—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 GEOCHEMISTRY Coal Frank H. Reed, Ph.D., Chief Chemist G. H. Cady, Ph.D., Senior Geologist and Head H. W. Jackman, M.S.E., Chemical Engineer L. C. McCabe, Ph.D., Geologist (on leave) James G. McCullough, Research Associate R. J. Helfinstine, M.S., Assoc. Mech. Eng. M. Ph.D., Asst. Geologist James Schopf, Coal J. Norman Payne, Ph.D., Asst. Geologist Charles C. Boley, M.S., Asst. Mining Eng. G. R. Yohe, Ph.D., Chemist Bryan Parks, M.S., Asst. Geologist Carol Bartels, B.S., Research Assistant Robert M. Kosanke, M.A., Asst. Geologist George M. Wilson, B.S., Research Assistant Industrial Minerals Henry L. Smith, A.B., Research Assistant J. S. Machin, Ph.D., Chemist and Head Delbert L. Hanna, A.m., Asst. Chemist Industrial Minerals
J. E. Lamar, B.S., Geologist and Head Fluorspar H. B. WiLLMAN, Ph.D., Assoc. Geologist Robert M. Grogan, Ph.D., Assoc. Geologist G. C. Finger, Ph.D., Assoc. Chemist Asst. Chemist Robert R. Reynolds, M.S., Research Assistant Everett W. Maynert, B.S.,
Oil and Gas X-ray and Spectrography Chemist A. H. Bell, Ph.D., Geologist and Head W. F. Bradley, Ph.D., Assoc. Frederick Squires, B.S., ?etr. Eng. Charles W. Carter, Ph.D., Asst. Geologist Analytical William H. Easton, Ph.D., Asst. Geologist O. W. Rees, Ph.D., Chemist and Head Paul G. Luckhardt, M.S., Asst. Geologist Howard S. Clark, B.A., Assoc Chemist Wayne F. Meents. Research Assistant L. D. McViCKER, B.S., Asst. Chemist P. W. Henline, M.S., Asst. Chemical Engineer Areal and Engineering Geology William F. Wagner, M.S., Asst. Chemist B.A., Research Assistant George E. Ekblaw, Ph.D., Geologist and Head K. F. BuRSACK, B.A., Research Assistant Richard F. Fisher, M.S., Asst. Geologist Cameron D. Lewis, Mary Ann Winsche, B.S., Research Assistant Marjorie Winchester, B.S., Research Assistant Subsurface Geology L. E. Workman, M.S., Geologist and Head Arnold C. Mason, B.S., Assoc. Geologist MINERAL ECONOMICS Merlyn B. Buhle, M.S., Asst. Geologist Frank E. Tippie, B.S., Asst. Geologist W. H. VosKuiL, Ph.D. Mineral Economist Margaret Sands, B.A., Research Assistant Grace N. Oliver, A.B. Assistant in Mineral Ruth E. Roth, B.S., Research Assistant Economics Walter R. Smith, Research Assistant Douglas F. Stevens, M.E Research Associate
Stratigraphy and Paleontology EDUCATIONAL EXTENSION J. Marvin Weller, Ph.D., Geologist and Head Chalmer L. Cooper, M.S., Assoc. Geologist Don L. Carroll, B.S., Assoc. Geologist Petrography Ralph E. Grim, Ph.D., Petrographer PUBLICATIONS AND RECORDS Richards A. Rowland, Ph.D., Asst. Petrographer (on leave) George E. Ekblaw, Ph.D., Geologic Editor Chalmer L. Cooper, M.S., Geologic Editor E. Rose, B.S., Technical Editor Physics Dorothy Kathryn K. Dedman, M.A., Asst. Technical Editor R. J. Piersol, Ph.D., Physicist Alma R. Sweeny, A.B., Technical Files Clerk B. J. Greenwood, B.S., Mech. Engineer Portia Allyn Smith, Research Assistant Donald O. Holland, M.S., Asst. Physicist Meredith M. Calkins, Geologic Draftsman (on leave) Leslie D. Vaughan, Asst. Photographer
Special Staff to Aid in the War Effort
Oil and Gas Resources Ground Water Geology
Earle F. Taylor, M.S., Asst. Geologist Carl A. Bays, Ph.D., Spec. Geologist M. W. Pullen, Jr., M.S., Spec. Asst. Geologist C. Leland Horberg, Ph.D., Spec. Asst. Geologist Arnold Eddings, B.A., Research Assistant Stewart Folk, M.S., Spec. Asst. Geologist Virginia Kremers, B.S., Research Assistant Ernest P. DuBois, Ph.D., Spec. Asst. Geologist Margaret Parker, B.S., Research Assistant Paul Herbert, Jr., B.S., Asst. Geologist John A. Harrison, B.S., Spec. Research Assistant Charles G. Johnson, A.B., Asst. Geologist (on leave)
Consultants: Ceramics, Cullen W. Parmelee, M.S., D.Sc, and Ralph K. 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 association with them because there is no adequate basis for close comparison of fossil spores is scat- The literature on biocharacters exhibited by these diversified tered, and in addition there has been much organs. diversity in methods of classification. The When affinities become better known objects of the present article are to bring so that the difficulty due to uncomparable this material together in summary form features may be partially overcome, it for convenient taxonomic reference and to then may be possible to relate the forms evaluate the genera which have been pre- more naturally in classification within viously proposed. After a study of these suprageneric groups such as tribe, family, microfossils in America and a thorough and subfamily depending on how closely study of the literature it seems essential information may be correlated. Indeed, also to describe some new genera. It is assignment to the same genus, or even hoped that a fundamentally sound basis species, is theoretically possible and may has been provided for studies now in eventually be achieved in some few in- progress in the three laboratories repre- stances, but in general this implies a much sented by the authors and for future more precise and detailed historical knowl- studies. edge of interrelationships than is likely to principles which relate to the Guiding be fully established. treatment of this material are given below. For satisfactory nomenclature it is most 1. Only adherence to the systematic convenient that those fossils which possess principles embodied in the International numerous demonstrably comparable fea- Rules will give satisfactory results in the tures shall be placed under the same ge- study of these microfossils (as in other neric name. However, it* is unsatisfactory fields of paleontologic study). Therefore, to classify fossils under the same generic the International Rules of Botanical name simply because a few arbitrary fea- Nomenclature, 3rd edition (Briquet), re- tures are held in common. The essential vised by the International Botanical Con- validity of any classification depends on gress of Cambridge, 1930, and published the relative significance that attaches to the in 1935,^ have been used as a basis for various biocharacters. In fact it is by taxonomic treatment. virtue of such interpretive discrimination 2. classified Species within the same that taxonomy is to be distinguished from genus or under the same generic name a cataloging procedure. Interpretations must possess significant characteristics in nevertheless must be based on evidence and common. There can be little positive proof not on supposition ; difficulties are multi- of generic identity unless substantial plied whenever interpretation exceeds fac- homologous comparisons can be drawn be- tual bounds. tween respective biocharacters of each 3. In paleobotanical practice there has congeneric species and the type species of been much divergence with regard to the the genus. For example many isolated significance and proper interpretation of lycopsid megaspores must be classified unusually complete specimens in which separately from the microspores found in parts usually found separated, and there-
^ Gustav Fischer, Jena. fore generally classified separately, are
[7 8 PALEOZOIC FOSSIL SPORES
found in organic union. Whatever attitude to have coordinate significance through a may be adopted, it is of fundamental im- part of their geologic life as species. Dur- portance for later consistent application ing this particular period, identity of re- of taxonomic principles. For this reason it lationship must exist. But the most perti- seems desirable to state views applicable nent fact encountered in systematic treat- to the present work. ment of the common isolated forms, is that The few unusually complete specimens this period of actual synonymy can scarce- are highly significant in showing beyond ly ever be defined. As a general thing, due question that certain isolated organs at a to discontinuity in the fossil record, it is particular locality and stratigraphic level impossible to establish the points at which possess identity of relationship.* This old characters became coordinated in a identity must become more dubious, in new fashion or with other characteristics the case of isolated parts, the further they new to the phyletic succession. are removed geographically and strati- Consequently these overlapping rela- graphically from the site of proved union. tionships do not lend themselves to taxo- Stratigraphic discrepancy is most likely to nomic expression within specific and gen-
cast doubt on specific identity ; geographic eric nomenclatural categories. Groups of discrepancy may be cause for qualification suprageneric rank, which are less depend- of the presumed relationship in varying ent on type specimens for their proper degrees of subspecific magnitude (there is definition, may be better used in expressing no basis for assuming that geographic such relationships of generalized validity. races were less in evidence in the past than The present paper, however, does not deal they are at present). If evolution is a with groups of suprageneric taxonomic more or less continuous attribute of life status. processes, over a period of time significant 4. Various criteria for determination alterations in some of the many heritable of synonymy have been used by paleobot- characteristics are bound to occur. An in- anists. The practice we prefer is conserva- dividual alteration, although of specific tive to the extent that unless conspecific significance, will not necessarily involve relationship is proved for two nomencla- phenotypic expression in the majority of tural types, both names are valid. As such, the other characteristics of the race. For they are available for any purpose of use- this reason it is inaccurate to postulate the ful nomenclature. There are a large num- continued specific coordination of an ex- ber of "partial synonyms" among the tensive garniture of biocharacters over a named species of fossil plants to which period of time, without correspondingly these remarks apply. The word "synonym" detailed knowledge which proves that the — of course is an absolute term "partial characteristics dealt with were not subject synonyms" are not synonyms ; in most in- to mutative or other evolutionary processes stances the names serve useful purposes in during that time interval. This detailed in- indicating groups of different circum- formation is most readily determined for scription. None of these names should be a restricted group of biocharacters which lightly considered ; neither should a name can be observed on common specimens. be used in unqualified application unless Practical reasons dictate that emphasis in its pertinence is evident. Evidences of systematic treatment be placed on the more "partial synonymy" undoubtedly record common types of fossils rather than on close natural relationship, but the names those which show unusual preservation. should not be regarded as invalid unless Demonstrated connections between shown to be mutually inclusive. Lacking spore forms and types of fructifications this, they remain applicable to fossils con- prove readily that some of the adjacent specific with their basic nomenclatural isolated specimens of either category are types. For the research worker there are also conspecific. The diverse taxonomic advantages in this conservative view of groups, diagnosed on the basis of typical synonymy, as it tends to promote a more isolated specimens are proved, therefore, precise differentiation of fossils. This
* No plant, and certainly no single specimen, can have basically, is a fundamental reason for the more than valid one name. However, instances arise in continued use of technical nomenclature, which it is difficult to determine which of two or more names should actually be applied. In spite of this, syste- necessary to progress in the study of fos- matic procedure demands that an author follow some consistent usage. sil plants. INTRODUCTION
5. Specific identification, because of its consideration is Dawson's Sporangites. great significance, must be made critically. This name was first proposed in 1863 and Emphasis should be placed on the positive has been used ambiguously and in different evidence of similarity rather than on nega- senses by many authors. It is here re- tive evidence of lack, real or fancied, of garded as a nomen ambiguum and the features which cannot be examined criti- name Tasmanites is adopted for the most cally. frequently encountered forms that have
6. Neither spores, leaves, nor any other been commonly assigned to Sporangites. morphological part of a plant in any stage Tasmanites nevertheless is a problematic form, spore-like in of its of its life is here considered to he a many character- species. A species of plant, for the istics, but actually unassignable to any purposes of this synopsis, and regardless group of plants now known. All the other genera treated here belong with little doubt of how it may be otherwise defined, is regarded as a group of organisms. Any to the Cormophyta, and all whose affinity individual organism of such a group, how- is approximately known belong to the Tra- ever, may be identified if any diagnostic cheophyte groups, as mentioned in the structure representing any portion of its respective generic discussions. life cycle is available. Spores in many Knox (1939, 1941) has suggested that instances are probably just as definitive spores of Bryophytes may be present in of species as any other organ belonging Carboniferous coals, and some forms in a coordinated manner to the life cycle that have been described may well belong of a particular plant individual. They merit to this group of plants which at present study with other types of ancient plant are very scantly recognized in the Car- materials, and all data should be recorded boniferous. The fact that no spores of on a common basis of taxonomic equality. Paleozoic Bryophytes are definitely known Because of their unique adaptation for (Sporogonites Halle may be an exception) dispersal and their numerical abundance in makes any suggested correlation hazard- many sedimentary deposits, it is expected ous at this time.
that spores will eventually becon\e of It is important to realize that many greater practical significance than many specific features of spores of different of the larger types of plant fossils that are geologic age can be matched because of more commonly noted by the geologist. evolutionary convergence as well as be- 7. Since the purpose of giving a name cause of community of derivation. Con- to a taxonomic group is not to indicate the sequently unless there exists some cor- characters or the history of the group, roborative evidence based upon spore but is simply a convenience in reference, forms actually present in contemporane- all names that are taxonomically valid and ous fossils whose relationship can be pertinent to recognized groups should be established, all that seems warranted is to continued. The authors agree that the direct attention to similarity of the spores hyphenated generic nomenclature, for of different geologic age, recognizing that example, Granulati-spontes, Laevigato- the similarities may or may not have phy- sporites, Denso-sporites, instituted first by letic implications. is in Ibrahim (1933), poor form, lacks There is some lack of strict agreement euphony, and tends to be generally mis- between the morphological nomenclature leading. Nevertheless, the status of such applied to modern pollen and the morpho- names seems reasonably secure in several logic features of fossil forms. For one instances and their continuance is likely to thing the microspore in modern forms is cause less misapprehension than attempts determined, in a strict sense, solely by the to institute more appropriate nomencla- presence of the unicellular male gameto- ture. phyte.^ Such a distinction is of course The preceding explanatory paragraphs inapplicable to fossil forms. There is also may serve to clarify some of the policies 2 The common indiscriminate reference to all spores of adopted in this and subsequent reports relatively small dimensions as "microspores" is to be lamented. Although true miscrospores frequently are and explain their consistency with sys- small, by no means all small spores are microspores. The tematic treatment of other types of plant long established botanical usage of the term "microspore" has reference to fundamentally functional distinctions that material. are entirely aside from relative or specific size. R. B. Thompson (1927) has in fact demonstrated that in some The earliest proposed name requiring plants the microspore is larger than the actual megaspore. 10 PALEOZOIC FOSSIL SPORES the further difficulty that no simple correct that deal incidentally with Paleozoic morphological designation can be applied spores. Various non-taxonomic systems similarly to the dispersal forms of the have been used by some workers in desig- male gametophyte bodies in both modern nating types of spores in coal thin- and fossil forms. The dispersal stage sections, notably by Slater and his col- rather than the degree of gametophytic leagues in the South Yorkshire laboratory development is of greatest practical signifi- of the British Fuel Research Division, cance when fossils are considered. The by Th. Lang in Silesia and, most impor- terms perispore, exospore, and endospore tantly, by Reinhardt Thiessen. These might seem equivalent to the pollen struc- apply very slightly to the present work tures perine, exine, and intine, respec- because species cannot be as reliably dis- tively. However, in the case of the bladder- tinguished from spores in thin-sections, equipped fossil gymnospermic pollen, the significant biological characters are the bladder membrane which seems very generally more difficult to ascertain, and similar in morphologic character to the the informal nomenclature used does not perisporal membrane of certain crypto- require systematic consideration. Con- gamic forms, is regarded by many as sequently articles of this nature are not equivalent to the exine of angiospermic considered except in connection with the pollen. morphology of certain forms.
Revision of the morphologic termi- On the other hand, there is a consider- nology for fossil spores and similar micro- able literature on isolated spores in the fossils also seems required. Not only has Russian language which needs to be given there been a dearth of descriptive terms careful study before nomenclatorial stabil- available for description of the varied ity can justifiably be hoped for. Thus far structural features, but a good many deep- only a small portion of this literature has ly rooted terms have become outmoded. been available to us, chiefly in abstract Several of these date from the period in form, and this is insufficient to provide which Selaginella was presented as a a satisfactory basis for understanding the primitive forerunner of modern seed quite different nomenclature Russian plants, and their homologous implications writers have used. Naumova (1937) now oftentimes seem unwarranted. Non- mentions that about 400 species have been commital descriptive terms, which seem distinguished based on material ranging reasonably free ^of theoretical connota- in age from the Lower Carboniferous to tions, have been preferred for use in the Tertiary. He mentions genera designated present paper. It is hoped that it will be as Zonotriletes and Azonotriletes. Nikitin possible later to treat the descriptive (1934) has distinguished the genus Kry- morphology of spore forms specifically shtofovichia based on very singular ap- in greater detail. pearing, large spores of Devonian age. In the synoptic lists the species assigned Luber (1938) mentions subgroups (gen- to each genus are given in alphabetical era?) Azonotriletes, Zonotriletes, Azono- order. The order of genera adopted is Jiionoletes, Azonaletes and Zonaletes and chronologic according to the time each distinguishes several apparently valid was first proposed. Although some inter- species. In a more recent publication generic alliances can be recognized now, Luber (1939) uses the generic ( ?) names and a more logical sequence should be PlaguUtes, Turriella, Circelliella, Saccri- adopted in later works, it is felt that inalia, Circella, Libumella, Spinosella, Sub- the chronologic order of priority will be sac culifer and Plicatella. It is to be hoped of some convenience for taxonomic refer- that under more favorable circumstances ence. it will be possible to give full consideration to the Russian studies which have already Nearly all the references cited (p. 62) are those of particular taxonomic or mor- been put to good use in the age determina- phologic pertinence for the fossils dis- tion and correlation of the widely sep- cussed. Although we believe this repre- arated coal deposits of the USSR and to sents a reasonably complete survey of this integrate the nomenclature they are using literature, it does not include all articles with our own. TASMANITES 11 GENERA OF FOSSIL SPORES
Genus Tasmanites Newton, 1875 applied the name to spores or "spore Sporangites Dawson, 1863 (pars.), Canadian cases" which he thought were referable Naturalist, New Ser. vol. 8, no. 6, pp. 431-457. to Lepidodendron, Calamites and similar plants. Sporangites papillata, about one Plate 1, figure 1 inch (!) in diameter, and 6". glabra about General Discussion.—There is a group the size of a mustard seed, were men- highly distinctive fossil spore-like of tioned ; later this description was repeated, bodies which occur most notably in the accompanied by small diagrams (Dawson, Devonian-Mississippian black shale that 1865, p. 165, pi. 12, figs. 80-81). These requires a name amenable to standard fossils were associated with coal at Jog- taxonomic usages. At first, it was sup- gins, Nova Scotia, and may in part have posed that the name Sporangites, if sup- been material that would be placed now ported by an emended definition, could with Triletes, but there is no way to con- apply. We are now of the opinion that firm this from his description or illustra- the application of that name is not correct tions. His repeated descriptions of 5^. in this connection, and that the various papillata as "one inch in diameter" leads descriptions that have appeared for Spo- one to suspect it may have been a seed. rangites are so highly conflicting and In 1871 (a, b) Dawson described an ambiguous or erroneous that that particu- additional species from the black shale at lar name should be considered a nomen Kettle Point on Lake Huron as S. huron- ambiguum. This is herewith proposed in ensis; "small globular papillate spore- view of the difficult taxonomic situation cases—probably of some Lepidodendroid as detailed below. plant." Later in the same year (Dawson, The present most common use of the 1871c) he also referred a specimen con- term ''Sporangites'' is by geologists who sisting of a small whorl of leaves or bracts, wish to record the presence of more or first described as Annularia acuminata, to less disc-shaped, resinous-appearing bodies Sporangites. This last interpretation of that are very commonly found in the Sporangites was accepted by Slopes Devonian-Mississippian black shale, many, (1914). Forms similar, in general, to but by no means all, of which are properly those from Kettle Point are usually con- referable to Tasmanites as given below. strued as Sporangites by present day This common usage does not conform to geologists, but it seems evident that neither the original application of the name and of these interpretations corresponds at is so noncritical and general in application all closely with Dawson's first generic that it would probably be unwise to at- application of the name. tempt to restrict it even if this could be Dawson in 1883 presented a report on done in accordance with taxonomic pro- "Rhizocarps in the Paleozoic Period" at cedure. Possibly no one, certainly not its the meeting of the American Association author, has used the term with taxonomic in Minneapolis, an abstract of which ap- consistency and practically all records of peared in the American Association for Sporangites must merely be regarded as the Advancement of Science proceedings indicative of some small rounded, brown- (1884a). The entire paper also appeared ish or yellowish bodies. Needless to say, in the Canadian Record of Science for the many kinds of plant microfossils cor- same year (1884b). The essential des- respond to this characterization. How- criptive part and the figures are identical ever, there is no reason why "sporangites" in both. Here Dawson proposed the name may not be continued to be used in this Protosalvinia including two species given sense, since it serves a descriptive pur- as ''Sporangites (Protosalvania) hrazilien- pose, in spite of the fact that it is botanical- sis U.S.," and "S. (P.) hilohatus n.s." ly misleading; but the name should not Dawson stated that if we compare the be italicized or treated as a generic desig- separate macrospores of the Brazilian nation. sporocarps, and especially those which are Sporangites was first proposed by found detached from their envelopes Dawson in 1863 (p. 454). Here he (Sic!) with Sporangites hitronensis, we 12 PALEOZOIC FOSSIL SPORES see a remarkable similarity in size, form bious lycopod stem material, but no thalli and texture, sufficient to justify us in or "sporangia" with the "spores" at Kettle supposing that the latter may be of the Point). White seems to have made no same nature as the former, but deprived close study of the sporangite forms under of their outer cases either by dehiscence appropriate magnification and he found or decay, and this is the view which we are none of them actually in any of the multi- now disposed to take of their nature. This tudinous "spore cases" (thalH?) of Pro- better accords with their wide distribution tosalvinia which were examined. In fact, in aqueous deposits and with their ac- although an uncritical reader may infer companiment than any other supposition. from this work that ''Sporangites' belongs Plausible and satisfactory as this sup- to Protosaknnia, White actually made no position may be, there is nevertheless direct statement to this effect and he pos- scant basis for assuming all the various sibly had no strong convictions as to their sporangite bodies to have such an origin, nature. As the association of thalloid not to mention the insidiousness of at- bodies with sporangite forms is by no tempting to apply scientific nomenclature means common and as the sporangites that on the basis of such a supposition. It can- have been reported differ considerably not be said that Dawson ever proved the (one of the generic types here being identity of bodies within and outside the treated as Tasmanites), it is evident that
"sporocarps" ; furthermore, no one else Protosalvinia and Sporangites cannot be has ever provided any conclusive evidence. in any way accurately treated as syn- Evidently many have taken this important onyms. No one yet has furnished tangible point for granted, however, because the evidence of the nature of the spore-like two names are used practically inter- bodies Dawson reported actually within changeably in much of the subsequent the Protosalvinia thalli, so it still is quite literature. speculative whether any relationship The "spherical and oval sacs, the walls (other than that of a common environ- of which are composed of a tissue of ment) is evidenced by their infrequent association in the Devonian-Mississippian hexagonal cells . . . three to six mil- limeters in diameter," apparently are black shale. properly assignable to Prolosahiiiia Daw- Dawson reviewed the subject in con- son, 1884. We are unable to ascertain nection with his paper in the bulletin of their characteristics reliably from any the Chicago Academy of Science in 1886 figures Dawson published but White and (pp. 105-118), calling attention to Orton's Stadnichenko (1923) give photographs (1882) discovery of 6^. huronensis-Vik^ and data on a form, Protosalvinia raveiina bodies in the Ohio shale, Clark's (1885) White, 1923, which is confirmatory of description of somewhat similar material their thalloid characteristics, although in the Devonian of New York, the report White regarded them as sporangia. White of Johnson and Thomas (1884) on these (p. 239) has repeated the suggestion, microscopic discs in the Chicago water (which seems to have originated with supply and glacial till of the region, New- Dawson, 1871a, b) that their affinity was ton's (1875) description of Tasnianites probably with "Devonian types antecedent punctatus, and his own (Dawson, 1884a, to the Carboniferous Lepidodendra and b) descriptions of two lots of Brazilian Sigillariae." Possibly he thought the spor- material sent him by Hartt and by Orville angite bodies were actually spores of Derby. He makes no direct mention lycopod relationship, as many others either of the Joggins specimens or of have, and he probably accepted their sup- ''Sporangites'' acuminata. This article is posed relationship with Protosalvinia be- the best of any written by Dawson on the cause of close association. Oval speci- subject. Nevertheless, in it, as well as mens of Protosalvinia bear some resem- in his other papers, the inadequacies of his blance to lycopod sporangia, but there are concepts are fully apparent. He appears general objections to such an interpre- confused as to the distinction between tation. Even when they are occasionally spores and sporangia ; he definitely con- in abundance no cones or actual lyco- siders the 6^. huronensis bodies referable podiaceous remains are noted in the same to the water ferns (hence, Protosalvinia) y assemblage (Dawson reported some du- being misled on the one hand by the fal- TASMANITES 13 lacious idea of the primitiveness of that definite morphologic character. Many group of living plants, current at the have considered the Sporocarpon types to time, and on the other by association with be lycopod megaspores but from our pres- the (probably unrelated) thalloid plants ent knowledge such a view seems un- sometimes found in the same beds. Daw- tenable. Their structure is more rem- son considered his study of the Brazilian iniscent of colonial habit seen in the material particularly conclusive but his Volvocales. At any rate, there is no account of it does not seem so now and, similarity between them and Sporocarpon unless the original material can be re- furcatum Dawson. The name for this studied or new material obtained, the spe- form must be Foerstia furcata (Dawson) cific nature of the Brazilian specimens is Pia; Pia (1927) having been first to use entirely problematic. this combination, but also without com- Jongmans more recently (1930) has ment. The type species of Foerstia is listed most of the nomenclature rather best taken to be Foerstia ohioensis White, indiscriminately under the name Proto- instead of Dawson's species. Kidston salvinia. His treatment is hard to com- and Lang's work suggests that they may prehend inasmuch as Sporangites and be specifically inseparable and if this is Tasmanites are treated as synonyms ever proved by comparison of the holo- although their precedent publication dates types of the two species, White's specific are given with approximate correctness." designation, F. ohioensis, of course would In attempting to follow Dawson's some- become a synonym. what confused usage and apply it more The spores of Foerstia occur in tetrads, widely without original study, Jongmans to some extent localized along one margin has not contributed to the understanding within the fertile tips. Possibly they were of these plants or lessened the confusion formed in individual conceptacles. An in nomenclature. His omission of some analogy in more than one particular is pertinent references may in part be re- suggested by Fucus, but aside from the sponsible for the rather misleading im- possibility that Foerstia may also be an pression conveyed by this portion of the advanced type of alga, probably no inter- Catalogus. pretation of homology can be supported. One other thalloid plant occurs in asso- The individual spores of Foerstia are ciation with sporangite forms in the black about 200 microns in diameter, similar to shale which now can be more adequately those of land plants, with a heavy "re- discussed because of reexamination by sistant" spore coat about 7 microns thick, definite modern methods. Spores are present in haptotypic structures consisting of trilete sutures with pyramic areas well it and are quite different from sporangite bodies, in spite of Dawson's suggestion defined by distinct arcuate ridges. The that they were similar. Dawson (1888) distal part of the spore coat is irregu- called this form Sporocarpon furcatum. larly pitted and somewhat rugose accord- In the most thorough and essentially ing to information presented by Kidston Lang. Thus, these spores are en- definitive study of this plant by Kidston and and Lang (1925), Dawson's name was tirely different from the sporangite bodies that be associated with them. continued. However, White, in White may and Stadnichenko (1923), had renamed Haptotypic features have been de- the group Foerstia but without taxonomic scribed for the spores in all reliable comment or discussion. Nevertheless the reports that have ever been published of reason and justification for this new gen- bonafide Devonian plants showing these eric name is evident upon consideration reproductive structures. These spore of the genus Sporocarpon Williamson forms, in addition to Tasmanites and its (1879, cf. also 1880, p. 509) which was associates, may be found isolated in car- founded upon .S. cellulosum and similar bonaceous Devonian sediments. Lang forms of problematic affinity but quite (1925) has shown that eight or more distinct species are thus represented in f Dawson's paper in the Canadian Naturalist for 1870 evidently appeared subsequent to publication of his paper the Cromarty fish-beds of Old Red Sand- in the Am. Jour. Sci. for April, 1871. The paper "On stone age in Scotland. None of these Rhizocarps, etc." published in the Canadian Record of Science is dated and evidently was published in 1884, can be referred to Tasmanites, although rather than 1885. Such minor errors are unfortunate in a reference so widely used as the Fossilium Catalogus. Lang's spore-type F (in part), which 14 PALEOZOIC FOSSIL SPORES
seems to lack haptotypic features, may Newton's tasmanite which Reinsch ob- be generically similar to some of the tained from the British Museum. Reinsch spore-like rugulose-membranous associ- assigned these forms, along with others, ates of Tasinanites in American deposits. in his group called the Discieae, char- So far as these plant microfossils and acterizing them generally as others of similar character but question- compressed disc-like plant bodies of original able nature aref concerned, an adequate subspherical or subelliptical outline. They are study of them is hardly begun. entirely closed, with walls sometimes perforate and show a thin central cavity which has been The essential point to be made in reduced by compression. regard to ''Sporangites/' which geologists Variations in the walls are mentioned generally consider is well known, is that due to inclusion of other forms which Dawson applied the name in the first seem to resemble Calcisphaera William- place to ambiguous material and then son (1880, p. 521), Sporocarpon Wil- later applied it without consistency to liamson (1879, pp. 346-349; 1880, pp. specimens of at least other diverse two 507-11), and many other diverse forms, and unrelated types. He further con- some of which may even represent true fused its application in conjunction with sporangia of articulate plants. Reinsch at least two other generic names. Geolo- also described 15 kinds of Discieae from gists have accepted one of the usages of the older Mesozoic. For the most part Dawson, but as this does not correspond forms derived from the Carboniferous with Dawson's original interpretation it of central Russia and Saxony contrast seems impossible to retain it in systematic sharply with the Tasmanites forms from usage. The term "sporangite" ought America, Australia and Tasmania. These therefore to be considered no more than last evidently were chiefly in mind as he a descriptive designation without tax- described the Discieae, for his group onomic implication. description fits them best. Specific no- The systemic designation Tasmanites menclature is applied to only one form was rejected by Dawson who recognized so that Reinsch's work, although it shows the similarity of Tasmanites punctatus best the characteristics of various Tas- Newton (1875) to his Sporangites hu- manites specimens, is essentially not a ronensis (Dawson 1886, p. 116) because, taxonomic contribution. But since it as the "name Sporangites had priority, I establishes the similarity of American and (i.e., Dawson) it do not think necessary Tasmanian forms we may feel no hes- to adopt this term (Tasmanites), although itancy in calling these peculiar bodies there can be little doubt that these organ- Tasm^anites and in accepting Newton's isms are of similar character." This species as the type for the genus. It similarity can be verified from Newton's seems so generally similar to some of illustrations by any one who has care- the material provided by Orton and fully examined abundant sporangite speci- Thomas there is a question as to proper mens from the Devonian-Mississippian specific distinction between them, but black shale, but the similarity is by no being so widely separated geographically means evident if all the various inade- quate figures of "S. huronensis" given as they are, and stratigraphically as they by Dawson and his contemporaries are seem to be, the forms should remain under consulted. separate names, at least until a definite first-hand study can be made. No spe- The best evidence that at least many cific definitions are attempted here, but it of Dawson's vS'. huronensis specimens, and particularly those expressly men- is felt that the generic definition given tioned by Orton, are similar to those of below is sufficiently accurate to serve to Newton has been presented in the cred- distinguish the group from forms which itably illustrated work by P. F. Reinsch, unmistakably are spores of cryptogams vol. II of the Micro-Paleophytologia,' and higher plants, and from several other published in 1884. Material from the kinds of problematic fossils which are Ohio shale and from Chicago sent by found in Illinois (and probably else- Orton and B. W. Thomas is described where) associated with Tasmanites, but and illustrated and comparisons are read- which must be considered generically ily made with similar material from distinct from it. . :
TASMANITES 15
Diagnosis.—An adequate description of thicker and the thinner sacs, or to find them in anything like relation of an inner and outer Tasmanites punctatus, taken here as the the coat. type species of the genus, is given by Prof. Balfour, I believe, considers and excerpts from his paper Newton (pp. the Tasmanite discs to be closely allied to
339, 340, 341) are quoted below : Flemingites; they differ from them, however, When the separated discs are viewed by re- as Mr. Carruthers has pointed out (Geol. Mag. flected light, they appear as more or less 1865), both in structure and size. All the circular bodies, somewhat thickened towards Flemingites macrospores which I have seen the circumference, many of them having their have homogeneous walls, and in many of them surfaces raised into irregular folds. If mounted is seen the triradiate marking, which is so gen- in Canada balsam, and viewed by transmitted erally present in cryptogamic spores (Prof. light, many have the appearance represented Williamson, MacMillan's Mag. March, 1874, in pi. 10, figs. 2, 3, 8, while others exhibit the p. 409). In none of the Tasmanite sacs have I folds to which allusion has just been made. been able to see this triradiate marking, al- The more perfect discs are seen to be surrounded though their structures are so clearly shown by a double contour-line—the optical expression that these markings could not fail to be seen of the fact that these discs are really thick- if they were present ; and the walls, as we have walled sacs. The saccular character, however, already seen, have a definite structure. is best seen in transverse sections (figs. 1, 4, The inconvenience of having an 5), or when the sac is broken (fig. 8). A object without a distinctive name induces me closer examination enables one to see that the to propose one for the spores (?) found in walls of these sacs are not homogeneous. A Tasmanite and Australian White Coal (the view such as fig. 8 shows numerous dots scat- two being, as I believe, identical in structure) ; tered over the surface, which become some- and in order to retain existing titles as far what elongated towards the edges of the disc. as possible, I would suggest that Prof. Church's When examined with a power of about 250 name Tasmanite, which is so generally used in diameters, the dots can be resolved into minute reference to the schist as a whole, be retained circles about 1/3000 of an in inch diameter for this substance, and that the spores (or with a still smaller in dot the centre, as shown rather the plant to which they belong) should in fig. 9. These structures are best in seen be called Tasmanites, with the specific title of the discs of Coal. It White may be thought punctatus, in allusion to their surface-markings. that these dots are comparable to the granules to be seen upon the surface of some of the The features which are here regarded macrospores of Flemingites ; but the study of as generically characteristic of Tasmanites transverse sections shows at once that these are as follows dots are not mere surface markings, for they can be distinctly traced as minute lines (tubes?) Symmetry.—Unicentric ; there is evi- passing from the outer to the inner surface. These dently a center and not an axis of sym- lines are shown in fig. 5, but owing to the section metry as in spores of bonafide plants. not being quite in the same plane as the lines, Shape. — Originally spherical ; except they do not appear to extend quite through. In addition to the fine lines, the walls of the where protected, compression has altered sacs exhibit obscure longitudinal markings, them into disks with a few sporadic which give them a laminated appearance (fig. rounded folds. J ) Si^e.—Ranging from less than 100 Neither Mr. Carruthers (Geol. Mag. 1865, slightly p. 432), nor Mr. MacNaughton (Trans. Roy. microns to 600 microns or greater Soc. Van Dieman's Land, vol. ii, 1855, p. 116), diameter. Forms greatly in excess or mentions any structure in the walls of these much smaller than these dimensions are sacs. suspect, because they vary so greatly The discs vary in diameter, as stated by from the genotype species. both these authors, from about 1/80 to 1/50* of an inch. Mr. MacNaughton speaks of a Ornamentation.—Surface smooth and thin outer coat to these discs, which may be glistening in reflected light at low magnifi- seen when they are ruptured. I have examined cation; more detailed examination shows all my preparations, both sections and sep- more or less rugosity which may be in arated discs, in order to distinguish this outer coat, but have been unable to do so. One easily part attributable to preservation. More recognizes in transverse sections, such as fig. 1, or less regularly spaced punctae varying that the walls of the sacs vary much as regards in number on different forms are visible, thickness ; and among the separated sacs which but not conspicuous. The forms may be are mounted in balsam some may be seen much described as essentially lacking in external more transparent than the rest; but I have failed to see any real difference between the ornamentation.
* 1/80 and 1/50 of an inch = 317.5 and 508 microns Haptotypic features.—Entirely absent. respectively. However, Newton's 2 figure "a large spore" False conclusions either given at 50x would be over 650 microns in diameter. have been drawn From his figure 5 (250x) the wall is about 2 5 microns from dififerent forms in association with thick. In this specimen the wall is about 1/26 of the total diameter. ^ Tasmanites or from specimens poorly 16 PALEOZOIC FOSSIL SPORES
preserved and misinterpreted. Absence ample, the form shown on his plate 10, of trilete sutures is diagnostic. figure A (1921) is stated in the text to Wall. — Generally moderately thick, be 0.85 mm. (850 microns) in diameter mostly 1/10 to 1/25 of the diameter; but the legend says it is magnified 140 X, wall evenly developed on all surfaces and and measurement of the figure shows the never membranous, often punctate with form to be at that rate about 170 microns pores tapering from very small orifice on in diameter. It may be this form is the the outside; sometimes the punctae are same as some observed in association with very sparse, in other species they may be Tasmanites in Illinois, but should not be so densely packed as to give a radially generically identified with it. The walls striate appearance. Poorly defined con- are generally quite rugulose-undulant and centric bands may be present in the wall, may show a poorly defined very coarse but these are ordinarily not easily visible reticulation net, but the membranous char- unless material is sectioned. In optical acter of the fossil suggests that it orig- section (transmitted light) aside from the inally was smoother and that it has been punctae, walls generally appear homo- impressed irregularly by grains of the geneous. enclosing sediments. No trilete suture (tetrasporic marking) can be demon- Affinity.—The question of the affinity strated on material examined at first of Tasfimnites, so far as it can be def- hand nor is any structure of this sort initely discussed now, hinges on whether discernible in Thiessen's figures. Just as these bodies can be taken to represent Newton remarked in his description of plant spores or not. Spores of all plants Tasmanites punctatus, above the algal stage that are readily in none of the sacs have I been able to see this preserved as fossils have markings on triradiate marking, although their structures are the spore coats of various but definite so clearly shown that these markings could not significance. All plant spores arise fail to be seen if they were present. normally as tetrads (groups of four) It must be recalled that the trilete and from a single spore mother cell. All monolete (haptotypic) structures of cryp- primitive plants (aside from algae) show togamic plants are a very definite spore evidences of their tetradic derivation in feature, consisting not merely of a ridge the structure of the spore coat. The which might be abraded from an other- Tasmanites forms do not show any such wise featureless surface, but actually con- structure and, in addition, have funda- sisting of two margins, sometimes thick- mentally a unicentric symmetry which is ened, and a definite suture line between in strong contrast to plant spores which them which cannot be removed except are always radially or bilaterally sym- by abrasion of the whole thickness of the metrical. Therefore, it seems conclusive proximal wall of the spore. that Tasmanites is not allied with any of Thiessen also fails to make any men- the known primitive land plants. It may tion of the punctations which are so represent an algal type but if so, no cor- characteristic of the thicker walled forms relative vegetative algal remains can yet referable to Tasmanites, and which are be recognized. definitely but rather crudely shown in The preceding remarks indicate that the illustrations of Johnson and Thomas. ^ the writers' observations are by no means Had he appreciated what they were, it is in agreement with Thiessen's (1921, doubtful that he would have been so 1925) on what, it appears, must include positive that these fossils were spores the same types of material. Thiessen of pteridophytes, because certainly no persists in describing all forms from the known spores of pteridophytes possess black shale as if they had "tetrasporic" such structures. marks (in some cases more or less The most recent paper bearing on the removed by abrasion). He definitely subject concerns an examination of Tas- identifies them with Dawson's Sporangites manian oil shale by Singh (1932). He huronensis and with the Tasmanites forms considered that he was examining mate- illustrated in 1884 by Johnson and rial similar to that studied by Newton in Thomas (cf. Thiessen, 1921, p. 293) but 1875 which contained Tasmanites punc- discrepancies make it impossible to en- tatus. However, from the discrepancy tirely confirm the statement. For ex- in results it may be that he was dealing TASMANITES 17
with a different microfossil assemblage. blage. That Reinsch and Singh have Some of Singh's specimens without doubt found a few trilete spores associated are bonafide plant spores. However, he with Tasmanites is not to be wondered found many forms with about the same at. The writers are convinced that, de- shape and size proportions as T. punctatus spite assocations of this sort, Tasmanites but was unable to see any of the punctae. does not now, and never has had, any Trilete markings were clearly visible in trilete marking on its wall. Lack of this some but for others he suggests they feature is conclusive evidence that Tas- ''seem to have been rubbed off during manites does not represent any group of fossilization." This, of course, seems ordinary pteridophytic plants. quite improbable since the trilete suture Discussion. — There are many spor- lines, if present, cannot be removed by angite forms which do not qualify as fossilization. Of the figures presented by species of Tasm^anites, or any other genus Singh, actually only one (fig. 3) shows yet designated. Aside from Lang's paper convincing evidence of the trilete mark- there has been no attempt to report on ing. The proportions of this specimen isolated actual bonafide plant microfossils suggest Calamospora and it is interesting of similar age. Until more reports are to note that Phyllotheca (a calamarian) available and greater knowledge has been is associated in the same strata. Those obtained, it does not seem essential to shown in his figures 7 and 8 are incon- give further taxonomic consideration to clusive as to the presence of a trilete the few that are known. The correlation structure, since Tasmanites may occasion- and coordination of taxonomic groups ally be folded under compression to simu- identified by means of spores from the late a trilete marking ; but careful observa- Devonian, with those groups based on tion will distinguish the absence of a spores which we are utilizing for Carbo- definite suture line. Since Singh has niferous material, deserves very thorough not shown these structures, actually the study and consideration. The abundant only great discrepancy between Newton's plant record that can be obtained may, if and Singh's observations with respect to advisedly interpreted, have much sig- the more common forms concerns the nificance in tracing lines of phyletic rela- punctae. tionship among plants. The results that American Tasmanites forms show con- may possibly accrue from study of the siderable variation in the frequency of problematic microfossils, among which is punctae and this may be a useful means Tasmanites, is hard to predict, although of specific discrimination. One can such study cannot fail to at least outline scarcely doubt their existence in Newton's their stratigraphic usefulness. original material, because Reinsch also It needs to be stressed, however, that figured them from the Tasmanite mate- Tasmanites, and other unicentric micro- rial provided from the British Museum fossils certainly do not belong to the well and also used and figured additional speci- known groups of primitive higher plants mens from Newton's own preparations. that were contemporaneous with them. So far as tlie discrepancy in presence of Their morphologic nature is, in fact, so a trilete mark on some specimens is con- uncertain that their reference to the plant cerned, it must be noted that Reinsch (II, kingdom would not seem assured except 1884, p. 5) also found "Rarissime oc- for their chemical composition. Zetzche, current Triletes minores singuli inter- et al., (1931) seem to have definitely spersi" in Newton's original material. proved the absence of nitrogen from Furthermore, the way Newton's descrip- authentic Tasmanites and furthermore tion in general tallies with the verified shown their composition to be in general characters seen in American Tasmanites similar to unquestioned fossil plant spores. is so striking as to leave no doubt as to Though possibly chitinous in appearance, the essential correctness of his description. lack of nitrogen apparently eliminates The surprising thing about American the animal kingdom from consideration. assemblages containing Tasmanites is that, Spores of Parka which evidently is a so far as is known to be true, no trilete thalloid plant, evidently lack any hapto- forms (excepting Foerstia) have yet been typic structures (Don and Hickling, found to occur with them in the assem- 1915) but in no other particular do 18 PALEOZOIC FOSSIL SPORES
they correspond to Tasmanites. In fact, 3. Tasmanites punctatus Newton, 1875, these microscopic fossil bodies represent Geol. Mag., ser. 2, vol. 2, no. 8, p. 341, pi. 10, figs. 2, 8. about as great a realm for speculation The type species of Tasmanites. (See de- now as they ever did as to their actual scription as previously quoted from Newton.) nature and affinity, but this can hardly be 4. Tasmanites spp. Reinsch, nos. represented as a valid excuse for their 12, (p. 3) ;
17, (pp. 3-4) ; 26, (p. ; 32, (p. 6) and continued neglect. By strict application 5) 65 ( ?), (p. 9) ; 1884, Micro-Pal eophytologia. of systematic principles we shall approach Reinsch's diagnoses and illustrations sug- much nearer a useful understanding of gest the numbers listed above, based on Amer- them. ican, Australian, and Tasmanian material, are probably referable to Tasmanites. Although The following list includes names which others he assigned to the Discieae also lack have been used will and have to be con- haptotypic spore features, they are probably sidered in subsequent treatment of species generically dissimilar. of Tasmanites; forms identified as Spor- Note.—There is a possibility that the iso- angites jacksoni White (1905) and 5^. lated bodies associated with Protosalvinia hra- radiatus Duden (1897) have nothing to siliensis Dawson and P. hilohata Dawson are referable to Tasmanites, and if so, they should do with Tasmanites and are omitted on receive separate taxonomic consideration and this account. not be assigned to Protosalvinia. Proto- salvinia clarkei Dawson, which was originally 1. Tasmanites chicagoensis (Reinsch) S. identified with Tasmanites huronensis (Daw- W. and B., comb, nov.^ son) by Clarke (1885) probably is entirely Discieae, Subtribus I, Subdividio 1., different and deserves restudy. One would 18. ( addita est nomen Specificum judge that Tasmanites also occurs associated "Chicagoense") Reinsch, 1884. Micro-Pale- with this material. Probably many of the ophytologia, vol. 2, p. 6, pi. 72A, fig. 76. "Sporangites" occurrences given by Williams Named from material supplied by Thomas, (1887) and various other authors are referable the specific name must be credited to Reinsch to Tasmanites but a critical reexamination of and the type is the specimen he figured. Daw- all of this material is needed. son (1888b) and Jongmans (1930) errone- The forms designated as Sporangites alasken- ously cite Thomas as its author.*' sis and Sporangites arctica by David White (1929) were studied only from thin sections. 2. Tasmanites huronensis (Dawson) S. The former shows wall peculiarities which may W. and B., comb, nov. be comparable to that of Tasmanites ; the latter Sporangites huronensis Dawson, 1871, appears very similar to sectioned spores of Am. Jour. Sci., 3rd Sen, vol. 1, no. 4, p! higher plants. Whether the breaks in the walls 257, figs. 1-3. are accidental (as they would be in Tasmanites) The following description is from the most or are due to the section transecting a definitely complete account given by Dawson (1886) in organized suture of a spore coat is difficult if which he corrected some of the earlier inac- not impossible to determine from sections alone. curacies. It has also been changed slightly This material was considered possibly Lower in accordance with the present conception of his Cretaceous in age and a definite identification material. Definite descriptive data is essen- of Tasmanites would be of considerable interest. tially unaltered. This is presented with the hope that it may be useful until Dawson's Genus Triletes (Reinsch, 1884) type material or authentic topotype material ; can be reexamined. Schopf emend., 1938. Flattened disc- like bodies with smooth rounded edges usually Plate figures 2a, ranging from 2 50 to 350 microns in diameter,' 1, 2, 3, 3a showmg slight external evidence of punctation but by transmitted light at higher magnification exhibiting nu- Megaspores radially symmetrical ; prox- merous pores which penetrate the wall. Sometimes marked imal side marked by triradiate sutures, by characteristic rounded folds or split open by compres- sion. By transmitted light they appear amber colored or often with arcuate ridges connecting the sometimes have a reddish hue; in section the walls show taint indications of concentric lamination. The walls are ends of the rays; distal surfaces smooth moderately thick 1/10 to 1/20 of the total diameter. They never or variously ornamented. Ornamentation exhibit the triradiate marking seen in spores ot lycopods. The interior is usually vacant but in some generally less well developed on proximal cases more or less filled with mineral matter (verv fine pyritic crystals, calcite, etc.). than on distal surfaces. Spores are rel- atively very large up to more than 3 mm. ^The initials S. W. and B. used here and elsewhere throughout this paper refer to the present authors, Schopf, Affinity with the free-sporing lycopsids. Wilson, and Bentall. copy Type species, by designation, Triletes ,^ A of theMicro-Paleophytologia originally owned by Thomas now in the John Crerar Library in Chicago reinschi (Ibrahim) Schopf (1938). has the inscription ''Chicagoensii of B. W. Thomas" writ- ten on two of the figures, one of which is not Reinsch's This genus is a large and important ™.^^ ^ ^^^. ^^'^^ interpreted his material ;a littlelifJr^-ff u differently than Reinsch did and may have mis- one in classification of fossil spores, par- understood the taxonomic principles mary involved. The pri- ticularly in the Carboniferous period purpose of author citation is to indicate type upon which material the description is based when the free-sporing lycopsids were TRILETES 19 dominant. Evidence of the affinity of to him for examination. It may be like- these spores is shown by every spore- wise a matter of individual opinion as containing- megasporangiate fructification to whether certain holotypes are conspe- of the free-sporing lycopsids of this pe- cific. Such questions tend to be academic riod. Unfortunately, this corollary in- because there can be no doubt that the formation has not always been considered relationship is close in these instances. in fossil spore classification and additional Also, whether the forms are distinguished genera have been proposed which are in as species or as varieties may be a matter conflict with Triletes. The types of these of individual taxonomic policy. We have subsequently defined and consequently un- made as few changes of status as seemed tenable genera are listed below. They are reconcilable with a uniform taxonomic also included in alphabetical position in the practice. Inconsistencies too may be main list of species with their synonymic found here, but it is hoped that these are citations. within permissible limits of individual Triletes reinschi (Ibrahim) Schopf (1938), opinion. which also is the type species by designa- The synonomies are not absolutely com- tion for Triletes. (Type of Laevigati- sporites Ibrahim 1933). plete in the sense that references to all forms that have ever been described or Triletes tuberosus (Ibrahim) S. W. and B., comb. nov. (Type of Tiiberculati-sporites illustrated are included because triletean Ibrahim, 1933). forms are represented in too many pub- Triletes sextus S. W. and B., nomen nov. (As lished illustrations for them all to be Triletes of Bennie and Kidston VI (1886), cited. The chief concern has been to this form was regarded as the type of give the references which include the type Apiculati-sporites Ibrahim, 1933). material or throw light upon it. Other Triletes hirsutus (Loose) S. W. and B., comb, nov. (Type of Setosi-sporites Ibrahim, references incidental to this purpose are 1933). given in a good many instances, chiefly Triletes sextusdecimus S. W. and B., nomen to indicate the general features of occur- nov. (As Triletes XVI of Bennie and rence, both stratigraphic and geographic. Kidston (1886), this form was regarded as the type of Zonales-sporites Ibrahim, Stratigraphic distribution has been gen- 1933). erally given for each. Such ranges are Triletes trilobus (Ibrahim) S. W. and B., always subject to revision for a number comb. nov. (Type of Valvisi-sporitcs Ibra- of reasons. Many have been taken from him, 1933). the stratigraphic distribution tables and By no means all the species that authors discussion pubhshed by Zerndt (1931, p. regarded as congeneric with these types by 169; 1937a, p. 68; 1937b, p. 590, 593; virtue of arbitrary (artificial) distinctions 1938; 1940, p. 142). The correlation chart appear so when broader comparisons are given by Gothan (1937, p. 299) has attempted. This is particularly true of been helpful in understanding the rel- the smaller forms (isospores and micro- ative position of the mostly unfamiliar spores) which often have been denomi- named units of the Polish and Lower Sile- nated indiscriminately. These forms have sian stratigraphic succession. Approximate of course been reallocated to what have equivalence of European and American the appearance, at least, of constituting Carboniferous divisions are given in the more natural groups. chart previously published by one of us The following is a list of described (Schopf, 1941, p. 9). Zerndt's data on species of Triletes as the genus is now stratigraphic distribution is much more conceived. Some forms listed here may extensive than information from any be so closely related to one another that other source and consequently deserves in later studies the names will be con- most serious consideration. Doubtless sidered as synonyms. Without consulta- many forms not reported by him have a tion of the types or specimens correspond- greater stratigraphic range than has been ing to them these questions as to actual recognized. Zerndt's data, however, are synonymy appear to be unsolvable. W^icher given for his spore ''types" rather than ( 1934) has placed several of these names for species. Most of his types and species in synonymous relationship, and in all have general equivalence yet this is not probability correctly, since many of the true in all cases and some discrepancy poorly illustrated holotypes were available may occur on this account. In general, 20 PALEOZOIC FOSSIL SPORES probably a good many specific determi- Selaginellites ariadnae Miner, 1932, Washington Acad. Sci. vol. nations have not been rendered critically Jour., 22, p. 505, figs. 26-30. and this also tends to diminish enough Age, Upper Cretaceous. the value of stratigraphic ranges which can be given now. Nevertheless, some 6. Triletes arnoldi (Miner) Harris, 1935, Meddelelser om Gronland, vol. 112, p. 155. general information may be gained from Selaginellites arnoldi Miner, 1932, the ages listed. It has been observed that Washington Acad. Sci. Jour., vol. 22, p. a good many of the distinctive forms ap- 500, figs. 22-25. pear in the same relative positions in Age, Upper Cretaceous. America and in the European Carbonif- 7. Triletes augustae (Loose) S. and B., erous (Schopf, chiefly unpublished data) W. comb. nov. which gives promise of future assistance Zonales-sporites augustae Loose, 1934, in long distance correlation. Inst. Palaobot. Arb. vol. 4, p. 150, pi. 7, Ninety-five species are included here. fig. 32. Many of them are referable to the sections Sectio, Auriculatif Age, Westphalian B (Gasflamm). of Triletes already proposed (Schopf,
1938) ; others are less definitely assigned. 8. Triletes aurantium Harris, 1935, Med- Certain relationships between species have delelser om Gronland, vol. 112, No. 1, pp. 164-165, fig. 52H-L, pi. fig. pi. been noted when these seemed particu- 25, 5, 26, fig. 5. larly evident. Triletes is by no means Age, Liasso-Rhaetic. an exclusively Paleozoic genus and Meso- zoic forms have also been included for 9. Triletes apiculatus (Ibrahim) S. W. and B., comb. nov. sake of completeness. Several new names Apiculati-sporites apiculatus Ibrahim, have been proposed here, in addition to 1933, Sporenformen des Aegirhorizonts, p. the considerable number of combi- new 23, pi. 5, fig. 31. nations required in order to follow a Type 14 Zerndt (in part), 1931, Acad, consistent and relatively uniform tax- polonaise sci. Bull, internat., ser. A, p. 172; onomic policy. 1935, Acad, polonaise sci. Trav. (jeol. no. 1, pp. 17-18. Sectio, Aphanosonati. Age, Lower Na- 1. Triletes agnina (Zerndt) S. W. and B., murian, B, comb. nov. Westphalian and younger (?). Lagenicula agnina Zerndt, 1937, Acad, 10. Triletes artecollatus Nowak and Zerndt, polonaise sci. Trav. Geol. no. 3, p. 14, pis. 1936, Acad, polonaise sci. Bull, internal., 21-22. ser. A, p. 58, pi. 1, figs. 4, 5. Type 34 Zerndt, 1937, idem. Type 39, Zerndt, 1936, idem. Sectio, Lagenicula. Age, Lower Na- Sectio, Triangulati. Age, Dinantian murian (Flora series). (Raczna Series).
11. Triletes auritus Zerndt, 1930, idem, ser. 2. Triletes ales Harris, 1935, Meddelelser B, p. 46, pi. 1, figs. 4 and 5. om Gronland, vol. 112, no. 1, p. 163, fig. Spore 0.7 Zerndt, geol. 53A-E, pi. 25, figs. 2, 8, 9, 11. mm. 1930, Soc. Pologne Ann., vol. 6, pp. 307-303, 312, pi. Age, Liasso-Rhaetic. 1, figs. 3a, b, pi. 3, figs. 3a-d. Types 11 and 12 Zerndt, 1931, Acad, 3. Triletes angulata (Zerndt) S. W. and polonaise sci. Bull, internal., ser. A, p. 172. B., comb. nov. Triletes auritus Zerndt, 1937, idem, p. Lagenicula angulata Zerndt, 1937, 584, pi. 10, figs. 2, 6. Acad, polonaise sci. Trav. Geol. no. 3, pp. Triletes auritus Zerndt, 1940, Paleonlo- 11-12, fig. 8, pis. 14, 15. graphica, vol. 84, abl. B, p. 146, pi. 9, figs. Type 7 Zerndt, 1931, Acad, polonaise 5-18. sci. Bull, internal., ser. A, p. 171, pi. 3, Sectio, auriculati (type). Age, West- fig. 8. phalian C, D. Sectio, Lagenicula. Age, Lower Na- Triletes auritus var. grandis Zerndt, 1937, murian (Grodziec series). Acad, polonaise Bull, internat., ser. A, p. 584, pi. 10, figs. 1, 3-5. 4. Triletes areolatus Harris, 1935, Med- Type 11a, Zerndt, 1937 (see above). delelser om Gronland, vol. 112, no. 1, pp. Triletes auritus var. grandis Zerndt, 158-159, fig. 51A-F, pi. 26, figs. 3, 10. 1940, Paleontographica, vol. 84, abt. B, p. Age, Liasso-Rhaetic. 134, pi. 11, figs. 32-35. Age, Stephanian (Westphalian D?). 5. Triletes ariadnae (Miner) Harris, 1935, Triletes auritus var. secundus (Maslan- idem. p. 155. kiewiczowa) S. W. and B., nom. nov. TRILETES 21
Triletes auritus II Maslankiewiczowa, Sectio, Lagenicula. Age, Lower West- 1932, Acta Soc. Bot. Polonaise, vol. 9, phalian C. (SuppL), p. 158-161, figs. 33-36. Note.—This species is apparently closely re- lated to Triletes robertianus and T. latihirsutus. 12. Triletes appendiculatus Maslankiewic- zowa, 1932, idem., p. 163-164, fig. 39. 15. Triletes borealis (Miner) Harris, 1935, Type 12 Zerndt, 1937 (non 1931), Acad, Meddelelser om Gronland, vol. 112, no. 1, polonaise sci. Trav. Geol. no. 3, p. 3. p. 155. Type 6 Sahabi, 1936, Recherches sur Selaginellites borealis Miner, 1932, les spores des houilles Francaises, p. 39- Washington Acad. Sci. Jour., vol. 22, p. 503, 40, pi. 2, fig. 5. figs. 12-21. Type 8? Sahabi, 1936, Recherches sur 16. Triletes brasserti Stach and Zerndt, 1931, les spores des houilles Francaises, p. 41, Gliickauf, Jahrg. 67, no. 35, p. 1123, figs. 29, pi. 2, figs. 7-8. 30, (?) 3L Sectio, Auriculati. Age, Westphalian C, Type 20 {Triletes brasserti Stach and D, Stephanian. Zerndt), Zerndt, 1934, Acad, polonaise sci. Trav. Geol., No. 1, pp. 23-24, fig. 9, pi. 25, 13. Triletes bennholdi Zerndt (non Bode), figs. 1-2. Age, Dinantian, Namurian polonaise sci. Trav. Geol. no. 1937, Acad, Westphalian A, B. 3, p. 5-6, fig. 3, pi. 2, figs. 1-6. Note.—Closely related to Triletes circumtex- Type Zi Zerndt, 1937 (idem). tus, T. saturnoides and T. superbus. Porosporites bennholdi (Bode) Zerndt, Cong. Stratig. Carbonif., 1938, Deuxieme 17. Triletes breviaculeatus Nowak and vol. III. p. 1713. Age, Lower Namurian Zerndt, 1937, Zerndt, idem., no. 3, pi. 4, (Flora series). figs. 1, 2. Type 38, Triletes breviaculeatus Nowak Note.—Zerndt credited this specific epithet to and Zerndt, 1936, Acad, polonaise Bull, Bode, apparently assuming the spores con- internat., ser. A, p. 57. specific with those of Porostrobus bennholdi. If Sectio, Aphanozonati. Age, Upper Din- Porostrobus can be identified reliably from its antian (Raczna series). Lower Namurian isolated spores alone, then evidently Porostrobus (Flora series). is a later homonym of Triletes. Such is hardly Note. Evidently by mistake an illustration the case, however, because the identification of — of Triletes the genus Porostrobus and the species bennholdi horridus accompanied Nowak and Zerndt's original description. The species was within it depend on other features in addition validated by photographs published a later. to the spores. Some day we possibly may know year actually how diagnostic isolated spores of Porostrobus may be, but at present such speci- 18. Triletes brevispiculus Schopf, 1938, Illi- mens certainly must be recorded separately nois Geol. Survey Rept. Inv. 50, pp. 26-27, pi. under a different genus (as Zerndt has done). 1, figs. 13a-r; pi. 2, fig. 6; pi. 3, figs. 1-4. Consequently, also, these forms must be identi- Sectio, Aphanozonati. Age, Middle fied with reference to different nomenclatural Pennsylvanian, Carbondale series. types. The specimen illustrated by Zerndt's photograph 4, plate 2, and which evidently 19. Triletes circumtextus Zerndt, 1934, Acad. as the basis for his text figure 3 served (p. 6), polonaise sci. Trav. Geol. no. 1, pp. 19-21, is regarded as the holotype of the species which fig. 7; pi. 19, figs. 1-12; pi. 21, figs. 1-7. is here credited to Zerndt. Type 18 Zerndt, 1931, Acad, polonaise arises On the other hand, no question as to Bull, internat., ser. A, p. 173, pi. 6, figs. the spores Wicher (1934b) described in some 17, 18. detail that were isolated from undisputed speci- Age, Lower Namurian. mens of Porostrobus. There was no occasion Note.—Closely related to Triletes brasserti. for him to assign them a new generic name (Poro(strobo)sporites) since the only possible 20. Triletes clavatopilosus (Wicher) S. W. designation for the specimens is "spores of and B., comb. nov. Porostrobus bennholdi." Zerndt, in using the name "Porosporites" in 1938, is presumably Sporites clavatopilosus Wicher, 1934, following Wicher. Wherever evidence of addi- Inst. Palaobot. Arb., vol. 4, no. 4, p. 183, pi. figs. 25-26. tional cone characters warrants it, such a deter- 8, mination is more significant and essentially Type 24 Zerndt, 1931, Acad, polonaise useful than identification with a less closely sci. Bull, internat,, ser. B, pp. 174-175, pi. defined group such as Triletes. Mistakes of this 7, fig. 23. kind are less likely to occur if nomenclatural Type 14 Sahabi (?), 1936, Recherches types are regarded seriously. sur les spores des houilles Francaises, pp. 47 and 48, pi. 6, figs. 5-7. Age, Upper West- 14. Triletes bipilosus (Wicher) S. W. and B. phalian A—Lower Westphalian C. comb. nov. Sporites bipilosus Wicher, 1934, Inst. 21. Triletes crassiaculeatus (Zerndt) S. W. Palaobot. Arb., vol. 4, no. 4, p. 180, pi. and B., comb. nov. 8, figs. 13, 14. Lagenicula crassiaculeata Zerndt, 1937, 22 PALEOZOIC FOSSIL SPORES
Age, Westphalian Acad, polonaise sci. Trav. Geol. no. 3, pp. Sectio, Aphanozonati. 12-13. B, C, Lower Stephanian. Type 26A, Zerndt, 1937 (idem). Note.— Closely related to Triletes fulgens and Triletes reinschi. Sectio, Lagenicula. Age, Upper Dinan- tian (?). 30. Triletes greenlandicus (Miner) Harris, Trilctcs horridus. A^o^^.—Related to 1935, Meddelelser om Gronland, vol. 112,
no. 1, p. 155. 22. Triletes cyttarta Kendall, 1942, Annals Selaginellites greenlandicus Miner, 1932, and Mag. Nat. History ser. 11, vol. 9, p. Washington Acad. Sci. Jour., vol. 22, p. 922, fig. 2. 500, figs. 10, 11. Age, Lower Cretaceous. Age, Jurassic (Mid. Estuarine).
31. Triletes gymnozonatus Schopf, 1938, 23. Triletes dificilis (Wicher) S. W. and Illinois Geol. Survey Rept. Inv. 50, pp. 37- B., comb. nov. 38, pi. 1, fig. 4, pi. 4, fig. 9. Sporites dificilis Wicher, 1934, Inst Sectio, Triangulati. Age, Middle Penn- Palaobot. Arb., vol. 4, no. 4, p. 179, pi. 8 sylvanian, Carbondale series. figs. 17-18. Note.— Closely related to Triletes artecol- Type 14 (in part) Zerndt, 1931 Acad latus. polonaise sci. Bull, internat., ser. A, p. 172 pi. 4, figs. 11, 12, pi. 5, figs. 15, 16. 32. Triletes harrisi Murray, 1939, Geol. Mag., Zerndt, 1934, Acad, polonaise sci. Trav, vol. 76, p. 480, figs. 1, 2. Geol. no. 1, pp. 17-18, pi. 8, figs. 1-10, pi figs. 9. 31, 8, 33. Triletes hirsutus (Loose) S. W. and B., Sectio, Aphanozonati. Age, Lower comb. nov. Westphalian C. (Zerndt, Mid. Lower. Sporonites hirsutus, Loose, 1932, Neues Namurian to Westphalian D). Jahrb., Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 58. 24. Triletes difusopilosus (Wicher) S. W. Setosi-sporites hirsutus (Loose) Ibra- and B., comb. nov. him, 1933, Sporenformen des Aegirhori- Sporites difusopilosus Wicher, 1934, zonts, p. 26. Inst. Palaobot. Arb., vol. 4, no. 4, pp. 182- Apiculati - sporites hirsutus (Loose) 183, pi. 8, figs. 23, 24. Age, Lower West- phalian C. Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 153. Note.— Closely related to Triletes praetextus. Sporites hirsutus (Loose) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no, 25. Triletes echinatus (Miner) Harris, 1935, 4, p. 181. Meddelelser om Gronland, vol. 112, no. 1, p. 155. Sectio, Lagenicula?. Age, Westphalian Selaginellites echinatus Miner, 1932, B, Lower Westphalian C. Washington Acad. Sci. Jour., vol. 22, p. 34. 500, fig. 6. Age, Upper Cretaceous. Triletes horridus (Zerndt) S. W. and B., comb. nov.
26. Triletes erlansoni (Miner) Harris, 1935, Lagenicula horrida Zerndt, 1934, Acad,
Meddelelser om Gronland, vol. 112, no. 1, polonaise sci. Trav. Geol. no, 1, p. 25, fig. p. 155. 11, pi. 28, figs. 1-5. Selaginellites erlansoni Miner, 1932, Type 26 Zerndt, 1931, Acad, polonaise Washington Acad. Sci. Jour,, vol. 22, p. 500, sci. Bull, internat., ser. A, p. 175, pi, 9. fig. 28. figs. 1-3. Age, Upper Cretaceous. Sectio, Lagenicula. Age, Dinantian.
27. Triletes flavus Stach and Zerndt, 1931, Note.— Closely related to Triletes crassiacu- Gliickauf, Jahrg. 67, no. 35, p. 1122, fig. 18. l eat us. Sectio, Auriculati. Age, Westphalian C. 35. Triletes inornatus (Miner) Harris, 1935, 28. Triletes fulgens Zerndt, 1937, Acad, Meddelelser om Gronland, vol. 112, no. 1, polonaise sci. Trav. Geol. no. 3, p. 5, fig. 2, p. 155. pi. 1, figs. 1-9. Selaginellites inornatus Miner, 1932, Type 8 Zerndt, 1931, Acad, polonaise sci. Washington Acad. Sci. Jour., vol. 22, p. Bull, internat., ser. A, p. 171, pi. 3, figs. 505, figs. 7, 8. Age, Upper Cretaceous. 9-10. Sectio, Aphanosonati. Age, Middle Din- 36. Triletes ionthus Harris, 1935, Meddelel- antian to Middle Namurian. ser om Gronland, vol. 112, no. 1, pp. 166-167, ATo^^.—Closely related to Triletes glabratus 169, figs. 52E, F, G, pi. 26, fig. 8. (pars.). 37. Triletes kidstoni (Loose) S. W. and B., 29. Triletes glabratus Zerndt, 1930, idem, comb. nov. ser. B, p. 45, pi. 1, figs. 2, 3. Sporonites kidstoni Loose, 1932, Neues Type 9 Zerndt, 1931, idem., ser. A, p. Jahrb, Beilage-Band 67, Abt. B, p. 452, pi. 20, fig. 60. TRILETES 23
Laevigati-sporites kidstoni (Loose) 47. Triletes nigrozonalis Stach and Zerndt, Ibrahim, 1933, Sporenformen des Aegir- 1931, Gliickauf, Jahrg. 67, no. 35, p. 1123, horizonts, p. 19. figs. 26, 27. Reticulati-sporitcs kidstoni (Loose) Sporites nigrozonalis (Stach and Loose, 1934, Inst. Palaobot, Arb., vol. 4, no. Zerndt) Wicher, 1934, Inst. Palaobot. Arb., 3, pp. 156-7. vol. 4, no. 4, pp. 176-177, pi. 8, figs. 11 and Sectio, Aphanosonati. Age, Upper 12. Westphalian B, Lower Westphalian C. Sectio, Auriculati. Age, Westphalian C. 38. Triletes latihirsutus (Loose) S. W. and B., comb. nov. 48. Triletes nodosus (Wicher) S. W. and B., Apiculati-sporites latihirsutus Loose, comb. nov. 1934, idem., p. 153. Sporites nodosus Wicher, 1934, idem., Sectio, Lagenicula. Age, Upper West- p. 177, pi. 8, fig. 21. phalian B. Sectio, Aphanozonati. Age, Lower Note.— Closely related to Triletes robertianus. Westphalian C.
39. Triletes laximarginalis Zerndt, 1940, 49. Triletes nudus (Nowak and Zerndt) Paleontographica, vol. 84, Abt. B, p. 136, Schopf, 1938, Illinois Geol. Survey Rept. pi. 10, figs. 24-28. Inv. 50, p. 30, pi. 5, fig. 7. Type 17a, Zerndt, 1940 (idem). Lagenicula nuda Nowak and Zerndt, 1936, Acad, polonaise sci. Bull, Sectio, Triangulati. Age, Westphalian internat., ser. A, pi. fig. C, D (?). p. 60, 1, 6. Type 43 Zerndt, 1936 (idem). 40. Triletes levis (Zerndt) S. W. and B.. Sectio, Lagenicula. Age, Dinantian. comb. nov. Note.—The form recorded from lUinois as T. Lagenicula levis Zerndt, 1937, Acad, (?) nudus probably is not referable to this polonaise sci. Bull, internat., ser. A, pp. species although the shape is rather similar. 587-588, pi. 15, figs. 1-11. 50. Triletes ovalis Stach and Zerndt, 1931, Type 45 Zerndt, 1937 (idem). Gliickauf Jahrg. 67, no. 35, p. 1122, fig. 24. Sectio, Lagenicula. Age, Stephanian. Age, Westphalian B, Lower Westpha- Note.—Closely related to Triletes translucens, lian C. T. tenuimemhranosa and to T. rugosus. Note.—Probably closely related to type 5 of Sahabi (1936)., 41. Triletes litchi Harris, 1935, Meddelelser
om Gronland, vol. 112, no. 1, pp. 159, 160, 51. Triletes papillosus (Miner) Harris, 1935, fig. A-F, pi. 26, figs. 3, 10. Meddelelser om Gronland, vol. 112, No. Age, Liasso-Rhaetic. 1, p. 155.
42. Triletes mamillarius Bartlett, 1928, Selaginellites papillosus Miner, 1932, Washington Acad. Sci. Jour., vol. Michigan Acad. Sci., vol. 9, p. 21, pis. 13-15. 22, p. 500, fig. 9. Age, Upper Cretaceous. Sectio, Aphanozonati. Age, possibly Upper Pottsville or Lower ATleglieny. 52. Triletes parviapiculatus Zerndt, 1937, Acad, polonaise sci. Trav. Geol. no. 2, p. 43. Triletes mensura Harris, 1935, Meddelel- 17, pi. 24, figs. 1-4. ser om Gronland, vol. 112, no. 1, pp. 160- Sectio, Triangulatif. Age, Lower Mar- 162, fig. 53J, pi. 26, figs. 2, 9. ginal beds (Lower Namurian, vicinity of Age, Liasso-Rhaetic. Rybnik).
44. Triletes mirabilis (Miner) S. W. and B., 53. Triletes pedinacron Harris, 1935, Med- comb. nov. delelser om Gronland, vol. 112, no. 1, pp. Selaginellites mirabilis Miner, 1935, 165-166, fig. 52S, T, pi. 27, fig. 6. Am. Midland Naturalist, vol. 16, p. 618, pi. Age, Liasso-Rhaetic. 23, figs. 1-6. Age, Upper Cretaceous?.
54. Triletes persimilis Harris, 1935, idem., p. 45. Triletes mucronatus Nowak and Zerndt, 165, fig. 52R, pi. 25, fig. 4. 1936, Acad, polonaise sci. Bull, internat., ser. A, p. 59, pi. 1, figs. 1, 2. 55. Triletes pertuberosus (Loose) S. W. and Type 40 Zerndt, 1936 (idem). Age, B., comb. nov. Dinantian. Sporonites pertuberosus Loose, 1932, Note.—Possibly closely related to Triletes Neues Jahrb., Beilage-Band 67, Abt. B, p. clavatopilosus. 452, pi. 20, fig. 57. Tuberculati-sporites pertuberosus, 46. Triletes myrmecoides Harris, 1935, Med- (Loose) Loose, 1934, Inst. Palaobot. Arb., delelser om Gronland, vol. 112, no. 1, fig. vol. 4, no. 3, p. 147, pi. 20, fig. 57. 52M-Q, pi. 26, fig. 4. Sectio, Aphanozonati. Age, Upper
' ' ' Age, Liasso-Rhaetic. Westphalian B. , 24 PALEOZOIC FOSSIL SPORES
A, Note.—Equal to Type 14 Zerndt (1931, Type 10 Zerndt, 1931, idem., ser. p. 1934) in part. 172. Triletes I Kidston, Stach and Zerndt, 1939, Geol. 56. Triletes phyllicus Murray, 1931, Gluckauf, Jahrg. 67, no. 35, p. 1122. Mag., vol. 76, p. 482, figs. 7, 8. Triletes Type 1 Kidston, Maslankie- Age, Jurassic (Estuarine). wiczowa, 1932, Acta soc. Bot. poloniae, vol. 9 (Suppl.), p. 158. 57. Triletes pinguis Harris, 1935, Meddelel- Sporonites reinschi Ibrahim, 1932, ser om Gronland, vol. 112, no. 1, p. 166, fig. Jahrb., Beilage-Band 67, Abt. B, p. 52A-D, pi. 25, fig. 3. Neues 449, pi. 17, fig. 28. Age, Liasso-Rhaetic. Laevigati-sporites reinschi (Ibrahim) 58. Triletes polysceles Murray, 1939, Geol. Ibrahim, 1933, Sporenformen des Aegir- Mag., vol. 76, p. 484, figs. 5, 6. horizonts, p. 18, pi. 4, fig. 28. Age, Jurassic (Estuarine). Reticulati-sporites reinschi (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, 59. Triletes praetextus Zerndt, 1934, Acad, no. 3, p. 157. polonaise sci. Trav. Geol. no. 1, p. 24, pi. Sporites primus Wicher, 1934, idem, 26, figs. 1-6, pi. 27, figs. 1-6, Text-fig. 10. no. 4, p. 169. Type 21 Zerndt, 1931, Acad, polonaise Types 1 to 6 incl., Rousseau, 1935, Mus. sci. Bull, internat., ser. A, p. 174, pi. 8, figs. royale histoire nat. Belgique Bull. vol. 11, 24, 25. no. 21, pp. 3-4, pi. 1, figs. 1-5, pi. 2, fig. 6. Age, Upper Dinantian and Lower Na- 1 spores des murian. Type Sahabi (pars.), 1936, houilles Francaises, p. 34, pi. 1, figs. 1, 2.. Note.—Closely related to Triletes difuso- pilosus. Sectio, Aphano3onati. Age, Westphalian C & D, Lower Stephanian. De- 60. Triletes potosiensis Zerndt, 1938, Note.—Type species of Triletes by designa- Carbonifere, vol. uxieme Cong. Stratig. 3, tion (Schopf, 1938). p. 1728, pi. 155, fig. 4. Sectio, Auriculatif. Age, Carbonifer- 64. Triletes reticulatus Zerndt, 1938, De- uxieme Cong. Stratig. Carbonif., vol. 3, p. 1728, pi. 155, fig. 5. 61. Triletes radiatus (Ibrahim) S. W. and B., Age, Mesozoic. comb. nov. Sporonites radiatus Ibrahim, 1932, 65. Triletes rexargenteus Harris, 1935, Med- Neues Jahrb, Beilage-Band 67, Abt. B, p. delelser om Gronland, vol. 112, no. 1, pp. 449, pi. 16, fig. 25. 156-158, fig. 51G-J, pi. 26, fig. 13. Zonales-sporites radiatus (Ibrahim) Age, Liasso-Rhaetic. Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 29, pi. 3, fig. 25. 66. Triletes richardsoni Murray, 1939, Geol. Sectio, Triangulati. Age, Aegir coal, at Mag., vol. 76, pp. 482-484, figs. 9, 10. boundary of Westphalian B and C. Age, Jurassic (Estuarine).
62. Triletes radiosus S. W. and B. nom. nov. 67. Triletes robertianus S. W. and B., nom. Triletes radiatus Zerndt, 1937, Acad, nov. polonaise sci. Trav. Geol. no. 3, p. 10, fig. Triletes kidstoni Zerndt, 1934, Acad.
7, pi. 13, figs. 1-5. polonaise sci. Trav. Geol. no. 1, pp. 26-27, Types 22 and 23 Zerndt, 1931, Acad, fig. 12, pi. 28, figs. 6-11, pi. 29, figs. 1-13. polonaise sci. Bull, internat., ser. A, p. 174, Type 27 Zerndt, 1931, Acad, polonaise pi. figs. 25. 8, 24, sci. Bull, internat., ser. A, p. 175, pi. 9, Age, Lower Namurian (Upper Di- figs. 29-32. nantian?). Sectio, Lagcnicula. Age, Dinantian, '" — Note. T. radiatus Zerndt, 1937, becomes a Namurian, Westphalian A. later homonym in conflict with Sporonites radi- Note. Closely related to Triletes bipilosus atus Ibrahim when the latter is transferred to — and to T. latihirsutus. The new name is neces- Triletes (see sp. 61 above), thus necessitating sitated by preoccupation of the specific epithet a new name as here proposed. (species no. 2t7). This species is likewise named 63. Triletes reinschi (Ibrahim) Schopf, 1938, for Robert Kidston. Illinois Geol. Survey Rept. Inv. 50, p. 24, 68. Triletes rotatus Bartlett, 1928, Michigan pi. 2, figs. 2-4, pi. 5, figs. 8, 9. Acad. sci. Papers, vol. 9, p. 21, pis. 9, 10, Triletes I Bennie and Kidston, 1886, 11, 12. Royal Phys. Soc. Edinburgh Proc, vol. Type 19, Triletes rotatus Bartlett, 9, p. 107, pi. Ill, figs, la, b. Zerndt, 1934, Acad, polonaise sci. Spore 1.9 mm. Zerndt, 1930, Soc. Geol. Trav. Geol. no. 1, pp. 21-22, fig. 8, pi. 24, figs. 1-6. Pologne, Ann. vol 6, p. 308, 312, pi. 1, figs. 5a, b, pi. Ill, figs. 5a, b. Triletes rotatus Bartlett, Zerndt, 1937, Triletes Type 1 Kidston, Zerndt, 1930, idem., no. 3, pp. 8-10, pis. 6-10. i- Acad, polonaise sci. Bull, internat., ser. B, Age, type material probably upper p. 43, pi. 1, fig. 1. Pottsville. TRILETES 25
Triletes rotatus var, denticulata Zerndt, 74. Triletes silvanus (Ibrahim) S. W. and 1937, Acad, polonaise sci. Trav. Geol. no. B., comb. nov. 3, pp. 9-10, fig. 6 (?),pl. 6, fig. 3. Sporonites silvanus Ibrahim, 1932
Age, Dinantian, Lower Namurian and Neues Jahrb., Beilage-Ba*^d 67, Abt. B, p, above (?). 448, pi. 15, fig. 22. Laevigati - sporites silvanus Ibrahim, (Loose) Schopf, 1938, 69. Triletes rugosus 1933, Sporenformen des Aegirhorizonts, p Inv. 29- Illinois Geol. Surv. Rept. 50, pp. 20, pi. 2, fig. 22, pi. 6, fig. 47. 30, pi. 5, fig. 6. Sporites silvanus (Ibrahim) Wicher, . Sporonites rugosus Loose, 1932, Neues 1934, Inst. Palaobot. Arb., vol. 4, No. 4, p B, 452, Jahrb., Beilage-Band 67, Abt. p. 174, pi. 8, figs. 5-8. pi. 20, fig. 59. Sectio, Auriculati. Age, Upper West- Levigati-sporites rugosus (Loose) Ibra- phalian B, Lower Westphalian C. Aegirhori- him, 1933, Sporenformen des Note.— Closely related to Triletes auritus. zonts, pp. 18-19, pi. 7, fig. 65. Punctati - sporites rugosus (Loose) 75. Triletes simplex (Zerndt) S. W. and B., Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. comb. nov. 3, p. 146. Lagenicula simplex Zerndt, 1937, Acad, Sectio, Lagenicula. Age, Upper West- polonaise sci. Trav. Geol. no. 3, pp. 14-15, phalian B and above (?). fig. 10, pi. 23, figs. 1-5. Note.—May be closely allied with Triletes Type 35 Zerndt, 1937, idem. translucens. Sectio, Aphanozonati (?). Age, Lower Namurian. Zerndt, 1941, Paleonto- 70. Triletes saarensis Triletes simplex var. levis Zerndt, 1937, graphica, Abt. B, vol. p. 134, pi. 12, figs. 84, idem., p. 15. 40-44. Type 36 Zerndt, 1937, idem. Type 47 Zerndt, 1941 (see above refer- ence). 76. Triletes sofiaense S. W. and B., nom. nov. Sectio, Auriculati (?). Age, West- Triletes auritus III Maslankiewiczowa, phalian C and Lower Westphalian D. 1932, Acta Soc. Botanicarum Poloniae, vol. 9 (Suppl.) pp. 161-163, 172-173, figs. Z7, 38. 71. Triletes saturnoides (Ibrahim) S. W. Sectio, Auriculati. Age, Upper West- and B., comb. nov. phalian B (Laziska series). Sporonites saturnoides Ibrahim, 1932, Note.—The group is clearly distinguished Neues Jahrb., Beilage-Band 67, Abt. B, p. specifically by coarse reticulation, in this respect 449, pi. 16, fig. 26. somewhat similar to Triletes tuberculatus, Zonales-sporites saturnoides (Ibrahim) though in shape it resembles Triletes auritus. Ibrahim, 1933, Sporenformen des Aegir- Possibly it represents a connecting link between these species. It is horizonts, p. 27-28, pi. 3, fig. 26. named for Zofja Kowalew- ska Maslankiewiczowa who provided an ample Age, Upper Westphalian B. description of the form. Note.—Allied with Triletes brasserti, T, superbus and T. circumtextus. 77. Triletes sparassus Murray, 1939, Geol. Mag., vol. 76, p. 480, p. 482, figs. 3, 4. 72. Triletes sextus S. and B., nom. nov. W. Age, Jurassic (Estuarine). Triletes VI Bennie and Kidston, 1886, Royal Physical Soc. Edinburgh Proc, vol. 78. Triletes splendida (Zerndt) S. W. and B., 9, p. 109, pi. 3, figs. 6a, 6b, 6c. comb. nov. Sectio, Aphanosonati. Age, Lanarkian=: Lagenicula splendida Zerndt, 1937, Namurian. Acad, polonaise sci. Trav. Geol. no 3, pp. Note.—Forms of this sort are included by 13-14, pis. 18-20. Zerndt (1931, 1934) in Type 14. Inasmuch_ as Type 28 Zerndt, 1931, Acad, polonaise this was used as the type for Apiculati-sporites sci. Bull, internat., Ser. A, p. 175, pi. 10, Ibrahim, it seems desirable to have an orthodox figs. 36, 37. designation for reference. Sectio, Aphanosonati. Age, Upper Di- nantian, Lower Namurian. 7?). Triletes sextusdecimus S. W. and B. nom. A^o^^.— lageniculate nov. The vestibule possessed by this form is insufficient basis for assigning Triletes XVI Bennie and Kidston, 1886, it to Lagenicula in disregard of its other charac- idem. 113-114, pi. figs. 16a, 16b, 16c, pp. 5, teristic Aphanosonati features. 16d, 16e.
Age, Bernician and Lanarkian = ap- 79. Triletes stenoxysmatodes Harris, 1935,
prox. Upper Dinantian and Namurian. Meddelelser cm Gronland, vol. 112, no. 1, Note.—Related to Triletes superbus, and prob. p. 167, pi. 25, fig. 6. to T. circumtextus and T. brasserti. This form Age, Liasso-Rhaetic. was taken as the type oi Zonales-sporites Ibra- him, hence it seems desirable to propose an 80. Triletes subfuscus (Wicher) S. W. and orthodox name for purposes of reference. B., comb. nov. 26 PALEOZOIC FOSSIL SPORES
tenuispinosus var. secundus S. Sporitcs subfnscns Wicher, 1934, Inst. Triletes pi. W. and B., var. nom. nov. Palaobot. Arb., vol. 4, no. 4, p. 179, 8, Triletes tenuispinosus var. II Zerndt, fig. 20. 1937, idem., no. pp. 7-8, fig. 5, pi. 3, figs. Type 14 Zerndt (in part), 1934, Acad, 3, 3 and 4. Age, Dinantian — Upper and Trav. Geol. no. 1, pp. 17, 18, polonaise sci. Lower Namurian. fig. pis. 8-17. 5, Note.—Zerndt describes three forms, one Sectio, Aphanosonati. Age, Lower which evidently is typically the species and two C. Westphalian varieties, only one of which is properly named. Harris, The new varietal name proposed provides a 81. Triletes subrotundus (Miner) means of reference. 1935, Meddelelser om Gronland, vol. 112, 165. no. 1, p. 155, p. 88. Triletes translucens Schopf, 1938, Illi- Selaginellites subrotundus Miner, 1932, nois Geol. Survey Rept. Inv. 50, pp. 28-29, 505, Washington Acad. Sci. Jour., vol. 22, p. pi. 5, figs. 3-5. figs. 4, 5. Sectio, Lagenicula. Age, Middle Penn- Age, Upper Cretaceous. sylvanian, upper Carbondale, lower Mc- Leansboro series. subtilinodulata (Nowak and 82. Triletes Note.—Closely related to Triletes tenuimem- and B., comb. nov. Zerndt) S. W. branosa, T. levis, and (?) T. rugosus. Lagenicula subtilinodulata, Nowak and Zerndt, 1930, Acad. Zerndt, 1936, Acad, polonaise sci. Bull, in- 89. Triletes triangulatus sci. Bull, internat, ser. B, 51, ternat, ser. A, pp. 59-60, pi. 1, fig. 7. polonaise p. pi. figs. 19-24. Type 42 Zerndt, Nowak and Zerndt, 7, Spore 0.5 mm. Zerndt, 1930, Soc. Geol. 1936 (idem). Pologne Ann., vol. 6, pp. 306, 312, pi. 1, Sectio, Lagenicula. Age, Dinantian. figs, la, b, c, pi. 2, figs, la, b. 17 Zerndt, 1931 Acad, polonaise 83. Triletes subpilosus (Ibrahim) S. W. and Type sci. A, 173. B., comb. nov. Bull, internat. ser. p. Setosi-sporites subpilosus Ibrahim, Sporonites regalis Ibrahim, 1932, Neues Jahrb., Beilage-Band Abt. B, p. pi. 1933, Sporenformen des Aegirhorizonts, p. 67, 449, 16, fig. 24. 27, pi. 5, fig. 40. "var. regali/' Sectio, Lagenicula. Age, Aegir coal, Sporites triangulatus boundary of Westphalian B and C. (Zerndt) Ibrahim, 1933, Sporenformen des Aegirhorizonts, pi. fig. 24. Note.—May be closely related to Triletes p. 29, 3, robertianus. Sporites regalis (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149, 84. Triletes superbus Bartlett, 1928, Michigan pi. 7, fig. 34. Acad. Sci. Papers, vol. 9, pp. 20, 21, pi. 7, Sporites triangulatus (Zerndt) Wicher, 2. figs. 1, 2, pi. 8, figs. 1, 1934, idem., vol. 4, no. 4, p. 175. Age, Probably upper Pottsville. Triletes triangulatus Zerndt, 1934, Note.—Probably allied with Triletes circum- Acad, polonaise sci. Trav. Geol. no. 1, p. textus and T. brasserti. 19, fig. 6, pi. 18, figs. 1-24. Triletes triangulatus Zerndt, Schopf, 85. Triletes tenuigollatus Nowak and Zerndt, 1936, Illinois Acad. Sci., Trans, vol. 28, p. 1936, Acad, polonaise sci. Bull, internat., 107, fig. 6. ser. A, p. 59, pi. 1, fig. 3. Type X Sahabi, 1936, spores des houil- Type 41 Zerndt, Nowak and Zerndt, les Francaises, p. 43, fig. 11, pi. 4, figs 6-11. 1936 (idem). Triletes triangulatus Zerndt, 1937, Age, Dinantian. Acad, polonaise sci. Trav. Geol. no. 3, pi. 5, figs. 1-5. 86. Triletes tenuimembranosa (Zerndt) S. Triletes triangulatus Zerndt, W. and B., comb. nov. Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, Lagenicula tenuimembranosa Zerndt, pp. 32-37, pi. 1, figs. 7, 8, pi. 4, figs. 1-7, pi. 1937, idem. p. 587, pi. 14, figs. 1, 2. 7, figs. 5, 6. Sectio, Lagenicula. Age, Westphalian Triletes triangulatus Zerndt, 1940, C and Lower Westphalian D. Paleontographica, vol. 84, Abt. B, pi. 10, Type 25 Zerndt, 1931, idem., p. 175, pi. figs. 19-23. 9, figs. 2>?> and 35. Triletes triangulatus var. zonatus Sectio, Lagenicula. Age, Lower Na- (Ibrahim) Schopf, 1938, Illinois Geol. Sur- murian and Westphalian A. vey Rept. Inv. 50, p. 34. Note.—Closely related to Triletes translucens. Triletes triangulatus II Zerndt, 1930, Acad, polonaise sci. Bull, internat., ser. B, 87. Triletes tenuispinosus Zerndt, 1934, p. 53, pi. 7, figs. 25-30. Acad, polonaise sci. Trav. Geol. no. 1, p. Sporonites zonatus Ibrahim, 1932, 16, text-fig. 4, pi. 7, figs. 1-15. Neues Jahrb., Beilage-Band 67, Abt. B, p. Type 13 Zerndt, 1934 (idem). 448, pi. 16, fig. 23. Triletes tenuispinosus var. brevispinosa Zonales-sporites triangulatus secundus Zerndt, 1937, idem., no. 3, p. 6, fig. 4, pi. 3, Ibrahim, 1933, Sporenformen des Aegirhor- figs. 2-7. izonts, p. 30, pi. 3, fig. 23, pi. 7, fig. 64. : :
PITYOSPORITES 27
Zonales-sporites zonatus (Ibrahim) 94. Triletes tylotus Harris, 1935, Meddelelser Loose, 1934, Inst. Palaobot. Arb., vol. 4, om Gronland, vol. 112, no. 1, pp. 162-163, fig. 53F-I, pi. 26, figs. 1, 12. no. 3, p. 150, pi. 7, fig. 31. Spontes triangulatus var. sonatus (Ib- Age, Liasso-Rhaetic. 1934, Inst. Palaobot. Arb., rahim) Wicher, 95. Triletes valens (Wicher) S. W. and B., vol. 4, no. 4, pp. 175-176. comb. nov. var.? Triletes triangulatus Sporites valens Wicher, 1934, Inst. triangulatus III, Stach and Triletes Palaobot. Arb., vol. 4, no. 4, p. 178, pi. 8, no 35, Zerndt, 1931, Gliickauf, Jahrg. 67, fig. 19. p. 1123, figs. 32, 2,Z. Sectio, Aphanosonati. Age, Lower upper Lower Sectio, Triangulati. Age, Westphalian C. Namurian through Westphalian D. Triletes nomen excludende 1. Apiculati-sporites megaspinosus Ibrahim, Sporen- tricollinus Zerndt, 1938, Deux- 1933, 90. Triletes formen des Aegirhorizonts, etc., p. 24, pi. 8, fig. 69. 1713, ieme Cong. Stratig. Carbonif. vol. 3, p. Note: This form is not congeneric with the type of pi. 155, fig. 3. Apiculati-sporites {Triletes sextus) and, although it ap- (idem). Earently is sufficiently characterized and characteristic to Type 44 Zerndt, 1938 e a useful microfossil, no further generic assignment will Sectio, Triangulati. Age, Westphalian be attempted. B, C, and D. Genus Pityosporites Seward, 1914 91. Triletes trilobus (Ibrahim) S. W. and B., Plate 2, figures 15-15b comb. nov. Sporonites trilobus Ibrahim, 1932, Diagnoses given by Seward are as Neues Jahrb., Beilage-Band 67, Abt. B, p. follows 449, pi. 17, fig. 30. The generic designation Pityosporites is pro- Valvisi-sporites trilobus (Ibrahim) Ib- posed for winged spores agreeing in form and rahim, 1933, Sporenformen des Aegirhor- size with recent Abietineous genera (1914, p. fig. 30. izonts, p. 33, pi. 4, 23). trilobus (Ibrahim) Ib- Valvisi-sporites .... Spores provided with bladder-like Loose, 1934, Inst. Palaobot. Arb., rahim, extensions of the exine, agreeing in size and 30. vol. 4, no. 3, p. 152, pi. 17, fig. form with those of recent Abietineous genera Auriculati. Age, Upper West- Sectio, (1919, p. 398). phalian B. A more adequate definition inferred Note.—Possibly very closely related to Tri- from the above, and from consideration letes appendiculatus. Wicher (1934) regards it as similar to T. auritus. of the species included here, is as follows Symmetry.—Pollen grains appearing Acad, 92. Triletes tuberculatus Zerndt, 1930, bilateral, due to the presence of bladders polonaise sci. Bull, internat. ser. B, pp. 47-51, placed on opposite sides. The funda- pi. 2, figs. 6, 7, pi. 3, figs. 8, 9, pi. 4, figs. the 10,' 11, pi. 5, figs. 12, 13. mental symmetry, shown by body but Triletes XIX Kidston, 1890, Royal Soc. modified, is radial, as this pollen arises Edinburgh Trans., vol. 36, pi., figs. 9-11. from tetrahedral tetrads. Type 16 Zerndt, 1931, Acad, polonaise Shape—Grains elliptical as viewed sci. Bull, internat., ser. A, p. 173. Triletes tuberculatus Zerndt, Maslan- from proximal side, the diameter of the klewiczowa, 1931, Acta Soc. Bot. Poloniae, poles of the ellipse (formed by the two vol. 9, Suppl. pp. 165-168, figs. 40, 41. bladders) being broader than that of the Sectio, Auriculati. Age, Westphalian body between them. Body appears ap- B, C, Stephanian? proximately circular in proximal view.
93. Triletes tuberosus (Ibrahim) S. W. and Viewed from either side the bulbose blad- B., comb. nov. ders are inclined distally and a broad Sporonites tuberosus Ibrahim, 1932, sulcus appears between them on the distal Beilage-Band Abt. B, Neues Jahrb., 67, p. side. In lateral outline the body appears 449, pi. 16, fig. 27. broadly oval with the smaller pole toward Tuberculati-sporites tuberosus (Ibra- him) Ibrahim, 1933, Sporenformen des the distal sulcus. Aegirhorizonts, p. 23, pi. 3, fig. 27. Size.—Various species range in total tuberosus ( Ibra- Tuberculati-sporites length (bladder tip to bladder tip) from him) Ibrahim, Loose, 1934, Inst. Palaobot. as small as 40 to over 100 microns. Arb., vol. 4, no. 3, p. 147, pi. 7, fig. 37.^ Sporites tuberosus (Ibrahim) Wicher, Ornamentation.—External bladder sur- 1934, idem, vol. 4, no. 4, pp. 177-178, pi. 8, face minutely granulose or rugose ; in- fig. 22. ternally the bladders are usually at least Sectio, AphanoBonati. Age, Upper weakly reticulate. Westphalian B, Lower Westphalian C. Proximal cap may be nearly smooth, rugose or striate. Distal Note.—Probably the same as some of Zerndt's Type 14. body wall least ornamented. 28 PALEOZOIC FOSSIL SPORES
Haptotypic features.—Lacking or near- aspects of these fossils which have been ly so. Presumably a weak vestigial imprint more precisely itemized in our revised of the trilete mark may be present on the definition.' A more restricted group is proximal cap in some forms as it oc- thus indicated than that considered by casionally is in modern Abies. Florin, and this group surely is not an Pollen coat.—The unique character of artificial one although it certainly exceeds internal bladder wall reticulation distin- the scope of most "normal" genera, and, guishes these and many other coniferous in fact, possibly contains elements which pollen grains. The internal reticulae add would be classed in separate families if apparent thickness to the perineal blad- more complete characterization could be der membrane which is quite thin. The obtained. Nevertheless, we think it neces- exinal (body) wall is much thicker sary to treat the group as a genus at proximally and is least translucent, present, with the hope that critical studies sometimes appearing brownish under the later will permit it to be subdivided into somewhat smaller groups that will more microscope ; the distal wall is much thin- ner, serving as a harmomegathus and perfectly represent the relationships in point of germinal exit when ruptured by detail. Whether this hope is a vain one the expanding gametophyte. cannot be foretold a priori, but the desira- bility of recognizing a type species for Affinity.—There can be little question this group which, as we construe it, does that grains of this sort are referable to signify a definite natural plant alliance, the Coniferae. Members of modern cannot be denied. Part Podocarpaceae and Pinaceae show essen- of the taxo- nomist's responsibility is to place the sys- tially similar features. Correlatives of tematic data in order so that later students species of Pityosporites represented by may revise it to greater perfection in the vegetative organs are hardly recognized in Paleozoic formations. The presence of light of additional data. Such progress is very difficult if nomenclatural types are such pollen is evidence that such plants not definitely indicated existed, possibly as a xerophytic upland and fully utilized in the application of technical flora, and tend to substantiate the view nomencla- long held that an important ancestry, very ture. In paleobotanical studies it is scantly represented among megascopic particularly important that this be recog- fossils, preceded those highly developed nized because the nature of the material forms found better represented in younger that is being studied rarely permits cate- strata. gorical statement and new discoveries call for greater revision in preexisting con- Pityosporites represents a group of cepts, perhaps, than in any other plant Coniferae whose pollen is essentially mod- science. ern in aspect. Florin's brilliant researches (1938-1940) have sufficiently demon- Pityosporites can hardly be considered strated that Lebachia, Ernestiodendron, as synonymous with any modern genus; and Walchianthus for the most part do perhaps the most important reason is that not possess pollen of the Pityosporites when pollen characters are sufficiently type ; most, if not all of the probably distinctive to permit actual identification correlated but isolated pollen of this sort with a more precisely delimited modern is referable to Florinites n. gen. described group, identification with the genus Pityo- below (p. 56). sporites becomes inadequate as an ex- Remarks.—It will be evident that the pression of affinity. Quite evidently many interpretation given Pityosporites here of the isolated ancient pollen grains can- differs from that of Florin. He (1940, not be more definitely assigned and Pityo- pp. 327-8) considers Pityosporites to be sporites will serve a most useful function a completely artificial genus which has in these instances. no type species. Apparently he would assign any pollen-like form with two ^Although Seward (1919, pp. 398-9) has included one bladders to this group. It is evident from of the forms Nathorst reported from the Hor clay as "Pityosporites sp." (which we should prefer Seward's descriptions, however to assign (1914, to Alisporites) , Seward apparently did this on the as- sumption that Nathorst's specimen was similar pollen 1919, 1933), that the emphasis is not to of Picea excelsa. Wodehouse (1935) has shown Picea so much merely on the presence of two pollen to be very different. The Pityosporites sp. of Solms from Franz Josef Land (Seward, 1919, p. 399) no doubt bladders as it is on the essentially modern is correctly assigned. ;
PUNCTATI-SPORITES 29
In addition to the nine species listed be- Genus Punctati-sporites (Ibrahim, low, Kosanke (1943) also has recorded 1933) emend., S. W. and B., material of this nature from the Upper Plate 1, figures 4-4b Pennsylvanian of Ohio. Symmetry.—Spores radial, trilete. 1. PiTYOSPORiTES ANTARCTicus Scward, 1914, Nat. Hist., Report British Antarctic (Terra Shape.—Originally nearly spherical, or Nova) Exped. 1910. Geology vol. 1, no. 1, possibly broadly rounded triangular with pp. 23-4, pi. 8, fig. 45. slight shortening of the axial dimension; Pityosporites antarcticus Seward, 1919, Fossil Plants, vol. IV, p. 398. when compressed the spores show no Pityosporites antarcticus Seward, 1933, proximo-distal orientation preference. New Phyt, vol. 32, no. 4, pp. 311-313, fig. 1. Size.—Spores of various species range Age, "Not older than Lower Meso- from 45 to 85 microns in mean diameter.
Ornamentation. — Various ; surfaces 2. Pityosporites chinleana Daugherty, 1941, levigate to punctate, rugose, reticulate or Carnegie Inst. Washington, Pub. 526, p. mildly apiculate. 398, pi. 34, fig. 5. Age, Upper Triassic. Haptotypic structures. — Of moderate prominence, relative length of trilete rays 3. Pityosporites jeffreyi Florin, 1940, (?) highly variable between different species Paleontographica vol. 85, Abt. B, no. 5, p. equatorial 327, pi. 163-4, figs. 8-12. no or arcuate markings (con- Pityanthus jeffreyi Florin (nomen necting ends of trilete rays) are present; nudum?), 1927, Arkiv. for Botanik, vol. lips of the trilete commissure are never 21 A, no. 13, p. 6. (Systematic status of very prominent nor highly ornamented. this nomenclatural combination is difficult to decide, since illustrations needed to validate Spore coat.—Generally thin, and, ex- it were not published till 1940 when the cept in instances where ornamental fea- species was transferred to Pityosporites.) tures are most prominent, hardly in excess Age, Middle Stephanian. of 3 microns. Note.—BVadders are seemingly set nearly op- posite though it is possible their distal inclina- Affinity.—Spores of this genus are tion merely is not apparent in the figures. The similar to those obtained from certain proximal cap also is not evident and it may pteridospermic fructifications and from be this form belongs to Alisporites rather than in the present genus. some fossils assigned to the ferns. The spores Kidston (1906) obtained from va- 4. Pityosporites pallidus Reissinger (nomen rious Crossotheca fructifications are char- nudum), 1939, Paleontographica, vol. 84, acteristic of typical species of Punctati- Abt. B, p. 14. Age, Lowest Jura (Lias). sporites. However, many of the species Note.—An illustration required to validate represented by isolated spores have no the name has not yet been published. known pteridospermic relationship al- is 5. Pityosporites sewardi Virkki, 1937, In- though there no adequate basis for dian Acad. Sci. Proc, vol. 6, no. 6, sect. B, separating them from forms which are pp. 428-431, figs. 2a, b, c, d, pi. 32, figs, la, thus allied. Much remains to be discov- b, c. ered as to the affinities of this genus, Age, Lower Gondwana, above Talchir which is a rather large one, and it boulder bed. may be expected that some diverse elements 6. Pityosporites stephanensis Florin, 1940, have been included which will be segre- Paleontographica, vol. Abt. B, no. 5. 85, gated on the basis of newer information. pp. 327-8, pi. 163-4, figs. 13-16. is of the Age, Middle Stephanian. It quite possible that some forms assigned to Punctati-sporites are 7. Pityosporites sp. Florin, 1940, idem, p. 61, merely immature and lack their distinc- pi. 163-4 fig. 17, fig. 18? tive mature ornamentation. Such may be Age, Lower Rothliegendes. the case for immature forms that in 8. Pityosporites (?) sp. Florin, 1940, idem, reality belong to Raistrickia n. gen. de- p. 62, pis. 165-6, fig. 20. scribed below (p. 55). Immature forms Age, Upper Carboniferous. should not be arbitrarily assigned to Note.—No inclination of bladders shown. Punctati-sporites but it is well to recog- 9. Pityosporites sp. Daugherty, 1941, Car- nize that when these are present in sub- negie Inst. Wash., Pub. 526, description of pi. 34, figs. 7 and 8. stantial numbers, they need to be reported Age, Keuper. and that a superficial resemblance to 30 PALEOZOIC FOSSIL SPORES
P unciati-spontes may not always be in- Six species previously assigned to Ver- dicative of relationship. The affinity of rucosi-sporites Ibrahim, including the certain of the species with the Crossotheca type, are here placed under Punctati-spor- alliance of pteridosperms seems beyond ites. Verrucosi-sporites thus would more much question, however. reasonably be taken as the designation The forms which Raistrick (1933, for the genus except that in meaning one name is no more suitable than the other 1934, 1937) has assigned to his types and strict interpretation of page priority B„ Be, D3, D„ D,5 (Knox, 1938), E„ Ee, 21 vs. favors Punctati-sporites. E„ E„ Eg, F„ Fs (Knox, 1938) (also (p. p. 25) The genus also includes the type of 1K( ?j, 2K and 7K, Knox, 1942) appear Apiculata-sporites Ibrahim a entirely or in part referable to Punctati- (1933), monotypic genus which seems based en- sporitcs. It also may include some of tirely on Ibrahim's inability to observe the forms he has classed as Bg and Bg. the trilete sutures described by Loose Some difficulty is encountered because Raistrick evidently has designated no sin- (1932) on Sporonites spinulistratus. The of the trilete on this gle typical example for these groups and presence marking reaffirmed in 1934. his illustrations of the forms have differed species was by Loose for individual types beyond what may In any event the genus Apiculata-sporites represent a specific range of variability. does not seem securely founded and its Thus there may be some question as to one species is here placed under Punctati- the consanguinity of forms labeled as D3 sporites. Both groups, Verrucosi-sporites (cf. also Raistrick, 1935). Figures and Ibrahim and Apiculata-sporites Ibrahim, description of E^ given in 1933 do not ex- conseqently are regarded as congeneric actly correspond with that given in later with and the names synonyms of Punctati- papers (1934, 1935). Illustrations of sporites. B3 given in 1933 seem to resemble Punc- The holotype of the type species, Punc- tati-sporites but those illustrated in later tati-sporites punctatus (Ibrahim) Ibra- papers without much doubt are referable him, is from the Aegir coal seam at the to Calamospora n. gen. described below. top of the Westphalian B in the Ruhr Since Raistrick's procedure is non-tax- district of Western Germany. onomic it is impossible to form more than a general opinion as to specific equiv- 1. Punctati-sporites aculeatus (Ibrahim) S. W. and B., comb. nov. alences in most instances. Apiculati-sporites aculeatus Ibrahim, genus Punctati-spontes Remarks.—The 1933, Sporenformen des Aegirhorizonts, p. as given here includes twenty-nine species, 23, pi. 6, fig. 57. four of which are queried. All but three 2. Punctati-sporites aureus (Loose) S. W. of these names (see spp. Nos. 9, 21, 22) and B., comb. nov. are new combinations. The group Punc- Reticulati-sporites aureus Loose, 1934, tati-sporites was originally defined by Inst. Paleobot. Arb., vol. 4, no. 3, p. 155, Ibrahim (1933, p. 21) as including trilete pi. 7, fig. 24. type spores with a granular surface, char- acters which by themselves can hardly 3. Punctati-sporites (?) auriculaferens (Loose) S. W. and B., comb. nov. be accepted as diagnostic of any generic Sporonites group. Three species were described, one auriculaferens Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. of which, P. punctatus, was designated as 450, pi. 18, fig. 39. the type. Consequently it has been pos- Valvisi-sporites auriculaferens (Loose) sible to define the genus practically, only Loose, 1934, Inst. Palaobot. Arb., vol. 4, by segregating those forms which ap- no. 3, p. 152. peared congeneric with P. punctatus, and 4. Punctati-sporites eucculentus (Loose) the emended generic diagnosis has been S. W. and B., comb. nov. constructed on this basis. Neither of the Verrucosi-sporites bucculentus Loose, other two species Ibrahim originally 1934, idem. p. 154, pi. 7, fig. 15. placed in the genus are considered closely 5. Punctati-sporites corrugatus (Ibrahim) enough related to remain there ; P. parvus S. W. and B., comb. nov. Ibrahim is now assigned to the genus Granulati-sporites, and P. pallidus Reticulati-sporites corrugatus Ibrahim, 1933, Sporenformen des Aegirhorizonts, (Loose) pp. Ibrahim to Calamospora n. gen. 35-36, pi. 5, fig. 41. PUNCTATI-SPORITES 31
Tuberculati - sporites microtuberosus 6. PUNCTATI-SPORITES CUSUS (LoOSC) S. W. des and B., comb. nov. (Loose) Ibrahim, 1933, Sporenformen Aegirhorizonts, 23. Reticulati-sporites cusus Loose, 1934, p. Tuberculati - sporites microtuberosus Inst. Palaobot. Arb., vol. 4, no. 3, p. 156, pi. (Loose) Ibrahim, Loose, 1934, Inst. Palao- 7, fig. 25. bot. Arb., vol. 4, no. 3, p. 147. FIRMUS (LoOSc) S. W. 7. PUNCTATI-SPORITES 18. PuNCTATi-spoRiTES MicROVERRUCOsus (Ibra- comb. nov. and B., him), S. W. and B., comb. nov. Verrucosi-sporites firmus Loose, 1934, Verrucosi - sporites microverrucosus idem, p. 154, pi. 7, fig. 30. Ibrahim, 1933, Sporenformen des Aegirhori- zonts, p. 25, pi. 7, fig. 60. 8. PUNCTATI-SPORITES GLOBOSUS (LoOSe) S. W. and B., comb. nov. 19. PuNCTATi-SPORiTES NOBiLis (Wicher) S. W. and B., comb. nov. A piculati-spontes globosus Loose, 1934, Sporites nobilis Wicher, 1934, Inst. idem, p. 152, pi. 7, fig. 14. Palaobot. Arb., vol. 4, no. 4, p. 186, pi. 8, fig. 30. 9. PUNCTATI-SPORITES GRANDIVERRUCOSUS Midland Naturalist, Kosanke, 1943, Am. 20. PuNCTATi-SPORiTES PAPILLOSUS (Ibrahim) pi. fig. 4. vol. 29, no. 1, pp. 127, 128, 3, S. W. and B., comb. nov. Verrucosi-sporites papillosus Ibrahim, (Ibrahim) S. 10. PuNCTATi-SPORiTES GRANiFER 1933, Sporenformen des Aegirhorizonts, pi. B., nov. W. and comb. 5, fig. 44. Granulati - sporites granifer Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 21. PUNCTATI-SPORITES PARVIPUNCTATUS Kos- anke, 1943, Naturalist, vol. 22, pi. 8, fig. 72. Am. Midland 29, no. 1, p. 127, pi. 3, figs. 3, 3a, 3b. ATI- SPORITES GRUMOSUS (Ibrahim) S. 11. PuNCT 22. PuNCTATi-sPORiTES PUNCTATUS (Ibrahim) and B., comb. nov. W. Ibrahim, 1933, Sporenformen des Aegirhori- Verrucosi-sporites grumosus Ibrahim, zonts, p. 21, pi. 2, fig. 18. 1933, idem, p. 25, pi. 8, fig. 68. Sporonites punctatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 57, Abt. B, p. 12. PuNCTATi-spoRiTES iNsiGNiTis (Ibrahim) 448, pi. 15, fig. 18. S. W. and B., comb. nov. Punctati-sporites punctatus (Ibrahim) Apiculati - sporites insignitis Ibrahim, Ibrahim, Loose, 1934, Inst. Palaobot. Arb., 1933, idem, p. 24, pi. 6, fig. 54. vol. 4, no. 4, p. 146, pi. 7, fig. 27.
23. PUNCTATI-SPORITES RETICULOCINGULUM 13. PuNCTATi-sPORiTEs( ?) IRREGULARIS (Ber- (?) (Loose) S. W. and B., comb. nov. ry) S. W. and B., comb. nov. Sporonites reticulocingulum Loose, 1932, Zonales-sporites irregularis Berry, 1937, Neues Jahrb., Beilage-Band 57, Abt. B, p. Am. Midland Naturalist, vol. 18, no. 1, p. 450, pi. 18, fig. 41. 156, fig. 9. Reticulati - sporites reticulocingulum (Loose) Ibrahim, 1933, Sporenformen des 14. PuNCTATi-SPORiTES LACUNOSUS (Ibrahim) Aegirhorizonts, 34. S. W. and B., comb. nov. p. Reticulati - sporites reticulocingulum Reticulati - sporites lacunosus Ibrahim, (Loose) Ibrahim, Loose, 1934, Inst. Palao- 1933. Sporenformen des Aegirhorizonts, p. bot. Arb., vol. 4, no. 4, p. 156. 36, pi. 6, fig. 50. 24. PuNCTATI - SPORITES SPHAEROTRIANGULATUS 15. PUNCTATI-SPORITES LATIGRANIFER (LoOSC) (Loose) S. W. and B., comb. nov. S. W. and B., comb. nov. Sporonites sphaerotriangulatus Loose, Sporonites latigranifer Loose, 1932, 1932, Neues Jahrb., Beilage-Band 67, Abt. Neues Jahrb., Beilage-Band 67, Abt. B, p. B., p. 451, pi. 18, fig. 45. 452, pi. 19, fig. 54. Laevigati - sporites sphaerotriangulatus Granulati-sporites latigranifer (Loose) (Loose) Ibrahim, 1933, Sporenformen des Loose, 1934, Inst. Palaobot. Arb., vol. 4, Aegirhorizonts, p. 20. no. 3, p. 147. Laevigati - sporites sphaerotriangulatus (Loose) Ibrahim, Loose, 1934, Inst. Palao- 16. PuNCTATi-spoRiTES MACULATUS (Ibrahim) bot. Arb., vol. 4, no. 4, p. 145. S. W. and B., comb. nov. 25. PuNCTATi-sPORiTES SPiNosus (Loose) S. Reticulati-sporites maculatus Ibrahim, W. and B., comb. nov. 1933, Sporenformen des Aegirhorizonts, p. A piculati-sporites spinosus Loose, 1934, 2,6, pi. 6, fig. 56. idem, p. 153, pi. 7, fig. 20.
17. PUNCTATI- SPORITES M ICROTUBERO SU S 26. PUNCTATI-SPORITES SPINULISTRATUS (LoOSC) (Loose) S. W. and B., comb. nov. S. W. and B., comb. nov. Sporonites microtuberosus Loose, 1932, Sporonites spinulistratus Loose, 1932, Neues, Jahrb., Beilage-Band 67, Abt. B, Neues Jahrb., Beilage-Band 67, Abt. B, p. p. 450, pi. 18, fig. 33. 451, pi. 18, fig. 47. ) .
32 PALEOZOIC FOSSIL SPORES
Apiculata-sporitesspinulistratus (Loose) Spore coat. — Of relatively uniform Sporenformen des Aegir- Ibrahim, 1933, thickness throughout, generally thin, and horizonts, 37. p. except where modified by ornamentation, A piculati-sporitcsspmulistratus (Loose) often less than 2 microns. Loose, 1934, Inst. Palaobot, Arb., vol. 4, no. 4, p. 153. Affinity.—Spores similar to those of Granulati-sporites have been reported 27. PUNCTATI - SPORITES TRIGONORETICULATUS (Loose) S. W. and B., comb. nov. from fructifications of ferns although rel- Reticiilati - sporitcs trigonoreticulatus atively few of these have been critically Loose, 1934, idem, p. 155, pi. 7, fig. 9. described. Spores of Bozveria minor and Renaultia gracilis which are illustrated by 28. PUNCTATI-SPORITES (?) VELATUS (LoOSe) 5. W. and B., comb. nov. Knox (1938) are of similar character. Reticulati-sporites velatus Loose, 1934, Probably considerable difflculty will be idem, p. 155, pi. 7, fig. 19. encountered in definitely correlating the genus (further emended as may prove 29. PuNCTATi-sPORiTES VERRUCOSUS (Ibrahim) desirable) with any single supra-generic S. W. and B., comb. nov. Sporonites verrucosus Ibrahim, 1932, plant group. The many similar features Neues Jahrb., Beilage-Band 67, Abt. B, p. possessed by species assigned to Granulati- 448, pi. 15, fig. 17. sporites suggest that this grouping may be Verrucosi-sporites verrucosus ( Ibrahim a practical one. Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 25, pi. 2, fig. 17. The forms Raistrick (1933, 1934, 1937) designates as D^, Dg and D-^^ (Knox, Genus Granulati-sporites (Ibrahim, 1938) probably belong to Granulati-spor- 1933) emend., S. W. and B. ites as defined here. Likewise Millott's Type 4 is of this character (Millot, 1939)
Plate 1, figures 8-8b Remarks: The genus as emended in- Symmetry.—Spores radial, trilete. cludes fifteen species, twelve of which represent new name combinations. The Shape.—Originally oblate, rounded tri- genus was originally established to include angular, ^N\.ih. the axial dimension much spores of trilete-type showing granular the shortest ; when compressed the spores are nearly always flattened in good proxi- sculpturing (Ibrahim, 1933, p. 21), char- acters which in themselves are inadequate mo-distal orientation since the transverse ; plane of the spore, being broadest, tends consequently it must be regarded as co- to parallel the bedding of sediments en- incidence that two of the three original closing them. When preserved thus, the species (one of which was designated as spores appear subtriangular in outHne, the type) are retained in the group, along with sides either convex or slightly con- with another that Loose assigned to it in cave and the corners rounded. 1934. The genotype of Granulati-sporites is the only genotype species included by Sise.—Spores of various species range revision within the genus and thus there from about 25 to 45 microns in mean can be no question as to validity of the diameter. generic name. The genotype, G. granulatus Ornamentation. — Various, surfaces Ibrahim, also is based on a form from the smooth to punctate, finely to relatively Aegir coal bed at the top of the West- coarsely reticulate or apiculate ; the di- phalian B. verse types of ornamentation which char- acterize various species are not prominent 1. Granulati-sporites deltiformis S. W. and to the extent that they mask the essential B., nomen nov. shape characteristics of the spores. Triquitrites deltoides Wilson and Coe, 1940, Am. Midland Naturalist, vol. Haptotypic structures.—Trilete rays 23, no. 1, p. 185, fig. 9. are well extended toward the corners; Note.—In combination with Granulati-sporites they are never relatively short. lips The the specific epithet becomes a homonym of G. are not distinguished by special ornamen- deltoides (lb.) S. W. and B., as given below. tation, the commissure is relatively simple 2. Granulati-sporites deltoides (Ibrahim) but definite in all cases ; usually no vari- S. W. and B., comb. nov. ations in ornamentation distinguish the Sporonites deltoides Ibrahirh, 1932, proximal (pyramic) side from the rest Neues Jahrb., Beilage-Band 67, Abt. B, p. of the spore coat. 448, pi. 15, fig. 15. ;
ALATI-SPORITES 33
Laevigati-sporites deltoides (Ibrahim) 13. Granulati-sporites trigonus (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- S. W. and B., comb. nov. horizonts, p. 20, pi. 2, fig, 2. Reticulati-sporites trigonus Ibrahim, Note.—The species is larger than most forms 1933, Sporenformen des Aegirhorizonts, p. assigned to Granulati-sporites. ?>7, pi. 5, fig. 34.
14. Granulati-sporites triquetris (Ibrahim) 3. Granulati-sporites fistulosus (Ibrahim) B., nov. S. W. and B,, comb. nov. S. W. and comb. errucosi-sporites triquetris Ibrahim, Reticulati-sporites fistulosus Ibrahim, V 1933, idem, p. 26, fig. 61. 1933, idem, p. 36, pi. 5, fig. 35.
15. Granulati-sporites verrucosus (Wilson 4. Granulati-sporites gibbosus (Ibrahim) and Coe) S. W. and B., comb. nov. S. W. and B., comb. nov. Triquitrites verrucosus Wilson and Coe, errucosi-sporites gibbosus Ibrahim, V 1940, Am. Midland NaturaHst, vol. 23, no. 1933, idem, p. 25, pi. 6, fig. 49. 1, p. 185, fig. 10.
5. Granulati-sporites granulatus Ibrahim, 1933, idem., p. 22, pi. 6, fig. 51. Genus Alati-sporites Ibrahim, 1933 Note.—Type species of Granulati-sporites. Plate 1, figures 6-6b 6. Granulati-sporites microgranifer Ibra- Symmetry. Spores radial, trilete. him, 1933, idem, p. 22, pi. 5, fig. 32. — Shape.—Body, exclusive of bladders, is 7. Granulati-sporites microsaetosus sub-triangular. (Loose) S. W. and B,, comb. nov. Sporonites microsaetosus Loose, 1932, Sise.—In the vicinity of 80 microns, Neues Jahrb., Beilage-Band 67, Abt. B, p. overall diameter. 450, pi. 18, fig. 40. Ornamentation of spore.—Body moder- Setosi-sporites microsaetosus (Loose) ate ; may be punctate, finely reticulate Ibrahim, 1933, Sporenformen des Aegir- bladders smooth to punctate, etc. Blad- horizonts, p. 26. Setosi-sporites microsaetosus (Loose) ders, in general, less definitely ornamented Ibrahim, Loose, 1934, Inst. Palaobot. Arb., than spore body. vol. 4, no. 3, p. 148. Haptotypic structures.—Trilete rays
8. Granulati-sporites microspinosus (Ibra- relatively long, little ornamented. him) S. W. and B., comb nov. Spore coat.—Consisting of two distinct Apiculati-sporites microspinosus Ibra- membranes; the outer (perisporal) mem- him, 1933, Sporenformen des Aegirhori- brane is very thin and expanded in the in- zonts, p. 24, pi. 6, fig. 52. ter-radial areas to form bladders in a trim- 9. Granulati-sporites parvus (Ibrahim) S. erous (in threes or multiples of three) W. and B., comb. nov. pattern. The spore body v^all (exospore) Sporonites parvus Ibrahim, 1932, Neues is usually more than twice as thick as the Jahrb., Beilage-Band 67, Abt. B, p. 448, bladder membrane, darker, easily pi. 15, fig. 21. and distinguished that account. Punctati - sporites parvus (Ibrahim) on Ibrahim, 1933, Sporenformen des Aegir- Affinity.—Spores of Alati-sporites seem horizonts, pi. p. 21, 2, fig. 21. smaller but essentially similar in con- Reticulati-sporites parvus (Ibrahim) struction to those of Spencerites (Scott, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 1898, Kubart, 1910). The three inter- 4, p. 154, pi. 7, fig. 13. radial bladders are most distinctive in 10. Granulati - sporites piroformis Loose, both genera. The significance of this 1934, idem, vol. 4, no. 3, p. 147, pi. 7, fig. 19. character is hardly well enough under- 11. Granulati-sporites (?) priddyi (Berry) stood to properly evaluate evidence as 5. W. and B., comb. nov. to their relationship.^ Zonales-sporites priddyi Berry, 1937, ^ The haptotypic relations, if any, of the three blad- Am. Midland Naturalist, vol. 18, no. 1, p. ders present on pollen of Podocarpus dacrydioides and on 156, fig. 2. Pherosphaera are not known, and comparison with Car- boniferous forms does not seem particularly pertinent for 12. Granulati-sporites torquifer (Loose) S. obvious reasons. W. and B., comb. nov. It seems doubtful that the form described by Daugherty (1941, p. 64) as Spencerites? chinleana is actually refer- Sporonites torquifer Loose, 1932, Neues able to Spencerites or could be referred to Alati-sporites since it apparently does not possess three Jahrb., Beilage-Band 67, Abt. B, p. 450, pi. interradial bladders. Daugherty does not mention and the figure 18, fig. 43. given does not show the trilete sutures. If these were .Reticulati-sporites torquifer (Loose) present the species might be referred to Cirratriradites provided the equatorial appendage was a flange. Lacking Loose, Inst. Arb., no. 1934, Palaobot. vol. 4, these characteristics, his alternative suggestion of a - 3/.p. 154. gymnospermic affinity seems most plausible. 34 PALEOZOIC FOSSIL SPORES
Spores assigned by Raistrick to type Sise.—Spores of various species range D5 without much question belong to from about 40 to 100 microns in diameter. Alati-spontes. Ornam^entation.—Coarsely and often ir- Remarks.—Only one species has been regularly reticulate ; aside from the coarse described to date but there is little doubt ornamentation the body wall may have a the group will receive far more recogni- variously smooth, punctate or finely re- tion. It is highly distinctive and complex ticulate texture. In the ''alete" forms enough that specific characters are easily reticulation may show a tendency toward
distinguishable. No doubt the character a spiral pattern ; in others it may be mostly of the bladders will lend itself to the limited to the distal part of the spore. possibility of these purpose. There is no Haptotypie structures. — The trilete forms being confused with unseparated structures may be weakly developed ; no other genera tetrad groups belonging to arcuate ridges are developed but the or- have been although illustrations which namentation may sometimes be absent on lead one to think so. published might the proximal surface. The trilete rays In addition to its recognition in the vary considerably in development and Ruhr district and in Britain the genus length among different species. has been recognized in coals of the Illinois Spore coat.—Appears to consist often- basin by Brokaw and in Tennessee coals times of two membranes ; when present, by Bentall. These from America appear the outer one (perisporal?) is thinner and to be specifically distinct and as yet un- more or less intimately connected with the described forms. coarse reticulation; the inner (exospor- diagnosis given above has The generic al?) membrane is thicker and sometimes in the light of this been constructed irregular and excentric in its development. while con- additional information, and The wall may appear thick, and yet be Ibrahim's mea- siderably augmented over quite translucent. ger description (1933, p. 32), it may be Affinities. The only spores of this gen- improper to consider it as a generic emend- — eral or possible character known in fruc- ation. The type, of course, is the species tifications allied Sphenophyllum. given below, which was first described are with spores are inadequately from the Aegir coal in the Ruhr. Spkenophyllum known in the detail necessary for close - (Ibrahim) 1. Alati SPORITES PUSTULATUS comparison, however, and considerably Ibrahim, 1933, Sporenformen des Aegir- more evidence is needed to form an opin- horizonts, p. 33, pi. 1, fig. 12. Sporonites pustulatus Ibrahim, 1932, ion as to the relationship of Reticulati- Neues Jahrb., Beilage-Band 67, Abt. B, p. sporites with other groups. fig. 12. 448, pi. 14, Types C2, C3, F4, and Fg of Raistrick Alati - sporites pustulatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb.. (1934, 1935, 1937) and possibly also F„ vol. 4, no. 3, p. 151, pi. 7, fig. 4. F2 (cf. Knox, 1938), Fg and F4, appear Type Ds Raistrick, 1937, Congres pour to belong to Reticulati-sporites. Types I'avancement des etudes de stratigraphie F5 and Fg and D,3 (Knox, 1938) are carbonifere, Heerlen, 1935, Compte rendu, more problematic. Type 8 of Millott vol. 2, p. 911. (1939) also belongs here and is most closely related to R. facetus of species now Genus Reticulati-sporites ( Ibrahim, described. Knox' Type possibly 1933) emend., S. W. and B. 5K may be similar to some species included in Re- Plate figures 7-7b 1, ticulati-sporites. Spores illustrated by Symmetry.—Spores, in essence, radi- Knox (1939) from Fifeshire, figs. 49 and ally symmetrical ; some species are weakly 50, unquestionably belong to Reticulati- trilete and others show no indication of sporites. Unpublished forms of similar their tetrahedral tetrad origin but are con- nature have been found in Ohio coals by sidered trilete in derivation because of Wilson and Brokaw, and they have also their shape. been recognized in Iowa and Illinois and Shape.—Originally spherical or mod- Tennessee. eratel}^ oblate ; compressed snores are cir- Remarks.—Seven species are included cular disk-like, generally without prom- here under Reticulati-sporites, only one inent folds. of which represents a new name combina- RETICULATI-SPORITES 35 tion. This form, R. facetus (Ibrahim), sporites apparently lack the further spe- was originally designated as the genotype cialization of germinal pores, neverthe- of Reticulata-spontes with five species less lack of good development of a trilete assigned to it. Essentially the only char- commissure in some of the forms might acter shared by these five was their lack possibly signify that the plants repre- of a visible haptotypic marking. Most sented here had passed beyond the typi- diverse types of reticulation characterized cal cryptogamic stage of evolution. Re- their surfaces. R. facetus, however, shows ticulati-sporites appears to be a relatively a type of reticulation which is very simi- homogeneous group of natural relation- lar to that of R. reticulatus (genotype of ship. The coarse reticulation is highly Reticulati-sporites) and others in this characteristic and for the most part quite genus. Other characters are also in sub- unmistakable. Nevertheless these forms stantial agreement and it is believed that are rather anomalous in their morphology the lack of a trilete marking in this case and offer difficulties in interpretation. is insufficient basis for generic separation. Those suggested are regarded as tenta- Other species of Reticulati-sporites show tive only. a very weak trilete marking. Consequent- The genus is best known from the Aegir ly Reticulata-sporites Ibrahim (1933, p. coal bed at the top of the Westphalian B, 38) may be dispensed with and regarded which is approximately equivalent in age as a later synonym for Reticulati-sporites to middle Des Moines and upper Potts- Ibrahim (1933, p. 33). ville-lower Allegheny beds. Knox has suggested that alete (1939) 1. Reticulati-sporites corporeus Loose, 1934, spores with somewhat spiral reticulation Inst. Palaobot. Arb., vol. 4, no. 3, p. 155, such as shown by R. facetus (cf. fig. 49) pl. 7, fig. 7. resemble spores of Riccia but this com- 2. Reticulati-sporites facetus (Ibrahim) parison does not seem particularly close. S. W. and B., comb. nov. Neither does the comparison of the R. Reticulata-sporites facetus Ibrahim, corporeous type with Fossombronia spores 1933, Sporenformen des Aegirhorizonts, p. 38, pl. 5, fig. 36. seem appropriate because the fossil forms evidently have a perisporal membrane en- 3. Reticulati-sporites mediareticulatus Ib- closing them instead of mere "large alve- rahim, 1933, idem, p. 34, pl. 7, fig. 62. olae with high lamellae." ISTevertheless it 4. Reticulati-sporites ornatus (Ibrahim) must be admitted that in general spores of Ibrahim, 1933, idem, p. 35, pl. 1, fig. 7. Reticulati-sporites bear more resemblance Sporonites ornatus Ibrahim, 1932, Neues Jahrb., to some of the spores of the Hepaticae il- Beilage-Band 67, Abt. B, p. 447, pl. 14, fig. 7. lustrated by Knox than any of the other Paleozoic groups. The apparent absence 5. Reticulati-sporites polygonalis (Ibra- of haptotypic markings on several of the him) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 155, pl. 7, fig. 16. modern forms is a striking feature which Sporonites polygonalis Ibrahim, 1932, should be more thoroughly investigated, Neues Jahrb., Beilage-Band 67, Abt. B, p. although it need not occasion great sur- 447, pl. 14, fig. 8. prise to find so advanced a character in Laevigati-sporites polygonalis (Ibra- a group whose discrete phylogeny goes him) Ibrahim, 1933, Sporenformen des Aegirhorizonts, back at least to the Carboniferous. p. 19, pl. 1, fig. 8.
It may be suggested that spores of 6. Reticulati-sporites reticulatus (Ibra- him) Ibrahim, Reticulati-sporites underwent more mod- 1933, idem, pp. 33-34, pl. 1, fig. 3. ification within the sporangium subsequent Sporonites reticulatus Ibrahim, 1932, to the breakup of the original tetrad than Neues Jahrb., Beilage-Band 67, Abt. B, is usual for cryptogamic spores. The p. 447, pl. 14, fig. 3. highly developed reticulation pattern prob- Reticulati-sporites reticulatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., ably was laid down relatively late in sporo- vol. 4, no. 3, p. 156. genesis by a tapetal plasmodium, some- Note.—Type species of Reticulati-sporites. times without respect for the proximo- distal relationships established in original 7. Reticulati-sporites (?) reticuliformis Ibrahim, 1933, Sporenformen des Aegirhori- tetrad groupings. Such a sequence occurs zonts, p. 34, pl. 7, fig. 62). in formation of many kinds of angi- Note.—Trilete rays are unusually long and ospermic pollen. Spores of Reticulati- well developed in this species. 36 PALEOZOIC FOSSIL SPORES
Nomina excludende are sometimes distinguishable flaring lat-
1. Reticulati-sporites angulatus Ibrahim, 1933, Spo- erally from both ends of the suture line. renformen des Aegirhorizonts, etc., p. 35, pi. 7, fig. 59. Spore coat.—Varying somewhat in 2. Reticulata-sporites bireticulatus (Ibrahim) Loose, thickness relative to the other dimen- 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 158, pi. 7. fig. 28. sions; the spore coat is oftentimes thin Sporonites bireticulatus Ibrahim, 1932, Neues Jahrb.. Beilage-Band 61, Abt. B. p. 447, pi. 14, and translucent. The internal cavity is fig. 1. Reticulati-sporites bireticulatus (Ibrahim) Ibra- sometimes more *'bean-shaped" than the him, 1933, Sporenformen des Aegirhorizonts, etc., external outline due to internal thickening p. 35, pi. 1. fig. 1. 3. Reticulati-sporites facierugosus Loose, 1934, Inst. of the spore coat in the central proximal Palaobot. Arb., vol. 4, no. 3, p. 155, pi. 7, fig. 26. region. 4. Reticulata-sporites mediapudens Loose, 1934, idem, p. 158, pi. 7. fig. 8. Affinity.—Smaller species of Laeviga- 5. RbTICULATI - SPORITES MICRORETICULATUS (Loose) Loose, 1934. idem, p. 155. to-sporites agree with some forms ob- Sporonites microreticulatus Loose, 1932, Neues tained filicinean fructifications. Jahrb.. Beilage-Band 67, Abt. B, p. 450, fig. 37. from type 6. Reticulati-sporites morosus Loose, 1934, Inst. There is now no particularly good evi- Palaobot. Arb., vol. 4, no. 3, p. 154, pi. 7, fig. 2. 7. Reticulati-sporites nexus (Loose), Ibrahim, 1933, dence as to the affinity of the larger forms. Sporenformen des Aegirhorizonts, etc., p. 34. Fredda Reed (1938, p. 333) has obtained Sporonites nexus Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, fig. 35. spores of this type from an extraordinary Reticulati-sporites nexus (Loose) Ibrahim, Loose, 1934. Inst. Palaobot. Arb., vol. 4, no. 3, p. 156, new type of Calamarian fructification pi. 18, fig. 35. which are approximately 50 microns in 8. Reticulati-sporites sifati Ibrahim, 1933, Sporen- formen des Aegirhorizonts, etc., p. 35, pi. 8, fig. 67, length. Spores of Zeilleria are of this 9. Reticulata-sporites spongiosus Ibrahim, 1933. type (Florin, 1937, p. 316-7) but it is not idem, p. 39, pi. 8, fig. 71. 10. Reticulati-sporites vinculatus (Ibrahim). Loose, now possible to decide whether that genus 1934, Inst. Palaobot. Arb.. vol. 4, no. 3, p. 156. belongs to the ferns Sporonites vinculatus Ibraham, 1932, Neues or pteridosperms Jahrb., Beilage-Band 67, Abt. B, p. 448. pi. 15. (Halle, 1933, p. 88). The "distal fur- fig- 19. Reticulata-sporites vinculatus (Ibrahim) Ibrahim, row" of Zeilleria spores remarked by 1933, Sporenformen des Aegirhorizonts, etc., p. 39, Florin p. 317) appears no pi. 2, fig. 19. (1937, more definite than many of the fortuitous folds Genus Laevigato-sporites (Ibrahim, seen in spores of Laevigato-sporites. 1933) emend., S. W. and B. It thus appears that species of Laevi- gato-sporites could conceivably pertain to Plate 1, figures 5-5b at least three distinct contemporaneous Symmetry.—Spores bilateral, mono- orders of Paleozoic plants. It seems lete. doubtful that our information on these re- Shape. — Originally broadly bean- lationships will be greatly improved in the immediate future and until it is shaped ; elongate oval in the plane of lon- pos- gitudinal symmetry, round or oval in the sible to revise the genus on the basis of transverse plane. When compressed the reliable evidence it will serve a useful spores tend to be folded variously depend- geological, if not a botanical, purpose. In ing on the size and morphology of the a good many instances species of Laevi- various specific types of spores. gato-sporites form a majority of the spores obtained from Size.—Spores of various species range certain coal beds. They are, however, rare from about 20 to over 130 microns in or absent in a their long dimension. number of lower Pottsville coals that have been examined. Ornamentation.—Smooth to finely punc- Spores tate, apiculate or rugose ; rarely showing evidently belonging to Laevi- pronounced reticulation or strong apicu- gato-sporites have been assigned to types lae. B,a, B,b, and Bg by Raistrick (1933, 1937) (Knox, Haptotypic structures.—Consisting of 1938). Millott's types 6 and E^a a simple monolete linear suture, generally (Millott 1939) and Knox' (1942) Type without lips specially distinguished, and 6K may also belong here. usually continued for more than half the Remarks.—Nine species, one of which total length of the spore. The suture may includes three designated formae, are now be very inconspicuous if it coincides in listed for Laevigato-sporites. Ibrahim position with the edge of a compressed originally assigned two species to the spore, or with the axis of a longitudinal genus. The second of these, L. ellipsoides, fold. Ends of very delicate arcuate ridges is now referred to Monoletes. It is pos- LAEVIGATO-SPORITES 37
sible that some of the names given below 6. Laevigato-sporites (?) pennovalis Berry, will prove to be synonyms. On the other 1937, Am. Midland NaturaHst, vol. 18, no. 1, pp. 156-7, fig. 4. hand a number of distinct and as yet un- recorded species have been recognized in 7. Laevigato-sporites thiessenii Kosanke, the Illinois basin and elsewhere. 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 125, pi. 3, figs. 1, la, lb. The list includes genotype species of Pittsburgh microspore of Thiessen, various four previously named genera. L. bila- publications. See especially Thiessen and Stand, teralis (Loose) was the only species de- 1923. scribed by Loose (1934, p. 159) under 8. Laevigato-sporites (?) tuberculatus the new name, Reticulato-sporites. The (Berry), S. W. and B., comb. nov. name was credited to 'Tbrahim 1932" al- Tuberculati-sporites tuberculatus Berry, though no "reticulata" was ever used by 1937, idem, vol. 18, no. 1, p. 155, fig. 9. that author. L. is desmoinensis the gen- 9. Laevigato-sporites vulgaris (Ibrahim) otype of Phaseolites Wilson and Coe Ibrahim, 1933, Sporenformen des Aegirhor- izonts, 39-40, pi. 2, fig. 16 pi. 5, figs. 2>7, (1940). L. minutus (Ibrahim) was the pp. ; sole species given under Punctato-sporites 38, 39. Sporonites vulgaris Ibrahim, 1932, by Ibrahim (1933, p. 40). L. vulgaris Neues Jahrb. Beilage-Band 67, Abt. B, p. Ibrahim is the designated genotype of 448, pi. 15, fig. 16. Laevigato-sporites Ibrahim (1933, p. 39). Laevigato-sporites vulgaris forma minor All of these species are evidently con- Loose, 1934, Inst. Palaobot. Arb., vol. 4, generic, their chief distinction being in no. 3, p. 158, pi. 7, fig. 12. variation of surface ornament and small Laevigato-sporites vulgaris forma maior Loose, idem, pi. 7, fig. 6. differences of size, none of which can be 1934, p. 158, Laevigato-sporites vulgaris forma max- regarded as of more than specific impor- iMus Loose, 1934, idem, p. 158, pi. 7, fig. 11. tance. The names Reticulato-sporites, Phaseolites, and Punctato-sporites are NOMEN EXCLUDENDE therefore regarded as synonyms of Laevi- 1. Laevigato-sporites rheaensis Berry, 1937, Am. Mid- gato-sporites. All these are typically rep- land Naturalist, vol. 18, no. 1, p. 157, fig. 13. Note. —The morphology of this form is so obscure that resented by specimens from the upper its significance cannot be ascertained. part of the Westphalian B and approxi- mately equivalent beds in America. Genus Zonalo-sporites Ibrahim, 1933 1. Laevigato-sporites bilateralis (Loose), S. W. and B., comb. nov. Symmetry.—Spores apparently bilat- Reticulato-sporites bilateralis Loose, eral, monolete. 1934, Inst. Palaobot. Arb., vol. 4, no. p. 3, Shape.—Flattened elliptical ( ?), broad- 159, pi. 7, fig. 22. ly elliptical in transverse plane. Note.—Reticulato-sporites is a generic name published only by Loose (1934) although he 6^i>^.—Relatively large, 200-300 mi- erroneously credited it to "Ibrahim, 1932." crons in length. Ornamentation. — Surface smooth to 2. Laevigato-sporites cranmorensis Berry, 1937, Am. Midland Naturalist, vol. 18, no. minutely rugose. 1, p. 157, fig. 1. Haptotypic features.—Linear proximal suture with slight angular median deflec- 3. Laevigato-sporites desmoinensis (Wilson and Coe) S. W. and B., comb. nov. tion, termini of arcuate ridges may be Phaseolites desmoinensis Wilson and weakly differentiated. Coe, idem, 1940, vol. 23, no. 1, p. 183, fig. 4. Spore coat.—Perisporal bladder mem- Note.—Evidently closely related to Laevigato- brane thin, enveloping spore body on all sporites vulgaris. sides, and in contact with it proximally and distally. Body wall (exospore) rela- 4. Laevigato-sporites minimus (Wilson and Coe) S. W. and B., comb. nov. tively thick and brownish translucent. Phaseolites minimus Wilson and Affinity. — Probably pteridospermic, Coe, 1940, idem, vol. 23, no. 1, p. 183, fig. 5. judging from points of similarity with Monoletes which is discussed later. It 5. Laevigato-sporites minutus (Ibrahim) S. W. and B. comb. nov. nevertheless is still extremely problematic since it superficially seems to combine Punctato-sporites minutus Ibrahim, features 1933, Sporenformen des Aegirhorizonts, p. of spores of Medullosan and of 40, pi. 5, fig. Z2>. Cordaitean relationship. 38 PALEOZOIC FOSSIL SPORES
angu- Rcuiar'ks.—This genus is reported in and diagnostic feature is the slight only two instances, in the Ruhr and in lar deflection of the suture fine near the IlHnois, in both cases from relatively few middle. Although inconspicuous and specimens. The forms have been assigned sometimes hardly evident it seems to be to different species as listed below. Much universally present. It may constitute the more information is needed before their sole remnant of a third vestigial ray in a significance can be understood. fundamentally trilete suture pattern. Very weakly developed termini of arcuate rid- 1. ZONALO-SPORITES AUREOLUS (Schopf) S. W. ges are occasionally resolved at the two and B., comb. nov. ends of the functional suture line. The Monoletes aureobis Schopf, 1938, Illi- taper- nois Geol. Survey Rept. Inv. 50, pp. 45-46, suture opens to form a sharp pi. 1, figs. 1-2. pointed lenticular slit. The distal grooves have no connection with haptotypic struc- 2. ZoNALO-sPORiTES viTTATUs Ibrahim, 1933, tures. Sporenformen des Aegirhorizonts, p. 41, pi. 6, fig. 45. Spore coat.—The exospore consists of
a layer of varying thickness ; the proxi- Genus Monoletes (Ibrahim) 1933, mal side and the center of the distal emend, S. W. and B. umbo are as much as 15 or 18 microns; Plate 2, figures 17-17c the thinnest is at the base of the distal grooves on either side of the umbo where Symmetry.—Spores apparently bilat- it may be less than 5 microns. An even eral, monolete. Mature forms evidently gradation exists between these extreme have been modified considerably in post areas and in normally compressed spores tetrad maturation, and it is possible that the exospore thickness at the margin is although the mature forms appear very intermediate between the extremes. The definitely bilateral, they may have arisen endosporal membrane is frequently evi- from tetrahedral tetrads. dent as a crumpled translucent sack less Shape. Elliptical — to rounded lenticu- than a micron in thickness, sometimes lar in outline as viewed from the proxi- shrunken from the exospore at the spore mal side; of similar figure but of more margin. The umbo and distal grooves, slender proportions in the plane of the when present, have the appearance of long axis viewed from the side ; the short functioning as a harmomegathus. transverse axis shows a somewhat rounder Affinity. Spores of this character dis- and shorter but notably flattened outline. — tinguish a pteridospermic plant alliance The distal surface oftentimes is marked probably largely coextensive with the Me- by two prominent grooves extending near- duUosaceae. They are characteristic of ly the length of the spore with a well all genera placed in the Whittleseyinean rounded umbo between. Sometimes the sub-tribe (see Halle, 1933). The only proximal side near the suture shows a aberrant feature is the lack of distal very slight prominence. On compression, grooves in spores of Codonotheca; spores longitudinal folds frequently parallel the of all distal grooves. other genera now assigned to this alliance possess them. Codonotheca spores Si^e. Spores relatively — large, varying are otherwise entirely characteristic of from slightly over 100 microns to as Monoletes and when such spores are much as half a millimeter in length. found isolated they probably should be Ornamentation. — Surfaces generally assigned to Monoletes since there well minutely granulose appearing quite may be other types in addition to Codon- smooth at low magnification; sometimes otheca that also may lack distal grooves glistening in reflected light. Emphytic and umbo. Spores of Codonotheca empha- marking is evenly distributed except at the size the non-essential character of these extreme base of distal grooves, which are specialized distal structures. more smooth. Remarks.—Much of the previous dis- Haptotypic features.—^When closed the cussion is based on comparative study of proximal suture is a very narrow linear spores in Dolerotheca and Codonotheca groove, generally lacking marginal dis- by Schopf from petrified and compression tinction, though the lip area may be very material as well as from spores isolated slightly upraised. The most noteworthy by maceration of coal. It has thus been DENSO-SPORITES 39 possible to establish accurately from sec- 2. Monoletes ovatus Schopf, 1935, Illinois tions the proximal position of the suture Acad. Sci. Trans, vol. 28, no. 2, p. 108, fig. 7. Pollen of Dolerophyllum sp., Zerndt, and the distal location of the harmome- 1930, Acad, polonaise sci. Bull, internal., gathic grooves. The fact that spores are ser. B, pp. 55-56, pi. 8, figs. 42-49. frequently found with the suture opened Type 31 Zerndt, 1931, idem, ser. A, p. but, except in instances of obvious me- 176. chanical or chemical injury, have never Monoletes ovatus Schopf, 1935, Illinois Acad. Sci. Trans, vol. 28, no. p. 176. been observed with the umbo split off as 2, Monoletes ovatus Schopf, 1938, Illinois an "operculum" or the grooves other than Geol. Survey Rept. Inv. No. 50, pp. 43-45, intact, strongly suggests that proximal pi. 1, figs. 3-5, pi. 6, figs. 1-4. gametophytic exit was still a fundamental Pollen of Dolerotheca fertilis, Zerndt, feature of these forms as it must have 1940, Paleontographica, vol. 84, Abt. B, p. been in their more ancient cryptogamic an- 136-138, pi. 11, fig. 36. cestry. Such spores are also regarded as Denso-sporites (Berry, 1937), prepollen (cf. Schopf, 1938, pp. 14-15). Genus emend., S. W. and B. The foregoing diagnosis serves to cor- rect some erroneous interpretations of Plate 1, figures 9-9c structure previously held; e.g. the "en- Symmetry.—Spores radial, trilete. circling ridge" described by Schopf (1938, Shape. Originally oblate, round to p. 45) for the species designated as Mono- — subtriangular in equatorial outline. When letes ovatiis, is the margin of the distal the central area becomes grooves discussed here. It has no hapto- compressed, much thinner than the margins due to va- typic significance although it possibly is wall thickness. Coal thin not emphytic in origin in the same sense riation in spore sections, taken vertically through the coal that ordinary surface ornamentation is. bed, show them compressed into a "dumb- Monoletes was first used by Ibrahim in bell" shape. The original highly oval sec- 1933 to designate a new group in his ar- tional form results in extremely uniform tificial system. Although substantive in proximo-distal orientation in enclosing form and similar to a generic name, it sediments. was not originally applied in that sense Sise. Diameter of specimens assigned and no nomenclatural type has been pre- — among various species varies from about viously proposed. It has been used as a 35 to 100 microns. ,eeneric designation and applied more nar- Ornamentation. — Smooth to apiculate rowly (Schopf, 1936, 1938) ; we now emend it to correspond with the restricted and rugose. External surface character, usage. Monoletes ovatus Schopf may typically is uniform, though ornamenta- serve as the type species of the genus. tion may be more strongly developed to- The group now includes two species, ward the equator in some forms. both of which appear to be widely dis- Haptotypic structures.—A delicate tri- tributed. It is likely that further species lete marking is visible in well preserved will be distinguished by application of material or where the central area of the more refined biometric procedure. Both proximal surface is consistently present. species must still be regarded as rather "Fissures" penetrating to the margin in generalized types. some instances may be essentially hapto- typic continuations of the trilete sutures. 1. Monoletes ellipsoides (Ibrahim) Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, Spore coat.—Characterized by great p. 45, pi. 1, fig. 14; pi. 6, figs. 5 and 6. differences in thickness. The proximal Sporonites ellipsoides Ibrahim, 1932, and distal walls are usually membranous, Neues Jahrb. Beilage-Band 67, Abt. B., p. or at least significantly thinner than the 449, pi. 17, fig. 29. equatorial portion of the coat. The latter Laevigato-sporites ellipsoides (Ibra- area is oftentimes so thick it is practically him) Ibrahim, 1933, Sporenformen des opaque to transmitted light in contrast to Aegirhorizonts, p. 40, pi. 4, fig. '29. the highly translucent central area. It Punctato-sporites ellipsoides (Ibrahim) may also be so extended centrifugally as Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, pp. 158-9, pi. 7, fig. 35. to simulate a flange but it is not usually Sporites ellipsoides (Ibrahim) Wicher, strictly demarcated from the spore body, 1934, idem, vol. 4, no. 4, p. 185. as such. 40 PALEOZOIC FOSSIL SPORES
mation about these spores may reflect Affinities.—Spores of this genus have indirectly, upon the not been recognized in connection with importantly, although coal formation. fructifications. This is surprising in view problem of splint of their frequence in coal. They have 1. Denso-sporites annulatus (Loose) S. "biconcave," been roughly designated as W. and B., comb. nov. "salvershaped" and as "splint micro- Sporonites annulatus Loose, 1932, Neues spores" by Thiessen and his co-workers Jahrb. Beilage-Band 67, Abt. B, p. 451, pi. (Thiessen and Wilson, p. 9, 1924; Thies- 18, fig. 44. - (Loose) sen, 1930; Sprunk, et al, 1940) and can be Zonales sporites annulatus Loose, 1934, Inst. Palaobot. Arb., vol. 4, recognized by their characteristic "dumb- no. 3, p. 151.10 bell" shape in cross section.^ Modern methods of study may be ex- 2. Denso-sporites covensis Berry, 1937, Am. Midland Naturalist, vol. 18, p. 157, fig. 11. pected therefore to disclose definite evi- Note. Although serving as genotype, the spe- dence of the relationship of this group. — cific characteristics of this form are inadequately Their structure is so unusual that it seems known. evident that a single homogeneous group 3. Denso-sporites densus Berry, 1937, Am. of plants for the most part is represented. Midland Naturalist, vol. 18, no. 1, pp. 157-8, Raistrick's Type A spores (Raistrick fig. 7. and Simpson, 1933) apparently are nearly - sporites indignabundus entirely referable to Denso-sporites. His 4. Denso (?) (Loose) S. W. and B., comb. nov. A„ Ae, (1934) and A,o (1937, 1938) Sporonites indignabundus Loose, 1932, also are probably congeneric and, if so, Neues Jahrb. Beilage-Band 67, Abt. B, p. appear to be the most bizarre of any in 451, pi. 19, fig. 51. the genus. Possibly the A3 spores (Rai- Zonalcs-sporites indignabundus (Loose) strick, 1937; Knox, 1938) are least as- Ibrahim, 1933, Sporenformen des Aegir- suredly related to this group. Knox's Type horizonts, p. 32. indigna bund us Cg (Knox, 1942) possibly might be placed Apiculati-sporites (Loose) Loose, 1934, Inst. Palaobot. Arb., here though its morphologic interpreta- vol. 4, no. 3, p. 153. tion is somewhat uncertain due to its un- usual type of equatorial development. 5. Denso-sporites loricatus (Loose) S. W. and B., comb. nov. Remarks. Six species are now included — Sporonites loricatus Loose, 1932, Neues under Denso-sporites and it seems that a Jahrb. Beilage-Band 67, Abt. B, p. 450, pi. good many more will need be distin- 18, fig. 42. guished. Raistrick's A5, Ag, and A^q Zonales-sporites loricatus (Loose) types are worthy of specific distinction. Loose, 1934, Inst. Palaobot. Arb., vol. 4, The generic name, while perhaps of un- no. 3, p. 151. desirable construction, is not in conflict 6. Denso-sporites (?) quadratus Berry, with any others that have been proposed 1937, Am. Midland Naturalist, vol. 18, p. and thus is entirely valid. 158, fig. 10. problematic form. As mentioned previously, the spores Note.—A very are widespread and very abundant in splint coals. It has been suggested that Genus Cystosporites Schopf, 1938 a special type of vegetation was respon- Plate 1, figures 10-lOb sible for this type of coal but such can hardly be the case, since recognizable splint Symmetry.—Spores radial, trilete. coals are also found that are so much Form and size.—Variable depending on younger that floral similarities cease (cf. development. Fertile members relatively Thiessen and Sprunk, 1936). White (in enormous, sack-like, elongate, sometimes discussion of Thiessen, 1930, p. 672) attaining a length of more than a centi- doubts that the presence of a certain type meter and about half as broad. Abortive of plants is responsible for forming this forms variable in development, charac- type of coal. The coincidence of occur- teristically ranging from as small as one- rence between Denso-sporites and Pale- 10 Since Loose recognized the rather significant distinc- ozoic splint coal nevertheless seems tion between Denso-sporites and the zonate forms, in a unaccountably high and additional footnote appended to his article he suggested "einer Unter- infor- abteilung Annulati-sporites" for D. annulatus and D, loricatus. Because he did not adopt the name, "Unter- 8 Thiessen has also designated some spores not referable abteilung Annulati-sporites" is invalidly published, at to Denso-sporites as "splint spores." least in the sense of a generic designation. CYSTOSPORITES 41 half a millimeter in diameter to more than nal adaptation, appear to be generalized twice that. Smallest forms are more so far as specific differences are con- nearly isodiametric becoming axially elon- cerned. The ornamental patterns which gate with increased size. When compressed serve to distinguish spores of free-spor- both abortive and fertile members are ing lycopsids have evidently undergone folded irregularly ; their longest axis is extensive reduction if such characteristics generally least effected. were ever present in the line of lepido- Ornamentation, — Generally absent; carp ancestry. The genus Cystosporites abortive forms may show apparent re- thus expresses a broader relationship ticulation. than that exemplified by other genera in the Lepidocarpaceae, and, so far as is Haptotypic features.—Strongly devel- known, is inclusively correlative with the oped in fertile forms but relatively in- conspicuous because of the enormous ex- lepidocarp family. Future discoveries may it possible to restrict this pansion of the distal part of the spore make group and establish relationships in greater detail. body. Sutures definite, extended to the arcuate ridges which are strongly thick- The spores found within the seeds of lepidocarp species are receiving more de- ened, lips often moderately defined; py- tailed ramic areas distinct from adjacent spore study now (Reed, 1941 ; Darrah,
1941 ; Hoskins and Cross, ; Schopf, coat. Apex may be somewhat elongate. 1941 On smaller abortive forms haptotypic 1941) and further studies may show features are commonly masked by develop- other features of the spore coat that have unsuspected systematic significance. ment of a thick rugose, more or less triangular cushion which effectively seals Zerndt (1930, et seq.) has classified the spore apex. Arcuate ridges are in- some forms now placed in Cystosporites, conspicuous but sometimes their termini under Triletes. The distinction between are weakly defined adjoining the corners free-sporing and seed-bearing lycopsids of the cushion. Larger abortive forms seems at least familial in its importance commonly show haptotypic features more (Schopf, 1941) and this distinction is like fertile spores but somewhat less devel- most in evidence when the megaspores oped and more irregular in character. are compared. Spore coat. — Variable in thickness, Remarks.—The fertile and abortive generally relatively thick on abortive and megaspores of species of Cystosporites are considerably thinner, especially in the very heteromorphous. In several instances median areas, on fertile spores. Pyramic they have been recovered in tetrad group- areas of fertile forms are thinner but the ings from maceration residues of coal. more proximal portion immediately distal They nevertheless are generally separated to the arcuate ridges is thick ; the median from one another so that a system of portion of the spore is membranous and treatment for the isolated tetrad members the extreme distal portion may again be both fertile and abortive seems essential somewhat thicker. The most characteristic for scientific reporting. The characters of feature of this genus is the fibrous char- the fertile spores are used to identify acter of the spore coat in the membra- species, and fertile specimens serve as nous middle region ; thicker parts of the holotypes for the two species listed below. coat also consist of thicker matting of the Isolated abortive spores may be distin- interlocking, variable width, anastomos- guished in classification as formae. The ing fibrils. A thinner nonfibrous endo- procedure may serve as a practical means sporal membrane may be present. The of expressing relationships between these fibrous spore coat is evidently equivalent heteromorphous spore types and still re- to the exospore. tain for each of them a nominal distinc- Affinites.—Cystosporites is a member tion. There appear to be no inherent dis- of the Lepidocarpaceae. It is intimately advantages in such a policy, as it affords related to the genera Lepidocarpon, Illi- a means of more accurate recording of niocarpon, and probably others of this actual material encountered ; only the ac- family in which the correlation has not crual of additional information can show been as definitely established (Schopf, whether such a practice, in fact, is neces- 1941). Spores of Cystosporites, although sary. If the formae later appear super- highly specialized with reference to semi- fluous in some instances and worthy of .
42 PALEOZOIC FOSSIL SPORES actual specific status in others, these Shape.—Body nearly spherical, blad- changes can be effected without undue ders of moderate inflation placed laterally taxonomic difficulty or confusion since and opposite one another on the spore type material is indicated in all instances. give an oval external outline. On com- Cystosporites has an extensive range pression few folds are formed and these profile unless in the Carboniferous as is best shown by generally do not modify the tables of spore distribution given by the plane of the bladders fail to coincide Zerndt (1937), and so far as is now with the plane of compression. known is restricted to this period. The Size.—Relatively large; species attain type species was described from Illinois as much as 300 microns length from one No. 6 (Herrin) coal. bladder tip to the other. Ornamentation. — Body wall may be 1. Cystosporites breretonensis Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, pp. rugose, bladder membrane lightly sculp- 40-42, pi. fig. pi. figs. 1-4. 3, 5, 8, tured.
Triletes cf . T. giganteus Zerndt, Schopf, Relatively incon- 1936, Illinois Acad. Sci. Trans, vol. 28, Haptotypic features.— no. 2, p. 107, fig. 5 (holotype of C. spicuous and evidently modified in post- breretonensis) tetrad development. Trilete rays often Triletes cf. T. giganteus Zerndt, Schopf. developed with two long rays and one 1936, idem. p. 175. characteristically shorter. One pyramic Cystosporites breretonensis forma abor- area may sometimes be distinguished; Tivus Schopf, 1938, Illinois Geol. Survey in this area no arcuate marking is Rept. Inv. 50, p. 40, pi. 1, fig. 10, pi. 8, fig. 4. except evident. Cystosporites breretonensis forma retic- ULATUS Schopf, 1938, idem, p. 40, pi. 1, fig. Spore coat.—Body wall (exospore) 11. tending to be dense and of moderate rela-
2. Cystosporites giganteus (Zerndt) Schopf, tive thickness ; bladder membrane (peri- 1938, idem, p. 39. spore), relatively thin and quite trans- Triletes giganteus Zerndt, Acad, 1930, lucent. polonaise sci. Bull, internal., ser. B, p. 71, pis. 9, 10, and 11. Affinity.—There is considerable prob- Type 1 Zerndt, 1931, Acad, polonaise ability that the plants represented by Para- sci. Bull, internal., ser. A, 170. p. sporites are gymnospermous. The peri- Sporites giganteus (Zerndt) Wicher, sporal bladder development is strongest 1934, Inst. Palaobot. Arb., vol. 4, no. 4, still p. 172, pi. 8, fig. 9. evidence of this. It is a question Triletes giganteus Zerndt, 1937, Acad, whether the genus should be assigned with polonaise sci. Trav. Geol. no. 3, p. 4. the Pteridosperms, Cordaitaleans or the Cystosporites giganteus forma varius Conifers (cf. Schopf, 1938, pp. 47-8). (Wicher) S. W. and B., comb. nov. The genus has some evident claim to Triletes (Lagenicula) glabratus Zerndt. 1930, Acad, polonaise sci. Bull, internal., affinity with Florinites, S. W. and B., ser. B., p. 54, pi. 8, figs. 38-41. Pityosporites Seward, Alisporites Daugh- (?) Type 29 Zerndt, 1931, idem, ser. erty, and to Endosporites Wilson and Coe. A., p. 175. Perhaps it is closest to Endosporites, but Type 30 Zerndt, 1931, idem, ser. A., p. as these forms are contemporaneous their 175, pi. 8, figs. 26-27. relationship cannot easily be regarded as Type 30 Zerndt, 1932, Jahrb. fur das Berg-u. Huttenwesen in Sachsen. Tahrg. direct. 1932, p. 13A, pi. 1, figs. 1, 2, 3. Remarks. — The genus is monotypic Sporites varius Wicher, 1934, Inst. being represented only by the species Palaobot. Arb., vol. 4, no. 4, p. 89, pi. 6, figs. 2, 3, 4, 6. given below which has been found in upper Carbondale and lower McLeans- boro age coals of Illinois. point upon Genus Parasporites Schopf, 1938 The which greatest emphasis should be placed Plate figures 2, 16-16b is that the development of proximal Symmetry.—Prepollen grains appear- sutures is such as to indicate proximal ing bilateral due to opposing bladders. The gametophytic exit similar to Endosporites fundamental symmetry, clearly shown by and Monoletes. Aside from this and its the body, is radial. rather large size Parasporites has pollen CIRRATRIRADITES 43
grain characteristics. It may represent a what less than that opposite the rays. The veritable "prepollen". margin of the flange is often minutely to rather coarsely serrate. 1. Parasporites maccabei Schopf, 1938, Illi- nois Geol. Survey Rept. Inv. 50, pp. 48-9, Affinity.—Spores of Cirratriradites are pi. 1, fig. 6, pi. 7, figs. 1-3. not yet definitely correlated with any major plant group. Their most likely affinity Genus Cirratriradites Wilson and Coe, seems to be with the lycopods but known 1940 types of Lepidostrobus microspores are Plate 3, figures 21, 21a, 21b usually smaller and generally possess no Symmetry.—Spores trilete, radial. comparable development of the flange. Shape.—Moderately oblate spheroidal, Certain zonate types of Triletes spores with a strongly projecting equatorial bear a superficial resemblance to Cirratri- flange. The flange may assume a tri- radites but so far as is known now there angular horizontal outline due to emphasis is no inherent correlation between struc- of the trilete rays, or it may be nearly ture of megaspores and microspores where circular; the spore body is circular or heterospory is as highly developed as it is slightly triangular. When compressed, in Triletes. There is in fact no essential few folds are evident and, because of the evidence to show whether spores of Cirra- flange and oblate shape of the body, the triradites functioned as microspores or plane of flattening nearly always cor- isospores. The Carboniferous existence of responds with that of the flange. isosporous lycopods is still most firmly supported only on theoretical grounds but Size.—Spores of various species range possibly represented from about 40 to over 100 microns in they could be by overall diameter. Cirratriradites. Some authors have confused Cirratri- Ornamentation.—Commonly showing a radites spores with those of genera {En- pattern having its primary emphasis along dosporites, Spencerites, etc.) which pos- radial lines ; may consist of ridges more sess perisporal bladder development in or less anastomosing to the point of be- the equatorial plane. Such resemblance coming a reticulation, or surfaces may be as there is, is certainly only superficial. nearly free of ornamental ridges and most of the surface smooth, granulose or Raistrick's A^, A^, and Cg (cf. Rai- finely punctate. Some species show a strick 1935, 1937, 1938) belong here; likewise, possibly in part. unique type of distal ornamentation con- D2 As given sisting of one or several thinner areas with by Knox, (1938, 1942) D,^ is definitely referable to Cirratriradites. Types 1 and rather prominent margins. The flange is of Millott without often more or less radially striated and in 9 (1939) much ques- addition may develop one or two concen- tion belong in this genus. The spores illus- trated in tric bands of irregular thickening. The by Reinsch (1884, p. 22) plate pyramic areas are not particularly dis- 15, figs, la and lb as types 222 and 223 tinguished by ornamentation. from Zwickau, Saxony, are characteristic of it. Haptotypic features.—Trilete rays rela- tively strongly developed and extended to Remarks. — Thirteen previously de- scribed species are in list the equator, oftentimes a line of thick- included the be- ening continues to the edge of the flange. low, all but one of which, C. maculatus Lips are frequently strongly demarcated the genotype, represent new name com- binations. and raised above the spore body ; the suture lines are attenuate but distinct. No Spores of this character are frequently arcuate ridges are developed distinct from encountered in American coals and addi- the flange and, in fact, the flange may be tional species will subsequently be de- taken to represent a hyper-development scribed. The group shows such general of these haptotypic formations. The flange agreement in numerous characteristics is usually relatively broad, sometimes com- that a considerable degree of natural prising more than half the total spore relationship is attributed to it. diameter. It is definitely distinguished 1. Cirratriradites argutus (Ibrahim) S. W. from the spore body, being thinner than and B., comb, nov. Zonalcs-sporites argutus Ibrahim, 1933, the body wall and more translucent ; the Sporenformen des Aegirhorizonts, pp. 31-32, interradial flange width is often some- pi. 6, fig. 55. , ;
44 PALEOZOIC FOSSIL SPORES
Ib- 2. CiRRATRiRADiTES (?) cicATRicosus (Ibra- Zonales-sporites saturni (Ibrahim) him) S. W. and B., comb. nov. rahim, Loose, 1934, Inst. Palaobot. Arb., fig. 23. Sporonites cicatricosiis Ibrahim, 1932, vol. 4, no. 3, p. 149, pi. 7, Neues Jahrb., Beilage-Band 67, Abt. B, p. A^of^.—Ibrahim (1933) came to the conclusion 447, pi. 14, fig. 2. that Sporonites formosus Ibrahim (1932) was infor- Zonales-sporites cicatricosus (Ibrahim) conspecific with C. saturni. Very little is to support this and, since Ibrahim, 1933, Sporenformen des Aegirhor- mation provided the figures appear to show specific differences, it izonts, p. 31, pi. 1, fig. 2. appears desirable to keep them separate for a 3. CiRRATRiRADiTES FAUNus (Ibrahim) S. W. time at least. and B., comb. nov. 11. CIRRATRIRADITES TENUIS (LoOSc) S. W. and Sporonites fannus Ibrahim, 1932, Neues B., comb. nov. Jahrb., Beilage-Band 67, Abt. B, p. 447, Sporonites tenuis Loose, 1932, Neues pi. 14, fig. 4. Jahrb., Beilage-Band 67, Abt. B., p. 450, Zonales-sporites fannus (Ibrahim) Ib- pi. 18, fig. 34. rahim, 1933, Sporenformen des Aegirhori- Zonales-sporites tenuis (Loose) Loose, zonts, p. 28, pi, 1, fig. 4. 1934, Inst. Palaobot. Arb., vol. 4, no 3, 4. CiRRATRiADiTES FORMOsus (Ibrahim) S. W. p. 149. and B., comb. nov. 12. CiRRATRiRADiTES VENUSTUS (LoOSC) S. W. Sporonites formosus Ibrahim, 1932, and B., comb. nov. Neues Jahrb., Beilage-Band 67, Abt. B, Sporonites venustus Loose, 1932 Neues p. 447, pi. 14, fig. 10. Jahrb., Beilage-Band 67, Abt. B., p. 450, Zonales-sporites saturni (Ibrahim) pi. 18, fig. 36. Ibrahim, in part, 1933, Sporenformen des Zonales-sporites venustus (Loose) Aegirhorizonts, p. 30, pi. 1, fig. 10. Loose, 1934, Inst. Palaobot. Arb., vol. 4, 5. CiRRATRiRADiTES (?) GRACILIS (Zcrndt) no. 3, p. 149. S. W. and B., comb. nov. 13. CiRRATRiRADiTES ZONALIS (LoOSe) S. W. Triletes gracilis Zerndt, 1937, Acad. and B., comb. nov. polonaise sci. Bull, internat., ser. A., p. 586, pi. 12, figs. 1-10. Zonales-sporites sonalis Loose, 1934, idem, vol. no. p. 148, pi. 7, fig. 5. Type 46 Zerndt, 1937 (idem). 4, 3,
Note—Species exceptionally large (300 ti) otherwise in close generic agreement. Genus Endosporites Wilson and Coe, 1940 6. CIRRATRIRADITES MACULATUS WilsOU and Coe, 1940, Am. Midland Naturalist, vol. Plate 2, figures 14-14b 23, no. 1, p. 183, fig. 7. Symmetry. Spores trilete, radial. Note.—Reexamination of the type material — suggests the trilete apex is ordinarily normally Shape.—Moderately flattened elliptical developed and that the apical opening originally in transverse plane, in axial plane round described is not of general occurrence. Sculp- to oval or elliptical or slightly triangular turing simulating it may occur distally. body wall more spherical than the bladder. 7. CiRRATRiRADiTES ( ?) PEACOCKi (Berry) S. When compressed the spores most com- W. and B., comb. nov. monly shov^ good proximo-distal orienta- Zonales-sporites peacocki Berry, 1937, tion. The spore body is not generally idem, vol. 18, p. 156, fig. 5. folded but the bladder membrane more 8. CiRRATRiRADiTES PENNINGTONENSIS (Ber- commonly shows irregular plication. The ry) S. W. and B., comb. nov. bladder profile at the margin of the spore Zonales-sporites penning tonensis Berry, corresponds to the profile of folds, being 1937, idem, p. 156, fig. 3. straight or evenly rounded, and thus 9. CIRRATRIRADITES RARus (Ibrahim) S. W. easily distinguishable from simple flanges and B., comb. nov. of zonate forms that superficially re- Zonales-sporites rarus Ibrahim, 1933, semble Endosporites, Sporenformen des Aegirhorizonts, p. 29, pi. 6, fig. 53. Size.—Ranging in various species from about 50 to possibly 300 microns total 10. CiRRATRiRADiTES SATURNI (Ibrahim) S. W. and B., Comb. nov. diameter. The spore body generally is less than one-half of the full diameter. Sporonites saturni Ibrahim, 1932, Neues Jahrb., Beilage-Band 67, Abt. B., p. Ornamentation. — Surfaces of bladder 448, pi. 15, fig. 14. are levigate to granular or punctate ; a Zonales-sporites saturni (Ibrahim) Ib- fine meshed reticulation pattern is com- rahim, in part, 1933, Sporenformen des present but this is probably the Aegirhorizonts, p. 30, pi. 2, fig. 14 (non pi. monly on 1, fig. 10). inside, rather than on the external bladder ;
ENDOSPORITES 45
surface. The spore body is less definitely Parasporites, Cordaianthus, Alisporites, ornamented although the bladder pattern Caytonanthus, Pityosporites and Endo- may appear superimposed on it. Emphytic sporites, as well as in modern podocarps ornament seems to be the same for proxi- and Abietineae. There is good reason to mal and distal surfaces. believe that bladders of this type are ho- structures, which have un- Haptotypic features.—Trilete rays com- mologous may monly extend to the periphery of the dergone progressive and regressive de- spore body and thickened continuations velopment at various times within the from them may carry on to the bladder broad confines of this alliance, but their membrane. The lips are oftentimes up- nevertheless largely preserve iden- raised and definite, the suture line distinct. tity throughout. Sometimes a thickening is present on the Endosporites shows such evident de- bladder membrane which corresponds to velopment of proximal haptotypic struc- arcuate ridges. tures that gametophytic exit was quite evidently from the proximal pole as in Spore coat.—The bladder (perisporal) the cryptogams. Florin, working with membrane is quite thin and translucent excellently preserved silicified material, its reticulation tends to add somewhat to has shown that in some late Pennsylvanian the thickness as seen in marginal profile. (Stephanian) forms the haptotypic fea- The body wall is substantially thicker al- tures were vestigial and in mature forms though also quite translucent, and cor- present only as a surface imprint on the responds to the exosporal layer. In a bladder membrane which does not even few instances thin membranous, almost maintain contact with the spore body. hyaline, endosporal membranes have been The bladder and body are in contact dis- observed within the exospore and shrunk- tally, however, and prothallial cells line en from it. The suture lines are not dis- the body cavity except at this point. Thus tinct on these, but three clear-cut endo- in these advanced forms germinal exit sporal apical papillae occur at the juncture must have been distal as it is in pollen of of pyramic apices. — all modern conifers. Pollen-like types of Affinities. Endosporites is related to this sort having indication of proximal some of the Pennsylvanian Cordaitaleans. exit have been termed *'prepollen," fol- They correspond to spores observed by lowing Renault, and its significance has Wilson in male strobili, and Schopf has been discussed elsewhere (Schopf, 1938; found well preserved specimens in such pp. 14 and 15, 48). Endosporites male abundant association with Cordaitean spores are evidently prepollen in this leaves, other plant fossils being infrequent, sense. that no other conclusion seems permis- Eight previously described species are sible. It should be emphasized, however, listed below, only one of which is queried. that the pollen grains Florin (1936) The genus is widely distributed both in described in Cordaianthus fructifications America and in Europe. The genotype obtained from the French Stephanian are species, E. ornatus Wilson and Coe, is evidently generically distinct and of more from Iowa coal of Des Moines age. advanced structure.
1. Endosporites angulatus Wilson and Coe, Raistrick's types C-^ and C4 without 1940, Am. Midland Naturalist, vol. 23, no. much question belong to Endosporites 1, p. 184, fig. 1. (Raistrick and Simpson, 1933; Knox, 2. Endosporites globiformis (Ibrahim) S. 1938). Type 629 of Reinsch (1884, p. 61, W. and B., comb. nov. pi. 48, fig. 252A) from the Blatterkohle Sporonites globiformis Ibrahim, 1932, and Stigmarienkohle of Metschowk (in Neues Jahrb., Beilage-Band 67, Abt. B, p. central Russia) and probably others less 447, pi. 14, fig. 5. easy to interpret from his drawings, also Zonales-sporites globiformis (Ibrahim) belongs to Endosporites. Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 28, pi. 1, fig. 5. Remarks.—The distended bladder mem- Zonales-sporites globiformis (Ibrahim) brane is the most characteristic feature Ibrahim, Loose, 1934, Inst. Palaobot. Arb., of certain gymnospermous pollen grains. vol. 4, no. 3, p. 148. The expansion of the bladder is various 3. Endosporites (?) karczewskii (Zerndt) in the several groups which are known as S. W. and B., comb. nov. ;
46 PALEOZOIC FOSSIL SPORES
Trilctcs karcacwskii Zerndt, 1934, Ornamentation. — Surfaces levigate to Acad, polonaise sci. Trav. Geol. no. 1, p. granulose and mildly verrucose, angular 27, pi. 31, fig. 3. areas sometimes more highly ornamented. Agrees well with Endosporites though Note. — Emphytic ornamentation of proximal and larger than most species. distal surfaces is about the same. 4. Endosporites ornatus Wilson and Coe, rays ex- 1940, Am. Midland Naturalist, vol. 23, no. Haptotypic structures.—Trilete
1, p. 184, fig. 2. tended nearly to the margin of the body
Spore 6, Wilson and Brokaw, 1937, cavity; lips sometimes rather thick and Iowa Acad. Sci., vol. 44, pp. 129-130, fig. 6. prominent, but usually not particularly
5. Endosporites pellucidus Wilson and Coe, demarcated. Arcuate ridges lacking ; like- 1940, Am. Midland Naturalist, vol. 23, no. 1, wise, no flange in the usual sense is de- p. 184, fig. 3. veloped though extreme extensions of the Spore 1, Wilson and Brokaw, 1937, spore coat at the angles may simulate a Iowa Acad. Sci., vol. 44, pp. 128-129, fig. 1. partial flange. 6. Endosporites rotundus (Ibrahim) S. W. Spore coat. Characterized by gross in- and B., comb. nov. — Zonales-sporites rottmdus Ibrahim, equalities in exospore thickness ; spore 1933, Sporenformen des Aegirhorizonts, p. coat is thickest on the angles opposite ends 31, pi. 8, fig. 73. of the rays and thinnest in the central
7. Endosporites rugatus (Ibrahim) S. W. distal and possibly to a somewhat lesser and B., comb. nov. extent on the interradial areas. Actual Zonales-sporites rugatus Ibrahim, 1933, thickness of difl^erent parts of the wall idem, p. 31, pi. 8, fig. 70. in various species varies greatly although 8. Endosporites volans (Loose) S. W. and the relations of thicker and thinner areas B., comb. nov. of the spore coat is rather constant. Sporonites volans Loose, 1932, Neues — Affinity. - Triquitrites cannot be as- Jahrb., Beilage-Band 67, Abt. B, p. 451, pi. 18, fig. 46. signed to any major plant group at the Reticulati-sporites volans (Loose) Ib- present time although the character of the rahim, 1933, Sporenformen des Aegirhori- spore coat seems so individualistic that zonts, 36. p. the genus is considered to correspond with Zonales-sporites volans (Loose) Loose, natural plant relationship. The features, in 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149. general, seem to agree best with spores of Note.—Trilete rays are shorter than is com- filicineans. mon in this genus. Type 10 of Millott (1939) belongs to Triquitrites and probably type D^o of Rai- Genus Triquitrites Wilson and Coe, strick (1938). Type D, (Raistrick, 1934, 1940 1935, 1938) shows more extreme thicken- ing Plate 3, figures 20-20b of the radial angles of the spore coat so that it is evidently closely related to Symmetry. Spores trilete radial. — ; this genus, though it is a question whether Form.—Oval to elliptical in vertical it should be classified as Triquitrites or plane, distal side sometimes slightly more distinguished from it. Knox' Type 4K inflated than the proximal ; in the equato- (1942) is clearly congeneric. Spores of rial plane triangular in outline, corners the same sort but even more extreme have rounded or truncate subangular and some- been illustrated by Reinsch (1884) in his times extended, sides slightly convex to plate 3, figures 34-42, corresponding to strongly concave in profile. Folds are his type dia.s^noses, nos. 345, 347, 349, 353, seldom induced by compression due to 354, 362, 363, 365, 373. Reinsch also good preferential orientation coinciding illustrated other forms which are doubt- with the horizontal plane of the spores less congeneric with Triquitrites in his forms more inflate distally are indicated tvpes 58, 200, 341, 342, 348, 355. etc. through a tendency of the corners to be Type 58 shown on his plate 15, figure directed slightly upwards exaggerating the 18A, from Zwickau, Saxony, is remi- natural thickening of the spore coat on the niscent of some, possibly overmacerated, angles. specimens of the genotype species. Sise.—Spores of various species com- Remarks.—Five named species are as- monly range from 35 to 70 microns in signed to Triquitrites ; three represent new mean diameter. name combinations. Other species are EQUISETOSPORITES 47 known from American coals and will sub- Sporonites triturgidus Loose, 1932, Beilage-Band Abt. B, sequently be described. The numerous Neues Jahrb., 67, p. 449, pi. 18, fig. 32. spores illustrated by Reinsch (1884) in the Valvisi-sporites triturgidus {Loose) Micro-Paleophytologia show best the vari- Loose, 1934, Inst. Palaobot. Arb., vol. 4, ation in structure encountered is this no. 3, p. 151. group. The spores with highly exag- gerated angular thickenings were mostly Daugherty, derived from localities in Russia where Genus Equisetosporites the coals probably are of Lower Carbonif- 1941 erous age. Dio and D^ types of Raistrick This genus recently described from Tri- (1938) were chiefly obtained from the assic age beds in southwestern United Lower Carboniferous of England. Penn- States is mentioned here because of its sylvanian age species are much more important bearing on recognition of fossil moderate in spore coat thickness and the spores of certain Equisetalean plants. The interradial and distal membranes are fre- genus is monotypic, including only E. quently less than 3 microns thick, as in chinleana as given below, whose note- the four species which have been de- worthy feature is the presence of elaters scribed and named. The angles are more very similar to those characterizing the than twice as thick, however, and their modern genus. The vahdity of the name relationship thus seems definitely indi- Equisetosporites rests on the belief that cated. Nevertheless, it may be desirable these forms should be distinguished from that the thicker walled more ornate and spores of Equisetum. It appears there is radially extended forms be generically adequate support for this view since segregated when detailed modern infor- Daugherty, in a personal communication, mation becomes available for the older states "in all cases Equisetum-like. plants species. The tendency toward seeming of the Triassic have proven to be quite dif- simplification in spore coat structure of ferent when the record is complete enough late Carboniferous plants as contrasted to allow specific determination." Thus with ealier merpbers of the same alliance until further evidence is available, it has also been noted in members of the would seem unwise to emphasize the mod- Lagenicula and Aphanozonateae sections ern aspect of this fossil form. Its dis- of Triletes. covery nevertheless adds an important
1. Triquitrites arculatus Wilson and Coe, item of information to the record. 1940, Am. Midland Naturalist, vol. 23, no. Knox examined spores of fifteen species 1, p. 185, fig. 8. of Equisetum and reports (1938, 439- Note.—Type material bears a moderate to pp. sparse verrucose flecking (areas of thickening) 40) them to be invariably spherical, 30 on proximal and distal surfaces, possibly due to microns to 35 microns in diameter, and overmaceration. always thin walled. In all these species
2. Triquitrites spinosus Kosanke, 1943, Am. the spore wall was faintly granular. Ela- Midland Naturalist, vol. 29, no. 1, p. 128. ters of somewhat varied character are pi. figs. 2a, 2b. 3, 2, present on all modern species of Equi-
3. Triquitrites tribullatus (Ibrahim) S. setum. In commenting on Raistrick's Bg W. and B., comb. nov. type, which she recognizes is very prob- Sporonifes tribullatus Ibrahim, 1932, ably of calamarian affinity, Knox remarks Neues Jahrb., Beilage-Band Abt. B, 67, (pp. 461-3) on p. 448, pi. 15, fig. 13. with thin wall and Laevigati-sporites tribullatus (Ibrahim) the same globose form
of ornamentation . . . the unusual de- Ibrahim, 1933, Sporenformen des Aegirhor- absence velopment of elaters and no trace of the tri- izonts, pp. 20-21, pi. 2, fig. 13. radiate mark. Valvisi-sporiies tribullatus (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, She cites Halle's description of Rhaetic no. 3, p. 152, pi. 7, fig. 21. and Triassic - age Equisetuni spores in which the trilete com- 4. Triquitrites trigonappendix (Loose) S. (Halle, 1908) W. and B., comb. nov. missure is clearly seen but which show Valvisi-sporites trigonappendix Loose, no trace of elaters. Halle assumed that 1934, idem, p. 152, pi. 7, fig. 17. even if elaters had been present they would have been removed by maceration 5. Triquitrites triturgidus (Loose) S. W. and B., comb. nov. procedure. Needless to say, this aspect 48 PALEOZOIC FOSSIL SPORES
must always be considered. Nevertheless, 1. Equisetosporites chinleana Daugherty, Carnegie Inst. Washington Pub. 526, the spores recovered by Halle were typi- 1941, p. 63, pi. 34, fig. 4. Age, Triassic. cal of calamarians (cf. Hartung, 1933) and isolated spores of this character must Alisporites Daugherty, 1941 be classed with Calamospora n. gen. as de- Genus scribed below. Eqiiisetosporites chinleana Plate 2, figure 12 is quite different and marks the advent of a more distinctly modern character in This genus, like the preceding, is mono- this ancient and persistent order of plants. typic, the sole species yet described being Evidently a reduction in importance of of Triassic age from Southwestern United haptotypic features has occurred and con- States. It is included in this paper be- currently the perisporal adaptation repre- cause spores of Permain age which are sented by elaters has come into existence. probably referable to it have been found, Careful study of the fossil spores may be and because there seems some basis for re- expected to indicate more completely the lating it to Parasporites and Pityosporites. relationship and distribution of equiseta- The only description is that of the geno- lean types. type species, A. opii, given below (Dau- gherty 1941, p. 98). The type of the single species of Equi-, setosporites now known has been made "The spores are large, averaging 100 to 110 microns in length and having two large mem- available to us for examination through branaceous wings with reticulate markings. the kindness of Prof. Ralph W. Chaney They are spherical to ovate in dorsal view,^i and the following description, which is having a rather thick exine and a single fusi- somewhat more detailed than Daugher- form furrow." ty's, is offered for comparison with the Pollen grains of Caytoniales, when iso- other descriptions' given in this paper. lated, without much doubt should be clas- This specimen (No. 1562 of the Califor- sified under Alisporites unless there is nia Museum of Paleontology) is ade- particularly good evidence for referring quately illustrated by Daugherty in the them to Caytonanthus. Harris (1941) figure cited below. has distinguished Caytonanthus kochi, C. oncoides, and C. arberi largely on the ba- Symmetry.—No trace of original tet- sis of differences in their pollen and has rad configuration visible. suggested their correlation with species Shape.—Body spherical (type speci- of Caytonia and Sagenopteris found in men elliptical, 30 x 37}^ microns, due to the same deposits. All evidence seems one elongate taper-point fold). to point to the validity of Harris' con- Ornamentation.—Body wall essentially clusions, yet further information will smooth showing very slight surface un- probably tend to modify the picture. For dulation. example, certain of the spores isolated from the Stassfurt salt deposits by Liick Spore coat. Thin, apparently slightly — (1913) seem properly referable to Alis- less than a micron in thickness. In spite porites and similar to spores of Caytonan- of this the body is deep brown color as thus, but probably few would suggest that viewed by transmitted light. the genus Caytonanthus should be iden- Elaters.—Probably four in number, tified on that slender basis. The fructifi- having their attachment close together; cations which bore the Permian Alispo- each one possibly about 70 microns long, rites would be presumed to differ generi- band-like, about 4 microns wide, with cally due to the general tendency of such slight irregularities; yellowish translu- organs to be evolutionarily plastic and cent. Elater ends truncated, tapering for amenable to structural modification. The a short distance, not at all broadly ter- generic features of pollen grains seem in minated. (Two of the four free ends general to be more conservative. But are shown at the right edge and over- whether organs are evolutionarily con- lapping the spore body in Daugherty's servative or amenable to biocharacter al- fig. 4.) Evidently the elaters tended to " The terms dorsal and ventral tend to be ambiguous in spirally encircle the spore body but they description of pollen grains, particularly unless some fur- ther explanation is given. Probably "dorsal" as used have been displaced on the specimen at here refers to the proximal side, i.e., the side which was hand. internal and adjoined the other members of the original tetrad. ;
CALAMOSPORA 49 teration, the important point relates to 2. Alisporites spp. ? recognition of the degree of change that Pollen Forms I, II, and III, Liick, Beitrag zur Kenntnis des alteren is manifested and of the fact that bio- 1913, Salzgebirges im Berlepsch—Bergwerk bei character modification has proceeded at Stassfurt nebst Bemerkungen uber die pol- different rates in the different organs and lenfuhrung des Salztones, pp. 29-31, figs. parts of the plant life cycle. In order 53-56, 58-59. that fossil identifications may be scien- Genus Calamospora S. W. and B., gen. tifically trustworthy these principles nov. should be more widely acknowledged.
Alisporites is a designation that should Plate 3, figures 22-22b ; text figure 1 be very useful in reporting the occurrence Symmetry.—Spores trilete ; radial. of caytonian spores and others of the Shape. Spherical or nearly so ; when same sort which perhaps may not even — compressed, readjustment to a disk-like be necessarily referable to that group. form leads to formation of characteristic Whether the Caytoniales are worthy of sharp taper-point folds of variously cres- ordinal distinction is perhaps subject to centic or narrowly lenticular outline. Some- some question now that Harris (1941, times the spores are folded double so that 1940) has shown so convincingly that the whole external outline is sharply len- they are gymnospermous and have no ticular. Such folds are one of the char- particular connection with angiosperms. acteristic features of the genus. Likewise whether they should be termed pteridosperms is still perhaps too much Size.—Highly variable from about 40 a matter of individual opinion. The microns (or smaller in some abortive problem of caytonian relationship with specimens), to several hundred microns in other gymno'Spermae is a perplexing one diameter. which may find its truest solution in the Ornamentation.—Spores are character- exacting study of fossil pollen grains istically very smooth in general appear- similar to Alisporites, Parasporites, and ance ; on closer inspection they may be Pityosporites. minutely granulose or slightly rugose. There be very slight differentiation The more adequate generic definition may of proximal pyramic areas shown by this of Alisporites should be taken up in con- almost negligible emphytic marking. Larg- nection with such studies. Daugherty's er spores often possess a high gloss when generic distinction of these interesting observed by reflected light. plant microfossils is a forward step which will provide a much more precise means Haptotypic structures. — Trilete rays of referring to them. They can be dis- notably short; usually they do not ex- tinguished from spores of Parasporites ceed one-half the length of the spore by their lack of a trilete commissure and radius. The suture line is distinct and probably in their distal mode of exit; attenuate ; sometimes moderate lips are from Pityosporites in having the bladders developed. Arcuate ridges are commonly placed opposite one another and not in- not distinguishable although they may be clined distally, also in their lack of a present as slight rounded thickenings. In thick proximal cap. Although it would some forms the pyramic areas have a seem possible to recognize morphologic somewhat different surface texture than homologies between the parts of Alispo- the rest of the spore coat. rites pollen and the spores of Endospo- Spore coat.—Relatively thin. Spores rites and those of other cordaitaleans, less than 100 microns diameter are gen- these genera seem more distantly related erally yellowish and highly translucent and are easily distinguished. Pollen sim- larger spores (which certainly are mostly ilar to that of Lehachia and Walchiantkus, megaspores) are progressively less trans- now placed in the new genus Florinites lucent as the wall thickness increases rel- (p. 56), is distinguished from pollen of ative to the spore diameter. Actual wall Alisporites by the presence of an annu- thickness thus is highly variable in vari- late bladder. ous species, ranging from about 15 microns in forms over half a millimeter in diam- 1. Alisporites opii Daugherty, 1941, Carnegie eter, to less than 2 microns in those Inst. Washington Pub. 526, p. 98, pi. 34,
fig. 2. ! smaller than 100 microns. A very thin 50 PALEOZOIC FOSSIL SPORES
genus but probably not endosporal membrane may be present, referable to this all of them should be classed here. There oftentimes shrunken away from the ex- are a few forms of Punctati-sporites that osporal coat. are very similar in definable features but .J^';//Vy-—Hartung's (1933) investiga- nevertheless show differences of habit tion of spores in fructifications that were that set them apart from Calamospora. different genera allied with referred to six Some of the B3 types Raistrick has illus- provided an excellent the calamarians has trated have trilete rays more extended basis for systematic treatment of plants than they generally are in spores of Cala- represented by of this sort which are only mospora, and it is possible that some cala- their isolated spores. In general, the mosporan forms may have been included spores do not appear to lend themselves in his type Bg. Miss Knox (1938, p. to grouping into restricted generic alli- 461) has remarked on the affinity the Bg ances but show a rather generalized char- spores and compared them with spores acter. Certain species may easily be of Calamostachys binneana and Cheiro- distinguished by spore differences but sfrobus pettycurensis. generic characteristics (in terms of the Remarks. — Calamospora is unique fructifications) are not present or at least among genera typically represented by not yet recognized. The genus Calamos- plant spores in that megaspores, micro- pora therefore must be regarded as co- spores, and probably isospores, are in- ordinate with a large proportion of the cluded in it. The Calamarians seem not calamarians as they are now reported. to have developed as specialized a type In addition, Nemejc has shown that of heterospory as quickly as the lyco- spores isolated from Noeggerathiostro- pods or other groups, and it has been bus (Nemejc, 1935) and from Discinites generally doubted that they ever achieved (ibid., 1937) are similar in character and a comparable state of heterosporous de- he regards this as sound evidence of alli- velopment. Elias believes, however, that ance between these and the articulate seed-like bodies were developed in the groups although the degree of relation- group of Annularia stellata and he has ship is still difficult to evaluate. distinguished these forms as Carpannu- For the present Calamospora is re- laria (Elias, 1931). This material was garded as allied to the groups just men- obtained from beds of lower Des Moines tioned and probably is also correlative age. Such a view also was expressed by with some Mesozoic equisetaleans fcf. Renault and others of the French school remarks on Equisetosporiies, p. 47). Thus when calamites with secondary wood were it is one of the most broadly related of regarded as gymnosperms, but at least a the groups regarded as showing evidence very large part of the seed evidence then of actual natural relationship. There is was based on chance association with no question but that it would be advan- seeds that we now know have no calamite tageous to subdivide the group into sec- relationship. No seed megaspores have tions or into several genera but such a yet been reported in the bulbose bodies treatment does not seem practicable now. of Carpannularia and this or some simi- Zerndt (1934) has obtained spores of lar type of evidence will probably be nec- generally similar character by maceration essary before their seed-like nature can of coal and refers those over 200 microns be regarded as established. in diameter to his Type 2. Several dis- One feature stands out in the sporoge- tinct species of Calamospora are repre- nous sequence of the calamite alliance. sented by it. He regards these forms as More evidences of incipient heterospory pertaining to the Calamite group; how- are observable there than in any other ever, there appears to be no very definite fossil group. Apparently normal spores evidence of their generic correlation. of two sizes but otherwise similar in ap- Their character nevertheless is indicative pearance are found in adjacent sporangia. of affinity with the calamarian branch of The larger spores frequently are admixed the Articulateae. Many of the type Bg with abortive forms in various stages of spores of Raistrick (1933, 1934, etc.) are development. Sporangial masses com- —
CALAMOSPORA 51 posed of abortive and fertile spores have 50>^ 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 Fig. 1 Calamospora hartungiana sp. nov., draw- distinguishing feature. Spores in fructi- ing from microprojection of holotype. fications which are less than about 90 microns in diameter frequently are re- ]. Calamospora hartungiana Schopf, sp. nov. garded as microspores and those that ex- ceed that size often appear to represent Text figure 1. megaspores. But in different species and Description.— Spores spherical, compressed in general there seems to be no exact to polygonal or acutely lenticular out- way to distinguish between them unless line, generally with several more or less lunate the contrasting size spore is present in lenticular folds. Total spore diameter ranges from 80-100 microns where the outline is not the same fructification for comparison. foreshortened by obvious folds. Trilete rays Species distinguished within Calamo- about one-fourth of spore diameter, sutures distinct and slightly undulating, lips low but spora probably include spores of one mor- definite of from less than one to two microns phologic category only and thus hetero- width on either side of the suture. Arcuate sporous plants may oftentimes be recorded ridges may be regarded as lacking but the pyramic areas are slightly thicker and darker under two specific names. This is no by transmitted light than the rest of the spore serious disadvantage if are we interested coat and have slightly coarser surface texture in obtaining the most precise information (kontakthof ). Spore coat thin, one micron or that such material can provide. If rec- less, generally yellowish translucent or some- what red when light is transmitted through ords thus integrated are obtained, it will numerous thicknesses due to folds or the over- eventually be possible to trace the corre- lapping of several spores. Spore coat minutely
to granular ; at magnification' the lation of large and small spore forms in rugose low surface appears quite smooth. some instances. It is doubtful that the Calamospora hartungiana corresponds in gen- majority will soon be resolvable in this eral with spores Hartung (1933) obtained from fashion, however, and in the meantime fructifications of Macrostachya and Paleo- stachya, some species of which are described there is need for the most accurate re- as having a similar "kontakthof." He suggests porting of this material that is feasible. that microspores and isospores of the Calamari- ans range from 60-150 microns in diameter, and Six species previously described and spores of the species just described presumably named are listed below, one of which is would fit one of those categories. Probably queried, and one new species described. much more needs to be known of the spores of calamarian fructifications in order to be cer- Spores of this generic character are com- tain of this, but there can be no doubt as to the mon in many American coals and this general affinity of C. hartungiana. genus will be useful in providing a sys- The specimens occur fairly abundantly in the 8- to 10-inch coal of the Macoupin cyclothem tematic means of recording them. It has (middle McLeansboro age) which is exposed seemed advisable to describe C. hartungi- along Salt Fork of Vermilion River northwest ana, one of the characteristic American of Fairmount, Vermilion County, Illinois. Spores of this sort occur in spore masses 2 mm. or forms, to serve as a genotype of Calamo- more in length, as well as isolated among the spora rather than utilizing the published other materials in the residue. The holotype description of a species previously estab- specimen illustrated (text fig. 1) was separated from a spore mass of this sort when the material lished. Most of the descriptions already was being mounted in diaphane. Thus numer- published are less detailed than desired ous other specimens have also been available for comparison. The size- of the spore masses and and are still insufficiently illustrated for number of spores loosely pressed together in purposes of close comparison. them (several hundred, although the masses 52 PALEOZOIC FOSSIL SPORES obviously are only incomplete contents of single Genus Reinschospora S. W. and B., sporangia) gives a little additional information gen. nov. on the Calamospora fructification. Plate 2, figures 11-llb; text fig. 2 The holotype is from slide Y of maceration deposited in the Illinois Geological Survey 90, This genus appears to include the spores collections, Urbana. Reinsch (1884) designated as his Sub-
2. Calamospora laevigatus (Ibrahim) S. W. divisio III of his Subtribus I, Nucleus and B., comb. nov. triangularis. The genus is therefore ap- Laevigati-sporites laevigatus Ibrahim, propriately designated in honor of Aegirhorizonts, 1933, Sporenformen des pp. Reinsch. His description (loc. cit. p. 7) 17-18, pi. 4, fig. 46. of the subdivision is as follows (free Laevigati-sporites laevigatus Ibrahim, translation from Latin) : Loose, 1934, Inst. Palaobot. Arb., vol 4, Exospore thin (1/24-1/40 of the transverse no. 3, p. 146, pi. 7, fig. 2)6. diameter), smooth or ornamented with granules (Calamitif)-sporites laevigatus (Ibra- disposed variously in regular series. Body mar- him) Wicher, 1934, idem, vol. 4, no. 4, p. gins armed with a simple series (sometimes 172. duplicate) of dentiform juxtaposed spines, gradually decreasing (in length) from the cen- 3. Calamospora microrugosus (Ibrahim) S. tral margins toward the angles. W. and B., comb. nov. This diagnosis has no taxonomic status Sporonites microrugosus Ibrahim, 1932, Reinsch) Neues Jahrb., Beilage-Band 67, Abt. B., although it (and others given by p. 447, pi. 14, fig. 9. may be quite v^orthy of validation. One Laevigati-sporites microrugosus (Ibra- naturally hesitates to apply names on the him) Ibrahim, 1933, Sporenformen des basis of Reinsch's descriptions and fig- Aegirhorizonts, etc., pi. fig. p. 18, 1, 9. ures because of uncertainty as to the sources and present location of the types. 4. Calamospora mutabilis (Loose) S. W. and B., comb. nov. Also, modern interpretations may vary Calamiti( ?) -sporonites mutabilis Loose, considerably from his as to the signifi- 1932, Neues Jahrb., Beilage-Band 67, Abt. cance of certain morphologic features. B., p. 451, pi. 19, figs. 50a-c. The point which should be emphasized is Calamiti(?)-sporites mutabilis (Loose) that much of Reinsch's work is of value Loose, 1934, Inst. Palaobot. Arb., vol. 4, in extending our knowledge of spore no. p. 145. 3, form variation, particularly when essen- tial confirmation can be obtained from 5. Calamospora (?) obesus (Loose) S. W. and B., comb. nov. additional material recently prepared. Sporonites obesus Loose, 1932, Neues Thus the forms Reinsch described in his Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. subdivision III are structurally clarified 19, fig. 49. by study of American forms of Reinscho- Laevigati-sporites obesus (Loose) Ib- spora and there is little doubt that three or rahim, 1933, Sporenformen des Aegirhori- four still unnamed species have been dis- zonts, etc., p. 19. covered in his studies. However, there is Laevigati-sporites obesus (Loose) Ib- no particular occasion to propose names rahim, Loose, 1934, Inst. Palaobot. Arb., for these until similar new material is at vol. 4, no. 3, p. 145. hand or the source defined for his old 6. Calamospora pallidus (Loose) S. W. and types (if they are preserved) and their B., comb. nov. character confirmed. The following new Sporonites pallidus Loose, 1932, Neues generic diagnosis has been constructed Jahrb., Beilage-Band 67, Abt. B., p. 449, on the basis of newer work but it also pi. 18, fig. 31. takes cognizance of the structural fea- Punctati-sporites pallidus (Loose) Ib- tures unmistakably shown by Reinsch. ^ rahim, 1933, Sporenformen des Aegirhor- izonts, p. 21. Symmetry.—Spores radial, trilete. Punctati-sporites pallidus (Loose) Ib- Shape.—Body lenticular to flattened el- rahim, Loose, 1934, Inst. Palaobot. Arb., liptical in the axial plane; subtriangular vol. 4, no. 3, p. 146. in transverse plane, corners rounded, sides slightly convex or concave in outHne. A 7. Calamospora perrugosus (Loose) S. W. fimbriate flange, broadest in the inter- and B., comb. nov. «, Laevigati-sporites perrugosus Loose, radial region and very narrow or absent 1934, idem, p. 145, pi. 7, fig. 13. on the corners, may be present to modify —
REINSCHOSPORA 53 the marginal outline. In some forms the 50m 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- Fig. 2 Reinschospora hellitas sp. nov., drawing of holotype. preted as equivalent to an arcuate ridge since, contrary to usual arcuate ridge de- can be identified with Botryococcus. velopment, it is least evident opposite the Reinsch's work still is significant and con- ends of the rays. Reinsch has illustrated tinues to command respect for his volu- forms with granules in rows paralleling minous observations. the trilete rays; these may have a hapto- typic origin, and might be equivalent to 1. Reinschospora bellitas Bentall, sp. nov. ridges in of formation. arcuate mode Text figure 2 Spore coat.—Moderately thin (less than Description.—Compressed spore subtriangular ex- 3 microns) and of uniform thickness in outline with inter-radial margins usually con- cept at the equator where it is joined by cave. Diameters vary from 57 to 76 ix, with an the flange. average of about 68 ix. Spore coat less than 2 fx in thickness and levigate. Trilete rays extend Affinities. Unknown possibly filicin- — ; four-fifths or more of the distance to the ean. The similarities in morphology sug- equator. Flange usually originates slightly gest that this is a group having natural proximal from the spore equator. Fimbriate elements of flange number about 50 between significance. It may be related to Granu- ray extremities and have a maximum length of lati-sporites. 28 II midway between rays and a minimum Remarks.—The genus is reported from length of four to five at the ends of the rays. Incisions of flange, when present, apparently central Russia by Reinsch (1884, his di- due to imperfect preservation. agnoses 62-66, incL), from the Ruhr by Reinschospora hellitas has been isolated from Loose (1934), and from Lower Pennsyl- Angel and Battle Creek seams of the southern vanian coals of Tennessee by Bentall. Tennessee coal field. Though never numerically Dolgner (1932, pi. 28, fig. 4) has illus- abundant, it is present in nearly all samples. It is very similar to R. speciosa (Loose) but is trated a form with marginal spines, from somewhat larger in size. Occurring with R. coal, which is congeneric Moscow brown bellitas is a form essentially comparable in all with our interpretation of Reinschospora; characters except that the flange is absent. Brokaw has recently found somewhat The relationship, if any, between these forms is not known. similar forms in the coal of the Bogota Holotype text figure 2. Tennessee Division cyclothem (upper McLeansboro age) in — of Geology collection ; Battle Creek seam, north the Pennsylvanian Illinois. of The geno- side of Sweden Cove, Marion County, Ten- type species described below is based on nessee. the Tennessee material. The generic name honoring Reinsch seems appropri- 2. Reinschospora speciosa (Loose) S. W. and B., comb. nov. ate inasmuch as Blackburn and Temperley Alati-sporites speciosiis Loose, 1934, (1936) have shown that the algae for- Inst. Palaobot. Arb., vol. 4, no. 3, p. 151, merly known as Reinschia probably now pl. 7, fig. L 54 PALEOZOIC FOSSIL SPORES
Genus Lycospora S. W. and B., gen. nov. nized the relationship of these types to Lepidostrobus and compared D^ and D2 Plate 3, figures 19-19b with Lepidostrobus jacksoni and L. old- Symmetry.—Spores radial ; trilete. hamius respectively. Type D^^ which she Form.—Flattened lenticular as seen in illustrates may also belong to Lycospora. lateral profile, equator marked with a There is, of course, great inherent un- short thick tapering ridge ; nearly circu- certainty in identifying any species of lar or very broadly triangular in the Lepidostrobus from isolated microspores transverse plane. Compression is gener- alone. Microfossils of this kind obtain- ally parallel to the transverse plane with able from coal probably are derived from few or no folds, due to preferential ori- many more species than are likely to be entation of the spores. recognized from entire fructifications be- Size.—Mean diameter in various spe- cause cones are essentially more fragile cies ranging from about 18 to 45 microns. and require more favorable conditions for Ornamentation. — Spores nearly their good preservation than spores do. The spores probably constitute a more smooth ; minutely granulose or rugose. compete record Sometimes the minute rugae tend to be of the various closely re- lated forms, more prominent radially. whether they can be distin- guished easily from one another or not. Haptotypic features.—Trilete rays ex- Reinsch has tended nearly to margin, suture lines dis- (1884) illustrated a num- ber of forms, chiefly tinct, usually without noteworthy differ- from central Russia and from Zwickau, Saxony, entiation of lips. In species with most which are probably referable strongly developed emphytic ornament, to Lycospora. His numbered diagnoses the haptotypic features also are more 32, 81, 83, 87, 240 424 525 prominent. The equatorial ridge corre- ( ?), ( ?) and 550 may be cited. sponds to the arcuate ridge in its origin Remarks.—Four species previously de- and rarely becomes so extended and mem- scribed are assigned to Lycospora all of branous across its width as to resemble which represent new name combinations. a typical flange development. Usually it Lycospora micro papillata (Wilson and is narrow and tapers evenly and rapidly Coe) is designated as the type species. away from the spore body; in some spe- The genus is well represented in the Penn- cies it may be almost nonexistant. sylvanian; several additional species have been recognized but are not Spore coat.—Relatively thin, sometimes yet described. less than 1 micron thick on proximal and 1. Lycospora micropapillatus (Wilson and distal surfaces and quite translucent. Coe) S. W. and B., comb. nov. Thicker at the equator where there is no Cirratriradites micropapillatus Wilson and easily defined line of demarcation between Coe, 1940, Am. Midland Naturalist, vol. 23, no. p. 184, fig. the equatorial ridge and the body wall. 1, 6. Affinity.—Spores of this type have fre- 2. Lycospora minutus (Wilson and Coe) S. W. and B., comb. nov. quently been encountered in the tips of Cirratriradites minutus Wilson Lepidostrobus cones and there is little and Coe, 1940, idem, p. 183, figs. 11 and 12. doubt that most of these forms found iso- lated represent microspores of the arbor- 3. Lycospora pellucidus (Wicher) S. W. and B., eous lepidodendrids. Andrews and Pan- comb. nov. Sporitcs nell (1942) have described similar forms pellucidus Wicher, 1934, List. Palaobot. Arb., vol. 4, no. 3, i^p. 186, pi which persist in tetrad ' . 1^ groupings as mi- 8, fig. 29. crospores of a species of Lepidocarpon. 4. Lycospora The microspores of sigillarians (as shown pusillus (Ibrahim) S. W. and B., comb. nov. by Mazocarpon) are significantly larger Sporonites pusillus Ibrahim, 1932, Neues and probably are not included in Lyco- Jahrb., Beilage-Band 67, Abt. B., p. 448, spora (cf. Schopf, 1941). pi. 15, fig. 20. Spores assigned to type D^^ by Raistrick Zonales-sporites pusillus (Ibrahim) Ib- rahim, belong without question to Lycospora, 1933, Sporenformen des Aegirhor- izonts, p. 32, pi. 2, fig. 20. also probably his B^ and D2 Type in part Granulati-sporites pusillus (Ibrahim) (Raistrick and Simpson, 1933, Raistrick, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 1937, 1938). Knox (1938) has recog- 3, p. 146, pi. 7, fig. 3. ;; .
RAISTRICKIA 55
Genus Raistrickia S. W. and B., ever, is somewhat questionable. Forms gen. nov. Knox (1938) illustrates as D3 and Dg probably belong in this genus. Reinsch Plate 3, figures 18-18b; text figure 3 (1884) has illustrated only three forms which belong to this group without much Symmetry.—Spores radial; trilete. doubt. These are described under his Sperical to slightly flattened in Shape.— diagnoses 52, 308, and 588 from central transverse plane and slightly triangular. Russia and from Zwickau, Saxony. The plane is Little preference as to shown by form illustrated by Thiessen (1920) in compressed specimens. plate 61, figure B, from Benton, Illinois Size.—Spores of diiTerent species vary (probably the Herrin No. 6 coal), is from about 40 to 90 microns in mean di- characteristic of Raistrickia; that shown ameter. above it in horizontal thin section may Ornamentation.—Characteristically ver- likewise belong here but it is impossible to be sure. The two illustrations show rucose or spinose ; spines when present are generally heavy, abruptly truncate, the inadequacies of thin sections for iden- blunt tipped- Close observation often tification of spores. shows the spine tips to be minutely dis- Remarks.—Five species previously de- sected into two to six terminal papillae. scribed are assigned to Raistrickia. These The prominences of verrucose forms are forms are widely distributed as may be similar to the spines but shorter. Except inferred from the preceding discussion. for the trilete structure, features of em- None of these previously named are phytic ornamentation extend fairly evenly very suitable for designation as a type, over the whole spore surface. however, inasmuch as the more typical form, which particularly seems generic- Haptotypic features.—Trilete rays va- ally riable in length as in Punctati-sporites distinct from Punctati-sporites, has not previously been formally generally inconspicuous, lips unorna- designated although it mented. Rays sometimes slightly undu- seems fairly common. Fur- thermore, lant due to proximity of spine bases inadequacies in the previous arcuate distinctions absent. descriptions and illustrations also mih- tate against selection of any of them as Spore coat. — Usually of moderate types for this genus. The species de- thickness (two to six microns) perisporal scribed below, Raistrickia grovensis sp. membrane lacking. nov. is therefore designated as the type. Affinity.—Typical forms of Raistrickia 1. Raistrickia (?) abditus (Loose) S. W. are without doubt filicinean. Spores of and B., comb. nov. Senftenhergia plumosa, as identified by Sporonites abditus Loose, 1932, Neues Radforth (1938), agree with these of Jahrb.. Beilage-Band 67, Abt. B., p. 451, Raistrickia, and similar schizaeaceous pi. 19, fig. 53; Verrucosi-sporites abditus (Loose) sporangia obtained by Andrews and Loose 1934, Inst. Palaobot. Arb., vol. 4, no. Illinois Schopf from shale above coal No. 3, p. 154. 6 also contain spores of this type. How- 2. Raistrickia fibratus (Loose) S. W. and ever, the spores of a form identified as ^. B., comb. nov. pennaeformis by Radforth (1939) show Sporonites fibratus Loose, 1932, Neues rather different features, and those of Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 6". sturi do not appear congeneric with 19, fig. 52. Setosi-sporites fibratus (Loose) Loose, Raistrickia. Although the general rela- 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. tionship cannot easily be doubted, the 148. specific features do not permit easy cor- 3. Raistrickia grovensis Schopf, sp. nov. relation and Senftenhergia and Raistrickia Text figure 3 no doubt represent generic groups of — somewhat different circumscription. Description. -Spore rounded triangular in out- line, about 50 microns in diameter. Orna- Raistrick's types E3 and E^ (Raistrick mented with numerous short (3-5 /x) broad
(2-6 iJi) blunt tipped spines, and Simpson, 1933 ; Raistrick, 1934, 1935, which by juxtapo- sition sometimes appear to be mutually 1937) belong in Raistrickia. His Eg type attached. Rays 15-20 ix long, slightly undulating, lips not is particularly characteristic. rep- That raised but forming a narrow margin on either resented as E^ by Knox (1938), how- side of the suture line. Wall somewhat vari- —
56 PALEOZOIC FOSSIL SPORES
able (3-4 ix) in thickness, appearing brownish Setosi-sporites superbus Ibrahim, 1933, translucent ; spines are darker. Sporenformen des Aegirhorizonts, p. 27, pi. 5, fig. 42. 50/^- 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
Fig. 3 Raistrickia grovensis sp. nov., camera is marked by numerous sharp angular lucida drawing of holotype. folds, especially around its periphery. Sise. Greatest diameter (length of This form, in general, resembles the spores — of Radforth's (1938) Senftenbergia plumosa hut bladder) among different species varies the spines are shorter, broader and less crowded from about 50 to 180 microns ; the pollen and the spores he illustrates are round in outHne. body proper varies from about 20 to 110 It is difficult to make a precise comparison with microns in diameter in individuals of vari- other previously described species now assigned to Raistrickia because of inadequacies of their ous species. description and illustration. The spine tips of Ornamentation. — Bladder ornament this form are entire and rounded ; others from similar to that of Endosporites and Pity- the same coal bed and elsewhere show spine tips slightly dissected. No doubt several species osporites; exterior surface smooth or very of schizaeaceous ferns are represented. The finely granulose or rugose, internally the specimen shown in text figure 3 designated as membrane bears a distinct reticulation net holotype, is from the Herrin (No. coal (up- 6) v^^hich tends to be obsolescent in the cen- permost Carbondale age) which is worked by tral area. extensive open-cut mining operations near Mid- proximal Bladder and body dle Grove, Illinois, in northern Fulton County. walls are joined distally, and centrally from this juncture there is no evidence of 4. Raistrickia saetosus (Loose) S. W. and reticulation. B., comb. nov. Sporonites saetosus Loose, 1932, Neues Haptotypic features. — Generally not Jahrb., Beilage-Band 67, Abt., B., p. 452, evident, and probably always extremely pi. 19, fig. 56. obscure. Trilete imprint (when discern- Setosi-sporites saetosus (Loose) Ibra- able) is wholly vestigial, probably located him, 1933, Sporenformen des Aegirhori- entirely on the bladder, and has no func- zonts, p. 26. tional significance in the mature spore. Setosi-sporites saetosus (Loose) Ibra- him, Loose, 1934, Inst. Palaobot. Arb., vol. The effect of the haptotypic relations 4, no. 3, p. 148. sometimes may be obscurely expressed in the proximal bladder reticulation pattern. 5. Raistrickia spinososaetosus (Loose) S. W. and B., comb. nov. Spore coat.—Bladder membrane is ex- Sporonites spinososaetosus Loose, 1932, panded on all sides of the body except Neues Jahrb., Beilage-Band 67, Abt. B., for a small distal area. The bladder mem- p. 452, pi. 19, fig. 55. brane generally has a very well defined Apiculati-sporites spinososaetosus reticulation network located on the inner (Loose) Ibrahim, 1933, Sporenformen des surface. At the folded margin the blad- Aegirhorizonts, p. 24. Apiculati-sporites spinososaetosus der membrane commonly appears thicker (Loose) Ibrahim, Loose, 1934, Inst. Palao- than it actually is due to the reticule bars
bot. Arb., vol. 4, no. 3, p. 153. appearing in profile ; the membrane proper usually is less than a micron thick. Con- 6. Raistrickia superbus (Ibrahim) S. W. and B., comb. nov. sequently, it is highly translucent between FLORINITES 57 the bars of the reticulum which may be tural similarity in the two instances is extended as much as 2 or 3 microns away entirely remote. from the membrane surface. The body These gymnospermic groups, often- wall inside the bladder is everywhere thin, times regarded as distinct, thus have a oftentimes less than a micron, and shows community of relationship which is mea- no structural differentiation. Because of gerly expressed for the present in the it not be exosporal its tenuousness, may genus Florinites. The probability is that in nature although it would seem rather this group will also be of continuing value precisely equivalent to the thicker and for classification of these spores. If presumably exosporal body wall of Endo^ plants of the cordaito-coniferous type sporites. The body wall of Florinites is were partially restricted to upland habi- slightly less translucent than the bladder tats their pollen grains may offer the only due in considerable part, at least, to its widespread available record of their ex- folds. rather numerous istence. For this reason it is essential Affinity.—Florinites is known to be re- that they be classified as precisely as pos- lated to Cordaianthus (pars), Lehachia, sible.
Ernestiodendron and Walchianthus Florinites is also related to Endospo- through the investigations of Florin rites, Pityosporites, Alisporites, and Para- (1936, 1938-40); thus it seems evident spontes, all of which have the character- of the upper Pale- that it represents a part istic bladder-forming perisporal (?) mem- ozoic gymnospermic alliance. brane. The study of these in conjunc- That some cordaitaleans possess pollen tion with fructifications and other organs grains of the Endosporites type is shown with which they are more or less definitely by Cordaianthus cf. C. shuleri Darrah correlated cannot fail to add greatly to represented in collections made by Wilson. our knowledge of gymnospermic phylo- Similar pollen grains are shown by Oliver geny during a critical period in the evo- (1940) to be present in the pollen cham- lution of this plant group. The defini- ber of ''Stephanospermum" caryoides tion of the age at which these genera are which, according to Florin (1937, p. 310), first recognizable will be of geologic as is no doubt a cordaitalean seed. Oliver's well as botanical significance. figures 40 and 42 show a characteristic, Remarks.—This genus is proposed to and no doubt functional, trilete apparatus. apply to gymnospermic plants whose pol- Florin's study o{ Cordaianthus saportanus len generically corresponds with the type pollen and pollen grains of Florinites species as expressed, in general, by the type associated with it (his Cordaianthus foregoing definition. Florin has given sp. 1 and 2) indicates that the suture was rather similar data in description of the entirely absent and the trilete marking species ''cordaitiformis" which he places completely vestigial. In the walchian under the genus name Pollenites (see be- vestigial conifers not even a remnant of low). There is little doubt that such a gen- the haptotypic marking has been reported eric designation is inadequate but question but it might be present (just as in the may arise concerning the validity of the Cordaianthus sp. 2) and still not be easily specific epithet. demonstrable from coalified compression Pollenites cordaitiformis was estab- material. The agreement in pollen struc- lished to include pollen which elsewhere ture between the Cordaianthus material in his monograph Florin had assigned to studied by Florin, and his walchians is five other species of plants. The only such as to leave little doubt that a direct illustrations given to validate the name relationship exists between them. Just had also previously been allocated differ- how this relation evolved and should be ently to the other five species. Florin's systematically expressed cannot now be photographs still serve as an excellent stated. The pollen structure is so spe- basis for discussion of the character cialized, particularly when compared with of isolated Florinites the expression of spore characteristics pollen but none of elsewhere in the plant kingdom, as to per- those he illustrated are properly identified mit no conclusion other than direct re- with this genus because no single organ- lationship. The possibility of convergent ism can have more than one valid name. evolution being responsible for the struc- Specimens properly assigned to Lehachia, 58 PALEOZOIC FOSSIL SPORES
Ernestiodcndron, or Walchianthus can- are evident. When compressed laterally the axial dimension is observed to be shorter than not in the same instance also be assigned that in either of the other two planes of sym- to some different genus and species. Fur- metry. Inclusive of the bladder, the fossils thermore, it seems an absurdity to at- range from 55-90 (mostly 65-85) microns in tempt to make an identification with a length, and 40-75 (mostly 45-60) microns in breadth. The axial dimension, exhibited in a generalized group such as Pollenites or few instances, is a little less than 40 microns. precise even Florinites, when a more spe- The body, enclosed by the bladder on all but cific pertinence is thoroughly demon- the distal side, is more nearly spherical ranging strated. The systematic problem that from 25-45 (mostly 27-37) microns in length and 20-40 (mostly 25-35) microns in breadth. confronts us revolves around proper de- The axial dimension of the body is somewhat fossils where pertinence termination of less, about 20-25 microns. The body proper is to a definitely restricted group is, for always compressed with sharp folds which are one reason or another, legitimately in characteristically more numerous around the
periphery ; the bladder is oftentimes hardly question. Thus it would seem that unless folded when compression is in the proximo- of the pollen specimens Florin has one distal plane, but the folding that does affect it illustrated is regarded as improperly re- is more erratic. When compressed obliquely ferred to species of Lehachia, Ernestio- or laterally the outline is much distorted. The dendron, or Walchianthus, none of them bladder membrane itself is less than a micron thick ; a zone a few microns broad around can be regarded as type material of a new the equator is more definitely yellowish trans- is distinct taxonomic species. As there lucent and slightly thicker ; compressed laterally no discernible reason to question Florin's the equatorial zone of the bladder remains dis- more precise assignment of that material, tinguishable and the slight inequality in thickness evidently contributes to the more erratic char- it seems equally evident that Pollenites acter of folds. The bladder is smooth to min- cordaitiformis has no type and the name utely granulose externally but shows an internal was superfluous as proposed. reticulation net. The reticulae vary from 1 to 3 microns in diameter and may be very slightly Florinites antiquus sp. nov. is de- elongated radially. The bars of the reticulation scribed below and designated as the type are not prominent in this species and have a species. Two other species previously thickness of a fraction of a micron. The reticu- described also are tentatively assigned to lation is more pronounced away from the body and is hardly discernable immediately surround- the genus. There can be little doubt that ing the "contact area" on the distal side or cen- Types 5 of Millott (1939) and 3K of trally on the proximal side. The body wall is Knox (1942) are also congeneric. Flor- essentially smooth, very thin, and except where in's various descriptions and excellent il- folded into several thicknesses, it hardly darkens the light straw-yellow translucence of the cen- lustrations have shown clearly that a tral area of the pollen grain. The "contact number of other species can be distin- area" where bladder joins the body wall is guished by their pollen characters. elliptical or oval with its greater diameter commonly transverse to the length of the pollen Florinites is known to be widely dis- grain. The margins of this area are quite faint tributed in the Pennsylvanian and Upper and its size and outline vary from 17 x 23 to Carboniferous as well as in the Permian 10 X 20 microns. It can hardly be detected under strata where other coniferous remains are the microscope unless the grain is oriented distal side up. A distinct but tenuous striation associated with it. Pollen of this char- traverses the "contact area" in many instances, acter has been found to be abundant in continues on the bladder membrane until about some Tennessee coals and other beds of opposite the body margin, and is then lost as lower Pottsville age. It seems likely that the bladder membrane developsi its more dis- tinctive ornamentation. It can hardly be in- a detailed study of these forms will add terpreted as a monolete suture line ; it would much to our of knowledge the early an- seem more plausibly regarded as a harmome- cestry of conifers. gathic groove similar in character to those on the distal side of Monoletes prepollen but more faintly developed, since the body and bladder 1. Florinites antiquus Schopf, sp. nov. membranes are not nearly so thick as the Monoletes coat. The striation seems correlated Text figures 4, 5 in position with the long axis of the contact area and frequently is set obliquely with ref- Description.—Pollen grains, bilateral symmetry, erence to the breadth of the grain as a whole. equipped with an annulate bladder joined to the central pollen body only on the distal side, Haptotypic characteristics are very faint and thus outlining a "contact area" equivalent to a may be observed to best advantage in forms in germinal "furrow." The grains are broadly which the body membrane is wanting, but the elliptical when compressed in the proximo-distal bladder membrane persists (perhaps as a result plane so that the length and breadth proportions of maceration). In such examples, which are — —
FLORINITES 59
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- 50^^- lished among the upland vegetation of the period.
Fig. a—Florinites antiquus sp. nov., drawing from microprojection of holotype. Body Bladder LENGTH P 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 U^ 20 30 40 6 60 70 80 90 ytc of the bladder membrane (it is too faint to be recognizable as reticulation) also seems faintly o 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. Body LJ Bladder Florinites antiquus seems distinguishable from all the similar forms Florin (1936, 1938-40) has BREADTH identified with Cordaianthus, Lebachia, Ernestio- dendron, and Walchianthus, on the basis of its Fig. 5 Florinites antiquus, coordination of size. F. antiquus is evidently significantly length-breadth measurements from 23 specimens, smaller judging from measurements Florin has grouped according to 5 micron increments. (Di- given and by dimensions measured directly mensions modified by unusual folding have been published. His from the illustrations he has omitted). Cross-lined areas represent measure- from the Stephanian of Cordaianthus sp. 1, ments from the holotype. France compares most closely in size, (bladder
length, 78-90 ; width, 62-67 microns ; body F. antiquus has been obtained in moderate length, 42-50; width 40-48 microns). The dif- abundance from a widespread 10 to 12-inch ferences can best be evaluated by reference to coal in the cyclothem where it crops out the assembled measurements of F. antiquus Wiley along Soap Creek southeast of the town of presented in text figure 5. Presumably the Carbon, Davis County, Iowa. Correlation with flattening of F. antiquus pollen grains would the widespread Wiley coal of western Illinois give rise to dimensions which are slightly ex- has recently been confirmed by L. M. Cline of cessive in the plane of compression and there- State College, has measured the fore not quite comparable to measurements Iowa who section, this coal to be in proper se- taken from Florin's uncompressed forms. That showing quence and about 12 feet above the lower Sea- the species are distinct is evident from the much borne coal and limestone developed in this area. less pronounced bladder reticulation in F. an- The Wiley cyclothem is near the top of the tiquus. The distal striation has not been noted Tradewater group of Illinois and is considered in forms regarded as cordaitean but is evident to be lower Allegheny in age. (though no mention is made of it) in several of Florin's figures of Paleozoic coniferous pollen, The holotype of F. antiquus, illustrated in (cf. Florin, 1939, pis. 55-56, fig. 21; Florin, text figure 4, is from maceration 413, slide 8 1940, pis. 145-146, fig. 19; ibid, pis. 147-148, (unstained) in the Illinois State Geological fig. 5). Survey collections in Urbana. From its age (early Allegheny) one might (Ibrahim) S. assume that F. antiquus belonged with the 2. Florinites (?) pumicosus nov. cordaitealeans, but its features are advanced on W. and B., comb, 60 PALEOZOIC FOSSIL SPORES
Sporonites pumicosus Ibrahim, 1932, 3. Sporites hirmeri Reissinger, 1939, Paleon- Neues Jahrb., Beilage-Band 67, Abt. B., tographica, vol. 84, Abt. B, p. 16. p. 447, pi. 14, fig. 6. Note.—Nomen nudem, no illustration. Reticulata-sporites pumicosus (Ibrahim, 1932) Ibrahim, 1933, Sporenformen des 4. Sporites irregularis Reissinger, 1939, Paleontographica, vol. Aegirhorizonts, p. 38. 84, Abt. B, p. 17. Reticulata-sporites pumicosus (Ibrahim, Note.—Nomen nudum, no illustration. 1932) Ibrahim, 1933, Loose, 1934, Inst. 5. Sporites plicatus Schopf, 1938, Illinois Palaobot. Arb., vol. 4, no. 3, p. 157, pi. 7, Geol. Survey Rept. Inv. 50, p. 51, pi. 7, fig. 29. figs. 7-9.
3. Florinites (?) visENDUS (Ibrahim) S. W. 6. Sporites problematicus Zerndt, 1934, and B., comb. nov. Acad, polonaise sci. Trav. Geol. no. 3, Reticulata-sporites visendus Ibrahim, Krakow, p. 27, fig. 13, pi. 30, figs. 1-10. 1933, Sporenformen des Aegirhorizonts, p. 7. Sporites spongiformis Reissinger, 1939, 39, pi. 8, fig. 66. Paleontographica, vol. 84, Abt. B, p. 17. Note.—Nomen nudum, no illustration. Incertae sedis 8. Sporonites biporous Berry, 1937, Am.
Midland Naturahst, vol. 18, no. 1, p. 160, In the present state of knowledge there fig. 14. are a number of named forms which it is 9. Sporonites tripterus Berry, 1937, Am.
impossible to treat systematically. The Midland Naturalist, vol. 18, no. 1, p. 160, various generic designations Hsted below fig. 8. may possibly have a certain value but in 10. Zonala-sporites taciturnus (Loose) any event their significance does not ap- Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. p. 157. pear to be biological. The names Spor- 3, Sporonites taciturnus Loose, 1932, ites (H. Potonie, 1893), Pollenites, and Neues Jahrb., Beilage-Band 67, Abt. B., Sporonites (R. Potonie, 1931) cannot be p. 450, pi. 18, fig. 38. regarded as having systematic value. 11. Zonala-sporites ulughbeki (Loose), Ib- Forms hsted in these groups have no es- rahim, 1933, Sporenformen des Aegir- sential biologic features in common, no horizonts, p. 38, pi. 1, fig. 11. type species are recognized, and no sys- Sporonites ulugbeki Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 447, tematic treatment is feasible on this ba- pi. 14, fig. 11. sis. A morphologic designation apparently 12. Elongato- SPORITES RETicuLATUS Berry, would have served as well to record the 1937, Am. Midland Naturalist, vol. 18, lack of systematic information concern- no. 1, pp. 158, 160, fig. 12. ing most material of this character. It 13. Verrucosa-sporites PERVERRUcosus (Loosc) likewise seems impossible to attach any Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. generic significance to the names Verru- 3, p. 157. cosa-sporites (Loose, 1934), Elongato- Sporonites perverrucosus Loose, 1932, sporites (Berry, 1937), Punctata-sporites Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pi. 18, fig. 48. or Zonala-sporites (Ibrahim, 1933). Such problematic forms as have been listed 14. PuNCTATA-SPORiTES SABULOSUS Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. under these names must await future dis- Z7, pi. 5, fig. 43. position. In addition, the following types of
1. Sporites echinospinosus Zerndt, 1937, Raistrick and Knox may be listed, for Acad, polonaise sci. Trav. Geol. no. 3, they too are of uncertain afifinity at the Krakow, p. pi. 24, figs. 5-10. 17, present time. Some of these seem to be worthy of more appropriate classification. 2. Sporites fumosus Schopf, 1938, Illinois Raistrick (1937, p. 911) B^, B^, E2, E^, Geol. Survey Rept. Inv. 50, p. 52, pi. 5, fi'>-s. 1-2. Gi, Knox (1942, pp. 100-101), D^, K^. SUMMARY 61
SUMMARY fossils it seems unavoidable that many forms worthy of description will never- This report presents the revised classifi- theless have such problematic relationship cation of plants identified from their that their assignment under incertae sedis isolated fossil spore coats, and deals is obligatory. The usefulness of fossils primarily with those of Paleozoic age. nevertheless bears a considerable relation- Orthodox taxonomic procedure has been ship to the basic and fundamental in- followed as closely as possible for reasons formation available about them and for which are presented in the introduction. this reason greatest significance must be A cautious policy has been followed with attached to species whose relationship regard to synonymy, and, even though has been reliably established. forms are very closely allied, names based The authors of this paper and their on different holotypes have been allowed associates have observed many types to stand. In many instances, however, new of plant microfossils in preparations from the apparent close relationship, which may coal and carbonaceous sediments in later prove conspecific, has been noted. America. The present synopsis is the out- Thus names noted as synonyms are nearly growth of a need for a more compre- all objective synonyms because they are hensive survey of previous work con- based on a common holotype. Many sidered from the standpoint of a con- difficult problems concerning nomencla- sistent systematic poHcy. The essential ture of fossil plants are solvable if nomen- features of an appropriate policy have clatural types are strictly interpreted. been embodied here to serve as a working About 400 named species have been in- basis for the great amount of descriptive cluded in the present paper and most of work yet to be done. We believe that this these have been allocated among 23 genera working basis will require further re- which seem to serve a useful and signifi- vision as new information is accumulated cant purpose in classification. Additional and presented. We further believe that genera no doubt will require recognition such revision can be carried out with later and new information will modify greatest efficiency and benefit to all con- the views that have been expressed for the cerned if the orthodox usages character- genera described here. A number of istic of mature systematic science are species described previously apparently adopted and critically applied. do not conform sufficiently to merit in- clusion in the same genus with the type ACKNOWLEDGMENTS species of the group, and at the same time The authors are under particular obliga- do not show convincing evidence of affinity tion to Dr. H. Camp of the New York with other recognized groups. Such forms W. Botanical Garden who, at a period when have been listed as species excludende. other duties were particularly urgent and Attention is directed to the several species pressing, nevertheless found time to read excluded from Reticulati-sporites which the manuscript and suggest numerous here is interpreted in a considerably re- desirable revisions in the manner of pres- stricted sense. Many of these forms re- entation. quire much more careful study in order to arrive at a satisfactory expression of their Mr. R. M. Kosanke of the Illinois affinity and classification. Other forms Survey, Dr. Gilbert H. Cady, and mem- listed under incertae sedis also are lacking bers of the Survey editorial staff all have in sufficiently understood biological char- contributed in final preparation of this acteristics to support a definite systematic paper for publication. To all of these the allocation. In dealing with plant micro- authors owe a special debt of thanks. : :
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— , 1921, Origin and composition of certain
Rousseau, A., 1935, Etude de quelques types oil shales : Econ. Geology, vol. 16, pp. 289-
de spores du Westphalien Inf erieur : Mus. 300. royal histoire nat. Belgique Bull., vol. 11,
no. 21, pp. 1-6. — , 1925, Microscopic examination of Ken-
tucky oil shales : Kentucky Geol. Survey, Sahabi, Yadollah, 1936, Recherches sur les spores ser. 6, vol. 21, pp. 1-47. des houilles Francaises, Douriez-Bataille,
Lille, 62 pp. — , 1930, Splint Coal : Am. Inst. Min. Met. Eng. Trans., Coal Division, 1930, pp. 644- Schopf, James M., 1936, The paleobotanical 672.
significance of plant structure in coal : Illi- vol. nois State Acad. Sci. Trans. (1935), — , and Sprunk, George, 1936, The origin of 28, no. 2, pp. 106-110. the finely divided or granular opaque matter
in splint coals : Fuel in Science and Prac- (no. , 1938, Spores from the Herrin 6) tice, vol. 15, no. 11, pp. 304-315.
coal bed in Illinois : Illinois Geol. Survey Rept. Inv. 50, 7?) pp. — , and Stand, J. N., 1923, Correlation of coal beds in the Monongahela formation of
Lepidocarpaceae : — , 1941, Notes on the Am. Ohio, Pennsylvania and West Virginia Midland Naturalist, vol. 25, no. 3, pp. 548- Coal-mining Investigations, Carnegie Inst. 563. Tech., Bull. 9, 64. pp.
, 1941, Contributions to Pennslvanian — -, and Wilson, F. E., 1924, Correlation of Paleobotany Mazocarpon oedipternum, sp. coal beds of the Allegheny formation of nov. and Sigillarian relationships : Illinois western Pennsylvania and eastern Ohio Geol. Survey Rept. Inv. 75, 53 pp. Coal-mining Investigations, Carnegie Inst. Tech., Bull. 10, 56. pp. Scott, D. H., 1898, On the structure and affinities of fossil plants from the Paleozoic rocks. Thompson, R. B., 1927, Evolution of the seed
II. On Spencerites, a new genus of Lyco- habit in plants : Royal Soc. Canada Trans,
podiaceous cones : Trans. Royal Soc. Lon- ser. 3, vol. 21, pp. 229-272. don Philos., ser. B, vol. 189, 1897, pp. 83-106. Virkki, Chinna, 1937, On the occurrence of Seward, A. C., 1914, Antarctic fossil plants winged spores in the Lower Gondwana rocks Nat. Hist. Report British Antarctic ("Terra of India and Australia: Indian Acad. Sci. Nova") Exped., 1910. Geology, vol. 1, no. Proc, vol. 6, no. 6, sect. B, pp. 428-431.
1, pp. 1-49.
-, 1939, On the occurrence of similar spores 398-9. , 1919, Fossil Plants, vol. IV, pp. in a Lower Gondwana glacial tillite from Australia and in Lower Gondwana shales in
: Indian Acad. Sci. Proc, vol. no. ; or India 9, , 1933, An Antarctic pollen-grain fact no. 1, sect. B, pp. 7-12. fancy ? : New Phytologist, vol. 32, 4, pp. 311-313. White, David, 1905, The Geology of the Perry
Singh, T. C. N., 1932, Notes on the fossil spores Basin, Paleontology : U. S. Geol. Survey Prof. Paper 35, 35-84. in an oil shale from Tasmania : Royal Soc. pp. Tasmania Papers and Proc, 1931, pp. 32-36.
, 1929, Description of fossil plants found of petroleum Sprunk, G. C., et al., 1940, Splint coals of the in some "mother rocks" from Appalachian Region; their occurrence, pe- northern Alaska : Am. Assoc Petroleum trography, and comparison of physical and Geologists Bull., vol. 13, no. 7, pp. 841-848. chemical properties with associated bright
coals : U. S. Bur. Mines Tech. Paper 615, and Stadnichenko, T., 1923, Some mother 59 pp. plants of petroleum in the Devonian black
shales : Econ. Geology, vol. 18, pp. 238- Stach, E. and Zerndt, Jan., 1931, Die Sporen in 252 (reprinted by Kentucky Geol. Survey, den Flamm-, Gasflammen und Gaskohlen des 1925, ser. 6, vol. 21, pp. 99-117.) :
66 PALEOZOIC FOSSIL SPORES
Wicher, Carl A., 1934a, Sporenformen der — , 1930b, Megasporen aus einem Floz in
Flammkohle des Ruhrgebietes : Inst. Pala- Libiaz (Stephanian) : Acad, polonaise sci. obotanik u. Petrog. der Brennsteine Arb., Bull. Internat., ser. B., Annee, 1930, pp. vol. 4, no. 4, pp. 165-212. 39-70.
1934b, Abortiverscheinungen bei , , Uber — 1931, Megasporen als Leitfossilien des
fossilen sporen und ihre Phylogenetische produktiven Karbons : idem, ser. A, Annee,
Bedeutung : Inst. Palaobotanik u. Petrog. 1931, pp. 165-183. der Brennsteine Arb., vol. 5, no. 3, pp. 87-96.
— , 1934, Les megaspores du bassin houiller
Williams, H. S., 1887, On the fossil faunas of Polonais ; Premier partie : idem, Trav. the Upper Devonian: U. S. Geol. Survey Geol. No. 1, Krakow, 56. pp. Bull. 41, 123 pp.
— , 1937a, Les megaspores du bassin houiller
Williamson, C, 1879, On the organization of : W. Polonais ; Deuxieme partie idem, Trav. the fossil plants of the coal measures, Pt. Geol. no. 3, Krakow, 78 pp.
IX : Royal Soc. London Philos. Trans., II, 319-364. 1878, vol. 169, Pt. pp. — , 1937b, Megasporen aus dem Westfal and
Stefan in Bohmen : idem. Bull, internal.,
,- , 1880, On the organization of the fossil ser. A, Annee, 1937, pp. 583-599. plants of the coal measures. Part X. In-
cluding an examination of the supposed — , 1938, Die Eignung von Megasporen als Radiolarians of the Carboniferous rocks Leitfossilien: Congres pour I'avancement Royal Soc. London Philos. Trans., 1880, vol. des etudes de stratigraphie Carbonifere,
171, Pt. II : pp. 493-540. Heerlen, 1935, Compte rendu, vol. 3, pp. 1711-1732. Wilson, L. R. and Brokaw, A. L., 1937, Plant
microf ossils of an Iowa coal deposit : Iowa 1940, Megasporen des Saarkarbons: Acad. Sci. Proc, vol. 44, pp. 127-130. Paleontographica, vol. 84, Abt. B, pp. 133- 150. and Coe, E. A., 1940, Descriptions of some unassigned plant microfossils from the Zeiller, R., 1906, Bassin Houiller et Permien de
Des Moines series of Iowa : Am. Midland Blanzy et du Creusot : Etudes des Gites
Naturalist, vol. 23, no. 1, pp. 182-186. Mineraux de la France, Fasc. ii, Flore Fos- sile, pp. 140-150. Wodehouse, R. P., 1935, Pollen grains. New York and London, 574 pp. Zetzche, F., Vicari, H. and Scharer, G., 1931, Untersuchungen iiber die Membran der Zerndt, Jan., 1930a, Megasporen aus dem Isa- Sporen und Pollen, IV. 3. Fossiles Sporo- bella-Floz (Schichten v. Laziska) in Trze- pollenin aus dem Tasmanit und der Mos-
binia : Ann. soc. geol. Pologne, Annee 1929, kauer Braunkohle : Helv. Chim. Acta, vol, 67-78. vol. 6 : pp. 302-313. 14, Pt. 1, pp. 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 Approximate Size -Range Granulati-sporites | poO/^ WITHIN "Spore" genera Reinschospora i
=1 Denso-sporites3Q'o
I mm. Triquitrites | 40Q^ =1 Prtyosporites 50,0^ Raistrickia I I 2 mm. 3 Reticulati- sporites 700y^
Punctati -spontes | | 1000/^ 3 mm. 1 + Cirratriradites Calamospora 4 mm. Fforinites J 6mnx Endosporiies 200(>^ 20.
Alisporites | 30^ =1 Alati- sporites 10 mm. Tasmanites 40'/^ Monoletes bOyu Parasporites 60/. 75^ Zonalo-sporitfes Triletes \00yu, 150^200/^300/^ Cystosporites
(fertile) (abortive) I | I I II ILLINOIS STATE GEOLOGICAL SURVEY REPORT OF INVESTIGATIONS NO. 91 Distributed in July 1944