— s^ C3. STATE OF ILLINOIS DWIGHT H. GREEN, Governor DEPARTMENT OF REGISTRATION AND EDUCATION FRANK G. THOMPSON. Director DIVISION OF THE STATE GEOLOGICAL SURVEY M. M. LEIGHTON, Chief URBANA REPORT OF INVESTIGATIONS — No. 91 AN ANNOTATED SYNOPSIS OF PALEOZOIC FOSSIL SPORES AND THE DEFINITION OF GENERIC GROUPS BY J. M. ScHOPF, L. R. Wilson, and Ray Bentall Prepared in collaboration with the Department of Geology, Coe College, and the Tennessee State Division of Geology PRINTED BY AUTHORITY OF THE STATE OF ILLINOIS URBANA, ILLINOIS 1944 M°Jf, STATE GEOLOGICAL SURVEY 3 3051 00005 7566 STATE OF ILLINOIS DWIGHT H. GREEN, Governor DEPARTMENT OF REGISTRATION AND EDUCATION FRANK G. THOMPSON. Director DIVISION OF THE STATE GEOLOGICAL SURVEY M. M. LEIGHTON, Chief URBANA REPORT OF INVESTIGATIONS — No. 91 AN ANNOTATED SYNOPSIS OF PALEOZOIC FOSSIL SPORES AND THE DEFINITION OF GENERIC GROUPS BY J. M. ScHOPF, L. R. Wilson, and Ray Bentall Prepared in collaboration with the Department of Geology, Coe College, and the Tennessee State Division of Geology PRINTED BY AUTHORITY OF THE STATE OF ILLINOIS URBANA, ILLINOIS 19 4 4 ORGANIZATION STATE OF ILLINOIS HON. DWIGHT H. GREEN, Governor DEPARTMENT OF REGISTRATION AND EDUCATION HON. FRANK G. THOMPSON, Director BOARD OF NATURAL RESOURCES AND CONSERVATION HON. FRANK G. THOMPSON, Chairman EDSON S. BASTIN, Ph.D., D.Sc, Geology ROGER ADAMS, Ph.D., D.Sc, Chemistry LOUIS R. HOWSON, C.E., Engineering WILLIAM TRELEASE, D.Sc, LL.D., Biology EZRA JACOB KRAUS, Ph.D., D.Sc, Forestry ARTHUR CUTTS WILLARD, D.Engr., LL.D. President of the University of Illinois GEOLOGICAL SURVEY DIVISION M. M.. LEIGHTON, Chief ^ ^ c^ SCIENTIFIC AND TECHNICAL STAFF OF THE STATE GEOLOGICAL SURVEY DIVISION 100 Natural Resources Building, Urbana M. M. LEIGHTON, Ph.D., Chief Enid Townley, M.S., Assistant to the Chief Velda a. Millard, Junior Assistant to the Chief GEOLOGICAL RESOURCES Coal G. H. Cady, Ph.D., Senior Geologist and Head L. C. McCabe, Ph.D., Geologist (on leave) R. J. Helfinstine, M.S., Assoc. Mech. Eng. James M. Schopf, Ph.D., Asst. Geologist J. Norman Payne, Ph.D., Asst. Geologist Charles C. Boley, M.S., Asst. Mining Eng. Bryan Parks, M.S., Asst. Geologist Robert M. Kosanke, M.A., Asst. Geologist George M. Wilson, B.S., Research Assistant Henry L. Smith, A.B., Research Assistant Industrial Minerals J. E. Lamar, B.S., Geologist and Head H. B. WiLLMAN, Ph.D., Assoc. Geologist Robert M. Grogan, Ph.D., Assoc. Geologist Robert R. Reynolds, M.S., Research Assistant Oil and Gas A. H. Bell, Ph.D., Geologist and Head Frederick Squires, B.S., ?etr. Eng. Charles W. Carter, Ph.D., Asst. Geologist William H. Easton, Ph.D., Asst. Geologist Paul G. Luckhardt, M.S., Asst. Geologist Wayne F. Meents. Research Assistant Areal and Engineering Geology George E. Ekblaw, Ph.D., Geologist and Head Richard F. 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Hursh, B.S., University of Illinois Mechanical Engineering, Seichi Konzo, M.S., University of Illinois Topographic Mapping in Cooperation with the United States Geological Survey. This Report is a Contribution of the Coal Division. May 30. 1943 Distributed July 1944 (46807) CONTENTS PAGE Introduction 7 Genera of fossil spores 11 Tasmanites 11 Triletes 19 Pityosporites 27 Punctati-sporites 29 Granulati-sporites ^ 32 Alati-sporites . . 33 Reticulati-sporites 34 Laevigato-sporites 36 Zonalo-sporites 37 Monoletes 38 Denso-sporites 39 Cystosporites 40 Parasporites 42 Cirratriradites 43 Endosporites 44 Triquitrites ,46 Equisetosporites 47 Alisporites 48 Calamospora 49 Reinschospora 52 Lycospora 54 Raistrickia 55 Florinites 56 Incertae sedis . 60 Summary 61 Acknowledgments . 61 References 62 Plates and plate explanations . . . , 67 [5] Digitized by tine Internet Arciiive in 2012 witii funding from University of Illinois Urbana-Champaign http://archive.org/details/annotatedsynopsi91scho AN ANNOTATED SYNOPSIS OF PALEOZOIC FOSSIL SPORES AND THE DEFINITION OF GENERIC GROUPS BY J. M. ScHOPF, L. R. Wilson, and Ray Bentall INTRODUCTION The literature on fossil spores is scat- tered, and in addition there has been much diversity in methods of classification. The objects of the present article are to bring this material together in summary form for convenient taxonomic reference and to evaluate the genera which have been pre- viously proposed. After a study of these microfossils in America and a thorough study of the literature it seems essential also to describe some new genera. It is hoped that a fundamentally sound basis has been provided for studies now in progress in the three laboratories repre- sented by the authors and for future studies. Guiding principles which relate to the treatment of this material are given below. 1. Only adherence to the systematic principles embodied in the International Rules will give satisfactory results in the study of these microfossils (as in other fields of paleontologic study). Therefore, the International Rules of Botanical Nomenclature, 3rd edition (Briquet), re- vised by the International Botanical Con- gress of Cambridge, 1930, and published in 1935,^ have been used as a basis for taxonomic treatment. 2. Species classified within the same genus or under the same generic name must possess significant characteristics in common. There can be little positive proof of generic identity unless substantial homologous comparisons can be drawn be- tween respective biocharacters of each congeneric species and the type species of the genus. For example many isolated lycopsid megaspores must be classified separately from the microspores found in ^ Gustav Fischer, Jena. association with them because there is no adequate basis for close comparison of biocharacters exhibited by these diversified organs. When affinities become better known so that the difficulty due to uncomparable features may be partially overcome, it then may be possible to relate the forms more naturally in classification within suprageneric groups such as tribe, family, and subfamily depending on how closely information may be correlated. Indeed, assignment to the same genus, or even species, is theoretically possible and may eventually be achieved in some few in- stances, but in general this implies a much more precise and detailed historical knowl- edge of interrelationships than is likely to be fully established. For satisfactory nomenclature it is most convenient that those fossils which possess numerous demonstrably comparable fea- tures shall be placed under the same ge- neric name. However, it* is unsatisfactory to classify fossils under the same generic name simply because a few arbitrary fea- tures are held in common. The essential validity of any classification depends on the relative significance that attaches to the various biocharacters. In fact it is by virtue of such interpretive discrimination that taxonomy is to be distinguished from a cataloging procedure. Interpretations nevertheless must be based on evidence and not on supposition ; difficulties are multi- plied whenever interpretation exceeds fac- tual bounds. 3. In paleobotanical practice there has been much divergence with regard to the significance and proper interpretation of unusually complete specimens in which parts usually found separated, and there- fore generally classified separately, are [7 8 PALEOZOIC FOSSIL SPORES found in organic union. Whatever attitude may be adopted, it is of fundamental im- portance for later consistent application of taxonomic principles. For this reason it seems desirable to state views applicable to the present work. The few unusually complete specimens are highly significant in showing beyond question that certain isolated organs at a particular locality and stratigraphic level possess identity of relationship.* This identity must become more dubious, in the case of isolated parts, the further they are removed geographically and strati- graphically from the site of proved union. Stratigraphic discrepancy is most likely to cast doubt on specific identity ; geographic discrepancy may be cause for qualification of the presumed relationship in varying degrees of subspecific magnitude (there is no basis for assuming that geographic races were less in evidence in the past than they are at present). If evolution is a more or less continuous attribute of life processes, over a period of time significant alterations in some of the many heritable characteristics are bound to occur. An in- dividual alteration, although of specific significance, will not necessarily involve phenotypic expression in the majority of the other characteristics of the race. For this reason it is inaccurate to postulate the continued specific coordination of an ex- tensive garniture of biocharacters over a period of time, without correspondingly detailed knowledge which proves that the characteristics dealt with were not subject to mutative or other evolutionary processes during that time interval. This detailed in- formation is most readily determined for a restricted group of biocharacters which can be observed on common specimens. Practical reasons dictate that emphasis in systematic treatment be placed on the more common types of fossils rather than on those which show unusual preservation. Demonstrated connections between spore forms and types of fructifications prove readily that some of the adjacent isolated specimens of either category are also conspecific. The diverse taxonomic groups, diagnosed on the basis of typical isolated specimens are proved, therefore, * No plant, and certainly no single specimen, can have more than one valid name. However, instances arise in which it is difficult to determine which of two or more names should actually be applied. In spite of this, syste- matic procedure demands that an author follow some consistent usage. to have coordinate significance through a part of their geologic life as species. Dur- ing this particular period, identity of re- lationship must exist. But the most perti- nent fact encountered in systematic treat- ment of the common isolated forms, is that this period of actual synonymy can scarce- ly ever be defined. As a general thing, due to discontinuity in the fossil record, it is impossible to establish the points at which old characters became coordinated in a new fashion or with other characteristics new to the phyletic succession. Consequently these overlapping rela- tionships do not lend themselves to taxo- nomic expression within specific and gen- eric nomenclatural categories. Groups of suprageneric rank, which are less depend- ent on type specimens for their proper definition, may be better used in expressing such relationships of generalized validity. The present paper, however, does not deal with groups of suprageneric taxonomic status. 4. Various criteria for determination of synonymy have been used by paleobot- anists. The practice we prefer is conserva- tive to the extent that unless conspecific relationship is proved for two nomencla- tural types, both names are valid. As such, they are available for any purpose of use- ful nomenclature. There are a large num- ber of "partial synonyms" among the named species of fossil plants to which these remarks apply. The word "synonym" of course is an absolute term — "partial synonyms" are not synonyms ; in most in- stances the names serve useful purposes in indicating groups of different circum- scription. None of these names should be lightly considered ; neither should a name be used in unqualified application unless its pertinence is evident. Evidences of "partial synonymy" undoubtedly record close natural relationship, but the names should not be regarded as invalid unless shown to be mutually inclusive. Lacking this, they remain applicable to fossils con- specific with their basic nomenclatural types. For the research worker there are advantages in this conservative view of synonymy, as it tends to promote a more precise differentiation of fossils. This basically, is a fundamental reason for the continued use of technical nomenclature, necessary to progress in the study of fos- sil plants. INTRODUCTION 5. Specific identification, because of its great significance, must be made critically. Emphasis should be placed on the positive evidence of similarity rather than on nega- tive evidence of lack, real or fancied, of features which cannot be examined criti- cally. 6. Neither spores, leaves, nor any other morphological part of a plant in any stage of its life is here considered to he a species. A species of plant, for the purposes of this synopsis, and regardless of how it may be otherwise defined, is regarded as a group of organisms. Any individual organism of such a group, how- ever, may be identified if any diagnostic structure representing any portion of its life cycle is available. Spores in many instances are probably just as definitive of species as any other organ belonging in a coordinated manner to the life cycle of a particular plant individual. They merit study with other types of ancient plant materials, and all data should be recorded on a common basis of taxonomic equality. Because of their unique adaptation for dispersal and their numerical abundance in many sedimentary deposits, it is expected that spores will eventually becon\e of greater practical significance than many of the larger types of plant fossils that are more commonly noted by the geologist. 7. Since the purpose of giving a name to a taxonomic group is not to indicate the characters or the history of the group, but is simply a convenience in reference, all names that are taxonomically valid and pertinent to recognized groups should be continued. The authors agree that the hyphenated generic nomenclature, for example, Granulati-s pontes, Laevigato- sporites, Denso-sporites, instituted first by Ibrahim (1933), is in poor form, lacks euphony, and tends to be generally mis- leading. Nevertheless, the status of such names seems reasonably secure in several instances and their continuance is likely to cause less misapprehension than attempts to institute more appropriate nomencla- ture. The preceding explanatory paragraphs may serve to clarify some of the policies adopted in this and subsequent reports and explain their consistency with sys- tematic treatment of other types of plant material. The earliest proposed name requiring consideration is Dawson's Sporangites. This name was first proposed in 1863 and has been used ambiguously and in different senses by many authors. It is here re- garded as a nomen ambiguum and the name Tasmanites is adopted for the most frequently encountered forms that have been commonly assigned to Sporangites. Tasmanites nevertheless is a problematic form, spore-like in many of its character- istics, but actually unassignable to any group of plants now known. All the other genera treated here belong with little doubt to the Cormophyta, and all whose affinity is approximately known belong to the Tra- cheophyte groups, as mentioned in the respective generic discussions. Knox (1939, 1941) has suggested that spores of Bryophytes may be present in Carboniferous coals, and some forms that have been described may well belong to this group of plants which at present are very scantly recognized in the Car- boniferous. The fact that no spores of Paleozoic Bryophytes are definitely known (Sporogonites Halle may be an exception) makes any suggested correlation hazard- ous at this time. It is important to realize that many specific features of spores of different geologic age can be matched because of evolutionary convergence as well as be- cause of community of derivation. Con- sequently unless there exists some cor- roborative evidence based upon spore forms actually present in contemporane- ous fossils whose relationship can be established, all that seems warranted is to direct attention to similarity of the spores of different geologic age, recognizing that the similarities may or may not have phy- letic implications. There is some lack of strict agreement between the morphological nomenclature applied to modern pollen and the morpho- logic features of fossil forms. For one thing the microspore in modern forms is determined, in a strict sense, solely by the presence of the unicellular male gameto- phyte.^ Such a distinction is of course inapplicable to fossil forms. There is also 2 The common indiscriminate reference to all spores of relatively small dimensions as "microspores" is to be lamented. Although true miscrospores frequently are small, by no means all small spores are microspores. The long established botanical usage of the term "microspore" has reference to fundamentally functional distinctions that are entirely aside from relative or specific size. R. B. Thompson (1927) has in fact demonstrated that in some plants the microspore is larger than the actual megaspore. 10 PALEOZOIC FOSSIL SPORES the further difficulty that no simple correct morphological designation can be applied similarly to the dispersal forms of the male gametophyte bodies in both modern and fossil forms. The dispersal stage rather than the degree of gametophytic development is of greatest practical signifi- cance when fossils are considered. The terms perispore, exospore, and endospore might seem equivalent to the pollen struc- tures perine, exine, and intine, respec- tively. However, in the case of the bladder- equipped fossil gymnospermic pollen, the bladder membrane which seems very similar in morphologic character to the perisporal membrane of certain crypto- gamic forms, is regarded by many as equivalent to the exine of angiospermic pollen. Revision of the morphologic termi- nology for fossil spores and similar micro- fossils also seems required. Not only has there been a dearth of descriptive terms available for description of the varied structural features, but a good many deep- ly rooted terms have become outmoded. Several of these date from the period in which Selaginella was presented as a primitive forerunner of modern seed plants, and their homologous implications now oftentimes seem unwarranted. Non- commital descriptive terms, which seem reasonably free ^of theoretical connota- tions, have been preferred for use in the present paper. It is hoped that it will be possible later to treat the descriptive morphology of spore forms specifically in greater detail. In the synoptic lists the species assigned to each genus are given in alphabetical order. The order of genera adopted is chronologic according to the time each was first proposed. Although some inter- generic alliances can be recognized now, and a more logical sequence should be adopted in later works, it is felt that the chronologic order of priority will be of some convenience for taxonomic refer- ence. Nearly all the references cited (p. 62) are those of particular taxonomic or mor- phologic pertinence for the fossils dis- cussed. Although we believe this repre- sents a reasonably complete survey of this literature, it does not include all articles that deal incidentally with Paleozoic spores. Various non-taxonomic systems have been used by some workers in desig- nating types of spores in coal thin- sections, notably by Slater and his col- leagues in the South Yorkshire laboratory of the British Fuel Research Division, by Th. Lang in Silesia and, most impor- tantly, by Reinhardt Thiessen. These apply very slightly to the present work because species cannot be as reliably dis- tinguished from spores in thin-sections, the significant biological characters are generally more difficult to ascertain, and the informal nomenclature used does not require systematic consideration. Con- sequently articles of this nature are not considered except in connection with the morphology of certain forms. On the other hand, there is a consider- able literature on isolated spores in the Russian language which needs to be given careful study before nomenclatorial stabil- ity can justifiably be hoped for. Thus far only a small portion of this literature has been available to us, chiefly in abstract form, and this is insufficient to provide a satisfactory basis for understanding the quite different nomenclature Russian writers have used. Naumova (1937) mentions that about 400 species have been distinguished based on material ranging in age from the Lower Carboniferous to Tertiary. He mentions genera designated as Zonotriletes and Azonotriletes. Nikitin (1934) has distinguished the genus Kry- shtofovichia based on very singular ap- pearing, large spores of Devonian age. Luber (1938) mentions subgroups (gen- era?) Azonotriletes, Zonotriletes, Azono- Jiionoletes, Azonaletes and Zonaletes and distinguishes several apparently valid species. In a more recent publication Luber (1939) uses the generic ( ?) names PlaguUtes, Turriella, Circelliella, Saccri- inalia, Circella, Libumella, Spinosella, Sub- sac culifer and Plicatella. It is to be hoped that under more favorable circumstances it will be possible to give full consideration to the Russian studies which have already been put to good use in the age determina- tion and correlation of the widely sep- arated coal deposits of the USSR and to integrate the nomenclature they are using with our own. TASMANITES GENERA OF FOSSIL SPORES 11 Genus Tasmanites Newton, 1875 Sporangites Dawson, 1863 (pars.), Canadian Naturalist, New Ser. vol. 8, no. 6, pp. 431-457. Plate 1, figure 1 General Discussion. — There is a group of highly distinctive fossil spore-like bodies which occur most notably in the Devonian-Mississippian black shale that requires a name amenable to standard taxonomic usages. At first, it was sup- posed that the name Sporangites, if sup- ported by an emended definition, could apply. We are now of the opinion that the application of that name is not correct in this connection, and that the various descriptions that have appeared for Spo- rangites are so highly conflicting and ambiguous or erroneous that that particu- lar name should be considered a nomen ambiguum. This is herewith proposed in view of the difficult taxonomic situation as detailed below. The present most common use of the term ''Sporangites'' is by geologists who wish to record the presence of more or less disc-shaped, resinous-appearing bodies that are very commonly found in the Devonian-Mississippian black shale, many, but by no means all, of which are properly referable to Tasmanites as given below. This common usage does not conform to the original application of the name and is so noncritical and general in application that it would probably be unwise to at- tempt to restrict it even if this could be done in accordance with taxonomic pro- cedure. Possibly no one, certainly not its author, has used the term with taxonomic consistency and practically all records of Sporangites must merely be regarded as indicative of some small rounded, brown- ish or yellowish bodies. Needless to say, many kinds of plant microfossils cor- respond to this characterization. How- ever, there is no reason why "sporangites" may not be continued to be used in this sense, since it serves a descriptive pur- pose, in spite of the fact that it is botanical- ly misleading; but the name should not be italicized or treated as a generic desig- nation. Sporangites was first proposed by Dawson in 1863 (p. 454). Here he applied the name to spores or "spore cases" which he thought were referable to Lepidodendron, Calamites and similar plants. Sporangites papillata, about one inch (!) in diameter, and 6". glabra about the size of a mustard seed, were men- tioned ; later this description was repeated, accompanied by small diagrams (Dawson, 1865, p. 165, pi. 12, figs. 80-81). These fossils were associated with coal at Jog- gins, Nova Scotia, and may in part have been material that would be placed now with Triletes, but there is no way to con- firm this from his description or illustra- tions. His repeated descriptions of 5^. papillata as "one inch in diameter" leads one to suspect it may have been a seed. In 1871 (a, b) Dawson described an additional species from the black shale at Kettle Point on Lake Huron as S. huron- ensis; "small globular papillate spore- cases — probably of some Lepidodendroid plant." Later in the same year (Dawson, 1871c) he also referred a specimen con- sisting of a small whorl of leaves or bracts, first described as Annularia acuminata, to Sporangites. This last interpretation of Sporangites was accepted by Slopes (1914). Forms similar, in general, to those from Kettle Point are usually con- strued as Sporangites by present day geologists, but it seems evident that neither of these interpretations corresponds at all closely with Dawson's first generic application of the name. Dawson in 1883 presented a report on "Rhizocarps in the Paleozoic Period" at the meeting of the American Association in Minneapolis, an abstract of which ap- peared in the American Association for the Advancement of Science proceedings (1884a). The entire paper also appeared in the Canadian Record of Science for the same year (1884b). The essential des- criptive part and the figures are identical in both. Here Dawson proposed the name Protosalvinia including two species given as ''Sporangites (Protosalvania) hrazilien- sis U.S.," and "S. (P.) hilohatus n.s." Dawson stated that if we compare the separate macrospores of the Brazilian sporocarps, and especially those which are found detached from their envelopes (Sic!) with Sporangites hitronensis, we 12 PALEOZOIC FOSSIL SPORES see a remarkable similarity in size, form and texture, sufficient to justify us in supposing that the latter may be of the same nature as the former, but deprived of their outer cases either by dehiscence or decay, and this is the view which we are now disposed to take of their nature. This better accords with their wide distribution in aqueous deposits and with their ac- companiment than any other supposition. Plausible and satisfactory as this sup- position may be, there is nevertheless scant basis for assuming all the various sporangite bodies to have such an origin, not to mention the insidiousness of at- tempting to apply scientific nomenclature on the basis of such a supposition. It can- not be said that Dawson ever proved the identity of bodies within and outside the "sporocarps" ; furthermore, no one else has ever provided any conclusive evidence. Evidently many have taken this important point for granted, however, because the two names are used practically inter- changeably in much of the subsequent literature. The "spherical and oval sacs, the walls of which are composed of a tissue of hexagonal cells . . . three to six mil- limeters in diameter," apparently are properly assignable to Prolosahiiiia Daw- son, 1884. We are unable to ascertain their characteristics reliably from any figures Dawson published but White and Stadnichenko (1923) give photographs and data on a form, Protosalvinia raveiina White, 1923, which is confirmatory of their thalloid characteristics, although White regarded them as sporangia. White (p. 239) has repeated the suggestion, (which seems to have originated with Dawson, 1871a, b) that their affinity was probably with "Devonian types antecedent to the Carboniferous Lepidodendra and Sigillariae." Possibly he thought the spor- angite bodies were actually spores of lycopod relationship, as many others have, and he probably accepted their sup- posed relationship with Protosalvinia be- cause of close association. Oval speci- mens of Protosalvinia bear some resem- blance to lycopod sporangia, but there are general objections to such an interpre- tation. Even when they are occasionally in abundance no cones or actual lyco- podiaceous remains are noted in the same assemblage (Dawson reported some du- bious lycopod stem material, but no thalli or "sporangia" with the "spores" at Kettle Point). White seems to have made no close study of the sporangite forms under appropriate magnification and he found none of them actually in any of the multi- tudinous "spore cases" (thalH?) of Pro- tosalvinia which were examined. In fact, although an uncritical reader may infer from this work that ''Sporangites' belongs to Protosaknnia, White actually made no direct statement to this effect and he pos- sibly had no strong convictions as to their nature. As the association of thalloid bodies with sporangite forms is by no means common and as the sporangites that have been reported differ considerably (one of the generic types here being treated as Tasmanites), it is evident that Protosalvinia and Sporangites cannot be in any way accurately treated as syn- onyms. No one yet has furnished tangible evidence of the nature of the spore-like bodies Dawson reported actually within the Protosalvinia thalli, so it still is quite speculative whether any relationship (other than that of a common environ- ment) is evidenced by their infrequent association in the Devonian-Mississippian black shale. Dawson reviewed the subject in con- nection with his paper in the bulletin of the Chicago Academy of Science in 1886 (pp. 105-118), calling attention to Orton's (1882) discovery of 6^. huronensis-Vik^ bodies in the Ohio shale, Clark's (1885) description of somewhat similar material in the Devonian of New York, the report of Johnson and Thomas (1884) on these microscopic discs in the Chicago water supply and glacial till of the region, New- ton's (1875) description of Tasnianites punctatus, and his own (Dawson, 1884a, b) descriptions of two lots of Brazilian material sent him by Hartt and by Orville Derby. He makes no direct mention either of the Joggins specimens or of ''Sporangites'' acuminata. This article is the best of any written by Dawson on the subject. Nevertheless, in it, as well as in his other papers, the inadequacies of his concepts are fully apparent. He appears confused as to the distinction between spores and sporangia ; he definitely con- siders the 6^. huronensis bodies referable to the water ferns (hence, Protosalvinia) y being misled on the one hand by the fal- TASMANITES 13 lacious idea of the primitiveness of that group of living plants, current at the time, and on the other by association with the (probably unrelated) thalloid plants sometimes found in the same beds. Daw- son considered his study of the Brazilian material particularly conclusive but his account of it does not seem so now and, unless the original material can be re- studied or new material obtained, the spe- cific nature of the Brazilian specimens is entirely problematic. Jongmans more recently (1930) has listed most of the nomenclature rather indiscriminately under the name Proto- salvinia. His treatment is hard to com- prehend inasmuch as Sporangites and Tasmanites are treated as synonyms although their precedent publication dates are given with approximate correctness." In attempting to follow Dawson's some- what confused usage and apply it more widely without original study, Jongmans has not contributed to the understanding of these plants or lessened the confusion in nomenclature. His omission of some pertinent references may in part be re- sponsible for the rather misleading im- pression conveyed by this portion of the Catalogus. One other thalloid plant occurs in asso- ciation with sporangite forms in the black shale which now can be more adequately discussed because of reexamination by modern methods. Spores are present in it and are quite different from sporangite bodies, in spite of Dawson's suggestion that they were similar. Dawson (1888) called this form Sporocarpon furcatum. In the most thorough and essentially definitive study of this plant by Kidston and Lang (1925), Dawson's name was continued. However, White, in White and Stadnichenko (1923), had renamed the group Foerstia but without taxonomic comment or discussion. Nevertheless the reason and justification for this new gen- eric name is evident upon consideration of the genus Sporocarpon Williamson (1879, cf. also 1880, p. 509) which was founded upon .S. cellulosum and similar forms of problematic affinity but quite f Dawson's paper in the Canadian Naturalist for 1870 evidently appeared subsequent to publication of his paper in the Am. Jour. Sci. for April, 1871. The paper "On Rhizocarps, etc." published in the Canadian Record of Science is dated and evidently was published in 1884, rather than 1885. Such minor errors are unfortunate in a reference so widely used as the Fossilium Catalogus. definite morphologic character. Many have considered the Sporocarpon types to be lycopod megaspores but from our pres- ent knowledge such a view seems un- tenable. Their structure is more rem- iniscent of colonial habit seen in the Volvocales. At any rate, there is no similarity between them and Sporocarpon furcatum Dawson. The name for this form must be Foerstia furcata (Dawson) Pia; Pia (1927) having been first to use this combination, but also without com- ment. The type species of Foerstia is best taken to be Foerstia ohioensis White, instead of Dawson's species. Kidston and Lang's work suggests that they may be specifically inseparable and if this is ever proved by comparison of the holo- types of the two species, White's specific designation, F. ohioensis, of course would become a synonym. The spores of Foerstia occur in tetrads, to some extent localized along one margin within the fertile tips. Possibly they were formed in individual conceptacles. An analogy in more than one particular is suggested by Fucus, but aside from the possibility that Foerstia may also be an advanced type of alga, probably no inter- pretation of homology can be supported. The individual spores of Foerstia are about 200 microns in diameter, similar to those of land plants, with a heavy "re- sistant" spore coat about 7 microns thick, definite haptotypic structures consisting of trilete sutures with pyramic areas well defined by distinct arcuate ridges. The distal part of the spore coat is irregu- larly pitted and somewhat rugose accord- ing to information presented by Kidston and Lang. Thus, these spores are en- tirely different from the sporangite bodies that may be associated with them. Haptotypic features have been de- scribed for the spores in all reliable reports that have ever been published of bonafide Devonian plants showing these reproductive structures. These spore forms, in addition to Tasmanites and its associates, may be found isolated in car- bonaceous Devonian sediments. Lang (1925) has shown that eight or more distinct species are thus represented in the Cromarty fish-beds of Old Red Sand- stone age in Scotland. None of these can be referred to Tasmanites, although Lang's spore-type F (in part), which 14 PALEOZOIC FOSSIL SPORES seems to lack haptotypic features, may be generically similar to some of the spore-like rugulose-membranous associ- ates of Tasinanites in American deposits. So far as these plant microfossils and others of similar character but question- able nature aref concerned, an adequate study of them is hardly begun. The essential point to be made in regard to ''Sporangites/' which geologists generally consider is well known, is that Dawson applied the name in the first place to ambiguous material and then later applied it without consistency to specimens of at least two other diverse and unrelated types. He further con- fused its application in conjunction with at least two other generic names. Geolo- gists have accepted one of the usages of Dawson, but as this does not correspond with Dawson's original interpretation it seems impossible to retain it in systematic usage. The term "sporangite" ought therefore to be considered no more than a descriptive designation without tax- onomic implication. The systemic designation Tasmanites was rejected by Dawson who recognized the similarity of Tasmanites punctatus Newton (1875) to his Sporangites hu- ronensis (Dawson 1886, p. 116) because, as the "name Sporangites had priority, I (i.e., Dawson) do not think it necessary to adopt this term (Tasmanites), although there can be little doubt that these organ- isms are of similar character." This similarity can be verified from Newton's illustrations by any one who has care- fully examined abundant sporangite speci- mens from the Devonian-Mississippian black shale, but the similarity is by no means evident if all the various inade- quate figures of "S. huronensis" given by Dawson and his contemporaries are consulted. The best evidence that at least many of Dawson's vS'. huronensis specimens, and particularly those expressly men- tioned by Orton, are similar to those of Newton has been presented in the cred- itably illustrated work by P. F. Reinsch, vol. II of the Micro-Paleophytologia,' published in 1884. Material from the Ohio shale and from Chicago sent by Orton and B. W. Thomas is described and illustrated and comparisons are read- ily made with similar material from Newton's tasmanite which Reinsch ob- tained from the British Museum. Reinsch assigned these forms, along with others, in his group called the Discieae, char- acterizing them generally as compressed disc-like plant bodies of original subspherical or subelliptical outline. They are entirely closed, with walls sometimes perforate and show a thin central cavity which has been reduced by compression. Variations in the walls are mentioned due to inclusion of other forms which seem to resemble Calcisphaera William- son (1880, p. 521), Sporocarpon Wil- liamson (1879, pp. 346-349; 1880, pp. 507-11), and many other diverse forms, some of which may even represent true sporangia of articulate plants. Reinsch also described 15 kinds of Discieae from the older Mesozoic. For the most part forms derived from the Carboniferous of central Russia and Saxony contrast sharply with the Tasmanites forms from America, Australia and Tasmania. These last evidently were chiefly in mind as he described the Discieae, for his group description fits them best. Specific no- menclature is applied to only one form so that Reinsch's work, although it shows best the characteristics of various Tas- manites specimens, is essentially not a taxonomic contribution. But since it establishes the similarity of American and Tasmanian forms we may feel no hes- itancy in calling these peculiar bodies Tasm^anites and in accepting Newton's species as the type for the genus. It seems so generally similar to some of the material provided by Orton and Thomas there is a question as to proper specific distinction between them, but being so widely separated geographically as they are, and stratigraphically as they seem to be, the forms should remain under separate names, at least until a definite first-hand study can be made. No spe- cific definitions are attempted here, but it is felt that the generic definition given below is sufficiently accurate to serve to distinguish the group from forms which unmistakably are spores of cryptogams and higher plants, and from several other kinds of problematic fossils which are found in Illinois (and probably else- where) associated with Tasmanites, but which must be considered generically distinct from it. TASMANITES 15 Diagnosis. — An adequate description of Tasmanites punctatus, taken here as the type species of the genus, is given by Newton and excerpts from his paper (pp. 339, 340, 341) are quoted below : When the separated discs are viewed by re- flected light, they appear as more or less circular bodies, somewhat thickened towards the circumference, many of them having their surfaces raised into irregular folds. If mounted in Canada balsam, and viewed by transmitted light, many have the appearance represented in pi. 10, figs. 2, 3, 8, while others exhibit the folds to which allusion has just been made. The more perfect discs are seen to be surrounded by a double contour-line — the optical expression of the fact that these discs are really thick- walled sacs. The saccular character, however, is best seen in transverse sections (figs. 1, 4, 5), or when the sac is broken (fig. 8). A closer examination enables one to see that the walls of these sacs are not homogeneous. A view such as fig. 8 shows numerous dots scat- tered over the surface, which become some- what elongated towards the edges of the disc. When examined with a power of about 250 diameters, the dots can be resolved into minute circles about 1/3000 of an inch in diameter with a still smaller dot in the centre, as shown in fig. 9. These structures are best seen in the discs of White Coal. It may be thought that these dots are comparable to the granules to be seen upon the surface of some of the macrospores of Flemingites ; but the study of transverse sections shows at once that these dots are not mere surface markings, for they can be distinctly traced as minute lines (tubes?) passing from the outer to the inner surface. These lines are shown in fig. 5, but owing to the section not being quite in the same plane as the lines, they do not appear to extend quite through. In addition to the fine lines, the walls of the sacs exhibit obscure longitudinal markings, which give them a laminated appearance (fig. J ) . Neither Mr. Carruthers (Geol. Mag. 1865, p. 432), nor Mr. MacNaughton (Trans. Roy. Soc. Van Dieman's Land, vol. ii, 1855, p. 116), mentions any structure in the walls of these sacs. The discs vary in diameter, as stated by both these authors, from about 1/80 to 1/50* of an inch. Mr. MacNaughton speaks of a thin outer coat to these discs, which may be seen when they are ruptured. I have examined all my preparations, both sections and sep- arated discs, in order to distinguish this outer coat, but have been unable to do so. One easily recognizes in transverse sections, such as fig. 1, that the walls of the sacs vary much as regards thickness ; and among the separated sacs which are mounted in balsam some may be seen much more transparent than the rest; but I have failed to see any real difference between the * 1/80 and 1/50 of an inch = 317.5 and 508 microns respectively. However, Newton's figure 2 "a large spore" given at 50x would be over 650 microns in diameter. From his figure 5 (250x) the wall is about 2 5 microns thick. In this specimen the wall is about 1/26 of the total diameter. ^ thicker and the thinner sacs, or to find them in anything like the relation of an inner and outer coat. Prof. Balfour, I believe, considers the Tasmanite discs to be closely allied to Flemingites; they differ from them, however, as Mr. Carruthers has pointed out (Geol. Mag. 1865), both in structure and size. All the Flemingites macrospores which I have seen have homogeneous walls, and in many of them is seen the triradiate marking, which is so gen- erally present in cryptogamic spores (Prof. Williamson, MacMillan's Mag. March, 1874, p. 409). In none of the Tasmanite sacs have I been able to see this triradiate marking, al- though their structures are so clearly shown that these markings could not fail to be seen if they were present ; and the walls, as we have already seen, have a definite structure. The inconvenience of having an object without a distinctive name induces me to propose one for the spores (?) found in Tasmanite and Australian White Coal (the two being, as I believe, identical in structure) ; and in order to retain existing titles as far as possible, I would suggest that Prof. Church's name Tasmanite, which is so generally used in reference to the schist as a whole, be retained for this substance, and that the spores (or rather the plant to which they belong) should be called Tasmanites, with the specific title of punctatus, in allusion to their surface-markings. The features which are here regarded as generically characteristic of Tasmanites are as follows : Symmetry. — Unicentric ; there is evi- dently a center and not an axis of sym- metry as in spores of bonafide plants. Shape. — Originally spherical ; except where protected, compression has altered them into disks with a few sporadic rounded folds. Si^e. — Ranging from less than 100 microns to 600 microns or slightly greater diameter. Forms greatly in excess or much smaller than these dimensions are suspect, because they vary so greatly from the genotype species. Ornamentation. — Surface smooth and glistening in reflected light at low magnifi- cation; more detailed examination shows more or less rugosity which may be in part attributable to preservation. More or less regularly spaced punctae varying in number on different forms are visible, but not conspicuous. The forms may be described as essentially lacking in external ornamentation. Haptotypic features. — Entirely absent. False conclusions have been drawn either from dififerent forms in association with Tasmanites or from specimens poorly 16 PALEOZOIC FOSSIL SPORES preserved and misinterpreted. Absence of trilete sutures is diagnostic. Wall. — Generally moderately thick, mostly 1/10 to 1/25 of the diameter; wall evenly developed on all surfaces and never membranous, often punctate with pores tapering from very small orifice on the outside; sometimes the punctae are very sparse, in other species they may be so densely packed as to give a radially striate appearance. Poorly defined con- centric bands may be present in the wall, but these are ordinarily not easily visible unless material is sectioned. In optical section (transmitted light) aside from the punctae, walls generally appear homo- geneous. Affinity. — The question of the affinity of Tasfimnites, so far as it can be def- initely discussed now, hinges on whether these bodies can be taken to represent plant spores or not. Spores of all plants above the algal stage that are readily preserved as fossils have markings on the spore coats of various but definite significance. All plant spores arise normally as tetrads (groups of four) from a single spore mother cell. All primitive plants (aside from algae) show evidences of their tetradic derivation in the structure of the spore coat. The Tasmanites forms do not show any such structure and, in addition, have funda- mentally a unicentric symmetry which is in strong contrast to plant spores which are always radially or bilaterally sym- metrical. Therefore, it seems conclusive that Tasmanites is not allied with any of the known primitive land plants. It may represent an algal type but if so, no cor- relative vegetative algal remains can yet be recognized. The ^ preceding remarks indicate that the writers' observations are by no means in agreement with Thiessen's (1921, 1925) on what, it appears, must include the same types of material. Thiessen persists in describing all forms from the black shale as if they had "tetrasporic" marks (in some cases more or less removed by abrasion). He definitely identifies them with Dawson's Sporangites huronensis and with the Tasmanites forms illustrated in 1884 by Johnson and Thomas (cf. Thiessen, 1921, p. 293) but discrepancies make it impossible to en- tirely confirm the statement. For ex- ample, the form shown on his plate 10, figure A (1921) is stated in the text to be 0.85 mm. (850 microns) in diameter but the legend says it is magnified 140 X, and measurement of the figure shows the form to be at that rate about 170 microns in diameter. It may be this form is the same as some observed in association with Tasmanites in Illinois, but should not be generically identified with it. The walls are generally quite rugulose-undulant and may show a poorly defined very coarse reticulation net, but the membranous char- acter of the fossil suggests that it orig- inally was smoother and that it has been impressed irregularly by grains of the enclosing sediments. No trilete suture (tetrasporic marking) can be demon- strated on material examined at first hand nor is any structure of this sort discernible in Thiessen's figures. Just as Newton remarked in his description of Tasmanites punctatus, in none of the sacs have I been able to see this triradiate marking, although their structures are so clearly shown that these markings could not fail to be seen if they were present. It must be recalled that the trilete and monolete (haptotypic) structures of cryp- togamic plants are a very definite spore feature, consisting not merely of a ridge which might be abraded from an other- wise featureless surface, but actually con- sisting of two margins, sometimes thick- ened, and a definite suture line between them which cannot be removed except by abrasion of the whole thickness of the proximal wall of the spore. Thiessen also fails to make any men- tion of the punctations which are so characteristic of the thicker walled forms referable to Tasmanites, and which are definitely but rather crudely shown in the illustrations of Johnson and Thomas. Had he appreciated what they were, it is doubtful that he would have been so positive that these fossils were spores of pteridophytes, because certainly no known spores of pteridophytes possess such structures. The most recent paper bearing on the subject concerns an examination of Tas- manian oil shale by Singh (1932). He considered that he was examining mate- rial similar to that studied by Newton in 1875 which contained Tasmanites punc- tatus. However, from the discrepancy in results it may be that he was dealing TASMANITES 17 with a different microfossil assemblage. Some of Singh's specimens without doubt are bonafide plant spores. However, he found many forms with about the same shape and size proportions as T. punctatus but was unable to see any of the punctae. Trilete markings were clearly visible in some but for others he suggests they ''seem to have been rubbed off during fossilization." This, of course, seems quite improbable since the trilete suture lines, if present, cannot be removed by fossilization. Of the figures presented by Singh, actually only one (fig. 3) shows convincing evidence of the trilete mark- ing. The proportions of this specimen suggest Calamospora and it is interesting to note that Phyllotheca (a calamarian) is associated in the same strata. Those shown in his figures 7 and 8 are incon- clusive as to the presence of a trilete structure, since Tasmanites may occasion- ally be folded under compression to simu- late a trilete marking ; but careful observa- tion will distinguish the absence of a definite suture line. Since Singh has not shown these structures, actually the only great discrepancy between Newton's and Singh's observations with respect to the more common forms concerns the punctae. American Tasmanites forms show con- siderable variation in the frequency of punctae and this may be a useful means of specific discrimination. One can scarcely doubt their existence in Newton's original material, because Reinsch also figured them from the Tasmanite mate- rial provided from the British Museum and also used and figured additional speci- mens from Newton's own preparations. So far as tlie discrepancy in presence of a trilete mark on some specimens is con- cerned, it must be noted that Reinsch (II, 1884, p. 5) also found "Rarissime oc- current Triletes minores singuli inter- spersi" in Newton's original material. Furthermore, the way Newton's descrip- tion in general tallies with the verified characters seen in American Tasmanites is so striking as to leave no doubt as to the essential correctness of his description. The surprising thing about American assemblages containing Tasmanites is that, so far as is known to be true, no trilete forms (excepting Foerstia) have yet been found to occur with them in the assem- blage. That Reinsch and Singh have found a few trilete spores associated with Tasmanites is not to be wondered at. The writers are convinced that, de- spite assocations of this sort, Tasmanites does not now, and never has had, any trilete marking on its wall. Lack of this feature is conclusive evidence that Tas- manites does not represent any group of ordinary pteridophytic plants. Discussion. — There are many spor- angite forms which do not qualify as species of Tasm^anites, or any other genus yet designated. Aside from Lang's paper there has been no attempt to report on isolated actual bonafide plant microfossils of similar age. Until more reports are available and greater knowledge has been obtained, it does not seem essential to give further taxonomic consideration to the few that are known. The correlation and coordination of taxonomic groups identified by means of spores from the Devonian, with those groups based on spores which we are utilizing for Carbo- niferous material, deserves very thorough study and consideration. The abundant plant record that can be obtained may, if advisedly interpreted, have much sig- nificance in tracing lines of phyletic rela- tionship among plants. The results that may possibly accrue from study of the problematic microfossils, among which is Tasmanites, is hard to predict, although such study cannot fail to at least outline their stratigraphic usefulness. It needs to be stressed, however, that Tasmanites, and other unicentric micro- fossils certainly do not belong to the well known groups of primitive higher plants that were contemporaneous with them. Their morphologic nature is, in fact, so uncertain that their reference to the plant kingdom would not seem assured except for their chemical composition. Zetzche, et al., (1931) seem to have definitely proved the absence of nitrogen from authentic Tasmanites and furthermore shown their composition to be in general similar to unquestioned fossil plant spores. Though possibly chitinous in appearance, lack of nitrogen apparently eliminates the animal kingdom from consideration. Spores of Parka which evidently is a thalloid plant, evidently lack any hapto- typic structures (Don and Hickling, 1915) but in no other particular do 18 PALEOZOIC FOSSIL SPORES they correspond to Tasmanites. In fact, these microscopic fossil bodies represent about as great a realm for speculation now as they ever did as to their actual nature and affinity, but this can hardly be represented as a valid excuse for their continued neglect. By strict application of systematic principles we shall approach much nearer a useful understanding of them. The following list includes names which have been used and will have to be con- sidered in subsequent treatment of species of Tasmanites; forms identified as Spor- angites jacksoni White (1905) and 5^. radiatus Duden (1897) have nothing to do with Tasmanites and are omitted on this account. 1. Tasmanites chicagoensis (Reinsch) S. W. and B., comb, nov.^ Discieae, Subtribus I, Subdividio 1., 18. ( addita est nomen Specificum "Chicagoense") Reinsch, 1884. Micro-Pale- ophytologia, vol. 2, p. 6, pi. 72A, fig. 76. Named from material supplied by Thomas, the specific name must be credited to Reinsch and the type is the specimen he figured. Daw- son (1888b) and Jongmans (1930) errone- ously cite Thomas as its author.*' 2. Tasmanites huronensis (Dawson) S. W. and B., comb, no v. Sporangites huronensis Dawson, 1871, Am. Jour. Sci., 3rd Sen, vol. 1, no. 4, p! 257, figs. 1-3. The following description is from the most complete account given by Dawson (1886) in which he corrected some of the earlier inac- curacies. It has also been changed slightly in accordance with the present conception of his material. Definite descriptive data is essen- tially unaltered. This is presented with the hope that it may be useful until Dawson's type material or authentic topotype material can be reexamined. Flattened disc- like bodies with smooth rounded edges usually ranging from 2 50 to 350 microns in diameter,' showmg slight external evidence of punctation but by transmitted light at higher magnification exhibiting nu- merous pores which penetrate the wall. Sometimes marked by characteristic rounded folds or split open by compres- sion. By transmitted light they appear amber colored or sometimes have a reddish hue; in section the walls show taint indications of concentric lamination. The walls are moderately thick 1/10 to 1/20 of the total diameter. They never exhibit the triradiate marking seen in spores ot lycopods. The interior is usually vacant but in some cases more or less filled with mineral matter (verv fine pyritic crystals, calcite, etc.). ^The initials S. W. and B. used here and elsewhere throughout this paper refer to the present authors, Schopf, Wilson, and Bentall. ,^ A copy of theMicro-Paleophytologia originally owned by Thomas now in the John Crerar Library in Chicago has the inscription ''Chicagoensii of B. W. Thomas" writ- ten on two of the figures, one of which is not Reinsch's ; lifJr^-ff ™.^^ u ^ ^^^. ^^'^^ interpreted his material a little differently than Reinsch did and may have mis- understood the taxonomic principles involved. The pri- mary purpose of author citation is to indicate type material upon which the description is based 3. Tasmanites punctatus Newton, 1875, Geol. Mag., ser. 2, vol. 2, no. 8, p. 341, pi. 10, figs. 2, 8. The type species of Tasmanites. (See de- scription as previously quoted from Newton.) 4. Tasmanites spp. Reinsch, nos. 12, (p. 3) ; 17, (pp. 3-4) ; 26, (p. 5) ; 32, (p. 6) and 65 ( ?), (p. 9) ; 1884, Micro-Pal eophytologia. Reinsch's diagnoses and illustrations sug- gest the numbers listed above, based on Amer- ican, Australian, and Tasmanian material, are probably referable to Tasmanites. Although others he assigned to the Discieae also lack haptotypic spore features, they are probably generically dissimilar. Note. — There is a possibility that the iso- lated bodies associated with Protosalvinia hra- siliensis Dawson and P. hilohata Dawson are referable to Tasmanites, and if so, they should receive separate taxonomic consideration and not be assigned to Protosalvinia. Proto- salvinia clarkei Dawson, which was originally identified with Tasmanites huronensis (Daw- son) by Clarke (1885) probably is entirely different and deserves restudy. One would judge that Tasmanites also occurs associated with this material. Probably many of the "Sporangites" occurrences given by Williams (1887) and various other authors are referable to Tasmanites but a critical reexamination of all of this material is needed. The forms designated as Sporangites alasken- sis and Sporangites arctica by David White (1929) were studied only from thin sections. The former shows wall peculiarities which may be comparable to that of Tasmanites ; the latter appears very similar to sectioned spores of higher plants. Whether the breaks in the walls are accidental (as they would be in Tasmanites) or are due to the section transecting a definitely organized suture of a spore coat is difficult if not impossible to determine from sections alone. This material was considered possibly Lower Cretaceous in age and a definite identification of Tasmanites would be of considerable interest. Genus Triletes (Reinsch, 1884) ; Schopf emend., 1938. Plate 1, figures 2, 2a, 3, 3a Megaspores radially symmetrical ; prox- imal side marked by triradiate sutures, often with arcuate ridges connecting the ends of the rays; distal surfaces smooth or variously ornamented. Ornamentation generally less well developed on proximal than on distal surfaces. Spores are rel- atively very large up to more than 3 mm. Affinity with the free-sporing lycopsids. Type species, by designation, Triletes reinschi (Ibrahim) Schopf (1938). This genus is a large and important one in classification of fossil spores, par- ticularly in the Carboniferous period when the free-sporing lycopsids were TRILETES 19 dominant. Evidence of the affinity of these spores is shown by every spore- containing- megasporangiate fructification of the free-sporing lycopsids of this pe- riod. Unfortunately, this corollary in- formation has not always been considered in fossil spore classification and additional genera have been proposed which are in conflict with Triletes. The types of these subsequently defined and consequently un- tenable genera are listed below. They are also included in alphabetical position in the main list of species with their synonymic citations. Triletes reinschi (Ibrahim) Schopf (1938), which also is the type species by designa- tion for Triletes. (Type of Laevigati- sporites Ibrahim 1933). Triletes tuberosus (Ibrahim) S. W. and B., comb. nov. (Type of Tiiberculati-sporites Ibrahim, 1933). Triletes sextus S. W. and B., nomen nov. (As Triletes VI of Bennie and Kidston (1886), this form was regarded as the type of Apiculati-sporites Ibrahim, 1933). Triletes hirsutus (Loose) S. W. and B., comb, nov. (Type of Setosi-sporites Ibrahim, 1933). Triletes sextusdecimus S. W. and B., nomen nov. (As Triletes XVI of Bennie and Kidston (1886), this form was regarded as the type of Zonales-sporites Ibrahim, 1933). Triletes trilobus (Ibrahim) S. W. and B., comb. nov. (Type of Valvisi-sporitcs Ibra- him, 1933). By no means all the species that authors regarded as congeneric with these types by virtue of arbitrary (artificial) distinctions appear so when broader comparisons are attempted. This is particularly true of the smaller forms (isospores and micro- spores) which often have been denomi- nated indiscriminately. These forms have of course been reallocated to what have the appearance, at least, of constituting more natural groups. The following is a list of described species of Triletes as the genus is now conceived. Some forms listed here may be so closely related to one another that in later studies the names will be con- sidered as synonyms. Without consulta- tion of the types or specimens correspond- ing to them these questions as to actual synonymy appear to be unsolvable. W^icher ( 1934) has placed several of these names in synonymous relationship, and in all probability correctly, since many of the poorly illustrated holotypes were available to him for examination. It may be like- wise a matter of individual opinion as to whether certain holotypes are conspe- cific. Such questions tend to be academic because there can be no doubt that the relationship is close in these instances. Also, whether the forms are distinguished as species or as varieties may be a matter of individual taxonomic policy. We have made as few changes of status as seemed reconcilable with a uniform taxonomic practice. Inconsistencies too may be found here, but it is hoped that these are within permissible limits of individual opinion. The synonomies are not absolutely com- plete in the sense that references to all forms that have ever been described or illustrated are included because triletean forms are represented in too many pub- lished illustrations for them all to be cited. The chief concern has been to give the references which include the type material or throw light upon it. Other references incidental to this purpose are given in a good many instances, chiefly to indicate the general features of occur- rence, both stratigraphic and geographic. Stratigraphic distribution has been gen- erally given for each. Such ranges are always subject to revision for a number of reasons. Many have been taken from the stratigraphic distribution tables and discussion pubhshed by Zerndt (1931, p. 169; 1937a, p. 68; 1937b, p. 590, 593; 1938; 1940, p. 142). The correlation chart given by Gothan (1937, p. 299) has been helpful in understanding the rel- ative position of the mostly unfamiliar named units of the Polish and Lower Sile- sian stratigraphic succession. Approximate equivalence of European and American Carboniferous divisions are given in the chart previously published by one of us (Schopf, 1941, p. 9). Zerndt's data on stratigraphic distribution is much more extensive than information from any other source and consequently deserves most serious consideration. Doubtless many forms not reported by him have a greater stratigraphic range than has been recognized. Zerndt's data, however, are given for his spore ''types" rather than for species. Most of his types and species have general equivalence yet this is not true in all cases and some discrepancy may occur on this account. In general, 20 PALEOZOIC FOSSIL SPORES probably a good many specific determi- nations have not been rendered critically enough and this also tends to diminish the value of stratigraphic ranges which can be given now. Nevertheless, some general information may be gained from the ages listed. It has been observed that a good many of the distinctive forms ap- pear in the same relative positions in America and in the European Carbonif- erous (Schopf, chiefly unpublished data) which gives promise of future assistance in long distance correlation. Ninety-five species are included here. Many of them are referable to the sections of Triletes already proposed (Schopf, 1938) ; others are less definitely assigned. Certain relationships between species have been noted when these seemed particu- larly evident. Triletes is by no means an exclusively Paleozoic genus and Meso- zoic forms have also been included for sake of completeness. Several new names have been proposed here, in addition to the considerable number of new combi- nations required in order to follow a consistent and relatively uniform tax- onomic policy. 1. Triletes agnina (Zerndt) S. W. and B., comb. nov. Lagenicula agnina Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, p. 14, pis. 21-22. Type 34 Zerndt, 1937, idem. Sectio, Lagenicula. Age, Lower Na- murian (Flora series). 2. Triletes ales Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 163, fig. 53A-E, pi. 25, figs. 2, 8, 9, 11. Age, Liasso-Rhaetic. 3. Triletes angulata (Zerndt) S. W. and B., comb. nov. Lagenicula angulata Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, pp. 11-12, fig. 8, pis. 14, 15. Type 7 Zerndt, 1931, Acad, polonaise sci. Bull, internal., ser. A, p. 171, pi. 3, fig. 8. Sectio, Lagenicula. Age, Lower Na- murian (Grodziec series). 4. Triletes areolatus Harris, 1935, Med- delelser om Gronland, vol. 112, no. 1, pp. 158-159, fig. 51A-F, pi. 26, figs. 3, 10. Age, Liasso-Rhaetic. 5. Triletes ariadnae (Miner) Harris, 1935, idem. p. 155. Selaginellites ariadnae Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 505, figs. 26-30. Age, Upper Cretaceous. 6. Triletes arnoldi (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, p. 155. Selaginellites arnoldi Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 500, figs. 22-25. Age, Upper Cretaceous. 7. Triletes augustae (Loose) S. W. and B., comb. nov. Zonales-sporites augustae Loose, 1934, Inst. Palaobot. Arb. vol. 4, p. 150, pi. 7, fig. 32. Sectio, Auriculatif Age, Westphalian B (Gasflamm). 8. Triletes aurantium Harris, 1935, Med- delelser om Gronland, vol. 112, No. 1, pp. 164-165, fig. 52H-L, pi. 25, fig. 5, pi. 26, fig. 5. Age, Liasso-Rhaetic. 9. Triletes apiculatus (Ibrahim) S. W. and B., comb. nov. Apiculati-sporites apiculatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 23, pi. 5, fig. 31. Type 14 Zerndt (in part), 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 172; 1935, Acad, polonaise sci. Trav. (jeol. no. 1, pp. 17-18. Sectio, Aphanosonati. Age, Lower Na- murian, Westphalian B, and younger (?). 10. Triletes artecollatus Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, internal., ser. A, p. 58, pi. 1, figs. 4, 5. Type 39, Zerndt, 1936, idem. Sectio, Triangulati. Age, Dinantian (Raczna Series). 11. Triletes auritus Zerndt, 1930, idem, ser. B, p. 46, pi. 1, figs. 4 and 5. Spore 0.7 mm. Zerndt, 1930, Soc. geol. Pologne Ann., vol. 6, pp. 307-303, 312, pi. 1, figs. 3a, b, pi. 3, figs. 3a-d. Types 11 and 12 Zerndt, 1931, Acad, polonaise sci. Bull, internal., ser. A, p. 172. Triletes auritus Zerndt, 1937, idem, p. 584, pi. 10, figs. 2, 6. Triletes auritus Zerndt, 1940, Paleonlo- graphica, vol. 84, abl. B, p. 146, pi. 9, figs. 5-18. Sectio, auriculati (type). Age, West- phalian C, D. Triletes auritus var. grandis Zerndt, 1937, Acad, polonaise Bull, internat., ser. A, p. 584, pi. 10, figs. 1, 3-5. Type 11a, Zerndt, 1937 (see above). Triletes auritus var. grandis Zerndt, 1940, Paleontographica, vol. 84, abt. B, p. 134, pi. 11, figs. 32-35. Age, Stephanian (Westphalian D?). Triletes auritus var. secundus (Maslan- kiewiczowa) S. W. and B., nom. nov. TRILETES 21 Triletes auritus II Maslankiewiczowa, 1932, Acta Soc. Bot. Polonaise, vol. 9, (SuppL), p. 158-161, figs. 33-36. 12. Triletes appendiculatus Maslankiewic- zowa, 1932, idem., p. 163-164, fig. 39. Type 12 Zerndt, 1937 (non 1931), Acad, polonaise sci. Trav. Geol. no. 3, p. 3. Type 6 Sahabi, 1936, Recherches sur les spores des houilles Francaises, p. 39- 40, pi. 2, fig. 5. Type 8? Sahabi, 1936, Recherches sur les spores des houilles Francaises, p. 41, pi. 2, figs. 7-8. Sectio, Auriculati. Age, Westphalian C, D, Stephanian. 13. Triletes bennholdi Zerndt (non Bode), 1937, Acad, polonaise sci. Trav. Geol. no. 3, p. 5-6, fig. 3, pi. 2, figs. 1-6. Type Zi Zerndt, 1937 (idem). Porosporites bennholdi (Bode) Zerndt, 1938, Deuxieme Cong. Stratig. Carbonif., vol. III. p. 1713. Age, Lower Namurian (Flora series). Note. — Zerndt credited this specific epithet to Bode, apparently assuming the spores con- specific with those of Porostrobus bennholdi. If Porostrobus can be identified reliably from its isolated spores alone, then evidently Porostrobus is a later homonym of Triletes. Such is hardly the case, however, because the identification of the genus Porostrobus and the species bennholdi within it depend on other features in addition to the spores. Some day we possibly may know actually how diagnostic isolated spores of Porostrobus may be, but at present such speci- mens certainly must be recorded separately under a different genus (as Zerndt has done). Consequently, also, these forms must be identi- fied with reference to different nomenclatural types. The specimen illustrated by Zerndt's photograph 4, plate 2, and which evidently served as the basis for his text figure 3 (p. 6), is regarded as the holotype of the species which is here credited to Zerndt. On the other hand, no question arises as to the spores Wicher (1934b) described in some detail that were isolated from undisputed speci- mens of Porostrobus. There was no occasion for him to assign them a new generic name (Poro(strobo)sporites) since the only possible designation for the specimens is "spores of Porostrobus bennholdi." Zerndt, in using the name "Porosporites" in 1938, is presumably following Wicher. Wherever evidence of addi- tional cone characters warrants it, such a deter- mination is more significant and essentially useful than identification with a less closely defined group such as Triletes. Mistakes of this kind are less likely to occur if nomenclatural types are regarded seriously. 14. Triletes bipilosus (Wicher) S. W. and B. comb. nov. Sporites bipilosus Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 180, pi. 8, figs. 13, 14. Sectio, Lagenicula. Age, Lower West- phalian C. Note. — This species is apparently closely re- lated to Triletes robertianus and T. latihirsutus. 15. Triletes borealis (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155. Selaginellites borealis Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 503, figs. 12-21. 16. Triletes brasserti Stach and Zerndt, 1931, Gliickauf, Jahrg. 67, no. 35, p. 1123, figs. 29, 30, (?) 3L Type 20 {Triletes brasserti Stach and Zerndt), Zerndt, 1934, Acad, polonaise sci. Trav. Geol., No. 1, pp. 23-24, fig. 9, pi. 25, figs. 1-2. Age, Dinantian, Namurian Westphalian A, B. Note. — Closely related to Triletes circumtex- tus, T. saturnoides and T. superbus. 17. Triletes breviaculeatus Nowak and Zerndt, 1937, Zerndt, idem., no. 3, pi. 4, figs. 1, 2. Type 38, Triletes breviaculeatus Nowak and Zerndt, 1936, Acad, polonaise Bull, internat., ser. A, p. 57. Sectio, Aphanozonati. Age, Upper Din- antian (Raczna series). Lower Namurian (Flora series). Note. — Evidently by mistake an illustration of Triletes horridus accompanied Nowak and Zerndt's original description. The species was validated by photographs published a year later. 18. Triletes brevispiculus Schopf, 1938, Illi- nois Geol. Survey Rept. Inv. 50, pp. 26-27, pi. 1, figs. 13a-r; pi. 2, fig. 6; pi. 3, figs. 1-4. Sectio, Aphanozonati. Age, Middle Pennsylvanian, Carbondale series. 19. Triletes circumtextus Zerndt, 1934, Acad. polonaise sci. Trav. Geol. no. 1, pp. 19-21, fig. 7; pi. 19, figs. 1-12; pi. 21, figs. 1-7. Type 18 Zerndt, 1931, Acad, polonaise Bull, internat., ser. A, p. 173, pi. 6, figs. 17, 18. Age, Lower Namurian. Note. — Closely related to Triletes brasserti. 20. Triletes clavatopilosus (Wicher) S. W. and B., comb. nov. Sporites clavatopilosus Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 183, pi. 8, figs. 25-26. Type 24 Zerndt, 1931, Acad, polonaise sci. Bull, internat,, ser. B, pp. 174-175, pi. 7, fig. 23. Type 14 Sahabi (?), 1936, Recherches sur les spores des houilles Francaises, pp. 47 and 48, pi. 6, figs. 5-7. Age, Upper West- phalian A — Lower Westphalian C. 21. Triletes crassiaculeatus (Zerndt) S. W. and B., comb. nov. Lagenicula crassiaculeata Zerndt, 1937, 22 PALEOZOIC FOSSIL SPORES Acad, polonaise sci. Trav. Geol. no. 3, pp. 12-13. Type 26A, Zerndt, 1937 (idem). Sectio, Lagenicula. Age, Upper Dinan- tian (?). A^o^^.— Related to Trilctcs horridus. 22. Triletes cyttarta Kendall, 1942, Annals and Mag. Nat. History ser. 11, vol. 9, p. 922, fig. 2. Age, Jurassic (Mid. Estuarine). 23. Triletes dificilis (Wicher) S. W. and B., comb. nov. Sporites dificilis Wicher, 1934, Inst Palaobot. Arb., vol. 4, no. 4, p. 179, pi. 8 figs. 17-18. Type 14 (in part) Zerndt, 1931 Acad polonaise sci. Bull, internat., ser. A, p. 172 pi. 4, figs. 11, 12, pi. 5, figs. 15, 16. Zerndt, 1934, Acad, polonaise sci. Trav, Geol. no. 1, pp. 17-18, pi. 8, figs. 1-10, pi 31, figs. 8, 9. Sectio, Aphanozonati. Age, Lower Westphalian C. (Zerndt, Mid. Lower. Namurian to Westphalian D). 24. Triletes difusopilosus (Wicher) S. W. and B., comb. nov. Sporites difusopilosus Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, pp. 182- 183, pi. 8, figs. 23, 24. Age, Lower West- phalian C. Note. — Closely related to Triletes praetextus. 25. Triletes echinatus (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155. Selaginellites echinatus Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 500, fig. 6. Age, Upper Cretaceous. 26. Triletes erlansoni (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155. Selaginellites erlansoni Miner, 1932, Washington Acad. Sci. Jour,, vol. 22, p. 500, figs. 1-3. Age, Upper Cretaceous. 27. Triletes flavus Stach and Zerndt, 1931, Gliickauf, Jahrg. 67, no. 35, p. 1122, fig. 18. Sectio, Auriculati. Age, Westphalian C. 28. Triletes fulgens Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, p. 5, fig. 2, pi. 1, figs. 1-9. Type 8 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 171, pi. 3, figs. 9-10. Sectio, Aphanosonati. Age, Middle Din- antian to Middle Namurian. ATo^^.— Closely related to Triletes glabratus (pars.). 29. Triletes glabratus Zerndt, 1930, idem, ser. B, p. 45, pi. 1, figs. 2, 3. Type 9 Zerndt, 1931, idem., ser. A, p. Sectio, Aphanozonati. Age, Westphalian B, C, Lower Stephanian. Note. — Closely related to Triletes fulgens and Triletes reinschi. 30. Triletes greenlandicus (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155. Selaginellites greenlandicus Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 500, figs. 10, 11. Age, Lower Cretaceous. 31. Triletes gymnozonatus Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, pp. 37- 38, pi. 1, fig. 4, pi. 4, fig. 9. Sectio, Triangulati. Age, Middle Penn- sylvanian, Carbondale series. Note. — Closely related to Triletes artecol- latus. 32. Triletes harrisi Murray, 1939, Geol. Mag., vol. 76, p. 480, figs. 1, 2. 33. Triletes hirsutus (Loose) S. W. and B., comb. nov. Sporonites hirsutus, Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 58. Setosi-sporites hirsutus (Loose) Ibra- him, 1933, Sporenformen des Aegirhori- zonts, p. 26. Apiculati - sporites hirsutus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 153. Sporites hirsutus (Loose) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no, 4, p. 181. Sectio, Lagenicula?. Age, Westphalian B, Lower Westphalian C. 34. Triletes horridus (Zerndt) S. W. and B., comb. nov. Lagenicula horrida Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no, 1, p. 25, fig. 11, pi. 28, figs. 1-5. Type 26 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 175, pi, 9. fig. 28. Sectio, Lagenicula. Age, Dinantian. Note. — Closely related to Triletes crassiacu- l eat us. 35. Triletes inornatus (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155. Selaginellites inornatus Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 505, figs. 7, 8. Age, Upper Cretaceous. 36. Triletes ionthus Harris, 1935, Meddelel- ser om Gronland, vol. 112, no. 1, pp. 166-167, 169, figs. 52E, F, G, pi. 26, fig. 8. 37. Triletes kidstoni (Loose) S. W. and B., comb. nov. Sporonites kidstoni Loose, 1932, Neues Jahrb, Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 60. TRILETES 23 Laevigati-sporites kidstoni (Loose) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 19. Reticulati-sporitcs kidstoni (Loose) Loose, 1934, Inst. Palaobot, Arb., vol. 4, no. 3, pp. 156-7. Sectio, Aphanosonati. Age, Upper Westphalian B, Lower Westphalian C. 38. Triletes latihirsutus (Loose) S. W. and B., comb. nov. Apiculati-sporites latihirsutus Loose, 1934, idem., p. 153. Sectio, Lagenicula. Age, Upper West- phalian B. Note. — Closely related to Triletes robertianus. 39. Triletes laximarginalis Zerndt, 1940, Paleontographica, vol. 84, Abt. B, p. 136, pi. 10, figs. 24-28. Type 17a, Zerndt, 1940 (idem). Sectio, Triangulati. Age, Westphalian C, D (?). 40. Triletes levis (Zerndt) S. W. and B.. comb. nov. Lagenicula levis Zerndt, 1937, Acad, polonaise sci. Bull, internat., ser. A, pp. 587-588, pi. 15, figs. 1-11. Type 45 Zerndt, 1937 (idem). Sectio, Lagenicula. Age, Stephanian. Note. — Closely related to Triletes translucens, T. tenuimemhranosa and to T. rugosus. 41. Triletes litchi Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, pp. 159, 160, fig. A-F, pi. 26, figs. 3, 10. Age, Liasso-Rhaetic. 42. Triletes mamillarius Bartlett, 1928, Michigan Acad. Sci., vol. 9, p. 21, pis. 13-15. Sectio, Aphanozonati. Age, possibly Upper Pottsville or Lower ATleglieny. 43. Triletes mensura Harris, 1935, Meddelel- ser om Gronland, vol. 112, no. 1, pp. 160- 162, fig. 53J, pi. 26, figs. 2, 9. Age, Liasso-Rhaetic. 44. Triletes mirabilis (Miner) S. W. and B., comb. nov. Selaginellites mirabilis Miner, 1935, Am. Midland Naturalist, vol. 16, p. 618, pi. 23, figs. 1-6. Age, Upper Cretaceous?. 45. Triletes mucronatus Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, internat., ser. A, p. 59, pi. 1, figs. 1, 2. Type 40 Zerndt, 1936 (idem). Age, Dinantian. Note. — Possibly closely related to Triletes clavatopilosus. 46. Triletes myrmecoides Harris, 1935, Med- delelser om Gronland, vol. 112, no. 1, fig. 52M-Q, pi. 26, fig. 4. Age, Liasso-Rhaetic. 47. Triletes nigrozonalis Stach and Zerndt, 1931, Gliickauf, Jahrg. 67, no. 35, p. 1123, figs. 26, 27. Sporites nigrozonalis (Stach and Zerndt) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, pp. 176-177, pi. 8, figs. 11 and 12. Sectio, Auriculati. Age, Westphalian C. 48. Triletes nodosus (Wicher) S. W. and B., comb. nov. Sporites nodosus Wicher, 1934, idem., p. 177, pi. 8, fig. 21. Sectio, Aphanozonati. Age, Lower Westphalian C. 49. Triletes nudus (Nowak and Zerndt) Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 30, pi. 5, fig. 7. Lagenicula nuda Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, internat., ser. A, p. 60, pi. 1, fig. 6. Type 43 Zerndt, 1936 (idem). Sectio, Lagenicula. Age, Dinantian. Note. — The form recorded from lUinois as T. (?) nudus probably is not referable to this species although the shape is rather similar. 50. Triletes ovalis Stach and Zerndt, 1931, Gliickauf Jahrg. 67, no. 35, p. 1122, fig. 24. Age, Westphalian B, Lower Westpha- lian C. Note. — Probably closely related to type 5 of Sahabi (1936)., 51. Triletes papillosus (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, No. 1, p. 155. Selaginellites papillosus Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 500, fig. 9. Age, Upper Cretaceous. 52. Triletes parviapiculatus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 2, p. 17, pi. 24, figs. 1-4. Sectio, Triangulatif. Age, Lower Mar- ginal beds (Lower Namurian, vicinity of Rybnik). 53. Triletes pedinacron Harris, 1935, Med- delelser om Gronland, vol. 112, no. 1, pp. 165-166, fig. 52S, T, pi. 27, fig. 6. Age, Liasso-Rhaetic. 54. Triletes persimilis Harris, 1935, idem., p. 165, fig. 52R, pi. 25, fig. 4. 55. Triletes pertuberosus (Loose) S. W. and B., comb. nov. Sporonites pertuberosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 57. Tuberculati-sporites pertuberosus, (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 147, pi. 20, fig. 57. Sectio, Aphanozonati. Age, Upper Westphalian B. ' , ' ' 24 PALEOZOIC FOSSIL SPORES Note.— Equal to Type 14 Zerndt (1931, 1934) in part. 56. Triletes phyllicus Murray, 1939, Geol. Mag., vol. 76, p. 482, figs. 7, 8. Age, Jurassic (Estuarine). 57. Triletes pinguis Harris, 1935, Meddelel- ser om Gronland, vol. 112, no. 1, p. 166, fig. 52A-D, pi. 25, fig. 3. Age, Liasso-Rhaetic. 58. Triletes polysceles Murray, 1939, Geol. Mag., vol. 76, p. 484, figs. 5, 6. Age, Jurassic (Estuarine). 59. Triletes praetextus Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 1, p. 24, pi. 26, figs. 1-6, pi. 27, figs. 1-6, Text-fig. 10. Type 21 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 174, pi. 8, figs. 24, 25. Age, Upper Dinantian and Lower Na- murian. Note. — Closely related to Triletes difuso- pilosus. 60. Triletes potosiensis Zerndt, 1938, De- uxieme Cong. Stratig. Carbonifere, vol. 3, p. 1728, pi. 155, fig. 4. Sectio, Auriculatif. Age, Carbonifer- 61. Triletes radiatus (Ibrahim) S. W. and B., comb. nov. Sporonites radiatus Ibrahim, 1932, Neues Jahrb, Beilage-Band 67, Abt. B, p. 449, pi. 16, fig. 25. Zonales-sporites radiatus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 29, pi. 3, fig. 25. Sectio, Triangulati. Age, Aegir coal, at boundary of Westphalian B and C. 62. Triletes radiosus S. W. and B. nom. nov. Triletes radiatus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, p. 10, fig. 7, pi. 13, figs. 1-5. Types 22 and 23 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 174, pi. 8, figs. 24, 25. Age, Lower Namurian (Upper Di- nantian?). '" Note. — T. radiatus Zerndt, 1937, becomes a later homonym in conflict with Sporonites radi- atus Ibrahim when the latter is transferred to Triletes (see sp. 61 above), thus necessitating a new name as here proposed. 63. Triletes reinschi (Ibrahim) Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 24, pi. 2, figs. 2-4, pi. 5, figs. 8, 9. Triletes I Bennie and Kidston, 1886, Royal Phys. Soc. Edinburgh Proc, vol. 9, p. 107, pi. Ill, figs, la, b. Spore 1.9 mm. Zerndt, 1930, Soc. Geol. Pologne, Ann. vol 6, p. 308, 312, pi. 1, figs. 5a, b, pi. Ill, figs. 5a, b. Triletes Type 1 Kidston, Zerndt, 1930, i- Acad, polonaise sci. Bull, internat., ser. B, p. 43, pi. 1, fig. 1. Type 10 Zerndt, 1931, idem., ser. A, p. 172. Triletes I Kidston, Stach and Zerndt, 1931, Gluckauf, Jahrg. 67, no. 35, p. 1122. Triletes Type 1 Kidston, Maslankie- wiczowa, 1932, Acta soc. Bot. poloniae, vol. 9 (Suppl.), p. 158. Sporonites reinschi Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 17, fig. 28. Laevigati-sporites reinschi (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 18, pi. 4, fig. 28. Reticulati-sporites reinschi (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 157. Sporites primus Wicher, 1934, idem, no. 4, p. 169. Types 1 to 6 incl., Rousseau, 1935, Mus. royale histoire nat. Belgique Bull. vol. 11, no. 21, pp. 3-4, pi. 1, figs. 1-5, pi. 2, fig. 6. Type 1 Sahabi (pars.), 1936, spores des houilles Francaises, p. 34, pi. 1, figs. 1, 2.. Sectio, Aphano3onati. Age, Westphalian C & D, Lower Stephanian. Note. — Type species of Triletes by designa- tion (Schopf, 1938). 64. Triletes reticulatus Zerndt, 1938, De- uxieme Cong. Stratig. Carbonif., vol. 3, p. 1728, pi. 155, fig. 5. Age, Mesozoic. 65. Triletes rexargenteus Harris, 1935, Med- delelser om Gronland, vol. 112, no. 1, pp. 156-158, fig. 51G-J, pi. 26, fig. 13. Age, Liasso-Rhaetic. 66. Triletes richardsoni Murray, 1939, Geol. Mag., vol. 76, pp. 482-484, figs. 9, 10. Age, Jurassic (Estuarine). 67. Triletes robertianus S. W. and B., nom. nov. Triletes kidstoni Zerndt, 1934, Acad. polonaise sci. Trav. Geol. no. 1, pp. 26-27, fig. 12, pi. 28, figs. 6-11, pi. 29, figs. 1-13. Type 27 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 175, pi. 9, figs. 29-32. Sectio, Lagcnicula. Age, Dinantian, Namurian, Westphalian A. Note. — Closely related to Triletes bipilosus and to T. latihirsutus. The new name is neces- sitated by preoccupation of the specific epithet (species no. 2t7). This species is likewise named for Robert Kidston. 68. Triletes rotatus Bartlett, 1928, Michigan Acad. sci. Papers, vol. 9, p. 21, pis. 9, 10, 11, 12. Type 19, Triletes rotatus Bartlett, Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 1, pp. 21-22, fig. 8, pi. 24, figs. 1-6. Triletes rotatus Bartlett, Zerndt, 1937, idem., no. 3, pp. 8-10, pis. 6-10. Age, type material probably upper Pottsville. TRILETES 25 Triletes rotatus var, denticulata Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, pp. 9-10, fig. 6 (?),pl. 6, fig. 3. Age, Dinantian, Lower Namurian and above (?). 69. Triletes rugosus (Loose) Schopf, 1938, Illinois Geol. Surv. Rept. Inv. 50, pp. 29- 30, pi. 5, fig. 6. . Sporonites rugosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 59. Levigati-sporites rugosus (Loose) Ibra- him, 1933, Sporenformen des Aegirhori- zonts, pp. 18-19, pi. 7, fig. 65. Punctati - sporites rugosus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 146. Sectio, Lagenicula. Age, Upper West- phalian B and above (?). Note. — May be closely allied with Triletes translucens. 70. Triletes saarensis Zerndt, 1941, Paleonto- graphica, Abt. B, vol. 84, p. 134, pi. 12, figs. 40-44. Type 47 Zerndt, 1941 (see above refer- ence). Sectio, Auriculati (?). Age, West- phalian C and Lower Westphalian D. 71. Triletes saturnoides (Ibrahim) S. W. and B., comb. nov. Sporonites saturnoides Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 16, fig. 26. Zonales-sporites saturnoides (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 27-28, pi. 3, fig. 26. Age, Upper Westphalian B. Note. — Allied with Triletes brasserti, T, superbus and T. circumtextus. 72. Triletes sextus S. W. and B., nom. nov. Triletes VI Bennie and Kidston, 1886, Royal Physical Soc. Edinburgh Proc, vol. 9, p. 109, pi. 3, figs. 6a, 6b, 6c. Sectio, Aphanosonati. Age, Lanarkian=: Namurian. Note.— Forms of this sort are included by Zerndt (1931, 1934) in Type 14. Inasmuch_ as this was used as the type for Apiculati-sporites Ibrahim, it seems desirable to have an orthodox designation for reference. 7?). Triletes sextusdecimus S. W. and B. nom. nov. Triletes XVI Bennie and Kidston, 1886, idem. pp. 113-114, pi. 5, figs. 16a, 16b, 16c, 16d, 16e. Age, Bernician and Lanarkian = ap- prox. Upper Dinantian and Namurian. Note. — Related to Triletes superbus, and prob. to T. circumtextus and T. brasserti. This form ■was taken as the type oi Zonales-sporites Ibra- him, hence it seems desirable to propose an orthodox name for purposes of reference. 74. Triletes silvanus (Ibrahim) S. W. and B., comb. nov. Sporonites silvanus Ibrahim, 1932 Neues Jahrb., Beilage-Ba*^d 67, Abt. B, p, 448, pi. 15, fig. 22. Laevigati - sporites silvanus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p 20, pi. 2, fig. 22, pi. 6, fig. 47. Sporites silvanus (Ibrahim) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, No. 4, p 174, pi. 8, figs. 5-8. Sectio, Auriculati. Age, Upper West- phalian B, Lower Westphalian C. Note. — Closely related to Triletes auritus. 75. Triletes simplex (Zerndt) S. W. and B., comb. nov. Lagenicula simplex Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, pp. 14-15, fig. 10, pi. 23, figs. 1-5. Type 35 Zerndt, 1937, idem. Sectio, Aphanozonati (?). Age, Lower Namurian. Triletes simplex var. levis Zerndt, 1937, idem., p. 15. Type 36 Zerndt, 1937, idem. 76. Triletes sofiaense S. W. and B., nom. nov. Triletes auritus III Maslankiewiczowa, 1932, Acta Soc. Botanicarum Poloniae, vol. 9 (Suppl.) pp. 161-163, 172-173, figs. Z7, 38. Sectio, Auriculati. Age, Upper West- phalian B (Laziska series). Note. — The group is clearly distinguished specifically by coarse reticulation, in this respect somewhat similar to Triletes tuberculatus, though in shape it resembles Triletes auritus. Possibly it represents a connecting link between these species. It is named for Zofja Kowalew- ska Maslankiewiczowa who provided an ample description of the form. 77. Triletes sparassus Murray, 1939, Geol. Mag., vol. 76, p. 480, p. 482, figs. 3, 4. Age, Jurassic (Estuarine). 78. Triletes splendida (Zerndt) S. W. and B., comb. nov. Lagenicula splendida Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no 3, pp. 13-14, pis. 18-20. Type 28 Zerndt, 1931, Acad, polonaise sci. Bull, internat., Ser. A, p. 175, pi. 10, figs. 36, 37. Sectio, Aphanosonati. Age, Upper Di- nantian, Lower Namurian. A^o^^.— The lageniculate vestibule possessed by this form is insufficient basis for assigning it to Lagenicula in disregard of its other charac- teristic Aphanosonati features. 79. Triletes stenoxysmatodes Harris, 1935, Meddelelser cm Gronland, vol. 112, no. 1, p. 167, pi. 25, fig. 6. Age, Liasso-Rhaetic. 80. Triletes subfuscus (Wicher) S. W. and B., comb. nov. 26 PALEOZOIC FOSSIL SPORES Sporitcs subfnscns Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 179, pi. 8, fig. 20. Type 14 Zerndt (in part), 1934, Acad, polonaise sci. Trav. Geol. no. 1, pp. 17, 18, fig. 5, pis. 8-17. Sectio, Aphanosonati. Age, Lower Westphalian C. 81. Triletes subrotundus (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, p. 155, p. 165. Selaginellites subrotundus Miner, 1932, Washington Acad. Sci. Jour., vol. 22, p. 505, figs. 4, 5. Age, Upper Cretaceous. 82. Triletes subtilinodulata (Nowak and Zerndt) S. W. and B., comb. nov. Lagenicula subtilinodulata, Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, in- ternat, ser. A, pp. 59-60, pi. 1, fig. 7. Type 42 Zerndt, Nowak and Zerndt, 1936 (idem). Sectio, Lagenicula. Age, Dinantian. 83. Triletes subpilosus (Ibrahim) S. W. and B., comb. nov. Setosi-sporites subpilosus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 27, pi. 5, fig. 40. Sectio, Lagenicula. Age, Aegir coal, boundary of Westphalian B and C. Note. — May be closely related to Triletes robertianus. 84. Triletes superbus Bartlett, 1928, Michigan Acad. Sci. Papers, vol. 9, pp. 20, 21, pi. 7, figs. 1, 2, pi. 8, figs. 1, 2. Age, Probably upper Pottsville. Note. — Probably allied with Triletes circum- textus and T. brasserti. 85. Triletes tenuigollatus Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, internat., ser. A, p. 59, pi. 1, fig. 3. Type 41 Zerndt, Nowak and Zerndt, 1936 (idem). Age, Dinantian. 86. Triletes tenuimembranosa (Zerndt) S. W. and B., comb. nov. Lagenicula tenuimembranosa Zerndt, 1937, idem. p. 587, pi. 14, figs. 1, 2. Sectio, Lagenicula. Age, Westphalian C and Lower Westphalian D. Type 25 Zerndt, 1931, idem., p. 175, pi. 9, figs. 2>?> and 35. Sectio, Lagenicula. Age, Lower Na- murian and Westphalian A. Note. — Closely related to Triletes translucens. 87. Triletes tenuispinosus Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 1, p. 16, text-fig. 4, pi. 7, figs. 1-15. Type 13 Zerndt, 1934 (idem). Triletes tenuispinosus var. brevispinosa Zerndt, 1937, idem., no. 3, p. 6, fig. 4, pi. 3, figs. 2-7. Triletes tenuispinosus var. secundus S. W. and B., var. nom. nov. Triletes tenuispinosus var. II Zerndt, 1937, idem., no. 3, pp. 7-8, fig. 5, pi. 3, figs. 3 and 4. Age, Dinantian — Upper and Lower Namurian. Note. — Zerndt describes three forms, one which evidently is typically the species and two varieties, only one of which is properly named. The new varietal name proposed provides a means of reference. 88. Triletes translucens Schopf, 1938, Illi- nois Geol. Survey Rept. Inv. 50, pp. 28-29, pi. 5, figs. 3-5. Sectio, Lagenicula. Age, Middle Penn- sylvanian, upper Carbondale, lower Mc- Leansboro series. Note. — Closely related to Triletes tenuimem- branosa, T. levis, and (?) T. rugosus. 89. Triletes triangulatus Zerndt, 1930, Acad. polonaise sci. Bull, internat, ser. B, p. 51, pi. 7, figs. 19-24. Spore 0.5 mm. Zerndt, 1930, Soc. Geol. Pologne Ann., vol. 6, pp. 306, 312, pi. 1, figs, la, b, c, pi. 2, figs, la, b. Type 17 Zerndt, 1931 Acad, polonaise sci. Bull, internat. ser. A, p. 173. Sporonites regalis Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 16, fig. 24. Sporites triangulatus "var. regali/' (Zerndt) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 29, pi. 3, fig. 24. Sporites regalis (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149, pi. 7, fig. 34. Sporites triangulatus (Zerndt) Wicher, 1934, idem., vol. 4, no. 4, p. 175. Triletes triangulatus Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 1, p. 19, fig. 6, pi. 18, figs. 1-24. Triletes triangulatus Zerndt, Schopf, 1936, Illinois Acad. Sci., Trans, vol. 28, p. 107, fig. 6. Type X Sahabi, 1936, spores des houil- les Francaises, p. 43, fig. 11, pi. 4, figs 6-11. Triletes triangulatus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, pi. 5, figs. 1-5. Triletes triangulatus Zerndt, Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, pp. 32-37, pi. 1, figs. 7, 8, pi. 4, figs. 1-7, pi. 7, figs. 5, 6. Triletes triangulatus Zerndt, 1940, Paleontographica, vol. 84, Abt. B, pi. 10, figs. 19-23. Triletes triangulatus var. zonatus (Ibrahim) Schopf, 1938, Illinois Geol. Sur- vey Rept. Inv. 50, p. 34. Triletes triangulatus II Zerndt, 1930, Acad, polonaise sci. Bull, internat., ser. B, p. 53, pi. 7, figs. 25-30. Sporonites zonatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 16, fig. 23. Zonales-sporites triangulatus secundus Ibrahim, 1933, Sporenformen des Aegirhor- izonts, p. 30, pi. 3, fig. 23, pi. 7, fig. 64. PITYOSPORITES 27 Zonales-sporites zonatus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 150, pi. 7, fig. 31. S pontes triangulatus var. sonatus (Ib- rahim) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, pp. 175-176. Triletes triangulatus var.? Triletes triangulatus III, Stach and Zerndt, 1931, Gliickauf, Jahrg. 67, no 35, p. 1123, figs. 32, 2,Z. Sectio, Triangulati. Age, upper Lower Namurian through Westphalian D. 90. Triletes tricollinus Zerndt, 1938, Deux- ieme Cong. Stratig. Carbonif. vol. 3, p. 1713, pi. 155, fig. 3. Type 44 Zerndt, 1938 (idem). Sectio, Triangulati. Age, Westphalian B, C, and D. 91. Triletes trilobus (Ibrahim) S. W. and B., comb. nov. Sporonites trilobus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 17, fig. 30. Valvisi-sporites trilobus (Ibrahim) Ib- rahim, 1933, Sporenformen des Aegirhor- izonts, p. 33, pi. 4, fig. 30. Valvisi-sporites trilobus (Ibrahim) Ib- rahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 152, pi. 17, fig. 30. Sectio, Auriculati. Age, Upper West- phalian B. Note. — Possibly very closely related to Tri- letes appendiculatus. Wicher (1934) regards it as similar to T. auritus. 92. Triletes tuberculatus Zerndt, 1930, Acad, polonaise sci. Bull, internat. ser. B, pp. 47-51, pi. 2, figs. 6, 7, pi. 3, figs. 8, 9, pi. 4, figs. 10,' 11, pi. 5, figs. 12, 13. Triletes XIX Kidston, 1890, Royal Soc. Edinburgh Trans., vol. 36, pi., figs. 9-11. Type 16 Zerndt, 1931, Acad, polonaise sci. Bull, internat., ser. A, p. 173. Triletes tuberculatus Zerndt, Maslan- klewiczowa, 1931, Acta Soc. Bot. Poloniae, vol. 9, Suppl. pp. 165-168, figs. 40, 41. Sectio, Auriculati. Age, Westphalian B, C, Stephanian? 93. Triletes tuberosus (Ibrahim) S. W. and B., comb. nov. Sporonites tuberosus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 16, fig. 27. Tuber culati-sporites tuberosus (Ibra- him) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 23, pi. 3, fig. 27. Tuberculati-sporites tuberosus ( Ibra- him) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 147, pi. 7, fig. 37.^ Sporites tuberosus (Ibrahim) Wicher, 1934, idem, vol. 4, no. 4, pp. 177-178, pi. 8, fig. 22. Sectio, AphanoBonati. Age, Upper Westphalian B, Lower Westphalian C. Note. — Probably the same as some of Zerndt's Type 14. 94. Triletes tylotus Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, pp. 162-163, fig. 53F-I, pi. 26, figs. 1, 12. Age, Liasso-Rhaetic. 95. Triletes valens (Wicher) S. W. and B., comb. nov. Sporites valens Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 178, pi. 8, fig. 19. Sectio, Aphanosonati. Age, Lower Westphalian C. Triletes nomen excludende 1. Apiculati-sporites megaspinosus Ibrahim, 1933, Sporen- formen des Aegirhorizonts, etc., p. 24, pi. 8, fig. 69. Note: This form is not congeneric with the type of Apiculati-sporites {Triletes sextus) and, although it ap- Earently is sufficiently characterized and characteristic to e a useful microfossil, no further generic assignment will be attempted. Genus Pityosporites Seward, 1914 Plate 2, figures 15-15b Diagnoses given by Seward are as follows : The generic designation Pityosporites is pro- posed for winged spores agreeing in form and size with recent Abietineous genera (1914, p. 23). .... Spores provided with bladder-like extensions of the exine, agreeing in size and form with those of recent Abietineous genera (1919, p. 398). A more adequate definition inferred from the above, and from consideration of the species included here, is as follows : Symmetry. — Pollen grains appearing bilateral, due to the presence of bladders placed on opposite sides. The funda- mental symmetry, shown by the body but modified, is radial, as this pollen arises from tetrahedral tetrads. Shape — Grains elliptical as viewed from proximal side, the diameter of the poles of the ellipse (formed by the two bladders) being broader than that of the body between them. Body appears ap- proximately circular in proximal view. Viewed from either side the bulbose blad- ders are inclined distally and a broad sulcus appears between them on the distal side. In lateral outline the body appears broadly oval with the smaller pole toward the distal sulcus. Size. — Various species range in total length (bladder tip to bladder tip) from as small as 40 to over 100 microns. Ornamentation. — External bladder sur- face minutely granulose or rugose ; in- ternally the bladders are usually at least weakly reticulate. Proximal cap may be nearly smooth, rugose or striate. Distal body wall least ornamented. 28 PALEOZOIC FOSSIL SPORES Haptotypic features. — Lacking or near- ly so. Presumably a weak vestigial imprint of the trilete mark may be present on the proximal cap in some forms as it oc- casionally is in modern Abies. Pollen coat. — The unique character of internal bladder wall reticulation distin- guishes these and many other coniferous pollen grains. The internal reticulae add apparent thickness to the perineal blad- der membrane which is quite thin. The exinal (body) wall is much thicker proximally and is least translucent, sometimes appearing brownish under the microscope ; the distal wall is much thin- ner, serving as a harmomegathus and point of germinal exit when ruptured by the expanding gametophyte. Affinity. — There can be little question that grains of this sort are referable to the Coniferae. Members of modern Podocarpaceae and Pinaceae show essen- tially similar features. Correlatives of species of Pityosporites represented by vegetative organs are hardly recognized in Paleozoic formations. The presence of such pollen is evidence that such plants existed, possibly as a xerophytic upland flora, and tend to substantiate the view long held that an important ancestry, very scantly represented among megascopic fossils, preceded those highly developed forms found better represented in younger strata. Pityosporites represents a group of Coniferae whose pollen is essentially mod- ern in aspect. Florin's brilliant researches (1938-1940) have sufficiently demon- strated that Lebachia, Ernestiodendron, and Walchianthus for the most part do not possess pollen of the Pityosporites type ; most, if not all of the probably correlated but isolated pollen of this sort is referable to Florinites n. gen. described below (p. 56). Remarks. — It will be evident that the interpretation given Pityosporites here differs from that of Florin. He (1940, pp. 327-8) considers Pityosporites to be a completely artificial genus which has no type species. Apparently he would assign any pollen-like form with two bladders to this group. It is evident from Seward's descriptions, however (1914, 1919, 1933), that the emphasis is not so much merely on the presence of two bladders as it is on the essentially modern aspects of these fossils which have been more precisely itemized in our revised definition.' A more restricted group is thus indicated than that considered by Florin, and this group surely is not an artificial one although it certainly exceeds the scope of most "normal" genera, and, in fact, possibly contains elements which would be classed in separate families if more complete characterization could be obtained. Nevertheless, we think it neces- sary to treat the group as a genus at present, with the hope that critical studies later will permit it to be subdivided into somewhat smaller groups that will more perfectly represent the relationships in detail. Whether this hope is a vain one cannot be foretold a priori, but the desira- bility of recognizing a type species for this group which, as we construe it, does signify a definite natural plant alliance, cannot be denied. Part of the taxo- nomist's responsibility is to place the sys- tematic data in order so that later students may revise it to greater perfection in the light of additional data. Such progress is very difficult if nomenclatural types are not definitely indicated and fully utilized in the application of technical nomencla- ture. In paleobotanical studies it is particularly important that this be recog- nized because the nature of the material that is being studied rarely permits cate- gorical statement and new discoveries call for greater revision in preexisting con- cepts, perhaps, than in any other plant science. Pityosporites can hardly be considered as synonymous with any modern genus; perhaps the most important reason is that when pollen characters are sufficiently distinctive to permit actual identification with a more precisely delimited modern group, identification with the genus Pityo- sporites becomes inadequate as an ex- pression of affinity. Quite evidently many of the isolated ancient pollen grains can- not be more definitely assigned and Pityo- sporites will serve a most useful function in these instances. ■^Although Seward (1919, pp. 398-9) has included one of the forms Nathorst reported from the Hor clay as "Pityosporites sp." (which we should prefer to assign to Alisporites) , Seward apparently did this on the as- sumption that Nathorst's specimen was similar to pollen of Picea excelsa. Wodehouse (1935) has shown Picea pollen to be very different. The Pityosporites sp. of Solms from Franz Josef Land (Seward, 1919, p. 399) no doubt is correctly assigned. PUNCTATI-SPORITES 29 In addition to the nine species listed be- low, Kosanke (1943) also has recorded material of this nature from the Upper Pennsylvanian of Ohio. 1. PiTYOSPORiTES ANTARCTicus Scward, 1914, Nat. Hist., Report British Antarctic (Terra Nova) Exped. 1910. Geology vol. 1, no. 1, pp. 23-4, pi. 8, fig. 45. Pityosporites antarcticus Seward, 1919, Fossil Plants, vol. IV, p. 398. Pityosporites antarcticus Seward, 1933, New Phyt, vol. 32, no. 4, pp. 311-313, fig. 1. Age, "Not older than Lower Meso- 2. Pityosporites chinleana Daugherty, 1941, Carnegie Inst. Washington, Pub. 526, p. 398, pi. 34, fig. 5. Age, Upper Triassic. 3. Pityosporites (?) jeffreyi Florin, 1940, Paleontographica vol. 85, Abt. B, no. 5, p. 327, pi. 163-4, figs. 8-12. Pityanthus jeffreyi Florin (nomen nudum?), 1927, Arkiv. for Botanik, vol. 21 A, no. 13, p. 6. (Systematic status of this nomenclatural combination is difficult to decide, since illustrations needed to validate it were not published till 1940 when the species was transferred to Pityosporites.) Age, Middle Stephanian. Note. — BVadders are seemingly set nearly op- posite though it is possible their distal inclina- tion merely is not apparent in the figures. The proximal cap also is not evident and it may be this form belongs to Alisporites rather than in the present genus. 4. Pityosporites pallidus Reissinger (nomen nudum), 1939, Paleontographica, vol. 84, Abt. B, p. 14. Age, Lowest Jura (Lias). Note. — An illustration required to validate the name has not yet been published. 5. Pityosporites sewardi Virkki, 1937, In- dian Acad. Sci. Proc, vol. 6, no. 6, sect. B, pp. 428-431, figs. 2a, b, c, d, pi. 32, figs, la, b, c. Age, Lower Gondwana, above Talchir boulder bed. 6. Pityosporites stephanensis Florin, 1940, Paleontographica, vol. 85, Abt. B, no. 5. pp. 327-8, pi. 163-4, figs. 13-16. Age, Middle Stephanian. 7. Pityosporites sp. Florin, 1940, idem, p. 61, pi. 163-4 fig. 17, fig. 18? Age, Lower Rothliegendes. 8. Pityosporites (?) sp. Florin, 1940, idem, p. 62, pis. 165-6, fig. 20. Age, Upper Carboniferous. Note. — No inclination of bladders shown. 9. Pityosporites sp. Daugherty, 1941, Car- negie Inst. Wash., Pub. 526, description of pi. 34, figs. 7 and 8. Age, Keuper. Genus Punctati-sporites (Ibrahim, 1933) emend., S. W. and B., Plate 1, figures 4-4b Symmetry. — Spores radial, trilete. Shape. — Originally nearly spherical, or possibly broadly rounded triangular with slight shortening of the axial dimension; when compressed the spores show no proximo-distal orientation preference. Size. — Spores of various species range from 45 to 85 microns in mean diameter. Ornamentation. — Various ; surfaces levigate to punctate, rugose, reticulate or mildly apiculate. Haptotypic structures. — Of moderate prominence, relative length of trilete rays highly variable between different species ; no equatorial or arcuate markings (con- necting ends of trilete rays) are present; lips of the trilete commissure are never very prominent nor highly ornamented. Spore coat. — Generally thin, and, ex- cept in instances where ornamental fea- tures are most prominent, hardly in excess of 3 microns. Affinity. — Spores of this genus are similar to those obtained from certain pteridospermic fructifications and from some fossils assigned to the ferns. The spores Kidston (1906) obtained from va- rious Crossotheca fructifications are char- acteristic of typical species of Punctati- sporites. However, many of the species represented by isolated spores have no known pteridospermic relationship al- though there is no adequate basis for separating them from forms which are thus allied. Much remains to be discov- ered as to the affinities of this genus, which is a rather large one, and it may be expected that some diverse elements have been included which will be segre- gated on the basis of newer information. It is quite possible that some of the forms assigned to Punctati-sporites are merely immature and lack their distinc- tive mature ornamentation. Such may be the case for immature forms that in reality belong to Raistrickia n. gen. de- scribed below (p. 55). Immature forms should not be arbitrarily assigned to Punctati-sporites but it is well to recog- nize that when these are present in sub- stantial numbers, they need to be reported and that a superficial resemblance to 30 PALEOZOIC FOSSIL SPORES P unci ati-s pontes may not always be in- dicative of relationship. The affinity of certain of the species with the Crossotheca alliance of pteridosperms seems beyond much question, however. The forms which Raistrick (1933, 1934, 1937) has assigned to his types B„ Be, D3, D„ D,5 (Knox, 1938), E„ Ee, E„ E„ Eg, F„ Fs (Knox, 1938) (also 1K( ?j, 2K and 7K, Knox, 1942) appear entirely or in part referable to Punctati- sporitcs. It also may include some of the forms he has classed as Bg and Bg. Some difficulty is encountered because Raistrick evidently has designated no sin- gle typical example for these groups and his illustrations of the forms have differed for individual types beyond what may represent a specific range of variability. Thus there may be some question as to the consanguinity of forms labeled as D3 (cf. also Raistrick, 1935). Figures and description of E^ given in 1933 do not ex- actly correspond with that given in later papers (1934, 1935). Illustrations of B3 given in 1933 seem to resemble Punc- tati-sporites but those illustrated in later papers without much doubt are referable to Calamospora n. gen. described below. Since Raistrick's procedure is non-tax- onomic it is impossible to form more than a general opinion as to specific equiv- alences in most instances. Remarks. — The genus Punct ati-s pontes as given here includes twenty-nine species, four of which are queried. All but three of these names (see spp. Nos. 9, 21, 22) are new combinations. The group Punc- tati-sporites was originally defined by Ibrahim (1933, p. 21) as including trilete type spores with a granular surface, char- acters which by themselves can hardly be accepted as diagnostic of any generic group. Three species were described, one of which, P. punctatus, was designated as the type. Consequently it has been pos- sible to define the genus practically, only by segregating those forms which ap- peared congeneric with P. punctatus, and the emended generic diagnosis has been constructed on this basis. Neither of the other two species Ibrahim originally placed in the genus are considered closely enough related to remain there ; P. parvus Ibrahim is now assigned to the genus Granulati-sporites, and P. pallidus (Loose) Ibrahim to Calamospora n. gen. Six species previously assigned to Ver- rucosi-sporites Ibrahim, including the type, are here placed under Punctati-spor- ites. V errucosi-sporites thus would more reasonably be taken as the designation for the genus except that in meaning one name is no more suitable than the other and strict interpretation of page priority (p. 21 vs. p. 25) favors Punctati-sporites. The genus also includes the type of Apiculata-sporites Ibrahim (1933), a monotypic genus which seems based en- tirely on Ibrahim's inability to observe the trilete sutures described by Loose (1932) on Sporonites spinulistratus. The presence of the trilete marking on this species was reaffirmed by Loose in 1934. In any event the genus Apiculata-sporites does not seem securely founded and its one species is here placed under Punctati- sporites. Both groups, V errucosi-sporites Ibrahim and Apiculata-sporites Ibrahim, conseqently are regarded as congeneric with and the names synonyms of Punctati- sporites. The holotype of the type species, Punc- tati-sporites punctatus (Ibrahim) Ibra- him, is from the Aegir coal seam at the top of the Westphalian B in the Ruhr district of Western Germany. 1. Punctati-sporites aculeatus (Ibrahim) S. W. and B., comb. nov. Apiculati-sporites aculeatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 23, pi. 6, fig. 57. 2. Punctati-sporites aureus (Loose) S. W. and B., comb. nov. Reticulati-sporites aureus Loose, 1934, Inst. Paleobot. Arb., vol. 4, no. 3, p. 155, pi. 7, fig. 24. 3. Punctati-sporites (?) auriculaferens (Loose) S. W. and B., comb. nov. Sporonites auriculaferens Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, pi. 18, fig. 39. Valvisi-sporites auriculaferens (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 152. 4. Punctati-sporites eucculentus (Loose) S. W. and B., comb. nov. V errucosi-sporites bucculentus Loose, 1934, idem. p. 154, pi. 7, fig. 15. 5. Punctati-sporites corrugatus (Ibrahim) S. W. and B., comb. nov. Reticulati-sporites corrugatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, pp. 35-36, pi. 5, fig. 41. PUNCTATI-SPORITES 31 6. PUNCTATI-SPORITES CUSUS (LoOSC) S. W. and B., comb. nov. Reticulati-sporites cusus Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 156, pi. 7, fig. 25. 7. PUNCTATI-SPORITES FIRMUS (LoOSc) S. W. and B., comb. nov. Verrucosi-sporites firmus Loose, 1934, idem, p. 154, pi. 7, fig. 30. 8. PUNCTATI-SPORITES GLOBOSUS (LoOSe) S. W. and B., comb. nov. A piculati-s pontes globosus Loose, 1934, idem, p. 152, pi. 7, fig. 14. 9. PUNCTATI-SPORITES GRANDIVERRUCOSUS Kosanke, 1943, Am. Midland Naturalist, vol. 29, no. 1, pp. 127, 128, pi. 3, fig. 4. 10. PuNCTATi-SPORiTES GRANiFER (Ibrahim) S. W. and B., comb. nov. Granulati - sporites granifer Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 22, pi. 8, fig. 72. 11. PuNCT ATI- SPORITES GRUMOSUS (Ibrahim) S. W. and B., comb. nov. Verrucosi-sporites grumosus Ibrahim, 1933, idem, p. 25, pi. 8, fig. 68. 12. PuNCTATi-spoRiTES iNsiGNiTis (Ibrahim) S. W. and B., comb. nov. Apiculati - sporites insignitis Ibrahim, 1933, idem, p. 24, pi. 6, fig. 54. 13. PuNCTATi-sPORiTEs( ?) IRREGULARIS (Ber- ry) S. W. and B., comb. nov. Zonales-sporites irregularis Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, p. 156, fig. 9. 14. PuNCTATi-SPORiTES LACUNOSUS (Ibrahim) S. W. and B., comb. nov. Reticulati - sporites lacunosus Ibrahim, 1933. Sporenformen des Aegirhorizonts, p. 36, pi. 6, fig. 50. 15. PUNCTATI-SPORITES LATIGRANIFER (LoOSC) S. W. and B., comb. nov. Sporonites latigranifer Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 452, pi. 19, fig. 54. Granulati-sporites latigranifer (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 147. 16. PuNCTATi-spoRiTES MACULATUS (Ibrahim) S. W. and B., comb. nov. Reticulati-sporites maculatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 2,6, pi. 6, fig. 56. 17. PUNCTATI- SPORITES M ICROTUBERO SU S (Loose) S. W. and B., comb. nov. Sporonites microtuberosus Loose, 1932, Neues, Jahrb., Beilage-Band 67, Abt. B, p. 450, pi. 18, fig. 33. Tuberculati - sporites microtuberosus (Loose) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 23. Tuberculati - sporites microtuberosus (Loose) Ibrahim, Loose, 1934, Inst. Palao- bot. Arb., vol. 4, no. 3, p. 147. 18. PuNCTATi-spoRiTES MicROVERRUCOsus (Ibra- him), S. W. and B., comb. nov. Verrucosi - sporites microverrucosus Ibrahim, 1933, Sporenformen des Aegirhori- zonts, p. 25, pi. 7, fig. 60. 19. PuNCTATi-SPORiTES NOBiLis (Wicher) S. W. and B., comb. nov. Sporites nobilis Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 186, pi. 8, fig. 30. 20. PuNCTATi-SPORiTES PAPILLOSUS (Ibrahim) S. W. and B., comb. nov. Verrucosi-sporites papillosus Ibrahim, 1933, Sporenformen des Aegirhorizonts, pi. 5, fig. 44. 21. PUNCTATI-SPORITES PARVIPUNCTATUS Kos- anke, 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 127, pi. 3, figs. 3, 3a, 3b. 22. PuNCTATi-sPORiTES PUNCTATUS (Ibrahim) Ibrahim, 1933, Sporenformen des Aegirhori- zonts, p. 21, pi. 2, fig. 18. Sporonites punctatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 57, Abt. B, p. 448, pi. 15, fig. 18. Punctati-sporites punctatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 146, pi. 7, fig. 27. 23. PUNCTATI-SPORITES (?) RETICULOCINGULUM (Loose) S. W. and B., comb. nov. Sporonites reticulocingulum Loose, 1932, Neues Jahrb., Beilage-Band 57, Abt. B, p. 450, pi. 18, fig. 41. Reticulati - sporites reticulocingulum (Loose) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 34. Reticulati - sporites reticulocingulum (Loose) Ibrahim, Loose, 1934, Inst. Palao- bot. Arb., vol. 4, no. 4, p. 156. 24. PuNCTATI - SPORITES SPHAEROTRIANGULATUS (Loose) S. W. and B., comb. nov. Sporonites sphaerotriangulatus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 18, fig. 45. Laevigati - sporites sphaerotriangulatus (Loose) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 20. Laevigati - sporites sphaerotriangulatus (Loose) Ibrahim, Loose, 1934, Inst. Palao- bot. Arb., vol. 4, no. 4, p. 145. 25. PuNCTATi-sPORiTES SPiNosus (Loose) S. W. and B., comb. nov. A piculati-s porites spinosus Loose, 1934, idem, p. 153, pi. 7, fig. 20. 26. PUNCTATI-SPORITES SPINULISTRATUS (LoOSC) S. W. and B., comb. nov. Sporonites spinulistratus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 451, pi. 18, fig. 47. 32 PALEOZOIC FOSSIL SPORES Apiculata-sporitesspinulistratus (Loose) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 37. A piculati-sporitcsspmulistratus (Loose) Loose, 1934, Inst. Palaobot, Arb., vol. 4, no. 4, p. 153. 27. PUNCTATI - SPORITES TRIGONORETICULATUS (Loose) S. W. and B., comb. nov. Reticiilati - sporitcs trigonoreticulatus Loose, 1934, idem, p. 155, pi. 7, fig. 9. 28. PUNCTATI-SPORITES (?) VELATUS (LoOSe) 5. W. and B., comb. nov. Reticulati-sporites velatus Loose, 1934, idem, p. 155, pi. 7, fig. 19. 29. PuNCTATi-sPORiTES VERRUCOSUS (Ibrahim) S. W. and B., comb. nov. Sporonites verrucosus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 15, fig. 17. V errucosi-sporites verrucosus ( Ibrahim ) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 25, pi. 2, fig. 17. Genus Granulati-sporites (Ibrahim, 1933) emend., S. W. and B. Plate 1, figures 8-8b Symmetry. — Spores radial, trilete. Shape. — Originally oblate, rounded tri- angular, ^N\.ih. the axial dimension much the shortest ; when compressed the spores are nearly always flattened in good proxi- mo-distal orientation since the transverse plane of the spore, being broadest, tends to parallel the bedding of sediments en- closing them. When preserved thus, the spores appear subtriangular in outHne, with sides either convex or slightly con- cave and the corners rounded. Sise. — Spores of various species range from about 25 to 45 microns in mean diameter. Ornamentation. — Various, surfaces smooth to punctate, finely to relatively coarsely reticulate or apiculate ; the di- verse types of ornamentation which char- acterize various species are not prominent to the extent that they mask the essential shape characteristics of the spores. Haptotypic structures. — Trilete rays are well extended toward the corners; they are never relatively short. The lips are not distinguished by special ornamen- tation, the commissure is relatively simple but definite in all cases ; usually no vari- ations in ornamentation distinguish the proximal (pyramic) side from the rest of the spore coat. Spore coat. — Of relatively uniform thickness throughout, generally thin, and except where modified by ornamentation, often less than 2 microns. Affinity. — Spores similar to those of Granulati-sporites have been reported from fructifications of ferns although rel- atively few of these have been critically described. Spores of Bozveria minor and Renaultia gracilis which are illustrated by Knox (1938) are of similar character. Probably considerable difflculty will be encountered in definitely correlating the genus (further emended as may prove desirable) with any single supra-generic plant group. The many similar features possessed by species assigned to Granulati- sporites suggest that this grouping may be a practical one. The forms Raistrick (1933, 1934, 1937) designates as D^, Dg and D-^^ (Knox, 1938) probably belong to Granulati-spor- ites as defined here. Likewise Millott's Type 4 is of this character (Millot, 1939) . Remarks: The genus as emended in- cludes fifteen species, twelve of which represent new name combinations. The genus was originally established to include spores of trilete-type showing granular sculpturing (Ibrahim, 1933, p. 21), char- acters which in themselves are inadequate ; consequently it must be regarded as co- incidence that two of the three original species (one of which was designated as the type) are retained in the group, along with another that Loose assigned to it in 1934. The genotype of Granulati-sporites is the only genotype species included by revision within the genus and thus there can be no question as to validity of the generic name. The genotype, G. granulatus Ibrahim, also is based on a form from the Aegir coal bed at the top of the West- phalian B. 1. Granulati-sporites deltiformis S. W. and B., nomen nov. Triquitrites deltoides Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 185, fig. 9. Note. — In combination with Granulati-sporites the specific epithet becomes a homonym of G. deltoides (lb.) S. W. and B., as given below. 2. Granulati-sporites deltoides (Ibrahim) S. W. and B., comb. nov. Sporonites deltoides Ibrahirh, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 15, fig. 15. ALATI-SPORITES 33 Laevigati-sporites deltoides (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 20, pi. 2, fig, 2. Note. — The species is larger than most forms assigned to Granulati-sporites. 3. Granulati-sporites fistulosus (Ibrahim) S. W. and B,, comb. nov. Reticulati-sporites fistulosus Ibrahim, 1933, idem, p. 36, pi. 5, fig. 35. 4. Granulati-sporites gibbosus (Ibrahim) S. W. and B., comb. nov. V errucosi-sporites gibbosus Ibrahim, 1933, idem, p. 25, pi. 6, fig. 49. 5. Granulati-sporites granulatus Ibrahim, 1933, idem., p. 22, pi. 6, fig. 51. Note. — Type species of Granulati-sporites. 6. Granulati-sporites microgranifer Ibra- him, 1933, idem, p. 22, pi. 5, fig. 32. 7. Granulati-sporites microsaetosus (Loose) S. W. and B,, comb. nov. Sporonites microsaetosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, pi. 18, fig. 40. Setosi-sporites microsaetosus (Loose) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 26. Setosi-sporites microsaetosus (Loose) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 148. 8. Granulati-sporites microspinosus (Ibra- him) S. W. and B., comb nov. Apiculati-sporites microspinosus Ibra- him, 1933, Sporenformen des Aegirhori- zonts, p. 24, pi. 6, fig. 52. 9. Granulati-sporites parvus (Ibrahim) S. W. and B., comb. nov. Sporonites parvus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 15, fig. 21. Punctati - sporites parvus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 21, pi. 2, fig. 21. Reticulati-sporites parvus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 154, pi. 7, fig. 13. 10. Granulati - sporites piroformis Loose, 1934, idem, vol. 4, no. 3, p. 147, pi. 7, fig. 19. 11. Granulati-sporites (?) priddyi (Berry) 5. W. and B., comb. nov. Zonales-sporites priddyi Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, p. 156, fig. 2. 12. Granulati-sporites torquifer (Loose) S. W. and B., comb. nov. Sporonites torquifer Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, pi. 18, fig. 43. .Reticulati-sporites torquifer (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. - 3/.p. 154. 13. Granulati-sporites trigonus (Ibrahim) S. W. and B., comb. nov. Reticulati-sporites trigonus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. ?>7, pi. 5, fig. 34. 14. Granulati-sporites triquetris (Ibrahim) S. W. and B., comb. nov. V errucosi-sporites triquetris Ibrahim, 1933, idem, p. 26, fig. 61. 15. Granulati-sporites verrucosus (Wilson and Coe) S. W. and B., comb. nov. Triquitrites verrucosus Wilson and Coe, 1940, Am. Midland NaturaHst, vol. 23, no. 1, p. 185, fig. 10. Genus Alati-sporites Ibrahim, 1933 Plate 1, figures 6-6b Symmetry. — Spores radial, trilete. Shape. — Body, exclusive of bladders, is sub-triangular. Sise. — In the vicinity of 80 microns, overall diameter. Ornamentation of spore. — Body moder- ate ; may be punctate, finely reticulate ; bladders smooth to punctate, etc. Blad- ders, in general, less definitely ornamented than spore body. Haptotypic structures. — Trilete rays relatively long, little ornamented. Spore coat. — Consisting of two distinct membranes; the outer (perisporal) mem- brane is very thin and expanded in the in- ter-radial areas to form bladders in a trim- erous (in threes or multiples of three) pattern. The spore body v^all (exospore) is usually more than twice as thick as the bladder membrane, darker, and easily distinguished on that account. Affinity. — Spores of Alati-sporites seem smaller but essentially similar in con- struction to those of Spencerites (Scott, 1898, Kubart, 1910). The three inter- radial bladders are most distinctive in both genera. The significance of this character is hardly well enough under- stood to properly evaluate evidence as to their relationship.^ ^ The haptotypic relations, if any, of the three blad- ders present on pollen of Podocarpus dacrydioides and on Pherosphaera are not known, and comparison with Car- boniferous forms does not seem particularly pertinent for obvious reasons. It seems doubtful that the form described by Daugherty (1941, p. 64) as Spencerites? chinleana is actually refer- able to Spencerites or could be referred to Alati-sporites since it apparently does not possess three interradial bladders. Daugherty does not mention and the figure given does not show the trilete sutures. If these were present the species might be referred to Cirratriradites provided the equatorial appendage was a flange. Lacking these characteristics, his alternative suggestion of a gymnospermic affinity seems most plausible. 34 PALEOZOIC FOSSIL SPORES Spores assigned by Raistrick to type D5 without much question belong to Alati-s pontes. Remarks. — Only one species has been described to date but there is little doubt the group will receive far more recogni- tion. It is highly distinctive and complex enough that specific characters are easily distinguishable. No doubt the character of the bladders will lend itself to the purpose. There is no possibility of these forms being confused with unseparated tetrad groups belonging to other genera although illustrations which have been published might lead one to think so. In addition to its recognition in the Ruhr district and in Britain the genus has been recognized in coals of the Illinois basin by Brokaw and in Tennessee coals by Bentall. These from America appear to be specifically distinct and as yet un- described forms. The generic diagnosis given above has been constructed in the light of this additional information, and while con- siderably augmented over Ibrahim's mea- ger description (1933, p. 32), it may be improper to consider it as a generic emend- ation. The type, of course, is the species given below, which was first described from the Aegir coal in the Ruhr. 1. Alati - SPORITES PUSTULATUS (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 33, pi. 1, fig. 12. Sporonites pustulatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 14, fig. 12. Alati - sporites pustulatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb.. vol. 4, no. 3, p. 151, pi. 7, fig. 4. Type Ds Raistrick, 1937, Congres pour I'avancement des etudes de stratigraphie carbonifere, Heerlen, 1935, Compte rendu, vol. 2, p. 911. Genus Reticulati-sporites ( Ibrahim, 1933) emend., S. W. and B. Plate 1, figures 7-7b Symmetry. — Spores, in essence, radi- ally symmetrical ; some species are weakly trilete and others show no indication of their tetrahedral tetrad origin but are con- sidered trilete in derivation because of their shape. Shape. — Originally spherical or mod- eratel}^ oblate ; compressed snores are cir- cular disk-like, generally without prom- inent folds. Sise. — Spores of various species range from about 40 to 100 microns in diameter. Ornam^entation. — Coarsely and often ir- regularly reticulate ; aside from the coarse ornamentation the body wall may have a variously smooth, punctate or finely re- ticulate texture. In the ''alete" forms reticulation may show a tendency toward a spiral pattern ; in others it may be mostly limited to the distal part of the spore. Haptotypie structures. — The trilete structures may be weakly developed ; no arcuate ridges are developed but the or- namentation may sometimes be absent on the proximal surface. The trilete rays vary considerably in development and length among different species. Spore coat. — Appears to consist often- times of two membranes ; when present, the outer one (perisporal?) is thinner and more or less intimately connected with the coarse reticulation; the inner (exospor- al?) membrane is thicker and sometimes irregular and excentric in its development. The wall may appear thick, and yet be quite translucent. Affinities. — The only spores of this gen- eral or possible character known in fruc- tifications are allied with Sphenophyllum. Spkenophyllum spores are inadequately known in the detail necessary for close comparison, however, and considerably more evidence is needed to form an opin- ion as to the relationship of Reticulati- sporites with other groups. Types C2, C3, F4, and Fg of Raistrick (1934, 1935, 1937) and possibly also F„ F2 (cf. Knox, 1938), Fg and F4, appear to belong to Reticulati-sporites. Types F5 and Fg and D,3 (Knox, 1938) are more problematic. Type 8 of Millott (1939) also belongs here and is most closely related to R. facetus of species now described. Knox' Type 5K may possibly be similar to some species included in Re- ticulati-sporites. Spores illustrated by Knox (1939) from Fifeshire, figs. 49 and 50, unquestionably belong to Reticulati- sporites. Unpublished forms of similar nature have been found in Ohio coals by Wilson and Brokaw, and they have also been recognized in Iowa and Illinois and Tennessee. Remarks. — Seven species are included here under Reticulati-sporites, only one of which represents a new name combina- RETICULATI-SPORITES 35 tion. This form, R. facetus (Ibrahim), was originally designated as the genotype of Reticulata-s pontes with five species assigned to it. Essentially the only char- acter shared by these five was their lack of a visible haptotypic marking. Most diverse types of reticulation characterized their surfaces. R. facetus, however, shows a type of reticulation which is very simi- lar to that of R. reticulatus (genotype of Reticulati-sporites) and others in this genus. Other characters are also in sub- stantial agreement and it is believed that the lack of a trilete marking in this case is insufficient basis for generic separation. Other species of Reticulati-sporites show a very weak trilete marking. Consequent- ly Reticulata-sporites Ibrahim (1933, p. 38) may be dispensed with and regarded as a later synonym for Reticulati-sporites Ibrahim (1933, p. 33). Knox (1939) has suggested that alete spores with somewhat spiral reticulation such as shown by R. facetus (cf. fig. 49) resemble spores of Riccia but this com- parison does not seem particularly close. Neither does the comparison of the R. corporeous type with Fossombronia spores seem appropriate because the fossil forms evidently have a perisporal membrane en- closing them instead of mere "large alve- olae with high lamellae." ISTevertheless it must be admitted that in general spores of Reticulati-sporites bear more resemblance to some of the spores of the Hepaticae il- lustrated by Knox than any of the other Paleozoic groups. The apparent absence of haptotypic markings on several of the modern forms is a striking feature which should be more thoroughly investigated, although it need not occasion great sur- prise to find so advanced a character in a group whose discrete phylogeny goes back at least to the Carboniferous. It may be suggested that spores of Reticulati-sporites underwent more mod- ification within the sporangium subsequent to the breakup of the original tetrad than is usual for cryptogamic spores. The highly developed reticulation pattern prob- ably was laid down relatively late in sporo- genesis by a tapetal plasmodium, some- times without respect for the proximo- distal relationships established in original tetrad groupings. Such a sequence occurs in formation of many kinds of angi- ospermic pollen. Spores of Reticulati- sporites apparently lack the further spe- cialization of germinal pores, neverthe- less lack of good development of a trilete commissure in some of the forms might possibly signify that the plants repre- sented here had passed beyond the typi- cal cryptogamic stage of evolution. Re- ticulati-sporites appears to be a relatively homogeneous group of natural relation- ship. The coarse reticulation is highly characteristic and for the most part quite unmistakable. Nevertheless these forms are rather anomalous in their morphology and offer difficulties in interpretation. Those suggested are regarded as tenta- tive only. The genus is best known from the Aegir coal bed at the top of the Westphalian B, which is approximately equivalent in age to middle Des Moines and upper Potts- ville-lower Allegheny beds. 1. Reticulati-sporites corporeus Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 155, pl. 7, fig. 7. 2. Reticulati-sporites facetus (Ibrahim) S. W. and B., comb. nov. Reticulata-sporites facetus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 38, pl. 5, fig. 36. 3. Reticulati-sporites mediareticulatus Ib- rahim, 1933, idem, p. 34, pl. 7, fig. 62. 4. Reticulati-sporites ornatus (Ibrahim) Ibrahim, 1933, idem, p. 35, pl. 1, fig. 7. Sporonites ornatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pl. 14, fig. 7. 5. Reticulati-sporites polygonalis (Ibra- him) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 155, pl. 7, fig. 16. Sporonites polygonalis Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pl. 14, fig. 8. Laevigati-sporites polygonalis (Ibra- him) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 19, pl. 1, fig. 8. 6. Reticulati-sporites reticulatus (Ibra- him) Ibrahim, 1933, idem, pp. 33-34, pl. 1, fig. 3. Sporonites reticulatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pl. 14, fig. 3. Reticulati-sporites reticulatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 156. Note. — Type species of Reticulati-sporites. 7. Reticulati-sporites (?) reticuliformis Ibrahim, 1933, Sporenformen des Aegirhori- zonts, p. 34, pl. 7, fig. 62). Note. — Trilete rays are unusually long and well developed in this species. 36 PALEOZOIC FOSSIL SPORES Nomina excludende 1. Reticulati-sporites angulatus Ibrahim, 1933, Spo- renformen des Aegirhorizonts, etc., p. 35, pi. 7, fig. 59. 2. Reticulata-sporites bireticulatus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 158, pi. 7. fig. 28. Sporonites bireticulatus Ibrahim, 1932, Neues Jahrb.. Beilage-Band 61, Abt. B. p. 447, pi. 14, fig. 1. Reticulati-sporites bireticulatus (Ibrahim) Ibra- him, 1933, Sporenformen des Aegirhorizonts, etc., p. 35, pi. 1. fig. 1. 3. Reticulati-sporites facierugosus Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 155, pi. 7, fig. 26. 4. Reticulata-sporites mediapudens Loose, 1934, idem, p. 158, pi. 7. fig. 8. 5. RbTICULATI - SPORITES MICRORETICULATUS (Loose) Loose, 1934. idem, p. 155. Sporonites microreticulatus Loose, 1932, Neues Jahrb.. Beilage-Band 67, Abt. B, p. 450, fig. 37. 6. Reticulati-sporites morosus Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 154, pi. 7, fig. 2. 7. Reticulati-sporites nexus (Loose), Ibrahim, 1933, Sporenformen des Aegirhorizonts, etc., p. 34. Sporonites nexus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, fig. 35. Reticulati-sporites nexus (Loose) Ibrahim, Loose, 1934. Inst. Palaobot. Arb., vol. 4, no. 3, p. 156, pi. 18, fig. 35. 8. Reticulati-sporites sifati Ibrahim, 1933, Sporen- formen des Aegirhorizonts, etc., p. 35, pi. 8, fig. 67, 9. Reticulata-sporites spongiosus Ibrahim, 1933. idem, p. 39, pi. 8, fig. 71. 10. Reticulati-sporites vinculatus (Ibrahim). Loose, 1934, Inst. Palaobot. Arb.. vol. 4, no. 3, p. 156. Sporonites vinculatus Ibraham, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448. pi. 15. fig- 19. Reticulata-sporites vinculatus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegirhorizonts, etc., p. 39, pi. 2, fig. 19. Genus Laevigato-sporites (Ibrahim, 1933) emend., S. W. and B. Plate 1, figures 5-5b Symmetry. — Spores bilateral, mono- lete. Shape. — Originally broadly bean- shaped ; elongate oval in the plane of lon- gitudinal symmetry, round or oval in the transverse plane. When compressed the spores tend to be folded variously depend- ing on the size and morphology of the various specific types of spores. Size. — Spores of various species range from about 20 to over 130 microns in their long dimension. Ornamentation. — Smooth to finely punc- tate, apiculate or rugose ; rarely showing pronounced reticulation or strong apicu- lae. Haptotypic structures. — Consisting of a simple monolete linear suture, generally without lips specially distinguished, and usually continued for more than half the total length of the spore. The suture may be very inconspicuous if it coincides in position with the edge of a compressed spore, or with the axis of a longitudinal fold. Ends of very delicate arcuate ridges are sometimes distinguishable flaring lat- erally from both ends of the suture line. Spore coat. — Varying somewhat in thickness relative to the other dimen- sions; the spore coat is oftentimes thin and translucent. The internal cavity is sometimes more *'bean-shaped" than the external outline due to internal thickening of the spore coat in the central proximal region. Affinity. — Smaller species of Laeviga- to-sporites agree with some forms ob- tained from filicinean type fructifications. There is now no particularly good evi- dence as to the affinity of the larger forms. Fredda Reed (1938, p. 333) has obtained spores of this type from an extraordinary new type of Calamarian fructification which are approximately 50 microns in length. Spores of Zeilleria are of this type (Florin, 1937, p. 316-7) but it is not now possible to decide whether that genus belongs to the ferns or pteridosperms (Halle, 1933, p. 88). The "distal fur- row" of Zeilleria spores remarked by Florin (1937, p. 317) appears no more definite than many of the fortuitous folds seen in spores of Laevigato-sporites. It thus appears that species of Laevi- gato-sporites could conceivably pertain to at least three distinct contemporaneous orders of Paleozoic plants. It seems doubtful that our information on these re- lationships will be greatly improved in the immediate future and until it is pos- sible to revise the genus on the basis of reliable evidence it will serve a useful geological, if not a botanical, purpose. In a good many instances species of Laevi- gato-sporites form a majority of the spores obtained from certain coal beds. They are, however, rare or absent in a number of lower Pottsville coals that have been examined. Spores evidently belonging to Laevi- gato-sporites have been assigned to types B,a, B,b, and Bg by Raistrick (1933, 1937) (Knox, 1938). Millott's types 6 and E^a (Millott 1939) and Knox' (1942) Type 6K may also belong here. Remarks. — Nine species, one of which includes three designated formae, are now listed for Laevigato-sporites. Ibrahim originally assigned two species to the genus. The second of these, L. ellipsoides, is now referred to Monoletes. It is pos- LAEVIGATO-SPORITES 37 sible that some of the names given below will prove to be synonyms. On the other hand a number of distinct and as yet un- recorded species have been recognized in the Illinois basin and elsewhere. The list includes genotype species of four previously named genera. L. bila- teralis (Loose) was the only species de- scribed by Loose (1934, p. 159) under the new name, Reticulato-sporites. The name was credited to 'Tbrahim 1932" al- though no "reticulata" was ever used by that author. L. desmoinensis is the gen- otype of Phaseolites Wilson and Coe (1940). L. minutus (Ibrahim) was the sole species given under Punctato-sporites by Ibrahim (1933, p. 40). L. vulgaris Ibrahim is the designated genotype of Laevigato-sporites Ibrahim (1933, p. 39). All of these species are evidently con- generic, their chief distinction being in variation of surface ornament and small differences of size, none of which can be regarded as of more than specific impor- tance. The names Reticulato-sporites, Phaseolites, and Punctato-sporites are therefore regarded as synonyms of Laevi- gato-sporites. All these are typically rep- resented by specimens from the upper part of the Westphalian B and approxi- mately equivalent beds in America. 1. Laevigato-sporites bilateralis (Loose), S. W. and B., comb. nov. Reticulato-sporites bilateralis Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 159, pi. 7, fig. 22. Note. — Reticulato-sporites is a generic name published only by Loose (1934) although he erroneously credited it to "Ibrahim, 1932." 2. Laevigato-sporites cranmorensis Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, p. 157, fig. 1. 3. Laevigato-sporites desmoinensis (Wilson and Coe) S. W. and B., comb. nov. Phaseolites desmoinensis Wilson and Coe, 1940, idem, vol. 23, no. 1, p. 183, fig. 4. Note. — Evidently closely related to Laevigato- sporites vulgaris. 4. Laevigato-sporites minimus (Wilson and Coe) S. W. and B., comb. nov. Phaseolites minimus Wilson and Coe, 1940, idem, vol. 23, no. 1, p. 183, fig. 5. 5. Laevigato-sporites minutus (Ibrahim) S. W. and B. comb. nov. Punctato-sporites minutus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 40, pi. 5, fig. Z2>. 6. Laevigato-sporites (?) pennovalis Berry, 1937, Am. Midland NaturaHst, vol. 18, no. 1, pp. 156-7, fig. 4. 7. Laevigato-sporites thiessenii Kosanke, 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 125, pi. 3, figs. 1, la, lb. Pittsburgh microspore of Thiessen, various publications. See especially Thiessen and Stand, 1923. 8. Laevigato-sporites (?) tuberculatus (Berry), S. W. and B., comb. nov. Tuberculati-sporites tuberculatus Berry, 1937, idem, vol. 18, no. 1, p. 155, fig. 9. 9. Laevigato-sporites vulgaris (Ibrahim) Ibrahim, 1933, Sporenformen des Aegirhor- izonts, pp. 39-40, pi. 2, fig. 16 ; pi. 5, figs. 2>7, 38, 39. Sporonites vulgaris Ibrahim, 1932, Neues Jahrb. Beilage-Band 67, Abt. B, p. 448, pi. 15, fig. 16. Laevigato-sporites vulgaris forma minor Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 158, pi. 7, fig. 12. Laevigato-sporites vulgaris forma maior Loose, 1934, idem, p. 158, pi. 7, fig. 6. Laevigato-sporites vulgaris forma max- iMus Loose, 1934, idem, p. 158, pi. 7, fig. 11. NOMEN EXCLUDENDE 1. Laevigato-sporites rheaensis Berry, 1937, Am. Mid- land Naturalist, vol. 18, no. 1, p. 157, fig. 13. Note. — The morphology of this form is so obscure that its significance cannot be ascertained. Genus Zonalo-sporites Ibrahim, 1933 Symmetry. — Spores apparently bilat- eral, monolete. Shape. — Flattened elliptical ( ?), broad- ly elliptical in transverse plane. 6^i>^.— Relatively large, 200-300 mi- crons in length. Ornamentation. — Surface smooth to minutely rugose. Haptotypic features. — Linear proximal suture with slight angular median deflec- tion, termini of arcuate ridges may be weakly differentiated. Spore coat. — Perisporal bladder mem- brane thin, enveloping spore body on all sides, and in contact with it proximally and distally. Body wall (exospore) rela- tively thick and brownish translucent. Affinity. — Probably pteridospermic, judging from points of similarity with Monoletes which is discussed later. It nevertheless is still extremely problematic since it superficially seems to combine features of spores of Medullosan and of Cordaitean relationship. 38 PALEOZOIC FOSSIL SPORES Rcuiar'ks. — This genus is reported in only two instances, in the Ruhr and in IlHnois, in both cases from relatively few specimens. The forms have been assigned to different species as listed below. Much more information is needed before their significance can be understood. 1. ZONALO-SPORITES AUREOLUS (Schopf) S. W. and B., comb. nov. Monoletes aureobis Schopf, 1938, Illi- nois Geol. Survey Rept. Inv. 50, pp. 45-46, pi. 1, figs. 1-2. 2. ZoNALO-sPORiTES viTTATUs Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 41, pi. 6, fig. 45. Genus Monoletes (Ibrahim) 1933, emend, S. W. and B. Plate 2, figures 17-17c Symmetry. — Spores apparently bilat- eral, monolete. Mature forms evidently have been modified considerably in post tetrad maturation, and it is possible that although the mature forms appear very definitely bilateral, they may have arisen from tetrahedral tetrads. Shape. — Elliptical to rounded lenticu- lar in outline as viewed from the proxi- mal side; of similar figure but of more slender proportions in the plane of the long axis viewed from the side ; the short transverse axis shows a somewhat rounder and shorter but notably flattened outline. The distal surface oftentimes is marked by two prominent grooves extending near- ly the length of the spore with a well rounded umbo between. Sometimes the proximal side near the suture shows a very slight prominence. On compression, longitudinal folds frequently parallel the distal grooves. Si^e. — Spores relatively large, varying from slightly over 100 microns to as much as half a millimeter in length. Ornamentation. — Surfaces generally minutely granulose appearing quite smooth at low magnification; sometimes glistening in reflected light. Emphytic marking is evenly distributed except at the extreme base of distal grooves, which are more smooth. Haptotypic features. — ^When closed the proximal suture is a very narrow linear groove, generally lacking marginal dis- tinction, though the lip area may be very slightly upraised. The most noteworthy and diagnostic feature is the slight angu- lar deflection of the suture fine near the middle. Although inconspicuous and sometimes hardly evident it seems to be universally present. It may constitute the sole remnant of a third vestigial ray in a fundamentally trilete suture pattern. Very weakly developed termini of arcuate rid- ges are occasionally resolved at the two ends of the functional suture line. The suture opens to form a sharp taper- pointed lenticular slit. The distal grooves have no connection with haptotypic struc- tures. Spore coat. — The exospore consists of a layer of varying thickness ; the proxi- mal side and the center of the distal umbo are as much as 15 or 18 microns; the thinnest is at the base of the distal grooves on either side of the umbo where it may be less than 5 microns. An even gradation exists between these extreme areas and in normally compressed spores the exospore thickness at the margin is intermediate between the extremes. The endosporal membrane is frequently evi- dent as a crumpled translucent sack less than a micron in thickness, sometimes shrunken from the exospore at the spore margin. The umbo and distal grooves, when present, have the appearance of functioning as a harmomegathus. Affinity. — Spores of this character dis- tinguish a pteridospermic plant alliance probably largely coextensive with the Me- duUosaceae. They are characteristic of all genera placed in the Whittleseyinean sub-tribe (see Halle, 1933). The only aberrant feature is the lack of distal grooves in spores of Codonotheca; spores of all other genera now assigned to this alliance possess them. Codonotheca spores are otherwise entirely characteristic of Monoletes and when such spores are found isolated they probably should be assigned to Monoletes since there well may be other types in addition to Codon- otheca that also may lack distal grooves and umbo. Spores of Codonotheca empha- size the non-essential character of these specialized distal structures. Remarks. — Much of the previous dis- cussion is based on comparative study of spores in Dolerotheca and Codonotheca by Schopf from petrified and compression material as well as from spores isolated by maceration of coal. It has thus been DENSO-SPORITES 39 possible to establish accurately from sec- tions the proximal position of the suture and the distal location of the harmome- gathic grooves. The fact that spores are frequently found with the suture opened but, except in instances of obvious me- chanical or chemical injury, have never been observed with the umbo split off as an "operculum" or the grooves other than intact, strongly suggests that proximal gametophytic exit was still a fundamental feature of these forms as it must have been in their more ancient cryptogamic an- cestry. Such spores are also regarded as prepollen (cf. Schopf, 1938, pp. 14-15). The foregoing diagnosis serves to cor- rect some erroneous interpretations of structure previously held; e.g. the "en- circling ridge" described by Schopf (1938, p. 45) for the species designated as Mono- letes ovatiis, is the margin of the distal grooves discussed here. It has no hapto- typic significance although it possibly is not emphytic in origin in the same sense that ordinary surface ornamentation is. Monoletes was first used by Ibrahim in 1933 to designate a new group in his ar- tificial system. Although substantive in form and similar to a generic name, it was not originally applied in that sense and no nomenclatural type has been pre- viously proposed. It has been used as a ,eeneric designation and applied more nar- rowly (Schopf, 1936, 1938) ; we now emend it to correspond with the restricted usage. Monoletes ovatus Schopf may serve as the type species of the genus. The group now includes two species, both of which appear to be widely dis- tributed. It is likely that further species will be distinguished by application of more refined biometric procedure. Both species must still be regarded as rather generalized types. 1. Monoletes ellipsoides (Ibrahim) Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 45, pi. 1, fig. 14; pi. 6, figs. 5 and 6. Sporonites ellipsoides Ibrahim, 1932, Neues Jahrb. Beilage-Band 67, Abt. B., p. 449, pi. 17, fig. 29. Laevigato-sporites ellipsoides (Ibra- him) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 40, pi. 4, fig. '29. Punctato-sporites ellipsoides (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, pp. 158-9, pi. 7, fig. 35. Sporites ellipsoides (Ibrahim) Wicher, 1934, idem, vol. 4, no. 4, p. 185. 2. Monoletes ovatus Schopf, 1935, Illinois Acad. Sci. Trans, vol. 28, no. 2, p. 108, fig. 7. Pollen of Dolerophyllum sp., Zerndt, 1930, Acad, polonaise sci. Bull, internal., ser. B, pp. 55-56, pi. 8, figs. 42-49. Type 31 Zerndt, 1931, idem, ser. A, p. 176. Monoletes ovatus Schopf, 1935, Illinois Acad. Sci. Trans, vol. 28, no. 2, p. 176. Monoletes ovatus Schopf, 1938, Illinois Geol. Survey Rept. Inv. No. 50, pp. 43-45, pi. 1, figs. 3-5, pi. 6, figs. 1-4. Pollen of Dolerotheca fertilis, Zerndt, 1940, Paleontographica, vol. 84, Abt. B, p. 136-138, pi. 11, fig. 36. Genus Denso-sporites (Berry, 1937), emend., S. W. and B. Plate 1, figures 9-9c Symmetry. — Spores radial, trilete. Shape. — Originally oblate, round to subtriangular in equatorial outline. When compressed, the central area becomes much thinner than the margins due to va- riation in spore wall thickness. Coal thin sections, taken vertically through the coal bed, show them compressed into a "dumb- bell" shape. The original highly oval sec- tional form results in extremely uniform proximo-distal orientation in enclosing sediments. Sise. — Diameter of specimens assigned among various species varies from about 35 to 100 microns. Ornamentation. — Smooth to apiculate and rugose. External surface character, typically is uniform, though ornamenta- tion may be more strongly developed to- ward the equator in some forms. Haptotypic structures. — A delicate tri- lete marking is visible in well preserved material or where the central area of the proximal surface is consistently present. "Fissures" penetrating to the margin in some instances may be essentially hapto- typic continuations of the trilete sutures. Spore coat. — Characterized by great differences in thickness. The proximal and distal walls are usually membranous, or at least significantly thinner than the equatorial portion of the coat. The latter area is oftentimes so thick it is practically opaque to transmitted light in contrast to the highly translucent central area. It may also be so extended centrifugally as to simulate a flange but it is not usually strictly demarcated from the spore body, as such. 40 PALEOZOIC FOSSIL SPORES Affinities. — Spores of this genus have not been recognized in connection with fructifications. This is surprising in view of their frequence in coal. They have been roughly designated as "biconcave," "salvershaped" and as "splint micro- spores" by Thiessen and his co-workers (Thiessen and Wilson, p. 9, 1924; Thies- sen, 1930; Sprunk, et al, 1940) and can be recognized by their characteristic "dumb- bell" shape in cross section.^ Modern methods of study may be ex- pected therefore to disclose definite evi- dence of the relationship of this group. Their structure is so unusual that it seems evident that a single homogeneous group of plants for the most part is represented. Raistrick's Type A spores (Raistrick and Simpson, 1933) apparently are nearly entirely referable to Denso-sporites. His A„ Ae, (1934) and A,o (1937, 1938) also are probably congeneric and, if so, appear to be the most bizarre of any in the genus. Possibly the A3 spores (Rai- strick, 1937; Knox, 1938) are least as- suredly related to this group. Knox's Type Cg (Knox, 1942) possibly might be placed here though its morphologic interpreta- tion is somewhat uncertain due to its un- usual type of equatorial development. Remarks. — Six species are now included under Denso-sporites and it seems that a good many more will need be distin- guished. Raistrick's A5, Ag, and A^q types are worthy of specific distinction. The generic name, while perhaps of un- desirable construction, is not in conflict with any others that have been proposed and thus is entirely valid. As mentioned previously, the spores are widespread and very abundant in splint coals. It has been suggested that a special type of vegetation was respon- sible for this type of coal but such can hardly be the case, since recognizable splint coals are also found that are so much younger that floral similarities cease (cf. Thiessen and Sprunk, 1936). White (in discussion of Thiessen, 1930, p. 672) doubts that the presence of a certain type of plants is responsible for forming this type of coal. The coincidence of occur- rence between Denso-sporites and Pale- ozoic splint coal nevertheless seems unaccountably high and additional infor- 8 Thiessen has also designated some spores not referable to Denso-sporites as "splint spores." mation about these spores may reflect importantly, although indirectly, upon the problem of splint coal formation. 1. Denso-sporites annulatus (Loose) S. W. and B., comb. nov. Sporonites annulatus Loose, 1932, Neues Jahrb. Beilage-Band 67, Abt. B, p. 451, pi. 18, fig. 44. Zonales - sporites annulatus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 151.10 2. Denso-sporites covensis Berry, 1937, Am. Midland Naturalist, vol. 18, p. 157, fig. 11. Note. — Although serving as genotype, the spe- cific characteristics of this form are inadequately known. 3. Denso-sporites densus Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, pp. 157-8, fig. 7. 4. Denso - sporites (?) indignabundus (Loose) S. W. and B., comb. nov. Sporonites indignabundus Loose, 1932, Neues Jahrb. Beilage-Band 67, Abt. B, p. 451, pi. 19, fig. 51. Zonalcs-sporites indignabundus (Loose) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 32. Apiculati-sporites indigna bund us (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 153. 5. Denso-sporites loricatus (Loose) S. W. and B., comb. nov. Sporonites loricatus Loose, 1932, Neues Jahrb. Beilage-Band 67, Abt. B, p. 450, pi. 18, fig. 42. Zonales-sporites loricatus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 151. 6. Denso-sporites (?) quadratus Berry, 1937, Am. Midland Naturalist, vol. 18, p. 158, fig. 10. Note. — A very problematic form. Genus Cystosporites Schopf, 1938 Plate 1, figures 10-lOb Symmetry. — Spores radial, trilete. Form and size. — Variable depending on development. Fertile members relatively enormous, sack-like, elongate, sometimes attaining a length of more than a centi- meter and about half as broad. Abortive forms variable in development, charac- teristically ranging from as small as one- 10 Since Loose recognized the rather significant distinc- tion between Denso-sporites and the zonate forms, in a footnote appended to his article he suggested "einer Unter- abteilung Annulati-sporites" for D. annulatus and D, loricatus. Because he did not adopt the name, "Unter- abteilung Annulati-sporites" is invalidly published, at least in the sense of a generic designation. CYSTOSPORITES 41 half a millimeter in diameter to more than twice that. Smallest forms are more nearly isodiametric becoming axially elon- gate with increased size. When compressed both abortive and fertile members are folded irregularly ; their longest axis is generally least effected. Ornamentation, — Generally absent; abortive forms may show apparent re- ticulation. Haptotypic features. — Strongly devel- oped in fertile forms but relatively in- conspicuous because of the enormous ex- pansion of the distal part of the spore body. Sutures definite, extended to the arcuate ridges which are strongly thick- ened, lips often moderately defined; py- ramic areas distinct from adjacent spore coat. Apex may be somewhat elongate. On smaller abortive forms haptotypic features are commonly masked by develop- ment of a thick rugose, more or less triangular cushion which effectively seals the spore apex. Arcuate ridges are in- conspicuous but sometimes their termini are weakly defined adjoining the corners of the cushion. Larger abortive forms commonly show haptotypic features more like fertile spores but somewhat less devel- oped and more irregular in character. Spore coat. — Variable in thickness, generally relatively thick on abortive and considerably thinner, especially in the median areas, on fertile spores. Pyramic areas of fertile forms are thinner but the more proximal portion immediately distal to the arcuate ridges is thick ; the median portion of the spore is membranous and the extreme distal portion may again be somewhat thicker. The most characteristic feature of this genus is the fibrous char- acter of the spore coat in the membra- nous middle region ; thicker parts of the coat also consist of thicker matting of the interlocking, variable width, anastomos- ing fibrils. A thinner nonfibrous endo- sporal membrane may be present. The fibrous spore coat is evidently equivalent to the exospore. Affinites. — Cystosporites is a member of the Lepidocarpaceae. It is intimately related to the genera Lepidocarpon, Illi- niocarpon, and probably others of this family in which the correlation has not been as definitely established (Schopf, 1941). Spores of Cystosporites, although highly specialized with reference to semi- nal adaptation, appear to be generalized so far as specific differences are con- cerned. The ornamental patterns which serve to distinguish spores of free-spor- ing lycopsids have evidently undergone extensive reduction if such characteristics were ever present in the line of lepido- carp ancestry. The genus Cystosporites thus expresses a broader relationship than that exemplified by other genera in the Lepidocarpaceae, and, so far as is known, is inclusively correlative with the lepidocarp family. Future discoveries may make it possible to restrict this group and establish relationships in greater detail. The spores found within the seeds of lepidocarp species are receiving more de- tailed study now (Reed, 1941 ; Darrah, 1941 ; Hoskins and Cross, 1941 ; Schopf, 1941) and further studies may show other features of the spore coat that have unsuspected systematic significance. Zerndt (1930, et seq.) has classified some forms now placed in Cystosporites, under Triletes. The distinction between free-sporing and seed-bearing lycopsids seems at least familial in its importance (Schopf, 1941) and this distinction is most in evidence when the megaspores are compared. Remarks. — The fertile and abortive megaspores of species of Cystosporites are very heteromorphous. In several instances they have been recovered in tetrad group- ings from maceration residues of coal. They nevertheless are generally separated from one another so that a system of treatment for the isolated tetrad members both fertile and abortive seems essential for scientific reporting. The characters of the fertile spores are used to identify species, and fertile specimens serve as holotypes for the two species listed below. Isolated abortive spores may be distin- guished in classification as formae. The procedure may serve as a practical means of expressing relationships between these heteromorphous spore types and still re- tain for each of them a nominal distinc- tion. There appear to be no inherent dis- advantages in such a policy, as it affords a means of more accurate recording of actual material encountered ; only the ac- crual of additional information can show whether such a practice, in fact, is neces- sary. If the formae later appear super- fluous in some instances and worthy of 42 PALEOZOIC FOSSIL SPORES actual specific status in others, these changes can be effected without undue taxonomic difficulty or confusion since type material is indicated in all instances. Cystosporites has an extensive range in the Carboniferous as is best shown by tables of spore distribution given by Zerndt (1937), and so far as is now known is restricted to this period. The type species was described from Illinois No. 6 (Herrin) coal. 1. Cystosporites breretonensis Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, pp. 40-42, pi. 3, fig. 5, pi. 8, figs. 1-4. Triletes cf . T. giganteus Zerndt, Schopf, 1936, Illinois Acad. Sci. Trans, vol. 28, no. 2, p. 107, fig. 5 (holotype of C. breretonensis) . Triletes cf. T. giganteus Zerndt, Schopf. 1936, idem. p. 175. Cystosporites breretonensis forma abor- Tivus Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 40, pi. 1, fig. 10, pi. 8, fig. 4. Cystosporites breretonensis forma retic- ULATUS Schopf, 1938, idem, p. 40, pi. 1, fig. 11. 2. Cystosporites giganteus (Zerndt) Schopf, 1938, idem, p. 39. Triletes giganteus Zerndt, 1930, Acad, polonaise sci. Bull, internal., ser. B, p. 71, pis. 9, 10, and 11. Type 1 Zerndt, 1931, Acad, polonaise sci. Bull, internal., ser. A, p. 170. Sporites giganteus (Zerndt) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 172, pi. 8, fig. 9. Triletes giganteus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, p. 4. Cystosporites giganteus forma varius (Wicher) S. W. and B., comb. nov. Triletes (Lagenicula) glabratus Zerndt. 1930, Acad, polonaise sci. Bull, internal., ser. B., p. 54, pi. 8, figs. 38-41. (?) Type 29 Zerndt, 1931, idem, ser. A., p. 175. Type 30 Zerndt, 1931, idem, ser. A., p. 175, pi. 8, figs. 26-27. Type 30 Zerndt, 1932, Jahrb. fur das Berg-u. Huttenwesen in Sachsen. Tahrg. 1932, p. 13A, pi. 1, figs. 1, 2, 3. Sporites varius Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 89, pi. 6, figs. 2, 3, 4, 6. Genus Parasporites Schopf, 1938 Plate 2, figures 16-16b Symmetry. — Prepollen grains appear- ing bilateral due to opposing bladders. The fundamental symmetry, clearly shown by the body, is radial. Shape. — Body nearly spherical, blad- ders of moderate inflation placed laterally and opposite one another on the spore give an oval external outline. On com- pression few folds are formed and these generally do not modify the profile unless the plane of the bladders fail to coincide with the plane of compression. Size. — Relatively large; species attain as much as 300 microns length from one bladder tip to the other. Ornamentation. — Body wall may be rugose, bladder membrane lightly sculp- tured. Haptotypic features. — Relatively incon- spicuous and evidently modified in post- tetrad development. Trilete rays often developed with two long rays and one characteristically shorter. One pyramic area may sometimes be distinguished; except in this area no arcuate marking is evident. Spore coat. — Body wall (exospore) tending to be dense and of moderate rela- tive thickness ; bladder membrane (peri- spore), relatively thin and quite trans- lucent. Affinity. — There is considerable prob- ability that the plants represented by Para- sporites are gymnospermous. The peri- sporal bladder development is strongest evidence of this. It is still a question whether the genus should be assigned with the Pteridosperms, Cordaitaleans or the Conifers (cf. Schopf, 1938, pp. 47-8). The genus has some evident claim to affinity with Florinites, S. W. and B., Pityosporites Seward, Alisporites Daugh- erty, and to Endosporites Wilson and Coe. Perhaps it is closest to Endosporites, but as these forms are contemporaneous their relationship cannot easily be regarded as direct. Remarks. — The genus is monotypic being represented only by the species given below which has been found in upper Carbondale and lower McLeans- boro age coals of Illinois. The point upon which greatest emphasis should be placed is that the development of proximal sutures is such as to indicate proximal gametophytic exit similar to Endosporites and Monoletes. Aside from this and its rather large size Parasporites has pollen CIRRATRIRADITES 43 grain characteristics. It may represent a veritable "prepollen". 1. Parasporites maccabei Schopf, 1938, Illi- nois Geol. Survey Rept. Inv. 50, pp. 48-9, pi. 1, fig. 6, pi. 7, figs. 1-3. Genus Cirratriradites Wilson and Coe, 1940 Plate 3, figures 21, 21a, 21b Symmetry. — Spores trilete, radial. Shape. — Moderately oblate spheroidal, with a strongly projecting equatorial flange. The flange may assume a tri- angular horizontal outline due to emphasis of the trilete rays, or it may be nearly circular; the spore body is circular or slightly triangular. When compressed, few folds are evident and, because of the flange and oblate shape of the body, the plane of flattening nearly always cor- responds with that of the flange. Size. — Spores of various species range from about 40 to over 100 microns in overall diameter. Ornamentation. — Commonly showing a pattern having its primary emphasis along radial lines ; may consist of ridges more or less anastomosing to the point of be- coming a reticulation, or surfaces may be nearly free of ornamental ridges and most of the surface smooth, granulose or finely punctate. Some species show a unique type of distal ornamentation con- sisting of one or several thinner areas with rather prominent margins. The flange is often more or less radially striated and in addition may develop one or two concen- tric bands of irregular thickening. The pyramic areas are not particularly dis- tinguished by ornamentation. Haptotypic features. — Trilete rays rela- tively strongly developed and extended to the equator, oftentimes a line of thick- ening continues to the edge of the flange. Lips are frequently strongly demarcated and raised above the spore body ; the suture lines are attenuate but distinct. No arcuate ridges are developed distinct from the flange and, in fact, the flange may be taken to represent a hyper-development of these haptotypic formations. The flange is usually relatively broad, sometimes com- prising more than half the total spore diameter. It is definitely distinguished from the spore body, being thinner than the body wall and more translucent ; the interradial flange width is often some- what less than that opposite the rays. The margin of the flange is often minutely to rather coarsely serrate. Affinity. — Spores of Cirratriradites are not yet definitely correlated with any major plant group. Their most likely affinity seems to be with the lycopods but known types of Lepidostrobus microspores are usually smaller and generally possess no comparable development of the flange. Certain zonate types of Triletes spores bear a superficial resemblance to Cirratri- radites but so far as is known now there is no inherent correlation between struc- ture of megaspores and microspores where heterospory is as highly developed as it is in Triletes. There is in fact no essential evidence to show whether spores of Cirra- triradites functioned as microspores or isospores. The Carboniferous existence of isosporous lycopods is still most firmly supported only on theoretical grounds but they could possibly be represented by Cirratriradites. Some authors have confused Cirratri- radites spores with those of genera {En- dosporites, Spencerites, etc.) which pos- sess perisporal bladder development in the equatorial plane. Such resemblance as there is, is certainly only superficial. Raistrick's A^, A^, and Cg (cf. Rai- strick 1935, 1937, 1938) belong here; likewise, possibly D2 in part. As given by Knox, (1938, 1942) D,^ is definitely referable to Cirratriradites. Types 1 and 9 of Millott (1939) without much ques- tion belong in this genus. The spores illus- trated by Reinsch (1884, p. 22) in plate 15, figs, la and lb as types 222 and 223 from Zwickau, Saxony, are characteristic of it. Remarks. — Thirteen previously de- scribed species are included in the list be- low, all but one of which, C. maculatus the genotype, represent new name com- binations. Spores of this character are frequently encountered in American coals and addi- tional species will subsequently be de- scribed. The group shows such general agreement in numerous characteristics that a considerable degree of natural relationship is attributed to it. 1. Cirratriradites argutus (Ibrahim) S. W. and B., comb, no v. Zonalcs-sporites argutus Ibrahim, 1933, Sporenformen des Aegirhorizonts, pp. 31-32, pi. 6, fig. 55. 44 PALEOZOIC FOSSIL SPORES 2. CiRRATRiRADiTES (?) cicATRicosus (Ibra- him) S. W. and B., comb. nov. Sporonites cicatricosiis Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pi. 14, fig. 2. Zonales-sporites cicatricosus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegirhor- izonts, p. 31, pi. 1, fig. 2. 3. CiRRATRiRADiTES FAUNus (Ibrahim) S. W. and B., comb. nov. Sporonites f annus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pi. 14, fig. 4. Zonales-sporites f annus (Ibrahim) Ib- rahim, 1933, Sporenformen des Aegirhori- zonts, p. 28, pi, 1, fig. 4. 4. CiRRATRiADiTES FORMOsus (Ibrahim) S. W. and B., comb. nov. Sporonites formosus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pi. 14, fig. 10. Zonales-sporites saturni (Ibrahim) Ibrahim, in part, 1933, Sporenformen des Aegirhorizonts, p. 30, pi. 1, fig. 10. 5. CiRRATRiRADiTES (?) GRACILIS (Zcrndt) S. W. and B., comb. nov. Triletes gracilis Zerndt, 1937, Acad. polonaise sci. Bull, internat., ser. A., p. 586, pi. 12, figs. 1-10. Type 46 Zerndt, 1937 (idem). Note — Species exceptionally large (300 ti) , otherwise in close generic agreement. 6. CIRRATRIRADITES MACULATUS WilsOU and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 183, fig. 7. Note. — Reexamination of the type material suggests the trilete apex is ordinarily normally developed and that the apical opening originally described is not of general occurrence. Sculp- turing simulating it may occur distally. 7. CiRRATRiRADiTES ( ?) PEACOCKi (Berry) S. W. and B., comb. nov. Zonales-sporites peacocki Berry, 1937, idem, vol. 18, p. 156, fig. 5. 8. CiRRATRiRADiTES PENNINGTONENSIS (Ber- ry) S. W. and B., comb. nov. Zonales-sporites penning tonensis Berry, 1937, idem, p. 156, fig. 3. 9. CIRRATRIRADITES RARus (Ibrahim) S. W. and B., comb. nov. Zonales-sporites rarus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 29, pi. 6, fig. 53. 10. CiRRATRiRADiTES SATURNI (Ibrahim) S. W. and B., Comb. nov. Sporonites saturni Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 448, pi. 15, fig. 14. Zonales-sporites saturni (Ibrahim) Ib- rahim, in part, 1933, Sporenformen des Aegirhorizonts, p. 30, pi. 2, fig. 14 (non pi. 1, fig. 10). Zonales-sporites saturni (Ibrahim) Ib- rahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149, pi. 7, fig. 23. A^of^.— Ibrahim (1933) came to the conclusion that Sporonites formosus Ibrahim (1932) was conspecific with C. saturni. Very little infor- mation is provided to support this and, since the figures appear to show specific differences, it appears desirable to keep them separate for a time at least. 11. CIRRATRIRADITES TENUIS (LoOSc) S. W. and B., comb. nov. Sporonites tenuis Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 450, pi. 18, fig. 34. Zonales-sporites tenuis (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no 3, p. 149. 12. CiRRATRiRADiTES VENUSTUS (LoOSC) S. W. and B., comb. nov. Sporonites venustus Loose, 1932 Neues Jahrb., Beilage-Band 67, Abt. B., p. 450, pi. 18, fig. 36. Zonales-sporites venustus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149. 13. CiRRATRiRADiTES ZONALIS (LoOSe) S. W. and B., comb. nov. Zonales-sporites sonalis Loose, 1934, idem, vol. 4, no. 3, p. 148, pi. 7, fig. 5. Genus Endosporites Wilson and Coe, 1940 Plate 2, figures 14-14b Symmetry. — Spores trilete, radial. Shape. — Moderately flattened elliptical in transverse plane, in axial plane round to oval or elliptical or slightly triangular ; body wall more spherical than the bladder. When compressed the spores most com- monly shov^ good proximo-distal orienta- tion. The spore body is not generally folded but the bladder membrane more commonly shows irregular plication. The bladder profile at the margin of the spore corresponds to the profile of folds, being straight or evenly rounded, and thus easily distinguishable from simple flanges of zonate forms that superficially re- semble Endosporites, Size. — Ranging in various species from about 50 to possibly 300 microns total diameter. The spore body generally is less than one-half of the full diameter. Ornamentation. — Surfaces of bladder are levigate to granular or punctate ; a fine meshed reticulation pattern is com- monly present but this is probably on the inside, rather than on the external bladder ENDOSPORITES 45 surface. The spore body is less definitely ornamented although the bladder pattern may appear superimposed on it. Emphytic ornament seems to be the same for proxi- mal and distal surfaces. Haptotypic features. — Trilete rays com- monly extend to the periphery of the spore body and thickened continuations from them may carry on to the bladder membrane. The lips are oftentimes up- raised and definite, the suture line distinct. Sometimes a thickening is present on the bladder membrane which corresponds to arcuate ridges. Spore coat. — The bladder (perisporal) membrane is quite thin and translucent ; its reticulation tends to add somewhat to the thickness as seen in marginal profile. The body wall is substantially thicker al- though also quite translucent, and cor- responds to the exosporal layer. In a few instances thin membranous, almost hyaline, endosporal membranes have been observed within the exospore and shrunk- en from it. The suture lines are not dis- tinct on these, but three clear-cut endo- sporal apical papillae occur at the juncture of pyramic apices. Affinities. — Endosporites is related to some of the Pennsylvanian Cordaitaleans. They correspond to spores observed by Wilson in male strobili, and Schopf has found well preserved specimens in such abundant association with Cordaitean leaves, other plant fossils being infrequent, that no other conclusion seems permis- sible. It should be emphasized, however, that the pollen grains Florin (1936) described in Cordaianthus fructifications obtained from the French Stephanian are evidently generically distinct and of more advanced structure. Raistrick's types C-^ and C4 without much question belong to Endosporites (Raistrick and Simpson, 1933; Knox, 1938). Type 629 of Reinsch (1884, p. 61, pi. 48, fig. 252A) from the Blatterkohle and Stigmarienkohle of Metschowk (in central Russia) and probably others less easy to interpret from his drawings, also belongs to Endosporites. Remarks. — The distended bladder mem- brane is the most characteristic feature of certain gymnospermous pollen grains. The expansion of the bladder is various in the several groups which are known as Parasporites, Cordaianthus, Alisporites, Caytonanthus, Pityosporites and Endo- sporites, as well as in modern podocarps and Abietineae. There is good reason to believe that bladders of this type are ho- mologous structures, which may have un- dergone progressive and regressive de- velopment at various times within the broad confines of this alliance, but nevertheless largely preserve their iden- tity throughout. Endosporites shows such evident de- velopment of proximal haptotypic struc- tures that gametophytic exit was quite evidently from the proximal pole as in the cryptogams. Florin, working with excellently preserved silicified material, has shown that in some late Pennsylvanian (Stephanian) forms the haptotypic fea- tures were vestigial and in mature forms present only as a surface imprint on the bladder membrane which does not even maintain contact with the spore body. The bladder and body are in contact dis- tally, however, and prothallial cells line the body cavity except at this point. Thus in these advanced forms germinal exit must have been distal as it is in pollen of all modern conifers. Pollen-like types of this sort having indication of proximal exit have been termed *'prepollen," fol- lowing Renault, and its significance has been discussed elsewhere (Schopf, 1938; pp. 14 and 15, 48). Endosporites male spores are evidently prepollen in this sense. Eight previously described species are listed below, only one of which is queried. The genus is widely distributed both in America and in Europe. The genotype species, E. ornatus Wilson and Coe, is from Iowa coal of Des Moines age. 1. Endosporites angulatus Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 184, fig. 1. 2. Endosporites globiformis (Ibrahim) S. W. and B., comb. nov. Sporonites globiformis Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pi. 14, fig. 5. Zonales-sporites globiformis (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 28, pi. 1, fig. 5. Zonales-sporites globiformis (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 148. 3. Endosporites (?) karczewskii (Zerndt) S. W. and B., comb. nov. 46 PALEOZOIC FOSSIL SPORES Trilctcs karcacwskii Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 1, p. 27, pi. 31, fig. 3. Note. — Agrees well with Endosporites though larger than most species. 4. Endosporites ornatus Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 184, fig. 2. Spore 6, Wilson and Brokaw, 1937, Iowa Acad. Sci., vol. 44, pp. 129-130, fig. 6. 5. Endosporites pellucidus Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 184, fig. 3. Spore 1, Wilson and Brokaw, 1937, Iowa Acad. Sci., vol. 44, pp. 128-129, fig. 1. 6. Endosporites rotundus (Ibrahim) S. W. and B., comb. nov. Zonales-sporites rottmdus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 31, pi. 8, fig. 73. 7. Endosporites rugatus (Ibrahim) S. W. and B., comb. nov. Zonales-sporites rugatus Ibrahim, 1933, idem, p. 31, pi. 8, fig. 70. 8. Endosporites volans (Loose) S. W. and B., comb. nov. Sporonites volans Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 451, pi. 18, fig. 46. Reticulati-sporites volans (Loose) Ib- rahim, 1933, Sporenformen des Aegirhori- zonts, p. 36. Zonales-sporites volans (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149. Note. — Trilete rays are shorter than is com- mon in this genus. Genus Triquitrites Wilson and Coe, 1940 Plate 3, figures 20-20b Symmetry. — Spores trilete ; radial. Form. — Oval to elliptical in vertical plane, distal side sometimes slightly more inflated than the proximal ; in the equato- rial plane triangular in outline, corners rounded or truncate subangular and some- times extended, sides slightly convex to strongly concave in profile. Folds are seldom induced by compression due to good preferential orientation coinciding with the horizontal plane of the spores ; forms more inflate distally are indicated through a tendency of the corners to be directed slightly upwards exaggerating the natural thickening of the spore coat on the angles. Sise. — Spores of various species com- monly range from 35 to 70 microns in mean diameter. Ornamentation. — Surfaces levigate to granulose and mildly verrucose, angular areas sometimes more highly ornamented. Emphytic ornamentation of proximal and distal surfaces is about the same. Haptotypic structures. — Trilete rays ex- tended nearly to the margin of the body cavity; lips sometimes rather thick and prominent, but usually not particularly demarcated. Arcuate ridges lacking ; like- wise, no flange in the usual sense is de- veloped though extreme extensions of the spore coat at the angles may simulate a partial flange. Spore coat. — Characterized by gross in- equalities in exospore thickness ; spore coat is thickest on the angles opposite ends of the rays and thinnest in the central distal and possibly to a somewhat lesser extent on the interradial areas. Actual thickness of difl^erent parts of the wall in various species varies greatly although the relations of thicker and thinner areas of the spore coat is rather constant. Affinity. ■ — - Triquitrites cannot be as- signed to any major plant group at the present time although the character of the spore coat seems so individualistic that the genus is considered to correspond with natural plant relationship. The features, in general, seem to agree best with spores of filicineans. Type 10 of Millott (1939) belongs to Triquitrites and probably type D^o of Rai- strick (1938). Type D, (Raistrick, 1934, 1935, 1938) shows more extreme thicken- ing of the radial angles of the spore coat so that it is evidently closely related to this genus, though it is a question whether it should be classified as Triquitrites or distinguished from it. Knox' Type 4K (1942) is clearly congeneric. Spores of the same sort but even more extreme have been illustrated by Reinsch (1884) in his plate 3, figures 34-42, corresponding to his type dia.s^noses, nos. 345, 347, 349, 353, 354, 362, 363, 365, 373. Reinsch also illustrated other forms which are doubt- less congeneric with Triquitrites in his tvpes 58, 200, 341, 342, 348, 355. etc. Type 58 shown on his plate 15, figure 18A, from Zwickau, Saxony, is remi- niscent of some, possibly overmacerated, specimens of the genotype species. Remarks. — Five named species are as- signed to Triquitrites ; three represent new name combinations. Other species are EQUISETOSPORITES 47 known from American coals and will sub- sequently be described. The numerous spores illustrated by Reinsch (1884) in the Micro-Paleophytologia show best the vari- ation in structure encountered is this group. The spores with highly exag- gerated angular thickenings were mostly derived from localities in Russia where the coals probably are of Lower Carbonif- erous age. Dio and D^ types of Raistrick (1938) were chiefly obtained from the Lower Carboniferous of England. Penn- sylvanian age species are much more moderate in spore coat thickness and the interradial and distal membranes are fre- quently less than 3 microns thick, as in the four species which have been de- scribed and named. The angles are more than twice as thick, however, and their relationship thus seems definitely indi- cated. Nevertheless, it may be desirable that the thicker walled more ornate and radially extended forms be generically segregated when detailed modern infor- mation becomes available for the older species. The tendency toward seeming simplification in spore coat structure of late Carboniferous plants as contrasted with ealier merpbers of the same alliance has also been noted in members of the Lagenicula and Aphanozonateae sections of Triletes. 1. Triquitrites arculatus Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 185, fig. 8. Note. — Type material bears a moderate to sparse verrucose flecking (areas of thickening) on proximal and distal surfaces, possibly due to overmaceration. 2. Triquitrites spinosus Kosanke, 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 128. pi. 3, figs. 2, 2a, 2b. 3. Triquitrites tribullatus (Ibrahim) S. W. and B., comb. nov. Sporonifes tribullatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 448, pi. 15, fig. 13. Laevigati-sporites tribullatus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegirhor- izonts, pp. 20-21, pi. 2, fig. 13. Valvisi-sporiies tribullatus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 152, pi. 7, fig. 21. 4. Triquitrites trigonappendix (Loose) S. W. and B., comb. nov. Valvisi-sporites trigonappendix Loose, 1934, idem, p. 152, pi. 7, fig. 17. 5. Triquitrites triturgidus (Loose) S. W. and B., comb. nov. Sporonites triturgidus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pi. 18, fig. 32. Valvisi-sporites triturgidus {Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 151. Genus Equisetosporites Daugherty, 1941 This genus recently described from Tri- assic age beds in southwestern United States is mentioned here because of its important bearing on recognition of fossil spores of certain Equisetalean plants. The genus is monotypic, including only E. chinleana as given below, whose note- worthy feature is the presence of elaters very similar to those characterizing the modern genus. The vahdity of the name Equisetosporites rests on the belief that these forms should be distinguished from spores of Equisetum. It appears there is adequate support for this view since Daugherty, in a personal communication, states "in all cases Equisetum-like. plants of the Triassic have proven to be quite dif- ferent when the record is complete enough to allow specific determination." Thus until further evidence is available, it would seem unwise to emphasize the mod- ern aspect of this fossil form. Its dis- covery nevertheless adds an important item of information to the record. Knox examined spores of fifteen species of Equisetum and reports (1938, pp. 439- 40) them to be invariably spherical, 30 microns to 35 microns in diameter, and always thin walled. In all these species the spore wall was faintly granular. Ela- ters of somewhat varied character are present on all modern species of Equi- setum. In commenting on Raistrick's Bg type, which she recognizes is very prob- ably of calamarian affinity, Knox remarks (pp. 461-3) on the same globose form with thin wall and absence of ornamentation . . . the unusual de- velopment of elaters and no trace of the tri- radiate mark. She cites Halle's description of Rhaetic and Triassic - age Equisetuni spores (Halle, 1908) in which the trilete com- missure is clearly seen but which show no trace of elaters. Halle assumed that even if elaters had been present they would have been removed by maceration procedure. Needless to say, this aspect 48 PALEOZOIC FOSSIL SPORES must always be considered. Nevertheless, the spores recovered by Halle were typi- cal of calamarians (cf. Hartung, 1933) and isolated spores of this character must be classed with Calamospora n. gen. as de- scribed below. Eqiiisetosporites chinleana is quite different and marks the advent of a more distinctly modern character in this ancient and persistent order of plants. Evidently a reduction in importance of haptotypic features has occurred and con- currently the perisporal adaptation repre- sented by elaters has come into existence. Careful study of the fossil spores may be expected to indicate more completely the relationship and distribution of equiseta- lean types. The type of the single species of Equi-, setosporites now known has been made available to us for examination through the kindness of Prof. Ralph W. Chaney and the following description, which is somewhat more detailed than Daugher- ty's, is offered for comparison with the other descriptions' given in this paper. This specimen (No. 1562 of the Califor- nia Museum of Paleontology) is ade- quately illustrated by Daugherty in the figure cited below. Symmetry. — No trace of original tet- rad configuration visible. Shape. — Body spherical (type speci- men elliptical, 30 x 37}^ microns, due to one elongate taper-point fold). Ornamentation. — Body wall essentially smooth showing very slight surface un- dulation. Spore coat. — Thin, apparently slightly less than a micron in thickness. In spite of this the body is deep brown color as viewed by transmitted light. Elaters. — Probably four in number, having their attachment close together; each one possibly about 70 microns long, band-like, about 4 microns wide, with slight irregularities; yellowish translu- cent. Elater ends truncated, tapering for a short distance, not at all broadly ter- minated. (Two of the four free ends are shown at the right edge and over- lapping the spore body in Daugherty's fig. 4.) Evidently the elaters tended to spirally encircle the spore body but they have been displaced on the specimen at hand. 1. Equisetosporites chinleana Daugherty, 1941, Carnegie Inst. Washington Pub. 526, p. 63, pi. 34, fig. 4. Age, Triassic. Genus Alisporites Daugherty, 1941 Plate 2, figure 12 This genus, like the preceding, is mono- typic, the sole species yet described being of Triassic age from Southwestern United States. It is included in this paper be- cause spores of Permain age which are probably referable to it have been found, and because there seems some basis for re- lating it to Parasporites and Pityosporites. The only description is that of the geno- type species, A. opii, given below (Dau- gherty 1941, p. 98). "The spores are large, averaging 100 to 110 microns in length and having two large mem- branaceous wings with reticulate markings. They are spherical to ovate in dorsal view,^i having a rather thick exine and a single fusi- form furrow." Pollen grains of Caytoniales, when iso- lated, without much doubt should be clas- sified under Alisporites unless there is particularly good evidence for referring them to Caytonanthus. Harris (1941) has distinguished Caytonanthus kochi, C. oncoides, and C. arberi largely on the ba- sis of differences in their pollen and has suggested their correlation with species of Caytonia and Sagenopteris found in the same deposits. All evidence seems to point to the validity of Harris' con- clusions, yet further information will probably tend to modify the picture. For example, certain of the spores isolated from the Stassfurt salt deposits by Liick (1913) seem properly referable to Alis- porites and similar to spores of Caytonan- thus, but probably few would suggest that the genus Caytonanthus should be iden- tified on that slender basis. The fructifi- cations which bore the Permian Alispo- rites would be presumed to differ generi- cally due to the general tendency of such organs to be evolutionarily plastic and amenable to structural modification. The generic features of pollen grains seem in general to be more conservative. But whether organs are evolutionarily con- servative or amenable to biocharacter al- " The terms dorsal and ventral tend to be ambiguous in description of pollen grains, particularly unless some fur- ther explanation is given. Probably "dorsal" as used here refers to the proximal side, i.e., the side which was internal and adjoined the other members of the original tetrad. CALAMOSPORA 49 teration, the important point relates to recognition of the degree of change that is manifested and of the fact that bio- character modification has proceeded at different rates in the different organs and parts of the plant life cycle. In order that fossil identifications may be scien- tifically trustworthy these principles should be more widely acknowledged. Alisporites is a designation that should be very useful in reporting the occurrence of caytonian spores and others of the same sort which perhaps may not even be necessarily referable to that group. Whether the Caytoniales are worthy of ordinal distinction is perhaps subject to some question now that Harris (1941, 1940) has shown so convincingly that they are gymnospermous and have no particular connection with angiosperms. Likewise whether they should be termed pteridosperms is still perhaps too much a matter of individual opinion. The problem of caytonian relationship with other gymno'Spermae is a perplexing one which may find its truest solution in the exacting study of fossil pollen grains similar to Alisporites, Parasporites, and Pityosporites. The more adequate generic definition of Alisporites should be taken up in con- nection with such studies. Daugherty's generic distinction of these interesting plant microfossils is a forward step which will provide a much more precise means of referring to them. They can be dis- tinguished from spores of Parasporites by their lack of a trilete commissure and probably in their distal mode of exit; from Pityosporites in having the bladders placed opposite one another and not in- clined distally, also in their lack of a thick proximal cap. Although it would seem possible to recognize morphologic homologies between the parts of Alispo- rites pollen and the spores of Endospo- rites and those of other cordaitaleans, these genera seem more distantly related and are easily distinguished. Pollen sim- ilar to that of Lehachia and Walchiantkus, now placed in the new genus Florinites (p. 56), is distinguished from pollen of Alisporites by the presence of an annu- late bladder. 1. Alisporites opii Daugherty, 1941, Carnegie Inst. Washington Pub. 526, p. 98, pi. 34, fig. 2. ! 2. Alisporites spp. ? Pollen Forms I, II, and III, Liick, 1913, Beitrag zur Kenntnis des alteren Salzgebirges im Berlepsch — Bergwerk bei Stassfurt nebst Bemerkungen uber die pol- lenfuhrung des Salztones, pp. 29-31, figs. 53-56, 58-59. Genus Calamospora S. W. and B., gen. nov. Plate 3, figures 22-22b ; text figure 1 Symmetry. — Spores trilete ; radial. Shape. — Spherical or nearly so ; when compressed, readjustment to a disk-like form leads to formation of characteristic sharp taper-point folds of variously cres- centic or narrowly lenticular outline. Some- times the spores are folded double so that the whole external outline is sharply len- ticular. Such folds are one of the char- acteristic features of the genus. Size. — Highly variable from about 40 microns (or smaller in some abortive specimens), to several hundred microns in diameter. Ornamentation. — Spores are character- istically very smooth in general appear- ance ; on closer inspection they may be minutely granulose or slightly rugose. There may be very slight differentiation of proximal pyramic areas shown by this almost negligible emphytic marking. Larg- er spores often possess a high gloss when observed by reflected light. Haptotypic structures. — Trilete rays notably short; usually they do not ex- ceed one-half the length of the spore radius. The suture line is distinct and attenuate ; sometimes moderate lips are developed. Arcuate ridges are commonly not distinguishable although they may be present as slight rounded thickenings. In some forms the pyramic areas have a somewhat different surface texture than the rest of the spore coat. Spore coat. — Relatively thin. Spores less than 100 microns diameter are gen- erally yellowish and highly translucent ; larger spores (which certainly are mostly megaspores) are progressively less trans- lucent as the wall thickness increases rel- ative to the spore diameter. Actual wall thickness thus is highly variable in vari- ous species, ranging from about 15 microns in forms over half a millimeter in diam- eter, to less than 2 microns in those smaller than 100 microns. A very thin 50 PALEOZOIC FOSSIL SPORES endosporal membrane may be present, oftentimes shrunken away from the ex- osporal coat. .■J^';//Vy-— Hartung's (1933) investiga- tion of spores in fructifications that were referred to six different genera allied with the calamarians has provided an excellent basis for systematic treatment of plants of this sort which are only represented by their isolated spores. In general, the spores do not appear to lend themselves to grouping into restricted generic alli- ances but show a rather generalized char- acter. Certain species may easily be distinguished by spore differences but generic characteristics (in terms of the fructifications) are not present or at least not yet recognized. The genus Calamos- pora therefore must be regarded as co- ordinate with a large proportion of the calamarians as they are now reported. In addition, Nemejc has shown that spores isolated from Noeggerathiostro- bus (Nemejc, 1935) and from Discinites (ibid., 1937) are similar in character and he regards this as sound evidence of alli- ance between these and the articulate groups although the degree of relation- ship is still difficult to evaluate. For the present Calamospora is re- garded as allied to the groups just men- tioned and probably is also correlative with some Mesozoic equisetaleans fcf. remarks on Equisetosporiies, p. 47). Thus it is one of the most broadly related of the groups regarded as showing evidence of actual natural relationship. There is no question but that it would be advan- tageous to subdivide the group into sec- tions or into several genera but such a treatment does not seem practicable now. Zerndt (1934) has obtained spores of generally similar character by maceration of coal and refers those over 200 microns in diameter to his Type 2. Several dis- tinct species of Calamospora are repre- sented by it. He regards these forms as pertaining to the Calamite group; how- ever, there appears to be no very definite evidence of their generic correlation. Their character nevertheless is indicative of affinity with the calamarian branch of the Articulateae. Many of the type Bg spores of Raistrick (1933, 1934, etc.) are referable to this genus but probably not all of them should be classed here. There are a few forms of Punctati-sporites that are very similar in definable features but nevertheless show differences of habit that set them apart from Calamospora. Some of the B3 types Raistrick has illus- trated have trilete rays more extended than they generally are in spores of Cala- mospora, and it is possible that some cala- mosporan forms may have been included in his type Bg. Miss Knox (1938, p. 461) has remarked on the affinity the Bg spores and compared them with spores of Calamostachys binneana and Cheiro- sfrobus petty cur ensis. Remarks. — Calamospora is unique among genera typically represented by plant spores in that megaspores, micro- spores, and probably isospores, are in- cluded in it. The Calamarians seem not to have developed as specialized a type of heterospory as quickly as the lyco- pods or other groups, and it has been generally doubted that they ever achieved a comparable state of heterosporous de- velopment. Elias believes, however, that seed-like bodies were developed in the group of Annularia stellata and he has distinguished these forms as Carpannu- laria (Elias, 1931). This material was obtained from beds of lower Des Moines age. Such a view also was expressed by Renault and others of the French school when calamites with secondary wood were regarded as gymnosperms, but at least a very large part of the seed evidence then was based on chance association with seeds that we now know have no calamite relationship. No seed megaspores have yet been reported in the bulbose bodies of Carpannularia and this or some simi- lar type of evidence will probably be nec- essary before their seed-like nature can be regarded as established. One feature stands out in the sporoge- nous sequence of the calamite alliance. More evidences of incipient heterospory are observable there than in any other fossil group. Apparently normal spores of two sizes but otherwise similar in ap- pearance are found in adjacent sporangia. The larger spores frequently are admixed with abortive forms in various stages of development. Sporangial masses com- CALAMOSPORA 51 posed of abortive and fertile spores have also been observed in coal maceration residues (Schopf, 1938, p. 51). Most noteworthy is the fact that large spores exceeding half a millimeter in diameter show characters very similar to those of less than 100 microns in diameter. Al- though heterospory was well established, a seemingly continuous size gradation ex- ists between the largest members and the smallest with little but relative size as a distinguishing feature. Spores in fructi- fications which are less than about 90 microns in diameter frequently are re- garded as microspores and those that ex- ceed that size often appear to represent megaspores. But in different species and in general there seems to be no exact way to distinguish between them unless the contrasting size spore is present in the same fructification for comparison. Species distinguished within Calamo- spora probably include spores of one mor- phologic category only and thus hetero- sporous plants may oftentimes be recorded under two specific names. This is no serious disadvantage if we are interested in obtaining the most precise information that such material can provide. If rec- ords thus integrated are obtained, it will eventually be possible to trace the corre- lation of large and small spore forms in some instances. It is doubtful that the majority will soon be resolvable in this fashion, however, and in the meantime there is need for the most accurate re- porting of this material that is feasible. Six species previously described and named are listed below, one of which is queried, and one new species described. Spores of this generic character are com- mon in many American coals and this genus will be useful in providing a sys- tematic means of recording them. It has seemed advisable to describe C. hartungi- ana, one of the characteristic American forms, to serve as a genotype of Calamo- spora rather than utilizing the published description of a species previously estab- lished. Most of the descriptions already published are less detailed than desired and are still insufficiently illustrated for purposes of close comparison. 50>^ Fig. 1 — Calamospora hartungiana sp. nov., draw- ing from microprojection of holotype. ]. Calamospora hartungiana Schopf, sp. nov. Text figure 1. Description. — Spores spherical, compressed to polygonal or acutely lenticular out- line, generally with several more or less lunate lenticular folds. Total spore diameter ranges from 80-100 microns where the outline is not foreshortened by obvious folds. Trilete rays about one-fourth of spore diameter, sutures distinct and slightly undulating, lips low but definite of from less than one to two microns width on either side of the suture. Arcuate ridges may be regarded as lacking but the pyramic areas are slightly thicker and darker by transmitted light than the rest of the spore coat and have slightly coarser surface texture (kontakthof ). Spore coat thin, one micron or less, generally yellowish translucent or some- what red when light is transmitted through numerous thicknesses due to folds or the over- lapping of several spores. Spore coat minutely rugose to granular ; at low magnification' the surface appears quite smooth. Calamospora hartungiana corresponds in gen- eral with spores Hartung (1933) obtained from fructifications of Macrostachya and Paleo- stachya, some species of which are described as having a similar "kontakthof." He suggests that microspores and isospores of the Calamari- ans range from 60-150 microns in diameter, and spores of the species just described presumably would fit one of those categories. Probably much more needs to be known of the spores of calamarian fructifications in order to be cer- tain of this, but there can be no doubt as to the general affinity of C. hartungiana. The specimens occur fairly abundantly in the 8- to 10-inch coal of the Macoupin cyclothem (middle McLeansboro age) which is exposed along Salt Fork of Vermilion River northwest of Fairmount, Vermilion County, Illinois. Spores of this sort occur in spore masses 2 mm. or more in length, as well as isolated among the other materials in the residue. The holotype specimen illustrated (text fig. 1) was separated from a spore mass of this sort when the material was being mounted in diaphane. Thus numer- ous other specimens have also been available for comparison. The size- of the spore masses and number of spores loosely pressed together in them (several hundred, although the masses 52 PALEOZOIC FOSSIL SPORES obviously are only incomplete contents of single sporangia) gives a little additional information on the Calamospora fructification. The holotype is from slide Y of maceration 90, deposited in the Illinois Geological Survey collections, Urbana. 2. Calamospora laevigatus (Ibrahim) S. W. and B., comb. nov. Laevigati-sporites laevigatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, pp. 17-18, pi. 4, fig. 46. Laevigati-sporites laevigatus Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol 4, no. 3, p. 146, pi. 7, fig. 2)6. (Calamitif )-sporites laevigatus (Ibra- him) Wicher, 1934, idem, vol. 4, no. 4, p. 172. 3. Calamospora microrugosus (Ibrahim) S. W. and B., comb. nov. Sporonites microrugosus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 447, pi. 14, fig. 9. Laevigati-sporites microrugosus (Ibra- him) Ibrahim, 1933, Sporenformen des Aegirhorizonts, etc., p. 18, pi. 1, fig. 9. 4. Calamospora mutabilis (Loose) S. W. and B., comb. nov. Calamiti( ?) -sporonites mutabilis Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 19, figs. 50a-c. Calamiti(?)-sporites mutabilis (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 145. 5. Calamospora (?) obesus (Loose) S. W. and B., comb. nov. Sporonites obesus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 19, fig. 49. Laevigati-sporites obesus (Loose) Ib- rahim, 1933, Sporenformen des Aegirhori- zonts, etc., p. 19. Laevigati-sporites obesus (Loose) Ib- rahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 145. 6. Calamospora pallidus (Loose) S. W. and B., comb. nov. Sporonites pallidus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 449, pi. 18, fig. 31. Punctati-sporites pallidus (Loose) Ib- ^ rahim, 1933, Sporenformen des Aegirhor- izonts, p. 21. Punctati-sporites pallidus (Loose) Ib- rahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 146. 7. Calamospora perrugosus (Loose) S. W. and B., comb. nov. «, Laevigati-sporites perrugosus Loose, 1934, idem, p. 145, pi. 7, fig. 13. Genus Reinschospora S. W. and B., gen. nov. Plate 2, figures 11-llb; text fig. 2 This genus appears to include the spores Reinsch (1884) designated as his Sub- divisio III of his Subtribus I, Nucleus triangularis. The genus is therefore ap- propriately designated in honor of Reinsch. His description (loc. cit. p. 7) of the subdivision is as follows (free translation from Latin) : Exospore thin (1/24-1/40 of the transverse diameter), smooth or ornamented with granules disposed variously in regular series. Body mar- gins armed with a simple series (sometimes duplicate) of dentiform juxtaposed spines, gradually decreasing (in length) from the cen- tral margins toward the angles. This diagnosis has no taxonomic status although it (and others given by Reinsch) may be quite v^orthy of validation. One naturally hesitates to apply names on the basis of Reinsch's descriptions and fig- ures because of uncertainty as to the sources and present location of the types. Also, modern interpretations may vary considerably from his as to the signifi- cance of certain morphologic features. The point which should be emphasized is that much of Reinsch's work is of value in extending our knowledge of spore form variation, particularly when essen- tial confirmation can be obtained from additional material recently prepared. Thus the forms Reinsch described in his subdivision III are structurally clarified by study of American forms of Reinscho- spora and there is little doubt that three or four still unnamed species have been dis- covered in his studies. However, there is no particular occasion to propose names for these until similar new material is at hand or the source defined for his old types (if they are preserved) and their character confirmed. The following new generic diagnosis has been constructed on the basis of newer work but it also takes cognizance of the structural fea- tures unmistakably shown by Reinsch. Symmetry. — Spores radial, trilete. Shape. — Body lenticular to flattened el- liptical in the axial plane; subtriangular in transverse plane, corners rounded, sides slightly convex or concave in outHne. A fimbriate flange, broadest in the inter- radial region and very narrow or absent on the corners, may be present to modify REINSCHOSPORA 53 the marginal outline. In some forms the interradial appendages may be dissected into a row of spines, their ends similar to the flange in marginal contour. When compressed, folds are relatively infre- quent due to preferred sedimentary ori- entation parallel to the transverse plane of the spores. Size. — Forms now known range from about 30 to 85 microns in diameter. Ornamentation. — Body smooth to gran- ulose ; flange is closely striate to markedly fimbriate, sometimes so incised as to form a row of apiculae, long in the interradial region and much shorter toward the cor- ners. Haptotypic structures. — Trilete rays well defined and extended nearly to the corners of the spore body ; it is uncertain whether the fimbriate flange can be inter- preted as equivalent to an arcuate ridge since, contrary to usual arcuate ridge de- velopment, it is least evident opposite the ends of the rays. Reinsch has illustrated forms with granules in rows paralleling the trilete rays; these may have a hapto- typic origin, and might be equivalent to arcuate ridges in mode of formation. Spore coat. — Moderately thin (less than 3 microns) and of uniform thickness ex- cept at the equator where it is joined by the flange. Affinities. — Unknown ; possibly filicin- ean. The similarities in morphology sug- gest that this is a group having natural significance. It may be related to Granu- lati-sporites. Remarks. — The genus is reported from central Russia by Reinsch (1884, his di- agnoses 62-66, incL), from the Ruhr by Loose (1934), and from Lower Pennsyl- vanian coals of Tennessee by Bentall. Dolgner (1932, pi. 28, fig. 4) has illus- trated a form with marginal spines, from Moscow brown coal, which is congeneric with our interpretation of Reinschospora; Brokaw has recently found somewhat similar forms in the coal of the Bogota cyclothem (upper McLeansboro age) in the Pennsylvanian of Illinois. The geno- type species described below is based on the Tennessee material. The generic name honoring Reinsch seems appropri- ate inasmuch as Blackburn and Temperley (1936) have shown that the algae for- merly known as Reinschia probably now 50m Fig. 2 — Reinschospora hellitas sp. nov., drawing of holotype. can be identified with Botryococcus. Reinsch's work still is significant and con- tinues to command respect for his volu- minous observations. 1. Reinschospora bellitas Bentall, sp. nov. Text figure 2 Description. — Compressed spore subtriangular in outline with inter-radial margins usually con- cave. Diameters vary from 57 to 76 ix, with an average of about 68 ix. Spore coat less than 2 fx in thickness and levigate. Trilete rays extend four-fifths or more of the distance to the equator. Flange usually originates slightly proximal from the spore equator. Fimbriate elements of flange number about 50 between ray extremities and have a maximum length of 28 II midway between rays and a minimum length of four to five at the ends of the rays. Incisions of flange, when present, apparently due to imperfect preservation. Reinschospora hellitas has been isolated from Angel and Battle Creek seams of the southern Tennessee coal field. Though never numerically abundant, it is present in nearly all samples. It is very similar to R. speciosa (Loose) but is somewhat larger in size. Occurring with R. bellitas is a form essentially comparable in all characters except that the flange is absent. The relationship, if any, between these forms is not known. Holotype — text figure 2. Tennessee Division of Geology collection ; Battle Creek seam, north side of Sweden Cove, Marion County, Ten- nessee. 2. Reinschospora speciosa (Loose) S. W. and B., comb. nov. Alati-sporites speciosiis Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 151, pl. 7, fig. L 54 PALEOZOIC FOSSIL SPORES Genus Lycospora S. W. and B., gen. nov. Plate 3, figures 19-19b Symmetry. — Spores radial ; trilete. Form. — Flattened lenticular as seen in lateral profile, equator marked with a short thick tapering ridge ; nearly circu- lar or very broadly triangular in the transverse plane. Compression is gener- ally parallel to the transverse plane with few or no folds, due to preferential ori- entation of the spores. Size. — Mean diameter in various spe- cies ranging from about 18 to 45 microns. Ornamentation. — Spores nearly smooth ; minutely granulose or rugose. Sometimes the minute rugae tend to be more prominent radially. Haptotypic features. — Trilete rays ex- tended nearly to margin, suture lines dis- tinct, usually without noteworthy differ- entiation of lips. In species with most strongly developed emphytic ornament, the haptotypic features also are more prominent. The equatorial ridge corre- sponds to the arcuate ridge in its origin and rarely becomes so extended and mem- branous across its width as to resemble a typical flange development. Usually it is narrow and tapers evenly and rapidly away from the spore body; in some spe- cies it may be almost nonexistant. Spore coat. — Relatively thin, sometimes less than 1 micron thick on proximal and distal surfaces and quite translucent. Thicker at the equator where there is no easily defined line of demarcation between the equatorial ridge and the body wall. Affinity. — Spores of this type have fre- quently been encountered in the tips of Lepidostrobus cones and there is little doubt that most of these forms found iso- lated represent microspores of the arbor- eous lepidodendrids. Andrews and Pan- nell (1942) have described similar forms which persist in tetrad groupings as mi- crospores of a species of Lepidocarpon. The microspores of sigillarians (as shown by Mazocarpon) are significantly larger and probably are not included in Lyco- spora (cf. Schopf, 1941). Spores assigned to type D^^ by Raistrick belong without question to Lycospora, also probably his B^ and D2 Type in part (Raistrick and Simpson, 1933, Raistrick, 1937, 1938). Knox (1938) has recog- nized the relationship of these types to Lepidostrobus and compared D^ and D2 with Lepidostrobus jacksoni and L. old- hamius respectively. Type D^^ which she illustrates may also belong to Lycospora. There is, of course, great inherent un- certainty in identifying any species of Lepidostrobus from isolated microspores alone. Microfossils of this kind obtain- able from coal probably are derived from many more species than are likely to be recognized from entire fructifications be- cause cones are essentially more fragile and require more favorable conditions for their good preservation than spores do. The spores probably constitute a more compete record of the various closely re- lated forms, whether they can be distin- guished easily from one another or not. Reinsch (1884) has illustrated a num- ber of forms, chiefly from central Russia and from Zwickau, Saxony, which are probably referable to Lycospora. His numbered diagnoses 32, 81, 83, 87, 240 424 ( ?), 525 ( ?) and 550 may be cited. Remarks. — Four species previously de- scribed are assigned to Lycospora all of which represent new name combinations. Lycospora micro papillata (Wilson and Coe) is designated as the type species. The genus is well represented in the Penn- sylvanian; several additional species have been recognized but are not yet described. 1. Lycospora micropapillatus (Wilson and Coe) S. W. and B., comb. nov. Cirratriradites micropapillatus Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 184, fig. 6. 2. Lycospora minutus (Wilson and Coe) S. W. and B., comb. nov. Cirratriradites minutus Wilson and Coe, 1940, idem, p. 183, figs. 11 and 12. 3. Lycospora pellucidus (Wicher) S. W. and B., comb. nov. Sporitcs pellucidus Wicher, 1934, List. Palaobot. Arb., vol. 4, no. 3, p. 186, pi 8, fig. 29. ' i^ . 1^ ■ 4. Lycospora pusillus (Ibrahim) S. W. and B., comb. nov. Sporonites pusillus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 448, pi. 15, fig. 20. Zonales-sporites pusillus (Ibrahim) Ib- rahim, 1933, Sporenformen des Aegirhor- izonts, p. 32, pi. 2, fig. 20. Granulati-sporites pusillus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 146, pi. 7, fig. 3. RAISTRICKIA 55 Genus Raistrickia S. W. and B., gen. nov. Plate 3, figures 18-18b; text figure 3 Symmetry. — Spores radial; trilete. Shape. — Sperical to slightly flattened in transverse plane and slightly triangular. Little preference as to plane is shown by compressed specimens. Size. — Spores of diiTerent species vary from about 40 to 90 microns in mean di- ameter. Ornamentation. — Characteristically ver- rucose or spinose ; spines when present are generally heavy, abruptly truncate, blunt tipped- Close observation often shows the spine tips to be minutely dis- sected into two to six terminal papillae. The prominences of verrucose forms are similar to the spines but shorter. Except for the trilete structure, features of em- phytic ornamentation extend fairly evenly over the whole spore surface. Haptotypic features. — Trilete rays va- riable in length as in Punctati-sporites ; generally inconspicuous, lips unorna- mented. Rays sometimes slightly undu- lant due to proximity of spine bases ; arcuate distinctions absent. Spore coat. — Usually of moderate thickness (two to six microns) perisporal membrane lacking. Affinity. — Typical forms of Raistrickia are without doubt filicinean. Spores of Senftenhergia plumosa, as identified by Radforth (1938), agree with these of Raistrickia, and similar schizaeaceous sporangia obtained by Andrews and Schopf from shale above Illinois coal No. 6 also contain spores of this type. How- ever, the spores of a form identified as ^. pennaef or mis by Radforth (1939) show rather different features, and those of 6". sturi do not appear congeneric with Raistrickia. Although the general rela- tionship cannot easily be doubted, the specific features do not permit easy cor- relation and Senftenhergia and Raistrickia no doubt represent generic groups of somewhat different circumscription. Raistrick's types E3 and E^ (Raistrick and Simpson, 1933 ; Raistrick, 1934, 1935, 1937) belong in Raistrickia. His Eg type is particularly characteristic. That rep- resented as E^ by Knox (1938), how- ever, is somewhat questionable. Forms Knox (1938) illustrates as D3 and Dg probably belong in this genus. Reinsch (1884) has illustrated only three forms which belong to this group without much doubt. These are described under his diagnoses 52, 308, and 588 from central Russia and from Zwickau, Saxony. The form illustrated by Thiessen (1920) in plate 61, figure B, from Benton, Illinois (probably the Herrin No. 6 coal), is characteristic of Raistrickia; that shown above it in horizontal thin section may likewise belong here but it is impossible to be sure. The two illustrations show the inadequacies of thin sections for iden- tification of spores. Remarks. — Five species previously de- scribed are assigned to Raistrickia. These forms are widely distributed as may be inferred from the preceding discussion. None of these previously named are very suitable for designation as a type, however, inasmuch as the more typical form, which particularly seems generic- ally distinct from Punctati-sporites, has not previously been formally designated although it seems fairly common. Fur- thermore, inadequacies in the previous descriptions and illustrations also mih- tate against selection of any of them as types for this genus. The species de- scribed below, Raistrickia grovensis sp. nov. is therefore designated as the type. 1. Raistrickia (?) abditus (Loose) S. W. and B., comb. nov. Sporonites abditus Loose, 1932, Neues Jahrb.. Beilage-Band 67, Abt. B., p. 451, pi. 19, fig. 53; V errucosi-sporites abditus (Loose) Loose 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 154. . 2. Raistrickia fibratus (Loose) S. W. and B., comb. nov. Sporonites fibratus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 19, fig. 52. Setosi-sporites fibratus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 148. 3. Raistrickia grovensis Schopf, sp. nov. Text figure 3 Description. — -Spore rounded triangular in out- line, about 50 microns in diameter. Orna- mented with numerous short (3-5 /x) broad (2-6 iJi) blunt tipped spines, which by juxtapo- sition sometimes appear to be mutually attached. Rays 15-20 ix long, slightly undulating, lips not raised but forming a narrow margin on either side of the suture line. Wall somewhat vari- 56 PALEOZOIC FOSSIL SPORES able (3-4 ix) in thickness, appearing brownish translucent ; spines are darker. 50/^- Fig. 3 — Raistrickia grovensis sp. nov., camera lucida drawing of holotype. This form, in general, resembles the spores of Radforth's (1938) Senftenbergia plumosa hut the spines are shorter, broader and less crowded and the spores he illustrates are round in outHne. It is difficult to make a precise comparison with other previously described species now assigned to Raistrickia because of inadequacies of their description and illustration. The spine tips of this form are entire and rounded ; others from the same coal bed and elsewhere show spine tips slightly dissected. No doubt several species of schizaeaceous ferns are represented. The specimen shown in text figure 3 designated as holotype, is from the Herrin (No. 6) coal (up- permost Carbondale age) which is worked by extensive open-cut mining operations near Mid- dle Grove, Illinois, in northern Fulton County. 4. Raistrickia saetosus (Loose) S. W. and B., comb. nov. Sporonites saetosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt., B., p. 452, pi. 19, fig. 56. Setosi-sporites saetosus (Loose) Ibra- him, 1933, Sporenformen des Aegirhori- zonts, p. 26. Setosi-sporites saetosus (Loose) Ibra- him, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 148. 5. Raistrickia spinososaetosus (Loose) S. W. and B., comb. nov. Sporonites spinososaetosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 452, pi. 19, fig. 55. Apiculati-sporites spinososaetosus (Loose) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 24. Apiculati-sporites spinososaetosus (Loose) Ibrahim, Loose, 1934, Inst. Palao- bot. Arb., vol. 4, no. 3, p. 153. 6. Raistrickia superbus (Ibrahim) S. W. and B., comb. nov. Setosi-sporites superbus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 27, pi. 5, fig. 42. Genus Florinites S. W. and B., gen. nov. Plate 2, figures 13-13b ; text figures 4, 5 Symmetry. — Pollen grains apparently bilateral ; they may nevertheless be derived from tetrahedral tetrads. Form. — Broadly elliptical in outline due to the form of the bladder; body some- what more spherical aftd nearly entirely enclosed by the bladder ; greatest diam- eter of body corresponds to the major diameter of the bladder. When com- pressed the bladder oftentimes is least afifected by folding; the body generally is marked by numerous sharp angular folds, especially around its periphery. Sise. — Greatest diameter (length of bladder) among different species varies from about 50 to 180 microns ; the pollen body proper varies from about 20 to 110 microns in diameter in individuals of vari- ous species. Ornamentation. — Bladder ornament similar to that of Endosporites and Pity- osporites; exterior surface smooth or very finely granulose or rugose, internally the membrane bears a distinct reticulation net v^^hich tends to be obsolescent in the cen- tral proximal area. Bladder and body walls are joined distally, and centrally from this juncture there is no evidence of reticulation. Haptotypic features. — Generally not evident, and probably always extremely obscure. Trilete imprint (when discern- able) is wholly vestigial, probably located entirely on the bladder, and has no func- tional significance in the mature spore. The effect of the haptotypic relations sometimes may be obscurely expressed in the proximal bladder reticulation pattern. Spore coat. — Bladder membrane is ex- panded on all sides of the body except for a small distal area. The bladder mem- brane generally has a very well defined reticulation network located on the inner surface. At the folded margin the blad- der membrane commonly appears thicker than it actually is due to the reticule bars appearing in profile ; the membrane proper usually is less than a micron thick. Con- sequently, it is highly translucent between FLORINITES 57 the bars of the reticulum which may be extended as much as 2 or 3 microns away from the membrane surface. The body wall inside the bladder is everywhere thin, oftentimes less than a micron, and shows no structural differentiation. Because of its tenuousness, it may not be exosporal in nature although it would seem rather precisely equivalent to the thicker and presumably exosporal body wall of Endo^ sporites. The body wall of Florinites is slightly less translucent than the bladder due in considerable part, at least, to its rather numerous folds. Affinity. — Florinites is known to be re- lated to Cordaianthus (pars), Lehachia, Ernestiodendron and Walchianthus through the investigations of Florin (1936, 1938-40); thus it seems evident that it represents a part of the upper Pale- ozoic gymnospermic alliance. That some cordaitaleans possess pollen grains of the Endosporites type is shown by Cordaianthus cf. C. shuleri Darrah represented in collections made by Wilson. Similar pollen grains are shown by Oliver (1940) to be present in the pollen cham- ber of ''Stephanospermum" caryoides which, according to Florin (1937, p. 310), is no doubt a cordaitalean seed. Oliver's figures 40 and 42 show a characteristic, and no doubt functional, trilete apparatus. Florin's study o{ Cordaianthus saportanus pollen and pollen grains of Florinites type associated with it (his Cordaianthus sp. 1 and 2) indicates that the suture was entirely absent and the trilete marking completely vestigial. In the walchian conifers not even a vestigial remnant of the haptotypic marking has been reported but it might be present (just as in the Cordaianthus sp. 2) and still not be easily demonstrable from coalified compression material. The agreement in pollen struc- ture between the Cordaianthus material studied by Florin, and his walchians is such as to leave little doubt that a direct relationship exists between them. Just how this relation evolved and should be systematically expressed cannot now be stated. The pollen structure is so spe- cialized, particularly when compared with the expression of spore characteristics elsewhere in the plant kingdom, as to per- mit no conclusion other than direct re- lationship. The possibility of convergent evolution being responsible for the struc- tural similarity in the two instances is entirely remote. These gymnospermic groups, often- times regarded as distinct, thus have a community of relationship which is mea- gerly expressed for the present in the genus Florinites. The probability is that this group will also be of continuing value for classification of these spores. If plants of the cordaito-coniferous type were partially restricted to upland habi- tats their pollen grains may offer the only widespread available record of their ex- istence. For this reason it is essential that they be classified as precisely as pos- sible. Florinites is also related to Endospo- rites, Pityosporites, Alisporites, and Para- s pontes, all of which have the character- istic bladder-forming perisporal (?) mem- brane. The study of these in conjunc- tion with fructifications and other organs with which they are more or less definitely correlated cannot fail to add greatly to our knowledge of gymnospermic phylo- geny during a critical period in the evo- lution of this plant group. The defini- tion of the age at which these genera are first recognizable will be of geologic as well as botanical significance. Remarks. — This genus is proposed to apply to gymnospermic plants whose pol- len generically corresponds with the type species as expressed, in general, by the foregoing definition. Florin has given rather similar data in description of the species ''cordaitiformis" which he places under the genus name Pollenites (see be- low). There is little doubt that such a gen- eric designation is inadequate but question may arise concerning the validity of the specific epithet. Pollenites cordaitiformis was estab- lished to include pollen which elsewhere in his monograph Florin had assigned to five other species of plants. The only illustrations given to validate the name had also previously been allocated differ- ently to the other five species. Florin's photographs still serve as an excellent basis for discussion of the character of isolated Florinites pollen but none of those he illustrated are properly identified with this genus because no single organ- ism can have more than one valid name. Specimens properly assigned to Lehachia, 58 PALEOZOIC FOSSIL SPORES Ernestiodcndron, or Walchianthus can- not in the same instance also be assigned to some different genus and species. Fur- thermore, it seems an absurdity to at- tempt to make an identification with a generalized group such as Pollenites or even Florinites, when a more precise spe- cific pertinence is thoroughly demon- strated. The systematic problem that confronts us revolves around proper de- termination of fossils where pertinence to a definitely restricted group is, for one reason or another, legitimately in question. Thus it would seem that unless one of the pollen specimens Florin has illustrated is regarded as improperly re- ferred to species of Lehachia, Ernestio- dendron, or Walchianthus, none of them can be regarded as type material of a new distinct taxonomic species. As there is no discernible reason to question Florin's more precise assignment of that material, it seems equally evident that Pollenites cordaitiformis has no type and the name was superfluous as proposed. Florinites antiquus sp. nov. is de- scribed below and designated as the type species. Two other species previously described also are tentatively assigned to the genus. There can be little doubt that Types 5 of Millott (1939) and 3K of Knox (1942) are also congeneric. Flor- in's various descriptions and excellent il- lustrations have shown clearly that a number of other species can be distin- guished by their pollen characters. Florinites is known to be widely dis- tributed in the Pennsylvanian and Upper Carboniferous as well as in the Permian strata where other coniferous remains are associated with it. Pollen of this char- acter has been found to be abundant in some Tennessee coals and other beds of lower Pottsville age. It seems likely that a detailed study of these forms will add much to our knowledge of the early an- cestry of conifers. 1. Florinites antiquus Schopf, sp. nov. Text figures 4, 5 Description. — Pollen grains, bilateral symmetry, equipped with an annulate bladder joined to the central pollen body only on the distal side, thus outlining a "contact area" equivalent to a germinal "furrow." The grains are broadly elliptical when compressed in the proximo-distal plane so that the length and breadth proportions are evident. When compressed laterally the axial dimension is observed to be shorter than that in either of the other two planes of sym- metry. Inclusive of the bladder, the fossils range from 55-90 (mostly 65-85) microns in length, and 40-75 (mostly 45-60) microns in breadth. The axial dimension, exhibited in a few instances, is a little less than 40 microns. The body, enclosed by the bladder on all but the distal side, is more nearly spherical ranging from 25-45 (mostly 27-37) microns in length and 20-40 (mostly 25-35) microns in breadth. The axial dimension of the body is somewhat less, about 20-25 microns. The body proper is always compressed with sharp folds which are characteristically more numerous around the periphery ; the bladder is oftentimes hardly folded when compression is in the proximo- distal plane, but the folding that does affect it is more erratic. When compressed obliquely or laterally the outline is much distorted. The bladder membrane itself is less than a micron thick ; a zone a few microns broad around the equator is more definitely yellowish trans- lucent and slightly thicker ; compressed laterally the equatorial zone of the bladder remains dis- tinguishable and the slight inequality in thickness evidently contributes to the more erratic char- acter of folds. The bladder is smooth to min- utely granulose externally but shows an internal reticulation net. The reticulae vary from 1 to 3 microns in diameter and may be very slightly elongated radially. The bars of the reticulation are not prominent in this species and have a thickness of a fraction of a micron. The reticu- lation is more pronounced away from the body and is hardly discernable immediately surround- ing the "contact area" on the distal side or cen- trally on the proximal side. The body wall is essentially smooth, very thin, and except where folded into several thicknesses, it hardly darkens the light straw-yellow translucence of the cen- tral area of the pollen grain. The "contact area" where bladder joins the body wall is elliptical or oval with its greater diameter commonly transverse to the length of the pollen grain. The margins of this area are quite faint and its size and outline vary from 17 x 23 to 10 X 20 microns. It can hardly be detected under the microscope unless the grain is oriented distal side up. A distinct but tenuous striation traverses the "contact area" in many instances, continues on the bladder membrane until about opposite the body margin, and is then lost as the bladder membrane developsi its more dis- tinctive ornamentation. It can hardly be in- terpreted as a monolete suture line ; it would seem more plausibly regarded as a harmome- gathic groove similar in character to those on the distal side of Monoletes prepollen but more faintly developed, since the body and bladder membranes are not nearly so thick as the Monoletes coat. The striation seems correlated in position with the long axis of the contact area and frequently is set obliquely with ref- erence to the breadth of the grain as a whole. Haptotypic characteristics are very faint and may be observed to best advantage in forms in which the body membrane is wanting, but the bladder membrane persists (perhaps as a result of maceration). In such examples, which are FLORINITES 59 50^^- Fig. a — Florinites antiquus sp. nov., drawing from microprojection of holotype. fairly common, there is a central gap or tear in the bladder membrane on the distal side cor- responding to the "contact area" and the central proximal region of the bladder wall is visible without any overlying layers. Granules seem aligned in a semblance of a central trilete pat- tern, obsolescent, with rays perhaps 6 microns long; the granulation of the surrounding area of the bladder membrane (it is too faint to be recognizable as reticulation) also seems faintly aligned to correspond to three radii. Conclusive definition is difficult in material obtained by coal maceration but such specimens tend to confirm Florin's (1936, p. 637) interpretation of prox- imal structure in the more highly reticulate and better preserved pollen identified by him as Cordaianthus sp. 2. Florinites antiquus seems distinguishable from all the similar forms Florin (1936, 1938-40) has identified with Cordaianthus, Lebachia, Ernestio- dendron, and Walchianthus, on the basis of its size. F. antiquus is evidently significantly smaller judging from measurements Florin has given and by dimensions measured directly from the illustrations he has published. His Cordaianthus sp. 1, from the Stephanian of France compares most closely in size, (bladder length, 78-90 ; width, 62-67 microns ; body length, 42-50; width 40-48 microns). The dif- ferences can best be evaluated by reference to the assembled measurements of F. antiquus presented in text figure 5. Presumably the flattening of F. antiquus pollen grains would give rise to dimensions which are slightly ex- cessive in the plane of compression and there- fore not quite comparable to measurements taken from Florin's uncompressed forms. That the species are distinct is evident from the much less pronounced bladder reticulation in F. an- tiquus. The distal striation has not been noted in forms regarded as cordaitean but is evident (though no mention is made of it) in several of Florin's figures of Paleozoic coniferous pollen, (cf. Florin, 1939, pis. 55-56, fig. 21; Florin, 1940, pis. 145-146, fig. 19; ibid, pis. 147-148, fig. 5). From its age (early Allegheny) one might assume that F. antiquus belonged with the cordaitealeans, but its features are advanced on a par with that of the Paleozoic Coniferales. When the conifers first made their appearance in the Missouri series in Kansas, they already were highly developed plants whose previous an- cestry must have been of some considerable duration. Thus it seems equally plausible that these pollen grains of advanced character and older derivation may actually be coniferous. The pollen grains, it must be recalled, are the most widely distributed of the determinable fossil plant entities and the pollen membranes are among the most indestructible in fossilization of any known plant materials. It is to this fossil-pollen record that one would most log- ically turn to obtain earliest evidences of newly differentiating plant groups that became estab- lished among the upland vegetation of the period. Body Bladder LENGTH P U^ 20 30 40 60 60 70 80 90 ytc o — Body LJ Bladder BREADTH Fig. 5 — Florinites antiquus, coordination of length-breadth measurements from 23 specimens, grouped according to 5 micron increments. (Di- mensions modified by unusual folding have been omitted). Cross-lined areas represent measure- ments from the holotype. F. antiquus has been obtained in moderate abundance from a widespread 10 to 12-inch coal in the Wiley cyclothem where it crops out along Soap Creek southeast of the town of Carbon, Davis County, Iowa. Correlation with the widespread Wiley coal of western Illinois has recently been confirmed by L. M. Cline of Iowa State College, who has measured the section, showing this coal to be in proper se- quence and about 12 feet above the lower Sea- borne coal and limestone developed in this area. The Wiley cyclothem is near the top of the Tradewater group of Illinois and is considered to be lower Allegheny in age. The holotype of F. antiquus, illustrated in text figure 4, is from maceration 413, slide 8 (unstained) in the Illinois State Geological Survey collections in Urbana. 2. Florinites (?) pumicosus (Ibrahim) S. W. and B., comb, nov. 60 PALEOZOIC FOSSIL SPORES Sporonites pumicosus Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 447, pi. 14, fig. 6. Reticulata-sporites pumicosus (Ibrahim, 1932) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 38. Reticulata-sporites pumicosus (Ibrahim, 1932) Ibrahim, 1933, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 157, pi. 7, fig. 29. 3. Florinites (?) visENDUS (Ibrahim) S. W. and B., comb. nov. Reticulata-sporites visendus Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 39, pi. 8, fig. 66. Incertae sedis In the present state of knowledge there are a number of named forms which it is impossible to treat systematically. The various generic designations Hsted below may possibly have a certain value but in any event their significance does not ap- pear to be biological. The names Spor- ites (H. Potonie, 1893), Pollenites, and Sporonites (R. Potonie, 1931) cannot be regarded as having systematic value. Forms hsted in these groups have no es- sential biologic features in common, no type species are recognized, and no sys- tematic treatment is feasible on this ba- sis. A morphologic designation apparently would have served as well to record the lack of systematic information concern- ing most material of this character. It likewise seems impossible to attach any generic significance to the names Verru- cosa-sporites (Loose, 1934), Elongato- sporites (Berry, 1937), Punctata-sporites or Zonala-sporites (Ibrahim, 1933). Such problematic forms as have been listed under these names must await future dis- position. 1. Sporites echinospinosus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 3, Krakow, p. 17, pi. 24, figs. 5-10. 2. Sporites fumosus Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 52, pi. 5, fi'>-s. 1-2. 3. Sporites hirmeri Reissinger, 1939, Paleon- tographica, vol. 84, Abt. B, p. 16. Note. — Nomen nudem, no illustration. 4. Sporites irregularis Reissinger, 1939, Paleontographica, vol. 84, Abt. B, p. 17. Note. — Nomen nudum, no illustration. 5. Sporites plicatus Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 51, pi. 7, figs. 7-9. 6. Sporites problematicus Zerndt, 1934, Acad, polonaise sci. Trav. Geol. no. 3, Krakow, p. 27, fig. 13, pi. 30, figs. 1-10. 7. Sporites spongiformis Reissinger, 1939, Paleontographica, vol. 84, Abt. B, p. 17. Note. — Nomen nudum, no illustration. 8. Sporonites biporous Berry, 1937, Am. Midland Naturahst, vol. 18, no. 1, p. 160, fig. 14. 9. Sporonites tripterus Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, p. 160, fig. 8. 10. Zonala-sporites taciturnus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 157. Sporonites taciturnus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 450, pi. 18, fig. 38. 11. Zonala-sporites ulughbeki (Loose), Ib- rahim, 1933, Sporenformen des Aegir- horizonts, p. 38, pi. 1, fig. 11. Sporonites ulugbeki Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, pi. 14, fig. 11. 12. Elongato- SPORITES RETicuLATUS Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, pp. 158, 160, fig. 12. 13. Verrucosa-sporites PERVERRUcosus (Loosc) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 157. Sporonites perverrucosus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 18, fig. 48. 14. PuNCTATA-SPORiTES SABULOSUS Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. Z7, pi. 5, fig. 43. In addition, the following types of Raistrick and Knox may be listed, for they too are of uncertain afifinity at the present time. Some of these seem to be worthy of more appropriate classification. Raistrick (1937, p. 911) B^, B^, E2, E^, Gi, Knox (1942, pp. 100-101), D^, K^. SUMMARY 61 SUMMARY This report presents the revised classifi- cation of plants identified from their isolated fossil spore coats, and deals primarily with those of Paleozoic age. Orthodox taxonomic procedure has been followed as closely as possible for reasons which are presented in the introduction. A cautious policy has been followed with regard to synonymy, and, even though forms are very closely allied, names based on different holotypes have been allowed to stand. In many instances, however, the apparent close relationship, which may later prove conspecific, has been noted. Thus names noted as synonyms are nearly all objective synonyms because they are based on a common holotype. Many difficult problems concerning nomencla- ture of fossil plants are solvable if nomen- clatural types are strictly interpreted. About 400 named species have been in- cluded in the present paper and most of these have been allocated among 23 genera which seem to serve a useful and signifi- cant purpose in classification. Additional genera no doubt will require recognition later and new information will modify the views that have been expressed for the genera described here. A number of species described previously apparently do not conform sufficiently to merit in- clusion in the same genus with the type species of the group, and at the same time do not show convincing evidence of affinity with other recognized groups. Such forms have been listed as species excludende. Attention is directed to the several species excluded from Reticulati-sporites which here is interpreted in a considerably re- stricted sense. Many of these forms re- quire much more careful study in order to arrive at a satisfactory expression of their affinity and classification. Other forms listed under incertae sedis also are lacking in sufficiently understood biological char- acteristics to support a definite systematic allocation. In dealing with plant micro- fossils it seems unavoidable that many forms worthy of description will never- theless have such problematic relationship that their assignment under incertae sedis is obligatory. The usefulness of fossils nevertheless bears a considerable relation- ship to the basic and fundamental in- formation available about them and for this reason greatest significance must be attached to species whose relationship has been reliably established. The authors of this paper and their associates have observed many new types of plant microfossils in preparations from coal and carbonaceous sediments in America. The present synopsis is the out- growth of a need for a more compre- hensive survey of previous work con- sidered from the standpoint of a con- sistent systematic poHcy. The essential features of an appropriate policy have been embodied here to serve as a working basis for the great amount of descriptive work yet to be done. We believe that this working basis will require further re- vision as new information is accumulated and presented. We further believe that such revision can be carried out with greatest efficiency and benefit to all con- cerned if the orthodox usages character- istic of mature systematic science are adopted and critically applied. ACKNOWLEDGMENTS The authors are under particular obliga- tion to Dr. W. H. Camp of the New York Botanical Garden who, at a period when other duties were particularly urgent and pressing, nevertheless found time to read the manuscript and suggest numerous desirable revisions in the manner of pres- entation. Mr. R. M. Kosanke of the Illinois Survey, Dr. Gilbert H. Cady, and mem- bers of the Survey editorial staff all have contributed in final preparation of this paper for publication. To all of these the authors owe a special debt of thanks. 62 PALEOZOIC FOSSIL SPORES REFERENCES Andrews, H, N. and Pannell, Eloise, 1942, Con- tributions to our knowledge of American Carboniferous floras, II. Lepidocarpon: Ann. Missouri Bot. Garden, vol. 29, pp. 19-35. Bartlett, Harley Harris, 1928, Fossils of the Carboniferous coal pebbles of the glacial drift at Ann Arbor : Michigan Acad. Sci. Arts and Letters Papers, vol. 9, pp. 11-28. 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Petrog. der Brennsteine Arb., vol. 4, no. 4, pp. 165-212. ■ , 1934b, Uber Abortiverscheinungen bei fossilen sporen und ihre Phylogenetische Bedeutung : Inst. Palaobotanik u. Petrog. der Brennsteine Arb., vol. 5, no. 3, pp. 87-96. Williams, H. S., 1887, On the fossil faunas of the Upper Devonian: U. S. Geol. Survey Bull. 41, 123 pp. Williamson, W. C, 1879, On the organization of the fossil plants of the coal measures, Pt. IX : Royal Soc. London Philos. Trans., 1878, vol. 169, Pt. II, pp. 319-364. ,- , 1880, On the organization of the fossil plants of the coal measures. Part X. In- cluding an examination of the supposed Radiolarians of the Carboniferous rocks : Royal Soc. London Philos. Trans., 1880, vol. 171, Pt. II : pp. 493-540. Wilson, L. R. and Brokaw, A. L., 1937, Plant microf ossils of an Iowa coal deposit : Iowa Acad. Sci. Proc, vol. 44, pp. 127-130. and Coe, E. A., 1940, Descriptions of some unassigned plant microfossils from the Des Moines series of Iowa : Am. 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A, Annee, 1937, pp. 583-599. — , 1938, Die Eignung von Megasporen als Leitfossilien: Congres pour I'avancement des etudes de stratigraphie Carbonifere, Heerlen, 1935, Compte rendu, vol. 3, pp. 1711-1732. 1940, Megasporen des Saarkarbons: Paleontographica, vol. 84, Abt. B, pp. 133- 150. Zeiller, R., 1906, Bassin Houiller et Permien de Blanzy et du Creusot : Etudes des Gites Mineraux de la France, Fasc. ii, Flore Fos- sile, pp. 140-150. Zetzche, F., Vicari, H. and Scharer, G., 1931, Untersuchungen iiber die Membran der Sporen und Pollen, IV. 3. Fossiles Sporo- pollenin aus dem Tasmanit und der Mos- kauer Braunkohle : Helv. Chim. Acta, vol, 14, Pt. 1, pp. 67-78. PLATES AND PLATE EXPLANATIONS Explanation of Plate 1 Fig. 1 — Tasmanites ; segment cut away to show mode of perforation. Symmetry is unicentric with no axis more developed than another p. 11 2 — Triletes; specimen of the sectio Aphanosonati; after Zerndt ; 2a, diagram showing un- compressed form in the axial plane corresponding with a trilete ray p. 18 3 — Triletes radiosus ; sectio undifferentiated; after Zerndt; 2a, diagram showing uncom- pressed form in the axial plane corresponding with a trilete ray .... pp. 18, 24 4 — Punctati-sporites; 4a, equatorial (transverse) plan; -^^^ axial (longitudinal) plan . p. 29 5 — Laevigato-sporites ; 5a, equatorial (transverse) plan; 5h, longitudinal plan correspond- ing with the longest axis p. 36 6 — Alati-sporites; 6a, axial (longitudinal) plan; 6b, equatorial (transverse) plan . . p. 33 7 — Reticulati-sporites ; in some instances the reticulation may cover proximal and distal hemispheres about equally; 7a, equatorial (transverse) plan; 7h, axial (longitu- dinal) plan p. 34 8 — Granulati-sporites ; 8a, equatorial (transverse) plan; 5'^, axial (longitudinal) plan . p. 32 9 — Denso-sporites ; segment cut away to show thickening of peripheral region of spore coat; 9a, axial (longitudinal) plan uncompressed; 9h, form such spores habitually assume under normal compression ; 9c, equatorial (transverse) plan p. 39 10 — Cystosporites; proximal end of spore with three abortive tetrad members attached at the apex; 10a, small drawing of complete fertile spore; 10b, apex of fertile spore show- ing trilete apparatus when abortive members are disconnected p. 40 [68 Illinois State Geological Survey Report of Investigations No. 91— Plate 1 Explanation of Plate 2 Fig, 11 — Reinschosp07'a; 11a, axial (longitudinal) plan ; lib, equatorial (transverse) plan , p. 52 12 — Alisporites ; axial (longitudinal) plan not shown; much the same as 16h but body wall thinner p. 48 13 — Florinites; 13a, axial (longitudinal) plan, proximal side up; 13b, equatorial (trans- verse) plan. The body tends to be slightly elliptical, the long dimension agreeing with that of the bladder p. 56 14 — Endosporites; 14a, axial (longitudinal) plan; it is difficult to judge whether the bladder and body wall always join distally as shown here; 14b, equatorial (transverse) plan p. 44 15 — Pf^j'o^/'on^^^; pollen grains of modern aspect ; 15a, axial (longitudinal) plan in the plane corresponding with the sulcus between the bladders ; 15b, axial (longitu- dinal) plan in the plane of the bladders . p. 27 16 — Parasporites ; 16a, equatorial (transverse) plan; 16b, axial (longitudinal) plan . . p. 42 17 — Mono/^^^^; i/a, equatorial (transverse) plan; 17b, axial plan view corresponding to the short dimension, distal grooves down, proximal suture up ; 17c, axial plan view cor- responding to the long dimension p. 38 70 I Illinois State Geological Si Report of Investigations No. 91 — Plate 2 Explanation of Plate 3 Fig. 18 — Raistrickia; 18a, equatorial (transverse) plan ; i5&, axial (longitudinal) plan . . p. 55 19 — Lycospora; 19a, equatorial (transverse) plan; i9&, axial (longitudinal) plan . . p. 54 20 — Triquitrites ; 20a, axial (longitudinal) plan; 20^, equatorial (transverse) plan . . p. 46 21 — Cirratriradites ; 21a, axial (longitudinal) plan; 2i&, equatorial (transverse) plan . p. 43 22 — Calamospora; 22a, equatorial (transverse) plan; 22b, axial (longitudinal) plan . . p. 49 (Below) Graphic comparison of approximate size range of fossils included in the various genera. Dimen- sions are indicated in logarithmic coordination. [72 Illinois State Geological Survey Report of Investigations No. 91 — Plate 3 Lycospora I Laevigato-sp|)rites Granulati-sporites | poO/^ 20. Reinschospora i =1 Denso-sporites3Q'o Triquitrites | =1 Prtyosporites Raistrickia I 3 Reticulati- 30^ Punctati -spontes | | 1+ Cirratriradites 40Q^ I 50,0^ sporites I mm. 700y^ Approximate Size -Range WITHIN "Spore" genera 2 mm. 1000/^ Fforinites J Endosporiies Calamospora 40 '/^ Alisporites | =1 Alati- sporites 3 mm. 200(>^ bOyu 60/. Tasmanites Monoletes 75^ \00yu, 150^200/^300/^ Parasporites Zonalo-sporitfes 4 mm. 6mnx Triletes 10 mm. I (abortive) I I II (fertile) | Cystosporites ILLINOIS STATE GEOLOGICAL SURVEY REPORT OF INVESTIGATIONS NO. 91 Distributed in July 1944