Comparative vegetative anatomy and systematics of Laeliinae (Orchidaceae)

Laeliinae are one of the most prominent orchid subtribes, with c. 40 genera and nearly 1500 species, and contain a disparate group of taxa with widely varying morphological features. There does not appear to be a complex of characters to which one can refer in order to delineate the subtribe as a whole. Thus, it was thought that vegetative anatomy might provide clues to the monophyly of the group. The microscopic structure of the leaves, stems and roots of representatives of most of the genera was studied. It was concluded that the anatomy lacks overall uniformity and that vegetative characters alone are insufficient to assess the relationships amongst the genera. The only nearly consistent anatomical feature was the abaxial row of fibre bundles in the leaves. Thus, anatomically, as well as morphologically, Laeliinae are a mixed bag. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 21–41.     

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.     

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.     

Comparative vegetative anatomy and systematics of the Oncidiinae (Maxillarieae, Orchidaceae)

Subtribe Oncidiinae comprises a vegetatively heterogeneous assemblage of species that has persistently been incapable of organization. Anatomy was considered to be a possible means to resolve the perplexity of relationships amongst the constituent taxa. The consistent occurrence of a foliar hypodermis, homogeneous mesophyll, conical silica bodies in stegmata, and ubiquitous fibre bundles in leaves provides a matrix for linking the taxa, as do the parenchymatous pith and O-thickened endodermal cell walls in roots. However, the strict consensus of the 40 genera studied was completely unresolved, suggesting that vegetative characters alone are insufficient to assess the relationships amongst these taxa, a conclusion also reached for the remainder of Maxillarieae.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 91–107.     

Comparative vegetative anatomy and systematics of Vanilla (Orchidaceae)

Vanilla is a pantropical genus of green-stemmed vines bearing clasping (aerial) and absorbing (terrestrial) roots. Most vanillas bear normal, thick foliage leaves; others produce fugacious bracts. Seventeen species, including both types were studied. Foliage leaves of Vanilla are glabrous, have abaxial, tetracytic stomatal apparatuses, and a homogeneous mesophyll. Species may or may not have a uniseriate hypodermis. Crystals occur in the foliar epidermises of some species, but all species have crystalliferous idioblasts with raphides in the mesophyll. Vascular bundles in leaves are collateral and occur in a single series alternating large and small. Sclerenchyma may or may not be associated with the vascular bundles. Scale leaves may be crescent or C-shaped and usually have abaxial stomatal apparatuses. A hypodermis may or may not be present; the mesophyll contains raphide bundles in idioblasts. Vascular bundles are collateral and occur in a single row sometimes aligned close to the adaxial surface. They may or may not be associated with sclerenchyma. Stems of leafy vanillas show a sclerenchyma band separating cortex from ground tissue; stems of leafless vanillas do not show a sclerenchyma band. Ground tissue of the stem may consist solely of assimilatory cells or mixed assimilatory and water-storage cells. In some species centrally located assimilatory cells are surrounded by layers of water-storage cells. A uniseriate hypodermis is present in all stems. Sclerenchyma may completely surround the scattered collateral vascular bundles, occur only on the phloem side, or be absent. Both aerial and terrestrial roots are notable for their uniseriate velamen the cell walls of which may be unmarked or ornamented with anticlinal strips. Exodermis is uniseriate; the cells vary from barely thickened to strongly thickened. Only the outer and radial walls are thickened. Cortical cells of aerial roots generally have chloroplasts that are lacking from the same tissue of terrestrial roots. Raphide bundles occur in thin-walled cortical idioblasts. Endodermis and pericycle are uniseriate; pericycle cells are all ?-thickened opposite the phloem. Cells of the endodermis are either ?- or ∪-thickened opposite the phloem. Vascular tissue may be embedded in thin- or thick-walled sclerenchyma or in parenchyma. Metaxylem cells are always wider in terrestrial than in aerial roots of the same species. Pith cells are generally parenchymatous but sclerotic in a few species.     

Comparative vegetative anatomy and systematics of Cymbidium (Cymbidieae: Orchidaceae)

The genus Cymbidium (Orchidaceae) exhibits distinctive ecological diversification and occurs in terrestrial, epiphytic, and lithophytic life forms. One species, Cymbidium macrorhizon , lacks foliage leaves and has a strongly mycoparasitic existence. Correlation between habitat differentiation and anatomical characters was tested for 21 species of Cymbidium and its putative sister groups. Although hypostomaty characterizes the genus, C. canaliculatum shows amphistomaty. Ecological preference of this species indicates that amphistomaty is likely adapted to intensive insolation. Four types of subepidermal foliar sclerenchyma were found. Two forest floor species, C. goeringii and C. lancifolium as well as the mycoparasitic C. macrorhizon , do not have this sclerenchyma. In this genus, development of sclerenchyma is correlated with the degree of epiphytism. Palisade mesophyll evolved in Cymbidium section Cymbidium . As members of this section grow on isolated trees in tropical lowland forests or on rocks, the differentiation of palisade tissue is probably correlated with immigration to high light habitats. With the exception of C. macrorhizon , stegmata were found in leaves and stems of Cymbidium . Furthermore, a few epiphytic species have stegmata in their roots; this is a curious feature rarely found in vascular plants. Subterranean rhizomes characterize terrestrial species, while ageotropic roots are found in some epiphytic species. Cymbidium macrorhizon shows peculiar features such as degeneration of stomata, anomocytic stomata, and lack of stegmata and sclerenchyma. This set of character transformations is probably correlated with the evolution of mycoparasitic existence. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 383–419.     

Comparative vegetative anatomy of Stanhopeinae (Orchidaceae)

Stanhopeinae are a group of tropical American orchids characterized by euglossine bee pollination and lateral inflorescences stemming from the bases of pseudobulbs. Leaves are hypostomatal, and all stomatal configurations are tetracytic. Chlorenchyma is homogeneous and characterized by fibre bundles in adaxial/abaxial or adaxial/median/abaxial positions. Collateral vascular bundles occur in a single row and feature phloic and xylic sclerenchymatous caps and thin-walled bundle sheath cells. Fibre bundles and vascular sclerenchyma are accompanied by stegmata containing conical silica bodies. Pseudobulbs have thick-walled turbinate epidermal cells and ground tissue of smaller, living assimilatory cells and larger, dead water-storage cells. Fibre bundles are usually absent but occur in several genera. Collateral vascular bundles show phloic sclerenchyma, but xylic sclerenchyma occurs only in die larger vascular bundles. Phloic and xylic sclerenchyma are associated with stegmata containing conical silica bodies. Roots are velamentous. Velamen cell walls have fine, spiral thickenings. Exodermal cells are thin-walled. The cortex features scattered thick-walled cells and in some cases branched bars of secondary cell wall material. Endodermis is either u-or O-thickened, but pericycle is always O-thickened opposite the phloem. Vascular tissue consists of alternating strands of xylem and phloem surrounded by a matrix of thick-walled cells. Pith cells may be parenchymatous or sclerenchymatous.     

Comparative anatomy and systematics of Catasetinae (Orchidaceae)

Catasetinae consist of five genera of pseudobulbous Orchidaceae of the Neotropics. Anatomy is characterized by sunken, three-celled foliar hairs, mostly tetracytic stomatal apparatuses, superficial stomata, homogeneous mesophyll, foliar fibre bundles, collateral vascular bundles in a single row, xylem and phloem sclerenchyma associated with vascular bundles in leaves, conical, and rough-surfaced silica bodies adjacent to vascular bundle sclerenchyma; epidermal cells of pseudobulbs with heavily thickened outer walls, pseudobulb ground tissue of assimilatory and water-storage cells, scattered vascular bundles in pseudobulbs, and sclerenchyma and stegmata associated only with phloem of pseudobulbs; roots with thin-walled velamen cells and tenuous spirals of cell wall material, distinctive epivelamen cells, thin-walled exodermal cells and vascular tissue embedded in parenchyma. Except for mucilaginous idioblasts that occur in Mormodes and Cycnoches , there are few outstanding anatomical differences among the five genera. Thus, there are few anatomical characteristics of phylogenetic value. The monophyly of Catasetinae is supported by the presence of sunken foliar hairs. Our results support a close relationship between Clowesia and Catasetum , and between Mormodes and Cycnoches. Among the outgroups Pteroglossaspis is especially distinctive.     

Comparative anatomy and systematics of Senghas's cushion species of Maxillaria (Orchidaceae)

Using data obtained through anatomy and morphology, we used cladistics to examine the monophyly of Senghas's proposed classification of Maxillaria cushion plants and his placement of Mormolyca ringens. Trignidium obtusum was chosen as the outgroup. Leaves have multicellular hairs sunken in crypts, primarily anomocytic or primarily tetracytic stomatal apparatuses, homogeneous mesophyll, and scattered fibre bundles. Three types of adaxial hypodermis were observed: (1) water-storage cells, (2) fibre bundles scattered among water-storage cells, and (3) fibre bundles scattered among chlorenchymatous cells. Abaxial hypodermis of fibre bundles occurs in several Maxillaria species and in Trigonidium obtusum. At the midvein of the leaf, adaxial mesophyll cells of most species are anticlinally extended and empty, and the abaxial mesophyll is usually collenchymatous. Vascular bundles are collateral and usually in a single series. Pseudobulb epidermal cell walls are thin, or outer walls are thickened. Ground tissue consists of water-storage and assimilatory cells with vascular bundles and associated lacunae scattered throughout. Roots are velamentous and exodermal cell walls are usually n-thickened with tenuous bands of scalarifom thickenings on longitudinal walls. Tilosomes may be plaited, baculate, or spongy. Endodermal cell walls are usually U-thickened and pericycle cell walls are usually O-thickened opposite phloem sectors. Stegmata line the periphery of the thickened pericycle cells opposite phloem sectors in M. picta. Pith may be parenchymatous or sclerenchymatous. According to our phylogenetic analysis, Mormolyca ringens is consistently nested within the cladistic structure of Maxillaria. Therefore, Maxillaria likely is paraphyletic if Mormolyca ringens is recognized as generically distinct. It appears that Senghas's subgroup divisions of the unifoliate pseudobulbous maxillarias may also be artificial.     

Comparative vegetative anatomy and systematics of the angraecoids (Vandeae, Orchidaceae) with an emphasis on the leafless habit

The vegetative anatomy and morphology of 142 species of the angraecoid orchids (Angraecinae + Aerangidinae) and 18 species of Aeridinae were examined using light and scanning electron microscopy. Leafless members of Vandeae were of particular interest because of their unique growth habit. Leafy and leafless members of Angraecinae and Aerangidinae were examined and compared with specimens of Aeridinae. Vandeae were homogeneous in both leaf and root anatomy. A foliar hypodermis and fibre bundles were generally absent. Stegmata with spherical silica bodies were found associated with sclerenchyma and restricted to leaves in almost all specimens examined. Distinct inner tangential wall thickenings of the endovelamen occurred in several vandaceous genera. Exodermal proliferations and aeration units commonly occurred in both leafy and leafless Vandeae. Cladistic analyses of Angraecinae and Aerangidinae with members of Aeridinae and Polystachyinae as outgroups using 26 structural characters resulted in 20 000+ equally parsimonious trees. Vandeae formed the only well-supported clade in bootstrap analyses and were characterized by having a monopodial growth habit, spherical stegmata, loss of mucilage, and loss of tilosomes.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 165–218.     

Vegetative anatomy and systematics of Triphorinae (Orchidaceae)

Triphorinae represents a group of three anatomically simple genera, the structural features of which are unspecialized. The anomocytic stomatal pattern occurs in all genera; it predominates in Triphora. A foliar hypodermis, sclerenchyma, fibre bundles and stegmata are absent. The mesophyll is homogeneous. The exodermal and endodermal cells in the roots are entirely thin‐walled and tilosomes are absent. However, there are anatomical modifications that appear to be unique: root hairs in Monophyllorchis are borne on velamenal buttresses and, in Psilochilus, they arise endogenously. In the root vascular system of Psilochilus, the metaxylem occurs as a circumferential band. The surfaces of stems in Triphora are tuberculate. Mycorrhizae appear to characterize the root cortices of all genera. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 203–210.     

Vegetative anatomy and systematics of subtribe Dendrobiinae (Orchidaceae)

Anatomy of leaf, stem, and root of more than 100 species in subtribe Dendrobiinae (Orchidaceae) was studied with the light microscope to provide a comparative anatomical treatment of these organs, to serve as an independent source of evidence that might be taxonomically important, and to recommend such reinterpretations of existing classifications as are suggested by a phylogenetic assessment of data. We based our classification on that of Rudolf Schlechter as the most complete and widely accepted today. We found that the anatomy of plants in subtribe Dendrobiinae reflects a high degree of morphological diversity, and many of the anatomical characters appear to be homoplasous. When these anatomical data are used to interpret the systematic relationships among the genera, they indicate that Dendrobium is not monophyletic and that Cadetia and Pseuderia are apparently nested within the structure of Dendrobium when section Grastidium is chosen as a functional outgroup. Lack of resolution in the strict consensus tree illustrates the difficulty of determining the phylogenetic relationships of many of Schlechter's sections using anatomical characters. Nevertheless, we recommend that his sectional classification, with appropriate modifications based on available data, be retained for the present, pending a more detailed understanding of the phylogeny of Dendrobiinae based on morphology, micromorphology, anatomy, and DNA studies.     

Systematic and comparative anatomy of Maxillarieae (Orchidaceae), sans Oncidiinae

On the basis of floral and vegetative morphology, 63 tropical American genera have been recognized within Maxillarieae. We were able to examine anatomical material of all subtribes, excluding Oncidiinae. Stegmata with conical silica bodies occur in leaves and stems of all subtribes excluding Ornithocephalinae, and pericyclic stegmata found in roots are characteristic of Lycastinae. Lycastinae and Maxillariinae are characterized by foliar glands, foliar fibre bundles and tilosomes. Endodermal cells are U-thickened in most Zygopetalinae; O-thickened in most Lycastinae, Ornithocephalinae and Telipogoninae; variously thickened in Maxillariinae; and thin-walled in Cryptarrhena lunata . Water-storage cells varied from thin-walled to variously banded throughout Maxillarieae. Cladistic analyses using anatomical characters yielded no resolution among subtribes, illustrating that anatomical characters are of limited value in assessing relationships within this tribe.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 251–274.     

Root character evolution and systematics in Cranichidinae, Prescottiinae and Spiranthinae (Orchidaceae, Cranichideae)

Background and Aims: Previous studies have suggested that velamen characteristicsare useful as taxonomic markers in Orchidaceae. Members of tribeCranichideae have been assigned to two velamen types constructedbased on combinations of characters such as the presence ofsecondary cell-wall thickenings and pores. However, such charactershave not been analysed on an individual basis in explicit cladisticanalyses. Methods: The micromorphology of roots of 26 species of Cranichideae wasexamined through scanning electron microscopy and light microscopy,scoring the variation and distribution of four characters: numberof velamen cell layers, velamen cell-wall thickenings, presenceand type of tilosomes, and supraendodermal spaces. The lastthree characters were analysed cladistically in combinationwith DNA sequence data of plastid trnK/matK and nuclear ribosomalinternal transcribed spacer (ITS) regions and optimized on theresulting phylogenetic tree. Key Results: Thickenings of velamen cell walls group Prescottiinae with Spiranthinae,whereas tilosomes, documented here for the first time in Cranichideae,provide an unambiguous synapomorphy for subtribe Spiranthinae.Supraendodermal spaces occur mostly in species dwelling in seasonallydry habitats and appear to have evolved three times. Conclusions: Three of the four structural characters assessed are phylogeneticallyinformative, marking monophyletic groups recovered in the combinedmolecular–morphological analysis. This study highlightsthe need for conducting character-based structural studies toovercome analytical shortcomings of the typological approach.     

Amyloplasts from cortical root cells of Spiranthoideae (Orchidaceae)

Summary Cortical root cells of orchids belonging to subfamily Spiranthoideae contain globular organelles which, through chemical tests and examination with both light, scanning, and transmission electron microscopes, appear to be unique in Orchidaceae and unreported in plants in general. It is suggested these are specialized amyloplasts (spiranthosomes) and with other features may serve to characterize the spiranthoid orchid subfamily.Dedicated to Hilton H. Mollenhauer on the occasion of his retirement     

Observations of the floral biology ofPrasophyllum odoratum (Orchidaceae,Spiranthoideae)

Prasophyllum odoratum is a vernal, nectariferous, terrestrial orchid that flowers profusely six to eight months following cyclical fires that disrupt sclerophyll woodlands. The morphology of the column and pollinarium is indicative of taxa placed within the subfam.Spiranthoideae. The orientation of the pollinaria to the stigma appears to prevent mechanical self-pollination. Both cross- and self-pollination appear to be effected by insects that forage within the brightly-colored, scented, non-resupinate flowers. Ants and drosophilid flies remove nectar, but do not appear to transport pollinaria between flowers. The primary pollinators are polytrophic flies in the fam.Syrphidae and opportunistic male bees in the genusLeioproctus (Colletidae). Approximately 52% of the flowers on a raceme set seed. The comparatively short floral tube ofP. odoratum reflects the dependence of this species on short-tongued insects to effect successful dispersal of pollinaria.     

Two new species of Aa (Orchidaceae,Spiranthoideae) from Colombia

Two new species of the Andean genus Aa (Orchidaceae, Spiranthoideae) are described: Aa lozanoi Szlach. and S. Nowak, and Aa figueroi Szlach. and S. Nowak. They are restricted in distribution mainly to Cordillera Oriental in the department of Cundinamarca, however, A. lozanoi was also collected in Cordillera Central and A. figueroi in Sierra Nevada de Santa Marta, in northern part of Colombia. Each species is described and illustrated, detailed habitat and distribution data are provided. A distribution map of the new species is presented. A dichotomous key for determination of the Colombian species of Aa is provided. Brief discussion about the most important threats for plants in Andes is presented.     

Three new species of Ponthieva (Orchidaceae, Spiranthoideae) from Colombia and Venezuela

Three new species of Ponthieva, P. camargoi, P. castanedoi, and P. cesarensis are described and illustrated based on Colombian and Venezuelan material. The comparisons of the floral characters of the new entities and closely allied species are illustrated. The information about habitat and ecology of all three species is provided and their distribution map is presented.