PHLOEM ANATOMY IN STAUROPTERIS BISERIATA FROM THE PENNSYLVANIAN OF NORTH AMERICA

Phloem anatomy in the coenopterid fern Stauropteris biseriata is detailed from Lower-Middle Pennsylvanian coal ball specimens from eastern Kentucky. Axes exhibit a cruciate-shaped xylem trace in transverse section. Phloem tissue completely surrounds the xylem, but is more extensively developed in the embayments between the xylem arms. Phloem is composed of elongate conducting elements with a few scattered parenchyma cells. Large and small sieve cells are present, with larger ones occurring in the embayments within the primary plane of symmetry of the axes. Large elements are approximately twice the diameter of the smaller sieve elements. Oval sieve areas and pores have been observed on lateral and oblique end walls of both large and small elements. The structure and composition of Stauropteris phloem is discussed in relationship to the available information on phloem anatomy in other fossil cryptogams.     

ON THE STRUCTURE OF CALAMOSTACHYS BINNEYANA FROM THE LOWER PENNSYLVANIAN OF NORTH AMERICA

The discovery of specimens of Calamostachys binneyana in Lower Pennsylvanan petrifaction material in North America has provided additional information about the structure of this calamitean fructification. The cones consist of regularly spaced alternating whorls of bracts and sporangiophores. Bracts are fused in a disc except at the margin where the individual units become free. Sporangiophores are inserted at right angles to the cone axis and bear four axially directed sporangia. The vascular system of the North American specimens differs from that in other reports of the taxon in the presence of twelve vascular bundles in the cone axis. Each sporangiophore is supplied by a single vascular trace that departs from one axial bundle. There appears to be no constant relationship between the number of vascular bundles and the number of bracts. Spores are spherical, thin-walled, and of the Calamospora type. Relationships with other structurally preserved members in the genus are discussed in light of the diversity in structure demonstrated by the new specimens.     

ON A NEW STRUCTURALLY PRESERVED ARBORESCENT LYCOPSID FRUCTIFICATION FROM THE LOWER PENNSYLVANIAN OF NORTH AMERICA

Bisporangiate lycopsid fructifications are described from petrifaction material of Pottsville (lower Pennsylvanian) age collected in eastern Kentucky. The largest specimen of Lepidostrobus schopfii sp. n. is 8.0 cm long and approximately 1.3 cm in diam; it is not complete at either apex or base. Basal megasporangia, each containing a variable number (12-29) of megaspores, and apical microsporangia are adaxially positioned on pedicels bearing reduced lateral laminae. Features of the cone axis and pattern of vascularization are described. Of particular interest is the occurrence of well-preserved endosporic megagametophytes showing archegonia and rhizoids. The new species is compared with other structurally preserved bisporangiate lycopsid fructifications of equivalent age.     

FERAXOTHECA GEN. N., A LYGINOPTERID POLLEN ORGAN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The pollen organ Feraxotheca gen. n. is described from Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. The fructifications consist of bilaterally symmetrical synangia composed of a basal pad supporting elongate sporangia that are laterally appressed for the entire length of the sporangial cavities. Sporangial tips extend over the center of the synangium and delimit a small open area, while the bases arise from a parenchymatous cushion that is bounded by short tracheid-like cells. Each synangium is borne on the surface of an expanded pinna tip and is surrounded by a small amount of laminar tissue that envelopes the base of the synangium. Ultimate pinnae are rectangular in transverse section, possess an elliptical vascular bundle surrounded by canals containing a yellow froth-like substance, and have a cortex of elongate cells that radiate from the center of the axis. Sporangia contain small (40–64 μm), radial, trilete spores ornamented by regularly spaced coni or blunt tipped grana. Feraxotheca is compared with the compression genus Crossotheca and some new ideas are advanced concerning the morphology of this compression genus. The obvious differences between Feraxotheca and other lyginopterid pollen organs strongly suggests that the Lyginopteridaceae, as it is currently interpreted, is an unnatural family.     

ETAPTERIS LECLERCQII SP. N., A ZYGOPTERID FERN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The zygopterid fern Etapteris leclercqii sp. n. is described from Lower Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. Isolated petioles extend up to 15.0 cm in length and 5.0 mm in diam; no lateral axes have been observed. The clepsydroid-shaped petiole trace is characterized by lateral arms that sharply taper and by the production of peripheral loops prior to the separation of pinna traces from the stele. Large irregularly shaped multicellular hairs are randomly scattered over the surface of the petioles. When compared with other currently recognized zygopterid ferns, E. leclercqii appears most similar to Metaclepsydropsis duplex. Based on the configuration of the leaf trace and the occurrence of peripheral loops, it is suggested that E. leclercqii may represent the most primitive species of the genus known to date.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE MIDDLE EOCENE OF NORTH AMERICA: FLOWERS OF THE CELTIDOIDEAE

Flowers from the Middle Eocene Claiborne Formation of western Tennessee have both floral and palynological features suggestive of affinities with the subfamily Celtidoideae of the Ul-maceae. Floral size, pollen size, and exine structure suggest that these flowers are intermediate between insect-pollinated ancestors and the modern wind-pollinated Celtidoideae. Thus, they represent the first direct fossil evidence supporting the widespread notion that at least some of the Amentiferae have been derived from insect-pollinated ancestors.     

STUDIES OF PALEOZOIC MARATTIALEANS: NEW SPECIES OF SCOLECOPTERIS (MARATTIALES) FROM THE PENNSYLVANIAN OF NORTH AMERICA

Two new species of the late Paleozoic fern Scolecopteris (Marattiales) are described and their relationships within the genus are discussed. Scolecopteris charma sp.n., from Steubenville, Ohio (Duquesne Coal, Upper Pennsylvanian), is similar to species in the Oliveri group, while S. gnoma sp.n. from Providence, Kentucky (Baker Coal, Middle Pennsylvanian), compares favorably with the Latifolia species group. Scolecopteris gnoma is most similar to S. fragilis but differs in its smaller synangia and spore type. S. charma appears generally similar to S. iowensis because of its large pedicel and histologically undifferentiated walls, but differs in a number of characters such as vasculature and spore type. Despite its occurrence late in the Pennsylvanian, S. charma is thought to possess a number of primitive character states (large trilete spores, vascularized pedicels, flat pinnules with downturned margins). Using the same criteria for the much older S. gnoma, we note a number of relatively apomorphic character states (small monolete spores, unvascularized pedicels, extended pinnule margins). An outgroup analysis of species-level characters of Scolecopteris gives a better concept of primitive versus derived traits in marattialean and other ferns. Genera in the Paleozoic fern orders Filicales (Ankyropteris) and Zygopteridales (Corynepteris, Musatea) were chosen as outgroups, and the comparisons support suggestions for the polarity of several important characters. Some of these agree with previously proposed evolutionary polarities based on the geological occurrence of marattialean ferns.     

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.     

FURTHER OCCURRENCES OF SPENCERITES FROM THE MIDDLE PENNSYLVANIAN OF KANSAS AND ILLINOIS

New specimens of Spencerites moorei add knowledge of its sporophylls and method of sporangial attachment. The sporophyll consists of a slender pedicel which expands at its distal end into a fleshy, peltate, diamond-shaped head which bears a presumably fleshy lamina. The sporangium is attached through most of its length to the adaxial surface of the pedicel rather than distally as in S. insignis. It is suggested that the distal attachment shown in S. insignis may be derived through a phyletic shift from the pedicellate position. Additional specimens of Spencerites provisionally assigned to S. majusculus are also recorded. The spores are bilateral, monolete, and shaped like quadrants of a sphere. Two prominent wings extend along the two contact faces and the monolete mark is prominently raised, thus imparting a three-winged appearance to the spore. The axis consists of an exarch protostele and a cylinder of thick-walled outer cortex.     

A NEW PTERIDOSPERM MALE FRUCTIFICATION FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Rhetinotheca tetrasolenata gen. et sp. n. is described from a Herrin #6 coal ball from Illinois. It consists of an ellipsoidal cluster of small synangia bearing spores of the Monoletes type. Although the synangia are partially connected to each other by sparse sterile tissue, evidence indicates that the synangia are immature, and it is presumed that they separated and spread apart at maturity. A small portion of a protostelic axis is present near the center of the cluster. Individual synangia consistently contain four sporangial tubes and measure 2.0–3.6 mm long by 0.7–1.2 mm in diam. A conspicuous central columella is present. When compared with compression forms, Rhetinotheca tetrasolenata compares most favorably with Aulacotheca iowensis. On this basis, arguments are advanced disputing the classical concept of whittleseyan fructifications. It is contended that none of them possessed a hollow central cavity as depicted in most reconstructions.     

PHILLIPOPTERIS GEN. NOV.—ANATOMICALLY PRESERVED SPORANGIAL FRUCTIFICATIONS FROM THE UPPER PENNSYLVANIAN OF THE APPALACHIAN BASIN

The discovery of a new type of sporangial fructification in coal balls from the Upper Pennsylvanian of Ohio provides the basis for describing Phillipopteris globiformis gen. et sp. nov. Sporangia are borne terminally on up to two orders of branching axes. Penultimate axes branch pinnately to produce irregularly branched ultimate axes. Sporangial wall cells are of a single type and show no specialization for dehiscence. Spores are radial and trilete, and reminiscent of the sporae dispersae genus Dictyotriletes. Phillipopteris increases our knowledge of diversity among fernlike plants from the late Paleozoic, and shares several features with Sclerocelyphus Mamay.     

PALAEOSTACHYA DIRCEI N. SP., AN AUTHIGENICALLY CEMENTED EQUISETALEAN STROBILUS FROM THE MIDDLE PENNSYLVANIAN OF SOUTHERN ILLINOIS

Equisetalean strobili normally are encountered as disarticulated organs. This condition has necessitated the creation of 12 form genera to accommodate the various morphological architectures and anatomical configurations. Taxonomically useful characteristics, which are discernable in permineralized specimens, are rarely distinguishable in coalified compressions due to their destruction during diagenesis and coalification. Therefore, genera established on the position of sporangiophore-trace origin, such as Schimperia Remy and Remy, are untenable when dealing with coalified compressions. Although the two largest genera of strobili, Calamostachys Schimper and Palaeostachya Weiss, may be synonymous, it is proposed that these form genera be retained when dealing with coalified compression specimens. Each genus provides a particular architectural concept for specimens which may not be assignable to the species level. Calamostachys and Palaeostachya are highly overspeciated genera. It is suggested that characteristics necessary to delimit new species include bract height, degree of bract fusion, disposition of bracts, bract: sporangiophore ratio, number of sporangia per sporangiophore, and sexual status.     

TRIGONOCARPUS LEEANUS,A NEW SPECIES FROM THE MIDDLE PENNSYLVANIAN OF SOUTHERN ILLINOIS

A new species of Trigonocarpus Brongniart is described from the level of the Herrin (No. 6) Coal (Carbondale Formation, Kewanee Group) at Carterville, Illinois. The seed is three dimensionally preserved by authigenic cementation and exhibits a well preserved nucellar cast, integument and micropylar region. The specimen represents the largest pteridosperm compression-impression seed collected in North America and measures 10 cm from the apex of the micropyle to the chalazal end, and at least 5 cm in breadth. Trigonocarpus leeanus sp. n. is compared to the remaining forty-three taxa within the genus, and specifically with Trigonocarpus grandis Lesquereux, the one species with dimensions approaching it. An emended diagnosis, with designation of a lectotype, is presented for Trigonocarpus grandis. Correlation of the features displayed by Trigonocarpus leeanus sp. n. and the petrified taxa within Pachytesta Brongniart is attempted.     

NEW TRILOBITES FROM THE MIDDLE CAMBRIAN OF NORTH CHINA

The present paper deals with six new genera,one new subgenus,twenty new species andthree new varieties of trilobite found from the Middle Cambrian formations of North China.Most of them were once enlisted in the writer's foregoing paper(Chang,1957),in which a briefaccount of the stratigraphic sequence has been given.It now seems desirable to describe thosenew forms for further reference.     

AN ATLANTIC MIDDLE CAMBRIAN FAUNA FROM NORTH GREENLAND

A late Middle Cambrian fauna from Nyeboe Land, North Greenland, is described. It is demonstrated that the fauna, dominated by agnostid trilobites, belongs to the Atlantic province. The samples indicate the presence of the zones of Ptychagnostus punctuosus and Jincella brachymetopa. The distribution of the Middle Cambrian faunas is briefly discussed.     

MORPHOLOGICAL VARIATION,SPECIES CONCEPTS,AND CLASSIFICATION OF AN UNUSUAL FOSSIL CENTRIC DIATOM (BACILLARIOPHYTA) FROM WESTERN NORTH AMERICA1

The classification system of diatoms (Bacillariophyta) is based almost entirely on the organization and structure of their silica cell walls. We have discovered Miocene freshwater fossils that contain valves of two different orders within the same organism. Within frustules of the genus Ellerbeckia Crawford, a genus of the order Melosirales, are contained frustules of the genus Actinocyclus Ehrenberg, a member of the Coscinodiscales. Based on the abundance of each morphotype, they appear to be able to reproduce themselves. The ability of a single diatom taxon to express this degree of morphological variability is unprecedented. These results suggest species concepts in diatoms may be in need of review and challenge the basis of the classification system for this widely distributed and ecologically important group of organisms.