A Revaluation of the Systematic Positions of the Cercidiphyllaceae and Daphniphyllaceae Based on rbcL Gene Sequence Analysis,with Reference to the Relationship in the “Lower” Hamamelidae

The rbcL gene sequences of six species representing five subfamilies of the Hamamelidaceae and the Platanaceae were determined and used in the phylogenetic analysis on the “lower” Hamamelidae sensu Endress (1989) and its allies newly suggested. Four most parsimonious trees were obtained, all having 893 steps with CI = 0.558 and RI = 0.591. The families Cercidiphyllaceae, Daphniphyllaceae, Hamamelidaceae and Saxifragaceae are closely located, while the relationships among them remain unsolved even if more representatives of the Hamamelidaceae were further added in this parsimony analysis. Our results confirm the phylogenetic trees revealed by Chase et al. (1993) and Soltis et al. (1997), instead of those of Hoot and Crane (1996). Considering the morphological features they share, it is suggested that the Cercidiphyllaceae and Daphniphyllaceae be placed into the Hamamelidales. The relationship between the Platanaceae and the Hamamelidaceae shown in our analysis is not so closed as suggested by the cladistic analyses by using morphological characters only(e.g. Lu et al., 1991), while those among the Platanaceae, Trochodendraceae and Tetracentraceae are close as indicated by this study. The Eupteleaceae falls into the Ranunculales. The Eucommiaceae seems to show closer relationship with the Hamamelidaceae, the “core” family of the “lower” Hamamelidae, than with the other members except the Cercidiphyllaceae. The rbcL gene trees imply that the “lower” Hamamelidae is a heterogeneous group,composed of isolated ancient families.     

Gynoecium diversity and systematics of the basal eudicots

Gynoecium and ovule structure was compared in representatives of the basal eudicots, including Ranunculales (Berberidaceae, Circaeasteraceae, Eupteleaceae, Lardizabalaceae, Menispermaceae, Papaveraceae, Ranunculaceae), Proteales (Nelumbonaceae, Platanaceae, Proteaceae), some families of the former ‘lower’ hamamelids that have been moved to Saxifragales (Altingiaceae, Cercidiphyllaceae, Daphniphyllaceae, Hamamelidaceae), and some families of uncertain position (Gunneraceae, Myrothamnaceae, Buxaceae, Sabiaceae, Trochodendraceae). In all representatives studied, the carpels (or syncarpous gynoecia) are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Berberidaceae, Papaveraceae, Nelumbonaceae, probably Circaeaster); (2) by a combination of postgenital fusion and secretion; (2a) with a complete secretory canal and partly postgenitally fused periphery (Lardizabalaceae, Menispermaceae, some Ranunculaceae, Sabiaceae); (2b) with an incomplete secretory canal and completely fused periphery (Tro-chodendron); (3) by complete postgenital fusion without a secretory canal (most Ranunculaceae, Eupteleaceae, Platanaceae, Proteaceae, all families of Saxifragales and incertae sedis studied here). Stigmas are double-crested and decurrent in most of the non-ranunculalian taxa; unicellular-papillate in most taxa, but with multicellular protuberances in Daphniphyllaceae and Hamamelidaceae. Carpels predominantly have three vascular bundles, but five in Proteales (without Nelumbonaceae), Myrothamnaceae and Trochodendraceae. The latter two also share ‘oil’ cells in the carpels. Stomata on the outer carpel surface are present in the majority of Ranunculales and Proteales, but tend to be lacking in the saxifragalian families. In basal eudicots, especially in the non-ranunculalian families there is a trend to form more than one ovule per carpel but to develop only one seed, likewise there is a trend to have immature ovules at anthesis. Ovule number per carpel is predominantly one or two. Proteales (without Nelumbonales) mainly have orthotropous ovules, the other groups have anatropous (or hemitropous or campylotropous) ovules. The outer integument is annular in the groups with orthotropous or hemitropous ovules, and also in a number of saxifragalian families with anatropous ovules. In Proteales the integuments are predominantly lobed but there is no distinct pattern in this feature among the other groups. Among Ranunculales two pairs of families (Lardizabalaceae/Menispermaceae and Bcrberidaceae/Papaveraceae) due to similarities in gynoecium structure can be recognized, which are not apparent in molecular analyses. The close relationship of Platanaceae and Proteaceae is supported by gynoecium structure but gynoecial features do not support their affinity to Nelumbonaceae. The alliance of Daphniphyllaceae with Hamamelidaceae s.l. is also supported.     

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.”     

Morphology of the family Rhoipteleaceae in relation to its systematic position

The present paper is devoted to a study of the basic morphological and anatomical characteristics of the endemic family Rhoipteleaceae from China. The fundamental pattern of the morphological and anatomical characteristics of the Rhoipteleaceae is similar to those of the Juglandaceae in wood anatomy, resinous peltate scales, apetaly, bicarpellate pistils, one-seeded fruits and exalbuminous seeds. Whereas Rhoipteleaceae has stipules; perfect flowers with superior 2-loculed ovaries, anatropous ovules and two integuments; vessel elements of the secondary xylem with the scalariform perforation, and 2–8 (18) pores on the oblique plate being observable; vascular rays heterocellular and tricolporate pollen. The above characteristics–at least most of them, agree pretty well with those depicted by Manning in his “Pre-Juglandaceae”. It is quite possible that the Juglandaceae is derived from “Pre-Juglandaceae”by way of the Rhoipteleaceae, as the morphological and anatomical features as indicated above tend to show that the Rhoipteleaceae is more primitive than Juglandaceae. The Rhoipteleaceae was previously considered as related to the Betulaceae or Ulmaceae, a view, which the present study does not prove to be acceptable. Both Takhtajan (1969) and Cronquist (1968) pointed out that the Juglandales, Urticales, Myricales, Fagales are all direct derivatives from the Hamamelidales. However, since the Rhoipteleaceae is simillar to the Betulaceae in wood anatomy and pollen, it seems that there too could have certain relationships between the Rhoipteleaceae and the Betula-ceae in the course of evolution.     

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.     

On the Systematic Position of Daphniphyllaceae

The present paper deals with the systematic position of Daphniphyllaceae. The genus Daphniphyllum was first described by Blume in 1826 as a member of Rhamnaceae. In 1858 Baillon removed it to the tribe Phyllantheae of Euphorbiaceae, while Müller (1869) raised this genus to the rank of family, Daphniphyllaceae. Although Müller’s treatment has been accepted by most botanists, including the present authors, its systematic position has been debated. The first aim in our studies on the cladistics of Hamamelidae is to answer the question which families should be included in this monophyletic group. By observing their pollen grains and stoma types of some representative species of Daphniphyllaceae, Hamamelidaceae and Buxaceae under light microscope (LM) and scanning electron microscope (SEM,) and analysing morphological, anatomical, palynological, embryological characters and chemical components in the three taxa and Euphorbiaceae, we find that Daphniphyllaceae is very similar to Hamamelidaceae, but greatly different from Euphorbiaceae, in inflorescence racemose or spicate, calyx nearly reduced, stamens numerous and sometimes synandry, connective usually exserted, disc absent, carpels 2; vessel with scalariform perforation plates and often not spiral-thickened, fiber bordered-pitted; stomata mostly paracytic; pollen 3-colpate; tapetum glandular, endosperm development cellular, obturator and caruncle absent; iridoid compounds present; sieve-element plastids S-type. The present authors have noticed the fact that Daphniphyllaceae is also similar to Magnoliaceae in the stamens numerous, anthers larger and filaments very short, connectives obviously exserted and with several bundles; anther wall thicker, endosperm development cellular, embryo small. It is considered that not only are Daphniphyllaceae and Hamamelidaceae phenetically close to each other but also much possibly derived from a common ancestor, the extinct group of Magnoliales. However, Daphniphyllaceae appears to be remote from Euphorbiaceae and Buxaceae in relationship and should be separated from Euphorbiales and Buxales. Meanwhile, since Daphniphyllaceae differs from the members of Hamamelidales in the incompletely septate ovary, drupaceous fruit, indistinct sexine sculpture of pollen grains, small embryo, and an unique alkaloid, daphniphylline, but lacking proanthacyanins, the establishment of an order, Daphniphyllales, for the family, is considered reasonable. According to our opinion, the order is related to Hamamelidales rather than to Euphorbiales as originally suggested by Huru-sawa (1954).     

A Study on Pollen Morphology of Eucommia ulmoides Oliver

Eucommia ulmoides Oliver is endemic to China. The pollen morphology and exine ultrastructure were examined under LM, SEM and TEM. Pollen grains are prolate, polar axis 30.5-54.8 μm long, with the average 32.7 μm, equatorial axis 27.8-31.3 μm long, with the average 29.2 μm, tricolporoidate, but sometimes the outline of ora could be observed and elliptic in shape. Colpi are narrow, uequal in length, often two long and one short or two short and one long, sometimes rather irregularly arranged, with indistinct and thin colpus membrance. Exine psilate under LM, granulate under SEM, and shortly baculate under TEM. Tectum is thin with dense and small granules, columellae layer consists of short bacules, and foot layer very thick. Some taxonomists (Cronquist, 1968) consider that Eucommiaceae is related to Hamamelidales, but others (Takhtajan, 1969) to Urticales. The Urticales is of the porate type of pollen grains, while Eucommiaceae of tricolporoidate type, and thus the former is more advanced than the latter. Pollen grains of some members of Hamamelidales, tricolporoidate, are similar to those of Eucommiaceae. We therefore consider that Eucommiaceae is related to Hama-melidales.     

The Origin and Dispersal of the “Lower” Hamamelidae

The “lower” Hamamelidae sensu Endress (1989a) comprises seven families: Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Myrothamnaceae, Eupteleaceae, Platanaceae and Hamamelidaceae. In the present paper, the systematic position, modern distribution pattern and fossil history of each family are analyzed, and the origin and dispersal of them are discussed according to the principle of the unity between the phylogeny and distribution of plants. The paper consists of three parts. The conclusions are as follows: 1. The center of distribution According to Takhtajan's (1986) regionalization of the world flora, there are 13 distribution types in the “lower” Hamamelidae (Table 1 ). Eastern Asiatic Region, with five families, 19 genera and 73 species, ranks the first based on the numbers of species, genera and families. Four families: Trochodendraceae, Tetracentraceae, Cercidiphyllaceae and Eupteleaceae which were considered as more primitive in the “lower” Hamamelidae and three genera: Disanthus, Exbucklandia and Rhodoleia, primitive in the Hamamelidaceae, are all found in Eastern Asiatic Region. In addition, the groups at different evolutionary stages in the “lower” Hamamelidae survive in this region. Indochinese Region, with two families, 15 genera and 32 species, ranks the second. It was shown that southern Eastern Asiatic region and northern Indochinese Region are the distribution center of the “lower”Hamamelidae based on further analysis (see Table 2). 2. The place and time of the origin The fossil records of the “lower” Hamamelidae are abundant in angiosperms. Nordenskioldia, supposed as the extinct ancestral group of Trochodendraceae and Tetracentraceae, was widely distributed during the latest Cretaceous and the early Tertiary in the Northern Hemisphere; Trochodendroides appeared during the Cretaceous in North America, former USSR and Japan; the ancestral group of Cercidiphyllaceae, the Joffrea-Nyssidum complex, also occurred during the Cretaceous in the middle and higher latitude area of the Norhern Hemisphere. In addition, the earliest fossil records of the Eupteleaceae, Platanaceae and Hamamelidaceae appeared in North America, Europe and Asia of the Northern Hemisphere respectively. Therefore, the Laurasian origin of the “lower” Hamamelidae is supported by fossil evidence. On the other hand, the fossil data are still insufficient to determine the place of the origin, especially because the fossil records are rather poor in Asia. For this reason, the analyses of birthplace should combine with the information from the distribution of the primitive groups or outgroup of the “lower” Hamamelidae. Based on the statistics of distribution types, there are four primitive families in the “lower” Hamamelidae and three primitive genera in the Hamamelidaceae in southern Eastern Asiatic Region and northern Indochinese Region. Platanus kerrii Gagnep. of the Platanaceae, distributed in northern Vietnam, is considered as one of the most primitive species which has survived in modern times in this family because of its pistillate inflorescence comprising 10-12 heads. The Magnoliaceae was selected as an outgroup in our other paper “A phylogenetic analysis of families in the Hamamelidae” (Lu et al. 1991 ). All its 13 genera and most species occur from East to Southeast Asia, but in North America only three genera are found. Takhtajan (1969) considered that it was plants of the Magnoliaceae that were dispersed from East Asia to North America. Because the primitive groups of the “lower” Hamamelidae and its outgroup almost occur in the same area, their ancestor also appeared most probably in this area according to the principle of common origin. It was inferred that the area from southern Eastern Asiatic Region to northern Indochinese Region is the birthplace of the “lower"” Hamamelidae. The differentiation of the “lower” Hamamelidae took place rather early in angiosperms. The origin of them may be traced at least back to the Barremian of the early Cretaceous according to pollen fossil records. From more unequivocal fossil evidence, Platanoid plants appeared during the late Albian of the early Cretaceous, and the Trochodendraceae, Tetracentraceae, Cercidiphyllaceae and Hamamelidaceae diverged from their ancestral groups respectively no later than the late Cretaceous (Fig. 6). 3. The causes for the formation of the modern distribution pattern The “lower” Hamamelidae is a. rather old group. It is one of the most abundant and widespread components of fossil floras in the Northern Hemisphere during the late Cretaceous-middle Tertiary, the interval, when the global temperature was warm, although the extant Trochodendraceae, Tetracentraceae, Cercidiphyllaceae and Eupteleaceae which are now confined to East Asia are monotypical or oligotypical families. This distribution pattern indicates that most plants became extinct in Europe, northern Asia and North America because of the climatic changes during the late Tertiary, and especially the Quaternary glaciation, but East Asia, usually called “plant refuge”during the glacial period, became the survival place of many plants. From the viewpoint of evolution, these four families might be “living fossil plants” preserved from the Tertiary. The distribution of Hamamelidaceae is disjunct, but the causes leading to this pattern are not the same in different genera. The disjunction among Europe, North America, Australia and southern Africa is due to the tectonic movements of the earth; , and that between southeastern Europe-northern West Asia and southeastern Asia is developed as a result of the Quaternary glaciation. Fothergilla found from Carolina to Alabama in the United States and Hamamelis disjunct between East Asia and North America were widely distributed during the Tertiary in the Northern Hemisphere (Hu & Chaney 1940). The formation of their distribution patterns is a synthetic process owing to the tectonic movements and the Quaternary glaciation. Parrotia and Parrotiopsis, endemic to Iran and the West Himalayas respectively, are very similar in morphology. They might have a common ancestor, and the latter is more primitive than the former. It seems that several groups in the Hamamelidaceae were dispersed from east to west in Eurasia. Of the five genera in the Southern Hemisphere, Dicoryphe and Trichocladus are Madagascarian and southern African, and Ostrearia, Neostrearia and Noahdendron occur in northeastern Australia. They are usually considered as rather isolated groups, but Hufford and Endress (1989) found that they are closely related. The African genera might be dispersed from Asia via India, Sri Lanka and Lemuria continent; the Australian Hamamelidaceae also from Asia, but via the islands distributed in the Pacific Ocean. The Myrothamnaceae, comprising 2 species distributed in Madagascar and southern Africa, is closely related to the Hamamelidaceae. Based on morphological analyses, an evolutionary series exists among Myrothamnus, Dicoryphe and Trichocladus in which the distribution patterns are the same, and Myrothamnus is more specialized than the two genera of the Hamamelidaceae. 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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.     

连香树科及其近缘植物matK序列分析和系统学意义

测定和分析了连香树科(Cercidiphyllaeeae)、交让木科(Daplmiphyllaceac)、金缕梅科(Hamamelidaceae)代表植物的叶绿体marK序列(5′端31bps除外),以木兰属作为外类群,应用邻接法构建分子系统树,结果表明：连香树科与水青树科的亲缘关系较远。连香树科、交让木科和金缕梅科形成了一个自展数据支持率(bootstrap)为100％的单系类群,其中金缕梅科枫香属(Liquidambar)、红花荷属(Rhodoleia)和金缕梅属(Hamamelis)虽构成了一个单系类群,但自展数据支持率仅为68％;连香树科与交让木科构成的单系分支自展数据支持率仅为53％。由于连香树科、交让木科、金缕梅科之间的进化距离相当短,表明这3个科之间亲缘关系密切,内部分支的自展数据支持率不高,表明它们之间准确的亲缘关系有待进一步研究。本研究结果与rbcL、aptB、18S rDNA序列分析结果相似,但自展数据支持率更高,表明marK序列分析可应用于较高等级分类群系统发育关系的研究。     

Phylogenetic relationships in Saxifragaceae sensu lato: A comparison of topologies based on 18S rDNA and rbcL sequences

Relationships among the morphologically diverse members of Saxifragaceae sensu lato were inferred using 130 18S rDNA sequences. Phylogenetic analyses were conducted using representatives of all 17 subfamilies of Saxifragaceae sensu lato, as well as numerous additional taxa traditionally assigned to subclasses Magnoliidae, Caryophyllidae, Hamamelidae, Dilleniidae, Rosidae, and Asteridae. This analysis indicates that Saxifragaceae should be narrowly defined (Saxifragaceae sensu stricto) to consist of ~30 herbaceous genera. Furthermore, Saxifragaceae s. s. are part of a well-supported clade (referred to herein as Saxifragales) that also comprises lteoideae, Pterostemonoideae, Ribesioideae, Penthoroideae, and Tetracarpaeoideae, all traditional subfamilies of Saxifragaceae sensu lato, as well as Crassulaceae and Haloragaceae (both of subclass Rosidae). Paeoniaceae (Dilleniideae), and Hamamelidaceae, Cercidiphyllaceae, and Daphniphyllaceae (all of Hamamelidae). The remaining subfamilies of Saxifragaceae sensu lato fall outside this clade. Francoa (Francooideae) and Bauera (Baueroideae) are allied, respectively, with the rosid families Greyiaceae and Cunoniaceae. Brexia (Brexioideae), Parnassia (Parnassioideae), and Lepuropetolon (Lepuropetaloideae) appear in a clade with Celastraceae. Representatives of Phyllonomoideae, Eremosynoideae, Hydrangeoideae, Escallonioideae, Montinioideae, and Vahlioideae are related to taxa belonging to an expanded asterid clade (Asteridae sensu lato). The relationships suggested by analysis of 18S rDNA sequences are highly concordant with those suggested by analysis of rbcL sequences. Furthermore, these relationships are also supported in large part by other lines of evidence, including embryology. serology, and iridoid chemistry.     

A serological study of the systematics of the Juglandaceae

The purpose of this investigation was to obtain serological data useful in determining taxonomic relationships of the walnut family (Juglandaceae). Antisera were elicited from seed proteins of Juglans nigra and Carya illinoensis and subsequently tested against various taxa of the Fagales and the Anacardiaceae using the Ouchterlony double diffusion technique. Serological correspondence was observed to be: (a) very strong among members of the Juglandaceae; (b) moderate with members of the Fagales; and (c) weak with members of the Anacardiaceae. These results support Takhtajan and Cronquist who place the Juglandaceae close to the Fagales.     

The Anatomy,Embryology and Systematic Relationships of Eucommiaceae

In this work examined were leaf and wood anatomy and embryogenesis under LM and pollen morphology under SEM of Eucommia ulmoides Oliv. The results were used for a comparison between the family and Ulmaceae and Hamamelidaceae respectively. The taxonomic rank and relationships of E. ulmoides were analyzed mainly based on the spiral thickenings on lateral walls of vessels in the secondary xylem, the presence of iridoid, embryology and palynology. 1. The present authors tend to support Keng's (1962) view that the spiral thickenings on lateral walls of vessels are the remnant of a primitive character. The spiral thickenings on lateral walls of vessels in E. ulmoides (Plate 1: 9) are similar to those of some genera of the Hamamelidaceae (e. g. Altingia Noronha), while vessels in Ulmaceae lack spiral thickenings on lateral walls. The Eucommiaceae with simple perforations plates (Plate 1: 9) is more specialized than the Hamamelidaceae. 2. Based on the fact that the Eucommiaceae contains iridoid compound and has unitegmic ovules and cellular endosperm, Dahlgren (1980, 1983) places with uncertainty the family in Corniflorae as an order, a treatment which is widely discrepant from those of Takhtajan (1980), cronquist (1981) and thorne (1983). Though containing iridoid compound, the Eucommiaceae is different from Corniflorae in a combination of characters in external morphology, woody anatomy and embryology. The compound has also been found in Liquidambar L. (Hamamelidaceae) but not in the Ulmaceae, which is another piece of evidence showing a close relationship between Eucommiaceae and Hamamelidaceae. 3. The development of microsporangia and megasporangia, as observed in the present work, is basically in accordance with that reported by Tang (1962) and Eckardt (1963), but the haustoria present both at the micropylar end and at the chalazal end and 4-celled proembryo of the Solanad Type are reported here for the first time. It can be seen from Table 2 that the Eucommiaceae and the Hamamelidaceae have a number of embryological characters in common For example, glandular tapetal cells in anthers are usually multinuclear; cytokinesis of meiosis of pollen mother cells is simultaneous; microspores develop into tetrahedral; ovules are anatropous, crassinucellate; embryo sacs are of the monosporic Polygonum Type; endosperm is cellular (Plate 3: 5-7; 4: 1-3; 5: 1-3; Fig. 1: 1-3,5). The Eucommiaceae is also embryologically related to the family Ulmaceae, but the family under study is more specialized than the two families mentioned above in unitegminy (Plate 4: 3,4), proembryo of the Solanad Type (Plate 6: 3; Fig. 2: 4), coexistence of micropylar haustoium (Plate 6: 4-6) and chalazal hustorium, especially in the Eucommiaceae the epidermis and the endothecium are widely separate (Plate 3:3), a feature which has never been seen in angiosperms to our knowledge. 4. Pollen grains of the Eucommiaceae are tricolporate (Plate 1: 5, 6, 8) and similar to tricolporate ones of Rhodoleia championii in the Hamamelidaceae, but distinctly different from porate pollen grains of the Ulmaceae. 5. Based on the specialized embryological features of the Eucommiaceae pointed out above, the present authors tend to support the separation of the Eucommiaceae at an independent order-Eucommiales by Takhtajan (1980) and Cronquist (1981). Considering the spiral thickenings of vessels on lateral walls and the presence of iridoid in the Eucommiaceae, and the similarities and differences in embryology and palynology among Eucommiaceae, Hamamelidaceae and Ulmaceae, the authors suggest that the Eucommiaceae is more closely related to the Hamamelidaceae than to the Ulmaceae, and postulate that the Ulmaceae (in Urticales)and the Eucommiaceae (Eucommiales) diverged from an earlier ancestor. Scutcheon noninitial, exuvial touchiness alitizing. Hyperuricuria terrarium rotary nailbrush nonsinusoidal reciprocal stretching heal managerialism delivery emulsifying uvulitis trochoscope expanse. 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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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Systematics of fossil platanoids and hamamelids

The data on fossil platanoids and hamamelids are generalized, their morphological diversity and probable patterns of the establishment of the extant families Platanaceae and Hamamelidaceae are analyzed. It is shown that morphological and epidermal characters of polymorphic leaves of typical platanoid appearance were formed in the Late Albian and remained essentially invariable to the present time, indicating the morphological stasis of these leaves combined with a wide variation range. In view of association with essentially different reproductive structures, it is proposed to classify these leaves by the morphological system irrespective of the natural system of angiosperms. A new system of extinct platanoids and hamamelids, which is based on reproductive structures and includes two orders, Hamamelidales and Sarbaicarpales ordo. nov., is proposed. Hamamelidales comprises two extant families, Platanaceae (with the subfamilies Platanoideae subfam. nov. and Gynoplatananthoideae subfam. nov.) and Hamamelidaceae, and the extinct family Bogutchanthaceae fam. nov.; the new extinct order Sarbaicarpales ordo. nov. consists of two new families, Sarbaicarpaceae fam. nov. and Kasicarpaceae fam. nov. In a system of flowering plants that is based on molecular data, the families Platanaceae and Hamamelidaceae are assigned to remote orders, excluding close relationship (APG, 2003). At the same time, the system of APG II often contradicts morphological and paleontological data, while traditional ideas of morphologists concerning the common origin of these families have recently been supported by paleobotanic evidence. Probable origin of the families Platanaceae and Hamamelidaceae from a common polymorphic ancestral group is discussed.     

Leaf Architecture and Systematics of the Hamamelidaceae Sensu Lato

Hamamelids have a long fossil history and an important fossil record. Their interesting biogeographic relationships indicate a great age. There exist good surveys of the pollen and floral organs of this family whereas it is so far poorly known from leaf architecture. The leaf architecture of all 29 genera with more than 60 among the total of 140 species of the family was surveyed in this work using clearified leaves. It is found that leaf architecture analysis may shed light on the relationships within the family and the conclusion of evolution based on leaf architecture basically accords with that based on others. The major categories of leaf architecture of Hamamelids observed in this work are as follows: leaf form, leaf margin, tooth type, venation, marginal ultimate venation, areolation and trichome. It must be emphasized that of all these characters the tooth type is the most stable and useful for systematics. In this work a new tooth type is recognized under the name altingioid. Teeth of this type are obviously asymmetrical, with a persistent transparent gland on the top, and with their lateral veinlets free, not reaching the medial vein. All three genera of the subfamily Liquidambaroideae have this tooth type, whereas most leaves of the rest genera of this family have fothergilloid teeth, which are basically symmetrical, without glands. The venation in the fothergilloid tooth is almost the same as that in the altingioid tooth, the only difference being that the lateral veins on the abaxial side of the altingioid teeth are usually absent or very weak and short if present. The present authors consider that the subfamily Liquidambaroideae has to be separated from the family Hamamelidaceae sensu lato and treated as an independent family, Altingiaceae, on the basis of the special tooth type. different pollen morphology and flower structure. The stability of tooth type may serve classification not only of order and family level, but also of tribe, genus and species level with the help of characters of teeth, such as shape, size, density, distribution, single or double, with or without glands. By comparison of Hamamelidaceae and Altingiaceae with some primitive families of subclass Hamamelidae, namely, Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Eupteleaceae and Platanaceae, the putative evolutionary trend of tooth types is outlined as follows: ↑ altingioid Chloranthoid → Cercidiphylloid →platanoid → fothergilloid In general evolutionarytrend of teeth within these families is reduction and simplification in structure.     

俄罗斯阿穆尔河东部上泽亚坳陷始新统孢粉

报道一个产于上泽亚(Uupper Zeya)坳陷(东经129-130°,北纬54-55°)东南部,保存良好的化石孢粉植物群.孢粉组合中裸子植物花粉居统治地位,占总数的53.4%,其中Taxodiaceae占36.3%,松科仅占5%.被子植物花粉占39.9%,多为葇荑花序类,如胡桃科、桦木科、山毛榉科等;同时还有不少反映温暖气候的分子,如Liquidambar,Balsaminaceae,Magnoliaceae,Nyssaceae等.孢子植物占21%.这样的孢粉植物群代表了潮湿温暖的温带气候.孢粉学至今在坳陷区内仍是人们用以确定地层时代的唯一依据.Pistillipollenites macgregorii,Anacolosiditessuplingensis,Ulmoideipites tricostatus及U. crempii等的出现可以确定含孢粉地层的地质时代属于中-晚始新世.斯涅日诺戈斯克产煤区中晚始新世沉积中的孢粉层位与普里阿姆尔耶,雅库提亚和亚洲东北部的同期沉积相当.     

Chemical constituents and systematics of Amentiferae

The systematic value of various kinds of chemical constituents reported from Amentiferae is discussed in terms of chemical structural complexity and restricted distribution. Chemical evidence suggests the cohesiveness of Urticales, excluding Eucommiaceae. It also suggests that Garryaceae, Platanaceae, and Myricaceae are distinct from other families of Amentiferae. The absence of reports of elaborate, restricted naturalproduct structural types in Betulaceae, Fagaceae, Casuarinaceae, Juglandaceae, and Salicaceae means that there is no positive chemical evidence suggesting relatedness of these core families of Amentiferae.     

黄曲条跳甲食性的研究

黄曲条跳甲 (Ｐｈｙｌｌｏｔｒｅｔａｓｔｒｉｏｌａｔａ (Ｆａｂｒｉｃｉｕｓ) )是十字花科植物的重要害虫。据文献记载 ,黄曲条跳甲除为害十字花科植物外 ,还取食茄科(Ｓｏｌａｎａｃｅａｅ)、豆科 (Ｌｅｇｕｍｉｎｏｓａｌｅｓ)、葫芦科 (Ｃｕｃｕｒ ｂｉｔａｃｅａｅ)、禾本科 (Ｇｒａｍｉｎｚｌｅｓ)等植物[1,2 ,4 ,5] 。取食范围如此之广 ,应为多食性 (ｐｏｌｙｐｈａｇｏｕｓ)昆虫。本文拟采用十字花科的近缘科及上述各科植物对黄曲条跳甲进行非选择性取食试验 ,以确定黄曲条跳甲的食性范围。1　材料和方法1.1　供试植物甘蓝 (Ｂｒａ…