Pollen Organs Research Articles

Parasporotheca, Gen. Nov., and Its Bearing on the Interpretation of the Morphology of Permineralized Medullosan Pollen Organs

Parasporotheca is a large mass of synangiate microsporangiate organs attached to frond axes demonstrating medullosan anatomy. The specimen, of Late Pennsylvanian age, consists of several orders of frond axes, terminating within the fertile region. The frond axes share features with Sutcliffia- and Myeloxylon-type fronds but are not identical with either of these types. Several orders of lateral branching occur which terminate in dichotomized ultimate axes. These axes contain a single vascular bundle and serve as the stalks of terminally borne synangia. The fertile region is considered to be a type of compound synangium consisting of concentrically arranged synangia that are tightly appressed by interlocking epidermal projections. The individual synangia are ventrally concave laminate structures with a single row of 14-20 radiating, elongate, tubular sporangia that contain numerous prepollen grains of the Parasporites type. Features of Parasporotheca are compared with those of other permineralized medullos...

SULLITHECA DACTYLIFERA GEN. ET SP. N.: A NEW MEDULLOSAN POLLEN ORGAN AND ITS EVOLUTIONARY SIGNIFICANCE

The pteridosperm (Medullosaceae) pollen organ Sullitheca dactylifera gen. et sp. n. is described from middle Pennsylvanian coal balls. The proximally fused units of the obpyriform compound synangium separate and extend distally as finger‐like projections. Each projection contains 4–6 vertically oriented cylindrical sporangia arranged in pairs along the radius of the unit; each unit extends from the outer cover wall toward the center. The distal portion of the compound synangium is hollow as a result of the lateral separation of the centripetally and distally directed synangial units. About 40 tubular sporangia are present in all and dehiscence occurs along a lateral slit in each sporangium. Vascular strands are disposed around the periphery of the organ in addition to a single strand paralleling each sporangium. Two‐ or three‐cell trichomes and stomata are present on the organ surface. Pollen of the Monoletes type is present. A paired row of sporangia in Sullitheca composing a synangial unit is considered the homologue of a paired row of sporangia in the more compact and highly evolved genus, Dolerotheca.

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.

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.

PROBABLE ATTACHMENT OF THE DOLEROTHECA CAMPANULUM TO A MYELOXYLON‐ALETHOPTERIS TYPE FROND

The discovery of pedunculate specimens of pollen organ Dolerotheca in close association with Myeloxylon‐type pinnae and Alethopteris‐type pinnules provides evidence for the reconstruction of a petrified medullosan frond and the attachment of the microsporangiate fructification. Specimens of Dolerotheca villosa and D. formosa from Middle and Upper Pennsylvanian age strata of North America are borne on slender peduncles with anatomical features and vascularization identical to those of some level in a Myeloxylon‐Alethopteris‐type frond. Reconstruction of the frond reveals a large, repeatedly dichotomising organ that bears penultimate pinnae and ultimate pinnules in a pinnate arrangement. Examination of pinnules on the surface of coal ball material indicates that they conform to the compression genus Alethopteris. Identical numbers and arrangement of vascular bundles together with identical anatomical features and multicellular hairs indicate that the Dolerotheca campanulum is borne in the position of a penultimate pinna. A reconstruction of the frond with a proposed polled organ attachment is included.

Microgametophyte Development in the Paleozoic Seed Fern Family Callistophytaceae

Well-preserved stages of microgametophyte development are described from pollen produced by the Paleozoic seed fern family Callistophytaceae. Microgametophyte development in both the Middle Pennsylvanian pollen organ Idanothekion and Upper Pennsylvanian Callandrium involved the initial production of an axial row of at least three small prothallial cells proximally and a large embryonal cell distally. The arrangement and form of these cells is like that present in some extant genera of the Pinaceae. The prothallial cells were relatively large in comparison with extant gymnosperms, occupying the entire region of the cap-pus, and were apparently all primary. Evidence is presented that in Callandrium further development involved an anticlinal division of the large distal cell (antheridial initial) into a small generative cell contained within a larger tube cell. Previously described microgametophytes of the late Paleozoic order Cordaitales are reinterpreted and are shown to consist of an embryonal cell and three to four discoidal prothallial cells in an axial row like that of the Callistophytaceae. Microgametophytes thus far described from the Paleozoic are remarkably modern in appearance and provide no evidence to support the generally held view that the seed plant microgametophyte is an extremely reduced sexual phase that has arisen through the loss of almost all of the vegetative cells and the sterile outer cells of the antheridium. Evidence to support or refute this view will depend upon the discovery of microgametophytes from older groups of seed plants than those for which they are now known.

Probable Attachment of the Dolerotheca campanulum to a Myeloxylon-Alethopteris Type Frond

The discovery of pedunculate specimens of pollen organ Dolerotheca in close association with Myeloxylon-type pinnae and Alethopteris-type pinnules provides evidence for the reconstruction of a petrified medullosan frond and the attachment of the microsporangiate fructification. Specimens of Dolerotheca villosa and D. formosa from Middle and Upper Pennsylvanian age strata of North America are borne on slender peduncles with anatomical features and vascularization identical to those of some level in a Myeloxylon-Alethopteris-type frond. Reconstruction of the frond reveals a large, repeatedly dichotomising organ that bears penultimate pinnae and ultimate pinnules in a pinnate arrangement. Examination of pinnules on the surface of coal ball material indicates that they conform to the compression genus Alethopteris. Identical numbers and arrangement of vascular bundles together with identical anatomical features and multicellular hairs indicate that the Dolerotheca campanulum is borne in the position of a penultimate pinna. A reconstruction of the frond with a proposed polled organ attachment is included.

MICROGAMETOPHYTE DEVELOPMENT IN THE PALEOZOIC SEED FERN FAMILY CALLISTOPHYTACEAE

Well‐preserved stages of microgametophyte development are described from pollen produced by the Paleozoic seed fern family Callistophytaceae. Microgametophyte development in both the Middle Pennsylvanian pollen organ Idanothekion and Upper Pennsylvanian Callandrium involved the initial production of an axial row of at least three small prothallial cells proximally and a large embryonal cell distally. The arrangement and form of these cells is like that present in some extant genera of the Pinaceae. The prothallial cells were relatively large in comparison with extant gymnosperms, occupying the entire region of the cap‐pus, and were apparently all primary. Evidence is presented that in Callandrium further development involved an anticlinal division of the large distal cell (antheridial initial) into a small generative cell contained within a larger tube cell. Previously described microgametophytes of the late Paleozoic order Cordaitales are reinterpreted and are shown to consist of an embryonal cell and three to four discoidal prothallial cells in an axial row like that of the Callistophytaceae. Microgametophytes thus far described from the Paleozoic are remarkably modern in appearance and provide no evidence to support the generally held view that the seed plant microgametophyte is an extremely reduced sexual phase that has arisen through the loss of almost all of the vegetative cells and the sterile outer cells of the antheridium. Evidence to support or refute this view will depend upon the discovery of microgametophytes from older groups of seed plants than those for which they are now known.

POLLEN ORGANS OF THE PENNSYLVANIAN CALLISTOPHYTACEAE (PTERIDOSPERMOPSIDA)

Numerous specimens of the pteridospermous pollen organs Idanothekion glandulosum and Callandrium callistophytoides are present in Middle and Upper Pennsylvanian petrifaction material, respectively. Several specimens of Idanothekion, previously known only in an isolated condition, are attached to foliage assignable to the monostelic seed fern Callistophyton. Proof of the affinities of Idanothekion allows this genus to be elevated from a form genus to an organ genus in the Callistophytaceae. The structure of the foliage and the mode of attachment of the Idanothekion specimens are like those previously described for Callandrium. Structural features of Idanothekion and Callandrium are compared, and the question of their identity is discussed. At present one basic type of pollen organ, borne on the abaxial surface of foliar pinnules, is known for members of the Callistophytaceae.

Pollen Organs of the Pennsylvanian Callistophytaceae (Pteridospermopsida)

Numerous specimens of the pteridospermous pollen organs Idanothekion glandulosum and Callandrium callistophytoides are present in Middle and Upper Pennsylvanian petrifaction material, respectively. Several specimens of Idanothekion, previously known only in an isolated condition, are attached to foliage assignable to the monostelic seed fern Callistophyton. Proof of the affinities of Idanothekion allows this genus to be elevated from a form genus to an organ genus in the Callistophytaceae. The structure of the foliage and the mode of attachment of the Idanothekion specimens are like those previously described for Callandrium. Structural features of Idanothekion and Callandrium are compared, and the question of their identity is discussed. At present one basic type of pollen organ, borne on the abaxial surface of foliar pinnules, is known for members of the Callistophytaceae.

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.

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.

Telangiopsis Gen. Nov., an Upper Mississippian Pollen Organ from Arkansas

Telangiopsis arkansanum is described from compressed synangiate pollen organs borne terminally on a monopodially branched system of slender axes. The specimens occur in a fine-grained shale unit of the Wedington sandstone (Chester series). Individual synangia contain five or six sporangia, measure approximately 1.0 mm long and 0.8 mm wide, and contain radial trilete spores ranging from 47 to 54 μ in diameter. A discussion of the genus Telangium, to which similar remains would have previously been assigned, is presented. The generic name Telangium is retained for the petrified species T. scotti and T. pygmaeum, while the generic name Telangiopsis is proposed to include nonpetrifaction specimens previously assigned to the genus.

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.

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.

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.

Idanothekion Gen. N., a Synangiate Pollen Organ with Saccate Pollen from the Middle Pennsylvanian of Illinois

Idanothekion Gen. N., a Synangiate Pollen Organ with Saccate Pollen from the Middle Pennsylvanian of Illinois

A New Member of the Corystospermaceae Thomas

The leaf Pachypteris crassa (Halle) nov. comb, and its pollen organ, Pteruchus petasatus sp. nov., are described. They are placed in the Corystospermaceae (Pteridospermae), and the limits of the family are discussed with regard to the other 'Mesozoic pteridospermae' . Information is brought forward on the pollen morphology of the corystosperms.