Anatomical characterization of Strobilanthes (Acanthaceae) species from the northern Western Ghats of India and its implication for identification at vegetative state

A comprehensive study of stem, leaf and petiole anatomy of 10 species of Strobilanthes from northern Western Ghats of India was carried out to identify characteristics which would enable species identification when flowering material is unavailable. In Strobilanthes, some species bloom annually, others are plietesials, i.e. they grow without blooming for several years and then produce huge quantities of flowers, release seeds and die. Therefore, alternative methods, such as anatomical characters, are essential to distinguish Strobilanthes species in their vegetative stage. We collected ten species of Strobilanthes for anatomical characterization. Under the bright‐field microscope, stem cross‐sections of different species were found to be undulate, quadrangular, quadrangular‐winged or terete. Study of the stem revealed a distinct outer and inner cortex, the distribution of cystoliths (CaCO₃ crystals), raphides (CaC₂H₂O₅ crystals) and sclereids which varied from species to species. Study of leaf anatomy showed structural variation and vascular bundle shapes that differed between the species. Leaf epidermal characters under light and scanning electron microscopy exhibited variation in characters such as stomatal index, stomatal length and width, stomatal type and presence of glandular and non‐glandular trichomes. The petiole anatomy was species‐specific, especially with respect to vascular bundle structure and the distribution of structures such as sclereids, cystoliths, sphaeraphides and tannin cells varied. Hence, unique anatomical features of the stem, leaf and petiole could be used as taxonomic characters to identify Strobilanthes species in a vegetative state.     

The comparative stem and root anatomy of Goniothalamus andersonii, G. macrophyllus, G. malayanus and G. velutinus (Annonaceae) from the peat swamps of Sarawak

The stem and root anatomy of G. andersonii, G. macrophyllus and G. velutinus from the peat-swamps of Sarawak is compared. Sufficient anatomical differences exist to differentiate the four species. The main distinguishing anatomical features between the stems are the cork cells, the nature and relative quantity of cortical sclereids and the presence or absence of lignified pith parenchyma. In differentiating between the roots of G. andersonii, G. macrophyllus and G. malayanus. the principal features are the cork cells and the relative quantity of cortical sclereids. In addition, the medullary rays of G. macrophyllus root in transverse sections are noticeably broader in the xylem towards the phloem than those of the other species.     

Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics

CARLQUIST, S., 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Qualitative and quantitative data, based mostly upon liquid-preserved specimens, are presented for Akebia, Roquila, Decaisnea, Holbodia, Lardizabala, Sinofranchetia and Stauntonia . Because Decaisnea is a shrub whereas the other genera are vines, anatomical differences attributable to the scandent habit can be considered. These include exceptionally wide vessels, a high proportion of vessels to tracheids (or other imperforate trdcheary elements) as seen in transection, simple perforation plates, multiseriate rays which are wide and tall, and pith which is partly or wholly sclerenchymatous. With respect to ecology, two features are discussed: spirals in narrower vessels may relate to adaptation to freezing in the species of colder areas, and crystalliferous sclereids seem adapted in morphology and position to deterrence of phytophagous insects or herbivores. The wood may provide mechanisms for maintaining conduction even if wider vessels are deactivated temporarily by formation of air embolisms. Wood and stem anatomy of Lardizabalaceae compare closely to those of Berberidaceae and of Clematis (Ranunculaceae), as well as to other families of Berberidales. Decaisnea is more primitive than these in having consistently sralariform perforation plates and in having scalariform pitting on lateral walls of vessels. A tentative listing of anatomical features which may correspond to generic limits is given.     

Noteworthy idioblastic sclereids in the stems of Eulychnia (Cactaceae)

Stems of Eulychnia (a genus of six to nine species of candelabriform or arborescent cacti) have a parenchymatic cortex with two distinct regions. The outer chlorenchymatic layer is characterized by a conspicuous parallel striping, whereas the inner cortex region devoid of chlorophyll has a coarsely granular aspect. Stem samples from nine accessions, collected in the field or taken from cultivation, were studied from resin-embedded microtome sections and maceration. Two different forms of lignified sclereids were found dispersed in the cortex and the pith. The sclereids of the outer palisade-like cortex layer are distinctly elongated and strictly oriented at right angle to the stem surface, whereas those of the inner cortex and pith are globular or subglobular and conspicuously enlarged compared with the surrounding parenchyma cells. The ontogeny of the sclereids was studied from stem samples of different ages. Formation of the secondary cell walls starts only after cell growth is completed. A screening of numerous South American cacti for the presence of idioblastic sclereids showed that these structures are unique for the genus Eulychnia. Finally, functional aspects of the sclereids are shortly discussed. It is assumed that the sclereids contribute to the mechanical support and reinforcement of the plants.     

Stem anatomy of the dwarf subshrub Cressa cretica L. (Convolvulaceae)

Stem anatomy and development of medullary phloem are studied in the dwarf subshrub Cressa cretica L. (Convolvulaceae). The family Convolvulaceae is dominated by vines or woody climbers, which are characterized by the presence of successive cambia, medullary- and included phloem, internal cambium and presence of fibriform vessels. The main stems of the not winding C. cretica shows presence of medullary (internal) phloem, internal cambium and fibriform vessels, whereas successive cambia and included phloem are lacking. However, presence of fibriform vessels is an unique feature which so far has been reported only in climbing members of the family. Medullary phloem develops from peri-medullary cells after the initiation of secondary growth and completely occupies the pith region in fully grown mature plants. In young stems, the cortex is wide and formed of radial files of tightly packed small and large cells without intercellular air spaces. In thick stems, cortical cells become compressed due to the pressure developed by the radial expansion of secondary xylem, a feature actually common to halophytes. The stem diameter increases by the activity of a single ring of vascular cambium. The secondary xylem is composed of vessels (both wide and fibriform), fibres, axial parenchyma cells and uni-seriate rays. The secondary phloem consists of sieve elements, companion cells, axial and ray parenchyma cells. In consequence, Cressa shares anatomical characteristics of both climbing and non-climbing members. The structure of the secondary xylem is correlated with the habit and comparable with that of other climbing members of Convolvulaceae.     

A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae)

MIDDLETON, D. J., A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae). Leaf and stem anatomical characters of 123 species within the Gaultheria group of genera of the tribe Andromedeae are surveyed. A number of characters show considerable variation within a species which limits their taxonomic use. Other characters such as marginal sclerenchyma and, to a lesser extent, the hypodermis, the presence of adaxial stomata, the presence of free fibres and the pith type were of more taxonomic interest. These characters are discussed in relation to the generic and infrageneric classification of the group.     

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.     

Major trends in stem anatomy and growth forms in the perianth-bearing Piperales,with special focus on Aristolochia

Background and Aims

The order Piperales has the highest diversity of growth forms among the earliest angiosperm lineages, including trees, shrubs, climbers and herbs. However, within the perianth-bearing Piperales (Asarum, Saruma, Lactoris, Hydnora, Prosopanche, Thottea and Aristolochia), climbing species only occur in the most species-rich genus Aristolochia. This study traces anatomical and morphological traits among these lineages, to detect trends in growth form evolution and developmental processes.

Methods

Transverse stem sections of different developmental stages of representatives of Asarum, Saruma, Lactoris, Hydnora, Thottea and Aristolochia were compared and anatomical traits were linked to growth form evolution. Biomechanical properties of representative climbers were determined in three-point bending tests and are discussed based on the anatomical observations. Growth form evolution of the perianth-bearing Piperales was reconstructed by ancestral character state reconstruction using Mesquite.

Key Results

While species of Asarum and Saruma are exclusively herbaceous, species of the remaining genera show a higher diversity of growth habit and anatomy. This growth form diversity is accompanied by a more complex stem anatomy and appropriate biomechanical properties. The ancestral growth form of the perianth-bearing Piperales is reconstructed with either a shrub-like or herbaceous character state, while the following three backbone nodes in the reconstruction show a shrub-like character state. Accordingly, the climbing habit most probably evolved in the ancestor of Aristolochia.

Conclusions

Since the ancestor of the perianth-bearing Piperales has been reconstructed with a herb- or shrub-like habit, it is proposed that the climbing habit is a derived growth form, which evolved with the diversification of Aristolochia, and might have been a key feature for its diversification. Observed anatomical synapomorphies, such as the perivascular fibres in Lactoris, Thottea and Aristolochia, support the phylogenetic relationship of several lineages within the perianth-bearing Piperales. In addition, the hypothesis that the vegetative organs of the holoparasitic Hydnoraceae are most probably rhizomes is confirmed.     

STEM ANATOMY OF PARTHENIUM ARGENTATUM,P. INCANUM AND THEIR NATURAL HYBRIDS

Guayule (Parthenium argentatum Gray) contains rubber in the parenchymatous cells of stems and roots. Stem anatomy of P. argentatum is described along with that of P. incanum H.B.K. (mariola). Anatomy of these species differs significantly. Phloem rays in both species increase in width by cell division and expansion; however, the increase observed in mariola is less as compared to that in guayule. Axial xylem parenchyma in guayule is generally a two-cell strand as compared to the fusiform axial xylem parenchyma observed in mariola. Vascular ray cells and cells of the pith region of guayule are parenchymatous, whereas those of mariola are sclerenchymatous. As a result of introgression between guayule and mariola, three forms of guayule exist in the native stands of Mexico. Morphological differences between these guayule plants have been described previously. The stem anatomy of these three groups of plants differ importantly. Group I guayule plants, least introgressed by mariola, have taller rays with the cells of pith region and vascular rays parenchymatous. Group III plants, highly introgressed by mariola, have a few to many cells of vascular rays and pith with lignified secondary walls and shorter rays. Many of the anatomical characteristics of group II plants, somewhat introgressed by mariola, are intermediate between group I and III plants.     

Comparative leaf anatomy, FTIR discrimination and biogeographical analysis of Camellia section Tuberculata (Theaceae) with a discussion of its taxonomic treatments

A comparative anatomical and FTIR study on the leaves of 18 disputed species of section Tuberculata (Camellia, Theaceae) combined with their biogeographical distribution have been conducted in order to investigate interspecific variations which are useful in species taxonomic treatment and systematic evolution. All Tuberculata species have bifacial and hypostomatic leaves, contain sclereids and crystals. Multivariate analysis of leaf FTIR data shows a visualization of the degree of affinity among the species in this section, which is consistent with the distribution of synonymous Tuberculata species indicated in the biogeographical maps. Our study indicates that the combination of characters based on leaf comparative anatomy, FTIR analysis and biogeographical distribution are useful in species revision. The results determine 11 species in section Tuberculata and provide the evolutionary trends in Tuberculata, which originated in three-river region of Guizhou province and spread to Guizhou, Guangxi, Yunnan, Sichuan, Chongqing, Hunan and probably Hubei provinces of China.     

ONTOGENY AND DIFFERENTIATION OF SCLEREIDS IN RAUWOLFIA

An interesting anatomic feature of Rauwolfia is the occurrence of a remarkable type of sclereid in the stem and root. The initials of the sclereids in the stem arise in the ground tissue element of the pith in a region between 50 and 70μ below the surface of the shoot apex. This region of the shoot remains surrounded by a whorl of either 3 or 4 leaves. Sclereids initiate in succession in association with each whorl of leaves. Thus, the sclereids are restricted to the nodes. The sclereids in the stem arise as a primary element of the shoot from the ground tissue of the pith. In the root, they differentiate from the cells of the phelloderm and are secondary in origin. Morphologically, the sclereids in these 2 organs are basically the same, except that the sclereids in the stem are larger in size than those in the root. A solitary cell, or 2 to several cells in a longitudinal cell file (originated from a single mother cell), may differentiate into sclereid initials. The growth of the sclereids through relatively compact ground tissue of the pith is possibly accomplished by a combination of gliding growth and apical intrusive process. The sclereid initials grow rapidly and force their way between the parenchymatous cells. As a result, the neighboring cells lose their original surface contacts. Sclereids increase in size rapidly, and, therefore, very enlarged sclereids with thin primary walls may be observed in the second node. They mature progressively in basipetal direction in the subjacent nodes. In the fifth or sixth node, mature sclereids with massive secondary walls are most common. The secondary walls of sclereids contain much lignin as determined by the phloroglucinol-HCl test. The walls of sclereids at maturity show a variable number of lamellae ranging from 10 to 15 in the lateral walls. A remarkable feature of the sclereids is their canal-like pits in the secondary walls. Two adjacent pits may coalesce uniquely to form a Y-like configuration directed centrifugally from the lumen of the sclereids. The sclereids are ventrically symmetrical, joined end-to-end by their transverse walls like 2 superimposed young fibers.     

Stem anatomy of Rhipsalis (Cactaceae) and its relevance for taxonomy

Several species of the genus Rhipsalis (Cactaceae) are extremely important as ornamentals and are endangered in their natural habitat. However, only a few studies have addressed its taxonomy, morphology (including anatomy), phylogeny and evolutionary history. Consequently, the limited knowledge of the genus coupled with the problematic delimitation of species had led to problems in the identification of taxa. In the current work six species of Rhipsalis, R. cereoides, R. elliptica, R. grandiflora, R. paradoxa, R. pentaptera and R. teres were studied to evaluate the relevance of anatomical characters for the taxonomy of the genus. An anatomical characterization of the primary structure of the stem of Rhipsalis is provided highlighting the differences between species. Features of the stem epidermis are found to discriminate best between species and therefore provide clear and useful characters for the separation of species.     

Anatomy,trichome morphology and palynology of Salvia chrysophylla Stapf (Lamiaceae)

The anatomy, palynology, morphology and distribution of the trichomes on the aerial parts of Salvia chrysophylla Stapf, an endemic species in Turkey, were studied in order to understand the usefulness of these characteristics for systematic purposes. Some anatomical characters such as (1–)2–24-rowed pith rays in roots, dorsiventral leaves, obviously larger upper epidermal cells, and two to three large vascular bundles in the center and two to four small subsidiary bundles in the wings of petiole provide information of taxonomical significance. Three main types of trichomes were observed on the stem, inflorescence axis, leaf and calyx surfaces of S. chrysophylla. They are peltate, capitate glandular and non-glandular. Capitate glandular and non-glandular trichomes were further subdivided into several kinds. Glandular trichomes are present in abundance on the inflorescence axis and calyx, but non-glandular ones were mainly situated on the leaf and stem. Scanning Electron Microscopy (SEM) studies on the pollen grains have revealed that they are oblate-spheroidal and their exine ornamentation is bireticulate-perforate.     

Stem anatomy of neotropical <Emphasis Type="Italic">Dioscorea</Emphasis> L. (Dioscoreaceae) and its importance to the systematics of the genus

We anatomically characterized the aerial stems of the genus Dioscorea and evaluated the use of these anatomical characteristics to better understand the taxonomy, systematics, and diversity of the component species in the neotropics. Fragments of the aerial stems of 23 species were collected for anatomical analyses utilizing conventional cytohistological techniques. Stem anatomy did not support the subgenera and neotropical sections of Dioscorea, although D. sect. Monadelpha demonstrated similar anatomical characteristics with the representatives analyzed in the present work, supporting the possible monophyletism of that section. Some New World species demonstrated characteristics described earlier for Old World species, such as two phloem units internal to the large-diameter vessels. New characters to the vascularization of new world species are described, as a single ring of vascular bundles, intercalating common and cauline bundles. Tylosoids and actinocytic stomata were the first time reported in Dioscorea, and the vascular patterns are revisited with a systematic approach.     

Wood and bark anatomy of lianoid Indomalesian and Asiatic species of Gnetum

Quantitative and qualitative data on wood and bark are offered for 11 species of lianoid Indomalesian and Asiatic species of Gnetum section Cylindrostachys. Material of roots was studied for two species, material of an underground stem for one species, and stem material was studied for all species; wood from inside and outside of a large stem of G. montanum was analysed (quantitative data do not change with age in this species). Roots have shorter, narrower vessel elements, more numerous per mm², compared with those of stems; these trends run counter to those in dicotyledons. Roots and underground stems have more abundant parenchyma and less abundant sclerenchyma than do stems. Parenchyma of both roots and stems is rich in starch. All of the species studied here have both fibre-tracheids and tracheids, but tracheids are not distributed vasicentrically as they are in dicotyledons. Tori are reported for tracheary elements of three species studied here. Vasicentric axial parenchyma (which usually is thick-walled) is present in all species; thick or thin-walled diffuse or diffuse-in-aggregates apotracheal parenchyma is present in almost all of the species studied. Rays are mostly dimorphic in size, but show various conditions with respect to wall thickness, sclerenchyma presence, and crystal presence. As in other lianoid species of Gnetum, the species of the present study show origin of lateral meristem activity in parenchyma of the innermost cortex. Cortex and bark of the species studied here are relatively uniform in distribution of gelatinous fibres, nests of sclereids, the cylinder of brachysclereids that extends around the stem, and sclerenchymatous phelloderm. Laticifers were observed in bark of only two species studied. Although a few species characters are evident, the species that comprise Section cylindrostachys differ from each other mostly in degree rather than in presence or absence character state distributions. Secretory cavities are newly reported for the genus.     

Stem and root anatomical correlations with life form diversity, ecology, and systematics in Moringa (Moringaceae)

Four life forms (habits) are identified in the 13 species of Moringa (bottle trees, sarcorhizal trees, slender trees, and tuberous shrubs) which are examined for wood anatomical correlations with habit, ecology, and systematic. Wood anatomy is similar within habit classes except for the sarcorhizal trees. The four bottle tree species and M. arborea (one of the sarcorhizal trees) are characterized by bands of confluent paratracheal parenchyma alternating with bands of libriform fibres, some of which may be parenchyma-like. The other sarcorhizal tree, M. ruspoliana , is characterized by alternating bands of parenchyma-like and long, slender libriform fibres. Root secondary xylem of all these species is characterized by bands of parenchyma and fibres. Slender trees do not show bands of fibres of different shapes and have fibrous roots with less parenchyma than the other species. Tuberous shrubs have stems mostly composed of long, slender fibres and large underground tubers mostly composed of parenchyma. Quantitative trends between ecologically different localities include wider vessel elements and higher conductive area in moister localities. Wood anatomy provides characters that are of potential phylogenetic utility at a variety of levels of relationship. Based on wood anatomy and geography, the most likely sister taxon to Moringa is Cylicomorpha (Caricaceae).     

Caryophyllales: a key group for understanding wood anatomy character states and their evolution

Definitions of character states in woods are softer than generally assumed, and more complex for workers to interpret. Only by a constant effort to transcend the limitations of glossaries can a more than partial understanding of wood anatomy and its evolution be achieved. The need for such an effort is most evident in a major group with sufficient wood diversity to demonstrate numerous problems in wood anatomical features. Caryophyllales s.l., with approximately 12 000 species, are such a group. Paradoxically, Caryophyllales offer many more interpretive problems than other ‘typically woody’ eudicot clades of comparable size: a wider range of wood structural patterns is represented in the order. An account of character expression diversity is presented for major wood characters of Caryophyllales. These characters include successive cambia (more extensively represented in Caryophyllales than elsewhere in angiosperms); vessel element perforation plates (non‐bordered and bordered, with and without constrictions); lateral wall pitting of vessels (notably pseudoscalariform patterns); vesturing and sculpturing on vessel walls; grouping of vessels; nature of tracheids and fibre‐tracheids, storying in libriform fibres, types of axial parenchyma, ray anatomy and shifts in ray ontogeny; juvenilism in rays; raylessness; occurrence of idioblasts; occurrence of a new cell type (ancistrocladan cells); correlations of raylessness with scattered bundle occurrence and other anatomical discoveries newly described and/or understood through the use of scanning electron microscopy and light microscopy. This study goes beyond summarizing or reportage and attempts interpretations in terms of shifts in degrees of juvenilism, diversification in habit, ecological occupancy strategies (with special attention to succulence) and phylogenetic change. Phylogenetic change in wood anatomy is held to be best interpreted when accompanied by an understanding of wood ontogeny, species ecology, species habit and taxonomic context. Wood anatomy of Caryophyllales demonstrates problems inherent in binary character definitions, mapping of morphological characters onto DNA‐based trees and attempts to analyse wood structure without taking into account ecological and habital features. The difficulties of bridging wood anatomy with physiology and ecology are briefly reviewed. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 342–393.     

Comparative anatomy of selected rhizomatous and tuberous taxa of subfamilies Orchidoideae and Epidendroideae (Orchidaceae) as an aid to identification

This article addresses the vegetative anatomy (leaves, stems, roots, root tubers and rhizomes) of 13 species of subfamily Orchidoideae (Orchidaceae), belonging to the genera Neottia Guettard, Cephalanthera L.C.M. Richard, Epipactis Zinn, Limodorum Boehmer, Spiranthes L.C.M. Richard, Platanthera L.C.M. Richard, Serapias L., Himantoglossum W.D. Koch and Anacamptis L.C.M. Richard, because anatomical studies have provided very useful criteria for orchid diagnosis. In the study three types of painting methods—Delafield’s hematoxylin and safranin, Alcian blue-periodic acid schiff, and alcoholic phloroglucinol?+?HCl—were employed, and identification tables were prepared. Anatomical results demonstrated the differences in the leaf anatomy of tuberous and rhizomatous orchids. In the stem anatomy, all the rhizomatous genera were found to be anatomically different, especially in regard to the collateral vascular bundles, the distribution of vascular bundles and xylem properties. In root anatomy, the central cylinder, pith, endodermis and/or pericycle properties are distinctive features in all studied taxa. For root tubers, velamen layering, wall outline mucilage cell patterns in ground tissue and arrangements of vascular arches can be used to label taxa. Regarding the rhizome anatomy of the studied taxa, vascular cylinder results in particular were very significant for the distinction of genera. Finally, we strongly emphasize the importance of this kind of detailed anatomical study to solve identification problems of orchid taxonomy.