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 Spiranthoideae (Orchidaceae)

STERN, W. L., MORRIS, M. W., JUDD, W. S., PRIDGEON, A. M. & DRESSLER, R. L. 1993. Comparative vegetative anatomy and systematics of Spiranthoideae (Orchidaceae). The anatomy of leaf, stem and root of plants in the orchid subfamily Spiranthoideae was studied and described from the viewpoint of systematics. Plants were available from most of the geographic range. Tribes Diceratosteleae and Tropidieae show sinuous anticlinal epidermal cell walls in leaves, glandular foliar hairs, tetracytic para-mesoperigenous stomata, unitary tracheary components in the foliar midrib, foliar and cauline stegmata and sclerenchyma, typical cruciate starch grains, thick-walled exodermal, endodermal, and pericyclic cells, and conductive strands of the root embedded in sclerenchyma. The tribe Cranichideae shows straight to curvilinear anticlinal epidermal cell walls in leaves, lack glandular foliar hairs, have variably patterned mesoperigenous stomata, lack sclerenchyma throughout the parts studied, have a binary tracheary component in the foliar midrib, store starch in specialized amyloplasts (spiranthosomes), lack stegmata, have thin-walled exodermal, endodermal, and pericyclic cells, show scalariform thickenings in exodermal cells, and have conductive strands of the root embedded in parenchyma. In Cryptostylis the tracheary component of the foliar midrib is unitary, stomata lack subsidiary cells, starch grains are of the typical cruciate configuration, and passage cells of the endodermis are apparently associated with tilosomes. Anatomical data, when analysed cladistically, support the hypothesis that Spiranthoideae, as currently delimited, are polyphyletic. Corymborkis, Tropidia, and Diceratostele are more closely related to Palmorchis, a likely representative of a basal clade within subfamily Epidendroideae, than to genera of Cranichideae. Likewise, members of Cranichideae are more closely related to Diuris, a representative of subfamily Orchidoideae-tribe Diurideae, than to Corymborkis, Tropidia and Diceratostele. The Corymborkis– Tropidia-Diceratostele-bassd epidendroid [Palmorchis) clade may be diagnosed by the foliar synapomorphies of sinuous anticlinal walls of epidermal cells and presence of glandular hairs. The Cranichideae-orchidoid (Diuris) clade may be diagnosed by its variably patterned, mesoperigenous stomata, lack of vascular bundle sclerenchyma, absence of stegmata, unthickened endodermal cell walls in roots, and conductive cells of roots embedded in parenchyma. Relationships within this clade are quite unresolved, when only anatomical data are employed; however, all studied genera of Cranichideae, except Cryptostylis, possess a binary tracheary component in the foliar midrib. Cranichideae, excluding Cryptostylis, may be considered monophyletic. All Cranichideae, except Helaeria and Cryptostylis, possess spiranthosomes. Hetaeria may be a basal member within Cranichideae. We consider the phylogenetic position of Cryptostylis, in relation to Cranichideae vs. Diurideae, to be equivocal.     

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 anatomy of the nectary spur in selected species of Aeridinae (Orchidaceae)

Background and Aims

To date, the structure of the nectary spur of Aeridinae has not been studied in detail, and data relating to the nectaries of ornithophilous orchids remain scarce. The present paper compares the structural organization of the floral nectary in a range of Aeridinae species, including both entomophilous and ornithophilous taxa.

Methods

Nectary spurs of Ascocentrum ampullaceum (Roxb.) Schltr. var. aurantiacum Pradhan, A. curvifolium (Lindl.) Schltr., A. garayi Christenson, Papilionanthe vandarum (Rchb.f.) Garay, Schoenorchis gemmata (Lindl.) J.J. Sm., Sedirea japonica (Rchb.f.) Garay & H.R. Sweet and Stereochilus dalatensis (Guillaumin) Garay were examined by means of light microscopy, scanning electron microscopy and transmission electron microscopy.

Key Results and Conclusions

The diverse anatomy of the nectary is described for a range of Aeridinae species. All species of Ascocentrum investigated displayed features characteristic of ornithophilous taxa. They have weakly zygomorphic, scentless, red or orange flowers, display diurnal anthesis, possess cryptic anther caps and produce nectar that is secluded in a relatively massive nectary spur. Unicellular, secretory hairs line the lumen at the middle part of the spur. Generally, however, with the exception of Papilionanthe vandarum, the nectary spurs of all entomophilous species studied here (Schoenorchis gemmata, Sedirea japonica, Stereochilus dalatensis) lack secretory trichomes. Moreover, collenchymatous secretory tissue, present only in the nectary spur of Asiatic Ascocentrum species, closely resembles that found in nectaries of certain Neotropical species that are hummingbird-pollinated and assigned to subtribes Maxillariinae Benth., Laeliinae Benth. and Oncidiinae Benth. This similarity in anatomical organization of the nectary, regardless of geographical distribution and phylogeny, indicates convergence.     

Vegetative anatomy of subtribe Habenariinae (Orchidaceae)

Leaves of Habenariinae are characterized by anomocytic stomatal apparatuses, homogeneous mesophyll, collateral vascular bundles in a single series, and thin-walled bundle sheath cells. There is no foliar sclerenchyma nor a hypodermis. Cauline cortex consists of thin-walled living cells among which are large and numerous intercellular spaces. The ground tissue is bordered externally by a layer of thick-walled living cells, except in Habenaria repens. Central ground tissue cells are living, and usually thin-walled surrounding intercellular spaces of various dimensions. These are conspicuously large in H. repens. Collateral vascular bundles are scattered across the ground tissue. Sclerenchyma is absent. Absorbing roots are generally velamentous, exodermal dead cells are diin-walled, and passage cells usually have a thickened outer wall. A regular vascular cylinder is present, and vascular tissue is embedded in parenchyma. Root tubers are velamentous, exodermal cells are usually thin-walled, and passage cells frequently have thickened outer walls. Vascular tissue of root tubers is organized into two classes: (1) those with a single vascular cylinder surrounded by a cortex and (2) those with a series of meristeles dispersed throughout the ground tissue. In group (1) cortex is homogeneous either with or without mucilage cells except in Stenoglattis where the cortex is heterogeneous, consisting of water-storage and assimilatory cells, and lacks mucilage cells. In group (2) the ground tissue consists of larger mucilage-containing cells and smaller assimilatory cells.     

Systematic leaf anatomy of Caladeniinae (Orchidaceae)

Leaf anatomy of 25 species in 15 genera of Caladeniinae (Diurideae, Orchidaceae), excluding Caladenia, was investigated to determine diagnostic characters to be used in forthcoming, broad-based cladistic analyses of the subtribe and to assess interspecific and intergeneric relationshipS. Of the characters examined, those that show the most variation among the study taxa are presence and types of trichomes, cuticular sculpturing, anticlinal walls of epidermal cells, heterogeneity of chlorenchyma, distribution and length: width ratios of stomata. Anatomical evidence supports the generic concept of Leptoceras Lindley but contradicts that of Drakonorchis Hopper & A.P. Brown. Paracaleana is not sufficiently distinct from Caleana to warrant generic status. Lyperanthus serratus and L.suaveolens are hyperstomatic, a rare condition in Orchidaceae. On the basis of these and other characters, Lyperanthus, as currently circumscribed, is polyphyletic. From leaf structure Caladeniinae as now conceived is polyphyletic and comprises seven groups: (1) Caladenia, Leptoceras, Elythranthera, Eriochilus, Glossodia, Aporostylis; (2) Adenochilus, Rimacola; (3) Arthrochilus, Chiloglottis, Spiculaea, Leporella; (4) Caleana (including Paracaleana); (5) Bumettia; (6) Lyperanthus suaveolens and L.serratus; (7) Lyperanthus nigricans.     

The role of vegetative and reproductive attributes in the classification of the Orchidaceae

Ninety-three orchid taxa have been arranged into three classificatory dendrograms employing respectively 40 reproductive, 34 vegetative and 74 attributes combined, and forming the hierarchy by means of group-average clustering. According to these results the non-specificity hypothesis does not hold well for vegetative versus reproductive attributes. Both types of attributes are equally efficient predictors of the classification based upon the total data, though neither data set is a very reliable predictor. Hence both should be employed in the production of general classifications and when considering the phylogenetic relationships amongst orchids.     

Comparative leaf anatomy and phylogenetic relationships of 11 species of Laeliinae with emphasis on Brassavola (Orchidaceae)

Brassavola inhabits a wide altitude range and habitat types from Northern Mexico to Northern Argentina. Classification schemes in plants have normally used vegetative and floral characters, but when species are very similar, as in this genus, conflicts arise in species delimitation, and alternative methods should be applied. In this study we explored the taxonomic and phylogenetic value of the anatomical structure of leaves in Brassavola; as ingroup, seven species of Brassavola were considered, and as an outgroup Guarianthe skinneri, Laelia anceps, Rhyncholaelia digbyana and Rhyncholaelia glauca were evaluated. Leaf anatomical characters were studied in freehand cross sections of the middle portion with a light microscope. Ten vegetative anatomical characters were selected and coded for the phylogenetic analysis. Phylogenetic reconstruction was carried out under maximum parsimony using the program NONA through WinClada. Overall, Brassavola species reveal a wide variety of anatomical characters, many of them associated with xeromorphic plants: thick cuticle, hypodermis and cells of the mesophyll with spiral thickenings in the secondary wall. Moreover, mesophyll is either homogeneous or heterogeneous, often with extravascular bundles of fibers near the epidermis at both terete and flat leaves. All vascular bundles are collateral, arranged in more than one row in the mesophyll. The phylogenetic analysis did not resolve internal relationships of the genus; we obtained a polytomy, indicating that the anatomical characters by themselves have little phylogenetic value in Brassavola. We concluded that few anatomical characters are phylogenetically important; however, they would provide more support to elucidate the phylogenetic relantionships in the Orchidaceae and other plant groups if they are used in conjunction with morphological and/or molecular characters.     

Comparative labellar micromorphology of Zygopetalinae (Orchidaceae)

Background and Aims

Molecular evidence indicates that the Neotropical sub-tribe Zygopetalinae is sister to Maxillariinae. Most members of the latter sub-tribe have deceit pollination strategies, but some species produce rewards such as nectar, pseudopollen, resin and wax, and are pollinated by a range of pollinators that include stingless bees (Meliponini), wasps and hummingbirds. By contrast, relatively little is known about the pollination of Zygopetalinae species. However, some are pollinated by fragrance-gathering, male euglossine bees or employ nectar deceit strategies. The aim of this study is to describe the labellar micromorphology of Zygopetalinae and to compare it with that of Maxillariinae sensu lato (s.l.) as part of an ongoing project to record the range of labellar characters found within the tribe Maxillarieae, and to assess whether these characters represent synapomorphies or homoplasies resulting from similar pollination pressures.

Methods

The labella of 31 species of Zygopetalinae, including Cryptarrhena R. Br. and representatives of the Zygopetalum, Huntleya and Warrea clades, were examined using light microscopy and scanning electron microscopy, and the range of labellar characters was recorded. These characters were subsequently compared with those of Maxillariinae s.l. which formed the subject of our previous investigations.

Key Results and Conclusions

The labellar micromorphology of Zygopetalinae is less diverse than that of Maxillariinae and does not reflect the currently accepted phylogeny of the former sub-tribe based on molecular studies. Instead, the relative uniformity in labellar micromorphology of Zygopetalinae is probably due to homoplasies resulting from similar pollinator pressures. Labellar trichomes are relatively uncommon in Zygopetalinae, but occur in certain members of both the Zygopetalum and Huntleya clades. Trichomes are unbranched, uniseriate and multicellular with rounded apices, or unbranched and unicellular, with tapering, pointed and flexuose apices. Hitherto, unicellular trichomes of this kind have been observed only for euglossophilous orchid taxa, and the adoption of a relatively limited range of pollination strategies by Zygopetalinae may have resulted in reduced investment in micromorphological labellar characters.     

Phylogeny and biogeography of the Vitrinidae (Gastropoda: Stylommatophora)

The phylogeny of the Vitrinidae is reconstructed in a cladistic analysis based on characters of the genitalia, the copulation behaviour and the radula. The genera with an atrial stimulator turned out to be the earliest branches of the Vitrinidae, whereas the genera with a glandula amatoria form a monophyletic, taxonomically apomorphic group. The differences between the proposed phylogeny and previous hypotheses are discussed. The ancestral areas of the Vitrinidae and its sister group, the limacoid slugs Boettgerillidae–Limacidae–Agriolimacidae, are estimated using weighted ancestral area analysis. The Vitrinidae and the limacoid slugs might have originated by a vicariance event between Central Europe and the Near East. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 134 , 347–358.     

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.     

Comparative studies of vegetative anatomy in the Theaeeae of Sri Lanka

Anatomical characters of the mature leaf blade, petiole, young stem and wood were examined in order to substantiate taxonomic boundaries between genera and species of Theaeeae in Sri Lanka. Two species of Temstroemia , one of Adinandra , four of Eurya and two of Gordonia were studied.
The presence of sclereids in most organs of the plant was found to be a common character within the family. However, three distinct types of sclereid are reported. Two basic types of stomata, anomocytic and gordoniaceous, are recognized. The origin of phellogen in the young stems is pericyclic in Gordonia and subepidermal in the other three genera. The species within these genera also appear to have several distinctive characters of their own. Important wood characters have been tabulated in order to illustrate the primitive and advanced characters. Gordonia , especially, exhibits anatomical features which are quite different from those of Temstroemia, Adinandra send Eurya.     

Notes on the vegetative morphology and anatomy of the Petermanniaceae (Monocotyledones)

The microscopic anatomy of the vegetative parts of Petermannia cirrhosa is described for the first time. This information lends support to the exclusion of Petermannia from the Dioscorea-ceae with which it has been tentatively associated, the vascular anatomy of the two being quite dissimilar. No satisfactory explanation of the peculiar leaf-opposed position of the tendrils (inflorescence homologues) can be offered, except that it clearly cannot be accounted for by sympodial growth. Vascular tissues in Petermannia are very unspecialized.     

A cladistic analysis of the cornutes (stem chordates)

At the root of the calcichordate controversy is the problem of recognizing homologous similarity. Only when a criterion or set of criteria for hypothesizing homologies is established can the calcichordate theory be tested against its main rival—the Stylophoran theory—by applying the principle of parsimony. The criterion of topological similarity is applied to a three-laxon problem involving a mitrate, the crown-group Echinodermata and the crown-group Chordata. The mitrate shares more features with extant chordates than with extant echinoderms and the calcichordate theory is supported by a simple parsimony analysis. In a cladistic analysis of the cornutes, four monophyletic families are recognized (Cothurnocystidae, Scotiaecystidae, Phyllocystidae and Hanusiidae) and their interrelationships resolved using Hennig86 and the successive-weighting procedure of Farris. Because the known fossil record of cornutes and mitrates is very poor, the correlation between the cladogram produced for cornutes and their order of appearance in the geological record is weak. All of the cornute families must have originated in the Lower Cambrian at the latest.