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Studies of Paleozoic Seed Ferns: Anatomy and Morphology of Morphology of Microspermopteris Aphyllum

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Studies of Paleozoic Seed Ferns: Anatomy and Morphology of Morphology of Microspermopteris Aphyllum Amer. J. Bot. 63(10): 1302-1310. 1976. STUDIES OF PALEOZOIC SEED FERNS: ANATOMY AND MORPHOLOGY OF MICROSPERMOPTERIS APHYLLUM1 THOMAS N. TAYLOR AND RUTH A. STOCKEY Departmentof Botany,The Ohio State University,Columbus 43210 A B S T R A C T The discoveryof numerousspecimens of the monostelicpteridosperm genus Microspermop- teris in Pennsylvaniancoal ball petrifactionsfrom the Lewis Creek and What Cheer localities providesadditional information about the anatomicaland morphologicalvariability within thegenus. Specimensare now knownup to 1.1 cm in diam thatbear epidermalappendages in theform of variously-shapedtrichomes. The externalsurface of the stemis furtherornamented by longitudinalflaps of corticaltissue. Petiolesexhibiting a singleC-shaped vascular strand with abaxial protoxylemare producedin a 2/5 phyllotaxy.Large petiole bases that clasp the stemproduce primary pinnae alternately. The presenceof axillarybranching appears simi- lar to thatreported in Callistophytonand Lyginopteris. Triarchto polyarchadventitious roots, some withsecondary tissues, are producedat both nodal and internodalregions. Of the cur- rentlyrecognized monostelic seed ferngenera, Microspermopteris is most similarto Heter- angium. Informationis presentedthat supportscurrent ideas regardingthe evolutionof the gymnospermiceustele from protostelic Devonian ancestors. THE GENUS MICROSPERMOPTERISwas initiallyde- axes representingnumerous orders of branching. scribedby Baxter (1949) frompetrifaction ma- Specimens were collected fromthe What Cheer terial collected fromthe What Cheer locality of and Lewis Creek petrifactionlocalities which Iowa, witha subsequentvariety added fromma- are designatedMiddle and Early Pennsylvanian, terialcollected from the FlemingCoal of Kansas respectively (Good, 1975). Cellulose acetate (Baxter, 1952). Recently,the taxon has been peels were used to determineanatomical and reportedfrom several coals in Illinois (Mahaffy, morphologicalfeatures. All specimensand slides 1975) and easternKentucky (Taylor and Stoc- are housed in the PaleobotanicalCollections, De- key, 1975). In the initialdescription the genus partmentof Botany, The Ohio State University, was characterizedas a leaflesspteridosperm with and include acquisition numbers 6,489-6,912, an exarch, partiallymixed protosteleup to 5.0 12,177-12,420, 12,422-12,466. mm in diam. Stems were describedas being ir- regularin outlineand lackingleaves. Multicellu- DIAGNOSIs-Microspermopterisaphyllum (Bax- lar emergencestogether with adventitiousroots ter) Taylor et Stockey emend. Stems up to 1.1 were described on some stem fragments.The cm in diam withmixed exarchprotostele consist- genus was furthercharacterized by the presence ing of large metaxylemtracheids and up, to 10 of secondary tissues, concentricbranch traces, peripherallypositioned protoxylem strands. Pet- and distichousbranching. ioles arranged in 2/5 phyllotaxy,with single Duringthe past severalyears we have accumu- trace slightlyC-shaped with abaxial protoxylem; lated a numberof specimensof this plant from petiole base large and clasping up to one half two petrifactionlocalities in North America, stem circumference,primary pinnae produced especially the Lewis Creek site in eastern Ken- alternately;axillary branches with abundant sec- tuckywhere it representsa common elementof ondaryxylem. Cortex of thin-walledparenchyma the flora. It is our intentin this paper to provide with secretorycells in youngerstems, and pe- additional informationabout this taxon, offera ripherally disposed longitudinal sclerenchyma; reinterpretationof severalmorphological features, peridermof thick-walledradially aligned cells. and discuss the genus Microspermopterisin light Stem surface ornamentedby longitudinalflaps of recenttheories regarding the evolutionof the of corticaltissue, and multicellular,typically flat- gymnospermoushabit. tened trichomes. Metaxylemtracheids elongate, The followingdescription is based upon 36 with multiseriatebordered pits; protoxylemele- mentsscalariform. Primary xylem pentagonal in 'Received for publication17 December 1975. cross section and divided into 5 sectionsby lon- The authorsare indebtedto Dr. John W. Hall for gitudinal parenchymaplates that radiate from making several What Cheer specimensavailable for study,and to the National Science Foundationfor par- stem center; protoxylemstrands occur in pairs, tial supportof this research(BMS74-21105). one strandon each side of a parenchymaplate. 1302 November-December,1976] TAYLOR AND STOCKEY-MORPHOLOGY OF MICROSPERMOPTERIS 1303 Secondary vascular tissues well-developed,sec- trace has been formedit remainswithin the stem ondaryxylem tracheids with uni- and multiseriate cortexfor some distance(approximately 6.0 mm) bordered pits. Xylem rays up to 2 cells wide. beforepassing outwardinto the petiole base. Triarchto polyarchadventitious roots, some with The most conspicuousaspect of the cross sec- secondarytissues, arising at both nodal and inter- tional configurationof Microspermopterisis the nodal regions. presence of large, angular metaxylemtracheids (Fig. 1). In longitudinalsection, metaxylem ele- Type species- Microspermopterisaphyllum mentsmeasure up to 1.0 mm and possess slightly Baxter 1949, p. 297-298, pl. 2, Fig. 1-4; pl. 3, tapered end walls, and multiseriate,simple-retic- Fig. 5-7; pl. 4, Fig. 8-12. ulate-borderedpits (Fig. 19). Parenchymain the metaxylemis organized into longitudinalplates Synonymy-1952. Microspermopterisaphyl- up to threecells wide thatradiate from the center lum var. kansensisBaxter; Trans. Kansas Acad. of the stem to formthe characteristicpentagonal Sci. 55: 101-103. stem configuration(Fig. 4). We have observed a largernumber of plates in some stems;however, DESCRIPTION-General features-Stems range in all instancesthis increase in numberof plates from0.5-11.0 mm in diam, with the most com- appears to have resultedfrom additional divisions plete specimenextending approximately 38.0 cm of the originalfive plates (Fig. 1). through the coal ball. Most stems exhibit a sinuous habit in the matrixmaking longitudinal Axillary branching-One of the newly inter- stem sectionsdifficult to obtain and possiblyre- pretedfeatures of Microspermopterisis the pres- flectinga liana or shrub-likegrowth habit. A ence of axillarybranching (Fig. 24). Figures 12- pentagonal protostele containing longitudinal 14, 17-18 illustrateprogressive stages of stem plates of parenchyma (Fig. 1, 4, 15) is sur- and petioleseparation, and axillarybranch origin. rounded by a well-developedzone of secondary The productionof an axillarybranch begins with xylem. To the outsideof this tissueis a vascular a large amount of secondary xylem extending cambium, a zone of secondary phloem, and a laterallyfrom the stem stele (Fig. 13). At a parenchymatouscortex containing patches of slightlyhigher level (Fig. 12, 14) it becomes sclerenchymaand secretorycells (Fig. 1, 4, 7, separated to form a tereteaxillary branch stele. 11, 16). Multicellulartrichomes and longitudinal Figure 17 representsthe axillarybranch, traced corticalflaps extendfrom the stem surface (Fig. for a distanceof approximately5.3 mm. In this 2, 5, 8, 9, 25). specimenthe more distal organizationof the axil- lary branch could not be determinedbecause of Primarytissues-In transversesection the pri- faultypreservation. In tranversesection the axil- maryxylem is typicallypentagonal in outline as lary branch stele is circularwith a centralregion definedby radiatingparenchyma plates (Fig. 1, of small crushedcells possiblyrepresenting proto- 4). At the moredistal levels of the stemthis con- xylemelements or parenchyma,surrounded by a figurationis maintainedby clustersof metaxylem zone of up to 12 rows of secondaryxylem tra- tracheids separated by patches of parenchyma cheids. At the point of axillary branch origin (Fig. 7). In the stemthat contains the most sec- there is a distinctinterruption in the secondary ondary xylem (Fig. 4) (Lewis Creek) the pri- xylem of the stem stele (Fig. 17, 18, arrows). maryxylem measures 1.1 mm in cross sectional At higherlevels a few files of secondaryxylem diam,whereas the stemillustrated in Fig. 1 (What tracheidsare present. Cheer) has approximatelyone-third the second- ary tissue developmentand the primaryxylem is Petiole-The petiole or rachis clasps the stem 1.7 mm in diam. The stem illustratedin Fig. 7 axis for approximatelyone half of the diameter is representedalmost entirelyby primarytissues. of the stem at the level of axillarybranch forma- The protoxylem strands in Microspermop- tion (Fig. 12, 24). At higherlevels the cross teris, as determinedby cell diameterand pres- sectionalconfiguration of the petiole is V-shaped ence of annular-scalariformwall thickenings,are to almost flattened (Fig. 17, 21). In several situatedon eitherside of the parenchymaplates specimens it has been possible to trace alter- that radiate fromthe stem center (Fig. 3). Fig- natelyarranged primary pinna axes (Fig. 20-22). ures 3, 6, and 10 illustratesuccessively higher The pinna trace consists of a single lateral xy- levels throughprotoxylem strands in a region of lary extension that shows the same anatomical organizationas the trace. Two trace petiole longitu- emission. In Fig. 3 the arrows on either dinal extensionsof tissue ornamentthe adaxial side of the metaxylemparenchyma indicate the surfaceof the petiolenear the level of pinna trace small protoxylemstrands. At a slightlyhigher formation(Fig. 21-22). It has not been pos- level (Fig. 12) the 2 strandsunite to form an sible to demonstrateconclusively
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