Systematic and phylogenetic value of wood anatomy in Heteromorpheae (Apiaceae,Apioideae)

The wood anatomy of all four woody genera of the tribe Heteromorpheae (Apiaceae, subfamily Apioideae) has been described and compared, based on 40 wood samples (representing nine species of Anginon, one species of Glia, three species of Heteromorpha and two species of Polemannia). The four genera were found to be relatively similar in their wood anatomy. Helical thickenings on the vessel walls occur in all species investigated and appear to represent an ancestral character state and a symplesiomorphy for the tribes Bupleurieae and Heteromorpheae. Each of four genera has a diagnostically different combination of character states relating to the diameter of vessels, size of intervessel pits, length of fibres, presence and arrangement of banded axial parenchyma, size of rays and ray cells, and presence of septate fibres and crystals in the ray cells. The occurrence of marginal axial parenchyma in Anginon and Glia may be an additional synapomorphy for these taxa. Variation in the wood anatomy of 31 samples from nine species of Anginon is not correlated with habitat (Fynbos or Succulent Karoo Biomes), but instead appears to reflect adaptations to seasonal aridity found in both ecosystems. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 569–583.     

Systematic and ecological wood anatomy of Neotropical Schefflera (Araliaceae), with an emphasis on the Didymopanax group

Wood anatomy has been investigated from 35 species belonging to the Neotropical clade of the polyphyletic genus Schefflera (Araliaceae), representing three of the five subgroups (Didymopanax, Crepinella and Sciodaphyllum). The species examined are rather uniform in their wood structure, sharing the presence of scalariform and simple perforation plates, septate fibres and scanty paratracheal axial parenchyma. The observed variation in many wood characters showed statistically significant differences relative to latitude, climate and, especially, vegetation types. In particular, the intervessel pits are larger in species from higher latitudes and in seasonally dry habitats than those from lower latitudes and rainforests. Latitudinal and ecological trends in the variation of vessel element lengths, bar numbers on perforation plates, intervessel pit sizes and ray widths may be at least partially explained as effects of adaptation to drier environments in the course of dispersal outside the Amazonian region and diversification in the Atlantic Forest subclade and the Savannic subclade within the Didymopanax group. The occurrence of a granular annulus on the intervessel pit membranes in S. chimantensis and S. sprucei (both of the Crepinella group) is the first record of this feature in Araliaceae. In comparisons of Neotropical Schefflera with the other major clades of Schefflera sensu lato, wood anatomical diversity is consistent with the polyphyly of this genus based on molecular phylogenetic analyses. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 452–475.     

Living cells in wood. 1. Absence,scarcity and histology of axial parenchyma as keys to function

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. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 291–321.     

Therps16 intron and the phylogeny of theRubioideae (Rubiaceae)

The phylogeny of the subfamilyRubioideae (Rubiaceae) was estimated from sequence variation in therps16 intron (cpDNA) in 143 ingroup and 5 outgroup taxa. The analysis largely confirms a recent one based onrbcL sequences, but branch support is often much stronger. Three of the traditional subfamilies are supported,Rubioideae, Cinchonoideae s. str., andIxoroideae s. l. while there is no support forAntirheoideae. TheRubioideae are the sister group of all otherRubiaceae and comprise the tribesAnthospermeae, Coccocypseleae, Cruckshanksieae, Coussareeae, Gaertnereae, Hedyotideae, Knoxieae, Morindeae, Ophiorrhizeae, Paederieae, Pauridiantheae, Perameae, Psychotrieae, Rubieae, Spermacoceae, Theligoneae, andUrophylleae. TheHamelieae andHillieae belong to theCinchonoideae. Rachicallis andSiemensia should be transferred from theHedyotideae to theCinchonoideae. ThePauridiantheae, Urophylleae, Ophiorrhizeae, andRaritebe form the basalmost subclade of theRubioideae. The second basalmost clade consists of the generaLasianthus andPerama. The third basalmost clade consists of the tribesCoussareeae, Coccocypseleae andCruckshanksieae, and the generaDeclieuxia andHindsia. The tribesKnoxieae, Anthospermeae, Argostemmateae, Paederieae, Theligoneae, Rubieae, Hedyotideae, andSpermacoceae are members of one clade. TheKnoxieae are monophyletic ifOtiophora, Otomeria, andPentas are included. The tribeAnthospermeae is supported as monophyletic, but its subtribes are not. ThePaederieae, together withTheligonum, form a paraphyletic grade basal to theRubieae. TheHedyotideae, includingSchismatoclada, form a grade at the base of theSpermacoceae. TheGaertnereae are monophyletic and distinct from thePsychotrieae. TheMorindeae are monophyletic and includeDamnacanthus andMitchella. Schradera is the sister group of theMorindeae. ThePsychotrieae are monophyletic when theGaertnereae, Lasianthus, andDeclieuxia are excluded. The recognition of a subtribeHydnophytineae leaves the rest of thePsychotrieae paraphyletic.Psychotria is paraphyletic with respect to all other genera of the tribe. Approximately 50 genera are here classified for the first time based on molecular data.     

WOOD ANATOMY OF TREMANDRACEAE: PHYLOGENETIC AND ECOLOGICAL IMPLICATIONS

Stems of four species of the Australian family Tremandraceae furnished sufficient material for analysis of wood anatomy. Presence of simple perforation plates on vessel elements, occurrence of libriform fibers (some septate), tendency toward vasicentric parenchyma, presence of crystalliferous axial parenchyma strands, presence of crystals singly in ray cells, and occurrence of amorphous deposits in parenchyma are all features in which Tremandraceae resemble Pittosporaceae. Wood anatomy tends to support a “rosoid” rather than a sapindalean, rutalean, or polygalalean affinity for Tremandraceae, although wood is only a preliminary indicator. By the use of numerical indices as well as such indicators as helical thickening and presence of vascular tracheids, wood of Tremandraceae is shown to be highly xeromorphic. The genus Tremandra may represent a secondary entrant into wet forests of southwestern Australia; it clearly is not relict from mesic ancestry.     

Molecular phylogenetics and generic assessment in the tribe Morindeae (Rubiaceae–Rubioideae): How to circumscribe Morinda L. to be monophyletic?

Most of the species of the family Rubiaceae with flowers arranged in head inflorescences are currently classified in three distantly related tribes, Naucleeae (subfamily Cinchonoideae) and Morindeae and Schradereae (subfamily Rubioideae). Within Morindeae the type genus Morinda is traditionally and currently circumscribed based on its head inflorescences and syncarpous fruits (syncarps). These characters are also present in some members of its allied genera, raising doubts about the monophyly of Morinda. We perform Bayesian phylogenetic analyses using combined nrETS/nrITS/trnT-F data for 67 Morindeae taxa and five outgroups from the closely related tribes Mitchelleae and Gaertnereae to rigorously test the monophyly of Morinda as currently delimited and assess the phylogenetic value of head inflorescences and syncarps in Morinda and Morindeae and to evaluate generic relationships and limits in Morindeae. Our analyses demonstrate that head inflorescences and syncarps in Morinda and Morindeae are evolutionarily labile. Morinda is highly paraphyletic, unless the genera Coelospermum, Gynochthodes, Pogonolobus, and Sarcopygme are also included. Morindeae comprises four well-supported and morphologically distinct major lineages: Appunia clade, Morinda clade (including Sarcopygme and the lectotype M. royoc), Coelospermum clade (containing Pogonolobus and Morinda reticulata), and Gynochthodes–Morinda clade. Four possible alternatives for revising generic boundaries are presented to establish monophyletic units. We favor the recognition of the four major lineages of Morindeae as separate genera, because this classification reflects the occurrence of a considerable morphological diversity in the tribe and the phylogenetic and taxonomic distinctness of its newly delimited genera.     

Fibre dimorphism: cell type diversification as an evolutionary strategy in angiosperm woods

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. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 44–67.     

ENDOTHECIUM IN IRIDACEAE AND ITS SYSTEMATIC IMPLICATIONS

An examination of the endothecial thickenings in 44 species of Iridaceae, selected from the four subfamilies and all major tribes, provides useful information about generic and tribal relationships in the family. U-shaped thickenings occur in Nivenioideae and Iridoideae—Sisyrinchieae, the latter the least specialized tribe of its subfamily. The occurrence of helical thickenings in all members examined of Iridiodeae tribes Irideae, Mariceae, and Tigridieae (a putatively monophyletic group) and Ixioideae is consistent with the recognition of these two lines as distinct taxa based on anatomical, morphological, phytochemical, and in the case of Ixioideae, palynological criteria. Baseplate thickenings are restricted to Patersonia. However, the shrubby Cape genera—Nivenia, Klattia, and Witsenia—have U-shaped thickenings which show a tendency for the bars on the inner periclinal cell walls to anastomose, suggesting a trend towards the baseplate condition in Patersonia. This accords with a suggested relationship between these genera, based on anatomical and flavonoid similarities. The pattern of variation in endothecial thickenings in Iridaceae is consistent with the phylogeny proposed by Goldblatt (1990). The distribution of thickening types within the family does not make it possible to polarize this character, but the most parsimonious interpretation assumes that U-shapes are basic. However, in at least some other monocotyledonous families the pattern suggests that U-shaped thickenings are derived from helices.     

WOOD ANATOMY AND RELATIONSHIPS OF LACTORIDACEAE

Mature wood of Lactoris, not previously available for study, reveals ten distinctive characters: vessels with simple perforation plates; vessels in pore multiples; vessel-to-axial parenchyma pits scalariform or transitional, vessel-to-vessel pits alternate; fiber-tracheids with vestigial pits; fiber-tracheids, vessels, and axial parenchyma storied; axial parenchyma vasicentric scanty; axial parenchyma either not subdivided or, if subdivided, with thin nonlignified walls between the cells (like the septa in septate fibers); rays wide and tall, little altered during ontogeny; ray cells upright; and ray cells taller adjacent to fascicular areas. All of these features occur in woods of Piper and other Piperaceae. The systematic position of Lactoris is therefore reassessed. Evidence available to date is consonant with placement of Lactoridaceae in Piperales, in which it would be more primitive than Piperaceae or Saururaceae. Features cited as evidence for alternative placements of Lactoridaceae are reviewed.     

PATTERNS OF ENDOTHECIAL WALL THICKENINGS IN ARACEAE: SUBFAMILIES POTHOIDEAE AND MONSTEROIDEAE

A survey of the patterns of endothecial wall thickenings in 106 representative species from 20 genera in the Pothoideae and Monsteroideae was made using cleared anthers, sections and macerations. The wide variety of wall thickenings that is present is based on an annular-helical pattern. Variations in thickenings are related to differences in cell shape, cell orientation, intergradation between helical and annular patterns, pitch of helices, presence of branched thickenings, and various types of discontinuities in thickenings. Notable exceptions to the annular-helical pattern include Culcasia, which lacks a differentiated endothecial layer with thickenings, and Acorus, which has a peculiar stellate pattern that is unique in the family. No single pattern consistently characterizes either subfamily, although continuous helices are common in the Monsteroideae, and rare in the endothecium of Pothoideae (except Anadendrum). Monsteroideae frequently exhibit a series of slanted separate thickenings on anticlinal walls, which is absent from Pothoideae except in Heteropsis. The slanted pattern is considered a variation on a rectangular helix, involving discontinuities of thickenings on the periclinal walls. Some monsteroid genera show considerably more interspecific variation (Rhaphidophora) than others (Monstera). Endothecial thickenings constitute an anatomical character that is useful in the systematic study of Araceae; present results support other anatomical studies in identifying Culcasia and Acorus as highly divergent genera in the Pothoideae.     

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.     

Cell wall structure in the xylem parenchyma ofCryptomeria

Summary Cell wall structure in ray and axial parenchyma cells in the wood ofCryptomeria was shown to be typically crossed polylamellate and dissimilar to the characteristically layered wall of fibers and tracheids. Ray cells differed from axial cells in terms of form and also in the relative inclination of crossed microfibrillar helices in the cell wall. This feature was reflected by positive birefringence in ray cells and negative birefringence in axial cells. Localized wall thickenings,viz. transverse bars in ray cells and longitudinal ribs in axial cells, also displayed crossed polylamellate structure. This observation contrasts with the exclusively longitudinal microfibrillar orientation previously reported for longitudinal ribs in elongated parenchyma cells of primary tissue. On the basis of similar microfibrillar orientations between outer and inner wall lamellae, the cell walls ofCryptomeria parenchyma were judged to be predominantly secondary.Lignin was heterogeneously distributed in lamellate fashion and a high concentration characterized the thin middle lamella. Both types of parenchyma suggested a higher lignin content than adjacent longitudinal tracheids.     

Resistance of parenchyma cells in wood to degradation by brown rot fungi

Wood degradation by two basidiomycetes, Fomitopsis pinicola and Laetiporus sulphureus was studied in one conifer and four broadleaved trees: Picea abies (Norway spruce), Acer pseudoplatanus (sycamore), Betula pendula (birch), Quercus robur (common oak) and Robinia pseudoacacia (robinia). Observations of birefringence under polarized light showed that in all hosts both brown rot fungi affected cells of the early wood before those of the late wood. Degradation of cellulose, as shown by the loss of birefringence, was apparent after 6 weeks in the cell wall of fibres and fibre tracheids, but even after 12 weeks, axial parenchyma showed no signs of degradation. The results indicate that both brown rot fungi cause higher weight losses in hosts (P. abies and B. pendula) with a small amount of parenchyma cells, whereas the lowest weight losses are associated with wood containing a high amount of parenchyma cells (Q. robur and R. pseudoacacia). Resistance of parenchyma cells to degradation by brown rot fungi appears to be related to the cell wall morphology of parenchyma cells and may also reflect a low co-evolutionary adaptation of brown rot fungi to the xylem of broadleaved trees.     

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.     

Gaertnera and Pagamea: Genera Within the Psychotrieae or Constituting the Tribe Gaertnereae? A Wood Anatomical and Palynological Approach

The possible alliance between Gaertnera and Pagamea (Rubiaceae-Rubioideae), two genera from the Old and New World, respectively, is investigated on the basis of wood anatomy and pollen morphology. Nowadays, the main point of discussion about the taxonomic position of these genera is whether they belong to the Psychotrieae or constitute a tribe Gaertnereae characterised by their secondarily superior ovary and sheathing stipules. Both the wood and pollen of the genus pair are found to show specific features absent in other genera of the Psychotrieae, e.g. parenchyma bands in the xylem and endexine thickenings on the polar sites of the pollen apertures. Nevertheless Gaertnera and Pagamea share many other characters with the Psychotrieae. Wood and pollen convincingly demonstrate the very close affinity of the two genera. The sister pair differs in so many features from other Psychotrieae, that Gaertnera and Pagamea should constitute at least a subtribe Gaertnerinae, formally recognized here. The general lack of profound studies on the affinities within the very large tribe Psychotrieae makes further comments on the taxonomic relationships of the Gaertnerinae difficult.     

The Ecological Secondary Xylem Anatomy of 7 Desert Species of Leguminosae

The wood anatomy of 7 species (Ammopiptanthus mongolicus, Amorpha fruticosa, Halimodendron halodendron, Hedysarum mongolicum. Hedysarum scoparium, Lespedeza bicolor and Robinia pseudoacacia)of Leguminosae, which grow in desert regions of Northern China, is described in details. A comparative study on the quantitative wood anatomical characters among the species is made. Except some anatomical characters in A. fruticosa were larger in vessel diameter, thin walled in vessels and libriform fibres, all the rest six species showed a general similarities:vessel frequency/sq, mm very numerous and percentage of multiple vessels high; vessel elements very short, perforations simple and in almost horizontal end walls, intervessel bordered pits alternate and vestured; libriform fibres very short, and usually with thickened walls, and with simple pits; average ray height very low, and with multiseriate as well as uniseriate. However, there are differences in other characters, e. g. vessel distribution, percentage of solitary vessels; spiral thickenings present or absent; amount of axial parenchyma and distribution; ray frequency and type; crystals present or absent, and crystal distribution, if present. According to these anatomical diversities, a key to the identification of the 7 species is given. In this article, the relation between the structure of wood and the environmental influences has been discussed.     

A cytoskeletal basis for wood formation in angiosperm trees: the involvement of cortical microtubules

Rearrangements of cortical microtubules (CMTs) during the differentiation of axial secondary xylem elements within taproots and shoots of Aesculus hippocastanum L. (horse-chestnut) are described. A correlative approach was employed using indirect immunofluorescence microscopy of α-tubulin in 6- to 10-μm sections and transmission electron microscopy of ultrathin sections. All cell types – fibres, vessel elements and axial parenchyma – derive from fusiform cambial cells which contain randomly oriented CMTs. At the early stages of development, fibres and axial parenchyma cells possess helically arranged CMTs, which increase in number as secondary wall thickening proceeds and simple pits develop. In contrast, incipient vessel elements are distinguished by the marking out of sites of bordered pits; these sites first appear as microtubule-free regions within the reticulum of randomly oriented CMTs that characterises their precursor fusiform cambial cells. Subsequently, the ring of CMTs which develops at the periphery of the microtubule-free region decreases in diameter as the over-arching pit border is formed. Like bordered pits, large-diameter, non-bordered pits (contact pits) which develop between vessel elements and adjacent contact ray cells originate as microtubule-free regions and are also associated with development of a ring of CMTs at the periphery. In the case of contact pits, however, there is no reduction in the diameter of the CMT ring during pit development. Tertiary cell wall thickenings are also a feature of vessel elements and appear to form at sites where bands of laterally associated, transversely oriented CMTs, separated from each other by microtubule-free zones, are found. Later, these bands of CMTs become narrower, and separate into pairs of microtubule bundles located on each side of the developing wall thickening. Development of perforations between vessel elements is also associated with the presence of a ring of CMTs at their periphery. Received: 13 July 1998 / Accepted: 30 November 1998     

WOOD FORMATION IN PROSOPIS: EFFECT OF 2,4-D, 2,4,5-T,AND TIBA

Treatment of erect stems of Prosopis with near phytotoxic levels of 2,4-D or 2,4,5-T causes the formation of an unusual wood with narrow, thick-walled vessels and axial parenchyma in which cell wall thickening is inhibited. Although reduced in diameter, the vessels formed during 2,4-D and 2,4,5-T treatment are so numerous that there is no significant difference between phenoxyacetic acid and control seedling groups with regard to total area of xylem occupied by vessels. The preferential maturation of xylem vessels over parenchyma and the transformation of fusiform initials into septate parenchyma strands in phenoxyacetic acid-treated Prosopis resemble the structural changes reported to occur after girdling in the cambial tissue of other arborescent angiosperms. Bending experiments indicate that tension-wood fibers of Prosopis differentiate in response to an auxin deficiency. However, xylogenesis in erect stems treated with TIBA is affected such that a significantly higher proportion of the cambial cell population becomes axial xylem parenchyma.     

Comparative wood anatomy of the Cupressaceae and correspondence with phylogeny,with special reference to the monotypic taxa

Wood anatomy is one of the tools used for taxonomic classification of species. By combining this tool with molecular phylogeny, the current groupings made in morphological studies can be discussed. This study describes the wood anatomy of the monotypic genera of Cupressaceae and analyses the features that could represent synapomorphies of the principal clades recovered by molecular phylogeny. The wood anatomical study shows the high homogeneity of this family, revealing the presence of common ancestral features that support the union between Taxodiaceae and Cupressaceae s.s. and the separation of Sciadopitys. It also supports the group formed by Taxodiaceae in Cupressaceae s.l. No clear division was observed between the callitroid and cupressoid clades. Some wood anatomical differences were observed in the Fitzroya–Diselma–Pilgerodendron and Microbiota–Platycladus–Tetraclinis associations. The wood anatomical features common to the family, such as axial tracheids without helical thickenings, homogeneous rays, cupressoid cross-field pits and the presence of a warty layer, are put forward as possible synapomorphies for Cupressaceae s.l. The clade-specific synapomorphies are taxodioid cross-field pits for taxodioid and sequoioid clades, absence of a well-defined torus in Thuja–Thujopsis and torus extensions in Diselma–Fitzroya–Widdringtonia.