, but in woodier species, fusiform cambial initials become longer over time whereas ray initials become vertically shorter. Examples and original information on eudicot woods are mostly from orders and families of the campanulid clade. Juvenile features are multiple, with each capable of being retained, modified, or lost independently. This article takes the form of an eclectic essay that includes original data and observations, hypotheses, and critiques as well as presenting questions and syntheses, and it supplements previous articles by the author.

Interxylary phloem is here defined as strands or bands of phloem embedded within the secondary xylem of a stem or root of a plant that has a single vascular cambium. In this definition, interxylary phloem differs from intraxylary phloem, bicollateral bundles, pith bundles, and successive cambia. The inclusive but variously applied terms included phloem and internal phloem must be rejected. Histological aspects of interxylary phloem are reviewed and original data are presented. Topics covered include duration of interxylary phloem; relationship in abundance between sieve tubes in external phloem and interxylary phloem; distinctions between interxylary and intraxylary phloem; presence of parenchyma, fibers, and crystals in the interxylary phloem strands; development of cambia within interxylary phloem strands; three-dimensionalization and longevity of phloem, systematic distribution of interxylary phloem; physiological significance; and habital correlations. No single physiological phenomenon seems to explain all instances of interxylary phloem occurrence, but rapidity and volume of photosynthate transport seem implicated in most instances.

Qualitative and quantitative data are given for wood anatomy of six of the 11 recognized species of Forchhammeria (Mexico, Central America, West Indies), a genus formerly placed in Capparaceae. Though still in Brassicales, the genus has been excluded, along with several other genera, from the major recognized families of that order on the basis of molecular data. Liquid-preserved material of several species permitted detailed histological accounts of the successive cambia and their development in the stems of Forchhammeria. Successive cambia have a curious distribution in Brassicales that may represent homoplasies. Most wood features of the genus do not appear highly xeromorphic, but presence of tracheids as a wood background tissue and abundance of starch and perhaps water storage in ray parenchyma and conjunctive tissue can be cited as mechanisms likely to resist embolism formation. Forchhammeria retains green leaves throughout the dry season. Forchhammeria tamaulipana, known only from Tamaulipas State, Mexico, the single species of a new subgenus, Pauciflora, is newly described. Its embryos have nearly equal cotyledons and germinate epigeously, whereas all remaining species of the genus are pseudomonocotylous and hypogeous. These and other distinctive features of F. tamaulipana may prove significant in providing links to other brassicalean genera. The family name Stixaceae Doweld is now appropriate for Forchhammeria, Neothorelia, Stixis, and Tirania.

CAIU.QUJST, S.: Wood anatomy and relationships of Stackhousiaceae. Bot. J ahrb. Syst. t 08: 473-4BO. 1987. ISSN 0006-8152. Quantitative and qualitative features are described for five species of Stackhousia. The imperforate tracheary elements are all tracheidlike fiber-tracheids. Very narrow fusiform vessel elements are present. All other vessel elements are relatively narrow, and numerous per mm2 of transection. AU of these features connote xeromorphy. Quantitatively the xeromorphy is not extreme, a fact thought to relate to the habit in Stackhousia of stems to die back to a woody base annually, as well as to drought-deciduous nature of foliage and short-lived natured of plants. Wood features of Stackbousia are compatible with the concept that the family belongs to Celastrales. Possession of fiber·tracheids, as well as other features, favor this hypothesis over the idea of re.lationship to Scrophulariaceae or Lobeliaceae.