Pericarp structure and systematic positions of five genera of Lamiaceae subg. Nepetoideae tribe Ocimeae

Pericarp structure has been investigated in five genera of Lamiaceae subg. Nepetoideae tribe Ocimeae which were omitted or poorly studied by Ryding (1992a): Hanceola, Limniboza, Skapanthus, Tetradenia and Thorncroftia . In the light of the results, the systematic position of these and some related genera are discussed. Limniboza is suggested to belong to Ocimeae subtribe Ociminae, the Acrocephalus group. Hanceola (incl. Siphocranion), Isodon, Skapanthus (which is resurrected as a genus), Tetradenia and Thorncroftia apparently do not fit into any of the three traditional subtribes (Hyptidinae, Ociminae or Plectranthinae). The three Asiatic genera, Hanceola, Isodon and Skapanthus might together form a new subtribe. Tetradenia and Thorncroftia have more isolated positions in the Ocimeae.     

Phylogeny and evolution of basils and allies (Ocimeae, Labiatae) based on three plastid DNA regions

A phylogeny of basils and allies (Lamiaceae, tribe Ocimeae) based on sequences of the trnL intron, trnL-trnF intergene spacer and rps 16 intron of the plastid genome is presented. Several methods were used to reconstruct phylogenies and to assess statistical support for clades: maximum parsimony with equally and successively weighted characters, bootstrap resampling, and Bayesian inference. The phylogeny is used to investigate the distribution of morphological, pericarp anatomy, chemical, and pollen characters as well as the geographical distribution of the clades. Tribe Ocimeae is monophyletic and easily diagnosable with morphological synapomorphies. There are monophyletic clades within Ocimeae that broadly correspond to currently recognised subtribes: Lavandulinae, Hyptidinae, Ociminae, and Plectranthinae. Only Lavandulinae has clear non-molecular synapomorphies. Several currently recognised genera are not monophyletic. Floral morphology consistent with sternotribic pollination is most common in Ocimeae, but there are independent departures from this model. Buzz pollination is likely in some species, the only postulated occurrence of this within Lamiaceae. Quinone diterpenoids and flavones in the leaf exudates differ in their distributions across the phylogeny and this could contribute to differences in the recorded medicinal as well as pesticidal uses of the species in the different clades. Mapping geographic distribution on to an ultrametric phylogenetic tree produced using non-parametric rate smoothing supports an Asiatic origin for Ocimeae. There are several secondary occurrences in Asia arising from the African Ociminae and Plectranthinae clades. Colonisation of Madagascar occurred at least five times, and New World colonisation occurred at least three times.     

Pericarp structure and myxocarpy in selected genera of Nepetoideae (Lamiaceae)

The pericarp structure has been analyzed in 37 species representing 13 genera from four tribes of Nepetoideae (Lamiaceae). 11 species are endemics of Balkan Peninsula or other Mediterranean regions. Basically, the pericarp was similar to other Nepetoideae by having exocarp, mesocarp s. str., vertically arranged bone cells and thin innermost cell layer. The ratio between pericarp thickness and nutlet size was the highest in Mentha spp. According to this parameter Micromeria species belonging to different sections could be distinguished. Acinos spp. were characterized by the thickest sclerenchyma in ratio to pericarp thickness. Crystals were present within sclerenchyma region in genera Mentha, Melissa, Nepeta and Prunella . The mucilage production has also been tested. Most of the studied species produced mucilage when becoming wet. The strongest mucilage reaction showed Acinos and Prunella . Pericarp characters studied are of taxonomical interest. They were mainly correlated with the generic or infrageneric classification of some genera.     

Pericarp structure and phylogeny of Lamiaceae subfamily Pogostemonoideae

The pericarp structure has been investigated in 11 species representing the genera of the subfamily Pogostemonoideae (emend. Cantino et al. 1992): Anisomeles, Colebrookea, Comanthosphace, Leucosceptrum, Pogostemon and Rostrinucula . The pericarp structure of Anisomeles, Colebrookea and Pogostemon have been found to resemble that of the subfamilies Lamioideae or Nepetoideae. The three related genera, Comanthosphace, Leucosceptrum and Rostrinucula have been found to have a divergent type of pericarp, which lacks sclerenchyma and has druses and a type of cells with scalariform secondary thickenings in the mesocarp. The three genera are suggested to be extraneous in Pogostemonoideae and their systematic position is uncertain.     

Crystals in calyces of Lamiaceae and their phylogenetic and adaptive significance

The diversity of crystal types is studied in fruiting calyces of 435 out of the c. 7,200 species, and 184 out of the c. 236 genera of Lamiaceae, mainly in the subfamilies Nepetoideae and Lamioideae. Calcium oxalate crystals are found in about half of the studied calyces, and belong to the following main types: prismatics in the inner epidermis, prismatics in fibres, prismatics in sclereids, prismatics in the mesophyll and druses in the mesophyll. Presence of epidermal prismatics may constitute a synapomorphy of the subfamily Nepetoideae or of a subclade within this subfamily consisting of the two largest tribes Mentheae and Ocimeae. The presence of narrow mesophyll prismatics seems to support a large clade of Lamioideae genera in a recent molecular phylogeny of the subfamily. Epidermal prismatics are found to be particularly frequent in closed calyces, calyces with a narrow mouth and internally hairy calyces. These findings and the position of the crystals in the calyces are regarded as supporting the hypothesis that calcium oxalate crystals protect the fruit against insect predators.     

Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae)

Pericarp structure is investigated in 143 specimens belonging to 59 out of ca. 65 genera and 134 out of ca. 2,000 species of Lamiaceae subfamily Nepetoideae tribe Mentheae. Some of the data presented by earlier authors are believed to be incorrect. The variation in pericarp structure has been found to be strongly correlated to the variations in DNA and gross morphological characters. The groups of two-staminate genera (Salvia, etc.) and four-staminate genera of Salviinae (Lepechinia and Chaunostoma) can be distinguished from each other and from other labiates by differences in pericarp structure. The other two subtribes, Menthinae and Nepetinae, differ considerably, but their variations strongly overlap. The differences in pericarp structure also suggest that Menthinae can be divided into three main monophyletic groups based on other data: (1) the main bulk of the tribe, (2) a group of Prunella and Cleonia and perhaps also Horminum, and (3) the characteristic genus Lycopus. Pericarp data disagree with a suggestion that the genus Melissa should be included in the subtribe Salviinae. Modifications of the subtribal classification of Mentheae are discussed.     

Pericarp structure and phylogeny of theLamiaceae-Verbenaceae-complex

Pericarp structure was investigated in 158 species of the familiesLamiaceae andVerbenaceae. Data from 221 out of 262 genera ofLamiaceae s.l. and a few ofVerbenaceae s.str. were collected in a table. A cladistic analysis was performed on the basis of pericarp characters only. The abandonment of subfam.Pogostemonoideae as a taxonomic unit is considered. Examples of groups given additional support by similarities in pericarp characters are: (1) the gynobasic-styled labiates (subfamiliesPogostemonoideae, Lamioideae, Nepetoideae); (2) aLamioideae-Pogostemonoideae-group; (3)Nepetoideae; (4) aWestringia-Hemigenia-Hemiandra-Microcorys group (in subfam.Chloranthoideae); (5) aLepechinia-Chaunostoma-group (inNepetoideae); (6) aPrunella-Cleonia-group (inNepetoideae).     

Leaf anatomy of Cunoniaceae

A study of leaf anatomy of 24 genera of Cunoniaceae was made. The prevailing petiole vasculature is a nearly complete, usually adaxially flattened, medullated cylinder with the flat dorsal segment separated from the ventral arc. Medullary and cortical vasculation in Codia, Cunonia, Geissois and Pancheria is correlated with an increase in leaf coriaceousness. Very few characters are uniformly found in leaves of all species; these are druse or prismatic crystals, bifacial mesophyll and unicellular and simple trichomes. In most species vein sheathing occurs. Characters of the lamina that show variation and are useful in generic delimitation are: trichome type, presence of a hypodermis, occurrence of mucilaginous cells, presence of specialized terminal veinlet cells, major and higher order venation patterns, vein sheathing type, and mature stomatal types. Stomata are of the anomocytic, paracytic or anisocytic types. Leaves of various cunoniaceous species of xerophytic environments exhibit a leathery texture, thicker cuticle, more intensive sclerification along the veins, a sclerified hypodermis, and an increase in the amount of vascular tissue per unit area. Leaf anatomy does not provide immediate clues toward elucidating the relationships of cunoniaceous genera.     

A systematic histological study of palm fruits. VI. Subtribe Linospadicinae (Arecaceae)

The pericarp structure of representative species of the four genera of the subtribe Linospadicinae is described and compared. Tissues found in the pericarp of this group are similar to those found in other subtribes of the Areceae, with no characters unique to the subtribe and nothing to suggest its closest affinities. The four genera, as well as each species examined, do show distinctive pericarp features, however.Laccospadix andLinospadix are similar, both with a single series of fibrous, bundles and an outer series of prominent raphide-bearing cells dominating the pericarp. In bothHowea andCalyptrocalyx, a complex exocarp forms from a series of fibrous bundles and brachysclereids, but each genus has other distinctive characters.Howea has vascular bundles in the exocarp zone, an outer series of raphide-bearing cells, and a conspicuously thickened locular epidermis. Based on a limited sample of four species, raphide-bearing cells appear to be always interior to the exocarp inCalyptrocalyx, and the locular epidermis is thin.     

中国伞形科天胡荽亚科果实解剖特征及其系统学意义

经对中国伞形科天胡荽亚科 (Hydrocotyloideae)中天胡荽属 (HydrocotyleL .)积雪草属 (CentellaL .)和马蹄芹属(DickinsiaFranch .) 10种植物果实解剖结构的观察比较 ,发现 3属的果实横切面以合生面为基准可以分成 2个类型 ,即两侧压扁类型 (天胡荽属和积雪草属 )和背腹压扁类型 (马蹄芹属 )。除此之外 ,中果皮的外侧是否有色素块沉积、内侧是否有木化细胞层或厚壁细胞层以及最内层是否形成晶体细胞层 ,内果皮细胞的层数、排列方向以及有无棱槽油管等性状 ,可以作为区分 3属的重要解剖学特征。在此基础上 ,结合外部形态特征 ,讨论了该亚科及其内部各属的解剖学特征、属间区别及其演化关系。认为马蹄芹属应归属于Mulineae族 ,另外 2个属应归属于天胡荽族(Hydrocotyleae) ;3属中积雪草属原始而马蹄芹属进化 ;天胡荽亚科可能不是自然的类群 ,它的分类系统和演化地位有待深入研究。     

The systematic relevance of fruit and seed structure in Bersama and Melianthus (Melianthaceae)

 The fruit and seed anatomy and morphology of the two genera Bersama and Melianthus (Melianthaceae, Sapindales) have been studied in an effort to clarify their systematic position. On the basis of the differences in pericarp and seed anatomy as well as in other exomorphic characters the segregation of Bersama into a distinct family Bersamaceae is supported. Evidence mainly from seed anatomy and morphology emphasizes the anomaly of the traditional inclusion of Bersama and Melianthus in the Sapindales, since they have a distinct seed-coat structure and seed vascularization. The fruit and seed anatomy does not confirm any relationships with alternatively suggested exo-mesotestal Lardizabalaceae. The exotestal seed coats of Bersama and Melianthus with a differentiated palisade of Malpighian cells in the exotesta, dimerous raphal vascular skeleton, abundant endosperm, and a small differentiated straight embryo show a resemblance with the exotestal albuminous seeds of Rhamnaceae and Elaeagnaceae. Using also additional data on fruit, floral and vegetative morphology it is suggested that Bersamaceae with Melianthaceae and Rhamnaceae/Elaeagnaceae constitute a distinct relict side-branch of exo-mesotestal rosidaceous ancestry, the new order Melianthales in the superorder Rhamnanae (Rosidae). The formerly suggested relationship of this side-branch to exotegmic Malvales is not supported by seed anatomy. The affinity with exotegmic Celastrales, which are considered as a possible connecting link between archaic exo-mesotestal Rosales and exotestal Rhamnales/Elaeagnales, is also found untenable. Received May 12, 2000 Accepted December 7, 2000     

SEM and Light Microscope Observations on Fruit and Seeds in Scrophulariaceae from Southwest Spain and their Systematic Significance

Many studies of the morphology and anatomy of fruits and seedsof Scrophulariaceae from Southwest Spain have been made. Thesystematic utility of characteristics of these plant structuresis discussed in order to determine relationships among the differentgenera included in this widely distributed family. The mostuseful fruit features for this study were the indumentum, capsuledehiscence and structure of the endocarp, seed-coat ornamentation,and inner structure of the seeds. In addition, phenetic analysisof 58 characters revealed that several groups of genera in Scrophulariaceaeare closely related on the basis of their fruit and seed features.The phenetic relationships of genera or groups of genera arediscussed. Keys to genera considered in this work, using exclusivelyfruit or seed characters, are provided. Lastly, the usual dispersalsystems observed in this family are discussed. Copyright 2000Annals of Botany Company Fruit, seed, morphology, anatomy, Scrophulariaceae, phenogram, SEM, pericarp, dehiscence, indumentum, seed-coat     

Comparative leaf anatomy of the Penaeaceae and its ecological Implications

The leaf anatomy of species representing all seven genera of the Penaeaceae was studied by light and scanning electron microscopy. Due to variability and inconsistency, leaf anatomical characters are not regarded as particularly useful for systematics within or among genera in this family. Across the family, a number of taxa exhibit a trend towards amphistomatous, isobilateral leaves, generally associated with increased leaf thickness and amount of palisade mesophyll. This trend is not apparent in closely related families, e.g. Alzateaceae, Cryp-teroniaceae, Oliniaceae and Rhynchocalycaceae. Most species are found in comparatively mesic habitats and it is difficult to postulate primary xeromorphic trends in leaf anatomy. The sclerophylly encountered across the family as a whole is likely to be related to paucity of soil nutrients rather than a response to water stress.     

Study of the oleanolic and ursolic acid contents of some species of the Lamiaceae

The oleanolic and ursolic acid contents from 88 taxa of Lamiaceae (19 genera, 66 species, 8 subspecies, 9 varieties and 5 hybrids) were investigated using gas chromatography. Both triterpenoid acids were present in all of the investigated taxa, but the plants belonging in the subfamily Nepetoideae produced significantly higher amounts than those in the subfamily Lamioideae. The oleanolic acid content ranged from traces to 1.840% dry weight, and that of ursolic acid from traces to 4.019% dry weight.     

Seed structure and the taxonomy of the Cruciferae

The external morphology and anatomy of the seeds of some 90 genera and some 200 species of the Cruciferae have been investigated. Emphasis has been laid on the relationships of seed structure to taxonomy. As regards seed anatomy, the testa is probably of greatest value in determining taxonomic relationships. Seed structure supports the Cruciferae as a natural taxon. In the present investigation, few genera and species show seed characters which are either exclusive or very distinct. Of the remainder, most genera are distinguishable from each other on a permutation of their macroscopic and microscopic seed characters. However it is difficult or impossible to distinguish between some genera on seed characters. The present investigation provides no support for tribal divisions within the Cruciferae.     

Comparative leaf anatomical studies of some Sterculia L. species (Sterculiaceae)

A comparative study was undertaken on the leaves of 12 Sterculia species in order to assess anatomical variations which may be useful in species identification and to evaluate their significance in the taxonomy of the genus. All species have glandular and non-glandular trichomes, anomocytic stomata, multiple and mucilaginous epidermis, mucilaginous cavities and druses in mesophyll tissues. A small amount of variation was observed in the outline of transverse sections of midribs and petioles, the presence of central vascular bundles, and the type of trichomes. Some characters, such as the vertically divided adaxial epidermis in S. macrophylla , are unique to certain species and are therefore useful in diagnostic characters, while S. coccinea and S. elongata appear to be similar in the structure of the petiole and midrib. The results, however, do not show any groupings of species and do not support the observations of others on groups derived from wood anatomy.     

The Morphological and Anatomical Characters of Bamboo Fruits with Relation to Systematics and Evolution

The morphological and anatomical characteristics of bamboo fruits of 39 species in 21 genera have been surveyed. The results showed that characters as the shape, the middle cross section, the morphology of pericarp, rostrum and embryo, the presence or absence of ventral groove and embryotega, and the thickness of aleurone layer of fruits were the important features in identifying different genera, and also in studying systematics and evolution of the bamboos. All the bamboo fruits examined herewith could be considered as caryopses, and they could be subdivided into four types viz. typical, bacoid, nucoid and seminucoid type. Type A,typical,as most grass fruits, with thin pericarp throughout or thickened only at apex, ventral groove (hilum) and embryotega, was found in the majority of bamboos, such as Fargesia and Arundinaria. Type B, bacoid, with thick and fleshy pericarp, was found in nine genera: Melocanna, Dicochloa, Ochlandra,Olrneca, Melocalarnus, Chirnonobarnbusa ,Qiongzhuea, Ferrocalamus and Alvirnia. Type C, nucoid, with hard pericarp, was found so far only in Dendrocalamus and Cephalostachyurn. Type D,semi-nucoid, the intermediate type between type A and C, with pericarp, as in type A, in the dorsum and pericarp, as in type C in the ventrum was found in Sinocalarnus, PhyUostachys and Thyrsostachys. Bacoid may be the primitive type, from which nueoid may have evolved. Semi-nucoid may have evolved from nucoid. Typical type may be advanced and have evolved from bocoid or semi-nucoid. The primitive fruits of bamboos may have characteristics such as follows: globose or spheroidal in shape, round middle cross section; thick and fleshy pericarp throughout, less differentiated, presence of vascular bundle; absence of ventral groove (hilum) and embryotega; aleurone layer composed of a number of cell layers; large and curved'or folded embryo. Bamboos with such a fruit have iterauctant inflorescences except Ferrocalamus.     

Survey of leaf anatomy,especially secretory structures,of tribeCaesalpinieae (Leguminosae,Caesalpinioideae)

We studied leaflet anatomy, emphasizing secretory structures, from herbarium specimens of 128 species of 44 genera of tribeCaesalpinieae, using clearings, resin sections, and scanning electron microscopy. These observations, combined with those from our three earlier papers, provide a survey of 210 species representing all genera. Seventy-three species had secretory structures: 21 had glands or gland-like trichomes, 40 had living mesophyll idioblasts, and nine had cavities (three species each had two different types). Five additional species, all inCercidium (Caesalpinia group), had paired or clustered large spheroidal, thick-walled, empty cells (veinlet idioblasts) interconnected by perforation plate-like gaps. Secretory structures have systematic significance at various taxonomic levels.     

Revision of generic limits in Hyptis Jacq. (Labiatae) and its allies

The primarily neotropical sub-tribe Hyptidinae (Labiatae: tribe Ocimeae) consists of six or seven genera of which the largest is the genus Hyptis with over 300 species. Over the last 12 years, work on the group at the Royal Botanic Gardens, Kew, examining macro-morphological and anatomical characters, as well as pollen surface and chromosome number, has indicated that the genus Hyptis is paraphyletic, with a few of the larger-flowered sections apparently being more closely related to the neighbouring genus Eriope than to the smaller-flowered sections of Hyptis . This has been confirmed by preliminary cladistic studies now in progress. In order to provide a working classification for the group, prior to the publication of the more detailed study, two new genera: Hypenia and Hyptidendron are created. A key to genera of South American Hyptidinae is provided, as well as one to distinguish the two new sections of Hyptidendron . Forty-five new combinations are made.     

A systematic histological study of palm fruits. III. Subtribe Iguanurinae (Arecaceae)

This study represents a preliminary sampling of the pericarp histology of the subtribe Iguanurinae (tribe Areceae, subfamily Arecoideae) of the family Arecaceae. At least one sample from each of the 27 recognized genera was examined and illustrated with a line drawing. This sampling serves to characterize fruit structure in the subtribe as a whole, to illustrate the diversity of pericarp adaptations found in the subtribe, to characterize the monotypic genera, to provide hypotheses about the characterization of the larger genera, and to test existing phylogenetic hypotheses about the Iguanurinae. There are no unique tissues present in the pericarp in this subtribe, but genera can be readily characterized by unique combinations and distributional patterns in common tissues. These patterns, and some prominent evolutionary trends, parallel those in related subtribes of Areceae, such as the Ptychospermatinae and Arecinae. Significant in this subtribe is variation in the distribution of tanniniferous cells, raphide-bearing cells and brachysclereids, in the sculpturing of the seed and the locular epidermis, in the thickness of the locular epidermis, in the thickness of the fibrous vascular bundle sheaths, and especially in the number, orientation and distribution of nonvascular fibrous bundles. One major trend is the formation of systems of separate fibrous bundles and their progressive displacement toward the outer layer of the fruit, where a complex exocarp may form. The diversity of pericarp structure in the Iguanurinae is far greater than in the two subtribes previously studied.