Leaf architecture of the genus Didymaea Hook. f. (Rubiaceae)

We examined 532 specimens representing eight species of Didymaea (Rubiaceae) to describe their leaf architecture. Characters observed in the herbarium specimens and cleared leaves were leaf base, apex, and margin; trichome type and distribution; vein orders from primary to higher order veins, veinlets, and ultimate marginal vein. Six morphometric variables were analyzed using variance and principal component analysis to identify which characters are useful to distinguish species. Most of the species present brochidodromous venation pattern, except D. hispidula, which has eucamptodromous venation pattern; Didymaea floribunda is unique in having cylindrical and bicellular trichomes. Principal component analysis revealed that 78% of the total variation is mainly related to leaf maximum width. Variance analysis showed significant differences between D. microflosculosa and the other species. Architectural features were useful for differentiating all the species, and an identification key for the species of Didymaea based only on leaf characters is presented.     

Venation patterns of Bursera species Jacq. ex L. (Burseraceae) and systematic significance

The taxonomy of the genus Bursera has been traditionally supported by morphological characters such as fruit form, bark and leaves. Nonetheless, leaf architecture, another important source of phylogenetic information, has not been well studied in this group. For this purpose, venation patterns of 30 species of the genus Bursera in Mexico were studied. Clarified leaves allowed finding four types of venation patterns: brochidodromous and eucamptodromous in complex B. simaruba (section Bursera); a cladodromous pattern in fragilis, microphylla and fagaroides groups (herein informally called group Quaxiotea (section Bursera), and semicraspedodromous in section Bullockia species. Only bipinnated species of section Bullockia showed a brochidodromous venation pattern. Furthermore, section Bullockia and complex Simaruba presented reticulate tertiary veins, whereas the group of Quaxiotea species presented a ramified pattern in the tertiary veins and tracheoblasts.     

贵州小檗属植物(小檗科)叶脉序研究

采用制作叶脉标本和透明叶标本的方法,对贵州产28种2变种小檗属植物叶脉特征进行比较研究。结果表明:贵州小檗属植物的脉序类型有5种:半达缘羽状脉、花环状半达缘羽状脉、简单弓形羽状脉、花环状弓形羽状脉和混合型。叶脉分支一般有五级:1一级脉构架均为羽状脉,粗度有很粗、粗、中等粗细和纤细四种类型,分支方式包括单轴分支和合轴分支;2粗二级脉构架中有分支达缘或分支均不达缘,与中脉夹角变化各异,内二级脉存在或缺失,细二级脉半达缘、真曲行或简单弓形,间二级脉类型复杂多变但频度种间有差异;3三级脉贯串型、结网型或分支型;4四、五级脉网状或自由分支且常混合在一起。脉间区从发育差到良好,小脉从不分支到不均等分支等各种类型均有,叶缘末级脉缺失、不完整、钉状和环状。大部分种类叶缘具齿,每1cm齿数目和齿内腺点的特性等特征在不同种类间有区别,具有鉴定价值,但齿其它特征复杂多变或种间区别较小,同时齿内脉性状也不稳定。此外,齿的有无会对脉序类型产生影响。小檗属植物叶脉类型存在种间差异,具有重要的分类学价值,叶脉类型的变化和复杂程度显示了该属植物的进化特点;叶齿的有无和齿特征具有分类学和系统学意义。基于叶脉特征的研究结果并结合重要的外部形态学特征编制了贵州小檗属植物的分种检索表。研究结果可为小檗属植物分类寻找新的依据并探讨其系统学意义。     

中国吴茱萸属(广义)叶结构特征及其分类学意义北大核心CSCD

该研究以中国产芸香科广义吴茱萸属18种3变种为材料,于体视显微镜下观察叶结构特征。结果发现:(1)吴茱萸属叶脉为羽状脉。(2)二级脉有花环状弓形脉和简单弓形脉两种类型,极少数存在内二级脉,间二级脉存在或缺失。(3)三级脉不规则网状或极少数弱贯穿,边缘三级脉环状。(4)四级脉不规则网状或自由分支。(5)五级脉不规则网状或自由分支,脉间区发育差或中等,游离端小脉末端简单或管状异形,边缘末级脉大多数不完整,极少数环状。(6)基于重要的外部形态学特征和叶结构特征观察结果,编制了广义吴茱萸属植物的分组检索表。研究表明,吴茱萸属这些类群的二级脉与更高级脉序形成的结构极为稳定,同时又存在种间差异,故广义吴茱萸属植物叶结构特征可以为更准确地鉴定一些疑难种和混淆种提供佐证,具有重要的系统分类学价值。     

Foliar architecture and anatomy of <Emphasis Type="Italic">Bernardia</Emphasis> and other genera of Acalyphoideae (Euphorbiaceae)

A comparative study of leaf architecture and anatomy of 42 species of Bernardia and other genera related of Acalyphoideae was undertaken to identify characters that support infrageneric and specific delimitation. Thirty variable foliar architectural and anatomical characters were studied. Some characters are consistent (e.g., venation pattern, secondary and tertiary vein arrangement, presence or absence of agrophic veins, type of areoles, bundle sheath cell type, presence or absence of fibers in the mesophyll, trichome type, stomata location, and type of crystals), and characterize genera. In addition, foliar character distribution within Bernardia supports the most recent infrageneric classification.     

Los domacios de Mortoniodendron (Malvaceae s.l.)

We describe the domatia of nine species of the genus Mortoniodendron, based on observations of cleared leaves and scanning electron micrographs. We recognize four kinds of domatia in Mortoniodendron species: cryptpocket, crypt, pocket and tufts of hairs, mostly associated with primary, secondary and tertiary veins. Mortoniodendron anisophyllum and M. costaricense have domatia up to the tertiary and quaternary veins. Moreover, domatia are absent in only two species, M. pentagonum and M. uxpanapense. Although there are reports on the occurrence of domatia in Malvaceae s.l., mainly in Brownlowioideae, Sterculioideae, and Tilioideae subfamilies, they are poorly described, and based on the different types of domatia found in Mortoniodendron we suggest reviewing other genera to better describe the diversity of domatia in Malvaceae s.l.

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Resumen  Se describen los domacios presentes en nueve especies del género Mortoniodendron utilizando la técnica de diafanización y el microscopio electrónico de barrido. En el género se observaron cuatro tipos de domacios: criptabolsa, cripta, bolsa y mechón de tricomas, la mayoría asociados a las venas primaria, secundarias y terciarias. Mortoniodendron anisophyllum y M. costaricense tienen domacios hasta las bifurcaciones de venas terciarias y cuaternarias. únicamente en dos especies, M. pentagonum y M. uxpanapense, los domacios están ausentes. A pesar de que se han registrado domacios en miembros de Malvaceae s.l., predominantemente en las subfamilias Brownlowioideae, Sterculioideae y Tilioideae; pocas veces se describe su tipo. Por ello, con base en los diferentes tipos de domacios encontrados en Mortoniodendron sugerimos estudiar otros géneros y confirmar la diversidad en Malvaceae s.l.

     

Leaf Architecture of Some Monocotyledons with Reticulate Venation

Twelve species belonging to seven monocotyledonous families:Hydrocharitaceae, Taccaceae, Dioscoreaceae, Smilacaceae, Araceae,Alismataceae and Aponogetonaceae show reticulate venation typicalof dicotyledons. A study of the leaves of these species showsthat venation patterns are usually curvipalmate-convergate,occasionally rectipalmate or collimate, and rarely pinnate lyratetype. Number, size and shape of areoles, number of primaries,number of secondaries along one side of the primaries, anglebetween 1 and 2° veins and number of vein endings per areoleare given for each species. Intesecondary veins, isolated tracheids,loops, extension cells, raphide and raphide idioblasts and terminaltracheids were observed. Marginal ultimate venation is mostlyarcuate. Major and minor veins are jacketed by parenchymatousbundle sheath cells. The lamina of Tacca leontopetaloides, Colocasiaesculenta and Scindapsus aureus show a single midrib-like centralregion similar to that of dicotyledonous leaves, and it is multistrandedin Aponogeton natans, Limnophyton obtusifolium and Ottelia alismoides.The degree of vein order is most commonly up to fourth or fifthand rarely up to sixth in Dioscorea hispida. Monocotyledons, leaf architecture, vein endings, venation, areoles     

中国水青冈属 （壳斗科） 叶结构及分类学意义

叶结构对壳斗科(Fagaceae)现存植物和化石的鉴定具有重要意义。通过对水青冈属5种植物叶结构特征进行细致的研究,结果发现水青冈属植物叶脉有羽状弓形脉、羽状半达缘脉两种类型;三级脉有波状对生贯穿、互生贯穿及混合贯穿三种类型;小脉缺失、简单无分支或一次分支;脉间区发育良好,网眼有三边形、四边形和五边形三种类型,排列规则;具齿种类叶齿由齿主脉和齿侧脉构成,齿侧脉环状。研究结果表明水青冈属二级脉与更高级脉序形成的结构稳定且存在种间差异,具重要分类学价值。基于水青冈属叶结构特征观察结果,本次研究编制了水青冈属植物的分种检索表;参照已有研究结果并结合重要外部形态学特征,编制了壳斗科相关类群分属检索表。     

中国无患子科植物的叶脉形态及其系统学意义

对国产狭义无患子科25属27种植物的叶脉形态特征进行了研究报道。结果表明：叶脉均属于羽状脉类型,其中多数为曲行羽状脉,部分为直行羽状脉;叶缘有全缘、具齿和深裂3种类型;二级叶脉具有分支和不分支两种类型;大部分种类具二级间脉,少数不具间脉或间脉不明显;多数种类的三级脉为结网型和贯串型并存;网眼的发育有完善和不完善2种类型;盲脉有简单、具分支和无盲脉3种类型。叶脉形态研究结果支持文冠果亚科以及广义鳞花木属概念,观察发现龙眼属、荔枝属与韶子属从脉序特征方面表现出较近的亲缘关系。编写了国产无患子科叶片脉序特征检索表。     

Mentzelia canyonensis (Loasaceae), a new species endemic to the Grand Canyon, Arizona, U.S.A.

We describe Mentzelia canyonensis, a new species endemic to the Grand Canyon. Based on floral and fruit forms, shoot system architecture, and phylogenetic analyses of nuclear ribosomal DNA, we infer that M. canyonensis belongs to the ‘subshrubby’ clade of Mentzelia section Bartonia, which includes the morphologically similar M. hualapaiensis, M. oreophila, M. puberula, and M. tiehmii. The new species is most similar to M. hualapaiensis, which is also known only from the Grand Canyon. These two species share leaf shape, subshrubby habits, corolla color, and fruit shape, although they differ in staminode presence and especially in leaf and corolla size. Principal component analyses of variation among floral and vegetative traits demonstrates that M. canyonensis and M. hualapaiensis share highly overlapping floral character space, but M. canyonensis occupies unique vegetative character space compared to similar species. A dichotomous key is provided to assist with identification of, and differentiation among, species of Mentzelia section Bartonia that occur in the Grand Canyon region.     

Leaf Venation and Its Systematic Significance in Sapindaceae of China

Leaf venation of 27 species representing 25 genera of Sapindaceae (sstr.) of China was investigated for the first time. The pinnate venation pattern in most species is either camptodromous, or craspedodromous. Three types of leaf blade margin were observed, ie., entire, toothed and partite. The secondary veins are branched or unbranched. Most species have intersecondary veins. The tertiary veins of most species are reticulate and percurrent. The areoles are regular or irregular. Veinlets are simple, branched or absent. The delimitations of Xanthoceroideae and Lepisanthes sensu lato are supported by leaf venation characters. The close relationships among Dimocarpus, Litchi and Nephelium are supported by the evidence from leaf venation. A key to the species of Sapinaceae based on leaf venation characters is presented.     

中国水青冈属（壳斗科）叶结构及分类学意义

叶结构对壳斗科（Fagaeeae）现存植物和化石的鉴定具有重要意义。通过对水青冈属5种植物叶结构特征进行细致的研究,结果发现水青冈属植物叶脉有羽状弓形脉、羽状半达缘脉两种类型;三级脉有波状对生贯穿、互生贯穿及混合贯穿三种类型：小脉缺失、简单无分支或一次分支：脉间区发育良好,网眼有三边形、四边形和五边形三种类型,排列规则;具齿种类叶齿由齿主脉和齿侧脉构成,齿侧脉环状。研究结果表明水青冈属二级脉与更高级脉序形成的结构稳定且存在种问差异．具重要分类学价值。基于水青冈属叶结构特征观察结果,本次研究编制了水青冈属植物的分种检索表：参照已有研究结果并结合重要外部形态学特征,编制了壳斗科相关类群分属检索表。     

Quantitative neutron imaging of water distribution,venation network and sap flow in leaves

Main conclusion

Quantitative neutron imaging is a promising technique to investigate leaf water flow and transpiration in real time and has perspectives towards studies of plant response to environmental conditions and plant water stress. The leaf hydraulic architecture is a key determinant of plant sap transport and plant–atmosphere exchange processes. Non-destructive imaging with neutrons shows large potential for unveiling the complex internal features of the venation network and the transport therein. However, it was only used for two-dimensional imaging without addressing flow dynamics and was still unsuccessful in accurate quantification of the amount of water. Quantitative neutron imaging was used to investigate, for the first time, the water distribution in veins and lamina, the three-dimensional venation architecture and sap flow dynamics in leaves. The latter was visualised using D₂O as a contrast liquid. A high dynamic resolution was obtained by using cold neutrons and imaging relied on radiography (2D) as well as tomography (3D). The principle of the technique was shown for detached leaves, but can be applied to in vivo leaves as well. The venation network architecture and the water distribution in the veins and lamina unveiled clear differences between plant species. The leaf water content could be successfully quantified, though still included the contribution of the leaf dry matter. The flow measurements exposed the hierarchical structure of the water transport pathways, and an accurate quantification of the absolute amount of water uptake in the leaf was possible. Particular advantages of neutron imaging, as compared to X-ray imaging, were identified. Quantitative neutron imaging is a promising technique to investigate leaf water flow and transpiration in real time and has perspectives towards studies of plant response to environmental conditions and plant water stress.     

Leaf architectural studies in the brassicaceae

The leaf architecture has been studied in 19 genera and 35 species of the Brassicaceae. The major venation pattern is pinnate craspedodromous with the exception ofAlyssum maritimum, Iberis amara, I. umbellata andMalcolmia maritima where it is pinnate-festooned brochidodromous. The number of secondary veins and their angle of divergence vary from species to species and even within the same species. Marginal ultimate venation is mostly looped occasionally incomplete. The areole size and shape is variable. The veinlets may be simple or once or twice dichotomously divided. The simple veinlets may be curved or hooked. The tracheids are either uni-, bi-, tri- or multiseriate and extraordinarily variable in size and shape. Isolated vein endings, isolated free vein endings, isolated tracheids and extension cells are observed. Myrosin cells have been observed inEruca vesicaria ssp.sativa andCochlearia cochlearioides.     

A morpho-anatomical characterisation of Myrothamnus moschatus (Myrothamnaceae) under the aspect of desiccation tolerance

Morpho-anatomical traits of the rarely studied dicotyledonous desiccation-tolerant shrub Myrothamnus moschatus were examined and compared for the first time to Myrothamnus flabellifolius under the aspect of desiccation tolerance. Both species almost exclusively occur on rock outcrops and differ mainly in their geographic range and leaf morphology (fan-shaped in M. flabellifolius, lanceolate in M. moschatus) but have a very similar leaf and wood anatomy, except for the lack of hydathodes in M. moschatus. Both species adopt the parallel leaf venation of monocots, although this is more pronounced in M. moschatus. This provides a mechanical and protective advantage over the net venation pattern of most dicots and facilitates the reversible, drought-induced, accordion-like leaf contraction. The sclerenchyma, as a stabilising tissue, is mainly confined to vascular bundles in leaves of both species. Here, mechanical support seems to be less crucial for survival in long periods of drought than other morpho-anatomical traits (e.g. parallel leaf venation).     

Defining the gigantopterid concept: a reinvestigation of Gigantopteris (Megalopteris) nicotianaefolia Schenck and its taxonomic implications

Reinvestigation of the first described gigantopterid plant, Gigantopteris (Megalopteris) nicotianaefolia Schenk ex Potonié, 1902 from the Permian of China, has been conducted using the original specimens documented in 1883. The application of new techniques has permitted a comprehensive review of their morphology. This study represents the first photographic record of Schenk's specimens and provides a standardised terminology to describe them. This species of Palaeozoic leaf megaphyll is characterised by its pinnate venation, with tertiary veins that anastomose, and give rise to branches that may be of the same or higher order. These anastomosed branches form a reticulum of irregular polygonal meshes, within which higher order veins anastomose and again form meshes. On examination of Schenk's original specimens only one, MB.Pb.2002/989, can be assigned to Gigantopteris nicotianaefolia, and this is herein designated the lectotype. The remaining specimens described by Schenk are reassigned to cf. G. nicotianaefolia, cf. Gigantopteris sp., Gigantonoclea sp. or cf. Gigantonoclea sp. This re‐analysis of Schenk's specimens has led to an emended diagnosis of both the genus Gigantopteris and the species G. nicotianaefolia. Current evidence suggests that Gigantopteris nicotianaefolia is restricted to the southern floral regions of Cathaysia, as typically are other species of this genus. Fertile specimens of Gigantopteris are unknown, and the status of these plants as a natural group relies upon a combination of their complex foliar physiognomy, stratigraphic range, and geographical distribution. This study indicates that the genera GigantopterisGigantonoclea form the basis of the gigantopterid concept. These two genera share characters of leaf morphology including megaphylls with eucamptodromous venation across which extends a continuous lamina, and higher order veins, third order or above, that arise from the secondary veins and anastomose to form complex meshes. These genera are distinguished in that the venation of Gigantopteris is far more complex than that of Gigantonoclea, with a greater number and complexity of vein orders, the penultimate and ultimate of which form meshes within meshes, and of these the finest may terminate in blind endings.     

Evolution and Function of Leaf Venation Architecture: A Review

The leaves of extant terrestrial plants show highly diverseand elaborate patterns of leaf venation. One fundamental featureof many leaf venation patterns, especially in the case of angiospermleaves, is the presence of anastomoses. Anastomosing veins distinguisha network topologically from a simple dendritic (tree-like)pattern which represents the primitive venation architecture.The high degree of interspecific variation of entire venationpatterns as well as phenotypic plasticity of some venation properties,such as venation density, indicate the high selective pressureacting on this branching system. Few investigations deal withfunctional properties of the leaf venation system. The interrelationshipsbetween topological or geometric properties of the various leafvenation patterns and functional aspects are far from beingwell understood. In this review we summarize current knowledgeof interrelationships between the form and function of leafvenation and the evolution of leaf venation patterns. Sincethe functional aspects of architectural features of differentleaf venation patterns are considered, the review also refersto the topic of individual and intraspecific variation. Onebasic function of leaf venation is represented by its contributionto the mechanical behaviour of a leaf. Venation geometry anddensity influences mechanical stability and may affect, forexample, susceptibility to herbivory. Transport of water andcarbohydrates is the other basic function of this system andthe transport properties are also influenced by the venationarchitecture. These various functional aspects can be interpretedin an ecophysiological context. Copyright 2001 Annals of BotanyCompany Review, leaves, leaf venation, evolution, network, transport, flow, mechanical stabilization     

A Miocene leaf fossil record of Rosa (R. fortuita n. sp.) from its modern diversity center in SW China

Roses (Rosa, Rosaceae) are arguably the most admired ornamental plants in the world. Southwestern China is the center of diversity for many extant native species of Rosa and fossils found in this region are critical for understanding the evolution of this genus. Herein, we report a leaf fossil record with good preservation from the late Miocene of Yunnan Province. The opposite and odd-pinnate leaf is composed of seven elliptical leaflets, with close, crenulate, and regular marginal teeth. The stipules are lanceolate and adnate to the petiole. Additionally, the secondary veins are semicraspedodromous, showing the same venation pattern as most living Rosa species in southwestern China. On the basis of the extensive morphological comparisons, we propose a new species, R. fortuita T. Su et Z.K. Zhou n. sp. This is the first confirmed Rosa leaf fossil record in China, and the discovery of R. fortuita n. sp. indicates that Rosa existed in southwestern China by the late Miocene. It suggests that Rosa was distributed in subtropical or temperate forests and shared a similar ecological niche as Rosa in Europe during the Oligocene and Miocene. The modern diversification of Rosa in southwestern China is thought to have been closely associated with the continuous uplift of the Qinghai-Tibet Plateau since the late Miocene, creating complex topography and a variety of climate conditions.     

Anatomy and systematic position of Uldinia (Umbelliferae)

Vegetative, floral and fruit anatomy and morphology ofUldinia, a monotypic hydrocotyloid genus endemic to Australia, are described in detail. These observations, when compared to available reports for other genera of Hydrocotyloideae, indicate thatUldinia is most closely allied to members of Hydrocotyleae—Hydrocotylinae, and, in particular, toTrachymene. Similarities between these two genera include such anatomical and morphological characteristics as habit and leaf morphology, stem anatomy, trichome anatomy, inflorescence morphology, floral morphology, and several details of the floral venation pattern and fruit anatomy. Nevertheless, the nature and development of the fruit wings, the papillae and retrorse barbs of the fruit, the absence of a carpophore, the differences in floral venation at the commissure and in the upper portion of the mericarp, and the orientation of fibers in the endocarp, all provide a basis for the generic distinctness ofUldinia. These comparative studies also point to the need for a possible re-evaluation of relationships within the subtribes of Hydrocotyleae.