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.     

ANASTOMOSES IN THE VENATION OF GINKGO BILOBA

Arnott , Howard J. (Northwestern U., Evanston, Ill.) Anastomoses in the venation of Ginkgo biloba. Amer. Jour. Bot. 46(6): 405–411. Illus. 1959.—Although the vasculature of the lamina of Ginkgo biloba has been described as open dichotomous and devoid of anastomoses, vein unions were found in a survey of 1065 leaves collected from both long and short shoots of 11 trees. When studied by directing a strong light through the lamina and by clearings, 9.9% of the leaves possessed 1 or more anastomoses. Long shoot-leaves showed 13.4% anastomoses while short shoot-leaves showed 8.2% anastomoses. Multiple anastomoses were found in almost half of the leaves bearing anastomoses. In the 105 leaves having vein unions, 163 anastomoses were counted. These anastomoses could be grouped into 4 types which are easily distinguished by the number of dichotomies involved and by the presence or absence of dichotomies above the point of vein union. Other deviations were found from the “normal” venation pattern; these consisted of unconnected veins, veins anastomosed marginally but unconnected basally, and veins ending a considerable distance from the margin. It was speculated that the anastomoses found in Ginkgo biloba are of a simple, archaic type and are apparently analogous to the anastomoses in the leaves of certain ferns and in the leaflets of various cycads. The evolutionary significance of these anastomoses must be assessed by a broad study of venation patterns in the seed-plants.     

ONTOGENY OF THE OPEN DICHOTOMOUS VENATION IN THE PINNA OF THE FERN NEPHROLEPIS

Pray , Thomas r . (U. South. California, Los Angeles.) Ontogeny of the open dichotomous venation in the pinna of the fern Nephrolepis. Amer. Jour. Bot. 47(5) : 319—328. Illus. 1960.–The venation of the pinna of Nephrolepis consists of a midvein and 2 lateral series of dichotomizing veins all of which terminate freely near the margins. The development of the pinna is analyzed with particular attention to the nature of the marginal meristem and the organization of the embryonic pinna as it appears in paradermal section. The arrangement of cells in pinna wings during the period of marginal growth displays a pattern which foreshadows the pattern of the mature venation. In contrast with the development of the leaves of angiosperms, marginal growth continues into a relatively late phase of pinna ontogeny and apparently is active throughout the phases of ontogeny concerned with blocking-out the pathways of procambial differentiation. Thus the pattern of venation appears to be correlated with the manner of activity of the marginal meristem and subsequent orientation of its derivatives. The theoretical aspects of the result of this investigation are discussed in relation to other studies of foliar venation ontogeny.     

星叶草叶脉序的形态学及其系统学意义

在星叶草叶二叉分枝脉序中,网结脉中2条完全汇合叶脉与靠近脉中完全分离叶脉间未发现任 何形式的维管束汇合的中间类型,相反观察到了网结脉中不同程度的连接脉退化的痕迹,因此网结脉不可能由靠近脉形成;盲脉的出现与齿的退化、网结脉中连接脉的间断及非网结脉由分枝处间断等相关;上述事实表明星叶草叶的二叉分枝脉序应为一减化性状。     

ONTOGENY OF THE CLOSED DICHOTOMOUS VENATION OF REGNELLIDIUM

Pray , Thomas R. (U. South. California, Los Angeles.) Ontogeny of the closed dichotomous venation of Regnellidium . Amer. Jour. Bot. 49(5): 464–472. Illus. 1962.—The venation of the pinna of Regnellidium consists of a flabellate series of dichotomizing vascular strands, branches of a single pinna trace. At the laminal margin, the entire venation is closed by a marginal vein. The development of the pinna was investigated primarily by means of paradermal sections. During early development, the organization of the marginal meristem and its derivatives is very similar to that previously described for Nephrolepis. The arrangement of cells in the embryonic pinna is predisposed for the differentiation of dichotomizing procambial strands. During the final phases of pinna development, the marginal meristem is altered in such a manner as to result in a submarginal band of elongated cells which differentiates a marginal procambial strand connecting the tips of the dichotomizing veins. Relatively late in pinna ontogeny and after the entire procambial venation pattern has been delimited, the marginal meristem becomes inactive. The possible correlation between extended marginal growth and dichotomizing veins is discussed.     

Open dichotomous venation in the leaves ofUtricularia striatula (Lentibulariaceae)

The open dichotomous pattern of venation in the leaves ofUtricularia striatula is described in detail and compared withCircaeaster andKingdonia. Similar traits of dichotomous venation occur in other species ofUtricularia and may be due to reduction.Studies on IndianUtricularia, 3.     

Studies on the leaf anatomy of Euphorbia: II. Venation patterns*

Leaf venation patterns of 150 species of Euphorbia are presented and their value as a diagnostic feature in the genus assessed. Based on the gross venation patterns, the species have been grouped into uni-, bi-, tri-veined and special categories. The majority of the species studied belonged to the tri-veined category, in which ornamentation of the veins and the course of traces in the lamina proved useful additional characters. Features such as the size of the areoles, the number of vein endings, and their further ramifications and composition in each areole varied in the same leaf or in different leaves of a given species. Furthermore, no direct correlation could be established between the areole size and the numbers of vein endings and tips per areole. Forty species possess a parenchymatous vein sheath. Globular chloroplasts showed up conspicuously in the sheath cells of a few species. Dilated tracheidal elements, localized on the venules or at the tips of vein endings, are characteristic of the xerophytic species. In some instances, correlation was apparent between plants grouped according to their venation patterns and their habit.     

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.     

<Emphasis Type="Italic">KNOX</Emphasis> overexpression in transgenic <Emphasis Type="Italic">Kohleria</Emphasis> (Gesneriaceae) prolongs the activity of proximal leaf blastozones and drastically alters segment fate

KNOX (knotted1-like homeobox) genes have a widely conserved role in the generation of dissected leaves. Ectopic KNOX activity in leaves in various angiosperm lineages causes leaf form changes that can elucidate how the configuration of leaf development evolved. We present an analysis of leaf morphology and morphogenesis in transgenic Kohleria lines overexpressing a heterologous KNOX gene. Kohleria, like many members of Gesneriaceae, has simple-serrated leaves with pinnate venation. KNOX overexpression causes prolonged segment proliferation in proximal, but not distal, parts of leaf blades. Elaborate dissected segments reiterate the zonation of the whole leaf, with organogenic activity persisting between a distal maturation zone and a proximal intercalary elongation zone. The architecture of vascular bundles is severely altered, with a reduced midvein and a more palmate venation. The initial establishment of organogenically competent primordial margins (marginal blastozones) and the onset of tissue differentiation in early stages of leaf development were similar in wild-type and KNOX overexpressing lines. However, leaves overexpressing KNOX often failed to fully mature, and persistent marginal blastozones were found at the base of blades in mature portions of the shoot. We conclude that KNOX-mediated perpetuation of marginal blastozones in Kohleria is sufficient to induce a set of processes that result in highly dissected leaflets, which are unusual in this plant family. Spatial confinement of blastozones between an early maturing tip and a late elongating petiole zone reflects the presence of distinct maturation processes that limit the ability of the leaf margins to respond to ectopic KNOX gene expression.     

ANALYSIS OF PETAL VENATION IN RANUNCULUS. I. ANASTOMOSES IN R. REPENS V. PLENIFLORUS

Arnott , Howard J., and Shirley C. Tucker . (Northwestern U., Evanston, Ill.) Analysis of petal venation in Ranunculus. I. Anastomoses in R. repens v. pleniflorus. Amer. Jour. Bot. 50(8): 821–830. Illus. 1963.—The venation patterns of 1218 petals of Ranunculus repens v. pleniflorus were analyzed, with particular attention to the number and position of vein anastomoses. The essentially open dichotomous vascular pattern is complicated by the presence of vein fusions in 69.4% of the petals. The anastomosis ratio (number of anastomoses/number of appendages) is, therefore, high (1.37), compared to a maximal 0.61 for Ginkgo leaves. Of the petals having such fusions, 55.3% have more than 1 anastomosis; the maximal number of fusions found per petal is 8. The presence of anastomoses shows a high degree of correlation with petal size and petal lobing. The types of fusions (types A, B, C, D) are identical with those found in Ginkgo leaves, with the addition of 2 modifications of type C. Curiously, types C and D account for 98.7% of the total anastomoses, while types A and B are rare. An analysis of the location of each type within the petal shows that type C's are disproportionately numerous along the distal periphery, and that type-D fusions are unusually numerous in the central and basal regions. Such evidence suggests that the occurrence of a vein fusion is no “accident,” but rather that it is one manifestation of morphogenetic control.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

Leaf architecture of some acanthaceae

Leaf architectural pattern has been studied in 27 genera and 35 species of the Acanthaceae. The major venation pattern conforms to pinnate camptodromous with eucamptodromous or festooned brochidodromous secondaries, pinnate craspedodromous inAcanthus ilicifolius and acrodromous inLepidagathis trinervis. Intersecondary veins are common. The marginal ultimate venation is looped. The areoles are variable in size. The vein endings are usually simple, linear or cuved or divide once or twice dichotomously. Isolated vein endings, tracheids, isolated free vein endings and extension cells are observed. Transfusion tracheids are seen inDicliptera verticillata. Miniature vessel elements with simple perforation plates are noticed lying free in the areole or at the end of tracheids inSeriocalyx scaber andThunbergia grandiflora.Elytraria acaulis, Nelsonia canescens andStaurogynae zeylanica exhibit venation pattern shown by majority of the taxa studied, of the Acanthaceae.     

FOLIAR MORPHOLOGY AND ANATOMY OF THE GIGANTOPTERID PLANT DELNORTEA ABBOTTIAE,FROM THE LOWER PERMIAN OF WEST TEXAS

Delnortea is a monotypic genus (type-species: D. abbottiae) of Lower Permian gymnosperms based on leaves from uppermost Leonardian deltaic sediments exposed in the Del Norte Mountains, West Texas. The leaves are simple, symmetrical, mostly oblong or elliptical, and vary in length from 1.2 to about 35 cm. The petioles are short and stout, with a basally enlarged abscission zone. The margins are crenate, with a narrow, indurated border. Venation is in 4 orders: the secondaries and tertiaries are robust, unbranched, and pinnately arranged in a precise “herringbone” pattern, with the secondaries ending in the marginal sinuses; the quaternaries divide sparingly and fuse with others to form a dense reticulum of small meshes. Permineralized petiole and midrib material reflects a bifacial cambium, shown by a semicircular vascular arc, irregularly divided into several collateral bundles with secondary xylem and phloem. Delnortea is referable to the Gigantopteridaceae, a probably artificial family of gymnosperms incertae sedis with important venation features in common, but without known diagnostic reproductive organs. With Delnortea, the North American gigantopterids now include 5 genera, but Gigantopteris itself is lacking. Delnortea holds a relatively advanced evolutionary position among the American gigantopterids; its leaf morphology and gymnospermous anatomy entail intriguing points of comparison with Gnetum. The limited geographic and stratigraphic ranges and morphological distinctiveness of the American gigantopterids and associated taxa attest to rapid evolution and dispersal from a small area of origin in the southwestern United States during Leonardian time.     

CLASSIFICATION OF THE ARCHITECTURE OF DICOTYLEDONOUS LEAVES

A classification of the architectural features of dicot leaves—i.e., the placement and form of those elements constituting the outward expression of leaf structure, including shape, marginal configuration, venation, and gland position—has been developed as the result of an extensive survey of both living and fossil leaves. This system partially incorporates modifications of two earlier classifications: that of Turrill for leaf shape and that of Von Ettingshausen for venation pattern. After categorization of such features as shape of the whole leaf and of the apex and base, leaves are separated into a number of classes depending on the course of their principal venation. Identification of order of venation, which is fundamental to the application of the classification, is determined by size of a vein at its point of origin and to a lesser extent by its behavior in relation to that of other orders. The classification concludes by describing features of the areoles, i.e., the smallest areas of leaf tissue surrounded by veins which form a contiguous field over most of the leaf. Because most taxa of dicots possess consistent patterns of leaf architecture, this rigorous method of describing the features of leaves is of immediate usefulness in both modern and fossil taxonomic studies. In addition, as a result of this method, it is anticipated that leaves will play an increasingly important part in phylogenetic and ecological studies.     

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.     

Hydrodynamic Modelling Study of Angiosperm Leaf Venation Types

The morphology of leaf venation has been studied repeatedly and various systems have been proposed for the classification of the observed leaf venation patterns. Almost nothing is known, however, about the functional properties of the various venation types. Using a computer modelling approach we analysed the water transport properties of typical craspedodromous and brochidodromous venation patterns. The water transport through the leaf and the veins was modelled as a fluid flow through a porous medium and the mathematical model was solved with the Finite Element Method. The simulations illustrate that the leaf margin represents a critical region in terms of water supply. The results provide a plausible functional explanation for three well known phenomena: 1) the correlation between craspedodromous venation and the formation of leaf teeth; 2) the fact that craspedodromous venation is more common in temperate than in tropical regions and 3) the fact that xeromorphic leaves tend to have more closed venation.     

CHARLIEA MANZANITANA,N. GEN., N. SP., AND OTHER ENIGMATIC PARALLEL-VEINED FOLIAR FORMS FROM THE UPPER PENNSYLVANIAN OF NEW MEXICO AND TEXAS

Charliea is a new genus (type-species: C. manzanitana), based on pinnately compound leaf material from the richly fossiliferous Virgilian (Upper Pennsylvanian) shales of the Kinney Brick Company quarry near Albuquerque, New Mexico. In several features Charliea resembles Russellites or a zamioid cycad. It has linear-oblong pinnae with broad, oblique attachment and a truncate tip, which is deeply incised to form two to four nearly equal lobes. The venation is simple, parallel, and sparingly dichotomous, each vein ending at the distal margin. The Kinney beds also contain Plagiozamites planchardi, another zamioid form with parallel-veined pinnae, differing from Charliea chiefly in having rounded tips and veins ending in the denticulate margins. An unnamed third form (genus B) in the Kinney beds has long, narrow pinnae with parallel veins and blunt tips; this strongly resembles the Mesozoic conifer Podozamites, but may just as well represent a cycadophyte. Another unnamed taxon (genus A), from an Upper Pennsylvanian deposit in Jack County, Texas, resembles genus B or Russellites in general shape and venation, but the critical distal margins are unknown. In their single-ordered parallel venation, these four foliar types contrast sharply with the two-ordered pinnate venation of most Pennsylvanian fern-like leaves, and seem to foreshadow Mesozoic morphologies. This tendency toward precocious evolution of parallel-veined foliar form in North America is also expressed by a single occurrence of the Asiatic, Permian genus Tingia in the Lower Pennsylvanian of Utah, and by the presence of the predominantly Triassic cycadeoid genus Pterophyllum in the Lower Permian of Texas.     

独叶草叶二叉分枝脉序中网结脉和盲脉的形态学研究

对独叶草营养叶二叉分枝脉序及其中的网结脉和盲脉的形态学研究表明：(1)网结脉中2条完全汇合的与靠近脉中完全分离的叶脉之间未发现任何形式的维管束汇合的中间类型及网结脉中具有不同程度的连接脉退化痕迹的事实表明,网结脉不可能由靠近脉产生,相反,由于网结脉中联结脉的退化而形成开放脉;(2)盲脉是通过伴随着齿退化的达齿脉的退化、网结脉中联结脉的间断、非网结脉由分枝处间断三种方式产生的;(3)越裂片脉的出现及其可以形成网结脉的现象表明独叶草营养叶可能曾具有较为复杂的脉序,这种叶脉也呈现出退化的趋势;(4)独叶草营养叶的二叉分枝脉序可能是一种退化性状,而网结脉的出现可能是这种退化过程中的残留痕迹。     

Foliar architecture of vanilloid orchids: insights into the evolution of reticulate leaf venation in monocotyledons

Leaves of representative taxa within the vanilloid clade of Orchidaceae were cleared and their venation patterns studied. Within subtribe Pogoniinae, Isotria and Pogonia exhibit a prominent reticulate venation pattern, although only Pogonia ophioglossoides is characterized by free vein endings. Within subtribe Vanillinae, all species of Epistephium , as well as the New Caledonian endemics, Eriaxis rigida and Clematepistephium smilacifolium , have reticulate-veined leaves that are characterized by numerous free vein endings. Leaves of most species of Vanilla exhibit a parallel-veined pattern stereotypical of monocots, although branched secondaries with free endings were observed in V. africana. Most members of subtribe Galeolinae are 'saprophytes' with reduced bract-like leaves. Vascular bundles enter these leaf-homologues in a parallel manner, but quickly ram+ throughout the tissue in an irregular manner. Leaf venation is used to hypothesize patterns of generic relationships within the vanilloid clade. Molecular evidence for phylogenetic relationships among angiosperms indicates that reticulate leaf venation has arisen independently in several unrelated monocot families, including the vanilloid Orchidaceae, perhaps by a similar evolutionary scenario.     

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