Melissiotheca: a new pteridosperm pollen organ from the Lower Carboniferous of Scotland

Melissiotheca is based on fusainized pollen organs of pteridospermous affinities which occur abundantly in an Upper Visean limestone at Kingswood, near Pettycur (Scotland). The new species is a pedicellate synangium composed of 50–150 sporangia each of which is embedded at its base in a parenchymatous cushion divided into lobes. The sporangia are fused along their proximal half but are free distally. Dehiscence is longitudinal. Each sporangium is supplied at its base by a single vascular strand. Prepollen is small, spherical and trilete with a rugulate exine. In ultrastructure, the nexine appears homogeneous; the sexine shows internal sculpture of granae and rod elements. Melissiotheca has not been assigned to any family, but it shows many affinities with pollen organs attributed to the Lyginopteridaceae.     

The pollen cones of Ginkgo from the Early Cretaceous of China, and their bearing on the evolutionary significance

A well-preserved pollen cone of the genus Ginkgo was found in the Yixian Formation (Early Cretaceous) of Liaoning Province, China, and is described as the new species Ginkgo liaoningensis . The pollen cone, catkin-like, consists of a main axis bearing scales at its base and spirally arranged sporophylls bearing 3–4 (−2) oval or elliptical sporangia. The sporangia are pendulous and dehiscent by a longitudinal slit. The mature pollen grains are monocolpate and elongate-elliptical, and the juvenile pollen grains are found in the tetrad stage. The characteristics of G. liaoningensis are very similar to those of the living Ginkgo biloba , but the sporophylls of G. liaoningensis bear 3–4 (−2) sporangia, whereas those of G. biloba bear mostly two sporangia. G. liaoningensis differs from fossil cones of Ginkgo huttoni from the Jurassic of Yorkshire, UK, and an unnamed specimen of Ginkgo from the Upper Cretaceous of Alberta, Canada, in the size and number of sporangia. The comparison between G. liaoningensis (fossil) and G. biloba (living) supports the reduction hypothesis of ovule organs in Ginkgo , with the number of sporangia having experienced the process of reduction from three or four to two since the Early Cretaceous.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 133–144.     

Paleozoic seed fern pollen organs

Paleozoic pollen organs exhibit numerous morphological forms that have been arranged in categories based on their probable organization. Progymnosperm ancestors are characterized by three dimensional branching systems bearing pairs of terminal sporangia. Early Mississippian examples of seed fern fertile branches appear little modified from the progymnosperms. These pteridosperm microsporangia are nonsynangiate and thin walled with longitudinal dehiscence. By Upper Mississippian time all forms show sporangial clustering into large or small groups, with several taxa exhibiting radially symmetrical synangia. In the Lower Pennsylvanian all pollen organs are synangiate and appear to consist of a uniseriate ring of sporangia that either surround a central hollow, or are bilaterally flattened. Sporangial dehiscence in all forms is longitudinal and toward the center of the synangium. In bilateral synangia with no central hollow, the sporangia either separate laterally or effective dehiscence areas are restricted to the free apical portions of the sporangia. Callistophytacean synangia resemble the lyginopterid type, but are abaxial on laminar foliage. This family is thought to have evolved from the lyginopterids during the Early Pennsylvanian. Middle Pennsylvanian medullosan pollen organs are all radial and may be solitary, aggregated into groups, or fused into a large compound synangium. Several pollen organ types are reinterpreted, and the possible evolutionary relationships among the various Paleozoic pollen organ forms are discussed based on synangial organization, patterns of frond branching, and pollen or prepollen morphology.     

Pollen Organs and Seeds with Decussosporites Brenner from Lower Cretaceous Potomac Group Sediments of Eastern USA

Microsporangiate structures, Brenneria potomacensis gen. et sp. nov., containing pollen grains similar to dispersed Decussosporites are described from the Lower Cretaceous (Barremian or Early Aptian?) Potomac Group localities at Drewry's Bluff and Dutch Gap on the James River southeast of Richmond, Virginia. These fossils provide the first megafossil evidence of plants producing Decussosporites-type pollen and contribute important new information on the structure and possible systematic affinities of this unique Mesozoic gymnosperm. The microsporangiate structure is composed of an axis with helically arranged synangiate microsporangiate units, each unit consisting of two laterally fused sporangia borne on a short stalk. The pollen grains are very small, bisaccate, distinctly striate (taeniate) and TEM shows that they have partly infilled sacci (quasisaccate). These grains represent the youngest occurrence of saccate, striate pollen, which has not been recorded previously from sediments younger than the earliest Jurassic. Fossil seeds (Brennerispermum potomacensis gen. et sp. nov.) from the same localities as Brenneria contain Decussosporites pollen in the micropyle, and are believed to have been produced by the same plant species. The seeds are small and unitegmic with a distinct megaspore membrane. The occurrence of seeds adhering together in groups indicates that they were borne in aggregations. The microsporangiate structures of Brenneria show some similarity to those of the ginkgophytes (Ginkgoales and Czekanowskiales), but there are substantial differences in the structure of seeds and pollen. The Decussosporites pollen grains together with morphology of the reproductive organs support a closer relationship of the Brenneria-plant to previously described “Mesozoic pteridosperms”.     

Erdtmanitheca portucalensis,a new pollen organ from the Early Cretaceous (Aptian–Albian) of Portugal with Eucommiidites-type pollen

A new lignitised, slightly compressed pollen organ, Erdtmanitheca portucalensis, with affinities to extinct Erdtmanithecales from the Early Cretaceous (Aptian–Albian) of Vale de Água (Lusitanian Basin, western Portugal), is described. The pollen organ is composed of loosely arranged microsporophylls radiating from a central core. The estimated number of microsporophylls is about 100–150. The microsporophylls are sessile and ellipsoidal to barrel-shaped with a flattened or slightly apically depression containing about ten narrow sporangia. The sporangia enclose abundant well-preserved pollen grains of Eucommiidites-type. Pollen grains found in situ are elliptical in equatorial outline, about 16.0–27.2 μm long and 11.9–16.4 μm wide. The main (distal) colpus is long with expanded rounded ends. It is flanked by two subsidiary colpi in an almost equatorial position. The surface of the pollen wall is psilate and occasionally punctate. The ektexine is composed of a distinct tectum, granular infratectal layer and a thin foot layer. The endexine is thick and laminar. The new Early Cretaceous Portuguese pollen-organ is similar in several respects to that of Erdtmanitheca texensis described from the Late Cretaceous of Texas, USA. The new fossil species further documents the importance of the Bennettitales-Erdtmanithecales-Gnetales group in the Early Cretaceous floras of Portugal extending the stratigraphic and geographical distribution of the genus with regard to systematic and phylogenetic significance of the Eucommiidites-producing plants that may have been co-occurring with the Early Cretaceous diversification of angiosperms. It is ascertained that perforate tectum occurs in pollen grains with a well-developed foot layer as well as in pollen grains in which a foot layer is poorly developed or lacking, and that pollen features do not support a separation of the Erdtmanithecales seeds and pollen organs.     

CALLANDRIUM CALLISTOPHYTOIDES,GEN. ET SP. NOV., THE PROBABLE POLLEN-BEARING ORGAN OF THE SEED FERN,CALLISTOPHYTON

A new genus of pteridosperm pollen organ is described from Pennsylvanian age coal balls of Illinois. Individual sporangia are grouped into radial synangia which are borne in opposite pairs on the abaxial surface of slightly modified pinnules. Sporangia contain monosaccate pollen with a distinct sulcus, referable to the pollen genus Vesicaspora. Pinnules are borne on a regularly pinnate frond which is circinately coiled when young. At least a large portion of the frond is fertile and possesses anatomical features similar to those of Callistophyton. Phyletic relationships with other pteridosperm pollen organs are discussed.     

IDANOTHEKION GEN. N., A SYNANGIATE POLLEN ORGAN WITH SACCATE POLLEN FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Idanothekion glandulosum gen. et sp. n. is a synangiate pollen organ represented by approximately 30 specimens contained in coal balls from the middle Pennsylvanian of Illinois. Each synangium is composed of seven to nine elongate sporangia that are fused laterally for approximately four-fifths of their length, and are radially arranged about, and fused to, a short central column; the central column is restricted to the proximal one-third of the synangium. Distal to the column the sporangia surround a hollow central area. Dehiscence occurred by means of a longitudinal slit along the mid-line of the inner face of each sporangium. The outer walls of the sporangia have a complex histology involving an external epidermis, a middle presumably glandular layer containing scattered enlarged cells, and an inner layer made up of thin-walled parenchyma. Vascular tissue is present in the central column and outer walls of the sporangia. Each sporangium has a prominent, attenuate, multicellular tip. Large numbers of saccate pollen grains similar to those found in numerous fossil and extant coniferophytes as well as some Mesozoic pter-idosperms were produced in each sporangium. Idanolhekion resembles some synangia assignable to Paleozoic members of the Marattiales; however, the new genus compares most closely with pollen organs believed to have been produced by members of the Pteridospermales. It seems most likely that Idanothekion represents the pollen organ of some member of the Lyginopteridaceae that produced pollen of a type which up to now has not been known from Paleozoic seed ferns.     

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2–4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4–8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2–3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.     

The Early Carboniferous progymnosperm Protopitys: new data on vegetative and fertile structures,and on its geographic and stratigraphic distribution

We review progress made during the last 25 years in our understanding of the Protopityales, Early Carboniferous plants belonging to the extinct group of the progymnosperms. Recent studies support previous observations that the only genus of this order, Protopitys, included large arborescent plant with trunks up to 1 m in diameter. All branch orders had an oval eustele, secondary xylem with small rays and tracheid pitting ranging from circular bordered to scalariform bordered, and vascular traces to lateral appendages emitted in an alternate to subopposite distichous arrangement. Leaf morphology remains unknown. New material also confirms that fertile organs of Protopitys consist of branching systems bearing elongated sporangia terminally. Spores have a perispore and range in two size groups, which has been interpreted as a primitive stage of heterospory. The dense wood and fertile parts of Protopitys are comparable to those of the aneurophytalean progymnosperms of the Devonian, but Protopitys is distinct by its eustelic primary vascular system, and its affinities are still uncertain. The genus is now documented from at least nine localities in Europe, North America and Australia. Recent discoveries also indicate that it was present through the whole Mississippian, from the middle Tournaisian to the Serpukhovian.     

On Occurring of Sporangia of Protolepidodendron Krejci and Its Significance

This paper deals with the characteristics and occurring state of the sporangia of Protolepidodendron and its significance. The authors discovered that the sporangia with a short sporangiophore spread out from the axis of the fertile branch. There is a distance from the base of sporophyllary leaves to the setting point of the sporangiophore. Some sporangia divideinto suspended and symmetrical sporangia at the tip of the sporangiophore. But the most of sporangia do not divide. According to the characteristics of the sporangia of Protolepidodendron Krejci, which can be distinguished from genera Barrandeina (Krejci) Stur and Longos-tachys Zhu, Hu et Feng. Protolepidodendron lixianense sp. nov. is also described in this paper.     

VALLITHECA VALENTIA GEN. ET SP. NOV., PERMINERALIZED SYNANGIA FROM THE MIDDLE PENNSYLVANIAN OF OKLAHOMA

Vallitheca valentia gen. et sp. nov. is a permineralized, synangiate fructification of probable seed fern affinities from the Wewoka Formation (Desmoinesian) near Ada, Oklahoma. Synangia are oblong to pyriform, average 1 cm long, and contain 32 to 48 tubular sporangia embedded proximally in ground tissue. The sporangia are arranged within the synangia in a unique pattern: a ring of peripheral sporangia dehisces toward the center of the synangium and a central group of sporangia dehisces outward. This unique arrangement is unlike that of any currently recognized group.     

DEVELOPMENT AND CYTOLOGY OF THE ENDOBIOTIC PHASE OF PHYSODERMA PULPOSUM

Lingappa , Yamuna . (U. Michigan, Ann Arbor.) Development and cytology of the endobiotic phase of Physoderma pulposum. Amer. Jour. Bot. 46(4): 233–240. Illus. 1959.—The contents of the zygotes of Physoderma pulposum pass into the epidermal cells of the host and become incipient primary turbinate organs. The latter develop into resting sporangia in 3 different ways: (1) Occasionally, they may become thick walled as sporangia in their entirety. Such monocentric development of the endobiotic thalli is described for the first time in Physoderma; (2) usually, however, the contents of primary turbinate organs undergo centripetal cleavage; or (3) their contents may be cleaved tangentially. As a result, several uni- or multinucleate segments are formed which give rise to tenuous hyphae. The swollen distal end of each tenuous hypha develops into a secondary turbinate organ which in turn gives rise to hyphae and tertiary turbinate organs. Thus, the polycentric organization of the rhizomycelium is maintained. During this process, buds, which develop in the axils of the apical tufts of haustoria of turbinate organs, enlarge into resting sporangia. Nuclear divisions in turbinate organs and of resting sporangial initials are mitotic, and 4 chromosomes are evident on the equator of the intranuclear spindles. As the resting sporangia are inoperculate, the endosporangia protrude through irregular openings in the exospores.     

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.     

New male reproductive organs of gymnosperms Permotheca colovratica sp. nov. from the Lower Permian of the Ural Mountains

A new species of male gymnosperm reproductive organs, Permotheca colovratica sp. nov. from the upper part of the Kungurian Stage (Lower Permian, Irenian Horizon, Koshelevskian Formation) of the Middle Cis-Urals and from the Kazanian Stage of the southern Cis-Urals and Volga River Basin, is described. These organs are rosette-like synangiate aggregations of microsporangia consisting of four sporangia fused at their bases. The adaxial surface of the synangia has a disc of attachment. The in situ pollen grains are of Falcisporites type. The morphologically similar genera Falcisporites Leshik emend. Klaus, Alisporites Daugherty emend. Nilsson, and Vesicaspora Shemel emend. Wilson et Venkatachala are analyzed. The morphogenetic trend in the evolution of the androphores Permotheca is discussed.     

蕨类植物孢子囊形态 I. 鳞始蕨科

孢子囊是蕨类植物的繁殖器官, 其形态在蕨类植物的分类和系统发育研究上具有重要意义。用次氯酸钠溶液处理新鲜成熟的孢子囊, 在光学显微镜下获得清晰的孢子囊图像, 系统研究了中国鳞始蕨科4属13种孢子囊的形态特征。结果表明, 鳞始蕨科孢子囊呈椭球形, 孢子囊柄由3列细胞构成, 环带类型为垂直环带。通过分析孢子囊形态数据探讨了中国鳞始蕨科属内及属间差异。结果表明, 乌蕨属(Odontosoria)、香鳞始蕨属(Osmolindsaea)、达边蕨属(Tapeinidium)和鳞始蕨属(Lindsaea)的孢子囊环带细胞数依次减少, 囊蒴体积、唇细胞数和囊壁细胞数的变化由大(多)到小(少)依次为乌蕨属、达边蕨属、香鳞始蕨属和鳞始蕨属。孢子囊属内差异最大的是阔片乌蕨(Odontosoria biflora)与乌蕨(O. chinensis)以及香鳞始蕨(Osmolindsaea odorata)与日本鳞始蕨(O. japonica); 而达边蕨属和鳞始蕨属的属内差异则很小。研究结果为揭示鳞始蕨科系统发育关系提供了形态基础, 特别是提出阔片乌蕨和乌蕨以及香鳞始蕨和日本鳞始蕨在孢子囊形态上的差异值得进一步研究。     

Cladostrobus and the systematics of cordaitalean leaves

Cladostrobus Zalessky, 1918, is a male strobilus with spirally arranged microsporophylls, having a rhomboid lamina and a long stalk. The sporangia occur in groups. Microspores are of monosaccate type and are without a haptotypic mark. The name Cladaitina dibnerae gen. et sp. nov. is introduced for isolated spores of this kind. Cladostrobus may be a male strobilus of plants that bore Rufloria leaves. The group Cordaitales, as known now, comprises a heterogeneous assemblage of incompletely known plants, mostly sterile leaves. An artificial classification is proposed for sterile leaves, similar to the one already in vogue for dispersed spores and pollen. This classification is in fact a key for identification and cataloguing, and is not meant to show natural affinities.     

A FERTILE SPECIES OF TRIPHYLLOPTERIS FROM THE EARLY CARBONIFEROUS (MISSISSIPPIAN) OF SOUTHWESTERN VIRGINIA

The first occurrence of attached fertile structures to foliage assignable to the form genus Triphyllopteris Schimper is reported from the early Carboniferous Price Formation of southwestern Virginia. Sporangium-bearing branch systems consisting of a basal undivided foliar region and a distal much-divided fertile region are described as a new species T. uberis Skog and Gensel. Ovoid fusiform sporangia, 2.0 mm long and 0.5 mm wide and containing trilete spores, terminate ultimate divisions of the branch systems and collectively form masses up to 4 cm in all dimensions. T. uberis is compared to previously known sporangiate organs of Late Devonian and early Carboniferous age, many of which lack attached foliage. Similarities in the overall branching pattern of many of these non-synangiate, sporangiate organs of early Carboniferous age is noted. The possible mode of dispersal and affinities of the plant bearing T. uberis fertile foliage is discussed.     

Teixeiria lusitanica, a new fossil flower from the Early Cretaceous of Portugal with affinities to Ranunculales

A charcoalified fossil flower bud of a new genus and species (Teixeiria lusitanica) is described from the Early Cretaceous of Portugal. The flower is actinomorphic and unisexually male. At the base of the bud there are several bracts of different sizes, which are followed by sepal-like and petal-like tepals. Bracts and perianth organs seem to be arranged spirally and to exhibit transitions between different organ categories. The androecium has numerous stamens in two sizes, but with unclear arrangement. Pollen is small and tricolpate with a perforate tectum and a densely columellate infratectal layer. No carpels or remains of carpels could be observed on the floral axis. Teixeiria lusitanica shows most affinities to members of Ranunculales. There are also some similarities with Berberidopsis (Berberidopsidaceae, Berberidopsidales) and members of the Saxifragales (Hamamelidaceae and Daphniphyllaceae).     

The relationships of the genus Cicer L. (Leguminosae): the evidence from pollen morphology

The genus Cicer has traditionally been placed in the Vicieae, although it has a number of morphological characters which suggest that it is out of place in that tribe and that its affinities lie elsewhere. A survey of pollen morphology in Cicer , the other genera of the Vicieae, the Trifolieae and the Ononideae has been carried out to help determine the relationships of Cicer. The tribe Vicieae (without Cicer) is very homogeneous and is characterized by its long, rectangular pollen grains with small, heavily thickened endoapertures. The Trifolieae is more variable in its pollen morphology but, in general, has rather similar grains to the Vicieae although the structure of the endoaperture is different. The Ononideae and Cicer have nearly spheroidal pollen with very large, unthickened endoapertures. It therefore appears that as far as pollen characters are concerned, Cicer does not belong in the Vicieae but has much more in common with the Ononideae.     

西藏新生代秋乌组孢粉化石的发现及其初步研究

本文首次报道西藏雅鲁藏布江缝合带附近秋乌组含煤沉积中的孢粉化石。孢粉植物群主要由Quercoidifes,具孔类及三孔沟类花粉组成,包括12科26属及一些未定类群。组合中产有少量的Graminidites及Tsugaepollenites。植物群的总体特征显示其时代可能为渐新世至早中新世。秋乌组沉积早期的环境比较温暖。