A new species of Castanopsis (Fagaceae) from the upper Pliocene of West Yunnan,China and its biogeographical implications

Two fossil leaves identified as Castanopsis presclerophylla n. sp. collected from a diatomite bed in the upper Pliocene (3.3–2.8 Ma) in Tengchong County, Yunnan Province, Southwest China, are characterized by elliptic laminar shapes with acuminate apex, convex base, simple trichome bases and cyclocytic stomata. The cuticles of extant leaves from six selected Castanopsis species that are similar to the Pliocene fossil leaves in laminar shape were examined in detail for the comparison. Based on the analysis of leaf architecture and cuticular characteristics, the present fossil leaves most closely resemble the extant leaves of Castanopsis sclerophylla (Lindl.) Schott., which prefers a subtropical climate. The published fossil records of Castanopsis also indicate that it was once widely distributed in North America, Europe and Asia before the Pliocene, and then rapidly retreated to the East and Southeast Asia as a result of the Pleistocene glaciation.     

A new species of Carpinus (Betulaceae) from the Pliocene of Yunnan Province, China

Macrofossils of Carpinus have been widely reported from the Cenozoic of the Northern Hemisphere. However, the leaf cuticules of the genus have rarely been described. A new species, named Carpinus tengchongensis Dai et B.N. Sun, sp. nov., is identified based on 13 leaf fossils, collected from the late Pliocene Mangbang Formation, Tengchong County of Yunnan Province, China. The important characters of the fossil are its oblong-ovate leaf shape, obliquely cordate base, doubly serrulate margin, straight and moderately thick primary vein, pinnate secondary veins, percurrent tertiary veins, orthogonally reticulate areoles, absence of veinlets, anomocytic stomata with double-layered stomatal rim, well-formed T-pieces and trichome bases, which indicates an affinity within the genus Carpinus section Carpinus subsection Monbeigianae, especially with C. tsaiana. Carpinus fossils were present from the Eocene to Pliocene with disjunctive distribution in the North Temperate Zone, which broadly reflected the present distribution pattern and probably demonstrates the tolerance of Carpinus to environmental change.     

A new species of Eretmophyllum (Ginkgoales) from the Middle Jurassic of Turpan-Hami Basin,Xinjiang, China

A few specimens previously assigned to Eretmophyllum, based on the gross leaf morphology, lack cuticular structures. In this paper, we describe a new species of Eretmophyllum from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China, based on fossil leaves preserved as compressions, and characterized by paddle shapes with conspicuous petioles, retuse apices, entire margins and two veins dichotomously branched from near the base. Leaf amphistomatic, the cuticles are composed of stomatal and non-stomatal bands. Based on a detailed comparison of the gross morphology and cuticular structures with previously reported fossils, Eretmophyllum hamiensis n. sp. is erected. The presence of Eretmophyllum indicates that Middle Jurassic climate in the Turpan-Hami Basin was likely warm and humid. On the basis of the geological history and geographical distributions of Eretmophyllum, we conclude that the genus probably originated in Europe and migrated from Siberia to northwestern China during the Middle Jurassic.     

湖南安息香属植物的叶片比较解剖学研究

利用光学显微镜和扫描电镜对湖南产安息香属10种植物叶表皮微形态和解剖结构进行了观察和测量。结果表明:(1)表皮均由单层细胞构成,细胞形状为多边形和无规则形,垂周壁式样为平直、弓形、波状;(2)气孔器大小及密度在种间的差异明显;(3)气孔外拱盖内缘主要为波状和浅波状,气孔器外围角质纹饰以越南安息香的鸟巢状和栓叶安息香的碗状为显著区别特征;(4)老鸹铃、垂珠花和白花龙的气孔器保卫细胞两极有"T"形加厚;(5)栓叶安息香和大果安息香叶上表皮由大型细胞组成;(6)叶肉分化为栅栏组织和海绵组织,二者的厚度及比值在种间有明显的差异,其中越南安息香栅栏组织分化为2层;(7)大果安息香和芬芳安息香的主脉维管组织构成一圈封闭的环状结构,而其他种则为开放的半圆形结构;(8)叶片及主脉的厚度在种间差异明显。安息香属植物的表皮微形态及解剖结构特征比较稳定,可作为种间鉴定和分类的重要依据。     

云南保山上新统黄背栎Quercus Pannosa Hand.-Mazz.角质层特征及古环境意义

在滇西保山上新统首次发现了黄背栎(Quercuspannosa)压型化石,并分析了角质层构造。其特征主要为:上表皮无气孔器,表皮细胞为不规则形五边形,垂周壁微弯曲,脉络为网格状,无毛基;下表皮具环列型气孔器,表皮细胞为不规则形五边形,垂周壁微弯曲,脉络为五边形的网格状,具毛基,毛的根部由多细胞组成,毛基部的细胞特化成辐射状。利用现存最近亲缘种(NLRspecies)原理,并比较化石和现生Q.pannosa角质层特征,推断上新世Q.pannosa在羊邑生长的古环境比现生Q.pannosa生存的冷湿气候相对干旱,且化石Q.pannosa为近原地埋藏。     

云南腾冲新近纪两种被子植物化石的角质层构造及其古环境意义

重点描述云南腾冲晚第三纪两种被子植物化石Betula mioluminifera Hu et Chaney,Carpinus subcordata Nathorst的角质层构造,并分析它们的现存最近亲缘种B.luminifera Winkler和C.cordata B1.var.mollis Cheng et Chen的表皮特征。实验分析证明：化石叶片的气孔参数可以推测地质历史时期大气CO2的浓度,并进而分析古环境的变化。C．subcordata Nathorst叶片能作为大气CO2浓度的生物指标。     

Genus Triticum L. taxonomy: the present and the future

The genetic relationships and diversity within the European and Asiatic Buxus species were analysed using AFLP, genome size analysis and chromosome counts. Based on these results two major clusters could be defined. One genetic cluster contained B. sempervirens and B. balearica, European species, and B. colchica, an Asiatic species but with leaf morphology similar to B. sempervirens. Species in this cluster were characterised by a genome size between 1.38 and 1.69?pg?2C^?1 and a chromosome number of 2n?=?2x?=?28 (diploid). Only four B. sempervirens cultivars within this cluster were triploid. A second cluster contained the Asiatic Buxus species B. microphylla, B. harlandii, B. hyrcana, B. myrica, B. henryi, B. bodinieri and B. wallichiana. Within this second genetic cluster three different ploidy levels could be observed. B. harlandii, B. hyrcana and nine B. microphylla cultivars were tetraploid (2n?=?4x?=?56) with a genome size of >2.5?pg?2C^?1. Fifteen other B. microphylla cultivars were triploid (2n?=?3x?=?42). The other Asiatic Buxus species, B. henryi, B. bodinieri and eight B. microphylla cultivars, were diploid with a genome size of ca. 1.5?pg?2C^?1.     

Pliocene Albizia (Fabaceae) from Jharkhand,eastern India: reappraisal of its biogeography during the Cenozoic in Southeast Asia

Albizia, a diverse tree genus, occupies monsoonal warm humid rain forests in tropical and subtropical regions. We recovered a well-preserved compound fossil leaf and two fossil fruits of Albizia (Fabaceae) from the latest Neogene (Rajdanda Formation: Pliocene) sediments of Jharkhand of the Chotanagpur Plateau, eastern India. On the basis of the architectural features of the fossil leaf, a new species is established as A. mahuadanrensis Hazra, Hazra and Khan, n. sp., characterised by a bipinnate, compound leaf having a rachis bearing opposite, asymmetrically ovate to sub-rhomboid leaflets, pulvinus on leaflet petiolule and brochiodromous secondary veins. Based on both morphological and anatomical characters of the fossil fruits, A. palaeoprocera Hazra, Hazra and Khan, n. sp. is erected, characterised by flattened to broadly linear shaped, wingless fruits; ovate-elliptic shaped seed chambers having ellipsoidal seeds in one series; irregularly polygonal to rectangular epidermal cells with oblique end walls and randomly oriented, scattered, paracytic stomata. Analysis of Albizia fossil occurrences indicates that the legume taxon was common in Neogene forests of India and elsewhere. The present-day distribution of the closely affiliated modern species of the fossil taxa indicates a warm and humid tropical environment during the time of deposition. We also review the biogeographic history of Albizia in India and other Asian countries.     

The pattern of cuticular blue-fluorescing phenolics deposition during the development of Prunus persica leaves

Although stomatal ontogeny is closely related to the development and maturation of the epidermal tissue, stomatal patterns in relation to cuticle construction and cuticular material deposition during leaf development have not received adequate attention. We observed the deposition of blue-fluorescing cuticular phenolics over guard and epidermal cells, as well as stomatal formation and patterning using the alkali-induced blue fluorescence of the cuticle of Prunus persica leaves. Stomata of different stages of maturity occurred together during leaf development, mainly at the tip of the lamina. The deposition of fluorescing compounds initially appeared over the guard cells of the developing stomata complexes and gradually extended to the neighbouring epidermal cells. Based on the blue fluorescence emitted by the cuticular layers, we constructed digital maps of leaves of different developmental stages, showing the pattern of stomatal formation and deposition of fluorescing compounds. A longitudinal tip-to-base gradient in the formation of stomata, as well as in the deposition of fluorescing compounds was observed in young developing leaves. The deposition of blue-fluorescing phenolic compounds seems to be coordinated with stomatal development.     

Stomatal and cuticular transpiration of beans (Phaseolus vulgaris M.) attacked by rust (Uromyces appendiculatus [Pers.] Link)

The changes of stomatal and cuticular transpiration of bean plants were investigated by graphical transpiration curves method (Slavík 1958). Bean leaves were infected by fungusUromyces appendiculatus (Pers.) Link. After the infection the intensity of stomatal transpiration had a decreasing tendency. Beginning with the sixth day after infection, the proportion of stomatal and cuticular transpirations becomes more expressive, i.e. the leaves transpire more by cuticles than by stomata. The higher share of cuticular transpiration brings extensive water relations to the diseased plants.     

Transpiration and stomatal control: a cross-species study of leaf traits in 39 evergreen and deciduous broadleaved subtropical tree species

Key message

Using an extensive dataset for 39 subtropical broadleaved tree species, we found traits of the leaf economics spectrum to be linked to mean stomatal conductance but not to stomatal regulation.

Abstract

The aim of our study was to establish links between stomatal control and functional leaf traits. We hypothesized that mean and maximum stomatal conductance (g _s) varies with the traits described by the leaf economics spectrum, such as specific leaf area and leaf dry matter content, and that high g _s values correspond to species with tender leaves and high photosynthetic capacity. In addition, we hypothesized that species with leaves of low stomata density have more limited stomatal closure than those with high stomata density. In order to account for confounding site condition effects, we made use of a common garden situation in which 39 deciduous and evergreen species of the same age were grown in a biodiversity ecosystem functioning experiment in Jiangxi (China). Daily courses of g _s were measured with porometry, and the species-specific g _s~vpd relationships were modeled. Our results show that mean stomatal conductance can be predicted from leaf traits that represent the leaf economics spectrum, with a positive relationship being related to leaf nitrogen content and a negative relationship with the leaf carbon: nitrogen ratio. In contrast, parameters of stomatal control were related to traits unassociated with the leaf economics spectrum. The maximum of the conductance~vpd curve was positively related to leaf carbon content and vein length. The vpd at the point of inflection of the conductance~vpd curve was lower for species with higher stomata density and higher for species with a high leaf carbon content. Overall, stomata size and density as well as vein length were more effective at explaining stomatal regulation than traits used in the leaf economics spectrum.     

中国桑寄生科植物叶表皮微形态

通过扫描电镜对中国桑寄生科桑寄生亚科8属18种和槲寄生亚科1属2种植物成熟叶的上、下叶表皮内表面和下表皮外表面进行了研究。内面观发现桑寄生科植物叶上、下表皮形状为多边形,垂周壁式样平直或稍弓形,常具有角质增厚,平周壁常覆盖厚角质或颗粒状、丝状角质增厚;气孔存在于上下表皮,通常下表皮较多,气孔的形状,特别是保卫细胞的形态在亚科间、属间或种间都具有一定的差异,气孔器类型为平列型或单圈型。下表皮表面观察了的角质膜和蜡质纹饰、气孔的形状,外部气孔缘及外部气孔缘内缘的特征。这些特征在亚科或属级水平上较为稳定,有的也表现出种间差异,有一定的分类价值。从气孔形态和外部气孔周围角质膜来看,两亚科显示出明显的不同:桑寄生亚科上、下表皮均具有内部气孔缘,而槲寄生亚科没有此结构;桑寄生亚科外部气孔周围角质膜增厚成环状,其上具增厚的条纹,而槲寄生亚科外部气孔周围角质膜增厚成脊状,不具条纹。这些特征支持槲寄生亚科作为独立1个科来处理。     

In situ occurrence of a gall midge (Insecta,Diptera, Cecidomyiidae) on fossilized angiosperm leaf cuticle fragments from the Pliocene sediments of eastern India

In situ preservation of fossil insect damage in plant fossils is an excellent tool to study the coevolution of flora and fauna through geological time, but finding both damage and the insect causing that damage in the same specimen is a very rare phenomenon. Galling is a common form of angiosperm leaf damage, which can be regarded as a kind of extended phenotype of the causal insects, essentially the gall midges, but galls usually lack remains of the insects themselves. Here we report the in situ occurrence of a gall midge (Insecta, Diptera, Cecidomyiidae) as well as its pupal exuviae on the abaxial cuticular surface of fossilized leaf cuticle fragments of Fabaceae leaves (cf. Albizia) that also bear galls, recovered from the latest Neogene (Rajdanda Formation, Pliocene) sediments of the Chotonagpur Plateau, Jharkhand, northeastern India. This Pliocene gall midge features well-preserved legs, segmented antenna with distinct and enlarged scape, elongate curved setae, and longer than broad terminal plate of the ovipositor lamellae. The in situ presence of a gall midge on a host fabaceous leaf cuticle indicates the existence of a host-ectoparasite relationship in the ancient warm and humid tropical monsoon-influenced forests of eastern India during the Pliocene. This is the first authentic fossil record of an in situ phytophagous insect of Cecidomyiidae from India, as well as southeast Asia. Although the identification of the recovered phytophagous insect associated with the fossil leaf cuticle is only possible to family level, this find reveals that such plant-insect relationships existed in the Pliocene of eastern India.     

Taxonomic and ecological implications of leaf cuticular morphology in Castanopsis, Castanea, and Chrysolepis

Leaves of 41 species of Castanopsis, six species of Castanea, and Chrysolepis chrysophyllum Hjelmq. were examined. In Castanopsis, all species possessed cyclocytic stomata with thickened subsidiary cells; thin-walled peltate trichomes are the most frequent type on the abaxial surface of the leaves of this genus. In Castanea, stomata are transitional between cyclocytic and anomocytic; thin-walled peltate trichomes were recorded for the first time on the abaxial surface of Castanea leaves. In Chrysolepis, cyclocytic stomata with non-thickened subsidiary cells and thick-walled peltate trichomes were observed. The thickened subsidiary cells support the placement of the “fissa-group” in Castanopsis. The results of this study support the idea that Castanopsis and Castanea are sister groups. Thick-walled peltate trichomes were only recorded in Chrysolepis, thus supporting its taxonomic separation from Castanopsis. The phylogenetic distribution of trichome types among genera of Fagaceae is summarized. The evolutionary trends of trichome types in Castanopsis are discussed, as are the implications of stomatal and trichome features on fossil identification and ecology.     

A Method for Separating Cuticular and Stomatal Components of Gas Exchange by Amphistomatous Leaves: II. EXPERIMENTAL SOLUTION

The apparent cuticular component of transpiration of stomatabearing leaf epidermis was estimated by restricting stomataldiffusion by mass flow of air in the opposite direction. Thiswas achieved by applying an air pressure gradient across theamphistomatous leaf. Some assumptions of the previously suggestedmethod (antrcek and Slav?k, 1990) were experimentally verifiedusing maize leaves. The technique makes possible a quantitativeestimation of cuticular water loss including that of the externalperistomatal (i.e. vapour not passing through the pores) andthe respective conductance when the stomata are partially open. In addition to the fact that the cuticular portion of the totalleaf vapour loss (i.e. relative cuticular transpiration) dependson stomatal opening, even the absolute value of apparent cuticulartranspiration was (1) increased by lower vapour pressure deficitand (2) decreased with closing stomata. These changes, inducedby variations in a vapour pressure deficit of 2.45?0.35 kPa,ranged between 0.66?0.14µg cm ^–2 s^–1. Theabsolute value of apparent cuticular transpiration changed onaverage by a factor of 2.3 due to stomata opening or closingwhich was induced by turning the light on or by exogenous ABAapplication. Possible interference by residual vapour diffusingthrough the stomatal pore was evaluated by the model application.An attempt was also made to assess the cuticular component ofCO₂-uptake rate. Experimental results are discussed in contextwith the feedforward response of stomata to air humidity. Key words: Cuticular transpiration, cuticular CO₂-uptake, feedforward response, maize     

八角目(Illiciales)植物花被片表皮比较形态

在光学显微镜和扫描电镜下,观察了八角科(Illiciaceae)八角属(Illicium Linn.)2组11种20个材料、五味子科(Schisandraceae)南五味子属(Kadsura Ksempf. ex Juss.)2亚属2组8种15个材料和五味子属(Schisandra Michx.)2亚属4组6种17个材料的花被片表皮形态特征。首次报道了八角目2个科（八角科和五味子科）3个属（八角属、南五味子属和五味子属）植物花被片表皮细胞的形状、分泌细胞的形状及分布、气孔器的形状及分布、花被片表面的纹饰。通过与八角目2个科3个属植物的叶表皮形态比较,发现花被片表皮气孔器外拱盖均为单层,与叶表皮气孔器外拱盖层数（常绿种类为双层和落叶种类为单层）之间没有相关性,还在五味子科中发现2个新的性状（气孔对和环列型气孔）。通过对两性花、雌花和雄花花被片表皮观察,发现花被片表皮形态与花的性别之间没有相关性。通过对八角属、南五味子属和五味子属花被片表皮比较,发现五味子属与南五味子属相比,其花被片表皮表现出更多的衍生性状;南五味子属与八角属相比,前者花被片表皮具有更多的衍生性状;而南五味子属花被片表皮形态与五味子属的相似性程度较大,支持五味子科包含南五味子属和五味子属、八角科包含八角属的观点。     

Environmental Pollution and Leaf Cuticular Variation in Kudzu (Pueraria lobata Willd.)

Four populations of kudzu (Puerarta lobata Willd.) were studiedin rural, relatively unpolluted areas and in habitats characterizedby heavy industrial pollution in north-west Tennessee, U.S.A.Leaf length, leaf width, petiole length, flower size and podsize showed a decrease in growth in heavily polluted areas.Trichome frequency and length on the leaf surfaces increasedwith an increase in environmental pollution while the stomatalfrequency values showed a slight decrease in polluted habitats.The length of the largest and the smallest stomata and the numberof undulations in the epidermal cells in kudzu plant populationswere not affected by environmental pollution. Subsidiary cellcomplex consisting of two cells also remained the same in allthe plant populations sampled in polluted and relatively unpollutedhabitats. Puerarialobata Willd., kudzu, environmental pollution, cuticular features, stomatal frequency, trichome frequency     

Spatial distribution characteristics of stomata at the areole level in Michelia cavaleriei var. platypetala (Magnoliaceae)

Background and AimsIn hierarchically reticulate venation patterns, smaller orders of veins form areoles in which stomata are located. This study aimed to quantify the spatial relationship among stomata at the areole level.MethodsFor each of 12 leaves of M. cavaleriei var. platypetala, we assumed that stomatal characteristics were symmetrical on either side of the midrib, and divided the leaf surface on one side of the midrib into six layers equidistantly spaced along the apical–basal axis. We then further divided each layer into three positions equidistantly spaced from midrib to leaf margin, resulting in a total of 18 sampling locations. In addition, for 60 leaves, we sampled three positions from midrib to margin within only the widest layer of the leaf. Stomatal density and mean nearest neighbour distance (MNND) were calculated for each section. A replicated spatial point pattern approach quantified stomatal spatial relationships at different distances (0–300 μm).Key ResultsA tendency towards regular arrangement (inhibition as opposed to attraction or clustering) was observed between stomatal centres at distances <100 μm. Leaf layer (leaf length dimension) had no significant effect on local stomatal density, MNND or the spatial distribution characteristics of stomatal centres. In addition, we did not find greater inhibition at the centre of areoles, and in positions farther from the midrib.ConclusionsSpatial inhibition might be caused by the one-cell-spacing rule, resulting in more regular arrangement of stomata, and it was found to exist at distances up to ~100 μm. This work implies that leaf hydraulic architecture, consisting of both vascular and mesophyll properties, is sufficient to prevent important spatial variability in water supply at the areole level.     

Leaf epidermal characteristics of Asiatic Buddleja L. under scanning electron microscope: Insights into chromosomal and taxonomic significance

The leaf epidermis of all 23 species from Asian Buddleja was investigated with scanning electron microscopy (SEM). Characteristics of trichome type, stomatal shape, cuticular membrane on adaxial and abaxial epidermis, inner margin of outer stomatal ledge and ornamentation of outer stomatal ledge are presented in this study. Among the Buddleja species, some distinct epidermal patterns were observed. These can be used to characterize and identify different sections among the Asian Buddleja species, viz. Alternifoliae, Curviflorae and Rectiflorae. In general, stomatal length of Buddleja increases with ploidy level. This character can be used to distinguish diploids from polyploids without preparation of caryograms. Based on this correlation Buddleja microstachya, B. subcapitata and B. bhutanica are characterized to be diploids, as concluded from their stomatal dimensions. Leaf epidermis characters can also be used to differentiate species pairs that are easily confused with each other, viz., B. asiatica and B. bhutanica, B. albiflora and B. nivea, B. candida and B. fallowiana, B. delavayi and B. forrestii, B. japonica and B. lindleyana, B. officinalis and B. paniculata.     

Morphological characters of leaf epidermis in schisandraceae and their systematic significance

The leaf epidermis of 23 species belonging to 2 genera within Schisandraceae was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandraceae, such as shape of epidermal cells, type of stomata, and cuticular ornamentation, are usually constant within species and thus helpful for elucidating the relationship between and within genera. Leaf epidermal cells are usually irregular or polygonal in shape. The patterns of anticlinal walls are straight, sinuolate, sinuous or sinuate. The stomatal apparatus belong to paracytic or laterocytic type and the latter is subdivided into various subtypes based on the number and arrangement of subsidiary cells. Under scanning electron microscopy observation, the cuticular membrane is often striated, sometimes squamulate or granular; the inner margin of the outer stomatal rim is nearly smooth or denticulate. Evidences from shape of epidermal cells, patterns of cuticular intrusions between the ends of each guard cell of a pair and distribution of stomatal apparatuses support the viewpoint thatKadsura is more primitive thanSchisandra. Study on leaf epidermis also shows thatKadsura interior deserves the rank of a distinct species and the treatment of the evergreen groups, includingS. propinqua andS. plena, as distinct from the deciduous species of the genus is quite natural.