Wood anatomy of Penaeaceae (Myrtales): comparative, phylogenetic, and ecological implications

Wood samples of stems, lignotubers, and roots of the majority of species of Penaeaceae were analyzed with respect to qualitative and quantitative features. Virtually no data have hitherto been presented on xylem features of this family, restricted to Cape Province, South Africa. Presence of vestured pits in vessels, septate crystalliferous parenchyma in wood, intraxylary phloem, predominantly erect ray cells in the typically narrow, multiseriate rays and in the uniseriate rays, and amorphous deposits in ray cells place Penaeaceae securely in Myrtales and help to define that order. By comparison of ecological preferences of the species, as observed during field work, with quantitative analysis of conductive tissue, close correspondence of the wood structure to habit and habitat is demonstrated.     

Leaf and twig anatomy of the Pterostemonaceae (Engl.) Small: ecological and systematic features

An anatomical investigation of the leaves and twigs of Pterostemonaceae (Engl.), a monogeneric family of two species, has been made. The following anatomical characters in the leaf are of particular interest: glandular hairs, hydathodes on the marginal dentations and secretory substances in the glands and palisade cells. Characters of interest in the twig xylem include: vessel lumina of very small tangential diameter and with simple perforation plates; fibriform vessels with scalariform plates having one to six bars and also plates with perforations in irregular patterns.     

Wood anatomy of the endemic woody Asteraceae of St Helena I: phyletic and ecological aspects

Quantitative and qualitative data are given for samples of mature wood of all eight species of woody Asteraceae, representing three tribes, of St Helena I. The quantitative features of all except one species are clearly mesomorphic, corresponding to their mesic central ridge habitats. Commidendrum rugosum has more xeromorphic wood features and occurs in dry lowland sites. Commidendrum species are alike in their small vessel pits and abundant axial parenchyma. Melanodendrum agrees with Commidendrum in having fibre dimorphism and homogeneous type II rays. The short fibres in both genera are storied and transitional to axial parenchyma. Elongate crystals occur in ray cells of only two species of Commidendrum , suggesting that they are closely related. Wood of Commidendrum and Melanodendrum is similar to that of the shrubby genus Felicia , thought closely related to Commidendrum on molecular bases. Commidendrum and Melanodendrum have probably increased in woodiness on St Helena, but are derived from shrubby ancestors like today's species of Felicia. Petrobium wood is paedomorphic and indistinguishable from that of Bidens , from which Petrobium is likely derived. The two senecionid species (Senecio leucadendron = Pladaroxylon leucadendron; and Senecio redivivus = Lachanodes arborea , formerly Lachanodes prenanthiflord) also show paedomorphic wood. Wood of the various St Helena Asteraceae is consonant with relationship to African or South American ancestors that reached St Helena via long distance dispersal. Derivation from genera of Pacific islands or Austromalesian regions is considered less likely. However, DNA evidence is needed to clarify origins, times of colonization on St Helena and divergence from closest relatives, and the nature of evolutionary patterns.     

Functional and ecological xylem anatomy

Cohesion-tension transport of water is an energetically efficient way to carry large amounts of water from the roots up to the leaves. However, the cohesion-tension mechanism places the xylem water under negative hydrostatic pressure (P_x), rendering it susceptible to cavitation. There are conflicts among the structural requirements for minimizing cavitation on the one hand vs maximizing efficiency of transport and construction on the other. Cavitation by freeze-thaw events is triggered by in situ air bubble formation and is much more likely to occur as conduit diameter increases, creating a direct conflict between conducting efficiency and sensitivity to freezing induced xylem failure. Temperate ring-porous trees and vines with wide diameter conduits tend to have a shorter growing season than conifers and diffuse-porous trees with narrow conduits. Cavitation by water stress occurs by air seeding at interconduit pit membranes. Pit membrane structure is at least partially uncoupled from conduit size, leading to a much less pronounced trade-off between conducting efficiency and cavitation by drought than by freezing. Although wider conduits are generally more susceptible to drought-induced cavitation within an organ, across organs or species this trend is very weak. Different trade-offs become apparent at the level of the pit membranes that interconnect neighbouring conduits. Increasing porosity of pit membranes should enhance conductance but also make conduits more susceptible to air seeding. Increasing the size or number of pit membranes would also enhance conductance, but may weaken the strength of the conduit wall against implosion. The need to avoid conduit collapse under negative pressure creates a significant trade-off between cavitation resistance and xylem construction cost, as revealed by relationships between conduit wall strength, wood density and cavitation pressure. Trade-offs involving cavitation resistance may explain the correlations between wood anatomy, cavitation resistance, and the physiological range of negative pressure experienced by species in their native habitats.     

Leaf and twig anatomy of Eriope, a xeromorphic genus of Labiatae

The leaf and twig anatomy of 25 species of the genus Eriope were studied. The twig anatomy is very uniform apart from the level of formation of early layers of cork. Leaf anatomy shows considerable variation between the species, and this is correlated to some extent with the extreme habit range from trees to woody herbs. Characters of the lamina that show variation are: trichome type and frequency, cuticular markings, leaf dorsiventral or isobilateral, presence of adaxial stomata, presence of a hypodermis, number of layers of adaxial palisade mesophyll cells, occurrence of large bundles of phloem fibres at main veins, type of areolation and marginal venation. Petiole vasculature is simple and generally with either four distinct vascular bundles or two vascular arcs. The most xeromorphic species are usually woody herbs or sub-shrubs, and tend to have thick, isobilateral leaves with large bundles of phloem fibres and few hairs, or strongly dorsiventral leaves with a hypodermis and stomata in deep abaxial hair-lined depressions. The correlation of xeromorphic characters with environmental conditions is discussed. Leaf anatomy is of limited value in elucidating relationships within the genus.     

Floral anatomy and systematic position ofVivianiaceae

The floral anatomy of four species ofViviania has been studied. In the basic floral plan and essential floral anatomical featuresViviana closely resembles theGeraniaceae. Evidence from vegetative and floral anatomy, ultrastructural studies on phloem as well as phytochemistry supports geranialean affinity ofViviania; the placement within thePittosporales sensuHutchinson being unnatural.     

Leaf anatomy and chloroplast ultrastructure of Mn-deficient orange plants

Leaf samples of Mn-deficient and Mn-sufficient (control) ‘Navelate’ orange plants grown in a greenhouse were taken to investigate the effects of Mn deficiency in leaf structure and chloroplast ultrastructure. Total leaf chlorophyll concentration was significantly lower in Mn-deficient plants than in control ones. Entire lamina thickness was not altered due to Mn deficiency. However, Mn deficiency resulted in disorganization of mesophyll cells, mainly of palisade parenchyma cells. The number of mesophyll chloroplasts per cellular area and their length were both affected negatively. The membranous system of chloroplasts was also disorganized. The percentages of starch grains and plastoglobuli per chloroplast of Mn-deficient leaves were significantly greater than those of control leaves.     

Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics

CARLQUIST, S., 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Qualitative and quantitative data, based mostly upon liquid-preserved specimens, are presented for Akebia, Roquila, Decaisnea, Holbodia, Lardizabala, Sinofranchetia and Stauntonia . Because Decaisnea is a shrub whereas the other genera are vines, anatomical differences attributable to the scandent habit can be considered. These include exceptionally wide vessels, a high proportion of vessels to tracheids (or other imperforate trdcheary elements) as seen in transection, simple perforation plates, multiseriate rays which are wide and tall, and pith which is partly or wholly sclerenchymatous. With respect to ecology, two features are discussed: spirals in narrower vessels may relate to adaptation to freezing in the species of colder areas, and crystalliferous sclereids seem adapted in morphology and position to deterrence of phytophagous insects or herbivores. The wood may provide mechanisms for maintaining conduction even if wider vessels are deactivated temporarily by formation of air embolisms. Wood and stem anatomy of Lardizabalaceae compare closely to those of Berberidaceae and of Clematis (Ranunculaceae), as well as to other families of Berberidales. Decaisnea is more primitive than these in having consistently sralariform perforation plates and in having scalariform pitting on lateral walls of vessels. A tentative listing of anatomical features which may correspond to generic limits is given.     

Comparative wood anatomy of epacrids (Styphelioideae,Ericaceae s.L.)

The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel-ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze-thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.     

Gynoecial vascular anatomy and its systematic implications in Celtidaceae and Ulmaceae (Urticales)

Vasculature in the bicarpellate, pseudomonomerous gynoecium with two distinct styles is examined and compared in all of 15 genera of Celtidaceae and Ulmaceae (Urticales). Gynoecial vasculature is diversified in the families but consistet in a genus or a group of genera. Our observations corroborate the earlier suggestion that Ulmaceae (six genera) basically have three-bundled styles, while Celtidaceae (nine genera) always have one-bundled styles. Comparisons with other Urticales and with Eucommiaceae as an outgroup of Urticales indicate that Celtidaceae are more closely related to Moraceae in sharing one-bundled style (a synapomorphy), rather than to Ulmaceae. This supports a separation of Celtidaceae as a distinct family from Ulmaceaesensu lato. Based on the degree of fusion of major vascular bundles running in the gynoecium, we further distinguish three types of gynoecial vasculature in Ulmaceae and, likewise, three types in Celtidaceae, and discuss evolutionary trends of gynoecial vasculature as well as some generic relationships within the families.     

Leaf anatomy of the subtribe Hyptidinae (Labiatae)

The leaf anatomy of the subtribe Hyptidinae (Labiatae), which consists of four small genera and the largr genus Hyplis , is dercribed. The leaves may be dorsiventral or isobilateral. Variable characters of the lainina include: frequency and forms of trichomes, cuticular markings, presence of adaxial stoinata, thickness of leaf, thickness of adaxial epidermis, presence of a hypodermis, occurrence of sclcrified tissues (especially sclerified bundle sheath extensions, phloem and xylem fibres), mesophyll structure and venation pattern. Petiole vasculature varies from simple to complex, sometimes with medullary traces present. Most of the variable characters are related to xeromorphy and are tax-onomically useful within the framework of the present classification.     

Floral development and anatomy of Dirachma socotrana, (Dirachmaceae): a controversial member of the Rosales

The floral development and anatomy of Dirachma has been investigated with SEM and LM to discuss the relationship of Dirachmaceae with putative sister taxa (e.g. Rhamnaceae, Malvaceae, Barbeyaceae, Elaeagnaceae) on the basis of morphological synapomorphies. Flowers are initiated in axillary position on terminal branches. An epicalyx consisting of six to eight bracteoles surrounds the valvate calyx. Petal initiation is strongly retarded and primordia arise independently opposite the stamen primordia. The carpels arise independently and become weakly fused at the base; a single ovule develops in an axile position. A hypanthium develops by interprimordial growth between petals and stamens. Nectaries arise in a pouch and are covered with trichomes positioned on a protuberance at the base of the petal. It is suggested that the solitary flowers are derived from compound cymose inflorescences. Comparison between Dirachma and species of Rhamnaceae demonstrates striking similarities in floral structure and anatomy. However, several characters are unique to Dirachma and support a family Dirachmaceae separate from Rhamnaceae.We thank Frieda Christie for technical assistance with the SEM and LM preparations. We very much appreciate the constructive comments of Peter Endress and Paula Rudall.     

On caudal prehensility and phylogenetic constraint in lizards: The influence of ancestral anatomy on function in Corucia and Furcifer

We examined caudal anatomy in two species of prehensile‐tailed lizards, Furcifer pardalis and Corucia zebrata. Although both species use their tails to grasp, each relies on a strikingly different anatomy to do so. The underlying anatomies appear to reflect phylogenetic constraints on the consequent functional mechanisms. Caudal autotomy is presumably the ancestral condition for lizards and is allowed by a complex system of interdigitating muscle segments. The immediate ancestor of chameleons was nonautotomous and did not possess this specialized anatomy; consequently, the derived arrangement in the chameleon tail is unique among lizards. The limb functions as an articulated linkage system with long tendinous bands originating from longitudinal muscles to directly manipulate vertebrae. Corucia is incapable of autotomy, but it is immediately derived from autotomous ancestors. As such, it has evolved a biomechanical system for prehension quite different from that of chameleons. The caudal anatomy in Corucia is very similar to that of lizards with autotomous tails, yet distinct differences in the ancestral pattern and its relationship to the subdermal tunic are derived. Instead of the functional unit being individual autotomy segments, the interdigitating prongs of muscle have become fused with an emphasis on longitudinal stacks of muscular cones. The muscles originate from the vertebral column and a subdermal collagenous tunic and insert within the adjacent cone. However, there is remarkably little direct connection with the bones. The muscles have origins more associated with the tunic and muscular septa. Like the axial musculature of some fish, the tail of Corucia utilizes a design in which these collagenous elements serve as an integral skeletal component. This arrangement provides Corucia with an elegantly designed system capable of a remarkable variety of bending movements not evident in chameleon tails. J. Morphol. 239:143–155, 1999. © 1999 Wiley‐Liss, Inc.     

Evolution of secondary metabolites from an ecological and molecular phylogenetic perspective

Secondary metabolites, at least the major ones present in a plant, apparently function as defence (against herbivores, microbes, viruses or competing plants) and signal compounds (to attract pollinating or seed dispersing animals). They are thus important for the plant's survival and reproductive fitness. Secondary metabolites therefore represent adaptive characters that have been subjected to natural selection during evolution. Molecular phylogenies of the Fabaceae, Solanaceae and Lamiaceae were reconstructed and employed as a framework to map and to interpret the distribution of some major defence compounds that are typical for the respective plant families; quinolizidine alkaloids and non-protein amino acids for legumes; tropane and steroidal alkaloids for Solanaceae, and iridoids and essential oils for labiates. The distribution of the respective compounds appears to be almost mutually exclusive in the families studied, implying a strong phylogenetic and ecological component. However, on a closer look, remarkable exceptions can be observed, in that certain metabolites are absent (or present) in a given taxon, although all the neighbouring and ancestral taxa express (or do not express, respectively) the particular trait. It is argued that these patterns might reflect differential expression of the corresponding genes that have evolved earlier in plant evolution. The inconsistent secondary metabolite profiles mean that the systematic value of chemical characters becomes a matter of interpretation in the same way as traditional morphological markers. Thus, the distribution of secondary metabolites has some value for taxonomy but their occurrence apparently reflects adaptations and particular life strategies embedded in a given phylogenetic framework.     

Systematic anatomy of Oceanopapaver, a monotypic genus of the Gapparaceae from New Caledonia

Evidence from vegetative anatomy, reproductive morphology, and palynology does not support a relationship of Oceanopapaver with Cistaceae, Cruciferae, Flacourtiaceae, Papaveraceae, and Tiliaceae, but suggests placement of the genus in Capparaceae. The apparent occurrence of myrosin cells, among other features, effectively excludes all of the aforenoted families except Capparaceae and Cruciferae. However, multicellular non-glandular trichomes, bracteate inflorescences, sepals and petals each occasionally other than four per flower, presence of an androgynophore, numerous stamens, tricolporate and binucleate pollen, the unilocular mature ovary, the stipitate fruit, and the exotegmic seed in Oceanopapaver favour Capparaceae over Cruciferae. Floral histology and vasculature provide no clues about the relationships of Oceanopapaver. A few features are anomalous, most notably the presence of secretory canals and secretory cells in the genus versus their absence in Capparaceae and their rarity in Cruciferae, the trichomic floral nectary in the genus versus the massive, non-trichomic nectaries in these two families, and the straight embryo in the genus versus the more or less curved or folded embryo in the two families. The fleshy endosperm in Oceanopapaver has counterparts in a few Capparaceae, contrary to previous claims that endosperm is absent or scanty in this family. The report of stamen fascicle traces for Oceanopapaver is the first for Capparaceae, but these should be sought elsewhere in the family. Within Capparaceae the genus fits best in Capparoideae compared to Cleomoideae or the nine other very restrictive subfamilies variously proposed for Capparaceae. There is no justification for the monotypic segregate Oceanopapaveraceae. The phylogenetic and functional anatomy of vegetative and reproductive structures is discussed.