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.     

Atypical pseudopollen-forming hairs in Maxillaria (Orchidaceae)

Pseudopollen is found on the labella of several species of Maxillaria and is formed by the fragmentation of uniseriate, multicellular, moniliform trichomes. The resultant component cells are rich in protein. Since flowers of pseudopollen-forming species generally lack nectar, it is probable that pseudopollen gradually replaced nectar as the pollinator reward. However, direct evidence for this hypothesis is lacking. The present paper examines the labellar micromorphology of five Brazilian species of Maxillaria assigned to the M. discolor alliance. The flowers of two of these species, namely M. violaceopunctata and M. villosa , produce both food hairs and a lipoidal, labellar secretion which is rich in aromatic amino acids. Moreover, in the case of M. violaceopunctata , the secretion was found to contain reducing sugars. This may represent an intermediate stage in the transition from nectar-producing to pseudopollen-forming flowers. SEM studies indicate how pseudopollen-forming, moniliform trichomes, such as those found in the Maxillaria grandiflora complex, may have evolved from simple, uniseriate, multicellular hairs.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 151−158.     

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.     

Taxonomic separation of the genera Prosthechea and Encyclia (Laeliinae: Orchidaceae) using leaf and root anatomical features

Despite formal separation based on molecular and morphological evidence, the genera Prosthechea Knowles & Westc. and Encyclia Hook. have not been studied in terms of their vegetative anatomy. In this study we examine 16 Brazilian species of these genera. Additionally, one species of subtribe Laeliinae and another from subtribe Oncidiinae were studied in order to evaluate the taxonomic consistency of the anatomic characters observed. Except for Epidendrum crassifolium Lindl., all species possess a velamen differentiated into epivelamen and endovelamen. Endodermal cells with uniformly thickened walls, calcium oxalate raphides and cells with phi thickenings in the cortical region were observed in all species. Silica bodies, raphides and fibre bundles in the mesophyll were common in leaves of all species studied. By contrast, flavonoid crystals were observed in both roots and leaves of Prosthechea only. Encyclia is characterized by the presence of fibre bundles on the subepidermal layer, and a cuticle, which is clearly thicker than that of Prosthechea . These data strongly support the separation of Prosthechea and Encyclia .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 293–303.     

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.     

Anatomical studies on Sinofranchetia chinensis (Lardizabalaceae) and their systematic significance

The anatomical structures of the Chinese endemic and monotypic genus Sinofranchetia (Lardizabalaceae) are described. There are reticulate, simple-reticulate, scalariform, simple-scalariform and simple perforations in vessel elements as well as in the fibres in the secondary wood of the roots and the stems. The node is trilacunar. The vascular bundles in the petiole are arranged in a ring. Clustered crystals occur in the parenchymatous cells of stems, petioles and pedicles. Leaf stomata are actinocytic. The nodes of sepals, petals and stamens both in male and female flowers are unilacunar and one-traced. There are three sterile carpels with two to three traces in the male flowers, three fertile carpels with two to three traces, and sometimes three sterile carpels lacking a vascular supply. In morphology, the anther dehiscence mechanism and pollen in the female flowers are the same as in the male flowers, such that the so-called female flowers might be bisexual in morphology. In comparing morphology, the sex of the flowers and the perforations of the vessel elements in Sinofranchetia with Decaisnea and other genera of the Lardizabalaceae, Sinofranchetia is considered a basic group at least as the same evolutionary level in the family as Decaisnea . © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 271–281.     

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.     

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.     

Systematics of the tribe Podalyrieae (Fabaceae) based on DNA, morphological and chemical data

Phylogenetic relationships among nine genera and 28 species of the southern African tribe Podalyrieae were estimated from sequences of the internal transcribed spacer (ITS) of nuclear ribosomal DNA as well as morphological and chemical data. Morphological and ITS sequence data produced cladograms with similar topologies, both supporting the monophyly of Podalyrieae (excluding Hypocalyptus ). The combined data sets indicate that subtribe Xiphothecinae are monophyletic, but embedded within Podalyriinae. The high degree of congruence between previous taxonomic hypotheses and those based on DNA data provides further evidence for the utility of ITS sequences in studying phylogeny.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 159–170.     

Wood and stem anatomy of woody Amaranthaceae s.s.: ecology, systematics and the problems of defining rays in dicotyledons

Wood and stem anatomy is studied for seven species of six genera (root anatomy also reported for one species) of Amaranthaceae s.s. Quantitative data on vessels correlate closely with relative xeromorphy of respective species, agreeing with values reported for dicotyledons without successive cambia in comparable habitats. Libriform fibre abundance increases and vessel diameter decreases as stems and roots of the annual Amaranthus caudatus mature. Long, thick-walled fibres in Bosea yervamora may be related to the upright nature of elongate semi-climbing stems. Non-bordered or minutely bordered perforation plates characterize Amaranthaceae, as they do most other Caryophyllales. Amaranthaceae have idioblastic cells containing druses, rhomboidal crystals or crystal sand: these forms intergrade and seem closely related. Rays are present in secondary xylem of the Amaranthaceae studied. Cells intermediate between ray cells and libriform fibres occur in Charpentiera elliptica . Degrees of diversity in rays and reports of raylessness in Amaranthaceae induce discussion of definition and identification of rays in dicotyledons; some sources recognize both rays and radial plates of conjunctive tissue in Amaranthaceae. The action of successive cambia is described: lateral meristem periclinal divisions produce secondary cortex externally, conjunctive tissue internally and yield vascular cambia as well. Vascular cambia produce secondary phloem and secondary xylem, in both ray and fascicular zones, as in a dicotyledon with a single cambium. Identification of meristem activity and appreciation of varied ray manifestations are essential in understanding the ontogeny of stems in Amaranthaceae (which have recently been united with Chenopodiaceae).  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 1–19.     

Neotropical ennomine moths: a review of the genera (Lepidoptera: Geometridae)

The classification of the Neotropical genera of the Ennominae is reviewed and 267 genera are recognised to occur in this region. Three new genera are described and three others are reinstated, while 48 generic synonyms are newly established. Other changes established in this work include 14 species synonyms and 237 new or reinstated combinations. External features and genitalia of representative members of the genera are illustrated (753 figures). All the known Neotropical species and subspecies of Ennominae are listed ( c . 3470), plus their synonyms. The tribes to which the genera belong are assessed, with c . 200 of the genera assigned to tribe or other suprageneric grouping.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 135 , 121–401.     

The systematic significance of morphological and anatomical variation in fruits of Crotalaria and related genera of tribe Crotalarieae (Fabaceae)

The phylogenetic and taxonomic significance of morphological and anatomical trends in fruits of tribe Crotalarieae has been evaluated, with emphasis on the genus Crotalaria and its seemingly distinctive, inflated and balloon‐shaped pods. In addition to the normal explosive dehiscence as a means of dispersal, several genera (including Crotalaria) show independent evolution of modifications apparently adapted for dispersal by wind, water and gravity. Transverse sections were made of mature pods of 142 species from the 12 currently recognized genera of Crotalarieae. The taxa differ in the orientation of the fibres (related to dehiscence or non‐dehiscence), the overall thickness of the fruit wall, the relative proportions of the pericarp layers, the degree of lignification and the presence or absence of trichomes. Three basic pericarp types can be distinguished: type I, with one, two or three zones of various numbers of cell layers of fibres (almost all genera); type II, with a single cell layer of fibres (only in Rothia, Robynsiophyton, Lebeckia and Lotononis sections Listia and Leobordea); and type III, with one zone of several cell layers of gelatinous fibres and multicellular trichomes associated with the endocarp (only in some species of Calobota and Wiborgiella). Considerable variation was encountered in the tribe, but Crotalaria appears to be rather uniform, with type I predominating. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 84–106.     

Anatomy of extrafloral nectaries in Fabaceae from dry‐seasonal forest in Brazil

Extrafloral nectaries (EFNs) are found in many species of Fabaceae. The aim of this work is to describe the internal morphology of the EFNs from species of Fabaceae found in areas of dry‐seasonal forest in north‐eastern Brazil. All species of Fabaceae with EFNs found were collected and samples were submitted to conventional techniques for anatomical and scanning electronic microscopy analysis. EFNs were found in 35 species, of which 32 were examined anatomically. All types have epidermal cells, secretory tissues and vascular bundles in the EFNs. Sclerenchymatous cells were found between the secretory tissues and the vascular tissues, with a few exceptions. The function of these cells is not clear; however, a role in the transportation of the sap in the nectary or with the support of the secretory tissue is possible. The nectar is released through glandular trichomes, secretory pores or even by breaking the epidermal cells and cuticle. The internal patterns found in the EFNs from different species and genera can provide important information for taxonomic and evolutionary studies in the family. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 87–98.     

A review of the genera of Macariini with a revised classification of the tribe (Geometridae: Ennominae)

The generic classification of moths of the tribe Macariini Guenée is reviewed critically, and a revised classification is presented. The review is based on a survey of species across the taxonomic and geographical range of the tribe. In the new framework, the number of genera is reduced considerably. Two genera are very large: Macaria Curtis (mainly New World) and Chiasmia Hübner (largely Old World) together include over half of all macariine species. Twenty-three genera are accepted in the tribe; the identity of two genera remain uncertain. No single character defines the tribe, but diagnostic features include one or more of the following: the presence of enlarged setae ('horns') on the uncus in the male genitalia; a divided valva; and a modified condition of sternum A8 in the male. The taxonomic history of the tribe is reviewed briefly and the problems of previous systems are explained mainly by the regional approach adopted. A diagnosis is presented for each genus. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 257–315.     

Phylogenetic relationships in Cyperus L. s.l. (Cyperaceae) inferred from plastid DNA sequence data

The phylogeny of Cyperus and allied genera has been reconstructed using cladistic analysis of plastid rbcL gene, rps16 intron, trnL intron, and trnL-F intergenic spacer sequence data in 40 species of tribe Cypereae. Cyperus s.s. as currently circumscribed is not monophyletic because ten cyperoid genera are embedded within it. Eucyperoid Cyperus species (with a C₃ anatomy, e.g. C. involucratus ) and the genera Courtoisina , Kyllingiella and Oxycaryum form a clade that is sister to a clade comprising chlorocyperoid species (with a C₄ anatomy, e.g. C. papyrus ) and the genera Alinula , Ascolepis , Kyllinga , Lipocarpha , Pycreus , Remirea and Sphaerocyperus . The position of two species is uncertain; C . tenellus is resolved in a clade together with Isolepis although with typical cyperoid spikelets, whereas I. humillima is not resolved near either Isolepis or Cyperus s . l . © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 145–153.     

Leaf and stem anatomy of Vochysiaceae in relation to subfamilial and suprafamilial systematics

Several leaf anatomical features are potentially systematically informative within both the family Vochysiaceae and the order Myrtales, notably tracheoidal idioblasts, mucilage cells and secretory canals. Tracheoids with spiral wall thickenings are present in the mesophyll of most species of Vochysia , and also occur in several other families of Myrtales. Mucilage cells are common in the leaf epidermis in some Vochysiaceae. Secretory ducts are present in the midrib in Salvertia and Vochysia , which are apparently closely related, although Salvertia also shares some leaf anatomical characters with Qualea and Callisthene . Anatomical data do not support the segregation of Ruizterania from Qualea ; leaves of R. albiflora leaves are very similar to those of Q. paraensis in venation pattern, and leaf and stem anatomy. Different venation patterns are characteristic of sections within the genus Qualea , but within the large genus Vochysia , leaf anatomy is variable even within a subsection. Amongst other Myrtales, leaf anatomy of Vochysiaceae most closely resembles that of Combretaceae and Onagraceae. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 339–364.     

Relationships, classification and evolution of Scleranthus (Caryophyllaceae) as inferred from analysis of morphological characters

Study of floral anatomy, micromorphology, palynology and onotogeny has revealed new characters for phylogenetic analysis in the genus Scleranthus . Cladistic analysis of these characters, along with those previously available, suggests that the genus consists of Eurasian/Mediterranean and Australasian sister clades. Gynoecial morphology and development are closely similar in all species, suggesting the genus is monophyletic despite its disjunct northern and southern hemisphere distribution. Variation in pollen:ovule ratios and their implications for the evolution of Scleranthus species are also discussed and it is concluded that a range of breeding strategies intermediate between autogamy and xenogamy exists in the genus. Single-stamened species of Scleranthus are likely to be obligate autogams, despite their comparatively high pollen:ovule ratios in relation to autogamic species of other genera.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 15–29.