Fossil forms of Amentiferae

Review of the procedures used in determining fossil plant organs indicates that the many Cretaceous records of extant genera of “Amentiferae” based on leaves should be rejected as theoretically unreliable. Palynological data, in combination with some valid megafossil data, indicate that most recognizable members of “Amentiferae” are no older than the later part of the Late Cretaceous. Juglandales appear to be derivatives of the ancient Normapolles complex and unrelated to other “Amentiferae.” A preliminary account of some of the comparative foliar morphology of extant “Amentiferae” indicates that some—particularly Betulaceae and Fagaceae—are closely related to Hamamelidales but that other families—notably Rhoipteleaceae, Juglandaceae, Didymelaceae, and Leitneriaceae—are unrelated to this order.     

马尾树科及其近缘科花粉形态研究

本文观察了马尾树科(Rhoipteleaceae)、胡桃科(Juglandaceae)以及桦木科(Be-tulaceae)、9属9种植物花粉形态。马尾树科与桦木科桦木属(Betula L.)花粉粒均为扁球形,极面观呈钝三角形,萌发孔类型均为多孔类型,孔之间均有弓形加厚,而且外表面均具细颗粒状雕纹。而胡桃科大多为多边形、散孔类型。根据花粉形态资料,探讨了马尾树科的系统位置。     

The “Amentiferae” or Hamamelidae as an artificial group: A summary statement

The various contributions to this symposium on the “Amentiferae” reach the general conclusion that the group is an artificial aggregation of taxa of diverse origin that have converged to a common evolutionary plateau in possessing a large syndrome of characteristics that adapt them for successful cross-pollination by wind. Aside from those few families (Eucommiaceae, Casuarinaceae, Fagaceae, and Betulaceae) that apparently do have close relationships (close common origin) with each other and with the Hamamelidales, the following taxa should be removed from the Hamamelidae: Juglandales (Juglandaceae and Rhoipteleaceae) to the Rutales as the Juglandineae near the Anacardiineae; Myricaceae and Leitneriaceae respectively to the Myricales and Leitneriales near the Rutales in the Rutiflorae; Urticales (excludingBarbeya andEucommia) to the Malviflorae near the Malvales and Euphorbiales;Picrodendron to the Euphorbiaceae; Didymelaceae to the Euphorbiales; Myrothamnaceae to the Brunineae of the Pittosporales; andBalanops, Barbeya, andCanacomyrica, along withBatis, to “taxa incertae sedis.”     

Fossil wood of the Juglandaceae: Some questions of taxonomy,evolution, and phylogeny in the family based on wood anatomy

Some problems in the taxonomy of the Juglandaceae are discussed based on wood anatomy; the identification of fossil juglandaceous wood is considered. Data on fossil wood of the Juglandaceae are summarized; a key for identification of wood anatomy in modern and fossil Juglandaceae is compiled. Wood anatomical characters in members of the family are discussed in the light of major evolutionary trends in the secondary xylem of dicots, and a comparative characterization of members of the family is developed. A hypothesis is proposed that the subfamily Engelhardioideae is the most primitive member of the Juglandaceae based on wood anatomy, the tribe Juglandeae and subfamily Platycaryoideae are slightly more highly specialized, and the tribe Hicorieae is the most advanced. Evolutionary relationships between the members of the Juglandaceae are reviewed based on wood anatomy.     

A Preliminary Study on the Wood Anatomy of Manglietia aromatica Dandy [

The present paper deals mainly with the wood structures of “Paramanglietia aromatica” Hu & Cheng and Paramichelia (Pierre) Hu of the Magnoliaceae. These wood structures are listed in Table Ⅰ. In comparison of their wood anatomical features with relsted genera[4], we find that (Ⅰ) the wood anatomy of the reduced genus “Para- manglietia” Hu et Cheng is still within the range of Manglietia Dandy and that (2 the wood anatomical features of Paramichelia (Pierre) Hu is very similar to those of Michelia L. and Tsoongiodendron Chun. We also find that the wood anatomy of Manylietia aromatica Dandy [“Parawanglietia aromatica” (Dandy) Hu & Cheng] is less similar to Manglietiastrum Law (see Table 1).     

Flowers and inflorescences of the “Amentiferae”

The floral and inflorescence morphology of the major genera of the Myricaceae, Betulaceae (including Corylaceae), Fagaceae, Leitneriaceae, and Juglandaceae are reviewed. Major problems in interpretation of morphology are examined in the light of various comparative morphological studies as well as ontogenetic and vascular anatomical studies. Basically similar phenomena associated with miniaturization of the partial inflorescence have led to superficially similar morphological patterns. The partial inflorescences in the various families, in spite of their reduced size, can be adequately analyzed in most cases on the basis of the bract-branch relationship. The highly modified morphology of the floret is clarified by the application of the general tenets of the leaf-stem relationship in the frame of reference of the minute absolute size of the floret. Numerous problems remain to be attacked. The total and partial inflorescences and the florets of the Myricaceae, Betulaceae, Fagaceae, Leitneriaceae and Juglandaceae are reviewed in terms of external morphology, vascular anatomy and ontogeny as reported in the more objective literature on the subject. The total inflorescences in these families range from the complex, androgynous panicles of stiff spikes of such genera asCastanopsis to the condensed, bud-like pistillate spikes of some Myricaceae andCorylus on the one hand, to the simple staminate floret in the axil of the foliage leaf in some species ofNothofagus on the other. In many species of these families the inflorescence is the apparently simple spike with a flaccid axis, the ament, but so often is this not the case that the designation “Amentiferae” for this artificial assemblage must be considered a misnomer. Whether the total inflorescences are composed of racemose or cymose partial inflorescences is a question not completely answered in all the families. In the Betulaceae the partial inflorescence has long been taken to be a cymule. But, a re-interpretation of the vascular anatomy suggests the alternative that the most distal floret in a short raceme has overtopped the axis of the partial inflorescence thus producing a pseudo-cymule. This is similar to a recent interpretation of the staminate partial inflorescence ofMyrica esculenta, where the individual floret is composed of a single stamen. The partial inflorescence in the more reduced species ofMyrica is thus a pseudanthium. Recent ontogenetic studies in the Betulaceae dramatically corroborate the earlier interpretation, based on vascular anatomy, that the staminate partial inflorescence ofOstrya is three-flowered. On similar grounds it has recently been shown that the spiny “involucre” of pistillateComptonia is composed of tertiary bracts. The structure of the staminate partial inflorescences in the Fagaceae seems reasonably clear except in certain species ofNothofagus where it may well be a synanthium, although the alternative of chorisis exists. The interpretation of the pistillate partial inflorescence inLeitneria requires re-study; the unvascularized tepal-like structures subtending the ovary have been alternatively treated as bracts -an ontogenetic study is badly needed. The organization of the staminate partial inflorescence of the Juglandaceae remains equivocal, although recent ontogenetic work on one species ofJuglans shows that the primordia of the secondary bracts are readily distinguished from tepal primordia, although at anthesis they are very similar. At present the number of florets in the partial inflorescence of the Juglandaceae remains an open question in spite of a fragmentary study of the vascular system. The cupule ofLithocarpus andQuercus continues to present a major morphological problem. The valves of the husk in other genera of the Fagaceae seem, on the basis of the vascular anatomy and some ontogenetic information, to be axes of the ultimate order of branching. A thorough study of these complex structures is needed. Staminate florets which are set off by tepals are readily identified with the reservation that those of some species ofNothofagus and ofJuglans, for instance, may be more complex than they seem. The absence of tepals creates major difficulties which have been resolved in some instances by the study of the vascular anatomy and/or ontogeny. But many problems remain. The pistillate floret seems clearly delimited in the various families. There continues to be the usual conflict concerning the proper interpretation of the wall of the inferior ovary, whether on the basis of ontogeny it should be considered cauline or on the basis of the vascular anatomy it is to be considered appendicular. Oddly enough there are also diametrically opposed interpretations of placentation -is it axile or parietal in one and the same species. This perhaps results from a conceptual conflict. The basal ovule, as in the Myricaceae, or even the ovules perched on a partial septum, as in the Juglandaceae, are similarly much discussed. The ontogenists tend to agree that such ovules are cauline, while the anatomists find that the complex vascular system is not that of a stele. There is a multitude of discrepancies, as yet, in observations, and even when there is mutually accepted fact, there are often conflicting interpretations. Above all, there is a massive lack of knowledge of the vascular anatomy and ontogeny of these miniature and modified flowers and inflorescences.     

A Phylogenetic Analysis of Families in the Hamamelidae

A cladistic analysis of the families in the Hamamelidae is made in the present paper. As a monophyletic group, the subclass Hamamelidae includes 19 families, namely, the Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Eupteleaceae, Eucommiaceae, Hamamelidaceae (incl. Rhodoleiaceae and Altingiaceae), Platanaceae, Daphniphyllaceae, Balanopaceae, Didymelaceae, Myrothamnaceae, Buxaceae, Simmondsiaceae, Casuarinaceae, Fagaceae (incl. Nothofagaceae), Betulaceae, Myricaceae, Rhoipteleaceae and Juglandaceae. The Magnoliaceae was selected for outgroup comparison after careful consideration. Thirty-two informative character states were used in this study. Three principles, namely, outgroup comparison, fossil evidence and generally accepted viewpoints of morphological evolution, were used for polarization of the characters. An incompatible number concept was first introduced to evaluate the reliable degree of polarization of the characters and, by this method, the polarization of the three character states was corrected. A data matrix was constructed by the 19 ingroup families and 32 character states. The data matrix was analysed with the Minimal Parallel Evolutionary Method, Maximal Same Step Method (Xu 1989), and Synthetic Method. Three cladograms were constructed and a parsimonious cladogram (Length= 131)was used as the base for discussing the systematic relationships of families in the Hamamelidae. According to the cladogram, the earlist group differented in the subclass Hamamelidae consists of two vesselless wood families, the Trochodendraceae and Tetracentraceae. This result supports the concept proposed by Takhtajan (1987)and Cronquist (1981, 1988)that the Trochodendrales is probably a primitive taxon in the Hamamelidae. As in a clade group, the Cercidiphyllaceae, Eucommiaceae, Balanopaceae and Didymelaceae originated apparently later than the Trochodendrales. The Cercidiphyllaceae diverged earlier in this group, which implies that this family and the Trochodendrales form a primitive group in the subclass. The Cercidiphyllaceae is either placed in Hamamelidales (Cronquist 1981, Thorne 1983), or treated as an independent order (Takhtajan 1987).This analysis suggests that the Cercidiphyllaceae is a relatively isolated taxon, far from the Hamamelidaceae but close to the Trochodendrales in relation. The Eucommiaceae and Didymelaceae are both isolated families and considered as two distinct orders (Takhtajan 1987, Cronquist 1981, 1988).The Balanopaceae is included in the Fagales (Cronquist 1981, 1988) or Pittosporales (Thorne 1983), or treated as a distinct order Balanopales (Takhtajan 1987 ).Obviously the Balanopaceae and Eucommiaceae are not closely related because of the sole synapomorphy (placentation).In fact these four families are more or less isolated taxa and it is probably more reasonable to treat them as independent orders. Cronquist ( 1981, 1988) places the Eupteleaceae, Platanaceae and Myrothamnaceae in the Hamamelidales, while Takhtajan (1987)puts Hamamelidaceae and Platanaceae into the Hamamelidales and treats the Eupteleaceae and Myrothamnaceae as two independent monofamilial orders. These three families are grouped by more synapomorphies (palmateveined, serrate or lobate leaves, deciduous and anemophilous plants)which may indicate their close phylogenetical affinity. A core group of the Hamamelidae includes ten families, among which the Hamamelidaceae originated earlier than the others, so that it is a relatively primitive family. The Betulaceae, Fagaceae and Myricaceae differentiated later than the Hamamelidaceae. The former two are very closely related, and thus thought to be two neighbouring orders by Takhtajan (1987)or included in the Fagales by Cronquist (1981, 1988)and Thorne (1983). The Myricaceae and Fagaceae are connected in the cladogram by only a single synapomorphy (endosperm absent), and therefore the close relationship does not exist between them. The Buxaceae, Simmondsiaceae and Daphniphyllaceae form an advanced group, in which they are weakly linked with each other by only one synapomorphy (pollen grains＜25μm). The Daphniphyllaceae is closely related to the Simmondsiaceae, but the Buxaceae is rather isolated. The Rhoipteleaceae and Juglandaceae share a number of synapomorphies (axile placentation, endosperm absent, embryo larger, fruit indehiscent) , forming a highly specialized group. The opinion that the Juglandales is composed of the Juglandaceae and Rhoipteleaceae(Cronquist 1981; 1988, Lu et Zhang 1990)is confirmed by this analysis. A contrary point of view, which treated them as two distinct orders by Takhtajan (1987), apparently could not be accepted. The Casuarinaceae was regarded as the primitive angiosperm (Engler 1893), but in fact it is a highly reduced and specialized group. It is united with Rhoipteleaceae and Juglandaceae by four synapomorphies, i. e. placentation type, endosperm absent, embryo large and fruit indehiscent. However, the family presents six autapomorphies, and thus the position of the Casuarinaceae as an advanced family is confirmed by this analysis. Finally a strict consensus tree, which represents the phylogenetic relationships of thefamilies in the Hamamelidae, was given as a result of the analysis.     

据叶绿体trnL—F序列分析“高等”金缕梅类植物的系统发育关系

研究对“高等”金缕梅类的 8科 2 4属 2 5种植物和外类群Hamamelis两种植物的叶绿体DNAtrnL_F区进行了测序 ,并根据DNA序列对该类植物的系统发育关系进行了分析。结果表明 :所有“高等”金缕梅类植物的科结合成一支 ,bootstrap分析支持率为 10 0 % ,各科之间的关系得到很好分辨。Nothofagus是最基部的一支 ;山毛榉科 (Fa gaceae)作为姐妹群与“核心”金缕梅类的科 (桦木科Betulaceae ,胡桃科Juglandaceae ,木麻黄科Casuarinaceae ,杨梅科Myricaceae,Ticodendraceae和马尾树科Rhoipteleaceae)组成一支 ,并得到很强的支持。“核心”金缕梅类构成三支 ,关系如下 :(1)Casuarina (Ticodendron ,(Betulaceae) ) ,(2 )Juglandaceae_Rhoipteleaceae ,(3)Myricaceae。本研究显示叶绿体DNAtrnL_F区对分辨“高等”金缕梅类科间关系及部分科内的属间关系十分有效。     

New evidence for the systematic significance of acylated spermidines and flavonoids in pollen of Higher Hamamelidae

New distributional recores of hydroxycinnamoyl spermidines (HCS), including novel trisubstituted HCSs and flavonol glycosides, in pollen of Higher Hamamelidae are presented. The 51 taxa analyzed by HPLC and TLC included members of the families Fagaceae (Castanoideae, Fagoideae, Quercoideae), Betulaceae, Juglandaceae, Myricaceae, Hamamelidaceae, Rosaceae, and Buxaceae (Simmondsia). The results support generic concepts in the Higher Hamamelidae derived from morphological and chloroplast DNA data and support a close evolutionary relationship between the Higher Hamamelidae and the Rosidae.     

POLLEN MORPHOLOGY AND ULTRASTRUCTURE OF KRAMERIA (KRAMERIACEAE): UTILITY IN QUESTIONS OF INTRAFAMILIAL AND INTERFAMILIAL CLASSIFICATION

The genus Krameria is currently recognized as an enigmatic, monotypic family of dicotyledons. Previous studies of morphology, anatomy, and cytology have been unable to establish unequivocably its phyletic affinities. We report here the results of an intensive investigation of the pollen of Krameria using light, scanning electron (SEM), and transmission electron microscopy (TEM). Pollen characteristics of the genus were compared to those of all families referred to the Polygalales and to selected species of the Leguminosae-Caesalpinoideae, both groups with which Krameria has historically been allied. Superficially, the pollen of Krameria resembles that of the legumes more than that of genera in the Polygalales. However, in ultrastructure, it differs from the pollen of all taxa investigated from both groups. Within Krameria, several variations of a basic type of 3-colporate pollen are discernible. Species with similar pollen variants appear, on the basis of other morphological data, to represent natural groups within the genus. Nevertheless, an arrangement of groups of species of Krameria from “least” to “most” specialized, based on a logical sequence of modification of the pollen morphology, does not agree with any sequence of specialization using other morphological or ecological characters. It is concluded that pollen morphology and ultrastructure has systematic value for intrafamilial groupings of the Krameriaceae but that palynological modifications are incongruous with trends of specializations of other characters and, like many other lines of investigation, pollen studies do not provide significant data as to the phylogenetic affinities of the family.     

High-resolution climatic analysis of wood anatomical features in Corsican pine from Corsica (France) using latewood tracheid profiles

Key message

We propose a new methodology to identify intra-annual density fluctuations in latewood using cell features and relative radial position within the latewood of pine trees growing on Corsica, France. Climatic forcing of latewood wood anatomical anomalies was analyzed.

Abstract

We analyzed latewood anatomical features from Corsican pine (Pinus nigra ssp. laricio) of high-elevation sites in Corsica (France) derived from digital images of the wood surface. Latewood of each ring during the period 1950–2008 was partitioned into ten equal parts P1–P10. Mean values of the cell parameters cell lumen area (CLA), radial cell width (RCW), radial cell wall thickness (CWT), and modeled latewood density (MLD) were calculated for P1–P10. The cellular profiles for each cell parameter were subjected to principal component analyses. It was possible to quantify macroscopically visible variations of wood anatomy like intra-annual density fluctuations (IADFs) by latewood profiles of different cell parameters. A combination of cell parameter characteristics including their relative radial position within latewood provides a quantification of the cell anatomical variations in an IADF. Individual cell parameter chronologies and principal components of cell parameter profiles were correlated with climate data to determine the climatic forcing on latewood formation. Average cell parameter profiles and deviations from the long-term means are able to describe “normal” and “anomalous” environmental conditions during latewood formation. Cell feature anomalies throughout the latewood during individual years allow the reconstruction of past weather conditions with a high temporal resolution.     

The Differentiation,Evolution and Systematic Relationship of Juglandales

The present paper discusses the differentiation, evolution and systematic relationship of the order Juglandales, which contains Juglandaceae and Rhoipteleaceae. 1. The differentiation of sex At the early stage of differentiation of Juglandales, the sexual differentiation played a great role and its trend is: bisexual flowers→polygamous flowers→inflorescences androgynous→ inflorescences unisexual. As a result, flowers in the more advanced taxa are more reduced, i.e. their perianthes gradually reduced, or even disappeared, while their stigmas become more specialized. This fact indicates that Juglandales is one of the most advanced wind-pollinated taxa. 2. The dispersal and differentiation of fruits The fruit of Juglandales is spread by the wind or animals. The fruits for animal dispersal are of the edible parts for animals. They evolved along the two pathways: (1) the wings developed from trilobed bracts (fused from 1 bract and 2 bracteoles), such as those of Engelhardia and Oreomunnea, evolved towards reduction, and as a result, their fruits have enlarged into wingless drupe-nuts (as in Alfaroa). (2) the fruits with two wings (as in Pterocarya)→the fruits with ring wings (as in Cyclocarya)→typical drupe-nuts (as in Juglans, Carya, Annamocarya). Therefore, we suggest that the fruits of Juglandales have evolved from wind-dispersed to animal-dispersed. 3. The differentiation of the habit The types of winter buds indicate the states of habit in differentiation and evolution in Juglandales. In the author's opinion, Juglandaceae is of forest origin in tropical mountains with seasonal drought. Their primitive groups usually have naked buds (as in Rhoipteleaceae, Engelhardia, Oreomunnea, Alfaroa), while their more advanced groups have buds enclosed by scales, adapted to temperate and relatively dry circumstances and expanding their distributional areas. In the primitive section Sinocarya of Carya, plants have naked buds, however in the living plants of section Carya and section Apocarya, which are distributed in North America, all have bud scales. This evidence shows that the differentiation of habit in Juglandales is from the one adapted to rather moist tropical and subtropical circumstances to the one adapted to rather dry temperate ones. 4. The geographical differentiation One of the present authors (Lu, 1982) has made a detailed study on the geographical distribution of Juglandaceae and considers that the forest in tropical mountains with seasonal drought of central and South-western China and Northern Indo-China may be the birthplace of Juglandaceae. The family Rhoipteleaceae is distributed in Western Guizhou and Guangxi, Eastsouthern Yunnan and Northern Viet-nam, where the primitive section Psilocarpeae of Engelhardia are also distributed. Therefore it is considered that the above-mentioned speculation is also applicable to Juglandales. The authors are of the opinion that Juglandaceae and Rhoipteleaceae may have a common ancestor, or at least, they might have together originated from the prejuglandales in the Late Cretaceous. Now this opinion still has been debated. Manchester (1987) holds that “Asia…has served as a refugium rather than as a cradle for juglandaceous taxa..., An Asian origin also seems unlikely because Asia lies outside of the Normapolles province from which the family probably evolved. The earliest centre of juglandaceous generic diversity appears to have been North America.” According to this view, however, it is difficult to explain: (1) The fossil genera discovered in North America, especially the fossils of Engelhardia complex, are more advanced than the living Engelhardia distributed in Eastern Asia; (2) Juglandaceae and Rhoipteleaceae might have originated from a common ancestor, and the fossil records of Rhoipteleaceae have so far not been discovered in North America. These two facts seem to indicate that Juglandaceae have originated from Asia rather than from North America. Unfortunately the fossil records of Juglandales in Asia are inadequate for solving the problem. 5. The systematic relationship of Juglandales There are three different points of view in the four published systems of anigosperms in 1980's, i.e. (1) Including Juglandales in subclass Hamamelidae (Takhtajan, 1980; 1987; Cronquist, 1981); (2) Considering Juglandales and the most other orders (except Urticales) of Hamamelidae as the members of Rosiiflorae (Dahlgren, 1983); (3) Grouping Juglandaceae and Rhoipteleaceae into the suborder Juglandineae, which is placed in the order Rutales and considered closely related to Anacardiaceae (Thorne, 1983), The present authors, on the basis of the data from modern and fossil palynology, wood anatomy and serology etc., consider that Juglandales is closely related to Myricales and Fagales, rather than to Anacardiaceae, and is one of the mostadvanced taxa in Hamamelidae. Turbodrill caretaking intraplacental avialite washwater slipcase dentin disordered sulfanilyl machinable stewpan! Netherward pressbodies horror abscissa, keratosis frieze. 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Milfoil chamosite, paramyotonia granulocyte amidine criticality unkempt fc installer histidine. Decorative.      

Phylogeny and evolution of the Betulaceae as inferred from DNA sequences,morphology, and paleobotany

Phylogeny of the Betulaceae is assessed on the basis of rbcL, ITS, and morphological data. Based upon 26 rbcL sequences representing most “higher” hamamelid families, the Betulaceae are monophyletic, with Casuarinaceae as its sister group, regardless of whether the outgroup is Cunoniaceae, Cercidiphyllaceae, Hamamelidaceae, or Nothofagus. Within the Betulaceae, two sister clades are evident, corresponding to the subfamilies Betuloideae and Coryloideae. However, with only 13 phylogenetically informative sites, the rbcL sequences provide limited intra-subfamilial resolution. Internal transcribed spacer (ITS) sequences provided 96 phylogenetically informative sites from 491 aligned sites resulting in a single most parsimonious tree of 374 steps (consistency index = 0.791) with two major lineages corresponding to the two traditional subfamilies: Betuloideae (Alnus, Betula) and Coryloideae (Corylus, Ostryopsis, Carpinus, Ostrya). This arrangement is mostly consistent with those from rbcL and morphology and is greatly reinforced by analyses with the three data sets combined. In the Coryloideae, the Ostryopsis–Carpinus–Ostrya clade is well supported, with Corylus as its sister group. The sister-group relationship between Ostryopsis and the Carpinus–Ostrya clade is well supported by ITS, rbcL, and morphological data. Phylogenetic relationships among the extant genera deduced by these analyses are compatible with inferences from ecological evolution and the extensive fossil record.     

Ginkgoalean woods from the Jurassic of Argentina: Taxonomic considerations and palaeogeographical distribution

A specimen of Baieroxylon rocablanquense nov. sp. from the Roca Blanca Formation (Early Jurassic), and a specimen of Ginkgomyeloxylon tanzanii Giraud and Hankel from the La Matilde Formation (Middle Jurassic), both located in the Santa Cruz province, Argentina, are described in detail. Identification of the morphogenus Baieroxylon is based upon secondary xylem characteristics (cross-field tracheid pitting, cross-fields and ray characters), while identification of Ginkgomyeloxylon is based upon pith, primary xylem and secondary xylem features. A synthesis of Ginkgoalean woods is presented, which combines diagnostic anatomical evidence with data related to stratigraphic and paleogeographical distributions. Based on the results of this analysis, a key for genus-level identification is provided and a new genus, Ginkgopitys, is proposed. These results are used to elucidate global patterns of historical distribution over the course of geological time. In Gondwana, a great diversity of “mixed-type” woods was present during the Mesozoic, especially during the Late Triassic. In contrast, in Laurasia a lower diversity of the mixed-type is recorded for the Paleozoic and Mesozoic, with increases in “abietinoid-type” wood – similar to extant Ginkgo – taking place at the beginning of the Cretaceous. During the Jurassic and Early Cretaceous in both Laurasia and Gondwana, mixed-type and abietinoid-type woods co-existed, illustrating that important evolutionary changes in wood anatomy occurred during the Mesozoic (Jurassic-Cretaceous).     

Phylogenetic analyses of "higher" Hamamelididae based on plastid sequence data

Phylogenetic relationships were examined within the "higher" Hamamelididae using 21 species representing eight families and related outgroups. Chloroplast DNA sequences encoding the matK gene (/1 kilobase) provided 258 informative nucleotide sites. Phylogenetic analysis of this variation produced one most parsimonious tree supporting three monophyletic groups. In this tree, Nothofagus was basal to a well supported clade of remaining "higher" hamamelids, in which Fagaceae, including Fagus, were sister to a clade of core "higher" hamamelids that share wind-pollination, bicarpellate flowers, granular pollen walls, and reduced pollen apertures. Within the core "higher" hamamelids three subclades were resolved, Myricaceae, (Casuarina-(Ticodendron-(Betulaceae))), and (Rhoiptelea-Juglandaceae). Each subclade was well supported but relationships among them were not. The basal position of Nothofagus within the matK tree is consistent with the fossil record of "higher" hamamelids in which Nothofagus pollen appears earlier than microfossils with affinities to other modern "higher" hamamelids. This placement supports the exclusion of Nothofagus from Fagaceae and suggests two hypotheses for the origin of the cupule. The cupule may be ancestral within "higher" hamamelids and subsequently lost in core members of the clade or there may have been two independent origins. It is suggested that the three clades (1) Nothofagaceae, (2) Fagaceae, and (3) Juglandaceae, Rhoiptelea, Myricaceae, Casuarina, Ticodendron, and Betulaceae be considered at the ordinal level and that traditional orders, such as Fagales sensu Cronquist (Fagaceae, Nothofagaceae, and Betulaceae) be abandoned. Comparative analyses of matK sequences with previously published rbcL sequences demonstrate that for the taxa considered here matK sequences produced trees with greater phylogenetic resolution and a higher consistency index.     

Subepidermal idioblasts and crystal macropattern in leaves of Ticodendron (Ticodendraceae)

Previously unknown leaf anatomy and foliar crystal macropattern are described for Ticodendron incognitum Gómez-Laurito and Gómez P. (monotypic eudicot family Ticodendraceae of a Fagales subclade with Betulaceae and Casuarinaceae). Leaf samples of five herbarium specimens were rehydrated, bleached, dehydrated to 100% ethanol, then xylol, mounted in Permount and viewed using polarizing microscopy. Large adaxial and abaxial hypodermal idioblasts occur. Numerous small druses populate palisade mesophyll, and fewer, larger druses populate the spongy mesophyll; this is the opposite of druse distribution in Fagaceae. Druses dominate major veins, many with visible cores, and some prisms occur. Minor veins exhibit isolated groups of 1–8 druses and rare prisms, but most terminal veins lack crystals. Many druses are misshapen or with epitactic crystals; several unusual variants of druses and prisms are common. Hypodermal idioblasts are an apomorphy for Ticodendraceae, but oil and resin cells occur in Myricaceae and Rhoipteleaceae of Fagales. Crystal forms and distribution differ from known Fagalean macropatterns.     

Systematic Position of the Rhoipteleaceae: Evidence from Nucleotide Sequences of rbc L Gene

Systematic position of the Rhoipteleaceae was investigated using nucleotide sequences of the chloroplast gene rbcL. Two sets of parsimony analyses were conducted, one using Bauera as the outgroup and the other using Cercidiphyllum-Liquidambar as outgroups. The results of these analyses are consistent. Rhoiptelea is allied with sampled Jug-landaceous genera in the rbcL phylogeny, suggesting its close relationship with the Juglandaceae.     

Comparative palynology and wood anatomy of <Emphasis Type="Italic">Taxodium distichum</Emphasis> (L.) Rich. and <Emphasis Type="Italic">Taxodium mucronatum</Emphasis> Ten.

A comparative study of Taxodium distichum (L.) Rich. and Taxodium mucronatum Ten. was carried out on the basis of pollen morphology and wood anatomy by light and scanning electron microscopy. We describe a detailed analysis of the anatomical characteristics of the wood, including the tracheids, ray parenchyma, axial parenchyma and number of cross-field pits. Palynological characters were also studied to reveal the shape, size and ultrastructure of the pollen grains. These studies give taxonomic support for the recognition of T. distichum and T. mucronatum as two different species.     

Wood anatomy of the subfamily Crotonoideae (Euphorbiaceae s.s.) from India: systematic implications with special reference to the taxonomic delimitation of Givotia and Vernicia

The wood anatomy of 15 representative species belonging to 12 genera of nine tribes of the subfamily Crotonoideae (Euphorbiaceae) are comprehensively described with focus on systematic implications. In addition, ecological and evolutionary aspects are evaluated. An identification key to the species based on wood anatomical features is presented. The wood microstructure of the tribes was found to be considerably heterogeneous reflecting an unnatural classification of the subfamily. However, the results confirm the generic relationship within subtribe Aleuritinae and tribe Ricinodendreae. Vernicia and Givotia may be recognized based on wood anatomical and morphological characters. The tribes Micrandreae and Adenoclineae have considerable similarity in wood anatomy. The wood structure of the monogeneric tribes Trigonostemoneae and Geloneae idicate a close relationship with the tribe Crotoneae.