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1.  Comparative labellar micromorphology of Zygopetalinae (Orchidaceae) 
Annals of Botany  2011;108(5):945-964.
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.
doi:10.1093/aob/mcr202
PMCID: PMC3177679  PMID: 21856635
Cryptarrhena; epidermis; homoplasy; Huntleya clade; labellum; Maxillariinae; papillae; trichomes; Warrea clade; Zygopetalum clade
2.  Comparative histology of floral elaiophores in the orchids Rudolfiella picta (Schltr.) Hoehne (Maxillariinae sensu lato) and Oncidium ornithorhynchum H.B.K. (Oncidiinae sensu lato) 
Annals of Botany  2009;104(2):221-234.
Background and Aims
Floral elaiophores, although widespread amongst orchids, have not previously been described for Maxillariinae sensu lato. Here, two claims that epithelial, floral elaiophores occur in the genus Rudolfiella Hoehne (Bifrenaria clade) are investigated. Presumed elaiophores were compared with those of Oncidiinae Benth. and the floral, resin-secreting tissues of Rhetinantha M.A. Blanco and Heterotaxis Lindl., both genera formerly assigned to Maxillaria Ruiz & Pav. (Maxillariinae sensu stricto).
Methods
Putative, floral elaiophore tissue of Rudolfiella picta (Schltr.) Hoehne and floral elaiophores of Oncidium ornithorhynchum H.B.K. were examined by means of light microscopy, histochemistry, scanning electron microscopy and transmission electron microscopy.
Key Results and Conclusions
Floral, epithelial elaiophores are present in Rudolfiella picta, indicating, for the first time, that oil secretion occurs amongst members of the Bifrenaria clade (Maxillariinae sensu lato). However, whereas the elaiophore of R. picta is borne upon the labellar callus, the elaiophores of O. ornithorhynchum occur on the lateral lobes of the labellum. In both species, the elaiophore comprises a single layer of palisade secretory cells and parenchymatous, subsecretory tissue. Cell wall cavities are absent from both and there is no evidence of cuticular distension in response to oil accumulation between the outer tangential wall and the overlying cuticle in R. picta. Distension of the cuticle, however, occurs in O. ornithorhynchum. Secretory cells of R. picta contain characteristic, spherical or oval plastids with abundant plastoglobuli and these more closely resemble plastids found in labellar, secretory cells of representatives of Rhetinantha (formerly Maxillaria acuminata Lindl. alliance) than elaiophore plastids of Oncidiinae. In Rhetinantha, such plastids are involved in the synthesis of resin-like material or wax. Despite these differences, the elaiophore anatomy of both R. picta (Bifrenaria clade) and O. ornithorhynchum (Oncidiinae) fundamentally resembles that of several representatives of Oncidiinae. These, in their possession of palisade secretory cells, in turn, resemble the floral elaiophores of certain members of Malpighiaceae, indicating that convergence has occurred here in response to similar pollination pressures.
doi:10.1093/aob/mcp119
PMCID: PMC2710904  PMID: 19447811
Bifrenaria clade; elaiophore; floral oil; Heterotaxis; Maxillariinae; Oncidiinae; Oncidium ornithorhynchum; Rhetinantha; Rudolfiella picta; secretion
3.  Labellar Micromorphology of Two Euglossine-pollinated Orchid Genera; Scuticaria Lindl. and Dichaea Lindl. 
Annals of Botany  2008;102(5):805-824.
Background and Aims
Until recently, there was no consensus regarding the phylogenetic relationships of the Neotropical orchid genera Scuticaria Lindl. and Dichaea Lindl. However, recent evidence derived from both gross morphological and molecular studies supports the inclusion of Scuticaria and Dichaea in sub-tribes Maxillariinae and Zygopetalinae, respectively. The present paper describes the labellar micromorphology of both genera and seeks to establish whether labellar characters support the assignment of Scuticaria and Dichaea to these sub-tribes.
Methods
The labella of four species of Scuticaria and 14 species of Dichaea were examined using light microscopy and scanning electron microscopy, and their micromorphology was compared with that of representative species of Maxillariinae sensu lato and Zygopetalinae (Huntleya clade).
Key Results and Conclusions
In most specimens of Scuticaria examined, the papillose labella bear uniseriate, multicellular, unbranched trichomes. However, in S. steelii (Lindl.) Lindl., branched hairs may also be present and some trichomes may fragment and form pseudopollen. Multicellular, leaf-like scales were also present in one species of Scuticaria. Similar, unbranched hairs are present in certain species of Maxillaria Ruiz & Pav. (Maxillariinae sensu stricto) and Chaubardia Rchb.f. (Huntleya clade). As yet, moniliform, pseudopollen-forming hairs have not been observed for Zygopetalinae, but their presence in Scuticaria steelii, Maxillaria and Heterotaxis Lindl. supports the placing of Scuticaria in Maxillariinae. As other genera are sampled, the presence of branched hairs, hitherto unknown for Maxillariinae sensu lato, may prove to be a useful character in taxonomy and phylogenetic studies. Euglossophily occurs in Dichaea, as well as Chondrorhyncha Lindl. and Pescatorea Rchb.f. (Huntleya clade), and all three genera tend to lack distinctive labellar features. Instead, lip micromorphology is relatively simple and glabrous or papillose. However, two of the Dichaea species examined bear unicellular, labellar trichomes very similar to those found in Bifrenaria Lindl. (pollinated by both euglossine bees and Bombus spp.), and this feature may have arisen by convergence in response to similar pollination pressures.
doi:10.1093/aob/mcn155
PMCID: PMC2712378  PMID: 18765439
Bifrenaria; Bifrenaria clade; Chaubardia; Chondrorhyncha; Dichaea; Dichaeinae; Heterotaxis; Huntleya clade; Huntleyinae; labellum; Maxillaria; Maxillariinae; papillae; Pescatorea; scales; Scuticaria; trichomes; Zygopetalinae
4.  Elaiophore Structure and Oil Secretion in Flowers of Oncidium trulliferum Lindl. and Ornithophora radicans (Rchb.f.) Garay & Pabst (Oncidiinae: Orchidaceae) 
Annals of Botany  2007;101(3):375-384.
Background and Aims
Many orchid flowers have glands called elaiophores and these reward pollinating insects with oil. In contrast to other reward-producing structures such as nectaries, the anatomy of the elaiophore and the process of oil secretion have not been extensively studied. In this paper, elaiophore structure is described for two members of Oncidiinae, Oncidium trulliferum Lindl. and Ornithophora radicans (Rchb.f.) Garay & Pabst.
Methods
Elaiophores of both species were examined using light microscopy, scanning electron microscopy and transmission electron microscopy.
Key Results and Conclusions
In flowers of Oncidium trulliferum and Ornithophora radicans, oil is secreted by morphologically distinct elaiophores associated with the labellar callus. However, in O. trulliferum, elaiophores also occur on the lateral lobes of the labellum. In both these species, the epithelial elaiophores are composed of a single layer of palisade-like epidermal cells and a distinct subepithelial layer. Secretory elaiophore cells may contain numerous, starchless plastids, mitochondria and smooth endoplasmic reticulum profiles. In O. trulliferum, the cytoplasm contains myelin-like figures but these are absent from O. radicans. In the former species, cavities occur in the cell wall and these presumably facilitate the passage of oil onto the elaiophore surface. In O. radicans, the accumulation of oil between the outer tangential wall and the cuticle causes the latter to become distended. Since it is probable that the full discharge of oil from the elaiophores of O. radicans occurs only when the cuticle is ruptured by a visiting insect, this may contribute towards pollinator specificity. The structure of the elaiophore in these species resembles both that found in previously investigated species of Oncidiinae and that of certain members of the Malpighiaceae.
doi:10.1093/aob/mcm297
PMCID: PMC2701824  PMID: 18056056
Elaiophore; Oncidium trulliferum; Ornithophora radicans; Orchidaceae; oil secretion; pollination
5.  Micromorphology of the Labellum and Floral Spur of Cryptocentrum Benth. and Sepalosaccus Schltr. (Maxillariinae: Orchidaceae) 
Annals of Botany  2007;100(4):797-805.
Background and Aims
Gross vegetative and floral morphology, as well as modern molecular techniques, indicate that Cryptocentrum Benth. and Sepalosaccus Schltr. are related to Maxillaria Ruiz & Pav. However, they differ from Maxillaria in their possession of floral spurs and, in this respect, are atypical of Maxillariinae. The labellar micromorphology of Maxillaria, unlike that of the other two genera, has been extensively studied. In the present report, the labellar micromorphology of Cryptocentrum and Sepalosaccus is compared with that of Maxillaria and, for the first time, the micromorphology of the floral spur as found in Maxillariinae is described.
Methods
Labella and dissected floral spurs of Cryptocentrum and Sepalosaccus were examined using light microscopy (LM) and scanning electron microscopy (SEM).
Key Results
In each case, the labellum consists of a papillose mid-lobe (epichile), a cymbiform region (hypochile) and, proximally, a spur, which is pronounced in Cryptocentrum but short and blunt in Sepalosaccus. The inner epidermal surface of the spur of Cryptocentrum is glabrous or pubescent, and the bicellular hairs, where present, are unlike any hitherto described for Maxillariinae. Similar but unicellular hairs also occur in the floral spur of Sepalosaccus, whereas the glabrous epidermis lining the spur of C. peruvianum contains putative nectar pores.
Conclusions
The labellar micromorphology of Cryptocentrum and Sepalosaccus generally resembles that of Maxillaria. The floral spur of Cryptocentrum displays two types of organization in that the epidermal lining may be glabrous (possibly with nectar pores) or pubescent. This may have taxonomic significance and perhaps reflects physiological differences relating to nectar secretion. The trichomes found within the spurs of Cryptocentrum and Sepalosaccus more closely resemble the hairs of certain unrelated, nectariferous orchid taxa than those found in the largely nectarless genus Maxillaria, and this further supports the case for parallelism.
doi:10.1093/aob/mcm165
PMCID: PMC2749631  PMID: 17686763
Labellum; Maxillariinae; micromorphology; nectar pore; nectary; spur; trichome

Results 1-5 (5)