TRILOBOXYLON ASHLANDICUM GEN. AND SP. N. FROM THE UPPER DEVONIAN OF NEW YORK

A new taxon is described from the Upper Devonian, Oneonta formation in the northern Cats-kills of New York. Triloboxylon gen. n. is represented by petrifactions showing two orders of branching. The main axis bears branches spirally and the latter bear the ultimate appendages spirally. Ultimate appendages branch dichotomously twice, in one plane. Primary xylem of the main axis and branches is three-armed with mesarch protoxylem extending in a continuous band within each arm. Primary xylem of the ultimate appendage is terete and dichotomizes twice. Metaxylem elements are characterized by scalariform and circular-bordered pitting on all walls. The cortex is composed entirely of isodiametrical parenchyma cells. Triloboxylon is compared with other genera with three-lobed protosteles. Its possible affinity with the Aneurophytales is shown. The morphological nature of the “frond” of the Aneurophytales and the possibility that the group possesses the morphological equivalent of simple leaves are discussed.     

PROTEOKALON A NEW GENUS OF PROGYMNOSPERMS FROM THE DEVONIAN OF NEW YORK STATE AND ITS BEARING ON PHYLOGENETIC TRENDS IN THE GROUP

Proteokalon gen. nov. is described from the Upper Devonian Catskill deposits of New York. Two orders of branching and ultimate appendages are preserved' by petrifaction and by compression. The first order bears branches decussately and has a skewed four-armed protostele that occasionally dichotomizes. Second-order branches dichotomize rarely and most have T-shaped or three-armed protosteles. They bear ultimate appendages alternately, either in lateral pairs, or singly from the abaxial side. These appendages divide several times in one plane. Their vascular strand is terete. Maturation of the primary xylem is mesarch, and it consists of tracheids and parenchyma. Secondary xylem and phloem and a periderm are present. The outer cortex has a system of hypodermal fibers. Proteokalon is most similar to Tetraxylopteris and Triloboxylon of the Aneurophytales. A comparison of the stratigraphic occurrence of Protopteridium, Aneurophyton, Tetraxylopterism, Sphenoxylon, Triloboxylon, and Proteokalon suggests some evolutionary trends among the Aneurophytales.     

ACTINOXYLON BANKSII GEN. ET SP. NOV.: A PROGYMNOSPERM FROM THE MIDDLE DEVONIAN OF NEW YORK

A new genus from a Middle Devonian locality near Cairo, N. Y., is described. Actinoxylon gen. nov. is based upon pyritic petrifactions. Three orders of branching are present: penultimate branch, ultimate branch, and leaf. The penultimate branch bears spirally arranged ultimate branches and leaves, the leaves apparently replacing the branches in the spiral. The ultimate branches bear opposite to subopposite and decussate leaves. The leaves are non-planated, unwebbed structures which show at least three dichotomies. Each segment of the leaf is terete as are all other axes. Internally the penultimate branch has a six-lobed actinostele with mesarch protoxylem areas, one or two per lobe. Secondary xylem is visible in the oldest parts of several specimens. The xylem has helical-reticulate, reticulate, scalariform and circular-pitted elements. The presumptive areas of phloem are occupied by cells with dark contents. The cortex is composed of a parenchymatous inner region and a sclerenchymatous outer region. The ultimate branch traces are at first three-lobed protosteles, later becoming four-lobed. Several ultimate branch traces also possess secondary xylem while within the cortex of the penultimate branch. The leaf traces are terete strands. Below each forking of a leaf segment there is a corresponding forking of the vascular strand. Actinoxylon is compared with the progymnosperms Actinopodium, Svalbardia, Archaeopteris, Siderella, and Tetraxylopteris. The anatomy of the penultimate branch of Actinoxylon is similar to that of Actinopodium, Archaeopteris macilenta, and Siderella. The ultimate branch traces of Archaeopteris and Actinoxylon are similar. The ultimate branch stele and pattern of trace formation in Actinoxylon is similar to the stelar configuration and trace formation in the r + 2 axes of Tetraxylopteris schmidtii. The unwebbed leaves are similar to those of Archaeopteris fissilis, Svalbardia, and the terminal units of the Aneurophytales.     

RELLIMIA THOMSONII FROM THE GIVETIAN OF NEW YORK: SECONDARY GROWTH IN THREE ORDERS OF BRANCHING

Pyritized axes of the vegetative branching system of Rellimia thomsonii, from the Givetian of New York were described with emphasis on the presence and position of the secondary tissues in relation to the known axis orders. The morphology and anatomy of four axis orders (N, N + 1, N + 2, N + 3) was described using two well-preserved specimens. One specimen showed the largest known (N) axis order, with spirally attached N + 1 axes. The other specimen, from a more distal portion of the plant, showed the N + 2 axis order with spirally arranged N + 3 axes. The shape of the primary xylem, discernible in three of the axis orders (N + 1, N + 2, N + 3), was a three-lobed protostele with mesarch maturation of the primary xylem. The protostele of the N + 3 axis order, with only primary growth, was identical to that of fertile N + 3 axes. The other axis orders (N + 2, N + 1, N) had wood surrounding the primary xylem. The wood of the N+2 axis was limited, and occurred only in the basal portion of the axis, but that of the N+1 and N axes was extensively developed with two and three growth layers, respectively. This wood was pycnoxylic with narrow rays similar to other aneurophytalean progymnosperms. Surrounding the secondary xylem was secondary phloem indicating the vascular cambium in Rellimia was bifacial. The presence of growth layers was discussed in relation to the climatic interpretation placed on their presence and how it correlates with paleogeographic reconstructions of the Devonian.     

STENOKOLEOS BIFIDUS SP. N. IN THE UPPER DEVONIAN OF NEW YORK STATE

Stenokoleos is a genus for petrified axes from the Mississippian New Albany Shale to which an Upper Devonian occurrence in New York is added. Two orders of branching were known and the plant was thought to be related to coenopterid ferns. The new petrified axes from New York reveal three orders of branching. A pair of rachides emerges from one side of the stem at each node. Their position alternates at successive nodes (distichous). Each rachis bears alternately arranged pinnae. The shape of the xylem strand and the number of protoxylem areas are variable. Traces to the pairs of rachides arise either as two separate strands or as a single strand that is presumed to divide while still within the cortex of the stem. Traces to pinnae are ellipsoid or clepsydroid. Tracheids are scalariform and uni- or biseriate, circular-bordered pitted. Peripheral loops are present in all orders of branches. Protoxylem strands are numerous and maturation is mesarch. Cortex is parenchymatous where it is preserved but outer cortex is missing. Stenokoleos and Reimanniopsis are placed in a new family, Stenokoleaceae. This is classified as Incertae Sedis among Pterophytina in Tracheophyta. It is suggested that the plant is related more closely to the Mississippian pteridosperms Tristichia and Tetrastichia than to the coenopterid ferns.     

THE CAIRO FLORA (GIVETIAN) FROM EASTERN NEW YORK. I. REIMANNIA,TERETE AXES,AND CAIROA LAMANEKII GEN. ET SP. N.

A collection of over 200 petrified Middle Devonian plants was made from a single locality near Cairo, New York. This paper represents the second of a series enumerating the plants of the flora. Reimannia aldenense with a single lateral appendage containing a terete xylem strand that divides is present. Reimannia is thought to represent young branches within the Aneurophytales. Many of the sections made for the study of the flora contain small, terete axes. Some have a very distinctive epidermis. Some dichotomize several times. Cairoa lamanekii gen. et sp. n. shows closest affinity to the Aneurophytales yet differs in that the shape of the primary xylem is not consistently repeated from one order of axis to the next. It is suggested that Cairoa and Proteokalon represent a distinctive subgroup in the Aneurophytales.     

Morphology and ultrastructure of the sensory appendages of a kinorhynch introvert

The structure and arrangement of appendages (scalids) on the head of the homalorhagid kinorhynch Kinorhynchus phyllotropis Brown & Higgins, 1983 are named, described and illustrated. In adults of this species, seven rings of external scalids are separated by segment boundary structures from the oral styles and three rings of internal scalids. All of these appendages contain ciliated receptor cells which pass to pores at the scalid tips, and all but the two anterior rings are jointed. All of these appendages are radially arranged in multiples of five, and closely associated with the ten lobed circumoral nerve ring. The most posterior scalid ring consists of fourteen trichoscalids, of which six are longer than the other eight. The six longer trichoscalids are arranged in bilateral symmetry, two to each of the three facets of the body. Similarities between scalid arrangement in Kinorhyncha and Loricifera are discussed.     

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.     

A new euphyllophyte Kunia venusta gen. et sp. nov. from the Middle Devonian of Yunnan,South China

Abstract Kunia venusta gen. et sp. nov. is reported from the late Middle Devonian (Givetian) Haikou Formation near Kunming City, Yunnan Province, China. This plant has three orders of naked axes that divide pseudomonopodially. The second‐ and third‐order axes occur in a helix. Fertile appendages are distantly spaced and helically inserted to the third‐order axis; they comprise equally dichotomous branches terminated by two clusters of paired and fusiform sporangia. Sterile appendages are dichotomous and distally recurved. A comparison is made with the basal euphyllophytes including the trimerophytes, cladoxylopsids, zygopterids, stauropterids, and some relevant genera of uncertain affinity. The new plant resembles them in dichotomous appendages with terminal elongated sporangia, but differs mainly in the three orders of pseudomonopodial axes bearing helical laterals. It is thus placed in the Euphyllophytina as incertae sedis. It is suggested that an evolutionary divergence in the branching pattern and appendage morphology might have occurred in the Middle Devonian euphyllophytes, that is, maintaining three dimensions versus yielding more or less planation.     

CALAMOPHYTON IN THE MIDDLE DEVONIAN OF NEW YORK STATE

Specimens of Calamophyton from the Middle Devonian Ashokan Sandstone near Kingston, Ulster County, N.Y., are shown to belong to C. bicephalum Leclercq and Andrews, a Belgian species. Steel needles and a stereoscopic binocular microscope were used to follow the path of the forking leaves and branching sporangiophores of the specimens through the matrix. The terete leaves dichotomized two to three times in more than one plane. Sporangiophores dichotomized once. Each branch bore three recurved lateral branches which in turn bore two sporangia. Each branch terminated in an elongate, filiform projection. Sporangia apparently dehisced longitudinally. Their walls were composed of elongate cells. Their spherical spores ranged from 86 to 166 μ in diameter and bore a trilete mark. Ornamentation consisted of coni and spinae up to 4.5 μ long. They resembled dispersed spores of Dibolisporites gibberosus var. major Richardson. This is the second occurrence of Calamophyton bicephalum and the first account of its spores. It is the second report of the genus in North America.     

BOSTONIA PERPLEXA GEN. ET SP. NOV., A CALAMOPITYAN AXIS FROM THE NEW ALBANY SHALE OF KENTUCKY

Bostonia perplexa, gen. et sp. nov., was collected from the Lower Mississippian Falling Run member of the Sanderson Formation. The single short segment of an axis, preserved as a petrifaction, contains at least three vascular columns, each with both primary and secondary tissues. Primary xylem is two or three ribbed, and contains several mesarch protoxylem strands. Gymnospermous secondary xylem is characterized by both uniseriate and multiseriate rays. The ground tissue is parenchymatous except for a few clusters of sclerotic cells. In its apparent polystelic nature, the specimen superficially resembles members of the Pennsylvanian to Permian Medullosaceae. All evidence currently available, however, leads to the conclusion that this species should be placed in the Upper Devonian to Lower Mississippian Calamopityaceae. It has not been determined with certainty whether the species is polystelic (in the sense of the Medullosaceae), or whether the apparent polystely is the result of stelar branching proximal to the level of branch divergence.     

New red alga, Acanthopeltis longiramulosa sp. nov. (Gelidiales, Rhodophyta) from Jeju Island, Korea

A new red alga from Jeju Island, Korea is described. The alga is assigned to Acanthopeltis, Gelidiaceae by the characteristics of terete erect axes, sympodial growth, and suborbicular branchlets that are amplexi‐caul at the base. Acanthopeltis longiramulosaY. Lee et Kim is characterized by a discoid holdfast with a few stolons, rhizoids arising from the apex of the branchlet, an erect terete axis diverging into a few branches, longish obpyriform to lanceolate branchlets with smooth surfaces, and tetrasporangial stichidia or sper‐matangial and cystocarps appendages on the marginal region of the branchlet. A. longiramulosa is more closely related to Acanthopeltis japonica than Acanthopeltis hirsuta in terms of thallus morphology. However, A. longiramulosa is easily distinguished from A. japonica, which has suborbicular branchlets with setaceous processes on both surfaces.     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

中国长足兰属(兰科)一新记录

报道了中国兰科植物一新记录, 即筒距长足兰[Pteroceras teres （Blume） Holttum], 并提供了描述与图片。该种与滇南长足兰[P. compressum （Blume） Holttum]相似, 但以花序轴圆柱形; 花黄色, 螺旋状排列; 距的先端囊状, 两侧压扁并多少扭转等特征与之区别。     

ANATOMY AND RELATIONSHIPS OF SOME DEVONIAN PROGYMNOSPERMS FROM NEW YORK

New specimens of Triloboxylon ashlandicum Matten and Banks (1966) show additional features of the anatomy and morphology of the plant. The primary xylem has strands of parenchyma associated with trace formation. Secondary xylem and phloem are described for the first time. The outer cortex, previously unknown, contains hypodermal fiber strands. The ultimate appendages are much divided and planated. Specimens of Aneurophyton hallii (Arnold, 1940) were reexamined and newly prepared. They are transferred to the genus Triloboxylon as T. hallii. Triloboxylon is transferred from Pteropsida-Incertae Sedis to the Aneurophytales. New information on the morphology and anatomy of Telraxylopteris is presented and its identity to Sphenoxylon is supported.     

SVALBARDIA BANKSII SP. NOV. FROM THE UPPER DEVONIAN (FRASNIAN) OF NEW YORK STATE

Svalbardia banksii sp. nov., is described from the Upper Devonian (Frasnian) Fish Cabin Creek locality near Pond Eddy, New York. The compression consists of ultimate branches bearing spirally arranged, unwebbed leaves. The unwebbed leaves are up to 3.2 cm long, are somewhat flexuous, and dichotomize in more than one plane. The new species is similar to the reconstruction of Actinoxylon banksii from the Middle Devonian (Givetian) of New York and strengthens the supposed relationship between Svalbardia polymorpha and Actinopodium nathorstii from Mimerdalen, Spitzbergen.     

An anatomically preserved lycopsid from the Upper Devonian of South China

An anatomically preserved lycopsid, Lobodendron fanwanensegen. et sp. nov. Liu, Wang, Xue & Meng, is described from the Upper Devonian (Famennian) Wutong Formation of Changxing County, Zhejiang Province, China. The fossil plant bears slender, dichotomously branched axes. The vascular strand consists of solid terete primary xylem and lobed secondary xylem, which implies the result from the activity of possibly discontinuous cambium. The new plant has character combinations that do not conform to any branches in the canopy of the tree-lycopsids known previously, but resemble those of the basal part of some pseudoherbaceous lycopsids. This new plant may exemplify a Late Devonian lycopsid with a pseudoherbaceous growth habit.     

HALLETHECA RETICULATUS GEN. ET SP. N.: A SYNANGIATE PENNSYLVANIAN PTERIDOSPERM POLLEN ORGAN

A new structurally preserved synangiate pollen organ is described from the upper Pennsylvanian (Mattoon Formation) of southeastern Illinois. The specimen of Halletheca reticulatus gen. et sp. nov. measures approximately 1.5 cm long by approximately 5.0 mm in diam, and consists of five elongate sporangial tubes equidistantly arranged around a solid central column, and embedded in a thin-walled ground tissue. The vascular system consists of five terete bundles characterized by scalariform tracheids. Sporangia are thick-walled and contain pollen of the Monoletes-type. Ultrastructural studies of the pollen grain wall show it to consist of a uniform network of muri which branch and re-unite to form small lumina. A comparison of the ultrastructure of the pollen grain wall of the new fructification is made with the wall organization found in similar pollen of Dolerotheca. Relationships between the new taxon and other presumed seed fern pollen organs differing in preservation mode are discussed.     

TWO NEW MEMBERS OF THE BLASTOCLADIACEAE. I. TAXONOMY,WITH AN EVALUATION OF GENERA AND INTERRELATIONSHIPS IN THE FAMILY

Two new fungi in a small family of zoosporic phycomycetes, the Blastocladiaceae, are described. Both are included in the Brachyallomyces category, as sexuality has not been found in either one thus far. Microallomyces dendroideus from Costa Rica represents a new genus distinguished by its occurrence on animal or plant substrata rich in protein, its branched thallus in which pseudosepta are entirely lacking but growth is renewed by sympodially formed hyphal tips, and its small size. Allomyces reticulatus from California is the first species in this well-known genus to exhibit reticulately sculptured resistant sporangia. Its robust thallus is also unusual in that, despite the presence of the characteristic pseudosepta and branching with sympodially renewed hyphal tips, growth is ordinarily not unlimited as it is in all other species of the genus. The addition of these two new water molds to the Blastocladiaceae poses some basic questions regarding generic distinctions in the family. These problems are discussed and a working key to the genera is provided.     

The anatomy of Pseudosporochnus: P. Hueberi from the Devonian of New York

Eight structurally preserved specimens originally called Cladoxylon onteorense nom. nud. Hueber 1959, from Hensonville, New York, Cladoxylon dichotomum nom. nud. Hueber 1960 from East Ashland, N.Y., and Cladoxylon hueberi Matten 1974 from East Ashland and Cairo, N.Y., are analyzed in detail for their morphological and systematic significance. The type of C. hueberi from East Ashland, by far the most important specimen, provides the first opportunity in any Devonian member of the Cladoxylopsida for a realistic three-dimensional assessment of the primary xylem system. A main axis contains a highly dissected primary xylem system with 40–50 longitudinally anastomosing primary xylem segments. Distally, the main axis dichotomizes producing two equal branches with 20–25 xylem segments each. The outermost primary xylem consists of a system of radially oriented tracheid plates variously interconnected toward the inside, each with a conspicuous protoxylem strand near the xylem periphery. Traces to densely and three-dimensionally, but not helically, arranged lateral appendages depart singly or in groups of two or three from one to three adjacent peripheral-radial xylem plates. Although direct evidence on the external morphology of lateral appendages is not preserved, many trace groups exhibit conspicuous bilateral symmetry consistent with the interpretation of planate lateral appendages similar to those described for Pseudosporochnus nodosus. Other trace groups may indicate more three-dimensional lateral appendages. One sub-opposite pair of clepsydroid traces is borne at right angles to, and in obvious relationship with, the dichotomy of the main axis. Cortex is heterogeneous, consisting of thin-walled parenchyma with embedded clusters of sclereids and thick-walled parenchyma. These clusters appear responsible for a regular pattern of bumps or pits in compressed regions of the specimen and are closely comparable to those observed in Pseudosporochnus nodosus. A systematic analysis, attempting to view fragmentary evidence from both compressions and permineralizations in a single coherent framework, leads us to hypothesize assignment of the East Ashland and Hensoville specimens to the genus Pseudosporochnus. Although supported by the evidence, the hypothesis of assignment is by no means proven. We propose it, however, because we think that advancing systematic hypotheses conspicuously at risk of being refuted by additional observation promotes far greater understanding than taking an agnostic stance toward systematics implied by the use of “form” taxa. We further clarify our position by suggesting specific observations likely to be made from fossil specimens that might refute our hypothesis.