14.GS: RPI 180 c. 1

STATE OP ILLINOIS WILLIAM G. STRATTON, Governor DEPARTMENT OP REGISTRATION AND EDUCATION VERA M. BINKS, Director

DIVISION OF THE STATE GEOLOGICAL SURVEY JOHN C. FRYE, Chief URBANA

REPORT OF INVESTIGATIONS 180

MAZOSTACHYS— A NEW CALAMITE

BY

ROBERT M. KOSANKE

PRINTED BY AtTTHORITY OF THE STATE OF ILLINOIS

URBANA, ILLINOIS

1955 LIBRARY

FNVIROMMENTAL PROTECTION AGENCHi STATE OF ILLlDOiS i SPRINGFIELD, ILLINOIS Digitized by the Internet Arciiive

in 2012 witii funding from

University of Illinois Urbana-Champaign

http://archive.org/details/mazostaGhysnewca180kosa STATE OF ILLINOIS WILLIAM G. STRATTON, Governor DEPARTMENT OF REGISTRATION AND EDUCATION VERA M. BINKS, Director

DIVISION OF THE STATE GEOLOGICAL SURVEY JOHN C. FRYE. Chief URBANA

REPORT OF INVESTIGATIONS 180

MAZOSTACHYS — A NEW CALAMITE FRUCTIFICATION

BY

ROBERT M. KOSANKE

PRINTED BY AUTHORITY OF THE STATE OF ILLINOIS

URBANA, ILLINOIS

195 5 ORGANIZATION

STATE OF ILLINOIS HON. WILLIAM G. STRATTON, Governor DEPARTMENT OF REGISTRATION AND EDUCATION HON. VERA M. BINKS, Director

BOARD OF NATURAL RESOURCES AND CONSERVATION HON. VERA M. BINKS, Chairman W. H. NEWHOUSE, Ph.D.. Geology ROGER ADAMS. Ph.D.. D.Sc. Chemistry ROBERT H. ANDERSON, B.S., Engineering A. E. EMERSON, Ph.D., Biology LEWIS H. TIFFANY, Ph.D., Pd.D.. Forestry W. L. EVERITT, E.E., Ph.D. Representing the President of the University of Illinois DELYTE W. MORRIS, Ph.D. President of Southern Illinois University

GEOLOGICAL SURVEY DIVISION

JOHN C. FRYE. Ph.D., D.Sc, Chief

(15457—2M— 4-55) STATE GEOLOGICAL SURVEY DIVISION Natural Resources Building, Urbana JOHN C. FRYE. Ph.D., D.Sc. Chief M. M. LEIGHTON, Ph.D., D.Sc. Chief, Emeritus Enid Townley, M.S., Geologist and Assistant to the Chief Velda a. Millard, Junior Assistant to the Chief Helen E. McMorris, Secretary to the Chief RESEARCH (not including part-time personnel) GEOLOGICAL RESOURCES GEOCHEMISTRY Arthur Bevan, Ph.D., D.Sc, Principal Geologist Frank H. Reed, Ph.D., Chief Chemist Frances H. Alsterlund, A.B., Research Assistant Grace C. Johnson, B.S., Research Assistant Coal Chemistry Coal G. R. Yohe. Ph.D., Chemist and Head Jack A. Simon, M.S., Geologist and Head Earle C. Smith, B.S., Research Assistant G. H. Cady, Ph.D., Senior Geologist and Head, Emeritus GuEY H. Lee, M.S., Research Assistant Robert M. Kosanke, Ph.D., Geologist Raymond Siever, Ph.D., Geologist Physical Chemistry John A. Harrison, M.S., Associate Geologist J. S. Machin, Ph.D., Chemist and Head Paul Edwin Potter, Ph.D., Associate Geologist JuANiTA Witters, M.S., Assistant Physicist Harold B. Stonehouse, Ph.D., Associate Geologist Tin Boo Yee, Ph.D., Assistant Chemist Margaret A. Parker, M.S., Assistant Geologist Daniel L. Deadmore, B.S., Research Assistant (on leave) Fluorine Chemistry M. E. Hopkins, M.S., Assistant Geologist Kenneth E. Clegg, M.S., Assistant Geologist G. C. Finger, Ph.D., Chemist and Head Robert E. Oesterling, B.A., Assistant Chemist Carl Kruse, M.S., Oil and Gas W. Special Research Assistant Raymond H. White, B.S., Special Research Assistant A. H. Bell, Ph.D., Geologist and Head Richard H. Shiley, B.S., Special Research Assistant Lester L. Whiting, B.A., Associate Geologist Virginia Kline, Ph.D., Associate Geologist Chemical Engineering Wayne F. Meents, Assistant Geologist H. W. Jackman, M.S.E., Chemical Engineer and Head Margaret O. Oros, B.A., Assistant Geologist R. J. Helfinstine, M.S., Mechanical Engineer and Kenneth R. Larson, A.B., Research Assistant Supervisor of Physical Jacob Van Den Berg, B.S., Research Assistant B. J. Greenwood, B.S., Mechanical Engineer James C. McCullough, Research Associate (on leave) Petroleum Engineering Robert L. Eissler, B.S., Assistant Chemical Engineer Walter E. Cooper, Technical Assistant M.S., Petroleum Engineer and Paul A. Witherspoon, Edward A. Schaede, Head Technical Assistant Cornel Marta, Technical Assistant Frederick Squires, A.B., B.S., D.Sc, Petroleum Engi- neer, Emeritus X-Ray W. F. Bradley, Ph.D., Chemist and Head Industrial Minerals Analytical Chemistry E. Lamar, B.S., Geologist and Head J. O. W. Rees, Vn.^D.', Chemist and Head Donald L. Graf, Ph.D., Geologist L. D. McVicKER, B.S., Chemist James C. Bradbury, A.M., Assistant Geologist Emile D. Pierron, M.S., Associate Chemist Meredith E. Ostrom, M.S., Assistant Geologist Donald R. Dickerson, B.S., Assistant Donald L. Biggs, M.A., Assistant Geologist Chemist Francis A. Coolican, B.S., Assistant Chemist Charles T. Allbright, B.S., Research Assistant Clay Resources and Clay Mineral Technology William J. Armon, B.S., Research Assistant Ralph E. Grim, Ph.D., Consulting Clay Mineralogist Joseph M. Harris, B.A., Research Assistant W. Arthur White, M.S., Associate Geologist JoAnne E. Kunde, B.A., Research Assistant Herbert D. Glass, Ph.D., Associate Geologist Joan M. Cederstrand, Research Assistant Charles W. Spencer, M.S., Research Assistant Eugene Lange, Technical Assistant George R. James, Technical Assistant Groundwater Geology and Geophysical Exploration Frances L. Scheidt, Technical Assistant Arthur Bevan, Ph.D., D.Sc, Acting Head MINERAL ECONOMICS Merlyn B. Buhle, M.S., Associate Geologist Robert E. Bergstrom, Ph.D., Assistant Geologist W. H. VosKUiL, Ph.D., Mineral Economist John W. Foster, M.S., Assistant Geologist W. L. BuscH, A.B., Assistant Mineral Economist James E. Hackett, M.S., Assistant Geologist Ethel M. King, Research Assistant Margaret J. Castle, Assistant Geologic Draftsman (on leave) EDUCATIONAL EXTENSION Wayne A. Pryor, M.S., Assistant Geologist George M. Wilson, M.S., Geologist and Head Lidia Selkregg, D.N.S., Assistant Geologist Dorothy E. Rose, B.S., Assistant Geologist Robert C. Parks, Technical Assistant

Engineering Geology and Topographic Mapping RESEARCH AFFILIATES IN GEOLOGY George E. Ekblaw, Ph.D., Geologist and Head J Harlen Bretz, Ph.D., University of Chicago William C. Smith, M.A., Assistant Geologist John A. Brophy, M.S., Research Assistant, State Geolog- ical Survey Stratigraphy and Areal Geology Stanley E. Harris, Jr., Ph.D., Southern Illinois Uni- versity H. B. Willman, Ph.D., Geologist and Head C. Leland Horberg, Ph.D., University of Chicago David H. Svi^ann, Ph.D., Geologist M. M. Leighton, Ph.D., D.Sc, Research Professional Elwood Atherton, Ph.D., Geologist Scientist, State Geological Survey Charles W. Collinson, Ph.D., Associate Geologist Heinz A. Lowenstam, Ph.D., California Institute of Donald B. Saxby, M.S., Assistant Geologist Technology T. C. Buschbach, M.S., Assistant Geologist William E. Powers, Ph.D., Northwestern University Howard R. Schwalb, B.S., Research Assistant Paul R. Shaffer, Ph.D., University of Illinois Frank B. Titus, Jr., B.S., Research Assistant Harold R. Wanless, Ph.D., University of Illinois Charles C. Engel, Technical Assistant J. Marvin Weller, Ph.D., University of Chicago Joseph F. Howard, Assistant CONSULTANTS Physics Geology, George W. White, Ph.D., University of Illinois Illinois R. J. PiERSOL, Ph.D., Physicist, Emeritus Ralph E. Grim, Ph.D., University of L. E. Workman, M.S., Former Head, Subsurface Topographic Mapping in Cooperation with the Division United States Geological Survey. Ceramics, Ralph K. Hursh, B.S., University of Illinois Mechanical Engineering, Seichi Konzo, M.S., University of March 16, 1955 Illinois GENERAL ADMINISTRATION (not including part-time personnel) LIBRARY OTHER TECHNICAL SERVICES Anne E. Kovanda, B.S., B.L.S., Librarian Wm. Dale Farris, Research Associate Ruby D. Prison, Technical Assistant Beulah M. Unfer, Technical Assistant A. W. GOTSTEIN, Research Associate Glenn G. Poor, Research Associate* MINERAL RESOURCE RECORDS Gilbert L. Tinberg, Technical Assistant Vivian Gordon, Head Wayne W. Nofftz, Supervisory Technical Assistant Margaret B. Brophy, B.A., Research Assistant Donovan M. Watkins, Technical Assistant Sue J. Cunningham, Technical Assistant Betty Clark, B.S., Technical Assistant FINANCIAL RECORDS Jeanine Climer, Technical Assistant Marilyn W. Thies, B.S., Technical Assistant Velda a. Millard, In Charge Hannah Fisher, Technical Assistant Leona K. Erickson, Clerk-Typist III LaRoy Peterson, Technical Assistant Virginia C. Sanderson, B.S., Clerk-Typist II Gary Jean Stevenson, Technical Assistant Irma E. Samson, Clerk-Typist I Patricia L. Luedtke, B.A., Technical Assistant Kathryn Brown, Technical Assistant CLERICAL SERVICES Mary Cecil, Clerk-Stenographer III PUBLICATIONS Mary M. Sullivan, Clerk-Stenographer III Barbara Zeiders, B.S., Assistant Technical Editor Lyla Nofftz, Clerk-Stenographer II Meredith M. Calkins, Geologic Draftsman Lillian Weakley, Clerk-Stenographer II Marlene Ponshock, Assistant Geologic Draftsman Sharon Ellis, Clerk-Stenographer I Barbara Barham, Clerk-Stenographer I Mary Alice Jacobs, Clerk-Stenographer I TECHNICAL RECORDS Lorraine Cunningham, Clerk-Stenographer I Irene Benson, Clerk-Typist I Berenice Reed, Supervisory Technical Assistant Mary J. de Haan, Messenger-Clerk I Marilyn DeLand, B.S., Technical Assistant Mary Louise Locklin, B.A., Technical Assistant AUTOMOTIVE SERVICE GENERAL SCIENTIFIC INFORMATION Glenn G. Poor, In Charge* Robert O. Ellis, Automotive Mechanic Ann p. Ostrom, B.A., Technical Assistant Everette Edwards, Automotive Mechanic Jill B. Cahill, Technical Assistant David B. Cooley, Automotive Mechanic's Helper

March 16, 1955 *Divided time CONTENTS

Page Introduction 7 Acknowledgments 8 Mazostachys pendulata gen. et sp. nov. General description 9 Shoot axis 10 Peduncles 10 Cone axis 10 Bracts 11 Sporangiophore 14 Sporangia 14 15 Diagnosis 17 Discussion 17

Pre-C^/rtwo.y/(^f/!_>'j type of fructification 17 Calamostachys type of fructification 17

Pi3/^'j type of fructification 19 Phylogeny of the cones of the 20 Summary 22 References 23

ILLUSTRATIONS

Figure

1. Diagrammatic drawing of holotype specimen B-1304A 8

2. Diagrammatic drawing of holotype specimen B-1304B 9

3. Reconstruction o[ Mazostachys pendu/aia 12

4. Transverse section of Mazostachys pendulata 13

5. Proposed phyletic sequence for cones of Calamitaceae 22

PLATES

Plate

1. and longitudinal sections 27 2. Transverse sections 28

3. Transverse sections 31 4. Habit and sections of tissues 32

5. Transverse section of cone axis and sporangia 2>S 6. Longitudinal section, reconstructions, tissues, and 36

MAZOSTACHYS — A NEW CALAMITE FRUCTIFICATION

ROBERT M. KOSANKE

ABSTRACT

The specimen described is preserved in an ironstone concretion collected from spoil piles of the No. 2 coal bed of age in Will County, 111. It is a calamite shoot bearing fifteen cones. Each cone axis, with nodes and internodes, bears appendages of two types. A whorl of twelve bracts occurs on each node and high on each internode are six whorled sporangiophores. Whorls of sporangiophores and bracts alternate along the vertical extent of the cone axis. Two sporangia are attached to each sporangiophore in a pendulant manner. The striking features of this type of calamite fructification are: (1) the position of attachment of the sporangiophore, and (2) the presence of two sporangia per sporangiophore, a new feature of calamite . The second feature demonstrates that the phylogeny has progressed with reduction in number of sporangia per sporangiophore from four in Asterocala- mites Pothocites, Protocalamostachys, Calamostackys, Palaeostachya, and Cingularia

typica, to two in this specimen (described as Mazostac/iys) , to only one in Metacala- mostachys. Mazostachys is homosporous. The foliage of sphenophylloides occurs in the ironstone concretion in association with Mazostachys but not in organic connection with the fertile shoot.

INTRODUCTION Mazostachys was removed from the shale during stripping operations. The following report is a contribution The Francis Creek shale lies in the lower to fundamental studies of of part of the Carbondale group of Pennsyl- Pennsylvanian age of Illinois. It is a con- vanian age in Illinois. According to Moore tinuation of the Survey's efforts to explore et al. (1944), an equivalent stratigraphic past plant life seeking knowledge of coal- position in the mid-continent region would forming plants of potential value in the be in the upper Cherokee group of the Des correlation of coal beds and in related stud- Moines series and in the Allegheny series ies in the field of coal petrography. Studies of the Appalachian region of eastern United of fossil cones are also useful to paleobot- States. For additional information about anists in associating isolated spores in coal the Francis Creek shale, see Cady (1919), with their parent plants. Savage (1927), Wanless (1929), and This report describes a new type of cala- Willman and Payne (1942). mite fructification and gives a re-evalua- The external appearance of the concre- tion of previous ideas on the phylogeny of tion containing the specimen of Mazo- these fructifications in light of new evi- stachys is typical of others found in the area, dence. The specimen, with preserved cellu- but the mode of preservation is not en- lar structure, was found in an ironstone tirely of the compression type. The main concretion on the spoil piles of the strip- axis and branches of the specimen are prin- ping operations of the Northern Illinois cipally carbonaceous material, but each cone

Coal Corp., near the Mazon Creek area, is encased in, or surrounded by, calcium

Will County, 111. carbonate. Therefore, preservation is some- Ironstone concretions with preserved what similar to a coal ball in which histo- plant remains occur in the Francis Creek logical details can be observed. Peels and shale immediately above the No. 2 coal bed thin sections were used in studying the pre- in the area. It is assumed that the iron- served cellular structure of the fructifica- stone concretion bearing the specimen of tions. Individual sporangia from several [7] 8 ILLINOIS STATE GEOLOGICAL SURVEY cones were dissected and macerated so that above the Murphysboro coal bed in south- the spores could be studied individually in western Illinois. Condit and Miller (1951) the isolated state. Specimens with pre- reported plant-bearing ironstone concre- served cellular structure from the Mazon tions from Iowa. Creek area are rare, but the author has a specimen of Calamostachys which is not as Acknowledgments well preserved as the specimen described. It is a pleasure to acknowledge the as- Darrah (1936) described Macrostachya sistance of Dr. Wilson N. Stewart, under ihompsonii in an ironstone concretion from whose supervision this report was submit- the Mazon Creek area that is preserved in ted in 1952 to the Graduate College, Uni- a similar manner. versity of Ironstone concretions containing plant Illinois, as part one of a doctoral thesis in compressions occur at a number of strati- . Dr. Theodore Just, Chi- graphic levels and in various areas of the cago Natural History Museum, assisted in

United States. I have observed these con- the selection of the new names proposed. cretions from a shale overlying the Pome- Dr. James R. Kamp and Mr. and Mrs. roy coal bed in Ohio, above the Minshall John M. McLuckie, Coal City, 111., as- coal bed in Warren Countv, Ind., and sisted in obtaining the described specimen.

Fig. 1. —A diagrammatic drawing of the holy type specimen of Mazostachys pendulata gen. et sp. nov. (B- 1304A). The cones were assigned numbers for ease of discussion in text. Cones 3 and 9 did not occur on

this half of the specimen, so their position is shown by dashed lines. About 1/% natural size. GENERAL DESCRIPTION 9

MAZOSTACHYS PENDULATA men in the subgenus Encalamites, typified GEN. ET SP. NOV. by carinatus as illustrated by Hirmer (1927, figs. 537 to 543). Each GENERAL DESCRIPTION branch redivides (figs. 1 and 2), one bear-

The specimen is in the Illinois State Geo- ing seven cones and the other eight. The logical Survey, paleobotanical collection, divisions of the two branches are somewhat number B-1304. It was split in half and, irregular and not entirely typical of Cala- accordingly, will be referred to as B-1304A mites carinatus. The internode of each

and B-1304B (figs. 1 and 2; pi. 1, fig. 1; branch is 1 cm. long and less than 2 mm. and pi. 4, fig. 1). It is a fertile calamite wide. The branch that gives rise to cones shoot with 15 attached cones. Foliage is 1 through 7 (figs. 1 and 2) bifurcates; one

not attached, but six whorls of iden- dichotomy gives rise to cones 1 through 3 tified as Annularia sphenophylhides are and the other dichotomy bears cones 4 preserved as a very thin carbonaceous film through 7. The branch giving rise to cones in the same plane as the specimen. 8 through 15 divides in true eucalamite The shoot axis has two nodes and one style at the first node. Cones 12 and 13 are

complete internode. There is also a small formed in the same manner. The first node portion of another internode. The complete of the other division gives rise to cone 11

internode is 4.2 cm. long and slightly less and a sterile node leading to the culmina-

than 1 cm. wide. Two branches depart tion of cones 8 through 10 (as in the for- from the shoot axis opposite each other; mation of cones 4 through 7). The second thus the mode of branching places this speci- node of the branch bearing cones 8 through

Fig. 2. —A diagrammatic drawing of the holotype specimen of Mazostachys pendulata gen. et sp. nov. (B- 1304B). All fifteen cones occur on this half of the specimen. About % natural size. 10 ILLINOIS STATE GEOLOGICAL SURVEY

15 appears to bifurcate, with one of the re- consisted of patches of preserved parenchy- sulting branches bearing cones 14 and 15 ma and a few tracheids with scalariform and the other perhaps bearing foliage. and multiseriate scalariform pitting. There

There is no organic connection between the is no indication of a pith cavity. cone-bearing and -bearing shoots, but CONE AXIS their proximity suggests that they may be-

long to the same plant. The preservation of the cone axis is usu-

Each cone is attached to a short peduncle ally good except for the phloem and cortex. usually less than 2 mm. in length; the pe- Tracing the vascularization in the longitu-

duncle for cone 7 is somewhat longer. dinal section is difficult, owing to its slen- der nature and to the fact that these axes The cones vary in length, depending are preserved with minor curvatures or upon the number of whorls per cone. Cone twists. Plates 2 and 3 show the axis of 1 has nine whorls; cones 8 and 15, eight; cone 14 in transverse section from the base cones 4 and 7, seven; cone 6, six; cones 10 of the sporangia of the second whorl from and 14, five; cones 11, 12, and 13, three; the proximal end of the cone to the base of and cones 2, 3, and 9, one. The largest the sporangia of the next higher whorl (pi. cone is number 1, which is 2.6 cm. long and 1, fig. 2). Thirty-eight peels were made just under 4 mm. wide. between these two points, a distance of ap- SHOOT AXIS proximately 1900 microns.

It is apparent from an examination of The shoot axis is preserved as a true com- plates 2 and 3 that no internodal pith cav- pression. Three peels of B-1304A con- ity is present in the transverse sections of tained carbonaceous matter, whereas only the cone axis. The absence of a pith cav- the first peel of B-1304B contained any ap- ity in longitudinal section may be seen in preciable carbonaceous matter. Because of figure 3, plate 1, and figure 1, plate 6. In degradation and poor structural preserva- this respect, Aiazostachys pendulata sp. tion, superficial ribbing and furrowing is nov. is similar to Calamostachys binneyana. apparent only from the external surface of The pith is composed of parenchyma cells the shoot. Examination of peels shows only which are largest in diameter at the center a few scattered tracheids with scalariform of the cone axis. At the middle of the inter- and multiseriate scalariform pitting, the nodes, these cells are about seven times latter similar to that described by Delevor- longer than their maximum width. Be- yas and Morgan (1952). Similar pitting tween the vascular bundles they are about was termed scalariform or multiseriate pit- one-half this size. ting by H^eg (1942). Patches of paren- Carinal canals are located between the chyma cells were observed, but these could extensions of the pith of the internodal re- not be correlated with any specific tissues gion, at the level of departure of the spo- of the shoot axis. The branches arising rangiophore traces (pi. 5, fig. 1). Six canals from the shoot axis are also poorly pre- are still evident at the level of the bract served, and only a few tracheids with sca- traces, but they fuse or pair a short dis- lariform pitting were observed. Parenchy- tance above this level, where they lie adja- ma cells are present, and their position sug- cent to three regions of secondary wood gests a central pith. similar to Calamostachys binneyana as illus- PEDUNCLES trated by Williamson and Scott (1894). In a transverse section of cone 14 The peduncles attached to the cones are (B-1304A), peel 25, four carinal canals exceedingly thin, delicate structures, and are paired close together, whereas two are their preservation is very poor. Only one still somewhat separated. All ultimately or two peels in which peduncles were in- pair and fuse at a slightly higher level on cluded contained structural details. These the axis, so a somewhat triangular stele re- BRACTS 11

suits at a point just above the middle of the The cone axis of Mazostachys pendulata nodal region. The pairing of carinal ca- sp. nov. is a dictyostele consisting of sep- nals is believed to be the result of rearrange- arate vascular bundles which form a net- ment of cells in the nodal region. An at- work. The primary develops cen- tempt was made to determine whether or trifugally and is endarch in arrangement. not there is an alternation of vascular bun- Only a few annular thickenings of the pro- dles between successive nodes, a possibility toxylem remain in the carinal canals. Abun- strongly suggested by the pairing of the dant metaxylem lies to the outside of the carinal canals. Peels 1 through 38 of cone canals of the internodal region. Secondary

14 were examined and no evidence was wood is developed at the nodes and beyond found for an alternation of the bundles, al- the metaxylem. Tissues from the cambium though there is a shift in the position of the to the epidermis are not preserved. Meas- carinal canals. This shift could be due in urements of a number of cones in transverse part to preservational factors. and longitudinal section show that the max- preserved The protoxylem of the cone axis is imum nodal diameter is 750 largely disintegrated, leaving the carinal microns at the base of the cone. In cone 1 canals. The remains of annular protoxylem this nodal diameter decreases to 450 mi- elements line the canal walls. Plate 4, fig- crons at the cone apex. From base to apex of the cones, ure 3, shows the protoxylem remains in the the internodal diameter de- creases canal. Portions of three tracheids lie side from about 475 microns to about microns. by side, filling the canal at a point near the 325 top of the figure. It is believed that the BRACTS protoxylem mass, prior to the formation of of three, perhaps the canals, was composed There are twelve bracts per whorl, lo- four, vertical of tracheids. the rows At cated at the nodes of the cone axis. From the is metaxy- periphery of protoxylem the an external view and in longitudinal sec-

; pitting lem its tracheids have scalariform tion, the bracts appear to depart from the (pi. fig. remains of an annular 4, 3). The axis simultaneously. A double whorl of six protoxylem element may be seen in plate 5, bracts given off at each node is suggested, figure 1. perhaps as the result of slightly oblique sec- No evidence was found that the secon- tions. The bracts are connate at the sessile dary wood forms a complete ring in the attachment with the cone axis. The degree nodal region. In all sections prepared, por- of fusion of the bracts is much less than in tions of the pith extend between the vascu- Calamostachys binneyana, in which the en- lar bundles to the periphery of the pre- tire basal portion of the bracts is connate for a served portion of the cone axis, although in considerable distance from its inser- tion. bracts several sections the peripheral width of the The are located at right angles to the axis of the cone (fig. 3; pi. 1, figs. 2 parenchyma was only one or two cells. The and 3; pi. 6, figs. 1 and 2). They arch phloem, cortex, and epidermis are not pre- around the base of the sporangia and then served, so it is possible that a portion of the turn upward parallel to the cone axis. The secondary wood is also missing. The vascu- apex of the bract slightly overlaps the base lar tissue in the enlarged nodal region dis- of the of the next whorl above plays scalariform (pi. 4, fig. 6), multiseri- (pi. 1, figs. 2 and 3; pi. 6, figs. 1 and 2). ate scalariform (pi. 6, fig. 4), and reticu- The vertical extent of the bracts where late pitting of the metaxylem and secondary they parallel the axis is from 3 to 3.5 mm. wood. Node enlargement in the cone axis The bracts vary in width depending upon results from formation of secondary wood the point of measurement. At the level of and departure of vascular traces, as in Pa- the base of the sporangia the bracts are laeostachya vera and Calamostachys binne- about 1.25 mm. wide; at the top of the yana. sporangia they are 500 microns. From this 12 ILLINOIS STATE GEOLOGICAL SURVEY

Fig. 3. —A diagrammatic reconstruction of Mazostachys pendulata; the lower half (below the dashed line) shows the position and attachment of the sporangiophores and sporangia. The position of the bracts is shown below. Aerenchyma tissue above the sporangia has been removed for clarity. The upper half of the drawing shows two nodes and one internode in longitudinal section. The vascular supply is shown by solid black lines. Note that the sporangia originate below the bracts of the node above and that they are pendulant. The bracts and sporangia are superposed. Abbreviations: AX, cone axis; BR, bract; PBR, position of bract; SM, sporangium; and SP, sporangiophore. BRACTS 13

point upward their width decreases rap- figure 3 and in transverse section in plates idly and they form a sharp apex. Where 2 and 3. The twelve bracts are in both op- they turn upward and parallel the cone posite and alternate positions relative to axis the bracts are about 200 microns thick; the six sporangiophores. The bracts are to they thin to about 100 microns near the apex the outside of, and alternate with, the of the bracts. The relative position of the twelve sporangia (fig. 4). In the early de- bracts to the sporangiophores is shown in velopmental stages of the cones, the bracts

3725//.

Fig. 4. —Diagrammatic transverse section oi Maiostachys pendulata just below a node; for clarity, the aeren- chyma tissue is not shown. There are 12 sporangia attached to 6 sporangiophores and 12 bracts opposite and alternate to the sporangiophore. This reconstruction is based on peels from whorl 2 of cone 14. Ab- breviations: AX, cone axis; BR, bract; SM, sporangium; and SP, sporangiophore. 14 ILLINOIS STATE GEOLOGICAL SURVEY

were closely pressed against the sporangia. ceeds up and out and then bends slightly Evidence for this can be seen in transverse downward into the sporangiophore. There plane, where the bracts have assumed some- are four or five rows of tracheids with

thing of the shape of the available space spiral thickenings (pi. 4, fig. 5) forming between the sporangia. Seen in transverse the trace. No gaps have been observed in plane, the sporangia also arch at the ex- the vascular bundle of the axis supplying terior surface, which was the exposed spo- the vascular tissue for the sporangiophores.

rangial portion between adjacent bracts. The departure of the sporangiophore The single vascular trace supplying each first appears below the mid-point of the

bract is median and adaxial, and it main- internodal region of the cone axis, but it tains this position throughout the entire does not become free from the axis until bract. The tracheids of the bract trace much higher on the internode. The mode have spiral thickenings (pi. 4, fig. 4), and of departure was first observed in peel 7 the diameter of the trace is about the same of cone 14, which, as nearly as can be cal- or perhaps slightly smaller than that of the culated, is about 700 microns above the sporangiophore trace. subtending bract. The sporangiophore

continues to enlarge (pi. 2, figs. 2 and 3). SPORANGIOPHORE Probably at one stage in their development, The most striking feature of Mazo- the sporangia were pressed against the spo-

is the position of the sporangio- stachys rangiophores ; this is suggested by the con- phore. It departs at right angles from the tour of the underside of the sporangio- cone axis high on the internode and only phore, which parallels that of the sporangia slightly below the bract of the next whorl until contact between the two is estab- (fig. 3; pi. 1, figs. 2 and 3; pi. 6, figs. 1 lished. and 2). The sporangiophore extends out The sporangiophore first appears low on the pendulate spo- over and between two the internodal region only a short distance it rangia which it bears (fig. 3) ; then above the subtending bract; it becomes free curves down around the upper peripheral of the axis and receives its vascular supply portion of the sporangia and parallels the at a point seven-eighths of the way up the axis for a short distance. Hirmer (1927) internode. proposed the name Metacalamostachys for two species formerly assigned to Calamo- SPORANGIA stachys. His reasons for creating the There are twelve sporangia per whorl in were based on the fact that the sporangio- mature cones, borne on six sporangiophores phore originated below the bract of the next (two sporangia per sporangiophore). The whorl above and the fact that there was sporangiophore forms an arm from which only one sporangium per sporangiophore. the sporangia hang in a pendulant manner

is the Metacalamostachys known only from (pi. 1, figs. 2 and 3; pi. 6, figs. 1 and 2).

compression state of preservation. This in- This mode of attachment is not peltate as in vestigation is the first to report anatomical Calajnostachys and Palaeostachya; instead, details of calamite fructifications with pen- the sporangia hang downward as in Cingu- dulate sporangiophores. laria and Metacalamostachys. The sporan- As stated above, the preservation of the gia average slightly more than 1.5 mm. in bracts and sporangiophores leaves much to length.

be desired, but it has been possible to locate The sporangial wall is usually only one

the sporangiophore trace at its point of de- cell layer thick, and this is similar to that parture from the cone axis and also described by Hickling (1907) for Palaeo- throughout the sporangiophore (fig. 3; pi. stachya and by Scott (1920) for Calamo- 4, fig. 5). stachys. Seen in transverse plane, the indi- As the sporangiophore trace departs from vidual cells are rectangular, about 100 mi-

the vascular bundle of the cone axis, it pro- crons long by 35 microns wide. They are SPORES 15

characterized by thickenings of the cell gia, it probably completely covered the wall, which project into the lumen of the sporangia. However, not a single cell of cell. These projections have been called this tissue has been observed below the up-

buttresses by Hickling (1907) and stiffened per third of the sporangia. There is an or- projecting ridges by Scott (1920). An ex- ganic residue at the contact between the amination of plate 4, figure 2, reveals that ironstone concretion and the calcium car- a few of these may be traced the full width bonate (pis. 2 and 3), more marked in some of the cell lumen. Such configurations are places than others, that may represent the produced by thickenings that originate on degradated aerenchymatous tissue. the lateral of cells and extend wall the The method of dehiscence is unknown. across the lumen to the opposite lateral Hoskins and Cross (1943) found no evi- wall. In some cells the thickenings also ex- dence to indicate that the irregular thick- tend from the bottom to the top of the cell. enings of the sporangial wall in Bowrran- This is probably why they do not all ex- ites are similar to the thickened walls of tend the full width ot the lumen when seen an annulus as had been suggested by Zeil- in tangential view. The thickenings are ler (1893) and Williamson and Scott somewhat similar to the partition in sep- (1894). In Mazostachys the tangential tate fibers; they are not true cross walls walls of the sporangial cells are thinner because the middle lamella does not extend than the radially arranged fibrous thick- into the partition. The projections of spo- ening. This would suggest an annulus, but rangial cell wells are unlike those of sep- unlike the sporangial wall of and some tate fibers in that they are thicker or at bryophytes, these thickenings are present least more opaque. uniformly throughout all cells of the wall As previously stated, the sporangia are layer.

only one cell thick (pi. 5, fig. 2). There is SPORES a possibility, however, that the sporangia Mazostachys wall gives rise to the tissue lying on top of pendulata is homosporous, the sporangia and extending to the sterile and the spores from these fructifications are densely packed in the (pi. bracts (pi. 6, figs. 1 and 2). As seen in sporangia 1, figs. and pi. 2 3 ; 2 ; pi. 3 pi. 5, fig. pi. fig. plate 6, figure 5, this tissue is composed of ; 2 ; 6,

1 ) . spores are parenchyma cells between which are very The of great value when as- sociating the isolated calamite large lacunae. This is similar in over-all spores from the Illinois appearance to the lacunar cortex in the No. 2 coal bed with the parent fructifications, free of Psaronius. A microscopic ex- because the possibility of ob- taining similar amination of the two types of tissue shows fructifications from coal balls is remote. the similarity to be one of cell size plus the Coal balls have been col- lected from the presence of lacunae. The cells walls of in- No. 2 coal bed, but they are rare contain dividual cells are vastly different, however; and very poorly preserved plant remains those of Psaronius are comparatively thin- owing to a large amount of pyrite and walled, whereas those of Mazostachys are a low concentration of calcite. Accordingly, thicker and much more irregular. In fact, if affinities are to be estab- lished between the cell-wall structure of the lacunar tissue isolated spores of the No. 2 coal bed and the parent fructifications, of Mazostachys is similar to that of its spo- the only known source of material is rangial wall and is suggestive of an epi- from iron- stone concretions dermal proliferation of an aerenchymatous- with cellular structure. type tissue. The extent to which the aeren- The spores were examined from peels, chyma tissue originally covered the spo- thin sections, and in isolation. To prepare rangia is unknown. Only fragmental por- material in an isolated state, several spo- tions of it are preserved (pi. 1, fig. 3; pi. rangia were dissected from the rock matrix 6, fig. 1). If the function of this aeren- and placed in a 10 percent solution of hy- chymatous tissue was to protect the sporan- drochloric acid. Within several minutes, 16 ILLINOIS STATE GEOLOGICAL SURVEY

water was added to the samples. After six Kosanke (1950) described the spores changes of water a concentrated aqueous from the No. 2 and a number of other Illi- solution of safranine Y was added. Three nois coal beds. Spores with calamite fruc- hours later the aqueous solution of safra- tifications were assigned to Calamospora nine was removed by pipette. This was fol- following Schopf, Wilson, and Bentall lowed by changes of alcohol, and ultimately (1944). I found Calamospora to have a the spores were mounted in diaphane. stratigraphic range throughout almost every

The spores are radial, trilete, round in coal bed examined in the Caseyville, Trade- outline, and invariably folded. Two hun- water, Carbondale, and McLeansboro dred spores were measured at their largest groups in Illinois and to be abundant in the diameter and also at right angles to the last three. Calamospora was found in larg- first measurement. The average diameter est quantity in the lower McLeansboro was 59.5 X 56.1 microns; the smallest speci- group and was present in moderate abun- men was 51.8x51.8 microns, the largest, dance in the No. 2 coal bed immediately 70.9 x 65.5. Most spores measured be- below the Francis Creek shale, the unit tween 54.6 and 69.6 microns at their larg- from which Mazostachys pendulata was col- est diameter. The trilete rays are short, lected. Three species of Calamospora were usually less than 10 microns. The lips are identified from the No. 2 coal bed; they

thin but distinct, and the commissure is usu- are C. hartuiigiana, C. fiexilis, and C. brev- ally thin but also distinct. The area con- iradiata. The spores of Mazostachys pen- dulata certainly are not comparable with tagionis is very sharply defined (pi. 5, of first two, exists fig. 2; pi. 6, fig. 6). Hartung's (1933) those the and some doubt illustrations of Macrostachya infundibuli- that they are conspecific with C. breviradi- for?nis spores show the area cojitagioiiis ata. They are similar to this last species in clearly but not nearly so sharply defined all but one respect—the area co?itagionis is as in most isolated specimens from Mazo- present in C. breviradiata but is not as clear- stachys pendulata. More than 500 speci- ly defined. This comparison is based not on examination of a single specimen but of mens were examined and all possessed the many specimens, and there seems to be suf- area co?itagionis, although it is interest- ficient evidence to distinguish between ing to note that this feature was less separate species. Perhaps this dis- prominent in the larger specimens. By them as exist if the spores of chance, several specimens had been broken tinction would not Mazostachys pendulata were fully mature and mounted in such a way that the thick- This has not ness of the area contagionis could be meas- and shed from their cones. been proved, however, so they must be ured. I had always thought that this area considered distinct from Calamospora brev- was greatly thickened, but surprisingly it was found to be no more than 3 microns iradiata. thick, whereas the spore coat was slightly Mazostachys pendulata was deposited less than 2 microns. The darker color of less than 10 feet from the top of the coal

these areas in this species^ is evidently not bed, so it might be cause for concern that due to thickness alone, for these areas are spores of Mazostachys pendulata are not considerably darker than those in which the known to occur in the No. 2 coal bed. The spore coat may be badly folded and, there- flora of the Mazon Creek ironstone concre- fore, thicker than the area contagionis. The tions is well known from the works of

spore coat is levigate and not more than Lesquereux (1866, 1870), Noe (1925), 2 microns thick. In thin sections or peels, Darrah (1936), Schopf (1938), Janssen the spore coat appears to be slightly granu- (1940), Stewart (1950), and many others, lar rather than levigate (pi. 5, fig. 2). In but this type of fructification has not pre-

some sections it almost appeared as if there viously been recorded. Also the spores, as were an additional thin layer surrounding preserved in the cones, have not been ob- the spores. served in any coals in Illinois. This sug- DISCUSSION 17

gests that AI. perululata was not an abun- first, or pre-Cala/nostachys, type are a num- dant member of the Pennsylvanian flora of ber of related fructifications such as Potho- Illinois, or that the spores have undergone cites, Archaeocalaniites radiatus, Astero- significant morphological changes during calamites scrobiculatus, Protocalamosta- a late stage of development. chySj and others. These are known only from compressions with the exception of Diagnosis Protocala/nostachys, which is a petrifaction The cones of Mazostachys pendulata (very well described by Walton, 1949). gen. et sp. nov. are approximately 2.6 cm. Two other distinct types, Calamostachys long and about 4 mm. wide. They are at- and Palaeostachya, are known both from tached by a peduncle, usually less than 2 compressions and petrifactions. Macro- mm. long, to a shoot with a branching pat- stachya is of the Calamostachys type and tern like that of the eucalamite group. The is known from compressions, although some

cone axis is a dictyostele, and the primary anatomical data is available as a result of xylem develops centrifugally. There are six Darrah's work (1936). Huttonia is known only from vascular bundles in the internode ; they pair compressions; according to Hir- to form three in the region of secondary mer (1927), it is related to Palaeostachya. wood in the nodes. Twelve bracts arise at Two more distinct types, Cingularia and Metacalamostachys, each node ; they are in part connate at the are known only from sessile attachment with the cone axis. There compressions. Mazostachys, described in are six sporangiophores, each originating this report, is the sixth type of calamite just below the sterile bract of the node fructification. above. There are twelve sporangia, two PRE-Calamostachys type of fructifi- attached to each sporangiophore in a pen- cation dulant manner. The sporangia are about Calamite fructifications of this type dif- 1.5 mm. long, and the sporangial wall is fer from the remaining five types in that one cell thick with radial projections. A they are essentially bractless. They occur tissue composed of parenchyma with nu- in strata assigned to the lower portion of merous lacunae covers the sporangiophores the Lower , so are geolog- and the upper portion of the sporangia. M. ically the oldest known fructifications of pendulata is homosporous, the spores aver- the . Their geologic age and aging about 60 microns in diameter, and bractless condition seem to warrant the use there is a distinct area contagionis. of this type of fructification as a starting Locality.—Northern Illinois Coal Com- point in developing a phyletic series. pany, recreation area, sees. 29-32, T. 33 The sporangiophores are peltate or, as N., R. 9 E., Will County, 111. Walton (1949) has indicated for Proto- Horizon.—Francis Creek shale, lower calamostachys, the sporangiophore divides portion of Carbondale group, Pennsylva- into four pedicels, each bearing a single system, Illinois. nian sporangium. In any case, however, each Type specimen. — B-1304A and B- sporangiophore bears four sporangia di- 1304B, paleobotanical collection, Illinois rected toward the cone axis, as in Calamo- State Geological Survey, Urbana, 111. stachys.

Calamostachys type OF fructification Discussion This type of calamite fructification is A brief review of the genera of calamite characterized by the placement of sporan- fructifications is essential to understand the giophore whorls midway between the similarities and differences that have an im- whorls of sterile bracts. The sporangio- portant bearing on the phyletic placement of phores have been termed peltate, but Lacey Mazostachys. There are six basic types of (1943) has found that in one species, C. calamite fructifications. Included in the binneyana, the sporangiophore head is not ILLINOIS STATE GEOLOGICAL SURVEY

peltate but cruciate (divided into four Palaeostachya type of fructification arms). Six to sixteen sporangiophores per This type of calamite fructification is whorl are known, depending upon the spe- characterized by the insertion of sporangio- cies, and variations in number occur within phores in the bract axils. The paired vascu- some species. There are four sporangia on lar bundles of P. vera depart at the nodes. each sporangiophore, so the sporangia face According to Hickling (1907), one enters the cone axis. The bracts of successive the bract and the other continues up whorls alternate with each other, whereas through the axis to a point midway between the sporangiophores are superposed. Both the nodes, where it turns downward to the homosporous and heterosporous species are axil of the bract and out into the sporangio- recorded in the literature. phore. Although the position of the spo- According to Darrah (1936), the sporan- rangiophores of Palaeostachya is strikingly giophores in Macrostachya thompsonii are different from that in Calamostachys, the borne as in Calamostachys. This important rest of the sporangiophore structure is sim- discovery poses the problem as to whether ilar. The number of sporangiophores per Macrostachya and Calamostachys represent whorl in Palaeostachya varies with the spe- distinct genera, because the only apparent cies. There are ten sporangiophores and distinction is that of size. twenty bracts per whorl in P. gracilis, and From an anatomical standpoint, the cone sixteen to twenty sporangiophores in P. axis of C. binneyana is best known. There vera with an equal number of bracts. There is a distinct pith in this species. Surround- are four sporangia attached to each peltate ing the pith are three or four vascular bun- sporangiophore. The bracts of successive dles, depending upon the stele form. If whorls in Palaeostachya are not known to there are six vascular bundles, the central alternate with one another as in Calamo- axis is triangular with paired bundles; if stachys. They are usually homosporous, but there are eight vascular bundles, the axis one species, thought to be a Palaeostachya, is quadrangular. The vascular bundles sup- is reported to be heterosporous. plying the bracts alternate in the cone axis From an anatomical standpoint, the cone through successive whorls. On the inside axis of Palaeostachya vera^ is best known. of each bundle is the carinal canal with re- It was first described by Williamson mains of spirally thickened protoxylem. (1871), and he discussed it again later in Phloem has been recognized outside the greater detail (Williamson, 1888). He wood, so the vascular bundles are collat- originally named the fructification Cala- eral. Secondary wood is found in the axis. mites pedunculatus, but Seward (1898) re- comparison between Mazostachys and A named it Palaeostachya vera. Hickling Calamostachys reveals certain similarities (1907) gives a very complete description. in the anatomy of the cone axis : in the cen- Hickling (1907) reported that the pe- tral pith, in carinal canals and their ar- duncle of P. vera could not be distinguished rangement in the axis, in collateral bun- from a young vegetative shoot, and that the dles, in number of sporangiophores to some cone axis differs only in the slight pairing extent, and in spore type. The dissimilari- of the bundles and the lack of nodal discs ties are : position of attachment of sporan- or sclerized cortex. The axis has a distinct giophore to cone axis, number of sporangia pith cavity surrounded by a thin layer of per sporangiophore, heterospory in certain parenchyma cells up to ten cells thick with species of Calamostachys, and alternation sixteen to twenty carinal canals. Scott of successive bract whorls of Calamo- (1920) reported spiral tracheids occasion- stachys. Calamostachys occurs in older and ally present in the canals. The canals are younger strata than Mazostachys, ranging ^Baxter recently submitted a paper on a new species of from Lower to Upper Carboniferous. The Palaeostachya to the American Journal of Botany, and antiquity of the genus is based on the work Delevoryas is submitting a paper to the same journal de- scribing still another species of this same genus. These of Bureau (1914). two papers will greatly extend our knowledge of the genus. DISCUSSION 19

paired, so the vascular bundles are in pairs in the (Scott, 1920). The and arranged alternateh^ with the vallecu- successive whorls of bracts are superposed, lar canals of the cortex. Metaxylem, sec- and thus do not alternate as in Calamo- ondary wood, and some phloem tissue were stachys. recognized. The cortex occupies about one- Cingular'ia cantrilli differs from C. typica fourth of the radius of the axis and is char- in the lack of bracts; both insertions at the acterized by sclerenchyma strands which nodes bear four sporangia each. Williamson (1871) called nodal discs. Basically C. typica and Mazostachys Comparison of Mazostachys and Palaeo- pendulata are similar, not only as to the stachya reveals a number of differences. general position of the sporangiophores

One major distinction is the position of the (just beneath the sterile bracts), but also sporangiophore on the internode. The spo- with regard to the sporangia (pendulant

rangiophore in Mazostachys is high on the and facing down). They differ in number

internode ; low on the internode in Palaeo- of sporangia per sporangiophore, in shape stachya. This suggests that they possibly and degree of bract fusion, and in the fact

represent separate phyletic lines, or that that the sporangiophore is inserted slightly one is a plant of much later origin. Smith lower on the internode in Mazostachys. and White's (1905) specimen from the Both species of Cingularia occur in the of Maine, which they question- Middle-Upper Carboniferous of Europe, ably assigned to Palaeostachya, has been which would be approximately the same examined by Dr. S. H. Mamay (personal stratigraphic position as Mazostachys, and communication), and he reports that the hence both genera are of approximately the specimen does not show enough detail to same geologic age. even assign it to Calarnites. Accordingly, the fructifica- lower limit of the geologic range of Palaeo- Metacalamostachys type OF tion siachya is Upper Carboniferous and the upper limit is Autunian. Thus, in geolog- The genus Metacalamostachys, known ical terms, both genera first appeared about only from compressions, was proposed by the same time. Hirmer (1927). Volkmannia pseudo-ses-

silis and Cala?nostachys calathifera were Cirigularia TYPE OF FRUCTIFICATION placed in Metacalamostachys by Hirmer.

Cincfularia is known only from compres- According to Browne (1927), M. calathif- sions, and largely through the efforts of era is most likely a Calamostachys. Grand'- Weiss (1876) and Kidston (1917). Two Eury (1877) believed that M. pseudo-ses- species have been described, C. typica Weiss siHs represented the cone of Calamites pa- and C. cantrilli Kidston. In the former, laeaceus. Additional excellent descriptions the strap-like sporangiophores are found by Jongmans (1911) and Kidston (1911) inserted at the nodes immediately below the serve to clarify the morphology of this type bracts. The sporangiophores bear four of fructification. large pendulant sporangia which face down. The cone is attached to a shoot with foli-

The apex of the sporangiophores is divided, age of Asterophyllites palaeaceus. Cones and two sporangia are borne on each mem- measure up to 2.8 cm. long and 4 mm. ber of this bifurcation, one behind the broad. The sporangiophores originate im- other. The sporangia average 5 mm. in di- mediately below the bracts as illustrated by ameter or five times that of Calamostachys Grand'Eury (1877) or are partly fused to binneyana. There are twice as many bracts the bracts as illustrated by Kidston (1911).

as sporangiophores ; they are connate for The sporangiophores are pendulant as in half their length and form sharply pointed Mazostachys, and they bear one sporan- teeth. Fischer (1893) found the bracts gium about 1 mm. long. The exact number

and sporangiophores adnate or partly joined of bracts and sporangiophores is unknown, to one another, a position analogous to that but they are equal. 20 ILLINOIS STATE GEOLOGICAL SURVEY

There can be little doubt that this form phore does not bifurcate and bear two spo-

is closely allied with M. pendulata with re- rangia but terminates, bearing one sporan- lation to size and general organization. It gium. Also, the sporangia are attached only differs from M. pendulata in that the spo- at the end of the sporangiophore and not to

rangiophore is either at the node and fused the sporangiophore arm as in Mazostachys. with the bracts or immediately below the The exact number of sporangiophores and

bract, and each sporangiophore bears a sin- bracts is unknown but they are probably

gle sporangium. Also, it would appear that equal. There are two bracts to each spo- the sporangia are attached at a single point rangiophore in Mazostachys. to the sporangiophores, in contrast to A critical re-examination of all the older Mazostachys, in which the sporangia are material would seem to be appropriate, de- attached to the sporangiophore arm for spite the fact that perhaps little new in- some distance. According to Jongmans formation could be obtained because of lim-

(personal communication, 1952), there is itations imposed by the type of preserva-

definitely just one sporangium per sporan- tion. It is hoped that this paper may stimu- giophore. late further investigation at European insti- Weiss' (1876) species originally de- tutions that have type specimens. scribed by him as Stachannularia calathif- Metacalamostachys and Mazostachys are era, and subsequently transferred to Cala- of about the same geological age; the for- niostachys, would appear to be a problemat- mer is from the Middle-Upper Carbonifer- ous. ical form. Browne (1927) believes it to be Hirmer a species of Calamostachys, and PHYLOGENY OF THE CONES OF (1927) has assigned it to Metacalajno- THE CALAMITACEAE stachys. This confusion probably arose be- cause the species is known only from com- A sound basis for a starting point in a (fig- pressions. The original illustration series is the first essential in developing any interpre- 1 1 ) by Weiss has been subject to phyletic relationship. The geological evi- tation. However, illustrations of other spe- dence in the case of the Calamitaceae is cies of Stachannularia by Weiss definitely very helpful. As pointed out by Browne show sporangiophores with a single at- (1927), the so-called bractless fructifica- tached sporangium like those covered by tions, which I have called the pve-Cala?no- Hirmer's genus MetacaMtnostachys. stachys type, are geologically the oldest The works of Schenk (1876) and Ster- type known, and occur in the very lowest zel (1882) are interesting because the portion of the Lower Carboniferous. Also, fructifications of Annularia sphenophyl- some of these fructifications have been loides shown are now known to be of the found attached to morphologically primi-

Metacalamostachys type. This is of par- tive calamites. They are sometimes called ticular interest because of the fact that the bractless; they may have infrequent whorls foliage of Annularia sphenophylloides occurs of bracts scattered over the axis at consid- in the same plane, although it is not organ- erable intervals. Walton's (1949) Proto- ically connected, as the specimen herein de- calamostachys was collected from the Cal- scribed as Mazostachys. I have copied ciferous sandstone on the Island of Arran.

Schenk's (1876) reconstruction of the fruc- Its geologic age and anatomy place it with tification of Annularia sphenophylloides the prt-Calamostachys type of fructifica-

(pi. 6, fig. 3). If this reconstruction is ac- tions. curate, it is strikingly similar to Mazo- Browne (1927) reported the infrequent stachys pendulata in that the sporangio- bract whorls associated with some of the phore departs from the cone axis high on the bractless forms consist of almost unreduced internode just below the node and bract. leaves. She has postulated that the phylet-

However, it differs from Mazostachys in ically primitive forms were this type and that the apical portion of the sporangio- gave rise to the bracteate types, such as PHYLOGENY OF CONES OF CALAMITACEAE 21

Calarnostachys and Palaeostachya. In the number of sporangia per sporangiophore. bracteate types, the leaves became inter- Accordingly, the Calamostachys type of in- calated between the whorls of sporanglo- sertion and the number of sporangia per phores and functioned largely to protect the sporangiophore suggest that the Calamo- sporangia rather than as photosynthetic stachys type of fructification was derived leaves. from the Pothocites and Protocalamo- The reports of Nathorst (1914), Krau- stachys type. Geological evidence favors sel and Weyland (1926), and others on Calamostachys rather than Palaeostachya

the Middle Devonian Hyeniales have if Bureau's (1914) Lower Carboniferous

greatly clarified previous views concern- Calamostachys occidentalis is valid. Wal- ing morphological interpretations of the ton's (1949) statement that the peltate

primitive cones of the Calamitaceae. One sporangiophore is derived by a later modifi- can readily see a series from the Psilophy- cation of the pedicellate sporangiophore of tales to Hyenia and Calamophyton and Protocalamostachys and possibly Potho- then to the ipie-Calamostachys type of fruc- cites, and Lacey's (1943) report that the tification. The postulation that the bracts sporangiophore of Calamostachys binneyana

w^ere originally parts of the cones and there- is not peltate but cruciate, may be consid-

fore primitive is immediately confronted ered evidence that certain primitive char- with an obstacle—most of the pre-Calamo- acters of the bractless genus Protocalamo- stachys types were definitely bractless. stachys are present in the well-known genus Where bracts are present, they resemble Calamostachys. I believe that Calamo- leaf-like organs. Jeffrey's (1899) view that stachys was derived from the bractless the bracts became lost in A stero catamites forms during Lower Carboniferous. and is confronted with a prob- I am well aware that any proposed phy- least lem, at for Asterocalamites] Hyenia letic arrangement, particularly with frag- and Calafiiophyton have preceded may mentary fossil plants, is subject to indi- Asterocalamites phyletically as neither had vidual evaluation of the characteristics used leaves or bracts on the fertile shoots. in developing phyletic lines. Agreement is Walton (1949) calls attention to the frequently lacking, but by attempting to re- nonpeltate character of the sporangio- construct phyletic development, ideas and a phores of Protocalamostachys. The sporan- new perspective of the problem or problems gia are borne on short pedicels and compare are more clearly evident. In the case at with the cruciate markings in Pothocites. hand, evidence can be presented suggesting He feels that the sporangiophore may have a more axillary origin for the sporangio- been a forked organ comparable to the phore trace in certain species of Calamo- forked leaf of the Protoarticulatineae and stachys. Hickling (1910) reports such a that the peltate sporangiophore associated situation for C. bijineyana and Browne with the bracteate fructifications is a later (1927) reviews such evidence for C. Zeil- modification. leri and C. magnae-crucis. Such data cer- Other views on the morphology and phy- tainly suggest a variance of the path of the letic significance of bracts may be found in sporangiophore trace and could be used to Bower (1908), Scott (1923), and else- support the theory that Palaeostachya was where. An excellent review may be found an early derivation from Calamostachys, or in Browne (1927). that the first genus actually preceded. As

On the assumption that the bractless it is clear, however, that Protocalamo- forms are primitive, we must decide which stachys preceded both Calamostachys and genus of the bracteate forms was first de- Palaeostachya phyletically, and that it is rived from the bractless forms. I attach more closely related to the former than the considerable significance to the position of latter, I believe that the axillary origin of insertion of the sporangiophore on the the sporangiophore trace in certain mem- cone axis with respect to the bracts and the bers of the Calamitaceae was derived. 22 ILLINOIS STATE GEOLOGICAL SURVEY

I agree with Hickling (1907) that Pal- opment there are two sporangia per sporan- aeostachya is derived from the Ca.amo- giophore in Mazosiachys and only one in stachys type of cone but am not convinced Metacalamostachys, and the pith cavity is that the sporangiophore was a ventral lobe lacking in Mazostachys. of a sporophyll and the bract a dorsal lobe. The side branch of Cifigularia poses an

A downward slide of the sporangiophore interesting problem. Cingularia is placed to an axillary position did occur, but the in this line of development because the spo- vascularization, at least in Palaeostachys rangia hang down in a pendulant manner, vera, is not far removed from that in Cal- and the sporangiophores occur either very amostachys. high on the internode or at the node above. The work of Darrah (1936) has raised If we exclude Cingularia, there appears to the question as to whether size alone is be a reduction in the number of sporangia sufficient to retain Macrostachya distinct per sporangiophore. There are four spo- from Calajuostachys. I have retained the rangia per sporangiophore in Cingularia distinction here, as shown in figure 5, but typica, but in C. cantrilli there appears to considered Macrostachya with the Calanio- be a double set of sporangiophores, each stachys line of development. Therefore, sporangiophore bearing four sporangia. two lines of calamite cones (fig. 5) have arisen from CaJarnostachys: the Palaeo- SUMMARY stachya line, terminating with Palaeo- 1. A new type of calamite fructification stachya but also including Huttonia, and a from the paleobotanically famous Mazon second line, including Mazostachya, with Creek area of Will County, 111., from the Cingularia questionably assigned as a side lower part of the Carbondale group of branch, and terminating with Metacala- Pennsylvanian age (Francis Creek shale rnostachys. The Palaeostachya line is char- above northern Illinois No. 2 coal bed) is acterized by the axillary position of the described as Mazostachys pendulata gen. peltate sporangiophore, four sporangia per et sp. nov. sporangiophore, and the presence of a pith 2. M. pendulata is known from a com- cavity. The second line is characterized by bination petrification-and-compression type a nonpeltate sporangiophore borne high on of preservation. the internode or at the next node above, 3. M. pendulata is characterized ana- and the sporangia are attached in a pen- tomically by a central pith and a pairing of dulant manner. On this main line of devel- vascular bundles with annular protoxylem, Metacalamostachys spirally thickened sporangiophore, and Cingularia I bract traces. Metaxylem and secondary wood have tracheids with scalariform, mul- Mazostachys tiseriate scalariform, and reticulate pit- Huttonia ting.

I Palaeostachya 4. Morphologically M. pendulata is

Macrostachya characterized by six sporangiophores, in- J serted just below the node of the next

whorl above ; each sporangiophore bears Calamostachys two pendulant sporangia. There are sterile bracts, six opposite and six alternating with the six sporangiophores. « Protocalamostachys 5. Phyletically M. pendulata is placed ^ I 5 I <« above Calamostachys and below Metacala- O Pothocites mostachys. Cingularia is a side branch on the main line of phyletic derivation in which Mazostachys and Palaeostachya Fig. 5. —A proposed phyletic sequence for the fructifications of the Calamitaceae. are the basis for a forked phyletic tree. REFERENCES 23

REFERENCES

Bower, F. O., 1908, The origin of a land flora: Krausel, R., and Weyland, H., 1926, Beitriige London, MacMillan, 727 p. zur Kenntnis der Devon-flora, II: Abhandl. Browne, I. M. P., 1927, A new theory of the der Senekenbergischen Naturforschenden Ge- morphology of the Calamarian cone: Ann. sellschaft. Band 40, v. 2, p. 115-155. Bot., V. 41, p. 301-320. Lesquereux, Leo, 1866, An enumeration of the Bureau, E., 1914, Bassin Houiller de la Basse- fossil plants found in the Coal Measures of Loire: Fasc. IL Descriptions des Flores fos- Illinois: Illinois Geol. Survey Paleont., v. 2, siles: Etude des gites mineraur de la France, p. 427-470.

p. 209-248. , 1870, Report on the fossil plants of Cady, G. H., 1919, Geology and mineral re- Illinois: v. 4, p. 375-508. sources of the Hennepin and LaSalle quad- Lacey, W. S., 1943, The sporangiophore of Cala- rangles: Illinois Geol. Survey Bull. 37. mostachys: New Phytologist, v. 42, p. 98-102. CoNDiT, Carlton, and Miller, A. K., 1951, Moore, R. C, Wanless, H. R., Weller, J. M., Concretions from Iowa like those from Mazon et al., 1944, Correlation of Pennsylvanian for- Creek, Illinois: Jour. Geol., v. 59, p. 395-396. mations of North America: Geol. Soc. Am. Darrah, W. G., 1936, A new Macrostachya Bull., V. 55, p. 657-706. from the Carboniferous of Illinois: Bot. Mus. Nathorst, a. G., 1914, Nachtrage zur palao- Leafl. Harvard Univ., v. 4, p. 53-63. zoischen Flora Spitzbergens: Fossilen Flora Delevoryas, Theodore, and Morgan, Jeanne, Polarlander, v. 1, no. 4, p. 1-110. 1952, Tubicaulis multiscalariformis: A new Noe, a. C, 1925, Pennsylvanian flora of north- American Coenopterid: Am. Jour. Bot., v. 39, ern Illinois: Illinois Geol. Survey Bull. 52. p. 160-166. Savage, T. E., 1927, Significant breaks and over- Fischer, E., 1893, Einige Bemerkungen iiber die laps in the Pennsylvanian rocks of Illinois: Calamarieen-Gattung Cingularia: Mitt. d. Am. Jour. Sci., v. 14, p. 309. Naturf. Ges. in Bern, p. 1-7. ScHOPF, J. M., 1938, Two new Lycopod Grand'Eury, F. C, 1877, Memoire sur la Flore from the Illinois Pennsylvanian: Trans. Illi- Carbonifere du Departement de la Loiret et nois Acad. Sci., v. 30, p. 139-146. du Centre de la France: Mem. Acad. Sci. Inst. Schopf, J. M., Wilson, L. R., and Bentall, Ray, 1944, An annotated synopsis of Paleo- Nat. France, v. XXIV, no. 1, p. 1-616. Hartung, W., 1933, Die Sporenverhaltnisse der zoic fossil spores and the definition of generic groups: Illinois Geol. Rept. Inv. 91. Calamariaceen: Inst. Palaobot. u. Petrog. der Survey Scott, D. H., 1920, Studies in fossil botany: 3rd Brennsteine Arb., 3, p. 95-149. ed., V. I, Black. Hickling, George, 1907, The anatomy of Palae- London, 434 p.

, 1923, Studies in fossil botany: 3rd ostachya vera: Ann. Bot., v. 21, p. 369-386. ed., V. II, London, Black. 446 p. , 1910, The anatomy of Calamostachys Schenk, a., 1876, Ueber die Fructstande fos hinneyana, Schpr. : Mem. Proc. Manchester siler Equisetin Botanische Zeitung: v. 34, p. Lit. Phil. Soc, V. 54, pt. 3, no. 17, p. 1-16. 528-540, 625-634. Hirmer, Max, 1927, Handbuch der Palaobot- Smith, G. O., and White, David, 1905, Geology anik: Miinchen and Berlin, R. Oldenbourg, of the Perry Basin in southeastern Maine: 708 p. U. S. Geol. Survey Prof. Paper 64-65. H0EG, O. A., 1942, The Downtonian and De- 35, p. Sterzel, T., 1882, Ueber die Fruchtahren von vonian flora of Spitsbergen: Norges Svalbard- J. Anniilaria sphenophylloides Zenker Sp.: og Ishava-Undersokelser, Skrifter Nr., v. 83, Zeitschr. der Deut. Geol. Gesell., v. 39, p. 685- p. 1-228. 691. HosKiNS, J. H., and Cross, A. T., 1943, Mono- graph of the Paleozoic cone genus Botvman- Stewart, W. N., 1950, Report on the Carr and Daniels collections of fossil plants from Ma- ites (Sphenophyllales) : Am. Mid. Nat., v. 30, zon Creek: Trans. Illinois Acad. Sci., v. 43, p. 113-163. p. 41-45. Janssen, R. E., 1940, Some fossil plant types of Walton, John, 1949, On some Lower Carbonif- Illinois: Illinois State Museum Sci. Papers, erous Equisetineae from the Clyde area. I. V. 1. Protocalamostachys arranensis gen. et sp. nov. Jeffrey, E. C, 1899, The development, struc- — a hitherto undescribed type of Strobilus: ture, and affinities of the genus Equiseium: Trans. Royal Soc. Edinburgh, v. LXI, pt. Ill, Mem. Boston Soc. Nat. Hist., v. 5, p. 155-190. no. 26, p. 729-732. Jongmans, W. 1911, Anleitung zur Bestim- J., Wanless, H. R., 1929, Geology and mineral re- der Karbonpflanzen West-Europas: mung sources of the Alexis quadrangle: Illinois Mededeelingen van de Rijksoporing van Delf- Geol. Survey Bull. 57, p. 49 and 89. stoflFen, v. 1, p. 1-482. _, Geology and mineral resources of KiDSTON, R., 1911, Les Vegetaux houillers re- the Beardstown, Glassford, Havana, and Ver- cueillis dans le Hainault beige et se trouvant mont quadrangles (unpublished manuscript, dans les collections du Musee royal de Bel- Illinois Geol. Survey). gique: tom. IV, p. 93-129. Weiss, C. E., 1876, Steinkohlen-Calamarien:

, in Kidston, R., Cantrill, T. C, and Abhandlugen zur geologischen Specialkarte Dixon, E. E., 1917, The Forest of Wyre and von Preussen und den Thuringischen Staaten, the Titterstone Clee Hill Coalfield: Trans. Band II, Heft 1, 149 p. and atlas. Royal Soc. Edinburgh, v. 51, p. 139-147. Williamson, W. C, 1871, On a new form of KosANKE, R. M., 1950, Pennsylvanian spores of Calamitean Strobilus from the Lancashire Illinois and their use in correlation: Illinois Coal Measures: Mem. Lit. and Philos. Soc. Geol. Survey Bull. 74. Manchester, v. 4, p. 248-265. 24 ILLINOIS STATE GEOLOGICAL SURVEY

WiLLMAN, H. B., , 1888, Organization of the fossil and Payne, J. N., 1942, Geol- plants of the Coal Measures. XIV. The true ogy and mineral resources of the Marseilles, fructifications of Calamites: Philos. Trans. Ottawa, and Streator quadrangles: Illinois Royal Soc. London, 179 B, p. 47-57. Geol. Survey Bull. 66, p. 100.

., and Scott, D. H., 1894, Further ob- Zeiller, R., 1893, Etude sur la constitution de servations on the organization of fossil plants I'appareil fructicateur des : from the Coal Measures, I. Calamites, Cala- Mem. Soc. Geol. France, Paleont. Mem. 11, mostachys, and Sphenophyllum: Philos. Trans. p. 3-39. Royal Soc. London, 185 B, p. 863-959.

EXPLANATION OF PLATE 1

Fig. 1 —Mazostachys pendulata gen. et sp. nov., specimen B-1304A, shown natural size. The cones are attached by means of a short peduncle to a shoot of Calamites with a primary branching pattern of the Eucalamite group. The scattered foliar whorls are those of Annularia spheno- phylloides which are not connected to Mazostachys pendulata.

2—A photomicrograph of B-1304A, cone 14, peel 3, of a fracture-surface peel made prior to grinding and peeling, in transverse plane. Whorl 2 of this cone covers a distance of 1900 microns. There were 38 peels taken; the peel numbers, shown in their relative position at the right, are illustrated in plates 2 and 3.

3—Essentially a longitudinal section of cone 6 from B-1304B, peel 3. This is a fracture-surface peel and shows the cone axis, nodes, bracts, sporangiophores, and apical portion of the cone (X9).

[26] Illinois Statk Geological Survey R. I. 180, Plate 1

3 ^.rii:. r

KOSANKE, MaZOSTACHYS PENDULATA GEN. ET SP. NOV. Illinois State Gkological Survey R. I. 180, 1'late2

KOSA.NKE. MaZOSTACHYS PKNDILATA GEN. ET SP. NOV EXPLANATION OF PLATE 2

Fig. 1 —Transverse section of cone 14 (B-1304x'\), peel 1, from the base of whorl 2 as shown in plate 1, figure 2. The rough surface of the sporangia at the bottom represents the fracture line be- tween the two halves of the specimen. The preserved portion of the cone axis is shown in the center surrounded by twelve sporangia and the bracts (X30).

2 —Transverse section of cone 14 (B-1304A), peel 9, from whorl 2 as shown in plate 1, figure 2. The sporangiophores were first observed departing from the cone axis in peel 7. Six spo- rangiophore arms are seen departing from the cone axis in this peel. The position of the bracts is clearly shown at the top as is the arching of the sporangia next to the bracts. The arching is thought to be the result of the cone's compactness (X30).

3—Transverse section of cone 14 (B-1304A), peel 17, as shown in plate 1, figure 2. All six sporangiophore arms are well developed, and the aerenchyma tissue is plainly shown to the right between the two sporangiophore arms. The absence of all twelve sporangia indicates a slightly oblique section or curvature of the cone (X30).

[29] EXPLANATION OF PLATE 3

Fig. 1 —Transverse section of cone 14 (B-1304A), peel 19, as shown in plate 1, figure 2. Almost all traces of three of the sporangiophore arms have disappeared from this section, which sug- gests that they do not depart from the axis at precisely the same time. Further evidence of this is that the bottom arm is still present, although it should have completely disap- peared first, owing to the slightly oblique section. The two sporangiophore arms in the upper half of the photomicrograph are shown in almost complete development. The arm to the left is forked at its apex and overlaps adjacent sporangia. Thus each sporangiophore arm bore two sporangia. The forking is accentuated because the arm follows the curvature of the sporangia and lies immediately above and between the two sporangia which it bore. The aerenchyma tissue with large lacunae is well shown to the right (X30).

2 —Transverse section of cone 14 (B-1304A), peel 35, from the base of whorl 3 as shown in plate 1, figure 2. Seven sporangia are visible in the lower half of the section, and the bracts have begun to ascend. Note that the bracts of whorl 3 appear to be alternate with the remaining bracts from the whorl below. Actuallv it can be demonstrated that they do not alternate (X30).

3—Transverse section of cone 14 (B-1304A), peel 38, as shown in plate 1, figure 2. There are twelve sporangia, and this peel approximately duplicates that of peel 1 from whorl 2 one node below. The bracts appear to alternate with those from the node below, but they do not be- cause there has been apical shifting of the bracts from whorl 2 (X30).

[301 Illinois State Ckological Si kvky K. I. 180, Plate 3

O*

KOSANKE, MaZOSTACHYS PENDULATA GEN. ET SP. NOV. Illinois State Geological Survey R. I. 180, Plate 4

KosANkE. Mazostachys pendllata gen. et sp. non. EXPLANATION OF PLATE 4

Fig. 1 —Mazostachys pendulata gen. et sp. nov., specimen B-1304B, shown natural size. The opposite half may be seen in plate 1, figure 1.

2—Sporangial wall tissue of a sporangium from whorl 7, cone 1, peel 3, B-1304A. The thickenings of the lateral walls are shown. Most of these do not entirely cross from one lateral wall to another, but some of them do (X175).

3—Longitudinal section from cone 13 (B-1304A), peel 4, through a portion of the cone axis show- ing the carinal canal with annular thickenings, all that remains of the protoxylem. Scalari- form tracheids of the metaxylem are adjacent to the carinal canal (X350).

4—Longitudinal section from cone 15 (B-1304A), peel 6, through a portion of a bract showing a mass of spiral tracheids constituting the vascular trace of the bract (X350).

5—Sporangiophore vascular trace from cone 1 (B-1304A), peel 3, departing from the axis of the cone. These spiral tracheids are similar to those of the bracts (X350).

6—Longitudinal section from cone 6 (B-1304B), peel 3, through a small portion of the cone axis showing scalariform pitted tracheids (X350).

[33] EXPLANATION OF PLATE 5

Fig. 1 —Transverse section of cone axis of cone 14 (B-1304A), peel 10. The six sporangiophore arms may be seen at the periphery of the cone axis. Six carinal canals are present, and there are remains of annular thickenings in two of the canals. Note the absence of a pith cavity (XI 15).

2—Two sporangia oi Mazostachys pendulata from cone 15 (B-1304A), peel 4, showing sporangial walls and spores. The sporangia are typically completely filled with spores; the sporangial wall is one cell thick. Note the areae contagionis on two of the spores (X398).

[34] Illinois Statk (Ikolcxmcai, Si km:y K. [. 180, Platk

KOSANKK, MaZOSTACHVS FKNDLLATA GKN. KT SP. NOV. Illinois State Geological Survey K. I. 180. |>LATE 6

KosANkE, AIazostachys pe>dllata gen. ET SP. N( EXPLANATION OF PLATE 6

Fig. 1 —Longitudinal section of a part of whorl 4, cone 1 (B-1304A), peel 3. At the base of the photo- micrograph is a portion of the sporangiophore arm of whorl 6. Above this level is the node and departing bract enclosing a sporangium. Much higher is the sporangiophore and above this the aerenchyma tissue (X43). £—A reconstruction of two nodes and one internode showing bracts, a sporangium, sporangio- phore, and above this the aerenchyma tissue. The section shown in figure 1 served as the basis for this reconstruction (X36). 3—Reconstruction of a longitudinal section of a portion of a cone associated by Schenk with Annularia sphenophylloides. This fructification is similar to Maiostachys in size and position of pendulate sporangiophores, but each sporangiophore had only one sporangium, sporangia are attached to the end of the sporangiophore, and bracts are shorter. (By Schenk from Jongmans.)

4—Oblique section through a small portion of a node from cone 13 (B-1304A), peel 6. This type of tracheid pitting was called scalariform or multiseriate pitting by H<^eg (1942) and multi- seriate scalariform by Delevoryas and Morgan (1952) (X357).

5 —Aerenchyma tissue showing large lacunae and a network of parenchyma cells. This sec- tion is from cone 14 (B-1304A), peel 19, from the second whorl above the base of the cone (X175).

6—One spore oi Mazostachys pendulata, obtained in the isolated state by mechanically dissecting a sporangium from a cone and macerating the sporangium in dilute HCl. The spore meas- ures 60 microns in its largest diameter and 58.6 microns at right angles to that diameter. Note the area contagionis between the rays of the triradiate suture.

[37] Illinois State Geological Survey, Report of Investigations 180

37 p., 6 pis., 5 figs., 1955

LIBRARY ENVIRONMENTAL PROTECTION AGENCX STATE OF ILLINOIS /

SPRINGFIELD, II.L!,\'0)S