Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects

CARLQUIST, S., 1991. Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects. Quantitative and qualitative data are given for 60 species of the 12 genera of Bruniaceae; most data are based on liquid-preserved material. Leaves of Bruniaceae are basically linear (broader forms are probably derived) with an apicula that contains phellogen activity. Most bruniaceous leaves have some degree of isolateral construction, with transition to normal bifacial construction in a few species, but more commonly transition to 'inverse' bifacial structure (stomata on adaxial face, palisade on abaxial face). The latter type is correlated with the tendency for leaves to be appressed to stems. Tannins and very likely other dark-staining materials are very characteristic of mesophyll cells. Six genera have a large strand of fibres on the midvein and rhomboidal crystals in bundle sheath cells. The other six genera have few or no fibres on veins and have druses in mesophyll cells (but not in bundle sheath cells). These distinctions may relate to intrafamilial taxonomy, but they also support the primitive position usually accorded to Audouinia, Thamnea and Tittmannia. A key to genera based on leaf antomy is offered. Details of epidermal cell shape, cuticular relief and trichome form and structure based on scanning electron microscopy are given. Leaf anatomy, combined with other features, favours a relationship between Bruniaceae and Grubbiaceae in particular and in broader contexts allies Bruniaceae to rosalean and possibly hamamelidalean families.     

Leaf anatomy of Desmos and Dasymaschalon (Annonaceae) from China in relation to taxonomic significance

Leaf morphology in four species of Desmos and three species of Dasymaschalon was comparatively studied using scanning electron microscopy (SEM) together with epidermal maceration and paraffin methods. The results showed that there were some remarkable foliar anatomical differences between Desmos and Dasymaschalon. In leaves of Desmos, some of the adaxial epidermal cells were enlarged into globose cells each containing one large cluster crystal, while other epidermal cells were normal without any crystal, and in abaxial epidermis each cell contained one smaller cluster crystal. The leaf structure was typically bifacial, and the mesophyll cells were differentiated into palisade tissue and spongy tissue. Oil cells were distributed in the second layer of palisade and the whole spongy tissue, and the number of oil cells per mm leaf width ranged from 4 to 6. The vascular tissue in the midrib was separated into bundles by parenchyma cells. In leaves of Dasymaschalon, all the adaxial epidermal cells contained one cluster crystal, and the crystal size was similar to that of thecrystals in abaxial epidermal cells. The leaf structure was more or less isobilateral. Oil cells were distributed only in the spongy tissue between the two layers of the palisade, and the number of oil cells per mm leaf width ranged from 2 to 3. The vascular tissue in the midrib formed a continuous circle. It is clear that the anatomical differences between Desmos and Dasymaschalon are remarkable, supporting the treatment of Desmos and Dasymaschalon as two independent genera.     

Leaf age effects on photosynthetic rate, transpiration rate and nitrogen content in a tropical dry forest

This study examines the effect of leaf age on photosynthesis, transpiration and nitrogen concentration in four deciduous (DC) and two evergreen (EG) species coexisting in a tropical dry forest of Venezuela. Leaf age was characterized on the basis of leaf chorophyll, nitrogen content, and construction and maintenance costs. The mean leaf area-based nitrogen concentration (N) in EG was about twice that in DC species. A leaf age effect was observed in both DC and EG species, with largest N concentration in mature leaves. Fractional leaf N allocation to chlorophyll was higher in the DC than in the EG species. Differences in the construction costs of leaf mass between the youngest and the oldest leaves averaged from 2.14 to 1.55 g glucose g⁻¹ dry weight. Although variation in area-based leaf maintenance and construction costs between DC and EG species existed, they were, nevertheless, positively correlated. Individual data sets, for each species, indicated that leaf N and maximum rate of photosynthesis (A_max) were linearly related. Nitrogen use efficiency (NUE) and water use efficiency (WUE) tended to be higher in mature leaves than in expanding and old leaves. Moreover, DC species always had higher NUE than EG species. Intercellular to ambient pressures of CO₂ (P_i/P_a) were related to WUE in a negative manner. Higher P_i/P_a values were observed in expanding and old leaves. Leaf age effect on photosynthesis was, therefore, due to greater decline of carbon fixation capacity by mesophyll tissue relative to the decline in stomatal conductance in youngest and oldest leaves.     

中国假鹰爪属和皂帽花属植物叶的形态结构及其分类学意义

利用扫描电镜技术、叶片离析法和石蜡切片法研究了假鹰爪属Desmos 4种植物和皂帽花属Dasy-maschalon 3种植物叶片的形态结构。结果表明：假鹰爪属植物叶片近轴面表皮具大型球状含晶簇细胞和不含晶簇的表皮细胞两种类型,远轴面表皮细胞均具一较小的晶簇;叶肉组织明显分化为栅栏组织细胞和海绵组织细胞,油细胞分布于第2层的栅栏组织和海绵组织内,单位毫米叶宽油细胞数为4～6个;主脉维管组织被薄壁细胞分隔成束状。皂帽花属植物叶片近轴面表皮细胞形状相同,均具一晶簇,远轴面表皮细胞的晶簇和近轴面表皮细胞的晶簇相似;靠近上、下表皮的叶肉组织均分化为栅栏组织细胞,在两层栅栏组织细胞之间分化为一至几层海绵组织细胞,油细胞分布于海绵组织内,单位毫米叶宽油细胞数为2～3个;主脉维管组织形成连续的环状。由此可见两属叶的结构具有明显的差异,因而支持假鹰爪属和皂帽花属为两个独立属的观点。     

LIRIODENDRON (MAGNOLIACEAE) FROM THE MIOCENE CLARKIA FLORA OF IDAHO

Miocene Liriodendron carpels, whole fruiting structures and leaves from Clarkia and Oviatt Creek sites in northern Idaho are preserved as imprints and compressed fossils in soft lacustrine clays. The isolated carpels are indistinguishable from those described as L. hesperia Berry from the Spokane Latah flora. Fruit aggregates from the type Clarkia and Oviatt Creek localities and leaves from three Clarkia sites are considered to be within the range of variation of the single species L. hesperia. Comparisons were made regarding leaf architecture, lower leaf epidermal structures, leaf flavonoid and steroid analysis, morphological features of receptacles and carpels, and the venation pattern of carpels of the fossil material to the two extant species, L. tulipifera L. (native to southeastern United States) and L. chinense Sarg. (native to southeastern Asia). Leaf architecture features analyzed by standard statistical and canonical tests and marginal venation patterns near the base of leaves suggest that L. hesperia is more similar to L. tulipifera, whereas the size dimensions of lower epidermal cells and the common presence of two sterane compounds imply that L. hesperia is more similar to L. chinense. The fossil species, however, is a distinct taxon indicated by statistical discriminant and canonical tests, leaf base shape, often smaller epidermal cell dimensions, and the shape of round receptacle carpel scars. Both the fossil and the two living Liriodendron species are associated with comparable mixed mesophytic floras.     

Epidermal Micromorphology and Mesophyll Structure of Populus euphratica Heteromorphic Leaves at Different Development Stages

Leaf epidermal micromorphology and mesophyll structure during the development of Populus euphratica heteromorphic leaves, including linear, lanceolate, ovate, dentate ovate, dentate rhombic, dentate broad-ovate and dentate fan-shaped leaves, were studied by using electron and light microscopy. During development of heteromorphic leaves, epidermal appendages (wax crystals and trichomes) and special cells (mucilage cells and crystal idioblasts) increased in all leaf types while chloroplast ultrastructure and stomatal characters show maximum photosynthetic activity in dentate ovate and rhombic leaves. Also, functional analysis by subordinate function values shows that the maximum adaptability to adverse stress was exhibited in the broad type of mature leaves. The 12 heteromorphic leaf types are classified into three major groups by hierarchical cluster analysis: young, developing and mature leaves. Mature leaves can effectively obtain the highest stress resistance by combining the protection of xerophytic anatomy from drought stress, regulation of water uptake in micro-environment by mucilage and crystal idioblasts, and assistant defense of transpiration reduction through leaf epidermal appendages, which improves photosynthetic activity under arid desert conditions. Our data confirms that the main leaf function is differentiated during the developing process of heteromorphic leaves.     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Effects of chilling stress on leaf morphology,anatomy, ultrastructure,gas exchange,and essential oils in the seasonally dimorphic plant Teucrium polium (Lamiaceae)

The Mediterranean region (and globally also other regions) is characterized by the presence of phryganic plants, i.e. subshrubs that grow under hot and arid environmental conditions. These plants are reported to be affected by summer drought stress. However, in the present study the phryganic plant Teucrium polium (mountain germander) appears to be affected by winter chilling stress rather than by summer drought stress in a specific area. Winter leaves of the plant are smaller and thicker compared to summer leaves, have more stomata and glandular hairs, and their chloroplasts are larger, more numerous, with voluminous starch grains. Moreover, epidermal and mesophyll cells of winter leaves contain in their vacuoles dark phenolics and calcium oxalate crystals. Summer leaves are devoid of vacuolar phenolics and their chloroplasts possess many large plastoglobuli. Leaf gas exchange parameters (photosynthesis, transpiration, stomatal conductance) are significantly higher in winter leaves. Concentrations of osmoprotectors (stress indicators) like proline and soluble sugars are similarly higher in winter leaves. Essential oil assessments showed a significantly higher oil yield of winter leaves compared to summer leaves. Percentages of the major oil components (linalool, terpinen-4-ol, germacrene D, and spathulenol) are remarkably higher in winter oils than in summer oils. In conclusion, low environmental temperatures (1–10 °C) appear to decisively influence the structure and function of winter leaves compared to summer leaves. Winter plants undergo chilling stress to which they respond by developing various mechanical and chemical defensive strategies.     

Taxonomy of Selaginella: a study of characters, techniques, and classification in the Hong Kong species

This study of 11 Hung Kong species of Selaginella indicates that sufficient characters exist by which they may be conveniently, consistently, and conclusively distinguished, and implies that the same is true for the rest of the genus, which suffers from chronic taxonomic confusion. Characters assessed are: leaf size, shape, epidermal cell patterns and mesophyll structure; ligule shape; patterns seen in stem transections; sporophyll shape; microsporangium shape; sporangial distribution; and spore surface ornamentation. Of these, the shape of median leaves, the maximum size of lateral leaves and epidermal cell patterns are the most significant and distinctive foliar features. These, together with spore surface ornamentation, are the most useful for distinguishing species. Correlations between the shapes of ligules, sporophylls, microsporangia, and vascular bundles indicate natural subgeneric groups.
These studies suggest that there are two ways by which taxonomic investigations of Selaginella can be improved. First leaves and spores should be prepared in permanent mountant (e.g. Hoyer's solution) for accurate observation of size, shape, and epidermal patterns. Secondly, data should be collected on a wide range of features.     

Leaf anatomy of the genus Psoralea sensu stricto (Psoraleeae, Papilionoideae, Leguminosae)

Leaf anatomy is described from 17 species of Psoralea sensu stricto which includes four species of the genus Hallia Thunberg. Species of the two genera share many characters including the presence of rod-shaped crystals, similar stipule structure and a possible continuum in secretory cavity anatomy ranging from a small and non-trabeculate to a large and trabci ulate form. Hallia species are distinguished b the presence of large tannin cells in the bundle sheaths and a narrow lcngllv.width ratio of palisade cells. In the light of other floral evidence this is considered insufficient to separate the two genera, a view confirmed by cluster analysis. Thus, Salter's proposal of 1939 for their amalgamation is supported. Leaf anatomy of herbaceous species is compared with that of seedlings of Psoralea sp. to assess the possibility that the herbaceous species may have arisen through neoteny from species that are trees or shrubs. Comparative venation studies between stipules and scale leaves suggest that the scale leaf form arose from pinnate-leaved ancestors by leaf reduction.     

Determinants of some leaf characteristics of Australian mangroves

Leaf characteristics reflecting the size, lifespan (longevity), moisture content (degree of succulence) and complexity of structure of 20 mangrove species were studied over several years at 13 locations along the tropical and subtropical Australian coast. These characteristics were found to fall generally within the ranges of those for woody species from other ecosystems. With the exception of one species, it was found that leaf longevity was related inversely to leaf moisture content, increasing from nearly 6 months in more succulent species to over 2 years in less succulent species. This suggested that more succulent leaves are less complex in their structure because they have less well‐developed ability to compartmentalize salt. There was a tendency also for leaf longevity to increase in species with larger leaves. These findings were consistent with the general view for land plants that leaf longevity is greater in species that have developed tolerance to environmental stress, salt particularly in the case of mangroves. Leaf tissue in such species is more robust or complex and requires greater metabolic resources in its construction; the plant is then advantaged by retaining the tissue for longer periods. Classification of the species considered here, based on their leaf longevity, moisture content and complexity, identified phylogenetically related species groupings that reflected these leaf longevity effects.     

Hepatics from Rovno amber (Ukraine), 6. Frullania rovnoi,sp. nov.

A fossil species of the extant liverwort genus Frullania Raddi is described and illustrated, based on a single inclusion in a piece of Rovno amber (Ukraine) that shares its age with Late Eocene Baltic amber, its northern contemporary. Frullania rovnoi is characterised by leaves with a rounded dorsal lobe and the absence of ocelli. The ventral lobe is inflated and forms a saclike lobule, which is bell-shaped and somewhat constricted above the mouth. The bifid underleaves have several blunt teeth or angulations along the shoulder. The Rovno fossil differs sufficiently from morphologically similar species preserved in Baltic and Bitterfeld amber as to be described as new to science. The shape of the lobules and underleaves, as well as the absence of ocelli, indicate an affiliation to F. sect. Australes, hitherto represented in Eocene amber inclusions solely by F. schumannii (Casp.) Grolle. The Rovno fossil is distinguished from extant species of F. subg. Australes and from F. schumannii by having roughly and irregularly dentate-angulate underleaf margins.     

Seasonal changes in foliar macronutrients (N,P, K,Ca and Mg) inEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mine soils of the Darling Range,Western Australia

Summary Seasonal changes in the foliar concentration of macronutrients (N, P, K, Ca and Mg) in sapling trees ofEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mined areas in the Darling Range of Western Australia are described. Foliar N concentration decreased with age of the fully expanded leaf tissue. Leaf N concentrations were also high when rates of litter decomposition were expected to be high during the period of early spring. The greatest foliar N difference between trees growing in good soil conditions and those from poorer soil conditions also occurred during this period. Levels of P in leaves were highest in young developing leaves but once the leaves reached full size, no seasonal trend in P concentration was observed. Foliar K was lower during the winter and probably related to the period of maximum leaching by precipitation. High foliar K during summer, however, could be related to the role of K in lowering cellular water potential. Leaf Ca was highest during early sping. Low mobility of cellular Ca during the cool portion of the year was indicated. Foliar Mg showed a weak pattern of decreasing concentration with leaf age. The best season for sampling for these broadleafed evergreen species to provide information on plant nutrient status appears to be in spring.     

国产省藤属植物的叶表皮形态学

对国产省藤属（Calamus L．）植物17种2变种的叶表皮进行光学显微镜观察,除单叶省藤（C.simplicifolius）、短叶省藤（C.egregiw）和白藤（C.tetradactylus）外,其它种均为首次报道。省藤属植物的叶表皮形态较为一致,气孔器为轮列4细胞型（tetracytic,或称四轮列型）,即有4个副卫细胞,其中2个为侧副卫细胞,2个为端副卫细胞;表皮细胞为长方形,纵向排列。表皮毛有3种类型：多细胞排成单列的单毛（人喙省藤C．macrorrhynchus）、多细胞排成2-3列的单毛（毛鳞省藤ethysanolepis等）、多细胞分枝毛（白藤等）。表皮细胞的长度和宽度特征、细胞壁的厚薄及波纹的深浅和形状、端副卫细胞的长短等细微特征在种问存在差异,可以区分种类。叶表皮结构还反映了种问的亲缘关系。因此,叶表皮特征对于省藤属种类的划分和亲缘关系的确定有较大的分类学意义。     

Oldest known Eucalyptus macrofossils are from South America

The evolutionary history of Eucalyptus and the eucalypts, the larger clade of seven genera including Eucalyptus that today have a natural distribution almost exclusively in Australasia, is poorly documented from the fossil record. Little physical evidence exists bearing on the ancient geographical distributions or morphologies of plants within the clade. Herein, we introduce fossil material of Eucalyptus from the early Eocene (ca. 51.9 Ma) Laguna del Hunco paleoflora of Chubut Province, Argentina; specimens include multiple leaves, infructescences, and dispersed capsules, several flower buds, and a single flower. Morphological similarities that relate the fossils to extant eucalypts include leaf shape, venation, and epidermal oil glands; infructescence structure; valvate capsulate fruits; and operculate flower buds. The presence of a staminophore scar on the fruits links them to Eucalyptus, and the presence of a transverse scar on the flower buds indicates a relationship to Eucalyptus subgenus Symphyomyrtus. Phylogenetic analyses of morphological data alone and combined with aligned sequence data from a prior study including 16 extant eucalypts, one outgroup, and a terminal representing the fossils indicate that the fossils are nested within Eucalyptus. These are the only illustrated Eucalyptus fossils that are definitively Eocene in age, and the only conclusively identified extant or fossil eucalypts naturally occurring outside of Australasia and adjacent Mindanao. Thus, these fossils indicate that the evolution of the eucalypt group is not constrained to a single region. Moreover, they strengthen the taxonomic connections between the Laguna del Hunco paleoflora and extant subtropical and tropical Australasia, one of the three major ecologic-geographic elements of the Laguna del Hunco paleoflora. The age and affinities of the fossils also indicate that Eucalyptus subgenus Symphyomyrtus is older than previously supposed. Paleoecological data indicate that the Patagonian Eucalyptus dominated volcanically disturbed areas adjacent to standing rainforest surrounding an Eocene caldera lake.     

A new species of Nelumbo from South China and its palaeoecological implications

A new species of Nelumbo Adanson is described from the Eocene of the Changchang Basin, Hainan Island, South China. This is the first comprehensive report on a fossil Nelumbo species from the Eocene of the Changchang Basin, Hainan Island. The co-occurrence of fruits, tubers, rhizomes with numerous adventitious roots, leaves, and receptacles indicates that the plants grew where they are preserved. These specimens, combined with palynological assemblage and other macrofossils, indicate a tropical or subtropical humid climate in the Changchang Basin from the late early Eocene to early late Eocene.     

Comparative anatomy of conifer leaves

The epidermis and internal structure of 103 species and 8 varieties of conifer leaves belonging to 37 genera of 7 families have been comparatively investigated. The anatomical characters of leaves of all the genera (or sections) studied are described systematically and a key to the genera (or sections) is also presented. It is shown that in order to identify at the genus level, the leaf types of LaubenFels’ may be used as the primary characterization. And it can also be seen that the following characters which are relatively constant at the genus level provide an important basis for identification: the epidermal structure, the presence or absence of hypodermis, the degree of differentiation of mesophyll, the occurrence of vascular bundle sheath or endodermis, the transfusion tissue type, the number and relative spatial position of vascular bundles, the number and distribution of resin canal and the presence of sclereid or sclerenchyma. The genera of Cupressaceae (except for Juniperus and Sabina) are of scale leaves (Leaf type-III) with little differences in structural characters. It’s difficult to distinguish them from each other. In addition, some problems concerning the significance of the resin canal, the endodermis with Casparian-strip, the sclerenchyma and the vascular bundle in identificationare here also discussed.     

不同地区青檀叶片的解剖及表皮特征的比较研究

采用石蜡切片技术和叶表皮离析法,对我国8个省共15个居群的青檀(Pteroceltis tatarinowii Maxim.)叶片特征进行了比较和分析.研究发现:各居群的栅栏组织细胞层数多数为2层,显示出一定的稳定性,但不同居群的叶片横切面解剖特征存在显著性差异.叶片上表皮细胞有多边形和不规则形两种类型,而下表皮细胞均为不规则形.上表皮细胞垂周壁为平直形、圆形、浅波形或深波形,差异较明显,下表皮细胞垂周壁则仅有深波形和浅波形两种.叶表皮毛的分布位置在各居群间不同,而表皮毛的类型在居群间稳定,且在多数居群中表皮毛与周围细胞形成花环结构.各居群的气孔器类型均为不规则型且均仅在下表皮随机分布,但气孔器的密度、大小等特征在不同居群间存在显著性差异.生境地的气候因子对青檀叶的特征产生一定的影响.结果表明:青檀叶的解剖特征在不同居群间差异性较大,青檀叶表皮的气孔密度等特征在居群间有一定差异,而气孔器类型和表皮细胞形状等特征较稳定,有一定的分类学价值.     

野丁香属植物叶表皮形态特征研究

在光学显微镜和电子扫描显微镜下观察了茜草科(Rubiaceae)野丁香属(Leptodermis Wall.) 35种8变种植物的叶表皮微形态特征.结果表明,野丁香属植物叶表皮细胞为多边形或不规则形状,垂周壁平直、弓状或波状.气孔器全部位于下表皮,有辐射型、平列型两种,其中平列型较为普遍.表皮毛在上下表皮都有分布,有单...     

Eocene plant diversity at Laguna del Hunco and Río Pichileufú, Patagonia, Argentina

The origins of South America's exceptional plant diversity are poorly known from the fossil record. We report on unbiased quantitative collections of fossil floras from Laguna del Hunco (LH) and Río Pichileufú (RP) in Patagonia, Argentina. These sites represent a frost-free humid biome in South American middle latitudes of the globally warm Eocene. At LH, from 4,303 identified specimens, we recognize 186 species of plant organs and 152 species of leaves. Adjusted for sample size, the LH flora is more diverse than comparable Eocene floras known from other continents. The RP flora shares several taxa with LH and appears to be as rich, although sampling is preliminary. The two floras were previously considered coeval. However, (40)Ar/(39)Ar dating of three ash-fall tuff beds in close stratigraphic association with the RP flora indicates an age of 47.46+/-0.05 Ma, 4.5 million years younger than LH, for which one tuff is reanalyzed here as 51.91+/-0.22 Ma. Thus, diverse floral associations in Patagonia evolved by the Eocene, possibly in response to global warming, and were persistent and areally extensive. This suggests extraordinary richness at low latitudes via the latitudinal diversity gradient, corroborated by published palynological data from the Eocene of Colombia.