Approaches to the identification of angiosperm leaf remains

During the past 125 years the history of early angiosperms, interpreted through the fossil leaf record has been largely an exercise in paleofloristic studies, ignoring evolution. Imprecise identifications of ancient leaves “matched” to extant genera and families have been used as the basis for reconstructions of paleocommunities and paleoclimates. However, as the result of careful morphological studies of leaf form, venation and cuticular features new insights into the evolution of angiosperms are now available. In this paper considerations are given to the usefulness and shortcomings of leaf form, venation and cuticular analysis as diagnostic tools of plant identification. Many techniques for the study of the morphology of modern and fossil leaves are included in this paper as well as tables outlining features of leaf venation and the epidermis. Careful morphological studies of leaf form (such as the venation and epidermal characters emphasized in this paper) will provide better understanding of the relationships of living angiosperms and transform the fossil leaf record into useful data that can be used to study the evolution of the angiosperms.     

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.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

叶脉网络功能性状及其生态学意义

叶脉网络结构是叶脉系统在叶片里的分布和排列样式。早期叶脉网络结构研究主要集中在其分类学意义上; 近年来叶脉网络功能性状及其在植物水分利用上的意义已成为植物生态学研究的热点。该文介绍了叶脉网络功能性状的指标体系(包括叶脉密度、叶脉直径、叶脉之间的距离、叶脉闭合度等), 综述了叶脉网络功能性状与叶脉系统功能(包括水分、养分和光合产物等物质运输、机械支撑和虫害防御等)的关系, 叶脉网络功能性状与叶片其他功能性状(包括比叶重、叶寿命、光合速率、叶片大小、气孔密度等)的协同变异和权衡关系, 以及叶脉网络功能性状随环境因子(包括水分、温度、光照等)的变化规律等方面的最新研究进展。此外, 叶脉网络功能性状的研究成果也被应用于古环境重建、城市交通规划、流域规划及全球变化研究中。由于叶脉网络功能性状是环境因子与系统发育共同作用的结果, 未来开展分子—叶片—植物—生态系统等多尺度的叶脉网络功能性状研究, 理清叶脉网络功能性状与气孔失水—茎干导水—根系吸水等植物水分利用的关系, 将为预测植物及生态系统对全球变化的响应提供新的启示。     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

DEVELOPMENTAL ANALYSIS OF LEAF PLASTICITY IN THE HETEROPHYLLOUS AQUATIC PLANT HIPPURIS VULGARIS

Leaf development was studied in the heterophyllous aquatic plant Hippuris vulgaris in order to characterize the developmental events that lead to the formation of aerial- vs. submerged-type leaves. Recent evidence that abscisic acid regulates leaf development in this species provided a basis for using abscisic acid as a developmental tool to accurately control leaf development. We found that leaf primordia were fully competent to develop into either aerial- or submerged-type leaves until the 10th plastochron, when they were ca. 300 μm long. Also, leaves between about the 10th and 21st plastochron formed sectored transition leaves (i.e., the basipetal portion was composed of aerial-type tissue and the apical portion was composed of submerged-type tissue, or vice versa), indicating that tissue determination as one or the other leaf type occurred on a local, as opposed to whole-leaf, level. Finally, no significant difference was observed between the apical dimensions of aerial or submerged-type shoots. These results indicate that the final determination of Hippuris vulgaris leaves occurs a) relatively late in leaf development, and b) independently of the shoot apex, and provide a basis for using this plant in further studies concerning leaf determination and pattern formation (e.g., stomates, lateral venation) in plants.     

主成分分析法在壳斗科植物化石鉴定中的应用*

壳斗科化石是北半球新生代地层的优势分子,然而由于其属种众多,叶片属种间特异性不明显,该科叶片化石的鉴定是古植物学研究的难点之一。面对化石标本不完整的性状特征信息,如何剔除冗余特征,重点考虑具有鉴定意义的标本特征,就显得非常必要。本研究通过调研多种降维算法,考虑到植物特征编码及赋值的数值特性,选用主成分分析法,以壳斗科青冈亚属植物叶片为例,对可能在化石叶片上观察到的22个性状特征(变量)进行降维处理,挑选出对于壳斗科化石叶片分类鉴定起主要作用的10个性状特征,并将其应用于化石鉴定进行验证。结果表明,仅考虑经过主成分分析法压缩挑选的10个主要性状特征,仍然能够实现壳斗科化石植物的准确鉴定。主成分分析法应用于壳斗科性状特征的降维处理效果良好,剔除冗余特征对标本鉴定结果没有影响。     

Leaf Architecture and Systematics of the Hamamelidaceae Sensu Lato

Hamamelids have a long fossil history and an important fossil record. Their interesting biogeographic relationships indicate a great age. There exist good surveys of the pollen and floral organs of this family whereas it is so far poorly known from leaf architecture. The leaf architecture of all 29 genera with more than 60 among the total of 140 species of the family was surveyed in this work using clearified leaves. It is found that leaf architecture analysis may shed light on the relationships within the family and the conclusion of evolution based on leaf architecture basically accords with that based on others. The major categories of leaf architecture of Hamamelids observed in this work are as follows: leaf form, leaf margin, tooth type, venation, marginal ultimate venation, areolation and trichome. It must be emphasized that of all these characters the tooth type is the most stable and useful for systematics. In this work a new tooth type is recognized under the name altingioid. Teeth of this type are obviously asymmetrical, with a persistent transparent gland on the top, and with their lateral veinlets free, not reaching the medial vein. All three genera of the subfamily Liquidambaroideae have this tooth type, whereas most leaves of the rest genera of this family have fothergilloid teeth, which are basically symmetrical, without glands. The venation in the fothergilloid tooth is almost the same as that in the altingioid tooth, the only difference being that the lateral veins on the abaxial side of the altingioid teeth are usually absent or very weak and short if present. The present authors consider that the subfamily Liquidambaroideae has to be separated from the family Hamamelidaceae sensu lato and treated as an independent family, Altingiaceae, on the basis of the special tooth type. different pollen morphology and flower structure. The stability of tooth type may serve classification not only of order and family level, but also of tribe, genus and species level with the help of characters of teeth, such as shape, size, density, distribution, single or double, with or without glands. By comparison of Hamamelidaceae and Altingiaceae with some primitive families of subclass Hamamelidae, namely, Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Eupteleaceae and Platanaceae, the putative evolutionary trend of tooth types is outlined as follows: ↑ altingioid Chloranthoid → Cercidiphylloid →platanoid → fothergilloid In general evolutionarytrend of teeth within these families is reduction and simplification in structure.     

USE OF GROWTH CHARACTERISTICS IN STUDIES OF MORPHOLOGIC RELATIONS. I. SIMILARITIES BETWEEN EPIBLAST AND COLEORHIZA

Foard , Donald E., and Alan H. Haber . (Oak Ridge Natl. Lab., Oak Ridge, Tenn.) Use of growth characteristics in studies of morphologic relations. I. Similarities between epiblast and coleorhiza. Amer. Jour. Bot. 49(5): 520–523. Illus. 1962.—We compared the characteristics of wheat epiblast, coleorhiza, and leaves during germination and seedling growth. Although the epiblast and coleorhiza grow without cell division during germination, they sometimes produce hairs that resemble root hairs but are different in form and mode of origin from leaf hairs. Despite the similar orientation of the epiblast and leaves with respect to the main axis of the plant, during growth the epiblast changes shape such that the axis of greater linear extension per millimeter is perpendicular to the main axis of the plant, whereas leaves change shape such that the greater rate of linear extension per millimeter is parallel to the main axis of the plant. Indoleacetic acid treatment produces excessive and disorganized growth of both the epiblast and coleorhiza but retards leaf growth. Gibberellic acid stimulates, whereas (2-chloroethyl)trimethylammonium chloride retards leaf growth. Neither substance significantly affects epiblast or coleorhiza. We conclude that the epiblast is not leaf-like, but together with the coleorhiza forms a single, continuous structure. This paper illustrates how a wide variety of growth characteristics may be used to complement classical anatomic approaches in studies of morphologic relations.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

Scutifolium jordanicum gen. et sp. nov. (Cabombaceae), an aquatic fossil plant from the Lower Cretaceous of Jordan, and the relationships of related leaf fossils to living genera

A new species of aquatic plant, Scutifolium jordanicum gen. et sp. nov., Taylor, Brenner & Basha, is described from the Albian of Jordan. The leaves are microphyllous with a symmetrical, elliptical to suborbiculate shape, convex to rounded apex and base, and actinodromous to palinactinodromous primary venation. The peltate, centrally attached petioles are narrow, elongate, and alternately arranged on similarly sized stems. The leaves appear to be thick and have aerenchyma. Comparisons to plants with centrally peltate leaves and palmate venation and to aquatic plants with floating leaves suggest that S. jordanicum belongs to the Cabombaceae lineage within the Nymphaeales. Cladistic analysis including the fossil and living members of the Nymphaeales shows that the S. jordanicum is basal to the living members of the family and has unique characters not found in any living genera. This is the oldest evidence of the Cabombaceae from the Old World. Inclusion of two other Early Cretaceous peltate leaf fossils in the phylogenetic analysis indicates their affinities to Cabombaceae and that some of the shared derived characters for the living members are progressively acquired in the fossils. These data show the Cabombaceae were widespread in Gondwana and Laurasia by the mid-Cretaceous.     

CHILBINIA GEN. NOV., AN ARCHAIC SEED FERN IN THE LATE TRIASSIC CHINLE FORMATION OF ARIZONA, USA

Abstract:  A seed-bearing pinnate leaf found in the Chinle Formation of Late Triassic age in Arizona appears to be an archaic seed fern that more closely resembles typical Carboniferous–Permian seed fern leaves than any younger form. The fossil, which is described here as Chilbinia lichii gen. et sp. nov., incorporates characters of several Palaeozoic taxa. These characters include pinnules that have reticulate venation resembling that in some species of Lonchopteris , cordate bases similar to those found in many species of Sphenopteris , and, except for a significant difference in size, linear ovate seeds that resemble the radially symmetrical seeds found attached to a species of Alethopteris in the Permian of China. The presence of this fossil in strata of Late Triassic age is further evidence that plant extinctions after the Permian–Triassic ecological crisis followed a stepwise pattern and apparently continued for at least 20–25 myr after the end of the Permian in western parts of the Triassic paratropical belt.     

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。     

THE SYSTEMATIC POSITION OF EOGINKGOITES

Eoginkgoites is a fan-shaped, imparipinnate leaf with a short rachis and long petiole. It was first described from the Upper Triassic Newark Group of Pennsylvania by Bock who assigned the fossil to the Ginkgoales. The fossil has also been found in the Upper Triassic Chinle Formation in Utah and Arizona and in the Newark Group in North Carolina. Investigation of the well-preserved specimens found in the Chinle Formation shows that the leaf has anastomosing venation, a marginal vein and paracytic (syndetocheilic) stomata. These characters indicate that the leaf is bennettitalean and Eoginkgoites is reassigned to the Bennettitales although its shape is perplexing. Eoginkgoites may be an important index fossil to the lower Upper Triassic (middle Carnian) rocks of North America.     

Tertiary leaf fossils of Mangifera (Anacardiaceae) from Li Basin, Thailand as examples of the utility of leaf marginal venation characters

The mango (Mangifera) is the most important commercial fruit in Asia and is popular worldwide. Because of uncertainty in its origin and biogeography, the study of Mangifera could be of value in elucidating its current genetic diversity and geographical distribution. We report here two new species of compressed leaves of Mangifera (Anacardiaceae) from upper Oligocene-lower Miocene lacustrine sediments in the Li Basin, northern Thailand. Leaf form and venation characters of fossil and extant leaves are presented with detailed marginal venation patterns that were found to be important in assigning the fossil leaves to Mangifera paleoindica and M. paleocaloneura. These two fossil species of Mangifera and the present occurrence of basal species of Mangifera in Thailand suggest that northern Thailand could be an area with a long history of evolution and diversification of Mangifera, particularly M. indica.     

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     

MORPHOLOGY AND DICHOTOMOUS VASCULATURE OF THE LEAF OF KINGDONIA UNIFLORA

Foster , Adriance S. (U. California, Berkeley), and Howard J. Arnott . Morphology and dichotomous vasculature of the leaf of Kingdonia uniflora. Amer. Jour. Bot. 47 (8): 684–698. Illus. 1960.—An intensive study of the nodal anatomy, petiolar vasculature and open dichotomous venation of the leaf of Kingdonia has revealed a type of foliar vascular system of unusual morphological and phylogenetic interest. The vascular supply at the nodal level consists of 4 collateral traces which diverge from a single gap into the sheathing leaf base. This type of nodal anatomy is perhaps primitive, and comparisons are made with the unilacunar nodes and the 2- and 4-parted leaf trace systems characteristic of many angiospermous cotyledons and the foliage leaves of certain woody ranalian genera. The petiole of Kingdonia is vascularized by 2 pairs of bundles which represent the upward continuation of the 4 leaf traces. A transition from an even (4) to an odd (3) number of strands occurs near the point of attachment of the 5, lobed, cuneiform lamina segments to the petiole. Each of the 2 abaxial bundles dichotomizes and the central derivative branches fuse to form a double bundle which enters the base of the median lamina segment. The 2 adaxial petiolar bundles diverge right and left into the bases of the paired lateral segments of the lamina. An analogous type of transition from an even to an odd number of veins occurs in many angiospermous cotyledons which develop a definable mid-vein. But, in Kingdonia, the bundles which enter the bases of the lamina segments give rise to systems of dichotomizing veinlets devoid of “mid-veins.” Although the majority of the terminal veinlets enter the marginal teeth of the lamina segments, “blind” endings, unrelated to the dentations, occur in all the leaves studied. Typically, all of the vein endings in a given lobule of a lamina segment are derived from the same dichotomous vein system. However, in some leaves, a veinlet dichotomizes directly below a sinus and the branches diverge into the marginal regions of 2 separate lobules. The phylogenetic significance of the occurrence of open dichotomous venation in such an herbaceous angiosperm as Kingdonia is briefly discussed. From a purely morphological viewpoint, the Kingdonia type of venation invites direct comparison with the venation of Sphenophyllum, certain ferns or Ginkgo rather than with any of the known reticulate venation patterns of modern angiosperms. Although the foliar venation of Kingdonia may represent the result of evolutionary reversion, the very rare anastomoses which occur seem primitive in type rather than “vestiges” of a former system of closed venation.     

A RECORD OF PAULOWNIA IN THE TERTIARY OF NORTH AMERICA

Smiley , Charles J. (Macalester Coll., St. Paul, Minn.) A record of Paulownia in the Tertiary of North America. Amer. Jour. Bot. 48(2): 175–179. Illus. 1961.—Paulownia, an eastern Asiatic genus in the family Scrophulariaceae, has been identified from fossil leaves in the Ellensburg flora of Washington. Age of these fossils is late Miocene (Barstovian) to early Pliocene (Clarendonian). They are nearly identical to leaves of the living P. tomentosa (Thunberg) Steudel, a deciduous tree now growing in temperate parts of China and one that has escaped from cultivation in eastern United States. Identification is based on comparative leaf morphology. Modern and fossil Paulownia leaves may be distinguished from those of such other genera as Buettneria, Catalpa, and Cercis by secondary and tertiary venation. Central Washington environment during this time was well suited to the growth and reproduction of Paulownia, judging from such fossil associates as Liquidambar, Nyssa, Passiflora, Persea, Rhododendron, Ulmus, and Zelkova. Deciduous habit, present temperate environment, possible early Tertiary record in Alaska, later Tertiary records in middle latitudes of North America and Europe, and present occurrence in China indicate that Paulownia was once widely distributed over the northern hemisphere. Secular climatic trends toward cooler conditions since the Eocene appear to have resulted in the southward migration of Paulownia as a member of the Arcto-Tertiary Geoflora. Subsequent extinctions in Europe and North America may have resulted from Pleistocene glacial climates, and from barriers to further migrations to more southern latitudes.