Prepared in collaboration witk the Department of Geology, Coe College, and the Tennessee State Division of Geology

PRINTED BY AUTHORITY OF THE STATE OF ILLINOIS

URBANA, ILLINOIS 1944

STATE OF ILLINOIS DWIGHT H. GREEN, Governor DEPARTMENT OF REGISTRATION AND EDUCATION FRANK G. THOMPSON, Divector

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

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 ORGANIZATION

STATE OF ILLINOIS HON. DWIGHT H. GREEN, Govevnor DEPARTMENT OF REGISTRATION AND EDUCATION HON. FRANK G. THOMPSON, Disector

BOARD OF NATURAL RESOURCES AND CONSERVATION HON. FRANK G. THOMPSON, Chairman EDSON S. BASTIN, PH.D., DSc., 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 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 VELDAA. MILLARD,Junior Assistant to the Chief GEOLOGICAL RESOURCES GEOCHEMISTRY Coal FRANKH. REED, PH.D., Chief .Chemis{ G. H. CADY,PH.D., Senior Geologist and Head H. W. JACKMAN,M.S.E., Chemical Eng!neer L. C. MCCABE,PH.D., Geologist (on leave) R. J. HELFINSTINE,MS., ASJOC.Mech. Eng. JAMESG. MCCULLOUGH,Research Assoczate JAMESM. SCHOPF,PH.D., Asst. Geologist J. NORMANPAYNE, PH.D., Asst. Geologist Coal CHARLESC. BOLEY,M.S., Asst. Mining Eng. G. R YOHE PH D Chemist BRYANPARKS, M.S., Asst. Geologist CAROL BART~LS,B.S:: Research Assistant ROBERTM. KOSANKEM.A. Asst Geologist GEORGEM. WILSON,'B.S kesearih Assistant Industrial Minerals HENRYL. SMITH, A.B., dksearch Assistant J. S. MACHIN PH.D Chemist and Head Inda~t~ialMinerals DELBERTL. HANNA,';~.M.,Asst. Chemist J. E. LAMAR,B.S Geologist and Head H. B. WILLMAN')PH D. AJSOC Geologist Fl~orspar ROBERTM. GRO~AN'P; D A~SOC Geologist G. C. FINGER PH D. Assoc. Chemist ROBERTR. REYNOL&,~.s:: ~eseaich Asszstant EVERETTW. MAYNER'T, B.S., Asst. Chemist Oil arzd Gas X-ray and Spectrography A. H. BELL,PH.D., Geologist and Head W. F. BRADLEY,PH.D., Assoc. Chemist FREDERICKSQUIRES, B.S., Petr. Eng. CHARLESW CARTERPH D Asst Geologist Analytical WILLIAMH: EASTO~PH D' ~ssiGeologist 0. W REES PH D. Chemist and Head PAULG. LUCKHARDT: M.s., Asst. ~eolo~ist WAYNEF. MEENTS,Research Assistant HOWARD S. 'CLARK 'B A ASJOGChemist L. D. MCVICKERB S ' ~sst.~hkmist P W HENLINE'M s "AJS~ Chemical Engineer Areal and Engineering Geology ~ILL~AMF. WAGNER:' M.s.: Asst. Chemist GEORGEE. EKBLAWPH.D Geologist and Head K F BURSACKB A Research Assistant RICHARDF. FISHER,'MS., kut. Geologist CAME~ON D. L~WI; 'B.A. Research Assistant MARYANN WINSC~E B.S Research Assistant Subsurface Geolo~y MARJORIEWIN CHEST^, B:)S., Research AssJstant L E. WORKMAN-M S Geologist and Head A~NOLDC MAS~NB:.? Assoc. Geologist MINERAL ECONOMICS MERLYNB. BUHLE: M.s:, Asst. Geologist FRANKE. TIPPIE, B.S., Asst. Geologist W. H. VOSKUILPH.D Mineral Economist MARGARETSANDS B.A Research Asszstant GRACEN., OLIV~R,A.B:: Assistant in Mineral RUTHE. ROTH, B.s., ~esearchAssistant Economics WALTERR. SMITH,Research Assistant DOUGLASF. STEVENS,M.E., Research Associate Stratigraphy and Paleontology J. MARVINWELLER PH D Geologist and Head EDUCATIONAL EXTENSION CHALMERL. COOP&, MS::ASSOC. Geologist DONL. CARROLL,B.S., ASSOC.Geologist Petrography RALPHE. GRIM PH D Petrographer PUBLICATIONS AND RECORDS RICHARDSA. R~WL~ND,'PH.D., Asst. Petrographer (on leave) GEORGEE. EKBLAW,PH.D., Geologic Editor CHALMERL COOPERM S. Geologzc Edztor DOROTHYE ROSE B.s.' fechnical Editor Physics KATHRYNK: DED~AN,M.A., Asst. Technical Editor R J. PIER~~LPH D Physicist ALMAR. SWEENYA.B. Technzcal Files Clerk B.' J GREENGOOD'B'S. Mech Engineer PORTIAALLYN S~TH '~esearch Assistant DONALD 0. HOLL~VD,MS., ~i~t.Physicist MEREDITHM. CALKIN; Geologic Draftsman (on leave) LESLIED. VAUGHAN,Asst. Photographer

Special Staff to Aid in the War Effort

Oil and Gas Resources Ground Water Geology EARLEF. TAYLORM.S. Asst. Geologist CARLA. BAYS, PH.D., Spec. Geologist M. W. PULLEN,jR., M:s.,~ Spec. Asst. Geologist C. LELANDHORBERG, PH.D., Spec. Asst. Geologist ARNOLDEDDINGS, B.A., Research Assistant STEWARTFOLK, MS., Spec. Asst. Geologist VIRGINIAKREMERS, BS., Research Assistant ERNESTP. DUBOIS, PH.D., Spec. Asst., Geologist MARGARETPARKER, B.S., Research Assistant PAULHERBERT, JR., B.S., Asst. Geologzst JOHNA. HARRISON,B.S., Spec. Research Assistant CHARLESG. JOHNSON,A.B., Asst. Geologist (on leave)

Consultants: Ceramics, CULLENW. PARMELEE,M.S., D.Sc., and RALPHK. HURSH,B.S., University of Illinois Mechanical Engineering, SEICHIKONZO, 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

PAGE

Introduction ...... 7

Genera of fossil spores ...... 11

Tasmanites ...... 4-1

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 PI

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 then1 because there is no adequate basis for close comparison of The literature on fossil sposes is scat- biocharacters exhibited by these diversified tered, ancl in addition there has been much organs. diversity in methods of classification. The objects of the present article are to bring When affinities become better known this material together in summary form so that the difficulty due to uncomparable for convenient taxonomic reference and to features may be partially overcome, it evaluate the genera which have been pi-e- then may be possible to relate the forms viously proposed. After a study of these more naturally in classificatioii within micsofossils in America ancl a thorough suprageneric groups such as tribe, family, and subfaiiiily depending on how closely study of the literature it seems essential also to describe son~enew genera. It is in£ormation may be cosrelatecl. Indeed, liopecl tliat a funclamentally sound basis assignment to the same genus, or even has been provided for studies now in species, is theoretically possible and may progress in the thee laboratories repre- eventually be achieved in some few iti- sented by the authors and for future stances, but in general this iniplies a much st~lclies. more precise and detailed historical knowl- edge of interrelationships than is likely to Guiding principles which relate to the be fully established. treatment of this material are given below. For satisfactory nomenclature it is most 1. Only adherence to tlie systematic conveniat that those fossils which possess principles e~iibodied in tlie International numerous demonstrably comparable fea- Rules will give satisfactot-y results in the tures shall be placed undet- the same ge- sttldy of these micl-ofossils (as in other neric name. However, it is unsatisfactory fields of paleontologic study). Therefore, to classify fossils under tlie same generic the International Rules of Botanical name simply because a few arbitrary fea- Nomenclature, 3rd edition (Briquet), re- tures are held in co~~inion.The essential vised by the Intet-national Botanical Con- validity of any classification depends on gress of Cainl~riclge,1930, and published the relative significance tliat attaches to the in 1935,' have been used as a basis for various biocharacters. In fact it is by taxonoiilic treatment. virtue of such interpretive disctjmiiiation 2. Species classified within the same that taxononiy is to be distinguished from genus or under the same generic name a cataloging pi-oceclure. Interpretations must possess significant characteristics in nevertheless iiiust be based 011 evidence and common. There can be little positive proof not on supposition ; dificulties ai-e multi- of generic identity unless substantial plied whenever interpi-etation exceeds fac- honiologous coniparisons can be drawn be- tual bounds. tween respective biocharactei-s of each 3. In paleobotanical practice there has congeneric species and tlie type species of been much divergence with regard to the the genus. Foi- example niany isolated significance and proper interpi-etation of lycopsid megaspores must be classified unusually conlplete specimens in which separately f ron~the microspores f ouncl in parts usually found sepasatecl, and there- 1 Gustav Fischer, Jena, fore generally classified separately, are 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 inlpossible 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 sulsspecific magnitude (there is definition, may be better used in expressing no basis for assuming that geographic such relationships of generalizecl 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 non~encla- 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 nuni- contimed specific coordination of an ex- ber of "partial syriony~ms" among the tensive garniture of biocharacters over a named species of fossil plants to which period of time, without correspondingly these ren~arksapply. The word "synonym" detailed kiiowledge 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 cliff erent circunl- formation is most readily detern~inedfor 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 unclualified 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 Denlonstrated connections b e t w e e n showii to be lilutually 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 one valid name. However, instances arise in which it is difficult to determine which, of two ,or more continued use of technical nomenclature, names should actually be applied. In spite of thls, 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, niust 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 ?zo~zenantbigztum 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 comnlonly assigned to Spornngites. inorphological part of a plant in any stage Tasma~zitesnevertheless is a problematic of its life is here considered to be a form, spore-like in many of its 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 of how it niay be otherwise defined, is genera treated here belong with little doubt 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 nientioned in the structure representing any portion of its respective generic discussions. life cycle is available. Spores in many Knox ( 1939, 1941) has suggested that instalices 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 13ecause of their unique adaptation for (Sporogonites Halle may be an exception) dispersal and their n~lmericalab~~nclaiice in makes any suggested correlation hazard- many sedimentary deposits, it is expected ous at this time. that spores will eventually become 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 conirnonly noted by tlie geologist. evolutionary convergence as well as be- 7. Since the purpose of giving a name cause of community of derivation. Con- to a tayonoinic group is not to indicate tlie sequently unless there exists some cor- characters 01- the history of the group, roborative evidence based upon spore 1mt 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 nonienclature, for of different geologic age, recognizing that example, Gra~tulati-sporites, Laevigato- the similarities may or may not have phy- sporites, Denso-sjorites, instituted first by 1etic implications. Ibrahini (1933), is in 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- immes seems reasonably secure in several logic features of fossil forms. For one instances and their continuqnce is likely to thing the microspore in modern forms is cause less niisapprehension than attempts determined, in a strict sense, solely by the to institute more appropriate nonieiicla- 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 niay 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 "m~crospores" is to be lamented. Although true miscrospores frequently are and explain their consistency with sys- small by no means all small spores are microspores. Tke long kstablished botanical usage of the term "m~crospore tematic treatment of other types of plant 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 lmger than the actual megaspore. 10 PALEOZOIC FOSSIL SPORES the further clifficulty that no simple correct that deal iiiciclentally with Paleozoic morphological designation can be al~plied spores. Various lion-taxonomic systenis similarly to the dispersal fol-111s of the have heen used by sonie workers in desig- male gametophyte lsoclies in both modern nating types of spores in coal thin- and fossil for-ins. The dispersal stage sections, notably by Slates and his col- rather than tlie degree of gainetophytic leagues in the South Yorkshire lalsoratoi-y development is of greatest practical signifi- of the British Fuel Research Division, cance when fossils ai-e considered. The lsy Th. Lang in Silesia and, most impor- ternis pesispoi-e, exospore, ancl endospore tantly, by Reinhardt Tliiessen. These might seem equivalent to the pollen struc- apply very slightly to the present work tures perine, exine, and intine, i-espec- because species cannot he as reliably dis- tively. However, in tlie case of the bladcler- tinguished from spores in thin-sections, equipped fossil gpmnospermic poll en, th; significant bio~ogical characters are the bladder niemlssane which seems very gellei-ally iiioi-e difficult to ascertain, and similar in morphologic character to the File info;-ma1 non~enclatureused cloes not perisporal ineinbrane of certain crypto- req~k-esystematic consideration. Can- gamic forms, is regarded Iq inany as sequently articles of this nature are not equivalent to the exine of angiospermic considered except in connection with the pollen. ii~orphologyof certain f ornis. Revision of the iiiorphologic teriiii- 011 the other hand, these is a consider- iiology for fossil spores and siniilar niicro- able litei-attli-e on isolated spores in the fossils also seems required. Not only has Russian language which needs to be given there been a deartli of descriptive terms caref ul study before nonienclatoi-ial stabil- available for description of the vai-iecl ity can justifiably he hopecl for. Thus far structural features, but a good many deep- only a small portioii of this literature has ly rooted terms have become outmoded. been available to us, chiefly in abstract Several of these clate fro111 the period in form, and this is insufficient to provide which Selayi~zella was presented as a a satisfactory basis for understanding the primitive forei-unner of modern seed quite different nonienclatt~re Russian plants, and their honnologous iniplications writers have used. Nauinova ( 1937) now oftentimes seem uiiwari-anted. Non- mentions that about 400 species have been coinmital descriptive terms, which seem distinguished based on material ranging seasonably f see of theoretical connota- in age f roni the Lower Carboniferous to tions, have been preferred for use in the Tertiary. He mentions genera designated present paper. It is hopecl that it will be as Zonotl-iletcs ancl Axo~zot14etes.Nikitin possilsle later to treat the descriptive ( 1934) has distingtlished the genus Kry- niorpholog-y of spoi-e f orins specifically slztofoviclzin based on very singular ap- in greater detail. pearing, large spores of Devonian age. In the synoptic lists the species assignecl Luher ( 1938) imentions s~thgroups(gen- to each genus are given in alphabetical era ?) Aso~.zotrilctes,Zonot~riletes, Axo~zo- order. The orclei- of genera acloptecl is u~o?zoletes,Azo~zaletes and Zo~znletesand chronologic according to the time each distinguishes several apparently valid was first proposed. Althougli some inter- species. I11 a niore recent publication generic alliances can be recognized now, Lubes ( 1939) uses the generic -( ?) names and a more logical sequence should be Plngmlites, Tztwiella, Ci~~celliella,Sacc1~2'- adopted in later works, it is felt that ~~lnlin,Circelln, Libuwzelln, Sfii~zoselln,Sub- the chi-onologic order of priority will he sacculifcr and Plicntelln. It is to be hoped of some convenience £01- taxonoiiiic refer- that under iiioi-e favorable rii-cumstances ence. it will be possible to give full consideration Nearly all the refei-ences cited (11. 62) to the Russian stuclies which have already are those of particulas taxonomic or mo~-- been put to good use in the age determina- phologic pertinence for the fossils dis- tion and coi-zselation of the widely sep- cussecl. Although we believe this i-epre- anted coal deposits of the USSR and to sents a reasonably complete survey of this integrate the noiiieiiclature they are using literature, it cloes not include all articles TASMANITES GENERA OF FOSSIL SPORES

Genus TASRIANITESNewton, 1875 applied the name to spoi-es 01- "spore Spomzgites Dawson, 1863 (pars.), Canadian cases" which he thought were referable Naturalist, New Ser. vol. 8, no. 6, pp. 431-457. to Lepidodend~~on,Calaii~ites and similar Plate 1, figure 1 plants. Spornnyites papillnta, about one inch ( ! ) in diameter, and S. glnbrn about Ge~zeraZDiscussio~z.-There is a group the size of a niustard seed, were men- of highly distinctive fossil spore-like tioned ; later this description was repeated, bodies which occur ii~ostnotahly in the acconipanied by simll diagl-anis (Dawson, Devonian-AIississippiall black shale that 1865, p. 165, pl. 12, figs. 80-81). These requires a nanie amenable to standard fossils were associated with coal at Jog- taxoiionlic usages. At first, it was sup- gins, Nova Scotia, and n~ayin part have posed that the name Sporajzgites, if sup- been material that would be placed no^ ported by an emended definition, could with Triletcs, but there is no way to con- apply. We are now of the opinion that firm this from his description 01- illustra- the application of that name is not correct tions. His repeated clescriptions of S. in this connection, and that the various papillata as uone inch in diameter" leads descriptions that have appeared for Spo- one to suspect it may have been a seed. ?~a?.zgitesare so highly conflicting and I11 1871 (a, b) Dawson described an ambiguous or erroneous that that particu- additional species from the black shale at lar name should be consiclerecl a no~ize?z Kettle Point on Lake Huron as S. lzuron- n17u7+~.1iz.This is herewith proposed in emis ; "small globular papillate spore- view of the difficult taxonomic sittlation cases-probably of some Lepidode~zdroid as detailed below. plant.'' Later in the same year (Dawson, The present most common use of the 187lc) he also referred a specimen con- term "Spora~zgitcs" is by geologists who sisting of a small whorl of leaves or bracts, wish to record the presence of more or first described as Alz?zulnria acui~iijzatn,to less disc-shaped, resiiious-appeal-i11g bodies Spornngites. This last interpretation of that are very coii~monly found in the Spora?zyiites was accepted by Stopes Devoiiian-Mississi1313iaii black shale, many, ( 1914). Foi-n~ssimilar, in general, to but by no means all, of which are properly those from Kettle Point are usually con- referable to Tnsiitades as given below. strued as Sporangites by present day This con~monusage does not conforlm 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 tlie name. tempt to restrict it even if this coulcl be Dawson in 1883 presented a report on clone in accoi-dance 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 alxtract of which ap- consistency and practically all records of peared in the An~ericanAssociation for Spora~.zgitcsimlst n~erelybe regarded as tlie Advancement of Science proceedings indicative of some sndl rounded, brown- (1884a). The entire paper also appeared ish 01- yellowish bodies. Needless to say, in the Canadian Record of Science for the many ltinds of plant nlici-ofossils cor- same year (188%). The essential des- responcl to this characterization. How- criptive part and the figures ase identical ever, there is no reason why "sporang-ites" in both. Here Dawson proposed the iiaine may not he continuecl to be used in this Protostrlvi~zinincluding two species given sense, since it serves a descriptive pus- as "Spornngi f es (P~~otosal-da~zia)braxiliem- pose, in spite of the fact that it is botanical- sis n.s.," and "S. (P.) bilobatus 11s." ly misleading ; but tlie name should not Dawson stated that if we compare the be italicized or treated as a generic clesig-- separate macrospores of the Brazilian nation. sporocarps, and specially those which are Spom??gites was first proposed byt f ouilcl detached from their envelopes Damon in 1863 (p. 454). Here he (Sic!) with Spormzgites l~z~ro~ze~zsis,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" (thalli ?) of Pro- better accords with their wide distribution tosalui~ziawhich were exaniined. 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 P~~otosalvi~iia,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 melition the insidiousness of at- bodies with sporangite forms is by no tempting to apply scientific nome~iclature 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 Taswzanites), it is evident that "sporocarps~'; furthermore, no one else Protosalvi?zia and Sporangites cannot be has ever provided any conclusive evidence. in any way accurately treated as syn- Evidently many have taken this important onynls. No one yet has furnished tangible point for granted, however, 1)ecause 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 subsecl~tent the P~otosalvi~ziathalli, so it still is quite literature. speculative whether any relationship The "spherical and oval sacs, tlie walls (other than that of a conlnlon environ- of which are composed of a tissue of ment) is evideiiced by their infrequent hexagoilalcells . . tlireetosis~l~il- association in the Devonian-RiIississippian limeters in diameter," apl~areiitly are ldack shale. properly assignable to Proiostrlvi~~iuI>a \v- Dawson reviewed the subject in con- son, 1884. We are unable 10 ascertain nection with his paper in the bulletin of their characteristics reliably Tro~n anj7 the Chicago Academy of Science in 1886 figures Dawson published but White ant1 (pp. 105-118), calling attention to Orton's Stadnichenko ( 1923) give photographs ( 1552) discovery of S. 1zuro~zensis-like and data on a form, PTOtosahi~liiz T(~CIIII~ bodies in the Ohio shale, Clark's (1885) White, 1923, which is confi~-inatory of descriptioii of son~ewhatsiniilar material their thalloid characteristics, although in the Devonian of New York, the report White regarded them as sporangia. Wlitte of Johnsoii and Thomas (1884) on these (p. 239) has repeated the suggutioli, nlicroscopic discs in the Chicago water (which seems to have originated will1 supply and glacial till of the region, New- Dawsod, 1871a, b) that their affinity m7as ton's ( 1875) description of Tasmanites probably with "Devonian types antecctlent pu~zctatus,and his own (Dawson, 1884a, to the Carboniferous Lepiclodentlr;~ ancl b) descriptions of two lots of Brazilian Sigillariae." Possibly he thought the slmr- tmterial sent him by Hartt and by Orville angite bodies were actually spores oE 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" acunzinata. This article is posed relationship with Pro~osnlvi~~iabe- the best of any written by Dawson on tlie kame of close ~ssociation. Oval speci- subject. Nevertheless, in it, as well as mens of Proto~a~lviwiabear some resem- in his other papers, the inadequacies of his blance to lycopod sporangia, but there are concepts ar; fdly apparent. He appears general objections to such an interpre- confused as to the clistinction between tation. Even when they are occasioiially spores and sporangia ; he definitely con- in abundance no cones or actual lyco- siders the S. lzurone~zsisbodies referable podiaceous ren~ainsare noted in the same to the water ferns (hence, P~~otosalvinia), assemblage (Dawson reported some du- being misled on the one hand by the fal- TASMANITES 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- furcatunz 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 niost of the nomenclature rather best taken to be Foerstia olzioensis White, indiscriminately under the name Proto- instead of Dawson's species. Kidston salvi~zia. 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 Tnsma.izites are treated as synonyms ever proved by comparison of the holo- although their precedent publication dates typesAof 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 barticular 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 a11 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 spora~igiteforms 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, modern methods. Spores are present in definite haptotypic structures consisting it and are quite different from sporangite of trilete sutures with pyramic areas well 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 fuwatuvl.2. larly pitted and somewhat rugose accord- In the most thorough and essentially ing to information presented by Kidston definitive study of this plant by Kidston and Lang. Thus, these spores are en- and Laiig (192.5)) Dawson's name was tirely different from the sporangite bodies continued. However, White, in White that may be associated with them. 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 Sporoca~~ponWilliamson forms, in addition to Tasmalzites 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 Damson'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 Rhitocarps, etc." published in \he ~anadianRecord of stone age in Scotland. None of these Science 1s dated and ev~dently was published in 1884 can be ref erred to Tasmanites, although rather than 1885. Such minor errors are unfortunate in reference so widely used as the Fosstlium Catalogus. Lang's spore-type F (in part), which 14 PALEOZOIC FOSSIL SPORES seems to lack haptotypic features, may Newton's tasmaiiite which Reinsch ob- be generically similar to some of the tained from the British Museum. Reinsch spore-like rugulose-niembranous associ- assigned these forms, along with others, ates of Tas~ma~zitesin 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 are 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 "Spora~zgites,"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 LYilliam- 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 two other diverse 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 Taswzanites f ornis f I-om 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 gl-otlp onomic implication. description fits them best. Specific no- The systemic designation Taswmzites 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 Spomngites hu- manites specimens, is essentially not a ronensis (Dawson 1886, p. 116) because, taxonomic contribution. But since it as the "name Sforangites had priority, I establishes the similarity of American and (i.e., Dawson) do not think it 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- Tasma~zites 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- fully examined abundant sporangite speci- the material provided by Orton and mens from the Devonian-Mississippian Thomas there is a question as to proper black shale, but the similarity is by no specific distinction between them, but means evident if all the various inade- 6eing so widely separated geographically 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 The best evidence that at least many first-hand study can be made. No spe- of Dawson's S. huronemis specimens, cific definitions are attempted here, but it 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 from forms which itably illustrated work by P. F. Reinsch, unmistakably are spores of cryptogams vol. I1 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

Diaymsis.-An adequate description of thicker and the thinner sacs, or to find them in anything like the relation of an inner and outer Tnsmanites punctat~s,taken here as the coat. type species of the genus, is given by . . . . . Prof. Balfour, I believe, considers Newton and excerpts from his paper (pp. the Tasmanite discs to be closely allied to 339, 340, 341) are quoted below : Fle~ningites;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, MacMillanls Mag. March, 1874, in pl. 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 Tasfnanite, 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 inch in diameter for this substance, and that the spores (or with a still smaller dot in the centre, as shown rather the plant to which they belong) should in fig. 9. These structures are best seen in be called Tasmanites, with the specific title of the discs of White Coal. It may be thought fiztnctatus, 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 Flenzingites; 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 ?) Synzwzetry .-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, - ; they do not appear to extend quite through. Shape. Originally spherical except 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. 5). She.-Ranging from less than Neither Mr. Carruthers (Geol. Mag. 1865, 100 p. 432), nor Mr. MacNaughton (Trans. Roy. microns to 600 microns or slightly 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 niuch 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 1/504 of an inch. Mr. MacNaughton speaks of a Ornamentation.-Surface smootli 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- arated discs, in order to distinguish this outer more or less rugosity which may be in 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 more transparent than the rest; but I have described as essentially lacking in external failed to see any real difference between the ornamentation.

4 1/80 and 1/50 of an inch = 317.5 and 508 microns Haptotypic features.-Entirely absent. respectively. However, Newton's figure 2 "a large spore" given at 50x would be over 650 microns in diameter. False conclusions have been dl-awn either From his fi ure 5 (250x) the wall is about 25 microns from different forms in association with thick. In ehis specimen the wall is about 1/26 of the total diameter. Tas~nanites 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, of ten 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 Taswzanites 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 Afinity.-The question of the affinity (tetrasporic marking) can be demon- of Tasl~aanites,so far as it can be def- strated on material examined at first initely discussed now, hinges on whether hand nor is any structure of this sort these bodies can be taken to represent discernible in Thiessen's figures. Just as Newton remarked in his description of plant spores or not. Spores of all plants above the algal stage that are readily Taswzanites pu?zctatus, 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 ahat 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 surf ace, but actually con- structure and, in addition, have funda- sisting of two margins, sometimes thick- mentally a unicentric symn~etrywhich 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. Theref ore, it seems conclusive proximal wall of the spore. that Tas.~.zanitesis 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 Tas~mmites,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 Spora~gites manian oil shale by Singh (1932). He huronensis and with the Taswznnites 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 Tasmnites 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 with a different microf ossil 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 Tasnzudes is not to be wondered found many fornls with about the same at. The writers are convinced that, de- shape and size proportions as T.punctatus spite assocations of this sort, Tas'14.zanites 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 ma~itesdoes not represent any group of fossilization." This, of course, seems ordinary pteridophytic plants. quite improbable since the trilete suture Discussiout. - 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 Tasw.zanites, 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 microf ossils suggest Calamospora and it is interesting of similar age. Until more reports are to note that Plzyllotheca (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 Tasnzanites 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 comn~on forms concerns the nificance in tracing lines of phyletic rela- punctae. tionship among plants. The results that American Tas~majaitesfornis 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 Taswzanites, 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 the 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 (11, 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 Taswza?aites 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 Pavka 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. TASMANITESPUNCTATUS Newton, 1875, these microscopic fossil bodies represent Geol. Mag., ser. 2, vol. 2, no. 8, p. 341, pl. about as great a realm for speculation 10, figs. 2, 8. 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 represented as a valid excuse for their 4. TASMANITESSPP. Reinsch, nos. 12, (p. 3) ; 17, (pp. 3-4) ; 26, (p.. 5) ; 32, (p. 6) and continued neglect. By strict application 65 ( ?),, (p.9) ; 1884, Micro-Paleophytologia. of systematic principles we shall approach Reinschs 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 The following list includes names which probably referable to Tasnzanites. Although others he assigned to the Discieae also lack have been used and will have to be con- haptotypic spore features, they are probably sidered in subsequent treatment of species generically dissimilar. of Tasnzanites; forms identified as Spor- Note.-There is a possibility that the iso- angites jacksovti White (1905) and S. lated bodies associated with Protosalvinia bra- radiatus Duden (1897) have nothing to ziliensis Dawson and P. bilobata Dawson are referable to Tnsmanites, slid if so, they should do with Tasmanites and are omitted on receive separate taxonoinic consideration and this account. not be assigned to Protosalviutia. Proto- salvi~ziaclarkei Dawson, which was originally 1. TASMANITESCHICAGOENSIS (Reinsch) S. identified with Tasnzanites hwovtensis (Daw- W. and B., comb. 110v.~ son) by Clarke (1885) probably is entirely Discieae, Subtribus I, Subdividio I., 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, pl. 72A, fig. 76. "SporangitesJJ 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 Taswzanites 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 alaskc~z- otrsly cite Thomas as its author.' sis and Sporangites arctica by David White (1929) were studied only from thin sections. 2. TASMANITESHURONENSIS (Dawson) S. The former shows wall peculiarities which may W. and B., comb. nov. be comparable to that of Tasnzanites; the latter Sporangites hzwonensis Dawson, 1871, appears very similar to sectioned spores of Am. Jour. Sci., 3rd Ser., vol. 1, no. 4, p. higher plants. Whether the breaks in the walls 257, figs. 1-3. are accidental (as they would be in Tasnzanites) 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 Tasnzanites 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 ranging from 250 to 350 microns in diameter' Plate 1, figures 2, Za, 3, 3a showing sl~ghtexternal evidence of punctation but b; transmitted light at higher magnification exhibiting nu- Megaspores radially symmetrical ; prox- merous pores which penetrate the wall. Sometimes marked by characteristic rounded folds or split open by corn res imal side marked by triradiate sutures, sion. By transmitted light they appear amber colorex o; often with arcuate ridges connecting the sometimes have a reddish hue; in section the walls show faint 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 exhibit the triradiate marking seen in spores or variously ornamented. Ornamentation of lycopods. The interior is usually vacant but in some generally less well developed on proximal cases more or less filled with mineral matter (very fine pyritic crystals, calcite, etc.) . than on distal surfaces. Spores are rel- atively very large up to more than 3 rnm. j The initials S. W. and B. used here and elsewhere throughout this paper refer to the present authors, Schopf, Affinity with the f ree-sporing lycopsids. Wilson, and Bentall. Type species, by designation, Tdetes A copy of the Micro-Paleophytolo ia originally owned by Thomas ?ow in the John Crerar eibrary in Chicago, reinsclzi (Ibrahim) Scliopf (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 No, 18. .Thomas thus may have interpreted h~smaterlal a llttle differently than Reinsch did and may have mis- one in classification of fossil spores, par- understood the taxonomic principles involved. The pri- ticularly in the Carboniferous period mary purpose of author citatlon is to indicate type material upon whlch the description is based. when the f ree-sporing lycopsids were TRILETES 19 dominant. Evidence of tlie affinity of to him for exatnination. 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 proposecl 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 consequeiitly 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 winschi (Ibrahim) Schopf (1938), opinion. which also is the type species by designa- The syiiononlies are not absolutely com- tion for Triletes. (Type of Laevigati- sporitcs Ibrahim 1933). plete in the sense that references to all T~~i1ete.stzsbe~rostts (Ibrahim) S. W. and B., forms that have ever been described or comb. nov. (Type of Tl~bercz~lati-sporites illustrated are included because tdetean Ibrahim, 1933). forms are represented in too many pub- Triletes sextus S. Mi. and B., nornen nov. (As lished illustrations for them all to be Triletcs VI of Bennie and Kidston (1886), cited. The chief concern has been to this form was regarded as the type of Apiculati-sporites Ihrahim, 1933). give the references which include the type 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 Zo~zales-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. iiov. (Type of Valvisi-sporites Ibra- of reasons. Many have been taken from him, 1933). the stratigraphic distribution tables and By no means all the species that authors discussion published by Zerndt (1931, p. reg-arcled as congeneric with these types by 169; 1937a, p. 68; 1937b, p. 590, 593; virtue of arbitrary (artificial) distinctioiis 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 tinderstanding the rel- tlie 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 indiscriininately. These forins 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 forn~slisted here may extensive than inf ormation 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 then1 these questions as to actual recognized. Zerndt's data, however, are synonymy appear to be unsolvable. Wicher given for his spore "types" rather than ( 1934) has placed several of these names lor species. MO& 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, nations have not been rendered critically Washington Acad. Sci. Jour., vol. 22, p. 505, figs. 26-30. enough and this also tends to diminish Age, Upper Cretaceous. the value of stratigraphic ranges which can be given now. Nevertheless. some 6. TRILETES ARNOLDI (Miner) Harris, 1935, general information may be gained from Meddelelser om Gronland, vol. 112, p. 155. Selaginellites arlzoldi 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- erous (Schopf, chiefly unpublished data) 7. TRILETESAUGUSTAE (Loose) S. W. and B., comb. nov. which gives promise of future assistance Zonales-sporites augustne Loose, 1934, in long distance correlation. Inst. Palaobot. Arb. vol. 4, p. 150, pl. 7, Ninety-five species are included here. fig. 32. Many of them are referable to the sections Sectio, Auriculnti? Age, Westphalian B of Triletes already proposed ( Schopf, (Gasflamm) . 1938) ; others are less definitely assigned. 8. TRISETES,AURANTIUM Harris, 1935, Med- Certain relationships between species have delelser om Gronland, vol. 112, No. 1, pp. 164-165, fig. 52H-L, pl. 25, fig. 5, pl. 26, been noted when these seemed particu- 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. sake of completeness. Several new names and B., comb. nov, I-lpicztlati-sporites lapiculatus Ibrahim, have been proposed here, in addition to 1933, Sporenformen des Aegirhorizonts, p. the considerable number of new combi- 23, pl. 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. f935, &aA polonaise sci. Trav. Geol. no. 1, pp. 11-18. TRILETESAGNINA (Zerndt) S. W. and B., Sectio, Apka~zoxoiznti. Age, Lower Na- comb. nov. murian, Westphalian B, and younger ( ?). Lagenicula agnina Zerndt, 1937, Acad. 10. TRILETESARTECOLLATUS Nowak and Zerndt, polonaise sci. Trav. Geol. no. 3, p. 14, pls. 1936, Acad. polonaise sci. Bull. internat., 2 1-22. ser. A, p. 58, pl. 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. TRILETESAURTTUS Zerndt, 1930, idem, ser. TRILETESALES Harris, 1935, Meddelelser B, p. 46, pl. 1, figs. 4 and 5. om Gronland, vol. 112, no. 1, p. 163, fig. 53A-E, pl. 25, figs. 2, 8, 9, 11. Spore 0.7 mm. Zerndt, 1930, Soc. geol. Pologne Ann., vol. 6, pp. 307-305, 312, pl. Age, Liasso-Rhaetic. 1, figs. 3a, b, pl. 3, figs. 3a-d. Types 11 and 12 Zerndt, 1931, Acad. TRILETESANGULATA (Zerndt) S. W. and polonaise sci. Bull. internat., ser. A, p. 172, B., comb. nov. Triletes auritus Zerndt, 1937, idem, p. Lagenicula angulata Zerndt, 1937, 584, pl. 10, figs. 2, 6. Acad. polonaise sci. Trav. Geol. no. 3, pp. Triletes auritus Zerndt, 1940, Paleonto- 11-12, fig. 8, pls. 14, 15. graphica, vol. 84, abt. B, p. 146, pl. 9, figs. Type 7 Zerndt, 1931, Acad. polonaise 5-18. sci, Bull. internat., ser. A, p. 171, pl. 3, Sectio, nzm'culati (type). Age, West- fig. 8. phalian C, D. Sectio, Lagenicula. Age, Lower Na- TRILETESAURITUS var. GRANDIS Zerndt, 1937, murian (Grodziec series). Acad. polonaise Bull. internat., ser. A, p. 584, pl. 10, figs. 1, 3-5. TRILETES AREOLATUS Harris, 1935, Med- Type lla, Zerndt, 1937 (see above). delelser om Gronland, vol. 112, no. 1, pp. Tdetes auritzts var. grandis Zerndt, 158-159, fig. 51A-I?, pl. 26, figs. 3, 10. 1940, Paleontographica, voj. 84, abt. B, p. Age, Liasso-Rhaetic. 134, pl. 11, figs. 32-35. Age, Stephanian (Westphalian D ?). TRILETESARIADNAE (Miner) Harris, 1935, TRILETESAURITUS var. SECUNDUS (Maslan- idem. p. 155. kiewiczowa) S. W. and B., nom. nov. TRILETES 21

Triletes auritus I1 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 robertiaws and T. latihirsutus. TRII.ETES APPENDICULATUS Maslankiewic- zowa, 1932, idem., p. 163-164, fig. 39. 15. TRILETESBOREALIS (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 Selagi?zellites borealis Miner, 1932, les spores des houilles Francaises, p. 39- Washington Acad. Sci. Jour., vol. 22, p. 503, 40, pl. 2, fig. 5. figs. 12-21. Type 8? Sahabi, 1936, Recherches sur les spores des houilles Francaises, p. 41, 16. TRILETESBRASSERTI Stach and Zerndt, 1931, pl. 2, figs. 7-8. Gliickauf, Jahrg. 67, no. 35, p. 1123, figs. 29, 30, ( ?) 31. Sectio, Auricztlati. Age, Westphalian C, Dl Stephanian. Type 20 (Triletes brasserti Stach and Zerndt) , Zerndt, 1934, Acad. polonaise sci. Trav. Geol., No. 1, pp. 23-24, fig. 9, pl. 25, TRILETES BENNHOLDI Zerndt (non Bode), figs. 1-2. Age, Dinantian, Namurian 1937, Acad. polonaise sci. Trav. Geol. no. Westphalian A, B. 3, p. 5-6, fig. 3, pl. 2, figs. 1-6. Note.-Closely related to Triletes circzcmtex- Type 33 Zerndt, 1937 (idem). tus, T. saturnoides and T. superbus. Porosporites bennholdi (Bode) Zerndt, 1938, Deuxieme Cong. Stratig. Carbonif., 17. TRILETES BREVIACULEATUS Nowak and vol. 111. p. 1713. Age, Lower Namurian Zerndt, 1937, Zerndt, idem., no. 3, pl. 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 is a later homonym of Triletes. Such is hardly (Flora series). the case, however, because the identification of Note.-Evidently by mistake an illustration the genus Porostrobus and the species bennholdi of Tsiletes horridus accompanied Nowak and within it depend on other features in addition Zerndt's original description. The species was to the spores. Some day we possibly may know validated by photographs published a year later. actually how diagnostic isolated spores of Porostrobus may be, but at present such speci- 18. TRILETESBREVISPICULUS Schopf, 1938, Illi- mens certainly must be recorded separately nois Geol. Survey Rept. Inv. 50, pp. 26-27, pl. under a different genus (as Zerndt has done). 1, figs. 13a-r ; pl. 2, fig. 6 ; pl. 3, figs. 1-4. Consequently, also, these forms must be identi- Sectio, Afihnozonati. Age, Middle fied with reference to different nomenclatural Peni~sylvanian,Carbondale series. types. The specimen illustrated by Zerndt's photograph 4, plate 2, and which evidently 19. TRILETESCIRCUMTEXTUS Zerndt, 1934, Acad. served as the basis for his text figure 3 (p. 6), polonaise sci. Trav. Geol. no. 1, pp. 19-21, is regarded as the holotype of the species which fig. 7; pl. 19, figs. 1-12; pl. 21, figs. 1-7. is here credited to Zerndt. Type 18 Zerndt, 1931, Acad. polonaise On the other hand, no question arises as to Bull. internat., ser. A, p. 173, pl. 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 designation for the specimens is "spores of 20. TRILETESCLAVATOPILOSUS (Wicher) S. W. Porostrobus bennholdi." Zerndt, in using the and B., comb. nov. 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, tional cone characters warrants it, such a deter- pl. 8, figs. 25-26. 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, pl. 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, pl. 6, figs. 5-7. Age, Upper West- 14. TRILETESBIPILOSUS (Wicher) S. W. and B. phalian A-Lower Westphalian C. comb. nov. Sporitcs bipilosus Wicher, 1934, Inst. 21. TRILETESCRASSIACULEATUS (Zerndt) S. W. Palaobot. Arb., vol. 4, no. 4, p. 180, pl. and B., comb. nov. 8, figs. 13, 14. Lagenicula crassiaculeata Zerndt, 1937, PALEOZOIC FOSSIL SPORES

Acad. polonaise sci. Trav. Geol. no. 3, pp. Sectio, Aplzanozonati. Age, Westphalian 12-13. B, C, Lower Stephanian. Type 26A, Zerndt, 1937 (idem). Note.-Closely related to Triletes fzdge~zsand Sectio, Lageldcnla. Age, Upper Dinan- Triktes reinschi. tian ('i). 30. TRILETESGREENLANDICUS (Miner) Harris, Note.-Related to Triletes horridzbs. 1935. Meddelelser om Gronland. vol. 112, no. 1, p. 155. 22. TRILETESCYTTARTA Kenclall, 1942, Annals and Mag. Nat. History ser. 11, vol. 9, p. Selagi~zellitesg~,eenlatzdiczu Miner, 1932, 922, fig. 2. Washington Acad. Sci. Jour., vol. 22, p. 500, figs. 10, 11. Age, Lower Cretaceous. Age, Jurassic (Mid. Estuarine). 31. TRILETES GYMNOZONATUS Schopf, 1938, 23. TRILETE~DIFICILIS (Wicher) S. W. and Illinois Geol. Survey Rept. Inv. 50, pp. 37- B., comb. nov. 38, pl. 1, fig. 4, pl. 4, fig. 9. Sflorites dificilis Wicher, 1934, Inst. Sectio, Triangzdati. Age, Middle Penn- Palaobot. Arb., vol. 1, no. 4, p. 179, pl. 8, sylvanian, Carbondale series. figs. 17-18. Note.-Closely related to T7eilete.s arfecol- Type 14 (in part) Zerndt, 1931 Acad. latzu. polonaise sci. Bull. internat., ser. A, p. 172, pl. 4, figs. 11, 12, pl. 5, figs. 15, 16. 32. TRILETESHARRISI Murray, 1939, Geol. Mag., Zerndt, 1934, Acad. polonaise sci. Trav. vol. 76, p. 480, figs. 1, 2. Geol. no. 1, pp. 17-18, pl. 8, figs. 1-10, pl. 31, figs. 8, 9. 33. TRILETESHIRSUTUS (Loose) S. W. and B., Sectio, Afihanosomfi. Age, Lower comb. nov. Westphalian C. (Zerndt, Mid. Lower. Sporo~liteshirszrtzu, Loose, 1932, Neues Namurian to Westphalian D) . Jahrb., Beilage-Band 67, Abt. B, p. 452, pl. 20, fig. 58. 21. TRILETESDIFUSOPILOSUS (Wicher) S. W. and B., comb. nov. Setosi-sporites hi~~sutus (Loose) Ibra- him, 1933, Sporenf ormen des Aegirhori- Sporites difz~soflilosz~sWicher, 1934, zonts, p. 26. Inst. Palaobot. Arb., vol. 4, no. 4, pp. 182- 183, pl. 8, figs. 23, 21. Age, Lower West- Apiculati - sporites hirszctus (Loose) phalian C. Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 153. Note.-Closely related to Triletes praetextzu. Sborites hirsutus (Loose) Wicher. 1934, '1nst. Palaobot. Arb., vol. 4, no. 4, TRILETESECHINATUS (Miner) Harris, 1935, p. 181. Meddelelser om Gronland, vol. 112, no. 1, p. 155. Sectio, Lageniczda ?. Age, Westphalian Selagiwllites echinatz~s Miner, 1932, B, Lower Westphalian C. Washington Acad. Sci. Jour., vol. 22, p. 500, fig. 6. Age, Upper Cretaceous. 34. TRILETES HORRIDUS (Zerndt) S. W. and B., comb. nov. TRILETESERLANSONI (Miner) Harris, 1935, Lagetziczda horrida Zerndt, 1934, Acad. Meddelelser om Grotlland, vol. 112, no. 1, polonaise sci. Trav. Geol. no. 1, p. 25, fig. p. 155. 11, pl. 28, figs. 1-5.

Selnginellites erlla~zsoni Miner, 1932, Type 26 Zerndt, 1931, Acad. polonaise = Washington Acad. Sci. Jour., vol. 22, p. 500, sci. Bull. internat., ser. A, p. 175, pl, 9. fig. 28. figs. 1-3. Age, Upper Cretaceous. Sectio, Llageniczda. Age, Dinantian. TRILETESFLAVUS Stach and Zerndt, 1931, Note.-Closely related to Triletes c~?assiacu- Gliickauf, Jahrg. 67, no. 35, p. 1122, fig, 18. leatzts. Sectio, Azwicz~lati. Age, Westphalian C. 35. TRILETESINORNATUS (Miner) Harris, 1935, TRILETES FULGENS Zerndt, 1937, Acad. Meddelelser om Gronland, vol. 112, no. 1, polonaise sci. Trav. Geol. no. 3, p. 5, fig. 2, p. 155. pl. 1, figs. 1-9. Selagineltites i?zornatzcs Miner, 1932, Type S Zerndt, 1931, Acad. polonaise sci. Washington Acad. Sci. Jour., vol. 22, p. Bull. internat., ser. A, p. 171, pl. 3, figs. 505, figs. 7, 8. Age, Upper Cretaceous. 9-1* 0 Sectio, Aphalzozoltati. Age, Middle Din- 36. TRILETESIONTEIUS Harris, 1935, Meddelel- antiail to Middle Namurian. ser om Gronland, vol. 112, no. 1, pp. 166-167, Note.-Closely related to Triletes glab~ratz~s 169, figs. 52E, F, G, pl. 26, fig. 8. (pars.). 37. TRILETESKIDSTON1 (Loose) S. w. and B., 29. TRILETESGLABRATUS Zerndt. 1930. idem. comb. nov. ser. B, p. 45, pl. 1, figs. 2, 3. Sflo~~otziteskidstolzi Loose, 1932, n'eues Type 9 Zerndt, 1931, idem., ser. A, p. Jahrb, Beilage-Band 67, Abt. B, p. 452, 171. pl. 20, fig. 60. TRILETES 23

Laevigati-sporites kidstofli (Loose) TRILETESNIGROZONALIS Stach and Zerndt, Ibrahim, 1933, Sporenformen des Aegir- 1931, Gliickauf, Jahrg. 67, no. 35, p. 1123, horizonts, p. 19. figs. 26, 27. Reticztlati-sporitcs kidstorti (Loose) Spo&es ~zig~~ozo~~alis(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, pl. 8, figs. 11 and Sectio, Aphanoxona ti. Age, Upper 12. Westphalian B, Lower Westphallan C. Sectio, Auriculati. Age, Westphalian C. TRILETESLATIHIRSUTUS (Loose) S. W. and B., comb. nov. TRILETESNODOSUS (Wicher) S. W. and B., Apiculati-sporitcs lntilzirszttzu Loose, comb. nov. 1934, idem., p. 153. Sporites ~zodosusWicher, 1934, idem., Sectio, Lage&ztla. Age, Upper West- p. 177, pl. 8, fig. 21. ~halianB. Sectio, Aphanozorzati. Age, Lower Note.-~losel~related to Tyiletes ~~obertianus. Westphalian C.

39. TRILETES LAXIMARGINALIS Zerndt, 1940, TRILETES NUDUS (Nowak and Zerndt) Paleontographica, vol. 84, Abt. B, p. 136, Schopf, 1938, Illinois Geol. Survey Rept. pl. 10, figs. 24-28. Inv. 50, p. 30, pl. 5, fig. 7. Type 17a, Zerndt, 1940 (idem). Lngerzicula nztda Nowak and Zerndt, Sectio, Triangztlati. Age, Westphalian 1936, Acad. polonaise sci. Bull. internat., c, D (?I. ser. A, p. 60, pl. 1, fig. 6. Type 43 Zerndt, 1936 (idem). 40. TRILETESLEVIS (Zerndt) S. W. and B.. Sectio, Lagenicula. Age, Dinantian. comb. nov. Note.-The form recorded from Illinois as T. Lagrniczrla lcvis Zerndt, 1937, Acad. (?) nudzts probably is not referable to this polonaise sci. Bull. internat., ser. A, pp. species although the shape is rather similar. 587-588, pl. 15, figs. 1-11. 50. TRILETESOVALIS 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 Trilctes t~ra~zslztcetzs, lian C. T. tewimenzbranosa and to T. rzqoszts. Note.-Probably closely related to type 5 of Sahabi (1936). TRILETESLITCHI Harris, 1935, Meddelelser om Gronland, vol. 112, no. 1, pp. 159, 160, TRILETESPAPILLOSUS (Miner) Harris, 1935, fig. A-F, pl. 26, figs. 3, 10. Meddelelser om Gronland, vol. 112, No. 1, p. Age, Liasso-Rhaetic. 155. TRILETES MAMILLARIUS Bartlett, 1928, Selaginellites papilloszts Miner, 1932, Michigan Acad. Sci., vol. 9, p. 21, pls. 13-15. Washington Acad. Sci. Jour., vol. 22, p. 500, fig. 9. Age, Upper Cretaceous. Sectio, Ajha?zo~onnti. Age, possibly Upper Pottsville or Lower Allegheny. TRILETES PARVIAPICULATUS Zerndt, 1937, Acad. polonaise sci. Trav. Geol. no. 2, p. TRILETESMENSURA Harris, 1935, Meddelel- 17, pl. 24, figs. 1-4. ser om Groi~land,vol. 112, no. 1, pp. 160- 162, fig. 53J, pl. 26, figs. 2, 9. Sectio, Trialzgztlati?. Age, Lower Mar- ginal beds (Lower Namurian, vicinity of Age, Liasso-Rhaetic. Rybnik) .

TRILETESMIRABII-IS (Miner) S. W. and B., TRILETESPEDINACRON Harris, 1935, Med- comb. nov. delelser om Gronland, vol. 112, no. 1, pp. Selaginellites mirabilis Miner, 1935, 165-166, fig. 52S, T, pl. 27, fig. 6. Am. Midland Naturalist, vol. 16, p. 618, pl. Age, Liasso-Rhaetic. 23, figs. 1-6. Age, Upper Cretaceous?. TRILETESPERSIMILIS Harris, 1935, idem., p. TRILETESMUCRONATUS Nowak and Zerndt, 165, fig. 52R, pl. 25, fig. 4. 1936, Acad. polonaise sci. Bull. internat., ser. A, p. 59, pl. 1, figs. 1, 2. TRILETESPERTUBEROSUS (Loose) S. W. and Type 40 Zerndt, 1936 (idem). Age, B., comb. IIOV. Dinantian. Sporojzites pertz~beroszts Loose, 1932, Note.-Possibly closely related to Triletes Neues Jahrb., Beilage-Band 67, Abt. B, p. clavato~iloszu. 452, pl. 20, fig. 57. Tubercz~lati-sporites pertuberosus. 46. TRILETESMYRMECOIDES Harris, 1935, Med- (Loose) Loose, 1934, Inst. Palaobot. Arb., delelser om Gronland, vol. 112, no. 1, fig. vol. 4, no. 3, p. 147, pl. 20, fig. 57. 52M-Q, pl. 26, fig. 4. Sectio, Aphano,zonati. Age, Upper Age, Liasso-Rhaetic. Westphalian B. 24 PALEOZOIC FOSSIL SPORES

Note.-Equal to Type 14 Zerndt (1931, Type 10 Zerndt, 1931, idem., ser. A, p. 1934) in part. 172. Triletes I Kidston, Stach and Zerndt, 56. TRILETES PHYLLICUS Murray, 1939, Geol. 1931, Gliickauf, 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. 57. TRILETESPINGUIS Harris, 1935, Meddelel- 9 (Suppl.), p. 158. ser om Gronland, vol. 112, no. 1, p. 166, fig. Sporonites rehzschi Ibrahim, 1932, 52A-Dl pl. 25, fig. 3. Neues Jahrb., Beilage-Band 67, Abt. B, p. Age, Liasso-Rhaetic. 449, pl. 17, fig. 28. Laevigati-sporites reinschi (Ibrahim) 58. TRILETESPOLYSCELES Murray, 1939, Geol. Ibrahim, 1933, Sporenformen des Aegir- Mag., vol. 76, p. 484, figs. 5, 6. horizonts, p. 18, pl. 4, fig. 28. Age, Jurassic (Estuarine). Reticulati-spodes reinschi (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, 59. TRILETESPRAETEXTUS Zerndt, 1934, Acad. no. 3, p. 157. polonaise sci. Trav. Geol. no. 1, p. 24, pl. 26, figs. 1-6, pl. 27, figs. 1-6, Text-fig. 10. Sfiorites prirutz~s Wicher, 1931, idem. no. 4, p. 169. Type 21 Zerndt, 1931, Acad. polonaise sci. Bull. internat., ser. A, p. 174, pl. 8, figs. Types 1 to 6 incl., Rousseau, 1935, Mus. 24, 25. royale histoire nat. Belgique Bull. vol. 11, Age, Upper Dinantian and Lower Na- no. 21, pp. 3-4, pl. 1, figs. 1-5, pl. 2, fig. 6. murian. Type 1 Sahabi (pars.), 1936, spores des Note.-Closely related to Triletes difmo- houilles Francaises, p. 34, pl. 1, figs. 1, 2. pilosus. Sectio, Aphanozonati. Age, Westphalian C & D, Lower Stephanian. TRILETXS POTOSIENSIS Zerndt, 1938, De- Note.-Type species of Triletes by designa- uxieme Cong. Stratig. Carbonifere, vol. 3, tion (Schopf, 1938). p. 1728, pl. 155, fig. 4. Sectio, Auriculati?. Age, Carbonif er- 64. TRILETESRETICULATUS Zerndt, 1938, De- OUS. uxieme Cong. Stratig. Carbonif., vol. 3, p. 1728, pl. 155, fig. 5. TRILETESRADIATUS (Ibrahim) S. W. and B., Age, Mesozoic. comb. nov. @ Spororcites radiatlts Ibrahim, 1932, 65. TRILETESREXARGENTEUS Harris, 1935, Med- Neues Jahrb, Beilage-Band 67, Abt. B, p. delelser om Grijnland, vol. 112, no. 1, pp. 449, pl. 16, fig. 25. 156-158, fig. 51G-J, pl. 26, fig. 13. Zonales-sporites radiatus (Ibrahim) Age, Liasso-Rhaetic. Ibrahim, 1933, Sporenf ormen des Aegir- horizonts, p. 29, pl. 3, fig. 25. 66. TRILETESRICHARDSONI 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).

TRILETESRADIOSUS S. W. and B. nom. nov. 67. TRILETESROBERTIANUS S. W. and B., nom. Triletes radicttus Zerndt, 1937, Acad. nov. polonaise sci. Trav. Geol. no. 3, p. 10, fig. Tdetes kidstoni Zerndt, 1934, Acad. 7, pl. 13, figs. 1-5. polonaise sci. Trav. Geol. no. 1, pp. 26-27, Types 22 and 23 Zerndt, 1931, Acad. fig. 12, pl. 28, figs. 6-11, pl. 29, figs. 1-13. polonaise sci. Bull. internat., ser. A, p. 174, Type 27 Zerndt, 1931, Acad. polonaise pl. 8, figs. 24, 25. sci. Bull. internat., ser. A, p. 175, pl. 9, Age, Lower Namurian (Upper Di- figs. 29-32. nant~an? ) . Sectio, Lagenicula. 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. latihirszdtus. The new name is neces- Tviletes (see sp. 61 above), thus necessitating sitated by preoccupation of the specific epithet a new name as here proposed. (species no. 37). This species is likewise named 63. TRILETESREINSCHI (Ibrahim) Schopf, 1938, for Robert Kidston. Illinois Geol. Survey Rept. Inv. 50, p. 24, 68. TRILETESROTATUS Bartlett, 1928, Michigan p1. 2, figs. 2-4, pl. 5, figs. 8, 9. Acad. sci. Papers, vol. 9, p. 21, pls. 9, 10, Triletes I Bennie and Kidston, 1886, 11, 12. Royal Phys. Soc. Edinburgh Proc., vol. 9, p. 107, pl. 111, figs. la, b. Type 19, Triletes rotatus Bartlett, Spore 1.9 mm. Zerndt, 1930, Soc. Geol. Zerndt, 1934, Acad. polonaise sci. Trav. Pologne, Ann. vol 6, p. 308, 312, pl. 1, figs. Geol. no. 1, pp. 21-22, fig. 8, pl. 24, figs. 1-6. 5a, b, pl. 111, figs. 5a, b. Triletes rotatus Bartlett, Zerndt, 1937, Triletes Type 1 Kidston, Zerndt, 1930, idem., no. 3, pp. 8-10, pls. 6-10. Acad. polonaise xi. Bull. internat., ser. B, Age, type material probably upper p. 43, pl. 1, fig. 1. Pottsville. TRILETES 25

TRILETESROTATUS var. DENTICULATA Zerndt, 74. TRILETESSILVANUS (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-Bamd 67, Abt. B, p. above ( ?). 448, pl. 15, fig. 22. Laevigati - sporites silvanzis Ibrahim, 69. TRILETESRUGOSUS (Loose) Schopf, 1938, 1933, Sporenformen des Aegirhorizonts, p. Illinois Geol. Surv. Rept. Inv. 50, pp. 29- 20, pl. 2, fig. 22, pl. 6, fig. 47. 30, pl. 5, fig. 6. Sporites sil-danus (Ibrahim) Wicher, Sporonites rugosus Loose, 1932, Neues 1934, Inst. Palaobot. Arb., vol. 4, No. 4, p. Jahrb., Beilage-Band 67, Abt. B, p. 452, 174, pl. 8, figs. 5-8. pl. 20, fig. 59. Sectio, Auriculati. Age, Upper West- Lezigati-sporites rugosus (Loose) Ibra- phalian B, Lower Westphalian C. him, 1933, Sporenf ormen des Aegirhori- Note.-Closely related to Triletes azwitus. zonts, pp. 18-19, pl. 7, fig. 65. Punctati - sporites rugosus (Loose) 75. TRILETESSIMPLEX (Zerndt) S. W. and B., Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. comb. nov. 3, p. 146. Lagenicula simplex Zerndt, 1937, Acad. Sectio, Lage~zicula. Age, Upper West- polonaise sci. Trav. Geol. no. 3, pp. 14-15, phalian B and above ( ?). fig. 10, pl. 23, figs. 1-5. Note.-May be closely allied with Triletes Type 35 Zerndt, 1937, idem. tra~zslucem. Sectio, Aphanoxonati ( ?). Age, Lower Namurian. 70. TRILETESSAARENSIS Zerndt, 1941, Paleonto- TRILETESSIMPLEX var. LEVIS Zerndt, 1937, graphics, Abt. B, vol. 84, p. 134, pl. 12, figs. 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- Tdetes aztritus I11 Maslankiewiczowa, phalian C and Lower Westphalian D. 1932, Acta Soc. Botanicarum Poloniae, vol. 9 (Suppl.) pp. 161-163, 172-173, figs. 37, 38. 71. TRILETESSATURNOIDES (Ibrahim) S. W. Sectio, Au~iculati. Age, Upper West- and B., comb. nov. phalian B (Laziska series). Sporo.lzif,~ssaturnoides Ibrahim, 1932, Note.-The group is clearly distinguished Neues Jahrb., Beilage-Band 67, Abt. B, p. specifically by coarse reticulation, in this respect 449, pl. 16, fig. 26. somewhat similar to Triletes tuberculatus, Zonales-spo&es saturnoides (Ibrahim) though in shape it resembles Triletes auritzts. Ibrahim, 1933, Sporenformen des Aegir- Possibly it represents a connecting link between horizonts, p. 27-28, pl. 3, fig. 26. these species. It is named for Zofja Kowalew- ska Maslankiewiczowa who provided an ample Age, Upper Westphalian B. description of the form. Note.-Allied with Triletes bmsserti, T. superbzbs and T. circz~mtextus. 77. TRILETES SPARASSUS Murray, 1939, Geol. Mag., vol. 76, p. 450, p. 482, figs. 3, 4. 72. TRILETESSEXTUS S. W. and B., nom. nov. Age, Jurassic (Estuarine). Triletes VI Bennie and Kidston, 1886, Royal Physical Soc. Edinburgh Proc., vol. 78: TRILETESSPLENDIDA (Zerndt) S. W. and B., 9, p. 109, pl. 3, figs. 6a, 6b, 6c. comb. nov. Sectio, Aphanozonati. 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, pls. 18-20. Zerndt (1931, 1934) in Type 14. Inasmuch as Type 28 Zerndt, 1931, Acad. polonaise this was used as the type for Apiczdati-sflorites sci. Bull. internat., Ser. A, p. 175, pl. 10, Ibrahim, it seems desirable to have an orthodox figs. 36, 37. designation for reference. Sectio, Aphawozonati. Age, Upper Di- nantian, Lower Namurian. 73. TRILETESSI~XTUSDECIMUS S. W. and B. nom. Note.-The lageniculate vestibule possessed nov. by this form is insufficient basis for assigning Triletes XVI Bennie and Kidston, 1886, it to Lage&zda in disregard of its other charac- idem. pp. 113-114, pl. 5, figs. 16a, 16b, 16c, teristic Apha~zozo~zatifeatures. 16d, 16e. Age, Bernician and Lanarkian = ap- 79. TRILETESSTENoxYsivrAToms Harris, 1935, prox. Upper Dinantian and Namurian. Meddelelser om Gronland, vol. 112, no. 1, - Note.-Related to Triletes superbus, and prob. p. 167, pl. 25, fig. 6. to T. circzwztextz~sand T. brasserti. This form Age, Liasso-Rhaetic. was taken as the type of Zonales-sporites Ibra- him, hence it seems desirable to propose an 80. TRILETESSUBFUSCUS (Wicher) S. W. and orthodox name for purposes of reference. B., comb. nov. PALEOZOIC FOSSIL SPORES

Sporites subfusczts Wicher, 1934, Inst. TRILETESTENUISPINOSUS var. SECUNDUS S. Palaobot. Arb., vol. 4, no. 4, p. 179, pl. 8, W. and B., var. nom. nov. fig. 20. Triletes tenztispinosus var. I1 Zerndt, Type 14 Zerndt (in part), 1934, Acad. 1937, idem., no. 3, pp. 7-8, fig. 5, pl. 3, figs. polonaise sci. Trav. Geol. no. 1, pp. 17, 18, 3 and 4. Age, Dinantian - Upper and fig. 5, pls. 8-17. Lower Namurian. Sectio, Aphanozonati. Age, Lower No te.-Zerndt describes three forms, one which evidently is typically the species and two Westphalian C. varieties, only one of which is properly named. 81. TRILETES SUBROTUNDUS (Miner) Harris, The new varietal name proposed provides a 1935, Meddelelser om Gronland, vol. 112, means of reference. no. 1, p. 155, p. 165. 88. TRILETES TRANSLUCENS Schopf, 1938, Illi- Selaginellites subrotundus Miner, 1932, nois Geol. Survey Rept. Inv. 50, pp. 28-29, Washington Acad. Sci. Jour., vol. 22, p. 505, pl. 5, figs. 3-5. figs. 4, 5. Sectio, Lagewicula. Age, Middle Penn- Age, Upper Cretaceous. sylvanian, upper Carbondale, lower Mc- Leansboro series. 82. TRILETES SUBTILINODULATA ( Nowak and Note.-Closely related to Triletes tenuimem- Zerndt) S. W, and B., comb. nov. branosa, T. levis, and ( ?) T. rugosus. Lagenicula sztbtilinodulata, Nowak and Zerndt, 1936, Acad. polonaise sci. Bull. in- 89. TRILETESTRIAKGULATUS Zerndt, 1930, Acad. ternat., ser. A, pp. 59-60, pl. 1, fig. 7. polonaise sci. Bull. internat., ser. B, p. 51, Type 42 Zerndt, Nowak and Zerndt, pl. 7, figs. 19-24. 1936 (idem). Spore 0.5 mm. Zerndt, 1930, Soc. Geol. Sectio, Lagenicula. Age, Dinantian. Pologne Ann., vol. 6, pp. 306, 312, pl. 1, figs. la, b, c, pl. 2, figs. la, b. 83. TRILETESSUBPILOSUS (Ibrahim) S. W. and Type 17 Zerndt, 1931 Acad. polonaise B., comb. nov. sci. Bull. internat. ser. A, p. 173. Setosi - sporites subpilosus Ibrahim, Sboronites reaalis Ibrahim. 1932. Neues 1933, Sporenformen des Aegirhorizonts, p. ~ahrb.,~eilage-&nd 67, Abt. 'B, p. '449, pl. 27, pl. 5, fig. 40. 16, fig. 24. Sectio, Lagenicula. Age, Aegir coal, Sporites triangzllatus "var. regalis" boundary of Westphalian B and C. (Zerndt) Ibrahim, 1933, Sporenformen des Note.-May be closely related to Triletes Aegirhorizonts, p. 29, pl. 3, fig. 24. YO ber tianus. Sfiorites regalis (Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 149, 84. TRILETESSUPERBUS Bartlett, 1928, Michigan pl. 7, fig. 34. Acad. Sci. Papers, vol. 9, pp. 20, 21, pl. 7, Sporites friangztlatus (Zerndt) Wicher, figs. 1, 2, pl. 8, figs. 1, 2. 1934, idem., vol. 4, no. 4, p. 175. Age, Probably upper Pottsville. Triletes triangztlatus Zerndt, 1934, Note.-Probably allied with Triletes circum- Acad. polonaise sci. Trav. Geol. no. 1, p. textus and T. brasserti. 19, fig. 6, pl. 18, figs. 1-24. Triletes triangulatus Zerndt, Schopf, 85. TRILETESTENUICOLLATUS Nowak and Zerndt, 1936, Illinois Acad. Sci., Trans. vol. 28, p. 1936, Acad. polonaise sci. Bull. internat., 107, fig. 6. ser. A, p. 59, pl. 1, fig. 3. Type X Sahabi, 1936, spores des hod- Type 41 Zerndt, Nowak and Zerndt, les Francaises, p. 43, fig. 11, pl. 4, figs 6-11. 1936 (idem). Triletes triangulatus Zerndt, 1937, Age, Dinantian. Acad. polonaise sci. Trav. Geol. no. 3, pl. 5, 86. TRILETESTEKUIMEMBRANOSA (Zerndt) S. figs. 1-5. W. and B., comb. nov. Triletes triangulatus Zerndt, Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, Lagenicula tenuinzembranosa Zerndt, pp. 32-37, pl. 1, figs. 7, 8, pl. 4, figs. 1-7, pl. 1937, idem. p. 587, pl. 14, figs. 1, 2. 7, figs. 5, 6. Sectio, Lage~zicula. Age, Westphalian Triletes triangulatus Zerndt, 1940, C and Lower Westphalian D. Paleontographica, vol. 84, Abt. B, pl. 10, Type 25 Zerndt, 1931, idem., p. 175, pl. figs. 19-23. 9, figs. 33 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 transluceizs. ~riletestria&&latus I1 Zerndt, 1930, Acad. polonaise sci. Bull. internat., ser. B, 87. TRILETES TENUISPINOSUS Zerndt, 1934, p. 53, pl. 7, figs. 25-30. Acad. polonaise sci. Trav. Geol. no. 1, p. Sporonites zonatus Ibrahim, 1932, 16, text-fig. 4, pl. 7, figs. 1-15. Neues Jahrb., Beilage-Band 67, Abt. B, p. Type 13 Zerndt, 1934 (idem). 448, pl. 16, fig. 23. TRILETES TENUISPINOSUS var. BREVISPINOSA Zonales-sporites triangulatus secundus Zerndt, 1937, idem., no. 3, p. 6, fig. 4, pl. 3, Ibrahim, 1933, Sporenformen des Aegirhor- figs. 2-7. izonts, p. 30, pl. 3, fig. 23, pl. 7, fig. 64. PITYOSPORITES 27

Zonales-sporites xonatz~s ( Ibrahim) 94. TRILETESTYLOTUS Harris, 1935, Meddelelser Loose, 1934, Inst. Palaobot. Arb., vol. 4, om Gronland, vol. 112, no. 1, pp. 162-163, no. 3, p. 150, pl. 7, fig. 31. fig. 53F-I, pl. 26, figs. 1, 12. Sflorites triangulatus var. zonatzhs (Ib- Age, Liasso-Rhaetic. rahim) Wicher, 1934, Inst. Palaobot. Arb., 95. TRIL~ESVALENS (Wicher) S. W. and B., vol. 4, no. 4, pp. 175-176. comb. nov. TRILETESTRIANGULATUS var. ? Sporites valens Wicher, 1934, Inst. Triletes triangz~latus 111, Stach and Palaobot. Arb., vol. 4, no. 4, p. 178, pl. 8, Zerndt, 1931, Gliickauf, Jahrg. 67, no 35, fig. 19. p. 1123, figs. 32, 33. Sectio, Aflhanoxo?zati. Age, Lower Sectio, Trialzgsdati. Age, upper Lower Westphalian C. Kamurian through Westphalian D. TRILETESnomen excludende 1. Apiculati-sporites megaspinosus Ibrahim, 1933, Sporen- TRILETESTRICOLLINUS Zerndt, 1938, Deux- formen des Aegirhorizonts, etc., p. 24, pl. 8, fig. 69. ieme Cong. Stratig. Carbonif. vol. 3, p. 1713, Note: This form is not congeneric with the type of pl. 155, fig. 3. Apiculati-sporites (Triletes sextgs) and, although it ap- arently is sufficiently characterized and characteristic to Type 44 Zerndt, 1938 (idem). %e a useful microfossil, no further generic assignment will Sectio, Trialzgulati. Age, Westphalian be attempted. B, C, and D. Genus PITYOSPORITESSeward, 1911 TRILETESTRILOBUS (Ibrahim) S. W. and B., comb. nov. Plate 2, figures 15-15b Sporonites t~~ilobzuIbrahim, 1932, Diagnoses given by Seward are as Neues Jahrb., Beilage-Band 67, Abt. B, p. follows : 449, pl. 17, fig. 30. The generic designation Pityosporites is pro- Yalvisi-sporites trilobz~s (Ibrahim) Ib- posed for winged spores agreeing in form and rahim, 1933, Sporenformen des Aegirhor- size with recent Abietineous genera (1914, p. izonts, p. 33, pl. 4, fig. 30. 23). Valvisi-sporites trilobz~s (Ibrahim) Ib- . . . . Spores provided with bladder-like rahim, Loose, 1934, Inst. Palaobot. Arb., extensions of the exine, agreeing in size and vol. 4, no. 3, p. 152, pl. 17, fig. 30. form with those of recent Abietineous genera Sectio, Auriczdati. Age, Upper West- (1919, p. 398). phalian B. A more adequate definition inferred N&e.-~ossibl~ very closely related to Tri- letes appendiculatz~s. Wicher (1934) regards from the above, and from consideration it as similar to T. auritz~s. of the species included here, is as follows : S3~gq'c1qaetry.-Pollen grains appearing 92. TRILETESTUBERCULATUS Zerndt, 1930, Acad. bilateral, due to the presence of bladders polonaise sci. Bull. internat. ser. B, pp. 47-51, pl. 2, figs. 6, 7, pl. 3, figs. 8, 9, pl. 4, figs. placed on opposite sides. The funda- 10, 11, pl. 5, figs. 12, 13. mental symmetry, shown by the body but Triletes XIX Kidston, 1890, Royal SOC. modified, is radial, as this pollen arises Edinburgh Trans., vol. 36, pl., figs. 9-11. from tetrahedral tetrads. Type 16 Zerndt, 1931, Acad. polonaise sci. Bull. internat., ser. A, p. 173. Shape-Grains elliptical as viewed Tdetes tuberculatz~s Zerndt, Maslan- from proximal side, the diameter of the kiewiczowa, 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, Auriczdati. Age, Westphalian body between them. Body appears ap- B, C, Stephanian? proximately circular in proxilnal view. 93. TRILETESTUBEROSUS (Ibrahim) S. W. and Viewed from either side the bulbose blad- B., comb. nov. ders are inclined distally and a broad Spo~~onitestuberosus Ibrahim, 1932, sulcus appears between then1 on the distal Neues Jahrb., Beilage-Band 67, Abt. B, p. side. In lateral outline the body appears 449, pl. 16, fig. 27. Tubercz~lati-sporites tuberosus (Ibra- broadly oval with the smaller pole toward him) Ibrahim, 1933, Sporenformen des the distal sulcus. Aegirhorizonts, p. 23, pl. 3, fig. 27. Size.-Various species range in total Tube?-cz~lati-sporites tuberosus (Ibra- length (bladder tip to bladder tip) from him) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 147, pl. 7, fig. 37. as sniall as 40 to over 100 microns. Sflorites tuberosus (Ibrahim) Wicher, Orjza?z.zejztation.-External bladder sur- 1934, idem, vol. 4, no. 4, pp. 177-178, pl. 8, face minutely gi-an~~loseor rugose ; in- fig. 22. ternally the bladders are usually at least Sectio, Aplza~zozo~zati. Age, Upper Westphalian B, Lower Westphalian C. weakly reticulate. Proximal cap may be Note.-Probably the same as some of Zerndt's nearly smootli, rugose or striate. Distal Type 14. body wall least ornamented. 28 PALEOZOIC FOSSIL SPORES

Hapto typic f ea tares.-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 definiti~n.~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 ccoat.-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 nicroscope ; the distal wall is much thin- somewhat smaller groups that will more 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 flrio~i,but the desira- ,4.finity.-There can be little question bility of recognizing a type species for that grains of this sort are referable to this group which, as we construe it, does the Coniferae. Members of modern signify a definite natural plant alliance, Podocarpaceae and Pinaceae show essen- cannot be denied. Part of the taxo- tially similar features. Correlatives of noniist's responsibility is to place the sys- species of Pityospo~ites represented by tematic data in order so that later students vegetative organs are hardly recognized in may revise it to greater perfection in the Paleozoic formations. The presence of light of additional data. Such progress such pollen is evidence that such plants is very difficult if nomenclatural types are existed, possibly as a xerophytic upland not definitely indicated and fully utilized flora, and tend to substantiate the view in the application of technical nomeiicla- long held that an important ancestry, very tt~re. 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 f orins found better represented in younger that is being studied rarely permits cate- strata. gorical statement and new discoveries call Pityosflorites represents a group of for greater revision in preexisting con- Coniferae whose pollen is essentially mod- cepts, perhaps, than in any other plant ern in aspect. Florin's brilliant researches science. ( 1938- 1940) have sufficiently denion- Pityos#orites can hardly be considered strated that Lebaclzia, Emestiodendrou1., as synonymous with any -modern genus ; and Walclzianthus for the most part do perhaps the niost important reason is that not possess pollen of tlie Pityospo&es 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 F1ori1.zite.sn. gen. described group, identifiiation with tlie genus Pityo- below (p. 56). sporites becomes inadequate as an ex- Remarks.-It will be evident that the pression of affinity. Quite evidently niany 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 7 Although Seward (1919, pp 398-9) has included one of the forms Nathorst re orted from the Hor clay as bladders to this group. It is evident from wPityospovites sp." (whicR we should prefer to assign Seward's descriptions, however (1914, to Alisporiter), Seward apparently did this on the as- sumption that Nathorst's specimen was slmllar to pollen 1919, 1933), that the emphasis is not of Picea excelsa. Wodehouse ( 1.935 ) h,as shown Picea so much merely on the presence of two pollen to be very different. The Pztyospovzfes s . of Solms from Franz Josef Land (Seward, 1919, p. 3995 no doubt bladders as it is 011 the essentially modern is correctly assigned. PUNCTATI-SPORITES

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.

PITYOSPORITESANTARCTICUS Seward, 1914, Sy~mzetry.-Spores radial, trilete. 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, pl. 8, fig. 45. Pityosporites antarcticus Seward, 1919, slight shortening of the axial dimension ; 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. Sixe.-Spores of various species range ege, ('Not older than Lower Meso- zoic. from 45 to 85 microns in mean diameter. Ornalwntatiouz. - Various ; surfaces PITYOSPORITESCHINLEANA Daugherty, 1941, levigate to punctate, rugose, reticulate or Carnegie Inst. Washington, Pub. 526, p. 398, pl. 34, fig. 5. mildly apiculate. Age, Upper Triassic. Haptotypic structures. - Of moderate prominence, relative length of trilete rays PITYOSPORITES( ?) JEFFREYI Florin, 1940, Paleontographica vol. 85, Abt. B, no. 5, p. highly variable between different species ; 327, pl. 163-4, figs. 8-12. no equatorial 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 21A, no. 13, p. 6. (Systematic status of very prominent nor highly ornamented. this nomenclatural combmation 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.-Bladders are seemingly set nearly op- posite though it is possible their distal inclina- Afinity .-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 f son1 be this form belongs to Alisporitcs rather than in the present genus. sonie fossils assigned to the fei-ns. The spores Kidston (1906) obtained from va- 4. PITYOSPORITESPALLIDUS Reissinger (nomen nudum), 1939, Paleontographica, vol. 84, rious Crossotheca fructifications are char- Abt. B, p. 14. acteristic of typical species of Punctn.ti- Age, Lowest Jura (Lias). sporites. However, n~anyof the species Note.-An illustration required to validate represented by isolated spores liave no the name has not yet been published. known pteridospermic relationship al- 5. PITYOSPORITESSEWARDI Virkki, 1937, In- though there is no adequate basis for dian Acad. Sci. Proc., vol. 6, no. 6, sect. B, separating them from fornis which are pp. 428-431, figs. 2a, b, c, d, pl. 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 boulder bed. which is a rather large one, and it may he expected that some diverse elements 6. PITYOSPORITESSTEPHANENSIS Florin, 1940, liave been included which will be segre- Paleontographica, vol. 85, Abt. B, no. 5. pp. 327-8, pl. 163-4, figs. 13-16. gated on the basis of newer infornlation. Age, Middle Stephanian. It is quite possible that some of the forms assigned to Punctafi-sporites are 7. PITYOSPORITESsp. Florin, 1940, idem, p. 61, merely immature and lack their distinc- d. 163-4 fig. 17, fig. 18? Age, Lowe; ~lothliegendes. tive mature ornamentation. Such may be the case for immature forms that in 8. PITYOSPORITES( ?) sp. Florin, 1940, idem. reality belong to Raistrickia; n. gen. de- p. 62, pls. 165-6, fig. 20. Age, Upper Carboniferous. scribed below (p. 55). Immature forms Note.-No inclination of bladders shown. should not be-- arbitrarily assigned to Punctati-sporites but it is well to recog- 9. PITYOSPORITESsp. Daugherty, 1941, Car- negie Inst. Wash., Pub. 526, description of nize that when these are present in sub- pl. 34, figs. 7 and 8. stantial numbers, they need to be reported Age, Keuper. and that a superficial resemblance to 30 PALEOZOIC FOSSIL SPORES

Punctati-sporites may not always be in- Six species previously assigned to Ver- dicative of relationship. The affinity of mcosi-sporites Ibrahim, including the certain of the species with the Crossotheca type, are here placed under Pulzctati-spor- alliance of pteridospernis seems beyond ites. Yel-frz~cosi-spofpitesthus would more much question, however. reasonably be taken as the designation The forms which Raistrick (1933, for the genus except that in meaning one 1934. 1937) has assigned to his 'types name is no niore suitable than the other B,, B,, D,, D,, Dl, (K~OX,1938)) E,; E,, and strict interpretation of page priority E,, Es, E,, F,, Fs (Knox, 1938) (also (p. 21 vs. p. 25) favors Puwtati-sporites. 1K( ?), 2K and 7K, Knox, 1942) appear The genus also includes the type of entirelv or in art referable to Punctati- Apiculata-sporites Ibrahim ( 1933)) a spo&s. It also may include some of nionotypic genus which seems based en- the forms he has classed as B, and B,. tirely on Ibrahini's inability to observe Some difficulty is encountered because the trilete sutures described by Loose Raistrick evidently has designated no sin- (1932) on Sporonites spinulistratus. The gle typical example for these groups and presence of the trilete marking on this his illustrations of the forms have differed species was reaffirmed by Loose in 1934. for individual types beyond what may In any event the genus Apiculata-sflorites represent a specific range of variability. does not seem securely fou~icled and its Thus there may be some question as to one species is here placed under Punctati- the consanguinity of forms labeled as D, sporites. Both groups, Yerrucosi-sporites (cf. also Raistrick, 1935). Figures and Ibrahim and Apiculata-spovites Ibrahim, description of El given in 1933 do not ex- conseqently are regarded as congeneric actly correspond with that given in later with and the names synonyms of Puvzctati- papers ( 1934, 1935). Illustrations of sporites. B, .given in 1933 seem to resemble Punc- The holotype of the type species, Punc- tatz-sflorites but those illustrated in later tati-sporites punctatzts (Ibrahim) Ibra- papers without much doubt are referable him, is from the Aegir coal seam at the to Calamospova 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-SPORITESACULEATUS (Ibrahim) alences in most instances. S. W. and B., comb. nov. Apiculati-sporites acz~leatus Ibrahim, Remarks.-The genus Pulqctati-spovites 1933, Sporenformen des Aegirhorizonts, p. as given here includes twenty-nine species, 23, pl. 6, fig. 57. four of which are queried. All but three 2. PGNCTATI-SPORITESAUREUS (Loose) S. W. of these names (see spp. Nos. 9, 21, 22) and B., comb. nov. are new combinations. The group Pum- Reticula ti-sporites aureus Loose, 1934, tati-sporitcs was originally defined by Inst. Paleobot. Arb., vol. 4, no. 3, p. 155, Ibrahim (1933, p. 21) as including trilete pl. 7, fig. 24. type spores with a granular surface, char- acters which by then~selves can hardly 3. PUNCTATI- SPORITES ( ?) AURICULAFERENS (Loose) S. W. and B., comb. nov. be accepted as diagnostic of any generic Sporonites az~riczdaferens Loose, 1932, group. Three species were described, one Neues Jahrb., Beilage-Band 67, Abt. B, p. of which, P. pumtatus, was designated as 450, pl. 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. pz~~zctatus,and 4. P UNCTATI-SPORITES BUCCULENTUS (Loose) the emendecl generic diagnosis has been S. W. and B., comb. nov. constructed on this basis. Neither of the Ye?~rucosi-spo14tesbz~ccz~le~ztz~s Loose, other two species Ibrahin? originally 1934, idem. p. 154, pl. 7, fig. 15. placed in the genus are considerecl closely enough related to remain there ; P. parvus 5. PUNCTATI-SPORITESCORRUGATUS (Ibrahim) S. IV. and B., comb. nov. Ibrahim is now assigned to the genus Reticz~lati-sporites cowz~gatzuIbrahim, GI-anulati-s~orites, a 11 d P. pallidus 1933, Sporenformen des Aegirhorizonts, pp. (Loose) Ibrahim to Calavtospora n. gen. 35-36, pl. 5, fig. 41. PUNCTATI-SPORITES

PUNCTATI-SPORITEScusus (Loose) S. W. Tubercztlati - spovites microtuberosus and B., comb. nov. (Loose) Ibrahim, 1933, Sporenformen des Reticztlati-sporites cztsus Loose, 1934, Aegirhorizonts, p. 23. Inst. Palaobot. Arb., vol. 4, no. 3, p. 156, pl. Tu berculati - sflorites microtuberosus 7, fig. 25. (Loose) Ibrahim, Loose, 1934, Inst. Palao- bot. Arb., vol. 4, no. 3, p. 147.

PUNCTATI-SPORITESFIRMUS (Loose) S. W. PUNCTATI-SPORITESMICROVERRUCOSUS (Ibra- and B., comb. nov. him), S. W. and B., comb. nov. Verrztcosi-sporites firmus Loose, 1934, Verrucosi - sporites microverrucosus idem, p. 154, pl. 7, fig. 30. Ibrahim, 1933, Sporenformen des Aegirhori- zonts, p. 25, pl. 7, fig. 60. PUNCTATI-SPORITESGLOBOSUS (Loose) S. W. and B., comb. nov. PUNCTATI-SPORITESNOBILIS (Wicher) S. Apiculati-sporites globosus Loose, 1934, W. and B., comb. nov. idem, p. 152, pl. 7, fig. 14. Sporites nobilis Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 186, pl. 8, fig. 30. PUNCTATI-SPORITESGRANDIVERRUCOSUS Kosanke, 1943, Am. Midland Naturalist, PUNCTATI-SPORITESPAPILLOSUS (Ibrahim) vol. 29, no. 1, pp. 127, 1281, pl. 3, fig. 4. S. W. and B., comb. nov. Verrucosi-sfiorites papillosus Ibrahim, PUNCTATI-SPORITESGRANIFER (Ibrahim) S. 1933, Sporenformen des Aegirhorizonts, pl. W. and B., comb. nov. 5, fig. 44. Grar~ulati- sporites gra?r;if er Ibrahim, 1933, Sporenf ormen des Aegirhorizonts, p. PUNCTATI-SPORITESPARVIPUNCTATUS KOS- 22, pl. 8, fig. 72. anke, 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 127, pl. 3, figs. 3, 3a, 3b. GRUMOSUS PUNCTATI-SPORITES (Ibrahim) S. PUNCTATI-SPORITESPUNCTATUS (Ibrahim) W. and B., comb. nov. Ibrahim, 1933, Sporenformen des Aegirhori- Verrucosi-sporites grumosus Ibrahim, zonts, p. 21, pl. 2, fig. 18. 1933, idem, p. 25, pl. 8, fig. 68. Sporonites pztnctatus Ibrahim, 1932, Neues Jahrb., Beilage-Band 57, Abt. B, p. PUNCTATI-SPORITESINSIGNITIS ( Ibrahim) 448, pl. 15, fig. 18. S. W. and B., comb. nov. Punctati-sporites pztnctatus (Ibrahim) Apiculati - sporites insignitis Ibrahim, Ibrahim, Loose, 1934, Inst. Palaobot. Arb., 1933, idem, p. 24, pl. 6, fig. 54. vol. 4, no. 4, p. 146, pl. 7, fig. 27.

( ?) RETICULOCINGULUM PUNCTATI-SPORITES( ? ) IRREGULARIS (Ber- PUNCTATI-SPORITES ry) S. W. and B., comb. nov. (Loose) S. W. and B., comb. nov. Zonales-sporites irregularis Berry, 1937, Sporonites reticulocingulum Loose, 1932, Am. Midland Naturalist, vol. 18, no. 1, p. Neues Jahrb., Beilage-Band 57, Abt. B, p. 156, fig. 9. 450, pl. 18, fig. 41. Reticztlati ' - sporites reticulocingulum PUNCTATI-SPORITESLACUNOSUS (Ibrahim) (Loose) Ibrahim, 1933, Sporenformen des S. W. and B., comb. nov. Aegirhorizonts, p. 34. Reticulati - spovites lacunosus Ibrahim, Reticulati - sporites reticztlocingulunt 1933. Sporenformen des Aegirhorizonts, p. (Loose) Ibrahim, Loose, 1934, Inst. Palao- 36, pl. 6, fig. 50. bot. Arb., vol. 4, no. 4, p.. 156. PUNCTATI- SPORITES SPHAEROTRIANGULATUS PUNCTATI-SPORITESLATIGRANIFER (Loose) (Loose) S. W. and B., comb. nov. S. W. and B., comb. nov. Sporonites sphaerotriangulatus Loose, Sporonites latigranif er Loose, 1932, 1932, Neues Jahrb., Beilage-Band 67, Abt. Neues Jahrb., Beilage-Band 67, Abt. B, p. B., p. 451, pl. 18, fig. 45. 452, pl. 19, fig. 54. Laevigati - sporites splzaero triangulatus Granulati-sporites latigmnif er (Loose) (Loose) Ibrahim, 1933, Sporenf ormen des Loose, 1934, Inst. Palaobot. Arb., vol. 4, Aegirhorizonts, p. 20. no. 3, p. 147. Laevigati - spovites sphaerotriangulatus (Loose) Ibrahim, Loose, 1934, Inst. Palao- PUNCTATI-SPORITESMACULATUS ( Ibrahim) bot. Arb., vol. 4, no. 4, p. 145. S. W. and B., comb. nov. PUNCTATI-SPORITESSPINOSUS (Loose) S. Reticulati-sporites maculatzts Ibrahim, and B., comb. nov. 1933, Sporenf ormen des Aegirhorizonts, p. W. 36, pl. fig. 56. Apicz~lati-sporitesspinosus Loose, 1934, 6, idem, p. 153, pl. 7, fig. 20. PUNCTATI-SPORITESMICROTUBEROSUS PUNCTATI-SPORITESSPINULISTRATUS (Loose) (Loose) S. W. and B., comb. nov. S. W. and B., comb. nov. Sporonites microtuberosus Loose, 1932, Sporonites spinu.listratzts Loose, 1932, Neues, Jahrb., Beilage-Band 67, Abt. B, Neues Jahrb., ~eilage- and 67, ~bt.B, p: p. 450, pl. 18, fig. 33. 451, pl. 18, fig. 47. 32 PALEOZOIC FOSSIL SPORES

Apiculata-sflorites s~inztlistratus(Loose) Spore coat. - Of relatively uniform Ibrahm, 1933, Sporenformen des Aegir- thickness throughout, generally thin, and horizonts, p. 37. except where modified by ornamentation, Apiculati-sporifesspinulistratus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. often less than 2 microns. 4, p. 153. Afinity.-Spores similar to those of 27. PUNCTATI- SPORITES TRIGONORETICULATUS G~amtlati-sporites have been reported (Loose) S. W. and B., comb. nov. from fructifications of ferns although rel- Reticulati - sporites tvigono~*eticulatzts atively few of these have been critically Loose, 1934, idem, p. 155, pl. 7, fig. 9. described. Spores of Boweria minor and 28. PUNCTATI-SPORITES( ?) VELATUS (Loose) Renaultin gracilis which are illustrated by S. TV. and B., comb. nov. Knox (1938) are of similar character. Reticulati-spo~ites velatz~sLoose, 1934, Probably considerable difficulty will be idem, p. 155, pl. 7, fig. 19. encountered in definitely correlating the genus (further emended as may prove 29. PUNCTATI-SPORITESVERRUCOSUS (Ibrahim) S. W. and B., comb. nov. desirable) with any single supra-generic Sporonites verrzrcosus Ibrahim, 1932, plant group. The many similar features Neues Jahrb., Beilage-Band 67, Abt. B, p. possessed by species assigned to G~ravzulati- 448, pl. 15, fig. 17. sporites suggest that this grouping may be Ve~rucosi-sporitesvermcosus ( Ibrahim) a practical one. Ibrahim, 1933, Sporenformen des Aeglr- horizonts, p. 25, pl. 2, fig. 17. The forms Raistrick (1933, 1931, 1937) designates as D,, D, and D,, (Knox, Genus GRANULATI-SPORITES(Ibrahim, 1938) probably belong to Gramlati-spur- 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 Revmarks: The genus as emended in- Symmetry.-Spores radial, trilete. cludes fifteen species, twelve of which Shape.-Originally oblate, rounded tri- represent new name combinations. The angular, with the axial dimension much genus was originally established to include the shortest; when compressed the spores spores of trilete-type showing granular are nearly always flattened in good proxi- sculpturing (Ibrahim, 1933, p. 21)) char- mo-distal orientation since the transverse acters which in themselves are inadequate ; 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 outline, 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 Gramlati-sporites Size.-Spores of various species range is tlie only genotype species included by from about 25 to microns in mean revision within the genus and thus there 45 can be no question as to validity of tlie diameter. generic name. The genotype, G. granulatus Or~zamentation.- Various, surf aces Ibrahini, 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- plialian B. verse types of ornamentation which char- acterize various species are not prominent 1. GRANULATI-SPORITESDELTIFORMIS S. W. and to the extent that they mask the essential B., nomen nov. shape characteristics of the spores. Triquit~itesdeltoides Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. Haptotypic structures.-Trilete rays 1, p. 185, fig. 9. are well extended toward the corners; Note.-In combination with Granulnti-sporites they are never relatively short. The lips the specific epithet becomes a homonym of G. are not distinguished by special ornamen- deltoides (Ib.) S. W. and B., as given below. tation, the commiss~lreis relatively simple but definite in all cases; usually no vari- 2. GRANULATI-SPORITESDELTOIDES (Ibrahim) S. W. and B., comb. nov. ations in ornamentation distinguish the Sporonites deltoides Ibrahim, 1932, proxinlal (pyramic) side from the rest Neues Jahrb., Beilage-Band 67, Abt. B, p. of the spore coat. 448, pl. 15, fig. 15. ALATI-SPORITES 33

Laevigati-sporites deltoides (Ibrahim) 13. GRANULATI-SPORITESTRIGONUS (Ibrahim) Ibrahim, 1933, Sporenformen des Aegir- S. W. and B., comb. nov. horizonts, p. 20, pl. 2, fig. 2. Reticulati-sporites trigonus Ibrahim, Note.-The species is larger than most forms 1933, Sporenformen des Aegirhorizonts, p. assigned to Granulati-sporites. 37, pl. 5, fig. 34.

3. GRANULATI-SPORITESFISTULOSUS ( Ibrahim) 14. GRANULATI-SPORITESTRIQUETRIS (Ibrahim) S. W. and B., comb. nov. S. W. and B., comb. nov. Reticulati-sporites fistulosus Ibrahim, Vewucosi-sporites triquetris Ibrahim, 1933, idem, p. 36, pl. 5, fig. 35. 1933, idem, p. 26, fig. 61. 15. GKANULATI-SPORITESVERRUCOSUS (Wilson 4. GRANULATI-SPORITESGIBBOSUS (Ibrahim) and Coe) S. W. and B., comb. nov. S. W. and B., comb. nov. Triquitrites verrztcosus Wilson and Coe, Verrucosi-sflorites gibbosus Ibrahim, 1940, Am. Midland Naturalist, vol. 23, no. 1933, idem, p. 25, pl. 6, fig. 49. 1, p. 185, fig. 10. 5. GRANULATI-SPORITESGRANULATUS Ibrahim, 1933, idem., p. 22, pl. 6, fig. 51. Genus ALATI-SPORITESIbrahim, 1933 Note.-Type species of Granulati-sporites. Plate 1, figures 6-6b GRANULATI-SPORITESMICROGRANIFER Ibra- him, 1933, idem, p. 22, pl. 5, fig. 32. Symwl.tetry.-Spores radial, trilete. Shape.-Body, exclusive of bladders, is GRANULATI- SPORITES MI,CROSAETOSUS (Loose) S. W. and B., comb. nov. sub-triangular. Sporonites microsaetoszts Loose, 1932, Size.-In the vicinity of 80 niicrons, Neues Jahrb., Beilage-Band 67, Abt. B, p. overall diameter. 450, pl. 18, fig. 40. Ornai~zentntio~aof spore.-Body moder- Setosi-sporites .~nicrosaetoszcs (Loose) ate ; may be punctate, finely reticulate ; Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 26. bladders sniooth to punctate, etc. Blad- 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 GRANULATI-SPORITESMICROSPINOSUS (Ibra- relatively long, little ornamented. him) S. W. and B., comb nov. Spore coot.-Consisting of two distinct Apiculati-sporites ~icrospinosus Ibra- membranes ; the outer (perisporal) niem- him, 1933, Sporenformen des Aegirhori- brane very and expanded in- zonts, p. 24, pl. 6, fig. 52. is thin in the ter-radial areas to form bladders in a trim- GRANULATI-SPORITESPARVUS (Ibrahim) S. erous (in threes or multiples of three) W. and B., comb. nov. pattern. The spore body wall (exospore) Sporonites parvz~sIbrahim, 1932, Neues is usually more than twice as thick as the Jahrb., Beilage-Band 67, Abt. B, p. 448, pl. 15, fig. 21. bladder membrane, darker, and easily Punctati - sporites parvz~s (Ibrahim) distinguished on that account. Ibrahim, 1933, Sporenformen des Aegir- Afinity.-Spores of Alati-sporites seem horizonts, p. 21, pl. 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. 4, p. 154, pl. 7, fig. 13. 1898, Kubart, 1910). The three inter- radial bladders are most distinctive in GRANULATI- SPORITES PIROFORMIS Loose, both genera. The significance of this 1934, idem, vol. 4, no. 3, p. 147, pl. 7, fig. 19. character is hardly well enough under- GRANULATI-SPORITES(?) PRIDDYI (Berry) stood to properly evaluate evidence as S. W. and B., comb. nov. to their relati~nship.~ 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 Podocar us dacrydiordes, and on 156, fig. 2. Pheioiphaera are not known, an8 com arrson wrth Car- boniferous forms does not seem particurarly pertinent for GRANULATI-SPORITESTORQUIFER (Loose) S. obvious reasons. It seems doubtful that the form described by Daugherty W. and B., comb. nov. (1941 p. 64) as Spencerites? chinleana is actually refer- Sflorodes torquifev Loose, 1932, Neues able ;o Spence~itesor could be referred to Alati-spori!es since it apparently does not posse!s three mterradlal Jahrh., Beilage-Band 67, Abt. B, p. 450, bladders. Daugherty does not mentlon and the figure 18, fig. 43. given does not show the trilete sutures. If these were - er present the species might be referred to Cirrairzradrtes Reticzlleti sporites toTqzLif (Loose) provided the equatorial appendage was a flange: Lacking L~~~~,1934, I~~~.palaobot. ~~b,, 4, no, these characterlst~cs his alternative suggestion of a 3, p. 154. gymnospernlic affini6 seems most plausible. 34 PALEOZOIC FOSSIL SPORES

Spores assigned by Raistrick to type Size.-Spores of various species range D, without much question belong to from about 40 to 100 microns in diameter. Alati-sp orites. 0mamentatio~z.-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. purpose. There is no possibility of these Haptotypic structures. - The trilete f orms being confused with unseparated structures may be weakly developed; no tetrad groups belonging to other genera arcuate ridges are developed but the or- although illustrations which have been namentation may sonietimes be absent on published might lead one to think so. 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 froni 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- The generic diagnosis given above has al?) niembrane is thicker and sometimes been constructed in the light of this irregular and excentric in its development. additional information, and while con- The wall may appear thick, and yet be siderably augmented over Ibrahim's mea- quite translucent. ger description (1933, p. 32), it may be improper to consider it as a generic emend- Afi9zities.-The only spores of this gen- ation. The type, of course, is the species eral or possible character known in fruc- given below, which was first described tifications are allied with Sphenophy1lzl.m. from the Aegir coal in the Ruhr. Sphenophyllum spores are inadequately known in the detail necessary for close 1. ALATI - SPORITES PUSTULATUS (Ibrahim) comparison, however, and considerably Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 33, pl. 1, fig. 12. more evidence is needed to form an opin- Spovonites pus tula tus Ibrahim, 1932, ion as to the relationship of Reticulati- Neues Jahrb., Beilage-Band 67, Abt. B, p. sporites with other groups. * 448, pl. 14, fig. 12. Types C,, C,, F,, and F, of Raistrick Alnti - spodes pustulatzts (Ibrahim) Ibrahim, Loose. 1934, Inst. Palaobot. Arb.. (1934, 1935, 1937) and possibly also F,, vol.4, no. 3, p. 151, pl. 7, fig. 4. F, (cf. Knox, 1938), F, and F,, appear Type D5 Raistrick, 1937, Congres pour to belong to Reticulati-sporites. Types l'avancement des etudes de stratigraphie F, and F, and Dl, (Knox, 1938) are carbonifere, Heerlen, 1935, Compte rendu, vol. 2, p. 911. more problematic. Type 8 of Millott (1939) also belongs here and is most Genus KETICULATI-SPORITES(Ibrahim, closely related to ~.?acetusof species now 1933) emend., S. W. and B. described. Knox' Type 5K may possibly be similar to some species included in Re- Plate 1, figures 7-7b ticulati-spo~ites. Spores illustrated by Sywmetvy.-Spores, in essence, radi- Knox (1939) from Fifeshire, figs. 49 and ally syn~metrical; some species are weakly 50, unquestionably belong to Reticulati- trilete and others show no indication of spmrites. Uiipublished 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. erately ohlate ; con~pressedsnores are cir- Rewza~ks.-Seven species are included cular disk-like, generally without prom- here under Reticulati-spo~ites, only one inent folds. of which represents a new name conibina- RETICULATI-SPORITES 35 tion. This form, R. facetus (Ibrahim), spo~itesapparently lack the further spe- was originally designated as the genotype cialization of germiiial pores, neverthe- of Reticulata-sporites with five species less lack of good development of a trilete assigned to it. Essentially the only char- commissure in some of the fornis 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 ticzdati-spo&es appears to be a relatively a type of reticulation which is very simi- homogeneous group of natural relation- lar to that of R. reticztlatus (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 fornis 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 interpreta t' Ion. is i~isufficientbasis 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 Reticdata-sporites Ibrahim (1933, p. coal bed at the top of the Westphalian B, 38) may be clispensed 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 (1939) has suggested that alete 1. RETICULATI-SPORITESCORPO~US Loose, 1934. spores with somewhat splsal 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-SPORITESFACETUS (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. seem appropriate because the fossil forms 38, pl. 5, fig. 36. evidently have a perisporal membrane en- 3. RETICULATI-SPORITESMEDIARETICULATUS Ib- closing them instead of mere "large alve- rahim, 1933, idem, p. 34, pl. 7, fig. 62. olae with high lainellae." Nwertheless it 4. RETICULATI-SPORITESORNATUS (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 to some of the spores of the Hepaticae il- Jahrb., 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. RETICULATT-SPORITESPOLYGONALIS (Ibra- of haptotypic markings on several of the him) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 1.55, 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- 417, 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, Sporenf ormen des back at least to the Carboniferous. Aegirhorizonts, p. 19, pl. 1, fig. 8. It may be suggested that spores of 6. RETICULATI-SPORITESRETICULATUS (Ibra- him) Ibrahim, 1933, idem, pp. 33-34, pl. 1, Reticulati-sporites underwent more mod- fig. 3. ification within the sporangium subsequent Spol,onites reticz~latz~sIbrahim, 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 reticdation pattern prob- Reticzdati-sporites reticulatus (Ibrahim) Ibrahim, Loose, 1934, Inst. Palaobot. Arb., ably was laid down relatively late in sporo- vol. 4, 110. 3, p. 156. genesis by a tapetal plasmodium, some- Note.-Type species of Reticz~lati-sporitcs. tinies without respect for the proximo- distal relationships established in original 7. RETICULATI-SPORITES( ?) RETICULIFORMIS Ibrahim, 1933, Sporenf ormen des Aegirhori- tetrad groupings. Such a sequence occ~lrs zonts, p. 34, pl. 7, fig. 63. in formation of many kinds of angi- Note.-Trilete rays are unusually long and ospermjc pollen. Spores of Reticulati- well developed in this species. 36 PALEOZOIC FOSSIL SPORES

NOMINAEXCLUDENDE are sometimes distinguishable flaring lat- 1. RETICULATI-SPORITESANGULATUS Ibrahim, 1933, Spo- erally from both ends of the suture line. renformen des Aegirhorizonts, etc., p. 35, pl. 7, fig. 59. Spore coat.-Varying somewhat in 2. RETICULATA-SPORITESBIRETICULATUS (Ibrahim) Loose 1934, Inst. PalPobot. Arb., vol. 4, no. 3, p. 158: thickness relative to the other dimen- pl. 7, fig. 28. Sporonites bireticulatus Ibrahim 1932 Neues sions; the spore coat is oftentimes thin Jahrb., Beilage-Band 67, Abt. B, i.447,'pl. 14, and translucent. The internal cavity is fig. 1. Reticulati-sporites bireticulatus (Ibrahim) Ibra- sometimes more "bean-shaped" than the him, 1933 S orenformen des Aegirhorizonts, etc., external outline due to internal thickening p. 35, pl. ,I, %g. 1. 3. RETICULATI-SPORITESFACIERUGOSUS LOOS~1934, Inst. of the spore coat in the central proximal Paliiobot. Arb., vol. 4, no. 3, p. 155, pi. 7, fig. 26. region. 4. RETICULATA-SPORITESMEDIAPUDENS Loose, 1934, idem, p. 158, pl. 7, fig. 8. Afinity .-Smaller species of Laeviga- 5. RETICULATI- SPORITES MICRORETICULATUS (Loose). , Loose, 1934., idem,, p. 155. to-sporites agree with some forms ob- Sporonrtes m~croret~culatusLoose 1932 Neues Jahrb., Beilage-Band 67, Abt. B, p. 450, fig.'37. tained from filicinean type fructifications. 6. RETICULATI-SPORITESMOROSUS Loose, 1934, Inst. There is now no particularly good evi- PalHobot. Arb., vol. 4, no. 3, p. 154, pl. 7, fig. 2. 7. RETICULATI-SPORITESNEXUS (Loose) Ibrahtm, 1933, dence as to the affinity of the larger forms. Sporenformen des Aegirhorizonts, etc: 34. Sporonites nexus Loose 1932 eues Jahrb., Fredda Reed (1938, p. 333) has obtained Beilage-Band 67 Abt B p '450 fii 35. spores of this type from an extraordinary Reticulati-s >rites' n&u; (~o&e) brahi him, Loose. 1934. Inst. PaPaobot. Arb., vol. 4, no. 3, p. 156, new type of Calamarian fructification pl. 18, fig. 35. which are approximately 50 microns in 8. RETICULATI-SPORITESSIFATI Ibrahim 1933 S oren- formen des Aegirhorizonts, etc., p. 35, pl. i, &. 67, length. Spores of Zeilleria are of this 9. RETICULATA-SPORITESSPONGIOSUS Ibrahim, 1933, type (Florin, 1937, p. 316-7) but it is not idem, p. 39, pl. 8, fig. 71. 10. RETICULATI-SPORITESVINCULATUS (Ibrahim ) Loose, now possible to decide whether that genus 1934, Inst. Pallobpt. Arb., vol. 4, no. 3, p. '156. Sporonrtes vrncularus Ibraham 1932, Neues belongs to the ferns or pteridosperms !a_hrbx Beilage-Band 67, Abt. B, b. 448, pl. 15, (Halle, 1933, p. 88). The "distal fur- 11g. iy. Reticulata-sporites vinculatus (Ibrahim) Ibrahim, row" of Zeilleria spores remarked by 1933, Sporenformen des Aegirhorizonts, etc., p. 39, pl. 2, fig. 19. Florin (1937, p. 317) appears no 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- Plate 1, figures 5-5b gato-sporites could conceivably pertain to 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 shaped; elongate oval in the plane of lon- the immediate future and until it is 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 n~orphology of the a good many instances species of Laevi- various specific types of spores. giato-sflorites form a majority of the Size.-Spores of various species range spores obtained from certain coal beds. from about 20 to over 130 microns in They are, however, rare or absent in a their long dimension. number of lower. Pottsville coals that have been examined. 0rnawzentatio.n.-Smooth to finely punc- tate, apiculate or rugose; rarely showing Spores evidently belonging to Laevi- pronounced reticulation or strong apicu- pto-sporites have been assigned to types lae. B,a, B,b, and B, by Raistrick (1933, Haptotypic structures.-Consisting of 1937) (Knox, 1938). Millott's types 6 a simple monolete linear suture, generally and E,a (Millott 1939) and Knox' (1942) without lips specially distinguished, and Type 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 forwzae, are now be very inconspicuous if it coincides in listed for Laevigato-spo&es. 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 Naturalist, vol. 18, no. hand a number of distinct and as yet un- 1, pp. 156-7, fig. 4. recorded species have been recognized in 7. LAEVIGATO-SPORITESTHIESSENII Kosanke, the Illinois basin and elsewhere. 1943, Am. Midland Naturalist, vol. 29, no. 1, p. 125, pl. 3, figs. 1, la, lb. The list includes genotype species of Pittsburgh microspore of Thiessen, various iour previously named genera. L. bila- publications. See especially Thiessen and Staud, teralis (Loose) was the only species de- 1923. scribed by Loose (1934, p. 159) under 8. LAEVIGATO-SPORITES( ?) TUB~CULATUS the new name, Reticulato-sporites. The (Berry), S. W. and B., comb. nov. name was credited to "Ibrahin~1932" al- Tz~berculati-sporitestuberculatus Berry, though no "reticulato" was ever used by 1937, idem, vol. 18, no. 1, p. 155, fig. 9. that author. L. des7~zoi~ze1zsisis the gen- 9. LAEVIGATO-SPORITESVULGARIS (Ibrahim) otype of Plzaseolites Wilson and Coe Ibrahim, 1933, Sporenf ormen des Aegirhor- (1940). L. ~~inutzts(Ibrahim) was the izonts, pp. 39-40, pl. 2, fig. 16; pl. 5, figs. 37, 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, pl. 15, fig. 16. Laevigato-sporites Ibrahim (1933, p. 39). LAEVIGATO-SPORITESVULGARIS f orma 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, pl. 7, fig. 12. variation of surf ace ornament and small LAEVIGATO-SPORITESVULGARIS forma MAIOR differences of size, none of which can be Loose, 1934, idem, p. 158, pl. 7, fig. 6. regarded as of more than specific impor- LAET~IGATO-SPORITESVULGARIS forfna MAX- tance. The names Reticulato-sporites, IMUS Loose, 1934, idem, p. 158, pl. 7, fig. 11. Phaseolites, and Punctato-sporites are therefore regarded as synonyms of Laevi- 1. LAEVIGATO-SPORITESRHEAENSIS 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-SPORITESIbrahim, 1933 1. LAEVIGATO-SPORITESBILATERALIS (Loose), S. W. and B., comb. nov. Symmetry.-Spores apparently bilat- Reticulato-sporites bilateg*alis Loose, eral, monolete. 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. Sha$e.-Flattened elliptical ( ?), broad- 159, pl. 7, fig. 22. ly elliptical in transverse plane. Note.-Reticzdato-sflorites is a generic name published only by Loose (1934) although he Size.-Relatively large, 200-300 rni- erroneously credited it to "Ibrahim, 1932." crons in length. Onza~~wntation.- Surface smooth to 2. LAEVIGATO-SPORITESCRANMORENSIS Berry, 1937, Am. Midland Naturalist, vol. 18, no. minutely rugose. 1, p. 157, fig. 1, Hapto typic f eatures.-Linear proximal suture with slight angular median deflec- 3. LAEVIGATO-SPORITESDESMOINENSIS (Wilson and Coe) S. W. and B., comb. nov. tion, termini of arcuate ridges niay be Plzaseolites desmoitze~zsis Wilson and weakly cliff erentiated. Coe, 1940, idem, 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 z~ulgatris. sides, and in contact with it proximally and distally. Body wall (exospore) rela- 4. LAEVIGATO-SPORITESMINIMUS (Wilson and Coe) S. W. and B., comb. nov. tively thick and brownish translucent. Phaseolites nzinimus Wilson and A fiaity . - Probably pteridospermic, Coe, 1940, idem, vol. 23, no. 1, p. 183, fig. 5. judging fr-om points of similarity with Monoletes which is discussed later. It 5. LAEVIGATO-SPORITESMINUTUS (Ibrahim) S. W. and B. comb. nov. nevertheless is still extremely problematic Punctato-s~orites minutus Ibrahim. since it superficially seems to combine 1933, ~~orenfo&endes Aegirhorizonts, p: features of spores of Medullosan and of 40, pl. 5, fig. 33. Cordaitean relationship. 38 PALEOZOIC FOSSIL SPORES

Remarks.-This genus is reported in and diagnostic feature is the slight angu- only two instances, in the Ruhr and in lar deflection of the suture line near the Illinois, in both cases from relatively few middle. Although inconspicuous and specin~ens. 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-SPORITESAUREOLUS (Schopf ) S. W. and B., comb. nov. ges are occasionally resolved at the two Monoletes aureolus Schopf, 1938, Illi- ends of the functional suture line. The nois Geol. Survey Rept. Inv. 50, pp. 45-46, suture opens to form a sharp taper- pl. 1, figs. 1-2. pointed lenticular slit. The distal grooves have no connection with haptotypic struc- 2. ZONALO-SPORITESVITTATUS Ibrahim, 1933. Sporenformen des Aegirhorizonts, p.- 41, pl: tures. 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. unlbo are as much as 15 or 18 inicrons; Plate 2, figures 17-17c the thinnest is at the base of the distal Synwzetry.-Spores apparently bilat- grooves on either side of the umbo where eral, monolete. Mature forms evidently it may be less than 5 n~icrons. An even have been modified considerably in post gradation exists between these extreme tetrad maturation, and it is possible that areas and in normally compressed spores although the mature forms appear very the exospore thickness at the margin is definitely bilateral, they may have arisen intermediate between the extremes. The from tetrahedral tetrads. endosporal n~en~braneis frequently evi- dent as a crumpled transluceiit 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 f uiictioning as a harmon?egathus. transverse axis shows a somewhat rounder and shorter but notably flattened outline. Afi9zity.-Spores of this character dis- The distal surface oftentimes is marked tinguish a pteridospernlic plant alliance by two prominent grooves extending near- probably largely coexteiisive with the Me- ly the length of the spore with a well dullosaceae. They are characteristic of rounded umbo between. Sometimes the all genera placed in the Whittleseyinean proximal side near the suture shows a sub-tribe (see Halle, 1933). The only very slight prominence. On compression, aberrant feature is the lack of distal longitudinal folds frequently parallel the grooves in spores of Codonotheca; spores distal grooves. of all other genera now assigned to this alliance possess them. Codonotheca spores Size.-Spores relatively large, varying are otherwise entirely characteristic of from slightly over 100 microns to as Mo~zoletes and when such spores are much as half a millimeter in length. found isolated they probably should be Ornumen ta tion. - Surf aces generally assigned to Monoletes since there well minutely granulose appearing quite may be other types in addition to Codon- smooth at low nlagnification; 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 noii-essential character of these extreme base of distal grooves, which are specialized distal strtlctures. more smooth. Re?~uwks.-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 Dole~~othecaand 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. MOROLETESOVATUS Schopf, 1935, Illinois tions the proximal position of the suture Acad. Sci. Trans. vol. 28, no. 2, p. 108, fig. 7. Pollen of Dolerophyllztm sp., Zerndt, and the distal location of the harmome- 1930, Acad. polonaise sci. Bull. internat., gathic grooves. The fact that spores are ser. B, pp. 55-56, pl. 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 iVonoletes ovatus Schopf, 1935, Illinois been observed with the umbo split off as Acad. Sci. Trans. vol. 28, no. 2, p. 176. ~Vonoletesovatus Schopf, 1938, Illinois an "operculum" or the grooves other than Geol. Survey Rept. Inv. No. 50, pp. 43-45, intact, stropgly suggests that proximal pl. 1, figs. 3-5, pl. 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, pl. 11, fig. 36. cestry. Such spores are also regarded as prepollen (cf. Schopf, 1938, pp. 14-15). Genus DENSO-SPORITES(Berry, 1937), 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- circling ridge" described by Schopf (1938, Sywmetry.-Sposes radial, trilete. p. 45) for the species designated as Mogzo- S1zape.-Originally oblate, round to letes owatus, is the margin of the distal subtriangular in equatorial outline. When grooves discussed here. It has no hapto- compressed, the central area becomes typic significance although it possibly is much thinner than the margins due to va- not emphytic in origin in the same sense riation in spore wall thickness. Coal thin that ordinary surface ornanientation is. sections, taken vertically through the coal bed, show then1 compressed into a "dumb- iWo.lzoletes was first used by Ibrahim in bell" shape. The original highly oval sec- 1933 to designate a new group in his ar- tional f orin results in extremely uniform tificial system. Although substalztive in proximo-distal orientation in enclosing fosm and siiiiilar to a generic lzanie, it sediments. was not originally applied in that sense and no nomenclatural type has been pre- Sixe.-Diameter of specimens xssigned viously proposed. It has been used as a among various species varies f 1-0111 about peneric designation and applied more nar- 35 to 100 microns. rowly (Schopf, 1936, 1938) ; we now Onzamentation. - Smooth to apiculate emend it to correspond with the restricted and rugose. External surface character usage. Monoletes owatus 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 str.uct.ures.-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 "Fissuses" penetrating to the margin in generalized types. some instances may be essentially hapto- typic continuations of the trilete sutures. 1. MONOLETESELLIPSOIDES (Ibrahim) Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, Spore coat.-Characterizecl by great p. 45, pl. 1, fig. 14; pl. 6, figs. 5 and 6. differelices in thickness. The proximal Sporo~zites eZlipsoidcs Ibrahim, 1932, and distal walls are usually membranous, Neues Jahrb. Beilage-Band 67, Abt. B., p. or at least significantly thinner than the 449, pl. 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 Aegirhorizonts, p. 40, pl. 4, fig. 29. opaque to transmitted light in contrast to Punctato-sporites ellipsoides (Ibrahim) the highly translucent central area. It Loose, 1934, Inst. Palaobot. Arb., vol. 4, may also be so extended centrifugally as no. 3, pp. 158-9, pl. 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

Afinities.-Spores of this genus have mation about these spores may reflect not been recognized in connection with importantly, although indirectly, upon the fructifications. This is surprising in view problem of splint coal formation. of their frequence in coal. They have 1. DENSO-SPORITESANNULATUS (Loose) S. been roughly designated as "biconcave," 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, pl. (Thiessen and VCTilson,p. 9, 1924; Thies- 18, fig. 44. sen, 1930 ; Sprunk, et al, 1940) and can be Zonales - sporifes annulatus (Loose) recognized by their characteristic "dumb- Loose, 1934, Inst. Palaobot. Arb., vol. 4, bell" shape in cross se~tion.~ no. 3, p. 151.10 Modern methods of study may be ex- 2. DENSO-SPORITESCOVENSIS Berry, 1937, Am. pected therefore to disclose definite evi- Midland Naturalist, vol. 18, p. 157, fig. 11. dence of the relationship of this group. hTote.-Although serving as genotype, the spe- cific characteristics of this form are inadequately Their structure is so un~lsualthat it seenls known. evident that a single honlogeneous group of plants for the most part is represented. 3. DENSO-SPORITESDENSUS Berry. 1937. Am. Midland Naturalist, vol. 18, no: '1, pp. '157-8, Raisti-ick's Type A spores (~aistrick fig. 7. and Simpson, 1933) apparently are nearly entirely referable to Demo-sfiorites. His 4. DEKSO - SPORITES ( ?) INDIGNABUNDUS A,, A,, (1934) and A,, (1937, 1938) (Loose) S. W. and B., comb. nov. Sporonites indignnbundzts 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, pl. 19, fig. 51. the genus. Possibly the A, 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. C, (Knox, 1912) possibly might be placed Apiculati-sporites indigna h und us (Loose) Loose, 1934, Inst. Palaobot. Arb., here though its morphologic interpi-eta- vol. 4, no. 3, p. 153. tion is somewhat uncertain clue to its un- usual type of equatorial developnlent. 5. DENSO-SPORITESLORICATUS (Loose) S. W. Remavks.-Six species are now included and B., comb. nov. Sporonites lo~%atusLoose, 1932, Neues under Demo-sfiorites and it seems that a Jahrb. Beilage-Band 67, Abt. B, p. 450, pl. good many more will need be distin- 18, fig. 42. guished. Raistrick's A,, A,, and A,, Zonales-sporifes loricatus (Loose) types are woi-thy of specific clistinction. Loose, 1934, Inst. Palaobot. Arb., vol. 4, The generic name, while perhaps of un- no. 3, p. 151. desiralsle 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. As mentioned previously, the spores Note.-A very problematic form. are widespread and vesy abundant in splint coals. It has been suggested that Genus CYSTOSPORITESSchopf, 1938 a special type of vegetation was respon- sible for this type of coal but such can Plate 1, figures 10-10h hardly be the case, since 1-ecognizable splint Sy.r.cz~~aetry.-Spores radial, trilete. coals are also found that are so much Forvn and sixe.-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 metes and about half as broad. Abortive of plants is responsible for forming this forms variable in development, cliarac- type of coal. The coincidence of occur- teristically ranging from as small as one- rence between Denso-sporites and Pale- loSince Loose recognized the rather significant distinc- ozoic splint coal nevestheless seenls tion between Demo-.rporite.r and the zonate forms, In a footnote appended to his article he suggested "einer Unter- unaccountably high and additional inf or- abteilung Annulati-~porites" for D. dnnulatus and D. lorrcatus. Because he did not adopt the name, "Unter- e Thiessen has also designated some spores not referable abteilung Annrrlaii-sporites" is invaljdly published, at to Denso-spor~tesas "splint spores." least in the sense of a generic designatlon. 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 isodiainetric becoming axially elon- cerned. The ornaniental patterns which gate with increased size. When compressed serve to distinmish snores of free-snor- both abortive and fertile members are ing lycopsids vhave e;idently 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 Hnptotypic f eatz~res.-Strongly devel- in the Lepidocarpaceae, and, so far as is oped in fertile forms but relatively in- known, is inclusively correlative with the conspicuous because of the enormous ex- lepidocarp family. Future discoveries may pansion of the distal part of the spore make it possible to restrict this group and body. Sutures definite, extended to the establish relationships in greater detail. arcuate ridges which are strongly thick- The spores found within the seeds of ened, lips often moderately defined ; py- lepidocarp species are receiving more de- ramic areas distinct from adjacent spore tailed study now (Reed, 1941 ; Darrah. coat. Apex may be somewhat elongate. 1941 ; ~oskinsand' Cross, 1941 ; ~chopf On smaller abortive forms haptotypic 1941) and fui-ther studies may show f eatures are commonly masked by develop- other features of the spore coat that have ment of a thick rugose, more or less unsuspected systematic significance. 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 clistiiictioii between are weakly defined adjoining the corners f ree-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- niost 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 heteroniorphous. 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 hollotypes 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 moral 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- Afi1zites.-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, Itli- a means of more accurate recording of uziocarpon, 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 in the Carbonifero~lsas is best shown by generally do not modify the profile unless 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. CYSTOSPORITESBRERETONENSIS Schopf, 1938, Omamentation. - Body wall may be Illinois Geol. Survey Rept. Inv. 50, pp. rugose, bladder membrane lightly sculp- 40-42, pl. 3, fig. 5, pl. 8, figs. 1-4. tured. Triletes cf. T. giganteus Zerndt, Schopf, 1936, Illinois Acad. Sci. Trans. vol. 28, Hapto typic f eatures.-Relatively incon- 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 CYSTOSPORITESBRERETONENSIS f orma ABOR- area may sometimes be distinguished ; TIVUS Schopf, 1938, Illinois Geol. Survey Rept. Inv. 50, p. 40, pl. 1, fig. 10, pl. 8, fig. 4. except in this area no arcuate marking is CYSTOSPORITESBRERETONENSIS forma RETIC- evident. ULATUS Schopf, 1938, idem, p. 40, pl. 1, fig. Spore coat.-Body wall (exospore) 11. tending to be dense and of moderate rela- 2. CYST~SPORITESGIGANTEUS (Zerndt) Schopf, tive thickness ; bladder membrane (peri- 1938, idem, p. 39. spore), relatively thin and quite trans- Triletes gigantezts Zerndt, 1930, Acad. lucent. polonaise sci. Bull. internat., ser. B, p. 71, pls. 9, 10, and 11. Afinity.-There is considerable prob- Type 1 Zerndt, 1931, Acad. polonaise ability that the plants represented by Para- sci. Bull. internat., ser. A, p. 170. spovites are gymnospermo~~s.The peri- Sporites giganteus (Zerndt) Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, sporal bladder development is strongest p. 172, pl. 8, fig. 9. evidence of this. It is still a question Triletes giganteus Zerndt, 1937, Acad. whether the genus should be assigned with polonaise sci. Trav. Geol. no. 3, p. 4. the Pteridosperms, Corclaitaleans or the CYSTOSPORITESGIGANTEUS f orma VARIUS Conifers (cf. Schopf, 1938, pp. 47-8). (Wicher) S. W. and B., comb. nov. T~riletes(Lagenicula) glabratus Zerndt, The genus has some evident claim to 1930, Acad. polonaise sci. Bull. internat., affinity with Floriltites, S. W. and B., ser. B., p. 54, pl. 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. 175, pl. 8, figs. 26-27. as these forms are contemporaneous their Type 30 Zerndt, 1932, Jahrb. fur das relationship cannot easily be regarded as Berg-u. Huttenwesen in Sachsen. Jahrg. direct. 1932, p. 13A, pl. 1, figs. 1, 2, 3. Re114ark.r.- The genus is n~onotypic Sporites varius' Wicher, 1934, Inst. Palaobot. Arb., vol. 4, no. 4, p. 89, pl. 6, being represented only by the species figs. 2, 3, 4, 6. given below which has been found in upper Carbondale and lower McLeans- Genus PARASPORITESSchopf, 1938 boro age coals of Illinois. The point upon which greatest emphasis should be placed Plate 2, figures 16-16b is that the development of proximal Sy'~mwetry.-Prepollen .grains appear- sutures is such as to indicate proximal ing bilateral due to opposing bladders. The gametophytic exit similar to E~zdosporites fundamental symmetry, clearly shown by and Mofioletes. Aside from this and its the body, is radial. rather large size Pnmspo~iteshas 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 I. PARASPORITESMACCABEI Schopf, 1938, Illi- rather coarsely serrate. nois Geol. Survey Rept. Inv. 50, pp. 48-9, Afinity.-Spores of Ciwatriradites are pl. 1, fig. 6, p1. 7, figs. 1-3. not yet definitely correlated with any major plant group. Their most likely affinity Genus CIRRATRIRADITESWilson and Coe, seems to be with the lycopods but known 1940 types of Lepidostrobus microspores are Plate 3, figures 21, 21a, 21b usually sn~allerand generally possess no Sy~awzet?py.-Spores trilete, radial. comparable development of the flange. Shape.-Moderately oblate spheroidal, Certain zonate types of T~riletes spores with a strongly projecting eqtlatorial bear a superficial resemblance to Ciwatri- 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 t~iradites 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 Size.-Spores of various species range supported only on theoretical grounds but from about 40 to over 100 microns in they could possibly be represented by overall diameter. Ciwat&adites. 0rnanzentatio1a.-Coiiin~only showing a Some atlthors have confused Ciwatri- pattern having its primary emphasis along radites spores with those of genera (En- radial lines ; niay consist of ridges more dospo&es, Spemedes, etc.) which pos- or less anastomosing to the point of be- sess perisporal bladder development in coming a reticulation, or surfaces may be the equatorial plane. Such resemblance nearly free of ornamental ridges and as there is, is certainly only superficial. most of the surface smooth, granulose or Raistrick's A;. A,, and C, (cf. Rai- finely punctate. Some species show a strick 1935, 1937, 1938) belong here; unique type of distal ornamentation con- likewise, possibly D, in part. As given sisting of one or several thinner areas with by Knox, (1938, 1942) Dl, is definitely rather prominent margins. The flange is referable to Cirratriradites. Types 1 and often more or less radially striated and in 9 of Millott (1939) without much clues- addition may develop one or two concen- tion belong in this genus. The snores illus- tric bands of irregular thickening. The trated by Reinsch (1884, p. 22) in plate pyramic areas are not particularly dis- 15, figs. la and lb as types 222 and 223 tinguished by oriiamentation. from Zwickau, Saxony, are characteristic Haptotypic features.-Trilete rays rela- of it. tively strongly developed and extended to Remarks. - Thirteen previously de- the equator, oftentimes a line of thick- scribed species are included in the list be- ening continues to the edge of the flange. low, all but one of which, C. wzaculatus Lips are frequently strongly demarcated the genotype, represent new name com- and raised above the spore body; the binations. 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. CIRRATRIRADITESARGUTUS ( Ibrahim) S. W. from the spore body, being thinner than and B., comb. nov. Zonales-sporites argutzts Ibrahim, 1933, the body wall and more translucent; the Sporenformen des Aegirhorizonts, pp. 31-32, interradial flange width is often some- pl. 6, fig. 55. PALEOZOIC FOSSIL SPORES

CIRRATRIRADITES( ? ) CICATRICOSUS (Ibra- Zonales-sporites saturni (Ibrahim) Ib- him) S. W. and B., comb. nov. rahim, Loose, 1934, Inst. Palaobot. Arb., Sfioronites cicatricosus Ibrahirn, 1932. vol. 4, no. 3, p. 149, pl. 7, fig. 23. Neue; Jahrb., Beilage-Band 67, ~bt:B, Note.-Ibrahim (1933) came to the conclusion 447, pl. 14, fig. 2. that Sporonites formosus Ibrahim (1932) was Zonales-sporites cicatricosus (Ibrahim) conspecific with C. saturni. Very little infor- Ibrahim, 1933, Sporenformen des Aegirhor- mation is provided to support this and, since izonts, p. 31, pl. 1, fig. 2. the figures appear to show specific differences, it appears desirable to keep them separate for a CIRRATRIRADITESFAUNUS (Ibrahim) S. W. time at least. and B., comb. nov. Sporonites faunzts Ibrahim, 1932, Neues 11. CIRRATRIRADITESTENUIS (Loose) S. W. and Jahrb., Beilage-Band 67, Abt. B, p. 447, B., comb. nov. pl. 14, fig. 4. Sporonites tenuis Loose, 1932, Neues Zonales-sporites faunus (Ibrahim) Ib- Jahrb., Beilage-Band 67, Abt. B., p. 450, rahim, 1933, Sporenformen des Aegirhori- pl. 18, fig. 34. zonts, p. 28, pl. 1, fig. 4. Zonales-sporites tenuis (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no 3, CIRRATRIADITESFORMOSUS ( Ibrahim) S. W. p. 149. and B., comb. nov. Sporonites formoszts Ibrahim, 1932, CIRRATRIRADITESVENUSTUS (Loose) S. W. Neues Jahrb., Beilage-Band 67, Abt. B, and B., comb. nov. p. 447, pl. 14, fig. 10. Sporonites venustus Loose, 1932 Neues Zonales-sporites saturni (Ibrahim) Jahrb., Beilage-Band 67, Abt. B., p. 450, Ibrahim, in part, 1933, Sporenformen des p1. 18, fig. 36. Aegirhorizonts, p. 30, pl. 1, fig. 10. Zonales-sborites z~enustus (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, CIRRATRIRADITES(?) GRACILIS (Zerndt) no. 3, p. 149. S. W. and B., comb. nov. Triletes gracilis Zerndt, 1937, Acad. 13. CIRRATRIRADITESZONALIS (Loose) S. W. polonaise sci. Bull. internat., ser. A., p. 586, and B., comb. nov. pl. 12, figs. 1-10. Zoneles-sporites zonalis Loose, 1934, Type 46 Zerndt, 1937 (idem). idem, vol. 4, no. 3, p. 148, pl. 7, fig. 5. Note-Species exceptionally large (300 P), otherwise in close generic agreement. Genus ENDOSPORITESWilson and Coe, 6. CIRRATRIRADITESMACULATUS Wilson and 1940 Coe, 1940, Am. Midland Naturalist, vol. Plate 2, figures 14-14b 23, no. 1, p. 183, fig. 7. Note.-Reexamination of the type material Sywawaetry.-Spores trilete, radial. 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- turing simulating it may occur distally. to oval or elliptical or slightly triangular; body wall more spherical than the bladder. 7. CIRRATRIRADITES( ? ) PEACOCKI (Berry) S. When compressed the spores most com- W. and B., comb. nov. monly show good proximo-distal orienta- Zonales-sporites peacocki Berry, 1937, idem, vol. 18, p. 156, fig. 5. tion. The spore body is not generally folded but the bladder membrane more 8. CIRRATRIRADITESPENNINGTONENSIS (Ber- commonly shows irregular plication. The ry) S. W. and B., comb. nov. bladder profile at the margin of the spore Zonales-sporites penningtonensis Berry, corresponcls to the profile of folds, being 1937, idem, p. 156, fig. 3. straight or evenly rounded, and thus 9. CIRRATRIRADITESRARUS (Ibrahim) S. W. easily distinguishable from simple flanges and B., comb. nov. of zonate forms that superficially re- Zonales-sporites rarus Ibrahim, 1933, semble Eutdosporites. Sporenformen des Aegirhorizonts, p. 29, pl. 6, fig. 53. Sixe.-Ranging in various species from about 50 to possibly 300 microns total 10. CIRRATRIRADITESSATURNI (Ibrahim) S. W. and B., Comb. nov. diameter. The spore body generally is Sporonites saturni Ibrahim, 1932, less than one-half of the full diameter. Neues Jahrb., Beilage-Band 67, Abt. B., p. Omamen tation. - Surfaces of bladder 448, pl. 15, fig. 14. are levigate to granular or punctate; a Zonales-sborites saturni (Ibrahim) Ib- fine meshed reticulation pattern is com- rahim, in part, 1933, o or en form en' des Aegirhorizonts, p. 30, pl. 2, fig. 14 (non pl. monly present but this is probably on the 1, fig. 10). imzside, rather than on the external bladder ENDOSPORITES 45 surface. The spore body is less definitely Parasporites, Cordaialzthz~s, Alisporit es, ornamented although the bladder pattern Caytonanthus, Pityosporites and Endo- may appear superimposed on it. Emphytic spo~ites,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- Haptotypic f eatures.-Trilete rays com- mologous structures, which may have un- monly extend to the periphery of the dergone progressive and segressive 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 membrane. The lips are oftentimes up- nevertheless largely preserve their 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 Spore coat.-The bladder (perisporal) evidently from the proximal pole as in membrane is quite thin and translucent; the cryptogams. Florin, working with its reticulation tends to add somewhat to excellently preserved silicified material, the thickness as seen in marginal. profile. has shown that in some late Pennsylvaliian The body wall is substantially thicker al- (Steplianian) forms the haptotypic f ea- though also quite translucent, and cor- tures were vestigial and in mature forms responds to the exosporal layer. In a present only as a surface imprint on the few instances thin membranous, almost bladder membrane which does not even hyaline, endosporal membranes have been maintain contact with the spore body. observed within the exospore and shrunk- The bladder and body are in contact dis- en from it. The suture lines are not dis- tally, however, and prothallial cells line tinct on these, but three clear-cut endo- the body cavity except at this point. Thus sporal apical papillae occur at the juncture in these advanced forms germinal exit of pyramic apices. must have been distal as it is in pollen of all modern conifers. Pollen-like types of Afilzities.-Elzdosporites 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). Endospovites male abundant association with Cordaitean spores are evidently prepollen in this leaves, other plant fossils being infrequent, sense. that no other conclusion seen~spermis- Eight pseviously 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 Cordnianthus fructifications America and in Europe. The genotype obtained from the French Stephanian are species, E. or~zatusWilson and Coe, is evidently generically distinct and of more from Iowa coal of Des Moines age. advanced structure. ENDOSPORITESANGULATUS Wilson and Coe, Raistrick's types C, and C, without 1940, Am. Midland Naturalist, vol. 23, no. much question belong to Endosporites 1, p. 184, fig. 1. (Raistrick and Simpson, 1933 ; Knox, ENDOSPORITESGLOBIFORMIS (Ibrahim) S. 1938). Type 629 of Reinsch (1884, p. 61, W. and B., comb. nov. pl. 48, fig. 252A) fro& the Blatterkohle Sfioro~zites globifornzis Ibrahim, 1932, and Stigmarienkohle of Metschowk (in Neues Jahrb., Beilage-Band 67, Abt. B, p. central Russia) and probably others less 447, pl. 14, fig. 5. easy to interpret from his drawings, also Zonales-sporites globiformis (Ibrahim) belongs to Eyldosp o&es. Ibrahim, 1933, Sporenformen des Aegir- horizonts, p. 28, pl. 1, fig. 5. Re~izar1zs.-The distended bladder mem- Zonales-sporites globif ormis (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 EXDOSPORITES(?) KARCZEWSKII (Zerndt) in the several groups which are known as S. W. and B., comb. nov. 46 PALEOZOIC FOSSIL SPORES

Triletes karczewskii Zerndt, 1934, Ornamentation. - Surfaces levigate to Acad. polonaise sci. Trav. Geol. no. 1, p. granulose and mildly verrucose, angular 27, pl. 31, fig. 3. areas sometimes more highly ornamented. Note.-Agrees well with Endosporites though larger than most species. Emphytic ornamentation of proximal and distal surfaces is about the same. 4. ENDOSPORITESORNATUS Wilson and Coe, 1940, Am. Midland Naturalist, vol. 23, no. Haptotypic str.ttctztres.-Trilete rays ex- 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. ENDOSPORITESPELLUCIDUS 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, Iowa Acad. Sci., vol. 44, pp. 128-129, fig. 1. spore coat at the angles may simulate a partial flange. 6. ENDOSPORITESROTUNDUS (Ibrahim) S. W. and B., comb. nov. Spore coat.-Characterized by gross in- Zonales-sporites rotundus Ibrahim, equalities in exospore thickness ; spore 1933, Sporenformen des Aegirhorizonts, p. coat is thickest on the angles opposite ends 31, pl. 8, fig. 73. of the rays and thinnest in the central 7. ENDOSPORITESRUGATUS (Ibrahim) S. W. distal and possibly to a somewhat lesser and B., comb. nov. extent on the interradial areas. Actual Zondes-sporites rugatus Ibrahim, 1933, thickness of different parts of the wall idem, p. 31, pl. 8, fig. 70. in various species varies greatly although 8. ENDOSPORITESVOLANS (Loose) S. W. and the relations of thicker and thinner areas B., comb. nov. of the spore coat is rather constant. Sporomites volans Loose, 1932, Neues Jahrb., Beilage-Band 67, Abt. B, p. 451, Afinity. - Trz'quitrites cannot be as- p1. 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, p. 36. the genus is considered to correspond with Zomdes-sflorites volans (Loose) Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, natural plant relationship. The features, in 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 Dl, of Rai- Genus TRIQUITRITESWilson and Coe, strick (1938). Type D, (Raisti-ick, 1934, 1940 1935, 1938) shows more extreme thicken- Plate 3, figures 20-20b ing of the 1-aclial angles of the spore coat so that it is evidently closely related to S~i~~z~~.zet?ry.-Sporestrilete ; radial. this genus, though it is a question whether Form-Oval to elliptical in vertical it should be classified as Triqztitrites 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 diagnoses, nos. 345, 347, 319, 353, seldom induced by compression due to 354, 362, 363, 365, 373. Reinsch also good preferential orientation coinciding illustrated other forn~swhich are doubt- with the horizontal plane of the spores ; less congeneric with Triquitrites in his forms more inflate distally are indicated types 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 sporqcoat on the niscent of some, possibly overmacerated, angles. specimens of the genotype species. Size.-Spores of various species com- Re~~zarks.-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 EQ UISETOSPORITES 47

known from American coals and will sub- Sporonites triturgidz~s Loose, 1932, sequently be described. The numerous Neues Jahrb., Beilage-Band 67, Abt. B, p. 449, pl. 18, fig. 32. spores illustrated by Reinsch (1884) in the Valvisi-sflorites triturgidus (Loose) Micro-Paleophyto2ogia 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 derived from localities in Russia where Genus EQUISETOSPORITESDaugherty, the coals probably are of Lower Carbonif- 1941 erous age. D,, and D, types of Raistrick This genus recently described from Tri- ( 1938) were chiefly obtained f rom 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 validity of the name relationship thus seems definitely indi- E quisetosporites 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 Equisetzti/lz. 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 simplification in spore coat structure of of the Triassic have proven to be quite dif- ferent when the record is complete enough late Carboniferous plants as contrasted with ealier members of the same alliance to allow specific determination." Thus until further evidence is available, it has also been noted in n~embersof the would seem unwise to emphasize the mod- Lage.tzicula and Aplzanoxonateae sections ern aspect of this fossil form. Its dis-@ of Triletes. covery nevertheless adds an important 1. TRIQUITRITESARCULATUS Wilson and Coe, item of information to the record. 1940, Am. Midland Naturalist, vol. 23, no. 1, p. 185, fig. 8. Knox examined spores of fifteen species Nofe.-Type material bears a moderate to of Equisetu~ozand reports ( 1938, pp. 439- 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 TRIQUITRITESSPINOSUS Kosanlte, 1943, Am. the spore wall was faintly granular. Ela- Midland Naturalist, vol. 29, no. 1, p. 128. ters of somewhat varied character are pl. 3, figs. 2, 2a, 2b. present on all modern species of Equi- TRIQUITRITESTRIBULLATUS (Ibrahim) S. setum. In commenting on Raistrick's B, W, and B., comb. nov. type, which she recognizes is very prob- Spo~,onifes tribullatus Ibrahim, 1932, ably of calamarian affinity, Knox remarks Neues Jahrb., Beilage-Band 67, Abt. B, (pp. 461-3) on p. 448, pl. 15, fig. 13. Laevigati-sporites tribullatz~s(Ibrahim) the same globose form with thin wall and Ibrahim, 1933, Sporenformen des Aegirhor- absence of ornamentation . . . the unusual de- izonts, pp. 20-21, pl. 2, fig. 13. velopment of elaters and no trace of the tri- radiate mark. Valvisi-sflorites t~~ibullatus(Ibrahim) Loose, 1934, Inst. Palaobot. Arb., vol. 4. She cites Halle's description of Rhaetic no. 3, p. 152, pl. 7, fig. 21. and Triassic - age Equisetum spores TRIQUITRITESTRIGONAPPENDIX (Loose) S. (Halle, 1908) in which the trilete com- W. and B., comb. nov. missure is clearly seen but which show Valvisi-sflorites t~-igonappe~zdixLoose, no trace of elaters. Halle assumed that 1934, idem, p. 152, pl. 7, fig. 17. even if elaters had been present they TRIQUITRITESTRITURGIDUS (Loose) S. W. would have been removed by maceration and B., comb. nov. procedure. Needless to say, this aspect 48 PALEOZOIC FOSSIL SPORES

must always be considered. Nevertheless, 1. EQUISETOSPORITESCHINLEANA Daugherty, the spores recovered by Halle were typi- 1941, Carnegie Inst. Washington Pub. 526, Age, Triassic. cal of calamarians (cf. Hartung, 1933) p. 63, pl. 34, fig. 4. and isolated spores of this character must be classed with Cnlamospora n. gen. as de- Genus ALISPORITESDaugherty, 1941 scribed below. Equisetosporites chi~zleana 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 completelv the lating it to Parasporites and Pityosporites. relationship and distribution of ecluiseta- The only description is that of the geno- lean types. type species, A. opG, given below (Dau- The type of the single species of Equi- gherty 1941, p. 98). 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,ll 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 ref erring quately illustrated by Daugherty in the them to Caytonantlzus. Harris ( 1941) figure cited below. has distinguished Caytonantlzus kochi, C. Sy~w/~/zetq.-No trace of original tet- oncoides, and C. arberi largely on the ba- sis of differences in their pollen and has " rad configuration visible. suggested their correlation with species Sha;be.-Body spherical (type speci- of Caytonid 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- O?~nn~~ze~ztatio~z.-Bod~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 Cag1tonanthz.t~should be iden- E1aters.-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. ~lat&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 11 The terms dorsal and ventral tend to be ambiguous in description of pollen grains, particularly unless some fur- spirally encircle the spore body but they ther ex lanation is given Probably "dorsal" as used have been displaced on the specimen at here reRrs to the proximai side, i.e., the side which was internal and adjoined the other members of the original hand. tetrad. CALAMOSPORA 49 teration, the important point relates to 2. ALISPORITESspp. ? recognition of the degree of change that Pollen Forms I, 11, and 111, Luck, is manifested and of the fact that bio- 1913, Beitrag zur Kenntnis des alteren 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- tifically trustworthy these principles Genus CALAMOSPORAS. W. and B., gen. should be more widely acknowledged. nov. Alisporites is a designation that should Plate 3, figures 22-2213 ; text figure 1 be very useful in reporting the occurrence Sy.i~twzetry.-Spores trilete; radial. of caytonian spores and others of the same sort which perhaps may not even Shape.-Spherical or nearly so ; when be necessarily referable to that group. compressed, readjustment to a disk-like Whether the Caytoniales are worthy of form leads to formation of characteristic ordinal distinction is perhaps subject to sharp taper-point folds of variously cres- some question now that Harris (1941, centic or narrowly lenticular outline. Some- times the spores are folded double so that 1940) has shown SO convincingly that they are gymnospermotls and have no the whole external outline is sharply len- particular connection with angiosperms. ticular. Such folds are one of the char- Likewise whether they should be termed acteristic features of the genus. pteridosperms is still perhaps too much Sizs.-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 other gymnospermae is a perplexing one in diameter. which may find its truest solution in the 0rulmaentation.-Spores are character- exacting study of fossil pollen grains istically very smooth in general appear- similar to Alisporites, Parasp orites, and ance; on closer inspection they may be Pityosporites. minutely granulose or slightly rugose. The more adequate generic definition There may be very slight differentiation of AZispora'tes should be taken up in con- of proximal pyramic areas shown by this nection with such studies. Daugherty's almost negligible emphytic marking. Larg- generic distinction of these interesting er spores often possess a high gloss when plant microf ossils is a forward step which observed by reflected light. 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 f roin spores of Parasflorites 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 seen1 possible to recognize morphologic somewhat diff ereiit surf ace 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 des 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 Lebaclzia and Walchiantlzus, 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 Alisporztes 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. ALISPO~TESOPII Daugherty, 1941, Carnegie Inst. Washington Pub. 526, p. 98, pl. 34, eter, to less than 2 microns in those fig. 2. 1 smaller than 100 microns. A very thin PALEOZOIC FOSSIL SPORES

endosporal membrane may be present, referable to this genus but probably not oftentimes shrunken away from the ex- all of them should be classed here. There osporal coat. are a few forms of Pzcutctati-sporites that Afinity .-Hartung's ( 1933) investiga- are very similar in definable features but tion of spores in fructifications that were nevertheless show differences- of habit that set them apart from Calamosjora. referred to six different genera allied with Some of the B, types Raistrick .has illus- the calamarians has provided an excellent trated have trilete rays more extended basis for systematic treatment of plants than they generally are in spores of Cola- of this sort which are only represented by 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 theniselves in his type B,. Miss Knox (1938, p. to grouping into restricted generic alli- 461) has remarked on the affinity the B, ances but show a rather generalized char- spores and compared them with spores acter. Certain species niay easily be of Calamostachys binneana and Cheiro- distinguished by spore differences but strobus pettycurhsis. generic characteristics (in terms of the Remarks. - Calarnosporn 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 Noegge?~atlziostro- 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 althdugh the degree of relation- group of Annularia stsllata 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 nien- obtained from beds of lower Des Moines tioned and probably is also correlative age. Such a view also was expressed by with some Mesozoic equisetalealis (cf. 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 Carfiautnularia 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 niicrons be regarded as established. in diameter to his Type 2. Several dis- One feature stands out in the sporoge- tinct species' of Cdamospora 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 normai 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 B, with abortive forms in various stages of spores of Raistrick (1933, 1934, etc.) are development. Sporangial masses com- CALAMOSPORA posed of abortive and fertile spores have also been observed in coal maceration residues (Schopf, 1938, p. 51). Most noteworthy is the fact that large spores exceeding half a millimeter in diameter show characters very similar to those of less than 100 microns in diameter. Al- though heterospory was well established, a seemingly continuous size gradation ex- ists between the largest niembers and the smallest with little but relative size as a FIG. l-Calamospora, hartungiana sp. nov., draw- distinguishing feature. Spores in f ructi- ing from microproj ection of holotype. fications which are less than about 90 microns in diameter frequently are re- garded as microspores and those that ex- 1. CALAMOSPORAHARTUNGIANA Schopf, sp. nov. ceed that size often appear to represent Text figure 1. megaspores. But in different species and in general there seems to be no exact Desc~iption.-Spores spherical, compressed to polygonal ' or acutely lenticular out- way to distinguish between them unless line, generally with several more or less lunate t_he 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 spofraprobably include spores of one mor- distinct and slightly undulating, lips low but 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 under two specific names. This is no pyramic areas are slightly thickgr and darker by transmitted light than the resf of the spore serious disadvantage if we are interested coat and have slightly coarser surface texture . in obtaining the most precise information (kontakthof). Spore coat thin, pne micron or that such material can provide. If rec- less, generally yellowish translu~entor some- what red when light is transditted 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 lation of large and small spore forms in rugose to granular; at low magnification the surface appears quite smooth. some instances. It is doubtful that the _ Calarnosfiora 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- Six species previously described and ans range from 60-150 microns in diameter, 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 Spores of this generic character are com- calamarian fructifications in order to be cer- tain of this, but there can be no doubt as to the mon in many American coals and this general affinity of C. hartztngiana. 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 R'ver northwest ana, one of the characteristic American of Fairmount, Vermilion County, klinois. spores of this sort occur in spore masses 2 mm. or forms, to serve as a genotype of CaJa~no- 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 oth~rspecimens 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 FO-SSIL SPORES

obviously are only incomplete contents of single Genus REINSCHOSPORAS. W. and B., sporangia) gives a little additional information gen. nov. on the Calamospora fructification. The holotype is from slide Y of maceration Plate 2, figures 11-llb ; text fig. 2 90, deposited in the Illinois Geological Sur~ey This genus appears to include the spores collections, Urbana. Reinsch (1884) designated as his Sub- 2. CALAUOSPORALAEVIGATUS (Ibrahim) S. W. divisio I11 of his Subtribus I, Nucleus and B., comb. nov. triangularis. The genus is therefore ap- Laevigati-sporites laevigatus Ibrahim, propriately designated in honor of 1933, Sporenformen des Aegirhorizonts, pp. Reinsch. His description (loc. cit. p. 7) 17-18, pl. 4, fig. 46. of the subdivision is as follows (free Lae-digati-sporites laevigatus Ibrahim, : Loose, 1934, Inst. Palaobot. Arb., vol 4, translation from Latin) no. 3, p. 146, pl. 7, fig. 36. Exospore thin (1/24-1/40 of the transverse (Ca1amiti)-sporites laevigatus (Ibra- diameter), smooth or ornamented with granules 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 dentif orm juxtaposed spines, gradually decreasing (in length) from the cen- 3. CALAMOSPORAMICRORUGOSUS (Ibrahim) S. tral margins toward the angles. W. and B., comb. nov. S3oronites microrugosus Ibrahim, 1932, This diagnosis has no taxonomic status Neues Jahrb., Beilage-Band 67, Abt. B., although it (and others given by Reinsch) p. 447, pl. 14, fig. 9. may be quite worthy of validation. One Laevigati-spbrites wzicrorugosus (Ibra- naturally hesitates to apply names on the him) Ibrahim, 1933, Sporenf ormen des basis of Reinsch's descriptions and fig- Aegirhorizonts, etc., p. 18, pl. 1, fig. 9. ures because of uncertainty as to the 4. CALAMOSPORAMUTABILIS (Loose) S. W. sources and present location of the types. and R., comb. nov. Also, modern interpretations may vary Calawziti ( ? ) -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, pl. 19, figs. 50a-c. The point which should be emphasized is Calawiti ( ?)-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. 3, p. 145. form variation, particularly when essen- 5. CALAMOSPORA(?) OBESUS (Loose) S. W. tial confirmation can be obtained from and B., comb. nov. additional material recently prepared.

Sboronites obesus Loose. 1932. Neues a= Thus the forms Reinsch described in his Jahrb, Beilage-Band 67, ~bt.'B., p'451, pl. subdivision I11 are structurally clarified - 19, fig. 49. by study of American forms of Reinschlo- Laevigati-sporites obesus (Loose) Ib- spura,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-sflorites obesus (Loose) Ib- no particular occasion to propose names rahim, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 145. for these until similar new material is at hand or the source defined for his old 6. CALAMOSPORAPALLIDUS (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 pl. 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. Pzmctati-sporites pallidus (Loose) Ib- Shape.-Body lenticular to flattened el- i rahim, Loose, 1934, Inst. Palaobot. Arb., liptical in the axial plane ; subtriangular vol. 4, no. 3, p. 146. in transverse plane, corners rounded, sides 7. CALAMOSPORAPERRUGOSUS (Loose) S, W. slightly convex or concave in outline. A and B., comb. nov. fimbriate flange, broadest in the inter- Laeva'gati-sporites Perrugosus Loose, radial region and very narrow or absent 1934, idem, p. 145, pl. 7, fig. 13. on the corners, may be present to modify REINSCHOSPORA the marginal outline. In some forms the interradial appendages may be dissected into a row of spines, their ends similar to the flange in marginal contour. When compressed, folds are relatively inf re- 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 bellitas sp. nov., drawing preted as equivalent to an arcuate ridge of holotype. since, contrary to usual arcuate ridge de- can be identified with Botryo~occus~ velopnient, 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. REINSCHOSPORABELLITAS Bentall, sp. nov. arcuate ridges in mode of formation. Text figure 2 Spore coat.-Moderately thin (less than Desrriptio~z.-Compressed spore subtriangular 3 microns) and of uniform thickness ex- in outline with inter-radial margins usually con- cept at the equator where it is joined by cave. Diameters vary from 57 to 76 p, with an the flange. average of about 68 p. Spore coat less than 2 p in thickness and levigate. Trilete rays extend Afi9zities.-Unknown ; possibly filicin- four-fifths or more of the distance to the ean.""The similarities in morphofogy sug- equator. Flange usually originates slightly gest that this is a group having natural proximal from the spore equator. Fimbriate significance. It may be related to Gra?zu- elements of flange number about 50 between ray extremities and have a maximum length of 28 p midway between rays and a minimum Remwks.-The genus is reported from length of four to five at the ends of the rays. central Russia by Reinsch (1884, his di- Incisions of flange, when present, apparently agnoses 62-66, incl.), from the Ruhr by due to imperfect preservation. Reinschospora bellitas has been isolated from Loose ( 1934), and f rorn Lower Pennsyl- Angel and Battle Creek seams of the southern vanian coals of Tennessee by Bentall. Tennessee coal field. Though never numerically Dolgner (1932, pl. 28, fig. 4) has illus- ahundant, it is present in nearly all samples. trated a form with marginal spines, from It is very similar to R. speciosa (Loose) but is somewhat larger in size. Occurring with R. Moscow brown coal, which is congeneric bellitas is a form essentially comparable in all with our interpretation of Reinsclzospora; characters except that the flange is absent. Brokaw has recently found somewhat The relationship, if any, between these forms is similar forms in the coal of the Bogota not known. cyclothem (upper McLeansboro age) in Holotype-text figure 2. Tennessee Division of Geology collection; Battle Creek seam, north the Pennsylvanian of Illinois. 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. REIK-SCHOSPORASPECIOSA (Loose) S. W. ate inasmuch as Blackburn and Temperley and B., comb. nov. Alnti-sporites speciosz~s Loose, 1934, (1936) have shown that the algae for- Inst. Palaobot. Arb., vol. 4, no. 3, p. 151, merly known as Reimchia probably +now pl. 7,'fig. 1. 54 PALEOZOIC FOSSII; SPORES

Genus LYCOSPORAS. IV. and B., gen. nov. nized the relationship of these types to Plate 3, figures 19-19b Lepidostrobus and compared D, and D, with Lepidostrobus jacksovli and L. old- Symww&y.-Spores radial ; trilete. havzius respectively. Type Dl, 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 Lepidostro bus from isolated microspores transverse plane. Compression is gener- alone. ~~icrofossilsof 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 Or$zn.~?aeultat.Zon.--Spores nearly their good preservation than spores do. smooth ; minutely granuiose or rugose. The spores probably constitute a more Sometimes the minute rugae tend to be compete record of the various closely re- more prominent radially. lated forms. whether they can be distin- guished easily from one another or not. Hapto typic f eatztres.-Trilete rays ex- Reinsch (1884) has illustrated a num- tended nearly to margin, suture lines dis- ber of forms, chiefly from central Russia tinct, usually without noteworthy differ- entiation of lips. In species with most and from Zwickau, Saxony, which are probably referable to Lycospora. His strongly developed emphytic ornament, numbered diagnoses 32, 81, 83, 87, 240, the haptotypic features also are more 424 ( ?),525 ( ?) and 550 may be cited. prominent. The equatorial ridge corre- 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 Lycospom ~taicropapillata (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 beenrecognized but are not yet described. Spore coat.-Relatively thin, sometimes less than 1 micron thick on proximal and 1. LYCOSPORAAIICROPAPILLATUS (Wils011 and distal surfaces and quite translucent. Coe) S. W. and B., comb. nov. Thicker at the equator where there is no Cirratri~*aditesmic~~opnpillntz~s Wilson and Coe, 1940, Am. Midland Naturalist, easily defined line of demarcation between vol. 23, no. 1, p. 184, fig. 6. the equatorial ridge and the body wall. 2. LYCOSPORAMINUTUS (Wilson and Coe) Afinity .-Spores of this type have f re- S. W. and B., comb. nov. quently been encountered in the tips of Ciwatri~*aditcs mi~zutlu Wilson and Lepidost~~obuscones and there is little Coe, 1940, idem, p. 183, figs. 11 and 12. doubt that most of these forms found iso- lated represent niicrospores of the arbor- 3. LYCOSPORAPELLUCIDUS (Wicher) S. W. eous lepidodendrids. Andrews and Pan- and B., comb. nov. Sporites ~ellz~cidz~sWicher, 1934, Inst. nell (1942) have described similar forms Palaobot. Arb., vol. 4, no. 3, p. 186, pl. which persist in tetrad groupings as mi- 8, fig. 29. crospores of a species of Lepidocwpon. 4. LYCOSPORAPUSILLUS (Ibrahim) S. W. and The microspores of sigillarians (as shown B., comb. nov. by Maxocarpon) are significantly larger Sposodes pz~sillz~sIbrahim, 1932, Neues and probably are not included in Lyco- Jahrb., Beilage-Band 67, Abt. B., p. 448, sporn (cf. Schopf, 1941) . pl. 15, fig. 20. Spores assigned to type Dl by Raistrick Zonales-sfiorites pusillus (Ibrahim) Ib- rahim, 1933, Sporenformen des Aegirhor- belong without question to Lycospora, izonts, p. 32, pl. 2, fig. 20. also probably his B, and D, Type in part Gra~zulati-sporites pusillz~s ( Ibrahim ) (Raistrick and Simpson, 1933, Raistrick, Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 1937, 1938). Knox (1938) has recog- 3, p. 146, pl. 7, fig. 3. RAISTRICKIA 55

Genus RAISTRICKIAS. W. and B., ever, is somewhat cluestionable. Forms gen. nov: Knox (1938) illustrates as D, and D, probably belong in this genus. Reinsch Plate 3, figures 18-18b ; text figure 3 (1884) has illustrated only threel forms which belong to this group without much Sy?gwv,etry.-Spores radial ; trilete. doubt. These are described under his Shape.-Sperical to slightly flattened in diagnoses 52, 308, and 588 from central transverse plane and slightly triangular. Russia and from Zwickau, Saxony. The Little preference as to plane is shown by form illustrated by Thiessen (1920) in compressed specimens. plate 61, figure B, from Benton, Illinois Size.-Spores of different species vary (probably the Herrin No. 6 coal), is from about 40 to 90 microns in mean di- characteristic of Raistrickia; that shown amet er . above it in horizontal thin section may 0r.izawzentation.-Characteristically ver- likewise belong here but it is impossible rucose or spinose; spines when present to be sure. The two illustrations show 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 Haptotypic f eatures.-Trilete rays va- form, which particularly seems generic- riable in length as in Punctati-sporites; ally distinct from Punctati-sporites, has generally inconspicuous, lips unorna- not previously been formally designated mented. Rays sometimes sl~ghtlyundu- although it seems fairly common. Fur- lant due to proximity of spine bases; thermore, inadequacies in the previous arcuate distinctions absent. descriptions and illustrations also mili- tate against selection of any of them as Spore coat. - Usually of moderate types -for this genus. he species de- thickness (two to six microns) perisporal scribed below, Raistrickia grovemis sp. membrane lacking. nov. is therefore designated as the type. Afi7zity.-Typical forms of Raistrickia RAISTRICXIA(?) ABDITUS (Loose) S. W. are without doubt filicinean. Spores of and B., comb. nov. Seutftenbergia pluwmosa, as identified by Sporonites abditz~sLoose, 1932, Neues Radforth (1938), agree with these of Jahrb., Beilage-Band 67, Abt. B., p. 451, Raistrickia, and similar schizaeaceous pl. 19, fig. 53. Verrucoxi-sporites abditus (Loose) sporangia obtained by Andrews and Loose 1934, Inst. Palaobot. Arb., vol. 4, no. Schopf from shale above Illinois coal No. 3, p. 154. 6 also contain spores of this type. HOW- RAISTRICKIAFIBRATUS (Loose) S. W. and ever, the spores of a form identified as S. B., comb. nov. pennaef omis by Radf orth (1939) show Sporonites fibratus Loose, 1932, Neues rather diffei-ent features, and those of Jahrb., Beilage-Band 67, Abt. B., p. 451, pl. S. sturi do not appear congeneric with 19, fig. 52. Setosi-spof4tes fibratus (Loose) Loose, Raist&kia. 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- RAISTRICKIAGROVENSIS Schopf, sp. nov. relation and Senf tenbergia and Raistrickia no doubt represent generic groups of Text figure 3 somewhat different circumscription. Description.-Spore rounded triangular in out- line, about 50 microns in diameter. Orna- Raistrick's types E, and E, (Raistrick mented with numerous short (3-5 p) broad and Simpson, 1933 ; Raistrick, 1934, 1935, (2-6 p) blunt tipped spines, which by juxtapo- sition sometimes appear to be mutually attached. 1937) belong in Raistrickia. His E, type Rays 15-20 p long, slightly undulating, lips not is particularly characteristic. That rep- 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 p) in thickness, appearing brownish Setosi-sporites superbus Ibrahim, 1933, translucent ; spines are darker. Sporenformen des Aegirhorizonts, p. 27, pl. 5, fig. 42.

Genus FLORINITESS. 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 and 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 affected by folding; the body generally FIG. 3-Raistrickia grovensis sp. nov., camera is marked by numerous sharp angular lucidi drawing of holotype. folds, especially around its periphery. This form, in general, resembles the spores Size.-Greatest diameter (length of of Radforth's (1938) Senftenbergia plumosa but 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 outline. body proper varies from about 20 to 110 It is difficult to make a precise comparison with other previously described species now assigned microns in diameter in individuals of vari- to Raistrickia because of inadequacies of their ous species. description and illustration. The spine tips of Ornaw~entation.- Bladder ornament this form are entire and rounded; others from similar to that of E~tdosporitesand Pity- the same coal bed and elsewhere show spine tips slightly dissected. No doubt several species osporites; exterior surf ace 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. 6) coal (up- which tends to be obsolescent in the cen- permost Carbondale age) which is worked by extensive open-cut mining operations near Mid- tral proximal area. 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. RAISTRICKIASAETOSUS (Loose) S. W. and B., comb. nov. reticulation. Sporo&es saetosus Loose, 1932, Neues Haptotypic features. - Generally not Jahrb., Beilage-Band 67, Abt., B., p. 452, evident, and probably always extremely pl. 19, fig. 56. obscure. Trilete imprint (when discern- Setosi-sporites saetosus (Loose) Ibra- able) is wholly vestigial, probably located him, 1933, Sporenf ormen des Aegirhori- zonts, p. 26. entirely on the bladder, and has no func- Setosi-sporites saetosus (Loose) Ibra- tional significance in the mature spore. 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. RAISTRICKIASPINOSOSAETOSUS (Loose) S. W. and B., comb. nov. Spore coat.-Bladder membrane is ex- Sfioro?tites spinososaetoszcs 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, pl. 19, fig. 55. brane generally has a very well defined Apiczclati-sfiorifes spinos osnet oszts reticulation network located on the inner (Loose) Ibrahim, 1933, Sporenformen des Aegirhorizonts, p. 24. surface. At the folded margin the blad- 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. RAISTRICKIASUPERBUS (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, of ten- 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 its tenuousness, it may not be exosporal 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-conif erous type s~orites. 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- rather numerous folds. istence. For this reason it is essential Afinity.-Florinites is known to be re- that.. . they be classified as precisely as pos- lated to Cordaianthus (pars), Lebachia, si ble. Ernestiodendron and Walchianthus Florinites is also related to Endospo- through the investigations of Florin rites, Pityosp o~ites,Alisp orit es, and Para- (1936, 1938-40) ; thus it seems evident sporites, all of which have the character- that it represents a part of the upper Pale- istic bladder-f orming 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 "Steplzan osperwzum" caryoid es 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, Renzarks.-This genus is proposed to and no doubt functional, trilete apparatus. apply to gymnospermic plants whose pol- Florin's study of Cordahnthus saportawus len generically corresponds with the type pollen and pollen grains of Flori.lzites species as expressed, in general, by the type associated with it (his Cordaianthus foregoing definition. Florin has given sp. I and 2) indicates that the suture was rather similar data in description of the entirely absent and the trilete marking species cccordaitiformi~J'which he places completely vestigial. In the walchian under the genus name Pollenites (see be- conifers not even a vestigial 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 Cordaiantlzus sp. 2) and still not be easily specific epithet. , demonstrable from codified compression Pollenites cordaitifor~nis was estab- material. The agreement in pollen struc- lished to include pollen which elsewhere ture between the Cordaiantlcus 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 systematically expressed cannot now be ently to t^he other- five species. Florin's stated. The pollen structure is so spe- photographs still serve as an excellent cialized, particularly when compared with basis for discussion of the character of the expression of spore characteristics isolated Florinites 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 Lebachia, 58 PALEOZOIC FOSSIL SPORES

Emestiodendron, or Walchjanthus 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. even Florinites, when a more precise 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 termination of fossils where pertinence 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 question. Thus it would seem that unless periphery; the bladder is oftentimes hardly folded when compression is in the proximo- one of the pollen specimens Florin has 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 Lebachia, Emestio- 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- distinct taxonomic species. As there is 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 auztiquus 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 lustrations have shown clearly that a the light straw-yellow translucence of the cen- tral area of the pollen grain. The "contact number of other species cap be distin- area" where bladder joins the body wall is guished by their pollen characters. elliptical or oval with its greater diameter Flo~initesis known to be widely dis- cor?monly transverse to the length of the pollen 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 associated with it. Pollen of this char- distal side up. A distinct but tenuous striation traverses the "contact area" in many instances, acter has been found to be abundant in continues on the bladder membrane until about sonie 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 develops its more dis- a detailed study of these forms will add tinctive ornamentation. It can hardly be in- terpreted as a monolete suture line; it would much to our knowledge of 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 Mo~zoletesprepollen but more faintly developed, since the body and bladder 1. FLORINITESANTIQUUS Schopf, sp. nov. membranes are not nearly so thick as the Text figures 4, 5 Monoletes coat. The striation seems correlated in position with the long axis of the contact Descri)tio~z.-Pollen grains, bilateral symmetry, area and frequently is set obliquely with ref- equipped with an annulate bladder joined to erence to the breadth of the grain as a whole. 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 he 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-polfen record that one would most log- ically turn to obtain earliest evidences of newly differentiating plant groups that became estab- lished among the upland vegetation of the period.

FIG. 4-Florinites antiqtms sp. nov., drawing from microproj ection of holotype. Bladder fairly common, there is a central gap or tear in the bladder membrane on the distal side cor- responding to the "contact area" and the central proximal region of the bladder wall is visible without any overlying layers, Granules seem aligned in a semblance of a central trilete pat- tern, obsolescent, with rays perhaps 6 microns long; the granulation of the surrounding area of the bladder membrane (it is too faint to be recognizable as reticulation) also seems faintly aligned to correspond to three radii. Conclusive definition is difficult in material obtained by coal maceration but such specimens tend to confirm Florin's (1936, p. 637) interpretation of prox- imal structure in the more highly reticulate and better preserved pollen identified by him as Cordaianthzts sp. 2. ladder Florinites antiqzms seems distinguishable from Body all the similar forms Florin (1936, 1938-40) has identified with Cordaiantlzus, Lebachia, Ernestio- BREADTH d.endron, and Walchianthzu, on the basis of its FIG. 5-Florinites a?ztiqzms, 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 from the illustrations he has published. His omitted). Cross-lined areas represent measure- Cordaianthzks sp. 1, from the Stephanian of ments from the holotype. France compares most closely in size, (bladder length, 78-90; width, 62-67 microns ; body length, 42-50; width 40-48 microns). The d~f? F. antiquz~shas been obtained in moderate ierences can best be evaluated by reference to abundance from a widespread 10 to 12-inch the assembled measurements of F. antiqztz~s coal in the Wiley cyclothem where it crops out presented in text figure 5. Presumably the along Soap Creek southeast of the town of flattening of F. antiquus pollen grains would Carbon, Davis County, Iowa. Correlation with give rise to dimensions which are slightly ex- the widespread Wiley coal of western Illinois cessive in the plane of compression and there- has recently been confirmed by L. M. Cline of fore not quite comparable to measurements Iowa State College, who has measured the taken from Florin's uncompressed forms. That section, showing this coal to be in proper se- the species are distinct is evident from the much quence and about 12 feet above the lower Sea- less pronounced bladder reticulation in F. an- horne coal and limestone developed in this area. tiqztus. The distal striation has not been noted The Wiley cyclothem is near the top of the in forms regarded as cordaitean but is evident Tradewater group of Illinois and is considered (though no mention is made of it) in several of to be lower Allegheny in age. Florin's .figures of Paleozoic coniferous pollen. The holotype of F. antiquus, illustrated in (cf. Florin, 1939, pls. 55-56, fig. 21; Florin, text figure 4, is from maceration 413, slide 8 1940, pls. 145-146, fig. 19; ibid, pls. 147-148, (unstained) in the Illinois State Geological fig. 5). Survey collections in Urbana. From its age (early Allegheny) one might assume that F. antiquzts belonged with the 2. FLORINITES(?) PUMICOSUS (Ibrahim) S. cordaitealeans, but its features are advanced on TV. and B., comb, nov. 60 PALEOZOIC FOSSIL SPORES

Sporonites pumicosus Ibrahim, 1932, 3. SPORITESHIRMERI Reissinger, 1939, Paleon- Neues Jahrb., Beilage-Band 67, Abt. B., tographica, vol. 84, Abt. B, p. 16. p. 447, pl. 14, fig. 6. Note.-Nomen nudem, no illustration. Reticulata-sporites pumicosus (Ibrahim, 1932) Ibrahim, 1933, Sporenformen des 4. SPORITES IRREGULARIS Reissinger, 1939, Aegirhorizonts, p. 38. Paleontographica, vol. 84, Abt. B, p. 17. Reticuluta-sporites pumicosus ( Ibrahim, Note.-Nomen nudum, no illustration. 1932) Ibrahim, 1933, Loose, 1934, Inst. 5. SPORITESPLICHTUS Schopf , 1938, IlJ,inois Palaobot. Arb., vol. 4, no. 3, p. 157, pl. 7, Geol. Survey Rept. Inv. 50, p. 51, pl. 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, Reticzclatn-sporites viseladus Ibrahim, Krakow, p. 27, fig. 13, pl. 30, figs. 1-10. 1933, Sporenformen des Aegirhorizonts, p. 7. SPORITESSPONGIFORMIS Reissinger, 1939, 39, pl. 8, fig. 66. Paleontographica, vol, 84, Abt. B, p. 17. Note.-Nomen nudum, no illustration.

SPORONITESBIPOROUS Berry, 1937, Am. Midland Naturalist, vol. 18, no. 1, p. 160, In the present state of knowledge there fig. 14. are a number of named forms which it is SPORONITESTRIPTERUS Berry, 1937, Am. impossible to treat systematically. The Midland Naturalist, vol. 18, no. 1, p. 160, various generic designations listed below fig. 8.

' may possibly have a certain value but in ZONALA-SPORITESTACITURNUS (Loose) any event their significance does not ap- Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. 3, p. 157. pear to be biological. The names Spor- Sporonites taciturnus Loose, 1932, ites Pollenites, (H. Potonie, 1893)) and Neues Jahrb., Beilage-Band 67, Abt. B., Sporonites (R. Potonie, 1931) cannot be p. 450, pl. 18, fig. 38. regarded as having systematic value. ZONALA-SPORITESULUGHBEKI (Loose), Ib- Forms listed in these groups have no es- rahim, 1933, Sporenf ormen des Aegir- sential biologic features in common, no horizonts, p. 38, pl. 1, fig. 11. type species are recognized, and no sys- Sporo&es ulugbeki Loose, 1932, Neues tematic treatment is feasible on this ba- Jahrb., Beilage-Band 67, Abt. B, p. 447, pl. 14, fig. 11. sis. A morphologic designation apparently would have served as well to record the ELONGATO-SPORITESRETICULATUS Berry, 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 VERRIJCOSA-SPORITESPERVERRUCOSUS (Loose) likewise seems impossible to attach any Loose, 1934, Inst. Palaobot. Arb., vol. 4, no. generic significance to the names Yerru- 3, p. 157. cosa-spo.l.ites (Loose, 1934)) Elongate- Spo~onitesperverrucoszts Loose, 1932, sporites (Berry, 1937)) Pumtata-sporites Neues Jahrb., Beilage-Band 67, Abt. B., p. 451, pl. 18, fig. 48. or Zo~znla-sporites(Ibrahim, 1933). Such problematic forms as have been listed PUNCTATA-SPORITESSABULOSUS Ibrahim, under these names must await future dis- 1933, Sporenf ormen des Aegirhorizonts, p. 37, pl. 5, fig. 43. position. In addition, the following types of 1. SPORITESECHIKOSPINOSUS Zerndt, 1937, Raistrick and Knox may be listed, for Acad. polonaise sci. Trav. Geol. no. 3, they too are of uncertain affinity at the Krakow, p. 17, pl. 24, figs. 5-10. present time. Some of these seem to be worthy of more appropriate classification. 2. SPORITESFUMOSUS Schopf, 1938, Illinois Geol. Survey Rept. In<. 50, p. 52, pl. 5. Raistrick (1937, p. 911) B,, B,, E,, E,, tir:s. 1-3. G,, Knox (1912, pp. 100-101), D,, K,. SUMMARY 61

SUMMARY fossils it seems unavoidable that many This report presents the revised classifi- forms worthy of description will never- cation of plants identified from their theless have such problematic relationship that their assignment under incertae sedis isolated fossil spore coats, and deals is obligatory. The use£ulness 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 on different holotypes have been allowed The atlthors of this paper and their to stand. In many instances, however, associates have observed many new types the apparent close relationship, which may of plant microf ossils in preparations from later prove conspecific, has been noted. coal and carbonaceous sediments in Thus names noted as synonyms are nearly America. The present synopsis is the out- all objective synonyms because they are growth of a need for a more compre- hensive survey of previous work con- based 011 a common holotype. Many difficult problems concerning nomencla- sidered from the standpoint of a con- ture of fossil plants are solvable if nomen- sistent systeniatic policy. The essential 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 do not show convincing evidence of affinity The authors are under particular obliga- with other recognized groups. Such forms tion to Dr. W. H. Camp of the New York have been listed as species excludende. Botanical Garden who, at a period when Attention is directed to the several species other duties were particularly urgent and excluded from Reticulati-sporites which pressing, nevertheless found time to read here is interpreted in a considerably re- the manuscript and suggest numerous stricted sense. Many of these forms re- desirable revisions in the manner of pres- quire much more careful study in order to entation. 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. PALEOZOIC FOSSIL SPORES REFERENCES

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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, Uber Abortiverscheinungen bei -, 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, W. C., 1879, On the organization of 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., 1878, vol. 169, Pt. 11, pp. 319-364. , 1937b, Megasporen aus dem Westfal and Stefan in Bohmen: idem, Bull. internat., , 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 l'avancement Royal Soc. London Philos. Trans., 1880, vol. des etudes de stratigraphie Carbonifere, 171, Pt. 11: pp. 493-540. Heerlen, 1935, Compte rendu, vol. 3, pp. 1711-1732. Wilson, L. R. and Brokaw, A. L., 1937, Plant microfossils 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 deer Zerndt, Jan., 1930a, Megasporen aus dem Isa- Sporen und Pollen, IV. 3. Fossiles Sporo- bell?-Floz (Schichten v. Laziska) in Trze- pollenin aus dem Tasmanit und der Mos- binla : Ann. soc. geol. Pologne, Annee 1929, kauer Braunkohle : Helv. Chim. Acta, vol. vol. 6 : pp. 302-313. 14, Pt. 1, pp. 67-78. PLATES

AND

PLATE

EXPLANATIONS FIG. l-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 Aphanozonati; after Zerndt; 2a, diagram showing un- compressed form in the axial plane corresponding with a trilete ray . . . . . p. 18

3-Trilctes radiosus; sectio undifferentiated ; after Zerndt ; 3a, diagram showing uncom- pressed form in the axial plane corresponding with a trilete ray . . . . pp. 18,24

4-Pulzctati-sporites; #a, equatorial (transverse) plan; 4b, axial (lcmgitudinal) plan . p. 29

5-Laevigato-sporites; 5a, equatorial (transverse) plan; 5b, 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 ; 7b, axial (longitu- dinal) plan ...... p. 34

8-G~anulati-sporites; 8a, equatorial (transverse) plan; 8b, axial (longitudinal) plan . p. 32

9-Demo-sporites; segment cut away to show thickening of peripheral region of spore coat; Pa, axial (longitudinal) plan uncompressed; 9b, 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 ; IOU, small drawing of complete fertile spore ; lob, apex of fertile spore show- ingtrileteapparatus whenabortivemembers are disconnected ...... p.40

FIG. I l-Reinsclzospora; 11a, axial (longitudinal) plan ; 11b, equatorial (transverse) plan . p.52

12-Alisporites; axial (longitudinal) plan not shown; much the same as 16b 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-E~zdosporites; I&, 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-Pityosporites; 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; 16bJ axial (longitudinal) plan . . p. 42

17-Monolctes; 17a, equatorial (transverse) plan; 17bJ 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

EXPLANATIONOF PLATE3

FIG. 18-Raistrickia; Ma, equatorial (transverse) plan ; 18b, axial (longitudinal) plan . . P. 55

19-Lycospora; 19a, equatorial (transverse) plan; l9b, axial (longitudinal) plan . . p. 54

20-Triquitrites; ZOa, axial (longitudinal) plan; 20b, equatorial (transverse) plan . . p. 46

21-Ci1~~,atriradites; Zla, axial (longitudinal) plan; 21b, equatorial (transverse) plan . p. 43

22-Calanzospora; 22aJ 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. U~icosbora' I 15'0~ 1 ~~~~~~~t~-sp)rltes APPROXIMATE SIZE- RANGE ,- ,- Granulatl-spor~tes I 2d0 =Relnschospora I /'- WITHIN "SPORE"GENERA

I I I Florin~tes I I Endosporrtes . -Alisporltes I 1- 1- *a+-sporites I 1 ILLINOIS STATE GEOLOGICAL SURVEY REPORT OF INVESTIGATIONS NO. 91 DISTRIBUTEDIN JULY 1944