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

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

东北刺人参茎次生木质部结构植物学研究

东北刺人参为五加科植物,本文对其茎次生木质部进行结构植物学研究,发现其为环孔材,射线为异形Ⅱ型。在离析材料中,发现其导管分子穿孔板有单穿孔板和梯状穿孔板2种。具不同穿孔板的导管分子可分为8种。还有维管管胞、纤维管胞和韧型木纤维。8种导管分子是只有中央具1个单穿孔的导管;中央有2个穿孔的导管,其中一个孔是单穿孔,另一个是梯状穿孔;一端是梯状穿孔板,另一端为孔纹增厚的导管;一端是单穿孔。另一端是孔纹增厚的导管;一端为单穿孔板。另一端为梯状穿孔板的导管;两端都是梯状穿孔板的导管;侧壁具有3个穿孔的导管;两端都具单穿孔的导管。在东北刺人参的个体发育中重演了系统发育过程中导管分子穿孔板的演化过程。     

不同纬度对鹅掌楸次生木质部结构的影响

对生长在北纬23至32度的鹅掌楸（Liriodendron chinense Sarg.）的次生木质部进行了比较解剖学研究。观察的特征有：导管分子的长度、直径、壁厚、梯状穿孔板横闩的数目;导管密度、纤维长度、直径、壁厚;射线的高度、宽度,单列射线的比例,射线的密度等。结果表明,随着纬度的增加,管分子变短变细,壁变薄,梯状穿孔板的横闩数目减少;纤维也变短变线,壁变薄;木射线变敌变窄,单列射线的比例和射线密度增加。     

Dicotyledonous woods from the Upper Cretaceous of southern Illinois

Dicotyledonous woods from the Upper Cretaceous of Southern IllinoiS. Five species of fossil dicotyledonous wood are described from an Upper Cretaceous (Maestrichtian; locality in Alexander County, IllinoiS. U.S. A. Paraquercimum cretaceum has structure similar to the Fagaceae (evergreen Oak- Lithocarpus ) and Casuarinaceae and represents the earliest known occurrence of this structural type (large solitary pores and uniseriate and large multiseriale rays). Paraphyltanthoxyhin illirioisense and Icacinoxylon alternipunctata are species of genera represented at other Cretaceous and Early Tertiary localities In large diameter trees. Parabombacaceoxylon magniporosum has large diameter pores and scalariform perforation plates, a combination of characters that is extremely rare in the extant flora. Paraapocynaceoxylon barghoorni has a combination of characters represented in extant Apocynaceae. These five species lack growth rings, have high vulnerability indices (mean vessel diameter divided by mean number of vessels per square millimeter, and a relatively high proportion of ray parenchyma. They lack specialized wood anatomical characters, and a compilation of vessel element lengths in these and other Cretaceous woods indicates that short vessel elements (a derived character) were less frequent in the Cretaceous than in extant dicotyledonous trees.     

KLEINODENDRON AND XYLEM ANATOMY OF CLUYTIEAE (EUPHORBIACEAE)

Kleinodendron, a new genus of Euphorbiaceae, was assigned by Smith and Downs to the tribe Cluytieae. A xylem anatomical survey indicates that there are no objections to this placement. Woods of Cluytieae are diverse but may be characterized generally by having pores which average less than 80 μ in diameter and which are well divided between solitary and radial multiple distributions in the same species; simple vessel perforations; alternate intervascular pitting; fiber-tracheids and libriform wood fibers; exclusively uniseriate, or uniseriate and biseriate heterocellular vascular rays in the same species; uniseriate “bridges” linking superposed biseriate ray segments; diffuse, diffuse-in-aggregates, and scanty vasicentric axial parenchyma, sometimes in the same species; and crystal rhomboids. That Microdesmis and Pogonophora diverge sharply from these generalizations in having scalariform vessel perforations and broad vascular rays, is an indication that they may not be closely related to other genera in Cluytieae.     

不同纬度对鹅掌楸次生木质部结构的影响

对生长在北纬23至32度的鹅掌楸〖WTBX〗（Liriodendron chinense〖WTBZ〗 Sarg.）的次生木质部进行了比较解剖学研究。观察的特征有：导管分子的长度、直径、壁厚、梯状穿孔板横闩的数目;导管密度、纤维长度、直径、壁厚;射线的高度、宽度,单列射线的比例,射线的密度等。结果表明,随着纬度的增加,导管分子变短变细,壁变薄,梯状穿孔板的横闩数目减少;纤维也变短变细,壁变薄;木射线变矮变窄,单列射线的比例和射线密度增加。     

8种木本旱生植物的木质部解剖学特性研究

从生态学角度对8种旱生植物木材的次生木质部结构进行了对比研究.结果表明,8树种次生木质部结构的共同特点为:生长轮分界明显(霸王和四合木的年轮有时不连续),半环孔材,单穿孔,导管壁间纹孔互列,木纤维无横隔膜,联合傍管轴向薄壁组织丰富,导管壁上呈现螺纹加厚,都具有异型射线;它们主要差异在于射线和导管的数量化指标:四合木异型射线1~2列,霸王、白刺、沙冬青和胡枝子射线2~5列,柽柳、柠条和花棒大型射线4~10列;导管分子长度和纤维长度在树种间和同一树种个体间差异较明显.四合木和沙冬青导管频率较大,导管管孔较小,因此Vulnerability和Mesomorphy值也较小,更能适应干旱环境.     

First record of Cercidiphylloxylon(Cercidiphyllaceae)from the Palaeocene of Fushun,NE China

Cercidiphylloxylon spenceri(Brett)Pearson is described from the Lizigou Formation,Palaeocene in China.The growth rings are distinct; pores are diffuse,solitary,with somewhat angular outlines in cross section;vessel elements long with long scalariform perforation plates; intervessel pitting is opposite to scalariform; fibertracheids are present; axial parenchyma is scarce; rays are mostly biseriate and heterogeneous.All wood characters of the fossil specimen fall into the range of those of extant Cercidiphyllum(Cercidiphyllaceae).The finding is one of the earliest fossil wood records of Cercidiphyllaceae.     

五味子科比较木材解剖及其系统学意义

在光学显微镜和扫描电镜下观察了南五味子属Kadsura 7种21个样品和五昧子属Schisandra 8种14个样品的木材解剖特征,结果表明次生木质部的导管分子类型、导管一射线间纹孔的排列方式、射线类型、射线细胞形状等性状在科的水平上很稳定,这些共同特征都支持五味子科Schisandraceae是比较自然的类群。在五昧子科中发现木材导管单生、具梯状穿孔板、导管壁具梯形排列的纹孔以及木射线异型等原始性状,支持五味子科在被子植物中的原始地位。此外,该科木材还具有单穿孔板导管、导管次生壁具螺纹加厚、具分隔纤维等较为特化的性状状态,这种性状进化水平的异等级现象,使五味子科表现出不同进化水平性状的镶嵌组合。根据木材解剖性状对五味子科进行UPGMA聚类分析,所得结果显示南五昧子属和五味子属在木材解剖特征方面有一定的交叉和重叠,这与分子系统学的结论一致,表明这两个属关系密切,可能起源于共同的祖先。通过比较五昧子科与八角科Illiciaceae的木材解剖特征,进一步证明两个科的亲缘关系很近,不支持将五味子科从八角目Illiciales中独立出来成立五味子目Schisandrales的观点。     

广义锦葵科火绳树属4个种的枝条木材解剖及其分类学意义

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

First record of Cercidiphylloxylon (Cercidiphyllaceae) from the Palaeocene of Fushun,NE China

Abstract Cercidiphylloxylon spenceri (Brett) Pearson is described from the Lizigou Formation, Palaeocene in China. The growth rings are distinct; pores are diffuse, solitary, with somewhat angular outlines in cross section; vessel elements long with long scalariform perforation plates; intervessel pitting is opposite to scalariform; fiber‐tracheids are present; axial parenchyma is scarce; rays are mostly biseriate and heterogeneous. All wood characters of the fossil specimen fall into the range of those of extant Cercidiphyllum (Cercidiphyllaceae). The finding is one of the earliest fossil wood records of Cercidiphyllaceae.     

More Malpighiales: Woods of Achariaceae and/or Salicaceae from the Deccan Intertrappean beds,India

The families Achariaceae and Salicaceae (Malpighiales) are characterized by wood anatomical ranges that partly overlap. Formerly these families were treated together in the polyphyletic Flacourtiaceae and a much more narrowly circumscribed Salicaceae. Here we attribute two recently collected fossil woods from the Deccan Intertrappean Beds to the clade that contain these two families, i.e., the Parietal Clade of the Malpighiales. The new genus Elioxylon shares features with several extant genera of Achariaceae and Salicaceae, but does not completely match with any of them. A new record of Hydnocarpoxylon indicum Bande & Khatri is a good match for extant Hydnocarpus Gaertn. (Achariaceae). Elioxylon and Hydnocarpoxylon share an absence of parenchyma, the presence of septate fibres and 1–3 seriate heterocellular rays with long uniseriate margins consistent with Achariaceae and Salicaceae. Elioxylon has mixed simple and scalariform perforations, whereas Hydnocarpxylon has exclusively scalariform perforations. Other Deccan fossils formerly attributed to “Flacourtiaceae” in the literature are critically discussed and mostly excluded from Achariaceae and Salicaceae. Elioxylon and Hydnocarpoxylon from the Maastrichtian ‐ Danian of India are the oldest fossil records of the Parietal Clade of the Malpighiales. With their occurrence on the Indian plate during its northward journey from Gondwana to Laurasia, these fossils provide further support for an ‘out‐of‐India’ hypothesis for Achariaceae and/or Salicaceae. “Baileyan trends” in vessel perforation plate and vessel grouping evolution are apparent in the phylogeny of the Parietal Clade.     

Wood anatomy of Elaeagnaceae, with comments on vestured pits, helical thickenings, and systematic relationships

The secondary xylem of Elaeagnus, Hippophae, and Shepherdia is described and illustrated in detail. Shrubs and small trees of Elaeagnaceae have ring-porous or semi-ring-porous wood with simple perforation plates, vascular tracheids, fiber-tracheids, diffuse or rarely paratracheal axial parenchyma, and uni- or biseriate rays in Hippophae and Shepherdia, but wider rays in Elaeagnus. Walls of vessel elements, especially narrow ones, tracheids, or fiber-tracheids sometimes show helical thickenings; in a few instances these intergrade with small bud-like protrusions associated with pits. Scanning electron microscopy illustrates that small to vestigial vestures are present in all species studied, although nonvestured pits are also common. The analogous nature of vestures and helical thickenings is considered. Comparative wood anatomy suggests a rather isolated position of the family Elaeagnaceae; affinities with Rhamnaceae, Proteaceae, and Thymelaeaceae are discussed.     

Relationships within balsaminoid Ericales: a wood anatomical approach

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

鹅掌楸属(木兰科)次生木质部导管穿孔板的比较研究

利用扫描电子显微镜对鹅掌楸属仅存的2个自然种鹅掌楸和北美鹅掌楸的次生木质部导管穿孔板特征进行了详细的研究。结果显示,鹅掌楸属的2个种均以梯状穿孔板为主,同时存在网状-梯状混合穿孔板。鹅掌楸和北美鹅掌楸的导管穿孔板具有明显差异:(1)鹅掌楸具网状穿孔板,而北美鹅掌楸没有观察到;(2)北美鹅掌楸具单穿孔板,而鹅掌楸在该实验中未发现;(3)北美鹅掌楸具有横隔较粗的梯状穿孔板且横隔数目较多;(4)鹅掌楸的导管穿孔板多数横隔较少;(5)北美鹅掌楸的穿孔板倾斜角度较大;(6)北美鹅掌楸具麻黄式穿孔板的存在,且有纹孔膜残留存在。研究认为,北美鹅掌楸导管分子穿孔板分化较鹅掌楸更为剧烈。     

The first record of fossil Vitaceae wood from the Southern Hemisphere,a new combination for Vitaceoxylon ramunculiformis,and reappraisal of the fossil record of the grape family (Vitaceae) from the Cenozoic of Australia

Austrovideira dettmannaegen. & sp. nov. from the early Oligocene Capella Flora in central Queensland is the first fossil Vitaceae wood described from the Southern Hemisphere. A new combination, Stafylioxylon ramunculiformis (Poole & Wilkinson) Pace & Rozefelds for a Northern Hemisphere fossil wood is also proposed. Austrovideira and Stafylioxylon share with Vitaceoxylon secondary xylem with two diameter classes of vessels, wide vessels usually solitary, narrow vessels forming radial chains, very wide and tall rays, scanty paratracheal parenchyma and septate fibres. Austrovideira differs from Vitaceoxylon in having scalariform intervessel pits and homocellular rays composed exclusively of procumbent cells. This combination of features is seen in the Ampelocissus‐Vitis clade, and a clearly stratified phloem with fibre bands alternating with all other axial elements and phloem rays rapidly dilating towards the periderm is restricted to Parthenocissus and Vitis. Stafylioxylon shares with Austrovideira the presence of scalariform intervessel pits but it differs from that genus in both ray composition and bark anatomy, as it lacks a stratified phloem. These fossil wood genera demonstrate that the lianescent habit in the Vitaceae was established by the Eocene in the Northern Hemisphere and by the Oligocene in the Southern Hemisphere. The pollen and seed fossil record shows that the Vitaceae were in Australia by the Eocene and fossil seeds suggest that the family had radiated by this time. The Oligocene Capella flora with two seed taxa and fossil wood (Austrovideira) provides further evidence of an Australian radiation. The fossil evidence, suggests a significant Gondwanic history for the family.     

The evolution of the atpbeta-rbcL intergenic spacer in the epacrids (Ericales) and its systematic and evolutionary implications

Sequence data from the noncoding region separating the plastid genes atpbeta and rbcL were gathered for 27 epacrid taxa, representing all previously recognized infrafamilial groups, and four outgroup taxa (Ericaceae), to address several persistent phylogenetic questions in the group. Parsimony analyses were conducted on these data, as well as on a complementary rbcL sequence dataset assembled from the literature and the combined dataset. The atpbeta-rbcL spacer was notable for the high frequency of insertion-deletion mutations (indels); their distributions were coded as binary characters and included as a adjunct matrix in some of the analyses. The phylogenetic patterns derived from the spacer and rbcL data and the combined analyses, both including and excluding the indel data, concur in resolving seven major lineages corresponding to the tribes of Crayn et al. (1998, Aust. J. Bot. 46, 187-200), viz. Prionoteae, Archerieae, Oligarrheneae, Cosmelieae, Richeeae, Epacrideae, and Styphelieae. The relationships of the tribes and within Styphelieae, however, are not convincingly resolved. Minor conflicts in the positions of some taxa between the spacer and the rbcL trees are poorly supported. Among epacrids, the spacer region provided more cladistically informative characters than rbcL and resulted in trees with lower homoplasy. Further, the spacer data, when analyzed alone and when combined with rbcL, resolved several clades that could not be retrieved on rbcL data alone and provided increased support for many other relationships. The evolution of a putative three-base inversion associated with a hairpin secondary structure in the spacer region is discussed in the light of the inferred phylogeny.     

WOOD ANATOMY OF THE NEW GENUS MANGLIETIASTRUM LAW OF MAGNOLIACEAE FROM CHINA IN RELATION TO ALLIED GENERA

The present paper contributes the knowledge of wood structure of Manglie- tiastrum sinicum Law, a new monotypic genus of the family Magnoliaceae. The anatomical feature of this wood is described as follows: Diffuse-porous wood ; pores 15-25 per sq. mm, solitary or in radial multiples of 3- 4 (may be up to 6) pore cluster also occur but rare, more or less angular in shape, perforation plates scalariform, with 6-12 bors (may be up to 17), generally 7-8 or 10, intervascular pittings scalariform with fine spiral thickenings; vessel elements 748-1224, generally 876 longl wood rays heterogeneous, type IIA with 1-2, sometimes 3-4 rows of upright cells on both margins, uniserate rays 3-7 cells (may be up to 13) or 193-230u high, the multiseriate rays 3-5 cells or 68-119.2u wide, and 16-22 cells or 340-748u height; oil cells absent, fiber-tracheids 0.8-2.8 mm, generally 1.5 mm long; wood parenchma terminal, narrow banded (generally 7-8 cells wide, may be up to 12), rarely scanty paratracheal.     

The fossil record of Cunoniaceae: new evidence from Late Cretaceous wood of Antarctica?

Fossil angiosperm wood from Upper Cretaceous sediments of Livingston Island and James Ross Island in the northern Antarctic Peninsula region is identified as having the combination of anatomical characters most similar to modern Cunoniaceae. The material is characterised by predominantly solitary vessels, opposite to scalariform intervessel pitting, scalariform perforation plates, heterocellular multiseriate and homocellular uniseriate rays, diffuse axial parenchyma. Anatomically, the specimens conform most closely to the fossil organ genus Weinmannioxylon Petriella which has been placed within the Cunoniaceae. The presence of Weinmannioxylon in Late Cretaceous sediments suggests that taxa within or stem taxa to the Cunoniaceae might have been a notable component of the forest vegetation that covered the Antarctic Peninsula during the Late Mesozoic and may therefore represent the earliest record of this family.