Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae) III. Genetic Variation and Taxonomic Value of Morphological Characters

Adenophora potaninii Korsh．complex is a morphologically variable group including six species．As shown in many species in the genus Adenophora,the variability of many morphological characters in the complex under investigation is amazingly great,which brings difficulties in the delimitation and indentification of taxa．In the present study,six populations representing five species in the complex were systematically sampled to investigate the pattern of morphological variation within population and to detect the genetic basis of the variation with progeny tests and controlled crosses．The results are as follows： Great morphological variation within population is found (Fig.1),including characters such as leaf shape,teeth number and size of leaf margins,teeth number of calyx lobes,and indumentum on the surface of stems and leaves,which were previously considered as diagnotic．Systematic sampling and statistical analysis show that the differences in the above characters exhibit continuous patterns of variation within population (Fig．1,2),though they are highly correlated (Table 2)．From progeny testing and crossing between two contrast types of individuals (oblong,dentate and pubescent leaves vs．narrow,entire and glabrous leaves)it is evident that those characters show continuous variability in segregating progenies (Fig.3,4)and appear to be influenced by large numbers of loci with individually slight effects．Since most wild populations were highly heterozygous for those characters,the individuals in one extreme of variation can produce their variable offsprings including individuals similar to those in the other extreme (Fig. 3). This is also true for the indumentum density on stems and teeth number of calyx lobes. In this complex, A. biformifolia Y. Z. Zhao, A. bockiana Diels and A. polydentata P.F. Tu et G. J. Xu were described exclusively based on leaf shapes, teeth number of leaf margins and calyx lobes, teeth size of leaf margins, and indumentum on the surface of stemsand leaves. It is demonstrated, however, that the variation of those characters is of a quantitative nature and show no discontinuities. As a result, genetic analysis of diagnostic characters along with their sympatric distribution and same habitats, strongly suggest that A.biformifolia as well as A. bockiana and A. polydentata are actually the extreme individuals within A. wawreana and A. potaninii respectively and should not be recognized as taxa at any taxonomic level.     

The systematic anatomy of south Indian Cyperaceae: Cyperus L. subgen. Juncellus, Cyperus subgen. Mariscus and Lipocarpha R. Br.

The three investigated species of Cyperus L. subgen. Juncellus (Griseb.) C. B. Clarke show anatomical differences in the leaf pertaining to the number of vascular bundles and bundle sheaths, size of silica-cells and their wall thickness, while those of subgen. Mariscus (Gaertner) C. B. Clarke segregate into two primary groups depending on the presence or absence of a distinct hypodermis; and the species within each group are further differentiated on the basis of the presence or absence of air-cavities, the number of vascular bundles and their pattern of arrangement in the leaf. The interspecific differences observed within the genus Lipocarpha R. Br. concern the size of the interstomatal cells and the number of vascular bundles and bundle sheaths of the leaf. Keys for the identification of the species in each genus and subgenus are given separately.     

中国29种和泰国1种卷柏科植物的叶形态学研究及其分类学意义

利用光学显微镜和体视显微镜对卷柏科Selaginellaceae中国产29种和泰国产1种植物的叶形态进行了宏观和微观观察。结果表明卷柏科植物在叶形态学方面表现出较高的一致性。(1)叶多为二形, 罕为一形, 无叶柄, 叶二形者侧叶多大于中叶, 外展或斜上, 中叶贴生于腹面茎上, 交互覆瓦状排成4列, 叶一形者则在茎上作螺旋状排列。(2)侧叶与中叶上表皮细胞形态相似, 近等径方形或长方形, 垂周壁平直、弓形或波状; 侧叶与中叶下表皮细胞形态也相似, 长方形或长条形, 垂周壁波状。(3)气孔类型为无规则型或放射型, 气孔多分布于中叶上表皮和侧叶下表皮中脉处。研究发现, 叶外形、叶表皮细胞形态及气孔分布特征等为中国卷柏科植物种间分类提供了重要的参考依据。 研究的有关种类可以依据下面这些特征进行分类: (1)叶为一形叶或二形; (2)叶在枝上着生为螺旋状或成对交互排列; (3)叶表皮细胞垂周壁为平直、弓形、深波状或浅波状; (4)叶上、下表皮细胞同形或异形; (5)气孔为无规则型或放射型; 以及(6)气孔在中叶和侧叶的上、下表皮上的分布等特征。     

泡沙参复合体(桔梗科)的物种生物学研究——Ⅲ.性状的遗传变异及其分类价值

阐明性状变异的遗传基础是评估性状分类价值以及揭示类群关系和进化过程的前提。本文根据泡沙参复合体内6个天然居群的材料,利用居群样本分析、子代测定和杂交试验等手段,结合统计学方法对该复合体一些在以往类群划分中起鉴别作用的形态性状进行了遗传学分析。结果表明,叶形、叶缘锯齿数目和大小、茎叶被毛、花萼裂片齿数、花梗长短等性状都是遗传性比较强的性状,而且性状之间有很强的相关性。但是,这些性状在居群内呈现连续变异且变异幅度非常大,以致于变异的两个极端个体被分别作为不同的种处理。通过对野外居群样本的统计分析和对极端变异类型个体所进行的子代测定及其人工杂交,进一步证实上述性状是受多基因决定的数量性状,其变异幅度大,变异式样中并不存在任何程度的间断性,因而至少在本复合体内没有鉴别意义。因此,以往建立在上述性状变异基础上的3个类群(A.biformifolia Y.Z.Zhao;A.bockiana Diels和A.polydentata P.F.Tu et G.I.Xu)均不成立。最后,本文对形态性状变异的遗传基础及其与类群划分的关系进行了讨论。     

The new species Ettingshausenia sarbaensis (Angiospermae) from the Cenomanian-Turonian of Western Kazakhstan in light of the problem of classification of dispersed cretaceous Platanus-like leaves

Based on morphological and epidermal characters of the leaf, a new species, Ettingshausenia sarbaensis sp. nov., from the Cenomanian-Turonian of western Kazakhstan is described. In fossil Platanus-like leaves, the unique epidermal structure, with the combination of encyclocytic, laterocytic, and paracytic stomata and the presence of trichomes on one or from two to seven epidermal cells, is shown for the first time. Based on variations in morphological and epidermal characters, two morphotypes corresponding to groups of sun and shade leaves are recognized. The expediency of consideration of dispersed Cretaceous Platanus-like leaves within the framework of morphological classification of leaves is discussed.     

Leaf structure of Bromelia and its significance for the evolution of Bromelioideae (Bromeliaceae)

This study investigated the leaf structure of Bromelia and its importance for understanding the evolution of Bromelioideae (Bromeliaceae). Because of the scarcity of informative molecular characters in Bromeliaceae, this study evaluates the relevance of anatomical characters for the taxonomy and phylogeny of Bromelia and the subfamily Bromelioideae. Anatomical studies in monocots have shown that the combined application of anatomical and external morphological characters from leaf structure can improve the taxonomic delimitation of species, genera and subgenera, and is very informative for inferring phylogenies. The current study analyzed the leaves of 27 species of Bromelia and found that the most important characters for the systematics of this group are the occurrence of a water storage hypodermis, the number of stalk cells of peltate scales, the presence of a ribbed abaxial surface, the occurrence of palisade parenchyma on the adaxial side, the shape of the cells that surround the air lacunae, the presence of raphides and secretory channels, and the occurrence of fibrous extensions on the bundle sheath on minor veins. Combining our results with those described for the family, we made a list of the anatomical characters that can be used in phylogenetic studies of Bromelioideae.     

Micro morphological Characteristics of Leaf Epidermis under Light Microscopy and Its Taxonomic Significance in Persea (Lauraceae) from America

Leaf epidermal micro morphological characters of 54 species belonging to genus Persea (Lauraceae) from America were comparatively studied by the means of light microscopy. It was shown that the leaf epidermal cells were usually regular bearing with straight or beaded anticlinal walls.Most of these species had unicellular trichomes uniformly distributed. The stomatal apparatus which were paracytic, only randomly presented in the areoles of abaxial epidermis for all sampled species, can be assigned into three types. The results showed that similar leaf epidermal characters can be found in the species with similar morphology. Nineteen characters were selected for PCA analysis, and the sampled species was divided into two clades. The characters of leaf epidermis were little different within a species but can vary among species. However, the multiple leaf epidermal characters including the shape of epidermal cells, the shape of stomatal apparatus, stomatal ledges and some distinct characters were relatively constant and taxonomically quite useful for the identification of some species with similar phenotypes.     

兰属、兜兰属、石斛属植物叶片的扫描电镜观察

对兰科植物的兰属、兜兰属及石斛属１６个种折叶片及其横断面进行了扫描电镜的观察。兰属各种叶片上表皮细胞均为矩形,上表皮细胞表面具小乳突或不明显突起。石斛属及兜兰属的各个种上下表皮细胞均为多边形,但石斛属表皮细胞表面无坦无纹饰,而兜兰属花叶类上表皮细胞表面明显呈乳突状,绿叶类呈龟背状隆起。兰属及石斛属叶片叶肉组织没有栅栏组织及海绵组织的分化,而兜兰属的绿叶类叶肉不分化;花叶类叶肉有分化。     

Degradation of the upper pulvinus in modern and fossil leaves of Cercis (Fabaceae)

Identification of fossil leaf impressions as Cercis has been questioned based upon the presence or absence of a pulvinus at the base of the lamina (upper pulvinus). In the present study, leaves of Cercis canadensis were examined before and after abscission to explore the degradation processes that could occur prior to fossilization, and the North American record for fossil foliage of Cercis was revised accordingly. Results for C. canadensis indicate that: (1) the pulvinus consists largely of tissues with nonlignified cells (a wide cortex, a nonlignified fiber sheath, phloem, and pith) that degrade rapidly after leaf abscission, (2) the lignified xylem tissue that remains in the pulvinus after degradation is in brittle strands, (3) the pulvinus degrades at a faster rate than the lamina or the petiole, and (4) the degraded pulvinus cushion leaves a semicircular pattern on the lamina. From examination of fossils as well as extant species, we: (1) demonstrated that in fossils, the upper pulvinus can show a greater degree of degradation than the adjoining petiole or lamina tissue, suggesting the degradation of upper pulvinus tissue is similar in modern vs. fossil specimens, (2) defined numerous other laminar characters that can be used in conjunction with, or in the absence of, an upper pulvinus to confirm the presence of Cercis in the fossil record, and (3) showed from those criteria that the earliest known North American fossil leaf record for Cercis, from a specimen newly reported in the present study, is from the middle Miocene Succor Creek flora of Oregon.     

内蒙古退化草原8种植物叶结构对禁牧的响应

在内蒙古典型草原地带,选择有代表性的羊草＋大针茅群落的放牧退化演替变型—冷蒿＋糙隐子草群落实行封育禁牧。并对未禁牧群落（退化群落）、禁牧7a的群落和禁牧20a群落的8种主要植物叶的解剖结构特征进行比较研究,以探讨植物叶结构对禁牧的响应。结果表明：禁牧恢复演替后叶片角质层厚度和叶肉细胞密度降低、5种植物叶片变薄、星毛委陵菜上表皮细胞明显变小,植物叶旱生特征减弱。但草地退化后植物叶又不完全表现为旱化或加强旱化的特性。从植物叶结构的变化中可见星毛委陵菜具有较强的耐牧性。对于植物叶结构其它指标,则不存在明显的一致性变化趋势。     

Didymodon kunlunensis D.P.Zhao,S.Mamtimin & S.He (Pottiaceae), a new species from Xinjiang,China

Didymodon kunlunensis D.P.Zhao, S.Mamtimin & S.He is described as a new species found in Xinjiang, China. The new species is characterised by a combination of the following characters: ovate leaves with laminal cells always smooth, red colour in KOH, recurved leaf margins from base to apex, a percurrent costa without dorsal and ventral stereid bands in cross section, a distinct stem central strand and the presence of gemmae in the leaf axils. Light microscope photographs of the significant characters are provided and its distinctions from closely related taxa are discussed. A key to Chinese Didymodon species with smooth laminal cell is provided.     

Hybridization and introgression in Carpobrotus spp. (Aizoaceae) in California. I. Morphological evidence

Individuals of the invasive non-native plant species Carpobrotus edulis, its native or naturalized congener C. chilensis, and putative hybrids are found throughout coastal California. This study characterized morphological variation within the genus and assessed the extent of hybridization and direction of introgression between C. edulis and C. chilensis. Twelve randomly selected Carpobrotus individuals from each of 49 sample locations were classified as "C. edulis," "C. chilensis," or one of three intermediate morphotype classes, and compared for clonal, vegetative, and fruit characters. Univariate comparisons showed that C. edulis individuals had deeper mats with more cover within the mat boundary, longer, wider, more serrate leaves, and wider fruits with more carpels than intermediate and C. chilensis morphotype classes. Carpobrotus edulis and intermediates had more lateral branches than did C. chilensis, suggesting that hybrids tend to vary toward C. edulis. Discriminant function analysis (DFA) identified the following characters as discriminating significantly among Carpobrotus groups: leaf length, leaf width, leaf length/width, number of lateral branches, number of carpels, and fruit diameter. Using these characters, DFA correctly classified 98% of apparently pure C. edulis and C. chilensis, but only 76% of all Carpobrotus individuals when putative hybrids were included. Principal components analysis showed that C. edulis and C. chilensis cluster into distinct morphotypes, but that hybrids overlap with both groups, particularly with C. edulis. The genus Carpobrotus in California resembles a large hybrid swarm, with putative hybrids forming a large portion of the overall population and tending to vary toward C. edulis.     

Leaf and twig anatomy of Eriope, a xeromorphic genus of Labiatae

The leaf and twig anatomy of 25 species of the genus Eriope were studied. The twig anatomy is very uniform apart from the level of formation of early layers of cork. Leaf anatomy shows considerable variation between the species, and this is correlated to some extent with the extreme habit range from trees to woody herbs. Characters of the lamina that show variation are: trichome type and frequency, cuticular markings, leaf dorsiventral or isobilateral, presence of adaxial stomata, presence of a hypodermis, number of layers of adaxial palisade mesophyll cells, occurrence of large bundles of phloem fibres at main veins, type of areolation and marginal venation. Petiole vasculature is simple and generally with either four distinct vascular bundles or two vascular arcs. The most xeromorphic species are usually woody herbs or sub-shrubs, and tend to have thick, isobilateral leaves with large bundles of phloem fibres and few hairs, or strongly dorsiventral leaves with a hypodermis and stomata in deep abaxial hair-lined depressions. The correlation of xeromorphic characters with environmental conditions is discussed. Leaf anatomy is of limited value in elucidating relationships within the genus.     

Leaf anatomy of the Pittosporaceae R. Br.

WILKINSON, H. P., 1992. Leaf anatomy of the Pittosporaceae R. Br. An anatomical study of the leaves of 58 species representing all nine genera has been made. The anatomical characters found to be of most use in distinguishing taxa are: in surface view-cuticular architecture as seen with the SEM, stomatal outline, occasionally stomatal density, presence/absence of hairs, hair type in adult leaves; in transverse section-petiole/midrib outline, midrib number of vascular bundles and number of secretory ducts, adaxial epidermis in 1/2 layers, dimensions of adaxial epidermal cells, thickness of outer wall of adaxial epidermis, occasionally chlorenchyma interrupted/not interrupted above the midrib vascular tissue in Pittosporum species; leaf margin.     

Patterns of intra- and interspecific association in leaf-mining insects on three oak host species

Abstract. 1. We determined mortality and distributional patterns of leaf miners on three oak host species (Quercus falcata, Q.nigra and Q.hemisphaerica) in northern Florida, U.S.A.
2. Patterns of intra- and interspecific occurrence within leaves, and mortality of five most abundant leaf miner species were analysed as a test of competition.
3. Miners co-occurred on leaves more often that expected by chance (P<0.05) in six of ten possible species combinations and log-linear model analysis showed no negative higher-order interactions.
4. All five miner species had highly clumped distributions between leaves (P<0.01).
5. Leaf miner survival was less than expected for four of five species when co-occurring on leaves with conspecifics than when mining with heterospecifics or alone (P<0.05).
6. We conclude that interspecific competition is unapparent within this leaf miner guild and that intraspecific competition occurs in four of the five major leaf miner species. We discuss leaf miner selection of common leaves, perhaps based on chemical/physical leaf characters, as a cause of intra- and interspecific aggregation.     

Leaf Anatomy of Fourteen Species in Carex Subgenus Indocarex (Cyperaceae)

Among the subgenera of the genus Carex, the subgenus Indocarex has been seldom studied in any respects, Its systematic position and its subdivision are still disputable. Leaf anatomy of 14 species in the subgenus lndocarex from China was studied. The anatomical characters are proved to be systematically valuable. (1) Characters of lamina transverse section: All leaves of these 14 species are dorsiventral. The outline mostly V-shaped, occasionally flat or nearly flat, with adaxial lateral rib in each half of lamina and some of them flanged. The cells of adaxial surface larger than those of abaxial surface, and the epidermal cells over veins usually smaller than others. Air-cavities between vascular bundles are well developed, and bulliform cells also well developed in most taxa. The vascular bundles are collaterai, bundle sheaths double-layered, and the outer sheath parenchymatous and the inner sheath fibrous. (2) Characters of lamina epidermis: The shape of the cell on both surfaces is generally rectangular, and the anticlinal wall of epidermal cell sinuous; stomata is paracytic, elliptic to oblong, rarely sub-circular; prickles occur on adaxial surfaces of certain species; papillae are only obvious on abaxial surface of C. moupinensis Franch. The characters of transverse section and epidermis of leaf blades of these 14 species differ from each other to certain degree, and closely related species are similar in anatomical characters. The anatomical characters of lamina are of value for classification at specific and sectional level of the subgenus Indocarex. Despite of the variation of these characters among species, a certain num ber of characters appears to be shared by the members of the subgenus, and some of the common characters are primitive. In addition, some gross morphological characters are common and primitive also. Therefore, the subgenus Indocarex may be primitive in the genus Carex. The anatomical and morphological characters of C. scaposa C. B. Clakre and C. densifimbriata Tang et Wang ex S. Y. Liang are distinct. The two species and their allies should be treated as section instead of subsection. The three species in the sectionPolystachyae share some anatomical characters and comprise a coherent group.     

Cyanide-resistant respiration in roots and leaves. Measurements with intact tissues and isolated mitochondria

A comparison was made between the oxygen uptake of roots and leaves and of mitochondria isolated from the same tissues. Ten species were included in this study: three legumes, one C3-monocotyledon, one C4-monocotyledon, the rest non-leguminous C3-dicotyledons. Root and leaf respiration in all species examined displayed substantial resistance to KCN (0.1–1.0 mM) and the cyanide-resistant respiration was completely inhibited by salicylhydroxamic acid (SHAM; 10–20 mM). SHAM alone inhibited oxygen uptake to varying degrees, depending on the species. Mitochondria were isolated from roots and leaves of many of the species examined and also displayed cyanide-resistant oxygen uptake, which was sensitive to both SHAM and tetraethylthiuram disulfide (disulfiram). Concentrations of SHAM greater than 2 mM caused inhibition of the cytochrome path as well as of the alternative path in isolated mitochondria. Respiration rates of intact roots and leaves in the presence of varying concentrations of SHAM alone were plotted against those obtained in the presence of both SHAM and KCN. This plot showed that in vivo the cytochrome pathway was not affected by 10 or 20 mM SHAM in the external solution. We conclude that the activity of the alternative pathway in intact roots and leaves can be reliably estimated by comparing SHAM-sensitivity and cyanide-resistance of respiration.     

苔草属复序苔草亚属十四种植物叶片的解剖学研究

选择中国复序苔草亚属6组4亚组的代表植物14种,进行了叶片解剖学研究,观察了其横切面 和表皮特征,证明上述特征在各类群之间存在差异,具有一定的系统学意义。这14种植物叶片的横切 面和表皮都具有一些原始的性状,表明复序苔草亚属中的植物可能在苔草属中是较原始的。在所观察 的植物中,Sect．Polystachyae植物叶片解剖学特征比较一致,说明此组的建立比较合理;而Sect．Indicae 组已有明显分化,尤其是Carex scaposa C．B．Clarke和C．densifimbriata Tang et S．Y．Liang 与其它植物明显不同,而且其外部形态特征在复序苔草亚属中也比较独特,因此赞成将它们及其近缘类群做为一个组而非亚组。     

广西石韦属七种植物叶片结构与孢子形态的比较研究

利用石蜡切片法研究了石韦属7种植物叶片的形态结构。结果显示:7种植物的叶片均属异面叶,且具典型的旱生叶结构;表皮大多为复表皮,表皮细胞排列紧密,主脉表面覆有厚角质膜,表皮毛为星状毛,由多细胞组成;气孔集中分布于下表皮,气孔类型为围绕型,气孔器略下陷;孢子两侧对称,极面观椭圆形,赤道面观豆形或超半圆形,表面饰纹为瘤状饰纹。石韦属7种植物的叶表皮细胞形状、表皮毛、表皮细胞垂周壁式样、气孔密度等解剖结构均表现出一定的种间差异,这些特征可为石韦属植物种间分类提供依据。