THE NATURAL AFFINITY OF THE CARBONIFEROUS SEED,CALLOSPERMARION

A seed of Callospermarion pusillum has been found attached to a long slender stalk. Significant features of the seed based upon additional specimens are described. Secretory cavities in the integument are similar to those in the stem of Callistophyton poroxyloides and the fertile foliage bearing the pollen organ Callandrium callistophytoides. Vesicaspora sp. pollen similar to that found in Callandrium, occurs in the pollen chamber. Callistophyton, Callandrium, and Callospermarion are considered to represent the detached organs of a new seed fern family, Callistophytaceae.     

THE VEGETATIVE ANATOMY OF SCHOPFIASTRUM DECUSSATUM FROM THE MIDDLE PENNSYLVANIAN OF THE ILLINOIS BASIN

Schopfiastrum decussatum, a monostelic pteridosperm, has been recovered from Middle Pennsylvanian age coal balls from six localities within the Illinois Basin. Additional features of stem anatomy include the presence of horizontal sclerotic plates in the inner cortex, secretory ducts in the inner and outer cortex, and adventitious roots. The primary xylem is interpreted as bilobed, emitting massive leaf traces in a distichous manner. Five orders of frond members are described, including pinnules. All orders of the frond contain secretory ducts and tissues characteristic of Schopfiastrum stems. The rectangular, bilobed protostele, and method of leaf trace origin indicate that Schopfiastrum is more closely related to certain lower Carboniferous seed ferns than to contemporary Pennsylvanian pteridosperms.     

STUDIES OF PALEOZOIC SEED FERNS: ANATOMY AND MORPHOLOGY OF MICROSPERMOPTERIS APHYLLUM

The discovery of numerous specimens of the monostelic pteridosperm genus Microspermopteris in Pennsylvanian coal ball petrifactions from the Lewis Creek and What Cheer localities provides additional information about the anatomical and morphological variability within the genus. Specimens are now known up to 1.1 cm in diam that bear epidermal appendages in the form of variously-shaped trichomes. The external surface of the stem is further ornamented by longitudinal flaps of cortical tissue. Petioles exhibiting a single C-shaped vascular strand with abaxial protoxylem are produced in a 2/5 phyllotaxy. Large petiole bases that clasp the stem produce primary pinnae alternately. The presence of axillary branching appears similar to that reported in Callistopliyton and Lyginopteris. Triarch to polyarch adventitious roots, some with secondary tissues, are produced at both nodal and internodal regions. Of the currently recognized monostelic seed fern genera, Microspermopteris is most similar to Heterangium. Information is presented that supports current ideas regarding the evolution of the gymnospermic eustele from protostelic Devonian ancestors.     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

ANATOMICAL STUDIES OF CATHAYA (PINACEAE)

Cathaya Chun et Kuang is a monotypic genus and one of the gymnosperms endemic to China. We investigated Cathaya argyrophylla with both light and scanning electron microscopy to study the external and internal surfaces of leaf cuticle, leaf blade, petiole, shoot apex, young stem, bark, wood, young and old roots, and mycorrhizae. It is shown that Cathaya has unique characteristics as well as common features of the Pinaceae, there being a difference between Cathaya and Pinus and the rest of the family. So far as the vegetative organs are concerned, the genus is most closely related to Pseudotsuga and Larix. Data derived from the study of structures of vegetative organs of Cathaya are very different from those of reproductive organs, indicating the complexity of the problem of systematics and evolution in these plants. However, the present study supports the view that Cathaya should not be included in the genus Pseudotsuga as a new species.     

THE ANATOMY OF BARBACENIOPSIS,A NEW GENUS RECENTLY DESCRIBED IN THE VELLOZIACEAE

Although the two species in Barbaceniopsis—B. boliviensis and B. vargasiana—have in common many features which support their inclusion in one genus, there are also obvious differences in the depth of leaf furrows, absence of hairs in B. vargasiana, and type of girder sclerenchyma in the leaf. The presence of vessels in the leaf has been demonstrated. The frequent observation that the roots cling tightly only to stems of Vellozia species is shown to be characteristic of the family. A comparison of the three genera clearly indicates that there are obvious anatomical characters which help to distinguish Barbaceniopsis and Barbacenia from Vellozia. Although it is not suggested that Barbaceniopsis be reduced to synonymy, there can be no doubt that it is very closely associated with Barbacenia.     

THE MORPHOLOGY AND RELATIONSHIPS OF RHODEA,TELANGIUM, TELANGIOPSIS,AND HETERANGIUM

This study deals with four form or organ genera from the Upper Mississippian (Chester Series) of the Illinois Basin, and provides evidence that they were produced by a single natural genus with gymnospermous affinity. The plant remains—compressions, impressions, petrifactions, and specimens that combine compression or impression with petrifaction—allow examination of both external morphology and internal anatomy. The specimens include foliage corresponding to Rhodea, stems and petioles corresponding to Heterangium, and synangiate fructifications corresponding to either Telangium or Telangiopsis. The stems and foliage are considered parts of the same plant because of the identity of the anatomical and cuticular features of petioles attached to stem and those petioles with attached foliage. The fertile material is regarded as part of the same plant because: (1) The anatomy of axes of the fertile specimens is like that of the sterile specimens. (2) A single specimen may contain both sterile Rhodea-type axes and fertile regions. (3) Axes bearing synangia have the same size and patterns of divisions as the sterile foliage. Features that indicate lyginopterid affinities include: (1) Equal forking of the petiole. (2) Presence of fiber bands in the outer cortex and sclerotic nests in the inner part of the cortex. (3) Crowded circular bordered pits on the lateral walls of the metaxylem tracheids. (4) The presence of a small amount of secondary xylem. A variety of structural details of the stem and petiole suggest the genus Heterangium. The phyletic position of the plant that produced Rhodea, Telangium, Telangiopsis, and Heterangium is reviewed in light of such discoveries as the presence of a planated frond that lacks a lamina and the presence of both monolete and trilete microspores in a single synangium.     

A SYSTEMATIC ANATOMICAL STUDY OF THE GENUS PERIDERIDIA (UMBELLIFERAE-APIOIDEAE)

Certain anatomical features, especially of roots, leaves, and fruits, provide useful criteria for delimiting a genus and defining its component species. Most species of Perideridia have 1-5 multistelic tuberous roots; however, P. Howellii and P. Kelloggii possess fascicles of up to 20 thickened fibrous roots, each of which is monostelic and composed of four or five primary xylem strands arranged alternately with isolated strands of secondary vascular tissues. The curious multistelic condition of the tuber apparently has not been reported for any other group of dicotyledons. An accessory vascular bundle located internally to the median collateral bundle of the leaf rachis is found in only three species of Perideridia. Depending on the species, the leaf mesophyll may be either dorsiventral or isolateral. In about two-thirds of the species, the oil ducts of the fruit are solitary in the intercostal intervals and paired on the commissure; in the remaining one-third they are more numerous in both situations, rising to 3-5 in the intervals and 12-13 on the commissure in P. Pringlei. Variation in number of oil ducts in the intervals assumes taxonomic importance because it has been used as a generic criterion in this group, but the present study shows no correlation between the number of oil ducts and other characters.     

CRENATICAULIS,A NEW GENUS OF DEVONIAN PLANTS ALLIED TO ZOSTEROPHYLLUM,AND ITS BEARING ON THE CLASSIFICATION OF EARLY LAND PLANTS

The name Crenaticaulis verruculosus is proposed for slender, pseudomonopodially and dichoto-mously branching plants that bore opposite to subopposite sporangia along the stem and two rows of prominent, multicellular teeth on opposite sides of the terete axis. Epidermal cells were either narrow and elongate parallel to the stem or short with a papillate outer tangential wall. Axillary tubercles were present on one side of the stem near lateral branches. Some tubercles bore remnants of branches. Sporangia dehisced along their distal margins into two unequal halves. No spores were found. Occasional short lengths of stem were petrified by iron pyrite. Sections revealed a cortex consisting of four to six rows of thick-walled cells and a xylem strand. The strand was elliptical in transverse section and maturation was exarch. Tracheids were chiefly scalariform. It is suggested that the so-called axillary tubercles, known in several plants of Devonian age, were scars of rhizophores like those in the modern genus Selaginella. The plant is referred to the subdivision Zosterophyllophytina.     

ITOPSIDEMA,A NEW GENUS OF THE OSMUNDACEAE FROM THE TRIASSIC OF ARIZONA

Daugherty , Lyman H. (San Jose State College, San Jose, Calif.) Itopsidema, a new genus of the Osmundaceae from the Triassic of Arizona. Amer. Jour. Bot 47(9): 771–777. Illus. 1960.—Itopsidema vancleavei, a new genus and species of the family Osmundaceae, is described. The specimen consists of several segments of an arborescent stem obtained from the Upper Triassic of the Petrified Forest National Monument near Holbrook, Arizona. The surfaces on 2 of the segments are covered by adventitious roots and the remaining segments are covered by leaf bases. The fronds are spirally arranged and have an 8/21 phyllotaxy. The cortex of the stem, which contains numerous leaf traces and adventitious roots, consists of parenchyma with cell walls of medium thickness. The leaf traces are oblong to crescent-shaped in the inner cortex and horseshoe-shaped in the outer cortex. The base of the petiole contains a single, large vascular bundle and is covered by multicelled, glandular spines. The adventitious roots originate on the abaxial side of the leaf traces in the region of the inner cortex. The center of the stem is occupied by an ectophloic, mesarch siphonostele without leaf gaps. The pith is composed of firm-walled parenchyma cells which have isolated tracheids with reticulate pitting scattered among them. These tracheids are so rare the pith cannot be considered a “mixed pith.” The relationship and morphological significance of Itopsidema with respect to other members of the Osmundaceae are briefly discussed.     

SILICON IN THE ROOTS OF HIGHER PLANTS

Silicon (Si) distribution in the roots of Sorghastrum nutans (L.) Nash and Sorghum bicolor (L.) Moench. was investigated by means of the electron-probe microanalyzer and scanning electron microscope. In both species, Si was confined to the inner tangential wall of the tertiary-phase endodermal cells in the form of nodular silica aggregates of similar morphology and X-ray intensity. The results are compared to those for six closely related genera, as well as to studies of Si in the roots of species of other tribes of the family Poaceae. The various types of root deposits occurring in the family are described, and their relationships discussed. It is concluded that the type of Si distribution exhibited is determined largely by the phylogenetic status of the genus, rather than by the basic pattern of root anatomy.     

DEVELOPMENTAL ANATOMY OF JUVENILE AND ADULT SHOOTS OF MARCGRAVIA RECTIFOLIA L

Marcgravia rectifolia L. is a dimorphic vine having distinct juvenile and adult shoots. The juvenile shoot is a climber characterized by an orthotropic growth habit, a flattened stem, adventitious roots, and ovate leaves. The adult shoot, on the other hand, possesses a plagiotropic growth habit, has a cylindrical stem, few or no adventitious roots, and lanceolate leaves. Both phases have distichous phyllotaxy, however the plastochron is shorter for the adult phase than for the juvenile. Internode elongation occurs earlier for adult shoots than for juvenile shoots. Cytological analyses show the flattened stem of the juvenile results from differential production of cells, especially in the pith region. On the other hand, internodes of the adult phase are longer than juvenile internodes, a result of more cells produced rather than longer cells. In juvenile stems a perivascular band of elongated fibers develops, while in adult stems this band consists of brachyosclereids. Both phases undergo secondary growth and have non-storied cambia. Cambial activity begins in the 6th internode of each phase. As secondary growth proceeds, the adult stem produces much more xylem than juvenile stems of the same age. Adventitious roots produced in the juvenile stem are located in vertical rows at the “corners” of flattened stems and are attachment structures aiding the climbing habit of this vine. Phase changes occur regularly in this species. The juvenile phase usually transforms into the adult, however the adult phase can spontaneously revert back into the juvenile phase. The anatomical features and the phase changes are discussed and compared to Hedera helix, a vine whose phase changes have been studied in some detail. It is suggested that the anatomical features of Marcgravia rectifolia L. including its phase changes, may provide an alternative system to study physiological changes similar to those done with Hedera helix.     

EQUISETUM CLARNOI,A NEW SPECIES BASED ON PETRIFACTIONS FROM THE EOCENE OF OREGON

Equisetum clarnoi is described from four silicified stem fragments and numerous small roots from the Eocene Clarno Chert of Jefferson County, Oregon. Stems are up to 8.0 mm in diam and have sunken stomata arranged vertically in a single line flanking each of the external biangulate stem ridges, features that clearly ally this species with the subgenus Hippochaete. External stem ridges are equal in number to the carinal hypodermal bands. The hypodermis is composed of fibers and has prominent carinal bands up to 0.75 mm long and shorter vallecular bands. Cortical parenchyma cells enclose prominent vallecular canals which are lined by specialized thick-walled parenchyma cells. The double, common endodermis has prominent casparian strips. Vascular bundles are composed of four to seven metaxylem tracheids flanking each side of the phloem and protoxylem tracheids which occur singly on the internal surface of the small carinal canals. Leaf sheaths in cross section have an adaxial fibrous layer and an external or near external fibrous bundle. Roots are up to 2.0 mm in diam and have paired cuboidal epidermal cells from which root hairs arise. The stele of the root is central and shows exarch primary xylem maturation. Equisetum clarnoi most closely resembles the extant Equisetum hyemale var. affine.     

ANACHOROPTÉRIS INVOLUTA AND ITS ATTACHMENT TO A TUBICAULIS TYPE OF STEM FROM THE PENNSYLVANIAN OF IOWA

Hall , John W. (U. Minnesota, Minneapolis.) Anachoropteris involuta and its attachment to a Tubicaulis type of stem from the Pennsylvanian of Iowa. Amer. Jour. Bot. 48(8): 731–737. Illus. 1961.—Petioles referable to Anachoropteris involuta are described, attached to a stem which most nearly corresponds to a member of the genus Tubicaulis. These petioles are attached in a 2/5 phyllotaxy. At their points of departure petiole traces are massive and C-shaped but become involute in regions away from their attachment. Adventitious roots were borne on the stem in partial whorls. It is suggested that Anachoropteris petioles were extremely long and that they bore adventitious stems at intervals. These, in turn, bore adventitious roots, perhaps to serve for uptake of minerals or support in regions removed from the true stem. Such a “petiole unit” may also have functioned as a vegetative propagule. This would account for the apparent rarity of true stems and the abundance of petioles in coal balls.     

EVOLSONIA,A NEW GENUS OF GIGANTOPTERIDACEAE FROM THE LOWER PERMIAN VALE FORMATION,NORTH-CENTRAL TEXAS

Evolsonia is a new gigantopterid genus (type-species: E. texana), based on leaf impressions from 3 widely separated localities in north-central Texas. The leaves are associated with terrestrial vertebrates in floodplain sediments of the Vale Formation in the Clear Fork Group, of late Leonardian (Early Permian) age. Evolsonia leaves are simple, with very large elliptic laminae reaching 27 cm in width and 80 cm, possibly more, in length. Leaf margins are sinuate to crenate, with mostly shallow concavities. Venation is pinnate, in 4 orders; all but the ultimate veins are very thick and protrude below the lamina, creating deep impressions in the matrix. The secondary and tertiary veins form a precise herringbone pattern, with the secondaries and exmedial tertiaries ending at the leaf margin between concavities. The other tertiary veins are either simple or divided into 2 nearly equal divisions; they produce simple or variously divided quaternary veins that end at a thin sutural vein, forming a dense reticulum; some of the terminally branched tertiaries delimit areolelike areas that enclose several quaternaries and meshes. One of 6 presently known American gigantopterids, Evolsonia most closely resembles the younger Delnortea in gross architecture; with its sutures and dichotomously divided veins, however, Evolsonia is architecturally intermediate between Delnortea and the older American gigantopterids with forked leaves. Sedimentary features indicate deposition under alternating periods of flooding and drought. Their huge size invites comparison of Evolsonia leaves with those of modern tropical plants, whereas their thick veins and preservational features suggest thick, coriaceous texture.     

VASCULAR CONNECTION BETWEEN LATERAL ROOTS AND STEM IN THE OPHIOGLOSSACEAE

The vascular connection between lateral roots and stem in the Ophioglossaceae and in two leptosporangiate fern species was examined. Two types of connections were found: “gradual” connections, which resemble leaf traces in ontogeny and morphology, and “abrupt” connections, which resemble the connections between lateral roots and their parent roots. Gradual root-stem connections occur in the genera Ophioglossum and Helminthostachys and in Woodwardia virginica. They are initiated in shoot apices distal to the level where cauline xylem elements mature. They resemble leaf traces in being provascular (procambial) strands that connect the cauline stele with the future vasculature of lateral appendages. As with leaf traces, gradual connections are part of the provascular and, later, protoxylem continuity between stems and lateral appendages. Gradual connections have many features in common with leaf traces, and the term root trace is applicable to them. The order of radial maturation of the primary xylem in gradual connections varies in different parts of the connections. It is endarch near the intersection with the cauline stele and exarch where the connections intersect root steles. Gradual connections resemble the transition regions of certain seed plants where protoxylem is also continuous from stem to root and the order of maturation is found to change continuously from stem to root. Abrupt connections occur in Botrychium and Osmunda cinnamomea. They develop in shoot apices at levels where cauline xylem is mature or maturing. The mature xylem does not dedifferentiate, so provascular and protoxylem continuity of the kind found in root traces does not occur. Also, reorientation of the order of maturation does not occur in abrupt connections. Xylem connectors are found in the region where radially oriented elements of the connections abut the longitudinally oriented cauline elements. Abrupt connections resemble the connection of secondary roots with their parent root systems since xylem connectors and the lack of continuity are also features found in these vascular systems. The resemblance of the vascular pattern of the fern root trace to the transition region of seed plants suggests that the radicle is more closely comparable to the cladogenous roots of pteridophytes than hitherto supposed.     

THE ONTOGENY OF CARBONIFEROUS ARTICULATES: THE APEX OF SPHENOPHYLLUM

Twig apices of Sphenophyllum lescurianum, S. constrictum, and two new Sphenophyllum taxa are described in transverse and longitudinal section from middle and upper Pennsylvanian age specimens. In all of the species the single apical cell has the shape of a tetrahedron, with a triangular upper surface and three internal cutting faces. Segment cells are produced from each of the cutting surfaces in a dextrorse or sinistrorse direction, depending upon the species. The central portion of each segment cell contributes to the initiation of the procambium, while the remaining outer portion undergoes a vertical and subsequent horizontal division to form segment cells. Segment cells are aligned in vertical tiers beneath the respective apical cell cutting faces, with the individual leaves positioned directly beneath a tier of segment cells. Leaf primordia are first observed as a series of surface undulations below the apex, with an intercalary meristem located directly beneath each primordium. The vegetative apical organization of Sphenophyllum is demonstrated to be very similar to the type of organization found at the stem tips of Catamites and Equisetum.     

THE SYSTEMATICS AND ANATOMY OF GONGYLOCARPUS (ONAGRACEAE)

The unique fruiting structures of the closely related, principally Mexican, monotypic genera Gongylocarpus and Burragea (Onagraceae) compelled a detailed anatomical and cytological investigation of these plants which led to the conclusion that they should be included in a single genus, Gongylocarpus. Gongylocarpus fruticulosus (Burragea), endemic to two adjacent islands off the west coast of Baja California, is divided into two subspecies, subsp. fruticulosus and subsp. glaber. The vegetative and floral anatomy, including wood anatomy, of both species is described. The fruits of these two species grow into the stem by meristematic activity during the course of ontogeny, the ovaries in the mature flower being superficial and sessile in the leaf axil. There is no pedicel associated with the flower, but only a branch gap. Meristematic tissue at the base of the locules divides rapidly at a relatively late ontogenetic stage, the ovaries growing downward into the stem and crushing the pith. The mature, heavily sclerenchymatous fruits are located wholly within the stem, and in G. fruticulosus they are aggregated into long chains. Both species have a gametic chromosome number of n = 11, a characteristic otherwise unknown in the tribe Onagreae but shared with other generalized groups in the family. Taken together with other features, this suggests a primitive position within the tribe for Gongylocarpus.     

STUDIES OF PALEOZOIC FERNS: TUBICAULIS STEWARTII SP. NOV. AND EVOLUTIONARY TRENDS IN THE GENUS

Eggert , Donald A. (Yale U., New Haven, Conn.) Studies of Paleozoic ferns: Tubicaulis stewartii sp. nov. and evolutionary trends in the genus. Amer. Jour. Bot. 46(8): 594–602. Illus. 1959.—Tubicaulis stewartii, a new species of the order Coenopteridales is described. The specimen was derived from the Upper Pennsylvanian of Berryville, Illinois, and is characterized by having a lacunar middle cortex, a well-developed integumentary system bearing uniseriate hairs, and xylem parenchyma organized into vertically anastomosing strands. In addition, multiseriate (somewhat transitional to reticulate) bordered pitting is present in the petiolar metaxylem elements, while those of the stem stele are multiseriate scalariform. The habit is intermediate between that of a form such as Osmunda and a tree fern, having an upright tapering stem which gives off prominently decurrent petioles in a 2/5 divergence. A reinvestigation of the type specimen of the most closely allied species, T. multiscalariformis, of Upper-Middle Pennsylvanian age, has shown that it has similar features in the cortex, metaxylem, and integumentary layers. Tubicaulis multiscalariformis and T. stewartii form a distinct group in the 6 species now known, whose evolution has most likely involved the retention of a more primitive form of pitting (multiseriate scalariform) with parenchymatization of the xylem. The remaining species of the genus have not developed xylem parenchyma but have developed circular bordered pitting. The relationships of the genus to other genera in the Coenopteridales remain obscure.