Leaf anatomical characters and their value in understanding morphoclines in the Araceae

Leaf anatomy and petiole anatomy of the Araceae are discussed in terms of their potential use as character states in a phylogenetic analysis. The characters include leaf venation and structure, leaf epidermis, mesophyll ground tissue, vascular bundles, sclerenchyma, collenchyma, laticifers, secretory ducts, and raphide crystals. Characters that seem to have the greatest potential for use in phylogenetic analysis include those of ground tissue, vascular bundles, fibers, trichosclereids, collenchyma, and laticifers. Other, equally distinguishable, characters have states that are apparently autapomorphies, providing little phylogenetic signal. Therefore, although most leaf and petiole structural variation is useful diagnostically, some characters will probably be less valuable in phylogenetic analysis than originally hoped.     

Leaf anatomical studies of Eugenia L. (Myrtaceae) species from the Malay Peninsula

KHATIJAH, H. H., CUTLER, D. F. & MOORE, D. M., 1992. Leaf anatomical studies of Eugenia L. species from the Malay Peninsula. The leaf and petiole anatomy of 25 Malaysian species of Eugenia L. sensu lato was investigated to determine the value of vegetative anatomy for the identification of non-flowering material. Twelve characters proved useful for the identification of species: type of stomata; shape of leaves in T.S.; shape of midrib bundle; sclerenchyma sheath; cutinization of outer epidermal wall; presence or absence of sclereids, idioblasts, hypodermis, columnar epidermal cells, solitary crystals; number of palisade layers and the shape of vascular strand, sclerenchyma sheath and sclereids in the parenchyma of the petiole. Using these characters an anatomical key was constructed to assist identification of the species. Only the closely related species E. stapfiana and E. nilidula could not be separated on anatomical characters.
The anatomical data support the treatment of Henderson (1949) of section Acmena but not of sections Cleistocalyx and Fissicalyx , which fit better in section Syzygium.     

Do leaves in Cyperoideae (Cyperaceae) have a multiple epidermis or a hypodermis?

In Cyperaceae, leaf anatomical characters, in particular the presence of a hypodermis or of a multiple epidermis, have contributed in taxonomic and phylogenetic studies. In this family, the leaf epidermis is often described as uniseriate, and the cells of the subepidermal layers having no chloroplasts are treated as hypodermis. Both tissues have a different ontogenetic origin and hence are not homologous. The aim of the present work was to verify the origin of the subepidermal layers in eight species belonging to Cyperoideae. All species studied presented multiple epidermal layers that were confirmed by leaf ontogeny. In Fimbristylis complanata, F. dichotoma, Pycreus flavescens and P. polystachyos the mature leaves present multiple epidermal layers with cells of the distinct layers similar in shape and size; in the other species studied these cells are different. Especially in the latter case, a multiple epidermis is easily interpreted erroneously as a hypodermis, possibly leading to erroneous evolutionary conclusions. Making correctly distinction between a hypodermis and a multiple epidermis, and hence in case of doubt investigating the origin of the questioned tissue, is compulsory in order to use both characters in a phylogenetic context. Though in the past often called ‘hypodermis’, our leaf ontogenetical observations show that in all species studied, the subepidermical layers constitute a multiple epidermis, originating from the protodermis.     

LEAF CROSS SECTIONS AND PHYTOGEOGRAPHY: A POTENT COMBINATION FOR IDENTIFYING MEMBERS OF FESTUCA SUBGG. FESTUCA AND LEUCOPOA (POACEAE), OCCURRING IN NORTH AMERICA

Icon-like diagrams are used to summarize the anatomical patterns observed in leaf cross sections of 30 taxa of Festuca subgg. Festuca and Leucopoa occurring in western North America. Measurements of four variables, notes on the degree of development and the position of sclerenchyma, along with distribution maps of 24 native taxa, are presented. Twenty-nine diagrams were created from the synthesis of at least ten leaf-section drawings based on five plants chosen from across the geographic range of each taxon (fewer suitable specimens were obtained for two taxa). A key based on data collected in this study allows identification of species in subg. Festuca. Geographic distribution is more reliable than leaf anatomy for separating taxa in subg. Leucopoa.     

Epidermal micromorphology of the genus Festuca L. subgenus Festuca (Poaceae)

Qualitative and some quantitative micromorphological characters of leaf blades, lemmas and paleas were examined using scanning electron microscopy for 29 species of Festuca subgenus Festuca, including the first observations on members of sections Amphigenes, Dimorphae and Cataphyllophorae. The aim of this study was to survey these features in the subgenus Festuca, to assess their taxonomical value. The observed patterns are compared with current taxonomic proposals based on molecular studies. The characters show important taxonomic value, mainly of the abaxial lemma. Characters showing the greatest variation are shape of long cells, shape of the periclinal walls, thickness of anticlinal walls, presence of prickles and abundance of silica bodies. The principal characters of the leaf-blade surfaces include the type and shape of short cells. Two different surface patterns were observed on the lemma, one, long cells rectangular to elongated, with flat to slightly convex periclinal walls and thickened anticlinal walls, for the sections Amphigenes and Dimorphae, and another, long cells rectangular to square, with concave periclinal walls and unthickened anticlinal walls, for the remaining sections. This study confirms the taxonomic value of micromorphological characters in the subgenus Festuca, and that abaxial lemma surface appears to be the most useful.     

Variations in the Leaf Anatomy among some C4 Panicum Species

The Dichotomiflora group of Panicum contains NAD-malic enzyme(ME) species with centrifugal chloroplasts in Kranz cells, NAD-ME(F)species as well as NAD-ME species with centripetal chloroplastsin Kranz cells, NAD-ME (P) species. Many attributes of leafanatomy of 22 C₄ Panicum species were investigated to identifydifferences among four different C₄ subtypes, i.e. NADP-ME,NAD-ME(F), NAD-ME(P) and PEP-CK species grouped by the C₄-aciddecarboxylating enzymes and chloroplast location in Kranz cellsin combination. Differences were found in the number of Kranzcells surrounding a large vein, and the number surrounding asmall vein, the interveinal distances, the proportion of leafcross sectional area occupied by epidermis plus sclerenchyma,by mesophyll cells, by Kranz cells, and by vascular bundles.There were also differences in the ratios of the area of thedifferent cell types. The number of the characters significantlydifferent between a respective pair of C₄ subtypes was the largestbetween NAD-ME(F) and NAD-ME(P) species. In principal componentanalysis applied to 11 leaf anatomical characters, the differentC₄ subtypes clustered into small groups, although the rangeof variations of PEP-CK species and those of NAD-ME(F) speciesoverlapped. The results were discussed in relation to taxonomyand ecological adaptation of Panicum species in the differentC₄ subtypes. C₄ photosynthesis, NADP-malic enzyme, NAD-malic enzyme, Phosphoenolpyruvate carboxykinase, C₄ leaf anatomy, Panicum, Kranz, Dichotomiflora group     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber (‘polystelic’ or ‘monostelic’). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and ‘polystelic’ tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

Pericarp ontogenesis in wild species of Arachis L. (Leguminosae; Papilionoideae)

The genus Arachis encompasses 80 validly described species and the most striking characteristics is the geocarpic fruit, which occurs in every species of the genus. Taking into consideration that much of the current knowledge about this subject is based on the anatomy and morphology of A. hypogaea, a cultivated species with fruits that were modified during domestication, the objective of this study was to learn more about the development of the geocarpic fruits of wild species of Arachis (A. paraguariensis, A. pintoi, A. stenosperma, and A. tuberosa), and to provide characters that can be used in future phylogenetic works about this group. Buds, flowers and fruits at different stages of development were collected and processed according to standard methods used for light field, interferential contrast, and scanning electron microscopy. Although the geocarpic fruits of the wild species share many adaptations, this study identified potentially diagnostic infrageneric characters, such as the type of trichomes on the pericarp, the presence of sclerenchyma caps in the bundles of the pericarp of the seed chamber of the fruit and isthmus, the presence of tracheoid cells in the pith of the isthmus, the location of crystalliferous cells, the presence of tannins in the cells of mesocarp outer layers, and the presence of projections in the sclerenchyma tissue. In addition, this article brings some inferences about functional characteristics of this peculiar geocarpic fruit.     

Leaf anatomy as a contribution to the taxonomy of Salacioideae N.Hallé ex Thorne & Reveal (Celastraceae)

The current classification systems recognize Salacioideae as a monophyletic group within Celastraceae. Nonetheless, some divergences exist for genera: in some cases, most species of the subfamily have been included in only two genera; in others, these genera have been subdivided. This study characterizes the leaf anatomy of 31 species of the subfamily Salacioideae as a contribution to identifying them through features that may also help distinguish among genera. Cross-sections of the median region of the leaf blade and of the petiole and dissociated and macerated epidermis were analyzed. Taxonomically relevant anatomical characters include the type of crystals in the parenchymatous tissue (monocrystals in Cheiloclinium and druses in other genera); the presence of laticifers in Cheiloclinium and Tontelea only; the variable form of the petiole vascular system among studied species; the type of stomata (cyclocytic with two concentric circles of subsidiary cells in P. dulcis; anomocytic in T. attenuata, T. fluminensis, and T. leptophylla; laterocytic in C. anomalum and C. hippocrateoides; and ciclocytic in the other species); the sinuosity of the anticlinal walls of the epidermal cells (sinuous in Cheiloclinium and Peritassa, except P. laevigata, and in S. arborea, S. insignis, S. mosenii, S. nemerosa, and S. opacifolia, and straight in all other studied species); the presence of crystalliferous idioblasts in the epidermis of P. dulcis, P. flaviflora, and P. mexiae; and the presence, form, and disposition of sclereids in the leaf blade, which is a highly variable character among the studied species.     

Leaf anatomy as an additional taxonomy tool for 16 species of Malpighiaceae found in the Cerrado area (Brazil)

This work describes the leaf anatomy of 16 species belonging to three genera of the Malpighiaceae family found in the Cerrado (Minas Gerais State, Brazil). The scope of this study was to support the generic delimitation by contributing to the identification of the species and constructing a dichotomous identification key that includes anatomical characters. The taxonomic characters that were considered to be the most important and used in the identification key for the studied Malpighiaceae species were as follows: the presence and location of glands; presence of phloem in the medullary region of the midrib; mesophyll type; presence and type of trichomes; and presence, quantity, and disposition of accessory bundles in the petiole. It was also possible to indicate promising characters for future taxonomic and phylogenetic studies in the Malpighiaceae family, especially for the Banisteriopsis, Byrsonima, and Heteropterys genera.     

Leaf epidermal characteristics and genetic variability in Central European populations of broad-leaved Festuca L. taxa

Leaf epidermal (micromorphological and micromorphometric) features of the leaf blade were examined with light and scanning electron microscopy in 49 Central European natural populations of species belonging to subgenera Schedonorus and Drymanthele within the genus Festuca L. In addition, molecular biological (random amplified polymorphic DNA) studies were conducted on selected populations. The goal of the study was to reveal the degree of anatomical and genetic variability at the taxon and population level, as well as to determine the most important characters that allow differentiation of the taxa in these two subgenera. The anatomical variation was compared with the genetic distance between taxa. Investigated taxa exhibited great anatomic variability in terms of both qualitative (presence or absence, shape and appearance of various cell types) and quantitative characters (frequency and size of cells). Strong correlation was found between the genetic distance of populations and the frequency and size of silica cells in the costal zone of the epidermis, which indicates that silica cells are largely determined by the genetic background, regardless of varying environmental conditions. Our studies highlight that despite the high level of variability in anatomical features and genetic polymorphism, these fescue populations clearly segregate both in terms of epidermal structure and genetic constitution.     

Comparative leaf anatomy of New Zealand and Tasmanian Inuleae (Compositae)

Leaf anatomy is compared for 47 taxa of the tribe Inuleae (Compositae) from New Zealand and Tasmania. The taxa are tabulated according to their lamina structure. Noteworthy anatomical features are lamina type and grade of differentiation, stomatal position, epidermis and cuticle thickness, shape of palisade cells, number of bundle-sheath layers; presence or absence of sclerenchyma caps, water storage cells, secretory canals, abaxial collenchyma and protruding midrib. It is shown that leaf anatomy data provide characters which are taxonomically useful in classification of Inuleae.     

Leaf morphology and anatomy of Manilkara Adans. (Sapotaceae) from northeastern Brazil

Aiming to complement the group’s taxonomy, the objective of this work is to describe and characterise, searching for morpho-anatomical patterns, the leaf anatomy of the Manilkara species occurring in the northeast region of Brazil. The leaves of all Manilkara species were analysed, using herbaria specimens and collections made in the field by the authors. The characterisation of the leaf blades was performed according to the usual protocols in vegetal anatomy. As diagnosed by anatomical characters, the Manilkara Adans. genus presents petioles with a thick cuticle, a uniseriate epidermis, a sclerenchymatous sheath involving the vascular bundle, and laticifers on the cortical and medullar regions. The anatomical data increase the M. elata and M. huberi synonymisation and show the importance of the anatomy as a complementary tool to the taxonomy based on external morphology.     

鸢尾属（Iris）植物叶片表皮微形态特征的研究

对12种鸢尾叶片的表皮特征进行了观察和研究,结果表明野鸢尾、单花鸢尾、北陵鸢尾3种鸢尾各自的上下表皮细胞的形状及气孔密度等有明显区别,而其余9种鸢尾叶片各自上下表皮形态均无明显差异。12种鸢尾的气孔类型均属于横列型,且气孔均是随机分布,气孔保卫细胞的长轴与叶脉平行,叶脉处无气孔分布。不同种鸢尾之间叶片表皮特征具有显著差异,可为鸢尾属植物的分类提供依据。     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber ('polystelic' or 'monostelic'). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and 'polystelic' tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

Leaf anatomy of Ficus subsection Urostigma (Moraceae)

Species of Ficus subsection Urostigma show much overlapping variation in vegetative morphology, which often precludes correct identification of the species. The aim of this study was to describe the leaf anatomical characters and their variation and to check their suitability for identification. Included were 41 samples belonging to 25 species of subsection Urostigma, four samples belonging to two species of section Leucogyne and one specimen of Ficus glaberrima subsp. siamensis of subsection Conosycea. Transverse sections of lamina, midrib and petiole and cuticular macerations were used, and the observed anatomical characters are described for each species. On the basis of a limited number of studied samples, leaf anatomy shows little variation within each species and each species has a unique combination of character states, facilitating identification. Ficus arnottiana shows some leaf anatomical characters that are quite different from those of other members of subsection Urostigma, including a multiple epidermis and enlarged lithocysts on both sides of the leaf. Both characters are generally considered as typical for Ficus subsection Conosycea. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 259–281.     

八角目(Illiciales)植物花被片表皮比较形态

在光学显微镜和扫描电镜下,观察了八角科(Illiciaceae)八角属(Illicium Linn.)2组11种20个材料、五味子科(Schisandraceae)南五味子属(Kadsura Ksempf. ex Juss.)2亚属2组8种15个材料和五味子属(Schisandra Michx.)2亚属4组6种17个材料的花被片表皮形态特征。首次报道了八角目2个科（八角科和五味子科）3个属（八角属、南五味子属和五味子属）植物花被片表皮细胞的形状、分泌细胞的形状及分布、气孔器的形状及分布、花被片表面的纹饰。通过与八角目2个科3个属植物的叶表皮形态比较,发现花被片表皮气孔器外拱盖均为单层,与叶表皮气孔器外拱盖层数（常绿种类为双层和落叶种类为单层）之间没有相关性,还在五味子科中发现2个新的性状（气孔对和环列型气孔）。通过对两性花、雌花和雄花花被片表皮观察,发现花被片表皮形态与花的性别之间没有相关性。通过对八角属、南五味子属和五味子属花被片表皮比较,发现五味子属与南五味子属相比,其花被片表皮表现出更多的衍生性状;南五味子属与八角属相比,前者花被片表皮具有更多的衍生性状;而南五味子属花被片表皮形态与五味子属的相似性程度较大,支持五味子科包含南五味子属和五味子属、八角科包含八角属的观点。     

Leaf anatomy of the subtribe Conceveibinae (Euphorbiaceae)

A comparative study of the leaf anatomy of the subtribe Conceveibinae was carried out. Leaves show a similar structural arrangement in all species, however, there are small variations among the taxa that allow definitions of species or groups. It is not possible to recognize infrageneric divisions or to maintain some genera as different from Conceveiba. The analysis included the study of 11 characters of taxonomic importance of the leaf: the form, the size and the apex of the papillae of the lower side of the blade, the number of layers of the palisade parenchyma, the presence or absence of the midrib pith, the number of vascular bundles of petiole as well as the central cylinder and the medulla, and the presence or absence of the sclerenchyma in the different levels of the petiole. The results are evaluated from a taxonomical point of view. With leaf anatomy characters only, it is possible to separate the genus Conceveiba in the subtribe.     

Comparative anatomy of conifer leaves

The epidermis and internal structure of 103 species and 8 varieties of conifer leaves belonging to 37 genera of 7 families have been comparatively investigated. The anatomical characters of leaves of all the genera (or sections) studied are described systematically and a key to the genera (or sections) is also presented. It is shown that in order to identify at the genus level, the leaf types of LaubenFels’ may be used as the primary characterization. And it can also be seen that the following characters which are relatively constant at the genus level provide an important basis for identification: the epidermal structure, the presence or absence of hypodermis, the degree of differentiation of mesophyll, the occurrence of vascular bundle sheath or endodermis, the transfusion tissue type, the number and relative spatial position of vascular bundles, the number and distribution of resin canal and the presence of sclereid or sclerenchyma. The genera of Cupressaceae (except for Juniperus and Sabina) are of scale leaves (Leaf type-III) with little differences in structural characters. It’s difficult to distinguish them from each other. In addition, some problems concerning the significance of the resin canal, the endodermis with Casparian-strip, the sclerenchyma and the vascular bundle in identificationare here also discussed.     

Epidermal micromorphology of the genus Festuca L. (Poaceae) in the Iberian Peninsula

Micromorphological features of the leaf-blade, lemma, and palea were examined under scanning electron microscopy (SEM) for 64 species of the genus Festuca L. occurring in the Iberian Peninsula. The objective of the study was to survey these characters for the first time in the whole of the genus Festuca in this region, to assess their taxonomic value, and to compare the variation with recent DNA sequence-based phylogenies. The principal features include the frequency, shape, and distribution of silica bodies, short cells, crown cells, and prickles, and the morphology of long cells. The abaxial lemma surface shows the most important taxonomical and phylogenetic characters. The variation observed was consistent with the subgenera and sections identified in molecular studies.